Beruflich Dokumente
Kultur Dokumente
Department of
Agriculture
Forest Service
Agriculture
Handbook
727
July 2008
The
Woody Plant
Seed Manual
The
Woody
Plant
Seed
Manual
Agriculture
Handbook
727
Cover photo
The scientific names for the seeds shown on the cover are identified by the key
and photo below. Seeds 2 and 7 have had their wings removed by cleaning.
1. Green ash (Fraxinus pennsylvanica Marsh.)
2. Ponderosa pine (Pinus ponderosa P. & C. Lawson)
3. Northern red oak (Quercus rubra L.)
4. Witch hazel (Hamame lis virginiana L.)
5. Service berry (Amelanchier arborea (Michx. F.) Fern.)
6. Persimmon (Diospyros virginiana L.)
7. White fir (Abies concolor (Gord. & Glend.) Lindl. Ex Hildebr.)
8. Tulip poplar (Liriodendron tulipifera L.)
4
5
8
7
Dedication
This handbook on the seeds of woody plants would not be possible if not for the pioneering work of many individuals in
past years. They worked without the modern laboratory and information retrieval services that we now routinely use and
take for granted. Their early efforts in the first half of the 20th century solved many seed problems and pointed the way for
later research on numerous subjects. Without them, our body of knowledge about the seeds of woody plants would not be
what it is today. Many contributed, but for their extensive work and leadership, we dedicate this book to the following
pioneers:
George S. Allen Canadian Forestry Service & University of British Columbia
Henry I. Baldwin New Hampshire Forestry Department
Lela V. Barton Boyce Thompson Institute
Claude E. Heit New York Agricultural Experiment Station
Nikolas T. Mirov USDA Forest Service
Paul O. Rudolf USDA Forest Service
Charles F. Swingle USDA Bureau of Plant Industry
W. R. Van Dersal USDA Soil Conservation Service
Philip C. Wakeley USDA Forest Service
Acknowledgments
The development and publication of this book was supported by the USDA Forest Services Research and Development
(R&D) and State and Private Forestry (S&PF) National Offices as well as by R&Ds Southern and Northeastern Research
Stations, which supported Dr. Bonner and Ms. Nisley, respectively. We especially thank Drs. Jerry Sesco and Robert Lewis,
Deputy Chiefs (retired) for Research and Development; Dr. Peter Roussopoulus, Director of the Southern Research Station;
Dr. Michael T. Rains, Director of the Northeastern Research Station; and Debra Dietzman, Communications Director at the
Northeastern Research Station, for their continued financial and personal support.
We also thank John K. Francis (retired after many years at the International Institute of Tropical Forestry), Robert P.
Karrfalt (S&PF, Regeneration, Nurseries, and Genetics Resources National Team), Susan E. Meyer (Rocky Mountain
Research Station), Peyton W. Owston (retired from the Pacific Northwest Research Station), and John C. Zasada (retired
from the (North Central Research Station), the regional coordinators who solicited and organized the authors of the 236
genera.
We are grateful to Dr. Stanley R. Krugman for getting this effort going and sharing his experiences from the production of
AH 450, Seeds of Woody Plants in the United States (1974). Sharon Friedman, Jacob L. Whitmore, Calvin Bey, Sam Foster,
and Marilyn Buford served as our liaisons to the R&D National Office; Karl Dalla Rosa and Hal Brockman were the
liaisons to S&PF; and Frank Burch to National Forest Systems. Becky Loth, at the National Seed Laboratory served
admirably as the webmaster of our interim website, and Laura Cricco, Phyllis Grinberg, Jean B. Holland, Pamela Huntley,
Barbara Johnson, and Kathy McManus, all of the Northeastern Research Station, provided much appreciated administrative
and computer support.
Contents
Introduction
vii
Invasives vii
Part 1
Chapter 1
Chapter 2
Chapter 3
Chapter 4
Chapter 5
Chapter 6
Chapter 7
Part 2
Part 3
Appendices
1199
Contents
iii
Introduction
The first comprehensive handbook on the seeds of trees
and shrubs produced by the USDA Forest Service was
USDA Misc. Pub. 654, Woody-Plant Seed Manual. The
manuscript was ready for publication in 1941, but World
War II delayed publication until 1948. The boom in tree
planting in the 1950s and 1960s created a large demand for
seeds and exposed the gaps in our knowledge concerning
production and quality of seeds of woody plants in
general.
Realization of this condition led to the revision and considerable expansion of the manual, resulting in publication
of USDA Agric. Handbk. 450, Seeds of Woody Plants in the
United States, in 1974. Seed data were presented for about
800 species, varieties, and sub-species in 188 genera, considerable more than the 420 species and 140 genera in the
1948 edition. The 1974 Handbook proved to be very popular both in this country and abroad, leading to five printings
and translations in several other languages. More than a
quarter-century after its publication, however, numerous
advances in tree seed technology have dictated that a new
revision is needed; the result is the current volume.
The major audience for this book, as for its two predecessors, is those who are involved in the growing and planting of trees and shrubs. Their involvement can be collection
and sale of seeds, production of nursery stock (both bareroot and container), or planting itself. Planting for commercial forest production is the traditional mainstay of tree
planting, but planting for wildlife food, watershed protection, urban environmental improvement, ornamental
enhancement, wetland mitigation, and carbon sequestration
are all on the increase. Ecosystem management, now commonly used in the management of many federal and other
governmental forest lands, has decreased the use of planting
to regenerate the forests and has increased the role of natural regeneration. Those who apply these practices will find
this book useful also in the data on flowering and seed production. Although the book is not intended to be a detailed
textbook on seed ecology and physiology, there is sufficient
scope and depth to the material included to make it useful to
anyone who studies seeds. For additional information on
these topics, readers should consult the recent works by
Baskin and Baskin (2000) and Farmer (1997).
The organization of this book follows that of the earlier
manuals. Part 1 comprises seven chapters that provide general principles on seed biology, genetic improvement, harvesting and conditioning, storage, testing, seed certification,
and nursery practices. The chapter on genetic improvement
combines two chapters from the 1974 Handbook but does
not include the extensive technical information provided in
1974. Genetic improvement of tree and shrub species is now
too common and widespread to be covered adequately in a
chapter in a seed manual. For complete treatments on this
subject, readers are referred to Zobel and Talbert (1984). In
Introduction
(seeds per square meter and seeds per square foot ). A table
of conversion factors is again included before the glossary
(page 1193).
The vast majority of the line drawings and photographs
used in the 1974 Handbook have been used again in this
volume. For new species, efforts were made to collect specimens for new photographs that are similar in style and
background to those of the 1974 Handbook. In most cases
this was done, but samples were not available for every new
species. As a result, line drawings from other publications
were utilized.
The glossary has been expanded by several dozen
terms; a few have been dropped. Others have been altered
to reflect current usage. Most terms of seed biology and
seed technology have been defined according to the glossary of the IUFRO Working Party S2.01.06 Seed
Problems (Bonner 1984).
Those who use this book should realize that much new
information on seeds of woody plants has appeared in print
since the current chapters were finalized. Efforts to improve
the utilization of our forest and range resources continue,
and new information is constantly discovered and put to
use. Like the seed manuals published before, this one will
not be the last, although the next revision may be a digital
data base. Until another revision takes place, however,
genus chapters will be periodically updated with new information on the website that was established during the
current revision: www.nsl.fs.fed.us.
References
Baskin CC, Baskin JM. 1998. Seeds: ecology, biogeography, and evolution of
dormancy and germination.. New York: Academic Press. 672 p.
Bonner FT. 1984. Glossary of seed germination terms for tree seed
workers. Gen.Tech. Rep. SO-49. New Orleans: USDA Forest Service,
Southern Forest Experiment Station. 4 p.
Bramlett DL, Askew GR, Blush TD, Bridgwater FE, Jett JB, eds. 1993.
Advances in pollen management. Agric. Handbk. 698. Washington, DC:
USDA Forest Service. 101 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation; from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Duryea ML, Dougherty PM, eds. 1991. Forest regeneration manual.
Dordrecht,The Netherlands: Kluwer Academic Publishers. 433 p.
Farmer RE Jr. 1997. Seed ecophysiology of temperate and boreal zone
forest trees. Delray Beach, FL: St. Lucie Press. 253 p.
Farr DF, Bills GF, Chamuris GP, Rossman AY. 1989. Fungi on plants and plant
products in the United States. St. Paul: American Phytopathological
Society. 1252 p.
GRIN [Germplasm Resources Information Network]. 2004. GRIN [available
at www.ars-grin.gov]. Beltsville, MD: USDA ARS, National Germplasm
Resources Laboratory.
Khullar P, Thapliyal RC, Beniwal BS,Vakshasya RK, Sharma A. 1991. Forest
seed. Dehra Dun, India: Indian Council of Forestry Research & Education.
409 p.
Landis TD,Tinus RW, McDonald SE, Barnett JP. 1990B95. The
container tree nursery manual,Volumes 25. Agric. Handbk. 674.
Washington, DC: USDA Forest Service. 523 p.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan. 1290 p.
Liegel LH,Venator CR. 1987. A technical guide for forest nursery management in the Caribbean and Latin America. Gen.Tech. Rep. SO-87. New
Orleans: USDA Forest Service, Southern Forest Experiment Station.
98 p.
Ng FSP. 1992. Manual of forest fruits, seeds, and seedlings.Volumes 1 and 2.
Kepong, Malaysia: Forest Research Institute of Malaysia. 997 p.
PLANTS. 2004. The PLANTS database [available at
http://plants.usda.gov/plants]. Baton Rouge, LA: USDA NRCS, National
Plant Data Center.
Rose R, Chachulski CE, Haase DL. 1998. Propagation of Pacific Northwest
native plants. Corvallis: Oregon State University Press. 248 p.
Schmidt L. 2000. Guide to handling of tropical and subtropical forest seed.
Humlebaek, Denmark: Danida Forest Seed Centre. 511 p.
Tompsett PB, Kemp R. (comp.) 1996. Database of tropical tree seed
research (DABATTS). Richmond, Surrey, UK: Royal Botanic Gardens
Kew. 263 p.
Vozzo JA, ed. 2003. Tropical tree seed manual. Agric. Handb. 721.
Washington, DC: USDA Forest Service. 899 p.
Williams RD, Hanks SH. 1976 [slightly revised 1994]. Hardwood nursery
guide. Agric. Handbk. 473. Washington, DC: USDA Forest Service. 78 p.
Zobel B,Talbert J. 1984. Applied forest tree improvement. New York: John
Wiley & Sons. 505 p.
Invasives
We remind our readers that listing species in this book
is not necessarily a recommendation to use them! Many
species that earlier in the twentieth century were recommended and even planted by various federal and state agencies as erosion control and wildlife plantsmultiflora rose,
autumn-olive, and Russian-olive, for exampleare now
considered invasive non-natives and are targets of eradica-
tion campaigns! Other plants that have escaped from horticultural usesbroom, burning bush, oriental bittersweet,
Japanese barberry, etc.are also members of this infamous
company. Finally, there are invasive plants such as ailanthus
and royal paulownia that arrived here accidentally. We sincerely hope that the information contained in this new manual may be of help in efforts to control woody invasives.
Introduction
vii
viii
Part 2
A
Abies 149
Acacia 199
Acer 204
Adenanthera 217
Aesculus 219
Ailanthus 224
Albizia (see also Pithecellobium) 227
Aleurites (see also Vernicia) 230
Alnus 232
Ambrosia 243
Amelanchier 245
Amorpha 250
Aralia 255
Araucaria 259
Arbutus 263
Arctostaphylos 266
Aronia 271
Artemisia 274
Asimina 281
Atriplex 283
B
Baccharis 291
Bauhinia 295
Berberis (see also Mahonia) 298
Betula 303
Bumelia (see Sideroxylon) 1047
530
G
Garrya 547
Gaultheria 550
Gaylussacia 557
Ginkgo 559
Gleditsia 562
Gordonia 565
Grayia (see also Zuckia)
Grevillea 573
Gutierrezia 575
Gymnocladus 578
567
H
Halesia 580
Hamamelis 582
Haplopappus (see Ericameria)
Heteromeles 585
Hippophae 588
Holodiscus 591
Hymenaea 595
497
D
Delonix 468
Dendromecon 470
Diospyros 472
Dirca 476
C
Callicarpa 311
Calocedrus 313
Campsis 319
Caragana 321
Carnegiea 324
Carpenteria 326
Carpinus 328
Carya 333
Castanea 338
Castanopsis (see Chrysolepis)
Casuarina 342
Catalpa 345
Ceanothus 348
Cedrus 357
Celastrus 363
Celtis 366
Cephalanthus 369
Ceratonia 371
Cercis 374
I
Ilex
J
Juglans 601
Juniperus 607
E
Ebenopsis (see also Pithecellobium)
482
Elaeagnus 484
Encelia 488
Enterolobium 490
Ephedra 492
Epigaea 495
Ericameria 497
Eriogonum 499
Eucalyptus 504
Euonymus 513
Eurotia (see Krasheninnikovia)
626
404
597
F
Fagus 520
Fallugia 525
Flindersia 528
K
Kalmia 615
Kalopanax 618
Kochia 620
Koelreuteria 624
Krascheninnikovia 626
L
Laburnum 631
Lagerstroemia 634
Larix 637
Larrea 651
Ledum 653
Lespedeza 655
Leucaena 658
Leucothoe 661
Introduction
ix
Picea 793
Pieris 807
Pinus 809
Pithecellobium (see also Albizia) 848
Platanus 850
Platycladus (see also Thuja) 854
Populus 856
Prosopis 872
Prunus 875
Pseudotsuga 891
Psorothamnus 907
Ptelea 910
Pterocarpus 913
Purshia 916
Pyrus 922
313
M
Maclura 697
Magnolia 700
Mahonia (see also Berberis)
Malosma (see Rhus) 954
Malus 712
Melia 718
Menispermum 720
Menodora 721
Metasequoia 723
Mitchella 726
Morella (see Myrica) 733
Morus 728
Myrica 733
706
Quercus
928
939
U
Ulex 1140
Ulmus 1143
Umbellularia 1150
V
Vaccinium 1154
Vernicia (see also Aleurites)
Viburnum 1162
Vitex 1168
Vitis 1171
749
P
Paraserianthes (see also Albizia)
764
Parkinsonia 766
Parthenocissus 769
Paulownia 772
Penstemon 774
Peraphyllum 778
Persea 781
Phellodendron 783
Philadelphus 786
Photinia (see Heteromeles) 585
Physocarpus 790
Woody Plant Seed Manual
Tamarix 1087
Taxodium 1089
Taxus 1092
Tectona 1099
Tetradymia 1102
Thespesia 1105
Thuja (see also Platycladus) 1108
Tilia 1113
Toona 1119
Torreya 1121
Toxicodendron (see Rhus) 954
Triadica 1125
Tristania (see Lophestemon) 689
Tsuga 1127
Nama 738
Nandina 740
Nemopanthus 743
Nyssa 745
Oemleria 749
Olea 753
Olneya 757
Osmaronia (see Oemleria)
Ostrya 759
Oxydendrum 761
Spiraea 1067
Styrax 1071
Swietenia 1075
Symphoricarpos 1078
Syringa 1083
Sabal 997
Salix 1000
Salvia 1010
Sambucus 1014
Sapindus 1019
Sapium (see Triadica) 1125
Sarcobatus 1022
Sassafras 1025
Schinus 1027
Sciadopitys 1030
Senna 1032
Sequoia 1034
Sequoiadendron 1037
Serenoa 1039
Shepherdia 1043
Sideroxylon 1047
Simmondsia 1049
Solanum 1052
Sophora 1055
Sorbaria 1057
Sorbus 1059
Spathodea 1065
W
Washingtonia
1160
1173
Y
Yucca
1175
Z
Zamia 1178
Zanthoxylum 1180
Ziziphus 1183
Zuckia (see also Grayia)
1185
Part 1
Principles and General Methods of
Producing and Handling Seeds
Chapter 1
Seed Biology
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Contents
Introduction 4
Flowering Plants 4
Reproductive Cycles 4
Flowering 4
Initiation 5
Phenology 5
Influencing factors 6
Temperature 6
Light 6
Moisture 6
Nutrition 7
Physiology 7
Manipulation of flowering 7
Juvenile Phase 7
Mature Phase 7
Structure and Development 8
Pollination 10
Pollen grain development 10
Pollen dispersal 11
Pollen viability and flower receptivity 11
Pollination in angiosperms 11
Pollination in gymnosperms 12
Fertilization 12
Reproductive Abnormalities 12
Polyembryony 12
Parthenocarpy 12
Agamospermy 13
Fruit and Seed Development 13
Morphological Development 13
Angiosperms 13
Gymnosperms 14
Physiological Development 16
Moisture content 16
Stored food reserves 17
Hormones 18
Factors That Influence Seed Production 20
Physiological factors 20
Weather 20
Biotic factor 20
Insects 20
Pathogens 21
Birds 22
Mammals 22
Maturity and Dispersal 23
Indices of maturity 23
Physical 23
Chemical 24
Shedding and dispersal 24
Dormancy 25
Types of Dormancy 25
Seedcoat (or external) dormancy 25
Embryo (or internal) dormancy 25
Morphological dormancy 26
Combined dormancy 26
Double dormacy 26
Secondary dormancy 26
Overcoming Dormancy 26
Seedcoat dormancy 26
Cold water soak 26
Hot water soak 26
Hot wire 26
Acid treatment 27
Mechanical treatments 27
Internal dormancy 27
Stratification (chilling) 27
Incubation and stratification 27
Chemical treatment 27
Combined treatments 27
Variation in Dormacy 28
Germination 28
Environmental Factors 28
Moisture 28
Temperature 29
Light 29
Aeration 30
Biochemical Changes 30
Physical Development 31
References 32
Introduction
Reproductive Cycles
Flowering
Flowering Plants
Initiation
In numerous woody plants, flower initiation and development is a lengthy process extending over several months.
During this period, environmental factors and the internal
physiological condition of the trees interact to produce the
flower crops. The effects of some environmental factors
have been observed through the years, and these relationships have been used to influence flowering and seed production in some species (see below). The internal factors
involved are still poorly understood, as are their interactions
with the environment.
Phenology. Flower buds on most trees and shrubs of
the temperate regions are initiated late in the growing season of the year preceding flowering (table 1). In species
with unisexual flowers, male flowers may start earlier and
differentiate more rapidly as well. Flowers may bloom from
late winter to fall, depending on the species and the location. In temperate trees, flowering is primarily seasonal, that
is, production only occurs in certain times of the year. Most
species bloom in the spring, but there are numerous exceptions to this rule. Witch-hazel (Hamamelis virginiana L.)
flowers from September to mid-November; California-laurel
(Umbellularia californica (Hook & Arn.) Nutt.) from
December to May; September elm (Ulmus serotina Sarg.) in
September; and deodar cedar (Cedrus deodara (Roxb.)
Loud.) in September to October. The times reported in this
book for flowering are typically expressed as a range of several months to allow for the latitudinal and elevational differences throughout the range of a given species. Local variations in weather may also affect the time of flowering from
year to year on the same tree.
Table 1Chapter 1, Seed Biology: times of flower initiation in selected species as determined from microscopic
examination of buds
Time of initiation
Female
Species
Location
Male
Acer pseudoplatanus L.
Betula papyrifera Marsh.
Carya illinoensis (Wangenh.) K. Koch
Larix occidentalis Nutt.
Picea glauca (Moench) Voss
Pinus elliottii Engelm.
P. monticola Dougl. ex D. Don
Populus tremuloides Michx.
Pseudotsuga menziesii (Mirb.) Franco
Taxodium distichum (L.) Rich.
Thuja plicata Donn ex D. Don
Tsuga heterophylla (Raf.) Sarg.
Indiana
NW Ontario
Georgia
British Columbia
Ontario
Florida
British Columbia
Connecticut
Oregon
Florida
British Columbia
British Columbia
June
Early May
May
June
Early Aug
Late JuneJuly
Late JuneAug
Early July
Apr
Early June
June
June
Late Juneearly July
Mar
June
Early Aug
Late Aug
Mid-Aug
Late June
Apr
Aug
July
July
Sources: Anderson and Guard (1964), Fraser (1962), Lester (1963), Macdonald and Mothersill (1987), Mergen and Koerting (1957), Owens and Molder (1974, 1977,
1979), Owens and Pharis (1971), Owens and Smith (1964),Takaso and Tomlinson (1990),Wetzstein and Sparks (1983, 1984).
Figure 1Chapter 1, Seed Biology: structure of a complete angiosperm flower (from Krugman and others 1974).
cones) without calyx, corolla, stamens, or pistils. These strobili characteristically have a central axis bearing a few to
numerous distinctly shaped scales and bracts (figure 2). In
staminate strobili, each scale (microsporophyll) bears 2
pollen sacs (microsporangia) on its lower surface. In ovulate
strobili, 2 inverted ovules (megasporangia) form on the
upper surface of each ovulate scale. Staminate strobili
often bright shades of yellow, red, or purple when fully
developedare numerous, short-lived, and highly productive of pollen. The less numerous, but infrequently colorful,
ovulate strobili develop into woody, relatively durable structures (cones) that contain a varying number of seeds.
Coniferous strobili are similarly arranged around the
central axes of cones (figure 2). Flowers of angiosperms, on
the other hand, have varied and distinctive floral arrangements. Some species bear a single flower on each peduncle,
for example, magnolia and tuliptree, but most others bear
flowers in groups or clusters called inflorescences. The general structure of an inflorescence is a central stem, with or
without branches, on which flowers, with or without
pedicels, develop. Examples of the common forms of inflorescences of woody plants (figure 3) include catkin (ament),
birch; raceme, serviceberry; spike, walnut (pistillate); head,
sycamore (Platanus L.); cyme, viburnum; panicle, sumac
(Rhus L.); and umbel, plum.
10
Weather conditions have a strong influence on pollination. Dry, warm weather will usually enhance pollen dispersal by wind. If winds are excessively dry, however, pollen of
white oaks may be shed before maturity (Sharp and
Chisman 1961). In contrast, rain or high humidity greatly
hinders anemophilous pollination. Complete seedcrop failures can occur locally if heavy rains dominate the weather
when anthesis is occurring. Late spring freezes can also kill
staminate flowers and cones and prevent any dissemination
of pollen in some species. Entomophilous pollination is not
as greatly affected by the weather, but low temperatures and
heavy rains will curtail the activities of insect pollinators.
Anemophilous pollen dispersal depends primarily on
weather factors and stand structures. Under near-calm conditions, pollen of many pines and hardwoods can be expected
to disperse only a few dozen meters (Sedgley and Griffin
1989), but in turbulent wind conditions, dispersal of pine
pollen for 1 km and more is likely (Griffin 1980; Lanner
1966). Entomophilous pollen dispersal distances are not
precisely known but probably are considerably less than
anemophilous dispersals.
Pollen viability and flower receptivity. For pollination to be successful, the pollen grains must remain viable
until they reach the stigma, and the female flowers or cones
must be receptive when the pollen arrives. Not much is
known about the length of viability of pollen in nature;
some pollens survive for only hours and others for weeks.
The pollen of many species, notably conifers, can be carefully dried to below 10% moisture content and stored below
freezing for several years (Copes 1987; Wang and others
1993). Like pollen viability, flower receptivity varies greatly
among species. For angiosperms, the receptive period for an
individual flower may last for less than a day, as noted earlier for southern magnolia (Thien 1974), or it may continue
for up to 10 days in some cherries (Stsser and Anvari
1982). Among gymnosperms, the receptive period ranges
from less than a day in japanese larch, Larix kaempferi
(Lam.) Carr. (Villar and others 1984) to 2 weeks or more in
true firs, hemlocks, and pines (Owens and Blake 1985).
Pollination in angiosperms. When pollen grains
reach the stigma of a receptive flower and germinate, the
pollination process is set into motion. Pollen grains are captured on the stigmas due to their own sticky surface characteristics or the nature of the stigma surface. The stigma
surface is naturally dry in some genera (maple; dogwood;
sweetgum, Liquidambar L.; elderberry, Sambucus L.; and
basswood, Tilia L.) and wet in others (hickory, eucalyptus,
holly, plum, and serviceberry) (Sedgley and Griffin 1989),
but there are no strong correlations between surface condition and other aspects of pollination and fertilization. Some
have suggested that pollen germination rates are quicker on
dry stigmas, but evidence for this is weak (Owens 1992).
Germination is rapid, usually occurring within a few hours
(Owens 1992), and is temperature-dependent. Luza and oth-
11
13
In most species, the perisperm, comprised of the maternal nucellar tissue in the ovule, fails to develop and is
absorbed by the developing embryo. In a few species, the
perisperm develops into a food storage tissue that is outside
of the embryo sac. In these cases there is no endosperm
development, and the perisperm becomes the major food
storage tissue. In this book, yucca is the only genus with a
fully developed perisperm.
Embryos differentiate and attain their full size in most
species by the time the fruits or seeds are shed. In fact, relative length of the embryo can be a good maturity index for
decisions on when to collect seeds of many species. In a few
species, howeverfor example, American holly (Ilex opaca
Ait.), European ash (Fraxinus excelsior L.), and common
snowberry (Symphoricarpos albus (L.) Blakeembryos are
still immature when seeds are shed from the trees, and full
size is only attained following a period of after-ripening.
This condition causes the very slow germination that is a
major problem in nursery production of these species.
As endosperm and embryo grow, the surrounding maternal tissues develop into the seed-covering structures, collectively called the seedcoat. Most seedcoats are composed of a
firm outer layer, the testa, and a generally thin, membranous
inner coat called the tegmen. There are many variations of
seedcoat structure, however, and many species do not fit the
model described above. In some genera, the testa is thin and
permeable, as in poplar (Populus L.) and willow. In others, it
may be thick and bony, as in hawthorn and apple. Some
hard seedcoats have special cutinized layers, as in redbud
and honeylocust (Gleditsia triacanthos L.). In some genera
both covering structures are membranous, as in elm; the
outer layer partially membranous and the inner one bony, as
in chastetree (Vitex L.); or the outer layer soft and fleshy and
the inner layer hard, as in magnolia.
Many species develop extended tissues on their seedcoats that play a role in dissemination of the seeds. These
extensions may be wings, as in ailanthus, Ailanthus altissima (Mill.) Swingle, and tuliptree; tufts of short, bristly hairs,
as in baccharis, Baccharis L., and sycamore; long soft hairs,
as in poplar and willow; wings with hairs, as in catalpa and
desertwillow, Chilopsis linearis (Cav.) Sweet; or various
other appendages, such as small points on sourwood
(Oxydendron arboreum (L.) DC.; and long, feathery styles
on cercocarpus, Cercocarpus H.B.K. Actually, the
appendages on baccharis, sycamore, and cercocarpus are on
the fruits, which are single-seeded achenes, commonly
called seeds.
It is useful to define and classify fruit types, although all
authorities do not completely agree on the results. The clas-
14
FROM INFLORESCENCES
or within their seedcoats: true fir, hemlock, and incensecedar (Calocedrus Endl.). The resin makes seeds sticky and
more difficult to handle in all phases of extraction and
cleaning. Most gymnosperm seeds are winged, but there are
exceptions: baldcypress, yew, torreya, and some pines.
These pines are often called the nut pines: Swiss stone
pine, piyon (Pinus edulis Engelm.), chilgoza pine (P. geradiana Wall.), etc. Wings may be loosely adhering structures
that are easily separated from the seeds, as in most pines, or
they may be integral parts of the seedcoat, as in Douglas-fir,
longleaf pine, and incense-cedar.
Cones of gymnosperms that require more than 1 year to
mature generally remain small during the first year after
flowering in the interval between pollination and fertilization. In a few species, such as western juniper (Juniperus
occidentalis Hook) and Alaska-cedar, the fruit grows before
fertilization occurs and attains almost full size during the
first growing season, a full year or more before the seeds are
physiologically mature. Seed collectors must be aware of
this condition to avoid collecting cones with immature
fruits. In Alaska-cedar, there are distinct color differences
between immature and mature cones (Harris 1990), and
position of cones on the branches is an indicator for both
species. Gymnosperm fruit classification is much simpler
than that of angiosperms (table 3) (modified from Krugman
and others 1974).
15
STROBILI
Cone
FLESHY STROBILI
Drupelike
Woody structures that generally open on the trees and release seeds at maturity, as in Abies, Picea, and
most Pinus; some Pinus cones remain closed at maturity and open only in fires or disintegrate over time
Enclosing a single seed, as in Ginkgo,Taxus,Torreya, and some Juniperus, or multiple seeds in
other Juniperus, that are shed from trees intact
Premature cone shedding can also be important in gymnosperms. It is most common several weeks after anthesis,
when pollination has not occurred, but can also result from
damage from frost, hail, drought, insects, or pathogens
(Owens and Blake 1985; Sedgley and Griffin 1989; Sweet
1973). There are also losses from what Bramlett (1972)
described as physiological drop, when there were no visible signs of external injury. In general, the physiology of
immature cone abscission is much less understood than premature fruit shedding in angiosperms (Sedgley and Griffin
1989).
Physiological Development
The growth of fruits that starts soon after fertilization (or
prior to fertilization in a few species) involves a complex
array of physiological processes and conditions. These
processes are generally similar for fruits of most temperate
trees, and they produce comparable trends in size, weight,
and moisture content. A typical pattern of development for
dry fruits is provided by the single-seeded samaras of green
ash (Fraxinus pennsylvanica Marsh.) (figure 6). Fresh
weight, dry weight, and moisture content increase slowly
through early summer. By the end of August, the embryo is
2 to 3 mm long. Over the next 6 weeks there are sharp
increases in dry weight and significant decreases in moisture
as embryo length increases 5-fold (Bonner 1973). In drupes
of the temperate zone, weight trends are similar, but moisture changes are somewhat different. In black cherry
(Prunus serotina Ehrh.), for example, moisture contents
decrease during spring to early summer, then increase again
as maturity approaches (figure 7). In temperate recalcitrant
seeds, such as acorns, the patterns are more like those of dry
fruits (figure 8). Similar trends occur in the maturation of
most tropical tree fruits also, but the changes are not always
correlated with the seasons as they are in temperate species.
Moisture content. Any discussion of seed moisture
must be based upon the 2 physiological classes of seeds in
16
tion of orthodox seeds of woody plants. Conditions are different in orthodox seeds from fleshy fruits, however, as they
are shed before complete desiccation. Desiccation occurs
later after the fleshy covering has dried or been removed
(eaten in many cases). Many of these species have complex
dormancy periods, and it can be hypothesized that there are
interactions between the dormancy and the delay in maturation drying of the seeds.
In recalcitrant seeds, there is no pronounced maturation
drying stage, because development never stops completely.
There are slight decreases in moisture content that are
apparently associated with shedding of fruits (figure 8), but
there is no true quiescent period with recalcitrant seeds.
Most species, especially tropical recalcitrant species, germinate soon after shedding, and some, including several
Quercus species, will germinate while still on the tree, an
event defined as vivipary.
Stored food reserves. As postfertilization growth proceeds, carbon fixed by photosynthesis is transferred to the
seeds in the form of sucrose. In the seeds the sucrose is
converted into many components, but most of it goes into
stored food reserves of carbohydrate, lipid, or protein
(Bewley and Black 1994). Many seeds have more than one
type of food reserve, but one is usually predominant (table
4). The type of food reserve has implications for seed storage (see chapter 4), and it has been suggested that there are
other important relationships. Korstian (1927), for example,
suggested that dormancy in the black oak group was related
to the high lipid content of these seeds, and that stratifica-
many enzyme systems, including those required for desiccation tolerance and germination when rehydration occurs
(Bewley and Black 1994). There are some data for tree
seeds (Finch-Savage and others 1994), but most of the work
in this area has been done on castor bean (Ricinis communis
L.) (Kermode and Bewley 1985) and cereal grains (Bewley
and Black 1994). There is no reason to doubt, however, that
the same physiological processes take place during matura-
17
Table 4Chapter 1, Seed Biology: some characteristic stored food reserves in tree seeds
(expressed as % of dry weight)
Species
Tissue
Seed
Samara
Seed
Husked fruit
Fruit
Seed
Cone
Seed
Seed
Seed
Seed
Seed
Fruit
Seed
Acorn
Acorn
Acorn
Seed
Fruit
Seed
Carbohydrate
Lipid
Protein*
41.2
42.9
13.0
18.3
10.6
79.8
11.6
3.1
2.3
2.9
20.8
5.1
46.6
25.8
67.1
12.3
13.6
8.9
37.6
1.5
1.9
37.4
20.5
36.2
6.8
26.2
44.2
28.1
20.5
18.5
4.9
37.2
2.9
20.3
20.8
9.0
46.6
15.3
13.9
17.0
8.2
5.9
4.0
12.6
5.6
25.3
23.8
24.4
21.9
13.8
7.8
4.6
3.8
6.6
38.7
17.1
27.4
Sources: Barnett (1976a), Bennett (1966), Bonner (1971, 1974a), Ching (1963), Pulliainen and Lajunen (1984),Waino and Forbes (1941).
* Most values obtained by multiplying total N by 6.25.
Total sugars only.
Insoluble N only multiplied by 6.25.
Figure 11Chapter 1, Seed Biology: changes in solublenitrogen (solid circles) and protein-nitrogen (open triangles) contents of maturing seeds of sweetgum
(Liquidambar styraciflua L.) (adapted from Bonner 1972).
19
Table 5Chapter 1, Seed Biology: some characteristic seed elemental compositions (expressed as % of dry weight)
Species
Acer rubrum L.
Callicarpa americana L.
Corylus avellana L.
Ilex vomitoria Ait.
Juglans regia L.
Picea abies (L.) Karst
Pinus sylvestris L.
Prunus serotina Ehrh.
Quercus pagoda Raf.
Q. stellata Wangenh.
Sassafras albidum (Nutt.) Nees
Ulmus alata Michx.
Vaccinium arboreum Marsh.
Tissue
Samara
Fruit
Seed
Fruit
Nut
Seed
Seed
Fruit
Acorn
Acorn
Fruit
Seed
Fruit
Ca
0.34
.26
.10
.24
.08
.02
.04
.14
.27
.25
.06
.51
.33
1.34
.73
1.25
.45
.79
.63
Mg
0.23
.19
.17
.31
.30
.09
.06
.06
.11
.20
.07
P
0.34
.13
.40
.11
.41
.66
.73
.14
.06
.08
.23
.52
.06
Sources: Bonner (1971, 1974a), Hastings (1966), Lott and Buttrose (19780, Pulliainen and Lajunen (1984).
20
Fungal pathogen
Species infected
21
as well. It can be found on larch, spruce, pine, and Douglasfir (Sutherland and others 1987). Pitch canker damages
shoots, cones, and seeds of pines in the South and East. In a
few short years, pitch canker has become a major disease
problem in seed orchards of all southern pines (BarrowsBroaddus and Dwinell 1985; Blakeslee and others 1980).
With the exception of species that attack trees with edible nuts, such as scab diseaseCladosporium caryigenum
(Ell. & Lang.) Gottwaldon pecan (Graves and others
1989), reduction of seedcrops in angiosperms by fungi is
generally not serious. There are, however, numerous fungi
that infect flowers and fruits and cause only incidental or
local damage to the seedcrop (table 7). For additional information on seed pathogens and other microorganisms and the
species on which they are found, readers are referred to
Mittal and others (1990).
Birds. Birds feed on flowers, fruits, and seeds, especially the latter. Many small birdssuch as finches, grosbeaks, and sparrows (Fringillidae), doves (Columbidae), and
quail (Phasianidae)feed on small seeds after they are shed,
but these losses are incidental to the total seedcrop. Larger
birds that feed on maturing fruits and seeds still on the trees
can have serious, though usually local, impacts on seed
yield. Acorns are a favorite of grackles (Quiscalus spp.),
jays (Corvidae), and woodpeckers (Picidae). The California
woodpecker (Balanosphyra formicivora) can devour enough
acorns, its favorite food, to severely reduce the crop within
its foraging range (Bent 1939). Pine seeds are a favorite of
Clarks nutcracker (Nucifraga columbiana) and piyon jays
(Cyanocephalus cyanocephalus), which specialize in piyon
seeds and even young cones (Bent 1946). Berries of various
juniper species are eaten in large numbers by jays, Clarks
fungi that cause minor or locally severe decreases to fruit crops of angiosperms
Fungus
Tissue attacked
Species infected
Botrytis spp
Ciboria acerina Whetz. & Buchew.
Coniothyrium spp.
Cytospora spp.
Gymnosporangium clavipes (Cooke & Peck) Cooke.& Peck
G. clavariiforme (Pers.) DC
Taphrina johansonii Sadeb.
T. occidentalis W.W. Ray
T. amentorum (Sadeb.) Rostr.
Flowers
Flowers
Seeds
Fruits
Fruits
Fruits
Catkins
Catkins
Catkins
Ilex opaca
Acer rubrum, A. saccharinum
Betula alleghaniensis
Prunus serotina
Amelanchier, Cotoneaster, Crataegus, Malus, Pyrus
Amelanchier, Cotoneaster, Crataegus, Malus, Pyrus
Populus spp.
Alnus spp.
Alnus spp.
22
Specific gravity
0.90
0.95
0.95
1.00
0.90
0.90
0.90
1.00
23
(Barnett 1979). Cone weight is estimated by water displacement of the floating cone, and volume is estimated by water
displacement of the submerged cone. Specific gravity is
equal to weight divided by volume (examples of cone specific gravities used to judge maturity are listed in table 8).
Other physical indices of seed maturity are easy cup
release from acorns of oak; a white, brittle embryo of some
ash species that breaks when bent at a sharp angle; and
white pine cone scales that flex open when cones are bent
double. For details on maturity indices of individual genera
or species, see part 2 of this book.
Chemical. Although chemical indices of maturity are
biologically sound, they are seldom practical to use in collection. Most potential chemical indicators are based on the
level of stored food reserves (table 9), but elemental phosphorus and IAA concentrations have been suggested as
indices for green ash (Bonner 1973) and English oak
(Michalski 1969), respectively.
Shedding and dispersal. The majority of temperate
genera shed their fruits and seeds in the fall or winter,
although manyfor example, birch and poplarshed theirs
in the spring. In some generafor example, maple, eucalyptus, willow, and elmthere are both spring-shedding and
fall-shedding species. Other species have seeds that mature
and are shed in mid-summerfor example, ceanothus
(Ceanothus L.).
The seeds of many species are shed or dispersed quickly
(within a few days) after they mature, and collectors must be
alert to the phenological characteristics of the species in
order to collect what they need. Some species that shed
fruits quickly when they mature are maples and elms. In
others, the fruits are persistent on the tree but open to disperse the seeds quickly after maturity; examples include
sweetgum, poplars, and willows. In still other species, fruit
opening and seed dispersal are very dependent on the weather. Cones of loblolly pine, for example, open readily in
warm, dry conditions and disperse their seeds. At night, they
close back up again when humidity rises. If a weather front
brings rain, the cones may close up completely and not
reopen for dispersal for several days. The primary seed dispersal agent of all of the above species is wind.
Drupes, berries, and other fleshy fruits are not usually
shed quickly, but they can be removed from the trees rapidly
by birds and animals. This can be a major problem for seed
collectors wishing to harvest the seeds of species such as
pawpaws (Asimina Adans.), hollies, plums, and prickly-ash
(Zanthoxylum L.). Seeds will usually have to be collected
exactly at the time of maturity on the trees, or the entire
crop may be lost. The same problem occurs for some fruits
that are not fleshy, for example, hickories, walnuts, and
oaks. These fruits are favorite foods of rodents, deer, and
other animals, and they must be collected from the ground
as soon as they are shed. Birds will also take many of these
fruits before shedding; for example, a flock of grackles can
completely strip a large willow oak (Q. phellos L.) of its
acorn crop in several hours.
The cones of most conifers disperse their seeds soon
after maturity. In true firs, dispersal occurs as the cone disintegrates on the trees, leaving the spike-like cone axis still
upright on the branches. In some pines, cedars, and hemlocks, the cones are slow to give up their seeds, and dispersal may take 3 to 12 months. Serotinous cones of several
speciessuch as jack (Pinus banksiana Lamb.), sand
(P. clausa (Chapm. ex Engelm.) Vasey ex Sarg.), pitch (P.
rigida Mill.), and lodgepole pinesdo not normally open on
the trees but open on the ground following fires that melt the
resin seals on the cone scales. Other pinesSwiss stone and
Siberian stone pines, etc.shed their cones while still
Species
Chemical
Crude fat
Crude fat
Crude fat
Crude fat
Insoluble CHO
Insoluble CHO
Crude fat
Reducing sugars
Sources: Bonner (1972, 1973, 1974b, 1976, Rediske (1961), Rediske and Nickolson (1965).
CHO=carbohydrates.
24
Dormancy
Once seeds have matured and been dispersed, survival of
the species requires that they germinate at a time and place
favorable for growth and survival of the seedlings. Plants
have evolved many mechanisms and processes that ensure
survival. Some species produce prodigious numbers of
seeds, so that even if only a tiny proportion germinate and
grow, some seedlings will survive. In others, germination at
unfavorable times is prevented by a mechanism that is commonly described as dormancy. Dormancy is defined as a
physiological state in which a seed disposed to germinate
does not, even in the presence of favorable environmental
conditions (Bonner 1984). Seeds are able to overcome dormancy and germinate when triggered by certain internal
processes that are usually induced by environmental
changes. There is a tremendous range in the degree of dormancy among woody species. Some seeds lie in the soil for
years before germinating, whereas other are delayed for only
a few weeks. The latter condition is sometimes described as
delayed germination to indicate something less than true
dormancy. In fact, the distinction between dormancy and
delayed germination is not at all clear, and among the
majority of species, the interval between maturity and germination (in natural conditions) is a continuum with no
distinct gradation.
Types of Dormancy
Many different classifications of dormancy have been
devised by seed scientists and there is no universal agree-
ment on the subject. Most tree seed workers accept the definitions of the Seed Problems Working Party of the IUFRO
International Union of Forest Research Organizations
(Bonner 1984)and these definitions will be used in this
discussion.
Seedcoat (or external) dormancy. Seedcoat dormancy has 3 primary modes of action. In the most common
mode, the seedcoats (or other covering structures) are impermeable to the entry of moisture or gases. Members of the
Leguminoseaefor example, acacia, albizia (Albizia
Durazz.), honeylocust, mesquite (Prosopis L.), black-locust,
sophora (Sophora L.)usually display this characteristic,
which is commonly called hardseededness by those who
work with seeds. Members of other families also have seedcoats that impose a similar dormancy, but seedcoat structures are different; some examples include American beautyberry (Callicarpa americana L.), hollies, sumacs, and
basswood.
The second mode of dormancy action attributed to seedcoats is the mechanical resistance to swelling of the embryo
as it absorbs moisture. This resistance delays full imbibition
and emergence of the radicle from within the seed.
Mechanical resistance frequently contributes to dormancy
and has been documented in big sagebrush (Artemesia tridentata Nutt.) (McDonough and Harniss 1974), pecan (Van
Staden and Dimalla 1976), loblolly pine (Barnett 1976b),
Korean pine (Hatano and Asakawa 1964), and water oak
(Peterson 1983). It does not appear to be the primary factor
in tree seed dormancy, however.
A third possible mode of seedcoat dormancy is the presence of germination inhibitors in the seedcoats (Bewley and
Black 1994; Nord and Van Atta 1960; Peterson 1983) that
may or may not play a significant role in dormancy. Some
of the phenolic substances in seedcoats that could possibly
be germination inhibitors could actually be beneficial by
inhibiting the growth of pathogenic microorganisms
(Mohamed-Yasseen and others 1994). In some herbaceous
species, there are inhibitors that must leach from the embryo
before germination can take place, and seedcoats prevent
this leaching (Bewley and Black 1994). There is no conclusive evidence of this condition in seeds of woody plants, but
success in stratifying seeds by placing them in porous sacks
in running water suggests that it may occur.
Embryo (or internal) dormancy. Embryo dormancy
arises from a condition within the embryo itself. The most
likely cause of embryo dormancy is the presence of germination inhibitors in the embryonic axis or in the food storage
tissues of the seed. For germination to occur, these
inhibitors must be metabolically inactivated, or their effect
25
26
27
Variation in Dormancy
As noted earlier, there is widespread variation in the
degree of dormancy, both among species, and within a
species. For some species, there are patterns of dormancy
that have been documented and that can have practical
application. For example, degree of dormancy appears to
increase with increasing elevation of seed source for black
cherry and red maple (Acer rubrum L.) in Tennessee
(Farmer and Barnett 1972; Farmer and Cunningham 1981).
Seeds from more northern sources generally require longer
stratification periods than seeds from southern sources. This
relationship has been reported for sugar maple (Kriebel and
Gabriel 1969), red maple (Farmer and Goelz 1984;
Tremblay and others 1996), sweetgum (Wilcox 1968), and
sycamore (Webb and Farmer 1968). In contrast, eastern
white pine showed just the reverse in a range-wide study
(Fowler and Dwight 1964): seeds from the southern sources
are more dormant, but this phenomenon may be related to
the higher altitudes of the natural stands of white pine at the
southern extremities of its range. Warmer climates during
seed maturation typically produce heavier and larger
embryos in seeds (Durzan and Chalupa 1968), presumably
because the growing seasons are longer. This conditions
suggests that degree of dormancy (or delayed germination)
is related to degree of physiological maturity in temperate
seeds, but the evidence for this is lacking.
Variation in dormancy among individual trees at the
same site has been documented for loblolly pine (McLemore
and Barnett 1966) and sweetgum (Rink and others 1979),
and can probably be assumed to occur in all woody plants.
Partial genetic control of dormancy is also obvious, because
most seed dormancy is related in some way to seedcoats or
other covering structures, all maternal tissues. The best way
to understand dormancy is to quantify it in mathematical
terms. A number of studies have attempted this for temperate trees (Bonner and Harrington 1993; Donald 1987;
Richter and Switzer 1982; Rink and others 1979; Sorensen
1983) and all of the proposed methods have application
under certain conditions.
Germination
Germination is defined as the resumption of active
growth in an embryo which results in its emergence from
the seed and development of those structures essential to
plant development (Bonner 1984). In another sense, it is
the culminating event of seed maturation, the establishment
of the seedling. It is useful to think of germination as occurring in overlapping events (Kramer and Kozlowski 1979):
28
1. Absorption of water
2. Increased respiration, enzymatic activity, and
assimilation of stored foods
3. Increased adenosine phosphate and nucleic acids
4. Cell growth and division
5. Differentiation of tissues
All of these events are influenced by environmental
conditions and events within the seeds themselves.
Environmental Factors
The most important environmental factors that influence
germination are moisture, temperature, light, and aeration.
Moisture. The typical pattern of moisture uptake by
seeds has 3 phases (Vertucci 1989): a rapid initial uptake, a
short lag period of extremely slow uptake, and another rapid
period of uptake just before germination. The first phase is
primarily imbibitional in nature and occurs in dead seeds as
well as live ones. It is a physical process of moisture moving
from a substance with high water potential (soil) to one with
a low water potential (dry seed). This uptake displaces gases
from dry seeds (Simon 1984) and is visually evident in the
bubbles that slowly escape from dry seeds when they are
submerged in water. The length of the second phase is related to the degree of dormancy or delayed germination in the
seeds. It can be practically absent in the rapidly germinating
seeds of oak (Bonner 1968) or extended in the case of very
dormant seeds. The third phase occurs when metabolism
becomes very active, and the seedcoats split, leading to
greater oxygen uptake.
There are minimum amounts of moisture required for
germination to proceed, and several studies have sought to
measure the moisture stresses that will retard or halt germination (table 10). Significant decreases in germination
occurred, in general, from 0.8 MPa and below, and germination was effectively stopped at stresses of 0.3 to 2.0
MPa. All of these studies used osmotic solutions to impose
stress, and there are concerns that this method may hinder
germination by inhibiting gas exchange. McDonough (1979)
used thermocouple psychrometer chambers to impose moisture stress on seeds of quaking aspen (Populus tremuloides
Michx.), however, and his results agree quite well with those
reported with osmotic solutions. Comparisons among
species should be made with caution, as methodology and
equipment varied widely in these studies. There were also
significant interactions with seed source, seed treatment, and
temperature for some species (Bonner and Farmer 1966;
Farmer and Bonner 1967; Moore and Kidd 1982).
Table 10Chapter 1, Seed Biology: critical levels of water potential (MPa) for germination within a 20 to 30 C range
of temperatures as determined with osmotic solutions*
Water potential (MPa)
Species
Strongly decreased
germination
0.1
0.29
0.4
0.4
0.5
0.8
0.6
0.81
0.81
0.4
0.8
0.1
1.01
0.5
Effectively stopped
germination
1.6
0.51
1.3
1.6
1.52
1.2
1.82
1.82
0.8
0.3
1.52
2.0
Sources: Barnett (1969), Bonner (1968), Bonner & Farmer (1966), Choinski and Tuohy (1991), Djavanshir and Reid (1975), Farmer and Bonner (1967), Kaufman and
Eckard (1977), Moore and Kidd (1982), Sabo and others (1979), Singh and Singh 1983
* Some data were converted from atmospheres and bars to MPa as follows: 1 bar = 0.1 MPa; 1 atm = 0.1013 MPa.
29
Physical Development
Physical changes in germinating seeds are practically the
same for all species. The first sign is usually swelling of the
seed from water uptake. Embryo elongation occurs second,
but unseen within the seeds covering structures. Then the
seedcoat splits, and the emerging radicle elongates. At this
point, germination in temperate species takes one of two
forms. One form is epigeal germination, in which the
hypocotyl elongates, arches upward, then straightens, pushing the cotyledons upward through the soil (figure 12). In
many species the seedcoats are still attached to the cotyledons after emergence and are not shed until the cotyledons
start growing. Genera that exhibit epigeal germination
include pine, cedar, eucalyptus, juniper, magnolia, and
mountain-ash.
In the second form, hypogeal germination, it is the epicotyl that elongates, pushing the young plumule through the
soil while the cotyledons remain below ground (figure 12).
There they remain attached to the seedling and supply
reserve foods for weeks or more. Genera that exhibit
hypogeal germination include buckeye, oak, walnut,
chestnut (Castanea Mill.), and torreya.
Germination form is normally the same for all species in
a genus, but like most things in seed biology of woody
plants, there is an exception. In cherries and plums, both
forms occur; common chokecherry (P. virginiana L.) is
epigeal, but the remaining species of the genus are hypogeal
(figure 12).
Some authorities recognize other forms of germination
in tropical species. Bunya-pine (Araucaria bidwillii Hooker)
and Parana-pine (A. angustifolia (Bert.) O. Kuntze) seeds
germinate on the surface of the soil, then the cotyledonary
stalks elongate and push the hypocotyl, plumule, and radicle
into the soil. The hypocotyl subsequently develops into a
tuber that serves to transfer the food reserves from the
female megagametophyte to the growing seedling. This type
of germination has been defined as cryptogeal (Burrows and
Stockey 1994), and these araucarias are the only species in
this book that exhibit this form of germination. Ng (1991)
also has defined durian germination in which the hypocotyl
elongates but the cotyledons remain within the seed. This
form of germination occurs in common durian (Durio
zibethinus Murr.), a popular, edible fruit of Southeast Asia
that is cauliferous.
31
References
32
Blakeslee GM, Dwinell LD, Anderson RL. 1980. Pitch canker of southern
pines: identification and management considerations. For. Rep. SA-11.
Atlanta: USDA Forest Service, Southeastern Area, State and Private
Forestry. 15 p.
Bonner FT. 1967. Germination of sweetgum seed in response to light.
Journal of Forestry 65: 339.
Bonner FT. 1968. Water uptake and germination of red oak acorns.
Botanical Gazzette 129: 8385.
Bonner FT. 1971. Chemical contents of southern hardwood fruits and
seeds. Res. Note SO-136. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 3 p.
Bonner FT. 1972. Maturation of sweetgum and American sycamore seeds.
Forest Science 18: 223231.
Bonner FT. 1973. Timing collections of samaras of Fraxinus pennsylvanica
Marsh in the southern United States. In: Proceedings, International
Symposium on Seed Processing; 1973 September 47; Bergen, Norway.
Volume 1. Stockholm: Swedish Royal College of Forestry. [not paged].
Bonner FT. 1974a. Chemical components of some southern fruits and
seeds. Res. Note SO-183. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 3 p.
Bonner FT. 1974b. Maturation of acorns of cherrybark, water, and willow
oaks. Forest Science 20: 238242.
Bonner FT. 1975. Maturation of black cherry fruits in central Mississippi.
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the breaking of dormancy in Acer pseudoplatanus L. Journal of
Experimental Botany 24: 741750.
Wetzstein HY, Sparks D. 1983. The morphology of pistillate flower differentiation in pecan. Journal of the American Society for Horticultural
Science 108: 9971003.
Wetzstein HY, Sparks D. 1984. The morphology of staminate flower differentiation in pecan. Journal of the American Society for Horticultural
Science 109: 245252.
Wilcox JR. 1968. Sweetgum seed stratification requirements related to
winter climate at seed source. Forest Science 14: 1619.
Willan RL, compl. 1985. A guide to forest seed handling with special reference to the tropics. For. Pap. 20/2. Rome: FAO. 379 p.
Williams PM, Ross JD, Bradbeer JW. 1973. Studies in seed dormancy: 4.The
abscisic acid content of the seeds and fruits of Corylus avellana L. Planta
110: 303310.
Willson MF, Burley N. 1983. Mate choice in plants. Princeton, NJ: Princeton
University Press. 251 p.
Ziller WG. 1974. The tree rusts of western Canada. Pub. 3129.Victoria, BC:
Canadian Forestry Service. 272 p.
Zobel B,Talbert J. 1984. Applied forest tree improvement. New York: John
Wiley and Sons. 505 p.17.1
Seed biology
37
38
Chapter 2
Contents
Introduction 40
Terminology 40
Allocation of Resources 40
Monoculture 40
Gene Conservation 40
Tree Improvement Versus Crop Improvement 41
The Biology of the Species 41
Concepts of Genetic Improvement 42
Phenotype and Genotype 42
The Genetic Code 43
Chromosome Numbers 43
Selection 43
Hybridization 43
Testing for Breeding Value 44
Screening for Fusiform Rust Resistance 44
Screening for White Pine Blister Rust Resistance 44
Advanced Generation Breeding 45
Starting a Tree Improvement Program 45
Establishing Objectives 45
Identifying the Raw Material To Be Used 45
Native versus exotic species 45
Successful introductions of exotics 45
Land races 45
Geographic variation 46
Utilizing the Raw Material 47
Seed production areas 47
Clonal seed orchards 48
50
39
Introduction
In this chapter, readers can gain a basic understanding of
why certain procedures are used to improve forest trees.
The references listed can be used as a source for obtaining
more detailed information, both historical background and
technical details. Examples are frequently cited from operational tree improvement programs to focus the chapter on an
applied level. Although many of the examples are taken
from the authors professional focus of over 40 years in the
southern United States, the principles they exemplify can be
used world-wide.
Terminology
Many of the terms used in this chapter are listed in the
Glossary; a more detailed glossary may be found in Snyder
(1972) and Wright (1976). Comprehensive references on
forest genetics include Dorman (1976), Wright (1976) and
Zobel and Talbert (1984).
Forest genetics is the general term often used for the
study of inheritance in forest trees, whereas forest tree
improvement usually refers to the applied use of forest
genetics to actually improve the quality of the trees. Tree
breeding is often used as a synonym for tree improvement,
but it also may be found referring to specific activities such
as controlled pollination. Zobel and Talbert (1984) define
forest tree breeding as activities geared to solve some specific problem or to produce a specifically desired product.
Tree improvement will be the term used most frequently in
this chapter.
It is important to understand that tree improvement is an
integral part of silviculture. Tree improvement provides the
raw material for artificial regeneration, which is one of the
most important weapons in the arsenal of the silviculturist.
Tree improvement provides a direct avenue to introduce
genetically improved seedlings (or cuttings) into the reforestation system with no additional handling fees. It costs
no more to plant a genetically improved seedling than a
woods-run seedling. (Note that although the costs of producing genetically improved planting stock are not insignificant, they can be viewed as an investment in future
increased productivity. Dividends accrue in terms of
increased growth, better form and wood quality, and
improved insect and disease resistance).
Allocation of Resources
One of the key elements of land management is allocation of resources. An ever-expanding world population
demands an ever-increasing supply of wood products. These
40
Field crops
Trees
12 months
46 months
12 years
520 years
10100 years
820 years
Chapter 2: Introduction
41
42
Genus
Juniperus, Nyssa, Sequoia, Thuja
Abies, Larix, Picea, Pinus, Quercus, Tsuga
Acer
Betula
Liquidambar
Carya, Juglans
Liriodendron, Populus, Salix
Platanus
Fraxinus
43
that they have little value as timber trees. There are two possible approaches to this problem. One is genetic engineering; the other is back-crossing to pure American chestnut.
The American Chestnut Foundation has produced many successful back-crosses with the potential of restoring this
grand tree to its former dominance in the eastern hardwood
forest.
Many tree species that coexist on the same sites maintain their status as separate species primarily by a separation
of flowering time. On transitional sites (ecotones) when one
species is accelerated or retarded in flowering time, hybrids
often result as in the case of the Coulter pine (Pinus coulteri
D. Don) x Jeffrey pine (Pinus jeffreyi Grev. & Balf.) hybrids
in California (Zobel 1951) and the pond pine (Pinus serotina
Michx.) x loblolly pine (P. taeda L.) mixtures in North
Carolina (Saylor and Kang 1973). Hybridization often
occurs near the edge of the range where the species is losing
its adaptive advantage. In southeastern Oklahoma and northeastern Texas, shortleaf (P. echinata P. Mill.) and loblolly
pines occupy many sites together and hybrids are not
uncommon (Abbott 1974).
Natural hybridization is a common phenomenon among
the oaks (Quercus L.) (Muller 1952), some birches (Barnes
and others 1974), and aspens (Pauley 1956).
Testing for Breeding Value
After the elite, select, or superior individuals have been
selected, some system of testing their genetic value must be
used. We have identified these trees as good phenotypes but
we do not know their genotypes and therefore we are uncertain as to their value as breeding stock. Sometimes the outstanding trees in a stand may be taller than their neighbors
due to an environmental advantage such as better soil or
more moisture. It is important to use only trees with better
than average genetic characteristics, as the environmental
differences will not be passed on to future generations. In
natural stands, it is critical to determine the age of individual
trees. Trees growing together may have a similar size, yet be
quite different in age. Obviously we would prefer that our
select trees not be outstanding merely based on the fact that
they are older than their neighbors.
The usual way to test vegetatively propagated trees is to
plant them in blocks and compare performance with a standard population. This may be a clone of known performance, or in some cases seedlings from a standard seedlot
may be used. Tests that are designed to evaluate the relative
performance of a specific clone are called clonal tests.
Trees propagated from seed are usually progeny tested
with one or more test designs modified from crop breeding.
44
45
Table 3Chapter 2, Genetic Improvement of Forest Trees: examples of exotic species used in plantation forestry
Location of planting
Origin
North America
Picea sitchensis (Bong.) Carr.
Pinus elliotti var. elliotti Engelm.
Pinus radiata D. Don
Pinus taeda L.
Populus L. spp.
Pseudotsuga menziesii (Mirb.) Franco
Central America
Pinus caribea Morelet
Pinus oocarpa Schiede ex. Schltdl.
Europe
Picea abies L.
Populus L. spp.
Asia
Gmelina arborea Roxb.
Tectona grandis L. f.
Australia/New Zealand
Eucalyptus L.Her. spp.
46
North
America
Central
America
South
America
X
X
X
Europe
Africa
Australia &
New Zealand
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Table 4Chapter 2, Genetic Improvement of Forest Trees: loblolly pine (Pinus taeda L.) plantation* performance illustrates the
importance of geographic sources of seed
Source
Louisiana (Livingston Co.)
Texas (Montgomery Co.)
Georgia (Clarke Co.)
Arkansas (Howard Co.)
Survival (%)
82
83
77
84
Height
m
ft
dbh
cm
in
14
12
11.5
11
12
13
13
12
46
41
38
36
6.7
5.2
5.2
4.7
Volume
m2/ha cords/ac
265
145
113
94
42
23
18
15
47
48
Genetic Gains
Realized genetic gains in volume growth from first generation southern pine clonal seed orchards have ranged from
6% with un-rogued loblolly and slash pine orchards to 17%
for rogued orchards of these species (Squillace 1989). The
gains from advanced generation loblolly orchards have been
predicted at 25% greater volume than unimproved material
for the second generation, 35% for the third generation, and
45% for the fourth generation (Zobel and Talbert 1984).
Advanced-Generation Breeding
Advanced-generation breeding often is designed to combine the best individuals from the best families in the first
generation with unrelated individuals from a separate breeding population. The Western Gulf Forest Tree Improvement
Program has developed a sub-line system separating the
breeding population into breeding groups that are crossed to
produce seeds only when a production orchard is established
(Lowe and van Buijtenen 1986). With this system, inbreeding is restricted to the breeding populations and the production populations are not affected.
A similar system has been used with northern red oak
Quercus rubra L.in Indiana. Coggeshall and Beineke
(1986) have designed 6 sub-lines with 30 clones in each for
a total of 180 clones. These sub-lines will be crossed only
when the production seed orchard is established.
49
50
Molecular Biology
Isozymes
Brewbaker (1967) was one of the first scientists to propose the study and use of isozymes in forestry. Since then,
isozymes have been widely used for taxonomic work, pollen
contamination estimates, heterozygosity estimates, and a
number of other uses. In isozyme analysis, a single gene
codes for production of a single protein that can be visually
distinguished as one band on an electrophoretic gel. The
band pattern on a stained gel may be interpreted as a direct
reflection of the genotype of the tree. Cotyledons, needles,
or embryos (all diploid tissues) may be used or pollen
grains and female gametophytes (haploid tissues) may be
used.
Isozymes have been used to compare the rates of heterozygosity and outcrossing (El-Kassaby and others 1986)
with Douglas-fir. These authors found no significant differences between clonal and seedling seed orchards in outcrossing rates. There were, however, significantly greater
proportions of homozygous progeny from the seedling
orchard.
Although isozyme analysis has been an effective tool for
many forest genetics studies, Libby and others (1997), summarizing a southern meeting on genetic diversity, found that
isozyme data have a number of limitations when used to
estimate the genetic variation within a single species.
However, isozyme analysis has been widely used to estimate the amount of pollen contamination in seed orchards
(Adams and Birkes 1989).
51
52
California
The California Tree Improvement Association, organized
in 1978 with 26 members, manages over 9 million acres of
forestland. Ponderosa pine was the first species selected, followed by Douglas-fir and sugar pine. Members include forest industry, the State of California, and the USDA Forest
Service. Local tree improvement associations were formed
to focus on one or more of the California tree seed zones.
The main objectives of the association are the selection of
superior trees, the establishment of clone banks, the establishment of progeny test sites, and the establishment of a
ponderosa pine seed orchard.
The Great Lakes Region
The Minnesota Tree Improvement Cooperative was
established in 1980 and currently has 18 full members and
7 supporting members. The cooperative is working with
black (Picea mariana (Mill.) B.S.P) and white spruces, and
jack, red, and white pines. There are 35 seed orchards occupying about 50 ha (125 ac). In 1995, about 30 hl (84 bu) of
cones were collected from 3 of these orchards. Six orchards
were approved for production of certified seed in 1995.
Gains in height growth have ranged from 3 to 9%.
The Future
The demand for wood products will continue to increase
worldwide. Computer and printing paper will be in great
demand, particularly in Southeast Asia where population
growth is expanding exponentially (Kellison 1997).
In the United States, timber harvesting on the forest land
base is being progressively restricted, which dictates that
wood production be concentrated on less land area each
year. This requires management for maximum wood growth
on our most productive sites. Fortunately this will result in
reduced pressure on average and marginal sites which often
have high value for recreation and aesthetic pursuits.
Tree improvement programs, combined with intensive
management, have dramatically increased wood yields. In
the Southeastern United States, genetically improved loblolly pine on good sites, under maximum cultural care, can be
expected to yield 10.8 to 14.4 m3 (3 to 4 cords) per 0.4 ha
(1 ac) per year. On the Pacific Coast, vegetatively propagated hybrid cottonwood grown under maximum culture on 6to 7-year rotations is producing 25.2 m3 (7 cords) per 0.4 ha
(1 ac) per year (Kellison 1997). In South America, southeast
Asia, and South Africa, plantations of acacia, eucalyptus,
and Gmelina from genetically improved sources are expected to yield about 25.2 m3 (7 cords) per 0.4 ha (1 ac) per
year. Many of these plantations can be managed with coppice rotations to further increase their economic value.
Chapter 2: Genetic Improvement of Forest Trees
53
54
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Chapter 3
Contents
Introduction 58
Harvesting 58
Planning and Preparation 58
Quantities to collect 58
Positive species identification 58
Seed structures 60
Seed maturity 60
Seed Acquisition 63
Collection 63
Purchase 65
Lot identity 66
Transportation 66
Seed Cleaning and Conditioning 66
Post-Harvest Storage 66
Seed Extraction 68
Fleshy fruits 68
Dry fruits 69
Conifer seeds 71
Cleaning and Upgrading 73
Removing trash with air 73
Removing trash with screens 73
Removing sticks and needles 75
Removing trash with static electricity 76
Removing trash by rolling and sliding 76
Spiral separators also use rollability 77
Improving quality with specific gravity tables 77
Upgrading quality with liquids 78
Conveying seeds 80
Quality Control 81
References 83
57
Introduction
Most forest trees and shrubs grown for artificial regeneration purposes, and some grown horticulturally, are reproduced from seeds. Seed quality is, therefore, of critical
importance in determining the many options and outcomes
in producing a crop of seedlings. Only high-quality seeds
that can be planted by machinery permit bareroot seedlings
to be grown at the uniform and controlled bed densities
needed to produce the desired seedlings at the most economical cost. Uniform and controlled bed densities facilitate
more efficient and mechanized methods of weed control,
root-pruning, and lifting of seedlings. Adverse consequences
occur in both labor costs and the genetic makeup of the
seedlings unless high-quality seeds are used and properly
managed. Therefore, a poor job of seed handling at harvest
and conditioning can have serious negative impacts on the
quality and availability of seedlings. However, a good job at
this point will have positive impacts on seedling status. It
also makes sense economically to focus effort at this point
in the process, because more efficiencies can then be
realized at later stages.
This chapter begins with what to consider in planning a
seed harvest, how to harvest, how to temporarily store seeds,
and how to extract, clean, and upgrade the finished seeds. A
discussion of quality control concludes the chapter.
H arvesting
Seed harvest is the first step in producing a high-quality
seedlot. This statement assumes that genetic considerations
have been properly addressed in planning the seed harvest
(see chapter 2). Quantities of seeds to collect, initial seed
quality, and the timing of collections are the key quality factors in seed harvest. Timing is important because maximum
seed viability and vigor occur at physiological maturity (figure 1). Collecting too early results in lower seed quality due
to seed immaturity and reduced yield. Collecting too late
can also be detrimental, because seeds may be lost to seed
shed, predation by animals or insects, or seed deterioration.
However, some species mature after natural seedfall; these
include various ash species (Fraxinus spp.) and ginkgo
(Ginkgo biloba L.). The initial seed quality must be assessed
to avoid collecting seeds that are empty, malformed, or damaged. Quantity of seeds to collect influences quality because
the seeds must be processed before deterioration becomes
measurable.
58
Question
How many plants are to be produced?
How many years of production is this?
What is the ratio of seedlings to viable seeds?
What is the viability of the seeds?
How many seeds are there per unit weight*?
What is the purity?
What volume of the raw collection unit
(that is, seeds, fruits, or cones) must be
collected to obtain the desired weight* of
pure seeds?
Value
200,000
3
80%
80%
99,000
95%
0.8 kg/hl
Calculated quantity
of seeds needed
Example
200,000 x 3
600,000 0.8
750,000 0.8
937,500 99,000
9.5 0.95
10 kg x 0.8 kg/hl
600,000
750,000
937,500
9.5 kg
10 kg
12.5 hl
59
Seed count
6
8
10
14
Table 3Chapter 3, Seed Harvesting and Conditioning: sound seed yield per cone for 4 pine species as estimated
from the number of sound seeds exposed when cones are bisected longitudinally
Sound seeds
exposed
Pinus palustris
(Louisiana)
Pinus taeda
(Louisiana)
Pinus elliottii
(Louisiana)
2
4
6
8
10
12
14
23
35
47
59
71
83
95
31
44
57
70
83
96
109
20
35
50
65
80
95
110
Sources:
60
Pinus elliottii
(Georgia-Florida)
31
50
69
87
106
124
143
Pinus echinata
(Virginia)
12
22
31
41
51
60
70
drop. This delay provides some margin for error in estimating maturity. Prompt action is, however, necessary once
maturity has been reached.
Immature fruits may be collected by mistake from
species having fruits that require 2 or 3 years for development. Alaska-cedar (Chamaecyparis nootkatensis (D. Don)
Spach) and western juniper (Juniperus occidentalis Hook.)
are examples of species that bear immature seed structures
that are nearly equal to mature ones in color and size.
Collecting intermingled mature and immature fruits should
be avoided, because these fruits are difficult to separate
(Stoeckeler and Slabaugh 1965). At the other extreme is the
possibility of collecting empty conifer cones. Cones that
have recently shed their seeds will close during rainy weather (Allen and Owens 1972), and workers must be careful not
to collect these closed empty cones by mistake.
Maturity for collection is most often judged subjectively
from the appearance of the cones, fruits, or seeds. Green
color changes to yellow-green, yellow, brown, reddish, or
purple. Fruits begin to soften. Scales or bracts begin to crack
or flex. A few early-maturing individuals begin to drop their
seeds. Such subjective indicators have a variety of shortcomings (Schubert and Adams 1971). The color changes may
not be the same with every individual or population of
plants. Weather can accelerate or modify the appearance of
the indicator. On the whole, however, these indicators have
proven to be reasonably practical. Their chief drawback is
their dependence on the experience and judgment of collectors. When in doubt, it is generally better to shorten the collection period than to collect immature seeds that will have
low viability and the tendency to produce low-vigor,
deformed seedlings (Heit 1961, Schubert 1956).
Some attempts have been made to develop more-objective maturity indices. Chemical constituents have been analyzed and related to maturity for Douglas-fir and noble fir
(Abies procera Rehd.) (Rediske 1968). This type of
approach has not been widely used, mostly because of the
difficulty of getting samples to a laboratory and then returning the information on a timely basis to the field.
Measuring specific gravity is used to evaluate the maturity of conifer cones. As a cone matures, it loses water and
its specific gravity decreases. The flotation procedure to
measure specific gravity can be done in the field by collectors. Cones placed in a liquid with the appropriate specific
gravity will sink if immature and float if mature. The specific gravity values for ripe pine and fir cones are listed in
tables in part 2 of this manual. Although simple in application, this version of the flotation procedure requires finding
a fluid with the correct specific gravity and carrying supplies
61
62
Seed Acquisition
Seed harvesting can begin once a seedcrop of appropriate genetic make-up, adequate quality, adequate quantity,
and proper maturity has been identified. However, seed
quality must be monitored at the time of collection, for the
situation may have changed since first determinations were
made. Unexpected seed dispersal, insect-feeding, or other
animal predation may have occurred. Certain seeds, such as
acorns, can dry out or germinate shortly after they are shed
(Bonner and Vozzo 1987; McDonald 1969; Olson 1957).
Cones, fruits, and seeds should be cut open and examined
again. Sometimes a 10 x hand lens is useful in making a
quick evaluation. Prompt collection reduces losses to fungi,
insects, and larger animals and birds. Some caution is
advised in collecting the first fruits or seeds to be shed from
the plant, which may be of poor quality and dropping
because of death rather than maturation (Aldhous 1972;
Stoeckler and Jones 1957).
Collection. The actual gathering of the seed or fruit
from the plant takes many forms depending on the botanical
characteristics of the mother plant. The first characteristic
important to seed conditioning is the seed-bearing structure.
In gymnosperms (for example, pines, spruces, firs, and ginkgo), this is the female strobilus or cone; in angiosperms (for
example, all the hardwood trees), it is the fruit. If this structure is not easily removed from the plant or shed naturally, it
is classified as persistent. Persistent cones and fruits require
more effort to remove them from the plant. Either hard
twisting and pulling, or a sharp cutting tool such as pruning
shears is needed to sever the connection (figure 5).
Alternatively, the seeds are allowed to shed naturally and
caught on netting (figure 6). When not persistent, the cones
or fruits can be quickly picked by hand or pulled off with
rakes, hooks, or vacuum. Another method is to gather cones
or fruits from the ground after shaking them from the plant
or after natural drop (figures 7 and 8). Yard tools such as
rakes, leaf blowers/vacuums, forks, and shovels can be useful in gathering seeds from the ground or off of low plants
(figure 8).
In some loblolly, shortleaf (Pinus echinata P. Mill.), and
eastern white pine seed orchards an extensive system of nets
is used to collect seeds. Loblolly cones are very persistent,
and much labor and equipment expense is saved by gathering seeds that shed naturally rather than using lift trucks or
other devices to pick cones. Problems related to seed maturity are almost totally avoided by relying on natural shedding
of the seeds. Avoiding such problems is the strongest attrac-
tion of the system in white pine orchards, because the window of time to collect is quite narrow for white pine. The
system has several disadvantages, however. There must be
enough dry weather during the collection period for natural
seed shed to finish by the desired collection date. Predation
from insects and larger animals can occasionally be too
great, and weed seeds can enter the seedlot from ground
plants and vines in the orchard and from droppings of birds
that roost in the orchard. Also, family identities are lost in
the bulk collection of seeds. This system can be used only
for seeds with at least moderate levels of dormancy; nondormant seeds germinate on the netting, resulting in great
loss of quality and quantity.
The netting is carpet backing with UV light inhibitors. It
is used in widths equal to the distance between rows of trees
in the orchard. The individual strips are then drawn together
at the edges and fastened with standard wire staples commonly used for paper (figure 9). If the ground is soft, tree
shakers are used to shake the seeds from the open cones of
loblolly pine, which has a hard seedcoat. As the trees are
shaken twice, the shaker actually drives over some seeds
that fell on the netting during the first shake. For this reason,
shakers cannot be used for eastern white pine, for their seedcoats are too fragile and can be mechanically damaged when
the seeds are driven over. Totally natural seed shed must be
used in this case. Alternatively, overflights with helicopters
can be used, although these are usually too expensive.
63
Once the seeds are shed from the trees, the netting is
rolled up. This is done by a net retrieval machine or simply
by drawing the net over itself and piling the seeds in a windrow. The windrow is then combined with a peanut combine
to separate the seeds from the bulk of the needles
64
65
66
Those seeds that can be dried are referred to as orthodox. Orthodox seeds are broken into 2 groups:
fleshy fruitsfor example, those of dogwoods (Cornus
spp.), cherries (Prunus spp., and junipers (Juniperus
spp.)
non-fleshy fruitsfor example, those of ash, elm
(Ulmus spp. L.), and pine
Fleshy fruits must be prevented from drying until the
pulp is removed, otherwise the pulp hardens and is then not
easily or adequately removed. Pulp is generally removed to
make handling easier and to control fungi or bacteria that
can grow in the pulp. As with recalcitrant species, the
fleshy-fruited orthodox seeds must be kept cool and ventilated to prevent the buildup of heat and subsequent deterioration. They may be treated basically the same as the recalcitrants; however, fleshy-fruited orthodox seeds might require
more ventilation because of higher moisture content.
The dry-fruited (also referred to as non-fleshy) orthodox
species should be allowed to dry to prevent deterioration.
High moisture in these fruits usually leads to heating, molding, and subsequently, loss of viability. Spreading the seeds
on screen racks is an economical way to dry these fruits.
Minimal drying is necessary during post-harvest storage
when the seeds are collected dry. Post-harvest storage conditions usually need to allow for the loss of moisture at a gradual rate and to protect the seeds from the weather. If the fruit
expands upon drying (for example, a pine cone), sufficient
space must be allowed for expansion. Otherwise the fruit
will become case-hardened and the seed locked inside the
fruit as discussed previously. Figure 12 shows wire-bottom
racks for air-drying cones under shelter. Alternatively, moisture must be maintained at a high level to prevent the expansion of cones. Because high moisture can lead to seed deterioration, it is important to evaluate a system that keeps
orthodox seeds moist to be sure there is no loss of seed
quality. Cones in full sacks or in bulk must be re-bagged
into half-full sacks or placed in ventilated storage. Drying
racks with cones spread out about two cones deep are very
good. Cones have also been successfully stored in temporary
cribs of snow fencing 8 to 10 feet in diameter. Storing cones
in this type of crib or in 35-liter (20-bushel) wire-bound
boxes ventilates the cones but also keeps them from drying
and loosing their seed. Both cribs and boxes are used out of
doors. Cones should never be stored in a large pile as this
will result in heating. Smaller piles will cause many surface
cones to open and the seeds will be lost. Also, piling cones
on the ground invariably leads to stones in the seedlot.
67
68
69
Table 4Chapter 3, Seed Harvesting and Conditioning: maximum recommended ambient relative humidity values for
drying seeds, cones, and fruits with heated air at various drier temperatures from 24 to 43 C
Ambient
air temp
(C)
4
7
8
9
11
16
18
20
22
24
27
70
27 C
29 C
32 C
35 C
38 C
41 C
100
100
100
94
100
77
68
60
50
45
38
35
100
100
100
100
88
78
65
58
49
45
38
100
100
100
100
100
94
84
71
65
55
47
100
100
100
100
100
100
95
81
69
63
54
100
100
100
100
100
100
100
95
82
74
64
75
86
65
58
51
42
38
31
30
43 C
100
100
100
100
100
100
100
100
100
87
75
tightly they press against the shell, and how long the seeds
are retained in the machine. The fruits need to be dry for the
hulling operation to work well. The effect of hulling with a
brush machine can be seen in figure 20. A partial list of genera and species that can be successfully de-winged or hulled
with the brush machine includes ash, maple, tuliptree
(Liriodendron tulipifera L.), southern catalpa (Catalpa
bignonioides Walt.), black locust, sycamore, big sagebrush
(Artemisia tridentata Nutt.), mountain-mahogany
(Cercocarpus montanus Raf.), and winterfat
(Krascheninnikovia lanata (Pursh) A.D.J. Meeuse & Smit)
Conifer seeds. Conifer seeds are usually de-winged
after tumbling from the cone. De-winging can be done either
wet or dry. Some species, white pines, for example, separate
easily when tumbled in a drum such as a concrete mixer.
Othersfor example, hard pines and sprucesrequire
adding a small amount of water to the seeds as they are tumbled to release the wings. In other machines, pressure can
be applied with brushes or paddles to remove wings from
dry seeds. A mortar mixer is an example of a machine commonly adapted to the dry de-winging of conifer seeds with a
modest amount of pressure. However, using pressure
increases the chance of mechanical damage. To minimize
the amount of mechanical injury, the paddles should be
slowed by changing gears on the mixer and de-winging the
seeds for only a limited time. A timer switch can be used to
easily control de-winging time. The Missoula small-lot pine
71
72
73
74
75
rolled against the sides of the beaker. After the trash has
clung to the sides of the beaker, the clean seeds can be
poured out. Plastic cups can substitute for the glass beaker.
Removing trash by rolling and sliding. Another
characteristic that can be used to separate seeds and trash is
surface texture, or the ability to slide or roll down a slope.
Conifer seeds, especially larch or white pines, can contain a
76
large amount of pitch. The pitch particles have a sticky surface whereas the seeds are relatively smooth. The vibratory
separator (figure 35) is often able to use this difference to
remove the pitch. This separator consists of a 22.9-cmsquare (9-inch-square) deck mounted on a variable-speed
vibrator. The deck has adjustable side and end tilt. The
vibration causes the pitch, which grips the deck surface
because it is tacky, to walk up the slope while the seeds,
which are smoother, slide down the slope. The rate at which
seeds are fed onto the deck, the speed of deck vibration, the
roughness of the deck, the degree of side and end tilts, and
the arrangement of the cut gates are all important. Trial and
error determine what adjustments are necessary to get a separation.
Not all pitch will separate on the vibratory separator.
Some may be too dry and, therefore, too smooth to stick to
the deck well enough. Other pieces of pitch may be too
round. Screening will sometimes work in these cases.
Another approach to surface texture is to modify it.
Sometimes pitch can be removed with a gravity table (see
section below on gravity tables), but only after it has been
very well dried or stiffened by placing the seedlot in a cooler or freezer (Zensen 1980). This drying or cooling keeps
the seeds from balling up around the pitch particles.
Another type of machine that separates by particles
ability to roll is the inclined draper (figure 36). This
machine is a variable speed belt that can be set at different
slopes. Particles that are round, or able to slide more easily,
go down the slope while the flatter particles with greater
friction ride up the hill on the moving belt and are placed in
the upper collection box. Separating juniper berries from
juniper leaves is an example of a separation that can be done
with the draper. A board with a piece of cloth over it makes
a simple draper for a small quantity of seeds. A handful of
seeds can be cleaned at one time with this board. The seeds
are allowed to roll down the board and then the needles are
manually dumped off.
Spiral separators also use rollability. These are made
of two concentric metal spirals. Seeds are poured into the
top of the inner spiral. As they slide down the spiral, the
round particles roll and gain momentum, causing them to fly
out of the inner spiral into the outer spiral. Trash can be
removed from dogwood and juniper berries in this manner
(Delany 1998).
Improving quality with specific gravity tables.
Specific gravity tables are another class of machine that separates by weight (figure 37). Gravity tables can be used on
almost any species of seed that flows freely. They have been
very successful in cleaning true firs, longleaf pine, tuliptree,
77
the lighter material, not toss it into the air. This more gentle
air flow results in a finer stratification of particles by weight
than is possible in the air columns of blowers and aspirators.
To pull the light and heavy particles apart, the deck shakes
back and forth and is tilted sideways to oppose the direction
of the shake. The shake of the deck pushes the heavy particles up the hill while the light particles drift down the slope,
floating on top of the heavier seeds and pulled down the
slope by gravity. This stratification of particles by weight
and separation by the use of the shake and slope continues
as the seed mass works its way across the deck, giving a
continuous gradation of particle weights until the heaviest
are at the top, intermediate weights are in the middle, and
the lightest are at the bottom. When seedlots are upgraded
on gravity table, the lightest particles can be empty seeds,
cone particles, straw, partially filled seeds, or even goodquality seeds that are lower in weight. The heavier particles
are usually the heaviest good-quality seeds but might also be
pitch, stones, dirt balls, or tramp metal that has fallen into
the seeds.
Dimensional grading of seedlots before using the gravity
table improves the effectiveness of the table and may even
be essential. Grading should be done both by width, thickness, and, if possible, length. The more dimensional grading
is done, the better the table will work. The table sorts by
density or dimension but never effectively for both at the
same time. For example, partially filled large-diameter seeds
and same-weight seeds that are completely filled but smaller
78
damaged seedcoats. The seeds are placed in water in a vacuum chamber and a vacuum is then drawn on the chamber to
break the surface tension and allow the water to wet the surface of the seeds. Water will enter seeds with damaged seedcoats rapidly, increasing their weight and causing them to
sink. These sinkers are discarded, for they have damaged
seedcoats and will be dead. The seeds that float are drawn
off the top and kept. Conducting the PREVAC procedure
before IDS prevents the mechanically damaged seeds from
sorting out with the good seeds in the IDS procedure.
The IDS procedure uses the fact that dead or weak seeds
lose water faster than living or more vigorous seeds. In the
first step, all seeds are allowed to imbibe water. With Scots
pine, this is done at 15 C for 8 to 12 days. Then the seeds
are dried in super-dry air with a relative humidity ranging
from 5 to 15%. The extremely rapid drying resulting from
the super-dry air maximizes the difference in drying rates
between viable and nonviable seeds. The living tissue in the
viable seeds holds water more tightly than the nonliving tissue in the nonviable seeds. Eventually, however, the viable
and nonviable seeds will both dry to the same moisture content. Meanwhile, though, samples of seeds are drawn at set
intervals through the drying period and placed in water to
determine the best length of time to dry. When the number
of seeds floating equals the number to remove, all the seeds
are placed into the water. The floaters are nonviable and are
discarded; the sinkers are viable and are kept (figure 39).
Unless sown immediately, the sinking seeds must be dried
thoroughly to a proper storage moisture content. The process
has been completely automated in Sweden using sophisticated machines. However, the basic principle can be followed
using pans of water and a kitchen sieve. The key is proper
incubation and very rapid rate of drying. In extremely cold
climates, it is possible to simply heat ambient air to near 40
C to achieve the desired relative humidity. However, it may
be necessary to dehumidify the air further by cooling it with
an air-conditioner before heating. The IDS process is protected by patent in Canada and the United States, and royalties must be paid to use the process commercially (Downie
and Bergsten 1991). Successful upgrades have been reported
for seedlots of Douglas-fir (Sweeney and others 1991) and
Pinus roxburghii Sarg. (Singh and Vozzo 1990). Attempts at
upgrading seedlots of white spruce, sitka spruce (Picea
sitkensis (Bong.) Carr.), and ponderosa pine (Pinus ponderosa P. & C. Lawson) gave mixed results (Downie and
Bergsten 1991; Downie and Wang 1992; Karrfalt 1996).
79
Conveying seeds. Moving seeds in forest seed processing plants is usually best accomplished by batch movement. In most cases, this means that a 4.5-liter (5-gallon)
pail must be lifted and emptied into a hopper that feeds a
conditioning machine about every 15 to 20 minutes. Larger
amounts are sometimes moved with a hopper and forklift.
The advantages of batch movement are greater control and
flexibility of seed flow, easier clean-out, and a simpler, less
expensive design. Greater control is provided because the
seeds can be seen more easily for continual inspection. If the
desired result is not obtained on one pass, the seeds can be
immediately rerun. In continuous flow systems the seeds
must flow to the next machine. Batch processing also provides greater flexibility, because the order of conditioning
can be altered to match the lot and kind of seeds. In continuous flow the order of processing is relatively fixed. Bucket
elevators have been used frequently in the past to move
seeds, but these can be difficult to clean out. With many elevator designs, as much as 0.45 to 2.3 kg (1 to 5 lb) of seeds
remain in the bottom or boot of the elevator and must be
cleaned out by hand. This is much work and highly impractical with lots of less than 23.7 kg (50 lb). Because of this
Table 5Chapter 3, Seed Harvesting and Conditioning: the application and interpretation of various seed
tests to seed conditioning
Test
Observation
Orthodox seeds
Above 10% MC
Below 10% MC
Recalcitrant seeds
Above 25%
Below 25%
Low (trash present will clog a seeder)
95% or better
Purity
Germination, tetrazolium
staining, or Excised
embryo
Seed weight*
X-radiography
Standard
Contrast agent
80
Interpretation/action
Quality Control
Quality control is very important in conditioning seeds.
Seed conditioning can be a highly complex operation, usually involving many steps and many different persons. A procedures manual is crucial, so that the role of everyone
involved can be clearly identified and thought out. Writing
out the steps in detail, from preparing for collection through
storage and planting, provides an opportunity for a careful
examination of all steps to identify potential problems and
inefficiencies. Also, when procedures are fully documented,
the same steps can be followed if there is a change in personnel. Many good techniques and small nuances gained
through years of seed conditioning can be lost if not documented in permanent form. Typically, plant managers are
not expected to publish their techniques, and knowledge is
lost at retirement or job transfer. It is therefore imperative
that the manager document all procedures used, in detail,
either on the managers own initiative or with the managers
supervisor. Sometimes an interview with someone who has
good writing ability will help a manager get thoughts on
paper.
The procedure manual should indicate all the types of
records to be kept and what to do when an error occurs.
Sooner or later a mistake inevitably will occur, and proce-
81
Visual inspection, cutting tests, and, ideally, x-radiography is used to monitor the conditioning operation. Visual
inspection leads us in improving purity. Cutting tests and xradiography reveal how many bad seeds still need to be
removed. X-radiography is the better method, for it can easily show mechanical damage to the seeds, is very fast, and is
more accurate than cutting tests in differentiating between
good and bad seeds. If an x-ray shows that 20% of the seeds
are bad and must be removed, the cleaning equipment
should then be adjusted to remove 20% of the seeds by
number or volume (figure 42). A second check can then be
made of the seeds after the machine has been adjusted to
ensure the adjustment has removed all the bad seeds it
should without removing too many good seeds. The separation may not be able to be completed with one setting.
In addition to daily monitoring, the conditioning procedures need to be verified as correct by full laboratory tests,
to be sure no harm is coming to the seeds and that the
desired quality is achieved. Purity tests tell how much of the
seed by weight is pure seed and how much is trash. A low
82
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current research in the chemical sciences:Proceedings, 3rd Annual
Southern Station Chemical Sciences Meeting;1990 February 78;
Alexandria, LA. Gen.Tech. Rep. SO-101. New Orleans:USDA Forest
Service, Southern Forest Experiment Station:710.
Stoeckler JH, Jones GW. 1957. Forest Nursery practice in the Lake States.
Agric. Handbk. 110.Washington, DC:USDA Forest Service. 124 p.
Stoeckler JH, Slabaugh PE. 1965. Conifer nursery practice in the Prairie
Plains.Agric. Handbk. 279.Washington, DC:USDA Forest Service. 93 p.
Sweeney JD, El-KassabyYA,Taylor DW, Edwards DGW, Miller GE. 1991.
Applying the IDS method to remove seeds infected with seed chalcid,
Megastigmus spermotrophusWachtl, in Douglas-fir, Pseduotsuga menziesii
(Mirb.) Franco. New Forests 5:327334.
Taylor AG, McCarthy AM, Chirco EM. 1982. Density separation of seeds
with hexane and chloroform. Journal of Seed Technology 7(1):7883.
Tylkowski T. 1984. The effect of storing silver maple (Acer saccharinum L.)
samaras on the germinative capacity of seeds and seedling growth.
Arboretum Kornikie Rocznik 29:131141.
Waldrip BT Jr. 1970. Artificial ripening of loblolly pine cones and seed.
Southeast. Nurserymens Conference Proceedings;1970.Atlanta:USDA
Forest Service, State and Private Forestry:8291.
Wilcox JR. 1966. Sweetgum seed quality and seedling height as related to
colleciton date. In:Proceedings, 8th Southern Conference on Forest
Tree Improvement;1965 June 16B17;Savannah, GA. Spons. Pub. 24.
Macon:[Georgia Forest Research Council, Committee on Southern
Forest Tree Improvement. 161 p.
Winston DA, Haddon BD. 1981. Effects of early cone collection and artificial ripening on white spruce and red pine germination. Canadian
Journal of Forest Research 11(4):817826.
Young JA, Budy JD, Evans R. 1983. Germination of seeds of wildland plants.
In:Proceedings, Intermountain Nurserymans Association Conference;
1983 August 811;LasVegas, NV. 93 p.
Zensen F. 1980. Improved processing techniques for western larch.Tree
Planters, Notes 31(4):2325.
83
84
Chapter 4
Storage of Seeds
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Contents
Introduction 86
Factors Affecting Longevity of Seeds
Seed Characteristics 86
Basic seed physiology 86
Seed morphology 88
Chemical composition 88
Seed maturity 89
Seed Handling Prior to Storage 89
Storage Environment 89
Moisture 89
Temperature 90
Atmosphere 91
86
Storage Facilities 92
Cold storage 92
Containers 92
Moisture Control 93
Storage Recommendation 93
Orthodox Seeds 93
Temperate-Recalcitrant Seeds 94
Tropical-Recalcitrant Seeds 94
Cryogenic Storage 94
Other Management Considerations
References 95
94
85
Introduction
In the simplest form of seed storage, mature seeds are
held for a short period until weather or other factors permit
sowing or planting. In the more comprehensive view, there
are at least 3 objectives for storing seeds, and each of them
dictates different strategies and procedures. These objectives
may be described as storage for the following periods:
1. Very short periods (overwinter) between collection and
sowing
2. Several years (10 or less) to ensure a reliable supply of
seeds in the absence of annual crops
3. Long periods (10 to 50+ years) for germplasm
conservation
The strategies employed will depend on all of the factors that influence seed longevity. Some of these factors
have been discussed in chapter 1, but now they will be
explored in the context of seed storage only. Following this,
recommendations will be made for storage procedures to
achieve the objectives listed here.
86
Table 1Chapter 4, Storage of Seeds: storage test results for some true orthodox species
Test conditions
Species
Sources:
Seed moisture
(%)
Temp (C)
Test results
0
48
10
24
15
[??]
3
3
5
5
35
6
5
24
3
48
24
24
4-5
0
04
5, 18
3
10
58
1623
616
8
8
48
6
7.5
510
79.5
11
<8
8
ca.12
8
10
Period
(years)
Viability
loss (%)
7
1.2
5.5
4
8
4
8
2
6
6
520
2
2.5
14
9
1.5
1324
1718
4
7
7
2
5
11
6
5
013
Little
Little
2
05
7
1
<5
<10
27
3
10
011
08
None
None
None
None
None
Bonner (1990), Clausen (1967), Jones (1987), Springfield (1968, 1973, 1974),Tylkowski (1987),Wang and others (1993).
Table 2Chapter 4, Storage of Seeds: storage test results for some sub-orthodox species
Test conditions
Species
Citrus limon (L.) Burm. F.
Fagus silvatica L.
Gmelina arborea Roxb.
Populus deltoides Bartr. ex Marsh.
P. grandidentata Michx.
P. tremuloides Michx.
Salix glauca L.
Test results
Temp (C)
20
10
5
20
18
18
10
5
10
610
610
1115
68
610
Period (years)
0.9
5
2
6
12
2
1.2
Sources: Bonner (1990), Fechner and others (1981),Wang and others (1982).
87
Table 3Chapter 4, Storage of Seeds: storage test results for some temperate recalcitrant species
Test conditions
Species
Acer saccharinum L.
Quercus macrocarpa Michx.
Q. pagoda Raf.
Q. robur L.
Q. rubra L.
Q. virginiana P. Mill.
Temp (C)
Test results
Seed moisture
3
1
3
1
1 to 3
2
Period (months)
50
44
35
4045
3845
18
6
30
29
17
12
8
None
6
3161
1846
35
Table 4Chapter 4, Storage of Seeds: storage test results for tropical recalcitrant species
Test conditions
Species
Araucaria hunsteinii K. Schum. & Hullrung
Azadirachta indica Adr. Juss
Dipterocarpus turbinatus C. F. Gaertn.
Hopea helferi (Dyer) Brandis
Shorea robusta C. F. Gaertn.
S. roxburghii G. Don
S. talura Roxb.
Symphonia globulifera L. f.
Temp (C)
19
26
16
15
13.5
16
23.5
15
2530
1018
4144
47
4050
40
47
Test results
Period (days)
54
56
161
37
30
270
105
270
Sources: Bonner (1990), Bras and Maury-Lechon (1986), Purohit and others (1982),Tompsett (1987).
88
drates and very little lipid. Even among the black oaks,
species with the highest lipid contents seem to store better,
even though there is no evidence of cause and effect. One
must conclude that among a wide range of species there is
no compelling argument for gross chemical composition as
the critical factor in seed longevity under proper storage
conditions. There is some evidence, however, that suggests
that the relative concentrations of particular carbohydrates
play key roles in desiccation tolerance, a critical property in
determining storage behavior of seeds (Lin and Huang
1994). This topic is obviously one that deserves more
research.
Seed maturity. Seeds of many orthodox species that
are immature when collected (or extracted from fruits) are
likely to fare poorly in storage (Stein and others 1974).
Experimental evidence has demonstrated this fact for Scots
(Pinus silvestris L.) (Kardell 1973), loblolly (P. taeda L.),
longleaf (P. palustris P. Mill.), and eastern white (P. strobus
L.) pines (Bonner 1991). The physiological basis for this
effect is not known, but it seems logical that immature seeds
have not been able to complete the normal accumulation of
storage food reserves, develop all needed enzymes and/or
growth regulators, or complete their full morphological
development and cell organization. For species with seeds
that are naturally dispersed while still physiologically immature, such as Fraxinus excelsior L., there is no apparent
damage to storage longevity (Willan 1985). The ability to
complete maturation naturally after separation from the
mother tree has apparently evolved with these species. For
conifers like the pines noted above, storage of immature
cones for several weeks prior to extraction of the seeds
appears to enhance seed maturity and viability retention
during storage (Bonner 1991).
Seed Handling Prior to Storage
Poor fruit or seed handling that damages seeds will often
lead to reduced viability in storage, especially in orthodox
seeds. The most common example of this is impact damage
to seeds during extraction and conditioning. Seeds can be
bruised by excessive tumbling of cones, running dry
dewingers too fast or too full, or poor transport systems
(Kamra 1967). During kiln drying of conifers, excessive
heat while seed moisture is still high can easily lead to damage that will show up later as reduced vigor and viability in
stored seeds (see chapter 3).
Another factor to consider in damage to seeds during
extraction and conditioning is cracks or other breaches of
the seedcoats that will allow microorganisms to enter.
Cracks in seedcoats that occur during seed conditioning are
89
potential
Potential effect
Germination can occur
Active fungal growth
Insect activity reduced
Best range for sealed storage
Desiccation injury possible in
some species
Table 6Chapter 4, Storage of Seeds: equilibrium moisture content at 4 to 5 C and 3 relative humidities
for some seeds
Species
Trees with orthodox seeds
Carya ovata (P. Mill.) K. Koch
Juglans nigra L.
Liquidambar styraciflua L.
Liriodendron tulipifera L.
Picea abies (L.) Karst.
Pinus sylvestris L.
P. taeda L.
Prunus serotina Ehrh.
Crops with orthodox seeds
Glycine max (L.) Merr.
Zea mays L.
Trees with recalcitrant seeds
Quercus alba L.
Q. nigra L.
Shorea robusta Gaertner f.
Sources: Bonner (1981), Bass (1978).
90
20% RH
45% RH
95% RH
6
6
10
11
8
10
8
8
10
9
15
20
20
19
17
17
6
8
8
12
19
20
37
17
50
29
35
Table 7Chapter 4, Storage of Seeds: cryogenic storage test results for some forest tree seeds
Species
<13
<13
Period (days)
6
6
6
6
6
6
112
112
180
180
91
Storage Facilities
Cold Storage
Facilities for seed storage will vary by the amount of seeds
to be stored and the projected length of storage. Small seedlotsa liter (quart) or lesscan be stored in household
refrigerators and freezers. Larger seedlots and quantities will
require a walk-in refrigerator or freezer (figure 3). These
units are usually assembled from prefabricated insulated
panels and can be made almost any size to fit the owners
needs. A suggested size for a nursery operation is one large
enough to hold a 5-years supply of seeds. The cold storage
at the USDA Forest Services W. W. Ashe Nursery in
Brooklyn, Mississippi (figure 3) has a capacity of 1,584 m3
(52,800 ft3). One cubic meter will hold from 125 to 140 kg
(275 to 310 lb) of seeds. Many orthodox and sub-orthodox
seeds show declining germination and vigor after a few
years in storage at temperatures just above freezing (Bonner
1991; Zasada and Densmore 1977), so freezers maintained
at about 18 to 20 C are preferred for any storage of sensitive species longer than 3 or 4 months. Because it would
be inconvenient to have separate facilities, most users just
place all orthodox species in freezers. For reasons discussed
92
a large walk-in
Storage Recommendations
Orthodox Seeds
All orthodox seeds should be stored in moisture-proof,
sealed containers with seed moisture contents of 5 to 10%.
93
If the period of storage will be 3 years or less for true orthodox species, or 2 years or less for sub-orthodox species,
temperatures of 0 to 5 C are sufficient. For longer periods
of storage for both types of orthodox species, freezers (18
to 20 C) should be used.
Temperate-Recalcitrant Seeds
Temperate recalcitrant seeds should be stored with moisture
contents at least as high as that present when the mature
seeds were shed from the tree. (Refer to genus chapters in
this manual for information on individual species.) This
moisture level must be maintained throughout storage,
which may require occasional rewetting of the seeds.
Temperatures should range from 0 to 5 C, although 1 or 2
degrees below freezing will not harm most species.
Containers should be basically impermeable to moisture
loss, but must allow some gas exchange with the atmosphere. Polyethylene bags with a wall thickness of 0.075 to
1.0 mm (3 to 7 mils) are suitable. Some oak acorns can be
stored for 3 years in this fashion (table 3), but some viability
will be lost. For other recalcitrant species, few data are
available.
Tropical-Recalcitrant Seeds
Storage of tropical recalcitrant seeds is done in the same
manner as storage of temperate species, except that temperatures must be kept at a high level. There are differences
among species but the lower limits are generally 12 to 20
C. Successful storage for more than 1 year should not be
expected.
Cryogenic Storage
For long-term germplasm conservation programs, true
orthodox and sub-orthodox seeds can be dried to moisture
contents of 5 to 10% and stored in liquid nitrogen. Such programs require special equipment and procedures and are
beyond the scope of this book.
94
References
4
Barnett JP,Vozzo JA. 1985. Viability and vigor of slash and shortleaf pine
seeds after 50 years of storage. Forest Science 31: 316320.
Berjak P, Pammenter NW. 1996. Recalcitrant (desiccation-sensitive) seeds.
In: Olesen K, ed. Innovations in tropical tree seed technology.
Proceedings, IUFRO Symposium of the Project Group P.2.04.00, Seed
Problems; 1995 September 710; Arusha,Tanzania. Copenhagen,
Denmark: National Tree Seed Programme: 1429.
Bonner FT. 1971. Chemical contents of southern hardwood fruits and
seeds. Res. Note SO-136. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 3 p.
Bonner FT. 1973. Storing red oak acorns.Tree Planters Notes 24(3): 1213.
Bonner FT. 1981. Measurement and management of tree seed moisture.
Gen.Tech. Rep. SO-49. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 11 p.
Bonner FT. 1990. Storage of seeds: potential and limitations for germplasm
conservation. Forest Ecology and Management 35: 3543.
Bonner FT. 1991. Effect of cone storage on pine seed storage potential.
Southern Journal of Applied Forestry 15: 216221.
Bonner FT. 1992. Unpublished data. USDA, Forest Service, Mississippi State,
Mississippi.
Bonner FT. 1994. Predicting seed longevity for four forest tree species with
orthodox seeds. Seed Science and Technology 22: 361370.
Bonner FT,Vozzo JA, Elam WW, Land SB Jr. 1994. Tree seed technology
training course: instructors manual. Gen.Tech. Rep. SO-106. New
Orleans: USDA Forest Service, Southern Forest Experiment Station.
160 p.
Bonnet-Masimbert M, Muller C. 1975. La conservation des faines est possible. Revue Forestiere Francaise 27: 129138.
Bras P, Maury-Lechon G. 1986. Graines forestieres tropicales de type fortement hydrate: la conservation et ses effets, exemple du Symphonis globulifera L.f. de Guyane Francaise. Bois et Forets des Tropiques 212: 3546.
Chin HF, Roberts EH. 1980. Recalcitrant crop seeds. Kuala Lumpur,
Malaysia:Tropical Press Sdn. Bhd. 152 p.
Clausen KE. 1975. Long-term storage of yellow and paper birch seed. Res.
Note NC-183. St. Paul: USDA Forest Service, North Central Forest
Experiment Station. 3 p.
Danielson HR,Tanaka Y. 1978. Drying and storing stratified ponderosa pine
and Douglas-fir seeds. Forest Science 24: 1116.
Eliason EJ, Heit CE. 1973. Red pine seed shows high germination after 42
years in storage. Journal of Forestry 71: 776.
Ellis RH, Hong TD, Roberts EH. 1990. An intermediate category of seed
storage behavior? 1. Coffee. Journal of Experimental Botany 41:
11671174.
Fechner GH, Burr KE, Myers JF. 1981. Effects of storage, temperature, and
moisture stress on seed germination and early seedling development of
trembling aspen. Canadian Journal of Forestry Research 11: 718722.
Hampton JG,TeKrony DM, eds. 1995. Handbook of vigour test methods.
3rd ed. Zurich: ISTA. 117 p.
Harrington JF. 1973. Problems of seed storage. In: Heydecker W, ed. Seed
ecology. University Park: Pennsylvania State University Press: 251263.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing: rules 1993. Seed Science & Technology 21, Supplement:
1259.
Jones L. 1967. Effect of storage at various moisture contents and temperatures on seed germination of silk oak, Australian pine, and Eucalyptus spp.
Res. Note SE-83. Asheville, NC: USDA Forest Service Southeastern
Forest Experiment Station 4 p.
Justice OL, Bass LN. 1978. Principles and practices of seed storage. Agric.
Handbk. 506. Washington, DC: USDA Science and Education
Administration. 289 p.
Kamra SK. 1967. Studies on storage of mechanically damaged seed of Scots
pine (Pinus silvestris L.). Studia Forestalia Suecica 42:119.
Kardell L. 1973. [in Swedish; with English summary: Studies on pine seeds
from northern Sweden: 2. Investigations on storage of pine cones and
pine seeds (Pinus silvestris L.) in northern Sweden]. Lund, Sweden:
Allmnna Frlaget. 70 p.
Lauridsen EB, Stubsgaard F. 1987. Longevity of hardcoated seed after scarification.Tech. Note 32. Humlebaek, Denmark: Danida Forest Seed
Centre. 4 p.
Lauridsen EB, Olesen K, Scholer E. 1992. Packaging materials for tropical
tree fruits and seeds.Tech. Note 41. Humlebaek, Denmark: Danida
Forest Seed Centre. 25 p.
Lin TP, Huang NH. 1994. The relationship between carbohydrate composition of some tree seeds and their longevity. Journal of Experimental
Botany 45: 12891294.
Martin SC. 1948. Mesquite seeds remain viable after 44 years. Ecology 29:
393.
Purohit AN, Sharma MM,Thapliyal RC. 1982. Effect of storage temperatures on the viability of sal (Shorea robusta) and talura (Shorea talura)
seed. Forest Science 28: 526530.
Roberts EH. 1973. Predicting the storage life of seeds. Seed Science and
Technology 1: 499-514.
Schroeder WR, Walker DS. 1987. Effects of moisture content and storage
temperatures on germination of Quercus macrocarpa acorns. Journal of
Environmental Horticulture 5(1): 2224.
Shrestha KB, Shepherd KR,Turnbull JW. 1985. Controlled atmosphere storage for Pinus radiata seed. Commonwealth Forestry Revue 64(2):
141150.
Simak M. 1966. [in Swedish, English summary: Chromosome changes in
ageing seed]. Skogen 53: 2830 [Forestry Abstracts 27: 3766; 1966].
Springfield HW. 1968. Cold storage not required for fourwing saltbush
seeds. Journal of Range Management 21: 335336.
Springfield HW. 1973. Cliffrose and mountainmahogany seeds retain viability 6 years in cold storage. Res. Note RM-236. Fort Collins, CO: USDA
Forest Service Rocky Mountain Forest and Range Experiment Station.
2 p.
Springfield HW. 1974. Winterfat seeds viable after 8 years refrigerated storage. Journal of Range Management 27(1): 78.
Stein WI, Slabaugh PE, Plummer AP. 1974. Harvesting, processing, and storage of fruits and seeds. In: Schopmeyer CS, tech. coord. Seeds of woody
plants in the United States. Agric. Handbk. 450. Washington, DC: USDA
Forest Service: 98125.
Stubsgaard F. 1992. Seed storage. Lecture Note C-9. Humlebaek, Denmark:
Danida Forest Seed Centre. 36 p.
Suszka B. 1975. Cold storage of already after-ripened beech (Fagus silvatica
L.) seeds. Arboretum Kornickie 20: 299315.
Tompsett PB. 1982. The effect of desiccation on the longevity of seeds of
Araucaria hunsteinii and A. cunninghamii. Annals of Botany 50: 693704.
Tompsett PB. 1987. Desiccation and storage studies on Dipterocarpus
seeds. Annals of Applied Biology 110: 371379.
Tylkowski T. 1984. The effect of storing silver maple (Acer saccharinum L.)
samaras on the germinative capacity of seeds and seedling growth.
Arboretum Kornickie 24: 131141.
Tylkowski T. 1987. Storing of Russian elm (Ulmus laevis Pall.) seed over
many years. Arboretum Kornickie 32: 297305.
Villiers TA. 1974. Seed aging: chromosome stability and extended viability of
seeds stored fully imbibed. Plant Physiology 53: 875878.
Wang BSP, Charest PJ, Downie B, comp. 1993. Ex situ storage of seeds,
pollen and in vitro cultures of perennial woody plant species. For. Paper
113. Rome: FAO. 83 p.
Willan RL., comp. 1985. A guide to forest seed handling, with special reference to the tropics. For. Paper 20/2. Rome: FAO. 379 p.
Zasada JC, Densmore RA. 1977. Changes in seed viability during storage
for selected Alaskan Salicaceae. Seed Science and Technology 5:
509518.
95
96
Chapter 5
Seed Testing
Robert P. Karrfalt
Mr. Karrfalt is the director of the USDA Forest Services National Seed Laboratory, Dry Branch, Georgia.
Contents
Introduction 98
Sampling 98
Sample Identification 100
Moisture Tests 100
Purity, Noxious Weed Content, and Seed Weight Tests 102
Purity Analysis 102
Noxious Weed Examination 103
Seed Weight Determination 103
Germination Testing 104
Vigor Testing 107
Chemical Staining for Viability 109
97
Introduction
Seed testing is the cornerstone of all other seed technologies. It is the means by which we measure the viability
and all the physical factors that regulate the use and maintenance of seeds. Everything that is done with seeds should
have some test information to guide the work and ensure
high quality. Seed tests tell if a crop of seeds is worth collecting, if handling procedures are correct, and how many
potential seedlings are available for regeneration.
The earliest form of seed analysis, the cut test, is still
often used today. Before seeds are collected in the field,
some seeds are cut open with a knife or razor blade to see if
their internal tissues are fully developed and undamaged.
This analysis is made more accurate in some cases by the
use of a hand lens. It is also used for simple analysis during
extraction and cleaning, or after germination to determine if
the ungerminated seeds have deteriorated or remained dormant. Although the cut test is often very good at producing
some information quickly, it is limited in the amount of
information it can supply and it lacks accuracy compared to
more sophisticated procedures. Therefore, it should never
be taken as a substitute for a formal laboratory analysis.
Sampling
Formal seed analysis begins with the sampling of the
seedlot. The Rules for Testing Seeds (AOSA 1996) and the
International Seed Testing Rules (ISTA 1996) both give
instructions on how to draw samples from a seedlot so that
the sample is representative of the entire seedlot.
Representative means that any tests conducted on this sample will accurately estimate the mean value of the lot
quality.
Sampling can be done with the hand or with a seed
probe, also known as a trier (figure 1). If a probe is used, it
must be long enough to reach to the farthest edge of the
container. A probe has gates that prevent seeds from entering until the probe is inserted the full dimension of the container. The probe should be inserted into the seed container
with these gates closed. Otherwise, seeds from the upper
layers will fill the probe as it is inserted and the bottom layers will not be sampled. Once the tip reaches the bottom or
far side of the container, the gates should be opened and the
probe gently turned back and forth to help the seeds fall in.
Then the gates should be closed gently, not forced, so that
any seeds that are caught in the opening and are preventing
the gates from closing fully (figure 2) are not crushed.
(Mechanically damaged seeds would bias the sample.) After
the probe has been withdrawn from the seed container, it
should be held horizontally, with the gates facing upward.
98
Then the gates should be opened gently and the probe shaken gently back and forth, so that seeds caught in the gates
will slip down into the probe and the gates can be safely
closed. Finally, the probe should be emptied by pouring the
seeds out the top of the probe and into a second container
(figure 3). This sample is the first primary sample.
If there is only 1 container, primary samples should be
taken until there are 5 of them. When more than 1 container
holds the seedlot, at least some of the other containers must
be sampled. When there are between 1 and 5 containers, all
containers should be sampled, at least 1 probe from each
container. When there are more than 5 containers, 5 of the
containers plus 10% of the remaining ones should be sampled. It is never necessary to sample more than 30 containers. (It would be rare that a forest seedlot would need 30
containers or more, or possibly even 20.) All of the primary
samples are then placed together to make up the composite
sample.
Sampling by hand is sometimes necessary when the
seeds will not flow into the probe because of their size,
shape, or surface texture. Sampling by hand can be done by
inserting the open hand (figure 4) into the seeds, closing it
once the point of sampling is reached, and then withdrawing
it closed. The seeds are then placed in a second container to
form the composite sample, just as in sampling with the
probe. At least 5 handfuls must be taken, and all levels must
be sampled. When the hand cannot be inserted into the
seedlot, the seeds can be poured from one container into a
second. The tester then should stop at a minimum of 5 evenly spaced intervals and remove a handful of the seeds for
the composite sample.
The composite sample, whether taken with a probe or
by hand, is usually too large to submit to a seed laboratory
for analysis. The composite sample is, therefore, mixed and
divided to obtain a submitted sample. This procedure is very
important and must be done correctly for the results to be
accurate.
The composite sample can be mixed either mechanically or by hand with rulers. Hand-mixing the composite sample is done by pouring the seeds into a cone on a flat, clean
surface. An open file folder makes a good work surface that
can be picked up to return the seeds to a container. With one
ruler held stationary against the seeds, the second ruler is
used to pull the outer edge of the pile up to the top of the
pile, allowing the seeds to roll down the sides and over the
top of the stationary ruler (figure 5). The full pile is thoroughly turned over and all layers mixed together. This procedure should be repeated for 1 full minute. Then the pile
should be divided by cutting the cone in half and then into
99
This amount is different for each species and the rules need
to be consulted to be sure the correct amount is submitted
for purity tests that are to be done according to the rules. A
smaller sample of seeds can be submitted, but the test will
not be according to the rules and the accuracy cannot be
assured to the same degree as a test that is done according
Figure 5Chapter 5, Seed Testing: seed can be handmixed before withdrawing a submitted sample from a
composite sample.
Sample Identification
Assignment of a test number is the first step in handling
every seedlot that is received in the laboratory. This number
allows for the orderly tracking of the test sample among the
other samples in the laboratory. A typical test number indicates the test year and an accession number. For example,
the 300th test conducted in 2005 would have a number such
as 05-300.
Figure 6Chapter 5, Seed Testing: the composite
sample is divided systematically into quarters, eighths,
sixteenths, and smaller fractions to obtain the submitted
sample at the seed storage plant or the working sample in
the laboratory.
Moisture Tests
Moisture tests must be the first tests conducted on samples when they arrive at the seed laboratory. Once a sample
container is opened and work begun, the seeds will likely
Figure 7Chapter 5, Seed Testing: a soil divider (left)
and a gamet divider (right), devices that systematically mix
and divide seed samples.
100
Some larger seeds and seeds with impermeable seedcoats need to be cut to make an accurate test (Bonner 1974,
1981, 1992). If the seed is not cut open, the moisture is not
freely released, and the moisture content is underestimated
(figure 10).
The oven method is not a direct measure of the content
of water. It measures weight loss that is assumed to be due
to the loss of water. A basic analytical procedure is required
to verify the temperatures and length of drying. The currently accepted procedure is the Karl Fisher procedure (figure
11) (Hart and Golumbic 1962). The moisture committee of
the ISTA uses this procedure in its work to standardize and
validate the oven procedures.
Another widely used method to measure seed moisture
is the electronic moisture meter. Although there are numerous brands of electronic moisture meters on the market, not
all of them will work for tree and shrub seeds (figure 12),
and those that do will not have calibrations for tree seeds.
Therefore, conversion charts must be developed for them by
testing samples with high to low moisture contents with
both the meter and the oven. A linear regression between
the oven and meter readings is calculated, and the conversion chart predicted from this regression (Bonner 1981; Hart
and Golumbic 1966; Jones 1960; Karrfalt 1987; Lanquist
1965). These meters provide quick results, are nondestructive to the seed, and are usually accurate to within 1% of
the moisture estimated by the oven method.
101
contains the minimum weight for conducting a purity analysis. Each species has its own specified minimum weight,
which has been determined to contain 2,500 seeds. The
mixing and dividing should be done in the same way as
described in the sampling section for drawing the submitted
sample from the composite sample. However, at this point it
is necessary to be very close to the minimum weight for
2 reasons. First, the analyst does not want to examine more
seeds than necessary, and second, the accuracy of the test is
evaluated using tolerance tables that were developed using
these minimum weights. Using substantially more seeds than
the minimum will invalidate the use of these tables.
Purity is determined differently by each of the 2 major
testing organizations. The ISTA rules specify a 3-part purity
and the AOSA rules specify a 4-part purity. The ISTA purity
values report percentages of pure seeds, other seeds, and
inert materials. The AOSA purity values report percentages
of pure seeds, weed seeds, other crop seeds, and inert materials. The pure-seed fraction consists of all those seeds that
are of the kind specified on the seedlots label. Specific
descriptions in the rules define pure seeds, but basically
the pure-seed fraction comprises whole seeds and seeds that
are not more than half broken away. Other seeds in the
ISTA rule are all kinds of seeds other than those listed on the
label. The AOSA rule makes a distinction between crop
seeds and weed seeds and uses a detailed list (AOSA
1995) to specify when a species is a weed and when it is a
crop. Weed seeds are mainly a problem in lots collected
from nets or directly from the ground. Contaminated cleaning equipment can also result in weed seeds entering a seed-
102
lot. Inert matter is all other material that is not classified as Figure 13Chapter 5, Seed Testing:
divided into its component parts.
crop seeds or other seeds. It could include soil particles,
stones, wire, small pieces of broken seeds, or other plant
parts. Purity is calculated by dividing the weight of the of
pure seeds by the total weight of all the fractions in the sample (figure 13) and is expressed as a percentage.
Purity work can often be tedious and very technical.
Devices such as the mechanical purity board (figure 14) can
speed up the procedure. The analyst must understand important taxonomy principles and accurately use the seed herbarium (figure 15) to identify all the kinds of seeds in the
sample.
Noxious Weed Examination
The noxious weed exam is a specialized purity examination. It is not a test traditionally associated with forest seeds
but may become more common as the commercial exchange
of native plants increases. A noxious weed is a highly
aggressive competitor or a plant with other highly objectionable characteristics, such as being poisonous. It is so offensive it has been put on a noxious weed list compiled by an
individual state or the federal government. A noxious weed
exam is made solely to identify the number of noxious weed
seeds found in the sample. Nothing else is noted in this
exam. The presence of any noxious weed seeds makes it illegal to sell the seeds until the noxious weeds have been
removed. The sample size for a noxious weed examination is
25,000 seeds.
Seed Weight Determination
The number of seeds per unit weight (kilogram and gram
or pound and ounce) is determined on the pure-seed fraction
from the purity test. This test is called the seed weight determination in the ISTA rules. It is made by counting out 8
replicates of 100 seeds and weighing them to the same precision as the weights for the purity test. The coefficient of
variation for these 8 values is computed. This coefficient
cannot be greater than 6 for chaffy seeds or greater than 4
for all other seeds. Otherwise, an additional 8 replications
need to be counted and weighed and combined with the first
8 weights. All 16 weights are then used to compute the
mean. Any weight diverging from the mean by more than 2
standard deviations is discarded; only the remaining weights
are used to compute the number of seeds per unit weight.
Seeds can be counted by hand, with a counting tray, a
shutter box, or a vacuum counter (figure 16). When seeds are
counted by hand, it is usually best to count out the appropriate number of piles of 10, 20, or 50 seeds, in order not to
lose ones place. A counting tray is simply a block of wood
a purity sample is
103
Seed weights are sometimes determined with an electronic counter (figure 18). The ISTA rule calls for counting
all pure seeds in the working sample when this is done. No
error-check is then made. A recent internal report made by
the Seed Count Committee of the Association of Official
Seed Analysts, augmented by the authors personal observations, suggests caution in the use of electronic counters for
seed weight determinations. A high potential for error in
counts exists. If carefully calibrated, these machines can
count quite accurately, but the machines need to be adjusted
and used correctly. A thorough evaluation of the degree of
desired accuracy and the amount of time required to achieve
it needs to be made before deciding to use the electronic
counter.
Figure 16Chapter 5, Seed Testing: seeds can be counted sometimes more quickly using a counting tray, a shutter
box, or a vacuum counter.
104
Germination Testing
Germination testing is designed to estimate the maximum number of seeds that will produce a normal seedling
and to give results that are as repeatable as possible. Without
uniform procedures, there would be no standard on which to
base the value of seedlots for commercial transactions and
the seed trade would be chaotic and filled with dispute.
Germination also tells a grower about a seedlots potential.
A seedlot with 80% germination cannot produce more than
80 seedlings per 100 seeds. Therefore, if 100 seedlings are
needed, a minimum of 125 seeds must be planted (100/0.80
= 125). How to use test data to compute sowing rates is presented in detail in chapter 7 (Nursery Practices) and later in
this chapter in the section on the use of test data.
The germination test is conducted on the pure-seed fraction from the purity test. Both the AOSA and ISTA prescribe the use of 4 replications of 100 seeds. These replications can either be planted 1 to a container (figure 19), 2 to
a container, or all on 1 tray. Alternatively, the 4 replications
can be further divided into smaller replications, but the total
number of seeds tested must remain 400 to remain in compliance with the rules. If fewer than 400 seeds are available,
then the number of seeds per replication should be reduced
so that an equal number of seeds is present in each of the 4
replications. Using fewer than 100 seeds in a replication is
not according to the rules, and the test would thus be unofficial. However, it is better statistically to have 4 replications
of 50 seeds each rather than 2 replications of 100 seeds
each. The 4 replications are then placed under optimal germination conditions for the period specified in the rules.
Germination is the number of normal seedlings produced
from 100 pure seeds expressed as a percentage. A normal
seedling has all the essential plant structures necessary for
the plant to continue to grow normally under favorable conditions (AOSA 1996; ISTA 1996).
Seeds can be planted in a number of ways. They can be
scattered or placed one at a time with forceps, although
more generally a vacuum counter or other type of planting
plate is used for speed and to ensure even spacing of the
seeds. The vacuum counter is the most expedient technique,
because it can handle a variety of seed sizes (figure 17).
Counting devices are described in the seed weight discussion above. Seeds should be hand-planted only when counting devices cannot be used in order to save time.
Seeds can be germinated on various media. Sand, sand
and perlite mixtures, potting mixtures, soil, and various
papersblue blotters, white blotters, or crepe-cellulose
papers (such as Kimpak)can be used (figure 20). Testing
rules, however, specify what is an acceptable medium for
the kinds of seeds tested. Specifying the medium helps
assure uniformity in test results. The blotters resist penetration by the roots of the plants, whereas the crepe-cellulose
paper allows for root penetration. Blotters offer the advantage of keeping the roots where the analyst can actually see
them for evaluation, but if a seedling is very large it will fall
over and tangle with other seedlings, making counts difficult. The media also differ in their water-holding ability.
Blotters usually need to be watered several times during the
test, whereas crepe-cellulose paper, sand, sand mixtures,
potting soils, and soil are absorbent enough to hold all the
water the seeds need for up to 3 months, if kept in a moisture-proof container. Watering the medium can be done by
hand or by machine. Watering by hand is usually done using
a squeeze bottle or a small hose from the tap and requires
subjectivity on the analysts part to estimate that the correct
amount of water has been applied. Too much or too little is
harmful, but in most cases there is wide latitude in the
amount that will give optimal results (Belcher 1975).
105
Figure 20Chapter 5, Seed Testing: seeds are germinated on various media, from right to left: crepe-cellulose
paper (such as Kimpak), blue blotters, sandperlite mixtures, and potting soil.
Figure 21Chapter 5, Seed Testing: an automatic pipetting machine can help to uniformly and rapidly water
germination dishes.
A species that does not require prechilling is called nondormant. If 10 to 14 days of prechilling are needed, the dormancy would be considered light. If 30 to 60 days of
prechilling are required to break the dormancy, it would be
considered moderate. More than 60 days of prechilling classifies the seedlot as highly or strongly dormant. The degree
of dormancy varies within the seedlot of even lightly dormant species; some seeds germinate without prechilling,
whereas other seeds in the same lot will not germinate until
they are prechilled. However, the term variable dormancy is
usually reserved for seedlots in which some seeds germinate
during prechilling, whereas other seeds in the same lot will
not germinate even after being placed in favorable germination conditions. Species that fit the deep and variable dormancy category are Rocky Mountain juniper (Juniperius
scopulorum Sarg.) and basswood (Tilia americana L.).
Because of the above-mentioned variation in dormancy,
seedlots will often be tested with and without prechilling or
with varying lengths of prechilling. Such tests are referred to
as paired or double tests; usually only 2 tests are done. More
tests, of course, can be and are done with some seedlots.
This type of testing can determine the presence of dormancy, the strength of dormancy, or a weakness in the seeds
(Belcher 1995). When the seedlot has the same germination
with and without prechilling, it is said to be nondormant.
When the germination is increased with prechilling, the
seedlot is classified as dormant; the longer the prechilling
period needed, the stronger the dormancy is said to be. A
decrease in germination with prechilling is an indication of
weakness in the seeds. This last condition is similar to the
situation of the type of vigor test known as the cold test,
which is described in the following section.
Prechilling is not the only treatment to break dormancy.
Light is useful to break dormancy and can reduce the need
for prechilling. Birches (Betula L.) and loblolly pine (Pinus
taeda L.) are prime examples where light helps break dormancy. Seedcoat dormancy is treated by scarifying the seedcoat with either acid, bleach or mechanical means. Chemical
stimulates such as gibberellins or potassium nitrate have
been little used with forest tree seeds.
Vigor Testing
Sometimes standardized laboratory germination procedures are criticized as not predicting field performance very
well (Moreno 1985; Stein 1967). These critics suggest using
a variety of test conditions to find an optimum for each
seedlot. The problem in predicting field germination is that
107
108
This greater leakage causes the water to have a higher conductivity, which can be measured with a conductivity meter
(figure 25). Bonner and Agmata-Paliwal (1992) reported on
the use of conductivity for tree seeds and found that results
have poor repeatability for precise estimates but possibly
would work for general estimates of classes as poor, low,
intermediate, or high viability.
Several statistics have been put forward to use speed of
germination as an indicator of vigor. The faster a seedlot
completes germination or reaches its peak, the more vigorous it is said to be. The simplest indicator is days to 90% of
total. For example, if the final germination is 88%, the indicator would be how many days it takes to reach 79% germination. A lot that reaches 79% in 12 days would be more
vigorous than one that takes 16 days. To use this statistic,
counts must be made quite frequently, even daily, or the
data must be interpolated to determine the number of days
to the specified germination.
Czabators factor (1962), developed for use with southern pines, combines the maximum daily average germination, called the peak value, and the average daily germination at the end of the test to form one statistic called the
germination value. Germination is counted frequently, at
least every third day, and the cumulative germination on
each day is divided by the number of days that the test has
been run in order to compute the mean daily germination
for that day. For example, if on day 22 the cumulative germination is 88, the mean daily germination is 4. This mean
Figure 24Chapter 5, Seed Testing: the accelerated
aging test is conducted by placing seeds in a plastic box
with a water reservoir and holding them at 40 C for
72 hours.
109
110
X-Radiography
X-radiography is very useful in forest seed analysis. It
provides a very rapid and accurate analysis of the internal
structure of seeds, identifying empty, insect-damaged, or
poorly developed seeds (figure 30). It is an immense help in
judging maturity, determining how many bad seeds should
be remove, and detecting any mechanical injury. It is more
accurate than cutting tests in many cases, requires much less
time, and is nondestructive (AOSA 1979; Simak and others
1989). X-radiography was first applied to tree seeds by
Simak in Sweden. The use of contrast agents has improved
the ability of the x-ray test to discriminate between viable
and nonviable seeds with some species (Kamra 1963; Simak
1957; Vozzo 1978). A contrast agent enters damaged areas
of the seed differentially from nondamaged areas, making
the damaged areas more radiopaque. They will then appear
as bright areas on the radiograph. Aqueous solutions of
heavy salts such as iodine or barium chloride and vaporous
agents such as chloroform have been used as contrast
agents.
Radiographs can be made on Polaroid film, x-ray
paper, or x-ray film. Polaroid film is useful if no darkroom
is available, because the film is developed in the light, just
like a Polaroid photograph. The disadvantages of Polaroid
are high cost, short shelf life, and lack of detail. X-ray paper
is fast to use but does require a simple darkroom. It is less
expensive than Polaroid, has a shelf life of several years in
cold storage (3 C), and much better resolution. The best
resolution is obtained with x-ray film. X-ray film, however,
111
seeds that failed to germinate. Empty seeds will never germinate, and damaged or poorly developed seeds will seldom
germinate. The excised embryo or tetrazolium test for difficult-to-cut seeds can be speeded up by x-radiography. The
seeds are first placed on the x-ray film or paper in a manner
that will allow the comparison of the exact image to the
exact seed. This is done by placing the seeds on an additional piece of paper before placing the paper on the x-ray film
or paper. If orientation of the seed is important, as in
double-seeded fruits such as dogwood, the seeds can be
placed on adhesive tape and that then laid on the paper. The
seeds should be oriented so that both seeds in the fruit can
be viewed and the tape prevents them from turning. After
the radiograph is made, the seeds are gently slipped off the
x-ray paper so that the seeds are kept in order for cutting.
Only those seeds that are morphologically sound in the
radiograph need to be cut.
Other Quick Tests
As stated in the introduction, cutting tests are very limited in their application. However, they can provide useful
information on full seed percentages and the condition of
the internal structures. For example, color of the tissue cannot be determined in a radiograph, which is only black and
white. Seeds that are cut and found to be dark are not likely
to germinate. New and unfamiliar images in a radiograph
require cutting the seed to determine what is actually in the
112
Sowing Rates
A sowing rate is the amount of seeds sown in a unit area
of nursery bed to produce the desired number of seedlings.
The following formulas show how seed test data are used to
determine this rate.
Weight of seeds to sow in a nursery bed
(width x length) is equal to
(bed width x bed length x seedlings desired per area)
(germination x seeds per weight x purity x survival factor)
Number of seeds needed to sow per area of nursery
bed is equal to
(seedlings desired per area) (germination x survival
factor)
In both of these formulas, the survival factor is the ratio
of the number of seedlings expected to the number of viable
seeds planted. It is derived from experience in the given
nursery and should be constantly updated with new information collected from history plots. History plots are permanent sample plots in a nurserybed used for carefully monitoring the number of seeds sown and the number and quality
of seedlings produced (Landis and Karrfalt 1987). For
example, if 100 seeds are sown on a square foot, germination is 80% in the laboratory, and 60 seedlings actually grow
on the square foot, then the survival factor is 60 80 or
.75 (75%).
Computing sowing rates for containers is somewhat different, because we must predict the probability of an empty
cell in the container. The probability that a container cell is
empty is equal to 1 minus the probability that at least 1
seedling is in the cell. Sowing 1 seed per cell, this probability is 1 minus germination. With a 90% germination, the
probability of an empty cell following single-seed sowing is
0.1. In sowing 2 seeds per cell, the probability of no
or
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114
to be measured: germination, seed weight, purity, and moisture content. Moisture needs to be monitored during storage
to be sure it is being properly maintained. If there are no
changes in moisture content, then seed weight and purity
will not change and viability will change very slowly, if at
all. Some annual monitoring of seed moisture is necessary
to ensure that storage conditions are being adequately maintained. The viability can be retested at 3- to 5-year intervals
with a current test always done no more than 6 to 9 months
before sowing. If a longer time passes before sowing, some
deterioration could occur, resulting in changes in germination. Determining viability in some seeds takes a long time,
and thus it is important to schedule adequate lead time into
the production schedule.
References
Anderson HW, Wilson BC. 1966. Improved stratification procedures for
western white pine seed. Pub 8. Olympia: Washington State Department
of Natural Resources.
AOSA [Association of Official Seed Analysts]. 1979. X-ray handbook.
Lincoln, NE: AOSA.
AOSA. 1983. Seed vigor testing handbook. In: Contrib. 32. Handbook on
seed testing. Lincoln, NE: AOSA. 88 p.
AOSA. 1996. Rules for testing seeds. Journal of Seed Technology 16(3):
1113.
AOSA. 1995. Uniform classification of weed and crop seeds. In: Larson AL,
Wiersema JH, Handwerker T, eds. Contrib. 25. Handbook on seed testing. Lincoln, NE: AOSA.
AOSA. 2000.Tetrazolium testing handbook. Contrib. 29. Handbook on seed
testing. Lincoln, NE: AOSA. 302 p.
Belcher EW. 1975. Influence of substrate moisture level on the germination
of seed of selected Pinus species. Seed Science and Technology 3(3/4):
597604.
Belcher EW. 1978. Aspects of seed quality. In: Proceedings, Western Forest
Nursery Council and Intermountain Nurserymans Association
Combined Nurserymans Conference and Seed Processing Workshop.
1978 October; Eureka, CA. D.54D.59.
Belcher EW. 1995.The effect of seed condition and length of stratification on
the germination of loblolly pine seed.Tree Planters Notes 46(4):
138142.
Belcher EW, Karrfalt RP. 1979. Improved methods for testing viability of
Russian olive seed. Journal of Seed Technology 4(1): 5764.
Bonner FT. 1972. Measurement of moisture content in seeds of some
North American hardwoods. Proceedings of the International Seed
Testing Association 37(3): 975983.
Bonner FT. 1974. Determining seed moisture in Quercus. Seed Science and
Technology 2(3): 399405.
Bonner FT. 1981. Measurement and management of tree seed moisture.
Pap. SO-177. New Orleans: USDA Forest Service, Southern Forest
Experiment Sation. 10 p.
Bonner FT. 1984. Tolerance limits in measurement of tree seed moisture.
Seed Science and Technology 12(3): 789794.
Bonner FT. 1992. Moisture content chapter In: Gosling PG. Report of the
Forest Tree and Shrub Seed Committee 19891992. Seed Science and
Technology 20(Suppl. 1): 8384.
Bonner FT. 1998. Testing tree seeds for vigor: a review. Seed Technology
20(1): 517.
Bonner FT, Agmata-Paliwal A. 1992. Rapid tests of seed quality in Picea
species by the leachate conductivity method. In: DeHayes DH, Hawley
GJ, eds. Genetics in forest biology; Proceedings, 1st Northern Forest
Genetics Association Conference; 1991 July 2325; Burlington,VT. Berea,
KY: Northern Forest Genetics Association: 6975.
Buszewicz G. 1962. A comparison of methods of moisture determination
for tree seeds. Proceedings of the International Seed Testing Association
27: 952961.
Ching TM, Parker MC. 1958. Hydrogen peroxide for rapid viability tests of
some coniferous tree seed. Forest Science 4: 128134.
Czabator FJ. 1962. Germination value: an index combining speed and completeness of pine seed germination. Forest Science 8: 386396.
Flemion F. 1948. Reliability of the excised embryo method as a rapid test
for determining the germinative capacity of dormant seeds.
Contributions of the Boyce Thompson Institute 15: 229241.
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116
Chapter 6
Contents
Introduction 118
Certification in Agriculture 118
Certification in Forestry 118
Pacific Northwest 120
Audit class 120
Source-identified class 120
Selected class 120
Tested class 120
Southeast 121
Source-identified material 121
Selected material 121
Cerified material 121
117
Introduction
Seed certification is a system that provides assurance to
buyers that the seeds being purchased are what they are represented to be by the producer or seller. This certification of
identity (and sometimes quality) is typically provided by an
independent third party for a fee that is charged to the producer and becomes part of the production costs. The system
is simple and effective; it is used all over the world in various, yet basically similar, forms for agricultural, horticultural, and forestry seeds or other propagules. This chapter will
briefly describe how seed certification developed in forestry
and how it is practiced today with seeds of woody plants.
More detailed historical accounts of agricultural and forest
tree seed certification in the United States can be found in
Hackleman and Scott (1990) and Rudolf (1974).
Certification in Agriculture
Certification of agricultural seeds has been practiced in
the United States since the early 1900s (Copeland and
McDonald 1995; Hackleman and Scott 1990) and has been
a positive force in the development of modern agriculture.
Certification in individual states is typically controlled by an
agency that is authorized by the state to carry out the procedures. The agencies are commonly called crop improvement
associations, but some carry other designations. The organizational structures of these agencies may vary, but their
goals are similar and they act cooperatively through the
Association of Official Seed Certification Agencies
(AOSCA 1994). AOSCA establishes minimum certification
standards for all types of plant materials. The member state
certification agencies may develop their own certification
standards for different materials, but their standards must
equal or exceed those of AOSCA. Agricultural seed certification is an assurance of the varietal (genetic) identity of the
material, but it is also normally a de facto assurance of
genetic quality. Developers of improved varieties of agricultural species (state land grant universities or private seed
companies) widely publicize the results of their field trials,
and the expected performance of new varieties is well
known and documented before they reach the market. Seed
buyers want the assurance from certification that their seeds
are really the variety that the producer says they are. The
slightly higher cost required for this assurance is gladly
paid.
Certification in Forestry
Certification of forest reproductive material has developed in a slightly different manner from that of agriculture
in this country. Most forest landowners do not have ready
118
(Rudolf 1974). The use of seed zones (figures 14) has been
very effective, and they are still widely used today. The
major impetus for forest seed certification, however, came
from the expanded reforestation programs and rapidly developing tree improvement programs in the 1950s and 1960s.
Establishment of seed orchards of major species with selected phenotypes and the subsequent progeny tests with their
offspring has led to wide-scale use of improved families and
clones in forest regeneration. When foresters wanted certification of this material, they turned to the state crop improvement associations, because these agencies were the only
ones legally permitted in most states to perform certification
services. As these services were extended to forestry material, the mechanisms for implementation developed differently
in different parts of the country.
South Dakota established the first forest tree certification program in 1952 for stock selected for shelterbelt use
(Rudolf 1974). Georgia established the next program in
1959 with comprehensive certification standards for tree
seeds (GCIA 1959). The AOSCA (then known as the
International Crop Improvement Association) adopted
almost identical standards also in 1959 (Rudolf 1974), with
later revisions in 1962, 1966, and 1970 (Hackleman and
Scott 1990). Other states were not far behind. In 1994,
AOSCA widened the scope of its tree seed certification standards to allow certification of material from all native
plantstrees, shrubs, vines, forbs, and grasses (AOSCA
1994). These standards were designated for pre-variety
germplasm certification and will be explained in a later
section.
119
Pacific Northwest
An organized effort to improve tree seed supplies in the
Pacific Northwest came about in the mid-1950s with the formation of the Northwest Forest Tree Seed Committee at
Corvallis, Oregon (Edwards 1981). This group later became
the Western Forest Tree Seed Council in affiliation with the
Western Forestry and Conservation Association. The bumper
cone crop of 1966 underlined the need for certification programs (Hopkins 1968), and the council and the Western
Reforestation Coordinating Committee of the Western
Forestry and Conservation Association took action. Through
their efforts, the Northwest Forest Tree Seed Certifiers
Association (NWFTSCA) was formed in 1966 to promote
seed certification. This organization developed seed zone
maps for Washington and Oregon and the framework for a
seed certification system (Edwards 1981).
Certification was jointly administered by the
Washington State Crop Improvement Association and the
Oregon Seed Certification Service, a division of the
Department of Crop and Soil Science at Oregon State
University. The NWFTSCA provided review and advice to
the agencies. Their system recognizes the following categories of reproductive material, which are indicated by standardized color-coded labels affixed to seed containers.
Audit class. Certifying authorities have reviewed
records indicating seed lot origin and collection documentation on where and when the seeds were collected. Origins
are usually less specifically identified than those in the
source-identified class. Labels placed on the seed containers are brown and white.
Source-identified class. Reproductive material comes
from a seed zone defined by a legal description and from
within a 154-m (500-ft) elevation band. The seed zones
were defined on the Tree Seed Zone Map issued by the
Western Forest Tree Seed Council in 1973. They are based
on physiographic and geological provinces of Washington
and Oregon as defined by Franklin and Dyrness (1973). Two
subclasses are recognized: (a) personally supervised productionboth the producer and the certifying agency have personal knowledge of the seed zone from which the seeds
were collected and (b) procedurally supervised
productiononly the buyer and not the certifying agency
determines if the collections are properly identified. Labels
for both subgroups are yellow.
Selected class. Reproductive material comes from
trees that were selected for a specific character(s). Two
classes are recognized: (a) reproductive material obtained
from selected trees recognized to be superior for any number
of traits, such as volume, form, or disease resistance and (b)
120
material from untested seed orchards and from seed production areas, and open-pollinated seeds from individual selected trees. In this class, only the female parent is known. This
class of material has promising traits that may be superior in
the offspring, but such superiority has not been determined
by testing. Details of the parents must be recorded. Labels
are green.
Tested class. Reproductive material comes from
selected trees that have been tested for performance of specific characteristics, as determined by progeny or other
applicable tests under specified conditions. The material
from this class that performs best in the tests is presumed to
be the ultimate in promised genetic superiority and is similar
to the agricultural class certified seed, indicating the highest degree of improvement. Labels are blue.
To be able to sell certified material, the producer or collector must submit an application to the appropriate seed
certifying agency, along with a fee, stating what material is
to be certified. The application must spell out how the material will be produced or collected. The certifying agency will
then notify the applicants if their plans are acceptable or in
need of modification. The agency performs field and seed
plant inspections, seed storage inventory audits, and whatever additional steps are necessary to ensure that the materials
meet the agencys standards and can be tagged with the official agency labels. Cone collectors and buyers have slightly
different registration and inspection procedures than seed
orchard producers, but the agency controls are comprehensive. Procedures may be amended from time to time, so
interested parties should check with their respective state
certifying agencies to get the current regulations.
The species of greatest interest for certified collections
have traditionally been Douglas-fir (Pseudotsuga menziesii
(Mirb.) Franco), western white pine (Pinus monticola
Dougl. ex D. Don), ponderosa pine (P. ponderosa P. & C.
Lawson), sugar pine (P. lambertiana Dougl.), and lodgepole
pine (P. contorta Dougl. ex Loud.). Another dozen or more
species, both hardwoods and conifers, are occasionally
certified.
Most of the tree seeds sold in the Pacific Northwest are
exported to countries in northern Europe that are members
of OECD (Organization for Economic Cooperation and
Development), a United Nationsbased international economic development organization that has set up the Scheme
for Control of Forest Reproductive Material Moving in
International Trade. European countries that import tree
seeds from the Pacific Northwest are required to have
OECD certificates on their seeds; consequently, procedures
were established in the Northwest to implement this scheme
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122
All of these types are defined very much like the corresponding classes in tree seed standards that were described
earlier. To satisfy OECD export standards, AOSCA will
allow a pink label to go on Selected material if it is equal to
the OECD class untested seed orchard. This class of material is collected from orchards for which progeny tests are not
yet complete. Even though the current demand is for sourceidentified materials, the mechanism now already exists for
certification of any improved native plant materials when
producers are ready. For information on certification of
native plant materials, including trees, consult the AOSCA
123
References
ACIA [Alabama Crop Improvement Association]. nd. Certification standards for forest reproductive material in Alabama. Auburn, AL: Alabama
Crop Improvement Association. 24 p.
AOSCA [Association of Official Seed Certification Agencies]. 1994.
Certification handbook. Pub. 23. Mississippi State, MS: Association of
Official Seed Certification Agencies. 288 p.
Copeland LO, McDonald MB. 1995. Principles of seed science and
technology. 3rd ed. New York: Chapman and Hall. 369 p.
Edwards DGW. 1981. The Western Forest Tree Seed Council. In:
Proceedings, Intermountain Nurserymans Association and Western
Forest Nursery Association Combined Meeting; 1980 Aug. 1214; Boise,
ID. Gen.Tech. Rep. INT-109. Ogden, UT: USDA Forest Service,
Intermountain Forest and Range Experiment Station: 2931.
Franklin JF, Dyrness CT. 1973. Natural vegetation of Oregon and
Washington. Gen.Tech. Rep. PNW-8. Portland: USDA Forest Service,
Pacific Northwest Forest and Range Experiment Station. 417 p.
GCIA [Georgia Crop Improvement Association]. 1959. Certification
standards for forest tree seed.Tree Planters Notes 36: 39.
Hackleman JC, Scott WO. 1990. A history of seed certification in the
United States and Canada. Raleigh, NC: Association of Official Seed
Certification Agencies. 97 p.
Hoekstra PE. 1976. Certification of source-identified tree seed under the
OECD scheme in Oregon and Washington, U.S.A. Washington, DC:
USDA Forest Service. 8 p.
Hopkins HG. 1968. Forest tree seed certification in the Pacific Northwest.
Journal of Forestry 66(5): 400401.
Karrfalt R. 1998. Personal communication. Dry Branch, GA: USDA Forest
Service, National Tree Seed Laboratory.
Lantz CW, Kraus JF. 1987. A guide to southern pine seed sources. Gen.
Tech. Rep. SE-43. Asheville, NC: USDA Forest Service, Southeastern
Forest Experiment Station. 34 p.
MSIA [Mississippi Seed Improvement Association]. 1979. Certification standards for forest reproductive material in Mississippi. Mississippi State, MS:
Mississippi Seed Improvement Association. 24 p.
124
Chapter 7
Nursery Practices
Thomas D. Landis
Dr. Landis retired from the USDA Forest Service, State and Private Forestrys Reforestation,
Nurseries, and Genetics Research National Team
Contents
Introduction 126
Terminology 126
The Target Seedling 127
Morphological Specification 127
Physiological Qualities 128
Genetic Considerations 129
Types of Nurseries 129
Propagation Options 130
Seed propagation 130
Direct seeding 130
Planting germinants 131
Transplanting emergents 132
Transplanting seedlings 132
Vegetative Propagation 133
125
Introduction
Plant propagation is both a science and an art. In this
chapter, we examine the science of plant propagation, which
consists of a knowledge of plant physiology, nursery cultural practices, and the biological characteristics of the particular plant that we want to grow. The art of plant propagation
cannot be taught, however, because it consists of certain
technical skills that must be acquired through experience
and often requires a certain feel. This special quality is
expressed in the saying that people who seem to be able to
grow plants have a green thumb (Landis and others 1999).
But before we get into the specific details of plant propagation, we first need to cover some basic nursery terms.
Terminology
A seedling is a plant grown from a seed, but the term is
commonly used generically for many types of nursery
stock, including transplants, rooted cuttings, and emblings
(plants that are produced through micropropagation). Forest
and conservation seedlings are traditionally divided into 2
basic stocktypes, depending on how they were propagated:
bareroot seedlings and container seedlings. Bareroot stock
is grown in soil in open fields (figure 1A), and the seedlings
are removed from the soil during harvesting (figure 1B).
Container seedlings are grown in an artificial growing medium in a controlled environment, such as a greenhouse (figure 2A), where most or all of the growth-limiting factors
can be manipulated. Because the volume of growing medium in containers is relatively small, roots bind the medium
into a cohesive plug by the time the seedlings are harvested (figure 2B). Therefore, container-grown stock are sometimes called plug seedlings.
Another stock type is the transplant, a seedling that has
been physically removed from its seedbed or container and
then replanted in another location for additional growth.
Traditionally, most transplants are bareroot seedlings that
were grown for 1 or 2 years and then replanted into a transplant bed and allowed to grow for another year or two.
Recently, container transplants are becoming much more
popular. This new stock type, also called a plug transplant,
is produced by transplanting a small container seedling into
the bareroot nursery for an additional year or two of growth.
Bareroot seedlings have been traditionally described
with a numerical code. The first number corresponds to the
number of years in the seedbed, and the second number
refers to the number of years in the transplant bed. Bareroot
seedlings are generally produced in 1 to 3 years (1+0 to
3+0), and transplants require 2 to 4 years (for example, 1+1
126
127
128
Genetic Considerations
Most seedlings grown for forest and conservation purposes are ordered by species, stock type, and seed zone or
seed source. A seed zone is a geographic area that is relatively similar in climate and soil and often is described by a
numerical code. Seed zones in mountainous terrain are also
stratified by elevation (figure 5A). For example, the geographically diverse state of California has more than 80 different seed zones, with numerous elevation bands within
each zone. All seeds and cuttings collected in a particular
zone are labeled with that source code so that all seedlings
produced from them will be planted back into the zone of
origin. When a seedling order is sown in the nursery, information on species, seed zone, and elevation is included into
a seedlot identification number. The seedlot number remains
with this group of seedlings throughout their entire nursery
tenure and is marked on the storage container when the
seedlings are harvested for outplanting (figure 5B).
Types of Nurseries
Once the target seedling has been defined, the next step
is to decide how best to grow it. Forest and conservation
stock is propagated in either bareroot or container nurseries,
and the choice is determined by several factors:
1. Cost. Container seedlings traditionally have been
more expensive than bareroot stock, although, in
recent years, the costs are becoming more comparable.
Container nurseries also are more cost-effective at low
seedling production levels.
2. Species characteristics. Most forest and conservation species can be grown as bareroot seedlings,
although some do better in containers.
3. Production time. Because container seedlings can
be produced more quickly than bareroot seedlings,
they are often used to reforest burns and other sites
that need to be planted quickly.
4. Outplanting site condition. Bareroot seedlings are
used on typical reforestation sites, but container
seedlings often are preferred for the more severe, hardto-plant sites. Container stock has a wider outplanting
window than bareroot stock.
5. Personal preference. Some customers tend to prefer
one stock type over the other.
Because bareroot seedlings are grown in open fields, the
soil, water supply, and climate of the nursery site must be
suitable for propagation. The growth rate of bareroot
seedlings and the length of the growing season are largely
B
controlled by the climate at the nursery site. Quality nursery
soils are difficult to find in convenient locations, and good
agricultural land is often expensive. Compared to container
nurseries, bareroot nurseries usually require considerable
capital to develop but have lower operating costs. A comprehensive discussion of site selection factors that should be
evaluated when locating a bareroot nursery is presented in
Duryea and Landis (1984) and Lantz (1985).
Container nurseries can be constructed on land with low
agricultural value that would be unsuitable for bareroot
seedling production. The amount of capital investment and
129
Propagation Options
To determine which type of propagation method will be
most effective and economical, both the biology of the plant
and the objectives of the outplanting project must be considered (figure 6). As mentioned in the target seedling section,
management objectives have a critical influence on the
selection of propagation system. Most of the commercially
important tree species used in plantation forestry can be
grown from seeds, but a few are vegetatively propagated on
Figure 6Chapter 7, Nursery Practices: nursery managers must consider many biological, operational, and economic factors before deciding on the best propagation system for a given plant species.The first and most important
decision is whether to use seed or vegetative propagation.
130
Table 1Chapter 7, Nursery Practices: operational considerations when choosing propagation methods
Considerations
MANAGEMENT
Seeds
Cuttings
Micropropagation
OBJECTIVES
Fast growth
Biodiversity
Availability of propagules
EASE
OF PROPAGATION
Difficulty
Relatively easy
Minimal
BR = seasonal
C = year-round
Low
which they are adapted. In the Pacific Northwest, some nurseries sow literally hundreds of different seedlots each year,
reflecting the many diverse environments in that mountainous terrain. In the South, some nurseries propagate by families and the seedlots from each family are sown and cultured
separately.
Many forest and conservation seeds have some type of
seed dormancy that keeps them from germinating when
placed under unfavorable environmental conditions. Growers
need to understand the dormancy characteristics of the seeds
that they are trying to germinate, because the type of presowing treatment differs for each. For example, some plants
exhibit seedcoat dormancy, which means that the seeds are
impermeable to the water and/or oxygen that the embryos
need to initiate germination. Culturally, there are a couple of
ways to overcome this problem. Scarificationany treatment that breaks down the seedcoats to allow penetration of
water and oxygencan be either mechanical or chemical.
Mechanical scarification consists of physically scratching
the seedcoat to reduce its thickness, and chemical scarification involves dissolving the seedcoat with caustic chemicals
such as acids. Hot water or steam can also be used to soften
hard seedcoats.
Another common presowing seed treatment is chilling or
stratification, which consists of keeping seeds under a cool,
moist environment for a specified period of time. The term
stratification comes from the practice of placing layers of
seeds between layers of insulating material that keep them
131
132
Bareroot or
container
Container
Planting germinants or
sowing sprouts
(pregerminated seeds are
sown from stratification
trays or bags)
Transplanting emergents
or pricking out (seeds are
sown into trays & then young
emergents are transplanted)
Transplanting seedlings
(established seedlings are
re-planted into a transplant
bed or container)
Container
Bareroot or
container
Direct seeding
(seeds are sown with
or without pretreatment)
Type of nursery
Propagation method
Best use
Advantages
Table 2Chapter 7, Nursery Practices: characteristics of seed propagation methods for forest and conservation species
Disadvantage
wheel with clips to place the transplant into the slit at the
proper spacing. Transplant beds have the same physical
dimension as seedbeds but seedlings are planted in fewer
rows and at much lower densities. Once they become established, transplants are fertilized and irrigated and are given
the same root culture treatments as bareroot seedlings.
The first container transplants were made from surplus
stock but now seedlings are grown specifically for this purpose. Typically, seedlings are grown in 33- to 66-cm3 (2- to
4-in3) containers for 4 to 6 months and are then transplanted
to the beds. Miniplugs, the newest type of plug transplant,
are grown in very small containers (around 16 cm3 or 1 in3)
specifically for transplanting. Plug transplants are cultured
and harvested in exactly the same manner as are bareroot
transplants. Another new larger container stock type is made
by transplanting miniplugs into much larger containersfor
example, 328 cm3 (20 in3).
A complete discussion of seed propagation is provided
in Landis and others (1999).
Vegetative Propagation
The second major plant propagation technique is vegetative propagation, which also is called asexual propagation
because 2 parents are not required. A clone is defined as a
group of genetically uniform individuals that were originally
derived from a single parent by asexual propagation. The
major benefit of vegetative propagation is that the offspring
will very closely resemble the parent because their genetic
code is identical (figure 7). Other benefits of vegetative
propagation include the following:
1. The ability to obtain a high degree of crop uniformity.
2. The elimination of problems with seed availability,
dormancy, and viability.
3. The ability to perpetuate genetically superior plants,
such as fast-growing or disease-resistant clones.
4. The ability to bulk-up valuable, genetically improved
seedlots.
In forest and conservation nurseries, rooted cuttings are
the most common type of vegetative propagation, and there
are 3 different types that are named for the type of tissue
used. Hardwood cuttings (figure 8A) are collected during
the dormant period from the last seasons growth, stratified
in cold storage, and planted (stuck) in containers or bareroot beds. Semi-hardwood cuttings are collected after the
active growth period from hardened woody tissue of the current seasons growth. Softwood cuttings are collected from
soft succulent new shoots of woody plants that have just
begun to harden, normally in spring, but also at any time of
Figure 7Chapter 7, Nursery Practices: plants propagated from seed appear different from their parents,
because they contain a mixture of genetic characteristics
(Top). Vegetative propagation, on the other hand, produces
exact duplicates of the parent plant (Bottom).
133
Figure 8Chapter 7, Nursery Practices: hardwood cuttings are the most common vegetative propagation method
used in forest and conservation nurseries (A). Hardwood
or semi-hardwood cuttings are typically treated with rooting hormones and planted into bareroot beds (B) or
containers (C).
Figure 9Chapter 7, Nursery Practices: growing schedule are used in crop planning to illustrate the time required
for each phase of the nursery cycle from seed procurement
to outplanting.
134
135
Figure 12Chapter 7, Nursery Practices: seed use efficiency, the ratio of seeds sown to seedlings harvested, is a
function of both seed quality and seedling losses during the
growing season (from Thompson 1984).
136
137
Figure 14Chapter 7, Nursery Practices: harvesting equipment lifts bareroot seedlings by undercutting them and
loosening soil from around the roots with vibration (A). During hand-lifting, workers pull seedlings from the seedbed and
place them in tubs, being careful to avoid excessive exposure and desiccation (B). Seedlings are then taken to the packing
shed where they are counted and graded (C) to predetermined morphological specifications that were agreed upon by
the nursery manager and the customer: shoot height, stem diameter (caliper), and some measure of root size and fibrosity
(D). Finally, the shippable seedlings are sealed into moisture-retentive bags or boxes for refrigerated storage (E).
139
Types of Containers
There are many different types of containers, with
capacities ranging from as small as 16 cm3 (1 in3) to more
than 492 cm3 (30 in3). The most commonly used container
types include Styrofoam blocks, book planters, and several
types made of molded hard plastic. Other growing containers, such as peat plugs and plastic bags, are sometimes used,
but these lack vertical ribs on their insides for controlling
root spiraling. The best type of container depends on available nursery equipment, the species of plant, and conditions
at the outplanting site. Hardwood species must be grown in
relatively larger containers than conifers because their large
leaves intercept irrigation and create more shade competition
with their neighbors. Foresters prefer seedlings grown in
smaller containers for moist outplanting sites, but demand
larger container stock for harsh dry conditions or sites with
heavy brush competition. New container types are continual140
Growing Media
Almost all container nurseries use some type of artificial
growing medium instead of native soil. An ideal medium
should be sterile, lightweight, porous, and consistent in
quality. Several different brands of media are commercially
available, and most are composed of sphagnum peat moss,
vermiculite, and sometimes perlite, composted bark, or sawdust. Some nurseries mix their own growing media. Larger
nurseries have specially designed mixers for blending the
components, and some have customized equipment using
cement mixers and so forth. Some components of growing
media, such as vermiculite and perlite, are inherently sterile;
sphagnum moss, however, may contain pathogenic fungi.
Chemical fumigants or steam heat are typically used to sterilize media. Fertilizers or other chemical amendments are
sometimes added to growing media during the mixing
process. Dolomitic limestone is used to supply calcium and
magnesium and raise the low pH. Slow-release fertilizers,
such as Osmocote, are composed of resin-coated pellets
that release mineral nutrients in response to temperature and
moisture.
Containers are filled with growing medium in several
different ways. Smaller nurseries fill containers by hand.
Automated filling machines that do everything from filling
and tamping the medium to sowing and covering the seeds
can also be used. A complete discussion of container types
and growing media can be found in Landis and others
(1990).
Sowing and Thinning
The number of seeds to sow per container cavity is calculated using the seed germination percentage, with the
objective of having no empty cavities. Containers can be
sown by hand, which is necessary for very large or irregularly shaped seeds, or with various sowing machines. The shutterbox (figure 16A) consists of a template with a set of
predrilled holes that correspond to the pattern of the individual container cavities. The size of the holes in the shutter
control the sowing rate, usually from 2 to 6 seeds per hole
depending on seed quality (figure 16B). Vacuum seeders
have plates or drums that hold a certain number of seeds
until they are released into the containers. Precision sowing
machines can accurately control the sowing density down to
141
Pest Management
factors for inducing hardiness and dormancy are cooler temperatures, mild moisture and nutrient stress, and shortened
photoperiod. Seedlings in fully enclosed greenhouses are
often moved to a shadehouse at this time, where the change
in temperature and humidity aid the hardening process.
Growers with shelterhouses permanently raise the sides to
expose the crop to ambient conditions. At the same time, the
photoperiod lights are shut off and the fertilizer mix
changed to a special low-nitrogen hardening formula.
Harvesting, Grading, Storing, and Shipping
The harvesting method is related to the type of seedling
storage. Some nurseries store their container seedlings outside to overwinter in sheltered storage, being particularly
careful to insulate the root systems against cold. Seedling
roots are much less cold-tolerant than shoots and can be
damaged or even killed at temperatures that are only a few
degrees below freezing. Other nurseries grade their
142
Figure 18Chapter 7, Nursery Practices: many container seedlings are harvested by pulling them from the containers (A), wrapping them in plastic film or placing them in
bags, and then storing them under refrigeration (B).
are a couple of good references that can help improve diagnostic skills. For bareroot seedlings, readers are referred to
Forest Nursery Pests (Cordell and others 1989), and for container seedlings, to volume five of the Container Tree
Nursery Manual (Landis and others 1989).
Once a pest has been confirmed and the population has
exceeded the allowable limit, growers should take immediate action. All controls should be part of an integrated pest
management (IPM) program that uses cultural as well as
chemical controls. Many pest problems, such as Botrytis
blight, can be almost completely controlled using proper
irrigation and sanitation measures. Fungicides and insecticides are often needed however, especially when a problem
has gotten out of hand. Pesticides are usually injected
through the irrigation system in container nurseries or with
tractor-drawn sprayers in bareroot beds. In recent years,
some new biocontrol agents have proven useful in controlling insect pests, such as fungus gnats in greenhouses.
Weeds are a much more serious concern in bareroot
nurseries, where they must be controlled either mechanically
Table 3Chapter 7, Nursery Practices: careful observation of disease development can aid in diagnosis
Characteristics
Abiotic disease
Biotic disease
HOSTS
SYMPTOMS
Patterns
Rate of development
Signs
Spread
143
Beneficial Microorganisms
Mycorrhizae develop from a symbiotic relationship
between a beneficial fungus and the roots of the host
seedling. Although mycorrhizae have been a popular topic
for many years, there seems to be a variety of opinions as to
their value in forest and conservation nurseries. Some people believe that mycorrhizae are essential for both nursery
culture and successful outplanting, whereas other nursery
and reforestation specialists are more skeptical. Almost
50,000 research studies have been done on mycorrhizae, and
most confirm the benefits of reducing root disease and
increasing seedling tolerance to drought and other environmental extremes. Results of operational nursery and field
trials have been more variable, however, depending on the
species of mycorrhizal fungus used and soil fertility and
types of indigenous fungi on the outplanting site.
Inoculation with mycorrhizal fungi can be worthwhile,
but the timing of inoculation and species of fungus should
be matched to nursery and outplanting objectives (table 4).
In particular, the fungal species should be selected for either
the nursery or the outplanting site. There is no all-purpose
fungus that will perform well under all conditions. Most
fungal species that are adapted to wildland conditions will
not survive under the high moisture and high nutrient nurs-
Summary
Anyone considering propagating forestry and conservation species must be familiar with the unique characteristics
of these plants. Unlike most other crops, seedlings from forest nurseries are typically outplanted on relatively harsh sites
without subsequent care. This difference is significant
because seedling quality is defined by environmental conditions on the outplanting site. There is no such thing as an
all-purpose tree seedling. Bareroot and container seedlings
have different applications, and the choice of approach
depends on the available resources, the nursery climate, and
the conditions on the outplanting site.
Table 4Chapter 7, Nursery Practices: nurseries and seedling customers must consider their reasons for inoculating
with mycorrhizal fungi because their objectives determine the species of fungus and the type and timing of inoculation
Type of mycorrhizal
inoculation
Before sowing
Before sowing
Establishment or
rapid growth phases
Hardening phase
During packing or
before outplanting
* Regardless of the biological objectives, mycorrhizal inoculation may have marketing advantages.
144
Objectives of inoculation *
Nursery
Outplanting
1) Increased growth
2) Disease prevention
1) Increased growth
2) Disease prevention
1) Increased growth
2) Disease prevention
1) Disease prevention
2) Increased growth
None
None
None
None
1) Increased survival
1) Increased survival
2) Increased growth
Forestry and conservations plants are typically propagated by seed, although vegetative propagation is used for some
species. Direct seeding is by far the most common, although
sowing germinants or transplanting is used for species that
have complex germination requirements or other operational
restrictions. Although specific practices differ for container
and bareroot seedlings, a successful cultural regime must be
designed to reflect the biological requirements of the species
and the available resources of the nursery.
Seedlings must be properly handled and stored from the
time they are harvested until they are outplanted. Exposure
of the root system is particularly damaging to seedling quality. Because warm temperatures rapidly bring seedlings out
of dormancy, refrigerated storage is recommended whenever
possible. The time between removing the seedlings from the
storage area and outplanting them is one of the most critical
in the entire nursery and reforestation process. Seedlings are
susceptible to many abuses, with desiccation from exposure
to direct sun and overly warm temperatures the most serious. Seedling users need to understand that seedling quality
only decreases after the seedlings leave the nursery and all
mistakes and abuses are cumulative.
References
Cordell CE, Anderson RL, Hoffard WH, Landis TD, Smith RS Jr,Toko HV, tech.
coords. 1989. Forest Nursery Pests. Agric. Handbk. 680. Washington,
DC: USDA Forest Service. 184 p.
Duryea ML, Landis TD, eds. 1984. Forest nursery manual: production of
bareroot seedlings. Boston: Kluwer Academic Publishers. 385 p.
Landis TD, Simonich EJ. 1984. Producing native plants as container
seedlings. In: Murphy PM, comp.The challenge of producing native plants
for the Intermountain Area. Proceedings, Intermountain Nurserymens
Association Conference; 1983 August 811; Las Vegas, NV. Gen.Tech.
Rep. INT-168. Ogden, UT: USDA Forest Service, Intermountain Forest
and Range Experiment Station. 96 p.
Landis TD,Tinus RW, McDonald SE, Barnett JP. 1989. The container tree
nursery manual.Volume 5,The biological component: nursery pests and
mycorrhizae. Agric. Handbk. 674. Washington, DC: USDA Forest Service.
171 p.
Landis TD,Tinus RW, McDonald SE, Barnett JP. 1990. The container tree
nursery manual.Volume 2, Containers and growing media. Agric. Handbk.
674. Washington, DC: USDA Forest Service. 87 p.
Landis TD,Tinus RW, McDonald SE, Barnett JP. 1994. The container tree
nursery manual.Volume 1, Nursery planning, development, and management. Agric. Handbk. 674. Washington, DC: USDA Forest Service. 188 p.
Landis TD,Tinus RW, McDonald SE, Barnett JP. 1999. The container tree
nursery manual.Volume 6, Seedling propagation. Agric. Handbk. 674.
Washington, DC: USDA Forest Service. 167 p.
Lantz CW, tech. coord. 1989. A guide to the care and planting of southern
pine seedlings. Mgmt. Bull. R8-M39. Atlanta: USDA Forest Service,
Southern Region. 44 p.
Lantz CW, ed. 1985. Southern pine nursery handbook. Atlanta: USDA
Forest Service, Southern Region. Unpaginated.
Lowman BJ, Landis TD, Zensen F, Holland B. 1992. Bareroot nursery equipment catalog. Prog. Rep. 9224-2839-MTDC. Missoula, MT: USDA Forest
Service, Missoula Technology and Development Center. 204 p.
South DB, Mexal JG. 1984. Growing the best seedling for reforestation
success. Auburn: Alabama Agricultural Experiment Station, Auburn
University, Forestry Department. 11 p.
Sutherland JR,Van Eerden E. 1980. Diseases and insect pests in British
Columbia forest nurseries. Joint Rep. 12.Victoria: British Columbia
Ministry of Forests and Canadian Forestry Service, Pacific Forest
Research Centre. 55 p.
Sutherland JR, Lock W, Benson LE. 1982. Diseases and insects and their
management in container nurseries. In: Scarratt JB, Glerum C, Plexman
CA, eds. Proceedings, Canadian Containerized Tree Seedling Symposium,
1981 September 1416,Toronto. Sault Ste. Marie, ON: Canadian
Forestry Service, Great Lakes Forest Research Centre: 215223.
Thompson BE. 1984. Establishing a vigorous nursery crop: bed preparation,
seed sowing, and early seedling growth. In: Duryea ML, Landis TD, eds.
Forest nursery manual: production of bareroot seedlings. Boston: Kluwer
Academic Publishers: 41 49.
Williams RD, Hanks SH. 1994. Hardwood nursery guide. Agric. Handbook
473. Washington, DC: USDA Forest Service. 78p
145
146
Part II
Specific Handling Methods and
Data for 236 Genera
PinaceaePine family
Abies P. Mill.
fir
D. George W. Edwards
Dr. Edwards retired from the Canadian Forest Services Pacific Forestry Centre
Growth habit, occurrence, and use. The name Abies
is derived from abed, the Old World Latin name for the
silver fir (Dallimore and Jackson 1967; Weber 1987).
Theophrastus (371286 BC) wrote of silver firs from Mt.
Ida (todays Kaz Dag, Turkey) being used in shipbuilding,
which may have been the lumber of A. equi-trojani (Thanos
2003b), but also may have been in reference to A. cephalonica Loud. and/or A. pectinata DC. (now A. alba P. Mill.)
(Amigues 1993, cited in Thanos (2003b). The name Abies
first appeared in Pliny the elders Historiae Naturalis from
about AD 77 (Liu 1971).
Firs are long-lived, on average achieving reproductive
maturity at 20 years, with an average life-span of 60 years
(Jacobs and others 1984). Fir trees in excess of 400 years
old have been recorded in several species (Earle 1999), and
noble firs 600 to 700 years old are known (Arno and
Hammerly 1977; Franklin 1979; Franklin and Dyrness
1973), but such life spans are modest compared to those of
other tree genera. Siberian fir (table 1) rarely, if ever, survives more than 200 years because the main stem decays out
(Vidakovic 1991). In numbers of species, fir is second only
to pine but lags behind spruce (Picea spp.) and Douglas-fir
(Pseudotsuga menziesii (Mirb.) Franco) in terms of overall
importance (Franklin 1982a).
All fir species are indigenous to the Northern
Hemisphere (table 1), being widely distributed over the
Eastern and Western Hemispheres (Liu 1971) chiefly in the
temperate and frigid regions, from sea level to altitudes of
4,700 m. More than 70 species have been variously
described (Liu 1971), although the number of those currently recognized is between 39 (Liu 1971) or 40 (Vidakovic
1991), 46 (Farjon 1990), ~50 (Welch 1991), and 55
(Rushforth 1987), depending on placements into varietal
categories. Firs are found in 4 extensive regions (Franklin
1974b; Liu 1971; Miller and Knowles 1989; Welch 1991;
Young and Young 1992):
149
150
A. alba P. Mill.
A. argentea DC.; A. candicans Fisch.
A. nobilis A. Dietr.; A. Pardei Gauss
A. pectinata DC.; A. picea Lindl.
A. taxifolia Desfont.; A. vulgaris Poir.
A. amabilis (Dougl. ex Loud.) Dougl. ex
Forbes
A. grandis A. Murr.
A. grandis var. densiflora Engelm.
A. balsamea (L.) P. Mill.
A. aromatica Rafn.
A. balsamifera Mich.
A. minor Duham. ex Gord.
A. bracteata (D. Don) D. Don ex Poit.
A. venusta (Dougl.) K. Koch
A. cephalonica Loudon
A. panachaica Heildr.; A. lusombiana Loudon
A. peloponesica Haage
A. cilicica (Antoine & Kotschy) Carrire
A. selinusia Carrire
A. concolor var. concolor (Gord. & Glend.)
Lindl. ex Hildebr.
A. lowiana (Gord.) A. Murr.
A. grandis var. lowiana (Gord.) Hoopes
A. concolor var. lowiana (Gord.) Lemm.
A. concolor var. lowiana (Gord. & Glend.)
Lemmon
A. lowiana (Gord.) A. Murr.
A. concolor (Gord. & Glend.)
A. concolor var. lasiocarpa Engelm. & Sarg.
A. grandis var. lowiana Mast.
A. firma Sieb. & Zucc.
A. bifida Sieb. & Zucc.; A. momi Sieb.
A. fraseri (Pursh) Poir.
``humilis La Pilaye
A. grandis (Dougl. ex D. Don) Lindl.
A. amabilis A. Murr.; A. excelsior (Franco)
A. gordoniana Carr.
A. lasiocarpa Lindl. & Gord.
A. guatemalensis Rehd.
A. tacanensis Lund.
A. guatemalensis var. tacanensis (Lund.) Mart.
A. guatemalensis var. jaliscans Mart.
Mtns of Guatemala, S Mexico, El Salvador; Honduras
(19301450N & 10491W)
Occurrence
Common name(s)
Turkey fir
A. holophylla Maxim.
Occurrence
Common name(s)
Abies
151
152
A. veitchii Lindl.
A. eichleri Lauche; A. sikokiana Nakai
The following recognized fir species are not included in the table for lack of sufficient data (common names are given when known):
MEDITERRANEAN
BASIN:
A. x borisii-regis Mattf. (Bulgarian fir, sometimes Macedonian fir, or King Boris fir); A. marocana Trab. (Moroccan fir); A. tazaotana Cheval. (Tazaotan fir).
EAST ASIA: A. beshanzuensis Wu (Baishan fir); A. chengii Rushf. (Cheng fir); A. chensiensis Van Tiegh. (Shensi fir); A. delavayi (Van Tiegh.) Franch. (Yunnan fir, or Delavay fir); A. densa
Griff. (Sikkim fir); A. fabri (Mast.) Craib (Faber fir, sometimes also Yunnan fir); A. fanjingshanensis Huang,Tu et Fang (Fanjingshan fir); A. fargesii Franch. (Farges fir); A. forrestii Coltm.Rog. (Forrest fir); A. kawakamii (Hay.) Ito (Taiwan fir); A. spectabilis (D. Don) Spach (east Himalayan fir or Webb fir); A. yuanbaoshensis Lu et Fu (Yuanbaoshan fir); A. ziyuanensis Fu et
Mo (Ziyuan fir).
CENTRAL AMERICA: A. colimensis sp. nov. Rushf. & Nar.; A. durangensis Mart. (Durango fir); A. flinckii sp. nov. Rushf.; A. hickelii Flous et Gauss. (Hickel fir); A. hidalgensis sp. nov. Debr.,
Rcz & Guz.; A. neodurangensis sp. nov. Debr., Rcz & Salaz.; A. zapotekensis sp. nov. Debr., Rcz & Ramr.; A. vejarii Mart. (Vejar fir).
Note:
Sources: Anon. (1998), Dallimore and Jackson (1967), Donahue and others (1985), Earle (1999), Farjon and Rushforth (1989), Franklin (1974b), Liu (1971), Puri and Gupta (1968).
* Spelling of Chinese place names has changed over time.This name is given as Hopeh in Liu (1971) but is currently Hebei.
Formerly spelled Szechuan or Szuchuan (Liu 1971).
Occurrence
Common name(s)
Abies
153
9 speciesPacific silver, balsam, white, Fraser, grand, subalpine, red, Shasta red and noble firsare now in regular
use throughout their native ranges.
With 2 exceptionsFraser fir, the remaining stands of
which are extremely valuable for watershed protection as
well as for their scenic beauty (Beck 1990), and the rare
bristlecone firall North American firs have become commercially valuable as timber and/or pulp species. In general,
fir wood is soft, odorless, and light in color and weight; it
lacks resin ducts and usually kiln-dries without checking or
collapse (but tends to warp). It is easily worked and finished
to a good surface, and it takes paint and polish well
(Dallimore and Jackson 1967). Although generally of low
durability (Franklin 1982a) unless treated with preservative,
fir wood can be used in projects that do not require high
structural strength; balsam fir is used extensively for cabin
logs. Noble fir wood (sometimes marketed as Oregon
larch) is the strongest (along with red fir) of fir woods and
is more durable than that of most firs. The frames of Royal
Air Force Mosquito fighter planes of World War II were
build with noble fir (Pojar and MacKinnon 1994). Grand fir
knots, steamed and carved, were made into fish hooks
(Turner 1998). The many other products made in North
America of fir wood include quality veneers, paneling, construction plywood, crates, container veneers, poles (after
preservative treatment), moldings, window sash and door
stock, Venetian blinds, ladder rails, and aircraft framing
(because of its high strength-to-weight ratio) (Bakuzis and
Hansen 1965; Frank 1990; Franklin 1974b, 1982a, 1990;
Smith 1982). In the late 19th century, clear lumber of red fir
was known as butter wood because, when made into
boxes for cheese and butter, it did not influence their flavor
(Young and Young 1992).
Japan, which imports large quantities of noble and
Pacific silver firs for construction (Franklin 1982a), uses its
indigenous Japanese fir for making boards, roof shingles,
door plates, matches, wooden clogs, musical instruments,
household utensils (furniture, packing boxes, and coffins), as
well as using it in ship-building and cooperage (Liu 1971).
The Yunnan and Faber firs (A. delavayi and A. fabri, see
table 1 footnotes) are used for temple construction in the
high mountains of Sichuan Province, China (Earle 1999).
European silver fir is widely used throughout Europe also
for construction, joinery, musical instruments, and (after preservative treatment) for poles. Guatemalan fir faces extinction in parts of its range (Donahue and others 1985; Salazar
1991) through overuse for building materials, roof shingles,
interior paneling, weaving looms and low-density furniture, shipping crates, charcoal, firewood (Anon. 1998;
154
divided into a total of 9 subsections, including 3 new subsectional names (Farjon and Rushforth 1989); an historical
review plus an evaluation of other attempts to classify firs
are included. The new scheme is diagrammatically represented in table 2.
For North American firs, section Bracteata retains the
single species A. bracteata as the type species, whereas section Amabilis includes A. amabilis as the type species.
Section Balsameae, subsection Laterales (type A.
kawakamii), includes A. balsamea, A. bifolia, and A. lasiocarpa, whereas subsection Medianae (type A. sachalinensis)
includes A. fraseri. Section Grandes includes A. grandis
(type) and A. concolor, as well as the Central American
species A. guatemalensis, A. durangensis, and a new species
A. flinckii (Rushforth 1989). Section Nobiles includes A.
procera (type) and A. magnifica. Section Oiamel, which is
divided into subsections Religiosae and Hickelianae,
includes the other known Central American firs, including
another new species A. colimensis (Rushforth 1989).
Note that this scheme places Fraser fir (Abies fraseri) in
subsection Medianae and balsam fir (A. balsamea) in subsection Laterales; this separation is based on whether bract
scales are exserted and the seed scales reniform (Medianae,
Fraser fir), or bract scales are included and seed scales are
cuneate-flabellate (Laterales, balsam fir) (Farjon and
Rushforth 1989). Natural hybrids between these 2 species
have been reported (see below) and bracts in balsam fir are
not always completely included (hidden) (Lester 1968), so
this separation does not appear to be justified.
Detailed taxonomy (as well as descriptions of cones,
pollen, seeds, and seedlings) of 11 European fir species can
be found in a recent monograph (Schutt 1991), whereas a
more general text (Vidakovic 1991) includes 26 fir species.
Other descriptions and drawings are available (Cope 1993;
Rehder 1958; Rushforth 1983, 1984, 1986; Farjon 1990;
Debreczy and Rcz 1995).
In North America, 2 sets of genetic complexesgrand
and white firs, and noble and California red firscreate significant taxonomic confusion for students, foresters, and
land-managers (Franklin 1982a). The geographic variation
of the first setgrand fir and white fir (section Grandes,
Farjon and Rushworth 1989; section Grandes, Liu 1971)
has been extensively studied. Although these 2 species are
morphologically, ecologically, and chemically distinct, they
are genetically plastic and intergrade and hybridize freely
over a wide area (Daniels 1969; Foiles and others 1990;
Hamrick 1966, cited by Franklin 1974b; Hamrick and Libby
1972; Klaehn and Winieski 1962; Laacke 1990a; Lacaze
1967; Steinhoff 1978). The variation can be continuous
Abies
155
156
Abies P. Mill.
Nobiles Engelmann
A. procera (type)
A. magnifica (includes var. shastensis)
Species
Oiamel Franco
(type: Abies religiosa)
Squamatae E. Murray
Laterales Patschke emended Farjon & Rushforth
Subsection
Section
Geographical location
Coastal lowlands of southern British
Columbia,Washington, Oregon, and
California, including lower elevations on
the western slopes of the Cascade Range
A. grandis
Eastern slopes and higher elevations in the
Cascade Range north of about 44 to
45N latitude
A. grandis
Northern Idaho and interior of southern
British Columbia
Intergrade
Klamath Mountains and Cascade Range of
southwestern Oregon and northern
California
Intergrade
Blue, Ochoco, and Wallowa Mountains of
northeastern Oregon, west central Idaho
A. concolor*
Sierra Nevada, California
A. concolor
Southern Rocky Mountains and southern
California
*Now recognized as Sierra white fir (table 1).
Noble, California red, and Shasta red firs form the second important interfertile complex of species (Franklin and
others 1978; Sorensen and others 1990). Noble and
California red firs readily produce hybrids (Barbour 1988;
Little 1979) with seed and seedling characteristics similar to
Shasta red fir where the ranges overlap (Franklin and others
1978; Sawyer and Thornburgh 1977; Silen and others 1965;
Sorensen and others 1990). Populations in southern Oregon
and northwestern California may represent hybrid swarms
between the 2 species (Franklin and others 1978).
Phenotypically, trees in southern Oregon to northwestern
California often resemble noble fir but behave ecologically
as Shasta red fir (Lfting 1966 and 1967, cited by Franklin
1974b). A latitudinal gradient in the Cascade Range, with a
major discontinuity around 44N, has been discerned
(Franklin and Greathouse 1968a). The 2 species can be artificially cross-pollinated without difficulty as long as red fir
is the female (ovuliferous) parent (Zavarin and others 1978).
Noble fir exhibits high self-fertility that does not appear to
affect germination but which can depress height growth
(Sorensen and others 1976). Although no races of noble fir
are known within its natural range, population differentiation and variation is reported (Maze and Parker 1983). Three
horticultural varietiescv. glauca, cv. prostrata, and cv.
robustifoliaare recognized (Franklin 1990). When noble,
Sakhalin, Maries, Japanese, and Grecian firs were used as
female parents, height, dbh, and crown area were greater in
the interspecific crosses than in intraspecific crosses
(Mergen and Gregoire 1988).
Of all the interspecific crosses, progeny of Maries fir (as
the female parent) showed the greatest growth; this species
also had the least, whereas Sakhalin fir had the greatest,
inbreeding depression (Mergen and Gregoire 1988). Effects
of these crosses on seed and seedling characteristics were
reported earlier (Mergen and others 1965). Geographic similarity (especially among Japanese, Korean, Maries, and
Sakhalin firs) was suggested as a positive influence on
hybrid survival and performance (Mergen and Gregoire
1988). Earlier, it had been suggested that a geographical,
rather than genetic or physiological, separation occurred as
the genus Abies evolved (Klaehn and Winieski 1962).
Possible causes for incompatibility and results from other
European inter- and intraspecific crossing experiments are
reported (Kantor and Chira, 1965, 1971, 1972). However,
many of the reported artificial crosses between noble fir and
other true firs including balsam, white, subalpine, Min (or
Min-kiang), and Sakhalin firs have not been repeated, and
Abies
157
158
Abies
159
160
Abies
161
Late Aprmid-May
June
Juneearly July
MayJune
Midlate June
June
Aug
Midlate Sept
May
After Oct 1
Septearly Oct
Midlate Sept
Late Sept
SeptOct
Late Sept
Late Sept
Septearly Oct
Mid-Septmid-Oct
Late Aug
Mid-Sept
Late Aug
SeptOct
Midlate Oct
Septmid-Oct
Aug
Octmid-Dec
Midlate Sept
Mid-Septearly Oct
Mid-Aug
Late Aug
Fruit ripening
SeptOct
Late Septearly Oct
SeptOct
OctNov
Early Oct
Early Oct
Oct
Late Septmid-Oct
Earlymid-Oct
Mid-Oct
SeptOct
Mid-Sept
Mid-Sept
Early Oct
Mid-Sept
SeptOct
Late Septlate Oct
Novmid-Dec
Midlate Sept
Late Septearly Oct
Mid-Sept
Early Sept
Early Oct
Mid-Septmid-Oct
Late AugSept
Seed dispersal
3055
1025
4060
4580
3055
2025
1020
1035
2560
3045
1025
3565
2030
2030
1035
1035
2545
3565
Tree
ht (m)
3545
3040
3040
50
1215
3040
30
2030
40
15
20
50
20
2530
30
Age
(yrs)*
23
57
23
24
36
24
23
28
56
24
35
39
46
3
23
23
57
23
24
23
36
Interval
(yrs)
Sources: Ahlgren (1957), Anon. (1950b, 1998), Bakuzis and Hansen (1965), Baron (1969), Beck (1990), Dallimore and Jackson (1967), Ebell and Schmidt (1964), Eis (1970), Eis and others (1965), Enescu (1960), Fowells (1965), Fowells
and Schubert (1956), Franklin (1968, 1974b), Franklin and Ritchie (1970), Gordon (1978), Haig and others (1941), Hetherington (1965), Hughes (1967), Laacke (1990a&b), Laacke and Fiske (1983), Legg (1953), Leloup (1956), Little and
DeLisle (1962), Lfting (1961), MacDonald and others (1957), MacLean (1960), Morris (1951), Munz and Keck (1959), Owens and Molder (1977b), Pearson (1931), Puri and Gupta (1968), Rudolf (1952), Sato (1940), Schmidt and Lotan
(1980), Singh and Singh (1984a&b),Talley (1974),Tulstrup (1952), USDA Forest Service (1948),Wappes (1932), Zon (1914).
* Minimum age for commercial seed bearing.
At higher elevations in central Europe, 4 to 6 years.
Occasionally (Talley 1974).
Includes A. delavayi, A. densa. and A. spectabilis (Puri and Gupta 1968).
A. sibirica
A. veitchii
A sachalinensis
A. shastensis
A. pindrow
A. pinsapo
A. procera
A. mariesii
A. nordmanniana
A. magnifica
A. guatemalensis
A. homolepis
A. lasiocarpa
A. firma
A. fraseri
A. grandis
A. bracteata
A. concolor
var. lowiana
Japan
Russian Georgia
Himalayas
Czech Republic & Soslovakia
Benton & Linn Cos., Oregon (1,3501,550)
Lewis Co.,Washington (1,600)
Hokkaido, Japan
SW Oregon (1,8502,000)
N. California coast ranges (2,000)
Shasta Co., California (1,7002,000)
W Siberia
Japan
Maymid-June
Late AprMay
Mid-MayJune
June
Midlate-May
Late Aprearly June
Late Aprearly May
MayJune
Late May
Mid-Mayearly June
Late Aprmid-May
Mid-Mayearly June
Mid-June
Mid-Aprilmid-May
Earlymid-June
Late Marchearly Apr
Mid-Maymid-June
Late June
Late Juneearly July
Earlymid-July
Late Mayearly July
Europe
W Washington & Oregon (150400)
Vancouver Is., British Columbia (500)
Lewis Co.,Washington (1,600)
Minnesota
Sta. Lucia Mtns, Monterey Co., California
A. alba
A. amabilis
A. balsamea
Flowering
Species
Table 3Abies, fir: phenology of flowering and fruiting, and major characteristics of mature trees
162
Figure 2Abies procera, noble showing the typical raspberry form (courtesy of Y. Tanaka.) fir: male strobili prior
to pollen shedding (courtesy of D. Pigott).
this twisting actively tears them from the axis. No such distortion occurs in noble fir, and seed dissemination requires
branch movement by the wind or other agents to disturb the
cone (Franklin and Ritchie 1970). Cone disintegration of
other species such as grand fir and subalpine fir are intermediate.
Thus, pollination, fertilization, seed ripening, and dissemination all occur in the same seasonin as little as 90 to
120 daysfollowing the year of strobilus initiation
(Franklin and Ritchie 1970). The chief agent of seed dispersal is the wind; seed rain density decreases as a function of
distance from the parent tree, seedling mortality increases,
and smaller-seeded species travel further (Carkin and others
1978; Franklin 1982b; Hofmann 1911; Houle 1992, 1995;
Isaac 1930b; McDonald 1980; Savcenko 1966;
Wolfenbarger 1946).
The majority of fir seeds are normally shed in
October/November (table 3). Frequently these have the highest seed weight, maximum germination capacity and lowest
occurrence of empty and immature seeds, plus higher
seedling survival rates, than seeds shed earlier/later. In several firs, seed dispersal may extend well into winter (Anon.
1950b; Aussenac 1966; Hetherington 1965; Houle and
Payette 1991; Roe 1946), the seeds becoming buried in, and
germinating in, snowbanks (see also Pregermination treatments). Up to 50% of a Maries fir seedcrop may lodge in
the foliage and only fall to the ground over winter (Smirnov
1991). The date of seed-fall of European silver fir in Italy
Abies
163
The role of resin has been linked to inhibiting precocious germination, that is, to promoting dormancy, of mature
fir seeds at the time of seedfall (Rohmeder 1951). It might
also provide some form of protection for the embryo and
megagametophyte against excessive drying (Gunia and
Simak 1970). Germination of non-stratified European silver
fir seeds was increased after resin removal by low-temperature vacuum distillation (Zentsch 1960), and resin extracted
from this species inhibited germination in pine and spruce
seeds (Dssler and Zentsch 1959; Rohmeder 1951).
Damaging the vesicles during processing of fresh European
silver fir seeds and allowing the resin to contaminate
undamaged seeds reduced their germination (Gunia and
Simak 1970). The germination-reducing effect of resin leak-
164
Figure 10Abies grandis, grand fir: longitudinally sectioned mature seeds showing embryos (e) occupying 90+%
of the corrosion cavity in the megagmetophytes (meg)
(endosperm. C = cotyledons; AP = apical meristem; R =
radicle; H= hypocotyl/shoot axis: V = resin vesicle; SC =
seedcoat. Scale bar is in millimeters (courtesy of D. Pigott).
ration (Eis 1973). Thus, large crops are unlikely in consecutive years. Other environmental requirements must be met
also, which is why lapses of several years between heavy
crops is more the rule. For example, the interval between
heavy crops of white fir in California is commonly 5 years
(McDonald 1992) but may vary from 3 to 9 years (Fowells
and Schubert 1956). Henderson (1982) found that for subalpine fir, only 1 year over a 28-year period produced a
bumper crop, whereas 4 other years were good. Several
true firs in Oregon and Washington produce good crops on a
3-year cycle (Franklin 1968), with noble fir averaging medium or better crops 50% of the time over its range, although
some sites may go as long as 6 years without significant
cone production (Franklin 1982b). Crop year can have a
large effect on seed weight and cotyledon number in noble
fir (Sorensen and Franklin 1977); cotyledon number in
Sakhalin fir was weakly correlated with provenance (Okada
1966).
Several methods for forecasting cone crops have been
devised. One, for Maries fir, is based on bud counts the previous year (Matsuura 1963); another uses visual estimates of
the number of cones on individual Sierra white fir trees and
the proportion of trees bearing cones (McDonald 1992); a
photographic method is more accurate than visual rating for
red fir (Gordon 1962). Crop production in grand fir can be
estimated using a regression equation that employs the number of cones on the top 2 whorls of the crown, and the num-
Abies
165
166
Abies
167
Insect*
Common name
bract feeder
fir cone moth
cone moth
Earomyia spp.
seed maggot
cone maggot
fir cone maggot
seed chalcid
Pegohylemia spp.
Ptilinus fur L.
Resseliella spp.
Spermatolonchaea viridana L.
Zeiraphera rufimitrana Foote
A. balsamea, fraseri
A. alba
A. concolor, lasiocarpa
A. alba, concolor, grandis, lasiocarpa, magnifica, nephrolepis
A. alba
A. alba
A. alba
A. concolor
A. concolor, grandis, lasiocarpa, procera
A. guatemalensis
A. alba, amabilis, balsamea, A. cephalonica, A. concolor,
A. grandis, nephrolepis, nordmanniana,
A. pindrow, pinsapo
A. alba, concolor, grandis, lasiocarpa, magnifica,
A. nordmanniana, procera
A. alba
A. concolor
A. alba, borisii-regis, cephalonica, sibirica
A. bracteata, concolor, grandis, lasiocarpa, nephrolepis
A. concolor, grandis, lasiocarpa, nephrolepis
A. alba, borisii-regis, cephalonica, concolor, magnifica
A. grandis
A. alba
A. alba, borisii-regis, cephalonica
A. alba
A. alba, amabilis, balsamea, borisii-regis,
A. bracteata, cephalonica, concolor, fraseri, grandis, guatemalensis,
A. lasiocarpa, magnifica, bornmuelleriana var. equitrojana
A. sibirica pinsapo, procera,
A. alba, balsamea
A. alba
A. alba, borisii-regis, cephalonica, cilicica, concolor,
A. grandis, nordmanniana
A. cilicica
A. alba
Sources: Androic (1960, 1976), Annila (1982), Arista and Talavera (1995), Bess (1946), Blais (1952), Bradley and others (1981), Bryant and Hudak (1968), Canakcioglu
(1969), Donahue and others (1985), Durzan (1979), Eremko and others (1989), Fang and others (1988, 1989), Fedde (1973a&b), Gagov (1976), Gonzalez and others (1983)
[in Donahue and others 1985]), Gordon (1970), Greenbank (1963), Hall (1981), Hedlin (1966), Hedlin and Ruth (1974), Hedlin and others (1980), Hussey (1954, 1957,
1960), Hussey and Klinger (1954), Jesperson and Lomholdt (1983), Kailidis and Georgevits (1970, 1972), Kayacik (1964), Keen (1968), Koerber (1963), Kolomiec (1950),
Kulhavy and Schenk (1976), Kulhavy and others (1976), Lanz (1942, 1943), Legg (1953), Mackay (1949), Matic (1972), Miller (1986), Miller and Ruth (1989), Moody (1988),
Nanu (1979b), Nanu and others (1986), Nekrasova (1978b), OConnor and OConnor (1984), Overhulser and Tanaka (1983), Pfister and Woolwine (1963), Powell (1973),
Pribylova (1975), Puri and Gupta (1968), Rahman and Chaudhry (1986), Schooley (1975, 1976, 1978), Scurlock and others (1982), Shea (1989a&b), (Skrzypczynska 1982,
1984, 1985, 1989a&b), Skrzypczynska and others (1988, 1990, 1995), Speers (1968, 1969),Talley (1974),Tanaka (1982),Toth (1973),Woodwell (1961).
* Insect names, in alphabetical order, are listed as cited by sources. No attempt has been made to rationalize synonyms, because sources rely on different nomenclature
authorities.
For simplicity and conciseness, where several species in a single genus have been identified, insects are grouped by genus, for example, Asynapta spp.
168
fungi and other organisms isolated from fir cones and seeds
Organism
Alternaria spp.
Aspergillus spp.
Botrytis cinerea Pers.: Fr.
Caloscypha fulgens (Pers.) Boud.
Cephalosporium spp.
Ciboria rufo-fusca (O.Weberb.) Sacc.
Cladosporium spp.
Cylindrocarpon spp.
Fusarium culmorum (Wm.G. Sm.) Sacc.
Fusarium moniliforme J. Sheld.
Fusarium oxysporum Schlechtend.: Fr.
Fusarium roseum Link: Fr.
Fusarium semitectum Berk.& Ravenel var. majus (Wollenweb.)
Fusarium spp.
Geniculodendron pyriforme G.A. Salt
Heterobasidion annosum (Fr.:Fr.) Bref.
Lirula macrospora (R. Hartig) Darker
Melanospora zamiae Corda
Mucor spp.
Papulospora spp.
Penicillium spp.
Phoma spp.
Rhacodium therryanum Theum.
Rhizoctonia solani Khn
Sclerotium spp.
Trichoderma spp.
Tricothecium roseum (Pers.:Fr.) Link
Truncatella hartigii (Tub.) Steyaert
Virus-like particles
Abies spp.*
Abies spp.
A. amabilis
A. grandis
Abies spp.
A. alba, nordmanniana
A. grandis, magnifica, shastensis
A. amabilis, sibirica, Abies spp.
Abies spp.
A. grandis, nordmanniana
A. grandis, procera
A. grandis, procera
A. amabilis
Abies spp.
A. amabilis, grandis
Abies spp.
Abies spp.
Abies spp.
Abies spp.
A. amabilis, grandis
A. amabilis, grandis, magnifica, shastensis, procera
Abies spp.
A. sachalinensis
A. balsamea, fraseri, grandis
A. mariesii, Abies spp.
A. amabilis, grandis, Abies spp.
A. grandis
Abies spp.
A. alba, homolepis
Sources: Anderson (1985), Bloomberg (1969), Buchwald and others (1961), Edwards and Sutherland (1979), Eremko and others (1989), Flachmann and others (1990),
Hayashi and Endo (1975), Heit and Natti (1969), Kolotelo (1994), Littke and Browning (1991), Ono (1974), Prisyazhnyuk (1960). Fungal nomenclature mainly according to
Farr and others (1989).
* Individual tree species not determined.
Abies
169
170
Species
Cones
Seeds
A. amabilis
A. balsamea
A. concolor
A. firma
A. fraseri
A. grandis
A. homolepis
A. lasiocarpa
A. magnifica
A. mariesii
A. procera
A. sachalinensis
A. veitchii
A. guatemalensis
Sources: Anon. (1998), Bakuzis and Hansen (1965), Donahue and others (1985), Eremko and others (1989), Franklin (1965, 1974b), Oliver (1974), Pfister (1967), Snyder
(1976), Speers (1962), Stoeckeler and Jones (1957).
* Using 10 lens.
Rediske and Nicholson (1965).
Abies
171
172
of the application of the technique, as well as aerial clipping/sawing, and aerial topping for cone collection, have
been comprehensively reviewed (Camenzind 1990).
Cone and seed processing. Seed germinability of a
number of species, including white (Oliver 1974), grand
(Pfister 1966), Nordmann (Muller 1971), and noble firs
(Edwards 1969; Franklin 1965; Rediske and Nicholson
1965) can be improved by storing the cones under cool,
moist conditions for several weeks after collection. In contrast, cones of red fir need to be collected as close as possible to seed fall (Oliver 1974). Artificial ripening of earlycollected seeds allows cone collections to be started sooner,
thus extending the collection period, so that immature cones
from logging operations can be used (Edwards 1982a). The
maximum period of collection prior to the onset of natural
seed dispersal appears to be around 6 weeks, but it is safer
Figure 15Abies grandis, grand fir: bagging aerially collected cones at a dump site; note the wooden box by pickers knee, this is a cone-volume measuring device (courtesy
of D. Pigott).
Abies
173
174
Figure 16Abies lasiocarpa, subalpine fir: viviparous germination, with seeds germinating in the cone before they
could be extracted (courtesy of D. Pigott).
Figure 17Abies lasiocarpa, subalpine fir: viviparous germination, with seeds germinating while still attached to the
ovuliferous scales (courtesy of D. Pigott).
Species
A. amabilis
A. balsamea
A. concolor
A. firma
A. fraseri
A. grandis
A. guatemalensis
A. homolepis
A. lasiocarpa
A. magnifica
A. mariesii
A. procera
A. sachalinensis
A
Kiln-drying period
Time (hr)
614
0
0
()
0
614
0
()
614
0
()
614
()
Temp (C)
3038
48
3038
48
3038
48
3038
48
Sources: Anon. (1998), Edwards (1982a), Franklin (1974b), Heit (1968a), Heit and Eliason (1940), Jones (1962), Leloup (1956), Speers (1967).
* At ambient air temperature; cooled (<10C) conditioning facilities are superior.
If air-drying not possible, but cones should not be processed immediately after harvest.
In a rotary kiln; seeds removed from heat as soon as they fall through the tumbler mesh.
In the shade.
Abies
175
176
28
35 2,7005,500 1,0002,000
3035
2,5002,700
9001,000
700
250
4250
800
300
2530
2632
850
312
3139
200
80
/bu
55
32
40
125
89
51
64
196
Seed wt/
cone wt
g/kg oz/100lb
2
3.7
2.93.6
1.32.5
2.53.7
1.92.5
1.7
1.21.9
1.4
4.85.8
1.73.6
25
48
400
3746
134
1732
185
3248
2432
115
6789
22
1624
18
5265
6275
2246
500
Seed wt/
cone vol Seeds/
kg/hl oz/bu cone
17,40041,000
17,20036,400
21,80045,900
66,150208,400
18,95039,100
20,50030.900
117,950173,650
26,25063,500
30,00043,000
32,20049,000
52,700108,700
38,80056,200
8,80019,600
42,10065,050
11,55019,000
20,30042,100
65,050118,000
11,25024,700
50,700173,750
/kg
/lb
/kg
/lb
10,200
11,000
13,800 *
59,600
11,100
11,400
60,800
18,400
20,200*
16,500
19,800
34,800
21,600*
22,300
6,400
23,000
7,100
13,500*
44,000
7,300*
45,000
Average
22,500
24,250
30,450*
131,400
24,500
25,150
134,050
40,600
44,550*
36,500
43,650
76,750
47,600*
49,200
14,100
50,700
15,650
29,800*
97,000
16,100*
99,200
Seeds/wt
7,90018,600
7,80016,500
9,90020,800
30,00094,500
8,60017,720
9,30014,000
53,50078,750
11,90028,800
13,60019,500
14,60022,200
23,90049,300
17,60025,500
4,0008,900
19,10029,500
5,7008,600
9,20019,100
29,50053,5000
5,10011,200
23,00078,800
Range
>72
66
8
42
46
>12
10
144
12
>2
19
19
4
8
36
>6
>24
>36
>29
36
17
Samples
Sources: Anon. (1998), Ching (1960), den Ouden and Boom (1965), Eis and others (1965), Fowells and Schubert (1956), Franklin (1974b), Ghent (1958), Heit (1968a), Lalu (1993), Lanquist (1946), Leloup (1956), MacDonald and others
(1957), Rafn (1915), Rafn and Son (nd), Roe (1948b), Seal and others (1965), Soljanik (1950), Speers (1962),Tulstrup (1952),Wappes (1932).
* Seeds were 100% sound, separated by x-radiography.
A. alba
A. amabilis
No. of cones
/hl
Cone wt/vol
kg/hl
lb/bu
36
A. balsamea
45
A. concolor
3945
A. firma
A. fraseri
A. grandis
A. guatemalensis
A. homolepis
5464
A. lasiocarpa
var. arizonica
A. magnifica
3239
A. mariesii
3341
A. nordmanniana 4050
A. procera
A. sachalinensis
A. x shastensis
A. veitchii
Species
Abies
177
experiences with seed storage conditions (recommended conditions are in bold face)
Moisture content
(% fresh wt)
A. alba
A. amabilis
A. balsamea
A. cephalonica
A. concolor
A. firma
A. fraseri
A. grandis
A. guatemalensis
A. homolepis
A. lasiocarpa
A. magnifica
A. mariesii
A. nordmanniana
A. procera
A. sachalinensis
A. shastensis
Storage
temp (C)
57
58
<9
68
58
68
911
58
610
1015
58
710
11
911
<9
6-8
58
911
911
<9
69
69
11
3 to 7
10 to 17
15
17
+0.5 to +4
17
+4
0 to 18
18
2 to 4
12
7
4 to 10
4
+4
15
+3 to +4
2 to +4
17
+5
2 to +4
+4
15
0 to 18
4
2 to +4
4
Possible storage
period (yr)
26
15
45
>5
5
13
12
7
3
>6
>2
3
10 +
12
>5
<1
>6
>5
5
>6
2
>5
7
> 10
>6
> 10
Sources: Allen (1957), Edwards (1982a), Franklin (1974b), Gradi (1966), Heit (1941, 1968b), Hofman and Vackova (1966), Holmes and Buszewicz (1962), Issleib (1956),
Jones (1962), Loffler (1985), Machanicek (1965), Mormann (1956), Radulescu (1968), Speers (1974b),Tillisch (1952),Tokarz (1974).
178
A. koreana
A. lasiocarpa
var. arizonica
A. homolepis
A. grandis
A. firmai
A. fraseri
A. concolor
A. bracteata
A. balsamea
A. amabilis
A. alba
Species
2860f
14d80h
14d80h
3060
6080
6080
0
30d80h
28d120h
0
0j42f,g
28d120h
28d120h
28d120h
4060
28d60h
28d60h
28d60h
Spring
Early Jan (1+0) for fall/winter lift
Febmid-Apr
30d120h
30d120h
0
2860h
0
1428
0
3 060
0
2860
28d60h
30d120h
30d120h
Sowing season
Fall
Mid-Marmid-Apr
MarApr
0
30d80
28f
Stratification
time (days)
220540
270
330
270430
215270
220540
22054
270430
270430
270540
330/row
2050
25
100
2540
2025
2050
2050
2540
0.3
0.30.5
2c
0.52
0.5
2c
2c
0.51
Abies .
1/
3/ c
4
1/ 3/
4 4
1/ 1/
8 4
3/ c
4
1/
1/ 1/
4 2
3/ c
4
Leaf mold
Sawdust
None, sawdust
pine needles
Pine needles,
peat moss, none
Strawg, none
None
Pine needles
Strawg
Bareroot production
Sowing depth
cm
in
Mulcha
2540
2550
100/row
Seedling density
/m2
/ft2
2+2
2+2
3+0, 4+0
2+0
3+0, 3+1
1+0, 2+0, 3+0
3+0, 3+1
Stock
type
Styro 2,5
Styro 2,5
Styro 2,5
313B, 410A
313A, 410A, 415B,D, 412A,
615A, Styro 2,5,7; Leach 1,2
415B, 412A
313B, 410A
415B, D, 615D
313A, 415B,D, 615D, Styro 2, 5, 7
415B, 412A
313B, 410A
313A, 410A, 415B,D, 615A
313A, 410A, 415B,D,
412A,615A,Leach 1,2,Styro 2,5,7
412A
Styro 2, 5
313B, 410A
313A&B, 410A, PCT410,
412A, 415B&D, 615A
313A, Styro 2, 5,& 7
415B,D, 412A
Styro 2, 5, 7
1+0
1+0
1+0, 2+0
1+0
1+0, 2+0, 2+1,
P+1
2+0
1+0
1+0
1+0, 2+0, P+1
2+0
1+0
1+0
1+0, 2+0, 2+1,
P+1
2+0, P+1
1+0
1+0
1+0, P+13
Container production
Container
Stock
typeb
type
179
180
0
0j42f
28d120h
28d120h
28d120h
28d120h
28d120h
28d120h
3060k
0j42f
30d45h
Jan
Late Marearly Apr
Fall
Spring
Mid-Marmid-Apr
Mid-Marmid-Apr
Fall
Spring
Early Marearly May
Early Jan (1+0) for fall/winter lift
Janearly Feb (1+0)
FebApr
30d60h
0
5070f
14d80h
30d80
30d120
30d120
3042f
30d120
30d120
Sowing season
2040
2040
(100/row)
2050
50
2050
3040
2035
215430
(330/row)
220540
540
220540
320430
220380
220430
11.5
2c
12.5
2c
0.51.5
0.51.5
3/ 1/
8 2
3/4c
3/81
3/ c
4
1/ 1/
4 2
1/ 1/
4 2
0.51.5 1/41/2
None
None
Bareroot production
Sowing depth
cm
in
Mulcha
Seedling density
/m2
/ft2
2+2
1+0, 2+0, 2+1, 3+0
3+0
Stock
type
415B,D, 615A
Styro 2,5,7
Styro 2,5,7
Styro 2,5
313B, 410A
313B, 410A, 415B,D, 615A
313B, 415B,D, 410A,
615A, Styro 2,5, Leach 1,2
313A
415B,D, 412A
313B, 410A
313B, 410A, 415B,D, 615A
313A,B, 410A, PCT410,
410A, 412A, 415B,D,415B, 615A
1+0
1+0, 2+0, P+1
1+0, 2+0
1+0
1+0
1+0
1+0, 2+0, 2+1,
P+1
2+0, P+1
2+0
1+0
1+0
1+0, 2+0, 2+1, P+1
1+0, 2+0, 2+1, P+1
Container production
Container
Stock
typeb
type
Sources: Adkins (1984),Adkins and others (1984),AOSA (1998),Asakawa (1968), Barton (1930), Bongio (1997), Bouvarel and Lemoine (1958), Curtis (1997), Fenimore (1997), Franklin (1974b), Garren (1997), Gates (1997), Hanson
(1997), Heit (1964, 1967, 1968b), Heit and Eliason (1940), Helson (1997), Henry and Blazich (1990), Holmsgaard and Kjaer (1951), ISTA (1993), Kusisto (1997), Lehar (1997), Leloup (1956), MacDonald (1998), Moore (1997),
Nagao and Asakawa (1963), NBV (1946), Pelton (1997), Rafn (1915), Rafn and Son (nd), Riskin (1997), Rutar (1991), Snyder (1997), Speers 1962, Stubley 1998,Thompson 1997,Toumey and Stevens (1928),Triebwasser
(1997),Trimble (1997),Tulstrup (1952), USDA Forest Service (1948),Vacowicz (1997),Wedman (1997),Wong (1997),Wright (1950), Zemanek (1997).
a Depth of mulch: peat moss, 0.51.5 cm (__ in); pine needles, 34.5 cm (1_1_ in); sawdust, 0.5 cm (_ in); straw, 5 cm (2 in).
b Containers are all PSB type; Styro = Styroblocks, PCT = copper treated.The various containers listed have the following volumes: 313A, 52 ml (3.6 in3); 313B, 65 ml (3.9 in3); 410A & PCT 410, 80 ml (4.9 in3); 415B, 93 ml
(6.3 in3); 412A, 126 ml
(7.7 in3); 415D, 172 ml (10.5 in3); 615A, 336 ml (20.0 in3); Styro 2, 39 ml (2.3 in3); Styro 5, 77 ml (4.7in3); Stryo 7, 121 ml (7.4in3); Leach 1, 50 ml (3 in3), Leach 2, 164 ml (10 in3).
c Seeds covered with 1 cm nursery soil plus 1 cm sand.
d For 2830 days only if intending to strat-redry.
e Some container transplants grown as P+2, P+3, P+4.
f Stratified in wet vermiculite, wet sand, or 1.5- to 2-day running-water soak and naked stratification.
g Used overwinter on first-year seedlings.
h Some free moisture left in plastic bag for long stratification.
i Light may be beneficial to germination.
j When not stratified, soaked 2 days before sowing.
k Alternatively, bury the seeds in snow for 50 days.
A. sachalinensis
A. x shastensis
A. pindrow
A. procera
A. nordmanniana
A. magnifica
var. magnifica
A. lasiocarpa
var. lasiocarpa
Species
Stratification
time (days)
extensive fungal and bacterial molding common to moreslowly-germinating unstratified seeds (Edwards 1969). In
noble fir, an increasing response to stratification as the seeds
matured suggested that dormancy increased also, and that
dormancy and maturity are interrelated (Edwards 1969).
Whereas much of the variability in dormancy among seedlots may be attributable to seed origin, crop year, and time
of collection, it may also be due to methods of cone processing, seed cleaning, and seed storage (Franklin 1974b;
Wang 1960).
Laboratory and nursery stratification is often performed
by refrigerating previously hydrated seeds in plastic bags or
other containersthe naked stratification method (Allen
and Bientjes 1954) favored in many nurseries for its ease of
seed handling. More traditionally, dry seeds (at storage
moisture contents) are placed on a moist medium (filter
paper, vermiculite, or wet sand) and refrigerated. The moist
filter paper method produced higher germination in noble fir
because it was believed that the preliminary water soak that
is the first step in the naked stratification procedure damaged the seeds by too-rapid tissue hydration, a phenomenon
well-documented in legumes (Jones and others 1991).
Soaking temperature in this noble fir study was 4 C.
However, no direct evidence for the damage, particularly its
location, was provided. It is unlikely that any damage
occurred in the tissues of the embryo. When noble fir seeds
were soaked in water at 25 C, after 48 hours most of the
water was still in the seedcoat: the outer region of the
megagametophyte had become moist, but the embryo was
still dry (Edwards 1969). It was found that noble fir
embryos require hydration of between 48 and 72 hours,
even at room temperature, before they absorb enough moisture to be safely excised (Edwards 1969). Furthermore,
when dry noble fir seeds are placed on a moist medium and
refrigerated, they absorb water slowly during the entire
chilling period and achieve a higher moisture content than
seeds soaked in water at room temperature for the same
length of time (Edwards 1971). Thus, in the above comparison, the moisture content of soaked seeds averaged 36%,
whereas that of seeds chilled on moist filter papr averaged
43% (Jones and others 1991). This difference, small as it
may appear, may have been significant due to the moisture
content in soaked seeds possibly being less than adequate
for optimal stratification to occur. In the development of the
stratification/redry method (see below), it was found that if
fir seeds were initially hydrated only to 35% moisture content (the same moisture content achieved after redrying),
subsequent stratification was far less effective (Edwards
1986). If noble fir seeds are sensitive to imbibitional damage as claimed (Jones and others 1991), then the stratification/redry methodwhich involves a preliminary soak at
room temperaturemust repair such damage since germination is greatly increased. However, no evidence for this
repair, or the initial imbibitional-damage phenomenon, has
been documented.
In any event, crop year, seed source, seed vigor (as distinct from seed quality), as well as chilling method and germination temperature played roles in the response of different seedlots of Pacific silver fir to stratification (Leadem
1986). Stratification response of Nordmann fir was also
believed to be strongly seedlot dependent (Poulsen 1996).
For balsam fir seeds, prolonged soaking in cold water containing a fungicide was deleterious (Kozlowski 1960), but
changing the water weekly produced germination similar to
that after stratification (Rudolf 1950). Best results with
Manchurian fir occurred when soaked seeds were stored in
snow for 1 to 2 months (Pavlenko 1972).
Stratification temperature range is often specified as 1 to
5 C (Franklin 1974b), although testing laboratories typically use a narrower window of 3 to 5 C. Stratifying grand
and subalpine fir seeds at 2 C was optimal (compared to
2, 5, and 7 C), especially during extended chilling
(Edwards 1982a). Fir seeds will germinate during stratification if left for a sufficient length of time (Allen 1960;
Edwards 1969; Blazich and Hinseley 1984; Roe 1948b;
Vabre-Durrieu 1956). Such observations reinforce the idea
that stratification is incipient germination. In this regard, it
should be remembered that late-dispersed seeds of numerous
high-elevation firs (plus some other conifers) germinate in
snow banks (Anon. 1951; Franklin and Krueger 1968;
Gordon 1970; Hetherington 1965; Irmak 1961; Roe 1946;
Stein 1951). Snow absorbs 99% of the infra-red (IR) radiation from sunlight, and dark-colored seeds embedded in
snow may reach several degrees above freezing by absorbing these IR rays. However, these germinants seldom establish as seedlings when the snow melts (Gordon 1970; Stein
1951).
Despite the fact that lower than normal levels of seed
moisture were known to benefit extended treatments of
hybrid fir seeds (Wright 1950), fir seed research continued
to focus on stratification temperature and duration and not
on moisture level during treatment. Since the 1980s it has
been demonstrated conclusively that seeds of Pacific silver,
grand, subalpine, and noble firs stratified at 2 to 5 C in
plastic bags for 4 weeks (moisture content 45% or higher),
then air-dried to moisture contents between 25 and 35%, can
be returned to the same refrigerator for (a) another 12
months (at 25%) without significant decreases in subsequent
germination or (b) a further 3 to 6 months (at 35%) with
greatly enhanced germination rate and germination capacity
(Edwards 1980b, 1981, 1982a,b, &c, 1986b, 1997; Leadem
1986, 1988b, 1989; Tanaka and Edwards 1986). When airAbies
181
The germination substrate is usually kept at its maximum moisture-holding capacity so the test samples are not
under any moisture stress but without excess free water
present. Full germination of Pacific silver and grand fir
seeds was unaffected unless the medium was moistened to
below 40% of maximum holding capacity (Edwards unpublished data). However, completeness of germination, and
germination rate of west Himalayan fir seeds was highly
sensitive to moistening the filter paper with PEG (polyethylene glycol) solution (Singh and others 1986). Many laboratories use a paper/blotter substrate as this allows easy evaluation of the radicles (figure 20 ), but porous mineral substrates such as perlite, vermiculite, and Sponge Rok may
be employed also. Tests conducted according to standard
laboratory prescriptions usually terminate after 21 or 28
days, although those on unstratified seeds may continue for
35 or 42 days. As a means of predicting operational sowing
requirements in nurseries, some agencies test stratified true
fir seeds in fumigated soils at temperatures of around 24 C
during the day and 18 C at night (Johnson 1984).
By the time newly harvested fir seeds have been
processed, there is often insufficient time to complete standard germination tests that require a minimum of 3 weeks
for completion, and more than twice this duration if the
seeds must be stratified, before they are required for sowing
the following spring. To provide more rapid estimates of
seed quality, several so-called quick tests have been developed. The simplest is the cutting test, but it is also the least
reliable because it fails to detect seeds damaged during
handling and processing or that have died during storage.
Abies
183
184
that the stratification/redry method (described earlier) broadens the range even further (Davidson and others 1984). This
temperature-range broadening is a sure sign of increased
vigor (Grabe 1976). One distinction between seed vigor and
seed germination can be seen in the effects of long-term
seed storage, which causes a reduction in plant percentage in
the nursery before it affects germination percentage
(Giannini and Murazio 1972; Muller 1977, 1980). Seed
vigor was related to germination rate, seed protein levels,
and seed respiration, all of which were thought to have
potential for development as quantifiable indices of this
variable in subalpine fir (Leadem 1988a&b, 1989).
Nursery practice. Fir seedlings are grown as both
bareroot and container stock. A 1997 survey found 20 nurseries growing almost 21 million seedlings of 16 (including 6
non-native) fir species for reforestation purposes. Several
other exotic firs are grown, especially in the northeastern
United States, for Christmas trees (Girardin 1997a&b). For
bareroot sowing in the past, most Pacific Northwest and
California nurseries stratified for 1 to 2 months (table 10) at
0 to 3 C, and sowed between mid-April to mid-May
(exceptionally as late as June), favoring a seedling density of
270 to 330 seedlings/m2 (25 to 30/ft2) (Lavender 1979)
(table 10). Bareroot sowing rates for Pacific silver, grand,
subalpine, and noble firs in British Columbian nurseries usually were lower220 to 240/m2 or 260 to 300/linear m of
seed bed (20 to 23/ft2 or 79 to 91/linear ft of seed bed)to
produce more open-grown plants (Arnott and Matthews
1982).
Although seeds of European silver, balsam, and Fraser
firs normally may be fall-sown in bareroot beds without
stratification (table 10) as are seeds of noble and white firs
raised in European nurseries (Franklin 1974b)spring-sowing of stratified seeds has been recommended for balsam
(Roe 1948b), and European silver firs (Neubacher 1959;
Paiero and Piussi 1964; Vlase and Iesan 1959). Fall-sowing
of freshly collected fir seeds may not be possible because
seed processing is incomplete, so sowing the following
spring provides the earliest opportunity. Spring-sowing of
stratified seeds is the traditional standard for most western
North American species (table 10), which minimizes losses
from birds, rodents, and adverse weather (Lanquist 1946).
Merely soaking grand fir seeds can be beneficial (Hofman
1966). Sowing unstratified seeds of grand and noble firs in
January to March or stratified seeds in April gave satisfactory results in the United Kingdom (Faulkner and Aldhous
1959). Most bareroot nurseries use a seedling caliper
between 2.5 and 5 mm (metric measure only) for culling
purposes.
species (table 10). Not every user succeeds with this technique, possibly due to differences in seedlot dormancy,
becauseas with routine stratificationthe
stratification/redry method has a greater effect on more-dormant fir seeds, less-dormant lots not benefiting as well.
Container seedlings of grand and noble firs grow quickly and evenly, so that 10- to 15-cm-tall plants can be
obtained about 20 weeks after sowing without using extended photoperiods. By artificially increasing daylengths to 18
hours, similarly sized Pacific silver fir seedlings (figure 21)
can be produced, but subalpine fir plants generally set bud
early and achieve no more than 6 cm of height (Arnott and
Matthews 1982; Gates 1994). When 5-month-old containergrown Fraser fir seedlings were naturally chilled outdoors
through mid-November (fluctuating temperatures and natural photoperiods), then returned to a greenhouse, at 15
months they were taller than conventionally grown 3+1 and
artificially chilled plants (Seiler and Kreh 1987).
Most containers are made of Styrofoam blocks with
cavities (Sjoberg 1974) or trays of individual plastic cells;
cavity and cell volumes vary widely (table 10). In general,
smaller containers are used for early sowing if the stock is to
be transplanted. Later sowings use bigger containers to produce bigger plants, some of which may be transplanted also
(table 10). The principles of container nursery technology
are well established (Landis and others 1989, 1990a&b,
1992, 1995), and the concept is now widely accepted.
Herbicides are not used at most container nurseries,
whereas bareroot facilities employ a range of chemicals;
recommendations for some of these (and for damping-off
control) have been published (Imai and others 1955; Roe
1948b; Sanftleben 1989; Sato 1962; Singh and Bhagat
Figure 21Abies amabilis, Pacific silver fir: seeds germinating in a container nursery; wooden toothpicks (left-rear
of cavities) were used to mark the progression of germination for a research trial (courtesy of C. L. Leadem).
Abies
185
186
References
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Northeast. Journal of Forestry 60: 9799.
Abbott HG, Hart AC. 1960. Mice and voles prefer spruce seeds. Sta.
Pap. 153. Upper Darby, PA: USDA Forest Service, Northeastern
Forest Experiment Station. 12 p.
Adkins CR. 1984. Role of stratification, temperature and light in Fraser
fir seed germination. Combined Proceedings of the International
Plant Propagators Society 33: 504515.
Adkins CR, Hinesley LE, Blazich FA. 1984. Role of stratification, temperature, and light in Fraser fir germination. Canadian Journal of Forest
Research 14: 8893.
Agee JK. 1982. True fir management for wilderness, water, recreation
and wildlife values. In: Oliver CD, Kenady RM, eds. Proceedings,
Symposium on Biology and Management of True Fir in the Pacific
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of Washington, Institute of Forest Resources: 227237.
Aguirre-Planter E, Furnier GR, Eguiarte LE. 2000. Low levels of genetic
variation within and high levels of genetic differentiation among
populations of species of Abies from southern Mexico and
Guatemala. American Journal of Botany 87: 362371.
Ahlgren CE. 1957. Phenological observations of nineteen native tree
species in northeastern Minnesota. Ecology 38: 622628.
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Abies
197
198
FabaceaePea family
Acacia L.
acacia
John K. Francis and Craig D.Whitesell
Dr. Francis retired from the USDA Forest Services International Institute of Tropical Forestry;
Dr.Whitesell retired from the USDA Forest Services Pacific Southwest
Forest and Range Experiment Station
Growth habit, occurrence, and use. The acacias
include about 1,200 species of deciduous or evergreen trees
and shrubs widely distributed in the tropics and warmer temperate areas (Guinet and Vassal 1978). Nearly 300 species
are found in Australia and about 70 in the United States.
Some 75 species are of known economic value, and about
50 of these are cultivated. Certain species of acacias
Cootamundra wattle (A. baileyana F. Muell.), Karoo thorn
(A. karroo Hayne), golden wattle (A. pycnantha Benth.), and
othersrank among the most beautiful of all flowering
trees, and many have been planted in the warmer regions of
the United States (LHBH 1976; Menninger 1962, 1964;
Neal 1965). Acacias produce many benefits: collectively
they yield lumber, face veneer, furniture wood, fuelwood,
and tannin; and such products as gum arabic, resins, medicine, fibers, perfumes, and dyes; some are useful for reclamation of sand dunes and mine spoils, and for shelterbelts,
agroforestry hedgerows, and forage; and some serve as a
host for the valuable lac insect (ACTI 1980; Prasad and
Dhuria 1989; Turnbull 1986). They are valuable not only to
the forest but also to pastures and agricultural crops for the
nitrogen that is fixed in their root nodules (Hansen and
others 1988).
Green wattle, introduced to Hawaii about 1890, has
been declared noxious for state land leases (Haselwood and
Motter 1966). A fast-growing tree of no local value, it
spreads rapidly by seeds and root suckers, crowding out
other plants. More than 90 years ago, Maiden (1908) commented on the pestiferous nature of several varieties of this
species in Australia. Only acacia species that do not spread
by suckering should be selected for planting. Also to be
avoided under most circumstances are the thorny acacias
such as sweet acacia and gum arabic treewhich are widely
dispersed rangeland pests. These 2 species are know to exert
allelopathic effects on plants growing near them (Hampton
and Singh 1979; Singh and Lakshminarayana 1992).
Reliable seed data are available on 8 species (table 1), all of
Acacia
199
Common names
A. auriculiformis A.
Cunningham ex Benth.
A. decurrens Willd.
A. decurrens var. normalis Benth.
A. farnesiana (L.) Willd.
Vachellia farnesiana (L.) Wright & Arn.
A. koa Gray
A. mangium Willd.
Mangium montanum Rumph.
A. mearnsii de Wildeman
A. decurrens var. mollis Lindl.
A. melanoxylon R. Br. ex Ait. f.
earleaf acacia
Australia
Australia
Native
Hawaii
Indonesia, New
Guinea, & Australia
Australia
Australia
US
Florida &
Puerto Rico
California
& Hawaii
S US, Puerto Rico,
& Virgin Islands
Hawaii
Hawaii &
Puerto Rico
California &
Hawaii
California &
Hawaii
Puerto Rico &
Virgin Islands
Height at
maturity
(m)
1230
818
35
2434
1230
15
2436
320
Source: Anderson (1968), Barrett (1958), Fagg (1992), Munoz (1959), Parrotta (1992),Turnbull (1987),Whitesell (1974).
longitudinal
200
Location
Florida
California
Puerto Rico
Hawaii
Puerto Rico
California
California
Hawaii
Puerto Rico
Flowering
Fruit ripening
Dispersal
MarApr
FebMar
NovFeb
JanJul
MarApr
Jun & later
FebJun
MayJun
Almost continuously
JunJul
AugDec
MarSep
Jun-Jul
JunOct
JulNov
MarDec
Feb; JunNov
MayAug
JunOct
JulDec or later
All year
All year
Species
A. auriculiformis
A. decurrens
A. farnesiana
A. koa
A. mangium
A. mearnsii
A. melanoxylon
A. nilotica
510
10
47
36
312
58
413
515
1.3
2.0
1.52.5
1.3
1.0
0.81.6
Cleaned seeds/wt
/kg
30,000158,000
53,00088,000
7,60013,000
5,30016,300
80,000110,000
33,00074,000
44,00088,000
5,00016,000
/lb
14,00072,000
26,00040,000
3,0006,000
2,0007,000
36,00050,000
15,00034,000
20,00040,000
2,0007,000
Sources: ACTI (1983), Fagg (1992), Goor (1968), Letourneux (1957), Magini and Tulstrup (1955), NFTA (1987a,b), Salazar (1989),Turnbull (1986),Whitesell (1974).
over 98% and grew 25% faster than control seedlings in the
first 3 months (De Zwaan 1978). Some species also appear
to require 2 to 4 months of after-ripening in dry storage
before good germination may be obtained (Whitesell 1974).
Germination is epigeal.
Germination testing. Prescriptions for official testing
for acacias call for clipping, nicking, or filing through the
seedcoats and soaking in water for 3 hours, or soaking seeds
in concentrated sulfuric acid for 1 hour, then rinsing thoroughly (ISTA 1993). Germination should then be tested on
moist blotter paper at alternating 20/30 C or constant 20 C
for 21 days. Germination tests of acacias can also be made
in flats with sand or soil. Results of tests for 8 species of
acacias are given in table 4.
Nursery and field practice. After proper pretreatment, the small-seeded acacias should be covered with 6 to
12 mm (1/4 to 1/2 in) of soil. Optimum sowing depth for
sweet acacia seeds was found to be 2 cm (3/4 in) (Scifres
1974). A 2:1 mixture of soil and sand proved to be a better
germination medium for gum arabic tree than other mix-
Acacia
201
Species
Seed source
Pretreatment
A. auriculiformis
Puerto Rico
Puerto Rico
Java
Puerto Rico
Hawaii
Australia
Tasmania
Tasmania
Victoria
Uruguay
Uruguay
Uruguay
None
Hot water
Warm water
Abrasion
Hot water
Hot water
Hot water
Hot water
Hot water
None
H2SO4
Abrasion
Hot water
Hot water
A. decurrens
A. farnesiana
A. koa
A. mangium
A. mearnsii
A. melanoxylon
A. nilotica
Medium
(C)
Soil
Soil
Soil
Paper
Soil
Soil
Paper
Paper
Paper
Soil
Soil
79
60
77
77
77
68
68
75
(days)
21
14
85
30
30
10
14
60
30
90
30
21
28
15
85
30
(%)
56
30
74
56
18
80
72
70
74
93
4
48
26
52
74
Sources: ACTI (1983), Francis and Rodriguez (1993), Newman (1989a), Parrotta (1992),Webb and others (1984), (1986),Whitesell (1974).
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Firewood crops, shrub and tree species for energy production.
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ACTI [Advisory Committee on Technology Innovation]. 1983.
Mangium and other fast-growing acacias for the humid tropics.
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Anderson RH. 1968. The trees of New South Wales, 4th ed. Sydney,
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Atchison E. 1948. Studies on the Leguminosae: 2. Cytogeography of
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Bahuguna VK, Pyare L. 1990. To study the effects of environment and
different soil mixture on germination of Acacia nilotica seed at
nursery stage. Journal of Tropical Forestry 5(1): 5156.
Barrett MF. 1956. Common exotic trees of south Florida
(Dictyledons). Gainesville: University of Florida Press. 414 p.
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germination of Acacia. Australian Journal of Botany 25(3): 269276.
De Zwaan JG. 1978. The effects of hot-water-treatment and scarification on germination of blackwood (Acacia melanoxylon) seed. South
African Forestry Journal 105: 4042.
Fagg CW. 1992. Acacia nilotica: pioneer for dry lands. NFTA 92-04.
Paia, HI: Nitrogen Fixing Tree Association. 2 p.
Francis JK, Rodrguez, A. 1993. Seeds of Puerto Rican trees and shrubs:
second installment, Res. Note SO-374. New Orleans: USDA Forest
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Galiana A, Chaumont J, Diem HG, Dommergues 1990. Nitrogenfixing potential of Acacia mangium and Acacia auriculiformis
seedlings inoculated with Bradyrhizobium and Rhizobium spp. Biology
and Fertility of Soils 68(4): 263272.
Goor AY, Barney CW. 1968. Forest tree planting in arid zones. New
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Guinet P,Vassal J. 1978. Hypothesis on the differentiation of the major
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509527.
Gunn BV. 1990. Germination pretreatments for selected acacia species
from the Pilbara region of Western Australia. ACIAR Proceedings
Series (Australia) 28: 4650.
202
Hampton CO, Singh SP. 1979. The presence of growth and germination inhibitors in the seeds of certain desert plants.Transactions of
the Kansas Academy of Science 82(2): 87.
Hansen AP, Stoneman G, Bell DT. 1988. Potential inputs of nitrogen by
seeder legumes to the Jarrah forest ecosystem. Australian Forestry
51(4): 226231.
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Huang LS. 1989. Study on the introduction and cultural techniques of
Acacia auriculiformis. Forest Science and Technology (China) 5:
1011.
ISTA [International Seed Testing Association]. 1993. Rules for testing
seeds: rules 1993. Seed Science and Technology 21 (Suppl.): 1259.
Judd CS. 1920. The koa tree. Hawaiian Forestry and Agriculture 17(2):
3035.
Kumar A, Gupta BB. 1990. Production of field plantable seedlings of
Acacia nilotica in fifty days. Indian Forester 116(4): 306322.
Kumar P, Purkayastha BK. 1972. Note on germination of the seeds of
lac hosts. Indian Journal of Agricultural Sciences 42(5): 430431.
Letourneux C. 1957. Tree planting practices in tropical Asia. For. Dev.
Pap. 11. Rome: FAO: 109111.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise
dictionary of plants cultivated in the United States and Canada.
New York: Macmillan: 47.
Mangini E,Tulstrup NP. 1955. Tree seed notes. For. Dev. Pap. 5. Rome:
FAO. 354 p.
Maiden JH. 1908. The forest flora of New South Wales.Volume 3.
Sydney, Australia: W.A. Gullick, Government Printer. 180 p.
Menninger EA. 1962. Flowering trees of the world. New York:
Hearthside Press. 336 p.
Menninger EA. 1964. Seaside plants of the world. New York:
Hearthside Press. 303 p.
Munz PA. 1959. A California flora. Berkeley: University of California
Press. 1861 p.
Natarajan N, Rai RSV. 1988. Studies to maximization of germination in
Acacia auriculiformis. Indian Journal of Forestry 11(4): 304306.
Neal MC. 1965. In gardens of Hawaii. Spec. Pub. 50. Honolulu: Bishop
Museum Press. 924 p.
Newman V. 1989a. Effects of pretreatments on germination of Acacia
mangium (Willd.) in Sabah. FRC publication 3/89. Sandakan, Sabah:
Forest Research Centre. 21 p.
Newman V. 1989b. Results of media trials using sawdust on germination of Acacia mangium (Willd.) FRC Publication 53. Sandakan,
Sabah: Forest Research Centre. 17 p.
NFTA [Nitrogen Fixing Tree Association]. 1987. Acacia auriculiformis:
the adaptable tropical wattle. NFTA 87-03. Waimanalo, HI: NFTA.
2 p.
NFTA [Nitrogen Fixing Tree Association]. 1987. Acacia mangium: a fastgrowing tree for the humid tropics. NFTA 87-04. Waimanalo, HI:
NFTA. 2 p.
NFTA [Nitrogen Fixing Tree Association]. 1992. Acacia nilotica: pioneer
for dry lands. NFTA 92-04. Waimanalo, HI: NFTA. 2 p.
Parrotta JA. 1992. Acacia farnesiana (L.) Willd., aroma, huisache. Res.
Note SO-ITF-SM-49. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 6 p.
Parrotta JA. 1994. Personal communication. Ro Piedras, PR: USDA
Forest Service, International Institute of Tropical Forestry.
Parry MS. 1956. Tree planting practices in tropical Africa. FAO For. Dev.
Pap. 8. Rome: FAO. 302 p.
Prasad R, Dhuria SS. 1989. Reclamation of iron-ore mined-out areas:
biomass production efficiency of species. Journal of Tropical Forestry
5(1): 5156.
Rana U, Nautiyal AR. 1989. Coat imposed dormancy of Acacia
farnesiana seeds. Seed Research 17(2): 122127.
Salazar R. 1989. Genetic variation of 16 provenances of Acacia
mangium at nursery level in Turrialba, Costa Rica. Commonwealth
Forestry Review 68(4): 263272.
Acacia
203
AceraceaeMaple family
Acer L.
maple
John C. Zasada and Terry F. Strong
Dr. Zasada retired from the USDA Forest Services North Central Research Station; Mr. Strong is a research
silviculturalist at the USDA Forest Services North Central Research Station, Rhinelander,Wisconsin
Growth habit, occurrence, and use. Maples
members of the genus Acerare deciduous (rarely evergreen) trees; there are 148 species (de Jong 1976; Van
Gelderen and others 1994). The majority of species originate
in central and eastern Asia, China, and Japan (de Jong 1976;
Van Gelderen and others 1994; Vertrees 1987). There are
several taxonomic treatments available for the genus.
Vertrees (1987) and Van Gelderen and others (1994) should
be consulted for a discussion and comparison of the different classifications. Van Gelderen and others (1994) recog-
Common name(s)
Occurrence
A. circinatum Pursh
A. ginnala Maxim.
A. glabrum var. glabrum Torr.
A. grandidentatum Nutt.
A. griseum (Franch.) Pax
A. macrophyllum Pursh
A. negundo L.
Negundo aceroides (L.) Moench.
A. palmatum Thunb.
A. pensylvanicum L.
A. striatum DuRoi.
A. platanoides L.
A. pseudoplatanus L.
A. rubrum L.
A. carolinianum Walt.
A. saccharinum L.
A. dasycarpum Ehrh.
A. saccharum Marsh.
A. saccharophorum K. Koch
A. spicatum Lam.
paperbark maple
bigleaf maple, broadleaf
maple, Oregon maple
boxelder, ashleaf maple,
California boxelder
Japanese maple
striped maple, moosewood
Norway maple
planetree maple, sycamore maple
red maple, soft maple,
swamp maple
silver maple, river
maple, soft maple
sugar maple, rock
maple, hard maple
mountain maple
Sources: De Jong (1976), Dirr (1990), Fischer (1990), Olson and Gabriel (1974), Rehder (1940),Van Gelderen and others (1994),Vertrees (1987),Viereck and Little
(1972).
* Introduced into subarctic interior Alaska, where it forms a small tree and produces viable seeds (Viereck 1996).
204
Species
A. circinatum
A. ginnala
A. glabrum var. glabrum
A. griseum
A. macrophyllum
A. negundo
A. palmatum
A. pensylvanicum
A. platanoides
A. pseudoplatanus
A. rubrum
A. saccharinum
A. saccharum
A. spicatum
Height (m)
at maturity
Year first
cultivated
Minimum
seed-bearing
age (yrs)
9
6
9
8
35
23
6
11
31
31
28
28
31
9
1826
1860
1882
1901
1812
1688
1820
1755
Long ago
Long ago
1656
1725
Long ago
1750
10
4
11
22
Years
between large
seedcrops
12
1
13
1
1
1
1
1
1
37
Sources: Burns and Honkala (1990), Dirr (1990), De Jong (1976), Olson and Gabriel (1974),Vertrees (1987).
Note: A. rubrum, A. negundo, A. pensylvanicum, and A. saccharinum are dioecious to varying degrees.The other species are monoecious, but male and female flowers may
occur in different parts of the tree.
actually classify the 4 species mentioned above as subspecies of sugar maple. Eight of the 16 sections of the genus
are represented in North America (Van Gelderen 1994).
Additionally, a number of species (table 1) have been introduced for use as ornamentals (Burns and Honkala 1990;
Dirr 1990; Dirr and Heuser 1987; Fischer 1990; Van
Gelderen and others 1994; Vertrees 1987). The native
species range in size from trees that dominate forest
canopies to medium to tall understory shrubs or small trees
(table 2). Boxelder has been introduced into Alaska, where it
survives and reproduces; however, it does dieback periodically under extreme winter temperatures (Viereck 1997).
The native maples all regenerate vegetatively by basal
sprouting, but the ability to do so varies among species and
with plant age (Burns and Honkala 1990; Fischer 1990).
Vine, Rocky Mountain, striped, and mountain maples frequently layer, giving them the potential to develop relatively
complex clones of varying size and morphology (Hibbs and
Fischer 1979; ODea and others 1995; Post 1969; Zasada
and others 1992).
Some species of maple are important sources of firewood, pulpwood, high-quality lumber, and veneer (Alden
1995; Burns and Honkala 1990). Four species have been
used to produce maple sugar and syrupsugar, black, red
(Jones 1832; USDA FS 1982), and bigleaf maple. Sugar
maple is the most important of these species because it has
the highest sugar content. In the western United States,
bigleaf maple produces adequate quantities of sap, but its
sugar content is low compared to the sap of sugar and red
maples, and the flow is erratic (Burns and Honkala 1990).
205
Flowering
Fruit ripening
Seed dispersal
A. circinatum
A. ginnala
A. glabrum var. glabrum
A. macrophyllum
A. negundo
A. palmatum
A. pensylvanicum
A. platanoides
A. pseudoplatanus
A. rubrum
A. saccharinum
A. saccharum
A. spicatum
MarJune
AprJune
AprJune
AprMay
MarMay
MayJune
MayJune
AprJune
AprJune
MarMay
FebMay
MarMay
MayJune
SeptOct
AugSept
AugOct
SeptOct
AugOct
AugSept
SeptOct
SeptOct
AugOct
AprJune
AprJune
SeptOct
SeptOct
OctNov
SeptJan
SeptFeb
OctMar
SeptMar
SeptOct
OctFeb
OctNov
SeptNov
AprJuly
AprJune
OctDec
OctDec
Sources:
206
Dirr (1990), Burns and Honkala (1990), Olson and Gabriel (1974).
Johnson 1992; Sipe and Linnerooth 1995). Each filled samara typically contains a single seed without endosperm
(figure 2). However, polyembryony has been observed in
sugar and bigleaf maples (Carl and Yawney 1972; Zasada
1996). Maple seeds turn from green or rose to yellowish or
reddish brown when ripe; the color of mature samaras can
vary among species. Pericarps have a dry, wrinkled appearance when fully mature (Albenskii and Nikitin 1956; Anon.
1960; Carl and Snow 1971; Harris 1976; Rehder 1940;
Sargent 1965; Vertrees 1987).
The embryo with associated seedcoats is contained
within the pericarp (figure 2). The surface of the pericarp is
usually glabrous (except that of bigleaf maple, which has
dense, reddish brown pubescence). The pericarp can be
extremely hard (particularly when it has dried out) and difficult to cut open. Development of the samara in black maple
has been described in detail by Peck and Lersten (1991).
Both the pericarp and seedcoat have been identified as causes of dormancy. The cavity (locule) in which the embryo
occurs may have concave or convex walls. There are 2 types
of embryo folding: (a) incumbent folding, in which the
hypocotyl is against the back of one cotyledon, and (b)
accumbent folding, in which the hypocotyl is against the
edges of the folded cotyledons. Of the native maples, vine
and sugar maples are classified as incumbent and the others
(except bigtooth maple, which was not classified) are
accumbent. The cotyledons may be green while still in the
pericarp (Carl and Yawney 1972; De Jong 1976; Dirr and
Heuser 1987; Olson and Gabriel 1974; Peck and Lersten
1991; Vertrees 1987).
Figure 2Acer circinatum, vine maple: longitudinal section of a seed showing bent embryo. On drying the seed
shrinks, leaving space between the seedcoat and the
pericarp.
Acer
207
/kg
A. circinatum
A. ginnala
A. glabrum var. glabrum
A. macrophyllum
A. negundo
A. pensylvanicum
A. platanoides
A. pseudoplatanus
A. rubrum
A. saccharinum
A. saccharum
A. spicatum
Source:
208
7,71012,220
22,98044,640
17,28044,860
5,9708,840
18,12045,080
21,43034,400
2,81010,300
6,48015,910
28,07084,420
1,9907,070
7,07020,110
33,81060,330
Average
/lb
3,4905,530
10,40020,200
7,82020,300
2,7004,000
8,20020,400
9,70015,600
1,2704,660
2,9307,200
12,70038,200
9003,200
3,2009,100
15,30027,800
/kg
10,210
37,570
29,680
7,180
29,610
24,530
6,320
11,290
50,520
3,930
15,540
48,910
/lb
4,620
17,000
13,430
3,250
13,400
11,100
2,860
5,110
22,860
1,780
7,030
22,130
Seeds/tree (thousands)
11.9
54.3
91.4
955.8
Species
vine maple
bigleaf maple
sugar maple
Weight/volume of samaras
kg/hl
lb/bu
15.3
11.9
5.9
4.6
13.1
10.2
Seed collection for most species occurs when the samaras are fully ripened and the wing and pericarp have turned
tan or brown in color (Carl 1982a; Carl and Yawney 1966).
However, for maples that are difficult to germinatesuch as
vine maple, striped maple, and the Japanese maplesit is
recommended that seeds be collected before they have dried
completely, when the wing has turned brown but the pericarp is still green (Dirr and Heuser 1987; Vertrees 1975,
1987).
Although the seeds of most maples are glabrous, those
of bigleaf maple are often densely pubescent. The pubescence may irritate the skin and cause some respiratory tract
congestion when airborne. Individuals who might be sensitive to this material should use rubber gloves and a face
mask.
Extraction and storage of seeds. Maple seeds are
generally not extracted from the fruits (samaras) after collection, except when seeds are used in research on seed dormancy or lots of particularly valuable seeds that are difficult
to germinate. De-winging reduces weightwings account
for about 15 to 20% of samara weight (Greene and Johnson
1992; Sipe and Linnerooth 1995)and bulk for storage. The
separation of filled and empty samaras for sugar maple can
be accomplished on small lots by floating the samaras in
n-pentane (filled seeds sink). This practice had no apparent
effect on long-term seed viability (Carl 1976, 1982a; Carl
and Yawney 1966). Removal of empty samaras, which can
be done readily on a gravity table, improves seed handling,
storage, sowing, and control of seedbed density.
After dispersal, maple seeds (with the exception of silver maple seeds and some red maple seeds) lie dormant in
Acer
209
210
Acer
211
Table 5Acer, maple: warm and cold stratification treatments for internal dormancy
Warm period
Cold period
Species
Temp (C)
Days
Temp (C)
Days
A. circinatum*
A. ginnala*
A. glabrum
A. macrophyllum
A. negundo*
A. palmatum (dry seeds)
A. palmatum (fresh seeds)
A. pensylvanicum
A. platanoides
A. pseudoplatanus
A. rubrum
A. saccharinum
A. saccharum
A. spicatum
2030
2030
2030
Warm water
3060
3060
180
12
3
5
35
15
5
18
18
5
5
15
3
15
5
90180
90150
180
4060
6090
60120
60120
90120
90120
4090
6090
0
4090
90120
Sources: Browse (1990), Dirr and Heuser (1987), Harris (1976), Olson and Gabriel (1974),Vertrees (1987).
Note: Even after standard pretreatment, seedlots of A. griseum may require 2 to 3 years for complete germination.
* Mechanical rupture of the pericarp may improve germination.This is necessary in A. negundo when seeds are very dry; a warm soak as for A. palmatum may suffice.
The benefit of warm incubation prior to stratification is not well-documented. Seeds may go through at least 1 warm/cold cycle before germinating under field conditions.
Water temperature at start of incubation is 40 to 50 C and allowed to cool gradually. Some recommend a 21 C incubation period following warm water treatment
and a 90-day stratification period.
Requirement for stratification is highly variable. In all seedlots, some seeds will germinate without stratification.
212
Species
A. circinatum
A. ginnala
A. glabrum
A. macrophyllum*
Source 1
Source 2
Source 3
A. negundo
A. pensylvanicum
A. platanoides
A. pseudoplatanus
A. rubrum
Low elevation (U)
Low elevation (S)
High elevation (U)
High elevation (S)
A. saccharinum
A. saccharum
A. spicatum
Germination rate
Amount
Time
(%)
(days)
Total
germination (%)
30
30
1016
20
20
1016
38
38
12
50
40
10
10
30
19
52
23
23
23
5
23
410
23
23
23
5
23
410
120
120
120
2460
90
60
1566
013
892
1467
2437
6090
6090
6090
1448
2097
100
100
100
2496
82
76
3081
5071
15
15
15
15
30
23
5
5
5
5
30
23
518
90
7291
80
32
313
75
31
55
89
13
54
9497
95
34
Sources: Olson and Gabriel (1974), Farmer and Goelz (1984), Farmer and Cunningham (1981),Vertrees (1987).
Notes: Germination rate indicates the number of seeds germinating in the time specified and total germination all of the seeds germinating in the test.The length of
germination tests are not same for all species.
Seeds of A. griseum and A. palmatum are very difficult to germinate and seed quality is usually poor. Cutting tests are recommended to determine potential viability.
Tetrazolium tests could be used to determine if seeds are alive; knowing this one can sow and wait several years for seeds to germinate. Because the delay in germination
appears related to a very hard pericarp, removing the pericarp can improve germination.
* Seed sources from central Oregon coastal range. Germination rate greatly increased when seeds moved to 20 to 25 C when germination in stratification begins
(Zasada 1996).
Germination of seeds with testa removed over radicles. Seeds with testae did not germinate at 23 C even after 5 months of stratification, whereas seeds kept at 5 C
germinated completely after 6 months (Wilson and others 1979).
Seed sources from Tennessee, total germination at higher temperatures was lower than shown here (Farmer and Cunningham 1981). Similar trends were observed with
red maple from Ontario (Farmer and Goelz 1984). U = stratified seeds, S = unstratified seeds.
Acer
213
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FabaceaePea family
Adenanthera pavonina L.
peronas
J. A. Vozzo
Dr.Vozzo retired from the USDA Forest Services Southern Research Station
seed.
Adenanthera
217
seedling,
cross section
References
Ahmed FU, Sharmila-Das, Hossain MA. 1983. Effect of seed treatment on
the germination of Rakta Kambal seeds. Bano-Biggyan-Patrika 12(1/2):
6265.
Bailey LH. 1941. Standard cyclopedia of horticulture. New York: Macmillan.
1200 p.
Francis JK. 1994. Personal communication. Ro Piedras, PR: USDA Forest
Service, International Institute of Tropical Forestry.
Francis JK, Liogier HA. 1991. Naturalized exotic tree species in Puerto
Rico. Gen.Tech. Rep. SO-82. New Orleans: USDA Forest Service,
Southern Forest Experiment Station. 12 p.
Francis JK, Rodrguez A. 1993. Seeds of Puerto Rican trees and shrubs: second installment. Res. Note SO-374. New Orleans: USDA Forest Service,
Southern Forest Experiment Station. 5 p.
Gunn CR. 1984. Fruits and seeds of genera in the subfamily
Mimosoidaceae (Fabaceae).Tech. Bull. 1681. Washington, DC: USDA
Agricultural Research Service. 194 p.
218
Little EL Jr, Wadsworth FH. 1964. Common trees of Puerto Rico and the
Virgin Islands. Agric. Handbk. 249. Washington, DC: USDA Forest Service.
548 p.
Neal MC. 1965. In gardens of Hawaii. Spec. Pub. 50. Honolulu: Bishop
Museum Press. 924 p.
Sandiford M. 1988. Burnt offerings: an evaluation of the hot-wire seed scarifier. Commonwealth Forestry Review 67(3): 285292.
Streets RJ. 1962. Exotic forest trees in the British Commonwealth. Oxford,
UK: Clarendon Press. 750 p.
Troup RS. 1921. Silviculture of Indian trees. Volume 2, Leguminosae
(Caesalpinieae) to Verbenaceae. Oxford, UK: Clarendon Press. 783 p.
Xu BM, Gu ZH. 1985. Effect of sulfuric acid treatment in breaking
dormancy in hard seeds. [Beijing]: Plant Physiology Communications
2: 2225.
HippocastanaceaeHorsechestnut family
Aesculus L.
buckeye
Paul O. Rudolf and Jill R. Barbour
Dr. Rudolph (deceased) retired from the USDA Forest Services North Central Forest Experiment Station;
Ms. Barbour is a germination specialist at the USDA Forest Services National Seed Laboratory
Dry Branch, Georgia
Growth habit, occurrence, and use. The buckeyes
which occur in North America, southeastern Europe, and
eastern and southeastern Asiainclude about 25 species of
deciduous trees and shrubs (Rehder 1940). They are cultivated for their dense shade or ornamental flowers, and the
wood of some species is occasionally used for lumber and
paper pulp. They also provide wildlife habitat. The shoots
and seeds of some buckeyes are poisonous to livestock
(Bailey 1939). Seven of the 9 species described (table 1) are
native to the United States. The horsechestnut was introduced into this country from southern Europe, and the
Himalayan horsechestnut occurs naturally in the Himalayas.
Seven of these 8 species are not used much in reforestation, but all are used for environmental forestry planting.
Himalayan horsechestnut is used extensively for reforestation and the nuts are fed to sheep and goats (Maithani and
Common name(s)
Occurrence
California buckeye
A. glabra Willd.
A. glabra var. arguta (Buckl.) B.L. Robins.
A. arguta Buckl.
A. glabra var. buckleyi Sarg.
A. buckleyi (Sarg.) Bush
A. hippocastanum L.
A. indica (Wall. ex. Cambess) Hook.
A. parviflora Walt.
A. pavia L.
A. sylvatica Bartr.
A. neglecta Lindl.
A. georgiana Sarg.
A. neglecta var. georgiana (Sarg.) Sarg.
Source:
horsechestnut,
chestnut, bongay
Himalayan horsechestnut
bottlebrush buckeye
red buckeye, scarlet buckeye,
woolly buckeye, firecracker plant
painted buckeye,
dwarf buckeye,
Georgia buckeye
Rudolph (1974).
Aesculus
219
220
longitudinal
soon after they have fallen. The fruits may be dried for a
short time at room temperature to free the seeds from any
parts of the capsules that may still adhere to them, but great
care must be taken not to dry them too long. When this
occurs, the seedcoats become dull and wrinkled and the
seeds lose their viability. There is ample evidence that buckeyes are recalcitrant in nature (Bonner 1969; Pence 1992;
Tompsett and Pritchard 1993). Moisture contents at the time
of shedding have been reported as 49% for horsechestnut
(Suszka 1966) and 56% for red buckeye (Bonner 1969). The
seeds of this genus should be sown at once in the fall or
stratified promptly for spring-sowing.
Buckeye seeds must be stored with moisture contents
close to what they are shed with, but even then their viability cannot be maintained very long. Initial viability of fresh
seeds of horsechestnut was maintained for 6 months when
they were stored in polyethylene bags at 1 C. This storage
condition is the same as cold moist stratification because of
the high moisture content of fresh seeds (Suszka 1966).
When seeds were stored at 1 C in sealed packages without
added moisture for 13 months, germination dropped from
85% to 60%; after 15 months, however, germination was
only 25% (Widmoyer and Moore 1968). Data on number of
cleaned seeds per weight are listed for 7 species in table 4.
Purity and soundness usually are close to 100% (Rudolf
1974). Nonviable seeds will float in water and can be discarded (Browse 1982).
Pregermination treatments. Seeds of Ohio, yellow,
and painted buckeyes and horsechestnut require stratification
or prechilling to induce prompt germination (Rudolf 1974).
Stratification has been done in moist sand or sand-peat mix-
Species
Location
Flowering
Fruit ripening
Seed dispersal
A. californica
A. flava
A. glabra
var. arguta
S California
Texas
Minnesota
Europe & NE US
SW Georgia, Alabama
South part of range
North part of range
Minnesota
AprSept
AprJune
MarMay
MarApr
May
Late Aprearly June
July-Aug
MarApr
MayJune
AprMay
May
SeptOct
Sept
Septmid-Oct
MayJune
SeptOct
Mid-Septearly Oct
OctNov
SeptOct
SeptOct
JulyAug
SeptOct
SeptOct
Mid-Septmid-Oct
OctNov
SeptNov
SeptNov
JulyAug
SeptOct
A. hippocastanum
A. parviflora
A. pavia
A. sylvatica
Sources: Brown and Kirkman (1990), Harrar and Harrar (1962), Little (1953), Loiseau (1945), NBV (1946), Radford and others (1964), Rehder (1940), Rudolf (1974),
Sargent (1965), Sus (1925),Turner (1969), van Dersal (1938),Vines (1960),Wyman (1947).
Height at
maturity Year 1st
(m)
cultivated
Species
A. californica
A. flava
A. glabra
var. arguta
A. hippocastanum
A. parviflora
A. pavia
A. sylvatica
Sources:
height, year first cultivated, flower color, seed-bearing age, seed crop frequency, and fruit
4.512
7.527
921
211
7.524
4.56
2.58.5
7.520
1855
1764
1809
1909
1576
1711
1826
Min
seedYears
bearing of large
age (yr) seedcrops
Flower color
White to rose
Yellow
Pale greenish yellow
Light yellowish green
White tinged with red
White
Bright red
Pale yellow, red
veins towards base
8
8
12
1+
12
1+
Ripe
color
Yellowish
Green
Yellow
Green
Yellow-green
Pale brown
Yellowish
Yellowish
Yellowish green
Yellowish brown
Light brown
Yellowish tan
Brown and Kirkland (1990), Rehder (1940), Rudolf (1974), Sargent (1965)
Average
Species
Place collected
/kg
/lb
/kg
/lb
A. californica
1836
816
26
127
6066
106148
71104
5175
4060
68126
2730
4867
3247
2334
1827
3157
62
128
88
64
51
117
88
28
58
40
29
23
53
40
A. flava
A. glabra
var. arguta
A. hippocastanum
A. parviflora
A. pavia
A. sylvatica
Aesculus
221
ommendation for germinating seeds of horsechestnut without stratification is to soak them in water for 48 hours and
cut off one-third of the seed at the scar end without removing the seedcoat. The portion with the scar should then be
germinated in sand flats for for 21 days at the same
30/20 C regime (ISTA 1993).
Nursery practice. Under natural conditions, seeds of
most buckeye species germinate in the early spring.
California buckeye, however, germinates just after winter
rains have begun, usually in November. In the nursery,
buckeye seeds usually are sown in the fall as soon after collection as possible to prevent drying and loss of viability. If
desired, however, the seeds of species having embryo dormancy can be stratified or placed in cold, moist storage
promptly and then sown in the spring (Rudolf 1974; Suszka
1966). Himalayan horsechestnut seeds without any treatment showed 80% germination after 133 days (Maithini and
others 1990). Seeds sown after 30 days of cold stratification
showed 68% germination in 78 days (Maithini and others
1990). The seeds should be sown about 5 cm (2 in) apart in
rows 15 cm (6 in) apart (NBV 1946) and covered with 2.5
to 5 cm (1 to 2 in) of soil. The seeds should be sown with
the scar underneath so that the radicle emerges in the correct
position to produce a normal seedling (Browse and Leiser
1982). If the seeds are variable in size, it is better to grade
them so that small sizes are discarded or sown separately, as
these rarely make large 1-year seedlings (Browse 1982).
Germination is hypogeal (figure 3) and usually is complete 3 to 4 weeks after spring sowing (NBV 1946). A tree
percentage of 70 has been obtained (Rudolf 1974). The beds
should not be over-watered because the seeds rot rather easily (Rudolf 1974). Ordinarily, 1+0 stock is large enough for
field planting.
Table 5Aesculus, buckeye: cold stratification periods, germination test conditions, and results
Species
A. californica
A. flava
A. glabra
var. arguta
A. hippocastanum
A. pavia
A. sylvatica
Cold
stratification*
(days)
Daily
light
(hrs)
Medium
Day
0
120
120
120
120
0
90
Sand
Sand
Sand
Sand
Sand
Kimpak
Sand
30
30
30
24
30
30
Temp (C)
Night
20
20
20
17
20
20
Sources:
May (1963), NBV (1946), Rudolf (1974), Suszka (1966), Widmoyer and Moore (1968).
* Cold stratification temperatures ranged from 0.5 to 5 C.
222
Days
20
40
40
30
30
30
30
Germinative
energy
Amount Time
Germinative
(%)
(days)
(%)
62
62
27
20
56
76
59
76
89
70
78
References
AOSA [Association of Official Seed Analysts]. 1998. Rules for testing seeds.
In: Proceedings of the Association of Official Seed Analysts: 1123.
Bailey LH. 1939. The standard encyclopedia of horticulture. New York:
Macmillan. 3639 p.
Bonner FT. 1969. Personnel communication. Starkville, MS: USDA Forest
Service, Southern Forest Experiment Station.
Brown CL, Kirkman KL. 1990. Trees of Georgia and adjacent states.
Portland, OR:Timber Press. 292 p.
Browse PM. 1982. The propagation of the hardy horse chestnuts and buckeyes. Plantsman 4(3): 150164.
Browse PM, Leiser AT. 1982. The California buckeye. Plantsman 4(1): 5457.
Harrar ES, Harrar JG. 1962. Guide to southern trees. 2nd ed. New York:
Dover Publishing. 709 p.
ISTA [International Seed Testing Association]. 1993. Rules for testing seeds:
rules 1993. Seed Science and Technology 21 (Suppl.): 1259.
Little EL Jr. 1953. Checklist of native and naturalized trees of the United
States (including Alaska). Agric. Handbk. 41. Washington, DC: USDA
Forest Service. 472 p.
Loiseau J. 1945. Les arbres et la fort. Paris. 204 p.
Maithani GP, Thapliyal RC, Bahuguna VK, Sood OP. 1990. Enhancement of
seed germination and seedling growth of Aesculus indica by stratification.
Indian Forester 116(7): 577580.
May C. 1963. Note on storage of buckeye and horsechestnut seed.
American Horticulture Magazine 42(4): 231232.
NBV [Nederlandsche Boschbouw Vereeniging]. 1946. Boomzaden: handleiding inzake het oogsten, behandelen, bewaren en uitzaaien van
boomzaden [Tree seed: handbook on the collection, extraction, storage,
and sowing of tree seed]. Wageningen,The Netherlands: Ponsen and
Looijen. 171 p.
Pence VC. 1992. Desiccation and the survival of Aesculus, Castanea, and
Quercus embryo axes through cryopreservation. Cryobiology 29:
391399 [Seed Abstracts 1994; 17(1): 21].
Radford AE, Ahles HE, Bell CR. 1964. Guide to the vascular flora of the
Carolinas. Chapel Hill: University of North Carolina Book Exchange.
383 p.
Rehder A. 1940. Manual of cultivated trees and shrubs hardy in North
America. 2nd ed. New York: Macmillan. 996 p.
Rudolf PO. 1974. Aesculus L., buckeye. In: Schopmeyer CS, tech. coord.
Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 195200.
Sargent CS. 1965. Manual of the trees of North America (exclusive of
Mexico). 2nd ed., corrected and reprinted. New York: Dover. 910 p.
Sus NI. 1925. Pitomnik [The forest nursery]. Moscow. 227 p.
Suszka B. 1966. Conditions for breaking of dormancy of germination of the
seeds of Aesculus hippocastanum L. Arbor. Krnicke. 11: 203220.
Tarabrin VP,Teteneva TR. 1980. Presowing treatment of seeds and its effect
on the resisitance of seedlings of woody plants against drought. Soviet
Journal of Ecology 10 (3): 204211.
Tompsett PB, Pritchard HW. 1993. Water status changes during development in relation to the germination and desiccation tolerance of
Aesculus hippocastanum L. seeds. Annals of Botany 71: 107116.
Turner BL. 1969. Personal communication. Austin: University of Texas.
Van Dersal WR. 1938. Native woody plants of the United States: their erosion control and wildlife values. Misc. Pub. 303. Washington, DC: USDA.
362 p.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
Widmoyer FB, Moore A. 1968. The effect of storage period temperature
and moisture on the germination of Aesculus hippocastanum seeds. Plant
Propagator 14(1) : 1415.
Wyman D. 1947. Seed collecting dates of woody plants. Arnoldia 7(9):
5356.
Aesculus
223
SimaroubaceaeQuassia family
224
Flowering and Fruiting. The tree is mainly dioecious, with some monoecious individuals (Dirr 1990; Miller
1990). Flowers are usually unisexual, but perfect flowers do
occur in some individuals (Feret 1973). Flowering has been
observed in seedlings 6 weeks after germination (Feret
1973).
Commercial seed consists of the 1-celled, 1-seeded,
oblong, thin, spirally twisted samaras. These samaras, with
seeds near the middle, are 8 to 12 mm wide and 33 to 48
mm long (Feret and others 1974) and light reddish brown in
color (figure 1). Flowering occurs from mid-April to July
(Little 1974). Seeds ripen in large panicles in September to
October of the same season and are dispersed from October
to the following spring (Illick and Brouse 1926). Ailanthus
is a prolific seeder: 15- to 20-year-old trees bear considerable quantities. Seeds have no endosperm (figure 2).
Collection of fruits; extraction and storage of seeds.
Ailanthus seeds have been found in soil seedbanks in stands
with no individuals present in the overstory. This suggests
that seeds may be stored in the soil for some period of time
after parent trees have disappeared from a site (Dobberpuhl
1980).
Figure 1Ailanthus altissima, ailanthus:
samara.
longitudinal
References
Alam MM, Anis M. 1987. Ethno-medicinal uses of plants growing in the
Bulandshahr district of northern India. Journal of Ethnopharmacology
19(1): 8588.
Albenskii AV, Nikitin PD, eds. 1956. Agrolesomeliortsiya, 3rd ed. Moscow:
Gosundarstvennoye Izdatelstvo Selskohozyaystvennoy Litrtsyury
[Handbook of afforestation and soil amelioration.Transl.TT 66-51098.
1967. Springfield,VA: USDC CFSTI. 516 p.].
Alden HA. 1995. Hardwoods of North America. Gen.Tech. Rep. FPL-83.
Madison, WI: USDA Forest Service, Forest Products Laboratory. 136 p.
Beniwal BS, Singh NB. 1990. Genetic improvement of forest trees in
Arunachal Pradesh. Indian Forester 116(1): 310.
Bicknell SH, Smith WH. 1975. Influence of soil salt, at levels characteristic
of some roadside environments, on the germination of certain tree
seeds. Plant and Soil 43(3): 719722.
Bordeau PE, Laverick ML. 1958. Tolerance and photosynthetic adaptability
to light intensity in white pine, red pine, hemlock, and ailanthus seedlings.
Forest Science 4(3): 196207.
Dirr MA. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture propagation and uses. Champaign, IL:
Stipes Publishing Co. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant
propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Dobberpuhl J. 1980. Seed banks of forest soils in east Tennessee
[MS thesis]. Knoxville: University of Tennessee. 219 p.
Feret PP. 1973. Early flowering in Ailanthus. Forest Science 19: 237239.
Ailanthus
225
226
Little S. 1974. Ailanthus altissima (Mill.) Swingle, ailanthus. In: Schopmeyer CS,
tech. coord. Seeds of woody plants in the United States. Agric. Handbk.
450. Washington, DC: USDA Forest Service: 201202.
Miller JH. 1990. Ailanthus altissima (Mill.) Swingle, ailanthus. In: Burns RM,
Honkala BA, tech. coords. Silvics of North America.Volume 2,
Hardwoods. Agric. Handbk. 654. Washington, DC: USDA Forest Service:
101104.
Rai SN. 1985. Notes on nursery and regeneration technique of some
species occurring in southern tropical evergreen and semi-evergreen
forests of Karnataka (India): 2. Indian Forester 111(8): 644657.
Ramakrishnan HB, Jacqueline AS,Vinaya Rai RS. 1990. Studies on the
ripeness index and presowing seed treatment in Ailanthus excelsa Roxb.
Seed Science & Techology 18(3): 491498.
Shumilina ZK. 1949. Podgotovka posevu semyan drevesnykh I kustarnikovykh porod.Vsesoyuznyy Nauchno-Issledovatelskiy Institut
Agrolesomelioratsii Goslesbumizdat Moscow [Preparation of tree and
shrub seed for sowing.Transl.TT 67-51300. 1967. Springfield,VA: USDC
CFSTI. 36 p.].
Van Dersal WR. 1938. Native woody plants of the United States: their erosion-control and wildlife values. Misc. Pub. 303. Washington, DC: USDA.
362 p.
FabaceaePea family
Albizia Durazz.
albizia
John A. Parrotta, Herbert L.Wick, and Gerald A.Walters
Dr. Parrotta is a research program leader at the USDA Forest Services Research and Development
National Office, Arlington,Virginia; Drs.Wick and Walters retired from the
USDA Forest Services Pacific Southwest Forest and Range Experiment Station
Common
name(s)
Occurrence
Native
US
Growth habit
silktree, albizia,
mimosa tree,
powder-puff tree
siris,
womans-tongue
Iran to Japan
Southern US
Deciduous
ornamental
Pakistan to Burma
Puerto Rico
& Hawaii
white siris,
tall albizia
India to Melanesia
& Hawaii
Puerto Rico
Deciduous
forest tree,
ornamental
Deciduous
forest tree
raintree, saman,
monkey-pod
S Florida &
Hawaii
Evergreen tree
(deciduous in
Hawaii)
Albizia
227
Species
Location
Flowering
Fruit ripening
Seed dispersal
A. julibrissin
A. lebbeck
A. procera
A. saman
S US
Puerto Rico
Puerto Rico
JuneAug
AprSept
AugSept
Springfall
SeptNov
All year
JanJune
Fallspring
SeptNov
All year
All year
All year
Sources: Little and Wadsworth (1964), Rock (1920),Wick and Walters (1974).
228
although a small sample of seeds kept in loosely corked bottles in a laboratory for almost 5 years had a germination rate
of almost 90% (Wick and Walters 1974).
Germination. Germination of albizia seeds is slow
because of their impermeable seedcoats (figure 2).
Dormancy can be broken either by mechanical scarification,
sulfuric acid scarification, or soaking in water (Francis and
Rodrguez 1993; Parrotta 1987a). The easiest, safest, and
usually most effective means for breaking dormancy in siris
and white siris is immersion of the seeds in boiling water for
1 to 3 minutes, soaking them in water at room temperature
for 24 hours, then sowing the seeds immediately (Parrotta
1987a). Germination rates for scarified seeds range from 50
to 99% and germination begins within 2 to 4 days after sowing (Francis and Rodrguez 1993; Parrotta 1987a&b).
Raintree seeds will often germinate without pretreatment,
but a 10-minute soak in sulfuric acid will increase the percentage and rate of germination (Walters and others 1974).
Germination as high as 92% has been reported for this
species (Neal 1965; Rock 1920). Germination in albizias is
epigeal (figure 3).
Nursery practice. Germination and seedling growth
of albizia is favored by shallow sowing, up to 2.5 cm (1 in)
longitudinal section
depth, in loose, moist soil under full sun (figure 3). Seedling
growth is rapid; siris and white siris seedlings reach
plantable size (20 cm in height) usually within 2 to 3
months after sowing under nursery conditions in Puerto
Rico (Parrotta 1987a). Raintree seeds are sown in March in
Hawaii for outplanting as 3/4 + 0 stock the following winter.
A sowing depth of 2.5 cm (1 in) and density of 160 to 215
seedlings/m2 (15 to 20/ft2) are recommended, with 75 to
85% shading of the beds (Walters and others 1974).
Plantations can be established by direct sowing (for siris and
white siris) or by using container seedlings (for all species).
Stumped seedlings or stem, branch, and root cuttings can
also be used to propagate siris and white siris (Parrotta
1987a).
Figure 3Albizia julibrissin, silktree: seedling development at 1, 3, 5, and 8 days after germination.
References
Seeds of Puerto Rican trees and shrubs: second installment. Res. Note SO374. New Orleans: USDA Forest Service, Southern Forest Experiment
Station. 5 p.
Little EL Jr, Wadsworth FH. 1964. Common trees of Puerto Rico and the
Virgin Islands. Agric. Handbk. 249. Washington, DC: USDA Forest
Service. 548 p.
Magini E,Tulstrup NP. 1955. Tree seed notes. For. Dev. Pap. 5. Rome: FAO.
354 p.
Neal MC. 1965. In gardens in Hawaii. Spec. Pub. 50. Honolulu: Bishop
Museum Press. 924 p.
Parrotta JA. 1987a. Albizia lebbek (L.) Benth, siris. Res. Note SO-ITF-SM-7.
New Orleans: USDA Forest Service, Southern Forest Experiment
Station. 5 p.
Parrotta JA. 1987b. Albizia procera (Roxb.) Benth., white siris, tall albizia.
Res. Note SO-ITF-SM-6. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 4 p.
Rock JF. 1920. The leguminous plants of Hawaii. Honolulu: Hawaii Sugar
Planters Association. 234 p.
Walters GA, Bonner FT, Petteys EQP. 1974. Pithecellobium Mart., blackbead.
In: Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service:
639640.
Wick HL, Walters GA. 1974. Albizia, albizzia. In: Schopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 203205.
Albizia
229
EuphorbiaceaeSpurge family
fruit (draw-
per fruit. The seeds are 2.5 to 3.5 cm long, and the shells are
hard, rough, and black (Dayan and Reaviles 1995; Little and
Skolmen 1989). Flowering and fruiting occurs intermittently
in Puerto Rico (Little and others 1974).
Collection, extraction, and storage. Fruits may be
collected from the ground after shedding or picked from the
trees. In the Philippines, it is common practice to let the
fruits decay for 3 to 5 days after collection and then remove
the husks by hand under running water. The seeds are then
dried in the sun for 3 or 4 days to a low moisture content;
there are about 116/kg (53/lb) (Dayan and Reaviles 1995).
Empty or deteriorated seeds can be removed by water flotation (Tamesis 1958; Eakle and Garcia 1977). There are no
long-term storage data on Indian-walnut, but the seeds are
apparently orthodox in storage characteristics. Dayan and
Reaviles (1995) reported that seeds dried to 10 to 12%
moisture can be successfully stored at room temperature for
7 months.
Germination. Indian-walnut germinates slowly,
apparently due to dormancy imposed by the hard seedcoat
(Eakle and Garcia 1977). Several pretreatments have been
References
Dayan MP, Reaviles RS. 1995. Seed technology manual of some reforestation species. Manila: National Forestation Development Office and
Ecosystem Research and Development Bureau. 60 p.
Eakle TW, Garcia AS. 1977. Hastening the germination of lumbang
[Aleurites moluccana (L.) Willd.] seeds. Sylvatrop 4: 291295.
Little EL Jr, Skolmen RG. 1989. Common forest trees of Hawaii (native and
introduced). Agric. Handbk. 679. Washington, DC: USDA Forest Service.
321 p.
Little EL Jr, Woodbury RO, Wadsworth FH. 1974. Trees of Puerto Rico and
the Virgin Islands.Volume 2. Agric. Handbk. 449. Washington, DC: USDA
Forest Service: 168170.
Tabat E. 1925. An efficient method of germinating lumbang. Makiling Echo
Philippines Bureau of Forestry, Division of Forest Investigation] 4(4):
1922.
Tamesis F. 1958. Lumbang culture in Tungao, Butuan City. Forest Leaves [July
1958]: 79, 40.
Aleurites
231
BetulaceaeBirch family
Alnus P. Mill.
alder
Constance A. Harrington, Leslie Chandler Brodie, Dean S. DeBell, and C. S. Schopmeyer
Dr. Harrington and Ms. Brodie are foresters on the silviculture research team at the USDA Forest Services
Pacific Northwest Research Station, Olympia,Washington; Dr. DeBell retired from the USDA Forest Services
Pacific Northwest Research Station; Dr. Schopmeyer (deceased) was the technical coordinator of the
previous manual
Growth habit and occurrence. Alderthe genus
Alnusincludes about 30 species of deciduous trees and
shrubs occurring in North America, Europe, and Asia and in
the Andes Mountains of Peru and Bolivia. Most alders are
tolerant of moist sites and thus are commonly found along
streams, rivers, and lakes and on poorly drained soils; in
addition, some species occur on steep slopes and at high elevations. The principal species found in North America are
listed in table 1. Many changes in the taxonomy of alder
have been made over the years; in this summary, species are
referred to by their currently accepted names although in
many cases the information was published originally under
the synonyms (and alternative common names) listed in
table 1.
Although some cultivated European alder is used commercially in the eastern United States, red alder is the largest
native species. It is also the most extensively utilized of the
native species. Management interest and research activity on
red alders have increased dramatically during the past 2
decades, and the resulting information accounts for the
majority of new information added to the previous summary
on alder seeds prepared by Schopmeyer (1974).
Alders are pioneer species favored by high light levels
and exposed mineral soils; in addition, their ability to fix
atmospheric nitrogen facilitates establishment on geologically young or disturbed sites with low levels of soil nitrogen
(Harrington and others 1994). Dense stands of naturally
regenerated red alders established quickly on mudflows
associated with the eruption of Mount St. Helens. The trees
grew rapidly and soon overtopped other pioneer species
such as poplars in the nitrogen-deficient soils (Heilman
1990). Sitka alder plays a similar role in primary succession
following deglaciation in Alaska.
Use. Seedlings have been planted successfully for
reforestation of coal mining spoil banks (Lowry and others
1962). Soil fertility is improved through fixation of atmospheric nitrogen by microorganisms in the root nodules
232
(Tarrant and Trappe 1971). Alders also have been planted for
wildlife food and cover (Liscinsky 1965) and for ornamental
use. European and red alders have been considered for use
in biomass plantings for energy (Gillespie and Pope 1994)
and are considered excellent firewood. In recent years, harvest and utilization of red alder has expanded greatly on the
Pacific Coast of North America, where the species is used
for paper products, pallets, plywood, paneling, furniture,
veneer, and cabinetry (Harrington 1984; Plank and Willits
1994). Red alder is also used as a fuel for smoking or curing
salmon and other seafood and its bark is used to make a red
or orange dye (Pojar and MacKinnon 1994). The soft, evengrained wood lacks odor or taste and has been traditionally
been used by native peoples, and more recently other woodworkers, to make bowls, eating utensils, and other items
(Pojar and MacKinnon 1994). In addition, alder exports
have grown from almost nothing in 1990 to more than
153,000 m3 (or 65 million board feet) of lumber annually
(Tarrant and others 1994). Several options exist for managing alder in both mixed (Miller and Murray 1978) and pure
stands (Tarrant and others 1983), and a summary of management principles and alternative strategies are available for
red alder (Hibbs and DeBell 1994).
Geographic races and hybrids. Considerable geographic variation exists among populations of red (Ager and
others 1993; Ager and Stettler 1994; Dang and others 1994;
Hamann and others 1988; Lester and DeBell 1989), speckled (Bosquet and others 1988), American green (Bosquet
and others 1987), and European alders (Funk 1990; Hall and
Maynard 1979). Disjunct populations of red alder have been
located in Idaho (Johnson 1968), and growth of such populations and those at the extremes of species range differs
markedly from that of most populations (Lester and DeBell
1989). Natural hybridization is common in alder, and zones
of introgression between some species can occur where
ranges overlap (Ager and Stettler 1994). Artificial hybridization has been conducted with numerous species, including
Common name(s)
Occurrence
European alder,
black alder,
European black alder
thinleaf alder,
mountain alder
but those of Nepal, red, and Sitka alders are larger, having
lengths of 12 to 24 mm (Carlson and Bryan 1959; Funk
1990; Harrington 1990; Krstinic 1994; Townsend and
Douglass 1994). They are produced in abundance before
trees reach 10 years of age in at least 2 species. European
alders can produce flowers by their second growing season,
and individual red alder trees are sexually mature at 3 or 4
years. Most dominant trees in a red alder stand will produce
seeds by age 6 to 8 years (Harrington and DeBell 1995;
Stettler 1978). Although the majority of seeds produced are
probably the result of outcrossing, both selfing and apomixis
occur in red alder (Stettler 1978). Seed production resulting
from selfing has been reported for European and mountain
alders; however, in many cases self-fertilization results in
Alnus
233
Common name(s)
Occurrence
Sitka alder
Siberian alder
Species
Location
Flowering
Fruit ripening
Seed dispersal
A. glutinosa
E US
S US & England
Europe
Canada, US
Idaho, Montana, Oregon
Hawaii
Oregon
Washington, Oregon
MarMay
(can start Jan)
MarMay
Mar-May
MarApr
Mar
Late winter
early spring
FebMay
Sept
FebApril
SeptNov
AugSept
OctFeb
Late Septearly Oct
AugOct
SeptDec
OctApr
SeptDec
Spring
AprJune
AprJune
Late Augmid-Oct
Mid Septearly Oct
SeptDec
A. incana
ssp. rugosa
ssp. tenuifolia
A. nepalensis
A. rhombifolia
A. rubra
A. serrulata
A. viridis
ssp. crispa
ssp. sinuata
E US, Alaska
Alaska,W Canada, & NW US
Sources: Densmore (1979), Fernald (1950), Funk (1990), Harrington (1990), Hitchcock and others (1964), Lewis (1985), McDermott (1953), McGee (1988), McVean
(1955), Schopmeyer (1974),White (1981).
* Flowering occurs during the period when leaves unfold.
Table 3Alnus, alder: growth habit, height, seed-bearing age, and seedcrop frequency
Species
A. glutinosa
A. incana
ssp. rugosa
ssp. tenuifolia
A. nepalensis (Hawaii)
A. rhombifolia
A. rubra
A. serrulata
A. viridis
ssp. crispa
ssp. sinuata
Growth habit
Height at
maturity (m)
Year first
cultivated
Minimum
seed-bearing
age (yrs)
Years
between large
seedcrops
Tree
Tree
Tree or shrub
Tree or shrub
Tree
Tree
Tree
Tree or shrub
to 35
to 20
to 8
19
1530
2025
1227
to 8
1866
1880
1916
1885
1884
1769
67
under 25
10
34
14
35
Shrub
Tree or shrub
to 3
to 12
1782
1903
Sources: Carlson and Bryan (1959), Fernald (1950), Funk (1990), Harrington (1990), Sargent (1965), Schopmeyer (1974).
235
longitudinal section
4293
70
2
1
1,277
998
2,816
2,200
6941,860
1,5304,101
71
70
2
5
650
776
1,430
1,712
613687
3501,400
1,3491,511
1,8432,700
1.0
1.3
0. 11.1
1.3
1.3
0.11.4
5
13
1.415
18
7
7
23
9
9
5
13
1.415
673
1,485
1.0
0.8
1.3
1.0
7
13
11
6
14
8
7
13
3060
299
660
3.7
4.8
51
123
668
1,470
437900
9611,980
810
1623
2130
810
39
7
86
321
352
706
774
257401
289639
565882
6351,406
1.2
1.5
Sources: Hibbs and Ager (1989), Liscinsky (1965), Mirov and Kraebel (1939), Schopmeyer (1974), Niemic and others (1995), USDA data on file at Olympia Forestry Sciences Laboratory.
* Yield data were determined on clusters of strobiles including stems.
Species
A. glutinosa
Pennsylvania
Europe
A. incana
Europe
A. incana
ssp. rugosa
ssp. tenuifolia
(fresh)*
(air dry)*
A. rhombifolia
(fresh)*
(air dry)*
A. rubra
A. viridis
ssp. crispa
ssp. sinuata
Soundness
Samples
(%)
Cleaned seeds (x1,000)/wt of strobiles
Range
Average
/kg
/lb
/kg
/lb
Cleaned seeds/
vol of strobiles
kg/hl
lb/bu
Cleaned seeds/
wt of strobiles
kg/100 kg lb/100 bu
Seed wt/
vol of strobiles
kg/hl
lb/bu
237
238
0
0
180
180+3
0
0
060
0
14
28
0
14
28
||
60
14
30
0
0
180
180+3
0
20
20
20
16
20
20
20
20
5
5
5
23
20
25
25
25
25
25
25
25
25
21
21
3040
21
30
7
28
28
28
28
56
56
56
10
26
21
21
21
21
21
21
21
21
30
28
28
5
59
56
18
21
42
49
0
17
54
27
21
12
25
38
21
9
27
35
12
7
14
7
7
7
7
7
21
21
21
2
13
5
5
5
5
5
5
5
5
Germination rate
Amount
(%)
Days
28
14
59
56
71
75
72
72
16
63
80
36
29
16
34
49
28
13
35
46
45
52
3
1
1
4
6
6
6
6
6
6
6
1
1
1
1
1
1
1
1
1
100
Germination
Avg (%)
Samples
Sources: ISTA (1993), McDermott (1953), Radwan and DeBell (1981), Schalin (1967), Schopmeyer (1974),Tanaka and others (1991), data on file at Olympia Forestry Sciences Laboratory.
Note: Day/night, 8 hrs/16 hours.
* Stratification, when used, was in a moist medium at 1 to 5 C.
180 days at 5 C, plus 3 days at 20 C,
No difference for 0, 30, or 60 days of stratification.
Light period was 10 housr/day at this temperature.
|| Seeds were stratified for an unspecified period.
A. glutinosa (Pennsylvania)
A. glutinosa (Finland)
fresh seed
dried seed
dried seed
dried seed
A. incana (Europe)
A. incana (Finland)
fresh seeds
dried seeds
dried seeds
dried seeds
A. i. ssp. tenuifolia
fresh seeds
A. rhombifolia
fresh seeds
A. rubra
dried seeds
fresh seeds
fresh seeds
fresh seeds
fresh seeds
fresh seeds
fresh seeds
A. serrulata
A. viridis
ssp. crispa
ssp. sinuata
Species
Cold
stratification
period* (days)
3040
65
87
87
87
87
87
87
45
45
45
45
43
43
43
43
Soundness
(%)
bareroot, 1+0 plug, and +0.5 (plug+transplant). Most nurseries sow in the spring when growing alder species (Ahrens
and others 1992; Schopmeyer 1974), but fall-sowing is mentioned by Heit (1968). Spring-sowing is sometimes delayed
until late spring to reduce seedling size. Sowing depths of 2
to 5 mm (.1 to .2 in) have been used for seeds of European
alder and red alder (Schopmeyer 1974). In California, seeds
of red alder have been mixed with 10 parts of vermiculite
and drilled 1 cm (.4 in) deep (Schopmeyer 1974). In
Oregon, seeds of red alder have also been sown on the soil
surface and covered with peat. Seeds of Nepal alder have
been mixed with sand and spread over the nursery beds. The
number of plantable seedlings obtained from 1 kg (2.2 lb) of
seed was 22,000 (10,000/lb) for European alder and 88,000
(40,000/lb) for hazel alder (Van Dersal 1938). Germination
is epigeal (figure 4).
Alder seedlings, particularly those of red alder, grow
rapidly and seedling densities should be lower than those
used for conifers. Seedlings grown at open-bed densities of
60 to 180 seedlings/m2 (5 to 15/ft2) or in large containers
result in much better outplanting performance than those
grown at greater densities or in small Styroblocks (Ahrens
1994). Inoculation of beds or container media with the
nodulating actinomycete Frankia can improve establishment
Figure 4Alnus glutinosa, European alder: seedling development at 1 and 7 days after germination (left); Alnus
incana ssp. tenuifolia, thinleaf alder: 2 older seedlings
(right).
Alnus
239
liter/10 m2 (or 0.5 pint/100 ft2) on bare soil and 0.38 liter
(0.7 pint) for the same area of sod (Liscinsky 1965). In
England, better stocking was obtained on a shallow blanket
bog with spot sowing of European alder than with broadcast
sowing. About 15 viable seeds were sown in each spot and
fertilized with about 60 g of rock phosphate (McVean 1959).
Seedling care. Information to guide lifting dates is
very limited, even for red alder (Ahrens 1994; Ahrens and
others 1992); current recommendations based on experience
in southwest Washington are to lift seedlings in January.
They are then stored at either +2 C or 2 C; the lower
temperature is recommended because it prevents budbreak
during storage (and possible Botrytis infection associated
with budbreak during storage) and reduces the tendency for
planted alders to break bud too soon after planting. Storage
in sealed bags will prevent desiccation. Because alder stems
are brittle and sensitive, seedlings must be handled carefully
during storage, transport, and outplanting to avoid damage
to stems, branches, and buds. At low elevations (< 300 m) in
western Washington, it has been recommended that
seedlings be planted between mid-March and mid-April.
The spring planting period should begin when the probability of severe frost is low and end before there is appreciable
soil drying (Dobkowski and others 1994).
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242
AsteraceaeAster family
mature seed.
2
0
5
0
10
4
15
26
20
21
25
18
30
10
Ambrosia
243
References
Bainbridge DA,Virginia RA. 1989. Restoration in the Colorado Desert:
species notes [Unpublished manuscript prepared for the California
Department of Transportation].
CALR [Center for Arid Lands Restoration]. 1995. Seed test data filed
19891995.Twentynine Palms, CA: USDI National Park Service, Joshua
Tree National Park. 24 p.
Graves WL. 1976. Revegetation of disturbed sites with native shrub species
in the western Mojave Desert. In: Test seeds of Mojave Desert shrubs.
Prog. Rep. BLM Contr. 53500-CT4-2(N): 1135.
Graves WL, Kay BL, Williams WA. 1975. Seed treatment of desert shrubs.
Agronomy Journal 67(NovDec): 773777.
Kay BL. 1975. Test of seeds of Mojave Desert shrubs: progress report. BLM
Contract 53500-CT4-2(N). 24 p.
Kay BL, Graves WL,Young JA. 1988. Long-term storage of desert shrub
seed. Mojave Reveg. Notes 23. Davis: University of California,
Department of Agronomy and Range Science.
244
Kay BL, Ross CM, Graves WL. 1977. Burrobush. Mohave Reveg. Notes 1.
Davis: University of California, Department of Agronomy and Range
Science.
Ruffner GA, Fedock DA, Carothers SW. 1985. Transplanting native Sonoran
Desert plants. Proceedings, 1st North American Riparian Conference,
Riparian Ecosystems and Their Management: Reconciling Conflicting
Uses; 1985 April 1618;Tuscon, AZ.
Went FW. 1979. Germination and seeding behavior of desert plants. In:
Arid land ecosystems: structure, functioning and management.
Cambridge, UK: Cambridge University Press: 477489.
Weiland PAT, Frolich EF, Wallace A. 1971. Vegetative propagation of woody
shrub species from the northern Mojave and southern Great Basin
Deserts. Madroo 21: 149152.
RosaceaeRose family
Amelanchier Medik.
serviceberry
Kenneth A. Brinkman and Terry F. Strong
Dr. Brinkman retired from the USDA Forest Services North Central Forest Experiment Station; Mr. Strong is
a research forester at the USDA Forest Services North Central Research Station, Rhinelander,Wisconsin
Growth habit, occurrence, and use. The serviceberriesthe genus Amelanchierinclude about 25 species
of small deciduous trees and shrubs native to North
America, Europe, and Asia. The distribution and chief uses
of 6 species are listed in table 1. Most species provide
browse and edible fruits for wildlife and many have attractive flowers. Saskatoon and common serviceberries have
been used to a limited extent for shelterbelt and wildlife
plantings and as a minor fruit crop, but other species also
should be considered for these and other environmental
uses. Native Americans have traditionally used most species
of serviceberry for food and medicine (Meeker and others
1993; Moerman 1986). Common and Saskatoon serviceberries are tolerant of temperatures to 60 C (Junttila and others 1983; Kaurin and others 1984; Lindstrom and Dirr
1989). Common serviceberry regenerates vegetatively and
by seed after clearcutting and burning (Scheiner and others
1988). Geographic races of Amelanchier have not been iden-
Common name(s)
Saskatoon serviceberry,
juneberry, western shadbush
Pacific serviceberry,
western serviceberry
common serviceberry,
downy serviceberry,
shadblow, serviceberry
Canadian serviceberry,
thicket shadblow, shadbush,
thicket serviceberry
Allegheny serviceberry,
juneberry, shadbush
roundleaf serviceberry,
roundleaf juneberry, shore
mtns. shadbush, Huron
serviceberry
Occurrence
Amelanchier
245
Location
Flowering
Fruit ripening
A. alnifolia
var. semiintegrifolia
Oregon (520 m)
Oregon (1,310 m)
A. arborea
A. canadensis
Carolinas
A. laevis
A. sanguinea
MayJune
Apr
May
May
MarJune
MarApril
May
MarJune
MayJune
JulyAug
Aug
Aug
JuneAug
MayJune
June
JuneAug
JulySept
Sources: Fernald (1950), Jones (1946), Mowat (1969), Plummer and others (1968), Radford and others (1964), Rehder (1940), St. Pierre and Steeves (1990),Van Dersal
(1938).
Height at
maturity (m)
Year first
cultivated
5
12
18
8
9
3
1826
1826
1623
1641
1870
1824
Color of
ripe fruit
Blue purple
Purplish black
Reddish purple
Nearly black (sweet)
Dark purple
Dark purple (sweet)
Sources: Fernald (1950), Jones (1946), Petrides (1958), Rehder (1940), Small (1933), Strausbaugh and Core (1953).
246
5081
80251
110178.6
2.8
2
2
1
Sources:
42
118
Oregon
Minnesota
A. alnifolia
var. semiintegrifolia
A. arborea
A. sanguinea
0.9
0.9
0.5
Average
/kg
/lb
/lb
Species
Seed wt/fruit wt
kg/45 kg
lb/100 lb
Fruit wt/vol
kg/hl
lb/bu
Place
collected
/kg
Range
Amelanchier
247
248
6174
120
60+
A. canadensis
A. laevis
Sources: Babb (1959), Brinkman (1974), Hilton and others (1965), McKeever (1938), Mclean (1967).
* Stratification was done in a moist medium at temperatures between 1 and 6 C.
In an additional test on excised embryos, germination was 82% (Brinkman 1974).
In an additional test on excised embryos, germination was 95% (Hilton and others 1965)
67
20
20
93
2
54
20
30
16
90120
A. alnifolia var.
semiintegrifolia
A. arborea
Sand or
sand & peat
Filter paper
2
10
31
20
30
2
10
70
62
50
30
70
20
21
30
21
Sand
Sand or
blotters
Kimpack
16
0
180+
120
98
3090
Medium
Species
A. alnifolia
Purity
(%)
Germination percentage
Avg (%)
Samples
Days
Germination rate
Amount
(%)
Days
Temp (C )
Day
Night
Daily
light
(hrs)
Cold
stratification*
(days)
Table 5Amelanchier, serviceberry: cold stratification period, germination test conditions, and results
References
Acharya SN, Chu SB, Hermesh R. 1989. Effects of population, environment
and their interaction on Saskatoon berry. Amelanchier alnifolia
Nutt seed germination. Canadian Journal of Plant Science 69: 277284.
Babb MF. 1959. Propagation of woody plants by seed. Alaska Agricultural
Experiment Station Bulletin 26: 112.
Bailey LH. 1935. The nursery manual. New York: Macmillan. 456 p.
Brinkman, KA. 1974. Amelanchier Med., serviceberry. In: Schopmeyer CS,
tech. coord. Seeds of woody plants in the United States. Agric. Handbk.
450. Washington, DC: USDA Forest Service: 212215.
Campbell CS, Greene CW, Dickinson TA. 1991. Reproductive biology in
subfam. Maloideae (Rosaceae). Systematic Botany 16(2): 333349.
Crocker W, Barton LV. 1931. After-ripening, germination, and storage of
certain rosaceous seeds. Boyce Thompson Institute Contributions 3:
385404.
Cruise JE. 1964. Studies of natural hybrids in Amelanchier. Canadian Journal
of Botany 42: 651633.
Fernald ML. 1950. Grays manual of botany. New York: American Book Co.
1632 p.
Flessner TR, Darris DC, Lambert SM. 1992. Seed source evaluation of four
native riparian shrubs for streambank rehabilitation in the Pacific
Northwest. In: Proceedings, Symposium on Ecology and Management of
Riparian Shrub Communities: 1991 May 2931; Sun Valley, ID. Gen.Tech.
Rep. INT-289. Ogden, UT: USDA Forest Service, Intermountain Forest
and Range Experiment Station: 155162.
Gorchov DL. 1985. Fruit ripening asynchrony is related to variable seed
number in Amelanchier and Vaccinium. American Journal of Botany
72(12): 19391943.
Heit CE. 1967. Fall planting of fruit and hardwood seeds. American
Nurseryman 126(4): 1213, 8590.
Heit CE. 1968. Thirty-five years testing of tree and shrub seed. Journal of
Forestry 66: 632634.
Hilton RJ, Joswal AS,Teskey BJ, Barabas B. 1965. Rest period studies on
seeds of Amelanchier, Prunus, and Sorbus. Canadian Journal of Plant
Science 45(1): 7985.
Jones GN. 1946. American species of Amelanchier. Illinois Biological
Monographs 20(2): 1126.
Junttila O, Stushnoff C, Gusta LV. 1983. Dehardening in flower buds of
saskatoon-berry, Amelanchier alnifolia, in relation to temperature,
moisture content, and spring bud development. Canadian Journal of
Botany 61(1): 164170.
Kaurin A, Stushnoff C, Junttila. 1984. Cold acclimation and dormancy of
Amelanchier alnifolia. Journal of the American Society of Horticultural
Science 109(2): 160163.
Lindstrom OM, Dirr MA. 1989. Acclimation and low-temperature tolerance
of eight woody taxa. HortScience 24(5): 818820.
McKeever DG. 1938. The effect of various methods of treatment on germination of seeds of some plants valuable for game and erosion
purposes [MS thesis]. Moscow, ID: University of Idaho School of
Forestry. 128 p.
McLean A. 1967. Germination of forest-range species from southern British
Amelanchier
249
FabaceaePea family
Amorpha L.
amorpha, indigobush
John C. Zasada and David Martineau
Dr. Zasada retired from the USDA Forest Services North Central Research Station; Mr. Martineau is the
plant division manager at the Prairie Nursery,Westfield,Wisconsin
from more southern seed sources grow more rapidly and are
taller than those from North Dakota sources, but they are
also more susceptible to winter damage (Lincoln Oakes
Nurseries 1996).
Leadplant and indigobush are reported to hybridize,
although hybrids are believed rare (Wilbur 1975). The
hybrid has the greatest affinity with leadplant and differs in
having a taller growth form as well as in several morphological traits (Wilbur 1975).
The growth form and stature of leadplant results from its
tendency to die-back to varying degrees each year. Regrowth
from basal stem and root collar buds maintains the above
ground stems. Under some conditions, stems will be relatively longer-lived and attain heights of 1.5 to 2 m (table 2).
Indigobush is taller than leadplant and its stem longevity is
like that of a true shrub.
Leadplant is palatable to domestic livestock and under
heavy grazing tends to disappear (Voigt and Mohlenbrock
nd); however its palatability for whitetail deer (Odocoileus
virginiana) was rated as low in a study in the Black Hills
(Rosario 1988). A primary use, at present, is for landscaping, where low-maintenance, drought-resistant plants are
desirable, and in restoration and reclamation projects
(Brown and others 1983; Cox and Klett 1984; Dirr 1990;
Common name(s)
Occurrence
A. californica Nutt.
A. canescens Pursh
A. fruticosa L.
A. nana Nutt.
A. microphylla Pursh
Sources: Brinkman (1974), Glad and Halsey (1993), Hickman (1993), Niering and Olmstead (1979), Rosario (1988),Voight and Mohlenbrock (nd),Wilbur (1975).
250
Height at
maturity (m)
Year first
cultivated
13
1218
13
1883
1724
1811
Sources: Brinkman (1974), Dirr (1990) Niering and Olmstead (1979), Rehder
(1940), Rosario (1988), Smith and Smith (1980),Vines (1960),Wilbur (1975).
Flowering
Fruit ripening
Seed dispersal
A. californica
A. canescens
A. fruticosa
A. nana
MayJuly
Junelate July
MayJune
MayJuly
JulySept
AugSept
Aug
July
AugSept
SeptOct
SeptOct
July
Sources: Brinkman (1974), Fernald (1950), Lincoln Oakes Nurseries (1996), Mirov and Kraebel (1939), Rehder (1940), Smith and Smith (1980),Van Dersal (1938).
Amorpha
251
Species
A. californica
A. canescens
A. fruticosa*
A. nana
/kg
194233
81205
88106
3793
211
114
133
96
52
60
Sources: Brinkman (1974), Lincoln Oakes Nurseries (1996), Prairie Nursery (1996), Salac and others (1978).
* One hundred pounds of dried fruit will produce about 60 pounds of clean seeds (Swingle 1939).
1966
272296
7282
84
624
170
38
284
77
Figure 3Amorpha canescens, leadplant: seedling development at 1, 2, 8, 20, and 52 days after germination.
Duration (days)
% Germination
30/20
30/20
30/20
5
1540
1540
3040
42
28
63
70
Sources: Blake (1935), Brinkman (1974), Christiansen (1967), Hutton and Porter (1937), Kraebel (1939), Lincoln Oakes Nurseries (1996), Martineau (1996), Pammel and
King (1928), Swingle (1939),Van Dersal (1938).
Note: Temperature is day/night regimen, photoperiod is 8 hours, based on Brinkman (1974).
* Germination of leadplant (Amorpha canescens) takes about 2 weeks when sown in nurserybeds in the spring (Lincoln Oakes Nurseries 1996; Martineau 1996).
Amorpha
253
References
Bailey LH. 1939. The standard cyclopedia of horticulture. New York:
Macmillan. 3639 p.
Blake AK. 1935. Viability and germination of seeds and early life history of
prairie plants. Ecology Monograph 5: 405460.
Brinkman KA. 1974. Amorpha, amorpha, false indigo. In: Schopmeyer CS,
tech. coord. Seeds of woody plants in the United States. Agric. Handbk
450. Washington, DC: USDA Forest Service: 216219.
Brown JE, Maddox JB, Splittstoesser WE. 1983. Performance of trees,
shrubs, and forbs seeded directly in the fall on minespoil and silt loam
soil. Journal of Environmental Quality 12: 523525.
Christiansen PA. 1967. Establishment of prairie species in Iowa by seeding
and transplanting [PhD thesis]. Ames: Iowa State University. 119 p.
Cox RA, Klett JE. 1984. Seed germination requirements of native Colorado
plants for use in the landscape. Plant Propagator 30(2): 610.
Curtis JT. 1959. The vegetation of Wisconsin: an ordination of plant communities. Madison: University of Wisconsin. 657 p.
Dirr MA. 1990. Manual of woody landscape plants: their identification,
ornamental characteristics, culture, propagation, and uses. Champaign IL:
Stipes Publishing. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Fernald ML. 1950. Grays manual of botany. New York: American Book Co.
1632 p.
Glad JB, Halse RR. 1993. Invasion of Amorpha fruticosa L. (Leguminosae)
along the Columbia and Snake Rivers in Oregon and Washington.
Madroo 40(1): 6263.
Henderson RA. 1995. Plant species composition of Wisconsin prairies.
Tech. Bull. 188. Madison: Wisconsin Department of Natural Resources.
58 p.
Hickman JC, ed. 1993. The Jepson manual: higher plants of California.
Berkeley: University of California Press. 1400 p.
Hutton ME, Porter RH. 1937. Seed impermeability and viability of native
and introduced species of Leguminoseae. Iowa State College Journal of
Science 12: 524.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing: rules 1993. Seed Science and Technology 21 (Suppl.): 159.
Johnson RG, Anderson RC. 1985. The seed bank of a tallgrass prairie in
Illinois. American Midland Naturalist 115(1): 123130.
Lee DK, Kim GT. 1986. Effects of artificial acid rain on seed germination
and seedling growth of several woody species. Seoul National University,
College of Agricultural Research, Bulletin of Kwanak Arboretum 7:
1521.
Lincoln Oakes Nurseries. 1996. Personal communication. Bismark, ND.
Martineau D. 1996. Unpublished data. Westfield, WI: Prairie Nursery.
Meeker JE, Elias JE, Heim JA. 1993. Plants used by the Great Lakes Ojibwa.
Odanah, WI: Great Lakes Indian Fish and Wildlife Commission. 440 p.
254
Mirov NT, Kraebel CJ. 1939. Collecting and handling seeds of wild plants.
For. Pub. 5. Washington, DC: USDA Forest Service, Civilian Conservation
Corps. 42 p.
Niering WA, Olmstead NC. 1979. The Audubon Society field guide to
North American wildflowers: eastern region. New York: Knopf. 887 p.
Pammel LH, King CM. 1928. Further studies of the germination and juvenile forms of some trees and woody shrubs, 1927. Proceedings of the
Iowa Academy of Science 35: 169183.
Prairie Nursery. 1996. Wildflowers and native grasses: catalog and growing
guide. Westfield, WI: Prairie Nursery. 47 p.
Rehder A. 1940. Manual of cultivated trees and shrubs hardy in North
America. New York: Macmillan. 996 p.
Richards MS, Landers RQ. 1973. Responses of species in Kalsow Prairie,
Iowa to an April fire. Proceedings of the Iowa Academy of Science
80(4): 159161.
Rock HW. 1981. Prairie propagation handbook. 6th ed. Hales Corner, WI:
Wehr Nature Center. 75 p.
Rogers CE, Garrison JC. 1975. Seed destruction in indigo amorpha by a
seed beetle. Journal of Range Management 28(3): 241242.
Rosario LC. 1988. Amorpha canescens. In: Fischer WC, comp.The Fire
Effects Information System [database]. Missoula, MT: USDA Forest
Service, Intermountain [now Rocky Mountain] Research Station,
Intermountain Fire Sciences Laboratory. 12 p.
Salac SS, Jensen PN, Dickerson JA, Gray RW Jr. 1978. Wildflowers for
Nebraska landscapes. Misc. Pub. 35. Lincoln: University of Nebraska,
Department of Horticulture. 27 p.
Smith JR, Smith BS. 1980. The prairie garden. Madison: University of
Wisconsin Press. 52 p.
Swingle CF. 1939. Seed propagation of trees, shrubs, and forbs for conservation planting. SCS-TP-27. Washington, DC: USDA Soil Conservation
Service. 198 p.
Van Dersal WR. 1938. Native woody plants of the United States: their erosion control and wildlife values. Misc. Pub. 303. Washington, DC: USDA.
62 p.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
Voigt JW, Mohlenbrock RH. [nd]. Prairie plants of Illinois. Springfield: Illinois
Department of Conservation, Division of Forestry. 272 p.
Weber GP, Wiesner LE. 1980. Tetrazolium testing procedures for native
shrubs and forbs. Journal of Seed Technology 5(2): 2334.
Wilbur RL. 1975. A revision of the North American genus Amorpha
(Leguminosae-Psoraleae). Rhodora 77 (811): 337409.
AraliaceaeGinseng family
Aralia L.
aralia
John C. Zasada and Barton M. Blum
Dr. Zasada retired from the USDA Forest Services North Central Research Station; Dr. Blum retired from
the USDA Forest Services Northeastern Forest Experiment Station
Scientific name
Common name(s)
Occurrence
A. hispida Vent.
bristly aralia,
wild-alder,
bristly sarsaparilla,
dwarf-elder
wild sarsaparilla,
small spikenard
Newfoundland to North
Carolina & W to
Minnesota & Indiana
1788
Newfoundland to North
Carolina & W to Manitoba
& Missouri
Pennsylvania to Florida,W to
SW Iowa & W Texas; range
extended by planting in
Massachusetts, Oregon,
Washington, & W Europe
Quebec to Manitoba, Great
Lakes region, New England,
& SE US
1731
A. nudicaulis L.
A. spinosa L.
devils-walkingstick,
angelica-tree,
Hercules-club,
prickly-ash
A. racemosa L.
spikenard, petty
morrel, life-of-man
1688
Growth
habit
Height at
maturity (m)
Subshrub or
perennial herb
0.30.9
Subshrub or
perennial herb
0.20.4
Tree
7.79.2
Subshrub or
perennial herb
0.53.0
Aralia
255
Figure 1Aralia nudicaulis, wild sarsaparilla: male inflorescence with 3 umbels, stamens just beginning to appear;
the larger vertical stem in the background is the leafbearing vegetative shoot.
256
nutlets
Flowering Dates
A. hispida
A. nudicaulis
A. spinosa
JuneJuly
MayJune
JulyAug
A
A
S
/kg
207,740218,790
185,640245,310
232,050346,970
Average
/lb
94,00099,000
84,000111,000
105,000157,000
/kg
/lb
Samples
203,320
218,300
288,850
92,000
99,000
131,000
2
3
2
Aralia
257
References
Barrett SCH, Helenrum K. 1981. Floral sex ratios and life history in Aralia
nudicaulis (Araliaceae). Evolution 35: 752762.
Barrett SCH,Thomson JD. 1982. Spatial pattern, floral sex ratios, and
fecundity in dioecious Aralia nudicaulis (Araliaceae). Canadian Journal of
Botany 60: 16621670.
Bawa KS, Keegan CR,Voss RH. 1982. Sexual dimorphism in Aralia nudicaulis
L. (Araliaceae). Evolution 36(2): 371378.
Blum BM. 1974. Aralia L., aralia. In: Schopmeyer CS, tech. coord. Seeds of
woody plants in the United States. Agric. Handbk. 450. Washington, DC:
USDA Forest Service: 220222.
Braun LE. 1961. The woody plants of Ohio. Columbus: Ohio State
University Press. 362 p.
Cheke AS, Nanakorn W, Yankoses C. 1979. Dormancy and dispersal of
seeds of secondary forest species under the canopy of a primary tropical rain forest in northern Thailand. Biotropica 11(2): 8895.
Crocker W. 1948. Growth of plants. New York: Reinhold Publishing. 90 p.
Curtis JT. 1959. The vegetation of Wisconsin. Madison: University of
Wisconsin Press. 657 p.
Dirr MA. 1990. Manual of woody landscape plants: their identification,
ornamental characteristics, culture, propagation and uses. Champaign, IL:
Stipes Publishing. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Edwards J. 1984. Spatial pattern and clone structure of the perennial herb
Aralia nudicaulis. Bulletin of the Torrey Botanical Club 111: 2833.
Graber RE,Thompson DF. 1978. Seeds in the organic layers and soil of
four beechbirchmaple stands. Res. Pap. NE-401. Upper Darby, PA:
USDA Forest Service, Northeastern Forest Experiment Station. 8 p.
Fernald, ML. 1950. Grays manual of botany. New York: American Book Co.
1632 p.
Harrar ES, George J. 1962. Guide to southern trees. New York: Dover
Publications. 709 p.
Hartmann HT, Davies DE, Davies FT Jr. 1990. Plant propagation: principles
and practice. 5th ed. Englewood Cliffs, NJ: Prentice Hall. 647 p.
Heit CE. 1967a. Propagation from seed: 11. Storage of deciduous tree and
shrub seeds. American Nurseryman 126(10): 1213, 8694.
Heit CE. 1967b. Propagation from seed: 6. Hardseedednessa critical
factor. American Nurseryman 125(10): 1012, 8896.
Heit CE. 1968. Propagation from seed: 15. Fall planting of shrub seeds for
successful seedling production. American Nurseryman 128(4): 810,
7080.
258
AraucariaceaeAraucaria family
Araucaria Juss.
araucaria
Gerald A.Walters and John K. Francis
Dr.Walters retired from the USDA Forest Services Pacific Southwest Forest and Range Experiment Station;
Dr. Francis retired from the USDA Forest Services International Institute of Tropical Forestry
Occurrence
US
Maximum
height (m)
Common name(s)
Native
Brazil, Argentina,
& Paraguay
Chile &
Argentina
Hawaii &
Puerto Rico
California, Oregon,
& Washington
Australia
California, Florida,
Hawaii, & Puerto Rico
Hawaii, Florida,
& Puerto Rico
California, Hawaii,
& Puerto Rico
California, Florida,
Hawaii, & Puerto Rico
Hawaii &
Puerto Rico
New Caledonia
New Guinea
& Australia
Norfolk Island
New Guinea
36
50
43
60
60
60
80
Araucaria
259
260
Table 2Araucaria, araucaria: phenology of flowering, seed development and dispersal, and seedcrop intervals
Species
Flowering
Seed ripening
Seed dispersal
A. angustifolia
A. bidwillii
A. columnaris
A. cunninghamii
Early-flowering races
Late-flowering races
A. heterophylla
SeptOct
DecJan
AprMay
JanFeb
DecFeb
MayAug
JanFeb
DecFeb
1
12
34
DecJan
AprMay
Sept
Dec
Apr
Dec
AprMay
45
34
/kg
6688
1,9802,640
3,3006,600
550620
2,0002,500
Average
/lb
/kg
3040
9001,200
1,5003,000
250280
9001,100
108
77
2,200
4,400
573
/lb
50
35
1,000
2,000
260
decreased significantly between 17 and 100 months of storage. However, after 100 months of storage, germination still
ranged from 25 to 44% (Shea and Armstrong 1978).
Tompsett (1984) found that seeds of monkey-puzzle-tree,
parana-pine, klinki-pine, and bunya-pine could not be safely
dried below 25 to 40% moisture content; seeds of cook-pine
and 2 other araucarias (A. nemorosa de Laubenfels and A.
scopulorum de Laubenfels) cannot be dried below 12%; and
seeds of hoop-pine could be dried to 2% without damage.
Seeds in the second 2 groups dried to moisture contents just
above the critical levels can be stored at 18 C and thus
appear to be orthodox. Parana-pine, monkey-puzzle-tree,
and bunya-pine seeds are classified as recalcitrant (Farrant
and others 1989; Ramos and others 1988). Plastic bags are
good containers (Ntima 1968). Seeds of hoop-pine can be
stored up to 8 years (Shea and Armstrong 1978).
Germination. No pregermination treatments are needed for araucaria seed (Ntima 1968; Walters 1974). Under
suitable moisture and temperature (21 to 30 C) conditions,
germination (which is cryptogeal in this genus) may begin
about 10 days after sowing. Germination is delayed by cooler temperatures, sometimes taking 50 days or more (Ntima
1968). Seed quality varies from year to year; if sufficient
pollen is available to the parent trees, seed quality is generally good (Walters 1974).
Twenty-nine and 45% of a large number of hoop-pine
and klinki-pine seeds germinated within 9 weeks in a germination test (Thong 1974). Klinki-pine seeds are pregerminated (incubated until the radicle begins to show) before sowing into containers. In a test with 3 replications of 1,200
seeds each, germination averaged 85% in 22 days. Of those
seeds not germinating, 54% were dead, 30% were rotten,
and 16% had not germinated yet. Survival of seedlings in
containers to outplanting size was 88%. Broadcasting seeds
on the surface of wet sawdust with a second shadecloth a
few centimeters above the bed gave better germination than
covering seeds with sawdust or germinating them without
the second shadecloth covering (Howcroft 1974). Tompsett
(1984) obtained 80 to 100% germination of 6 species tested
when seed moisture contents were optimal.
Nursery practice. Araucarias can be grown under
high shade or low shade. For both types of shade, seeds are
sown during spring. Norfolk-Island-pine seeds are placed on
a bed of sandsoilpeat mix to germinate with the pointed
end of the seed slightly embedded. About 70% of fresh
seedlots germinate in 4 to 12 days (Logsdon 1973). Seeds
Araucaria
261
References
Dallimore W, Jackson AB. 1954. A handbook of Coniferae and
Ginkgoaceae. London: Edward Arnold, Publisher. 686 p.
El-Lakany MH, Kamara AM, Badran OA, Attia YG. 1981. Seed pathology of
Araucaria spp.: 2. Fungal species associated with Araucaria heterophylla
seed. Australian Forestry Research 11(3/4): 275281.
Farrant PM, Pammenter NW, Berjak P. 1989. Germination-associated events
and the desiccation sensitivity of recalcitrant seeds: a study on three
unrelated species. Planta 178(2): 189198.
Francis JK. 1988. Araucariaceae in Puerto Rico.Turrialba 38(3): 202207.
Haines RJ, Nikles DG. 1987. The Araucaria araucana gene resource in Chile.
For. Occ. Pap. 1976/1. Rome: FAO: 26.
Howcroft NHS. 1974. Pregermination techniques for Araucaria hunsteinii.
Trop. For. Res. Note SR.27. Boroko, Papua New Guinea: Department of
Forests. 10 p.
Howcroft NHS. 1978a. Exploration and provenance seed collection in
Papua New Guinea 1976/1977. For. Occ. Pap. 1978/2. Rome: FAO: 511.
Howcroft NHS. 1978b. Data sheet on Araucaria hunsteinii K. Schumann. For.
Occ. Pap. 1978/2. Rome: FAO: 3137.
Kamara AM, El-Lakany MH, Badran OA, Attia YG. 1981. Seed pathology of
Araucaria spp.: 1. A survey of seed-borne fungi associated with four
Araucaria spp. Australian Forest Research 11(3/4): 269274.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan: 98.
Logsdon BB. 1973. Growing the Norfolk Island pine.Tree Planters Notes
24(1): 3336.
Menninger EA. 1964. Seaside plants of the world. New York: Hearthside
Press. 303 p.
Ntima OO. 1968. Fast growing timber trees of the lowland tropics: the
araucarias. Oxford: Commonwealth Forestry Institute. p. 139.
262
EricaceaeHeath family
made into bobbins and spools. This species was first cultivated in 1827 and has been planted occasionally as an ornamental tree in Europe and the United States (McMinn and
Maino 1959).
Flowering and fruiting. Flowers, which bloom from
March to June, are formed on a panicle 12 to 15 cm long.
The 8-mm flowers consist of 5 sepals fused at the base with
5 fused urn-shaped petals and 10 stamens. The anthers split
open when ripe, the awns are elongate, and the superior
ovary is rough and bumpy with 5 chambers (Hickman
1993). The fruit is a berry, also rough and bumpy, less than
12 mm in diameter (figure 1). The generic name derives
from arboise, a Celtic word for rough fruit (Roy 1974).
The thin-skinned berry has rather dry, mealy flesh and generally is 5-celled (figures 1 and 2). McDonald (1978) found
that, in northern California, the number of seeds per berry
ranged from 2 to 37, with an average of 20. The berries
ripen in September through November but often remain on
the trees through December. Fully ripe berries are bright red
Arbutus
263
264
References
EDA [Economic Development Administration]. 1968. The Hoopa Valley
Reservation hardwood study report. Washington, DC: USDC. 154 p.
Hickman JC, ed. 1993. The Jepson manual of higher plants of California.
Berkeley: University of California Press. 1400 p.
Koch M. 1973. Santa Cruz County: parade of the past. Fresno, CA:Valley
Publishers: 3343.
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
McDonald PM. 1978. Silviculture-ecology of three native California
hardwoods on high sites in north-central California [PhD dissertation].
Corvallis: Oregon State University, Department of Forest Science. 309 p.
McDonald PM. 1992. Estimating seed crops of conifer and hardwood
species. Canadian Journal of Forest Research 22: 832838.
McDonald PM,Tappeiner JC II. 1990. Arbutus menziesii Pursh, Pacific
madrone. In: Burns RM, Honkala BH, tech. coords. Silvics of North
America.Vol. 2, Hardwoods. Agric. Handbk. 654. Washington, DC: USDA
Forest Service: 124132.
McDonald PM, Minore D, Atzet T 1983. Southwestern OregonNorthern
California hardwoods. In: Burns RM, tech. comp. Silvicultural systems for
the major forest types of the United States. Agric. Handbk. 445.
Washington, DC: USDA Forest Service: 2932.
McMinn HE, Maino E. 1959. An illustrated manual of Pacific Coast trees.
Berkeley: University of California Press. 490 p.
Mirov NT, Kraebel CJ. 1939. Collecting and handling seeds of wild plants.
For. Pub. 5. Washington, DC: USDA Forest Service, Civilian Conservation
Corps. 5 p.
Arbutus
265
EricaceaeHeath family
Arctostaphylos Adans.
manzanita
Susan E. Meyer
Dr. Meyer is a research ecologist at the USDA Forest Services Rocky Mountain Station,
Shrub Sciences Laboratory, Provo, Utah
Growth habit, occurrence, and uses.
The shrub genus
Arctostaphylos, or manzanita, comprises about 50 species,
90% of which are endemic to California and adjacent areas
(Munz and Keck 1959). Three speciesgreenleaf manzanita, Mexican manzanita, and rosybract manzanitaare widely distributed in the southwestern United States and Mexico.
One speciesbearberry or kinnickinnickis circumboreal
in distribution (table 1). The manzanita habit varies from
mat-forming (bearberry) to nearly arborescent (bigberry
manzanita). About a quarter of the species have subterranean
burls that generate new sprouts both after fire and throughout the long life of the plant (Keeley 1992; Wells 1969). The
leaves of manzanitas are leathery, entire, and evergreen.
They are major components of chaparral and are also common understory species in montane coniferous forest types,
especially ponderosa (Pinus ponderosa Dougl. ex Laws.)
and Jeffrey (P. jeffreyi Grev. & Balf.) pines. They are most
Common name(s)
Habit
Habitat
Distribution
A. canescens Eastw.
hoary manzanita
Eastwood manzanita
A. glauca Lindl.
bigberry manzanita
Ponderosa pine
forest, chaparral
Ponderosa pine
forest, chaparral
Chaparral, joshua
tree woodland
N California to Oregon
A. glandulosa Eastw.
A. patula Greene
greenleaf manzanita
A. pungens Kunth
Mexican manzanita,
pointleaf manzanita
Shrubby,
without burl
Shrubby,
with burl
Shrubby or
treelike,
without burl
Shrubby,
with burl
Shrubby,
without burl
A. pringlei Parry
rosybract manzanita,
Pringle manzanita
Shrubby,
without burl
A. uva-ursi (L.)
Spreng.
bearberry,
kinnickinnick
Mat forming,
without burl
266
Ponderosa pine
forest
Ponderosa pine forest,
chaparral, pinyon-juniper
woodland
Ponderosa pine forest,
chaparral, mixed warm
desert shrubland
Coniferous forest
mostly at high elevation
California to Oregon
S California to Baja California
California to Oregon,
Arizona, & Colorado
S California, E to Utah &
Texas & S into Mexico
S California, S to Baja
California & E to
Arizona & SW Utah
Circumboreal, S to California,
New Mexico, Illinois, &
Georgia
Arctostaphylos
267
may be checked in the field by cutting the fruits transversely, preferably before the endocarp hardens (Berg 1974).
Kelly and Parker (1991) reported a mean set (percentage of
ovules forming seeds) of 62% for 14 California species,
with a range from 50 to 80%.
To clean manzanita seeds, the fruits should be soaked in
water, then macerated by hand or in a macerator to separate
the pulp from the stones. The pulp may be removed by
flotation, or the material may be dried, after which nutlets
may be separated from the dried pulp using screens or a fanning mill (Berg 1974). Seedlots may be cleaned to high
purity (Belcher 1985). Representative seed unit weights are
given in table 2. Seed unit weights are highly variable even
within a seedlot because a seed unit may be single or
multiple-seeded, depending on the degree of coalescence of
the nutlets.
Manzanita seeds form persistent seed banks and are
apparently long-lived under field conditions (Kelly and
Parker 1990). There is little information on longevity in
warehouse storage, but it is probable that seedlots would
maintain viability over periods of 10 years or more.
Germination and seed testing. In natural stands,
new seedlings (figure 3) of most species of manzanita grow
only after fire, and the seeds of these species are considered
to be refractory, that is, germinating only in response to firerelated environmental cues (Keeley 1991, 1995). But unlike
the refractory seeds of most chaparral shrubs, manzanita
seeds apparently do not become germinable through heat
shock (Kauffman and Martin 1991; Keeley 1987a). There is
evidence that charate leached from incompletely burned
wood can trigger germination in manzanita seeds, but the
maximum percentages attained using recently collected
seeds were not high (13% for Eastwood manzanita and 19%
for greenleaf manzanita (Keeley 1987a, 1991). It is probable
that, under field conditions, the seeds change in some way
following dispersal (perhaps through dry after-ripening at
Seed unit
/kg
Seeds/weight
/lb
Filled seeds
(%)
Sample
A. glandulosa
A. glandulosa
A. glauca
A. patula
A. patula
A. uva-ursi
12 seeded
13 seeded
Entire stone
Variable
1-seeded
1-seeded
66,15097,020
55,125
9901,760
36,69055,125
44,100
59,53590,405
30,00044,000
25,000
450800
18,00025,000
20,000
27,00041,000
58
83
85
2
2
5
1+
1
3+
Sources: Belcher (1985), Berg (1974), Keeley (1977, 1991), Keeley and Hayes (1976), Meyer (1997).
268
References
Belcher E. 1985. Handbook on seeds of browse-shrubs and forbs.Tech.
Pub. R8-8. Atlanta: USDA Forest Service, Southern Region. 246 p.
Berg AR. 1974. Arctostaphylos Adans., manzanita. In: Schopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 228231.
Carlson JR, Sharp WC. 1975. Germination of high elevation manzanitas.
Tree Planters Notes 26(3): 1011, 25.
Dirr MA. 1983. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation, and uses. Champaign, IL:
Stipes Publishing. 826 p.
Emery DE. 1988. Seed propagation of native California plants. Santa
Barbara, CA: Santa Barbara Botanic Garden. 107 p.
Fulton RE, Carpenter FL. 1979. Pollination, reproduction, and fire in
California Arctostaphylos. Oecologia 38: 147157.
Giersbach J. 1937. Germination and seedling production of Arctostaphylos
uva-ursi. Contributions to the Boyce Thompson Institute 9: 7178.
Kauffman JB, Martin RE. 1991. Factors influencing scarification and germination of three montane Sierra Nevada shrubs. Northwest Science 65:
180187.
Keeley JE. 1977. Seed production, seed populations in soil, and seedling
production after fire for two congeneric pairs of sprouting and
nonsprouting chaparral shrubs. Ecology 58: 820829.
Keeley JE. 1987a. Role of fire in seed germination of woody taxa in
California chaparral. Ecology 68: 434443.
Keeley JE. 1987b. Ten years of change in seed banks of the chaparral shrubs
Arctostaphylos glauca and A. glandulosa. American Midland Naturalist 117:
446448.
Keeley JE. 1991. Seed germination and life history syndromes in the
California chaparral. Botanical Review 57: 81116.
Keeley J. 1992. Recruitment of seedlings and vegetative sprouts in
unburned chaparral. Ecology 73: 11941208.
Arctostaphylos
269
270
Kelly VR, Parker VT. 1991. Percentage seed set, sprouting habit, and ploidy
level in Arctostaphylos (Ericaceae). Madroo 38 (4): 227232.
Meyer SE. 1997. Personal observation. Provo, UT: USDA Forest Service,
Rocky Mountain Research Station.
Munz PA, Keck DD. 1959. A California flora. Berkeley: University of
California Press. 1681 p.
Parker VT, Kelly VR. 1989. Seed banks in California chaparral and other
Mediterranean climate shrublands. In: Leck MA, Parker VT, Simpson RL,
eds. Ecology of soil seed banks. San Diego: Academic Press: 231253.
Wells PV. 1969. The relation between mode of reproduction and extent of
speciation in woody genera of the California chaparral. Evolution 23:
264267.
RosaceaeRose family
Aronia Medik
chokeberry
John D. Gill, Franz L. Pogge, and Franklin T. Bonner
Dr. Gill and Mr. Pogge retired from the USDA Forest Services Northeastern Forest Experiment Station;
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Growth habit, occurrence, and uses. The chokeberriesgenus Aroniadiscussed here are 2 closely related
species (red and black chokeberries) and 1 hybrid deciduous
shrub (purple chokeberry) (table 1). Black chokeberry is
small, only 0.5 to 1 m tall. Red chokeberry and purple
chokeberry are medium sized, 3 to 4 m tall. Red and black
chokeberries hybridize readily and may be difficult to distinguish. Red and purple chokeberries are practically identical
ecologically (Van Dersal 1938), and the only satisfactory
way to distinguish between them is by the color of their ripe
fruit. Both have pubescence on younger branches, leaf
stems, and lower leaf surfaces. In contrast, black chokeberry
is smooth or has only a few scattered hairs on these parts
(Gleason 1963). The combined ranges of these 3 include
most of the eastern United States and southern parts of adjacent Canadian provinces (table 1). All are moderately tolerant of shading and prefer moist soils, which usually are
acidic. The most likely habitats are bogs and swamps, low
woods, clearings, and damp pine barrens. However, each
Common name
Occurrence
Height (m)
red chokeberry
black chokeberry,
gueles noires
Newfoundland to Minnesota
& S to Tennessee & South
Carolina
0.51
purple chokeberry,
hybrid chokeberry
14
14
Aronia
271
Species
Location
Flowering
Fruit ripening
A. arbutifolia
Texas
West Virginia
North
South
North
West Virginia
MarApr
MarMay
AprJuly
MarJune
AprJuly
June
AprJuly
OctNov
SeptOct
SeptNov
Aug
AugOct
SeptOct
AugOct
A. melanocarpa
A. x prunifolia
Sources: Ammons (1975), Fernald (1950), McDonald (1960), Mahlstede and Maber (1957),Van Dersal (1938).
leaf and
Table 3Aronia, chokeberry: cold stratification periods, germination test conditions and results
Species
A. arbutifolia
A. melanocarpa
A. x prunifolia
Sources:
Cold stratiication
period (days)
Day
90
90120
60
20
30
20
30
30
30
Germinative capacity
Amount
(%)
Samples
94
22
96
4
4
2
References
Adams J. 1927. The germination of the seeds of some plants with fleshy
fruits. American Botanist 15(8): 415428.
Ammons N. 1975. Shrubs of West Virginia. Grantsville, WV: Seneca Books.
127 p.
Chadwick LC. 1935. Practices in propagation by seeds: stratification treatment for many species of woody plants described in fourth article of
series. American Nurseryman 62(12): 39.
Crocker W, Barton LV. 1931. After-ripening and storage of certain roseaceous seeds. Contributions of the Boyce Thompson Institute 3: 385404.
Dehgan B, Gooch M, Almira F, Kane M. 1989. Vegetative propagation of
Florida native plants: 3. Shrubs. Proceedings of the Florida State
Horticultural Society 102: 254266 [Seed Abstracts 1991; 14(4): 1155].
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seeds to tissue culture. Athens, GA:Varsity Press. 239 p.
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1632 p.
Gill JD, Pogge FL. 1974. Aronia Medik, chokeberry. In: Schopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 232234.
Gleason HA. 1963. The new Britton and Brown illustrated flora of the
Northeastern United States and adjacent Canada. 3 vol. New York:
Hafner Publishing.
Aronia
273
AsteraceaeAster family
Artemisia L.
sagebrush
Susan E. Meyer
Dr. Meyer is a research ecologist at the USDA Forest Services Rocky Mountain Research Station,
Shrub Sciences Laboratory, Provo, Utah
Growth habit, occurrence, and use. Sagebrush
Artemisia L.species are probably the most common
shrubs in western North America. Big sagebrush alone occupies an estimated 60 million ha as a landscape dominant or
codominant in the semiarid interior, and related species of
the subgenus Tridentatae are estimated to occupy an additional 50 million ha (Beetle 1960; McArthur and Stevens in
press). Sagebrush-dominated vegetation occurs mostly under
semiarid climatic regimes characterized by cold winters and
predominantly winter precipitation. The genus is circumboreal in distribution and consists of about 400 species of
mostly evergreen shrubs, subshrubs, and herbaceous perennials.
The 20 or so shrubby sagebrush species in the United
States differ widely in their growth form, ecology, distribution, and abundance (table 1). Big, black, silver, and low
sagebrushes are widely distributed, polymorphic species of
relatively broad ecological amplitude, whereas most of the
remaining species are either more geographically restricted
or more specialized in their habitat requirements. The subshrub fringed sagebrush, common and widespread in both
the Old and New Worlds, may be the most widely distributed sagebrush taxon. Sand sagebrush is an important
species on sandy soils on the Great Plains and in the
Southwest, whereas the summer-deciduous subshrub budsage is the principal sagebrush species of salt desert shrub
vegetation in the Great Basin.
Because of their status as regional dominants, sagebrush
speciesespecially those of the subgenus Tridentatae
have been the object of a great deal of study (McArthur and
Welch 1986). Many have long been regarded as undesirable
plants by the ranching industry because of their perceived
low palatability to livestock and propensity for increase
under conditions of abusive grazing. However, they provide
a principal source of browse on winter ranges for both wild
and domestic ungulates, and undoubtedly are central to the
habitat requirements of many other wildlife species.
274
Table 1Artemisia, sagebrush: distribution and ecology of principal shrubby species in the United States
Scentific name
Common names(s)
Distribution
Habitat
SUBGENUS TRIDENTATAE
A. arbuscula Nutt.
low sagebrush
Widely distributed,
mostly intermountain
SW deserts
A. bigelovii Gray
A. cana Pursh
Bigelow sagebrush,
rimrock sagebrush
silver sagebrush
A. nova A. Nels.
black sagebrush
A. pygmaea Gray
pygmy sagebrush
stiff sagebrush,
scabland sagebrush
big sagebrush
OTHER SUBGENERA
A. filifolia Torr.
A. frigida Willd.
A. spinescens D.C. Eat.
Picrothamnus desertorum
Nutt.
NW Great Plains, N
intermountain region
& N Sierras
Widely distributed,
mostly intermountain
Utah & adjacent parts
of Nevada & Colorado
Columbia Plateau,
E Washington & Oregon
Widely distributed,
W North America
See species
See species
See species
threetip sagebrush
Columbia Plateau E
into Wyoming
sand sagebrush,
old man sagebrush
fringed sagebrush
budsage
Flowering and fruiting. Most North American sagebrush species flower in late summer or autumn and ripen
fruit from September through December. Seeds of high-elevation populations generally ripen earlier than those of lowelevation populations. Budsage, which flowers in March or
April and sets seed in May or June before entering summer
dormancy, is a major exception. The tiny yellowish or
brownish flowers are wind-pollinated and are borne in
groups of about 2 to 70 (depending on species) in small
heads enclosed in overlapping bracts with thin, dry margins. The numerous heads are arranged in spikelike or open
panicles that occur terminally on the branches of currentseason growth. Each fertile floret within a head may develop into a small, 1-seeded fruit (achene) that lacks any special appendages for dispersal (figure 1). The pericarp of the
achene is papery and membranous, whereas the seedcoat of
the enclosed seed is firmer and somewhat shiny. The
endosperm is reduced to a membrane fused to the inner
wall of the seedcoat, whereas the embryo is well-developed
and fills the interior of the seed. Mucilaginous nerves on
Artemisia
275
276
Range
Species
/kg
/lb
/kg
/lb
A. arbuscula
A. bigelovii
A. cana
A. nova
A. pygmaea
A. rigida
A. tridentata
spp. tridentata
spp. vaseyana
spp. wyomingensis
A. tripartita
A. filifolia
A. frigida
A. spinescens
1.81
5.54
2.87
2.03
1.04
1.10
0.82
2.52
1.30
0.92
0.47
0.50
1.132.15
1.814.90
2.002.12
0.150.98
0.822.23
0.910.96
5.26*
4.30
4.72
4.87
3.20
10.0
3.06
2.38*
1.95
2.14
2.21
1.45
4.55
1.39
4.255.67*
4.234.36
4.005.42
2.253.70
1.932.58*
1.921.98
1.822.46
1.021.68
Sources: Belcher (1985), Deitschman (1974), McArthur and others 2004, Meyer (1990).
* Subspecies not distinguished.
Species
A. arbuscula
A. bigelovii
A. cana,
A. nova
A. tridentata
ssp. tridentata
ssp. vaseyatia
ssp. wyomingensis
A. filifolia
A. spinescens
Days to 50%
germination at 1 C (light)
Mean
Range
Lots #
100
100
100
92.3
81.5
21.2
100
75100
7588
357
38.2
56.0
47.6
38
5458
1780
1
1
2
5
94.6
85
98.4
100
92.7
18.6
12.2
13.4
72.6
84100
6494
94100
8798
046
024
246
5293
54.0
49.2
55.2
45.5
2795
1698
1898
3853
5
5
5
1
2
Sources: All data from Meyer (1990) except for A. tridentata lots stored 4 months (Meyer and others 1990).
* Expressed as percentage of viable seeds.
Artemisia
277
others 1959; Wilson 1982) probably stems from the fact that
sagebrush seeds have no need for protection from germination at summer temperature, as they almost never encounter
summer regimes. Budsage, a species with seeds that ripen in
early summer but do not germinate until the following early
spring, shows strong germination suppression at summer
temperatures (Meyer and Kitchen 1997).
Germination testing for sagebrush species is a relatively
straightforward process. We recommend a 21-day test at 15
or 20 C with light as the standard for big sagebrush and
black sagebrush, with a 2-week chill (stratification) for more
dormant lots (AOSA 1993; Meyer and others 1988a, 1988b).
Because many dormant sagebrush seeds will not germinate
in response to a short chilling, the viability of ungerminated
seeds should be evaluated with tetrazolium.
Tetrazolium staining also represents an alternative to the
germination test for evaluating the viability of sagebrush
seeds. The fruits are pierced with a needle through the center of the cotyledon region of the embryo (figure 2) and
immersed in buffered 1% tetrazolium chloride solution for 6
hours at 25 C. The pericarp and seedcoat are then slit with
a needle at the cotyledon end, and the embryos are squeezed
out. Embryos stained a uniform bright red may be classed as
viable.
The principal source of inconsistent results in sagebrush
seed testing comes from decisions made during the purity
evaluation. The inclusion of non-viable half-filled and aborted fruits in the pure seed fraction has little effect on the
value for percentage purity but can affect the viability per-
278
longitudinal
success, perhaps through re-inoculation with essential symbionts such as mycorrhizae (Monsen and Richardson 1984).
Fertilization per se usually favors herbaceous competitors
over the shrub seedlings and is not generally recommended.
Reports on seedling competitiveness in sagebrush are
somewhat contradictory. In the era of sagebrush control on
rangelands, managers often remarked on the ability of sagebrush to reestablish in perennial forage grass plantings
(Pechanec and others 1944). Follow-up moisture in the summer appears to facilitate shrub seedling survival in competition with perennial grasses. Success in mixed seedings may
be enhanced by separating the seeds spatially, for example,
References
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing
seeds. Journal of Seed Technology 16(3): 1113.
Bai Y, Romo JT. 1994. Germination of previously-buried seeds of fringed
sage (Artemisia frigida). Weed Science 42: 390397.
Beetle AA. 1960. A study of sagebrush: the section Tridentatae of
Artemisia. Wyoming Agriculture State Bulletin 368: 183.
Belcher E. 1985. Handbook on seeds of browse-shrubs and forbs.Tech.
Pub. R8-8. Atlanta: USDA Forest Service, Southern Region. 246 p.
Billings WD. 1990. Bromus tectorum, a biotic cause of ecosystem impoverishment in the Great Basin. In: Woodell GM, ed.The Earth in transition:
patterns and processes of biological impoverishment. Cambridge, UK:
Cambridge University Press: 301322.
DAntonio CM,Vitousek PM. 1992. Biological invasions by exotic grasses,
grass-fire cycle, and global change. Annual Review of Ecology and
Systematics 23: 6388.
Deitschman GH. 1974. Artemisia, sagebrush. In: Schopmeyer CS, tech
coord. Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 235237.
Fairchild JA. 1991. Plant community and shrub vigor responses to one-way
and two-way chaining of decadent big sagebrush on a critical mule deer
winter range in east central Utah [PhD dissertation]. Provo, UT:
Brigham Young University.
Hassan MA, West NE. 1986. Dynamics of soil seed pools in burned and
unburned sagebrush semi-deserts. Ecology 67: 269272.
Jacobsen TLC, Welch BL. 1987. Planting depth of Hobble Creek mountain
big sagebrush. Great Basin Naturalist 47: 497499.
Long LE. 1986. Container nursery production of Artemisia and Chrysothamnus species. In: McArthur ED, Welch BL, comps. Proceedings,
Symposium on the Biology of Artemisia and Chrysothamnus. 1984 July
913; Provo, UT. Gen.Tech. Rep. INT-200. Ogden, UT: USDA Forest
Service, Intermountain Forest and Range Experiment Station: 395396.
McArthur ED, Welch BL, eds. 1986. Proceedings, Symposium on the
Biology of Artemisia and Chrysothamnus. 1984 July 913; Provo, UT. Gen.
Tech. Rep. INT-200. Ogden, UT: USDA Forest Service, Intermountain
Forest and Range Experiment Station. 398 p.
McArthur ED, Blauer AC, Plummer AP, Stevens R. 1979. Characteristics
and hybridization of important Intermountain shrubs: 3. Sunflower family. Res. Pap. INT-220. Ogden, UT: USDA Forest Service, Intermountain
Forest and Range Experiment Station. 82 p.
McArthur ED, Stevens R, Shaw NL. 2004. Composite shrubs. In: Monsen
SB, Stevens R, eds. Restoring western ranges and wildlands. Ogden, UT:
USDA Forest Service Rocky Mountain Research Station.
McDonough WT, Harniss RO. 1974. Effects of temperature on germination
in three subspecies of big sagebrush. Journal of Range Management 27:
204205.
Meyer SE. 1990. Unpublished data. Provo, UT: USDA Forest Service, Rocky
Mountain Research Station.
Meyer SE. 1990. Seed source differences in germination under snowpack
in northern Utah. In: Munkshower F, ed. Fifth Billings Symposium on
Disturbed Land Reclamation.Volume 1. 1990 March 2530; Billings, MT.
Pub. 9003. Bozeman: Montana State University Reclamation Research
Unit: 184191.
Meyer SE. 1994. Germination and establishment ecology of big sagebrush:
implications for community restoration. In: Monsen SB, Kitchen SG, eds.
Proceedings, Ecology and Management of Annual Rangelands. 1992 May
1822; Boise, ID. Gen.Tech. Rep. INT-313. USDA Forest Service,
Intermountain Research Station: 244251.
Meyer SE, Kitchen SG. 1997. Unpublished data. Provo, UT: USDA Forest
Service, Rocky Mountain Research Station.
Meyer SE, Monsen SB. 1990. Seed-source differences in initial establishment
for big sagebrush and rubber rabbitbrush. In: McArthur ED, Romney EM,
Smith SD,Tueller PT, eds. Symposium, Cheatgrass Invasion, Shrub Die-off
and Other Aspects of Shrub Biology and Management. 1989 April 57;
Las Vegas. Gen.Tech. Rep. INT-276. Ogden, UT: USDA Forest Service,
Intermountain Research Station: 200208.
Meyer SE, Monsen SB. 1991. Habitat-correlated variation in mountain big
sagebrush (Artemisia tridentata ssp. vaseyana: Asteraceae) seed
germination patterns. Ecology 72: 739742.
Meyer SE, Monsen SB. 1992. Big sagebrush germination patterns: subspecies
and population differences. Journal of Range Management 45: 8793.
Meyer SE, Kitchen SG, Wilson GR, Stevens R. 1988a. Proposed rule for
Artemisia nova. Association of Official Seed Analysts Newsletter 62(1):
1617.
Meyer SE, Kitchen SG, Wilson GR, Stevens R. 1988b. Proposed rule for
Artemisia tridentata. Association of Official Seed Analysts Newsletter
62(1): 1718.
Meyer SE, Monsen SB, McArthur ED. 1990. Germination response of
Artemisia tridentata to light and chill: patterns of between-population
variation. Botanical Gazette 152: 176183.
Monsen SB. 1995. Personal communication. Provo, Utah: USDA Forest
Service Rocky Mountain Research Station.
Monsen SB, Meyer SE. 1990. Seeding equipment effects on establishment of
big sagebrush on mine disturbances. In: Munkshower F, ed. Fifth Billings
Symposium on Disturbed Land Rehabilitation.Volume 1, Hardrock waste,
analytical, and revegetation. 1990 March 2530; Billings, MT. Pub. 9003.
Bozeman: Montana State University, Reclamation Research Unit:
192199.
Monsen SB, Richardson BL. 1984. Seeding shrubs and herbs on a semiarid
minesite with and without topsoil. In:Tiedemann AR, McArthur ED, Stutz
HC, Stevens R, Johnson KL, eds. Proceedings, Symposium on the Biology
of Atriplex and Related Chenopods. 1983 May 26; Provo, UT. Gen.Tech.
Rep. INT-172. Ogden, UT: USDA Forest Service, Intermountain Research
Station: 298305.
Monsen SB, Meyer SE, Carlson SL. 1992. Sagebrush establishment enhanced
by snowfencing. In: Rangeland Technology and Equipment Council, USDA
Forest Service Technology and Development Program 2200-Range: 1992
Annual Report: 6-8.
Pechanec JF, Plummer AP, Robertson JH, Hull AC. 1944. Eradication of big
sagebrush (Artemisia tridentata). Res. Pap. 10. Ogden, UT: USDA Forest
Service, Intermountain Forest and Range Experiment Station. 23 p.
Plummer AP, Christensen DR, Monsen SB. 1968. Restoring big game range
in Utah. Pub. 68-3. Salt Lake City: Utah Division of Fish and Game
183 p.
Stevens R, Jorgensen KR, Davis JN. 1981. Viability of seed from thirty-two
shrub and forb species through fifteen years of warehouse storage.
Great Basin Naturalist 41: 274277.
Artemisia
279
Wagstaff FJ, Welch BL. 1991. Seedstalk production of mountain big sagebrush enhanced through short-term protection from heavy browsing.
Journal of Range Management 44: 7274.
Walton TP, White RS, Wambolt CL. 1986. Artemisia reproductive strategies:
a review with emphasis on plains silver sagebrush. In: McArthur ED,
Welch BL, eds. Symposium on the Biology of Artemisia and
Chrysothamnus. 1984 July 913; Provo, UT. Gen.Tech. Rep. INT-200.
Odgen, UT: USDA Forest Service, Intermountain Forest and Range
Experiment Station: 6774.
Welch BL. 1995. Beyond twelve percent purity. In: Roundy BA, McArthur
ED, Haley JS, Mann DK, comps. Proceedings, Symposium on Arid Lands
Ecology and Restoration. 1993 October 1921; Las Vegas. Gen.Tech.
Rep. INT-315. Ogden, UT: USDA Forest Service, Intermountain Research
Station: 126129.
Welch BL, Nelson DL. 1995. Black stem rust reduces big sagebrush seed
production. Journal of Range Management 48: 398401.
Welch BL, McArthur ED, Nelson DL, Pederson JC, Davis JN. 1986. Hobble
Creeka superior selection of low-elevation mountain big sagebrush.
Res. Pap. INT-370. Ogden, UT: USDA Forest Service, Intermountain
Forest and Range Experiment Station. 10 p.
280
Welch BL, Wagstaff FJ, Jorgensen GL. 1990. Hobble Creek mountain big
sagebrush seed production. In: McArthur ED, Romney EM, Smith S,
Tueller PT, eds. Symposium on Cheatgrass Invasion, Shrub Die-off, and
Other Aspects of Shrub Biology and Management. 1989 April 57; Las
Vegas. Gen.Tech. Rep. INT-276. Ogden, UT: USDA Forest Service,
Intermountain Research Station: 166170.
Weldon LW, Bohmont DW, Alley HP. 1959. The interrelation of three environmental factors affecting germination of sagebrush seed. Journal of
Range Management 12: 236238.
Wilson RG. 1982. Germination and seedling development of fringed
sagewort (Artemisia frigida). Weed Science 30: 102105.
Young JA, Evans RA. 1975. Germinability of seed reserves in a big sagebrush (Artemesia tridentata) community. Weed Science 23: 358364.
Young JA, Evans RA. 1989. Dispersal and germination of big sagebrush
(Artemisia tridentata) seeds. Weed Science 37: 201-06.
Young JA, Martens E. 1991. Importance of hypocotyl hairs in germination
of Artemisia seeds. Journal of Range Management 43: 358366.
Young JA, Evans RA, Palmquist DE. 1989. Big sagebrush (Artemisia
tridentata) seed production. Weed Science 37: 4753.
AnnonaceaeCustard-apple family
Asimina Adans.
pawpaw
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Growth habit and use. Of the 9 species of the genus
Asimina, seed data are available only for small-flower pawpaw and pawpaw (table 1). Both form shrubs or small deciduous trees (Vines 1960). Their fruits provide food for
wildlife and are also eaten by humans. There is some interest in commercial fruit production of pawpaw, and cultivar
selections have been made since the early part of the century
(Peterson 1990).
Flowering and fruiting. Flowers of the pawpaw
genus are solitary, perfect, and greenish purple. They appear
in the spring during March to May, about the same time as
the leaves. In natural stands of pawpaw, pollination and
seed set are very poor (Norman and others 1992; Willson
and Schemske 1980), conditions that discourage commercial
productions. In central Illinois, pawpaw averaged 3.5 to 10.5
seeds/fruit (Willson and Schemeske 1980). Pawpaw fruits
are 5 to 17 cm long, whereas those of small-flower pawpaw
are 5 to 12 cm long (Halls 1973). Pawpaw fruits are greenish yellow before maturity and turn brown to black as they
ripen in July to August and fall to the ground in August and
September. Seeds of small-flower pawpaw mature while the
fruit coat is still green (Norman and others 1992). The fruits
are fleshy berries that contain several dark brown, shiny
seeds (figure 1). The fleshy part of the fruit is considered
edible, but there appear to be 2 different fruit types. Those
with white flesh are barely edible, whereas others are larger
Figure 1 Asimina, pawpaw: fruits and seeds of A. parviflora, small-flower pawpaw (top) and A. triloba, pawpaw
(bottom).
Common name(s)
Occurrence
small-flower pawpaw,
small-fruited pawpaw,
small custard-apple,
dwarf pawpaw
pawpaw, custard-apple
common pawpaw
Texas E to Florida;
N to Virginia
3.5
12
Asimina
281
small-flower pawpaw
pawpaw
Cleaned seeds/wt 2,860/kg (1,300/lb) 1,540/kg per (700/lb)
Purity (%)
98
100
Sound seeds (%)
94
96
There is no storage information available on these
species.
Germination. Germination is usually very slow
because seeds have dormant embryos, and seedcoats are
slowly permeable. Moist stratification for 60 days at 5 C
resulted in germination of 50, 62, and 82% for 3 samples of
pawpaw seeds (Bonner and Halls 1974). Stratification for
100 days has been recommended, but germination still may
be slow and irregular. Fall-sowing of untreated seeds does
not improve results (Bonner and Halls 1974). No specific
test conditions have been reported, but alternating temperatures of 20 C during the day and 30 C at night on a moist
medium have been satisfactory for most species of the
northern temperate zone.
Nursery practice. Pawpaw seeds may be sown in the
fall without pretreatment, or stratified and sown in the
spring. Seeds should be covered about 20 mm (3/4 in) deep.
Some shade is helpful to germinating seedlings (figure 3).
Another method is to plant fresh seeds, before they dry, in
pots of sand and then to keep them in a cool cellar or similar
place. As the seeds sprout, they can be picked out and transplanted into nursery beds. Pawpaws can also be propagated
by layering and root cuttings (Bonner and Halls 1974) but
apparently not by stem cuttings (Dirr and Heuser 1987).
References
Bonner FT, Halls LK. 1974. Asimina, pawpaw. In: Schopmeyer CS, tech.
coord. Seeds of woody plants of the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 238239.
Dirr MA, Heuser CW. 1987. The reference manual of woody plant propagation. Athens, GA:Varsity Press. 239 p.
Halls LK. 1973. Flowering and fruiting of southern browse species. Res. Pap.
SO-90. New Orleans: USDA Forest Service, Southern Forest Experiment
Station. 10 p.
Norman EM, Rice K, Cochran S. 1992. Reproductive biology of Asimina
parviflora (Annonaceae). Bulletin of the Torrey Botanical Club 119: 15.
Peterson RN. 1990. Pawpaw (Asimina). In: Moore JN, Ballington JR, eds.
Genetic resources of temperate fruit and nut crops. Acta Horticulturae
290: 567600.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
Willson MF, Schemske DW. 1980. Pollinator limitation, fruit production, and
floral display in pawpaw (Asimina triloba). Bulletin of the Torrey Botanical
Club 107: 401408.
282
ChenopodiaceaeGoosefoot family
Atriplex L.
saltbush
Susan E. Meyer
Dr. Meyer is a research plant ecologist at the USDA Forest Services Rocky Mountain Research Station,
Shrub Sciences Laboratory, Provo, Utah
Growth habit, occurrence, and use. The genus
Atriplex L.saltbushis cosmopolitan in distribution and
comprises about 250 species of annual and perennial herbs,
subshrubs, and shrubs (McArthur and Sanderson 1984).
Most species are halophytic (at least to some degree) and
occupy salt desert, coastal strand, or saltmarsh habitats.
Shrubby species are important in arid and semiarid regions
throughout the world, with centers of diversity in south central Asia, Australia, temperate South America, and western
North America. Western North America is an area of particularly high genetic diversity, with more than 20 principal
species of shrubs and subshrubs as well as countless hybrids
and variants; 12 of these species are described here (table 1).
The genus is in a state of active evolution in the Intermountain region (Stutz 1978, 1984). The drying up of
Pleistocene lakes 10,000 or so years ago opened up vast
areas of unexploited salt-desert habitat. Shrubby saltbush
species migrated in rapidly from several directions and
hybridized freely, giving rise to the rich complex of forms in
the region today.
In terms of areal extent, the most important species are
probably shadscale and Gardner saltbushes (Blauer and others 1976). These species are regional dominants over millions of hectares in the Intermountain and northwestern
Great Plains regions, respectively. Shadscale saltbush mostly
occurs with winterfat (Krascheninnikovia lanata (Pursh)
Guldenstaedt.); budsage (Artemesia spinescens D.C. Eaton);
and other salt-desert shrubs, whereas Gardner saltbush is
able to maintain codominance with perennial grasses (Stutz
1978). In the Mojave Desert, desert-holly is an upland landscape dominant, particularly in the Death Valley region,
whereas allscale saltbush is a dominant species on playa
fringes. Fourwing saltbush is the most widely distributed
shrubby saltbush in North America and is often an important
component of grassland communities, especially in the
Chihuahuan Desert and western Great Plains. Sickle and
basin saltbushes are inconspicuous but common components
of northern Intermountain salt-desert vegetation.
Shrubby saltbush species are extremely important as forage plants for livestock and wildlife in arid and semiarid
regions worldwide (Goodall 1982). They provide palatable
and nutritious feed on a year-round basis and are especially
important on winter ranges. As a consequence, they have
been studied and used in range rehabilitation far more extensively than most other shrubs (Jones 1970; McArthur and
Monsen in press; Osmond and others 1980; Tiedemann and
others 1984). There is also considerable interest in utilizing
saltbush species as irrigated forage crops on marginal, salinized agricultural land (Glenn and others 1992; Watson and
OLeary 1993). Some shrubby saltbush species are also used
extensively for the stabilization of drastically disturbed land
because of their ability to establish and grow on harsh sites.
Geographic races and hybrids. An important feature
of infraspecific variation in many saltbush species is the
presence of series of races at different ploidy levels
(Sanderson and others 1990; Stutz 1978; Stutz and
Sanderson 1979). Polyploid races often show dwarfing and
adaptation to extremely harsh environments. The tendency
to evolve polyploid races has also been important in facilitating the formation and stabilization of interspecific
hybrids. Saltbush species possess a wealth of genetic variability, both within and among ploidy levels for numerous
traits that may be important for survival both of local populations in nature and of the products of artificial seedings.
Hybrid forms, even those that have not yet formed stabilized
populations in nature, may possess attributes that make them
useful in specific disturbed land rehabilitation applications
(Stutz 1995).
Common garden studies with fourwing saltbush have
demonstrated ecotypic variation in growth form, growth
rate, winter-greenness, drought and cold hardiness, palatability, nutrient status, seed size, and seed germination and
establishment traits (McArthur and others 1983; Springfield
1970; Van Epps 1975; Welch and Monsen 1981, 1984). It is
Atriplex
283
A. cuneata A. Nels.
Sources:
N Great Basin
Ansley and Abernathy (1985), Meyer (1996), Mikhail and others (1992),Young and others (1980).
A. semibaccata R. Br.
A. tridentata Kuntze
A. obovata Moq.
A. polycarpa (Torr.) S.Wats.
Colorado Plateau N to
Red Desert of Wyoming
Colorado Plateau
Geographic distribution
Common name(s)
Scientific name
Saline flats
Saline & subsaline slopes & flats
Ecology
(figure 1). The bracteoles are not fused to the utricle, but in
native species they usually enclose it so completely that
threshing is not possible. In Australian saltbush, the bracteoles are not fused across the top and the utricles may be
threshed free (figure 2) (Foiles 1974).
The saltbush seed itself is contained within the utricle
and is generally not separable from it (figure 3). The diskshaped seed has a curved embryo on its outer perimeter and
a scanty provision of storage tissue (in this case perisperm)
in the center. In most native species, the ovule (and thus the
seed) is inverted within the fruit, meaning that the radicle
end points upward. This facilitates radicle emergence from
between the bracteoles, which often have their only opening
or weakest point at the tip. The degree of woody thickening
of the bracteole walls varies among and within species and
may be linked to the persistent seed dormancy often encountered.
Seed collection, cleaning and storage. Saltbush seeds
are harvested by stripping or beating the ripe fruits into
shoulder hoppers, boxes, or bags, or onto tarps spread under
the bushes. Vacuum or reel-type harvesters may also be used
(McArthur and others 2004). In field cultivation, Wytana
fourwing and Gardner saltbushes have been cut and windrowed with a hay-swather and then combine-harvested.
Seeds shattered during combining were salvaged using a
vacuum harvester (Carlson and others 1984).
Seed collections of fourwing and shadscale saltbushes
are commonly hammermilled to remove the bracteole wings
(McArthur and others 2004). This reduces bulk by half and
facilitates cleaning, handling, and seeding through conventional drills. Hammermilling has little effect on dormancy of
Atriplex
285
Figure 3Atriplex semibaccata, Australian saltbush: exteror views in 2 planes of utricles removed from their bracts
and a longitudinal section through a utricle.
fourwing saltbush but may speed the germination of nondormant seeds somewhat (Gerard 1978; Springfield 1970).
Collections of wingless small-fruited species such as
Gardner, sickle, and mat saltbushes do not require hammermilling. Seed collections of all species may be cleaned by
screening and blowing in a fanning mill (McArthur and
others 2004).
Even relatively high-quality seedlots of saltbush may
average only 50% fill. A cut test to determine fill is often
286
A
Fruit (x1,000) /weight
Range
Species
/kg
A. canescens
intact
de-winged
A. confertifolia
A. corrugata
A. cuneata
A. falcata
A. gardneri
A. hymenelytra
A. lentiformis
A. obovata
A. polycarpa
A. semibaccata
A. tridentata
17120
29326
65277
210262
9002,000
7851,370
165317
120370
Average
/lb
855
13148
30126
95119
409909
357623
75144
55168
/kg
/lb
68
118
142
174
180
434
233
477
1,957
457
1,078
280
31
54
65
79
82
197
106
217
890
208
490
128
A. confertifolia
A. gardneri
A. hymenelytra
A. lentiformis
A. obovata
A. polycarpa
A. semibaccata
Storage
(months)
3
24
3
24
3
36
3
36
3
15
8
8
24
24
8
8
8
24
24
Incubation
treatment
Germination (%)
Mean
Range
15 C
15 C
4 wk @ 115 C
4 wk @ 115 C
5/15 C
5/15 C
16 wk @ 15/15 C
16 wk @ 15/15 C
Mean multiple treatments
Mean multiple treatments
5/15 C
10/20 C
Mean multiple treatments
Best treatment 10/25 %C
10/20 C & 0.05 M NaCl
10/20 C
20/30 C
Mean multiple treatments
Best treatment 10/25 C
32
54
41
69
0
2
16
46
26
48
33
56
39
68
42
53
50
41
69
496
10100
493
21100
01
06
047
483
2971
3940
1194
2179
3746
Samples
23
23
23
23
15
15
15
15
1
1
1
3
2
1
1
2
2
3
1
Sources: Ansley and Abernathy (1985), Meyer (unpublished data), Mikhail and others (1992),Young and others (1980).
Note: Germination period is 28 days and germination is expressed as percentage of filled fruits, except for A. gardneri data, where germination is 14 days, and for A.
lentiformis and A. semibaccata data, where germination is expressed as percentage of total fruits.
weakening the bracteole walls. Actual rupture of the membranous utricle wall is usually damaging to the seed (Sabo
and others 1979). After-ripening may also act on the bracteole walls, as evidenced by work with a seedlot of the South
American species A. repanda Phil., for which optimum time
for sulfuric acid scarification decreased from 7 to 2 hours
during 5 years in dry storage (Fernandez 1978). The bracte-
Atriplex
287
288
References
Ansley RJ, Abernethy RH. 1984. Seed pretreatments and their effects on
field establishment of spring-seeded gardner saltbush. Journal of Range
Management 37: 509513.
Ansley RJ, Abernethy RH. 1985. Environmental factors influencing Gardner
saltbush seed dormancy alleviation. Journal of Range Management 38:
331335.
Beadle NCW. 1952. Studies in halophytes: 1.The germination of the seed
and establishment of the seedling of five species of Atriplex in Australia.
Ecology 33: 4962.
Belcher E. 1985. Handbook on seeds of browse-shrubs and forbs.Tech.
Pub. R8-TP8. Atlanta: USDA Forest Service Southern Region. 246 p.
Blauer AC, Plummer AP, McArthur ED, Stevens R, Giunta BC. 1976.
Characteristics and hybridization of important Intermountain shrubs: 2.
Chenopod family. Res. Pap. INT-177. Ogden, UT: USDA Forest Service,
Intermountain Forest and Range Experiment Station. 42 p.
Bleak AT, Frischknecht NC, Plummer AP, Eckert RE. 1965. Problems in artifical and natural revegetation in the arid shadscale vegetation zone of
Utah and Nevada. Journal of Range Management 18: 5963.
Briggs JA. 1984. Seed production of Atriplex canescens in southern Arizona.
In:Tiedemann AR, McArthur ED, Stutz HC, Stevens R ,Johnson KL, compilers. Proceedings, Symposium on the Biology of Atriplex and Related
Chenopods; 1983 May 46; Provo, UT. Gen.Tech. Rep. INT-172. Ogden,
UT: USDA Forest Service, Intermountain Forest and Range Experiment
Station: 187190.
Carlson JR. 1984. Atriplex cultivar development. In:Tiedemann AR,
McArthur ED, Stutz HC, Stevens R, Johnson KL, comps. Proceedings,
Symposium on the Biology of Atriplex and Related Chenopods; 1983
May 46; Provo, UT. Gen.Tech. Rep. INT-172. Ogden, UT: USDA Forest
Service, Intermountain Forest and Range Experiment Station: 176182.
Carlson JR, Scheetz JG, Oaks WR. 1984. Seed production techniques of
two chenopods: Gardner saltbush and winterfat. In:Tiedemann AR,
McArthur ED, Stutz HC, Stevens R, Johnson KL, comps. Proceedings,
Symposium on the Biology of Atriplex and Related Chenopods; 1983
May 46; Provo, UT. Gen.Tech. Rep. INT-172. Ogden, UT: USDA Forest
Service, Intermountain Forest and Range Experiment Station: 191195.
Cornelius DR, Hylton LO. 1969. Influence of temperature and leachate on
germination of Atriplex polycarpa. Agronomy Journal 61: 209211.
Edgar RL, Springfield HW. 1977. Germination characteristics of broadscale:
a possible saline-alkaline site stabilizer. Journal of Range Management 30:
296298.
Everett RL, Meewig RO, Stevens R. 1978. Deer mouse preference for seed
of commonly planted species, indigenous weed seed, and sacrifice foods.
Journal of Range Management 31: 7073.
Ferguson RB. 1980. Potting media for Atriplex production under greenhouse conditions. Res. Note INT-301. Ogden, UT: USDA Forest Service,
Intermountain Forest and Range Experiment Station. 4 p.
Fernandez HG. 1978. Aumento de la germinacion en Atriplex repanda: 2.
Efecto de diferentes tratamientos quimicos al pericarpio. Phyton 36:
123127.
Foiles MW. 1974. Atriplex, saltbush. In: Schopmeyer CS, ed. Seeds of woody
plants in the United States. Agric. Handbk. 450. Washington, DC: USDA
Forest Service: 240243.
Garvin SC, Meyer SE, Carlson SL. 1996. Seed germination studies in
Atriplex confertifilia (Torr. and Frem.) Wats. In: Barrow JR, McArthur ED,
Sosebee RE,Tausch RJ, comps. Proceedings, Shrubland Ecosystem
Dynamics in a Changing Environment; 1995 May 2325; Las Cruces, NM.
Gen.Tech. Rep. INT-338. Fort Collins, CO: USDA Forest Service, Rocky
Mountain Forest and Range Experiment Station: 165169.
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289
Gerard JB. 1978. Factors affecting fruit fill and seed germination of fourwing
saltbush (Atriplex canescens (Pursh) Nutt.). In: Hyder DN, ed.
Proceedings, 1st International Rangeland Congress. Denver: Society for
Range Management.
Glenn EP, Watson MC, OLeary JW, Axelson RD. 1992. Comparison of salt
tolerance and osmotic adjustment of low-sodium and high-sodium
subspecies of the C4 halophyte, Atriplex canescens. Plant, Cell and
Environment 15: 711718.
Goodall DW. 1982. Chenopod shrubland communities: a global perspective. International Journal of Ecology and Environmental Science 9:
8599.
Graves WL, Kay BL, Williams WA. 1975. Seed treatment of Mojave Desert
shrubs. Agronomy Journal 67: 773777.
Haws BA, Bohart GE, Meadows RW, Coombs EM, Roe AR. 1984. Status of
information concerning insects associated with selected species of
Atriplex. In:Tiedemann AR, McArthur ED, Stutz HC, Stevens R, Johnson
KL, comps. Proceedings, Symposium on the Biology of Atriplex and
Related Chenopods; 1983 May 46; Provo, UT. Gen.Tech. Rep. INT-172.
Ogden, UT: USDA Forest Service, Intermountain Forest and Range
Experiment Station: 226236.
Jones R. 1970. The biology of Atriplex. Canberra, Australia: CSIRO. 128 p.
Kay BL, Brown CR, Graves WL. 1977a. Desert saltbush. Mojave Reveg.
Notes 18. Davis: University of California, Department of Agronomy and
Range Science. 7 p.
Kay BL, Brown CR, Graves WL. 1977b. Fourwing saltbush. Mojave Reveg.
Notes 17. Davis: University of California, Department of Agronomy and
Range Science. 12 p.
McArthur ED, Sanderson SC. 1984. Distribution, systematics, and evolution
of Chenopodiaceae: an overview: In:Tiedemann AR, McArthur ED, Stutz
HC, Stevens R, Johnson KL, comps. Proceedings, Symposium on the
Biology of Atriplex and Related Chenopods; 1983 May 46; Provo, UT.
Gen.Tech. Rep. INT-172. Ogden, UT: USDA Forest Service, Intermountain
Forest and Range Experiment Station: 1423.
McArthur ED, Blauer AC, Noller GL. 1984. Propagation of fourwing saltbush by stem cuttings. In:Tiedemann AR, McArthur ED, Stutz HC,
Stevens R, Johnson KL, comps. Proceedings, Symposium on the Biology of
Atriplex and Related Chenopods; 1983 May 46; Provo, UT. Gen.Tech.
Rep. INT-172. Ogden, UT: USDA Forest Service, Intermountain Forest
and Range Experiment Station: 261264.
McArthur ED, Freeman CD, Luckinbill LS, Sanderson SC, Noller GL. 1992.
Are trioecy and sexual lability in Atriplex canescens genetically based?
Evidence from clonal studies. Evolution 46: 17081721.
McArthur ED, Monsen SB, Shaw NL. 2004. Chenopod shrubs. In: Monsen
SB Stevens R. Shaw NL, eds. Restoring western ranges and wildlands.
USDA Forest Service, Intermountain Research Station.
McArthur ED, Plummer AP, Van Epps GA, Freeman DC, Jorgensen KR. 1978.
Producing fourwing saltbush seed in seed orchards. In: Heyder DC, ed.
Proceedings, 1st International Rangeland Congress. Denver: Society for
Range Management: 406410.
McArthur ED, Stevens R, Blauer AC. 1983. Growth performance comparisons among 18 accessions of fourwing saltbush (Atriplex canescens) at
two sites in central Utah. Journal of Range Management 36: 7881.
Meyer SE, Allen PS, Wilson GR, Davis TD, Davis JN, Stevens R, Jorgensen K.
1986. Proposed rule for Atriplex canescens. Association of Official Seed
Analysts Newsletter 60(1): 1415.
Meyer SE, Carlson SL, Garvin SC. 1998. Seed germination regulation and
field seed bank carryover in shadscale (Atriplex confertifolia:
Chenopodiaceae). Journal of Arid Environments 38: 255267.
Mikhiel G, Meyer SE, Pendleton RL. 1992. Variation in germination response
to temperature and salinity in shrubby Atriplex species. Journal of Arid
Environments 22: 3949.
Munz PA. 1974. A flora of southern California. Berkeley: University of
California Press. 1086 p.
Nord EC, Whitacre JE. 1957. Germination of fourwing saltbush seed
improved by scarification and grading. Forest Research Notes 125: 15.
Nord EC, Hartless PF, Nettleton WD. 1971. Effects of several factors on
saltbush establishment in California. Journal of Range Management 24:
216223.
Nord EC,Van Atta GR. 1960. Saponin: a seed germination inhibitor. Forest
Science 6: 350353.
Osmond CB, Bjorkman O, Anderson DJ. 1980. Physiological processes in
plant ecology: toward a synthesis with Atriplex. Berlin: Springer-Verlag.
Plummer AP, Monsen SB, Christensen DR. 1966. Fourwing saltbush, a shrub
for future game ranges. Pub. 66-4. Salt Lake City: Utah State Department
of Fish and Game: 112.
Plummer AP, Christensen DR, Monsen SB. 1968. Restoring big game range
in Utah. Pub. 68-3. Salt Lake City: Utah Division of Fish and Game. 183 p.
290
Richardson SG, Barker JR,Van Epps GA. 1979. Factors affecting rooting
stem cuttings of salt desert shrubs. Journal of Range Management 32:
280283.
Sabo DG, Johnson DV, Martin WC, Aldon EF. 1979. Germination requirements of 19 species of arid land plants. Res. Pap. RM-210. Fort Collins,
CO: USDA Forest Service, Rocky Mountain Forest and Range
Experiment Station: 126.
Sanderson SP, Pendleton RL, McArthur ED, Harper KT. 1986. Saponin effect
on small mammal forage preference in a planting of Atriplex canescens. In:
Provenza FJ, Flinders JT, McArthur ED, comps. Proceedings, Wildland
Shrub Symposium on PlantHerbivore Interactions. Gen.Tech. Rep. INT222. Ogden, UT: USDA Forest Service, Intermountain Forest and Range
Experiment Station: 7477.
Sanderson SC, Stutz HC, McArthur ED. 1990. Geographic differentiation in
Atriplex confertifolia. American Journal of Botany 77: 490498.
Shaw N, Monsen SB. 1984. Nursery propagation and outplanting of bareroot chenopod seedlings. In:Tiedemann AR, McArthur ED, Stutz HC,
Stevens R, Johnson KL, comps. Proceedings, Symposium on the Biology of
Atriplex and Related Chenopods; 1983 May 46; Provo, UT. Gen.Tech.
Rep. INT-172. Ogden, UT: USDA Forest Service, Intermountain Forest
and Range Experiment Station: 251260.
Springfield HW. 1970. Germination and establishment of fourwing saltbush
in the southwest. Res. Pap. RM-55. Fort Collins, CO: USDA Forest
Service, Rocky Mounntain Forest and Range Experiment Station: 148.
Stevens R, Jorgensen KR, Davis JN. 1981. Viability of seed from thirty-two
shrub and forb species through fifteen years of warehouse storage. Great
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Stroh JR,Thornberg AA. 1969. Culture and mechanical harvest of fourwing
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Stutz HC. 1984. Atriplex hybridization in western North America. In:
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University.
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Tiedemann AR, McArthur ED, Stutz HC, Stevens R, Johnson KL, comps. 1984.
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UT: USDA Forest Service, Intermountain Forest and Range Experiment
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and chloride removal. Journal of Range Management 8: 218220.
Van Epps GA. 1975. Winter injury to fourwing saltbush. Journal of Range
Management 28: 157159.
Vest ED. 1952. A preliminary study of some of the germination characteristics of Atriplex confertifolia [MS thesis]. Salt Lake City: University of Utah.
46 p.
Warren DC, Kay BL. 1984. Pericarp inhibition of germination in Atriplex
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Watson MC, OLeary JW. 1993. Performance of Atriplex species in the San
Joaquin Valley, California, under irrigation and with mechanical harvests.
Agriculture, Ecosystems and Environment 43: 255-266.
Welch BL, Monsen SB. 1981. Winter crude protein differences among
accessions of fourwing saltbush grown in a common garden. Great Basin
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UT. Gen.Tech. Rep. INT-172. Ogden, UT: USDA Forest Service,
Intermountain Forest and Range Experiment Station: 138144.
Young JA, Kay BL, George H, Evans RA. 1980. Germination of three species
of Atriplex. Agronomy Journal 72: 705709..
AsteraceaeAster family
Baccharis L.
baccharis
hairs 13 mm long or less (figures 1 and 2). Achenes are dispersed by wind soon after ripening (table 2). Seedcrops are
borne annually.
Quantities of seeds produced on an individual plant can
be very high in full sunlight. A single plant of eastern baccharis has been estimated to produce over 1 million seeds
(Westman and others 1975). Dense shade (3% of full sunlight) reduced seed production dramatically but did not
totally eliminate it (Westman and others 1975).
Collection of fruits; extraction and storage of seeds.
The ripe fruits of baccharis are either collected by hand or
brushed onto cloth or plastic sheets spread beneath the
shrubs. The fruits should be spread out to dry in a warm
well-ventilated room or in the sun, protected from the wind.
When dried, the fruits may be rubbed between the hands or
treated in bulk to remove the pappus. Alternatively, full
inflorescences can be fed into a brush machine, where the
fruit is threshed from the stems and the pappus removed.
The seeds can then be cleaned with air, screens, or other
equipment described in the seed handling chapter.
Sometimes the entire fruits are used without removing the
pappus. The number of fruits per weight for coyotebrush is
about 180,800/kg (82,000/lb) (1 sample); for mulefat
baccharis, about 110,250/kg (50,000/lb) (1 sample) (Olson
1974). Cleaned seeds of baccharis species can be stored dry
at 1.7 to 4.5 C in airtight containers (McBride 1964). Data
published by Westman and others (1975) indicate that seeds
of eastern baccharis could be stored for 1 to 4 months at
room temperature. After 4 months of room temperature storage, the final germination was actually slightly higher than
with seeds stored for only 1 month. Panetta (1979) found
that seeds stored in an atmosphere of 33% relative humidity
maintained their germination of 98% for 12 months at 20 C
but that their percentage germination had dropped to 67%
by 24 months. For seeds stored in the laboratory in a constant 70% relative humidity, germination began to drop at 6
Baccharis
291
Common name(s)
Occurrence
B. angustifolia Michx.
saltwater falsewillow,
narrowleaf baccharis
Bigelow falsewillow
shortleaf baccharis
broombush falsewillow
saltmarsh baccharis
Emory baccharis
silverling
eastern baccharis
B. bigelovii Gray
B. brachyphylla Gray
B. dioica Vahl
B. douglasii DC
B. emoryi Gray
B. glomeruliflora Pers.
B. halimifolia L.
B. halimifolia var. angustior DC.
B. havardii Gray
B. myrsinites (Lam.) Pers.
B. neglecta Britt.
B. pilularis DC.
B. pilularis ssp.
consanguinea (DC.) C.B.Wolf
B. pilularis var.
consanguinea (DC.) Kuntze
B. plummerae Gray
B. pteronioides DC.
B. salicifolia (Ruiz & Pavon) Pers.
B. viminea DC.
Molina salicifolia Ruiz & Pavon
B. glutiosa Pers.
B. sarothroides Gray
B. sergiloides Gray
B. texana (Torr. & Gray) Gray
Linosyris texana Torr. & Gray
B. thesioides Kunth
B. vanessae Beauchamp
B. glutinosa
B. wrightii Gray
Sources:
Harvard falsewillow
Santo Domingo falsewillow
Rooseveltweed
coyotebrush,
kidneywort baccharis
Plummer baccharis
yerba de pasmo
mulefat baccharis
California
Arizona, New Mexico, & Texas
California E to Texas & Utah
desertbroom
squaw waterweed baccharis
prairie falsewillow
Arizona baccharis
Encinitis falsewillow
Wright baccharis
292
nation than no prechilling or prechilling at 0 C when eastern baccharis was germinated at 10, 15, or 20 C with continuous light. In a greenhouse test of eastern baccharis,
there was no apparent reduction in germination under 56.7,
23.6, or 17.4% of full sunlight (Panetta 1990). Embryo excision was found to speed embryo germination in both
Encinitis falsewillow and eastern baccharis (Kuti and others
1990), demonstrating that there is some inhibitory effect
from the seedcoat.
Nursery practice. Seeds may be sown in the fall or
early spring in flats or seedbeds using a sandy soil mixture,
or one of the vermiculite, perlite, or sphagnum moss seeding media (Everett 1957). Seeds usually germinate within 7
to 15 days. Plants large enough for 10-cm (4-in) pots can be
taken from outside seedbeds within 4 months (Everett 1957)
(figure 3). Rooseveltweed seeds sown in 15-cm-deep (6-indeep) pots germinated slowly, requiring 1 month to establish seedlings (Van Auken and Bush 1990).
seedling
Flowering
Fruit ripening
Seed dispersal
B. angustifolia
B. pilularis
B. salicifolia
SeptOct
JulyOct
MayJuly
SeptOct
SeptDec
MayJuly
Oct
Fall
MayJuly
Sources: McBride (1964), Mirov and Kraebel (1939), Olson (1974), Radford and others (1964).
Baccharis
293
Species
Medium
B. angustifolia
B. halimifolia
B. pilularis
Kimpak
Moist paper
Moist paper
Moist paper
B. salicifolia
15.6
19
19
17.3
7.225
20
55
10
10
1530
30
1530
Germination
Average
(%)
Samples
21
92
93
92
4054
7582
2
1
1
1
28
3
Sources: McBride (1964, 1969), Mirov and Kraebel (1939), Olson (1974), Panetta (1979).
References
Barkley TM. 1986. Flora of the great plains. Lawrence: University Press of
Kansas. 181 p.
BONAP. 1996. The digital checklist of the vascular flora of North America
[access website at
http://shanana.berkeley.edu/bonap/checklist_intro.html].
Correll DS, Johnston MC. 1970. Manual of vascular plants of Texas.
Renner:Texas Research Foundation. 1881 p.
Day AD, Ludeke KL. 1980. Reclamation of copper mine wastes with
shrubs in the southwestern U.S.A. Journal of Arid Environments 3:
107112.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant
propagation: from seed to tissue culture. Athens, GA:Varsity Press.
239 p.
Emery D. 1964. Seed propagation of native California plants. Leaflets of
the Santa Barbara Botanic Garden 1(10).
Everett P C. 1957. A summary of the culture of California plants at the
Rancho Santa Ana Botanic Garden, 19271950. 223 p.
Kuti JO, Jarvis BB, Mokhtari-Rejali N, Bean G.A. 1990. Alleochemical regulation of reproduction and seed germination of two Brazilian Baccharis
species by phytotoxic tricothecenes. Journal of Chemical Ecology
16(12): 34413453.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third. New York:
Macmillan. 1290 p.
McBride JR. 1964. Invasion of park grasslands by Baccharis pilularis DC [MS
thesis]. Berkeley: University of California.
294
FabaceaePea family
Bauhinia L.
bauhinia
Kristina F. Connor
Dr. Connor is a plant physiologist at the USDA Forest Services Southern Research Station,
Auburn University, Alabama
Growth habit, occurrence, and use. There are about
600 species of the bauhinia genus found in the tropical
regions of the world (Larson 1974). The genus includes
trees, vines, and shrubs that are frequently planted for their
showy flowers and ornamental foliage (Bailey 1941; Neal
1965). Practical usage of the bark of orchidtree as an astringent in tanning and dyeing and of the leaves and flower buds
as a vegetable has been reported (Bailey 1941). Seeds of
some bauhinia species have served as a human food source
(malucreeper, B. vahlii Wight & Arn.) (Ramasastri and
Shenolikar 1974); a source of vitamin A (butterfly bauhinia)
(Essien and Fetuga 1989); and as a possible pest control
agent (malucreeper) (Freedman and others 1979). Butterfly
bauhinia is used for fuelwood on Puerto Rico and for fences
on Jamaica (Little and Wadsworth 1964), but it is considered
a weed on Guam (McConnell and Muniappan 1991). Four
species, all small evergreen or deciduous trees, have been
planted in the continental United States (table 1). Hawaii has
13 species of introduced bauhinias (Neal 1965), whereas
Puerto Rico has at least 5 (Francis and Liogier 1991).
Flowering and fruiting. The large 5-petaled orchidlike flowers of bauhinias occur in racemes and range in
color from white to deep purple and yellow. The fruits
flowers and
Common name(s)
Occurrence
B. megalandra Griseb.
B. multinervia (Kunth) DC.
B. monandra Kurz
B. kappleri Sagot
Caspareopsis monandra (Kurz) Britt. & Rose
B. purpurea L.
Phanera purpurea (L.) Benth.
Caspareopsis purpurea (L.) Pittier
B. variegata L.
bauhinia
petite flambouyant
butterfly bauhinia,
pink bauhinia,
pink orchidtree
purple bauhinia
Carribean basin
orchidtree,
poor-mans-orchid,
mountain-ebony
Sources: Francis and Liogier (1991), Little and others (1974), Neal (1965).
Bauhinia
295
Flowering time
Petal color
Fertile stamens/flower
B. monandra
All year
B. purpurea
34
B. variegata
Autumn to spring
Purple variegated
with red & yellow
56
stamen per flower and a calyx splitting along one side (Little
and Wadsworth 1964; table 2). Flowers of purple bauhinias
have 3 to 4 fertile stamens and a 2-parted calyx, whereas
those of orchidtrees have 5 to 6 fertile stamens/flower and a
calyx that splits on one side (Little and others 1974; Neal
1965). Information on pollinators is scarce, but Heithaus and
others (1982) report that B. ungulata L. is pollinated by
bats and that 59.4% of flowers examined show evidence of
herbivory.
Butterfly bauhinia seeds are elliptic, flat, and 1 cm long;
fruits are present throughout the year (Little and Wadsworth
1964). Purple bauhinia seeds are shiny-brown, rounded, flat,
and range in length from 1.3 to 1.6 cm; flowering and fruiting occur in autumn and winter months (Little and others
1974). Orchidtree seeds are fairly large, about 1.3 cm in
diameter, and the fruits mature in late spring or early summer. Bauhinia megalandra seeds are shown in figure 2
and 3. Rugenstein and Lersten (1981) report the presence of
stomata on the seeds and pods of purple bauhinias and
orchidtrees. In general, bauhinia seeds contain high amounts
of linoleic and oleic fatty acids and low amounts of myristic
and linolenic fatty acids (Balogun and Fetuga 1985;
Legume
1530 cm long, pointed at apex,
twists as opens
1530 cm long, black, thin, twists as opens
1330 cm long, thin,
pointed on both ends
296
seed.
longitudal
Seeds/wt
/kg
/lb
5,680
4,670
4,950
2,576
2,118
2,245
Germination *
Period (days) Percentage
4
4
4
100
99
77
References
Athaya CD. 1985. Ecological studies of some forest tree seeds: 2. Seed storage and viability. Indian Journal of Forestry 8(2): 137140 [Forestry
Abstracts 47: 3250; 1986].
Babeley GS, Kandya AK. 1986. Excised-embryo test of seed germinability:
an evaluation through the seeds of six dry-deciduous tropical forest tree
species. Journal of the Japanese Forestry Society 68(5): 197199
[Forestry Abstracts 47: 5543; 1986].
Bailey LH. 1941. The standard cyclopedia of horticulture. New York:
MacMillan. 1200 p.
Balogun AM, Fetuga BL. 1985. Fatty acid composition of seed oils of some
members of the Leguminosae family. Food Chemistry 17(3): 175182
[Nutritional Abstracts and Review Series A 55: 6146; 1985].
Essien AI, Fetuga BL. 1989. Beta-carotene content and some characteristics
of under-exploited seed oils of forest trees in Nigeria. Food Chemistry
32(2): 109116 [Forestry Abstracts 50: 5676; 1989].
Francis JK, Liogier HA. 1991. Naturalized exotic tree species in Puerto
Rico. Gen.Tech. Rep. SO-82. New Orleans: USDA Forest Service,
Southern Forest Experiment Station. 12 p.
Francis JK, Rodrguez A. 1993. Seeds of Puerto Rican trees and shrubs:
second installment. Res. Note SO-374. New Orleans: USDA Forest
Service, Southern Forest Experiment Station. 5 p.
Freedman B, Nowak LJ, Kwolek WF, Berry EC, Guthrie WD. 1979. A
bioassay for plant-derived pest control agents using the European corn
borer. Journal of Economic Entomology 72(4): 541545 [Rev. Applied
Entomology, Series A: 1973+].
Heithaus ER, Stashko E, Anderson PK. 1982. Cumulative effects of
plantanimal interactions on seed production by Bauhinia ungulata, a
neotropical legume. Ecology 63(5): 12941302 [Herbage Abstracts
1973+].
Bauhinia
297
BerberidaceaeBarberry family
Berberis L.
barberry
Don Minore and Paul O. Rudolf
Dr. Minore retired from USDA Forest Services Pacific Northwest Research Station;
Dr. Rudolf (deceased) retired from the USDA Forest Services North Central Forest Experiment Station
Growth habit, occurrence, and use. The barberries
include about 500 species of spiny or unarmed, evergreen or
deciduous shrubs (rarely small trees) native to Asia, Europe,
North Africa, and to North, Central, and South America
(Ahrendt 1961). Some authorities consider that the genus
Mahonia, consisting of about 100 species that closely
resemble the barberries, should be a section of Berberis
(Hitchcock and others 1964), whereas others consider
Mahonia to be a separate genus (Ahrendt 1961). The USDA
plant nomenclature system (USDA NRCS 1999) separates
them into 2 separate genera, and that is the authority used
for this manual (table 1). Thus, Mahonia is treated separately in a later chapter. The barberry genus is essentially
diploid, with 2n = 28 (Cadic 1992). Many interspecific
hybrids are known, such as those between Japanese and
Table 1Berberis, barberry: nomenclature, height, growth habit, and color of ripe fruit
Common name(s)
Height at
maturity (m)
B. buxifolia Lam.
B. candidula (C.K. Schneid.)
C.K. Schneid.
B. circumserrata (C.K. Schneid.)
C.K. Schneid.
B. darwinii Hook.
B. gagnepainii C.K. Schneid.
B. gilgiana Fedde
B. julianiae Schneid.
boxleaf barberry
paleleaf barberry
B. koreana Palibin.
B. sargentiana C.K. Schneid.
B. thunbergii DC.
B. trifoliata Moric.
B. tricanthophora Fedde
B. vernae C.K. Schneid.
B. verruculosa Hensl. &
E.H.Wilson
B. vulgaris L.
Growth habit
Color of
ripe fruits
0.62.1
0.61.2
Deciduous
Evergreen
Pruinose blue
Purplish, bloomy
cutleaf barberry
0.60.9
Deciduous
Pale red
Darwin barberry
black barberry
wildfire barberry
Julian barberry,
wintergreen barberry
Korean barberry
Sargent barberry
Japanese barberry
1.52.4
0.91.8
1.82.4
1.83.0
Evergreen
Evergreen
Deciduous
Evergreen
Pruinose blue
Pruinose blue
Reddish
Bluish-black
1.21.8
1.82.7
0.91.8
Deciduous
Evergreen
Deciduous
Bright red
Black
Bright red
threespine barberry
Verna barberry
warty barberry
0.91.5
0.91.2
0.91.8
Evergreen
Deciduous
Evergreen
Bluish-black
Pale red
Violet-black
common barberry,
European barberry
1.83.0
Deciduous
Scarlet or purple
Sources: Ahrendt (1961), Dirr (1990), Dirr and Heuser (1987), Garrett (1969), Hitchcock and others (1964), McMinn (1951), Rehder (1940), Rudolf (1974),Vines (1960).
298
Berberis
299
Species
Origin
Location
Flowering
Fruit ripening
B. koreana
B. thunbergii
Korea
Japan
B. vulgaris
Europe
Mayearly June
AprJune
MarApr
MayJune
AprJune
SeptOct
Oct
MaySept
SeptNov
SeptOct
Sources: Bailey (1939), Loiseau (1945), McMinn (1951), Mirov and Kraebel (1939), NBV (1946), Ohwi (1965), Plummer and others (1965), Radford and others (1964),
Rudolf (1974),Van Dersal (1938),Vines (1960),Wappes (1982),Wyman (1947).
* Fruits of these 3 species often remain on bushes over winter.
To 1,525 m in the Alps.
300
Germination tests. Germination of seeds from several barberry species has been tested in sand-filled flats, in
petri dishes, on paper or blotters, or in standard germinators.
Day temperatures of 16 to 30 C, night temperatures of
13 to 21 C, and germination periods of 20 to 95 days have
been used. Results are summarized in table 4. For Japanese
and common barberries, the Association of Official Seed
Analysts (AOSA 1993) recommends germination of excised
embryos in covered petri dishes at temperatures of 18 to
22 C for 10 to 14 days. This method may be satisfactory
for other barberry species.
Nursery practice. Whole berries or (preferably)
cleaned seeds may be sown in the fall, or stratified seeds
may be sown in the spring. Injury from molds is more likely
if whole berries are used (Chadwick 1936). Fall-sown beds
should be mulched until germination begins (NBV 1946).
The seeds should be covered with 0.3 to 1.3 cm (1/8 to
1/ in) of soil plus 0.6 cm (1/ in) of sand (Rudolf 1974).
2
4
Germination is epigeal (Terabayashi 1987), and seedlings
develop rapidly (figure 2). In a sowing of common barberry,
22% of the seeds survived to produce shrubs (Swingle
1939). Barberries may be field-planted as 2+0 stock (Rudolf
1974).
The barberries can be propagated from rooted stem cuttings. Several deciduous species are best rooted when propagated from softwood cuttings collected in the summer, but
many of the evergreen species root better when hardwood
cuttings are collected in the autumn or winter (Dirr and
Heuser 1987). Both should be treated with indole butyric
acid (IBA) rooting hormone in talc or in solution.
B. koreana
B. thunbergii
B. vulgaris
5582
7590
/kg
16
60
90
40
Sand or perlite
Wet paper or sand
Wet paper or sand
Medium
16
24
24
16
13
13
20
40
40
Germination rate
Amount
(%)
Days
88
90
91
1
4
2+5
% germination
Avg
(%)
Samples
84
64
84
97
93
Purity
(%)
38
29
38
Sources: Davis (1927), Heit (1968a,b), McLean (1967), Mirov and Kraebel (1939), Morinaga (1926), Plummer and others (1968), Rafn and son (nd), Rudolf (1974), Swingle (1939),Vines (1960).
* Cold stratification temperatures ranged from -1 to 5 C.
Twenty-one to 27 C during the day and 10 to 16 C during the night.
B. koreana
B. thunbergii
B. vulgaris
Species
Daily
light
(hrs)
Cold
stratification*
(days)
/lb
2537
3441
Range
Table 4Berberis, barberry: stratification periods, germination test conditions, and results for 3 species
39
915
Carver Co., MN
US
US
Species
30
1625
Place
collected
95
95
96
Soundness
(%)
2
5+
2
Samples
Berberis
301
References
302
Malasi CB, Chauhan JS, Paliwal GS. 1989. Influence of growth substances on
pollen germination, fruit-set and fruit growth in Berberis asiatica Roxb.
Indian Journal of Forestry 12(1): 2933.
McLean A. 1967. Germination of forest range species from southern
British Columbia. Journal of Range Management 20(5): 321322.
McMinn HE. 1951. An illustrated manual of California shrubs. Berkeley:
University of California Press. 663 p.
Millet B. 1976. Kinetic study of the curve of Mahonia aquifolium (Pursh)
Nutt. stamens electrically stimulated. Annales Scientifique de lUniversite
de Besancon, Botanique 3(17): 7580 [Biological Abstracts 68(5): 3095,
Ref. 31123].
Millet B. 1977. A scanning electron microscope survey of the surface of
the staminal filament in some Berberidaceae: Comparison with other
sensitive organs. Compte Rendu des Seances de la Societe de
Biologique et des Ses Filiales 171(3): 58084 [Biological Abstracts 65(8):
4336, Ref. 44154].
Minore D. 1994. Unpublished observation. Corvallis, OR: USDA Forest
Service, Pacific Northwest Research Station.
Mirov NT, Kraebel CJ. 1939. Collecting and handling seeds of wild plants.
For. Pub. 5. Washington, DC: Civilian Conservation Corps. 42 p.
Morinaga T. 1926. Effect of alternating temperatures upon the germination
of seeds. American Journal of Botany 13: 141158.
Mosch J, Zeller W. 1989. Bekampfung des Feuerbrandes (Erwinia amylovora)
mit ausgewahlten Pflanzenextrakten. Nachrichtenblatt des Deutschen
Pflanzenschutzdienstes [Braunschweig] 41(8/9): 149151 [Horticultural
Abstracts 1991 061-01390].
NBV [Nederlandsche Boschbouw Vereeniging]. 1946. Boomzaden:
Handleiding inzake het oogsten, behandelen, bewaren en uitzaaien van
boomzaden. Wageningen,The Netherlands: Ponsen and Looijen. 171 p.
Obeso JR. 1989. Fruit removal and potential seed dispersal in a southern
Spanish population of Berberis vulgaris ssp. Australis (Berberidaceae).
Acta Oecologica, Oecologica Plantarum 10(3): 321328 [Biological
Abstracts 89(1):AB-281, Ref. 2707].
Ohwi J. 1965. Flora of Japan. Washington, DC: Smithsonian Institution.
1067 p.
Pitea M, Petcu P, Goina T, Preda N. 1972. Dnnschictchromatographische
Untersuchungen der Alkaloide von Berberis vulgaris. Planta Medica
21(2): 177181 [Biological Abstracts 54(6): 3187, Ref. 33072].
Plummer AP, Christensen DR, Monsen SB. 1968. Restoring big-game range
in Utah. Pub. 68-3. Salt Lake City: Utah Division of Fish and Game.
182 p.
Radford AE, Ahles HE, Bell CR. 1964. Guide to the vascular flora of the
Carolinas. Chapel Hill: University of North Carolina Book Exchange.
333 p.
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18871927. Copenhagen: Udfrt paa Statsfrkontrollen i Kbenhavn.
5 p.
Rehder A. 1940. Manual of cultivated trees and shrubs hardy in North
America. 2nd ed. New York: Macmillan. 996 p.
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North Central Forest Experiment Station.
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Schlosser WE, Blatner KH, Zamora B. 1992. Pacific Northwest forest lands
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4455.
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5356.
BetulaceaeBirch family
Betula L.
birch
Robert P. Karrfalt
Mr. Karrfalt is director of the USDA Forest Services National Seed Laboratory,
Dry Branch, Georgia
Growth habit, occurrence, and use. The birchgenusBetulaconsists of about 40 to 50 species of deciduous trees and shrubs occurring in the cooler parts of the
Northern Hemisphere (Weaver 1978). Several species produce valuable lumber. Other species are useful for ornamental plantings because of their attractive growth habit, foliage,
and bark. Nearly all species provide food and cover for
wildlife, and some are valuable because they seed-in
promptly on harvested or burned lands. The 14 species
native to the Unites States are listed in table 1, along with
several species that are introduced or are referenced in the
seed literature.
Flowering and fruiting. The flowers are monoecious
and borne in catkins. Staminate catkins are formed in late
summer or autumn, remain naked during winter, and open
after considerable elongation in the spring (table 2). The pistillate catkins, which are cone-like with closely overlapping
scales, are born terminally on short, spur-like lateral branches and appear with the leaves (table 2). When the female
catkins (strobiles) ripen in late summer or autumn (table 2),
they become brown and woody and are either erect or pendulous (figure 1). Each scale may bear a single small,
winged nut (seed) (figures 2 and 3) that is oval, with 2 persistent stigmas at the apex. The seeds turn from greenish tan
to light brown or tan when mature (Brinkman 1974). Seeds
disperse from late fall until the following spring (Houle and
Payette 1990; Matlack 1989). Although seeds can begin to
disperse in late summer, these early-shed seeds may be of
poor quality. Seeds of yellow birch shed in August were
found to not be viable. Viable seeds were not released in
meaningful amounts until September, with the maximum of
good seeds being released in October (Houle and Payette
1990). After seedfall, the strobiles slowly disintegrate on the
trees, with the axes persisting on the branchlets.
Seed production. Birch tends to flower at the relatively young age of 10 to 15 years (Lepisto 1973) (table 3).
Betula
303
Common name(s)
Occurrence
B. alleghaniensis Britt.
B. lutea Michx. F.
B. borealis Spach
yellow birch
B. davurica Pall.
B. ermanii Cham.
Dahurian birch
Erman birch
B. lenta L.
northern birch
304
Murray birch
bog birch, swamp birch,
dwarf birch
Michigan
Newfoundland to Alaska, S to higher mtns
of California, Colorado, & Maine
Alaska birch
Common name
Occurrence
B. pubescens Ehrh.
B. alba L.
B. tortusa Ledeb.
B. pumila L.
B. pumila var. glandulifera
Regel (Gleason)
B. glandulifera (Regel) Butler
B. nana var. glandulifera (Regel) Boivin
B. glandulosa var. glandulifera
(Regel) Gleason
B. uber (Ashe) Fern.
Betula x utahensis Britt. (pro sp.)
B. andrewsii A. Nels.
B. piperi Britt.; B. ( commixta Sarg.
B. occidentalis var. fecunda Fern.
B. papyrifera var. subcordata
(Rydb.) Sarg.
downy birch
roundleaf birch
northwestern paper birch
Smyth Co.,Virginia
Yukon Territory, S through British Columbia,
Alberta, Saskatchewan,Washington, Idaho,
Montana, Oregon,Wyoming, & Utah
Source:
Brinkman (1974)
Location
Mid-range
Japan
Mid-range
Mid-range
N part of range
Mid-range
Russia & Finland
Mid-range
Germany & Finland
Mid-range
Flowering
Fruit ripening
Seed dispersal
AprMay
May
AprMay
JuneAug
AprMay
AprJune
AprJune
AprMay
MayJune
MayJune
AugOct
Oct
AugSept
AugOct
MayJune
AugSept
JulyAug
SeptOct
AugSept
SeptOct
SeptSpring
SeptNov
SeptMar
MayJune
AugSpring
JulySept
Oct to mid-winter
FallWinter
OctMar
Sources: Ahlgren (1957), Brinkman (1974), Damberg (1915), Fernald (1950), NBV (1946), Sarvas (1952),Van Dersal (1938),Wappes (1932).
directly into bags rather than allowed to fall onto the ground
or tarps, which can result in loss of the seeds. However,
seeds can also be collected from paved surfaces in urban
areas.
Seed extraction. Freshly collected strobiles can be
subject to heating because they usually are at least somewhat green. They should be spread out to dry for several
weeks until they begin to disintegrate. Low relative humidity
is the most important factor in drying the strobiles. Matlack
(1989) found that sweet birch strobiles released their seeds
at low humidity anywhere in the temperature range of 14 to
16 C. Once the strobiles begin to fall apart, they can be
shattered by rubbing or shaking, and the seeds can be separated from most of the scales and debris by screening and
fanning. Round-hole screens of the following sizes have
Betula
305
longitudinal section
Table 3Betula, birch: height, seed-bearing age, and seed crop frequency
Species
B. allenghaniensis
B. davurica
B. lenta
B. nana
B. nigra
B. papyrifera
B. pendula
B. populifolia
B. pubescens
B. pumila
Height at
maturity (m)
30
19.5
24
1.8
30
21
19.5
12
19.5
3
Year first
cultivated
Minimum
seed-bearing
age (yr)
1800
1883
1759
1880
1736
1750
Long
1750
1789
1762
40
40
15
15
8
15
Years between
large seedcrops
2
2
12
2
23
1
23
12
Species
B. alleghaniensis
B. davurica
B. lenta
B. nana
B. nigra
B. papyrifera
B. pendula
(de-winged)
(winged)
B. populifolia
B. pubescens
B. pumila
Seeds/fruit vol
kg/hl
lb/bu
1.34.5
2.69.4
1.03.5*
2.03.4*
Range
/kg
6121,995
1,5181,672
9752053
6,54711,253
6311,206
1,3429,064
278907
690760
443933
2,9765,115
287548
6104,120
990
1,595
1,421
8,446
825
3,036
450
725
646
3,839
375
1,380
24
2+
13
3
13
28
3,33211,088
1,6061,892
7,87810,846
1,6509,900
3,0727,634
1,5105,040
730860
3,5814,930
7504,500
1,3963,470
5,317
1,749
9,363
3,784
5,328
2,417
795
4,256
1,720
2,422
154+
10
2
45
4
Betula
307
Moisture
content (%)
Prechilling
Percent
(days)
germination
5.0
7.0
0
63
63
0
0
21
45
70
67
32
56
58
7.9
30
63
63
0
63
54
7.0
0
63
63
0
63
0
76
82
87
4
16
18
8.9
72
67
45
37
Prechill
period
(days)
300
None
None
4070
(over
winter)
None
3060
None
6075
None
3040
None
6090
3060
None
None
Daily
light
(hr)
8+
8+
8+
8+
20
8+
20
8+
8+
8+
8+
8+
8+
8+
20
Germination conditions
Temp (C)
Medium
Day
Night
Days
Sand
Sand
Sand
32
30
30
32
30
15
20
20
15
20
3040
1428
148
30
30
27
59
18
43
24
22
3
4
13
1
56
60
72
Perlite
Sand
Perlite
Sand
Paper pads
Sand
Sand
Perlite
24
30
24
32
30
30
30
25
24
18
20
18
15
30
30
30
3040
40
3040
40
30
30
30
3
34
73
47
30
36
64
40
87
31
5
13
35
6
10+
143
3
44
17
4
42
24
68
69
38
20
20
20
15
18
Germination
Avg
(%)
Samples
Purity
(%)
Sources: Black and Waring (1954, 1955), Brinkman (1974), Heit (1968), Gorshenin (1941),Yelenosky (1961).
Figure 4Betula populifoliaa, gray birch: seedling development at 1, 10, and 40 days after germination.
Betula
309
References
310
Johnson EA. 1975. Buried seed populations in the subarctic forest east of
Great Slave Lake, Northwest Territories. Canadian Journal of Botany 53:
29332941.
Komarova TA. 1986. Role of forest fires in germination of seeds dormant in
the soil. Soviet Journal of Ecology 16(6): 311-315.
Lepisto M. 1973. Accelerated birch breeding in plastic greenhouses. Forestry
Chronicle 49(4): 172-173.
Matlack GR. 1989. Secondary dispersal of seed across snow in Betula lenta, a
gap-colonizing tree species. Journal of Ecology 77: 853869.
Moore JM, Wein RW. 1977.Viable seed populations by soil depth and potential site recolonization after disturbance. Canadian Journal of Botany 55:
24082412.
Nagata H, Black M. 1977. Phytochrome-chilling interaction in the control of
seed dormancy of Betula maximowicziana Reg. Journal of the Japanese
Forestry Society 59(10): 368371.
Nagata H,Tsuda Y. 1975. Action of far-red and blue lights on the germination of Japanese white birch seed. Journal of the Japanese Forestry
Society 57(5): 160163.
NBV [Nederlandsche Boschbouw Vereeniging]. 1946. Boomzaden:
Handleiding inzake het oogsten, behandelen, bewaren en uitzaaien van
boomzaden. 171 p. Wageningen,The Netherlands: Ponsen and Looijen.
Odani K, Anma Y. 1986. Betula ermani seed germination regulated by gibberellin. Journal of the Japanese Forestry Society 68(12): 511513.
Patterson C F, Bruce AC. 1931. Rapid methods of determining the percentages of fertility and sterility in seeds of the genus Betula. Science
Agriculture (Ottawa) 11: 704708.
Perala DA, Alm AA. 1989. Regenerating paper birch in the lake states with
the shelterwood method. Northern Journal of Applied Forestry 6:
151153.
Pratt CR Jr. 1986. Environmental factors affecting seed germination of gray
birch (Betula populifolia) collected from abandoned anthracite coal mine
spoils in northeast Pennsylvania. Annals of Applied Biology 108:
649658.
Rafn J & Son. [nd, circa 1928]. Skovfrkontorets franalyser gennem 40 Aar,
18871927. Copenhagen. Udfrt paa Statsfrkontrollen i Kbenhavn.
5 p.
Sarvas R. 1952. On the flowering of birch and the quality of the seed crop.
Commonwealth Institute For. Fenn. 40(7): 138.
Scherbatskoy T, Klein RM, Badger GJ. 1987. Germination responses of forest
tree seed to acidity and metal ions. Environmental and Experimental
Botany 27(2): 157164.
Steijlen I, Zackrisson O. 1986. Long-term regeneration dynamics and successional trends in a northern Swedish coniferous forest stand. Canadian
Journal of Botany 65: 839848.
Vaartaja O. 1956. Photoperiodic response in germination of seed of certain
trees. Canadian Journal of Botany 34: 377388.
Van Dersal WR. 1938. Native woody plants of the United States: their
erosion control and wildlife values. Misc. Pub. 303. Washington, DC:
USDA 362 p.
Vanhatalo V, Leinonen K, Rita H, Nygren M. 1996. Effect of prechilling on the
dormancy of Betula pendula seeds. Canadian Journal of Forest Research
26: 12031208.
Wappes L. 1932. Wald und Holz ein Nachschlagebuch fr die Praxis der
Forstwirte, Holzhndler und Holzindustriellen.Vol. 1, Berlin: J. Neumann.
872 p.
Wearstler KA Jr., Barnes BV. 1977. Genetic diversity of yellow birch
seedlings in Michigan. Canadian Journal of Botany 55: 27782788.
Weaver RE Jr. 1978. The ornamental birches. Arnoldia 1978. 38(4):
117131.
Wheeler AG. 1976. Life history of Kleidocerys resedae on European white
birch and Ericaceous shrubs. Annals of Entomological Society of America
69(3): 459463.
Weinberger P, Vladut R. 1981. Comparative toxic effects of some xenobiotics
on the germination and early seedling growth of jack pine (Pinus
banksiana Lamb.) and white birch (Betula papyrifera Marsh.). Canadian
Journal of Forest Research 11: 796-804.
Weinberger P, Pomber L, Prasad R. 1978. Some toxic effects of fenitrothion
on seed germination and early seedling growth of jack pine, spruce, and
birches. Canadian Journal of Forest Research 8: 243246.
Yelenosky G. 1961. Birch seeds will germinate under a waterlight treatment
without pre-chilling. Res. Note 124. Upper Darby, PA: USDA Forest
Service, Northeast Forest Experiment Station. 5 p.
VerbenaceaeVerbena family
Callicarpa americana L.
American beautyberry
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Other common names. French-mulberry, Spanishmulberry, sour-bush, sow-berry.
Growth habit, occurrence, and uses. American
beautyberryCallicarpa americana L.is a small, woody
shrub of the pine forests in the southern coastal plain. It seldom grows taller than 2 or 3 m. The shrub is common
underneath the pine overstory and along roads and forest
edges, where it grows best. It is found from Virginia to
Florida and west to Texas and Oklahoma; it also occurs in
the West Indies (Vines 1960). American beautyberry is an
important food plant for wildlife, especially birds and eastern white-tailed deer (Odocoileus virginianus) (Blair and
Epps 1969; Grelen and Duvall 1966; Halls 1973). The
shrubs well-branched root system and drought resistance
make it desired for erosion control in some areas (Brown
1945), and it is frequently grown as an ornamental because
of the colorful fruits (Dirr and Heuser 1987).
Flowering and fruiting. The small, inconspicuous
flowers are borne in axillary, dichotomous cymes about 8 to
36 mm long. Flowering starts in early June and may continue into the fall months, even as the fruits mature in August
to November (Dirr and Heuser 1987; Vines 1960). The fruit
is a berrylike, globose drupe, about 3 to 6 mm in diameter,
that is borne in conspicuous axillary clusters on the current
seasons growth. The rose to purple, or sometimes white
(Brown 1945), fruit color gives this plant its ornamental
value. A single fruit cluster may contain as many as 300
fruits, although about 100 is typical. Each fruit usually contains 4 small flattened seeds that are light brown in color
and about 1 to 1.5 mm in length (Grelen and Duvall 1966;
Vines 1960) (figures 1 and 2). Plants begin to bear fruit as
early as 2 years of age, and mature plants may yield over
1/ kg (about 1 lb) annually (Halls 1973).
2
Collection of fruits; extraction and storage of seeds.
Fruits can be easily collected by hand in autumn, when their
rose to purple color indicates maturity. The soft fruits quickly disintegrate when they are macerated with water. Filled
Callicarpa
311
seeds sink in water, and the pulp can be floated off. Any type
of macerator should work, even laboratory or kitchen
blenders for small lots. There are about 600 seeds/g
(17,000/oz), and good cleaning should yield a purity of practically 100%. There are no known storage data for this
species, but soil seed bank studies show that the seeds will
survive for at least 1 year buried in the soil. This fact, plus
the hard seedcoat, suggest that these seeds are orthodox in
storage behavior. Long-term storage at temperatures near or
below freezing should be successful with seeds that are dried
to below 10% moisture content.
Pregermination treatments and germination tests.
The seeds have a hard seedcoat, and germination is relatively
slow. One sample stratified for 30 days yielded only 22%
germination in 90 days when tested at an alternating temperature of 20 C at night and 30 C in the light. Untreated
seeds sown in the fall, however, were reported to give excellent germination in the spring (Dirr and Heuser 1987). There
are no official test prescriptions for American beautyberry.
312
References
Blair RM, Epps EA Jr. 1969. Seasonal distribution of nutrients in plants of
seven browse species in Louisiana. Res. Paper SO-51. New Orleans:
USDA Forest Service, Southern Forest Experiment Station. 35 p.
Brown CA. 1945. Louisiana trees and shrubs. Bull. 1. Baton Rouge: Louisiana
Forestry Commission. 262 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Grelen HE, Duvall VL. 1966. Common plants of longleaf pinebluestem
range. Res. Pap. SO-23. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 96 p.
Halls LK. 1973. Flowering and fruiting of southern browse species. Res.
Paper SO-90. New Orleans: USDA Forest Service, Southern Forest
Experiment Station. 10 p.
Vines RA. 1960. Trees, shrubs and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
CupressaceaeCypress family
Calocedrus
313
the winter and early spring... (Sargent 1961). The inconspicuous pale yellow ovulate flowers also develop singly at
tips of twigs. Flowering has been reported to occur as early
as December and as late as May (Britton 1908; Hitchcock
and others 1969; Mitchell 1918; Peattie 1953; Sargent 1961;
Sudworth 1908), but it is not clear how well observers distinguished between flower appearance and actual pollen dissemination. Unopened staminate flowers and open or nearly
open ovulate flowers were present on branches collected in
the first week of April west of Klamath Falls, Oregon (Stein
1974).
Individual cones (figure 1), each containing up to 4
seeds, are scattered throughout the crown, and mature in a
single growing season. As they ripen, their color changes
from a medium green to a yellowish green or yellow tinged
with various amounts and shades of brown. During opening,
the cone becomes reddish brown and acquires a purplish
cast. Insect-attacked cones are among the first to change
color. Generally, cones of many color shades are found on a
tree as opening commences.
Seed dispersal may extend over a lengthy period, from
late August through November or later (Fowells and
Schubert 1956; McDonald 1992; Mitchell 1918; Powers and
Oliver 1990; Sudworth 1908). For example, in 1937 and
1940, respectively, 11 and 32% of the seed had fallen by
early October at 1 or 2 California locations, yet 47 and 66%
of the total fell after November 11 (Fowells and Schubert
1956). Cutting tests have shown that 14 to 65% of the naturally dispersed seeds appear sound, with higher values coincident with heavy crops (Fowells and Schubert 1956).
The oft-repeated generalization that incense-cedars bear
some seeds every year and abundant crops frequently (Betts
Figure 1Calocedrus decurrens, incense-cedar: cones
hang singly from branch tips well-dispersed over the crown
and contain up to 4 seeds each.
314
/kg
Average
/lb
Figure 3Calocedrus decurrens, incense-cedar: longitudinal section showing the radicle located at the narrow end of
the seed.
Range
/kg
/lb
Samples
27,270
12,368
24,82029,960
11,26013,588
41
31,820
14,433
21,54045,330
9,76820,562
36
33,420
15,160
24,12038,760
10,94017,583
Calocedrus
315
316
Figure 4Calocedrus decurrens, incense-cedar: germinating seed with radicle and hypocotyl emerging from the
winged end.
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New York: McGraw-Hill. 510 p.
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480 p.
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discovered pest of coniferous seed. Annals of the Entomological Society
of America 56: 229234.
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Little EL Jr. 1979. Checklist of United States trees, native and naturalized.
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
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deer herds, their ranges and management problems. Bull. 6. Sacramento:
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Martin AC, Zim HS, Nelson AL. 1951. American wildlife and plants. New
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Calocedrus
317
318
Schubert GH. 1954. Viability of various coniferous seeds after cold storage.
Journal of Forestry 52: 446447.
Schubert GH, Adams RS. 1971. Reforestation practices for conifers in
California. Sacramento: California Department of Conservation,
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Show SB. 1930. Forest nursery and planting practice in the California pine
region. Circ. 92. Washington, DC: USDA. 74 p.
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logging. Journal of Forestry 63: 456457, 460461.
Stein WI. 1963. Comparative juvenile growth of five western conifers [PhD
thesis]. New Haven, CT: Yale University. 194 p.
Stein WI. 1974. Libocedrus decurrens Torr., incense-cedar. In: Schopmeyer CS,
tech coord. Seeds of woody plants in the United States. Agric. Handbk.
450. Washington, DC: USDA Forest Service: 494499.
Stone EC. 1957. Dew as an ecological factor: 2.The effect of artificial dew
on the survival of Pinus ponderosa and associated species. Ecology 38:
414422.
Sudworth GB. 1900. The forest nursery: collection of tree seeds and propagation of seedlings. Bull. 29. Washington, DC: USDA Division of Forestry.
63 p.
Sudworth GB. 1908. Forest trees of the Pacific slope. Washington, DC:
USDA Forest Service. 441 p.
Tevis L Jr. 1953. Stomach contents of chipmunks and mantled squirrels in
northeastern California. Journal of Mammalogy 34: 316324.
Tillotson CR. 1925. Growing and planting coniferous trees on the farm.
Farm. Bull. 1453. Washington, DC: USDA. 38 p.
Toumey JW, Korstian CF. 1942. Seeding and planting in the practice of
forestry. 3rd ed. New York: John Wiley & Sons. 520 p.
Van Dersal WR. 1938. Native woody plants of the United States, their erosion-control and wildlife values. Misc. Pub. 303. Washington, DC: USDA.
362 p.
van Pelt R. 2001. Forest giants of the Pacific Coast. Seattle: University of
Washington Press: 8085.
BignoniaceaeTrumpet-creeper family
Campsis
319
References
Bertin RI. 1990. Effects of pollination intensity in Campsis radicans.
American Journal of Botany 77: 178187.
Bertin RI, Sullivan M. 1988. Pollen interference and cryptic self-fertility in
Campsis radicans. American Journal of Botany 75: 11401147.
Bonner FT. 1974. Campsis radicans, common trumpetcreeper. In:
Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service:
260261.
Dirr MA, Heuser CWJr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
320
FabaceaePea family
seed.
Caragana
321
322
germinators for 14 to 60 days at the same alternating temperatures (Dietz and Slabaugh 1974; Hamm and Lindquist
1968). Germination after 25 to 41 days averaged 45 to 72%,
and 55 to 100% after 60 days (Dietz and Slabaugh 1974).
Nursery practice. Seeds of Siberian peashrub may be
drilled or broadcast in late summer or spring. In a North
Dakota nursery, Siberian peashrub is seeded during the last
week in July or the first week in August. A cover crop of
oats is seeded between the tree rows early enough to give
winter protection. The shrubs are large enough to dig the
following fall (Dietz and Slabaugh 1974). Many nurseries
recommend drilling 80 to 160 seeds/m (25 to 50/ft) at 6, 9,
or 12 mm (1/4 to 1/2 in) depth; percentages of seeds growing
into seedlings have varied from 35 to 50 (Dietz and
Slabaugh 1974; Lindquist and Cram 1964).
Grading seeds for size has greatly increased the percentage of plantable seedlings. To be plantable, seedlings should
be 30 cm (12 in) or more in height at the time of lifting.
Only 87% of the seedlings grown from seeds measuring
2.5 mm in diameter were plantable, whereas 77% of seeds
measuring 4.0 mm in diameter were plantable (Lindquist
and Cram 1967). Inoculation of seeds with Rhizobium ssp.
before sowing has been recommended (Wright 1947), but
other workers report no significant effect on 1+0 seedlings
(Cram and others 1964). Commercial nurseries have recommended anywhere from 1+0 to 3+0 stock for outplanting
(Dietz and Slabaugh 1974).
Spraying to control insects in the nursery may be necessary. Grasshoppers are especially destructive to Siberian
peashrub, sometimes completely defoliating the plants
(Kennedy 1968). Plants have also been extensively damaged
by deer browsing.
Vegetative propagation of Siberian peashrub is also possible. Untreated cuttings taken in late July rooted 80% in
sand, whereas cuttings taken earlier (May to June) responded well to indole-butyric acid (IBA) in talc (Dirr and Heuser
1987).
References
Cram WH. 1956. Research. In: 1956 summary report of the Forest
Nursery Station. Indian Head, SK: Canadian Department of Agriculture,
Experimental Farms Service: 9394.
Cram WH. 1969. Breeding and genetics of Caragana. Forestry Chronicle
45(6): 400401.
Cram WH,Thompson AC, Lindquist CH. 1964. Nursery production investigations. In: 1964 Summary report for the tree nursery. Indian Head, SK:
Canadian Department of Agriculture, Prairie Farm Rehabilitation
Administration: 1925.
Dietz DR, Slabaugh PE. 1974. Caragana arborescens Lam., Siberian
peashrub. In: Schopmeyer CS, tech. coord. Seeds of woody plants in the
United States. Agric. Handbk. 450. Washington, DC: USDA Forest
Service: 262264.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
George EJ. 1953. Tree and shrub species for the northern Great Plains.
Agric. Circ. 912. Washington, DC: USDA. 46 p.
Graham EH. 1941. Legumes for erosion control and wildlife. Misc. Pub. 412.
Washington, DC: USDA. 153 p.
Hamm JW, Lindquist CH. 1968. Research for production and distribution
programs: performance and propagation. In: 1968 summary report for
the Tree Nursery. Indian Head, SK: Canadian Department of Agriculture,
Prairie Farm Rehabilitation Administration: 1220.
ISTA [International Seed Testing Association]. 1993. Rules for testing seeds:
rules 1993. Seed Science and Technology 21 (Suppl.): 1259.
Kelsey HP, Dayton WA. 1942. Standardized plant names. 2nd ed.
Harrisburg, PA: J. Horace McFarland Co. 675 p.
Kennedy PC.1968. Insects and diseases of Siberian peashrub (Caragana) in
North Dakota, and their control. Res. Note RM-104. Ft. Collins, CO:
USDA Forest Service, Rocky Mountain Forest and Range Experiment
Station. 4 p.
Lindquist CH. 1960. Notes on the moisture requirements of the stratifying
media for the seed of Caragana arborescens Lam. Canadian Journal of
Plant Science 40: 576577.
Lindquist CH, Cram WH. 1964. Tree improvement and propagation studies.
In: 1964 summary report for the tree nursery. Indian Head, SK: Canadian
Department of Agriculture, Prairie Farms Rehabilitation Administration:
1519.
Lindquist CH, Cram WH. 1967. Propagation and disease investigations. In:
1967 summary report for the tree nursery. Indian Head, SK: Canadian
Department of Agriculture, Prairie Farms Rehabilitation Administration:
2126.
Rehder A. 1940. Manual of cultivated trees and shrubs. 2nd ed. New York:
Macmillan. 966 p.
Ross NM. 1931. Tree-planting on the prairies of Manitoba, Saskatchewan,
and Alberta. Bull. 1, 8th ed. Ottawa: Canadian Department of Interior,
Forest Service. 64 p.
Wright PH. 1947. Caragana needs inoculation.Your Garden and Home
[Toronto] 1(5): 19.
Caragana
323
324
seeds.
longitudinal
References
Alcorn SL, Martin SC. 1974. Cereus giganteus Engelm., saguaro. In:
Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service:
313314.
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
Despain DG. 1974. The survival of saguaro (Carnegiea gigantea) seedlings
on soils of differing albedo and cover. Journal of the Arizona Academy of
Science 9: 102107.
Hevly RH. 1979. Dietary habits of two nectar and pollen feeding bats in
southern Arizona and northern Mexico. Journal of the ArizonaNevada
Academy of Science 14: 1318.
Kearney TH, Peebles RH. 1960. Arizona flora. Berkeley: University of
California Press. 1085 p.
Munz PA. 1974. A flora of southern California. Berkeley: University of
California Press. 1086 p.
Nobel PS. 1980. Morphology, nurse plants, and minimum apical temperatures
for young Carnegiea gigantea. Botanical Gazette 141: 188191.
Steenbergh WF, Lowe CH. 1969. Critical factors during the first years of life
of the saguaro (Cereus giganteus) at Saguaro National Monument,
Arizona. Ecology 50: 825834.
Steenbergh WF, Lowe CH. 1976. Ecology of the saguaro: 1.The role of
freezing weather in a warm desert population. In: Research in the parks.
Symp. Ser. 1. Washington DC: USDI National Park Service: 4992.
Steenbergh WF, Lowe CH. 1977. Ecology of the saguaro: 2. Reproduction,
germination, establishment, growth, and survival of the young plant. Sci.
Monogr. Ser. 8. Washington, DC: USDI National Park Service. 248 p.
Steenbergh WF, Lowe CH. 1983. Ecology of the saguaro: 3. Growth and
demography. Sci. Monogr. Ser. 17. Washington, DC: USDI National Park
Service: 228 p.
Carnegiea
325
HydrangeaceaeHydrangea family
Growth habit. Carpenteria (bush-anemone or treeanemone)Carpenteria californica Torr.is an erect evergreen shrub that is 1 to 3 (sometimes 4 m) tall with large
showy white flowers. Plants are usually multi-stemmed and
about as wide as tall. The leaves are oblonglanceolate on
short petioles and placed opposite. The leaves are leathery
and in response to moisture stress, they turn yellow and
twist and their edges roll under. They return to normal
appearance and color when moisture is available (Kottcamp
1983; Sanwo 1997).
Occurrence. The range of carpenteria is extremely
limited, occurring only between 300 and 1,500 m on the
west slope of the Sierra Nevada, between the San Joaquin
and Kings Rivers in eastern Fresno County, California. The
shrub is found in scattered stands over an area 20 by 30 km
or about 60,000 ha. The total number of plants has been
estimated to less than 5,000 (Clines 1995).
Most stands are in small drainages on dry rocky slopes,
mixed with Digger pine (Pinus sabiniana Dougl. ex Dougl.),
interior live oak (Quercus wislizeni A. DC.), chaparral
whitethorn (Ceanothus leucodermis Greene), and other representatives of the foothill woodlands at the lower elevational limits of its range. At their upper elevational limit, carpenteria plants can be found growing with ponderosa pine
(Pinus ponderosa P. & C. Lawson), interior live oak, and
other species of the lower yellow pine zone.
About two-thirds of the existing plants occur on lands of
the USDA Forest Services Sierra National Forest, and carpenteria is classified as a sensitive plant by the Forest
Service. It receives some protection on 2 areas set aside by
the Sierra National Forest and 1 owned by The Nature
Conservancy. Carpenteria is a threatened species under the
California Endangered Species Act, and in October of 1994
it was proposed for listing as endangered under the Federal
Endangered Species Act (Federal Register 1994).
Natural reproduction. Until recently, all observed
natural reproduction was by stump-sprouting after fire
(Stebbins 1988; Wickenheiser 1989). However, after a large
wildfire in 1989, an abundance of naturally occurring
326
References
Cheatham DH. 1974. Carpenteria, the mystery shrub. Fremontia 1(4): 38.
Clines J. 1994. Reproductive ecology of Carpenteria californica
(Philadelphaceae) [MA thesis]. Fresno: California State University. 80 p.
Clines J. 1995. Personal communication. Clovis, CA.
Federal Register 1994. 59(191): 5054050549. October 4.
Kottcamp K. 1983. Water relations of Carpenteria californica [MA thesis].
Fresno: California State Univeristy. 80 p.
Laclergue R. 1995. Personal communication. Auberry, CA.
Mirov NT, Kraebel CJ. 1939. Collecting and handling seeds of wild plants.
Pub. 5. Washington, DC: USDA Civilian Conservation Corps. 42 p.
Sanwo MM. 1987. Leaf anatomy and pigment analysis of Carpenteria
california [MA thesis]. Fresno: California State University. 111 p.
Stebbins J. 1988. A petition to the State of California Fish and Game
Commission to list tree anemone (Carpenteria californica). 8 p.
Wickenheiser LP. 1989. Report to the Fish and Game Commission on
the status of tree anemone (Carpenteria californica). 19 p.
seed.
Carpenteria
327
BetulaceaeBirch family
Carpinus L.
hornbeam or ironwood
Paula M. Pijut
Dr. Pijut is a plant physiologist at the USDA Forest Services North Central Research Station,
Hardwood Tree Improvement and Regeneration Center,West Lafayette, Indiana
Growth habit, occurrence, and use. The hornbeam
genusCarpinus L.includes about 35 species of deciduous, monoecious, small to large trees, that are native to the
Northern Hemisphere from Europe to eastern Asia, south to
the Himalayas, and in North and Central America (Furlow
1990; Hillier 1991; Krssmann 1984; LHBH 1976; Suszka
and others 1996). Five species are considered here (table
1). Hornbeams occur mainly as understory trees in rich,
moist soils on bottomlands and on protected slopes
(Metzger 1990; Rudolf and Phipps 1974). European hornbeam is an important forest tree species throughout Europe
(Furlow 1990). In Mexico and Central America, Carpinus
tropicalis (J.D. Sm.) Lundell forms a dominant canopy
component (Furlow 1990). American hornbeams, which are
native to the eastern United States and Canada, are smaller
trees that grow in the mixed hardwood forest understory
(Furlow 1990; Metzger 1990). Several geographic races of
American hornbeam exist in North America (Fernald 1935;
Furlow 1990). The races are morphologically variable and
difficult to distinguish on the basis of independent characters. Furlow (1987a), using multivariate analysis, analyzed
this geographical variation. The northern American hornbeam species is divided into the subsp. caroliniana from
along the Atlantic and Gulf Coastal Plains of the southeastern United States and the subsp. virginiana of the
Table 1Carpinus, hornbeam: nomenclature, occurrence, height at maturity, and date of first cultivation
Scientific name
Common name(s)
Occurrence
C. betulus L.
C. caroliniana Walt.
European hornbeam
American hornbeam,
musclewood, blue
beech, ironwood
heartleaf hornbeam
Japanese hornbeam
Oriental hornbeam
C. cordata Blume
C. japonica Blume
C. orientalis Mill.
Sources: Dirr (1990), Hillier (1991), Krssmann (1984), LHBH (1976), Metzger (1990).
328
Height at
maturity (m)
Year first
cultivated
1221
69
1800s
1812
615
69
68
1879
1895
1739
Carpinus
329
Location
Flowering
Fruit ripening
Seed dispersal
C. betulus
C. caroliniana
Europe & NE US
NE US
AprMay
MarJune
AugNov
AugOct
Novspring
Novspring
Table 3Carpinus, hornbeam: seed-bearing age, seedcrop frequency, seed weight, and fruit ripeness criteria
Species
Minimum
seed-bearing
age (yrs)
Years between
large seedcrops
C. betulus
C. caroliniana
1030
15
12
35
Average no.
cleaned seeds
/kg
/lb
28,660
66,138
13,000
30,000
Preripe
color
Green
Green
Ripe
color
Brown
Greenish brown
Species
20
20
20
2030
20
C. betulus
C. caroliniana
C. orientalis
Cold period
Temp
(C)
Days
28
14
30
60
60
35
5
4
5
4.5
5
C
Percentage
germination
90112
210
120
60
126
90120
NS
65
65
10
58
NS
Sources: Allen (1995), Blomme and Degeyter (1977), Bretzloff and Pellet (1979), Bugala (1993), Rudolf and Phipps (1974), Suszka and others (1996).
NS = not stated.
Table 5Carpinus, hornbeam: germination test conditions and results with stratified seed
Species
C. betulus
C. caroliniana
Test conditions*
Temp (C)
Day
Night Days
20
27
20
16
70
60
Germination rate
%
Days
30
2
7
12
% Germination
Avg
Samples
1890
15
50
2
Purity Soundness
(%)
(%)
97
96
60
62
Carpinus
331
References
Allen DH. 1995. Personal communication. Sandwich, MA: F.W. Schumacher.
Bassuk N, Miske D, Maynard B. 1985. Stock plant etiolation for improved
rooting of cuttings. Combined Proceedings of the International Plant
Propagators Society 34: 543550.
Beckett JL. 1994. American hornbeam (Carpinus caroliniana). Morton
Arboretum 30(2): 2325.
Blomme R, Degeyter L. 1977. Problemen bij de kieming vanzaden van
Carpinus betulus (Haagbeuk) [in Dutch: Problems with the germination
of seeds of Carpinus betulus].Verbondsnieuws voor de Belgische Sierteelt
21(13): 429432.
Bretzloff LV, Pellet NE. 1979. Effect of stratification and gibberellic acid on
the germination of Carpinus caroliniana Walt. HortScience 14(5):
621622.
Bugala W, ed. 1993. Grab zwyczajny: Carpinus betulus L. In: Nasze Drzewa
Lesne [in Polish: chapter summaries in English]. Monogr. Pop. 9. Kornik:
Polish Academy of Sciences, Institute of Dendrology. 352 p.
Cesarini J. 1971. Propagation of Carpinus betulus Fastigiata. Combined
Proceedings of the International Plant Propagators Society 21: 380382.
Chalupa V. 1990. Micropropagation of hornbeam (Carpinus betulus L.) and
ash (Fraxinus excelsior L.). Biologia Plantarum 32(5): 332338.
Chavagnat A. 1978. Utilisation des tests topographiques au ttrazolium
pour dterminer la viabilit des semences darbres et darbustes dornement [in French:The use of topographical tetrazolium tests to determine
the seed viability of ornamental trees and shrubs]. LHorticulture
Francaise 95: 36.
Davies RJ. 1987. A comparison of the survival and growth of transplants,
whips and standards, with and without chemical weed control.
Arboricult. Res. Note 67. Surrey: UK Department of the Environment.
5 p.
Davies RJ. 1988. Sheet mulching as an aid to broadleaved tree establishment: 1.The effectiveness of various synthetic sheets compared. Forestry
(Journal of the Institution of Chartered Foresters, London) 61(2):
89-105.
Dirr MA. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation, and uses. Champaign, IL:
Stipes Publishing Co. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Fernald ML. 1935. Midsummer vascular plants of southeastern Virginia.
Rhodora 37: 378413, 423454.
Furlow JJ. 1987a. The Carpinus caroliniana complex in North America: 1. A
multivariate analysis of geographical variation. Systematic Botany 12(1):
2140.
Furlow JJ. 1987b. The Carpinus caroliniana complex in North America: 2.
Systematics. Systematic Botany 12(3): 416434.
Furlow JJ. 1990. The genera of Betulaceae in the southeastern United
States. Journal of the Arnold Arboretum 71(1): 167.
Gordon AG, Gosling P, Wang BSP. 1991. Tree and shrub seed handbook.
Zurich: International Seed Testing Association. 9.19.19.
332
Hartmann HT, Kester DE, Davies FT. 1990. Plant propagation: principles and
practices. Englewood Cliffs, NJ: Prentice Hall. 647 p.
Hillier Nurseries (Winchester) Ltd. 1991. The Hillier manual of trees and
shrubs. Melksham, Wiltshire, UK: Redwood Press. 704 p.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing. Rules 1993. Seed Science and Technology 21 (Suppl.):
1259.
Krssmann G. 1984. Manual of cultivated broad-leaved trees and shrubs.
Vol. 1, AD. Beaverton, OR:Timber Press. 448 p.
Leiss J. 1985. Seed treatments to enhance germination. Combined
Proceedings of the International Plant Propagators Society 35: 495499.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third, a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan. 1290 p.
Macdonald B. 1986. Practical woody plant propagation for nursery
growers. Portland, OR:Timber Press. 669 p.
MacMillan-Browse P. 1974. No system is foolproof in propagating hornbeam. Nurseryman and Garden Centre 159(21): 649, 651652.
Maynard BK, Bassuk NL. 1987. Stockplant etiolation and blanching of
woody plants prior to cutting propagation. Journal of the American
Society for Horticultural Science 112(2): 273276.
Maynard BK, Bassuk NL. 1991. Stock plant etiolation and stem banding
effect on the auxin dose-response of rooting in stem cuttings of
Carpinus betulus L. Fastigata. Plant Growth Regulation 10: 305311.
Maynard BK, Bassuk NL. 1992. Stock plant etiolation, shading, and banding
effects on cutting propagation of Carpinus betulus. Journal of the
American Society for Horticultural Science 117(5): 740744.
Maynard BK, Bassuk NL. 1996. Effects of stock plant etiolation, shading,
banding, and shoot development on histology and cutting propagation of
Carpinus betulus L. fastigata. Journal of the American Society for
Horticulural Science 121(5): 853860.
Metzger FT. 1990. Carpinus caroliniana Walt. In: Burns RM, Honkala BH, tech.
coord. Silvics of North America,Volume 2, Hardwoods. Agric. Handbk.
654. Washington, DC: USDA Forest Service: 179185.
Obdrzalek J. 1987. Produkce mladych rostlin listnatych stromu a kvetoucich
keru z letnich rizku ve foliovych krytech [in Czech, summary in English:
Production of young plants of broad- leaved trees and flowering shrubs
from summer cuttings in plastic houses]. Forestry Abstracts 1988. 49(8):
5003.
Rudolf PO, Phipps H. 1974. Carpinus, hornbeam. In: Schopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 266268.
Suszka B, Muller C, Bonnet-Masimbert M. 1998. Seeds of forest broadleaves: from harvest to sowing. Paris: Institut National de la Recerche
Agronomique. 294 p.
Vrgoc P. 1994. Fraxinus rotundifolia Mill., Pinus nigra ssp. austriaca (Hoess)
Vid.; Carpinus orientalis Mill. u bonsai kultur [in Croatian; figures, tables,
and summary in English]. Glasnik za Sumske Rokuse 31: 135197.
JuglandaceaeWalnut family
Carya Nutt.
hickory
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Common name
Occurrence
Carya
333
Flowering
Fruit ripening
Seed dispersal
C. alba
C. aquatica
C. cordiformis
C. glabra
C. illinoensis
C. laciniosa
C. myristiciformis
C. ovata
C. pallida
AprMay
MarMay
AprMay
AprMay
MarMay
AprJune
AprMay
AprJune
MarApr
SeptOct
SeptNov
SeptOct
SeptOct
SeptOct
SeptNov
SeptOct
SeptOct
SeptOct
SeptOct
OctDec
SeptDec
SeptOct
SeptOct
SeptOct
SeptOct
SeptOct
SeptOct
334
Table 3Carya, hickory: height, seed-bearing age, seedcrop frequency, and year first cultivated
Species
Height at
maturity (m)
Year first
cultivated
Minimum
seed-bearing
age (yrs)
Years between
seedcrops
30
30
1530
2427
3443
37
2430
2130
12-30
1766
1800
1689
1750
1766
1800
1911
25
20
30
30
1020
40
30
40
23
12
35
12
12
12
23
13
23
C. alba
C. aquatica
C. cordiformis
C. glabra
C. illinoensis
C. laciniosa
C. myristiciformis
C. ovata
C. pallida
Species
C. alba
C. aquatica
C. cordiformis
C. glabra
C. illinoensis
C. laciniosa
C. myristiciformis
C. ovata
Source:
Mississippi
Mississippi
Mississippi
Mississippi
Texas
Mississippi
& Arkansas
Wisconsin
Mississippi
Fruits/vol
/hl
/bu
Range
/kg
/lb
Average
/kg
/lb
5,040
10,100
20,800
14,500
1,776
3,552
7,330
5,110
57
51
51
44
40
40
75249
71106
305419
275408
386496
121353
333384
5577
207375
34113
3248
138140
125185
175225
55160
151174
2535
94170
200
79
360
344
441
143
220
357
311
66
273
90
36
164
156
200
65
100
162
141
30
124
17,600
12,100
6,200
4,264
3849
3038
176331
80150
220
291
207
100
32
94
Carya
335
Table 5Carya, hickory: stratification period, germination test conditions, and results
Species
C. alba
C. aquatica
C. cordiformis
C. glabra
C. illinoensis
C. laciniosa
C. myristiciformis
C. ovata
Cold
stratification
(days)
90150
3090
90
90
90120
3090
3090
30
90120
60120
90150
60120
Days
Germination
Rate
(%)
Days
93
63
250
50
3045
4560
60
3597
4560
60
4560
60
54
76
40
60
80
53
75
65
Temp (C)
Day
Night
30
2732
30
27
30
30
30*
32
30
30*
30
30*
20
21
20
21
20
20
20
21
20
20
20
20
64
28
30
50
33
50
40
35
Germination %
Avg
(%)
Samples
66
92
55
60
85
50
91
75
60
80
73
4
1
3
1
2
9
6
2
2
6
2
References
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management. Louisiana Agriculture 20(2): 1415.
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Journal of Seed Technology 16 (3): 1113.
Bonner FT. 1971. Chemical contents of southern hardwood fruits and
seeds. Res. Note SO-136. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 3 p.
Bonner FT. 1974. Chemical components of some southern fruits and seeds.
Res. Note SO-183. New Orleans: USDA Forest Service, Southern Forest
Experiment Station. 3 p.
Bonner FT. 1976a. Effects of gibberellin on germination of forest tree seeds
with shallow dormancy. In: Hatano I, Asakawa S, Katsuta M, Sasaki S,
Yokoyama T, eds. Proceedings, 2nd International Symposium on
Physiology of Seed Germination; 1976 October 1830; Fuji, Japan.Tokyo:
Government Forest Experiment Station: 2132.
336
Madden GD, Malstrom HL. 1975. Pecans and hickories. In: Janick J, Moore
JN, eds. Advances in fruit breeding. West Lafayette, IN: Purdue University
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Mexico). 2nd ed., corrected and reprinted. New York: Dover Publications.
934 p.
Short HL, Epps EA Jr. 1976. Nutrient quality and digestibility of seeds and
fruits from southern forests. Journal of Wildlife Management 40:
283289.
van Staden J, Dimalla GG. 1976. Regulation of germination of pecan, Carya
illinoensis. Zeitschrift fr Pflanzenphysiologie 78: 6675.
Williams RD, Hanks SH. 1976. Hardwood nurserymans guide. Agric.
Handbk. 473. Washington, DC: USDA Forest Service. 78 p.
Yates IE, Reilly CC. 1990. Somatic embryogenesis and plant development in
eight cultivars of pecan. HortScience 25: 573576.
Yates IE, Sparks D. 1990. Three-year-old pecan pollen retains fertility. Journal
of the American Society for Horticultural Science 115: 359363.
Carya
337
FagaceaeBeech family
Castanea P. Mill.
chestnut
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Common name(s)
Occurrence
Japanese chestnut
American chestnut
Chinese chestnut
Allegheny chinkapin
European chestnut,
Spanish chestnut
Japan
S Maine to Michigan; S to S Mississippi & Georgia
China & Korea
Pennsylvania S to central Florida & W to E Texas & Oklahoma
S Europe,W Asia, & N Africa
338
Table 2Castanea, chestnut: height, year first cultivated, and seed weights
Species
C. crenata
C. dentata
C. mollissima
C. pumilla
C. sativa
Height at
maturity (m)
10
2025*
21
15
21
Pre 1880
C
Cleaned seeds/weight
/kg
33
220360
50220
300
33
/lb
15
100162
23100
136
15
fruit
Castanea
339
References
Ackerman WL, Jayne HT. 1980. Budding the epicotyls of sprouted chestnut
seeds. HortScience 15: 186187.
Aldous JR. 1972. Nursery practice. For. Comm. Bull. 43. London: Her
Majestys Stationery Office. 184 p.
ACF [American Chestnut Foundation]. 2002. Bennington,VT: ACF [website
available at www.acf.org/
].
Anagnostakis SL. 1990. An historical reference for chestnut introductions
into North America. Northern Nut Growers Annual Report (1989) 80:
132143.
Berry FH. 1960. Germination of Chinese chestnut seed. Northern Nut
Growers Association Annual Report 51: 4042.
Brown CL, Kirkman LK. 1990. Trees of Georgia and adjacent states.
Portland, OR:Timber Press. 292 p.
340
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Gibson LP. 1985. Description and key to larvae of Curculio spp. of eastern
United States and Canada (Coleoptera: Curculionidae). Proceedings of
the Entomological Society of Washington 87: 554563.
Hardy MB. 1948. Chestnut growing in the Southeast. Northern Nut
Growers Association Annual Report 39: 4050.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing. Rules 1993. Seed Science and Technology 21 (Suppl.):
1259.
Jaynes RA. 1975. Chestnuts. In: Janik J, Moore JN, eds. Advances in fruit
breeding. West Lafayette, IN: Purdue University Press: 490503.
Little EL, Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Payne JA, Wells JM. 1978. Postharvest control of the small chestnut weevil
in inshell chestnuts. Journal of Economic Entomology 71: 894895.
Payne JA, Miller G, Johnson GP, Senter SD. 1994. Castanea pumila (L.) Mill.:
an underused native nut tree. HortScience 29: 62, 130131.
Pritchard HW, Manger KR. 1990. Quantal response of fruit and seed germination rate in Quercus robur L. and Castanea sativa Mill. to constant temperatures and photon dose. Journal of Experimental Botany 41:
15491557.
Rehder A. 1940. Manual of cultivated trees and shrubs hardy in North
America. 2nd. ed. New York: Macmillan. 996 p.
Sander IL. 1974. Castanea, chestnut. In: Schopmeyer CS, tech. coord. Seeds
of woody plants in the United States. Agric. Handbk. 450. Washington,
DC: USDA Forest Service: 273275.
Castanea
341
CasuarinaceaeCasuarina family
Casuarina Rumph. ex L.
casuarina
David F. Olson, Jr., E. Q. P. Petteys, and John A. Parrotta
Dr. Olson retired from the USDA Forest Service; Dr. Petteys is the district forester of
Kauai, Hawaii Department of Lands and Natural Resources, Division of Forestry and Wildlife;
Dr. Parrotta is a research program leader at the USDA Forest Services Research and Development National
Office, Arlington,Virginia
Growth habit, occurrence, and use. The genus
Casuarinathe only genus in the Casuarina familycomprises about 50 species, chiefly Australian, with a few having native ranges extending from Bangladesh to Polynesia.
Casuarina trees are evergreen angiosperms, resembling
conifers, with thin crowns of drooping branches and leaves
reduced to scales (Little 1949; Little and Wadsworth 1964).
Three species of this genus have been introduced successfully into continental United States, Hawaii, and Puerto Rico
(table 1) (Bailey 1949; Rockwood and others 1990). Beach
she-oak, especially, is planted as a windbreak throughout its
native and introduced ranges and as an ornamental in parks
and gardens (Parrotta 1993; Rockwood and others 1990). It
was first introduced into Hawaii in 1882 (Neal 1965). The
bark has been used in tanning, in medicine, and for the
extraction of dye (Parrotta 1993). The fruits are made into
novelties and Christmas decorations (Little and Wadsworth
1964). The wood is hard and heavy and is difficult to work,
hence the common name ironwood. It was once heavily
used for building poles and firewood but now is seldom used
commercially in the United States (Parrotta 1993). Beach
and gray she-oaks are considered invasive pests in southern
Florida and gray she-oak in Hawaii.
Common name(s)
C. cunninghamiana Miq.
C. tenuissima Hort.
C. equisetifolia L.
C. litorea L.
Casuarina glauca Sieb. ex Spreng.
342
Occurrence
(native & introduced)
Australia & New Caledonia;
Hawaii, S US, & California
Burma through Australia & Polynesia;
Hawaii, Florida, & Puerto Rico
Australia; Hawaii
plastic sheets. Seeds reach maximum weight and germinability 18 weeks after anthesis, or when cones change in
color from green to brown (Rai 1990). The samaras, which
are used as seeds, may be separated from the fruits by shaking and screening (Olson and Petteys 1974). Cones placed
in trays, covered by a thin cloth, and dried under full sunlight will soon begin to release their seeds, usually within 3
Casuarina
343
References
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing
seeds. Journal of Seed Technology 16(3): 1113.
Bailey LH. 1949. Manual of cultivated plants most commonly grown in the
continental United States and Canada. New York: Macmillan. 1116 p.
Halos SC. 1983. Casuarinas in Philippine forest development. In: Midgeley
SJ;Turnbull, RD, Johnston RD, eds. Casuarina Ecology, Management and
Utilization: Workshop Proceedings; 1990 January 1520; Cairo, Egypt.
Cairo: American University in Cairo, Desert Development Center:
102109.
Jones L. 1967. Effect of storage at various moisture contents and temperatures on seed germination of silk oak, Australian pine, and Eucalyptus
spp. Res. Note SE-83. Asheville, NC: USDA Forest Service, Southeastern
Forest Experiment Station. 4 p.
Kondas S. 1990. Casuarina equisetifolia: a multipurpose tree cash crop in
India. In: Midgeley SJ;Turnbull, RD; Johnston RD, eds. Casuarina Ecology,
Management and Utilization: Workshop Proceedings; 1990 January
1520; Cairo. Cairo: American University in Cairo, Desert Development
Center: 6676.
Little EL Jr. 1949. Fifty trees from foreign lands. In:Yearbook of Agriculture
(1949). Washington, DC: USDA: 815832.
Little EL Jr; Wadsworth FH. 1964. Common trees of Puerto Rico and the
Virgin Islands. Agric. Handbk. 249. Washington, DC: USDA Forest
Service. 548 p.
344
Magini E;Tulstrup NP. 1955. Tree seed notes. For. Dev. Pap. 5. Rome: FAO.
354 p.
Neal MC. 1965. In gardens in Hawaii. Spec. Pub. 50. Honolulu: Bishop
Museum Press. 924 p.
Olson DF Jr, Petteys EQP. 1974. Casuarina, casuarina. In: Schopmeyer CS,
tech. coord. Seeds of woody plants in the United States. Agric. Handbk.
450. Washington, DC: USDA Forest Service: 278280.
Parrotta JA. 1993. Casuarina equisetifolia L. ex J.R. & G. Forst.: casuarina,
Australian pine. Res. Note SO-ITF-SM-56. New Orleans: USDA Forest
Service, Southern Forest Experiment Station. 11 p.
Rai RSV. 1990. Seed management in Casuarina equisetifolia. In: Midgeley SJ;
Turnbull, RD, Johnston RD, eds. Casuarina Ecology, Management and
Utilization: Workshop Proceedings; 1990 January 1520; Cairo. Cairo:
American University in Cairo, Desert Development Center: 7884
Rockwood DL, Fisher RF, Conde LF, Huffman JB. 1990. Casuarina L. ex
Adans. In: Burns RM, Honkala BH, tech. coord. Silvics of North America.
Volume 2, Hardwoods. Agric. Handbk. 654. Washington, DC: US
Department of Agriculture: 240243.
Turnbull JW, Martensz PN. 1990. Seed production, collection and germination in Casuarinaceae. In: Midgeley SJ;Turnbull, RD, Johnston RD, eds.
Casuarina Ecology, Management and Utilization: Workshop Proceedings;
1990 January 1520; Cairo. Cairo: American University in Cairo, Desert
Development Center: 126132.
BignoniaceaeTrumpet-creeper family
Catalpa Scop.
catalpa
Franklin T. Bonner and David L. Graney
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station, Mississippi State,
Mississippi; Dr. Graney is a research forester at the USDA Forest Services
Southern Research Station, Fayetteville, Arkansas
Sources:
Common name(s)
Occurrence
Year first
cultivated
1726
1754
Catalpa
345
seed.
longitudinal
346
Species
Place
collected
C. bignonioides
Florida
C. longissima
C. speciosa
Puerto Rico
Minnesota
Prairie states
Wt yield of
seeds/
100wt fruit
35
1025
2535
C
Cleaned seeds/weight
Range
/lb
/kg
32,60040,10
30,90081,600
572,000618,000
29,45048,300
30,00080,700
35,30066,150
14,80018,200
14,00037,000
259,460280,325
13,35921,910
13,60036,600
16,00030,000
Average
/kg
/lb
36,400
44,100
600,000
46,300
16,500
20,000
272,160
21,000
References
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
Bonner FT, Graney DL. 1974. Catalpa, catalpa. In: Schopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 281283.
Brown CL, Kirkman LK. 1990. Trees of Georgia and adjacent states.
Portland, OR:Timber Press. 292 p.
Fosket EB, Briggs WR. 1970. Photosensitive seed germination in Catalpa
speciosa. Botanical Gazette 131(2): 167172.
Francis JK. 1990. Catalpa longissima (Jacq.) Dum. Cours., yokewood. Silvics
Manual SO-ITF-SM-37. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 4 p.
Francis JK. 1993. Seeds of Puerto Rican trees and shrubs: second installment. Res. Note SO-374. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 5 p.
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Little EL, Delisle AL. 1962. Flowering, size, growth rate and life span: forest
trees, North American. In: Altman PL, Dittmer DS, eds. Biological handbook on growth. Washington, DC: Federation of American Societies for
Experimental Biology: table 103.
Rehder A.1940. Manual of cultivated trees and shrubs. 2nd ed. New York:
Macmillan. 996 p.
Sargent CS. 1965. Manual of the trees and of North America (exclusive of
Mexico). 2nd ed., corrected and reprinted. New York: Dover. 934 p.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
Williams RD, Hanks SH. 1976. Hardwood nurserymans guide. Agric.
Handbk. 473. Washington, DC: USDA Forest Service. 78 p.
Catalpa
347
RhamnaceaeBuckthorn family
Ceanothus L.
ceanothus
Susan G. Conard and Merton J. Reed
Dr. Conard is a program leader at the USDA Forest Services Research and Development National Office,
Arlington,Virginia; Dr. Reed retired from the USDA Forest Services Southwestern Forest and Range
Experiment Station
New-Jersey-tea, Jersey-tea,
C. arboreus Greene
C. arboreus var. glabra Jepson
C. cordulatus Kellogg
feltleaf ceanothus,
island myrtle, Catalina ceanothus
mountain whitethorn,
snowbush, whitethorn ceanothus
C. crassifolius Torr.
hoaryleaf ceanothus
buckbrush ceanothus,
wedgeleaf ceanothus,
hornbrush, buckbrush
Monterey ceanothus
snowbrush ceanothus,
mountain balm, sticky-laurel,
tobacco brush
varnish-leaf ceanothus,
Hooker ceanothus, snowbrush
C
Occurrence
Dry woods, Ontario to Manitoba, Maine to
North Dakota, S to Florida & Texas
Larger islands of Santa Barbara Channel, California (up
to 300 m on dry slopes & chaparral)
Baja California & mtns of S California, N to SW Oregon,
E to Nevada (9002,900 m on rocky ridges & ponderosa
pine to red fir forests)
Cis-montane southern California & Baja California (to
1,100 m on dry slopes & ridges, chaparral)
Inner Coast Range & Sierra Nevada foothills, California
into Oregon, S to Baja California (1001,800 m in
chaparral & ponderosa pine forests)
San Luis Obispo Co., N through Mendocino Co.,
California (up to 200 m on coastal bluffs, in closed-cone
pine forests)
Westside Sierra Nevada, spotty in northern Coast Range,
California (9001,800 m, under oaks & pines)
South Dakota to New Mexico, Arizona, & Mexico (1,500
to 3,000 m, in ponderosa pine to dry Douglas-fir forests)
W Texas to S California, Utah, & N Mexico (3002,300 m,
chaparral & desert chaparral)
Coastal areas in Santa Barbara & San Luis Obispo Cos.,
California (to 200 m in dry, sandy flats & slopes)
N California, Oregon,Washington to S California,
Arizona, & New Mexico (3002,100 m, in ponderosa pine
to western hemlock, white fir forests; chaparral in SW)
S California to N Baja California (to 1,800 m in chaparral,
dry slopes)
California
Coast Ranges, San Luis Obispo & Santa Barbara Cos.
& San Gabriel Mtns to Humboldt Co., California
(to 1,300 m in chaparral)
Sierra Nevada & N Coast Range S to Calaveras Co.,
California; higher mtns of Oregon & Washington,W Nevada
(9002,200 m, common in ponderosa & Jeffrey pine forests)
N California, Oregon, Idaho,Washington, & W Montana
to S British Columbia (around 1,200 m in ponderosa pine
Douglas-fir/mixed conifer, western hemlock zones)
Coast Range in Los Angeles & Riverside Cos., Parry to
Humboldt Co., California (1501,000 m, in chaparral)
Coastal mountains Santa Barbara Co., California, to
Douglas Co., Oregon (from sea level to 600 m in coast
redwood, mixed-evergreen, Douglas-fir forest, & chaparral
Coast Ranges, British Columbia to Marin Co.,
California, Siskiyou Mtns, Sierra Nevada/Cascade axis
E to SW Alberta, Montana, South Dakota, & Colorado
(to 3,000 m, in many forest types, ponderosa pine to
subalpine)
Same as above, but more common near coast
Sources: Franklin and others (1985), Hitchcock and others (1961), Lenz and Dourley (1981), McMinn (1964), Munz and Keck (1968), Reed (1974), Sampson and
Jesperson (1963), Schmidt (1993).
Ceanothus
349
Figure 1Ceanothus, ceanothus: capsules of C. americanus, New-Jersey-tea (top) and C. velutinus, snowbrush
(bottom).
hoaryleaf ceanothus at 5 years (Everett 1957), desert ceanothus at 6 to 8 years (Zammit and Zedler 1992), and snowbrush ceanothus at 6 to 10 years (McDonald and others
1998). Thus it appears that most species can be expected to
begin producing seeds by about 5 to 10 years of age.
Fendler ceanothus has been reported to bear good seedcrops
annually (Reed 1974). However, in hoaryleaf ceanothus,
desert ceanothus, chaparral whitethorn, and other species,
350
Table 2Ceanothus, ceanothus: phenology of flowering and fruiting, height, and year of first cultivation
Species
Flowering
Fruit-ripening
C. americanus
C. arboreus
C. cordulatus
California
Oregon
C. crassifolius
C. cuneatus
C. cuneatus var. rigidus
C. diversifolius
C. fendleri (Arizona)
C. greggii (Arizona*)
C. impressus
C. integerrimus
C. leucodermis
C. oliganthus
C. prostratus
C. sanguineus
C. sorediatus
C. thyrsiflorus
C. velutinus
California
N Idaho
W Montana
SW Oregon
Utah
MayJuly
FebAug
Augearly Oct
Mayearly Oct
MayJune
JuneJuly
JanJune
MarJune
DecApr
Spring
AprOct
MarApr
FebApr
AprAug
FebApr
AprJune
AprJune
MarApr
JanJune
JulySept
AugSept
MayJune
AprJuly
MayJune
JuneJuly
AugDec
July
June
JuneAug
JulyAug
MayJune
July
JuneJuly
MayJuly
AprJuly
JuneAug
May 20July 25
June 25July 15
MayJuly
JulyAug
July 15Aug 1
Aug 10Sept 10
JulySept
Aug 1Aug 30
Height at
maturity (m)
0.51
39
0.62.5
1.23
14.5
12.1
0.3 or less
0.21
0.61.8
15.5
1.27.5
.05.15
1.53
15.5
1.28
0.62.4
Year first
cultivated
1713
1911
1927
1848
1847
1941
1893
1850
1812
1837
1853
Sources: Evans and others (1987), Furbush (1962), Hitchcock (1961), Hubbard (1958), Kearney (1951), McMinn (1964), Mirov and Kraebel (1939), Plummer and others
(1968), Reed (1974), Rowntree (1948), Sampson and Jesperson (1963), Swingle (1939),Van Dersal (1938),Van Rensslaer (1942).
Elevations: * 9001,500 m. 700 m. 1,650 m.
Ceanothus
351
/kg
C. americanus
C. arboreus
C. cordulatus
C. crassifolius
C. cuneatus
C. cuneatus var. rigidus
C. diversifolius
C. greggii
C. impressus
C. integerrimus
C. oliganthus
C. prostratus
C. sanguineus
C. sorediatus
C. thyrsiflorus
C. velutinus
212291
106110
311396
73143
80123
128179
137161
8298
282291
267269
106400
135335
Average
/lb
96132
4850
141179
3365
3656
5881
6273
3745
128132
121122
48181
61152
/kg
247
108
366
117
108
159
185
51
245
154
148
90
287
207
/lb
Samples
112
49
166
53
49
72
84
23
111
70
67
41
130
94
5
2
4
3
3
1
1
1
2
2
3
2
2
Sources: Emery (1964), Hubbard (1958), Mirov and Kraebel (1939), Plummer and others (1968), Quick (1935), Quick and Quick (1961), Reed (1974), Swingle (1939).
general, longer periods of cold stratification are more effective than short ones. For example, Radwan and Crouch
(1977) observed increasing germination of redstem ceanothus as cold stratification was increased from 1 to 3 or 4
months; no germination occurred without stratification.
Similar patterns were observed by Quick and Quick (1961)
for deerbrush ceanothus (increased germination up to 2
months of stratification) and Bullock (1982) for mountain
whitethorn (increased germination up to 3 months). In lieu
of cold stratification, a chemical treatment with gibberellin
and thiourea was used to induce germination of buckbrush
ceanothus (Adams and others 1961). Treatment with potassium salts of gibberellin also sucessfully replaced cold stratification in germination tests on redstem ceanothus seeds
(Radwan and Crouch 1977). Following chemical treatments,
seeds may then be germinated or dried again and stored
(Adams and others 1961). Although emphasis has been on
more natural methods of stimulating germination, seeds of
snowbrush ceanothus and other species can be germinated
quite successfully with acid scarification followed by a gibberellin treatment (Conard 1996).
Specific germination test conditions have not been well
defined for most species of ceanothus. Sand or a mixture of
sand and soil has been used as the moisture-supplying medium in most of the reported germination tests (Emery 1964;
Quick 1935; Reed 1974), but filter paper has also been used
successfully (Keeley 1987a). Diurnally alternating tempera-
Species
C. americanus
C. arboreus
C. cordulatus
C. crassifolius
C. cuneatus
C. megacarpus
C. oliganthus
C. prostratus
C. sanguineus
C. sorediatus
C. thyrsiflorus
C. velutinus
Pregermination treatments
Hot water soak
Cold
Temp
Time*
stratification
(C)
(min)
(days)
77100
7191
90
85
80
71
71
71
120
100
70
71
77100
100
77100
85
71
120
100
70
120
100
70
71
100
77100
100
100
100
100
100
100
71
71
90
71
0
ttc
ttc
ttc
ttc
ttc
ttc
ttc
ttc
5
5
60
0
ttc
ttc
0
1
ttc
ttc
ttc
5
5
60
0
5
5
60
0
ttc
0.5
ttc
1
5
15
15
5
5
ttc
ttc
ttc
ttc
90
60
0
94
94
94
90
0
90
30
30
30
30
0
60
0
3060
60
56
90
30
30
30
30
30
30
30
30
0
115
90
120
120
120
0
90
0
90
0
6384
90
Germination
test days
50
30
40112
35
35
35
2190
90
2190
21
21
21
21
60112
60
17
30
20
21
21
21
21
21
21
21
21
70
30
32
32
32
32
30
30
60
60
30
C
Germination rate
Avg
(%)
Samples
65
32
90
74
90
74
76
48
92
28
38
3
10
85
61
16
51
73
100
85
68
50
47
7
88
53
54
6
62
92
71
97
92
41
0
100
38
83
73
82
70
4
1
3+
4
4
4
1+
1+
1+
3
3
3
3
2+
1+
1+
1
1+
3
3
3
3
3
3
3
3
1+
1+
3
3
3
3
1+
1
1+
1
1
2+
Sources: Emery (1964), Keeley (1987a), Mirov and Kraebel (1939), Quick (1935), Quick and Quick (1961), Radwan and Crouch (1977), Reed (1974),Van Dersal (1938).
* ttc = time to cool (to room temperature) varied from several hours to overnight.
Results reported here are for dry heat treatments, with germination in the dark; see Keeley (1987) for data on light germination.
Ceanothus
353
354
References
Adams L.1962. Planting depths for seeds of three species of ceanothus.
Res. Note PSW-194. Berkeley, CA: USDA Forest Service, Pacific
Southwest Forest and Range Experiment Station. 3 p.
Adams L, Stefanescu E, Dunaway DJ. 1961. Gibberellin and thiourea break
seed dormancy in California ceanothus. Res. Note PSW-178. Berkeley,
CA: USDA Forest Service, Pacific Southewest Forest and Range
Experiment Station. 4 p.
Barro SC, Poth M. 1987. A possible advantage for the obligate seeding
strategy in Ceanothus [unpublished manuscript on file]. Riverside, CA:
USDA Forest Service, Pacific Southwest Research Station, Forest Fire
Laboratory.
Binkley D, Cromack K Jr, Fredriksen RL. 1982. Nitrogen accretion and availability in some snowbrush ecosystems. Forest Science 28(4): 720724.
Binkley D, Husted L. 1983. Nitrogen accretion, soil fertility, and Douglas-fir
nutrition in association with redstem ceanothus. Canadian Journal of
Forest Research 13: 122125.
Brickell C, Zuk JD. 1997. American Horticultural Society AZ encyclopedia
of garden plants. New York: DK Publishing: 239241.
Bullock S. 1982. Reproductive ecology of Ceanothus cordulatus [MA thesis].
Fresno: California State University.
Conard SG. 1996. Unpublished data. Washington, DC: USDA Forest
Service, Research and Development.
Conard SG, Jaramillo AE, Cromack K Jr, Rose S, comps. 1985. The role of
the genus Ceanothus in western forest ecosystems. Gen.Tech. Rep.
PNW-182. Portland, OR: USDA Forest Service, Pacific Northwest Forest
and Range Experiment Station. 72 p.
Delwiche CC, Zinke PJ, Johnson CM. 1965. Nitrogen fixation by Ceanothus.
Plant Physiology 40(6): 10451047.
DFFW [Department of Forester and Fire Warden]. 1985. Emergency
revegetation in Los Angeles County 19191984. Los Angeles: County
Fire Department. 93 p.
Emery DE. 1964. Seed propagation of native California plants. Leaflets of
the Santa Barbara Botanical Garden 1(10): 9091.
Emery DE. 1988. Seed propagation of native California plants. Santa
Barbara, CA: Santa Barbara Botanical Garden. 115 p.
Evans RA, Biswell HH, Palmquist DE. 1987. Seed dispersal in Ceanothus
cuneatus and C. leucodermis in a Sierran oakwoodland savanna.
Madroo 34(4): 283293.
Everett PC. 1957. A summary of the culture of California plants at the
Rancho Santa Ana Botanic Garden 19271950. Claremont, CA: Rancho
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Hellmers H, Kelleher JM. 1959. Ceanothus leucodermis and soil nitrogen in
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Hubbard RL. 1958. Hot water and thiourea break dormancy of wedgeleaf
ceanothus seed. Res. Note PSW-143. Berkeley, CA: USDA Forest
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Keeley JE. 1977. Seed production, seed populations in soil, and seedling
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Keeley JE. 1987a. Role of fire in seed germination of woody taxa in
California chaparral. Ecology 68(2): 434443.
Keeley JE. 1987b. Fruit production patterns in the chaparral shrub
Ceanothus crassifolius. Madrono 34(4): 273282.
Keeley JE. 1991. Seed germination and life history syndromes in the
California chaparral. Botanical Review 57(2): 81116.
Kruckeberg AR. 1982. Gardening with native plants of the Pacific
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Lanini WT, Radosevich SR. 1986. Response of three conifer species to site
preparation and shrub control. Forest Science 32: 6177.
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and environmental use in southern California and adjacent areas.
Claremont, CA: Rancho Santa Ana Botanic Garden. 232 p.
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planted deerbrush and greenleaf manzanita seedlings. Res. Note PSW422. Albany, CA: USDA Forest Service, Pacific Southwest Research
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University of California Press: 278320.
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shrubs and herbs in southern California chaparral. Ecology 72(6):
19932004.
Munz PA, Keck DD. 1968. A California flora with supplement. Berkeley:
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Perry B. 1992. Landscape plants for western regions, an illustrated guide to
plants for water conservation. Claremont, CA: Land Design Publishing.
318 p.
Ceanothus
355
356
Swingle CF, comp. 1939. Seed propagation of trees, shrubs and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA Soil
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erosion-control and wildlife values. Misc. Pub. 303. Washington, DC:
USDA. 362 p.
Van Rensslaer M, McMinn HE. 1942. Ceanothus. Berkeley, CA: Gillick Press.
308 p.
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109112.
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175187.
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Ecology 81: 499511.
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(Ceanothus velutinus) in the Oregon Cascades. Ecology 49(6): 1134-1145.
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1967 November 28December 1. Washington, DC: National Academy
of Sciences: 4053.
PinaceaePine family
Cedrus Trew
cedar
Paula M. Pijut
Dr. Pijut is a plant physiologist at the USDA Forest Services North Central Research Station,
Hardwood Tree Improvement and Regeneration Center, West Lafayette, Indiana
Table 1Cedrus, cedar: nomenclature, occurrence, height at maturity, and date first cultivated
Height at
maturity (m)
Scientific name
Common name
Occurrence
C. atlantica (Endl.) G.
Manetti ex Carriere
Atlas cedar
Cyprian cedar
C. libani A. Rich.
deodar cedar
cedar of Lebanon
Year first
cultivated
940
Before 1840
824
1879
1550
1831
1540
Pre-1650
Cedrus
357
Species
C. atlantica
C. brevifolia
C. deodara
C. libani
Cone characteristics
Length
Width
Ripe color
(cm)
(cm)
Light brown
Light brown
Reddish brown
Grayish brown
358
58
510
713
812
35
36
59
36
Seed size
Length
Width
(mm)
(mm)
813
814
1015
1014
46
56
57
46
Leaf characteristics
Length
No. in
(cm)
whorls
12.5
0.51.6
26.0
13.5
2045
1520
2030
2040
mature cone.
seeds with
Cedrus
359
Flowering
Cone
ripening
Seed
dispersal
C. atlantica
C. deodara
C. libani
JuneSept
SeptOct
JuneSept
SeptOct
SeptNov
AugOct
Fallspring
SeptDec
Fallspring
Species
C. atlantica
C. brevifolia
C. deodara
C. libani
C. libani ssp.
stenocoma
Avg no.
cleaned seeds
/kg
/lb
13,900
13,000
8,150
11,700
17,600
6,300
5,890
3,700
5,300
8,000
360
Commercial
seed purity (%)
89
85
87
References
Adams RP. 1991.
Chandra JP, Ram A. 1980. Studies on depth of sowing deodar (Cedrus deodara) seed. Indian Forester 106(12): 852855.
Chaney WR. 1993. Cedrus libani, cedar of Lebanon. Arbor Age 13 (1):
2627.
Demetci EY. 1986. Toros sediri (Cedrus libani A. Richard) odununun bazi
fiziksel ve mekanik zellikleri zerine arastirmalar [in Turkish; summary in
English: Studies on the some physical and mechanical properties of
cedar (Cedrus libani A. Richard) wood].Teknik Blten Serisi 180.
Ormancilik Arastirma Enstits Yayinlari. 60 p.
Dirr MA. 1990. Manual of woody landscape plants: their identification,
ornamental characteristics, culture, propagation, and uses. Champaign, IL:
Stipes Publishing Company. 1007 p.
Dirr MA, Heuser Jr CW. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Dirr MA, Lindstrom OM, Lewandowski R,Vehr MJ. 1993. Cold hardiness
estimates of woody taxa from cultivated and wild collections. Journal of
Environmental Horticulture 11(4): 200203.
Doty JC. 1982. Seedling production: Cedrus deodara. Combined
Proceedings of the International Plant Propagators Society 31: 6163.
Erkuloglu OS. 1995. Kayin, goknac ve sedir tohumlarini: uzun sure saklama
olanaklari uzerine arastirmalar [in Turkish, summary in English: Possiblities
for long-term storage of beech, fire and cedar seeds]. Ic Anadolu
Ormancilik Arastirma Enstitusu Dergisis 77: 4587.
Farjon A. 1990. Pinaceae: drawings and descriptions of the genera Abies,
Cedrus, Pseudolarix, Keteleeria, Nothotsuga, Tsuga, Cathaya, Pseudotsuga,
Larix, and Picea. Knigstein, Germany: Koeltz Scientific Books. 330 p.
Guehl JM, Falconnet G, Gruez J. 1989. Caractristiques physiologiques et
survie aprs plantation de plants de Cedrus atlantica levs en conteneurs sur diffrents types de substrats de culture [in French, summary
in English: Physiological characteristics and field survival of Cedrus
atlantica seedlings grown on different container growth media]. Annales
des Sciences Forestieres 46: 114.
Cedrus
361
Hartmann HT, Kester DE, Davies FT. 1990. Plant propagation: principles and
practices. Englewood Cliffs, NJ: Prentice Hall. 647 p.
Heit CE. 1968. Propagation from seed: 16.Testing and growing Cedrus
species. American Nurseryman 128(6): 1213, 8794.
Hillier Nurseries (Winchester) Ltd. 1991. The Hillier manual of trees and
shrubs. Melksham, Wiltshire: Redwood Press Ltd. 704 p.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing. Rules 1993. Seed Science & Technology 21 (Suppl.): 1259.
Koller GL. 1982. Introducing Cedrus deodara Shalimar. Arnoldia 42(4):
153157.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan. 1290 p.
Liu M. 1990. Deodara cedar. In: Chen Z, Evans DA, Sharp WR, Ammirato
PV, Sndahl MR, eds. Handbook of plant cell culture,Volume 6, Perennial
crops. New York: McGraw-Hill. 506 p.
Loureiro AM. 1990. Cultura da Cedrus atlantica (Endl.) Carr. [in Portuguese,
English summary: Silviculture of Cedrus atlantica]. Srie Didctica 5.
Universidade de Trs-os-Montes e Alto Douro, Cincias Aplicadas. 61 p.
Loureiro A[M]. 1994. Algumas notas sobre a cultura de Cedrus atlantica
(Endl.) Carr. [in Portuguese, summary in English: Some notes on the culture of Cedrus atlantica]. Informacao Florestal 5: 811.
Lyon L. 1984. Winter grafting of cedar, spruce, and ornamental cherry.
Combined Proceedings of the International Plant Propagators Society
33: 5455.
Macdonald B. 1986. Practical woody plant propagation for nursery growers. Portland, OR:Timber Press. 669 p.
Maheshwari P, Biswas C. 1970. Cedrus. Bot. Monogr. 5. New Delhi: Council
of Scientific and Industrial Research. 112 p.
Mittal RK. 1983. Studies on the mycoflora and its control on the seeds of
some forest trees: 1. Cedrus deodara. Canadian Journal of Botany 61:
197201.
Nicholson R. 1984. Propagation notes Cedrus deodara Shalimar and
Calocedrus decurrens. Plant Propagator 30: 56.
Panetsos KP, Christou A, Scaltsoyiannes A. 1992. First analysis on allozyme
variation in cedar species (Cedrus sp.). Silvae Genetica 41(6): 339342.
Piola F, Rohr R. 1996. A method to overcome seed and axillary bud dormancy to improve Cedrus libani micropropagation. Plant Tissue Culture
and Biotechnology 2(4): 199201.
Piola F, Label P,Vergne P, von Aderkas P, Rohr R. 1998. Effects of endogenous ABA levels and temperature on cedar (Cedrus libani Loudon) bud
dormancy in vitro. Plant Cell Reports 18: 279283.
362
CelastraceaeBittersweet family
Celastrus scandens L.
American bittersweet
G.W. Wendel, Jill R. Barbour, and Robert P. Karrfalt
Dr.Wendel retired from the USDA Forest Services Northeastern Forest Experiment Station;
Ms. Barbour is a germination specialist at and Mr. Karrfalt is director of the USDA Forest Services
National Seed Laboratory, Dry Branch, Georgia
Other common names. climbing bittersweet, shrubby
bittersweet.
Growth habit, occurrence, and use. American bittersweet is a deciduous climbing or twining shrub of eastern
North America (Brizicky 1964; Fernald 1950) that occurs in
thickets, in stands of young trees, along fence rows, and
along streams, usually in rich soil. It occurs naturally from
southern Quebec; west to southern Manitoba; and south to
Oklahoma and central Texas, Arkansas, Tennessee, northern
Alabama, and western North Carolina (Brizicky 1964).
Some authors (Fernald 1950; USDA FS 1948) reported it in
Louisiana, New Mexico, Georgia, and Mississippi, but its
occurrence has not been verified in Georgia, Louisiana, or
Mississippi (Brizicky 1964).
The plant is valuable for ornamental purposes and game
food and cover; the bark has been used for medicinal purposes (USDA FS 1948). Among the animals and birds feeding on American bittersweet are the bobwhite quail (Colinus
virginianus), ruffed grouse (Bonasa umbellus), ringneck
pheasant (Phasianus colchicus), eastern cottontail
(Silvilagus floridanus), fox squirrel (Sciurus niger), and various songbirds (Van Dersal 1938). It was introduced into
cultivation in 1736 (USDA FS 1948).
By 1970, oriental or Asiatic bittersweetC. orbiculatus
Thunb.had become naturalized on at least 84 sites from
Georgia to Maine and west to Iowa (McNab and Meeker
1987), occupying many of the same sites as American bittersweet. It is listed as an invasive plant by the United States
Government (USDA NRCS 1999). In some locales, the
species is found mainly along fence lines, resulting from the
germination of seeds contained in the droppings from frugivorous birds (McNab and Meeker 1987). The stem, leaves,
and berries of oriental bittersweet are reported to be toxic
for human consumption (McNab and Meeker 1987).
Flowering and fruiting. The small greenish, polygamodioecious or dioecious flowers open from May to June and
are borne in raceme-like clusters at the end of branches
Celastrus
363
364
References
Brizicky GK. 1964. The genera of Celastrales in the southeastern United
States. Journal of the Arnold Arboretum 45: 206234.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Fernald ML.1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1632 p.
Hart HT. 1928. Delayed germination in seed of Peltandra virginica and
Celastrus scandens. Puget Sound Biology Sation Publication 6: 255261.
Heit CE.1967. Propagation from seed: 11. Storage of deciduous tree and
shrub seeds. American Nurseryman 126(10): 1213, 8694.
Heit CE.1968. Thirty-five years testing of tree and shrub seed. Journal of
Forestry 66: 632634.
Heit CE, Nelson C. 1941. Approximate germination tests of dormant seeds
by excising embryos. Proceedings of the Association of Official Seed
Analysts 194: 8789.
Celastrus
365
UlmaceaeElm family
Celtis L.
hackberry
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Common name(s)
Occurrence
C. laevigata Willd.
C. mississippiensis Spach
C. laevigata var. reticulata
(Torr.) L. Benson
C. reticulata Torr.
C. occidentalis L.
C. crassifolia Lam.
Flowering
Fruit ripening
Seed dispersal
C. laevigata
C. laevigata var. reticulata
C. occidentalis
AprMay
MarApr
AprMay
SeptOct
Late fall
SeptOct
OctDec
Fallwinter
Octwinter
366
Figure 3Celtis laevigata, sugarberry: seedling development at 1, 2, and 5 days after germination.
endocarp
seedcoat
cotyledons
hypocotyl
radicle
367
Species
Height at
maturity (m)
Year first
cultivated
1824
914
940
1811
1890
1656
C. laevigata
C. laevigata var. reticulata
C. occidentalis
Minimum
seed-bearing
age (yrs)
Preripe
15
Fruit color
Ripe
Green
Orange-red
Species
C. laevigata
C. laevigata var. reticulata
C. occidentalis
4,850
4,520
Range
/kg
2,200
2,050
8,15015,600
5,15014,500
7,70011,900
Cleaned seeds/weight
Average
/lb
/kg
/lb
3,7007,080
2,3406,600
3,5005,400
13,200
10,500
9,500
6,000
4,870
4,300
Species
C. laevigata
C. laevigata var. reticulata
C. occidentalis
30
30
30
20
20
20
60
60
60
Germination rate
Amount
(%)
Days
3050
39
2530
37
Germination %
Avg
Samples
55
37
47
6+
7
7
References
Bonner FT. 1974. Celtis, hackberry. In: Schopmeyer CS, tech. coord. Seeds of
woody plants in the United States. Agric. Handbk. 450. Washington, DC:
USDA Forest Service: 298300.
Bonner FT. 1984. Germination tests for sugarberry (Celtis laevigata Willd.).
AOSA Newsletter 58(2): 2426.
Kennedy HE Jr. 1990. Celtis laevigata Willd., sugarberry. In: Burns RM,
Honkala BH, tech. coord. Silvics of North America.Volume 2.
Hardwoods. Agric. Handbk. 654. Washington, DC: USDA Forest Service:
258261.
Krajicek JE, Williams RD. 1990. Celtis occidentalis L., hackberry. In: Burns RM,
Honkala BH, tech. coord. Silvics of North America.Volume 2.
Hardwoods. Agric. Handbk. 654. Washington, DC: USDA Forest Service:
262265.
Little EL Jr. 1950. Southwestern trees, a guide to the native species of new
Mexico and Arizona. Agric. Handbk. 9. Washington, DC: USDA Forest
Service. 109 p.
368
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Preston RJ Jr. 1947. Rocky Mountain trees. 2nd ed. Ames: Iowa State
College Press. 285 p.
Rehder A. 1940. Manual of cultivated trees and shrubs. 2nd ed. New York:
Macmillan. 996 p.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA Soil
Conservation Service. 198 p.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
Vora RS. 1989. Seed germination characteristics of selected native plants of
the lower Rio Grande Valley,Texas. Journal of Range Management 42:
3640.
Williams RD, Hanks SH. 1976. Hardwood nurserymans guide. Agric.
Handbk. 473. Washington, DC: USDA Forest Service. 78 p.
RubiaceaeMadder family
Cephalanthus occidentalis L.
buttonbush
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station
Mississippi State, Mississippi
Cephalanthus
369
Mississippi
Medium
Water
Blotter paper
Temperature (C)
2434
30
Yes
No
30
10
Germination (%)
86
78
Light
No. of samples
References
Bonner FT. 1974a. Chemical components of some southern fruits and
seeds. Res. Note SO-183. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 3 p.
Bonner FT. 1974b. Cephalanthus occidentalis L., common buttonbush. In:
Schopmeyer CS, tech. coord. Seeds of woody plants in the United States.
Agric. Handbk. 450. Washington, DC: USDA Forest Service: 301302.
DuBarry AP, Jr. 1963. Germination of bottomland tree seed while
immersed in water. Journal of Forestry 61: 225226.
Imbert FM, Richards JH. 1993. Protandry, incompatibility, and secondary
pollen presentation in Cephalanthus occidentalis (Rubiaceae). American
Journal of Botany 80: 395404.
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Maisenhelder LC. 1958. Understory plants of bottomland forests. Occ. Pap.
165. New Orleans: USDA Forest Service, Southern Forest Experiment
Station. 26 p.
Van Dersal WR. 1938. Native woody plants of the United States: their erosion-control and wildlife values. Misc. Pub. 393. Washington, DC: USDA.
362 p.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
370
FabaceaePea family
Ceratonia siliqua L.
carob
Wayne D. Shepperd
Dr. Shepperd is a principal silviculturist at the USDA Forest Services Rocky Mountain Research Station,
Fort Collins, Colorado
371
seed.
longitudinal sections
372
References
Alexander RR, Shepperd WD. 1974. Ceratonia siliqua L., carob. In:
Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service:
303304.
Bailey LH. 1947. Standard cyclopedia of horticulture. 2nd ed. New York:
Macmillan. 717718.
Binder RJ, Coit JE, Williams KT, Brekke JE. 1959. Carob varieties and composition. Food Technology 13: 213216.
Catarino F. 1993. The carob tree: an exemplary plant. Naturopa 1993(73):
1415.
Coit JE. 1951. Carob or St. Johnsbread. Journal of Economic Botany 5(1):
8296.
Coit JE. 1961. Carob varieties. Fruit Varieties and Horticultural Digest 15(4):
7577.
Coit JE. 1962. Carob culture in the semiarid southwest.Vista, CA: J. Eliot
Coit. 6 p.
Goor AY, Barney CW. 1968. Forest tree planting in arid zones. New York:
Ronald Press. 409 p.
Griffiths C. 1952. Locust kernel gum. Food 21: 5859.
Karschon R. 1960. Studies in nursery practice for carob (Ceratonia siliqua
L.). Leafl. 14. Ilanot: Israel Department of Forestry. 8 p. [Forestry
Abstracts 22: 3017, 1961].
Loock EEM. 1940. The carob or locust tree (Ceratonia siliqua L.). Journal
South African Forestry Association 4: 7880 [Forestry Abstracts 34.27:
12.2, 1941].
Rhizopoulou S, Davies WJ. 1991. Influence of soil drying on root development, water relations and leaf growth of Ceratonia siliqua L. Oecologia
88(1): 4147.
Roukas T. 1994. Solid-state fermentation of carob pods for ethanol production. Applied Microbiology and Biotechnology 41(3): 296301.
Toth J. 1965. The forest aspect of a plantation in the Sahara. Revue
Forestiere Francaise 17: 674695 [in French; Forestry Abstracts 27: 3883,
1966].
Ceratonia
373
FabaceaePea family
Cercis L.
redbud
Valerie A. Banner and William I. Stein
Ms. Banner is a general biologist and Dr. Stein is a forest ecologist emeritus at the
USDA Forest Services Pacific Northwest Research Station, Corvallis, Oregon
tall; the tallest one on record is 8.8 m and the tallest Texas
redbud is about the same (AFA 1996). Eastern redbud
occurs on many soils in moist open woodlands, flood plains,
river thickets, and borders of small streams, whereas the
variety, Texas redbud, often inhabits drier locations, primarily paleozoic limestone formations such as xeric pastures,
hills, outcrops, and bluffs (Hopkins 1942). California redbud
is unevenly distributed at elevations of 70 to 1,524 m along
foothill streams, flats, draws, low slopes and canyons and on
dry gravelly and rocky soils (Chamlee 1983; Jepson 1936;
Sudworth 1908).
Redbuds are valued particularly for their showy buds
and flowers that appear before the leaves (Clark and
Bachtell 1992; McMinn and Maino 1937). They exhibit
caulifloryflowering directly along older branches and
trunkswhich is rare among temperate species (Owens and
Ewers 1991) and contributes greatly to flowering showiness.
Flowers typically are a deep reddish purple (magenta) but
vary among localities (Coe 1993; Smith 1986) and species
(table 3). Some white ones occur naturally, and cultivars
have been developed for particular flower and leaf colors
(Raulston 1990). Ornamental uses are extensive within each
species indigenous range and several species have proven
hardy more extensively (McMinn and Maino 1937;
Common name(s)
Occurrence
C. canadensis L.
eastern redbud,
redbud, Judas-tree
Texas redbud
Mexican redbud
E-central Mexico
California redbud,
Arizona redbud,
western redbud
Sources: Clark and Bachtell (1992), Hopkins (1942), Hosie (1969), Little (1979), Robertson and Lee (1976), Sargent (1933).
374
Table 2Cercis, redbud: growth habit, height, legume color, and size
Height at
maturity (m)
Legume color
C
Legume size
Length
Width
(cm)
(mm)
Seed
diameter
(mm)
Species
Growth habit
C. canadensis
C. canadensis
var. texensis
C. orbiculata
Tree or shrub
712
Reddish brown
510
818
45
Shrub or tree
Shrub or tree
410
26
Reddish brown
Reddish purple, dull
red, to reddish brown
610
49
825
1325
45
34
Sources: Fernald (1950), Hopkins (1942), Hosie (1969), Jepson (1936), McMinn (1939), Munz and Keck (1959), Sargent (1933).
Flowering
Flower color
Fruit ripening
C. canadensis
C. canadensis
var. texensis
C. orbiculata
MarMay
Magentapurplish pink
Julyearly autumn
MarApr
FebMay
Magenta pink
Magenta pinkreddish purple
AugSept
JulySept
Sources: Abrahms (1944), Clark and Bachtell (1992), Fernald (1950), Hopkins (1942), Jepson (1936), Mirov and Kraebel (1939),Van Dersal (1938).
Robertson 1976). Where redbuds are numerous, they provide valued bee pasture in early spring (Magers 1970). The
buds, flowers, and legumes (pods) of redbuds are edible and
have been used in salads or batter (Coe 1993). Native
Californians used the roots and bark of California redbud in
basketry (Coe 1993; Jepson 1936); remedies for diarrhea
and dysentery were also made from the astringent bark
(Balls 1962).
Redbuds also are used for borders, erosion control,
windbreaks, and wildlife plantings. Eastern redbud is
browsed by white-tail deer (Odocoileus virginiana) and the
seeds are eaten by birds, including bobwhite (Colinus
virginianus) (Van Dersal 1938). California redbud is moderately important as fall and spring browse for deer but has
been rated only fair to poor for goats and poor or useless for
other livestock (Sampson and Jespersen 1963).
Two fungal diseases affect the flowering and attractiveness of eastern redbudverticillium wilt (Verticillium sp.)
and botryosphaeria canker (Botryosphaeria dothidea
(Moug.:Fr.) Ces. & De Not.)by causing die-back of
branches. The canker has become more common and
destructive in the eastern United States, appearing to attack
trees that are under stress (Geneve 1991a; Raulston 1990;
Vining 1986).
Flowering and fruiting. Flowering occurs from
February to May, varying somewhat by species and location
(table 3). The bisexual redbud flowers are brilliant pink to
Cercis
375
4 to 10
376
Figure 2Cercis canadensis, eastern redbud: the flattened seed in transverse section (above) and longitudinal
section (below).
25 C. Seeds of California redbud have been stored satisfactorily for 12 years or more under the same conditions as
many conifers at a moisture content of 5 to 9% and temperature of 18 C (Lippitt 1996).
Seeds of California redbud average about half again as
heavy as those of eastern redbud but seed weight varies
widely among lots for both species (table 4).
Pregermination treatments and germination tests.
Redbud seeds generally require pregermination treatment to
overcome dormancy attributable both to a hard, impermeable seedcoat and to some demonstrated, but not fully identified, embryo dormancy (Afanasiev 1944; Geneve 1991b;
Hamilton and Carpenter 1975; Heit 1967a7b; Jones and
Geneve 1995; Profumo and others 1979; Rascio and others
1998; Riggio-Bevilacqua and others 1985; Tipton 1992;
Zins 1978). Test results indicate that the level of dormancy
varies by species, seed source, seedlot, age of seeds, and
perhaps other factors. Given such variable dormancy, pretreatment might involve using the one demonstrated to be
most broadly applicable, or determining sufficiently the
nature of dormancy in local lots and applying a customized
pretreatment.
Three pretreatments have proven satisfactory for overcoming redbuds seedcoat impermeabilitymechanical
scarification, immersion in sulfuric acid, or in hot water
(table 5). In comparison tests, the acid treatment has generally produced more consistent or slightly better results
(Afanasiev 1944; Liu and others 1981), but good imbibition
of water has resulted after all 3 treatments. Acid treatment
involves immersing redbud seeds in concentrated sulfuric
acid for 15 to 90 minutes at room temperature followed by
thorough washing in water (Afanasiev 1944; Frett and Dirr
1979; Liu and others 1981). Length of treatment required
can be determined on a small sample; if immersion is too
short, seedcoats remain impermeable, if too long, the seeds
are damaged. Well-rinsed, acid-scarified seeds can be placed
immediately in stratification or surface-dried and stored several months until sown by hand or seeder (Heit 1967a).
Abrading, clipping, or piercing the seedcoat to expose
the endosperm and allow ready entry of water (Hamilton
and Carpenter 1975; Riggio-Bevilacqua and others 1985;
Zins 1978) can be done easily for small test lots but not as
readily in quantity. Immersing small or large quantities of
seeds in hot or boiling water can be done easily, but results
have been more variable than for acid treatmentsometimes
reasonably good (Fordham 1967; Mirov and Kraebel 1939),
other times poor to mediocre (Afanasiev 1944; Liu and others 1981). Hot water treatment clearly makes redbud seedcoats permeable but may cause internal damage. Better calibration of timetemperature effects appears needed
Species
C. canadensis
C. orbiculata
Seeds/100 wt
of legumes
2035
44
Seed wt/
legume vol
kg/hl
lb/bu
2.10
1.64
Average
/kg
/lb
39,570
27,460
17,950
12,455
Cleaned seeds/wt
Range
/kg
30,87055,100
20,95040,100
/lb
Samples
14,00025,000
9,50018,169
18
24
Table 5Cercis, redbud: scarification, stratification, germination test conditions, and test results*
Species
C. canadensis
C. canadensis
var. texensis
C. orbiculata
Scarification
TreatTime
ment
(min)
H2SO4
H2SO4
H2SO4
H2SO4
H2SO4
Mech.
H2SO4
Mech
H2SO4
H2SO4
H2SO4
H2SO4
H2SO4
Heat
Heat
H2SO4
45
45
30
2530
30
30
30
1560
3090
62
62
Overnight
9
60
Stratification
Temp
Days
(C)
Var.
Var.
42
3591
60
60
60
60
60
0
28
35
35
90
Peat
Peat
Cotton
Cotton
Sand
Peatperlite
Peatperlite
Peatperlite
Peatperlite
Vermiculite
Soil
Paper
Paper
Paper
Paper
Vermiculite
Vermiculite
Cotton
5
5
5
37
5
5
5
5
5
1
1
5
5
24
21
2030
25
25
25
25
1821
2027
2030
2030
21
28
48
107
8
30
24
24
24
24
42
14
28
28
14
14
118
118
10
Germination
(%)
77
78
97
88100
80
90
88
82
91
87
6772
43
83
95
100
38
52
84
Samples
2
2
2
7
2
5
5
5
5
3
12
1
1
25
81
1
1
1
Sources: Afanasiev (1944), Flemion (1941), Frett and Dirr (1979), Hamilton and Carpenter (1975), Heit (1967a), Liu and others (1981), Roy (1974),Tipton (1992), USDA
FS (1948, 1996),Williams (1949).
* Only the better results for each test series are listed. In several studies, only full seeds were tested.
Best results from a set of tests on each of 7 seedlots.
Test combinations used to develop a response surface.
Moist heat applied by immersing seeds in 82 C water that cooled gradually.
// Dry heat applied in oven at 121 C.
Cercis
377
378
excised embryo test reveals that the seeds maximum potential and generally is higher than indicated by a germination
test (Flemion 1941; Hamilton and Carpenter 1975; USDA
FS 1996).
Nursery practice. Redbuds are propagated most readily from seeds sown either in the fall or spring. Fall-sown
seeds may or may not be scarified, and stratification occurs
naturally in the seedbeds (Lippitt 1996; Raulston 1990). In
one reported instance, immature seeds of eastern redbud collected, extracted, and sown before the seedcoats hardened
yielded 90% germination the following spring (Titus 1940).
When seeds need to be scarified for either fall- or springsowingacid treatment for 15 to 60 minutes, rinsing, and a
24-hour soak in water; a boiling water dip for 15 seconds or
more followed by a 24-hour soak in cooler water; or immersion overnight in 88 C hot water that gradually coolscan
be generally used to overcome seedcoat dormancy (Frett and
Dirr 1979; Heit 1967b; Lippitt 1996; Raulston 1990;
Robertson 1976; Smith 1986). After scarified seeds have
imbibed water, they may need to be sorted to separate those
not swollen and still impermeable for further treatment.
When necessary, seeds are stratified at 1 to 5 C for 30 to 90
days. Stratification requirements are uncertain for 2 reasons:
variability among seedlots and the unknown stratification
effect produced by low temperature storage of the seeds.
Surface-dried seeds are drill- or broadcast-sown and
covered 0.6 to 2.5 cm (0.2 to 1 in) deep with soil, sand, sawdust, or bark. Some nurseries fumigate, then reinoculate
before sowing seedbeds. Presence of an endomycorrhizal
fungus is important; inoculation with Glomus fasciculatum
(Thaxter) Gerdemann and Trappe has increased first-season
growth of eastern redbud as much as 72% (Maronek and
Hendrix 1978). Mulching of fall-sown beds can be beneficial but the mulch must be removed when germination
starts. Germination is epigeal (figure 3).
Seedling return from nursery sowings is very variable.
For eastern redbud, an average of 2,425 usable seedlings
(range 617 to 7,055) were produced per kilogram of seeds
(1,100 usable seedlings/lb, range 280 to 3,200/lb) (Roy
1974). Germination is fairly consistent year to year for
California redbud, averaging 54 to 60% (Lippitt 1996).
Under favorable conditions, seedling height growth of eastern redbud can be rapid: about 0.5 m (20 in) in reinoculated
soil (Maronek and Hendrix 1978), 1 m (40 in) or more
under an intensive nitrogen fertilizer schedule, and about
2 m (80 in) if started in January in a greenhouse under longday conditions and transplanted outdoors after the danger
from frost is over (Raulston 1990).
References
Abrams L. 1944. Illustrated flora of the Pacific States.Volume 2,
Polygonaceae to Krameriaceae. Stanford, CA: Stanford University Press.
635 p.
Afanasiev M. 1944. A study of dormancy and germination of seeds of
Cercis canadensis. Journal of Agricultural Research 69(10): 405420.
AFA [American Forestry Association]. 1996. 199697 National register of
big trees. American Forests 102(1): 2051.
Balls EK. 1962. Early uses of California plants. Berkeley: University of
California Press. 103 p.
Bennett L. 1987. Tissue culturing redbud. American Nurseryman 166(7):
8591.
Chamlee H. 1983. The redbud in southern California. Fremontia 10(4): 27.
Clark R, Bachtell KR. 1992. Eastern redbud (Cercis canadensis L.). Morton
Arboretum Quarterly 28(1): 610.
Coe FW. 1993. Redbuds and Judas trees. Pacific Horticulture 54(2): 3741.
Donselman HM. 1976. Variation in native populations of eastern redbud
(Cercis canadensis L.) as influenced by geographic location. Proceedings
of the Florida State Horticultural Society 89: 370373.
Donselman HM, Flint HL. 1982. Genecology of eastern redbud (Cercis
canadensis). Ecology 63(4): 962971.
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Company. 1632 p.
Flemion F. 1941. Further studies on the rapid determination of the germinative capacity of seeds. Contributions from Boyce Thompson Institute
11: 455464.
Fordham AJ. 1967. Hastening germination of some woody plant seeds with
impermeable seed coats. Combined Proceedings of the International
Plant Propagators Society 17: 223230.
Frett JL, Dirr MA. 1979. Scarification and stratification requirements for
seeds of Cercis canadensis L., (redbud), Cladrastis lutea (Michx. F.) C. Koch
(yellowwood), and Gymnocladus dioicus (L.) C. Koch (Kentucky coffee
tree). Plant Propagator 25(2): 46.
Geneve RL. 1991a. Eastern redbud (Cercis canadensis L.) and Judas tree
(Cercis siliquastrum L.). Biotechnology in Agriculture and Forestry 16:
142151.
Geneve RL. 1991b. Seed dormancy in eastern redbud (Cercis canadensis).
Journal of the American Society for Horticultural Science 116(1): 8588.
Hamilton DF, Carpenter PL. 1975. Regulation of seed dormancy in Cercis
canadensis L. Journal of the American Society for Horticultural Science
100(6): 653656.
Heit CE. 1967a. Propagation from seed: 6. Hardseededness: a critical factor.
American Nurseryman 125(10): 1012, 8896.
Heit CE. 1967b. Propagation from seed: 8. Fall planting of fruit and hardwood seeds. American Nurseryman 126(4): 1213, 8590.
Hopkins M. 1942. Cercis in North America. Rhodora 44: 193211.
Hosie RC. 1969. Native trees of Canada. 6th ed. Ottawa: Canadian
Forestry Service, Department of Fisheries and Forestry. 380 p.
Jepson WL. 1936. A flora of California.Volume 2, Capparidaceae to
Cornaceae. Berkeley: University of California. 684 p.
Cercis
379
380
RosaceaeRose family
Cercocarpus Kunth
mountain-mahogany
Stanley G. Kitchen
Mr. Kitchen is a botanist at the USDA Forest Services Rocky Mountain Research Station,
Shrub Sciences Laboratory, Provo, Utah
recovery following fire. Recovery of curlleaf mountainmahogany stands following fire is from seed only and can
be extremely slow. Because they are long-lived, produce an
extensive root system, and survive well on dry steep slopes,
mountain-mahogany plants play an important role in erosion
control. Nitrogen fixation in root nodules has been described
for both curlleaf (Lepper and Fleschner 1977) and true
mountain-mahoganies (Hoeppel and Wollum 1971), suggesting a significant role by these species in improving fertility
in otherwise infertile soils. The wood of curlleaf mountainmahogany is extremely dense and heavy and has had limited
use, primarily as fuel wood (Johnson 1970).
Geographic races and hybrids. Two distinct subspecies or varieties of curlleaf mountain-mahogany occur in
the western United States (Stutz 1990). Although considerable overlap in distribution exists, C. ledifolius ssp. ledifolius (formerly ssp. intercedens) has a more northeastern
distribution, whereas the distribution of ssp. intermontanus
is centered to the west of its sister taxon. In northern Idaho,
northern Wyoming, and southern Montana, ssp. ledifolius is
the only mountain- mahogany taxon present (Stutz 1990).
The leaves of ssp. ledifolius plants differ from those of ssp.
intermontanus in being narrower, more strongly involute,
and densely pubescent ventrally. The leaves of ssp. intermontanus are broadly elliptic and glabrous. Habit of ssp.
Common name(s)
C. ledifolius Nutt.
curlleaf mountain-mahogany,
curlleaf cercocarpus, curlleaf mahogany,
desert mahogany
true mountain-mahogany, mountain
cercocarpus, birchleaf cercocarpus,
birchleaf mountain-mahogany,
alderleaf mountain-mahogany,
blackbrush, deerbrush, tallowbrush
C. montanus Raf.
C. betuloides Nutt.
C. parvifolius Nutt.
C. flabellifolius Rydb.
Occurrence
Washington & Oregon E to Montana
& Wyoming, S to Arizona, California,
& Mexico (Baja)
Oregon E to South Dakota S to Mexico,
incl. parts of Wyoming, Colorado, Nebraska,
Kansas,Texas, New Mexico, Arizona,
Utah, & California
Cercocarpus
381
382
held hoppers using a beating stick. During harvest and handling, short hairs dislodge from the fruits. These hairs cause
considerable discomfort to eyes and skin, thus the cowboy
epithet of hell feathers (Plummer and others 1968). Fruits
may collect in harvestable depths on the ground during
years of superior production. However, collections from
ground accumulations are often of poor quality due to the
removal of viable seeds by rodents.
Cleaning and storage. Highest purity values are
obtained by removing most broken branches from fruits during collection. For large collections, empty fruits, styles, and
fine hairs are best removed using a variable-speed debearder
and a 9.5-mm (#2) screen fanning mill (figure 2).
Hammermilling causes excessive breakage and should not
be used. Minimum standards accepted by the Utah Division
of Wildlife Resources for both species are purity values of
95%, and viability values of 85% (Jorgensen 1995).
Cleaned-fruit sizes differ by species, ecotype, and year
of collection. In one study, average number of fruits per
weight for curlleaf (8 collections) and true mountainmahoganies (10 collections) was 106,000 and 88,000/kg
(48,000 and 40,000/lb), respectively (Kitchen and others
1989a&b). These fruit weights were either equivalent to or
somewhat heavier than those previously reported
(Deitschman 1974). Curlleaf and true mountain-mahogany
fruits stored under warehouse conditions experienced no significant loss of viability during 15 and 7 years, respectively
(Stevens and others 1981).
Germination. Reported germination responses to
moist chilling for curlleaf mountain-mahogany range from
no response after 12 weeks (Young and others 1978), to
good germination with 4 weeks (Heit 1970). In most of
these studies, interpretation of results is difficult because
fruit fill percentage was not determined. Dealy (1975)
reported 20% germination in response to 60 days of moist
Figure 4 Cercocarpus montanus, true mountainmahogany: seedling with primary leaves and welldeveloped secondary leaves.
Cercocarpus
383
Nursery and field practice. Curlleaf and true mountain-mahoganies were first cultivated in 1879 and 1872,
respectively (Deitschman and others 1974). Bareroot and
container nursery stock are commercially available for both
species, generally as 1- or 2-year-old stock. Unless nondormant collections are used, cleaned fruits are either prechilled
or fall-sown. Seedbeds should be kept moist until seeds have
germinated (Deitschman and others 1974). Deep-rooting
containers filled with a minimum of 0.2 liter (13 in3) stan-
dard potting mix is recommended for container stock production (Landis and Simonich 1984). With optimum rearing
conditions a minimum of 4 to 6 months is required to develop an adequate root system. Figure 4 illustrates a seedling
with well-developed secondary leaves. Direct seeding of
mountain-mahogany should be carried out in fall or early
winter in conjunction with seedbed preparations that minimize competition to first-year seedlings (Plummer and
others 1968).
References
Davis JN. 1990. General ecology, wildlife use, and management of the
mountain mahoganies in the Intermountain West. In: Johnson KL, ed.The
genus Cercocarpus. Proceedings, 5th Utah Shrub Ecology Workshop;
1988 July 1314; Logan, UT. Logan: Utah State University: 113.
Dealy JE. 1975. Ecology of curlleaf mountain-mahogany (Cercocarpus ledifolius Nutt.) in Oregon and adjacent areas [PhD dissertation]. Corvallis:
Oregon State University. 162 p.
Deitschman GH, Jorgensen KR, Plummer AP. 1974. Cercocarpus H. B. K.,
cercocarpus (mountain-mahogany). In: Schopmeyer CS, tech. coord.
Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 309312.
Heit CE. 1970. Germination characteristics and optimum testing methods
for twelve western shrub species. Proceedings, Association of Official
Seed Analysts 60: 197205.
Hoeppel RE, Wollum AG II. 1971. Histological studies of ectomycorrhizae
and root nodules from Cercocarpus montanus and Cercocarpus paucidentatus. Canadian Journal of Botany 49: 13151318.
Johnson CM. 1970. Common native trees of Utah. Spec. Rep. 22. Logan:
Utah State University, Cooperative Extension Service. 109 p.
Jorgensen KR. 1995. Personal communication. Ephraim, UT: Utah Division
of Wildlife Resources.
Kitchen SG, Meyer SE, Wilson GR, Stevens R. 1989a. Addition of
Cercocarpus ledifoliuscurlleaf mountain-mahoganyto the rules.
Association of Official Seed Analysts Newsletter 63: 2628.
Kitchen SG, Meyer SE, Wilson GR, Stevens R. 1989b. Addition of
Cercocarpus montanustrue mountain-mahoganyto the rules.
Association of Official Seed Analysts Newsletter 63: 2830.
384
Kitchen SG, Meyer SE. 1990. Seed dormancy in two species of mountainmahogany (Cercocarpus ledifolius and Cercocarpus montanus). In: Johnson
KL, ed. The genus Cercocarpus. Proceedings, 5th Utah Shrub Ecology
Workshop; 1988 July 1314; Logan, UT. Logan: Utah State University:
2742.
Landis TD, Simonich EJ. 1984. Producing native plants as container seedlings.
In:The challenge of producing native plants for the Intermountain area.
Proceedings, Intermountain Nurserymans Association Conference; 1983
August 811; Las Vegas, NV. Gen.Tech. Rep. INT-168. Ogden, UT: USDA
Forest Service, Intermountain Research Station: 1625.
Lepper MG, Fleschner M. 1977. Nitrogen fixation by Cercocarpus ledifolius
(Rosaceae) in pioneer habitats. Oecologia (Berl.) 27: 333338.
Plummer AP, Christensen DR, Monsen SB. 1968. Restoring big-game range
in Utah. Pub. 68-3. Utah Division of Fish and Game. 183 p.
Schultz BW,Tueller PT,Tausch RJ. 1990. Ecology of curlleaf mahogany in
western and central Nevada: community and population structure.
Journal of Range Management 43: 1320.
Stevens R, Jorgensen KR, Davis JN. 1981. Viability of seed from thirty-two
shrub and forb species through fifteen years of warehouse storage.
Great Basin Naturalist 41: 274277.
Stidham ND, Ahring RM, Powell J, Claypool PL. 1980. Chemical scarification,
moist prechilling, and thiourea effects on germination of 18 shrub
species. Journal of Range Management 33: 115118.
Stutz HC. 1990. Taxonomy and evolution of Cercocarpus in the western
United States. In: Johnson KL, ed.The genus Cercocarpus. Proceedings, 5th
Utah Shrub Ecology Workshop; 1988 July 1314; Logan, UT. Logan: Utah
State University: 1525.
Young JA, Evans RA, Neal DL. 1978. Treatment of curlleaf cercocarpus
seeds to enhance germination. Journal of Wildlife Management 42:
614620.
RosaceaeRose family
achene (left)
longitudinal
Chamaebatia
385
References
Adams RS. 1969. How to cure mountain misery. Sacramento: California
Division of Forestry. 23 p.
Bailey LH. 1928. The standard cyclopedia for horticulture. New York:
Macmillan. 3639 p.
Dayton WA. 1931. Important western browse plants. Misc. Pub. 101.
Washington, DC: USDA. 214 p.
Emery D. 1964. Seed propagation of native California plants. Santa
Barbara Botanical Garden Leaflet 1(10): 8196.
386
RosaceaeRose family
Chamaebatiaria
387
follicle.
seeds.
388
longitudi-
Source
Nampa, ID
Elevation
(m)
Origin
831
Unknown
Sun Valley, ID
1,773
Unknown
Santa Fe, NM
2,134
W New Mexico
Cold,
wet chill
(days)*
% Germination
15 C
20/10 C
Incub
Incub
0
28
0
28
0
28
3
72
12
33
9
58
1
65
20
44
22
60
Seed
fill (%)
100
100
100
100
100
100
Seed
viability
(%)
96
96
86
86
85
85
* Chilling temperature = 3 to 5 C.
Incub = incubation time = 28 days; seeds were exposed to 8 hours of light (PAR = 350 M m/sec) each day with temperatures of either constant 15 C or 8 hours of
20 C and 16 hours of 10 C. In the alternating temperature regime, plants were exposed to light during the high-temperature period.
Based on the percentage of viable seeds to germinate normally.
The Nampa and Sun Valley, ID, plants were grown from the same unknown seed source.
References
Albee BJ, Schultz LM, Goodrich S. 1988. Atlas of the vascular plants of
Utah. Occ. Pub. 7. Salt Lake City: Utah Museum of Natural History.
670 p.
Bailey LH. 1902. Cyclopedia of American horticulture.Volume 4, RZ. New
York: Macmillan: 1686 p.
Davis RJ. 1952. Flora of Idaho. Provo, UT: Brigham Young University Press.
836 p.
Eggler WA. 1941. Primary succession on volcanic deposits in southern
Idaho. Ecological Monographs 3: 277298.
Everett PC. 1957. A summary of the culture of California plants at the
Rancho Santa Ana Botanic Garden. Claremont, CA: Rancho Santa Ana
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Fikenscher LH, Hegnauer R, Ruijgrok HWL. 1981. Distribution of hydrocyanic-acid in Cormophyta: 15. New observations on cyanogenesis in
Rosaceae. Planta Medica 41: 313327.
Hawksworth FG, Mathiasen RL. 1978. Hosts of juniper mistletoe at
Walnut Canyon National Monument, Arizona, USA. Great Basin
Naturalist 38: 89.
Hickman JC, ed. 1993. The Jepson manual higher plants of California.
Berkeley: University of California Press. 1400 p.
Hitchcock CL, Cronquist A, Ownbey M,Thompson JW. 1961. Vascular plants
of the Pacific Northwest: 3. Saxifragaceae to Ericaceae. Seattle: University
of Washington Press. 614 p.
Hurd EG. 1995. Unpublished data, 19941995. Boise, ID: USDA Forest
Service, Intermountain Research Station.
Chamaebatiaria
389
Kirkwood JE. 1930. Northern Rocky Mountain trees and shrubs. Stanford,
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Mackie F. 1995. Personal communication. Sun Valley, ID.
McArthur ED. 1984. Natural diversity of western range shrubs. In: Cooley
JL, Cooley JH, eds. Proceedings, Workshop on Natural Diversity in
Forest Ecosystems; 1982 November 29December 1; Athens, GA.
Athens: University of Georgia, Institute of Ecology: 193209.
McArthur ED, Sanderson SC. 1985. A cytotaxonomic contribution to the
western North American rosaceous flora. Madroo 32: 2428.
McArthur ED, Stutz HC, Sanderson SC. 1983. Taxonomy, distribution, and
cytogenetics of Purshia, Cowania, and Fallugia (Rosoideae, Rosaceae). In:
Tiedemann AR, Johnson KL, comps. Proceedings, Symposium on
Research and Management of Bitterbrush and Cliffrose in Western
North America; 1982 April 1315; Salt Lake City, UT. Gen.Tech. Rep.
INT-152. Ogden, UT: USDA Forest Service, Intermountain Forest and
Range Experiment Station: 424.
McDorman W. 1994. Unpublished data. Ketchum, ID: High Altitude
Gardens.
Merkle J. 1952. An analysis of a pinyonjuniper community at Grand
Canyon, Arizona. Ecology 33: 375384.
Mozingo HN. 1987. Shrubs of the Great Basin. Reno: University of
Nevada Press. 342 p.
Munz PA, Keck DD. 1959. A California flora. Berkeley: University of
California Press. 1681 p.
390
Peck ME. 1961. A manual of the higher plants of Oregon. 2d ed. Portland,
OR: Binfords & Mort. 936 p.
Peters J. 2000. Tetrazolium testing handbook. Contrib. 29. Las Cruces, NM:
Association of Official Seed Analysts. unpaged.
Phillips J. 1949. Southwestern landscaping with native plants. Santa Fe:
Museum of New Mexico Press. 140 p.
Rehder, A. 1940. Manual of cultivated trees and shrubs. New York:
Macmillan. 996 p.
Shaw NL. 1995. Unpublished data, 19921995. Boise, ID: USDA Forest
Service, Intermountain Research Station.
Van Dersal WR. 1938. Native woody plants of the United States: their erosion control and wildlife values. Misc. Pub. 303. Washington, DC: Civilian
Conservation Corps. 362 p.
Welsh SL, Atwood ND, Higgins LC, Goodrich S, eds. 1987. A Utah flora.
Great Basin Naturalist Memoirs 9: 1894.
Wolfe JA, Shorn HE. 1989. Paleoecologic, paleoclimatic, and evolutionary
significance of the Oligocene Creede flora, Colorado. Paleobiology 15:
180198.
Wolfe JA, Wehr W. 1988. Rosaceous Chamaebatiaria-like foliage from the
Paleogene of western North America. Aliso 12: 177200.
Young JA,Young CG. 1986. Collecting, processing and germinating seeds of
wildland plants. Portland, OR:Timber Press. 236 p.
Young JA,Young CG. 1992. Seeds of woody plants in North America.
Portland, OR: Dioscorides Press. 407 p.
CupressaceaeCypress family
Chamaecyparis Spach
white-cedar
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Growth habit, occurrence, and use. The genus
Chamaecyparis occurs naturally on the Atlantic and Pacific
Coasts of North America and in Japan and Taiwan. Three
species are native to North America, 2 to Japan, and 1 to
Taiwan (Sargent 1965). The North American species
(table 1) are long-lived evergreens that attain large size.
Port-Orford-cedar, the largest, has reached diameters of
more than 1 m and heights of near 70 m in old-growth
stands (Zobel 1990a). Branching is distinctive, with manybranched twigs and small paired scalelike leaves arranged in
fernlike sprays. Another common name is false cypress
(Little 1979); they are not true cedars (Cedrus spp.).
Because of their somber beauty and variety of form, whitecedars are often used for ornamental plantings, hedges, and
windbreaks (Rehder 1940). They produce valuable timber,
the wood being sought for poles, posts, construction timbers,
specialty items, and other uses where durability is desired.
Atlantic white-cedar wood is especially popular for boats,
outdoor furniture, posts, and utility poles (Kuser and
Zimmerman 1995).
Geographic races and hybrids. Two geographic
races of Atlantic white-cedar have been proposed: var.
henryae (Li) Little in Georgia, Florida, Alabama, and
Mississippi and var. thyoides in the area from South
Carolina to Maine (Little 1966). Great variation exists within the genus, and numerous horticultural selections have
been made of the 3 North America species as well as the
Asian ones (Dirr and Heuser 1987; Harris 1990; Little and
Garrett 1990; Zobel 1990a). Both interspecific and intergeneric crosses have been successful with certain of the
cedars. Alaska-cedar and 2 of the Asian species have been
crossed (Yamamoto 1981), and Alaska-cedar has also been
crossed with several species of Cupressus (Harris 1990). The
most well-known of these crosses is with Monterey cypress
(Cupressus macrocarpa Hartw. ex Gord.) to produce the
widely planted Leyland cypress (Cupressocyparis leylandii).
Flowering and fruiting. White-cedars are monoecious. Their tiny inconspicuous yellow or reddish male
pollen-bearing flowers and greenish female flowers are
borne on the tips of branchlets (Harris 1974). Staminate
flowers of Atlantic white-cedar, for example, are about
3 mm long, and the pistillate flowers are approximately
3 mm in diameter (Little and Garrett 1990). Pollination
occurs generally from March to May, and cones ripen in
September to October. Cones are slow to open fully, and
seed dispersal occurs from fall into the following spring
(table 2). Cones of Port-Orford-cedar and Atlantic white-
Common name(s)
Occurrence
Atlantic white-cedar,
white-cedar, swamp-cedar,
southern white-cedar
Chamaecyparis
391
Location
Flowering
Cone ripening
Seed dispersal
C. lawsoniana
C. nootkatensis
C. thyoides
Oregon
Pacific Coast
Atlantic Coast
March
AprMay
MarApr
SeptOct
SeptOct*
SeptOct
SeptMay
Octspring
Oct 15Mar 1
cedar mature the same year that they are pollinated, whereas
cones of Alaska-cedar, in most of the species range, take a
second year to complete maturation (Harris 1974, 1990). In
the extreme southern portion of the range of Alaska-cedar,
cones may mature in only 1 year (Owens and Molder 1975).
This condition even occurs on trees from more northern origins or from higher elevations when established in seed
orchards in warm, southern, coastal sites (El-Kassaby and
others 1991). The seeds from these 1-year cones are of size
and germination quality equal to seeds from 2-year cones.
The white-cedars bear cones at an early age5 to 20
years for Port-Orford-cedar (Zobel 1990a) and 3 to 5 years
for Atlantic white-cedar (Little and Garrett 1990). Sprays of
gibberellin (primarily GA3) will induce flowering in even
younger seedlings of Port-Orford-cedar and Alaska-cedar
(Owens and Molder 1977; Pharis and Kuo 1977). The use
of GA3 and supplemental pollination on container-grown
Port-Ordford-cedar trees 4 to 6 years from rooting or grafting has shown good potential to produce a large amount of
seeds in a short period (Elliott and Sniezko 2000). Mature
cones are 6 to 12 mm in diameter, spherical, and are borne
erect on branchlets (figure 1). Cones have from 6 to 12
scales, each bearing from 1 to 5 seeds with thin marginal
wings (figures 2 and 3) (Harris 1974). The average number
of seeds per cone is 7 for Alaska-cedar (Harris 1990) and 8
for Atlantic white-cedar, but less than a third of these seeds
may be filled. With controlled crosses in a seed orchard,
Port-Orford-cedar averaged as high as 8.6 filled seeds per
cone (Elliott and Sniezko 2000). Cone ripeness is normally
determined by their exterior color (table 3).
Seedcrops of both western white-cedars can be damaged by larvae of the seedworm Laspeyresia cupressana
(Kearfott) feeding on seeds in the cones. Larvae of the
incense-cedar tip mothArgyresthia libocedrella Busck
mine the cones and seeds of Port-Orford-cedar and can
destroy practically the entire seedcrop (Hedlin and others
1980).
Collection of cones. Cones may be collected by hand
or raked from the branchlets of standing or felled trees. As
392
Table 3Chamaecyparis, white-cedar: height, seed-bearing age, seed crop frequency, and color of ripe cones
Species
C. lawsoniana
C. nootkatensis
C. thyoides
Height at
maturity
(m)
Year first
cultivated
to 73
to 53
1227
1854
1851
1727
320
35
4 or more
1 or more
Sources: Little (1950), Korstian and Brush (1931), Ouden (1965), Rehder (1940), Sargent (1965).
8- to 10-year-old plantations of Atlantic white-cedar produced good seedcrops that were easy to collect (Bonner and
Summerville 1999). When collecting cones of Alaska-cedar
in the northern part of the range, precautions are needed to
limit the collection to mature, second-year cones. The smaller, greenish-blue, immature, first-year cones are often present on the same branches with the yellow-brown mature
cones (Harris 1974).
Extraction, cleaning, and storage of seeds. Cones of
white-cedars may be dried by spreading them in the sun or
in a warm room, or they may be kiln-dried at temperatures
below 43 C (Harris 1974). Over 90% of the seeds can be
recovered from cones of Atlantic white-cedar dried at 35 to
40 C if 2 or 3 cycles of drying, interspersed with re-wetting
of the cones, are used (Bonner and Summerville 1999).
Each time the cones are redried, they open a little more.
Mature cones of all white-cedars open when dried properly,
and their seeds may be extracted by gentle shaking or tumbling. The thin-coated seeds of all species are easily injured
and de-winging should not be attempted (Harris 1974).
Chamaecyparis
393
Seed wt/
cone wt
Range
/kg
20
20
176,4001,323,000
145,500396,900
926,1001,102,500
Average
/lb
80,000600,000
66,000180,000
420,000500,000
/kg
/lb
463,000
238,140
1,014,300
210,000
108,000
460,000
Table 5Chamaecyparis, white-cedar: stratification periods and germination test conditions and results
Test conditions
Stratification (days) Temp (C)
Species
C. lawsoniana
C. nootkatensis
C. thyoides
Warm*
0
0
58
0
0
0
0
Cold
0
0
30
3090
0
0
90
Day Night
30
30
30
30
30
30
30
20
20
20
20
20
20
20
Germ energy
Days
Test results
Germ
capacity Soundness
(%)
Days
(%)
(%)
Samples
44
24
10
0
0
14
34
11
48
52
12
0
0
84
48
51
57
54
9
60
1
3
8
11
28
60
22
41
2855
60
28
References
Aldhous JR. 1972. Nursery practice. For. Comm. Bull. 43. London: Forestry
Commission. 184 p.
Allen GS. 1957. Storage behavior ofconifer seeds in sealed containers held
at 0 F., 32 F., and room temperature. Journal of Forestry 55: 278281.
Bonner FT, Summerville KO. 1999. Production and quality of Atlantic
white-cedar seed in coastal North Carolina. In: Shear TH, Summerville
KO, eds. Symposium, Atlantic white-cedar: Ecology and Management.
1997 August 67; Newport News,VA. Gen.Tech. Rep. SRS-27. Asheville,
NC: USDA Forest Service, Southern Research Station. 7679.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Elliott L, Sniezko RA. 2000. Cone and seed production in a Port-Orfordcedar containerized orchard. In: Hansen EM, Sutton W, eds. Phytophthora
diseases of forest trees. IUFRO WP7.02.09. Proceedings, 1st
International Meeting on Phytophthoras in Forest and Wildland
Ecosystems. 1999 Auugust 30September 3; Grants Pass, OR. Corvallis:
Oregon State University: 105106.
El-Kassaby YA, Maze J, MacLeod DA, Banerjee S. 1991. Reproductive cycle
plasticity in yellow cedar (Chamaecyparis nootkatensis). Canadian Journal
of Forestry Research 21: 13601364.
Harris AS. 1974. Chamaecyparis Spach, white-cedar. In: Schopmeyer CS,
tech. coord. Seeds of woody plants in the United States. Agric. Handbk.
450. Washington, DC: USDA Forest Service: 316320.
Harris AS. 1990. Chamaecyparis nootkatensis (D. Don) Spach, Alaska-cedar.
In: Burns RM, Honkala BH, tech. coord. Silvics of North America.
Volume 1, Conifers. Agric. Handbk. 654. Washington, DC: USDA Forest
Service: 97102.
Hedlin AF,Yates HO III,Tovar DC, Ebel BH, Koerber TW, Merkel EP. 1980.
Cone and seed insects of North American conifers. FAO North
American Forestry Commission, Study Group on Forest Insects and
Diseases. Ottawa: Canadian Forestry Service. 122 p.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing. Rules 1993. Seed Science and Technology 21 (Suppl.):
1259.
Karlsson I. 1982. Propagation of Alaska yellow cedar (Chamaecyparis
nootkatensis [D. Don] Spach.) by rooted cuttings for production planting. Proceedings of the International Plant Propagators Society
31(1981): 112116.
Korstian CF, Brush WD. 1931. Southern white cedar. Agric.Tech. Bull. 251.
Washington, DC: USDA. 75 p.
Kuser JE, Zimmerman GL. 1995. Restoring Atlantic white-cedar swamps:
techniques for propagation and establishment.Tree Planters Notes
46(3): 7885.
Little EL Jr. 1966. Varietal transfers in Cupressus and Chamaecyparis.
Madroo 18(6): 161167.
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Little S Jr. 1940. Seed fall of Atlantic white-cedar. Tech. Note 26.
Philadelphia: USDA Forest Service, Allegheny Forest Experiment Station.
1 p.
Little S Jr. 1950. Ecology and silviculture of white-cedar and associated
hardwoods in southern New Jersey. Yale University School of Forestry
Bulletin 56: 1103.
Little S Jr, Garrett PW. 1990. Chamaecyparis thyoides (L.) B.S.P., Atlantic
white-cedar. In: Burns RM, Honkala BH, tech. coord. Silvics of North
America.Volume 1, Conifers. Agric. Handbk. 654. Washington, DC: USDA
Forest Service: 103108.
Ouden PD. 1965. Manual of cultivated conifers.The Hague: Martinus
Nijhoff. 526 p.
Owens JN, Molder M. 1975. Pollination, female gametophyte, and embryo
and seed development in yellow cedar (Chamaecyparis nootkatensis).
Canadian Journal of Botany 53: 186199.
Owens JN, Molder M. 1977. Cone induction in yellow cypress
(Chamaecyparis nootkatensis) by gibberellin A3 and the subsequent
development of seeds within the induced cones. Canadian Journal of
Forest Research 7: 605613.
Pharis RP, Kuo CG. 1977. Physiology of gibberellins in conifers. Canadian
Journal of Forest Research 7: 299325.
Rehder A. 1940. Manual of cultivated trees and shrubs hardy in North
America. 2nd ed. New York: Macmillan. 996 p.
Sargent CS. 1965. Manual of the trees of North America (exclusive of
Mexico). 2nd ed., corrected and reprinted. New York: Dover. 934 p.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA Soil
Conservation Service. 198 p.
Van Dersal WR. 1938. Native woody plants of the United States: their erosion-control and wildlife values. Misc. Pub. 303. Washington, DC: USDA.
362 p.
Yamamoto C. 1981. [in Japanese, English summary: Possibilities of interspecific hybridization between Chamaecyparis lawsoniana and some other
Chamaecyparis species]. Journal of the Japanese Forestry Society 63:
311319.
Zobel DB. 1979. Seed production in forests of Chamaecyparis lawsoniana.
Canadian Journal of Forest Research 9: 327335.
Zobel DB. 1990a. Chamaecyparis lawsoniana (A. Murr.) Parl., Port-Orfordcedar. In: Burns RM, Honkala BH, tech. coord. Silvics of North America.
Volume 1, Conifers. Agric. Handbk. 654. Washington, DC: USDA Forest
Service: 8896.
Zobel DB. 1990b. Effects of low temperature, seed source, and seed age
on germination of Chamaecyparis lawsoniana. Canadian Journal of Forest
Research 20: 10531059.
Chamaecyparis
395
BignoniaceaeTrumpet-creeper family
Synonyms. C. saligna D. Don, C. linearis var. originaria Fosberg, C. linearis var. glutinosa (Engelm.) Fosberg,
C. linearis var. arcuata Fosberg.
Other common names. false-willow, jano, floweringwillow, desert catalpa, catalpa-willow.
Growth habit, occurrence, and use. Desert-willow
grows along dry washes and streams in the desert between
450 and 1,500 m of elevation from southern California
through southern Utah to western Texas and southward into
Mexico and Lower California. It is a deciduous shrub or
small tree that attains heights of 3 to 7.5 m or occasionally
more. Growth can be rapid, up to 1 m annually (Munz
1979). The plant is useful for wildlife cover, erosion control,
restoration, stream stabilization, and ornamental plantings in
arid regions (McMinn 1959; Bainbridge and Virginia 1989;
Munz 1959). Seed pods and flowers are edible, but the
major use for Native American people was the wood (for
house frames, granaries, and bows) and the fibrous bark (for
weaving nets, shirts, and breechclouts) (Bainbridge and
Virginia 1989).
Flowering and fruiting. Desert-willow produces perfect flowers between April and August throughout its range
(Magill 1974; McMinn 1959). The fruit is a 2-celled capsule
about 6 mm in diameter and from 10 to 30 cm long. It
ripens from late summer to late fall (Afanasiev 1942) and
persists through winter (Little 1950). The numerous lightbrown oval seeds are about 8 mm long and have a fringe of
soft white hairs on each end (figures 1 and 2).
Collection, extraction, and storage. Seedpods can be
hand-picked after late September and through the winter
months. Care must be taken not to pick unripened fruits
the fruits on a tree may mature unevenly because of their
long flowering period (Engstrom and Stoeckeler 1941). Seed
396
seed.
Nursery field practice and seedling care. Desertwillow seeds may decay unless sown in spring soon after the
soil warms up. Sowing depth should be 6 mm (1/4 in).
A ratio of seven times as many viable seeds as the desired
number of usable seedlings is required to grow nursery
stock (Magill 1974). Damping-off can be a problem.
Desert-willow may also be propagated from cuttings
(Magill 1974); cuttings should be handled carefully and
allowed to produce an extensive rootball before transplanting. Mature plants grown in ~57-liter (15-gal) pots and
0.8-m (30-in) tubes have been successfully outplanted as
windbreaks and for desert restoration at Joshua Tree
National Park (CALR 1993).
longitudinal
References
Chilopsis
397
PyrolaceaeShinleaf family
Chimaphila Pursh
chimaphila
Don Minore
Dr. Minore retired from the USDA Forest Services Pacific Northwest Research Station
Common name(s)
Distribution
C. japonica Miq.
Japanese chimaphila
striped pipsissewa,
striped princes-pine,
spotted wintergreen
little pipsissewa,
little princes-pine
pipsissewa,
princes-pine
Sources: Barrett and Helenurm (1987), Blake (1914), Camp (1939), Fernald (1950), Hill (1962), Nordal and Wischmann (1989), Ohwi (1965), Prain (1960),Traulau (1981).
398
1968). They have 5-parted calyxes, 5 petals, 10 stamens, 5chambered ovaries, and short thick styles with wide, 5pointed stigmas (Krssmann 1984). The ovary is superior,
and there is a well-developed, collar-like disk at the base of
the pistil that secretes nectar. Placentation is central-axile,
with a massive, 2-lobed placenta intruding into each locule
(Pyykko 1968). Those placentae are beset with numerous
minute ovules (Copeland 1947). The 1 to 3 (little pipsissewa
and Japanese chimaphila) or 2 to 6 (striped pipsissewa and
pipsissewa) flowers are borne in pendulous, terminal inflorescences (Krssmann 1984; Ohwi 1965). In pipsissewa
those inflorescences are corymbs; in striped and little pipsissewas, they are cyme-like clusters (Copeland 1947). Flowers
are pink or white and slightly fragrant.
Chimaphila pollen grains are packed into polyads composed of indefinite numbers of tetrads (Knudsen and Olesen
1993; Takahashi 1986). Pollination is by insects. In pipsissewa, bumble bees are the most important pollinators but the
flowers also are visited by staphylinid beetles (Barrett and
Helenurm 1987; Knudsen and Olesen 1993), and there is a
small amount of self-pollination (Barrett and Helenurm
1987). Differences in the flower preferences of the bumble
bee species involved may help to prevent interbreeding
between pipsissewa and striped pipsissewa during a short
overlap in flowering periods where these inter-fertile species
grow together (Standley and others 1988).
An average pipsissewa flower produces 308,800 pollen
grains and 5,587 ovulesa pollen to ovule ratio of 58
(Barrett and Helenurm 1987). In central New Brunswick,
Canada, anthesis occurs over a 30-day period in July
(Helenurm and Barrett 1987). In the Pacific Northwest, pipsissewa flowers may be found from June until August
(Hitchcock and others 1959). The fruits matured in about 70
days in New Brunswick, where an average fruit weighed
23 mg, and fruit set was 75% for both self-pollinated and
cross-pollinated flowers (Barrett and Helenurm 1987).
The chimaphila fruit is a 5-celled, loculicidally dehiscent capsule that contains very large numbers of tiny seeds
(Barrett and Helenurm 1987; Copeland 1947; Pyykko 1968)
that sift out of the capsule openings to be borne away by the
wind. The embryos of those seeds develop no distinct parts
during seed development, but they eventually absorb all of
the endosperm except a single layer of cells. The inner
integumental cells die and remain in existence as more or
less shriveled empty spaces at the ends of the seeds
(Copeland 1947). The seedcoat consists only of the outer
cell layer of the integument, which loses protoplasm and
tannin to become transparent (Pyykko 1968). Although the
inner periclinal walls of those transparent testa cells are
smooth or slightly pitted in all species, intraspecific differences occur in pipsissewa (Takahashi 1993).
Chimaphila seeds are characterized by very small size,
few cells, and little differentiation (figure 1). Each contains
a central ellipsoidal mass consisting of an embryo covered
by a single layer of endosperm cells, surrounded by a transparent seedcoat that is hollow and shriveled at each end.
Ripe seeds are 0.6 to 0.9 mm long and 0.1 mm wide. There
are about 1,500,000 seeds/g (42,524,250/oz) (Minore 1994).
Collection and storage of seeds. Seeds can be collected in the field by tapping recently dehisced capsules to
dislodge the tiny seeds, which can then be captured in a jar
or plastic bag as they drift downward. Recently dehisced
capsules may be difficult to find, however, because mature
capsules often are not open or have already lost their seeds.
Mature closed capsules can be collected, dried, and macerated to recover the seeds. Unfortunately, this latter procedure
creates debris that is difficult to separate from the seeds, and
seed maturity is not assured. Optimal storage conditions and
seed longevity are unknown.
Figure 1Chimaphila umbellata, pipsissewa: seeds
(0.1 mm wide and ~ 0.7 mm long, at center) with central
embryo and elongate, transparent seedcoat.
Chimaphila
399
References
Altschul SVR. 1973. Drugs and foods from little-known plants. Notes in
Harvard University Herbaria. Cambridge, MA: Harvard University Press.
366 p.
Barrett SCH, Helenurm K. 1987. The reproductive biology of forest herbs:
1. Breeding systems and pollination. Canadian Journal of Botany 65(10):
20362046.
Blake SF. 1914. A new Chimaphila from San Domingo. Journal of Botany
(British and Foreign) 52: 169.
Camp WH. 1939. Studies in the Ericales: 4. Notes on Chimaphila, Gaultheria
and Pernettya in Mexico and adjacent regions. Bulletin of the Torrey
Botanical Club 66: 728.
Copeland HF. 1947. Observations on the structure and classification of the
Pyroleae. Madroo 9(3): 65102.
DiModica G,Tira S, Borello E. 1953. Su sostanze estratte da Chimaphila
corymbosa Pursh: struttura delia Chimafillina. Gazetta Chimica Italia
83(6): 393401 [Biological Abstracts 28: 679, Ref. 6848].
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1632 p.
Haber E, Cruise E. 1974. Generic limits in the Pyrolideae (Ericaceae).
Canadian Journal of Botany 52(4): 877883.
Helenurm K, Barrett SCH. 1987. The reproductive biology of forest herbs:
2. Phenology of flowering and fruiting. Canadian Journal of Botany
65(10): 20472056.
Hill AW, ed. 1962. Index Kewensis plantarum phanerogamarum: supplementum septimum. Oxford, UK: Oxford University Press. 260 p.
Hitchcock CL, Cronquist A, Ownbey M,Thompson JW. 1959. Vascular
plants of the Pacific Northwest: 4. Ericaceae through Campanulaceae.
Seattle: University of Washington Press. 510 p.
Holm T. 1927. The flower of Chimaphila. Rhodora 29(337): 16.
Knudsen JT, Olesen JM. 1993. Buzz-pollination and patterns in sexual traits
in North European Pyrolaceae. American Journal of Botany 80(8):
900913.
Kruckeberg AR. 1982. Gardening with native plants of the Pacific
Northwest: an illustrated guide. Seattle: University of Washington Press.
252 p.
Krssmann G. 1984. Manual of cultivated broad-leaved trees and shrubs.
Volume I, AD. Beaverton, OR:Timber Press. 448 p.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. Revised.
New York: Macmillan. 1290 p.
Lucia F. 1991. Princes pine medicinal action traced to specific chemical
compounds. In Good Tilth 1991 (August): 6.
400
OleaceaeOlive family
Chionanthus virginicus L.
white fringetree
John D. Gill, Franz L. Pogge, and Franklin T. Bonner
Dr. Gill and Mr. Pogge retired from the USDA Forest Services Northeastern Forest Experiment Station;
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Forest Experiment Station,
Mississippi State, Mississippi
fruit
Chionanthus
401
References
Barton LV. 1961. Seed preservation and longevity. Plant Science
Monographs. London: Leonard Hill Ltd. 216 p.
Brown CL, Kirkman LK. 1990. Trees of Georgia and adjacent states.
Portland, OR:Timber Press. 292 p.
Carpenter WJ, Ostmark ER, Sheehan TJ. 1992. Recommendations for germinating fringetree Chionanthus virginicus L. seed. Proceedings of the
Florida State Horticultural Society 104: 337340 [Seed Abstracts 1995;
18(5): 1542].
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seeds to tissue culture. Athens, GA:Varsity Press. 239 p.
Flemion F. 1941. Further studies on the rapid determination of the germinative capacity of seeds. Contributions of the Boyce Thompson Institute
11: 455464.
Gill JD, Pogge FL. 1974. Chionanthus virginicus L., fringetree. In: Schopmeyer
CS, tech. coord. Seeds of woody plants in the United States. Agric.
Handbk. 450. Washington, DC: USDA Forest Service: 323325.
Gleason HA. 1963. The new Britton and Brown illustrated flora of the
northeastern United States and adjacent Canada. 3 vol. New York:
Hafner Publishing.
Goodrum PD, Halls LK. 1961. Fringetree. In: Halls LK, Ripley TH, eds. Deer
browse plants of southern forests. New Orleans: USDA Forest Service,
Southern and Southeastern Forest Experiment Stations: 1011.
Heit CE. 1955. The excised embryo method for testing germination quality
of dormant seeds. Proceedings of the Association of Official Seed
Analysts 1955: 108117.
Heit CE. 1967. Propagation from seed: 8. Fall planting of fruit and hardwood seeds. American Nurseryman 126 (4): 1213, 8590.
Krochmal A, Walters RS, Doughty RM. 1969. A guide to medicinal plants of
Appalachia. Res. Pap. NE-138. Upper Darby, PA: USDA Forest Service.,
Northeastern Forest Experiment Station. 291 p.
Lay DW. 1961. Fruit production of some understory hardwoods.
Southeastern Association of Game and Fish Commissions Proceedings
15: 3037.
Rehder A. 1940. Manual of cultivated trees and shrubs. 2nd ed. New York:
Macmillan. 996 p.
Schumacher FW. 1962. How to grow seedlings of trees and shrubs. 2nd
ed. Sandwich, MA: F. W. Schumacher. 14 p.
Sheat WG. 1948. Propagation of trees, shrubs and conifers. London:
Macmillan. 479 p.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA Soil
Conservation Service. 187 p.
Van Dersal WR. 1938. Native woody plants of the United States: their erosion-control and wildlife values. Misc. Pub. 303. Washington, DC: USDA.
362 p.
Chionanthus
403
FagaceaeBeech family
Chrysolepis Hjelmqvist
chinquapin
Philip M. McDonald
Dr. McDonald is a research silviculturist at the USDA Forest Services Pacific Southwest Research Station,
Redding, California
Synonyms. The 2 species of Chrysolepis found in the
United States are distinct from their Asian relatives in the
genus Castanopsis and were placed by Hjelmqvist (1948) in
their current genus. This genus was accepted in Hickmans
extensive and long-researched flora of California (1993).
These American species, which have a floral morphology
that is intermediate between Castanopsis and Lithocarpus,
represent the ancient condition of the family Fagaceae
(McKee 1990). The common name also has changed
throughout the years. Early workers called the species chinquapin. Later, it became chinkapin but more recently it
was changed back to chinquapin (Hickman 1993; KeelerWolf 1988).
Occurrence and growth habit. In North America, the
genus Chrysolepis consists of 2 species and 1 variety
(Hickman 1993), all located in the Pacific Coast region.
Giant chinquapinChrysolepis chrysophylla var. chrysophylla (Dougl. ex. Hook) Hjelmqvist is a tree that ranges
from southwestern Washington southward to San Luis
Obispo County in the Cascade, Klamath, and Coastal
Mountains of California. A remnant stand also exists in El
Dorado County in the north central Sierra Nevada. This
species achieves its best form from Marin County,
California, northward (Griffin and Critchfield 1972) to Lane
County, Oregon. Giant chinquapin also has a shrub form
C. chrysophylla var. minor (Benth.) Munz, often called
golden chinquapinthat is found throughout the range of
its taller brethren.
The second speciesC. sempervirens (Kellogg)
Hjelmqvistwhich is always a shrub, has the common
name bush chinquapin. This species is found from the
Cascade Mountains of southern Oregon westward in the
Siskiyou Mountains of northern California, and southward
along the east-facing slopes of the north Coast Range and
the west-facing slopes of the Sierra Nevada, San Jacinto,
and San Bernardino Mountains (McMinn 1939). Throughout its range, the habitat is characterized as being of low
quality with shallow, rocky, and often droughty soils. In
404
Use. The light, fairly hard, and strong wood of chinquapin has been used for veneer, paneling, cabinets, furniture, turned products, pallets, and fuel (EDA 1968).
Flowering and fruiting. The flowers of giant chinquapin, which bloom from June through midwinter, and the
flowers of the shrubs, which bloom throughout the summer,
are unisexual, with staminate and pistillate flowers being
borne on the same plant. The staminate flowers are borne in
groups of 3 in the axils of bracts, forming densely flowered,
erect cylindrical catkins 2.5 to 7.6 cm long; 1 to 3 pistillate
flowers are borne in an involucre, usually at the base of the
staminate catkins or borne in short separate catkins. At the
time of peak blooming in June, each tree is covered with
erect creamy white blossoms, which provide a pleasing contrast to the more somber foliage (Peattie 1953).
The fruit consists of 1 to 3 nuts (figures 1 and 2)
enclosed in a spiny golden brown bur. The nuts mature in
fall of the second year (Hickman 1993). The minimum seedbearing age (from root crown sprouts) is 6 years (McKee
1990). Giant chinquapin trees have been reported as producing seeds at 40 to 50 years of age but probably do so before
this age (McKee 1990). Controversy exists over seed productivity. Sudworth (1908) reported that the tree form is an
abundant seeder, but Peattie (1953) noted that although
flowering is abundant, fruiting is strangely shy. Insects,
squirrels, and birds often consume most of a given crop.
Indeed, Powell (1994) observed tree squirrels (Sciurus spp.)
cutting burs of large chinquapins during a bumper seed year.
By late fall, the ground beneath the trees was covered with
burs.
Collection, extraction, and storage. Because of
heavy predation by many animals, collectors should handpick the burs in late summer or early fall, after they ripen
but before they open (Hubbard 1974). The collected burs
should be spread out to dry in the sun or in a warm room.
After drying, the nuts can be separated from the burs
mechanically. The following number of nuts per weight
have been recorded (Hubbard 1974; McMinn 1939):
giant chinquapin
(C. chrysophylla var. chrysophylla)
golden chinquapin
(C. chrysophylla var. minor)
bush chinquapin
(C. sempervirens)
Nuts/kg
1,8262,420
Nuts/lb
8301,100
1,540
700
2,640 1,200
longitudinal section
Chrysolepis
405
seedling at 1 month
References
EDA [Economic Development Administration]. 1968.
Nursery practice. Little is known about raising chinquapins in nurseries. In a study at the Rancho Santa Ana
Botanic Gardens in California, the 3 native species were
raised in pots. Some survived through 1 or more potting
stages, but none survived after outplantings (Hubbard 1974).
Propagation by layering, grafting, or budding is feasible
(Hubbard 1974).
Seedling care. Natural regeneration of giant chinquapin usually is sparse or totally lacking. Powell (1994)
noted that not a single seedling was present on ground covered with burs beneath large seed trees. McKee (1990) also
inferred that regeneration was lacking in environments of
deep litter and dense understory vegetation. Sudworth
(1908) noted that regeneration was best if seeds were covered, apparently by eroded soil. Keeler-Wolf (1988) found
sexually reproduced seedlings and saplings to average about
19/ha (7/ac) in the Klamath Mountains but only in shaded
mesic environments. In the Oregon Cascade Mountains,
McKee (1990) noted that chinquapin reproduction occurred
in light leaf mulch in partial shade, with plantlets that were
15 to 45 cm tall at ages 4 to 12. For bush chinquapin in the
northern Sierra Nevada on 10 study areas over a 10-year
period, not 1 seedling was found. Although tiny plants
looked like seedlings, a gentle tug showed that they were
connected to parent-plant root systems. The number of new
sprouts averaged over 39,000/ha (16,000/ac) 6 years after
site preparation by bulldozer bared the ground (McDonald
and others 1994).
406
The Hoopa
Valley Reservation hardwood study report. Washington, DC: U.S.
Department of Commerce: 154 p.
Griffin JR, Critchfield WG. 1972. The distribution of forest trees in
California. Res. Pap. PSW-82. Berkeley, CA: USDA Forest Service, Pacific
Southwest Forest and Range Experiment Station: 118 p.
Hickman JC, ed. 1993. The Jepson manual: higher plants of California.
Berkeley: University of California Press. 1400 p.
Hjelmqvist H. 1948. Studies on the floral morphology and phylogeny of the
Amentiferae. Botanical Notes Supplement 2(1): 1171.
Hubbard RL. 1974. Castanopsis (D.Don) Spach, chinkapin. In: Schopmeyer
CS. tech. coord. Seeds of woody plants in the United States. Agric.
Handbk. 450. Washington, DC: USDA Forest Service: 276277.
Keeler-Wolf T. 1988. The role of Chrysolepis chrysophylla (Fagaceae) in the
Pseudotsugahardwood forest of the Klamath Mountains of California.
Madrono 35(4): 285308.
McDonald PM, Abbott CS, Fiddler GO. 1994. Response of young ponderosa pines, shrubs, and ferns to three release treatments. Western
Journal of Applied Forestry 9(1): 2428.
McDonald PM, Minore D, Atzet T. 1983. Southwestern Oregonnorthern
California hardwoods. In: Burns RM, tech. comp. Silvicultural systems for
the major forest types of the United States. Agric. Handbk. 445.
Washington, DC: USDA Forest Service: 2932.
McKee A. 1990. Castanopsis chrysophylla (Dougl.) A. DC, giant chinkapin. In:
Burns RM, Honkala BH, tech. coords. Silvics of North America.Vol. 2,
Hardwoods. Agric. Handbk. 654. Washington, DC: USDA Forest Service:
234239.
McMinn HE. 1939. An illustrated manual of California shrubs. Berkeley:
University of California Press. 663 p.
Mirov NT, Kraebel CJ. 1937. Collecting and propagating the seeds of
California wild plants. Res. Note 18. Berkeley: USDA Forest Service,
California Forest and Range Experiment Station. 27 p.
Peattie DC. 1953. A natural history of western trees. Boston: HoughtonMifflin. 751 p.
Powell C. 1994. Personal communication. Big Bar, CA: USDA Forest
Service.
Sudworth GB. 1908. Forest trees of the Pacific slope. Washington, DC:
USDA Forest Service: 441 p.
Zobel DB, McKee A, Hawk GM, Dyrness CT. 1976. Relationships of environment to composition, structure, and diversity of forest communities
of the central western Cascades of Oregon. Ecological Monographs 46:
135156.
AsteraceaeAster family
Chrysothamnus Nutt.
rabbitbrush
Susan E. Meyer
Dr. Meyer is a research ecologist at the USDA Forest Services Intermountain Research Station,
Shrub Sciences Laboratory, Provo, Utah
Chrysothamnus
407
Table 1Chrysothamnus, rabbitbrush: ecology and distribution of some common species and subspecies
Taxon & species
SECTION CHRYSOTHAMNUS
C. linifolius Greene
Ericameria linifolia (Greene) L.C.Anders.
C. viscidiflorus (Hook.) Nutt.
E. viscidiflora (Hook.) L.C.Anders.
C. v. ssp. lanceolatus (Nutt.) Hall & Clements
E. viscidiflora spp. lanceolata (Nutt.) L.C.Anders.
C. v. ssp. viscidiflorus (Hook.) Nutt.
E. viscidiflora (Hook.) L.C.Anders.
Common name(s)
Geographic distribution
Habitat
spearleaf rabbitbrush,
alkali rabbitbrush
green rabbitbrush
Colorado Plateau N
to Montana
Intermountain
Douglas rabbitbrush
Intermountain
Montane
low rabbitbrush
Intermountain
Low to mid-elevation
rubber rabbitbrush
W North America
Wide amplitude
mountain whitestem
rubber rabbitbrush
basin whitestem rubber
rabbitbrush
threadleaf rubber rabbitbrush
Mostly Intermountain
Rocky Mtn
Mostly Great Basin
SECTION NAUSEOSI*
C. nauseosus (Pallas ex Pursh) Britt.
E. nauseosa (Pallas ex Pursh) Nesom & Baird
C. n. ssp. albicaulis (Nutt.) Hall & Clements
C. n. ssp. hololeucus (Gray) Hall & Clements
C. n. ssp. consimilis (Greene) Hall & Clements
E. nauseosa ssp. consilimus (Greene) Nesom & Baird
C. n. ssp. graveolens (Nutt.) Piper
C. n. ssp. salicifolius (Rydb.) Hall & Clements
C. parryi (Gray) Greene
E. parryi (Gray) Nesom & Baird
SECTION PUNCTATI*
C. teretifolius (Dur. & Hilg.) Hall
E. teretifolia (Dur. & Hilg.)
Mojave rabbitbrush,
Jepson green rabbitbrush
Mojave Desert
Rocky washes;
hot desert
Sources: Anderson (1995), Deitschman and others (1974), USDA NRCS (2001).
408
Chrysothamnus
409
Range
/kg
/lb
/kg
/lb
1.8
1.8
1.8
1.8
1.5
0.8
0.8
0.8
0.8
0.7
1.52.0
1.12.2
1.12.2
0.70.9
0.51.0
0.51.0
1.7
1.1
1.5
1.3
1.5
1.5
1.7
1.3
0.9
0.9
0.8
0.5
0.7
0.6
0.7
0.7
0.8
0.6
0.4
0.4
1.52.0
0.91.4
1.11.5
0.92.4
0.91.1
0-91.1
0.70.9
0.40.6
0.50.7
0.41.1
0.40.5
0.40.5
1.3
0.6
SECTION CHRYSOTHAMNUS
C. linifolius
C. viscidiflorus
ssp. lanceolatus
ssp. viscidiflorus
SECTION NAUSEOSI
C. nauseosus
ssp. albicaulis
ssp. hololeucus
ssp. consimilis
ssp. graveolens
ssp. salicifolius
C. parryi
SECTION PUNCTATI
C. teretifolius
Sources: Belcher (1985), Deitschman and others (1974), Meyer (1995, 1997), McArthur and others (2004).
* 100% purity.
410
less water than most other shrubs and should not be overwatered or fertilized excessively (Monsen 1966).
Rabbitbrushes are among the easiest shrubs to establish
from direct seeding, and most plantings on wildland sites
use this method. Minimal seedbed preparation is required.
Surface planting onto a firm but roughened seedbed in late
fall usually results in adequate stands. This planting may be
accomplished through aerial seeding; hand-broadcasting; or
seeding with a thimble seeder, seed dribbler, browse seeder,
or a drill with the drop tubes pulled so that the seed is
placed on the disturbed surface behind the disk furrow openers (McArthur and others 2004). Seeds should not be planted too deeply. One millimeter of soil coverage is sufficient.
Seeding rates of about 20 to 30 live seeds/m2 (2 to 3/ft2) are
usually adequate. This is equivalent to about 200 g/ha (ca 3
oz/ac) on a pure live seed basis for a seedlot that averages
1.5 million seeds/kg (680,400/lb). If the seedlot is cleaned to
high purity, it may be necessary to dilute it with a carrier
such as rice hulls in order to achieve uniform seeding rates.
Seedlings emerge in early spring, and young plants grow
rapidly, often producing seeds in their second growing
season.
Table 3Chrysothamnus, rabbitbrush: germination percentage (as percentage of total viable seeds) after 28 days at 15 C
or at 25 C, and days to 50% of total germination during 20 weeks at 3 C for some common species and subspecies
Species
SECTION CHRYSOTHAMNUS
C. linifolius
C. viscidiflorus
ssp. lanceolatus
ssp. viscidiflorus
Germination percentage
at 15 C
Mean
Range
No.
Germination percentage
at 25 C
Mean
Range
No.
21
37
58
2949
3198
3
3
64
68
5870
4096
3
3
60
60
3582
3582
5
5
37
75
70
76
25
2945
1797
2696
28100
955
2
8
6
7
6
85
91
91
91
65
7892
7496
80100
58100
4492
2
8
6
7
6
41
21
33
33
89
34
988
770
5108
10105
60100
1254
2
12
17
17
6
4
SECTION NAUSEOSI
C. nauseosus
ssp. albicaulis
ssp. hololeucus
ssp. consimilis
ssp. graveolens
ssp. salicifolius
C. parryi
SECTION PUNCTATI
C. teretifolius
Sources: Meyer (1997), Meyer and McArthur (1987), Meyer and others (1989).
Chrysothamnus
411
References
Anderson LC. 1986. An overview of the genus Chrysothamnus. In:
McArthur ED, Welsh BL, comp. Proceedings, Symposium on the Biology
of Artemisia and Chrysothamnus. Gen.Tech. Rep. INT-200. Ogden, UT:
USDA Forest Service, Intermountain Forest and Range Experiment
Station: 2945.
Anderson LC. 1995. The ChrysothamnusEricameria connection. Great
Basin Naturalist 55: 8488.
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
Belcher E. 1985. Handbook on seeds of browse-shrubs and forbs.Tech.
Pub. R8-TP8. Atlanta: USDA Forest Service, Southern Region. 246 p.
Deitschman GH, Jorgensen KR, Plummer AP. 1974. Chrysothamnus, rabbitbrush. In: Schopmeyer CS, ed. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington DC: USDA Forest Service:
326328.
Frischknecht NC. 1963. Contrasting effect of big sagebrush and rabbitbrush on the production of crested wheatgrass. Journal of Range
Management 16: 7074.
Johnson KL. 1987. The genus Chrysothamnus. In: Proceedings, 4th Utah
Shrub Ecology Workshop. Logan: Utah State University. 59 p.
Khan MA, Sankhla N, Weber DJ, McArthur ED. 1987. Seed germination
characteristics of Chrysothamnus nauseosus ssp. viridulus (Astereae,
Asteraceae). Great Basin Naturalist 47: 220226.
Long LE. 1986. Container nursery production of Artemisia and
Chrysothamnus species. In: McArthur ED, Welch BL, comps. Proceedings,
Symposium on the Biology of Artemisia and Chrysothamnus. Gen.Tech.
Rep. INT-200. Ogden, UT: USDA Forest Service, Intermountain Forest
and Range Experiment Station: 395396.
McArthur ED, Meyer SE. 1987. A review of the taxonomy and distribution
of Chrysothamnus. In: Johnson KL, ed. Proceedings, 4th Utah Shrub
Ecology Workshop.The genus Chrysothamnus. Logan: Utah State
University: 918.
McArthur ED, Stevens R, Shaw NL. 2004. Composite shrubs. In: Monsen
SB, Stevens R, eds. Restoring western ranges and wildlands. Ogden, UT:
USDA Forest Service, Rocky Mountain Research Station.
McArthur ED, Welch BL, comps. 1986. Proceedings, Symposium on the
Biology of Artemisia and Chrysothamnus. Gen.Tech. Rep. INT-200. Ogden,
UT: USDA Forest Service, Intermountain Forest and Range Experiment
Station. 398 p.
McArthur ED, Meyer SE, Weber DJ. 1987. Germination rate at low temperature: rubber rabbitbrush population differences. Journal of Range
Management 40: 530533.
McArthur ED, Blauer AC, Plummer AP, Stevens R. 1979. Characteristics and
hybridization of important Intermountain shrubs: 3. Sunflower family. Res.
Pap. INT-220. Ogden, UT: USDA Forest Service, Intermountain Forest
and Range Experiment Station: 182.
Meyer SE. 1990. Seed source differences in germination under snowpack in
northern Utah. In: Munkshower F, ed. 5th Billings Symposium on
Disturbed Land Reclamation.Volume 1. Pub. 9003. Bozeman: Montana
State University, Reclamation Research Unit: 184191.
Meyer SE. 1997. Ecological correlates of achene mass variation in
Chrysothamnus nauseosus (Asteraceae). American Journal of Botany 84:
471477.
412
Meyer SE, McArthur ED. 1987. Studies on the seed germination biology of
rubber rabbitbrush. In: Johnson KL, ed.The genus Chrysothamnus.
Proceedings, 4th Utah Shrub Ecology Workshop. Logan: Utah State
University: 1926.
Meyer SE, Monsen SB. 1990. Seed-source differences in initial establishment
for big sagebrush and rubber rabbitbrush. In: McArthur ED, Romney EM,
Smith SD,Tueller PT, comps. Proceedings, Symposium on Cheatgrass
Invasion, Shrub Die-off and Other Aspects of Shrub Biology and
Management. Gen.Tech. Rep. INT-276. Ogden, UT: USDA Forest Service,
Intermountain Research Station: 200208.
Meyer SE, McArthur ED, Jorgensen GL. 1989. Variation in germination
response to temperature in rubber rabbitbrush (Chrysothamnus nauseosus: Asteraceae) and its ecological implications. American Journal of
Botany 76: 981991.
Meyer SE, Wilson GR, Stevens R. 1989. Proposed rule for Chrysothamnus
nauseosus. Association of Official Seed Analysts Newsletter 63(1): 3032.
Monsen SB. 1996. Unpublished data on file. Provo. UT: USDA Forest
Service, Rocky Mountain Research Station.
Monsen SB, Meyer SE. 1990. Seeding equipment effects on establishment of
big sagebrush on mine disturbances. In: Munkshower F, ed. Fifth Billings
Symposium on Disturbed Land Rehabilitation.Volume 1. Pub. 9003.
Bozeman: Montana State University, Reclamation Research Unit:
192199.
Monsen SB, Stevens R. 1987. Seed and seeding characteristics of rabbitbrush. In: Johnson KL, ed.The genus Chrysothamnus. Proceedings, 4th
Utah Shrub Ecology Workshop. Logan: Utah State University: 4149 p.
Romo JT, Eddleman LE. 1988. Germination of green and gray rabbitbrush
and their establishment on coal mined land. Journal of Range
Management 41: 491494.
Stevens R, Jorgensen KR, Davis JN. 1981. Viability of seed from thirty-two
shrub and forb species through fifteen years of warehouse storage.
Great Basin Naturalist 41: 274277.
Stevens R, Jorgensen KR, Davis JN, Monsen SB. 1986. Seed pappus and
placement influences on white rubber rabbitbrush establishment. In:
McArthur ED, Welch BL, comps. Proceedings, Symposium on the Biology
of Artemisia and Chrysothamnus. Gen.Tech. Rep. INT-200. Ogden, UT:
USDA Forest Service, Intermountain Forest and Range Experiment
Station: 353357.
USDA NRCS [USDA National Resources Conservation Service]. 2001.
The PLANTS database, version 3.1. Baton Rouge, LA: National Plant
Data Center [http://plants.usda.gov].
Weber DJ, Hegerhorst DR, Davis TD, McArthur ED. 1987. Potential uses of
rubber rabbitbrush. In: Johnson KL, ed.The genus Chrysothamnus.
Proceedings, 4th Utah Shrub Ecology Workshop. Logan: Utah State
University: 2734.
Whisenant S. 1987. Improving herbicidal control of rubber rabbitbrush. In:
Johnson KL, ed.The genus Chrysothamnus. Proceedings, 4th Utah Shrub
Ecology Workshop. Logan: Utah State University: 3540.
Young JA, Evans RA. 1974. Population dynamics of green rabbitbrush in
disturbed big sagebrush communities.
FabaceaePea family
Cladastris
413
legumes.
References
414
Bailey LH. 1949. Manual of cultivated plants most commonly grown in the
continental United States and Canada. Rev. ed. New York: Macmillan.
1116 p.
Barton LV. 1947. Special studies on seed coat impermeability. Contributions
of the Boyce Thompson Institute 14: 355362.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1632 p.
Frett JL, Dirr MA. 1979. Scarification and stratification requirements for
seeds of Cercis canadensis L. (redbud), Cladrastis lutea (Michx. f.) K. Koch
(yellowwood), and Gymnocladus dioicus (L.) K. Koch. (Kentucky coffeetree). Plant Propagator 25(2): 46.
Harris JA. 1917. The weight of seeds as related to their number and
position.Torreya 17: 180182.
Heit CE. 1967. Propagation from seed: 6. Hard-seedednessa critical
factor. American Nurseryman 125(10): 1012, 8896.
Jenkins EM. 1936. Seed practices in nursery. American Nurseryman 63(11):
911.
Olson DF Jr, Barnes RL. 1974. Cladrastis lutea, yellowwood (Michx. f.) K.
Koch. In: Schopmeyer CS, tech. coord. Seeds of woody plants in the
United States. Agric. Handbk. 450. Washington, DC: USDA Forest
Service: 329330.
Radford AE, Ahles HE, Bell CR. 1964. Guide to the vascular flora of the
Carolinas. Chapel Hill: University of North Carolina Book Exchange.
Rivera R, Popp HW, Dow RB. 1937. The effect of high hydrostatic pressure
upon seed germination. American Journal of Botany 24: 508513.
Sargent CS. 1965. Manual of the trees of North America. 2nd ed.,
corrected and reprinted. New York: Dover. 934 p.
Wyman D. 1953. Seeds of woody plants. Arnoldia 13: 4160.
RanunculaceaeButtercup family
Clematis L.
clematis
John C. Zasada and Paul O. Rudolf
Dr. Zasada retired from the USDA Forest Services North Central Forest Research Station;
Dr. Rudolf (deceased) retired from the USDA Forest Services North Central Forest Experiment Station
Growth habit, occurrence, and use. The genus
Clematis includes more than 200 species of climbing vines,
and erect or ascending perennial herbs (sometimes woody)
widely that are distributed through the temperate regions,
chiefly in the Northern Hemisphere (Rehder 1940). Clematis
is subdivided into 3 sectionsFlammula (western and eastern virgins-bowers), Atragene (western blue clematis and C.
occidentalis (C.L. Hitchc.) Pringle), and Viorna (travelersjoy). The taxonomy and distribution of section Atragene are
described by Pringle (1971). Many horticultural varieties are
grown for ornamental purposes (Dirr 1990; Lloyd 1977;
Markham 1935). The 8 species included here (table 1) are
also useful for erosion control, ground cover, and wildlife
food (Bailey 1939; Dirr 1990; Fernald 1950; Rehder 1940;
Van Dersal 1938).
Species occupy different site types within their range. In
Wisconsin, for example, eastern virgins-bower was found in
13 community types but was most abundant in the wet alder
thicket community. Rock clematis is present in 2 communities and most abundant in northern dry mesic forests (Curtis
1959). Western species seem to be more common on drier
well-drained sites than species native east of the Mississippi
(table 1).
Geographic races. Two varieties of western virginsbowerC. ligusticifolia var. californica Wats. and var. brevifolia Nutt.are separated geographically within the
species range (Vines 1960). These and a variety of eastern
virgins-bowerC. virginiana var. missouriensis (Rydb.)
Palmer & Steyrm.may be geographic races. Wild plants
intermediate between Drummond clematis and western virgins-bower may be of hybrid origin (Vines 1960). Several
hybrids of Italian clematis are known (Rehder 1940).
Flowering and fruiting. There are both monoecious
and dioecious species. Eastern virgins-bower and western
virgins-bower (section Flammula) are dioecious, but their
female flowers have non-functional stamens. Species in the
sections Atragene and Viorna are monoecious (Fernald
1950). Flower size differs significantly among species, for
example, eastern virgins-bower flowers occur in clusters
(panicles) containing several flowers, and their sepals are
about 0.5 cm in diameter, whereas rock clematis flowers are
borne singly, and their sepals are about 4 cm. Fruits are
borne in heads of 1-seeded achenes with persistent feathery
styles. Achenes (figures 1 and 2) are produced annually
(Rudolf 1974) and are dispersed by wind in late summer or
fall. Some species have been shown to produce viable seeds
the first year after sowing (neoteny) (Beskaravainya 1977).
Common name(s)
Occurrence
western virgins-bower,
western clematis, travelers-joy
rope-vine
eastern virgins-bower,
Virginia virgins-bower, eastern clematis
travelers-joy, old-mans-beard
Italian clematis, vine-bower
C. flammula L.
C. pallasii J. F. Gmel.
C. ligusticifolia Nutt.
C. brevifolia Howell
C. pauciflora Nutt.
C. virginiana L.
C. catesbyana Pursh
C. vitalba L.
C. viticella L.
Clematis
415
Species
Location
Flowering
Fruit ripening
C. columbiana
C. drummondii
C. flammula
C. ligusticifolia
SW US
California
Texas
Colorado & Utah
California
Minnesota
NE US
France
NE US
MayJune
MarSept
AugOct
MarApr
MarSept
MayAug
MarApr
JulySept
JuneJuly
JulySept
JuneJuly
JuneAug
JulyAug
AugOct
AugOct
MayAug
AugNov
OctDec
MayJuly
JulySept
AugSept
JulySept
SeptOct
JuneAug
C. pauciflora
C. virginiana
C. vitalba
C. viticella
Sources: Fernald (1950), Loiseau (1945), McMinn (1951), Mirov and Kraebel (1939), Radford and others (1964), Rehder (1940), Rosendahl (1955), Rydberg (1922),Van
Dersal (1938),Vines (1960).
Table 3Clematis, clematis: size, year first cultivated, and flower color
Species
Flower color
C. columbiana
C. drummondii
C. flammula
C. ligusticifolia
C. pauciflora
C. virginiana
C. vitalba
C. viticella
2.8
3.14.6
0.912.3
3.76.2
10.2
4.6
1797
1509
1880
Before 1935
1726
Long cultivated
1597
Purple
White
White
White
White
Creamy white
White
Purplish
Sources: Fernald (1950), McMinn (1951), Rehder (1940), Rosendahl (1955),Vines (1960).
Place
collected
Minnesota
Europe
California
Utah
California
Baraga Co., Michigan
Europe
Europe
Range
/kg
663,000724,880*
402,220446,420
48,620-103-870
Average
/lb
300,000328,000*
182,000202,000
22,000-47,000
/kg
141,440
55,250
205,530
696,150*
187,850
424,320
707,200
59,670
/lb
64,000
25,000
93,000
315,000*
85,000
192,000
320,000
27,000
Sources: Mirov and Kraebel (1939), Rafn & Son (1928), Rudolf (1974).
* Styles removed.
Styles presumably removed.
Clematis
417
Test
Germination
duration (days) capacity (%)
40
200
60
76
1184
36
32
# Tests
1
8
1
1
Sources: Mirov and Kraebel (1939), Plummer and others (1968), Rudolf (1974),
References
Bailey LH. 1939. The standard cyclopedia of horticulture. New York:
Macmillan. 3639 p.
Beskaravainya MA. 1977. Neoteny in some clematis species [abstract].
Byulleten Gosudarstvennogo Nikitskogo Botanicheskogo Sada 32:
2629.
Blair FM. 1959. Raising from seed large-flowered clematis. Garden Journal
9(1): 11, 1415, 29.
Clark WS, Beattie DJ, Arteca RN. 1989. A growth inhibitor in Clematis viticella L. seeds. Journal of Plant Physiology 134: 492495.
Curtis JT. 1959. The vegetation of Wisconsin. Madison: University of
Wisconsin Press, 657 pp.
Czekalski M. 1987. Seed germination capacity and seedling morphology in
Clematis vitalba [abstract]. Prace Komisji Naukowe Rolniczych i Komisji
Naukowe Lesnych [published 1991] 63:1520.
Dirr MA. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation, and uses. Champaign, IL:
Stipes Publishing. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Evison RJ. 1977. Propagation of clematis. Proceedings of the International
Plant Propagators Society 27: 436440.
Fernald ML. 1950. Grays manual of botany. New York: American Book Co.
1632 p.
Fordham A. 1960. Propagation of woody plants by seed. Arnoldia 20:
3340.
Goi M, Senda Y, Ihara Y. 1975. Studies on flowering behavior in clematis: 1.
The flowering behavior of Clematis jackii cv Comtesse de Bouchaud
[abstract]. Kagawa Daigaku Nogakuba Gakujutsu Hokoku 26: 94100.
Hartmann HT, Kester DE, Davies FT Jr. 1990. Plant propagation: principles
and practice. 5th ed. Edgewood Cliffs, NJ: Prentice Hall. 647 p.
Heit CE. 1968. Thirty-five years testing of tree and shrub seed. Journal of
Forestry 66(8): 632634.
Lloyd C. 1977. Clematis. London: Collins. 208 p.
Loiseau J. 1945. Les arbres et la fort. Paris. 204 p.
Lush WM, Kaye PE, Grooves RH. 1984. Germination of Clematis microphylla
seeds following weathering and other treatments. Australian Journal of
Botany 32(2): 121129.
Markham E. 1935. Clematis. New York: Charles Scribner. 116 p.
418
Matlack GR. 1987. Diaspore size, shape, and fall behavior in wind-dispersed
plant species. American Journal of Botany 74(8): 11751160.
McMinn HE. 1951. An illustrated manual of California shrubs. Berkeley:
University of California Press. 663 p.
Mirov NT, Kraebel CJ. 1939. Collecting and handling seeds of wild plants.
For. Pub. 5. Washington, DC: USDA Civilian Conservation Corps. 42 p.
Plummer AP, Christensen DR, Monsen SB. 1968. Restoring big-game range
in Utah. Pub. 68-3. Salt Lake City: Utah Division of Fish and Game.
182 p.
Pringle JS. 1971. Taxonomy and distribution of Clematis, sect. Atragene
(Rannunculaceae), in North America. Brittonia 23: 361393.
Radford AE, Ahles HE, Bell CR. 1964. Guide to the vascular flora of the
Carolinas. Chapel Hill: University of North Carolina Book Exchange.
383 p.
Rafn J, & Son. [circa 1928]. Skovfrkontorets Franalyser Gennem 40 Aar,
18871927. Copenhagen: Udfrt paa Statsfrkontrollen i Kbenhavn.
5 p.
Rehder A. 1940. Manual of cultivated trees and shrubs hardy in North
America. New York: Macmillan. 996 p.
Rosendahl CO. 1955. Trees and shrubs of the upper Midwest. Minneapolis:
University of Minnesota Press. 411 p.
Rudolf PO. 1974. Clematis L., clematis. In: Schopmeyer CS, tech. Coord.
Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 331334.
Rydberg PA. 1922. Flora of the Rocky Mountains and adjacent plains,
Colorado, Utah, Wyoming, Montana, Saskatchewan, Alberta, and neighboring parts of Nebraska, South Dakota, and British Columbia. New
York: New York Botanical Garden. 1143 p.
Stribling IL. 1986. Clematis armandii propagation by seed. Plant Propagation
32(2): 1011.
Swingle CF. 1939. Seed propagation of trees, shrubs, and forbs for conservation planting. SCS-TP-27. Washington, DC: USDA Soil Conservation
Service. 198 p.
Van Dersal WR. 1938. Native woody plants of the United States: their erosion-control and wildlife values. Misc. Publ. 303. Washington, DC: USDA.
362 p.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
Clethra L.
sweet pepperbush, summersweet
Jason J. Griffin and Frank A. Blazich
Dr. Griffin is assistant professor at Kansas State Universitys Department of Horticulture, Forestry, and
Recreation Resources, Manhattan, Kansas; Dr. Blazich is alumni: distinquished graduate professor of plant
propagation and tissue culture at North Carolina State Universitys Department of Horticultural Science,
Raleigh, North Carolina
Growth habit, occurrence, and uses. The genus
Clethra L. comprises about 30 species native to eastern
Asia, eastern North America, and Madeira (Huxley 1992;
LHBH 1976). Of those, cinnamon-bark clethra and sweet
pepperbush are native to eastern North America, occurring
from southern Maine to Florida and west to Texas (LHBH
1976). Some taxonomists consider woolly summersweet to
be a separate species in this same range, but others consider
it to be a variety of sweet pepperbush (Huxley 1992; Kartesz
1994; Radford and others 1968). Japanese clethra, a native
of Japan, is commonly cultivated in North America (Koller
1974). Specific geographic regions of occurrence differ
among these species (table 1).
North American species of Clethra are deciduous shrubs
or small trees with heights ranging from 3 to 10 m in their
natural settings (Krssmann 1984). Species generally grow
as rounded, multi-stemmed plants that can be shaped easily
into attractive small trees (Bir 1992b).
Table 1Clethra, sweet pepperbush: nomenclature and occurrence of species cultivated in North America
Scientific name & synonym(s)
Occurrence
C. acuminata Michx.
cinnamon-bark clethra,
mountain sweetpepperbush
sweet pepperbush,
summersweet, coastal
sweetpepperbush
Japanese clethra,
Asiatic sweet pepperbush
woolly summersweet
C. alnifolia L.
C. alnifolia var. paniculata (Ait.) Rehd.
C. paniculata Ait. C. tomentosa Lam.
C. alnifolia var. pubescens Ait.
C. alnifolia var. tomentosa (Lam.) Michx.
C. barbinervis Sieb. & Zucc.
C. canescens Forbes & Hemsl.
C. kawadana Yanagita
C. barbinervis var. kawadana (Yanagita) Hara
C. repens Nakai
C. tomentosa Lam.
C. alnifolia var. pubescens Ait.
C. alnifolia var. tomentosa (Lam.) Michx.
Clethra
419
420
Figure 2Clethra, sweet pepperbush: seeds of C. acuminata, cinnamon-bark clethra (top left); C. alnifolia, sweet
pepperbush (top right); C. barbinervis, Japanese clethra
(bottom left); and C. tomentosa, woolly summersweet
(bottom right).
longitudinal
Nursery practice. When seedlings are grown in a typical azalea growing medium of 3 parts pine bark to 1 part
peat (vol/vol)amended with 4.2 kg/m3 (7.0 lb/yd3)
dolomitic limestone and fertilized following recommendations for azaleasseedlings grow well, filling a 3.8-liter
(1-gal) pot by the end of a growing season (Bir 1992b).
Although naturally occurring as an understory species,
seedlings of cinnamon-bark clethra show no symptoms of
stress when grown in full sun and are visually no different
than seedlings maintained under 50% shade (Bir 1992b).
The species is well adapted to dry soils once established.
However, if seedlings are exposed to drought conditions
before a sufficient root system has developed, high mortality
can be expected (Bir 1992b).
Asexual propagation of species of summer-sweet is very
easy and is widely used for propagation of particular cultivars. Species listed in table 1 are propagated readily by stem
cuttings taken during the summer, as well as by root cuttings
taken during December and January (Dirr and Heuser 1987).
References
Bir RE. 1992a. Growing and propagating showy native woody plants.
Chapel Hill: University of North Carolina Press. 192 p.
Bir RE. 1992b. Native spice. American Nurseryman 175(3): 5152, 5459.
Bir RE. 1993. Clethra. Fine Gardening 43(30): 5254.
Dirr MA. 1994. Confessions of a Clethra-phile. Nursery Manager 10(11):
1421.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Everett TH, ed. 1981. The New York Botanical Garden illustrated
encyclopedia of horticulture.Volume 3. New York: Garland Publishing:
7051058.
Huxley A, ed. 1992. The new Royal Horticultural Society dictionary of gardening.Volume 1. New York: Stockton Press. 815 p.
Jordan RA, Hartman JM. 1995. Safe sites and the regeneration of Clethra
alnifolia L. (Clethraceae) in wetland forests of central New Jersey.
American Midland Naturalist 133: 112123.
Kartesz JT. 1994. A synonymized checklist of the vascular flora of the
United States, Canada, and Greenland. 2nd ed.Volume 1. Portland, OR:
Timber Press. 622 p.
Koller G. 1974. Shrub profiles: Clethra barbinervis. Morris Arboretum
Bulletin 25(1): 23.
Krssmann G. 1984. Manual of cultivated broad-leaved trees & shrubs.
Volume 1. Portland, OR:Timber Press. 624 p.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise
dictionary of plants cultivated in the United States and Canada. 3rd ed.
New York: Macmillan Publishing Co. 1290 p.
Ohwi J. 1984. Flora of Japan. Washington, DC: Smithsonian Institution.
1067 p.
Radford AE, Ahles HE, Bell CR. 1968. Manual of the vascular flora of the
Carolinas. Chapel Hill: University of North Carolina Press. 1183 p.
Sleumer H. 1967. Monographia Clethracearum. Botanische Jahrbcher fr
Systematik, Pflanzengeschichte und Pflanzengeographie 87(1): 36116.
Small JK. 1933. Manual of the southeastern flora. Chapel Hill: University of
North Carolina Press. 1554 p
Clethra
421
RosaceaeRose family
422
fruits with
However, the vigor of older seeds (10+ years) in field plantings has not been determined.
Germination. Fresh blackbrush seedlots are 68 to
95% dormant and remain dormant in laboratory storage for
the first year after collection (table 1). Seed dormancy
increases with increasing elevation of the seed source. Fiveyear-old seeds are essentially nondormant and will readily
germinate at cool temperatures (15 C) (Pendleton and others 1995). Stratification of fresh seeds for 4 to 6 weeks at
1 C will produce rapid maximum germination of all collections when seeds are removed from chill and placed at temperatures between 5 and 25 C. Under field conditions,
seeds are nondormant by October, and germination can
occur at this time given proper moisture conditions and cool
soil temperatures. Field germination typically occurs during
the winter and early spring (figure 3) (Meyer and Pendleton
2002).
Nursery and field practice. Container stock can
readily be produced from seeds. However, both stratified
and unstratified seeds will germinate and emerge over a long
period of time (up to 1 year). The most efficient method to
synchronize germination and produce uniform-aged plants is
to plant germinated seeds. Fresh seeds should be stratified
between moist blotter paper for 6 weeks at 1 C. When
seedlots are removed from 1 C and kept at room temperature, more than 75% will germinate in 24 to 48 hours. Seed
collections 3 years old or older, stratified for 3 to 4 weeks,
produce similar results. Germinated seeds, with radicals 2 to
10 mm (0.1 to 0.4 in) long, should be planted about 2 cm
(0.8 in) deep in a well-draining soil mix. However, using a
soil medium that retains moisture often leads to problems
with damping-off diseases. Under experimental greenhouse
conditions, blackbrush responds positively to inoculation
with arbuscular mycorrhizal fungi (Pendleton and Warren
Seed age
Fresh
Stratified
Nonstratified
1 year
Stratified
Nonstratified
5 years
Nonstratified
Range
Percent germination
Mean
532
8498
17.6
91.8
10
10
013
8595
6.5
92.0
10
10
8296
90
Collections
Coleogyne
423
424
References
Beatley JC. 1974. Phenological events and their environmental triggers in
Mojave desert ecosystems. Ecology 55: 856863.
Bowns JE, West NE. 1976. Blackbrush (Coleogyne ramosissima Torr.) on
southwestern Utah rangelands. Res. Rep. 27. Logan: Utah Agricultural
Experiment Station.
Graham T. 1991. Unpublished data. Moab, UT: USDI Geological Survey,
Forest and Range Ecosystem Science Center.
Holden MK. 1994. Mojave plant information. Newsletter of the Mojave
Native Growers Association (Summer): 25.
Hughes H, Weglinski E. 1991. Blackbrush, Coleogyne ramosissima, propagation and revegetation of disturbed sites. In: Plumb GE, ed. 15th annual
report, University of Wyoming, National Park Service Research Center:
6774.
Meyer SE, Pendleton BK. 2002. Regeneration biology of blackbrush
(Coleogyne ramosissima: Rosaceae): 2. Field germination and establishment experiments [in review].
Monsen SB. 2004. Coleogyne ramosissima. In: Monsen SB, Stevens R, eds.
Restoring western ranges and wildlands. Gen.Tech. Rep. RM-136. Shaw
NL. Fort Collins, CO: USDA Forest Service, Rocky Mountain Research
Station.
Nord EC. 1962. Bitterbrush seed harvesting: when, where, and how.
Journal of Range Management 16: 258260.
Pendleton BK, Meyer SE. 2002. Unpublished data. Provo, UT: USDA Forest
Service, Rocky Mountain Research Station.
Pendleton BK, Meyer SE. 2002. Regeneration biology of blackbrush
(Coleogyne ramosissima: Rosaceae): 1. Habitat-correlated variation in
seed germination responses [in review].
Pendleton BK, Meyer SE, Pendleton RL. 1995. Blackbrush biology: insights
after three years of a long-term study. In: Roundy BA, McArthur ED,
Haley JS, Mann DK, comps. Proceedings, Wildland Shrub and Arid Land
Restoration Symposium. Gen.Tech. Rep. INT-315. Ogden, UT: USDA
Forest Service, Intermountain Forest and Range Experiment Station:
223227.
Pendleton BK, Pendleton RL. 1998. Pollination biology of Coleogyne ramosissima (Rosaceae). Southwestern Naturalist 43: 376380.
Pendleton RL, Warren SD. 1996. The effects of cryptobiotic soil crusts and
VA mycorrhizal inoculation on growth and nutrient content of five
rangeland plant species. In: West NE, ed. Rangelands in a Sustainable
Biosphere: Proceedings, 5th International Rangeland Congress. Denver:
Society for Range Management: 436437.
USDoE [United States Department of Energy]. 1994. Unpublished data.
Las Vegas, NV.
Wallace A, Romney EM. 1972. Radioecology and ecophysiology of desert
plants at the Nevada test site. Oak Ridge,TN: US Atomic Energy
Commission,Technical Information Service.
Wallace A, Romney EM, Ashcroft RT. 1970. Soil temperature effects on
growth of seedlings of some shrub species which grow in the transitional area between the Mojave and Great Basin deserts. BioScience 20:
11581159.
Welsh SK, Atwood ND, Goodrich S, Higgins LC, eds. 1993. A Utah flora.
2nd ed. Great Basin Naturalist Memoirs 9. Provo, UT: Brigham Young
University. 986 p.
Coleogyne
425
FabaceaePea family
Colutea L.
bladder-senna
Paula M. Pijut
Dr. Pijut is a research plant physiologist at the USDA Forest Service, North Central Research Stations
Hardwood Tree Improvement and Regeneration Center,West Lafayette, Indiana
Table 1Colutea, bladder-senna: morphological characteristics, height at maturity, and date first cultivated
Scientific name
Leaflets
C. arborescens
C. orientalis
C. x media
913
711
1113
426
Flowers/
raceme
Height at
maturity (m)
Year first
cultivated
68
25
Varies
1.84.5
2
1.83.0
1570
1710
1809
References
Allen DH. 1995. Personal communication. Sandwich, MA: F.W. Schumacher
Co.
Allue Andrade JL. 1983a. Aspectos ecologicos, fenologicos y biometricos de
Colutea arborescens L. [in Spanish, with English summary: Ecological, phenological, and biometric aspects of Colutea arborescens L.]. Anales del
Instituto Nacional de Investigaciones Agrarias Seria Forestal 7: 111127.
Allue Andrade JL. 1983b. Morfoligia, clases, atributos, dificultades y
tratamientos en la produccion y germinacion de las semillas de Colutea
arborescens L. [in Spanish, English summary: Morphology, types, attributes,
difficulties and treatments in production and germination of seeds of
Colutea arborescens L.]. Anales del Instituto Nacional de Investigaciones
Agrarias Seria Forestal 7: 129154
Browicz K. 1963. The genus Colutea L.: a monograph. Monographiae
Botanicae 14: 1136.
Browicz K. 1967. A supplement to the mongraph of the genus Colutea L.
Arboretum Kornickie 12: 3343.
Dirr MA. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation, and uses. Champaign, IL:
Stipes Publishing Company. 1007 p.
Dirr MA, Heuser CW. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Gonzalez-Benito ME, Caze-Filho J, Perez C. 1994. Cryopreservation of
seeds of several legume species. Plant Varieties and Seeds 7: 2327.
Grosvenor PW, Gray DO. 1998. Coluteol and colutequinone B, more
antifungal isolavonoids from Colutea arborescens. Journal of Natural
Products 61(1): 99101.
Hillier Nurseries (Winchester) Ltd. 1991. The Hillier manual of trees and
shrubs. Melksham, Wiltshire, UK: Redwood Press. 704 p.
Iriondo JM, Perez C, Perez-Garcia F. 1992. Effect of seed storage in liquid
nitrogen on germination of several crop and wild species. Seed Science
and Technology 20: 165171.
Krssmann G. 1984. Manual of cultivated broad-leaved trees and shrubs.
Volume 1, AD. Beaverton, OR:Timber Press. 448 p.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan. 1290 p.
Rudolf PO. 1974. Colutea arborescens L., bladder-senna. In: Schopmeyer CS,
tech. coord. Seeds of woody plants in the United States. Agric. Handbk.
450. Washington, DC: USDA Forest Service: 335.
Colutea
427
CornaceaeDogwood family
Cornus L.
dogwood
Kenneth A. Brinkman and Victor Vankus
Dr. Brinkman retired from the USDA Forest Services North Central Experiment Station;
Mr. Vankus is a botanist at the USDA Forest Services, National Seed Laboratory, Dry Branch, Georgia
Figure 1Cornus, dogwood: cleaned seeds of C. alternifolia, alternate-leaf dogwood (top left); C. amomum, silky
dogwood (top center); C. sericea ssp. orientalis, California
dogwood (top right); C. drummondii, roughleaf dogwood
(middle left); C. florida, flowering dogwood (middle
center); C. nuttallii, Pacific dogwood (middle right); and
C. racemosa, gray dogwood (bottom).
Figure 2Cornus sericea, red-osier dogwood: longitudinal section through an embryo of a stone (left); transverse
section of a stone containing 2 embryos (right top) and
transverse section of a stone containing 1 embryo (right
bottom).
Common name(s)
Occurrence
C. alba L.
C. tatarica Mill.
C. alternifolia L. f.
Swida alternifolia (L.f.) Small
C. amomum P. Mill.
C. canadensis L.
Chamaepericlymenum
canadense(L.) Aschers & Graebn.
Cornella canadensis (L.) Rydb.
C. controversa Hems.
C. drummondii C. A. Mey.
C. priceae Small
C. florida L.
Cynoxylon floridum (L) Raf. ex. B.D. Jackson
C. kousa Hance
Tatarian dogwood
alternate-leaf dogwood,
blue dogwood, pagoda dogwood
silky dogwood, kinnikinnik, red-willow
bunchberry, bunchberry
dogwood, dwarf cornel
Newfoundland to SE Manitoba, S to
Missouri & E Arkansas, E to Georgia
Maine to Indiana, S to Georgia & Florida
S Greenland to Alaska, S to
Maryland,W to South Dakota,
New Mexico, & California
giant dogwood
roughleaf dogwood
C. macrophylla Wall.
C. mas L.
C. nuttallii Audubon
ex Torr. & Gray
C. officinalis Siebold & Zucc.
C. racemosa Lam.
C. foemina ssp. racemosa (Lam.)
C. circinata LHerit. J.S.Wilson
C. paniculata LHerit.
C. rugosa Lam.
C. circinata LHerit.
C. sanguinea L.
C. sanguinea var. viridissima Dieck
Swida sanguinea (L.) Opiz
C. sericea L.
C. stolonifera Michx.
C. baileyi Coult. & Evans
Suida stolonifera (Michx.) Rydb.
C. sericea ssp. occidentalis
(Torr.& Gray) Fosberg
roundleaf dogwood,
roundleaved dogwood, roundleaf cornel
bloodtwig dogwood, common
dogwood, dogberry, pegwood
Newfoundland to Alaska, S to
California, New Mexico, & Nebraska,
in NE US from Wisconsin to New York
429
Flowering
Fruit ripening
Seed dispersal
C. alba
C. alternifolia
C. amomum
C. canadensis
C. controversa
C. drummondii
C. florida
C. kousa
C. macrophylla
C. mas
C. nuttallii
C. officinalis
C. racemosa
C. rugosa
C. sanguinea
C. sericea
C. sericea ssp. occidentalis
MayJune
MayJuly
MayJuly
MayJuly
MayJune
MayJune
Mar & Apr (S US)May (N US)
MayJune
JulyAug
FebMarch
AprilMay
FebMar
late MayJuly
MayJuly
MayJune
MayJuly, JuneAug (N US)
AprAug
AugSept
JulySept
AugSept
Aug
AugSept
AugOct
Sept (N US)Oct (S US)
AugOct
AugSept
SeptOct
Sept
JulyOct
AugSept
AugSept
JulyOct
JulyNov
JulySept
Sept
AugOct
Oct
Augwinter
SeptNov
SeptOct
SeptOct
Octwinter
Sources: Asakawa, (1969), Billington (1943), Brinkman (1974), Dirr (1990), Fernald (1950), Forbes (1956), Holweg (1964), Gordon and Rowe (1982), Lakela (1965),
McMinn (1951), Ohwi (1965), Rehder (1940), Rosendahl (195), Rydberg (1932), Steyermark (1963),Van Dersal (1938),Vimmerstedt (1965),Weaver (1976),Wyman (1947).
Table 3Cornus, dogwood: height, seed-bearing age, seedcrop frequency, and fruit ripeness criteria
Species
Height at
maturity
(m)
C. alba
3
C. alternifolia
58
C. amomum
3
C. canadensis
0.3
C. controversa
918
C. drummondii
814
C. florida
612
C. kousa
8
C. macrophylla
811
C. mas
8
C. nuttallii
624
C. officinalis
69
C. racemosa
4
C. rugosa
3
C. sanguinea
25
C. sericea
36
C. sericea ssp. occidentalis
5
Year first
cultivated
1741
1760
1658
1880
1836
1731
1875
1827
Ancient
1835
1877
1758
1784
1656
Years between
large
seedcrops
45
10
12
2
Sources: Dirr (1990), Fernald (1950), Gordon and Rowe (1982), McMinn (1951), Rehder (1940),Weaver (1976).
Stones/fruit wt
kg/100 kg
lbs/100 lb
C. alba
C. alternifolia
C. amomum
C. canadensis
C. drummondii
C. florida
C. kousa
C. mas
C. nuttallii*
C. racemosa
C. rugosa
C. sanguinea
C. sericea
C. sericea ssp. occidentalis
1518
1624
1741
13
11
1622
1318
1720
1827
1946
15
12
1825
1520
Range
/kg
27,90040,900
13,00020,500
22,40030,800
129,800169,400
18,90046,200
7,30013,600
14,30018,300
3,5007,500
8,80013,400
22,40033,700
16,10026,000
30,40058,700
/lb
12,70018,600
5,9009,300
10,20014,000
59,00077,000
8,60021,000
3,3006,200
6,5008,300
1,6003,400
4,0006,100
10,20015,300
7,30011,800
13,80026,700
Average
/kg
/lb
33,000
17,600
26,800
147,400
34,500
9,900
21,300
5,000
10,300
28,600
41,800
20,200
40,700
73,500
Samples
15,000
8,000
12,200
67,000
15,700
4,500
9,700
2,300
4,700
13,000
19,000
9,200
18,500
33,400
33
6
6
2
5
11
3
22
4
11
1
70
9
1
Sources: Asakawa (1969), Brinkman (1974), Edminster (1947), Forbes (1956), Gordon and Rowe (1982), Gorshenin (1941), Heit (1969), Mirov and Kraebel (1939),
Mugford (1969), NBV (1946), Stevenson (1969), Swingle (1930).
* 0.036 cubic meters (1 bu) of fruit clusters weighed 15 kg (33 lb) and yielded 2 kg (4 lb) of stones (Brinkman 1974).
Cornus
431
Species
C. alba
C. alternifolia
C. amomum*
C. canadensis
C. controversa
C. drummondii
C. florida
C. kousa
C. macrophylla
C. mas
C. nuttallii
C. officinalis
C. racemosa
C. rugosa
C. sanguinea
C. sericea //
Sand
Sand
Sand, peat, or vermiculite
Soil or vermiculite
Peat
Sand
Soil
Sand
Sand
3020
25
2127
2030
1522
2030
Days
Cold period
Temp (C)
60
3060
6090
1
3060
90150
120
120150
60
60
5
5
35
5
5
15
113
3
5
Outdoors
25
5
Duration (days)
90120
60
2128
90120
6090
120150
6090
30
3060
120
40120
90
30120
90
90
60, 120
Overwinter
6090
6090
60-90
Sources: Billington (1943), Brinkman (1974), Dirr and Heuser (1987), Emery (1988), Guan and others (1989), Goodwin (1948), Gordon and Rowe (1982), Heit (1967,
1968b), Jack (1969), Nichols (1934), Ohwi (1965), Pammel and King (1921), Peterson (1953), Soljanik (1961), Swingle (1939),
* Seeds were soaked for 3 hours in water at room temperature before stratification (Heit 1968b).
Seeds were soaked for 1 hour in sulfuric acid before stratification (Dirr and Heuser 1987).
Seeds were mechanically scarified before stratification (Brinkman 1974).
Seeds were soaked for 4 hours in sulfuric acid before stratification (Emery 1988).
// Seeds were soaked for 1 hour in sulfuric acid before stratification (Brinkman 1974).
Species
C. alba
C. alternifolia
C. amomum
C. canadensis
C. drummondii
C. florida
C. kousa
C. macrophylla
C. mas
C. nuttallii
C. racemosa
C. rugosa
C. sericea
8
824
8
8
824
8
8
60
1428
6090
50
60
30
47
60
60+
6090
Germination
rate
Amt
(%)
Days
8
86
6
14
1445
57
2230
8
35
50
11
26
34
1520
16
14
15
1318
Germination %
Average
(%)
Samples
Purity
(%)
10
70
16
25
35
85
57
81
20
46
57
2
6
5
3
7
2
6
2
8
4
18
63
91
90
89
97
95
100
83
95
99
Sources: Adams (1927), Asakawa (1969), Brinkman (1974), Heit (1968a&b, 1969), McKeever (1938), Nichols (1934), Peterson (1953), Soljanik (1961), Swingle (1939),Titus
(1940).
*Temperatures were 30 C for 8 hours and 20 C for 16 hours each day. Sand was the medium used on all listed species. Additional tests were made on wet paper in
germinators with seeds of C. amomum, C. kousa, and C. nuttallii (Brinkman 1974; Heit 1969).
One test.
432
References
Adams J. 1927. The germination of the seeds of some plants with fleshy
fruits. American Journal of Botany 14: 415428.
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing
seeds. Journal of Seed Technology 16(3): 1113.
Asakawa S. 1969. Correspondence, June 17 and November 27, 1969.
Tokyo: Japanese Ministry of Agriculture and Forestry.
Billington C. 1943. Shrubs of Michigan. Cranbrook Institute of Science
Bulletin 20: 1249.
Brinkman KA. 1974. Cornus. L. dogwood. In: Schopmeyer CS, tech. coord.
Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 503504.
Brock S. 1997. Personal communication. Macon: Georgia Forestry
Commission.
Dirr MA. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture propagation and uses. Champaign, IL:
Stipes Publishing Co.
Dirr MA, Heuser CW. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Edminster FC. 1947. The ruffed grouse: its life story, ecology and
management. New York: Macmillan. 385 p.
Edminster FC. 1950. Use of shrubs in developing farm wildlife habitat.
North American Wildlife Conference Transactions 15: 519550.
Edminster FC, May RM. 1951. Shrub plantings for soil conservation and
wildlife cover in the Northeast. Circ. 887. Washington, DC: USDA. 68 p.
Emery D. 1988. Seed propagation of native California plants. Santa Barbara,
CA: Santa Barbara Botanic Garden. 115 p.
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1632 p.
Flemion F. 1948. Reliability of the excised embryo method as a rapid test
for determining the germinative capacity of dormant seeds.
Contributions of the Boyce Thompson Institute 15(4): 229242.
Forbes RD, ed. 1956. Forestry handbook, Sect. 9. New York: Ronald Press:
2831.
Furuta T. 1960. Alabama study evaluates production methods, polyethylene
packing. Florist Exchange 135(1): 2224, 2627, 2930, 32.
Goodwin RH. 1948. How to grow dogwood from seed. Plants and
Gardens 4(4): 236238.
Gordon AG, Rowe DCF. 1982. Seed manual for trees and shrubs. For.
Comm. Bull. 59. London: United Kingdom Forestry Commission. 132 p.
Gorshenin NM. 1941. Agrolesomelioratsiya [in Russian: Agro-forest
melioration]. 392 p.
Guan KL, Fan X, Zhen G. 1989. [in Chinese: Causes of seed dormancy of
Cornus officinalis and conditions for germination]. Plant Physiology
Communications 5: 2427 [Seed Abstracts 1992; 15(12): 3972].
Heit CE. 1955. The excised embryo method for testing germination quality
of dormant seed. Proceedings of the Association of Official Seed
Analysts 45: 108117.
Heit CE. 1967. Propagation from seed: 11. Storage of deciduous tree and
shrub seeds. American Nurseryman 126(10): 1213, 8694.
Heit CE. 1968a. Propagation from seed: 15. Fall planting of shrub seeds for
successful seedling production. American Nurseryman 128(4): 810,
7080.
Heit CE. 1968b. Thirty-five years testing of tree and shrub seed. Journal of
Forestry 66: 632634.
Heit CE. 1969. Correspondence, Feb. 14. Geneva, NY: Cornell University,
New York State Agricultural Experiment Station Department of Seed
Investigations.
Holweg AW. 1964. Some shrubs and vines for wildlife food and cover. New
York Conservation 19(2): 2227.
ISTA [International Seed Testing Association]. 1993. Rules for testing seeds:
rules 1993. Seed Science and Technology 21 (Suppl.): 1259.
Jack RA. 1969. Personal communication. Silverton, OR: Silver Falls Nursery.
Lakela O. 1965. A flora of northeastern Minnesota. Minneapolis: University
of Minnesota Press. 541 p.
Litvinenko SN. 1959. The Ukrainian gibberellin, an effective growth stimulant. Doklady Akademii Nauk SSSR Seriya Biologiya 126: 13681370.
McKeever DG. 1938. The effect of various methods of treatment on germination of seeds of some plants valuable for game and erosion purposes
[unpublished MS thesis]. Moscow, ID: University of Idaho.132 p.
McMinn HE. 1951. An illustrative manual of California shrubs. Berkeley:
University of Califfornia Press. 663 p.
Mirov NT, Kraebel CJ. 1939. Collecting and handling seeds of wild plants.
For. Pub. 5. Washington, DC: USDA Civilian Conservation Corps. 42 p.
Mugford D. 1969. Personal communication. Licking, MO: Missouri
Conservation Department, George White Nursery.
Murphy K. 1997. Personal communication. Planis, MT: Lawyer Nursery.
NBV [Nederlandsche Boschbouw Vereeniging]. 1946. Boomzaden:
Handleiding inzake het oogsten, behandelen, bewaren en uitzaaien van
boomzaden. Wageningen,The Netherlands: Ponsen and Looijen. 171 p.
Nichols GE. 1934. The influence of exposure to winter temperatures upon
seed germination in various native American plants. Ecology 15:
364373.
Ohwi J. 1965. Flora of Japan. Washington, DC: Smithsonian Institution.
1067 p.
Pammel LH, King CM. 1921. Studies in the germination of some woody
plants. Proceedings of the Iowa Academy of Science 28: 273282.
Peterson RA. 1953. Comparative effect of seed treatments upon seedling
emergence in seven browse species. Ecology 34: 778785.
Rehder A. 1940. Manual of cultivated trees and shrubs hardy in North
America. New York: Macmillan. 996 p.
Rosendahl CO. 1955. Trees and shrubs of the Upper Midwest. Minneapolis:
University of Minnesota Press. 411 p.
Rydberg PA. 1932. Flora of the prairies and plains of Central North
America. New York: New York Botanical Garden. 969 p.
Shumilina ZK. 1949. Podgotovka posevu semyan drevesnykh i kustarnikovykh porod.Vses. Nauchnoissled. Inst. Agrolesomelior.,
Goslesbumizdat, Moskva-Leningrad [Preparation of tree and shrub seed
for sowing.Transl.TT 67-51300. 1967. Springfield,VA: USDC CFSTI. 36
p.].
Soljanik I. 1961. Proizvodnja sadnica od nedozrelog sumskog semena [in
Croation]. 11 p. Savez. Inz.Teh. Sum. Drvne Ind. Sad. Beograd [English
transl. for USDA, 1968: Producing seedlings from unripe forest seed.].
Stevenson H. 1969. Personal communication. Elsberry, MO: Forrest Keeling
Nursery.
Steyermark JA. 1963. Flora of Missouri. Ames: Iowa State University Press.
1728 p.
Sus NI. 1925. Pitomnik [in Russian:The forest nursery]. 227 p.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP27. Washington, DC: USDA Soil
Conservation Service. 198 p.
Titus GR. 1940. So-called 2-year seeds germinated first year. American
Nurseryman 72(11): 22.
Tylkowski T. 1992. Thermal conditions for the after-ripening and germination
of Cornelian cherry (Cornus mas L.) seeds. Arboretum Kornickie 36:
165172.
Van Dersal WR. 1938. Native woody plants of the United States: their erosion-control and wildlife values. Misc. Pub. 303. Washington, DC: USDA.
362 p.
Vimmerstedt JP. 1965. Flowering dogwood (Cornus florida L.). In: Silvics of
forest trees of the United States. Agric. Handbk. 271. Washington, DC:
USDA Forest Service: 162166.
Weaver RE. 1976. The cornelian cherries. Arnoldia 36(2): 5056.
Wyman D. 1947. Seed collecting dates of woody plants. Arnoldia 7(9):
5356.
Cornus
433
BetulaceaeBirch family
Corylus L.
hazel
Jill Barbour and Kenneth A. Brinkman
Ms. Barbour is a germination specialist at USDA Forest Services National Seed Laboratory, Dry Branch,
Georgia; Dr. Brinkman retired from the USDA Forest Services North Central Forest Experiment Station
Common name(s)
Occurrence
C. americana Walt.
C. avellana L.
C. cornuta Marsh.
C. rostrata Ait.
C. cornuta var. california
Marsh. (A.DC.) Sharp
Source: Brinkman (1974).
434
California hazel,
California filbert
Location
Flowering
Fruit ripening
C. americana
C. avellana
C. cornuta
var. california
Europe
Tennessee
California
MarMay
FebApr
JanFeb
JanFeb
JulySept
SeptOct
AugSept
SeptOct
Sources: Fernald (1950), Loiseau (1945), Munz and Keck (1959), NBV (1946), Rosendahl (1955), Sus (1925),Van Dersal (1938),Vines (1960),Wappes (1932), Zarger
Place of
Seed wt/fruit wt
collection
kg/45 kg lb/100 lb
Europe
California
1114
27
2530
60
Range
/kg
4341,623
3531,180
9371,490
882922
Average
/lb
197736
160535
425676
400418
/kg
/lb
1,083
803
549
410
491
364
249
186
Samples
11
244
3
Sources: Brinkman (1974), Gorshenin (1941), NBV (1946), Rafn (1928), Swingle (1939), Vines (1960), Zarger (1968).
Because some dormancy is apparently induced by drying the nuts, seeds of hazel species were once thought to be
recalcitrant and intolerant of any drying (Hong and Ellis
1996). Recommendations usually were to keep the hazelnuts
moist after collection and store them moist over winter
(stratification) before planting in the spring (Heit 1967;
NBV 1946). Seeds of hazel species are now considered as
orthodox in storage behavior, even though moist storage will
prevent deep embryo dormancy for at least several months.
Seeds of this genus will also remain viable for a year in
unsealed containers at room temperature. Most of the viability of American hazelnut and some of beaked hazelnuts
(Brinkman 1974) will be retained if seeds are stored in
sealed containers at 5 C. There are no long-term storage
data for hazelnuts.
Pregermination treatments. Newly harvested hazelnuts are not dormant, but inhibitors present in the testa and
pericarp are carried to the cotyledons and subsequently
through the cotyledonary petioles into the embryonic axis
(Bradbeer 1978; Jarvis 1975). Numerous studies have been
Corylus
435
436
Species
C. americana
C. avellana
C. cornuta
var. californica
Germinative
energy
Amt
(%)
Days
Germinative capacity
Average
Purity
(%)
Samples
(%)
30
30
20
20
60
60
10
30
13
69
2
13
96
95
30
30
20
20
60
90
26
1
20
1
1
99
62
References
Arias I, Williams PM, Bradbeer JW. 1976. Studies in seed dormancy: 9.The
role of gibberellin biosynthesis and the release of bound gibberellin in
the post-chilling accumulation of gibberellin in seeds of Corylus avellana
L. Planta 131: 135139.
Bradbeer JW, Pinfield NJ. 1966. Studies in seed dormancy: 3.The effects of
gibberellin on dormant seeds of Corylus avellana L. New Phytologist 66:
515523.
Bradbeer JW, Arias IE, Nirmala HS. 1978. The role of chilling in the breaking
of seed dormancy in Corylus avellana L. Pesticide Science 9: 184186.
Brinkman KA. 1974. Corylus L., hazel, filbert. In: Schopmeyer SC, tech. coord.
Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 343345.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1632 p.
Gorshenin NM. 1941. Agrolesomelioratsiya [in Russian: Agro-forest
melioration]. 392 p.
Heit CE. 1967. Propagation from seed: 11. Storage of deciduous tree and
shrub seeds. American Nurseryman 126 (10): 1213, 8694.
Heit CE. 1968a. Propagation from seed: 15. Fall planting of shrub seeds for
successful seedling production. American Nurseryman 128(4): 810,
7080.
Heit CE. 1968b. Thirty-five years testing of tree and shrub seed. Journal of
Forestry 66: 632634.
Hong TD, Ellis RH. 1996. A protocol to determine seed storage behavior.
Tech. Bull. 1. Rome: International Plant Genetic Resources Institute. 62 p.
Hora B. 1981. The Oxford encyclopedia of trees of the world. Oxford, UK:
Oxford University Press. 288 p.
Horvath D. 1999. Personal communication.Topeka, IL: Mason State
Nursery.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing. Seed Science and Technology 21 (suppl.): 1288.
Jarvis BC. 1975. The role of seed parts in the induction of dormancy of
hazel (Corylus avellana L.). New Phytologist 75: 491494.
Jarvis BC, Wilson DA. 1978. Factors influencing growth of embryonic axes
from dormant seeds of hazel (Corylus avellana L.). Planta 138: 189191.
Jeavons RA, Jarvis BC. 1984. The breaking of dormancy in hazel seed by
pretreatment with ethanol and mercuric chloride. New Phytologist 96:
551554.
Li L, Ross JD. 1990. Starch synthesis during dormancy breakage in oilseed
of Corylus avellana. Annals of Botany 66: 507512.
Loiseau J. 1945. Les arbres et la foret. 204 p. Paris.
Maloney J. 1999. Personal communication. Licking, MO: George O. White
Nursery.
Munz PA,Keck DD. 1959. A California flora. Berkeley: University of
California Press. 1681 p.
NBV [Nederlandsche Boschbouw Vereeniging]. 1946. Boomzaden: handleiding inzake het oogsten, behandelen, bewaren en uitzaaien van
boomzaden [in Dutch:Tree seed: handbook on hte collection, extraction, storage, and sowing of tree seed]. Wageningen,The Netherlands:
Ponsen and Looijen. 171 p.
Rafn J & Son. [circa] 1928. Skovfrkontorets Franalyser gennem 40 Aar,
18871927. Udfrt paa Statsfrkontrollen i Kbenhavn. Copenhagen.
5 p.
Rosendahl CO. 1955. Trees and shrubs of the Upper Midwest.
Minneapolis: University of Minnesota Press. 411 p.
Shumilina ZK. 1949. Stratifikatsiya semyan drevesnykh i kustarnikovykh
porod.Vsesoyuznoe Nauchno-Issledovatelskii Institut Agrolesomelior.
Goslestekhizdat, Moskva. [Stratification of seeds of trees and shrubs.
Transl. OTS-60-51012, 1961. Springfield,VA: USDC CFSTI, NTIS, 64 p.].
Sus NI. 1925. Pitomnik [in Russian:The forest nursery]. 27 p.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting.TP-27. Washington, DC: USDA Soil Conservation
Service. 198 p.
Van Dersal WR. 1938. Native woody plants of the United States: their
erosion-control and wildlife values. Misc. Pub. 303. Washington, DC:
USDA. 362 p.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
Wappes L. 1932. Wald und Holz ein Nachschlagebuch fr die Praxis der
Forstwirte, Holzhndler und Holzindustriellen.Volume 1. Berlin: J.
Neumann. 872 p.
Zarger TG. 1968. Personal communication.Tennessee Valley Authority,
Division of Forests & Development.
Corylus
437
AnacardiaceaeSumac family
Cotinus P. Mill.
smoketree or smokebush
Paula M. Pijut
Dr. Pijut is a research plant physiologist at the USDA Forest Services North Central Research Station,
Hardwood Tree Improvement and Regeneration Center,West Lafayette, Indiana
var accumulates anthocyanin pigments in response to ultraviolet light of wavelengths between 300 and 400 nm and
low temperatures (Oren-Shamir and Levi-Nissim 1997).
American smoketree is a large, upright shrub or small,
round-headed tree with bluish to dark green leaves that turn
a brilliant yellow, orange, red, and reddish purple color in
the fall (Dirr 1990). The bark of the American smoketree is
a beautiful gray to gray-brown, and scaly mature trunks (that
is, with a fishlike scale effect), providing pattern and detail
in the winter landscape (Dirr 1990; Koller and Shadow
1991). For a review of Cotinus and discussion of selected
cultivars, see Tripp (1994).
Flowering and fruiting. The small, usually infertile,
yellowish flowers, which bloom in June to July (April to
May for American smoketree), are borne in large, terminal
panicles (Krssmann 1984). The pedicels and peduncles
lengthen after flowering and are clad with fine hairs, creating the smokelike effect that gives the plant its common
name (LHBH 1976). The plumelike inflorescences often
persist through September (Dirr 1990). The fruit (figures 1
and 2) is a dry, reticulate drupe about 3 to 6 mm in length,
light red-brown in color (ripening to near black), containing
a thick, bony stone (Rudolf 1974). Seedcrops are produced
annually but are often poor. The kidney-shaped drupe ripens
in the fall, which is usually August to October for common
Table 1Cotinus, smoketree: nomenclature, occurrence, growth habit, height at maturity, and date first cultivated
Growth
habit
Height
(m)
Year first
cultivated
Shrub
2.54.6
1656
Tennessee, S to Alabama
& Missouri,W to Texas
Tree
6.19.1
1882
Scientific name(s)
Common name(s)
Occurrence
C. coggygria Scop.
C. americanus Nutt.
C. cotinoides (Nutt.
ex Chapm.) Britt.
C. obovatus Raf.
common smoketree,
smokebush, European
smoketree,Venetian sumac
American smoketree,
yellowwood
438
seeds.
longi-
Cotinus
439
Scarification
in H2SO4 (min)
Moist medium
30
30/60
20/80
20/40
Sand
Sphagnum moss
Peat
Plastic bag
C. coggygria
C. obovatus
Stratification treatments
Temp (C)
Days
3
5
3
3
4560
90
6080
60
Sources: Dirr and Heuser (1987), Gonderman and ORourke (1961), Heit (1968) cited by Rudolf (1974), Stilinovic and Grbic (1988).
Table 3Cotinus, smoketree: germination test conditions and results with pretreated seed
Germination test conditions
Temp (C)
Species
C. coggygria
C. obovatus
Medium
Germinator
Sphagnum
Kimpak in
germinator
Day
20
21
30*
Germination rate
Night
Days
20
21
20
30
21
46
37
Germination
Sound-
Days
ness (%)
11
80
93
39
Samples
2
2
3
70
60
References
Allen DH. 1994. Personal communication. Sandwich, MA: FW Schumacher
Co., Inc.
Blakesley D, Weston GD, Elliott MC. 1991. Endogenous levels of indole-3acetic acid and abscisic acid during the rooting of Cotinus coggygria cuttings taken at different times of the year. Plant Growth Regulation 10(1):
112.
Blakesley D, Weston GD, Elliot MC. 1992. Increased rooting and survival of
Cotinus coggygria cuttings from etiolated stock plants. Journal of
Horticultural Science 67(1): 3337.
Dirr MA. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation, and uses. Champaign, IL:
Stipes Publishing Company. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant
propagation: from seed to tissue culture. Athens, GA.:Varsity Press.
239 p.
Enescv V. 1991. The tetrazolium test of viability. In: Gordon, AG, Gosling P,
Wang BSP, eds.Tree and shrub seed handbook. Zurich: International Seed
Testing Association: 9, 119.
440
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan Publishing. 1290 p.
Macdonald B. 1986. Practical woody plant propagation for nursery
growers. Portland, OR:Timber Press. 669 p.
Oren-Shamir M, Levi-Nissim A. 1997. UV-light effect on the leaf pigmentation of Cotinus coggygria Royal Purple. Scientia Horticulturae 71(1/2):
5966.
Rudolf PO. 1974. Cotinus, smoketree. In: Schopmeyer CS, tech. coord. Seeds
of woody plants in the United States. Agric. Handbk. 450. Washington,
DC: USDA Forest Service: 346348.
Siftar A. 1981. Ucinek kinetina (6-furfurilaminopurin), IBA (indol maslene
kisline) in fenofaz na induciranje adventivnih korenin pri zelenih potaknjencih rdecelistnega ruja (Cotinus coggygria Scop. Royal Purple) v
pogojih meglenja [in Slovenian, with English summary: Effect of kinetin
(6-furfurilamino purin), IBA (indolebutyric acid) and phenophases on
induction of adventitious roots in Venetian sumach softwood cutting
(Cotinus coggygria Scop. Royal Purple) under mist spray conditions].
Zbornik Biotehniske Fakultete Univerze E.K. v Ljubljani 37: 177212.
Spellerberg B. 1985. Verbesserung des Vermehrungserfolges bei Schwer
vermehrbaren Laubgeholzen: 1. Der Einfluss des Vermehrungsklimas auf
Inhaltsstoffe und weiteres Wachtum der bewurzelten Stecklinge [in
German, with English summary: Improved propagation of difficult-to-root
deciduous ornamental shrubs: 1. Influence of climate on carbohydrate
contents and continuous growth of rooted cuttings].
Gartenbauwissenschaft 50(2): 7177.
Cotinus
441
RosaceaeRose family
Cotoneaster Medik.
cotoneaster
Paul E. Slabaugh and Nancy L. Shaw
Dr. Slabaugh (deceased) retired from the USDA Forest Services Rocky Mountain Forest and Range
Experiment Station; Dr. Shaw is a research botanist at the USDA Forest Services Rocky Mountain Research
Station, Forestry Sciences Laboratory, Boise, Idaho
others 2004; Slabaugh 1974). They require little maintenance and provide ground cover, soil stabilization, snow
entrapment, and aesthetic values. Peking cotoneaster provides food and cover for wildlife (Johnson and Anderson
1980; Kufeld and others 1973; Leach 1956; Miller and others 1948). Six species used in conservation plantings are
described in table 1 (Hoag 1965; Nonnecke 1954; Plummer
and others 1977; Rheder 1940; USDA SCS 1988; Zucker
1966). Use of cotoneasters in some areas may be limited due
to their susceptibility to fire blight (infection with the bacterium Erwinia amylovora), borers (Chrysobothris femorata
(Olivier)), lace bugs (Corythucha cydonia (Fitch)), and red
spiders (Oligonychus platani (McGregor)(Griffiths 1994;
Krssmann 1986; Wyman 1986).
Cotoneasters are apomictic and will, therefore, propagate true from seed (Wyman 1986). However, because of the
apomictic habit, many variants occur within each species
(Everett 1982). This variability has been exploited in cultivar
Common name(s)
Occurrence
C. acutifolius Turcz.
C. acutifolia Turcz.
C. pekinensis Zab.
C. apiculatus Rehd. & Wilson
C. apiculata Rehd. & Wilson
C. horizontalis Dcne.
C. davidiana Hort.
C. integerrimus Medic.
C. vulgaris Lindl.
C. lucidus Schltdl.
C. acutifolia Lindl., not Turcz.
C. sinensis Hort.
C. niger (Thunb.) Fries
C. melanocarpus Lodd.
Peking cotoneaster
hedge cotoneaster
black cotoneaster,
darkseed cotoneaster
cranberry cotoneaster
442
fruits.
Cotoneaster
443
Location
Flowering
Fruit ripening
Seed dispersal
C. acutifolius
C. apiculatus
C. horizontalis
C. integerrimus
C. lucidus
C. niger
N Great Plains
S Michigan
Great Plains
North Dakota
MayJune
MayJune
June
MayJune
MayJune
MayJune
SeptOct
AugSept
SeptNov
AugSept
Sept
Septwinter
Fallwinter
Septwinter
Sources: Krssmann (1986), Macdonald (1986), Slabaugh (1974), USDA SCS(1988), Zucker (1966).
Table 3Cotoneaster, cotoneaster: height, year first cultivated, and color of flowers and ripe fruit
Species
C. acutifolius
C. apiculatus
C. horizontalis
C. integerrimus
C. lucidus
C. niger
Height at
maturity (m)
Year first
cultivated
1.83.9
0.31.5
0.91.2
1.23.6
1.82.7
1.52.4
1883
1910
1880
1840
1829
Flower color
Pink
Pink
White-pink
Pinkish
White, tinged w/pink
Pinkish-white
Black
Scarlet
Light to dark red
Red
Black
Blackish red
Sources: Griffiths (1994), Hoag (1958, 1965), LHBH (1976), Leslie (1954), Krssmann (1986), Rehder (1940), Rosendahl (1955), USDA SCS (1988).
Average
/kg
/lb
/kg
/lb
48,46658,212
21,98426,405
59,300
141,094
35,274
51,560
26,900
64,000
16,000
23,390
Sources: Cumming (1960), McDermand (1969), Plummer and others (1968), Slabaugh (1974), Uhlinger (1968, 1970), USDA SCS (1988).
Immersion time in
conc H2SO4 (min)
1090
60120
90180
120
520
1090
Medium
Wet prechill at 4 C
Period (days)
Peat
Sand & peat
Peat
3090
6090
90120
120*
3090
3090
Sources: Dirr and Heuser (1987), Fordham (1962), Leslie (1954), McDermand (1969), Slabaugh (1974), Smith (1951), Uhlinger (1968, 1970), USDA SCS (1988).
* Wet prechilling was preceded by 90 days of warm incubation at 21 C.
444
Species
C. acutifolius
C. horizontalis
C. lucidus
C. niger
Daily
light (hrs)
9
24
24
9
9
Medium
Wet paper
Wet paper
Sand
Wet paper
Wet paper
Percentage germination
Day
25
27
30
25
25
Temp (C)
Night
10
20
10
10
Days
Avg
(%)
100
7080
100
30
70
80
Samples #
5+
Cotoneaster
445
References
Cullum FG, Gordon AG. 1994. Dormancy release of tree and shrub seeds
using a compost activator pretreatment. Combined Proceedings of the
International Plant Propagators Society 43: 125130.
Cumming WA. 1960. Germination studies with Cotoneaster lucida being
conducted at the Canada Experimental Farm, Morden, Manitoba.
Western Canadian Society of Horticulture Report Proceedings 16:
4344.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant
propagation. Athens, GA:Varsity Press. 239 p.
Everett TH, ed. 1982. The New York Botanical Garden illustrated encyclopedia of horticulture.Volume 10. New York: Garland Publishing: 893897.
Fordham AJ. 1962. Methods of treating seeds at the Arnold Arboretum.
Plant Propagators Society Proceedings 1962: 157163.
Griffiths M. 1994. Index of garden plants. Portland, OR:Timber Press.
1234 p.
Heriteau J. 1990. The National Arboretum book of outstanding garden
plants. New York: Simon and Schuster. 292 p.
Hinds LW. 1969. Personal communication. Bismarck, ND: Lincoln-Oakes
Nurseries.
Hoag DG. 1958. Hardy cotoneasters for North Dakota. Bulletin of the
North Dakota Agricultural Experiment Station 20: 3033.
Hoag DG. 1965. Trees and shrubs for the northern plains. Minneapolis:
Lund Press. 376 p.
Huxley A, ed. 1994. The new Royal Horticultural Society dictionary of gardening. London: Macmillan. 1298 p.
ISTA [International Seed Testing Association]. 1993. Rules for testing seeds:
rules 1993. Seed Science and Technology 21 (Suppl.): 1259.
Johnson TK, Anderson ES. 1980. Conservation planting handbook for
Wyoming and Colorado. Laramie: University of Wyoming, Agriculture
Extension Service. 123 p.
Jorgensen K. 1996. Personal communication. Ephraim, UT: Utah Division of
Wildlife Resources.
Krssman G. 1986. Manual of cultivated broad-leaved trees & shrubs.
Volume 3, PRUZ. Portland, OR:Timber Press. 510 p.
Kufeld RC, Wallmo OC, Feddema C. 1973. Foods of the Rocky Mountain
mule deer. Res. Pap. RM-111. Fort Collins, CO: USDA Forest Service,
Rocky Mountain Forest and Range Experiment Station. 31 p.
Leach HR. 1956. Food habits of the Great Basin deer herds in California.
California Fish and Game 42: 243308.
Leslie WR. 1954. Propagation of cotoneaster. Western Canadian Society of
Horticulture Report Proceedings 10: 88.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third. New York:
Macmillan. 1290 p.
446
RosaceaeRose family
Crataegus L.
447
Common name(s)
Occurrence
312
Arnold hawthorn,
anomalous hawthorn
barberry hawthorn,
bigtree hawthorn
blueberry hawthorn,
blue haw, pomette bleu
Brainerd hawthorn
510
fireberry hawthorn,
roundleaf or golden-fruit
hawthorn
510
Kansas hawthorn,
Eggert thorn
cockspur hawthorn,
Newcastle thorn,
hog-apple
Indiana to Kansas, S to
Arkansas & Oklahoma
Quebec to Michigan &
Kansas, S to Florida & Texas
47
C. crus-galli L.
C. acutifolia Sarg.; C. bushii Sarg.
C. canbyi Sarg.; C. cherokeensis Sarg.
C. mohrii Beadle; C.operata Ashe; C.palmeri Sarg.
C. regalis Beadle; C. sabineana Ashe
C. salicifolia Medik. C. signata Beadle
C. subpilosa Sarg.; C. vallicola Sarg.
C. warneri Sarg.
C. dilatata Sarg.
broadleaf hawthorn,
C. conspecta Sarg.
apple-leaf hawthorn
C. locuples Sarg.
C. douglasii Lindl.
black hawthorn, Douglas
C. columbiana Howell
or western black hawthorn,
black thornberry
C. erythropoda Ashe
cerro, chocolate hawthorn
C. cerronis A. Nelson
C. flabellata (Spach) Kirchn.
fanleaf hawthorn
C. densiflora Sarg.; C. grayana Egglest.
C. flava Ait.
yellow hawthorn
summer haw
C. cullasagensis Ashe
C. greggiana Egglest.
Gregg hawthorn
C. harbisonii Beadle
Harbison hawthorn
C. intricata Lange
thicket hawthorn, entangled
or Allegheny hawthorn
C. lacrimata Small
Pensacola hawthorn,
weeping or sandhill hawthorn
C. laevigata (Poir.) DC.
English hawthorn, English
C. oxyacantha L., in part
midland or English woodland
C. oxycanthoides Thuill.
hawthorn
C. marshallii Egglest.
parsley hawthorn, parsley
C. apiifolia (Marshs.) Michaux
haw
C. mollis (Torr. & Gray) Scheele
C. albicans Ashe
C. arkansana Sarg.
C. brachyphylla Beadle
C. cibaria Beadle
C. coccinea var. mollis Torr. & Gray
C. invisa Sarg.;
C. lacera Sarg.;
C. limaria Sarg.
C. monogyna Jac.
C. oxyacantha L. ssp. monogyna
511
615
27
46
510
Quebec to Michigan, S to
New York, Kentucky, & Missouri
48
Alaska to S California,
Ontario to Dakotas, S to
Michigan & Nevada
Wyoming to Washington,
S to New Mexico & Arizona
Maine to Quebec to Michigan,
S to Florida & Louisiana
Maryland & West Virginia,
S to Florida & Mississippi
712
26
46
58
36
24
Virginia to Illinois, S to
Florida & Texas
28
612
oneseed hawthorn,
single-seed or common
512
36
38
17
C
Table 1Crataegus, hawthorn: nomenclature, occurrence, and heights at maturity (continued)
Table 1Crataegus, hawthorn: nomenclature, occurrence, and heights at maturit
Scientific name & synonym(s)
Common name(s)
rufous mayhaw
C. saligna Greene
C. sanguinea Pall.
willow hawthorn
Siberian hawthorn
C. spathulata Michx.
C. microcarpa Lindl.
C. succulenta Schrad. ex Link
C. florifera Sarg.; C. laxiflora Sarg.
C. tracyi Ashe ex Egglest.
C. montivaga Sarg.
C. triflora Chapman
C. uniflora Mnchh.
C. bisculcata Ashe; C. choriophylla Sarg.
C. dawsoniana Sarg.; C. gregalis Beadle
C. viridis L.
C. amicalis Sarg.
C. ingens Beadle
Occurrence
Height at
maturity (m)
712
W Florida to Texas, N to
Arkansas
Maine to Michigan, S to
Virginia & Illinois; South
Carolina & Florida also
New York to Ontario, S to
Pennsylvania & Ohio
New Jersey to Missouri,
S to Florida, Mississippi &
Louisiana
610
British Columbia, S to
48
710
410
46
28
Florida to Mississippi
48
510
Missouri to Kansas, S to
Arkansas & Texas
N Florida, SW Georgia, &
SE Alabama
Colorado
E Russia & Siberia, S to
Mongolia & China
Virginia to Missouri, S to
Florida to Texas
Nova Scotia to Montana, S to
North Carolina & Utah
Texas & NE Mexico
18
Tennessee, S to Georgia
& Louisiana
New York to Missouri,
S to Florida & NE Mexico
Pennsylvania to Kansas,
S to Florida & Texas
39
46
58
58
58
35
46
1/ 4
2
512
Sources: Beadle (1913), Brinkman (1974), Dirr (1998), Flint (1997), Foote and Jones (1989), Griffiths (1994), Jacobson (1996), Little (1980a&b), Palmer (1950, 1952),
Phipps (1988, 1995, 1998a&b), Phipps and OKennon (1998), Phipps and others (1990), Sargent (1933), Strausbaugh and Core (1978),Tidestrom (1933),Vines (1960),
Wasson (2001),Weakley (2002).
Crataegus
449
450
Table 2Crataegus, hawthorn: phenology of flowering and fruiting, and color of ripe fruit
Species
Flowering
Fruit ripening
C. aestivalis
C. x anomala
C. berberifolia
C. brachyacantha
C. brainerdii
C. calpodendron
C. chrysocarpa
C. coccinoides
C. crus-galli
C. dilatata
C. douglasii
C. erythropoda
C. flabellata
C. flava
C. greggiana
C. harbisonii
C. intricata
C. lacrimata
C. laevigata
C. marshallii
C. mollis
C. monogyna
C. nitida
C. opaca
C. pedicellata
C. persimilis
C. phaenopyrum
C. piperi
C. pruinosa
C. pulcherrima
C. punctata
C. reverchonii
C. rufula
C. saligna
C. sanguinea
C. spathulata
C. succulenta
C. tracyi
C. triflora
C. uniflora
C. viridis
Mar
May
MarApr
AprMay
MayJune
MayJune
MayJune
May
June
May
May
AprMay
May
Apr
Apr
May
MayJune
Apr
AprMay
AprMay
May
May
May
FebMar
May
MayJune
May
MayJune
MayJune
AprMay
MayJune
May
MarApr
May
May
AprMay
MayJune
AprMay
May
AprMay
AprMay
MayJune
SeptOct
Oct
Aug
SeptOct
SeptOct
AugSept
Oct
Oct
Sept
AugSept
Oct
Sept
Oct
OctNov
Oct
Oct
Aug
SeptOct
Oct
AugSept
SeptOct
Oct
May
Sept
Oct
SeptOct
AugSept
OctNov
SeptOct
SeptOct
Oct
JuneJuly
Oct
AugSept
SeptOct
SeptOct
SeptOct
Oct
SeptOct
SeptOct
Lustrous, scarlet
Bright crimson
Orange with red face
Bright blue with white wax
Red
Orange-red to red
Yellow to orange to crimson
Glossy, dark crimson
Dull red
Scarlet with dark spots
Lustrous, black to chestnut-brown
Red to wine purple, brown, or black
Crimson
Dark orange-brown or yellow
Bright red
Bright red or orange-red
Greenish or reddish brown
Dull yellow or orange or red
Deep red
Bright scarlet
Scarlet with large dark dots
Bright red
Dull red covered with white wax
Lustrous scarlet with pale dots
Glossy, scarlet
Bright red
Lustrous scarlet
Salmon-orange to scarlet
Dark purple-red
Red
Dull red or bright yellow
Shiny or dull red
Red
Red to blue-black
Bright red
Red
Bright red
Orange-red
Red, hairy
Yellow to dull red to brown
Bright red, orange-red, yellow
Sources: Beadle (1913), Brinkman (1974), Dirr (1998), Everett (1981), Flint (1997), Foote and Jones (1989), Jacobson (1996), Little (1980a&b), Palmer (1950, 1952), Phipps
(1988, 1998a), Phipps and OKennon (1998), Sargent (1933),Vines (1960).
* Color of ripe fruit is highly arbitrary and varies in interpretation amoung authors due to lack of standardization. Accurate determinations of fruit color cannot be
ascertained from herbarium specimens.
Plants growing in Boston, Massachusetts, not in native habitat.
Crataegus
451
452
longitudinal section of a
Provenance
C. chrysocarpa
C. douglasii
C. phaenopyrum
C. punctata
C. sanguinea
C. succulenta
South Dakota
Washington, Idaho, Oregon
Minnesota
Russia
C
Seed wt/fruit wt
kg/kg
lb/100 lb
0.15
0.11
0.15
15.2
11.3
15.0
41,200
10,750
22,600
29,800
4,700
20,600
Samples
1
6
1
2
Crataegus
453
warm temperate climates) that will germinate either in shorter time periods without the cumbersome waiting periods
involved in cold stratification or through innovative seed pretreatment techniques such as fermentation.
Research on vegetative propagation of hawthorns by
stem cuttings is limited. Dirr and Heuser (1987) reported
previous efforts as being rarely successful, whereas Dirr
Species
C. anomala
C. crus-galli
C. douglasii
C. mollis
C. monogyna
C. pedicellata
C. persimilis
C. phaenopyrum
C. punctata
C. sanguinea
C. succulenta
C. tracyi
Scarification*
(hrs)
4.5
0
23
0
0.53
2
0
0.52
2
4
0
0
2
0
0.5
0, 0.5, 2.5, 4.5
Stratification treatments
Warm period
Cold period
Temp (C)
Days
Temp (C)
Days
2127
2125
21
25
30
25
20
20
20
21
2125
2025
2127
3090
21
120
90
21
90
1428
28
1428
120
21
30
29
29
Low
7
5
5
10
35
24
24
24
510
5
5
47
4
4
180
90180
21135
135
84112
120
180
270
7084
84
7084
135
135
21
110140
0, 20, 100
Sources: Brinkman (1974), Felipe Isaac and others (1989), Qrunfleh (1991), St John (1982),Tipton and Pedroza (1986),Young and Young (1992).
* Immersion time in sulfuric acid (H2SO4).
Outdoor winter temperatures.
Species
Medium
C. anomala
C. crus-galli
C. douglasii
C. mollis
C. phaenopyrum
Soil
Soil
Peat or sand
Soil
Soil
Peat
Peat
Peat
Peat
Soil
Germination blotters
C. punctata
C. sanguinea
C. succulenta
C. tracyi
8
21
21
21
21
5
21
21
4
16
2
21
21
21
21
5
21
21
7
16
454
Days
180
21
3545
135
21
21
30
28
Germination
Avg (%)
Samples
35
73
30
4250
71
92
60
73
50
3540
0
1
1
6
3
2
1
1
1
2
2
2
and (b) bark-slippage occurs over a long season (late summer to early fall) (Dirr and Heuser 1987). Cultivars budded
onto Washington hawthorn can be expected to grow 0.9 to
1.2 m in the growing season following budding (Dirr and
Heuser 1987). Cultivars of European species (for example,
English and oneseed hawthorns) should be budded onto
rootstocks of European species, whereas hawthorns native to
North America should be budded onto rootstocks of North
American species (Dirr and Heuser 1987; Hartmann and
others 2002). Aside from these constraints, T-budded
hawthorns appear to be highly compatible across many
species.
Several grafting procedures are employed (rather than
budding procedures) in production of plants of mayhaw.
Cleft grafts for larger rootstocks or whip-and-tongue grafts
for small diameter rootstocks are used widely in late winter
(Payne and Krewer 1990). In Louisiana, cleft grafting is the
most popular grafting method used for western mayhaw
(Bush and others 1991). Other species, such as parsley,
cockspur, Washington, and yellow hawthorns, also can be
used as rootstocks for mayhaws (in particular, western mayhaw) due to graft compatibility (Payne and Krewer 1990).
Brinkman (1974) called for additional trials on
hawthorns to acquire more knowledge on seed biology.
However, little comprehensive research has been conducted
in the intervening 30 years on this subject. Much work
remains to be done before a comprehensive understanding of
propagation of hawthorns will be possible.
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Usher G. 1974. A dictionary of plants used by man. London: Constable.
619 p.
Vines RA. 1960. Trees, shrubs, and woody vines of the southwest. Austin:
University of Texas Press. 1104 p.
Wasson E. 2001. Trees and shrubs: illustrated AZ of over 8500 plants.
Willoughby, NSW, Australia: Global Book Publishing: 239241.
Weakley AS. 2002. Flora of the Carolinas and Virginias [website available at
www.bio.unc.edu/faculty/peet/lab/PEL/weakley_flora].
Young JA,Young CG. 1992. Seeds of woody plants in North America:
revised and enlarged edition. Portland, OR: Dioscorides Press. 407 p.
TaxodiaceaeRedwood family
seed.
Cryptomeria
457
seeds kept in the light than seeds kept in the dark (Chettri
and others 1987). Official test prescriptions for sugi call for
germination on top of moist blotters at alternating temperatures of 20 and 30 C for 28 days; no pretreatment is necessary (ISTA 1993).
Nursery and field practice. Sugi seeds are sown in
Hawaii from November to March. Sowing is by the broadcast method or by using a planter that has been adjusted to
the proper seed size. The planter places seeds in rows about
15 to 20 cm (6 to 7 in) apart. Seeds are covered with 3 to 6
mm (1 to 21/2 in) of soil (Ohmasa 1956; Walters 1974). No
mulch is used in Hawaii (Walters 1974), but a single layer
of straw is used in Japan (Ohmasa 1956). The seedbeds are
given about 75% shade for about 2 months (Walters 1974).
Seedling density in the beds is about 220 to 330
seedlings/m2 (20 to 30/ft2). Frost damage to seedlings in
early winter can be avoided by shading or shortening the
daily period of exposure to solar radiation (Horiuchi and
Sakai 1978). Seedlings are outplanted as 1+0 stock in
Hawaii (Walters 1974). Sugi can be started from cuttings
(Carlson 1959). In an experiment in which the trees were
measured after more than 26 years, there was no significant
difference in any measure of growth between trees started
from seeds and those started from cuttings (Yang and Wang
1984).
longitudinal
References
Carlson NK, Bryan LP. 1959. Hawaiian timber for the coming generations.
Honolulu:Trustees of the Bernice Bishop Estate. 112 p.
Chettri R, Rai B, Basu PK. 1987. Ecological distribution of species in plant
communities of the Sukhia Pokhari Forest, Darjeeling District.
Environment and Ecology 5(3): 590594.
Dallimore W, Jackson A. 1967. A handbook of Coniferae and Ginkgoaceae.
4th ed., rev. New York: St. Martins Press. 729 p.
Hashizume H. 1973. Fundamental studies on mating in forest trees: 5.
Flowering and pollination in Cryptomeria japonica. Bulletin of the Faculty
of Agriculture,Tottori University 25: 8196.
Horiuchi T, Sakai A. 1978. Deep supercooling may be important in frost
avoidance mechanisms of 16 native species. In: Li PH, Sakai A. Plant cold
hardiness and freezing stress: mechanisms and crop implications. St. Paul:
University of Minnesota, Department of Horticulture and Landscape
Architecture. 416 p.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing. Rules 1993. Seed Science and Technology 21 (Suppl.):
1259.
Itoo S, Katsuta M. 1986. Seed productivity in the miniature seed orchard of
Cryptomeria japonica D. Don. Journal of the Japanese Forestry Society
68(7): 284288.
Ohmasa M. 1956. Tree planting practices in temperate Asia: Japan. For. Dev.
Pap. 10. Rome: FAO. 156 p.
Parry MS. 1956. Tree planting practices in tropical Africa. For. Dev. Pap. 8.
Rome: FAO. 302 p.
458
CupressaceaeCypress family
Cupressus L.
cypress
LeRoy C. Johnson and Robert P. Karrfalt
Mr. Johnson retired from the Pacific Southwest Forest and Range Experiment Station; Mr. Karrfalt is director
of the USDA Forest Services National Seed Laboratory, Dry Branch, Georgia
459
Common names
Occurrence
C. abramsiana C.B.Wolf
C. goveniana var. abramsiana (C.B.Wolf) Little
C. arizonica Greene
Arizona cypress
Arizona smooth
cypress
Mendocino cypress,
pygmy cypress
Piute cypress
Cuyamaca cypress
Gowen cypress
C. macnabiana A. Murr.
Guadalupe cypress
Mexican cypress,
cedar-of-Gog,
Portuguese-cedar
MacNab cypress
Monterey cypress
C. sargentii Jepson
stands
C. sempervirens L.
C. sempervirens var. stricta Aiton
C. sempervirens var. horizontalis
(Mill.) Gord.
Cupressus torulosa D.Don
C. torulosa var. corneyana Carr.
Sargent cypress
Sources:
Italian cypress,
Mediterranean cypress
spreading Italian cypress
Mediterranean area
Himalayan cypress,
surai
460
Table 2Cupressus, cypress: growth habit, height, cone and seed ripeness criteria
Height at
Year first
maturity (m) cultivated Cones
Species
Growth habit
C. arizonica
1521
1882
ssp. arizonica
ca. 1909
ssp. nevadensi
ssp. stephensonii
C. bakeri
C. forbesii
C. goveniana
ssp. pygmaea
C. guadalupensis
C. lusitanica
C. macnabiana
C. macrocarpa
C. sargentii
C. sempervirens
var. horizontalis
C
Color ripeness criteria
Seed
1930
1900
1917
1927
Light tan
Rich dark brown
1846
Dull brown
1854
Brownish gray
1838
Brown
Medium brown to
glaucous brown
Dark brown
1908
B.C.
B.C.
ca. 1879
ca. 1670
Dark brown
Sources: Dallimore and Jackson (1967), Raizada and Sahni (1960), Sargent (1965), Sudworth (1915),Wolf and Wagener (1948).
Cupressus
461
/kg
C. arizonica
ssp. arizonica
ssp. nevadensis
ssp. stephensonii
C. bakeri
C. forbesii
C. goveniana
ssp. pygmaea
C. guadalupensis
C. lusitanica
C. macrocarpa
C. macnabiana
C. sargentii
C. sempervirens
C. torulosa
387.6103.2
210.365.1
201.186.7
123.295.0
387.2316.8
112.684.5
313.3253.4
246.4-232.3
356.4100.1
200.6147.8
147.898.6
149.6118.8
238-200
Seeds (x1,000)/weight*
Average
/lb
/kg
/lb
176.246.9
95.629.6
91.439.4
56.043.2
176.0144.0
51.238.4
142.4115.2
112.0105.6
162.045.5
91.267.2
67.244.8
68.054.0
10891
182.6
121
126.5
109.1
359.9
98.6
283.4
239.4
55.0
261.8
167.4
174.2
123.2
137.9
219
83.0
55.0
57.5
49.6
163.6
44.8
128.8
108.8
25.0
119.0
76.1
79.2
56.0
62.7
99
Samples
77+
22+
11+
2
4
2
2
2
1
1
20
2
2
9
Scales/cone
Seeds/cone
68
510
68
68
68
610
35
810
8-10
6-10
812
68
610
814
90120
90100
93
100125
50-85
90110
130
>100
75
140
75105
100
64280
Sources: Goggans and Posey (1968), Rafn (1915), Raizada and Sahni (1960),Toumey and Stevens (1928),Von Carlowitz (1986),Wolf and Wagener (1948).
* Figures are for samples that have foreign matter (twigs, leaves, cone scales, etc.) removed but no attempt was made to separate sound from hollow and other nonviable
seeds.
462
longitudi-
463
Species
C. arizonica
ssp. nevadensis
C. bakeri
C. forbesii
C. goveniana
ssp. pygmaea
C. macnabiana
C. macrocarpa
C. sargentii
C. sempervirens
Days
20
6
30
30
30
30
30
30
30
30
Germination
Average
(%)
Samples
26
6
12
12
22
31
1
24
14
13
9
1
2
2
2
2
1
15
4
37
2
Soundness
Average
(%)
Samples
30
38
36
54
93
5
2
82
41
27
4
1
2
1
2
2
9
Because of variable dormancy, AOSA (1998) also recommends paired tests for Arizona cypress, using unstratified
and stratified (21 days) samples for each lot. Official test
prescriptions have not been developed for the other cypress
species, but similar conditions should be sufficient. For
unstratified Himalayan cypress seeds (Rao 1988), the use of
the alternating germination temperatures of 21 C daytime
and 9 C night time gave 60% germination compared to
33% germination at constant 25 C. Although light appears
to be important, prechilling and alternating temperatures are
the more significant promoters of germination. Light did not
appear necessary for seeds of Arizona cypress (Goggans
1974). The seeds can be watered throughout the test with a
mild solution of fungicide (the same formulation used
above) with no phytotoxicity. Germination test results (table
4) have been low primarily because of the low percentages
of sound seed that are common among seed lots of cypress.
Good estimates of germination can be made with x-ray
analysis of fresh seeds of Italian, Mexican, and Arizona
cypresses (Bergsten and Sundberg 1990; Chavagnat and
Bastien 1991).
Nursery practice. Fall-sowing of cypress seeds has
been recommended (Johnson 1974; Wolf and Wagener
464
References
AOSA [Association of Official Seed Analysts]. 1998. Rules for testing seeds.
Lincoln, NE: Association of Official Seed Analysts. 123 p.
Bailey LH. 1923. The cultivated evergreens. 434 p. New York: Macmillan.
Bannister MH. 1962. Prospects for selection in the cypresses. New Zealand
Journal of Forestry 8: 545559.
Bannister MH, Orman HR. 1960. Cupressus lusitanica as a potential timber
tree for New Zealand. New Zealand Journal Forestry 8: 203217.
Bergsten U, Sundberg M. 1990. IDS-sedimentation of Cupressus lusitanica
seeds. In: Turnbull JW, ed.Tropical tree seed research: Proceedings,
International Workshop; 1989 August 2124; Gympie, Queensland,
Australia. Canberra: Australian Centre for International Agricultural
Research: 99102.
Bolotin M. 1964a. Contributions to the arboreal flora of Israel: Cupressus
sempervirens L. La-Yaaran 14(4): 17.
Bolotin M. 1964b. Segregation in progenies of Cupressus sempervirens L. LaYaaran 14(2): 4648.
Ceccherini L, Raddi S, Andreoli C. 1998. The effect of seed stratification on
germination of 14 Cupressus species. Seed Science and Technology 26:
159168.
Chavagnat A, Bastien JC. 1991. Determination de la qualite des graines de
Cupressus sempervirens L. et Cupressus arizonica E. Greene par la radiographie aux rayons X. Seed Science and Technology 19: 139146.
Dallimore W, Jackson AB. 1967. A handbook of Coniferae and
Ginkgoaceae. 4th ed., rev. by Harrison SG. New York: St. Martins Press.
729 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seeds to tissue culture. Athens, GA:Varsity Press. 239 p.
Gaussen H. 1968. Extant and fossil gymnosperms. Fasc. X.The
Cupressaceae [in French].Travaux du Laboratoire Forestiere de Toulouse
Tome II, Sec. 1,Vol. 1, Part 2, 326 p.
Goggans J F, Posey CE. 1968. Variation in seeds and ovulate cones of some
species and varieties of Cupressus. Circ. 160. Auburn University, AL:
Auburn University Agricultural Experiment Station. 23 p.
Goggans JF, Jones L, Lynch KD. 1974. Germination rate of Arizona cypress
improved by better cone collection techniques and seed germination
treatments.Tree Planters Notes 25(1): 34.
Griffin JR, Stone CO. 1967. MacNab cypress in northern California: a
geographic review. Madroo 19: 1927.
Grigsby HC. 1969. Exotic trees unsatisfactory for forestry in southern
Arkansas and northern Louisiana. Res. Note SO-92. New Orleans:
USDA Forest Service, Southern Forest Experiment Station. 5 p.
Hardham CB. 1962. The Santa Lucia Cupressus sargentii groves and their
associated northern hydrophilous and endemic species. Madroo 16:
173179.
Hedlin AF,Yates HO III,Tovar DC, Ebel BH, Koerber TW, Merkel EP. 1980.
Cone and seed insects of North American conifers. Ottawa: Canadian
Forestry Service; Washington, DC: USDA Forest Service; and Mexico
City: Secretaria de Agricultura y Recursos Hidraulicos, Mexico. 122 p.
Hepting GH. 1971. Diseases of forest and shade trees of the United States.
Agric. Handbk. 386. Washington, DC: USDA Forest Service. 658 p.
ISTA [International Seed Testing Association]. 1993. Rules for testing seeds:
rules 1993. Seed Science and Technology 21 (Suppl.): 1259.
Johnson LC. 1974. Cupressus, cypress. In: Schopmeyer CS, tech coord. Seeds
of woody plants in the United States. Agric. Handbk. 450. Washington,
DC: USDA Forest Service: 363369.
Linnartz NE. 1964. Arizona cypress for Christmas tree production in
Louisiana. For. Note 56. Baton Rouge: Louisiana State University. 4 p.
Little EL Jr. 1966. Varietal transfers in Cupressus and Chamaecyparis.
Madroo 18: 161192.
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
McMillan C. 1952. An experimental inquiry into the edaphic restrictions of
certain members of the California flora [thesis]. Berkkeley: University of
California.
Magini E,Tulstrup NP. 1955. Tree seed notes. For. Dev. Pap. 5. Rome: FAO.
354 p.
Mittal RK, Anderson RL, Mathur SB. 1990. Microorganisms associated with
tree seeds: world checklist 1990. Info. Rep. PI-X-96E/F. Ottawa: Canadian
Forestry Service, Petawawa National Forestry Institute. 57 p.
Paterson DN. 1963.The production of sawn timber from small cypress
thinnings in Kenya. Commonwealth Forestry Review 42: 211-216.
Pharis RP, Morf W. 1967. Experiments on the precocious flowering of
western red cedar and four species of Cupressus with gibberellins A3 and
A4/A7 mixture. Canadian Journal of Botany 45: 15191524.
Posey CE, Goggans JF. 1967. Observations on species of cypress indigenous
to the United States. Circ. 153. Auburn University, AL: Auburn University
Agricultural Experiment Station. 19 p.
Rafn J. 1915. The testing of forest seeds during 25 years, 18871912.
Copenhagen: Langkjaers Bogtrykkeri. 91 p.
Raizada MB, Sahni KC. 1960. Living Indian gymnosperms: 1. Cycadales,
Ginkgoales and Coniferales. Indian Forest Research 5(2): 1150.
Rao PB. 1988. Effects of environmental factors on germination and seedling
growth in Quercus floribunda and Cupressus torulosa, tree species of central Himalaya. Annals of Botany 61: 531540.
Sargent CS. 1965. Manual of the trees of North America (exclusive of
Mexico). 2nd corrected ed. New York: Dover. 910 p.
Schubert GH. 1954. Viability of various coniferous seeds after cold storage.
Journal of Forestry 52: 446447.
Shehaghilo IM. 1987. Some studies on the collection and extraction of
Cupressus lusitanica seed at Lushoto,Tanzania. In: Kamra SK, Ayling RD,
compl. & eds. Proceedings, International Symposium of Forest Seed
Problems in Africa; 1987 August 23September 2; Harare, Zimbabwe.
Umea: Swedish University of Agricultural Science: 280286.
Sudworth GB. 1915. The cypress and juniper trees of the Rocky Mountain
region.Tech. Bull. 20. Washington, DC: U.S. Department of Agriculture.
36 p.
Sudworth GB. 1967. Forest trees of the Pacific slope, with new foreword
and changes in nomenclature. New York: Dover. 455 p.
Toumey JW, Stevens CL. 1928. The testing of coniferous tree seeds at the
School of Forestry,Yale University, 19061926. Bull. 21. New Haven, CT:
Yale University School of Forestry. 46 p.
Von Carlowitz PG. 1986. Multipurpose tree & shrub seed directory.
Nairobi: International Council for Research in Agroforestry. 265 p.
Wolf CB, Wagener WW. 1948. The New World cypresses.Volume 1. El
Aliso, CA: Rancho Santa Ana Botanic Garden. 444 p.
Zeide B. 1977. Germination of cypress seeds in the field. Forest Ecology
and Management 1: 141147.
Cupressus
465
FabaceaePea family
References
Abdullah MM, Jones RA, El-Beltagy AS. 1989. A method to overcome dormancy in Scotch broom (Cytisus scoparius). Environmental and
Experimental Botany 29: 499505.
Bossard CC. 1991. The role of habitat disturbance, seed predation, and ant
disperal on the establishment of the exotic shrub Cytisus scoparius.
American Midland Naturalist 126: 113.
Bossard CC. 1993. Seed germination in the exotic shrub Cytisus scoparius
(Scotch broom) in California. Madroo 40: 4761.
Bossard CC, Rejmanek M. 1994. Herbivory, growth, seed production, and
resprouting of an exotic invasive shrub Cytisus scoparius. Biological
Conservation 67: 193200.
Bravo L. 1980. We are losing the war against broom. Fremontia 15: 2729.
Gill JD, Pogge FL. 1974. Cytisus scoparius (L.) Lk., Scotch broom. In:
Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service:
370371.
Gonzales-Andres F, Ortiz M. 1997. Phenology of species belonging to the
genus Cytisus and allies (Genisteae: Leguminosae). Israel Journal of Plant
Science 45: 5969.
ISTA [International Seed Testing Association]. 1993. Rules for testing seeds:
rules 1993. Seed Science and Technology 21 (Suppl.): 1259.
Munz PA, Keck DD. 1959. A California flora. Berkeley: University of
California Press. 1681 p.
Parker IB. 1997. Pollinator limitation of Cytisus scoparius (Scotch broom),
an invasive exotic shrub. Ecology 78: 14571470.
Tarrega R, Calvo L,Trabaud L. 1992. Effect of high temperature on seed
germination of two woody Leguminosae.Vegetatio 102: 139147.
Weiss FE. 1909. The dispersal of the seeds of the gorse and the broom by
ants. New Phytologist 8: 8190.
Wyman D. 1986. Wymans gardening encyclopedia. Expanded 2nd ed.
New York: Macmillan. 1221 p
Cytisus
467
FabaceaePea family
mature seed.
longitudinal section
seedling at 10 days
References
Duarte O. 1974. Improving royal poinciana seed germination. Plant
Propagator 20(1): 1516.
Francis JK. 1994. Personal communication. Ro Piedras, PR: USDA Forest
Service, International Institute of Tropical Forestry.
Francis JK, Liogier HA. 1991. Naturalized exotic tree species in Puerto Rico.
Gen.Tech. Rep. SO-82. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 12 p.
Little EL Jr, Wadsworth FH. 1964. Common trees of Puerto Rico and the
Virgin Islands. Agric. Handbk. 249. Washington, DC: USDA Forest
Service: 176177.
Marrero J. 1949. Tree seed data from Puerto Rico. Caribbean Forester 10:
1130.
Menninger EA. 1962. Flowering trees of the world. New York: Hearthside
Press. 336 p.
Delonix
469
PapaveraceaePoppy family
Dendromecon Benth.
bushpoppy
W. Gary Johnson and Donald L. Neal
Mr. Johnson retired from the USDA Forest Services National Seed Laboratory; Dr. Neal retired from the
USDA Forest Services Pacific Southwest Forest and Range Experiment Station
Growth habit, occurrence, and use. Bushpoppies
(also known as treepoppies) are openly branched, evergreen
shrubs from 0.6 to 2.5 m high, sometimes to 6 m. They have
a woody base with gray or white shreddy-barked stems. The
2.5- to 10-cm-long leaves are mostly lanceolate, 3 to 8 times
longer than wide (LHBH 1976). Environmental factors and
shoot growth pattern affect leaf size (Bullock 1989). The
2 species considered here grow on dry chaparral slopes,
ridges, and washes below 1,830 m. One species is found in
Californias Channel Islands and the other in the coastal
range, from Sonoma County to the Sierra San Pedro Martir,
Baja California, Mexico, and in the west foothills of the
Sierra Nevada, from Shasta County to Tulare County (table
1). Bush-poppies rely on seed production to propagate. No
lignotuber is formed on sprouts that appear after burning, so
regrowth after fire is rare (Bullock 1989). The genus is useful for watershed protection (Sampson and Jespersen 1963)
and for forage. Goats are especially fond of bushpoppies,
and deer (Odocoileus spp.) and sheep eat the sprouts after
fire.
Flowering and fruiting. Flowers are bisexual, yellow,
showy, and solitary on stalks. At several locations, the
shrubs first flowered in their second spring (Bullock 1989).
Flowers appear in April through June and sometimes into
August (Munz and Keck 1959). Bullock (1989) reports that
the shrubs flower profusely from February through April in
the Santa Monica Mountains. Several populations produce a
Common name(s)
Occurrence
D. harfordii Kellogg
Dendromecon rigida ssp. harfordii (Kellogg) Raven
D. rigida spp. rhamnoides (Greene) Thorne
D. rigida Benth.
island bushpoppy,
Harford tree-poppy
470
longi-
1974). Vivrette reported no germination in of 9 samples tested for 21 days at 15 C. A few seeds germinated on blotters
moistened with 400 ppm GA3 (gibberellic acid). Total viable
seeds as determined by staining in tetrazolium chloride
ranged from 11 to 50%, average 27% (Vivrette 1996).
Emery recommended fire treatment or 1 1/2 to 2 months of
stratification and stated that 3 months of stratification with a
diurnal fluctuation from 8 to 21 C may improve germination (Emery 1988).
Nursery practice. Fire-treated bushpoppy seeds give
the most reliable germination in nurseries (Emery 1988;
Everett 1957). Seeds to be fire-treated should be sown in the
fall in a slightly moist nurserybed. The seeds should be then
covered with a layer of milled peat or sand 1 to 2 times as
thick as the seeds diameter and not watered. Then, a 10- to
15-cm (4- to 6-in) layer of dry pine needles or excelsior
should be placed over the bed and burned. The seedbed
should be watered after it has cooled. If wooden flats are
being used, 2 layers of aluminum foil will protect the wood
during the burning (Emery 1988).
References
Berg RY. 1966. Seed dispersal of Dendromecon: its ecologic, evolutionary
and taxonomic significance. American Journal of Botany 53: 6173.
Bullock SH. 1989. Life history and seed dispersal of the short-lived chaparral shrub Dendromecon rigida (Papaveraceae). American Journal of
Botany 76(10): 15061517.
Emery D. 1988. Seed propagation of native California plants. Santa Barbara,
CA: Santa Barbara Botanical Garden. 115 p.
Everett PC. 1957. A summary of the culture of California plants at the
Rancho Santa Ana Botanic Garden 17271950. Claremont, CA: Rancho
Santa Ana Botanical Gardens. 223 p.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States. New York: Macmillan.
1290 p.
Mirov NT, Kraebel CJ. 1939. Collecting and handling seeds of wild plants.
For. Pub. 5. Washington, DC: USDA Civilian Conservation Corps. 42 p.
Munz PA, Keck DD. 1959. A California flora. Berkeley: University of
California Press. 1681 p.
Neal DL. 1974. Dendromecon rigida Benth., stiff bushpoppy. In: Schopmeyer
CS, tech. coord. Seeds of woody plants in the United States. Agric.
Handbk. 450. Washington, DC: USDA Forest Service: 372.
Sampson AW, Jespersen BS. 1963. California range brushlands and browse
plants. Manual 33. University of California Extension Service, California
Agricultural Experiment Station. 162 p.
Vivrette NJ. 1996. Personal communication. Carpinteria, CA: Ransom Seed
Laboratory.
Dendromecon
471
EbenaceaeEbony family
Diospyros L.
persimmon
David F. Olson, Jr., R. L. Barnes, and W. Gary Johnson
Drs. Olson and Barnes retired from the USDA Forest Services Southeastern Forest Experiment Station;
Mr. Johnson retired from the USDA Forest Services National Seed Laboratory
Common name(s)
Occurrence
D. kaki L.f.
D. chinensis Blume
D. lotus L.
D. japonica Siebold & Zucc.
D. texana Scheele
Japanese persimmon,
kaki, keg fir, date-plum
date-plum
Texas persimmon,
black persimmon
common persimmon,
eastern persimmon
D. virginiana L.
D. mosieri Small
Sources: LHBH (1976), Sargent (1965).
472
S Connecticut to SE Nebraska,
S to Gulf of Mexico
Diospyros
473
increased to 10 weeks (Oh and others 1988). No pretreatment is needed to germinate Texas persimmon seeds (Vora
1989).
Germination tests. Germination of stratified common
persimmon seeds was tested in sand or peat flats at diurnally
alternating temperatures of 20 to 30 C. Germinative energies ranging from 54 to 94% were obtained in 20 to 34
days; and germinative capacities at 60 days varied from 62
to 100% (Olson and Barnes 1974). Payne achieved 90% uniform germination on common persimmon and date-plum by
stratifying the seeds for 60 to 90 days in wet vermiculite
after lightly dusting them with a fungicide. Scratching the
seedcoat can shorten the stratification period (Payne 1996).
Fresh Japanese persimmon seeds taken from ripe fruits
and sown immediately germinate best. Germination ranged
from 20 to 77% in a study of 18 cultivars with fresh seeds
sown immediately (Dirr and Heuser 1987). Date-plum seeds
germinated best without light at alternating 18 to 30 C with
10 weeks stratification at 5 C. Germination of seeds stratified for 2 weeks was increased by treating them with 500
ppm gibberellin (GA3) (Oh and others 1988). Fresh Texas
persimmon seeds sown immediately after extraction germinated 33%. Germination was reduced with all other treatments (Dirr and Heuser 1987).
The tetrazolium chloride staining test is often used to
estimate the viability of common persimmon and date-plum
seeds due to the long stratification period needed to overcome dormancy. Clipping the radicle end of a seed with toenail clippers and soaking the seed for several days in water
or 500 ppm GA3 will soften it. Then it should be cut lengthwise to expose the embryo and storage tissue for staining.
Nursery practice. Common persimmon seeds may be
fall-sown or stratified and sown in the spring. In Missouri,
fall-sowing at a depth of 5 cm (2 in) is the normal practice,
and seedbeds are mulched. Steavenson (Olson and Barnes
1974) reported a tree percent of 50%; an average tree percent of 25 to 33% is easily attainable. Seedlings of this
/kg
3,0153,790
1,4603,880
Average
/lb
/kg
/lb
Samples
1,3701,720
6651,765
1,550
8,910
2,640
3,400
4,040
1,200
2
1
474
species have a strong taproot (figure 3) and should be fieldplanted at the end of the first season. Root wrenching will
cause the seedlings to form a compact, fibrous root system
(Myatt and others 1988).
Little EL Jr, Skolmen RG. 1989. Common forest trees of Hawaii (native and
introduced). Agric. Handbk. 679. Washington, DC: USDA Forest Service.
321 p.
Little EL Jr, Woodbury RO, Wadsworth FH. 1974. Trees of Puerto Rico and
the Virgin Islands. 2nd volume. Agric. Handbk. 449. Washington, DC:
USDA Forest Service. 1024 p.
Morris RC. 1965. Common persimmon (Diospyros virginiana L). In: Silvics of
forest trees of the United States. Agric. Handbk. 271. Washington, DC:
USDA Forest Service: 168170.
Myatt A, Huffman G, Odell J Jr. 1988. Seed processing manual. Washington,
OK: Oklahoma Department of Agriculture, Forestry Division.
Oh JH, Kim SK, Ahn HK. 1988. Studies on seed germination of Diospyros
species. Journal of the Korean Society for Horticultural Science 29(4):
297303.
Olson DF Jr, Barnes RL. 1974. Diospyros virginiana L., common persimmon.
In: Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service:
373375.
Payne JA. 1996. Personal communication. Byron, GA: USDA Agricultural
Research Service.
Radford AE, Ahles HE, Bell CR. 1964. Guide to the vascular flora of the
Carolinas. Chapel Hill: University of North Carolina Book Exchange.
Sargent CS. 1965. Manual of the trees of North America. 2nd ed., corrected and reprinted. New York: Dover. 934 p.
Thornhill HB. 1968. Propagation of woody ornamentals by seeds. American
Nurseryman 127(6): 18, 8689.
Vora RS. 1989. Seed germination characteristics of selected native plants of
the lower Rio Grande Valley,Texas. Journal of Range Management 42(1):
3640.
References
Aroeira JS. 1962. Dormencia e conservacao de sementes de algumas plantas frutiferas. Experientiae 2: 541609.
Blomquist HL. 1922. Dormancy in seeds of persimmon. Elisha Mitchell
Scientific Society Journal 38: 14.
Crocker W. 1930. Harvesting, storage, and stratification of seeds in relation
to nursery practice. Boyce Thompson Institute Professional Paper 1:
114120.
Dirr M, Heuser C Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Engstrom HE, Stoeckler JH. 1941. Nursery practice for trees and shrubs
suitable for planting on the prairie-plains. Misc. Pub. 434. Washington, DC:
USDA. 159 p.
Everitt JH. 1984. Germination of Texas persimmon seed. Journal of Range
Management 37: 189192.
Harlow WM, Harrar ES. 1958. Textbook on dendrology. New York:
McGraw-Hill. 561 p.
Kotobuki K. 1978. Seed storage of Japanese persimmon, Diospyros kaki. In:
Long term preservation of favourable germ plasm in arboreal crops.
Ibaraki-Ken, Japan: Fruit Tree Research Station, Ministry of Agriculture and
Forestry: 3642.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concinse
dictionary of plants cultivated in the United States and Canada. New
York: Macmillan. 1290 p.
Little EL Jr, Delisle AL. 1962. Time periods in development: forest trees,
North America. In: Biological handbook on growth. Washington, DC:
Federation of American Societies for Experimental Biology: table 104.
Diospyros
475
ThymelaeaceaeMezereum family
Dirca palustris L.
eastern leatherwood
John C. Zasada, David S. Buckley, Elizabeth Ann Nauertz, and Colleen F. Matula
Dr. Zasada retired from the USDA Forest Services North Central Research Station; Dr. Buckley is an
ecologist and Ms. Nauertz is a biological technician at the USDA Forest Services North Central Research
Station, Grand Rapids, Minnesota; Ms. Matula is a botanist at the USDA Forest Services Ottawa National
Forest, Bessemer Ranger District, Bessemer, Michigan
Growth habit, occurrence, and use. Eastern leatherwoodDirca palustris L.is also known as moosewood,
rope-bark, and wicopy. Its natural distribution extends from
New Brunswick to Ontario in the north and from northern
Florida to Louisiana in the south (Fernald 1950). Within this
range, the distribution is restricted to very specific site conditions. It is found almost exclusively in mesic, relatively
rich hardwood forests or mixed coniferhardwood forests
(Alban and others 1991; Curtis 1959; Fernald 1950; Ferrari
1993; Jones 2000; Kotar and others 1988; Meeker and others 1993; Neveling 1962; Rooney 1996; Soper and
Heimberger 1982; Voss 1985; Weir-Koetter 1996). In aspen
ecosystems across the upper Great Lakes region, leatherwood is present in stands with a relatively high aspen site
index and a significant northern hardwood component
(Alban and others 1991). In Ontario, the northern limit of
distribution is similar to that of beech (Fagus grandifolia
Ehrh.) and sugar maple (Acer saccharum Marsh.)(Soper and
Heimberger 1982). The distribution of plants on a particular
site can vary from apparently random to aggregated (Jones
2000). Forests in which leatherwood is common are characterized by a dense overstory that permits relatively little
light to reach the forest floor during the growing season. It
is often the only true understory shrub in these stands; the
other woody understory species are tolerant to mid-tolerant
treesfor example, sugar maple, ironwood (Ostrya virginiana (Mill.) K. Koch.), white ash (Fraxinus americana L.),
eastern hemlock (Tsuga canadensis (L.) Carr.), and balsam
fir (Abies balsamea (L.) Mill.)having the capacity to grow
into the overstory (Alban and others 1991; Buckley 1996;
Ferrari 1993; Jones 2000).
Western leatherwoodD. occidentalis Grayis very
similar to eastern leatherwood (Neveling 1962). Its distribution is limited to the wooded hills of the San Francisco Bay
region (Vogelman 1953). Flower descriptions and morphological comparisons of the 2 species are provided by
Vogelman (1953). A related species in the Thymelaeaceae
Daphne mezereum L.is an introduced species that has
476
become established in some areas. The information presented here is for eastern leatherwood; some of it may also
apply to western leatherwood.
In its natural habitat, eastern leatherwood reaches a
height of 3 to 4 m and basal diameters of 5 to 10 cm. Crown
width and depth of larger plants can be as much as 2 to 3 m;
the largest crown volumes that we have measured are in the
range of 15 to 25 m3. Crown architecture can be fairly complex, with frequent branching and numerous apical growing
points (figure 1). The largest individuals that we have
observed are in old-growth northern hardwood forests where
logging is prohibited and in older hardwood stands managed
under a single-tree selection system. The maximum age
attained by leatherwood is not known, but 30- to 50-yearold plants occur in older hardwood forests.
Figure 1Dirca palustris, eastern leatherwood: mature
forest-grown plant in full flower, with plant about 1.3 m tall.
Dirca
477
478
ripe fruits
Table 1Dirca palustris, eastern leatherwood: flowering and fruit production in 2 forest types in northern Wisconsin
Stand type*
Hardwood forest
1995
1995
1996
Pine forest
1995
1995
1996
No. 3-flower
clusters/shrub
Mean
Range
% cluster
w/13
fruits
Fruits/plant
Mean
Range
153
198
5515
21695
23
55
22
20
48
41
30
19
40
194
0386
13840
59
85
0255
0325
23
0
14
77
57
23
29
36
045
0260
Note: fpc = fruits per cluster; these populations did not have 4-flower or 4-fruit clusters in the 2 years of observation.
* Based on 15 randomly selected shrubs in each stand.
To obtain total number of flowers, multiply by 3.
Dirca
479
Figure 6Dirca palustris, eastern leatherwood: generalized longitudinal section of mature seed, based on Neveling
(1962) and Buckley (1996).
480
wall, develops between the fleshy fruit wall and the hard
inner seed coat, making it fairly easy to hand-clean small
quantities of seeds. The stony seedcoat is easily broken with
the pressure of a fingernail and the seed can be squashed by
squeezing between the thumb and forefinger with moderate
pressure. Consequently, any type of mechanical cleaning
must be done with care.
No information was found on the best ways to handle
fruits or store seeds. Seeds remain viable in the forest floor
from the time of dispersal until they germinate in the spring
(del Tredici 1984), suggesting that storage for at least 8 to
10 months is possible. Seeds are exposed to a fairly wide
range of temperature and moisture conditions between dispersal and germination.
Germination. Detailed information on the effects of
environmental conditions on germination was not found. del
Tredici (1984) used a number of standard methods to stimulate gemination, but untreated seeds planted in a nursery bed
soon after they were collected were the only ones that produced seedlings (67% germination). Dirr and Heuser (1987)
reported that both cleaned and uncleaned (fleshy fruit wall
removed) seeds produced seedlings. In controlled environment studies, a seedlot was observed to produce germinants
over at least a 3-year period (Zasada and others 1996).
Nursery. Based on the limited information available,
we recommend planting seeds soon after collection with and
without the fleshy fruit wall in order to provide the range of
conditions under which seeds appear to germinate naturally;
seeds sown this way will germinate the next spring (del
Tredici 1984; Dirr and Heuser 1987). Dirr and Heuser
(1987) reported finding a number of young plants under a
mature plant growing in a landscaped area, indicating that it
might be possible to obtain some small seedlings from these
situations. The growth rate of seedlings under open conditions is not documented. In its natural habitat, seedlings
grow to a height of 20 to 30 cm in 5 to 10 years.
There has been little or no success in stimulating rooting
in stem cuttings (Dirr and Heuser 1987). Layering occurs
under natural conditions, suggesting that air-layering is a
potential option for propagating leatherwood. However,
Hendricks (1985) reported that air-layered stems did not
produce roots or callus during an 8-week period. Our observations of layering of branches and the main stem under natural conditions suggest that it might take longer than 8
weeks for rooting to occur.
References
Alban DH, Perala DA, Jurgensen MF, Ostry ME, Probst JR. 1991. Aspen
ecosystems properties in the upper Great Lakes. Res. Pap. NC-300. St.
Paul: USDA Forest Service, North Central Forest and Range Experiment
Station. 47 p.
Alden HA. 1995. Hardwoods of North America. Gen.Tech. Rep. FPL-83.
Madison, WI: USDA Forest Service, Forest Products Laboratory. 136 p.
Curtis JT. 1959. The vegetation of Wisconsin. Madison: University of
Wisconsin Press. 657 p.
Buckley DS. 1996. Unpublished data. Rhinelander, WI: USDA Forest
Service, North Central Research, Forestry Sciences Laboratory.
del Tredici P. 1991. Propagating leatherwood: a lesson in humility. Arnoldia
51 (4): 6266.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Dirr MA. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation, and uses. Champaign, IL:
Stipes Publishing. 1007 p.
Esson JG. 1949. Leatherwood for early spring bloom. Journal of the New
York Botanical Gardens 50: 5759.
Fernald ML. 1943. The fruit of Dirca palustris. Rhodora 45: 117119.
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1632 p.
Ferrari JB. 1993. Spatial patterns of litterfall, nitrogen cycling, and understory
vegetation in a hemlockhardwood forest [PhD thesis]. St. Paul:
University of Minnesota. 180 p.
Hendricks DR. 1985. Air layering native woody plants. Proceedings of the
International Plant Propagators Society 34: 528531.
Dirca
481
FabaceaePea family
482
legume.
seeds.
longitudinal
Nursery practice. There is little information on nursery practices for Texas-ebony. Nurserybed densities of 160
to 215/m2 (15 to 20/ft2) appear to be suitable for raintree
(Albizia saman (Jacq.) F. Muell.), a similar species (Walters
and others 1974), and the same is suggested for Texasebony. Optimum planting depth in a greenhouse was reported to be 1 cm (0.4 in) (Alaniz and Everitt 1978). Direct
seeding in old fields in Texas was improved by mulching the
seeds with a commercial straw blanket (Vora and others
1988), and either mulching or shading would seem to be
beneficial in nursery beds in that region.
References
Alaniz MA, Everitt JH. 1978. Germination of Texas ebony seeds. Journal of
the Rio Grande Valley Horticultural Society 32: 95100. [Horticultural
Abstracts 49(12): 9556; 1979].
Little EL, Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
Vora RS. 1989. Seed germination characteristics of selected native plants of
the lower Rio Grande Valley,Texas. Journal of Range Management 42:
3640.
Vora RS, Schumacher RW, Labus Z. 1988. Planting seeds of four south
Texas native species under mulch cover.Texas Journal of Science 40:
456458.
Walters GA, Bonner FT, Petteys EQP. 1974. Pithecellobium Mart., blackbead.
In: Schopmeyer CS,Tech. Cord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service:
639640.
Ebenopsis
483
ElaeagnaceaeOleaster family
Elaeagnus L.
elaeagnus
David F. Olson, Jr., and Jill R. Barbour
Dr. Olson retired from the USDA Forest Services Southeastern Forest Experiment Station; Ms. Barbour is a
germination specialist at the USDA Forest Services National Seed Laboratory, Dry Branch, Georgia
Common name(s)
Occurrence
Russian-olive, oleaster,
narrow-leafed oleaster
silverberry, wolfberry
autumn-olive,
autumn elaeagnus
Sources: Fernald (1950), Harrington (1954), Olson (1974), Rehder (1940), Small (1933).
Species
Location
Flowering
Fruit ripening
Seed dispersal
E. angustifolia
E. commutata
E. umbellata
June
JuneJuly
MayJune
AugOct
AugSep
AugOct
All winter
SepNov
SepNov
1213
89
68
Source: Borell (1962), Dietz (1969), Hora (1981), McDermand (1969), Radford and others (1964), Rehder (1940).
fruit.
Figure 2Elaeagnus, elaeagnus: achenes with fleshy perianth removed of E. angustifolia, Russian-olive (left) and
E. commutata, silverberry (right).
Figure 3Elaeagnus angustifolia, Russian-olive: longitudinal section through an achene enclosed in the fleshy perianth.
485
Table 3Elaeagnus, elaeagnus: height; seed-bearing age, seedcrop frequency, and fruit ripeness criteria
Species
Height at
maturity (m)
E. angustifolia
469
E. commutata
E. umbellata
1.84.6
0.93.7
Minimum
seed-bearing
age (yr)
Years
between large
seedcrops
Long cultivated
1813
1830
12
Year first
cultivated
Silver-gray outer;
emon-yellow inside
Silver
Red-pink brown scales
Species
E. angustifolia
E. commutata
E. umbellata
Seed wt/fruit
wt ratio
1560
510
/kg
7,65015,400
5,95010,140
46,52584,670
Cleaned seeds/weight
Range
/lb
3,4706,990
2,7004,600
21,10038,400
Average
/kg
/lb
Samples
11,380
8,380
62,180
5,160
3,800
28,200
15
5
30
Sources: Belcher and Washburn (1965), Carroll (1971), Harrington (1954), Heit (1970), Hinds (1967), McDermand (1969), Mickelson (1968), Molberg (1969), Mowry
(1971), Olson (1974), Schumacher (1968), Zarger (1968).
486
both ends to open the embryo cavity. After 48 hours of soaking in 1% tetrazolium chloride, the seeds should be cut longitudinally to expose the embryos. The radicle tips and as
much as one-third of the distal cotyledons can be unstained,
and the seeds still considered viable. A secondary procedure
calls for longitudinal cuts at the beginning.
Nursery practice. Seeds may be sown 13 to 25 mm
(1/2 to 1 in) deep in the late summer or fall without stratification, or in the spring after 10 to 90 days of cold stratification (Baker 1969; Growl 1968; Hinds 1967; Jack 1969;
McDermand 1969; Mickelson 1968; Molberg 1969; Olson
1974; Zarger 1968). July seeding after 90 days of stratification gave excellent germination of Russian-olive in southeast
Saskatchewan (Cram and Elliott 1966). In Michigan,
autumn-olive is seeded by broadcasting 1.7 kg of fresh fruit/
10 m2 of bed area (1 lb/25 ft2) (Carroll 1971). At the Los
Lunas Plant Material Center, Russian-olive is sown at a rate
of 200 seeds, or 40 g (1.4 oz) of clean seeds/m, which yields
150 usable plants. In areas with a large population of mice,
the pulp should be removed and cleaned seeds used for sowing (Carroll 1971). Russian-olive seedlings are susceptible
to damage from rabbits and must be protected if these
rodents are a problem.
Soil splash, which coats the pubescent leaves of newly
emerged seedlings, is an important cause of mortality, and
consequently, nursery beds should be mulched to cover the
soil and prevent rain spattering (Carroll 1971; Growl 1968;
Table 5Elaeagnus, elaeagnus: germination test conditions and results for stratified seedlots*
Species
E. angustifolia
E. commutata
E. umbellata
E. umbellata
30
30
30
30
30
20
20
20
20
10
60
2140
28
28
50
28
Germinative
energy
Amt (%)
Days
776
27
1032
10
52
13
Germinative capacity
Average
Samples
779
5490
30
68
60
41
93
32
11
1
19
1
57
Sources: Belcher and Washburn (1965), Heit (1968), Molberg (1969), Olson (1974).
* Seeds were stratified for 10 to 90 days at 1.1 to 10 C.
Elaeagnus
487
AsteraceaeAster family
Encelia Adans.
encelia
Jane E. Rodgers and Carol Miller
Ms. Rodgers is at the USDI Point Reyes National Seashore, Point Reyes, California; Ms. Miller was a
vegetation specialist at the USDI Joshua Tree National Parks Center for
Arid Lands Restoration and Native Plants Nursery,Twentynine Palms, California
Other common names. brittlebush, bush-sunflower.
Growth habit, occurrence and uses. The brittlebush
genusEnceliaincludes 14 species of low branching
shrubs native to western America. The plants are suffrutescent, often with a pungent odor (Benson and Darrow 1954).
Ray flowers (sometimes absent) are yellow, usually conspicuous when present, and produce neither pollen or fertile
seeds. Disk flowers are yellow or purple (Benson and
Darrow 1954). Species frequently hybridize, especially in
disturbed areas. Species commonly found in the southwestern United States are listed in table 1.
The brittle wood secretes a clear resin used by Native
Americans as a glue. In some parts of Mexico, the resin has
been burned as incense for religious ceremonies (Benson
and Darrow 1954). The Cahuilla of the southwestern United
States have used gum from this plant as a medicine; the gum
was heated and applied to the chest to relieve pain (Bean
and Saubel 1972).
Flowering and fruiting. Flowering can begin in
February and continue through July, weather conditions permitting. Most encelia flowerheads are yellow or a brown- or
yellow-purple. The achenes are densely compressed, obovate or wedge-shaped, with edges that are long-ciliate and
faces that are glabrous or short-hairy (figures 1 and 2)
(Jepson 1993).
Collection, extraction, and storage. Timing of seed
collection is critical, as the achenes are easily blown from
the plant after maturity (Kay and others 1977). Seeds may
be hand-harvested and stored successfully for several years.
Cleaning is difficult, for seeds are often mixed with dry
flower and plant parts of similar size and weight. Studies on
long-term storage of seeds of rayless encelia and Acton brittlebush showed good germination after 4 and 14 years,
respectively (Kay and others 1988). Seeds of both species
that were stored under 4 conditions (15 C, 4 C, room
temperature, and warehouse temperatures) also showed significantly poorer germination rates in the warehouse after
3 years. Storage conditions studied by Padgett and others
(1999) showed that seedlots stored for 6 months in a stan-
achenes.
Common names
Occurrence
E. californica Nutt.
E. farinosa Gray ex Torr.
E. frutescens (Gray) Gray
E. virginensis A. Nels.
E. virginensis var. actonii
(Elmer) B.L.Turner
California brittlebush
brittlebush, incienso, goldenhills
rayless encelia, green brittlebush
Virgin River encelia, brittlebush
Acton brittlebush
488
longitudinal
0 1 47 65 55
References
Bean LJ, Saubel KS. 1972. Temalpakh (from the earth): Cahuilla Indian
knowledge and usage of plants. Morongo Indian Reservation, CA: Malki
Museum Press. 225 p.
Benson L, Darrow RA. 1954. The trees and shrubs of the southwestern
deserts.Tuscon: University of Arizona Press. 437 p.
CALR [Center for Arid Lands Restoration]. 1993. Seed test data filed
19891993.Twentynine Palms, CA: National Park Service, Joshua Tree
National Park.
Emery DE. 1988. Seed propagation of native California plants. Santa
Barbara, CA: Santa Barbara Botanical Garden. 115 p.
Jepson WL. 1993. The Jepson manual of higher plants of California.
Hickman JC, ed. Berkeley: University of California Press. 1400 p.
Kay BL. 1975. Test of seeds of Mojave Desert shrubs. Prog. Rep. BLM
Contract 53500-CT4-2(N). 24 p.
Kay BL, Brown CR, Graves WL. 1977. Virgin River encelia. Mojave Reveg.
Notes 4. Davis: University of California Department of Agronomy and
Range Science.
Kay BL, Graves WL,Young JA. 1988. Long-term storage of desert shrub
seed. Mojave Reveg. Notes 23. Davis: University of California
Department of Agronomy and Range Science.
Padgett P,Vazquez L, Allen EB. 1999. Seed viability and germination
behavior of the desert shrub Encelia farinosa. Madroo 46(3): 126133
Encelia
489
FabaceaePea family
490
Collection, cleaning, and storage. Seeds can be collected in quantity by picking up the legumes after they have
fallen to the ground. They can be separated by macerating
the legumes and then washing them to remove the sticky
syrup or by picking the seeds from the tough legumes with
the point of a knife (Francis 1988). One thousand to a few
thousand seeds are produced per tree. The 1.3- to 1.9-cm
(1/2- to 3/4-in) seeds (figure 2) number 1,100/kg (500/lb)
(Janzen 1983; Neal 1965). The seeds store well according to
Bauer (1982).
Germination. Without scarification, a moderate percentage of the seeds germinate over a span of several
Figure 1Enterolobium cyclocarpum, guanacaste:
and legume.
seeds
longitu-
References
Bauer J. 1982. Especies con potential para la reforestacin en Honduras,
resumenes.Tegucigalpa, Honduras: Corporacin Hondurea de
Desarrollo Forestal. 42 p.
Bertoni VR, Juarez VM. 1980. Comportamento de nueve especies forestales
tropicales plantadas en 1971 en el campo experimental El Tormento.
Revista Ciencia Forestal 25(5): 439.
Chudnoff M. 1984. Tropical timbers of the world. Agric. Handbk. 607.
Washington, DC: USDA Forest Service. 466 p.
Fournier LA, Herrera de Fournier ME. 1977. La sucescin ecologica como
un metodo eficaz para la recuparacin del bosque de Costa Rica.
Agronoma Costarricense 1(1): 2329.
Francis JK. 1988. Enterolobium cyclocarpum (Jacq.) Griseb., guanacaste, earpod tree. Res. Note SO-ITF-SM-15. New Orleans: USDA Forest Service,
Southern Forest Experiment Station. 4 p.
Francis JK, Rodrguez A. 1993. Seeds of Puerto Rican trees and shrubs: second installment. Res. Note SO-374. New Orleans: USDA Forest Service,
Southern Forest Experiment Station. 5 p.
Guridi LI. 1980. La madera en las artisanas del estado de Michoacana.
Boletn Divulgativo 50. Distrito Federal, Mxico: Instituto Nacional de
Investigaciones Forestales. 128 p.
Holdridge, LR, Poveda LR. 1975. Arboles de Costa Rica.Volume 1. San Jos,
Costa Rica: Centro Cientfico Tropical. 546 p.
Hughes CE, Styles BT. 1984. Exploration and seed collection of multiplepurpose dry zone trees in Central America. International Tree Crops
Journal 3: 131.
Janzen DH. 1983. Costa Rican natural history. Chicago: University of
Chicago Press. 816 p.
Little EL Jr, Woodbury RO, Wadsworth FH. 1974. Trees of Puerto Rico and
the Virgin Islands,Volume 2. Agric. Handbk. 449. Washington, DC: USDA
Forest Service. 1024 p.
Neal MC. 1965. In gardens of Hawaii. Spec. Pub. 50. Honolulu: Bernice P.
Bishop Museum. 924 p.
Pennington TD, Sarukhan J. 1968. Manual para la identificatin de campo de
los principales rboles tropicales de Mexico. Distrito Federal, Mxico:
Instituto Nacional de Investigaciones Forestales. 413 p.
Salazar R. 1985. Tcnicas de produccin de lea en fincas pequeas.
Turrialba, Costa Rica: Centro Agronmico de Investigaciones y
Enseanza. 459 p.
Enterolobium
491
EphedraceaeEphedra family
Ephedra L.
ephedra or Mormon-tea
Susan E. Meyer
Dr. Meyer is a research ecologist at the USDA Forest Services Rocky Mountain Research Station,
Shrub Sciences Laboratory, Provo Utah
Table 1Ephedra, ephedra: habit, habitat, and geographic distribution of some species used in revegetation
Scientific name
Common name(s)
Habit
Habitat
Distribution
E. nevadensis S.Wats.
Nevada Mormon-tea,
gray Mormon-tea, gray
ephedra, gray Nevada joint-fir
Torrey Mormon-tea,
Torrey ephedra,
Torreys joint-fir
green Mormon-tea,
Brigham tea, green
ephedra, Mormon-tea
Sprawling, gray-green,
leaves in pairs, bases
W US
E. torreyana S.Wats.
E. viridis Coville
492
Sprawling, gray-green,
leaves in whorls of 3,
bases gray
Erect, broomlike, bright
green, leaves in pairs,
bases black
Colorado Plateau,
Chihuahuan Desert
W US
seed.
/g
3357
108128
2862
Mean
/oz
9351,615
3,0603,630
7901,760
/g
43
118
47
/oz
% viability
Range
Mean
1,220
3,345
1,330
8494
7691
46100
89
83
89
Sources: Belcher (1985), Kay and others (1977), Meyer (1995), Meyer and others (1995), Monsen (1995).
Ephedra
493
seeds
long-
References
otherwise nondormant seeds from precocious summer germination. Ephedra seeds germinate readily during prolonged
chilling. Kay and others (1977) reported 76% germination
during a 30-day stratification at 2 C for a Mojave desert
collection of Nevada Mormon-tea. In chilling experiments
with the 6 seedlots mentioned above, weeks to 50% germination at 1 C varied from 6 to 7 weeks for the Torrey
Mormon-tea collections and from 8 to 9 weeks for collections of the other 2 species. All viable seeds germinated during chilling within 12 weeks.
Official rules for testing green Mormon-tea call for a 4week test at 15 C, with the option of a 4-week prechill for
494
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
Belcher E, ed. 1985. Handbook on seeds of browse shrubs and forbs.Tech.
Pub. R8-TP8. Atlanta: USDA Forest Service, Southern Region. 246 p.
Freeman DC, Klikoff LG, Harper KT. 1976. Differential resource utilization
by the sexes of dioecious plants. Science 193: 597599.
Kay BL, Ross CM, Graves WL, Brown CR. 1977. Gray ephedra and green
ephedra. Mojave Reveg. Notes 19. Davis: University of California,
Department of Agronomy and Range Science. 8 p.
Meyer SE. 1995. Unpublished data. Provo, UT: USDA Forest Service, Rocky
Mountain Research Station.
Meyer SE, Kitchen SG, Wilson GR, Stevens R. 1988. Proposed rule: Ephedra
viridis green mormon tea. Association of Official Seed Analysts
Newsletter 62(1): 1819.
Monsen SB. 1995. Unpublished data. Provo, UT: USDA Forest Service,
Rocky Mountain Research Station.
Stevens R, Jorgensen KR, Davis JN. 1981. Viability of seed from thirty-two
shrub and forb species through fifteen years of warehouse storage.
Great Basin Naturalist 41: 274277.
Welsh SL, Atwood ND, Higgins LC, Goodrich S. 1987. A Utah flora. Great
Basin Naturalist Memoirs: 1894.
Young JA, Evans RA, Kay BL. 1977. Ephedra seed germination. Agronomy
Journal 61: 209211.
EricaceaeHeath family
Epigaea repens L.
trailing-arbutus
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Epigaea
495
When the seedlings have 3 to 5 leaves above the cotyledons, they may be transplanted to individual pots. High
humidity should be maintained until the plants are well
established (Barrows 1936). In 1 year, the plants develop
into rosettes about 10 cm (4 in) in diameter (Blum and
Krochmal 1974). Plants will tolerate a fairly wide range of
acidity. Wild plants in Connecticut grew on soils ranging in
pH from 7.67 to 4.65, but the larger plants occurred on the
more acid soils (Barrows 1936; Coville 1911; Lemmon
1935; Steffek 1963).
Trailing-arbutus thrives best in association with mycorrhizal fungi. Including soil that was collected near healthy
wild plants in soil mixtures will introduce the necessary fungus (Barrows 1936; Coville 1911, 1915). The mycorrhizal
fungus also appears to be essential for propagation from cuttings (Barrows 1936). Stem cuttings taken in August have
given 94% rooting in a sandpeat mixture without any treatment (Dirr and Heuser 1987).
References
Bailey LH. 1949. Manual of cultivated plants. New York: Macmillan. 851 p.
Barrows FL. 1936. Propagation of Epigaea repens L.: 1. Cuttings and seeds.
Contributions of the Boyce Thompson Institute 8: 8197.
Blum BM, Krochmal A. 1974. Epigaea repens L., trailing-arbutus. In:
Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service:
380381.
Clay K. 1983. Myrmecochory in the trailing arbutus (Epigaea repens L.).
Torrey Botanical Club Bulletin 110: 166169.
Coville FV. 1911. The use of acid soil for raising seedlings of the mayflower,
Epigaea repens. Science 33(853): 711712.
Coville FV. 1915. The cultivation of the mayflower. National Geographic
Magazine 27: 518519.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation. Athens, GA:Varsity Press. 239 p.
496
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1632 p.
Jacobs ML, Burlage HM. 1958. Index of plants of North Carolina with
reputed medicinal uses. Chapel Hill, NC. 322 p.
Lemmon RS. 1935. The trailing arbutus in home gardens. Horticulture 13:
101102.
Lincoln WC Jr. 1980. Laboratory germination of Epigaea repens, mayflower
or trailing arbutus. AOSA Newsletter 54(1): 7273.
Steffek EF. 1963. Wild flowers and how to grow them. 5th ed. New York:
Crown. 192 p.
Stupka A. 1964. Trees, shrubs, and woody vines of the Great Smoky
Mountains National Park. Knoxville: University of Tennessee Press.
186 p.
AsteraceaeAster family
achene
longi-
Ericameria
497
Germination. Parish goldenweed seeds are not dormant, and no pretreatments are required to stimulate germination (Emery 1964). Seeds sown on sand began germinating in 4 days, and a maximum of 95% was obtained (Mirov
and Kraebel 1937). Germination is, however, usually much
lower (about 20%) because of a high percentage of defective
seeds (Ratliff 1974). Parish goldenweed may also be propagated by cuttings (Jepson 1951).
498
References
Emery D. 1964. Seed propagation of native California plants. Santa Barbara
Botanical Garden Leaflet 1(10): 8196.
Everett PC. 1957. A summary of the culture of California plants at the
Rancho Santa Ana Botanic Garden 19271950. Claremont, CA: Rancho
Santa Ana Botanic Garden. 223 p.
Jepson WL. 1951. A manual of the flowering plants of California. Berkeley:
University of California Press. 1025 p.
Mirov NT, Kraebel CJ. 1937. Collecting and propagating the seeds of
California wild plants. Res. Note 18. Berkeley, CA: USDA Forest Service,
California Forest and Range Experiment Station. 27 p.
Munz PA, Keck DD. 1959. A California flora. Berkeley: University of
California Press. 1681 p.
Ratliff RD. 1974. Haplopappus parishii, Parish goldenweed. In: Schopmeyer
CS, tech. coord. Seeds of woody plants in the United States. Agric.
Handbk. 450. Washington, DC: USDA Forest Service: 445.
PolygonaceaeBuckwheat family
Eriogonum Michx.
wild-buckwheat, buckwheatbrush
Susan E. Meyer
Dr. Meyer is a research ecologist at the USDA Forest Services Rocky Mountain Research Station,
Shrub Sciences Laboratory, Provo, Utah
499
500
Habitat
Source:
E. shockleyi S.Wats.
PULVINATE/ MAT-FORMING
E. bicolor M.E. Jones
E. ovalifolium Nutt.
pretty buckwheatbrush
cushion wild-buckwheat,
roundleaf buckwheat
Shockley wild-buckwheat,
mat buckwheat
E. jamesii Benth.
Wyeth wild-buckwheat,
parsnipflower buckwheat
James wild-buckwheat
shortstem wild-buckwheat
lace buckwheatbrush,
buckwheatbrush, crisp-leaf buckwheat
Mojave buckwheatbrush, California
buckwheatbrush, flat-top buckwheatbrush
Heermann buckwheatbrush,
molecule model plant
Common name(s)*
E. heracleoides Nutt.
SUBSHRUBS
E. brevicaule Nutt.
E. fasciculatum Benth.
SHRUBS
E. corymbosum Benth.
Species
Central Utah
Widespread,W North America
Range
Figure 2Eriogonum fasciculatum, Mojave buckwheatbrush: longitudinal section through a seed excised from
an achene.
ly, as the radicle end of the achene is often slender and easily damaged. Achene weights vary both among and within
species but are usually in the range of 350 to 1,360/g
(10,000 to 39,000/oz) (table 2). Seed quality is also variable
(table 2).
There are few published reports of viability evaluation
beyond germination percentages obtained without pretreatment, which may underestimate viability if there is a dormant fraction. Stevens and others (1996) report that viabilities of >90% may be expected from sulfurflower and Wyeth
wild-buckwheats in an agronomic setting if seeds are harvested when fully mature; these values are comparable to
those for wild-collected lots of many species (table 2).
Insects may damage 10 to 35% of the fruits prior to harvest,
but damaged seeds are normally eliminated in cleaning.
Post-harvest damage from insect infestations is also possible
(Stevens and others 1996). There is little information on
maintenance of viability during storage for species of wildbuckwheat. Stevens and others (1996) report high viability
for sulfurflower and Wyeth wild-buckwheats during 10 to 15
years in warehouse storage, which would indicate orthodox
storage behavior. Other species are perhaps comparable.
Seed germination and testing. Germination is
epigeal (figure 3). Seedlots of many species of wild-buckwheats contain at least a fraction that will germinate without
any pretreatment (tables 2 and 3) (Young 1989). The size of
this fraction depends on species and on the particular lot
involved. Stevens and others (1996) report that seeds of
sulfurflower and Wyeth wild-buckwheats lose dormancy
during short periods of dry storage, and Mojave buckwheatbrush seeds are also reported to dry after-ripen (Kay and
others 1977). Dormant seeds of most species we have examined lose dormancy during chilling at 1 C for periods of 8
to 12 weeks (table 3).
To date there are no formal procedures for evaluating
the seed quality of wild-buckwheat species, and tetrazolium
(TZ) staining is probably the procedure most commonly
employed. To evaluate using TZ, achenes are soaked
overnight in water, clipped through both pericarp and seed
coat at the cotyledon end (the wide end or hilum), and
placed in 1% TZ solution for several hours at room temperature. Achenes are bisected longitudinally for evaluation
(Belcher 1985).
Field seeding and nursery practice. Wild-buckwheats are generally readily established from direct seeding
(Ratliff 1974; Stevens and others 1996; Tiedemann and
Driver 1983; Zamora 1994). They establish best when seeded at a depth of 2 to 5 mm (1/16 to 3/16 in), either by drilling
or by broadcasting followed by covering (for example, raking). Seeding should take place before the season of maximum precipitation, which is generally fall or early winter in
Eriogonum 501
Achenes/weight
/lb
/g
SHRUBS
E. corymbosum
E. fasciculatum
E. heermannii
SUBSHRUBS
E. brevicaule
E. heracleoides
E. jamesii
E. niveum
E. umbellatum
Viability
Test
900
2,000
1,330
5201,085
660
410,000
900,000
600,000
236,000490,000
300,000
93
434
2046
95
Germination %, no pretreatment
Germination %, no pretreatment
Post-chilling cut test
700
350
310
350
1,2901,360
470
265
320,000
160,000
141,000
160,000
585,000620,000
213,000
120,000
84
95
87
5272
86
Germination %; no pretreatment
Post-chilling cut test
960
990
750
436,000
450,000
340,000
47
95
86
PULVINATE/MAT-FORMING
E. bicolor
E. ovalifolium
E. shockleyi
Sources: Belcher (1985), Kay and others (1977), Meyer and Paulsen (2000), Stevens and others (1996),Tiedemann and Driver (1983).
* Post-chilling cut tests (AOSA 1996) are considered accurate for recently harvested seedlots; however, tetrazolium staining (TZ) is required for seedlots stored for more
than 2 years.
Samples
2
3
3
2
2
4
No chill
3
28
4
54
22
7
65
100
30
91
98
74
86
100
55
94
98
99
16 weeks
96
100
77
100
100
100
lifted carefully. Other woody wild-buckwheats could probably be produced as bareroot stock, but no published information is available. Wild-buckwheats may also be produced
as container stock; book planters or tube containers such as
those used for producing conifer seedlings are most appropriate. Nondormant lots may be direct-sown, whereas seedlots requiring chilling may be sown as chilled seed or as
young germlings (Landis and Simonich 1984). Seedlings of
many species grow rapidly and should not be held in small
containers for more than a few months. Many species flower
the first year and may even form flowering stalks while still
in small tube containers.
Figure 3Eriogonum fasciculatum, Mojave buckwheatbrush: very young seedling (left) and older seedling
(right).
References
AOSA [Association of Official Seed Analysts]. 1996. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
Belcher E, ed. 1985. Handbook on seeds of browse-shrubs and forbs.Tech.
Pub. R8-TP8. Atlanta: USDA Forest Service, Southern Region. 246 p.
Kay BL, Ross CM, Graves WL. 1977. California buckwheat. Mojave Reveg.
Notes 5. Davis: University of California, Department of Agronomy and
Range Science. 4 p.
Landis TD, Simonich EJ. 1984. Producing native plants as container seedlings.
In: Murphy PM, comp.The challenge of producing native plants for the
intermountain area. Proceedings, Intermountain Nurserymans
Association 1983 Conference; 1983 August 811; Las Vegas, NV. Gen.
Tech. Rep. INT-168. Ogden, UT: USDA Forest Service, Intermountain
Forest and Range Experiment Station: 1625.
Meyer SE, Paulsen A. 2000. Chilling requirements for seed germination of
ten Utah species of wild buckwheat (Eriogonum Michx.: Polygonaceae).
Native Plants Journal 1: 1824.
Ratliffe RD. 1974. Eriogonum fasciculatum Benth., California buckwheat. In:
Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service:
382383.
Shaw NL. 1984. Producing bareroot seedlings of native shrubs. In: Murphy
PM, comp.The challenge of producing native plants for the intermountain area. Proceedings, Intermountain Nurserymans Association 1983
Conference; 1983 August 811; Las Vegas, NV. Gen.Tech. Rep. INT-168.
Ogden, UT: USDA Forest Service, Intermountain Forest and Range
Experiment Station: 615.
Stevens R, Jorgensen KR,Young SA, Monsen SB. 1996. Forb and shrub seed
production guide for Utah. Logan: Utah State University Extension
Service. 51 p.
Tiedemann AR, Driver CH. 1983. Snow eriogonum: a native halfshrub to
revegetate winter game ranges. Reclamation and Revegetation Research
2: 3139.
Tiedemann AR, Lambert SM, Carlson JR, Perry CJ, Shaw NL, Welch BL, Driver
CH. 1997. Umatilla snow buckwheat for rangeland restoration in the
interior Pacific Northwest. Rangelands 19(3): 2225.
Young JA. 1989. Germination of seeds of sulphur flower. Journal of Seed
Technology 13: 3138.
Zamora BA. 1994. Use of Eriogonum for reclamation. Hortus Northwest
5(1): 911, 47.
Eriogonum
503
MyrtaceaeMyrtle family
Eucalyptus LHer.
eucalyptus
Stanley L. Krugman and Craig D.Whitesell
Dr. Krugman retired from the World Bank and from the USDA Forest Services Washington Office;
Dr.Whitesell retired from the USDA Forest Services Pacific Southwest Research Station
504
Common name(s)
Natural range
Extension
Australia
California, Hawaii,
& Arizona
California & Hawaii
Central & N
Queensland, Australia
SE Australia
SE Australia
California
SE Australia
California
SE Australia
California
SE Australia
California, Hawaii,
& Arizona
E Australia
E Australia
SE Australia
California, Florida,
& Hawaii
California
California & Hawaii
SE Australia
California
E Australia
E Australia
SE Australia
E Australia
California, Florida,
Hawaii, & West Indies
W Australia
E Australia
SE Australia
SE Australia
Sources: Chippendale and others (1969), Johnston and Marryat (1965), Krugman (1974).
Jacobs 1979; Penford and Willis 1961; Pryor 1979). A number of hybrids have been described, but their value for planting in the United States must still be demonstrated. When
grown under plantation conditions outside their natural habitat, species hybridization will occur more often, and seed
collections from small plantations of closely related species
should be discouraged if hybrid seeds are not desired
(Boden 1964).
Flowering and fruiting. The flower clusters develop
enclosed within an envelope formed by 2 bracteolessmall
leafy structures. These bracteoles split and are shed during
development, revealing the flower buds (Boland and others
1980; Penford and Willis 1961). The perfect flowers are
white, yellow, or red, often in axillary umbels, corymbose,
Eucalyptus
505
Table 2Eucalyptus, eucalyptus: height at maturity and phenology of flowering and fruiting of trees grown in
California
Species
Height at
maturity (m)
Flowering
Fruit ripening
Seed dispersal
1836
2439
1836
3083
1860
412
4554
4254
3045
3090
1575
2442
3660
52105
2427
915
1545
1230
1545
FebApr
NovJan
JuneAug
AprJune
AprMay
JulyAug
NovApr
SeptNov
DecFeb
AprJuly
AprJuly
FebMay
DecMar
AprJuly
JanMar
JanMar
AprJune
JuneSept
All year
JulyOct
MayAug
AugOct
AprJuly
JulyAug
MaySept
OctMar
MayJune
MayAug
JanApril
JuneSept
OctDec
OctNov
MayJuly
NovFeb
OctMar
MayJune
All year
E. camaldulensis
E. citriodora
E. dalrympleana
E. delegatensis
E. fastigata
E. glaucescens
E. globulus
E. grandis
E. microcorys
E. nitens
E. obliqua
E. paniculata
E. pilularis
E. regnans
E. robusta
E. rudis
E. saligna
E. sideroxylon
E. viminalis
Source: Krugman (1970).
Species
E. camaldulensis
E. citriodora
E. delegatensis
E. fastigata
E. glaucescens
E. globulus
E. nitens
E. obliqua
E. regnans
E. robusta
E. saligna
E. sideroxylon
E. viminalis
506
0.751.75
4.25
1.253.75
1.253.25
1.252.5
2.25
1.252.5
1.02.0
1.252.5
1.5
1.25
1.02.0
1.252.5
0.51.0
2.5
1.01.75
0.51.25
1.01.75
1.75
1.01.75
0.751.25
0.51.25
0.75
1.0
0.751.5
1.5
Seed color
Chaff color
Yellow-brown
Black
Pale brown or brown
Pale brown or brown
Black or dark brown
Dark brown
Black or dark brown
Dark brown
Pale brown or brown
Dark brown
Black
Dark brown or black
Black or dark brown
Yellow-brown to orange
Brownish red
Pale brown or brown
Pale brown or brown
Pale red-brown
Brownish red
Pale red-brown
Orange-brown or brown
Pale brown or brown
Brownish red
Brownish red
Orange-brown
Pale red-brown
longitudinal
507
Kiln-drying
Temp
(C)
Time
(days)
Temp
(C)
Time
(hrs)
32
32
21
32
32
21
21
1
3
5
3
3
4
6
59
59
59
59
3
6
5
6
1958b). A few speciessuch as tingiringy-gum and alpineash, brown-barrel, shining, and mountain-ash eucalyptusesare normally dormant at the time of collection and
will require pretreatment. Stratification of moist seeds stored
in a plastic bag at temperatures of 3 to 5 C for a period of 3
weeks will break the dormancy of these 5 species, except for
alpine-ash eucalyptus, which should be stratified for 4
weeks (Boland and others 1980; Grose 1969). Longer stratification periods of 6 to 8 weeks are often recommended.
Dormancy between different seedlots of the same
species can vary considerably. In addition, different methods
of extraction and storage can induce dormancy in nondormant seed or strengthen primary dormancy in normally dormant seeds (Krugman 1970). If the seeds fail to germinate
after the recommended shorter stratification periods, then a
longer period should be tried before the seeds are considered
nonviable. Because most seeds are stored before they are
used, stratification for 3 to 4 weeks at a temperature of 3 to
5 C is recommended for all eucalyptus seeds to ensure
faster and more uniform germination (Hinkle 1968;
Krugman 1970).
In a few cases, chemicals have been employed to overcome seed dormancy. The germination of unstratified and
dormant seeds of alpine-ash, brown-barrel, and mountainash eucalyptuses was improved by germinating the seeds in
a solution of gibberellic acid (Bachelord 1967). However,
not all seedlots of the same species responded to gibberellic
acid (Krugman 1970).
Germination tests. Standard methods for testing germination in other seeds are not used for eucalyptus seeds
because of their small size and the presence of so much
chaff, which can exceed the weight of viable seeds. Instead,
samples for germination are of equal weight, not number
(Boland and others 1980; Grose and Zimmer 1958b; ISTA
1993; Turnbull and Doran 1987). Such methods as the
excised-embryo and tetrazolium tests are impractical
(Boland and others 1980; Grose and Zimmer 1958b). The
Species
E. camaldulensis
E. citriodora
E. dalrympleana
E. delegatensis
E. fastigata
E. glaucescens
E. globulus
E. grandis
E. microcorys
E. nitens
E. obliqua
E. paniculata
E. pilularis
E. regnans
E. robusta
E. rudis
E. saligna
E. sideroxylon
E. viminalis
/g
652,100
80220
65285
40125
90210
40120
2070
2001,200
530900
230550
20160
65340
785
20530
220700
2701,100
85915
65440
265445
1,80059,500
2,2006,200
1,8008,100
1,1003,500
2,5005,900
1,0003,000
5009,100
5,60034,000
1,50025,600
6,60015,700
5004,500
1,8009,600
2002,400
60015,000
6,20020,000
7,60031,000
2,40026,000
1,80012,500
7,50012,600
770
140
195
75
150
35
150
700
85
385
85
75
35
315
390
600
460
240
350
Samples #
21,900
4,000
5,500
2,100
4,300
2,000
2,500
20,000
6,800
10,900
2,400
5,000
1,000
8,900
11,000
17,000
13,000
6,800
10,000
41
15
7
13
6
2
10
13
22
7
18
8
28
11
12
9
9
16
6
germination test
Daily light
exposure (hrs)
Temp
(C)
24
0
24
0
0
24
24
0
24
24
0
24
0
24
24
24
24
24
24
35
25
25
20
25
20
25
25
20
20
20
20
20
25
20
35
25
20
25
Days
14
14
14
14
14 or 21
14 or 21
14
14
28
14 or 21
14
28
14
21
28
14
28
14
14
Eucalyptus
509
% Germination
E. citriodora
E. grandis
E. microcorys
E. pilularis
E. robusta
E. sideroxylon
51
98
76
11
84
100
69
Duration (days)
15
29
24
29
18
21
49
510
5,000
1,700
2,830
800
180
160
400
3,160
References
Bachelord EP. 1967. Effects of gibberellic acid, kinetin, and light on the germination of dormant seeds of some eucalypt species. Australian Journal
of Botany 15: 393401.
Barnard RC. 1967. Some garden eucalypts. Australian Plants 4(30): 6970,
7172.
Blakely WF. 1955. A key to the eucalypts. 2nd ed. Canberra: Forest and
Timber Bureau. 359 p.
Blomstedt C, Cameron J, Whiteman P, Chandler SF. 1991. Micropropagation
of juvenile Eucalyptus regnans (mountain ash). Australian Journal of
Botany 39(2): 179186.
Boden RW. 1964. Hybridization in eucalyptus. Indian Forester 90(9):
581586.
Boland DJ, Brooker MIH,Turnbull JW. 1980. Eucalyptus seed. Sydney,
Australia: Hogbin Poole Pty. Ltd. 191p.
Chippendale GM, Johnston, RD, Kelly S. 1969. Eucalypts. Melbourne,
Australia: Thomas Nelson (Australia), Ltd. 82 p.
Eucalyptus
511
Geary TF, Meskimen GF, Franklin EC. 1983. Growing eucalypts in Florida for
industrial wood production. Gen.Tech. Rep. SE-23. USDA Forest Service,
Southeastern Forest Experiment Station. 43 p.
Grose RJ. 1969. Personal communication. Melbourne, Australia:
Commonwealth Forestry.
Grose RJ, Zimmer WJ. 1958a. A description of the seeds of 70 Victorian
eucalypts. Bull. 8. Melbourne, Australia: Commonwealth Forestry. 24 p.
Grose RJ, Zimmer WJ. 1958b. The collection and testing of seed from some
Victorian eucalypts with results of viability tests. Bull. 10. Melbourne,
Australia: Commonwealth Forestry. 14 p.
Hall N, Johnston RD, Marryatt R. 1963. The natural occurrence of the eucalypts. Leafl. 65, 2nd ed. Canberra, Australia: Forestry and Timber Bureau.
122 p.
Hamilton WD. 1979. The landscape eucalyptus tree: an evaluation study
19711978. Journal of Arboriculture 5(9): 210216.
Hinkle EH. 1968. Eucalyptus in Taiwan. Quarterly Journal of Chinese
Forestry 1(2): 8798.
Holmes DA, Floyd AG. 1969. Nursery techniques for raising eucalypts in
Jiffy pots on the New South Wales north coast. Res. Note 22. Sydney,
Australia: Commonwealth Forestry. 15 p.
Hunt R, Zobel B. 1978. Frost-hardy eucalyptus grow well in the Southeast.
Southern Journal of Applied Forestry 2(1): 6-10.
ISTA [International Seed Testing Association]. 1966. International rules for
seed testing. Proceedings of the International Seed Testing Association
31(1): 52106.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing: rules 1993. Seed Science and Technology 21 (Suppl. ):
1259.
Jacobs MR. 1955. Growth habits of the eucalypts. Canberra, Australia:
Forestry and Timber Bureau. 262 p.
Jacobs MR. 1979. Eucalypts for planting. For. Ser. 11. Rome: FAO. 677 p.
Johnston RD, Marryatt R. 1965. Taxonomy and nomenclature of eucalypts.
Leafl. 92. Canberra, Australia: Forestry and Timber Bureau. 24 p.
Karschon R. 1967. Ecotypic variation in Eucalyptus camaldulensis Dehn. In:
Contributions on eucalypts in Israel. [National University Institute of
Agricultural, Ilanot, and Land Development Authority]: Kiriat Hayim III:
3553.
King JP, Krugman SL. 1980. Tests of 36 Eucalyptus species in northern
California. Res. Pap. PSW-152. Berkeley, CA: USDA Forest Service, Pacific
Southwest Forest and Range Experiment Station: 6 p.
Krugman SL. 1970. Observations recorded 19651970. Berkeley, CA:
USDA Forest Service, Pacific Southwest Forest and Range Experiment
Station. 11 p.
512
Krugman SL. 1974. Eucalyptus, eucalyptus. In: Schopmeyer CS, tech. coord.
Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 384392.
Larsen E. 1965. Germination of Eucalyptus seed. Leafl. 94. Canberra,
Australia: Forestry and Timber Bureau. 16 p.
LeBarron RK. 1962. Eucalypts in Hawaii: a survey of practices and research
programs. Misc. Pap. 64. Berkeley, CA: USDA Forest Service, Pacific
Southwest Forest and Range Experiment Station. 24 p.
Ledig FT. 1989. Improvement of eucalypts for fuel and fiber in California. In:
Pereira JS, Landsberg JJ, eds. Biomass production by fast growing trees.
Proceedings, NATO Advanced Research Workshop, Series E. Dordrecht,
The Netherlands: Kluwer Academic Publishers: 231245.
Linnard W. 1969. Cultivation of eucalypts in the USSR. Forestry Abstracts
30(2): 199209.
Metcalf W. 1961. Progress with eucalypts in North America [unpublished
office report]. Section A, United States mainland. National Report, 2nd
World Eucalyptus Conference, Brazil: 118.
Miles M. 1990. Principles of species and clonal selection for farm forestry.
California Eucalyptus Grower 5(1): 8, 9, 11.
Penford AR, Willis JL. 1961. The eucalypts: botany, cultivation, chemistry, and
utilization. New York: Interscience Publishers. 551 p.
Pryor ID. 1979. Reproductive habits of the eucalypt. Unasylva 30(119/120):
4246.
Pryor LD, Byrne OR. 1969. Variation and taxonomy in Eucalyptus camaldulensis. Silvae Genetica 18(3): 6471.
Scott L. 1972. Viability testing of eucalpt seeds. Leafl. 116. Camberra,
Australia: Forestry and Timber Bureau. 24 p.
Toyama S, Moromizato S. 1957. Breeding of eucalyptus trees (report 1):
characteristics of eucalyptus seed [in Japanese]. Government Forest
Experiment Station (Tokyo) Bulletin 103: 124.
Turnbull JW, Doran JC. 1987. Germination of Australian native plant seed.
In: Langkamp PJ, ed. Melbourne, Australia: CSIRO, Division of Forest
Research: 4657, 186198.
Uhr SC. 1976. Eucalypt: the wonder tree. American Forests 82(10): 4243,
5963.
Whitesell CD, DeBell DS, Schubert TS, Strand RF, Crabb TB. 1992. Shortrotation management of eucalyptus: guidelines for plantations in Hawaii.
Gen.Tech. Rep. PSW- 137. Albany, CA: USDA Forest Service Pacific
Southwest Research Station. 30 p.
Willan, RL. 1985. A guide to forest seed handling. For. Pap. 20/2. Rome:
FAO. 379 p.
Zon R, Briscoe JM. 1911. Eucalypts in Florida. Bull. 87. Washington, DC:
USDA Forest Service. 47 p.
CelastraceaeBittersweet family
Euonymus L.
euonymus, spindletree
John C. Zasada and Paul O. Rudolf
Dr. Zasada retired from the USDA Forest Services North Central Forest Research Station;
Dr. Rudolf (deceased) retired from the USDA Forest Services North Central Forest Experiment Station
Common name(s)
Occurrence
E. atropurpurea Jacq.
E. bungeana Maxim.
E. europaea L.
E. hamiltoniana spp.
maackii (Rupr.) Komarov
E. obovata Nutt.
E. verrucosa Scop.
winterberry euonymus
European spindletree,
European euonymus
Maack euonymus
running strawberry-bush,
running euonymus
warty-bark euonymus,
warty spindletree
Euonymus
513
Height at
maturity (m)
0.93.1
0.91.8
1.86.2
4.06.2
3.17.1
1.55.2
0.30.6
0.92.2
Figure 1Euonymus, euonymus: top views of open capsules of E. americana, American strawberry-bush (left) and
E. atropurpurea, eastern wahoo (right).
Year first
cultivated
1860
1697
1756
1883
Long ago
1880
1820
1763
Figure 2Euonymus americana, American strawberrybush: seeds enclosed in their fleshy arils.
tent is about 50% (fresh weight basis) (Lee and others 1991;
Nielsen and Iroquoian 1988). Lee and others (1991)
described the seeds of European spindletree as poisonous
and little used by birds; however, Snow and Snow (1988)
document a substantial use of fruits by birds, stating that the
seeds are not consumed.
Collection of fruits. Seeds may be collected in late
summer or fall by picking the ripe fruits from the bushes or
trees by hand or by shaking them onto an outspread canvas.
They should then be spread out to dry for several days in a
warm room but need not be completely dry to be cleaned
(Myatt and others 1991).
Extraction and storage of seeds. Seeds can be
processed with a macerator (Stein and others 1974). The
plate on the separator should be set slightly larger than the
seeds and adjusted as necessary to prevent too many seeds
from being lost with the pulp (Myatt and others 1991).
Smaller seedlots can be extracted by beating the fruits
in a canvas bag and then rubbing them through a coarse,
round-hole grain screen. The fruits may be macerated in
water and the seeds extracted by flotation (Rudolf 1974).
Following dry extraction, the chaff can be removed by
Species
Location
Flowering
Fruit ripening
E. alata
E. americana
E. atropurpurea
E. bungeana
E. europaea
E. hamiltoniana
ssp. maackii
E. obovata
E. verrucosa
NE US
NE US & Europe
MayJune
MayJune
MayJune
June
MayJune
SeptOct
SeptOct
SeptOct
SeptOct
AugNov
NE US
NE US & Germany
June
AprilJune
MayJune
Oct
AugOct
AugOct
Sources: Fernald (1950), Lancaster (1981), Radford and others (1964), Rehder (1940), Sus (1925), Snow and Snow (1988),Voss (1985),Wappes (1932),Wyman (1947).
Table 4Euonymus, euonymus: fruit form and color of flowers, fruits, and seeds
Color
Species
Fruit form
Flower
Ripe fruit
Seed
Aril
E. alata
Yellowish
Reddish-purplish
Brown*
Orange-red
Reddish green
greenish purple
Purple
Pink-rose
Scarlet
Pinkpurple
Yellowish
white
Light brown
Yellowish
Whitish or
pinkish
White
Orange
Yellowish green
Yellowish
pinkish white
Rose red-pink
Orange
Yellowish
Greenish purple
Brownish
Pink
Crimson
Yellowish red
Red
Tan
Black
Orange
Orange-scarlet
Orange-red
E. americana
E. atropurea
E. bungeana
E. europaea
E. hamiltoniana
ssp. maackii
E. obovata
E. verrucosa
Smooth, deeply 3- to
4-lobed, 4-celled
Deeply 4-lobed
& 4-angled
Smooth, 4-lobed,
3 to 5celled
4-lobed
Usually 3-lobed
Deeply 4-lobed
Scarlet
Sources: Bailey (1939), Dirr (1990), Fernald (1950), Rehder (1940), Snow and Snow (1988).
* Black, in one variety.
Whitish, in one variety.
Seed not wholly covered by aril.
515
Pregermination treatments. Seeds of most euonymus species have dormant embryos. Cold stratification is
adequate to break dormancy for some species, but warm
stratification followed by a cold period is needed for maximum germination for other species (table 7) (Dirr 1990; Dirr
and Heuser 1987; Nikolaeva 1967; Singh 1985; Yu and others 1976). The length of the warm period should be adjusted, depending on the temperature used for cold stratification. For example, Nikolaeva (1967) suggests a 2- to 3month period of warm stratification if cold stratification is at
0 to 3 C. Table 7 provides the range of temperatures for
warm and cold stratification that have been effective for
breaking dormancy. Nikolaeva (1967) provides a thorough
discussion of the effects of temperature, water availability,
seed maturation, and storage alone and in combination on
germination. There may also be some variation in germination among European spindletree seeds formed under different climatic conditions (Dawidowicz-Grezgorzewska and
Beranger-Novat 1989).
Variation in dormancy can be significantly different
among plants. Nielsen (1988), for example, reported that
germination of European spindletree seeds collected from 10
different plants varied from 0 to 30% following stratification
Place collected
E. alata
E. americana
E. atropurpurea
NE US
Durham Co., North Carolina
Carver Co., Minnesota; Cole
Co., Missouri; & rangewide
US
Russia, Netherlands, & NE US
Clinton Co., Michigan
Russia
E. bungeana
E. europaea
E. obovata
E. verrucosa
/kg
Cleaned seeds/weight
Average
/lb
/kg
/lb
41,11069,620
63,300100,113
18,60031,500
30,00045,300
55,250
77,571
25,000
35,100
2+
3+
19,22788,400
18,78535,360
35,95058,870
8,70040,000
8,50016,000
16,30026,700
37,349
29,835
29,393
49,725
45,084
16,900
13,500
13,300
25,500
20,400
8
1+
32+
1
10+
Sources: Barnes (1969), Gorshenin (1941), Heit (1968a), NBV (1946), Nielsen and Eriksen (1988), Rudolf (1974), Sus (1925), Swingle (1939).
Species
E. atropurpurea*
E. europaea
E. obovatas*
E. verrucosa
Air-dry
Dry
Moist
Air-dry
Air-dry
Moist
1.13.3
1520 or 2.8
1.13.3
20
1520 or 2.8
Sources: Gorshenin (1941), Heit (1967), NBV (1946), Nikolaeva (1967), Rudolf (1974), Sus (1925), Swingle (1939).
* Seed stored in closed containers.
516
Samples
12
7
2
7
medium
E. alata
E. bungeana
E. europaea
Sand or peat
Perlitepeat mix
Sand
Sand
Sand, peat, or filter paper
Sand, peat, or filter paper
E. americana
E. atropurpurea
Warm period
Cold period
Temp (C)
Days
Temp (C)
Days
2030
2025
1520
0
0
60
0
0
6090
0
0
6090
010
5
5
2.85
2.810
3250
2.85
010
2.85
010
90100
139
60
60180
61120
60120
60120
6090
60150
120150
Sources: Heit (1968a), Nikolaeva (1967), Rudolf (1974), Shumailina (1949), Swingle (1939).
at 4 to 6 C. This variation may have been significantly different if seeds had been subjected to warm stratification
before cold stratification (table 7) (Nikolaeva 1967).
The morphological changes that occur during pretreatment are important indicators of the adequacy of the
pregermination treatment. An increase in seed volume,
cracking of the seedcoat, and protrusion of the tip of the
hypocotyl occur during warm stratification or warm stratification hastens these changes when seeds are moved to cold
stratification. Completeness of germination depends on these
changes in seed morphology (Nikolaeva 1967).
There seems to be little information on the natural germination pattern of euonymus seeds. Untreated seeds of
517
Species
E. americana
E. atropurpurea
E. bungeana
E. europaea
E. hamiltoniana ssp. maackii
E. verrucosa
Days
Avg germ
capacity
(%)
Purity
(%)
Soundness (%)
Samples
21.1
30
10
25
21.1
20
0
20
14
61
60
60
15
40
20
71
75
75
88
96
1
2
1
22+
20
15
60
75
3+
20
12
60
70
75
96
7+
References
Bailey LH. 1939. The standard cyclopedia of horticulture. New York:
Macmillan 3639 p.
Curtis JT. 1959. The vegetation of Wisconsin: an ordination of plant communities. Madison: University of Wisconsin Press. 657 p.
Dawidowicz-Grzegorzewska A, Beranger-Novat N. 1989. Ultrastructural
studies on development in mature seeds: embryos of Euonymus europea
L. dormant or cultured at +25 C. Journal of Experimental Botany 40:
913918.
Dirr MA. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation, and uses. Champaign, IL:
Stipes Publishing Co. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seeds to tissue culture. Athens, GA: Varsity Press. 239 p.
Fernald ML. 1950. Grays manual of botany. New York: American Book Co.
1632 p.
Foster S, Duke JA. 1990. A field guide to medicinal plants: eastern and central North America. New York: Houghton Mifflin. 366 p.
Gleason HA, Cronquist A. 1963. Manual of vascular plants of northeastern
United States and adjacent Canada. Boston: Willard Grant Press. 810 p.
Gorshenin NM. 1941. Agrolesomelioratsiya [Agro-forest melioration].
Moscow. 392 p.
Heit CE. 1966. Propagation from seed: 2.Testing extremely dormant seeds.
American Nurseryman 124(10): 10, 11, 38, 40.
Heit CE. 1967. Propagation from seed: 11. Storage of deciduous tree and
shrub seeds. American Nurseryman 126(10): 1213, 8694.
Heit CE. 1968a. Propagation from seed: 15. Fall planting of shrub seeds for
successful seedling production. American Nurseryman 128(4): 810,
7080.
Heit CE. 1968b. Thirty-five years testing of tree and shrub seeds. Journal of
Forestry 66(8): 632634.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing: rules 1993. Seed Science and Technology (suppl.): 1259.
Krssman G. 1960. Handbuch der Laubgehlze.Volume 1 and Volume 2.
495 & 608 p.
Lancaster R. 1981. An account of Euonymus in cultivation and its availability
in cultivation. Plantsman 3: 133166.
Lee WG, Grubb PJ, Wilson JB. 1991. Patterns of resource allocation in
fleshy fruits of nine European tall shrubs. Oikos 61: 307315.
Myatt A, Huffman G, Odell J. 1991. Seed processing manual. Stillwater:
Oklahoma Department of Agriculture, Forestry Division, Forest
Restoration Center. 1 p.
NBV [Nederlandsche Boschbouw Vereeniging]. 1946. Boomzaden: handleiding inzake heto ogsten, behandelen, bewaren en uitzaaien van
boomzaden. Wageningen, the Netherlands: Ponsen and Looijen. 171 p.
Nielsen KK. 1988. Dormancy in seeds from different positions on individual
plants. Acta Horticulturae 226(I): 255261.
Nielsen KK, Ericksen EN. 1988. Dry weight and moisture content in seeds
from different positions in individual plants. Acta Horticulturae 226(II):
525528.
518
Euonymus
519
FagaceaeBeech family
Fagus L.
beech
Franklin T. Bonner and William B. Leak
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station, Mississippi State
Mississippi; Dr. Leak is a silviculturist at the USDA Forest Services Northeastern Research Station
Durham, New Hampshire
Growth habit, occurrence, and use. The beeches
the genus Fagusincludes 10 species of medium-sized,
deciduous trees native to the temperate regions of the
Northern Hemisphere (Rehder 1940). Only 1 species,
American beech, is native to North America, although
another, the European beech, has been widely planted as an
ornamental in the Northeast (table 1). Some authorities have
argued that there are separate northern and southern species
of American beech, but this view is not widely supported
(Tubbs and Houston 1990). Beeches that grow in northeastern Mexico are now classified as a variety of American
beechF. grandifolia var. mexicana (Martinez) (Little
1965). There is some evidence of geographic races of
European beech in that species native range (Rudolf and
Leak 1974). Beech wood is used for flooring, furniture,
veneer, plywood, ties, charcoal, and many specialty products. The trees are highly valued for ornamental plantings,
and the mast is widely utilized by numerous birds and animals (Tubbs and Houston 1990).
Flowering and fruiting. Beech flowers are monoecious. The minute male and female flowers appear in the
spring when the leaves are about one-third grown (table 2).
The staminate flowers occur in densely clustered, drooping
heads 8 mm wide, whereas the pistillate flowers are generally paired on stout stalks about 2.5 cm long (Brown and
Kirkman 1990). Flowers of European beech are quite vulnerable to late spring frosts (Matthews 1955). The fruit is a
prickly bur approximately 2 cm long, which opens soon
after maturity in the fall (figure 1).
Each fruit contains 2 or 3 yellowish-brown or chestnutbrown, unevenly triangular nuts, 1 to 1.5 cm long (figures 2
and 3). Times of flowering, fruiting, and seed dispersal for
the 2 species are listed in table 2. Natural seed dispersal is
chiefly by gravity and by animals such as rodents and blue
jays (Cyanocitta cristata) (Johnson and Adkisson 1985;
Tubbs and Houston 1990). Information on height at maturity, minimum seed-bearing age, and interval between good
seedcrops is shown in table 3.
Common name
Occurrence
F. grandifolia Ehrh.
F. sylvatica L.
Flowering
Fruit ripening
Seed dispersal
F. grandifolia
F. sylvatica*
MarchMay
AprMay
SeptNov
SeptOct
Sources: Brown and Kirkman (1990), Rudolf and Leak (1974),Tubbs and Houston (1990).
* Dates are similar for western Europe and the northeastern United States.
520
nuts
nut.
longitudinal
Fagus
521
Species
2137
2030
F. grandifolia
F. sylvatica
Minimum
seed-bearing
age (yrs)
Year first
cultivated
1800
Long ago
40
4080*
Years between
large seedcrops
Time
Location
23
45
58
912
310
1520
Wisconsin
Mtn areas
Great Britain
Species
F. grandifolia
F. sylvatica
Fresh fruits
Air-dried fruits
Fruit wt/vol
kg/hl
lb/bu
Range
/kg
Cleaned seeds/weight
Average
/lb
/kg
/lb
Samples
12
2,8505,110
1,2902,320
3,500
1,600
10
5053
3945
3941
3035
4,0006,200
1,8002,800
4,630
2,100
24+
Bonnet-Masimbert 1982; Suszka 1974). Poulsen (1993) recommends that drying should be done at temperatures below
20 C. This behavior would seem to put beeches into the
orthodox class of storage behavior, although there is evidence that beeches fit somewhere between the orthodox and
recalcitrant classes (Gosling 1991) or in the sub-orthodox
class (Bonner 1990). The high lipid content of 40.7% reported for kernels of European beech (Prasad and Gulz 1989)
would seem to support the sub-orthodox classification. The
seeds are basically orthodox, however, and 5 years of storage is long enough for operational storage. There are no
comparable data for American beech, but there are no reasons to suspect that this species cannot be treated in the
same way. Beech seeds require cold stratification
(prechilling) for prompt germination, and current practices
with European beech have combined stratification and storage into a coordinated procedure. The first step is to determine how much stratification is needed to overcome dormancy (Suszka and Zieta 1977). Samples of fresh seeds are
brought to maximum moisture content or mixed with moist
sand and stored at 3 C until about 10% of the seeds have
started germination (radicles are visible). This period is
assumed to be the amount of time required to overcome dormancy in that particular lot. The remainder of the seeds are
522
Species
Test conditions
Temp (C)
Day
Night
Germination rate
Amount Period
(%)
(days)
Medium
90
42
Sand
Sand, paper
30
30
20
20
8447
85
81
140
150
Sand + peat
Sand + peat
1
5
1
5
56120
60110
100
100
F. grandifolia
F. sylvatica
F. sylvatica
Fresh seeds
Stored seeds
Germination
(%)
Fagus
523
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524
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RosaceaeRose family
achene with
Fagus
525
achene with
longitudinal
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Fallugia
527
RutaceaeRue family
528
seed.
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Commonwealth of Australia Forestry and Timber Bureau. 469 p.
Little EL Jr, Skolmen RG. 1989. Common forest trees of Hawaii. Agric.
Handbk. 679. Washington, DC: USDA Forest Service. 321 p.
Neil MC. 1965. In gardens in Hawaii. Spec. Pub. 50. Honolulu: Bishop
Museum Press. 924 p.
Swain EHF. 1928. The timbers and forest products of Queensland. Brisbane,
Australia: Queensland Forest Service. 500 p.
Wick HL. 1974. Flindersia, Queensland-maple In: Schopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 409410.
Flindersia
529
RhamnaceaeBuckthorn family
Frangula P. Mill.
buckthorn
Andrew Youngblood
Dr.Youngblood is a research forester at the USDA Forest Services Pacific Northwest Research Station
La Grande, Oregon
Growth habit, occurrence, and use. The buckthorn
genusFrangulaand the closely related genus Rhamnus
have until recently been treated as a single genus (Rhamnus)
consisting of more than 125 species of evergreen or deciduous shrubs and trees with alternate branches and simple
leaves with prominent pinnate veins (Hickman 1993).
Kartesz and Gandhi (1994), however, used floral morphology and leaf venation, as well as anatomical features of
xylem vessels, to support segregation of Frangula. Under
their treatment, Frangula species lack winter bud scales, the
pinnate leaf nerves are almost straight rather than arcuate,
and thorns are absent. Both Rhamnus and Frangula are
native to the temperate region of North America, Europe,
and Asia and also occur in the Neotropics and southern
Africa as shrubs and trees up to 1.5 m dbh and over 60 m
tall (Johnston and Johnston 1978; Krssmann 1985). The
common name, buckthorn, is probably misapplied and is
based on European species of Rhamnus that are thorny
(Mozingo 1987; USDA 1937). At least 16 species and subspecies are distributed within the United States (table 1)
(USDA NRCS 2001).
Glossy buckthorn, which is native to Europe, North
Africa, and western Europe, also is naturalized in northeastern and central United States and southern Canada, where it
grows to a height of 6 m and is often used for hedges. The
fruits are eaten by American robins (Turdus migratorius),
Bohemian waxwings (Bombycilla garrulus), cedar
waxwings (B. cedrorum), rose-breasted grosbeaks
(Phencticus ludovicianus), and starlings (Sturnus vulgaris).
Dispersal of seeds by birds and subsequent germination and
establishment represents a rapidly increasing problem; for
example, this non-native invasive shrub has replaced natural
open and semi-open wetland communities in southern
Ontario (Catling and Porebski 1994).
Beechleaf buckthorn is a low-growing shrub with dark
green leaves found in rock crevices, hanging gardens, and
desert shrub communities in the Southwest (Welsh and others 1990).
530
Common name(s)
Occurrence
F. alnus P. Mill.
Rhamnus frangula L.
R. frangula L. var. angustifolia Loud.
glossy buckthorn
beechleaf buckthorn,
birchleaf buckthorn
obovate buckthorn
California buckthorn
California buckthorn
California
California buckthorn
California
California buckthorn
California buckthorn
California
California buckthorn
531
Common name(s)
Occurrence
Carolina buckthorn,
yellow buckthorn, yellowwood
Nevada buckthorn
Nevada
obtuse buckthorn
red buckthorn,
Sierra buckthorn, coffeeberry
Yosemite buckthorn
fruit.
California
cool-white fluorescent light followed by 14 hours of darkness at 20 C (Radwan 1976). Dormant seeds responded
favorable to applications of 500 ppm of potassium gibberellate (K-GA3) when light was available during germination
and may represent a practical alternative to artificial cold
stratification for breaking dormancy (Radwan 1976). Clean
seeds of glossy buckthorn have been treated with sulfuric
acid (H2SO4) for 20 minutes to break dormancy; the acid
treatment should be done carefully because soaking the
seeds of other buckthorns was harmful (Hubbard 1974).
There are no officially prescribed germination tests procedures for buckthorns. Viability tests by tetrazolium staining have been suggested for European species (Enescu
1991). Seeds should be soaked in water for 24 hours,
cracked open in a vise, then re-soaked overnight. Staining
should take place in a 1% tetrazolium solution for 24 hours
at 30 C (Dirr 1990). To be considered viable, the embryos
must be completely stained, with the exception of the
extreme third of the distal ends of the radicle and
cotyledons.
Nursery and field practice. Detailed nursery techniques have not been developed for most Frangula species.
ments (Hubbard 1974; Keeley 1981, 1987). During laborato- The available information suggests that for most of the
ry tests involving 1 month of stratification at 5 C, however, species, the seeds should be sown in the spring at a depth of
10 to 40 mm (0.4 to 1.6 in) after they have been treated to
more than 75% of the total germination occurred after 7
break dormancy (Hubbard 1974). In contrast, cascara seeds
days of incubation at 23 C in the dark. Germination
increased to 90% when seeds were incubated with an initial may germinate faster and produce more vigorous plants
when seeds are sown at a depth of 3 mm (0.1 in) (Radwan
heat treatment of 100 C for 5 minutes and then placed on
1976). Germination is epigeal, with thick, straight cotylesoil containing 0.5 g powdered charred wood (charate) of
dons (Kartesz and Gandhi 1994). Cascara has also been
the chaparral shrub chamiseAdenostoma fasciculatum
Hook. & Arn. This treatment is designed to simulate condi- propagated by cuttings, and glossy buckthorn by grafting
(Hubbard 1974).
tions after a chaparral fire (Keeley 1987). Seeds of cascara
germinated best when stratified in the dark for 112 days at 5
C, then incubated for 28 days at 30 C for 10 hours under
long-
Frangula
533
References
Arno SF, Hammerly RP. 1977. Northwest trees. Seattle, WA:The
Mountaineers. 222 p.
Catling PM, Porebski ZS. 1994. The history of invasion and current status of
glossy buckthorn, Rhamnus frangula, in southern Ontario. Canadian FieldNaturalist 108: 305310.
Conrad CE. 1987. Common shrubs of chaparral and associated ecosystems
of southern California. Gen.Tech. Rep. PSW-99. Berkeley, CA: USDA,
Forest Service, Pacific Southwest Forest and Range Experiment Station.
86 p.
Conrad CE. 1996. Personal communication. Honolulu: Institute of Pacific
Islands Forestry.
Dirr MA. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation and use. Champaign, IL: Stipes
Publishing Co. 1007 p.
Enescu V. 1991. The tetrazolium test of viability. In: Gordon AG, Gosling P,
Wang BSP, eds.Tree and shrub seed handbook. Zurich: International
Seed Testing Association: 9.19.19.
Heiser CB, Jr. 1993. Ethnobotany and economic botany. In: Flora of North
America north of Mexico.Volume 1, Introduction. New York: Oxford
University Press: 199206.
Hickman JC, ed. 1993. The Jepson manual: higher plants of California.
Berkeley: University of California Press. 1400 p.
Hubbard RL. 1974. Rhamnus, buckthorn. In: Schopmeyer CS, tech. coord.
Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 704708.
Johnston MC, Johnston LA. 1978. Flora Neotropica. Monograph 20. New
York: New York Botanical Garden. 96 p.
Kartesz, JT, Gandhi, KN. 1994. Nomenclatural notes for the North
American Flora. XIII. Phytologia 76: 441457.
Keeley JE. 1981. Reproductive cycles and fire regimes. In: Mooney HA,
Bonnicksen TM, Christensen NL [and others], tech. coords.: Proceedings,
Fire Regimes and Ecosystem Properties Conference; 1978 December
1115; Honolulu, HI. Gen.Tech. Rep. WO-26. Washington, DC: USDA
Forest Service: 231277.
534
TheaceaeTea family
capsules
Franklinia
535
seeds.
longitudinal
References
Regardless of temperature, germination in the dark was negligible for nonstratified seeds. However, in the presence of
light, cumulative germination at 16/7 C, 24/16 C, or
29/24 C was 2, 75, and 61%, respectively. Stratification
enhanced germination by accelerating the rate of germination and reducing sensitivity of the seeds to light. After 4
weeks of stratification, total germination in the presence of
light at 16/7 C, 24/16 C, and 29/24 C was 5, 87, and
91%, respectively, in comparison to 2, 31, and 85%, respectively, for seeds in darkness. Germination following stratification for 8 weeks was similar to that of 4 weeks of stratification. Additional stratification for 12 weeks resulted in an
increase in dark germination at 24/16 C to 53% and a large
increase in germination at 16/7 C with dark and light germination of 32 and 52%, respectively.
536
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Elias TS. 1989. Field guide to North American trees. Danbury, CT: Grolier
Book Clubs. 948 p.
Everett TH. 1981. The New York Botanical Garden illustrated encyclopedia
of horticulture. New York: Garland Publishing. 3601 p.
Farmer Jr. RE. Chase SB. 1977. Germination and container production of
Franklinia. HortScience 12(1): 43.
Hartmann HT, Kester DE, Davies Jr FT, Geneve RL. 1997. Plant propagation:
principles and practices. 6th ed. Upper Saddle River, NJ: Prentice Hall.
770 p.
Jacobson AL. 1996. North American landscape trees. Berkeley:Ten Speed
Press. 722 p.
Judd WH. 1930. The fruiting of Franklinia. Horticulture 8(5): 103.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. 3rd ed.
New York: Macmillan. 1290 p.
Sargent CS. 1949. Manual of the trees of North America. New York: Dover
Publications. 910 p.
Schneider R. 1988. Franklinia alatamaha. American Nurseryman 167(2):
146.
Small JK. 1933. Manual of the southeasern flora. Chapel Hill: University of
North Carolina Press. 1554 p.
Wildman A. 1996. Franklinia alatamaha: Franklin tree. Arbor Age 16(6): 20.
OleaceaeOlive family
Fraxinus L.
ash
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Common name(s)
Occurrence
Fraxinus
537
cluster of
Table 2Fraxinus, ash: flowering habit and phenology of flowering and fruiting
Species
Location
Flowering
Flowering habit
Fruit ripening
Seed dispersal
F. americana
F. caroliniana
F. dipetala
F. excelsior
F. latifolia
F. nigra
F. ornus
F. pennsylvanica
F. profunda
F. quadrangulata
F. uhdei
California
NE US
Hawaii
Puerto Rico
AprMay
FebMar
AprMay
AprMay
AprMay
MayJune
MayJune
MarMay
AprMay
MarApr
MarMay
NovJan
MarApr
Dioecious
Dioecious
Perfect
Polygamous
Dioecious
Polygamous
Polygamous
Dioecious
Dioecious
Perfect
Dioecious
Dioecious
OctNov
AugOct
JulySept
AugSept
AugSept
JuneSept
SeptOct
SeptOct
JuneOct
JulySept
Aug
Sept
SeptDec
Winterearly spring
SeptOct
JulyOct
Octspring
OctDec
JulySept
F. velutina
Sources: Bonner (1974), Francis (1990), Harms (1990), Kennedy (1990), Owston (1990), Rehder (1940), Schlesinger (1990),Vines (1960),Wright and Rauscher (1990).
538
longitudinal
Species
F. americana
F. caroliniana
F. dipetala
F. excelsior
F. latifolia
F. nigra
F. ornus
F. pennsylvanica
F. profunda
F. quadrangulata
F. uhdei
F. velutina
Height at
maturity (m)
2124
612
26
2938
1824
1224
620
21+
37
49
37
15
Year first
cultivated
1724
Long ago*
1800
pre-1700
1824
1823
1900
1900
Minimum
seed-bearing
age (yr)
Years
between large
seedcrops
20
15
30
20
9
25
15
35
12
35
34
1
but they must be completely dry for the process to be successful. Smaller seedlots, such as those used for research or
testing, can be de-winged in laboratory blenders operated at
low speeds about half-full of water. Seed yield data for
ashes are summarized in table 4.
Long-term storage studies with seeds of the ashes are
few, but these seeds are definitely orthodox in their storage
characteristics. Studies by Barton (1945) showed no loss in
viability for 7 years for green and European ash seeds stored
in sealed containers at 5 C with seed moisture contents of 7
to 10%. Similar conditions have proved successful for flowering ash (Heit 1967) and Shamel ash (Bonner 1974).
Fraxinus
539
540
10
F. profunda
F. quadrangulata
F. uhdei
F. velutina
24,25054,250
35,06074,150
6,8307,270
13,00015,430
34,20038,600
28,85061,740
7.2
F. latifolia
F. nigra
F. pennsylvanica
F. caroliniana
F. dipetala
F. excelsior
51
9
5
2+
10
6
17,260
20,950
3,200
16,500
20,600
38,850
46,200
7,050
36,380
45,420
11,00024,000
15,90035,630
3,0003,300
5,9007,000
15,50017,500
13,00028,000
11
13
78
7
2
1
3+
10+
4
13,120
8,470
5,744
5,900
8,100
28,930
18,680
13,660
13,000
17,860
5,54018,185
5,71211,288
5,0008,800
4,7507,000
4,0007,000
10,00014,060
6,1009,500
12,22040,100
12,60024,900
11,02519,400
10,47015,430
8,80015,430
22,00031,000
13,45020,950
12.4
12.5
14
16*
16*
18
Midwest
Mississippi
Arkansas
California
Europe
USA
Hawaii
F. americana
Species
Place fruit
collected
Seeds/vol
kg/hl
lb/bu
/kg
Range
Cleaned seeds/weight
Average
/lb
/kg
/lb
Samples
Seed
moisture (%)
Medium
F. americana
Sand
Plastic bag*
Plastic bag*
Sand, peat
Sand
Sand, peat, or plastic bag*
Sand
Peat
Soil
Moist substrate
Plastic bag*
Moist paper
Sand
Sand, soil
F. caroliniana
F. dipetala
F. excelsior
F. nigra
F. ornus
F. pennsylvanica
F. profunda
F. quadrangulata
F. uhdei
F. velutina
Warm period
Temp (C)
Days
2030
20
2030
21
Warm
20
2030
30
6090
60
126
30
60
60
0
Cold period
Temp (C)
Days
5
3
3
25
Cool
45
5
4
Cold
05
25
5
5
25
60
5684
60
90
480540
6090
90
90
90
210
60150
60
90
0
90
Sources: Bonner (1974), Bonner (1975), Francis (1990), Mirov and Kraebel (1939), Soljanik (1961), Steinbauer (1937),Vanstone and LaCroix (1975),Walle (1987).
* Naked stratification in plastic bags.
In outdoor pits.
Exact temperatures not given.
For seeds from southern sources, 2 or 3 months is enough, but for seeds from northern sources, 5 months is needed.The warm period is helpful but not essential for
southern sources (Bonner 1974; Eliason 1965).
Fraxinus
541
Table 6Fraxinus, ash: germination test conditions and results for stratified seed
Germination test conditions
Species
F. americana
F. caroliniana
F. dipetala
F. excelsior
F. nigra
F. ornus
F. pennsylvanica
F. profunda
F. quadrangulata
F. uhdei
F. velutina
Daily
light
period
(hrs)
8
8
8
16
NDL
NDL
Medium
Sand
Paper
Kimpak
Sand
Soil
Paper
Paper
Sand
Soil
Sand
Kimpak
Sand
Temp (C)
Day
Night
30
25
30
30
30
30
30
30
30
30
20
15
20
20
20
20
16
20
20
20
Days
Germination
rate
Amt
(%)
Days
2440
56
60
40
3034
42
30
45
56
40
49
54
70
71
32
43
66
24
14
18
20
21
20
40
21
Germination
percentage
Avg
(%)
Samples
54
68
61
71
61
20
49
76
80
89
48
44
69
33
3
1
3
2
4
6
3
6
3
3
1
1
4
5
Sources: Bonner (1974), Bonner (1975), Cram (1984), Mirov and Kraebel (1939), Soljanik (1961),Tinus (1982).
NDL = Natural daylength in a greenhouse.
542
References
Aldhous JR. 1972. Nursery practice. Bull. 43. London: Forestry Commission.
194 p.
Arrillaga I, Marzo T, Segura J. 1992. Embryo culture of Fraxinus ornus and
Sorbus domestica removes seed dormancy. Hortscience 27: 371.
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
Barger JH, Davidson RH. 1967. A life history of the ash seed weevils,
Thysanocnemis bischoffi Blatchley and T. helvola Leconte. Ohio Journal of
Science 67(2): 123127.
Barton LV. 1945. Viability of seeds of Fraxinus after storage. Contributions of
the Boyce Thompson Institute 13: 427432.
Bonner FT. 1963. Some southern hardwoods can be air-layered. Journal of
Forestry 61: 923.
Bonner FT. 1973. Timing collections of samaras of Fraxinus pennsylvanica
Marsh. in the southern United States. In: International Symposium on
Seed Processing.Volume 1, Seed processing (IUFRO W.P. S2.01.06). 1973
September; Bergen, Norway. Stockholm: Royal College of Forestry: 7 p.
Bonner FT. 1974. Fraxinus L., ash. In: Schopmeyer CS, tech. coord. Seeds of
woody plants in the United States. Agric. Handbk. 450. Washington, DC:
USDA Forest Service: 411416.
Bonner FT. 1975. Germination temperatures and prechill treatments for
white ash (Fraxinus americana L.). Proceedings of the Association of
Official Seed Analysts 65: 6065.
Brown CL, Kirkman LK. 1990. Trees of Georgia and adjacent states.
Portland, OR:Timber Press. 292 p.
Clausen KE. 1984. Survival and early growth of white ash provenances and
progenies in 19 plantations. Canadian Journal of Forest Research 14:
775782.
Clausen KE, Kung FH, Bey CF, Daniels RA. 1981. Variation in white ash.
Silvae Genetica 30(2/3): 9397.
Cram WH. 1984. Presowing treatments and storage for green ash seeds.
Tree Planters Notes 35(1): 2021.
Cram WH, Lindquist CH. 1982. Germination of green ash is related to seed
moisture content at harvest. Forest Science 28: 809812.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation. Athens, GA:Varsity Press. 239 p.
Eliason EJ. 1965. Treatment of forest tree seed to overcome dormancy
prior to direct seeding. In: Direct Seeding in the Northeast: A
Symposium. Amherst: University of Massachusetts Experiment Station:
8791.
Francis JK. 1990. Fraxinus uhdei (Wenzig) Lingelsh. Fresno, tropical ash. In:
Silvics manual. SO-ITF-SM-28. New Orleans: USDA Forest Service,
Southern Forest Experiment Station. 4 p.
Gendel SM, Fosket DE, Miksche JP. 1977. Increasing white ash seed germination by embryo dissection. Res. Note NC-220. St. Paul, MN: USDA
Forest Service, North Central Forest Experiment Station. 3 p.
Harms WR. 1990. Fraxinus profunda (Bush) Bush, pumpkin ash. In: Burns RM,
Honkala BH, tech. coords. Silvics of North America.Volume 2,
Hardwoods. Agric. Handbk. 654. Washington, DC: USDA Forest Service:
355357.
Heit CE. 1967. Propagation from seed: 11. Storage of deciduous tree and
shrub seeds. American Nurseryman 126(10): 1213, 8694.
Hendrix KW, Lowe WJ. 1990. Geographic variation of green ash in the
Western Gulf region. Silvae Genetica 39(3/4): 95103.
Houston DB. 1976. Determining the quality of white ash seed lots by x-ray
analysis.Tree Planters Notes 27(2): 8, 23.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing: Rules 1993. Seed Science & Technology 21 (Suppl.): 1259.
Karrfalt RP. 1992. Increasing hardwood seed quality with brush machines.
Tree Planters Notes 43(2): 3335.
Kennedy HE Jr. 1990. Fraxinus pennsylvanica Marsh., green ash. In: Burns RM,
Honkala BH, tech. coords. Silvics of North America.Volume 2,
Hardwoods. Agric. Handbk. 654. Washington, DC: USDA Forest Service:
348354.
Kentzer T. 1966. Gibberellin-like substances and growth inhibitors in relation
to the dormancy and after-ripening of ash seeds (Fraxinus excelsior L.).
Acta Societatis Botanicorum Poloniae 35(4): 575585.
Krinard RM. 1989. Stand parameters of 11- to 15-year old green ash plantings. Res. Note SO-352. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 4 p.
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Marshall PE. 1981. Methods for stimulating green ash seed germination.Tree
Planters Notes 32(3): 911.
McBride JR, Dickson R. 1972. Gibberellic, citric acids and stratification
enhance white ash germination.Tree Planters Notes 23(3): 12.
Mirov NT, Kraebel CJ. 1939. Collecting and handling seeds of wild plants.
Fraxinus
543
SterculiaceaeSterculia family
Fremontodendron Coville
fremontia, flannelbush
Susan E. Meyer
Dr. Meyer is a research ecologist at the USDA Forest Service, Rocky Mountain Research Stations
Shrub Sciences Laboratory, Provo, Utah
Common name(s)
Distribution
544
after emergence. Seed production is reportedly better in cultivated than in naturally occurring individuals (Nord 1974).
The ripened seed may be retained in the capsule for up to a
month, but it is best to collect seeds when the first capsules
begin to split open (Nord 1974). Capsules are collected by
hand stripping or beating into containers. Gloves are recommended to protect hands against the irritating capsule bristles. Capsules that do not open soon after collection should
be soaked in water for a few minutes, then dried before
extraction. Capsules may be broken up in a hammermill or
other threshing device, and the seeds cleaned out by screening and fanning (Nord 1974). Seed weight varies among and
within species (table 2). Fremontia species form persistent
seed banks in the field and are probably long-lived in storage (orthodox). In field seed bank experiments with eldorado fremontia, there was little loss of viability over a 7-year
period (Boyd and Serafini 1992).
Species
/kg
F. californicum
30,87055,125
Maximum
/lb
Fill %
germination %
14,00025,000
53
50
F. decumbens
26,460
12,000
100
72
F. mexicanum
44,10066,150
20,00030,000
100
55
Sources:
Fremontodendron
545
References
Boyd RS. 1994. Pollination biology of the rare shrub Fremontodendron
decumbens (Sterculiaceae). Madroo 41: 277289.
Boyd RS. 1996. Ant-mediated seed dispersal in the rare chaparral shrub
Fremontodendron decumbens (Sterculiaceae). Madroo 43: 299315.
Boyd RS, Serafini LL. 1992. Reproductive attrition in the rare chaparral
shrub Fremontodendron decumbens Lloyd (Sterculiaceae). American
Journal of Botany 79: 12641272.
Emery DE. 1988. Seed propagation of native California plants. Santa
Barbara: Santa Barbara Botanic Garden. 107 p.
Holmes R, ed. 1993. Taylors guide to natural gardening. Boston: Houghton
Mifflin: 158, 338.
546
GarryaceaeSilktassel family
Garrya
547
Common name
Occurrence
Elevation (m)
Growth form
G. buxifolia Gray
dwarf silktassel
Brushy shrub
G. elliptica Dougl.
ex Lindl.
wavyleaf silktassel
G. flavescens S.Wats.
ashy silktassel
G. fremontii Torr.
bearbrush
G. glaberrima
Wangerin
G. grisea Wiggins
G. laurifolia Benth.
laurel-leaf silktassel
G. longifolia Rose
G. ovata Benth.
eggleaf silktassel
G. salicifolia
Eastwood
G. veatchii Kellog
canyon silktassel
2302,600
Shrub
G. wrightii Torr.
Wright silktassel
9142,133
Shrub
Shrub (< 6 m)
Shrub (< 6 m)
Shrub
Small tree
Shrub (24.6 m)
Tree (< 11 m)
Small tree
Clumped shrub
(24.6 m)
Small tree
548
berry (left)
longitudinal sec-
References
Dahling GV. 1978. Systematics and evolution of Garrya. Contributions from
the Gray Herbarium of Harvard University 209: 1104.
Kartesz JT. 1994. A synonymized checklist of the vascular flora of the
United States, Canada, and Greenland. 2nd ed. Portland, OR:Timber
Press. 813 p.
Moerman DE. 1986. Medicinal plants of native America. Ann Arbor:
University of Michigan, Museum of Anthropology. 910 p.
Mirov NT, Kraebel CJ. 1937. Collecting and propagating seeds of California
wild plants. Res. Note 18. Berkeley, CA: USDA Forest Service, Pacific
Southwest Forest and Range Experiment Station. 27 p.
Mulligan BO, Nelson EC. 1980. Garrya issaquahensis Nelson (G. elliptica
Lindl. G. fremontii Torr.) in cultivation in western USA and Ireland. UW
Arboretum Bulletin 43(3): 1015.
Rehder A. 1940. Manual of cultivated trees and shrubs. 2nd ed. New York:
Macmillan. 996 p.
Reynolds HG, Alexander RR. 1974. Garrya, silktassel. In: Schopmeyer CS,
tech. coord. Seeds of woody plants in the United States. Agric. Handbk.
450. Washington, DC: USDA Forest Service: 420421.
Ridgeway D. 1973. Production of Garrya elliptica. Combined Proceedings of
the International Plant Propagators Society, 23: 177180.
Ridgeway D. 1985. Propagation and production of Garrya elliptica.
Combined Proceedings of the International Plant Propagators Society,
34: 261265.
Roth WB, Carr ME, Davis EA, Bagby MO. 1985. New sources of guttapercha in Garrya flavescens and G. wrightii. Phytochemistry 24(1):
183184.
Garrya
549
EricaceaeHeath family
Gaultheria L.
wintergreen
David W. Huffman, John C. Zasada, and William I. Stein
Dr. Huffman is a research associate at Northern Arizona State Universitys Ecological Restoration Institute,
Flagstaff, Arizona; Dr. Zasada retired from the USDA Forest Services
North Central Research Station; Dr. Stein is a forest ecologist emeritus at the USDA Forest Services
Pacific Northwest Research Station, Corvallis, Oregon
Growth habit, occurrence, and uses. Of the 100 to
150 species of the genus Gaultheria, commonly called wintergreen, most are found in Asia, Australia, and South
America. Only 6 speciescreeping snowberry, alpine wintergreen (G. humifusa (Graham.) Rydb.), G. miqueliana
Takeda, G. ovatifolia A. Gray, checkerberry, and salal
occur in North America north of Mexico (Abrams 1951;
Chou 1952; Hitchcock and others 1959; Viereck and Little
1972). The 3 species considered here (table 1) are evergreen
shrubs. Both creeping snowberry and checkerberry have a
prostrate or creeping form (Fernald 1950) and have been
described as semi-herbaceous or almost herbaceous
(Fernald 1950; Rosendahl 1955). Salal has a distinctly
woody stem and grows 1 to 3 m tall.
Over its wide range in the United States and Canada,
creeping snowberry is most common in bogs and wet
forested conditions (Curtis 1959; Gleason 1952;
MacKinnon and others 1992). Checkerberry tolerates site
conditions ranging from dry to poorly drained and grows
well on many acidic soil types, including peat, sand, sandy
loam, and coal mine spoils (Robinette 1974). Salal also
grows on a variety of sites, including shallow rocky soils,
sand dunes, glacial till, and peat (Haeussler and Coates
1986). It is most common on well-drained slopes and ridges
Common name(s)
Occurrence
creeping snowberry,
creeping pearlberry,
moxieplum
G. procumbens L.
checkerberry, wintergreen,
mountain-tea
salal, Oregon wintergreen
G. shallon Pursh
Sources: Abrams (1951), Fernald (1950), Gleason (1952), Hitchcock and others (1959).
550
racemes bearing
Gaultheria
551
Flowering
G. hispidula
G. procumbens
G. shallon
AprAug
MaySept
MarJuly
AugOct*
AugJune
JulyDec
Sources: Fernald (1950), Gleason (1952), Hitchcock and others (1959), McMinn (1951), Robinette (1974), Rosendahl (1955),Van Dersal (1938).
* Actual ripening time uncertain.
Fruit of this species notably persistent and reportedly increase slightly in size during winter (Van Dersal 1938).
Figure 3Gaultheria shallon, salal: the ripe fruit is a purplish black, somewhat-fuzzy pseudo-berry.
552
Table 3Gaultheria, wintergreen: growth form, height at maturity, and fruit characteristics
Species
Growth habit
G. hispidula
G. procumbens
G. shallon
Prostrate*
Creeping
Tall shrub
Height at
maturity (cm)
2040*
520
25300
Fruit diameter
(mm)
310
510
610
G
Color of ripe fruit
Clearwhite
Scarletbright red
Dark purple to bluish black
Sources: Fernald (1952), Gleason (1952), Hitchcock and others (1959), Rehder (1940), Rosendahl (1955), Stein (1995).
* Length.
/kg
6.756.89
6.3310.67
5.678.33
Average
/lb
/kg
/lb
3.063.13
2.874.84
2.573.78
6.82
8.50
7.14
3.09
3.86
3.24
longitu-
553
after germination), or under shade cloth (Huffman and others 1994b; McKeever 1938; Rogers 1994). Even without
such measures, up to 73% germination has been obtained in
soil-filled flats (Mirov and Kraebel 1939). Light for 8
hours/day is recommended. Some propagators expose seeds
to an additional 2 hours of light during the dark period
(Rogers 1994). A potted plant of checkerberry can be propagated from seeds in 4 months (Rogers 1994). Salal seedlings
raised in outdoor nursery beds grew 11 to 17 cm (4.3 to 6.7
in) tall in 2 years (Huffman and others 1994b). They exhibited poor apical dominance, however, developing 6 to 12 aerial stems. Light shade (70% light) produced seedlings with
greater biomass, greater canopy size, and more aerial stems
compared to those under 20 or 50% or full sun. Under all
light intensities, some seedlings produced rhizomes in 2
years.
All 3 species are readily propagated vegetatively from
layers, suckers, division of plants, stem or root cuttings,
stolons, or rooting at the nodes (Brown and Hafenrichter
1962; Dimock and others 1974; Huffman and others 1994b;
Rogers 1994; Sabhasri 1961; Van Dersal 1938). In the
Northwest, salal is presently propagated almost entirely by
rhizome cuttings (Dimock and others 1974). Cultured rhizome cuttings can produce 5 or more new rhizomes and
over 7 aerial shoots/year under light shade during the first 2
years after planting (Huffman and others 1994b). Moist,
acid conditions under partial shade are beneficial for young
plants of all 3 species raised from either cuttings or seed.
Species
G. hispidula
G. procumbens
G. shallon
Cold
stratification
(days*)
83
0
60
0
30120
0
0
0
0
60
60
60
150
150
30
30
30
30
21
21
16
10
21
16
10
21
10
20
20
20
20
21
16
10
4
16
10
4
16
4
Germination rate
Days
%
98
213
56
61
55
28
60
60
60
60
60
60
60
60
7
16
28
26
74
42
28
0
39
31
23
55
32
Total
germination
Days
%
59
15
27
37
22
30
30
30
30
30
30
30
30
8
16
38
31
76
51
51
41
45
40
50
59
40
Seeds of wintergreen appear to have little innate dormancy under field conditions. No evidence of delayed emergence of salal was observed in 2 replicated studies subsequent to sowing test plots (Huffman and others 1994a;
Tappeiner and Zasada 1993). Salal seedlings establish most
readily on rotten logs and stumps under partial shade
(Huffman and others 1994a; Huffman and Tappeiner 1997).
There is evidence that this is also true for checkerberry
(Matlack and Good 1989). Forest floor disturbance that
exposes mineral soil enhances survival of salal seedlings
(Huffman and others 1994a; Tappeiner and Zasada 1993).
Predation of seeds did not appear to be a significant factor in
a seedling establishment study in the Oregon coastal range
(Tappeiner and Zasada 1993). Under field conditions,
growth of salal seedlings is slow; they attained average
heights of 2 to 4 cm (.8 to 1.6 in) in 2 years but can grow to
20 cm (7.9 in) (Huffman and others 1994a). Seedlings begin
vegetative expansion in 4 to 6 years (Huffman and others
1994a). Young seedlings are susceptible to late spring frost
(Sabhasri 1961).
Most field regeneration of wntergreen is vegetative
(Bunnell 1990; Huffman and others 1994a; Huffman and
Tappeiner 1997; Matlack and Good 1990; Matlack and others 1993; Sabhasri 1961). Checkerberry expands by growth
of rhizomes and layering of creeping stems where conditions permit (Matlack and others 1993; Robinette 1974).
Maximum expansion rates can be 10 cm (3.9 in) per year or
more (Sobey and Barkhouse 1977). Clones of creeping
snowberry develop as a result of layering of prostrate stems;
maximum expansion rates range from 2 to 7 cm (.8 to 2.8
in) per year (Sobey and Barkhouse 1977). Rhizome expansion rates of 44 cm (17 in) per year have been reported for
salal; the maximum observed was 93 cm (37 in)/year
(Huffman and others 1994a). Individual clones of salal can
occupy areas up to 29 m2 (312 ft2)with up to 218 m (715 ft)
of interconnected rhizomes (figure 1) (Huffman and others
1994a). Salal populations rapidly recover after logging
(Halpern 1988; Messier 1992; Messier and Kimmins 1991;
Stein 1995) and can severely compete with commercially
important tree species (Price and others 1986; Weetman and
others 1990; Messier and Kimmins 1990). Clonal assemblages persist by vegetative regeneration of aerial shoots that
replace older, dying stems (Huffman and others 1994a).
Although shade-tolerant, salal loses vigor with increasing
overstory density and clones disintegrate into smaller fragments (Huffman and others 1994a). An estimated minimum
light requirement for salal survival ranges from 0.3 to 3.3%
of full sunlight (Messier and others 1989).
References
Abrams L. 1951. Illustrated flora of the Pacific States.Volume 3. Stanford,
CA: Stanford University Press. 866 p.
Brown RL, Hafenrichter AL. 1962. Stabilizing sand dunes on the Pacific
coast with woody plants. Misc. Pub. 892. Washington, DC: USDA Soil
Conservation Service. 18 p.
Bunnell FL. 1990. Reproduction of salal (Gaultheria shallon) under forest
canopy. Canadian Journal of Forest Research 20: 91100.
Chou YL. 1952. Floral morphology of three species of Gaultheria. Botanical
Gazette 114: 198221.
Curtis JT. 1959. The vegetation of Wisconsin. Madison: University of
Wisconsin Press. 657 p.
Dimock EJ II, Johnson WF, Stein WI. 1974. Gaultheria L., wintergreen. In:
Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service:
422426.
Dirr MA. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation and uses. Champaign, IL:
Stipes Publishing. 1007 p.
Douglass BS. 1970. Special forest products1969 harvesting report,
Oregon and Washington. Portland, OR: USDA Forest Service, Pacific
Northwest Region. 39 p.
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Company. 1632 p.
Fraser L,Turkington R, Chanway CP. 1993. The biology of Canadian weeds:
102. Gaultheria shallon Pursh. Canadian Journal of Plant Science 73:
12331247.
Gleason HA. 1952. Illustrated flora of the northeastern United States and
adjacent Canada.Volume 3. Lancaster, PA: Lancaster Press. 589 p.
Gunther E. 1945. Ethnobotany of western Washington. Seattle: University of
Washington Press. 61 p.
Haeussler S, Coates D. 1986. Autecological characteristics of selected
species that compete with conifers in British Columbia: a literature
review. Land Mgmt. Rep. 33.Victoria, BC: British Columbia Ministry of
Forests. 180 p.
Halpern CB. 1988. Early successional pathways and the resistance and
resilience of forest communities. Ecology 69: 17031715.
Gaultheria
555
556
EricaceaeHeath family
Gaylussacia
557
558
References
Bonner FT, Halls LK. 1974. Gaylussacia baccata (Wangh.) K. Koch, black
huckleberry. In: Schopmeyer CS, tech. coord. Seeds of woody plants in
the United States. Agric. Handbk. 450. Washington, DC: USDA Forest
Service: 427428.
Radford AE, Ahles HE, Bell CR. 1968. Manual of the vascular flora of the
Carolinas. Chapel Hill: University of North Carolina Press. 1183 p.
Van Dersal WR. 1938. Native woody plants on the United States: their
erosion-control and wildlife values. Misc. Pub. 303. Washington, DC:
USDA. 362 p.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
GinkgoaceaeGinkgo family
Ginkgo biloba L.
ginkgo
Wayne D. Shepperd
Dr. Shepperd is a research silviculturist at the USDA Forest Services Rocky Mountain Research Station,
Fort Collins, Colorado
Ginkgo
559
longitudinal section
References
AGINFO. 1994. Plant database [available at www.ags.udel.edu]. Newark,
DE: University of Delaware, College of Agricultural Sciences.
Alexander RR. 1974. Ginkgo biloba L., ginkgo. In: Schopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 429430.
Bailey LH. 1923. Cultivated evergreens. New York: Macmillan: 177178.
Bailey LH. 1947. Standard cyclopedia of horticulture. 2nd ed. New York:
Macmillan. 338 p.
Dallimore W, Jackson AB. 1948. Handbook on coniferae. 3rd ed. London:
Edward Arnold Co.: 229233
Davis SH, Henery JT. 1942. A Xylaria pathogenic to Ginkgo biloba (L.)
seeds. Phytopathology 32: 9192.
560
Porterfield W. 1940. Chinese vegetable foods in New York: 11. Seeds of the
Ginkgo. New York Botanical Garden Journal 41: 186188.
Rehder A. 1940. Manual of cultivated trees and shrubs. 2nd ed. New York:
Macmillan. 996 p.
Sakisaka M. 1927. On the seed bearing leaves of Ginkgo. Japanese Journal of
Botany 4: 219236.
Seward AC, Gowan J. 1900. The maidenhair tree (Ginkgo biloba L.). Annals
of Botany 14(53): 109164.
Swingle CF. 1939. Seed propagation of trees, shrubs, and forbs for conservation planting. SCS-TP-27. Washington, DC: USDA Soil Conservation
Service. 198 p
Willan RL. 1985. A guide to forest seed handling with special reference to
the tropics. For. Pap. 20/2. Rome: FAO.
Ginkgo
561
FabaceaePea family
Gleditsia L.
honeylocust
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Figure 1Gleditsia, honeylocust: legumes of G. triacanthos, honeylocust (top left), G. aquatica, water honeylocust
(bottom left), and G. texana,Texas honeylocust (right).
Table 1Gleditsia, honeylocust: nomenclature, occurrence, height at maturity, and year first cultivated
Height (m)
at maturity
Scientific name
Common name(s)
Occurrence
G. aquatica Marsh.
waterlocust,
swamp honeylocust.
G. texana Sarg.
Texas honeylocust,
Texas locust
honeylocust,
sweet-locust,
thorny-locust
G. triacanthos L.
562
Year first
cultivated
1218
1723
40
1900
2143
1700
The seeds are close to the same size (figure 2) and contain a thin, flat embryo surrounded by a layer of horny
endosperm (figure 3). Phenology of flowering and fruiting is
summarized in table 2. Seedbearing starts at about age 10,
with optimum production between 25 and 75 years (Blair
1990). Good crops are borne almost every year (Bonner and
others 1974).
Collection of fruits. Fruit color changes from green
to a deep reddish brown, or even brownish black at maturity
(Brown and Kirkman 1990; Gordon 1966). Legumes may be
picked from the trees after they dry or from the ground after
natural dissemination, which may last into late winter (Blair
1990). Collection from the ground should be completed
early to avoid losses to wildlife and to disintegration of the
legumes in late winter or spring. Moist legumes should be
spread for thorough drying before extraction. Tree shakers
have been used to collect honeylocust fruits in Russia, with
as much as 90 to 100% of the crop recovered (Kiktev and
others 1977).
Extraction and storage. Dried legumes may be run
through macerators or other mechanical threshers to extract
the seeds; hand flailing will also work. The Forest Service
macerator can extract 180 to 270 kg (400 to 600 lb) of clean
seeds per day (Bonner and others 1974). Small trash can be
removed with fans, air-screen cleaners, or water flotation,
which will also remove empty, insect-damaged, and incompletely developed seeds. Seeds can be separated from large
trash, such as legume fragments, with screens.
A seed yield of 44 to 77 kg/100 kg (20 to 35 lb/100 lb)
of honeylocust legumes and a purity of 95% and soundness
of 98% have been reported (Bonner and others 1974). In 36
seed samples of this species, there was an average of 6,100
seeds/kg (2,800/lb) with a range of 3,800 to 9,000 (1,750 to
4,050). Seeds of Texas honeylocust are generally larger, with
4,000 seeds/kg (1,830/lb) in 1 sample (Bonner and others
1974).
Seeds of honeylocust species are orthodox in storage
characteristics. Their viability can probably be maintained
for many years if seeds are stored at low temperatures with
moisture contents below 10%, although no long-term storage studies have been done. The food reserves in the seeds
are primarily carbohydrates and proteins (Felker and
Bandurski 1977; Mazzini and Cerezo 1979).
G. aquatica
waterlocust
G. X texana
Texas honeylocust
G. triacanthos
honeylocust
longitudinal section
Flowering
Fruit ripening
Seed dispersal
G. aquatica
G. texana
G. triacanthos
MayJune
AprMay
MayJune
AugOct
AugSept
SeptOct
SeptDec
SeptDec
Septlate winter
Gleditsia
563
References
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
Blair RM. 1990. Gleditsia triacanthos L., honeylocust. In: Burns RM, Honkala
BH, tech. coords. Silvics of North America.Volume 2, Hardwoods. Agric.
Handbk. 654. Washington, DC: USDA Forest Service: 358364.
Bonner FT, Burton JD, Grigsby HC. 1974. Gleditsia L., honeylocust. In:
Schopmeyer CS, tech. coord. Seeds of woody plants in the United States.
Agric. Handbk. 450. Washington, DC: USDA Forest Service: 431433.
Brown CL, Kirkman LK. 1990. Trees of Georgia and adjacent states.
Portland, OR:Timber Press. 292 p.
Davies DJG, Macfarlane RP. 1979. Multiple-purpose trees for pastoral farming in New Zealand: with emphasis on tree legumes. New Zealand
Agricultural Science 13(4): 177186 [Forestry Abstracts 42(1): 252;
1981].
Dirr MA, Heuser CW. 1987. The reference manual of woody plant propagation. Athens, GA:Varsity Press. 239 p.
Felker P, Bandurski RS. 1977. Protein and amino acid composition of tree
legume seeds. Journal of the Science of Food and Agriculture 28:
791797.
Felker P, Bandurski RS. 1979. Uses and potential uses of leguminous trees
for minimal energy input agriculture. Economic Botany 33(2): 172184.
Gordon D. 1966. A revision of the genus Gleditsia (Leguminosae) [PhD dissertation]. Bloomington: Indiana University. 114 p.
564
Heit CE. 1942. Acid treatment of honey locust. Notes For. Invest. 42.
Albany: New York Conservation Department. np.
Kiktev YN, Mitrofanov AS, Gaziev FM. 1977. [Machine for collecting seeds
from standing trees]. Lesnoe Khozyaistvo 7: 5657 [Forestry Abstracts
39(7): 2764; 1978].
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Liu NY, Khatamian H, Fretz TA. 1981. Seed coat structure of three woody
legume species after chemical and physical treatments to increase seed
germination. Journal of the American Society for Horticultural Science
106: 691694.
Mazzini MN, Cerezo AS. 1979. The carbohydrate and protein composition
of the endosperm, embryo and testa of the seed of Gleditsia triacanthos.
Journal of the Science of Food and Agriculture 30: 881891.
Singh DP, Hooda MS, Bonner FT. 1991. An evaluation of scarification
methods for seeds of two leguminous trees. New Forests 5: 139145.
Stubsgard F. 1986. Pretreatment of Acacia and Prosopis seed: two mechanical methods.Tech. Note 27. Humlebaek, Denmark: DANIDA Forest
Seed Centre. 8 p.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
Williams RD, Hanks SH. 1976. Hardwood nurserymans guide. Agric.
Handbk. 473. Washington, DC: USDA Forest Service. 78 p.
TheaceaeTea family
ommendations for germination testing, but the ease of germination suggests that the common alternating regime of
20/30 C would suffice. Germination of 70 to 80% within
10 days in petri dishes in sunlight has been reported
(Gresham and Lipscomb 1990). Germination is epigeal. This
species is also very easy to propagate vegetatively. Cuttings
taken in June through August rooted 90 to 100% in
peatperlite and mist with 3,000 ppm of IBA applied (Dirr
and Heuser 1987).
Figure 1Gordonia lasianthus, loblolly-bay:
capsules.
seeds.
Gordonia
565
longitudinal
References
Brown CK, Kirkman LK. 1990. Trees of Georgia and adjacent states.
Portland, OR:Timber Press. 292 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation. Athens, GA:Varsity Press. 239 p.
Gresham CA, Lipscomb DJ. 1990. Gordonia lasianthus (L.) Ellis, loblolly-bay.
In: Burns RM, Honkala BH, tech. coords. Silvics of North America.Volume
2. Hardwoods. Agric. Handbk. 654. Washington, DC: USDA Forest
Service: 365369.
Little EL, Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Vines RA. 1960. Trees, shrubs and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
566
ChenopodiaceaeGoosefoot family
Common name(s)
Occurrence
Sources: Collotzi (1966), Dayton (1931), Hitchcock and Cronquist (1973), Kay (1977), Shaw (1992a&b), Smith (1974),Welsh and others (1987).
Grayia
567
longitudinal
bracted utricle
568
hygroscopic bracts absorb water rapidly if exposed to environments with increased humidity.
Seed extraction and cleaning. Preliminary separation
of harvested seedlots with an air-screen machine removes
twigs, large leaves, and other coarse material. Some empty
bracts can also be separated by this process. Bracteoles may
be removed, if necessary, by threshing them with a hammermill (King 1947) or a barley de-bearder (Jorgensen 1992). A
seed scarifier, seed de-winger, or rubbing board may be used
to thresh small collections (Shaw and Haferkamp 1990).
Threshing generally results in complete removal of bracteoles and partial to complete removal of the pericarp, leaving
seeds as the product. Some embryos may be damaged during threshing as the radicle tip is vulnerable to abrasion
(figure 4).
Threshed seeds may be separated from chaff using an
air-screen machine or a seed blower. Removing the chaff is
necessary only when it is desirable to reduce bulk for storage or shipping. Otherwise, the chaff can serve as a diluent
for the small seeds as it will feed through most seeding
mechanisms when dry. Smith (1974) obtained 1.2 kg (2.6 lb)
of cleaned seeds from 45.4 kg (100 lb) of fruits. Number of
bracted utricles and seeds per weight and seed fill data are
provided in table 3.
Storage. Kay (1976) and Kay and others (1977, 1984,
1988) found that total germination percentage of seeds dried
to a water content of 5.1% and stored at 15 or 4 C or
room temperature in sealed glass containers containing a sil-
ica gel desiccant did not decline from the initial value of
42% after 14 years (Kay 1976; Kay and others 1977, 1984,
1988). Germination of air-dried seeds stored in cloth bags in
a warehouse decreased to about 20% after 1.5 years and to
0% after 7 years. All germination tests were conducted at 15
C. Thus, for long-term storage, it is recommended that
seeds be dried to a water content below 10% and kept in
sealed containers.
Pre-germination treatments and germination tests.
Dormancy of freshly harvested utricles of many woody
chenopods can be reduced by dry after-ripening, whereas the
response to wet prechilling and temperature is regulated by
the environmental conditions in which they were produced
(Ansley and Abernethy 1985; Kay and others 1988;
Springfield 1972). However, the response of spiny hopsage
seeds to dry after-ripening is poorly known and may vary
with seedlot and with seed age. Shaw and others (1994)
found that field germination and seedling establishment of
2 spiny hopsage seed collections from the northern shrub
steppe were similar after 2 and 4 years of dry storage at
room temperature. By contrast, King (1947) found that an
additional 2 years of dry after-ripening decreased the wet
prechilling (5 C) requirement for eastern Washington seeds
from 12 weeks for 4-year-old seeds to 2 weeks for 6-yearold seeds.
Spiny hopsage seeds produced in the northern shrub
steppe generally have a requirement for wet prechilling;
seeds produced in the Mojave Desert do not (Shaw 1992a;
Flowering
Fruit ripening
Seed dispersal
Mar
FebApril
MarMay
April & June
AprilMay
Mar
MarApr
May
May & July
MayJune
Mar
Apr
MayJune
May & Aug
MayJune
Sources: Ackerman and Bamberg (1974), Ackerman and others (1980), Blauer and others (1976), Branson and others (1967), Everett and others (1980), Goodrich and
Neese (1986), Plummer and others (1968), Shaw (1992b),Wallace and Romney (1972).
Table 3 Grayia spinosa, spiny hopsage: fruit and seed numbers per weight
Seeds/weight
Bracted utricles/weight
/kg
/lb
337,000447,000
152,900202,800
Range
/kg
339,000930,000
692,6001,031,600
Average
/lb
153,800421,800
314,200468,000
/kg
/lb
500,000
1,219,500
227,000
553,200
Sources: Belcher (1985), Kay and others (1977), King (1947), Plummer and others (1968), Shaw (1992b), Smith (1974), Swingle (1939).
Note: Filled seeds (%) = 18 to 95.
Grayia
569
570
Figure 5Grayia spinosa, spiny hopsage: seedling development at 1, 9, and 14 days after germination.
References
Ackerman TL, Bamberg SA. 1974. Phenological studies in the Mojave
Desert at Rock Valley (Nevada Test Site). In: Leith H, ed. Phenology and
seasonality modeling, ecological studies. New York: Springer-Verlag 8:
215226.
Ackerman TL, Romney EM, Wallace A, Kinnear JE. 1980. Phenology of
desert shrubs in southern Nye County, Nevada. Great Basin Naturalist
Memoirs 4: 423.
Ansley RJ, Abernethy RH. 1985. Environmental factors influencing Gardner
saltbush seed dormancy alleviation. Journal of Range Management 38:
331335.
Augustine G, Augustine J, Bach D, Backhaus R, Corgan J, and others 1979.
Soil mixes for greenhouse and nursery growth of desert plants. Desert
Plants 1: 8288.
Beall K. 2000. Personal communication. Boise, ID: USDA Forest Service,
Boise National Forest, Lucky Peak Nursery.
Belcher E. 1985. Handbook on seeds of browse-shrubs and forbs.Tech.
Pub. R8-8. Atlanta: USDA Forest Service, Southern Region. 246 p.
Blauer AC, Plummer AP, McArthur ED, Stevens R, Giunta BC. 1976.
Characteristics and hybridization of important Intermountain shrubs: 2.
Chenopod family. Res. Pap. INT-335. Ogden, UT: USDA Forest Service,
Intermountain Forest and Range Experiment Station. 78 p.
Branson FA, Miller RF, McQueen IS. 1967. Geographic distribution and factors affecting the distribution of salt desert shrubs in the United States.
Journal of Range Management 20: 287296.
Collotzi AW. 1966. Investigations in the genus Grayia, based on chromatographic, morphological, and embryological criteria [MS thesis]. Logan:
Utah State University. 42 p.
Dayton WA. 1931. Important western browse plants. USDA Misc. Pub.
101. Washington, DC: USDA. 214 p.
Drobnick R, Plummer AP. 1966. Progress in browse hybridization in Utah.
Proceedings of the Conference of Western State Game and Fish
Commissioners 46: 211213.
Dueleheimer C. 1992. Unpublished data. Boise: Idaho State Seed
Laboratory.
Grayia
571
Everett RL,Tueller PT, Davis JB, Bruner AD. 1980. Plant phenology in
galletashadscale and galletasagebrush associations. Journal of Range
Management 33: 446450.
Ferguson RB. 1980. Potting media for Atriplex production under greenhouse conditions. Res. Note INT-301. Ogden, UT: USDA Forest Service,
Intermountain Forest and Range Experiment Station. 7 p.
Ferguson RB, Frischknecht NC. 1981. Shrub establishment on reconstructed soils in semiarid areas. In: Stelter LH, DePuit EJ, Mikol SJ, tech. coords.
Shrub establishment on disturbed arid and semiarid lands; 1980
December 23; Laramie, WY. Cheyenne: Wyoming Game and Fish
Department: 5762.
Ferguson RB, Frischknecht NC. 1985. Reclamation on Utahs Emery and
Alton coal fields: techniques and plant materials. Res. Pap. INT-335.
Ogden, UT: USDA Forest Service, Intermountain Forest and Range
Experiment Station. 78 p.
Ferguson RB, Monsen SB. 1974. Research with containerized shrubs and
forbs in southern Idaho. In:Tinus RW, Stein WI, Balmer WE, eds.
Proceedings, North American Containerized Forest Tree Seedling
Symposium; 1974 August 2629; Denver, CO. Pub. 68. Lincoln, NB: Great
Plains Agricultural Council: 349358.
Frischknecht NC, Ferguson RB. 1984. Performance of Chenopodiaceae
species on processed oil shale. In:Tiedemann AR, McArthur ED, Stutz
HC, Stevens R, Johnson KL, comps. Proceedings, Symposium on the
Biology of Atriplex and Related Chenopods; 1983 May 26; Provo, UT.
Gen.Tech. Rep. INT-172. Ogden, UT: USDA Forest Service,
Intermountain Forest and Range Experiment Station: 293297.
Goodrich S, Neese E. 1986. Uinta Basin flora. Ogden, UT: USDA Forest
Service, Intermountain Region, Ashley National Forest; and USDI Bureau
of Land Management,Vernal District. 320 p.
Hitchcock CL, Cronquist A. 1973. Flora of the Pacific Northwest. Seattle:
University of Washington Press. 730 p.
Hunter RB, Wallace A, Romney EM. 1980. Fencing enhances shrub survival
and growth for Mojave Desert revegetation. Great Basin Naturalist
Memoirs 4: 212215.
Jorgensen K. 1992. Unpublished data. Ephraim: Utah Division of Wildlife
Resources.
Kay BL. 1976. Test of seeds of Mojave Desert shrubs. Prog. Rep. BLM contract 535000-CT4-2(N). Davis: University of California. 48 p.
Kay BL, Graves WL,Young JA. 1988. Long-term storage of desert shrub
seed. Mojave Reveg. Notes 23. Davis: University of California. 22 p.
Kay BL, Pergler CC, Graves WL. 1984. Storage of seed of Mojave Desert
shrubs. Journal of Seed Technology 9: 2028.
Kay BL, Ross CM, Graves WL. 1977. Hop-sage. Mojave Reveg. Note 6.
Davis: University of California. 5 p.
King JE. 1947. The effect of various treatments to induce germination of
seed of some plants valuable for soil conservation and wildlife [MS
thesis]. Moscow, ID: University of Idaho. 97 p.
Manning SJ, Groeneveld DP. 1990. Shrub rooting characteristics and water
acquisition on xeric sites in the western Great Basin. In: McArthur ED,
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Cheatgrass Invasion, Shrub Die-off, and Other Aspects of Shrub Biology
and Management; 1989 April 57; Las Vegas, NV. Gen.Tech. Rep. INT276. Ogden, UT: USDA Forest Service, Intermountain Research Station:
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McArthur ED, Sanderson SC. 1984. Distribution, systematics, and evolution
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Stevens R, Johnson KL, comps. Proceedings, Symposium on the Biology
of Atriplex and Related Chenopods; 1983 May 26; Provo, UT. Gen.Tech.
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McCullough DR. 1969. The Tule elk: its history, behavior, and ecology. Pub.
Zool. 88. Berkeley: University of California Press. 191 p.
572
Peters J, ed. 2000. Tetrazolium testing handbook. Contrib. 29. In: Handbook
on seed testing. [Las Cruces, NM]: Association of Official Seed Analysts.
Plummer AP, Christenson DR, Monsen SB. 1968. Restoring big-game range
in Utah. Pub. 68-3. Salt Lake City: Utah Division of Fish and Game. 183 p.
Rehder A. 1940. Manual of cultivated trees and shrubs. New York:
Macmillan. 996 p.
Rickard WH, Keough RF. 1968. Soil-plant relationships of two steppe
desert shrubs. Plant and Soil 19: 205212.
Rickard WH, Warren JL. 1981. Response of steppe shrubs to the 1977
drought. Northwest Science 55: 108112.
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National Symposium on Radioecology; 1971 May 10; Oak Ridge,TN. Los
Angeles: University of CaliforniaLos Angeles, School of Medicine.
Volume 2: 10151073.
Shaw NL. 1992a. Germination and seedling establishment of spiny hopsage
(Grayia spinosa [Hook.] Moq.) [PhD dissertation]. Corvallis: Oregon
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Shaw NL. 1992b. Unpublished data. Boise, ID: USDA Forest Service,
Forestry Sciences Laboratory.
Shaw NL, Haferkamp MR. 1990. Field establishment of spiny hopsage. In:
McArthur ED, Romney ER, Smith SD,Tueller PT, comps. Proceedings,
Symposium on Cheatgrass Invasion, Shrub Die-off, and Other Aspects of
Shrub Biology and Management; 1989 April 57; Las Vegas, NV. Gen.
Tech. Rep. INT-276. Ogden, UT: USDA Forest Service, Intermountain
Research Station: 193199.
Shaw NL, Haferkamp MR, Hurd EG. 1994. Germination and seedling establishment of spiny hopsage in response to planting date and seedbed
environment. Journal of Range Management 47: 165174.
Shaw NL, Hurd EG, Haferkamp MR. 1996. Spiny hopsage fruit and seed
morphology. Journal of Range Management 49: 551553.
Smith JG. 1974. Grayia H.&A., hopsage. In: Shopmeyer CS, tech. coord.
Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 434436.
Springfield HW. 1972. Winterfat fruits undergo afterripening. Journal of
Range Management 25: 6970.
Swingle CF. 1939. Seed propagation of trees, shrubs, and forbs for conservation planting. SCS-TP-27. Washington, DC: USDA Soil Conservation
Service. 198 p.
Tueller PT, Bruner AD, Everett R, Davis JB. 1974. The ecology of Hot Creek
Valley, Nevada and nonradiation effects of an underground nuclear detonation. USAEC Rep. 96. Reno: University of Nevada, Max C. Fleischmann
College of Agriculture. 51 p.
USDA SCS. 1968. Management and uses of spiny hopsage in the State of
Washington. Washington Plant Sci. Handbk. Spokane, WA: USDA Soil
Conservation Service, Plant Materials Section. 2 p.
Wallace A, Romney EM. 1972. Radioecology and ecophysiology of desert
plants at the Nevada Test Site. USAEC Rep.TID-25954. Riverside:
University of CaliforniaLos Angeles, Departments of Soil Science and
Agricultural Engineering; Laboratory of Nuclear Medicine and Radiation
Biology; Agricultural Sciences; and Environmental Radiation Division.
439 p.
Wallace A, Romney EM. 1974. Feasibility and alternate procedures for
decontamination and post-treatment management of PU-contaminated
areas in Nevada. Los Angeles: University of CaliforniaLos Angeles,
School of Medicine: 251337.
Wallace A, Romney EM, Hunter RB. 1980. The challenge of a desert: revegetation of disturbed desert lands. Great Basin Naturalist Memoirs 4:
216225.
Welsh SL, Atwood ND, Goodrich S, Higgins LC, eds. 1987. A Utah flora.
Great Basin Naturalist Memoirs 9: 1894.
Wood MK, Knight RW,Young JA. 1976. Spiny hopsage germination. Journal
of Range Management 29: 5356.
ProteaceaeProtea family
573
longitudinal section
germination rates ranging from 60 to 70% when seed moisture was maintained below 10% during storage in airtight
containers (Jones 1967).
Germination. Testing procedures for official purposes
call for 21 days of testing at alternating temperatures of
20/30 C with no pretreatment (AOSA 1993). Two
pregermination treatments have been found to increase substantially the germination of stored seeds (ETSL 1969). A
48-hour water soak and stratification at 3 C for 30 days
were equally effective. Pretreated seeds were germinated on
moist Kimpak at diurnally alternating temperatures of 30 C
during an 8-hour light period and 20 C during the dark
period. The average germination rate of 8 samples was 38%
after 17 days and 70% after 72 days. Germination of stored,
untreated seeds, however, was only 26% (ETSL 1969).
Fresh seed in Australia had a germination rate of 60 to 80%
(Goor and Barney 1968). Germination of fresh, unstratified
seeds require about 20 days (Skolmen 1990). Germination in
silk-oak is epigeal.
Nursery practice. Nursery practices vary among
countries where silk-oak is grown (Skolmen 1990). In some
countries 4- to 6-week-old wildlings are lifted and potted
and later replanted. Seedlings grown in Sri Lanka are outplanted when they are about 40 cm (16 in) tall, whereas
those in Jamaica are outplanted when about 60 cm (24 in)
tall (Streets 1962). Elsewhere, plants are grown to 45-cm
(18 in) heights in large baskets so that they can compete
more effectively when outplanted. In Hawaii, silk-oak seeds
are sown at a depth of 1.25/cm (1/2 in) without mulch
(Wong 1974). Seedbeds are treated with insecticides and
fungicides before sowing. Seedling density ranges from 200
to 300 seedlings/m2 (19 to 28/ft2). Seedlings grown in flats
are outplanted when they are about 9 months old (Wong
1974), whereas container-grown seedlings reach a plantable
size of 20 cm (8 in) in height in 12 to 14 weeks (Skolmen
1990).
References
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
ETSL [Eastern Tree Seed Laboratory]. 1969. Germination and sowing rate
report for 1969. Macon, GA: USDA Forest Service and Georgia Forest
Commission.
Goor AY, Barney CW. 1968. Forest tree planting in arid zones. New York:
Ronald Press. 409 p.
Haselwood EL, Motter GG. 1966. Handbook of Hawaiian weeds. Honolulu:
Hawaii Sugar Planters Association Experiment Station. 479 p.
Jones L. 1967. Effect of storage at various moisture contents and temperatures on seed germination of silk oak, Australian pine, and Eucalyptus spp.
Res. Note SE-83. Asheville, NC: USDA Forest Service, Southeast Forest
Experiment Station. 4 p.
Little, EL, Jr, Skolmen RG. 1989. Common forest trees of Hawaii. Agric.
Handbk. 670. Washington, DC: USDA Forest Service. 321 p.
Magini E,Tulstrup NP. 1955. Tree seed notes. For. Dev. Pap. 5. Rome: FAO.
574
354 p.
Neal MC. 1965. In gardens in Hawaii. Spec. Pub. 50. Honolulu: Bishop
Museum Press. 924 p.
Parry MS. 1956. Tree planting practices in tropical Africa. For. Dev. Pap. 8.
Rome: FAO. 302 p.
Schaefer C. 1991. Storage of tree seeds in Kenya: recommendations and
problems. In: Schnier P, ed. Proceedings, 1st National Tree Seed
Workshop; 1991 July 15; Nairobi, Kenya. Nairobi: Kenya Forestry
Research Centre: 99113.
Skolmen RG. 1990. Grevillea robusta A. Cunn. In: Burns RM, Honkala BH,
tech. coords. Silvics of North America.Volume 2, Hardwoods. Agric.
Handbk. 654. Washington, DC: US Department of Agriculture, Forest
Service: 370373.
Streets RJ. 1962. Exotic trees of the British Commonwealth. Oxford:
Clarendon Press. 765 p.
Wong WHC Jr. 1974. Grevillea robusta, A. Cunn., silk-oak. In: Schopmeyer
CS, tech. coord. Seeds of woody plants in the United States. Agric.
Handbk. 450. Washington, DC: USDA Forest Service: 437438.
AsteraceaeAster family
Gutierrezia Lag.
snakeweed
Kirk C. McDaniel and Ballard Wood
Dr. McDaniel is a professor and Mr. Wood a research assistant at New Mexico State Universitys
Department of Animal and Range Sciences, Las Cruces, New Mexico
Figure 1Gutierrezia sarothrae, broom snakeweed: flowering head (center); ray (left) and disk (right) flower with
attached achene (adapted from Ruffin 1974).
575
References
Kruse WH. 1970. Temperature and moisture stress affect germination of
Gutierrezia sarothrae. Journal of Range Management 23: 143144.
Lane MA. 1985. Taxonomy of Gutierrezia (Compositae: Astereae) in North
America. Systematic Botany 10: 728.
Mayeux HS. 1983. Effects of soil texture and seed placement on emergence of four subshrubs. Weed Science 31: 380384.
Mayeux HS. 1989. Snakeweed seed characteristics and germination
requirements. In: Huddleston EW, Pieper RD, eds. Snakeweed: problems
and perspectives. Bull. 751. Las Cruces: New Mexico State University,
Agricultural Experiment Station. 225 p.
Mayeux HS, Leotta L. 1981. Germination of broom snakeweed and
threadleaf snakeweed seed. Weed Science 29: 530534.
McDaniel KC, Pieper RD, Donart GB. 1982. Grass response following thinning of broom snakeweed. Journal of Range Management 35: 219222.
Oshman A, Pieper RD, McDaniel KC. 1987. Soil seed banks associated with
individual broom snakeweed plants. Journal of Range Management 40:
441443.
Gutierrezia
577
FabaceaePea family
References
Ball j, Kisor R. 1985. Acid scarificationrequrements of Kentucky coffeetree
(Gymnocladus diocious (L). K. Koch) seeds from
southcentral Minnesota.Tree Planters Notes 36(2); 23.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Engstrom HE, Stoeckler JH. 1941. Nursery practice for trees and shrubs
suitable for planting on the Prairie-Plains. Misc. Pub. 434. Washington, DC:
USDA. 159 p.
Evans CS, Bell EA. 1978. Uncommon amino acids in the seeds of 64
species of Caesalpinieae. Phytochemistry 17: 11271129.
Harr WH. 1927. Some studies on the structure and behavior during germination of Gymnocladus canadensis Lam. University of Kansas Science
Bulletin. 17(5): 331365.
Harrar ES, Harrar JG. 1946. Guide to southern trees. New York: McGrawHill Book Company. 712 p.
Liu NY, Khatamian H, Fretz TA. 1981. Seed coat structure of three woody
legume species after chemical and physical treatments to increase seed
germination. Journal of the American Society for Horticultural Science
106: 691694.
Phillips GR. 1931. Collection, care, and planting of tree seeds. Pub. 10.
Oklahoma City: Oklahoma Forest Service. 16 p.
Rehder A. 1940. Manual of cultivated trees and shrubs hardy in North
America. 2nd ed. New York: Macmillan. 996 p.
Rehr SS, Bell EA, Janzen DH, Feeny PP. 1973. Insecticidal amino acids in
legume seeds. Biochemical Systematics 1(1): 6367.
Sander IL. 1974. Gymnocladus dioicus (L.) K. Koch., Kentucky coffeetree. In:
Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service:
439440.
Sargent CS. 1965. Manual of the trees of North America (exclusive of
Mexico). 2nd ed., corrected and reprinted. New York: Dover. 934 p.
Van Dersal WR. 1939. Native woody plants of the United States: their erosion-control and wildlife values. Misc. Pub. 303. Washington, DC: USDA
362 p.
Weisehuegel EG. 1935. Germinating Kentucky coffeetree. Journal of
Forestry 33: 533534.
Gymnocladus
579
StyracaceaeStorax family
Halesia carolina L.
Carolina silverbell
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station
Mississippi State, Mississippi
Other common names. opossum-wood, silverbell,
snowdrop-tree.
Growth habit, occurrence, and uses. Carolina silverbellHalesia carolina L.is a small, deciduous tree found
naturally from West Virginia and southern Illinois south to
South Carolina, northwestern Florida, and Alabama, with
small pockets as far west as Arkansas and Oklahoma
(Sargent 1965; Sluder 1990). It has been successfully planted in Massachusetts and California and also to some extent
in northern and central Europe. Carolina silverbell was first
cultivated in 1756 (Bonner and Mignery 1974). It is highly
valued as an ornamental, and its fruits are a source of food
for wildlife.
Flowering and fruiting. The perfect, white (sometimes pink), bell-shaped, axillary flowers of Carolina silverbell are borne in fascicles of 1 to 5 in March to May (Brown
and Kirkman 1990; Sluder 1990). The species is precocious
and seedlings may flower when only a little over 1 m in
height (Dirr and Heuser 1987). The fruit, which matures in
August and September, is an oblong or oblong-ovate, dry, 4winged, reddish brown, corky drupe 2.5 to 5 cm long. The
ovary is a 4-celled ellipsoid stone, 13 to 16 mm long, usually containing only 1 seed (figures 1 and 2) (Bonner and
Mignery 1974; Brown and Kirkman 1990; Sluder 1990).
The fruits are persistent on the branches, and dispersal
occurs well into the winter.
Collection, extraction, and storage. Carolina silverbell fruits may be collected from the trees in late fall and
early winter. De-winging can be done mechanically
(Thornhill 1968) and is recommended to reduce bulk and
facilitate handling. Complete extraction of stones from the
fruits is not necessary. Bonner and Mignery (1974) found
about 2,600 to 5,500 de-winged fruits/kg (1,200 to 2,500/lb)
using 2 samples. Although no data on long-term storage are
available, dry cold storage of cleaned fruits has been recommended (Chadwick 1935).
580
fruits.
longitudi-
the flats are then moved to an outdoor cold frame for the
cold part of the after-ripening treatment. The flats are protected by mulch or by board covers on the coldframes
(Bonner and Mignery 1974).
Carolina silverbell can also be propagated by cuttings
taken in mid-summer after shoot elongation has ceased but
before fall hardening sets in. It is best to treat cuttings with
2,500 or 10,000 ppm indole butyric acid (IBA) solution and
place them in peat or perlite under mist. Rooting success at
levels of 80 to 100% can be expected (Dirr and Heuser
1987; Sluder 1990). Micropropagation techniques are also
under study (Brand and Lineberger 1986).
References
seedling
Bonner FT, Mignery AL. 1974. Halesia carolina var. carolina, Carolina silverbell. In: Schopmeyer CS, tech. coord. Seeds of woody plants in the
United States. Agric. Handbk. 450. Washington, DC: USDA Forest
Service: 441442.
Brand MH, Lineberger RD. 1986. In vitro propagation of Halesia carolina L.
and the influence of explantation timing on initial shoot proliferation.
Plant Cell,Tissue and Organ Culture 7: 103113.
Brown CL, Kirkman LK. 1990. Trees of Georgia and adjacent states.
Portland, OR:Timber Press. 292 p.
Chadwick LC. 1935. Practices in propagation by seed. American
Nurseryman 62(12): 39.
Dirr MA. 1977. The silverbells. American Nurseryman 146(3): 1214, 42,
44, 46.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation. Athens, GA: Varsity Press. 239 p.
Sargent CS. 1965. Manual of the trees of North America (exclusive of
Mexico). 2nd ed., corrected and reprinted. New York: Dover. 934 p.
Sluder ER. 1990. Halesia carolina L., Carolina silverbell. In: Burns RM,
Honkala BH, tech. coords. Silvics of North America.Volume 2,
Hardwoods. Agric. Handbk. 654. Washington, DC: USDA Forest Service:
374378.
Thornhill HB. 1968. Propagation of woody ornamentals by seed. American
Nurseryman 127(6):
Halesia
581
HamamelidaceaeWitch-hazel family
Hamamelis L.
witch-hazel
Jill R. Barbour and Kenneth A. Brinkman
Ms. Barbour is a germination specialist at the USDA Forest Services National Seed Laboratory,
Dry Branch, Georgia; Dr. Brinkman retired from the USDA Forest Services
North Central Forest Experiment Station
582
Common name
Japanese witch-hazel
Chinese witch-hazel
Ozark witch-hazel
American witch-hazel
Occurrence
Japan
W central China
Ozark Mtns of Missouri & Arkansas
E US & Canada
Height (m)
10
10
2
710
Hamamelis
583
Nursery practice. Witch-hazel seeds may be fallsown in the nursery as soon as collected, or stratified seeds
may be sown in the spring. Limited trials show that seeds
collected as early as August and sown by early October
results in as much as 90% germination the following spring
(Heit 1968; Sandahl 1941). Fall-sowing is recommended;
the seedbeds should be mulched over winter and uncovered
at germination time in the spring. For spring-sowing of
stratified seeds, seedbeds should be prepared as early as soil
conditions permit. Sowing in drills spaced 20 to 30 cm
(8 to 12 in) apart will facilitate weeding and cultivating.
Secondary leaves may develop on a seedling within 21 days
after germination (figure 3). Propagation by layering also is
possible.
584
seedling at
References
Brinkman KA. 1974. Hamamelis virginiana L., witch-hazel. In: Schopmeyer
CS, tech. coord. Seeds of woody plants in the United States. Agric.
Handbk. 450. Washington, DC: USDA Forest Service: 443444.
DeSteven D.1982. Seed production and seed mortality in a temperate forest shrub (witch-hazel, Hamamelis virginiana) Journal of Ecology 70:
437443.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1631 p.
Fernald ML. 1970. Grays manual of botany. 8th ed. New York:Van
Nostrand Co. 1632 p.
Fordham AJ. 1976. Propagation of some Hamamelidaceae (witch-hazel
family) Combined Proceedings of the International Plant Propagators
Society 26: 296298.
Fordham AJ. 1991. Propagation techniques of Cornus kousa and Hamamelis
taxa: 1940s vs. 1980s. Combined Proceedings of the International Plant
Propagators Society 1990 40: 524527.
Heit CE. 1968. Propagation from seed: 15. Fall planting of shrub seeds for
successful seedling production. American Nurseryman 128(4): 810,
7080.
Hora B. 1981. The Oxford encyclopedia of trees of the world. Oxford, UK:
Oxford University 288 p.
Johnson H. 1973. The international book of trees. New York: Simon &
Schuster. 288 p.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan. 1290 p.
Little EL Jr. 1953. Checklist of native and naturalized trees of the United
States (including Alaska). Agric. Handbk. 41. Washington, DC: USDA
Forest Service. 472 p.
Moore RP. ed. 1985. Handbook on tetrazolium testing. Zurich: International
Seed Testing Association. 99 p.
Rehder A. 1940. Manual of cultivated trees and shrubs. 2nd ed. New York:
Macmillan. 996 p.
Rivera R, Popp HW, Dow RB. 1937. The effect of high hydrostatic pressures upon seed germination. American Journal of Botany 24: 508513.
Sandahl PL. 1941. Seed germination. Parks and Recreation 24(11): 508.
Van Dersal WI. 1938. Native woody plants of the United States: their erosion-control and wildlife values.
RosaceaeRose family
seeds.
Christmasberry seed weight is apparently highly variable. Magill (1974) reported a mean seed weight of 19 mg
and count of 52,630 seeds/kg (23,900/lb), whereas Keeley
(1991) reported a seed weight of 5.5 mg and count of
181,820/kg (82,500/lb). Martineja and Bullock (1997) examined Christmasberry seed weight as a function of habitat
variables. They found no correlation with latitude or annual
precipitation but did find a significant increase in seed
weight with increasing elevation. Overall mean seed weight
for their 12 Christmasberry populations was 36 mg and seed
count was 27,800 seeds/kg (12,600/lb), with weight ranges
of 28 to 49 mg and counts of 20,400 to 35,700 seeds/kg
(9,200 to 16,200/lb).
Christmasberry seeds have limited longevity at room
temperature, but they are probably orthodox in storage
behavior. Keeley (1991) reported a shelf life of less than 1
year in laboratory storage. The seeds were also damaged or
killed by high temperature treatments. One hour at 70 C
reduced viability from 99 to 33%, and 5 minutes at 120 C
resulted in essentially complete mortality (Keeley 1987).
Magill (1974) recommended sealed storage at low temperaHeteromeles
585
ture but did not give any data on viability retention under
these conditions.
Germination and seed testing. Christmasberry seeds
are reported to be nondormant at dispersal (Emery 1988;
Keeley 1987; Magill 1974), whereas seeds that have been
stored are rendered secondarily dormant and require 3
months of chilling at 3 to 5 C in order to germinate. Under
field conditions, Christmasberry seeds germinate within a
few months of dispersal and do not form a persistent seedbank (Parker and Kelly 1989). Germination is epigeal (figure 3). Recruitment of new individuals is rarely observed.
Although winter seedling emergence is a common occurrence, the seedlings almost invariably die, either from herbivory or summer drought (Parker and Kelly 1989). Because
of the transient seed bank, there can be no recruitment after
fire, and new recruitment is in fact restricted to chaparral
stands that have been free of fire for at least 50 years
(Keeley 1992). The seedlings are not very drought-tolerant
and seem to need the shade and the deep litter that develops
under adult shrub canopies in old stands in order to survive.
Recently harvested lots of Christmasberry seeds that are
well-cleaned to remove unfilled seeds generally have high
fill and high viability. Keeley (1987) reported germination of
99% at 23 C. Such lots should be relatively easy to evaluate, either with a germination test or with tetrazolium staining. A 3-month chill at 5 C followed by a germination test
of 28 days at 20 or 25 C should give maximum germination. Christmasberry seeds have no endosperm, and the
embryo completely fills the seed cavity (figure 2). A procedure similar to that used for apple (Malus spp.) or bitter-
586
young
References
Emery DE. 1988. Seed propagation of native California plants. Santa
Barbara, CA: Santa Barbara Botanic Garden. 107 p.
Greever PT. 1979. Propagation of Heteromeles (Photinia) arbutifolia by softwood cuttings or by seed. Plant Propagator 25 (2): 1011.
Keeley JE. 1987. Role of fire in seed germination of woody taxa in
California chaparral. Ecology 68: 434443.
Keeley JE. 1991. Seed germination and life history syndromes in the
California chaparral. Botanical Review 57: 81116.
Keeley J. 1992. Recruitment of seedlings and vegetative sprouts in
unburned chaparral. Ecology 73: 11941208.
Magill AW. 1974. Photinia arbutifolia Lindl., Christmasberry. In: Schopmeyer
CS, tech. coord. Seeds of woody plants in the United States. Agric.
Handbk. 450. Washington, DC: USDA Forest Service: 582583.
Martineja NE, Bullock SH. 1997. Geographic variation in seed mass in the
chaparral shrub Heteromeles arbutifolia (Rosaceae). Southwestern
Naturalist 42: 119121.
Parker VT, Kelly VR. 1989. Seed banks in California chaparral and other
Mediterranean climate shrublands. In: Leck MA, Parker VT, Simpson RL,
eds. Ecology of soil seed banks. San Diego: Academic Press: 231255.
Phipps JB. 1992. Heteromeles and Photinia (Rosaceae, subfam. Maloideae) of
Mexico and Central America. Canadian Journal of Botany 70:
21382182.
Heteromeles
587
ElaeagnaceaeOleaster family
Hippophae rhamnoides L.
common seabuckthorn
Richard T. Busing and Paul E. Slabaugh
Dr. Busing is an ecologist at the USDI Geographical Survey, Corvallis, Oregon; Dr. Slabaugh (deceased)
retired from the USDA Forest Services Rocky Mountain Forest and Range Experiment Station
588
References
Avanzanto D, Magherini R, Lodoli E. 1987. Studies on the germination
potential of seeds and the rooting ability of cuttings of Hippophae rhamnoides L. Ministero dellAgricoltura e delle Foreste 1987: 411419.
Bogdon N, Untaru E. 1967. The substitution of Hippophae rhamnoides on
eroded sites in Vrancea [in Romanian]. Revue Pdurilor 82(5): 238243.
Cram WH, Nagv MJ, Lindquist CH. 1960. Propagation research. In: 1960
Summary report for the Forest Nursery Station. Indian Head, SK:
Canada Department of Agriculture, Research Branch: 1618.
Demenko VI, Potemkina GA, Medvedkova LA. 1983. Biological aspects of
fruit growth and shedding in sea buckthorn. Biologicheskie Nauki
(Moscow) 10: 7883.
Eliseev IP, Mishulina IA. 1972. New data on the biology of germination of
Hippophae rhamnoides seeds.Trudy Gorkovskogo
Selskokhozyaistvennogo Instituta 38: 107109.
Eliseev IP, Mishulina IA. 1977. Changes in the biological and biochemical
properties of Hippophae rhamnoides seeds during ripening.Trudy
Gorkovskogo Selskokhozyaistvennogo Instituta 105: 1522.
Hippophae.
589
Papp L. 1982. The importance of vegetative propagation of the sea buckthorn (Hippophae rhamnoides). Erdo 31(7): 309312.
Pearson, MC, Rogers JA. 1962. Hippophae rhamnoidea L. Journal of Ecology
50: 501513.
Rehder A. 1940. Manual of cultivated trees and shrubs. 2nd ed. New York:
Macmillan. 996 p.
Rohmeder E. 1942. Keim-und Saatversuche mit Sanddorn (H. rhamnoidea).
Forstwissenschaftliches Centralblatt 64: 241245.
Slabaugh PE. 1974. Hippophae rhamnoides L., common seabuckthorn. In:
Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service:
446447.
Smirnova NG,Tikhomirova NI. 1980. Combined use of x-ray photography
and the tetrazolium method for assessing seed viability. Byulleten
Glavnogo Botanicheskogo Sada 117: 8185.
590
RosaceaeRose family
Common name(s)
Occurrence
creambush ocean-spray,
creambush, creambush
rockspirea, hardhack,
Indian arrow-wood, ocean-spray
S British Columbia,Washington,
Oregon,W Montana, N Idaho,
NE Nevada, & California
gland ocean-spray,
bush ocean-spray, bush
rockspirea, mountain-spray,
rock-spirea, creambush
Sources: Archer (2000), Flessner and others (1991), Hitchcock and others (1971), Ley (1943), McMurray (1987b).
Holodiscus
591
achenes of H. discolor,
rare and costly. In addition, the achenes are difficult to handle because of their pubescence and small size. Achenes are
hand-stripped from inflorescences in late summer or autumn
(table 2) (Monsen 1996). Large debris in air-dried collections can be removed with a fanning mill. Small lots may be
cleaned by hand-rubbing and sieving (Link 1993).
Sound achenes are identified by examining imbibed achenes through a dissecting microscope for the presence of an
embryo. Using this method, King (1947) found that only 7%
of ocean-spray achenes collected were sound. In creambush
ocean-spray from British Columbia, viability was greater for
achenes collected in October or November than for those
collected in August or September (Marchant and Sherlock
1984).
Storage requirements for ocean-spray have not been
examined. The achenes appear to be orthodox in storage
behavior and can probably be stored for several years at low
water contents and temperatures.
Germination. There are no official testing prescriptions for this genus. Germination of creambush ocean-spray
seeds is enhanced by wet prechilling at 2 to 5 C for 15 to
18 weeks (King 1947; Marchant and Sherlock 1984). King
Location
Flowering
Fruit ripening
Seed dispersal
H. discolor
California
N Idaho
N Idaho
Great Basin
Utah
MayAug
July
July
JuneAug
JuneAug
Aug
Late Julyearly Sept
Aug
AugNov
Aug
H. dumosus
Sources: Drew (1967), Jorgensen (2004), Mozingo (1987), Munz and Keck (1973), Orme and Leege (1980),Welsh and others (1987).
592
be lifted as 1+0 or 2+0 stock, depending upon size specifications and growing conditions.
Container seedlings are propagated by planting several
wet-prechilled achenes in each container and thinning or by
planting germinants. Kruckeberg (1982) reported that oceanspray can be propagated by fall-sowing achenes in boxes
outdoors and covering them lightly with soil. Flessner and
others (1992) planted wet prechilled (4 months at 4 C)
creambush ocean-spray achenes in shallow flats in a greenhouse. Seedlings emerged after 16 to 30 days of incubation
at a minimum temperature of 21 C. Developing seedlings
were fertilized and treated with a fungicide as necessary.
After 2 months they were transferred to larger containers in
a lathhouse and held overwinter for planting as 1+0 stock.
Kruckeberg (1982) reported that creambush ocean-spray
planting stock is easily obtained by potting wildlings, which
are often abundant. Morgan and Neuenschwander (1988)
observed high densities of creambush ocean-spray wildlings
following severe burns, but Wright and others (1979) and
Stickney (1996) concluded that the species exhibits poor
seedling regeneration following fire in sagebrush (Artemesia
spp.) and conifer communities of the intermountain and
northern Rocky Mountain regions.
Ocean-spray can be grown from cuttings, but rooting of
both species varies widely among clones, cutting types, and
propagation techniques (Antieau 1987; Link 1993).
Softwood cuttings may be treated with rooting hormones
and propagated in a greenhouse with a mist system (Antieau
1987; Marchant and Sherlock 1984). Success with semihardwood cuttings is variable (Everett and others 1978;
Kruckeberg 1982). Fall-harvested hardwood cuttings are cut
to 15-cm (6-in) lengths and treated with 0.8% indole-3butyric acid (IBA) powder and a fungicide (Macdonald
1986). Hardwood cuttings stored in straw-bale bins or cold
frames will develop calluses (Macdonald 1986; Marchant
and Sherlock 1984). When fall-planted, these cuttings root
rapidly. Layers and suckers have also been propagated successfully (Kruckeberg 1982).
Field practice. Fresh achenes broadcast over a rough
seedbed in fall are covered by natural soil sloughing (Shaw
and Monsen 2004; Van Dersal 1938). Achenes may be
mixed with seeds of other shrub species, but they should not
be sown with more competitive grasses or forbs. Planting
areas should be selected carefully to make the best use of
seed supplies, as seeding results are often erratic. Native
creambush ocean-spray seedlings develop slowly and are
poor competitors (Wright and others 1979).
Holodiscus
593
Creambush ocean-spray can be established by transplanting. Youtie (1992) reported good survival of rooted
cuttings on biscuit scablands in Oregons Columbia River
Gorge. Marchant and Sherlock (1984) found that planted
seedlings grew slowly the first year. Low survival on western Montana roadcuts was attributed to poor soils and
unhealthy planting stock (Hungerford 1984).
References
Anderson BA, Holmgren AH. 1969. Mountain plants of northeastern Utah.
Circ. 319. Logan: Utah State University. 148 p.
Antieau CJ. 1986. Patterns of natural variation in ocean-spray (Holodiscus
discolor) (Rosaceae). HortScience 21: 120.
Antieau CJ. 1987. Field notes: Holodiscus discolor. American Nurseryman
166: 110.
Archer AJ. 2000. Holodiscus discolor. In: Fire Effects Information System,
available online at http://www.fs.fed.us/database/feis/]. Missoula, MT:
USDA Forest Service, Rocky Mountain Research Station, Fire Sciences
Laboratory.
Atthowe H. 1993. Propagation of riparian and wetland plants. In: Landis TD,
tech. coord. Proceedings, Western Forest Nursery Association; 1992
September 1418; Fallen Leaf Lake, CA: Gen.Tech. Rep. RM-221. Fort
Collins, CO: USDA Forest Service, Rocky Mountain Forest and Range
Experiment Station: 7881.
Daubenmire R. 1970. Steppe vegetation of Washington.Tech. Bull. 62.
Pullman: Washington State University.
Drew LA. 1967. Comparative phenology of seral shrub communities in the
cedar/hemlock zone [thesis]. Moscow, ID: University of Idaho. 108 p.
Everett PC. 1957. A summary of the culture of California plants at the
Rancho Santa Ana Botanic Garden. Claremont, CA: Rancho Santa Ana
Botanic Garden. 223 p.
Everett RL, Meeuwig RO, Robertson JH. 1978. Propagation of Nevada
shrubs by stem cuttings. Journal of Range Management 31: 426429.
Ferguson RB. 1983. Use of rosaceous shrubs for wildland plantings in the
Intermountain West. In: Monsen SB, Shaw N, comps. Proceedings;
Managing Intermountain RangelandsImprovement of Range and
Wildlife Habitats; 1981 September 1517;Twin Falls, ID, & 1982 June
2224; Elko, NV: Gen.Tech. Rep. INT-157. Ogden, UT: USDA Forest
Service, Intermountain Forest and Range Experiment Station: 136139.
Flessner TR, Darris DC, Lambert SC. 1992. Seed source evaluation of four
native riparian shrubs for streambank rehabilitation in the Pacific
Northwest. In: Clary WP, McArthur ED, Bedunah D, Wambolt CL,
comps. Proceedings, Symposium on Ecology and Management of
Riparian Shrub Communities; 1991 May 2931; Sun Valley, ID: Gen.Tech.
Rep. INT-289. Ogden, UT: USDA Forest Service, Intermountain
Research Station: 155162.
Goldblatt P. 1979. Miscellaneous chromosome counts in Angiosperms: 2.
Including new family and generic records. Annals of Missouri Botanic
Gardens 66: 856861.
Halverson NM,Topik C,Van Vickle R. 1986. Plant association and management guide for the western hemlock zone: Mt. Hood National Forest.
R6-ECOL-232A. Portland, OR: USDA Forest Service, Pacific Northwest
Region. 111 p.
Harrington HD. 1954. Manual of the plants of Colorado. Denver: Sage
Books. 666 p.
Hitchcock CL, Cronquist A, Ownbey M,Thompson JW. 1961. Vascular
plants of the Pacific Northwest: 3. Saxifragaceae to Ericaceae. Seattle:
University of Washington Press. 614 p.
Hopkins WE, Kovalchik BL. 1983. Plant associations of the Crooked River
National Grassland. R6-ECOL-133-1983. Portland, OR: USDA Forest
Service, Pacific Northwest Region. 98 p.
Hungerford RD. 1984. Native shrubs: suitability for revegetating roadcuts in
northwestern Montana. Res. Pap. INT-331. Ogden, UT: USDA Forest
Service, Intermountain Forest and Range Experiment Station. 13 p.
Hurd EG. 1996. Unpublished data. Boise, ID: USDA Forest Service,
Intermountain Research Station.
Jorgensen K. 2004. Appendix I. In: Monsen SB, Stevens R, comps. Restoring
western ranges and wildlands. Fort Collins, CO: USDA Forest Service,
Rocky Mountain Research Station.
King JE. 1947. The effect of various treatments to induce germination of
seeds of some plants valuable for soil conservation and wildlife [thesis].
Moscow, ID: University of Idaho. 97 p.
Kruckeberg, AR. 1982. Gardening with native plants of the Pacific
Northwest. Seattle: University of Washington Press. 252 p.
Ley A. 1943. A taxonomic revision of the genus Holodiscus (Rosaceae).
Bulletin of the Torrey Botanical Club 70: 275288.
Link E. 1993. Native plant propagation techniques for National Parks: interim guide. East Lansing, MI: Rose Lake Plant Materials Center. 240 p.
594
FabaceaePea family
Hymenaea courbaril L.
courbaril
J. A.Vozzo
Dr.Vozzo is a research plant physiologist at the USDA Forest Services Southern Research Station
Mississippi State, Mississippi
Flowering and fruiting. Large trees with full, overhead light usually flower during spring and summer.
Terminal racemes bear white flowers about 4 cm wide.
Mature legumes (figure 1) measure 5 to 10 cm long, 2 to 3.5
cm wide, and 2.5 cm thick and fall during the following
spring. The thick, hard legume does not open naturally, but
protects 3 or 4 large seeds (figures 2 and 3) encased in a
powdery, cream-colored pulp (Liogier 1978). Small animalssuch as agouties (Tayassu spp.) and peccaries
(Dasyprocta spp.)open the legumes to eat the pulp and
seeds. Legumes have a protective gum that delays rotting for
several months, until the seeds begin to take up moisture for
germination; otherwise the seeds would rot in their legumes
(Jansen 1983).
Collection and storage. Seeds collected in Puerto
Rico average about 253/kg (115/lb) (Francis 1990), whereas
those collected in Brazil yield 475/kg (215/lb) (Pereira
1982). A single tree may produce 100 legumes per year but
not necessarily each year. Because of the height of the trees,
the legumes are usually picked manually from the ground,
and seeds are obtained from fresh legumes that have fallen
in spring (Jansen 1983). After-ripening causes an actual
legume.
Hymenaea
595
seeds.
longitudinal
References
Allen ON, Allen EK. 1981. The Leguminoseae: a source book of characteristics, uses, and nodulation. Madison: University of Wisconsin Press. 812 p.
Chudnoff M. 1984. Tropical timbers of the world. Agric. Handbk. 607.
Washington, DC: USDA Forest Service. 466 p.
Decelle J. 1979. Pygiopachymerus lineola (Chevr.), coleoptere bruchide
neotropical introduit a Tahiti. Bulletin et Annales de la Societe Royale
Belge dEntomologie.Tervuren, Belgium: Musee Royale de lAfrique
Centrale.
Francis JK. 1990. Hymenaea courbaril (L.). SO-ITF-SM 27. New Orleans:
USDA Forest Service, Southern Forest Experiment Station. 5 p.
Francis JK, Rodrguez A. 1993. Seeds of Puerto Rican trees and shrubs: second installment. Res. Note SO-374. New Orleans: USDA Forest Service,
Southern Forest Experiment Station. 5 p.
Jansen DL. 1975. Behavior of Hymenaea courbaril when its predispersal
seed predator is absent. Science 189(4194): 145147.
Jansen DL. 1983. Costa Rican natural history. Chicago: University of
Chicago Press. 816 p.
596
AquifoliaceaeHolly family
Ilex L.
holly
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station
Mississippi State, Mississippi
Growth habit, occurrence, and use. The hollies
genus Ilexinclude almost 400 species of deciduous or evergreen shrubs and trees that occur in temperate and tropical
regions of both hemispheres (Brown and Kirkman 1990).
About 20 species are native to eastern North America. Of the
7 species included in this book (table 1), most are highly
valued for ornamental plantings and all are good food
sources for wildlife. More than a thousand cultivars of
American holly have been selected for their ornamental features (Grelen 1990). This species also hybridizes with
dahoon (Ilex cassine L.) to produce Topel holly (I. attenuata Ashe) (Little 1979). The wood of American holly is also
used in cabinetry and for construction of novelties and specialized wood products (Vines 1960).
Flowering and fruiting. The small, axillary, white or
greenish white, dioecious flowers appear in the spring on the
current seasons growth (table 2). Holly fruits are rounded,
berrylike drupes that range from 6 to 13 mm in diameter
(figure 1). Each fruit contains 2 to 9 bony, flattened seeds
that are botanically defined as nutlets, or pyrenes (figure 2).
The fruits mature in the fall (table 2), turning from green to
various shades of red, yellow, or black (table 3). The seeds
Common name(s)
Occurence
I. aquifolium L.
English holly
I. decidua Walt.
Ilex
597
Location
Flowering
Fruit ripening
Seed dispersal
I. aquifolium
I. decidua
I. glabra
I. montana
I. opaca
I. verticillata
I. vomitoria
Appalachian Mtns
MayJune
MarMay
MarJune
MayJune
AprJune
JuneJuly
AprMay
Sept
Fall
Fall
Sept
SeptOct
SeptOct
SeptOct
Winterspring
Winterspring
Winterspring
Mar
Fallwinter
Dec
Sources: Bonner (1974), Halls (1973), Little and Delisle (1962), Stupka (1964),Vines (1960).
598
longitudinal
Table 3Ilex, holly: height, seed-bearing age, and color of ripe fruit
Species
I. aquifolium
I. decidua
I. glabra
I. montana
I. opaca
I. verticillata
I. vomitoria
Height at
maturity (m)
1524
69
4
12
30
8
38
Year first
cultivated
Ancient times
1759
1870
1744
1736
512
47
Sources: Bonner (1974), Brown and Kirkman (1990), Grelen and Duvall (1966), Halls (1973), Little and Delisle (1962), Maisenhelder (1958), Rehder (1962),Vines (1960).
/kg
48,50080,150
88,200284,450
Average
/lb
/kg
/lb
Samples
22,00036,350
40,000129,000
125,700
43,6000
63,900
77,200
62,700
202,860
83,350
57,000
19,800
29,000
35,000
28,430
92,000
37,800
1
1
1
1
4
4
1
599
References
I
Afanasiev M. 1942. Propagation of trees and shrubs by seed. Circ. C-106.
Stillwater: Oklahoma Agricultural Experiment Station. 43 p.
Bonner FT. 1974. Ilex L., holly. In: Schopmeyer CS, tech. coord. Seeds of
woody plants in the United States. Agric. Handbk. 450. Washington, DC:
USDA Forest Service: 450543.
Brown CL, Kirkman LK. 1990. Trees of Georgia and adjacent states.
Portland, OR:Timber Press. 292 p.
Barton LV,Thornton NC. 1947. Germination and sex population studies of
Ilex opaca Ait. Contributions of the Boyce Thompson Institute 14:
405410.
Dirr MA, Heuser Jr CW. 1987. The reference manual of woody plant propagation. Athens, GA:Varsity Press. 239 p.
Giersbach J, Crocker W. 1929. Germination of Ilex seeds. American Journal
of Botany 16: 854855.
Grelen HE. 1990. Ilex opaca Ait., American holly. In: Burns RM, Honkala BH,
tech. coords. Silvics of North America.Volume 2, Hardwoods. Agric.
Handbk. 654. Washington, DC: USDA Forest Service: 379385.
Grelen HE, Duvall VL. 1966. Common plants of longleaf pinebluestem
range. Res. Pap. SO-23. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 96 p.
Halls LK. 1973. Flowering and fruiting of southern browse species. Res. Pap.
SO-90. New Orleans: USDA Forest Service, Southern Forest
Experiment Station. 10 p.
Hartmann HT, Kester DE. 1968. Plant propagation: principles and practices.
2nd ed. Englewood Cliffs, NJ: Prentice-Hall, Inc.
Hu CY. 1975. In vitro culture of rudimentary embryos of eleven Ilex species.
Journal of the American Society for Horticultural Science 100: 221225.
Hu CY. 1977. Advances in Ilex embryo culture. Proceedings, 54th Meeting
of the Holly Society of America 1977: 56 [Horticultural Abstracts
600
JuglandaceaeWalnut family
Juglans L.
walnut
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Common name(s)
Occurrence
J. ailantifolia Carriere
J. sieboldiana maxim.
J. californica S.Wats.
Japanese walnut,
Siebold walnut
California walnut, southern
California walnut, black walnut
butternut, oilnut,
white walnut
Japan
J. cinerea L.
Wallia cinerea (L.) Alef.
J. hindsii (Jepson) Jepson ex R.E. Sm.
J. californica var. hindsii Jepson
J. major (Torr.) Heller
J. rupestris var. major Torr.
J. microcarpa var. major (Torr.) L. Benson
J. elaeopyren Dode
J. microcarpa Berl.
J. rupestris Englem. ex Torr.
J. nigra L.
Wallia nigra (L.) Alef.
J. regia L.
Juglans
601
Flowering
Fruit ripening
Seed dispersal
J. ailantifolia*
J. californica
J. cinerea
J. hindsii
J. major
J. microcarpa
J. nigra
J. regia
MayJune
MarApr
AprJune
AprMay
Spring
MarApr
AprJune
MarMay
AugOct
Fall
SeptOct
AugSept
Fall
AugSept
SeptOct
SeptNov
Oct
Fall
After leaf-fall
SeptOct
Fall
Fall
After leaf-fall
Fall
602
Table 3Juglans, walnut: height, seed-bearing age, seedcrops frequency, and fruit ripeness criteria
Species
Height at
maturity (m)
Year first
cultivated
Minimum
seed-bearing
age (yrs)
Years
between large
seedcrops
20
12
30
24
15
6
46
27
1860
1889
1633
1878
1894
1868
1686
Long cultivated
10
5-8
20
9
20
12
8
13
23
23
J. ailantifolia
J. californica
J. cinerea
J. hindsii
J. major
J. microcarpa
J. nigra
J. regia
Light green
Greenish bronze
Light yellow-green
Light green
Light yellowish green
Dark brown
Greenish brown
Dark brown to black
Yellowish green
Black
Species
J. ailantifolia
J. californica
J. cinerea
J. hindsii
J. major
J. microcarpa
J. nigra
J. regia
Place
collected
Japan
California
Shasta Co.,
California
Coconino Co.,
Arizona
California
Fruit wt/
fruit vol
kg/hl
lb/bu
Seed wt/
fruit vol
kg/h lb/bu
/kg
Cleaned seeds/weight
Range
Average
/lb
/kg
/lb
Samples
130175
65165
3388
6080
3075
1540
155
110
66
70
50
30
2
2
13
47
36
16
12.5
64175
2980
100
45
62
48
170225
170235
25220
66110
77102
78107
11100
3050
200
203
88
88
90
92
40
40
10
2
20+
10+
Juglans
603
Table 5Juglans, walnut: stratification period, germination test conditions and results
Species
Cold
stratification
period*
(days)
J. ailantifolia
J. californica
J. cinerea
J. hindsii
J. major
J. microcarpa
J. nigra
J. regia
0
156
90120
156
120190
190
90120
30156
8+
8+
8+
30
30
30
30
30
30
20
20
20
20
20
20
42
30
5080
30+
49
3060
1540
40
54
10
68
60
58
28
14
24
Germination %
Avg
Purity
(%)
Samples
(%)
75
58
65
41
64
46
50
82
3
3+
7
4
5
7
14+
4
96
94
87
High
Days
J. californica Peat
150
J. cinerea
Sand
90120
J. hindsii
J. major
Sand
90150
or peat
J. microcarpa
J. nigra
Sand
90100
J. regia
Sand
30+
Sowing
season
Seedlings/area
/m2
/ft2
Type
Mulch
Depth
cm
in
Spring
Spring
Fall
Fall
Spring
6568
700732
5
2.55
2.55
2.5
5
2
12
12
1
2
Sawdust
None
Vermiculite
2.5
2.5
1
1
Fall
Spring
Spring
3565
3565
377700
377700
2.55
2.55
5
12
12
2
Sawdust
2.5
Sources: Brinkman (1974), Schultz and Thompson (1990),Williams and Hanks (1976).
* Outdoors during the winter or in a cold room at 1 to 5C.
Seeds were soaked in water at 88 C for 11/2 to 2 minutes before sowing.
604
Sowing
depth
cm
in
Juglans
605
References
606
Rink G. 1988. Black walnut cultivars. In: Burde EL, ed. Walnut Notes, #1.06.
St. Paul, MN: USDA Forest Service, North Central Forest Experiment
Station. 4 p.
Rink G. 1990. Juglans cinerea L., butternut. In: Burns RM, Honkala BH, tech.
coord. Silvics of North America.Volume 2, Hardwoods. Agric. Handbk.
654. Washington, DC: USDA Forest Service: 386390.
Rink G, Kung FH. 1995. Age trends in genetic control of Juglans nigra L.
height growth. In: Gottschalk KW, Fosbroke SLC, eds. 1995 March 58;
Morgantown, WV. Gen.Tech. Rep. NE-197. Radnor, PA: USDA forest
Service, Northeastern Forest Experiment Station: 247255.
Rink G, Phelps JE. 1989. Variation in heartwood and sapwood properties
among 10-year-old black walnut trees. Wood and Fiber Science 21:
177182.
Rink GH, Zhang G, Jinghua Z, Kung FH, Carroll ER. 1994. Mating parameters in Juglans nigra L. seed orchard similar to natural population estimates. Silvae Genetica 43: 261263.
Sargent CS. 1965. Manual of the trees of North America (exclusive of
Mexico). 2nd ed., corrected and reprinted. New York: Dover. 934 p.
Schultz RC,Thompson JR. 1990. Nursery practices that improve hardwood
seedling root morphology.Tree Planters Notes 41(3): 2132.
Stuke W. 1960. Seed and seed handling techniques in production of walnut
seedlings. International Plant Propagators Society Proceedings 10:
274277.
Van Dersal WR. 1938. Native woody plants of the United States: their erosion-control and wildlife values. Misc. Pub. 303. Washington, DC: USDA.
362 p.
Van Sambeek JW. 1988a. Grafting. In: Burde EL, ed. Walnut Notes, #1.05.
St. Paul: USDA Forest Service, North Central Forest Experiment
Station. 4 p.
Van Sambeek JW. 1988b. Growing containerized seedlings. In: Burde EL, ed.
Walnut Notes, #1.03. St. Paul: USDA Forest Service, North Central
Forest Experiment Station. 4 p.
Van Sambeek JW. 1988c. Nut production In: Burde EL, ed. Walnut Notes,
#4.01. St. Paul: USDA Forest Service, North Central Forest Experiment
Station. 2 p.
Van Sambeek JW, Preece JE, Lindsay TL, Gaffney GR. 1990. In vitro studies
on black walnut embryo dormancy. Northern Nut Growers Association
Annual Report 80: 55-59.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
Vozzo JA. 1978. Radiopaque agents for seed research. In: Bonner FT, ed.
Proceedings, Flowering and Seed Development in Trees: A Symposium.
1978 May 1518; Starkville, MS. Mississippi State: Mississippi State
University: 272280.
Williams RD. 1971a. Stratified walnut seed still viable after four years in
storage.Tree Planters Notes 22(4): 12.
Williams RD. 1971b. Storing black walnut seed. Northern Nut Growers
Association Annual Report 62: 8789.
Williams RD. 1990. Juglans nigra L., black walnut. In: Burns RM, Honkala BH,
tech. coords. Silvics of North America.Volume 2, Hardwoods. Agric.
Handbk. 654. Washington, DC: USDA Forest Service: 391399.
Williams RD, Hanks SH. 1976. Hardwood nurserymans guide. Agric.
Handbk. 473. Washington, DC: USDA Forest Service. 76 p.
Williams RD, Rink G, Funk DT. 1985. Planting site and genotype affect black
walnut seedling development more than nursery environment. Canadian
Journal of Forestry Research 15: 1417.
Wyman D. 1947. Seed collecting dates of woody plants. Arnoldia 7(9):
53-56.
CupressaceaeCypress family
Juniperus L.
juniper
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Genetic variation and hybridization. Junipers exhibit considerable natural variation in their growth habit and
appearance, and studies have established marked differences
in color, crown form, growth rate, and disease resistance in
eastern redcedar (Henderson and others 1979; Minckler and
Ryker 1959; Seidel and Watt 1969; Tauer and others 1987;
Van Deusen 1979), Rocky Mountain juniper (Tauer and others 1987), and western juniper (Matthews 1945). Where
ranges of the junipers overlap, natural hybridization
abounds. This condition probably explains the large number
of reported varieties of North American junipers (Dealey
1990; Fassett 1945; Hall 1952; Hall and others 1961;
Lawson 1990; Noble 1990; Ross and Duncan 1949; Vines
1960; Wilhite 1990).
Flowering and fruiting. The small, inconspicuous
flowers are borne in the spring (table 2) on the ends of short
branchlets or along the branchlets. The flowers are dioecious
or occasionally monoecious in oneseed juniper and some
sources of western juniper (Dealy 1990; Johnsen and
Alexander 1974). Pollen cones are yellow, terminal, and
about 3 to 4 mm long; ovulate cones are composed of pointed scales, 3 to 8 in number, that fuse to form a fleshy cone
6 to 8 mm long (figure 1) (Brown and Kirkman 1990). The
fleshy cones are commonly called berries. Cones are usually
greenish in color when immature and change to a bluish
black or reddish brown as they mature in the autumn (table
2). Most are covered with a conspicuous glaucous bloom.
Cones of alligator, Utah, and common junipers require 2
years to reach full maturity, but those of common juniper
may require 3 years in some parts of its range (Johnsen and
Alexander 1974; Vines 1960). Cones of the other junipers
mature in the fall of the first year (table 2). The outer skins
of the cones may be thin and resinous, as in Virginia redcedar and Rocky Mountain and oneseed junipers, or dry and
leathery or mealy, as in alligator and Utah junipers (Johnsen
and Alexander 1974).
Juiperus
607
Common name(s)
Occurrence
J. ashei Buchh.
J. sabinoides (H.B.K.) Nees
J. mexicana Spreng.
J. monticola Martinez
J. californica Carr.
California juniper,
desert white-cedar
common juniper, dwarf
juniper, prostrate juniper
J. communis L.
J. sibirica Burgsd.
J. deppeana Steud.
J. mexicana Schlecht. & Cham.
J. pachyphlaea Torr.
J. deppeana var. pachyphlaea
(Torr.) Martinez
J. monosperma (Engelm.) Sarg.
J. occidentalis var. monosperma Engelm.
(Engelm.) Cory
J. mexicana var. monosperma
J. occidentalis Hook
J. osteosperma (Torr.) Little
J. californica var. utahensis Engelm.
J. utahensis (Engelm.) Lemmon
J. pinchotii Sudworth
J. monosperma var. pinchotii
(Sudworth) Van Melle
J. texensis Van Melle
J. scopulorum Sarg.
J. scopulorum var. columnaris Fassett
alligator juniper,
checkered-bark juniper,
western juniper (lumber)
oneseed juniper,
cherrystone juniper,
redberry juniper,
west Texas juniper, sabina
western juniper,
Sierra juniper
Utah juniper, bigberry juniper,
western juniper (lumber), sabina
Pinchot juniper,
redberry juniper
Rocky Mountain juniper,
Rocky Mountain redcedar,
redcedar, river juniper
J. virginiana L.
J. viriginiana var. crebra Fern. & Grisc.
eastern redcedar,
red juniper, savin
southern redcedar,
eastern redcedar
608
Location
Flowering
Fruit ripening
Seed dispersal
J. ashei
J. communis
J. deppeana
J. monosperma
J. occidentalis
J. osteosperma
J. pinchotii
J. scopulorum
J. virginiana
J. virginiana
var. silicicola
Arizona
Oregon
Arizona
Texas
Nebraska
South Carolina
JanApr
AprMay
FebMar
MarApr
Mid-Aprmid-May
MarApr
Spring
MidAprmid-June
MidMarmid-May
JanFeb
SeptNov
AugOct
AugOct
AugSept
Mid-Sept
Sept (2nd year)
OctNov
Mid-Septmid-Dec
SeptNov
OctNov
Fallwinter
Persists for 2 yrs (2nd3rd yr)
Persists for 2 seasons (2nd yr)
OctNov (persists 12 yrs)
Persists for 2 yrs
Persists for 2 yrs
Year-round
October (persists 23 yrs)
FebMar (1st yr)
Juniperus
609
Species
Height
Year
at maturity
first
Seeds/
(m)
cultivated cone
Years
between large
seedcrops
36
15
115
320
1925
1560
1873
12
12
13
24
Irregular
J. monosperma
38
1900
12
25
J. occidentalis
J. osteosperma
J. pinchotii
59
512
15
1840
1900
23
12
1
Green-blue
Green glaucous
Green with light bloom
J. scopulorum
J. virginiana
J. virginiana
var. silicicola
615
930
7
1936
1664
12
12
12
25
23
Green with
Green
Green
J. ashei
J. californica
J. communis
J. deppeana
Green
Bluish w/dense bloom
Red
Green
Deep blue
Reddish brown
Bluish to black, glaucous
Bluish to reddish brown,
glaucous
Copper to dark blue with
white waxy bloom
Bluish black, glaucous
Reddish brown,
Copper to red to reddish
brown
Blue w/white waxy bloom
Blue
Dark blue
610
Species
J. ashei
J. communis
J. deppeana
J. monosperma
J. occidentalis
J. osteosperma
J. pinchotii
J. scopulorum
J. virginiana
Place
collected
Arizona
Arizona & New Mexico
Oregon
Arizona
Sonora & Texas
Arizona
Great Plains
/kg
Cleaned seeds/weight
Range
/lb
56,120120,170
19,84034,400
33,65044,100
17,64034,970
7,94015,660
21,28030,650
39,36092,830
81,580121,270
25,45054,500
9,00015,600
15,26020,000
8,00015,860
3,6007,100
9,65013,900
17,85042,100
37,00055,000
Average
/kg
22,270
80,480
28,270
40,350
27,120
10,910
24,230
59,760
96,140
/lb
Samples
10,100
36,500
12,820
18,300
12,300
4,950
10,990
27,100
43,600
1
8
5
10
15
2
36
34
Sources: Johnsen and Alexander (1974), Stoeckler and Slabaugh (1965),Vines (1960).
Juniperus
611
Species
J. ashei
J. communis
J. deppeana
J. monosperma
J. osteosperma
J. pinchotii
J. scopulorum
J. virginiana
Stratification
period (days)
Warm*
Cold
0
0
6090
0
0
0
120
0
30
120
0
4
0
120
120
90+
0
3060
0
120
0
60
120
30120
90
45
8
8
Sand
Sand
Paper, sand
Paper, sand
Sand, peat
Sand, peat, soil
Sand, soil
Perlite
Perlite
Paper, sand
Paper, sand
Perlite
Kimpak
86
50
86
86
86
86
86
60
60
86
50
58
60
68
50
68
68
68
68
68
60
60
68
77
58
60
Germination
rate
Days
%
Germination
percentage
Days
%
60
60
2030
40
3040
3070
70
36+
2030
2030
60
66
10
29
815
924
30
43
30
27
531
674
84
70
Sources: Cotrufo (1963), Johnsen (1959), Johnsen and Alexander (1974), Meagher (1943), Riffle and Springfield (1968).
* 30 to 20 C alternated diurnally.
5 C.
Seeds soaked in sulfuric acid 45 minutes.
Seeds soaked in 1% citric acid for 4 days.
612
33
36
775
16-30
45
2075
849
63
53
22
76
87
78
Samples
1
1
10+
2
1
34
8
4
4
7
16
3
2
References
Adams RP. 1987. Investigation of Juniperus species of the United States for
new sources of cedarwood oil. Economic Botany 41: 4854.
Afanasiev M, Engstrom A, Johnson EW. 1959. Effects of planting dates and
storage on survival of eastern redcedar in central and western
Oklahoma. Sta. Bull. B-527. Stillwater: Oklahoma Agricultural
Experiment Station. 19 p. [Forestry Abstracts 21(1): 445; 1960].
Barton LV. 1951. Germination of seeds of Juniperus virginiana L.
Contributions of the Boyce Thompson Institute 16(8): 387393
[Forestry Abstracts 13(4): 2925; 1952].
Benson DA. 1976. Stratification of Juniperus scopulorum.Tree Planters
Notes 27(2): 11, 23.
Brown CL, Kirkman LK. 1990. Trees of Georgia and adjacent states.
Portland, OR:Timber Press. 292 p.
Burkhardt JW,Tisdale EW. 1976. Causes of juniper invasion in southwestern Idaho. Ecology 57: 472484.
Chavez-Ramirez F, Slack RD. 1994. Effects of avian foraging and post-foraging behavior on seed dispersal patterns of Ashe juniper. Oikos 71:
4046.
Cotrufo C. 1963. Stimulation by citric acid of germination of eastern redcedar (Juniperus virginiana L.). Nature 199: 9293.
Dealy JE. 1990. Juniperus occidentalis Hook., western juniper. In: Burns RM,
Honkala BH, tech. coords. Silvics of North America.Volume 1, Conifers.
Agric. Handbk. 654. Washington, DC: USDA Forest Service: 109115.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant
propagation: from seed to tissue culture. Athens, GA:Varsity Press.
239 p.
Djavanshir K, Fechner GH. 1976. Epicotyl and hypocotyl germination of
eastern redcedar and Rocky Mountain juniper. Forest Science 22:
261266.
Edson JL, Wenny DL, Dumroese RK, Leege-Brusven A. 1996. Mass propagation of Rocky Mountain juniper from shoot cuttings.Tree Planters
Notes 47(3): 9499.
Juniperus
613
Johnsen TN Jr, Alexander RA. 1974. Juniperus L., juniper. In: Schopmeyer CS,
tech. coord. Seeds of woody plants in the United States. Agric. Handbk.
450. Washington, DC: USDA Forest Service: 460469.
Jones L. 1962. Recommendations for successful storage of tree seed.Tree
Planters Notes 55: 920.
Lawson, ER. 1990. Juniperus virginiana L., eastern redcedar. In: Burns RM,
Honkala BH, tech. coords. Silvics of North America.Volume 1, Conifers.
Agric. Handbk. 654. Washington, DC: USDA Forest Service: 131140.
Little EL Jr. 1971. Atlas of United States trees.Volume 1, Conifers and
important hardwoods. Misc. Pub. 1146. Washington, DC: USDA Forest
Service. 209.
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Livingston RB. 1972. Influence of birds, stones and soil on the establishment
of pasture juniper, Juniperus communis, and red cedar, J. virginiana, on
New England pastures. Ecology 53: 11411147.
McPherson GR, Wright HA. 1990. Establishment of Juniperus pinchotii in
western Texas: environmental effects. Journal of Arid Environments 19:
283287 [Forestry Abstracts 52(9): 6566; 1991].
Matthews OV. 1945. The Robinson juniper. Journal of Forestry 43: 755756.
Meagher GS. 1943. Reaction of pinyon and juniper seedlings to artificial
shade and supplemental watering. Journal of Forestry 41: 480482.
Meines MK. 1965. Juniper germination simplified.Tree Planters Notes 70:
67.
Minckler LS, Ryker RA. 1959. Color, form, and growth variations in eastern
redcedar. Journal of Forestry 57: 347349.
Noble DL. 1990. Juniperus scopulorum Sarg., Rocky Mountain juniper. In:
Burns RM, Honkala BH, tech. coords. Silvics of North America.Volume 1,
Conifers. Agric. Handbk. 654. Washington, DC: USDA Forest Service:
116126.
Otta JD, Fiedler DJ, Lengkeek VH. 1980. Effect of benomyl on Phomopsis
juniperovora infection of Juniperus virginiana. Phytopathology 70: 4650.
Peterson GW. 1973. Infection of Juniperus virginiana and J. scopulorum by
Phomopsis juniperovora. Phytopathology 63: 246251.
Peterson GW. 1977. Epidemiology and control of a blight of Juniperus virginiana caused by Cercospora sequoiae var. juniperi. Phytopathology 67:
234238.
Peterson GW, Wysong DS. 1968. Cercospora blight of junipers: damage and
control. Plant Disease Reporter 52: 361362.
Rehder A. 1956. Manual of cultivated trees and shrubs hardy in North
America. 2nd ed. New York: Macmillan. 996 p.
Rietveld WJ. 1989. Variable seed dormancy in Rocky Mountain juniper. In:
Landis TD, tech. coord. Proceedings, Intermountain Forest Nursery
Association. 1989 August 1418; Bismark, ND. Gen.Tech. Rep. RM-184
Fort Collins, CO: USDA Forest Service, Rocky Mountain Forest and
Range Experiment Station: 6064.
614
EricaceaeHeath family
Kalmia latifolia L.
mountain-laurel
Frank A. Blazich and Mark C. Starrett
Dr. Blazich is alumni distinguished graduate professor of plant propagation and tissue culture at North Carolina
State Universitys Department of Horticultural Science, Raleigh, North Carolina; Dr. Starrett is
associate professor at the University of Vermonts Department of Plant and Soil Science, Burlington,Vermont
Kalmia
615
616
seeds.
longitudinal
References
Blazich FA. 1996. Unpublished data. Raleigh: North Carolina State
University.
Bridwell FM. 1994. Landscape plants: their identification, culture, and use.
Albany, NY: Delmar Publishers. 560 p.
Dirr MA. 1990. Manual of woody landscape plants: their identification,
ornamental characteristics, culture, propagation and uses. 4th ed.
Champaign, IL: Stipes Publishing Co. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1632 p.
Fordham AJ. 1960. Propagation of woody plants by seed. Arnoldia 20(6):
3340.
Fryxell PA. 1957. Mode of reproduction in higher plants. Botanical Review
23: 135233.
Glenn CT, Blazich FA, Warren SL. 1998. Influence of storage temperatures
on long-term seed viability of selected ericaceous species. Journal of
Environmental Horticulture 16(3): 166172.
Jaynes RA. 1971. Seed germination of six Kalmia species. Journal of the
American Society for Horticultural Science 96: 668672.
Jaynes RA. 1997. Kalmia, mountain laurel and related species. Portland, OR:
Timber Press. 295 p.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. 3rd ed.
New York : Macmillan. 1290 p.
Kalmia
617
AraliaceaeGinseng family
cleaned
Pregermination treatments. Under natural conditions, castor-aralia seeds require a 2-year germination period
(Sato 1998). Dormancy of the seed is caused by neutral
(coumarin) and acid (abscisic) inhibitors present in the seedcoat and endosperm, and by an impermeable seedcoat (Dirr
1990; Huang 1987a&b). Warm temperatures of 15 to 25 C
for 3 to 5 months followed by cold stratification at 0 to 5 C
for 2 to 3 months will overcome seed dormancy and give
reasonable germination (Dirr and Heuser 1987; Huang 1986,
1987b; Sato 1998; Xu and Han 1988). Soaking the seeds in
sulfuric acid for 30 minutes will substitute for the warm
stratification period (Dirr and Heuser 1987; Rudolf 1974).
Germination tests. Tests in germinators or sand flats
for 60 days is suggested (Rudolf 1974).
Nursery practice and seedling care. Fresh seeds that
have been cleaned and dried can be sown in the fall but will
not germinate for 2 years (Dirr and Heuser 1987; Satoo
1992). Stratified seeds should be sown in the spring (Rudolf
1974). The seeds should be sown in well-prepared beds at a
rate of 1,760 to 3,300/m2 (164 to 307/ft2) to give 200 to 300
seedlings/m2 (19 to 28/ft2) (Satoo 1992). Castor-aralia can
be propagated by root cuttings (Dirr and Heuser 1987;
Macdonald 1986). Root cuttings, 7.6 to 10.2 cm (3 to 4
inches) in length, should be dug soon after frost and then
placed upright (proximal end) in a medium kept in a cool
References
Dirr MA. 1990. Manual of woody landscape plants: their identification,
ornamental characteristics, culture, propagation, and uses. Champaign, IL:
Stipes Publishing. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Hillier Nurseries (Winchester) Ltd. 1991. The Hillier manual of trees and
shrubs. Melksham, Wiltshire, UK: Redwood Press. 704 p.
Huang YG. 1986. Study on embryonic dormancy of Kalopanax septemlobus
seeds [in Chinese; summary in English]. Journal of North-East Forestry
UniversityChina 14(1): 3944.
Huang YG. 1987a. A preliminary study on neutral and acid inhibitors in the
seed of Kalopanax septemlobus [in Chinese; summary in English]. Acta
Botanica Sinica 29(3): 283292.
Huang YG. 1987b. Inhibiting substances play a role in dormancy of
Kalopanax septemlobus seeds [in Chinese; summary in English]. Journal of
North-East Forestry UniversityChina 15(2): 1825.
KRRT [Korea Republic, Research Team on the Wood Properties of Native
Hardwoods]. 1987. Studies on the wood properties of native hardwoods of major importance: 4. Wood properties of 2 species of genus
Acer, 2 species of Prunus and 1 species of genus Kalopanax. Research
Reports of the Forestry Research InstituteSeoul, Korea 34: 93109.
Krssmann G. 1984. Manual of cultivated broad-leaved trees and shrubs.
Volume 2, EPro. Beaverton, OR:Timber Press. 445 p.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan. 1290 p.
Liu GP, Sun JH, Lin GP, Sun JH. 1998. Analysis of nutrient elements of
Kalopanax septembolus [in Chinese; English summary]. Journal of
Northeast Forestry University 26(3): 6567.
McDonald B. 1986. Practical woody plant propagation for nursery growers.
Portland, OR: timber Press. 669 p.
Ohashi H. 1994. Nomenclature of Kalopanax septemlobus (Thunberg ex
Murray) Koidzumi and classification of its intraspecific taxa (Araliaceae).
Journal of Japanese Botany 69 (1): 2831.
Porzel A, Sung TV, Schmidt J, Lischewski M, Adam G. 1992. Studies on the
chemical constituents of Kalopanax septemlobus. Planta Medica 58 (5):
481482.
Kalopanax 619
ChenopodiaceaeGoosefoot family
Kochia Roth
kochia
Stanley G. Kitchen and Stephen B. Monsen
Mr. Kitchen and Mr. Monsen are botanists at the USDA Forest Services Rocky Mountain Research Station,
Shrub Sciences Laboratory, Provo, Utah
Growth habit, occurrence, and use. Two woodybased non-rhizomatous sub-shrub species of kochias are
found in the western United States. The widely distributed
native, gray molly, and its introduced and closely related
counterpart (Blackwell and others 1978), forage kochia, are
found in salt-desert, sagebrush, pinyonjuniper, and dry
mountain brush communities (table 1). Erect to steeply
ascending annual stems growing from a woody base and
long-lived prostrate branches attain heights of 5 to 50 cm for
gray molly (Welch and others 1987) and 10 to 100 cm for
forage kochia (Baylan 1972).
Each species provides nutritious winter forage for livestock (Baylan 1972; Blauer and others 1976). Variability in
preference by livestock (Shishkin 1936) and mule deer
(Odocoileus hemionis) (Davis and Welch 1985) has been
observed among ecotypes of forage kochia.
Both species have potential for use in revegetation of
saline and alkaline soils on arid and semi-arid sites (Blauer
and others 1976; Clarke and West 1969; Francois 1976;
Romo and Haferkamp 1987) and forage kochia has been
used successfully in stabilizing mine spoils (Frischknecht
and Ferguson 1984). Perhaps the greatest potential use of
forage kochia is in providing cover and forage on degraded
western cold-deserts (McArthur and others 1974; Monsen
and Turnipseed 1990; Pendleton and others 1992). Where
established, it effectively competes with weeds such as
Common name(s)
Occurrence
K. americana S.Wats.
K. vestita (S.Wats.) Rydb.
K. americana var. vestita S.Wats.
K. prostrata (L.) Schrad.
K. suffruticulosa Lessing
Salsola prostrata L.
620
virescens), has resulted from this research (Stevens and others 1995). Immigrant has been planted on several thousand
acres in Utah, Idaho, and surrounding states. Small plantings
of other accessions (ssp. grisea) also exist (Blauer and others 1993; McArthur and others 1990; Monsen and Kitchen
1995; Monsen and Turnipseed 1990).
Flowering and fruiting. Kochia flowering structures
are described as one to several, mostly perfect, inconspicuous, sessile flowers occurring in axils of foliose bracts
(Welsh and others 1987). Stems are potentially floriferous
throughout (Blackwell and others 1978). Flowering is indeterminate from May to August for gray molly and from July
to September for forage kochia (Shishkin 1936). The fruit is
a 1-seeded utricle that is enclosed in a thin, fragile perianth
(figure 1). The perianth is horizontally winged at maturity.
Perianth pubescence for forage kochia is highly variable
(Baylan 1972). Seeds are oval to orbicular in shape and 1 to
2 mm in diameter. The embryo is bent into roughly the
shape of a horseshoe, a common configuration for this family (figure 2).
Fruit collection and cleaning. Fruits of forage kochia
ripen from September to November (Baylan 1972) whereas
those of gray molly are generally ripe by mid-October.
Fruits are easily dislodged when fully ripened and dry. They
are hand-harvested by stripping individual stems or by beating seeds into a hand-held hopper with a badminton racket
or similar device. Mechanical harvest techniques for forage
kochia seeds include mowing stems just before fruits are
ready to shatter, drying, and combining. Vehicle-mounted
mechanical sweepers are also used to harvest fully ripened
fruits from solid stands (Stevenson 1995). Although harvesting the fruits before the seeds are fully ripened can reduce
losses to shattering, it also results in lower seed viability
Figure 1Kochia prostrata, forage kochia:
perianth.
fruits in
longitudinal
Kochia
621
Table 2Kochia prostrata, forage kochia: mean germination times as affected by temperature of dry storage
Accession*
Fresh
314929
343101
356818
356826
358941
Mean
72
51
53
108
81
71
a
a
a
a
a
a
60 a
46 a
51 a
86 b
63 b
61 a
5C
67 a
55 a
58 a
90 b
76 a
69 a
622
References
Baylan GA. 1972. Prostrate summer cypress and its culture in Kirghizia.
Isdatelstvo, Frunze, Kirghizistan [translated from Russian. 1979.
Washington, DC: USDA and National Science Foundation].
Blackwell WH, Baechle MD, Williamson G. 1978. Synopsis of Kochia
(Chenopodiaceae) in North America. SIDA 7: 248254.
Blauer AC, McArthur ED, Stevens R, Nelson, SD. 1993. Evaluation of roadside stabilization and beautification plantings in south-central Utah. Res.
Pap. INT-462. Ogden, UT: USDA Forest Service, Intermountain Research
Station. 65 p.
Blauer AC, Plummer AP, McArthur ED, Stevens R, Giunta BC. 1976.
Characteristics and hybridization of important Intermountain shrubs: 2.
Chenopod family. Res. Pap. INT-177. Ogden, UT: USDA Forest Service,
Intermountain Research Station. 42 p.
Clarke LD, West NE. 1969. Germination of Kochia americana in relation to
salinity. Journal of Range Management 22: 286287.
Davis JN, Welch BL. 1985. Winter preference, nutritive value and other
range use characteristics of Kochia prostrata (L.) Schrad. Great Basin
Naturalist 45: 778783.
Francois, LE. 1976. Salt tolerance of prostrate summer cypress (Kochia
prostrata). Agronomy Journal 68: 455456.
Frischknecht NC, Ferguson RB. 1984. Performance of Chenopodiaceae
species on processed oil shale. In:Tiedemann AR, McArthur ED, Stutz
HC, Stevens R, Johnson KL, comp. Proceedings, Symposium on the
Biology of Atriplex and Related Chenopods; 1983 May 26; Provo, UT.
Gen Tech. Rep. INT-172. Ogden, UT: USDA Forest Service,
Intermountain Research Station: 293297.
Haferkamp MR, Ganskopp DC, Marietta KL, Knapp BW. 1990.
Environmental influences on germination of utricles and seedling establishment of Immigrant forage kochia. Journal of Range Management 43:
518522.
Jorgensen KR, Davis JN. 1984. A technique for retaining seed viability in
Kochia prostrata. In:Tiedemann AR, McArthur ED, Stutz HC, Stevens R,
Johnson KL, comps. Proceedings, Symposium on the Biology of Atriplex
and Related Chenopods; 1983 May 26; Provo, UT. Gen Tech. Rep. INT172. Ogden, UT: USDA Forest Service, Intermountain Research Station:
166167.
Kitchen SG, Monsen SB. 1999. Unpublished data. Provo, UT: USDA Forest
Service, Rocky Mountain Research Station.
McArthur ED, Blauer AC, Stevens R. 1990. Forage kochia competition with
cheatgrass in central Utah. In: McArthur ED, Romney EM, Smith SD,
Tueller PT, comps. Proceedings, Symposium on Cheatgrass Invasion,
Shrub Dieoff, and Other Aspects of Shrub Biology and Management;
1989 April 57; Las Vegas, NV. Gen.Tech. Rep. INT-276. Ogden, UT:
USDA Forest Service, Intermountain Research Station: 5665.
McArthur ED, Guinta BC, Plummer AP. 1974. Shrubs for restoration of
depleted ranges and disturbed areas. Utah Science 34: 2833.
Monsen SB,Turnipseed D. 1990. Seeding forage kochia into cheatgrass
infested ranges. In: McArthur ED, Romney EM, Smith SD,Tueller PT,
comps. Proceedings, Symposium on Cheatgrass Invasion, Shrub Dieoff,
and Other Aspects of Shrub Biology and Management; 1989 April 57.
Las Vegas, NV. Gen.Tech. Rep. INT-276. Ogden, UT: USDA Forest
Service, Intermountain Research Station: 6671.
Pendleton RL, Frischknecht NC, McArthur ED. 1992. Long-term survival of
20 selected plant accessions in a Rush Valley, Utah, planting. Res. Note
INT-403. Ogden, UT: USDA Forest Service, Intermountain Research
Station: 7 p.
Pope CL, McArthur ED. 1977. IOBP chromosome number reports: 55.
Chenopodiaceae.Taxon 26: 109.
Romo JT, Haferkamp MR. 1987. Forage kochia germination response to
temperature, water stress, and specific ions. Agronomy Journal 79: 2730.
Shishkin BK, ed. 1936. Flora of the U.S.S.R.Volume 6, Centrospermae.
Moscow: Isdatelstvo Akademii Nauk SSSR [translated from Russian.
1970. Washington, DC: Smithsonian Institution and National Science
Foundation]. 731 p.
Stevens R,Van Epps GA. 1984. Seeding techniques to improve establishment of forage kochia [Kochia prostrata (L.) Schrad.] and fourwing saltbush [Atriplex canescens (Pursh) Nutt.]. In:Tiedemann AR, McArthur ED,
Stutz HC, Stevens R, Johnson KL. comps. Proceedings, Symposium on the
Biology of Atriplex and Related Chenopods; 1983 May 26; Provo, UT.
Gen.Tech. Rep. INT-172. Ogden, UT: USDA Forest Service,
Intermountain Research Station: 269272.
Stevens R, Jorgensen KR, McArthur ED, Davis JN. 1985. Immigrant forage
kochia. Rangelands 7: 2223.
Stevenson R. 1995. Personal communication. Ephraim, UT: Stevenson
Intermountain Seed.
Stewart A, Anderson VJ, Kitchen SG, 1999. Immigrant forage kochia (Kochia
prostrata) seed germination as affected by storage conditions. In:
McArthur ED, Ostler WK, Wambolt CL, comps. Proceedings, Shrubland
Ecotones; 1998 August 1214; Ephraim, UT. RMRS-P11. Ogden, UT:
USDA Forest Service, Rocky Mountain Research Station: 274279.
Waller SS, Britton CM, Schmidt DK, Stubbendieck J, Sneva FA. 1983.
Germination characteristics of two varieties of Kochia prostrata (L.)
Schrad. Journal Range Management 36: 242245.
Welsh SL, Atwood ND, Higgins LC, Goodrich S. 1987. A Utah flora. Great
Basin Naturalist Memoirs 9: 1894.
Young JA, Evans RA, Stevens R, Everett RL. 1981. Germination of Kochia
prostrata seed. Agronomy Journal 73: 957961.
Kochia
623
SapindaceaeSoapberry family
624
magnetic field of 100 gauss for 4.3 seconds increased germination after scarification from 56 to 97% (Maronek 1975).
Germination tests. Germination is epigeal (figure 3).
Germination should be tested in sand flats or germinators
for 5 to 10 days at 20 (night) to 30 C (day), using 200 to
400 seeds that were acid treated and then stratified for each
test. One test of untreated seeds gave a germination rate of
only 2% in 29 days, whereas seeds of the same sample gave
52% after the acid plus stratification treatment recommended above (Rudolf 1974). In another test, 74% of untreated
seeds germinated in 54 days, compared with 91% of
mechanically scarified seeds in 23 days (Zentsch and Kaul
1968). Official seed testing agencies recommend tetrazolium
staining for germination tests of panicled golden raintree.
The suggested procedure is to soak the seeds in water for 18
hours, then remove the seedcoat before staining for 24 hours
at 30 C in a 1% solution (ISTA 1993).
Nursery practice. Untreated seeds may be sown in
the fall and scarified seeds can be sown in the spring (some
seedlots may require stratification after scarification) and
covered with 6 to 13 mm (1/4 to 1/2 in) of soil. Seedlots
sown immediately after collection in fall usually give reasonably good results (Swingle 1939). A target bed density is
about 300 to 315 seedlings/m2 (30/ft2). Tree survival is
about 70% (Jack 1969). Seedlings should be lifted as 2+0
stock (Jack 1969).
This species should be planted only in sunny locations,
but it is not particular as to soil type (Bailey 1939). It may
also be propagated by layers, cuttings, or root cuttings
(Bailey 1939).
References
Bailey LH. 1939. The standard cyclopedia of horticulture. New York:
Macmillan. 3639 p.
Garner JL. 1979. Overcoming double dormancy in golden-rain tree seeds.
Plant Propagator 25(2): 68.
Garner JL, Lewis AJ. 1980. An evaluation of techniques used for germinating
goldenrain tree seeds. American Nurseryman 151(8): 12.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing: rules 1993. Seed Science and Technology 21 (Suppl.):
1259.
Jack RA. 1969. Personal communication. Silverton, OR: Silver Falls Nursery.
Krssmann G. 1960. Handbuch der Laubgehlze.Volume 1. 495 p.;
Volume 2. 608 p.
Maronek DM. 1975. Electromagnetic seed treatment increased germination
of Koelreuteria paniculata Laxm. HortScience 10(3): 227228.
Ohwi J. 1965. Flora of Japan. Washington, DC: Smithsonian Institution.
1067 p.
Pammel LH, King CM. 1930. Germination and seedling forms of some
woody plants. Proceedings of the Iowa Academy of Science 37:
131141.
Koelreuteria
625
ChenopodiaceaeGoosefoot family
626
fruiting
cleaned
year and produce abundant crops annually (table 1). A 10year-old stand has produced 78 to 90 kg/ha (70 to 80 lb/ac)
of fruit
(diaspores) (Springfield 1974b). Good seed quality depends
on the mother plants maintaining transpiration rates during
seed and diaspore development (Booth 1990a).
Seed collection and storage. Seeds are harvested by
stripping the diaspores from the bushes or by cutting and
drying the fruiting spikes. Harvest time is mid-September in
Saskatchewan (Romo 1995) to mid-October or early
November at lower latitudes (Strickler 1956; Wasser 1945;
Wilson 1931). Mechanized harvest methods have been tried
(Springfield 1974b), but most collectors have found it more
efficient to hand-harvest. However, Majerus (2003)
described harvesting winterfat seeds with a combine. Dry
diaspores should be stored without threshing or other processing to prevent accelerated aging. Harvested material will
contain unfilled diaspores, but there are no practical methods for separating these from the germinable diaspores
(Allen and others 1987). Percentage diaspore fill may be
determined by threshing small samples. This is quickly done
using equipment described by Booth and Griffith (1984).
sectional
Krascheninnikovia
627
Table 1 Krascheninnikovia lanata, winterfat: diaspore weights by source and harvest year
Diaspores/weight
Source
Collections
Colorado
New Mexico
Nevada
Saskatchewan
Utah
Wyoming
Total
3
3
1
1
2
2
12
Years
harvested
1982, 84, 94
1984
1983
1994
1982, 84
1994
Mean*
Range
/g
/oz
147
231
175
147
212
177
5.2
8.1
6.2
5.2
7.5
6.2
/g
137203
208270
175
147
167257
173181
/oz
4.87.1
7.39.5
6.2
5.2
5.99.0
6.16.4
Germination is most suitably tested by imbibing diaspores at 0 to 5 C for 4 or 5 days followed by incubation at
15 C. A longer cold treatment, 6 to 15 days, may increase
the germination rate and seedling vigor for some seedlots,
especially those that are less than 3 months or more than 12
months after harvest. Seeds less than 3 months from harvest
may require after-ripening (Springfield 1972). Germination
is not affected by light (Hilton 1941) and is rapid at warm
temperatures.
Nursery and field practice. In the past, it was considered important to thresh the seed from the diaspore to
simplify seeding with mechanized equipment (Springfield
1974b). However, that practice is no longer recommended
because the bracts aid in seedling establishment, and threshing damages the seeds (Booth 1984, 1989a&b, 1990b;
Booth and Schuman 1983). Broadcasting diaspores results
in good establishment in depressions, in litter, and in protected sites (Stevens and others 1977). Diaspores can also be
sown with a cultipacker (Luke and Monsen 1984), with a
hydroseeder (Pellant and Reichert 1984), as pelleted diaspores, and in seed tapes (Booth 1987a&b). Use of the casedhole punch seeder (Booth 1995) is effective and allows diaspores to be sown through fabric mulch. Natural establishment occurs with cool temperatures and high surface moisture and with a mat of diaspores on the soil surface (figure
3) (Booth 1987b, 1989a, 1990b; Gasto 1969; Wilson 1931;
Woodmansee and Potter 1971). Fall-seeding is recommended (Zabek and Romo 1998). Under-snow germination produces vigorous seedlings and contributes to seeding success.
Winterfat seeds and seedlings can show freeze-tolerance
(Bai and others 1997a; Booth 1987b, 1989a; Hilton 1941;
Stricker 1956; Woodmansee and Potter 1971), but reduced
germination or loss of seedlings can also occur (Bai and oth-
628
References
Allen PS, Meyer SE, Davis TD. 1987. Determining seed quality of winterfat
[Ceratoides lanata (Pursh) J.T. Howell]. Journal of Seed Technology 11:
714.
Asay KH. 1959. A study of Eurotia lanata [MS thesis]. Laramie: University of
Wyoming. 34 p.
Agustrina R. 1995. The influence of imbibition temperature on germination,
mitochondria structures, and proteins of winterfat (Ceratoides lanata
(Pursh) J.T. Howell) [PhD dissertation]. Laramie: University of Wyoming.
125 p.
Bai Y, Booth DT, Romo JT. 1998a. Low temperature exotherm did not mark
the death of hydrated winterfat (Eurotia lanata (Pursh) Moq.) seeds.
Annals of Botany 81: 595602.
Bai Y, Booth DT, Romo JT. 1998b. Developmental stages of winterfat germinants related to survival after freezing. Journal of Range Management 51:
709713.
Bai Y, Booth DT, Romo JT. 1999. Imbibition temperature affects winterfat
(Eurotia lanata (Pursh) Moq.) seed hydration and cold-hardiness
response. Journal of Range Management 52: 271174.
Booth DT. 1984. Threshing damage to radicle apex affects geotrophic
response of winterfat. Journal of Range Management 37: 222225.
Booth DT. 1987a. Diaspores of rangeland plants: ecology and management.
In: Frasier GW, Evans RA, eds. Proceedings, Symposium on Seed and
Seedbed Ecology of Rangeland Plants; 1987 April 2123;Tucson, AZ.
Washington, DC: USDA Agricultural Research Service: 202211.
Booth DT. 1987b. Contributions to the reproductive autecology of winterfat [Eurotia lanata (Pursh) Moq] with notes on direct seeding methods
[PhD dissertation]. Laramie: University of Wyoming. 187 p.
Booth DT. 1988. Winterfat diaspore morphology. Journal of Range
Management 41:351-337.
Booth DT. 1989a. A model of freeze tolerance in winterfat germinants. In:
Wallace A, Haferkamp MR, McArthur ED, comps. Proceedings,
Symposium on Shrub Ecophysiology and Biotechnology; 1987 June
30July 2; Logan, UT. Gen.Tech. Rep. INT-256. Ogden, UT: USDA Forest
Service, Intermountain Research Station: 8389.
Booth DT. 1989b. Seedbed ecology of winterfat: cations in diaspore bracts
and their effect on germination and early plant growth. Journal of Range
Management 42: 178182.
Booth DT. 1990a. Seedbed ecology of winterfat: effects of mother-plant
transpiration, wind stress and nutrition on seedling vigor. Journal of
Range Management 43: 2024.
Booth DT. 1990b. Plant diaspore functions. Journal of Seed Technology 14:
6173.
Booth DT. 1992. Seedbed ecology of winterfat: imbibition temperature
affects post-germination growth. Journal of Range Management 45:
159164.
Booth DT. 1994. Unpublished data. Cheyenne, WY: USDA Agricultural
Research Service.
Booth DT. 1995. Cased-hole punch seeder: a tool for increasing plant
diversity. Abstracts, 48th Annual Meeting of the Society for Range
Management; 1995 January 1420; Phoenix, AZ: 8.
Booth DT, Griffith LW. 1984. Evaluation of air threshing for small lots of
winterfat fruits. Journal of Range Management 37: 286287.
Booth DT, Haferkamp MR. 1995. Morphology and seedling establishment.
In: Sosebee R, Bedunah DJ, eds. Management of grazing lands: importance of plant morphology and physiology to individual plant and community response. Denver: Society for Range Management. 239290.
Booth DT, Schuman GE. 1983. Seedbed ecology of winterfat: fruits versus
threshed seeds. Journal of Range Management 36: 387390.
Booth DT, McDonald MB. 1994. Cation concentration in post-imbibed
winterfat seeds as influenced by imbibition temperature. Journal of
Range Management 47: 485488.
Krascheninnikovia
629
Pursh F. 1814. Flora Americae Septentrionalis. 2nd ed. London: James Black,
and Son. 751 p.
Riedl WA, Asay KH, Nelson JL, Delwar GM. 1964. Studies of Eurotia lanata
(winterfat). Bull. 425. Laramie: Wyoming Agricultural Experiment Station.
18 p.
Romo JT. 1995. Personal communication. Saskatoon: University of
Saskatchewan.
Shaw, N, Monsen, SB. 1984. Nursery propagation and outplanting of bareroot chenopod seedlings. In:Tiedemann AR, and others, comps.
Proceedings, Symposium on the Biology of Atriplex and Related
Chenopods; 1983 May 26; Ogden, UT. Gen.Tech. Rep. INT-172. Ogden,
UT: USDA Forest Service, Intermountain Forest and Range Experiment
Station: 251260.
Springfield HW. 1968a. Germination of winterfat seeds under different
moisture stresses and temperatures. Journal of Range Management 21:
314316.
Springfield HW. 1968b. Age and year of collection affect germination of
winterfat seeds. Res. Note RM-112. Fort Collins, CO: USDA Forest
Service, Rocky Mountain Forest and Range Experiment Station. 2 p.
Springfield HW. 1968c. Cold storage helps winterfat seeds retain viability.
Journal of Range Management 21: 401402.
Springfield HW. 1972. Winterfat seeds undergo after-ripening. Journal of
Range Management 25: 479480.
Springfield HW. 1973. Winterfat fruits and seeds retain high viability 3 years
in cold storage. Res. Note RM-233. Fort Collins, CO: USDA FS, Rocky
Mountain Forest and Range Experiment Station. 3 p.
Springfield HW. 1974a. Winterat seeds viable after 8 years refrigerated
storage. Journal of Range Management 27: 78.
Springfield HW. 1974b. Eurotia lanata (Pursh) Moq., winterfat. In:
Schopmeyer, CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service:
398400.
Strickler GSS. 1956. Factors affecting the growth of white-sage (Eurotia
lanata (Pursh) Moq.) [MS thesis]. City: University of Nevada. 125 p.
630
Stevens R, Monsen SB. 1988. Hatch winterfat: a quality shrub for ranges
and wildlands. Rangelands 10: 104105.
Stevens R, Giunta BC, Jorgensen KR, Plummer AP. 1977. Winterfat
(Ceratoides lanata). Pub. 77-2. [Salt Lake City]: Utah State Division of
Wildlife Resources. 41 p.
Stutz H. 1995. Personal communication. Provo, UT: Brigham Young
University.
Tutin TG, Heywood VH, Burgess NA,Valentine DH, Walters SM, Webb DA,
Ball PW, Chater AO, eds. 1964. Flora Europaea.Volume 1. Cambridge
UK: Cambridge University Press. 464 p.
Wasser CH. 1945. High protein content makes winterfat valuable forage
for Colorado ranges. Colorado Farm Bulletin 7: 67, 13.
Welsh SL. 1974. Andersons flora of Alaska. Provo, UT: Brigham Young
University Press. 724 p.
Wiersema J. 2000. Personal communication. Beltsville, MD: USDA
Agricultural Research Service.
Wilson CP. 1931. The artificial reseeding of New Mexico ranges. Bull. 189.
New Mexico Agricultural Experiment Station.
Woodmansee RG, Potter LD. 1971. Natural reproduction of winterfat
(Eurotia lanata) in New Mexico. Journal of Range Management 24:
2430.
Workman JP, West NE. 1967. Germination of Eurotia lanata in relation to
temperature and salinity. Ecology 48: 659661.
Workman JP, West NE. 1969. Ecotypic variation of Eurotia lanata populations in Utah. Botanical Gazette 130: 2635.
Zabek C, Romo JT. 1995. Seed-seedbed ecology and establishment of winterfat. In: Romo JT, project leader. Seedbed requirements and cold tolerance of winterfat seedlings: a adapted forage for the Canadian Prairies.
Prog. Rep. 3. Saskatoon: University of Saskatchewan, Department of
Crop Science and Plant Ecology: 715.
FabaceaePea family
Laburnum Medik.
laburnum
Paula M. Pijut
Dr. Pijut is a research plant physiologist at the USDA Forest Services North Central Research Station,
Hardwood Tree Improvement and Regeneration Laboratory,West Lafayette, Indiana
Table 1Laburnum, laburnum: nomenclature, occurrence, growth habit, height at maturity, and date of first cultivation
Scientific name
& synonym(s)
Common name(s)
Occurrence
Scotch laburnum,
alpine goldenchain
common laburnum,
goldenchain tree
Waterer laburnum,
goldenchain tree
S Alps, N Apennines,
Tree
NW Yugoslovia, S
Slovakia & Czech Republic, Tree
& Central & S Europe
Observed (1856) wild
Tree/shrub
in Tyrol & Switzerland,
now in cultivation
Sources:
Growth
habit
Height at
Year first
maturity (m) cultivated
6.1
1596
6.19.1
1560
3.74.6
1875
Laburnum
631
632
References
Allen DH. 1994. Personal communication. Sandwich, MA: F.W. Schumacher
Co.
Dirr MA. 1990. Manual of woody landscape plants: their identification,
ornamental characteristics, culture, propagation, and uses. Champaign, IL:
Stipes Publishing Company. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Gillis S, Debergh P. 1992. Preliminary results on micropropagation of
Laburnum watereri Vossii. Mededlingen van de Faculteit
Landbouwwetenschappen Rijksuniversiteit Gent 57(4a): 15531555.
Greinwald R, Bachmann P, Witte L, Czygan FC. 1990. Cytisine-12-carboxyethylester, a quinolizidine alkaloid from Laburnum watereri and its occurrence in the leguminosae. Phytochemistry 29(1): 35533554.
Hartmann HT, Kester DE, Davies FT. 1990. Plant propagation: principles and
practices. Englewood Cliffs, NJ: Prentice Hall. 647 p.
Hillier Nurseries (Winchester) Ltd. 1991. The Hillier manual of trees and
shrubs. Melksham, Wiltshire: Redwood Press Ltd. 704 p.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing: rules 1993. Seed Science & Technology 21 (Suppl.): 1259.
Krssmann G. 1984. Manual of cultivated broad-leaved trees and shrubs.
Volume 2, EPro. Beaverton, OR: Timber Press. 445 p.
Laroppe E, Muller C, Boulet-Gercourt B. 1996. Levee de dormance des
graines de trios legumineuses arbores [Breaking dormancy of three leguminous tree species]: Robinia pseudoacacia, Laburnum anagyroides, and
Cytisus scoparius. Foret Entreprise 109: 4751.
Leyland A. 1981. Laburnum (Cytisus laburnum) poisoning in two dogs.
Veterinary Record 109(13): 287.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan. 1290 p.
Macdonald B. 1986. Practical woody plant propagation for nursery
growers. Portland, OR:Timber Press. 669 p.
NBV [Nederlandsche Boschbouw Vereeniging]. 1946. Boomzaden:
Handleiding inzake het oogsten, behandelen, bewaren en uitzaaien van
boomzaden [in Dutch]. Wageningen,The Netherlands: Ponsen and
Looijen. 171 p.
Rudolf PO. 1974. Laburnum anagyroides, laburnum. In: Schopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 476477.
Scheller VH. 1974. Die Bestimmungsmerkmale der kultivierten LaburnumArten. Mitteilungen der Deutschen Dendrologischen Gesellschaft 67:
4551.
Schubert J. 1955. Zr Prfung hartschaligen Saatgutes von Robinia pseudoacacia L. Archiv fr Forstwesen 4 (2/3): 241269.
Stilinovic S, Grbic M. 1988. Effect of various presowing treatments on the
germination of some woody ornamental seeds. Acta Horticulturae
226(1): 239245.
Wasson E. 2001. Trees and shrubs: illustrated AZ of over 8500 plants.
Willoughby, NSW, Australia: Global Book Publishing. 928 p.
Whalley DN, Loach K. 1981. Rooting of two genera of woody ornamentals
from dormant, leafless (hardwood) cuttings and their subsequent establishment in containers. Journal of Horticultural Science 56(2): 131138.
Whalley DN, Loach K. 1983. Establishment in containers of woody ornamentals propagated from dormant leafless cuttings. Combined
Proceedings of the International Plant Propagators Society 32: 186199.
Laburnum
633
LythraceaeLoosestrife family
Lagerstroemia L.
crape-myrtle
Jean-Jacques B. Dubois and Frank A. Blazich
Dr. Dubois is a plant physiologist at the USDA Agricultural Research Services Plant Science Research
Unit, Raleigh, North Carolina; Dr. Blazich is alumni distinguished graduate professor of plant propagation and tissue culture at North Carolina State Universitys Department of Horticultural Science, Raleigh, North Carolina.
Occurrence, growth habit, and uses. There are
about 55 species in the crape-myrtle genusLagerstroemia.
They are indigenous primarily to the Asian and Pacific
island tropics but also occur in China, India, Korea, Japan,
and Australia (Brner 1962; LHBH 1976). Many are important timber species, producing wood of quality suitable for
cabinetry and construction that is also highly resistant to
decay and destructive insects (Brner 1962; Howard 1948).
Three species are cultivated in North America, all for their
horticultural interest (table 1).
One species of crape-myrtle, Lagerstroemia indica L.,
and its hybrids with L. fauriei Koehneboth of which are
called crape-myrtleare used widely in landscape plantings
in warmer parts of the continental United States, particularly
the South. Lagerstroemia indica is indigenous to China,
whereas L. fauriei was introduced in 1956 by Creech (1958,
1985) from seeds collected on Yakushima Island, Japan.
Many cultivars have been named, including a few of strict
L. fauriei parentage (Dirr 1998; Egolf and Andrick 1978;
Raulston and Tripp 1995). Cultivars of crape-myrtles are
typically cold hardy to USDA Zone 7, but some cultivars
have withstood temperatures of 23 C without injury
(Egolf 1990b). There have been reports of tropical species,
particularly Queens crape-myrtle, growing in frost-free portions of the United States corresponding to USDA Zones 10
and 11 (Egolf and Andrick 1978; Everett 1981; Menninger
1962).
Common name(s)
Occurrence
L. fauriei Koehne
L. indica L.
L. elegans Wallich ex Paxt.
L. speciosa (L.) Pers.
L. flos-reginae Retz.
crape-myrtle, crapemyrtle
crape-myrtle, crapemyrtle
Queens crape-myrtle,
pride-of-India
634
open fruit
seeds.
longitudinal
Lagerstroemia
635
References
Babele GS, Kandya AK. 1986. Use of TTC for rapid testing of the viability
of Lagerstroemia parviflora (Roxb.) seeds. Journal of Tropical Forestry
2(3): 226231.
Brner J. 1962. Die Nutzhlzer der Welt.Volume 3. Weinheim, Germany:
Verlag von J. Cramer. 803 p.
Chudnoff M. 1980. Tropical timbers of the world. Madison, WI: USDA
Forest Service, Forest Products Laboratory. 831 p. [reprinted in 1984 as
Agric. Handbk. 607. Washington, DC: USDA Forest Service].
Creech JL. 1958. Exploring southern Japan for ornamental plants. National
Horticultural Magazine 37(2): 7594.
Creech JL. 1985. The National Arboretum does more than gather seeds.
American Nurseryman 161(2): 8182.
Dirr MA. 1998. Manual of woody landscape plants: their identification,
ornamental characteristics, culture, propagation and uses. Champaign, IL:
Stipes Publishing Company. 1187 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Egolf DR. 1981a. Muskogee and Natchez Lagerstroemia. HortScience
16(4): 576577.
Egolf DR. 1981b. Tuscarora Lagerstroemia. HortScience 16(6): 788789.
Egolf DR. 1986a. Tuskegee Lagerstroemia. HortScience 21(4): 10781080.
Egolf DR. 1986b. Acoma, Hopi, Pecos, and Zuni Lagerstroemia.
HortScience 21(5): 12501252.
Egolf DR. 1987a. Biloxi, Miami, and Wichita Lagerstroemia. HortScience
22(2): 336338.
Egolf DR. 1987b. Apalachee, Comanche, Lipan, Osage, Sioux, and Yuma
Lagerstroemia. HortScience 22(4): 674677.
Egolf DR. 1990a. Caddo and Tonto Lagerstroemia. HortScience 25(5):
585587.
Egolf DR. 1990b. Choctaw Lagerstroemia. HortScience 25(8): 992993.
636
PinaceaePine family
Larix P. Mill.
larch
Raymond C. Shearer
Dr. Shearer is a research forester at the USDA Forest Services Rocky Mountain Research Station
Forestry Sciences Laboratory, Missoula, Montana
as demonstrated at the Wind River Arboretum in southwestern Washington, where 7 larch species, some with several
varieties, and 1 hybrid were planted from 1913 to 1939
(Silen and Olson 1992). European larches there are doing
better than Asian species in this warm, moist Washington
state climate. The native western larch specimens from more
continental climates with lower humidity are doing poorly.
In 1992, a larch arboretum containing all species, several
varieties, and 3 hybrids of larch was established at Hungry
Horse, Montana, within the natural range of western larch
(Shearer and others 1995).
Growth habit. Larix is one of the few conifer genera
with deciduous needles. The trees are valued for their light
green hues in the spring and shades of yellow to gold in the
Common name(s)
Occurrence
L. decidua P. Mill.
L. europaea DC.
L. larix Karst
L. gmelinii (Rupr.) Rupr.
L. dahurica Turcz. ex Trautv.
L. cajanderi Mayr
L. kaempferi (Lamb.) Carr.
L. leptolepis (Sieb. & Zucc.) Gord.
L. japonica Carr.
L. laricina (Du Roi) K. Koch
European larch
Dahurian larch
Japanese larch
L. lyallii Parl.
subalpine larch,
alpine larch, tamarack
L. occidentalis Nutt.
L. sibirica Ledeb.
L. russica (Endl.) Sabine ex Trautv.
L. europaea var. sibirica (Ledeb.) Loud.
Larix
637
638
Conversely, low genetic variation occurs among populations of western larch for growth, phenology, and cold hardiness (Rehfeldt 1982) compared with other Rocky Mountain
conifers. Rehfeldt (1983) identified an 11% variation associated with the elevation of the seed source and recommended
that seedlots not be transferred more than 29 m or 2
contour bands. Based on genetic variation in allozymes of
western larch seeds, Fins and Seeb (1986) cautioned transferring seeds from eastern Washington to north Idaho and
recommended that seedlots for planting should include seeds
from a diversity of locations within an area. Hall (1985)
concluded that yields of cones and seeds from interspecific
and intraspecific crosses and open-pollinated seeds of
European larch were reduced in hybrid crosses compared to
non-hybrid crosses. Wide variation in yield suggests that
both genetic and environmental factors are important in controlling yield of seeds.
Hybrids. Larches hybridize readily (Rudolf 1974;
Lewandowski and others 1994; Young and Young 1992), and
geographic isolation is a major factor for lack of hybridization. Natural hybrids of western and subalpine larches occur
where their ranges overlap (Carlson and Blake 1969;
Carlson and others 1990). Seeds from natural hybrid trees
closely resemble those of western larch (Carlson and
Theroux 1993). Reciprocal cross pollinations between western and subalpine larches were successful, and germination
of seeds from these crosses was higher than that of seeds
from either parent (Carlson 1994).
L. kaempferi decidua, known as L. eurolepis A.
Henry and commonly called Dunkeld larch, originated about
1900. It has been planted extensively in northwestern
Europe and to a lesser extent in the eastern United States
and Canada because it combines desirable characteristics of
both parent species and grows faster than either (Eliason
1942; MacGillivray 1969). L. kaempferi sibirica, known
as L. marschlinsii Coaz, was originated in 1901. L. laricina decidua, known as L. pendula Salisb. or weeping larch,
was originated before 1800 (Rehder 1940). Many other larch
hybrids are known. Several larch species and hybrids were
tested as potential short-rotation fiber crops for the
Northeast and the Great Lakes region (Einspahr and others
1984) and in Wisconsin (Riemenschneider and Nienstaedt
1983); Dunkeld larch showed best growth in both studies.
Seeds from a single provenance of Japanese larch and 6
provenances of European larch had, after 5 years, 3 times
the growth potential of seeds from native red pine (Pinus
resinosa Ait.) in another Wisconsin study (Lee and Schabel
1989).
Larix
639
640
seed with
641
Atmospheric fluorides can reduce the size of seeds, percentage germination, numbers of seeds per cone, and numbers of cones per tree. Reproductive failure and mortality of
tamarack in Newfoundland have resulted in their replacement by more tolerant species (Sidhu and Staniforth 1986).
Micropropagation and genetic engineering.
Micropropagation techniques can supplement reliance on
larch seeds for a broad range of tree improvement and
regeneration needs. Karnosky (1992) suggests biotechnology
can help produce genetically superior larch by (1) mass
propagation, (2) disease screening, and (3) transfer of genetic information through genetic engineering techniques.
Organogenesis from young and mature larch callus tissues is
reported (Bonga 1984; Chapula 1989). Lelu and others
(1993) developed somatic embryogenesis techniques for
several species and hybrids of larch. Full-sib immature
zygotic embryos were produced from induction of embryonal masses for European and Dunkeld larches and Larix
leptoeuropaea (Lelu and others 1994a). Thompson and von
Aderkas (1992) successfully regenerated western larch from
immature embryos. Protoplasts of Dunkeld larch can be
effectively isolated from embryonal mass and cultured to
produce somatic plantlets (Charest and Klimaszewska
1994). Further, Lelu and others (1994b) showed that the
number of mature somatic embryos of Larix leptoeuropaea produced per gram (fresh weight) of embryonal mass
was influenced by embryogenic line, sucrose concentration,
and abscisic acid concentration. No universal maturation
medium was recommended because of the interactive effects
of these 3 factors. High plantlet survival was achieved in the
greenhouse through either of 2 acclimatization methods
(Lelu and others (1994c). In gymnosperms, gene transfer
was first accomplished in European larch; transfer was
mediated by Agrobacterium rhizogenes and subsequent
regeneration of the transgenic plants (Huang and others
1991). Shin and others (1994a&b) reported that transgenic
European larch plants were produced that use Agrobacterium-mediated single gene transfer to promote insect (Bt
toxin gene) and herbicide (aroA gene) resistance.
Collection of cones. Larch cones should be collected
as soon as they ripen; different species ripen at various times
from August to December (table 2). Larch cones are picked
from trees in forests, seed production areas, seed orchards,
and potted tree collections or they can be gathered from
felled trees, slash, or squirrel caches. In Tyrol, European
larch seeds were picked from the snow by hand; they can
also be gathered in late winter from canvas spread on the
ground before the trees were shaken to release the seeds
(Rudolf 1974). In most species, ripe cones are brown. Tests
show that seedcoats are hard and that female gametophytes
are firm. Often seeds mature earlier than expected and the
period for cone collection for tamarack (Smith 1981) and
western larch (Shearer 1977) can be expanded. Cones of
Siberian larch should be harvested when needles start to turn
yellow to assure high-quality seeds (Lobanov 1985). Data
on height, seed-bearing age, seed crop frequency, and
ripeness criteria are listed in tables 3 and 4.
Exraction of seeds. Freshly collected cones should be
spread out in thin layers to dry in the sun or in well-ventilated cone sheds. The cones can be opened by solar heat, by
heating in a cone kiln or room, or by tearing them apart
mechanically (Rudolf 1974; Tkachenko and others 1939).
Recommended kiln schedules are 8 hours at 49 C for
tamarack and 7 to 9 hours at 43 C for western larch
(Rudolf 1974).
After opening, cones should be run through a shaker to
remove the seeds. Sometimes equipment must be modified
to extract larch seeds (Saralidze and Saralidze 1976). Seeds
can then be de-winged by a de-winging machine, by treading in a grain sack, or by hand-rubbing. The integument,
which attaches the wing to the seed, is difficult to remove in
normal processing without damage (Edwards 1987). Finally,
Location
Flowering
Fruit ripening
Seed dispersal
L. decidua
L. gmelinii
MarMay
MayJune
Late Aprearly June
AprMay
Late Aprmid-May
MayJune
AprJune
AprMay
SeptDec
SeptNov
Sept
Earlylate Sept
Sept
Midlate Oct
AugSept
AugSept
SeptNov
Septspring
FebMar
Winter
Sept
SeptOct
SeptMar
L. kaempferi
L. lyallii
L. occidentalis
L. sibirica
Sources: Arno and Habeck (1972), Asakawa and others (1981),Chinese Academy of Sciences (1978), Kaigorodov (1907), Ohmasa (1956), Rudolf (1974), Shearer (1990),
Tkachenko and others (1939).
642
Species
L. decidua
L. gmelinii
L. kaempferi
L. laricina
L. lyallii
L. occidentalis
L. sibirica
Height at
maturity (m)
Year first
cultivated
940
2030
3040
920
925
3055
??40
1629
1827
1861
1737
1904
1881
1806
Minimum
seed-bearing
age (yrs)
10
1415
15
1216
40
30
25
12
L
Years
between large
seedcrops
310
24
3
48
36
210
210
35
Sources: Arno and Habeck (1972), Asakawa and others (1981), ODLF (1962, 1966), Schmidt and Shearer (1990),Tulstrup (1952).
Preripe color
Ripe color
L. decidua
L. gmelinii
Light brown
L. kaempferi
L. laricina
L. lyallii
L. occidentalis
L. sibirica
Greenpurple
Green-brown-purple
Length (mm)
1938
1925
1727
1624
1724
26
1932
1319
3851
2538
2538
Sources: Raevskikh (1979), Rehder (1940), Rudolf (1974), Shearer (1977), Suo (1982)
50% were sound; most of the unsound seeds had incompletely developed embryos and endosperm (Farmer and
Reinholt 1986). Hall and Brown (1977) found similar conditions among seeds of European and Japanese larches and
their hybrids. Seeds of western larch also have a high proportion of embryo failures (Owens and Molder 1979b). Use
of X-radiography was recommended to evaluate the quality
of tamarack seeds because flotation in 95% ethanol killed
52% of germinable seeds (Eavy and Houseweart 1987). A
purity of 80% and a viability (germinative capacity) of 20%
were recommended in 1966 as minimum standards for western larch (WFTSC 1966). Current standards for tree seeds to
be certified under OECD Certification in Ontario require a
minimum of 95% purity for tree seeds, resulting in an average germinability of 75 to 80% at 15 years for tamarack
(Wang 1995).
Storage of seeds. Because larch seeds can be stored
for long periods at seed moisture contents of 5 to 10% in
sub-freezing temperatures, Bonner (1990) classifies them as
true orthodox seeds. Gordon (1992) found that larch seeds
can be stored at 6 to 8% moisture content at 1 to 3 C for 25
years with little or no loss of germination quality. European
Larix
643
larch seeds keep well for a year or two if stored in the cones
(Rudolf 1974). Tamarack seeds store very well at 2 C for
10 years (Wang 1982). Details on seed storage for 6 species
are shown in table 7. There was no significant difference in
viability of European larch seeds stored at 0 C or in liquid
nitrogen (196 C) for 1 to 6 days (Ahuja 1986). European
larch seeds (Sudeten source) collected in 1956 and stored at
9% moisture content showed little decrease in germination,
if any at all, over a 12-year period (Hill 1976).
Pregermination treatments. Seeds of most larch
species germinate without pretreatment, but stratification in
Place collected
L. decidua
NE US Ontario, &
Europe
L. gmelinii
L. kaempferi
Japan
Japan & Europe
L. laricina
L. occidentalis
L. sibirica
kg/hl
lb/bu
kg/hl
31
2435
26.3
35.537
2435
32
32
24
1927
20
2829
1927
25
25
1.16
.96
0.962.57
0.961.28
0.96
0.71
64
*
lb/bu
0.90
0.75
0.752.00
0.751.00
0.75
0.55
0.50
Sources: Asakawa and others (1981), Eliason (1942), Eremko and others (1989), NBV (1946), Ohmasa (1956), ODLF (1966), Rudolf (1974),Tulstrup (1952).
* Here, 1.81 kg of seeds were extracted from 45.36 kg of cones (Gorshenin 1941).
Species
Place collected
L. decidua
Alps*
Tatra Mtns (Slovakia)
Sudeten Mtns
Romania
Europe & NW US
Sakhalin
Korea
Japan
NE US
Japan
Europe
Japan
Ontario
NW US
NW US
Europe
L. gmelinii
L. kaempferi
L. laricina
L. lyallii
L. occidentalis
L. sibirica
/kg
93214
161269
205265
150229
152225
93269
176465
359425
203331
241551
170302
150503
126335
117333
463926
494723
231359
216434
68163
154
198
229
187
179
159
265
390
236
249
265
254
190
701
556
313
302
97
/lb
Samples
70
90
104
85
81
72
120
177
107
113
120
115
86
318
252
142
137
44
141+
20
4
12
4
190+
21
5
12
14
68+
17+
16
10+
4
131+
71+
Sources: Asakawa and others (1981), Eliason (1942), Heit and Eliason (1940), NBV (1946), Ohmasa (1956), ODLF (1966), Rudolf (1974), Shearer (1961, 1977), Simak
(1967).
* Alpine race.
Sudeten race.
644
Seed moisture
content (%)
Temp (C)
Viable
period (yr)
9
7.5
6.2
7
5.59.8
12.1
48
69
6
68
6
910
24
24
2
24
24
-18
-18
4
13
24
12
14
15
10
1718
23
16*
25
13
Figure 3Larix laracina, tamarack: seedling development at 1 (right) and 8 (left) days after germination.
Sources: Heit (1967), Kiaer (1950), Rudolf (1974), Schubert (1954),Wang (1982),
Wang and others (1993).
* Viability of 5% retained after 16 years of storage.
645
646
0
0
0
030
21
60
0
0//
30
042
21
0
Moist
Moist
Moist
Moist
Moist
Sand
Moist
Moist
Soil
Moist
Moist
paper or blotters
paper
paper
paper
paper
paper or blotters
paper, sand
paper
paper or blotters
Medium
30
30
30
30
30
30
30
18
30
30
30
20
20
20
26
20
20
20
18
20
20
20
Days
Fall or spring
Fall
Spring
Spring
Spring
Species
L. decidua
L. laricina
L. leptolepis
L. occidentalis
L. sibirica
431538
269
753861
323592
323431
4050
25
7080
3035
3040
Seedlings
/m2
/ft2
36
6
36
36
36
0.13.25
0.25
0.13.25
0.13.25
0.13.25
Sowing depth
mm
in
Sowing
season
Type
Sources: AOSA (1993), Carlson and Blake (1969), Heit and Eliason (1940), ISTA (1993), Rudolf (1974), Shearer (1961), Simancik (1968).
* Daily light period was 8 to 16 hours.
Constant temperatures at 26 C and at 20 C also were used.
Cold stratification generally recommended for at least 21 days.
Constant temperatures at 24 C and at 20 C also were used.
// Seeds were soaked in USP 3% H2O2 for 24 hours in lieu of stratification.
L. sibirica
L. lyallii
L. occidentalis
L. laricina
L. gmelinii
L. kaempferi
L. decidua
Species
Cold
stratification
(days)
10
.38
Depth
mm
in
47
25
33
10
35
1020
40
30
Tree
percent
18
20
29
21
Germination rate
Amount
(%)
Days
36
52
43
47
14
15
57
Avg
(%)
Outplanting
age (yrs)
368
23
179
16
1
1
104
Samples
Germination
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ZygophyllaceaeCaltrop family
Larrea tridentata
(Sess & Moc. ex DC.) Coville
creosotebush
Richard T. Busing
Dr. Busing is an ecologist at the USDI Geological Survey, Corvallis, Oregon
Synonyms. Larrea tridentata (DC.) Cov.
Other common names. greasewood, gobernadora,
hediondilla.
Growth habit and occurrence. Creosotebush
Larrea tridentata (Sess & Moc. ex DC.) Covilleis an
evergreen shrub native to the arid subtropical regions of the
southwestern United States, Mexico, Argentina, and Chile
(Benson and Darrow 1945). Whether the North American
species L. tridentata is distinct from the South American
species L. divaricata Cav. has been unclear (Benson and
Darrow 1945), but most recent authors recognize L. tridentata as a separate species. It is the dominant shrub in all 3
warm deserts of the United States: the Mojave, Sonoran, and
Chihuahuan Deserts (Barbour and others 1980). Although
creosotebush can grow on a variety of substrates, it is most
abundant on calcareous soils (Musick 1978). Stands vary in
density and stature, depending on the aridity of the site
(Woodell and others 1969). Under very low rainfall, shrubs
are smaller and more widely spaced than those in stands
under more mesic conditions. Morphological and physiological adaptations of the genus Larrea to growth under xeric
conditions are well studied (Barbour and others 1974, 1977).
Despite dominance of the species in xeric sites, the emergence and growth of seedlings is favored by mesic conditions. Moisture, neutral pH, low salinity, and moderate temperatures are conducive to successful germination and
seedling establishment (Barbour and others 1977).
Use. Creosotebush is not browsed by livestock.
Although an edible livestock feed has been made from
creosotebush and a valuable antitoxidant has been extracted
from the shrub (Duisberg 1952), no economically feasible
program for gathering and using large amounts of creosotebush has been developed. Creosotebush, like other common
plants with peculiar odor or taste, has been used in traditional medicine to cure various ills (Benson and Darrow 1945).
In arid and semiarid parts of the Southwest, creosotebush is
used for landscaping and reclamation of disturbed lands
(Day and Ludeke 1980; Graves and others 1978; Williams
and others 1974).
single carpel.
Larrea
651
longitudinal
References
Ackerman TL. 1979. Germination and survival of perennial plant species in
the Mojave Desert. Southwestern Naturalist 24: 399408.
Barbour MG. 1968. Germination requirements of the desert shrub Larrea
divaricata. Ecology 49: 915923.
Barbour MG, Burk JH, Pitts, WD. 1980. Terrestrial plant ecology. Menlo Park,
CA: Benjamin/Cummings Publishing. 604 p.
Barbour MG, Cunningham GL, Oechel WC, Bamberg SA. 1977. Growth
and development, form and function. In: Mabry TJ, Hunziker JH, Difeo
DR, eds. Creosote bush. Stroudsburg, PA: Dowden, Hutchinson, and
Ross: 4891.
Barbour MG, Diaz DV, Breidenbach RW. 1974. Contributions to the biology
of Larrea species. Ecology 55: 11991215.
Benson L, Darrow RA. 1945. A manual of southwestern trees and shrubs.
Biol. Sci. Bull. 6. [Tucson]: University of Arizona. 411 p.
Boyd RS, Brum GD. 1983. Postdispersal reproductive biology of a Mojave
Desert population of Larrea tridentata (Zygophyllaceae). American
Midland Naturalist 110: 2536.
Day AD, Ludeke KL. 1980. Reclamation of copper mine wastes with shrubs
in the southwestern U. S. A. Journal of Arid Environments 3: 107112.
Duisberg PC. 1952. Development of a feed from creosotebush (Larrea
divaricata) determination of its nutritive value. Journal of Animal Science
11: 174180.
Graves WL, Kay BL, Williams WA. 1975. Seed treatment of Mojave Desert
shrubs. Agronomy Journal 67: 773777.
Graves WL, Kay BL, Williams WA. 1978. Revegetation of disturbed sites in
the Mojave Desert with native shrubs. California Agriculture 32: 45.
652
EricaceaeHeath family
Ledum L.
Labrador-tea
Tricia L.Wurtz
Dr. Wurtz is a research ecologist at the USDA Forest Services Pacific Northwest Research Station,
Boreal Ecology Cooperative Research Unit, Fairbanks, Alaska
ter and flower the following spring, in late May and early
June (Reader 1982). Flowers are white, with protruding stamens; they occur in numerous umbel-like clusters. Fruits
occur as drooping clusters of dry capsules (figure 1). A large
number of seeds are produced per flower. Sumner (1964)
found a range of 34 to 181 seeds per fruit in her study of
marsh Labrador-tea in interior Alaska. Extensive flowering
is common. Seeds are small (bog Labrador-tea, 1.8 to 3.0
mm by 0.2 to 0.3 mm; marsh Labrador-tea, 1.4 to 2.0 mm
by 0.2 to 0.3 mm) (Karlin and Bliss 1983). Seedcoats are
golden and translucent, with a loose, elongated testa that
aids wind dispersal (Densmore 1997). Calmes and Zasada
(1982) found that only 45% of bog Labrador-tea seeds were
filled.
Extraction, cleaning, and storage of seeds. Seed
capsules open as they dry, readily releasing seeds. Empty
capsules can be separated from seed with a fine-mesh sieve.
Most seed viability is lost within 1 year of collection. When
seeds were stored for 22 months at 4 C, germination
dropped from 58 to 16% (Karlin and Bliss 1983).
Pre-germination treatments. Labrador-tea does not
require cold stratification for germination, but most data
suggest that stratification improves germination. In a study
of marsh Labrador-tea, seeds exhibited shallow dormancy
(Calmes and Zasada 1982); 30 days of cold stratification
Common name
Occurrence
L. glandulosum Nutt.
L. palustre L. ssp. decumbens (Ait.) Hulten
L. palustris ssp. groenlandicum (Oeder) Hulten
western Labrador-tea
marsh Labrador-tea
L. groenlandicum Oeder.
L. decumbens (Ait.) Lodd ex Steud.
bog Labrador-tea
N Europe
SE & interior Alaska; Canada E to
Newfoundland, S to New Jersey,
Ohio, Minnesota, & Washington
Alaska & E through Canada to Greenland;
S to Labrador & Hudson Bay; also
N Europe & Asia
Ledum
653
References
654
Calmes MA, Zasada JC. 1982. Some reproductive traits of four shrub
species in the black spruce forest type of Alaska. Canadian FieldNaturalist 96: 3540.
Densmore RV. 1997. Effect of day length on germination of seeds collected
in Alaska. American Journal of Botany 84(2): 274278.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Hulten E. 1968. Flora of Alaska and neighboring territories. Stanford, CA:
Stanford University Press.
Junttila O. 1972. Effect of gibberellic acid on dark and light germination at
different temperatures of Calluna, Ledum and Rhododendron seeds.
Physiologia Plantarum 26: 239243.
Karlin EF, Bliss LC 1983. Germination ecology of Ledum groenlandicum and
Ledum palustre ssp. decumbens, in accordance with latitudinal change.
Arctic and Alpine Research 15(3): 397404.
MacKinnon A, Pojar J, Coupe R. 1992. Plants of northern British Columbia.
Edmonton, AB: British Columbia Ministry of Forests and Lone Pine
Publishing, 345 pp.
McGraw, JB Shaver, GR. 1982. Seedling density and seedling survival in
Alaskan cotton grass tussock tundra. Holarctic Ecology 5(2): 212217.
Reader RJ. 1982. Variation in the flowering date of transplanted ericacious
shrubs in relation to their flowering season. Journal of Biogeography. 9:
397410.
Reichardt PB, Bryant JP, Anderson BJ, Phillips D, Clausen TP, Meyer M, Frisby K.
1990. Germacrone defends Labrador tea from browsing by snowshoe hares. Journal of Chemical Ecology 16(6): 19611970.
Sheat WG. 1948. Propagation of trees, shrubs and conifers. London:
Macmillan and Co. 479 p.
Sumner PC. 1964. Ecology of Ledum groenlandicum Oeder in interior
Alaska [MS thesis]. Fairbanks: University of Alaska. 174 p.
Viereck LA, Little EL Jr. 1972. Alaska trees and shrubs. Agric. Handbk. 410.
Washington, DC: USDA Forest Service. 265 p.
Young JA,Young CG. 1992. Seeds of woody plants in North America.
Portland, OR: Dioscorides Press. 405 p.
Ziller WG. 1974. The tree rusts of western Canada. Pub. 1329.Victoria, BC:
Canadian Forestry Service. 272 p.
FabaceaePea family
Lespedeza Michx.
lespedeza
Richard T. Busing and Willis G.Vogel
Dr. Busing is an ecologist with the USDI Geological Survey, Corvallis, Oregon; Mr.Vogel retired
from the USDA Forest Services Northeastern Research Station
Common name(s)
Occurrence
L. bicolor Turcz.
shrub lespedeza,
bicolor lespedeza
leafy lespedeza, shrub lespedeza
Thunberg lespedeza
L. cyrtobotrya Miq.
L. thunbergii (DC.) Nakai
L. sieboldii Miq.
L. racemosa Dipp.
L. formosa Koehne
L. japonica Bailey
Desmodium penduliflorum Oudem.
Lespedeza
655
656
References
Allen ON, Allen EK. 1981. The Leguminosae. Madison: University of
Wisconsin. 812 p.
Byrd M,Young WC, Davison VE. 1963. Seed yields of shrub lespedezas in
Arkansas. Journal of Wildlife Management 27: 135136.
Clewell AF. 1966. Identification of the lespedezas in North America and a
selected bibliography on lespedeza. Bull. 7. FL:Tall Timbers Research
Station. 29 p.
Crider FJ. 1952. Natob: a new bush lespedeza for soil conservation. Circ.
900. Washington, DC: USDA Soil Conservation Service. 10 p.
Davison VE. 1954. Lespedezas for quail and good land use. Leafl. 373.
Washington, DC: USDA Soil Conservation Service. 8 p.
Gabrielson FC, Cross EA, Bradshaw DK, Carter OC. 1982. Seed size influence on germination and plant growth of Kobe lespedeza and other
species used for surface mine revegetation. Reclamation and
Revegetation Research 1: 271281.
Gleason HA, Cronquist A. 1991. Manual of vascular plants of northeastern
United States and adjacent Canada. 2nd ed. New York: New York
Botanical Garden. 910 p.
Graetz KE. 1951. Shrub lespedeza requires insect pollination. Soil
Conservation 16: 224226.
Hensen PR. 1957. The lespedezas: Part 1. Advances in Agronomy 9:
113122.
Isely D. 1990. Vascular flora of the southeastern United States. Chapel Hill:
University of North Carolina Press. 258 p.
Lespedeza
657
FabaceaePea family
658
seed.
longitudinal
659
References
Brewbaker JL, Plucknett DL, Gonzales V. 1972. Varietal trials of Leucaena
leucocephala (koa haole) in Hawaii. Res. Bull. 166. Honolulu: University
of Hawaii, College of Agriculture, Hawaii Agricultural Experiment
Station. 29 p.
Daguma B, Kang BT, Okali DUU. 1988. Factors affecting germination of
leucaena (Leucaena leucocephala (Lam.) de Wit.) seed. Seed Science and
Technology 16(2): 489500.
Francis JK. 1993. Leucaena leucocephala established by direst seeding in
prepared seed spots under difficult conditions. Nitrogen Fixing Tree
Reports 11: 9193.
Little EL Jr, Wadsworth FH. 1964. Common trees of Puerto Rico and the
Virgin Islands. Agric. Handbk. 249. Washington, DC: USDA Forest
Service. 548 p.
NAS [National Academy of Sciences]. 1984. Leucaena: promising forage
and tree crop for the tropics. 2nd ed. Washington, DC: National
Academy of Sciences. 100 p.
Neal MC. 1965. In gardens of Hawaii. Spec. Pub. 50. Honolulu: Bishop
Museum Press. 924 p.
660
Oaks AJ, Skov O. 1967. Yield trials of Leucaena in the US Virgin Islands.
Journal of Agriculture of the University of Puerto Rico 51: 176181.
Parrotta JA. 1992. Leucaena leucocephala (Lam.) de Wit: leucaena, tantan.
Res. Note SO-ITF-SM-52. New Orleans: USDA Forest Service,
Southern Forest Experiment Station. 8 p.
Sherman M,Tamashiro M. 1956. Biological control of Araecerus levipennis
Jordan (Coleoptera: Anthribidae). Proceedings of the Hawaii
Entomological Society 16: 138148.
Takahashi M, Ripperton JC. 1949. Koa haole (Leucaena glauca): its establishment, culture, and utilization as a forage crop. Res. Bull. 100.
Honolulu: University of Hawaii, College of Agriculture. Hawaii
Agricultural Experiment Station. 56 p.
Van den Beldt RJ, Brewbaker JL, eds. 1985. Leucaena wood production and
use. Waimanalo, HI: Nitrogen Fixing Tree Association. 50 p.
Whitesell CD. 1974. Leucaena leucocephala, leucaena. In: Schopmeyer CS,
tech. coord. Seeds of woody plants in the United States. Agric. Handbk.
450. Washington, DC: USDA Forest Service: 491493.
EricaceaeHeath family
Leucothoe
661
References
Bir RE. 1987. Native plant production using a multidisciplinary approach.
American Nurseryman 166(12): 7483.
Blazich FA, Warren SL, Acedo JR, Whitehead, RO. 1991. Seed germination
of Leucothoe fontanesiana as influenced by light and temperature. Journal
of Environmental Horticulture 9(2): 7275.
Bridwell FM. 1994. Landscape plants: their identification, culture, and use.
Albany, NY: Delmar Publishers. 560 p.
Dirr MA. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation and uses. 4th ed. Champaign,
IL: Stipes Publishing. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Fordham AJ. 1960. Propagation of woody plants by seed. Arnoldia 20(6):
3340.
662
Glenn CT, Blazich FA, Warren SL. 1998. Influence of storage temperatures
on long-term seed viability of selected ericaceous species. Journal of
Environmental Horticulture 16(3): 166172.
Halfacre RG, Shawcroft AR. 1975. Carolina landscape plants. 2nd ed.
Raleigh, NC: Sparks Press. 325 p.
Ingram J. 1961. Studies in the cultivated Ericaceae: 1. Leucothoe. Baileya 9:
5766.
Melvin NC III. 1981. Additional comments on Leucothoe (Ericaceae). Baileya
21(3): 127130.
Odenwald N,Turner J. 1987. Identification, selection, and use of southern
plants for landscape design. Baton Rouge, LA: Claitors Publishing
Division. 660 p.
Wyman D. 1953. Seeds of woody plants. Arnoldia 13(7/9): 4160.
OleaceaeOlive family
Ligustrum L.
privet
Daniel A. Mikowski and William I. Stein
Mr. Mikowski is a general biologist and Dr. Stein is a forest ecologist emeritus at the USDA Forest Services
Pacific Northwest Research Station, Corvallis Oregon
Common name(s)
Occurrence
L.ovalifolium Hassk.
California privet
L. japonicum Thunb.
Japanese privet
L. lucidum Ait. f.
L. sinense Lour.
glossy privet
Chinese privet,
trueno de seto
European privet,
common privet
L. vulgare L.
Table 2Ligustrum, privet: height, leaf habit, color, and size of mature fruit
Species
L. ovalifolium
L. japonicum
L. lucidum
L. sinense
L. vulgare
Height at
maturity (m)
5
212
310
410
5
Leaf habit
Fruit color
Deciduous or half-evergreen
Evergreen
Evergreen
Deciduous or half-evergreen
Deciduous or half-evergreen
Purple-black, black
Purple-black, blue
Purple-black, blue-black
Purple-black, blue-black
Lustrous black
Sources: Bean (1978), McMinn and Maino (1937), Ohwi (1965), Radford and others (1968), Rehder (1940),Vines (1983).
Ligustrum
663
664
Flowering
Fruit ripening
L. ovalifolium
L. japonicum
L. lucidum
L. sinense
L. vulgare
JuneJuly
JuneSept
JulySept
MarJuly
JuneJuly
SeptNov
SeptNov
SeptOct*
SeptNov*
SeptOct*
Ligustrum
665
References
AFA [American Forestry Association]. 1996. 199697 National register of
big trees. American Forests 102(1): 2051.
Bailey LH. 1947. The standard cyclopedia of horticulture.Volume 2. New
York: MacMillan: 12012421.
Bean WJ. 1978. Trees and shrubs hardy in the British Isles.Volume 2, 8th
ed., rev. London: John Murray Ltd. 784 p.
Burrows FJ, Kohen J. 1983. Germination of Ligustrum lucidum W.T. Ait. and L.
sinense Lour. at different temperatures. Australian Weeds 2(4): 130132.
Catanzaro CJ, Skroch WA, Henry PH. 1993. Rooting performance of hardwood stem cuttings from herbicide-treated nursery stock plants. Journal
of Environmental Horticulture 11(3): 128130.
Chadwick LC. 1935. Practices in propagation by seed. American
Nurseryman 62(12): 39.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press: 146147.
Heit CE. 1968. Propagation from seed: 15. Fall planting of shrub seeds for
successful seedling production. American Nurseryman 128(4): 810,
7080.
Howe TK, Woltz SS. 1981. Symptomology and relative susceptibility of
various ornamental plants to acute airborne sulfur dioxide exposure.
Proceedings of the Florida State Horticultural Society 94:121123.
ISTA [International Seed Testing Association]. 1996. International rules for
seed testing: 1996. Seed Science and Technology 24 (Suppl.): 1335.
Keever GJ, Cobb GS, Mills DR. 1989. Effects of floral buds, flowers, and fruit
on the propagation of four woody ornamentals from stem cuttings.
Applied Agricultural Research 4(4); 285287.
Lee WG, Grubb PJ, Wilson JB. 1991. Patterns of resource allocation in
fleshy fruits of nine European tall-shrub species. Oikos 61: 307315.
Little EL Jr. 1979. Checklist of United States trees, native and naturalized.
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Martin AC, Zim HS, Nelson AL. 1951. American wildlife and plants. New
York: McGraw-Hill. 500 p.
McMinn HE, Maino E. 1937. An illustrated manual of Pacific Coast trees.
Berkeley: University of California Press. 409 p.
Meikle RD. 1958. British trees and shrubs. London: Eyre and Spottiswoode.
132133.
Morita M, Iwamoto S, Higuchi H. 1979. Interrelated effects of thermo- and
photo-periodism on growth and development of ornamental woody
plants: 4.The effects of chilling in winter and photoperiod on growth and
development of several ornamental woody plants. Journal of the
Japanese Society for Horticultural Science 48(2): 205212.
666
LauraceaeLaurel family
Lindera Thunb.
spicebush
Victor Vankus, Kenneth A. Brinkman, and H. M. Phipps
Mr.Vankus is a botanist at the USDA Forest Services National Seed Laboratory, Dry Branch, Georgia; Drs.
Brinkman and Phipps retired from the USDA Forest Services North Central Forest Experiment Station
Common name(s)
Occurrence
Japanese spicebush
bog spicebush
Lindera
667
seed.
668
longitudi-
Species
L. benzoin
L. melissifolia
L. obtusiloba
Stratification (days)
Warm
Cold*
30
28
90
105
120
90
91
56
119
90
Percentage
germination
8590
100
84
88
Sources: Barton and Crocker (1948), Brinkman and Phipps (1974), Dirr and
Heuser (1987), Olney (1960), Schroeder (1935).
* 1 to 5 C. 25 C. 20 to 30 C.
References
Barton LV, Crocker W. 1948. Twenty years of seed research at Boyce
Thompson Institute for Plant Research. London: Faber and Faber. 148 p.
Boyle M. 1997. Personal communication. Willow Springs, MO: Hicks Seed
Co.
Bremness L. 1994. Herbs. New York: Dorling Kindersley. 304 p.
Brinkman KA, Phipps HM. 1974. Lindera (L.) Blume, spicebush. In:
Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service:
503504.
Cipollini ML, Wallace-Senft DA, Whigham DF. 1994. A model of patch
dynamics, seed dispersal, and sex ratio in the dioecious shrub Lindera
benzoin (Lauraceae). Journal of Ecology 82(3): 621633.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1632 p.
Grimm WC. 1957. The book of shrubs. Harrisburg, PA: Stackpole Books.
522 p.
Huxley A, Griffiths M, Levy M, eds. 1992. The new Royal Horticultural
Society dictionary of gardening.Volume 3. New York: Stockton Press: 89.
Laurie A, Chadwick LC. 1931. The modern nursery, a guide to plant
propagation, culture and handling. New York: Macmillan. 494 p.
Murphy K. 1997. Personal communication. Plains, MT: Lawyer Nursery.
Niesenbaum RA. 1992. Sex ratio, components of reproduction, and pollen
deposition in Lindera benzoin (Lauraceae). American Journal of Botany
79(5): 495500.
Olney HO. 1960. Growth and translocation during the after ripening of
seeds with resting embryos [PhD thesis]. Newark, DE: University of
Delaware.
Rehder A. 1940. Manual of cultivated trees and shrubs hardy in North
America. New York: Macmillan. 996 p.
Schroeder EM. 1935. Dormancy in seeds of Benzoin aestivale L.
Contributions of the Boyce Thompson Institute 7: 411419.
USDA Forest Service. 1948. Woody-plant seed manual. Misc. Pub. 654.
Washington, DC: USDA Forest Service. 416 p.
Van Dersal WR. 1938. Native woody plants of the United States: their erosion-control and wildlife values. Misc. Pub. 303. Washington, DC: USDA.
362 p.
Lindera
669
HamamelidaceaeWitch-hazel family
Liquidambar styraciflua L.
sweetgum
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services, Southern Research Station
Mississippi State, Mississippi
670
fruiting
seed.
longitudinal
Liquidambar
671
8
8
0
30
30
30
20
20
30
Days
28
25
30
Germination rate
%
Days
76
86
21
14
Germination
Avg (%) Samples
85
95
85
14
23
13
testing (table 1) (AOSA 1993). Light is not absolutely necessary for germination of stratified seeds (Bonner 1967), but
it is normally used in all testing. Tetrazolium staining
(Bonner and Gammage 1967), radiography (Belcher and
Vozzo 1979), and the excised embryo method (Bonner and
Gammage 1967; Flemion 1948) also provide reliable tests of
viability. Germination is epigeal (figure 4). For moisture
testing, duplicate samples of 4 to 6 g each should be dried
for 17 1 hour at 103 2 C (ISTA 1993), or electric
meters can be used for rapid measurements (Bonner 1981).
Nursery practice. Stratified seeds should be broadcast or drilled in the spring to achieve an initial seedling
density of 100 to 160/m2 (9 to 15/ft2) (Barham 1980).
Aluminum powder may be mixed with wet stratified seeds
at a rate of 15 ml/45 kg of seeds (4 tablespoons/100 lb) to
achieve easy flow in seeders (Bonner 1974). The seeds
should be sown on the surface and lightly into the soil with
a roller. A 6- to 12-mm (1/4- to 1/2-in) mulch of sawdust,
sand, or chopped pine straw should be applied (Bonner
1974; Coleman 1965; Vande Linde 1964), although some
nurseries have reported better results with wood fiber
mulches at rates of 1,400 to 2,900 kg/ha (1,250 to 2,600
lb/ac) (Barham 1980).
Ornamental cultivars of sweetgum are usually propagated vegetatively. Cuttings taken in early June will root, and
budding is common also (Dirr and Heuser 1987).
672
References
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
Barham RO. 1980. Nursery-grown sweetgum production improved by
hydromulching.Tree Planters Notes 31(2): 2830.
Belcher E,Vozzo J. 1979. Radiographic analysis of agricultural and forest tree
seeds. Association of Official Seed Analysts Handbk. No. 31. 27 p.
Bonner FT. 1967. Germination of sweetgum seed in response to light.
Journal of Forestry 65: 339.
Bonner FT. 1970. Artificial ripening of sweetgum seeds.Tree Planters Notes
21(3): 2325.
Bonner FT. 1972. Maturation of sweetgum and American sycamore seeds.
Forest Science 18: 223231.
Bonner FT. 1974. Liquidambar styraciflua L., sweetgum In: Schopmeyer CS,
tech. coord. Seeds of woody plants in the United States. Agric. Handbk.
450. Washington, DC: USDA Forest Service: 505507.
Bonner FT. 1981. Measurement and management of tree seed moisture.
Res. Paper SO-177. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 11 p.
Bonner FT. 1987. Collection and care of sweetgum seed. New Forests 3:
207214.
Bonner FT. 1994. Predicting seed longevity for four forest tree species with
orthodox seeds. Seed Science and Technology 22: 361370.
Bonner FT, Farmer RE. 1966. Germination of sweetgum in response to
temperature, moisture stress, and length of stratification. Forest Science
12: 4043.
Bonner FT, Gammage JL. 1967. Comparison of germination and viability
tests for southern hardwood seed.Tree Planters Notes 18(3): 2123.
Brown CK, Kirkman LK. 1990. Trees of Georgia and adjacent states.
Portland, OR:Timber Press. 292 p.
Coleman MC. 1965. Georgias experience in hardwood seedling production. In: Proceedings, Region 8 Forest Nurserymens Conferences; 1964
August 1920; Morganton, NC; and September 1617; Oklahoma City,
OK. Atlanta: USDA Forest Service, State and Private Forestry: 2532.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant
propagation. Athens, GA:Varsity Press. 239 p.
Ebel BH, Summerville KO. 1983. An evaluation of feeding by a seed bug,
Leptoglossus oppositus (Say), on seed yield of sweetgum, Liquidambar
styraciflua L. Journal of the Georgia Entomological Society 18: 316320.
Flemion F. 1948. Reliability of the excised embryo method as a rapid test
for determining the germinative capacity of dormant seeds.
Contributions of the Boyce Thompson Institute 15: 229241.
Liquidambar
673
MagnoliaceaeMagnolia family
Liriodendron tulipifera L.
tuliptree
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
674
longitudinal
Liriodendron
675
Place collected
Mississippi
Warren Co.
Oktibbeha Co.
North Carolina*
E Tennessee
New York
Cone wt/
cone vol
kg/hl lb/bu
39
32
30
25
Seed wt/
cone vol
kg/hl
lb/bu
8
924
719
/kg
Cleaned seeds/weight
Range
Average
/lb
/kg
/lb
9,45517,200
32,43075,080
15,17030,980
22,05052,920
4,2887,804
14,71034,050
6,88014,050
10,00024,000
12,680
41,200
23,440
30,870
5,750
18,700
10,630
14,000
Samples
3
676
References
Adams RE. 1968. Are alternating temperatures more beneficial than constant temperatures during stratification of yellow poplar seeds? Tree
Planters Notes 19(3): 1617.
Affeltranger CE. 1969. An evaluation of a soil fumigation treatment for controlling cylindrocladium root rot in a forest nursery. Forest Pest Cont.
Rep. 70-1-1. Atlanta: USDA Forest Service, Southeastern Area, State and
Private Forestry. 7 p.
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
Beck DE. 1990. Liriodendron tulipifera L., yellow-poplar. In: Burns RM,
Honkala BH, tech. coords. Silvics of North America.Volume 2,
Hardwoods. Agric. Handbk. 654. Washington, DC: USDA Forest Service:
406416.
Belcher E,Vozzo J. 1979. Radiographic analysis of agricultural and forest tree
seeds. AOSA Handbk. 31. Association of Official Seed Analysts. 27 p.
Bonner FT. 1976a. Effects of gibberellin on germination of forest tree seeds
with shallow dormancy. In: Hatano K, Asakawa S, Katsuta M, Sasaki S,
Yokoyama T, eds. Proceedings, 2nd International Symposium on
Physiology of Seed Germination; 1976 October 1830; Fuji, Japan.Tokyo:
Government Forest Experiment Station: 2132.
Bonner FT. 1976b. Maturation and collection of yellow-poplar seeds in the
Midsouth. Res. Paper SO-121. New Orleans: USDA Forest Service,
Southern Forest Experiment Station. 8 p.
Bonner FT, Russell TE. 1974. Liriodendron tulipifera L., yellow-poplar. In:
Schopmeyer CS, tech. coord. Seeds of woody plants of the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service:
508511.
Bonner FT, Switzer GL. 1971. Upgrading yellow-poplar seeds. Res. Note
SO-129. New Orleans: USDA Forest Service, Southern Forest
Experiment Station. 4 p.
Boyce SG, Hosner JF. 1963. Alternating storage temperatures increase the
germination of yellow-poplar seed. Journal of Forestry 61: 731733.
Boyce SG, Kaeiser M. 1961. Why yellow-poplar seeds have low viability.
Tech. Pap. 168. Columbus, OH: USDA Forest Service, Central States
Forest Experiment Station. 16 p.
Carvell KL, Korstian CF. 1955. Production and dissemination of yellowpoplar seed. Journal of Forestry 53: 169170.
Chadwick LC. 1936. Improved practices in propagation by seed. American
Nurseryman 62(12): 39.
Cech FC, Keys RN. 1987. Collection of yellow-poplar seed-heads by
shaking. Northern Journal of Applied Forestry 4(2): 7881.
Goebel NB, McGregor WHD. 1973. Seedfall of three bottomland hardwood species. For. Bull. 11. Clemson, SC: Clemson University,
Department of Forestry. 5 p.
Guard AT. 1943. The development of the seed of Liriodendron tulipifera L.
Indiana Academy of Science Proceedings 53: 7577.
Guard AT, Wean RE. 1941. Seed production in the tulip poplar. Journal of
Forestry 39: 10321033.
Griswold M. 1999. Washingtons gardens at Mount Vernon: landscape of the
inner man. Boston: Houghton Mifflin. 192 p.
Heit CE. 1942. Tulip poplar (Liriodendron tulipifera): propagation and seed
date. Notes For. Invest. Cornell: New York Conservation Department 44.
1 p.
Houston DB, Joehlin KA. 1989. Are pollination bags needed for controlled
pollination programs with yellow-poplar? Silvae Genetica 38: 137140.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing (rules 1993). Seed Science and Technology 21 (Suppl.):
1259.
Kaeiser M, Boyce SG. 1962. X-ray negatives for sorting yellow-poplar
samaras with filled and empty seeds. Journal of Forestry 60: 410411.
Kavanagh K, Carleton TJ. 1990. Seed production and dispersal patterns in
populations of Liriodendron tulipifera at the northern edge of its range in
southern Ontario, Canada. Canadian Journal of Forestry Research 20:
14611470.
Limstrom GA. 1959. Yellow-poplar seed quality varies by seed trees, stands,
and years. Sta. Res. Note 134. Columbus, OH: USDA Forest Service,
Central States Forest Experiment Station. 2 p.
Little EL Jr. 1978. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Little EL Jr, Delisle AL. 1962. Time periods in development: forest trees,
North American. In: Altman PL, Dittmer, eds. Biological handbook on
growth. Washington, DC: Federation of American Societies for
Experimental Biology:Table 104.
Paton RR. 1945. Storage of tulip tree seed. Journal of Forestry 43: 764765.
Sluder ER. 1964. Quality of yellow-poplar planting stock varies by mother
tree and seedbed density. Tree Planters Notes 65: 1619.
Steavenson HA. 1940. The hammer mill as an important nursery implement. Journal of Forestry 38: 356361.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA, Soil
Conservation Service. 198 p.
Taft KA Jr. 1962. The effect of controlled pollination and honeybees on
seed quality of yellow-poplar (Liriodendron tulipifera L.) as assessed by Xray photography.Tech. Rep. 13. Raleigh: North Carolina State University
School of Forestry. 21 p.
Whipple SD. 1968. Yellow-poplar regeneration after seed tree cutting and
site preparation. Bull. 384. Auburn, AL: Auburn University Agricultural
Experiment Station. 15 p.
Williams RD, Hanks SH. 1976. Hardwood nurserymans guide. Agric.
Handbk. 473. Washington, DC: USDA Forest Service. 78 p.
Williams RD, Mony CC. 1962. Yellow-poplar seedling yield increased by
seed stratification. Journal of Forestry 60: 878.
Liriodendron
677
FagaceaeBeech family
Occurrence and growth habit. This evergreen hardwood species, the sole representative of its genus in North
America, is considered a link between the chestnuts
(Castanea) and the oaks (Quercus) (McMinn 1939). Tanoak
(also known as tanbark-oak)Lithocarpus densiflorus
(Hook. & Arn.) Rehd.has flowers that resemble those of
the chestnuts, but acorns that resemble those of the oaks.
Tanoak is found from just north of the Umpqua River in
southwestern Oregon southward throughout the coastal
ranges to the eastern end of the Santa Ynez Mountains in
western Ventura County, California. Its range then extends
eastward to near Grants Pass, Oregon, and the lower slopes
of Mt. Shasta, and then intermittently southward along the
western slopes of the Sierra Nevada to Mariposa County,
California (Griffin and Critchfield 1972).
A striking characteristic of the tanoak species is that,
throughout its range, the tree form is found where moisture
is presentfrom the soil, from fog, or from high relative
humidity (McDonald and Tappeiner 1987). Another characteristic of the species is that shade is a requirement, but the
amount varies by reproductive mode. Seedlings from acorns
need shade to become established and grow. Sprouts from
root crowns, which are often found in burned or otherwise
severely disturbed areas, grow best in full sunlight, but only
until the crowns close. From then on, and whether from
acorns or sprouts, only toplight is needed. Indeed, full sunlight is then deleterious.
Because partial shade is necessary, tanoak is often found
in dense stands, usually in mixture with several conifer and
hardwood species. Pacific madrone (Arbutus menziesii
Pursh) and Douglas-fir (Pseudotsuga menziesii (Mirb.)
Franco) are its most common associates. The species is particularly abundant in a belt surrounding the redwood forest
in northern coastal California and in Yuba and Butte
Counties in the Sierra Nevada (Sudworth 1908). But even
though abundant, it is reported to never form pure stands
(Jepson and others 1911). However, extensive pure stands of
678 Woody Plant Seed Manual
acorns.
longitudinal
Lithocarpus
679
seedling devel-
References
EDA [Economic Development Administration]. 1968. The Hoopa Valley
Reservation hardwood study report. Washington, DC: USDC EDA:
154 p.
Griffin JR, Critchfield WG. 1972. The distribution of forest trees in
California. Res. Paper PSW-82. Berkeley, CA: USDA Forest Service,
Pacific Southwest Forest and Range Experiment Station. 118 p.
Hickman JC, ed. 1993. The Jepson manual, higher plants of California.
Berkeley: University of California Press. 1400 p.
Huber DW, McDonald PM. 1992. Californias hardwood resource: history
and reasons for lack of a sustained hardwood industry. Gen.Tech. Rep.
PSW-135. Albany, CA: USDA Forest Service, Pacific Southwest Research
Station. 14 p.
Jepson WL, Betts HS, Mell CD. 1911. California tanbark oak. Bull. 75.
Washington, DC: USDA Forest Service. 34 p.
McDonald PM. 1978. Silviculture-ecology of three native California hardwoods on high sites in north-central California [PhD dissertation].
Corvallis: Oregon State University, Department of Forest Science. 309 p.
McDonald PM. 1992. Estimating seed crops of conifer and hardwood
species. Canadian Journal of Forest Research 22: 832838.
McDonald PM, Litton RB Jr. 1987. Enhancing the roadside view: creating
pleasing views can be expensive. Journal of Forestry 85(11): 1923.
McDonald PM,Tappeiner JC II. 1987. Silviculture, ecology, and management
680
of tanoak in northern California. In: Plumb TR, Pillsbury NH, tech. coord.
Gen.Tech. Rep. PSW-100. Proceedings, Symposium on Multiple-Use
Management of Californias Hardwood Resources; 1986 November
1214; San Luis Obispo, CA. Berkeley, CA: USDA Forest Service, Pacific
Southwest Forest and Range Experiment Station: 6470.
McDonald PM, Fiddler GO, Smith WH. 1989. Mulches and manual release
fail to enhance Douglas-fir seedling survival and growth. In: Proceedings,
10th Annual Forest Vegetation Management Conference; 1988
November 13, Eureka, CA: 140153.
McMinn HE. 1939. An illustrated manual of California shrubs. San Francisco:
J.W. Stacey, Inc. 689 p.
Mirov NT, Kraebel CJ. 1937. Collecting and propagating the seeds of
California wild plants. Res. Note 18. Berkeley, CA: USDA Forest Service,
California Forest and Range Experiment Station. 27 p.
Peattie DC. 1953. A natural history of western trees. Boston: HoughtonMifflin. 751 p.
Roy DF. 1974. Lithocarpus densiflorus (Hook. & Arn.) Rehd., tanoak. In:
Lithocarpus
681
CaprifoliaceaeHoneysuckle family
Lonicera L.
honeysuckle
Jean-Jacques B. Dubois and Frank A. Blazich
Dr. Dubois is a plant physiologist at the USDA Agricultural Research Services Plant Science Research Unit,
Raleigh, North Carolina; Dr. Blazich is alumni distinguished graduate professor of plant propagation and tissue
culture at North Carolina State Universitys Department of Horticultural Science, Raleigh, North Carolina
Occurrence, growth habit, and uses. Honeysuckles
include about 180 species of deciduous or evergreen, bushy,
scandent, twining, or creeping shrubs, found throughout the
Northern Hemisphere, south to Mexico and North Africa,
Java, and the Philippines (Huxley 1992). Many species are
cultivated for their attractive, often-fragrant flowers and for
their ornamental fruits. Some species furnish food and cover
for wildlife, whereas others are valuable for erosion control
and shelterbelt planting (Brinkman 1974; Huxley 1992).
They are valued also for their extreme cold hardiness
(Herman and Davidson 1997). Many species introduced in
the United States have escaped cultivation and have become
naturalized within this century (table 1). Japanese, Amur,
and Tatarian honeysuckles are now considered invasive
weeds (Luken and Thieret 1997)
Scientific and common names. The nomenclature of
honeysuckles has been the object of many revisions over
time. Many species were once classified as varieties, and
vice versa, making synonyms common. The names currently
accepted for species native to, naturalized, or in cultivation
in the United States are listed in table 1.
Geographic races and hybrids. Erstad (1991)
demonstrated extensive genetic variation between black
twinberry plants from various northwestern American provenances. Thirty North American honeysuckles have been
assigned varietal status (Kartesz 1994). Among the 74 cultivated honeysuckles reported by the Liberty Hyde Bailey
Hortorium (1976), 10 species are of hybrid origin.
Flowering and fruiting. The small, perfect flowers
vary from white or yellow to pink, purple, or scarlet. They
are borne in axillary pairs or sessile, 6-flowered whorls in
terminal spikes or panicles. Time of flowering varies not
only among species but also by geographic locality within
species (table 2). The attractive fruits are berries, white, red,
orange, blue, or black at maturity (table 2). They occur often
in coalescent pairs that ripen in the spring, summer, or early
fall (figure 1). Depending on the species, each berry contains a few to many small seeds that measure about 4 mm in
diameter (figures 2 and 3) (Brinkman 1974; Huxley 1992).
682
fruit
L. flava Sims
L. flava Sims var. flavescens Gleason
L. flavida Cockerell ex Rehder
L. fragrantissima Lindl. & Paxton
Xylosteon fragrantissimum
(Lindl. & Paxton) Small
L. hirsuta Eat.
L. hirsuta Eat. var. interior Gleason
L. hirsuta Eat. var. schindleri B. Bovin
L. hispidula (Lindl.) Dougl.
ex Torr. & Gray
L. interrupta Benth.
L. involucrata Banks ex Spreng.
L. etrusca Santi
Nebraska N to Saskatchewan,
E to Pennsylvania & Quebec
California to British Columbia
California to Oregon & Arizona
California N to Alaska, E to
New Mexico & New Brunswick
hairy honeysuckle
California honeysuckle
chaparral honeysuckle
black twinberry, bearberry
honeysuckle, inkberry,
skunkberry, twinberry
honeysuckle
China
winter honeysuckle,
sweet-breath-of-spring
Scandent to 4 m
California to
British Columbia
2m
3.5 m
2m
2.5 m
1.5 m
1.5 m
5.5 m
4m
Scandent to 6 m
1.5 m
Oklahoma N to Illinois, E
to South Carolina & Ohio
limber honeysuckle,
mountain honeysuckle
Etruscan honeysuckle
L. dioica L.
California N to British
Columbia, E to Utah & Montana
California N to Washington,
E to Nebraska & Idaho
Arizona N to British Columbia, E to
Georgia, Quebec, & Missouri
Mediterranean region
NE Asia to Japan
yellow honeysuckle
L. conjugialis Kellogg
fly honeysuckle
New Jersey to Massachusetts
& Nova Scotia
No occurrence except
in cultivation
2m
California N to British
Columbia, E to Pennsylvania
& Newfoundland
Italian woodbine,
Italian honeysuckle
coralline honeysuckle,
honeysuckle
orange honeysuckle
L. caerulea L.
Tennessee N to Iowa, E to
Georgia & Nova Scotia
Europe to W Asia
5.5 m
2.5 to 3 m
Arizona honeysuckle
Belle honeysuckle,
whitebell honeysuckle
bearberry honeysuckle,
sweetberry honeysuckle
Scandent to 4 m
Height (m)
N Am occurrence
of introduced species
Common names(s)
Native occurrence
scientific and common names, native occurrence and North American occurrence of introduced species and height at maturity
Lonicera
683
684
SE China
blueleaf honeysuckle
Amur honeysuckle
Morrow honeysuckle
woodbine honeysuckle,
European honeysuckle
grape honeysuckle
Manchurian honeysuckle
trumpet honeysuckle,
coral honeysuckle
Standish honeysuckle
L. morrowii Gray
L. periclymenum L.
L. reticulata Raf.
L. prolifera (Kirchn.) Booth
ex Rehder
L. prolifera (Kirchn.) Booth
ex Rehder var. glabra Gleason
L. sullivantii Gray
L. ruprechtiana Regel
L. x muscaviensis Rehder
L. sempervirens L.
L. xylosteum L.
China
1.5 m
1m
3m
Missouri N to Ontario,
E to Virginia & Quebec
2.5 m
4m
3.5 m
4m
6m
4m
1.5 m
3m
5m
3m
Scandent to
10 m
2m
Height (m)
California N to Alberta, E to
Virginia & Nova Scotia
BONAP (1996), Brickell and Zuk (1997), Brinkman (1974), Cullina (2002), Dirr (1990), Dwelley (1980), FNPS (2002), Huxley (1992), Kartesz (1994), LHBH (1976).
Utah honeysuckle
L. utahensis S.Wats.
Sources:
southern honeysuckle
Tatarian honeysuckle
L. standishii Jacques
No occurrence except in
cultivation
Texas to South
Carolina & Ontario
New Mexico N to Saskatchewan,
E to Virginia & New Brunswick
E Asia
Japanese honeysuckle,
gold-and-silver-flower
California N to Oregon,
E to Texas & N to Nebraska,
E to Florida & Maine
Coastal California
Michigan N to Manitoba, E to
Nova Scotia
Europe, N Africa, & W Asia
Native occurrence
Common names(s)
N Am occurrence
of introduced species
Table 1Lonicera, honeysuckle: scientific and common names, native occurrence and North American occurrence of introduced species and height at maturity (continued)
Location
Arizona
NE US
Japan
L. ciliosa
L. conjugialis
L. dioica
L. etrusca
L. flava
L. fragrantissima
L. hirsuta
L. hispidula
L. interrupta
L. involucrata
E US
N Rocky Mtns
W US
L. involucrata var. California
ledebourii
L. japonica
L. korolkowii
L. maackii
NE US
Japan
L. morrowii
NE US
Japan
L. oblongifolia
L. periclymenum
L. reticulata
L. ruprechtiana
L. sempervirens
L. standishii
L. subspicata
L. tatarica
L. utahensis
W US
L. villosa
L. xylosteum
L.
L.
L.
L.
L.
L.
L.
albiflora
arizonica
x bella
caerulea
canadensis
caprifolium
chrysantha
Flowering
Fruit ripening
Spring
JuneJuly
Early summer
Spring
AprJuly
MayJune
June
Early summer
MayJuly
Summer
Late spring
Winterearly spring
MayAug
Summer
Early summer
June
JuneJuly
AprAug
MarJuly
JulyAug
JulySept
JulyAug
JulySept
SeptOct
Aug
JulyAug
Orange
Red
Red
Dark blue
Red, orange-red
Orange-red
Coral red, dark red
Red
Red
Red
Red
Red
Red
Yellow, red
Red
Red
Purple-black, glossy
Black
Summer
Late spring
June
May
MayJune
May
MayJune
Summer
Late spring
Summer
Early spring
MayJune
AprilJune
June
Late spring
SeptNov
AugSept
JuneAug
AugSept
JulyAug
JulyAug
JuneSept
JulyAug
Black
Bright red
Dark red, black
Sources: Bailey (1949), Brickell and Zuk (1997), Brinkman (1974), Dwelley (1980), FNPS (2002), , Hriteau (1990), Huxley (1992), Krssmann (1985), Las Pilitas Nursery
(2002), LHBH (1976), Maisenhelder (1958).
685
for 60 days at 20 to 30 C (table 4). He indicated that without such treatments, germination may be prolonged over a
period of 6 months or longer.
Germination tests. Germination is epigeal (figure 4),
and germination tests can be conducted in flats or in a germinator. Light is not necessary, at least for Tatarian honeysuckle. For most species, alternating temperatures of 30 and
20 C yield satisfactory results (table 5). Brinkman (1974)
reported conflicting results in 2 studies conducted to determine the optimal germination temperature of Tatarian honeysuckle. One study reported that 20 C or less was required
for complete germination, whereas the other reported that 18
to 20 C was the minimum needed, with the most rapid germination occurring at 25 to 27 C. Swingle (1939) reported
that tests by the USDA Soil Conservation Service (which is
now called the Natural Resources Conservation Service)
showed no correlation between seed viability, as estimated
by cutting tests, and germination rates as measured by germination tests. There are no official testing protocols for
honeysuckle species.
Nursery practice and seedling care. Seeds of species
of honeysuckle that only exhibit embryo dormancy can be
involucrata
maackii
morrowii
oblongifolia
tatarica
/kg
500,0001,050,000
260,000430,000
250,000440,000
520,000530,000
260,000440,000
686
Average
/lb
227,000477,000
116,000194,000
114,000191,000
234,000239,000
116,000198,000
/kg
720,000
330,000
335,000
520,000
310,000
/lb
326,500
148,000
152,000
236,000
142,000
Warm period
Species
L.
L.
L.
L.
hirsuta
maackii
oblongifolia
tatarica
Cold period
Temp (C)
Days
Temp (C)
Days
2030
2030
60
60
5
010
5
5
60
6090
90
3060
Species
L.
L.
L.
L.
L.
L.
L.
canadensis
chrysantha
dioica
hirsuta
involucrata
oblongifolia
tatarica
Germination conditions
Temp (C)
Medium
Day
Night
Sand
Sand
Sand
Sand
Sand
30
30
30
30
30
20
20
20
20
20
Days
90
80100
100
60
6090
Germination rate
Amount
(%)
Days
33
58
25
33
Total
germination
(%)
100
7891
95
43
83
37
85
Figure 4Lonicera tatarica, Tatarian honeysuckle: seedling sown either broadcast or by drills in the fall, or cold-stratidevelopment at 1, 3, 13, and 31 days after germination.
fied and sown in early spring. Seeds of species believed to
have an impermeable seedcoat as well, however, should be
sown as soon as possible after collection to ensure germination the next spring. Nondormant seeds may be sown in the
spring without pretreatment. Seeds should be covered with 3
to 6 mm (1/8 to 1/4 in) of nursery soil. Mulching with 5.0 to
7.5 cm (2 to 3 in) of straw prevents excessive drying of the
soil and seeds. Germination of Tatarian honeysuckle usually
is complete in 40 to 60 days after spring-sowing. This time
can be shortened by soaking seeds in water for 2 to 3 days
before sowing. About 15% of sown seeds of Tatarian honeysuckle result in usable seedlings. One- or 2-year-old
seedlings of this species and Amur honeysuckle are suitable
for field planting (Brinkman 1974).
Vegetative propagation of honeysuckles by stem cuttings
is also possible. Most species can be propagated readily by
softwood, semi-hardwood, or hardwood cuttings (Dirr and
Heuser 1987; Hartmann and others 2002).
Lonicera
687
References
Bailey LH. 1949. Manual of cultivated plants, rev. ed. New York: Macmillan.
1116 p.
Belcher CR, Hamer DW. 1982. Improved technique for harvesting Amur
honeysuckle seeds.Tree Planters Notes 33(3): 1719.
BONAP. 1996. The digital checklist of the vascular flora of North America
[website: http://www.bonap.org].
Brickell C, Zuk J, eds. 1997. The American Horticultural Society AZ
encylcopedia of garden plants. New York: DK Publishing. 1092 p.
Brinkman KA. 1974. Lonicera L., honeysuckle. In: Schopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk 450.
Washington, DC: USDA Forest Service: 515519.
Cram WH. 1982. Seed germination of elder (Sambucus pubens) and
honeysuckle (Lonicera tatarica). HortScience 17(4): 618619.
Cullina W. 2002. Native trees, shrubs, & vines: a guide to using, growing, and
propagating North American woody plants. Boston: Houghton Mifflin.
354 p.
Dirr MA. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation and uses. Champaign, IL:
Stipes Publishing Co. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Dwelley MJ. 1980. Trees and shrubs of New England. Camden, ME: Down
East Books. 275 p.
Erstad JLF. 1991. Annual shoot growth in different populations of Lonicera
involucrata collected in North America and grown in Norway. Euphytica
53(3): 165171.
FNPS [Flagstaff Native Plant and Seed]. 2002. Catalog of native plants and
seeds [website available at www.nativeplantandseed.com]. Flagstaff, AZ:
FNPS.
Hriteau J. 1990. The National Arboretum book of outstanding garden
plants. New York: Simon and Schuster. 292 p.
688
Hartmann HT, Kester DE, Davies FT Jr., Geneve RL. 2002. Hartmann and
Kesters plant propagation: principles and practices. 7th ed. Upper Saddle
River, NJ: Prentice Hall. 880 p.
Herman DE, Davidson CG. 1997. Evaluation of Lonicera taxa for honeysuckle aphid susceptibility, winter hardiness and use. Journal of
Environmental Horticulture 15(4): 177182.
Huxley A, ed. 1992. The new Royal Horticultural Society dictionary of gardening.Volume 3. London: Macmillan. 790 p.
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United States, Canada, and Greenland.Volume 1. Portland, OR:Timber
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Krssmann G. 1985. Manual of cultivated broad-leaved trees and shrubs. 3
vol. Portland, OR:Timber Press.
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www.laspilitas.com]. Escondido & Santa Margarita, CA: Las Pilitas
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LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. 3rd ed.
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Luken JO, Goessling B. 1995. Seedling distribution and potential persistence
of the exotic shrub Lonicera maackii in fragmented forests. American
Midland Naturalist 133(10): 124130.
Luken JO,Thieret JW, eds. 1997. Assessment and management of plant
invasions. New York: Springer-Verlag. 324 p.
Maisenhelder LC. 1958. Understory plants of bottomland forests. Occ. Pap.
165. New Orleans: USDA Forest Service, Southern Forest Experiment
Station. 40 p.
Munson RH. 1986. Extracting seeds from fleshy fruits. Plant Propagator
32(2): 1415.
Romanyuk VV. 1989. [Seed dormancy in species of the genus Lonicera
(Caprifoliaceae)]. Botanicheskii Zhurnal 74(9): 1328-1332 [in Russian;
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Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP-27. Washington DC: USDA Soil
Conservation Service. 198 p.
MyrtaceaeMyrtle family
seed.
longitudinal
are dry, simple agitation will separate the seeds from the
capsules. There are almost 5 million seeds/kg (2.2
million/lb), but as few as 2 or 3% of these may be viable
(Petteys 1974). The seeds are orthodox in storage behavior,
Lophostemon
689
690
References
Bailey LH. 1906. Encyclopedia of American horticulture.Volume 4. New
York: Macmillan. 2016 p.
Carlson NK, Bryan LW. 1959. Hawaiian timber for coming generations.
Honolulu:Trustees of the Bernice P. Bishop Estate. 112 p.
Francis WD. 1951. Australian rain-forest trees. Commonwealth of Australia
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Little RL Jr, Skolmen RG. 1989. Common forest trees of Hawaii (native and
introduced). Agric. Handbk. 679. Washington, DC: USDA Forest Service:
250254.
Maiden JH. 1904. The forest flora of New South Wales.Volume 1. Sidney,
Australia: W.A. Gullick Govt. Printer. 218 p.
Neal MC. 1965. In gardens of Hawaii. Spec. Pub. 50. Honolulu: Bernice P.
Bishop Museum Press. 924 p.
Petteys EQP. 1974. Tristania conferta R. Br., brushbox. In: Schopmeyer CS,
tech. coord. Seeds of woody plants in the United States. Agric. Handbk.
450. Washington, DC: USDA Forest Service: 817818.
Streets RJ. 1962. Exotic trees in the British Commonwealth. Oxford, UK:
Clarendon Press. 765 p.
FabaceaePea family
Lupinus L.
lupine
Don E. Riemenschneider,Tim D. Davis,Wayne A. Mackay, and Raymond D. Ratliff
Dr. Riemenschneider is a research geneticist at the USDA Forest Services North Central Research
Station, Rhinelander,Wisconsin; Dr. Davis is professor and director at Texas A&M Universitys Agricultural
Experiment Station, Dallas, and head of the Department of Horticultural Sciences, College Station,Texas; Dr.
Mackay is associate professor of environmental horticulture and associate resident director at Texas A&M
Universitys Agricultural Experiment Station; Dr. Ratliffe retired from the USDA Forest Services Pacific
Southwest Forest and Range Experiment Station
Common name(s)
Occurrence
Sources: Davidson and Barbour (1977), Everett (1957), Gadgil (1971a,b&c), Hickman (1993), USDA SCS (1982).
Lupinus
691
longitudinal
Storage
(yrs)
Wet chilling
(days)
2
0
0
0
72
0
0
0
Sources: Davidson and Barbour (1977), Mirov and Kraebel (1937), Ratliff (1974).
692
Test duration
(days)
6
10+
95
Germination
percentage
90
92
92
445
Tests
1
1
1
3
References
Christofolini G. 1989. A serological contribution to the systematics of the
genus Lupinus (Fabaceae). Plant Systematics and Evolution 166: 265178.
Davidson G, Barbour MG. 1977. Germination, establishment, and demography of coastal bush lupine (Lupinus arboreus) at Bodega Head, California.
Ecology 58: 592600.
Emery D. 1964. Seed propagation of California native plants. Leaflets of the
Santa Barbara Botanic Garden 1(10): 8196.
Everett PC. 1957. A summary of the culture of California plants at the
Rancho Santa Ana Botanic Garden 19271950. Claremont, CA: Rancho
Santa Ana Botanic Garden. 263 p.
Gadgil RL. 1971a. The nutritional role of Lupinus arboreus in coastal sand
dune forestry: 1.The potential influence of undamaged lupin plants on
nitrogen uptake by Pinus radiata. Plant and Soil 34: 357367.
Gadgil RL. 1971b. The nutritional role of Lupinus arboreus in coastal sand
dune forestry: 2.The potential influence of damaged lupin plants on
nitrogen uptake by Pinus radiata. Plant and Soil 34: 575593.
Gadgil RL. 1971c. The nutritional role of Lupinus arboreus in coastal sand
dune forestry: 3. Nitrogen distribution in the ecosystem before tree
planting. Plant and Soil 35: 113126.
Hickman JC, ed. 1993. The Jepson manual: higher plants of California.
Berkeley: University of California Press. 1400 p.
Lupinus
693
SolanaceaePotato family
Lycium L.
wolfberry
Paul O. Rudolph and Richard T. Busing
Dr. Rudolph (deceased) retired from the USDA Forest Services North Central Forest Experiment Station;
Dr. Busing is an ecologist at the USDI Geological Survey, Corvallis, Oregon
694
screens of suitable sizes (Glazebrook 1941). After extraction, the seeds should be dried and stored in sealed containers at 5 C (NBV 1946; Rudolf 1974), or stratified in moist
sand (Glazebrook 1941; NBV 1946). Stratified seeds of
matrimony vine will maintain good viability for 6 months
(NBV 1946), but there is no information on long-term storage of dry seeds. They appear to be orthodox, however, so
storage should not be a problem. Seed data are listed in
table 4.
Germination. Dormancy in wolfberry seeds is variable. Seed samples of Anderson wolfberry and Arizona
desert-thorn germinated well without pretreatment. They had
germination of 68 and 94% (Swingle 1939). Germination of
matrimony vine seeds, however, was hastened and improved
by stratification in moist sand for 60 to 120 days at 5 C.
After cold stratification, the average germination capacity
for 19 samples was 74% (Glazebrook 1941; NBV 1946;
Rudolf 1974). These tests were run in sand flats for 30 to 40
days at diurnally alternating temperatures of 30 to 20 C.
Germination after 18 days was 54%. Seeds of Rich wolfberry probably would benefit from similar pretreatment,
Figure 1Lycium barbarum, matrimony vine:
seed.
cleaned
Common name(s)
Occurrence
L. andersonii Gray
Anderson wolfberry,
Anderson desert thorn,
& water jacket, squawberry
matrimony vine,
boxthorn, European desert thorn
Chinese wolfberry,
Chinese matrimony-vine,
Chinese desertthorn
Arizona desert thorn
Rich wolfberry,
Baja desert thorn
L. barbarum L.
L. halimifolium P. Mill.
L. chinense P. Mill.
L. exsertum Gray
L. fremontii var. bigelovii Gray
L. richii Gray
L. palmeri Gray
L. pringlei Gray
Location
Flowering
Fruit ripening
L. andersonii
W US
SW US
California
Arizona
Holland, NE US
NE US
Japan
Arizona
California
AprJune
JanMay
NovApr
FebApr
JuneSept
JuneSept
AugNov
JanFeb*
MaySept
AugSept
AugOct
AugOct
JuneOct
L. barbarum
L. chinense
L. exsertum
L. richii
Sources: Bailey (1939), Kearney and Peebles (1942), McMinn (1951), Mirov and Kraebel (1939), NBV (1946), Ohwi (1965), Rehder (1940),Van Dersal (1938),Vines (1960),
Wyman (1947).
* Most abundant then, but flowers throughout the year (Kearney and Peebles 1942).
Table 3Lycium, wolfberry: height, length of cultivation, flower color, and fruit characteristics
Species
L. andersonii
L. barbarum
L. chinense
L. exsertum
L. richii
Height at
maturity (m)
0.33
16
12*
14
14
Year first
cultivated
Before 1935
Long cultivated
Before 1709
Before 1935
Before 1935
Flower color
Seeds/fruit
Red or white
Scarlet to orange-red
Very many
320
Scarlet to orange-red
Orange or red
Bright red
2030
3050
Sources: Bailey (1939), Benson and Darrow (1954), Hitchcock (1932), Kearney and Peebles (1942), McMinn (1951), Rehder (1940), Standley (1924),Vines (1960).
* Up to 4 m long as a prostrate rambler.
Lycium
695
longitudinal
Species
L. chinense
L. barbarum*
L. richii
Seed
soundness
(%)
99
98
Cleaned seeds/weight
Range
/kg
555,600586,400
Average
/lb
/kg
/lb
Samples
252,000266,000
377,000
573,000
3,022,600
171,000
260,000
1,371,000
1
3
1
References
Bailey LH. 1939. The standard cyclopedia of horticulture. New York:
Macmillan. 3639 p.
Benson L, Darrow RA. 1954. The trees and shrubs of the southwestern
deserts. Tucson: University of Arizona Press/Albuquerque: University of
New Mexico Press. 437 p.
Chiang F. 1983. Nomenclatural changes for new sectional delimitation in
Lycium (Solanaceae) of the New World.Taxon 32: 456458.
Glazebrook TB. 1941. Overcoming delayed germination in the seed of
plants valuable for erosion control and wildlife utilization [MS thesis].
Moscow, ID: University of Idaho, School of Forestry. 97 p.
Hitchcock CL. 1932. A monographic study of the genus Lycium of the
Western Hemisphere. Annals of the Missouri Botanical Garden 19:
179364.
Kearney TH, Peebles RH. 1942. Flowering plants and ferns of Arizona.
Misc. Pub. 423. Washington, DC: USDA. 1069 p.
Laurie A, Chadwick LC. 1934. Commercial flower forcing: the fundamentals
and their practical application to the culture of greenhouse crops.
Philadelphia: Blakistons. 519 p.
McMinn HE. 1951. An illustrated manual of California shrubs. Berkeley:
University of California Press. 663 p.
Mirov NT, Kraebel CJ. 1939. Collecting and handling seeds of wild plants.
For. Pub. 5. Washington, DC: USDA Civilian Conservation Corps. 42 p.
NBV [Nederlandsche Boschbouw Vereeniging]. 1946. Boomzaden: handleiding inzake het oogsten, behandelen, bewaren en uitzaaien van
boomzaden [Tree seed: handbook on the collection, extraction, storage,
and sowing of tree seed]. Wageningen,The Netherlands: Ponsen and
Looijen. 171 p.
696
MoraceaeMulberry family
227/hl
80/bu
No. of seeds/volume
70,000/hl
24,650/bu
2.9kg/hl
2.25lb/bu
30,900/kg
14,000/lb
15,40035,300/kg
7,00016,000/lb
of fruit
Seed weight/volume
of fruit
Cleaned seeds weight
Average
Range
Maclura
697
aggregate
exercised
698
References
Barnett JP, Burton JD. 1997. Osage-orange: a pioneering stewardship
species.Tree Planters Notes 48(3/4): 81-86.
Bonner FT, Ferguson ER. 1974. Maclura pomifera (Raf.) Schneid., Osageorange. In: Schopmeyer CS, tech. coord. Seeds of woody plants in the
United States. Agric. Handbk. 450. Washington, DC: USDA Forest
Service: 525526.
Burton JD. 1990. Maclura pomifera (Raf.) Schneid., Osage-orange. In: Burns
RM, Honkala BH, tech. coords. Silvics of North America.Volume 2,
Hardwoods. Agric. Handbk. 654. Washington, DC: USDA Forest Service:
426432.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant
propagation: from seed to tissue culture. Athens, GA:Varsity Press.
239 p.
Engstrom HE, Stoeckler JH. 1941. Nursery practice for trees and shrubs
suitable for planting on the Prairie-Plains. Misc. Pub. 434. Washington,
DC: USDA. 159 p.
Little EL. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Myatt A, Huffman G, Odell J. 1991. Seed processing manual. Stillwater:
Oklahoma Department of Agriculture, Forestry Division.
Sargent CS. 1965. Manual of the trees of North America (exclusive of
Mexico). 2nd ed., corrected and reprinted. New York: Dover. 934 p.
Williams RD, Hanks SH. 1976. Hardwood nurserymans guide. Agric.
Handbk. 473. Washington, DC: USDA Forest Service. 78 p.
Maclura
699
MagnoliaceaeMagnolia family
Magnolia L.
magnolia
Jill R. Barbour
Ms. Barbour is a germination specialist at the USDA Forest Services Tree Seed Laboratory,
Dry Branch, Georgia
The genus Magnolia comprises about 80 species of trees
naturally distributed throughout eastern North America and
southeastern Asia (Callaway 1994). It is the largest genus in
the family Magnoliaceae. There are 10 species and varieties
native to the United States (Callaway 1994) and 2 others
native to Puerto Rico (table 1) (Figlar 1982, 1984a).
Cucumber magnolia is the only species native to Canada.
There are no indigenous magnolias in Europe (Johnson
1973). Sweetbay was the first native American magnolia to
be cultivated in Europe in 1688 (Hora 1981).
Based on records of early fossil pollen and leaves, the
magnolias are considered the most ancient of all flowering
plants (FNAEC 1993). These plants are the base from which
all other angiosperms have evolved (FNAEC 1993).
Common name(s)
Occurrence
M. acuminata (L.) L.
Tulipastrum acuminatum (L.) Small
cucumber magnolia,
cucumbertree, yellow
cucumber magnolia
Ashe magnolia
M. ashei Weatherby
M. fraseri Walt.
M. auriculata Lam.
M. grandiflora L.
M. foetida (L.) Sarg.
M. macrophylla Michx.
M. splendens Urban
M. tripetala (L.) L.
Fraser magnolia,
mountain magnolia
southern magnolia,
evergreen magnolia, bull bay
bigleaf magnolia,
greatleaf(ed) magnolia,
large-leaf cucumbertree
Puerto Rico magnolia
pyramid magnolia,
ear-leaf(ed) magnolia,
ear-leaf umbrellatree
shining magnolia
umbrella magnolia
M. virginiana L.
M. australis Ashe
M. glauca L.
sweetbay, swamp-laurel,
sweetbay magnolia, southern
sweetbay, evergreen sweetbay
M. portoricensis Bello
M. pyramidata Bartr.
Sources: Callaway (1994), Figlar (1984a, b), Fordham (1960), LHBH (1978), Sargent (1965),Wasson (2001)..
* Sometimes spelled Ochlokonee.
700
Floral biology. The large, perfect flowers of the magnolias are borne singly at the ends of the branches in the
spring and summer. The flowers appear after the leaves
between April and June, except for cucumber magnolia,
which flowers before leaf bud-break. In section
Rhytidospermum, the flowers have 6 to 9 tepals (sepals and
petals), in section Magnolia, 8 to 12 tepals and in section
Tulipastrum, 9 to 12 tepals (table 2) (Fernald 1970). The
flowers have a pleasant fragrance, except those of umbrella
magnolia, which have an unpleasant odor (Burns and
Honkala 1990).
Magnolia flowers are highly specialized for cantharophilypollination by beetles, which predate the other pollinators, that is, bees, wasps, butterflies, and moths (Peigler
1988). Beetle-pollinated flowers are characterized by their
large size, white or pink color, lack of nectar, and abundance
of pollen (Peigler 1988). The flowering is protanderous to
prevent the flower from being pollinated with its own
pollen. Magnolia flowers close at night. The beetles (members of the Mordellidae and Nitidulidae families) chew
through the tepals with their strong mandibles to feed on the
flower parts (Peigler 1988). While feeding, the beetles get
covered with pollen. When the flower opens, the stigmas are
no longer receptive, and the stamens have dehisced and
detached from the gynandrophore (central axis of flower).
The beetles, covered with pollen, leave the flower to feed on
another flower, thus effecting cross-pollination (Thien
1974). The self-incompatible speciessuch as Fraser and
pyramid magnolias, sweetbay, and cucumber magnolia
cannot receive pollen from other flowers on the same tree
(McDaniel 1963). Nonviable seeds may have been collected
from trees that are self-incompatible. It is best to select other
trees for future collections.
Seed biology. The fruits develop from the gynandrophore into a follicetum (figure 1) (Callaway 1994). The
individual fruits are referred to as follicles and usually con-
Ploidy
Flower color
M. acuminata
M. ashei
M. fraseri
M. grandiflora
M. macrophylla
M. portoricensis
M. pyramidata
M. splendens
M. tripetala
M. virginiana
2N=76
2N=38
2N=38
2N=114
2N=38
2N=114
2N=38
2N=114
2N=38
2N=38
Yellow-green, yellow
White
Creamy white
Creamy white
White
Creamy white
Creamy white
Creamy white
White
Creamy white
Tepals
912
9
69
915
9
912
69
912
69
812
Magnolia
701
Figure 3Magnolia grandiflora, southern magnolia: longitudinal section through a seed (left) and seed with sarcotesta removed (right).
702
M
Cleaned seeds/weight
Range
Species
M. acuminata
M. fraseri
M. grandiflora
M. macrophylla
M. portoricensis
M. tripetala
M. virginiana
/kg
6,40014,500
5,47012,460
12,80015,000
Average
/lb
/kg
/lb
Samples
2,9006,600
2,4805,650
5,8006,800
12,020
10,030
14,220
9,550
7,410
16,540
16,600
5,450
4,550
6,450
4,330
3,360
7,500
7,530
15
12
8
1
1
1
5
Sources: Bonner (2002), Dirr and Heuser (1987), Francis and Rodriguez (1993), Heit (1968), Olson and others (1974).
Table 4Magnolia, magnolia: germination test conditions and results for stratified seeds
Species
Medium
M. acuminata
M. fraseri
M. grandiflora
M. macrophylla
M. portoricensis
M. virginiana
Sandperlite
Sandperlite
Sandvermiculite
Potting mix
Sandpeat
Kimpac
1520
20
20
22
30
18
20
3560
40100
3050
35
44
33
4761
Germinative capacity
Avg (%)
Samples
886
821
4390
71
64
93
3250
4
6
9
35
1
4
Sources: Afanasiev (1937), Del Tredici (1981), Francis and Rodriguez (1993), Hanchey and Kimbrough (1954), Jones, (1969), Olson and others (1974), Seitner (1981),
Toumey (1942).
Magnolia
703
References
Afanasiev M. 1937. A physiological study of dormancy in seed of Magnolia
acuminata. Cornell University Agricultural Experiment Station Memoir
208. 37 p.
AOSA [Association of Official Seed Analysts]. 2001. Rules for testing seeds.
AOSA. 125 p.
Bonner FT. 2002. Personal communication. Starkville, MS: USDA Forest
Service, Southern Research Station.
Bosch L. 1996. Personal communication. Jonesboro, LA: Bosch Nursery.
Bouman F. 1977. Integumentary studies in the Polycarpicae: 4. Liriodendron
tulipfera L. Acta Botanica Neerlandica 26(3): 213223.
Browse PM. 1986. Notes on the propagation of magnolias from seed.
Plantsman 8(1): 5863.
Buchanan M. 1996. Personal communication. Day, FL: Central Land and
Timber Co.
Burns RM, Honkala BH, tech. coords. 1990. Silvics of North America.
Volume 2, Hardwoods. Agric. Handbk. 654. Washington, DC: USDA
Forest Service. 877 p.
Callaway DJ. 1994. The world of magnolias. Portland, OR:Timber Press.
308 p.
Del Tredici P. 1981. Magnolia virginiana in Massachusetts. Arnoldia 41(2):
3649.
Dirr MA. 1990. Manual of woody landscape plants: their identification,
ornamental characteristics, culture, propagation, and uses. Champaign, IL:
Stipes Publishing. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant
propagation from seed to tissue culture. Athens, GA:Varsity Press. 239
p.
Durio B. 1996. Personal communication. Opelousas, LA: Louisiana Nursery.
Earle TT. 1938. Origins of the seedcoats in magnolia. American Journal of
Botany 25: 221222.
Evans CR. 1933. Germination behaviour of Magnolia grandiflora L. Botanical
Gazette 94: 729754.
704
Fernald ML. 1970. Grays manual of botany. 8th ed. New York:Van
Nostrand. 1632 p.
Figlar RB. 1982. Magnolia splendens: Puerto Ricos lustrous magnolia. Journal
of the Magnolia Society 18(1): 1316.
Figlar RB. 1984a. Magnolia portoricensis: Puerto Ricos other magnolia.
Journal of the Magnolia Society 19(2): 13.
Figlar RB. 1984b. Magnolia splendens. Journal of the Magnolia Society 20(1):
2324.
FNAEC [Flora of North America Editorial Committee]. 1993. Flora of
North America, North of Mexico.Volume 1, Introduction. New York:
Oxford University Press. 372 p.
Fordham AJ. 1960. Propagation of woody plants by seed. Arnoldia 20:
3340.
Francis JK, Rodriguez A. 1993. Seeds of Puerto Rican trees and shrubs: second installment. Res. Note SO-374. New Orleans: USDA Forest Service,
Southern Forest Experiment Station. 5 p.
Galle FC. 1953. The propagation of magnolias by seed. Proceedings of the
International Plant Propagators Society 3: 105107.
Hanchey RH, Kimbrough WD. 1954. Magnolia grandiflora seed germination
tests. In: Proceedings, Association of Southern Agricultural Workers; 1954
Feb. 13; Dallas,TX: 140.
Heit CE. 1939. Seed treatment and nursery practice with cucumber
(Magnolia acuminata). New York State Department of Conservation
Notes on Forest Investigations 20. 2 p.
Heit CE. 1955. The excised embryo method for testing germination quality
of dormant seed. Proceedings of the Association of Official Seed
Analysts 45: 108117.
Heit CE. 1968. Personal communication. Geneva, NY: New York State
Agricultural Experiment Station.
Hora B. 1981. The Oxford encyclopedia of trees of the world. Oxford, UK:
Oxford University Press. 288 p.
Jones L. 1969. Personal observation. Atlanta: USDA Forest Service,
Southeastern Area, State and Private Forestry.
Johnson H. 1973. The international book of trees. New York: Simon &
Schuster. 288 p.
Magnolia
705
BerberidaceaeBarberry family
Mahonia Nutt.
Oregon-grape
Don Minore, Paul O. Rudolf, and Franklin T. Bonner
Dr. Minore retired from USDA Forest Services Pacific Northwest Research Station, Corvallis, Oregon;
Dr. Rudolf (deceased) retired from the USDA Forest Services North Central Forest Experiment Station;
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Growth habit, occurrence, and use. Mahoniathe
Oregon-grapeis a genus of about 100 evergreen shrubs
native to Asia, Europe, North Africa, and the Americas
(Ahrendt 1961). Some authorities (Hitchcock and others
1964) place these species in the genus Berberis, and that
nomenclature was accepted in the 1974 edition of this book
(Rudolf 1974). However, most authorities (LBHB 1976;
USDA NRCS 1999) now separate the genera by placing the
evergreen species with compound leaves in Mahonia. The
distinction is far from clear, however: barberry is a common name for some Mahonia species (table 1) and intergeneric hybrids have been reported (Ahrendt 1961).
Several Oregon-grape species are valued as ornamentals
because of their foliage, flowers, or fruits (Bailey 1939; Dirr
Common name(s)
hollyleaf barberry,
Oregon grapeholly
Beale Oregongrape,
leatherleaf mahonia
Chinese mahonia
Height at
maturity (m)
0.63.0
Blue-black, bloomy
1.83.0
1.51.8
Purple-black
Fremont mahonia
0.94.6
Bluish black
red barberry
0.93.7
Blood red
Japanese mahonia
Cascades Oregongrape,
Cascades barberry
Nevin barberry
1.83.0
0.31.8
Blue
Deep blue, bloomy
0.91.8
cluster mahonia
2.43.0
Pruinose blue
Oregongrape,
creeping barberry
0.32.4
Purple, bloomy
Sources: Ahrendt (1961), Dirr (1990), Dirr and Heuser (1987), Hitchcock and others (1964), McMinn (1951), Rehder (1940), Rudolf (1974), USDA NRCS (1999),Vines
(1960).
706
Occurrence
M. aquifolium
M. fremonti
M. haematocarpa
M. nervosa
M. nevinii
M. repens
Oregon-grape contain alkaloids that are used in folk medicine in Asia (Zhao and others 1991), and seeds of hollyleaf
barberry contain tertiary alkaloids of note (Kostalova and
others 1986).
Flowering and fruiting. Perfect yellow flowers are
borne in the spring in racemes, panicles, umbels, fascicles,
or individually, depending on the species (Ahrendt 1961).
Stamens are contact-sensitive, and they respond to a tactile
stimulus by snapping toward the stigma (Millet 1976, 1977).
The fruit (figure 1) is a berry with one to several seeds (figures 2 and 3). A single sample of 100 fruits indicated that
most Cascade Oregon-grape berries have about 3 seeds
(Minore 1994). Good fruit crops are borne almost annually;
the fruits ripen in the summer and autumn (table 3). Seed
dispersal by both birds and mammals is widespread (Rudolf
1974; Vines 1960).
Collection of fruit; extraction and storage of seeds.
Ripe fruits may be picked by hand (with gloves) or flailed
onto cloths or receptacles spread beneath the bushes. The
fruits may be run through a macerator or blender with water
and the pulp then screened out or floated off. The seeds
should then be dried superficially and either sown immediately or stored in sealed containers at temperatures slightly
above freezing (Heit 1967; NBV 1946; Rudolf 1974). Seed
purity and soundness can be in the 90% range (Rafn and
Son nd; Rudolf 1974). Seeds of Fremont mahonia and
Oregon-grape did not loose viability for 5 years when stored
in unsealed containers in an unheated shed in a temperate
climate (Plummer and others 1968). Fruit yields, seed
yields, and numbers of seeds per weight are listed in table 4.
707
Figure 2Mahonia, Oregon-grape: seeds of M. aquifolium, hollyleaf barberry (top left); M. nervosa, Cascades mahonia
(top right); M. nevins, Nevin barberry (bottom left); and M. repens, Oregon-grape (bottom right).
Flowering
Fruit ripening
M. aquifolium
California
Black Hills, South Dakota (1,830 m)
M. fremontii
M. haematocarpa
M. nervosa
M. nevinii
M. repens
Sources: Bailey (1939), Loiseau (1945), McMinn (1951), Mirov and Kraebel (1939), NBV (1946), Ohwi (1965), Plummer and others (1965), Radford and others (1964),
Rudolf (1974),Van Dersal (1938),Vines (1960),Wappes (1982),Wyman (1947).
708
Mahonia
M. aquifolium
44
39
34
43*
8495
119157
/kg
90
0
90
196
Species
M. aquifolium
M. fremontii
M. nevinii
M. repens
Daily
light
(hrs)
Sand or perlite
Soil
Wet paper
Medium
30
21
20
21
30
95
10
62
12
150
Germination rate
Amount
(%)
Days
25
85
77
74
1
2+
1
1
95
90
90
Purity
(%)
33
41
42
103
23
30
57
62
99
90+
Soundness
(%)
1
2
1+
1
1
2
1
2
Samples
Sources: Heit (1968a&b), McLean (1967), Mirov and Kraebel (1939), Morinaga (1926), Plummer and others (1968), Rafn and Son (nd), Rudolph (1974), Swingle (1939),Vines (1960).
* Cold stratification temperatures ranged from 1 to 5 C.
Maximum germination was obtained with 4 months of warm stratification at 20 C, followed by 4 months of cold stratification at 2 to 4 C (Dirr and Heuser 1987).
Successive stratification periods were 30 days at 1 C, 60 days at 21 C, and 196 days at 1 C. During the final cold period, 62% of the seeds germinated. An additional 12% germinated after the temperature was again raised to 21
C for a total of 74%.
Cold
strati
fication*
(days)
73
90
93
227
51
66
126
136
Percent germination
Avg (%) Samples
3843
5471
Table 5Mahonia, Oregon-grape: stratification periods, germination test conditions, and percentage germination
M. nevinii
M. repens
M. fremontii
M. haematocarpa
M. nervosa
Place collected
Species
709
longitudinal
References
Ahrendt T. 1961. Berberis and Mahonia: a taxonomic revision. Journal of the
Linnean Society of London, Botany 57(369): 1410.
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
Bailey LH. 1939. The standard cyclopedia of horticulture. New York:
Macmillan. 3639 p.
Baskin CC, Baskin JM, Meyer SE. 1993. Seed dormancy in the Colorado
plateau shrub Mahonia fremontii (Berberidaceae) and its ecological and
evolutionary implications. Southwestern Naturalist 38(2): 9199.
Chadwick LC. 1936. Improved practices in propagation by seed. American
Nurseryman 62(8): [3]4; (9): 56; (10): 78; (12): [3]9.
Decker SR, Pekins PJ, Mautz WW. 1991. Nutritional evaluation of winter
foods of wild turkeys. Canadian Journal of Zoology 69(8): 21282132.
Dirr MA. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation, and uses. 4th. ed. Champaign,
IL: Stipes Publishing Co. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Heit CE. 1967. Propagation from seed: 8. Fall planting of fruit and hardwood seeds. American Nurseryman 126(4): 1213, 8590.
Heit CE. 1968a. Propagation from seed: 15. Fall planting of shrub seeds for
successful seedling production. American Nurseryman 128(4): 810,
7080.
Heit CE. 1968b. Thirty-five years testing of tree and shrub seeds. Journal of
Forestry 66(8): 632634.
Hitchcock CL, Cronquist A, Ownbey M,Thompson JW. 1964. Vascular
plants of the Pacific Northwest: 2. Salicaeae to Saxifragaceae. Seattle:
University of Washington Press. 597 p.
ISTA [International Seed Testing Association]. 1993. Rules for testing seeds:
rules 1993. Seed Science and Technology 21 (Suppl.): 1259.
Kern FD. 1921. Observations of the dissemination of the barberry. Ecology
2: 211214.
Kostalova D, Hrochova V,Tomko J. 1986. Tertiary alkaloids of Mahonia
aquifolium (Pursh) Nutt. III. Chemical Papers 40(3): 389394.
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Macmillan. 1290 p.
710
Schlosser WE, Blatner KH, Zamora B. 1992. Pacific Northwest forest lands
potential for floral greenery production. Northwest Science 66(1):
4455.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA Soil
Conservation Service. 198 p.
Terabayashi S. 1987. Seedling morphology of the Berberidaceae. Acta
Phytotaxonica Geobotanica 38(0): 6374 [Biological Abstracts 85(7):
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Baton Rouge, LA: USDA National Resources Conservation Service,
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Department of Agriculture. 362 p.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
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Forstwirte, Holzhndler und Holzindustriellen.Volume 1. Berlin: J.
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Wyman D. 1947. Seed collection dates of woody plants. Arnoldia 7(9):
5356.
Zhao TF, Wang XK, Rimando AM, Che CT. 1991. Folkloric medicinal plants:
Tinospora sagittata var. cravaniana and Mahonia bealei. Planta Medica 57:
5, 505.
Mahonia
711
RosaceaeRose family
Malus Mill.
apple
Paul D. Anderson and John A. Crossley
Dr. Anderson is a plant physiologist at the USDA Forest Services Pacific Northwest Research Station,
Corvallis, Oregon; Mr. Crossley retired from the USDA Forest Services
Northeastern Forest Experiment Station
712
and forbs. It also occurs in the coastal fringe forest as scattered, slow-growing individuals on rocky shorelines and
inland passages where it is protected from wind and salt
spray. Oregon crab apple is somewhat tolerant of brackish
water and short-term inundation.
In the upper mid-West of the United States, prairie and
Great Lakes crab apples occur in open areas, on welldrained soils, near forest margins, in abandoned pastures, in
oak savannahs, and at prairie margins (Kromm 1996; Little
1980; Yanny 1996). Common associates include shrubs of
hawthorn (Crataegus spp.) and bur (Quercus macrocarpa)
and black (Q. velutina) oaks. In southeastern Wisconsin, the
native crab apples occur with greater frequency on clay and
loam soils than on sandy soils. However, in contrast to
Oregon crab apple, neither prairie or Great Lakes crab
apples occur in wet areas (Kromm 1996).
In the southeastern United States, southern crab apple
occurs at low altitudes on moist soils of valley bottoms and
lower slopes. It is found in abandoned fields, along fence
rows, and at forest margins, often forming dense thickets
(Little 1980).
Native apples have served as a supply of food for both
wildlife and humans. Indigenous peoples in both the eastern
and western regions of North America have consumed crab
apples (Pojar 1996; Vierdeck and Little 1972; Yarnell 1964).
The occurrence of crab apples may be locally abundant in
areas traditionally inhabited by indigenous peoples, but it is
not known whether the trees were cultivated or grew from
discarded fruit remains (Pojar 1996).
Consumption of fruit by birds and mammals is common.
Known consumers include grouse (Bonasa umbellus),
pheasant (Phasianus colchicus), rabbits (Sylvilagus spp.),
squirrels (Sciurus spp.), opossum (Didelphis virginiana),
raccoon (Procyon lotor), skunks (Conepattus spp.) and
foxes (Vulpes vulpes) (Little 1980). The abundance of crab
apples along fencelines and riparian areas is thought to
reflect dispersal by wildlife. However, the large fruit size
and retention of the stem upon falling make transport by
Common name(s)
Occurrence
Japanese flowering
crab apple
Oregon crab apple, Pacific
crab apple, western crab
apple, wild crab apple
Asia
Sargent apple
European crab apple,
apple
Japan
Europe & W Asia
from the Siberian crab apple, but these varieties are usually
propagated vegetatively. Common apple and European
crabapple are most often used as the rootstock for cultivars
of crab apple (Fiala 1994).
Flowering and fruiting. The pink to white perfect
flowers appear in the spring with or before the leaves.
Flowering time varies among species from March to June
(table 2). The fruit is a fleshy pome in which 3 to 5 carpels,
usually 5, are embedded. Each carpel contains 2 seeds or 1
by abortion (figure 1) (Sargent 1965). Seeds have a thin lining of endosperm (figure 2), except in the cultivated, or
common, apple, which has almost no endosperm (Martin
1946). Depending upon species, fruits ripen as early as
August or as late as November (table 2). The fruits drop to
the ground soon after ripening. Color of ripe fruit varies
among the species (table 2).
Good crops of fruits and seeds generally occur every 2
to 4 years (Crossley 1974); however, good seed production
Malus
713
Table 2Malus, apple: phenology of flowering and fruiting, color of ripe fruit, and height of mature trees
Species
Flowering
Fruit ripening
M. baccata
M. coronaria
M. floribunda
M. fusca
May
MarMay
May
AugOct
SeptNov
OctNov
M. ioensis
M. pumila
M. x robusta
M. sargentii
MayJune
May
AprMay
SeptOct
AugOct
Color of
ripe fruit
Red or yellow
Yellow-green
Red or yellow
Green-yellow
to yellow & red
Greenish waxy
Yellow to red
Red or yellow-green
Red
Height of
mature trees (m)
12.2
9.2
3.610.0
812.2
15.4
1.82.5
Year first
cultivated
1784
1724
1862
1836
1885
Ancient times
1815
Sources: Callahan (1996), Crossley (1974), Fiala (1994), Krssmann (1960), Nielsen (1988), Rehder (1940), Sudworth (1908),Van Dersal (1938).
seeds.
longitu-
mal crop of crab apples were produced, yet the fruits bore
no seeds (Kromm 1996).
Biennial bearing is a problem for commercial apple production (Williams 1989). Alternate-year fruit production
arises from competitive effects of vegetative production,
fruit development, and flowering. Trees bearing fruit with a
large complement of seeds tend to initiate fewer flowers.
Chemical methods, including the post-bloom application of
thinning agents or growth regulators, have been used in
manipulating fruit set and fruit quality (shape, firmness,
russeting, and seed set) in commercial cultivars of apple
(Greene 1989; Looney and others 1992; Williams 1989).
Collection of fruits; extraction and storage of seed.
Ripe apples may be collected either by picking the fruits
in refrigerator in a 50:50 sand and peat mixture for an additional 3 months. As with seeds of commercial varieties of
apple, seeds from native crab apples may germinate in cold
storage, resulting in difficult sowing.
Apple seeds are orthodox in storage behavior; long-term
storage of seeds can be accomplished by drying the seedlot
to lower moisture contents. Seeds dried to a moisture content less than 11% have been stored in a sealed container at
2 to 10 C for over 2 years without loss of germinative
capacity or seedling vigor (Solovjeva and Kocjubinskaja
1955). Decline in seed viability as a function of storage temperature and seed moisture content has been modeled for
cultivars of cultivated apple (Dickie and Bowyer 1985).
They determined that seedlots dried to 14.5% moisture content (fresh-weight basis) and stored at 6 C lose half their
viability in 323 days. With further drying to 5%, the estimated storage life increases to 37 years at 5 C storage temperature or 100 years at 18 C storage temperature (Dickie
1986).
Germination. Apple seeds display dormancy which
has been overcome by cold stratification (table 4).
Stratification is achieved by placing the seeds in a moist
medium and storing at a temperature of 2 to 5 C. A minimum of about 30 days under stratification conditions is
required to remove embryonic dormancy (Zhang and
Lespinasse 1991). After stratification, apple seeds exposed
to diurnally alternating day/night temperatures of 30/20 C,
germinated in 30 to 60 days (table 4). Germination is
epigeal (figure 3).
The application of growth-regulating chemicals, including gibberellin A3 (GA3), ethephon (E), and benzylaminopurine (BAP), has been used to obtain germination from
non-stratified seeds (AOSA 1965; Litvinenko 1959; Sinska
1989; Zhang and Lespinasse 1991). Chemical treatments
often involve soaking excised embryos in growth regulator
solutions for periods of 1 to 24 hours. Variations in the concentration of growth substance and duration of soaking have
Species
M. bacatta
M. coronaria
M. fusca
M. floribunda
M. ioensis
M. pumila
M. x robusta
Seeds/fruit wt
/100 kg
/100 lb
1.2
0.5
1.2
0.4
2.5
1
2.4
0.75
/kg
10,85042,000
3,46013,300
7,00027,000
30,000
6,350
24,500
26,800
13,600
9,100
7,700
66,000
14,000
54,000
59,000
30,000
20,000
17,000
Samples
5
1
1
1
5+
Malus
715
significant impacts on both the percentage embryo germination and the quality of the resulting plants. Germination of
nearly 100% of non-stratified embryos has been obtained
with GA3 applied at concentrations of 12.5 to 50.0 mg/liter
for periods of 1 to 24 hours and with BAP applied at 12.5 to
100 mg/liter for periods of 6 to 24 hours. Such treatments
have resulted in 50 to 60% germination in less than 10 days
and nearly 100% germination in 30 days (Zhang and
Lespinasse 1991). Some plants produced from treated
embryos demonstrate reduced growth, abnormally thick
stems, or poorly developed roots. The percentage of abnormal plants produced tends to increase with increasing
growth regulator concentration or increasing period of soaking. Successful application of growth regulator treatments as
a substitute for stratification requires careful attention to
protocol and is beyond the needs of most propagators as
germination percentages of 90% or greater are commonly
achieved using the relatively simple cold stratification
process.
Nursery practice. Seedlings for use in landscape
restoration or as apple rootstocks are often grown from
seeds in nurseries (Richardson 1966; Callahan 1996).
Untreated seeds have been sown in late fall (Bakke and others 1926; Callahan 1996; Kromm 1996; Yanny 1996) and
stratified seeds have been sown in the spring (Crossley
1974; Yanny 1996). In a Washington nursery, seeds are stratified by first soaking them in water for 5 to 7 days, then
placing sacks of seeds between layers of ice in a sawdust pit
for 6 to 8 weeks. Seeds are subsequently dried only enough
to flow freely through a mechanical planter (Crossley 1974).
Seeds are sown in rows 20 cm (8 in) wide and 106 cm (42
in) apart (Davis 1940), 1.25 to 2.5 cm (1/2 to 1 in) deep on
loose friable soil. A thin sawdust mulch aids seedling emergence on soils that form a crust after watering. Seedlings
may be sprayed weekly with a fungicide to control powdery
mildew. By the end of the growing season most of the
Table 4Malus, apple: effects of cold stratification and germination test conditions on germination results
Species
M. bacatta
M. coronaria
M. fusca
M. floribunda
M. ioensis
M. pumila
M. x robusta
Cold
stratification
(days)
30
120
90
60120
60
60
60120
Germination conditions
Temp (C)
Day
Night
Days
30
10
30
30
20
10
20
20
30
30
10
60
Germinative energy
%
days
48
93
48
8
19
716
Germinative
capacity (%)
Samples
54
96
58
65
2
1
1
1+
soil temperatures are less than 4.5 C, and the seedlings are
generally hardy with respect to spring frost. Seedlings may
be grown for 2 years without undercutting, reaching a size
References
AOSA [Association of Official Seed Analysts]. 1965. Rules for testing seed.
Proceedings of the Association of Official Seed Analysts 54(2): 1112.
Bakke AL, Riches NW, Reeves K. 1926. Germination and storage of apple
seeds. Iowa Agricultural Experiment Station Research Bulletin 97:
243255.
Brown JE, Maddox JB, Splittstoesser WE. 1983. Performance of trees,
shrubs, and forbs seeded directly in the fall on minespoil and silt loam
soil. Journal of Environmental Quality 12: 523525.
Callahan F. 1996. Personal communication. Central Point, OR: Callahan
Seed.
Crossley JA. 1974. Malus Mill., apple. In: Schopmeyer CS, tech. coord. Seeds
of woody plants in the United States. Agric. Handbk. 450. Washington,
DC: USDA Forest Service: 531533.
Davis LL. 1940. Germination and growth of crab apple seedlings as
influenced by fungicidal treatment. Proceedings of the American
Horticultural Society 37: 359360.
Dickie JB. 1986. A note on the long-term storage of apple seeds. Plant
Genetic Resources Newsletter 65: 1315.
Dickie JB, Bowyer JT. 1985. Estimation of provisional seed viability constants
for apple (Malus domestica Borkh. cv. Greensleeves). Annals of Botany
56: 271275.
Fiala JL. 1994. Flowering crabapples: the genus Malus. Portland, OR:Timber
Press.
Green T. 1996. Personal communication. Macomb, IL: Western Illinois
University.
Greene DW. 1989. CPPU influences McIntosh apple crop load and fruit
characteristics. HortScience 24: 9496.
Heit CE. 1967. Propagation from seed: 11. Storage of deciduous tree and
shrub seeds. American Nurseryman 126(10): 1213, 8694.
Hickman JC. 1993. The Jepsen manual: higher plants of California. Berkeley:
University of California Press.
Kallio TK. 1962. Seed stratification with special reference to apples. Puntarha
65: 6263 [Horticultural Abstracts 32(4277), 1962].
Kromm D. 1996. Personal communication. Reeseville, WI: Reeseville Ridge
Nursery.
Krssmann G. 1960. Handbuch der Laubgeholze.Volume 1, 495 p.; volume
2, 608 p.
Little EL Jr. 1980. The Audubon Society field guide to North American
trees: eastern region. New York: Knopf. 714 p.
Litvinenko SN. 1959. [in Russian:The Ukrainian gibberellin, an effective
growth stimulant]. Doklady Akademii Nauk SSSR [Horticultural
Abstracts 30(73), 1960].
Looney NE, Granger RL, Chu CL, Mander LN, Pharis RP. 1992. Influences of
giberellins A4, A4+7 and A4+iso-A7 on apple fruit quality and tree productivity: 2. Other effects on fruit quality and importance of fruit position within the tree canopy. Journal of Horticultural Science 67:
841847.
Malus
717
MeliaceaeMahogany family
Melia azedarach L.
chinaberry
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station
Mississippi State, Mississippi
fruit and
longitudinal
References
Al-Sharook Z, Balan K, Jiang Y, Rembold H. 1991. Insect growth inhibitors
from two tropical Meliaceae: effects of crude seed extracts on mosquito
larvae. Journal of Applied Entomology 111: 425430.
Amata-Archachai P, Wasuwanich P. 1986. Mechanical extraction and
cleaning of nuts of some tropical species. Embryon 2(1): 18.
Atwal AS, Pajni HR. 1964. Preliminary studies on the insecticidal properties
of drupes of Melia azedarach against caterpillars of Pieris brassicae L.
(Lepidoptera: Pieridae). Indian Journal of Entomology 26: 221227.
Bonner FT, Grano CX. 1974. Melia azedarach L., chinaberry. In: Schopmeyer
CS, tech. coord. Seeds of woody plants in the United States. Agric.
Handbk. 450. Washington, DC: USDA Forest Service: 535536.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Guha SRD, Negi JS. 1965. Writing and printing paper from Melia azedarach
Linn. (Persian lilac). Indian Forester 91: 867869.
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Nair NC. 1959. Studies of Meliaceae: 2. Floral morphology and embryology
of Melia azedarach Linn.a reinvestigation. Journal of the Indian
Botanical Society 38: 367378.
Moncur MW, Gunn BV. 1990. Seed development and germination responses of Melia azedarach var. australasica. In:Turnbull JW, ed.Tropical tree
seed research. 1989 August 2124; Gympie, Australia. AICAR Proc. 28.
Canberra: Australian Centre for International Agricultural Research:
2428.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
Melia
719
MenispermaceaeMoonseed family
Menispermum canadense L.
common moonseed
Kenneth A. Brinkman and H. M. Phipps*
Drs. Brinkman and Phipps retired from the USDA Forest Services North Central Research Station
Growth habit, occurrence, and use. Common moonseedMenispermum canadense L.is a climbing woody
vine growing to a height of 3.6 m (Rehder 1940) that is
capable of spreading from underground stems (Wyman
1949). It is native from Quebec and western New England to
southeastern Manitoba, south to Georgia, Alabama, and
Oklahoma (Fernald 1950). The plants are seldom eaten by
livestock (Van Dersal 1938), but the fruits are of value to
wildlife, although reportedly poisonous to humans
(Kingsbury 1964). This species has been cultivated since
1646 for its attractive foliage and fruit (Rehder 1940).
Flowering and fruiting. The dioecious flowers
appear from May to July and the bluish-black drupes ripen
from September to November (Grimm 1966; Redher 1940).
The seeds are flattened stones in the form of a crescent or
ring (figures 1 and 2).
Collection of fruits and seed extraction. Fruits may
be collected from September to November (Rehder 1940).
Seeds may be extracted by washing the macerated fruits in
water. One sample showed 16,758 seeds/kg (7,600/lb)
(Brinkman and Phipps 1974).
Germination. Stratification of one seedlot at 5 C for
60 days resulted in 65% germination in 11 days and 98% in
26 days. An unstratified seedlot showed germination of 83%
in 28 days and 92% in 60 days (Brinkman and Phipps
1974). Germination was tested in sand under light at alternating temperatures of 30 (day) and 20 C (night).
Figure 1Menispermum canadense L., common moonseed:
fruit (top) and seeds (bottom).
720
OleaceaeOlive family
longitudi-
seed.
Menodora
721
but declined at warmer temperatures. Seeds of showy menodora germinated in the dark, but both germination rate and
germination percentage were greater in a light regime with
12 hours light (Fulbright and Flenniken 1986).
References
Fulbright TE, Flenniken KS. 1986. Effects of temperature and presowing
treatments on showy menodora seed germination. Journal of Range
Management 39(4): 310313.
Krugman SL. 1974. Menodora scabra A. Gray, rough menodora. In:
Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States: Agric. Handbk. 450. Washington, DC: USDA Forest Service: 539.
Munz PA, Keck DD. 1965. A California flora. Berkeley: University of
California Press. 1681 p.
Sabo DG, Johnson GV, Martin WC, Aldon EF. 1979. Germination requirements of 19 species of arid land plants. Res. Pap. RM-210. Fort Collins,
CO: USDA Forest Service. 23 p.
Turner BL. 1991. An overview of the North American species of Menodora.
Phytologia 71(5): 340356.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
TaxodiaceaeRedwood family
Growth habit, occurrence, and use. Dawn-redwoodMetasequoia glyptostroboides Hu & W.C. Cheng
is the only known living example of its genus (Hu and
Cheng 1948). It is often called a living fossil because until
1946 it was known only from the fossil record (Hu 1948;
Merrill 1945; Shao 1982). The natural range is quite restricted: a few trees are found near the village of Mo-tao-chi in
eastern Szechuan Province and the bulk of the native groves
are found in Shui-hsa-pa Valley (south of Mo-tao-chi) in the
northwestern corner of Hupeh Province, Peoples Republic
of China (Chu and Cooper 1950; Shao 1982). It has been
introduced to many other parts of China, as well as the
United States and Europe, for a total of 50 other countries
(Shao 1982).
Since its introduction into the United States in 1948,
this deciduous conifer has mostly been planted as an ornamental, especially at museums and in arboreta. The wood is
soft, weak, and brittle, so it has little value as a source of
lumber (Wyman 1968), although it is used for building timbers in China (Shao 1982). Pulping characteristics are similar to, and its fibers are stronger than, southern pines
(Wyman 1968). In the United States, Wyman (1968) reported height growth was as much as 18 m in 20 years. In
China, Shao (1982) described 4-year-old dawn-redwood
trees averaging 7 m tall and 11 cm dbh. In its natural range,
dawn-redwood grows in the submontane zone at elevations
between 100 and 1500 m. The species is hardy in Massachusetts, where the winter temperatures may drop to
-34 C, and thrives in Placerville, California, where summer
temperatures often exceed 35 C and there is usually no
summer rainfall (Johnson 1974).
Geographic races. Although great phenotypic diversity exists between planted trees, no geographic races are
known to exist. Several cultivars have been described
(Broekhuizen and Zwart 1967; DeVos 1963). Of the 6 trees
growing at the USDA Forest Services Institute of Forest
Genetics at Placerville, California, half are of the normal
single-stemmed conifer shape and the others are bushshaped with no single branch showing dominance. Johnson
723
724
References
Broekhuizen JT, Zwart FN. 1967. A contribution to the knowledge of
Metasequoia glyptostroboides [English summary]. Agric. Comm. 10.
Wageningen,The Netherlands: University of Wageningen Institute of
Forest Resources: 439463.
CDF [China Department of Forestry, Nanking College of Forest Products].
1977. A preliminary observation on the flowering and seed development of Metasequoia glyptostroboides Hu et Cheng. Acta Botanica Sinica
19(4): 252256.
Chu K, Cooper WS. 1950. An ecological reconnaissance in the native home
of Metasequoia glyptostroboides. Ecology 31(2): 260275.
DeVos F. 1963. Metasequoia glyptostroboides National. American
Horticulture Magazine 42(3): 174177.
Em H. 1972. Metasequoia glyptostroboides and its growth in the Skopje
basin. Sumarstvo 24:515.
Hu H. 1948. How Metasequoia, the living fossil, was discovered. Journal of
the New York Botanical Garden 49(585): 201207.
Hu H, Cheng W. 1948. On the new family Metasequoiaceae and on
Metasequoia glyptostroboides, a living species of the genus Metasequoia
found in Szechuan and Hupeh. Bulletin of Fan Memorial Institute of
Biology NS1(2): 153161.
Hwa CT. 1945. UCB. M 186893. [Herbarium Specimen] Metasequoia
glyptostroboides. Shui-hsa-pa Valley, 3550 ft. March 10, 1945.
Hwa CT. 1948. UCB. M 186884. [Herbarium Specimen] Metasequoia glyptostroboides. In ravine; tree. Li-Chuan, Shui-hsa-pa Valley, 3500 ft. Hupeh
Province. March 10, 1948.
Metasequoia
725
RubiaceaeMadder family
Mitchella repens L.
partridgeberry
Kenneth A. Brinkman, G. G. Erdmann, and Jill R. Barbour
Drs. Brinkman and Erdmann retired from the USDA Forest Services North Central Forest Experiment
Station; Ms. Barbour is a germination specialist at the USDA Forest Services National Seed Laboratory,
Dry Branch, Georgia.
Growth habit, occurrence, and use. Partridgeberry
Mitchella repens L., also called two-eyed berry or runningfoxis an evergreen vine or herb with fruit valuable as food
for birds, raccoons (Procyon lotor), and red foxes (Vulpes
vulpes) (Van Dersal 1938). The natural range is from Texas
to Florida, north to southwest Newfoundland, and west to
Ontario and Minnesota (Fernald 1950). This attractive plant
was introduced into cultivation in 1761 and is often used in
rock gardens (Rehder 1940).
Flowering and fruiting. The distylous flowers appear
from June to August and can be separated into 2 genetic
compatibility groups (Rehder 1940). Plants with short-styled
flowers (thrums) have exserted stamens 15 mm above the
ovary and stigmas 10 mm above the ovary; whereas plants
with long-styled flowers (pins) have stamens 11 mm
above the ovary and exserted stigmas 16 mm above the
ovary (Ganders 1975). The only pollinations that are compatible are those between anthers and stigmas of the same
height, that is, between pin and thrum and thrum and pin.
The genetic control is by a single gene (S), with thrums the
heterozygotes (Ss) and pins the recessive homozygotes (ss)
(Allard 1960). Thrums contribute more than three-quarters
of all genes transmitted through ovules, and pins more than
three-quarters of all genes transmitted through pollen (Hicks
and others 1985). Pins and thrums contribute almost equally
to gene flow via pollen and ovules.
The flowers occur in pairs on a short peduncle with the
base of the calyces fused. Each flower has 1 style and 4 stamens (Fernald 1950). Fruit-set occurs when both flowers of
a pair have been pollinated. Bumble bees (Bombus spp.) are
the principal pollinators of partridgeberry. They fly around a
patch of partridgeberry for a mean of 2.3 2.3 minutes, visiting 34 43 inflorescences per minute (Hicks and others
1985).
Fruits are scarlet drupaceous berries 7 to 10 mm wide
that ripen in July but usually persist overwinter (Petrides
1958). The maximum number of seeds that a single full
726
seed.
longitudinal
References
Allard RW. 1960. Principles of plant breeding. New York: John Wiley and
Sons. 485 p.
Barton LV, Crocker W. 1945. Twenty years of seed research. London: Faber
and Faber. 148 p.
Brinkman KA, Erdmann GG. 1974. Mitchella repens L., partridgeberry. In:
Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service: 543.
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1632 p.
Ganders FR. 1975. Fecundity in distylous and self-incompatible homostylous
plants of Mitchella repens (Rubiaceae). Evolution 29(1): 186188.
Hicks DJ, Wyatt R, Meacher TR. 1985. Reproductive biology of distylous
partridgeberry, Mitchella repens. American Journal of Botany 72(10):
15031514.
Keegan CR,Voss RH, Bawa KS. 1979. Heterostyly in Mitchella repens
(Rubiaceae). Rhodora 81: 567573.
Petrides GA. 1958. A field guide to trees and shrubs. Boston: Houghton
Mifflin. 431 p.
Rehder A. 1940. Manual of cultivated trees and shrubs. 2nd ed. New York:
Macmillan. 996 p.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA Soil
Conservation Service. 198 p
Van Dersal WR. 1938. Native woody plants of the United States: their erosion-control and wildlife values. Misc. Pub. 303. Washington, DC: USDA.
362 p.
Mitchella
727
MoraceaeMulberry family
Morus L.
mulberry
Jill R. Barbour, Ralph A. Read, and Robert L. Barnes
Ms. Barbour is a germination specialist at the USDA Forest Services National Seed Laboratory, Dry Branch,
Georgia; Dr. Read retired from the USDA Forest Services Rocky Mountain Research Station;
Dr. Barnes retired from the USDA Forest Services Southeastern Forest Experiment Station
Growth habit, occurrence, and use. The mulberry
genusMoruscomprises about 12 species of deciduous
trees and shrubs native to temperate and subtropical regions
of Asia, Europe, and North America (Rehder 1956). Seeds
of 2 native species and 2 naturalized species are described
here (table 1). White (sometimes called Russian) mulberry
was introduced to the United States by Mennonites from
Russia in 1875. The United States Prairie States Forestry
Project planted an average of over 1 million trees of this
species annually from 1937 through 1942 for windbreaks in
the Great Plains from Nebraska to northern Texas (Read and
Barnes 1974). The high drought resistance of white mulberry makes it well suited for shelterbelt planting (Read and
Barnes 1974).
There are 2 mulberries indigenous to North America.
Littleleaf mulberry occurs in Arizona, New Mexico,
Oklahoma, Texas, and Mexico and has not been cultivated
(table 1). Red mulberry has a wide range that covers most of
the eastern United States, Great Lakes region, and the southern Great Plains. Though once common, red mulberry is
decreasing over its range, possibly because of an unidenti-
Common name(s)
Occurrence
M. alba L.
M. alba var. tatarica (L.) Ser.
M. microphylla Buckl.
M. nigra L.
M. rubra L.
728
Morus
729
longitudinal
cleaned seeds.
Location
Flowering
Fruit ripening
M. alba
E US
Nebraska
Oklahoma
SW US
E US
May
May
Apr
AprMay
AprMay
JulyAug
JuneAug
Late MayJune
JuneAug
JuneAug
M. microphylla
M. rubra
Sources: Engstrom (1969), FNAEC (1997), Little and Delisle (1962), Read and Barnes (1974), Rehder (1956).
730
Table 3Morus, mulberry: height, seed-bearing age, seedcrop frequency, and fruit ripeness criteria
Height at
maturity (m)
M. alba
M. microphylla
M. nigra
M. rubra
314
7.5
10
12
Year first
cultivated
Minimum
seed-bearing
age (yr)
Years
between
large crops
1700s
1548
1629
10
Yearly
23
Sources: Little and Delisle (1962), Read and Barnes (1974), Rehder (1956), Sargent (1940), Small (1933).
Morus
731
/kg
286770
4401,100
Average
/lb
/kg
/lb
130350
200500
517
792
235
360
Samples
18+
4
Sources: Engstrom and Stoeckler (1941), Read and Barnes (1974), Swingle (1939).
References
Afanasiev M. 1942. Propagation of trees and shrubs by seed. Circ.C106.
Oklahoma Agricultural Experiment Station. 43 p.
Burton PJ, Bazzaz FA. 1991. Tree seedling emergence on interactive temperature and moisture gradients and in patches of old-field vegetation.
American Journal of Botany 78(1): 131149.
Core EL. 1974. Red mulberry, Morus rubra L. In: Gill JD, Healy WM, comp.
Shrubs and vines for northeastern wildlife. Gen.Tech. Rep. NE-9. Upper
Darby, PA: USDA Forest Service, Northeastern Forest Experiment
Station: 106107.
Davis WC, Wright E, Hartley C. 1942. Diseases of forest-tree nursery
stock. For. Pub. 9. Washington, DC: USDA Civilian Conservation Corps.
79 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Dirr MA. 1998. Manual of woody landscape plants: their identification,
ornamental characteristics, culture, propagation and uses. 5th ed.
Champaign, IL: Stipes Publishing. 1187 p.
Engstrom A. 1969. Personal communication. Oklahoma City: Oklahoma
Department of Agriculture, Forestry Division.
Engstrom HE, Stoeckler JH. 1941. Nursery practice for trees and shrubs
suitable for planting on the Prairie-Plains. Misc. Pub. 434. Washington,
DC: USDA. 159 p.
Fernald ML. 1970. Grays manual of botany. 8th ed. New York:Van
Nostrand Co. 1632 p.
FNAEC [Flora of North America Editorial Committee]. 1997. Flora of
North America North to Mexico.Volume 3, Magnoliophyta: Magnoliidae
and Hamamelidae. New York: Oxford University Press: 390392.
Griggs WH, Iwakiri BT. 1973. Development of seeded and parthenocarpic
fruits in mulberry (Morus rubra L.). Journal of Horticultural Science
48: 8397.
Halls LK. 1973. Flowering and fruiting of southern browse species. Res. Pap.
SO-90. New Orleans: USDA Forest Service, Southern Forest
Experiment Station. 10 p.
Heit CE. 1968. Thirty-five years testing of tree and shrub seed. Journal of
Forestry 66: 632634.
Hora B, ed. 1981. The Oxford encyclopedia of trees of the world. Oxford,
UK: Oxford University Press. 288 p.
Huffman GR. 1996. Seed handling and propagation of hardwood trees
and shrubs at Oklahoma Forestry Services Forest Regeneration Center.
In: Landis TD, South DB, tech coords. National Proceedings, Forest and
Conservation Nursery Associations. Gen.Tech. Rep. PNW-389. Portland,
OR: USDA Forest Service, Pacific Northwest Research Station: 4348.
ISTA [International Seed Testing Association]. 1999. International rules for
testing seeds: rules 1999. Seed Science and Technology 27 (Suppl.)
732
333 p.
Korves T. 1969. Correspondence. Fremont, NE: Plumfield Nursery.
Lamson NI. 1990. Morus rubra L., red mulberry. In: Burns RM, Honkala BH.
tech. coords. Silvics of North America.Volume 2, Hardwoods. Agric.
Handbk 654. Washington, DC: USDA Forest Service: 470473.
Little EL Jr, Delisle AL. 1962. Time periods in development: forest trees,
North American. In: Altman PL, Dittmer DS, eds. Growth including
reproduction and morphological development. Washington, DC:
Federation of American Societies for Experimental Biology: 382386.
Myatt A, Huffman G, Odell J. 1991. Seed processing manual. Washington,
OK: Oklahoma Department of Agriculture, Forestry Division, Forest
Regeneration Center.
Moore DM,Thomas WP. 1977. Red mulberry, Morus rubra L. In: Halls LK, ed.
Southern fruit-producing woody plants used by wildlife. Gen.Tech. Rep.
SO-16. New Orleans: USDA Forest Service, Southern Forest
Experiment Station: 5556.
Ottman Y. 1987. Rediscovering the realm of fruiting mulberry varieties. Fruit
Varieties Journal 41(1): 47.
Peaslee A. 2002. Personal communication. Jackson, NJ: New Jersey Forest
Tree Nursery.
Radford AE, Ahles HE, Bell CR. 1968. Manual of the vascular flora of the
Carolinas. Chapel Hill: University of North Carolina Press. 1183 p.
Read RA, Barnes RL. 1974. Morus L., mulberry. In: Schopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk 450.
Washington, DC: USDA Forest Service: 544547.
Rehder A. 1956. Manual of cultivated trees and shrubs hardy in North
America. 2nd ed. New York: Macmillan. 996 p.
Reich L. 1992. Native American fruits. Organic gardening 39(2): 52, 5457.
Sargent CS. 1940. Manual of the trees of North America (exclusive of
Mexico). 2nd ed., corrected and reprinted. New York: Dover. 934 p.
Small JK. 1933. Manual of the southeastern flora. New York: J.K. Small.
1554 p.
Stapanian MA. 1982. A model for fruiting display: seed dispersal by birds
for mulberry trees. Ecology 63(5): 14321443.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA Soil
Conservation Service. 198 p.
Taylor CA. 1941. Germination behavior of tree seeds. Washington, DC:
USDA Forest Service, Prairie States Forestry Project. 64 p.
USDA FS [USDA Forest Service]. 2002. Unpublished data. Dry Branch,
GA: National Tree Seed Laboratory.
Wasson E. 2001. Trees and shrubs: illustrated AZ of over 8500 plants.
Willoughby, NSW, Australia: Global Book Publishing: 483484.
Wright E. 1944. Damping-off in broadleaf nurseries of the Great Plains
region. Journal of Agricultural Research 69: 7794.
MyricaceaeBayberry family
Table 1Morella and Myrica, bayberry and wax-myrtle: nomenclature and occurrence
Scientific name & synonym(s)
Common name(s)
Occurrence
California wax-myrtle,
California bayberry,
Pacific bayberry
southern wax-myrtle,
southern bayberry
waxberry, candleberry
candleberry-myrtle
northern bayberry
bayberry, candleberry
Alaska to Newfoundland, S to
Pennsylvania,W to Wisconsin,Washington,
& Oregon; isolated areas of Tennessee &
North Carolina (swampy soils)
Coastal plain from Newfoundland
& Nova Scotia S to North Carolina
Myrica gale L.
Gale palustris Chev.
Myrica palustris Lam.
734
Table 2Morella and Myrica, bayberry and wax-myrtle: flowering and fruiting characteristics
Species
Flowering
Fruit
ripening
Morella californica
MayJune
Sept
Morella cerifera
Morella faya
Myrica pensylvanica
Mar-June
Variable
AprJuly
AugOct
AugNov
SeptOct
Myrica gale
MarApr
Oct
Diam of ripe
fruit (mm)
M
Cleaned seeds/wt
/kg
/lb
6
3
5
48,000
185,000
22,000
84,000
4
3
121,000
55,000
Sources: Fordham (1983), Huxley (1992), Krochmal (1974), Krssmann (1984), Schwintzer and Ostrofsky (1989), Walker (1990).
735
736
Nursery practice and seedling care. For field production, seeds can be sown in fall or spring. Fall-sowing
should be sufficiently late to avoid germination before winter, and seedbeds should be mulched. Spring-sowing should
follow a period of stratification at 5 C for 90 days
(Krochmal 1974). If container production is desired, seeds
may be sown indoors in early spring, and the seedlings
repotted before moving outdoors for further growth.
Germination is epigeal (figure 3) (Young and Young 1992).
Asexual propagation has been successful to varying
degrees depending on species. Blazich and Bonaminino
(1984) reported that terminal stem cuttings of southern waxmyrtle, in a transitional growth stage between softwood and
semi-hardwood, rooted in high percentages. Cuttings treated
with solutions of indolebutyric acid (IBA) at 0, 1,000
(0.1%), 2,000 (0.2%), or 4,000 ppm (0.4%) resulted in rooting of 87, 97, 87, and 90%, respectively. Cutting propagation of northern bayberry is more challenging. However,
softwood cuttings can be rooted successfully when treated
with a solution of 5,000 ppm (0.5%) IBA (Dirr and Heuser
1987). Most bayberry species produce root suckers and can
be propagated by division as well as by root cuttings (Dirr
and Heuser 1987).
References
Bir RE. 1992. Growing and propagating showy native woody plants. Chapel
Hill: University of North Carolina Press. 192 p.
Blazich FA, Bonaminio VP. 1984. Propagation of southern waxmyrtle by
stem cuttings. Journal of Environmental Horticulture 2(2): 4548.
Bornstein AJ. 1997. Myricaceae. In: Flora of North America Editorial
Committee, eds. Flora of North America north of Mexico.Volume 3,
Magnoliophyta: Magnoliidae and Hamamelidae. New York: Oxford
University Press: 429434.
Brackin RL. 1991. Wax myrtle named Lane. United States Plant Patent
7555.
Dirr MA, Heuser Jr. CW. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Elias TS. 1971. The genera of Myricaceae in the southeastern United States.
Journal of the Arnold Arboretum 52(2): 305318.
Everett TH. 1981. The New York Botanical Garden illustrated encyclopedia
of horticulture.Volume 7. New York: Garland Publishing: 21312492.
Fordham AJ. 1983. Of birds and bayberries: seed dispersal and propagation
of three Myrica species. Arnoldia 43(4): 2023.
Hamilton DF, Carpenter PL. 1977. Seed germination of Myrica pensylvanicum L. HortScience 12(6): 565566.
Huxley A, ed. 1992. The new Royal Horticultural Society dictionary of
gardening.Volume 3. New York: Stockton Press. 790 p.
Kartesz JT. 1994. A synonymized checklist of the vascular flora of the
United States, Canada, and Greenland. 2nd ed.Volume 1. Portland, OR:
Timber Press. 622 p.
Krochmal A. 1974. Myrica, bayberry. In: Schopmeyer CS, tech. coord. Seeds
of woody plants in the United States. Agric. Handbk. 450. Washington,
DC: USDA Forest Service: 548550.
Krssmann G. 1984. Manual of cultivated broad-leaved trees and shrubs.
Volume 1. Portland, OR:Timber Press. 624 p.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. 3rd ed.
New York: Macmillan. 1290 p.
Radford AE, Ahles HE, Bell CR. 1968. Manual of the vascular flora of the
Carolinas. Chapel Hill: University of North Carolina Press. 1183 p.
Schwintzer CR, Ostrofsky A. 1989. Factors affecting germination of Myrica
gale seeds. Canadian Journal of Forest Research 19(9): 11051109.
Small JK. 1933. Manual of the southeastern flora. Chapel Hill: University of
North Carolina Press. 1554 p.
Walker LR. 1990. Germination of an invading tree species (Myrica faya) in
Hawaii. Biotropica 22(2): 140145.
Weakley AS. 2000. Flora of the Carolinas and Virginia: working draft of 15
May 2000. Chapel Hill, NC: Nature Conservancy, Southeast Regional
Office, Southern Conservation Science Department [available in draft
from the author].
Wilbur RL. 1994. The Myricaceae of the United States and Canada: genera,
subgenera, and series. SIDA 16(1): 93107.
Young JA,Young CG. 1992. Seeds of woody plants in North America.
Portland, OR:Timber Press. 407 p.
737
HydrophyllaceaeWaterleaf family
738
seed.
longitudinal section
References
McDonald PM, Fiddler GO. 1995. Development of a mixed shrubponderosa pine community in a natural and treated condition. Res. Pap.
PSW-224-web. Albany, CA: USDA Forest Service, Pacific Southwest
Research Station. 19 p.
McDonald PM, Oliver WW. 1984. Woody shrubs retard growth of ponderosa pine seedlings and saplings. In: Proceedings, 5th Annual Forest
Vegetation Management Conference; 1983 November 23; Redding, CA.
Redding, CA: Forest Vegetation Management Conference: 6589.
Nord EC, Goodin JR. 1970. Rooting cuttings of shrub species for plantings
in California wildlands. Res. Note PSW-213. Berkeley, CA: USDA Forest
Service, Pacific Southwest Forest and Range Experiment Station. 4 p.
Nord EC, Leiser AT. 1974. Nama lobbii Gray, woolly nama. In: Schopmeyer
CS, tech. coord. Seeds of woody plants in the United States. Agric.
Handbk. 450. Washington, DC: USDA Forest Service: 551552.
Nama
739
BerberidaceaeBarberry family
740
seeds.
longitudinal sec-
Nandina
741
References
Afanasiev M. 1943. Germinating Nandina domestica seeds. American
Nurseryman 78 (9): 78.
Barr B. 1987. Propagation of dwarf nandina cultivars. Combined
Proceedings of the International Plant Propagators Society 37: 507508.
Bean WJ. 1976. Trees and shrubs hardy in the British Isles. 8th ed.Volume 3.
London: John Murray Co. p. 1.
Briggs BA, McCulloch SM. 1983. Progress in micropropagation of woody
plants in the United States and western Canada. Combined Proceedings
of the International Plant Propagators Society 33: 239248.
Coats AM. 1992. Garden shrubs and their histories. New York: Simon and
Schuster. 223 p.
Dehgan B. 1984. Germination of Nandina domestica seed as influenced by
GA3 and stratification. Proceedings of the Florida State Horticultural
Society 97: 311313.
Dirr MA. 1990. Manual of woody landscape plants: their identification,
ornamental characteristics, culture, propagation, and uses. 4th ed.
Champaign, IL: Stipes Publishing Co. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Flint HL. 1997. Landscape plants for eastern North America: exclusive of
Florida and the immediate Gulf Coast. 2nd ed. New York: John Wiley and
Sons. 842 p.
Gibson G. 1982. The bamboo alternative. American Horticulturist 61(10):
4, 710, 36.
742
AquifoliaceaeHolly family
Nemopanthus
743
References
N
(Adams 1927; Schopmeyer 1974). Cold stratification alone
did not increase germination rate (Adams 1927; Nichols
1934). Dirr and Heuser (1987) recommended 5 months of
warm followed by 3 months of cold stratification.
Propagation by greenwood cuttings is feasible (Bailey 1937;
Dirr and Heuser 1987).
744
Adams J. 1927. The germination of the seeds of some plants with fleshy
fruits, American Journal of Botany 14: 415425.
Baas P. 1984. Vegetative anatomy and the taxonomic status of Ilex collina
and Nemopanthus (Aquifoliaceae). Journal of the Arnold Arboretum 65:
243250.
Bailey LH. 1937. The standard cyclopedia of horticulture. 3 vol. New York:
Macmillan. 3639 p.
Begin J, Belanger L, Pfalzgraf J, Pineau M. 1990. Qualite de production dans
les erablieres rouges de la plaine Drummondville, Quebec. Forestry
Chronicle 66(4): 377386.
Bonner FT. 1974. Ilex L., holly. In: Schopmeyer CS, tech. coord. Seeds of
woody plants in the United States. Agric. Handbk. 450. Washington DC:
USDA Forest Service: 450453.
Curtis JT. 1959. The vegetation of Wisconsin: an ordination of plant
communities. Madison: University of Wisconsin Press: 655 p.
Cram H. 1988. A focus on peatlands and peat mosses. Ann Arbor:
University of Michigan Press. 306 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Farrar JL. 1995. Trees in Canada. Markham, ON: Fitzhenry and White. 502
p.
Gorchov DL. 1990. Pattern adaptation and constraint in fruiting synchrony
within verterbrate dispersed woody plants. Oikos 58: 169180.
LeBlanc CM, Leopold DJ. 1992. Demography and age structure of a central
New York shrub-carr 94 years after fire. Bulletin of the Torrey Botanical
Club 119(1): 5064.
Nichols GE. 1934. The influence of exposure to winter temperatures upon
the seed germination in various native American plants. Ecology 15:
364373.
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Rep. 88 (26.1), Northeast Region (region 1). Washington DC: USDI US
Fish and Wildlife Service, Northeast Region.
Rehder A. 1940. Manual of cultivated trees and shrubs. 2nd ed. New York:
Macmillan. 996 p.
Schopmeyer CS. 1974. Nemopanthus mucronatus (L.) Trel., mountain holly.
In: Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington DC: USDA Forest Service: 553.
Vitt DH, Slack NG. 1975. An analysis of the vegetation of sphagnum-dominated kettle-hole bogs in relation to environmental gradients. Canadian
Journal of Botany 53: 332359.
NyssaceaeSour-gum family
Nyssa L.
tupelo
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Common name(s)
Occurrence
N. aquatica L.
N. uniflora Wangenh.
2430
Ogeechee tupelo,
Ogeechee-lime, sour tupelo,
sour tupelo-gum, white tupelo
black tupelo, blackgum,
sourgum, tupelo-gum, pepperidge
1215
1518
N. biflora Walt.
N. sylvatica var. biflora (Walt.) Sarg.
N. sylvatica var. ursina
(Small) Wen & Stuessy
N. ogeche Bartr. ex. Marsh.
N. acuminata Small
N. sylvatica Marsh.
40
Flowering
Fruit ripening
Fruit drop
N. aquatica
N. biflora
N. ogeche
N. sylvatica
MarApr
AprJune
MarMay
AprJune
SeptOct
AugOct
JulyAug
SeptOct
Dark purple
Blue-black
Red
Blue-black
OctDec
SeptDec
NovDec
SeptNov
Sources: DeBell and Hook (1969), Kossuth and Scheer (1990), Radford and others (1964), Vande Linde (1964).
Nyssa
745
Figure 1Nyssa, tupelo: fruits of N. aquatica, water tupelo (upper left); N. sylvatica, black tupelo (upper right);
N. ogeche, Ogeechee tupelo (bottom).
746
longitudinal sec-
dried may float also (Kossuth and Scheer 1990), but fruits of
the other tupelos do not (McGee and Outcalt 1990). External
fruit color is the best index of maturity in the field (table 2).
To extract the seeds, the fruits should be run through a macerator with running water to float off the pulp. Small samples may be de-pulped by rubbing the fruits over a largemeshed screen, such as hardware cloth. For water tupelo,
observed numbers of fruits per weight have been from 340
to 600/kg (155 to 270/lb). Fifty kilograms (100 lb) of black
tupelo fruits should yield 12 kg (25 lb) of cleaned seeds
(Bonner 1974). Seed weights are listed in table 3.
Water tupelo seeds are orthodox in storage behavior.
They can be stored for at least 30 months in polyethylene
bags at either 3 or 10 C, if seed moisture contents are
<20% or <10%, respectively (Bonner and Kennedy 1973).
Seeds of black tupelo can be stored satisfactorily over 1
winter in cold, moist stratification in sand or in just cold
storage (Vande Linde 1964). Removal of the pulp did not
appear to be essential for retention of viability in either condition. There are no published storage data for other tupelo
species, but it is probable that the same methods would be
successful for them also.
Pregermination treatment. Tupelo seeds exhibit
moderate embryo dormancy, and they benefit from cold,
moist stratification. Treatment in moist sand and in plastic
bags without medium have been used successfully (Bonner
1974; DeBell and Hook 1969). Good germination has been
reported after only 30 days of stratification, but periods up
Collection place
N. aquatica
N. biflora
N. ogeche
N. sylvatica
South Carolina
North Carolina
Midwest
Avg
/kg
/lb
/kg
/lb
Samples
2,3003,100
4,1008,820
5,7508,500
1,0401,420
1,8504,000
2,6103,860
1,000
5,320
2,700
7,280
7,450
5,500
456
2,415
1,230
3,300
3,380
2,492
10
2
5
10
2+
Table 4Nyssa, tupelo: germination test conditions and results on stratified seeds
Species
N. aquatica
N. biflora
N. ogeche
N. sylvatica var.
sylvatica
Daily
light
(hr)
8
0
ND
8
8
Kimpak
Water in petri dish
Sand
Kimpak
Kimpak
30
29
30
30
20
29
20
20
27
28
60
70
27
Germination rate
Amt
(%)
Days
87
57
69
18
14
12
Germination %
Avg
(%) Samples
97
79
51
85
71
5
24
1
8
Purity
(%)
100
99
Nyssa
747
Figure 4Nyssa sylvatica, black tupelo: seedling development at 1, 4, and 39 days after germination.
Shading with tobacco shade cloth can help keep beds moist
and aid the newly emerged seedlings (Vande Linde 1964).
Germination is epigeal (figure 4). Desirable seedbed densities for water and black tupelos are 100 to 150 seedlings/m2
(9 to 14/ft2) (Williams and Hanks 1976. Vegetative propagation of tupelos is possible by softwood cuttings and grafting
(Dirr and Heuser 1987).
References
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
Bonner FT. 1974. Nyssa L., tupelo. In: Schopmeyer CS, tech. coord. Seeds of
woody plants in the United States. Agric. Handbk. 450. Washington, DC:
USDA Forest Service: 554557.
Bonner FT, Kennedy HE Jr. 1973. Storage of water tupelo seeds.Tree
Planters Notes 24(4): 78.
Brown CL, Kirkman LK. 1990. Trees of Georgia and adjacent states.
Portland, OR:Timber Press. 292 p.
DeBell DS, Hook DD. 1969. Seeding habits of swamp tupelo. Res. Pap. SE47. Asheville, NC: USDA Forest Service, Southeastern Forest Experiment
Station. 8 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant
propagation. Athens, GA:Varsity Press. 239 p.
DuBarry AP Jr. 1963. Germination of bottomland tree seed while
immersed in water. Journal of Forestry 61: 225226.
Earle FR, Jones Q. 1962. Analyses of seed samples from 113 plant families.
Economic Botany 16: 221250.
Heit CE. 1967. Propagation from seed: 8. Fall planting of fruit and hardwood seeds. American Nurseryman 126 (4): 1213, 8590.
Johnson RL. 1990.
Nyssa aquatica L., water tupelo. In: Burns RN, Honkala
BH, tech. coords. Silvics of North America.Volume 2, Hardwoods. Agric.
Handbk. 654. Washington, DC: USDA Forest Service: 474478.
748
Kossuth SV, Scheer RL. 1990. Nyssa ogeche Bartr. ex Marsh., Ogeechee tupelo. In: Burns RM, Honkala BH, tech. coord. Silvics of North America.
Volume 2, Hardwoods. Agric. Handbk. 654. Washington, DC: USDA
Forest Service: 479481.
Little EL Jr. 1978. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
McGee CE, Outcalt KW. 1990. Nyssa sylvatica Marsh., black tupelo. N. sylvatica Marsh. var. sylvatica; black tupelo (typical), N. sylvatica var. biflora
(Walt.) Sarg., swamp tupelo. In: Burns RM, Honkala BH, tech. coords.
Volume 2, Hardwoods. Agric. Handbk. 654. Washington, DC: USDA
Forest Service: 482489.
Priester DS. 1979. Stump sprouts of swamp and water tupelo produce
seeds. Southern Journal of Applied Forestry 3: 149151.
Radford AE, Ahles HE, Bell CR. 1964. Guide to the vascular flora of the
Carolinas. Chapel Hill: University of North Carolina Book Exchange.
1183 p.
Schneider RL, Sharitz RR. 1988. Hydrochory and regeneration in a bald
cypresswater tupelo swamp forest. Ecology 69: 10551063.
Vande Linde F. 1964. Nursery practices for southern oaks and gums.Tree
Planters Notes 65: 2426.
Williams RD, Hanks SH. 1976. Hardwood nurserymans guide. Agric.
Handbk. 473. Washington, DC: USDA Forest Service. 78 p.
RosaceaeRose family
Oemleria cerasiformis
(Torr. & Gray ex Hook. & Arn.) Landon
osoberry
William I. Stein
Dr. Stein is a forest ecologist emeritus at the USDA Forest Services Pacific
Northwest Research Station, Corvallis, Oregon
Oemleria
749
Flowering and Fruiting. Anatomical and natural population studies have confirmed strongly that osoberry is
dioecious, with male and female plants similar in size,
growth form, morphology of vegetative structures, and
microhabitats occupied (Allen and Antos 1988, 1993, 1999;
Antos and Allen 1990a; Sterling 1964). Flowering period in
osoberry is relatively short and varies with latitude and elevation from January to May concurrent with leaf development (Allen 1986; Haskin 1967; Hitchcock and others 1961;
McMinn 1970). Both male and female plants flower frequently except in low light; male plants are generally more
abundant and may have up to 3 times as many flowers as
female plants (Allen 1986; Allen and Antos 1988, 1993).
Male plants start flowering earlier than female plants but
reach peak abundance and finish flowering later (Allen
1986). First flowering has occurred 2 years after germination on male plants raised from seed (Allen and Antos
1993). The 5-petaled flowers are white, fragrant, and borne
on drooping racemes (figure 2). Osoberry pollen is sculptured and distinctive among Rosaceae pollens studied in
western Canada (Hebda and others 1991).
Pistillate flowers may yield up to 5 thin-fleshed, singleseeded drupes per flower, but generally fewer than 60% of
pistils on a plant bear fruit; production from 10 to 20 of pistils has been reported (Antos and Allen 1994, 1999). Higher
light levels favorably influence fruit set; exposure to light is
gained by early flowering before deciduous associates leaf
out (Allen and Antos 1988). Fruits develop and ripen in 10
to 12 weeks near Victoria, British Columbia (Antos and
Allen 1994). Developing fruits become peach colored, then
reddish, and finally deep blue-black under a whitish bloom
when ripe (figure 1). In the Pacific Northwest, dispersal by
gravity, birds, and mammals may begin in May and be nearly finished in July (Dimock and Stein 1974), substantially
750
white flowers
seeds have a
seedlings at 40
following year, they may germinate as early as midFebruary. Seeds collected in July, cleaned, and stored at
room temperature until sown outdoors in flats in late
December began germinating in March in Victoria, British
Columbia; second-year germination started in early
February and varied from 0 to 70% of total germination for
individual seedlots (Allen and Antos 1995). Total germination ranged from 1 to 96% among the 25 lots of 100 seeds
each representing 5 plants at each of 5 collection areas in
British Columbia and Washington.
Germination
during stratification (%)
Total
germination (%)
60
90
120
160
180
0
21
80
94
95
1
37
14
0
0
1
58
94
94
95
Oemleria
751
References
Abrams L. 1944. Illustrated flora of the Pacific States. Stanford, CA:
Stanford University Press.Volume 2. 635 p.
Allen GA. 1986. Flowering pattern and fruit production in the dioecious
shrub Oemleria cerasiformis (Rosaceae). Canadian Journal of Botany 64:
12161220.
Allen GA, Antos JA. 1988. Relative reproductive effort in males and
females of the dioecious shrub Oemleria cerasiformis. Oecologia 76:
111118.
Allen GA, Antos JA. 1993. Sex ratio variation in the dioecious shrub
Oemleria cerasiformis. American Naturalist 141: 537553.
Allen GA, Antos JA. 1995. Personal communication.Victoria, BC: University
of Victoria.
Antos JA, Allen GA. 1990a. A comparison of reproductive effort in the
dioecious shrub Oemleria cerasiformis using nitrogen, energy, and biomass
as currencies. American Midland Naturalist 124: 254262.
Antos JA, Allen GA. 1990b. Habitat relationships of the Pacific Coast shrub
Oemleria cerasiformis (Rosaceae). Madroo 37: 249260.
Antos JA, Allen GA. 1994. Biomass allocation among reproductive structures in the dioecious shrub Oemleria cerasiformisa functional
interpretation. Journal of Ecology 82: 2129.
Antos JA, Allen GA. 1999. Patterns of reproductiive effort in male and
female shrubs of Oemleria cerasiformis: a six-year study. Journal of
Ecology 87: 7784.
Dayton WA. 1931. Important western browse plants. Misc. Pub. 101.
Washington, DC: USDA. 214 p.
Dimock EJ II, Stein WI. 1974. Osmaronia (Torr. & Gray) Greene, osoberry.
In: Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service:
561563.
752
OleaceaeOlive family
Olea europaea L.
olive
George C. Martin
Dr. Martin is professor of pomology emeritus at the University of California, Davis, California
later than expected. When each leaf axil maintains a developing inflorescence, there are hundreds of flowers per twig.
Each inflorescence contains between 15 and 30 flowers,
depending on developmental processes for that year and the
cultivar.
The flowers are borne on the inflorescence and are
small, yellow-white, and inconspicuous. Each contains a
short, 4-segmented calyx and a short-tubed corolla containing 4 lobes. The 2 stamens are opposite on either side of the
2-loculed ovary that bears a short style and capitate stigma.
Two types of flowers are present each season: perfect flowers, containing stamen and pistil, and staminate flowers,
containing aborted pistils and functional stamens. The proportion of perfect and staminate flowers varies with inflorescence, cultivar, and year. Large commercial crops occur
when 1 or 2 perfect flowers are present among the 15 to 30
flowers per inflorescence. As a rule, more staminate flowers
than pistillate flowers are present.
The perfect flower is evidenced by its large pistil, which
nearly fills the space within the floral tube. The pistil is
green when immature and deep green when open at full
bloom. Staminate flower pistils are tiny, barely rising above
the floral tube base. The style is small and brown, greenish
white, or white, and the stigma is large and plumose as it is
in a functioning pistil.
The olive fruit is a drupe, botanically similar to almond,
apricot, cherry, nectarine, peach, and plum fruits. The olive
fruit consists of carpel, and the wall of the ovary has both
fleshy and dry portions. The skin (exocarp) is free of hairs
and contains stomata. The flesh (mesocarp) is the tissue
eaten, and the pit (endocarp) encloses the seed. Fruit shape
and size and pit size and surface morphology vary greatly
among cultivars.
The mature seed (figure 1) is covered with a thin coat
that covers the starch-filled endosperm (figure 2). The latter
surrounds the tapering, flat leaflike cotyledons, short radicle
Olea
753
stone.
Olea
755
Small
Medium
Large
/kg
/lb
706
198
99
320
90
45
353
265
220
Seeds/weight
/kg
/lb
4,410
1,654
992
2,000
750
450
sand or vermiculite and then placed in the dark in a controlled environment. The temperature is kept at 15 C for 30
days. Stratification is thought to reduce abscisic acid (which
inhibits germination) within the embryo or seedcoat. After
stratification, pits can be planted outdoors if the weather is
suitable; severe weather can cause losses. Pits can be planted in a greenhouse maintained at a 21 to 27 C daytime temperature. Bottom heat is necessary. Germination should
occur within 1 month. Transplanting seedlings from the
greenhouse to the nursery should include steps to harden the
seedlings, such as partial shade provided by a lathhouse.
Adequate irrigation and fertilization are recommended to
ensure continued rapid growth.
Nursery practice and seedling care. Virtually all
olive trees are produced from rooted cuttings. Seed handling
difficulties, low germination percentage, and slow initial
seedling growth rate make seedling production impractical.
References
Crisosto C, Suffer EG. 1985. Improving Manzanillo olive seed germination.
HortScience 20: 100102.
del Rio C, Caballero JM. 1994. Preliminary agronomical characterization of
131 cultivars introduced in the Olive Germplasm Bank of Cordoba in
March 1987. Acta Horticulturae 356: 110115.
Denney JO, Martin GC, Kammereck R, Ketchie DO, Connell JH, Krueger WH,
Osgood JW, Sibbeft GS, Nour GA. 1993. Freeze damage and coldhardiness in olive: findings from the 1990 freeze. California Agriculture 47:
112.
Lagarda A, Martin GC, Kester DE. 1983a. Influence of environment, seed
tissue and seed maturity on Manzanillo olive seed germination.
HortScience 18: 868869.
756
FabaceaePea family
dark, used for carvings, and will not float, its density being
1.22. The tree is threatened by introduced pasture grasses,
urbanization, and illegal harvesting for charcoal and artists
wood.
Flowering and fruiting. Flowering occurs from April
to June (Munz 1984; Shreve and Wiggins 1964). The pinkish to pale rose-purple flowers, 8 to 9 mm long, produce a
legume (pod) that may contain 1 to 2, or sometimes 3 or 4
or more seeds. The legume is light brown, rounded, and
hairy, and measures 4 to 6 cm in length (Munz 1984; Shreve
and Wiggins 1964). The seeds are chestnut brown to blackish, shiny, ovoid, and 8 to 9 mm long (figure 1) (Irving and
Becker 1985).
Collection and storage of fruits. Legumes on the tree
may be picked in June or July or fallen legumes and seeds
may be hand-gathered. The legumes dehise easily (Felker
1981). Many seeds are infested with insect larvae when collected, so the seeds should be stored cold or fumigated. Seed
Olneya
757
758
References
Becker R. 1983. Nutritional evaluation and chemical composition of seeds
from desert ironwood Olneya tesota. International Tree Crops Journal 2
(3/4): 297312.
Emery D. 1964. Seed propagation of native California plants. Leaflet. Santa
Barbara Botanical Garden 1(10): 8196.
Everett PC. 1957. A summary of the culture of California plants at the
Rancho Santa Ana Botanic Garden 19271950. Claremont, CA: Rancho
Santa Ana Botanical Garden. 223 p.
Felger RS, Moser, MB. 1985. People of the desert and sea: ethnobotany of
the Seri Indians. University of Arizona Press. 435 p.
Felker P. 1981. Uses of tree legumes in semiarid regions. Economic Botany
35(2): 174186.
Felker P, Clark PR. 1981. Nodulation and nitrogen fixation (acetylene
reduction) in desert ironwood (Olneya tesota). Oecologia 48(2):
292293.
Irving DW, Becker R. 1985. Seed structure and composition of potential
new crops. Food Microstructure 4(1): 4353.
Krugman SL. 1974. Olneya tesota, tesota. In: Schopmeyer CS, tech. coord.
Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 560.
Munz PA. 1974. A flora of Southern California. Berkeley: University of
California Press. 1086 p.
Nabhan GP, Carr JL, eds. 1994. Ironwood: an ecological and cultural keystone of the Sonoran Desert. Occ. Pap. 1. Washington DC: Conservation
International. 92 p.
Shreve F, Wiggins IL. 1964. Vegetation and flora of the Sonoran Desert.
Volume 1. Stanford, CA: Stanford University Press. 840 p.
Suzan H, Nabhan GP, Patten DT. 1994. Nurse plant and floral biology of a
rare night-blooming cereus Peniocereus striatus (Brandegee) f. Buxbaum.
Conservation Biology 8(2): 461470.
BetulaceaeBirch family
and Kirkman 1990; Sargent 1965). The fruits ripen from the
end of August in Michigan to October in the South. Nuts are
dispersed after ripening when the strobiles fall apart. The
buoyancy of the papery sacs aids dispersal by wind
(Metzger 1990). Trees do not produce seeds abundantly until
they are about 25 years old (Schopmeyer and Leak 1974).
Seed production in the northern part of the range has averaged 124,000 seeds/ha (50,200/ac) (Metzger 1990).
Collection, extraction, storage. The strobiles may be
hand-picked from the trees when they are a pale greenish
brown in color. At this stage, they are not yet dry enough to
fall apart. When completely ripe, they are light gray to
greenish brown (Schopmeyer and Leak 1974). The fruits
should be thoroughly dried before seeds are extracted by
thrashing or rubbing the dried fruits over screens. Seeds can
be separated from the chaff with air-screen cleaners or fractionating aspirators or by fanning. One hectoliter of fruit
will yield about 2.5 kg of seed (1 bu yields 2 lb). The number of seeds per weight (5 samples) ranged from 55,100 to
77,200/kg (25,000 to 35,000/lb), with an average of
66,100/kg (30,000/lb). Purities (percentages) in the high 90s
are easily obtained with good cleaning. The proportion of
sound seeds will vary widely, especially due to insect dam-
Ostrya
759
References
Brown CL, Kirkman LK. 1990. Trees of Georgia and adjacent states.
Portland, OR:Timber Press. 292 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation. Athens, GA:Varsity Press. 239 p.
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 451. Washington, DC: USDA Forest Service. 375 p.
Metzger FT. 1990. Ostrya virginiana (Mill.) K. Koch, eastern hophornbeam. In:
Burns RM, Honkala BH, tech. coord. Silvics of North America.Volume 2,
Hardwoods. Agric. Handbk. 654. Washington, DC: USDA Forest Service:
490496.
760
EricaceaeHeath family
Oxydendrum
761
762
Nursery practice. Johnson (1978) described a commercial method for seed propagation, in which seeds are
sown in November soon after harvest. Seeds are spread
lightly on the surface of a flat containing fine milled sphagnum and vermiculite (1:1, by vol.) and misted. Then, the flat
is wrapped in a clear plastic bag, with supports to keep the
bag from touching the surface of the medium, and placed
under continuous light, provided by cool-white fluorescent
lamps. Typically, the germination medium is maintained at
22 C using bottom heat. The medium surface should never
be allowed to dry. Seeds germinate within 2 weeks, and
seedlings develop rapidly. At the 2- to 3-leaf stage, seedlings
can be transplanted into peat pots or individual containers
containing an acidic, organic medium. After 6 months,
seedlings can be potted into 3.8-liter (1-gal) containers containing a well-drained, acidic, organic medium. Growth of
0.6 m (2 ft) can be obtained in 9 months following this production protocol. Blazich and others (1994) reported that
commercial production of seedlings of sourwood may be
accelerated by utilizing a pine bark medium and a day/night
cycle of 26/22 C or 30/26 C with long-day conditions.
Stem cuttings are reported as difficult to root (Dirr and
Heuser 1987). However, sourwood can be propagated vegetatively by micropropagation (Banko and Stefani 1989).
References
Bailey LH. 1977. Manual of cultivated plants. New York: Macmillan. 1116 p.
Banko TJ, Stefani MA. 1989. In vitro propagation of Oxydendrum arboreum
from mature trees. HortScience 24(4): 683685.
Barton SS, Bonaminio VP. 1985. Influence of light and temperature on
germination of sourwood (Oxydendrum arboreum (L.) DC.). Journal of
Environmental Horticulture 3(3): 108111.
Barton SS, Bonaminio VP. 1986. Influence of stratification and light on
germination of sourwood (Oxydendrum arboreum (L.) DC.). Journal of
Environmental Horticulture 4(1): 811.
Blazich FA. 1996. Unpublished data. Raleigh: North Carolina State
University.
Blazich FA, Warren SL,Thetford M. 1994. Growth response of sourwood as
influenced by temperature, photoperiod, and media. Proceedings,
Southern Nurserymens Association Research Conference, 39th Annual
Report: 137.
Bridwell FM. 1994. Landscape plants: their identification, culture and use.
Albany, NY: Delmar Publishers. 560 p.
DeWolf Jr. GP. 1987. Taylors guide to trees. New York: Chanticleer Press.
479 p.
Oxydendrum
763
FabaceaePea family
Germination. Seeds of peacock-plume exhibit seedcoat dormancy that can be overcome with acid scarification,
mechanical scarification, or hot-water soaking (Khullar and
others 1992; Wick and Walters 1974). The first 2 methods
have often produced slightly better results, but hot water
soaking is less likely to damage the seeds. Ten to 15 minutes
in concentrated sulfuric acid, followed by washing and then
15 minutes of soaking in water has been recommended
(Wick and Walters 1974). In hot-water soaking, seeds are
immersed in boiling water for 1 to 3 minutes, then soaked in
cool water at room temperature for 24 hours immediately
before sowing (Parrotta 1990). In a similar method, seeds
are immersed in boiling water that is then removed from the
heat source and allowed to cool at room temperature; the
seeds should remain in the water for 24 hours. Proper treat-
References
Desch HE. 1941. Manual of Malayan timbers,Volume 1. Kuala Lumpur:
Malayan Forestry Department. 328 p.
Gerhards CC. 1966. Physical and mechanical properties of Molucca albizia
grown in Hawaii. Res. Pap. FPL-55. Madison, WI: USDA Forest Service,
Forest Products Laboratory. 9 p.
Khullar P, Thapliyal RC, Beniwal BS,Vakshasya RK, Sharma A. 1992. Forest
seed. Dehra Dun, India: Indian Council of Forestry Research & Education.
409 p.
Little EL Jr, Skolmen RG. 1989. Common forest trees of Hawaii (native and
introduced). Agric. Handbk. 679. Washington, DC: USDA Forest Service.
321 p.
Little EL Jr, Wadsworth FH. 1964. Common trees of Puerto Rico and the
Virgin Islands. Agric. Handbk. 249. Washington, DC: USDA Forest
Service. 548 p.
Parrotta JA. 1990. Paraserianthes falcataria (L.) Nielsen: batai, Moluccan sau.
Res. Note SO-ITF-SM-31. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 5 p.
Rock JF. 1920. The leguminous plants of Hawaii. Honolulu: Hawaii Sugar
Planters Association. 234 p.
Wick HL, Walters GA. 1974. Albizia Durraz., albizia. In: Schopmeyer CS,
tech. coord. Seeds of woody plants in the United States. Agric. Handbk.
450. Washington, DC: 203205.
Paraserianthes
765
FabaceaePea family
Parkinsonia L.
palo verde
Kristina F. Connor, Jane E. Rodgers, and Carol Miller
Dr. Connor is a research plant physiologist at the USDA Forest Services Southern Research Station,
Auburn University, Alabama; Ms. Rodgers is now stationed at the Point Reyes National Seashore,
Point Reyes, California; Ms. Miller is a former propagationist at the Joshua Tree National Park,
Twentynine Palms, California
Common name(s)
Occurrence
P. aculeata L.
SW US
766
Bainbridge and Virginia (1989), Little (1979), Little and Wadsworth (1964).
SW US
seed.
Parkinsonia
767
longitudinal
References
Bainbridge DA,Virginia RA. 1989. Restoration in the Colorado Desert:
Species Notes. Unpublished manuscript prepared for the California
Department of Transportation.
Bean LJ, Saubel KS. 1972. Temalpakh (from the earth): Cahuilla Indian
knowledge and usage of plants. Morongo Indian Reservation, CA: Malki
Museum Press. 225 p.
Dean WRJ, Milton SJ. 1991. Patch disturbances in arid grassy dunes: antelope, rodents and annual plants. Journal of Arid Environments 20(2):
231237 [1992 Herbage Abstracts 62: 2115].
Delorit RJ, Gunn CR. 1986. Seeds of continental United States legumes
(Fabaceae). Park Falls, WI: F.A. Weber & Sons. 134 p.
Ebeling W. 1986. Handbook of Indian foods and fibers of arid America.
Berkeley: University of California Press. 971 p.
Everitt JH. 1983. Seed germination characteristics of two woody legumes
(retama and twisted acacia) from south Texas. Journal of Range
Management 36(4): 411414 [1984 Forestry Abstracts 45: 4409].
Felger RS, Moser MB. 1985. People of the desert and the sea.Tucson:
University of Arizona Press. 435 p.
Francis JK, Liogier HA. 1991. Naturalized exotic tree species in Puerto Rico.
Gen.Tech. Rep. SO-82. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 12 p.
Francis JK, Rodrguez A. 1993. Seeds of Puerto Rican trees and shrubs: second installment. Res. Note SO-374. New Orleans: USDA Forest Service,
Southern Forest Experiment Station. 5 p.
Jaegar EC. 1940. Desert wildflowers. Stanford, CA: Stanford University
Press. 322 p.
Jain KL, Kapil RP. 1980. Foraging rhythm of megachilid bees in relation to
the flowering of Medicago sativa L. and Parkinsonia aculeata L. Indian Bee
Journal 42(2): 3538 [Apicultural Abstracts 1973+].
768
VitaceaeGrape family
Parthenocissus Planch.
creeper
John D. Gill, Franz L. Pogge, and Franklin T. Bonner
Dr. Gill and Mr. Pogge retired from the USDA Forest Services Northeastern Forest Experiment Station;
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Common name(s)
Occurrence
Virginia creeper,
woodbine
Japanese creeper,
Boston ivy
Flowering
Fruit ripening
Fruit drop
P. quinquefolia
P. tricuspidata
JuneAug
JuneJuly
JulyOct
SeptOct
AugFeb
Sources: Fernald (1950), Gill and Pogge (1974), Rehder (1949),Van Dersal (1938).
Parthenocissus
769
770
References
Adams J.1927. The germination of the seeds of some plants with fleshy
fruits. American Journal of Botany 15 (8): 415428.
Bailey LH. 1914. The standard cyclopedia of horticulture. 6 vol. New York:
Macmillan. 3639 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seeds to tissue culture. Athens, GA:Varsity Press. 239 p.
Edminster FC. 1947. The ruffed grouse: its life story, ecology and management. New York: Macmillan. 385 p.
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1632 p.
Fisher PL, Briggs AH, Elkins WA, Roe EL. 1935. Propagation of game food
and cover plants of the Lake States. St. Paul: USDA Forest Service, Lake
States Forest Experiment Station. 81 p.
Flemion F. 1948. Reliability of the excised embryo method as a rapid test
for determining the germinative capacity of dormant seeds.
Contributions of the Boyce Thompson Institute 15: 229241.
Gill JD, Pogge FL. 1974. Parthenocissus Planch., creeper. In: Schopmeyer CS,
tech. coord. Seeds of woody plants in the United States. Agric. Handbk.
450. Washington, DC: USDA Forest Service: 568571.
Heit CE. 1955. The excised embryo method for testing germination quality
of dormant seed. Proceedings Association of Official Seed Analysts 1955:
108117.
Howard WL. 1915. An experimental study of the rest period in plants:
seeds. Res. Bull. 17. Columbia: Missouri Agricultural Experiment Station.
58 p.
Rehder A. 1949. Manual of cultivated trees and shrubs. Rev. ed. New York:
Macmillan. 1116 p.
Robinson FB. 1960. Useful trees and shrubs: reference cards [a card file of
data on hardy woody plants in common use as ornamentals].
Champaign, IL: Garrard Press. 427 cards.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA Soil
Conservation Service. 187 p.
Van Dersal WR. 1938. Native woody plants of the United States: their erosion-control and wildlife values. Misc. Pub. 303. Washington, DC: USDA.
362 p.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
Parthenocissus
771
ScrophulariaceaeFigwort family
8,800/hl
3,100/bu
2,033
2.8 kg/hl
2.2 lb/bu
6,200/g
175,770/oz
Percent moisture
content (fresh weight)
7%
References
Barnhill MA, Cunningham M, Farmer RE. 1982. Germination characteristics
of Paulownia tomentosa. Seed Science and Technology 10: 217221.
Beckjord PR. 1982. Containerized and nursery production of Paulownia
tomentosa. Tree Planters Notes 33(1): 2933.
Bonner FT. 1990. Paulownia tomentosa (Thunb.) Sieb. & Zucc. ex Steud.,
royal paulownia. In: Burns RM, Honkala BH, tech. coords. Silvics of North
America.Volume 2, Hardwoods. Agric. Handbk. 654. Washington, DC:
USDA Forest Service: 501502.
Bonner FT, Burton JD. 1974. Paulownia tomentosa (Thunb.) Sieb. & Zucc.,
royal paulownia. In: Schopmeyer CS, tech. coord. Seeds of woody plants
in the United States. Agric. Handbk. 450. Washington, DC: USDA Forest
Service: 572573.
Borthwick HA,Toole EH,Toole VK. 1964. Phytochrome control of Paulownia
seed germination. Israel Journal of Botany 13: 122133.
Carpenter SB, Smith ND. 1979. Germination of paulownia seeds after
stratification and dry storage.Tree Planters Notes 30(4): 46.
Paulownia
773
ScrophulariaceaeFigwort family
Penstemon Schmidel
penstemon, beardtongue
Susan E. Meyer
Dr. Meyer is a research ecologist at the USDA Forest Services Rocky Mountain Research Station,
Shrub Sciences Laboratory, Provo, Utah
Growth habit, occurrence, and use. The genus
Penstemon comprises about 230 species of perennial herbs
and subshrubs, most of which are found in western North
America. Although most of the species are herbaceous, there
are many more subshrubby species than are treated here.
Several shrubby species from California that were formerly
included in the genus Penstemon have been transferred to
the closely related genus Keckiella Straw. In the previous
edition of the Seed Manual, Hylton (1974) treated these
species under the name Penstemon.
Subshrubby penstemon species occur in most vegetation
types of the western United States, from warm desert shrublands to alpine fell-fields (table 1). They are most often
found on well-drained, rocky or sandy, infertile soils with
sunny exposure. Some species, such as Bridges penstemon,
are widely distributed and of wide ecological amplitude,
whereas others, such as crevice penstemon, are restricted
Common name(s)
Habitat*
Geographic distribution
P. ambiguus Torr.
moth penstemon,
bush penstemon,
gilia beardtongue
shrubby penstemon,
bush penstemon
Leonard penstemon,
Leonards beardtongue
toadflax penstemon
S Nevada to S Utah,
Kansas, & Oklahoma
P. fruticosus (Pursh)
Greene
P. leonardii Rydb.
P. linarioides Gray
P. petiolatus Brandeg.
P. platyphyllus Rydb.
P. rostriflorus Kellogg
P. bridgesii Gray
P. sepalulus A. Nels.
crevice penstemon,
petiole beardtongue
sidehill penstemon,
broadleaf beardtongue
Bridges penstemon
littlecup penstemon,
littlecup beardtongue
/g
/oz
/g
Range
/oz
Mean %
viability
Samples
1,000
3,500
1,250
810
2,700
1,460
2,260
1,700
28,000
98,000
35,000
23,000
77,000
42,000
64,000
48,000
8201,270
3,2303,850
9002,220
720900
2,6402,800
1,3901,590
1,5602,940
1,3502,000,
23,00036,000
90,000108,000
25,00062,000
20,00025,000
74,00078,000
39,00045,000
44,00082,000
38,00056,000
90
68
84
84
98
95
87
85
6
4
8
4
2
3
14
6
Penstemon
775
0 wk
4 wk
8 wk
12 wk
35
6
0
0
100
1
18
3
21
19
1
0
55
17
25
19
14
22
1
66
54
37
10
19
82
6
100
99
83
83
16 wk
16
40
80
12
100
97
86
24 wk
Samples
31
83
80
10
100
100
98
100
3
4
2
2
1
7
3
2
Sources: Kitchen and Meyer (1991), Meyer (2002), Meyer and others (1995).
* Germination percentage for seeds subjected to 0 to 24 weeks of chilling at 1 to 2 C followed by 4 weeks of incubation at 10/20 C.
The germination requirements of penstemon seeds generally change very little in dry storage; dormancy status is
affected only by conditions during time spent in the imbibed
state (Meyer and others 1995). For species and populations
from middle elevations in the West, there is rarely a natural
dormancy-breaking treatment that will remove dormancy in
all seeds of a lot. For those that respond positively to chilling, this is manifest as a fraction that will not respond to
chilling of any duration. These seeds form a persistent seedbank under natural conditions, and it is not known how they
eventually become germinable. Treatment with gibberellic
acid can remove seed dormancy or shorten the chilling
requirement for many (but not all) species of penstemon
(Kitchen and Meyer 1991). This method may or may not be
feasible in a production setting, depending on the degree to
which gibberellic acid affects seedling quality. Penstemon
seeds germinate best at cool temperatures, and germination
at temperatures higher than 20 C may be completely suppressed, a fact to keep in mind when attempting to produce
plants from direct sowing in the greenhouse (Allen and
Meyer 1990). Light usually has little effect (Meyer and others 1995).
The quality of penstemon seeds may be evaluated using
tetrazolium (TZ) staining, a germination test with chilling or
gibberellic acid, or a combination of these. A general seedtesting rule for the genus has been adopted by the Association of Official Seed Analysts (Kitchen and others 1999).
This test calls for 2 separate procedures. In the first procedure, seeds are placed on blotters saturated with 500 ppm
gibberellic acid, chilled at 2 to 5 C for 60 days, and incubated at 15 or 10/20 C for 14 days. Post-test viability of
ungerminated seeds is then determined with TZ staining.
This procedure is used to determine total viability for the
seedlot, and TZ staining on non-incubated seeds may be
substituted. In the second procedure, seeds are incubated at
776
15 or 10/20 C in the light for 28 days. This second procedure is used to determine the size of the nondormant fraction.
Tetrazolium staining is carried out by allowing the seeds
to imbibe water for 24 hours, piercing their seedcoats with a
needle, immersing them in 1% TZ for 12 hours at room temperature, and then bisecting them longitudinally for evaluation. The embryo is a small, sausage-shaped body embedded
in the copious endosperm. Non-viable embryos usually do
not stain at all but remain a yellowish color. Even light pink
staining indicates a viable embryo, as evidenced by comparisons with maximum germination percentages for numerous
seedlots (Kitchen and Meyer 1991). A simple cut test can be
used in place of TZ staining for recently harvested seeds, as
the presence of a firm, white, viable embryo is quite evident.
Field planting and nursery practice. Most penstemon species can be established from direct seeding. Seeds
should be broadcast on a firm seedbed and lightly covered,
raked, or pressed in. Planting should take place in late fall or
early winter for most species, except in the summer rainfall
areas of the Southwest, where seeds of nondormant species
should be sown just before summer rains. Nondormant
species may be spring-seeded in the North but may require
supplemental water to establish. Penstemons are susceptible
to fusarium wilt diseases in cultivation and should not be
fertilized or overwatered. They are more likely to survive in
coarse, rapidly draining soils that have not previously been
used for agriculture. The young seedlings cannot survive
heavy competition from weeds or aggressive perennial
grasses.
Penstemons may also be readily produced from seeds in
container culture. They grow well in a coarse medium in
elongated containers such as those used to produce conifer
seedlings. Seeds of nondormant species may be direct-sown,
whereas chilled seeds or germlings may be planted for those
References
Allen PS, Meyer SE. 1990. Temperature requirements for seed germination
of three Penstemon species. HortScience 25: 191193.
Cronquist, A. Holmgren, AH, Holmgren NH, Reveal JL, Holmgren PK. 1984.
Intermountain flora.Volume 4, Subclass Asteridae (except Asteraceae).
Bronx, NY: New York Botanical Garden. 573 p.
Hylton LO. 1974. Penstemon Mitch., penstemon. In: Schopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 574575.
Kitchen SG, Meyer SE. 1991. Seed germination of intermountain penstemons as influenced by stratification and GA3 treatments. Journal of
Environmental Horticulture 9: 5156.
Kitchen SG, Meyer SE, Stevens R, Wilson GR. 1999. Proposed rule: a generalized rule for the genus Penstemon, beardtongues. Seed Technologist
Newsletter 72(1): 52.
Meyer SE. 2002. Unpublished data. Provo, UT: USDA Forest Service, Rocky
Mountain Research Station.
Penstemon
777
RosaceaeRose family
778
seeds.
value for seeds stratified only 35 days. Viable seeds from the
same lot kept in low-temperature stratification (1 C)
showed 50% germination by day 82 and about 95% by day
120. Results from seeds collected the next year (1995) were
similar. Smith (1974) reported somewhat lower germination
percentages at longer chill durations. When seeds were tested at 30/20 C (8/16 hours, day/night) after 0, 30, 60, and 90
days of cold stratification, germination of squaw-apple averaged 9, 9, 16, and 51% of total seeds.
Nursery practice. Squaw-apple is grown in nursery
beds only occasionally, usually when requested for transplantation at age 1 to 2 years into native-plant gardens (Prag
and Prag 1996). In the greenhouse, squaw-apple seedlings
emerge in 6 to 12 days from seeds planted about 5 mm
(0.2 in) deep and covered with a thin layer of fine sand
(Smith 1974). Overwatering and transplantation during the
growing season increase the risk of seedling mortality (Prag
and Prag 1996). Establishment is rated fair and persistence
is very good (Plummer and others 1968; Shaw and Monsen
2004).
Peraphyllum
779
References
Auger J. 1994. Viability and germination of seeds from seven fleshy-fruited
shrubs after passage through the American black bear (Ursus
americanus) [MS thesis]. Provo, UT: Brigham Young University. 49 p.
Auger J. 2002. Unpublished data. Reno: University of Nevada.
Auger J, Black HL, Meyer SE. 1995. Seed dispersal ecology of squawapple
(Rosaceae: Peraphyllum ramosissimum). Final report, 29 October 1995 on
file. Provo, UT: USDA Forest Service, Intermountain Research Station,
Shrub Sciences Laboratory.
Dayton WA. 1931. Important western browse plants. Misc. Pub. 101.
Washington, DC: USDA. 214 p.
Harrington HD. 1954. Manual of the plants of Colorado. Denver: Sage
Books.
Hitchcock CL, Cronquist A, Ownbey M, Thompson JW. 1961. Vascular
plants of the Pacific Northwest. Part 3. Seattle: University of Washington
Press. 614 p.
780
LauraceaeLaurel family
fruits.
cleaned seed.
781
longitudinal section
References
Brendemuehl RH. 1990. Persea borbonia (L.) Spreng., redbay. In: Burns RM,
Honkala BH, tech. coords. Silvics of North America.Volume 2,
Hardwoods. Agric. Handbk. 654. Washington, DC: USDA Forest Service:
503506.
Brown CL, Kirkman LK. 1990. Trees of Georgia and adjacent states.
Portland, OR:Timber Press. 292 p.
King MW, Roberts EH. 1980. Maintenance of recalcitrant seeds in storage.
In: Chin HF, Roberts EH, eds. Recalcitrant crop seeds. Kuala Lumpur:
Tropical Press: 5389.
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Radford AE, Ahles HE, Bell CR. 1968. Manual of the vascular flora of the
Carolinas. Chapel Hill: University of North Carolina Press. 1183 p.
Sargent CS. 1965. Manual of the trees of North America (exclusive of
Mexico). 2nd. Ed., corrected and reprinted. New York: Dover. 934 p.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
782
RutaceaeRue family
Growth habit, occurrence, and use. Amur corktreePhellodendron amurense Rupr.is native to northern
China, Manchuria, Korea, and Japan. This small to medium
deciduous tree25 to 50 feet tallhas been cultivated in
the Far East and eastern Europe. It was introduced into the
United States around 1865, and its thick, corky bark and
massive, irregular branches have made it of special interest
for landscape and environmental plantings in the northern
and western United States (Blackburn 1952; Everett 1964;
Hoag 1965; Lewis 1957). In tests in Kansas, however, the
tree did not perform well and was not recommended for
general use (Hensley and others 1991). It is a potential
source of industrial cork (Izmodenov 1972; Ota and others
1965), important as a nectar-bearing species in bee-keeping
areas of the Russian Far East (Necaev and Pelemenev 1965),
and of possible importance for the insecticidal properties of
its fruit oils (Schechter 1943). In Byelorussia it is considered
a soil builder when mixed with Scots pinePinus
sylvestris L. (Letkovskij 1960). It tolerates a wide range of
soil conditions, pH, drought, and pollution; it is easily transplanted and generally free of pests.
Flowering and fruiting. The species is dioecious, and
female plants develop tend to have a bushier form than
males (Hensley and others 1991). Small, yellowish green
flowers, in large clusters of terminal panicles, appear in May
and June (Krecetova 1960; Rehder 1940; Schechter 1943).
Climate affects the time of day when flowers open, pollination, and the longevity of flowers (Starshova 1972).
Fruits are subglobose drupes about 1 cm in diameter
(figure 1), green to yellowish green, turning black when ripe
in September and October (Read 1974). They remain on the
terminal panicles long after the leaves have dropped. Fruits
are borne singly on short stalks (figure 1) and are very oily
and aromatic. Each fruit usually contains 2 or 3 full-sized
seeds and 3 or 4 aborted seeds (Read 1974). Seeds are
brown to black, 5 mm long, 2 mm wide, and about 1 mm
thick (figures 2 and 3); they have a moderately hard, stony
coat (Gorokhova 1981; Read 1974).
fruit
Phellodendron
783
Germination for a seedlot (for which the handling and storage history was not described) was best following cold
moist, underground stratification for 166 days (Timm 1989).
In another study, seeds stratified for 8 weeks had a higher
germination rate and the same germination percentage as
seeds stratified 4 weeks; germination of unstratified seeds
was less than half that of stratified seeds (Mukai and
Yokoyama 1985). Based on the information available, it is
recommended that seeds be stratified if the history of collection, handling, and storage is not documented. Seeds of
other Phellodendron spp. vary in their requirements for
stratification (Dirr and Heuser 1987; Lin and others 1994).
Seeds germinate best at alternating temperatures. Both
Lin and others (1979) and Mukai and Yokoyama (1985)
reported germination of 3% or less at constant temperatures
and 75 to 90% at alternating temperatures. The best temperature regimes were 35/5 C and 35/15 C (day length and
high temperature for 8 hours).
Nursery practice and natural regeneration. In its
natural range, natural regeneration sometimes occurs in
dense groups. Although the corktree has been reported to
sucker from its roots (Dirr and Heuser 1987), this dense
regeneration is believed to be from seeds present in the forest floor (Soludukin 1977). Light fire or disturbances that
result in drying and warming of the forest floor are believed
to promote this development. There was no indication
regarding the longevity of the seeds in the forest floor environment, however the endocarp is moderately hard and
might facilitate longevity under these conditions (Soludukin
1977).
In the nursery, untreated seeds may be sown in the fall
(Read 1974) or stratified through winter for spring-seeding
(Yerkes 1945). It is suggested that the best time for spring
sowing is when the mean daily soil temperature has reached
8 to 10 C (Antonyuk 1987). Trees may also be propagated
vegetatively by root cuttings or shoot cuttings (Bailey 1947;
Dirr and Heuser 1987).
References
Antonyuk ED. 1987. Growth of seedlings in a polythene greenhouse in
relation to sowing times. Lesnoe Khozyaistvo 4: 3637.
Atkimockin NG. 1960. [in Russian:Trials with Phellodendron spp. at the
Forest-Steppe Experiment Station [abstract]. Byulleten Glavnogo
Botanicheskogo Sada [Moskva] 37: 3033.
Bailey LH. 1947. Standard cyclopedia of horticulture. 2nd ed. New York:
Macmillan. 1388 p.
Blackburn BC. 1952. Trees and shrubs of eastern North America. New
York: Oxford University Press. 358 p.
Dirr MA. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation, and uses. Champaign, IL:
Stipes Publishing. 1007 p.
784
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant cultivation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Everett TH. 1964. New illustrated encyclopedia of gardening.Volume 8.
New York: Greystone Press: 14551456.
Gorokhova GI. 1981. Fruiting of far-eastern species of plants under conditions of the forest steppe zone of western Siberia. Soviet Journal of
Ecology 12 (4): 219222 [translated and published 1982].
Gorokhova GI. 1986. Effect of drought on the seed quality of exotic plants
from the Soviet East [abstract]. Byulleten Glavnogo Botanicheskogo
Sada [Moskva] 139: 8589.
Hensley DL, Wiest SC, Long CE, Pair JC, Gibbons FD III. 1991. Evaluation
of 10 landscape trees for the Midwest. Journal of Environmental
Horticulture 9(3): 149155.
Hoag DG. 1965. Trees and shrubs of the northern plains. Minneapolis: Lund
Press. 376 p.
Izmodenov AG. 1972. Multiple use of Pinus koraiensis forests [English
abstract]. Moscow: Lenaya Promyshlenoost. 122 p.
Krecetova NV. 1960. [in Russian:The biology of flowering and fertilization in
P. amurense. abstract]. Botanicheskii Zhurnal 45(9): 13361340.
Letkovskij AI. 1960. [in Russian: Planting Phellodendron amurense to increase
the productivity of Scots pine forest (in Belorussia.)] Lesnaya Hortus
12(1): 3637 [Forestry Abstracts 23: 548, 1962].
Lewis CE. 1957. American elm substitutes. American Nurseryman 106(10):
1011, 5051.
Lin TP,TW Hu, Leu JJ. 1979. The effect of alternating temperatures on the
germination of the seed of Phellodendron spp. [abstract]. Bulletin of the
Taiwan Forest Research Institute 322: 110.
Lin TP, Chen MH, Lin CH. 1994. Dormancy in seeds of Phellodendron wilsonii
is mediated in part by abscisic acid. Plant Cell Physiology 35(1):
115119.
Mukai Y, Yokoyama T. 1985. Effect of stratification on seed germination of
Phellodendron amurense [abstract]. Journal of the Japanese Forestry
Society 67(3): 103104.
Necaev AD, Pelmenev VK. 1965. [in Russian:The importance of Phellodendron amurense as a nectar-bearing species.] Rastitelnye Resursy
[Moskva] 1(3): 419423 [Forestry Abstracts 28: 621; 1967.
Ota M and others. 1965. [in Japanese: A study of the manufacturing conditions and properties of SIS hardboard from the inner bark of Kihada
(Phellodendron amurense Rupr.).] Wood Industries Tokyo 20(6): 1316
[Forestry Abstracts 27: 1475; 1966].
Read RA. 1974. Phellodendron amurense Rupr., Amur corktree. In:
Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service:
578579.
Rehder A. 1940. Manual of cultivated trees and shrubs. 2nd ed. New York:
Macmillan. 996 p.
Schechter MS. 1943. The insecticidal principle in the fruit of Amur corktree
(Phellodendron amurense). Journal of Organic Chemistry 8: 194197.
Soludukhin ED. 1977. Features of the regeneration of Phellodendron
amurense [abstract]. Lesnoe Khozyaistvo 4: 3233.
Starshova NP. 1972. The anthecology of Phellodendron amurense in the
middle Volga region: 1. Ecology of flowering. Botanicheskii Zhurnal
57(11): 14021412.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP-27, Washington, DC: USDA Soil
Conservation Service. 198 p.
Timm GT. 1989. Effect of stratification on the field germination rate of
some tree seed. Journal of the Korean Forestry Society 78(1): 2629.
Yerkes GE. 1945. Propagation of trees and shrubs. Farm. Bull. 1567 (rev.)
Washington, DC: USDA. 54 p.
Phellodendron
785
HydrangeaceaeHydrangea family
Philadelphus L.
mock orange
Nancy L. Shaw, Emerenciana G. Hurd, and Peter F. Stickney
Dr. Shaw is a research botanist at the USDA Forest Services Rocky Mountain Research Station, Forestry
Sciences Laboratory, Boise, Idaho; Dr. Hurd and Mr. Stickney retired from the USDA Forest Services
Rocky Mountain Research Station
Common name(s)
Occurrence
P. lewisii Pursh
P. californicus Benth.
P. gordonianus Lindl.
P. columbianus Koehne
P. gordonianus var. columbianus Rehd.
P. microphyllus Gray
P. nitidus A. Nels.
P. stramineus Rydb.
Sources: Conquist and others (1997), Davis (1952), Hickman (1974), USDA ARS (2002),Welsh and others (1987).
* This common name, although widely used, creates some confusion, as it also the generic name of the lilacs, to which the mock orange bears some resemblance.
786
Service 1937). Quail (Callipepla spp.) and squirrels consume Lewis mock orange seeds (Van Dersal 1938). Mule
deer (O. hemionus) browse littleleaf mock orange (Patton
and Ertl 1982).
Mock oranges are valuable plants for revegetating disturbances on steep, rocky, unstable slopes within their native
ranges (Stevens and others 2004). Seedlings or larger stock
are recommended for planting such sites. Mock oranges are
also useful for planting in drier areas of degraded riparian
zones.
Mock orange species and their cultivars are used as
ornamentals. Lewis mock orange was first cultivated in 1823
or 1884, and littleleaf mock orange in 1883 (Rehder 1940).
Both are used as borders, screens, and hedges or as isolated
specimens in sunny areas. They can also be used for lowmaintenance landscaping and in recreational area plantings
(Kruckeberg 1982, Sutton and Johnson 1974, Wright 1980).
They do well on a wide variety of soils and require little
maintenance. Plants grow vigorously, flower reliably, and
are generally free of insect and disease problems.
Native Americans used stems of Lewis mock orange for
making arrows (USDA FS 1937). Flowers are used in
preparing perfumes and teas (Taylor 1972).
Genetic variation and hybridization. Natural variability in mock orange floral and vegetative characteristics is
extensive and has contributed to development of the complex synonymy for each species (Cronquist and others 1997;
Hickman 1993; Hitchcock and others 1961; Holmgren and
Reveal 1966; Hu 1955; Rydberg 1905). This variability has
been exploited to develop numerous hybrids (Rehder 1940;
Rydberg 1905) and several ornamental cultivars (Wright
1980). Waterton Lewis mock orange, selected from the
Waterton Lakes area of Alberta, is a hardy, bushy shrub with
flowers scattered over the crown of the plant (Taylor 1972).
The P. lemoinei (P. coronarius P. microphyllus) group of
hybrids exhibit the pineapple scent and beauty of their littleleaf mock orange parent (Sutton and Johnson 1974; Wright
1980).
Flowering and fruiting. Mock orange flowers are
white, fragrant, and showy, with 4 (5) petals and numerous
stamens. They are produced in few-flowered cymes at the
ends of shoots formed the previous year. Western species
flower from May to July (Hitchcock and others 1961; Munz
and Keck 1973; Orme and Leege 1980). Fruits are woody,
turbinate, loculicidal capsules that mature in late summer
(figure 1); those of Lewis mock orange dehisce in
September or October (Marchant and Sherlock 1984; Orme
and Leege 1980). The seeds are dispersed by wind and
gravity.
Philadelphus
longi-
787
Seed fill
(%)
Viability
Wet
(%)
prechill (days)
Percentage germination
15 C
20/10 C
98
98
100
100
100
100
99
99
96
96
92
92
90
90
96
96
0
28
0
28
0
28
0
28
1
57
12
41
8
47
2
34
1
16
6
23
5
33
2
33
98
98
63
63
60
28
5
44
20
788
References
Atthowe H. 1993. Propagation of riparian and wetland plants. In: Landis TD,
tech. coord. Proceedings, Western Forest Nursery Association; 1992
September 1418; Fallen Leaf Lake, CA. Gen.Tech. Rep. RM-221. Fort
Collins, CO: USDA Forest Service, Rocky Mountain Forest and Range
Experiment Station: 7881.
Cronquist A, Holmgren NH, Holmgren PK. 1997. Intermountain flora,
vascular plants of the Intermountain West, U.S.A.Volume 3, Part A subclass Rosidae (except Fabales). Bronx., NY: New York Botanical Garden.
446 p.
Dirr MA, Heuser CW. 1987. The reference manual of woody plant propagation. Athens, GA:Varsity Press. 239 p.
Everett PC. 1957. A summary of the culture of California plants at the
Rancho Santa Ana Botanic Garden. Claremont, CA: Rancho Santa Ana
Botanic Garden. 223 p.
Glazebrook TB. 1941. Overcoming delayed germination in the seed of
plants valuable for erosion control and wildlife utilization [thesis].
Moscow, ID: University of Idaho. 97 p.
Harrington HD. 1954. Manual of the plants of Colorado. Denver: Sage
Books. 665 p.
Hartmann HT, Kester DE, Davies FT Jr. 1990. Plant propagation principles
and practices. 5th ed. Englewood Cliffs, NJ: Prentice-Hall. 657 p.
Hickman JC. 1993. The Jepson manual: higher plants of California. Berkeley:
University of California Press. 1400 p.
Hitchcock CL. 1943. The xerophyllous species of Philadelphus in southwestern North America. Madroo 17: 3556.
Hitchcock CL, Cronquist A, Ownbey M,Thompson JW. 1961. Vascular
plants of the Pacific Northwest: 3. Saxifragaceae to Ericaceae. Seattle:
University of Washington Press. 614 p.
Holmgren AH, Reveal JL. 1996. Checklist of the vascular plants of the
Intermountain Region. Res. Pap. INT-32. Ogden, UT: USDA Forest
Service, Intermountain Forest and Range Experiment Station.
Hopkins WE, Kovalchik BL. 1983. Plant associations of the Crooked River
National Grassland. R6 Ecol. 133-1983. Portland, OR: USDA Forest
Service, Pacific Northwest Region. 98 p.
Hu SY. 1955. A monograph of the genus Philadelphus. Journal of the Arnold
Arboretum 36: 52109.
Hurd EG. 1995. Unpublished data. Boise, ID: USDA Forest Service,
Intermountain Research Station.
Jorgensen K. 2004. Appendix II. In: Monsen SB, Stevens R, Shaw NL, comps.
Restoring western ranges and wildlands Gen.Tech. Rep. RMRS-136. Fort
Collins, CO: USDA Forest Service, Rocky Mountain Research Station.
Kruckeberg AR. 1982. Gardening with native plants of the Pacific
Northwest. Seattle: University of Washington Press. 252 p.
Kufeld RC. 1973. Foods eaten by the Rocky Mountain elk. Journal of Range
Management 26: 106113.
Leege TA. 1968. Prescribed burning for elk in northern Idaho. Proceedings
of the Annual Tall Timbers Fire Ecology Conference 8: 235253.
Leege TA, Hickey WO. 1971. Sprouting of northern Idaho shrubs after
prescribed burning. Journal of Wildlife Management. 35(3): 508515.
Macdonald B. 1986. Practical woody plant propagation for nursery
growers.Vol. I. Portland, OR:Timber Press. 669 p.
Marchant C, Sherlock J. 1984. A guide to selection and propagation of
some native woody species for land rehabilitation in British Columbia.
For. Res. Rep. RR-84007-HQ.Victoria, BC: Ministry of Forests. 117 p.
Mirov NT, Kraebel CJ. 1939. Collecting and handling seeds of wild plants.
For. Pub. 5. Washington, DC: USDA Civilian Conservation Corps. 42 p.
Munz PA, Keck DD. 1973. A California flora and supplement. Berkeley:
University of California Press. 1681 p.
Orme ML, Leege TA. 1980. Phenology of shrubs on a north Idaho elk
range. Northwest Science 54: 187198.
Patton DR, Ertl MG. 1982. Run wild: wildlife/habitat relationships. Wildlife
Unit Tech. Ser. Washington, DC: USDA Forest Service. 49 p.
Rehder A. 1940. Manual of cultivated trees and shrubs. 2d ed. New York:
Macmillan. 996 p.
Rydberg PA. 1905. North American Flora 22: 162175.
Shaw NL. 1995. Unpublished data. Boise, ID: USDA Forest Service,
Intermountain Research Station.
Stevens R, Monsen SB, Shaw NL. 2004. Shrubs of other families. In: Monsen
SB, Stevens R, comps. Restoring western ranges and wildlands Gen.Tech.
Rep. RMRS-136. Fort Collins, CO: USDA Forest Service, Rocky
Mountain Research Station.
Stickney PF. 1974. Philadelphus lewisii Pursh, Lewis mockorange. In:
Shopmeyer CS, tech. coord. Seeds of woody plants in the United States.
USDA Agric. Handbk. 450. Washington, DC: USDA Forest Service:
580581.
Sutton R, Johnson CW. 1974. Landscape plants from Utahs mountains. EC368. Logan: Utah State University. 137 p.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA Soil
Conservation Service. 198 p.
Taylor S. 1972. Philadelphus lewisii Pursh mock orange or syringa.
Davidsonia 3: 47.
USDA ARS [USDA Agricultural Research Service]. 2001. Germplasm
Information Network (GRIN)[online datbase]. Beltsville, MD: National
Genetic Resources Program.
USDA Forest Service. 1937. Range plant handbook. Washington, DC. 512 p.
USDA NRCS [USDA National Resources Conservation Service]. 2001. The
PLANTS Database, version 3.1[http://plants.usda.gov]. Baton Rouge, LA:
USDA NRCS National Plant Data Center.
Van Dersal WR. 1938. Native woody plants of the United States: their erosion control and wildlife values. Misc. Pub. 303. Washington, DC: USDA.
362 p.
Welsh SL, Atwood ND, Higgins LC, Goodrich S. 1987. A Utah flora. Great
Basin Naturalist 9: 1894.
Wright D. 1980. Philadelphus. Plantsman 2: 104116.
Youtie BA. 1991. Native plants delight visitors at Columbia Gorge plot.
Park Science 11: 45.
Youtie BA. 1992. Biscuit scabland restoration includes propagation studies
(Oregon). Restoration & Management Notes 10: 7980
Philadelphus
789
RosaceaeRose family
790
Common name
Occurrence
dwarf ninebark
California to Nevada
Amur ninebark
1856
Pacific ninebark
mallow ninebark
First cultivated
1827
1897
mountain ninebark
1889
common ninebark
Maine to Minnesota, S
to Tennessee & Florida
1687
Atlantic ninebark
1908
Missouri
Physocarpus
791
References
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Everett TH. 1981. The New York Botanical Garden illustrated encyclopedia
of horticulture. New York: Garland Publishing. 3601 p.
Gill JD, Pogge FL. 1974. Physocarpus, ninebark. In: Schopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 584586.
Higginbotham JS. 1990. New plants for 1990. American Nurseryman
171(4): 2640.
Lea SM, Morgan P. 1993. Resprouting response of ninebark (Physocarpus
malvaceus) shrubs to burning and clipping. Forest Ecology and
Management 56: 199210.
Link E, ed. 1993. Native plant propagation techniques for National Parks,
interim guide. East Lansing, MI: USDA Soil Conservation Service, Rose
Lake Plant Materials Center. 240 p.
Krssmann G. 1985. Manual of cultivated broad-leaved trees and shrubs.
Vol. II. Portland, OR:Timber Press.
Strausbaugh PD, Core EL. 1978. Flora of West Virginia, 2nd ed. Grantsville,
WV: Seneca Books. 1079 p.
Stokes DW. 1981. The natural history of wild shrubs and vines. New York:
Harper & Row. 246 p.
Straley GB. 1989. A new cultivar of Physocarpus capitatus (Rosaceae).
Baileya 23(1): 5153.
USDA NRCS USDA Natural Resources Conservation Service]. 2001.
PLANTS,Version 3.1 [database available at http://plants.usda.gov]. Baton
Rouge, LA: National Plant Data Center.
Wheller AG, II, Hoebeke ER. 1985. The insect fauna of ninebark,
Physocarpus opulifolius (Rosaceae). Proceedings of the Entomological
Society of Washington 87(2): 356370.
Yoder WG. 1995. Personal communication. Licking, MO: George O. White
State Forest Nursery.
PinaceaePine family
Picea A. Dietr.
spruce
Andrew Youngblood and Lawrence O. Safford
Dr.Youngblood is a research forester at the USDA Forest Services Pacific Northwest Research Station,
La Grande, Oregon; Dr. Safford retired from the USDA Forest Services
Northeastern Forest Experiment Station
Growth habit, occurrence, and use. The spruce
genusPiceaincludes 40 to 50 species of evergreen
conifers native to the temperate and boreal regions of the
Northern Hemisphere, occurring in Europe, Asia Minor, the
Caucasus, Siberia, China, Japan, the Himalayas, and North
America (table 1). The genus evolved from primordial
ancestors in the northeastern mainland of Asia, with presentday Korean spruce likely the most primitive species. Most
North American species probably arose through eastward
migrated and mutation of Ezo spruce (Wright 1955). More
recent work suggests a strong relation between the Old
World Serbian spruce and the New World black spruce
(Fowler 1980). At least 12 species occur in China (Li and
others 1990). Seven species are native in North America,
excluding the rare and localized occurrence of Chihuahua
spruceP. chihuahuana Martinezin northwest Mexico
(Rushforth 1986; Patterson 1988). The genus name is
derived from the Latin pix or picis, pitch, referring to the
resinous qualities of the trees (Everett 1981) or of a pitch
pine, probably Scots pine (Pinus sylvestris L.) (Little 1979).
The genus includes medium to tall conifers that range in
height at maturity from 9 to over 70 m. Crowns of most
species appear conical in outline. The generally small
branches occur in whorls with common internodal branches.
The needle-like leaves are borne on peg-like projections
(pulvini) on the twigs, have angled or flattened cross section, and persist for several years. Needles fall readily from
twigs on drying. The slender boles gradually taper along
their entire length, sometimes from a buttressed base. The
thin and scaly bark sometimes has furrows at the base of old
trees. The generally shallow root systems have many long,
stringy, and tough rootlets. Open-grown trees retain live
branches to the ground, and in black spruce and sometimes
Norway, Ezo, and white spruces, layering occurs when
branch tips come in contact with moist soil, take root, and
develop into full-size trees (Nienstaedt and Zasada 1990;
Nikolov and Helmisaari 1992; Stone and McKittrick 1976;
Viereck and Johnston 1990).
Picea
793
Table 1Picea, spruce: nomenclature and occurrence of native and cultivated species in North America
Scientific name & synonym(s)
Common name(s)
Occurrence
Norway spruce
P. asperata Mast.
P. crassifolia Kamarov
P. breweriana S.Wats.
P. engelmannii Parry ex Engelm.
P. columbiana Lemmon
P. glauca ssp. engelmann (Parry
ex Engelmann) T.M.C.Taylor
P. engelmanii var. glabrata Goodman
P. glauca (Moench) Voss
P. alba (Aiton) Link
P. alba var. albertiana (S. Brown) Beiss.
P. albertiana S. Brown;
P. canadensis B.S.P.
P. canadensis var. albertiana (S. Brown) Rehder
P. glauca var. albertiana (S. Brown) Sarg.
P. glauca var. posildii Raup
P. nigra var. glauca Carr.
P. glehnii (Fr. Schmidt) Mast.
dragon spruce,
Chinese spruce
Brewer spruce, weeping spruce
Engelmann spruce,
mountain spruce
P. obovata Ledeb.
P. abies var. obovata Lindquist
P. omorika (Pancic) Purk.
P. pungens Engelm.
P. commutata Horton P. parryana Sarg.
P. rubens Sarg.
P. australis Small
P. nigra (Ait.) Link var. rubra (Du Roi) Engelm.
P. rubra (DuRoi) Link (not A. Dietrich)
P. sitchensis (Bong.) Carr.
P. sitchensis Bong.
P. falcata (Rafin.) Suringar
P. menziesii (D. Don) Carr.
Abies falcata Rafin.
A. menziesii (D. Don) Lindley
Pinus menziesii Douglas
P. smithiana (Wall.) Boss.
P. morinda Link
P. ramaleyi A. Nels.
Sakhalin spruce
Serbian spruce
blue spruce, Colorado spruce,
Colorado blue spruce
red spruce, West Virginia
spruce, eastern spruce,
yellow spruce, he-balsam
Himalayan spruce,
west Himalayan spruce
794
are planted as ornamentals. Many cultivars featuring variations or extremes in crown height, shape and symmetry, or
thickness; rate of height growth; branch angle and degree of
twig droop; and needle color exist (Everett 1981, Huxley
1992). In general, spruce species do not tolerate droughty
sites but do thrive on slightly acidic and moist but welldrained soils. Of all the species, Serbian spruce may best
tolerate industrial air pollution (Dallimore and Jackson
1967).
Geographic races and superior strains. The wide
ranges and diverse environments to which the spruce species
have adapted provide an array of individual, ecological, and
geographic variations. Natural hybridization and introgression commonly occur where ranges of compatible species
overlap. Hybridization between white spruce and Sitka
spruce (first reported by Little 1953 as P. lutzii), occurs in
British Columbia and throughout the Kenai Peninsula in
Alaska (Copes and Beckwith 1977). This hybrid has demonstrated a genetically based resistance to attack by the Sitka
spruce weevilPissodes strobi Peckwhich causes severe
height growth and stem form reduction in Sitka spruce
(Mitchell and others 1990). Hybridization between white
spruce and Engelmann spruce occurs in northern Montana
and British Columbia (Daubenmire 1974). Artificial crosses
of Engelmann spruce with Sitka spruce and with blue spruce
suggest the close relatedness of these North American
species (Fowler and Roche 1977). Electrophoresis has yet to
clearly identify hybrids of Engelmann spruce and blue
spruce along a 1,200-m elevational transect in the front
range of the Colorado Rocky Mountains, where the species
grow together. Morphological similarity between the
2 species, such as number of bud scales, number of stomatal
rows, and location of resin sacs, however, suggests either
convergent evolution or the influence of environmental variation on the morphological characters (Mitton and Andalora
1981). Natural introgression between the maritime Sitka
spruce and the more interior complex of white and
Engelmann spruces occurs in a portion of British Columbia,
and the hybrid fraction was estimated by restriction fragment length polymorphisms of the nuclear ribosomal RNA
genes (Sutton and others 1994). Differences in monoterpene
composition from black spruce oleoresin (including
pinene, 3-carene, and terpinolene) vary among geographic origins in an eastwest pattern, except for seeds
from sources in New England that have close affinity in
monoterpene composition to red spruce (Chang and
Hanover 1991). Natural introgression of black spruce into
red spruce may result in greater height and diameter growth
in New Brunswick, yet the hybrid performed unpredictably
in managed stands (Fowler and others 1988).
Early crosses within the genus have provided a thorough
background in potential crossability of the genus, including
specimens from artificial crosses between nonsympatric
species (Wright 1955). Analysis of morphological characteristics and monoterpene composition from artificial crosses
795
796
reproductive cycle (from Owens and Molder 1984, used with permission of
Brown
Cinnamon brown
Pale brown
Brown
Brown
Brown
Sources: Alden (1955), Alexander (1987), Alexander and Shepperd (1984), Edwards (1980), Fechner (1985), Nikolov and Helmisaari (1992), Safford (1974), Zasada and Viereck (1970).
* Cones of P. mariana are semi-serotinous and release seeds throughout the year for several years.
Green
Green
Green
Crimson
Green
Green
Bluish black
Green
Green
Yellow-green
Bright green
413
2
26
213
24
4
1213
13
38
34
SeptApr
SeptOct
SeptOct
AugMay
Oct*
OctMar
AugSept
OctNov
1018
813
36
35
6
58
24
5
710
1018
SeptNov
SeptOct
AugSept
Aug
Sept
Sept
SeptOct
AugSept
OctNov
AprJune
MayJune
May
MayJune
May
AprJune
AprMay
May
AprMay
4060
2030
1540
30
2025
10
20
3050
20
20
3060
45
2530
2530
1530
30
3045
18
927
30
2150
2130
1873
61
Cone ripening
Flowering
Species
P. abies
P. asperata
P. breweriana
P. engelmannii
P. glauca
P. glehnii
P. jezoensis
P. koraiensis
P. mariana
P. obovata
P. omorika
P. pungens
P. rubens
P. sitchensis
P. smithiana
Ripe
cone color
Preripe
cone color
Dispersal
Years
between large
seedcrops
Cone
size (cm)
Minimal seedbearing age (yr)
Mature
height (m)
Table 2Picea, spruce: height, seed-bearing age, and phenology of flowering and fruiting
bright purple, or crimson male strobili have ovoid to cylindrical shape and uniform distribution over the crown. Each
scale (microsporophyll) bears 2 pollen sacs (microsporangia) and are spirally arranged on a central axis. Male strobili
dry out and fall off soon after pollen-shedding.
The timing of female strobili differentiation is similar
for most species of spruce that have been studied; female
strobili become anatomically determined at the end of the
period of bud-scale initiation and the end of lateral shoot
elongation (Owens 1986). Female strobili arise near the
apex of shoots on upper branches in crowns of Engelmann,
Sitka, and white spruces; the seed-cone zone in black spruce
occurs on the most vigorous 1-, 2-, and 3-year-old branches
at the top of the tree (Caron and Powell 1992). Initially the
female strobili are erect, yellowish green, crimson, or purple; cylindrical; and 5 to 20 mm in diameter. The ovuliferous scales are spirally arranged on a central axis and each
bears 2 ovules (megasporangia) at the base. Each species
has a characteristic number of spirals per cone, and the number of seeds per cone depends in part on the pitch and diameter of the spirals as well as the length of the cones (Fogal
and Alemdag 1989). The size of the preceding cone crop
and climatic conditions at the time of cone bud differentiation influence the number of reproductive buds formed in
white spruce (Zasada and others 1978). Checking female
strobili in the fall preceding the seed year provides an early
means of predicting potential size of the cone crop
(Eis 1973).
Female strobili receive pollen when fully open, a period
that lasts only a few days. Fechner (1974) determined that
female strobili of blue spruce become receptive 1 to 5 days
after the first pollen release, depending on elevation, and
that cones tip over and become pendent within 3 to 4 weeks
of initial receptivity. Fertilization may follow pollination
within a few days or may be delayed until after cones
become pendent (Fechner 1974); cones mature in late summer or autumn, depending on summer growing conditions
(table 2). Embryo development (figure 2) of white spruce
seeds in Alaska generally proceeds rapidly during July after
completion of shoot, stem, and cone growth, although on
any specific date, embryo length, percentage of embryo
length, cotyledon length, and relative cotyledon length will
differ among trees within a stand (Zasada 1988). Embryos
of white spruceEnglemann spruce hybrids in British
Columbia typically fill the embryo cavity well before the
seeds mature (Eremko and others 1989). Cotyledon number
between species differs from 4 to 15 (Dallimore and Jackson
1967) and may be under strong maternal control (Diebel and
Fechner 1988).
The size of a cone crop for individual trees and stands
tends to follow the phenomenon of alternate bearing, with
heavy crops followed by light or no crops, because cones
develop in terminal positions on the shoots, leaving fewer
available locations for flower production the year after a
Picea
797
good crop (Edwards 1986; Fechner 1985). Annual production of cones and seeds differs considerably, however, with
the intervals between good cone crops ranging from 2 years
in Brewer spruce to as long as 13 years in white and
Norway spruces (table 2). Between 1969 and 1994,
Engelmann spruce in central Colorado produced good cone
crops in 8 of the 26 years (Shepperd 1995). Between 1957
and 1978, irregular production of white spruce cones and
seeds in interior Alaska varied with environmental factors
such as temperature during cone initiation; nutrient deficiencies; and losses to insects, diseases, and squirrels (Zasada
1980; Zasada and Viereck 1970).
At maturity, the pendent cones open to shed seeds during autumn and winter (table 2). Persistent cone scales on
mature cones may have rounded, pointed, irregular, notched,
or reflexed ends. Most species shed cones at the end of the
season, but some cones may remain on the tree throughout
the next growing season. Cones should be harvested before
inclement weather reduces workers productivity or losses to
squirrels increase (Curran and others 1987). Cones may be
collected before they are fully ripe and, if artificially
ripened, release seeds with maximal germination capacity
(Edwards 1980). Cones may be collected from standing
trees, slash, or animal caches, although cones that have been
in contact with the forest floor may acquire seed-killing
fungi. Seeds generally reach maturity before cones show
their characteristic ripe color (table 2). Time of ripening
varies among cones on an individual tree and among trees
in a single stand (Fechner 1974; Jensen and others 1967;
Zasada 1988). Various measures of estimating seed maturity
have emphasized (1) physical attributes such as color and
firmness of cones; (2) moisture content and specific gravity
of cones; (3) color of testa and brittleness of seeds (Crossley
1953; Edwards 1980); (4) a cone moisture content of 30%
or less for Norway spruce; (5) specific gravity between 0.78
798
Figure 3Picea, spruce: seeds with wings of P. breweriana, Brewer spruce; P. engelmannii, Engelmann spruce;
P. glauca, white spruce; P. mariana, black spruce; P. rubens, red
spruce; and P. sitchensis, Sitka spruce (left to right).
Picea
799
800
P. engelmannii
P. glauca
P. engelmannii, P. glauca, P. sitchensis,P. mariana
P. mariana, P. glauca, P. engelmannii, P. pungens,
P. sitchensis, P. glauca
P. mariana, P. glauca, P. engelmannii, P. sitchensis
P. engelmannii, P. glauca, P. mariana, P. pungens, P. rubens, P. sitchensis
P. glauca
All native Picea
P. mariana
Damage
Common name
Insect species
Affected species*
801
Sources: Dirr and Heuser (1987), Jeglum and Kennington (1993), Nikolov and Helmisaari (1992), Safford (1974), Stein and others (1986),Willan (1985).
Note: TB = top of blotters; P = petri dishes covered with blotters, filter paper, or sand.
Prechill
Prechill; light; sensitive to excess moisture; if dormant, use KNO3
Prechill 1421 days at 35 C; light
Prechill 21 days at 35 C
Prechill 21 days at 35 C
Light
Prechill or soak; light
Light; sensitive to excess moisture
Prechill
Light
Soak; prechill; light; sensitive to excess moisture; if dormant, use KNO3
16
16
21
14
14
21
16
16
28
21
TB
TB,P
TB
TB,P
TB,P
TB
TB
TB
TB,P
TB
TB,P
47,000209,700
70,00075,000
51,00074,000
69,000322,000
135,000401,000
179,200230,600
95,000110,000
335,000664,000
125,600171,200
80,000163,000
100,000289,000
155,000400,000
24,00040,000
105,600462,300
154,300165,400
112,500163,200
152,200710,000
298,000884,200
395,100508,500
209,500242,500
739,0001,464,100
277,000377,500
176,400359,000
220,500637,000
342,000882,000
53,00088,200
P. abies
P. asperata
P. breweriana
P. engelmannii
P. glauca
P. glehnii
P. jezoensis
P. koraiensis
P. mariana
P. omorika
P. pungens
P. rubens
P. sitchensis
P. smithiana
Additional directions
Test
(days)
Substrate
/lb
Seeds/weight
/kg
Species
Table 4Picea, spruce: weight of cleaned seeds, methods of testing for laboratory germination, and additional directions
for 14 months showed only a slight loss of percentage germination; under these conditions black spruce lost about
one-third of its percentage germination and germination of
all 3 species declined to about 10% of the original capacity
after 27 months (MacGillivary 1955). Prechilling, or cold
stratification, may widen the range of temperatures over
which seeds can subsequently germinate, increase the maximal percentage germination at some temperatures, and
increase the rate of germination at almost any temperature
(Gosling and Rigg 1990). Prechilling of white spruce seeds
at 2 to 4 C for 6 weeks results in high percentage germination (Caron and others 1990). Other researchers have
prechilled white spruce seeds by soaking them in cold running water for 24 hours, blotting them dry, and then refrigerating them at 4 C for 3 weeks (Chanway and others 1991).
Storage of cones at 5 C for 4 weeks, however, may eliminate any subsequent need for stratification of white spruce
seeds (Winston and Haddon 1981). White spruce seeds from
high-latitude sites in Alaska (> 55 latitude) do not undergo
dormancy, and stratification is detrimental for mature seeds
(Alden 1995). Before nursery sowing, Engelmann spruce
seeds need to be primed for 24 hours, then prechilled for 6
to 8 weeks at 2 C in loosely closed polyethylene bags
(Tanaka and others 1986). Unstratified seeds of black spruce
incubated for 24 days at 3 or 20 C germinated completely
within18 days with 14:10 (light:dark) hours of fluorescent
light, whereas moist seeds prechilled for 24 hours at 3 C in
a polyethylene bag in the dark reached 95% germination
within 12 days when incubated at 5 to 30 C, regardless of
lighting regime (Farmer and others 1984). Priming black
spruce seeds for 5 to 6 days in water (until the radicles nearly emerge) and surface-drying before sowing accelerated
germination by about 1 week (Malek 1992). Seeds of black
spruce from high latitudes in Alaska, collected and immediately extracted in the spring, will germinate in 2 to 6 days
after becoming fully imbibed with water; no dormancy
exists and stratification is not required (Alden 1995).
Dormancy of Sitka spruce seeds is broken and 95% germination is possible after priming for 72 hours (until seeds
reach 30% moisture content), then chilling for 6 weeks in
loosely closed polyethylene bags at 4 C (Gosling and Rigg
1990).
Treating seedlots with various fumigants, insecticides,
fungicides, and rodent repellents in storage or before sowing
may reduce germination of seeds. Germination of white
spruce seeds treated with a finely ground rodent repellent
mixed with graphite declined slightly from that of untreated
seeds after more than 5 years of storage (Radvanyi 1980).
Aluminum powder, which is used as a lubricant on Sitka and
white spruce seeds in bareroot nurseries in Canada, may
decrease the percentage germination of treated seeds and
reduce first-year survival of seedlings (Sutherland and others
1978). Embedding black spruce seeds in pellets may discourage their consumption by small mammals, depending
Picea
803
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EricaceaeHeath family
807
seeds.
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808
PinaceaePine family
Pinus L.
pine
809
Common name(s)
Occurrence
P. albicaulis Engelm.
whitebark pine
P. aristata Engelm.
bristlecone pine,
P. balfouriana var. aristata (Engelm.) Engelm. foxtail pine, hickory pine
P. arizonica Engelm.
P. ponderosa var. arizonica (Engelm.) Shaw
P. armandii Franch.
P. attenuata Lemmon
P. tuberculata Gord.
foxtail pine,
Balfour pine
P. banksiana Lamb.
jack pine, scrub pine,
P. divaricata (Ait.) Dum.-Cours.
banksiana pine,
black/gray pine,
Hudson Bay pine
P. brutia Tenore
Calabrian pine
P. halepensis var. brutia (Ten.) Elwes & Henry
P. canariensis C. Smith
Canary Island pine,
Canary pine
P. caribaea Morelet
Caribbean pine
P. bahamensia Griseb.
P. hondurensis Loock
P. cembra L.
Swiss stone pine,
P. montana Lam.
cembrian pine,
arolla pine
P. cembroides Zucc.
Mexican piyon,
nut pine, pinyon
P. clausa (Chapman ex
Engelm.) Vasey ex Sarg.
Bolander pine
810
Common name(s)
P
Occurrence
P. elliottii var. densa Little & Dorman South Florida slash pine
P. elliottii var. elliottii Engelm.
P. caribaeu Morelet
P. engelmannii Carr.
P. latifolia Sarg.
P. apacheca Lemmon
P. flexilis James
P. gerardiana D. Don
P. aucklandii Lodd.
P. chilghoza Ehh.
P. glabra Walt.
P. halepensis P. Mill.
P. alepensis Poir.
P. heldreichii Christ
P. heldreichii var. leucodermis (Ant.) Markgr.
P. eucodermis Ant.
P. nigra var. leucodermis (Ant.) Rehd.
P. jeffreyi Grev. & Balf.
P. ponderosa var. jeffreyi (Grev. & Balf.)
E. Murr.
P. kesiya Royle & Gordon
P. khasya Royle
P. insularis Endl.
P. koraiensis Sieb. & Zucc.
Heldreich pine,
Balkan pine, Bosnian pine,
graybark pine
P. lambertiana Dougl.
sugar pine,
pio real
Chihuahua pine,
yellow pine,
pio real
Merkus pine,
Tenasserim pine
singleleaf piyon,
nut pine,
pinyon, pion
western white pine,
Idaho white pine,
silver pine
P. mugo Turra
P. montana Miller
Swiss mountain
pine, mugho
(or mugo) pine,
dwarf mountain pine
P. muricata D. Don
P. remorata Mason
bishop pine,
prickle-cone pine,
Santa Cruz Island pine
Austrian pine,
European black
pine, black pine
P. nigra Arnold
P. nigra var. austriaca (Hoess)
Aschers. & Graebn.
Jeffrey pine
Khasi pine,
Benguet pine
Korean pine,
cedar pine
Pinus
811
Common name(s)
Occurrence
P. palustris P. Mill.
P. australis Michx. f.
longleaf pine,
southern pine,
longstraw pine
Japanese white pine
P. roxburghii Sarg.
P. longifolia Roxb.
Chir pine,
longleaf Indian pine
P. serotina Michx.
P. rigida var. serotina (Michx.) Clausen
P. sibirica Du Tour
P. cembra var. sibirica Loud.
P. strobiformis Engelm.
P. flexilis var. reflexa Engelm.
P. reflexa (Engelm.) Engelm.
P. ayacahuite var. brachyptera Shaw
southwestern white
pine, border limber
pine, Mexican white pine
812
Table 2 Pinus, pine: mature tree height, earliest cultivation, seed bearing age, and interval between large cone crops
Species
P. albicaulis
P. aristata
P. arizonica
P. armandii
P. attenuata
P. balfouriana
P. banksiana
P. brutia
P. canariensis
P. caribaea
P. cembra
P. cembroides
P. clausa
P. contorta
var. contorta
var. latifolia
var. murrayana
P. coulteri
P. densiflora
P. echinata
P. edulis
P. elliottii
var. densa
var. elliottii
P. engelmannii
P. flexilis
P. gerardiana
P. glabra
P. halepensis
P. heldreichii
P. jeffreyi
P. kesiya
P. koraiensis
P. lambertiana
P. leiophylla var. chihuahuana
P. merkusii
P. monophylla
P. monticola
P. mugo
P. muricata
P. nigra
P. palustris
P. patula
P. parviflora
P. peuce
P. pinaster
P. pinea
P. ponderosa
var. ponderosa
var. scopulorum
P. pumila
P. pungens
P. quadrifolia
P. radiata
Mature tree
height (m)
Year first
cultivated
633
615
2327
1837
515
1118
1730
2030
30
1830
1023
58
524
1852
1861
1895
1847
1852
1783, earlier
Age at onset
seed bearing (yr)
Years between
cone crops
1746
1830
1832
2030
20
1520
20
58
20
315
710
1520
2530
35
102
23
1
56
34
1
34
610
58
12
612
846
1530
923
2137
230
312
1855
1853
1832
1854
1726
1848
48
510
48
820
2030
520
2575
1
1
1
36
2
310
25
826
2430
1521
624
1524
2427
1524
1830
1855
3046
2746
3069
924
1830
615
2761
212
1227
2050
2437
1834
530
1030
2737
1423
1861
1839
1683
1865
1853
1861
1827
1848
1851
1779
1846
1759
1727
1861
1863
1660, earlier
Long history
812
710
2830
2040
10
1520
8
510
1540
4080
2830
1020
2025
720
10
56
1540
20
1215
1230
1015
15
3
34
24
24
1
15
35
34
12
12
37
1
23
25
57
45
34
35
1870
1535
0.32.5
918
59
246
1826
1807
1804
1885
1833
1620
620
510
25
25
15
1
Table 2 Pinus, pine: mature tree height, earliest cultivation, seed bearing age, and interval between large cone crops (continued)
Species
P. resinosa
P. rigida
P. roxburghii
P. sabiniana
P. serotina
P. sibirica
P. strobiformis
P. strobus
P. sylvestris
P. taeda
P. thunbergiana
P. torreyana
P. virginiana
P. wallichiana
P. washoensis
Mature tree
height (m)
2146
630
4655
1224
1224
40
838
2467
2440
2734
3040
818
1530
1546
1846
Year first
cultivated
1756
1759, earlier
1807
1832
1713
1837
1840
1705
Long history
1713
1855
1853
1739
1827
Age at onset
seed bearing
2025
34
1540
1025
410
2535
15
510
515
510
640
1218
5
1520
1520
Years between
cone crops
37
49
24
24
1
38
34
310
46
313
1
1
12
25
Sources: Altman and Dittmer (1962), Bailey (1970), Bates (1930), Britton and Shafer (1908), Carlisle and Brown (1968), Cooling (1968), Dallimore and Jackson (1967),
Day (1967), den Ouden and Boom (1965), Dimitroff (1926), Duff (1928), Fowells (1965), Fritts (1969), Goor (1955), Harlow and Harrar (1950), Iroshnikov (1963), Iroshnikov
and others (1963), Jacaline and Lizardo (1958), Krugman and Jenkinson (1974), Little EL (1941a, 1950), Loock (1950), Luckhoff (1964), Magini and Tulstrip (1955), McIntyre
(1929), Mirov (1956), NBV (1946), Otter (1933), Pearson (1931), Poynton (1961), Pravdin (1963), Rehder (1940), Rohmeder and Loebel (1940), Rossi (1929), Sargent (1905,
1965), Sudworth (1908),Troup (1921),Veracion (1966),Wahlenberg (1946),Wakeley (1954),Wakeley and Barnett (1968),Wappes (1932),Wellner (1962).
Note: See table 3 for cone ripening and seed dispersal dates and table 4 for cone ripeness criteria.
tests follows a largely clinal pattern that is linked to environmental gradients of latitude and length and temperature of
the growing season at seed origin (Rudolph and Laidly
1990; Rudolph and Yeatman 1982).
Pinus brutiaCalabrian pine has 2 known varieties,
both in the northernmost parts of its range. The var. pithyusa
Stev. occurs along the north central and northeast shores of
the Black Sea and var. eldarica Medw. occurs in the central
Caucasus Mountains (Magini and Tulstrup 1955). Trees
from an Afghanistan source related to var. eldarica outgrew
trees of var. pithyusa, had good form, and were both frost
and drought hardy in California (Harris and others 1970;
Krugman and Jenkinson 1974). Altitudinal variation in a
number of seed and seedling traits is manifested in Greece
and in Turkey (Isik 1986; Panetsos 1986).
Pinus canariensisCanary Island pine is native only to
the Canary Islands, where it is found at 640 to 2,195 m
above sea level (Magini and Tulstrup 1955). Seedlings
grown from seeds from the various islands and an array of
elevations showed marked differences in winter cold hardiness when grown at the USDA Forest Services Institute of
Forest Genetics nursery near Placerville, California.
Seedlings from a seed source at 1,220 m on Tenerife Island
showed more cold damage than those from a source at 1,890
m there. Seedlings from a source from 1,890 m on Palma
Island, however, were badly damaged, suggesting that
Pinus
815
818
Pinus
819
820
Pinus
821
that of the planting site generally yield the best survival and
growth rates. Southern seed sources produce fast-growing
trees on southern sites but on northern sites these trees grow
slowly and suffer winter injury (Dorman 1976; Genys 1966;
Genys and others 1974). Genetic variation in growth rate,
stem form, wood traits, and monoterpene content in
Kentucky and Tennessee occurs mainly among and within
stands, rather than among geographic origins (Carter and
Snow 1990).
Pinus wallichianaBlue pine ranges through the
Himalayan region in discontinuous distribution from eastern
Afghanistan to eastern-most India, north Burma, and adjoining China. Although no distinct varieties have been reported,
at least 7 broad provenances have been proposed, including
4 in the western Himalayas and 3 in the eastern Himalayas
(Dogra 1987).
Pinus washoensisWashoe pine ranges in limited areas
along the western edge of the Great Basin in western
Nevada and northern California, notably on the east slope of
Mount Rose in the Carson Range in Nevada, in the Bald
Mountain Range in the northern Sierra Nevada, and in the
South Warner Mountains and several intervening areas in
northeastern California (Critchfield 1984; Critchfield and
Allenbaugh 1965; Niebling and Conkle 1990; Smith 1981).
Closely related in appearance to and often wrongly identified as Pacific ponderosa pine, this rare pine ranges at higher
altitudes than ponderosa pine, the same as Jeffrey pine.
Washoe pine flowers in July, and its male and developing
second-year female cones are purple to purplish black. The
latter mature in AugustSeptember, turn to a dull purple,
purplish brown, or light brown, and open in September.
Cones are assessed and processed like those of Pacific ponderosa pine. Stored seeds germinate quickly after 60 days of
moist, cold stratification (Jenkinson 1980; Krugman and
Jenkinson 1974).
Hybrids. Pine hybrids are myriad. More than 260
first- and second-generation hybridsas well as backcrosses, crosses between varieties of the same species, and crosses that involve 3 or more different specieseither occur naturally or have been produced artificially (Critchfield 1963,
1966a, 1977, 1984; Critchfield and Krugman 1967;
Krugman and Jenkinson 1974; Little and Righter 1965;
Mirov 1967; Vidacovic 1991; Wright 1962). We provide no
yield statistics for hybrids because such data are highly variable. Yields of sound seeds depend on species and individual
trees, as well as on the environmental conditions under
which the cross is made.
Natural hybrids are common, and we list but few of the
many known. P. palustris taeda (Sonderegger pine) occurs
822
frequently in Louisiana and east Texas and is the bestknown southern pine hybrid (Baker and Langdon 1990;
Chapman 1922). Most natural hybrids occur infrequently in
the overlaps of their parent species ranges. Some of the
better-known examples include the following:
P. contorta var. murrayana banksiana in western
Canada (Zavarin and others 1969)
P. ponderosa jeffreyi, P. jeffreyi coulteri, and P.
radiata attenuata in California (Critchfield and
Krugman 1967; Mirov 1967)
P. flexilis strobiformis in north central Arizona and
north central New Mexico (Steinhoff and Andresen
1971)
P. taeda echinata in east Texas
P. taeda serotina throughout the species common
range in southern United States (Critchfield 1963;
Smouse 1970; Wright 1962)
The hybrid P. densiflora thunbergiana, natural in
Japan, has formed spontaneously in plantations of its parent
species in Michigan (Krugman and Jenkinson 1974). In
Europe, Scots pine crosses occasionally with Austrian pine
and with mugo pine where the species are planted near one
another (Wright 1962), and Austrian pine crosses with
Heldreich pine var. leucodermis where they overlap in
Herzegovina (Vidacovic 1991).
Several pine hybrids have been produced in relatively
large numbers by controlled pollination methods. They
include P. rigida taeda in Korea where the hybrid is
important in plantation forestry (Hyun 1962), and P. attenuata radiata, tested in California and Oregon (Griffin and
Conkle 1967). Many other pine hybrids have been produced
in small numbers and tested for fitness in various parts of
the United States (Burns and Honkala 1990).
Flowering and fruiting. Reproductive structures in
certain pines first form when the trees are 5 to 10 years old,
as in knobcone, jack, sand, lodgepole, and Monterey pines,
among others (table 2). In other pines, reproductive structures form when the trees are 25 to 30 years old (as in Swiss
stone and Siberian stone pines; piyon; and Apache and
Chihuahua pines) or 40 years old (as in sugar pine).
Pines are monoecious. Male and female flowers
properly strobili (microsporangiate and megasporangiate
strobili)are borne separately on the same tree. Male strobili predominate on the basal part of new shoots, mostly
those on older lateral branches in the lower crown. Female
strobili occur most often in the upper crown, chiefly at the
apical ends of main branches in the position of subterminal
Species
Location
Flowering
Cone ripening
Seed dispersal
P. albicaulis
P. aristata
P. arizonica
P. armandii
P. attenuata
P. balfouriana
P. banksiana
P. brutia
P. canariensis
P. caribaea
P. cembra
P. cembroides
P. clausa
P. contorta
var. contorta
var. latifolia
var. murrayana
P. coulteri
P. densiflora
P. echinata
P. edulis
P. elliottii
var. densa
var. elliottii
P. engelmannii
P. flexilis
P. gerardiana
P. glabra
P. halepensis
P. heldreichii
P. jeffreyi
P. kesiya
P. koraiensis
P. lambertiana
P. leiophylla var.
chihuahuana
P. merkusii
P. monophylla
P. monticola
P. mugo
P. muricata
P. nigra
P. palustris
P. parviflora
P. patula
P. peuce
P. pinaster
P. pinea
P. ponderosa
var. ponderosa
var. scopulorum
P. pumila
P. pungens
P. quadrifolia
P. radiata
California
Arizona
Arizona
California
California
California
Great Lakes
California
California
Cuba
Germany
California
Florida
July
JulyAug
May
AprMay
Apr
JulyAug
MayJune
MarMay
AprMay
JanFeb
May
MayJune
SeptDec
AugSept
SeptOct
SeptOct
Aug
JanFeb
SeptOct
Sept
JanMar
Sept
JulyAug
AugOct
NovDec
Sept
Not shed
SeptOct
Oct
AugSept
Closed cone
SeptOct
Sept
Closed cone
SeptOct
Sept
Not shed
NovDec
Sept
California
Rocky Mtns
California
California
California
South Carolina
Arizona
MayJune
JuneJuly
MayJune
MayJune
Apr
MarApr
June
SeptOct
AugSept
SeptOct
AugSept
AugSept
OctNov
Sept
Fall
SeptOct
SeptOct
Oct
SeptOct
OctNov
SeptOct
Florida
Florida
Arizona
California, Montana
India, California
Mississippi
France
Italy, California
California
Philippines
Japan
California
JanApr
JanFeb
May
JuneJuly
MayJune
FebMar
MayJune
MayJuly
JuneJuly
JanMar
MayJune
JuneJuly
AugSept
SeptOct
NovDec
AugSept
SeptOct
Oct
Sept
AugSept
AugSept
OctJan
Sept
AugSept
SeptNov
Oct
NovFeb
SeptOct
Nov
OctNov
Fall
SeptOct
SeptOct
FebMar
Oct
AugOct
California
Thailand
California, Nevada
Idaho
Europe
California
Ontario, Canada
SE US
Japan
Mexico, California
Europe
Europe, California
Europe
MayJune
Jan
May
JuneJuly
MayJune
AprJune
MayJune
FebMar
MayJune
FebApr
May
Apr
MayJune
Nov*
AprJun
Aug
Aug
Oct
Sept
SeptNov
SeptOct
Sept
Dec
Fall
NovDec
Late summer*
DecJan
MayJul
SeptOct
AugSept
NovDec
Midwinter
OctNov
OctNov
Nov
Midwinter
Fall
DecJan
Late summer
California
South Dakota, Colorado
Russia
West Virginia, California
California
California
AprJune
MayJune
July
MarApr
June
JanFeb
AugSept
AugSept
Fall
Sept
Nov
AugSept
SeptJan
Fall or not shed
Fall
SeptOct
Midwinter
Pinus
823
Location
Flowering
Cone ripening
Seed dispersal
P. resinosa
P. rigida
P. roxburghii
P. sabiniana
P. serotina
P. sibirica
P. strobiformis
P. strobus
P. sylvestris
P. taeda
P. thunbergiana
P. torreyana
P. virginiana
P. wallichiana
P. washoensis
AprJune
May
FebApr
MarApr
MarApr
May
June
MayJune
MayJune
FebApr
AprMay
FebMar
MarMay
AprJune
July
AugOct
Sept
Winter
Oct
Sept
AugSept
Sep
AugSept
SeptOct
SeptOct
OctNov
JunJul*
SeptNov
AugOct
AugSept
OctNov
Fall
AprMay
Oct II
Spring
Not shed
SeptOct
AugSept
DecMar
OctDec
NovDec
Sept II
OctNov
SeptNov
Sept
Sources: Andreev (1925), Britton and Shafer (1908), Carlisle and Brown (1968), Cocozza (1961), Cooling (1968), Critchfield (1966b), Dallimore and Jackson (1967),
Dimitroff (1926), Dorman and Barber (1956), Duffield (1953), Fowells (1965), Goor (1955), Jacaline and Lizardo (1958), Krugman and Jenkinson (1974), Lamb (1915),
Letourneux (1957), Little EL (1938a & b, 1940), Little S (1941, 1959); Loiseau (1945), Loock (1950), Luckhoff (1964), Mikhalevskaya (1960), Mirov (1962), NBV (1946),
Ohmasa (1956), Pearson (1931), Rehder (1940), Rohmeder and Loebel (1940), Sargent (1905), Snow (1960), Sudworth (1908),Tkachenko (1939),Troup (1921),Veracion
(1964),Vines (1960),Wahlenberg (1946).
* Cones and seeds mature in the third year.
Seeds are dispersed when the detached cones disintegrate.
Cones open after several years, if at all. Seed dispersal normally requires fire.
Many cones remain closed for several months or years.
II Cones are massive, open slowly, and shed seeds over several months.
or lateral buds. Exceptions to this general scheme are common. Trees of knobcone, jack, sand, and Monterey pines are
multinodal in the bud, and female strobili occasionally arise
in secondary whorl positions. Trees of knobcone, Monterey,
and Virginia pines usually produce female strobili in all
parts of the crown. In temperate climates, the earliest stages
of male and female strobili can be detected in the developing buds in summer or fall. Male strobili appear from 1 to
several weeks before the female strobili (Fowells 1965;
Gifford and Mirov 1960; Krugman and Jenkinson 1974;
Mirov 1967).
Male and female strobili of the southern and tropical
pines emerge from buds in late winter, as in slash pine var.
densa and var. elliottii) and spruce and longleaf pines (table
3). Strobili of other pines emerge from the winter bud in
early spring or in late spring and early summer.
Male strobili are arrayed in indistinct spirals in clusters
that range from 13 to 51 mm in length (Dallimore and
Jackson 1967; Pearson 1931; Shaw 1914; Sudworth 1908).
Immature male strobili are green or yellow to reddish purple; mature male strobili at the time of pollen shed are light
brown to brown. In most pines, male strobili fall soon after
ripening.
Female strobili emerge from the winter bud shortly after
the male strobili and are green or red to purple (Dallimore
824
Species
Pre-ripe color
Ripe color
P. albicaulis
P. aristata
P. arizonica
P. armandii
P. attenuata
P. balfouriana
P. banksiana
P. brutia
P. canariensis
P. caribaea
P. cembra
P. cembroides
P. clausa
P. contorta
var. contorta
var. latifolia
var. murrayana
P. coulteri
P. densiflora
P. echinata
P. edulis
P. elliottii
var. densa
var. elliottii
P. engelmannii
P. flexilis
P. gerardiana
P. glabra
P. halepensis
P. heldreichii
P. jeffreyi
P. kesiya
P. koraiensis
P. lambertiana
P. leiophylla
var. chihuahuana
P. merkusii
P. monophylla
P. monticola
P. mugo
P. muricata
P. nigra
P. palustris
P. patula
P. parviflora
P. peuce
P. pinaster
P. pinea
P. ponderosa
var. ponderosa
var. scopulorum
P. pumila
P. pungens
P. quadrifolia
P. radiata
P. resinosa
P. rigida
P. roxburghii
P. sabiniana
Dark purple
Green to brown-purple
Green
Green
Greenish brown
Deep purple
Green
Green
Yellow-green
Green
Greenish violet
Green
Green
Purple-green
Purple-green
Purple-green
Green
Green
Green
Green
Green
Green
Brownish purple-green
Green
Green
Green
Green
Yellow-green
Dark purple to black
Green
Green
Green
Brown
Brown-yellow to brown
Light brown
Lustrous yellowish brown to light brown
Brown
Green
Lustrous yellowish brown or reddish brown
Yellowish or light brown to dull brown
Dull purple to light brown
Bright brown to dark brown
Yellowish brown
Lustrous greenish brown to light brown
Green
Green
Green
Green to purple-black
Violet purple
Green to purple
Yellowish green
Green
Green
Yellow-green
Green to yellow
Purplish
Green
Light brown
Light brown
Shiny deep russet brown
Yellowish beige-brown to reddish, dark brown
Lustrous tawny yellow to dark brown or cinnamon brown
Shiny light chestnut brown
Shiny yellow brown to light brown
Green to dull brown
Yellow ochre to nut brown
Leathery-woody brownish red to reddish brown
Tawny yellow to light brown
Lustrous light brown
Shiny nut brown
Pinus
825
Pre-ripe color
Ripe color
P. serotina
P. sibirica
P. strobiformis
P. strobus
P. sylvestris
P. taeda
P. thunbergiana
P. torreyana
P. virginiana
P. wallichiana
P. washoensis
Green to yellow
Green
Green
Green
Green
Green
Deep lustrous purple
Green to dark violet
Green
Green
Purple to purplish black
Sources: Bailey (1939), Barnett and McLemore (1967b), Britton and Shafer (1908), Cerepnin (1964), Cooling (1968), Dallimore and Jackson (1967), den Ouden and
Boom (1965), Fowells (1965), Jacaline and Lizardo (1958), Krugman and Jenkinson (1974), Little EL (1940, 1950), Lizardo (1950), Loock (1950), Luckhoff (1964), McIntyre
(1929), McLemore (1961a&b), Pravdin (1963), Rehder (1940), Sargent (1965), Sudworth (1908),Troup (1921),Wahlenberg (1946),Wakeley (1954).
Note: See table 2 for intervals between large cone crops and table 3 for cone ripening and seed dispersal dates.
Jenkinson 1974; Rudolph and others 1959). The closedcone habit results from 3 factors: an extremely strong adhesion between adjacent, overlapping cone scales beyond the
ends of the winged seeds (LeBarron and Roe 1945, Little
and Dorman 1952a); cone structure; and the nature of the
scale tissue systems described. The scales appear to be
bonded by a resinous substance. The melting point of this
resin seal is between 45 and 50 C for Rocky Mountain
lodgepole pine (Critchfield 1957). Heat, especially that of
fire, melts the resin so that the cones open. Still other pines,
such as whitebark, Japanese stone, Swiss stone, and
Siberian pines, shed partly opened or unopened cones, and
their seeds are dispersed only when the cones disintegrate
on the ground (Dallimore and Jackson 1967; Mirov 1967;
Pravdin 1963; Sudworth 1908).
Cones that open on the tree are usually shed within a
few months to a year after the seeds are shed. Pines such as
knobcone, Rocky Mountain lodgepole, and pitch pine, how-
2
3
4
5
A
1
2
3
4
5
6
Pinus
827
longitudinal
Specific gravity
of ripe cones
0.59.80
.88.97
.43.89
1.10
.88
.80.86
<.89
<.95
.88
.81.86
.70.80
1.00
.80.89
.80.89
.84.86
<.85.86
<1.00
.80.94
.88
.85.95
.90.97
.881.00
.88.90
<1.00
Flotation
liquid
Kerosene
SAE 20 oil
Kerosene
SAE 20 oil
SAE 20 oil
SAE 20 oil
SAE 30 oil
Kerosene
SAE 20 oil
Kerosene
SAE 30 oil
Kerosene
Water
Kerosene
95% ethanol
Linseed oil
SAE 20 oil
scale tips, and in eastern white pine, when they turn from
green to yellow-green with brown on the scale tips, or to
light brown. In certain other pines, however, cone color
changes too late to be a useful index to ripeness. In longleaf
pine, for example, ripe cones are still green in color and
may have already started to open before turning brown
(Wakeley 1954).
In species for which cone color changes may not be useful, seed maturity may be assessed by flotation tests of the
cones specific gravity (table 5). Although the crucial factor
in cone ripening is moisture content, not specific gravity,
measuring specific gravity is the quickest and easiest way to
estimate cone moisture content (Bonner 1991). To determine
whether cones have reached a desired specific gravity, sam-
Pinus
829
Table 6Pinus, pine: cone processing schedules and safe times to cold- or freeze-store dry, mature seeds
Cone processing schedule*
Species
P. albicaulis
P. aristata
P. arizonica
P. armandii
P. attenuata
P. balfouriana
P. banksiana
P. brutia
P. canariensis
P. caribaea
P. cembra
P. cembroides
P. clausa
P. contorta
var. contorta
var. latifolia II
var. murrayana
P. coulteri
P. densiflora
P. echinata
P. edulis
P. elliottii
var. densa
var. elliottii
P. engelmannii
P. flexilis
P. gerardiana
P. glabra
P. halepensis
P. heldreichii
P. kesiya
P. jeffreyi
P. lambertiana
P. merkusii
P. monophylla
P. monticola
P. mugo
P. muricata
P. nigra
P. palustris
P. parviflora
P. patula
P. pinaster
830
Boiling
water dip
(sec)
Air-drying
(day)
Kiln-drying
(hr)
0
0
0
0
1530
0
0
1030
0
0
0
0
1030
1530
28
15
13
28
310
320
210
28
1
0
0
60
0
48
0
24
0
0
10
0
0
24
43
49
66
54
63
8
9
16
16
1718
18
3
>1
0
0
3060
0
0
0120
0
030
0
0
220
230
220
315
34
0
96
0
68
0
0
72
0
48
0
49
60
49
41
17
20
17
5
25
35
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1530
0
0
42
1530
15
310
520
520
57
57
57
23
57
310
515
12
410
810
0
810
0
60
0
0
48
10
0
0
0
24
0
24
0
0
0
4
0
48
48
24
0
48
0
48
49
49
43
38
54
49
49
43
49
49
46
38
46
46
Kiln
temp
(C)
Seed
storage
(yr)
50
>1
10
18
21
20
>10
510
21
11
Table 6Pinus, pine: cone processing schedules and safe times to cold- or freeze-store dry, mature seeds (continued)
Cone processing schedule*
Species
P. pinea
P. ponderosa
var. ponderosa
var. scopulorum
P. pumila
P. pungens
P. quadrifolia
P. radiata
P. resinosa
P. rigida
P. roxburghii
P. sabiniana
P. serotina
P. sibirica
P. strobiformis
P. strobus
P. sylvestris
P. taeda
P. thunbergiana
P. torreyana
P. virginiana
P. washoensis
Boiling
water dip
(sec)
Air-drying
(day)
18
0
0
0
0
0
0
0
60120
60120
0
0
0
0
0
0
0
0
0
030
0
0
0
412
412
30
28
0
37
14
37
520
520
412
3
0
2
0
72
0
0
4872
0
1320
48
48
0
1520
1016
0
48
0
0
2
0
49
74
49
49
54
49
41
54
49
41
77
18
>15
21
30
11
>4
5
>2
10
15
>9
11
6
>5
8
Kiln-drying
(hr)
Kiln
temp
(C)
Seed
storage
(yr)
Sources: Barnett (1969, 1970), Barnett and McLemore (1967a&b), Bonner (1990), Church and Sucoff (1960), Dent (1947), FAO (1993), Goor and Barney (1968), Heit
(1967b), Jones (1962, 1966), Kamra (1967), Karschon (1961), Krugman and Jenkinson (1974), LeBarron and Roe (1945), Little and Dorman (1952a), Lizardo (1950), Luckhoff
(1964), McLemore (1961b), Mirov (1946), Nather (1958), NBV (1946), Nyman (1963), Ohmasa (1956), Schubert (1952), Schubert and Adams (1971), Simak and others
(1961), Steinhoff (1964), Stoeckeler and Jones (1957), Swingle (1939),Troup (1921),Wakeley (1954),Wakeley and Barnett (1968).
* Air drying temperatures are 15.6 to 32.2 C. Kiln drying, if used, should follow air drying. Recommended air drying time is 3 to 7 days, where none is listed.
Seed germination was at least 50% after storage. Seeds of most pines were stored at 0.5 to 5 C or 15.6 to 18.8 C. Freezing is preferred. Seed moisture contents
were between 5 and 10%.
Cones of P. albicaulis, P. cembra, P. pumila, and P. sibirica must be broken up to free and extract the seeds.
An alternate extraction method for P. cembroides, P. edulis, P. monophylla, and P. quadrifolia is to shake the trees mechanically and collect shed seeds from cloths spread on
the ground.
II Time needed in boiling water is estimated. Reported treatment was 5 to 10 minutes in water at 64 C or higher.
Cones were soaked in water overnight and dried in a warm room.
831
832
P. brutia
P. coulteri
P. echinata
P. echinata
P. elliottii var. elliottii
P. glabra
P. halepensis
P. nigra
P. palustris
P. pinaster
P. pinea
P. sabiniana
P. strobus
P. sylvestris
P. taeda
Water
Water
95% ethanol
Water
Water
95% ethanol
95% ethanol
95% ethanol
Pentane
95% ethanol
Water
Water
100% ethanol
Petroleum ether
Water
Species
P. albicaulis
P. aristata
P. arizonica
P. armandii
P. attenuata
P. balfouriana
P. banksiana
P. brutia
P. canariensis
P. caribaea
P. cembra
P. cembroides
P. clausa
P. contorta
var. contorta
var. latifolia
var. murrayana
P. coulteri
P. densiflora
P. echinata
P. edulis
P. elliottii
var. densa
var. elliottii
P. engelmanni
P. flexilis
P. gerardiana
P. glabra
P. halepensis
P. heldreichii
P. jeffreyi
P. keisya
P. koraiensis
P. lambertiana
P. leiophylla var.
chihuahuana
P. merkussi
P. monophylla
P. monticola
P. mugo
P. muricata
P. nigra
P. palustris
P. parviflora
P. patula
P. peuce
P. pinaster
P. pinea
Place
collected
Seed weight
/cone wt
/cone vol
g/kg
oz/lb
kg/hl
lb/bu
Lot
Idaho
Arizona
Arizona
France
California &
Oregon
California
Great Lakes
Europe
South Africa
Germany
Florida
40
923
0.6
.14.35
1.42
.901.3
1.1
.71.0
4.86.6
3942
2426
2.64.1
2.23.0
1719
1119
1.21.9
5.7
40
25
3.5
2.6
18
11
1.6
3
4
10
2
10
35
.15
.52
0.13
.26.9
.771.2
.1
.2.7
8.89.9
.6.9
3371
3149
156551
1726
4.04.5
5281
3.55.1
1.44.6
143187
1532
1422
71250
7.611.6
9.3
2437
1.62.3
0.62.1
6585
56
37
289
20.1
4.2
68.3
4.4
2.4
165
25.4
17
131
9.1
10
31
2.0
1.1
75
6
3
423
5
>10
>10
5
>10
California
Montana to
Colorado
California
California
Japan
Arizona
.641.5
.51.2
245364 111165
298
135
28
810
5.5
22
20
2030
28
.1.15
.1
.33
.3
.3.45
.4
.261.0
.26
1.03
.641.0
.521.4
3.3
.2.8
.2
.8
.5.8
.411.1
4.25
174251 79114
256262 116119
2.63.5
1.21.6
79141
3664
71161
3273
3.35.5
1.52.5
207
258
3.1
115
102
4.2
94
117
1.4
52
46
1.9
39
4
8
26
144
9
S Florida
Arizona
India
Louisiana
Italy
California
SE Asia
Korea
California
1020
11
35
37
.15.3
.16
.52
.56
.641.3
.771.0
.52
.131.3
1.22.6
1.92.6
.51.0
.6.8
.4
.11.3
.92.0
1.52.0
3137
2143
7.115.0
2.42.9
88115
4888
3571
5.811.7
4476
1.62.0
3.36.0
1417
1019
3.26.8
1.11.3
4052
2240
1632
2.65.3
2034
0.70.9
1.52.7
34
30
22
10.8
2.4
101
62
46
8.2
59.5
1.8
4.6
15
13
10
10.0
4.9
1.1
46
28
3.7
27
3.7
2.1
30
404
1
44
10
8
>10
18
26
>5
3
27
Nevada
Utah
Germany
California
Louisiana
South Africa
& Mexico
Europe
10
4
2040
21
.15
.3.6
.32
.901.2
2.26.0
.41.03
1.03
.26
.521.5
1.03
.7.9
1.74.7
.3.8
.8
.2
.41.2
.8
2859
2.32.6
3171
126201
86112
3186
6.615.4
6.810.1
1327
1.01.2
1432
5791
4050
1439
37
3.14.6
88
40
2.4
59
152
103
57
10.8
8.6
40
18.2
1.1
27
69
46.8
26
4.9
3.9
>1
11
2
>99
10
3
>159
220
>3
3555
.53.82
88132
2231
1529
1.11.6
4060
1014
713
0.50.7
116
24
22
1.3
53
11
10
0.6
>3
6
16
4
Longleaf pine must be de-winged dry. In a few pines, dewinging can be simplified by first wetting the seeds and then
drying them. Proper redrying precludes loss in seed quality
(Wang 1973). The loose seedwings can be fanned out
(Stoeckeler and Slabaugh 1965; Wakeley 1954). Mechanical
Pinus
833
Species
P. ponderosa
var. ponderosa
var. scopulorum
P. pumila
P. pungens
P. quadrifolia
P. radiata
P. resinosa
P. rigida
P. roxburghii
P. sabiniana
P. serotina
P. sibirica
P. strobiformis
P. strobus
P. sylvestris
P. taeda
P. thunbergiana
P. torreyana
P. virginiana
P. wallichiana
Place
collected
Black Hills
West Virginia
California
California
Lake States
Pennsylvania/
New York
India
California
SE US
Siberia
Arizona
Europe, E US
Japan, Korea, NE US
California
India
Seed weight
/cone wt
/cone vol
g/kg
oz/lb
kg/hl
lb/bu
2070
39
30
9
1020
.31.0
.6
.45
.14
1.5.3
.82.6
1.93
.52
.39
.641.0
.62.0
1.5
.4
.3
.5.8
2030
80
2030
20
2030
30
.3.45
11.2
.3.45
.3
.3.45
.45
1.03
.52
3.5
.42.2
.50.8
.81.7
.261.0
.641.2
.8
.4
2.7
.31.7
.4.6
.6.63
.2.8
.5.9
6884
1.82.6
2335
66166
Lot
723
1015
3138
0.81.2
1016
3076
26
29
24
75
2.1
29
115
12
13
11
34
1.0
13
52
185
74
11
3
3
7
529
94181
4282
6.825
311
1.21.4 0.50.6
104139
4763
3.54.6 1.62.1
5.56.4 2.52.9
39117 17.553
74245 33.8111
2758
1226
57110
2650
8.817.6
.4.8
101201
4691
1623
710
136
12
1.3
119
4.0
6.0
58
165
40
75.0
11.0
122
20
62
25
0.6
54
1.8
2.7
26
75
18
34
0.5
55
9
10
36
3
4
>10
10
300
>346
652
50
7
30
163
Sources: Barnett and McLemore (1967b), Cooling (1968), Curtis (1955), Debazac (1964), Delevoy (1935), Eliason (1942), Heit (1963, 1969), Karschon (1961), Krugman
and Jenkinson (1974), Letourneux (1957), Luckhoff (1964), Magini and Tulstrup (1955), Miller and Lemmon (1943), Mirov (1936), Nather (1958), NBV (1946), Ohmasa
(1956), Poynton (1961), Pravdin (1963), Rafn (1915), Read (1932), Sen Gupta (1936), Steinhoff (1964), Stoeckeler and Jones (1957), Sudworth (1908), Swingle (1939),
Takayama (1966),Troup (1921), Wappes (1932).
2030
060
030
90
60
60180
60
0
3060
060
120150
0
030
720
60
030
0
60120
030
6090
60120
60
1590
3060
3060
3090
30
1590
60
Pinus
835
Seeds
treated
+
+
P. aristata
P. arizonica
P. attenuata
P. banksiana
+
P. canariensis
P. caribaea
P. cembra
+
+
+
+
P. cembroides
P. clausa
P. contorta
var. contorta
var. latifolia
+
+
+
var. murrayana
P. coulteri
+
+
P. densiflora
+
P. echinata
+
+
+
P. edulis
P. elliottii
var. densa
var. elliotti
+
P. engelmannii
P. flexilis
+
+
+
P. gerardiana
P. glabra
P. halepensis
836
+
+
Germinative energy
Rate
Total
amt
Time
(mean
(%)
(days)
%) Samples
Daily
light
(hr)
Seed
medium
8
0
0
24
0
0
24
8
8
8
8
0
8
>8
0
0
0
0
8
8
A, P
pe, s
A, P
P
pe, s
P, pl
A, P
s
A, P
s
s
A
A
P
P, s
A
s
B, P
K
s+v
30
21
30
32
35
22
30
30
30
30
20
30
30
30
22
20
21
20
10
20
21
20
24
26
20
20
20
20
20
20
20
20
20
20
20
20
1618
28
365
14
22
30
20
30
120
14
8
23
28
21
21
28
90
60
28
21
30
75
52
79
69
86
54
72
58
63
2142
55
86
85
10
5
10
10
5
9
20
7
1737
28
14
20
30
91
86
75
92
85
87
69
75
74
76
72
37
62
90
90
74
>7
8
1
4
14
29
6
9
4
3
8
1
19
>8
0
8
0
0
10
0
8
8
0
0
0
>8
8
8
0
0
A, P, v
pe, s
A, P
s
s
s
P, s
s, v
A, P
s
s
A
A, P
s+v
pl + s
A
P
30
30
30
28
28
26
30
30
30
30
30
30
22
22
30
32
20
20
20
14
14
26
20
20
20
20
20
24
20
22
22
20
21
28
50
21
62
62
30
28
28
21
30
2460
30
28
28
27
28
16
73
75
54
88
81
80
10
15
15
14
10
60
80
80
73
65
75
37
87
83
74
90
90
96
3
29
10
9
12
3
3
4
20
139
148
>8
16
>8
16
16
0
0
8
8
8
0
8
16
16
0
0
0
s+v
K
A, P
K
K
P
B, P
A
s+v
A
A
A, P
s+v
s+v
A, P
A
A
22
22
30
22
22
32
30
30
22
30
30
22
22
22
22
20
22
22
21
20
20
22
20
21
20
22
22
20
2022
1819
32
28
28
26
26
16
21
21
27
30
3060
21
30
30
28
30
30
3079
86
80
75
70
69
85
46
5066
6580
711
7
10
9
4
14
13
30
20
1620
3282
87
89
84
88
42
82
47
98
98
79
89
30
28
392
83
4
1
1
2
25
30
5
12
Temp (C)*
Light
Dark
Days
Table 10Pinus, pine: seed germination test conditions and results (continued)
Germination test condition
Species
Seeds
treated
P. heldreichii
+
+
P. jeffreyi
+
+
P. kesiya
P. koraiensis
P. lambertiana
+
+
+
+
+
P. leiophylla var.
chihuahuana
P. merkusii
P. monophylla
P. monticola
+
+
P. mugo
P. muricata
+
+
P. nigra
P. palustris
P. parviflora
P. peuce
+
+
+
P. pinaster
P. pinea
P. ponderosa
var. ponderosa
+
+
+
+
+
+
var. scopulorum
P. pumila
P. pungens
P. quadrifolia
P. radiata
+
+
P. resinosa
P. rigida
+
Daily
light
(hr)
Seed
medium
Temp (C)*
Light
Dark
Days
P
Germinative energy
Rate
Total
amt
Time
(mean
(%)
(days)
%) Samples
8
8
8
24
0
8
0
>8
0
24
A, P
A
A
A, P, s
P
v
A
s
P
P, s
v
P, s
30
30
30
22
30
30
30
30
30
30
22
20
20
21
20
26
20
20
20
20
20
20
26
28
40
40
28
30
21
28
60
28
28
28
30
95
82
14
49
14
25
21
72
69
99
79
86
18
8595
59
55
14
1
1
5
1
1
4
5
1
0
8
8
24
0
8
8
0
8
0
0
24
8
0
16
16
0
8
>8
0
0
8
8
0
0
8
0
pl
A
A
pl
P, s
pl + s
A, P
A
A
A
v
P
A, P
A
P
s+v
P
s
A
s
A
A
A, P
A
s
A
A
30
20
75
30
22
30
30
30
30
2226
30
30
20
22
22
30
25
30
20
20
20
20
24
20
20
21
20
22
20
20
20
20
20
20
20
20
22
18
20
25
20
22
20
20
20
20
20
18
22
21
21
35
28
71
1421
30
20
35
21
30
14
30
21
30
1014
28
28
28
30
35
28
30
21
21
22
60
3950
39
25
76
70
91
54
90
71
41
70
30
88
11
10
10
7
10
10
10
35
7
20
7
14
70
59
67
8690
44
45
80
38
85
92
86
95
80
69
79
83
81
98
3
1
1
2
11
30
30
5
1
49
49
100
8
5
4
3
8
>8
0
16
0
>8
0
0
24
8
>8
>8
0
0
8
0
8
A
A
pe, s
K
s
A
pe + s
pe, s
A
A
P
v
A, P
s
A, P
A
A
30
30
29
22
20
30
2226
30
20
30
30
30
30
30
30
20
20
18
22
30
20
2129
24
20
20
20
20
20
20
20
20
21
21
3060
30
3060
21
4560
40
30
28
25
28
14
30
14
30
45
1487
84
50
46
16
69
2575
77
24
60
729
11
19
10
725
10
10
18
67
59
86
64
77
55
65
69
81
67
83
75
86
47
70
100
186
4
40
9
3
9
1
9
15
23
551
6
6
19
Pinus
837
Table 10Pinus, pine: seed germination test conditions and results (continued)
Germination test condition
Seeds
treated
Species
P. roxburghii
P. sabiniana
P. serotina
P. sibirica
+
+
+
P. strobiformis
P. strobus
+
+
+
P. sylvestris
P. taeda
P. thunbergiana
P. torreyana
P. virginiana
P. wallichiana
+
+
Daily
light
(hr)
Seed
medium
0
0
0
24
8
0
0
0
0
>8
>8
>8
8
8
24
8
>8
16
>8
>8
0
0
>8
>8
0
>8
0
A
s
s
A
pl + s
s
A
A
pl
A, P
K
K
A, P
A
s
A
A, P
pl + s
A, P
A
A
pe, s
A
K
pe, s
A, P
A
Temp (C)*
Light
Dark
2226
22
20
20
30
30
30
22
30
30
2225
20
30
22
30
24
25
27
30
22
30
30
24
2022
22
22
30
22
20
24
20
20
22
20
20
20
20
22
20
24
21
18
20
22
20
20
21
Days
30
30
30
30
21
60
>60
35
46
21
40
40
14
30
50
30
28
30
21
30
28
60
21
28
30
28
60
Germinative energy
Rate
Total
amt
Time
(mean
(%)
(days)
%) Samples
79
90
33
78
46
1899
90
50
69
87
44
10
10
10
10
14
17
10
10
10
20
83
81
76
13
73
7
40
94
39
100
93
81
59
8999
2199
90
75
76
85
81
90
65
64
5
135
3
1
1
1
4
8
31
20
28
99
99
36
18
481
4
19
100
21
29
5
12
Sources: Andresen (1965), Asakawa (1957), AOSA (1965, 1996), Graber (1965), Heit (1958, 1963), Heit and Eliason (1940), ISTA (1966), Kamra (1969), Krugman and
Jenkinson (1974), Luckhoff (1964), Magini (1955), McIntyre (1929), McLemore (1961a), Nather (1958), Ohmasa (1956), Rafn (1915), Rohmeder and Loebel (1940), Rossi
(1929), Sen Gupta (1936), Simak and others (1961).
Note: The + symbol shows that the seeds were pretreated, usually by moist chilling. Letters show the seed germination media that were used, as follows: A = absorbent
paper (filter, blotter), B = blotters supporting and covering seeds, K = Kimpak, P = absorbent medium in covered petri dish, pe = peat, pl = perlite, s = sand or soil,
v = vermiculite.
* Alternating periods of high and low temperatures typically were 8 and 16 hours (8/16).The light period normally coincided with the warmer temperature.Temperatures
of 10, 15, 20, 25, 30, and 35 C equal 50, 59, 68, 77, 86, and 95 F, respectively.
Seeds were extracted from old cones.
testing for the very dormant stone pines (ISTA 1996), but
they are used on other species for rapid tests only.
Tetrazolium test estimates are highly dependent on the analysts experience and seed age, and too often exceed the percentages attained in standard germination tests (Stoeckeler
and Slabaugh 1965; Tanaka 1984).
Nursery and field practices. Pines are grown successfully in diverse soil types in most regions of the United
States. Various regional handbooks, manuals, and reports on
forest tree nursery and reforestation practices describe bareroot seedling production, illustrate typical nursery equipment and facilities, and provide critical guides on soil management, bed preparation, seed treatment, seed sowing,
seedling cultural regimes and pest control, undercutting,
wrenching, lifting, packing, transplanting, cold and freeze
storage, shipping, field handling, and safe planting times
(Cleary and others 1978; Duryea and Landis 1984; Heit
1964; ICIA 1963; Jenkinson 1980; Jenkinson and others
1993; Lowman and others 1992; Schubert and Adams 1971;
Schubert and others 1970; Stoeckeler and Jones 1957;
Stoeckeler and Slabaugh 1965; Wakeley 1954). Together,
these references and work cited therein capture the bulk of
knowledge and practical experience gained on pine seedling
production and planting in the United States.
Productive nursery soils are invariably deep, arable, fertile, and drain rapidly to ensure root aeration. Most of the
large mechanized nurseries fumigate their soils (in late summerearly fall) to control soil-borne diseases, insects, nematodes, and weeds before the scheduled sowing in fall or
spring (Thompson 1984). Nursery climates, soils, seed
sources, and their interactions have resulted in a wide range
of cultural regimes and practices. Recommended practices
for 35 different species and varieties show the wide ranges
encountered in seed chilling time, in sowing time, density,
and depth, in bed mulches, and in yields and types of planting stock produced (table 11).
In temperate regions, seeds can be sown in fall or
spring. Seeds with embryo dormancy can be sown in fall
without pretreatment. Normally, both dormant seeds and
nondormant seeds are sown in spring more often than in fall.
Dormant seeds in spring-sowings must be pretreated to
enable rapid and complete germination. Applying the same
treatment to the dormant seeds of all pines is inadvisable.
Success of the treatment applied depends on species and
seed source, and the one applied should be the one that
achieves the highest germination and seedling emergence for
the particular seedlot. Seedlings produced in fall-sowings
after 1 growing season are often larger and better developed
than those produced in spring-sowings. But fall-sowings are
inherently risky. They typically incur excessive overwinter
Pinus
839
Species
P. attenuata
P. banksiana
60
0
Sponge Rok
None
613
3550
70
1+0
1+0, 2+0, 1/2+11/2,
1+1, 1+2
1+0, 2+0, 1/2+11/2
1+0
3
3
10
13
36
Press
6
None
Sawdust
Peat mossII
None
Sand
Pine straw
Sawdust
13
613
13
48
60
7075
80
60
80
1+0
2+0
1+0, 2+0
1+0
2+0, 3+0, 1+1
1+0
2+0
35
3035
Press
Press
13
80
5874
1+0
1+0
Spring
Spring
Spring
Spring
Spring
Spring
Spring
Fall
Spring
Spring
Spring
2530
30
3035
2530
35120
50
3075
5060
15
30
610
10
1013
13
610
10
319
1319
Press
613
Pine straw
Sawdust or
pine straw
None
None
None
Sawdust
Sawdust II
Peat moss
Peat moss II
Peat moss
Pine straw
613
613
613
613
613
5880
50
2180
33
3290
55
3760
6065
75
1+0
1+0
Spring
2546
610
None
4880
3540
5065
36
13
None
None
70
70
2033
613
Straw
1928
1+0, 2+0
2575
3050
319
610
Peat moss II
None
613
3470
6580
3035
2050
3060
40
50100
Press
313
16, or
Press
313
Press
3
3035
5585
2570
60
31
3035
6 or
press
6390
1+0, 2+0
0
1520
60
Fall
Spring
Spring
cast
2530
13
610
Sand
3
Sawdust, or
36
wood fiber
Peat moss II
613
Pine straw
Burlap or straw
in winter
Sand, pine
613
Needles or
sawdust
None
None
1+0
2+0, 3+0, 2+1,
2+2
2+0
1+1, 2+1
2+0, 3+0, 2+1,
2+2
1+0, 2+0, 2+1,
2+2
1+0
2+0, 1+1
0
14
Spring
Fall or
spring
Fall or
spring
Spring
Fall or
spring
Spring
Spring
Fall
Spring
Fall
Spring
Spring
Fall or
spring
Fall
Spring
46
5080
1+1, 1+1+1
1+0, 2+0
P. palustris
P. pinaster
P. ponderosa
var. ponderosa 2860
P. radiata
P. resinosa
3545
0
P. rigida
P. roxburghii
P. strobus
0
0
0
30
0
260
3060
0
P. sylvestris
P. taeda
P. thunbergiana
P. virginiana
P. wallichiana
P. washoensis
25
30
10
6
6
613
Spring
Spring
Spring
Spring
Spring
Spring
Spring
30
48
2560
25
50
40
30
Spring
Spring
None
None
Type
80
5060
2860
0
90
90
4290
4050
2840
0
3545
0
0
Spring
Fall or
spring
Spring
Spring
Planting stock
Depth
Yield
(mm)
(%)
P. canariensis
0
P. clausa
0
P. contorta
var. contorta
28
var. latifolia
2835
var. murrayana 3060
P. coulteri
60
P. densiflora
0
P. echinata
1560
P. edulis
60
P. elliottii
var. densa
0
var. elliotti
0
P. jeffreyi
P. kesiya
P. lambertiana
P. monophylla
P. monticola
P. mugo
P. muricata
P. nigra
Seed sowing
Seedbed mulch
Density Depth
(mm)
Material
Season
(/ft2)
Sources: Claveria (1953), Derr (1955), Goor (1955), Heit (1968a), Krugman and Jenkinson (1974), Letourneux (1957), Magini and Tulstrup (1955), NBV (1946), Schubert
and Adams (1971), Shoulders (1961), Stoeckeler and Jones (1957), Stoeckeler and Rudolf (1956), Stoeckeler and Slabaugh (1965),Troup (1921),Veracion (1964, 1966).
* Seeds were chilled in a moist medium at 0.5 to 5 C.
Multiply number per square foot by 10.758 to convert to number per square meter.
The word press indicates that seeds were pressed flush to the soil surface.
Type of planting stock codes the number of growing seasons and transplant operations for seedlings in the nursery: 1+0 stock is lifted and shipped to the field for
outplanting after 1 nursery growing season, and 2+0 stock, after 2 seasons; 1+1 stock is transplanted after its first growing season and shipped to the field after its second
season.
II Mulch was not always used.
840
Pinus
841
References
Allen R, Wardrop AB. 1964. The opening and shedding mechanism of the
female cones of Pinus radiata. Australian Journal of Botany 12: 125134.
Altman PL, Dittmer DS. 1962. Biological handbook of growth. Washington,
DC: Federation American Society Experimental Biology. 608 p.
Andreev VN. 1925. Pinus peuce Griseb. [in Russian]. In: Dendrology.Volume
1, Gymnosperms. Kiev: Ukraine State Printing Office: 96
Andresen JW. 1963. Germination characteristics of Pinus rigida seed borne
in serotinous cones. Broteria 32(3/4): 151178.
Andresen JW. 1965. Stratification of seed of Pinus strobiformis.Tree Planters
Notes 72: 57.
Asakawa S. 1957. Studies on hastening germination of the seeds of fiveleaved pines [in Japanese]. Government Forest Experiment Station,
Meguro (Tokyo) Bulletin 100: 4154.
AOSA [Association of Official Seed Analysts]. 1965. Rules for testing seeds.
Proceedings Association of Official Seed Analysts 54(2): 12112.
AOSA [Association of Official Seed Analysts]. 1996. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
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Pinus
847
FabaceaePea family
848
Pulpy aril
1.5 cm
Seed
References
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348353 [Weed Abstracts 28(5): 1618; 1979].
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639640.
Pithecellobium
849
PlatanaceaePlanetree family
Platanus L.
sycamore
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Common name(s)
Occurrence
P. occidentalis L.
P. occidentalis var. Sarg. glabrata (Fern.)
P. orientalis L.
P. racemosa Nutt.
Location
Flowering
Fruit ripening
Seed dispersal
P. occidentalis
P. orientalis
P. racemosa
E US
NE US
MarMay
May
SeptNov
SeptOct
JuneAug
JanApr
JuneDec
850
Figure 1Platanus, sycamore: fruiting heads of P. occidentalis, American sycamore (top) and P. racemosa, California
sycamore (bottom).
Figure 2Platanus, sycamore: single achenes of P. occidentalis, American sycamore (top) and P. racemosa,
California sycamore (bottom).
long-
Platanus
851
10% and temperatures of 0 to 5 C are suitable for shortterm storage of up to 5 years. For longer storage periods,
sub-freezing temperatures (18 C) at the same moisture
content are recommended (Bonner 1979). The upper limit of
storage potential for sycamore is not yet known, but current
research suggests that it will be far beyond 10 years under
optimum conditions (Bonner 1994). To maintain low seed
moisture in moist surroundings, the dried seeds must be
stored in moisture-proof containers, such as polyethylene
bags or fiber drums with plastic liners (Bonner 1979).
Several species of Aspergillus fungi have been identified as
pathogens that harm viability of sycamore seeds in storage
(Fakir and others 1971), but they have never been a major
problem.
Pregermination treatments. Moist stratification for
60 to 90 days at 5 C in sand, peat, or sandy loam has been
reported as beneficial for germination of California
sycamore (Bonner 1974). The other sycamores have no dormancy, and pregermination treatments are usually not
required for prompt germination (Bonner 1972a; Webb and
Farmer 1968). Germination rate of sycamore can be
increased by treating with gibberellin (GA3) at 100 to 1,000
mg/liter, but this increase seems to be simple growth stimulation that is not involved in seed dormancy (Bonner 1976).
Germination tests. Germination can be easily tested
on wet paper or sand or even in shallow dishes of water
(table 4). Official testing prescriptions call for alternating
day/night temperatures of 30/20 C on the top of moist blotters for 14 days (AOSA 1993). A large percentage of the
sound seeds will usually germinate, but the great variation in
number of sound seeds among lots will result in varied germination percentages. Surface sterilization of the seeds with
a 30-second dip in a 1% commercial bleach solution is often
beneficial to laboratory germination (Mullins 1976). Rapid
viability tests can also be made on sycamore with tetrazolium staining and x-radiography (Bonner 1974).
Species
Place of collection
/kg
P. occidentalis
LouisianaMississippi*
SE US
Denmark
United States
294,370589,620
192,340500,100
178,600357,200
249,160370,440
P. orientalis
852
133,500267,400
87,2330226,800
81,000162,000
113,000168,000
Average
/kg
426,160
330,530
282,240
308,700
/lb
193,270
149,900
128,000
140,000
Samples
100+
28
8
2+
neutral to slightly alkaline soils, damping-off may be a problem. Root pruning in midsummer is recommended to promote growth of smaller roots, and some nurseries prune
seedling tops in late July or August to reduce size. Seedlings
should not be both root- and top-pruned during the growing
season (Briscoe 1969). Sycamore is usually outplanted as
1+0 stock, and oriental planetree is often planted as 1+1 or
2+0 seedlings in Europe (Bonner 1974). The sycamores are
easy to propagate vegetatively by dormant or greenwood
cuttings (Dirr and Heuser 1987), and many plantations of
sycamore have been established in the South by these techniques. Some tests show no difference in growth after 1 year
between seedlings and cuttings (Garrett 1975).
References
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
Beland JW, Jones L. 1967. Self-incompatibility in sycamore. In: Proceedings, 9th
Southern Conference on Forest Tree Improvement; 1967 June 89;
Knoxville,TN. Spons. Publ. 28. Macon, GA: USDA Forest Service, Eastern
Tree Seed Laboratory: 5658.
Bonner FT. 1972a. Laboratory germination testing of American sycamore.
Proceedings of the Association of Official Seed Analysts 62: 8487.
Bonner FT. 1972b. Maturation of sweetgum and American sycamore seeds.
Forest Science 18: 223231.
Bonner FT. 1974. Platanus L., sycamore. In: Schopmeyer CS, tech. coord.
Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 641644.
Bonner FT. 1976. Effects of gibberellin on germination of forest tree seeds
with shallow dormancy. In: Hatano K, Asakawa S, Katsuta M, Sasaki S,
Yokoyama T, eds. Proceedings, Second International Symposium on
Physiology of Seed Germination; 1976 October 1830; Fuji, Japan.Tokyo:
Government Forest Experiment Station: 2132.
Bonner FT. 1979. Collection and care of sycamore seeds. Southern Journal
of Applied Forestry 3(1): 2325.
Bonner FT. 1994. Predicting seed longevity for four forest tree species with
orthodox seeds. Seed Science and Technology 22: 361370.
Bonner FT, Switzer GL. 1974. Mechanical separation of full and empty
sycamore seeds. In: 1974 Southeastern Nurserymens Conferences; 1974
July 1718; Nacogdoches,TX and August 68; Gainesville, FL. Atlanta:
USDA Forest Service, Southeastern Area State & Private Forestry:
95100.
Briscoe CB. 1969. Establishment and early care of sycamore plantations.
Res. Pap. SO-50. New Orleans: USDA Forest Service. Southern Forest
Experiment Station. 18 p.
Cooper DT, Filer TH Jr, Wells OO. 1977. Geographic variation in disease
susceptibility of American sycamore. Southern Journal of Applied
Forestry 1(4): 2124.
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Misc. Pub. 434. Washington, DC: USDA. 159 p.
Fakir GA, Welty RE, Cowling EB. 1971. Prevalence and pathogenicity of
fungi associated with achenes of sycamore in the field and in storage.
Phytopathology 61: 660668.
Ferguson RB, Land SB Jr, Cooper DT. 1977. Inheritance of growth and
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Karrfalt RP, Helmuth RE. 1984. Preliminary trials on upgrading Platanus occidentalis with the Helmuth electrostatic seed separator. In: Murphy PM,
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7981.
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Washington, DC: USDA Forest Service. 30 p.
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Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
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occidentalis L.) [MSc thesis]. Starkville: Mississippi State University, School
of Forest Resources. 57 p.
Swingle CF (comp.). 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA Soil
Conservation Service. 198 p.
Vande Linde F. 1960. Nursery practice for growing sycamore seedlings.
Tree Planters Notes 41: 1516.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
Webb CD, Farmer RE Jr. 1968. Sycamore seed germination: the effects of
provenance, stratification, temperature, and parent tree. Res. Note SE100. Asheville, NC: USDA Forest Service, Southeastern Forest
Experiment Station. 6 p.
Wells OO, Schmidtling RC. 1990. Platanus occidentalis L., sycamore. In:
Burns RM, Honkala BH, tech. coords. Silvics of North America.Volume
2, Hardwoods. Agric. Handbk. 654. Washington, DC: USDA Forest
Service: 511517.
Williams RD, Hanks SH. 1976. Hardwood nurserymans guide. Agric.
Handbk. 473. Washington, DC: USDA Forest Service. 78 p.
Platanus
853
CupressaceaeCypress family
854
tained for at least 5 years. For longer storage periods, subfreezing temperatures of about 18 C are recommended
(Bonner 1991).
Germination tests. For most seedlots, pretreatments
are not needed for germination tests, although beneficial
effects have been reported for cold stratification for 1 to
1 1/2 months and for short soaks in weak solutions of gibberellic acid (Dirr and Heuser 1987; Schopmeyer 1974).
Both AOSA (1993) and ISTA (1993) recommend testing
oriental arborvitae on the top of moist paper media at a constant 20 C for 21 days; no pretreatments are prescribed.
Germination is epigeal.
Nursery practice. Sowing outdoor nursery beds
should take place in the spring at a depth of 6 to 9 mm (1/4
to 3/8 in) and no mulch. A bed density of 375 to 430/m2 (35
to 40/ft2) has been recommended (Schopmeyer 1974).
Under most conditions, seedlings are grown for 2 years
before outplanting, although this may vary according to
individual nursery practices. Container production in greenhouses is also possible, using techniques that are successful
with the closely related Thuja species.
Vegetative reproduction, while more difficult than it is
with Thuja species, is possible with oriental arborvitae and
is widely used for ornamental cultivars. Dirr and Heuser
(1987) recommend taking cuttings from June to August,
treating with indole-butyric acid (IBA) in talc and Benlate,
then rooting with mist and bottom heat in a peat and perlite
mixture (2:1, v/v).
References
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 159.
Bonner FT. 1991. Seed storage. In: Gordon AG, Gosling P, Wang BSP, eds.
Tree and shrub seed handbook. Zurich: International Seed Testing
Association: 13: 14.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
ISTA [International Seed Testing Association]. 1993. Rules for testing seeds:
rules 1993. Seed Science and Technology 21 (Suppl.): 1259.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan. 1290 p.
Rushforth KD. 1987. Conifers. New York: Facts on File. 232 p.
Schopmeyer CS. 1974. Thuja, arborvitae. In: Schopmeyer CS, tech. Coord.
Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 805809.
Vidakovic M. 1991. Conifers: morphology and variation. Zagreb, Croatia:
Graficki zavod Hrvatske. 754 p.
Platycladus
855
SalicaceaeWillow family
Populus L.
poplar, cottonwood, aspen
Gary W. Wyckoff and John C. Zasada
Mr. Wyckoff is a forester at Mead-Westvaco in Escanaba, Michigan;
Dr. Zasada retired from the USDA Forest Services North Central Research Station
856
Common name(s)
Occurrence
lanceleaf cottonwood,
lanceleaf poplar, smooth-bark
cottonwood, & Andrews poplar
white poplar, abele
P. alba L.
P. angustifolia James
P. balsamifera L.
P. balsamifera L. spp. trichocarpa
(Torr. & Gray ex Hook.) Brayshaw
P. trichocarpa Torr. & Gray
P. x canescens (Ait.) Sm. (pro sp.)
P. deltoides Bartr. ex Marsh.
P. deltoides Bartr. ex Marsh.
ssp. monilifera (Ait.)
P. deltoides var. occidentalis Ryb.
P. deltoides Bartr. ex Marsh. ssp.
wislizeni S.Wats.
P. fremontii var. wislizeni S.Wats.
P. euphratica Olivier
P. fremontii S.Wats.
P. grandidentata Michx.
P. heterophylla L.
P. laurifolia Lebed.
P. maximowiczii A. Henry
P. nigra L.
P. x petrowskiana R.I. Schrod. ex Regel
P. sieboldii Miq.
P. simonii Carriere
P. tremula L.
P. tremuloides Michx.
narrowleaf cottonwood,
narrowleaf poplar, black cottonwood,
mountain cottonwood, lamo
balsam poplar, tacamahac poplar,
cottonwood, hackmatack, tacamahac
black cottonwood, California
poplar, cottonwood, balsam
cottonwood, western balsam poplar
gray poplar
eastern cottonwood,
eastern poplar
plains cottonwood, plains
poplar, cottonwood,Texas
cottonwood
Rio Grande cottonwood, Rio
Grande poplar, cottonwood,
valley cottonwood,Wislizenus
cottonwood, lamo
Euphrates poplar, bahan,
Gharab-Palk-Saf-Saf
Fremont cottonwood, Fremont
poplar, cottonwood, Arizona
cottonwood, Macdougal cottonwood,
lamo
bigtooth aspen, largetooth
aspen, aspen, poplar, popple
swamp cottonwood, swamp
poplar, cottonwood, black
cottonwood, river cottonwood,
downy poplar
laurel poplar
Japanese poplar
black poplar, European black poplar
Petrowsky poplar, Russian poplar
Siebold aspen, Japanese aspen
Simon poplar
European aspen, tremble,
Zitterpappel
quaking aspen, quaking asp,
aspen, golden aspen, mountain
aspen, trembling aspen,Vancouver
aspen poplar, popple, alamo blanco
Populus
857
western United States and Canada, where they are the only
tree cover, cottonwoods provide critical habitat for some
species (Friedman and others 1997; Rood and others 1995;
Stromberg 1993). They provide bark and male flowerbuds
for specialist species such as beaver (Castor canadensis) and
ruffed grouse (Bonasa umbellus) as well as forage for generalist browsers such as moose (Alcea alcea), elk (Cervus elaphus), and deer (Odocoileus spp.) (Burns and Honkala 1990;
Dickmann and Stuart 1983; Dickmann and others 2001;
Graham and others 1963; MacKinnon and others 1992;
Peterson and Peterson 1995; Viereck 1987).
Poplars have long been important as ornamentals and in
horticulture (tables 1 and 2). They were commonly used for
amenity plantings in urban areas as ornamentals and landscaping. In rural areas, they have been used in shelterbelts
and as ornamentals. The most rapidly increasing use of the
species today is in intensive culture for wood fiber and biomass for energy in such diverse climates as the lower
Mississippi Valley, western and eastern Washington, and
western Minnesota. Shortages of fiber due to regulation of
harvesting to protect critical wildlife habitat and old-growth
forests, declining traditional forest land base, and concerns
about effects of carbon dioxide (CO2) from combustion of
fossil fuels have renewed interest in intensive culture.
Table 2Populus, poplar, cottonwood, aspen: height at maturity, first cultivation, minimum seed-bearing age, and seed
crop frequency
Species
P. x acuminata
P. alba
P. angustifolia
P. balsamifera
P. balsamifera spp.
trichocarpa
P. x canescens
P. deltoides
spp. monilifera
spp. wislizeni
P. fremontii
P. grandidentata
P. heterophylla
P. laurifolia
P. maximowiczii
P. nigra
P. simonii
P. tremula
P. tremuloides
Years
between large
seedcrops
Year first
cultivated
10.715.4 (18.5)
15.442.1
10.715.4 (18.5)
18.536.3
15.461.5
1898
LC
1893
Before 1689
1892
510
1015
810
10
29.230.8 (40)
24.658.5
12.330.8
2.330.8 (40)
15.430.8
9.227.7 (30.8)
24.627.3 (30.8)
to 15.4
to 30.2
18.530.8
to 12.3+
21.538.5
15.427.7 (30.8)
LC
Before 1750
1908
1894
1904
1772
1656
1830
Before 1890
LC
1862
LC
1812
815
10
10
5
510
1020
10
810
10
812
10
810
1020
1
1
1
1
45
1
1
1
4.5
45
858
Minimum
seed-bearing
age (yr)
Height at
maturity* (m)
Populus
859
Location
Flowering
P. x acuminata
P. alba
Nebraska
Nebraska
NE US
S Michigan
Alberta
Lake States
Interior Alaska
Vancouver Island, British Columbia
Great Lakes region
Lower Mississippi Valley
NE US
Alberta
Lincoln, Nebraska
Albuquerque, New Mexico
Central Arizona
Syracuse, New York
S Michigan
New England
N Great Lakes region
Mississippi
Rochester, New York
Rochester. New York
N Great Lakes region
Great Lakes region
Alberta, Canada
New England
Interior Alaska
N Great Lakes region
May
AprMay
AprMay
MarApr
Late Apr
AprMay
MayJune
AprJune
Late Apr
Early Marearly Apr
Aprearly May
AprMay
AprMay
mid-Febmid-Mar
Mid-Marmid-Apr
Late Marmin-May
Mid-MarApr
Late Aprearly May
MarMay
Late Apr
Apr
Mid-Apr
Late Marearly May
Early Aprearly May
Mid-MarApr
AprMay
Mid-Apr
July
Early June
MayJune
MayJune
Early Juneearly July
MayJune
July
Late Maymid-July
Late May
Mid-Maylate Aug
Maymid-June
late Juneearly Aug
June
JuneJuly
MarApr
Mid-Maylate May
early Maylate June
Mayearly June
Late Mayearly June
AprJuly
Aug
Late May
Mid-May
Mid-Maymid-June
Mid-Maymid-June
Mayearly June
Late Mayearly June
Mid-May
P. balsamifera
P. balsamifera spp. trichocarpa
P. x canescens
P. deltoides
spp. monilifera
spp. wislizeni
P. fremontii
P. grandidentata
P. heterophylla
P. maximowiczii
P. nigra
P. tremula
P. tremuloides
P. tremula
860
Catkins (no.)
Seeds (no.)
8,700
25
1,200
1,275,000
25
500
205,000
45
10,000
3,300,000
100
40,000
54,000,000
tree at
861
Flowering of 24- to 36-year-old grafted clones of bigtooth aspen (3 female and 7 male) was also followed in the
above-mentioned Wisconsin arboretum. The female clones
flowered in 35% of the years in a 20- to 25-year observation
period and the males in 18% of the years. Flowering of individual female clones occurred in from 29 to 40% of the
years; male clones flowered in 0 to 50% of the years
(Wyckoff 1996).
Mature catkins are made up of many capsules, each
developed from an individual flower (figures 1 and 2). The
number of viable seeds per capsule has been reported to
vary from 2 to 7 for quaking aspen (Brown 1989; Henry and
Barnes 1977; Nagaraj 1952; Spies 1978); 2 to 10 for bigtooth aspen (Henry and Barnes 1977); 2 to 4 for
bigtoothquaking aspen hybrids (Henry and Barnes 1977);
about 1 for white poplar (Spies 1978); 1 to 2 for white
poplar hybrids with bigtooth aspen (Spies 1978); and 8 to 15
(Nagaraj 1952), 32 (Bessey 1904), and 40 to 60 for eastern
cottonwood (Farmer and Nance 1968). Estimates of seed
production per catkin for quaking aspen have varied considerably among studies and locations: 500 in central Alberta
(Brown 1989), 77 in northern Wisconsin (Rudolph 1978)
280 to 290 in southern Michigan, (Henry and Barnes 1977;
Spies 1978), and 150 to 300 in central Wisconsin (Einspahr
and Benson 1964). Henry and Barnes (1977) reported 500
seeds per catkin for bigtooth aspen. Spies (1978) reported
about 11 seeds per catkin for white poplar, 281 for quaking
aspen, and 102 for the hybrid P. rouleauiana Boivin.
Figure 2Populus, poplar: catkins consisting of mature
but unopened capsules.P.deltoides ssp. monilifera, plains
cottonwood (top); P. fremontii, freemont cottonwood
(middle); P. freemontii ssp. wislizeni, Rio Grande cottonwood
(bottom).
862
863
Table 4Populus, poplar, cottonwood, aspen: seed weights by soil sieve size (20 to 50 mesh)*
Seeds (millions)/weight
28-mesh
40-mesh
/g
/oz
/g
/oz
20-mesh
/g
/oz
Species
P. alba
P. balsamifera
spp. trichocarpa
P. x canescens
P. deltoides
P. grandidentata
P. tremula
P. tremuloides
50-mesh
/g
/oz
5.09
2.71
2.93
2.31
1.23
1.33
8.89
7.90
4.54
7.50
4.92
13.19
4.03
3.58
2.06
3.40
2.23
5.98
19.71
8.69
13.87
21.52
17.46
18.46
8.94
3.94
6.29
9.76
7.92
8.37
39.58
23.62
17.95
10.70
4.81
4.34
4.43
3.00
3.20
2.18
1.97
2.01
1.36
1.45
10.61
18.43
7.74
10.52
7.08
5.18
6.90
4.54
4.83
4.81
8.36
3.51
4.77
3.21
2.35
3.13
2.06
2.19
17.68
44.10
16.27
13.38
23.09
15.79
19.45
10.36
60.86
8.02
20.00
7.38
6.07
10.47
7.16
8.82
4.70
27.60
38.68
35.57
47.83
35.10
30.96
31.00
17.54
16.13
21.69
15.92
14.04
14.06
864
Species
/kg
Seeds (x million)/weight
Average
/lb
kg
P. deltoides
P. deltoides spp. monilifera
P. grandidentata
P. heterophylla
P. tremula
P. tremuloides
0.443.31
0.551.06
0.310.36
5.8916.65
5.516.62
0.201.50
0.250.48
0.140.16
2.667.55
2.53.0
Range
0.77
6.62
0.34
8.09
6.00
/lb
0.425
3.00
0.15
3.67
2.75
Samples
6+
4+
1
4
30
1980), although Wang (1982) reported good long-term storage at 11 to 15% moisture content (dry-weight basis). Seed
viability is maintained at lower moisture contents, but there
is an indication that seedling vigor is reduced under these
conditions (Simak 1980). Recommendations for drying time
to achieve these approximate moisture contents have varied:
for example, 2 to 3 days at 21 C (Moss 1938); 3 to 8 days
at 24 C (Faust 1936); air-drying for 7 days (Tauer 1979); 1
day at 35 to 40 C, and 1 day of drying over calcium chloride (CaCl2) (Simak 1980). If achieving a specified moisture
content is important, a method should be used that attains
the desired content as quickly as possible while not affecting
seed viability.
A number of research trials have compared storage
under vacuum and with various desiccants to storage in
sealed containers; unfortunately, the results are not definitive. In some cases, they work and in others they seem to be
of no benefit or may even have a negative effect on viability.
Benson and Harder (1972) compared 4-year germination
results from 40-mesh aspen and aspen hybrid seeds stored
over calcium chloride in a desiccator at 4.4 C and 24 C.
Freezer storage maintained higher levels of viability in each
of the 4 years with the exception of 2 hybrids of bigtooth
aspen during the first year. By year 4, all seedlots stored at
4.4 C were either nonviable or had considerably reduced
germination. Freezer-stored seeds maintained a high level of
germinative ability at the end of 4 years, with the exception
of gray poplar hybrids. Benson and Harders study (1977,
1996) indicates that germination of freezer-stored seed of
quaking aspen dropped from an average of 98% to 65% over
a 10-year period. Hybrid seeds and 50-mesh seeds generally
had lower germination than quaking aspen and 40-mesh
seeds after 10 years of freezer storage. Seed viability of 3 of
4 open-pollinated cottonwood seedlots stored at 24 C over
calcium chloride did not decrease during 8 years of storage.
Populus
865
Figure 6Populus, poplar: stages in the normal germination of a seed (adapted from Simak (1980) and McDonough
(1979).
Populus
867
868
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Populus
869
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Populus
871
FabaceaePea family
Prosopis L.
mesquite
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
legume.
Common name(s)
Occurrence
P. glandulosa Torr.
honey mesquite
872
873
Scarification
treatment
Germination
medium
Nicking
Nicking
None
H2SO4
Wet
Wet
Wet
Wet
paper
paper
paper
sand
Day
Temp (C)
Night
Avg %
germination
27
27
27
30
27
27
27
20
98
60
18
88
Figure 3Prosopis juliflora, mesquite: seedling development at 1, 2, 5, 10, and 25 days after germination.
874
References
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factors influencing Acacia constricta and Prosopis velutina establishment in
the Sonoran Desert. Journal of Range Management 46: 4348.
Davis CA, Barkley RC, Haussamen WC. 1975. Scaled quail foods in southeastern New Mexico. Journal of Wildlife Management 39: 496502.
Day AD, Ludeke KL. 1980. Direct seeding of tree species on copper mine
wastes in the southwestern United States. Journal of Environmental
Quality 9: 304306.
Felker P, Clark PR. 1981. Rooting of mesquite (Prosopis) cuttings. Journal of
Range Management 34: 466468.
Felker P, Clark PR, Osborn JF, Cannell GH. 1984. Prosopis pod production:
comparison of North American, South American, Hawaiian, and African
germplasm in young plantations. Economic Botany 38: 3651.
Ffolliott PF,Thames JL. 1983. Handbook on taxonomy of Prosopis in Mexico,
Peru and Chile. Rome: FAO. 31 p.
Glendening GE, Paulsen HA Jr. 1955. Reproduction and establishment of
velvet mesquite, as related to invasion of semidesert grasslands.Tech.
Bull. 1127. Washington, DC: USDA Forest Service. 50 p.
Gonzlez-Benito ME, Caz-Filho J, Prez C. 1994. Cryopreservation of
seeds of several legume species. Plant Varieties and Seeds 7:2327 [Seed
Abstracts 18(2): 499; 1995].
Goor AY, Barney CW. 1968. Forest tree planting in arid zones. New York:
Roland Press. 344 p.
Harden ML, Zolfaghari R. 1988. Nutritive composition of green and ripe
pods of honey mesquite (Prosopis glandulosa, Fabaceae). Economic
Botany 42: 522532.
Johnson CD. 1983. Handbook on seed insects of Prosopis species. Rome:
FAO. 55 p.
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Martin SC. 1948. Mesquite seeds remain viable after 44 years. Ecology 29:
393.
Martin SC, Alexander RR. 1974. Prosopis juliflora (Sw.) DC., mesquite. In:
Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington, DC: USDA Forest Service:
656657.
Sargent CS. 1965. Manual of the trees of North America (exclusive of
Mexico). 2nd ed. New York: Macmillan. 996 p.
Skolmen RG. 1990. Prosopis pallida (Humb. & Bonpl. ex Willd.) H.B.K.,
kiawe. In: Burns RM, Honkala BH, tech. coords. Silvics of North America.
Volume 2, Hardwoods. Agric. Handbk. 654. Washington, DC: USDA
Forest Service: 583586.
Solbrig OT, Cantino PD. 1975. Reproductive adaptations in Prosopis
(Leguminosae, Mimosoideae). Journal of the Arnold Arboretum 56:
185210.
Vines RA. 1960. Trees, shrubs and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
Von Carlowitz PG. 1986. Multipurpose tree and shrub seed directory.
Nairobi: International Council for Research in Agroforestry. 265 p.
RosaceaeRose family
Prunus L.
cherry, peach, and plum
Ted J. Grisez, Jill R. Barbour, and Robert P. Karrfalt
Dr. Grisez retired from the USDA Forest Services Northeastern Forest Experiment Station; Ms. Barbour is a
germination specialist at and Mr. Karrfalt is the director of the USDA Forest Services National Tree Seed
Laboratory, Dry Branch, Georgia
Growth habit, occurrence, and use. The genus
Prunusoften called the stone fruitsis one of the most
important genera of woody plants. Its 5 well-marked subgenera include the plums and apricots (Prunophora), the
almonds and peaches (Amygdalus), the umbellate cherries
(Cerasus), the deciduous racemose cherries (Padus), and the
evergreen racemose or laurel cherries (Laurocerasus). Plums
can be distinguished from peaches and almonds by lack of a
terminal bud, multiple flowers from a bud, and an elongated
pedicel (Janick and Moore 1996). Plums can be distinguished from cherries by the lack of a terminal bud, the
presence of a suture, a waxy bloom on the fruit, and a flatter
pit.
Nearly 200 speciesranging from prostrate shrubs to
trees over 30 m tallare found in the Northern Temperate
Zone, with a few in Central and South America (Harlow and
Harrar 1958; LHBH 1978; Rehder 1940). By far the greatest
number of species of cherries occur in eastern Asia (Hedrick
1915), but most of the long-cultivated food-producing
species originated in Europe and western Asia (table 1).
Over 100 species have been brought under cultivation, mostly as food crops or ornamentals (Rehder 1940), and 32 of
the more important species for planting in the United States
are described in table 1.
Many of the stone fruits have been cultivated since
ancient times for their edible fruits and a few for edible
seeds (almonds). Wild species have also been a source of
food for Native Americans and early European settlers in
this country and are still used to some extent. Many selections of wild plums have been propagated for fruit production. Several species are useful as ornamentals because of
their showy flowers, variety of growth habits, relatively fast
growth and ease of cultivation, and adaptability to a wide
variety of soils and climates (Hedrick 1915; Olson and
Nagle 1965; Rehder 1940; Strausbaugh and Core 1964).
Trees for fruit production and many ornamentals are
propagated by budding or grafting, but seed production is
875
Table 1Prunus, cherry, peach, and plum: nomenclature, growth habit, and occurrence
Scientific name synonym(s)
Common name(s)
Growth habit
Occurrence
P. alleghaniensis Porter
Allegheny plum,
sloe, Allegheny sloe,
Porter plum
American plum, wild
yellow plum, red plum,
goose plum, hog plum
Chickasaw plum,
sand plum
Tree or shrub
Tree or shrub
Tree or shrub
P. americana Marsh.
P. angustifolia Marsh.
P. armeniaca L.
Armeniaca vulgaris Lam.
P. avium (L.) L.
P. cerasus avium L.
Cerasus avium Moench
P. caroliniana (P. Mill.) Ait.
P. cerasifera Ehrh.
P. domestica var. myrobalan L.
P. myrobalana Loisel.
P. korolkowi Vilm.
P. cerasus L.
Cerasus vulgaris Mill.
P. domestica L.
P. damascena Dierb.
P. communis Huds.
P. domestica var. insititia
(L.) Fiori & Paoletti
P. domestica insititia
Fiori & Paoletti
P. dulcis (P. Mill) D.A.
Webber
Prunus amygdalus Batsch
Amygdalus dulcis P. Mill.
P. communis (L.) Arcang.
Amygdalus communis L.
P. emarginata (Dougl. ex
Hook.) D. Dietr.
P. mollis Walpers
Cerasus prunifolia Greene
P. fasciculata (Torr.) Gray
P. fremontii S.Wats
P. gracilis Engelm. & Gray
P. hortulana Bailey
P. ilicifolia (Nutt. ex
Hook & Arn.) D. Dietr.
apricot
Tree
mazzard cherry,
sweet cherry, gean,*
bird cherry*
Carolina laurel
cherry, wild orange
myrobalan plum,*
cherry plum,
marianna plum,
Tree
Tree
(evergreen)
Tree
flowering plum
sour cherry,
pie cherry
garden plum, plum,
European plum
Tree
Tree
bullace plum,
damson, damson plum
Tree or shrub
almond
Tree
bitter cherry,
wild cherry,
narrowleaf cherry
Shrub or tree
desert almond
desert apricot
Oklahoma plum
hortulan plum
Shrub
Shrub
Shrub
Tree
Tree or shrub
(evergreen)
California to Utah
S California
Arkansas to Texas
S Indiana to Iowa, Oklahoma,
Arkansas, Alabama & W Tennessee
Pacific Coast region, central to S California
& in N Lower California, & Mexico
have large stones and little flesh, while the pits are smaller
and there is more flesh with the opposite extremes in environment. The weights of black cherry seeds increase with
latitude (Pitcher 1984), ranging from 7 g in Florida to 14 g
in northern Michigan. There is a significant negative correlation (r = 0.35) between seed size and germination in that
smaller seeds have better germination (Pitcher 1984).
Table 1Prunus, cherry, peach, and plum: nomenclature, growth habit, and occurrence (continued)
Common name(s)
Growth habit
Occurrence
P. laurocerasus L.
laurel cherry,
cherry-laurel
mahaleb cherry,
mahaleb, St. Lucie
cherry, perfumed cherry
beach plum
wildgoose plum,
Munson plum
European bird
cherry, mayday tree
Tree
(evergreen)
Tree
SE Europe, SW Asia
Tree or shrub
Tree
sand cherry
Shrub
Bessey cherry,
western sand cherry,
Rocky Mountain cherry
Shrub
Tree
sloe, blackthorn
Shrub or tree
Klamath plum,
Pacific plum, Sierra
plum, western plum
Manchu cherry,
Nanking cherry
downy cherry
hog plum
Tree or shrub
P. mahaleb L.
Cerasus mahaleb Mill.
P. maritima Marsh.
P. munsoniana W.Wight
& Hedrick
P. padus L.
P. racemosa Lam.
Padus racemosa (Lam.) Schneid.
Cerasus padus (L.) DC.
P. pensylvanica L. f.
P. persicifolia Desf.
P. montana Marsh.
P. lanceolata Willd.
P. persica (L.) Batsch
Amygdalus persica L.
Persica vulgaris Mill.
P. pumila L.
P. depressa Pursh
P. pumila var. besseyi
Bailey (Gleason)
P. prunella Daniels
P. pumila besseyi (Bailey) Waugh.
P. susquehanae Willd.
Cerasus canadensis Mill.
P. serotina Ehrh.
P. virginiana L.
Padus virginiana (L.) Mill.
Padus serotina Borkh.
P. spinosa L.
P. subcordata Benth.
P. tomentosa Thunb.
P. trichocarpa Bge.
Cerasus tomentosa Wall.
P. umbellata Ell.
P. virginiana L.
P. nana DuRoi
P. demissa (Nutt.) D. Dietr.
Padus nana (DuRoi) Borkh.
common choke
cherry
Shrub
Tree or shrub
Tree
Shrub
Tree
Tree or shrub
intermediate (Tukey 1927). The final stage of fruit development is the rapid growth of the pericarp, and in early ripening cultivars of these species and peach, this stage begins
before the embryo reaches full size (Tukey 1936). Garden
plum also shows wide variation in germination capacity
among cultivars tested under identical conditions (Suszka
1967). Immature embryos have been brought to a germinable stage by (1) growing excised embryos in artificial
Prunus
877
culture, (2) storing whole fruit (Hesse and Kester 1955), and
(3) not picking until the fruit is over-mature (Zielinski
1958). Current technology allows the successful culture of
ovules as small as 0.6 mm (Janick and Moore 1996). Only
32% of embryos <10 mm long produce plants, compared to
78% for larger embryos (Ramming 1990).
Non-viability of apparently normal seeds derived from
crossbreeding cherries or plums has been a problem
(Cochran and others 1961). The Duke cherrieshybrids of
mazzard cherry and sour cherryoften have empty seeds
(Hedrick 1911).
Flowering and fruit-ripening dates vary among cultivars
of a species grown in the same location (Hedrick 1911,
1915; Kester 1969). Individual trees of black cherry vary in
a similar manner (Grisez 1974), and the same variation can
be expected in other wild species. The food quality of fruit
varies greatly among wild plants of Bessey cherry and the
plums. Selections of 15 species of plums have been grown
under cultivation for their fruit (Hedrick 1911, 1915).
American plum seeds from northern Minnesota germinate much better at a temperature of 10 C than at higher
temperatures, whereas those from Nebraska germinated as
well and more rapidly at 21 C (night) to 27 C (day)
(Grisez 1974).
In apricots, the variety called Russian apricot is hardier
than the typical form (Grisez 1974). Mazzard cherry cultivars require 5 to 6 days longer to begin germination in stratification than wild mazzard cherry (Suszka 1967).
In a provenance study of black cherry, Cech and
Kitzmiller (1968) found that the pattern of variation for seed
traits is random throughout most of its range. However,
seeds from the southern and southwestern parts of the range
in the United States were characteristically lighter in weight
and smaller in diameter as well as having thinner endocarps
than seeds from other areas. Geographic locations and mother trees contributed about equally to the variability in total
germination.
Flowering and fruiting. The flowers of nearly all
species are bisexual. They normally have 5 white or pink
petals and 15 to 20 or more stamens. In general, the pistil
matures 3 or 4 days before the stamens (Hedrick 1915). The
flowers are solitary, in umbel-like clusters or racemes, and
usually appear before or with the leaves. The flowers are
insect-pollinated. Except for plums and sweet cherries, most
species are self-fertile and thus a tree will set fruit without a
cross-pollinator (Janick and Moore 1996).
Of the 2 ovules, only 1 normally develops, resulting in
a 1-seeded drupe. The drupe is thick and fleshy (except in
the almonds) and has a hard, bony endocarp surrounding the
seed itself (figures 13) (Fernald 1950; LHBH 1976; Kester
878
1969; Knuth 190609; Rehder 1940). The developed endocarp and seed are commonly called the stone or pit. Dates of
flowering and fruiting are listed in table 2. Seeds are distributed mainly by birds and mammals (Grisez 1974). Fruit
diameters of most species are between 5 and 25 mm, but
Figure 1Prunus, cherry, peach, plum: seeds of P. americana, American plum (upper left), P. angustifolia, Chickasaw
plum (upper middle); P. armenica, apricot (upper right);
and P. persica, peach (bottom).
Table 2Prunus, cherry, peach, and plum: phenology of flowering and fruiting
Species
Location
Flowering
Fruit ripening
Seed dispersal
P. alleghaniensis
P. americana
P. angustifolia
P. armeniaca
Scattered
California
USSR
NE US
SE US
Geneva, New York
USSR
SE US
Nebraska
Central Mississippi
N Pennsylvania
USSR
Cheyenne,Wyoming
Bismarck, N Dakota
SE United States
E US
Warren Co., Pennsylvania
California
Late AprMay
MarMay
MarApr
FebMar
MarApr
AprMay
MarApr
May 12
AprMay
AprMay
May 818
May 1221
May
Late AprMay
May 1621
Mid FebMar
AprJune
MarMay
MarMay
AprMay
AprMay
AprJune
MarMay
May 2024
End Aprearly May
Mayearly June
Late Marearly July
May 115
AprMay
FebApr
MayJuly
AprMay
Early Apr
Late Mayearly June
Late AprJune 10
AprMay
MarMay
Early May
May 1015
MarApr
Late Aprearly June
May 1020
AugSept
JuneOct.
MayJuly
MayJune
July
JuneJuly
SeptOct
July 15Aug 10
Aug
JuneJuly
JuneJuly
July 15Oct 1
Aug
AugSept
Aug 20Oct 1
Late AugOct*
JulySept
AugOct
SeptOct*
JulyAug
July
SeptOct
JulySept*
July 15Sept 10
Late JuneJuly
JuneAug
JulySept
Late Julyearly Aug
JulySept
MayAug
JulySept
JulySept
JuneJuly
Late AugSept
JuneSept.
AugSept
AugSept
Late July
July 1015
AugSept
JulyOct
Early Aug
AugSept*
JuneOct
MayJuly
JulyAug*
AugSept*
OctDec
Aug*
JulySept
July
Aug 20Sept; rarely Nov
July 1Sept
Sept
Early Aug
July 15Sept 1
P. avium
P. caroliniana
P. cerasifera
P. cerasus
P. domestica
P. d. var. insititia
P. dulcis
P. emarginata
P. hortulana
P. ilicifolia
P. laurocerasus
P. mahaleb
P. maritima
P. munsoniana
P. padus
P. pensylvanica
P. persica
P. pumila
P. pumila. var. besseyi
P. serotina
P. spinosa
P. subcordata
P. tomentosa
P. umbellata
P. virginiana
Sources: Altman and Dittmer (1962), Bailey (1976), Bonner (1975), Fernald (1950), Grisez (1974), Hedrick (1911), Hedrick (1915), Hitchcock and others (1961), Kester
(1969), Koreisho and Morozov (1955), Long (1923), McMinn (1959), Mirov and Kraebel (1937, 1939), Munz and Keck (1959), Pane (1966), Petrides (1958), Radford and others (1964), Rehder (1940), Sudworth (1908),Van Dersal (1938).
* Collecting dates.
Average dates of height of bloom for one to several cultivars.
Prunus
879
880
Table 3Prunus, cherry, peach, and plum: height, seed-bearing age, seedcrop frequency, fruit color, and fruit size
Species
Height at
Min seedYears
maturity Year first bearing between large
(m)
cultivated age (yrs)
crops
P. alleghaniensis
P. americana
P. angustifolia
P. armeniaca
P. avium
4.9
39
4.37.6
10.4
930.5
P. caroliniana
P. cerasifera
P. cerasus
P. domestica
P. d. var. insititia
P. dulcis
P. emarginata
P. fasciculata
P. fremontii
P. gracilis
P. hortulana
P. ilicifolia
P. laurocerasus
P. mahaleb
P. maritima
P. munsoniana
P. padus
P. pensylvanica
P. persica
P. pumila
P.var. besseyi
P. serotina
P. spinosa
P. subcordata
P. tomentosa
P. umbellata
P. virginiana
5.512
8.2
915
912
67.6
39
115
12.4
1.53.7
4.6
9
7.69
5.5
610
3
69
15
312
37.6
0.32.4
0.31.2
15.333.5
4
37.6
1.83
1.89
1889
1768
~ 1874
Early
Early
4
4
2
5
67*
1
12
2
1
Early
Early
Early
Early
Early
1918
Pre-1925
Early
Pre-1909
Early
1773
Early
1756
1892
1629
Early
~ 1850
1870
1724
3
6-7*
5
4,67
3
3
3
3
3
2
3
23
5
23
3
23
1
12
12
15
12
2
12
Yellowish
Red
Red to yellow
Purple or black
Purple to black
Black
Purple
Red or yellow
Black in typical variety
Light red
Yellow to red
Purple-black
Purple to black
Black
Blue-black
Red or yellow
Red
Black, red, yellow
Red-purple to dark purple
1013
1625
825
30+
25+
30+
812
13
1520
13
25
1317
10
810
1325
2030
68
57
3060
10
15
710
1015
2030
1031
1015
8
1013
1520
3060
13
25
813
610
20
1525
3075
10
610
15
1530
15
1013
Sources: Bailey (1976), Everett (1957), Fernald (1950), Giersbach and Crocker (1932), Grisez (1974), Gysel and Lemien (1964), Hedrick (1911, 1915), Huntzinger (1971),
Kester (1969), Koreisho and Morozov (1955), Munz and Keck (1959), Peck (1961), Petrides (1958), Rehder (1940), Strausbaugh and Core (1964), Van Dersal (1938).
* Minimum commercial seed-bearing age.
Ages are for seedling stock; grafted or budded stocks bear seeds 1 or 2 years younger (Wright 1966).
ly requires only a few hours (Huntzinger 1968). The moisture content of black cherry seeds has been reduced from
about 14% to 5% by drying at 32 C for 3 hours
(Huntzinger 1971).
Sealed containers are preferred for Prunus seeds if the
moisture content is to be closely controlled. Seeds of mazzard cherry were dried to a moisture content of 11% and
stored in sealed bottles at 1 C for 4 1/2 years. During this
period, viability decreased from 93% to 84% (Suszka 1970).
Plastic bags have been satisfactory for storage of black
cherry seeds for at least 3 years at cold temperatures
(Huntzinger 1971). Cloth sacks may serve for short periods
in cool temperatures (Suszka 1967). Maheleb cherry and
Prunus
881
Species
Seed weight/45 kg
(100 lb) of fruit
Range
Average
kg
lb
kg
lb
P. alleghaniensis
P. americana
315
734
P. angustifolia
414
830
P. armeniaca
USA
1418
3040
USSR
57
1015
P. avium
USA
311
725
USSR
78
1518
P. cerasifera
5
P. cerasus
P. domestica
P. d. var. insititia
P. dulcis
P. emarginata
P. ilicifolia
P. mahaleb
911
2025
P. munsoniana
P. padus
P. pensylvanica
712
1627
P. persica
P. pumila
typical
var. besseyi
713 1528
10
P. serotina
fresh seeds
P. subcordata
P. tomentosa
35
712
P. virginiana
811
1825
Range
/kg
Cleaned seeds/weight
Average
/lb
/kg
/lb Samples
9
7
19
16
5501,500
7701,530
250680
350694
2,950
870
1,030
338
395
467
1
27+
14+
200560
270495
91254
123225
317
382
144
173
10+
10
9
5
3
11
12
1,4503,000
1,6402,770
7821,330
1,5104,000
416907
6251,920
126225
4,1208,790
200240
4,8005,600
9002,240
6,60012,300
8,00021,800
72244
6581,361
3,6166,108
355603
6851,815
189411
284871
57102
1,8693,987
91109
2,1772,540
4081,016
2,9945,580
3,6299,889
33111
2,360
4,740
994
2,910
597
1,380
181
7,020
220
5,200
1,690
8,910
14,200
156
1,070
2,150
451
1,320
271
626
82
3,184
100
2,359
767
4,042
6,442
71
9+
7+
6+
5+
3+
3+
6+
2+
3+
5+
6+
6+
20
10
7
25
20
20
21
2,4604,000
1,5004,000
1,1161,815
6811,815
2,920
2,400
1,325
1,090
4+
10+
9
10
5
5
9
20
21
10
10
20
2,8006,040
2,84013,800
1,9702,670
450631
1,7306,400
3,0108,400
1,2702,740
1,2886,260
8941,211
204286
7852,903
1,3153,810
4,240
5,370
2,240
556
4,740
4,790
1,923
2,436
1,016
252
2,150
2,173
68
197
4+
9+
19
Sources: Benjdl (1954), Cech and Kitzmiller (1968), Chittenden (1927), Cumming and others (1933), Defler (1937), Engstrom and Stoeckeler (1941), Everett (1957),
Glazebrook (1941), Grisez (1974), Huntzinger (1971), King (1947), Koreisho and Morozov (1955), Krefting and Roe (1949), Krier (1948), Mirov and Kraebel (1937, 1939),
Swingle (1939), USDA (1961),Van Dersal (1938).
myrobalan plum can be stored up to 2 winters at room temperature in jute sacks without loss of viability (Grzeskowiak
and others 1983). Pin cherry seeds retained high viability
after 10 years of storage at 1 to 3 C under sealed conditions
(Dirr and Heuser 1987).
Normally, storage temperatures should be within the
range 0.6 to 5 C, although American plum, mazzard cherry,
and mahaleb cherry have been successfully stored at room
temperatures for 2 to 5 years. American plum seeds can be
stored at room temperature up to 30 months without loss of
germinative capacity (Giersbach and Crocker 1932).
Manchu cherry seeds stored for 21 months at room temperature did not lose viability (Dirr and Heuser 1987). Dried
seeds of mazzard and mahaleb cherries and myrobalan plum
can be stored up to 3 winters at 1 or 3 C without signifi-
882
Tables 5Prunus, cherry, peach, and plum: germination of seeds after dry storage*
Species & storage period
P. americana
18 months
53 months
18 months
53 months
18 months
53 months
P. avium
7 years
15 years
55 months
55 months
8-12 months
207 days
214 days
570 days
571 days
935 days
213 days
214 days
568 days
P. pensylvanica
2 months
6 years
10 years
P. serotina
1 year
2 years
3 years
5 years
8 years
1 year
2 years
3 years
5 years
8 years
1 year
2 years
3 years
5 years
8 years
1 year
2 years
3 years
5 years
8 years
18
18
18
18
18
18
18
18
18
18
P
Germination (%)
70
45
72
16
62
0
5
5
1
1
3
3
3
3
3
1
3
3
3
~10?
~10?
11
11
911
8.6
9.0
8.6
9.5
8.3
9.0
8.9
8.6
9197
98
84
88
98100
100*
99
100
94
99
99
98
100
18
13
13
Dry
Low
Low
95
74
76
to 14
to 14
to 14
to 14
to 14
to 14
to 14
to 14
to 14
to 14
0.55
0.55
0.55
0.55
0.55
0.55
0.55
0.55
0.55
0.55
46
46
46
46
46
1113
1113
1113
1113
1113
46
46
46
46
46
1113
1113
1113
1113
1113
52
81
81
47
66
4
7
1
4
0
63
81
90
69
56
72
88
77
0
0
Sources: Ellis and Hong (1986), Giersbach and Crocker (1932), Grisez (1976), Heit (1967), Huntzinger (1971), Laidlaw (1983), Michalska and Suszka (1980c&d), Solovieva
(1966, 1978), Suszka (1970).
Viability determined by indigo carmine embryo-staining test (2 hours in 0.05% solution at 20 C).
883
884
Table 6Prunus, cherry, peach, and plum: stratification periods, germination test conditions, and results
Species
P. alleghaniensis
P. americana
P. angustifolia
P. armeniaca
endocarp removed
endocarp intact
endocarp intact
P. avium
endocarp removed
endocarp intact
endocarp intact
P. caroliana
P. cerasifera
P. cerasus
P. domestica
P. domestica var. insititia
P. dulcis
P. emarginata
P. ilicifolia
fresh seed
stored seed
P. laurocerasus
P. mahaleb
P. maritima
P. munsoniana
P. padus
fresh seeds
stored seeds
P. pensylvanica
P. persica
endocarp intact
endocarp removed
P. pumila var. besseyi
P. serotina
P. spinosa
P. subcordata
P. tomentosa
P. virginiana
Recommended
stratification (days)
Warm*
Cold
Avg
germination
Viability
(%)
Samples
(%)
0
0
0
150
90150
60120
10
10
10
10
60
60
60
25
60
55
7
21
74
90
0
14
0
0
189
8090
3
5
7
3
5
14
95
95
90
3
4
0
0
14
0
0
14
0
0
0
0
0
90125
120180
189
3060
196
90150
189
120150
90
84112
65
90126
21
21
3
2
18
2
24
21
21
3
2
18
2
24
28
60
91
76
88
65
56
89
10+
2
82
15
85
91
7
90
3+
69
0
0
0
0
14
0
0
0
90
6090
80100
189
90
80100
89
55
100
24
5
3
39
10
0
14
60
100120
210
90
3
25
3
10
60
85
50
62
3
2
91
0
0
0
0
14
0
0
0
0
98105
70105
120
120
189
170
90
6090
120160
5(41)
5(41)
26
3
25
5(41)
5(41)
10
3
10
4060
100
40
32
82
60
86
90
1
11
77
8
8
72
32
3
90
92
80
86
62
Sources: Afanasiev (1940, 1942), Albenshkii and Nikitin (1956), Chadwick (1935), Chao and Walker (1966), Coe and Gerber (934) Crocker (1927, 1931), Defler (1937),
Dirr and Heuser (1987), Emery (1964), Engstrom and Stoeckeler (1941), Everett (1957), Fogle (1958), Fogle and McCrory (1960), Glazebrook (1941), Giersbach and
Crocker (1932), Grisez (1974), Haut (1932, 1938), Havis and Gilkeson (1949), Hesse and Kester (1955), Heit (1938), Kester (1969), Koreisho and Morozov (1955), Krefting
and Roe (1949), Morov and Kraebel (1937), Pollock (1959), Probocskal (1963), Roe (1941), Suszka (1964, 1967), Swingle (1939),Tukey (1924), USDA (1961).
* Seeds were in a moist medium at a constant temperature of 20 C or at a temperature alternating diurnally from 30 C (8 hours) to 20 C (16 hours).
Seeds were in a moist medium at a temperature between 0.6 C and 5 C; 2.8 to 5 C was better.
Germination occurred during the stratification period.
Results were similar at 10 C.
II Adequate germination was reported at unspecified temperatures.
Prunus
885
886
Prunus
887
medium or large sized fruits, but plants with much branching, very thin shoots, and small leaves are likely to have
small fruit and fruit at an older age (Janick and Moore
1996). Nursery practices that have been successful are listed
in table 7.
Species
P. americana
P. angustifolia
P. armeniaca
P. avium
P. cerasifera
P. cerasus
P. domestica
P. mahaleb
P. padus
P. persica
P. pumila var. besseyi
P. serotina
P. spinosa
P. tomentosa
P. virginiana
120
1520
90
120
90
60
120
85
120
120
170
60
120160
Seeds
sown/ft2
4
1
9
13
10
21
13
50
0.75
67
1020
1728
1530
25
Sowing
depth (in)
12
33
2
12
2
2
12
1/21ll
2
1 1/22ll
12
1ll
_
Tree %
3350
1
50
64
45
77
783
7075
72
334
Outplanting
age (yr)
1
1
1
1
1
1
1
or 2
1
or 2
or 2
or 2
1
or 2
1
or 2
Sources: Afanasijev (1962), Albenskii and Nikitin (1956), Bailey (1969), Bejdl (1954), Engstrom and Stoeckeler (1941), Grisez (1974), Heit (1938, 1967), Huntzinger (1971),
Koreisho and Morozov (1955), Nyholm (1951), Rudolf (1961), Schaaf (1938, 1940), Shoemaker and Teskey (1959), Shumilina (1940, 1949), Stoeckeler and Jones (1957),
Swingle (1939),Talbert (1946).
* Stratified in a moist medium at a temperature between 2.8 and 5 C.
Add a 10 to 15 cm (4- to 6-in) soil ridge to the nurserybed.
Or stratify from time of collection to time of sowing when fresh seeds are used.
Germination percent (not tree percent).
ll On fall-sown beds, add ~8 cm (3 in) of straw or moss for a mulch.
References
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trees. Oklahoma Agricultural Experiment Station Biennial Report
193840: 124126.
Afanasiev M. 1942. Propagation of trees and shrubs by seed. Circ. C-106.
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Afanasijev D. 1962. Seed stimulation [translated from Serbo-Croatian].
Washington, DC: National Science Foundation and USDA. 15 p.
Albenskii AV, Nikitin PD, eds. 1956. Agrolesomeliortsiya. 3rd ed. Gos. Izd. Skh. Lit., Moskva. [Handbook of afforestation and soil amelioration.Transl.
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AOSA [Association of Official Seed Analysts]. 1996. Rules for testing seeds.
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Altman PL, Dittmer DS. 1962. Biological handbook on growth. Washington,
DC: FASEB [Federation of American Societies for Experimental Biology].
608 p.
Bejdl R. 1954. Tresen, cilova drevina blizke budoucnosti. Lesnicka Prace 33:
354357 [Prunus avium, the tree of the future. Forestry Abstracts 16:
199].
Heit CE. 1967a. Propagation from seed: 6. Hard seedednessa critical factor. American Nurseryman 125(10): 1012, 8896.
Heit CE. 1967b. Propagation from seed: 8. Fall planting of fruit and hardwood seeds. American Nurseryman 126(4): 1213, 8590.
Heit CE. 1967c. Propagation from seed: 11. Storage of deciduous tree and
shrub seeds. American Nurseryman 126(10): 1213, 8694.
Hesse CO, Kester DE. 1955. Germination of embryos of Prunus related to
degree of embryo development and method of handling. Proceedings of
the American Society of Horticultural Science 65: 251264.
Hitchcock CL, Cronquist A, Ownbey M,Thompson JW. 1961. Vascular
plants of the Pacific Northwest: 3. Saxifragaceae to Ericaceae. Seattle:
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Huffman GR. 1996. Seed handling and propagation of hardwood trees and
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Conservation Nursery Associations. Gen.Tech. Rep. PNW-389.Portland,
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Huntzinger HJ. 1968. Methods for handling black cherry seed. Res. Pap. NE102. Upper Darby, PA: USDA Forest Service, Northeastern Forest
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Huntzinger HJ. 1971. Long-term storage of black cherry seedis it effective? Tree Planters Notes 22(4): 34.
ISTA [International Seed Testing Association]. 1996. International rules for
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Janick J, Moore JN. 1996. Fruit breeding.Volume 1,Tree and tropical fruits.
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Jones L. 1963. Effect of various pregermination treatments on germination
of black cherry seed. Res. Note SE-8. Asheville, NC: USDA Forest
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Kester DE. 1969. Almonds. In: Handbook of North American nut trees.
Knoxville,TN: Northern Nut Growers Association: 302314.
King JE. 1947. The effect of various treatments to induce germination of
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Knuth P. 19061909. Handbook of flower pollination. 3 vol. Oxford:
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germination. Ecology Monographs 19: 269286.
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Lockley GC. 1980. Germination of chokecherry (Prunus virginiana) seeds.
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Long B. 1923. Naturalized occurrence of Prunus padus in America. Rhodora
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McMinn HE. 1959. An illustrated manual of California shrubs. Berkeley:
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Michalska S, Suszka B. 1980c. Testing of the effectivness of a warm-followed-by-cold stratification of Prunus avium L. seeds, with a 3-week
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Prunus
889
890
Shumilina ZK. 1949. Podgotovka posevu semyan drevesnykh i kustarnikovykh porod.Vses. Nauchnoissled Inst. Agrolesomelior.,
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PinaceaePine family
Pseudotsuga Carr.
Douglas-fir
William I. Stein and Peyton W. Owston
Dr. Stein is a forest ecologist emeritus at the USDA Forest Services Pacific Northwest Research Station,
Corvallis, Oregon; Dr. Owston retired from USDA Forest Services Pacific Northwest Research Station
Common name(s)
Occurrence
bigcone Douglas-fir,
bigcone-spruce, desert-fir
Mtns of SW California
interior Douglas-fir,
blue Douglas-fir, Rocky
Mountain Douglas-fir,
Colorado Douglas-fir,
inland Douglas-fir
coastal Douglas-fir,
green Douglas-fir,
Oregon Douglas-fir,
Douglas-fir,
Douglas-spruce
Sources: Griffin and Critchfield (1976), Hermann (1982), Hermann and Lavender (1990), Little (1952, 1971, 1979), McDonald (1990), Minnich (1982).
Pseudotsuga
891
892
insect and disease resistance, cold hardiness, wood characteristics, and chemical composition (Campbell 1986;
Campbell and Sorensen 1978; Campbell and Sugano 1993;
Ching and Bever 1960; Griffin and Ching 1977; Joly and
others 1989; Li and Adams 1989; Read and Sprackling
1976; Rehfeldt 1979, 1989; Schowalter 1988; Silen 1978;
Sorensen 1983; St. Clair and Adams 1991; Strauss and Tsai
1988). Because the survival and growth of Douglas-fir at a
given location varies by the genetic source used, much effort
has been expended in defining guidelines and zones for
informed use of seeds and stock beyond the local geographic and climatic area of origin (Campbell 1991; Kleinschmit
and Bastien 1992; Randall 1996; Rehfeldt 1981, 1983a&b.)
Inspection systems have been developed to ensure that
Douglas-fir seedlots and stock are correctly labeled and their
origins certified (Portlock 1992; Schrumpf and Pfeifer 1993.)
Efforts to genetically improve Douglas-fir began over 50
years ago (Isaac 1949) and have developed into large longterm cooperative programs in western North America and
abroad. In the Pacific Northwest, most of the seeds used in
reforestation of coastal Douglas-fir now come from seed
orchards (Daniels 1995). The species use in the breeding
zone of origin and other breeding zones is guided by results
of outplanting tests and general rules for seed transfer
(Randall 1996). A 10% gain in juvenile height growth is
predicted for selections representing 6 low-elevation breeding zones in western Oregon and Washington (Stonecypher
and others 1996). An international program for genetic
improvement of Douglas-fir was started in 1967 and is continuing under IUFRO agencies involving 59 institutions in
36 countries (Kleinschmit and Bastien 1992). Other genetic
improvement efforts involve selection for Christmas trees
(Douglass and TerBush 1975; Silen 1978) hybridization
(Rehfeldt 1977), and clonal propagation (Silen 1978). Many
cultivars of Douglas-fir have been propagated by horticulturists (Hermann 1982).
Flowering and fruiting. Male and female strobili
burst bud during late winter and spring (table 2), about a
year after their initiation as axillary bud primordia (Allen
and Owens 1972). Male strobili are generally borne abundantly over much of the crown on the proximal half of yearold shoots; these strobili become somewhat pendant when
mature and are about 2 cm long. Female strobili develop
more distally on year-old shoots that are located primarily in
the upper half of the crown. The female strobili are erect at
the time of pollen shedding and measure about 3 cm long;
their appearance is dominated by large trident bracts (Allen
and Owens 1972). The color of female strobili (seed cones)
ranges from deep green to deep red, and that of male strobili
Species
Location
Flowering
Cone ripening
Seed dispersal
P. macrocarpa
P. menziesii var. glauca
S California
Central Colorado
(elev. 2,0602,880 m)
Montana
(elev. 7001,700 m)
Northern Idaho
(elev. 8201,000 m)
Central Oregon
Coastal British Columbia
W-central Oregon &
W Washington
S Oregon
N California
Februarymid-Apr
Mid-Aprearly May
Late Augmid-Sept*
Mid-Aug
Early Sept*
Mid-Maymid-June
Late Marmid-May
Mid-Marearly June
Aug
Aug
Mid-Augmid-Sept*
Septlate Mar
Late Auglate Mar
August
Mid-Aug*
Septearly winter
Sources: Allen (1942), Allen and Owens (1972), Ching and Ching (1962), Gashwiler (1969), Gause (1966), Griffith (1968), Isaac (1943), McDonald (1990, 1992), Morris
(1952), Owston and Stein (1974), Roeser (1942), Silen (1963), Sorensen and Campbell (1971), Sudworth (1908).
* Beginning dates only.
Upslope progression in pollen shedding and female receptivity averaged 23.5 m of elevation per day on 4 transects in western Oregon and Washington (Silen 1963).
About 70 to 90% of the seeds usually disperse in September and October; most of the remainder disperse between November and March (Allen 1942; Isaac 1943).
Seedfall usually greatest in October (McDonald 1992).
cones (Willis 1917). In interior Douglas-fir, filled-seed numbers ranged from 20 to 30 per cone in some collections
(Owston and Stein 1974). For coastal Douglas-fir, filledseed numbers ranged from 4 to 54 per cone in 309 collections from individual 35-year-old trees in northwestern
Oregon (Olson and Silen 1975). Numbers averaged 16 per
cone in collections from 127 individual trees in 10 westside
areas located from north-central Washington to south-central
Oregon (Willis and Hofmann 1915) and 19.4 per cone
(range 2 to 35) in collections made between 1966 and 1978
Pseudotsuga .
893
894
Table 3Pseudotsuga, Douglas-fir: height, seed-bearing age, crop frequency, and cone length
Species
P. macrocarpa
P. menziesii
var. glauca
var. menziesii
Mature
height (m)
Minimum
seed-bearing
age (yr)
Years
between large
seedcrops
P
Cone length
(cm)
930
20*
1117
2349
2791
20
710
310
211
47
610
Sources: Allen (1942), Boe (1954), Gause (1966), Hermann and Lavender (1990), Lowry (1966), McArdle and others (1961), McDonald (1990), Roeser (1942), Sudworth
(1908).
* Occasionally earlier on good, open sites (Gause 1966).
The minimum age for commercial collections has been considered 20 to 25 years (Allen 1942).
Pseudotsuga
895
896
Table 4 Pseudotsuga, Douglas-fir: cone drying schedules and seed yield data
Species
P. marocarpa
P. menziesii
var. glauca
var. menziesii
Seed yield
Seed wt/cone vol
Ratio seed wt/
kg/hl
lb/bu
100 cone wt
810
3239
2530
1.03
0.8
2.8
460
860
210
1648
3843
3243
3277
3964
2560
3050
0.651.03
0.261.03
0.50.8
0.20.8
1.01.3
0.52.0
Sources: Brown (1983), Owston and Stein (1974), Radcliffe (1952), Swingle (1939).
Pseudotsuga
897
Range
/lb
Samplesa
/kg
/lb
/kg
7,145
7,716
11,001
6,748
3,241
3,500
4,990
3,061
6,1378,115
6,6149,921
9,25913,927
2,8003,681
3,0004,500
3,4606,317
2
2
3
1
70,768
83,555
85,539
88,405
102,354
97,665
119,905
32,100
37,900
38,800
40,100
46,427
44,300
54,388
52,91175,619
72,31290,830
73,41496,122
79,80799,869
101,834103,093
62,832117,506
104,166163,398
24,00034,300
32,80041,200
33,30043,600
36,20045,300
46,19146,762
28,50053,300
47,24974,116
8
8
33
14
3
19
23
86,882
70,028
71,752
67,250
39,409
31,764
32,546
30,504
80,257104,058
55,61878,493
33,951116,845
40,858109,171
36,40447,200
25,22835,604
15,40053,000
18,40949,519
4b
5c
41
20d
72,305
65,538
56,906
76,278
110,498
99,503
83,333
78,626
84,336
86,589
87,268
81,183
77,499
116,822
86,999
130,891
93,584
32,797
30,181
25,812
34,599
50,121
45,134
37,799
35,664
38,254
39,276
39,584
36,824
35,153
52,990
39,462
59,371
42,449
63,26893,268
54,78081,359
48,02869,867
58,34399,109
84,104126,263
69,589182,150
62,501109,409
59,00098,000
77,162125,664
73,634102,458
76,128110,972
75,07487,336
74,99979,999
82,576153,845
65,00199,999
107,411173,012
79,807120,593
28,69842,605
24,84836,904
21,78531,691
26,46444,955
38,14957,272
31,56582,622
28,35049,627
26,76244,452
35,00057,000
33,40046,474
34,53150,336
34,05339,615
34, 01936,287
37,45669,783
29,48445,359
48,72178,749
36,20054,700
62
29
37
8d
10
309e
39
8f
127g
97
23
2d
2f
46
4f
21
16
Sources: Allen (1942), Bialobok and Mejnartowicz (1970), Ching and Bever (1960), Griffin and Ching (1977), Heit (1968), Lippitt (1996), Olson and Silen (1975), Owston
and Stein (1974), Rafn (1915), Randall (1997), St. Clair and Adams (1991), Sweet (1965),Willis and Hofmann (1915).
a Data represent seedlots of unknown size and number of trees unless otherwise noted.
b Four locations representing 41 elevation points (stands) totaling 87 trees.
c Five locations representing 44 elevations points (stands) totaling 94 trees.
d Each provenance (sample) represented by 10 trees under 50 years of age.
e Collected over the entire season (August 12 to September 15) from individual 35-year-old trees growing at 122 to 518 m of elevation.
f Each sample collected from 14 to 89 trees within a 40-km radius.
g From individual open-pollinated parent trees.
and some that need it are harmed by too lengthy stratification (Allen 1960; Allen and Bientjes 1954; Gosling 1988;
Jensen and Noll 1959; Taylor and others 1993). Based on
accumulated experience, seeds of coastal Douglas-fir generally require or benefit from stratification, those of interior
Douglas-fir from northern sources may benefit, those from
southern sources may not benefit, and the response to stratification by bigcone Douglas-fir is unknown (Allen 1960,
1962c; Heit 1968).
898
Stratification overcomes different degrees of seed dormancy that may be related to seed source and family, year of
collection, cone and seed drying conditions, length and kind
of storage, and other causes (Allen 1960; Allen and Bientjes
1954; El-Kassaby and others 1992; Jensen and Noll 1959;
Sorensen 1991). Several pretreatments used with or in lieu
of stratification also stimulate germination or reduce
pathogen damage. These include a light rinse with cold or
hot water, fungicide, bleach, hydrogen peroxide, ethanol,
polyethylene glycol, or ethylene (Borno and Taylor 1975;
Species
P. macrocarpa
P. menziesii
var. glauca
P. menziesii
var. menziesii
Stratification*
(days)
28
21
0
30
0
2040
0
21
0
2040
0
28
Germination conditions
Moist
Temp (C)
medium
Day Night
Vermiculite
Sand
Sponge Rok
Sponge Rok
Paper
Paper
Vermiculite
Vermiculite
Sponge Rok
Sponge Rok
Vermiculite
Vermiculite
30
27
30
30
30
30
25
25
30
30
25
25
30
20
21
20
20
20
20
25
25
20
20
25
25
20
Days
28
100
2090
1421
2135
17
29
30
50
1435
2835
30
50
28
Germinative energy
Amt
Time
Germ. capacity (%)
(%)
(days)
Avg
Range
Samples
14
60
40
70
76
55
54
60
9
12
10
9
10
17
31
28
30
68
60
78
84
95
93
81
75
87
84
84
1436
1557
2483
2775
86100
8898
3895
2993
34100
42100
6698
3
3
5
8
8
3
3
6
6
194
194
20
20
129
Sources: Allen (1962c), Lippitt (1996), Owston and Stein (1974), Rafn (1915).
* Seeds stratified naked (Allen and Bientjes 1954) or on moist Sponge Rok, vermiculite, or paper at 0 to 5 C.
Alternating temperatures included 8 hours of light during the high temperature period; light apparently was provided with constant temperature.
Var. glauca from north-central Colorado seed sources.
Pseudotsuga 899
900
stored at 7 to 3 C for 9 months or more retained the viability and stratification effect (Allen 1962b; Belcher 1982;
Danielson and Tanaka 1978; Jones and Gosling 1994;
Muller and others 1999). In other trials, viability was
retained by partial drying of stratified seeds followed by low
temperature storage, but the stratification effect was lost
(Hedderwick 1970; Malavasi and others 1985). Seeds cannot
be held in stratification indefinitely; seeds of some lots will
deteriorate and those of others will germinate (Allen 1960;
Danielson and Tanaka 1978; Sorensen 1980, 1991).
For bareroot production, most Douglas-fir seeds are
drill-sown at a depth of 3 to 6 mm (0.1 to 0.2 in). The larger
seeds of bigcone Douglas-fir are usually sown at a depth of
13 mm (0.5 in). Seedbed density varies depending on the
stock size desired. For 2+0 Douglas-fir stock, seedling densities vary from 161 to 323/m2 (15 to 30/ft2); for 1+1s, bed
densities of first-year seedlings range from 538 to 753/m2
(50 to 70/ft2) (Thompson 1984). Seedbed density has more
effect on size of seedlings produced, and on their subsequent
field survival and growth, than does irrigation frequency or
undercutting and wrenching (Stein 1988).
Irrigation is used in bareroot nurseries to supply moisture to seeds and seedlings, prevent overheating or frost
damage, promote growth, and augment other practices such
as fertilizing and root wrenching (Duryea 1984). Carefully
planned irrigation regimes are also used to control moisture
stress, harden seedlings, and initiate dormancy. Specific irrigation regimes should be developed for each nursery and
kind of stockseedlings can be harmed by either too much
or too little water.
The fertilizer mix and the timing and number of applications must also be developed for each bareroot nursery.
Physical and chemical characteristics of the soil and irrigation water and the density of the seedling crop are critical
influencing factors. A pH of 5.0 to 6.0 has been suggested
for nurseries in the Pacific Northwest and 5.5 to 6.5 for
nurseries in Intermountain region, as well as concomitant
target levels for key mineral elements (Youngberg 1984).
Most nurseries stop fertilizing in July or early August to
promote seedling hardening (Duryea 1984).
Almost all bareroot nurseries undercut Douglas-fir
seedlings to stimulate root growth in the upper soil layers
(Duryea 1984). Timing, frequency, and depth vary by nursery and stock type. Eighty percent of nurseries in the Pacific
Northwest also wrench the roots of Douglas-fir seedlings to
promote fibrous root systems, stress and harden seedlings,
control shoot height, and aerate the soil.
Many other cultural techniques including transplanting,
weed and pest control, top pruning, and mycorrizhal inocu-
Pseudotsuga
901
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902
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903
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Pseudotsuga
905
906
FabaceaePea family
Psorothamnus Rydb.
indigobush
Jane E. Rodgers and Carol Miller
Ms. Rodgers is stationed at the Point Reyes National Seashore, Point Reyes, California; Ms. Miller was a
propagationist at the USDI National Park Services Joshua Tree National Park
Common name(s)
Occurrence
indigobush,*
Mojave dalea
Mojave indigobush,
Saunder dalea
Johnson dalea
California dalea
dyeweed,* dyebush
Fremont dalea
Nevada dalea,
Nevada smokebush
indigobush, Schott dalea
Mojave Desert
smoketree, smokebush
Psorothamnus
907
908
Table 2Psorothamnus, indigobush: seed weight, initial and best germination, and storability of seeds
Species
Seeds/weight
/kg
/lb
P. emoryi
P. fremontii
P. polydenius
P. schottii
P. spinosus
600
35
460
22
50
275
16
210
10
23
Percentage germination
Initial
Best
58
41
2
90
22
75
97
99
88
58
Storability
Stores well
50% hard seed, stores well
90% hard seed, stores well
Good storage
1747% hard seed, stores well
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Psorothamnus 909
RutaceaeRue family
Ptelea trifoliata L.
common hoptree
Kenneth A. Brinkman, R. C. Schlesinger, and Jill R. Barbour
Drs. Brinkman and Schlesinger retired from the USDA Forest Services North Central Forest Experiment
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Dry Branch, Georgia
910
fruit
longitudinal
911
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912
FabaceaePea family
Pterocarpus Linn.
padauk, narra
John K. Francis
Dr. Francis is a research forester at the USDA Forest Services International Institute of Tropical Forestry,
Ro Piedras, Puerto
913
4 cm
914
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Gen.Tech. Rep. SO-82. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 12 p.
Francis JK, Rodrguez A. 1993. Seeds of Puerto Rican trees and shrubs: second installment. Res. Note SO-374. New Orleans: USDA Forest Service,
Southern Forest Experiment Station. 5 p.
Hundley HG. 1956. Padauk, Pterocarpus macrocarpus Kurz. Burma Forester
7(1): 638.
Pterocarpus
915
RosaceaeRose family
Common name
P. glandulosa Curran
P. tridentata var. glandulosa (Curran) M.E. Jones
P. mexicana (D. Don) Henrickson
Cowania mexicana D. Don
P. tridentata (Pursh) DC.
desert bitterbrush
cliffrose
916
antelope bitterbrush
Geographic distribution
Table 2Purshia, bitterbrush and cliffrose: seed yield data (seeds/weight) for mechanically cleaned seeds*
Mean
Species
/kg
P. glandulosa
P. mexicana
P. tridentata
Range
/lb
50,850
129,000
35,000
/kg
26,540
58,600
15,750
/lb
45,00090,000
108,000210,000
29,00051,000
20,30040,900
49,00095,000
13,40023,200
Sources: Alexander and others (1974), Belcher (1985), Deitschman and others (1974), Meyer (2002), Meyer and others (1988).
2 wk
10
6
6
2
13
56
33
64
43
60
81
83
91
88
100
93
100
94
100
98
100
65
100
32
100
36
12 wk
100
32
19
37
Samples
1*
1
6
1
13
1
Sources: Deitschman and others (1974), Meyer (2002), Meyer and Monsen (1989),Young and Evans (1981).
Note: Values are expressed as percentage of initially viable seeds after moist chilling at to 2 C for 0 to 12 weeks followed by incubation at 15 C or 10/20 C for 4
weeks.
* These seeds were chilled at 3 to 5 C and germination was scored during chilling.
Decrease in germination percentage after 6 weeks was due to seed mortality during the test.
918
Purshia
919
weeks of age are usually large enough (Alexander and others 1974; Shaw 1984). On more level terrain, a conventional
tree-planter may be used (Alexander and others 1974).
Transplanting should be carried out at a time and in such a
way as to assure that the transplants will have adequate
moisture for root development for 4 to 6 weeks after planting. This may be accomplished by planting in very early
spring or by watering at the time of planting. Fall-planted
seedlings may require supplemental watering. Controlling
competition from weedy annual or perennial grasses before
planting will enhance survival and first-season growth.
References
Alexander RR, Jorgensen K, Plummer AP. 1974. Cowania mexicana var.
stansburiana (Torr.) Jeps., cliffrose. In: Schopmeyer CS, tech. coord. Seeds
of woody plants in the United States. Agric. Handbk. 450. Washington,
DC: USDA Forest Service: 353355.
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
Austin DD, Urness PJ. 1983. Summer use of bitterbrush rangelands by
mule deer. In:Tiedemann AR, Johnson KL, comps. Proceedings, Research
and Management of Bitterbrush and Cliffrose in Western North
America; 1982 May 1315; Salt Lake City. Gen.Tech. Rep. INT-152.
Ogden, UT: USDA Forest Service, Intermountain Forest and Range
Experiment Station: 203212.
Belcher E. 1985. Handbook on seeds of browse-shrubs and forbs.Tech.
Pub. R8-8. Atlanta: USDA Forest Service, Southern Region. 246 p.
Bishop CJ, Garton EO, Unsworth JW. 2001. Bitterbrush and cheatgrass
quality on 3 southwest Idaho winter ranges. Journal of Range
Management 54:595-602.
Bond G. 1976. Observations on the root nodules of Purshia tridentata.
Proceedings of the Royal Society of London (Series B) 192: 127135.
Booth DT. 1980. Emergence of bitterbrush seedlings on land disturbed by
phosphate mining. Journal of Range Management 33: 439441.
Booth DT. 1995. Cased-hole punch seeder: a tool for increasing plant
diversity. In: Abstracts, 48th Annual Meeting. Denver: Society for Range
Management: 8.
Booth DT. 1999. Imbibition temperatures affect bitterbrush seed dormancy
and seedling vigor. Journal of Arid Environments 48: 3539.
Booth DT, Morgan DR. 1993. Post-germination growth related to time-togermination for four woody plants. Journal of Seed Technology 16:
3038.
Booth DT, Sowa S. 2001. Respiration in dormant and non-dormant
bitterbrush seeds. Journal of Arid Environments 43: 91101.
Davis KM, Englert JM, Kujawski JL. 2002. Improved conservation plant
materials released by NRCD and cooperators through September 2002.
Beltsville, MD: USDA Agricultural Research Service. 57 p.
Deitschman GH, Jorgensen KR, Plummer AP. 1974. Purshia, bitterbrush. In:
Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington DC: USDA Forest Service:
686688.
Dreyer DL and Trousdale EK. 1978. Cucurbitacins in Purshia tridentata.
Phytochemistry 17:325-326.
Evans RA,Young JA, Cluff GJ, McAdoo JK. 1983. Dynamics of antelope bitterbrush seed caches. In:Tiedemann AR, Johnson KL, comps.
Proceedings, Research and Management of Bitterbrush and Cliffrose in
Western North America. 1982 May 1315; Salt Lake City. Gen.Tech.
Rep. INT-152. Ogden, UT: USDA Forest Service, Intermountain Forest
and Range Experiment Station: 195202.
920
Everett RL, Meeuwig RO. 1975. Hydrogen peroxide and thiourea treatment of bitterbrush seed. Res. Note 196. Ogden, UT: USDA Forest
Service, Intermountain Forest and Range Experiment Station. 6 p.
Ferguson RB, Basile JV. 1967. Effect of seedling numbers on bitterbrush
survival. Journal of Range Management 20: 380382.
Geier-Hayes K. 1987. Occurrence of conifer seedlings and their microenvironments on disturbed sites in central Idaho. Res. Pap. INT-383. Ogden,
UT: USDA Forest Service, Intermountain Forest and Range Experiment
Station. 12 p.
Giunta B, Stevens R, Jorgensen K, Plummer AP. 1978. Antelope bitterbrush:
an important wildland shrub. Pub. 78-12. Salt Lake City: Utah State
Division of Wildlife Resources. 48 p.
Hideyuki I,Tsutomu H, Masateru M,Takashi Y, Mutsuo K, Harukuni T, Takuo O.
1995. Tannins from Cowania mexicana, and their inhibitory effects on
TPA-induced EBV activation. In: Brouillard R, Jay M, Scalbert A, eds.
Polyphenols 94, Paris: Institut National de la Recherche Agronomique
(INRA): 419420.
Ito H, Miyake M, Nishitani E, Mori K, Hatano T, Okuda T, Konoshima T,
Takasaki M, Kozuka M, Mukainaka T,Tokuda H, Nishino H,Yoshida T. 1999.
Anti-tumor promoting activity of polyphenols from Cowania mexicana
and Coleogyne ramosissima. Cancer Letters 143: 513.
Jabbes M. 2000. Hybridization and its evolutionary consequences in Purshia
and Cowania [dissertation]. Moscow, ID: University of Idaho. 185 p.
Krannitz PG. 1997a. Variation in magnesium and nitrogen content in seeds
of antelope bitterbrush (Purshia tridentata (Rosaceae)). American
Midland Naturalist 138: 306321.
Krannitz PG. 1997b. Seed weight variability of antelope bitterbrush (Purshia
tridentata: Rosaceae). American Journal of Botany 84: 17381742.
Kyle NE, Righetti TL. 1996. Observations of shoots and roots from interspecific grafted rosaceous shrubs. Journal of Range Management 49:
350354.
Landis TD, Simonich EJ. 1984. Producing native plants as container
seedlings. In: Murphy PM, ed.The challenge of producing native plants for
the Intermountain area. Gen.Tech. Rep. INT-168. Ogden, UT: USDA
Forest Service, Intermountain Forest and Range Experiment Station:
1625.
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
McArthur ED, Stutz HC, Sanderson SC. 1983. Taxonomy, distribution, and
cytogenetics of Purshia, Cowania, and Fallugia (Rosoideae, Rosaceae). In:
Tiedemann AR, Johnson KL, comps. Proceedings, Research and
Management of Bitterbrush and Cliffrose in Western North America;
1982 April 1315; Salt Lake City. Gen.Tech. Rep. INT-152. Ogden, UT:
USDA Forest Service, Intermountain Forest and Range Experiment
Station: 424.
McCarty EC, Price R. 1942. Growth and carbohydrate content of important mountain forage plants in central Utah as affected by clipping and
grazing.Tech. Bull. 818. Washington, DC: USDA. 51 p.
Meyer SE. 1989. Warm pretreatment effects on antelope bitterbrush
(Purshia tridentata) germination response to chilling. Northwest Science
63: 146153.
Meyer SE. 2002. Unpublished data. Provo, UT: USDA Forest Service,
Intermountain Research Station.
Meyer SE, Monsen SB. 1989. Seed germination biology of antelope bitterbrush (Purshia tridentata). In: Wallace A, McArthur ED, Haferkamp MR,
comps. Proceedings, Symposium on Shrub Ecophysiology and
Biotechnology; 1987 June 30July 2; Logan, UT. Gen.Tech. Rep. INT-256.
Ogden, UT: USDA Forest Service, Intermountain Forest and Range
Experiment Station: 147157.
Meyer SE, Kitchen SG, Wilson GR, Stevens R. 1988. Proposed rule for
Cowania mexicanacliffrose. Association of Official Seed Analysts
Newsletter 63: 2425.
Nakanishi T, Nishi M, Somekawa M, Murata H, Mizuno M, Iinuma M,Tanaka T,
Murata J. 1994. Structures of new and known cyanoglucosides from a
North American plant, Purshia tridentata DC. Chemical Pharmaceutical
Bulletin 42:22512255.
Nelson DL. 1983. The occurrence and nature of Actinorhizae on Cowania
stansburiana and other Rosaceae. In:Tiedemann AR, Johnson KL, comps.
Proceedings, Research and Management of Bitterbrush and Cliffrose in
Western North America; 1982 April 1315; Salt Lake City. Gen.Tech.
Rep. INT-152. Ogden, UT: USDA Forest Service, Intermountain Forest
and Range Experiment Station: 225239.
Ngugi KR, Powell J, Hinds FC, Olsen RA. 1992. Range animal diet composition in southcentral Wyoming. Journal of Range Management 45:
542545.
Nord EC. 1965. Autecology of bitterbrush in California. Ecological
Monographs 35: 307334.
Pearson BO.1957. Bitterbrush seed dormancy broken with thiourea.
Journal of Range Management 10: 4142.
Plummer AP, Christensen DR, Monsen SB. 1968. Restoring big game range
in Utah. Pub. 68-3. Salt Lake City: Utah Division of Fish and Game.
183 p.
Righetti TL, Chard CH, Munns DN. 1983. Opportunities and approaches
for enhancing nitrogen fixation in Purshia, Cowania and Fallugia. In:
Tiedemann AR, Johnson KL, comps. Proceedings, Research and
Management of Bitterbrush and Cliffrose in Western North America;
1982 April 1315; Salt Lake City. Gen.Tech. Rep. INT-152. Ogden, UT:
USDA Forest Service, Intermountain Forest and Range Experiment
Station: 214224 .
Sargent CS. 1965. Manual of the trees of North America (exclusive of
Mexico). 2 volumes; 2nd corr. ed. New York: Dover. 934 p.
Scholten GC. 1983. Bitterbrush management on the Boise wildllife management area. In:Tiedemann AR, Johnson KL, comps. Proceedings,
Research and Management of Bitterbrush and Cliffrose in Western
North America; 1982 April 1315; Salt Lake City. Gen.Tech. Rep. INT152. Ogden, UT: USDA Forest Service, Intermountain Forest and Range
Experiment Station: 153162.
Purshia
921
RosaceaeRose family
Pyrus L.
pear
Several Asian species have fruits with the size and shape
of a pea. The Japanese and Korean pea pears and the evergreen pear are considered by some to be varieties or subspecies of Callery pear (Rehder 1986; Yu 1979). The birchleaf pear has the smallest sized fruit of the pea pears.
The common pear has naturalized in the United States
(Gill and Pogge 1974). The Ussuri pear, introduced from
Asia about 1855, has been grown on the northern Great
Plains in shelterbelt and environmental plantings and in New
England. It has contributed genes for cold-hardiness and
resistance to fire blight in pear breeding programs (Stushnoff
and Garley 1982). Other traits inherent in this species
include vigor, dense growth, attractive glossy foliage, and
scarlet autumn leaf color. Pear cultivars adapted to warm
winter areas have been derived from the Pashia pear of central Asia. The pendulous form of the willow-leaf pear makes
it a unique ornamental landscape plant. Flowering ornamental selections of the Callery pear and the evergreen pear are
widely planted as street trees in the United States. The use of
the evergreen pear is limited to warm-winter areas such as
California and the more southerly states. These species are
often referred to as flowering pears in the urban landscape.
The Callery pear has become naturalized in the eastern
United States and is now considered a weed in some areas
such as the Maryland suburbs of Washington, DC.
Pears are deciduous, rarely evergreen, sometimes thorny
trees or shrubs. Their leaves are serrate, crenate, or entire;
rarely lobed. The petioles are stipulate and the buds are involute, with imbricate scales.
Flowering and fruiting. Pear species are cross-compatible sexual diploids (x = 17). Individual genotypes are
generally self-incompatible. The perfect flowers bloom on
2-year or older spurs, between March and April in the
Northern Hemisphere and appear before or with the new
leaves (table 2). The inflorescence consists of 6 to 8 flowers
occurring in umbel-like racemes. Petals are white, or rarely
pinkish with reflexed or spreading sepals, 20 to 30 pink, red,
Common name(s)
Growth habit
P. amygdaliformis Vill.
P. sinaica Dom.-Cours.
P. betulifolia Bunge
P. calleryana Decne.
almond-leaf pear
Shrub to small
tree, 12 m
Large tree, 56 m
Medium tree, 35 m
Mediterranean Europe
& Asia Minor
Central & N China
Central & S China
W to SE Europe,
Turkey; in world-wide
cultivation
Caucasus pear
Large tree, 56 m
SE Europe, Greece
heart-leaf pear,
Plymouth pear
Algerian pear
SW England,W France,
Spain, & Portugal
Algeria
Medium tree, 34 m
Japan
elaeagnus-leaf pear
Medium tree, 34 m
Korea
P. communis L.
P. asiae-mediae Popov; P. balansae Decne
P. boissieriana Buhse; P. elata Rubtzov
P. medvendevii Rubtzov
P. communis ssp. caucasica
(Fed.) Browicz
P. caucasica Fed.
P. cordata Desv.
P. cossonii Rehder
P. longipes Coss, S. Dur.
P. dimorphophylla Makino
P. calleryana var. dimorphophylla
(Makino) Koidz
P. elaeagrifolia Pall.
P. kotschyana Boiss ex Deone
P. fauriei C.K. Schneid.
P. calleryana var. fauriei (Schneid.) Rehd.
P. gharbiana Trab.
P. glabra Boiss.
P. koehnii C.K. Schneid
P. korshinskyi Litv.
P. pyraster Burgsd.
P. communis var. pyraster
P. mamorensis Trab.
P. nivalis Jacq.
P. pashia Buch.-Ham. ex D.Don
P. kumaoni Decne
P. varoilosa Wall ex G. Don.
P. wilhelmii C. Schneid.
P. pseudopashia T.T.Yu
P. pyrifolia (Burm.f.) Nakai
P. serotina Rehd.
P. regelii Rehder
P. heterophylla Regel G.Schmalh
P. salicifolia Poll.
P. syriaca Boiss.
P. ussuriensis Maxim.; P. lindleyi Rehd.
P. ovoidea Rehd.
P. sinensis Lindley
P. xerophylla T.T.Yu
Sources:
birch-leaf pear
Callery pear, pea pear,
Chinese pea pear
common pear,
European pear,
cultivated pear
evergreen pear
wild European pear
Small tree
Thornless medium tree,
34 m
Medium tree, 34 m
Morocco
W Central & S Europe
Kansu pear
sand pear, Japanese
pear, Chinese pear
Regel pear
Tree
Medium to large tree,
35 m
Shrub or tree to
12 m
Small tree, 12 m
Small tree, 12 m
Small to medium
tree, 13 m
Tree
N China
willow-leaf pear
Syrian pear
Ussuri[an] pear,
Harbin pear,
Manchurian pear
LHBH (1976), Bell (1991), Hedrick (1921), Lombard and Westwood (1987), Rehder (1986).
Pyrus
923
Bloom season
Ripening season
P. amygdaliformis
P. betulifolia
P. calleryana
P. communis (wild types)
P. communis (cultivars)
P. cordata
P. cossonii
P. dimorphophylla
P. elaeagrifolia
P. fauriei
P. gharbiana
P. glabra
P. hondoensis
P. koehnei
P. korshinskyi
P. mamorensis
P. nivalis
P. pashia
P. pyrifolia (wild types)
P. pyrifolia (cultivars)
P. regelii
P. salicifolia
P. syriaca
P. ussuriensis (wild types)
P. ussuriensis (cultivars)
MML
MMLL
EEMM
EMMML
EEMMMLL
MMLL
MMLL
EMML
EMMML
EMMML
ML
EM
EMMML
EEMMML
EMMML
ML
ML
EEMMMLL
EMM
EMMML
MML
EMMML
EMM
EEM
EEMM
L
L
L
EMMMLL
EEMMMLL
MLL
MMLL
L
ML
MLL
ML
MML
MMLL
L
EMMMLL
L
MLL
L
MMLL
EMMMLL
ML
MLL
MLL
EMMM L
MMLL
* Observations made at the USDA ARS National Clonal Germplasm Repository in Corvallis, OR, 1988 through 1994.
Average full bloom: E = March 13March 23, EM = March 24April 2, M = April 3April 7, ML = April 8April 17, L = April 18April 26.
Average fruit ripening: E = before July 6, EM = July 6August 8, M = August 9August 25, ML = August 26September 28, L = after September 28.
or purple anthers, 2 to 5 free styles that are closely constricted at the base, and 2 ovules per locule.
The fruit is a globose or pyriform pome with persistent
or deciduous calyx. Most Asian species, with the exception
of the Ussuri pear, have deciduous calyxes. The fruit of different species ranges from about 0.5 to 20 cm in length and
are quite diverse (figure 1). The extracarpellary tissue, which
comprises the bulk of the fruit flesh, may contain sclerenchyma, that is, stone cells. The ground-color of the fruit skin
may change from green to yellow or red during maturation,
and russeted lenticels may be prominent on some species.
Environmental conditions, such as humidity, may cause russetting or browning of the maturing skin. The ripening season for cultivated pears in the Northern Hemisphere ranges
from June through December (table 2). Fruit from some
species can be eaten directly from the tree, whereas others
may require a period of cold storage to ripen or soften the
fruit before it can be eaten. Common pears growing wild in
Russia are reported to be biennial producers (Albenskii and
Nikitin 1956).
924
fruit and seed of P. ussuriensis, Ussuri pear (left); seeds of P. calleryana, Callery pear (right).
longitudinal
Pyrus
925
Species
P. amygdaliforms
P. betulifolia
P. calleryana
P. communis ssp. caucasica
P. communis (domestic)
P. cordata
P. dimorphophylla
P. elaeagrifolia
P. fauriei
P. gharbiana
P. koehnii
P. mamorensis
P. nivalis
P. pashia
P. pyrifolia
P. regelii
P. salicifolia
P. syriaca
P. ussuriensis
2527
5586
3087
130
90
6588
90127
3888
6078
5058
110
1543
120170
100
Best
chilling Length
temp (C) (mm)
7
4
7
4
4
4
7
4
7
7
7
7
4
10
4
4
7
7
6.7
4.0
5.2
7.7
8.4
4.6
5.2
6.7
4.7
4.6
4.4
8.9
10.0
6.5
8.7
11.3
7.2
9.3
7.4
Seed size
Width
(mm)
L/W
ratio
/kg
/lb
4.2
2.3
2.6
4.2
4.8
2.6
2.8
4.2
2.9
2.4
2.4
5.9
4.3
3.1
4.4
7.6
4.6
6.2
4.5
1.60
1.74
2.00
1.83
1.77
1.77
1.86
1.6
1.62
1.92
1.83
1.51
2.32
2.10
1.98
1.49
1.59
1.50
1.64
24,000
90,000
55,000
40,000
22,000
86,000
77,000
22,000
57,000
99,000
79,000
11,000
18,000
55,000
26,000
7,000
24,000
9,000
20,000
11,000
41,000
25,000
18,000
10,000
39,000
35,000
10,000
26,000
45,000
36,000
5,000
8,000
25,000
12,000
3,000
11,000
4,000
9,000
Seeds/wt
Sources: Gill and Pogge (1974), Lombard and Westwood (1987), Rudolph (1949), Swingle (1939),Westwood and Bjornstad (1968),Yerkes (1930),Young and Young (1992)
Figure 4Pyrus communis, common pear: seedling development at 1, 2, 3, 6, and 12 days after germination.
926
embryos should be germinated for 10 to 14 days at alternating temperatures of 18/22 C (AOSA 1993).
Nursery practice. Seeds are planted thickly, about 13
mm (1/2 in) deep in a seedbed, and allowed to grow for 1
season. The following spring, plants are dug, their roots and
top are cut back, and they are transplanted to nursery rows.
After a second season, the rootstock are of correct size for
budding in the fall (Hartmann and others 1990). Seedlings
of 1+0 nursery stock can be either field-planted or rootpruned at a depth of 15 to 20 cm (6 to 8 in) and transplanted
for 1 year (Gill and Pogge 1974). Common pear seedlings
may be subject to powdery mildew, which is caused by
Podosphaera leucotricha (Ellis & Everh.) E.S. Salmon, and
by root rots.
Cultivars are propagated by budding or grafting onto
rootstocks. Bench-grafting dormant scions onto bareroot
rootstocks is no longer common in large-scale nursery production. Nursery trees can be produced more efficiently by
T-budding onto field-grown rootstocks in late summer when
the bark is slipping. Chip-budding is an alternative technique for seasons when the rootstock bark is not slipping
(Frecon 1982). A whip-and-tongue graft or cleft-graft is
commonly used when top-working growing trees in early
spring. Scions can be grafted a few centimeters off the
ground on a young rootstock, as in side-grafting, or multiple
grafts can be placed higher up onto scaffold branches to
convert an older tree over to a different cultivar, that is, topworking.
ible with a number of other genera in the Maloideae subfamily, including serviceberry (Amelanchier), cotoneaster
(Cotoneaster), hawthorn (Crataegus), apple (Malus), medlar
(Mespilus), squaw-apple (Peraphyllum), mountain-ash
(Sorbus), and others (Lombard and Westwood 1987;
Postman 1992). The common quince (Cydonia oblonga
Mill.) has traditionally been used as a dwarfing rootstock for
edible European pears.
References
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S-kh. Lit., Moskva. [Handbook of afforestation and soil amelioration.
Transl.TT 66-51098, 1967. Springfield,VA: USDC CFSTI, 516 p.].
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing
seeds. Journal of Seed Technology 16(3): 1113.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan. 1290 p.
Bell RL. 1991. Pears (Pyrus). In: Moore JN, Ballington JR Jr, eds. Genetic
resources of temperate fruit and nut crops. Wageningen,The
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Ellis RH, Hong TD, Roberts EH. 1985. Handbook of seed technology for
genebanks.Volume 2, Compendium of specific germination and test recommendations. Rome: FAO International Board of Plant Genetic
Resources.
Gill JD, Pogge FL. 1974. Pyrus, pear. In: Seeds of woody plants in the United
States. Agriculture Handbook 450. Washington, DC: USDA Forest
Service: 584586.
Hartmann HT, Kester DE, Davies F. 1990. Plant propagation: principles and
practices. 5th ed. Englewood Cliffs, NJ: Prentice Hall. 662 p.
Hedrick UP. 1921. The pears of New York. Geneva, NY: New York
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ISTA [International Seed Testing Association]. 1993. Rules for testing seeds:
rules 1993. Seed Science and Technology 21 (Suppl.): 1259.
Kikuchi A. 1946. Speciation and taxonomy of Chinese pears [in Japanese].
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Lombard PB, Westwood MN. 1987. Pear rootstocks. In: Rom RC, Carlson RF,
eds. Rootstocks for fruit trees. New York: Wiley: 145183.
Macdonald B. 1986. Practical woody plant propagation for nursery
growers. Portland, OR:Timber Press. 660 p.
Postman JD. 1992. Graft compatibility of different pome fruit genera.
Pomona 25(3):1517.
Rehder A. 1986. Manual of cultivated trees and shrubs. 2nd ed, Portland,
OR: Dioscorides Press: 401406.
Rudolf PO. 1949. First the seed, then the tree. In: USDA Yearbook of
Agriculture (1949):Trees: 127135.
Stushnoff C, Garley B. 1982. Breeding for cold hardiness. In: van der Zwet T,
Childers NF, eds. The pear. Gainesville, FL: Horticultural Publications:
189205.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA Soil
Conservation Service. 187p.
Teng Y,Tanabe K,Tamura F, Itai A. 2002. Genetic relationships of Pyrus
species and cultivars native to east Asia revealed by randomly amplified
polymorphic DNA markers. Journal of the American Society for
Horticultural Science 127(2): 262-270.
Westwood MN. 2002. Personal communication. Corvallis, OR: Oregon
State University.
Westwood MN, Bjornstad HO. 1968. Chilling requirements of dormant
seeds of 14 pear species as related to their climatic adaptation.
Proceedings of the American Society of Horticultural Science 92:
141149.
Yamamoto T, Kimura T, Sawamura Y, Manabe T, Kotobuki K, Hayashi T, Ban Y,
Matsuta N. 2002. Simple sequence repeats for genetic analysis in pear.
Euphytica 124:129-137.
Yeo DY, Reed BM. 1995. Micropropagation of three Pyrus rootstocks.
HortScience 30(3): 620623.
Yerkes GE. 1930. Raising root stocks from seed in the United States.
International Horticulture Congress Proceedings 9: 8391.
Young JA,Young CG. 1992. Seeds of woody plants in North America. Rev.
ed. Portland, OR: Dioscorides Press. 407 p.
Yu T. 1979. Taxonomy of the fruit tree in China [in Chinese]. Beijing: China
Agriculture Press.
Pyrus
927
FagaceaeBeech family
Quercus L.
oak
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Growth habit, occurrence, and use. The oaks
members of the genus Quercusinclude numerous species
of deciduous and evergreen trees and shrubs and make up
the single most economically important genus of hardwoods
in North America. Quercus is also the largest genus of trees
native to the United States (Little 1979) and has recently
been designated as the national tree by the National Arbor
Day Foundation. About 500 species are widely distributed
throughout the temperate regions of the Northern Hemisphere in both the Eastern and Western Hemispheres as well
as southward through Central America to the mountains of
Colombia and through Turkey to Pakistan (Sargent 1965).
There are about 58 tree and 10 shrub species native to the
United States, 104 species in Mexico, and another 30 in
Central America and Colombia. At least 70 hybrids have
been described, and there are probably many more (Little
1979). Information on hybrids and genetic variation has
been summarized for 25 species in Burns and Honkala
(1990).
Oaks are divided into 2 subgenera: Lepidobalanus
(white oaks) and Erythrobalanus (black oaks). These subgenera differ in several ways, but most importantly for seed
considerations, they differ in time required for fruit maturation, chemical composition of their stored food reserves, and
degree of dormancy. In this book, 48 taxa are considered
(table 1). Oaks are valuable for a very wide range of products and uses: construction timber, furniture, interior trim,
and flooring; watershed protection, wildlife habitat and
food, and ornamental plantings; as well as tannins and other
extractives and cork. Consequently, many oak species are
widely planted for a variety of purposes. For additional
information on growth habit, uses, ecology, and silviculture
of individual oak species, consult Burns and Honkala
(1990).
Flowering and fruiting. Flowering is monoecious.
The staminate flowers are borne in clustered aments
(catkins) and the pistillate flowers in solitary (or in 2- to
many-flowered) spikes in the spring (February to May)
928 Woody Plant Seed Manual
Figure 1 Quercus, oak: acorns of (top row, left to right) Q. alba, white oak; Q. falcata, southern red oak; Q. kelloggii,
California black oak; Q. lyrata, overcup oak. (second row, left to right) Q. macrocarpa, bur oak; Q. marilandica, blackjack
oak; Q. michauxii, swamp chestnut oak. (third row, left to right) Q. muehlenbergii, chinkapin oak; Q. nigra, water oak;
Q. pagoda, cherrybark oak; Q. phellos, willow oak. (fourth row, left to right), Q. rubra, northern red oak; Q. shumardii,
Shumard oak; Q. sinuata, Durand oak; Q. stellata, post oak. (bottom row, left to right), Q. texana, Nuttall oak;
Q. velutina, black oak; Q. wislizeni, interior live oak.
Quercus
929
Group*
Common names
Occurrence
Q. acutissima Carr.
Q. agrifolia Ne
white
black
Q. alba L.
white
Q. arizonica Sarg.
white
Q. bicolor Willd.
white
Q. cerris L.
white
Q. chrysolepis Liebm.
white
Q. coccinea Muenchh.
black
white
Q. dumosa Nutt.
white
Q. ellipsoidalis E. J. Hill
black
Q. emoryi Torr.
black
Q. falcata Michx.
Q. triloba Michx.
Q. gambelii Nutt.
Q. vreelandii Rydb.
Q. utahensis (A. DC.) Rydb.
Q. garryana Dougl. ex Hook.
black
Q. grisea Liebm.
white
sawtooth oak
California live oak,
coast live oak; encina
white oak, fork-leaf
white & stave oaks
Arizona white oak,
Arizona oak; roble
swamp white oak,
cow oak
European turkey oak,
turkey oak
canyon live oak, canyon,
maul, goldcup, & live oaks
scarlet oak, black &
Spanish oaks
blue oak, California blue,
iron, & mountain white oaks
California scrub
oak, scrub oak
northern pin oak,
black, jack, & Hill oaks
Emory oak, black
oak, bellota, roble negro
southern red oak,
Spanish & red oaks
Gambel oak, Rocky
Mtn. white & Utah
white oaks; encino
Oregon white oak, Garry,
post, Oregon, Brewer, & shin oaks
gray oak
Q. ilicifolia Wangenh.
black
Q. imbricaria Michx.
black
Q. incana Bartr.
black
Q. kelloggii Newb.
black
Q. laevis Walt.
Q. catesbaei Michx.
Q. laurifolia Michx.
black
Q. lobata Ne
white
Q. lyrata Walt.
white
Q. macrocarpa Michx.
white
930
white
white
black
Group*
Common names
Occurrence
Q. marilandica Muenchh.
black
Q. michauxii Nutt.
Q. prinus L.
white
blackjack oak,
barren & jack oaks; blackjack
swamp chestnut oak,
cow & basket oaks
Q. muehlenbergii Engelm.
white
Q. nigra L.
black
Q. pagoda Raf.
Q. falcata var pagodaefolia Ell.
black
Q. palustris Muenchh.
Q. petraea (Mattusch) Liebl.
Q. sessiliflora Salisb.
Q. phellos L.
Q. prinus L.
Q. montana Willd.
Q. robur L.
black
white
black
white
white
Q. rubra L.
Q. borealis Michx.f.
black
Q. shumardii Buckl.
black
Q. sinuata Walt.
Q. durandii Buckl.
Q. stellata Wangenh.
white
Q. suber L.
Q. texana Buckl.
Q. nuttallii Palmer
Q. turbinella Greene
white
black
white
white
white
white
black
black
Q. virginiana P. Mill.
white
Q. wislizenii A. DC.
black
Quercus
931
longitudinal
valley, blue, and California black oaks and canyon live and
coast live oaks (Koenig and others 1994) found no mast
production patterns at the population level. Crop failures did
occur frequently but they were probably more related to
lack of pollination and fertilization success than to inherent
patterns. Cecich (1993) concluded that most of the potential
seedcrop in oaks in Missouri is lost when pistillate flowers
abort between the time of pollination and fertilization.
Really good crops of California black oak acorns were
found to occur only every 8 years or so (McDonald 1992).
The following yield averages on an area basis have been
reported: 3.2 to 1,620 kg/ha (2.9 to 1,448 lb/ac) for white
oak in Illinois (Johnson 1975); 208 kg/ha (186 lb/ac) for
southern Appalachian oaks (Beck 1977); and 560 kg/ha
(500 lb/ac) for Oregon white oak in California (Stein 1990).
Collection and cleaning of acorns. Collecting acorns
of high quality requires an awareness of the indices of acorn
maturity. Natural dissemination from the tree is a sure sign
of maturity, of course, but collections are often made before
this time to reduce losses to deer, rodents, and other predators that quickly eat fallen acorns. Good indices of maturity
for most species are (1) change in pericarp color from green
to yellow, brown, or black; (2) a cup scar colored pink,
lemon, orange, or white; and (3) cups that slip easily from
the acorns without resistance (Bonner and Vozzo 1987;
Lotti 1959). Ripe acorns may be collected from August to
December from the ground or they can be shaken from trees
onto canvas or plastic sheets after ripening. Mechanical tree
shakers can be very effective with oaks where the terrain or
932
Height at
maturity (m)
15
23
30
12
30
30
30
30
18
6
21
18
27
15
21
20
6
21
12
26
9
27
30
24
30
15
30
24
24
34
24
30
30
24
34
30
34
23
18
24
30
3
1
24
27
18
18
Year first
cultivated
1862
1849
1724
1800
1735
1877
1691
1902
1763
1873
1800
1724
1878
1834
1786
1874
1786
1811
1737
1822
1723
1904
1770
Long
1723
1688
Long
1724
1907
1819
1699
1923
1895
1861
1905
1739
1874
5
15
20
20
20
20
25
25
30
15
25
35
20
20
25
20
40
20
20
20
25
25
25
12
5
20
Q
Years between
large seedcrops
410
35
24
35
23
24
12
23
24
23
12
1
23
34
23
35
12
12
12
57
1
23
24
35
23
23
24
34
35
2
23
1
57
Sources: Burns and Honkala (1990), Olson (1974), Sargent (1965), Smith (1993), Sork and others (1993),Vines (1960).
Quercus
933
Species
Q. acutissima
Q. agrifolia
Q. alba
Q. bicolor
Q. cerris
Q. chrysolepis
Q. coccinea
Q. douglasii
Q. dumosa
Q. ellipsoidalis
Q. falcata
Q. garryana
Q. ilicifolia
Q. imbricaria
Q. incana
Q. kelloggii
Q. laevis
Q. laurifolia
Q. lobata
Q. lyrata
Q. macrocarpa
Q. michauxii
Q. muehlenbergii
Q. nigra
Q. pagoda
Q. palustris
Q. petraea
Q. phellos
Q. prinus
Q. robur
Q. rubra
Q. shumardii
Q. sinuata
Q. stellata
Q. suber
Q. texana
Q. turbinella
Q. vaccinifolia
Q. variabilis
Q. velutina
Q. virginiana
Q. wislizenii
Seed weight/
fruit vol
kg/hl
lb/bu
58129
3977
4264
50
3945
5180
6066
5772
5960
28134
64
53
69
67
33
5363
71
36
45100
3060
3350
39
3035
4062
4751
4456
4647
22104
50
47
54
52
26
4149
55
28
Range
/kg
210245
155465
200385
130320
110310
230890
120330
450640
7051,730
165220
6951,750
5001,500
115325
8601,520
165525
285340
90300
75430
5801,145
5101,545
9251,640
7051,190
130650
6001,530
120430
200495
165565
170280
4401,400
110220
125315
660770
16302,910
165275
275882
5301,125
100152
Cleaned seeds/weight
Average
/lb
/kg
/lb
95110
70210
90175
60145
50150
105405
55180
205290
320785
75100
315795
225680
52145
90690
75237
130154
40135
35195
265520
230700
420745
320540
60295
270695
55195
90225
75255
80130
200635
50100
55145
300350
7401,320
75125
125400
240510
100150
85
200
98
265
240
520
220
220
540
1,190
185
1545
915
210
870
1,235
285
265
165
125
870
640
690
475
375
835
220
285
235
220
6,400
840
165
220
715
2,270
230
540
775
275
187
440
215
120
110
235
100
100
245
540
85
700
415
95
395
560
130
120
75
55
395
290
312
220
170
380
100
130
105
100
290
380
75
100
325
1,030
105
245
350
125
Samples
2
1
23
3
4
4
4
1
11
9
3
1
11
49
1
3
4
6
8
35
4
226
41
33
9
183
5
10
55
27
1
9
13
83
2
2
12
7
4
3
Sources: Burns and Honkala (1990), Olson (1974),Toumey and Korstian (1942),Van Dersal (1938).
934
1980). Acorns of the black oak group exhibit variable dormancy that is apparently imposed by the pericarp, the
embryo, or both (Hopper and others 1985; Jones and Brown
1966; Peterson 1983), and stratification is usually recommended before spring-sowing or certain types of germination tests. Epicotyl dormancy has been reported in at least
1 black oak speciesbear oak (Allen and Farmer 1977). If
proper procedures are followed for storage of black oak
acorns, the storage conditions will also serve to complete the
stratification requirement, and additional treatment is not
necessary (Bonner and Vozzo 1987). If additional stratification is needed, imbibed acorns should be held for 4 to 12
weeks at temperatures of 2 to 5 C. The acorns may be
mixed with peat or other media, but this is not necessary.
Most managers stratify in plastic bags without medium,
turning the bags each week or so to prevent pooling of
excess moisture in the bags (Bonner and Vozzo 1987).
Acorns of the black oak group sown in the fall or early winter need not be stratified before to sowing.
Germination tests. In the standard official laboratory
test procedure for all oaks, the acorns should be soaked in
water for 48 hours; then a third of the acorn at the cup scar
end should be cut off and the pericarp removed from the top
half and placed on thick, moist blotters at alternating temperatures of 20 to 30 C (ISTA 1993). No other pretreatments are necessary, and germination should be complete
within 14 days. Germination can also be tested with intact
acorns in sand, peat, or other media in greenhouse flats. In
such tests, stratification may be necessary for black oak
species (table 4). Germination is hypogeal (figure 3) and is
generally complete in 3 to 5 weeks. Rapid estimates of viability can also be made with cutting tests, radiography, or
tetrazolium staining (Belcher and Vozzo 1979; Bonner and
Vozzo 1987). Cutting tests are reliable on freshly collected
acorns, and radiography is very good for quick determination of insect infestation. Tetrazolium staining can also provide information on seed vigor, but acorn chemistry and
morphology present some problems in this test (Bonner
1984).
Nursery practice. Numerous research studies have
shown that success in planting oaks depends on production
of vigorous seedlings through low sowing densities and
undercutting in the beds (Schultz and Thompson 1990).
Container production in greenhouses is also practiced for a
few species (Tinus 1980). Fall-sowing acorns is preferable
to spring-sowing in many instances if weather allows bed
preparation in the fall. Fall-sowing eliminates the need for a
large storage capacity for acorns and avoids the problems of
fungi and early germination in storage. One disadvantage to
Quercus
935
Species
Q. acutissima
Q. agrifolia
Q. alba
Q. bicolor
Q. cerris
Q. chrysolepis
Q. coccinea
Q. douglasii
Q. durmosa
Q. ellipsoidalis
Q. falcata
Q. gambelii
Q. garryana
Q. ilicifolia
Q. imbricaria
Q. kelloggii
Q. laevis
Q. laurifolia
Q. lyrata
Q. macrocarpa
Q. marilandica
Q. michauxii
Q. muehlenbergii
Q. nigra
Q. pagoda
Q. petraea
Q. phellos
Q. prinus
Q. robur
Q. rubra
Q. shumardii
Q. sinuata
Q. stellata
Q. suber
Q. texana
Q. turbinella
Q. vaccinifolia
Q. variabilis
Q. velutina
Q. virginiana
Q. wislizenii
Cold
stratification
(days)
0
0
0
0
060
3060
0
3090
6090
3090
14
0
60120
3060
3045
6090
0
1490
0
42
3060
90
0
30
0
3060
60120
0
3090
0
0
0
3045
70
60120
0
0
0
6090
0
0
3060
0
3060
Germinative
rate
Avg (%)
Days
Kimpac
Sand
Germinator
Peat/loam
Kimpac
Sand
Sand
Sand
Sand
Loam
Sand/perlite
Sand
Sand
Sand
Soil
Sand
Sand
Sand
Sand
Soil
Soil
Kimpac
Sand/peat,
Kimpac
Sand/perlite
Sand
Soil, Kimpac
Soil
Sand
Sand
Sand
Sand/peat
Soil, Kimpac
Kimpac
Sand, Kimpac
Sand
Soil
Sand
Loam
Sand
Sand
Kimpac
Sand/peat
1540
3098
60240
30
5660
3060
30
28
3060
3057
90
3681
30
3040
7
108
3090
160
128
40
5084
50
45
5273
3993
6595
97
8093
6274
92
82
28-85
2348
86
95
5480
1041
80120
16
1826
2236
15
100
25-45
4060
22
8
3173
98
73
5099
7898
3376
5675
9499
7072
8090
95
75100
92
77100
8694
2866
95
82
50
4592
84
82
45
91
49
98
98
6094
1
1
21
3
3
2
7
4
3
5
8
1
4
12
2
1
2
1
6
1
4
11
1
2
1
4
12
3040
30
45100
90
60
3060
4060
20
2950
30
4560
2030
5887
180
28
3050
69
8590
41
83
7278
3985
80
5366
81
4293
38
55
92
2138
55
47
40
1342
10
2128
21
1045
30
28
8698
6574
67
89
82
81
58
100
7282
87
5498
73-100
6069
95
43
11
7
4
1
3
4
11
1
3
4
7
5
20
2
1
2
5
4
1
30
2135
22
30
30
30
30
30
2327
30
30
24
30
27
20
1016
20
20
20
20
20
21
2327
21
20
16
21
23
27
2135
27
30
32
32
30
3032
23
1016
23
20
21
21
20
2021
30
30
32
32
27
25
30
20
32
30
30
27
32
38
23
25
27
30
30
20
20
21
21
18
16
20
20
21
20
20
27
21
5
19
18
20
20
Germination
(%)
Samples
47
97
75
Sources: Dirr and Heuser (1987), Korstian (1927), Larsen (1963), Olson (1974), Swingle (1939).
936
seedling growth
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Jones EW. 1958. The storage of acorns in water. Forestry 31: 163166.
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von Steilund Traubeneichen (Quercus robur und Quercus petraea): 3.
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Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
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Lotti T. 1959. Selecting sound acorns for planting bottomland hardwood
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California oak species. American Midland Naturalist 122: 6676.
938
McCreary D, Koukoura Z. 1990. The effects of collection date and pre-storage treatment on the germination of blue oak acorns. New Forests 3(4):
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McDonald PM. 1990. Quercus kelloggii Newb., California black oak. In: Burns
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McDonald PM. 1992. Estimating seed crops of conifer and hardwood
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Mignery AL. 1975. Direct-seeding oaks on the Cumberland Plateau in
Tennessee. Res. Pap. SO-107. New Orleans: USDA Forest Service,
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Gambel oak in Utah. American Journal of Botany 67: 426428.
Oliver AD, Chapin JB. 1984. Curculio fulvus (Coleoptera: Curculionidae) and
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nigra L. seeds. Annals of Botany 52: 8192.
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Radford AE, Ahles HE, Bell CR. 1964. Guide to the vascular flora of the
Carolinas. Chapel Hill: University of North Carolina. 383 p.
Rink G, Williams RD. 1984. Storage technique affects white oak acorn
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RhamnaceaeBuckthorn family
Rhamnus L.
buckthorn
Andrew Youngblood
Dr.Youngblood is a research forester at the USDA Forest Services
Pacific Northwest Research Station, La Grande, Oregon
Rhamnus
939
Common name(s)
Occurrence
R. alnifolia LHr.
alder buckthorn
R. arguta Maxim.
R. cathartica L.
European buckthorn,
waythorn, common buckthorn
R. crocea Nutt.
R. pilosa (Trel.) Abrams
R. davurica Pallas
Dahurian buckthorn
hollyleaf redberry
Japanese buckthorn
lanceleaf buckthorn
lanceleaf buckthorn
island redberry
sawleaf buckthorn
Smith buckthorn
Chinese buckthorn
R. pirifolia Greene
R. crocea Nutt. var. pirifolia (Greene) Little
R. crocea Nutt. ssp. pirifolia (Greene) C.B.Wolf
R. serrata Humb. & Bonpl. ex
J.A. Schultes
R. fasciculata Greene
R. smithii Greene ssp. fasciculata (Greene)
C.B.Wolf
R. smithii Greene
R. smithii Greene ssp. typica C.B.Wolf
R. utilis Dcne.
940
Rhamnus
941
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in central New York. Ecological Monographs 61: 183205.
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Botany 15: 139143.
Guitian P. 1995b. Reproductive biology of Rhamnus legionensis Rothm., a
dioecious species endemic to the northwest Iberian peninsula. Flora
(1995) 190: 345352.
Hickman JC, ed. 1993. The Jepson manual: higher plants of California.
Berkeley: University of California Press. 1400 p.
Hitchcock CL, Cronquist A, Ownbey M,Thompson JW. 1961. Vascular
plants of the Pacific Northwest:Volume 3, Saxifragaceae to Ericaceae.
Seattle: University of Washington Press. 614 p.
Hubbard RL. 1974. Rhamnus, buckthorn. In: Schopmeyer CS, tech. coord.
Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 704-708.
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New York Botanical Garden. 96 p.
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American flora. XIII. Phytologia 76: 441457.
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Bonnicksen TM, Christensen NL [and others], tech. coords. Proceedings,
Fire Regimes and Ecosystem Properties Conference; 1978 December
1115; Honolulu, HI. Gen.Tech. Rep. WO-26. Washington, DC: USDA
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California chaparral. Ecology 68: 434443.
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Canadian Forestry Service, Department of the Environment. 272 p.
EricaceaeHeath family
Rhododendron L.
rhododendron and azalea
Frank A. Blazich and D. Bradley Rowe
Dr. Blazich is alumni distinguished graduate professor of plant propagation and tissue culture at North
Carolina State Universitys Department of Horticultural Science, Raleigh, North Carolina; Dr. Rowe is
associate professor at Michigan State Universitys Department of Horticulture, East Lansing, Michigan
Occurrence. The genus rhododendronRhododendron L.is indigenous mainly to the Northern Hemisphere,
with large concentrations in the mountain ranges of China,
Tibet, and upper Burma as well as in Japan and the eastern
United States. Plants are found commonly in regions with
highly organic soils, high rainfall, high humidity, and a temperate climate (Cox 1990). Species range from tiny, prostrate, alpine shrubs only 5 cm tall to trees with enormous
leaves that reach heights of 24 m (Leach 1961). Species
native to North America are listed in table 1.
Growth habit. There are over 900 species of rhododendrons and numerous cultivars (Davidian 1992). They
include many of the most spectacular flowering trees and
shrubs and are one of the most important and diverse groups
of ornamental plants in cultivation (Dirr and Heuser 1987).
The genus comprises both rhododendrons and azaleas.
General characteristics are listed in table 2; however, these
distinct characters are now known to be part of a continuum
of gradation. Therefore, there are no clear delineations
between azaleas and rhododendrons.
Uses. Besides their aesthetic appeal, rhododendrons in
the wild provide erosion control for steep watersheds and
protection for wildlife. In addition, some Himalayan species
have been utilized for medicinal purposes, as a tea substitute, or for incense (Cox 1990). Under cultivation, the
species are recognized as one of the most important plants
available due to their attractive foliage and extremely showy
flowers. For landscaping, rhododendrons are unsurpassed
with their variations in form, flower color, texture, and leaf
morphology. Those with larger leaves should be planted in a
woodland or similar setting. Rosebay rhododendron is ideal
as a woodland shrub or for tall evergreen backgrounds, but
its texture is much too coarse and its stature entirely too
large for home foundation plantings. Catawba rhododendron
and its western relative, west coast rhododendron, are also
well suited for woodland plantings, although in cooler climates Catawba rhododendron occurs often in full sun.
943
Common name(s)
Occurrence
Carolina rhododendron,
Carolina azalea
Catawba rhododendron,
Catawba rosebay, mountain
rosebay, purple laurel,
Chapmans rhododendron
Alabama azalea
Alabama
Cascade azalea
smooth azalea,
sweet azalea
coast azalea,
dwarf azalea
Florida flame azalea,
orange azalea
flame azalea,
yellow azalea
Kamchatka rhododendron
rhodora
Newfoundland to Pennsylvania
Cumberland rhododendron
Oconee azalea,
Piedmont azalea
Lapland rhododendron,
Lapland rosebay
Texas azalea
western azalea
S Oregon to S California
or cultivar, and does not come true from seed. Thus, vegetative propagation is essential, as seed propagation results in
inevitable variation among individuals. Generally, hybrids
are more adaptable because they possess a combination of
those genes required by their parents to withstand the environments where they originated. As a group, hybrids flower
944
Common name(s)
Occurrence
pinxterbloom, pinxter
flower, honeysuckle,
pink azalea
rose-shell azalea,
early azalea, piedmont azalea,
mayflower azalea
plumleaf azalea,
plum-leaved azalea
pink-shell azalea
North Carolina
Rhododendrons
Azaleas
Evergreen
Coriaceous
Scaly or punctate
Entire
Deciduous
Membranous
Pubescent
Ciliate or ciliolate
Campanulate
10 or more
Scaly or tomentose
Funnelform
5
Setose
Rhododendron
945
EVERGREEN RHODODENDRONS
R. carolinianum
Compact shrub; to 1.8 m
R. catawbiense
Spreading, rounded in the open; generally wider
than tall to 3 m, sometimes small tree to 6 m
R. chapmanii
Shrub to 1.8 m
R. macrophyllum
Open tree-like shrub; often erect to 39 m
R. maximum
Shrub in cultivation; to 4.6 m
(sometimes to 12 m in the wild)
R. minus
2.8 m
DECIDUOUS RHODODENDRONS
R. alabamense
Low stoloniferous shrub; to 0.62.4 m
R. albiflorum
Erect shrub; from 0.92.1 m
R. arborescens
From low spreading bushes in open
to tall and leggy in shade; up to 6 m
R. atlanticum
Stoloniferous shrub, forms branching
sprays when well established; 0.31.5 m
R. austrinum
Stiff and upright; from 3.03.6 m
R. calendulaceum
Stiff and upright; to 3.6 m
R. camtschaticum
Very small shrub; to 0.2 m
R. canadense
Much branched shrub; to 0.9 m
R. canescens
Sparingly branched shrub; to 4.6 m
R. cumberlandense
Low and twiggy, often stoloniferous
shrub; to 2.4 m but rarely over 1.8 m
R. flammeum
Mounding form; to 2.5 m
R. lapponicum
Dwarf, procumbent shrub; to 0.3 m
R. oblongifolium
Upright, somewhat stoloniferous shrub; to 1.8 m
R. occidentale
Rounded, occasionally upright or low shrub; to 1.04.6 m
R. periclymenoides
Usually tall, vigorous and much- branched
shrub; to 2.7 m & up to 4.5 m in wild
R. prinophyllum
Upright, well branched shrub; to 2.5 m
R. prunifolium
Tall, rounded-topped; up to 3.6 or 5.5 m in wild
R. vaseyi
Upright shrub to 3.6 m
R. viscosum
Form various: large & upright to dwarf, small tree; from 36 m,
rounded or straggly shrub, stoloniferous form to 4.6 m
Sources: Davidian (1992), Leach (1961), LHBH (1976).
946
Flowering
Flower color
May
MayJune
May
MayJune
JuneJuly
Rose
Purplish rose, white
White, pink, purplish red
June
Rose, white
AprMay
JuneJuly
JuneJuly
White
Creamy white, yellow
White
May
White, pink
Apr
MayJune
May
Apr
AprMay
JuneJuly
Yellow-orange
Yellow, orange, scarlet, pink
Reddish purple
Rose-purple, white
Pink, white
Yellow, orange, scarlet
May
Apr
June
AprAug
May
May
JulyAug
AprMay
JulyOct
Figure 2Rhododendron, rhododendron: seeds of R. calendulaceum, flame azalea (upper left); R. carolinianum, Carolina
rhododendron (upper right); R. catawbiense, Catawba rhododendron (center); R. chapmanii, Chapmans rhododendron
(lower left); and R. maximum, rosebay rhododendron (lower right).
Rhododendron
947
Figure 3Rhododendron macrophyllum, west coast rhododendron: seeds in external view (top left), longitudinal
section (center), and cross section (bottom right).
948
Table 4Rhododendron, rhododendron and azalea: variation in seed size among species and seed source
Species
Seed source
R. calendulaceum
R. carolinianum
R. catawbiense
R. chapmanii
R. macrophyllum
R. maximum
Elevation (m)
Seed moisture
content (%)
1,400
720
1,100
1,860
1,860
1,954
67
320
950
950
6
6
4
7
6
10
9
7
5.5
6
5
Cleaned seeds/wt
/g
/oz
4,350
29,460
23,930
6,070
6,070
6,780
5,700
5,000
29,100
4,460
11,790
11,430
122,000
825,000
670,000
170,000
170,000
190,000
160,000
140,000
815,000
125,000
330,000
320,000
Sources: Arocha and others (1999), Blazich and others (1991, 1993), Glenn and others (1998), Malek and others (1989, 1990), Olson (1974), Rowe and others (1994a).
Figure 5Rhododendron macrophyllum, west coast rhododendron: seedling development at 1, 9, 40, and 60 days
after germination.
for the light requirement for some species (Toole and others
1955). However, this inhibition usually dissipates by the end
of 30 days of germination (Blazich and others 1991, 1993;
Rowe and others 1994a).
A test of seeds of flame azalea collected from the Blue
Ridge Mountains of western North Carolina demonstrated
that (at a constant temperature of 25 C) increasing photoperiods increased germination, with maximum germination
(85%) occurring by day 12 under continuous light (Malek
and others 1989). An 8/16-hour thermoperiod of 25/15 C
enhanced germination when light was limiting. Maximum
germination of 84 to 91% was reached by day 24 for all
photoperiods 1/2 hour, although at photoperiods 4
hours, comparable germination was noted at day 18 (Malek
and others 1989). Similar results were reported for seeds of
949
from all provenances were similar. Regardless of temperature, seeds required light for germination, and daily
photoperiods as short as 1/2 hour maximized germination.
The major difference in germination response among provenances was related to seed vigor. Seeds from the Yancey
County (higher-elevation) provenance germinated at a faster
rate with greater cumulative germination than seeds from
lower elevation provenances.
In studying effects of irradiance on seed germination of
rosebay rhododendron, Romancier (1970) provided a range
of irradiance levels to seeds during 16-hr photoperiods at
22 C. He reported zero germination in total darkness but
found no significant differences in germination with light
intensities ranging from 1.6 mol/m2/sec (0.13 klux or 12
foot-candles) to 21.9 mol/m2/sec (1.72 klux or 160 footcandles), indicating that very low levels of irradiance will
stimulate germination. All seeds, including those in total
darkness, had been exposed to light before the test began, so
it is during the period following imbibition that light is
essential. Glenn and others (1999) reported that dormancy
was induced in seeds of Catawba and rosebay rhododendrons by not subjecting seeds immediately to light following
inbibition. However, the degree of dormancy varied depending on (a) the length of time imbibed seeds were maintained
in darkness and (b) the temperature at which the dark treatments were imposed and the seeds were germinated.
Nursery practice and seedling care. Rhododendrons
may be propagated by seeds, stem cuttings (Dirr and Heuser
1987; Hartmann and others 2002), layering (Wells 1985),
grafting (Wells 1985), and micropropagation (tissue culture)
(Anderson 1984; McCown and Lloyd 1983). Commercially,
plants usually are propagated by stem cuttings, although
rooting ability is genotype specific. Procedures developed
for micropropagation are currently being used with great
success. Nevertheless, seed propagation is still practiced to
develop new hybrids, raise understocks for grafting, and
propagate wild species.
Seeds should be sown in January or as early as local
conditions will allow. This is important to allow maximum
growth the first year. The longer the growing period before
mid-July (when growth normally ceases), the larger the
seedlings will be at the end of the first season (Leach 1961).
Many materials have been used as a germination medium,
including vermiculite, perlite, sawdust, peat, and various soil
mixes. Flats filled with peat moss and sand or perlite mixtures topped with 6 mm (1/4 in) of slightly firmed shredded
sphagnum moss work well (Wells 1985). Many propagators
are convinced that a medium consisting solely of shredded
sphagnum moss provides the best results (Leach 1961; Wells
950
long days at 16 different day/night temperature combinations, Rowe and others (1994b) found that a day/night cycle
of 22/22 C to 26/22 C was optimal for seedling growth,
whereas cycles ranging from 30/22 C to 26/22 C optimized net photosynthesis (Rowe and others 1994c). Similar
results were reported for flame azalea (Malek and others
1992b). Throughout propagation and subsequent culture,
plants should be examined frequently for insect and disease
problems. Rhododendrons can be raised successfully with
proper handling of the tender and delicate young seedlings
by using a porous, well-drained acidic medium high in
organic matter, and by maintaining ample moisture at all
times.
References
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Anderson A, Anderson S. 1994. How to grow rhododendrons from seed.
Journal of the American Rhododendron Society 48(1): 10.
Arocha LO, Blazich FA, Warren SL,Thetford M, Berry JB. 1999. Seed germination of Rhododendron chapmanii: influence of light and temperature.
Journal of Environmental Horticulture 17(4): 193196.
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
Blazich FA, Warren SL, Acedo JR, Reece WM. 1991. Seed germination of
Rhododendron catawbiense and Rhododendron maximum: influence of
light and temperature. Journal of Environmental Horticulture 9(1): 58.
Blazich FA, Warren SL, Starrett MC, Acedo JR. 1993. Seed germination of
Rhododendron carolinianum: influence of light and temperature. Journal of
Environmental Horticulture 11(2): 5558.
Bowers CG. 1960. Rhododendrons and azaleas. 2nd ed. New York:
Macmillan. 525 p.
Cho MS, Jung JH,Yeam DY. 1981. Studies on seed germination of rhododendron plants. Journal of the Korean Society of Horticultural Science
22: 107120.
Cox PA. 1990. The larger rhododendron species. Portland, OR:Timber
Press. 389 p.
Davidian HH. 1992. The Rhododendron species.Volume 3. Elepidotes
continued, Neriiflorum-Thomsonii, Azaleastrum and Camtschaticum.
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Dirr MA, Heuser Jr CW. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
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Gibson WH. 1901. Blossom hosts and insect guests. New York: Newson
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Glenn CT, Blazich FA, Warren SL. 1998. Influence of storage temperatures
on long-term seed viability of selected ericaceous species. Journal of
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Glenn CT, Blazich FA, Warren SL. 1999. Secondary seed dormancy of
Rhododendron catawbiense and Rhododendron maximum. Journal of
Environmental Horticulture 17(1): 14.
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Leach DG. 1961. Rhododendrons of the world and how to grow them.
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LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
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Malek AA, Blazich FA, Warren SL, Shelton JE. 1989. Influence of light and
temperature on seed germination of flame azalea. Journal of
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Malek AA, Blazich FA, Warren SL, Shelton JE. 1990. Influence of light and
temperature on seed germination of flame azalea. Journal of the
American Rhododendron Society 44(4): 215217.
Malek AA, Blazich FA, Warren SL, Shelton JE. 1992a. Growth response of
seedlings of flame azalea to manual and chemical pinching. Journal of
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Malek AA, Blazich FA, Warren SL, Shelton JE. 1992b. Initial growth of
seedlings of flame azalea in response to day/night temperature. Journal
of the American Society for Horticultural Science 117(2): 216219.
McCown BH, Lloyd GB. 1983. A survey of the response of Rhododendron
to in vitro culture. Plant Cell,Tissue and Organ Culture 2: 7785.
Odenwald N,Turner J. 1987. Identification, selection, and use of southern
plants for landscape design. Baton Rouge, LA: Claitors Publishing. 660 p.
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dissertation]. Durham, NC: Duke University. 189 p.
Rowe DB, Blazich FA, Warren SL, Ranney TG. 1994a. Seed germination of
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temperature. Journal of Environmental Horticulture 12(3): 155158.
Rowe DB, Warren SL, Blazich FA. 1994b. Seedling growth of Catawba rhododendron: 11.Temperature optima, leaf area, and dry weight distribution. HortScience 29(11): 12981302.
Rowe DB, Warren SL, Blazich FA, Pharr DM. 1994c. Seedling growth of
Catawba rhododendron: II. Photosynthesis and carbohydrate accumulation and export. HortScience 29(11): 13031308.
Toole EH,Toole VK, Borthwick HA, Hendricks SB. 1955. Interaction of
temperature and light in germination of seeds. Plant Physiology 30:
473478.
Wells JS. 1985. Plant propagation practices. Chicago: American
Nurseryman Publishing. 367 p.
Rhododendron
951
RosaceaeRose family
952
fruit.
References
Rhodotypos
953
AnacardiaceaeSumac family
Rhus L.
sumac
D. Bradley Rowe and Frank A. Blazich
Dr. Rowe is associate professor at Michigan State Universitys Department of Horticulture, East Lansing,
Michigan; Dr. Blazich is alumni distinguished graduate professor of plant propagation and tissue culture at
North Carolina State Universitys Department of Horticultural Science, Raleigh, North Carolina
Table 1Rhus, sumac; Toxicodendron, poison-ivy, etc.; Malosma, laurel-sumac: nomenclature and occurrence
Scientific names & synonym(s)
Common name(s)
Occurrence
R. aromatica Ait.
R. canadensis Marsh.
R. choriophyllum Woot. & Standl.
R. copallina L.
R. glabra L.
Schmaltzia glabra Small
R. borealis Greene
R. hirta (L.) Sudworth
R. typhina L.
R. integrifolia (Nutt.) Benth. & Hook.
f. ex Brewer & S.Wats.
R. kearneyi Barkl.
R. lanceolata (Gray) Britt.
R. copallina var. lanceolata Gray
R. michauxii Sarg.
Schmaltzia michauxii M. Small
R. microphylla Engelm. ex Gray
R. ovata S.Wats.
R. ovata var. traskiae Barkl.
R. trilobata Nutt.
Schmaltzia anisophylla Greene
S. trilobata var. anisophylla (Greene) Barkl.
R. virens Lindheimer ex Gray
SW US
poison-oak
poison-ivy
RELATED TAXA
Toxicodendron diversilobum
(Torr. & Gray) Greene
R. diversiloba Torr. & Gray
R. toxicodendron ssp. diversilobum
Torr. & A. Gray) Engl.
T. radicans ssp. radicans (L.) Kuntze
R. radicans L.; R. toxicodendron L.
T. vernix (L.) Kuntze
R. vernix L.
Malosma laurina (Nutt.) Nutt. ex
Abrams
R. laurina Nutt.
Sources:
Rhus
955
Table 2Rhus, sumac; Toxicodendron, poison ivy, etc.; Malosma, laurel-sumac: growth habit, flowers, and fruits
Speicies
Flowers
Fruits
R. aromatica
Shrub to 2.5 m
R. choriophylla
R. copallina
R. glabra
Shrub or tree to 6 m
R. hirta
R. integrifolia
Shrub or tree to 9 m,
twigs densely pubescent
Evergreen shrub or tree to 9 m
R. kearneyi
R. lanceolata
Thicket-forming shrub
or small tree to 10 m
R. michauxii
R. microphylla
R. trilobata
Shrub, to 2 m, rarely
treelike to 5 m
Evergreen shrub to 3 m,
rarely a tree to 4.5 m
Shrub to 2 m
R. virens
Shrub
R. ovata
RELATED TAXA
T. diversilobum
T. radicans ssp.
radicans
T. vernix
Greenish, in slender
panicles 820 cm long
M. laurina
Shrub, 36 m
Whitish
Whitish, berrylike 56 mm
across, in axillary clusters; early
summer, persisting into winter
Greenish white in pendent
axillary panicles to 20 cm long;
pedicles persist through winter
Whitish; early summer
and empty seeds. Such thorough cleaning is seldom practiced except for skunkbush; seeds of other species are sown
with pieces of the fruit wall still attached (Brinkman 1974).
Trials have shown that about 99% of the empty seeds of
smooth sumac can be removed by flotation, as empty seeds
float and filled ones sink (Johnson and others 1966).
However, the flotation method of separating empty seeds is
not always successful with seeds of staghorn sumac
(Brinkman 1974). Number of seeds per unit weight and seed
yields vary among species (table 3).
Storage. Seeds of sumac are orthodox in storage
behavior and can be stored over winter and possibly for
years without special treatment (Dirr and Heuser 1987).
Seeds of smooth sumac stored at room temperature for 10
956
longitudinal
Rhus
957
Species
R. copallina
R. glabra
R. hirta
R. integrifolia
R. ovata
R. trilobata
M. laurina
50.6105.6
66.0
6.6
37.4
15.419.8
198.0
23.048.0
30.0
3.0
17.0
7.09.0
90.0
/kg
81.4173.8
52.8277.2
107.1148.7
15.017.6
41.157.2
23.366.0
125.4
107.8
117.3
16.7
44.7
285.1
57.0
49.0
53.3
7.6
20.3
129.6
Samples
4
28
5
2
2
9
1
Figure 4Rhus hirta, staghorn sumac: seedling development at 2, 4, and 17 days after germination.
Rhus
959
References
Brinkman KA. 1974. Rhus L., sumac. In: Schopmeyer CS, tech. coord. Seeds
of woody plants in the United States. Agric. Handbk. 450. Washington,
DC: USDA Forest Service: 715719.
Cross RE Sr. 1982. Propagation and production of Rhus typhina Laciniata,
cutleaf staghorn sumac. Combined Proceedings of the International Plant
Propagators Society 31: 524527.
Cross RE Sr. 1988. Persistence pays: a Minnesota nursery spent years refining its propagation method for cutleaf staghorn sumac but says it was
well worth the effort. American Nurseryman 168(12): 63, 6567.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Duncan WH. 1935. Root systems of woody plants of old fields of Indiana.
Ecology 16: 554567.
Elias TS. 1989. Field guide to North American trees. 2nd ed. Danbury, CT:
Grolier Book Clubs. 948 p.
Farmer RE, Lockley GC, Cunningham M. 1982. Germination patterns of the
sumacs, Rhus glabra and Rhus coppalina: effects of scarification time, temperature and genotype. Seed Science and Technology 10(2): 223231.
Gill JD, Healy WH. 1974. Shrubs and vines for northeastern wildlife. Gen.
Tech. Rep. NE-9. Broomall, PA: USDA Forest Service Northeastern
Forest Experiment Station. 180 p.
Hampton CO, Singh SP. 1979. The presence of growth and germination
inhibitors in the seeds of certain desert plants. Kansas Academy of
Science Transactions 82(2): 87.
Hartmann HT, Kester DE, Davies FT Jr, Geneve RL. 2002. Hartmann and
Kesters plant propagation: principles and practices. 7th ed. Upper Saddle
River, NJ: Prentice-Hall. 880 p.
Haywood JD. 1994. Seed viability of selected tree, shrub, and vine species
stored in the field. New Forests 8(2): 143154.
Heit CE. 1967. Propagation from seed: 7. Successful propagation of six
hardseeded group species. American Nurseryman 125(12): 1012,
3741, 4445.
Hubbard AC. 1986. Native ornamentals for the U.S. Southwest. Combined
Proceedings International Plant Propagators Society 36: 347350.
Humphrey EG. 1983. Smooth sumac tested for growth on mine spoils.
USDA Soil Conservation Service 4(6): 8.
Johnson AG, Foote LE, Smithberg MH. 1966. Smooth sumac seed
germination. Plant Propagator 12(3): 58.
Jonsson GB, Zak JM. 1975. Propagation of sumac (Rhus) species for
Massachusetts roadsides. American Nurseryman 142(6): 1415,
24, 26, 28.
960
Krugman SL, Stein WI, Schmitt DM. 1974. Seed biology. In: Schopmeyer CS,
tech. coord. Seeds of woody plants in the United States. Agric. Handbk
450. Washington, DC: USDA Forest Service: 5-40.
Lee JJ, Weber DE. 1979. The effect of simulated acid rain on seedling emergence and growth of eleven woody species. Forest Science 25(3):
393398.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. 3rd ed.
New York: Macmillan. 1290 p.
Matveev NM, Krisanov GN, Lyzhenko II. 1975. The role of plant exudates in
formation of herbaceous cover under Robinia pseudoacacia and sumac
protective tree strips in the steppe zone. Nauchnye Doklady Vysshei
Shkoly, Biologicheskie Nauki 10: 8084.
Norton CR. 1985. The use of gibberellic acid, ethephon and cold treatment
to promote germination of Rhus typhina L. seeds. Scientia Horticulturae
27: 163169.
Norton CR. 1986. Seed germination of Rhus typhina L. after growth
regulator treatment. Plant Propagator 32(2): 5.
Norton CR. 1987. Seed technology aspects of woody ornamental seed
germination. Acta Horticulturae 202: 2334
Rasmussen GA, Wright HA. 1988. Germination requirements of flameleaf
sumac. Journal of Range Management 41(1): 4852.
Rehder A. 1990. Manual of cultivated trees and shrubs hardy in North
America. 2nd ed. Portland, OR: Dioscorides Press. 996 p.
RHS [Royal Horticultural Society]. 1994. The new Royal Horticultural
Society dictionary index of garden plants. Griffiths M, ed. London:
Macmillan Press. 1234 p.
Smith NE. 1990. Water-soluble extracts from leaves of shining sumac inhibit
germination and radicle growth of loblolly pine.Tree Planters Notes
41(3): 3334.
Strauss SY. 1988. Determining the effects of herbivory using naturally
damaged plants. Ecology 69(5): 16281630.
Tipton JL. 1990. Vegetative propagation of Mexican redbud, larchleaf goldenweed, littleleaf ash, and evergreen sumac. HortScience 25(2): 196198.
Tipton JL. 1992. Requirements for seed germination of Mexican redbud,
evergreen sumac, and mealy sage. HortScience 27(4): 313316.
Washitani I. 1988. Effects of high temperature on the permeability, and
germinability of the hard seeds of Rhus javanica L. Annals of Botany 62:
1316.
Weber GP, Wiesner LE, Lund RE. 1982. Improving germination of
skunkbush sumac and serviceberry seed. Journal of Seed Technology
7(1): 6071.
GrossulariaceaeCurrant family
Ribes L.
currant, gooseberry
Robert D. Pfister and John P. Sloan
Dr. Pfister is a professor of forestry at the University of Montana, Missoula;
Mr. Sloan is at the USDA Forest Services Lucky Peak Nursery, Boise, Idaho
Growth habit, occurrence, and use. The currant and
gooseberry genusRibes includes about 150 species of
deciduous, (rarely) evergreen, shrubs that grow in the colder
and temperate parts of North America, Europe, Asia, and
South America. The unarmed species are commonly called
currants; the prickly species are gooseberries. Of the more
important species for which seed data are available, 16 are
native to the United States and 1 was introduced from
Europe (table 1). These species generally occur as rather
low-growing shrubs, although 3 species can attain heights
of 3 to 4 m (table 2).
Six of the more showy speciesalpine, American
black, golden, wax, clove, and winter currantsare cultivated for their colorful fruit, attractive flowers, and ornamental
foliage. Berries are made into jam, jelly, pie, juice, and
syrup. All native species are valuable as food and cover for
wildlifeand many provide browse for livestock (Plummer
and others 1968). Golden and clove currants have been used
in shelterbelt plantings in the prairieplains and intermountain regions. The former also has been widely planted for
erosion control (Pfister 1974). Golden, wax, white-stem,
and gooseberry currants are valued as ornamentals in the
United States and Canada (Barnes 1986). Currants are
shade tolerant (Quick 1954). Many species regenerate vegetatively as well as from seed (Dittberner and Olson 1983;
Wasser 1982). Most are rhizomatous (Lotan and others
1981). Seeds of currants remain viable in soil for long periods of time (Lyon and Stickney 1976).
Germination is stimulated by disturbances such as fire
(Lotan and others 1981; Morgan and Neuenschwander
1985; Young 1983). Consequently, currants are common
pioneer species on hot burns occurring on xeric sites
(Hopkins and Kovalchik 1983). However, their seedcoats
are relatively thin and may be destroyed by severe fires.
Moist mineral soil with high amounts of humus provides a
good seedbed for currants. Seeds are often introduced to the
seedbank by birds and mammals that cannot digest the
Ribes
961
Common name(s)
Occurrence
R. alpinum L.
R. opulifolium L.
R. americanum P. Mill.
R. floridum LHerit.
R. aureum Pursh
Chrysobotrya aurea (Pursh) Rydb.
R. flavum Colla; R. tenuiflorum Lindl.
R. aureum var. villosum DC.
R. odoratum H.Wendl.
Chrysobotrya odorata (Wendl.) Rydb.
R. cereum Dougl.
R. churchii A. Nels & Kenn.
R. inebrians Lindl.; R. pumilum Nutt.
R. cynosbati L.
Grossularia cynosbati (L.) Mill.
R. gracile Michx.
R. hudsonianum Richards.
R. petiolare Dougl.
R. inerme Rydb.
Grossularia inermis (Rydb.) Cov. & Britt.
R. divaricatum Dougl. var. inerme (Rydb.) McMinn
R. purpusii Koehne ex Blank.
R. lacustre (Pers.) Poir.
Limnobotrya lacustris Rydb.
R. echinatum Dougl.; R. grossulariodes Michx.
R. parvulum Rydb.
R. missouriense Nutt.
Grossularia missouriensis (Nutt). Cov. & Britt.
R. gracile Pursh, not Michx.
R. montigenum McClatchie
Limnobotrya montigena McClatchie Rydb.
R. lacustre var. molle Gray.
R. lentum Cov. & Rose; R. molle Howell
R. nevadense Kellogg
R. ascendens Eastw.; R. grantii Heller
R. oxyacanthoides ssp. irriguum
(Dougl.) Sinnott
R. irriguum Dougl.
R. divaricatum var. irriguum (Dougl.) Gray
Grossularia irrigua (Dougl.) Cov. & Britt.
R. roezlii Regel
Grossularia roezlii (Regel) Cov. & Britt.
R. amictum Greene; R. aridum Greene
R. urlsonianum Greene
R. rotundifolium Michx.
Grossularia rotundifolia (Michx.) Cov. & Britt.
R. triflorum Willd.
R. sanguineum Pursh
Calobotrya sanguinea (Pursh) Spach
Coreosma sanguinea (Pursh) Spach
R. glutinosum Benth.
R. viscosissimum Pursh
Coreosma viscosissima (Pursh) Spach
R. halli Jancz.
alpine currant
Europe to Siberia
962
golden currant,
slender golden currant,
flowering currant
clove currant,
buffalo currant
wax currant,
squaw currant
pasture gooseberry,
eastern prickly gooseberry
prickly currant,
swamp gooseberry,
swamp black currant
Alaska to Newfoundland, S to
California, South Dakota, & Pennsylvania
Missouri gooseberry
Minnesota to Connecticut, S
to Tennessee, Arkansas, & Kansas
gooseberry currant,
alpine prickly currant,
mountain gooseberry
Sierra currant
Idaho gooseberry,
inland black gooseberry
British Columbia, S to NE
Oregon & E to W Montana
Sierra gooseberry
roundleaf gooseberry,
Appalachian gooseberry
winter currant,
red flowering currant,
Oregon currant,
blood currant
sticky currant
Table 2Ribes, currant, gooseberry: growth habit, height at maturity and year of first cultivation
Species
Growth habit
R. alpinum
R. americanum
R. aureum
R. aureum var. villosum
R. cereum
R. cynosbati
R. hudsonianum
R. inerme
R. lacustre
R. missouriense
R. montigenum
R. nevadense
R. oxyacanthoides spp. irriguum
R. roezlii
R. rotundifolium
R. sanguineum
R. viscosissimum
Height at
maturity (m)
Year first
cultivated
0.92.4
0.61.8
0.93.0
0.93.0
0.31.5
1.5
0.31.8
0.92.1
0.31.8
0.31.8
0.30.9
0.91.8
0.32.4
0.61.5
0.9
0.93.6
0.31.8
1588
1727
1806
1812
1827
1759
1899
1899
1812
1907
1905
1907
1920
1899
1809
1818
1827
Location
Fruit ripening
Flowering
R. alpinum
R. americanum
R. aureum
R. aureum var.villosum
Europe
Wyoming
Kansas
Oregon
AprMay
AprJune
AprMay
Late May
MidApr
AprJune
AprJune
Aprearly June
MayJuly
MayJune
AprJuly
AprMay
Late JuneJuly
MayJuly
AprJune
MayJune
AprMay
AprMay
MarJune
MayJune
JulyAug
JuneSept
JuneJuly
Late Aug
June
JuneAug
Aug
Late JulySept
Aug
JuneSept
AugSept
JulySept
JulyAug
AugSept
R. cereum
R. cynosbati
R. hudsonianum
R. inerme
R. lacustre
R. missouriense
R. montigenum
R. nevadense
R. oxyacanthoides ssp. irriguum
R. roezlii
R. rotundifolium
R. sanguineum
R. viscosissimum
Sources: Fernald (1950), Hitchcock and others (1955), Krssmann (19601962), Loiseau (1945), Munz and Keck (1965), NBV (1946), Petrides (1955), Pfister (1974),
Rehder (1940), Stephens (1969), Steyermark (1963), Symonds (1963),Wyman (1949).
963
Table 4Ribes, currant, gooseberry: fruit characteristics and seed storage conditions for air-dried seeds
Species
Surface
R. alpinum
R. americanum
R. aureum
R. aureum var. villosum
R. cereum
R. cynosbati
R. hudsonianum
R. inerme
R. lacustre
R. missouriense
R. montigenum
R. nevadense
Glabrous
Glabrous
Glabrous
Smooth
Glandular
Glandular
Smooth
Smooth
Glandular
Smooth
Glandular
Glandular
Glandular
Smooth
Glandular
Glandular
Smooth
Glandular
Glandular
Fruit characteristics
Diam (cm)
Ripe color
0.6
0.6
1.0
0.6
1.0
0.6
0.6
1.3
0.6
1.0
1.3
1.3
0.6
1.0
1.3
Scarlet
Black
Red, black, or yellow
Black, golden, or reddish brown
Dull to bright red
Reddish purple
Black
Reddish purple
Purple to black
Purple to black
Red
Blue to black
Blue to black
Bluish purple
Purple or deep reddish brown
Purple or deep reddish brown
Purple
Blue to black
Black
Temp
(C)
6
21
21
21
21
21
21
Soil
21
Soil
2
21
21
Storage conditions
Duration
Viability
(yr)
at end (%)
4
17
17
27
7
17
11
4
4
13
12
17
22
38
89
32
4
8
40
80
81
88
82
45
23
7
Sources: Hitchcock (1955), Jepson (1925), Ketchum and others (1968), Munz (1965), Pfister (1974), Quick (1945, 1947, 1954), Rehder (1940), Stephens (1969).
964
(4 lb) for golden currant, 3.6 kg (8 lb) for clove currant, and
1.8 kg (4 lb) for winter currant (Pfister 1974). One liter of
berries from winter currant weighs about 0.5 kg (1 bu
weighs about 40 lb). Each prickly currant plant produces
around 50 to 75 berries, and each berry has 8 seeds (Moss
and Wellner 1953).
Storage. Currant seeds are orthodox and remain
viable for long periods when stored in sealed containers at a
low moisture content. Temperature is evidently not critical.
Samples of Sierra gooseberry seeds buried in soil in inverted
open containers for 13 years exhibited 70 to 94% viability
(Quick 1947b). Seeds of several species stored dry at room
temperature also maintained high viability for periods up to
17 years (table 4).
Germination. In nature, currant seeds normally germinate in spring following dispersal, although some seeds
may remain dormant for many years (Moss and Wellner
1953; Quick 1954). The best seedbed appears to be mineral
soil well supplied with humus. Germination is epigeal (figure 4). In the laboratory, seeds are slow to germinate except
for those of Hudson Bay currant and roundleaf gooseberry.
Most species require at least 1 stratification period of fairly
long duration to break embryo dormancy (Rudolf 1949).
Stidham and others (1980) achieved good germination of
golden currant after 10 weeks of wet chilling in distilled
water. Impermeable seedcoats also appear to be involved in
dormancy of some seedlots of clove and American black
currants (Pfister 1974). Germination rate and total can be
increased by wet prechilling in sand, peat, or vermiculite or
in a mixture of these media. Seed losses from damping-off
fungi can be prevented by applying 646 mg of copper
Species
Place
collected
R. americanum
R. aureum
R. aureum var. villosum
R. cereum
R. cynosbati
R. hudsonianum
R. inerme
R. lacustre
R. missouriense
R. montigenum
R. nevadense
R. roezlii
R. sanguineum
R. viscosissimum
North Dakota
California
Idaho
Idaho & California
California
Utah
California
California
Oregon
Idaho & California
Range
/kg
544741
441628
234395
443624
417487
1,3892,703
780877
344370
650935
388650
562769
156168
295424
176295
255344
690
514
368
553
452
2,127
807
1,135
357
313
862
520
626
657
Samples
313
233
167
251
205
965
366
515
162
142
391
236
284
298
4
4
8
5
2
12
5
1
2
1
10
10+
1
5
Ribes
965
Table 6Ribes, currant, gooseberry: pregermination treatments and germination test results
Pregermination
treatment
Temp
(C)
Species
R. alpinum
R. americanum
R. aureum
R. aureum var. villosum
R. cereum
R. cynosbati
R. hudsonianum
0 to 10
2 to 2
2 to 2
20/0 (D/N)
2 to 0
2 to 5
NP
0 to 2
R. inerme
0
R. lacustre
0
R. missouriense
2 to 5
R. montigenum
0
0
R. nevadense
0
R. oxyacanthoides ssp. irriguum
0 to 5
R. roezlii
0
R. rotundifolia
2 to 0
R. sanguineum
02
R. viscosissimum
2 to 0
Days
90+
90120
60
120
120150
90150
NP
90120
120200
120200
90+
200300
120150
120
90
100150
90+
100140
140
Germination
under test
conditions*
(%)
80
68
60
94
61
69
57
69
60
48
73
53
8
78
79
80
80
61
58
Germination
capacity
(%)
Samples
76
63
98
72
72
85
76
74
61
33
87
81
87
81
64
67
10
39
19
3
61
19
116
42
54
64
3
6
15
43
11
200
10
55
88
Sources: NBV (1946), Pfister (1974), Quick (1939, 1940, 1941, 1943, 1945, 1947).
Note: D/N = day/night, NP = no pretreatment.
* Virtually all of the tested seeds were stratified and germinated in sand moistened with nutrient solution.The germination tests were conducted under greenhouse conditions for periods of 30 to 40 days.
Germination capacity was determined by retrial stratification and a repeat germination test with conditions about the same as used initially.
966
are sown because fresh seeds will not germinate, even after
stratification (Quick 1939). If fall-sowing is not possible, the
seeds should be stratified before spring-sowing using the
procedures summarized in table 6. Seeds should be sown at
a rate of 646 to 860/m2 (60 to 80/ft2) (NBV 1946) or 130
viable seeds/m of row (40/ft) and covered to a depth of 3 to
6 mm (1/8 to 1/4 in) (Pfister 1974). Seeds of Sierra gooseberry and wax and Sierra currants may be covered up to 1.3 cm
(1/2 in) (Quick 1939a, 1940).
The only reported experience in nursery stock production is for clove currant (Pfister 1974). Seedbeds are fallsown, mulched to a depth of 5 to 8 cm (2 to 3 in) and covered with snow fence. About 20,000 seedlings are produced
per kilogram of seeds (9,000/lb) and the normal outplanting
age is 2 years. Most species can be propagated readily from
hardwood cuttings taken in autumn (Pfister 1974).
References
Barnes HW. 1986. Five Ribes species of the U.S. and Canada Rocky
Mountains. Plant Propagator 32(2): 910.
Blasse W, Hofman S. 1988. Studies on the biology of fertilization in Ribes
rubrum L. and Ribes nigrum L. Archiv fr Gartenbau 36(7): 437448.
Dittberner PL, Olson MR. 1983. The plant network data base (PIN):
Colorado, Montana, North Dakota, and Wyoming. FWS/OBS-83/86.
Washingotn, DC: USDI Fish and Wildlife Service. 786 p.
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1632 p.
Fivaz AE. 1931. Longevity and germination of seeds of Ribes, particularly R.
rotundifolium, under laboratory and natural conditions.Tech. Bull. 261.
Washington, DC: USDA. 40 p.
Heisey RM. 1982. Allelopathic effects of Trichostema lanceolatum (Labiatae)
[PhD thesis]. Davis: University of California. 156 p.
Heit CE. 1968. Propagation from seed: 15. Fall planting of shrub seeds for
successful seedling production. American Nurseryman 128(4): 810.
Hitchcock CL, Cronquist A, Ownbey M,Thompson JW. 1955. Vascular
plants of the Pacific Northwest: 3. Saxifragaceae to Ericaceae. Seattle:
University of Washington Press. 614 p.
Hopkins WE, Kovalchik BL. 1983. Plant associations of the Crooked River
National Grassland. R6-Ecol 133-1983. Portland, OR: USDA Forest
Service, Pacific Northwest Region. 98 p.
Jepson WL. 1925. A manual of the flowering plants of California. Berkeley,
CA: Associated Students Store. 1238 p.
Ketcham DE, Wellner CA, Evans SS Jr. 1968. Western white pine management programs realigned on northern Rocky Mountain National
Forests. Journal of Forestry 66: 329332.
Krssmann G. 19601962. Handbuch der Laugbehlze. 2 volumes. Berlin:
Parey. 495 & 608 p.
Loiseau J. 1945. Les arbres et la fort.Volume 1. Paris:Vigot. 204 p
Lotan JE, Alexander ME, Arno SF, French RE, Langdon OG, Loomis RM,
Norum RA, Rothermel RC, Schmidt WC,Van Wagtendonk JW. 1981.
Effects of fire on flora: a state-of-knowledge review. Gen.Tech. Rep.
WO-16. Washington, DC: USDA Forest Service. 71 p.
Lyon JL, Stickney PF. 1976. Early vegetal succession following large northern
Rocky Mountain wildfires. In: Proceedings,Tall Timbers Fire Ecology
Conference #14 and Intermountain Fire Council and Land Management
Symposium; 1974 October 810; Missoula, MT. Tallahassee, FL:Tall
Timbers Research Station 14: 355375.
Miller DR. 1931. Ribes seed germination studies. In: Blister rust control
work in the Far West. Annu. Rep. 1931. Washington, DC: USDA Bureau
of Entomology and Plant Quarantine, Division of Blister Rust Control.
431 p.
Ribes
967
Quick CR. 1947a. Experimental germination of Ribes and pine seeds, 1945.
Serial 135. Berkeley, CA: USDA Bureau of Entomology and Plant
Quarantine. 35 p.
Quick CR. 1947b. Some experimental aspects of Ribes seed longevity. Serial
137. Berkeley, CA: USDA Bureau of Entomology and Plant Quarantine.
8 p.
Quick CR. 1954. Ecology of the Sierra Nevada gooseberry in relation to
blister rust control. Circ. 937. Washington, DC: USDA. 30 p.
Rehder A. 1940. Manual of cultivated trees and shrubs. 2nd ed. New York:
Macmillan. 2996 p.
Rudolf PO. 1949. First the seed, then the tree. In: USDA Yearbook of
Agriculture (1948):Trees. 127135.
Stephens HA. 1969. Trees, shrubs, and woody vines in Kansas. Lawrence:
University Press of Kansas. 250 p.
Steyermark JA. 1963. Flora of Missouri. Ames: Iowa State University Press.
1725 p.
968
Stidham ND, Ahring RM, Claypool PL. 1980. Chemical scarification, moist
prechilling, and thiourea effects on germination of 18 shrub species.
Journal of Range Management 33(2): 115118.
Symonds GWD. 1963. The shrub identification book. New York: M. Barrows
and Co. 379 p.
Wasser CH. 1982. Ecology and culture of selected species. In: Revegetating
Disturbed Lands in the West. FWS/OBS-82/56. Washington, DC: USDI
Fish and Wildlife Service: 263265.
Wyman D. 1949. Shrubs and vines for American gardens. New York:
Macmillan. 442 p.
Young RP. 1983. Fire as a vegetation management tool in rangelands of the
Intermountain region. In: Monsen SB, Shaw N, eds. Managing
Intermountain Rangelands: Improvement of Range and Wildlife Habitats.
Gen.Tech. Rep. INT-157. Ogden, UT. USDA Forest Service,
Intermountain Forest and Range Experiment Station: 1831.
FabaceaePea family
Robinia L.
locust
David F. Olson, Jr., and Robert P. Karrfalt
Dr. Olson retired from the USDA Forests Southeastern Forest Experiment Station; Mr. Karrfalt is director of
the USDA Forest Services National Seed Laboratory, Dry Branch, Georgia
Robinia
969
Common name(s)
Occurrence
locust
North Carolina
bristly locust
bristly locust
Kelsey locust
bristly locust
mossy locust,
bristly locust
Holdt locust
locust
Margarett locust
Rusby locust
black locust
clammy locust
Hartweg locust
BONAP (1996).
970
971
Flowering
Fruit ripening
Seed dispersal
R. hispida
R. hispida var. fertilis
R. neomexicana
R. pseudoacacia
MayJune
Early June
JulySept
Sept
Sept
SeptOct
OctNov
SeptOct
SeptApr
9
6.815
20
1533
Cleaned seeds/weight
/kg
/lb
50,715
61,080
47,630
52,920
23,000
27,700
21,600
24,000*
972
a problem along highways and in dry countries where irrigation must be practiced in agroforestry applications. Under
salinity levels of 0.05 to 0.80%, black locust germination
was reduced and occurred more slowly (Bangash 1977;
Bicknell and Smith 1975).
References
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
Bangash SH. 1977. Salt tolerance of forest tree species as determined by
germination of seeds at different salinity levels. Pakistan Journal of
Forestry 27(2): 9397.
Bicknell SH, Smith WH. 1975. Influence of soil salt, at levels characteristic of
some roadside environments, on the germination of certain tree seeds.
Plant and Soil 43(3): 719722.
BONAP [Biota of North America Project]. 1996. The digital checklist of the
vascular flora of North America [website available at
http://www.bonap.org].
Brown JH. 1973. Site factors and seeding methods affecting germination
and survival of tree species direct-seeded on surface-mined areas. Bull.
620. Morgantown: West Virginia University Agricultural Experiment
Station. 25 p.
Chaney WR, Kozlowski TT. 1974. Effects of antitranspirants on seed germination, growth, and survival of tree seedlings. Plant and Soil 40(1):
225229.
Chapman AG. 1936. Scarification of black locust seed to increase and
hasten germination. Journal of Forestry 34: 6674.
Cox RA, Klett JE. 1984. Seed germination requirements of native Colorado
plants for use in the landscape. Plant Propagator 30(2): 610.
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1632 p.
Heit CE. 1967. Propagation from seed: 6. Hard seedednessa critical
factor. American Nurseryman 125 (10): 1012, 8896.
Heit CE. 1968. Thirty five years testing of tree and shrub seed. Journal of
Forestry 66: 632634.
ISTA [International Seed Testing Association]. 1993. Rules for testing seeds:
rules. Seed Science and Technology 21 (Suppl.): 1259.
Karrfalt RP. 1990. Personal observation. Dry Branch, GA: USDA Forest
Service, National Tree Seed Laboratory.
Little EL Jr, Delisle AL. 1962. Time periods in development: forest trees of
North America, table 104. In: Biological handbook on growth. Altman PL,
Dittmer D, eds. Washington, DC: Federation of American Societies for
Experimental Biology.
McWilliams JL. 1970. Arnot bristly locust. Harrisburg, PA: USDA Soil
Conservation Service. 9 p.
Meginnis HG. 1937. Sulphuric acid treatment to increase germination of
black locust seed. Agric. Circ. 453. Washington, DC: USDA. 34 p.
Myatt A. 1991. Personal communication. Washington, OK: Oklahoma
Department of Agriculture, Forestry Division.
Olson DF. 1974. Robinia, locust. In: Schopmeyer CS, tech coord. Seeds of
woody plants in the United States. Agric. Handbk. 450. Washington, DC:
USDA Forest Service: 728731.
Radford AE, Ahles HE, Bell CR. 1964. Guide to the vascular flora of the
Carolinas. Chapel Hill: University of North Carolina Book Exchange,
383 p.
Roach BA. 1965. Black locust (Robinia pseudoacacia L.). In: Burns RM,
Honkala BH, eds. Silvics of forest trees of the United States. Agric.
Handbk. 271. Washington, DC: USDA Forest Service: 642648.
Robertson C. 1928. Flowers and insects. Lancaster, PA: Science Press. 221 p
Sargent CS. 1965. Manual of trees of North America (exclusive of Mexico).
2nd ed. New York: Dover. 934 p.
Singh DP, Hooda MS, Bonner FT. 1991. An evaluation of scarification methods for seeds of two leguminous trees. New Forests 5(2): 139145.
Small JK. 1933. Manual of the southeastern flora. New York: J.K. Small.
1554 p.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP 27. Washington, DC: USDA Soil
Conservation Service. 198 p
Whitaker TW. 1934. A karyo-systematic study of Robinia. Journal of the
Arnold Arboretum 15: 353357.
Wilson JK. 1944. Immersing seeds of species of Robinia in boiling water hastens germination. Journal of Forestry 42: 453454.
Wooten EO. 1913. Trees and shrubs of New Mexico. Bull. 87: State
College: New Mexico Agricultural Experiment Station. 159 p.
Wyman D. 1953. Seeds of woody plants. Arnoldia 13: 4160.
Robinia
973
RosaceaeRose family
Rosa L.
rose, briar
Susan E. Meyer
Dr. Meyer is a research ecologist at the USDA Forest Services Rocky Mountain Research Station
Shrub Sciences Laboratory, Provo, Utah
and act as seed dispersers (Gill and Pogge 1974). Wild roses
are also utilized as browse by many wild and domestic
ungulates. Rose hips are an excellent source of vitamin C
and may also be consumed by humans (Densmore and
Zasada 1977). Rose oil extracted from the fragrant petals is
an important constituent of perfume. The principal use of
roses has clearly been in ornamental horticulture, and most
of the species treated here have been in cultivation for many
years (Gill and Pogge 1974).
Many roses are pioneer species that colonize disturbances naturally. The thicket-forming species especially
have potential for watershed stabilization and reclamation of
disturbed sites. If roses are to be used for these purposes, it
is greatly preferable to utilize species native to the region
rather than exotics, which can become serious pests. An
Table 1Rosa, rose: scientific names and geographic distribution for 12 species native or naturalized in the United States
Scientific name
Common name(s)
Geographic distribution
R. acicularis Lindl.
prickly rose
R. blanda Ait.
meadow rose,
smooth rose
California rose
dog rose
974
sweetbriar rose,
eglantine
baldhip rose,
dwarf rose
multiflora rose,
Japanese rose
Nootka rose
rugosa rose,
hedgerow rose
prairie rose,
climbing rose
wichura rose,
memorial rose
Woods rose
Figure 1Rosa, rose: fruits (hips) of R. eglanteria, sweetbriar rose (top); R. multiflora, multiflora rose (bottom left);
R. nutkana, Nootka rose (bottom center); and R. setigera,
prairie rose (bottom right).
Rosa
975
should either be refrigerated or spread out to dry, as otherwise they can overheat and the seeds become damaged. The
hips should be soaked in water if they have been allowed to
dry prior to processing, then macerated using a macerator or
similar device. Small lots can be macerated by rubbing the
hips through screens. The achenes may be separated from
the pulp by flotation or the material may be dried and the
achenes cleaned out using a fanning mill. Achene weights
vary from 5 to 17 mg (1.84 to 6.04 oz) and they number
59,530 to 185,220/kg (27,000 to 84,000/lb), depending on
species and seedlot (table 2). Rose seeds may have a limited
storage life, with some loss of viability in laboratory or
warehouse dry storage after as little as 2 to 3 years (Crocker
and Barton 1931; Gill and Pogge 1974), but they are almost
certainly orthodox in storage behavior. Seeds of Woods rose
have been reported to retain viability in open warehouse
storage for 15 years (Stevens and others 1981). Sealed storage of air-dried seeds at low temperature is recommended
(Gill and Pogge 1974).
Germination and seed testing. Rose seeds are normally dormant at maturity and require some form of pretreatment in order to germinate. Release from dormancy is a
complex process that may involve changes at the pericarp,
testa, and embryo levels. The degree of dormancy and the
principal level of dormancy control varies among species,
cultivars, seedlots, and even among hips within a single
bush. Because the achenes have a thick, hard pericarp and
do not swell when placed in water, it is often assumed that
they are water-impermeable. Work by Svejda (1972) and
others has shown that this is not the case. The achenes do
take up water, although the mechanical restriction presented
by the pericarp can sometimes prevent full imbibition.
Tincker and Wisley (1935) showed, for 10 rose species, that
longitudinal section
mg
2528
912
4
13 (817)
15
16
69
815
69
9
5
9 (713)
Achenes/weight
oz
0.91.0
0.30.4
0.1
0.5 (0.30.6)
0.5
0.6
0.20.3
0.30.5
0.20.3
0.3
0.2
0.3 (0.20.5)
Sources: Belcher (1985), Gill and Pogge (1974), Mirov and Kraebel (1939).
976
/kg
/lb
35,94040,130
81,580116,860
224,910
59,530119,070
68,355
61,740
110,250180,810
66,150132,300
114,660163,170
110,250
185,220
77,170143,320
16,30018,200
37,00053,000
102,000
27,00054,000
31,000
28,000
50,00082,000
30,00060,000
52,00074,000
50,000
84,000
35,00065,000
the sutures, whether with acid or through microbial digestion. Responsiveness to warm plus cold stratification can
also be increased in R. dumetorum Laxa by vacuum-infiltrating the achenes with growth hormones such as gibberellic acid or benzyladenine (Foster and Wright 1983), which
suggests that something other than simple mechanical
restriction may be involved. Similarly, in the relatively nondormant multiflora rose, the achenes may be induced to germinate without chilling either by treatment with macerating
enzymes that weaken pericarp sutures or by leaching with
activated charcoal to remove inhibitors from the incubation
solution (Yambe and Takeno 1992; Yambe and others 1992).
By using macerating enzymes to remove dormancy, these
workers were able to demonstrate a phytochrome-mediated
light requirement for germination in this species (Yambe and
others 1995). Acid scarification (but not mechanical scarification) is reported to substitute for warm pretreatment in the
cultivated rose R. gallica L. (Svejda 1968).
Chilling is the treatment most often applied to remove
rose seed dormancy, and the achenes of most species will
stratification
Days
Temp (C)
118
60
90
128
60
60
25
20
20
18.5
20
20
Cold stratification
Days
Temp (C)
365
90
90
270
90
60
150
570
450
90
90
180
120
365
128
128
90
90
210
120
90
60
45
120
90
5
5
5
5
5
4
4
5
5
5
5
5
5
4.5
4.5
4.5
3
3
4
5
4.4
5
5
3
3
Germination
temp (C)
5
20, 10/20
13, 18
13, 18
5
5
1518
1518
5
4.5
18.5
18.5
2029
2029
20
1518
18.3
1518
18.3
Incubation
(%)
57*
90*
7
53
62
47
34
24
40
43
45
60
72
65
48
72
32
60
85
90
48
75
76
0
49
Sources: Crocker and Barton (1931), Densmore and Zasada (1977), Gill and Pogge (1974), McTavish (1986), Mirov and Kraebel (1939), Rowley (1956), Semeniuk and
Stewart (1962, 1964, 1966), Stewart and Semeniuk (1965), Svejda (1968),Tillberg (1983),Tinker and Wisley (1935).
* Based on total viable seeds.
Total viability known to be about 55%; all other percentages based on total seeds, viability unknown.
Rosa
977
above the compensating temperature was to induce secondary dormancy at the embryo level. Interestingly, this dormancy could be alleviated only by chilling whole achenes;
chilling the embryos did not alleviate their dormancy.
Other species, such as prickly, Nootka, and Woods
roses, show much increased germination percentages in
response to chilling periods corresponding to a single winter
if the chilling period is preceded by a period of warm incubation (table 3). This requirement for warm incubation
before chilling would effectively postpone seedling emergence in the field until the second spring after seed production (Densmore and Zasada 1977). The temperature and
duration of the warm treatment is sometimes important. In
rugosa rose, a warm pretreatment of 60 days at 20 C before
90 days of chilling at 3 C increased germination over chilling alone, but longer periods resulted in decreased germination (Svejda 1968). The effect of warm pretreatment on
chilling response has been formally documented for only a
few rose species, but it is likely that high-viability lots of
any species that show minimal germination after 6 months
of chilling would be benefitted by a warm pretreatment.
Exactly what changes take place in rose seeds during
warm pretreatment or chilling is not known. In many cases,
the warm pretreatment seems to have effects at the seed
level rather than simply providing an opportunity for pericarp weakening (Densmore and Zasada 1977). Hormonal
balance has been implicated in the imposition of dormancy
in rose seeds by several workers. Substances leached from
dormant rose achenes or obtained from them by grinding
have been shown to suppress germination of otherwise nondormant excised rose embryos (Jackson and Blundell 1963,
1965; Svejda and Poapst 1972). Excised seeds with physically disrupted testas showed much lower germination than
embryos with testas removed, suggesting that inhibitors
leaching from the testa suppressed germination (Jackson and
Blundell 1963). Other workers have shown that, although
inhibitory substances are present in dormant achenes and
may disappear during dormancy loss, their removal alone is
not sufficient to induce germination (Julin-Tegelman 1983;
Tillberg 1983).
Variation in dormancy-breaking requirements both
within and among lots of any rose species make it difficult
to predict effective treatments. One of the causes of this
variation has been quite well-studied in cultivated tea roses,
and the results probably apply to wild species as well. Von
Abrams and Hand (1956) were the first to demonstrate that
seeds of a given cultivar matured in the field at warmer temperatures were less dormant (that is, had a shorter chilling
requirement) than seeds matured at cooler temperatures.
978
The preferred method in official testing is also tetrazolium staining (ISTA 1993), although stratification for 28 days
at 3 to 5 C is suggested for multiflora rose (AOSA 1993).
For other rose species, the international rules (ISTA 1993)
suggest an alternate method of 12 months of stratification,
followed by germination in sand at 20 C for 70 days.
Germination is epigeal (figure 4).
Field seeding and nursery practice. Woods rose has
been fall-seeded as a part of mixes for revegetation of deer
winter ranges in pinyon-juniper and mountain brush communities of the Intermountain West (Plummer and others
1968). It is recommended for areas with more than 300 mm
of annual precipitation, and should be broadcast-seeded or
drilled with other small-seeded shrubs at rates of 0.5 to 1
kg/ha (0.45 to 0.9 lb/ac). It reportedly is relatively easy to
establish from seeds and persists very well after initial
establishment. Other native rose species could probably also
be direct-seeded successfully in wildland settings.
Planting rose seeds in a nursery setting may be carried
out in fall for outdoor cold stratification or in summer for
warm followed by cold stratification. Seedlings will emerge
the following spring. For spring plantings, the achenes must
be appropriately stratified or otherwise pretreated prior to
planting. Recommended planting depth is 5 to 10 mm
(1/5 to 2/5 in), depending on seed size. Bareroot plants may
be produced successfully as 1+0 stock, and container stock
Figure 4Rosa blanda, meadow rose: seedling development at 1, 3, 6, 26, and 41 days after germination.
References
Anderson WL, Edminster FC. 1954. The multiflora rose for fences and
wildlife. Leaflet 374. Washington, DC: U.S. Department of Agriculture.
8 p.
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
Belcher E. 1985. Handbook on seeds of browse-shrubs and forbs.Tech.
Pub. R8-8. Atlanta: USDA Forest Service, Southern Region. 246 p.
Crocker W, Barton LV. 1931. Afterripening, germination, and storage of certain rosaceous seeds. Contributions to the Boyce Thompson Institute
3: 385404.
Cullum FJ, Bradley SJ, Williams ME. 1990. Improved germination of Rosa
corymbifera Laxa seed using a compost activator. Proceedings of the
International Plant Propagators Society 40: 244250.
Densmore R, Zasada JC. 1977. Germination requirements of Alaskan Rosa
acicularis. Canadian Field Naturalist 9: 5862.
DeVries DP, Dubois LAM. 1987. The effect of temperature on fruit set,
seed set, and seed germination in Sonia Hadley hybrid tea rose
crosses. Euphytica 36: 117-120.
Foster TC, Wright CJ. 1983. The germination of Rosa dumetorum Laxa.
Scientific Horticulture 34: 116125.
Gill JD, Pogge FL. 1974. Rosa L., rose. In: Schopmeyer CS, tech. coordinator.
Seeds of woody plants in the United States. USDA Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 732737.
Gudin S, Arene L, Chavagnat A, Bulard C. 1990. Influence of endocarp
thickness on rose achene germination: genetic and environmental factors. HortScience 25: 786788.
ISTA [International Seed Testing Association]. 1993. Rules for testing seeds:
rules 1993. Seed Science and Technology 21 (Suppl.): 1259.
Jackson GAD, Blundell JB. 1963. Germination in Rosa. Journal of
Horticultural Science 38: 310320.
Jackson GAD, Blundell JB. 1965. Germination of Rosa arvensis. Nature 205:
518519.
Rosa
979
Semeniuk P, Stewart RN. 1966. Effect of temperature and duration of afterripening period on germination of Rosa nutkana seeds. American Society
for Horticultural Science Proceedings 89: 689693.
Semeniuk P, Stewart RN, Uhring J. 1963. Induced secondary dormancy of
rose embryos. Proceedings of the American Society of Horticultural
Science 83: 825828.
Shaw N. 1984. Producing bareroot seedlings of native shrubs. In: Murphy
PM, comp.The challenge of producing native plants for the
Intermountain area. Gen.Tech. Rep. INT- 168. USDA Forest Service,
Intermountain Forest and Range Experiment Station: 615.
Stewart RN, Semeniuk P. 1965. The effect of the interaction of temperature with after-ripening requirements and compensating temperature on
germination of seeds of 5 species of Rosa. American Journal of Botany
52: 755760.
Stevens R, Jorgensen KR, Davis JN. 1981. Viability of seed from thirty-two
shrub and forb species through fifteen years of warehouse storage.
Great Basin Naturalist 41: 274277.
Svejda F. 1968. Effect of temperature and seed coat treatment on the
germination of rose seeds. HortScience 3: 184185.
Svejda FJ. 1972. Water uptake of rose achenes. Canadian Journal of Plant
Science 52: 10431047.
980
ArecaceaePalm family
Common name(s)
Occurrence
Sources: Francis (1992), Little (1979), Little and Wadsworth (1964), Neal (1965),West and Arnold (1952).
Roystonea
981
ers (Little and Wadsworth 1964), and pinnae grow from the
rachis in 2 planes (LHBH 1977). Pinnae grow in several
planes along the rachis of each Cuban royal palm leaf. Little
and Wadsworth (1964) contend that another characteristic
distinguishing Puerto Rico royal palm from Cuban royal
palm is that the latter lacks the swollen trunk of the former;
however, both West and Arnold (1952) and Neal (1965)
report the swollen baseand Neal (1965) and Braun (1983)
the swollen middle trunkin Cuban royal palm.
Flowers and fruits. Flowers of both species develop
from buds formed at the base of the leaves. Whitish male
and female flowers form on the same panicle, with male
flowers of each tree opening and falling before the female
flowers to prevent self-fertilization. Generally, each female
flower forms between 2 male flowers on the panicle (Francis
1992; Little and Wadsworth 1964). The male flowers have 3
small broad sepals and 3 blunt-pointed petals; the females
have 3 small broad sepals and a tubular corolla (Little and
Wadsworth 1964).
In Puerto Rico royal palm, the twice-branched drooping
panicles develop from large narrow buds. The panicles
develop inside a dark brown sheath that is 0.9 to 1.5 m long
(Francis 1992; Little and Wadsworth 1964). According to
LHBH (1977) and Braun (1983), one feature that distinguishes this species from the Cuban royal palm is the presence of scales on the axes bearing the flowers (rachillae).
The length of the inflorescence also seems to differ, with
that of Puerto Rico royal palm reaching up to 1 m (Little
and Wadsworth 1964) and that of Cuban royal palm reaching only 60 to 80 cm (Braun 1983; West and Arnold 1952).
The panicle of Puerto Rico royal palm bears stalkless male
flowers measuring 13 mm across, smaller female flowers,
982
References
Broschat TK, Donselman H. 1988. Palm seed storage and germination
studies. Principles 32(1): 312.
Braun A. 1983. Palmas para interiores, parques y avenidas. Caracas:
Venezuelan: Instituto Nacional de Parques, INPARQUES. 83 p.
Ellis RH, Hong TD, Roberts EH, Soetisna U. 1991. Seed storage behaviour
in Elaeis guineensis. Seed Science Research 1(2): 99104.
Francis JK. 1992. Puerto Rico royal palm. Res. Note SO-ITF-SM-55. New
Orleans: USDA Forest Service, Southern Forest Experiment Station. 5 p.
Francis JK, Rodriguez A. 1993. Seeds of Puerto Rican trees and shrubs: second installment. Res. Note SO-374. New Orleans: USDA Forest Service,
Southern Forest Experiment Station. 5 p.
Holdridge LR. 1967. Life zone ecology. San Jos, Costa Rica:Tropical Science
Center. 206 p.
LHBH (Liberty Hyde Bailey Hortorium). 1977. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan. 1290 p.
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington DC: USDA Forest Service. 375 p.
Little EL Jr, Wadsworth FH. 1964. Common trees of Puerto Rico and the
Virgin Islands. Agric. Handbk. 249. Washington, DC: USDA Forest Service:
4445.
Moscoso RM. 1945. Palmas dominicanas. Santo Domingo, Dominican
Republic: Universidad de Santo Domingo. 82 p.
Moore HE Jr. 1973. The major groups of palms and their distribution. Ithaca,
NY: Cornell University, New York State College of Agriculture and Life
Sciences, L.H. Bailey Hortorium. 115 p.
Neal MC. 1965. In gardens of Hawaii. Honolulu: Bishop Museum Press.
924 p.
Wolcott GN. 1946. A list of woods arranged according to their resistance
to the attack of the West Indian dry-wood termite, Cryptotermes brevis
(Walker). Caribbean Forester 7(4): 329334.
West E, Arnold LE. 1952. The native trees of Florida. Gainesville: University
of Florida Press. 2112 p.
Roystonea
983
RosaceaeRose family
Rubus L.
blackberry, raspberry
John C. Zasada and John C.Tappeiner III
Dr. Zasada retired from at the USDA Forest Services North Central Research Station; Dr.Tappeiner is a
professor at Oregon State Universitys College of Forest Science, Corvallis, Oregon
984
Common names
Occurrence
Allegheny blackberry,
sow-teat blackberry
smooth blackberry,
thornless blackberry,
mountain blackberry
swamp dewberry,
running blackberry
cutleaf blackberry,
evergreen blackberry
Himalayan blackberry
trailing blackberry,
Pacific blackberry
red raspberry
R. occidentalis L.
blackcap raspberry,
black raspberry, thimbleberry
salmonberry
R. spectabilis Pursh
R. stenopetalus Cham.
SUBGENUS: Chamaemorus (cloudberry)
R. chamaemorus L.
cloudberry, bake-apple
SUBGENUS: Anoplobatus (flowering raspberries)
R. odoratus L.
fragrant thimbleberry,
Rubacer odoratus (L.) Rydb.
flowering raspberry,
purple-flowering raspberry
R. parviflorus Nutt.
thimbleberry, western
thimbleberry
SUBGENUS: Cyclatis (Arctic berries)
R. arcticus L.
nangoon berry, arctic
bramble, wineberry
Sources: Brinkman (1974), Curtis (1959), Fernald (1950), Hickman (1993), Jennings (1988), MacKinnon and others (1992),Viereck and Little (1972).
985
Table 2Rubus, blackberry, raspberry: height or length at maturity and fruit ripeness criteria
Species
Growth
habit
Height or
length at
maturity
(m)
Year first
cultivated
SUBGENUS: Eubatus
R. allegheniensis
R. canadensis
R. hispidus
R. laciniatus
R. procerus
R. ursinus
Shrub
Shrub
Vine
Vine
Vine
Vine
1.8
2.8
1.82.5
2.84.6
6.29.2
4.66.2
1905
1727
1770
1890
Red, hard
Red, hard
Red, hard
Dull red
Red, hard
Red, hard
Black-purple
Black, soft
Reddish purple to black
Black, sweet, shining
Black, soft
Black, shining, soft
Shrub
Shrub
Shrub
2.2
1.52.2
2.84.6
1696
1827
Pink, hard
Bright red, hard
Pink, hard
Red, sweet
Purple-black, soft
Orange or red, soft
Perennial forb,
below-ground rhizome
0.10.2
Red, hard
Orange, soft
Shrub
Shrub
1.8
0.52.5
1635
Pink, hard
Pink, hard
Red, soft
Red, soft
SUBGENUS: Idaeobatus
R. idaeus
R. occidentalis
R. spectabilis
SUBGENUS: Chamaemorus
R. chamaemorus
SUBGENUS: Anoplobatus
R. odoratus
R. parviflorus
Sources: Brinkman (1974), Fernald (1950), Jennings (1988), MacKinnon and others (1992), Viereck and Little (1972).
Figure 1Rubus, blackberry, raspberry: general structure of ramets in populations with different growth habits.
Diagram (left) for speciesin this case red raspberry, a biennial cane speciesin which clone development occurs in the
soil by development of root (for example, red raspberry) or rhizome (for example, salmonberry) systems. Diagram
(right) for species in which clones expand by layering of above ground stems (for example, trailing raspberry).
KEY: a = primocanes, b = florocanes, c = dead canes, d = part of stem or root system that is either dead or nonflowering. (Drawings are based on observations by Whitney (1982, 1986), Suzuki (1987, 1989, 1990), and the authors.)
986
Rubus
987
Location
Flowering
Fruit ripening
Seed dispersal
R. allegheniensis
R. canadensis
R. laciniatus
R. hispidus
R. procerus
R. ursinus
SUBGENUS: Idaeobatus
NE US
Washington
Pacific Coast
MayJuly
JuneJuly
JuneAug
Juneearly Sept
JuneAug
JuneJuly
AugSept
JulySept
JulyOct
Mid-AugOct
AugSept
AugSept
AugSept
JulySept
SeptOct
AugOct
OctNov
R. idaeus
R. occidentalis
R. spectabilis
Rangewide
Alaska
OregonWashington
Late MayJuly
AprJune
MayJune
AprMay
Late JuneOct
JuneAug
JuneAug
MayJuly
JulyOct
JuneAug
JuneAug
JuneJuly
JuneJuly
JulyAug
AugSept
Pacific Northwest
JuneSept
MayJune
JulySept
JuneJuly
JulySept
JulyAug
SUBGENUS: Eubatus
SUBGENUS: Chamaemorus
R. chamaemorus
SUBGENUS: Anoplobatus
R. odoratus
R. parviflorus
Sources: Barber (1976), Brinkman (1974), Coladonato (1990a), Hippa and Koponen (1976),Viereck and Little (1972),Whitney (1978).
Species
Range
1.3 (0.82.4)
0.371.09
62
40
190
36
63
13
32
25
28128
1765
127246
2847
27103
1035
2.5
1.26.8
11
14
11
8
8
8
18
10
56
713
1016
318
2783
R. spectabilis
R. parviflorus
R. idaeus
General (N = 8 cv)
Restricted pollination
Open-pollination
R. arcticus
R. chamaemorus
Full light
Shade
Hand-pollination
Open-pollination
No defoliation
50% defoliation
General
General
Rubus subgen. Eubatus
Seeds/fruit
Avg
Source
W Oregon
SE Alaska
W Oregon
British Columbia & N Alberta
Norway
Norway
Norway
Finland
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Finland
Alaska
Arkansas
Sources: gren (1989), Moore and others (1974a), Nybom (1986), Rantala (1976), Redalen (1977), Ryynnen (1973), Staniforth and Sidhu (1984), Suzuki (1990),Van
Adrichem (1972),Willson (1996),Whitney (1978), Zasada (1996).
have used seed.] Each drupelet is a complete fruit, a miniature version of a cherry or plum (which are drupe-type
fruits). Each aggregate fruit is the product of 1 flower and
the number of drupelets per aggregate varies with species,
pollination success, and environmental conditions (figure 3
and table 4). Ripening occurs 30 to 36 days and 40 to 70
days after pollination in raspberries and blackberries,
988
respectively. Drupelets within an aggregate fruit ripen uniformly, but there can be considerable variation among fruits.
Three phases of development are recognized: rapid fruit
growth following pollination, slow growth as the seed develops, and a final period of rapid growth before the fruit is
fully mature (Jennings 1988). In natural populations, the
interaction between microclimate and genetic variation in
The breeding system in Rubus is often described as versatile because seeds are formed sexually and asexually. The
relative importance of these two types of seeds varies within
and among subgenera and species and may differ within a
plant depending on the pollen source. In the Idaeobatus
Figure 2Rubus, blackberry, raspberry: red raspberry
group, seeds are normally formed sexually. In Eubatus
clone showing distribution of ramets (circles) as they were
species, seeds are produced sexually and asexually
in a clone excavated on an upland site in central Alaska.This
(Jennings 1975; Nybom 1985, 1986, 1988). In most cases,
plant was about 5-years-old and originated from seed. Red
raspberry clones develop by expansion of the root system.
pollen is required to produce seed asexually, but the embryo
Salmonberry, thimbleberry, cloudberry, and other species
is not produced by the fusion of male and female gametes
may develop clones with similar ramet distribution, but
(pseudogamy). Parthenogenesis (seed formation without polclone expansion occurs by the growth of rhizomes. Ramet
lination) occurs in some species. Seeds of both sexual and
longevity in these latter species is also different.
asexual origin may be present in the same fruit (Jennings
1975; Nybom 1985, 1986, 1988).
The abscission layer that develops as the fruit ripens differs in raspberries and blackberries. Fruits may drop from
the plant or be removed by various animal species. The
number of drupelets or entire aggregates removed at any one
time depends on the size of the fruit and the size and eating
habits of the animal (Snow and Snow 1988). Seeds are
usually deposited with other materials in the feces. Large
animals such as the grizzly bear (Ursus arctos), may deposit
50,000 to 100,000 salmonberry seeds in a single pile of
feces. Seeds may be secondarily consumed or moved from
the feces piles by small rodents and birds. Brunner and others (1976), Jordano (1984), Gervais (1996), and Traveset
and Willson (1997, 1998) discuss other aspects related to
selection and dispersal of Rubus seeds by animals. The
amount of fruit removed has been found to vary from near
100 to 40% and will depend on habitat type and type of animal feeding on the fruits (Jordano 1982; Snow and Snow
Figure 3Rubus, blackberry, raspberry: fruits of R.
alleghensis, Allegheny blackberry (upper far left); R. canaden- 1988). In British Columbia, forest silvicultural practices are
being altered in coastal riparian areas to provide for adesis, smooth blackberry (upper middle left); R. hispidus,
swamp dewberry (upper middle right); R. procerus,
quate fruit production by salmonberry and other species that
Himalayan blackberry (upper far right); R. parviflorus,
are important food sources for grizzly bear (McLennan and
thimbleberry (lower left); and R. ursinus, trailing
Johnson 1993).
blackberry (lower right).
Although fruit consumption is often viewed as a loss of
seeds, in Rubus spp. consumption of seeds is important to
the reproductive biology of the plant. Several examples are
described below. Dispersal of seeds away from parent plants
depends on animals. The distribution of seeds in space and
time depends on the size and eating habits of the animal (for
example, bears deposit large quantities of seeds in one place,
whereas small birds deposit only a few seeds at a time), and
the movement habits of the animal following feeding. Seeds
that pass through the digestive tract of animals receive varying degrees of scarification (for example, salmonberry seeds
in bear feces may have had the fleshy fruit wall completely
removed or be little affected, as evidenced by the presence
flowering and fruit ripening usually spreads the timing of
aggregate maturation over a period of several weeks or
more.
Rubus
989
990
Species
SUBGENUS: Eubatus
R. allegheniensis
R. canadensis
R. hispidus
R. laciniatus
R. procerus
R. ursinus
Rubus
(general European)*
SUBGENUS: Idaeobatus
R. idaeus
Place
collected
R
Seeds (x1,000)/weight
Range
/kg
/lb
Seed wt/fruit wt
g/kg lb/100 lb
Average
/kg
/lb
Washington
Washington
40
40
58
4
4
0.7
5.8
370724
458495
282513
168329
208225
128233
574
476
408
301
323
845
262
216
185
137
147
384
Sweden
359869
163395
480
219
Minnesota
30
British Columbia/Alberta
46
R. occidentalis
Minnesota
3080
R. spectabilis
Oregon 1
Oregon 2
Oregon 3
Oregon 4
Alaska
SUBGENUS: Chamaemorus
R. chamaemorus
Sweden/Finland
59
Alaska
SUBGENUS: Anoplobatus
R. odoratus
Pennsylvania
R. parviflorus
Oregon
Washington
SUBGENUS: Cyclatis
R. arcticus
Sweden/Finland
76
3
4.6
3-8
667845
469794
629845
251528
189321
27045
216298
303384
213397
286384
115240
87146
123157
98135
722
632
735
354
240
316
265
315
328
288
334
162
109
144
120
143
(5.9)
80101
3745
122
90
56
40
357806
7191201
162367
327546
1,085
611
20
493
278
418
(7.6)
980
446
Sources: Brinkman (1974), Lautenschlager (1990), Nybom (1980), Rantala (1976), Zasada (1996).
* Seeds of 20 species in Sweden.
Schott., the individual seed weight with the highest frequency was 2 to 2.5 mg, whereas weights ranged from 1 to 5 mg
(Jordano 1984).
Extraction and storage of seeds. Seeds may be
extracted by macerating the fruits in water then floating off
or screening out the pulp and empty seeds (Brinkman 1974).
Because of the high strength of the endocarp (figure 5),
maceration does not damage the seeds (Rose 1919). Small
lots of fruit may be covered with water and macerated in a
blender until the pulp and fiber are separated (Morrow and
others 1954). Additional water is then added, the sound
seeds allowed to settle, and the pulp and empty seeds
decanted. Several changes of water will yield cleaner seeds.
Seed yield data are presented in tables 4 and 5.
The cleaned seeds should be dried before storage. Clark
and Moore (1993) reported that seeds from raspberry cultivars germinated well after storage for 26 years at 4 to 5 C.
Rubus seeds can be present in the forest floor of many
forest types in North America (Barber 1976; Graber and
Thompson 1978; Granstrom 1982; Maxwell 1990; McGee
991
992
10
20
30
69 (1894)
93 (8496)
60 (4088)
28 days
6 days
33 days
48
53
21
21
6
0
21
21
10
0
21
21
8
0
2128
2128
2128
2128
0
0
62
81
2128
2128
2128
0
25
73
2128
Variable
Variable
Variable
<1
310
3031
Sources: Barber (1996), Dale and Jarvis (1983), Lautenschlager (1990), Lundergan and Carlisr (1984); Moore and others (1974); Nesme (1985), Rantala (1976).
* Seeds were treated as follows: surface sterilized in 1% sodium hypochlorite (NaClO) for 10 minutes, nicked to expose radicle; soaked for 3 min in 1% NaClO, and
incubated in the dark for 1 year.
Seeds were extracted and air-dried, treated for 20 minutes with sulfuric acid and 7 days with calcium hypochlorite; stratified at 5 C or unstratified. Fruits were
collected 43 days after anthesis. Seeds were from fruits collected earlier or later differed in germination response.
Seeds were from natural populations in Maine; they were stratified for 2 to 6 months after acid treatment; tests were conducted for 30 days.
Fresh seeds from Washington state populations; stratified at 2 to 5 C.
II Seeds stratified at 1 C and germinated monthly in a mist propagation chamber.
Rubus
993
994
seedling
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996
ArecaceaePalm family
Sabal Adans.
palmetto
Common name(s)
Occurrence
Florida
Texas
dwarf palmetto,
Sonoran palmetto
cabbage palmetto,
cabbage palm, palmetto
Sabal
997
998
longitudinal
Species
S. etonia
S. palmetto
/kg
758763
Cleaned seeds/weight
Range
/lb
/kg
1,6681,682
Average
1,280
1,675
/lb
Samples
Moisture
content (%)
581
7,600
8
2
9.8
19.3
References
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Carpenter WJ. 1987. Temperature and imbibition effects on seed germination of Sabal palmetto and Serenoa repens. HortScience 22(4): 660.
Carpenter WJ. 1988. Temperature affects seed germination of four Florida
palm species. HortScience 23(2): 336337.
Dickie JB, Balick MJ, Linington IM. 1993. Studies on the practicality of ex situ
preservation of palm seeds. Princepes 37(2): 9498 [Seed Abstracts
1996; 9(4): 1304].
Ellis RH, Hong TD, Roberts EH. 1985. Handbook of seed technology for
genebanks. Volume 2, Compendium of specific germination information
and test recommendations. Rome: International Board for Plant Genetic
Resources. 667 p.
Sabal
999
SalicaceaeWillow family
Salix L.
willow
John C. Zasada, D.A. Douglas, and W. Buechler
Dr. Zasada retired from the USDA Forest Services North Central Research Station;
Dr. Douglas is an emeritus professor and Mr. Buechler is an affiliate faculty member at
Boise State Universitys Department of Biology, Boise, Idaho
Growth habit, occurrence and use. The willow
genusSalixincludes 350 to 400 species (Argus 1996).
The majority are in the Northern Hemisphere, from arctic
through temperate latitudes (table 1). Three species are
native south of Mexico (Dorn 1976) and 67 found in the
contiguous 48 United States, where tree and shrub forms
predominate. The 39 species found in Alaska are mostly tall
to medium shrubs; prostrate growth forms are mostly on the
tundra. Shrub and prostrate shrub forms constitute a dominant portion of the vegetation of the Circumpolar Arctic
(defined as north of the treeline) and include about 29
species in the North American Arctic (Argus 1996). These
tundra species are segregated into a variety of habitats
(Argus 1973; Viereck and Little 1972). General information
on the genus worldwide can be found in Warren-Wren
(1972) and Newsholme (1992); the taxonomy and distribution of American species is covered by Argus (1973, 1986,
1995), Dorn (1976), MacKinnon and others (1992), and
Viereck and Little (1972). General reviews of ecological
characteristics and effects of fire on more than 20 species
are available in the Fire Effects Information System
Database (Fisher 1992). Seed characteristics of poplar
(Populus, the other North American genus in the Salicaceae)
are very similar to those of willows, and information for
poplar is applicable to willow (Schreiner 1974; Zasada and
Wyckoff 2002). The uniformity in seed characteristics, particularly germination, in the Salicaceae is remarkable considering that the family comprises several hundred species.
The importance of willows as a component of regional
vegetation varies geographically and with the mix of habitat
types within the region (Fisher 1992). In particular, willows
become more important with increasing latitude in North
America. In the boreal forests of northern Canada and
Alaska, willows are the most common tall and intermediate
shrubs; they fill niches occupied by hazel (Corylus spp.),
maple (Acer spp.), and cherry and plum (Prunus spp.) in
more southern parts of the boreal forest. In the tundra,
1000
Common name
Occurrence
S. alaxensis (Anderss.)
Coville
feltleaf willow
S. amygdaloides Anderss.
peachleaf willow
S. arctica Pallas
S. babylonica L.
arctic willow
weeping willow
S. bebbiana Sarg.
Bebb willow
S. boothii Dorn.
Booth willow
S. caroliniana Michx.
S. discolor Muhl.
S. eriocephala Michx.
S. cordata Muhl.; S. rigida Muhl.
S. exigua Nutt.
cordate willow
S. geyerana Anderss.
Geyer willow
S. glauca L.
white willow
S. interior Rowlee
sandbar willow
S. lasiolepis Benth.
arroyo willow
S. lucida Muhl.
[incl. S. lasiandra Benth.]
S. lutea Nutt.
Pacific willow
S. nigra Marsh.
black willow
S. petiolaris Sm.
meadow willow
S. planifolia Pursh.
S. repens L.
S. scouleriana Barratt
ex Hook.
diamondleaf willow
creeping willow
Scouler willow
coyote willow
yellow willow
Sources: Argus (1973, 1975), Brinkman (1974), Cooper and Van Havern (1994), Hillier and sons (1989), Little (1979), MacKinnon and others (1992), Newsholme
(1972, 1992),Viereck (1987),Viereck and Little (1972),Vogel (1990).
buds located at the base of the stem. Other types of vegetative regeneration found in a limited number of species
include sprouting from roots, layering of stems, and rooting
of broken stem and branch segments. In riparian areas,
whole plants are sometimes dispersed by water after being
washed out by erosion. Artificial regeneration can be
achieved by seeding or by planting seedlings and stem cuttings. Willows regenerate quickly after natural disturbances
such as flooding (Krasny and others 1988; Shafroth and
others 1994; Viereck 1970) and fire (Lyon and Stickney
1976; Viereck and Dyrness 1979; Zasada and others 1983;
Zedler and Sheid 1988). They also regenerate on sites dis-
Salix
1001
1002
is no definitive biochemical test or molecular genetics technique available for distinguishing male and female plants.
Mosseler (1987) and Mosseler and Zsuffa (1989) found
highly skewed sex ratios resulting from controlled inter- and
intraspecific crosses. Sex ratios in naturalized exotic
speciesfor example, S. rubens Schronk (pro spp.) and
S. alba spp. vitellina (L.) Arcang. in riparian areas in
Coloradoare often highly skewed toward one sex because
of vegetative reproduction (Shafroth and others 1994).
There have been reports of differences in vegetative
characteristics and growth rate between male and female
plants, but these differences are not well-established
(Alliende and Harper 1989; Crawford and Balfour 1983,
1990). Male plants usually produce more flowers per unit of
crown area than female plants (Kay and Chadde 1992;
Zasada 2000).
Although the dioecious trait is universal across the
genus, hermaphrodite plants (individuals with separate male
and female flowers) and catkins (male and female flowers
on the same catkin) have been observed in a number of
species (Alliende and Harper 1989; Crawford and Balfour
1983; Mosseler and Zsuffa 1989). Mosseler and Zsuffa
(1989) observed hermaphroditic plants in both natural populations and in controlled inter- and intraspecific crosses.
Plants that are hermaphroditic initially sometimes become
completely male as they mature sexually (Mosseler and
Zsuffa 1989).
Seed-bearing age in willows depends on species and site
conditions. Following disturbances such as fire and logging,
willows of vegetative origin (for example, stump sprouts
developing from the basal bud bank) flower sooner than
plants of seed origin. Sprouts often produce seeds 1 to 2
years after a fire that kills the mature plant, whereas
seedlings of the same species require 5 to 10 years before
the first seeds are produced. In controlled environments,
Mosseler and Zsuffa (1989) reported that coyote willow
plants flowered several months after germination in a controlled environment and Mosseler (1996) found that 6 of 7
native willows flowered within 2 years of germination.
Zasada (2000) has also observed flowering in 1-year-old
creeping willow seedlings.
Catkins bearing several to many staminate or carpellate
flowers (figure 1) appear before or after leaf appearance,
depending on the species (Mosseler 1987; Viereck and Little
1972). Each carpellate flower contains 2 carpels. The number of ovules per carpel may vary considerable within and
among species. Argus (1996) observed the following variation in ovules per carpel: feltleaf willow, 6 to 9; peachleaf
willow, 8 to 11; arctic willow, 6 to 7; Bebb willow, 3 to 8;
Salix
1003
Location
S. alaxensis
S. amygdaloides
S. arctica
S. bebbiana
S. boothii
S. discolor
S. exigua
S. geyeriana
S. lucida
S. lutea
S. petiolaris
S. repens
Capsules in catkin
with seeds
Seeds/capsule
Seeds/catkin
45 (3561)
71 (6180)
119 (92137)
98 (82109)
8 (78)
9 (810)
10 (811)
7 (68)
16 (1418)
333 (245427)
673 (488800)
1,174 (7361,280)
600 (492763)
24 (840)
65 (6070)
7 (212)
37 (2448)
64 (4379)
18 (1229)
74 (6978)
4 (34)
18 (1025)
9 (810)
12 (518)
6 (57)
6
10 (812)
25 (1536)
5 (46)
17 (1220)
11 (1112)
3 (25)
22 (1925)
432
595
84
218 (144311)
400 (286427)
81 (42171)
841 (754925)
82 (50110)
Sources: Jones (1995), Kay and Chadde (1992), Moore (1982), Mosseler (1987), Zasada (2000).
Note: Values are means with ranges in parentheses.
1004
a wide altitudinal or elevational range (table 3). Once capsules are ripe and begin to open, the rate of seed dispersal is
determined by weather conditions: under warm, dry, windy
conditions all seeds may be dispersed within a few days.
Under wetter, cooler conditions, dispersal may be spread out
over a month. If a small amount of seeds is all that is
required, stems with immature catkins can be collected and
placed in a greenhouse in water; seeds can then be collected
when the capsules open (Marten and Young 1992).
After catkins have been removed from the plant they
should be placed in a paper bag that allows the catkin-drying
process to continue during transport. Catkins should not be
packed tightly because air circulation may be restricted.
Bags containing catkins must be kept out of direct sunlight.
To obtain seeds from a specific inter- or intraspecific
cross, dormant stem cuttings (50 cm or less in length) with
reproductive buds should be obtained from female and male
plants in late winter (Mosseler 1987). These cuttings should
then be placed in a greenhouse or growth chamber, where
they will flower within 2 weeks. Male clones are often
forced to flower before females and the pollen stored until
the females are ready for pollination. This avoids pollination
from unwanted sources. Willow pollen may be frozen for 1
to 2 months without losing its viability. There is a period of
3 to 6 days, depending on species, during which flowers can
be pollinated (Mosseler 1987). Catkins will produce viable
Species
Location
Flowering
Fruit ripening
Seed dispersal
S. alaxensis
AlaskaBrooks Range
AlaskaTanana River
Alaskacentral Interior*
NE Minnesota
Canadian high Arctic
Ellesmere Island
Interior Alaska
MayJune
AprMay
MayJune
MayJune
JuneJuly
May
JuneJuly
JulyAug
MayJune
JulyAug
July
JuneJuly
AprJune
MarApril
May
MarApril
AprJune
AugSept
JulyAug
MayJune
MarApr
AprMay
June
AugSept
MayJune
AprMay
June-July
MayJuly
AprMay
JuneJuly
MayJune
JuneJuly
JuneJuly
MayJune
JulyAug
JulyAug
JulyAug
Aug 13
JuneAug
AprMay
JuneJuly
JuneJuly
MayJuly
JuneJuly
MayJune
SeptNov
SeptNov
SeptNov
JuneAug
AprMay
JuneJuly
JuneJuly
MayJuly
S. amygdaloides
S. arctica
S. bebbiana
S. caroliniana
S. discolor
S. eriocephala
(as S. rigida)
S. exigua
S. fragilis
S. glauca
S. interior
S. lucida
S. nigra
S. petiolaris
S. scouleriana
US & Europe
AlaskaBrooks Range
Alaskamid-boreal forest
AlaskaDenali National Park
N Ontario
Idaho
In north
In south
General
General
AprMay
FebApril
MayJune
MayJune
AprJune
Sources: Brinkman (1974), Densmore and Zasada (1983), Jones (1995),Viereck and Little (1972).
* High elevation. As S. lasiandra.
Salix
1005
seeds, divisions
Place collected
S. amygdaloides
S. bebbiana
S. caroliniana
S. exigua
S. fragilis
S. lasiandra
S. petiolaris
S. scouleriana
Minnesota
Idaho (770 m)
South Carolina
Washington (615 m)
Minnesota
Idaho (770 m)
Minnesota
Idaho (770 m)
S
Cleaned seeds (x 1,000)/weight
/kg
/lb
5,720
5,500
18,260
22,000
7,040
25,300
1,100
14,300
2,600
2,500
8,300
10,000
3,200
11,500
500
6,500
Samples
1
2
1
1
1
1
1
1
Figure 7Salix, willow: generalized patterns of temperature and stratification effects on germination of seeds of
summer- and fall-dispersing willows; numbers indicate
germination temperatures (C).
1007
References
Alliende MC, Harper JL. 1989. Demographic studies of a dioecious tree: 1.
Colonization, sex and age structure of a population of Salix cinerea.
Journal of Ecology 77: 10291047.
Argus GW. 1973. The genus Salix in Alaska and the Yukon. Pub. Bot. 2.
Ottawa: National Museum of Natural Sciences.
Argus GW. 1974. An experimental study of hybridization and pollination in
Salix (willow). Canadian Journal of Botany 52(7): 16131619.
Argus GW. 1986. The genus Salix (Salicaceae) in the southeastern United
States. Systematic Botany Monographs 9: 1170.
Argus GW. 1995. Salicaceae, part 2: Salix. Journal of the ArizonaNevada
Academy of Science 29(1): 3961.
Argus GW. 1996. Personal communication. Ottawa: Canadian Museum of
Nature.
Begin Y, Payette S. 1991. Population structure of lakeshore willows and icepush events in subarctic Quebec, Canada. Holarctic Ecology 14: 917.
Bewley JD, Black M. 1994. Seeds: physiology, development, and germination. New York: Plenum Press. 445 p.
Bishop SC, Chapin FS. 1989. Establishment of Salix alaxensis on a gravel
pad in arctic Alaska. Journal of Applied Ecology 26: 575583.
Bliss LC, Cantlon JE. 1957. Succession on river alluvium in northern Alaska.
American Midland Naturalist 58(2): 454469.
Brinkman KA. 1974. Salix L., willow. In: Schopmeyer CS, tech. coord. Seeds
of woody plants in the United States. Agric Handbk. 450. Washington
DC: USDA Forest Service: 746750.
Chmelar J, Meusel W. 1979. Die Weiden Europas. Wittemberg, Germany: A.
Ziemsen Verlag. 143 p.
Chosa JA, Shetron SG. 1976. Use of willow cuttings to revegetate the
slime areas of iron mine tailings basins. Res. Note 21. Houghton:
Michigan Technological University. 6 p.
Cooper DJ,Van Haveren. 1994. Establishing feltleaf willow from seed to
restore Alaskan, U.S.A. floodplains. Arctic and Alpine Research 26(1):
4245.
Crawford RMM, Balfour J. 1983. Female predominant sex ratios and physiological differentiation in arctic willows. Journal of Ecology 71: 149160.
Crawford RMM, Balfour J. 1990. Female-biased sex ratios and differential
growth in arctic willows. Flora 184: 291302.
Densmore R. 1979. Aspects of the seed ecology of woody plants of the
Alaskan taiga and tundra [PhD thesis]. Durham, NC: Duke University.
285 p.
Densmore R, Zasada JC. 1978. Rooting potential of Alaska willow cuttings.
Canadian Journal of Forest Research 8: 477479.
Densmore R, Zasada JC. 1983. Seed dispersal and dormancy patterns in
northern willows: ecological and evolutionary significance. Canadian
Journal of Botany 61: 32073216.
1008
Schreiner EJ. 1974. Populus L., poplar. In: Schopmeyer CS, tech. coord. Seeds
of woody plants in the United States. Agric. Handbk. 450. Washington,
DC: USDA Forest Service: 645655.
Sennerby-Forsse L, ed. 1986. Handbook for energy forestry. Uppsala:
Swedish University of Agricultural Sciences, Department of Ecology and
Environmental Research.
Shafroth PB, Scott ML, Friedman JM. 1994. Establishment, sex structure, and
breeding system of an exotic riparian willow, Salix rubens. American
Midland Naturalist 132: 159172.
Simak M. 1980. Germination and storage of Salix caprea L. and Populus
tremula L. seeds. In Proceedings, International Symposium on Forest Tree
Seed Storage; 1980 September 2327; Chalk River, ON. Ottawa:
Canadian Forestry Service: 142160.
Siren G, Sennerby-Forsse L, Ledin S. 1987. Energy plantations: short rotation forestry in Sweden. In: Hall DE, Overend RP, eds. Biomass. London:
John Wiley and Sons: 119143.
Toepfer AD. 1915. Salices Bavariae,Versuch einer Monographie der bayerischen Weiden. Berichte der Bayerischen Botanischen Gesellschaft zur
Erforschung der Heimischen Flora 15.
Viereck LA, Little EL Jr. 1972. Alaska trees and shrubs. Agric. Handbk. 410.
Washington, DC: USDA Forest Service. 265 p.
Viereck LA. 1970. Forest succession and soil development adjacent to the
Chena River in interior Alaska. Arctic and Alpine Research 2: 126.
Viereck LA, Dyrness CT, eds. 1979. Ecological effects of the Wickersham
Dome fire near Fairbanks, Alaska. Gen.Tech. Rep. PNW-90. Portland, OR:
USDA Forest Service, Pacific Northwest Forest and Range Experiment
Station.
Viereck EG. 1987. Alaskas wilderness medicines: healthful plants of the far
north. Edmonds, WA: Northwest Publishing Company. 107 p.
Vogel VJ. 1990. American Indian medicine. Norman: University of
Oklahoma Press. 578 p.
Vroege PW, Stelleman P. 1990. Insect and wind pollination in Salix repens L.
and Salix caprea L. Israel Journal of Botany 39(1/2): 125132.
Walker LR, Zasada JC, Chapin FS. 1986. The role of life history processes in
primary succession on an Alaskan floodplain. Ecology 67(5): 12431253.
Warren-Wren SC. 1972. Willows. Newton Abbot, UK: David and Charles
Publishers. 179 p.
Westoby JC. 1975. China uses trees to halt sands. World Wood 16(13):
2022.
Wyckoff G, Zasada JC. 2005 . Populus L., poplar. In: Bonner F, tech. coord.
Woody plant seed manual. Agric. Handbk. 727. Washington, DC: USDA
Forest Service: 856871.
Zasada JC. 2000 Unpublished data. Grand Rapids, MN: USDA Forest
Service, North Central Research Station.
Zasada JC, PI. Coyne 1975. Unpublished data. Grand Rapids, MN: USDA
Forest Service, North Central Research Station.
Zasada JC, Densmore RA. 1977. Changes in viability during storage for
selected Alaskan Salicaceae. Seed Science and Technology 5: 509518.
Zasada JC, Densmore R. 1979. A trap to measure Populus and Salix seedfall. Canadian Field-Naturalist 93(1): 7779.
Zasada J[C], Densmore R[A]. 1980. Alaskan willow and balsam poplar seed
viability after 3 years storage.Tree Planters Notes 31: 910.
Zasada JC,Viereck LA. 1975. The effect of temperature and stratification
on germination in selected members of the Salicaceae in interior Alaska.
Canadian Journal of Forest Research 5: 333337.
Zasada JC, van Veldhuizen RM, Norum RA,Teutsch CE. 1983. Artificial
regeneration of trees and tall shrubs in experimentally burned upland
black spruce/feathermoss stands in Alaska. Canadian Journal of Forest
Research 13(5): 903913.
Zedler PH, Scheid GA. 1988. Invasion of Carpobrotus edulis and Salix lasiolepis after fire in a coastal chaparral site in Santa Barbara county.
Madrono 35(3): 196201.
Zsuffa L, Sennerby-Forsse L, Weisgerber H, Hall RB. 1993. Five strategies
for clonal forestry with poplars, aspens, and willows. In: Ahuja MR, Libby
WJ, eds. Clonal forestry.Volume 2, Conservation and Application. Berlin:
Springer-Verlag.
Salix
1009
LamiaceaeMint family
Salvia L.
sage
Susan E. Meyer
Dr. Meyer is a research ecologist at the USDA Forest Service's Rocky Mountain Research Station,
Shrub Sciences Laboratory, Provo, Utah
Scientific name
S. apiana Jepson
white sage
Subshrub
S. dorrii (Kellogg)
Abrams
Dorr sage,
purple sage
Shrub
S. mellifera Greene
black sage
Subshrub
S. sonomensis Green
creeping sage
Mat-forming
suffrutescent
Source:
1010
Habitat
Coastal sage scrub, chaparral,
pondersoa pine forest, warm
desert shrub margins
Warm & cold desert shrub
communities, piyonjuniper
woodland
Coastal sage scrub, chaparral,
warm desert shrub margins
Chaparral, oak woodland,
ponderosa pine forest
Distribution
Coastal & cis-montane S
California S to Baja California
W Washington to SE California,
Arizona, Utah, & Idaho
Coastal & cis-montane central
California S to Baja California
Coastal & cis-montane California,
mostly in N
nutlets.
longitudinal
Salvia
1011
Seeds/weight
Species
mg
oz
S. apiana
1.4
1.9
2.4
1.1
1.5
0.8
.05
.07
.08
.04
.05
.03
S. dorrii
S. mellifera
S. sonomensis
Source:
/kg
Max
germination %
/lb
714,400
511,600
416,800
911,000
685,800
1,212,800
324,000
233,000
189,000
413,000
311,000
550,000
42
43
53
69
90
70
Kay and others (1988), Keeley (1986, 1987), Nord and Gunter (1974).
Light
Alternating temp
Chilling
Heat shock
Dry storage
Charate
GA
+
+
nd
+
+
nd
+
+
+
+
+
nd
+
nd
nd
nd
+
nd
+
nd
+
Sources: Emery (1988), Kay and others (1988), Keeley (1986, 1987), Nord and Gunter (1974).
Notes: nd = no data; see text for details.
1012
References
ANPS [Arizona Native Plant Society]. 1990. Desert shrubs.Tucson: Arizona
Native Plant Society, Urban Landscape Committee. 35 p.
Belcher E[W Jr]. 1985. Handbook on seeds of browse-shrubs and forbs.
Tech. Pub. R8-8. USDA Forest Service, Southern Region. 246 p.
Capoh B, Maxwell GL, Smith PH. 1978. Germination responses to temperature pretreatment of seeds from ten populations of Salvia columbariae
in the San Gabriel Mountains and Mojave Desert, California. Aliso 9:
365-373. Correll DS, Johnson MC. 1970. Manual of the vascular plants of
Texas. Renner:Texas Research Foundation. 1881 p.
Emery DE. 1988. Seed propagation of native California plants. Santa
Barbara, CA: Santa Barbara Botanic Garden. 107 p.
Jepson WL. 1951. A manual of flowering plants of California. Berkeley:
University of California Press. 1238 p.
Kay BL, Graves WL,Young JA. 1988. Long-term storage of desert shrub
seed. Mojave Reveg. Notes 23. Davis: University of California
Department of Agronomy and Range Science. 22 p.
Salvia
1013
CaprifoliaceaeHoneysuckle family
Sambucus L.
elder
Kenneth A. Brinkman and W. Gary Johnson
Dr. Brinkman retired from the USDA Forest Services North Central Forest Experiment Station;
Mr. Johnson retired from the USDA Forest Services National Seed Laboratory
(figure 1) containing 3 to 5 one-seeded nutlets or stones (figures 2 and 3). When ripe, the fruits vary from red to nearly
black, depending on species (table 3). Dispersal is chiefly by
birds and animals.
Geographic races. Regional floras do not agree on
the taxonomy of this genus. There is great confusion on
delimiting the species, subspecies, and varieties (LHBH
1976). Two varieties of blue elder have definite geographic
limits and these may be climatic races. Both American and
scarlet elders have developed varieties, but these do not
appear to be related to climate.
Collection of fruits. Elder fruits are collected by
stripping or cutting the clusters from the branches.
Collection should be made as soon as the fruits ripen to
reduce losses to birds. If the seeds are not to be extracted
immediately, the fruits should be spread out in thin layers to
prevent heating. Scarlet elder fruits can be harvested before
Common name(s)
Occurrence
Flowering
Fruit ripening
Seed dispersal
S. nigra
spp. canadensis
spp. cerulea
S. racemosa var. racemosa
JuneJuly
MayJuly
AprJuly
JulySept
AugSept
JuneAug
AugOct
AugOct
JuneNov
Sources: Harris (1969), Hitchcock and others (1959), LHBH (1976), Plummer and others (1968), Rehder (1940),Van Dersal (1938).
1014
fruit
Figure 2Sambucus, elder: seeds of S. racemosa var. racemosa, scarlet elder (top); S. nigra spp. canadensis, American
elder (center); and S. nigra spp. cerulea, blue elder
(bottom).
maturity and the seeds will still germinate (Dirr and Heuser
1987). Cram (1982) found that seeds harvested on August 5
in Saskatchewan before the fruit was color ripe showed 95%
germination after 90 days of cold stratification, whereas
seeds harvested on August 25 showed only 76%. Increased
dormancy or harder seedcoat may be the cause of the lower
germination in the seeds harvested August 25 (Cram 1982).
Specific gravity or moisture content of the fruits did not
decrease during maturation but fluctuated up and down,
apparently in response to rainfall (Cram 1982).
Extraction and storage of seeds. The fruits (figure 1)
may be (a) dried; (b) run through a macerator with water
and the pulp and empty seeds floated off (Plummer and others 1968); or (c) crushed, dried, and used without further
cleaning. Commercial seedlots may consist of either dried
fruits or cleaned seeds (figures 2 and 3). After a short period
of drying, lots of freshly extracted seeds may be fanned or
screened to remove debris. Excessive drying should be
avoided. Seed yields and number of seeds per weight vary
among species (table 4).
Small lots of fruit may be cleaned in a food blender
(Morrow and others 1954). The berries are covered with
water, the blender is run long enough to macerate them, and
more water is added to float off the pulp and debris. Several
changes of water will result in cleaner seeds. The seeds are
separated from the last change of water in a filter-line funnel
and can be dried on the filter paper.
Elder seeds may be stored dry at 5 C for several years
(Morrow and others 1954). Seeds of American elder showed
Figure 3Sambucus racemosa var. racemosa, scarlet elder:
longitudinal section through a nutlet.
Sambucus
1015
Table 3Sambucus, elder: height, seedcrop frequency, and fruit ripeness criteria
Height at
maturity (m)
Species
S. nigra
spp. canadensis
spp. cerulea
S. racemosa var. racemosa
2.7
9
3.0
Year first
cultivated
Years between
large seedcrops
1761
1850
1812
Sources: Brinkman (1974), Fernald (1950), LHBH (1976), Rehder (1940), Rydberg (1965).
Species
Seeds/wt of fruit
kg /45 kg
lbs/100 lbs
S. nigra
spp. canadensis
spp. cerulea
S. racemosa var. racemosa
3.28.2
2.7
1.8
718
6
4
Range
/kg
385713
257570
422829
Cleaned seeds/weight
Average
/lb
/kg
/lb
175324
117259
192377
510
451
629
232
205
286
Samples
14
23
6
Sources: Brinkman (1974), McKeever (1938), Plummer (1968), Rydberg (1965), Swingle (1939).
1016
Species
S. nigra
spp. canadensis
spp. cerulea
S. racemosa var. racemosa
stratification treatments
Warm period
Temp (C)
Days
Medium
Sand
Sand
Sand
2030
Room*
2030
Cold period
Temp (C)
Days
60
450*
3060
5
5
5
90150
98
90150
Species
S. nigra
ssp. canadensis
Sand or soil
spp. cerulea
Sand
S. racemosa var. racemosa
Sand
Germination speed
Percent
Days
Germination avg
(%)
Samples
Purity
(%)
30
21
20
21
60
35
32
55
16
12
63
79
7
3
80
8090
30*
20*
60
50
27
47
97
Sources: Adams (1927), Brinkman (1974), Davis (1927), McKeever (1938), Plummer (1968).
* Day and night temperature of 77 and 50 F were used on some samples.
at least 16 hours of light, the need for light has not been
established. Germination is epigeal (figure 4).
Nursery practice. Elder seeds can be sown in the fall
soon after collection, or they can be stratified and sown in
the spring. In either case, germination often is not complete
until the second spring. At the USDA Forest Services nursery in Coeur dAlene, Idaho, dried seeds of blue elder usually are soaked in water for 3 days, then stratified in vermiculite for 3 months at 1 C before spring-sowing (Brinkman
References
Adams J. 1927. The germination of seeds of some plants with fleshy fruits.
American Journal of Botany 14: 415428.
Brinkman KA. 1974. Sambucus, elder. In: Schopmeyer CS, tech. Coord. Seeds
of woody plants in the United States. Agric. Handbk. 450. Washington,
DC: USDA Forest Service: 372.
Cram WH. 1982. Seed germination of elder (Sambucus pubens) and honeysuckle (Lonicera tatarica). Hortscience 17(4): 618619.
Davis OH. 1927. Germination and early growth of Cornus florida,
Sambuscus canadensis, and Berberis thunbergii. Botanical Gazette 84:
225263.
Dirr M, Heuser C Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1632 p.
Harris AS. 1969. Unpublished observation. Juneau, AK: USDA Forest
Service, Pacific Northwest Forest and Range Experiment Station.
Heit CE. 1967. Propagation from seed: 7. Successful propagation of 6 hardseeded group species. American Nurseryman 125(12): 1012, 3741,
4445.
Hitchcock CL, Cronquist A, Ownbey M,Thompson JW. 1959. Vascular
plants of the Pacific Northwest: 4. Ericaceae through Campanulaceae.
Seattle: University of Washington Press. 510 p.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan. 1290 p.
McKeever DG. 1938. The effect of various methods of treatment on the
germination of seeds of some plants valuable for game and erosion control purposes [unpublished masters thesis]. Moscow, ID: University of
Idaho School of Forestry. 132 p.
Morrow EB, Darrow GM, Scott DH. 1954. A quick method of cleaning
berry seed for breeders. Proceedings of the American Society of
Horticultural Science. 63: 265.
Sambucus
1017
1018
Rydberg PA. 1965. Flora of the prairies and plains of central North
America. New York: Hafner. 969 p.
Swingle CF, comp. 1939. Seed propagation of trees shrubs and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA Soil
Conservation Service. 198 p.
USDA NRCS [USDA National Resources Conservation Service]. 1996.
PLANTS [database]. Baton Rouge, LA: USDA NRCS, National Plant
Data Center.
Van Dersal WR. 1938. Native woody plants of the United States: their erosion-control and wildlife values. Misc. Pub. 303. Washington, DC: USDA.
362 p.
SapindaceaeSoapberry family
translucent, globular drupe measuring 10 to 14 mm in diameter, usually contains a single, dark brown, hard-coated seed
(figures 1 and 2), but occasionally 2 or 3 seeds are present
(Khatamian and Abuelgasim 1986; Preston 1940). The fruits
ripen during September to October and persist on the tree
until late winter or spring. Seedcrops are usually abundant
each year (Engstrom and Stoeckeler 1941).
Collection, extraction, and storage. Fruits may be
collected any time during late fall or winter by hand-picking
or flailing them from the trees onto canvas. Although fruits
are fairly dry by this time, they still need to be spread in
shallow layers to keep them from heating. A bushel of fresh
fruits from a central Oklahoma source had a calculated
weight of 18.6 kg (41 lb) (Read 1974). Seed extraction is
facilitated by sprinkling the fruits with water twice daily
until pulp softens. Pulp can then be removed and floated
away by running the fruits through a macerator with water.
After drying, the seeds are ready for storage or use (Read
1974).
Forty-five kilograms (100 lb) of fruit will yield 13.6 to
16 kg (30 to 35 lb) of clean seeds (Read 1974) with a maximum of 37.2 kg (82 lb) reported (Swingle 1939). There are
950 to 1430 fruits/kg (430 to 650/lb) (Read 1974). Clean
seeds per weight (based on 8 samples) varied from 1,510 to
4,360/kg (685 to 1,980/lb) (Engstrom and Stoeckler 1941;
Read 1974; Swingle 1939). A fresh collection from central
Oklahoma ran 1,540 seeds/kg (700/lb), with 104% moisture
(percentage of dry weight) after 7 days of water-soaking followed by a de-pulping treatment (Reed 1974). Soundness of
12 samples averaged 77% (Read 1974). No data are available on seed longevity in cold storage, but it is likely that
dry storage at low temperatures would be satisfactory.
Pregermination treatments. Germination of stored
seeds may be slow and delayed. The chief cause is embryo
dormancy, often accompanied by an impermeable seedcoat.
Some seedlots require only stratification, whereas others
Sapindus
1019
References
Afanasiev M. 1942. Propagation of trees and shrubs by seed. Circ. C-106.
Stillwater: Oklahoma Agricultural Experiment Station. 43 p.
Davis RE, Whitcomb CE. 1974. Effects of propagation container size on top
and root development of tree seedlings. Res. Rep. P-704. Stillwater:
Oklahoma Agricultural Experiment Station: 1213.
Dirr MA. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation and uses. Champaign, IL:
Stipes Publishing Co. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Engstrom HE, Stoeckeler JH. 1941. Nursery practice for trees and shrubs
suitable for planting on the Prairie-Plains. Misc. Pub. 434. Washington,
DC: USDA. 159 p.
Khatamian H, Abuelgasim Z. 1986. Auxins aid soapberry cuttings. American
Nurseryman 164(4): 6572.
Little EL. 1950. Southwestern trees. Agric. Handbk. 9. Washington, DC:
USDA Forest Service. 109 p.
Mehra PN, Khosla PK, Sareen TS. 1972. Cytogenetical studies of Himalayan
Aceraceae, Hippocastanaceae, Sapindaceae and Staphyleaceae. Silvae
Genetica 21: 96102.
Munson RH. 1984. Germination of western soapberry as affected by
scarification and stratification. HortScience 19(5): 712713.
Phillips GR, Gibbs FJ. 1953. Forest trees of Oklahoma. Pub. 1 (rev. ed., #8).
Oklahoma City: Oklahoma Planning and Resources Board, Division of
Forestry. 135 p.
Preston RJ Jr. 1940. Rocky Mountain trees. Ames: Iowa State College Press.
285 p.
Read RA. 1974. Sapindus drummondii Hook. and Arn., western soapberry. In:
Schopmeyer CS, comp. Seeds of woody plants in the United States.
Agric. Handbk. 450. USDA Forest Service. Washington, DC: 758759.
Sheikh MI. 1979. Tree seeds respond to acid scarification. Pakistan Journal of
Forestry 29(4): 253254.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA Soil
Conservation Service. 198 p.
Tirmenstein DA. 1990. Sapindus saponaria var. drummondii. In: Fischer WC,
comp.The Fire Effects Information System [electronic database].
Missoula, MT: USDA Forest Service, Intermountain Research Station.
Vora RS. 1989. Seed germination characteristics of selected native plants of
the lower Rio Grande Valley,Texas. Journal of Range Management 42(1):
3640.
Vora RS. 1992. Restoration of the native vegetation in the lower Rio
Grande Valley, 198487. Restoration and Management Notes 10 (2):
150157.
Watson L, Dallwitz Mj. 1992 [onwards]. The families of flowering plants:
descriptions, identification, illustrations, and information retrieval.Version
14 December 2000 [website available at
http://biodiversity.uno.edu/delta/].
Sapindus
1021
ChenopodiaceaeGoosefoot family
1022
Table 1Sarcobatus vermiculatus, black greasewood: germination tests, conditions, and results
Seed source
Montana
Montana
New Mexico
Filter paper
10
10
525
11
11
Germination rate
Amt (%)
Days
30
30
30
25
26
67
91
66
2
10
Average %
germination
100
98
8093
100
88
Samples
4
4
4
Sources: Eddleman (1979), Romo and Eddleman (1985), Sabo and others (1979).
References
Blauer AC, Plummer AP, McArthur ED, Stevens R, Guinta BC. 1976.
Characteristics and hybridization of important Intermountain shrubs: 2.
Chenopod family. USDA Forest Service, Res. Pap. INT-177. Ogden, UT:
Intermountain Forest and Range Experiment Station. 42 p.
Branson FA, Miller RF, McQueen IS. 1967. Geographic distribution and factors affecting the distribution of salt desert shrubs in the United States.
Journal of Range Management 20: 287296.
Eddleman LE. 1977. Indigenous plants of southeastern Montana: 1.Viability
and suitability for reclamation in the Fort Union Basin. Spec. Pub. 4.
Missoula: University of Montana, Montana Forest and Conservation
Experiment Station. 122 p.
Eddleman LE. 1979. Germination in black greasewood (Sarcobatus vermiculatus (Hook.) Torr.). Northwest Science 53: 289294.
Sarcobatus
1023
1024
Shantz HL, Piemeisel RL. 1940. Types of vegetation in the Escalante Valley,
Utah, as indicators of soil conditions.Tech. Bull. 713. Washington, DC:
USDA. 32 p.
Stephens HA. 1973. Woody plants of the north central plains. Lawrence:
University of Kansas Press. 530 p.
Stubbendieck J, Hatch SL, Butterfield CH. 1994. North American range
plants. Lincoln: University of Nebraska Press. 493 p.
Van Dersal WR. 1938. Native woody plants of the United States, their erosion, control, and wildlife values. Misc. Pub. 303. Washington, DC: USDA.
362 p.
Welsh SL, Atwood ND, Higgins LC, Goodrich S. 1987. A Utah flora. Great
Basin Naturalist Memoirs 9: 1894.
LauraceaeLaurel family
rubbing the fruits over hardware cloth by hand or by breaking them up with mechanical macerators and washing the
debris away with water. In the South, there are about 6,200
fruits/kg (2,800/lb) (Halls 1973). In the North, seeds collected and cleaned averaged 13,000/kg (5,900/lb). In Pennsylvania, 45 kg (100 lb) of fruit yielded about 14 kg (31 lb) of
cleaned seeds (Bonner and Maisenhelder 1974).
There are no known storage tests for sassafras, but the
seeds can apparently be stored successfully for a few years
at 2 to 4 C and low moisture contents (Bonner and
Maisenhelder 1974). This behavior should place sassafras in
the orthodox seed storage grouping, although the very high
lipid content (47%) of the seeds (Bonner 1971) suggests that
long-term storage would be difficult. Soil seedbank studies
have demonstrated that seeds buried in litter retained viability for 4 years in Louisiana (Haywood 1994) and for 6 years
in West Virginia (Wendel 1977).
Sassafras
1025
References
Bonner FT. 1971. Chemical contents of southern hardwood fruits and
seeds. Res. Note SO-136. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 3 p.
Bonner FT, Maisenhelder LC. 1974. Sassafras albidum (Nutt.) Nees, sassafras. In: Schopmeyer CS, tech. coord. Seeds of woody plants in the
United States. Agric. Handbk. 450. Washington, DC: USDA Forest
Service: 761762.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Griggs MM. 1990. Sassafras albidum (Nutt.) Nees, sassafras. In: Burns RM,
Honkala BH, tech. coords. Silvics of North America.Volume 2,
Hardwoods. Agric. Handbk. 654. Washington, DC: USDA Forest Service:
773777.
Halls LK. 1973. Flowering and fruiting of southern browse species. Res. Pap.
SO-90. New Orleans: USDA Forest Service, Southern Forest
Experiment Station. 10 p.
Haywood JD. 1994. Seed viability of selected tree, shrub, and vine species
stored in the field. New Forests 8: 143154.
1026
AnacardiaceaeSumac family
Schinus L.
peppertree
Jill R. Barbour
Ms. Barbour is a germination specialist at the USDA Forest Services National Seed Laboratory,
Dry Branch, Georgia
Common name(s)
Occurrence
S. molle L.
W South America
Brazil; naturalized in S Florida & Hawaii
Schinus
1027
1028
Germination tests. The seed germination characteristics that differentiate Peruvian peppertree and Christmasberry tree may be the factors inhibiting the naturalization of
Christmasberry tree in southern California (Nilsen and
Muller 1980). The latter species may be excluded from the
California vegetation because of its slow germination rate,
which could preclude its establishment during the periods of
brief rainfall and intermittent drought (Nilsen and Muller
1980).
Germination treatments carried out in the dark at 24 C
were performed on the seeds of both species. Peruvian peppertree seeds germinate best after soaking for 5 minutes in a
10% solution of sulfuric acid (H2SO4) (Nilsen and Muller
1980); imbibition for 24 hours and stratification (30 days at
2 C) did not break dormancy in seeds of this species.
Seeds of Christmasberry tree germinate equally well
with imbibition or acid treatment, and a significant amount
of germination occurred after 30 days of 2 C stratification
(Nilsen and Muller 1980). Seeds of Peruvian peppertree are
more inhibited by cold conditions than are those of
Christmasberry tree. Seeds of neither species germinated
when incubated to 70 C for 1 hour. Laboratory germination
of Peruvian peppertree began on the 17th day and that of
Christmasberry tree began on day 28.
Germination tests performed in native soil delayed germination of Peruvian peppertree and inhibited that of
Christmasberry tree. In both greenhouse soil mix or field
soil, seeds of Peruvian peppertree germinated better than
those of Christmasberry tree, except when seeds were germinated in a winter-regimen growth chamber (15/2 C)
(Nilsen and Muller 1980a).
Nursery practices. Schinus seeds are usually grown
in pots filled with sandy clay loam. In southern California,
the seeds germinate in the winter at about 50% relative
humidity and 18/10 C temperatures (Nilsen and Muller
1980a). Christmasberry tree has a higher root net accumulation ratio, which gives it a greater drought tolerance than
Peruvian peppertree. Both Schinus species show positive relative growth rates within and below the common irradiances
under canopies of southern California coastal communities
(Nilsen and Muller 1980a).
References
Anderson E. 2002. Personal communication. Escondido, CA: Anderson
Seed Co.
Correll DS. Johnston MC. 1970. Manual of vascular plants of Texas. Renner:
Texas Research Foundation: 1881.
Eisikowitch D, Masad Y. 1980. Nectar-yielding plants during the dearth season in Israel. Bee World 61: 1118.
Eizenbrand A. 2002. Personal communication. Bet Nehemia, Israel: Keren
Kayemet LeIsrael.
Ewel JJ. 1979. The ecology of Schinus. In: Workman RW, ed. SchinusA
Technical Proceedings of Techniques for Control of Schinus in South
Florida: A Workshop for Natural Area Managers. Sanibel, FL: SanibelCaptiva Conservation Foundation: 721.
Ewel JJ. 1986. Invasibility: lessions from south Florida. Ecological Studies:
Analysis and Synthesis 58: 214230.
Ewel JJ, Ojima DS, Karl DA, DeBusk WF. 1982. Schinus in successional
ecosystems of Everglades National Park. Rep.T-676. Homestead: South
Florida Research Center. 141 p.
Habeck DH, Bennett FD, Grissell EE. 1989. First record of a phytophagous
seed chalcid from Brazilian peppertree in Florida. Florida Entomologist
72(2): 378379.
Johnson H. 1973. The International book of trees. New York: Simon and
Schuster. 288 p.
Jones DT, Doren RF. 1997. The distribution, biology and control of Schinus
terebinthifolius in southern Florida with special reference to Everglades
National Park. In: Brock JH, Pysek P, eds. Green plant invasions: studies
from North America and Europe. Leiden,The Netherlands: D. Backhuys,
Publishers: 8193.
Schinus
1029
TaxodiaceaeRedwood family
1030
seed
not easy to grow and is extremely slow-growing when propagated from seeds (Halladin 1991). It has a tendency to form
several leaders. Field planting has been done with 3+2 and
4+2 stock (Dallimore and Jackson 1967). Umbrella-pine can
also be propagated by layers or by cuttings of half-ripened
wood in summer (Bailey 1939). A nursery in Oregon propagates solely by cuttings because of faster results; Halladin
(1991) describes the technique in detail.
References
Asakawa S. 1973. Germination behavior of Sciadopitys verticillata seeds. In:
Proceedings, International Symposium on Seed ProcessingSeed
Problems; 1973; Bergen, Norway. IUFRO. 8 p.
Bailey LH. 1939. The standard cyclopedia of horticulture. New York:
Macmillan. 3639 p.
Barton LV. 1930. Hastening the germination of some coniferous seeds.
American Journal of Botany 17: 88115.
Dallimore W, Jackson AB. 1967. A handbook of Coniferae and
Ginkgoaceae. 4th ed. 4, Harrison SG, rev. New York: St. Martins Press.
729 p.
Halladin P. 1991. Sciadopitys vertricillata. American Nurseryman 174(8): 73.
Hatano K. 1972. Dormancy of Sciadopitys verticillata seed. Journal of the
Japanese Forestry Society 54(8): 264268.
McClintock E. 1992.
Trees of Golden Gate Park. Pacific Horticulture
53(3): 38.
Sciadopitys
1031
FabaceaePea family
1032
mature seed.
longitudinal section
100 seeds in damp paper toweling placed in a growth chamber at 15 C. Test conditions were maintained for 28 days,
with germination percentages recorded every 7 days; initial
germination rate for senna was 75%. Germination tests, conducted annually to test the effects of storage, were then averaged to a best germination of 92%. These annual tests
consisted of 4 replications of 50 seeds using the same initial
testing methods. The effects of temperature on germination
rates were also tested, with the following results (Kay and
others 1988):
Temperature (C) 2 5 10
Germination (%) 0 0 19
15
41
20
46
25
20
30 40
28 0
Nursery practice and seedling care. In direct-seeding trials, germination in Nevada seedlots was best at 15 to
20 C, but seeds collected at lower elevations may need
higher temperatures; no germination was observed at 5 or
40 C (Bainbridge and Virginia 1989). Nursery stock has
been outplanted at JTNP using 3.8-liter (1-gal) and 6.8-liter
(1.8-gal) containers that were 35 to 37 cm (14 to 15 in)
deep. The plants monitored after 10 months (before late
winter precipitation) had respective survival rates of 7 and
14% (CALR 1995). Monitoring continues at the site, and
figures may be different after the winter and spring rains.
Senna seedlings were noted to be susceptible to rot and
should be planted into a well-drained soil with conservative
watering (CALR 1995).
References
Kay BL. 1975. Tests of seeds of Mojave Desert shrubs [unpublished manuscript prepared for the California Department of Transportation]. Prog.
Rep. BLM Contract 53500 CT4-2(N). 24 p.
Kay BL, Graves WL,Young JA. 1988. Long-term storage of desert shrub
seed. Mojave Reveg. Notes 23. Davis: University of California
Department of Agronomy and Range Science.
Kay BL, Ross CM, Graves WL. 1977. Armed senna. Mojave Reveg. Notes 2.
Davis: University of California Department of Agronomy and Range
Science.
Perry B. 1987. Trees and shrubs for dry California landscapes. San Dimas,
CA: Land Design Publishing. 184 p.
Stark N. 1966. Review of highway planting information appropriate to
Nevada. Bull. B-7. Reno: University of Nevada, College of Agriculture,
Desert Research Institute. 71 p.
Senna
1033
TaxodiaceaeRedwood family
usually fertilized in May (Buchholz 1939). Dry weather during pollination permits better pollen dispersal and improves
seed viability (Olson and others 1990). Cone ripening time
can range from late September to mid-January, depending
on latitude, elevation, and weather (Lippitt 1996).
Trees begin to bear seeds at 5 to 15 years of age (Boe
1974). Good seedcrops occur every 5 to 7 years (Lippitt
1996), with light crops intervening. Fair to abundant crops
occurred for 5 consecutive years in north-coastal California
(Boe 1968); however, this is unusual (Lippitt 1996). Further
south in the redwood type, some stands produce seed poorly
and irregularly, whereas others frequently have fair to abundant crops (Muelder and Hansen 1961). A mature seed has a
brown wing and a slightly darker seedcoat. The wing, which
is part of the seedcoat, is about equal in width to the seed
(figure 1). Embryos have 2 cotyledons (figure 2). Opened
cones often persist through the next growing season (Boe
1974). Cones have the following quantitative characteristics
(Lott 1923; Munz 1959; Olson 1990; Roy 1965):
Seeds per cone scale
25
Average seeds per cone
60
Cone length
1.32.8 cm
Cone diameter
1.32.5 cm
Average fresh cones per weight 500/kg (227/lb)
Because of the polyploid chromosomal make-up of redwood, care should be taken to avoid in-breeding in seed
orchards. Seedlings from self-crossed seeds had lower nursery survival under stress and grew slower in the field than
those from out-crossed seeds (Libby and others 1981).
A technique for making controlled pollinations on
detached redwood cuttings has become practical. Pollination
was most effective when dry pollen was brushed between
the open scales of the female strobili. In subsequent years,
cuttings that have rooted successfully can be pollinated and
will produce viable seeds (Libby and McCutchan 1978;
Libby and others 1972).
seed.
nation rate. Experience with a magnified x-ray makes it possible to distinguish tannin-filled seeds and filled but less
viable seeds from seeds that will produce vigorous germinants.
The following seed data have been noted by Lippitt
(1996) for cleaned seedlots:
Cleaned seeds per weight
Low
167,372/kg
longitudinal
(75,920/lb)
High
259,297/kg
(117,617/lb)
Average (N = 37)
194,000/kg
(88,000/lb)
Purity
98%
Germination
60%
Sequoia
1035
or 2+0 planting stock. All seedbeds are fumigated as standard practice. Mineral soil provides the best seedbed, but
seeds will germinate in duff, on logs, and on most moist surfaces. Seeds can germinate in either shade or sunlight
(Olson and others 1990), but frost protection is important
(Lippitt 1996). Redwood seedlings need a greater supply of
soil water than most associated species (Fritz 1966). The
roots lack root hairs and, therefore, are not efficient in
References
Boe KN. 1968. Cone production, seed dispersal, germination in old-growth
redwood cut and uncut stands. Res. Note PSW-184, Berkeley, CA:
USDA Forest Service, Pacific Southwest Forest and Range Experiment
Station. 7 p.
Boe KN. 1974. Sequoia sempervirens (D. Don) Endl., redwood. In:
Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric. Handbk. 450. Washington DC: USDA Forest Service:
764766.
Buchholz JT. 1939. The embryogeny of Sequoia sempervirens with a
comparison of sequoias. American Journal of Botany 26(4): 248257.
Fritz E, Rydelius JA. 1966. Redwood reforestation problems: an experimental approach to their solution. San Francisco: Foundation for American
Resource Management: 130 p.
Harlow WM, Harrar ES. 1969. Textbook of dendrology. New York:
McGraw-Hill: 171176.
ISTA [International Seed Testing Association]. 1993. Rules for testing seeds:
rules 1993. Seed Science and Technology 21 (Suppl.): 1259.
Libby WJ, McCutchan BG. 1978. Taming the redwood. American Forests.
84(8): 3739.
Libby WJ, Kiang YT,Yinhart YB. 1972. Control pollination seeds from cutting
of coast redwood. Silvae Genetica 21(1/2): 1720.
Libby WJ, McCutchan BG, Miller CI. 1981. Inbreeding depression in selfs of
redwood. Silvae Genetica 30(1): 1525.
Lippitt L. 1996. Unpublished data. Davis: California Department of Forestry
and Fire Protection, Lewis A. Moran Reforestation Center.
Lott H C. 1923. The production and viability of redwood (Sequoia sempervirens) seed [masters thesis]. Berkeley: University of California. 32 p.
1036
TaxodiaceaeRedwood family
2540
39
230
Cone length
59 cm
5 oz/bu
113,625/kg
51,530/lb
High
199,810/kg
90,620/lb
173,100/kg
78,500/lb
Sequoiadendron
1037
Table 1Sequoiadendron, giant sequoia: pregermination treatments, germination test conditions, and results
Cold
stratification
period (days)
Daily
light (hr)
0
0
0
0
28
0
<16
24
24
24
15
5
20
30
20
20
1038
Days
Avg %
germination
Samples
32
32
32
32
28
28
43.1
4.1
40.9
5.6
38.5
30.3
10
10
10
10
2
3
ArecaceaePalm family
1039
Cultural treatments have been used to stimulate flowering. Fertilizer (10% N, 5% P2O5, 5% K2O) and dolomite
lime (49% CaCO3), 36% MgCO3, and 10% Mg) applied at a
rate of 155 g per plant around the plants drip line, did not
influence flowering. But when crowns were clipped and
plants fertilized, there was a significant elevation in flowering during the second growing season: 18% flowering in
treated plants compared to 4% flowering in control plants
(Abrahamson 1999). Saw-palmettos that were only clipped
had a 22% flowering response.
The fruit is a drupe measuring about 1.5 to 3 cm long
and 15 to 20 mm in diameter, that is ovoid-oblong, green or
yellow before ripening, and bluish to black when ripe
(Hilmon 1968; McCurrach 1960; Vines 1960) (figure 1).
Immature green fruits are present from May through July,
turn orange by August, and ripen to bluish-black in
September and October (Carrington and others 2001). Each
drupe contains a single globose seed (figures 1 and 2), and
the embryo is laterally oriented (Bailey 1976).
Each inflorescence typically produces 4 to 5 kg (9 to 11
lb) of fruits (Vines 1960) and can produce up to 12 kg (27
lb) in a good year (Carrington and others 2000). The average fruit yield for a site is 200 kg/ha; yields can vary from
150 kg/ha to over 1,500 kg/ha (Carrington and others 1997).
In Florida, anthracnose infectionby Colletotrichum
gloeosporioides (Penz.) Penz. & Sacc.has been identified
as a major factor (90%) in fruit loss. The remaining 10% of
the loss is caused by a caterpillar (Atheloca sp.) (Carrington
and others 2001).
Collection of fruits. In south Florida, fruit harvesting
begins in August, when fruits turn orange. In south Georgia,
harvesting begins in early September, when fruits begin to
turn black. The fruits are collected by snapping the panicles
by hand, cutting them with pruning shears, or shaking the
attached fruits into burlap bags, plastic sheets, or the bed of
a truck. Seeds are available commercially within the natural
range of the species.
Extraction and storage of seeds. Palmetto fruits can
be dried in the sun or in indoors in bins or tobacco barns.
The fruits are piled about 0.6 to 1.0 m high (Carrington and
others 2000). Fruits are dried at 54 C (not to exceed 60 C )
for about 3 days; fruits in bins are turned every 12 hours
(Carrington and others 2000). The initial moisture content is
about 66% fresh weight; the fruits are then dried to a maximum of 10% for storage.
Large suppliers store the freshly harvested fruits in wet
tanks holding 100,000 kg (250,000 lb). The berries are conveyed to a stainless steel dryer with a capacity of drying
300,000 kg (750,000 lb) per day. The dryer takes an hour to
dry a batch of fruits, thus preserving more of the fatty acids
1040
longitudinal
Serenoa
1041
References
Abrahamson WG. 1984. Species responses to fire on the Florida Lake
Wales Ridge. American Journal of Botany 7 (1): 3543.
Abrahamson WG 1999. Episodic reproduction in two fire-prone palms,
Serenoa repens and Sabal etonia (Palmae). Ecology 80(1): 100115.
Bailey LH. 1939. The standard cyclopedia of horticulture. New York:
Macmillan. 3639 p.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan. 1290 p.
Carpenter WJ. 1986. Temperature and imbibition effects on seed germination of Sabal palmetto and Serenoa repens. Horticultural Science [1987
August] 22(4): 660.
Carpenter WJ. 1987. Seed germination of Serenoa repens. Proceedings of
the Annual Meeting of the Florida State Horticultural Society [1986] 99:
158159.
Carrington ME, Mullah JJ, Roma F. 1997. Saw palmetto: a fountain of youth.
Proceedings of the American Forage and Grassland Council [1997] 6:
233237.
Carrington ME, Mullahey JJ, Krewer G, Boland B, Affolter J. 2000. Saw palmetto (Serenoa repens): an emerging forest resource in the southeastern
United States. Southern Journal of Applied Forestry 24(3): 129134.
1042
Carrington ME, Roberts PD, Urs RR, McGovern RJ, Seijo TE, Mullahey JJ. 2001.
Premature fruit drop in saw palmettos caused by Colletotrichum
gloeosporioides. Plant Disease 85(2): 122125.
Fisher JB,Tomlinson PB. 1973. Branch and inflorescence production in saw
palmetto (Serenoa repens). Principes 17: 1019.
Ganzera M, Croom EM, Khan IA. 1999. Determination of the fatty acid
content of pumpkin seed, pygeum, and saw palmetto. Journal of
Medicinal Food [1999 November 1] (1): 2127.
Hilmon JB. 1964. Plants of the Caloosa Experimental Range. Res. Pap. SE12. Asheville, NC: USDA Forest Service, Southeastern Forest
Experiment Station. 24 p.
Hilmon JB. 1968. Autecology of saw-palmetto (Serenoa repens (Bartr.)
Small) [unpublished PhD dissertation]. Durham, NC: Duke University.
McCurrach JC. 1960. Palms of the world. 290 p. Harper Bros., New York.
Perkins D. 2002. Personal communication. LaBelle, FL: Perkins Nursery.
Radford AE, Ahles HE, Bell CR. 1964. Manual of the vascular flora of the
Carolinas. Chapel Hill: University of North Carolina Press. 1183 p.
SPHC [Saw Palmetto Harvesting Company]. 2002. International supplier of
saw palmetto (Serenoa repens) raw materials... [website: www.sawpalmetto.com/]. Frostproof, FL.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest.
Austin: University of Texas Press. 1104 p.
ElaeagnaceaeOleaster family
Shepherdia Nutt.
buffaloberry
Scott C.Walker
Mr. Walker is the project leader for the Utah Division of Wildlife Resources
Great Basin Research Center, Ephraim, Utah
food for small and large game animals (Knudson and others
1990). This species is regarded as poor to fair forage for
sheep, deer (Odocoileus spp.), and elk (Cervus elaphus), and
generally considered worthless for cattle. The fruits provide
abundant and nutritious food and are highly sought after by
birds (Mozingo 1987; Wasser 1982). The berries are edible
and were used by Native Americans and are still commonly
used as they make excellent jelly (Borland 1994; Knudson
and others 1990; Lackschewitz 1991).
Russet buffaloberry is a thornless, deciduous, small to
medium shrub with a characteristically spreading growth
form, 1 to 3 m tall at maturity (Lackschewitz 1991;
Mozingo 1987; Stubbendieck and others 1986). Twigs are
slender, round, and densely scurfy with rusty, bran-like
scales. Leaves, which are paired, have a bright green upper
surface and paler lower surface with conspicuous brown
scales (Lackschewitz 1991; Welsh and others 1987). This
species is very cold and drought hardy and it can grow in a
variety of habitat types. It is typically found along the banks
of streams, and on moist open wooded slopes at 1,000 to
3,400 m. It can also be found on sandy or rocky, often sterile, soils. At its southern extremity, it is confined to the higher vegetation zones in the mountains (Link 1993; Mozingo
1987; Thilenius and others 1974).
Common name
Occurrence
S. rotundifolia Parry
Lepargyrea rotundifolia (Parry) Green
S Utah, N Arizona
Shepherdia
1043
Russet buffaloberry has little or no browse value for cattle and is only fair for sheep before frost. The berries are bitter and though not highly palatable to humans are eaten by
birds and other wildlife (Lackschewitz 1991; Stubbendieck
and others 1986).
Roundleaf buffaloberry has a low sprawling habit and is
mainly 1 to 2 m tall, and 1 to 4 m wide. The thornless
brachlets are covered with small white to yellowish hairs
often appearing silver. The thick, persistent, somewhat evergreen leaves are silvery green above, and pale densely
scurfy beneath, and as the name implies, circular or oval in
outline (Welsh and others 1987). This species inhabits
warm, dry, sandy or rocky slopes and occurs from southern
Utah into the Grand Canyon region of Arizona throughout
the saltbrush, sagebrush, and piyon zones. Welsh and others (1987) describe roundleaf buffaloberry thusly: This is a
beautiful shrub. It festoons slopes with silvery clumps. It is
reported to have some value as a winter browse in southeastern Utah.
Flowering and fruiting. Buffaloberries are dioecious.
The small, petal-less, yellow to yellowish green flowers are
borne single or clustered at the nodes. Plants resume growth
in very early spring, usually soon after snowmelt. Flowering
occurs quite early in the season (March to April), before or
soon after the leaves appear. Fruits mature in late summer
and fall (late June to September), varying with environment
and source of planting stock (Borland 1994; Mozingo 1987;
Thilenius and others 1974; Vories 1981; Wasser 1982).
Fruits are 3.2 to 8.5 mm in diameter and drupe-like,
with a solitary smooth achene or small nutlet enveloped in a
fleshy perianth. Color of mature fruits vary from orange-red
(silver buffaloberry), red-yellow (russet buffaloberry), to silvery (roundleaf buffaloberry) (McTavish 1986; Mozingo
1987; Smith 1987; Wasser 1982; Welsh and others 1987).
Cleaned achenes are used as seeds (figures 1 and 2).
Minimum seed-bearing age is 4 to 6 years (Thilenius and
others 1974).
Seedcrop quality and quantity can vary from year to
year. McTavish (1986) reports that one of the major propagating problems with russet buffaloberry is poor seed quality. Researchers have obtained widely varying germination
percentages from year to year under identical treatments.
This seems to be due to poor embryo development.
Therefore, it is suggested that seed collectors check the
seeds before collection to ensure that proper embryo development has taken place (McTavish 1986).
Collection of fruits. The fruits may be harvested by
stripping or flailing them from the bushes onto canvas; they
may also be picked by hand or collected from the ground.
The use of mechanical shakers has shown to be effective in
harvesting the seed of silver buffaloberry (Halderson 1986).
Heavy gloves should be used when collecting this species to
avoid injury from the thorns. Care should be taken when
1044
The seeds are orthodox and should be stored dry, in cool the fall, but seeds that are prechilled for 3 months can be
conditions, optimally at 5 C. Seed can be stored for 4 to 5
sown in spring, or probably later where late summer moisyears while maintaining good viability (Thilenius and others ture is more reliable, or with irrigation (Thilenius and others
1974; Vories 1981). For short-term storage, seed extraction
1974).
is not necessary. The fruits may be spread out in a thin layer
Silver buffaloberry can be propagated by cuttings, and
and dried. For short-term storage of fruits, place them in
wildlings can be transplanted successfully. Success of propaopen plastic bags under cool-dry conditions. Care should be gating russet buffaloberry from cuttings can vary. Vories
taken to prevent heating of the collected fruits (Link 1993;
(1981) reports that it roots well from cuttings, whereas
Thilenius and others 1974). Seed quality has not been stanMcTavish (1986) reports that attempts at propagation by cutdardized. Minimum standards established by the USDI Fish
tings were largely unsuccessful. Roundleaf buffaloberry is
and Wildlife Service (Wasser 1982) are 90% purity and
generally grown from seeds because cuttings do not do well
about 60% germination.
(Borland 1994; Vories 1981; Wasser 1982).
Germination. A physiologically dormant embryo, and
physical dormancy due to impenetrable seedcoats, are the
Figure 3Shepherdia argenta, silver buffaloberry:
major problems affecting germination (McTavish 1986;
seedling
development at 1, 9, and 38 days after germination.
Thilenius and others 1974). Two generally accepted methods
of breaking dormancy are scarification with sulfuric acid
and moist cold stratification (table 2). After pretreatment,
the majority of viable seeds of silver buffaloberry germinate
in 20 days. Some seeds do delay germination up to 60 days
(Wasser 1982). Germination is epigeal (figure 3).
Nursery practice and seeding. In nursery practice,
seeds are planted 6 mm (1/4 in) deep and covered with up to
2.5 cm (1 in) of mulch. This suggests that seeds could be
planted, perhaps to advantage, at depths up to 2 cm (3/4 in)
in coarse, dry, and loose soil or in fall under wildland conditions. About half of the viable seeds sown produce usable
1+0 seedlings in nurseries, whereas only 5 to 15% establishment would be good survival from seeding under dryland
field conditions (Thilenius and others 1974; Vories 1981;
Wasser 1982).
The recommended seeding rate for wildland seedings is
1.1 to 2.2 kg/ha (1 to 2 lb/ac) in seeding mixtures totaling
11 to 34 kg/ha (10 to 30 lb/ac) (Wasser 1982). In nursery
row plantings, seeds can be sown in rows at a rate of 100 to
160 viable seeds/m (30 to 50/ft). Seeds should be sown in
Pretreatment
Germination treatment
S. argentea
S. canadensis
S. rotundifolia
170 days)
30 days)
170 days)
170 days)
Percent germination
93
7186
94
89
80
80
8090
8090
86
Sources: Borland (1994, 1996), Jorgensen (1995), McTavish (1986),Thilenius and others (1974),Vories (1981).
Shepherdia
1045
References
Borland J. 1994. Shepherdia rotundifolia. American Nurseryman 179(7): 106.
Borland J. 1996. Personal communication. Denver, CO: Genesee Open
Space.
Halderson JL, Howard CG. 1986. Seed harvester development for rangeland shrubs. Pap. 86-1092. American Society of Agricultural Engineers.
Jorgensen K. 1995. Personal communication. Ephraim UT: Utah Division of
Wildlife Resources.
Knudson MJ, Haas RJ,Tober DA, Darris DC, Jacobson ET. 1990.
Improvement of chokecherry, silver buffaloberry, and hawthorn for conservation use in the northern plains. In: McArthur ED, Romney EM,
Smith SD,Tueller PT, comps. Proceedings, Symposium on Cheatgrass
Invasion, Shrub Die-off, and Other Aspects of Shrub Biology and
Management; 1989 April 57; Las Vegas, NV. Gen.Tech. Rep. INT-276.
Ogden, UT: USDA Forest Service, Intermountain Research Station:
291299.
Lackschewitz K. 1991. Vascular plants of west-central Montana: identification
guidebook. Gen.Tech. Rep. INT-277. Ogden, UT: USDA Forest Service,
Intermountain Research Station. 224 p.
Link E, ed. 1993. Native plant propagation techniques for national parks.
East Lansing, MI: USDA NRCS, Rose Lake Plant Materials Center. 197 p.
1046
SapotaceaeSapodilla family
seed
1012 kg/50 kg
(1012 lb/50 lb)
12,500/kg (5,700/lb)
94%
88%
Sideroxylon
1047
References
Afanasiev M. 1942. Propagation of trees and shrubs by seed. Circ. #106.
Stillwater: Oklahoma Agricultural Experiment Station.
Bonner FT, Schmidtling RC. 1974. Bumelia lanuginosa (Michx.) Pers., gum
bumelia. In: Schopmeyer CS, tech. coord. Seeds of woody plants in the
United States. Agric. Handbk. 450. Washington, DC: USDA Forest
Service: 258259.
1048
Clark R. 1940. A hardy woody plant new to horticulture. Bull. 28. St. Louis:
Missouri Botanical Garden: 216220.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
SimmondsiaceaeJojoba family
Simmondsia
1049
Figure 1Simmondsia chinensis, jojoba: longitudinal section through a seed (top) and exterior view of a seed
(bottom).
conducive to this activity, a problem that is solved in cultivation by bottom-pruning (Yermanos 1974). For small lots,
seeds can be collected by beating the branches over a hopper
or by hand-stripping them when still slightly green, the
hard-dough stage (Nord and Kadish 1974). Seeds picked
green should be allowed to dry in a shady, well-ventilated
area. A pneumatic device has been developed for commercial harvest (Coates and Yacizi 1991). If collected intact, the
capsules may be broken up using a barley de-bearder or
hammermill. The seeds can then be cleaned of debris and
unfilled seeds in a conventional fanning mill or air-screen
cleaner. The purity and viability of cleaned seedlots are usually high (Nord and Kadish 1974).
Jojoba seeds are quite variable in size, both within and
among seedlots (Yermanos 1979). Nord and Kadish (1974)
report an among-lot mean seed weight range of 660 to
3,300/kg (300 to 1,500/lb). Ismail (1988) sorted seeds of a
single lot into 3 size classes with the following mean values:
Seeds/weight
/kg
/lb
Length
cm
in
small
2,300
1,045
0.93
1/3
medium
1,300
590
1.38
1/2
large
1,060
480
1.84
3/4
Figure 2Simmondsia chinensis, jojoba: seedling development at 3, 7, and 14 days after germination.
References
Al-Ani HA, Strain BR, Mooney HA. 1972. The physiological ecology of
diverse populations of the desert shrub Simmondsia chinensis. Journal of
Ecology 60: 4157.
Brooks WH. 1978. Jojobaa North American desert shrub: its ecology,
possible commercialization, and potential as an introduction into other
arid regions. Journal of Arid Environments 1: 227236.
Castellanos AE, Molina FE. 1990. Differential survivorship and establishment
in Simmondsia chinensis (jojoba). Journal of Arid Environments 19: 6576.
Coates W,Yazici E. 1991. Pneumatic device for harvesting jojoba seeds.
Transactions of the ASAE 34: 23042308.
Dunstone R. 1990. Jojoba (Simmondsia chinensis). Plant Varieties Journal 3:
1416.
Dunstone R. 1991. Jojoba (Simmondsia chinensis) variety: Wadi wadi. Plant
Varieties Journal 4: 1920.
Foster KE. 1980. Environmental consequences of an industry based on
harvesting the wild desert shrub jojoba. Bioscience 30: 255258.
Ismail AMA. 1988. The ecological and agronomic role of seed polymorphism in Simmondsia chinensis. Journal of Arid Environments 14: 3542.
Kumari A, Chikara J, Iyengar ERR. 1991. Variability in Simmondsia chinensis
under semiarid conditions. Indian Journal of Genetics and Plant Breeding
51: 8588.
Milthorpe PL. 1989. The potential of jojoba (Simmondsia chinensis) in New
South Wales: 2. Some factors affecting yield. Australian Journal of
Experimental Agriculture 29: 389395.
Munz PA. 1974. A flora of southern California. Berkeley: University of
California Press. 1086 p.
Muthana KD. 1981. Jojoba: an oil-yielding desert shrub. Indian Farming 31:
2831.
Niklas KJ, Buchmann SL. 1985. Aerodynamics of wind pollination in
Simmondsia chinensis (Link) Scheider. American Journal of Botany 72:
530539.
Nimir MN, Ali-Dinar HM. 1991. Jojoba, a new cash crop in marginal lands.
Acta Horticulturae 270: 369372.
Nord EC, Kadish A. 1974. Simmondsia chinensis (Link.) C. K. Schneid., jojoba.
In: Schopmeyer CS, tech. coord. Seeds of woody plants in the United
States. Agric.Handbk. 450. Washington, DC: USDA Forest Service:
774776.
Palzkill DA,Younes MH, Hogan L. 1989. AT-1310, AT-1487, and AT-3365:
clonal jojoba germplasm for horticultural use. HortScience 24: 526527.
Rao PHV, Iyengar ERR. 1982. Studies in seed morphology and germination
in jojoba (Simmondsia chinensis Link). Current Science 51: 516518.
Sherbrooke WC. 1976. Differential acceptance of toxic jojoba seed
(Simmondsia chinensis) by four Sonoran Desert heteromyid rodents.
Ecology 57: 596602.
Sherbrooke WC. 1977. First year seedling survival of jojoba (Simmondsia
chinensis) in the Tucson Mountains, Arizona. Southwestern Naturalist 22:
225234.
Yermanos DM. 1974. Agronomic survey of jojoba in California. Economic
Botany 28: 161174.
Yermanos DM. 1979. Jojoba: a crop whose time has come. California
Agriculture 33: 411.
Simmondsia 1051
SolanaceaeNightshade family
Solanum dulcamara L.
bitter nightshade
John C. Zasada and John A. Crossley
Dr. Zasada retired from the USDA Forest Services North Central Research Station; Dr. Crossley retired
from the USDA Forest Services Northeastern Forest Experiment Station
Growth habit, occurrence, and use. Bitter nightshadeSolanum dulcamara L., also known as European bittersweetis a climbing perennial vine, somewhat woody at
the base. It grows to a height of 1.8 to 3.6 m. It is native in
Europe, northern Africa, and eastern Asia. In its natural
range in Europe, it occurs on sites ranging from wet and
shaded to dry and exposed. Its presence indicates a habitat
in which the moisture regime may fluctuate from moist to
waterlogged. It occurs on mineral to peat soils characterized
by a high nitrogen supply and with a pH range of 4.8 to 7.9
(Pegtel 1985). Pegtel (1985) has briefly summarized many
aspects of the species biology within its natural range.
Naturalized in North America, it is often found in moist
thickets, from Nova Scotia to Minnesota, south to North
Carolina and Missouri (Curtis 1959; Gleason 1958) and
from Idaho to Washington and California (Crossley 1974).
Bitter nightshade has been cultivated since 1561, chiefly for
ornamental purposes, but is now often considered invasive.
The fresh berries are poisonous to most humans and fatal to
rabbits, but some birds and other wildlife eat them with
impunity. Gunn and Gaffney (1974) state that any medicinal
values are offset by the poisonous properties of the fruits
and berries. Recommendations for medicinal use are only
for external application; it has been used as an ingredient in
ointments.
Leaves of the typical variety are minutely pubescent or
nearly glabrous. Many plants from Nova Scotia to Ontario,
however, have distinctly hairy leaves and branches. These
plants have been segregated as the variety villosissimum
Desv. (Gleason 1958). Mathe and Mathe (1973) found that
plants from western and eastern European sources differ in
their alkaloid chemistry, suggesting the presence of chemical
taxa within the species.
Bitter nightshade is 1 of 1,200 species in the genus,
most of which occur in the tropical and subtropical regions
of both hemispheres (Crossley 1974). The genus contains
economically important agricultural speciessuch as potato
(S. tuberosum L.) and eggplant (S. melongena L.)that have
been domesticated through plant breeding and for which
1052
there is a large amount of information available. (The tomato genusLycopersiconis also a member of the
Solanaceae.) The nightshade genus also contains a number
of agricultural weed species that affect the production of
crops such as sorghum, soybeans, and cotton, and for which
there is a significant amount of information available on various aspects of seed biology (for example, Rogers and Ogg
1981). Some of the information may be useful for understanding the seed biology of bitter nightshade, but we did
not review this information in detail. Seed characteristics of
42 economically important Solanum spp., including bitter
nightshade, have been described (Gunn and Gaffney 1974).
Flowering and fruiting. The violet flowers, which
occur in long peduncled cymes, bloom from July to August.
Bumble bee speciesBombus spp.are important pollinators (Liu and others 1975). The ovoid to ellipsoid scarlet
berries ripen from August to October.
The fruit is a juicy berry 8 to 11 mm in diameter that
contains from 40 to 60 seeds. The seeds are 2 to 3 mm by
1.7 to 2.5 mm by 0.7 to 1 mm, strongly flattened, tannish
pink, irregular disks, and dully glistening as if coated with
fine sugar. The embryo is coiled within the seed (figure 1)
(Gleason 1958; Gunn and Gaffney 1974). In cross-section,
the embryo is seen as 4 small round structures within the
endosperm; the presence of 2 or 3 sections of embryo in a
cross-section of the seed is common in the genus (Gunn and
Gaffney 1974). Good seedcrops are borne almost annually.
Collection of fruits; extraction and storage of seeds.
Seeds may be collected from July to September by handpicking the ripe berries (Crossley 1974). The fruits may be
rubbed through a 10-mesh (2-mm) soil screen, and the pulp
and empty seeds floated off with water. Large-scale extraction can be done in a macerator. Parts of the fruit may
adhere to the seed (Gunn and Gaffney 1974). Crossley
(1974) found in 1 collection that there were about 700,00
seeds/kg (350,000/lb) and that, after careful cleaning, purity
should be 99 to 100% and soundness from 92 to 99%. Seeds
from genetically transformed plants had seed weights that
Stratification
(days)
0
0
0
0
1
3
6
0
1
3
6
Germination conditions*
Days
Temp (C)
ND
ND
ND
ND
ND
ND
ND
ND
ND
ND
ND
20/30
20/30
20/30
25
25
25
25
15
15
15
15
Total
germination (%)
95
95
95
5
80
85
75
0
5
30
65
Solanum
1053
References
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
Boyd JW. 1981. Effects of certain environmental factors on the growth and
development of silverleaf nightshade (Solanum eleagnifolium).
Dissertation Abstracts International [B] 42(9): 35133514.
Crossley JA. 1974. Solanum dulcamara L., bitter nightshade. In: Schopmeyer
CS, tech. coord. Seeds of woody plants in the United States. Agric.
Handbk. 450. Washington, DC: USDA Forest Service: 777778.
Curtis JT. 1959. The vegetation of Wisconsin. Madison: University of
Wisconsin. 657 p.
Gleason HA. 1958. The new Britton and Brown illustrated flora of the
northeastern United States and adjacent Canada. New York: New York
Botanical Garden.
Gunn CR, Gaffney FB. 1974. Seed characteristics of 42 economically important species of Solanaceae in the United States.Tech. Bull. 1471.
Washington, DC: USDA. 34 p.
Lee JY, Davey MR. 1988. Morphological variation of Solanum dulcamara
plants transformed with Ri plastids [abstract]. Research Report of the
Office of Rural Development (Korea) [Biotechnology] 30(3): 2127.
1054
Liu HJ, Macfarlane RP, Pengelly DH. 1975. Relationships between flowering
plants and four species of Bombus (Hymenoptera: Apidae) in southern
Ontario [abstract]. Canadian Entomologist 107: 577588.
Mathe I, Mathe I Jr. 1973. Data to the European area of the chemical taxa
of Solanum dulcamara L. [abstract]. Acta Botanica Academiae Scientiarum
Hungaricae 19(1/4): 441451.
Pegtel DM. 1985. Germination in populations of Solanum dulcamara L. from
contrasting habitats. New Phytologist 100(4): 671679.
Roberts HA, Lockett PM. 1977. Temperature requirements for germination
of dry-stored, cold-stored and buried seeds of Solanum dulcamara L.
New Phytologist 79(3): 505510.
Rogers BS, Ogg AG Jr. 1981. Biology of weeds in the Solanum nigrum complex (Solanum section Solanum) in North America [abstract]. Agric. Rev.
Manual (West. Ser.) ARM-W-23. Washington, DC: USDA Science and
Education Administration. 30 p.
FabaceaePea family
Sophora L.
sophora
Kristina F. Connor
Dr. Connor is a plant physiologist at the USDA Forest Services Southern Research Station,
Auburn University, Alabama
Growth habit, occurrence and use. There are 50 to
70 woody and herbaceous species of the sophora genus
found in the warm temperate and tropical regions of the
world (Little 1979). Of the species found in the United
States, 2 are discussed in detail here.
Mescalbean (also known as frijolito and Texas mountain-laurel)Sophora secundiflora (Ortega) Lag. ex DC.
is a small tree of western Texas, New Mexico, and northern
Mexico (Little 1976; Ruter and Ingram 1990). Mamane
Sophora chrysophylla (Salisb.) Seem.is found in the
forests of Hawaii. Mescalbean is a favored tree for landscaping in areas with alkaline soils and moderate drought (Ruter
and Ingram 1990). Its seeds reportedly have hallucinogenic
properties (Murakoshi and others 1986) and contain many
alkaloids (Hatfield and others 1977; Izaddoost 1975; Keller
1975; Keller and others 1976). Vines (1960) lists 2 other
southwestern species: silverbush (Sophora tomentosa L.), an
evergreen shrub, and Texas sophora (Sophora affinis Torr. &
Gray), a small tree. The Japanese pagoda tree (or Chinese
scholar tree) (S. japonica L.) is planted in the United States
as an ornamental (LHBH 1976; Wasson 2001).
seed.
Sophora
1055
References
Hatfield GM,Valdes LJJ, Keller WJ, Merrill WL, Jones VH. 1977. An investigation of Sophora secundiflora seeds (mescalbeans). Lloydia 40(4): 374383
[Horticultural Abstracts 1973+].
Izaddoost M. 1975. Alkaloid chemotaxonomy of the genus Sophora.
Phytochemistry 14(1): 203204 [Horticultural Abstracts 1973+].
Keller WJ. 1975. Alkaloids from Sophora secundiflora. Phytochemistry 14(10):
23052306 [Herbage Abstracts 1973+].
Keller WJ, Hatfield GM,Valdes LJ. 1976. Isolation of lupinine and DELTA 5dehydrolupanine from Sophora secundiflora. Lloydia 39(6): 472
[Horticultural Abstracts 1973+].
LHBH [Liberty Hyde Bailey Hortorium]. 1975. Hortus third: a concise dictionary of plants cultivated in the United States. New York: Macmillan.
1290 p.
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Little EL Jr., Skolmen RG. 1989. Common forest trees of Hawaii (native and
introduced). Agric. Handbk. 679. Washington, DC: USDA Forest Service.
321 p.
1056
RosaceaeRose family
seeds.
longitudinal sec-
Sorbaria
1057
1058
References
Krssmann G. 1960. Handbuch der Laubgehlze [in German]. 2 vols. Berlin:
Parey. 495 and 608 p.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan. 1290 p.
Rehder A. 1940. Manual of cultivated trees and shrubs hardy in North
America. 2nd ed. New York: Macmillan. 996 p.
Rosendahl CO. 1955. Trees and shrubs of the upper Midwest. Minneapolis:
University of Minnesota Press. 411 p.
Rudolf PO. 1974. Sorbaria sorbifolia, Ural false spirea. In: Schopmeyer CS,
tech. coord. Seeds of woody plants in the United States. Agric. Handbk.
450. Washington, DC: USDA Forest Service: 779.
Schnelle MA. 1990. Sorbaria sorbifolia, Ural false spirea. American
Nurseryman 171(June 1): 122.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA Soil
Conservation Service. 198 p.
RosaceaeRose family
Sorbus L.
mountain-ash
William I. Stein
Dr. Stein is a forest ecologist emeritus at the USDA Forest Services Pacific Northwest Research Station,
Corvallis, Oregon
Growth habit, occurrence, and uses. The mountainash genusSorbusincludes 80 to 100 species of deciduous trees and shrubs that are distributed in the Northern
Hemisphere (Chalupa 1992; Little 1979). Mountain-ash, like
other genera in the Rosaceae, is a plastic genus comprised of
poorly defined taxa that show extensive introgression where
their ranges meet or overlap (Calder and Taylor 1968;
McAllister 1985). Geographic races probably have developed, especially in European mountain-ash, as evidenced by
its wide range and its several forms and varieties. Hybrids
are common among species of mountain-ash, yet the seeds
of several species are produced asexually and the resulting
progeny are always true to the parent (McAllister 1985;
Wright 1981). Hybrids between species of mountain-ash and
chokeberry (Aronia) or serviceberry (Amelanchier) also
occur (Rehder 1940).
Common name(s)
Occurrence
S. americana Marsh.
Pyrus americana DC.
American mountain-ash,
mountain-ash,
small-fruited mountain-ash
European mountain-ash,
rowan-tree, rowan
Newfoundland W to Manitoba, S to E
Tennessee, N Georgia, & N South Carolina
S. aucuparia L.
Pyrus aucuparia (L.) Gaertn.
S. decora (Sarg.) Schneid.
Pyrus americana DC. var. decora Sarg.
S. americana var. decora (Sarg.) Sarg
Pyrus decora (Sarg.) Hyland
S. scopulina Greene
S. cascadensis G.N. Jones
Pyrus scopulina (Greene) Longyear
S. andersonii G.N. Jones
S. sitchensis M. Roemer
S. californica Greene
Pyrus sitchensis (Roem.) Piper
S. cascadensis G.N. Jones
showy mountain-ash,
mountain-ash,
large-fruited mountain-ash
Greene mountain-ash,
western mountain-ash
Sitka mountain-ash,
Pacific mountain-ash,
western mountain-ash,
California mountain-ash
Sources: Chalupa (1992), Little (1979), Rosendahl (1955),Viereck and Little (1972).
Sorbus
1059
a cluster
Flowering
Fruit ripening
Seed dispersal
S. americana
S. aucuparia
S. decora
S. scopulina
S. sitchensis
MayJuly
MayJuly
MayJuly
MayJuly
JuneAug
AugOct
AugOct
AugNov
July
AugOct
AugMar
Aug winter
AugMar
JulyDec
Auglate winter
Sources: Fernald (1950), Harris and Stein (1974), Hitchcock and others (1961), Miller and others (1948). Rehder (1940), Rosendahl (1955),Van Dersal (1938),Viereck and
Little (1972).
Table 3Sorbus, mountain-ash: growth habit, height, fruit diameter, and color of ripe fruit
Species
Growth habit
S. americana
S. aucuparia
S. decora
S. scopulina
S. sitchensis
Shrub
Tree
Shrub
Shrub
Shrub
or tree
or tree
or tree
or tree
Height at
maturity (m)
410
520
612
16
16
Fruit
diameter (mm)
46
810
812
810
1012
Sources: Fernald (1950), Hitchcock and others (1961), Rehder (1940), Rosendahl (1955),Viereck and Little (1972).
1060
Figure 3Sorbus aucuparia, European mountain-ash: longgitudinal section through a seed, showing large cotyledons.
1061
Species
S. americana
S. aucuparia*
S. decora
S. sitchensis
52
40
Seed yield/
fruit vol
kg/hl
lb/bu
1.32.6
1.3
12
Cleaned seeds
Range
/kg
/lb
183,000521,000
229,300374,800
146,400385,000
83,000236,300
104,000170,000
66,400174,600
Average
/kg
/lb
319,400
286,200
280,400
290,800
144,900
129,800
127,200
131,900
Samples
9
10
1
8
Sources: Harris and Stein (1974), King (1947), McKeever (1938), Mirov and Kraebel (1939), Swingle (1939),Van Dersal (1938).
* Data represent seeds only from North American sources.
The number of dry fruits in one sample was 13,690/kg (6,210/lb) (Mirov and Kraebel 1939).
the total time required, but did not improve the high total
germination. Lenartowicz (1988) reached a similar conclusion but recommended that European mountain-ash seeds be
stratified at 20 C for 6 weeks and then germinated at 3 C
to gain the benefits of a shortened period during which most
germination occurs. Mechanical or chemical scarification of
the seeds has not shortened the stratification or germination
period (Flemion 1931; Harris and Hilton and others 1965;
Stein 1974) but sulfuric acid treatment increased total germination in one instance (Hilton and others 1965).
The standard germination test procedure for mountainash species requires pre-chilling the seeds for 4 months at 3
to 5 C followed by germination for 28 days at alternating
temperatures of 20 and 30 C (ISTA 1996). It is noteworthy
that the highest germination (90% or higher) for mountainash seeds was obtained in various lengthy comparison tests
at markedly lower germination temperatures than those prescribed in the standard testat 1 to 3 C (table 5). In fact,
the same substrate and temperature were used during many
tests for both stratification and germination. Given enough
time, seeds of American and European mountain-ashes completed germination under moist low-temperature conditions
(Flemion 1931; Harris and Stein 1974; Lenartowicz 1988;
Zentsch 1970), a capability also demonstrated by other
species of mountain-ash (Nikolaeva 1967).
Review of the methods employed and results obtained in
comparison studies leads to the conclusion that the standard
germination test for mountain-ash needs a firmer foundation.
Hints among published results point to 2 aspects that
deserve further investigationafter-ripening and germination temperature. After-ripening requirements may vary,
depending on when and where fruits are collected, as indicated by collections made over an altitudinal gradient
(Barclay and Crawford 1984). As early as 1931, Flemion
reported that dry storage at room temperature shortened the
time needed for cold stratification and that germination tem-
1062
Table 5Sorbus, mountain-ash: stratification and germination test conditions and results*
Species
S. americana
S. aucuparia
S. decora
S. scopulina
S. sitchensis
Stratification
Temp
Moist
Days
(C)
medium
60
90
CSG
CSG
CSG
CSG
CSG
120
CSG
30180
CSG
70
126
115
90
120
140
5
5
5
1
1
1
1
2
3
20
3
20
2
35
3
0
5
Sand
Sand
Peat moss
Peat moss
Peat moss
Peat moss
Paper
Sand
Sand
Peat & sand
Peat & sand
Peat
Peatverm
Paper
Sand
Germination test
Temp
(C)
Days
2030
10
5
1
1
1
1
20
3
3
3
3
20
2030
20
2030
21
14
60
150
127
82
120
120
150210
150210
560
350
42
60
38
53
50
17
Germination
(%)
Days
13
11
15
70
69
58
57
68
60
86
61
30
15
8
22
132
99
61
90
90
90
420
300
17
10
11
S
Total
germination
(%) Samples
13
12
16
94
96
93
90
76
95+
90+
88
93
30
10
61
30
21
15
4
4
4
1
1
6
4
1
26
52
1
1
3
4
1
1
1
1
Sources: Barclay and Crawford (1984), Flemion (1931), Harris and Stein (1974), Hilton and others (1965), Lenartowicz (1988), McKeever (1938), Mirov and Kraebel
(1939),Trindle (1996), Zentsch (1970).
CSG = stratification and germination as a continuum under the same conditions; verm = vermiculite.
* Only the better test results for each species are listed.
Stratification in moist sand, peat moss, soil, or petri dish.
Reached 33% germination in 330 days (USDA FS 1948).
Ten-minute soak in H2SO4 before stratification.
lowing spring for production of seedlings in the same growing season. Fall-sowing involves the following considerations: (1) sowing should perhaps be done as early as midsummer, because some seedlots or species benefit from
moist warm conditioning prior to the moist chilling that is
supplied by winter weather (Heit 1968); (2) the seeds are
subject to predation by rodents, insects, and birds for a long
time and may need protection; and (3) stored seeds must be
available because time for after-ripening may not be sufficient for freshly collected seeds. Sowing outdoors in late
winter or spring requires use of cold-stratified seeds or sowing early enough that natural cold stratification can still
occur. Cold stratification at or near 1 C for 60 to 120 days
is needed for best results (Barclay and Crawford 1984;
Flemion 1931; Hilton and others 1965; Taylor and Gerrie
1987). If seeds are sown late or are not sufficiently preconditioned, or conditions are too warm, germination will be
delayed until the second or even third year (Fabricius 1931;
Flemion 1931; Harris and Stein 1974). Mountain-ash
seedlings can also be container-grown in greenhouses where
good germination and growth conditions are readily maintained (McDonald 1989).
1063
Mountain-ash seedlings are hardy and not very susceptible to insects or disease; however, they are subject to nipping by deer and moose (Fabricius 1931; Van Dersal 1938).
For field planting, 1+1 transplants are best, but 2+0
seedlings are also suitable (Harris and Stein 1974).
References
AOSA [Association of Official Seed Analysts]. 2000. Tetrazolium testing
handbook. Lincoln, NE: Association of Official Seed Analysts. 21+ p.
Barclay AM, Crawford RMM. 1984. Seedling emergence in the rowan
(Sorbus aucuparia) from an altitudinal gradient. Journal of Ecology 72(2):
627636.
Calder JA,Taylor RL. 1968. Flora of the Queen Charlotte Islands: 1.
Systematics of the vascular plants. Monogr. 4, Pt. 1. Ottawa: Canada
Department of Agriculture, Research Branch. 659 p.
Chalupa V. 1992. Micropropagation of European mountain ash (Sorbus aucuparia L.) and wild service tree (Sorbus torminalis (L.) Cr.). In: Bajaj YPS, ed.
High-tech and micropropagation II.Volume 18, Biotechnology in
Agriculture and Forestry. Berlin: Springer-Verlag: 211226.
Englund R. 1993. Fruit removal in Viburnum species: copious seed predation
and sporadic massive seed dispersal in a temperate shrub. Oikos 67(3):
503510.
Fabricius L. 1931. Die Samenkeimung von Sorbus aucuparia L.
Forstwissenschaftliches Zentralblatt 53: 413418.
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1632 p.
Flemion F. 1931. After-ripening, germination, and vitality of seeds of Sorbus
aucuparia L. Contributions from the Boyce Thompson Institute 3:
413439.
Flemion F. 1938. A rapid method for determining the viability of dormant
seeds. Contributions from Boyce Thompson Institute 9: 339351.
Granstrom A. 1987. Seed viability of fourteen species during five years of
storage in a forest soil. Journal of Ecology 75(2): 321331.
Harris AS, Stein WI. 1974. Sorbus L., mountain-ash. In: Schopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 780784.
Heit CE. 1955. The excised embryo method for testing germination quality
of dormant seed. Proceedings of the Association of Official Seed Analysts
45: 108117.
Heit CE. 1967a. Propagation from seed: 8. Fall planting of fruit and hardwood seeds. American Nurseryman 126(4): 1213, 8590.
Heit CE. 1967b. Propagation from seed: 11. Storage of deciduous tree and
shrub seeds. American Nurseryman 126(10): 1213, 8694.
Heit CE. 1968. Propagation from seed: 15. Fall planting of shrub seeds for
successful seedling production. American Nurseryman 128(4): 810,
7080.
Hilton RJ, Jaswal AS,Teskey BJE, Barabas B. 1965. Rest period studies on
seeds of Amelanchier, Prunus, and Sorbus. Canadian Journal of Plant
Science 45: 7985.
Hitchcock CL, Cronquist A, Ownbey M,Thompson JW. 1961. Vascular
plants of the Pacific Northwest. Part 3. Seattle: University of Washington
Press. 614 p.
ISTA [International Seed Testing Association]. 1996. International Rules for
Seed Testing: rules 1996. Seed Science and Technology 24 (Suppl.):
1335.
King JE. 1947. The effect of various treatments to induce germination of
seeds of some plants valuable for soil conservation and wildlife [MS
thesis]. Moscow, ID: University of Idaho. 97 p.
Kronenberg HG. 1994. Temperature influences on the flowering dates of
Syringa vulgaris L. and Sorbus aucuparia L. Scientia Horticulturae 57:
5971.
Lenartowicz A. 1988. Warm-followed-by-cold stratification of mountain-ash
(Sorbus aucuparia L.) seeds. Acta Horticulturae 226: 231238.
Little EL Jr. 1979. Checklist of United States trees, native and naturalized.
Agric. Handbk 541. Washington, DC: USDA Forest Service. 375 p.
MacDonald B. 1989. Introducing new and recommended plantsthe
British Columbia way. International Plant Propagators Society
Combined Proceedings 38: 413416.
McAllister H. 1985. The Aucuparia section of Sorbus. Plantsman 6(4):
248255.
McKeever DG. 1938. The effect of various methods of treatment on the
germination of seeds of some plants valuable for game and erosion
purposes [MS thesis]. Moscow, ID: University of Idaho. 132 p.
Miller HW, Ball CC, Knott NP. 1948. The comparative value of woody
plants as food for upland game birds. Biol. Bull. 8. Olympia: Washington
State Game Department. 39 p.
Mirov NT, Kraebel CJ. 1939. Collecting and handling seeds of wild plants.
For. Pub. 5. Washington, DC: USDA Civilian Conservation Corps. 42 p.
NBV [Nederlandsche Boschbouw Vereeniging]. 1946. Boomzaden: handleiding inzake het oogsten, behandelen, bewaren en uitzaaien van
boomzaden. Wageningen,The Netherlands: Ponsen and Looijen. 171 p.
Nikolaeva MG. 1967. Fiziologiya glubokogo pokoya semyan. Akademiya
Nauk SSSR., Botanicheskii Institut IM.V. L. Komarova. Leningrad:
Izdatelstvo Nauka [Physiology of deep dormancy in seeds [Translation
TT 68-50463. 1969. Springfield,VA: USDC CFSTI. 220 p.].
Pojar J, MacKinnon A, eds. 1994. Plants of the Pacific Northwest Coast.
Redmond, WA: Lone Pine Publishing. 527 p.
Rehder A. 1940. Manual of cultivated trees and shrubs hardy in North
America. 2nd ed. New York: Macmillan. 996 p.
Rohwer SA. 1913. Technical papers on miscellaneous forest insects: 6.
Chalcidids injurious to forest-tree seeds.Tech. Ser. 20, Part VI.
Washington, DC: USDA Bureau of Entomology: 157163.
Rosendahl CO. 1955. Trees and shrubs of the upper Midwest. Minneapolis:
University of Minnesota Press. 411 p.
Shoemaker JS, Hargrave PD. 1936. Propagating trees and shrubs from seed.
Circ. 21. Edmonton: University of Alberta, College of Agriculture. 22 p.
Stein WI, Slabaugh PE, Plummer AP. 1974. Harvesting, processing, and storage of fruits and seeds. In: Schopmeyer CS, tech. coord. Seeds of woody
plants in the United States. Agric. Handbk. 450. Washington, DC: USDA
Forest Service: 98125.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA Soil
Conservation Service. 198 p.
Taylor CW, Gerrie WA. 1987. Effects of temperature on seed germination
and seed dormancy in Sorbus glabrescens Cardot. Acta Horticulturae
215: 185192.
Trindle JDC. 1996. Personal communication. Corvallis, OR: USDA Natural
Resources Conservation Service, Plant Materials Center.
USDA FS [USDA Forest Service]. 1948. Woody-plant seed manual. Misc.
Pub. 654. Washington, DC: USDA Forest Service. 416 p.
Van Dersal WR. 1938. Native woody plants of the United States, their erosion-control and wildlife values. Misc. Pub. 303. Washington, DC: USDA.
362 p.
Viereck LA, Little EL Jr. 1972. Alaska trees and shrubs. Agric. Handbk. 410.
Washington, DC: USDA Forest Service. 265 p.
Wright D. 1981. Sorbus: a gardeners evaluation. Plantsman 3(2): 6598.
Zentsch W. 1970. Stratification of Sorbus aucuparia L. seeds. In: Bialobok S,
Suszka B, eds. Proceedings, International Symposium on Seed Physiology
of Woody Plants; 1968 September 38; Kornik, Poland. Kornik, Poland:
Institute of Dendrology and Kornik Arboretum: 127132.
BignoniaceaeTrumpet-creeper family
Spathodea
1065
References
Eggeling WJ. 1947. Working plan for the Budongo and Siba forests.
Entebbe: Uganda Protectorate. 66 p.
Francis JK. 1990. African tulip tree (Spathodea campanulata Beauv.). Res.
Note SO-ITF-SM-32. New Orleans: USDA Forest Service, Southern
Forest Experiment Station. 5 p.
Francis JK, Rodrguez A. 1993. Seeds of Puerto Rican trees and shrubs: second installment. Res. Note SO-374. New Orleans: USDA Forest Service,
Southern Forest Experiment Station. 5 p.
Holridge LR. 1942. Trees of Puerto Rico. Occ. Pap. 2. Ro Piedras, PR:
USDA Forest Service,Tropical Forest Experiment Station. 105 p.
Irvine FR. 1961. Woody plants of Ghana. London: Oxford University Press.
868 p.
Liogier HA, Martorell LF. 1982. Flora of Puerto Rico and adjacent islands: a
systematic synopsis. Ro Piedras, PR: Editorial de la Universidad de
Puerto Rico. 342 p.
Little EL Jr, Wadsworth FH. 1964. Common trees of Puerto Rico and the
Virgin Islands. Agric. Handbk. 249. Washington DC: USDA Forest
Service. 548 p.
Mahecha Vega GE, Echeverri Restrepo R. 1983. Aborales del Valle del
Cauca. Bogota: Litografia Arco. 208 p.
McConnell J, Muniappan R. 1991. Introduced ornamental plants that have
become weeds on Guam. Micronesia 3 (Suppl.): 4749.
RosaceaeRose family
Spiraea L.
spirea
John C. Zasada and Peter F. Stickney
Dr. Zasada retired from the USDA Forest Services North Central Research Station; Dr. Stickney retired
from the USDA Forest Services Rocky Mountain Forest and Range Experiment Station
Common name(s)
Occurrence
S. alba Du Roi
meadowsweet
meadowsweet
birchleaf spirea
birchleaf spirea
Douglas spirea
Beauverd spirea,
Alaska spirea
hardhack, steeplebush
S. virginiana Britt.
Virginia spirea,
Appalachian spirea
Sources: Curtis (1959), Habeck (1991), MacKinnon and others (1992), Ogle (1991a&b),Viereck and Little (1972).
Spiraea
1067
Germination. Seeds germinate readily with no pretreatment, particularly if sown before there has been any significant drying (Dirr and Haeuser 1987). Birchleaf spirea
seeds germinate at 0 to 2 C when kept under such conditions for more than 120 days (McLean 1967). This suggests
that seeds sown in the fall and overwintering under the snow
will germinate at about the time of snowmelt to take best
advantage of conditions favorable for seedling development.
Unstratified seeds of Beauverd spirea germinated only at
25 C. Germination of stratified seeds (30 days at 2 C) was
greater than 95% between 10 to 25 C and 40% at 5 C.
Neither stratified nor unstratified seeds germinated to any
degree in the dark (Calmes and Zasada 1982). Filled seeds
made up 68 and 85% of seedlots of birchleaf and Beauverd
spireas, respectively (Calmes and Zasada 1982; McLean
1967).
Nursery practice and natural regeneration. Natural
regeneration following disturbance appears to be mostly by
basal sprouting or from rhizomes. Only very severe fires or
soil disturbances can eliminate vegetative reproduction
(Calmes and Zasada 1982; Morgan and Neuenschwander
1988; Ogle 1991; Stickney 1986, 1989).
Seed regeneration of birchleaf spirea occurs 2 to 3 years
after fire, when seeds are abundant following the mass flowering in the first post-fire growing season (Stickney 1989).
This appears to be the main window for seed regeneration,
as seed availability and seedbed conditions are best at this
time (Stickney 1986, 1989, 1990). However, recent germinants and 1- to 2-year-old seedlings are not common (Miller
1996; Morgan and Neuenschwander 1988; Stickney 1990).
Plants can be produced from seeds or by vegetative
propagation. Seeds should be sown immediately after collection for the most rapid germination. Stored seeds may
require some stratification for best germination, but unstratified seeds germinate well. Softwood or hardwood cuttings
of horticultural varieties can be rooted and grow fairly rapidly, filling a 3.8-liter (1-gal) container in a single growing
season. Softwood cuttings appear to be used most commonly (Dirr and Heuser 1985). Shoot explants and micropropagation can be used to increase desirable clones; performance
and vigor of plants produced in this way varies with season
of the year and the number of times vegetative material is
subcultured (Norton and Norton 1988 a&b).
Spiraea
1069
References
Batta J. 1977. Studies of the genus Spiraea. Rep. 62. s, Norway: Agricultural
University of Norway, Department of Dendrology and Nursery
Management. 12 p.
Calmes MA, Zasada JC. 1982. Some reproductive traits of four shrub
species in the black spruce forest type of Alaska. Canadian Field
Naturalist 96(1): 3540.
Corns IGW, Annas RM. 1986. Field guide to forest ecosystems of west-central Alberta. Edmonton, AB: Canadian Forest Service, Northern Forestry
Centre.
Curtis JT. 1959. The vegetation of Wisconsin: an ordination of plant communities. Madison: University of Wisconsin Press. 657 p.
Dirr MA. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation, and uses. Champaign, IL:
Stipes Publishing. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Drew LA. 1967. Comparative phenology of seral shrub communities in the
cedar/hemlock zone [unpublished MS thesis]. Moscow, ID: University of
Idaho, College of Forestry. 108 p.
Esser LL. 1995. Spiraea douglasii. In: Fischer WC, comp.The Fire Effects
Information System [data base]. Missoula, MT: USDA Forest Service,
Intermountain Research Station, Fire Science Laboratory.
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1632 p.
Fowler WB,Tiedemann AR. 1980. Phenological relationships of Spiraea
betulifolia Pall. and Apocynum androsaemilfolium L. Northwest Science
54(1): 1725.
Goi M, Kawanishi T, Ihara Y. 1975. Studies on the acceleration of flowering in
woody ornamentals by low temperature treatments: 5.The flowering
behavior of Spiraea cantonensis [abstract]. Kagawa Daigaku Nogakubu
Gakujutsu Hokoku 26: 8493.
Goi M, Hasegawa A, Kashiwaga Y. 1974. Studies on the acceleration of flowering in woody ornamentals by low temperature treatments: 4.
Flowering behavior of Spiraea thunbergii. Kagawa Daigaku Nogakubu
Gakujutsu Hokoku 25: 3542
Graber RE,Thompson DF. 1978. Seeds in the organic layers and soil of four
beechbirchmaple stands. Res. Pap. NE-401. Upper Darby, PA: USDA
Forest Service, Northeastern Forest Experiment Station. 8 p.
Habeck RJ. 1991. Spiraea betulifolia. In: Fischer WC, comp.The Fire Effects
Information System [data base]. Missoula, MT: USDA Forest Service,
Intermountain Research Station, Intermountain Fire Science Laboratory.
StyracaceaeStorax family
Styrax L.
snowbell
W. Gary Johnson and Mark S. Roh
Mr. Johnson retired from the USDA Forest Services National Seed Laboratory
Dr. Roh is a horticulturist at the USDA Agricultural Research Services U.S. National Arboretum,
Floral and Nursery Plants Research Unit, Beltsville, Maryland
Growth habit, occurrence and use. The genus
Styraxthe snowbellscomprises about 100 species of
trees and shrubs in the warm temperate and tropical regions
of the Northern Hemisphere (LHBH 1976). The snowbells
in the United States are shrubs or small trees planted for
their showy flowers (table 1). Southeastern Asian species are
the source of the balsamic resin benzoin (LHBH 1976).
American snowbell grows to 5 m and 9 cm dbh, with 9cm-long pubescent leaves. Even though American snowbell
is common, it rarely grows large enough to be considered a
tree. It grows under 200 m of elevation in moist to wet
places, such as bottomland woods, flood plains, swamps,
and stream banks (Duncan and Duncan 1988; LHBH 1976).
Bigleaf snowbell reaches 8 m and 10 cm dbh, with 18cm-long gray pubescent leaves. It grows to 1,000 m elevation in deciduous or mixed woods, usually in well-drained
areas. Bigleaf snowbell rarely reaches tree size (Duncan and
Duncan 1988; LHBH 1976). The USDI Fish and Wildlife
Service has designated Texas snowbell as an endangered
species.
Several Asian snowbells are grown in the United States.
Japanese snowbell grows to 9 to 10 m with 8-cm-long
Common name(s)
Occurrence
S. americanus Lam.
S. americanus var. pulverulentur (Michx.) Perkins ex Rehd.
S. pulverulentus Michx.
S. grandifolius Ait.
S. japonicus Sieb. & Zucc.
American snowbell,
mock-orange*
bigleaf snowbell
Japanese snowbell,
Japanese snowdrop tree,
snowbell tree
fragrant snowbell,
bigleaf snowbell
Texas snowbell
Styrax 1071
/kg
11,200
8,000
2,950
/lb
Samples
5,090
3,630
1,340
1
2
2
Harvest date
Mean peak
germination* (%)
Mean no. of
weeks until
peak germ.
1999
23
14
27
10
25
8
22
5
26
June
July
July
Aug
Aug
Sept
Sept
Oct
Oct
0
0
0
7
67
80
83
84
80
19
2
16
30
13
4
19
July
Aug
Aug
Aug
Sept
Oct
Oct
0
0
5
16
65
73
88
d
c
b
ab
a
b
3.0 b
6.0 a
7.0 a
4.0 b
4.5 ab
3.0 b
e
d
c
b
a
2.3 c
7.0 a
8.0 a
4.7 b
6.0 ab
2000
Moist storage
at 18.5 C
Germination %
upon harvest to avoid any possible reduction in the germination percentage by dry storage. At least 1 month of warm
stratification is required, followed by 3 months of cold stratification to improve germination. Japanese snowbell seeds
harvested 12 to 14 weeks after flowering are mature and
respond to germination-promoting treatments. One month of
warm stratification at 18.5/18 C followed by 2 months of
cold stratification at 5.5 C resulted in germination higher
than 73%, while the maximum germination percentage was
98% after 2 months of warm stratification, followed by 3
months of cold stratification (figure 2).
1999
0
1
2
3
4
0
70.8 2.4 b
85.0 2.4 a
80.8 2.4 a
85.0 2.4 a
0
1
2
3
4
0
72.5 2.4 b
75.8 2.4 b
52.5 2.4 c
53.3 2.4 c
2000
Styrax
1073
Table 5Styrax japonicus, Japanese snowbell: effect of storage duration and moisture on seed germination, as evidenced by the number of weeks to reach peak germination
Months in
storage
0
1
2
3
4
Moisture
during storage
Germination
peak (%)
Dry
Moist
Dry
Moist
Dry
Moist
Dry
Moist
Dry
Moist
0
0
70.8 2.2 a
53.3 2.2 c
59.2 2.2 bc
65.0 2.2 ab
41.7 2.2 d
61.7 2.2 b
28.3 2.2 e
66.7 2.2 a
Weeks to peak
germination
6.3
6.3
7.0
5.0
6.7
5.3
5.3
5.7
0.45
0.45
0.45
0.45
0.45
0.45
0.45
0.45
References
Delaney J. 2002. Personal communication. Lecompte, LA: Louisiana Forest
Seed Co.
Dirr M, Heuser C Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Dirr M. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation and uses. Champaign, IL:
Stipes Publishing. 1007 p.
Duncan W, Duncan M. 1988. Trees of the southeastern United States.
Athens, GA: University of Georgia Press. 322 p.
Johnson W. 1997. Unpublished data. Dry Branch, GA: USDA Forest Service,
National Tree Seed Laboratory.
Kwon HJ. 1995. Ecological characteristics, propagation method, and new
cultivar of Korean Styrax japonicus Sieb. et. Zucc. for landscape tree uses
[PhD thesis]. Suwon, Korea: Sung Kyun Kwan University.
Kiew R. 1982. Germination and seedling survival in keminyan, Styrax
benzoin. Malaysian Forester 45(1): 6980.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan. 1290 p.
Ng FSP. 1976. The fruits, seeds and seedlings of Malayan trees: XIIXV.
Malaysian Forester 39(3): 110146.
Roh MS, Bentz JA. 2003. Germination of Styrax japonicus seeds as influenced by storage and sowing conditions. Acta Horticulturae 630:
411416
Roh MS, Bentz JA, Wang P, Li E, Koshioka M. 2003. Maturity and temperature stratification affects the germination of Styrax japonicus seeds
[unpublished data, submitted to Seed Technology].
Wasson E. 2001. Trees and shrubs: illustrated AZ of over 8500 plants.
Willoughby, NSW, Australia: Global Publishing: 725.
MeliaceaeMahogany family
Swietenia Jacq.
mahogany
John K. Francis
Dr. Francis retired from the USDA Forest Services International Institute of Tropical Forestry
Common name(s)
S. humilis Zucc.
S. macrophylla King
S. candollei Pittier
S. mahagoni (L.) Jacq.
S. macrophylla x mahagoni
Sources: Blake (1920), Francis (1991), Lamb (1966),Whitmore and Hinjosa (1977).
* Arose spontaneously from the introduced parent species.
Median fruit
dimensions (cm)
S. humilis
S. macrophylla
S. mahagoni
S. macrophylla x mahagoni
17 x 11
15 x 8
7x 4
12 x 7
Seeds/weight
Seeds/fruit
50 +
5070
3560
4565
/kg
/lb
1,500
1,4002,400
5,4007,800
1,9003,000
680
6401,100
2,5003,500
8601,400
Sources: Blake (1920), Francis and Rodrguez (1993), Marraro (1949), Montalvo and others (1991),Whitmore and Hinojosa (1977).
Swietenia
1075
germinating seedling.
References
Blake SF. 1920. Revision of mahoganies. Journal of the Washington
Academy of Science 10(10): 286297.
Burgos JA. 1954. Un estudio de la silvicultura de algunas especies forestales
de Tingo Maria, Peru. Caribbean Forester 15(1/ 2): 1453.
Campbell de Araujo U. 1971. Sobre a germinaco do mogno (aguano)
Swietenia macrophylla King. Acta Amazonica 1(3): 5969.
Chudnoff M. 1984. Tropical timbers of the world. Agric. Handbk. 607.
Washington, DC: USDA Forest Service. 464 p.
Francis JK. 1991. Swietenia mahagoni Jacq., West Indies mahogany. Res. Note
SO-ITF-SM-46. New Orleans: USDA Forest Service, Southern Forest
Experiment Station. 7 p.
Francis JK, Rodrguez A. 1993. Seeds of Puerto Rican trees and shrubs: second installment. Res. Note SO-374. New Orleans: USDA Forest Service,
Southern Forest Experiment Station. 5 p.
Jacalne DV, Heimban JR,Tadle JF. 1957. A study of the stump planting of
mahogany (Swietenia macrophylla King). Philippine Journal of Forestry
13(1/ 2): 6380.
Lamb FB. 1966. Mahogany of tropical America. Ann Arbor: University of
Michigan Press. 220 p.
Marrero J. 1943. A seed storage study of some tropical hardwoods.
Caribbean Forester 4(3): 99106.
Marrero J. 1949. Tree seed data from Puerto Rico. Caribbean Forester
10(1): 1136.
Mondala CA. 1977. Depth and position of sowing large-leaf mahogany
seeds. Sylvatrop 2(2): 131137.
Montalvo JM, Pena A, Castillo E, Sordo L. 1991. Caracteristics de la calidad
intrinsica de las semillas de Swietenia macrophylla. Revista Baracoa
21(2/3): 7584.
Ricardi M,Torres F, Hernandez C, Quintero R. 1977. Morfologa de plantulas de arboles Venezolanos: 1. Revista Forestal Venezolana 27: 1556.
Schmidt PB. 1974. Sobre a profundidade ideal de semeadura do mogno
(Aguano): Swietenia macrophylla King. Brasil Florestal 5(17): 4247.
Vivekanandan K. 1978. Retention of viability of mahogany seed through
cold storage. Sri Lanka Forester 13(3/4): 6768.
Whitmore JL, Hinjosa G. 1977. Mahogany (Swietenia) hybrids. Res. Paper
ITF-23. Ro Piedras, PR: USDA Forest Service, Institute of Tropical
Forestry. 8 p.
Zamoni-Mendiburu CA. 1975. Propagacin vegetativa por estacas de ocho
especies forestales [MS thesis].Turialba, Costa Rica: University of Costa
Rica. 100 p.
Swietenia
1077
CaprifoliaceaeHoneysuckle family
Symphoricarpos Duham.
snowberry
Scott C.Walker
Mr.Walker is the project leader at the Utah Division of Wildlife Resources
Great Basin Research Center, Ephraim, Utah
Common name(s)
Natural occurrence*
common snowberry
S. orbiculatus Moench
S. oreophilus var. oreophilus Gray
S. oreophilus var. utahensis
(Rydb.) A. Nels.
S. utahensis Rydb.
S. vaccinoides Rydb.
S. rotundifolia var. parishii
(Rydb.) Dempster
S. parishii Rydb.
garden snowberry,
Columbia snowberry
western snowberry,
buckbrush
coralberry, Indian currant,
snowberry
mountain snowberry,
Utah snowberry
Parish snowberry
be picked by hand. Those collected in early fall contain considerable moisture and therefore require careful handling to
prevent heating (Evans 1974; Vories 1981). Weight of fruits
per volume for western snowberry (fresh weight) are 3.6
kg/liter (58 lb/bu) and for coralberry (dry weight) are 0.81
kg/liter (13 lb/bu) (Evans 1974).
var. laevigatus
S. occidentalis
S. orbiculatus
S. oreophilus
Flowering
Fruit ripening
Michigan
Michigan
Idaho
700 m
Missoula Co., Montana
1,000 m
1,300 m
1,650 m
2,000 m
Pennington Co., South Dakota
750 m
June 1July 31
May 1Sept 30
Sept 1Oct 31
Aug 1Oct 31
June 5Aug 5
Aug 1 Sept 5
June 20Aug 15
July 1July 30
July 15Aug 30
July 25Sept 5
Aug 15Sept
Aug 25Sept
Sept 10Oct
Sept 25Oct
June 1July 31
July 1Aug 31
Sept 1Oct 31
Sept 1heavy frost
June 17June 26
June 22June 30
July 2July 8
Aug 20Sept 18
Aug 17Sept 12
Aug 21Sept 26
June 5June 10
July 15Aug 3
30
20
5
25
Sources: Billington (1943), Costello and others (1939), Evans (1974),Willard (1971).
Symphoricarpos
1079
Figure 3Symphoricarpos albus var. albus, common snowberry: seedling development at 5, 7, 13, and 20 days after
germination.
Figure 2 Symphoricarpos albus var. albus, common snowberry: longitudinal section through a nutlet.
Species
S. albus
var. albus*
var. laevigatus
S. occidentalis
S. orbiculatus
S. oreophilus
var. oreophilus
Seed wt/
fresh fruit wt
Range
/kg
Seeds (1,000s)/weight
Average
/lb
/kg
510
7
119250
86144
114217
298317
54113
3965.2
5298.7
135144
117165
53-75
167.5
122
164
308
141
/lb
76
55.4
74.4
140
53.9
Samples
10
5+
6+
2
1
S. orbiculatus
Immersion
in H2SO4 (min)
60
75
20
0
60
0
60
0
30
30
Stratification (days)
Warm*
Cold
60
20
0
112
84
91
0
120
20
120
180
180
60
182
168
182
140
120
120
180
Germination
(%)
35
74
1
45
69
87
32
72
58
81
Symphoricarpos
1081
References
Akagi ST. 1996. Personal communication. Salt Lake City: Utah Department
of Agriculture and Food.
Auger J. 1994. Viability and germination of seeds from seven fleshy-fruited
shrubs after passage through the American black bear (Ursus
americanus) [zoology MS thesis]. Provo, UT: Brigham Young University.
48 p.
Banister R. 1991. Snowberry. Rangelands 13(1): 3334.
Billington C. 1943. Shrubs of Michigan. Cranbrook Institute of Science
Bulletin 20: 1249.
Costello DF, Price R. 1939. Weather and plant-development data as
determinants of grazing periods on mountain range.Tech. Bull. 686.
Washington, DC: USDA. 31 p.
Cronquist A, Holmgren AH, Holmgren NH, Reveal JL, Holmgren PK. 1984.
Intermountain flora: vascular plants of the Intermountain West.Volume
4. Bronx, NY: New York Botanical Garden. 573 p.
Evans KE. 1974. Symphoricarpos, snowberry. In: Schopmeyer CS, tech. coord.
Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 787790.
Flemion F. 1934. Physiological and chemical changes preceding and during
the after ripening of Symphoricarpos racemosus seeds. Contributions of
the Boyce Thompson Institute 6: 91102.
Flemion F, Parker E. 1942. Germination studies of seeds of Symphoricarpos orbiculatus. Contributions of the Boyce Thompson Institute 12:
301307.
GBRC [Great Basin Research Center]. 1985. Unpublished data. Ephraim:
Utah Division of Wildlife Resources.
Grimm WC. 1957. The book of shrubs. Harrisburg, PA: Stackpole Books.
533 p.
Hidayati SN, Baskin JM, Baskin CC. 2001. Dormancy-breaking and germination requirements for seeds of Symphoricarpos orbiculatus
(Caprifoliaceae). American Journal of Botany 88(8): 14441445.
Landis TD, Simonich EJ. 1984. Producing native plants as container seedlings.
In: Murphy PM, comp.The challenge of producing native plants for the
Intermountain area. Proceedings, Intermountain Nurserymans
Association; 1983 August 811; Las Vegas, NV: 1625.
Link E. 1993. Native plant propagation techniques for national parks: interim guide. East Lansing, MI: USDA NRCS Rose Lake Plant Materials
Center. 197 p.
Monsen SB. 1984. Use of shrubs on mine spoils. In: Murphy PM, comp.The
challenge of producing native plants for the Intermountain area.
Proceedings, Intermountain Nurserymans Association; 1983 August
811; Las Vegas, NV: 2631.
Mozingo HN. 1987. Shrubs of the Great Basin: a natural history. Reno:
University of Nevada Press: 251254.
Plummer AP, Christiansen DR, Monsen SB. 1968. Restoring big game range
in Utah. Pub. 68-3. Salt Lake City: Utah Division of Fish and Game.183 p.
Shaw N. 1984. Producing bareroot seedlings of native shrubs. In: Murphy
PM, comp.The challenge of producing native plants for the
Intermountain area. Proceedings, Intermountain Nurserymans
Association; 1983 August 811; Las Vegas, NV: 615.
Shiell R. 1992. Symphoricarpos orbiculatus. American Nurseryman 176(10):
130.
SCS [Soil Conservation Service]. 1982. National list of scientific plant
names.Volume 1, List of plant names. SCS-TP-159. Washington, DC:
USDA Soil Conservation Service. 416 p.
Stevens R. 1994. Interseeding and transplanting to enhance species composition. In: Monsen SB, Kitchen SG, comps. 1994. Proceedings, Ecology and
Management of Annual Rangelands; 1992 May 1821; Boise, ID. Gen.
Tech. Rep INT-GTR-313. Ogden, UT: USDA Forest Service,
Intermountain Research Station. 416 p.
Stevens R, Jorgensen KR. 1994. Rangeland species: germination through 25
and up to 40 years of warehouse storage. In: Monsen SB, Kitchen SG,
comps. 1994. Proceedings: Ecology and Management of Annual
Rangelands; 1992 May 1821; Boise, ID. Gen.Tech. Rep INT-313. Ogden,
UT: USDA Forest Service, Intermountain Research Station. 416 p.
Thames JL, ed. 1977. Reclamation and use of disturbed land in the
Southwest.Tucson: University of Arizona Press. 682 p.
Vories KC. 1981. Growing Colorado plants from seed: a state of the art.
Volume 1, Shrubs. Gen.Tech. Rep INT-103. Ogden, UT: USDA Forest
Service, Intermountain Research Station.
Wasser CH. 1982. Ecology and culture of selected species useful in revegetating disturbed lands in the West. FWS/OBS-82/56. Washington, DC:
USDI Fish and Wildlife Service. 347 p.
Weber GP, Wiesner LE. 1980. Tetrazolium testing procedures for native
shrubs and forbs. Journal of Seed Technology 5(2): 2334.
Welsh SL, Atwood ND, Higgins LC, Goodrich S. 1987. A Utah flora. Great
Basin Naturalist Memoirs 9: 1894.
Willard EE. 1971. Some factors involved in activation of sprouting in little
rabbitbrush and snowberry on summer range [dissertation]. Logan: Utah
State University. 116 p.
OleaceaeOlive family
Syringa L.
lilac
Paul O. Rudolf, Paul E. Slabaugh, and Nancy L. Shaw
Dr. Rudolf (deceased) retired from the USDA Forest Services North Central Forest Experiment Station;
Dr. Slabaugh retired from the USDA Forest Services Rocky Mountain Forest and Range Experiment Station;
Dr. Shaw is a research botanist at the USDA Forest Services Rocky Mountain Research Station,
Forestry Sciences Laboratory, Boise, Idaho
Growth habit, occurrence, and use. The lilac genus
comprises about 30 species of deciduous shrubs or small
trees with opposite, usually undivided leaves. The genus
nameSyringais derived from the Greek word syrinx, a
pipe, and refers to the hollow shoots. Lilacs are native to
temperate Asia and southeastern Europe (Everett 1982) and
were probably introduced to America before 1700 (Heriteau
1990; Wyman 1986). They are grown primarily as ornamentals because of their large, showy, and often fragrant inflorescences (Rehder 1940). Lilacs are generally hardy and
long lived (Everett 1982). At least 3 species are used in shelterbelts and windbreaks. Four species or varieties grown for
conservation purposes in the United States are discussed in
this chapter (table 1); their heights at maturity and years of
first cultivation are also listed (Hoag 1965; Rehder 1940).
Hybrids and cultivars. Numerous lilac hybrids and
cultivars have been developed for horticultural use. These
selections exhibit variation in such characteristics as flower
color, period of flowering, and growth habit. Krssmann
(1986) reported that more than 900 cultivars are grown,
including more than 800 developed from common lilac (S.
vulgaris L.). The largest collections and numbers of varieties
Common name(s)
Occurrence
S. persica L.
S. reticulata ssp. amurensis (Rupr.)
P.S. Greene & M.C. Chang
Persian lilac
Amur lilac,
Manchurian lilac
Iran to NW China
S. amurensis Rupr.
S.reticulata var. mandschuria (Maxim.) Hara
S. villosa Vahl.
late lilac,
S. bretschneideri Lemoine
villous lilac
S. vulgaris L.
common lilac
Height at
maturity (m)
Year first
cultivated
1.53.0
1614
N China to Himalayas
3.03.9
1882
SE Europe
3.07.0
1563
SE Siberia in Amur
River region
Syringa
1083
Location
Flowering
Fruit ripening
S. persica
S. reticulata var. amurensis
NE US, Kansas
North Dakota
Manitoba
NE US & Europe
Kansas
W US
MayJune
Early June
JuneJuly
AprJune
Late Marearly May
Late Marmid-May*
Late MarApr
SeptOct
AugOct
S. vulgaris
Sources: Caprio and Snyder (1989), Cummings (1963), Hoag (1965), Hulbert (1963), LHBH (1976), NBV (1946), Rehder (1940),Walker (1968).
*First flowering.
fruits (capsules).
% germination
Average
Range
72
77
61
44
6480
7084
3385
Tests
5
2
13
61
Sources: Heit (1968a&b, 1974), Junttila (1974b), Rafn and Son (nd), Rudolf and
Syringa
1085
References
Caprio JM, Snyder RD. 1989. Within-station analysis of hourly thermal and
solar thermal phenological models for flowering of Syringa vulgaris L. In:
Driscoll D, Box EO, eds. Proceedings, 11th International Society of
Biometeorology Congress.The Hague: SPB Academic Publishing: 5365.
Cram WH, Nagy MH, Lindquist CH. 1960. Propagation research. Forest
Nursery Station Summary Report. Indian Head, SK: Canadian
Department of Agriculture, Research Branch: 1618.
Cumming WA. 1963. Late flowering lilacs for the prairies. Prairie Gardening
20: 95.
Everett TH, ed. 1982. The New York Botanical Garden illustrated encyclopedia of horticulture.Volume 10, STEZY. New York: Garland Publishing:
32253601.
Fordham AJ. 1959. Propagation and care of lilacs. Arnoldia 19: 3644.
Griffiths M. 1994. Index of garden plants. Portland, OR:Timber Press.
1234 p.
Hartmann HT, Kester DE, Davies FT Jr. 1990. Plant propagation principles
and practices. 5th ed. Englewood Cliffs, NJ: Prentice-Hall. 647 p.
Heit CE. 1967. Propagation from seed: 11. Storage of deciduous tree and
shrub seeds. American Nurseryman 126: 1213, 8694.
Heit CE. 1968a. Propagation from seed: 15. Fall planting of shrub seeds for
successful seedling production. American Nurseryman 128: 810.
Heit CE. 1968b. Thirty-five years testing of tree and shrub seed. Journal of
Forestry 66: 632634.
Heit CE. 1974. Laboratory germination and testing method for Syringa
villosa (late lilac) seed. Newsletter of the Association of Official Seed
Analysts 48: 2628.
Heriteau J. 1990. The National Arboretum book of outstanding garden
plants. New York: Simon and Schuster. 292 p.
Hoag DG. 1965. Trees and shrubs for the Northern Plains. Minneapolis:
Lund Press. 376 p.
Hulbert LC. 1963. Gates phenological records of 132 plants at Manhattan,
Kansas, 19261955.Transactions of the Kansas Academy of Science 66:
82106.
ISTA [International Seed Testing Association]. 1966. International rules for
seed testing. Proceedings of the International Seed Testing Association
31: 1152.
Isely D, Everson LE, eds. 1965. Rules for testing seeds. Proceedings of the
Association of Official Seed Analysts 54: 1112.
Junttila O. 1970a. Effects of gibberellic acid and temperature on the germination of Syringa vulgaris L. and S. reflexa Schneid. Meldinger fra Norges
Landbrukshgskole 49(36): 18.
Junttila O. 1970b. Effects of stratification, growth substances and temperature on the germination of some Syringa species. Meldinger fra Norges
Landbrukshgskole 49(35): 158.
Junttila O. 1971. Effects of mother plant temperature on seed development
and germination in Syringa reflexa Schneid. Medlinger fra Norges
Landbrukshgskole 50(10): 116.
Junttila O. 1973a. The mechanism of low temperature dormancy in mature
seeds of Syringa species. Physiologia Plantarum 29: 256263.
Junttila O. 1973b. Seed and embryo germination in Syringa vulgaris and S.
reflexa as affected by temperature during seed development. Physiologia
Plantarum 29: 264268.
Junttila O. 1974a. Effects of low oxygen concentrations on the germination
of Syringa seeds. Zeitschrift fr Pflanzenphysiologie 74: 168171.
Junttila O. 1974b. Seed quality and seed production of woody ornamentals
in Scandinavia. Medlinger fra Norges Landbrukshgskole 53: 141.
Krssmann G. 1986. Manual of cultivated broad-leaved trees and shrubs.
Volume 3, PRUZ. Handbuch der Laubgehlze. Portland, OR:Timber
Press. 510 p.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan. 1290 p.
Macdonald B. 1993. Practical woody plant propagation for nursery
growers.Volume 1. Portland, OR:Timber Press. 669 p.
NBV [Nederlandsche Boschbouw Vereeniging]. 1946. Boomzaden:
Handleiding inzake het oogsten, behandelen, bewaren en witzaaien van
boomzaden. Wageningen,The Netherlands: Ponsen and Looijen. 171 p.
Rafn J and Son. [nd, circa 1928]. Skovfrkontorets Franalyser gennem 40
Aar, 18871927. Udfrt paa Statsfrkontrollen I Kbenhavn. 5 p.
Rehder A. 1940. Manual of cultivated trees and shrubs hardy in North
America. 2nd ed. New York: Macmillan. 996 p.
Rudolf PO, Slabaugh PE. 1974. Syringa L., lilac. In: Schopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 791793.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA Soil
Conservation Service. 198 p.
Walker J. 1968. Immature vs. mature seed. Western Canadian Society of
Horticulture Proceedings 24: 7072.
Wyman D. 1986. Wymans gardening encyclopedia. New York: Macmillan.
1221 p.
TamaricaceaeTamarix family
longitudinal
Tamarix
1087
References
Baum BR. 1967. Introduced and naturalized tamarisks in the United States
and Canada. Quarterly Journal of Horticulture Taxonomy 15(1): 1925.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation. Athens, GA: Varsity Press. 239 p.
Everitt BL. 1980. Ecology of saltcedar: a plea for research. Environmental
Geology 3(2): 7784.
Gary HL, Horton JS. 1965. Some sprouting characteristics of five-stamen
tamarisk. Res. Note RM-39. Fort Collins, CO: USDA Forest Service,
Rocky Mountain Forest and Range Experiment Station. 7 p.
Horton HL. 1964. Notes on the introduction of deciduous tamarisks. Res.
Note RM-16. Fort Collins, CO: USDA Forest Service, Rocky Mountain
Forest and Range Experiment Station. 7 p.
Horton JS, Campbell CJ. 1974. Management of phreatophyte and riparian
vegetation for maximum multiple use values. Res. Pap. RM-117. Fort
Collins, CO: USDA Forest Service, Rocky Mountain Forest and Range
Experiment Station. 23 p.
Horton JS, Mounts FC, Kraft JM. 1960. Seed germination and seedling
establishment of phreatophyte species. Stn. Pap. 48. Fort Collins, CO:
USDA Forest Service, Rocky Mountain Forest and Range Experiment
Station. 26 p.
Merkel DL, Hopkins HH. 1957. Life history of saltcedar.Transactions of the
Kansas Academy of Science 60(4): 360369.
Reynolds HG, Alexander RR. 1974. Tamarix pentandra Pall., five-stamen
tamarisk. In: Schopmeyer CS, tech. coord. Seeds of woody plants in the
United States. Agric. Handbk. 450. Washington, DC: USDA Forest
Service: 794795.
Robinson TW. 1965. Introduction, spread, and areal extent of salt-cedar
(Tamarix) in the western states. Prof. Pap. 491-A. Washington DC: USDI
Geological Survey. 11 p.
Turner RM. 1974. Quantitative and historical evidence of vegetation
changes along the upper Gila River, Arizona. Prof. Pap. 655-H. Washington
DC: USDI Geologic Survey: 120.
TaxodiaceaeRedwood family
Common name(s)
Occurrence
T. ascendens Brongn.
T. distichum var. imbricarium (Nutt.) Croom
T. distichum var. nutans (Ait.) Sweet
T. distichum (L.) Rich.
Taxodium
1089
1090
longitudinal
Two insect pests destroy significant amounts of baldcypress and pondcypress seedssouthern pine coneworm
(Dioryctria amatella (Hulst)) and baldcypress coneworm
(D. pygmaeella Ragonot). The baldcypress seed midge
(Taxodiomyia cupressi Schweinitz) forms small round galls
inside the cones of baldcypress (Hedlin and others 1980;
Merkel 1984). The seed midge apparently does little damage to seeds, but the galls are difficult to separate from the
seeds and become a quarantine problem for seed exporters.
Collection, extraction, and storage. Mature, dry
cones can be picked by hand from standing or felled trees
and spread in a thin layer for air-drying. The dried cones
should be broken apart by flailing or dry maceration. The
resin in the cones presents a major problem in separation
and cleaning because it causes seeds and cone fragments to
stick together. The resin also gums up mechanical macerators. One possible solution is to place the dried seeds and
cone fragments in a freezer to harden the resin, then run
them through a macerator again while the resin is still in a
solid state. Resin can be cleaned from equipment with
alcohol or other organic solvents.
The number of seeds per cone volume for baldcypress
averages about 58 kg/hl (45 lb/bu) of fresh cones. About 50
kg of seeds can be obtained from 100 kg (110 lb/220 lb) of
fresh cones, and there are 7,300 to 10,000 cones/hl (2,600
to 3,550 cones/bu) (Bonner 1974). For baldcypress, the
average number of cleaned seeds per weight determined
from 26 samples was 11,500/kg (5,200/lb) with a range of
5,600 to 18,500/kb (2,540 to 8,400/lb). One sample of
pondcypress from Florida contained about 9,000 seeds/kg
4,100 seeds/lb) (Bonner 1974). Baldcypress seeds keep well
in dry storage at 2 to 5 C for at least 3 years. Because they
appear to be orthodox in storage behavior, longer storage
under the same conditions will probably succeed.
Germination. Baldcypress seeds exhibit a moderate
amount of dormancy that can be overcome by cold stratification (table 2). For germination testing, moist stratification
for 30 days at 3 to 5 C is recommended, followed by 28
days of testing at alternating temperatures of 20 C for 16
hours (dark) and 30 C for 8 hours (light) (ISTA 1993).
Studies with collections from the Gulf Coast region suggested that dormancy in both species is regulated by the
seedcoat, and any treatment that softens or weakens the
coats will increase rate of germination. Soaking for 4 hours
in concentrated sulfuric acid was recommended as the easiest treatment (Murphy and Stanley 1975). An alternative
method for nursery use has been to soak the seeds in water
Table 2Taxodium, baldcypress: germination test conditions and results on stratified seedlots
Species
T. ascendens
T. distichum
Kimpak
Kimpak
30
30
20
20
Germinative
energy
Days
(%)
30
30
76
67
Germinitive
capacity
Days
(%)
8
17
93
74
Samples
4
7
References
Bonner FT. 1974. Taxodium distichum (L.) Rich., baldcypress. In: Schopmeyer
CS, tech. coord. Seeds of woody plants in the United States. Agric.
Handbk. 450. Washington, DC: USDA Forest Service: 796798.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Faulkner S,Toliver J. 1983. Genetic variation of cones, seed, and nurserygrown seedlings of baldcypress (Taxodium distichum (L.) Rich.) provenances. In: Proceedings, 17th Southern Forest Tree Improvement
Conference; 1983 June 79; Athens, GA. Spons. Pub. 39. Southern
Forest Tree Improvement Committee: 281288.
Hedlin AF,Yates HO III,Tovar DC, Ebel BH, Koerber T, Merkel EP. 1980.
Cone and seed insects of North American conifers; North American
Forestry Commission, Study Group on Forest Insects and Diseases.
Ottawa: Canadian Forestry Service. 122 p.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing: rules 1993. Seed Science and Technology 21 (Suppl.):
1259.
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Merkel EP. 1984. The baldcypress coneworm, Dioryctria pygmaeella
Ragonot (Lepidoptera: Pyralidae). In: Yates HO III, ed. Proceedings,
Cone and Seed Insects Working Party Conference; Working Party S207.01; 1983 July 31August 6; Athens, GA. Asheville, NC: USDA Forest
Service, Southeastern Forest Experiment Station: 8590.
Murphy JB, Stanley RG. 1975. Increased germination rates of baldcypress
and pondcypress seed following treatments affecting the seed coat.
Physiologia Plantarum 35: 135139.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
Wilhite LP,Toliver JR. 1990. Taxodium distichum (L.) Rich., baldcypress. In:
Burns RM, Honkala BH, tech. coords. Silvics of North America.Volume
1, Conifers. Agric. Handbk. 654. Washington, DC: USDA Forest Service:
563572.
Taxodium
1091
TaxaceaeYew family
Taxus L.
yew
Nan C.Vance and Paul O. Rudolf
Dr. Vance is a research plant physiologist at the USDA Forest Services Pacific Northwest Research Station,
Corvallis, Oregon; Dr. Rudolf (deceased) retired from USDA Forest Service, North Central Research Station
Growth habit. The yewsmembers of the genus
Taxus of the family Taxaceaeare non-resinous evergreen
gymnosperms that are widely distributed throughout the
moderate zone of the Northern Hemisphere (table 1). They
grow primarily in the understory of moist, forested habitats
in cool, temperate to subtropical climates (Price 1990). The
growth form may be a tree or a shrub. In the understory, the
yews sprawling branchiness and spreading crown enable it
to capture light gaps in the canopy. The tree may convert to
shrub form if the main shoot is injured or declines and is
replaced by lateral branches or new growth. Shrubbiness
may also be sustained by frequent browsing. Crowns of
these shrubby forms may attain as much as 24 m in diameter
(Bugala 1978). The main stem of the yew tree can become
quite stout in proportion to its height. Often the large diameter is attained by multiple stems that have fused over time.
English yew has reached great age (1,000+ years) and girth,
Common name
Occurrence
T. baccata L.
T. baccata ssp. eubaccata Pilger.
T. brevifolia Nutt.
T. baccata ssp. brevifolia Pilger.
T. canadensis Marsh.
T. baccata ssp. canadensis Pilger.
English yew,
common yew
Pacific yew
Canada yew,
eastern yew,
ground hemlock
Chinese yew,
Maire yew,
Yunnan yew
Japanese yew
Florida yew
Honduran yew,
Guatemalan yew,
Mexican yew
Himalayan yew
1092
Taxus
1093
1094
longitudinal
English yew seeds was maintained for up to 4 years by storing them in moist sand or acid peat at low temperatures
(Rudolf 1974).
Pregermination treatments. Yew seeds are slow to
germinate; natural germination usually does not take place
until the second spring after seedfall (Suszka 1978). Viable
seeds of Pacific yew have been found in soil seedbanks for
several years (Minore 1994). Although a variety of birds and
small mammals eat, digest, and disperse yew seeds
(Bartkowiak 1978), germination does not appear to be hastened by their passing through the alimentary canal of birds.
Yew seeds have a strong but variable dormancy that can be
broken by warm-plus-cold stratification (Suszka 1978). One
recommendation is to hold the seeds for 150 to 210 days at
16 to 18 C, then for 60 to 120 days at 2 to 5 C (Heit 1967,
1969). The ISTA rules specify prechilling yew seeds for 270
days at 3 to 5 C. Steinfeld (1993a) reported on 2 groups of
seeds collected in the fall in Oregon that were stratified during the fall and winter. One group was chilled for 1 month
and the other was kept at warm temperatures for 5 months
and then chilled for 2 months. The seeds were sown in bareroot beds covered with mulch the following spring.
Location
Flowering
Seed dispersal
T. baccata
T. brevifolia
T. canadensis
T. cuspidata
T. floridana
W Europe
Washington & Oregon
Minnesota & Wisconsin
Japan
NW Florida
MarMay
MarMay
Apr
AprJune
JanMar
AugOct
JulyOct
AugSept
SeptOct
AugOct
AugOct
JulyOct
AugSept
Oct
AugOct
Species
Place collected
T. baccata
Western Europe
NE US
Carson & Skamania Cos.,Washington
S Cascades, Oregon
Central Cascades, Oregon
Upper mid-West
Minnesota & Wisconsin
Japan
NE US
T. brevifolia
T. canadensis
T. cuspidata
/kg
13,90018,000
13,20015,000
32,40036,200
23,80025,900
26,33039,950
33,00062,400
35,70038,460
24,70043,000
14,84019,300
Cleaned seeds/weight
Range
Average
/lb
/kg
/lb
6,3008,200
6,0006,800
14,70016,500
10,80011,800
12,00018,200
15,00028,400
16,20017,500
11,20019,500
6,7008,800
17,000
14,100
33,100
24,950
31,077
46,300
37,000
31,300
16,300
Samples
7,700
6,400
15,000
11,300
14,100
21,000
16,800
14,200
7,400
14
3
2
10
4
4
7
3
Taxus
1095
Figure 4Taxus baccata, English yew: seedling development 1, 8, 12, 22, and 39 days after germination.
Table 4Taxus, yew: stratification periods, germination test conditions, and results
Germination test conditions
Stratification (days)
Temp (C)
Warm
Cold
Day
Night
Species
T. baccata
T. brevifolia
T. cuspidata
120
120
365
365
1096
16
1016
30
1016
10
1016
20
1016
Days
60
60
60
Germinative capacity
Avg
Range
(%)
(%)
67
47
55
68
4770
5099
Samples
12
1
3
1
References
Alaback P, Antos J, Goward T, Lertzman K, MacKinnon A, Pojar J, Pojar R, Reed
A,Turner N,Vitt D. 1994. Plants of the Pacific Northwest Coast.
Redmond, WA: Lone Pine Publishing. 527 p.
Allison TD. 1990. Pollen production and plant density affect pollination and
seed production in Taxus canadensis. Ecology 7(2): 516522.
Allison TD. 1991. Variation in sex expression in Canada yew (Taxus
canadensis). American Journal of Botany 78(4): 569578.
Allison TD. 1995. Personal communication. Bethesda, MD: National Science
Foundation.
Ambasta SP. 1986. The useful plants of India. New Delhi: Council of
Scientific and Industrial Research, Publications and Information
Directorate. 622 p.
Arno SF, Hammerly RP. 1977. Northwest trees. Seattle, WA: The
Mountaineers. 222 p.
Bolsinger CL, Jaramillo AE. 1990. Taxus brevifolia Nutt., Pacific yew In: Burns
RM, Honkala BH, tech. coord. Silvics of North America. Agric. Handbk.
654. Washington, DC: USDA Forest Service: 573579.
Bugala W. 1978. Systematics and variability. In: Bialobok S, ed.The yew: Taxus
baccata L. [Markiewicz H, trans.]. Nasze Lesne, Warsaw: National Center
for Scientific,Technical and Economic Information Service. 149 p.
Chadwick LC, Keen RA. 1976. A study of the genus Taxus. Res. Bull. 1086.
Wooster, OH: Ohio Agricultural Research and Development Center.
56 p.
Copes DL. 1977. Influence of rooting media on root structure and rooting
percentage of Douglas-fir cuttings. Silvae Genetica 26: 102106.
Crawford, R. 1983. Pacific yew community ecology in north-central Idaho
with implications to forest land management [PhD thesis]. Moscow, ID:
University of Idaho. 109 p.
Dallimore W, Jackson AB. 1967. A handbook of Coniferae and
Ginkgoaceae. 4th ed. 4 [rev. by SG Harrison]. New York: St. Martins
Press. 729 p.
DiFazio, SP, Wilson MV,Vance, NC. 1996. Variation in sex expression of
Taxus brevifolia in western Oregon. Canadian Journal of Botany 74:
19431946.
DiFazio SP,Vance NC, Wilson MV. 1997. Strobilius production and growth
of Pacific yew under a range of overstory conditions in western Oregon.
Canadian Journal of Forest Research 27: 986993.
DiFazio SP, Wilson MV,Vance NC. 1998. Factors limiting seed production in
Taxus brevifolia (Taxaceae) in western Oregon. American Journal of
Botany 85: 910918.
Doede DL, Carroll E, Westfall R, Miller R, Kitzmiller JH, Snader KM. 1993.
Geographic variation in allozymes and taxol, and propagation by rooted
cuttings in Pacific yew. In: Temple CR, ed. International Yew Resources
Conference Extended Abstracts; 1993 March 1213. Berkeley, CA:
University of California: 1011.
Edwards DGW. 1987. Methods and procedures for testing tree seeds in
Canada. For. Tech. Rep. 36. Ottawa: Canadian Forestry Service. 31 p.
Flores T, Wagner LJ, Flores HE. 1993. Embryo culture and taxane production
in Taxus spp. In Vitro Cellular and Developmental Biology Plant 29:
160165.
Harlow WM, Harrar ES. 1958. Textbook of dendrology. 4th ed. New York:
McGraw-Hill. 561 p.
Harmann HT, Kester DE, Davies FT, Jr. 1990. Plant propagation principles and
practices. 5th ed. Englewood Cliffs, NJ: Regents/Prentice-Hall. 647 p.
Hartzell H. 1991. The yew tree. Eugene, OR: Hulogosi Press. 319 p.
Heit CE. 1967. Propagation from seed, part 10: storage methods for conifer
seeds. American Nurseryman 126(8): 1415, 3854.
Heit CE. 1969. Propagation from seed: 18.Testing and growing seeds of
popular Taxus forms. American Nurseryman 179(2): 1011, 118128.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing: rules 1993. Seed Science and Technology 21 (Suppl.): 1259.
Krssmann G. 1983. Manual of cultivated conifers. 2nd rev. ed. Berlin:Verlag
Paul Parey. 361 p.
Lee S. 1973. Forest botany of China, supplement. November 30, 1973: 12.
Lewington A, Parker E. 1999. Ancient trees: trees that live for 1000 years.
London: Collins and Brown: 6676.
Little EL Jr. 1971. Atlas of United States trees.Volume 1, Conifers and
important hardwoods. Misc. Pub. 1146. Washington, DC: USDA Forest
Service. 9 p. + 200 maps.
Miller RW 1980. A brief survey of Taxus alkaloids and other taxane derivatives. Journal of Natural Products 43: 425437.
Minore D, Weatherly HG, Cartmill M. 1996. Seeds, seedlings, and growth of
Pacific yew (Taxus brevifolia). Northwest Science 70: 223229.
Nisley RG. 2002. Personal observation. Avon, CT: USDA Forest Service,
Northeastern Research Station.
Owens JN, Simpson S. 1986. Pollen from conifers native to British Columbia.
Canadian Journal of Forest Research 16: 955967.
Pilz D. 1996a. Propagation of Pacific yews from seed: Part 1. American
Conifer Society Bulletin 13(1): 1319.
Taxus.
1097
1098
VerbenaceaeVerbena family
Tectona grandis L. f.
teak
T. H. Schubert and John K. Francis
Dr. Schubert retired from the USDA Forest Services Washington Office Timber Management Staff;
Dr. Francis retired from the USDA Forest Services International Institute of Tropical Forestry
Growth habit, occurrence, and use. Native to
Southeast Asia in India, Myamar (Burma), Thailand, and
Indochina, teak is the only important species of the 3 in the
genus Tectona (Schubert 1974). It is a large deciduous tree
that reaches maximum heights of 30 to 40 m. It grows best
in warm, moist tropical climates with 1,250 to 3,000 mm of
mean annual precipitation and a marked dry season of 3 to 6
months (Webb and others 1984). Teak has probably been
cultivated for centuries in Asia and has been planted for timber production in India and Burma since at least 1840
(Troup 1921). In the Western Hemisphere, teak has been
planted since about 1900, beginning in the Caribbean region
(Marshall 1929; Moldenke 1935). Because it is a tropical
species, in the continental United States, it grows successfully only in southern Florida. Adaptability trials have been
successful in Hawaii (Whitesell and Walters 1976). About
130 ha of teak plantations have been established in Puerto
Rico and the U.S. Virgin Islands (Weaver 1993). Teak wood
is famous the world over for its strength, durability, dimensional stability, working qualities, and the fact that it does
not cause corrosion when in contact with metal (Kukachka
1970; Troup 1921). It is currently used for shipbuilding, fine
furniture, trim, decorative objects, veneer for decorative plywood, posts, poles, and fuel (Kukachka 1970; Webb and
others 1984).
Geographical races of teak have been distinguished by
differences in stem form and rate of growth (Champion
1933). These are not recognized botanically even as varieties, but it is most important when establishing plantations
to use seeds from a race that will grow well under local conditions (Beard 1943; Champion 1933; Laurie 1938). In
Trinidad, trees grown from seeds of Burmese origin have
been more satisfactory than those grown from seeds of
Indian origin (Beard 1943).
Flowering and fruiting. The small white, perfect
flowers of teak are borne on short pedicels, in large erect
terminal panicles, about 2 months after the dry season has
ended and the large obovate leaves have emerged. The dates
vary somewhat depending on the climatic regime, but flow-
Tectona
1099
longitudinal section
Germination tests. Cut tests of fruits on 56 collections from across the range of teak revealed a potential mean
viability of 71% and ranged from 40 to 96% (Danish/FAO
Forest Tree Seed Centre 1973). Laboratory germination tests
should be carried out in sand at a constant 30 C for 28 days.
Pretreatment to stimulate germination should be 6 repetitions of soaking the fruits in water, followed by 3 days of
drying (ISTA 1993). Germination in nursery beds in various
parts of the world has varied from 0 to 96% in periods varying from 10 days to 3 months. Seeds extracted from the
fruits and treated with fungicide gave a germination of 54%
in 12 days (Dabral 1976). Because it is difficult to extract
teak seeds from their fruits and untreated teak fruits give
protracted, often low and unpredictable germination, some
pre-treatment is usually applied to fruits. Various pretreatments to hasten or improve germination have been used.
Soaking the fruits in water for several days, or alternate wetting and drying as in laboratory testing, have proven effective (Schubert 1959; Troup 1921; White and Cameron nd).
In one test, clean fruits were pretreated by 5 cycles of alternate soaking in water for 24 hours and drying in the sun for
48 hours and then sown. Germination began 18 days after
sowing and continued to increase for 15 days, after which it
gradually decreased. Germination 68 days after sowing was
61% of the total number of fruits sown (Schubert 1974).
Weathering of the epicarp and mesocarp aids germination.
Seeds inoculated with Scytalidium sp. (a cellulolytic fungus
isolated from teak litter), 0 and kept moist for 21 days had
96% germination compared to 20% for uninoculated control
(Dadwal and Jamaluddin 1988). Increases in germination of
5 to 12% over controls (21% germination) were obtained
with treatments of IAA and GA alone and in combination at
various concentrations (Uanikrishnan and Rajeeve 1990). A
novel method reported from Thailand is to expose the fruits
to ants for 1 to 2 weeks: they attack and remove the felty
covering and thus speed up germination without loss of viability (Bryndum 1966). Soaking fruits from 11 Indian provenances in a nutrient solution resulted in a higher seedling
yield (34%) than control (18%), water soak (30%) or scarification (28%). It is felt that nutrient deficiencies in some of
the sources resulted in lower germination or early seedling
failure (Gupta and Pattanath 1975). A temperature of 30 C
appears to be optimal for germinating teak seeds (Dabral
1976). Some seeds that were stored for several months germinated better than fresh seeds (Champion and Brasnett
1958; Mahapol 1954; Troup 1921), probably because seeds
need a period of after-ripening (Coster 1933). Because they
tend to have a greater number of seeds per fruit, larger fruits
yield a significantly higher number of seedlings per fruit. It
is recommended that fruits smaller than 14 mm in diameter
be culled (Banik 1977). Seeds from dry regions frequently
References
Banik RL. 1977. Studies on grading of teak fruits: 1. Fruit size is a factor in
germination of teak seeds. Bano Biggyan Patrika 6(1): 17.
Beard JS. 1943. The importance of race in teak, Tectona grandis L. Caribbean
Forester 4(3): 135139.
Bryndum K. 1966. The germination of teak. Bulletin of the Natural History
Society of Siam (Thailand) 21: 7586.
Chable AC. 1969. Reforestation in the Republic of Honduras, Central
America. Ceiba 13(2): 156.
Champion HG. 1933. The importance of the origin of seed used in
forestry. Indian Forest Records 17(5): 176.
Champion HG, Brasnett NV. 1958. Choice of tree species. For. Dev.
Pap. 13. Rome: FAO. 307 p.
Dabral SL. 1976. Extraction of teak seeds from fruits, their storage and
germination. Indian Forester 102(10): 650658.
Dadwal VS, Jamaluddin. 1988. Role of fungi in weathering of teak fruits.
Indian Forester 114(6): 328330.
Danish/FAO Forest Tree Seed Centre. 1973. Provenance collections of
teak. For. Occ. Pap. 1973/2. Rome: FAO: 5461.
Gupta BN, Kumar A. 1976. Estimation of potential germination of teak
(Tectona grandis L.f.) fruits from twenty three Indian sources by cutting
test. Indian Forester 102(11): 808813.
Gupta BN, Pattanath PG. 1975. Factors affecting germination behavior of
teak seeds of eighteen Indian origins. Indian Forester 101(10): 584588.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing: rules 1993. Seed Science and Technology 21 (Suppl.):
1259.
Keiding H. 1985. Teak, Tectona grandis, Linn. f. Seed Leaflet 4. Humlebaek,
Denmark: DANIDA Forest Seed Centre. 21 p.
Kukachka BF. 1970. Properties of imported tropical woods. Res. Pap. FPL125. Madison, WI: USDA Forest Service, Forest Products Laboratory.
68 p.
Kushalappa KA. 1977. Teak plantations in Thailand. Indian Forester 103(5):
323328.
Laurie MV. 1938. Branching and seed origin in Coorg teak plantations.
Indian Forester 64(10): 596600.
Mahapol S. 1954. Teak in Thailand. R-16. Bangkok: Ministry of Agricultural
Research, Forest Department. 31 p.
Marshall RC. 1929. Growing teak in Trinidad.Tropical Woods 19: 13.
Moldenke HN. 1935. A monograph of the genus Tectona as it occurs in
America and in cultivation. Phytologia 1(4): 154164.
Parry MS. 1956. Tree planting practices in tropical Africa. For. Dev. Pap. 5.
Rome: FAO. 302 p.
Schubert TH. 1974. Tectona grandis, teak. In: Schopmeyer CS, tech. coord.
Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 803804.
Troup RS. 1921. Silviculture of Indian trees.Volume 2. Oxford, UK:
Clarendon Press: 337783.
Uanikrishnan K, Rajeeve KP. 1990. On germination of Indian teak (Tectona
grandis L.f.). Indian Forester 102(10): 650658.
Weaver PL. 1993. Tectona grandis L.f., teak. Res. Note SO-ITF-SM-64. New
Orleans: USDA Forest Service, Southern Forest Experiment Station.
18 p.
Webb DB, Wood PJ, Smith JP, Henman GS. 1984. A guide to species selection for tropical and sub-tropical plantations.Trop. For. Pap. 15. Oxford,
UK: Oxford University, Commonwealth Forestry Institute, Unit of
Tropical Silviculture. 256 p.
White KJ, Cameron AL. [no date] Silvicultural techniques in Papua and
New Guinea forest plantations. Bull. 1. Port Moresby,Territory of Papua
and New Guinea: Division of Silviculture, Department of Forests. 99 p.
Whitesell CD, Walters GA. 1976. Species adaptability trials for man-made
forests in Hawaii. Res. Pap. PSW-118. Berkeley, CA: USDA Forest
Service, Pacific Southwest Forest and Range Experiment Station. 30 p.
Tectona
1101
AsteraceaeAster family
Tetradymia DC.
horsebrush
Lee E. Eddleman
Dr. Eddleman retired from Oregon State Universitys Department of Rangeland Resources,
Corvallis, Oregon
Common name
Occurrence
striped horsebrush,
striped cottonthorn
longspine horsebrush
gray horsebrush,
spineless horsebrush,
common horsebrush
hairy horsebrush
threadleaf horsebrush
smooth horsebrush,
littleleaf horsebrush
Nuttall horsebrush
spiny horsebrush,
cottonthorn horsebrush,
catclaw horsebrush,
shortspine horsebrush,
thorny horsebrush
Mojave horsebrush
cotton horsebrush,
four-part horsebrush,
dune horsebrush
T. axillaris A. Nels.
T.a. var. axillaris
T.a. var. longispina (M. E. Jones) Strother
T. canescens DC.
T. inermis Nutt.; T. multicaulis A. Nels.
T. linearis Rydb.
T. comosa Gray
T. filifolia Greene
T. glabrata Torr. & Gray
T. nuttallii Torr. & Gray
T. spinosa Nutt.
x T. permixta Payson
T. spinosa Hook. & Arn.
Lagothamnus ambiguus Nutt.
L. microphyllus Nutt.
T. stenolepis Greene
T. tetrameres (Blake) Strother
T. comosa Gray ssp. tetrameres Blake
Sources: Cronquest (1994), McArthur and others (1979), Mozingo (1987), Strother (1974).
1102
Tetradymia
1103
References
Cronquist A, Holmgren AH, Holmgren NH, Reveal JL, Holmgren PK. 1994.
Intermountain flora.Volume 5, Asterales. Cronquest A, ed. Bronx, NY:
New York Botanical Garden. 496 p.
Eddleman LE. 1977. Indigenous plants of southeastern Montana. 1.Viability
and suitability for reclamation in the Fort Union Basin. Spec. Pub. 4.
Missoula: University of Montana Montana, Forest and Conservation
Experiment Station. 122 p.
Hartman H. 1984. Ecology of gall-forming lepidoptera on Tetradymia. 1. Gall
size and shape. Hilgardia 52: 116.
Johnson AE. 1974. Experimental photosensitization and toxicity in sheep
produced by Tetradymia glabrata. Canadian Journal of Comparative
Medicine 38: 408410.
Kingsbury JM. 1964. Poisonous plants of the United States and Canada.
Englewood Cliffs, NJ: Prentice-Hall. 626 p.
McArthur ED, Blauer AC, Plummer AP, Stevens R. 1979. Characteristics and
hybridization of important intermountain shrubs. 3. Sunflower family. Res.
Pap. INT-220. Ogden: USDA Forest Service, Intermountain Forest and
Range Experiment Station. 82 p.
1104
Mozingo HN. 1987. Shrubs of the Great Basin: a natural history. Reno:
University of Nevada Press. 342 p.
Stark N. 1966. Review of highway planting information appropriate to
Nevada. Bull. B-7. Reno: University of Nevada, College of Agriculture,
Desert Research Institute. 209 p.
Strother JL. 1974. Taxonomy of Tetradymia (Compositae: Senecioneae).
Brittonia 26: 177202.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA Soil
Conservation Service. 198 p.
Young JA,Young CG. 1992. Seeds of woody plants in North America.
Portland, OR: Dioscorides Press. 407 p.
MalvaceaeMallow family
Thespesia
1105
Germination. No pregermination treatments are necessary. Seeds of portiatree should be sown in sandy media
and lightly covered (Parrotta 1994). From 65 to 79% of
fresh seeds germinate, beginning in 8 days and continuing
over a 9-week period (Francis and Rodrguez 1993; Ricardi
and others 1977; Parrotta 1994). Maga seeds may be sown
and lightly covered in ordinary potting mix. Marrero (1942)
reported that, although 70 to 80% of fresh seeds germinated,
only 20% of seeds stored at room temperature for 2 weeks
germinated. Francis and Rodrguez (1993) reported 80%
germination beginning 6 days after sowing. Germination of
both species is epigeal (figure 3) (Francis 1989; Parrotta
1994).
Nursery practice. Ordinary nursery practice is to germinate seeds in germination trays or beds and transplant
seedlings into containers (pots or plastic nursery bags) after
the first true leaves emerge. Portiatree seedlings reach 15 cm
(6 in) in height about 3 months after sowing (Parrotta 1994).
Moving portiatree seedlings into full sunlight after they are
established in the pots is recommended. Rooted cuttings are
also used to produce portiatree stock. Maga seedlings develop rapidly in partial shade, reaching 20 cm (8 in) in 3
months and 40 cm (16 in) in 6 months (Francis 1989). Maga
seedlings should be moved into full sun a few weeks before
outplanting. Seedling stock of either species from 15 to 50
cm (6 to 20 in) can be used to establish plantations. Trees
destined to become ornamentals are often grown in pots
until they attain 1 to 2 m (39 to 79 in) in height. Wildlings
are sometimes collected, potted, and allowed to rebuild their
root system before outplanting.
seed cut in
seedcoat
cotyledons
hypocotyl
radicle
1106
germination and
References
Francis JK. 1989. Thespesia grandiflora (DC.) Urban, maga. Res. Note
SO-ITF-SM-21. New Orleans: USDA Forest Service, Southern Forest
Experiment Station. 4 p.
Francis JK, Rodrguez A. 1993. Seeds of Puerto Rican trees and shrubs: second installment. Res. Note SO-374. New Orleans: USDA Forest Service,
Southern Forest Experiment Station. 5 P.
Howard RA. 1989. Flora of the Lesser Antilles, Leeward and Windward
Islands.Volume 5. Jamaica Plain, MA: Harvard University, Arnold
Arboretum. 604 p.
Little EL Jr, Skolmen RG. 1989. Common forest trees of Hawaii (native and
introduced). Agric. Hdbk. 679. Washington, DC: USDA Forest Service:
194.
Little EL Jr, Wadsworth FH. 1964. Common trees of Puerto Rico and the
Virgin Islands. Agric. Handbk. 249. Washington, DC: USDA Forest
Service. 548 p.
Marrero J. 1942. A seed storage study of maga. Caribbean Forester 3(4):
173184.
Marrero J. 1949. Tree seed data from Puerto Rico. Caribbean Forester 10:
1130.
Neal MC. 1965. In gardens of Hawaii. Spec. Pub. 50. Honolulu: Bernice P.
Bishop Press. 924 p.
Parrotta JA. 1994. Thespesia populnea (L.) Soland. ex Correa: portiatree,
emajagilla. Res. Note SO-ITF-SM-76. New Orleans: USDA Forest
Service, Southern Forest Experiment Station. 5 p.
Rashid MA, ed. 1975. The silviculture of Indian trees. rev. ed. Volume 1.
Delhi: Government of India Press. 307 p.
Ricardi M,Torres F, Hernandez C. Quintero R. 1977. Morfologa de plantulas de rboles Venezolanos. Revista Forestal Venezolana 27: 1556.
Von Carlowitz PG. 1986. Multipurpose tree and shrub seed directory.
Nairobi: International Council for Research in Agroforestry. 265 p.
Thespesia
1107
CupressaceaeCypress family
Thuja L.
arborvitae
Gary J. Brand and C. S. Schopmeyer
Mr. Brand is a research forester at the USDA Forest Service, North Central Forest Experiment Station,
St. Paul, Minnesota; Dr. Schopmeyer (deceased) was technical coordinator for the previous version of this manual
Common name(s)
Occurrence
T. occidentalis L.
T. obtusa Moench
T. odorata Marshall
T. plicata Donn ex D. Don
T. plicata D. Don; T. plicata Donn
T. plicata Donn ex D. Don in Lamb.
T. gigantea Nutt.
T. menziesii Dougl. ex Endl.
T. lobbii Hort. ex Gord.
T. standishii (Gord.) Carr.
T. japonica Maxim.
Thujopsis standishii Gord.
T. koraiensis Nakai
T. kongonsis Nakai
T. sutchuensis Frachet
Japan
Korea
Sichuan thuja
China
Sources: Cope (1986), Kartesz (1994a&b), Little (1979), Rushforth (1987),Vidakovic (1991).
1108
Thuja
1109
1110
Thuja
1111
References
Bower RC, Dunsworth BG. 1988. Provenance test of western red cedar on
Vancouver Island. In: Smith NJ, ed. Western red cedar: does it have a
future? 1987 July 13;Vancouver, BC.Vancouver: University of British
Columbia: 131135.
Briand CH, Posluszny U, Larson DW. 1992. Comparative seed morphology
of Thuja occidentalis (eastern white-cedar) from upland and lowland sites.
Canadian Journal of Botany 70: 434438.
Colangeli AM, Owens JN. 1990. The relationship between time of pollination, pollination efficiency, and cone size in western red cedar (Thuja plicata). Canadian Journal of Botany 68: 439443.
Cope EA. 1986. Native and cultivated conifers of northeastern North
America: a guide. Ithaca, NY: Cornell University Press. 231 p.
Curran MP, Dunsworth BG. 1988. Coastal western red cedar regeneration:
problems and potential. In: Smith NJ, ed. Western red cedar: does it have
a future? 1987 July 13;Vancouver, BC.Vancouver: University of British
Columbia: 2032.
Curtis JD. 1946. Preliminary observations of northern white cedar in
Maine. Ecology 27: 2326.
Dirr MA. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation, and uses. 4th ed. Champaign,
IL: Stipes Publishing. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Edwards DGW. 1986. Cone production, collection, and processing. In:
Shearer RC, comp. Proceedings, Conifer Tree Seed in the Inland
Mountain West Symposium; 1985 August 5; Missoula, MT. Gen.Tech. Rep.
INT-203. Ogden, UT: USDA Forest Service, Intermountain Forest and
Range Experiment Station: 78102.
Edwards DGW, Leadem CL. 1988. The reproductive biology of western
red cedar with some observations on nursery production and prospects
for seed orchards. In: Smith NJ, ed. Western red cedar: does it have a
future? 1987 July 13;Vancouver, BC.Vancouver: University of British
Columbia: 102113.
Gordon AG, Gosling P, Wang BSP, ed. 1991. Tree and shrub seed handbook.
Zurich: ISTA. 14 chapters.
Harlow WM, Harrar ES, Hardin JW, White FM. 1991. Textbook of dendrology: covering the important forest trees of the United States and
Canada, 7th ed. New York: McGraw-Hill. 501 p.
Henchell C. 1994. Personal communication. Carson, WA: USDA Forest
Service.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing: rules 1993. Seed Science and Technology 21 (Suppl.):
1259.
Jeffers RM. 1976. Survival and height growth of northern white-cedar from
18 provenances. In: Beineke WF, ed. Proceedings, 10th Central States
Forest Tree Improvement Conference; 1976 September 22; West
Lafayette, IN. West Lafayette, IN: Purdue University, Department of
Forestry and Natural Resources: 152156.
Johnston WF. 1990. Thuja occidentalis. In: Burns RM, Honkala BH, tech.
coords). Silvics of North America.Volume 1, Conifers. Agric. Handbk.
654. Washington, DC: USDA Forest Service: 580589.
1112
TiliaceaeLinden family
Tilia L.
linden or basswood
D. Bradley Rowe and Frank A. Blazich
Dr. Rowe is associate professor at Michigan State Universitys Department of Horticulture, East Lansing,
Michigan; Dr. Blazich is alumni distinguished graduate professor of plant propagation and tissue culture at
North Carolina State Universitys Department of Horticultural Science, Raleigh, North Carolina
bark that becomes fissured on old trees (table 2). The winter
form is striking, with stiff, erect branches growing upward at
30 angles from a thick trunk (Burgess 1991). Considerable
differences in growth habit exist among cultivars of littleleaf
linden, ranging from the very dense, formal pyramidal habit
of Greenspire, the dense upright oval shape of
Chancellor, to the more open, informal oval habit of
Fairview (Pellett and others 1988).
There is much disagreement among taxonomists as to
correct identification of species, and there are numerous
names in the literature that are no longer recognized by
many botanists. For example, T. monticola Sarg. and T.
michauxii (Nutt.) Sarg. are sometimes seen in the literature
or listed as specimens in botanical gardens, but they are now
considered to be varieties of white basswoodT. americana
var. heterophylla (Venten.) Loud.recognized previously as
T. heterophylla Venten. (Ayers 1993; Rehder 1990).
Common name(s)
Occurrence
T. americana L.
T. glabra Venten.
T. americana var. caroliniana
(P. Mill.) Castigl.
T. americana var. heterophylla
(Venten.) Loud.
T. cordata P. Mill.
T. parviflora J. F. Ehrh. ex Hoffm.)
T. euchlora K. Koch
T. cordata x T. dasystyla
T. europaea L.
T. cordata x T. platyphyllos
T. intermedia DC.
T. vulgaris Hayne
T. mexicana Schldl.
T. petiolaris DC.
T. platyphyllos Scop.
T. europaea var. grandiflora Hort.
T. tomentosa Moench
T. argentea DC.
white basswood
littleleaf linden, small-leaved
lime, European linden
Crimean linden, Caucasian lime
SE US
West Virginia to Florida,W to
Indiana & Alabama
Europe
SE Europe & SW Asia
Europe
Mexican basswood
pendent silver linden,
pendent white lime, weeping lime
bigleaf linden, large-leaved
lime, largeleaf linden
silver linden, European
white linden
Mexico
SE Europe & W Asia
Europe to SW Asia
SW Europe & Asia
Sources: Dirr (1990), LHBH (1976), Plotnik (2000), Rehder (1990), RHS (1994).
Tilia
1113
General comments
T. americana
T. a. var. caroliniana
T. a. var. heterophylla
T. cordata
T. euchlora
T. europaea
T. mexicana
T. petiolaris
T. platyphyllos
T. tomentosa
Sources: Dirr (1990), LHBH (1976), Plotnik (2000), Rehder (1990), RHS (1994).
seed.
longitudinal
Tilia
1115
Species
T. americana
T. cordata
T. x europaea
T. platyphyllos
T. tomentosa
34.1
36.3
1116
75
80
/kg
6.617.6
2032.1
24.938.3
48.865.1
23.329.7
25.130.6
20.025.1
3.08.0
9.114.6
11.317.4
22.229.6
10.613.5
11.413.9
9.111.4
6.6
11
30.4
3
5
13.8
Samples
2
15+
57+
good germination for all species and seedlots have not been
developed. Much variability exists among species and seedlots in regards to permeability of the pericarp and seedcoat,
as well as stratification requirements.
In Europe, dormancy in littleleaf linden is overcome by
the use of warm incubation or acid scarification, followed by
stratification (Suszka and others 1996). Fully imbibed seeds
are first stored for 4 months at 20 to 25 C (or scarified with
concentrated sulfuric acid for 12 minutes), then stratified at
3 C for 14 to 18 weeks. Stratification should be stopped
when the first seeds start to germinate.This complete process
may take 8 or 9 months.
Germination tests. Germination is epigeal (figure 4).
Optimum germination occurs at temperatures above 20 C
(68 F), but seeds will germinate at temperatures as low as
2 C once stratification requirements have been satisfied
(Spaeth 1934). Thus, seeds should be checked periodically
for radicle emergence during stratification. Light is not
required for germination (Heit 1967). The use of any stratification procedure requires far too much time to be used in
routine germination testing, however, so rapid estimates of
viability are recommended for this purpose. This can be
done with tetrazolium staining, indigo-carmine staining, or
excised embryo tests (ISTA 1996; Suszka and others 1996).
However, these tests require removal of the pericarp and the
seedcoat without damaging the embryo. Tetrazolium staining is the most common test. It requires soaking seeds in
water for 18 to 24 hours, removing all or a large part of the
seedcoat, and soaking the seeds in a 1% tetrazolium solution
for 24 to 48 hours at 30 C.
Pitel and Wang (1988) found that both the rate and percentage of germination of seeds of American linden were
increased by treating scarified seeds with a solution of
kinetin (1 mg/liter) and gibberellic acid (GA3, 500 mg/liter.
Over 90% germination was obtained after 60 to 80 days at
4 C. However, GA did not improve germination percentage
of seedlots of littleleaf, bigleaf, and silver linden (Magherini
and Nin 1993). The conflicting results are likely due to the
level of gibberellin present. Natural levels of GA exist in
dormant, nonstratified seeds and a sudden increase in the
quantity of gibberellin is observed from the sixth week of
stratification (Nagy 1980). It is likely that a specific quantity, rather than just the presence of free gibberellins, is
required to break dormancy and stimulate germination.
Traditionally, for an accurate germination test, the outer
pericarp must be removed and the hard seeds must undergo
scarification and stratification. However, excised embryos of
American linden that were separated from the seedcoat and
endosperm were able to germinate and grow when placed on
Tilia
1117
References
Ayers GS. 1993. Reconsidering the basswoods: 11.The native American
basswoods. American Bee Journal 133(5): 337340.
Bremness L. 1994. Herbs. London: Dorling Kindersley: 88.
Brinkman KA. 1974. Tilia L., basswood, linden. In: Schopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk 450.
Washington, DC: USDA Forest Service: 810812.
Burgess KS. 1991. Tilia tomentosa. Public Garden: Journal of the American
Association of Botanical Gardens and Arboreta 6(1): 39.
Chalupa V. 1990. Plant regeneration by somatic embryogenesis from cultured immature embryos of oak (Quercus robur L.) and linden (Tilia cordata Mill.). Plant Cell Reports 9(7): 398401.
Dirr MA. 1998. Manual of woody landscape plants: their identification,
ornamental characteristics, culture, propagation and uses. 5th ed.
Champaign, IL: Stipes Publishing Co. 1187 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Flemer W III. 1980. Linden propagation: a review. Combined Proceedings of
the International Plant Propagators Society 30: 333336.
Haller JM. 1995. Tilia americana, linden: a neglected jewel. Arbor Age 15(7):
3233.
Hartmann HT, Kester DE, Davies FT Jr, Geneve RL. 2002. Hartmann and
Kesters plant propagation: principles and practices. 7th ed. Upper Saddle
River, NJ: Prentice-Hall. 880 p.
Heit CE. 1967. Propagation from seed: 7. Successful propagation of six
hardseeded group species. American Nurseryman 125(12): 1012,
3741, 4445.
Heit CE. 1977. Propagation from seed: 27. Collecting, testing and growing
tilia linden species. American Nurseryman 146(7): 1011, 100110.
Howard BH. 1995. Opportunities for developing clonal rootstocks from
natural seedlings of Tilia spp. Journal of Horticultural Science 70(5):
775786.
ISTA [International Seed Testing Association]. 1996. International rules for
seed testing. Seed Science and Technology 24(suppl.): 335.
Kunneman BPAM, Albers MRJ. 1991. Linden trees (Tilia spp.). In: Bajaj YPS,
ed. Biotechnology in agriculture and forestry: 16.Trees III Berlin:
Springer-Verlag:152163.
1118
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan. 1290 p.
Magherini R., Nin S. 1993. Experiments on seed germination of some Tilia
spp. Acta Horticulturae 331: 251258.
Nagy M. 1980. Dormancy in fruits of Tilia platyphyllos Scop.: 4. Changes in
the endogenous gibberellin content during stratification. Acta
Agronomica 29: 111.
Nagy M., Keri A. 1984. Role of the embryo in the cytolysis of the
endosperm cells during the germination of the seeds of Tilia platyphyllos
Scop. Biochemie und Physiologie der Pflanzen 179: 145148.
Pellett H,Vogel K, McNamara S, Zuzek K. 1988. Relative growth rate and
plant habit of linden taxa. Journal of Environmental Horticulture 6(2):
4852.
Pigott CD, Huntley JP. 1981. Factors controlling the distribution of Tilia
cordara at the northern limits of its geographical range: 3. Nature and
causes of seed sterility. New Phytologist 87(4): 817839.
Pitel JA, Cheliak WM, Wang BSP. 1989. Some biochemical changes associated with stratification and germination of basswood seeds. Seed Science
and Technology 17(1): 5771.
Pitel JA, Wang BSP. 1988. Improving germination of basswood (Tilia americana L.) seeds with gibberellic acid. Seed Science and Technology 16(1):
273280.
Plotnik A. 2000. The urban tree book: an uncommon field guide for city
and town. New York:Three Rivers Press: 211215.
Rehder A. 1990. Manual of cultivated trees and shrubs hardy in North
America. 2nd ed. Portland, OR: Dioscorides Press. 996 p.
RHS [Royal Horticultural Society]. 1994. The new Royal Horticultural
Society dictionary index of garden plants. Griffiths M, ed. London:
Macmillan. 1234 p.
Spaeth JN. 1934. A physiological study of dormancy in Tilia seed. Cornell
University Agricultural Experiment Station Memoir 169: 171.
Suszka B, Muller C, Bonnet-Masimbert M. 1996. Seeds of forest
broadleaves, from harvest to sowing. Gordon A, trans. Paris: Institut
National de la Recherche Agronomique. 294 p.
Vanstone DE. 1978. Basswood (Tilia americana L.) seed germination.
Combined Proceedings of the International Plant Propagators Society
28: 566570.
Vanstone DE. 1982. Seed germination of American basswood in relation
to seed maturity. Canadian Journal of Plant Science 62(3): 709713.
MeliaceaeMahogany family
seed.
longitudinal sec-
Toona
1119
References
Carlson NK, Bryan, LW. 1959. Hawaiian timber for coming generations.
Honolulu: Trustees of the Bernice P. Bishop Estate. 112 p.
Chudnoff M. 1984. Tropical timbers of the world. Agric. Handbk. 607.
Washington, DC: USDA Forest Service. 464 p.
Francis WD. 1951. Australian rain-forest trees. Sydney: Commonwealth of
Australia, Forestry and Timber Bureau. 469 p.
Streets RJ. 1962. Exotic forest trees in the British Commonwealth. Oxford,
UK: Clarendon Press. 765 p.
Viga WM. 1976. Comportamento do cedro australiano, Toona ciliata M.
Roem. var. australis (F. & M.) D. DC., face a suceptibilidade do ataque de
Hypsipyla grandella (Zeller). Silvicultura em So Paulo 10: 109118.
1120
Walters GA. 1974. Toona australis, Australian toon. In: Schopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 813814.
Webb DB, Wood PJ, Smith JP, Henman GS. 1984. A guide to species selection for tropical and sub-tropical plantations.Trop. For. Pap. 15. Oxford,
UK: Oxford University, Commonwealth Forestry Institute, Unit of
Tropical Silviculture. 256 p.
Whitmore JL, Medina Gaud S. 1974. White peach scale attack on toon in
Puerto Rico. Journal of Agriculture of the University of Puerto Rico
58(2): 276278.
TaxaceaeYew family
Torreya Arn.
torreya
William I. Stein
Dr. Stein is a forest ecologist emeritus at the USDA Forest Services Pacific Northwest
Research Station, Corvallis, Oregon
Growth habit, occurrence, and uses. The genus
Torreya includes 7 species of conifer trees found in North
America and eastern Asia (Little 1979; Price 1990). These
species of limited distribution represent a genus that in earlier geologic times was widespread in the Northern
HemisphereEurope, Greenland, Alaska, British Columbia,
Oregon, Colorado, Virginia, and North Carolina (Abrams
1955; Boeshore and Gray 1936; Florin 1963; Schwartz and
Hermann 1993a). Two species are native in the United
States: Florida torreya is endemic to a small area in Florida
and Georgia, and California torreya to central California
(table 1). Little genetic variability has been found among
populations of Florida torreya in contrast to those of
California torreya (Schwartz 1993). Although growing in
markedly different climates, the 2 species responded similarly in water stress tests (Schwartz and others 1995).
California torreya is a slow-growing, medium-sized tree
found along streams and in canyon bottoms and other moist
locations (Griffin and Critchfield 1976; Storer and Usinger
1963; Sudworth 1908). In its shrub form, it is found on serpentine soils and in chaparral. In elevation, California torreya ranges from coastal lowlands to almost 2,130 m in the
southern Sierra Nevada. Under very favorable conditions,
trees may grow to 23 m or more in height and 60 to 90 cm
in diameter (Sudworth 1908). The tallest tree now on record
Common name(s)
Occurrence
T. californica Torr.
T. myristica Hook.
Tumion californicum (Torr.) Greene
T. taxifolia Arn.
Tumion taxifolium (Arn.) Greene
California torreya,
California-nutmeg,
stinking-yew, stinking-cedar
Florida torreya,
Florida-nutmeg, stinking-cedar
Sources: Griffin and Critchfield (1976), Kurz (1938), Little (1979), Stalter (1990), Sudworth (1908).
Torreya
1121
Peattie 1953). Both species were used locally for such purposes as shingles, fence posts, and firewood. They grow satisfactorily outside of their native range and have received
moderate use as ornamentals (Burke 1975; Sargent 1875;
Wilson 1938). Fruits of California torreya were collected for
food by native Californians, and the characteristics of its oil
compare favorably with those of pine-nut oil for cooking
purposes (Burke 1975). Squirrels have been observed eating
fruits and seeds of Florida torreya and antler-rubbing scars
provide evidence of use by deer (Bronaugh 1996; Nicholson
1990; Schwartz and Hermann 1993a).
Flowering and fruiting. Torreyas are dioecious. The
male flowers are small, budlike, and clustered on the under
sides of twigs in axils of leaves produced the previous year
(Abrams 1955; Jepson 1925; Sargent 1933; Sudworth 1908).
The female flowers are less numerous and occur on the
lower sides of the current years twigs. After fertilization,
they develop singly into sessile, thin-fleshed arils that
mature during the second season as green to purplish drupes
25 to 44 mm long (figure 1). When mature, the leathery
cover eventually releases a 25- to 30-mm yellow-brown seed
(Munz and Keck 1959) (figure 2). The thick woody inner
seedcoat is irregularly folded into the female gametophyte,
and the embryo is minute (figure 3).
Both species flower in March and April, with some
flowers of Florida torreya appearing as early as January and
some of California torreya extending into May (Rehder
1940; Sargent 1933; Stalter 1990; Weidner 1996). Under
favorable growing conditions, Florida torreya produces male
and female flowers about age 20 (Stalter 1990); in greenhouse conditions, 5-year-old sprouts produced pollen
(Schwartz 1996).
1122
the fruit is
Figure 3Torreya californica, California torreya: longitudinal section through a seed showing the folds of the inner
seedcoat extending into the endosperm.
then held in open storage until the outer cover turned dark;
then the pulp was softened in water and removed by rubbing
fruits against hardware cloth (Weidner 1996).
Fruit production of California torreya is common and
widespread enough to forestall concerns about shortage;
several hundred pounds may be collected in single commercial collections (Callahan 1996). The fruits are generally
picked from the trees but are sometimes collected after they
have been shed. Seed extraction is about the same as for
Florida torreya, with the softened pulp removed by water
pressure and some hand rubbing (Callahan 1996). Care is
needed to avoid damage to the relatively tender seedcoat.
Seed quality of California torreya is generally good and can
be improved sometimes by separating light seeds through
flotation.
Storage experience is short-term and fragmentary
because torreya seeds are generally used as available. Based
on incidental observations, the seeds may be recalcitrant, as
high moisture content appears necessary to maintain their
viability. California torreya has been stored in moist vermiculite or sphagnum moss at 2 to 7 C for up to 3 years
(Callahan 1996). Some seeds of both species will germinate
in lengthy cool or warm stratification (Callahan 1996;
Weidner 1996).
Seeds of California torreya averaged 324/kg (147/lb),
with a range of 243 to 421/kg (110 to 191/lb) in 3 samples
(Roy 1974). Florida torreya had 496 seeds/kg (225/lb) in
1 sample at a moisture content of 8.6% (Roy 1974).
Pregermination treatments and germination tests.
Standard germination test procedures have not been developed yet for torreya seeds. Both species require lengthy
after-ripening and stratification, but efforts made to date
have not identified methods for timely germination testing
of fresh or stored seeds.
As available, fresh seeds of Florida torreya have been
tested at Alfred B. Maclay State Gardens according to the 9
variations of methodology specified in the recovery plan for
1985
69
77
100
85
77
62
80
85
86
58
44
35
54.0
54.0
70.3
60.3
48.7
45.0
Torreya
1123
The urgency of conserving Florida torreya has stimulated development of its reproduction by cuttings (Bailo and
others 1998; Nicholson 1988, 1993). Up to 91% of cuttings
collected in November from trees throughout the species
native range rooted in a mixture of pumice, peat, and perlite
mixture. The cuttings were potted and grown for 2 years and
then shipped to botanic gardens and research institutions. A
database on living Florida torreya material is maintained by
The Center for Plant Conservation, headquartered at the
Missouri Botanical Garden, St. Louis, Missouri.
References
Abrams L. 1955. Illustrated flora of the Pacific states: Washington, Oregon,
and California.Volume I. Stanford, CA: Stanford University Press. 538 p.
AFA [American Forestry Association]. 2000. The national register of big
trees 20002001. American Forests 106(1): 2264.
Bailo BG, Determann R, Nicholson R, Sojkowski S. 1998. The ex situ conservation of stinking cedar. Public Garden 13(3): 911.
Boeshore I, Gray WD. 1936. An Upper Cretaceous wood: Torreya antiqua.
American Journal of Botany 23: 524528.
Bronaugh W. 1996. Champions on the brink. American Forests 102(1):
1619.
Burke JG. 1975. Human use of the California nutmeg tree, Torreya californica, and of other members of the genus. Economic Botany 29: 121139.
Callahan F. 1996. Personal communication. Central Point, OR: Callahan
Seeds.
Florin R. 1963. The distribution of conifer and taxad genera in time and
space. Acta Horti Bergiani 20(4): 121312 [Torreya: 262266].
Godfrey RK. 1988. Trees, shrubs, and woody vines of northern Florida and
adjacent Georgia and Alabama. Athens, GA: University of Georgia Press.
734 p.
Griffin JR, Critchfield WB. 1976. The distribution of forest trees in California.
Res. Pap. PSW-82 (reprinted with supplement). Berkeley, CA: USDA
Pacific Southwest Forest and Range Experiment Station. 118 p.
Harrar ES, Harrar JG. 1962. Guide to southern trees. 2nd ed. New York:
Dover. 709 p.
Jepson WL. 1925. A manual of the flowering plants of California. Berkeley:
University of California. 1238 p.
Kurz H. 1938. Torreya west of the Apalachicola River. Proceedings of the
Florida Academy of Sciences 3: 6677.
Little EL Jr. 1979. Checklist of United States trees, native and naturalized.
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Metcalf W. 1959. Native trees of the San Francisco Bay region. Berkeley:
University of California Press. 72 p.
Mirov NT, Kraebel CJ. 1939. Collecting and handling seeds of wild plants.
For. Pub. 5. Washington, DC: USDA Civilian Conservation Corps. 42 p.
Moore RP, ed. 1985. Handbook on tetrazolium testing. Zurich: International
Seed Testing Association: 73.
Munz PA, Keck DD. 1959. A California flora. Berkeley: University of
California Press. 1681 p.
Nicholson R. 1990. Chasing ghosts: the steep ravines along Floridas
Apalachicola River hide the last survivors of a dying tree species.
Natural History 12/90: 8, 1013.
Nicholson R. 1993. Rooting Torreya taxifolia, an endangered conifer of the
Florida panhandle. Botanic Gardens Conservation News 2(2): 13.
Nicholson RG. 1988. Propagation of some woody endemic plants of eastern North America. Combined Proceedings of the International Plant
Propagators Society 37: 468473.
1124
EuphorbiaceaeSpurge family
30,300/hl (10,700/bu)
6,100/kg (2,780/lb)
90
seed.
longitudinal sec-
1125
References
Bonner FT. 1974. Sapium sebiferum (L.) Roxb., tallowtree. In: Schopmeyer
CS, tech. coord. Seeds of woody plants in the United States. Agric.
Handbk. 540. Washington, DC: USDA Forest Service: 760.
Bringi NV. 1988. Progress in the chemistry and technology of non-traditional oilseeds/oils. Journal of the Oil Technologists Association of India
20(1): 19 [Seed Abstracts 16: 392; 1993].
Samson WD,Vidrine CG, Robbins JWD, Case JI. 1985. Chinese tallow seed
oil as a diesel fuel extender.Transactions of the American Society of
Agricultural Engineers 28: 14061409.
1126
PinaceaePine family
Tsuga Carr.
hemlock
Jill R. Barbour, Robert H. Ruth, and Robert P. Karrfalt
Ms. Barbour is a germination specialist at the USDA Forest Services National Seed Laboratory,
Dry Branch, Georgia; Dr. Ruth (deceased) retired from the Pacific Northwest Research Station;
Mr. Karrfalt is director of the USDA Forest Services National Seed Laboratory, Dry Branch, Georgia
Growth habit, occurrence, and use. Trees of the
hemlock genusTsuga spp.are tall, straight, late successional climax evergreens with conical crowns and slender,
horizontal to pendulous branches. Fourteen species have
been reported; 4 of these are native to the United States and
the others to the Himalayas, China, Taiwan, and Japan. The
name tsuga is a Japanese word meaning tree-mother (Dirr
1998). Native American names for the North Country (that
is, Canada), hoe-nadia, and for the lands of upper New
York, oh-neh-tah, both mean land of the hemlock
(Dirr 1998).
Of the 4 native species in the United States (table 1),
both eastern and western hemlocks are used commercially
for lumber and pulpwood. The bark of eastern hemlock has
been a source of tannin for the leather industry. In central
and southern Oregon and some other areas, mountain hemlock has become an important part of the softwood sawtimber volume.
Much of eastern hemlock has been severely affected by
the hemlock woolly adelgidAdelges tsugae Annandin
New England and the midAtlantic region (Dirr 1998). The
hemlock woolly adelgid has also been noted on Carolina
hemlock in the Tallulah Gorge in northeastern Georgia
(Price 2002). Although the hemlock woolly adelgid occurs
Common name(s)
Occurrence
eastern hemlock,
Canada hemlock, hemlock
Carolina hemlock
T. caroliniana Engelm.
T. heterophylla (Raf.) Sarg.
T. mertensiana (Bong.) Carr.
western hemlock,
Pacific hemlock, hemlock
mountain hemlock,
black hemlock
Tsuga
1127
1128
can trap more than 100 pollen grains, the average pollen
grain count per bract from controlled pollinations being 34,
with a range from 2 to 116 (Colangeli and Owens 1989a).
The ovuliferous scales elongate over the bracts, trapping the
pollen between the bracts and scales. About 4 to 7 days after
pollen germination, the pollen tubes grow into the micropyles; usually 1 to 6 pollen tubes and sometimes up to 10
pollen tubes have been found in each micropyle (Colangeli
and Owens 1989a). In western hemlock, pollen is not essential for seed cone enlargement and unpollinated ovules can
continue seedcoat development, but the seed will not have
an embryo or gametophytic tissue (Colangeli and Owens
1990a).
Cones mature in 1 season and are small, pendant, globose to ovoid or oblong, with scales longer than the bracts
(figure 1). Carolina hemlock has the largest seeds of the
native hemlocks, followed by mountain hemlock and eastern
hemlock, with western hemlock having the smallest seeds
(table 2; figure 2). Eastern hemlock has the smallest cones;
they measure 1.5 to 2.5 cm by 1 to 1.5 cm. Eastern hemlock
trees grown from eastern and southern sources have larger
cones than do those grown from northern and western
sources (Godman and Lancaster 1990). Western hemlock
cones measure 1 to 3.0 cm by 1to 2.5 cm; Carolina hemlock
cones measure 2.5 to 4 cm by 1.5 to 2.5 cm. Mountain hemlock have the largest cones, which measure 3 to 6 cm by 1.5
to 3 cm (FNAEC 1993; Harlow and Harrar 1968; Hough
1947; Sargent 1933).
Cone production of hemlock usually begins when trees
are 20 to 30 years of age, a little later if trees are shaded. All
4 species of hemlock bear some cones almost every year and
large crops are frequent (table 3). Cones often remain on the
hemlocks well into the second year, being especially conspicuous on the tops of mountain hemlock. Wisconsin had
good eastern hemlock cone crops on 61% of the 32 years
recorded (Godman and Lancaster 1990). Eastern hemlock
trees as old as 450 years have been seen bearing cones.
/kg
273794
167213
4171,120
132459
Average
/lb
/kg
/lb
124360
7697
189508
60208
412
573
251
187
260
114
Samples
69
2+
106
6
Sources: Burns and Honkala (1990); Buszewicz and Holmes (1961), Hill (1969), Rafn (1915),Toumey and Korstian (1952),Toumey and Stevens (1928), Ruth (1974).
Tsuga
1129
Table 3Tsuga, hemlock: height, seed-bearing age, seedcrop frequency, and cone ripeness criteria
Height at
maturity Year first
(m)
cultivated
Species
T. canadensis
1830
1736
2030
T. caroliniana
T. heterophylla
T. mertensiana
1221
1875
7.545
1881
1851
1854
2030
2030
23
15
58
15
Yellow-green
Green
Purple
Green with purple tips
Yellow-green to brown
Purple-brown
Tan to brown
Light brown
Brown with red-brown tips
Brown
Sources: Burns and Honkala (1990), den Ouden and Boom (1965), Franklin (1968), Frothingham (1915), Harlow and Harrar (1968), Harris (1969), Hough (1947), Merrill
and Hawley (1924), Olson and others (1959), Ruth (1974), Ruth and Berntsen (1955), Sudworth (1908).
Location
Flowering
Fruit ripening
Seed dispersal
T. canadensis
T. caroliniana
T. heterophylla
Aprearly June
MarApr
AprMay
Late May-June
Mid to late Apr
May 27June 5
June
Junemid-July
Aug
SeptOct
AugSept
SeptOct
Sept 15
SeptOct
Aug
Late SeptOct
Late Sept-Nov
Septwinter
OctMay
OctJune
Oct
SeptMay
Sept 17winter
T. mertensiana
Sources: Allen (1957), Burns and Honkala (1990); Ebell and Schmidt (1963), Frothingham (1915), Garman (1951), Gashwiler (1969), Godman (1953), Green (1939), Harris
(1967), Harris (1969), Heusser (1954), Hough (1947), James (1959), Leiberg (1900), Radford and others (1964), Ruth (1974), Ruth and Berntsen (1955).
1130
longitu-
1131
Seed
moisture (%)
Temp (C)
Viable period
(yrs)
68
79
68
8
8
5
13
3
18
0
18
18
21
57
5+
3+
23
Sources: Allen (1957), Barton (1954b, 1961), Jones (1962), Ruth (1974).
1132
Medium
T. canadensis
T. caroliniana
T. heterophylla
T. mertensiana
Temp (C)
15
35
15
12
5
Time (days)
30120
3090
2190
2156
90
Sources: Allen (1958), Babb (1959), Deffenbacher (1969), Devitt and Long (1969), Eide (1969), Olson and others (1959), Ruth (1974), Swingle (1939),Walters and others
(1960),Ward (1969),Weyerhaeuser (1969).
* Seeds were presoaked in tap water for 24 to 36 hours.
Germination of eastern hemlock seeds declines depending on the frequency and degree of drying following the
imbibition phase and on the intensity of light. Eastern hemlock seeds incubated in open petri dishes at a low light level
(645 lux) showed various germination values, from 50.2%
with decomposed birch medium to 0% with filter paper.
Seeds incubated in open petri dishes with decomposed birch
medium that were exposed to a moderate light level (4,682
lux) exhibited delayed initial germination and significantly
reduced total germination to half that at low light conditions
(Coffman 1978). The intensity of light had no effect on
seeds in covered petri dishes where a high moisture content
was maintained.
Seeds of western hemlock stratified for 3 weeks at 1 C
germinated faster than untreated seeds; longer stratification
periods caused additional but smaller increases in the rate of
germination (Bientjes 1954; Ching 1958). Stratification of
western hemlock seeds apparently has its main effect on
speed of germination; it has only a minor effect on total germination percentage. Seedlots stratified for 1 week reached
R50 (the number of days to reach 50% germination) 2.5 days
sooner than did unstratified seedlots. Seedlots stratified for
4 weeks reached R50 4.5 days sooner, and seedlots stratified
for 16 weeks reached R50 10.5 days sooner than did unstratified seedlots (Edwards 1973). Unstratified seedlots of western hemlock required nearly 2.5 weeks (18 days) to produce
the same number of germinants as did seedlots stratified for
3 months in 10 days (Edwards 1973). Western hemlock
seeds stratified for 1 week in plastic bags germinated about
1 day sooner than seeds stratified on filter paper (Edwards
1973). Presoaking the seeds for 48 hours was as effective in
reducing the germination rate as was 1 week of stratification
on filter paper (Edwards 1973). Immature western hemlock
seeds tend to have lower total germination as a result of
stratification (Allen 1958).
Experiments in Great Britain showed slightly increased
rates of germination following stratification when western
1133
Table 7Tsuga, hemlock: stratification period, germination test conditions, and results
Species
T. canadensis
T. caroliniana
T. heterophylla
T. mertensiana
Cold
stratification*
(days)
Daily
light
period
(hrs)
60120
030
2130
20
40
90
0
2130
0120
0
090
0
28
090
90
0
28
90
8
8
8
8
8
8
16
8
8
8
Dark
8
8
8
8
20
22
16
15
15
15
20
20
22
20
11
15
15
20
20
20
20
20
20
20
Days
60
28
28
28
28
28
28
34
2835
30
35
35
2530
28
28
60
28
28
28
Germination rate
(%)
(days)
1055
662
49
38
6275
61
1530
28
21
2030
1620
16
Sources: AOSA (2001), Buszewicz and Holmes (1961), Edwards and El-Kassaby 1996, Hill (1969), ISTA (1999), Ruth (1974), USDA FS (2002)
* Temperatures were 16 to 15 C.
Moisture-holding media were either blotters, Kimpak, sand, or peat.
1134
Germination
(%) Samples
38
1066
60
45
61
4080
5157
8291
53
56
86
86
47
91
90
62
81
97
72
15
912
3
3
9
9
3
5
146
25
44
43
4
19
19
1
4
3
5
1135
1136
References
Allen GS. 1957. Storage behavior of conifer seeds in sealed containers held
at 0 F, 32 F, and room temperature. Journal of Forestry 55: 278281.
Allen GS. 1958. Factors affecting the viability and germination behavior of
coniferous seed: 1. Cone and seed maturity, Tsuga heterophylla (Rafn)
Sarg. Forestry Chronicle 34: 266274.
AOSA [Association of Official Seed Analysts]. 2001. Rules for testing seeds.
Association of Official Seed Analysts. 126 p.
Babb MF. 1959. Propagation of woody plants by seed. Alaska Agricultural
Experiment Station Bulletin 26: 111.
Baldwin HI. 1930. The effect of after-ripening treatment on the germination
of eastern hemlock seed. Journal of Forestry 28: 853857.
Baldwin HI. 1934. Further notes on the germination of hemlock seed.
Journal of Forestry 32: 99100.
Barton LV. 1954a. Effect of subfreezing temperatures on viability of conifer
seeds in storage. Contributions of the Boyce Thompson Institute 18:
2124.
Barton LV. 1954b. Storage and packeting of seeds of Douglas-fir and western hemlock. Contributions of the Boyce Thompson Institute 18: 2537.
Barton LV. 1961. Seed preservation and longevity. New York: Interscience
Publishers. 216 p.
Benzian B. 1965. Experiments on nutrition problems in forest nurseries.
Volume 1. For. Comm. Bull. 37. London: Her Majestys Stationery Office.
251 p.
Bientjes W. 1954. The effects of temperature, seed moisture, and stratification on the germination behavior of western hemlock seed. For. Club
Res. Note 11.Vancouver: University of British Columbia. 7 p.
Bramlett DL, Belcher EW, DeBarr GL, Hartel GD, Karrfalt RP, Lantz CW, Miller
T, Ware KD,Yates HO. 1977. Cone analysis of southern pines: a guidebook. Gen.Tech. Rep. SE-13. Asheville, NC: USDA Forest Service,
Southeastern Forest Experiment Station.
Burns RM, Honkala BH. 1990. Silvics of North America.Volume 1, Conifers.
Agric. Handbk. 654. Washington, DC: USDA Forest Service. 1675 p.
Buszewicz G, Holmes GD. 1957. Seven years seed testing experience with
the tetrazolium viability test for conifer species. In: Report of the
Forestry Research Committee. Great Britain Forestry Commission
(London) 1957: 142151.
Tsuga
1137
1138
Ruth RH. 1974. Tsuga, hemlock. In: Schopmeyer CS, tech. cord. Seeds of
woody plants in the United States. Agric. Handbk. 450. Washington, DC:
USDA Forest Service: 819827.
Ruth RH, Berntsen CM. 1955. A 4-year record of Sitka spruce and western
hemlock seed fall. Res. Pap. 12. Corvallis, OR: USDA Forest Service
Pacific Northwest Forest and Range Experiment Station. 13 p.
Sargent CS. 1933. Manual of the trees of North America. Boston:
Houghton Mifflin. 910 p.
Schubert GH. 1954. Viability of various coniferous seeds after cold storage.
Journal of Forestry 52: 446447.
Smith M. 1997. Personal communication. Olympia: Washington
Department of Natural Resources, Webster Forest Nursery.
Stearns F, Olson J. 1958. Interactions of photo period and temperature
affecting seed germination in Tusga canadensis. American Journal of
Botany 45: 5358.
Sudworth GB. 1908. Forest trees of the Pacific Slope. Washington, DC:
USDA Forest Service. 441 p.
Swartley JC. 1945. Variations and uses of Canada hemlock. Cornell
Plantations 2: 36.
Swingle CF, comp. 1939. Seed propagation of trees, shrubs, and forbs for
conservation planting. SCS-TP-27. Washington, DC: USDA Soil
Conservation Service. 198 p.
Toumey JW, Korstian CF. 1952. Seeding and planting in the practice of
forestry. 3rd ed. New York: John Wiley & Sons. 520 p.
Toumey JW, Stevens CL. 1928. The testing of coniferous tree seeds at the
School of Forestry,Yale University, 19061926. For. Bull. 21. New Haven,
CT: Yale University School of Forestry. 46 p.
USDA Forest Service. 2002. Official seed testing results. Dry Branch, GA:
National Tree Seed Laboratory.
Vabre-Durrieu A. 1954a. Lhybride TsugaPicea hookeriana et ses parents:
des plantules.Travaux du Laboratoire Forestier de Toulouse.Tome 1
(vol 5, art. 15) 8 p.
Vabre-Durrieu A. 1954 b. Lhybride TsugaPicea hookeriana et ses parents:
etude chromosomique et caryologique.Travaux du Laboratoire
Forestier de Toulouse,Tome 1 (vol 5, art. 17) 4 p.
Walters J, Soos J, Haddock PG. 1960. The selection of plus trees in the
University of British Columbia Research Forest, Haney, B.C. Res. Pap. 33.
Vancouver: University of British Columbia Faculty of Forestry. 12 p.
Ward H. 1969. Personal correspondence. Olympia: Washington State
Department Natural Resources, L.T. Mike Webster State Forest
Nursery,
WFTSC [Western Forest Tree Seed Council]. 1966a. Sampling and service
testing western conifer seeds. Western Reforestation Coordinating
Committee of the Western Forestry and Conservation Association.
36 p.
WFTSC [Western Forest Tree Seed Council]. 1966b. Tree seed zone map.
Portland, OR: Western Forest Tree Seed Council.
Weyerhaeuser Co. 1969. Written correspondence.Tacoma, WA.
Tsuga
1139
FabaceaePea family
Ulex europaeus L.
common gorse
George P. Markin
Dr. Markin is a research entomologist at the USDA Forest Services Rocky Mountain
Research Station, Bozeman, Montana
Growth habit, occurrence, and use. Gorse is a leafless, spined shrub introduced from western Europe. In its
homeland, it grows 1 to 2 m tall and is primarily a nonaggressive invader of disturbed areas that is recognized as
useful for wildlife protection, soil stabilization, and revegetation. It has also been cultivated as an ornamental and as
forage for livestock, which feed on the soft, new growing
shoots. Its major use in the past, however, was for
hedgerows to contain livestock before barbed wire (Jobson
and Thomas 1964). As a useful plant, European settlers carried gorse to many parts of the world where it quickly
escaped from cultivation and formed aggressive feral populations. These feral plants grow 3 to 5 m tall in dense, spiny,
impenetrable stands that exclude desirable vegetation in pasture lands (Hill 1983; Sandrey 1985) and, in open forests,
interfere with reforestation and forest management
(Balneaves and Zabkiewicz 1981; Zabkiewicz 1976). Gorse
is presently recognized as one of the worst weeds in New
Zealand, Chile, and Tasmania and is recognized as a weed in
at least 15 other countries or island groups around the world
(Holm and others 1979).
In North America, gorse is still used to a limited extent
as an ornamental for its dense yellow flowers. In the eastern
United States, scattered feral populations have been recorded, but apparently these are not of an aggressive nature. By
contrast, along the Pacific Coast, gorse is found scattered
along the coastline from San Francisco, California, north to
Vancouver, British Columbia (Markin and others 1994).
Through most of this area, it is found in small, scattered
populations that are usually targeted for intensive control
programs to keep them from expanding. The major outbreak
along the southwestern coast of Oregon covers at least
15,000 ha and is a major problem in forest management.
This gorse population interferes with reforestation and,
because of the plants highly flammable nature, creates an
extreme fire hazard (Herman and Newton 1968). Gorse also
infests 14,000 ha at higher elevations on the islands of
Hawaii and Maui in Hawaii (Markin and others 1988).
1140
seed.
Other methods of germination include soaking in concentrated sulfuric acid for 1/2 to 11/2 hours, and mechanical
scarification (Buttler 1976), most simply done with emery
paper (Moss 1959).
Germination tests. Because of the noxious nature of
gorse in North America, standardized germination tests for
quality control have not been developed. In Europe, where
gorse is a beneficial native plant, germination tests at one
time were apparently developed in which seeds were tested
in germinators or sand flats at 20 C for 30 days using 400
pretreatment seeds/test (Rudolf 1974). Researchers have
reported no problem in obtaining germination by planting
scarified seeds 1 cm deep in different media. First signs of
germination are usually seen within 10 days of planting. In
15 to 25 days, seedlings are small rosettes with true leaves,
approximately 1.5 cm in diameter. Small leaves continue to
form until the plant is approximately 5 cm tall, at which
time the first spines are produced. During the remainder of
its life, the plant produces no more leaves, only spines. The
juvenile stage of the plant, from seed germination until
spines begin to form, requires 4 to 6 months in the field. In
Europe, large-scale germination in pots and direct seeding
into the field have been practiced in the past (Rudolf 1974).
Ulex
1141
References
Balneaves JM, Zabkiewicz JA. 1981. Gorse control: a review. In: Chavasse
CGR, ed. Proceedings, Forest Nursery and Establishment Practice in
New Zealand; 1982 March 2327; Rotorua, NZ: New Zealand Forest
Service, Forest Research Institute: 92105.
Butler JHB. 1976. A preliminary investigation of the hard seed characteristics of Ulex europaeus with emphasis on laboratory scarification treatments [thesis]. Palmerston North, New Zealand: Massey University. 82 p.
Chater EH. 1931. A contribution to the study of the natural control of
gorse. Bulletin of Entomological Research 22: 225235.
Hermann RK, Newton M. 1968. Tree plantings for control of gorse on the
Oregon coast. Res. Pap. 9. Corvallis: Oregon State University, Forestry
Research Laboratory. 12 p.
Hill RL 1983. Prospects for the biological control of gorse. New Zealand
Weed and Pest Control Conference: 5658.
Holm L, Pancho JV, Herberger JP, Plucknett DL. 1979. A geographical atlas
of world weeds. New York: Wiley-Interscience: 373 p.
Jobson HT, Thomas B. 1964. The composition of gorse (Ulex europaeus).
Journal of the Science of Food and Agriculture 15: 652656.
Markin GP, Yoshioka ER. 1989. Present status of biological control of the
weed gorse (Ulex europaeus L.) in Hawaii. In: Delfosse ES, ed.
Proceedings, 7th Symposium on Biological Control of Weeds; 1988
March 611; Rome. East Melbourne, Australia: CSIRO Publications:
357362.
Markin GP, Yoshioka ER, Brown RE. 1994. Gorse. In: Nechols JR, ed.
Biological control in western United States. Pub. 3361. Oakland:
University of California, Division of Agriculture and Natural Resources:
299302.
Markin GP, Dekker LA, Lapp JA, Nagata RF. 1988. Distribution of the weed
gorse (Ulex europaeus L.) a noxious weed in Hawaii. Newsletter of the
Hawaiian Botanical Society 27: 110117.
1142
UlmaceaeElm family
Ulmus L.
elm
Jill R. Barbour and Kenneth A. Brinkman
Ms. Barbour is a germination specialist at the USDA Forest Services National Seed Laboratory,
Dry Branch, Georgia; Dr. Brinkman retired from the USDA Forest Services
North Central Forest Experiment Station
Common name(s)
Occurrence
U. alata Michx.
Japanese elm
U. americana L.
U. crassifolia Nutt.
U. glabra Huds.
U. scabra Mill.
U. montana With.
U. cammpestris L. in part
U. japonica (Sarg. ex Rehd.) Sarg.
U. campestris var. japonica Rehd.
U. davidiana var. japonica (Rehd.) Nakai
U. laevis Pall.
U. pedunculata Pall.
U. effusa Willd.; U. racemosa Borkh.
U. minor Mill.
U. carpinifolia Gled.
U. parvifolia Jacq.
U. chinensis Pers.
U. procera Salisb.
U. pumila L.
U. rubra Mhl.
U. fulva Michx.
U. serotina Sarg.
U. thomasii Sarg.
U. racemosa Thomas
Ulmus
1143
Location
Flowering
Fruit ripening
Seed dispersal
U. alata
U. americana
U. crassifolia
U. glabra
U. japonica
U. laevis
U. parvifolia
U. pumila
U. rubra
U. serotina
U. thomasii
From S to Canada
SE US
Europe & Asia Minor
Japan
Massachusetts
NE US
E central US
F S to Canada
SE US
NE US
FebApr
FebMay
AugSept
MarApr
AprMay
AprMay
AugSept
MarApr
FebMay
Sept
MarMay
Apr
Late FebJune
SeptOct
MayJune
June
MayJune
SeptOct
AprMay
AprJune
Nov
MayJune
Apr
Mid-Marmid-June
Oct
MayJune
MayJune
SeptOct
AprMay
AprJune
Nov
MayJune
Seed
size (mm)
68
13
613
1525
1015
10
1014
1218
1013
1325
Sources: Asakawa (1969), Brinkman (1974), Burns and Honkala (1990), Dirr (1998), FNAEC (1997), Hora (1981), Little and Delisle (1962), Loiseau (1945), Pammel and
King (1930), Petrides (1958), Rehder (1940), Spector (1956), Stoeckeler and Jones (1957), Sus (1925) ,Vines (1960),Wappes (1932),Wyman (1947).
1144
longitudinal section
Table 3Ulmus, elm: height, seed-bearing age, seed crop frequency, and fruit ripeness criteria
Species
U. alata
U. americana
U. crassifolia
U. glabra
U. japonica
U. laevis
U. parvifolia
U. pumila
U. rubra
U. serotina
U. thomasii
Height at
maturity (ft)
50
120
100
130
100
100
80
80
70
60
100
Year first
cultivated
1820
1752
Long cultivated
1895
Long cultivated
1794
1860
1830
1903
1875
Minimum
seed-bearing
age (yr)
15
3040
3040
8
15
20
Years
between large
seedcrops
23
2
23
45
24
23
34
Yellow-brown
Brown
Yellow
Green
Light green to brownish
Yellow or brownish
Sources: Brinkman (1974), Burns and Honkala (1990), Dore (1965), FNAEC (1997), George (1937), Little and Delisle (1962), McDermott (1953),Van Dersal (1938),Vines
(1960),Wappes (1932).
1145
Seed
moisture (%)
Storage
temp (C)
Viable
period (yr)
Air-dried
34
Air-dried
Air-dried
Air-dried
Air-dried
1015
35
Air-dried
4
4
4
4
110
22
0
24
Cold
1
15
2
1
0.5
0.5
0.5
8
Sources: Barton (1939, 1953), Brinkman (1974), Heit (1967a&b), Kirby and
Santelmann (1964), Rohmeder (1942), Sus (1925).
Species
Place
collected
U. alata
U. americana
U. crassifolia
U. glabra
U. japonica
U. laevis
U. parvifolia
U. pumila
U. rubra
U. serotina
U. thomasii
Mississippi
Mississippi
Europe
Japan
Russia
US, Japan
Fruit/vol
kg/hl
lb/bu
5.8
46.5
7.710.3
4.5
35
68
112
71
67
40
6
63
121
72
41
149
7
222269
106240
130135
6699
117205
250372
88261
77119
1115
/lb
101119
48109
5961
3045
5393
114169
40119
3554
57
Samples
4
14
5
12+
2+
20+
6+
35+
10
Sources: Asakawa (1969), Brinkman (1974), Engstrom and Stoeckeler (1941), Goor (1955), Gorshenin (1941), Heit (1969), Rafn and Son (1928), Stoeckeler and Jones
(1957), Sus (1925), Swingle (1939),Taylor (1941),Van Dersal (1938),Wappes (1932).
1146
months (Dirr and Heuser 1987). Seeds of slippery elm, especially from northern sources, also may show dormancy;
70% of fresh seeds germinated and 57% germinated after 2
months of cold, moist stratification (Dirr and Heuser 1987).
Stratification at 5 C for 60 to 90 days before sowing
improves germination of cedar, smoothleaf, and September
elms (Brinkman 1974; Dirr and Heuser 1987; Maisenhelder
1968).
Winged, Scots, Japanese, English, Russian, Siberian,
and rock elms have no pregermination requirements (Dirr
and Heuser 1987). Fresh seedlots of Scots elm germinated at
98%, but after 2 months of cold, moist stratification, only
88% germinated (Dirr and Heuser 1987). English elm rarely
produces seeds, but fresh seeds will germinate at 100% with
or without 2 months of stratification (Dirr and Heuser 1987).
Fresh Siberian elm seeds germinated 96% and cold stratification did not improve germination (Dirr and Heuser 1987).
Fresh lacebark elm seeds will germinate without pretreatment, but once dried they require 1 to 2 months of cold,
moist stratification (Dirr and Heuser 1987).
Germination tests. Official testing rules for American
elm call for alternating temperatures of 30 C (day) for 8
hours and 20 C (night) for 16 hours for 14 days on wet
blotters and 10 days at a constant 20 C for Chinese and
Siberian elms (AOSA 2001). American elm seeds can also
germinate well at alternating temperatures of 21 C (day)
and 10 C (night) (Burns and Honkala 1990). The
International Seed Testing Association (1999) suggests testFigure 3Ulmus americana, American elm: seedling
development at 1, 3, and 21 days after germination.
ing for 14 days on wet blotters for all 3 species. ISTA also
suggests removal of the pericarp if germination is slow.
Germination tests of most species may also be made on sand
or peat in germinators at alternating temperatures of 30 C
(day) and 20 C (night). Rock elm seeds germinated 70 to
80% in a peat moss medium (Burns and Honkala 1990).
Light requirements may vary among species (table 6).
American elm can germinate in darkness but germination is
increased with the addition of light (Burns and Honkala
1990).
Germination is epigeal (figure 3); it usually peaks
within 10 days. Seedlots of stratified seeds complete germination in 10 to 30 days. With American elm seeds, germination can extend up to 60 days; seeds can lay on flooded
ground for a month without adversely affecting germination
(Burns and Honkala 1990). Radicles of rock elm emerge in
2 to 3 days in a petri dish and are 2.5 to 3.8 cm (1 to 1.5 in)
long by the 5th day; cotyledons opened about the 5th or 6th
day (Burns and Honkala 1990). Winged elms cotyledons are
oval with shallowly notched apexes and heart-shaped bases
and may persist 1 to 2 months on the seedling with primary
leaves appearing 1 week after germination in natural forest
conditions (Burns and Honkala 1990).
Nursery practice. Seeds of elm species ripening in
the spring are usually sown immediately after collection,
whereas seeds of fall-ripening species or of species requiring stratification are usually planted the following spring
(table 7). Beds should be kept moist until germination is
complete; shading is not usually necessary. From 5 to 12%
of the viable cedar elm seeds sown can be expected to produce plantable stock (Burns and Honkala 1990). One-yearold seedlings usually are large enough for field planting.
Rock elm seedlings have a persistent dormant bud, so
seedlings rarely develop more than a single pair of true
leaves in the first growing season (Burns and Honkala
1990). In northern Wisconsin, rock elm 1.5+0 nursery stock
averaged 27 cm (10.6 in) in height 5 years after planting and
52 cm (20.5 in) in height 10 years after planting; first-year
survival was 85% and 10th-year survival was 32% (Burns
and Honkala 1990). To improve survival in semiarid regions,
trees often are transferred into containers after 1 year in the
seedbeds (Goor 1955). Slippery elm is commonly used as
rootstock when grafting hybrid elms (Burns and Honkala
1990).
Ulmus
1147
Species
U. alata
U. americana
U. crassifolia
U. glabra
U. laevis
U. parvifolia
U. pumila
U. rubra
U. serotina
U. thomasii
Soil
32
Paper pads
30
Kimpak
30
Soil
32
Germinator or
sand
2130
Germinator or
sand
21
Paper pads
2029
Paper pads
Kimpak
30
Germinator or
sand
2030
Sand
30
Soil
32
Sand or petri dish
30
Days
Germinative Germinative
energy
capacity
Amount
Period Avg
(%)
(days)
(%)
Samples
Purity
(%)
21
20
20
21
15
14
28
1360
80
76
55
56
7
78
91
67
64
56
1
15
2
92
2025
3060
44
72+
21
20
20
30
1060
10
28
65
55
81
22+
2+
85
64
20
20
21
20
30
60
30
30
55
21
68
77
10
10
20
8
76
23
72
81
48
5
1
11
90
94
95
Sources: Arisumi and Harrison (1961), AOSA (2001), Engstrom and Stoeckeler (1941), Gorshenin (1941), Heit (1967a&, 1968), ISTA (1999), Johnson (1946), Kirby and
Santelman (1964), Maisenhelder (1968), McDermott (1953), NBV (1946), Rafn and Son (1928), Rohmeder (1942), Spector (1956), Stoeckeler and Jones (1957), Sus (1925),
Swingle (1939), USDA FS (2002),Wappes (1932).
* Light for 8 hours or more per day is recommended for American elm (AOSA 2001; ISTA 1999; McDermott 1953). Light is neither required nor inhibitory for germination
of winged elm (Loiseau 1945), and Chinese and Siberian elms (AOSA 2001; ISTA 1999).
Sowing
season*
U. alata
U. americana
U. crassifolia
U. glabra
U. laevis
U. parvifolia
U. pumila
U. rubra
U. serotina
U. thomasii
Summer
Spring
Spring
Summer
Summer
Spring
Summer
Spring
Spring
Spring
Seedlings/area
/m2
/ft2
2530
25
1538
23
14
Sowing depth
mm
in
06.4
6.4
06.4
06.4
0-6.4
4.86.4
6.4
6.4
06.4
6.4
01/4
1/
4
01/4
01/4
01/4
3/ 1/
16
4
1/
4
1/
4
01/4
1/
4
Tree
percent
Out-planting
age (yrs)
12
6
1220
37
1
1
1
12
12
12
1
1
2
Sources: Baker (1969), Deasy (1954), Engstrom and Stoeckeler (1941), George (1937), Kirby and Santelman (1964), Rohmeder (1942), Stoeckeler and Jones (1957), Sus
(1925), Swingle (1939),Toumey and Korstian (1942).
* Spring-sowing was preceded by stratification in sand or in a plastic bag at 4 to 5 C for 60 days.
1148
References
Arisumi T, Harrison JM. 1961. The germination of rock elm seeds. American
Nurseryman 114(7): 10.
Asakawa S. 1969. Personal correspondence. Meguro, Japan: Ministry of
Agriculture and Forestry.
AOSA [Association of Official Seed Analysts]. 2001. Rules for testing seeds.
Association of Official Seed Analysts. 126 p.
Baker LA. 1969. Personal correspondence. Elkton, OR: Oregon State
Forestry Department, Dwight L. Phipps Forest Nursery.
Barnard ES. 2002. New York City trees: a field guide for the metropolitan
area. New York: Columbia University Press: 212.
Barton LV. 1939. Storage of elm seeds. Contributions of the Boyce
Thompson Institute 10(2): 221233.
Barton LV. 1953. Seed storage and viability. Contributions of the Boyce
Thompson Institute 17(2): 87103.
Brinkman KA. 1974. Ulmus L., elm. In: Schopmeyer CS, tech. coord. Seeds of
woody plants in the United States. Agric. Handbk 450. Washington, DC:
USDA Forest Service: 829834.
Burns RM, Honkala BH. 1990. Silvics of North America.Volume 2,
Hardwoods. Agric. Handbk 654. Washington, DC: USDA Forest Service.
877 p.
Cram WH, Lindquist CH,Thompson AC. 1966. Seed viability studies:
American and Siberian elm. Summary Report Tree Nursery
Saskatchewan. 1965: 89.
Deasy JJ. 1954. Notes on the raising of forest trees in the nursery. Irish
Forestry 11(1): 1019.
Dirr MA. 1998. Manual of woody landscape plants: their identification,
orna mental characteristics, culture, propagation and uses. 5th ed.
Champaign, IL: Stipes Publishing. 1187 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant
propagation: from seed to tissue culture. Athens, GA: Varsity Press.
239 p.
Dore W. 1965. Ever tried rock elm seeds for eating? Canada Audubon
27(3): 9091.
Engstrom HE, Stoeckeler JH. 1941. Nursery practice for trees and shrubs.
Misc. Pub. 434. Washington, DC: USDA Forest Service. 159 p.
Fernald ML. 1970. Grays manual of botany. New York:Van Nostrand.
1632 p.
FNAEC [Flora of North America Editorial Committee]. 1997. Flora of
North America north of Mexico.Volume 3, Magnoliophyta: Magnoliidae
and Hamamelidae. New York: Oxford University Press. 590 p.
George EJ. 1937. Storage and dewinging of American elm seed. Journal of
Forestry 35(8): 769772.
Goor AY. 1955. Tree planting practices for arid areas. For. Dev. Pap. 6.
Rome: FAO. 126 p.
Gorshenin NM. 1941. Agrolesomelioratsiya [in Russian: Agro-forest melioration]. 392 p.
Heit CE. 1967a. Propagation from seed: 8. Fall planting of fruit and hardwood seeds. American Nurseryman 126(4): 1213, 8590.
Heit CE. 1967b. Storage of deciduous tree and shrub seeds. American
Nurseryman 126(10): 1213, 8694.
Heit CE. 1968. Thirty-five years testing of tree and shrub seed. Journal of
Forestry 66: 632634.
Heit CE. 1969. Personal communication. Geneva, NY: New York State
Agricultural Experiment Station.
Hora B. 1981. The Oxford encyclopedia of trees of the world. Oxford, UK:
Oxford University Press. 288 p.
ISTA [International Seed Testing Association]. 1999. International rules for
seed testing rules 1999. Seed Science and Technology 27(Suppl.). 333 p.
Johnson H. 1973. The international book of trees. New York: Simon and
Schuster. 288 p.
Johnson LPV. 1946. Effect of humidity on the longevity of Populus and
Ulmus seeds in storage. Canadian Journal of Research 24(Sec. C):
298302.
Kirby B, Santelmann PW. 1964. Germination and emergence of winged elm
seed. Weeds 12(4): 277279.
Krssmann G. 1960. Die Nadelgehlze. 2nd ed. Berlin. 335 p.
Lee MT, Lester DT. 1974. Floral receptivity in American elm. Canadian
Journal of Forest Research 4: 416417.
Little EL Jr, Delisle AL. 1962. Time periods in development: forest trees,
North American. In: Altman PL, Dittmer D, eds. Biological handbook on
growth. Washington, DC: Federation of American Societies for
Experimental Biology:Table 104.
Loiseau J. 1945. Les arbres et la fret. Paris. 204 p.
Maisenhelder LC. 1966. Unpublished data. Stoneville, MS: USDA Forest
Service, Southern Forest Experiment Station.
Ulmus
1149
LauraceaeLaurel family
1150
leaves, and seeds have been sold for pharmaceutical purposes such as treating catarrh, nervous disorders, rheumatism,
meningitis, intestinal colic, and dyspepsia (Peattie 1953;
Sargent 1895; Stuhr 1933).
California-laurel is used to a moderate extent as an ornamental evergreen. It has thick, glossy, medium-to-dark green
persistent leaves that turn orange or yellow before they drop
individually and contrasting pale yellow flowers. The very
dense aromatic foliage often shapes naturally into a pleasing, symmetrical, rounded crown. Since it was first cultivated in 1829 (Rehder 1940), it has demonstrated the ability to
grow well far outside its natural range (Stein 1958). It can
be grown as a decorative potted plant for lobbies and patios
and will tolerate moderate pruning (Kasapligil and Talton
1973).
California-laurel also has wildlife valuesyoung
sprouts are choice browse in spring and summer. Year-long
use is rated by Sampson and Jespersen (1963) as good to
fair for deer (Odocoileus spp.) and fair to poor for cattle,
sheep, and goats. Longhurst and others (1952) list it as a
principal browse species for deer in the north coastal ranges
of California. Silver gray-squirrels (Sciurus griseus), duskyfooted wood rats (Neotoma fuscipes), and Stellers jays
(Cyanocitta stelleri) feed on the seeds extensively (Bailey
1936; Van Dersal 1938). Hogs eat both seeds and roots
(Jepson 1910; Van Dersal 1938).
Flowering and fruiting. California-laurel flowers regularly and often profusely. The small, pale yellow, perfect
flowers grow on short-stemmed umbels that originate from
leaf axils or near the terminal bud (figure 1). Flower buds
develop early; those for the following year become prominent as current-year fruits are maturing. Within its long
northsouth range, California-laurel has been reported to
flower in all months from November to May, beginning
before new leaves appear (Jepson 1910; Kasapligil and
Talton 1973; Rehder 1940; Unsicker 1974). The flowering
period may stretch into late spring and summer with the
occasional appearance of flowers originating in axils of the
current years developing leaves (Sargent 1895). Californialaurel flowers at an early age; flowers have been observed
on short whiplike shrubs and on 1-year-old sucker growth
that originated on a long broken stub. Small insects appear
to be the chief pollinaters (Kasapligil 1951).
Seedcrops are abundant in most years (Stein 1974).
Although umbels bear 4 to 9 flowers each, generally only 1
to 3 fruits set (Jepson 1910). The age when a tree first bears
fruit, the age for maximum production, and the average
quantity produced have not been reported. Seeds are produced in abundance after trees are 30 to 40 years old
Umbellularia
1151
References
AFA [American Forestry Association]. 2000. The national register of big
trees, 200001. American Forests 106 (1): 2264.
Bailey V. 1936. The mammals and life zones of Oregon. North American
Fauna 55. Washington, DC: USDA Bureau of Biological Survey. 416 p.
Britton NL. 1908. North American trees. New York: Henry Holt &
Co. 894 p.
Chesnut VK. 1902. Plants used by the Indians of Mendocino County,
California. United States National Herbarium Contributions 7(3):
295408.
Coombes AJ. 1992. Trees: eyewitness handbook. New York: Dorling
Kindersley: 189.
Davis EM. 1947. Machining of madrone, California laurel, tanbark oak, and
chinquapin. Pub. R1727. Madison, WI: USDA Forest Service, Forest
Products Laboratory. 6 p.
Drake ME, Stuhr ET. 1935. Some pharmacological and bactericidal properties of umbellulone. Journal of the American Pharmaceutical Association
24(3): 196207.
Eastwood A. 1945. New varieties of two well-known Californian plants.
Leaflets of Western Botany 4(6): 166167.
Eliot WA. 1938. Forest trees of the Pacific Coast. New York: G. P. Putnams
Sons. 565 p.
Harlow WM, Harrar ES, White FM. 1979. Textbook of dendrology. 6th ed.
New York: McGraw-Hill Book Co. 510 p.
Harvey AG. 1947. Douglas of the fir. Cambridge, MA: Harvard University
Press. 290 p.
Jepson WL. 1910. The silva of California. Memoirs of the University of
California 2: 1480.
Kasapligil B. 1951. Morphological and ontogenetic studies of Umbellularia
californica Nutt. and Laurus nobilis L. University of California Publications
in Botany 25(3): 115239.
Kasapligil B,Talton B. 1973. A flowering calendar for native California
shrubs in the San Francisco Bay region. California Horticultural Journal
34(1): 1230.
Kellogg A. 1882. California laurel or bay tree. In: Forest trees of California.
Sacramento: California State Mining Bureau: 137140.
Lippitt L. 1995. Personal communication. Davis: California Department of
Forestry and Fire Protection, Lewis A. Moran Reforestation Center.
Longhurst WM, Leopold AS, Dasmann RF. 1952. A survey of California
deer herds, their ranges and management problems. Game Bull. 6.
Sacramento: California Department of Fish and Game. 136 p.
McBride JR. 1969. Plant succession in the Berkeley Hills [PhD thesis].
Berkeley: University of California.
Umbellularia
1153
EricaceaeHeath family
Vaccinium L.
blueberry, cranberry
Jason J. Griffin and Frank A. Blazich
Dr. Griffin is assistant professor at Kansas State Universitys Department of Horticulture, Forestry, and Recreation
Resources, Manhatten, Kansas; Dr. Blazich is alumni distinguished graduate professor of plant propagation and tissue
culture at North Carolina State Universitys Department of Horticultural Science, Raleigh, North Carolina
1154
Table 1Vaccinium, blueberry and cranberry: nomenclature, plant height, and natural occurrence
Plant height
(cm)
Common name(s)
V. angustifolium Ait.
V. lamarckii Camp
V. nigrum (Wood) Britt.
V. angustifolium
var. hypolasium Fern.
var. laevifolium House
var. nigrum (Wood) Dole
var. brittonii Porter ex Bickn.
V. arboreum Marsh.
V. arboreum var. glaucescens (Greene) Sarg.
Batodendron andrachniforme Small
Batodendron arboreum (Marsh.) Nutt.
V. corymbosum L.
V. constablaei Gray
V. corymbosum var. albiflorum (Hook.) Fern.
V. corymbosum var. glabrum Gray
Cyanococcus corymbosus (L.) Rydb.
Cyanococcus cuthbertii Small
V. macrocarpon Ait.
Oxycoccus macrocarpus (Ait.) Pursh
lowbush blueberry,
late sweet blueberry,
low sweet blueberry
V. myrtilloides Michx.
V. angustifolium var myrtilloides
(Michx.) House
V. canadense Kalm ex A. Rich.
Cyanococcus canadensis (Kalm
ex A. Rich) Rydb.
V. ovatum Pursh.
Canadian blueberry,
velvet-leaf blueberry,
velvetleaf huckleberry,
sour-top blueberry
35 14
California huckleberry,
evergreen huckleberry,
shot huckleberry
small cranberry
82 42
V. oxycoccos L.
V. palustre Salisb.
Oxycoccus palustris Pers.
Oxycoccus quadripetalus Gilib.
V. virgatum Ait.
V. virgatum var. ozarkense Ashe
V. virgatum var. speciosum Palmer
V. parviflorum Gray; V. amoenum Ait.
V. ashei Rehd.; V. corymbosum
var. amoenum (Ait.) Gray
Cyanococcus virgatus (Ait.) Small
Cyanococcus amoenus (Ait.) Small
V. vitis-idaea L.
18 9
sparkleberry,
farkleberry
Occurrence
Labrador & Newfoundland;W to Manitoba
& Minnesota; S to Illinois, Delaware, &
Pennsylvania; mtns of Virginia &
West Virginia
311 102
230 60
highbush blueberry,
American blueberry,
swamp blueberry
cranberry, large
cranberry, American
cranberry
rabbiteye blueberry,
smallflower blueberry
63
21
300 100
lingonberry, cowberry,
foxberry, mountain cranberry
73
Newfoundland,W to Minnesota, S to N
Illinois, N Ohio, & central Indiana;
Appalachian Mtns to Tennessee &
North Carolina
Central Labrador to Vancouver Island,
Northwest Territories SE to
Appalachian Mtns
be placed in a food blender, covered with water, and thoroughly macerated using several short (5-second) power
bursts. More water is added to allow the pulp to float while
the sound seeds (figures 2 and 3) settle to the bottom.
Repeating this process several times may be necessary to
achieve proper separation of seeds and pulp (Galletta and
Ballington 1996). Seed weights are listed in table 3.
Vaccinium
1155
Table 2Vaccinium, blueberry and cranberry: phenology of flowering and fruiting for cultivated species
Species
Flowering
Fruit ripening
V. angustifolium
V. arboreum
V. corymbosum
V. macrocarpon
V. myrtilloides
V. ovatum
V. virgatum
V. vitis-idaea
MayJune
MayJune
June
MayJune
MarJuly
MarMay
MayJune
JulyAug
OctDec
JuneAug
SeptOct
AugSept
JuneAug
Aug
Species
V. angustifolium
V. arboreum
V. corymbosum
V. macrocarpon
V. myrtilloides
V. ovatum
V. oxycoccos
V. virgatum
V. vitis-idaea
Berry
diameter
(mm)
8
8
8
12
7
7
9
12
9
1
1
1
2
1
1
2
4
1
Cleaned
seeds/weight
/kg
/lb
3.90 x 106
1.01 x 106
2.20 x 106
1.09 x 106
3.81 x 106
2.99 x 106
1.46 x 106
3.54 x 104
1.45 x 106
4.59 x 105
1.00 x 106
4.95 x 105
1.73 x 106
1.36 x 106
6.62 x 105
1.61 x 104
Vaccinium
1157
Figure 2Vaccinium, blueberry: seeds of V. angustifolium, lowbush blueberry (A); V. arboreum, sparkleberry (B);
V. virgatum, rabbiteye blueberry (C); V. corymbosum, highbush blueberry (D); V. macrocarpon, cranberry (E);
V. myrtilloides, Canadian blueberry (F); and V. ovatum, California huckleberry (G).
1158
References
Aalders LE, Hall IV. 1975. Germination of lowbush blueberry seeds stored
dry and in fruit at different temperatures. HortScience 10(5): 525526.
Aalders LE, Hall IV. 1979. Germination of lowbush blueberry seeds as
affected by sizing, planting cover, storage, and pelleting. Canadian Journal
of Plant Science 59(2): 527530.
Aalders LE, Hall IV, Brydon AC. 1980. Seed production and germination in
four lowbush blueberry clones. HortScience 15(5): 587588.
Ballington JR. 1998. Personal communication. Raleigh: North Carolina State
University.
Ballington JR, Galletta GJ, Pharr DM. 1976. Gibberellin effects on rabbiteye
blueberry seed germination. HortScience 11(4): 410411.
Brissette L,Tremblay L, Lord D. 1990. Micropropagation of lowbush blueberry from mature field-grown plants. HortScience 25(3): 349351.
Crossley JA. 1974. Vaccinium, blueberry. In: Schopmeyer CS, tech. coord.
Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 840843.
Devlin RM, Karczmarczyk SJ. 1975. Effect of light and gibberellic acid on the
germination of Early Black cranberry seeds. Horticultural Research 15:
1922.
Devlin RM, Karczmarczyk SJ. 1977. Influence of light and growth regulators
on cranberry seed dormancy. Journal of Horticultural Science 52(2):
283288.
Devlin RM, Karczmarczyk SJ, Deubert KH. 1976. The influence of abscisic
acid in cranberry seed dormancy. HortScience 11(4): 412413.
Dirr MA. 1990. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation and uses. Champaign, Ill:
Stipes Publishing. 1007 p.
Dirr MA, Heuser Jr. CW. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Dweikat IM, Lyrene PM. 1988. Adventitious shoot production from leaves
of blueberry cultured in vitro. HortScience 23(3): 629.
Dweikat IM, Lyrene PM. 1989. Response of highbush blueberry seed germination to gibberellin A3 and 6N-benzyladenine. Canadian Journal of
Botany 67(11): 33913393.
Everett TH. 1981. The New York Botanical Garden illustrated encyclopedia
of horticulture.Volume 10. New York: Garland Publishing: 32253601.
Vaccinium
1159
EuphorbiaceaeSpurge family
cross-section of a
major problem. There are no definitive storage data for tungoil tree seeds, but they are considered short-lived and are
normally planted the spring following harvest. Their high
oil content suggests that storage for long periods may be
difficult.
References
Bailey LH. 1949. Manual of cultivated plants. rev. ed. New York: Macmillan.
1116 p.
Brown CA. 1945. Louisiana trees and shrubs. Bull. 1. Baton Rouge: Louisiana
Forestry Commission. 262 p.
Brown CL, Kirkman LK. 1990. Trees of Georgia and adjacent states.
Portland, OR:Timber Press. 292 p.
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
McCann LP. 1942. Development of the pistillate flower and structure of the
fruit of tung (Aleurites fordii). Journal of Agricultural Research 65:
361378.
Potter GF, Crane HL. 1951. Tung production. Farm. Bull. 2031. Washington,
DC: USDA. 41 p.
Vines RA. 1960. Trees, shrubs and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
Vernicia
1161
CaprifoliaceaeHoneysuckle family
Viburnum L.
viburnum
Franklin T. Bonner, John D. Gill, and Franz L. Pogge
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi; Dr. Gill and Mr. Pogge retired from the USDA Forest Services Northeastern
Forest Experiment Station
Growth habit, occurrence, and use. Among the 135
or so viburnum species, 12 that are either native to North
America or have been introduced are discussed here (table
1). All 12 species are deciduous shrubs or small trees. Their
characteristics place the viburnums among the most important genera for wildlife food and habitat and environmental
forestry purposes. The attractive foliage, showy flowers, and
fruits of viburnums have ensured their widespread use as
ornamental plants as well. The fruits of most species are
Common name(s)
Occurrence
V. acerifolium L.
V. dentatum L.
V. pubescens (Ait.) Pursh
V. lantana L.
V. lantanoides Michx.
V. alnifolium Marsh.
V. grandifolium Ait.
V. lentago L.
V. nudum var. nudum L.
V. cassinoides L.
V. nudum var. cassinoides
(L.) Torr. & Gray
V. opulus L
V. opulus var. amerieanum Ait.
V. trilobum Marsh.
V. prunifolium L.
V. rafinesquianum J. A. Schultes
V. affine Bush ex Schneid.
V. affine var. hypomalacum Blake
V. recognitum Fern.
V. rufidulum Raf.
witherod viburnum,
wild-raisin, witherod
European cranberrybush,
cranberrybush, Guelder rose,
highbush-cranberry
blackhaw, stagbush, sweethaw
downy arrowwood,
Rafinesque viburnum
smooth arrowwood,
arrowwood
rusty blackhaw, southern
blackhaw, bluehaw, blackhaw,
southern nannyberry
1162
cluster of fruits
Location
Flowering
Fruit ripening
Seed dispersal
V. acerifolium
Midrange
West Virginia
South
Midrange
Extremes
Midrange
Midrange
West Virginia
New York
Midrange
Extremes
South
Midrange
Extremes
Midrange
Extremes
Midrange
Extremes
Midrange
Extremes
North
South
South
North
MayAug
AprMay
MayJune
JuneAug
MayJune
MayJune
May
MayJune
AprJune
AprJune
JuneJuly
MayJuly
MayJune
MayJuly
AprMay
AprJune
JuneJuly
MayJune
MayJune
AprMay
MarApr
MayJune
JulyOct
Late Oct
Late July
SeptOct
JulyNov
AugSept
AugSept
Late Sept
AugSept
SeptOct
Mid July
SeptOct
SeptOct
JulyOct
AugSept
SeptOct
SeptOct
JulyAug
SeptOct
JulySept
AugSept
JulyAug
SeptOct
Fall
NovDec
FallSpring
to Dec
to Feb
SeptFeb
Fall
OctNov
AugOct
OctMay
FallSpring
OctNov
MarMay
OctMay
to Mar
OctApr
Oct
to Dec
to Feb
Dec
V. dentatum
V. lantana
V. lantanoides
V. lentago
V. nudum var. nudum
V. nudum var.
cassinoides
V. opulus
V. prunifolium
V. rafinesquianum
V. recognitum
V. rufidulum
Sources: Brown and Kirkman (1990), Donoghue (1980), Gill and Pogge (1974).
Viburnum
1163
longitudinal
Table 3Viburnum, viburnum: growth habit, height, seed-bearing age, and seedcrop frequency
Species
Growth habit
V. acerifolium
V. dentatum
V. lantana
V. lantanoides
V. lentago
V. nudum var. nudum
V. nudum var. cassinoides
V. opulus
V. prunifolium
V. rafinesquianum
V. recognitum
V. rufidulum
Erect shrub
Erect shrub
Shrub or tree
Erect or trailing shrub
Shrub of tree
Shrub or tree
Erect shrub
Erect shrub
Shrub or tree
Shrub
Erect shrub
Shrub or tree
Height at
maturity (m)
Year first
cultivated
Seed-bearing
age (yrs)
1736
1736
1820
1761
1761
1727
1830
23
34
35
810
56
5
2
5
5
3
10
1.8
3
4
5
2
3
3.5
V
Years
between large
seedcrops
1
3 or 4
1
Table 4Viburnum, viburnum: fruit and seed weight and yield data
Cleaned seeds/weight
Species
V. acerifolium
V. dentatum
V. lantana
V. lantanoides
V. lentago
V. nudum var. cassinoides
V. opulus
V. prunifolium
V. rufidulum
Dried fruits/wt
/kg
/lb
10,600
16,700
4,850
6,600
12,100
5,200
4,800
7,580
2,200
3,000
5,500
2,360
Range
/kg
24,05036,600
32,20071,900
9,25029,100
4,85027,350
55,10063,950
20,70039,250
8,80013,230
/lb
10,90016,600
14,60032,600
4,20013,200
2,20012,400
25,00029,000
9,40017,800
4,0006,000
/kg
28,000
45,000
19,200
25,350
13,000
60,850
30,000
10,600
Average
/lb
13,100
20,400
8,700
11,500
5,900
27,6003
13,600
4,800
Samples
5
6
2
11
21
12
5
Viburnum
1165
180510
0
150
150270
60
6090
150270
360510
360510
180360
60120
0
75
60120
90
3060
3060
60120
75
0
Germination
test duration
60+
60
100
120
120
60
60+
60+
Germination percentage
Avg (%)
Samples
32
43
51
67
60
75
69
3
3
2
3+
2
that seeds need not be handled after their roots emerge during the warm stratification period (Rollins 1970). Seeds of
species with more shallow dormancy can be sown in the fall
shortly after collection and extraction. For the several
species that may be handled in this manner, the latest sowing dates for optimum seedling percentages in the ensuing
year are listed in table 6. Sowing done somewhat earlier
than these dates gave nearly as good results, but sowing at
later dates reduced germination percentages.
1166
Figure 6Viburnum lentago, nannyberry: seedling development from stratified seedroot development during
warm stratification (about 150 days) (left); very little development during ensuing cold stratification (about 120 days)
for breaking epicotyl dormancy (middle); subsequent
development at germinating temperatures (right).
Table 6Viburnum, viburnum: latest allowable dates for sowing in nurserybeds and seedling percentages obtained
in the following year
Species
Location
Latest allowable
sowing date*
Seedling %
V. acerifolium
V. lantana
V. lentago
V. opulus
V. prunifolium
V. recognitum
New York
Ohio
Ohio
New York
New York
New York
May 1
Oct 21
Oct 7
July 1
May 1
May 1
55
90
75
87
26
32
References
Barton LV. 1951. A note on the germination of Viburnum seeds. University
of Washington Arboretum Bulletin 14: 1314, 27.
Barton LV. 1958. Germination and seedling production of species of
Viburnum. Proceedings of the International Plant Propagators Society 8:
15.
Brown CL, Kirkman LK. 1990. Trees of Georgia and adjacent states.
Portland, OR:Timber Press. 292 p.
Chadwick LC. 1935. Practices in propagation by seeds: stratification treatment for many species of woody plants described in fourth article of
series. American Nurseryman 62: 39.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seeds to tissue culture. Athens, GA:Varsity Press. 239 p.
Donoghue M. 1980. Flowering times in Viburnum. Arnoldia 40: 222.
Edminster FC. 1947. The ruffed grouse: its life story, ecology and management. New York: Macmillan. 385 p.
Fedec P, Knowles RH. 1973. Afterripening and germination of seeds of
American highbush cranberry (Viburnum trilobum). Canadian Journal of
Botany 51: 17611764.
Fernald ML. 1950. Grays manual of botany. 8th ed. New York: American
Book Co. 1632 p.
Giersbach J. 1937. Germination and seedling production of species of
Viburnum. Contributions of the Boyce Thompson Institute 9: 7990.
Gill JD, Pogge FL. 1974. Viburnum L., viburnum. In: Schopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 844850.
Gould WP. 1966. The ecology of Viburnum alnifolium Marsh [PhD thesis].
Syracuse: State University of New York, College of Forestry. 246 p.
Halls LK. 1973. Flowering and fruiting of southern browse species. Res. Pap.
SO-90. New Orleans: USDA Forest Service, Southern Forest
Experiment Station. 10 p.
Heit CE. 1967. Propagation from seed: 11. Storage of deciduous tree and
shrub seeds. American Nurseryman 126: 1213, 8694.
Hottes AC. 1939. The book of shrubs. New York: DeLa Mare Co. 435 p.
ISTA [International Seed Testing Association]. 1993. International rules for
seed testing: rules 1993. Seed Science and Technology 21 (Suppl.):
1259.
Krochmal A, Walters RS, Doughty RM. 1969. A guide to medicinal plants of
Appalachia. Res. Pap. NE-138. Upper Darby, PA: USDA Forest Service,
Northeast Forest Experiment Station. 291 p.
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Martin AC, Zim HS, Nelson AL. 1951. American wildlife and plants: a guide
to wildlife food habits. New York: Dover. 500 p.
Miliczky ER, Osgood EA. 1979. Insects visiting bloom of withe-rod
Viburnum cassanoides L. in the Orono, Maine, area. Entomological News
90(3): 131134.
Rollins JA. 1970. Viburnums [unpublished document]. Amherst: University of
Massachusetts, Department of Botany. 21 p.
Smith BC. 1952. Nursery research at Ohio State. American Nurseryman
95: 15, 9496.
Spinner GP, Ostrum GF. 1945. First fruiting of woody food plants in
Connecticut. Journal of Wildlife Management 9: 79.
Vines RA. 1960. Trees, shrubs and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
Viburnum
1167
VerbenaceaeVerbena family
Vitex agnus-castus L.
lilac chastetree
John C. Zasada and C. S. Schopmeyer
Dr. Zasada retired from the USDA Forest Services North Central Research Station; Dr. Schopmeyer
(deceased) retired from the USDA Forest Services National Office,Washington, DC
Other common names. chaste-tree, monks-pepper
tree, hemptree (Bailey 1949).
Growth habit, occurrence, and use. The genus Vitex
occurs in both hemispheres in the tropical and subtropical
zones. About 380 taxa have been described (Bredenkamp
and Botha 1993). Lilac chastetree, a deciduous, strongly
aromatic shrub or small tree, is one of the few species in the
genus that is native to the temperate zones, but it is not
native to North America (Bailey 1949). It has, however, naturalized in much of the southeastern United States.
In Washington on the west side of the Cascades, it
attains a height of 1.8 m, increasing in more southerly latitudes to a height of 7.7 m in the low desert of southern
California (Williamson 1967). Multiple stems support a
broad spreading form, but shade trees with a single stem can
be developed by pruning (Williamson 1967).
In the eastern United States, the species is hardy as far
north as New York (USDA Hardiness Zone 6), but marginally so; it performs better further south, in USDA Hardiness
Zones 89 (LHBH 1076; Dirr 1990; Moldenke 1968). This
species is less hardy than negundo chastetree (Vitex negundo
L.), which is also planted as an ornamental (Dirr 1990) and
has been cultivated as an ornamental in southern Europe, the
Middle East, India, and Brazil (Moldenke 1968). Lilac
chastetree was introduced as an ornamental into the United
States in 1570 (Rehder 1940). The species has value in shelterbelt plantings (Engstrom and Stoeckeler 1941).
Since the days of Dioscorides in the first century AD,
seeds of this species have been noted for their ability to subdue sexual urges in men, hence the name chastetree
(Moldenke 1968; Polunin and Huxley 1966). This property
was recognized as being useful to celibates and this in turn
led to the name monks-peppertree. However, these properties are questioned today. There is evidence that phytomedicines from the chastetree are useful in the treatment of
menstrual disorders in women (Bohnert and Hahn 1990).
Because of the aromatic pungency of fresh seeds, however,
1168
Vitex
1169
References
Bailey LH. 1949. Manual of cultivated plants most commonly grown in the
continental United States and Canada. New York: Macmillan. 1116 p.
Bohnert KJ, Hahn G. 1990. Phytotherapy in gynecology and obstetrics: Vitex
agnus-castua. Acta Medica Emperica 9: 494502.
Bredenkamp CL, Botha DJ. 1993. A synopsis of the genus Vitex L.
(Verbenaceae) in South Africa. South African Journal of Botany 59(6):
611622.
Dirr MA. 1990. Manual of woody landscape plants: their identification,
ornamental characteristics, culture, propagation and uses. Champaign, IL:
Stipes Publishing Co. 1007 p.
Dirr MA, Heuser Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Engstrom HE, Stoeckeler JH. 1941. Nursery practice for trees and shrubs
suitable for planting on the prairie-plains. Misc. Pub. 434. Washington,
DC: USDA Forest Service. 159 p.
King CM. 1932. Germination studies of woody plants with notes on some
buried seeds. Proceedings of the Iowa Academy of Science 39: 6576.
1170
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan: 11611162.
Moldenke HN. 1968. Additional notes on the genus Vitex. 7. Phytologia
16(6): 487502.
Polunin H. 1966. Flowers of the Mediterranean: 154155 [quoted by
Moldenke 1968].
Rehder A. 1940. Manual of cultivated trees and shrubs hardy in North
America. New York: Macmillan. 996 p.
Schopmeyer CS. 1974. Vitex agnus-castus L., lilac chastetree. In: Schopmeyer
CS, tech. coord. Seeds of woody plants in the United States. Agric.
Handbk. 450. Washington, DC: USDA Forest Service: 851852.
Verma SC, Misra PN. 1989. Biomass and energy production in coppice
stands of Vitex negundo L. in high density plantations on marginal lands.
Biomass 19: 189194.
Williamson JF. 1967. Sunset western garden book. Menlo Park, CA: Lane
Magazine and Book Co. 448 p..
VitaceaeGrape family
Vitis labrusca L.
fox grape
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station
Mississippi State, Mississippi
seed.
longitudinal section
Vitis
1171
References
AOSA [Association of Official Seed Analysts]. 1993. Rules for testing seeds.
Journal of Seed Technology 16(3): 1113.
Bonner FT, Crossley JA. 1974. Vitis labrusca L., fox grape. In: Schopmeyer
CS, tech. coord. Seeds of woody plants in the United States. Agric.
Handbk. 450. Washington, DC: USDA Forest Service: 853854.
Cawthon DL, Morris JR. 1982. Relationship of seed number and maturity to
berry development, fruit maturation, hormonal changes, and uneven
ripening of Concord (Vitis labrusca L.) grapes. Journal of the American
Society for Horticultural Science 107: 10971104.
1172
Simonov IN. 1963. [The influence of micro-elements and growth substances on seed germination and seedling growth of vines.] Venodelie I
Venogradarstvo 23(4): 3537 [Horticultural Abstracts 34(518); 1964].
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
Vories KC. 1981. Growing Colorado plants from seed: a state of the art.
Gen.Tech. Rep. INT-103. Ogden, UT: USDA Forest Service,
Intermountain Forest and Range Experiment Station. 80 p.
ArecaceaePalm family
Washingtonia
1173
References
Bomhard ML. 1950. Palm trees in the United States. Agric. Info. Bull. 22.
Washington, DC: USDA. 26 p.
DeLeon NJ. 1958. Viability of palm seeds. Principes 2: 9698.
DeMason DA. 1988. Embyro structure and storage reserves histochemistry in the palm Washingtonia filifera. American Journal of Botany 75(3):
330337.
Emery D. 1969. Correspondence. Santa Barbara, CA: Santa Barbara Botanic
Garden.
Jepson WL. 1910. The silva of California.Volume 2. Berkeley: University of
California Press. 283 p.
Khan MI. 1982. Allelopathic potential of dry fruits of Washingtonia filifera:
inhibition of seed germination. Physiologia Plantarum 54(3): 323328.
Loomis HF. 1950. The preparation and germination of palm seeds.
Principes 2: 98102.
1174
AgavaceaeCentury-plant family
Yucca L.
yucca
Robert R. Alexander, Floyd W. Pond, and Jane E. Rodgers
Mr. Alexander and Mr. Pond retired from the USDA Forest Services Rocky Mountain Forest and Range
Experimental Station; Ms. Rodgers is at the Point Reyes National Seashore, Point Reyes, California
Common name(s)
Occurrence
Y. brevifolia Engelm.
Y. elata (Engelm.) Engelm.
Y. radiosa (Engelm.) Trel.
Y. glauca Nutt.
Y. angustifolia Pursh
Y. schidigera Roezl ex Ortgies
Yucca
1175
Species
Location
Flowering
Fruit ripening
Seed dispersal
Y. brevifolia
Y. elata
Y. glauca
Y. schidigera
Mar 1Apr 1
May 15July 15
May 15June 30
Late Marearly May
July 1Aug 1
Aug 1late Sept
JulyAug
AugSept
SeptOct
Sept
Sources: Kay and others (1977), Kearney and Peebles (1969), McKelvey (1937),Webber (1953).
longitudinal sec-
(McCleary and Wagner 1973). Seeds do not require scarification for germination (CALR 1995; Went 1948). Kay and
References
Alexander RR, Pond FW. 1974. Yucca, yucca. In: Schopmeyer CS, tech.
coord. Seed of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 857858.
Arnott, H. 1962. The seed, germination, and seedlings of Yucca. Pub. Bot.
35(1). Berkeley: University of California Press. 164 p.
Bailey LH. 1947. Standard encyclopedia of horticulture. 2nd ed. New York:
Macmillan: 35293530.
Bean LJ, Saubel KS. 1972. Temalpakh (from the earth): Cahuilla Indian
knowledge and usage of plants. Morongo Indian Reservation, CA: Malki
Museum Press. 225 p.
Campbell RS, Keller JG. 1932. Growth and reproduction of Yucca elata.
Ecology 13(4): 364375.
CALR [Center for Arid Lands Restoration]. 1995. Data filed 19891995.
Twentynine Palms, CA: National Park Service, Joshua Tree National Park.
Cornett JW. 1991. The Joshua tree. Nat. Sci. Pub. 1-91. Palm Springs, CA:
Palm Springs Desert Museum.
Ellis MM. 1913. Seed production of Yucca glauca. Botanical Gazette 56:
7278.
Hester W. 1933. Yucca from seed. Flower Grower 20(9): 405.
Yucca
1177
ZamiaceaeSago-palm family
Zamia pumila L.
coontie
W. Gary Johnson
Mr. Johnson retired from the USDA Forest Services National Seed Laboratory
1178
References
Dehgan B. 1995. The few, the proud, the cycads. American Nurseryman
182(12): 7487.
Dehgan B. 1996. Permian permanence. American Nurseryman 183(2):
6681.
Dehgan B, Johnson CR. 1983. Improved seed germination of Zamia floridana (sensu lato) with H2SO4 and GA3. Scientia Horticulturae 19:
357361.
FNAEC [Flora of North America Editorial Committee]. 1993. Zamiaceae.
In:The flora of North America north of Mexico.Volume 2, Pteridophytes
and gymnosperms. New York: Oxford University Press: 347349.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. New York:
Macmillan: 1180.
Smith GS. 1978. Seed scarification to speed germination of ornamental
cycads (Zamia spp.). HortScience 13(4): 436438.
Witte TW. 1977. Storage and germination of Zamia seed. Proceedings of
the Florida State Horticultural Society 90: 8991.
Zamia
1179
RutaceaeRue family
Zanthoxylum L.
prickly-ash
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station,
Mississippi State, Mississippi
Common name(s)
Occurrence
Z. americanum Mill.
common prickly-ash,
toothache-tree,
northern prickly-ash
Hercules-club, toothachetree, southern prickly-ash,
tingle-tongue, pepperbark
espino rubial, pino macho,
aya, yellow hercules, bos
Z. clava-herculis L.
Z. martinicense (Lam.) DC.
1180
Height at
maturity (m)
8
915
2025
single
Flowering
Fruit ripening
Z. americanum
Z. clava-herculis
Z. martinicense
AprMay
AprJune
AprMay*
JuneAug
JulySept
AugSept
Species
Place
collected
Z. americanum
Z. clava-herculis
Z. martinicense
Minnesota
Mississippi
Puerto Rico
Seed
moisture
(%)
10
Cleaned seeds/weight
Range
/kg
48,10072,590
33,10037,050
Average
/lb
21,80032,900
15,00016,800
/kg
56,490
35,000
75,000
/lb
25,600
15,900
34,020
Samples
3
2
Zanthoxylum
1181
Species
Pregermination
treatment
Z. americanum
Z. clava-herculis
Stratified*
H2SO4
24
8
Sand
Blotterpaper
30
30
20
20
Days
60
45
Germination
rate
Amt
(%)
Days
20
29
20
19
Germination %
Avg
(%)
Samples
24
31
1
3
Nursery practice. Until more effective pregermination treatments are developed, fall sowing of untreated seed
immediately after collection is recommended. Germination
is epigeous (figure 4). Vegetative propagation from root cuttings and suckers is also possible (Dirr and Heuser 1987).
References
Bailey LH. 1949. Manual of cultivated plants most commonly grown in the
continental United States and Canada. Rev. ed. New York: Macmillan.
1116 p.
Bonner FT. 1974. Zanthoxylum L., prickly-ash. In: Schopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 859861.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Francis JK. 1991. Zanthoxylum martinecense (Lam.) DC., espino rubial. SOITF-SM-42. New Orleans: USDA Forest Service, Southern Forest
Experiment Station. 5 p.
Little EL Jr. 1979. Checklist of United States trees (native and naturalized).
Agric. Handbk. 541. Washington, DC: USDA Forest Service. 375 p.
Sargent CS. 1965. Manual of the trees of North America. New York: Dover.
934 p.
Vines RA. 1960. Trees, shrubs, and woody vines of the Southwest. Austin:
University of Texas Press. 1104 p.
1182
RhamnaceaeBuckthorn family
Ziziphus P. Mill.
jujube
Franklin T. Bonner
Dr. Bonner is a scientist emeritus at the USDA Forest Services Southern Research Station
Mississippi State, Mississippi
Growth habit and occurrence. There are about 100
species of this genus, which is composed of trees, shrubs,
and lianas found chiefly in the tropical and subtropical
regions of the world (Johnston 1963). There are 7 species
native to the United States and Mexico, but none of them
are of economic importance (Lyrene 1979). However, 2
exotic species, which are small deciduous trees, have been
planted in this country for fruit production, wildlife food,
and watershed protection (table 1). Common jujube
Ziziphus jujuba Mill.the most commonly planted species,
may grow to heights of 15 m at maturity (Vines 1960). This
species has been cultivated for about 4,000 years in China
and grown in this country for over 150 years (Bonner and
Rudolf 1974; Lyrene 1979; Mowry and others 1953). Both
common jujube and Christ-thornZ. spina-christi Willd.
are highly valued for fruit production and numerous agroforestry uses in Africa and Asia (Von Carlowitz 1986),
where there are many selected cultivars.
Flowering and fruiting. The perfect, yellow flowers
of common jujube appear in March to May in the United
States, and the reddish-brown fruits mature from July to
November. The fruits are globose to slender, fleshy drupes,
which turn from green to dark reddish brown at maturity. If
left on the tree, the fruits will turn black (Bailey 1939;
Vines 1960). Common jujube drupes are oblong and 2.5 to
5 cm in length. They contain a 2-celled and 2-seeded pointed stone that is deeply furrowed, reddish brown to deep
gray, oblong, and 2 to 2.5 cm long (figure 1) (Bonner and
Rudolf 1974; Mowry and others 1953). Trees bear fruit as
early as 1 to 4 years after planting (Lyrene 1979). Good
crops are borne annually, and although they are popular for
Cleaned seeds/weight
kg (lb)
1,650 (750)
1,500 (680)
No conclusive storage data are available for this genus,
but dry storage at room temperature has been successful for
Christ-thorn (Goor 1955). Because these seeds appear to
be orthodox, storage at low moisture contents at 5 C is
suggested.
Pregermination treatments. Jujube seeds are moderately dormant and require treatment for prompt germination.
Stratification recommendations for common jujube are 60 to
90 days in moist sand at 5 C (Bonner and Rudolf 1974) or
3 months warm incubation, followed by 3 months cold stratification (Dirr and Heuser 1987). Some growers recommend
scarification in sulfuric acid for 2 to 6 hours, followed by
stratification at 5 C for 60 to 90 days (Lyrene 1979). Very
Common name(s)
Occurrence
Z. jujuba Mill.
common jujube,
jujube, Chinese date
Z. spina-christi Willd.
Christ-thorn
Ziziphus
1183
Figure 1Ziziphus jujuba, common jujube: longitudinal section through 2 seeds in a stone (left), exterior view of a seed
after removal from a stone (center), exterior view of a seed (right).
Species
Z. jujuba
Z. spina-christi
Sand
30
38
Days
21
38
50
4
Germination rate
Amt
(%)
Days
65
Germination %
Avg
(%)
Samples
31
85
2
4
References
Bailey LH. 1939. The standard cyclopedia of horticulture. New York:
Macmillan. 3639 p.
Bonner FT, Rudolf PO. 1974. Ziziphus Mill., jujube. In: Schopmeyer CS, tech.
coord. Agric. Handbk. 450. Seeds of woody plants in the United States.
Washington, DC: USDA Forest Service: 862863.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant propagation: from seed to tissue culture. Athens, GA:Varsity Press. 239 p.
Gindel I. 1947. Ricerche sui semi di specie forestali indigene ed introdotte
in Palestina: primi risaltati [in Italian: Research on seeds of forest species
indigenous and introduced to Palestine: initial results]. Florence: Instituto
Selvicoltura Universita Firenze. 48 p.
Goor AY. 1955. Tree planting practices for arid areas. For. Dev. Pap. 6.
Rome: FAO. 126 p.
1184
ChenopodiaceaeGoosefoot family
Zuckia
1185
bracted utricles.
Location
Flowering
Fruit ripening
Seed dispersal
Z. brandegei
Central Utah
Uinta Basin, Utah
Sanpete Co., Utah
Mid-Junemid-Aug
MayJune
Mid-MayJuly
Jan or later
Sources: Blauer and others (1976), Goodrich and Neese (1986), Pendleton and others (1988), Plummer and others (1968).
1186
bracted utricle (top left), seed (top right), utricle (bottom left), and embryo
Normal seedlingEpigeal, with thin, 10- to 15-mmlong hypocotyls; small, narrow cotyledons; short epicotyl; and well-developed root hairs (figure 4).
Excised embryoSeeds soaked in water at 28 C for
12 hours and then drained can have their embryos
excised with sharp needles; these embryos germinate
rapidly at 15/5 or 15 C and should be evaluated for
presence of normal seedlings.
ViabilitySeeds soaked in water at 28 C for 12
hours, and then drained can be pierced through the
perisperm with a sharp probe or needle, then they are
Zuckia
1187
252349
191360
418561
190 254
527
721
16
15
seedling
1188
555769
420794
Sources: Pendleton and others (1988), Plummer and others (1968), Smith (1974).
129
332
284
732
119142
169481
263312
3721,061
Z. brandegei
Z. brandegei var. arizonica
Species
/kg
Range
Seeds (x1,000)/weight
Average
/lb
/kg
/lb
Filled seed %
Range
Average
References
Blauer AC, Plummer AP, McArthur ED, Stevens R, Giunta BC. 1976.
Characteristics and hybridization of important Intermountain shrubs: 2.
Chenopod family. Res. Pap. INT-177. Ogden, UT: USDA Forest Service,
Intermountain Forest and Range Experiment Station. 42 p.
Dorn RD. 1988. Vascular plants of Wyoming. Cheyenne, WY: Mountain
West Publishing. 340 p.
Drobnick R, Plummer AP. 1966. Progress in browse hybridization in Utah.
Proceedings of the Conference of Western State Game and Fish
Commissioners 46: 211213.
Goodrich S, Neese E. 1986. Uinta Basin flora. Ogden, UT: USDA Forest
Service, Intermountain Region, Ashley National Forest and USDI Bureau
of Land Management,Vernal District. 320 p.
Pendleton RL, Freeman DC, McArthur ED, Sanderson SC. 2000. Gender
specialization in heterodichogamous Grayia brandegei. American Journal
of Botany 87: 508516.
Plummer AP, Christensen DR, Monsen SB. 1968. Restoring big-game range
in Utah. Pub. 68-3. Salt Lake City: Utah Division of Fish and Game. 183 p.
Sanderson SC. 2000. Personal communication. Provo, UT: USDA Forest
Service, Rocky Mountain Research Station, Shrub Sciences Laboratory.
Shaw NL. 1992. Germination and seedling establishment of spiny hopsage
(Grayia spinosa [Hook.] Moq.) [PhD dissertation]. Corvallis: Oregon
State University [Dissertation Abstracts 9229768].
Smith JG. 1974. Grayia Hook. & Arn., hopsage. In: Shopmeyer CS, tech.
coord. Seeds of woody plants in the United States. Agric. Handbk. 450.
Washington, DC: USDA Forest Service: 434436.
Stevens R, Jorgensen KR. 1994. Rangeland species germination through 25
and up to 40 years of warehouse storage. In: Monsen SB, Kitchen SG,
comps. Proceedings, Ecology and Management of Annual Rangelands;
1992 May 1822; Boise, ID. Gen.Tech. Rep. INT-GTR-313. Ogden, UT:
USDA Forest Service, Intermountain Research Station: 257265.
Stevens R, Davis JN, Jorgensen KR. 1981. Viability of seed from thirty-two
shrub and forb species through fifteen years of warehouse storage.
Great Basin Naturalist 41(3): 274277.
Stutz HC. 1995. Personal communication. Provo, UT: Brigham Young
University.
Stutz HC, Sanderson SC, McArthur ED, Chu G. 1987. Chromosome races
of Grayia brandegei (Chenopodiaceae). Madroo 34: 142149.
Welsh SL. 1984. Chenopodiaceae. Great Basin Naturalist 44: 183209.
Welsh SL, Atwood ND, Goodrich S, Higgins LC, eds. 1987. A Utah flora.
Great Basin Naturalist Memoirs 9: 1894.
Zuckia
1189
1190
Part III
Appendices
Conversion Factors
Glossary
1192
1193
1202
1199
1206
1224
Appendices
1191
Metric to English
To convert from
To
Multiply by
millimeters
millimeters
millimeters
millimeters
millimeters
millimeters
millimeters
centimeters
meters
number per hectoliter
kilograms per hectoliter
grams per hectoliter
number per kilogram
number per gram
number per gram
number per square meter
number per linear meter
degrees Centigrade (C)
hectares
sixteenths of an inch
eighths of an inch
fifths of an inch
fourths of an inch
thirds of an inch
halves of an inch
inches
inches
feet
number per bushel
pounds per bushel
ounces per bushel
number per pound
number per pound
number per ounce
number per square foot
number per linear foot
degrees Fahrenheit (EF)
acres
0.6301
0.3150
0.1968
0.1574
0.1181
0.07874
0.03937
0.3937
3.281
0.3524
0.777
0.0124
0.4536
453.6
28.35
0.0929
0.3048
(1.8 H C) + 32
2.471
To convert from
To
Multiply by
sixteenths of an inch
eighths of an inch
fifths of an inch
fourths of an inch
thirds of an inch
halves of an inch
inches
feet
number per bushel
pounds per bushel
ounces per bushel
number per pound
number per pound
number per ounce
number per square foot
number per linear foot
degrees Fahrenheit (EF)
acres
millimeters
millimeters
millimeters
millimeters
millimeters
millimeters
centimeters
meters
number per hectoliter
kilograms per hectoliter
grams per hectoliter
number per kilogram
number per gram
number per gram
number per square meter
number per linear meter
degrees Centigrade (C)
hectares
1.587
3.175
5.080
6.350
8.467
12.70
2.540
0.3048
2.838
1.287
80.44
2.205
0.002205
0.03527
10.76
3.281
0.55 H (F - 32)
0.4047
English to metric
1192
Glossary
abortive imperfectly or incompletely developed, as
abortive seed.
abscission natural separation of leaves, flowers, and fruit
from plants generally associated with deterioration of a
specialized layer of thin-walled cells.
achene small, dry, indehiscent, 1-seeded fruit with seed
attached to ovary wall at only 1 point as in Cowanina and
Eriogonum; or pericarp fused with calyx tube and embryo,
completely filling the ovarian cavity as in Artemesia and
Chrysothamnus.
after-ripening biochemical or physical processes occurring in seeds, bulbs, tubers, and fruit after harvesting; often
necessary for germination or resumption of growth.
agamospermy a type of apomixis in which seeds develop
from female gametophyte tissue without fertilization as in
Amelanchier, Cotoneaster, Crataegus, and Rubus.
aggregate fruit formed from a cluster of ripened ovaries of
separate pistils of a single flower, as in Maclura, Magnolia,
and Rubus. (Compare multiple fruit and simple fruit;
synonym = syncarp).
allele an alternative form of a gene (at a given locus) differing in DNA sequence. If the array contains more than 2
genes, the genes are called multiple alleles. Multiple alleles
arise by repeated mutations of a gene, each with different
effects. No more than 2 alleles can be present in a given
(diploid organism).
ament see catkin.
anatropous having an ovule inverted at an early stage of
growth, so that the micropyle points toward the funicle, as in
Eriogonum.
angiosperm member of the group of vascular flowering
plants having seeds that develop in a carpellary ovary
(compare gymnosperm).
anthesis 1. stage of full flower expansion. 2. bursting of
pollen sacs with release of pollen.
apomixis any form of reproduction involving generative tissue, but without fertilization (compare agamospermy).
apophysis 1. an enlargement or swelling of the surface of
an organ. 2. visible portion of a scale in a closed cone.
aril exterior covering of appendage of certain seeds that
develops after fertilization as an outgrowth from the point of
attachment of the ovule as in Celastrus and Euonymus.
asexual reproduction reproduction without fertilization;
reproduction by purely vegetative means accomplished in
woody plants usually by rooting stem cuttings, air-layering,
grafting, or budding.
autogamy self-fertilization; pollination of a flower with
its own pollen; may occur in Kalmia, for example.
berry fleshy indehiscent fruit developed from a single pistil and containing 1 or more seeds as in Berberis,
Diospryros, and Ribes.
Glossary
1193
1194
fruit wall outer layer of fruits in which pericarp is not distinguishable from the seedcoat as in the achenes of
Baccharis (synonym = pericarp).
full seeds those filled with tissue having a normal appearance as distinguished from empty or partially empty seeds
(compare sound seeds).
funiculus stalk of an ovule.
fusiform radicles spindle-shaped radicles formed in
cryptogeal germination as in Araucaria.
gametophyte the haploid generation in organisms that
alternate haploid (n) and diploid (2n) generations.
geitonogamy pollination of a flower by pollen from
another flower on the same plant.
gene the smallest transmissible unit of genetic material
consistently associated with a single primary genetic effect.
genetic diversity the genetic variability within a population or a species.
genetic gain average improvement among progeny over
the mean for the parents with respect to the characteristics
used in selecting the parents.
genome a complete haploid set of chromosomes as found
in a gamete.
genotype 1. an individuals hereditary (genetic) constitution; it interacts with the environment to produce the phenotype. 2. Individual(s) characterized by a certain genetic
constitution (compare phenotype).
geographic race a race native to a geographic area.
germination resumption of active growth in an embryo
which results in its emergence from the seed and the development of structures essential to plant development.
germination percentage see germinative capacity.
germination, real percentage of sound seeds that
germinate.
germinative capacity proportion of seeds that germinate
normally during a period of time when germination is practically complete; usually expressed as a percentage
(synonym = germination percentage).
germinative energy that proportion of germination that
has occurred up to the time of peak germination, the time of
maximum germination rate, or some other preselected point.
glabrous smooth; without hairs or other projections.
glaucous having a whitish or waxy coating that give a
frosted appearance and tends to rub off.
globose approximately or completely spherical; globular.
gymnosperm members of the subdivision of plants having
seeds not enclosed in an ovary (naked seeds) borne on the
scales of a cone, on the megasporophylls of other types of
strobile, or singly with arils as in Torreya and Taxus.
(compare angiosperm).
haploid having 1 complete set of chromosomes per cell.
hardwood cutting
cuttings for vegetative propagation
that are collected during the dormant period from last
seasons growth.
Glossary
1195
1196
root growth potential test a test that estimates the physiological condition of seedlings by their ability to produce
new roots when growing in an ideal environment.
roguing systematic removal of individuals not desired for
perpetuation of a population.
samara dry, indehiscent, winged fruit; 1-seeded as in
Fraxinus and Ulmus, or sometimes with 2 samaras fused
together as in Acer.
scarification pregerminative disruption of seedcoats, usually by mechanical abrasion or by brief treatment in a strong
acid, to increase their permeability to water and gases, or to
lower their mechanical resistance.
seed matured ovule containing an embryo and nutritive
tissue enclosed in layers of protective tissue (seedcoat).
seed certification guaranty of seed identity and quality by
a recognized agency, usually evidenced by a certificate
including such information as certification category, species
and variety, year of collection, origin, purity, soundness, and
germinative capacity. See also certified seeds, selected
seeds, source-identified seeds.
seedcoat protective outer layer of a seed derived from the
integuments of the ovule. When 2 coats are present, the
thick, tough outer coat is the testa and the thin, delicate
inner coat is the tegmen.
seedcoat dormancy dormancy as a result of seedcoat conditions: impermeability to water or gas exchange or mechanical restrictions.
seed zone a designated area having defined boundaries
and altitudinal limits within which soil and climate are sufficiently uniform to indicate high probability for maintaining
a relatively uniform genetic composition as determined by
progeny-testing various seed sources.
seedlot a specified quantity of seeds having reasonably
uniform quality. It may comprise seeds from a specific location or a single seed collection zone, all collected in the
same year.
seed orchard a plantation of clones or seedlings from
selected trees for early and abundant production of seed and
to promote balanced, random mating.
seed-production area an existing stand that is usually
upgraded and opened by removal of phenotypically undesirable trees and then cultured for early and abundant seed
production.
seed source the locality where a seedlot was collected
(compare provenance).
selected seed a seedlot derived from clearly defined and
carefully chosen natural stands or plantations that conform
to specified standards and have been approved and registered by a designated authority.
serotinous 1. flowering or fruiting late in the growing season. 2. pertaining to cones that remain closed on a tree for
several months or years after maturity and are therefore late
in dispersing seeds.
Glossary
1197
1198
strobilus (plural strobili) conelike male or female fruiting bodies, composed of compact bracts or scales, of the
conifers.
style neck of the pistil which connects the stigma with the
ovary.
sub-shrub a shrub, usually small, the woody parts of
which normally die back at least partially during winter.
suture the line of dehiscence on fruits that opens naturally
to disperse seeds.
syncarp see aggregate fruit.
target seedling a seedling ideally suited to planting for a
specific management objective on a particular site.
Production of such seedlings is a major management goal in
nurseries, and requires matching genetic characteristics,
environmental factors of the intended planting site, and
cultural practices in the nursery.
tegmen the inner seedcoat, usually thin and delicate.
testa the outer seedcoat, usually thick and tough.
tree a woody perennial plant, typically large, and with a
well-defined central stem or stems with branches forming a
more or less definite crown (compare shrub).
tree percent number of trees in a nursery bed at time of
lifting expressed as a percentage of the number of viable
seeds sown.
trichome an outgrowth of the epidermis, as a hair or
scale, which is variable in shape, size, and function.
triploid having 3 times (3n) the haploid (n) number of
chromosomes.
umbel
a flat-topped inflorescence with flower stalks
arising from a common point, as in Rhamnus caroliniana;
frequently compound as in the paniculate umbels of Aralia
spinosa.
unisexual individual flowers of 1 sex, either staminate or
pistillate (compare bisexual).
unitegmic having only 1 integument as the ovules of the
composite family.
utricle a bladdery, 1-seeded, usually indehiscent fruit, consisting of an achene surrounded by bracts, as in Eurotia and
Grayia.
variety a category usually intermediate between species
(or subspecies) and forma, given a Latin name preceded by
var. based on fewer correlated characters than are used to
differentiate species or subspecies, and having a more
restricted geographical occurrence.
viability the state of being capable of germination and
subsequent growth and development of the seedling.
viscid fruits covered with sticky secretions as in
Ceanothus.
Chilopsis D. Don
Spathodea Beauv.
C
CactaceaeCactus family
Carnegiea Britt. & Rose
CaprifoliaceaeHoneysuckle family
Lonicera L.
Sambucus L.
Symphoricarpos Duham.
Viburnum L.
CasuarinaceaeCasuarina family
Casuarina Rumph. ex L.
CelastraceaeBittersweet (Staff-tree)
family
Celastrus L.
Euonymus L.
ChenopodiaceaeGoosefoot family
Atriplex L.
Grayia Hook. & Arn.
Kochia Roth
Krascheninnikovia Guldenstaedt
Sarcobatus Nees
Zuckia Standl.
ClethraceaeClethra (White alder)
family
Clethra L.
CornaceaeDogwood family
Cornus L.
CupressaceaeCypress family
Calocedrus Kurz
Chamaecyparis Spach
Cupressus L.
Juniperus L.
Platycladus Spach
Thuja L.
E
EbenaceaeEbony family
Diospyros L.
ElaeagnaceaeOleaster (Elaeagnus)
family
Elaeagnus L.
Hippophae L.
Shepherdia Nutt.
EphedraceaeEphedra (Mormon-tea)
family
Ephedra L.
EricaceaeHeath family
Arbutus L.
Arctostaphylos Adans.
Epigaea L.
Gaultheria L.
Gaylussacia Kunth
Kalmia L.
Ledum L.
Leucothoe D. Don
Oxydendrum DC.
Pieris D. Don
Rhododendron L.
Vaccinium L.
EuphorbiaceaeSpurge family
Aleurites J.R. & G. Forst.
Triadica Lour.
Vernicia Lour
F
FabaceaePea family
Acacia L.
Adenanthera L.
Albizia Durz.
Amorpha L.
Bauhinia L.
Caragana Fabr.
Ceratonia L.
Cercis L.
Cladrastis Raf.
Colutea L.
Cytisus Desf.
Delonix Raf.
Ebenopsis Britt. & Rose
Enterolobium Mart.
Gleditsia L.
Gymnocladus Lam.
Hymenaea L.
Laburnum Medik.
Lespedeza Michx.
Leucaena Benth.
Lupinus L.
Olneya Gray
Paraserianthes I. Nielsen
Pithecellobium Mart.
Prosopis L.
1199
Psorothamnus Rydb.
Pterocarpus Jacq.
Robinia L.
Senna P. Mill.
Sophora L.
Ulex L.
FagaceaeBeech family
Castanea P. Mill.
Chrysolepis Hjelmquist
Fagus L.
Lithocarpus Blume
Quercus L.
G
GarryaceaeSilk tassel family
Garrya Dougl. ex Lindl.
GinkgoaceaeGinkgo family
Ginkgo L.
GrossulariaceaeCurrant family
Ribes L.
H
HamamelidaceaeWitch-hazel
family
Hamamelis L.
Liquidambar L.
HippocastanaceaeHorsechestnut
family
Aesculus L.
HydrangeaceaeHydrangea family
Carpenteria Torr.
Philadelphus L.
HydrophyllaceaeWaterleaf family
Nama L.
J
Juglandaceae Walnut family
Carya Nutt.
Juglans L.
L
LamiaceaeMint family
Salvia L.
LauraceaeLaurel family
Lindera Thunb.
Persea P. Mill.
Sassafras Nees & Eberm.
Umbellularia (Nees) Nutt.
LythraceaeLoosestrife family
Lagerstroemia L.
1200
M
MagnoliaceaeMagnolia family
Liriodendron L.
Magnolia L.
MalvaceaeMallow family
Thespesia Soland. ex Correa
MeliaceaeMahogany family
Melia L.
Swietenia Jacq.
Toona (Endl.) Roemer
MenispermaceaeMoonseed family
Menispermum L.
MoraceaeMulberry family
Maclura Nutt.
Morus L.
MyricaceaeBayberry (Wax-myrtle)
family
Myrica L.
Morella Lour.
MyrtaceaeMyrtle family
Eucalyptus LHer.
Lophostemon Schott
N
NyssaceaeSour gum family
Nyssa L.
O
OleaceaeOlive family
Chionanthus L.
Fraxinus L.
Ligustrum L.
Menodora Bonpl.
Olea L.
Syringa L.
P
PapaveraceaePoppy family
Dendromecon Benth.
PinaceaePine family
Abies P. Mill.
Cedrus Trew
Larix P. Mill.
Picea A. Dietr.
Pinus L.
Pseudotsuga Carr.
Tsuga Carr.
PlatanaceaePlane-tree (Sycamore)
family
Platanus L.
PolygonaceaeBuckwheat family
Eriogonum Michx.
ProteaceaeProtea family
Grevillea R. Br. ex Knight
PyrolaceaeShinleaf family
Chimaphila Pursh
R
RanunculaceaeButtercup family
Clematis L.
RhamnaceaeBuckthorn family
Ceanothus L.
Frangula P. Mill.
Rhamnus L.
Ziziphus P. Mill.
RosaceaeRose family
Amelanchier Medik.
Aronia Medik.
Cercocarpus Kunth
Chamaebatia Benth.
Chamaebatiaria (Porter) Maxim.
Coleogyne Torr.
Cotoneaster Medik.
Crataegus L.
Fallugia Endl.
Heteromeles M. Roemer
Holodiscus (K. Koch) Maxim.
Malus P. Mill.
Oemleria Reichenb.
Peraphyllum Nutt.
Physocarpus (Camb.) Raf.
Prunus L.
Purshia DC. ex Poir.
Pyrus L.
Rhodotypos Sieb. & Zucc.
Rosa L.
Rubus L.
Sorbaria (Ser. ex DC.) A. Braun
Sorbus L.
Spiraea L.
RubiaceaeMadder family
Cephalanthus L.
Mitchella L.
RutaceaeRue family
Flindersia R. Br.
Phellodendron Rupr.
Ptelea L.
Zanthoxylum L.
S
SalicaceaeWillow family
Populus L.
Salix L.
SapindaceaeSoapberry family
Koelreuteria Laxm.
Sapindus L.
SapotaceaeSapodilla (Sapote) family
Sideroxylon L.
ScrophulariaceaeFigwort family
Paulownia Sieb. & Zucc.
Penstemon Schmidel
SimaroubaceaeQuassia (Ailanthus)
family
Ailanthus Desf.
SimmondsiaceaeJojoba family
Simmondsia Nutt.
Solanaceae Potato (Nightshade)
family
Lycium L.
Solanum L.
SterculiaceaeCacao (Sterculia)
family
Fremontodendron Coville
StyracaceaeStorax (Snowball)
family
Halesia Ellis ex L.
Styrax L.
U
UlmaceaeElm family
Celtis L.
Ulmus L.
V
VerbenaceaeVerbena family
Callicarpa L.
Tectona L. f.
Vitex L.
Vitaceae Grape family
Parthenocissus Planch.
Vitis L.
Z
ZamiaceaeSago-palm family
Zamia L.
ZygophyllaceaeCreosote-bush
(Caltrop) family
Larrea Cav.
T
TamaricaceaeTamarix family
Tamarix L.
TaxaceaeYew family
Taxus L.
Torreya Arn.
TaxodiaceaeRedwood family
Cryptomeria D. Don
Metasequoia Miki ex Hu &
W.C. Cheng
Sciadopitys Siebold & Zucc.
Sequoia Endl.
Sequoiadendron Buchh.
Taxodium L.C. Rich.
TheaceaeTea family
Franklinia Bartr. ex Marsh.
Gordonia Ellis
ThymelaeaceaeMezereum family
Dirca L.
TiliaceaeLinden family
Tilia L.
1201
Index of Authors
A
Alexander, Robert R.
Yucca
Anderson, Paul D.
Malus
Auger, Janene
Peraphyllum
B
Banner, Valerie A.
Cercis
Barbour, Jill R.
Aesculus
Celastrus
Cladrastis
Corylus
Elaeagnus
Hamamelis
Magnolia
Mitchella
Morus
Prunus
Ptelea
Serenoa
Tsuga
Ulmus
Barnes, R. L.
Cladrastis
Diospyros
Morus
Sabal
Serenoa
Becker, Robert
Olneya
Blazich, Frank A.
Clethra
Franklinia
Kalmia
Lagerstroemia
Leucothoe
Lonicera
Morella & Myrica
Nandina
Oxydendrum
Pieris
Rhododendron
Rhus
1202
Tilia
Vaccinium
Blum, Barton M.
Arali
Boe, Kenneth N.
Sequoia
Sequoiadendron
Bonner, Franklin T.
Chapter 1, Seed biology
Chapter 4, Storage
Chapter 6, Certification
Aleurites
Aronia
Asimina
Callicarpa
Campsis
Carya
Castanea
Catalpa
Celtis
Cephalanthus
Chamaecyparis
Chionanthus
Epigaea
Ericameria
Fagus
Fraxinus
Gaylussacia
Gledistia
Gordonia
Gymnocladus
Halesia
Ilex
Juglans
Juniperus
Liquidambar
Liriodendron
Lophostemon
Maclura
Melia
Nyssa
Ostrya
Paraserianthes
Parthenocissus
Paulownia
Persea
Pithecellobium
Platanus
Prosopis
Quercus
Sapium
Sassafras
Sideroxylon
Taxodium
Vernicia
Viburnum
Vitis
Zanthoxylum
Ziziphus
Booth, D. Terrance
Krascheninnikovia
Purshia
Brand, Gary J.
Platycladus
Thuja
Brinkman, Kenneth A.
Amelanchier
Amorpha
Cornus
Corylus
Hamamelis
Lindera
Menispermum
Mitchella
Ptelea
Sambucus
Ulmus
Brodie, Leslie Chandler
Alnus
Buckley, David S.
Dirca
Buechler, W.
Salix
Busing, Richard T.
Hippophae
Larrea
Lespedeza
Lycium
C
Connor, Kristina F.
Bauhinia
Parkinsonia
Roystonea
Sophora
Spathodea
Conard Susan G.
Ceanothus
Crossley, John A.
Malus
Solanum
D
Davis, Tim D.
Lupinus
Debell, Dean S.
Alnus
Dietz, Donald R.
Caragana
Douglas, D.A.
Salix
Dubois, Jean-Jacques B.
Lagerstroemia
Lonicera
E
Eddleman, Lee E.
Sarcobatus
Tetradymia
Edwards, D. George W.
Abies
Erdmann, G.G.
Mitchella
F
Francis, John K.
Acacia
Araucaria
Cryptomeria
Enterolobium
Pterocarpus
Roystonea
Spathodea
Swietenia
Tectona
Thespesia
Toona
G
Gabriel, W. J.
Acer
Gill, John D.
Aronia
Chionanthus
Parthenocissus
Physocarpus
Viburnum
Graney, David L.
Catalpa
Griffin, Jason T.
Clethra
Franklinia
Myrica
Vaccinium
Grisez, Ted J.
Prunus
H
Haferkamp, Marshall R.
Grayia
Harrington, Constance A.
Alnus
Huffman, David W.
Gaultheria
Hummer, Kim E.
Pyrus
Hurd, Emerenciana G.
Chamaebatiaria
Grayia
Holodiscus
Philadelphus
J
Jenkinson, James L.
Pinus
Johnson, LeRoy C.
Metasequoia
Johnson, W. Gary
Dendromecon
Diospyros
Sambucus
Styrax
Zamia
Jull, Laura G.
Nandina
K
Karrfalt, Robert P.
Chapter 3, Harvesting &
Conditioning
Chapter 5, Testing
Baccharis
Celastrus
Prunus
Robinia
Sabal
Tsuga
Kitchen, Stanley G.
Cercocarpus
Kochia
Krugman, Stanley L.
Eucalyptus
Menodora
Pinus
Sequoiadendron
Washingtonia
L
Landis, Thomas D.
Chapter 7, Nurseries
Lantz, Clark W.
Chapter 2, Genetics
Lasseigne, F. Todd
Crataegus
Leak, William B.
Fagus
Ostrya
Leiser, Andrew T.
Nama
Little, Silas
Ailanthus
M
McDaniel, Kirk C.
Gutierrezia
McDonald, Philip M.
Arbutus
Chrysolepis
Lithocarpus
Mackay, Wayne A.
Lupinus
Magill, Arthur W.
Chamaebatia
Chilopsis
Mangold, Robert D.
Chapter 6, Certification
Markin, George P.
Ulex
Martin, George C.
Olea
Martineau, David
Amorpha
Index of Authors
1203
Matula, Colleen A.
Dirca
Meyer, Susan E.
Arctostaphylos
Artemisia
Atriplex
Carnegiea
Chrysothamnus
Cytisus
Ephedra
Eriogonum
Fallugia
Fremontodendron
Heteromeles
Penstemon
Purshia
Rosa
Salvia
Simmondsia
Michler, Charles H.
Koelreuteria
Mikowski, Daniel A.
Ligustrum
Miller, Carol
Ambrosia
Chilopsis
Encelia
Parkinsonia
Psorothamnus
Minore, Don
Berberis
Chimaphila
Monsen, Stephen B.
Kochia
N
Nauertz, Elizabeth Ann
Dirca
Neal, Donald L.
Carpenteria
Dendromecon
Nord, Eamor C.
Nama
O
Olson, David F., Jr.
Baccharis
Casuarina
Cladrastis
Diospyros
Elaeagnus
1204
Robinia
Sabal
Serenoa
Owston, Peyton W.
Pseudotsuga
Rhodotypos
Sciadopitys
Sorbaria
P
Parrotta, John A.
Albizia
Casuarina
Flindersia
Grevillea
Leucaena
Paraserianthes
Pendleton, Burton K.
Coleogyne
Pendleton, Rosemary
Zuckia
Petteys, E.
Casuarina
Lophostemon
Pfister, Robert D.
Ribes
Phipps, Howard M.
Lindera
Menispermum
Pijut, Paula M.
Carpinus
Cedrus
Colutea
Cotinus
Kalopanax
Laburnum
Pogge, Franz L.
Aronia
Chionanthus
Parthenocissus
Physocarpus
Viburnum
Pond, Floyd W.
Yucca
Postman, Joseph D.
Pyrus
R
Ratliff, Raymond D.
Ericameria
Lupinus
Read, Ralph A.
Morus
Phellodendron
Sapindus
Riemenschneider, Don E.
Lupinus
Rodgers, Jane E.
Ambrosia
Chilopsis
Encelia
Parkinsonia
Psorothamnus
Senna
Yucca
Roh, Mark S.
Styrax
Rowe, D. Bradley
Rhododendron
Rhus
Tilia
Rudolf, Paul O.
Aesculus
Berberis
Clematis
Euonymus
Koelreuteria
Lycium
Rhodotypos
Sciadopitys
Sorbaria
Syringa
Ruth, Robert H.
Tsuga
S
Safford, Lawerence O.
Picea
Schlesinger, R. C.
Ptelea
Schmidtling, R. C.
Sideroxylon
Schopmeyer, C. S.
Alnus
Nemopanthus
Platycladus
Thuja
Vitex
Schubert, T. H.
Tectona
Shaw, Nancy L.
Chamaebatiaria
Cotoneaster
Grayia
Holodiscus
Philadelphus
Purshia
Syringa
Zuckia
Shearer, Raymond C.
Larix
Shepperd, Wayne D.
Ceratonia
Garrya
Ginkgo
Tamarix
Slabaugh, Paul E.
Caragana
Cotoneaster
Hippophae
Syringa
Sloan, John P.
Metasequoia
Ribes
Sequoia
Smith, Justin G.
Peraphyllum
Starrett, Mark C.
Kalmia
Leucothoe
Oxydendrum
Pieris
Stein, William I.
Calocedrus
Cercis
Gaultheria
Ligustrum
Oemleria
Pseudotsuga
Sorbus
Torreya
Umbellularia
Stickney, Peter F.
Holodiscus
Philadelphus
Spiraea
Strong, Terry F.
Acer
Amelanchier
T
Tappeiner, John C.
Rubus
V
Vance, Nan C.
Taxus
Vankus, Victor
Cornus
Lindera
Vogel, Willis G.
Lespedeza
Vozzo, Jack A.
Adenanthera
Delonix
Hymenaea
W
Walker, Scott C.
Shepherdia
Symphoricarpos
Walters, Gerald A.
Albizia
Araucaria
Cryptomeria
Toona
Wendel, G. W.
Celastrus
Whitesell, Craig D.
Acacia
Eucalyptus
Leucaena
Wick, Herbert L.
Albizia
Flindersia
Wong, Wesley H. C., Jr.
Grevillea
Wood, Ballard
Gutierrezia
Wurtz, Tricia L.
Ledum
Wyckoff, Gary W.
Populus
Y
Youngblood, Andrew
Frangula
Physocarpus
Picea
Rhamnus
Z
Zasada, John C.
Acer
Ailanthus
Amorpha
Aralia
Clematis
Dirca
Euonymus
Gaultheria
Menodora
Nemopanthus
Phellodendron
Populus
Rubus
Salix
Sapindus
Solanum
Spiraea
Vitex
Index of Authors
1205
thinleaf
wavyleaf
western
white
algarrobo, Hymenaea
aliso, Alnus
aliso, Platanus
Allegheny chinkapin, Castanea
almond, Prunus
desert
alpine goldenchain, Laburnum
American beautyberry, Callicarpa
amorpha, Amorpha
California
Amur corktree, Phellodendron
angelica-tree, Aralia
Ao-todomatsu, Abies
Aomori-todo-matsu, Abies
aotodo, Abies
Apache-plume, Fallugia
apes earring, Ebenopsis
apple, Malus
American crab
Biltmore crab
common
Dunbar crab
European crab
Great Lakes crab
Iowa crab
Japanese flowering crab
midwest crab
narrow-leaf crab
Oregon crab
Pacific crab
paradise
prairie crab
red Siberian crab
Sargent
Siberian crab
southern crab
western crab
wild crab
wild sweet crab
applebush, Grayia
applebush, Zuckia
apricot, Prunus
desert
aralia, Aralia
bristly
araucaria, Araucaria
columnar
Arauco-pine, Araucaria
arborvitae, Thuja
eastern
giant
Japanese
Korean
swamp-cedar
arbutus, Arbutus
arctic bramble, Rubus
aroma, Acacia
arrowwood, Viburnum
arrowwood
downy
mapleleaf
roughish
smooth
southern
ash, Fraxinus
Arizona
basket
Biltmore
Biltmore white
black
blue
brown
Carolina
Darlington
desert
European
flowering
foothill
green
hoop
leatherleaf
Modesto
Oregon
pop
pumpkin
red
Shamel
smooth
swamp
Toumey
tropical
two-petal
velvet
water
white
Asiatic sweet pepperbush, Clethra
aspen, Populus
bigtooth
European
golden
Japanese
largetooth
mountain
quaking
Siebold
trembling
Canada
double fir
healing
western
white
barberry, Berberis
black
boxleaf
common
cutleaf
Darwin
European
Japanese
Julian
Korean
paleleaf
Sargent
threespine
Verna
warty
wildfire
wintergreen
barberry, Mahonia
Cascades
creeping
hollyleaf
Nevin
red
basswood, Tilia
Carolina
Mexican
white
batai, Paraserianthes
bauhinia, Bauhinia
butterfly
petite flambouyant
pink
purple
bay, Gordonia
bay, Umbellularia
bayberry, Myrica/Morella
California
northern
Pacific
southern
bead tree, Melia
bearberry, Arctostaphylos
bearbrush, Garrya
bearclover, Chamaebatia
beardtongue, Penstemon
broadleaf
gilia
Leonards
littlecup
petiole
beargrass, Yucca
bearmat, Chamaebatia
southern
bee-tree, Tilia
beech, Fagus
American
European
bellota, Quercus
Benjamin bush, Lindera
benzoin tree, Styrax
big-leaved ivy, Kalmia
bigcone-spruce, Pseudotsuga
bigtree, Sequoiadendron
bilsted, Liquidambar
biota, Platycladus
birch, Betula
Alaska
Asian white
black
bog
canoe
cherry
Dahurian
downy
dwarf
dwarf white
Erman
European white
glandulose
gray
Japanese white
monarch
Murray
northern
northwestern paper
paper
river
roundleaf
silver
swamp
sweet
water
white
wire
yellow
bitter nightshade, Solanum
bitterbrush, Purshia
antelope
desert
bitternut, Carya
bittersweet, Celastrus
American
climbing
shrubby
black alder, Ilex
black greasewood, Sarcobatus
black laurel, Gordonia
black thornberry, Crataegus
black twinberry, Lonicera
1207
black-snap, Gaylussacia
blackberry, Rubus
Allegheny
cutleaf
evergreen
Himalaya
mountain
Pacific
running
smooth
sow-teat
thornless
trailing
blackbrush, Cercocarpus
blackbrush, Coleogyne
blackgum, Nyssa
blackhaw, Viburnum
rusty
southern
blackthorn, Prunus
blackwood, Acacia
bladder-senna, Colutea
common
blue beech, Carpinus
blue blossom, Ceanothus
blue bush, Ceanothus
blueberry, Vaccinium
American
Canadian
highbush
late sweet
low sweet
lowbush
rabbiteye
smallflower
sour-top
swamp
velvet-leaf
bluegum, Eucalyptus
bluehaw, Viburnum
bodark, Maclura
bog-myrtle, Myrica/Morella
bois-darc, Maclura
bongay, Aesculus
Boston ivy, Parthenocissus
bos, Zanthoxylum
bottlebrush
California
dwarf
fetid
Georgia
Ohio
painted
red
scarlet
sweet
1208
Texas
yellow
woolly
bowwood, Maclura
boxelder, Acer
boxthorn, Lycium
Brayley flindersia, Flindersia
Brazilian-pine, Araucaria
Brigham tea, Ephedra
Brisbane-box, Lophostemon
brittlebrush, Encelia
California
green
broad-leaved laurel, Kalmia
brushbox, Lophostemon
buckbrush, Ceanothus
buckbrush, Symphoricarpos
buckeye, Aesculus
big
buckthorn, Frangula
beechleaf
birchleaf
California
Carolina
glossy
Modoc
Nevada
obovate
obtuse
Pursh
red
Sierra
yellow
Yosemite
buckthorn, Rhamnus
alder
Chinese
common
Dahurian
European
Japanese
lanceleaf
redberry
sawleaf
sharp-tooth
Smith
buckthorn, Sideroxylon
woolly
buckwheat, Eriogonum
crisp-leaf
mat
parsnipflower
roundleaf
buckwheatbrush, Eriogonum
California
flat-top
Heerman
lace
Mojave
pretty
budsage, Artemisia
buffaloberry, Shepherdia
Canadian
roundleaf
russet
silver
thornless
bull bay, Magnolia
bullberry, Shepherdia
bullnut, Carya
bunchberry, Cornus
bunya-bunya, Araucaria
bunya-pine, Araucaria
Burma-cedar, Toona
Burmese rosewood, Pterocarpus
burning-bush, Euonymus
burrobush, Ambrosia
white
burroweed, Ambrosia
bursage, Ambrosia
white
bursting-heart, Euonymus
bush rockspirea, Holodiscus
bush-anemone, Carpenteria
bushpoppy, Dendromecon
island
stiff
butternut, Juglans
buttonball-tree, Platanus
buttonbush, Cephalanthus
common
buttonwood, Platanus
C
calico-bush, Kalmia
California boxelder, Acer
California fan-palm, Washingtonia
California Washington-palm,
Washingtonia
California washingtonia,
Washingtonia
California-holly, Heteromeles
California-laurel, Umbellularia
California-nutmeg, Torreya
California-olive, Umbellularia
California-palm, Washingtonia
calmoun, Kalmia
candelabra tree, Araucaria
candleberry, Myrica/Morella
candleberry-myrtle, Myrica/Morella
candlenut-tree, Aleurites
canoe-cedar, Thuja
caragana, Caragana
carob, Ceratonia
bitter
black
common choke
downy
European bird
evergreen
fire
hollyleaf
laurel
mahaleb
Manchu
mazzard
Nanking
narrowleaf
pie
pin
perfumed
Rocky Mountain
rum
sand
sour
St. Lucie
sweet
western sand
wild
wild black
wild red
chestnut, Aesculus
chestnut, Castanea
American
Chinese
European
Japanese
Spanish
Chilean-pine, Araucaria
Chinese arborvitae, Platycladus
Chinese date, Ziziphus
Chinese tallowtree, Triadica
chittamwood, Sideroxylon
checkerberry, Gaultheria
cherrioni, Sapindus
cherry-laurel, Prunus
Carolina
Chimaphila, Chimaphila
Japanese
China-tree, Koelreuteria
chinaberry, Melia
chinatree, Melia
chinquapin, Chrysolepis
bush
giant
golden
chittam, Frangula
chittamwood, Sideroxylon
chokeberry, Aronia
black
hybrid
purple
red
Christmas-berry, Ilex
Christ-thorn, Ziziphus
Christmasberry, Heteromeles
cigar-tree, Catalpa
cinnamon-bark clethra, Clethra
clematis, Clematis
Drummond
eastern
Italian
mountain
plume
purple
rock
western
clethra, Clethra
cinnamon-bark
Japanese
cliffrose, Purshia
cloudberry, Rubus
coastal sweetpepperbush, Clethra
coffeeberry, Frangula
coffeetree, Frangula
common gorse, Ulex
common moonseed, Menispermum
common seabuckthorn, Hippophae
common trumpetcreeper, Campsis
comptie, Zamia
Cook-pine, Araucaria
coontie, Zamia
copaltree, Ailanthus
coralberry, Symphoricarpos
corkbush, Euonymus
cornelian-cherry, Cornus
cotoneaster, Cotoneaster
black
cranberry
darkseed
European
hedge
Peking
rock
rock spray
cotton-gum, Nyssa
cottonwood, Populus
Arizona
balsam
black
eastern
Fremont
lanceleaf
Macdougal
mountain
narrowleaf
plains
Rio Grande
1209
river
smooth-bark
swamp
Texas
valley
Wislizenus
courbaril, Hymenaea
cowberry, Vaccinium
cowitch vine, Campsis
coyotebrush, Baccharis
cranberry, Vaccinium
American
large
mountain
small
cranberrybush, Viburnum
European
crape-myrtle, Lagerstroemia
crape-myrtle
Queens
crapemyrtle, Lagerstroemia
creambush, Holodiscus
creambush rockspirea, Holodiscus
creeper, Parthenocissus
Japanese
Virginia
creeping snowberry, Gaultheria
creeping pearlberry, Gaultheria
creosotebush, Larrea
crocus, Epigaea
cryptomeria, Cryptomeria
Japanese
cuapinol, Hymenaea
cucumbertree, Magnolia
Cunningham beefwood, Casuarina
curlleaf mahogany, Cercocarpus
currant, Ribes
alpine
alpine prickly
American black
blood
buffalo
clove
flowering
golden
gooseberry
Hudson Bay
northern black
Oregon
prickly
red flowering
Sierra
slender golden
squaw
sticky
swamp black
wax
1210
wild black
winter
custard-apple, Asimina
cypress, Chamaecyparis
false
Lawson
Sitka
yellow
cypress, Cupressus
Arizona
Arizona smooth
Baker
Cuyamaca
Gowen
Guadalupe
Himalayan
Italian
MacNab
Mediterranean
Mendocino
Mexican
Modoc
Monterey
Piute
pygmy
Santa Cruz
Sargent
Siskiyou
spreading Italian
tecate
cypress, Taxodium
common bald
gulf
red
tidewater red
white
yellow
D
Dake-momi, Abies
dalea, Psorothamnus
California
Fremont
Johnson
Mojave
Nevada
Saunder
Schott
damson, Prunus
date-plum, Diospyros
dawn-redwood, Metasequoia
deer brush, Ceanothus
deerbrush, Cercocarpus
desert catalpa, Chilopsis
desert ironwood, Olneya
desert mahogany, Cercocarpus
desert sweet, Chamaebatiaria
Douglas-spruce, Pseudotsuga
drooping leucothoe, Leucothoe
dwarf cornel, Cornus
dwarf-elder, Aralia
dyebush, Psorothamnus
dyeweed, Psorothamnus
E
ear-leaf umbrellatree, Magnolia
earpod-tree, Enterolobium
eastern leatherwood, Dirca
eastern wahoo, Euonymus
ebony blackbead, Ebenopsis
eglantine, Rosa
Egyptian thorn, Acacia
elaeagnus, Elaeagnus
autumn
elder, Sambucus
American
blackberry
blue
blueberried
blueberry
common
red
redberried
scarlet
sweet
elderberry, Sambucus
blue
elm, Ulmus
American
basket
cedar
Chinese
cork
dwarf Asiatic
English
European white
field
grey
Japanese
lacebark
leatherleaf
red
rock
Russian
Scotch
Scots
September
Siberian
slippery
smoothleaf
soft
southern rock
spreading
water
winged
white
Wych
emajagilla, Thespesia
empress tree, Paulownia
encelia, Encelia
rayless
Virgin River
encina, Quercus
encino, Quercus
ephedra, Ephedra
gray
green
Torrey
espino rubial, Zanthoxylum
eucalyptus, Eucalyptus
alpine-ash
beakpod
blackbutt
bluegum
brown-barrel
cuttail
dalrymple
delegate
desert
gray ironbark
lemon
lemon-gum
long-beak
manna
messmate stringybark
moitch
mountain-ash
mountain-gum
mulga ironbark
redironbark
ribbon
river redgum
robusta
rosegum
saligna
shining
Sidney bluegum
swamp-gum giant
tallowwood
Tasmanian blue
tooler
euonymus, Euonymus
brook
European
Maack
running
warty-bark
winged
winterberry
European bittersweet, Solanum
F
false indigo, Amorpha
false spirea, Sorbaria
Ural
falsewillow, Baccharis
Bigelows
broombrush
Encinitis
Harvards
prairie
saltwater
Santo Domingo
false-willow, Chilopsis
farkleberry, Vaccinium
fern-bush, Chamaebatiaria
fernbush, Chamaebatiaria
fetterbush, Leucothoe
fetterbush, Pieris
feverbush, Lindera
filbert, Corylus
American
beaked
California
common
European
fir, Abies
Abete delle Nebrodi
alamo de le sierra
Algerian
Algerian silver
alpine
amabilis
Amur
Ao-todomatsu
Aomori-todo-matsu
aotodo
akatodo
Arizona
balsam
blister
bristlecone
California red
California white
Cascades
Caucasian
Cephalonian
Chinese silver
Cilician
Colorado white
common silver
concolor
corkbark
Crimean
dake-momi
eastern
European silver
feather cone
1211
flaky
Fraser
golden
grand
great silver
Grecian
Greek silver
Guatemalan
Hinggan
Japanese
Japanese silver
Khingan
Korean
linpi lengshan
lovely
Low silver
Low white
lowland white
magnificent
Manchurian
Maries
Mayr Sakhalin
Mexican silver
Min
Min-kiang
momi
Mt. Enos
needle
Nikko
Nikko-momi
noble
noble red
Nordmann
Oregon
oyamel
O-shirabiso
Pacific silver
Pacific white
Pindrow
pio real blanco
pitch silver
real blanco de la sierras
red
red bark
Rocky Mountain alpine
Rocky Mountain subalpine
Rocky Mountain white
sacred
sacred Mexican silver
Sakhalin
Santa Lucia
sapin concolore
sapin du Vancouver
sapin gracieux
sapin grandissime
Shasta
Shasta red
1212
shirabe
shirabiso
Siberian
Siberian silver
Siberian white
Sicilian
Sierra white
silver
silvertip
Sino-Korean
southern balsam
Spanish
Spanish silver
subalpine
todo-matsu
Todomatsu
Turkey
urajiro-momi
Veitch
Veitch silver
west Himalayan
west Himalayan silver
western white
white
yellow-fruited
fir pine, Abies
firecracker plant, Aesculus
five-stamen tamarisk, Tamarix
flamboyan, Delonix
flametree, Delonix
flannelbush, Fremontodendron
California
Mexican
flooded-gum, Eucalyptus
Florida arrowroot, Zamia
Florida pinxter, Rhododendron
Florida-nutmeg, Torreya
flowering-ash, Chionanthus
flowering-willow, Chilopsis
fountain tree, Spathodea
fox grape, Vitis
Northern
foxberry, Vaccinium
fragrant false indigo, Amorpha
Franklin tree, Franklinia
franklinia, Franklinia
fremontia, Fremontodendron
California
eldorado
Mexican
French-mulbery, Callicarpa
fresno, Fraxinus
frijolito, Sophora
fringed sage, Artemisia
fringed spruce, Abies
G
gallberry, Ilex
garland-tree, Malus
gean, Prunus
Gharab-Palk-Saf-Saf, Populus
giant cactus, Carnegiea
giant sequoia, Sequoiadendron
giant-cedar, Thuja
ginkgo, Ginkgo
globe-flowers, Cephalanthus
goatnut, Simmondsia
gobernadora, Larrea
goddess-of-mercy-fir, Cryptomeria
gold-and-silver-flower, Lonicera
goldenchain tree, Laburnum
goldenhills, Encelia
gooseberry, Ribes
Appalachian
eastern prickly
Idaho
inland balck
Missouri
mountain
pasture
roundleaf
Sierra
swamp
white-stem
gordonia, Gordonia
grandfather-graybeard, Chionanthus
grape, Vitis
plum
swamp
gravel plant, Epigaea
gravel weed, Epigaea
Grays saltbush, Grayia
graybeard, Clematis
grayia, Grayia
greasewood, Larrea
greasewood, Sarcobatus
ground hemlock, Taxus
ground-laurel, Epigaea
guamchil, Pithecellobium
guanacaste, Enterolobium
Guelder rose, Viburnum
gueles noires, Aronia
gum arabic tree, Acacia
gum bumelia, Sideroxylon
gum elastic, Sideroxylon
H
hackberry, Celtis
common
netleaf
northern
sugar
western
hackmatack, Larix
hackmatack, Populus
hardhack, Holodiscus
hardhack, Spiraea
Harford tree-poppy, Dendromecon
haw, Crataegus
apple
blue
dwarf
green
parsley
red
summer
yellow
hawthorn, Crataegus
Allegheny
anomalous
apple
apple haw
apple-leaf
Arnold
barberry
beautiful
bigtree
black
blueberry
Brainerd
broadleaf
cerro
chocolate
cockspur
Columbia
common
dotted
Douglas
downy
English
English midland
English woodland
entangled
fanleaf
fireberry
flat-topped
fleshy
frosted
glossy
golden-fruit
green
Gregg
Harbison
Kansas
large-fruited
littlehip
longspine
may
mountain
one-flowered
oneseed
Ontario
parsley
pasture
pear
Pensacola
Piper
plumleaf
Reverchon
riverflat
roundleaf
sandhill
scarlet
shining
Siberian
single-seed
small-fruited
southern
succulent
sugar
summer
sunny
tall
Texas
thicket
three-flower
Tracy
Virginia
Washington
waxy-fruited
weeping
western black
willow
yellow
hazel, Corylus
American
beaked
California
European
he-balsam, Picea
hearts-a-busting, Euonymus
heavenly-bamboo, Nandina
hedge, Maclura
hediondilla, Larrea
hemlock, Tsuga
black
Canada
Carolina
eastern
mountain
Pacific
western
hemptree, Vitex
Hercules-club, Aralia
Hercules-club, Zanthoxylum
hickory, Carya
big shagbark
bigleaf shagbark
bitter water
bitternut
mockernut
nutmeg
pale
pallid
pignut
sand
scalybark
shagbark
shellbark
swamp
water
white
whiteheart
highbush-cranberry, Viburnum
highland doghobble, Leucothoe
hobblebush, Viburnum
hog-apple, Crataegus
hoghaw, Crataegus
hognut, Carya
holly, Ilex
American
deciduous
English
evergreen
mountain
swamp
white
holly-bay, Gordonia
hollywood, Heteromeles
honey-balls, Cephalanthus
honeylocust, Gleditsia
swamp
Texas
honeysuckle, Lonicera
Amur
Arizona
bearberry
Belle
blueleaf
California
chaparral
coral
coralline
dwarf
Etruscan
European
European fly
fly
grape
hairy
Italian
Japanese
limber
Manchurian
1213
Morrow
mountain
mountain fly
orange
purple flower
southern
Standish
swamp fly
sweetberry
Tatarian
trumpet
twinberry
Utah
western white
whitebell
winter
woodbine
yellow
honeysuckle, Rhododendron
swamp
hoop-pine, Araucaria
hophornbeam, Ostrya
American
eastern
hopsage, grayia
hoptree, Ptelea
common
woolly common
hornbeam, Carpinus
American
European
heartleaf
Japanese
oriental
hornbeam, Ostrya
hornbrush, Ceanothus
horse-apple, Maclura
horsebean, Parkinsonia
horsebrush, Tetradymia
catclaw
common
cotton
cottonthorn
dune
four-part
gray
hairy
littleleaf
longspine
Mojave
Nuttall
shortspine
smooth
spiny
spineless
striped
thorny
1214
threadleaf
horsebush, Grayia
horsechestnut, Aesculus
American
Himalayan
horsetail beefwood, Casuarina
huckleberry, Gaylussacia
black
highbush
huckleberry, Vaccinium
California
evergreen
shot
velvetleaf
huisache, Acacia
I
incense-cedar, Calocedrus
California
incienso, Encelia
Indian arrow-wood, Holodiscus
Indian arrow-wood, Philadelphus
Indian currant, Symphoricarpos
Indian lilac, Melia
Indian peach, Oemleria
Indian plum, Oemleria
Indian soap-plant, Sapindas
Indian-bean, Catalpa
Indian-walnut, Aleurites
indigobush, Amorpha
dwarf
indigobush, Psorothamnus
Mojave
inkberry, Ilex
inkberry, Lonicera
ironbark, Eucalyptus
ironwood, Carpinus
ironwood, Casuarina
ironwood, Olneya
ironwood, Ostrya
island myrtle, Ceanothus
islay, Prunus
Italian woodbine, Lonicera
ivy, Kalmia
ivy-bush, Kalmia
J
jaboncillo, Sapindus
jano, Chilopsis
Japanese cornelian-cherry, Cornus
Japanese snowdrop tree, Styrax
Japanese-cedar, Cryptomeria
Jersey-tea, Ceanothus
Jerusalem-thorn, Parkinsonia
jetbead, Rhodotypos
Jim brush, Ceanothus
jimbrush, Ceanothus
jojoba, Simmondsia
Joshua tree, Yucca
Joves fruit, Lindera
Judas-tree, Cercis
juneberry, Amelanchier
jujube, Ziziphus
common
jumbie-bead, Adenanthera
juneberry, Amelanchier
juniper, Juniperus
alligator
Ashes
bigberry
California
checkered-bark
cherrystone
common
dwarf
Mexican
oneseed
Pinchot
prostrate
red
redberry
river
Rocky Mountain
Sierra
Utah
west Texas
western
jutaby, Hymenaea
K
kaki, Diospyros
keg fir, Diospyros
keminyan, Styrax
Kentucky coffeetree, Gymnocladus
Kew-tree, Ginkgo
kiawe, Prosopis
kingnut, Carya
kinnickinnick, Arctostaphylos
kinnikinnik, Cornus
Klinki-pine, Araucaria
koa, Acacia
koa haole, Leucaena
kochia, Kochia
forage
prostrate
kukui, Aleurites
L
Labrador-tea, Ledum
bog
marsh
western
laburnum, Laburnum
common
Scotch
Waterer
lacewood, Grevillea
lama, Aleurites
larch, Larix
alpine
American
Dahurian
eastern
European
Japanese
Montana
mountain
Russian
Siberian
subalpine
western
large-leaf cucumbertree, Magnolia
laurel, Umbellularia
laurel-leaves, Kalmia
laurel-sumac, Rhus
leadplant, Amorpha
leadtree, Leucaena
lemon-gum, Eucalyptus
lemonade berry, Rhus
lensscale, Atriplex
lentisco, Rhus
lespedeza, Lespedeza
bicolor
leafy
shrub
Thungerg
leucaena, Leucaena
leverwood, Ostrya
life-of-man, Aralia
lilac, Syringa
Amur
common
late
Manchurian
Persian
villous
lilac chastetree, Vitex
lily-of-the-valley tree, Oxydendrum
lime, Tilia
American
Caucasian
large-leaved
pendent white
small-leaved
weeping
linden, Tilia
American
bigleaf
common
Crimean
European
European white
largeleaf
littleleaf
pendent silver
silver
lingonberry, Vaccinium
linpi lengshan, Abies
little princes-pine, Chimaphila
Lobb fiddleleaf, Nama
loblolly-bay, Gordonia
locust, Ceratonia
locust, Robinia
black
bristly
clammy
Hartweg
Holdt
Kelsey
Margarett
mossy
New Mexican
Rusby
longleaf ironwood, Casuarina
lost camellia, Franklinia
lost gordonia, Franklinia
lumbang, Aleurites
lupine, Lupinus
Inyo bush
longleaf bush
Pauma
silver
Sims bush
whiteface
M
Madras thorn, Pithecellobium
madrone, Arbutus
madroo, Arbutus
maga, Thespesia
magnolia, Magnolia
Ashe
bigleaf
cucumber
ear-leaf(ed)
evergreen
Fraser
greatleaf(ed)
mountain
Puerto Rico
pyramid
shining
southern
sweetbay
umbrella
yellow cucumber
mahala mat, Ceanothus
mahaleb, Prunus
mahogany, Swietenia
bigleaf
Honduras
hybrid
littleleaf
Pacific coast
West Indies
mahonia, Mahonia
Chinese
cluster
Fremont
Japanese
leatherleaf
maibao, Alnus
maidenhair-tree, Ginkgo
mamane, Sophora
mangium, Acacia
manzanita, Arctostaphylos
bigberry
Eastwood
greenleaf
hoary
Mexican
pointleaf
Pringle
rosybract
maple, Acer
Amur
ashleaf
bigleaf
bigtooth
broadleaf
dwarf
hard
Japanese
mountain
Norway
Oregon
paperbark
planetree
red
river
rock
Rocky Mountain
Siberian
silver
soft
striped
sugar
swamp
sycamore
vine
maple-silkwood, Flindersia
matrimony vine, Lycium
Chinese
may, Crataegus
mayday tree, Prunus
Index of Common Names
1215
mayflower, Epigaea
mayhaw, Crataegus
eastern
rufous
western
meadow-fern, Myrica/Morella
meadowsweet, Spiraea
mescalbean, Sophora
mesquite, Prosopis
honey
mesquite
screwbean
velvet
milo, Thespesia
mimosa tree, Albizia
mock locust, Amorpha
mock orange, Philadelphus
desert
Lewis
little-leaf
littleleaf
wild
mock-orange, Styrax
mockernut, Carya
molecule model plant, Eriogonum
molly, Kochia
gray
green
Molucca-albizia, Paraserianthes
momi, Abies
monkey-puzzle, Araucaria
monkey-puzzle-tree, Araucaria
monkeypod, Albizia
monkeypod, Pithecellobium
monkspeppertree, Vitex
moosewood, Acer
moosewood, Dirca
moosewood, Viburnum
Moreton-Bay-pine, Araucaria
Mormon-tea, Ephedra
gray
green
Nevada
Torrey
mountain andromeda, Pieris
mountain balm, Ceanothus
mountain cedar, Juniperus
mountain fetterbush, Pieris
mountain ivy, Kalmia
mountain pieris, Pieris
mountain sweeetpepperbush, Clethra
mountain whitethorn, Ceanothus
mountain-ash, Sorbus
American
California
European
Greene
1216
large-fruited
Pacific
showy
Sitka mountain
small-fruited
western
mountain-ebony, Bauhinia
mountain-holly, Nemopanthus
mountain-laurel, Kalmia
mountain-mahogany, Cercocarpus
alderleaf
birchleaf
curlleaf
true
mountain-misery, Chamaebatia
San Diego
Sierran
mountain-pink, Epigaea
mountain-spray, Holodiscus
moxieplum, Gaultheria
mulberry, Morus
black
littleleaf
mountain
Persian
red
Russian
silkworm
Texas
white
musclewood, Carpinus
myrtlewood, Umbellularia
N
namboca, Juglans
nandina, Nandina
nangoon berry, Rubus
nannyberry, Shepherdia
nannyberry, Viburnum
nanten, Nandina
narra, Pterocarpus
narrow-leafed oleaster, Elaeagnus
Nevada joint-fir, Ephedra
New-Jersey-tea, Ceanothus
Nikko-momi, Abies
ninebark, Physocarpus
Amur
Atlantic
common
dwarf
mallow
mountain
Pacific
nogal, Juglans
nogal silvestre, Juglans
nogalito, Juglans
Nootka yellow-cypress,
Chamaecyparis
Norfolk-Island-pine, Araucaria
northern muscadine, Vitis
nuez, Aleurites
nuez de India, Aleurites
nuez encarcelada, Carya
O
O-shirabiso, Abies
oak, Querecus
Ajo
Arizona
Arizona white
barren
basket
bastard
bear
black
blackjack
blue
bluejack
bluff
bottomland red
Brewer
bur
California black
California blue
California live
California scrub
Californian white
canyon
canyon live
Catesby
cherrybark
chestnut
chinkapin
coast live
common red
cork
cow
Darlington
Durand
Durand white
durmast
eastern red
Elliot
Emory
English
European turkey
fork-leaf white
Gambel
Garry
goldcup
gray
highland live
Hill
huckleberry
interior live
iron
jack
Kellogg
laurel
live
maul
mossy-overcup
mossycup
mountain white
northern pin
northern red
Nuttall
Oregon
Oregon white
oriental
overcup
peach
pedunculate
pin
possum
post
quercitron
red
Red River
rock
rock chestnut
Rocky Mtn. white
sandjack
sawtooth
scarlet
Schneck
scrub
sessile
shin
shingle
shrub live
Shumard
Shumard red
Sierra live
smooth-bark
southern red
Spanish
spotted
stave
swamp
swamp chestnut
swamp post
swamp red
swamp Spanish
swamp white
swamp willow
tanbark
turbinella
turkey
Utah white
valley
valley white
Virginia live
water
water white
weeping
white
willow
yellow
yellow chestnut
yellow-bark
ocean-spray, Holodiscus
bush
creambush
gland
Ogeechee-lime, Nyssa
oilnut, Ilex
oilnut, Juglans
old-mans-beard, Chionanthus
old-mans-beard, Clematis
oleaster, Elaeagnus
olive, Olea
olneya, Olneya
opossum-wood, Halesia
orchidtree, Bauhinia
pink
Oregon grapeholly, Mahonia
Oregon larch, Abies
Oregon-cedar, Chamaecyparis
Oregon-grape, Mahonia
Beale
Cascades
Oregon-myrtle, Umbellularia
Oregon-tea tree, Ceanothus
oriental arborvitae, Platycladus
Osage-orange, Maclura
osoberry, Oemleria
oyamel, Abies
P
Pacific madrone, Arbutus
padauk, Pterocarpus
Burma
India
palm, Sabal
cabbage
Puerto Rico hat
palmetto, Sabal
cabbage
dwarf
etonia
Mexican
Oaxaca
Puerto Rico
Rio Grande
scrub
Sonoran
palmilla, Yucca
1217
snow
Syrian
Ussuri
wild European
willow-leaf
pecan, Carya
bitter
sweet
pecky cedar, Calocedrus
pegwood, Cornus
pencil cedar, Calocedrus
penstemon, Penstemon
Bridges
bush
crevice
Leonard
littlecup
moth
shrubby
sidehill
toadflax
pepperbark, Zanthoxylum
pepperidge, Nyssa
pepperwood, Umbellularia
peronias, Adenanthera
persimmon, Diospyros
black
common
eastern
Japanesse
Texas
petty morrel, Aralia
pignut, Carya
pinabete, Abies
pine, Pinus
Aleppo
Apache
Arizona
Arizona longleaf
Arizona ponderosa
Arizona yellow
Arkansas
Armand
arolla
Austrian
Balfour
Balkan
banksiana
bay
beach
Benguet
Bhutan
big-cone
bishop
black
blackjack
blue
1218
Bolander
border limber
Bosnian
bottom white
bristlecone
bull
Calabrian
Canary
Canary Island
Caribbean
cedar
cembrian
Chiapas white
Chihuahua
chilgoza
Chir
cluster
coast
Coulter
Del Mar
Digger
dwarf mountain
dwarf Siberian
eastern white
European black
foxtail
Gerard
gray
graybark
Greek stone
hard
Heldreich
hickory
Himalayan
Honduras
Hudson Bay
Idaho white
Italian stone
jack
Japanese black
Japanese red
Japanese stone
Japanese white
Jeffrey
Jersey
Jerusalem
Khasi
knobcone
Korean
limber
loblolly
lodgepole
longleaf
longleaf Indian
longstraw
Macedonian
maritime
marsh
Merkus
Mexican weeping
Mexican white
Monterey
mountain
Muhgo
North Carolina
northern white
Norway
nut
oldfield
Pacific ponderosa
pinaster
piyon
pitch
pocosin
pond
prickle-cone
prickly
radiata
red
rock
Rocky Mountain lodgepole
Rocky Mountain ponderosa
Rocky Mountain white
sand
Santa Cruz Island
Scots
Scotch
scrub
shore
shortleaf
Siberian stone
Sierra Nevada lodgepole
silver
slash
soft white
Soledad
South Florida slash
southern
southern yellow
southwestern white
spruce
stone
sugar
swamp
Swiss mountain
Swiss stone
Table Mountain
tamarack
Tenasserim
Torrey
umbrella
Virginia
Washoe
whitebark
western white
western yellow
Weymouth
yellow
yellow slash
pinemat, Ceanothus
pio macho, Zanthoxylum
pio real, Pinus
pion, Pinus
pinxter flower, Rhododendron
pinxterbloom, Rhododendron
piyon, Pinus
Colorado
Mexican
Parry
singleleaf
two-needle
pipissewa, Chimaphila
little
striped
planetree, Platanus
American
California
oriental
plum, Prunus
Allegheny
American
beach
bullace
cherry
Chickasaw
damson
European
flowering
garden
goose
hog
hortulan
Klamath
marianna
Munson
myrobalan
Oklahoma
Pacific
Porter
red
sand
Sierra
western
wild yellow
wildgoose
poison elder, Rhus
poison-ivy, Rhus
poison-oak, Rhus
poison-sumac, Rhus
pondcypress, Taxodium
pondberry, Lindera
poor-mans-orchid, Bauhinia
popinac, Leucaena
poplar, Liriodendron
poplar, Populus
Andrews
balsam
black
California
downy
Euphrates
European black
Fremont
gray
Japanese
lanceleaf
laurel
narrowleaf
Petrowsky
plains
Rio Grande
Russian
Simon
swamp
tacamahac
western balsam
white
popple, Populus
Port-Orford-cedar, Chamaecyparis
portiatree, Thespesia
Portuguese-cedar, Cupressus
possumhaw, Ilex
possumhaw, Viburnum
powder-puff tree, Albizia
prairie shoestrings, Amorpha
prickly-ash, Aralia
prickly-ash, Zanthoxylum
common
northern
southern
pride-of-India, Lagerstroemia
pride-of-India, Melia
princes-pine, Chimaphila
princess tree, Paulownia
privet, Ligustrum
California
Chinese
common
European
glossy
Japanese
prostrate summer cypress, Kochia
purple laurel, Rhododendron
Q
quailbush, Atriplex
quaking asp, Populus
Queensland-maple, Flindersia
quercitron, Quercus
quickthorn, Crataegus
quinceberry, Cotoneaster
R
rabbitbrush, Chrysothamnus
alkali
basin whitestem rubber
Douglas
green
green rubber
low
Mojave
mountain whitestem rubber
Parry
rubber
spearleaf
threadleaf rubber
willowleaf rubber
raintree, Albizia
raspberry, Rubus
black
blackcap
flowering
purple-flowering
red
real blanco de la sierras, Abies
red heat, Acacia
red-beech, Flindersia
red-gum, Eucalyptus
red-ironbark, Eucalyptus
red-willow, Cornus
redbay, Persea
redberry, Rhamnus
hollyleaf
island
spiny
redberry, Shepherdia
redbud, Cercis
Arizona
California
eastern
Mexican
Texas
western
redcedar, Juniperus
eastern
Rocky Mountain
southern
redcedar, Thuja
Pacific
western
redgum, Liquidambar
redroot, Ceanothus
redwood, Sequoia
California
coast
Index of Common Names
1219
trident
saltbush, Zuckia
saltcedar, Tamarix
saman, Albizia
sandbur, Ambrosia
sandthorn, Hippophae
sapin concolore, Abies
sapgum, Liquidambar
sarsaparilla, Aralia
bristly
wild
sassafras, Sassafras
sau, Paraserianthes
savin, Juniperus
saw-palmetto, Serenoa
Scotch broom, Cytisus
screwbean, Prosopis
scrub-box, Lophostemon
seaside mahoe, Thespesia
Seminole-bread, Zamia
senna, Senna
armed
bladder
spiny
serviceberry, Amelanchier
Allegheny
Canadian
common
downy
Huron
Pacific
roundleaf
Saskatoon
western
thicket
shadblow, Amelanchier
thicket
shadbush, Amelanchier
shore
western
shagbark, Carya
she-balsam, Abies
she-oak, Casuarina
beach
gray
river
sheepberry, Viburnum
sheepfat, Atriplex
shellbark, Carya
big
bottom
shinglewood, Thuja
shirabe, Abies
shirabiso, Abies
shorebay, Persea
Siberian peashrub, Caragana
Sierra redwood, Sequoiadendron
silk-oak, Grevillea
silktassel, Garrya
ashy
canyon
dwarf
eggleaf
wavyleaf
Wright
silktree, Albizia
silkwood, Flindersia
siltbush, Zuckia
silver pine, Abies
silver-oak, Grevillea
silver-top shining-gum, Eucalyptus
silverbell, Halesia
silverberry, Elaeagnus
silverberry, Shepherdia
silverling, Baccharis
siris, Albizia
white
skunkberry, Lonicera
skunkbush, Rhus
sloe, Prunus
Allegheny
small custard-apple, Asimina
smokebush, Cotinus
smokebush, Psorothamnus
Nevada
smoketree, Cotinus
American
common
European
smoketree, Psorothamnus
smooth gallberry, Ilex
snakeweed, Gutierrezia
broom
perennial
threadleaf
snow eriogonum, Eriogonum
snowbell, Styrax
American
bigleaf
drug
fragrant
Japanese
styrax
Texas
snowbell tree, Styrax
snowberry, Symphoricarpos
Columbia
common
garden
mountain
Parish
Utah
western
snowbush, Ceanothus
snowdrop-tree, Halesia
soapberry, Sapindus
soapberry, Shepherdia
soapolalillie, Shepherdia
soapweed, Yucca
sophora, Sophora
sorrel-tree, Oxydendrum
sour tupelo-gum, Nyssa
sour-bush, Callicarpa
sourberry, Rhus
sourgum, Nyssa
sourwood, Oxydendrum
southern nannyberry, Viburnum
sow-berry, Callicarpa
Spanish-bayonet, Yucca
Spanish-dagger, Yucca
Spanish-mulberry, Callicarpa
sparkleberry, Vaccinium
spicebush, Lindera
bog
common
Japanese
northern
southern
spikenard, Aralia
small
spindletree, Euonymus
European
warty
winged
spineless hopsage, Zuckia
spiny hopsage, Grayia
spiny-sage, Grayia
spirea, Spiraea
Alaska
Appalachian
Beauverd
birchleaf
Douglas
Virginia
spoonwood, Kalmia
spotted wintergreen, Chimaphila
spruce, Picea
Alaska
Alberta
black
Black Hills
blue
bog
Brewer
Canadian
cat
Chinese
coast
Colorado
Colorado blue
dragon
eastern
Engelmann
Ezo
Himalayan
Korea
Koyama
mountain
Norway
Porsild
red
Sakhalin
Serbian
Siberian
Sitka
skunk
swamp
tideland
weeping
west Himalayan
West Virginia
western
white
yeddo
yellow
yezo
squaw mat, Ceanothus
squaw plum, Oemleria
squaw-apple, Peraphyllum
squaw-carpet, Ceanothus
squawberry, Lycium
squawbush, Cornus
St. Johns bread, Ceratonia
stagbush, Viburnum
steeplebush, Spiraea
sticky-laurel, Ceanothus
stinking-cedar, Torreya
stinking-yew, Torreya
storax, Styrax
strawberry-bush, Euonymus
American
running
striped cottonthorn, Tetradymia
striped princes-pine, Chimaphila
styraxtree, Styrax
sugarberry, Celtis
sugarbush, Rhus
sugi, Cryptomeria
sulfur wildbuckwheat, Eriogonum
sumac, Rhus
desert
dwarf
evergreen
false poison
fragrant
ill-scented
Kearney
lemon
1221
lemonade
Mearns
mountain
prairie
scarlet
scrub
shining
small-leaf
smooth
staghorn
sugar
swamp
sweet-scented
tobacco
velvet
wing-rib
winged
summer cypress, Kochia
summersweet, Clethra
woolly
surai, Cupressus
swallow-thorn, Hippophae
swamp black-gum, Nyssa
swamp dewberry, Rubus
swamp-cedar, Chamaecyparis
swamp-laurel, Magnolia
swamp-mahogany, Eucalyptus
swampbay, Persea
swampbay persea, Persea
swamphaw, Viburnum
sweet gale, Myrica/Morella
sweet pepperbush, Clethra
Asiatic
coastal
mountain
sweet pignut, Carya
sweet-birch, Ceanothus
sweet-breath-of-spring, Lonicera
sweet-locust, Gleditsia
sweetbay, Magnolia
evergreen
southern
sweetgum, Liquidambar
American
sweethaw, Viburnum
Swiss pine, Abies
switch ivy, Leucothoe
sycamore, Platanus
American
California
western
syringa, Philadelphus
T
tallowbrush, Cercocarpus
tallowtree, Triadica
Chinese
1222
tamarack, Larix
western
tan bay, Gordonia
tanbark-oak, Lithocarpus
tangle legs, Viburnum
tanoak, Lithocarpus
tarweed, Chamaebatia
Tasmania bluegum, Eucalyptus
Tasmanian blackwood, Acacia
teak, Tectona
tesota, Olneya
Texas locust, Gleditsia
Texas mountain-laurel, Sophora
Texas possum-haw, Viburnum
Texas-ebony, Ebenopsis
thespesia, Thespesia
thimbleberrry, Rubus
fragrant
western
thorn, Crataegus
dwarf
Eggert
green
hedge
Newcastle
parsley-leaf
shining
southern
Washington
thorny-locust, Gleditsia
thuja, Thuja
Japanese
Korean
Sichuan
tingiringy-gum, Eucalyptus
tingle-tongue, Zanthoxylum
tobacco brush, Ceanothus
todo-matsu, Abies
Todomatsu, Abies
toona, Toona
toothache-tree, Zanthoxylum
tornillo, Prosopis
Torreys joint-fir, Ephedra
torreya, Torreya
California
Florida
toyon, Heteromeles
trailing-arbutus, Epigaea
trappers-tea, Ledum
travelers-joy, Clematis
tree-anemone, Carpenteria
tree-of-heaven, Ailanthus
tree-poppy, Dendromecon
tremble, Populus
trueno de seto, Ligustrum
trumpet-flower, Campsis
trumpetvine, Campsis
tulipan Africano, Spathodea
tulip-poplar, Liriodendron
tuliptree, Liriodendron
tung-oil tree, Vernicia
tupelo, Nyssa
black
Ogeechee
sour
swamp
water
white
tupelo-gum, Nyssa
turkey-apple, Crataegus
tutui, Aleurites
two-eyed berry, Mitchella
U
umbrella chinaberry, Melia
umbrella-pine, Sciadopitys
Japanese
umbrella-tree, Melia
urajiro-momi, Abies
utis, Alnus
V
varnish tree, Koelreuteria
venetian sumac, Cotinus
viburnum, Virburnum
arrowwood
hobblebush
mapleleaf
Rafinesque
sweet
witherod
vine-bower, Clematis
vinegar-tree, Lophostemon
virgilia, Cladrastis
virgins-bower, Clematis
eastern
Texas
Virginia
western
W
wafer-ash, Ptelea
wahoo, Euonymus
wahoo, Ulmus
walnut, Juglans
American
Arizona
Arizona balck
black
California
Carpathian
eastern black
English
Hinds
Hinds black
Japanese
little
northern California
Persian
river
Siebold
southern California
Texas
Texas black
white
water jacket, Lycium
waterlocust, Gleditsia
wattle, Acacia
black
green
Sally
Sidney black
wax-myrtle, Myrica/Morella
California
southern
waxberry, Myrica/Morella
wayfaringtree, Viburnum
waythorn, Rhamnus
western Catawba-tree, Catalpa
western soapberry, Sapindas
white fringetree, Chionanthus
white poplar, Liriodendron
white sassafras, Sassafras
white-cedar, Chamaecyparis
Atlantic
Port-Orford
southern
white-cedar, Thuja
eastern
Northern
white-gum, Eucalyptus
white-sage, Krascheninnikovia
whitethorn, Crataegus
whitewood, Liriodendron
wicopy, Dirca
wild allspice, Lindera
wild China-tree, Sapindus
wild lilac, Ceanothus
wild orange, Prunus
wild vine, Vitis
wild-alder, Aralia
wild-buckwheat, Eriogonum
cushion
James
Shockley
shortstem
snow
sulfurflower
Wyeth
wild-oleaster, Shepherdia
wild-olive, Shepherdia
wild-raisin, Viburnum
willow, Salix
arctic
arroyo
Bebb
black
Booth
coastal plain
cordate
coyote
creeping
diamondleaf
feltleaf
Geyer
meadow
Pacific
peachleaf
pussy
sandbar
Scouler
weeping
white
yellow
wineberry, Rubus
winter-pink, Epigaea
winterberry, Ilex
common
mountain
winterfat, Krascheninnikovia
wintergreen, Gaultheria
mountain-tea
Oregon
wintersage, Grayia
witch-hazel, Hamamelis
American
Chinese
Japanese
Ozark
witch-hobble, Viburnum
witherod, Viburnum
wolfberry, Elaeagnus
wolfberry, Lycium
Anderson
Chinese
Rich
wolfberry, Symphoricarpos
womans-tongue
woodbine, Parthenocissus
woolly common hoptree, Ptelea
woolly nama, Nama
wristwood, Viburnum
yellow-cedar, Chamaecyparis
Alaska
yellow-poplar, Liriodendron
yellowwood, Cladrastis
Kentucky
American
yellowwood, Cotinus
yellowwood, Frangula
yerba de pasmo, Baccharis
yew, Taxus
Canada
Chinese
common
eastern
English
Florida
Guatemalan
Himalayan
Honduran
Japanese
Maire
Mexican
Pacific
Yunnan
yokewood, Catalpa
yucca, Yucca
Great Plains
Mojave
soaptree
tree
Z
zarcilla, Leucaena
Zitterpappel, Populus
Y
yaupon, Ilex
yellow hercules, Zanthoxylum
1223
Addendum
Table of Contents
Cercidium floridum
Dirca paulustris
1225
1225
Enterolobium contrisilignum
Frangula alnus
1225
1225
Persea americana
1225
1226
Styrax japonicus
1226
1227
1228
Quercus lamellosa
Quercus lineata
Quercus palustris
1228
1228
1228
Addendum
1224
Addendum
1225
Addendum
1226
Addendum
1227
1228