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VOLUME LV NUMBER 2

THE

BOTAN ICAL GAZETTE

FEBR UAR Y 1913

THE MORPHOLOGY OF ARAUCARIA BRASILIENSIS

I. THE STAMINATE CONE AND MALE GAMETOPHYTE
L. LANCELOT BURLINGAME

(WITH ELEVEN FIGURES AND PLATES IV AND V)

Introduction
Although wood of the araucarian type, strikinglyresembling
that of the ancientCordaitales,has been knownfora long timeand
has been used as evidenceof the antiquityof the araucariansand as
a proofof theirrelationshipto the Cordaitales, both of these opin-
ions have been vigorouslychallenged. It has been asserted that
theyare less ancientthan theAbietineaeand are derivedfromthem
(7, 9); that possibly theyare not related to othergymnospermsat
all and have been derived froma lycopod ancestry (I7, i8, 20);
that " the geologicalclaim forthe great antiquityof the Abietineae
thus fails on critical study of the two formsupon which it is
based" (27); that "the ancient geological and widely separated
geographical distribution (of Araucarineae), the large micro-
sporangiatecones in comparisonwith the megasporangiatecones,
the evident transitionbetween the sporophyllsand the foliage
leaves are indications of an interestingand probably primitive
group. The anatomy of the microsporophyllsand megasporo-
phylls indicates that they are homologousstructures,functionally
differentiated"(26); that "unfortunately,no teratologicalphe-
nomena, on which he always laid great stress,were known in the
Araucariae, but they were in the Abietineae and showed that
97
98 BOTANICAL GAZETTE [FEBRUARY

the single ovule was a modificationof an axillary shoot bearing

sporangia" (30, 3I).
In the face of opinionsso diverse,the desirabilityof additional
factsis very evident. We know already that the pollen grainsand
pollen tubes of Araucaria contain numerousnuclei, most of which
are prothallial or vegetative (13, 24, 25, 26), and that the same
conditionobtains in Agathis (24), with the added informationthat
there are definitely"two large male nuclei" (6). Among the
Podocarpineae the same conditions are known to be true for
Podocarpus (I, 2, 5, 8, 22, 33), Dacrydium (8, 23, 32, 33), Phyllo-
cladus (II, 33), Saxegothaea (I5, 29), and Microcachrys(28).
The presentinvestigationwas undertakenin the belief that addi-
tional knowledgeof the morphologyof the Araucarineaewould aid
very materially in clarifyingour ideas concerningthe origin (or
origins) of Coniferalesand the relationshipof the Pinaceae, par-
ticularly the Abietineae, and the Podocarpineae (and Taxaceae
generally).
The materialsforthis investigationhave been secured through
the kindnessof Mr. JAMES C. FLOOD of San Francisco, California,
fromhis countryplace, Linden Towers, at Menlo Park, California,
to whom the writerwishes to express his sense of obligation and
appreciation. He is also under obligationfor actual assistance in
makingthe collectionsto the generousassistanceof Mr. ROACH, the
gardener at Linden Towers, to whom also his thanks are due.
Collectionswere begun in March i9io and made weeklyup to the
middle of November, when they were interruptedby illness until
Februaryi9ii. Collectionswereagainmade fromNovemberI5,
i9iI, to February I9I2, thus completingthe series.
Various killingand fixingagents have been tried. A solution
recommendedby JUEL (IO) gave as satisfactoryresultsas any for
most stages. Flemming'sstrongersolution,as well as the chrom-
acetic mixturerecommendedin CHAMBERLAIN'S Methodsin plant
histology,also gave good resultsforsomestages,but failsto penetrate
the microsporangiumduring the developmentand division of the
mothercells, owingpartly to the nearlyimperviousepidermisthat
develops about this time,and partly,perhaps,to the presencethen
in the pollen sacs of a sortof mucilage,apparentlyproducedby the
19I3] BURLINGAME-ARA UCARIA BRASILIENSIS 99

degenerationof thewallsof themothercells. At certaintimesin

mid-winter practicallyall stagesof the cones are presenton the
treesat thesametime. Notwithstanding thefactjust mentioned,
stagesbetweenthe prophaseof the mothercell and tetradswere
collectedonlyfourtimes,thoughdiligently soughtfor. The same
is trueof thestagesin the development of thegametophyte from
maturemicrospores to thosein whichtheprimary spermatogenous

ma n..~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

FIG. i.-Forest of Araucaria brasiliensison the Rio Tibagy, State of Parana,

southernBrazil.-Photograph by J. C. BRANNER.

cell has alreadydivided. The reasonsforthisstateof affairsare

notveryapparent,thoughit may be that thesestagesare passed
through veryrapidlyor thatdivisionoccursduringthenight.
The habitof the treeis shownin textfig.i, representing old
treesin theirnativehabitat,and in textfig.2, an ovulatetreefrom
thegardensat LindenTowers.
The staminatecone
AmongthenumerousspecimensofAraucariain thesegardens,
to at leastfourdifferent
belonging species,thereare threestaminate
IOO BOTANICAL GAZETTE [FEBRUARY

ones ofA. brasiliensis.They are about 30 yearsold and rangein

heightfrom25 feetto about 6o feet. The largestone is about 20
inchesin diameter. Most of the collectionsweremade fromthe
smallesttree because it has had the crownbrokenout, and on
this account fruitsmuch nearerthe ground,a matterof some

FIG. 2.-Ovulatetreefrom
gardensat LindenTowers,showing
retention
oflower
branchesin youngtrees.

consequencewhenone considers waysand meansofsecuringcones

froma "monkeypuzzle" tree. This tree was selectedfor the
additionalreasonthat the rangeof stageson it at any timewas
greater. It had moreover thepeculiarity
offormingitsconesmuch
earlierin the fall and continuingto formn
new ones later in the
1913] BURLINGAME-ARA UCARIA BRASILIENSIS ioi

spring. Numerouscollectionsweremade fromone of the other

in theconesfromall
treesin orderto makesurethatdevelopment
thetreeswas thesame.

... ...... ....

tippedwith terminalbuds, in some of which,dist-

*rasiliensis .
.

..

.. .........

. ... . . . . . ... . ... ... .. .

. .~...
.....-v

LInfulth fal(uusbrnc
FIG!. 3.-tmne
oOtbe)oefid
of.,A.....,..
; rasi~iensis
..X..X,bern
;S, ,' tw old coe
h hotof preiou
fA

one can findtheveryyoung

guishedby beinglargerand plumper,
cones. As soonas thebranchhas developedfromsucha bud,one
I02 BOTANICAL GAZETTE [FEBRUARY

usually findsabout 3-6 staminate cones scattered along the new

shoot (text fig. 3). Each cone terminatesa short leafy branch,
which may occasionally continueits growththe followingseason,
but usually the cone driesup, turnsbrown,and falls offthe winter
followingthesheddingofits pollen (textfig.3). The mature cones
are very large, immediatelyafter having shed their pollen being
frequentlyas much as I5 cm. long and 3 cm. in diameter; theyare
about one-thirdsmaller before shedding. Text fig. 4 shows an
unopened cone, a cone sheddingits pollen, and the apical view of
a maturecone brokenin two,about natural size. Such a cone as is
shown in the photographhas about iooo sporophylls,disposed in
about I2 spiral rows, each of which makes 2 or 2 .5 turns. There
are usually IO-I5 sporangia,pendent (text figs.5 and 6) fromthe
expanded distal portion of the sporophyllon its morphologically
contains500-I000
abaxial side. Each sporangium pollengrains.
A little calculation will show that the output of a terminalbud
may easily reach a billionpollen grains. The pollen is shed in the
springfromthe firstof April to the latterpart of May forthe most
part, although I have found cones shedding in December and as
late as July i. The sporangia dehisce throughlongitudinalslits
on the sides of the sacs facingthe other row of sporangia,so that
the pollen is shed into the space between the innerand outer rows
of sporangia.
By dissectingthe terminalbuds already referredto, one can
recognizethe young cones when they are merelyrounded growing
points. Each one is enveloped in two or three layers of delicate
leaves and is slightlyflattenedon the side next the main axis of the
bud. Plate fig. i shows a slightlyolder cone, on which the first
sporophyllsare forming. A sporophyllis firstrecognizable as a
group of meristematic cells (plate fig. 2) just beneath a slight
emergenceof the smoothsurfaceof the growingpoint (plate fig.i).
This emergenceincreases in size rapidly by growth throughout.
Verysoon its distal end expandsuntil thewhole structuresomewhat
resemblesa mushroomwith a very shortstalk and one-halfof the
pileus removed. The stalk is so short that the abaxially placed
swollen portionof the sporophyllis in contact with the cone axis.
From its surface grow the sporangia, elongatingtoward the cone
1913] BURLINGAME-ARAUCARIA BRASILIENSIS 103

I0 -

- ~~~~~~~~~bb
-~~~~~~~~~~14h
|

FIG. 4.-Cone sheddingpollen (to right),matureunopenedcone,and cross-section

of maturecone; naturalsize.
I04 BOTANICAL GAZETTE [FEBRUARY

axis as fast as the axis of the sporophyllitselfelongates. Like the

sporophyllitself,the sporangia are firstdistinguishableas meri-

vb

6~~~~~~~~

V 7~~~~~~~~~~~~~~~~~~~~~~~~~~~

9~~~~~~~~~~~~~~~~~~~~

8
FIGS. 5-9.-Fig. 5, cross-sectionof sporophyll,showingrelationof sporangiato
one another and to the stalk; fig. 6, longitudinalsection of a sporophyll,showing
positionof sporangiaand course of theirvascular supply; fig. 7, cross-sectionof a
vascular bundle of sporophyll; fig. 8, cross-sectionof a sporangiumat mothercell
stage, showingsporogenouscells (s), tapetal cells (t), 4-6 layersof wall cells, tannin-
filledepidermalcells, and thincells wheredehiscencewill take place; X 250.; fig.9,
annulus and stomiumjust beforedehiscence; X 250.

stematic outgrowthsconsistingof several cells. Plate fig.3 shows

part of three sporangia adjacent to one. anotherin the same row,
and plate fig.4 shows a section at rightangles to it, showingthe
19I3] BURLINGAME-ARA UCARIA BRASILIENSIS I05

beginningsof two sporangia adjacent to one anotherbut in differ-

ent rows.
Occasionally one can distinguish what appears to be the
archesporiumalmost as soon as the sporangiumitself is recog-
nizable, but much more commonlyit cannot be detected until the
sporangiumis I2-I5 cells in lengthand 8-io cells wide (plate figs.
4, 5, 6). Not infrequentlysporangia as far developed as that
shownin plate fig.7 do not show any differentiation of sporogenous
cells. Ordinarilyone can recognizeall the primaryregionsof the
sporangiumat this stage. The cells of the tapetum seem to be
derived more or less indifferently from those of the wall layers
outside,or fromderivatives of the inner and presumablysporoge-
nous cells. However,I shouldnot like to be too dogmaticin regard
to this. Multiplication of cells continues until the sporogenous
tissue consistsof a mass some 20 cells in cross-section(plate fig.8)
and 40-50 cells long. The tapetumremainsone or two cells thick,
the ultimate cells being elongated radially and retaining their
positionand structuretill some timeduringthe developmentof the
male gametophyte. At this time thereare 4-6 layers of wall cells
radially flattened,with very little cell contents. The epidermisis
at this time filledwith a densely staining substance, resembling
tannin in its staining reactions, and presentinga very effective
barrierto the ready penetrationof the ordinaryreagents. In the
furtherdevelopmentof the sporangiumthe contentsof theseepider-
mal cells disappear, and the radial walls, seen in a cross-sectionof
a sporangium,thickenin the fashionof a fernannulus, except at
the point of future dehiscence. The wall cells are eventually
crushed and more or less destroyedand the tapetum finallydis-
integrates(text figs.8 and 9).

Sporogenesis
Afterthe last divisionof the sporogenouscells, the mothercells
begin enlarging,until at the prophase of the heterotypicdivision
theyhave attaineda diametermore than twice that of the sporoge-
nous cells. This eight or ten times increase of volume is largely
water, the cytoplasmbecoming greatlyvacuolated as the growth
increases. There is a correspondingenlargementof the nucleus,
io6 BOTANICAL GAZETTE [FEBRUARY

though not proportionallyso great (plate fig. 9). Once this

increasein size has been effectedin the mothercells and theirwalls,
the cell contents appear to round up and shrink away fromthe
walls more or less; just how much I am unable to say owing to the
difficultyof determiningwhetherpart or all of the shrinkagemay
not be due to the difficultiesof securingsatisfactoryfixation. The
walls themselveshave a tendencyto persistuntil afterthe young
sporeshave been formed,but apparentlytheychangetheirchemical
condition and become more or less mucilaginous in character.
Sometimes no walls at all are visible, but the spaces among the
mothercells are filledwitha thinsortofmucilage. The greaterthe
amount of this mucilage present,the poorer the fixation.
Owing to the difficulties already mentioned,I have but little
trustworthy informationconcerningthe courseof events duringthe
reduction divisions. Stages in the formationof the presynaptic
spireme,synapsis,and diakinesiswereobserved. At themetaphase
of the heterotypicdivision thereare 8 bivalent chromosomes,but
the material did not permit one to follow the method of their
formation. The spindle fibersare attached to theirapices (plate
fig. ii), and they are drawn apart as short stubby masses and
collect at the pole in a close mass. They appear at the spindle of
the homeotypicdivision as much longerrods curved into more or
less U-shaped chromosomes. No walls are formed,apparently,
until afterthe spore nuclei have passed into the restingcondition,
when a systemof fibersis presentbetweenthe nuclei,on which the
plasmatic membranes separating the young spores arise (plate
figs.I3 and 4) .
The young spores now formwalls around themselves,entirely
within the old mother cell wall if it is still present. The spores
then begin a slow enlargementand thickeningof the walls. At
firstthe growthconsistsmerelyin enlargementof the wall without
any apparentincreaseof cytoplasmor nucleus. There is eitherone
verylarge vacuole almost entirelyfillingthe spore or many smaller
ones separated by only the most tenuous cytoplasmicwalls. At
this timeno inclusionsof starchor otherfood materialsare visible.
After the spore wall has reached its mature size, but not final
thickness,the cytoplasmbecomes more abundant and the nucleus
enlarges(plate figs.i6 and I7). In the largenucleus thereis a large
I9I3] BURLINGAME-ARAUCARIA BRASILIENSIS I07

nucleolus and ratherscanty chromatic. From this is organized a

ratherlong,loose spireme,apparentlycontainingscantychromatin.
It segments to form8 chromosomes,which contract into rather
small dense oval masses (plate fig.i8).

Male gametophyte
The results of this firstdivision are a primaryprothallial cell
and a freenucleus (plate fig.i9). The latter at once divides to
produce a second prothallial cell, which takes its place over the
preceding. The next divisionyields the freetube nucleus and the
primaryspermatogenouscell (plate fig. 2 I). Meantime the pro-
thallial cells have divided (plate figs. 20, 2I, 22). The primary
spermatogenouscell now divides, yielding the usual stalk and
body cell. The stalk cell is very evanescent, usually becoming
confusedwith the general cytoplasmvery quickly. Plate fig. 25
shows one of the fewpreparationsin whichit could still be distin-
guished as a distinctcell. Anotherinterestingcase was observed
in which thereseemed to be two freeand equal cells in the unshed
pollen grain. The usual conditionof a mature pollen grain,with
I5-25 free prothallial nuclei, a recognizable tube nucleus, and a
singlebody cell, is shown in plate fig. 27.
It is in this conditionthat the pollen is shed in April (usually).
The potentialnumberof pollen grainsis fourto eighttimesas great
as the actual numberformed. It has already been remarkedthat
the ordinarynumberof mothercells is about iooo, while the usual
number of pollen grains that mature in a sporangium is only
500-Iooo. At the time theyare shed the pollen grainshave a two-
layeredwall, the outer coat of whicheasily separatesfromthe inner
(plate figs.25, 26, 27), but whichdoes not appear to do so naturally
to formwings. At the time they are shed they contain a great
many very large starch grains,a few of which are shown in plate
fig. 27. They are usually so crowdedwith it that the microtome
knife scatters the contents all about in cutting sections of shed
pollen.
Pollination
The ovulate cones are firstrecognizablein late April. At this
time thereis no trace of ovules on the scales and the pollen lodges
somewherenear the freeedge of the so-called ligule (text fig. io).
io8 BOTANICAL GAZETTE [FEBRUARY

Whether the pollen shed in the early part of the season finds a
lodgmentin a position to become effectiveis uncertain. Neither
does there appear to be any available data as to whether this
precocioussheddingof pollen occursin its native habitat. Though
Californiaand Brazil exhibita somewhatrough correspondencein
their seasons of rainfalland plant growth,yet it is probably not
exact to permitof any verysatisfactoryinferencesas to
sufficiently
the correspondingbehavior in
the two habitats.
I have not yet ascertainedcer-
tainly how long a time elapses
.9'
(S~t3\ \ after the pollen falls on the
scale beforegerminationoccurs.
/\'i \ Grains that have germinated
can be found in the latter part
of summer after the fogs have
set in. Meantime during the
/, summer the ovule is forming
and the stigmatic nucellus is
10 11 usually ready to receive the ad-
FIGS. IO, I-Fig. io, ovule and
vancing pollen tubes some time
scale in December, showing course of in Septemberor October.
pollen tubes; fig. II, course of pollen I have not so far succeeded in
tube throughnucellusto archegonium. germinatingpollen to any very
advanced stage,nor have I been
able to followwith any certaintythe courseof events in the pollen
tube beforeit reaches the nucellus. When the ovuliferousscales
are pulled apart, very numerous pollen tubes are usually found
sewed back and forthbetween the two adjacent surfacesof the
scales. One can sometimesisolate one without completely de-
stroyingit, but I have learned from such preparationsnothing
more than that thereare numerousnuclei and a body cell present.
One could inferthat theywould be there,inasmuch as theywere
in the pollen grain,and they are afterwardpresent in the tube
when it enters the nucellus. Whether divisionof the prothallial
nuclei occurs in the tube or not is uncertain,owing to the fact
that I have never been able to obtain an entire tube in which I
I9I3] BURLINGAME-ARA UCARIA BRASILIENSIS IO9

could count thenuclei and be sure of havingseen themall. These

nuclei do increasemarkedlyin size, and much of the starch which
enteredthe tube at germinationdisappearsbeforepenetrationof the
tube into the nucellus.
At the timethat the tube entersthenucellusthereare numerous
nuclei and the body cell present. I was unable to distinguishthe
tube nucleus any longer fromthe others. At firstonly a few of
thesenucleientertheportionof the tube in thenucellus. The body
cell is usually in that portion just outside of the nucellus or just
barely within it. There is apparently very little activity in the
developmentof the tube fromthe timeit firstpenetratesthenucellus
fora shortdistance until spring. I did not succeed in findingthe
mitosisconcernedin the divisionof the body cell, though the two
male cellswerefoundtwo or threetimesstillimbeddedin a common
mass of dense cytoplasm. The two nuclei frequentlydifferin size
markedly. The body cell and its nucleus have increasedin volume
afterleaving the pollen grain some six or eight timesby the time
the pollen tube has penetratedthe nucellus in October (plate fig.
28). This increase continueson an even greaterscale up to the
timeof division. Afterthe divisionof the body cell, one, at least,
of the pieces organizesitselfinto a large male cell or perhaps a real
sperm. These vary considerablyin size and in the definitenessof
theirorganization. They all agree in being verylarge and in being
more or less completelydelimited from the cytoplasm in which
they are imbedded (plate fig. 29). Almost invariably the large
nucleus is at the extremeend of the cytoplasmand the cytoplasm
is very oftenevidentlyarranged in such a way as to lead one to
surmisethat the nucleus actively changes its position in reference
to the cytoplasm. The nucleus is invariably at that extremity
which would lead in the direction that the "sperm" would be
inferredto be moving. The cytoplasmicbody is sometimesbent,
almostat a rightangle in one instanceobserved,in such a way as to
remindone stronglyof the creepingmovementsof an amoeba. In
one instance of two "sperms" lying in a common mass of cyto-
plasm, apparentlyhaving just divided,therewas presenta peculiar
fibrillarstructureremindingone of a nuclear spindle without the
chromosomesand with an aster at either pole. Whether this is
i iO BOTANICAL GAZETTE [FEBRUARY

an indicationof the presence of a blepharoplast and whetherthe

"sperm" is actually motile, I hope to be able to ascertain during
the comingspring. The divisionprobablyoccursin the latterpart
of Februaryand in the part of the tube outside of the nucellus.
During this time the pollen tube has advanced through the
nucellar tissues in a more or less irregularand branchingcourse.
When it reachestheupperpart of the femalegametophyteit usually
turnsaside fromthe apex, which is protectedby a cap of crushed
nucellarcells, and creepslaterallyalong the surfaceof the gameto-
phyteuntilit reachesthenarrowopeningleadingdown to one of the
archegonia,down whichit descendsto the neck of the archegonium.
Here it becomes perforated. The "sperm" passes through the
openingand crowdstheneck cells of the archegoniumapart without
destroyingthem,and plungesdown into the egg so vigorouslyas to
leave a verydistinctwake behindit in the cytoplasm. Fertilization
occurs in the latter part of March or firstweek or two of April.

Discussion
It is not proposedat thistimeto enterintoany generaldiscussion
regardingthe broader questions that promptedthe investigation,
but to let that await the issue of furtherinvestigationof other
phases of thelifehistoryof thisspeciesand of the otherthreespecies
of which material is available. However, it may not be amiss to
point out that this added informationconcerningthe male gameto-
phyte goes far toward inducinga beliefin the primitivecondition
of the araucarians. It certainlystrengthensthe resemblanceto the
podocarps and tends to increase our confidencein their genetic
connection. Barringthe abnormallylarge "sperms" of Araucaria
and the greaternumberof prothallialnuclei,its male gametophyte
exhibits an almost identical structureeven in small details with
that of the Podocarpineae. On the contrary,it is clear that the
type of male gametophytefound in these two tribesis essentially
different fromthatfoundin CupressusGoveniana(io) and Juniperus
communis(I4). In thesewe have a multiplicationof spermatoge-
nous cells, which may be induced, as has been suggested,by the
opportunityformore than one sperm to function,but in any case
is doubtless a reversionto an ancient habit (io, I4). This view
1913] BURLINGAME-ARAUCARIA BRASILIENSIS III

is strengthenedby the occurrenceof more than two sperms in

Microcycas(3) and Ceratozamia(4). The depositionof the pollen
at a distance from the nucellus has also been recordedforSaxe-
gothaeaconspicua (I5), and serves to strengthenthe likenessof the
araucarians and podocarps.
By long odds the most interestingand suggestivefeatureof the
male gametophyteof the araucarians yet knownis the remarkably
large "sperms" and the possibility of their motility. The one
figured(plate fig. 29) is about I50 i- long. It is by no means the
largest one observed, but seemed to be the one most definitely
organized. Such dimensionsare only known elsewhereamong the
actively swimmingsperms of the Cycadales and Gingko. It is of
interestto note that the kauri (6) appears to have two large nuclei
also, thoughit is not stated that theyconstitutepart of organized
cells. Inasmuch as Agathis has usually been held to be more
primitivein most respectsthan Araucaria, it may be possible that
they will yet be found there. The aster-likefibersfound in the
dividingbody cell are of course very suggestiveof a blepharoplast,
and one might perhaps legitimatelyexpect such structuresto be
foundin connectionwith such large and possiblymotilemale cells.
Again theymightbe only such structuresas have been recordedby
LAND (I2) in the egg of Ephedra.

Summary
i. The staminate cones are extraordinarilylarge and have
numerous sporophyllswith an indefinitenumber of pollen sacs
pendent fromthe abaxial side of the swollen apex.
2. An almost incomprehensibly great number of pollen grains
is produced.
3. The method of differentiating the sporogenous tissue is
variable and indefinite. The size of the sporangium and the
numberof microsporesis subject to wide fluctuations.
4. The structuresconcernedin dehiscenceare very fernlike.
5. The chromosomenumberin the male gametophyteis 8.
6. Prothallialtissueis formedin a manneralmostidenticalwith
that in the Podocarpineae, but the number of cells so formedis
greater.
II2 BOTANICAL GAZETTE [FEBRUARY

7. The pollenis shed withnumerousprothallialnuclei,stalk

nucleus,tubenucleus,and bodycellfreein it.
8. The pollendoes not fall upon the nucellusbut upon the
ovuliferousscale at a considerabledistancefromthe positionof
theovule.
9. Abouta yearelapsesbetweenpollination and fertilization.
iO. No ovulehas yet been formed on the scale at the timeof
pollinationand thepollentubedoesnotreachtheovulefor5 or 6
monthsafterward.
i i. The bodycellis single,
at leastusually,and large.
I2. Two male cells (spermsperhaps). are formedand are
usuallyunequal.
i3. There may be a blepharoplast-like body associatedwith
thedivisionof thebodycell.
I4. Fertilizationoccursabout thefirstof April.
STANFORD UNIVERSITY
CALIFORNIA

LITERATURE CITED
I. BROOKS, F. T., and STILES, WALTER, The structure of Podocarpusspinu-
losus. Ann. Botany 24:305-3I8. pI. 2I. I9I0.
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3. CALDWELL, OTIS W., Microcycascalocoma. BOT. GAZ. 44:II8-I4I. pis.
I0-I3. figs. I4. I907.
4. CHAMBERLAIN, C. J., Morphology of Ceratozamia. BOT. GAZ. 53:I-I9.
pl. I. figs.7. 19I2.
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BOT. GAZ. 33:89-iO7. ps. 5-7. I902.
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i6o. I9I2.
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GAZ. 42: I-I5. pIs. I, 2. I906.
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4I:355-364. figs. 5. I907.
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44:435-444.PIs. 28-30. I907.

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19I3] BURLINGAME-ARA UCARIA BRASILIENSIS I I3

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II4 BOTANICAL GAZETTE [FEBRUARY

EXPLANATION OF PLATES IV AND V

All figureshave been drawn under Zeiss apochromaticobjectives and
compensating ocularswiththe aid of a cameralucida. Figs. 2-8 weredrawn
with4 mm.obj. and compensating ocular6; figs.9-27, underobj. 3 mm. and
ocular I2; figs.28 and 29, under3 mm.obj. and compensating ocular6.
Fig. i.-Longitudinal sectionof a youngcone froma terminalbud: sc,
scale leaf; sp, youngsporophyll;vb,vascularbundle; XI2.
FIG. 2.-Primordial meristemof veryyoungsporophylldrawnfromsame
cone as fig.I; X250.
FIG. 3.-Cross-sectionofa cone showingthe sporangialmeristems;X 250.
FIG. 4.-Longitudinal sectionof cone of same age as preceding: section
passes throughthe stalk of the sporophylland throughthe primordiaof one
sporangiumin each row; c.ax, axis of cone; X 250.
FIG. 5.-Transversesectionofconeand longitudinal sectionofsporangium:
sporogenoustissuenot defined; X 250.
FIG. 6.-Same as fig.5, but showingusual differentiation of sporogenous
tissue(spt); X 250.
of tapetum(t); X 250.
FIG. 7.-Usual stage of differentiation
FIG. 8.-Cross-section of sporangiumafter last sporogenousdivision;
X250.
FIG. 9.-Somewhat shrunkenmothercell showingeighttimesincreasein
volume; X770-
FIG. io.-Portion of a nucleusin whichthe heterotypic chromosomes are
forming;X 770.
FIG. II .-The 8 heterotypic chromosomes;X 770.
FIG. I 2.-Second divisionofmothercell; X 770.
FIGS. I3 and I4.-Tetrads, showingcell division; X 770.
FIG. I5.-A groupof youngmicrospores;X 770.
FIGS. I6 and I7.-Mature microspores;X 770.
FIG. I8.-Division of microspore nucleus; X 770.
FIG. I9.-Showing firstprothallialcell (pc'); X 770.
FIG. 20.-Mitosis of a prothallialcell; X 770.
FIG. 2I.-Showing twotiersof prothallialcells (pc' pc"), primarysperma-
togenouscell (p.s.c.), and tube nucleus(t); X 770.
FIG. 22.-Division ofprimaryspermatogenous cell; X770.
FIGS. 23-26.-Series of transverse sectionsofa pollengrain: pc', basal tier
of prothallialcells; pc", secondtier; tn,tube nucleus; bcn,body cell nucleus;
bc,body cell; sc, stalkcell; scn,stalk cell nucleus; X 770.
FIG. 27.-Sheddingstageofpollengrain; X 770.
FIG. 28.-Body cell about OctoberI5; X385.
FIG. 29.-A male cell just beforefertilization;X385.
BOTANICAL GAZETTE, LV PLATE IV
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