A Revision of Mouriri, Section Nesophytum

Author(s): Thomas Morley

Source: Brittonia, Vol. 9, No. 2 (Jul. 30, 1957), pp. 109-131
Published by: Springer on behalf of the New York Botanical Garden Press
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1957] MORLEY: MOURIRI 109

4.5-5.5 mmlongae suborbicularesapice rotundataevel late obtusae,extus basim

versus sparse furfuraceae. Hypanthium glabrum 5 mm longum (ad torum);
calycistubus 1.2-1.3 mmaltus, lobis vix conspicuisca. 0.8 X 3.1-3.5 mm.Petala
11.5 X 7.5-8.5 mm obovata apice rotundato-truneata inconspicueciliolata (ciliis
ca. 0.1 lum longis) apicali-centralitermodice punctata. Antherae inter se ad-
haerentes; filamenta10.5 mm; thecae 9.8 mm longae lineares apice dorsaliter
porosae, connectivodorsaliterin calear hebes 1.1 X 0.6 mm prolongato. Stylus
13.6 X 0.5-0.6 mm; stigma truneatum; ovarium liberum glabrum 6-loculare
apice truneatumet coneavum,fovea ca. 0.6 mmalta.
Type collection:Earl L. Core 677 (holotypeUS 2105274; isotypeNY), col-
lected on the Anori-Crucesroad along the Rio Anornabout 12 km northof Anori,
elev. 1000 m, Dept. Antioquia, Colombia,May 11, 1944. "Small tree 25 ft high
overhangingcamino,flowerspink.'"
Paratype: A. E. Lawrence 744 (NY), from El Hiumbo,Boyaca, Colombia,
elev. 1100 m, April, 1933.
The leaf-bladesof the Lawrence collectionare slightlythickerthall in the
Antioquia material. Lawrence's label notesgive the habit as a vine 100 ft long.
The leaf blades of T. subscaberula are thinlycoriaceous and the pedicels at
anthesisare only4-6 mmlong. To T. subscaberulaI have referredthe following
collections,all fromeitherthe Pacific coast of El Valle or the Choco,with speci-
mens at both F and NY: Cuatrecasas 16226, 16602, 17948, 21394, and Killipp&
Cuatrecasas39106. Othernear relativesof T. membranaceaare T. praecox Glea-
son (with much more distinctlylobed calyx), T. regeliana Cogn. (with slightly
thickerleaf-blades,shorterpedicels,and solitaryor few-fasciculateflowers),and
T. superba Naud. (with much larger, thickerleaves and wider secondaryvein
spacing, longer hypanthia, and long ovarial collar). To T. saperba, I have
referred(ex char.) Little 7949 (NY, US), fromSan Antonio,Huila, and (with
some reservations) Cuatrecasas 20526 (F, NY), from the Rio Bugalagrande,
El Valle.

A REVISION OF MOURIRI, SECTION NESOPHYTUM

THOMAS MORLEY
Department of Botany, University of Minnesota
Minneapolis

The small West Indian sectionNesophytumhas been one of the least under-
stood sectionsof Mouriri (Melastomataceae). For this reason, and because the
area is relativelyaccessiblefromthe United States, a detailed study of the group
was undertaken. This will formpart of a proposedmonographof the genus.
The sectionNesophytumbelongsin the subgenusMoiuriri,as delimitedby me
in 1953. Of the several closelyrelated sectionsin this subgenus,Nesophytu,m is
the only one confinedto the islands of the Caribbean. Species of this sectionare
primarilydistinguishedby relativelysmall leaves whichoftentend to be obovate
withroundedto emarginatetips; bv stomatalcryptswhichwhen well developed
have the formof a shortand very broad T in cross section (Type III) ; and by
the regular presence of a hypodermiswhich does not contain wall thickenings
of the "mucilaginous" type. In the presentstudy it was learned that elliptic
and even ovate leaves are morecommonthan had been previouslysuspected,and
1 This work has been supported in part by a research grant from the Graduate School of
the Universityof Minnesota. The lists of exsieeati are published with the aid of a contribution
by the author.
110 BRITTONIA [VOL. 9

that the smaller cryptsusually have a simaplerform (Type II) than the larger
ones,but nothingwas foundthat mightinvalidate or alter the boundariesof the
section.
Section Nesophytunit was proposed2 to include twelve iianaed species. Con-
sideringtheirWest Indian distribution,it is not surprisingto learn that eight
weredescribedby Urban and Ekman, or by Urban alone, and threeby Grisebach;
one was named by Britton. Charactersprimarilyemployedby these authorsfor
the twelve"species" in questionwere shape and size of leaves, numberof flowers
per infloreseence,length of pedicel, shape of calyx-lobe,shape of ovary, and
numberof locules.
Regardless of the distinctioniclaimed for these species, inaiy of thein look
superficiallyvery much alike, so that it was difficultto avoid the beforehand
conclusionthat mnany should be combined. In order to reevaluate the species of
the sectiona surveywas made of as mnany variables as possible,in an effortto
broadenthe basis of comparison.Anatomicalcharactersof vegetativeparts were
especially desirable,in yiew of the genlerallysimilar foliage of the differentspe-
cies, the intergradingof several floralcharacters,and the fact that many of the
collectionsare sterileor in fruitonly: the fruithas so far provenof little value
in classificationin Mouriri except for its size at maturityand except for the old
ealyx-lobesit may bear. Yet in this section some of the species are based only
on sterileor fruitingspecimens.
Pressed materialwas exaniinedfrontthe herbaria listed below,eitheron loan
or at the source; I am very gratefulto the curatorsof these collectionsfor the
use of their material. Abbreviationsfor the institutionalherbaria are those of
Lanjouw and Stafleuin the Index herbariorumn (1956) : Arnold Arboretum(A)
Chicago Natural History Museum (F); Gray Herbarium,Harvard University
(GH); Herbario de la Salle, Havana (LS); Universityof Minnesota,Minne-
apolis (MIN); Missouri Botanical Garden (MO); New York Botanical Garden
(NY); NaturhistoriskaRiksmuseumn, Stockholm (S) ; Estacion Experimental
Agronomica,Santiago de las Vegas, Cuba (SV) ; Universityof California,Berke-
ley (UC); and United States National Museum (US). In addition to these,ma-
terial fromtwo privateherbariawas consulted. Ownersof theseherbariaand the
designatedsymbolsare as follows: Dr. Jose de Js. Jimenez(JIM), Calle Maximo
Gomez38-40,Santiago de los Caballeros,Dominican Republic; and Juan T. Roig
(ROIG), whose herbarium is housed at Estacion Experimental Agronomica,
Santiago de las Vegas, Cuba. Finally, the symbolGOET refersto the System-
atie-GeobotanicInstitute,Universityof G6ttingen,which has reportedto me on
the presenceof certain specimensin its herbarium.
Regarding the herbariummaterial, it will be noted below that the Cuban
plants have been much more intensivelycollected than those from Hispaniola
and Puerto Rico, and are thereforebetterunderstood. Moutririgonavensisand
M. hellerivar. helleriin particular need furthercollecting.
Field observationsweremade of both Cuban species of sect. Nesophytum,and
preservedspecimensof their wood and leaves were taken. Flowers and fruits
were also foundand preservedof one of these species,and seeds sent to the Uni-
versityof Minnesota. A few of the seeds germinatedsuccessfullybut none lived
past the seedling stage. No material suitable for chromosomestudies was
obtained.
2 Morley, Thomas. The genus Mouriri (Melastomaeeae). A sectional revision based on
anatomy and morphology. Univ. Calif. Publ. Bot. 26: 223-312. 1953.
1957] MORLEY: MOURIRI 1ll

In preparationfor microscopicstudy the leaves of a imajorityof specimens

seen were sectioned,and whole pieces fromalmost all were cleared and stained.
Flowers of mostspecimenswere cleared and stained,and sectionscut 125mthick
afterwholemountshad been made of the various appendages. The methodsused
were those described previously (1953).
I should like to express my appreciation particularly to Brother Alain of
the Herbario de la Salle forhis generosityin sendingmaterial,and to Dr. R. A.
Howard of the Arnold Arboretumand Dr. Jose de Js. Jimernez of Santiago for
their patience and helpfulnessin answeringmy queries.
DISCUSSION OF CERTAIN CRITERIA

VegetativeStructure.Shape and size of leaves prove to be extremelyvariable,

and do not warrant the emphasis of previouLsauthors. Nonetheless,one species
is partly characterizedby its unusually small leaves, and slight differencesin
shape are oftenuseful in distinguishingthe others.
An anatomicalcharacterof considerablevalue is that of the stomata.lcrypts.
Cryptshave been found on all specimensexamined except one, Ekmnan15810;
this sterilespecimenis here interpretedas an atypical memberof M. spathulata.
The variable featuresof cryptsthat concernus here are those of size and fre-
*quency (fig. 1), best referredto as the single character size-frequency.Size-
frequencyhas a characteristicrange of variation in each species. Only in M.
gonavensisis the range entirelywithinthat of anotherspecies, and the number
of collectionsis too small to give morethan a very roughindicationof the actual
range. In one species, M. spathulata, the two varieties have conspicuouslydif-
ferentranges of variation; in M. emarginatathe varieties have no real differ-
ences in this respect. Collectionsof M. helleriand M. gonavensisare too few to
show differences within the species. In figure1 the total range of variation is
seen to illustratethe close inverse correlationbetweensize and frequency,and
to suggesta hyperbolawithM. spathulata at one extremeand M. emarginataat
the other. These are the two most widely separated species of the section,con-
sideringall characters,the othertwo being more or less intermediate.Thus the
cryptsize-frequencymightserve as a single characterfor the identificationof
eitherof these two species, within the section,if it should fall in an area not
overlappedby one of the otherranges. The size-frequencyappears to be corre-
lated withthe environment to some extent. As indicatedin the generaldiscussion
below,the rainfall in the range of the species presentis higherin Puerto Rico
and presumablyin Hispaniola than in Cuba; and the larger and fewer crypts
are found in Cuba.
In the lowerleftcornerof figure1 will be seen a miscellaneousclusterof size-
frequenciesnot included in the outlinesof the different ranges of variation. All
four species are representedin this group. It is curious that in some collections
the cryptsmay be veryfew,or theymay be distributedin a very irregularman-
ner over the leaf surface,in some places few or absent and in othersnumerous.
The irregularitiesmay be apparently random, or, as in Ekman H10601, the
cryptsmay be absent except at the verytip of the lamina. In such leaves I have
read the frequencyas a rough average of the extremes. To me it seems reason-
able to regard such leaves as abnormal,so far as cryptdevelopmentis concerned,
and these size-frequenciesare thereforenot ineluded in the outlines drawn,
althoughdoing so would not materiallyaffectresults. It appears probable that
these seemingabnormalitiesare at least in part reflectionsof unusual environ-
ment,eitherof the whole plant or of parts of it. An example in point is that of
112 BRITTONIA [VOL. 9

170
\-M.
0 - M. spathulata
gonovensis 2
160 \ -M. helleri
0-M. emorginata
150 0

sb
140

130 0

E 1200
E

Ss
if 1mm
M 100 so0
@0

90 0

80

70h0

60

50~~~~~~~~~
E
40~~~~~~~~~
20
z 0~~

U)~~~
10 0 Q 0@ ~ 00a~ ~
30,u 40Op 50Ou 60Op 70, 80,uo 90)a

Diagram oflongitudinalStomotol crypt diametenr

West East H-ispani-

Cuba Cuba ala
e

o0
2ak an enlvda eolein b s h aite rcyoaadsahlt fM ptuaa

08

FIS 2, 3. M. sah4a(ka930)foesinbdreptilyaerxasonn5
0

e 1400

130'-

FIG. 1. Stomatal crypt size anid frequenceyis Mouriri sect. Nesophytutm. Eachi symbol
niarks an i-ndividualcollection. sb, Ss, the varieties brachypoda and spathulata of M. spathulata;
hi,M. helleri; g, M. govavensis; ee, er, the varieties ermarginataand rostrata of M. emargin~ata.
FIGS. 2, 3. M. spathutlata (Ekman 9350) flowers in bud, respectively after expansion in 5
per cent NaOH and dry. FiGs. 4, 5. M. emarginata (Wright 2467) as in figs. 2, 3. FIG. 6.
Diagram of longitudinal section of ovary sho'wingangle 'loll. PIGS. 7, 8. Cross sections of
leaves showvinguppeir epidermis. e, epidermis; h, hypodermis; p, palisade layer. FIG. 7. M.
1957j MORLEY: MOURIRI 113

Morley 869, M1.enarginata var. rostra1a. This came from a small tree about
71?2ft high withfewbranchesand leaves, apparentlynot in robusthealth,grow-
ing in light shade; one leaf on a small sucker shoot 11/2 ft fromthe base had
crypts0-12/mm2,40,uin diameter; anotherleaf fromthe same shoothad crypts
30/mm2and 45/A in dianmeter; a leaf fromhalf way up had crypts20/mm2anid
47juin diameter;two leaves fromthe crownhad respectivelycrypts60,/mm2and
4Op.in diameter,anid55/mM2and 50Ain diameter. In this extremeexample the
firstleaf mentionedis certainly abnormal in crypt frequency,and it is my
opinion that the crownileaves representthe nornm for that individual growing
under those conditions. A similar example was found in a tree of the related
,enus Coryp,hadeniaMorley,C. monantha,, a sucker shoot of which at the very
base had feweranidsmallercryptsthan the leaves higherup. On the otherhand,
amongtheleaves withwell-developedcryptsthesituatiolnis altered: in fivediffer-
ent individuals of M. spathtulata,leaves fromthe lower, more shaded branches
were found to have slightlymore and slightlysmaller cryptsthan leaves of the
crown. So far as the taxonomistis concerned,variationlcaused by environmentis
probably fairly well taken into account by the more or less random collection
methodsthat have built up the herbaria.
At firstit was believedthat the nuLmber of surface stomata,those not within
erypts,mightbe a reliable supportingchariaeter.This expectationwas lnotreal-
ized, however; the numberis too variable, and these stomata,moreover,do not
always stain well enough to be seen.
Anothervaluable leaf characteris that of epidermisconstruction.The epi-
dermisvaries fromsingleto irregularlydouble or occasionallytriple,apparently
as a resultof perielinaldivisionsin the epidermalinitials. This characteris best
observedin the upper epidermis. In M1.emnarginata (fig. 8) at least 40 per cent
and usually 50 per cent or more of the upper epidermisis double in any leaf,
countingonly directlyopposite pairs as seen in cross section. The double areas
are of varyingsize and are evidentlyrandomlydistributed.A second epidermal
characterof M. emarginatais a thickeningof the inner walls of at least some of
the upper or lower epidermal cells or both with a pearly-whitemucilaginous
substance. In the other species of the section the upper epidermisis predomi-
nantlyone cell thick (fig. 7), very rarely (in 31. spathulata) as much as 40 per
cent double,and mucilage-thickened walls have not been found. Moutririemargi-
nata can thus be distinguishedon the basis of epidermisalone. I have founldno
correlationbetween variatiolnsin environmentand the presence of muLcilage-
thickenedwalls.
The terminalfoliar selereidswere checkedcarefully,but differences in form
were few, about the only character of diagnostic value being the degree of
elongationof the main selereidbody. The elongationis greatestin M. spathulata,
least in MI. emarginata,and more or less intermediatein the othertwo species.
Exceptions are common,however. Differencesin arm lengthand branchingare
occasionallyof use.
Minor differences, sometimesuseful, have been found in the level of the
smallest veinlets in the leaf. In M. spathulata var. brachypoda these veinlets
are found at a level 0.38 to 0.46 of the distance from the bottomto the top.
In M. gonavensis,they are 0.45 to 0,48 of the distance,and in M. helleri from
0.40 to 0.52. Weak thoughthis criterionis, it has proved of value occasionally

spathulata (E7cman 9350). FIG. 8. M. emarginata (Morley 869). FIG. 9. Angle "'ol" in
individual collectionisby geographie area; legend in fig. 1.
114 BRITTONIA [vOL. 9

as a supplement. Veinlet level was not measured for the Cuban plants, where
the epidermal characteris conclusive.
Floral Structure. The numberof flowersin the infloreseeneeproveson exami-
nation to be a veryinconstantfeature. Some species do tend to have more fully
developed infloreseencesthan others,and differencesin the number of nodes
presentin the main axis of the infloreseence
can be detectedbetweensomespecies,
but these differences are of secondaryimportance.
Length of pedicel is anotherfeaturethat proves to be highlyvariable within
the species,and of littlereliability.
With regard to the floweritself,one of the most obvious differencesis in
over-allproportion (figs. 2, 4); however,the width decreases on drying (figs.
3, 5) in such an erraticmannerthat this characterhas not been used. Another
variable in the general shape is that of the curvature of the base of the hy-
panthiumwhereit is adnate to the ovary. This characteris very consistentand
has proved to be independentof calyx proportion,and so has been used, even
thoughit too may change somewhatas the flowerdries. In order to expressthis
feature numerically,measurementswere made of the angle, here designated
angle "o," betweentangentsdrawn at the most vertical part of the ovary wall
and at the part nearest the horizontal (figs. 6, 9). These measurementswere
made feasible by the use of projectiontracings,the flowersof almost all collec-
tions being already sectioned and mounted on slides. The angles of opposing
sides were averaged to give the figureused. Representativemeasurementscan be
obtained fromflowerstaken during and before anthesis, even very immature
flowersgiving results that appear characteristic. Of the two extremes,cup-
shaped and obconic,the latter can probablybe consideredthe more specialized
owing to its association with flowershaving a smaller number of locules and
ovules.
The calyx-lobesprovide some of the muost valuable criteria. In Momririthe
hypanthiumis prolongedabove the inferiorovary to the point of stamenattach-
ment,whencethe calyx lobes diverge. These vary considerablyin shape, and this
variation is one of the features that most attracted the attentionof previous
authors. Variation involveslength,width,and the shape of apex and shoulders
(figs. 10-17). Shape within the species is highly variable, but nonethelessis
sometimesdistinctive.Direct measurementsof length (fig. 18) and the length-
widthratio (fig.19) have each someadvantages,but both showthe relativesepa-
ration of the two species in Cuba, and the lack of it among the species in His-
paniola. By using, instead of the latter ratio, the ratio of total calyx-length
(calyx lobe,freehypanthium,and inferiorovary) to lobe-width,an inconsistency
of groupingin the Hispaniola species is eliminated,but this ratio has the draw-
back of being restrictedin use to floweringmaterial.
A bettercalyx characteris that of fusionbetweenlobes. In the bud, adjacent
lobes are fused at least a shortdistance above the point of petal attachmentin
almost all individuals of the section. Down the outside of the fused part there
is a crease along which splitting takes place at anthesis. Measurementshave
been made fromthe upper edge of the stamen sear rather than fromthe petal
sear because'the formeris small and clearlydefined,while the latter is large and
variable in size and with limits that are sometimesdifficultto determine.
Measurementswere made with a microscopeeyepiecemicrometer.The heightto
which the lobes are fused is a variable of major importance(figs. 20, 22, 23).
Even aftersplittinghas taken place, the heightof previousfusion can be deter-
minedby microscopicexamination.Althoughthe differences are small,the char-
1957j]MORLEY: MOURIRI 115

10 I - |2
1 13

West East Hispon,- Puerto West East Hispont- Puerto

Cuba Cubo ola Rico Cuba Cuba ola Rico 14
3.0 00 0
0

2.5 1.5 0 *

0 ~~~~~~~~~0g
0~~~~~~~
0~~~~~~~~~
2.0 rnm5
0 22
0 1.0 0 0

1.5 Q:p 6 ( 0

1.0 17 .5 0
*
0~~~~~~~~~~~~~~~~~~
Calyxotbe length in mm. RaFtia of calyx ambe length to width

0 ',5'f'
West East Hispani- Puerto West East Hispaoni- a
Cuba Cuba ala Rica Cuba Cuba ala 2mm-
0
2.5 - 0 -

1.5 0 0

00 2.0
M. *ayiaavr
0
0 mrint
eCabl 0
07.0FI. 14 enr*nt
.eagiaavr
2
0
1.0 0 @ -
0 00 Q ...
0

5 @00It0o

0~~~~~~~~~.
Fusion distance of calys lobes Ovary height in mm. C ) C D C
in mm.
20 21 23
FiGs. 10-17. Calyx-lobes of Mouriri fromithe outer sidle. Dotted lines represenitpotential
iiiies of separationi,bars inidicate level of stamienattachment. FIGS. 10, 11. M. spathulata.
FIG. 10. Acuiia, April 16, 1945 (SV 12603). FIG. 11. Victorin,,Clemente 4- Alain 21835.
FIGS. 12-14. M. emarginata. FIG. 12. M. emar-ginatavar. rostrata (Le,6n 22383). FIG. 13.
M. ernar-ginatavar. emarginata (Cvtrbelo 307). FIG. 14. M. enmarginatavar. enitarginata
(Acwtha,April 11, 1945 (SV 12601). F'IGS. 15, 1.6. M21. gonavensis. FIG. 15. M. gonavs?nsis
var. goniavensis (Ekman H8712'). FIG. 16. M. gonavensis var. hottensis (Ekman H10399).
FIG. 17. M. helleri (Abbott 2233). FIGS. 18-21. Character distribution showing individual
collectionisby geographic area; legend in fig. 1. FIGS. 22, 23. Diagrams showing the inner
side of calyx-lobes. p, petal- scar; s, stamen scar. FIG. 22. Type with very little fusion be-
tween aidjacent lobes. FIG. 23. Type with considerable fusion between adjacent lobes.
116 BRITTONIA [Vol,. 9

acter appears to be a veryreliable one. Several points of interestare illustrated

in figure20: the apparentlywider ranges of variation in the Cuban species as
comparedwith the Hispaniolan ones; the greater overall range of variation in
easternCuba as comparedto Hispaniola; the distinctgap that sets apart the two
species in eastern Cuba and the similar one settingoffM. gonavensis in His-
paniola; and the differencebetween the western and eastern varieties of M.
emarginata. Using instead of directmeasurementsthe ratio of fusiondistanceto
lobe width the results are only slightlydifferent.Calyces with greater fusioll
distancesare here consideredto be more specialized than those with smaller,in
accordance with generallyaccepted principlesof fusion of parts.
Ratios betweenmany differentcharactershave been tested,but except for
those mentionedthey were found generallyless satisfactorythan direct meas-
urements.
Color of petals is evidentlyone of the more variable featuresof the section.
In by far the majorityof individuals the color is fromwhiteto violet,or some-
times pink or red. In two collections,however,one of M. spa,thulata(Howard
5933) and one of M. emarginata (Shafer 589), "flower" color is reportedto be
yellow. The frequencyof yellow-flowered individuals is difficult
to judge owing
to the failure of most collectorsto note petal color on their labels. So far as I
can tell, this characteroffersno basis for segregationat any level.
The structureof the ovary offerssome distinctionsof value. Ovary height
(fig.21) is one examDle. The numberof locules in the ovary has sometimesbeen
stressed in descriptions,primarily,I believe, because it is a relatively stable
characterin mostgenera and was assumedto be so here also. Numberof locules
is, however,highlyvariable in this section,and only ranges of variation have
any meaning. In M. spathulata,for example,the numbervaries fromtwo to five,
and in one specimenalone (Wright1234) locule numbersof two,three,and four
were found. The numberof ovules is also quite variable, but may be relatively
less so than locule number; in the specimenjust cited the respectivenumbersfor
the threeflowerswere ten,nine,and eleven. As pointedout previously(footnote
2, op cit.,p. 255), a locule numberof fiveseemsto be primitivefor the genus.
GENERAL DISCUSSION

The primaryresult of this reevaluationhas beelna reductionin the number

of species fromtwelveto four,includingtwo varietiesin each species. The dis-
tributionof six of the principal distinguishingcharactersis summarizedin fig.
24. In this diagramangle " o" would have been preferableto the length-to-width
ratio of calyx-lobesas one of the coordinates,but to use it would have meant
omittingseveral importantcollectionsfor which this characteris not available.
The greatervariabilityamong the longer calyx measurementshas resulted in a
scatteringof the M. emazrginata plots. This diagram and figures1, 9, and 18-21
illustratethe points discussedbelow.
The general picture given by the four species is of a group containingtwo
basic types,one representedby M. spathulata,the otherby M. helleri,M. gona-
vensis,and M. emarginata. The latter type is distinguishedfromthe formerby
a generallymore obconic ovary with fewer locules, by relativelylonger calyx
measurements,and by stomatal cryptsthat tend to be larger and fewer. The
distinctionsbetweenspecies of the second type tend accordinglyto be somewhat
less clear-cutthan betweenany of themand M. spathulata.
Of the two types,M. spathulata,appears to be the more primitive. Reasons
for this conclusionare its smaller crypts,often of Type II (a less specialized
19571 MORLEY: MOURIRI 117

conditionaccordingto the progressionsuggestedin Morley,op. cit., p. 233) ; its

generallylesser fusion distance between calyx-lobes; and its locule number of
2-5, as comparedto 1-3 forthe otherthreespecies. The obconicovary,construed
as a specialization,would support this interpretation.Within each basic type,
it can be seen that the stomatalcryptsare smaller and more numerousin His-
paniola (and also in Puerto Rico forthe secondtype) than in Cuba (figs.25-28),
althoughthe total range of crypt variation, including all species, is about the
same in the two differentareas. The fusion distance of the calyx-lobes,while

1.9

1.8 _:

'.7

1.6 4
_ 1.5 Simmature)
4-
3 1.4 4
0 :

,1.2
.

.0 bo
0~~~~~~~~~~
1.0I h

o _
.9

4-
0
.8

.7 - b
.6 _
-b

.5

.4, .
.4 .5 .6 .7 .8 .9 1.0 1.1 1.2 1.3 1.4 1.5 1.6 1.7 1.8 1.9

24 Fusion distance of calyx lobes above stamen scar

* - M. spathulata Upper epidermis Angle " oa (see fig. 6)

b -var. brachypoda 0-< 30% cells doubled, no O- <1570
- M. helleri mucilage-thickened walls >157?
s - var. samanensis
0 M.
gonavensis=0-30- 40% cells doubted,no
--M. gonavensis mucilage-thickened walls Ovule number
h - var. hottensis =O->40% cells doubled, mucilage- 0 - 12-18
O -M. emarginata thickened walls present O- iI
r _ var. rostrata
O _ 10
these characters Ovary height 9: -7
not available 0 - 1.1- 1.5 mm
0-
?7 or 8
1.55- 1.95
0- 2.0 - 2.6

FIG. 24. Diagrammatic summationof six principal characters of Mouriri sect. Nesophyturn.
See text. M. spathulata symbol unmarked means Ml. spathulata var. spathulata, and so with
the other species.
118 BRITTONIA [VOL. 9

changin-ig little in MJ.spatlttlata,in the othertype progressivelyiniereasesfrom

east to west, with the exceptionof M. emnarginata var. rostrata. The develop-
mentof a specialized doubled epidermisis also conspicuousin the Cuban repre-
sentativesof the second type. These trendssuggestthat the eastern representa-
tives of both types may be less specialized than the westernolnes. Olnthe other
hand, the small leaves of ll. helleri in the east can probablybe regarded as a
speeialization; and angle "o" appears to show a slight over-all increase from
west to east, taking both types into aecount. However, the significanceof the
last characteris uincertaiii,ancdthe preponderanceof available evidence may-
thereforebe said to support the hypothesisthat the eastern representativesare
in generalthe less specialized. At the same time,and for the same reasons,these
eastern representativesresenmble each other nmoreclosely in over-all characters
than do the westernones. These facts and interpretations, if correct,are prob-
ably related to the originand migrationalhistoryof the group,but in just what
manner must remain a matter of speculation. If ancestors of the group first
enteredthe area fromCentral America, as appears to have been the case with
most plants now restrictedto the GreaterAntilles,the sectionmightbe said to
supportMatthew'shypothesisthat the mostconservativemembersof a group of
organismswould be foundfarthestfromtheircenterof origin (Ann. N. Y. Acad.
24: 171-318. 1915).
Distributionof species in the sectionNesophytutm is noteworthyin that the
four principal isolated areas occupied, iindicatedin figures26-28, are all linked
to one anotherin turn by the distributionranges of different helleri
species: M11.
occurs in Puerto Rico and Hispaniola; M. spathulata in Hispaniola and eastern
Cuba; and M1.emarginatain eastern and westernCuba. In each instance the
membersof the species in question are sufficiently distincton opposite sides of
the gap to be recognizedas separate varieties,this termbeing used in this paper
to denote a morphologicallyrecognizablesubspecificgroup with its own distri-
bution pattern. The area occupied by Mll.gonavensis,althoughnot distantfrom
the range of M. spathulata var. brachypoda,mightbe considereda fifthisolated
area; even here the few specimensavailable suggesta differentiation betweenthe
island and mainland memiibers. The effectof isolation is thus well illustrated,
as is the axiom that a group's closestrelativeis usually not sympatricwith it.
The geologicalhistoryof the regionhas probablyplayed a large part in bring-
ing about the presentdistributionpattern. Section Nesophytutm is restrictedto
the GreaterAntilles,islands whichat one time were all connectedby land with
each other as well as with Central America (Schuchert). The Lesser Antilles
were not connected either with each other, with the Greater Antilles, or
with South America, according to the same author. The absence of species of
Nesophyttum from any islands not once connected by land with the Greater
Antilles,as well as fromseveral nearby islands of the latter archipelago itself,
suggeststhat theirmeans of over-watertransportis poor at best. It is possible
that this distributionis simply due to an inabilityof these plants to grow else-
where,but in view of the generallysimilar clinmate throughoutthe area and the
wide range of soil toleranceof the plants this explanationseems to me unlikely.
If one assumes the absence of over-watertransport,approximatelyequal rates
of divergentevolution,and the absence of major isolating mechanismsother
than geographical, then the present distributioncould be accounted for by
changesinvolvingeitherland or water barriers. The presentgeographyand the
geological historyboth suggest that the latter means has been of primaryim-
portance,altlhoughit does not appear that such changes can have been solely
1957] MORLEY: IMOURIRI 119

responsiblefor the presenitdistribution.Certain minimumsteps in the geologic

historywould have been necessary to account for the present distributionof
section Nesophytum: (1) a period during which Cuba, Hispaniola, and Puerto
Rico were all connected,the ancestorsof M. spathulata became establishedin
northernHispaniola, and the ancestersof the M. helleri-M. emarginita group
became spread over the entirearea except perhaps for the northernhalf of His-
paniola; (2) an early separationof Cuba fromHispaniola; (3) submergenceof
all but west and east Cuba, and separation of Hispaniola from Puerto Rico;
(4) reelevationof centralCuba. Periods similarto thesesteps are already estab-
lished on a geologicbasis, as illustratedby Schuchert (Historical geologyof th.e
Antillean-Caribbeanregion, pl. 11-16), beginninog with the Lower Oligocene,
althoughwith interveningchanges that could not be indicated by the present
distributionof section Nesophytum. Whether or not the early spread of the
ancestorsof the section actually occurred as far back as the Oligocene is of
courseunanswerable. In addition to the steps listed above, the Nesophytumdis-
tributionsuggeststhat northHispaniola may once have been separated fromthe
rest of that presentisland; that Hispaniola may have been joined to east Cuba
at a timewhenthe latterwas separated fromotherparts of Cuba; and that the
southwestpart of Hispaniola may once have been separated fromthe rest of that
island (at present,M. goinavensisis apparentlyisolated partly by a geographic-
environmentalbarrier,and the initial isolationmighthave been of this nature).
Of thesepossibilitiesonly the last is shownby Schuchert,as existingboth in the
Middle Oligoceneand the Lower and Middle Miocene. It is noteworthythat the
area now occupied by M. spathulata var. spathulata correspondsrather closely
withthe emergentpart of eastern Cuba during the Upper Pliocene and Pleisto-
cene, as shownby Schuchert. PresumablybothM. spathulata,and M. ema,rginata

300 2
miles

C \

26 4} 2

*O 0

28

FIGS. 25-28. Geographic distribution of the species of Mouriri sect. Nesophytum; legend
in fig. 1. FIG. 25. Relative positions of the three islands involved. FIG. 26. Cuba. FIG. 27.
Rispainiola. FIG. 28. Puerto Rico.
 1120 BRITTONIA [ VOL. 9

were onithis islanidarea, and ornlyM. emaryitnata succeeded in spreadiingfar

fromthe originalarea afterrecessionof the seas. Much of the above is specula-
tion,but nonethelessthe distributionof the sectionNesophytumand the sequence
of changes it suggestsadd their small bit to the vast amount of biogeographic
evidencebearing on the geological historyof the area, and may play a part in
furtherelucidatingits details.
Habitat requirementsof species of section Nesophytunm appear to be froml
verybroad to fairlyrestricted.In Cuba, bothMoloririemnarginata anidM. spathut-
lata are found fromthe low scrub forestsand thicketsthat oceur on poor soil
to the montebravo and montefresco; the latter termsare those used by E. E.
Smith (The forests of Cuba. 1954) to designiaterespectivelyrelatively slow-
growingforestsin areas of mediumn rainfallor rainfall effectiveness,anldrapidly
growingforestsin areas of high rainfall. In addition Mfouriri spathutlataoccurs
in the pine lands, near Moa and Punta Gorda. Both these species are found at
elevationsfromsea level to at least 800 In; judging by Chamberlin's rainfall
nmaps(Month. Weath. Rev. 68: 4-10. 1940), they grow in areas of about 45-65
inehesannual rainfall. In Hispaniola and Puerto Rico the species presentappear
to be mostlyrestrictedto the "moist" forests,as designatedby Holdridge (in
Verdoorn,F. Plants aindplant science in Latin Amnericca 76-78, 81-83. 1945).
In Puerto Rico, at least (Lobeek, Sci. Surv. Porto Rico & Virgin Is. 1: 301-379,
1922), and apparently in Hispaniola (Hydrogr. OfficePubl. 530. 1947), this
meansan annual rainfallof about 70-110 inehes. In Hispaniola, M. helterigrows
in the northeasternpart of the island where the moist belt comes close to sea
level, and is found fromsea level to 400 mnelevation; M. spathtlata, running
almostthelengthof the island,is reportedfrom200 to 700 m; and M. gonacvensis,
in the southwestwherethe moistzone is apparentlyhigher,from700 to 1100 m.
In Puerto Rico M. helleri has been collected fromsea level to an elevation of
about 300 m.
Rock sources of the soil vary greatlyin the range of the sectioln.Limestone,
nlietamorphic rocks,serpentinecontainingup to 80-per cent magnesiumsilicate
and 5-6 per cent iron, and probablyothersare found. Soil type seems to have
littleor no effecton distributionof species of sect.Nesophytrni,as in manyforest
species of these regions. In Cuba there is a suggestionthat a differencein soil
tolerance may exist between M. emarginata and MI. spathutlata. Both species
occur on calcareous and othersoils, but only M. emarginata.has been reported
on serpentinebarrensor on little-modified serpentinesoil, whereit can be found
in both easternand westernCuba; and only M. spathtlata is knownon the red
serpentine-derived laterite soil of northeastOriente which contains as low as
2 per centmnagnesium silicateand up to 52 per centiron,althoughM. entargitnata
does grow in a pocket of red soil of probable limestoneorigin on a limestone
mountain in Pinar del Rio. The evidence is inconclusive; data on labels are
inadequate, and detailed informationis hard to get. For Hispaniola and Puerto
Rico available information is completelyinadequate to detectany soil preferences.
Regardingpatternsof variationin the section,it may be said in general that
the species of the sectionare ratherwell distinguishedfromeach other,and that
variabilityis great, although probably not unusually so. Most of the variants
appear to be withina normalvariationrange,and in fact onlyone givessomesug-
gestionof possiblehybridizationbetweenadjoining species. Other than this pos-
sible hybridonlyone specimenis so unusual as to be difficult to place, and thatis a
sterilecollection,EkmtanIf10601, named by Urban M. plinioides,and discussed
underM. gonavensis. Only furthercollectingcan resolvethe problemit presents.
1957] MORLEY: M11OURIIsI 121

TAXONOMIC TREATMENT

Mouriri sect. NesophytumMorley,Univ. Calif. Publ. Bot. 26: 277-278. 1953.

Yotunigfirstyear twigs rather quadranguLlar, the angles smoothto conspicu-
ously wing-lined,the twigs soon roumdingand the wings disappearing; leaves
small, usually less than 8 cm long, rarely (in M. spathulata) to 10.5 em lolng;
blades rarelylanceolate or ovate, usually ovate-ellipticto obovate,the base nar-
rowlyto broadlyacute or attenuateto the petiole,the apex sometimesacute, usu-
ally roundedto emarginate,oftenmucronuLlate; midribnot groovedabove; upper
surfaces of dried leaves snmooth to minutelyrugulose with irregular ridges to
punctulatewith tiny raised dots caused by the foliar selereidswithiln;stomatal
cryptsusually type III but when very small type II, sometimesabsent; upper
epidermissingleor irregularlydoubled,countingas double only directlyopposite
pairs of cells; innerwalls of lowerand upper epidermalcells sometimesmucilage-
thickened; hypodermispresen-t,without mucilaginous walls; foliar terminal
selereids type I, IT, or 113. Infloreseences1-5 in the leaf axils, each a small
opposite-branched racemewith1-3 joinltsin the centralaxis (inclludingthe joint
at the pedicel base of the terminalflower) and 1-7 flowers,each lateral flower
terminalon a distinctbranchthe end of whichis markedby anlarticulationand
two bracts; bracts deciduoLus, each leaving after falling a tiny crescent-shaped
sheath; infloreseenceaxes, pedicels, anid flowersglabrous or minuLtely puLbern-
lent, the puberulence centeringo on the pedicel and extendingmore or less to
adjacent parts. Calyx-lobesfree early in the bud or connate to 23 their length
and separatingat anthesis; anthersaverage in form (for Molriri), 1.75-2.7 mm
long; ovary 1-5-locular,7-18-ovuled; placentationaxile-basal to axile, the ovules
all at one level.

Key to Species of Mouriri section Nesophytum

Length of calyx in total (including the inferior ovary) 5.8 mm or more.
Petioles usually winged, 2-8 mm long, usually 0.1 or more the length of the blade;
calyx-lobeshalf elliptic to triangular; Hispaniola. M. gonavensis.
Petioles not noticeably winged, 0.5-3.5 mm long, 0.05 or less the lelngthof the blade;
calyx-lobes triangular to acuminate; CuLba. M. emarginata.
Length of calyx less than 5.8 mm.
Fusion distance of adjacent calyx-lobes,measured from the upper edlge of the stamen
attachment,more than 1 m-m(the lobes separating at anthesis); Cuba. M. emarginata
Fusion distance of adjacenit calyx-lobes1 mm or less.
Length of calyx less than 5 mm; calyx-lobes less thani1.7 mm lonig; fusion distance
of calyx-lobes 0.6 mm or less.
Leaf-blades not more than 3.7 cm long; Hispaniola and Puerto Rico. M. helteri.
Leaf-blades mostly or at least somreof them more than 3.7 cm long; easterlnCuba
and Hispaniola. M. spathmlata
Lenigthof calyx 5 mm or more; calyx-lobes 1.7 mm long or more; fusion distance
of calyx-lobesmore than 0.6 mm.
Ratio of length of calyx to width of calyx-lobe less than 2.6; ratio of length of
calyx-lobe to width less than 0.85.
Leaf-blades not longer than 3.7 cm; Hispaniola and Puerto Rico. M. helleri.
Leaf-blades mostly or at least some of them longer than 3.7 cm; easterniCuba.
M. spathutlatavar. spathulata.

Albl.
3 Foster, A. C. Comiparativemorphologyof the foliar selereids in the genus Montririaz
Jour. Arnold Arb. 27: 253-271. 1946.
 122 BRITTONIA [VOL. 9

Ratios more than those given above.

Leaf-blades not longer than 3.7 cm; bracts of the inflorescenece sonmetimes per-
sistent to the fruiting stage; fusion distance of calyx-lobes less than 0.7
mm; ratio of fusion distance to width of lobe less than 0.45; calyx-lobes
1.7 mm long or less; Hispaniola and Puerto Rico. M. helleri.
Leaf-blades most'y or at least some of them more than 3.7 em long; bracts of
the infloreseenceearly deciduous; fusion distance of calyx-lobes 0.7 mm
or more; ratio of fusion distance to width of lobe 0.45 or more; calyx-lobes
1.7 mm long or more; western Cuba. M. emarginata var. rostrata.
1. Mouriri spathulata Grisebach.
Small tree to ca. 10 m high and 10 em in diameterat breast height; older
individuals commonlywith a strongbasal shoot,sometimesbecomingsomewhat
shrubby,oftensproutingfromthe stump; bark thin,light to dark brown,finely
checked vertically with fewer cross-checks;angles of young first-yeartwigs
sometimeswinged; petioles 1-7 mm long; blade coriaceous,rarely lareeolate to
ovate to elliptic or rhombie-ellipticto obovate,3-10.5 cm long, 1.5-5.5 cm wide;
marginoftenrevolute,the base cuneate, attenuateto the petiole,the apex nar-
rowlyacute to broadly rounded to slightlyemarginate,sometimesmucronulate;
midribusually roundedbelow,sometimesrectangularat the middle,very rarely
with tiny incompletewings; lateral veins visible above and below in very large
leaves, bothfreshand dried,usually visible only above; youngleaves when fresh
a smooth,glossy green above, pale green and scarcely shining below; older
leaves less shinyabove and dull below; dried leaves sometimesshiny above, dull
below,thesurfacessmoothor punctulate; stomatalcryptswhenregularlypresent
30-170/mm2, theircavitiesaveraging30-65t in diameter;centralbodies of foliar
terminalselereidstypicallyelongate,2-4 times as long as wide, but sometimes
isodiametric;upper epidermispredominantlysingle, rarely with up to 40 per
cent of the cells doubled,the inner walls of the epidermalcells never mucilage-
thickened; fresh leaves withoutaromatic odor when crushed,faintly speckled
when held to the light. Racemes 1-5 per leaf axil, 2-26 mm high to the ovary
base of the terminalflower,the joints in the central axis 1-3, the flowersper
raceme1-7, the bractsat the joints (rarely one also at the ovarybase) triangular
or ovate-triangular, acute to acuininate,1.0 mm long, soonldeciduous; lengthof
the true pedicel (ovary to firstjoint below) 1.5-6 mm. Inferior ovary 1.2-1.7
mm high, campanulate to cup-shaped or approachingturbinate,the angle "o"
(fig.6) 1200-1550;lengthof the calyx (includingovaryheight) 3.7-5.6 mm (dry
measurements3.5-5.1 mm); ratio of lengthof calyx to width of calyx-lobe1.5-
3.1; calyx-lobes0.9-2.1 mm high fromthe stamen attachment,1.5-4 mm wide,
0.4-1.05 times as high as wide, truncate with rounded shoulders to broadly
roundedto low-triangularto rarely minutelyapiculate, the shoulderssometimes
expanded into rounded lobes (figs. 10, 11), the marginsminutelyfringedwith
shorthairs; fusion distance of adjacent lobes above the stamen attachment0.4-
0.9 mm,the ratio of fusioll distance to width of lobe 0.15-0.45; petals yellow,
white,pink,or violet,broadlyovate,oftencrinkledbelow,5-8 mm long, 4-7 mm
wide,the base roundedto truncateor slightlycordate to a shortclaw 0.8-1.0 mm
long and 1 mmwide,the apex acute; filamentswhite; pollen sacs yellow; anthers
1.75-2.5 mm long; stamensand style oftenslightlyzygomorphic,most stamens
orientedwiththe gland upward; ovary (2-) 3-5-locular,9-18-ovuled; ripe fruits
yellow to red-orange,1-4-seeded, approximatelyglobose when 1-seeded, then
12-15 mm in diameter,lobed accordingto the numberof seeds, up to 30 mm in
diameter with 2-4 seeds, their dry measurements3-4 mm less; fruit thinly
fleshed,withslightlysweet,insipid flavor.
1957] MORLEY: MOURIRI 123

la. Mouriri spathulata var. spathulata.

Motriri spathulata Griesbach,Mem. Am. Acad. 11. 8: 183. 1861.
M"riri lanceolata Grisebach,Mem. Am. Acad. II. 8: 183. 1861.
Aulacocarpus wrightiiGrisebach,Cat. P1. Cub. 90. 1866.
Mowtriri snaestralisUrban, Symb. Ant. 9: 126. 1923'.
Stomiatalcryptswhen regularlypresent 30-115/mm2,the eavity averagilng
34-65k,in diameter; raceme to the ovary base of the terminalflower5-26 mm
high; flowersper raceme 1-7; lengthof the true pedicel 2-6 mmll;lengthof the
calyx 4.2-5.6 mm (dry measureluents3.7-5.1 mm); ratio of length of calyx to
widthof calyx-lobe1.5-2.5; ratio of width to heightof ovary (not dried) 1.35-
1.7; locules (2-)3-5; ovules 9-18.
Type collectionand locality: Wright1234, Cuba, Orienteprovince,La Perla,
propevillamMonteVerde dietam,May 28, 1859. (Other Wrightcollectionsnuzm-
bered 1234 exist, some from Monte Libano and others without data. The La
Perla locationis specifiedby Grisebach. For a briefreviewof Wright's localities
in easterlnCuba, see Jervis,Asa Gray Bull. II. 2: 29-40. 1953). Holotype at
GOET.
Distribution: southeasternhalf of the provilce of Oriente in eastern Cuba
(fig.26).
Representativespecimenis:CUBA: Linden 2147 (paratypeof M. spathulata; NY); La
Perla, propevillamMonteVerde dietam,Wright 1234 (isotypes: GH, MO; GOET specimen
not seen); prope villam Monte Verde dietam, at the farallons, Wright 12?35 (type collection
of M. lanceolata: GH, MO; GOET not seen); Farallon de la Perla, Shafer 8796 (NY); Monte
Libano, Wiight 1234, 1860-1864 (two writtenslips in pocket, one describing floweringmaterial
and dated Mav 2, the other describing fruiting material and dated Apr. 30; GH); Wright
1234, 1860-1864 (S); Wrtight 1234 (NY); Guant'anamo, in monte Libanon, near Monteru's,
ca. 7-800 in, Ekman 15810 (NY, S, US); Valparaiso, Monte Libano, Wright 2465, 1860-1864
(type collection of Aulacocarpits wrightii; GH, GOET not seen); east of Puinta Gorda Ca. 1/3
of way to Alto Quemado Negro from Rio Cayoguan oii road, Morley,881 (MIN): Woods,
Delta Mine, near Cayoguan, Moa, Victorin, Clemente, 4 Alain 21830 (LS); east side of Rio
Yagrumaje on road from Moa to Puiita Gorda, Morley 891 (MIN, US); between Moa and
Yagrumajes, Aug. 30, 1917, Roig 61 (NY, ROIG, SV 6565); Gran Tierra, Moa, Acwiah,April
16, 1945 (NY, ROIG 8792, SV 12603); Moa, chemin de Cayo Chiquito, Clemente 3652 (GH,
LS, MIN); new road to La Brefia, Moa, Clemente & Chrysogone4422 (LS, MIN); near air-
strip, Moa, Clemente 3546 (LS, MIN); Playa de Moa, Le6n, Clemente 4f Howard 20271 (LS,
MIN); near the village of Moa, Howard 5933 (GH); ca. 1/2 mile east of the Moa pier, ca.
200 yds from ocean, Morley 888 (MIN, US); Playa La Vaca, Moa, Clemrnente & Chrysogone
6140 (LS, MIN); El Purio, Cabonico, Roig, Sept. 15, 1917 (SV 6633); Levisa, Roig, Sept. 15,
1917 (ROIG 1635); Sierra de Nipe, Ekman 6031 (S); Sierra de Nipe ad Loma Mensura, ca.
725 m alt., Eklman 9907 (NY, S, US); Palma Soriano, Curbelo, June 2, 1932 (NY, ROIG
6232); Sierra Maestra ad Manacal, 8-900 m alt., Ekman,9350 (type Collectionof M. maestralis;
NY, S, US; photo of NY specimen,LS); Sierra Maestra, Santiago de Cuba, Turquino region,
'near Cueva del Aura," Bitcher 174 (ROIG 5317, US).

lb. Mouririspathulatavar. brachypoda(Urban & Ekman) Morleycomb. nov.

MouriribrachypodaUrban & Ekman, Ark. Bot. 22A(17): 67. 1929.
Stomatal cryptswheii regularlypresent 90-170/mm2,the cavity averaging
30-331uin diameter; raceme to the ovary base of the terminalflower2-9 mm
high; flowersper raceme1-3; lengthof the true pedicel 1-4.5 mm; lengthof the
calyx ca. 3.7-5.0 mm (drv measurements3.5-4.7 mm); ratio of length of calyx
to width of calyx-lobe2.1-3.1; ratio of width to heigfhtof ovary (not dried)
1.15-1.45; locules 2-4; ovules 9-12.
Type collectionand locality: EkmnanH6064, Haiti, Massif du Nord, Hinche,
betweenCerea-Carvajal and Bois-Charles,at Riviere Samana, ca. 500 m, metam.
rocks,May 12, 1926. Lectotypeat S.
 124 BRITTONIA [ VOL. 9

Distribution: Northernhalf of the island of Haiti, fromn1earGonaives to

the vicinityof the Bay of Samana (fig.27).
Specimens examined: HAITI: Dept. de L'Artibonite: Massif des Cahos, Hinche, Morne
Vaillecite, 700 m, Ekman f6123 (paratype:S). Dept. du Nord: Massif du Nord, Hinche, be-
tween Cerca-Carvajal and Bois-Charles, at Riviere Samaana, Ekmnan f6064 (A, NY, S-
type, US).
DOMINICAN REPUBLIC: Libertador: Cordillera Central, prov. de Monte Cristi, Res-
tauraci6n, at Rio Gurabo, ca. 525 m alt., Ekinan f6257 (paratypes: A, S). Santiago Rodri-
guez: Sabana de Inaje, Carlos Gonzales 2042, April 17, 1950 (JIM); Moncion, at Rio Mao,
ca. 200 m, Ekman f12650 (S, US). Santiago: San Jose de las Matas, edge of Rlo Inoa, ca.
350 mn,EkmnanH14610 (S). Samand: Peninsula de Samaana, Sanchez, ca. 300 mn,Ekman
f14694 (NY, 5); Cordillera Central, Los Haitises, La Llanada, Ekinan f15480 (A, NY, S).
A close scrutinyof this well-distingauished but highlyvariable species shows
the presenceof two subgroupsthat can be accordedno morethan varietal status.
Only one clear-cut distinctionsets the two varieties apart, the character of
stomatal crypt size-frequency;otherwisethe effectsof the sharp isolation are
seen onlyin the different but overlappingranges of variationof the various char-
acters indicated. In the Cuban variety, certain tendencies can sometimesbe
associatedwith certainareas; for example,the plants of the northcoastal region
and nearbymountainstend to have relativelylarge leaves, and these are thicker
than is usual in the Moa and Punta Gorda area; leaves from the vicinity of
Punta Gorda in particulartend to have a rathercharacteristienarrowlyrhombic-
elliptic form; and not far to the south of the latter area a lanceolate leaf is
apparently common. This variation appears very irregular,however,with no
clear separationsand no furtherbasis for segregation.In the Punta Gorda area
whereI have seen them,someindividualshave a fairlyconsistentrhombie-elliptic
leaf; otherindividualsnearbyhave broad, elliptic-obovate leaves; and still others
have all possibleformson the same plant. Othercharacters,such as those of the
infloreseenceand flower,also appear to be nearly random in variation. In the
Hispaniolan varietythereis also considerablevariation,but the limitednumber
of collectionsdoes not allow accurate assessmentof it. So far as I can tell, the
great variabilityof M. spathulcatais all self-generated.In the Cuban collections
I have seen no specimensthat suggest interbreedingwith the sympatricspecies
M. emarginata,even thoughthe two sometimesoccur in the same locality. In the
Dominican Republic one sterile collectionof M. helleri var. samanensisis sug-
gestiveof M. spathulata in formand size of leaves. The influenceof any cross-
ing that may have occurred does not appear to have passed in the opposite
direction.
2. Mouriri helleri Britton.
Small to mediumtree,sometimesshrub-like;angles of young first-year twigs
narrowlybut prominentlywinged; petioles 0.5-2 (-3) inm long; blade ovate to
narrowlyor broadly elliptic or almost round to obovate,1.0-t.4 (-3.7) cm long,
0.5-1.9(-2.4) cm wide (the parentheticalmeasurementsall refer to Ekman
115047); margin oftensomewhatrevolute,the base narrowlyto broadly acute,
oftencuneate,the apex narrowlyto broadly acute (the acutenesssometimesac-
centuatedby inrolled margins on both sides) to rounded to emarginate,often
mucronulate;midribrounded to flat below, rarely (Ekman H15047) somewhat
rectangularin themiddle,not winged; lateral veinsnot visible,or veryindistinct,
above and below; upper surfacesof dried leaves smoothto minutelyruguloseor
less oftenpunctulate,the lower surfaces smoother;stomatal cryptswhen regu-
larly present 30-85/mm2,their cavities averaging 40-60, in diameter; central
1957] MORLEY: MOURIRI 125

bodies of foliar ternminalselereidswhen elongate up to 3 times as long as wide,

the armsusually prominent,theirlengthup to twieethe body thickness,5--40per
cent of the selereidshaving the body very shortand with a stellate appearance;
upper epidermispredominantlysingle, sometimesup to 6 per cent of the cells
doubled, the inner walls of the epidermal cells not mucilage-thickened(a few
doubtfulinstancesseen in Abbott2233). Racemes 1 per leaf axil, 5-12 mmhigh
to the ovary base of the terminalflower;joints in the central axis 2, the flowers
per racemie1-3, the bractsat the joints ovate-trianoular,acute, 0.8-1.3 mm long,
deciduous early or late, oftenpresenltwhen the fruit is imature;length of the
true pedicel (ovary to firstjoint below) 1.5-3.5 mm. Inferiorovary ca. 1.6 mm
high, turbinate,the sides only slightlyconvex, angle "o" (fig. 6) 1600-1640;
lengthof the calyx (includingovaryheight) ca. 5.3 fmnr; ratio of lengthof calyx
to widthof calyx-lobeca. 3.15; calyx-lobesca. 1.0-1.7 mm high fromthe stamein
attachment,ca. 1.6-2.1 mm wide, 0.5-1.0 times as high as wide, the free ends
triangularor broadlyso, acute (fig.17), the marginsminutelyfringedwithshort
hairs; fusiondistanceof adjacent lobes above the stameenattachmelent 0.4-0.7 mm,
the lobes usually separating at anthesis; ratio of fusion distance to width of
lobe 0.2-0.4; maturepetals unknowii,the immatureones ovate-triangular, acute,
4 mmlong, 3 mmwide, the base truncateor with a claw ca. 0.3 mmlong and 1.5
mm wide; anthers2.25-2.45 mm long; ovary 2-locular,7-9-ovuled; ripe fruits
orange,1-4-seeded,more or less globose when 1-seeded,their dry measurements
then ca. 5-8 mm in diameter,5-9 mm long, lobed according to the numberof
seeds,up to ca. 10 mm or morethickwhen morethan 1-seeded.
2a. Mouriri helleri var. helleri.
MouririhelleriBritton, Torreya 2: 10. 1902.
Calyx-lobes1-1.3 mmhigh,ca. 2.15 mmwide, averagingca. 0.56 timesas high
as wide; fusiondistanceof adjacent calyx-lobesabove the stamenattachmentca.
0.4 mm; ratio of fusion distance to width of lobe ca. 0.2.
Type collectionand locality: Heller 1372, Puerto Rico, Cataio, in sandy soil
near a mangroveswamp, Mav 23, 1899. Holotype at NY.
Distribution: Puerto Rico (fig.28).
Specimens examined: Catafio, in sandy soil near a mangrove swamp, Heller 1372 (F, NY-
type); prope Hatillo in scopulosis sylvae primaevae ad Guajanes, Sintenis 6195 (paratypes:
A, F, GH, MO, NY, S, US); Maricao, Whetzel & Olive, March 22, 1916 (NY).
2b. Mouriri hellerivar. samanensis (Urban) Morley,comb.nov.
Moturiri samanensisUrban,Repert.Sp. Nov. 20: 342. 1924.
MouriribuxifoliaUrban, Ark. Bot. 24A(4): 31-32. 1932.
Calyx-lobes1.3-1.7 mmhigh,1.6-1.7 mmwide, 0.8-1.0 timesas high as wide;
fusiondistanceof adjacenltcalyx-lobesabove the stamenattachmenlt 0.6-0.7 mm;
ratio of fusiondistanceto widthof lobe ca. 0.4.
Type collectionand locality: Abbott2233, Domillican Republic, San l:orenzo
Bay and vicinity,south coast of Samanal Bay; sea level; April 26, 1922. Lecto-
type at US.
Distribution: Dominican Republic, Sarnana Peninsula and south coast of the
Bay of Sanmana,and near the south coast of the province of Trujillo Valdez
(fig.27).
Specinmensexamined: Samana: Peninsula de Samana, Cabo Cabron, light forest at La
Herradura, Ekman H15047 (NY, S); Cabo Saman'a, in forest, Ekman H14895 (A, NY, S);
slope of Pan de Azuecar,ca. 400 m, EkrmanH15180 (A, NY, S). San Lorenzo Bay and vicinity,
south coast of Samana Bay, sea level, Abbott 2233 (GH, MO, NY, US-type); C'ordillera
Central, Los Haitises, La Llanada, rocks at sea level, Ekrman H15497 (S). Trujillo Valdez:
Llano Costero, prov. Saiito Domiiigo, El Manielito, hillsides, Ekman H11295 (type collection
of M. buxivfolia:A, NY, S).
126 BRITTONIA [ VOL. 9

The outstandingcharacteristicof this species is its general smallness,espe-

cially of leaf but also of flower. The paucity of floweringmaterial has been a
serious drawbackto this study; however,thereis reasonable coneurrenceamong
the limitedmaterials available, so that the chances seem good that it is fairly
representative.Motriri helleri is the only species in the section combiningthe
short calyx-lobefusion distance characteristicof M. spathulata with the high
angle "o" and relativelylarge ratio of calyx-lengthto widthof lobe of M. gona-
qensis and M. emnarginata.Otherwiseits measurementsare mostlyintermediate
betweenthoseof M. spathulactaand the othertwo species. Variation in the group
is not unusual except apparently in some parts of the Peninsula of Samana.
A niote,evidentlyEkman's, appended to his collectionH15180, says, "Somewhat
doubtful. The Mouririae of Samana are very difficult." This particular speci-
men appears to me to be withinthe normalrange of variation of the species,but
his sterilecollectionH15047 fromCabo Cabro6nhas a larger and moreacute leaf
than usual, one that is suggestiveof MT.spathulata var. brachypoda. Of this
specimellEkman says, "Mouriria samanensisgrew togetherwith this, and ap-
peared to be distinct. I saw, however,only a few trees of this." The stomatal
cryptfrequencyand size of H15047 are thoseof M. helleri. The nearesteollection
of M. spalthulatacomes fromabout 25 mailesaway, although the species might
easily occur closer. There is no wav of furthercheekingthe source of variability
AwTithoutmore field work.
Concerningthe two varieties of 1'. helleri, it must firstbe noted that no
floweringmaterialhas been seen of M. hellerivar. helleri,and that it is therefore
possiblethat the Puerto Rican plants are more distinctthan is indicated by this
treatment. However, their very great similarity to the eastern Dominican Re-
public plants in bothvegetativeand fruitcharactersleaves no doubt in my mind
as to their close relationship. Quite possibly not even varietal status is war-
ranted.

3. Mouriri gonavensisUrban & Ekman.

Smiiallto mediumtree; angles of younc first-year twigswith or withoutnar-
row wings; petiolesusually winged,2-8 mmlong, usually 0.1 or more the blade
lengthwhenmeasuredto the abrupt curve at the blade base; blade ovate-elliptic
to narrowlyor broadlyellipticto slightlyobovate,2.5-6.5 cm long, (1.0-) 1.4-3.8
cm wide, usually mnore thall half as wide as long; margin oftensomewhatrevo-
lute, the base usually abruptlyaeuminateinto the winged petiole,the apex nar-
rowly to broadly rounded to broadly and slightlyemarginate,sometimesmu-
cronulate; midrib below flat to rounded to rectangular,often the two angles
withverynarrowwings; lateral veins sometimesvisibleabove and below in dried
material; dried leaf-surfaceabove minutelyrugulose or rarely punctulate,the
lower surface smoother; stomiatalcrypts when regularly present 25-50/mm2,
their cavities averaging 55-60,uin diameter; central bodies of foliar terminal
selereidswhen elongate2.5-3 timesas long as wide, theirarms prominent,their
length0.5-1.5 timesthe body thickness,5-40 per cent of the selereidshaving the
body veryshortand with a stellateappearance; upper epidermispredominantly
single,rarelywith up to 20 per cent of the cells doubled,the inner walls of the
epideriilalcells nevermucilage-thickened.Racenles 1 per leaf axil, 9-25 mmhigh
to the ovarybase of the terminalflower,the joints in the centralaxis 2 or 3, the
flowersper racemne 1-3(-6), the bracts at the joints ovate-triangular,acute, 1.5
nmm long, soon (leciduous; length of the true pedicel (ovary base to firstjoint
 1957] MORLEY: MOURIRI 127

below) 2.5-5.5 mm. Inferior ovarv 2-2.6 mm high, turbinate,the sides only
slightlyconvex,angle "o" (fig. 6) 158'-166' or more; lengthof the calyx (in-
eludingthe ovaryheight) ca. 6.5 mm fresh,6.0 mm dry; ratio of lengthof calyx
to width of calyx-lobe3.2-3.6; calyx-lobes1.7-2.5 mm high fromthe stamen at-
tachment,1.7-2.1 mmwide, 1-1.2 timesas long as wide, the free ends triangular
to half-elliptic(figs. 15, 16), rarely almost truneate,the margins fringedwith
shorthairs; fusiondistanceof adjacent lobes above the stamenattachment0.85-
1.0 (av. 0.95) mnm, the lobes separating at anthesis; ratio of fusion distance to
width of lobe 0.45-0.57; petals white to pale pink, purple on the midrib,ovate-
cordate to broadly oblong to orbicular,6-7 mm long, 4.5-6 mm wide, the base
witha claw to 1 mmlong and ca. 0.7 mmwide, the apex apiculate or shortlyand
abruptlyacuminate,the marginirregularlycrisped below; anthers2.45-2.7 mm
long; ovary2- or 3-locular,8-11-ovuled; fruitunknown.
3a. Mouriri gonavensisvar. gonavensis.
Mouriri gonavensisUrban & Ekman, Ark. Bot. 22A(10): 102, 103. 2 June
1929.
Wing-lineson youngtwigs absent or barely visible; midribbelow roundedto
flat,onlyrarelywithtraces of wing-margins;heightof infloreseence to the ovary
base of the terminalflower9.5-16 mm; calyx-lobeshalf-elliptic(fig. 15), rarely
almosttruncate,2.1-2.5 mmlong.
Type collectionand locality: Ekman H8712, Haiti, Insula Gonave, Morne
Mouri-Corps,patch of forest,700 m, July28, 1927. Lectotypeat S. (Other speci-
mens examinedat NY, US.)
Distribution: Known onlv fromthe type locality on the island of Gonave,
Haiti (fig. 27).
3b. Mouriri gonavensisvar. hottensis(Urban & Ekman) Morley,comb nov.
MouririhottensisUrban & Ekman, Ark. Bot. 22A(17): 67, 68. 19 N 1929.
MouririplinioidesUrban, Ark. Bot. 24A(4): 32. 1932.
Wing-lineson young growingtwigs conspicuous; midrib below rectangular,
prominent,with or withoutnarrowwings at the angles; height of infloreseence
to the ovary base of the terminalflower11-25 mmn;calyx-lobestriangular (fig.
16), 1.7-2.0 mm long.
Type collectionand locality: Ekman H10399, Haiti, Massif de la Hotte, west-
ern group, Jeremie,on the ridge between LopineauLand M. Pain-de-Suere, ca.
1100 m, July 22, 1928. Lectotypeat S.
Distribution: Haiti, Dept. du Sud, westernhalf of Massif du Sud (Massif de
la Hotte; fig.27).
Specimensexamined:Les Roseaux,Nan-Patales,in forest,1000 m,EkmanH10711 (para-
type: S); Jeremie,ridge toward Morne Pain-de-Sucre, 1100 m, Ekman 110399 (A, NY, S-
type,US); Tiburon, Morne Sentier, ca. 800 m, Ekman H10601 (type collection of M. pUniioides;
NY, S).
This species appears closely related to both M. helleri and M. emarginata,
and servesas an intermediatebetweenthe two. Mouriri gonavensisis primarily
distinguishedfromthe formerby larger leaves, greaterinfloreseenceand flower
measurements,and bv the promptlydeciduous bracts of the infloreseence;from
the latter it differsmostlyin its single epidermis withoutmucilage-thickened
walls, its longer petioles,a combinationof calyx-lobefusion distance plus angle
"o,'" and its range of calyx-lobeshapes. The similarityof M. goncvensisto M.
helleriis particularlygreat,mostof the differenlees
betweenthe two being only in
size. Within these two species, the varieties M. gonavensis hottensisand M.
helleri sanavensis approach each other the most nearly. The sterile collection
128 BRITTONIA [vOI,. 9

Ekman H10601, nlamedby Urban M. plinioides,seemsto representan -unusually

narrowleaf in M. gonavensiswhich closelyresemblesleaves of M. helleri. Were
it not for the winged midriband the lengthof leaves in this collectionit might
easily be taken for a typical collectionof M. helleri var. samanensis. However,
differencesbetween these two groups are sufficiently great to make it appear
unwise to place both under one specificname, even though they are evidently
more closely related than other species of the section. Variability is hard to
judge in M. gontavensig, owingto the very small numberof collectionsavailable.
One can only hope that the two extant collectionswith flowersare reasonably
representativeof that structure. Leaf formis fairly consistentin three speci-
mens,in one (Ekmnan H10399) rathercloselyresemblingthat of M. emarginatas;
but the fourthcollection,Ekmman f10601, differssharply, as described above.
However,close examinationof Ekman H10711 shows similar tendencies,so that
I considerit probable that Ekma,nH10601 is no more than an extremevariant
of the same species. Further collectingis needed to verifythis point. Regarding
the two varieties of the species, it can only be said that the existing material
suggeststhe presenttreatment.The differences betweenthe two appear in some
respectsratherdecisive; a measure of support for the decision to considerthem
as one species is found in the fact that the charactersin question have similar
ranges of variationin anotherspecies,M. spathulata. I kniowof only one collec-
tion of M. gonavensisvar. gonavensis,and that may very well be the only one
ever to be made. In 1927 Ekman found the island of Gonave badly cut over,
with no sign of the abatementof destruction(Eyerdam, Field & Lab. 22: 85-
106. 1954).
4. Mouriri emarginataGrisebach.
Small to mediumtree,in favorablesituationsup to 16 m high andl20 cmlin
diameterat breast height,sometimessproutingfromthe stump,the bark brown,
finelycorrugatedvertically; young first-yeartwigs consistentlywinged at the
angles; petioles0.5-3.5 mmlong; blade coriaceous,ovate-ellipticto obovate,2.6-
8 cm lonlg,1.3-4.5 cm wide; marginsometimesrevolute,the base acute or attenu-
ate to the petiole,the apex rarelybroadly acute to roulndedto emarginate,often
mucronate;midribbelow rectangularor the two angles with very narrowwings
at least in the mid-region;lateral veins invisiblebelow,usually visible above only
in very large leaves; young leaves when fresha smoothvery shiny dark green
above,less shinyand lighterbelow; older leaves duller; dried leaves only rarely
shinyabove, dull below,the upper surface minutelyrugulose,the lower less so;
stolmiatalcryptswhen regularlypresent15-60/mm2, theircavities averaging40-
95/,tin diameter; central bodies of the foliar terminalselereids isodiametricto
twice as long as broad, sometimeslonger; upper epidermispredominantlydou-
ble, more than 40 per cent of the cells doubled, the ilnnerwalls of some of the
inner cells nucilage-thickenedas well as those of some of the lower epidermlial
cells, or only the latter so thickened; fresh leaves not aromatic whenicrushed,
faintlyspeckledwhenheld to the light. Racemes 1-3 per leaf axil, 6-18 mmhigh
to the ovary base of the terminalflower;joints in the central axis 1 or 2, the
flowersper raceme1-3 (-4), the bracts at the joints ovate-triangular, acute, 1-1.5
mnm lono, soon deciduous; lengthof the true pedicel (ovary to firstjoint below)
2-8 mm. Inferiorovary1.4-2.5 mlnhigh,narrowlycup-shapedto niearlyobconic,
the anglle"o" (fig. 6) 145?-168?; lengthof the calyx (ineluding ovary height)
5.0-6.7 mm (dry measurementsca. 4.8-6.5 mlim),ratio of length of calyx to
widthof calyx-lobe2.7-4.3; calyx-lobes1.7-3.0 mmhighfromthe stamenattach-
ment,1.2-2.3 mm wide, 0.85-1.9 timesas bigh as wide, the free ends triangular
 1957] MORLEY: MOURIRI 129

to apiculate to cuspidate to acuminate (figs. 12-14), the marginsminutelyand

irregularlyfringedwithshorthairs; fusion distance of adjacent lobes above the
stamelnattachment0.7-1.9 mm,the lobes separatingat anthesis; ratio of fusion
distalnceto width of calyx-lobe0.45-1.25; petals yellow,white,light purple, or
red, ovate to narrowlyovate-triangular, 4.5-6.5 mmlong, 3-5 mmwide, the mar-
gins oftenirregularlyand minutelycrenulate,the base truncateor with a short
claw ca. 0.7 mli wide and to 0.7 mm long, the apex acute to acuminate; anthers
1.95-2.5 mml7long; ovary1-3-locular,7-11-ovuled; fruitglobose,1-seeded [Urban
in his descriptionof M. rostrata (1926) described its fruits as having 1 or 2
seeds; I have been unable to verifythis],-itsdrymeasurements8-10 mmin diam-
eter,9-12 mm lolng,thinlyfleshed.

4a. Mouriri emarginatavar. emarginata.

MottririemarginataGrisebach,Cat. P1. Cub. 92. 1866.
Very narrow wings usually present at least in the center of the iimidribbe-
low; truepedicels 2-4.5 mmlong; lelngthof the calyx (ineludinginferiorovary)
ca. 5.5-6.7 mm (dry measurementsca. 0.2 mm less); length of calyx-limb(in-
cluding the free hypanthium) 3.9-4.9 mm; fusion distance of adjacent lobes
above the stamenattachment1.2-1.9 mm; angle "o" 158?-168?.
Type collectionand locality: Wright 2467, Cuba, Oriente provilnce,Mount
Friendship,1860-1864. Holotype at GOET.
Distributioll: Cuba, scatteredthroughthe provilnceof Orieliteexcept for the
east quarter, mostlyin mountainousareas, and west into Camaguieynear the
lorthcoast as far as a point northof the city of Camaguiey(fig. 26).
Specimens examilned: Oriente: Playa Vaca, Moa, Acultia,April 11, 1945 (SV 12601, UC);
Regi6n de la Sierra cle Cristal, Mayari, Charrascal del Saca Lengua, Alain, Acu,ia, & L6pez
F. 5407 (LS, MIN); Region de la Sierra Cristal, Mayari, woods, near Rio Miguel, Alain,
Acwita,4- L6pez F. 5921 (LS, MIN); Mount Friendship, Wright 2467, 1860-1864 (isotypes:
MO, GH, S; GOET specimen not seen); La Maya, Bucher, Oct. 1929 (ROIG 4928); Bayate
in forest,Ekman 6086 (S, US); Miranda, Cainizarez, July, 1949 (SV 15580); Corojo in jugo
mont. inter Pinar del Media et Treinta pinos (in sylva "Monte bueno-malo''), ca. 800 m alt.,
Ekman 5155 (S); Sierra Maestra, Pinar di Papayo, in forest near the pinelands, Ekman 9280
(NY, 8, US); Papayo (of the Sevilla tract), in the Mandinga hill, Ekman 9298 (S); Sierra
Maestra via Turquino, Acii4a, May, 1948 (two collections; SV 18815, 18816); Sierra Maestra,
on the ascent fromRRioYara called "Subida de Catasus", ca. 600 m, Ekman 14204 (NY, S);
Sierra Maestra, inter Punta de Palmamocha et Rio Yara in collibus siccis, Ekrnan 5604 (S);
High Maestra, Pico Turquilno,Le6n 11084 (LS, MIN, NY); Sierra Maestra, fromupper edge of
drier forests, Bucher 191 (ROIG 5329); Holguin in charrascales ad " El Paraiso, " Ekman
7613 (S); Cayo Largo, Puerto Padre, Cur-belo307 (LS-frag. on sheet of M. emarginata var.
rostrata, Le6n, Ekcman,Johnston, & Roig 9097; NY; ROIG 5638); Galbis (ad limit. prov.
Camagiiey) ill silva, Ekqnan 7443 (NY, S), 7460 (NY, S, US). Camagiiey: Colombia, Shafer
589 (NY, US); Camino de la Cueva del Circo; Roig, Luaces, 4; Arango, May 11, 1915 (ROIG
820F); La Gloria, Roig g' Ballou, May 18, 1915 (ROIG 1306).

4b. Mouriri emarginatavar. rostrata(Urban) Morley,comb.nov.

MouririrostrataUrban, Repert. Sp. Nov. 22: 240, 241. 1926.
Midrib below usually rectangularat least in the centerbut usually without
ncarrowwilngs;true pedicels 3.5-8 mm long; length of the calyx (including the
inferiorovary) 5.0-5.8 mm (dry measurementsca. 0.2 mm less) ; length of the
calyx-limb(including the freehypanthium)3.3-3.7 mm lolng;fusiondistance of
adjacent calyx-lobesabove the stamelnattachment0.7-1.0 mm; angle "o" 145?-
152?.
Type collectionand locality: Ekman 14069, Cuba, Havana province,at Rio
Qutezada,June 18, 1922. Lectotypeat S.
130 BRITTONIA [VOL. 9

Distribution: West Cuba, low mountainsof northcentral Havana province;

low to high mountainsnear the easternend of the Sierra del Rosario, Pinar del
Rio province,and low areas in the southwestend of the same province (fig. 26).
Specimens examined: Havana: Loma de la Pita, San Miguel de Casanova: Leon 11566
(LS, MIN-both dated Dec. 6, 1923; NY-dated 6-10-1924); Leon Ekman, Johnson,& Roig
9097, May 20, 1920 (LS, NY); Loma de la Pita, San Miguel del Padr6n (error for San Miguel
de Casanova), 1oig, May 20, 1920 (probably same as preceding; NY as Roig 4, ROIG 2032,
SV 7542); crest of ridge ca. 150 feet lower than and west of the highest point of the Loma,
ca. 200 m elevation, Morley 858 (MIN, US); Campo Florido, in forests on serpentine at Rio
Quezada (the part of the Rio Guanabo above Campo Florido is known locally as the Rio Que-
zada), Ekman 13245 (paratypes: NY, S); at Rio Quezada, Ekman 14069 (GH, NY, S-type,
US; photo of NY specimen,LS). Pinar del Rio: Prope Las Pozas ad Rio del Medio in sylvis
fruticosis solo serpentino,Ekman 17467 (NY, 5); low ridge between and ca. 200 yards above
confluenceof two small streams,ca. 1 mile south of Las Pozas-La Mulata highway, ca. 1/4 mile
west of small road leading south from highway and joining highway ca. /2 mile west of Rio
del Medio, low forest on serpentine,Morley 869 (MIN, US); Falda Norte del Pan de Guajai-
b6n, Alain 4+ Acufia 3004 (LS, SV 18585); ca. half way up north side of Pan de Guajaibon
on trail made by survey engineers,ca. 500 m elevation, in large pocket of red soil, Morley 865
(MIN, US); Rangel, Leon 22383 (LS, MIN); Guane, El Frances, Remates, A. Fors, March
25, 1929 (ROIG 4833); Sabalo, terrenospedregosos bajos, Roig, Nov. 24, 1915 (ROIG 1083, SV
18814).
The outstandingcharacteristicsof this species are its mostlydouble epidermis
with mucilage-thickened walls, its relativelyfew and large stomatal crypts,and
its long calyx. Mouririgonavensisis its nearestrelative; the primarydistinctions
betweenthe two are listed thereunder.In M. emarginata,leaf formis somewhat
more constantthan in the otherspecies of the section; otherwiseit too is quite
variable. The range in size of the plants is outstanding;this is apparentlylargely
a responseto growthconditions. Some trees on poor soil, oftenserpentine,are
so small and bushyas to be called shrubsby somecollectors,whereasothersin bet-
ter locationsmay grow up to 16 m tall. Also highlyvariable is the shape of the
calyx-lobes(figs.12-14). I have been unable to detectany basis for segregation
in thevariationof theseor othercharacters,exceptin the two varietiespresented.
Regarding these varieties,a comparisonof their descriptionsshows them to be
unusually well separated as contrastedwith the varieties of the other species.
They are best given onlyvarietal rank,in my opinion,because of theirverygreat
similarityin other features. Moreover,only three floweringspecimens of M.
emarginatavar. rostratahave been collected; more would probably expand the
knownrange of variation of the critical charactersso that more of themmight
overlap those of the eastern variety. Finally, there is a questionablespecimen,
Clemente573 (572?), 1920, Loma del Gato, Sierra Maestra, Oriente (LS), which
is very similar to specimensfromHavana in all charactersI have been able to
check. I suspect that there has been a confusionof labels, as the questioned
numbersuggests,and that the specimenis really fromHavana. If it is correctly
labeled, however,then the proposed differencesbetween eastern alnd western
membersare largely invalid. The contentionthat the separation between east
and westis a natural one, and not one merelybroughtabout by extensivecutting
and burningof the interveningareas, is supportedby the apparent absence of
M. emarginatafromthe mountainsof central Cuba, where original forestscan
still be found.

SUMMARY

The twelve nlamedspecies in the West Indian section Nesophyturm are here
reducedto four,ineludingtwovarietiesforeach of the four. Similarityof leaves,
1957] MANNING: JUGLANS 131

floralcharacters,and existenceof many sterilespecimensillake the

iiiter(gradinog
applicationof anatomicalcharactersdesirable. The charactersof stomatalcrypts,
epidermalconstrLction, externalshape of the inferiorovary,and the calyx have
proved mostuseful. The four species appear to be divisibleinltotwo basic types
which are most similar and least specialized in the eastern part of the range.
Their distributionacross Cuba, Hispaniola, anid Puerto Rico reflectsin part
the geologiealhistoryof the region.

A BOLIVIAN WALNUT FROM PERU GROWING IN COSTA RICA

WAYNE E. MANNING
Bucknell University
Lewisburg, Pennsylvania

In 1950 Dr R. J. Seibert reportedthe planting in 1948 in Costa Rica of nluts

of a walnLutcollectedin Peru ("A black walnut for Central Amierica." Ceiba
1:190-192). These were planted at the U.S.D.A., Bureau of Plant Industry,
CooperativeRubber Plant Field Station at Turrialba, Costa Rica; this is now
"ICA/IIAA" (International CooperationiAdministration,Institute of Inter-
American Affairs). Dr. Seibert suggestedthat the niutswere probablyJuglans
'neotropica.
Throughthe courtesyof Dr ErlnestP. linle, fornierDirector of the Rubber
Plant Field Station at Turrialba in 1950, and of Dr JorgeLe6n, of the Instituto
Interamericanode Ciencias Agricolas at Turrialba in 1955, specimenswere sent
to the writer for identification.These were both JutglaCtsboliviana (C.DC.)
Dode, definitelymatchingphotographsof the type seelnby the writer,and are
not J. neotropicaDiels. Both species are in Dode's sectionRhysocaryon,which
is typifiedby J. vigra, the black wallnut,and have the sanie general qualities of
wood and nuts.
Dr Seibert reportedthat the nuts were collectedfromthe La Merced region
of the Upper Perene valley of Peru, which is in the departmentof Junin (see
the National Geographicmap of South Anierica of 1950). The writerhas seen
anotherspecimenof ,J. botiviansacollected not far northeastof La Merced.
Although Juglans neotropica is essentially the only species of the genlus
whichoccurs in northernPeru, Ecuador, and Colombia,J. boliviana seems to be
the prinicipalspecies in centralanldsouthernlPeru as well as in Bolivia.
Very few specimensof J. boliviana have beenlcollected,and few if any that
inleludeleaves, flowers,and nuts. The nuts selntfromPeru to Dr Seibert,some
of whichwere sent by the latterto the writer,vary frolmi large, 4.5 ecmin diame-
ter, deeply and rather sharply ridged, to medium,3.0 cm in diameter,almost
striate,resemblingthe nuts of J. molltsof Mexico. The two extremesappear to
be fromdifferent species; this is, of course,a possibility,and there is, further-
more,some ulncertainty over which type germiinated.
There is a hope that thesetrees will colntiilueto grow in Costa Rica, and will
make it possibleto have rathereasily accessibletrees of this little-knowni species
available for futurestudy. As Dr Seibert suggested,this species may also pro-
vide for Costa Rica and othercountriesof Central America a tree of value for
wood and nuts.