Peculiarities of the Inflorescence in the Euphorbiaceae

Author(s): Leon Croizat

Source: Botanical Gazette, Vol. 103, No. 4 (Jun., 1942), pp. 771-779
Published by: The University of Chicago Press
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 PECULIARITIES OF THE INFLORESCENCE
IN THE EUPHORBIACEAE
LEON CROIZAT
(WITH FOURTEEN FIGURES)

The cyathiumof the Euphorbiaceae Euphorbieae is a structureunmatchedby

otherinflorescences in the spermatophytes.It is the purpose of this paper to dis-
cuss certain of its aspects, concludingwith a cursoryreview of the epicarp of
Ptychopyxis,a genus whichbelongs to the same familyif not to the same tribe.
In a laterpaper attentionwillbe given to the inflorescence
of certainCelastraceae
which suggestsconsiderationsof a general nature on the significanceof flower-
bearingaxes.
I. Cyathiumof Pedilanthus
The cyathiumof Pedilanthusconsistsof two chambers (fig. iA), one superior
and smaller,containinga variable numberof glands,the otherinferior and larger,
mostly tubular in its outline, holding the "ovary," that is, the Y flowerand
numerousstamenswhichin realityare monandrousc flowers.This "flower"has
been interpretedand describedby studentsof the Euphorbieae (BOISSIER in DC.
Prodr. I5(2):4. i862; MILLSPAUGHin Field Mus. Publ. Bot. Ser. 2:353. I9I3) in
a mannerwhichis hardlysatisfactory.For instance,BOISSIER concludesthat the
upper lip is merelyan externalcucullate appendage becomingsaccate at its base,
includingthe glands and having no communicationwith the "tube" surrounding
thesexualorgans.
To elucidatethe featuresof the cyathiumof Pedilanthus
whichhave proved
embarrassingto BOISSIER and MILLSPAUGH, it is necessaryonly to tilt this pe-
culiar inflorescenceuntil the gynophorestands almost vertical. This done, it is
necessaryto suppose that the upper chamberis separated fromthe lower by a
slightelongationof the pedicel. The diagramwhichfiguresthe outcomeof these
assumptions(fig.2) is strikinglyreminiscentof the two-storieddispositionchar-
acteristic for the inflorescenceof another euphorbiaceous genus, Dalechampia
(fig.3). The floralstructuresofPedilanthusand Dalechampia differbasically only
as follows: (a) in Pedilanthusthe upper chamber containsa variable numberof
glands but no active sexual organs; in Dalechampia the upper chamber contains
active e flowersin additionto moreor less evolute glands; (b) in Pedilanthusthe
lower chambercontains one 9 flowerand numerousa flowers;in Dalechampia
thelowerchamberis occupiedby three 9 flowersonly. On thisbasis, ifthe upper
chamberof the inflorescence ofDalechampia is deprivedofits a flowersand these
77I] [BotanicalGazette,vol. I03

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772 BOTANICAL GAZETTE [JUNE

are placed around the central 9 flowerof the lower chamber (the lateral ones
being suppressed),the cyathiumof Pedilanthusis seen to appear. The changes
hypothetizedhere to effecta passage fromthe inflorescenceof one to that of the

A~~3 A

FIGS. I-8. Fig. i, inflorescenceof Pedilanthustithymaloides;A, sectionthroughcyathiumshowing

upperchamberwithtwoglands,lowerwith 9 flower(ovary) and monandrousflowers(stamens). Fig. 2,
diagramillustratingthe presumedmannerof formationof cyathiumof Pedilanthus,the upperchamber
beingproducedabove the lowerin the arrangementcharacteristicof the inflorescence of Daleciampia.
of Dalechampia; A, ? flower.Fig. 4, cyathiumof Pedilanthustehuacanus.Fig. 5,
Fig. 3, inflorescence
same of P. linearifolius;A, section throughgland. Fig. 6, diagram illustratingeuphorbioidcyathium
(with several glands) at apex of main stem of Euphorbia geniculata,followedabove by pedilanthoid
cyathium(withone gland) in forkof firstdichotomyofumbellaster.Fig. 7, E. clava: apex of mainstem
bearingfalselyapical cyathiumset upon extensionof podaria of main axis. Fig. 8, normalapex of main
stem of E. clava.

othergenus are consistentwith the morphologyand the sexual expressionof the

Euphorbiaceae,a familywhichfrequentlyyields monoecious,dioecious,or polyg-
in the same genus and even in the same species.
amous inflorescences

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I942] CROIZAT-INFLORESCENCES 773

The theorythat the appendix of the cyathiumof Pedilanthusis homologous

with the upper bract that inclosesthe androeciumof Dalechampia, and that the
saccate chamberclosed above by thisappendix is actually a sterilizedandroecium
but potentially a i- or 2-sexual inflorescence,invites the objection that the
hypotheticelongationof the pedicel above and beyond the lowerchamber(fig.2)
is not to be taken forgranted. Proofshould be given that the stelarcontinuation
of the pedicel reaches the upper chamber of the inflorescence.In view of this
objectionit must be admittedthat the matterdoes not seem to have receivedas
yet the attentionof a specializedmorphologist.However,a carefulstudentof the
vascular anatomy of the cyathiumof Euphorbia (HABER in Ann. Bot. 39:704.
192S) has concluded that in this inflorescence the glands are "a pair of modified
secondarybranchesof a lateral inflorescence";that is, the glands are primarily
caulinar bodies, unlikeappendages of foliarorigin. Since the glands of Euphorbia
are oftenset upon the rimof the involucrein a mannerhardlysuggestiveof stelar
bodies, while those of Pedilanthusare oftenfound at the bottomof the cavity of
the upper chamber,it is not imprudentto assume, until proofto the contraryis
given, that the supply of these glands is a branch offthat of the pedicel of the
inflorescence.Such an assumptionhas in its favor broad phylogeneticand mor-
phologicalconsiderations,not less than the fact that the glands of Euphorbiaand
Pedilanthusare admittedlyhomologous.
Two modificationsof the inflorescence of Pedilanthusare worthyof notice. In
P. tehuacanusthe tube is sharplytruncated(fig.4), the openingbeingalmostverti-
cal. The appendixis deeplycleft,and, so faras known,no glandsare presentin the
upper chamber. The ovary is roundedas it usually is in Euphorbia,not elongate
as in typical representativesof Pedilanthus (for instance,P. tithymaloides). In
the
P. linearifolius upper chamber is exceedinglyreduced,appearingas a glandu-
lar saccate foldon the side ofthe vertical,baglikeinvolucre(fig.5). The tendency
inherentin the cyathiumof P. tehuacanusto depart fromzygomorphism withthe
concomitantreductionof the upper chambersuggeststhat a furtherevolutionin
this type of cyathium may yield a vertical, exappendiculate and eglandular
perianth,such as existsin the c cyathiumof E. plagiantha. This suggeststhat a
shiftin the positionof the floralaxes may have far-reaching consequencesupon
the nature of specialized reducedinflorescences.
The cyathium of P. linearifolius,a species which some treat as Cubanthus
linearifolius,does not differin its essential characters from the cyathium of
Euphorbia (Poinsettia)pulcherrima.These two species agree closelyin floraland
vegetativecharactersand to all appearances are consanguineous.It is possible
that the solitary nectary of the cyathium of E. pulcherrimais not absolutely
homologouswiththe saccate lateral appendage of that ofP. linearifolius, and that
theformeris a muchmorereducedand specializedstructurethan the latter. Both

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774 BOTANICAL GAZETTE [JUNE

these "glands" have the functionof hydatodes,however,whichsets themphysio-

logicallyon a par; it should not be surprisingthat theydifferin morphologicalde-
tails, because relevant differencesseparate the various kinds of nectaries and
glands prevalentin Euphorbia. The glands of E. esula, E. jacquiniiflora,and E.
tridentata are most unlike,but differences of the kind are correctlyinterpretedas
indicatinglittlethat has absolute value in a phylogeneticstudy. To judge froma
sum of characters,P. linearifoliusand E. pulcherrimaare the links connecting
Pedilanthusand more distantlyDalechampia with tne aggregateof species com-
monlytreatedby taxonomistsas Euphorbia. The phylogeneticsignificanceof the
so-called section Poinsettia of Euphorbia is certainlygreaterthan most taxono-
mists appear to believe, as it is throughthis sectionthat the prevailinglyactino-
morphiccyathiumof Euphorbia can be connectedin phylogenywith the zygo-
morphicinflorescenceof Pedilanthus,and -more remotely-with the ancestral
inflorescence commonto all the Euphorbieae. In this connectionit is interesting
that the species of section Poinsettia are closely related with Africanand Aus-
tralianformsin the vicinityofE. crotonoides and E. eremophila, formingan aggre-
gate (in the broad sense) that enjoys practicallya pandemic range.
In view of the positionoccupied by the species of sectionPoinsettia among the
Euphorbieae,it is not surprisingto findin thesespeciesdimorphiccyathiaofcon-
siderableinterest.In E. geniculata,forinstance,theprimarymain stem oftenends
witha cyathiumthat may have eightto tenglands; nextabove, at the firstdichot-
omy of the umbellaster,cyathia occur which are typical of the section and bear
only one gland, being the same as those of E. pulcherrima.This peculiar arrange-
ment (fig.6) suggeststhat in the speciesof thissectiona progressivespecialization
takes place at different levels on the florigerous
axes, the same specieshavingfirst
a euphorbioidcyathium (with more than fourglands), then poinsettoidcyathia
(with only one gland). Such a specialization,as has been noticed,also occurs in
Pedilanthus(fig.2), in which the lower chamberis sexuallycompleteand active,
theupper sterile,and occupied onlyby relicbodies ofthe natureofglands. More-
over,the firstcyathiumto appear in species of Euphorbiaoutside of sectionPoin-
settia is oftenunlike the cyathia that followit. This firstcyathiummay be uni-
sexual, have nearlyfreeglands and lobes, and especiallyeight to ten glands and
lobes instead of the normal numberof fourto five. In one species,E. capitulate,
no umbellastersform,each fertileshoot being terminatedby a single cyathium.
This cyathiumhas a status and positioncloselycorrespondingto thoseof the first
cyathiumof otherspecies of Euphorbia. It arises at the originof the umbellaster,
and, like it, has usuallyeightto ten lobes and glands,a peculiaritywhichSCHMIDT
has used to erecta monotype,Diplocyathium.Apparentlya cyathiumwith four
to fivelobes and glandsis not necessarilycharacteristicofthespeciesofEuphorbia,
and the cyathia of this genus and of the Euphorbieae in general tend toward
dimorphism.

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I942] CROIZAT-INFLORESCENCES 775

In her study of the cyathiumof Euphorbia,HABER has concluded (Ann. Bot.

39:694. 1925) that the articulationof the monandrouspseudo-stamen of this
peculiar inflorescence is "the receptacleupon whichthe floweris situated,and in
functionis similar to that of other receptacles." Since the receptacle may be
understoodas a regionin which earliergrowthcomes to a stop and new growth
begins,it is logical to suppose that a flowerthat is apparentlyterminalpast an
articulationis not necessarilyapical. This flowermay arise, in fact,froma meri-
stem that is lateral upon the receptacle,occupyingan apparent terminalposition
on the axis on account of the failureof the true apical meristemto develop a
flower.An articulation,consequently,may be analogous if not homologouswith
the scar that marks the insertionof a scion upon the stock to which it has been
grafted,and it is not surprisingthat it shouldbe anatomicallycomplex (LECOMTE
in Mus. Nat. Hist. Paris, Nouv. Arch., 5 ser. 2:I2I-242. i9io). In theoryat
least, an articulationdoes not requirethat the axis come to an abrupt end, estab-
lishinga sharp horizontalreceptacle. It may formwhen a lateral meristemac-
quirespreponderantgrowth,as when,forinstance,a lateral bud of Tilia or Ulmus
becomes "terminal" on the shoot on account of the dying-outof the apex of the
branchletupon which this bud is borne. The formationof an articulationmay
consequentlybe regardedas the resultof a theoreticalreversalin the normalratio
of developmentbetweenthe bud gap and the stele,the stele being ultimatelydis-
posed of by too continuousa proliferationof lateral buds. In the physiological
sense, this happens particularlywhen dormant structuresgain the upper hand
against active ones at the end ofa lengthor time ofgrowth. There does not seem
to be a sharp line to separate such true articulation,as that of the monandrous
flowerof Euphorbia, described by HABER, from the abscission layer and scar
which end a branchletof Ulmus or Tilia next above a falselyterminalbud.
The cyathium of Euphorbia is apparentlyterminal,and its pedicel does not
exhibita manifestarticulation. It is very likely,however,that this cyathiumis
not trulyapical. Euphorbiaclava,a succulentwhichis oftencultivatedas an orna-
mental,has main axes which normallydo not bear apical cyathia (fig.8), these
beingcarriedupon lateralspecializedflorigerous axes. In some cases, however,the
main stems taper to a peduncle-like end (fig. 7), crowned by a cyathium. Of
course this cyathiumis not truly apical, because it rests upon a structureof
spirallyascendingpodaria (= decurrentsucculentpetioles) and is in realityan
intercalaryinflorescence(PARKINin Jour. Linn. Soc. Bot. 42:556-558. I9I4).
What seldom happens in E. clava normallytakes place in E. caput-medusae,in
which all cyathia are borne upon peduncles of spirallyascendingpodaria. In a
speciesin the vicinityofE. clava (probablyE. loricata)I have foundby dissection
that thebottomofthe involucreactually slantsupon the supportingaxis, showing
that the cyathium is not really apical. Thus evidence is forthcomingthat the

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776 BOTANICAL GAZETTE [JUNE

cyathia of the Euphorbieae are not necessarilyand trulyapical, despite theirap-

pearingin a terminalpositionupon the axes.
Once it is realized that thesepeculiar inflorescences
are essentiallyintercalary,
that is, that theybecomefalselyterminalby the extinctionof theaxis whichbears

^~~ ~ 31 ~ ~ ~~~~ 14
FIGS. 9-I4.-Fig. 9, diagramillustratingcomponentsofcyathiumofEuphorbiageniculata;A, outline
of gland occupyingplace of sexual organsin upperchamber(=upper chamberof cyathiumof Pedilan-
ofEuphorbiaceaeEuphorbieae;A,
presumedancestralinflorescence
thus;see fig.2). Fig. io, illustrating
detail of glomerule. Fig. iI, Pavonia hastata: diagram of flowerin longisection.Fig. I 2, Cleistanthus
decurrens:diagramof oe flowerwithpistillode. Fig. I3, showingpossibleevolutionof the a' flowerof
C. decurrens by returnto functionalityof 9 organsand persistencyupon theovaryof sterilizedstamens
(= staminodesor processesof glandularnature). Fig. I4, flowerof Ptychopyxis thwaitesii(Podadenia
thwaitesii)showingpresenceofactual glandson epicarp.

themin a lateral position,it is not difficult

to reconstructthe hypotheticalances-
tral inflorescencewhich gave origin to any and all formsof cyathium,that of
Pedilanthus together with that of Poinsettia and Euphorbia. The dimorphic

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I942] CROIZAT-INFLORESCENCES 777

cyathia of E. geniculata(fig.6), forinstance,may be homologizedto threelateral

inflorescences (fig.9). Of these three,one yields an euphorbioidcyathiumby the
simpleprocessof fusionof glands and bracts (lobes), fourto ten glands becoming
adnate and alternate with as many bracts. On the contrary,the pedilanthoid
cyathiumwithone gland is formedby the specializationand subsequentfusionof
two inflorescences in a mannercharacteristicof Pedilanthus(figs.I, 2). In brief,
the dimorphismobserved in the cyathiumof the Euphorbieae in general is the
resultof the specializationof the inflorescences ancestralto the cyathium,these
inflorescencesbecomingcyathia eitheras singleunits or by fusionof two units,it
beingprobable that the euphorbioidcyathiumwithmany glands arises fromone
inflorescenceand thepedilanthoidcyathiumwithno apparentglands or one gland
fromthe fusionof two inflorescences.The fundamentalunitsin all these inflores-
cences are the following:(a) a central 9 flowerwith a much reduced calyculus;
(b) a variable numberof e flowers,with or withouta calyculus,bearingone to
severalstamens;(c) glandsrepresenting abortive6 flowers;(d) a variable number
of appendages. On this basis the ancestralinflorescence of the Euphorbieae and
Dalechampieae, and possibly of the Pereae, mighthave resembledthe inflores-
cence shown in figureio. Such an inflorescenceis aptly described (BAILLON in
Adansonia 6:354. i866) as being composed of "Cymes pluriparesa fleurfemelle
centrale,"and some of its earlieststages may not have been altogetherdifferent
fromthe floralarrangementofLongetiabuxoidesas definedby BAILLON. All these
inflorescencesrelateback to the capitula of the Tiliaceae and to the cymesof the
Malvaceae, modifiedby sexual specialization and by extensive fusion among
partial inflorescences and singleflowers.Thus the phylogenyof the inflorescences
ofthe Euphorbieaeconfirms what is knownon the strengthofotherevidence,that
the Euphorbiaceae and the Sterculiaceaeare closelyrelated,both familiesarising
fromthe malvoid ancestralgroup as a case of sexual specialization.

I. Epicarp of Ptychopyxis
The corolla of malvaceous flowersis described in most textbooksas dialipe-
talous. This is erroneous,as the corollain theseflowersis intimatelyconnatewith
the staminal column, the petals and the androeciumfallingtogether,calyptra-
wise, afteranthesis. Such flowersin longitudinalsection have an outline similar
to that shownin figurei i.
Pavonia hastata,a species of the Malvaceae, is knownto turn "cleistogamous"
at certaintimesof the year (GOEBEL in Organ. Pflanzen.3 ed. 3: I97I [fig.20351.
I933, CROIZATinDarwiniana5:4I9. I94I). This "cleistogamy" actuallyconsists
ofa reductionof thecorollaand staminalcolumnwhich,insteadofevolvingin full
blossom,fail to develop and remain inclosed withinthe barely opened or even
unopened calyx. That this is hardlya case of true cleistogamyneed not be em-
phasized. What actually takesplace in Pavonia is this: in summerthe corollaand

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778 BOTANICAL GAZETTE [JUNE

the staminalcolumndevelop in full,but forsome reason or otherthe gynoecium

remainssterileand no seeds are produced; in winter,on the contrary,the corolla
and the staminalcolumnbecome abortivebut the pollen and the ovules are active,
so that normalseeds capable ofgerminatingare produced. The backgroundofthis
differentialgrowth would seem to be metabolic because it is connected with
changesin average temperature.The tentativeconclusionmay be advanced that
P. hastata-and withit otherMalvaceae such as Sida spp.-tends towarda mode
of sexual expressionwhich involvesa departurefromthe usual bisexual arrange-
ment of the family. This tendencycalls into play apetaly and unisexuality.
A malvoid entitywhichbecomes completelyapetalous and unisexualis turned,
ipsofacto,intoa sterculiaceousand euphorbiaceousplant because the Sterculiaceae
in part and the Euphorbiaceae as a whole differessentiallyfromthe Malvaceae in
beingapetalous and unisexual. No sharpmorphologicalline can be drawnto sepa-
rate the Malvaceae, Tiliaceae, Sterculiaceae,and Euphorbiaceae, for they differ
in tendenciesmorethan in characters. Thus Pavonia and Sida illustratethe begin-
ning of a type of floral evolution which eventually ends by establishingthe
cyathiumofEuphorbia. The firststep in thisevolutionis probablytakenwhenthe
corolla and the staminal columnbecome intimatelyconnate,forit is then that a
type of flowerwhich is found in the Theaceae and, fartherstill, in the Mag-
noliaceae, becomes actually malvoid. As a matter of fact, the entire evolution
may be tentativelyoutlined as follows: (a) the syncarpic, conelike gynoecium
becomes reduced,only the basal row of carpels survivingeitherdialicarpicallyor
syncarpically;(b) the stamens fuse togetherand with the corolla; (c) eitherthe
corollaor the staminalcolumnevolves,ultimatelyleadingto unisexuality;(d) uni-
sexuality being attained, the flowerundergoesfurtherreductionin detail. All
these steps merelypostulate concentrationand specializationof the various floral
organs,that is, an essentialprogressionin reduction. It is not surprisingthat in
the familiesin whichunisexualityand reductionprevail,such as the Sterculiaceae
and the Euphorbiaceae, nectariesare frequentlyfound togetherwith pistillodes,
these organsrepresentingsurvivingbut degenerateparts of the androeciumor of
the gynoecium.
In Pavonia hastatathe gynoeciummay become sterile,eitheranatomicallyor
physiologically,this being a point upon whichno informationseems as yet avail-
able, and the androecium (corolla and staminal column) may become reduced,
simulatingcleistogamyand actually presentinga formof apetaly. In numerous
Euphorbiaceae the 9 organsare abortiveand reducedto pistillodesor "glands,"
while the c' organs are fully developed, the phylogeneticlink between the
Malvaceae and the Euphorbiaceae beingfurnishedby the Sterculiaceae. An espe-
ciallyrevealingexampleofflowerin whicha pistillodeis surroundedby stamensis
that of Cleistanthusdecurrens(fig.I2), a euphorbiaceousplant.

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I942] CROIZAT-INFLORESCENCES 779

In thelightof the tendenciesprevalentin the malvoid-euphorbioid phylumit is

permissibleto suppose that the pistillode of C. decurrensmight again become
sexuallyactive and anatomicallyperfectin connectionwith the process of ovule-
and seed-bearing,the stamensbeingturnedintostaminodes,that is, into sterilized
stamensor glandularbodies. As the resultofthissuppositiona structureis visual-
ized of the kind shown in figureI3, in which the staminode or its equivalent, a
gland, persistsupon the ovary and the fruitas a process or an appendage, the
androecium(staminal columnand corolla) becomingpart of the epicarp.
The hypothesisthat the processesupon the capsule of certainEuphorbiaceae
are actually relicsfroman abortiveandroeciummay not be truein everycase, but
it certainlyexplains manifestationswhich are otherwisedifficultto account for.
In certainspecies of Codiaeum,forinstance,the epicarp is beset withglands that
do not differin the slightestfromfloralnectaries. Such nectariesare numerous
upon the ovary and fruitof Ptychopyxisthwaitesii(fig.W and are in no wise dis-
tinctfromthe staminodesand glands normallysituated next to the calyx. It is
wellknown,moreover,that in Ricinus thepricklesupon the capsule have a vascu-
lation of theirown whichis not suggestiveof epidermalprocessesas such. In con-
clusion: some evidence is forthcoming that in the malvoid flowerand its deriva-
tive structures,the carpelsas well as the stamensmay become intimatelyconnate
and adnate withthe corolla,whichis fullyin harmonywith the tendencytoward
reductionand unisexualityprevailingin the malvoid and euphorbioidphylum.
Since it is admitted-on the mere strengthof visual evidence-that the abortive
gynoeciumof the Euphorbiaceae persistsas a pistillode (Cleistanthus)or as dis-
creteglands (Tetracoccus,Securinega)in the c flower,it should not be surprising
that theandroeciummay persistas glands or otherprocesses,freefromthe epicarp
(Croton)or intimatelyconnate with it (Ptychopyxis)in the 9 flower.

Summary
i. The cyathia of Euphorbia and Pedilanthus representdifferent degrees of
adaptation and fusionin a single type of compound lateral inflorescence.Such
was probablyintercalaryin its origin.
an inflorescence
2. A tendency toward the reduction of floral parts involving apetaly and

asexualityappears in the Malvaceae and becomes dominantin the Sterculiaceae

and the Euphorbiaceae.
3. The glands and processes on the epicarp of Codiaeum, Ptychopyxis,and
Ricinus are apparentlyrelicsfromthe androeciumthat became intimatelyfused
with the carpels. Glands of the same nature probably appear also in other
families(Sabiaceae).
ARNOLD ARBORETUM
JAMAICA PLAIN, MASSACHUSETTS

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