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 STAMINATE STROBILUS OF TAXUS CANADENSIS
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY 255
A. W. DUPLER

(WITH PLATES XXIV-XXVI AND TWENTY-TWO FIGURES)

Introduction
In a previouspaper(8) thewriter describedthegametophytes of
Taxus canadensisMarsh.,withthestatementthatotherphasesof
themorphology wouldbe treatedinlaterpapers. In thispaperthe
staminatestructures with respectto developmentand vascular
anatomyare described. The lack of detailed information con-
cerning thesestructureshas seemedto thewritersufficient
justifica-
tionfortheinvestigation herereported. In viewof thegenerally
recognizedconservativecharacterof the staminatestructures in
conifers,it seemsthata moreextended ofthem,in the
investigation
groupas a whole,wouldbe worthwhile. The description of the
ovulatestructures willbe givenin anotherpaper.
The generalstatement in thepreviouspaperas to materialand
methodswill also apply here. The writeris underobligationsto
ProfessorW. L. EIKENBERRY,of theUniversityof Kansas, forsome
materialcollectedin northernIllinois a numberof years ago.
Acknowledgmentsare also due ProfessorsJOHNM. COULTERand
C. J. CHAMBERLAIN, under whose direction the study of Taxus
canadensiswas begun.
Historical
Whilethe male gametophyte and its attendantfeatureshave
receivedconsiderableattention,apart fromthe generalmore
obviousfeaturesverylittleis foundin theliteraturedealingwith
Taxusas to themorphology ofthestaminatestrobilus itself. The
earlierworkerswho studiedthe staminatestructures of conifers
wereconcerned largelyinattemptsto interpretthemin termsofthe
angiosperm flower,naturallyleadingto confusion as to the true
natureof the structures.These earlierviews have been sum-
marized by VONMOHL (26) in perhaps one of the most important
345] [BotanicalGazette,vol. 68
346 BOTANICAL GAZETTE [NOVEMBER

of the earlypapers dealingwith the "male flowers"of conifers,and

to whichwe are indebtedforpart of the followingstatementregard-
ing this earlyinterpretationof the staminate strobilusof Taxus.
LINNAEUS (i6) regarded the entirestrobilusas a single flower,
with the stamensin a cylinder,the perianthlacking and replaced
by bud scales. JUSSIEU(Ii) held that the strobiluswas a monadel-
phousflower;whileLINDLEY (I5) consideredthe strobilusas a true
cone with naked monadelphous flowers,each sporophyll repre-
sentinga flower. RICHARD(ig) went still farther,with the rather
unique view that therewere 5-8 flowersunder each scale to which
the stalk of the floweris attached on the underside. Accordingto
thisview the pollen sac representeda "flower,"and he had a similar
interpretationfor the sporophyllsof Thuja and Juniperus. Zuc-
CARINI (28), regarding the reproductivestructuresas modified
portionsof the stem, comparable with the phylloclads of Phyllo-
cladus, described the antherof Taxus as 7-8-lobed around the tip
of a central column. He considered that Taxus has the most
complete male flowerin conifers,in other formsthe anther folds
growingon only one side of the central column, the other side
growingout into a scale. VON MOHL opposes the idea of the stem
characterof the "flower" of conifers,and objects to the view that
the "anthers" of other conifershave been derived from such a
structureas that of Taxus, because "we have yet no certain data
with which we can determinewithcertaintywhetherthe antherof
Taxus arises fromone leaf or froma whorlof leaves."
As comparedwithour presentideas these earlyviews are rather
strange,having largely only a historicalinterest,with very little
bearingon the real morphologyof the structuresconcerned. Con-
siderably later STRASBURGER (22) made some observationson T.
baccata, describingthe spiral arrangementof the scales of the
strobilusand the grosserfeaturesof the developmentof the sporo-
phylls. He held that the peltate stamen of Taxus representsthe
"extreme formof stamen," and found that it begins as a rounded
knob about the firstof August,becomes lobed by lateral swellings
due to internalgrowth,and that pollen mothercells formin these
lateral swellingsand produce pollen by tetrad divisions. He also
describesthe pollen regionas separated fromthe epidermisby two
,9i9] DUPLER-TAXUS 347

layersofirregularcells,andstatesthatdehiscence is accomplishedby
theruptureofcellsat thebase and sidesofthepollensac. CHAM-
BERLAIN(3) describesthemicrosporangium of T. canadensisat the
mothercell stage(Octoberi, i897), at whichtimethenucleiare
stillrathersmallin comparison withthesizeofthecell,thetapetum
beingsharplydifferentiated, and its cells showingno tendencyto
plasmolyzelike the cells of the sporangiumwall. PILGER(i8)
describesthegeneralexternalfeatures, largelyfromthetaxonomic
viewpoint,speakingof the "flower"as consistingonlyof sporo-
phyllssurroundedat the base by a scale envelopewhichcom-
pletelyinclosestheflower in thebud state. He regardsthe"leafy
structure"of theanther,whichis yetto be recognized in Torreya
and Cephalotaxus, as being"entirely lost" in Taxus.
In the relatedformsthestaminatestructures of Torreyahave
been describedin a general way by PILGER,based on T. nucifera;
in more detail by Miss ROBERTSON (20) for T. californica; and
by COULTER and LAND (5) for T. taxifolia. In Cephalotaxus
have beendescribed
someofthefeaturesofthespermatogenesis by
STRASBURGER (23), and by ARNOLDI (I). STRASBURGER (24)
pointedout thatthepollengraindividesin thesporangium
before
shedding; LAWSON(14) also confirmsthis in C. drupacea; and
WORSDELL (27) of the generalfeaturesof the
gives a description
"male flower,"based on C. Fortunei,comparingit withthoseof
otherforms(Phyllocladus especiallyin thesporophyll
and Ginkgo),
features.
Strobilibuds
In theaxilsoftheleavesoftheshootofa givenseason3 types
of structures are produced: (i) the vegetativebuds fromwhich
developthelateralleafyshootsof thenextseason; (2) theyoung
staminatestructures, maturing thenextseason; and (3) theovule-
also
bearingstructures, maturing thenextseason. Duringthefirst
seasonall of thesestructuresare in bud form,the staminatebuds
duringthelatterpartofthesummer andwinterbeingmoreglobular
than the othertwo kinds,whichare so nearlyalike in external
appearanceas to maketheirdistinction uncertainexceptby very
carefulexamination.
348 BOTANICAL GAZETTE [NOVEMBER

The rudiments beginto developverysoon

of thesestructures
ofgrowth
afterthebeginning oftheterminalbud in thespring,the

FIGS. I-6.-Longitudinalsectionsofyoungbuds: fig.-I,youngvegetative shoot

withbudrudiment budwithconicalapex; fig.3,
inaxilofyoungleaf;fig. 2, vegetative
staminatebud with broadenedapex; fig.4, ovulate bud, showingvegetativetip (to
left)and rudimentofovulate strobilusin axil ofscale; fig.5, young staminatestrobi-
lus, showingprimordiaofstamensand tip ofaxis; fig.6, youngstaminatestrobiluswith
primordiaofsporophylls,axis apex not evident; X36.

rudiments appearingas conicalprojectionsin the axils of young

leaves(fig.I). By themiddleofJunetheseaxillarystructures have
1919] DUPLER-TAXUS 349

attained a lengthaveragingabout i . 5 mm., consistingof the main

axis surroundedby compactlyarrangedscales. In this early stage
one cannot distinguishthesestructuresfromone another,eitherby
externalappearanceor in section. Early in July,however,one can
recognize in median longitudinal sections the beginning of the
differentiationwhichis now takingplace, the apex of the vegetative
bud remainingconical (fig.2), as is characteristicof the vegetative
stem tip (fig. i), the apex of the staminate structurebecoming
broadened(fig.3), while the ovule-bearingstructureis recognizable
by the rudimentof the ovulate strobilusappearing in the axil of
one of the scales near the tip of the primaryshoot (fig.4). All 3
kinds of buds may occur on the same shoot; in fact,thisis theusual
occurrence,with the staminatebuds generallythe more numerous,
the vegetativebuds nearestthe tip,and one to several ovulate buds
a short distance below the vegetative ones, the staminate buds
occupyingthe older portionof the shoot.
The buds arise only on the currentseason's growth,and in case
of the staminate structuresalways mature the next season. No
cases were observed in which staminatestrobiliwere produced on
older growth,nor were any cases found in which the buds remain
dormant for a time and then mature. Miss ROBERTSON,in her
study of Torreyacalifornicafromtrees growingin England, found
that whilethe staminatestrobiliare formedin the axils of the leaves
of a currentseason, theymay remaindormantforas longas 3 years.
In Taxus buds may be found on older growth,but they are either
dormantvegetative buds or persistingprimaryshoots of the ovu-
liferousstructuresof a formerseason, as will be described more
fullyin the paper dealing with these structures.

Sporophylls
PRIMORDIA

The broadened apex (fig. 3) is the firstindication of the true

nature of the staminate strobilusbud, and can be recognizedfirst
about July i. STRASBURGER(22) was able to recognize the stami-
nate structureof T. baccataabout August i, and in Torreyataxifolia
COULTERand -LAND (5) firstobserved the staminate buds in July,
350 BOTANICAL GAZETTE [NOVEMBER

but the primordiaof the sporophyllsdid not begin to appear until

August. The greatermeristematicactivityin some regionsof this
rounded apex than in othersmarks the position of the primordia
of the sporophylls. These soon become rounded lobes above the
general surface (figs. 5, 6). The nature of the growth of the
primordiumwould indicate that it arises froma group of meriste-
matic cells ratherthan froma definiteinitial; at least no defined
sporophyllinitial could be recognized.
The sporophyllsare probably spirally arranged,although this
is somewhatindefinite,and indicationswere found in a few cases
that theymay arise in acropetalsuccession(fig.5); but if thisis the
case it is very soon obscured in the uniformdevelopmentof the
primordiaas the sporophyllsdevelop, no trace of the axis apex
being recognizable after the very early beginningsof the sporo-
phylls. The earlydevelopmentof the primordiumis uniformin all
directionsfromits centralaxis, at least until the differentiationof
the archesporialinitials takes place. The strobilus in this stage
showsa seriesofroundedsporophyllprimordia(figs.5, 6, 23). The
later developmentof the sporophyllis so intimatelybound up with
the developmentof the sporangia as to best be described in con-
nection with them. In fact, the development of the sporangia
determinesthe shape and characterof the sporophyll,as aside from
the sporangia the sporophyll consists of practically nothing
exceptingthe shortcentralaxis and the epidermis.

MICROSPORANGIUM
ARCHESPORIAL INITIALS.-HOFMEISTER(II) seems to have been
the first to publish with referenceto the microsporangiumof
conifers,reportingthe spore mothercell stage as being reached in
Pinus maritima in November. GOEBEL (9) traced the arche-
sporiumofPinus to a singlehypodermalcell, and claimed a similar
originforthe archesporiumof Thuja. His mostimportantobserva-
tion on thispointwas that the developmentofthe microsporangium
is like that of the eusporangiateferns. COKER (4) in Taxodium
distichum,and NICHOLS (I7) in Juniperuscommunisvar. depressa,
also found a hypodermal origin of the archesporium,in the
latter case consistingof "a plate of radially elongated cells, 4-6 in
I919] DUPLER-TAXUS 35I

number, when viewed in longitudinal section." COULTER and

LAND, interpretingthe structuresin abortive sporangia, conclude
that in Torreyataxifoliathereis "a singlehypodermalarchesporial
cell." Miss ROBERTSONdid not get the originof the sporogenous
tissue in T. californica.
In Taxus canadensis the development follows the usual eu-
sporangiate method, the 4-8 (usually 5-7) archesporial initials
arising fromthe hypodermallayer of the sporophyllprimordium
while this structureis yet quite small (fig. 23), being uniformly
distributedalong its margin,and, dividingby periclinalwalls, form
theprimarywall cell and the primarysporogenouscell (figs.27-29),
as in Torreyataxifolia(5) and mostotherforms. These initialsare
firstto be recognizedby the size of the cells and of their nuclei
(figs.23-26). One initial cell seems to be the rule, althoughcases
werefoundin which the archesporiumconsistsof 2 cells (fig.26).
SPOROGENOUS TISSUE.-The primary sporogenous cell or cells
soon divide periclinally (fig. 30) or anticlinally before or after
the divisionof the primarywall cell,and by successivedivisionsthe
mass ofthesporogenouscells is increased(figs.3 I-35), theformation
and growthof which result in the lobed peltate structureof the
sporophyll,the sporangia being uniformlydistributedaround the
centralaxis whichcontinuesthe veryshortstalk of the sporophyll.
As thetissueincreasesthereis a corresponding growthof the epider-
mis and the sporangiumwall (to be describedlater), the completion
of whichresultsin the separationof the sporogenoustissuefromthe
otherportionof the sporophyll(fig.34). The tapetumis differen-
tiated from the peripheral layer, and the remainingsporogenous
mass increasesin amountuntilthemothercell stage is reached early
in October,as described by CHAMBERLAIN (3) and the writer(8).
This has been given (6), and even quite recently(7), as the winter
conditionof the microsporangium, and has frequentlybeen quoted
by writers. As the author has already pointed out (8), microspore
formationtakes place during the early part of October,collections
covering a number of years and from several localities in the
northern United States bearing out the statement that the
microsporeis the wintercondition of Taxus canadensis. STRAS-
BURGER (25) foundmicrosporeformationin T. baccatatakingplace
352 BOTANICAL GAZETTE [NOVEMBER

in February in I904, duringunusually warm weather, indicating

that in thisformthe sporogenoustissue remainsin the mothercell
stage until spring. It would be of interestto know the behavior
in the extremenorthernpart of the range of T. canadensis,as it is
possible that in regionsfartherto the north the microsporestage
mightnot be reached beforewinter. The microsporemothercell
stage is the winterconditionof Torreyacalifornica(20) in England

FIGS. 7, 8.-Median longitudinalsections of older strobili: fig. 7, at time of

completionof sporangiumwall, showingoval areas of young sporogenoustissue;
vascular tissue of axis and upper scales, shownin black, embryonicvascular tissueof
upper portionin outline; fig.8, winterconditionof strobilus,showingglobularchar-
acter of bud and microspores;vascular tissueas in precedingfigure; X36.

and of T. taxifolia (5) in Florida. During this developmentthe

strobilushas grownconsiderablyin size (cf. figs.7 and 8), becoming
more pronouncedlyglobular,and it remainsin this conditionuntil
the renewedgrowthof springtakes place.
No cases of abortive sporangiawere found,and it seems a safe
assumptionthat a sporangiumdevelops fromeach initial or initial
group. The adult sporangia show some variation in size, but not
enoughto indicate any tendencyto abortionof any of them. This
19I9] DUPLER-TAXUS 353

is in marked contrastwiththe behaviorin the related Torreya,in

whicha resincavity resultsfromthe abortionof some of the poten-
tially sporogenoustissue of the sporophyll,the abortion beginning
at the primarysporogenouscell stage, as pointed out by COULTER
and LAND for T. taxifolia. This resultsin the sporangiaoccurring
on onlyone side ofthe otherwisepeltatesporophyll. Miss ROBERT-
SONalso findsthat normallythereare 4 sporangiaon the side of the
sporophyllof T. californicaand a resincavity on the otherside, but
that the strobilus axis sometimesterminatesin a radially sym-
metrical sporophyll, like that of Taxus, with 6 or 7 mature
sporangia. Whethera resincavityis presentin such a sporophyll
is not stated, the inferencebeingthatmost or all of the sporangium
initialsreached maturity. A similarabortion of sporangia,in the
formationof mucilage cavities, is indicated by Miss STARR'S(2i)
work on Ginkgobiloba. COULTERand LAND findin Pinus Laricio
resin cavities related to sporangia, exactly as are the lateral
sporangia to the two middle ones in Torreya,and say "there is
evidenta tendencyto reduce the numberof sporangiaby abortion,
a reductionthat has proceeded fartherin Pinus than in Torreya,
and in the latterfartherthan in Taxus." It seems that when resin
or mucilage cavities are presentin the sporophyllthe sporangium
initialsare involved,and whenabsent theseinitialsmay all function
normally,as in Taxus. Whetherthis can be made as a general
statementfor all formswith resin cavities in the sporophyllmust
wait formore extensiveworkon otherforms. Cephalotaxushas a
sporophyllsimilarin generalappearance to that of Torreya,but it
is not known whetherany abortiontakes place.
SPORANGIUM WALL.-The initial development of the wall is
fromthe primarywall cell, which by a periclinaldivision formsa
tierof 2 cells. As the sporogenoustissue develops, these wall cells
divideanticlinally(fig.3I), increasingthe extentof the wall layers.
Only a portion of the wall is derived fromthe primarywall cell,
however,as othercellsabuttingtheyoungsporogenoustissuedivide
periclinallyand add to the wall, firston the outerside (figs.3I-33)
and thenon the innerside as well (fig.34), thus completelyenvelop-
ing the young sporogenous tissue. The wall usually consists of
2 layers of cells, although 3 or even more layers may be present,
354 BOTANICAL GAZETTE [NOVEMBER

especially at the angles formed by the mutual pressure of the

sporangiallobes.
As the sporogenoustissueincreasesin size thereis pressureupon
the wall cells, and theybecome flattenedand extended,so that at
the timeof theirmaximumdevelopmenttheyare broad thinplates.
By the timeof sporeformationtheyare usually quite flattened(fig.
36), and duringthe furthergrowthof the spores become more or
less disorganized,so that by the time the spores have reached
maturity,just before shedding,the wall has become a very thin
layerabuttingthe epidermis,whichhas now become, in effect,the
functionalsporangiumwall.
TAPETUM.-At the timewhenthe sporangiumwall is completed
the sporogenoustissue inclosed withinit is uniformin appearance
(fig.34). Soon, however,theperipherallayerof thistissuebecomes
differentiatedas a tapetum (fig. 35), thus originatingfrom the
sporogenoustissue and not fromthe innerlayer of the sporangium
wall, as in some forms. The tapetum, however, has its chief
significance from a physiological standpoint, being generally
regarded as a nutritivelayer, its origin seeming to be of little
morphological significance. The tapetal cells are usually uni-
nucleate, but not infrequentlyare binucleate (figs.35-36). The
tapetum is quite distinctduringlater phases of the developmentof
the sporogenoustissue,is sharplydifferentiated at the sporemother
cell stage, as pointedout by CHAMBERLAIN (3), and remainsdistinct
during the early winter (fig. 36). With the growth of the
microsporesin the springit be-comesless and less prominent,until
near pollinationit consistsof only a very thinlayerof disorganized
material surroundingthe spore mass.

EPIDERMIS

From the beginningof theprimordiumto thematuresporophyll

the epidermisremainsas a distinctlayer, the sporangiumdevelop-
ing fromhypodermaltissues,as already stated. During the early
growthof the sporophyllthe epidermisis meristematicthroughout,
dividinganticlinally(fig.24), its surfacearea thuskeepingpace with
the increasein the mass of the sporogenoustissue. An occasional
periclinaldivisionresultsin the epidermisbecoming2 cells thickat
i9g9] DUPLER-TAXUS 355

some points. The meristematicability, however, soon becomes

limited to the base of the sporangium,the epidermalcells of the
remainder of the sporophyllbecoming largerand with less dense
contents, the cells at the base remainingisodiametricand rich in
cytoplasm (figs. 32-34). As the sporangium increases in size,
causing a more pronouncedlobingof the sporophyll,the necessary
increase in epidermalsurfaceis effectedby the enlargementof the
non-meristematiccells and the addition to them of cells fromthe
basal meristematicregion. The enlarged cells become filledwith
an amorphoussubstance and the walls become thicker.
By the timethe sporangiaare maturethe epidermishas become
the functionalwall of the sporangium,owing to the practical dis-
integrati6nof thetruesporangiumwall. At maturitytheepidermal
cells are devoid of contents and have the markingscharacteristic
of the walls of many sporangia (figs. II-12), these thickenings
exercisinga hygroscopiceffect,usefulin the liberationof the spores.
JEFFREY (12) regards this thickeningof the epidermalcells of the
sporophyll,in a mechanical dehiscingdevice, as the result of the
invasion of the epidermis by mechanical tissues of fibrovascular
origin. There are no indicationsin Taxus of mechanical elements
elsewherein the sporophyll. COULTERand LAND foundnumerous
stomata in the epidermisof Torreya. In Taxus canadensis there
is a singlestoma on a sporophyll,at the centerof the peltate disk,
occupying the bottom of the depression caused by the enlarged
sporangia (fig. Io). GOEBEL (IO) shows a similar situation in
T. baccata.
Mature strobilus
The scales at the base of the strobilusare small and decussate,
increasingin size and becoming spiral in arrangementabove, the
uppermostones being considerablylargerthan the lowerones, and
functionas bud scales in the immatureconditionof the strobilus.
The scales are brownish in color, with heavily cutinized outer
epidermal walls, especially on the abaxial surface, the stomata
occurringonlyon the innersurface(fig.37), reversingthe condition
on the vegetativeleaves of the plant, wherethe stomata occur only
on the lower (abaxial) surface. The midrib is marked by the
356 BOTANICAL GAZETTE [NOVEMBER

100
15.-
t
- ----- - -- - (
t7~~~~~~~~~~~~~~~~o
20----- ~ ~ ----- O

21.--. e

221-__ __ _

9 2
~~~~~~~~~~~1
FIGS. 9-i2.-Fig. 9, median longitudinalsection of maturestrobilusjust before
pollen shedding,showing"elongating region" of axis and 4. sporophylls; xylemof
bundle black; note that xylembecomescentrallyplaced in upper portionsand does
not extendas farintostalksas phloem; numbersat left(W3-22) indicateapproximately
levelsofcross-sectionsofstrobilusshownin figs.I3-22; fig.IO, mediansectionofopen
sporophyll,showingelongatedstalk,open sporangia,and solitarystoma in centerof
depressionofdisk; fig.II, tangentialsectionof matureepidermis,showingmechanical
thickeningson walls; fig. i2, cross-sectionof matureepidermis,with microspore,at
timeof shedding; figs.9, IO, X36; figs.II, I2, X475.
19191 DUPLER-TAXUS 357

vascular bundle, when present,and occasionallyby sclerenchyma-

like cells along the outer margin of the midrib. In the young
strobilusthe mesophyllof the scale is compact,but as the strobilus
matureslarge air spaces develop. In additionto the solitarystoma
foundon the sporophyll,stomata occuron thestrobilusaxis between
the bases of the sporophyllswithrathersurprisingfrequency,being
foundonlyon thisportionoftheaxis and not on theportionbetween
the upper scales and the lower sporophylls. While the functional
characterof thesestomata mightbe open to question,owingto their
positionratherthan to theirstructure,theirchiefinterestprobably
lies in theirmorphologicalsignificanceas hereditarystructuresfrom
a morehighlyvegetativeancestralstrobilus.
During the autumn, winter,and early springthe strobilushas
the appearance of a globular "bud," the stamensbeing surrounded
by the uppermostscales (fig.7). The axis betweenthe upper scales
and the bases of the lower sporophyllsis veryshortand remainsso
until a few days beforematurity,duringthe latterpart of Aprilin
central Pennsylvania, at which time there is a rapid enlargement
and elongationof this portion of the strobilus,the effectbeing to
push the sporophyll-bearing portionbeyond the scales (fig.9). A
similarelongationof this regionis reportedfor Torreyacalifornica
(20). COULTER and LAND describedan enlargedpith regionin the
axis of the strobilusin T. taxifolia,which the authorssuggestmay
be "an important storage region for the strobilus." No such
enlarged region was found in T. canadensis. In addition to the
elongation of this portion of the strobilus axis there is also an
elongationof thestalkofthe sporophyll(cf.figs.9 and io), resulting
in the separationof the sporophyllsfromone another.
The sporangia do not hang freelyfrom the underside of the
disk, but are fused with the stalk on the inner side (fig. 9), and
laterallyare separatedfromone anotheronlyby thinpartitions,the
externalfurrowsbetweenthe sporangianot extendingall theway to
the center,the sporophylland sporangia thus constitutinga very
compact structure. RICHARD (Ig), STRASBURGER (22), and
GOEBEL (IO) gave accounts of the dehiscenceof the sporangiumof
T. baccata,in which theypointed out the ruptureof the sporangia
at the base and the umbrella-likemovementof the epidermalwall.
358 BOTANICAL GAZETTE [NOVEMBER

The processis the same in T. canadensis,the breakingof the thin-

walled epidermalcells at the base of the sporangiumin a circle
around the base of the stalk, the ruptureof some of the cells at the
side of the sporangium,and the hygroscopicrole of the thickened
epidermalcells resultingin the wall of the sporangia spreadingout
in umbrellaform,the thinpartitionsbetweenthe several sporangia
also beingbrokenin the process.
When young the strobilirudimentsare erect in the axils of the
leaves, but as they develop theybecome orientedin such a way as
to hang pendenton the lowerside of the shoot,the fertileportionof
the strobilusbeing directeddownward. GOEBEL (io) regardsthe
position and the method of dehiscencesuch as to secure the most
advantageous distributionof the pollen.

Vascular features
Since the reproductiveorgans, and especially the staminate
structures,are regarded as among the most conservativeof plant
organs,a considerationof the vascular anatomy of the staminate
strobilus is not without interest. While the ovulate strobili of
conifershave been the subject of considerable investigationand
discussion,in theirvascular as well as in otherfeatures,the stami-
nate strobili have not received much attention in their vascular
anatomy, probably not as much as they deserve in view of the
conservativenature generallyassigned to them on other grounds.
The only reference to this feature of Taxus is by STRASBURGER(22),
who gave the arrangementof the scales of T. baccata and states
that each stamen contains a bundle which passes into the stalk.
Like any otherbranch,the strobilusaxis receives2 bundlesfrom
the cylinderof the leafy shoot. These are semicircularin outline,
and by meetingat theiredges soon forma closed cylinder,broken
here and thereby the gaps formedby the weak bundle tracesof the
scales. In the lower portionof the strobilus,where the scales are
small and decussate, the small traces oftenend in the cortex and
do not reach the scale itself. The traces for the upper scales are
betterdeveloped and extendforsome distanceinto themidribof the
scale, especiallyin the 2 or 3 uppermostscales. Althoughthe axis
cylinder,as well as the cortical portion of the scale traces, are
1919] DUPLER-TAXUS 359

collateral endarch,in theirterminalportionstheycontain not only

centripetal xylem, but are also accompanied by transfusiontis-
sue which may be both dorsal and lateral to the xylemelements
(fig.38).
At the level of the uppermostscales the cylinderconsistsof 3 or
4 large bundles (figs.22, 48) whichextendinto the fertileportionof
the strobilus,wheretheybranch,givingofffinallya branchto each
sporophyll,the bundle extendinga little way into the base of the
sporophyllstalk. In a young strobilusthese bundles are repre-
sented only by elongated thin-walledelements, evidently pro-
cambiumstrands,whichtraversetheregionbetweenthe base of the
sporophyllsand the level of the upper scales (figs. 7, 8). These
strands remain in this embyronicconditionuntil near maturity,
whentheyelongateand take on theirvascularfeaturesin connection
with the growthof the "elongating region" of the strobilusaxis.
In this "elongating region" the pith becomes larger in diameter
than in the lowerportionof the strobilus,but shows no evidenceof
being in any way a storageregion; in fact,therewould be littleuse
of a storagetissue at this stage in the developmentof the strobilus.
The several large bundles of the strobilusaxis extendforsome dis-
tance into the " elongatingregion" and thengive offbranchesto the
various sporophylls,each of the large bundles supplyingseveral
sporophyllsin this way (figs. I3-22). Some of the branchesmay
unite and then separate (see the behavior of bundles Aiand e, and
also of 1,c, and m,in figs.i8-22), althoughusually the bundlespass
ratherdirectlyto the base of the sporophyll(figs. 14-I7, 44-47).
Throughoutthe entireaxis there is relativelya strongerdevelop-
mentof the phloemthan of thexylem,the latterforminga narrower
zone than the former(fig.48). Both xylemand phloemreach their
greatestdevelopmentnear the level of the upper scales (figs.9, 22),
above this the xylem formingonly a very narrowportion of the
bundle. Throughout the strobilusthe xylem consists of spirally
thickenedtracheidswith borderedpits, the tracheidsbeing rather
short,however,although in the elongated region of the axis they
are somewhatlongerthan at a lowerlevel and the borderedpits are
fewer in number. The phloem of the portion above the scales
shows very little of the pittingpresentat a lower level, consisting
360 BOTANICAL GAZETTE [NOVEMBER

of elongatedcellssimilarto thoseof the youngerconditionof the

thexylemextendsa shortdistanceintothe
strobilus. Occasionally
stalkofthesporophyll,thebundlehere,however, usuallyconsisting

Gd Q

14 15 16
0/k

Genj
Oi9 ced (>04

17 18 19

e~~~~~~~~~~~~~~~~~~ln

20 21 22
FIGS. 13-22.-Cross-sectionsofmaturestrobilusat approximately levelsindicated
by numbersto leftof strobilusshownin fig.9; branchesof bundlesto various sporo-
phyllsindicatedby a, b, c, etc., xylemindicatedby black; bundles a and b supply
terminalsporophylls;union of bundles indicatedby combiningletters,as jd in figs.
I7-I9; fig.i9 showscompletecylinderbelow lowermostsporophyll;in fig.i8 1and c
united,in fig.i9 separated,in fig.20 c and m united,and in fig.22 lcn one of the 3
large strandsfromsterileportionof axis; fig. 22 also shows traces to 3 uppermost
scales, Sci, SC2,and sc3; note concentriccharacterof terminalportions,as in c, d, and
e, in fig.I5; X36.

onlyofthephloemportion(fig.39), thexylemusuallyendingwithin
thecortexoftheaxis.
The bundlesof thisregionare collateralendarchin the lower
portions. In theupperportions, however,thebundlesfrequently
showcentripetalxylem(figs.40, 44), givingmesarchbundles,and
19191 DUPLER-TAXUS 36i

in some cases the smallerxylemelementsare on the outside of the

bundle, indicating a possibilityof exarch structure(fig. 42). In
addition, the xylem elements, in the terminal portions of the
bundles, become more and more placed toward the center of the
bundle, giving virtually a concentricbundle of a few xylem cells
surrounded by the phloem portion of the bundle (fig. 41). No
transfusion tissue was found in
elsewherethan the scales.

Discussion
Perhaps the two most important features of the staminate
strobilusof Taxus are the peltate sporophyllsand the characterof
the vascular bundles of the scale and sporophylls. The peltate
(epaulet) type of stamen occurredamong the Paleozoic Cycado-
filicales,in the Crossothecaforms,but the sporangia were bilocular
and dehisced by a longitudinalslit along the adaxial face, the
bilocular character being differentfromthat of the moderngym-
nosperms. Peltate stamens are not known in Bennettitales,and
none occur in the Cycadales. The peltate stamen has been carried
forwardto modern plants throughthe Cordaitalean line, in all
probability,althoughso faras is known the stamensin the Cordai-
tales bore terminal erect sporangia. As COULTER and CHAM-
BERLAIN state, however,"it cannot be supposed that the stamens
of so great a group were uniformin type," and it is very possible
that peltate stamens occurred there also. The sporophyll of
Ginkgogives a suggestionof thepeltate type of stamen,in occasion-
ally havingmorethan 2 sporangia,in the regularoccurrenceofmore
than 2 sporangiain fossilforms,and in the possibility,pointed out
by Miss STARR,that the mucilage cavity replaces abortive spo-
rangia. Among Coniferalesthere is a suggestionof the peltate
stamenin the Araucarineae,and stamensof truepeltate formoccur
in such formsas Widdringtonia,Torreya,and Taxus. In Torreya
the truepeltate characteris generallyobscuredin the adult sporo-
phyll owingto the developmentof the resincavity from3 of the 7
sporangiumbeginnings. Hence it is seen that peltate stamens,in
one formor another,are scatteredfromCycadofilicalesto modern
conifers,and there is no necessityof regardingsuch a sporophyll
as that of Taxus as being of recentevolution. Assumingpeltate
362 BOTANICAL GAZETTE [NOVEMBER

sporophyllsin Cordaitalesas probable, theircontinuationin Gink-

goales and Coniferalesis quite possible, abortion of some of the
sporangia in the formationof mucilage or resin cavities, in such
formsas Ginkgo and Torreya, obscuring their true nature, but
showingthetruepeltate characterwhenall ofthe sporangiadevelop,
as in Taxus. WORSDELL, following the view put forward by
CELAKOVSKY (2), considersthe peltate sporophyllof Taxus to have
been derivedfromsuch a formas occurs in the Cordaitales, where
the pollen sacs are "erect and terminalon the radial sporophyll,"
throughsuch formsas foundin Cephalotaxusand Torreya,wherethe
pollen sacs are "sub-terminal and pendulous, owing to a slight
prolongationof the axis of the sporophyll,betweenand beyond the
sacs, in a small protuberance,"this condition being intermediate
between the Cordaitales situationand Taxus, "where the extended
terminalportionhas become enlargedand flattenedout into a very
distinct peltate structure." "Taxus thus representsan advance
from the earlier types of Cephalotaxus,Ginkgo,etc., toward the
subpeltate dorsiventraltype of sporophyllof the true Coniferae."
One must question the necessityof such an explanation foreither
the peltate sporophyllof the taxads or the dorsiventralone of most
conifers,in view of the historicaloccurrenceof both of these types
in formsmore primitivethan even the Cordaitales.
The significantfeaturesof the vascular anatomy of the strobilus
are the mesarch character of the terminal portion of the scale
bundles,as well as the appearance of centripetalxylemin the termi-
nal portionof the sporophyllbundle, where the bundle is not only
mesarch at times, but may also be exarch and concentric. This
indicates the very conservativenature of the staminate strobilus.
These primitive features, however, occur only in the terminal
portionsof the strobilus,whichmay be regardedas an argumentin
favorof the "advanced" characterof Taxus, comparedwithforms
with more abundant centripetalxylem.

Summary
i. The staminate strobilioccur in the axils of the leaves. The
buds can firstbe distinguishedfromother types of buds by the
broad apex.
r9g9] DUPLER-TAXUS 363

2. The sporophyllprimordia firstappear as slightlyrounded

lobes above the general surface and may arise in acropetal

succession.
3. The archesporial initials are hypodermalcells and develop
according to the eusporangiatemethod. There are 4-8 of them,
distributedaround the marginof the primordium.
4. The sporogenoustissue reaches the mothercell stage about
October i, and formsmicrosporesabout 2 weeks later. There is
no abortion of sporangia such as occurs in Torreya,the sporangia
occurringin a circlearound the stalk of the sporophyll.
5. The sporangium wall is usually 2-layered. The tapetum
arises from the peripheral layer of the sporogenous tissue and
persistsuntil aftermegasporeformation.
6. The epidermis of the sporangiumremains alive and thin-
walled at the base, dehiscence being accomplishedby the rupture
of these cells at maturity, by the elongation of the stalk of
the sporophyll. Owing to the disintegrationof the sporangium
wall, the epidermisis the functionalwall in the later stages.
7. The strobilusmatures the latter part of April. Just before
maturity there is an enlargementand elongation of the axis,
pushingthe sporophyllsbeyond the scales.
8. The strobiliof Taxus canadensisare somewhatsmaller than
those of T. baccata.
9. The strobilusbundles are collateralendarch,exceptingin the
terminalportionsof the scale bundles and the sporophyllbundles,
wherethey may be mesarch,and in the latter show indicationsof
occasional exarch structure,the terminalportionof these bundles
also being concentric.
HUNTINGTON,PA.

LITERATURE CITED
i. ARNOLDI,W., Beitrage zur Morphologie der Gymnospermen.III.
Embryogenie Fortunei. Flora 87:46-63. p/s.I-3. 1900.
von Cephalotaxus
2. CELAKOVSKY, L., Die Gymnospermen:einemorphologisch-phylogenetische
Studie. Abhandl. K6nigl. Bohm. Gesell. Wiss. VII. 4:i-48. i890.
3. CHAMBERLAIN, C. J., Wintercharactersof certainsporangia. BOT. GAZ.
25:I25-I28. pl. ii. I898.
4. COKER,W. C., On the gametophytesand embryoof Taxodium. BOT.
GAZ.36:I-27, II4-I40. pls. I-II. I903.
364 BOTANICAL GAZETTE [NOVEMBER

5. COULTER,JOHNM., and LAND,W. J.G., Gametophytesand embryoof

Torreyataxifolia. BOT. GAZ.39: I6I-I78. pIs. I-3. I905.
6. COULTER, JOHN M., and CHAMBERLAIN,C. J.,Morphology of gymno-
sperms. I9IO.
7. , Morphologyof gymnosperms.Revised edition. I917.
8. DUPLER,A. W., The gametophytesof Taxus canadensisMarsh. BOT.
GAZ.64:II5-I36. pis. II-I4. I9I7.
9. GOEBEL,K., Beitrage zur vergleichendenEntwickelungsgeschichte der
Sporangien. Bot. Zeit. 39:697-706, 7I3-720. pI. 6. i88i.
xo. , Morphologischeund biologischeBemerkungen. I3. lVber die
bei einigerGymnospermen.Flora 91: 237-263. figs.I9.
Pollerientleerung
I902.
W., tVberdie Entwicklungdes Pollens. Bot. Zeit. 6:425-
xI. HOFMEISTER,
434, 649-658,670-674. pis. 4-6. I848.
12. JEFFREY,E. C., The anatomyof woodyplants. I9I7.
13. JUSSIEU, A. L. DE, GeneraPlantarum.I789.
A. A., The gametophytes,
14. LAWSON, and embryoof Cephalo-
fertilization,
taxus drupacea. Ann. Botany 2i: I-23. pis. 1-4. I907.
I5. LINDLEY, J., Natural systemof botany. 2d ed.
x6. LINNAEUS, GeneraPlantarum. 6thed. I764.
17. NICHOLS, GEORGE E., A morphologicalstudy of Juniperuscommnunis
var. depressa. Beih. Bot. Centralbl. 25: 20I-24I. pis. 8-I7. figs. 4.
I9IO.
i8. PILGER, R., Taxaceae in ENGLER'S Das Pflanzenreich.I903.
x9. RICHARD,L. C., Commentatio botanicade Conifereset Cycadeis. Posthu-
mous workeditedby A. Richard. pp. 20. pI. 2. i826.
20. ROBERTSON, AGNES,Sporeformation in Torreyacalifornica. New Phytol.
3: I33-I48. pis. 3, 4. I904.
21. STARR, ANNA M., The microsporophylls
of Ginkgo. BOT. GAZ. 49:5I-55.
pl. 7. I910.
22. STRASBURGER, E., Die Coniferenund die Gnetaceen. I872.
23. Die Angiospermen
, und die Gymnospermen. I879.
24. , tber das Verhaltendes Pollensund die Befruchtungsvorgangebei
die Gymnospermen. I892.
25. , Anlage des Embryosackesund Prothalliumbildung bei der Eibe
nebstanschliessenden Er6rterungen.Festschrift zumsiebzigstenGeburts-
tage von ERNST HAECKEL. pp. i8. pis. 2. Jena. I904.
26. VON MOHL, HUGO, Uber die mannlichenBluthender Coniferen. Verm.
Bot. Schriften,pp. 45-6I. I845; published as a dissertation, I837.
27. WORSDELL,W. C., The morphologyof the "flowers" of Cephalotaxus.
Ann. Botany I5:637-652. PI. 35. I901.
28. ZUCCARINI, , Beitrage zur Morphologie den Coniferen. Abhandl.
Acad. MunchenIII. p. 794 (fromVON MOHL).
1919] DUPLER-TAXUS 365

EXPLANATION OF PLATES XXIV-XXVI.

All drawingsweremade witha cameralucida; textfigs.i-io and I3-22
are drawnto the same scale, witha magnification in reproductionof approxi-
mately36; text figs. II-I2 and all plate figs.are drawn to the same scale,
reducedone-halfin reproduction,havinga magnification ofapproximately 475.

PLATE XXIV

FIG. 23.-Young sporophyllprimordium, showing2 archesporialinitials

in hypodermallayerof roundedprimordium.
FIG. 24.-Archesporialinitialin hypoderm;divisionofepidermalcell.
FIG. 25.-Archesporialinitialin tangentialsection.
FIG. 26.-Archesporiumof 2 cells.
FIG. 27.-Metaphase in divisionof archesporial-initial.
FIG. 28.-Late stage in divisionof archesporialinitial.
FIG. 29.-Primary wall cell (outer cell) and primarysporogenouscell
(innercell), resultingfromdivisionofinitial.
FIG. 3o.-Primary wall cell and divisionof primarysporogenouscell.
FIG. 3i.-Primary wall cell has formed2 tiers of wall cells, in one of
which division is taking place; primarysporogenouscell has dividedanti-
clinally,forming2 sporogenouscells.
FIG. 32.-Lobe of youngsporophyll, showingsmall mass of sporogenous
tissue,2-layeredsporangiumwall formed on outerside, and meristematic basal
portionof epidermisdifferentiated fromremainderof epidermis.
FIG. 33.-Somewhat olderstage thanfig.32.
FIG. 34.-Sporangium wall complete,entirelysurrounding sporogenous
mass; latterpart of July.
FIG. 35.-Older sporangium,showingdifferentiation of tapetum from
sporogenoustissue.
FIG. 36.-Portion of sporophyll, showingepidermis,2-layeredsporangium
wall with narrowflat cells, tapetum (I cell binucleate), and microspores;
wintercondition.
PLATE XXV

FIG. 37.-Transverse sectionof portionof lowerscale, showingstoma on

innersurfaceand heavilycutinizedepidermalwalls,especially.onoutersurface.
FIG. 38.-Transverse section of portionof upperscale, showingvascular
bundleand innerepidermisofscale; in vascularbundlenotecentripetalxylem
and 2 transfusioncells,I dorsal,I lateralto xylem.
FIG. 39.-Transverse sectionof bundlea of fig.I3, showingphloemchar-
acter of bundle; no xylempresent.
FIG. 40.-Bundle a at level of fig.I5, showingmesarchcharacter.
FIG. 4I.-Bundle e offig.I5; singlexylemcellsurrounded by phloem.
366 BOTANICAL GAZETTE [NOVEMBER

FIG. 42.-Bundle e at a lowerlevel; largexylemcell centripetalto smaller

ones, indicatingpossibleexarchcondition.
FIG. 43.-Bundle gbat leveloffig.I 7,showingcollateralendarchcharacter.

PLATE XXVI

FIGS. 44-47.-Fusion of bundlesf and a (see figs.I5-I7): in fig.44f is

concentric,a, mesarchcollateral; section6o u below level of fig.I5; fig.45,
enlargedviewofbundlefa offig.i6, 8o u belowfig.44; fig.46, 2 bundlesnear
together20 / belowfig.45; fig.47, fusionbundlefa 30 u belowfig.46.
FIG. 48.-Transverse sectionofbundle1cmnear level of fig.22.
BOTANICAL GAZETTE, LX VIII PLA TE XXIV

\J~k~0OQ
~ 24 25 26

0 27~~0

0 ~ ~ ~ ~ 0

~~~~ 3

33~~~~43

DUPLER on TAXUS
BOTANICAL GAZETTE, LXVIII PLA TE XXV

370

40

39am4
0~~~~~~~4

41

DUPLER on TAXUS
BOTANICAL GAZETTE, LX VIII PLA TE XXVI

45

48~~~~4

DUPLER on TAXUS