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A STUDY OF TARGIONIA HYPOPHYLLA
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY I56
HERMANN DEUTSCH

(WITH THIRTEEN FIGURES)


Considerable work has already been done on the morphology
of Targionia hypophylla,LEITGEB (i) and CAMPBELL (2) being
foremost among the investigators. Recently CAVERS (3) also
published a paper on the same species; but with all this there still
remains some little ground which has not been covered at all, as
well as some which has been covered but superficially. These
pointsit is the aim of thispaper to tryto clear up. .
The materialforthisstudywas collectedin i908 by Drs. BARNES
and LAND along the steep slopes of the canyon of the Rio Santiago
in westernMexico, and also on the eastern slope of Mt. Orizaba.
In both regionsit was found only at an altitude of I500 meters.

Gametophytebody
The thallus in this group is about as complex as in any of
the Marchantiales. This statementis not based on any one of the
several charactersthat usually distinguisha thallus as simple or
complex,but on an average of the total amount of differentiation
and complexitypresent.
In the firstplace, the thallusis formedby the segmentationof a
single,cuneate apical cell (figs. i and 2), cuttingoffsegmentson
fourfaces. CAVERS (3) reportsa row of initials at the apex. In
none ofthepreparationsstudiedin thisparticularinstance,however,
could this reportbe verified. On the contrary,they seemed to
show verydistinctlya singleapical cell, distinguishedfromits sur-
roundingsegmentsboth as to its size, and also as to the size, plane,
and position of its nucleus. Relative to the apical cells found in
the othergenera of the Marchantiales, the apical cell of Targionia
is rathersmall.
The developmentof the air chamberis one featurewhich,as yet,
has not been described. This proceeds along the lines reportedby
Botanical Gazette, vol. 53] 492
I9I2] DEUTSCH-TARGIONIA 493

BARNES and LAND (4) as typical for the Marchantiales. The air
chambers arise by the splittingapart of the superficialcells just

FIG. I FIG. 2

FIGS. I, 2. Fig. I, median longitudinalsection throughthe apical cell; fig.2,


sectionin horizontalplane of thallusthroughthe apical cell.

back of the growingpoint (fig. 3). However, in the other Mar-


chantiales described,this splittingoriginatesin an angle between
the epidermaland hypodermallayersof cells,and proceedsoutward
toward the surface; in Targionia the pro-
cess is reversed,the crackingapart starting
at the surfacebetween two epidermalcells
and proceedinginward. Subsequent divi-
sions enlargethe space thus formed,as well
as the breadthof the roof.
Early in its development the pore is
closed by rapid divisions in the roof cells,
and it remains closed until the chamber
has almost reached maturity. The chloro-
phyllose filamentsdevelop comparatively
early in the historyof the chamber (figs. FIG. 3.-Nearly median

4-6); as soon as the chamber is 3-5 cells section through growing


broad, the cells of the floorbegin to project point, showing origin of
as papillae. Thesepapillaeare dividedby ai chambers.
transversewalls into filaments,which at maturityare 2-6 cells in
length. They branchprofusely,and in the mature chambervery
494 BOTANICAL GAZETTE [JUNE

often are so long that they grow snuglyup against the roof,thus
giving the impression,in section,as though therewere filaments

j~
dependingfromthe roofofthe chamber,as well as standingup from
the floor. Directly beneath the pore
the filaments
are slightly
modified,
the distal cell being hyalineand con-
taining no chloroplastsexcept one or
two lying along the bottom wall.
There is no modificationin the formof

FIG. 4

FIG. 5 FIG. 6

FIGS. 4-6.-Fig. 4, young air chamber before appearance of the chlorophyllose


filaments;fig.5, air chamberwithtwo chlorophyllosefilamentsand primordiumof a
third; fig.6, later stage of air chamber,showingdistortiondue to unequal elongation
of cells of the thallus.

these hyalinecells,as thereis in Conocephalus; theyretainthe same


ovoid shape foundin the other cells of the filament(figs.7 and 8).
The cells surroundingthe air pore are arranged in a series of
concentricrings,raised crater-likea little above the dorsal surface
of the thallus. The innermostring is composed of dead cells,
collapsed and highlycutinized; and is not, as stated by CAVERS, a
hardenedmembranousringwhich has been put forthby the inner-
most layer of cells surroundingthe pore (fig.9).
The ventral scales are placed in two rows on eitherside of the
midriband are ofan intensedark purple or red-browncolor. They
are insertedon the posteriormargin,and arrangedon the ventral
surfacein a wonderfully exact and regularfashion. On the anterior
margin of the scale is borne a curious little appendage (fig. io)
I9I2] DEUTSCH-TARGIONIA 495

which in the younger scales overlaps the growingpoint of the


thallus,and may serve as a protectivecovering. The two rows of

FIG. 7.-Mature air chamberin median longitudinalsection

scales are separatefromthe first,arisingfromyoungsegmentsclose


to the apical cell (CAMPBELL 2).
Both the pegged and the
smooth rhizoids are present.
They are extremelylong, and
most curiouslyswollen and dis-
torted at the distal end. Ac-
cording to CAVERS (3) the
smoothrhizoidsare borneon the
midrib and pass directly into
theground. The pegged rhizoids
arise in great profusionin the
axils of thescales and pass back-
ward along either side of the
midrib.
The solid,colorlesstissue of FIG. 8.-Transverse section through
thethallus
the thallu is
is composed
composed of of twoairchambers, showingtheirextreme
narrowness.
elongatedand, forthemost part,
highly vacuolate cells. These begin their elongation directly
behind the growingpoint, and this process goes on so rapidly
496 BOTANICAL GAZETTE [JUNE

that it pulls both the filamentsand the vertical walls of the air
chambers diagonally backward, givingthe chambera more or less
distortedappearance (figs.4-7).
A really surprisingamount of differentiation is seen in the cells
of the solid portionof the thallus. The commonestand most usual
is a strand of stumpy cells, filled
with oil globules and food gran-
ules, which passes longitudinally
throughthe centerof the thallus,
and ends around the foot of the
sporophyte. Besides this,thereis
frequentlya strand composed of
the ordinaryelongated cells, with
their walls thickened by an
FIG. 9.-One memberof the ringof irregularly wound tangle of fibers,
collapsed cells which surroundthe air also running the entirelengthof
pore. the thallus.

Vegetativereproduction.-The dichotomousbranching,which is
so common in the other members of the Marchantiales, is here
almost entirely replaced by the oc-
currenceof branches arising fromthe
ventral surface. These brancheshave,
forthe most part, at maturitya stalk-
like base, throughthe dying away of
which the branchesare set freeas inde-
pendent plants, and will then them-
selves multiplyin the same fashion.
In their origin,these adventitious
branches have absolutely nothing to FIG. iO.-Appendage to ven-
tral scale, which overlapsthe
do with the apical cell. This was growingpoint.
clearlyseen in several of the prepara-
tions studied,where,on a plant bearing two young branches,the
older one was placed betweenthe apical cell and the younger.
Archegonia. The archegoniaare borne terminalon the thallus.
They followso closelythe generalline ofindividualdevelopmentfor
the Marchantiales,that it is not necessaryto repeat it here. They
arise in two rowsin acropetal succession. The surface,or pad, on
I9I2] DEUTSCH-TARGIONIA 497

whichtheyare borne slopes forwardand downwardat the time the


archegoniaare mature. The developmentofthis" fruitingsurface"
runs as follows. The archegonia, arising as they do in acropetal
succession, check to a great extent the vegetative growthbehind
the growingpoint. The apical cell, however,is not immediately
concerned,and continuesits segmentationwith practicallyno inter-
ruption. The natural resultof this is that the apical cell is carried
forwardand out. Thus thearchegonialsurfaceis finallybroughtto
lie in a sort of pit or depressionin the anteriorend of the thallus.
On the lowerand outer marginof this pit is the apical cell, which,
when this stage is reached,ceases to function.
The involucre. The developmentof the involucreis so closely
allied to that of the archegonialsurface,that it is extremelydifficult
to say just where the one leaves offand the other begins. It is
simply the continued forwardgrowthof the tissues immediately
surroundingthe shallowpit at the anteriorend of the thallus,with
thenaturalresultthat theopeningto thispit is narrowed,and begins
to close up. It is never completelyclosed, however,although in
the earlier stages the edges of the wings are brought very close
together. As the involucrematures,the outer surface,as well as
the innermarginof the wings,becomes highlycutinized.
That the involucreis not, as reportedby CAVERS (3), a result
of the stimulusgiven by the act of fertilizationis shown in one of
the preparations,whereit was completeand well developed, while
the eggs of the archegoniait surroundedwere stillunfertilized.
Calyptra. The calyptrais a simpleand very delicate structure,
soon rupturedby the growthof the sporophyte. At maturityit
is 2-4 layers thickat the base.

Sporophytebody
Capsule. The capsule is a comparativelysimple affair. The
wall consists of a single layer of cells, uniformlythickenedwith
spiral and annular bands. There is no special mechanism for
dehiscence. In his recentreporton Targionia, CAVERS(3) figures
and describes a rudimentaryelaterophore,representedby a few
spirallythickenedcells depending fromthe distal portion of the
capsule. This statementcould not be verifiedin any of the sections
498 BOTANICAL GAZETTE [JUNE

studiedhere,nor do either CAMPBELL (2) or LEITGEB (i) reportsuch


a condition.
True elaters are present,long, slender,fusiform, and thickened
by usually two spiral bands.
The sporesthemselveshave fairlythickwalls, highlysculptured,
and are produced in great numbers (fig. ii). CAVERS (3) has
described the spore mother cells as lobing
deeply beforedivision,as happens in many
of the Jungermanniales. All of the prepara-
tions studied here, however,show the ordi-
1N~t; nary tetrad formed froma spherical spore
mothercell.
Seta and foot.-Both seta and foot are
i-Tbcat
FIGre. rather well developed. While small as com-
pared with the size of the capsule, theyare
neverthelessrather massive, and well differentiatedfrom each
other. The foot is club-
shaped, and bluntly
pointed at the lower end
(fig.I 2). The surfacecells
project as short papillae,
and are haustorialin func-
tion. These cells, as well
as those of the calyptra
surroundingthem,show in
sectiona verymuch darker
stain than do the cells of
the surroundingtissues.
Between the calyptra and
foot is interposed a fairly
thicklayerofmucilaginous FIG. I2.-Foot and seta
material.
Although6-iO archegoniaare produced, only one egg is fertil-
ized. The remainingarchegonia, however, persist, and are still
visible afterthe sporophytehas matured; soon afterthe sporophyte
has begun its development,however, their contents break down
into a darklystainingmucilaginousmass.
I9I2] DEUTSCH-TARGIONIA 499

Classification
The Marchantiaceae are at presentclassifiedaccordingto three
schemes. LEITGEB (i) gives the following:
Marchantiaceae
a) Astroporae(Clevea, etc.)
b) Operculatae(Plagiochasma,Fimbriaria,etc.)
c) Targioniae(Targionia,Cyathodium)
d) Compositae(Marchantia,Lunularia,etc.)
CAMPBELL (2, p. 67) gives the following:
Marchantieae
a) Corsinieae
b) Targionieae
c) Marchantieae
GOEBEL has proposed still another classification. In this he trans-
poses the Riellaceae to the Marchantiagroup,as follows:
a) Corsiniaceae
b) Targioniaceae
c) Riellaceae
d) Marchantiaceae

Without concerningourselveswith the relative meritsof these


schemes,it mightbe well to take up the one family,the Targionieae.
This family at present contains but two genera, Targionia and
Cyathodium, and it is withregardto the meritsof placing these two
genera withinthe same familythat a question may be raised. In
orderthe more clearly to presentthis question, the accompanying
diagrams (fig. I3) have been prepared. In the key to these
diagrams it will be seen that each generation (sporophyte and
gametophyte)has been divided into fourfeatures,selected because
of their bearing on a natural scheme of classification. Each of
these divisions has been subdivided into five stages of as nearly
equal importanceas it was possible to find.
From the diagram it will be seen that the two lines coincide in
but two points. The one of these is D4 and the other E4, which
representrespectivelythe apical positionof the archegoniaaccom-
panied by a checkingof the growthof the thallusat thispoint,and
the commoninvolucre.
As for the former(D4), the character is not peculiar to this
500 BOTANICAL GAZETTE [JUNE

group. In several other genera the archegonia are borne in a


terminal cluster, notably Grimaldia, Reboulia, and Clevea. This
narrowsthe situation down to a singlecharactercommon to these
two genera (Targionia and Cyathodium)and not found in others.
This is the common involucre enclosing the terminal group of
archegonia.
One glance at the diagramswill show how widelythe two genera
differin all other respects save this. Targionia has perhaps as
complex a thallus as is foundin the entiregroup of Marchantiales;
A B C D B Y G H A B C D E F G H
5 5
4 4
3 3
2 ~~~~~~~~~~~2
Targionia Targionia

A B C B E V G H A B C B H F C H

4 4
3 ~~~~~~~~~~~3

Cyathodium Targionia and Cyathodi=x

FIG. I3.-Diagrams comparing Targioniaand Cyathodium

Cyathodiumhas perhaps the simplest,both as regardsstructureand


the amount and character of differentiation.The thallus of
Cyathodiumconsists of a simple ribbon, two layers of cells thick,
the two layersslightlyseparated to forman air space, the air pores
being simpleopeningsin the upper layer of cells.
In Targionia the antheridiaare borne on a special portion of a
special branch; in Cyathodiumthey are scatteredin clustersalong
the edge of the thallus. In Targionia the foot and seta of the
sporophyteare massive and well differentiated;in Cyathodium
both foot and seta are representedby a single filament,four cells
in length.
In Targionia the elaters are true elaters, long pointed, and
spirallybanded; in Cyathodiumthe elaters,while spirallybanded,
are littlemorethan dead nursecells, shortand stumpy.
In but one character is Targionia less advanced than
Cyathodium,and that is in the differentiation of the capsule wall
I9121 DEUTSCH-TARGIONIA 50I

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502 BOTANICAL GAZETTE [JUNE

in the mature sporophyte. In Cyathodium,only the cells of the


upper part of the capsule are spirallythickened,so that dehiscence
occurs throughthe formationof 6-8 fairlyregularteeth.
Of course there are several difficultiesin the way of such a
determiningscheme as this. In the firstplace, it is next to
impossibleto select charactersthat will be absolutely determining.
In the second place, it is impossibleto select groups of characters
whosedetermining value willbe equal. Again, thereis the difficulty
of deciding as to which of two charactersis the more advanced.
And finally,thereis the conflictas to the relative value of sporo-
phytic and gametophyticcharacters. Is a complex gametophyte
with a simple sporophytemore advanced than a simple gameto-
phytebearinga complexsporophyte,or vice versa; to say nothing
of the differentgradationsin combiningthe two.
Leaving aside, forthe present,the difficultieswhich a practical
applicationof this schemepresentsin general,it does seem to apply
in the specificcase under discussion; no matterwhich of the two
is the higher,it seems to be fairlycertain that they are widely
different. Accordingto the diagrams the two genera have but the
one salient featurewhich is common to them and to them only, a
single involucreenclosinga group of terminallyborne archegonia.
Now it certainlydoes not seem as thoughsuch a characteras this
should be sufficientto bind two genera differingso widely in all
otherrespectsinto one family.
As to where Cyathodiumbelongs, if not with the Targionieae,
the Corsinieae suggest themselvesreadily. But such a matter as
this cannot be settled definitelywithout a much more extended
study of Cyathodium,as well as the familyCorsinieae,with its two
genera (Corsinia and Funicularia), than has been given them.

Thanks are due ProfessorJOHN M. COULTER and Dr. W. J. G.


LAND forassistancerenderedduringthe progressof the work.
THE UNIVERSITY OF CHICAGO
I9I2] DEUTSCH-TARGIONIA 503

LITERATURE CITED
i. LEITGEB, H., Untersuchungen
uiberdie Lebermoose6: I3I-I36. i88i.
2. CAMPBELL,D. H., Mosses and ferns.I905.
3. CAVERS,F., Contributions to the biology of the Hepaticae. Part I. Tar-
gionia,Reboulia,Preissia,Monoclea. I904.
4. BARNES, C. R., and LAND, W. J. G. The originofair chambers. BOT. GAZ.
44: I97-2I3. I907.