Torrey Botanical Society

Studies on American Hepaticae-I. Revision of the Genus Thysananthus

Author(s): Margaret Fulford
Source: Bulletin of the Torrey Botanical Club, Vol. 68, No. 1 (Jan., 1941), pp. 32-42
Published by: Torrey Botanical Society
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STUDIES ON AMERICAN HEPATICAE-I. REVISION OF THE
GENUS THYSANANTHUS

MARGARET FULFORD

(WITII FIFTY-ONE FIGURES)

Three species of Thysanantthus, a gelnus of the subgroup Holostipae of

the Lejeuneae, have beeln reported from the Americas and a fourth is here
proposed as lnew. Spruce1 has described several species under Lejemuiea,
subogenusVIII, Thysano-Lejeitnea, lnamely L. arnazonica from the Amazon
coulntry,L. dissoptera from Guialna, and L. pterobryoides from Ecuador. In
addition to these, Nees von- Eselnbeek2 described a plant from Jamaica as
PhragnmiconiaLehmanniana, which Stephanii later transferred to Thysanan-
thts. Verdoorln3suggested that this plant was a Catdalejeunea. All examina-
tiolnof a portion of the tvpe material shows that it is idelntical with Cautdale-
jeurnea Lehmanlriaria (Gottsche) Evalns4 and it should therefore be reduced
to synOilivnlv.Taylor5 described plalnts collected in Ecuador bv Professor W.
Jameson unider the name Thysananthuts mexicanius. His plalnts do lnot have
the characteristics of the genus Thysananthus as it has been delimited by
recent moliographers so that the species is a member of some other gellus of
the Lejeuneae.
The genus6 as ulnderstoodby Evalns, Verdoorl,8 ald others may be char-
acterized as follows: the female ilnfloreseenceis terminal olnthe maii stem or
prilncipal branch, with ininovatiolnsproceeding from one or both sides below;
the perianth is 3-angled in transverse section, with the ventral keel sharp
andl distinct; there are no secondary folds or ridges; carinal and surface
wilngs are developed in some species; the female bracts and bracteoles, and
the keels of the periaiith are toothed to a greater or lesser degree; -and the
leaves and underleaves usually show some indicatiolns of teeth along the mar-
ginls,particuLlarlyin the apical region.
The plants are large and grow in depressed mats or among other bryo-
phytes on the trunks and bases of trees and over logs. The stem is robust and
1 Hepaticae of the Amiiazonand Andes. Tranis. Bot. Soc. (Edinburgh) 15: 105-110.
1884.
2 In Gottsebe,Liiidenbelg & Nees von Eseiibeek, Syii. Hep. 302. 1845.
3 Die Frullaniaceis XV. Die Lejeuiieaceae Holostipae der Indomalaya uniter Be-
riicksichtigulng siimtliceleiaus Asien, Australieln,Neu-seelalndulnd Ozealnien alngefiilrten
Alten. XIII. Thlysananthots.Anin.Bryol. Supl. 4: 163-188. 1934.
4 Hepaticae of Puerto Rico VIII. Cautdalejeatnea. Bull. Tolrey Club 34: 554-557.
pl. 33. fig. 1-12. (1907.) 1908.
5 On some new Musei collected by Professol W. Jamesoli oln Piehilnela. Hookel 's
Jour. Bot. 7: 187-199. 1848.
6 For synlonymy in the gelnus Thtysavanthlais see Verdoorn, 1. c.
7Hepaticae of Pueirto Rico VIII. Symibiezidihtw,Marchesinia, Mastigolejellnea,
Caatdalejeainea and Br-yopterii. Bull. Tolrey Club 34: 533-568. pl. 31-33. (1907.) 1908.
32

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1941] FULFORD: THYSANANTHUS 33

is differelitiatedinto a prostrate primary caudex which is appressed to the

sulbstratulm,and ascendin(g or upright secondary stems which are usually
abulncdantlybranched. The branches are olive or dark green alid often beconle
deeply-pigmeintedwith browi.
Evans8l describes the secondary stem of T. anuazoniicts as beilig 3-8 cm.
in lenoth aind about 0.15 mm. in diameter. The tralnsverse sectioli shows a
uiiforlil orange-brown pigmentation except for the slightly darker middle
lamellae. The cortex is made up of 30-33 rows of thick-walled cells ranging
from 10 to 20 p in width, aid from 10 to 15 p in thickness. The cells of the
medulla are quite similar, except that they have distinctly larger cell-eavi-
ties, so that the medulla appears somewhat more open than the cortex.
The branchilng of the subfloral innovations, other vegetative branches,
and the male branches seem always to be of the Radutla type.9
The lobule is small in proportion to the lobe. The keel is rounded and the
upper edoge,wvhichis entire, tends to be tightly appressed to the lobe, so that
a definite,ilnflatedwater-sac is usually formed. The apical tooth is from one
to seven cells long and may be curved.
Although the genus, as indicated by the stem structure, is clearly a
member of the Holostipae, which are characterized by entire uiiderleaves,
several species have retuse or distinctly bifid underleaves. Three of the Amer-
ican species show this characteristic. The additional generic characteristics
are discussed uiider the descriptions of the separate species.
The writer wishes to acknowledge the helpful criticism of Dr. A. W.
Evans of Yale University, and the courtesies extended by the New York
Botanieal Gardeln during the preparation of this paper.
The following abbreviations have been used to desiglnate the location of
specimenisin the citations under the individual species: M, Missouri Botan-
ical Gardeii; NY, New York Botanical Garden; and Y, Herbarium of Yale
University, including the private collection of Dr. A. WV.Evans.
KEY TO THE SPECIES

1. Undeileaves obovate, rarelytruncate,the apical margin serrate . 4. T. cowIosits

1. Uindelleavessubquadrate-cuneate,from retuse to deeply bifid.
2. Lobule oblong, the tooth if preselntof only olie oi a few cells;
leaf cells elliptical-angularin outline.. 3. T. aviazoniciis
2. Lobule more or less subqiuadrate,the tooth long, curved,of 5-7
cells; leaf cells quadrate in outline; underleaves distinctly
bifid.
3. Margilis of the leaves and uliderleaves dentate with coarse
teeth.2.lT T.
pterob.yoides
terobe.............................................................................................................................t....e2.
3. Margins of the leaves and underleaves from obscurelyden-
tate to denticulate...... 1. T. Evansii
8 Anatomy of the stem in the Lejeuneae. Bull. Torrey Club 62: 187-214; 259-280.
8 fig. 1935.
9 For a description of the various types of branching see Evalns, Bralncllinigin the
Leafy Hepaticae. Ann. Bot. 26: 1-37. 36 fig. 1912.

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34 BULLETIN OF THE TORREY CLUB [VOL. 68

Thysananthus Evansii Fulford, sp. nov. Caules robusti, olivacei, pinnate

ramosi, feminini dichotomi; folia imbricata, divaricata, ovata, 1-1.3 mm.
longa, superne obscure serrata, cellulis 8-10 p dianm.,lobulo parvo, dente
apicali longo; foliola imbricata, subquadrato-cuneata, bifida; flores dioici;
bracteae femininae foliosae, lobulis magnis, planis, ovatis, bracteola magna,
ovata, carinata, crasse dentata.
Plants robust, olive-green, becoming darker green in the older portions:
secondary stems stout, to 4 cm. or more in length, with leaves to 2 mm. broad,
ascending, irregularly pinnate, the branches obliquely spreading, the female
branchsystemshowingapparentlyregulardichotomy:leaf insertioncurved
in theupper part; theleaves imbricated,spreading,becominga littledeflexed
whendry,unsymmetrically ovate,1-1.3 mm.long,0.5 mm.broad at thebase,
obseurelyserrateabove the middle,the dorsal base cordate,the ventralmar-
gin straight;the lobule small, inflated,ovate, 0.12 mm. long, 0.1 mm.high,
the free margin entire,appressed, the apical region of the lobe 8-10 p in
diameter,cells of the base much longer,a vitta not differentiated, the cell
walls uniformlythickened,the cell luminarounded,trigonesnot evident,the

FIGS. 1-12. T. Evansii Fulford. FIG. 1. Diagram of the branching patterli of a

female plant. FIG. 2. Portion of a plant, ventral view, x 15. FIG. 3. Portion of a leaf and
stem, dorsal view, x 30. FIG. 4. A leaf, x 30. FIG. 5. A cell from the apical portion of a
leaf, x 400. FIG. 6. Outline of the apical margin of a leaf, x 300. FIG. 7. Lobules, one of
them has developed two apical teeth, x 90. FIG. 8. Apical teeth of lobules, x 400. FIG. 9.
Underleaves, x 30. FIG. 10. Outline of the apical margin of a tooth of an underleaf,x 300.
FIG. 11. Female bract, x 30. FIG. 12. Female bracteole, x 30. Drawn fromthe type mate-ial.

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1941] FULFORD: THYSANANTHUS 35

cuticle smooth to fainitlyv verruculose: underleaves inmbricated,attached in a

straight lilne,subquadrate-euiieate, 0.7 mm. long, 0.5-0.6 mm. broad in the
upper part, retuse or bifid to olne-fifththe lenigth,the broad and poilnted divi-
siolns altd the obtuse to lunulate sinus serrate, the lateral margins convex,
elntire,the cells as in the leaf: dioicous: female branches short, oln the main
stem or branches, always subtendled by two subfloral inllovations, a pair of
leaves below the ilnvoluere intermediate in form betweeln the bracts alnd the
liormal leaves; the braets leaf-like in outlilne,1 mm. or more long, the mar-
gills oftelnmore strolngly serrate thaniin the leaf, the lobule ovate, much en-
larged, plane, mostly 0.7 mm. lolng,the apical tooth not evident; the brac-
teoles broadly ovate, 1.3 mm. or more long, the maroini coarsely dentate, the
apex obscurely bifid, keeled nearly to the apex, the alngle broad: male
brainehes,perialnthsalnd sporophytes not seeln.
Habitat: On bark of trees, lowland forests.
Distributioln: BRITISII HONDU7RAS: Pulnta Gorda, withouitcollector's lnalue,
TYPE (M).

The distinguishing characteristics of the species are its olive-greelncolor

andl dichotomously bralnehed femualestems; the long, curved tooth of the
lobule of the leaf; the bifid ulnderleaves serrate above the middle; alnd the
long,,keeled, more or less bifid, delntate,female bracteoles. See figures 1-12.

THYSANANTIIIJS PTEROBRY0IDES (Spruce) Stephani, Spee. Hep. 4: 786.

1912. Lejeunea (Thysano-Lejeunea) pterobryoides Spruce, Trans. & Proe.
Bot. Soc. Edilnb. 15: 109. 1884. Bryopteris Wallisii Stephani, Hedwigia 24:
89. pl. 1, fig. 1-8. 1885. Thysanolejeutnea pterobryoides Stephaini (syn.),
Spee. Hep. 4: 786. 1912.
Plants olive ogreen,becoming darker green in the older portiolns: see-
ondlary stems to 8 ciii. or more in lenogth,with leaves 2 mm. broad, aseend-
ing, more or less reogularlyand simply pinnate wrhensterile, infloreseences
produced oln primary branches of pilnnate branch systems; leaf insertion
curved in the upper part; the leaves imbricated, spreading, becoming a little
deflexed when dry, 1-1.3 mm. long, 0.4-0.6 mm. broad at the base, unsym-
metrically ovate, coarsely dentate above the middle, the dorsal base cordate,
extelndino aeross the stem arid beyond, the ventral margil nmore or less re-
curved, entire, the lobtile inflated,ovate, 0.7-0.12 mm. long, 1.8-0.1 mm. high,
the free margin appressed, elntire,the apical tooth five to seven cells long,
eirved, the sinus deep, luimilate; the cells of the lobe averaging 8-10 p in
diameter, those of the basal area longer, a vitta lnot differentiated,the cell
walls uniformlv thiekened, the cell lumina rounded, trigones not evident:
ulnderleaves imbricated, attached in a straight line, coarsely dentate, sub-
quadrate-tuneate, 0.56-0.7 mm. long, 0.5-0.63 mm. broad in the upper part,
bifid to one-fourth the length, the sinulslunulate, the teeth broad, acute, the
cells as in the leaf: dioieous: female branches short, always with two sub-
floral innovations, a pair of leaves below the involuere intermediate ill form
betweeii the bracts aind normal leaves; bracts leaf-like in outline, to 1.3 mm.
or nmorelong,,the lobule lanceolate, plane, more than half the length of the
lobe, the apical tooth not evident, the lobe and lobule coarsely dentate above
the middle; the bracteole oblono, to 1.5 mm. in length, bifid and coarsely
toothed above the middle as in the underleaf, keeledl nearly to the apex, the

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36 BULLETIN OF THE TORREY CLUB [VOL. 68

angle acute: perianthto 2.5 mm.long,thekeelssharp,crenlulate,occasionally

serrate,thebeak short:male branchesand sporophytesnot seen.
Habitat: On bark of treesin forests.
Distribution:COSTA RICA: San Jos', Lehmalnn,cited by Stephani.10PAN-
AMA: Island of Coiba, Seemann (Y, NY). COLOMBIA: Cordoba,Killip 11776,
11801 (Y, NY); withoutlocality,Wallace (NY). ECUADOR: Pastaza River,
Spruce 1885, 109; Manabi, IVallis, cited by Stephaui (1885, 89), TYPE of
Bryopteris Wallisiti.
The distinguishingcharacteristicsof the species are its olive-greencolor,
the simply and regularlypinnate branching,the long apical tooth of the

fI
26
27
13

14 21

18 ~ ~ 6

22 --~~~~~~~~~~~~~~2

FIGS. 13-27. T. pterobryoides (Spruce) Steph. FIG. 13. Diagram of branching pat-
tern of a sterile stem. FIG. 14. Diagram of the branching pattern of a female stem. FIG.
15. Portion of a plant, ventral view, x 15. FIG. 16. A leaf, x 30. FIG. 17. Outline of the
apical margin of a leaf, x 300. FIG. 18. A cell from the apical portion of a leaf, x 400.
FIG. 19. Cells from the basal portion of a leaf, x 300. FIG. 20. A lobule, x 90. FIG. 21. An
apical toothof a lobule, x 400. FIG. 22. Underleaves, x 30. FIG. 23. Outline of the apical
margin of a tooth of an underleaf, x 300. FIG. 24. Female bract, x 30. FIG. 25. Female
bracteole, x 30. FIG. 26. Upper portion of a perianth, dorsal view, x 30. FIG. 27. Upper
portion of a perianth, ventral view, x 30. Drawn from material collected by Dr. Killip in
Colombia.

10 In Cryptogamae Centrali-Americanae.Hepaticae. Bull. Herb. Boiss. 2: 402-403.

1894.

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1941] FULFORD: THYSANANTHUS 37

lobule of the leaf, the bifid underleaves, alnd the coarsely dentate marglins of
the leaves and underleaves. See figures 13-27.
T. pterobryoides all d T. Eva nsiihave many characters ill co0n1oi0,lnamely
the color alid size of the plalits, the size of the leaves and ulnderleaves, the
size anid shape of the leaf cells, the lobules of the leaves, and the production
of pairs of subfloral inniovationissubtending the female infloreseeinees.They
differin that in T. pterobryoidesthe secondary stems are often very lonc,
the branehinigis simply and regularly pinnate, and the female infloreseences
are produced on primary branehes of a pinnate branch system, while in T.
Evansii the branches are less regularly pinnate and not so lonc, and the
female plants always show an apparently regular dichotomy. (Compare figs.
13, 14 with fig. 1.)
The margins of the leaves and underleaves, aiid the female bracts and
bracteoles of the two are striking,lydifferent.(Compare figs. 15, 16, 17, 22-25
with figs. 2, 4, 6, 9, 10-12.) The shape of the female bracts is quite similar in
the two species, but the bracteoles of T. Evansii are broadly ovate aind the
angle of the keel is broad, while in T. pterobryoides the bracteoles are oblong
and the angle of the keel is acute. Unfortunately no perianths have been
founldin T. Evansii, so that a comparison of this structure in the twvospecies
cannot be made at this time.
Not oinlydo T. pterobryoidesand T. Evansii have many charaeteristies
in common but they also make up a unit within the genus which is,different
not only from the other South American forms but also from any of the
Asiatic species with which I am familiar. All the other species have a deep
brown pigmentation and have large leaf cells which are more or less elonigate
and hexagoonal or anigular-elliptical in outline, with conspicuous trigolnes,
and often additional intermediate thickeninogs.
THYSANANTHUSAMAZONICUS(Spruce) Stephanii, Spec. Hep. 4: 784. 1912.
Lejeitnea (Thysano-Lejeutnea) amazonica Spruce, Trans. Bot. Soc. Edinb.
15: 106. 1884. Thysaunolejeunea amazonica Stephaiii (svn.) I.c.
Plants dark greeiuishbrown, becoming very dark browiinin the older por-
tions; seconidary stemrscoarse, 6 em. or more in length, with leaves to 3 mm.
broad, irregularly pinnlate, the branehes oblique, ofteii braliched, female
branehes with only one subfloral ininovation: leaf iiisertion curved in the
upper parts; the leaves imbricated, spreading, tightly appressed to the stem
when dry, averaging 1.4-1.6 mm. longy,0.8 mm. broad at the base, unsym-
metrically ovate, entire except for an occasional serration, the dorsal base
cordate, covering the stem and extendinlgbeyond, the veiitral margin entire,
recurved; the lobule small, inflated,oblong in outline, 0.3-0.42 mm. lonig,0.15
mm. high, the free margin entire, appressed, the apical tooth sometimes evi-
dent, of one or two cells, the slime papilla not seen, the silius shallow, llitulate;
cells of the marginal alid apical region and of the lobuile 16-20 p x 8 , those
of the base loliger, a vitta Hot differentiated,the cell walls thin, the trigones
colispicuous, with convex sides, soon becomrig confluienit through the deposi-
tion of secondary material, intermediate thickenilig frequent, the cell ltmilna

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38 BULLETIN OF THE TORREY CLUB [VOL. 68

angular-oblongin outline,the cuticlesmoothto verrueulose:underleavesim-

bricated,attached in a straightline, cuneate,retuse to shortlybifid,aver-
aging 0.77 mm.long, 0.7 mm.broad in the upper part, the teethvery short
and broad, the sinus shallow, lunulate, the margins entire,sometimesob-
seurelyserrateon underleavesnear the tip of the stem,the cells as in the
leaf: autoicous: male infloreseencesterminalon shortlateral branches,the
bractsin twoto fourpairs,at thetip of a branch,imbricated,thelobe broadly
ovate, the lobule ovate, smaller,the apical tooth conspicuous,the margins
sparselydentate; the bracteolesimbricated,similarto the underleaves; an-
theridiain pairs: femalebranchesshort,occurringsinglyor several approxi-

39/
34 34

28

40

29 ~<
y

32

,135
37
~~~33
FIGS. 28-40. T. amazonicus (Spruce) Steph. FIG. 28. Diagram of branching pattern
of a stem. FIG. 29. Portion of a plant, ventral view, x 15. FIG. 30. Portion of the apical
margin of a leaf, x 30. FIG. 31. A cell from the apical portion of a leaf, x 400. FIG. 32.
Cells fromthe basal portion of a leaf, x 300. FIG. 33. Lobule, x 90. FIG. 34. Lobules, x 30.
FIG. 35. Underleaves, x 15. FIG. 36. Female bract, x 15. FIG. 37. Female bracteole, x 15.
FIG. 38. Outline of a cross section of the perianth. FIG. 39. Upper portion of a perianth,
dorsal view, x 30. FIG. 40. Upper portion of a perianth, ventral view, x 30. Drawn from
type material.

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1941] FULFORD: THYSANANTHUS 39

mate, appearing lateral, a suibfloralinnovation from onilyonieside, the bracts

similar in outline to the leaves, averaging 2 mm. long, the lobule oblong,
ovate, plane, 0.8 mm. long, the margins of both coarsely toothed in the upper
part; the bracteole oblong, bifid, 1.6 mm. or more in lenigth,averagillng0.9
mm. broad in the upper part, the teeth broad, acute, the silluLsacuite, the
margins of the teeth serrate and coarsely dentate: perianth to 2 mm. long,
the beak short,the keels serrate and dlentatewith scattered teeth: sporophyte
not seen.
Habitat: On bark of trees in forests.
Distribuition: TRINIDAD: Mora Forest, E. G. Britton 2878 (Y, NY) ; with-
out locality, Fendler (NY). COLOMBIA: Without locality, Wier (NY). BRAZIL:
Para, Spruice, Hepat. Sprue., TYPE (Y, NY). BRITISH GUIANA: nlear Bartica,
Richards188, 510 (Y).
The distinguishing characteristics of the species are the large, coarse,
irregularly branched, dark greenish brown stems; the large, ovate, entire
leaves with oblong lobules which have a 1-3-celled apical tooth; the long-
angular cell lumina, large trigones alnd intermediate thickeninigs ancd the
large eiliueate underleaves which are retuse or somewhat bifid at the apices.
See figures 28-40.
T. arniazonicuts
is readily distinguished from the preceding species because
of its atutoicousinfloreseence,the deep brown pigmentation, the entire mar-
gins of the leaves and underleaves, the short apical tooth of the lobule, an-d
the anoglar-oblonli outline of the leaf cells.

THYSANANTHUS COMOSUS ilindenberg, in LehmanllnPug. P1. 8: 25. 1844.

Lejeanieaa contosa Mitten, Jour. uinn.Soc. Bot. 5: 109. 1861. Lejeutnea (Tlhy-
sano-Lejeiuea) comosa Spruce, Trans. Bot. Soc. Edinb. 15: 108. 1884. Thy-
sanaithtusdissopteruts
Stephani, Spec. Hep. 4: 784. 1912. Thysanolejeutnea
dissopter-aStephani (syn.) I.e.
Plants dark greenish browli, becoming very dark brown in the older por-
tionis; stemnscoarse, 5 cm. or more in lenlgth,with leaves to 2.5 mm. broad,
irre2ularly piniate or bipinnate, microphyllous branches frequelt, female
branehes with olnly one subfloral iniovation: leaf insertioni eurved in the
upper part; the leaves imbricated, spreading, aseending, and appressed to
the stemnwhen dry, averag,inig1.4 mm. long, 0.85 mm. broad at the base, un-
symmetricallyovate, entire except for an occasional serratioli, the dorsal base
cordate, coverilio the stem and extelndingbeyonld,the ventral maroin enitire,
recuirved: the lobule small, inflated, ovate in outlilne, 0.3-0.4 mm. long, 0.2
mm. hioh, the free margin elntire,appressed, the apical tooth one to several
cells long, the inier (proximal) tooth sometimes evident, of one or two cells,
the silnuisvery shallow; cells of the lobule and the margin and apical reoions
averaoino 20 [1 long, 12 [p wide, those of the base loniler, a vitta not differen-
tiated. the cell walls thin, the trigone conspicuous, with convex sides, ofteni
becomingocoalesced, intermediate thickeniingsfrequent, the cell lumina an-
gular-obloiio in outline, the cuticle smooth or faintly verruculose: undlder-
leaves imbricated, attached in a straight line, obovate to cuneate, averagilln
0.7 mm. lono, 0.7 mm. broad in the upper part, the maroin serrate, the cells
as in the leaf: autoiceous ( ?) or dioicous: female branches short, occurrinlg

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40 BULLETIN OF THE TORREY CLUB [VUL. 08

sinoly, with one subfloral innovation, thus appearing lateral, the bracts simi-
lar in outline to the leaves, averaging 1.3 mm. long, the ventral maroin deln-
tate, the lobule large, 0.85 mm. long, plane, more or less obovate, obscutrely
bifid, the margins denitate; the bracteole eunleate, similar to the underleaf,
0.9-1.2 mm. long, the margins serrate and coarsely dentate: the perialnth
mostly 1.5 mm. long, the beak short, the keels delnsely set with large irregu-
larly tooth laciniae, the dorsal face smooth, the venitral faces each w-ithonle
or more groups (in one or two rows) of laciniae similar to those of the keels:
male branches and sporophytes not seen.
Habitat: On bark of trees in forests.
Distribution: GUIANA: Hb. Hooker, (Y, NY). BRAZIL: Tatnat. Spruice
(NY). Also in the Ilndo-Malayani Region.

The distinguishing characteristics of the species are its robust lhabit,dark

color, the lnearly entire leaves, alnd small lobules with a short apical tooth;
the long(-ang,ularcell lumina and conspicuous trigolnes; the obovate under-
leav es serrate along the upper margins; and the laciniate willnS of the peri-
auth. See figures41-51.
T. anazonicuts and T. conosius are very similar in maniyrespects. niamiely,
size of plant, growth habit, and, in a genieralway, the shape of the leaf and of
the leaf cells. The underleaves of T. comosstS, when well dev-eloped, are
obovate with a serrate margin (figs. 43, 48), while in T. a)niazon1icu(s (figs. 29,
35), although they are more or less obovate, the apical reoion is distilnctly
retuse or more or less 2-toothed. The apical margin is serrate to a greater or
lesser degree.
is to be found in the parts of tlle felmale
An additiolnal striking distilnctioni
involucres of the two species. The bracts of T. amazonicus are dlentate at the
apices, and the lobules are long, niarrow, and conispicuously bifid with the
margins occasionally toothed in the upper part (fig. 36), while those of T.
COIOSitS are delntate along the ventral margins, and the lobules are lolng,
ovate, obseurely bifid, with the maroins coarsely dentate to t.he base (fig.
49). The bracteoles of T. aniazonicuts are oblong, bifid, and coarsely-dentate
in the upper part, while those of T. coniosutsare cuneate, andl from serrate
to dentate througThout.(Compare figs. 37 and 50.) The laciniate wingoson
the keels of the perianths of T. coralosuts will immediately distinogish that
species from T. amazoPic?ts, in which the perianth keels are only sparingly
dentate (figs. 39-40, 51).
The leaves and unlderleaves in the vieinity of the female bralnelhesanld
sometimes at the tips of the stems are usually more strongly toothed than
elsewhere on the plant. When toothed leaves ocecur in the region of the
female branehes they are ilntermediatein fornmbetweeniordinary-leaves and
the bracts and bracteoles.
All the South American nmaterialexamined was dioicous. Spruce de-
scribes the South American material (under Thysano-Lejenaca dissoptera)

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1941] FULFORD: THYSANANTHUS 41

41

I p~~~~~51)

F^IGS. 41-51. Tr.cosmnos Lindenb. FIG. 41. Diagram of branching patternlof a sterile
stem. FIG. 42. Diagram of branching pattern of a female plant. FIG. 43. Portion of a
plant, ventral view, x 15. FIG. 44. A leaf, x 30. FIG. 45. A cell from the apical portion
of a leaf, x 400. FIGL. 46. Cells from the basal portion of a leaf, x 300. FIG. 47. Lobule,
X 30. FIG. 48. Underleaf, x 30. FIG. 49. Female bract, x 15. FIG. 50. Female bracteole, x 30.
FIG. 51. Ventral faces of a perianth,x 30. Drawn fromthe orsiginalcollection fromGuiana.

as "dioica (?)"; Stcphani11describesboth T. dissopterusand T. comlosus

as " diojeous," while Verdoorn's description11states that the species is
"usually monoicous." The portionof the type materialfromPaulo-Penang
which I examinedhad many well developed perianthsbut I could findno
male branches.
There has been somedifference of opinionas to whetherthe South Amer-
ican plants were actually T. cornosus,a species widespreadin Judo-Malaya.
Ljindenbergin the original descriptioncited plants in "Paulo Penang, ubi
legit cel. Wallich," also " in Guiana lectam accepi ab illustr. Hooker."
Spruce13consideredthis interpretationto be incor-rect and separated the
11 Spec. Hep. 4: 784, 787. 1912.
12 Verdoorn,op. cit. p. 176.
13 Trans. Bot. Soc. Edinb. 15: 107-109. 1885.

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42 BULLETIN OF THE TORREY CLUB [ VOL. 68

South American plauits from the "composite species," T. coinosits, anld

called the South Americaii plants Lejetnea (Thysano-Lejea nea) dissoptera.
He also described the material from Paulo Penan(g, retaining for it the spe-
On the other hanid, Mitten14 considered the two to be
cific epithet comitosuts.
identical. Stephanii's comments15 concernilng the Guiaiia material are
obscure, for he says (discussing T. cornosuts),that only Nos. 85 anid 86 rep-
resent the true species, moreover the American No. 84 is very similar. Ver-
doornl16 has followed Liindenberg in conisidering the South Americaii and
the Asiatic material idelitical. In my observationis of a portioinof the type
material from both Guiaana and Paulo-Penang in the Mittein collectioll I
have not been able to discover any variations of sufficientdegree an-d uiii-
formityby which the two could be separated. The two seem to be idelntical.
Except for the original material from Guiana, I have seen oiilv onie other
collection from South America, one made by Spruce at Tauau, Brazil.
Spruce makes no melition of this collectioniin his " Hepaticae of the Amazon
alid Andes. "
DEPARTMENT OF BOTANY,
UNIVERSITY OF CINCINNATI.

14 Hepaticae Indiae Orielitalis. Jour. Liluln.Soc. Bot. 5: 89-128.

15 Die Gattung Lejeunea iin Herbarium Lindenberg. Hedwvigia29: 1-23, 1890.
16 Verdoorn,op. cit. 175.

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