Organization for Flora Neotropica

Calymperaceae
Author(s): William D. Reese
Reviewed work(s):
Source: Flora Neotropica, Vol. 58, Calymperaceae (Jan. 22, 1993), pp. 1-101
Published by: New York Botanical Garden Press on behalf of Organization for Flora Neotropica
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FLORA NEOTROPICA

MONOGRAPHNUMBER58

CALYMPERACEAE

by

William D. Reese

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NEOTROPICAFP0
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Publishedfor
Organization for Flora Neotropica
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Issued 22 January 1993
 CALYMPERACEAE
WILLIAM D. REESE

TABLE OF CONTENTS
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1
Resumen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1
Introduction . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
Historyof Calymperaceae ..........................................2
Morphologyand Terminology ........................................4
Phylogeny and Evolution ........................................... 4
SystematicTreatment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. .5
Descriptionof Calymperaceae ....................... . . . .... 5
IntroductoryKey to NeotropicalCalymperaceae............................ . .5
Descriptionof Syrrhopodon . ................................. ..... 6
Key to the Species of Syrrhopodon .................................... .7
Descriptionsof Species of Syrrhopodon .................................. 11
Descriptionof Calymperes .................................... . .. .68
.
Key to the Species of Calymperes .................. ................ . 69
Descriptionsof Species of Calymperes ................. ............... . 70
ExcludedTaxa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
Acknowledgments ........................ .............. 96
LiteratureCited ............................ ....... . .. 98
Index of Scientific Names ........................................... 99
Addendumto Key B, page 10 ..................................... 101

ABSTRACT
Reese, William D. (Biology Department,Box 42451, Universityof SouthwesternLoui-
siana, Lafayette,LA 70504-2451, U.S.A.). Calymperaceae.Flora Neotrop. Monogr. 58:
1-102. 1993.-The Calymperaceae(Musci) are acrocarpousmosses, mostly dioicous and
with erect stems, predominantlycorticolous, and largely restrictedto the tropics and
subtropicsworldwide.The family is characterizedby the uniformpresencein the clasping
leaf bases of cancellinae(fields of enlarged,hyaline,exterally andinterally porose cells),
lack of stem centralstrand,mostly thickenedandtoothedleaf margins,peristomesingle (or
lacking), and commonproductionof uniseriategemmaeon the leaves. The Calymperaceae
aremost closely relatedto the Pottiaceae,fromwhich they differ in no single character.
The traditionalfamily Calymperaceaeis representedin the neotropicsby 55 specific and
subspecific taxa in two genera:Calymperes(16 species) and Syrrhopodon(39 species and
taxonomic varieties). Nine species of Calymperesand 30 species and varieties of
Syrrhopodonareendemicto theneotropics;theothertaxaalso occuroutsideof theneotropics
includingsome thatare more or less pantropicalandsome disjunctin Africa.

RESUMEN
Reese, William D. (Biology Department,Box 42451, Universityof SouthwesternLoui-
siana, Lafayette, LA 70504-2451, U.S.A.). Calymperaceae.Flora Neotrop. Monogr. 58:
1-102. 1993.-Los Calymperaceae(Musci) son musgos acrocipicos, su mayoriadioicos
y con tallos erectos, predominantemente
corticicolas,y en granparterestrictosa las zonas

of Biology,Box42451,Universityof SouthwesternLouisiana,
Department Lafayette,Louisiana70504-2451,U.S.A.

1
 ~~~~~~~~~~~~~2 ~~~Flora Neotropica

tropicalesy subtropicalesdel mundo.La familiase caracterizapor la presenciauniformede

cancelinos (grandescelulas hialinas,extera e intemamenteporosas)en la base cinientede
las hojas, ausenciade cord6ncentralen el tallo, margende hojasprincipalmenteengrosado
y dentado,peristomasimple (o ausente)y comun producci6nde yemas uniseriadasen las
hojas. Los Calymperaceaese parecenmas a los Pottiaceae,de los que no se diferencianen
una caracteristicacomin.
La familia Calymperaceaetradicionalesta representadaen el neotr6pico por 55 taxa
especificas o subespecificasen dos generos:Calymperes(16 especies) y Syrrhopodon(39
especies y variedades taxon6micas). Nueve especies de Calymperes y 30 especies y
variedadesde Syrrhopodonson end6micasdel neotr6pico;las otras taxa tambienocurren
fuera del neotr6pico, incluyendo algunas que son mis o menos pantropicalesy algunas
dislocadasen Africa.

INTRODUCTION differentiationof one from the other, imply a

commonphylogeny.
The Calymperaceaeare a large and diverse Reese et al. (1986) discussed generic concepts
in the Calymperaceae. In brief, Reese et al.
family of acrocarpousmosses of worldwideoc-
currence,primarilyin the tropicsand subtropics, (1986) consideredthe family to include only the
but with a few species ranging into temperate generatraditionallyassigned to it-Calymperes,
latitudes.Worldwidethereappearto be approxi- Mitthyridium,and Syrrhopodon-and suggest
mately 40 taxa of Calymperes,20 of Mitthyrid- that Hypodontiumbe excluded, which was for-
ium, and 90 of Syrrhopodon (Reese 1987a,b; mally doneby Reese andZander(1988) by trans-
Reese et al. 1986). Sixteenspecies of Calymperes ferringit to the Pottiaceae. Reese et al. (1986)
and 39 taxa of Syrrhopodonare recognized for also suggested that the question of most appro-
the neotropics in this treatment.Although one priate classification of genera with "leucobrya-
ceous"leaves is still unresolvedin spite of varied
species of Mitthyridiumhas long been attributed
to western South America (cf. Nowak 1980), no proposalson the matterin recentyears (see Reese
recently collected specimens exist and it seems et al. 1986 for review).The presenttreatmentin-
likely thatthe specimen(s)uponwhich therecord cludesonly thetwo traditionalgeneranotedabove.
is based were mislabeled as to origin. Although most species of Calymperaceaein-
The Calymperaceae are characterizedby a habit tropical lowlands, many-particularly in
combinationof characters,includingthe acrocar- Syrrhopodon-range into or characteristically
occur at moderateto high elevations, to above
pous and mostly corticolous habit, cancellinae
(fields of enlarged,hyalinecells withinternaland 3000 m in the case of S. gardneri in Nepal.
externalpores in the leaf bases), lack of a central However, the typical habitatfor most species in
strand in the stem, thickened and toothed leaf the family is moist, tropical-subtropicalforest at
low to moderateelevations. The majorityof the
margins, often reduced single peristome (lack-
ing in Calymperes and some Syrrhopodon), species of the family are corticolousor epixylic,
and common production of uniseriate gemmae but some grow occasionallyor characteristically
on the leaves, mostly at the tips. In the genus on rock and soil. In the neotropics members of
Calymperes, many species bear characteristic the familyoccuras faras approximately41 ?N(in
intramarginal files of differentiated cells, the easternUnited States) and 35?S (in Argentina),
teniolae, which are largely lacking in butmost taxahave muchmorerestrictedlatitudi-
Syrrhopodon. nal ranges.
The Calymperaceaeare closely related to the
Pottiaceae,and no single characterdistinguishes HISTORY OF CALYMPERACEAE
the two families. Reese and Zander(1988) com-
pared and contrastedmany features of the two Calymperes(1813) was the first genus of the
families, and also provided a key to distinguish present-dayfamily Calymperaceaeto be recog-
them. The many similarities between the nized, followed by Syrrhopodon(1824) and Thy-
Calymperaceaeand Pottiaceae, and the subtle ridium (1868-segregated from Syrrhopodon),
Calymperaceae 3

the latter name a later homonym and now re- and Syrrhopodontaceaesensu Fleischer), it has
placed by Mitthyridium. not been so used in recentyears.
The name Calymperaceaewas first used in Since 1924authorshave generallyfollowed the
print by Muller (1849), at the rank of subtribe, concept of Brotherus for the family
andincluded Calymperes,Syrrhopodon,andEn- Calymperaceae, uniting Calymperes with
calypta (the latternot consideredclosely related Syrrhopodonandits segregates,ratherthanplac-
to Calymperaceaetoday). It next appearedin ing them in separatefamilies. Althoughpropos-
printin 1873 when Jaegerand Sauerbeckused it als have been made from time to time to enlarge
as the name of their tribe Calymperaceae,still the scope of the Calymperaceaeby including
including Calymperes, Syrrhopodon,and En- genera with "leucobryaceous"leaves, there do
calypta. Meanwhile, Mitten (1869) had pub- not seem to be compelling reasons for doing so
lished the name Syrrhopodonteaefor a tribe and the questionof most appropriateclassifica-
including only Calymperesand Syrrhopodon. tion of such plantsis unresolved.See Reese et al.
Kindberg (1897: 11) was the first author to (1986) for a review of generic concepts in
designate Calymperaceae as a family name. Calymperaceae.
However, he included only the genus Following the world-wide compendium of
Calymperes in his family, and classified mosses by Miiller (1849), the next significant
Syrrhopodonin the family Weisiaceae(= Pottia- contribution to knowledge of neotropical
ceae). A few years later Muller (1901) became Calymperaceaeis the monumentalwork of Mit-
the firstauthorto use thenameat the familylevel ten (1869), who included nine species of
in its modem sense, emending its concept to Calymperesand 40 of Syrrhopodonfor the con-
includeCalymperes,Codonoblepharum(a segre- tinentof SouthAmerica.Bescherelle (1896) and
gate of Syrrhopodon), Hypodontium, and Roth (1912) reviewed the genus Calymperes
Syrrhopodon.Brotherus(1901, but later in the world-wide, giving rise to considerable confu-
year than Muller) also used the name sion by using a largelyartificialbasis for classi-
Calymperaceaeat the family level and included fication.
only Calymperesand Syrrhopodon. Morefocusedstudy of neotropicalCalympera-
In 1904 Fleischer used the name ceae beganwith Williams(1920), who published
Calymperaceaein the same sense thatit was used a revision of the species known to occur in the
by Kindberg, including only the genus North AmericanFlora area, preliminaryto for-
Calymperes.Fleischer coined the family name mal treatmentof the family for the North Amer-
Syrrhopodontaceaefor the genera Syrrhopodon icanFloraseries. However,a formaltreatmentof
and Thyridium.Brotherus(1924) again used the Calymperaceaehas never been publishedin the
name Calymperaceaein its extended sense, in- North AmericanFlora series. Williams recog-
cluding the genera Calymperes,Calymperopsis nized 12 species of Calymperes and 19 of
(a segregate of Syrrhopodon), Hypodontium, Syrrhopodonin his study, which excluded South
Syrrhopodon,and Thyridium. America.
Dixon, in 1930, recognizedthese mosses at the Bartram (1949) included five species of
ordinal level by creating the Syrrhopodontales, Calymperes and six of Syrrhopodonin his
including the single family Syrrhopodontaceae, "Mosses of Guatemala,"and Crum and Steere
which included two subfamilies-Calymperoi- (1957) treatedseven species of Calymperesand
deae and Syrrhopodontoideae.Dixon did not 11 of Syrrhopodonin theirstudyof the mosses of
mention the name Calymperaceae. Syrrhopo- PuertoRico and the Virgin Islands.
dontaceaeused in the sense of Dixon is illegiti- In 1961Reese publisheda revisionof the genus
mate because it includes the type genus Calymperesfor the neotropics,including 16 spe-
(Calymperes)of a name previouslypublishedat cies. This was followed by Florschiitz (1964),
the rank of family. Although the family name who treated 14 species of Calymperes and 11
SyrrhopodontaceaeFleisch. is available for use species and varieties of Syrrhopodonfor Sur-
by authors who might prefer to recognize iname.Florschiitzperformeda greatservice in his
Calymperesand Syrrhopodonas comprisingsep- publicationby includinga most useful "Guideto
arate families (Calymperaceaesensu Kindberg theliteratureon mossesof LatinAmerica."Finally,
4 FloraNeotropica

in 1977 and 1978, Reese publishedrevisions of slit-like perforationsof the calyptraare likewise
the neotropicaltaxa of Syrrhopodon-the elimb- closedwhenwetandopenwhendry.Inthedrystate
ate and limbate taxa, respectively- includinga of the capsule-calyptrasystem of Calymperes,
totalof 35 species andtaxonomicvarieties. sporescan escapethroughthe gapingperforations
Subsequentstudies on neotropicalCalympera- of the calyptra(cf. Edwards,1980;Reese, 1961).
ceae-e.g., Reese 1983a-have refined our Fritsch(1991) includedn = 13 for Syrrhopodon
knowledge of the family. See also Reese (1992) gardneribasedon two counts from India,n = 13
for a treatmentof the Calymperaceaeof Central for S. incompletus,on a count fromFlorida, and
America. n = 13 for S. texanuson a count from Alabama.
L. E. Anderson(pers. comm. 1968) also found
the numbern = 13 for Calymperespalisotiifrom
MORPHOLOGY AND Florida.
TERMINOLOGY
PHYLOGENY AND EVOLUTION
Other than the usually distinct division of the
leaf into an "upperlamina"and an often shoul- Since Miller's treatment (1849), the
dered "lower lamina," in general structure, Calymperaceaehave generallybeen classified in
mosses of the Calymperaceaearesimilarto many alliancewith the Pottiaceaein recognitionof the
otheracrocarpousmosses. However,severalspe- manysimilaritiesandsharedcharacteristics.This
cializationsshould be mentioned.The most con- is doubtlessthe correctview, and it is likely that
spicuousmorphologicalfeatureofcalymperaceous both families share a now vanished common
mosses is in the usually clasping leaf base, which ancestor. Reese (1987a,b) proposed that
is largelycomposedof large,empty,hyalinecells Syrrhopodonis ancestralto bothCalymperesand
forming the cancellinae. The cells of the can- Mitthyridium,and that the morphotype of
cellinae areadaptedfor short-termwaterstorage; Syrrhopodonwith leaves borderedby elongate,
they have large external and internalpores, so hyaline cells is ancestral to the forms whose
that they communicatewith one anotheras well leaves lack such borders. Reese's analyses
as with the environment.Waterheldin the cells of (1987a, 1987b) of world ranges of Calymperes
the cancellinaepresumablyextends the periodof andSyrrhopodonshow thatalthougha handfulof
metabolicactivityof the plantsafterwetting.Sim- taxaaremoreor less pantropical,the majorityof
ilar poresare knownin basalcells of othermosses species occur either in the neo- or paleotropics
including at least Encalypta and members of but not in both. Furthermore,differentsuites of
Dicranaceae, Leucophanaceae, Pottiaceae, and morphologies and species complexes have
Leucobryaceae (cf. Reese & Zander 1988), evolved independentlyin the neo- andpaleotrop-
probably indicating multiple evolutionary ori- ics, indicatinga long periodof geographicisola-
gins of the pores. In many species of the genus tion following the evolutionary origin of the
Calymperes the leaves characteristically bear precursorsof the family, which evidently took
intramarginalfiles of elongate,often thick-walled place priorto dispersalof the presentday conti-
cells. The files of cells are termedteniolae;they nentallandmasses.
presumablyfunctionin supportand,perhaps,water It seems likely that the few species of
transport. Calymperaceaethat have pantropicalranges or
The most strikingmorphologicalfeatureof the large disjunctions-i.e., South America-Af-
Calymperaceaeoccurs in the genus Calymperes rica-achieved such rangesby long-distancedis-
andnowhereelse amongmosses-the persistent, persal, although the plants of this family are
perforatedcalyptra,enclosing the entirecapsule poorly designed for travel (cf. Reese 1987a,b).
and gripping the operculum by the tip of its The problemof interpretingdisjunctionsis exac-
rostrum(Fig. 72E). Capsulesof Calympereslack erbatedby ranges such as that of Syrrhopodon
a peristomeandinsteadspore releaseis regulated incompletusvar.incompletus,which is very wide
by the calyptra,with the operculumfunctioning in the neotropics and also known from a few
as a stopper in closing the mouth of the capsule collections from eastern(but not western) tropi-
when wet and opening it when dry. The vertical cal Africa(Reese & Orban1986).
Calymperaceae 5

SYSTEMATIC TREATMENT but sometimes borne along costa at midleaf or

lower, clavate, fusiform, or filiform-uniseriate.
Dioicous (rarelymonoicous).Perichaetialleaves
Description of Calymperaceae
scarcely differentiated. Setae smooth, mostly
Calymperaceae(C. Mill.) Kindberg,Generaof elongate; capsules cylindrical, mostly exserted
but rarely immersed; stomata scanty, phan-
European and Northamerican Bryineae
(Mosses) Synoptically Disposed, eropore;peristomessingle or lacking, often ves-
11. 1897. Type: CalymperesSw. tigial, the teeth variously papillose externally,
mostly smooth internally,with weak transverse
Syrrhopodontaceae Fleischer, Die Musci der Flora von bars;opercularostrate.Calyptraenaked,smooth
Buitenzorg. 1: 193. 1904. Type: Syrrhopodon or papillose, cucullate and deciduous or rarely
Schwaegr. conical-mitrate(Syrrhopodon),or clasping the
seta below the capsule and persistent
Plants erect (rarelyprostrateandwith erect-as-
(Calymperes),the spores then escaping through
cending branches),with simple or highly forked vertical fissures in the calyptra.Spores mostly
stems in loose to dense mats or cushions;stems
sphericaland finely papillose.
lacking central strand;rhizoidsbrown to red or
Mostlytropicalto subtropical;a few temperate.
purple.Leaves sometimesdimorphic(gemmifer-
ous and nongemmiferous),straightto crispateor Epiphytic,epixylic, or less commonly terrestrial
or epilithic. Calymperes, with 16 species and
variously contorted or twisted when dry; in-
Syrrhopodon,with 39 species and varieties,
tramarginalfiles of differentiatedcells (teniolae) occurin the neotropics.
present (some Calymperes) or lacking (most Fertile specimens of Calymperes and
Syrrhopodon);costa single and strong,in section
showing medianguide cells with dorsalandven- Syrrhopodonareeasy to recognizebecauseof the
tralbandsof stereidcells, the latterpoorlydevel- difference in the calyptra.Sterile specimens of
oped in some Calymperes; basal part of leaf Calympereswhose leaves have teniolae are also
(lower lamina)mostly sheathingandwithevident easy to recognize, although leaves of
shoulders, including usually conspicuous fields Syrrhopodonsteyermarkiiand some of the vari-
(cancellinae)of enlarged,hyaline,externallyand eties of S. incompletusbearteniola-likefeatures.
internallyporose cells; upperpartof leaf (upper Sterile specimens of Syrrhopodonwhose leaves
lamina)shorterto many times longer thanlower areborderedwith elongatehyaline cells are like-
lamina,its marginsmostly variouslythickenedor wise readilyrecognized.However, sterile speci-
bordered, or both, but rarely undifferentiated, mens of Calymperesthat lack teniolae in their
mostly toothed at least in part; cells of upper leaves, and sterile specimens of elimbate
lamina isodiametricto vertically or transversely Syrrhopodon,may be difficult to place in the
elongate, smooth to bulging to mammillose to correct genus. The following key attempts to
uni- or pluripapillose;gemmae frequentlypres- guide the userto the propergenus for both fertile
ent on leaves in Calymperes, less common in and sterilespecimens.
Syrrhopodon,mostly ventralon costaat leaf apex

IntroductoryKey to NeotropicalCalymperaceae

1. Sporophytes present.
2. Calyptra cucullate, deciduous; peristome present or lacking. .............. 1. Syrrhopodon.
2. Calyptra clasping seta below capsule, persistent; peristome lacking. ........... 2. Calymperes.
1. Sporophytes lacking.
3. Margins of upper lamina with border of elongate hyaline cells (bordersometimes incomplete to
nearly lacking) ...................... .. . . 1. Syrrhopodon.
3. Margins of upper lamina lacking border of elongate hyaline marginal cells, variously bordered other-
wise or undifferentiated.
~~~~~~~~~~~~6 ~~~Flora Neotropica

4. Teniolaepresent(atleastin someleaves).
....................... Calymperes-butsee also Syrrhopodon incompletusvars.
4. Teniolaelacking.
5. Marginsof upperlaminanotdifferentiated, unistratose(occasionallyirregularly bistratose
abovein Syrrhopodon theriotii).
6. Plantssmall,oftenpurplish,to 1 cm tall;leavesmostlyless than2 mmlong,
marginscrenate. ................ .. . 16. Calymperesrubiginosum.
6. Plantsto 8 cm tall,neverpurple;leaves 15-20 mmlong,marginsirregularly toothed.
................................. 28. Syrrhopodon theriotii.
5. Marginsof upperlaminaclearlydifferentiated, bistratoseor thicker.
7. Upperlaminalong andnarrowlylinear,its cells mostlytransversely elongate;costain section
showing2 or morerowsof guidecells. ....................... 2. Calymperes.
7. Upperlaminavariouslylanceolateto ovateorbroadlylinear,cells mostlyisodiametricto verti-
cally elongate;costain sectionmostlyshowinga singlerowof guidecells.
8. Lowerlaminagolden-brown to reddish;cancellinaemostlyfragileanderoded(butmostlyin-
tactin Syrrhopodon cryptocarposandS. rigidus). .............. 1. Syrrhopodon.
8. Lowerlaminahyaline,mostlynot coloredas above(butsometimesyellowishin
Syrrhopodonincompletus); cancellinaeintactandpersistent(butsometimessomewhateroded
in S. cryptocarpos).
9. Marginsof upperlaminacoarselyserrate-toothed in rows,the teethforming3 or moreirregular
wingsfromshoulderto apex;cells of cancellinaeornamented withfainttransversebands.
................................. 30. Syrrhopodon cryptocarpos.
9. Marginsof upperlaminaentireorbearingat most2 rowsof teeth;cells of cancellinae
lackingbands.
10. Marginsof lowerlaminabearingstout,sharp,spreadingto recurvedspinoseteeth;
rhizoidsdark-red. ....................... 20. Syrrhopodon gardneri.
10. Marginsof lowerlaminaentireto serrateor dentate,butlackingspinose-recurved
teeth;rhizoidsbrownto reddish.
11. Marginsof lowerlaminaclearlyborderedby severalto manyrowsof elongate,
thick-walledcells.
12. Cells of upperlaminapapillosedorsally;marginsof upperlaminaentireor
finelyserrateabove. ................ . . . . 2. Calymperes.
12. Cells of upperlaminasmoothto slightlybulgingdorsally;marginsof
upperlaminadoubly-toothed above.
13. Borderof elongatecells 2-6 cells wide .... 10. Calymperessmithii.
13. Borderof elongatecells 8 or morecells wide.
. 21.Syrrhopodon incompletusvars.lanceolatusandperangustifolius.
11. Marginsof lowerlaminalackingwell-definedborderof elongate,thick-walled
cells.
14. Plantstiny,to 2-3 mmtall;leaf marginsuni-or bistratose;gemmaeborne
all aroundapexof costa. ............... 9. Calymperestenerum.
14. Plantscoarse,to 2-3 cm tall;leaf marginsmultistratose; gemmaeborne
only on ventralsurfaceof costatip. .... 21. Syrrhopodon incompletusvars.

1. Syrrhopodon Schwaegrichen, Sp. Muse. mostly isodiametric,rarelytransverselyor verti-

Suppl.2(1):10. 1824. Typespecies:S.gardneri cally elongate; leaf margins variously toothed,
(Hooker) Schwaegrichen(designatedby E. G. mostly thickened,borderedentirelyor in partby
Britton, Fl. Bermuda 436. 1918). The name elongate hyaline cells, or unbordered;teniolae
Syrrhopodonis derivedfrom two Greekwords andgemmareceptaclesmostly lacking;gemmif-
meaning"toclose the eye" and"tooth,"in refer- erousleaves mostlynot muchdifferentfromveg-
ence to the narrow,connivent,horizontalperi- etative leaves; capsules mostly exserted, rarely
stome teeth of some species that "close" the immersed;peristomesmostly imperfectlydevel-
mouthof the capsulewhen dry. oped, sometimes lacking; calyptraecucullate or
rarelymitrate-campanulate, deciduous.
Plants mostly tuftedor gregariouson treetrunks, Syrrhopodon occurs worldwide,mostly in the
logs, rock, and humus; stems erect or rarelyre- tropics and subtropics, but with some species
pent, simple or variously branched;leaf cells ranginginto temperateregions. A few species of
 Calymperaceae 7

Syrrhopodon are more or less pantropicalbut SubgenusMacropyxidiumBroth.

most have more restrictedranges in the neo- or Syrrhopodonlycopodioides(Brid.) C. Mull.
paleotropics,orboth.Some arenarrowendemics.
The center of distributionfor the genus in the SubgenusPorodictyonBroth.
New World is northeastern South America. SyrrhopodonhelicophyllusMitt.
Thirty-ninetaxa of Syrrhopodonareknownfrom SubgenusSyrrhopodon
the neotropics.Most of these mosses arecortico- Syrrhopodoncircinatus (Brid.) Mitt., S. elatus
lous but some grow characteristicallyon logs, Mont., S. gardneri (Hook.) Schwaegr., S.
humus, rock, and soil. Many are plants of warm hornschuchiiMart.,S. incompletusSchwaegr.,S.
humidlowland habitatsbut some grow at higher rigidus Hook. & Grev., S. rupestris Mitt., S.
elevations, to 2000 m or more. steyermarkiiRobins., S. theriotii Bartr., S.
Syrrhopodonis a large and evidently entirely xanthophyllusMitt.
naturalgroup of plants with adequatelyuniform
SubgenusCalymperopsishas been recognized
sporophytic and gametophytic morphology to at the genericlevel by some authors,e.g., Broth-
meritrecognitionas a genus. Variedevolutionary
erus(1924), Fleischer(1904), and Tixier (1967),
trends within the genus clearly deserve to be
but retainedin Syrrhopodonby, e.g., Crumand
recognized at the subgenericlevel. Anderson(1981), Florschiitz(1964), and Reese
Of the eight formallyrecognizedsubgeneraof
et al. (1986). See Reese (1978) and Reese et al.
Syrrhopodonworldwide, (see Reese 1987b for
(1986) for discussion. More recently subgenus
review), six (includingtwo monotypicendemics)
have representativesin the Americas.The Amer- Heliconema,recognized by Mitten (1869) as a
ican subgeneraand theirspecies are listed alpha- section, was raised to generic rank by Ellis and
Eddy (Ellis, 1989). Because the name
betically below. Heliconema was already in use at the generic
Subgenus Calymperopsis(C. Mull.) Broth. level for anotherplant,Ellis and Eddy replacedit
SyrrhopodoncymbifoliusC. Mill., S.flexifolius withChameleion(Eddy1990).In thepresenttreat-
Mitt., S. graminicola Williams, S. parasiticuls mentHeliconemais recognizedas a subgenuswith
(Brid.) Besch. a single species in the Americas,S. cryptocarpos.
SyrrhopodonxanthophyllusMitt., also treatedby
Subgenus Heliconema(Mitt.)Broth. Ellis andEddyin Heliconema,is hereassignedto
SyrrhopodoncryptocarposDozy & Molk.
subgenus Syrrhopodon.See discussion below
Subgenus HyalolimbataeOrbain underS. cryptocarposandS. xanthophyllus.
Syrrhopodon annotinus Reese & Griffin, S. Syrrhopodonis easy to differentiate from
brasiliensisReese, S. elongatusSull.,S. erubescens Calymperesin the case of plantswith sporophytes
Bartram,S.fimbriatusMitt.,S.gaudichaudiiMont., or with leaves borderedby elongatehyalinecells.
S. isthmiReese, S. leprieuriiMont., S. ligulatus However, sterile plants lacking borderedleaves
Mont., S. prolifer Schwaegr.,S. texanusSull., S. may be difficult to distinguish from sterile
tortilisHampe,S. simmondsiiSteere Calymperesspp. whose leaves lack teniolae.

Key to Neotropical Syrrhopodon

Upperlaminaof leaves borderedentirelyor in partby elongatehyalinecells (borderincompletein some taxaor lack-

ing or nearlyso on some leaves). KEYA
Upperlaminaof leaves lackingelongatehyalinecells on margins. KEYB

Key A. Limbate Syrrhopodon

1. Leafshouldersregularlytoothedor ciliate,at leaston someleaves,or witha regulartendencyto be toothed
or ciliate.
2. Leavesdimorphic, vegetativewithbroadupperlamina,gemmiferous withupperlaminareducedto narrow
wings alongcosta;distributionNorthAmerican,notknownsouthof NuevoLe6n,Mexico. 4. S. texanus.
8 Flora Neotropica

2. Leavesbasicallymonomorphic; distribution tropicalAmerican.

3. Cells of upperlaminasmoothdorsally.
4. Leavesmostly2-3(-3.5) mmlong,acute,ciliate-toothed only at shoulders;plantsmostly
terrestrial; distributionin Amazonia. .... ..... ... ... ....... 2. S. annotinus.
4. Leavesmostly3-4(-6) mmlong, acuteto broadlyrounded,cilia sometimesextending
nearlyto leaf tip;plantscorticolous;distribution in CentralAmericaandPuertoRico.
......................................... 11. S. erubescens.
3. Cells of upperlaminavariouslypapillosedorsally.
5. Leavesmostlyshorterthan4 mm,mostlystraight.
6. Upperlaminalingulate,aboutlengthof lowerlaminaorshorter,cells of upperlaminaeach
witha single,large,pointeddorsalpapilla;leavesoftenciliatenearlyall around.
. .... ... .... .... .... .... ... .... .... ... 13..S.S.fimbriatus.
6. Upperlaminalinearto lanceolate,1.5-2 timeslengthof lower;cells of upperlaminanearly
smoothdorsallyto uni-orpluripapillose; leavesciliate-toothed only at shoulders.
7. Leafshouldersbearingdelicatecilia .............. 8. S. simmondsii.
7. Leafshouldersbearingstoutcilia or teeth.
8. Plantsprimarilyterrestrial; costashowingenlargedcells on dorsalsurfacein
section. . . . . . . . . . . ............ 2. S. annotinus.
8. Plantsmostlyon deadwood;costalackingenlargedcells on dorsalsurfaceas
seen in section. . . . ................ . 6. S. brasiliensis.
5. Leavesmostlylongerthan4 mm,mostlyevidentlyflexed.
9. Leavesonlymoderatelyflexed;cells bulging-papillose; shouldersof at leastsome
leavesirregularly toothed ........................ 3. S. tortilis.
9. Leavesmostlystronglyflexed;cells various;shouldersstronglyto weaklyciliate.
10. Cellsdorsallyminutelypluripapillose, oreachwitha singlelarge,sometimes
branched, papilla,ventrallymammillose, pluripapillose, orwitha singlelarge
spinosepapilla;plantspaleandsoft,appearing glaucous,usuallyfreelyforked.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. S. leprieurii.
10. Cells dorsallystronglypluripapillose orbulging-pluripapillose; stemsusually
not orlittleforked.
11. Cellsmerelypluripapillose dorsally,notbulging;leaveslackingcon-
spicuousrhizoids;shoulderswithsmallciliaordenticulations; plantsoften
slenderanddelicate,appearingsilky;leaves oftenremotealongstem.
...................... 1e. S.prolifervar.cincinnatus.
11. Cellsbulgingpluripapillose; leavesoftenwith conspicuousredrhizoids
at baseandapex;shouldersmostlywithstoutcilia;plantsrobust.
12. Upperlaminamostlyabouttwice lengthof lower,straightand
erect-ascending whendry,twistedbutnot crispate.
.................. 12a.S. elongatusvar.elongatus.
12. Upperlaminaaboutas long as loweror a littlelonger,
crispate-contorted whendry. 12b.S. elongatusvar.glaziovii.
1. Leafshouldersentire(occasionalteethirregularly presentin sometaxa).
13. Upperlaminamoreor less tightlyhelicallytwistedwhendry,its cells large,smooth,andconspicuously
porose ...................................... ... 18. S. helicophyllus.
13. Upperlaminamostlynottwisted;if so, its cells small,notconspicuouslyporose,andmostlypapillose.
14. Upperlaminalingulate,aboutlengthof lowerlaminaor shorter,cells eachwitha single,large,
pointeddorsalpapilla ............................... 13. S.fimbriatus.
14. Upperlaminamostlylongerthanlower,ligulateto linearor lanceolate;cells smoothorvariously
papillosedorsally.
15. Leavesincompletelyor irregularly borderedwithelongatehyalinecells.
16. Leafapexbluntlyroundedorretuse;cellsof upperlaminapluripapillose; leavesligulate,
oftenfolded-falcateon slide;sometimesbluntlytoothedat shoulders. . . 7. S. ligulatus.
16. Leafapexacute;cells smoothto pluripapillose; leavesoblongto broadlylinearor
lanceolate,notfalcate-foldedon slide,shouldersentire(exceptsometimesmoreorless
toothedin S. annotinus).
17. Plantson soil or rock(rarelydeadtreees);apexof manyleaveswithabaxiallyre-
flexedapiculation;cells of upperlaminasmoothto uni-orpluripapillosedorsally.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. S. annotinus.
17. Plantsepiphytic(sometimesepiphyllous);leaf apexlackingreflexedapiculation;
cells of upperlaminasmoothto unipapillose.
18. Cellsof upperlaminamostly10-14 x 14-17 pGn,dorsallysmoothorsomecells
inconspicuously bluntlyunipapillose. ............ 15. S.flexifolius.
Calymperaceae 9

18. Cellsof upperlaminamostly7-10 pm diameter,dorsallysmoothto sharply

unipapillose.
19. Comaeof enlarged,oftendeltoid,leavesbearingfilamentousgemmae
alongcostaoftenpresent;cells of upperlaminasmoothto unipapillose
dorsally;marginsof upperlamina-where unbordered-essentially
entire;plantssmallto robust,to 20 mmtall. ..... 14. S. parasiticus.
19. Comaeof enlargedleavesrare;cells of upperlaminaunipapillose
dorsally;marginsof upperlamina-where unbordered-serrulate by
papillaeandcell angles;plantstiny,3-4(-5) mmtall.
............................ 17. S. cymbifolius.
15. Leavesborderedessentiallyall aroundwithelongatehyalinecells.
20. Cellsof upperlamina-at leastin part-transversely elongate;marginsof upperlamina
usuallycoarselytoothedwithlarge,oftenpairedteeth. ....... 19. S. lycopodioides.
20. Cells of upperlaminaisodiametric orverticallyelongate;marginsof upperlamina
toothedor entire,if toothed,thentheteethmostlysingle.
21. Cellsof upperlaminaessentiallysmoothdorsally(occasionallywith low
inconspicuous papillae).
22. Marginsandcostaconspicuouslyciliate-spinoseat leaf tip;cancellinae
bulgingventrallyat distalends. ................. 10. S. isthmi.
22. Marginsandcostaentireabove(marginssometimesremotelydenticulatein
S.flexifolius);cancellinaeplaneventrally.
23. Plantsmostlyepiphytic,sometimeson logs, commonlywithcomaeof
enlargedleavesbearingfilamentousgemmaealongcosta;leaves
crispate-flexuouswhendry. .............. 15. S.flexifolius.
23. Plantsmostlyterrestrial, rarelyon deadtrees,lackingcomaeof enlarged
leaves;gemmaefusiform,on leaf tips;leavesinvoluteandcurvedwhen
dry,notcrispate. ................ 2.. 2. S. annotinus.
21. Cells of upperlaminadistinctlypapillosedorsally.
24. Upperlaminatightlycrispateto helicallytwistedwhendry,foldedandfalcate
on slide,oftennotmuchlongerthanlowerlamina.
25. Upperlaminaligulate,apexbroad,bluntto retuse,lackingteeth;
hyalineborderof laminaweak,incomplete,orlackingdistally;
costaoftenendingbelowleaf tip. ............. 7. S. ligulatus.
25. Upperlaminalanceolate-acuminate, apexpointed,usuallywithcoarse
teeth;hyalineborderof laminacomplete;costapercurrent.
26. Plantssoft;stemsshort(1-2 cm),repeatedlyforked;leaf bases
usuallyloosely divergentfromstems;leaf tipsandbaseslacking
conspicuoustanglesof rhizoids. ........ 5. S. gaudichaudii.
26. Plantsfirm;stemstall (to 15 cm),simple;leaf basestightly
claspingstems;leaf tipsandbasesoftenwithconspicuous
tanglesof rhizoids. ......... 12b.S. elongatusvar.glaziovii.
24. Upperlaminastraight,variouslycurvedor flexuous,or somewhatcrispate
whendry,butnottightlycrispate,rarelyhelicallytwisted,mostlylying flat
andnotfalcate-foldedon slide,mostly2-severaltimeslongerthanlower
lamina.
27. Costanearapexvirtuallysmooth;gemmaeventralon costaat midleaf;
marginsof upperlaminatoothed;upperlaminaca. 2 timeslengthof
lower. ......................... 16. S. graminicola.
27. Costanearapexusuallypapillose-spinosedorsallyandventrally;
gemmaeventralon leaf apex;marginsof upperlaminatoothedto
entire;upperlaminato manytimeslengthof lower.
28. Leavesusuallywidelyspacedon stem,conspicuouslyflexedat
45?ormore;upperlaminaattenuate,oftenhelicallytwistedbut
straightwhendry,nototherwisecontorted;stemsoftensimple.
. . . . . . ............. 1e. S. prolifer var. cincinnatus.
28. Leavescrowdedon stems,flexedor straight;upperlamina
linearto lanceolate,rarelyattenuate,straightor variously
contortedwhendry,rarelytwisted;stemsmostlyrepeatedly
forked.
29. Cellsof upperlaminadistinct,thick-walled,bulging-papillose.
30. Leaves mostly only 2-2.3 mm long. . .6. S. brasiliensis.
30. Leavesmostly5 mmor morelong.
10 Flora Neotropica

31. Plantsrathercoarse,palegreenor darker,neither

glaucousnorin grass-likecolonies;bordersof lower
laminaverybroad,on atleastsomeleavesbearinga
few low stoutdenticulations. ....... 3. S. tortilis.
31. Plantsslender,paleyellow-green,oftenappearing
glaucous,oftenin grass-likecolonies;marginsof lower
laminaentireat shoulders.
32. Longestleavesto 5 cm or more;leavesmostly
straight,oftenproliferousat tips;costaof longest
leavesoftenshowingaccessoryguidecells in
section;notcommon;mostlyabove 1000m.
............ .ld. S.prolifervar.tenuifolius.
32. Longestleavesto ca 2 cm;leavesstraightor flexu-
ous,notcommonlyproliferous;costalackingac-
cessoryguidecells;commonandabundantfrom
sealevelto to over1000m. lb. S. prolifervar.scaber.
29. Cellsof upperlaminaobscure,flat,wallsnot conspicuously
thickened,withlow papillae.
33. Leavesusuallystronglyflexedat shoulders,upper
laminaacuminateorattenuate,taperingquicklyand
evenlyto acuteapex;cancellinaeusuallybulging
ventrallyat distalends;plantsoftendark-colored,
reddishorpurple,glossy,almostexclusivelyon rock.
............ Ic. S.prolifervar.acanthoneuros.
33. Leavesusuallystraight,upperlaminalinearto broadly
linearorlanceolate,nottaperingexceptat apex,
cancellinaeplaneventrallyat distalends;plantsmostly
green,usuallyon treesandlogs butsometimeson rock
or soil. ............ la. S.prolifervar.prolifer.

Key B. Elimbate Syrrhopodon (See also page 101)

1. Lowerlaminagolden-brownto reddish;cancellinaecommonlyfragileanderoded(butmostlyintactand
persistentin S. cryptocarposandS. rigidus).
2. Cancellinaeextendingwell up intolowerlamina,to ornearlyto shoulders,usuallyintactandpersistent.
3. Leavesnot morethanca. 4 mmlong, marginswith3 ormoreirregular rowsof teeth;cells of
cancellinaebearingfainttransversebands;capsuleimmersed;highlymodifiedgemmiferous
leaveslacking. ................................ 30. S. cryptocarpos.
3. Leavesto 10 mm or morelong, marginswithat most2 rowsof teeth;cells of cancellinaelacking
transversebands;capsuleslong-exserted; highlymodifiedgemmiferousleavescommonlypresent.
. . . ......25.
. . . . . . . . . . .... . . . . . . . S. rigidus.
2. Cancellinaeendingfarbelowshoulders,usuallyeroded.
4. Upperlaminagraduallyandnarrowlyacuminate,usuallyterminating in a narrow,spinosetip;
costain sectionshowingaccessoryguidecells.
5. Distalportionof lowerlaminawithprominentteniola-likefeatures,its marginsentire.
. ................................... . 27. S. steyermarkii.
5. Distalportionof lowerlaminalackingteniola-like features,itsmarginsfinelyandevenlyserrate.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26. S. rupestris.
rupestris.
4. Upperlaminaligulateto lanceolateor broadlyor narrowlyacuminate,apexoftenacutebutnot
narrowandspinose;costalackingaccessoryguidecells.
6. Marginsof upperlaminaentireto slightlyirregular,notcoarselytoothedwithpairedteeth;
leavesmostlyless than4 mmlong;capsulesimmersed;highlymodifiedgemmiferousleaves
lacking. .............. 29. S. xanthophyllus.
6. Marginsof upperlaminausuallycoarselytoothedwithpaired(ortriple)teeth;leaves mostly
5-6 mm long;capsuleexsertedon long seta;modifiedgemmiferousleavesoftenpresent.
7. Marginsof lowerlaminaof at leastsomeleaves-often all-more or less regularly
dentate-serrate withatleastsometeethspreadingorrecurved;costaof vegetative
leavesmostlysmoothabovedorsallyor occasionallywith 1-2(-3) low teeth;
cancellinaeveryshort,at baseof lowerlaminaonly. . ..... 24. S. hornschuchii.
7. Marginsof lowerlaminaentireor somewhatirregularly serrate,teethneitherspreading
norrecurved;costaof vegetativeleavesusuallywith2-severalprominentteethabove
dorsally;cancellinaeextending1/3-1/2 up intolowerlamina. ..... 23. S. circinatus.
Calymperaceae 11

1. Lowerlaminahyaline,mostlynotcoloredas above(sometimesyellowishin S. incompletus); cancellinaeintact

andpersistent
or,if somewhateroded(S.cryptocarpos, S.gardnen),cellsof upperlaminasharplypapillosedorsally.
8. Marginsof upperlaminaentirelyor mostlyunistratose................... 28. S. theriotii.
8. Marginsof upperlaminaconspicuouslythickenedfromshouldersto apex.
9. Marginsof lowerlaminaat shouldersbearingsharp,spreadingto recurvedspinoseteeth;leaf cells
sharplypapillosedorsally. . ....................... . 20. S. gardneri.
9. Marginsof lowerlaminaentireto serrateor dentatebutnotspinose-toothed, teeth-if present-not
spreading-recurved; leaf cells smoothorpapillosedorsally.
10. Marginsof upperlaminaserrate-toothed in rows,theteethforming3 or moreirregularwings
fromshouldersto apex;leaf cells papillosedorsally;capsuleimmersed. 30. S. cryptocarpos.
10. Marginsof upperlaminabearingat most2 rowsof teeth;leaf cells mostlysmoothdorsally
(sometimessomewhatpapillose);capsuleexsertedon longseta.
11. Marginsof upperlaminabearingpredominantly singleteeth.
. . . . ...................... 2 . 21b. S. incompletusvar.berteroanus.
11. Marginsof upperlaminabearingpredominantly pairedteeth.
12. Marginsof lowerlaminaat shoulderregionuniformlyandconspicuouslybordered
by a bandof elongate,thick-walledcells;upperlaminanarrowlylinearto linear-
lanceolate.
13. Leaveslinear-lanceolate, appressedto stemwhendry,marginsin section
showingfew orno stereids. ....... 21d. S. incompletusvar. lanceolatus.
13. Leavesnarrowlylinear,spreading-ascending or somewhatflexuouswhendry,
marginsin sectionshowingseveralto manystereids.
................... 21e. S. incompletus var.perangustifolius.
12. Marginsof lowerlaminanot or only irregularly bordered;upperlaminanarrowly
linearto broadlylanceolate.
14. Upperlaminaplanewhendry,notor littlecontorted;marginsof lower
laminaat shouldersentireornearlyso; upperandmedianleaf cells plane
ventrally(as seen in section). .................. 22. S. elatus.
14. Upperlaminainvoluteorfoldedwhendry,oftencontorted;marginsof lower
laminausuallyserrateatshoulders;upperandmediancells conspicuously
highlybulgingventrally(as seen in section).
15. Costasmoothdorsally;upperlaminaalwaystronglytoothedabove,
linearto narrowlylanceolateor subulate,its upperandmediancells
mostlyuniformlyovalto rounded-rectangular; lowerlamina
uniformlywithstronglyflaringshoulders,usuallywithintramarginal
teniola-likedownwardextensionsof borderof upperlamina;supra-
cancellinarcells smoothventrally. ... 21c. S. incompletusvar.luridus.
15. Costaslightlyto stronglypapillosedorsally(atleaston someleaves);
upperlaminastronglyorweaklytoothedabove,broadlyto narrowly
lanceolate(occasionallysubulate),its upperandmediancells square,
rounded-square, orrounded-rectangular intermixed; lowerlaminawith
orwithoutteniola-like features;supra-cancellinarcellsusuallybulging-
mammilloseattheirdistalends. .. 21a. S. incompletus var.incompletus.

la. Syrrhopodon prolifer Schwaegrichenvar. SyrrhopodoncompactulusC. Miiller, Flora 83: 332.

1897. Type.Venezuela.MoritzIn. 57 (lectotype,here
prolifer Fig. 1 designated,NY).
Syrrhopodon prolifer Schwaegrichen, Sp. muse. frond. Syrrhopodon calochlorusC. Miiller,Hedwigia39: 263.
suppl.2(2): 99. 1827. Type.Brazil.Serrados Orgaos, 1900. Type.Brazil.Rio de Janeiro:Tijuca,Ule 1669
Jan1823,Beyrichs.n. (holotype,G, n.v.;isotypes,BM, (lectotype,heredesignated,H-BR).
GOET,JE,NY). Syrrhopodon persordidusC. Miiller,Hedwigia39: 264.
Syrrhopodonparvulus Schimper exC. Muller, Syn. muse. 1900. Type.Brazil.MinasGerais:Serrade OuroPreto,
frond. 1: 544. 1849. Type. Mexico. Mirador, s.c., Ule 1383 (lectotype,heredesignated,H).
(lectotype,heredesignated,NY;isolectotype,S). Syrrhopodonassimilis Brotherus,Akad. Wiss. Wien,
Syrrhopodon longifolius Lindberg ex Aongstrom, Ofvers. Math.-Naturwiss.KI., Denkschr. 83: 280. 1926. Type.
F6rh.Kongl.SvenskaVetensk.-Akad.33(4): 7. 1876. Brazil. Sao Paulo: Raiz de Serra, Schiffner 1391 (holo-
Type. Brazil. Caldas, 18/7/1854, Lindberg s.n. (holo- type, H-BR; isotype, BM).
type, S; isotype,NY). SyrrhopodonminutusBrotherus,Akad. Wiss. Wien,
Syrrhopodon laeviusculus Brotherus, Hedwigia 34: 122. K1.,Denkschr.83: 280. 1926. Type.
Math.-Naturwiss.
1895. Type. Brazil.Goyaz:im Waldedes Corumba, Brazil.SaoPaulo:distr.Itapecirica,
Schiffner1041 (ho-
Ule 1544 (lectotype,H-BR;isolectotypes,NY, S). lotype,H-BR;isotype,BM).
12 Flora Neotropica

A
I A 2

Xd B A

\FIGS.1. B. Cs at m ro

FIGS. 1-5. Syrrhopodon. 1. S. prolifer var. prolifer. A. Leaf outlines. B. Cells at midleaf. From Vitt21446 (LAF).
2. S. prolifer var. scaber. A. Leaf outlines. B. Cells at midleaf. FromReese 12603 (LAF). 3. S. prolifer var.
acanthoneurosA. Leafoutlines.B. Cells at midleaf.FromVital10588 (LAF).4. S. prolifervar.tenuifolius.A. Leaf
outlines.B. Cells at midleaf.FromChurchillet al. 15395 (NY). 5. S. prolifervar.cincinnatus.A. Leaf outlines.B.
Cells at midleaf.FromE. & P. Hegewald9412 (LAF).Scalebars:a = 1 mm (Figs. 3A, 5A); b = 2 mm (Fig. 4A); c =
0.05 mm (all cells); d = 1 mm (Figs. 1A, 2A).
 Calymperaceae 13

Plants pale- to dark-green,in loose to dense M. 2824 (NY, U). MAGDALENA: Parque Nac. de la Sierra
Nevada de Santa Marta, Griffin, III et aL 1284 (FLAS,
clumps; rhizoids red to brownish;stems usually NY). SANTANDER: Along river Luisito, Ireland 23479
much forked, to 1-severalcm tall. Leaves to ca. (CANM, NY).
1 cm long, straightor contortedwhen dry, linear VENEZUELA. AMAZONAS: Dpt. Rio Negro, 5.3 km
to narrowly lanceolate from scarcely broader NE of Pico Phelps, Sastre-de Jesuis 396 (NY). ARAGUA:
base; margins borderedall aroundwith hyaline 11.5 km N of Pie de Cerro, Luteyn et al. 8274-A, 8276
cells, mostly entire below, variously toothed (NY). BOLIVAR: Ptari-tepui Steyermark 59879 (NY).
LIBERTADOR: Sierra Nevada Merida, Griffin, III et al.
above; cancellinaevery distinct;mediancells of 017454 (FLAS, NY). Merida: Distr. Campo Elias,
upperlamina obscure, densely pluripapilloseon Fransdn 1217 (NY). ZULIA:near San Jose de Los Altos,
both surfaces,quadrateto rectangular,mostly ca. Griffin, III 69 (FLAS, LAF).
5 x 5-12 pm; gemmae not common, on ventral TRINIDAD. ST. ANDREW: near Hollis Reservoir,
surface of leaf tip. Seta reddish,5-12 mm long; Crosby 2364 (DUKE, LAF).
SURINAME. Tafelberg, Maguire 24186M (NY).
capsule 1-2 mm long;peristomeof fragile,often ECUADOR. ZAMORA: Ortega U. 530 (LAF, Q).
imperfect, blunt or pointed, granular-papillose GALAPAGOS:Isla Santa Cruz, Gradstein & Weber M-35
teeth to ca. 190 pm long; operculum1 mm long. (COLO, LAF, U).
PERU. SAN MARTIN: Strasse Moyobamba-
Spores 9-12 pm, granular. Calyptra 1.5-3 mm
Chachapoyas km 403, Frahm et al. 595 (LAF, U).
long. BRAZIL. AMAZONAS: upper slopes of Serra
Selected illustrations. Florschiitz (1964, fig. Curicuriari,
Buck2482 (INPA,LAF, NY). BAHIA: Mun.
60); Reese (1978, fig. 1-3). de Contas, Harley 26244 (K, LAF). DISTRITO FEDERAL:
Distribution. (Fig. 6A). Nearly pantropical; Immediately W of Sobradinho, Irwin et al. 29750 (NY).
20kmN of Alto do Paraiso,Irwin eta133076 (NY).
essentially throughout the neotropics; on tree Goias:
MATOGROSSO:Aripuana, Dardanelos, margem do Rio
trunksandbases, roots,stumps,humus,in forests
Aripuana, Lisboa et al. 599 (FLAS, INPA, LAF). MINAS
to ca. 2000 m. GERAIS: Serrado NE Belo Horizonte,
Cipo, Schafer-Ver-
wimp 9950 (herb. Schafer-Verwimp, LAF). PARA: Drain-
Selected specimens examined. MEXICO.CHIAPAS: age of Rio Mapuera,Smith2951 (NY). PARANA: ca. 20 km
Cercade Jaltenangode la Paz,DelgadilloM.3548 (LAF, SW of PontaGrossa on BR 376, Vitt21446 (ALTA, LAF).
MEXU). HIDALGO:Near Apulco, Sharp 5626 (NY, Rio DEJANEIRO: [without locality] Bandeira 29 (NY).
TENN).NAYARIT: Norris& Taranto14661(HSC,LAF). RONDONIA: vic. first rapids on Rio Pacaas Novos, Reese
PUEBLA: Near Zacopoaxtla,Sharp4211 (NY, TENN). 13594 (INPA, LAF, MICH, MO, NY, US). SANTACATA-
VERACRUZ: Cerrode San Crist6bal,nearOrizaba,Sharp RINA:Ueleira-Ararangua, Reitz 1498 (S). SAOPAULO:37
5560 (MICH,TENN). km E of Brotas, Vital 4842 (FLAS, LAF, SP).
GUATEMALA.SOLOLA: Volcin Atitlin,Steyermark BOLIVIA. LA PAZ: Prov. S. Yungas, basin of Rio
47986a (F). Bopi,Kinkoff10354(NY).TARIJA:O'Connor,aboveand
NICARAGUA.MuelleReal, Rio Siguia,6 Jan 1941, W of Entre Rios, Lewis 79-480 (F, LAF). Bergwald von
La Rues.n. (NY). Santiago de Chiquitos, May 1907, Herzog s.n. (JE).
COSTA RICA. ALAJUELA: Los Angeles de San PARAGUAY. ALTO PARANA: Reserva Biologica del
Vic.Tortuguero, Itab6,Buck12409(NY).
Ram6n,Brenes17130p.p.(NY).LIM6N:
OTS Station at San
Steere CR-125 (NY). PUNTARENAS:
VitodeJava,Bowers517-f(LAF,TENN).SANJOSt:4 km
SE de Carttgo,Griffin, II& MoralesC149 (FLAS,NY). Discussion. Variety proliferis quite variable in
PANAMA. CHIRIQUI: N of Boquete, Crosby 3990 size of plants and shape and length of leaves;
(LAF,MO). COCL9: Above El Cope,Brako8473 (NY). however,thelinearleaves andsmall, obscureleaf
COL6N: end of Santa Rita Ridge Road, Crosby10365
(LAF,MO).
cells-densely papillose with low papillae-
CUBA. Lomadel Gato,Dec 1893, Hiorams.n. (BM). make it easy to tell in most cases. It grades into
JAMAICA.ST.ANDREW: 1/4 mi. SE of Morce'sGap, the var. tenuifolius. In the very common var.
Crosby2991 (DUKE,LAF). scaber, also with linear leaves, the leaf cells are
HAITI. Morne des Commissaires, Mackaness60 distinct and highly bulging-pluripapillose on
(MICH). bothsurfaces,andin varietiesacanthoneurosand
WINDWARD ISLANDS. MARTINIQUE:
Deux-Choix,
Stehle'3878(NY). cincinnatus the leaves are acuminate. Orban and
COLOMBIA. ANTIOQUIA: 20 km NE de Medellin, Reese (1990) reviewedthe S.prolifercomplex on
Cerca
Churchilletal. 13144 (HUA, LAF, NY). CAQUETA: a world-widebasis.
del Rio Pato, 2?30-33'N, 74?45'W, Churchill &
Bentancur17024 (LAF,NY). CAUCA:Mun.de Popayan,
Churchill & Franco 16569, 16584 (LAF, NY). lb. Syrrhopodon prolifer Schwaegrichenvar.
-La Primavera,"Hammen & Jaramillo-
CUNDINAMARCA: scaber (Mitt.)Reese, Comb. Nov. Basionym:
14 Flora Neotropica

TROPICAL AMERICA Flra Newopoca base map no 1

.
~---... ..,.,-w -
7. ....

FIG. -d - -.
p- -

A. S. prolifervar.prolifer.B. S. prolifervar. scaber.

FIG. 6. Distributionsof AmericanSyrrhopodon.
Calymperaceae 15

Syrrhopodonscaber Mitten,J. Linn. Soc., Bot. Illustrations. Crumand Anderson(1981, Fig.

12: 119. 1869. Type. Cuba. Wright46 (holo- 111); Reese (1978, Fig. 4-6).
type, NY; isotypes, H-BR, L, S). Fig. 2. Distribution. (Fig. 6B). Widespread and
weedy throughout tropical and subtropical
C. Miller, Bot. Zeitung(Berlin) America,from the southeasternU.S. to southern
Syrrhopodonpapillosus
6: 766. 1848. Syrrhopodonprolifer Schwaegr.var. Brazil. On tree trunks and bases, stumps, logs,
papillosus(C. Mull.)Reese,Bryologist81: 196. 1978,
nom.illeg. incl.typevar.prior.Type.Venezuela.Prov. humus,andsoil in forestsfromsea level to about
Caracas:pr. Galipan,alt. 5000 ft. in arboribus,1845, 2000m.
Funck& Schlim358 (lectotype,heredesignated,NY;
isolectotypes,BM,G, H-BR,L, S). Selected specimens examined. MEXICO. CHIAPAS:
Syrrhopodonflavescens C. Miiller,Syn. muse.frond.1: Guey-tepec prope S. Crist6bal, Munch 7390 (H, NY).
541. 1849.Type.Trinidad.MonteTocuche,Criigers.n. HILDAGO: Cerro de Tutotepec, Sharp 5226 (MICH,
(lectotype,here designated,NY; isolectotypes,BM, TENN). OAXACA:N of Niltepec, Sharp 5769 (MICH,
H-BR,JE, S). TENN). PUEBLA: Below Huauchinango, Sharp 3047 (NY,
SyrrhopodonschwaneckeanusC. Miiller,Bot. Zeitung TENN). SANLouis POTOSI:Above Xilitla, Sharp 5823
(Berlin)13: 763. 1855.Type.PuertoRico.Schwanecke (MICH, TENN). TAMAULIPAS: Rancho del Cielo, Sierra
s.n. (lectotype,heredesignated,NY; isolectotypes,H- de Guatemala,Crum 1661 (MICH). VERACRUZ: 9 km SW
BR, S). de Tlapacoyan, Delgadillo M. 3882 (LAF, MEXU).
SyrrhopodoncryptodusMitten,J. Linn. Soc., Bot. 12: GUATEMALA. ALTA VERAPAZ:Prope Coban,
119. 1869.Type.Peru.Campana,Spruce7(NY). Turckheim 7539 (H, NY). Baja Verapaz: Civija, Sharp
Syrrhopodoncalymperidianus Bescherelle,Ann.Sci.Nat. 5199 (NY, TENN).
Bot. 6, 3: 195. 1876. Type. Guadeloupe.L'Herminier BELIZE. ELCAYO:San Agustin, Mains 3929 (MICH).
s.n., "inherb.Schimp.sub S.flavescensHpe.,"(holo- HONDURAS. FRANCISCO MORAZAN: Cerro La Tigre,
type, BM;isotypes,H-BR,NY, S). Richards et al. 3530 (LAF, NY).
Syrrhopodonschwaneckeanusfo. major Bescherelle, NICARAGUA. PEARLKEY, [without data] ex "Her-
Ann.Sci. Nat.Bot.6, 3:195.1876. Type.Guadeloupe. barium de Thiimen," (NY). ZELAYA:1 km E of Cerro
L'Herminiers.n.,'in herb.Schimp."(BM). Livico, Stevens 18722 (MO).
Syrrhopodon subviridisBescherelle,Ann.Sci. Nat.Bot. COSTA RICA. ALAJUELA: Reserva Biological de
6, 3: 196. 1876. Type. Guadeloupe.Husnots.n., P1. Bosque Nuboso de Monte Verde, Crosby 10007 (LAF,
des Antilles122(holotype,BM;isotypes,H-BR,L,NY,
MO). CARTAGO: La Estrella, Standley 39377 (NY). SAN
REN,S). Jose: vie. El General, Skutch2888 (NY).
Syrrhopodonscaber var. breviligulatusC. Miiller, PANAMA. BOCASDELTORO:Cerro Colorado, Allen
Hedwigia37: 235. 1898. Type. PuertoRico. Utuado, 5274 (MO, NY). COL6N:end of Santa Rita Ridge Road,
1887, Sinteniss.n. (lectotype,heredesignated,H-BR; Crosby 10367 (LAF, MO).
isolectotype,S). CUBA. Sierra de Cante6, Ledn & Clement 5514
Syrrhopodon dussii Brotherusin Urban,Symb.Antill.3:
(MICH).
422. 1898. Type. Martinique.Mome Rouge,Duss 83 JAMAICA. ST. ANDREW: 1-2 mi. NW by N Hardwar
(holotype,H-BR;isotype,NY). Gap, Crosby 3191 (DUKE, LAF).
Syrrhopodonmacrophyllus Brotherus in Herzog, HAITI. Mt. La Salle, Mackaness 24 (MICH).
Biblioth.Bot. 87: 28. 1916. Type. Bolivia.Bergwald DOMINICAN REPUBLIC. SEIBo:Between Miches
bei TresCruces,Herzog3515 (holotype,JE;isotypes, and El Seibo, Norris et al. B6945 (NY).
FH,H-BR,L, NY, S),nonS.macrophyllus Broth.,horn. PUERTO RICO. CordilleraCentral,trail downto river
illeg. at km 14 on Hwy 120, Reese 14602 (LAF, NY).
SyrrhopodonsubmacrophyllusBrotherus in Herzog, VIRGIN ISLANDS. ST.JOHN:Bordeaux, N. L Britton
Biblioth.Bot. 87: 29. 1916. Type.Bolivia.Bergwald & Shafer 549 (NY). TORTOLA: Lafe Mt., Fishlock 82
bei TresCruces,Herzog3516 (holotype,JE;isotypes,
(NY).
H-BR,L, S). LEEWARD ISLANDS. MONTSERRAT: Pond Mt., Sha-
Syrrhopodon longifoliusLindbergin Aongstr6mvar.al- 4 Jan 1885, Suringar s.n.
ticaulis Brotherus, Akad. Wiss. Wien, Math.- fer 830 (NY). ST. EUSTATIUS:
Naturwiss.Ki., Denkschr.83: 279. 1926.Type.Brazil. (H, NY).
WINDWARD ISLANDS. GUADELOUPE: Bois de
Sao Paulo:pr.CampoGrandead"Sio PauloRailway,"
Grand-Etang,Duss s.n. (MICH). DOMINICA: Roseau Val-
Schiffner645 (holotype,H-BR;isotype,BM).
ley, Elliott 17 (BM). GRENADA: Morne au Camp, Eggers
6212 (H-BR, NY). MARTINIQUE: [without locality] Duss
16, 83 (NY). ST. LUCIA:Zwischen Castries und Dennery,
Variety scaber is commonly yellowish-green E. & P. Hegewald 9510 (herb. Hegewald, LAF).
and often grows in low grass-liketufts;its leaves COLOMBIA. ANTIOQUIA: Mun. Campamento, 6.6-
14.6 km de Campamento, Churchill et al. 15017 (HUA,
arelinearandnot or only slightly contortedwhen
LAF, NY). NORTEDE SANTANDER: Barrio Chucarina,
dry. Its median leaf cells are distinctand highly Steere 7358 (NY). VALLE:Dagua Valley, Killip 11782a
bulging-papillosedorsallyand ventrally. (NY).
16 FloraNeotropica

VENEZUELA. AMAZONAS: Dpto. Rio Negro, 5.3 km SyrrhopodoncylindrotheciusC. Miiller, Linnaea 42:
NE of Pico Phelps, Sastre-de Jesus 441 (NY). ARAGUA: 484. 1879. Type. Venezuela,Valencia,Fendlersn.
Parque Nac. Henry Pittier, Steyermark & Espinoza (lectotype,heredesignated,NY; isolectotye,S).
105806 (NY). DISPUTED AREA BETWEEN LARA AND Syrrhopodonepapillosus C. Miiller, Linnaea 42:
FALCON:Cerro Cerron, Liesner et al. 8264B (LAF, MO). 483. 1879. Type. Venezuela.Valencia,Fendlers.n.
MIRANDA: Cerros del Bachiller, near east end, Steyermark (lectotype,heredesignated,NY).
& Davidse 116695 (MO). Syrrhopodonciliolatus Geheeb& HampeinHampe,Flora
TRINIDAD. ST. GEORGE: trail up Cerrodel Aripo, 64: 338. 1881. Type. Brazil.Sao Paulo:pr. Apiahy,
Crosby 2210 (DUKE, LAF). Puiggari395 (holotype,BM;isotypes,H-BR,L,S),non
SURINAME.Wilhelminagebergte, Schultz& Wessels S.ciliolatusParis& Brotherus nec S.ciliolatusHerzog.
Boer 10246 (LAF, U). Syrrhopodon argenteusBrotherus,Acta Soc. Sci. Fenn.
FRENCH GUIANA. ST-LAURENT-DU-MARONI: Can- 19(5): 12. 1891. Type.Brazil.MinasGeris: Caraca,
tondeMaripasoula, ca.6 kmN of Saiil,200m, Buck18650 Apr 1885, Wainios.n. (holotype,H-BR;isotypes,NY,
(CAY,NY). S).
ECUADOR. ZAMORA: Ortega U. 547b (LAF, Q). GA- Syrrhopodon carassensisBrotherus, Acta Soc. Sci. Fenn.
LAPAGOS:Isla San Cristobal, Gradstein & Weber M-115 19(5): 13. 1891. Type. Brazil.MinasGeris: Caraca,
(COLO, LAF, U). Apr1885,Wainios.n.(holotype,H-BR;isotypes,NY,S).
PERU. JUNfN:Pichis trail, Yapas, Killip & Smith25572 Syrrhopodongoyazensis Brotherus, Hedwigia 34:
(NY). 122. 1895. Type. Brazil. Goyaz: Passo Tempo, Ule
BRAZIL. AMAZONAS: km 240 on Manaus-Porto Velho 1540(H-BR).
road, Lleras et al. P19598 (INPA, NY). MINASGERAIS: SyrrhopodonmacroproliferC. Miiller, Flora 83:
Vicosa, roadto Sao Miguel, Mexia5368a (NY). PARA: 332. 1897. Type.Venezuela.Cumbrede SanHilario,
Serrado Cachimbo,km 1224 on Cuiaba-Santarem road, Goebels.n. (lectotype,heredesignated,NY).
Reese 16852 (INPA, LAF, MO, NY). PARANA: Carvalho, Syrrhopodonarenarius C. Miiller, Hedwigia 39:
Dusen13014 (NY). Rio de Janeiro:Glaziou7154 (NY). 262. 1900. Type. Brazil.MinasGerais:Caraca,Ule
RONDONIA: SeringalSao Luis,just below firstrapidson 1387(lectotype,heredesignated,NY; isolectotype,H-
Rio PacaasNovos, Reese13726 (INPA,LAF,MO,NY). BR).
SANTA CATARINA: Ilha de SantaCatarina,Yano2524 (LAF, SyrrhopodonrubicundusC. Miiller, Hedwigia 39:
SP). SAOPAULO:Ilha do Cardoso, Yano459 (LAF, SP). 263. 1900. Type. Brazil. Minas Gerais: Serra de
BOLIVIA.NearAten, Williams1859 (NY). Caraca,Ule 1379 (lectotype,here designated,NY;
isolectotype,H-BR).
Discussion. The variety scaber is a weedy Syrrhopodon macrophyllusBrotherus,Rev. Bryol. 47:
5. 1920. Type. Ecuador.Confl. riv. Gualaquizacum
moss in much of its range. Its generallystraight Bomboiza,14 Oct 1910,Allionis.n. (holotype,H-BR;
leaves when dry and distinctive highly bulging- isotype,NY), horn.illeg., non S. macrophyllusBroth.
pluripapilloseleaf cells distinguishit from var. in Herzog.
subscaber Brotherus, Rev. Bryol. 47:
prolifer. Like the var.prolifer, it gradesinto var. Syrrhopodon
5. 1920. Type. Ecuador.Gualaquiza,"Rio de Re-
tenuifolius. See Orbainand Reese (1990) for a mate,"ad pontemCuchaipamba, Jul 1909,Allionis.n.
review of the S. prolifer complex. In my revision (H-BR).
of the limbate taxa of Syrrhopodon(1978), I Syrrhopodon luetzelbergiiHerzog,Arch.Bot. Sao Paulo
this taxon as var. 1:59. 1923. Type.Brazil.Bahia:S. Goncalo,Rio das
incorrectly designated
Femeas,Luetzelberg 514/F (JE).
papillosus (C. Mill.) Reese, overlooking an al- Syrrhopodon allionii Brotherusin Engler& Prantl,Nat.
readyavailablenameat the varietallevel, created Pflanzenfam.ed. 2, 10: 231. 1924, nom. nov. Type.
autonymicallywhen Muller describedS. scaber Thetypeof S. macrophyllus Brotherus.
var. breviligulatus,as pointed out to me by Wil- Syrrhopodonrupicola Brotherus,Akad. Wiss. Wien,
liam R. Buck (1991, pers. comm.). Math.-Naturwiss.Kl.,Denkschr.83:281. 1926. Type.
Brazil.SaoPaulo:Pr.Faxina,Schiffner1850(holotype,
H-BR;isotype,BM).
Ic. Syrrhopodon prolifer Schwaegrichenvar. SyrrhopodontenuipapillosusTh6riot,Mem. Soc. Cub.
acanthoneuros (C. Muller) C. Muller, Syn. Hist.Nat. "FelipePoey"13: 222. 1939. Type.Cuba.
musc. frond. 1: 542. 1849. Fig. 3. Oriente:SierraMaestre,Ekman5320 ["5820'in proto-
logue](holotype,S; isotype,NY).
Syrrhopodon acanthoneuros C. Muller, Bot. Zeitung
(Berlin)2: 727. 1844. Type. Brazil.Serrade Nativid- Plants of variety acanthoneurosare generally
ade,Dec 1839,Gardner50 (lectotype,heredesignated, similar to those of var. prolifer but differ in
BM;isolectotypes,G, GOET,JE, L, NY). having the leaves mostly strongly flexed at the
Syrrhopodon elatior Hampe, Vidensk. Meddel. Dansk
Naturhist. Foren. Kjo/benhavn, ser. 3, 6: shoulders,and in their usually darkercolor and
132. 1875. Type.Brazil.RiodeJaneiro,Glaziou7142 taperingor attenuateupperlamina.The leaf cells
(holotype,BM;isotypes,H-BR,NY, S). have low papillae,as in var.prolifer.
Calymperaceae 17

TROPICAL AMERICA Flora Neotrop.ca base map no 1

FIG 7 Distributions of American Syrrhopodon. A..........

prolifer var. acanthoneuros B. . prolifer var. tenufoius.
18 FloraNeotropica

Illustration. Reese (1978, fig. 7-8). tenuifolius (Sullivant) Reese, Bryologist 81:
Distribution. (Fig. 7A). Tropical America, 199. 1978. Fig. 4.
tropical east Africa. Central America;Cuba; Ja-
maica; Colombia; Venezuela; Suriname;Ecu- Calymperes tenuifoliumSullivant,Proc. Amer. Acad.
Arts5:280. 1861. Syrrhopodon tenuifolius(Sullivant)
ador (including the Galapagos);Brazil; Bolivia.
Mitten,J. Linn.Soc., Bot. 12: 117. 1869. Type.Cuba.
Almost exclusively on rock, but sometimes on Decayedwood, Wright45 (holotype,FH;isotypesBM,
tree trunks,logs, and soil; forests from near sea L, NY).
level to ca. 1700 m. Syrrhopodon subintegerLindbergex Aongstr6m,Ofvers.
Selectedspecimensexamined.GUATEMALA.BAJA Forh. Kongl. Svenska Vetensk.-Akad. 33:
Civija, Sharp 5203 (NY, TENN).
VERAPAZ: 7. 1876. Type. Brazil.Sao Paulo:Serrade Cubatan,
Los Angeles de San
COSTA RICA. ALAJUELA: Feb 1855,Lindbergs.n.(S).
Ram6n, Brenes 17130, (F, NY). SANJOSE:Vie. El Gen- Syrrhopodoncapillaceus Hampe, Vidensk. Meddel.
eral, Skutch 2888 (NY). Dansk Naturhist. Foren. Kj0benhavn, ser. 4, 1:
CUBA. The type of Syrrhopodon tenuipapillosus 81. 1879. Type.Brazil.Rio de Janeiro,Glaziou9248
Theriot. (holotype,BM;isotypes,NY, S).
JAMAICA. Lower Buzza, Harris 12 (NY).
COLOMBIA. MAGDALENA: The variety tenuifoliusis characterizedby its
Parque Nacional de la Si-
erra Nevada de Santa Marta,Griffin,111et al. 50235
extremelylong leaves, to ca. 5.5 cm long, often
(FLAS,LAF). with accessoryguide cells in the costa (as seen in
VENEZUELA. BARINAS: Distr. Pedraza, 8?37'N,
70?40'W, Dorr et al. 4870 (NY). BOLiVAR: section); the leaves are often proliferousat their
Cordillera
tips. The median leaf cells are mostly like those
Epicara,base of SE escarpmentof CerroPit6n,Maguire
et al. 53616 (LAF, NY). LARA:S of Las Sabenetas, of thevar.scaber,butin some collections arelike
Steyermark55396a (NY). those of the var. prolifer. In some specimens
SURINAME. In montibus Emmaketen,Daniels & leaves of both
Jonker 1026 (LAF, U). types may be present.See Orban
ECUADOR.ORIENTE: Confluent.RioQualaquiza
cum and Reese (1990) for a review of the S. prolifer
fl. Bamboiza, 10 Dec 1910, Allioni s.n. (H, NY). complex.
GALPAGOS: Isla Isabela, Sipman M-258 (COLO, LAF, U).
BRAZIL. BAHIA:ca. 15 km NW of Lenqois,Boom& Illustration. Reese (1978, fig. 9).
Mori 1080 (LAF, NY). CALDAS: Piedrar Blanca, Mosen
Distribution. (Fig. 7B). Tropical America;
91 (NY, S). DISTRITO FEDERAL: Near Cachoeira do Lago
Azul, Vital 13507 (LAF, SP). ESPiRITO
Central America, West Indies, northernSouth
SANTO:Reserva
Florestal Pedra Azul, Schafer-Verwimp 10149 (herb.America,and disjunctto southernBrazil. Trees,
Schafer-Verwimp,LAF).GoIAs:6 km SE of Cristalina, stumps,logs, soil, humusin forests,mostly above
VitalDV-6271 (LAF,SP). MATOGROSSO:Veude Noiva, 1000 m, to ca. 2300 m.
Pranceetal. 19092 (INPA,NY). MINASGERAIS: Mun.de Selected specimens examined. BELIZE. EL CAYO
Conceiqodo MatoDentro,Vital7585 (LAF,SP). PARA: DISTR.:San Agustin, Mains3976 (MICH). TOLEDO:Maya
km 774 on Cuiabai-Santarem road,Reese16019 (INPA, Mountains, Davidse & Brant 32095 (MO).
LAF, MICH, MO, NY, US). PARANA: 11 km SE of NICARAGUA. RIVAS:NW slope of Volcan Maderas,
Jaguariaiva,Vital10588(LAF,SP). PIAUI:
ParqueNacio- Stevens 6546 (MO).
nalde Sete Cidades,Vital5397 (FLAS,LAF,SP).Rio DE COSTA RICA. CARTAGO: 25 km SE of Cartago,
JANEIRO: Serra dos Orgaos, Schafer- Verwimp7441 (herb.Crosby & Crosby 6317 (LAF, MO).
Schafer-Verwimp, LAF). Rio GRANDEDOSUL:Morro d. PANAMA. PANAMA:S. slope of CerroJefe, Tyson2486
Pedras, Sehnem 285 (NY). RONDONIA: vic. first rapids on
(NY). PANAMA-COLOMBIAFRONTIER:Cerro Mali Mori
Rio Pacais Novos, Reese 13688 (INPA, LAF, MICH, & Gentry 4322 (LAF, MO).
MO, NY). SANTACATARINA: Orleaes, Reitz 2277 (NY). CUBA. Sierra Maestra, Ledn 11232 (NY).
SAo PAULO:Ilha de Sao Sebastiao, Schdfer-Verwimp7347 JAMAICA. ST. ANDREW:1-2 mi. NW by N of
(herb.Schiifer-Verwimp, LAF). HardwarGap, Crosby 3201 (DUKE, LAF).
BOLIVIA. NearAten, Williams1859 (NY). DOMINICAN REPUBLIC. SANTIAGO: Pico Diego
Ocampo, Judd 1564 (NY).
PUERTO RICO. Maricao Insular Forest, Steere 5544
Discussion. The variety acanthoneuros is usu-
(NY).
ally easy to recognize by its habitat, dark color, LEEWARDISLANDS.ST.KITTS: Craterof Mt.Mis-
flexed tapering leaves, and obscure leaf cells with ery, 1841,Breutels.n.(NY).
low papillae. See Orbainand Reese (1990) for a WINDWARD ISLANDS. DOMINICA:
St. George,
review of the S. prolifer complex. Girondel,E.&P. Hegewald9567(herb.Hegewald,LAF).
COLOMBIA. ANTIOQUIA: La Unionhacia San Miguel,
Churchill et al 15395 (HUA, LAF, NY). CAUCA:Mun.
ld. Syrrhopodon prolifer Schwaegrichen var. de Piendamo,Churchill& Franco 16543 (LAF, NY).
Calymperaceae 19

NORTEDE SANTANDER: Barrio Chucarima, Steere 7378, oftenpresentandtheplantsaregenerallysoft and

7404 (NY). VALLE:Mun. de Call NO de CalL Churchill
etal. 15161 (NY). silky in appearance.See OrbanandReese (1990)
VENEZUELA. BOLIVAR: Base of Mt. Roraima, SW for a review of the S. prolifercomplex.
side, Buck& Brewer15675 (NY). MtRIDA:Above La Illustration. Reese (1978, fig. 10-11).
Azulita, Steyermark56144 (NY). NUEVA ESPARTA: Isla de Distribution. (Fig. 9A). Endemic to tropical
Margarita, Steyermark et aL 131080(FLAS,LAF). CentralAmerica;West Indies; north-
TRINIDAD. ST. GEORGE: S slopes of Cerro del Aripo, America;
Crosby2216 (DUKE,LAF).
ern South Americaanddisjunctto southernBra-
ECUADOR. CARCHI: SW slope of Volcan de Chiles, zil. Rotted wood, trees, rock, in forests at
Steere 9012 (LAF, NY). NAPO-PASTAZA: Sibunday, Steere moderateelevations,to 1400 m.
8630 (NY).
BRAZIL. MINAS GERAIS: Mun.Capara6Novo, Vital Selected specimens examined. BELIZE. TOLEDO:
& Buck 11513 (NY, SP). Rio DEJANEIRO: Corcovado, RichardsonCreek, Bladen Branch,Davidse & Brant
Mosen 194 (NY, S). Rio GRANDE DOSUL:Mun. Cambara 31891
(MO).
do Sul, Vital&Buck12251(NY, SP).SAoPAULO: Ilhado NICARAGUA. GRANADA: Upper slopes of Volcan
Cardoso,Vital10534 (LAF,SP). Mambacho,Croat39148 (MO).
COSTARICA.CARTAGO: I.C.E.TapantiProjectArea,
Discussion. Plants of var. tenuifolius are sim- Crosby& Crosby6076 (MO).
ilar in most respects to those of vars. prolifer and PANAMA. CHIRIQUI:vie. Fortuna Dam, Allen 5480
(LAF,MO).
scaber; it is only the greatly elongate upper CUBA. CobreRangeof SierraMaestra,Leonet al.
leaves and the accessory guide cells that distin- 10248(NY).
guish them. The leaves of this moss-to 5.5 JAMAICA.NewHavenGap,Nichols124c(MICH,S).
cm-rank among the longest known for any moss PUERTO RICO. Sierra de Naguabo, Barria de
in the world. Leaves of Syrrhopodon loreus Maizales,N. L Britton& Hess 2269 (NY).
WINDWARD ISLANDS. DOMINICA: St. George,
(Sande Lac.) Reese, of Asia and Oceania, may be zwischen FreshwaterLake und Boeri Lake, E. &. P.
up to to 6 cm long (Reese et al. 1986), which may Hegewald 9412 (herb. Hegewald, LAF). GUADELOUPE:
be the record. See Orbainand Reese (1990) for a [without locality] Duss 316a (NY). MARTINIQUE: Crags
review of the S. prolifer complex. of CarbetnearDeuxChoux,Stehle3599 (MICH).
Mun. de Guatap6, Chur-
COLOMBIA. ANTIOQUIA:
chill 16413(LAF,NY).
VENEZUELA. AMAZONAS:Cano Manteco, Bermtdez
le. Syrrhopodon prolifer Schwaegrichen var. PA-00477B (herb. Bermfidez,LAF). BOLIVAR: Distr.
cincinnatus (Hampe) Reese, Bryologist 81: Steyermarket al. 132043
Piar, Camarcaibarai-Tepui,
200. 1978. Fig. 5. (NY). NUEVA ESPARTA: La Sierra, Bermidez NN-0007
(herb.Bermuidez,
LAF).
TRINIDAD.[Withoutfurtherdata]Criigers.n.(BM).
SyrrhopodoncincinnatusHampe, Vidensk. Meddel. GUYANA. UpperMazaruniRiver region, Boom &
Dansk Naturhist. Foren. Kjobenhavn, ser. 3, 6:
131. 1875. Type. Brazil. In vie. Rio de Janeiro, Gopaul7445p.p.(NY).
PERU. SAN MARTIN: Strasse Chachapoyas-
Glaziou7136 (holotype,BM;isotypes,H-BR,NY, S). km387, Frahmet al 1874 (LAF,U).
Moyobamba
Syrrhopodon husnotiiBescherelle,Ann.Sci. Nat.Bot.6,
3: 195. 1876. Type. Guadeloupe.L'Herminiers.n. slopes and summit of Serra
BRAZIL. AMAZONAS:
Curicuriari,Buck 2465 p.p. (INPA, LAF, NY). SAo
(lectotype,BM). Martinique.MontagnePelee, sur le PAULO:
rocherset les arbres,alt. 700-850 m, Husnots.n., P1. propeCampoGrandead 'Sao Paulo Railway,"
des Antilles123 (isosyntypes,BM, H-BR,NY, S). Schiffners.n.,12.VII.1901(H-BR).
SyrrhopodonleptophyllusBrotherus,Akad.Wiss. Wein,
Math.-Naturwiss. Kl.,Denkschr.83:280. 1926. Type. Discussion. The var. cincinnatus is usually
Brazil. Parana:9 km ad orienturbisCurityba,1900, easy to recognize by the characteristicsnoted
Lalouettes.n. (lectotype,H-BR;isolectotype,S). Bra- above. It is not a common taxon anywherein its
zil. Sao Paulo:PropeCampoGrandead 'Sao Paulo
Railway"in silvulis campestribusad truncos,ca. 700 range.See Orbainand Reese (1990) for a review
m.s.m., Schiffner568 (syntype, H-BR). of the S. prolifer complex.

The variety cincinnatus is distinguished by its 2. Syrrhopodon annotinus Reese & Griffin,
leaves flexed, often ratherwell separated along the Bryologist79:518. 1976[1977]. Type. Brazil.
commonly simple stem, the upper lamina attenuate Amazonas:Rio Lages, km 130 along Manaus-
and often helically twisted, and shoulders often Caracarairoad,Griffin,III et al. 723 (holotype,
toothed. Distinctive purple to violet rhizoids are INPA;isotypes, FLAS, LAF, NY). Fig. 8.
20 Flora Neotropica

? Sn7 C
,'CO

^-^ '.*'*' 0o\

B f7O

a ________

oVf"/V~Cc~/~:: C
aO
0.. <0 C
D

FIG. 8. Syrrhopodon annotinus. A. Leaf outlines. B. Leaf margin at shoulders, entire. C. Leaf margin at shoulders,
ciliate. D. Portions of leaf sections. A from Griffin, III et al. 731 (LAF); B from Griffin, III et al. 723 (LAF); C from
Berg etal. P19521 (LAF); D from Buck 1872 (LAF). Scale bars: a = 1 mm (A); b = 1 mm (B); c = 0.1 mm (C-D).

Plants small, glossy; rhizoidspurple;stems to Distribution. (Fig. 9B). Endemic to

1 cm tall. Leaves lanceolate above scarcely Amazonia; Venezuela and Brazil. Mostly on rock
broaderbase, mostly 2-3 mm long, involute and and soil, rarely on dead wood, at low elevations.
incurved when dry; margins of upper lamina
Selectedspecimensexamined.VENEZUELA.AMA-
entire,borderedall aroundwith elongatehyaline ZONAS:CafnoManteco, Bermidez P1-00677a (herb.
cells; marginsof lower laminaentire to ciliate at Bermndez,LAF).
shoulders;medianleaf cells isodiametric,mostly estrada Manaus-Boa Vista, km
BRAZIL. AMAZONAS:
ca. 10-12 m, smooth to bulging-papillose dor- 45, Buck 2130 (INPA, LAF, NY). PARA: Serra do
Cachimbo,at Base A6reado Cachimbo,Reese 16200
sally and ventrally;gemmae infrequent,on ven- (INPA,LAF,MICH,MO,NY, US). RORAIMA: alongBR
tral surface at leaf tips. Sporophytes nearly 174,6 kmS of the Equator,Buck1873 (INPA,LAF,NY).
immersed;seta 2-2.5 mm long; capsule 1 mm
long; peristome lacking; operculum 0.75 mm Discussion. The leaves of S. annotinus are
long. Spores 33-38 pm, finely and bluntlypapil- quite variable in that cilia may be present or
lose. Calyptra mitrate-campanulate, 1.1 mm absent at the shoulders, and the leaf cells range
long, papillose above. from smooth to bulging-papillose. Nonetheless,
Illustrations. Reese and Griffin (1976, figs. this moss is easy to recognize by its low stature
1-10); Reese (1981, figs. 16-19). and soil habitat. Its mitrate-campanulate calyptra
is very unusual in Syrrhopodon.
Calymperaceae 21

TROPICALAMERICA
AMERICA -- -_
__TROPICAL-____ Neotropoca
basemapno
F__ 1
A ------- - . ........ kxalo

--- .... ......

------?--
--------------

j _ j

FIG. 9. Distributions of American Syrrhopodon. A. S. prolifer var. cincinnatus. B. S. annotinus (lines in South
America); texanus (lines
America); S. texanus dots in North
(linesanddots NonthAmerica); brasiliensis (asterisks).
America); S. brasiliensis (asterisks).
22 Flora Neotropica

AI I

10

a
b

X_

/X c,
H W<11 0

DL1aoo
0000

IC); c = 0.
(Fig. I11C); 0.55 n-m
mm (Fig. I10B).
OB).
Calymperaceae23

3. Syrrhopodon tortilis Hampe, Vidensk. Ilhado Cardoso,

Lowy1391BR(LAF,LSU).SAOPAULO:
Meddel. Dansk Naturhist.Foren.Kjobenhavn, Vital9729 (LAF,SP).
ser. 3, 4: 38. 1872. Type. Brazil. Rio de Ja- Discussion. This species was treatedunderthe
neiro: Glaziou 5188 (holotype, BM; isotypes, name S. ulei C. Mull. by me in 1978; I later
realizedthatI hadincorrectlyplaced S. tortilis-
H-BR, S). Fig. 10. the oldest name for this species-in the synon-
Syrrhopodonulei C. Muller, Hedwigia 39: 265. ymy of S. gaudichaudii. The purple rhizoids,
1900. Type.Brazil.Sa. Catarina:PedrasGrandes,Ule toothed leaf shoulders, large, bulging-
1135 (lectotype,here designated,NY; isolectotypes, pluripapilloseleaf cells, and heavily thickened
H-BR,S). leaf marginscombine to make this species dis-
SyrrhopodonrosulatusC. Miller, Hedwigia39: 264. tinct. In general,it is like a coarse version of S.
1900. Type. Brazil.Rio de Janeiro:Tijuca,Ule 1666
(lectotype,heredesignated,NY;isolectotype,H-BR). prolifervar.scaber.
Syrrhopodonulei C. Miller var. percrispaC. Muller,
Hedwigia39: 265. 1900. Type.Brazil.RiodeJaneiro:
GaviaIn, Ule 1667(lectotype,heredesignated,H-BR). 4.
Syrrhopodon texanus Sullivant,Musci U.S.
SyrrhopodondensifoliusHerzog,Beih.Bot.Centralbl. 26:
103. 1856. Type.U.S.A. Texas: 1845, Wright
60. 1909. Type. Bolivia. Bei "El Carmen,"
(Chiquitos),May 1907, Herzog s.n. (holotype,JE; s.n.(FH). Fig. 11.
isotypes,L, NY).
Plants dull, sordid,cespitose, often appearing
bristly due to stiffly erect gemmiferous leaves;
Plants coarse, pale-green to darker,in dense,
sometimes wiry, clumps; rhizoidspurple;stems rhizoidsscanty, reddish-brown;stems to ca. 10
to 1-2 cm tall. Leaves4-5(-7) mm long, crowded mm tall. Leaves strongly dimorphic,vegetative
ones linear-lanceolatefrom broader base, 3-5
at stem tips, linear-lanceolate,more or less erect
and little contortedwhen dry but sometimes he- mm long, contorted when dry, gemmiferous
lically coiled; marginsheavily borderedwith hy-leaves very narrow,erectwet and dry,marginsof
aline cells, spinose-toothed above, at least onupperlamina entire to irregularlytoothed;mar-
gins of lower laminacoarsely toothedespecially
some leaves stoutly denticulate-toothedat shoul-
ders; cancellinae distinct;cells of upperlaminaatshoulders,teethoftenspreading-recurved; can-
obscure, 5-9 x 9-14 pm, bulging and pluripa- cellinae mostly rounded distally; median leaf
pillose dorsally and ventrally; gemmae infre- cells quadrate, obscure, ca. 5-10 pm,
quently produced, borne on ventral surface of pluripapillosedorsally and ventrally; gemmae
costa at leaf tips. Sporophytesunknown. seasonally abundant,on tips of leaves. Seta 15
Illustration. Reese (1978, figs. 44-46). mm long; capsule 2.2-5 mm long; peristome
Distribution. (Fig. 12A). Mexico, Central teeth ca. 120 pm tall, blunt,segmented;opercu-
America,Jamaica,South Americaincludingthe lum 1-1.3 mm long. Spores 12-15 pm, granular.
Galapagos.Shaded,moist, rockandsoil at eleva- Calyptrasmooth.
tions to ca. 1700 m. Selected illustrations. Crum and Anderson
(1981, fig. 110 A-G); Reese (1978, figs. 47-50).
Selected specimens examined. MEXICO.CHIAPAS: Distribution. (Fig. 9B). Endemicto the south-
above MapastepecnearPaval,Xolocotzi& Sharp4419 eastern part of the North American continent;
(CANM,TENN). easternU.S. andnorthernMexico.
GUATEMALA. BAJAVERAPAZ:
Sharp 5212a (LAF,
Sharp5281 (LAF,NY, TENN).
NY, TENN);ELQUICHE: Specimen examined. MEXICO. NUEVO LEON:Mon-
BELIZE.ELCAYO:Mains3929 (MICH). Jun 1908, Pringle s.n. (S).
Cerrode la Muerte,Croat
COSTA RICA. CARTAGO: terrey,
815a (MO).
Cerro Azul, Castillo 112B (MO).
PANAMA. PANAMA: Discussion. This is the only member of the
COLOMBIA. META:Cordillera Oriental, S of Calymperaceaeendemic to continental North
Villavicencio,Steere7681b(LAF,NY). America.It is easy to recognize by its strongly
VENEZUELA. ARAGUA:Dall'Aglis 448 (herb.
Dall'Aglis,LAF).
dimorphicleavesborderedall aroundwithhyaline
ECUADOR. GALAPAGOS:Isla Santa Cruz, Itow s.n. cells andstoutlytoothedattheshoulders.Thismoss
(COLO). has not been recollectedin Mexico since Pringle
BRAZIL. GUANABARA:
Parque Itatiaia, Veu da Noiva, foundit thereandmayno longerexist in Mexico.
24 Flora Neotropica

TROPICAL AMERICA FloraNeotopca base map no 1

FIG. 12. Distributions

of American Syrrhopodon. A. . trtl. B. S. gaudchaud.

FIG. 12. Distributions of American Syrrhopodon. A. S. tortilis. B. S. gaudichaudii.

Calymperaceae 25

5. Syrrhopodon gaudichaudii Montagne, Ann. Selected illustrations. Crum and Anderson

Sci. Nat. Bot. 2, 2: 376. 1834. Type. Brazil. (1981, fig. 112);Florschiitz(1964, fig. 59); Mag-
Gaudichauds. n. (PC?, n.v.; isotypes BM, G, L, ill (1981, fig. 46: 13-21, as S. uncinifolius
NY). Fig. 13. C.Miill.; fig. 46: 1-12, as S. obliquirostris
C.Miill.);Reese (1978, figs. 12-16).
Syrrhopodon undulatusC. Miiller,Bot. Zeitung(Berlin) Distribution. (Fig. 12B). Throughoutthe neo-
2: 727. 1844. Type. Brazil. Gardner45 (lectotype, southeasternU.S. (Florida) to northern
heredesignated,BM),nonS.undulatus(Dozy& Molk) tropics;
Lindb. Argentina; Hawaii; Africa. Tree trunks,rotting
Syrrhopodoninflexus Mitten, J. Linn. Soc., Bot. 12: wood, sometimes on humus, soil, or rock, from
117. 1869. Type.Colombia.AndesBogotenses,Weir sea level to ca. 2500 m.
152 (holotype,NY; isotypes,BM, H-BR).
Syrrhopodon sartoriiC. Miiller,Linnaea38: 633. 1874. Selected specimens examined. MEXICO. CHIAPAS:
Type.Mexico.Mirador,Sartoriuss.n. (lectotype,here road from San Crist6bal las Casas to Tenejapa, Sharp
designated,NY-mostlyS. gardneri). M59124e (LAF, TENN). OAXACA:25 mi above Valle
S. spininervisLindbergex Aongstr6m,Ofvers. F6rh. Nacional in Sierra Juarez, Sharp et al. 3067-c (LAF,
Kongl. Svenska Vetensk-Akad.33: 7. 1876. Type. TENN).
Brazil.Caldas,PedraBranoa,Lindbergs.n. (holotype, GUATEMALA. BAJAVERAPAZ: Civija, Sharp 5212b
S; isotype,NY). (NY, TENN). SOLOLA:Volcan Atitlan, Steyermark
Syrrhopodon argentinicusLorentzex C. Miiller,Linnaea 47400B(F).
42: 356. 1879. Type. Argentina.Argentiniasubtrop., Richardson Creek, Bladen Branch,
BELIZE. TOLEDO:
Rio Seccopr.Orin, Lorentzs.n. (lectotype,heredesig- Davidse& Brant31932 (MO).
nated,NY; isolectotypes,H-BR,REN,S). COSTARICA.HEREDIA: SWslopesof VolcanBarba,
Syrrhopodonflexiareolatus C. Miiller,Linnaea42: 484. La Fuenta,Alfaro 40c
Crosby9713 (LAF, MO). PERALTA:
1879. Type.Venezuela.Valencia,Fendlers.n. (ecto- (MICH,NY). SANJOSE:5-7 miles from San Rafaelde
type,heredesignated,NY; isolectotypes,H-BR,S). Herediaon VolcanBarba,Bowers233-a (LAF,TENN).
Syrrhopodongracilescens Brotherus,ActaSoc.Sci.Fenn. PANAMA.COLON: end of SantaRita Road, Crosby
19(5): 12. 1891. Type. Brazil.MinasGerais:Lafay- 10472 (LAF, MO). CHIRIQUi: Las Cumbres, near town of
ette,1885,Wainios.n.(holotype,H-BR;isotypes,NY,S). Cerro Punta,Croat & Porter 16178 (LAF, MO). DARItN:
Syrrhopodonterebellatulus C. Miiller,Hedwigia39: 264. PirreMassif,just W of Cana,1200-1350 m, Allen 8901
1900. Type. Brazil. Rio de Janeiro:Tijuca, Ule s.n. Cerro Azul, Arrocha 128 (NY, PMA).
(MO). PANAMA:
(lectotype,heredesignated,NY;isolectotype,S). BAHAMA ARCHIPELAGO. ANDROS ISLAND:
Near
Syrrhopodon ochroleucusHerzog,Biblioth.Bot. 87: 29. Nicholl'sTown,Northrop344 (NY).
1916. Type. Bolivia. Im unterenCoranital,Herzog CUBA. PINAR DELRio: Rio Guao,N. L. Brittonet al.
4673 (holotype,JE;isotype,L). 10125(MICH,NY).
SyrrhopodoncrispulusSehnem, Pesquisas,Bot. 29: 7. JAMAICA. ST. ANDREW:
below Morce's Gap, Crosby
1972. Type. Brazil.SantaCatarina:SantaCatarinaIs- 3543 (DUKE,LAF).
land, Sehnem3189 (isotype,NY), hom. illeg. non S. Massif de la Hote, 44 km S of
HAITI. GRAND'ANSE:
crispulusTaylorin Mitten. Roseaux,Buck9150 (NY).
La Nevera, 47
DOMINICAN REPUBLIC. PERAVIA:
Plants in loose to dense clumps, much- km S of Constanza,Reese15686 (LAF,NY).
PUERTORICO.Caribbean NationalForest,Luquillo
branched, dark-green, with lower laminae of Div., trailfromHwy. 186 to top of El Toro,Reese14411
leaves pale and conspicuous; rhizoids purple; (LAF,NY).
stems to 1-2 cm tall but often much shorter. WINDWARD ISLANDS. GUADELOUPE:
Morne
Leaves 3-4 mm long, ligulate-acuminate, loosely i'Echelle,Duss210 (H, NY).
Mun. Medellin, 5 km be-
COLOMBIA. ANTIOQUIA:
to tightly crispate or coiled when dry, character-
yondAltode SantaElena,Luteyn& Callejas10052(LAF,
istically folded and falcate on the slide; margins Cerca de Los Andes, arribade Rio Pato,
NY). CAQUETA:
entire except at leaf tip, rarely toothed at shoul- 2?30-45'N, 74?10'W, Churchill& Bentancur16906
ders; cancellinae distinct, conspicuous when dry; (LAF, NY). CAUCA:Mun. de Popayan, Churchill &
Franco 16569 p.p. (HUA, LAF, NY). CUNDINAMARCA:
median cells obscure, ca. 7-10 pm, bulging-papil-
MundoNuevo,NE of Bogota,Steere7830 (LAF, NY).
lose dorsally and ventrally; gemmnnae infrequently NARINO:Pasto, 1?10'N, 77?07'W, Churchill& Rengifo M.
produced, ventral on leaf tips. Seta 4-5 mm long, 17501 (LAF, NY). VALLE:W slope of Cordillera Central,
reddish; capsule 1.5-2 mm long; peristome teeth Steere 7872 (LAF, NY).
VENEZUELA. AMAZONAS: Cerro Duida, Buck &
yellow to dark-red, segmented, ca. 100 pm tall; Brewer 15579 (NY). BOLIVAR:Mun. Urdaneta, Sastre-de
operculum to 1 mm long. Spores 16 pm, granu- Jestus 591 (NY). LARA: Sierrade Portugesa,Griffin&
lar. Calyptra 2-2.5 mm long, slightly roughened L6pez F. 393 (FLAS, LAF).' MONAGAS: E of Santa In6s,
above. Pursell 8981 (LAF, PAC). MtRIDA: Distr. Campo Elias,
26 Flora Neotropica

1:3S i 4

A,A

B i?8 -1'
0lfc
in/)C CN i )|;

0.05mm d mm(Fig.14A).
(Figs.14B-D); I

FIGS. 13-14. Syrrhopodon. 13.S. gaudichaudii. A. Outlines of four leaves. B. Leaf apex. From Haumann s.n. (JE).
14. S. brasiliensis. A. Leaf outlines. B. Margins and cells at leaf shoulders. C. Margin and cells at midleaf. D. Section
at midleaf. A from Vital 10328 & 10329 (LAF) (four leaves at right-two above, two below), Uk 1388 (NY) (three
leaves at bottom center). B-D from Vital 10328 (LAF). Scale bars: a = 0.1 mm (Fig. 13B); b = 1 mm (Fig. 13A); c =
0.05 mm (Figs. 14B-D); d = 1 mm (Fig. 14A).
Calymperaceae27

Griffin,1l et al. 019254 (FLAS,LAF).TACHIRA: 18 km tapering above somewhat broader shoulders,

SE of SantaAna, Steyermark& Liesner119001 (LAF, rounded to
MO). TRUJILLO: Distr. Bocono, Griffin,II et al. 1093 nearly spatulate at tips, 2-2.3 mm
(FLAS,LAF). ZULIA: near SanJose de Los Altos,Griffin, long, folded or channeled and variously con-
11192 (FLAS, LAF). tortedwhen dry,marginsof upperlaminaentire,
SURINAME. Voltzberg, Mennega 56 (LAF, U). borderedall aroundwith hyaline cells, margins
ECUADOR. GALAPAGOS ISLANDS:Duncan Island, of lower lamina entire or serrate; cancellinae
Stewart3164 (NY); Isla Isabela,Gradstein& Sipman
M-258(LAF,U). LOJA: 12kmS of Yangana,Luteyn8045 often erodedin older leaves, broadlyroundedto
(NY). NAPO-PASTAZA: Cordillera Central, E of El Pun, acute distally; median leaf cells rounded-quad-
Steere 8917 (LAF, NY). ORIENTE:Cordillera Central, rateto verticallyelongate,ca. 10-14 x 12-17 pm,
regionof El Topo,Steere9333 (LAF,NY). bulging-pluripapillose dorsally and ventrally;
BRAZIL.AMAZONAS: AlongRioNegrojustbelowRio
Uaupes, Buck 2605 (INPA, LAF, NY). BAHIA:15 km NW
gemmae clavate, slender, small, ventral on leaf
of Lencois, Boom & Mori 1089 (LAF, NY). DISTRITO tips. Seta 2.3 mm long; capsule 1 mm long;
FEDERAL:Irwin et al. 33230 (NY). ESPIRITO SANTO:Var- peristomeof low, imperfect,segmentedteeth,ca.
gem Alta, Schafer-Verwimp 10366 (herb. Schafer-Ver- 96 pm tall; operculumca. 1 mm long. Spores
wimp, LAF). GoIAs: Alto Paraiso, Schdfer-Verwimp9893
(herb. Schifer-Verwimp, LAF). MARANHAO:Mun. Barra
granular,12-14 pm. Calyptra1 mm long, papil-
do Corda, Schatz et al. 840B (NY). MATOGROSSO: Serra lose on rostrum.
do Cachimbo, 763 km N of Cuiaba on BR 163, Reese Illustration. Reese (1983b, figs. 1-11).
16050 (INPA, LAF, NY). MINASGERAIS:PoCos de Distribution. (Fig. 9B). Endemic to southern
Caldas, Schafer-Verwimp 7061 (herb. Schiifer-Verwimp, Brazil.
LAF). PARA:Serra do Cachimbo, Base Aerea do
Cachimbo,Reese16337(INPA,LAF,MO,NY).PARANA:
6 km ESE of Senges, Vital10710 (LAF, SP). Rio DE Specimens examined. The type specimens.
Serrados Orgaos,Schafer-Verwimp
JANEIRO: 7446(herb.
Schafer-Verwimp, LAF). Rio GRANDE
DOSUL:Colonia Discussion. This little moss is similar in a
Sto. Angelo, Lindman175 (H, S). SANTA
CATARINA:19 generalway to S. gaudichaudiibut the plantsare
km S of Lajes,Vital5658 (FLAS,LAF,SP). SAoPAULO: much
Ilhado Cardoso,Vital9727 (LAF,SP). smaller, have larger leaf cells, and less
BOLIVIA. Im Bergwald bei Tres Cruces, Herzog3563 for the leaves to
tendency be folded and falcate
(NY, S). NearAten, Williams1856 (NY). on the microscopeslide. The blunt,roundedleaf
PARAGUAY. ALTO PARANA: 31 km N of tip also differs from the characteristicallymore
Buck12347 (NY).CANENDIYi:
Hemandareis, 2.5 kmE of
Rio Corrientes, Buck 12452 (NY). PARAGUARI:
pointedone of S. gaudichaudii.
Parque
Nac. Ybycui,Buck12049 (NY).
ARGENTINA. TUCUMAN:
Valle de los Sosas, Steere 7. Syrrhopodon ligulatus Montagne,Syll. gen.
60-296 (NY). sp. crypt. 47. 1856. Type. French Guiana.
Leprieur1384 (holotype,PC; isotypes BM, K,
Discussion. Syrrhopodongaudichaudii is quite NY). Fig. 16.
variable and may intergrade with the S. prolifer
complex, from which typical specimens are dis- Syrrhopodoncrispus Austin, Bot. Gaz. 2:109. 1877, horn.
tinct by their crispate leaves with upper lamina illeg., non S. crispus Dozy & Molkenboer. Calymperes
crispum (Austin) Austin, Bot. Gaz. 4:151. 1879. Type.
short and folded-falcate when moist on the slide. U.S.A. Florida: 1877, Smith s.n. (NY).
See comments under S. ligulatus, a somewhat Syrrhopodonparaguensis Bescherelle, J. Bot. (Morot) 5:
similar species with leaves crisped when dry and 349. 1891. Type. Paraguay. Balansa 3673 (holotype,
with conspicuously pale leaf bases. BM; isotypes, G, S).
Syrrhopodonanomalus Brotherus, Bih. Kongl. Svenska
Vetensk. Akad. Handl. 21 Afd. 3, 3: 19. 1895. Type.
6. Syrrhopodon brasiliensis Reese, Bryologist Brazil. Minas Gerais: in Serra de Caldas, Mosen 403
86: 354. 1983. Type. Brazil. Minas Gerais: Im (holotype, H-BR; isotypes, FH, S).
Walde bei Caraqa, Ule 1388 (holotype, HBG;
isotypes, H, NY). Paratypes. Brazil. Sio Paulo: Plants very small, in compact sods, pale-green,
Ilha do Cardoso, Vital 10328, 10349 (LAF, with conspicuous leaf bases; rhizoids dark-red;
SP). Fig. 14. stems mostly less than1 cm tall. Leavesca. 2 mm
long, tightly crispedwhen dry, often folded-fal-
Plants gregarious, very small; rhizoids red-pur- cate on the slide, upperlaminalingulate,rounded
ple; stems to 5 mm tall. Leaves linear or slightly to retuse at apex; marginscrenateby projecting
28 FloraNeotropica

TROPICAL AMERICA Fkora Nvpc base map no

.'

-----

~ ~~~~~~~~~~~~~~~'

. ..

-uy. _ ----
FIG. o A c o .A.S. a.B..sn ..
1..--\.5.Dist
o ~ ~~~~~~~~~
?

6- ~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ -

...

FIG. 15. Distributions of American Syrrhopodon. A. S. ligulatus B S. immonds (line) S(asterisk)....

isthmi

FIG. 15. Distributionsof AmericanSyrrhopodon.

A. S. ligulatus.B. S. simmnondsii
(line);S. isthmti(asterisk).
Calymperaceae 29

cells, border of hyaline cells usually present on at CalQoene,Mori & Cardoso 17381 (NY). AMAZONAS:
least some leaves at and just above shoulders, Along Rio Negro at Ilha da Costa Arirarrn,Buck2193
(INPA,LAF,NY). BAHIA: 26 km N of Seabra, Irwinet
rarelyextendingbeyond midleafor lacking;can- al. 30844a(NY).MATO GROSSO: Serrado Cachimbo,763
cellinae distinct;mediancells obscure,quadrate, km N of Cuiabaon BR 163, Reese 16052 (INPA, LAF,
mostly ca. 7 pm, pluripapillosedorsallyandven- MO, NY). MINASGERAIS:
zwischen Piumhi und Vargem
trally; gemmae borne ventrally along costa to- Bonita, Schdfer-Verwimp10422 (herb. Schifer-Ver-
wardapex.Rarelyfertile;seta yellowish-red,3-4 wimp, LAF).PARA: Serrado Cachimbo,Base Aereado
Cachimbo,Reese 16360 (H, INPA, LAF, MICH,MO,
mm long; capsule ca. 1 mm long; peristome NY). ROND6NIA: vic. firstrapidson Rio Pacais Novos,
fragile, irregular,teeth blunt; operculum ca. 1 Reese 13589 (INPA, LAF, MO, NY). RORAIMA:Boca da
mm long. Spores 17-19 pm, granular.Calyptra Mata, 216 km N of Boa Vista, Buck 2017 (INPA, LAF,
2 mm long, rough above. NY).
BOLIVIA. Santiago de Chiquita, 1907, Herzog s.n.
Selected illustrations. Bartram (1949. fig. (JE, L, NY, S).
31E-F); Crum and Anderson (1981, fig. 110 PARAGUAY. PARAGUARI:Parque Nac. Ybycui, Buck
H-K); Crum and Steere (1957, fig. 17b); 12077 (NY). San Bernadino, Lindman B346 (NY, S).
Florschiitz (1964, fig. 57); Reese (1978, figs.
17-20). Discussion. The small stature,tightly crisped
Distribution. (Fig. 15A). Widespread but (when dry) lingulate leaves with bluntly
never common or abundantin tropicaland sub- rounded-sometimes retuse-apices, andlackof
tropical America, including the southeastern a strong borderof elongate hyaline cells on the
United States (along the Gulf Coast,Floridaand upperlamina,make this moss easy to recognize.
Georgia to Louisiana). On bark, rotting wood, Small plants of S. gaudichaudii may rather
rarelyrock, from sea level to ca. 1200 m. closely resemble S. ligulatus at first glance but
differ, among other ways, in their leaves com-
Selected specimens examined. MEXICO.JALISCO:pletely borderedwith hyaline cells and with a
Arroyo de la Resolana, Crum 631 (MICH, NY, S). sharpapical toothat the leaf tip.
NAYARIT:
near Loma de Garcia, Norris & Taranto 13482
(HSC, LAF). TAMAULIPAS: Sierra de Guatemala, near
8. Syrrhopodon simmondsii Steere, Bryologist
Ranchodel Cielo, Crum1754 (MICH).
GUATEMALA. BAJA VERAPAZ:
Mts. near Jacaro, 49: 8. 1946.Type. Trinidad.St. George:Top of
Sharp 2704 (MICH, TENN). Arima-Blanchisseuse road, Simmonds 69
COSTA RICA. CARTAGO:nearTurrialba,Hoshizaki (MICH). Fig. 17.
598-c (LAF,TENN).
PANAMA. PANAMA: Hills NE of HaciendaLa Joya, brevisetusFlorschiitz,Mossesof Suriname
Syrrhopodon
Dodge et al. 1691 7a, 16918a (MO). 1: 148. 1964.Type.Suriname: Tibitisavanne,Lanjouw
CUBA. PINARDELRio: Rio Guao, N. L. Britton et al. & Lindeman1780 (holotype,U; isotype,LAF).
10123 (NY).
DOMINICAN REPUBLIC. LA VEGA:Vie. Piedra Syrrhopodongriffinii Robinson, Phytologia 21: 390.
1971.Type.Peru.Loreto: 25 kmoestede Iquitos,14 Jul
Blanca, Allard 17156 (NY). 1965, D. & N. Griffin,II1s.n. (holotype,US; isotypes,
PUERTO RICO. Luquillomountains,Reese 14849 FLAS,LAF).
(NY).
WINDWARD ISLANDS. GUADELOUPE: Bois du Nez-
Cass, Duss 1617 (NY). Plants small, gregariousor in sods; rhizoids
VENEZUELA. AMAZONAS: CafnoManteca, Bermtidez purple; stems to 10 mm tall. Leaves linear to
PA-00677bp.p. (herb. Benrmdez, LAF). APURE: Reserva lanceolateabove narrowerbase, mostly 2-4 mm
Forestal San Camilo, Steyermark et al. 101807 (NY).
BOLIVAR:Along Rio Kavak, 5?50'N. 62?25'W, Buck &
long, erect-spreadingor a little curvedwhen dry;
Brewer 15640 (NY). marginsciliate at shoulders,cilia sometimes ex-
TRINIDAD. ST. GEORGE: Las Lapas Trace, Crosby tendingnearlyto leaf tips, borderof hyaline cells
2159 (DUKE,LAF). often weak, ending well below apex on many
GUYANA. Drainageof TakutuRiver, Smith3311a leaves; cancellinae narrow,ending in acute an-
(NY).
SURINAME. NICKERIE:Vic. Kabalebo Dam, gles distally; median leaf cells thick-walled,
Florschiitz-de Waard & Zielman 5586 (NY, U). quadrate to rounded-quadrateor rectangular,
FRENCH GUIANA. ST-LAURENT-DU-MARONI:Can- mostly 7-9 x 9-12 pm, smooth, mammillose,or
ton de Maripasoula, ca. 6 km N of Saul, 200 m, Buck18629 unipapillosedorsally,bulgingventrally;gemmae
(NY). scanty, ventralon leaf tips. Rarely fruiting;seta
BRAZIL. AMAPA:Mun. Calqoene, 30 km S of
30 Flora Neotropica

_/_l
a16 r

,, do B

!111

FIGS. 16-18. Syrrhopodon. 16. S. ligulatus. A. Leaf outlines. B. Leaf apex. A from Norris & Taranto 13482 (LAF).
B from Pursell & Reese 4164 (LAF). 17. S. simmondsii. A. Leaf outline. B. Margin at leaf shoulder. From Simmonds
2 73 (MICH). 18. S. leprieurii. A. Leaf outlines. B. Margin and cells at leaf shoulder. A from Leprieurs.n. (BM, isotype)
(two leaves at left), Mori & Kallunki 5296 (LAF) (leaf at right); B from Leprieurs.n. (BM, isotype). Scale bars: a = 1
mm (Figs. 16A, 18A); b = 0.1 mm (Figs. 17B, 18B); c = 0.5 mm (Figs. 16B, 17A).
Calymperaceae 31

short, barely exceeding perichaetialleaves; cap- SyrrhopodonrecurvulusMitten,J. Linn.Soc., Bot. 12:

sule nearly cyathiform, 0.5-1 mm long; peri- 120. 1869;Syrrhopodon gaudichaudiiMontagnevar.
recurvulus (Mitten) Th6riot, Mem. Soc. Cub. Hist. Nat.
stome lacking;operculum1 mm long. Sporesca. "FelipePoey"13: 266. 1939. Type. Cuba.Wright47
30 pm, granular-papillose.Calyptrabroadlycu- (holotype,NY; isotypes,BM, GOET,H-BR,L, REN,
cullate, 1.5 mm long. S).
Selected illustrations. Florschiitz(1964, fig. Syrrhopodon sylvaticusMitten,J. Linn.Soc., Bot., 12:
as S. Reese 121.1869.Type.Ecuador.MontafiadeCanelos,Spruce
56, brevisetus); (1978, figs. 51-52). s.n. (NY).
Distribution. (Fig. 15B). Endemicto northern Syrrhopodonscaberrimus C. Muller, Malpighia 10: 513.
South America;Colombia,Trinidad,Venezuela, 1896.Type.Guyana.Pr.Georgetown,1896,Quelchs.n.
Suriname,FrenchGuiana,Peru,Brazil, Bolivia. (lectotype,heredesignated,NY; isolectotype,BM, ex
Not common but sometimes locally abundant; K).
on sandy soil, humus, rotting wood, and trees, Syrrhopodon serpentinus C. Miller, Nuovo Gior. Bot.
Ital.,n.s. 4(1): 48. 1897.Type.Bolivia.Cochabamba,
to 600 m. pr.Choquecamata, Jun 1889, Germains.n. (lectotype,
heredesignated,NY;isolectotypes,H-BR,JE,S).
Selected specimens examined. COLOMBIA. SyrrhopodonmartiniiHerzog,Beih. Bot. Centralbl.26:
VAUPtS: Rio Piraparani, Schultes & Cabrera 17269 60. 1909. Type. Bolivia. Cerro de Santiago
(CAN). (Chiquitos),May 1907, Herzog s.n. (holotype, JE;
VENEZUELA.AMAZONAS:
Dpto. Rio Negro, along isotypes,L, NY).
Rio Mawarinuma, Sastre-de Jests 264 (LAF, NY). Syrrhopodon aculeociliatus Bartram, J. Washington
FRENCH GUIANA. ST-LAURENT-DU-MARONI: Can- Acad.Sci. 24: 469. 1934.Type.CostaRica.SanJose:
ton de Maripasoula, ca. 6 km N of Saiil, 200 m, 'Buck La Hondura,1200m. alt., 15 Aug 1933, Valerio369a
18294 (CAY, NY), 18373 (NY). (holotype,FH;isotypeMICH).
PERU. LORETO: bei Iquitos, P. & E. Hegewald 6365 Syrrhopodon cristatusBartram,Contr.U.S. Natl. Herb.
(herb. Hegewald, LAF). 26: 72. 1928.Type.CostaRica.Heredia:YerbaBuena,
BRAZIL. AMAZONAS: along Rio Uatuma off Igarape 2000 m, 22-28 Feb. 1926,Standley& Valerio49947a
Caititu, Buck 2932 (INPA, LAF, NY). PARA:Serra do (holotype,FH;isotypeNY).
Cachimbo, Base A6rea do Cachimbo, Reese 16308
(FLAS, INPA, LAF, MICH, MO, NY, US). RONDONIA: Plants small to robust,pale yellowish-greenor
vic. first rapids on Rio Pacaas Novos, Reese 13646
(COLO,DUKE, FLAS, INPA, LAF, MICH,MO, NY, darker,often appearingglaucous, tinged with
US). pink below; rhizoidsscanty, purple;stems to 5
BOLIVIA. BENI:Vie. of Guayaramerin, Reese 12772 cm tall, branched, sometimes repent. Leaves
(COLO, DUKE, FLAS, INPA, LAF, MICH, MO, NY, flexed at shoulders,only a little curved or flexu-
US).
ous when dry,3-5 mm long, tubuloseabove wet
and dry, acuminate to acute or broadly acute
Discussion. A distinct species, S. simmondsii
is recognized by its small stature;conspicuous, spinose apex; marginsentire to spinose-toothed
above shoulders, usually strongly ciliate at
delicate marginalcilia; narrowcancellinae;vari-
shouldersbut merely denticulate in some spec-
able leaf cells (at least some-often most-stel-
imens; cancellinae distinct; median cells very
late-papillose,especially distally);weak border;
narrow, nonflexed leaves; and compact habit
thick-walled,mostly ca. 5-8 x 9-12 tm, nearly
with erect or slightly curved leaves.
smooth to minutely pluripapillose to long-spi-
nose-unipapillosedorsally, highly mammillose
9. Syrrhopodon leprieurii Montagne, Ann. Sci.
to remarkably spinose-unipapillose or
Nat. Bot. 2, 2: 379. 1834. Type.FrenchGuiana. pluripapilloseventrally;gemmae not seen. Seta
Leprieur s.n. (holotype, PC; isotypes, BM, L,
red, 6-10 mm long; capsule 1.5-2.2 mm long;
peristome teeth slender,pale, pointed,papillose,
NY). Fig. 18. to ca. 250 tm tall; operculum 1-1.2 mm long.
Syrrhopodon iridans Mitten, J. Linn. Soc., Bot 12: 117. Spores ca. 14 tm, granular.Calyptra2-2.5 mm
1869. Type. Ecuador. Seemans.n., exhb. Hooker (NY). long.
Syrrhopodonpallidus Mitten, J. Linn. Soc., Bot. 12: 121. Selected illustrations. Bartram(1928, fig. 13,
1869. Type. Ecuador. Montafia de Canelos, Spruce 11 as S. cristatusBartr.);Florschiitz(1964, fig. 55);
(holotype, NY; isotype, BM). Reese (1978, fig. 21-25).
Syrrhopodonpusillus Mitten, J. Linn. Soc., Bot. 12: 120.
1869. Type. Colombia. Andes Bogotensis, Montafiade Distribution. (Fig. 20A). Endemic to and
Sogamoza,Weir153 (holotype,NY;isotypes,BM,FH). widespreadin the neotropics;CentralAmerica,
32 Flora Neotropica

D Q
o

/0 0

VI -i

*,\, A / /'00r80
%o70U'~,

o 0

isthmi.A. Leafoutlines.B. Leafbase.C. Marginandcells at midleaf.D. Portionsof sections

FIG. 19. Syrrhopodon
of leaf. From Crosby 4532 (holotype, MO). Scale bars: a = 0.05 mm (C-D); b = 1 mm (A-B).
Calymperaceae 33

Jamaica, Cuba, northern South America. INPA, LAF, MICH, MO, NY, US). RORAIMA: km 328 on
Manaus-Venezuela road, Buck et al. 1851 (INPA, LAF,
Humus, trees, rotting logs, moist rock, from sea
NY).
level to 2400 m; most frequent at higher eleva- BOLIVIA. Near Apolo, Williams 1858 (H, NY).
tions.
Discussion. This is a very variablespecies but
Selected specimens examined. COSTA RICA. AL-
AJUELA: Reserva Biologica de Bosque Nuboso de Monte readily recognizable by
the pale color of the
Verde, Crosby 10019 (LAF, MO). SANJOSe:17 km NNE plants, flexed tubulose leaves with denticulateto
of San Jose, Crosby 10872 (LAF, MO). ciliate(rarelyentire)shoulders,andthick-walled,
PANAMA. PANAMA/COLOMBIA FRONTIER: west ridge often heavily armedmediancells.
of Cerro Tacarcuna massif, Mori & Gentry 4428 (LAF,
MO). CHIRIQUi: 10 miles NW of Los Planes de Horito,
Antonio 4189A (LAF, MO). DARIEN: Pirre Massif, just W 10. Syrrhopodon isthmi Reese, Bryologist 85:
of Cana, 1200-1350 m, Allen 8854 (MO). PANAMA: near 85. 1982. Type. Panama.Prov. Panamai:Cerro
summit of Cerro Jefe, Crosby 10047 (LAF, MO). VERA- Jefe, Crosby 4532 (holotype, MO; isotype,
GUAS: below summit of CerroTute, Mori & Kallunki5296 PMA). Fig. 19.
(LAF, MO).
CUBA. Banao Hills, Clement 17(NY).
JAMAICA. ST.ANDREW: vic. Sir John's Peak, Crosby Plants robust, pink-tinged below, branched,
3040 (DUKE, LAF). more or less pendent; rhizoids violet-purple;
COLOMBIA. AMAZONAS: Surestedelapistadepespe- stemsto 5 cm tall. Leavesacuminateabove broad
gue (este de La Pedrera), Churchill et al. 16102 (HUA, shoulders, 5-6 mm long, flexuous-spreadingand
LAF, NY). ANTIOQUIA: Mun. Guatape, Sastre-de Jesis et
al. 1252 (HUA, LAF, NY). CUNDINAMARCA: Cerro de
tubulose when dry; marginsnarrowlybordered
Villeta, Steere 7774 (NY). MAGDALENA: ParqueNacional all around with hyaline cells, conspicuouslycili-
dela SierraNevada de Santa Marta,Griffin,I1 etal. 50149 ate-spinose toward leaf tip, entire below; can-
(FLAS, LAF, U). META: Cordillera Oriental, W of cellinae distinct, ventrally bulging distally;
Villavicencio, Steere 7652 (LAF, NY). NORTE DESANTAN- mediancells 5-7 x 12-16
DER:Cordillera Oriental, valley of Rio Valegri, Steere pm, smooth dorsally,
7321 (NY). SANTANDER: NW of Bucaramanga, Steere bulging ventrally;gemmae in tightclustersat leaf
7644 (NY). VALLE:Mun. Cali, Churchill et al. 15168 tips. Sporophyteunknown.
(CUVC, LAF, NY). VAUPtS:Rio Piraparana,Schultes & Illustration. Reese (1982, fig. 1-10).
Cabrera 17269 (FH, NY). Distribution. (Fig. 15B). Endemicto Panama.
VENEZUELA. AMAZONAS: Dpto. Rio Negro, 5.3 km on tree in forest.
NE of Pico Phelps, Sastre-de Jesus 358 (LAF, NY). Epiphytic
BOLIVAR:Cerro Uroi, Maguire et al. 53762 (LAF, NY). Specimen examined. The type of S. isthmi.
PORTUCGUESA: Distr. Sucre, La Divisoria dela Concepci6n, Discussion. This species is known only from
van der Werff et al. 761 '4 (MO). TACHIRA:on Rio San the original collection. Although superficially
Buena, 10 km W of La Fundacion, Liesner et al. 9555 similar to S. leprieuriiand S. erubescens, it dif-
(LAF, MO). ZULIA:near San Jos6 de Los Altos, Griffin,
II1 149 (FLAS, LAF). fers-among otherways-in lacking cilia on the
TRINIDAD. ST.GEORGE: S slopes of Cerrodel Aripo, shoulders and in having ventrally bulging can-
Crosby 2219 (DUKE, LAF). cellinae.
GUYANA. Pr. Georgetown, Quelch 1292 (NY).
SURINAME. In montibus, qui dicuntur Nassau, 11. Syrrhopodon erubescens Bartram,Bryol-
Lanjouw & Lindenman2831 (LAF, U).
FRENCH GUIANA. 1.5 km S of Charvein, Benoist ogist 54: 233. 1951. Type. CostaRica. Tilaran,
467 (LAF, U). 700 m, 15 Mar 1934,Alfaro 120a (holotype,F;
ECUADOR. NAPO: Rio Aguarico, Tangoy, Holm- isotype, MICH).Paratype,Tilaran,580 m, Al-
Nielsen et al. 20099 (AAU, NY). NAPO-PASTAZA: E slope faro 178 (FH).
of Cordillera Oriental, E of San Gabriel, Steere 9121
Fig. 21.
(LAF, NY). ORIENTE:Cordillera Oriental, near Shell- Crum& Steere,Bryologist59:
Syrrlopodonborinquensis
Mera, Steere 9373 (LAF, NY). 249. 1956.Type.PuertoRico.Sierrade Luquillo,along
PERU. LORETO:80 km NE of Iquitos, Timme 4851 Coco River,Steere7027 (holotype,MICH?,n.v.; iso-
(herb. Timme, LAF). MADREDEDios: Oct 1893, Jay s.n. type, FH). Paratypes:PuertoRico. Variouslocalities:
(NY). SANMARTN: zwischen Rioja und Balsapata, Frahm Steere 4851, 4855, 5333, 5386, 6260, 6277, 6733
et al. 2208 (LAF, U).
BRAZIL. ACRE:Rio Moa, Prance et al. 12646 (NY).
(MICH?,n.v.).
AMAZONAS: Along the Rio Marie at Manauna, Buck 2357
(INPA, LAF, NY). ROND6NIA:vie. first rapids on the Rio Plants small to medium, pale-green, tinged
Pacaas Novos, Reese 13634 (COLO, DUKE, FLAS, with pink below; rhizoidsdark-purple;stems to
34 FloraNeotropica

TROPICAL AMERICA Rom Neoteopca base map no 1

'~
----
;,-------

- -
~':- -.--
.."" -,

FIG. 20. Distributions of American Syrrhopodon. A. S. leprieurii. B. S. erubescens (ines); S. elongatus var.
elongatus (asterisk); S. elongatus var. glaziovii (dots).
Calymperaceae 35

B
21

w
fE~/i
m X 00

00
~~A o
00

00

\23 Qo O

A
A

FIGS. 21-23. Syrrhopodon. 21. S. erubescens. A. Leaf outlines. B. Margin at leaf shoulder. A from Steere 7027
(FH, isotype of S. borinquensis) (leaf at left), Alfaro 120a (FH, isotype of S. erubescens) (two leaves at right). B from
Alfaro 120a (FH). 22. S. elongatusvar. elongatus. Leaf outlines. From Ekman3799 (NY) (left), Wright49 (NY) (right).
23. S. elongatus var. glaziovii. A. Leaf outlines. B. Margin and cells at leaf shoulder. A from Irwin et al. 32324 (LAF),
B from Pringle s.n. (S). Scale bars: a = 1 mm (all leaves); b = 0.1 mm (Figs. 21B, 23B).
36 FloraNeotropica

7' cm tall but mostly much less, mostly simple. lowerlaminastronglyappressedto stem;margins
Leaves ligulate to lanceolate,3-6 mm long, vari- of upper lamina strongly bordered all around,
ously curved and contorted when dry; margins entireor tootheddistally, mostly strongly ciliate
ciliate below, cilia sometimes extending nearly at shouldersbutsometimesonly dentateornearly
to leaf tip or reducedor lacking in some leaves; entire, margins of lower lamina entire; median
cancellinae distinct; median cells mostly 6-7 x leaf cells obscure,thick-walled,isodiametric,5-
9-12 pm, dorsally smooth, ventrally bulging; 7 x 9-12 pm, highly bulging-pluripapillosedor-
gemmaeventralon leaf tips.Seta red,5 mm long; sally and ventrally; gemmae unknown. Seta
capsule 1 mm long; peristome of low, blunt, 1.5-2 mm long;capsule 2-3 mm long;peristome
imperfect teeth; operculumrostrate.Spores not teeth slender,papillose, to 150 pm tall; opercu-
seen. Calyptra2 mm long. lum 1.5 mm long. Spores 12 pm, granular.Ca-
Illustrations. Crumand Steere (1957, fig. 18, lyptra3-3.5 mm long.
as S. borinquensisCrum& Steere);Reese (1978, Illustration. Reese (1978, figs. 39-40)
figs. 56-60). Distribution. (Fig. 20B). Endemic to Cuba.
Distribution. (Fig. 20B). CostaRica (Heredia,
Puntarenas, Tilaran); Panama (Panama), and Specimenexamined.CUBA. Cuchillasde Moa, Pico
El Toldo, P6cs 9178/L (FLAS, LAF, VBI). ORIENTE:
PuertoRico. Trees in forests, to 700 m elevation.
Sierra Nipe, Shafer 3337a (NY).

Specimens examined. COSTA RICA. HEREDIA: La

Virgen, Finca la Tigra, 4 Dec 1982, Young s.n. (LAF, Discussion. This taxon is very similar to the
UWSP). PUNTARENAS: Osa Peninsula, Marenco Biologi- var. glaziovii but with the upper lamina usually
cal Preserve, Karlin 9006-0405 (NY). about twice as long as the lower lamina, and
PANAMA. PANAMA: along El Llani-Carti road, lead- twisted but not crispate when dry. Most speci-
ing N from Inter-American H., 9?15'N, 79?00'W, Crosby
10425 (LAF, MO). mens bear stout cilia at the shouldersbut occa-
PUERTO RICO. Sierra de Cayey, Reserva Forestal sional specimensareonly dentateor more or less
Carite, Reese 14667 (NY). eciliate,such as those cited by Theriotin describ-
ing his var. anomalous.
Discussion. This species is easy to recognize
by its dorsally smooth leaf cells and usuallycili- 12b. Syrrhopodon elongatus var. glaziovii
ate margins. Specimens of S. leprieurii may be (Hampe)Reese Fig. 23.
similar but almost always have the leaf cells
variouslypapillosedorsally;S. isthmiis also sim- Syrrhopodon glazioviiHampe,Vidensk.Meddel.Dansk
ilar but lacks cilia on the leaf shouldersand has Naturhist.Foren.Kjobenhavn,ser. 3, 6: 133. 1874.
Type.Brazil.Rio de Janeiro,Glaziou7134 (holotype,
distally bulging cancellinaeventrally. BM;isotypes,NY, S).
12a. Syrrhopodon elongatus Sullivant var. Thevar.glaziovii differsfromvar.elongatusas
elongatus Fig. 22. noted above, particularlyin its shorter leaves
which are crispate-contortedwhen dry.
Syrrhopodon elongatus Sullivant, Proc. Amer. Acad. Illustration. Reese (1978, figs. 41-43).
Arts 5: 280. 1861. Type. Cuba. Wright 49 (holotype,
FH; isotypes, BM, GOET, L, NY). Distribution. (Fig. 20B). Mexico (see Reese,
Syrrhopodon elongatus var. anornalus Th6riot, Mem. 1978 for discussionof this possibly dubious oc-
Soc. Cub. Hist. Nat. "Felipe Poey" 13: 267.1939. Type.
Cuba. Oriente: Taco Bay, minas de Iberia, Ekman3799
currence);Venezuela;Guyana;southernBrazil.
(lectotype, S; isolectotypes, NY, US). Syntype. Cuba. Selected specimens examined. MEXICO. OAXACA:
Sierra Maestra, Ekman 7122 (NY, PC, S).
Fucha fan, Oct 1898, Pringle s.n. (NY, S).
VENEZUELA. BOLIVAR: Steyermark93396 (LAF, US).
Plants in dense, deep tufts and cushions, pale GUYANA. Upper Mazaruni River region, Boom &
glaucous-green above; rhizoids purple, usually Gopaul 7672 (NY).
BRAZIL. BAHIA:Summit of Morro do Chap6u, Irwin
conspicuous on leaf tips and bases; stems to 15 et aL 32324 (LAF, NY). PARANA:Icarehy, Dusen 6594
cm tall, little branched.Leaves broadly acumi-
(NY). Rio DEJANEIRO:Glaziou 7138 (NY, S). SANTA
nate above broaderbase, strongly flexed, 9-11 CATARINA: Joinville, 1904, Grossman s.n. (NY). SAo
mm long, crispate to spirally twisted when dry, PAULO: Cardoso Island, Vital 3166 (LAF, SP).
Calymperaceae 37

jrY/1 / a _________

/ b ....
C
d

C B

FIGS. 24-25. Syrrhopodon. 24. S.fimbriatus. A. Leaf outlines. B. Margin and cells at leaf shoulder. A from P. & E
Hegewald 6345 p.p. (hb. Hegewald) (left), Liitzelburg 22914 (JE) (right). B from Liitzelburg 22914 (JE). 25. S.
parasiticus. A. Outlines of noncomal leaves. B. Outline of comal leaf with gemmae. C. Distal portion of gemma. A
from Hoehne s.n. (JE), B-C from Reese 16805 (LAF). Scale bars: a = 0.05 mm (Fig. 25C); b = 1 mm (Figs. 25A-B);
c = 0.1 mm (Fig. 24B); d = 0.5 mm (Fig. 24A).
38 FloraNeotropica

Discussion. The Steyermarkspecimen of var. to recognizeby its characteristichabitand small,

glaziovii from Venezuela,cited above,andothers lingulate, usually ciliate leaves. The occasional
I have seen fromBolivar, Venezuela,have some specimens whose leaves lack cilia are still easy
or all of their leaves more or less eciliate at the to recognizeby theirappearanceand leaf shape.
shoulders,as in some of the Cubanspecimensof
the var. elongatus. However, usually at least 14. Syrrhopodon parasiticus (Bridel)
some leaves in each specimen bearstout teethor Bescherelle, Ann. Sci. Nat. Bot. 8, 1: 298.
cilia at the shoulders. 1895. Fig. 25.

13. Syrrhopodon fimbriatus Mitten, J. Linn. BryumparasiticumBridel,Muse. Rec. 2(3): 54. 1803;
Soc., Bot. 12: 122. 1869. Type. Brazil. In Anictangium parasiticum(Bridel)Bridel,Spec. muse.
1: 270. 1806; Brachypodiumparasiticum(Bridel)
arboribusad Panure, flumi Vaupes, Spruce 9 Bridel, Bryol. univ. 1: 149. 1826; Calymperes
(NY). Fig. 24. parasiticum(Bridel)Hooker& Greville,EdinburghJ.
Sci. 1: 131. 1824; Calymperopsisparasitica(Bridel)
Plants very small, yellowish-green, in low, Brotherusin Engler& Prantl,Nat. Pflanzenfam.ed. 2.
10:235. 1924;Encalyptaparasitica (Bridel)Swartz,Fl.
compact sods or gregarious; rhizoids purple; Ind.occid.3: 1759. 1806;Glyphomitriumparasiticum
stems to 10 mm tall but mostly much shorter. (Bridel) Bridel, Mant. muse. 31. 1819; Weissia
Leaves lingulate from scarcely broaderbase, 1- parasitica(Bridel)Mohr,Ann.Bot. (K6nig& Sims)2:
1.5 mm long, tightly crisped when dry; margins 545. 1806.Type.Jamaica.Swartzs.n. (holotype,BM;
of upperlaminanarrowlyborderedwith hyaline isotypeNY).
CalymperesswartziiAmott, Mem Soc. Linn. Paris 5:
cells, ciliate from below shouldersto leaf tip, or 233. 1827,nom.illeg. incl.sp. pr.
cilia lacking in distal part of leaf (cilia rarely CalymperesfiligerumAustin, Bot. Gaz. 4: 151. 1879.
lacking altogetherand the leaves nearlyor quite Syrrhopodonfiligerus (Austin)Williams,Bull. Torrey
entire);cancellinaeroundedabove or sometimes Bot.Club47:384. 1920.Type.U.S.A.Florida:Caloosa,
narrowand acute;medianleaf cells thick-walled, Mar1878,Smith& Austins.n. (NY).
Syrrhopodonwainioi Brotherus,Acta Soc. Sci. Fenn.
5-7 x 7-12 pm, each with a single, large blunt 19(5): 14. 1891. Calymperopsiswainioi (Brotherus)
papilla dorsally (rarely mammillose), bulging- Fleischer,Biblioth.Bot. 80: 5. 1913. Type. Brazil.
mammillose or papillose ventrally;gemmae not MinasGerais: Sitio,1885,Wainios.n.(holotype,H-BR;
seen. Seta 9 mm long; capsule ca. 1 mm long; isotype,S).
Calymperessemilimbatulum C. Miiller, Hedwigia 39:
peristomeof imperfect,bluntteethca. 70 pm tall; 261. 1900.Type.Brazil.Novo-Friburgum, Peterss.n.
operculum 0.75 mm long. Spores 14-17 pm, (lectotype,heredesignated,NY).
granular.Calyptra 1.75 mm long. SyrrhopodonmartinicensisBrotherusin Urban,Symb.
Illustration. Reese (1978, figs. 53-55). antill. 3(3): 422. 1903; Calymperopsismartinicensis
Distribution. (Fig. 27A). Endemicto northern (Brotherus) Brotherusin Engler & Prantl, Nat.
Pflanzenfam.ed. 2. 10: 235. 1924. Type. Martinique.
Amazonia; Colombia, Venezuela, Peru, Brazil. Mome Rouge, Duss 372 (holotype,H-BR; isotype,
On trees (sometimes in the crowns), rarely on NY).
logs and sandstone,mostly at low elevations,but Calymperesfiligera Mitten ex Tixier, Rev. Bryol.
collected to 1000 m. Lich6nol.35: 279. 1967,nom nud.in syn.

Selected specimens examined. COLOMBIA. AMA- Plants dark-green,solitary,in clustersof a few

ZONAS:Oeste de La Pedrera,a lo del lado sur del Rio or gregarious;rhizoidsbrown;stems to 2 cm tall
Caqueta, Churchill etal. 16079 (LAF, NY). META:W. of but mostly much shorter, not much forked.
Villavicencio,Steere7652 (NY).
VENEZUELA. AMAZONAS: Dpt. Rio Negro, Cerro de
Leaves to 5 mm or more long, involute and in-
Neblina, Buck 11615 (NY). BOLIVAR: CerroGuaiquinima, curvedwhendry,basicallydimorphic,vegetative
5?40'N, 63'34'W, Sipman 26622 (B, NY). ones lingulate-lanceolate, gemmiferous ones
PERU. LORETO: ca. 5 km SE of Iquitos airport,Timme
4684 (herb. Timme, LAF).
broadlylanceolateto triangularand clusteredin
BRAZIL.ACRE:Slopesof CerrodaMoa,Pranceet al. comae at stem tips; marginsentire, mostly bor-
12131 (INPA, NY). AMAZONAS: km 380 along Manaus- deredat least in partwith elongate,hyaline cells,
Porto Velho road, Prance et al. 22869 (INPA, LAF, NY). sometimesunbordered;cancellinaedistinct,dis-
tally acute; median cells ca. 7 ,Lm,smooth to
Discussion. This is a beautiful little moss, easy unipapillose dorsally and ventrally; gemmae fil-
Calymperaceae 39

amentous,borne ventrallyalong costa from near TRINIDAD. ST. ANDREW: Quare Road near gate of
Hollis Reservoir, Crosby 2367 (DUKE, LAF).
cancellinaeto leaf tip. Seta reddish,3-4 mm long; SURINAME. near Paramaribo,J. & P. Florschiitz 873
capsule 1-2 mm long; peristometeeth irregular, (LAF, U).
blunt;operculumca. 1mmlong.Spores24-36 pm, FRENCH GUIANA. ST-LAURENT-DU-MARONI: Can-
irregularin shape,granular.Calyptra3 mm long. tonde Maripasoula,ca. 6 km N of Sail, 200 m, Buck 1 8365
(NY), 18368 (CAY, NY).
ECUADOR. GALAPAGOS ISLANDS: Is-
Selected illustrations. Bartram (1949, fig. land, Mt. Crocker, Howell 106 (FH). Indefatigable PASTAZA: Puyo,
35DG); Crum and Anderson (1981, Fig. 113); Steere E-69 (NY). PICHINCHA: 18 km NW de San Miguel
Crum and Steere (1957, fig. 17c); Florschiitz de las Bancos, Buck 10300 (NY).
PERU. PASCO:ca. 5 km von Oxapampa, P. & E.
(1964, fig. 58ah); Reese (1978, figs. 28-31); 8443 (herb. Hegewald, LAF).
Reese et al. (1986, figs. 51-52). Hegewald
BRAZIL. AMAZONAS: km 45 on Manaus-Boa Vista
Distribution. (Fig. 27B). Pantropical;through- road, Buck 2151 (INPA, LAF, NY). BAHIA: Itabuna,
out tropical and subtropical America. A strict CEPLAC, Yano 2310 (LAF, SP). MATOGROSSO:Near
Rio Arica, Vital 9951 (LAF, SP). PARA: Serra do
epiphyte on twigs, branches,trunks,and leaves Base Aerea do Cachimbo, Rio Formiga, Reese
in forests; to ca. 1600 m but most frequentat Cachimbo, 16291 (DUKE, FLAS, INPA, LAF, MICH, MO, NY).
lower elevations. PARANA: Mun. Medianeira, Vital& Buck 12106 (NY, SP).
Rio DEJANEIRO: Glaziou 9237 (BR, G). RORAIMA: km 328
Selected specimens examined. MEXICO.JALISCO:on Manaus-Venezuela road (BR 174), Buck et al. 1848
Arroyo de la Resolana, Crum 615a (MICH). VERACRUZ: (INPA, LAF, NY). SANTACATARINA: 6 km E of Palma
20 mi. W of Tuxpan, Reese & Pursell 4776 (LAF, MEXU, Sola, Vitt21229 (ALTA, LAF). SAOPAULO: between Sao
PAC, US). YUCATAN:Chichankanal,Gaumer 1278 (LAF, Paulo and Santos, km 42, Vitt20662 (ALTA, LAF).
NY). BOLIVIA. BENI:Vic. Guayaramerin, Reese 12728b
GUATEMALA. ALTAVERAPAZ: Between Tactic and (NY).
divide on road to Tamahu, Standley 90775a (F). PETEN: PARAGUAY. ALTOPARANA: 31 km N of Heman-
Chicbul, La Libertad, Lundell 4455 (MICH). darias, Buck 12349 (NY). PARAGUARI: Parque Nac.
BELIZE. EL CAYO:Valentin, Mains 3645 (MICH). Ybycui, Buck 11947 (NY).
HONDURAS. ATLANTIDA: Near Tela, Crosby 4089
(MO).
Discussion. This is a well-markedtaxon andis
NICARAGUA. 'Eastern Nicaragua," Cama River, easy to recognize by its very acute cancellinae,
Cukra Hill zone, Shanks 5b (LAF, NY). dimorphicleaves with filamentous gemmae on
COSTA RICA. CARTAGO:vie. of Turrialba,Godfrey the
66222 (LAF, NY). LIM6N: La Asunci6n, Rio Banano,
largerones, and often incomplete borderof
Gutierrez 151 (FH, NY). hyalinecells-completely lackingin some spec-
PANAMA. CANALZONE:Barro Colorado Island, imens. Thebroadgemmiferousleaves often form
Arrocha 393 (PMA). CHIRIQUI: Alhajuela, within 10 distinctivecomae at the stem tips. Syrrhopodon
miles of Boquete, Corman 3014a (FH). COCLt: 5 miles N graminicolais similarbuthas pluripapillosecells
of El Valle, Tyson& Godfrey 2476A (LAF, NY). PANAMA: and a
area of summit of Cerro Jefe, Crosby 10067 (LAF, MO).
complete distally toothed border.
CUBA. PINARDELRio: NE of San Diego de los Bafio, Syrrhopodonflexifolius
is also similar but has a
Le6n 3564 (NY). pinkishtinge in lower parts,usually completely
JAMAICA. ST. ANDREW-PORTLAND BORDER: 0.5 mi. N borderedleaves, and much larger cells that are
of Newcastle, Crosby 3102 (DUKE, LAF). neverdistinctlypapillose dorsally(althoughlow
DOMINICAN REPUBLIC. PEDERNALES: Las remoteindistinctdorsalpapillae are rarelypres-
Abajas, Reese 14939 (NY).
PUERTO RICO. CaribbeanNational Forest, Luquillo ent).Syrrhopodon cymbifoliusis like a miniatureS.
Division, along trail 1.9 mi. S of entrance to El Verde parasiticus
but only rarelyhas comae of enlarged
Biological Station, Reese 14864 (LAF, NY). leaves. Small forms of S.parasiticusmay be diffi-
WINDWARD ISLANDS. GUADELOUPE: More du cultto distinguishfromS. cymbifoliuswhensterile.
Bananier, Duss 1643 (NY). DOMINICA:St. George, Roseau See commentsunderthe lattername.
Tal, E. & P. Hegewald 9300 (herb. Hegewald, LAF).
COLOMBIA. ANTIOQUIA:Mun. Santo Domingo,
Churchill et al. 14714 (NY). 15. Syrrhopodon flexifolius Mitten, J. Linn.
VENEZUELA. AMAZONAS: Dpto. Rio Negro, Soc., Bot. 12: 118. 1869. Type. Brazil. Sao
00?50'N,66?10'W, Buck 12777(NY). DISTR.FED.CARA- Gabriel,Spruces.n. (lectotype, NY). Panure,
CAS: Ramirez-Reyes 427 (Herb. Instituto Pedagogico Ca-
Spruce 11 (syntype,NY). Fig. 26.
racas, LAF). MONAGAS:vic. Caripe, Pursell et al. 9200
(LAF, PAC). SUCRE:entre Cumanacoa y Satillo, Syrrhopodon parasiticus (Bridel) Bescherelle var.
Bermiudez008-77 (herb. Bermudez, FLAS, LAF). flexifolius (Mitten) Reese, Bryologist 84: 245. 1981.
40 Flora Neotropica

I
I ' c,

": '" A ; '

(A-B);b 0. mm (C-D).
26 Syrrhopodonflexifolius.
FIG.
FIG. 26. Syrrhopodonflexfoiu. A.A. Noncomal
Noncomalleaves. B. Comal
leaves.B. Comalleaves, twowith
leaves,two withgemmae. C. Margin
gemma. C. andcells
Marginand cells at
at
midleaf. D. Portions
midleaf. D. of sections
Portions of of leaves.
sections of leaves. A-C from Buck
A-C from 2442 (NY),
Buck 2442 (NY), D fromi
froni Buck
Buck 2636 (LAF). Scale
2636 (LAF). bars: a = 1 mm
Scale bars: mm
(A-B); b = 0.1 mm (C-D).

Plants tufted,soft, glossy, green to yellowish- gular or rounded, mostly 10-14 x 14-17 um;
green above, darkerin older parts, often tinged gemmae long-filamentous, borne ventrally along
with pink below; rhizoids purple-brownto red- the costa. Sporophytes unknown.
dish, sometimes conspicuous in lower parts of Illustration. Reese (1981, fig. 8-15).
stems;stems to 2 cm tall, oftenrepentandbranch- Distribution. (Fig. 27B). Costa Rica; Panama;
ing. Leaves dimorphic, 1.54 mm long, crispate- Venezuela (Amazonas, Bolivar); Brazil (Amazo-
flexuous when dry, spreadingto recurvedwhen nas). Tree trunks, branches, twigs, logs, in forests
moist, oblong to lanceolate;gemmiferousleaves to 1200 m.
short and broad,formingcomae, sometimes del-
toid; margins usually borderedall aroundwith Specimens exaniined. COSTA RICA. PUNTARENAS:
elongate hyaline cells, mostly entire, sometimes 9.5 km NW of Rinc6n de Osa, Bowers 327-a (LAF,
with blunt denticulationsabove; hyaline border TENN).
sometimes reducedor lackingdistally,especially PANAMA. DARI:N: Pirre Massif, just W of Cana,
800-1200 m, A lien 8903 (MO).
in gemmiferous leaves; cancellinae broad and VENEZUELA. AMAZONAS: Dpto. Rio Negro, along
distinct, distally acute, pale and conspicuously Rio Mawarinuma, Buck 11254 (NY). BOLIVAR: Cerro
exposed when dry;mediancells smoothand pel- Guaiquinima, 5?49'N, 63?32'W, Sipnman27072 (B, NY).
BRAZIL. AMAZONAS:vic. first cachoeira of Igarape
lucid, in vertical files, mostly collenchymatously
Foibara off Rio Negro, Buck 2636 (INPA, LAF, NY).
thickened,often porose, plane dorsallyor rarely
with low remoteindistinctpapillae,nearlyplane
to slightly bulging ventrally,quadrateto rectan- Discussion. This rare moss is known only from
Calymperaceae 41

TROPICAL AMERICA Fora Neotropca base map no 1

------------------
-.------------------

.1

- .. --- -- ---------- ~

FIG. 27. Distributions of American Syrrhopodon. A. S. fimbriatus (line); S elatus (dots in South America); S.
S. theriotii
steyerrniarkii(asterisks); S.
steyermazrkii theriotii (dot in Costa Rica). B.
in Costa S.parasiticuls
B. S. S.flexifolius
parasiticus (lines); S.fiexifolius (dots).
42 Flora Neotropica

a relatively few collections from small area of in size and shape, 21-47 um diam., spherical to
northernAmazonia and a single collection each ellipsoidal, dark-green, finely granular, some
from Costa Rica and Panama. It is generally small, colorless, empty, and collapsed. Calyptra
similar to S. parasiticus but differs consistently 2.5 mm long, divided into several narrow seg-
as noted under that taxon. In my revision of ments frombase to rostrum.
limbate Syrrhopodon (1978), I treated S. Selected illustrations. Florschiitz (1964, fig.
flexifolius as a synonymof S.parasiticus(follow- 58 i-j), Reese (1978, figs. 32-34), both as S.
ing Florschiitz, 1964 and Tixier, 1967), but later parasiticusvar. disciformis.
(1981) recognized it at the varietal level. Here I Distribution. (Fig. 31A). Tropical America;
restore the taxon to specific rank based on its Belize, Panama,Cuba,Hispaniola,Jamaica,Co-
unique qualities as noted above, which do not lombia,Guyana,Suriname,FrenchGuiana,Ecu-
intergradewith S. parasiticus. ador,Brazil (Amazonas,Para, Sao Paulo). Tree
trunksand twigs; to ca 500 m.
16. Syrrhopodon graminicola Williams, Bull.
Torrey Bot. Club 47: 379. 1920. Type. Ja- Selected specimens examined. BELIZE. EL CAYO:
maica. Woodstock,nearBeaufort,E. G.Britton SanAgustin,Mains3982 (MICH).
PANAMA. VERAGUAS: Taylor 1107, 1121 (MICH).
579 (NY). Fig. 28. CUBA. ORIENTE: Sierra near
Nipe, Woodfred,Shafer
3732a (NY).
Calymperesdisciforme C. Miiller,Linnaea21:183. 1848; JAMAICA.Thetype.
Calymperopsisdisciformis(C. Miller) Tixier, Rev. HAITI.CampPenrin,Curtis45 (NY).
Bryol. Lich6nol. 35: 290. 1967, hom. illeg., non DOMINICAN REPUBLIC. LA VEGA:vic. Piedra
Calymperopsis disciformis (Dus6n) Fleischer; Blanca,Allard18899 (NY).
Syrrhopodon parasiticus(Bridel)Bescherellevar.dis- COLOMBIA. AMAZONAS: E of Villazul, Isla
ciformis(C.Miiller)Florschiitz,Mossesof Suriname1: Mariniamoin riverCaqueti,0?45'S,72?06'W,Sipman&
154. 1964. Type. Suriname,Kegel 505 (GOET). Duivenvoorden28153 (B, NY).
SyrrhopodonschiffneriBrotherus,Akad. Wiss. Wien, GUYANA.UpperMazaruni district,5 52'N,60?37'W,
Math.-Natur. K1.,Denkschr.83: 281. 1926.Type.Bra- 17063 (U).
zil. Sao Paulo:Pr. Conceicode Itanha6m,3 Jul 1901, Aptroot
SURINAME.BrownsbergPlateau,J. & P. Florschiitz
Schiffners.n.(holotype,H-BR;isotype,BM). 4636A(U).
FRENCH GUIANA. ST-LAURENT-Du-MARoNI:
Can-
Plants dull-green, solitary, gregarious, or ton de Maripasoula,ca. 6 km N of Saiil, 200 m, Buck
tufted; rhizoids scanty, reddish-brown;stems to 18344,18653(NY);Sauil,3?32'N,53?12'W,Montfoort&
1 cm tall but mostly shorter. Leaves dimorphig Ek 788 (U).
ECUADOR.NAPO:Near SantaClara,Steere E-77
vegetative acuminateto linear or broadlylinear (NY).
above scarcely broaderbase, acute, 2-3.5 mm BRAZIL.AMAZONAS: along Rio Uatuma,Buck2791
long, crispate-contortedwhen dry, erect-spread- (INPA,LAF,NY).PARA: Serrado Cachimbo,BaseAerea
ing to squarrosewhen moist, often folded-falcate do Cachimbo,Reese16437 (INPA,LAF,NY).
on microscope slide; margins of upper lamina
undulate,borderedall around,toothed in upper Discussion. Some collections of this moss lack
portion, margins of lower lamina entire; can- comae of enlarged gemmiferous leaves, which
cellinae acute to broadly acute distally; median areso distinctivewhen present.AlthoughMiiller
leaf cells obscure, isodiametri9mostly 6-10 x describedthe sporophyteof his Calymperesdis-
7-10 pm, pluripapillosedorsally and ventrally; ciformein the type description,the type material
gemmiferous leaves oblong-acuteto nearly del- at GOETis sterile. The descriptionof the sporo-
toid, gemmae filamentous, borne along costa at phyte above is based on the only two fruiting
midleaf. (Description of sporophyte based on specimens I have seen, from French Guiana
Buck 18344, 18653, NY.) Seta reddish-brown, (Buck18344, 18653, NY). The variablespores of
2-2.5 mm long; capsule ovoid-cylindric,brown, Buck 18653-the only specimen with spores-
1-1.5 mm long, set somewhat obliquely on the may indicate that the sporophytewas a hybrid.
seta, stomatanot seen; peristometeethimperfect, Hybridization is a possibility because S.
withyellow-orangetips,smooth,segmented,52- graminicola shares the same habitat with the
65 pumtall, bases fused into a pale membrane; relatedS. parasiticus,althoughthe latterwas not
operculum1 mm long. Sporesextremelyvariable presentin Buck 18653.
Calymperaceae 43

, 2 ,
\^I'

A B A

B.

~'30 3

FIGS. 28-30. Syrrhopodon.28. 5. graminicola. A. Noncomal leaves. B. Comal leaf. A from E. G. Britton 579 (NY)
(left), Schiffner s.n. (H) (right). B from Schiffner s.n. (H). 29. 5. cymbifolius. A. Leaf outlines. B. Margin and cells at
leaf shoulder. C. Portion of leaf section. D. Sporophyte with detached operculum. A-C from Buck 2 752 (LAF), D from
Ule 2272 (H, holotype of S. ramicola Broth.). 30. S. helicophyllus. A. Leaf outline. B. Cells at midleaf. C. Distal portion
of gemma. A from Griffin et al. 407 (LAF), B-C from Griffin et al. 542 (LAF). Scale bars: a = 1 mm (all leaves and
Fig. 29D); b = 0.11 mm (Figs. 29B-C, 30B-C).
= 0. 30B-C).
44 FloraNeotropica

Plants of S. graminicola are grossly similar to erous leaves sometimes presentin comae, vege-
those of S. parasiticus but differ consistentlyin tativeleaves oblong-lingulateto broadlylanceo-
having pluripapilloseleaf cells and the leaf mar- late, tubulose and secund when dry, 2-3.5 mm
gins completely borderedwith hyaline cells and long;marginsentirewherebordered,serrulateby
toothed above. Further,the leaf marginstend to papillaeand cell angles where unbordered,nar-
be undulate,especially when dry,a conditionnot rowly borderedin lower 1/2-3/4 with hyaline
seen in S. parasiticus where the margins are cells, or borderirregularand restrictedto shoul-
plane. This a very rare taxon known from rela- der region or more or less lacking (the hyaline
tively few collections. It is easily distinguished cells sometimes intramarginaland teniola-like);
from S. parasiticus, which it most closely resem- cancellinaeending distally in acute angles; me-
bles, as noted above. The undulateleaf margins dianleaf cells thick-walled,isodiametricto short-
allow easy recognition when the plants are dry. rectangular,mostly 6-8 x 9-10 pm, sharply
The oldest name for this species is Calymperes unipapillose dorsally, strongly mammillose to
disciformeC. Mull. (1848), but the combination mammillose-papillose ventrally, lumina often
in Syrrhopodonis preemptedby S. disciformis triangularas seen in section; gemmae unknown.
Dus. (1895), which representsanotherspecies. I Monoicous; sporophytes minute, equalled or
followed Florschiitz (1964) in my treatmentof slightly exceeded by perichaetialleaves; setae
this taxon (1978) as a variety of S. parasiticus, 1-3 perperichaetium,1.5-2.2 mm long; capsule
but based on furtherexperience, including field 1-1.3 mm long;peristometeethbrown, delicate,
observation,I now recognize it as a distinctspe- transverselybarred,ca. 85 pm tall;operculumca.
cies. It does not intergradewith S. parasiticus. 0.75 mm long. Spores smooth, 16.8-21.6 pm.
Tixier (1967) included Africa in the range of Calyptracucullate,deeply lobed at base, sharply
this taxon [which he treated under the name papillose above, ca. 1.8 mm long.
Calymperopsis disciformis (C. Mill.) Tixier, Illustration. Reese (1981, figs. 1-7).
hom. illeg.] and listed severalnamesin its synon- Distribution. (Fig. 31A). A rare endemic of
SouthAmerica;Brazil(Amazonas,EspiritoSanto,
ymy based on Africantypes. While specimensof
the African and Americanpopulationsare cer- Goias,Parai,Sio Paulo);Colombia(Antioquia).
tainly similar, they differ in thatthe leaves of the Selected specimens examined. COLOMBIA. ANTI-
formerare generally narrowerand more acumi- OQUIA:Mun.GomezPlata,Churchillet al. 14805 (HUA,
nate,longer(longest leaves 3.5-4 mm for African LAF,NY).
materialvs. 3-3.5 mm for Americanspecimens), BRAZIL. AMAZONAS: along Rio Uatuma between
and have smaller leaf cells (5-8 x 5-9 pm for cachoeirasMiritiand Morena,Buck2752 (INPA,LAF,
African specimens vs. 6-10 X 7-10 pm for NY). ESPRITO SANTO: 18?54'S, 40?3'W, Schdfer-Ver-
wimp& Verwimp8824 (herb.Schifer-Verwimp,LAF).
Americanspecimens). Therefore,the two popu- PARA:Serrado Cachimbo,vie. Rio Jamanxim,km 1023
lations can be recognized as distinct.The oldest on Cuiaba-Santaremroad(BR 163),Reese16700 (INPA,
namein Syrrhopodonfor the Africanpopulations LAF,NY). SAOPAULO: bei Bertioga,Schdfer-Verwimp&
is S. africanus (Mitt.) Paris (basedon Thyridium Verwimp 11749 (herb.Schifer-Verwimp,LAF).
africanumMitt.).
Discussion. This tinymoss is remarkablein the
17. Syrrhopodon cymbifolius C. Muller, Calymperaceae for being monoicous, in contrast
to the usual dioicous condition.It closely resem-
Hedwigia 39: 262:1900. Type. Brazil. Goyaz bles small forms of S.
[Goias]: flum. Corumba,Ule 1571 (lectotype, smaller parasiticus but differs in
here designated,NY). size, smaller spores, usual lack of en-
Fig. 29.
largedgemmiferousleaves in comae, andin hav-
Syrrhopodon ramicola Brotherus, Hedwigia 45: 270. ing stomata at the base of the capsule.
1906.Type.Brazil.Amazonas:JuruaMiry,Rio Jurui,
Ule 2272 (holotype,H; isotype,S). 18. Syrrhopodon helicophyllus Mitten, J. Linn.
Soc., Bot. 12: 119. 1869. Type. Brazil. Rio
Plants tiny, gregarious;rhizoidspurple-brown, Negro, Spruce 8 (lectotype, here designated,
inconspicuous; stems 3-4(-5) mm tall. Leaves NY; isolectotypes, BM, BR). Panure, Spruce
crowded, rarely dimorphi9short broadgemmif- s.n. (syntype,NY). Fig. 30.
Calymperaceae 45

TROPICAL AMERICA FloraNeotrolpcabase map no 1

~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.
' ...

lf<~?-- \--? K t--

e\ ^--- L- I-.

1.Ditrbuios f meicn
FI. lns;S
yrhpodn .Srmncl ybfludt) .S eiohlu
line;S
dase yooiies(otmu ie)

FIG. 31. Distributions of American Syrrhopodon.A. S. graminicola (lines); S. cymbifolius (dots). B. S. helicophyllus
(dashed line); 5. lycopodioides (continuous lines).
46 FloraNeotropica

Plants robust, glossy above, wiry, in tangled Plants robust,yellowish-green above, darker
mats; rhizoids dark-red;stems to ca. 3 cm tall, below, in dense, spreadingclumps or pendent,
often repentand rhizome-like,sometimesmatted branched;rhizoidsscanty,brown;stems to 15 cm
with rhizoids. Leaves linearto tapering,3-4 mm tall. Leaves linear to linear-lanceolate from
long, upper lamina often helically coiled when scarcelybroaderbase, 7-10 mm long, spreading
dry; margins entire all around, bordered all to erect-appressedwhen dry; margins promi-
aroundwith thickbandof elongatehyaline cells; nently bordered by elongate cells, coarsely
cancellinaepersistent,very distinct;median leaf toothed above, leaf base usually bordered by
cells thick-walled, porose, pellucid, square to severalto manyrows of elongate,hyaline,porose
rectangular,mostly 12 x 36 pm, smooth and flat cells; cancellinae distinct, supra- and para-
dorsally, slightly bulging ventrally;gemmae fil- cancellinarcells usually with heavily thickened,
amentous,septate,to ca. 160 pm long, dorsaland porosewalls;medianleaf cells mostly squarebut
ventralalong costa on old, brown, lower leaves. sometimes some or all transversely elongate,
Seta red, 4 mm long; capsule 1.5 mm long;peri- mostly ca. 12-15 pm, bulging mammillose-pa-
stome ca. 145 pm tall,fairlywell developed,teeth pillose dorsallyand ventrallyor nearly smooth;
slender,articulate,verticallystriateon outersur- gemmaeinfrequent,brown,filamentous,on ven-
face; operculum 1.4 mm long. Spores 17 pm, traland dorsalsurfacesof costa from tips nearly
granular.Calyptra2-2.2 mm long. to shoulders, or restricted to distal portion of
Illustration. Reese (1978, figs. 65-66). costa. Seta dark, 10-11 mm long; capsule 3-4
Distribution. (Fig. 31B). A rare endemic of mm long; peristome of fragile, imperfect teeth
northernSouth America. Colombia(Amazonas, scarcely projecting above rim of capsule
Vaupes);Venezuela(Amazonas,Bolivar);Brazil mouth; operculum 1.5-2 mm long. Spores
(Amazonas). In dense, springy mats on tree granular, ca. 20 pm. Calyptra 2-3 mm long,
trunks,rarelyon soil, at elevations to ca. 200 m. smooth.
Selected specimens examined. COLOMBIA.AMA- Selected illustrations. Bartram (1949, fig.
ZONAS:Oestede LaPedrera,a lo del ladodel RioCaqueta, 31G-H);Reese (1978, figs. 61-64).
Churchill et al. 16802 (LAF, NY). VAUPtS: Rio Distribution. (Fig. 31B). Mexico, Central
Piraparani,Schultes& Cabrera17552 (FLAS,LAF). America, West Indies, South America. Tree
VENEZUELA. AMAZONAS: Rio Negro, Along Rio
trunksandbases, rottingwood, rocks, soil in wet
Mawarinuma,Buck 11054 (LAF, NY). BOLIVAR: Rio
49
Uriman,Holmquist (FLAS,LAF,MY). montane forests, from ca. 150 to 3000 m, but
BRAZIL. AMAZONAS: km 60 along Manaus-Carcarai uncommonbelow 1000 m.
road,Griffin,IIIet al. 542 (FLAS,LAF,SP).
Selected specimens examined. MEXICO. VERACRUZ:
Discussion. This very distinctivemoss is char- 3 km N of Santa Rita, Delgadillo M. 3476 (LAF, MEXU).
acterized by its large, smooth, porose leaf cells GUATEMALA. BAJAVERAPAZ:Over mt. from Finca
and its often helically coiled leaves. Its gemmae Bucaral, Sharp 2760 (MICH, TENN).
COSTA RICA. ALAJUELA: 7 miles N of San Ram6n,
are almost identicalto those of S. parasiticus and Bowers 394 (LAF, TENN). LIMON: Rio Banana, Gutierrez
S. flexifolius, but are borne in a very different 192 (NY). SANJOSE:La Palma, Standley38160/b (JE).
way, on both surfaces of the leaf. PANAMA. COCL.: Near El Valle de Anton, Brako
8412 (NY). PANAMA: areaof summit of CerroJefe, Crosby
10054 (LAF, MO).
19. Syrrhopodon lycopodioides (Bridel) C.
CUBA. Pico Turquino, Leon 11266 (NY).
Miller, Syn. musc. frond.1: 538. 1849. JAMAICA. ST. ANDREW:ca. 2 mi. N of Cinchona,
Fig. 32. Crosby 2988 (DUKE, LAF).
PUERTO RICO. Luquillo Mountains, vic. Pico de
Bryum lycopodioides Bridel, Muscol. recent. 2(3): 54. Este, Reese 14778 (NY).
1803; Dicranum lycopodioides (Bridel) Swartz, Fl. Ind. WINDWARD ISLANDS. GUADELOUPE: 1862,
occid. 3: 1766. 1806; L'Herminier s.n. (GOET, REN). DOMINICA: St. George,
Orthothecalycopodioides (Bridel) Bridel, Bryol. univ. 1: zwischen Freshwater and Boeri lakes, E. & P. Hegewald
778.1827. Type. Jamaica. Swartzs.n. (holotype, B,n.v.; 9414 (herb. Hegewald, LAF). MARTINIQUE: Montagne
isotypes, BM, NY, S). Pele, Duss 297 (NY).
SyrrhopodonrhizogonioidesC. Miiller, Hedwigia39: COLOMBIA. ANTIOQUIA: Mun. Guatap6, Sastre-de
266. 1900. Type. Brazil. Minas Gerais: Caraca, Ule Jesus et al. 1275 (HUA, LAF, NY). BOYACA:Mun.
1389 (lectotype, here designated, NY; isolectotype, H). Duitama, Ireland 23566 (CANM, LAF). MAGDALENA:
Calymperaceae 47

ParqueNacionalde la Sierrade SantaMarta,Griffin,11 ventrally,mostly ca. 8-11 pm;gemmae not seen.

etal. 50096 (FLAS,LAF,U). VALLE:Mun.Cali, 17 km Seta brown, 7-9 mm long; capsule 2-2.5 mm
NO de Call, Churchill et al. 15151 (CUVC, LAF, NY).
VENEZUELA. AMAZONAS: Dpto. Rio Negro, 2.8 km long;peristomeof erodedfragments;operculum
NE Pico Phelps, Buck 12618 (NY). LARA:[without local- very slenderly rostrate, 1.5 mm long. Spores
ity] Steyermark55264 (NY). PORTUGUESA: Distr. Sucre, granular, 17-19 pm. Calyptra 3.5 mm long,
LaDivisoriade la Concepci6n,vander Werffetal. 7626A
Distr. Bocono, Griffin, 111et al. 1099 roughenedat tip.
(MO). TRUJILLO: Illustrations. Magill (1981, fig. 46: 22-29, as
(FLAS, LAF). ZULIA:near San Jose de los Altos, Griffin,
III 225 (FLAS, LAF). S. gomesii P. Varde);Reese (1977, figs. 51-52,
GUYANA. UpperMazaruniDistr.,N slope of mount as S. quintasii Broth.); Reese et al. (1986, figs.
Roraima,Gradstein5329 (LAF,U). 4142).
ECUADOR.CARCHI: SW slope of Volcainde Chiles, Distribution. (Fig. 35A). Pantropical.Mexico
CordilleraOccidental, Steere 9053 (LAF, NY). IM-
BABURA:W slope of Cordillera Occidental, San Pablo, (Chiapas, Jalisco, Oaxaca, Tamaulipas, Vera-
Steere 9237 (LAF, NY). LOJA:SE of Loja, Laegaard cruz), CentralAmerica (Costa Rica), Colombia
53628C (LAF, NY). NAPO-PASTAZA: valley of Rio (Meta, Norte de Santander,Vaupes), Trinidad,
Chingual E of El Pun, Steere 8419 (LAF, NY). ORIENTE: Brazil (Amazonas,Bahia, Goias, Mato Grosso,
Rio Pastaza near Shell-Mera, Steere 9378 (LAF, NY).
BRAZIL. MINASGERAIS:The type of Syrrhopodon
Minas Gerais), Bolivia (Sta. Cruz). Trees, soil,
rhizogonioidesC. Mill. rock in forests,at moderateto high elevations.
BOLIVIA.Bei Tablas,Herzog4542 (JE,L, NY, S).
Selectedspecimens examined. MEXICO.CHIAPAS:
slope NE of Cerro
Venado, Breedlove67551
(CAS,LAF).
Discussion. Syrrhopodonlycopodioides,with JALISCO: 10 mi SW of Autlan, Crum 1342 (MICH). OA-
its long, narrow,prominentlytoothedandheavily XACA: 12 mi. aboveValle Nacionalon Hwy. 175, Sharp
borderedleaves, is unmistakable.It has no close et al. 4522-a (LAF, TENN). TAMAULIPAS: Rancho del
relatives anywhere in the world and cannot be Cielo, above G6mez Farias,Smithet al. 172A (LAF,
confused with any otherspecies of Syrrhopodon. TENN).
COSTA RICA. HEREDiA: Vic. of Vara Blanca, Croat
20. Syrrhopodon gardneri (Hooker)Schwaeg- 35538c (MO).
COLOMBIA. META: Cordillera Oriental, W of
richen, Spec. musc. frond. Suppl. 2(1): 110. Villavicencio, Steere 7655 (LAF, NY). NORTEDESANTAN-
1824. Fig. 33. DER: CordilleraOriental,E of Rio Valegra,Steere 7354
(LAF, NY). VAUPES:Rio Apaporis, Schultes 11794
Calymiperes gardneriHooker,Musciexot. 2: 146. 1819. (FLAS,LAF).
Type.Nepal.Hon.D. Gardners.n.,comm.Dr.Wallich TRINIDAD.Mt.Tamana,18 Apr 1920,N. L Britton
(BM). et al. s.n. (NY).
Syrrhopodon quintasiiBrotherus,Bol. Soc. Broteriana
8: BRAZIL. BAHIA:Mun Rio deContas, Harley 27622 (K,
178. 1890.Type. Africa.Sao Tom6Island:F. Quintas LAF).GOIAs: Alto Paraiso,Chapadados Veadeiros,Can-
s.n. (holotype,H-BR,n.v.; isotype,NY). yon des Rio Preto,Schafer-9892(herb.Schfifer-Verwimp,
Syrrhopodon hobsonii(Greville)Hooker& Grevillevar. LAF).MATO GROSSO: Gorgeof VeudeNoiva,Chapada dos
spinulosusHerzog,Beih. Bot. Centralbl.26: 69. 1909, Guimaries,Pranceet al 19098(INPA,LAF,NY). MINAS
nomn.nud. Originalcollection.Bolivia. An schattigen GERAIS: Sera da Nantiqueira,
S. Tomedasletras,Schdfer-
SandsteinfelsendesCerrode Santiago(Chiquitos),May Verwimp 6806(herb.Schifer-Verwimp, LAF).
1907, Herzogs.n. (JE,L). BOLIVIA. The types of Syrrhopodonciliolatus
Syrrhopodonciliolatus Herzog,Biblioth.Bot. 87: 166. HerzogandS. hobsoniivar.spinulosusHerzog.
1916, hom. illeg., non S. ciliolatus Geheeb & Hampe in
Hampe. Type. Bolivia. Im Bergwald,ohne nahere Discussion. At first glance S. gardneri may
Fundortsnotiz,
Herzog4203/a (JE). look much like S. incompletusvar. incompletus,
but the sharp,spreadingteeth on the shoulders,
Plants slender to robust,in tufts and cushions,
dorsallypapillosemediancells, and usually con-
green above, brown or blackish below; rhizoids spicuousdark-redto purplerhizoidsarediagnos-
dark-redto purple; stems to 4 cm tall. Leaves tic. This pantropicalspecies has been known in
fragile,contortedwhen dry,4-5 mm long, linear- the Americasas S. quintasiiin recentyears.
lanceolate from broader base; margins thickened,
coarsely toothed,at shoulderswith sharpspread- 21a. Syrrhopodon incompletus Schwaegrichen
ing or ascending teeth; cancellinae distinct, var. incompletus Fig. 34.
sharply demarcated distally from adjacentgreen SyrrhopodonincompletusSchwaegrichen,Spec. muse.
cells; median cells of upper lamina square, frond.Suppl.2(1): 119. 1824.Type.Cuba.Poeppigs.n.
pluripapillose dorsally, mammillose-papillose (holotype,G, n.v.;isotypes,BM, JE,PC).
48 Flora Neotropica

32 . 3
,33 o

Ai %, i

d
A
\ /A 1 ^

*
~I 'i)0000
0~0 oO

FIGS. 32-34. Syrrhopodon. 32. S. lycopodioides. A. Leaf outlines. B. Leaf apex. A from Standley 38160b (JE) (two
leaves at left), Ule 1389 (NY) (leaf at right), B from Ule 1389 (NY). 33. S. gardneri. A. Leaf outline. B. Margin and
cells at leaf shoulder. A from Herzog s.n. (L), B from Herzog 4203a (JE). 34. S. incompletus var. incompletus. A. Leaf
outlines. B. Leaf apex. C. Margin and cells at leaf shoulder. A from Florschutz & Maas 2328 (U) (left), Stoffers 3698
(U) (right), B-C from Florschutz & Maas 2328 (U). Scale bars: a = 0.1 mm (Figs. 32B, 34B); b = 1 mm (Fig. 32A);
c = 1 mm (Figs. 33A, 34A); d = 0.1 mm (Fig. 33B); e = 0.1 mm (Fig. 34C).

Calymperes hobsonii Greville, Ann. Lyceum Nat. Hist. SyrrhopodonsemicompletusSchwaegrichen, Spec. Musc.
New York 1(2): 271. 1825; Syrrhopodonhobsonii (Gre- Suppl. 2(2): 97. 1827, nom. illeg. incl. type spec. prior.
ville) Hooker & Greville, Edinburgh J. Sci. 3: 224. Syrrhopodonfloridanus Sullivant in Gray, Man. Bot. ed.
1825. Type. Guiana. Hobson s.n. (holotype, BM-K; 2, 631. 1856. Type. U.S.A. Florida: Apalachicola,
isotype, NY). Drummnonds.n.,Musci Americani (Souther States) 37
Calymperaceae 49

TROPICALAMERICA For Neotopcabase apno1

..... ,--------- -
---------------e

...~ . ?
......................... .~
...........

'
i ,* f xc " 1 X e w ,c

'~ry ~i --..---~ ----

FIG. 35. Distributions of American Syrrhopodon. A. S. gardneri (continuous lines); S. rupestris (dashed line). B.
S. incompletus var. incompletus (continuous lines); S. incompletus var. luridus (dashed lines); S. incompletus var.
perangustifolius (dots).
50 FloraNeotropica

(as S. albovaginatusSchwaegr.),(holotype,FH?,n.v.; on costa at leaf tips. Seta yellowish-brown,7-12

isotypes,BR, NY, REN). mm long; capsule 1.5-2.5 mm long; peristome
SyrrhopodonmohriiC. Miiller,Linnaea38: 633. 1874.
Type. Mexico. Cordoba,1857, Mohrs.n. (ectotype, lacking or rudimentaryand of fragile, eroded
heredesignated,NY; isolectotypes,BM, MICH). teeth of only a few segments; operculum slen-
Calymperesglaziovii Hampe,Vidensk Meddel.Dansk derly long-rostrate.Spores 14-19 pm, nearly
Naturhist.Foren.Kj0benhavn3,9-10:252.1877.Type. smooth to finely roughened. Calyptra 2.5-3.5
Brazil.in viciniis Rio de Janeiro,Glaziou7449 (holo-
mm long, smooth.
type, BM?,n.v.; isotypes,G, S).
Syrrhopodonbernoullii C. Miiller,Bull.Herb.Boissier5: Selected illustrations. Bartram(1949, fig. 34:
189. 1897. Type. Guatemala.Mazatenango,Bernoulli G-I); CrumandAnderson(1981, fig. 109);Crum
& Cario 134 (lectotype, here designated, H-BR; andSteere(1957, fig. 17a);Florschiitz(1964, fig.
isolectotype,GOET). 62: i-j); Reese (1977, figs. 1-4).
SyrrhopodonbrachystelioidesC. Miiller,Nuovo Giom.
Bot. Ital. n. ser. 4: 48. 1897. Type. Bolivia. Distribution. (Fig. 35B). Throughouttropical
Cochabamba, propeChoquecamata, Jun1889,Germain and subtropicalAmerica,including the southern
s.n. (lectotype,heredesignated,JE;isolectotypes,NY, coastal plain of southeasternUnited States; Af-
PC, S). rica. Tree trunks,logs, humus, especially com-
Syrrhopodon decoloransC. Miiller,Bull. Herb.Boissier
5: 188. 1897. Type.Guatemala.Bernoulli& Cario89 mon on palm stems, in forests, to ca. 1700 m,
(lectotype,here designated,GOET;isolectotypes,H- most commonat lower elevations.
BR, NY, PC).
Syrrhopodonbrotheri C. Miiller,Hedwigia39:266.1900. Selected specimens examined. MEXICO. CAMPE-
Type.Brazil.Goias:Pr.Massamedes,Ule 1542 (holo- CHE:4 km W of Escircega, Davidse et al. 20297 (LAF,
type, H-BR;isotype,NY). MO). CHIAPAS:9 kmN of Ocozocoautla, Breedlove67160
Syrrhopodonjuruensis Brotherus,Hedwigia45: 269. (CAS, LAF). JALISCO:Near Sapotillo, Crum 620 (MICH,
1906. Type. Brazil.Amazonas:Jurui,Bocca de Tejo, NY). NAYARIT:ca. 6 mi E of San Bias, Norris & Taranto
Ule s.n., Bryothecabrasiliensis253 (holotype,H-BR; 13359 (NY). OAXACA:12 mi. above Valle Nacional on
isotypes,JE, L, NY, REN,S). Hwy. 175, Sharp et aL 3131 (LAF, TENN). PUEBLA: El
SyrrhopodonsubdecoloransBrotherus,Hedwigia 45: Cerro de Cuhuatepec Sharp 1376a (MICH, TENN).
269. 1906. Type. Brazil.Amazonas:JuruaiMiry,Rio QUINTANAROO:25 km N de Tomas Garrido, Delgadillo
Jurua,Ule 2271 (holotype,H- BR;isotype,S). M. 4303 (MEXU, NY). SANLUISPoTOSi:El Sal, Sharp
CalymperesarmatumBrotherus,Akad. Wiss. Wien, 1657 (TENN). TABASCO: Mun. del Centro, Dos Montes,
Math.-Naturwiss. K1.,Denkschr.83: 282. 1926.Type. Delgadillo M. 1397 (MEXU, NY). TAMAULIPAS:Rancho
Brazil.Sao Paulo:Pr.RioGrande,Schiffner883(lecto- del Cielo, Sharp 8680-b (LAF, TENN). VERACRUZ:1.2 mi
type,heredesignated,H-BR).Syntype.Brazil.Pr.San- S of Rio Tecolutla onroad to Nautla, Reese & Pursell4619
tos, Schiffner910 (H-BR). (LAF, US). YUCATAN:11 mi E of Valladolid, McFarland
Syrrhopodon hioramiiBizot & Theriotin Th6riot,Mem. & Sharp 9285 (LAF, TENN).
Soc. Cub.Hist.Nat."FelipePoey"13:268. 1939.Type. GUATEMALA. BAJA VERAPAZ:Civija, Sharp 5212,
Cuba.LomaSanJuan,Hioram11809 (holotype,PC?, 5229 (NY, TENN). PETEN:Fallab6n-Yaxha road, Lundell
n.v.; isotype, MICH). 2035 (MICH).
Calynperes meyeri Reese, Bryologist60: 336. 1957. BELIZE. British Honduras-Guatemala Border Sur-
Type. Panama.Canal Zone: BarroColoradoIsland, vey, Camp 34, 6 May 1934, Schipp s.n. (LAF, NY). EL
Meyer7 (TENN). CAYO:Bartlett 12085 (NY). TOLEDODISTRICT:Near Or-
ange Point, Gentle 7476 (MO).
Plants dark-greento brownish, tufted or gre- HONDURAS. ATLANTIDA:Tela, Steyermark54284 (F).
NICARAGUA. Rio Siguia, 6 Jan 1941, La Rue s.n.
garious; rhizoids brown to purple; stems to 2-3
(NY).
cm tall but often much shorter, often freely COSTA RICA. ALEJUELA: Los Angeles de San
forked. Leaves basically monomorphic,lanceo- Ramon, Brenes 17114 (NY). LIM6N: near Tortuguero,
late to linear-lanceolateor nearly subulatefrom Steere CR-120 (LAF, NY).
PANAMA. CANALZONE:Barro Colorado Island,
broad, clasping base, 4-8 mm long, variously 3938
curved-contortedto straightand appressedwhen M-1 (LAF, (LAF,
Crosby MO). DARItN: Rio Espare, Gentry
MO). PANAMA:upper Bayano near Maje, 25
dry;marginsthickened,with mostly pairedteeth Mar 1969, CorreaA. s.n. (LAF, MO).
above; leaf shoulders often with intramarginal BERMUDA. Paget Marsh, E. G. Britton 1856 (LAF,
teniola-like features;cancellinaepersistent,hya- NY).
line (rarely yellowish); cells of upper lamina BAHAMAS ARCHIPELAGO. GRAND BAHAMA:
Lucaya National Park, Reese 18000 (DUKE, LAF, NY,
obscure, quadrateto elliptical, mostly ca. 5-7 x
U).
5-12 pm, smooth or somewhat papillose dor- CUBA. ISLE OF PINES: Sierra de la Casas, N. L Britton
sally, bulging ventrally; gemmae borne ventrally & Wilson15766 (NY). ORIENTE:Moa, Jervis 1687 (NY).
Calymperaceae51

JAMAICA. ST. THOMAS: S slope of Winchester Peak, Stroessner, Buck 12364 (NY). Nova Germania, Wassan,
Crosby 13706 (NY). 1 Oct 1896, Tetrepliscz s.n. (NY). PARAGUARI: Parque
HAITI. Savanne Zombi, Mornes de Commissaires, Nac. Ybycui, Buck 11982 (NY).
Mackaness 94a (MICH). ARGENTINA. MISIONES: Dept. Iguazi,
DOMINICAN REPUBLIC. LA VEGA:ESE of Bonao, Naturschutzparkdes Rio Iguazu, Hosseus 213 (JE).
Zanoni etal. 15788 (NY).
PUERTO RICO. CaribbeanNational Forest, Luquillo Discussion. This is a weedy moss. Itsplantsare
Division, El Verde Biological Station, Reese 14283 (LAF,
NY). dimorphic and occur in strict- and flexuous-
LEEWARD ISLANDS. MONTSERRAT: Shafer 807 leavedformsthatmay appearstrikinglydifferent
(NY); ST.EUSTATIUS:The Quill, Florschiitz3207(LAF, U). from one another.However, under the micro-
WINDWARD ISLANDS. GUADELOUPE: Ile Saint-
scope the leaves of the two forms are indistin-
Barthelemy, Le Gallo 1035a (NY). MARTINIQUE:Riviere The leaf bases of both sometimes bear
du Prehaux, Duss 368 (H). ST. LUCIA:Basse de l'Isle, S guishable.
of Piton Flore, Simmonds 194a (NY). teniola-like features at the shoulders. Variety in-
SOUTHERN DUTCH ANTILLES. CURACAO: completus differs from var. berteroanus in hav-
Christoffelley, Hummelinck 15124 (BM). ing mostly paired teeth on the upper margins and
COLOMBIA. ANTIOQUIA:Mun. Maceo, 5 km NNE de the
upper margin usually lanceolate rather than
Maceo, Churchill et al. 14351 (NY). CAQUETA:Cerca de the two forms in-
Rio Pato, 2?30-33'N, 74'45'W, Churchill & Betancur commonly subulate; however,
16980 (LAF, NY). MAGDALENA: Sierra Nevada de Santa tergrade. The var. incompletuis differs from S.
Marta, Rangel & van Reenen 1135 (FLAS, LAF). META: circinatus and S. rigidus, among other ways, in
Rio Gianirare,Schultes 12007 (FLAS, LAF). SANTANDER: its monomorphic leaves and persistent, hyaline
NW of Bucaramanga, Steere 7643 (NY).
VENEZUELA. AMAZONAS:
(but sometimes yellowish) cancellinae.
Dpto. Rio Negro, Along is similar but
Rio Mawarinuma, Buck 12559 (NY). ARAGUA:Parque Syrrhopodongardneri superficially
Nacional "Henry Pittier," Sharp 9929 (LAF, TENN). differs prominently in have sharp spreading teeth
SUCRE:Rio Sabacual, Griffin, III & Bermddez 021326 at the leaf shoulders, dark-red rhizoids, and
(FLAS, LAF). sharply papillose leaf cells.
TRINIDAD. ST. ANDREW: ca. 2 mi. E of Valencia,
Crosby 2339 (LAF, MO).
GUYANA. NW slopes of Karuku Mtns., Smith 3565 21b. Syrrhopodon incompletus var.
(FH). berteroanus (Bridel) Reese, Bryologist 80: 9.
SURINAME. Raleighvallen, J. & P. Florschiitz 4540 1977. Fig. 36.
(LAF, U).
FRENCH GUIANA. ST-LAURENT-DU-MARONI:Can- Dicranumberteroanum Bridel, Bryol. univ. 1: 445. 1826;
ton de Maripasoula, ca. 6 km N of Saul, 200 m, Buck18392 Orthotheca berteroana (Bridel) Bridel, Bryol. univ. 1:
(CAY, NY). 778. 1827; Syrrhopodon berteroanus (Bridel) C.
ECUADOR. GALAPAGOS ISLANDS: DuncanIsland,Stew- Muller, Syn. musc. frond. 1: 539. 1849; Syrrhopodon
art 3165 (NY). NAPO:ca. 75 km E of Puerto Franciscode berteroiKindberg,Enum.Bryin.exot. 76. 1888,nom.
Orellana,Churchill & Sastre-deJesus 13801 (LAF, NY). illeg. incl. spec. prior.Type. PortoRico. Berteros.n.
PERU. LORETO: Maynas, between Rio Napo and Rio (holotype, B-Bridel; isotypes, JE, NY).
Amazona, Timme4845 (herb. Timme, LAF). SANMARTN: Syrrhopodonmalouinensis C. Miiller, Syn. musc. frond.
km 95, Strasse Yurimaguas-Tarapoto, Frahm et al. 1634 1: 539. 1849. Type. Insulae Malouinae. Dumont
(LAF, U). d'Urville s.n. (B).
BRAZIL. ACRE:15 km E of Rio Branco, Reese & Syrrhopodonberteroanus var. strictior Hampe, Linnaea
McPherson 13160 (INPA, LAF, MO, NY). AMAZONAS: 25: 360. 1853, nom. nud.
Cano Tucano, Rio Cauaburi, Maguire et al. 60208 (NY). Syrrhopodonlaevidorsus Bescherelle, Rev. Bryol. 18: 75.
MATOGROSSO: Mun. de Pedro Gomes, Vital6596 (FLAS, 1891. Type. Guadeloupe. Marie 754 (holotype, BM;
LAF, SP). MATO GROSSODO SUL: 70 km ESE de isotypes, H-BR, NY).
Corumba, Vital 8615 (NY, SP). MINASGERAIS: [Without Syrrhopodon vaginans Brotherus, Bih. Kongl. Svenska
data] 1890, Rudolph s.n. (H). PARA:Serra do Cachimbo, Vetensk.-Akad. Handl.21 Afd. 3(3): 20. 1895. Type.
km 777 on Cuiba-Santaremhighway, Reese 160 71 (INPA, Brazil.Santos,Mosen46 (ectotype,H-BR).
LAF, MO, NY). ROND6NIA: 2-4 km above the first rapids SyrrhopodoncurvatusC.Miller, Hedwigia39: 267.1900.
on Rio Pacaas Novos, Reese 13442 (INPA, LAF, MO, Brazil.Sa. Catarina:Blumenau,Ule 534 (lectotype,
NY). RORAIMA: 216 km N of Boa Vista, Buck et al. 2020 NY;isolectotype,H-BR).
(INPA, LAF, NY). SANTACATARINA: Sao Francisco do Syrrhopodonperhorridus C. Miller, Hedwigia 39: 267.
Sul, Yano 2373 (LAF, SP). SAo PAULO: Ilha do Cardoso, 1900. Type. Brazil. Sa. Catarina: Nova Venezia, Ule
Vital 10297 (LAF, SP). 1133 (lectotype, H-BR).
BOLIVIA. BENI: vic. of Guyaramenrn,Reese 12931 Syrrhopodonberteroanus var. planifolius Theriot, Mem.
(INPA, LAF, MO, NY, US). Soc. Cub. Hist. Nat. 'FelipePoey" 13: 268. 1939. Type.
PARAGUAY. ALTAPARANA: 4.5 km W of Pto. Pte. Cuba. Loma San Juan, Hioram 13701 (PC).
52 Flora Neotropica

Plants of var. berteroanusare dimorphicwith WINDWARD ISLANDS. DOMINICA: Laudat, Lloyd

381 (NY). GUADELOUPE: Sofia, Questel3483 (NY). MAR-
strict- and flexuous-leaved forms. They differ
TINIQUE:[without locality] M. & H. Stehle 3869 (NY). ST.
from var. incompletus primarily in the leaves LUCIA:Zwischen Castries und Dennery, E. & P.
mostly narrower(commonly subulate)and with Hegewald 9611 (herb. Hegewald, LAF). ST. VINCENT:
more broadly flaring bases, and in bearingsin- Mariaqua Valley, Elliott 20 (BM). GRENADA:Grand
gle-rather than paired-marginal teeth above; Etang, Richards 948 (NY).
COLOMBIA. MAGDALENA: Parque Nacional de la Si-
the margins of the upper lamina are sometimes erra Nevadade Santa Marta,Griffn, 111et al. 50214
unistratosein part. The sporophytesare as in S. (FLAS,LAF). NARINO: ReservaNaturalde La Planada,
incompletus var. incompletus. The var. Ramirez2061 (LAF,NY, PSO).
berteroanusintergradesto some extent with var. VENEZUELA.ZULIA: Sierrade Perija,nearSanJose
de Los Altos,Griffin,11120(FLAS,LAF,NY).
incompletusandit is not alwayspossible to assign TRINIDAD.ST.GEORGE: Crosby2167 (LAF,MO).
a varietalname with ease. However, most speci- TOBAGO.Moriah,24 Feb 1892, Weisss.n. (NY).
mens arereadilyidentifiableto the varietallevel. ECUADOR:Los Rios:HaciendaClementina,Harling
Illustration. Reese (1977, figs. 8-10). 2152 (NY). PASTAZA:
2?22'S, 76?40'W, 0llgaard et al.
Distribution. (Fig. 41A). Endemic to tropi- 35289 (AAU, NY). PICHINCHA: 16 km E of Santo Do-
cal America; Mexico, Central America, Carib- mingo,Steere& Balslev25979 (NY).
PERU. LORETO: Maynas, 12 In SW of Iquitos airport,
bean, northern South America and disjunct to Timme4770 (herb. Timme, LAF).
southernBrazil. Tree trunksand logs in forests BRAZIL. AMAPA:Mun. Oiapoque, Mori & Cardoso
to about 1700 m; more abundantat higher ele- 17124 (NY). PARANA: Serrado Prada,24 Aug 1910, Dusen
s.n. (US). Rio DEJANEIRO:
[without locality] Glaziou 9278
vations.
(NY). SANTACATARINA: Tubarao, Ule 951 (NY). SAo
PAULO:Ilha do Cardoso, Vital10507(LAF, SP).
Selected specimens examined. MEXICO. OAXACA:
Near Niltepec Sharp 5764 (MICH, TENN). Discussion. Althoughgenerallysimilarto var.
GUATEMALA. IZABEL: Near Entre Rios, Standley
72778 (F, MICH, NY). incompletus, var. berteroanus is usually easily
BELIZE. Stann Creek Valley, Mountain Cow Ridge, distinguishedby the single teeth on the leaf mar-
Gentle 3279 (LAF, NY). gins, but the two varieties intergrade.Like var.
NICARAGUA. BLUEFIELDS: 3.6 km SE Cerro San
incompletus, its leaves often bear teniola-like
Isidro, Proctor et al. 27197 (NY). ZELAYA:near Rama, featuresat theshoulders.See commentsunderthe
Kofron 3a (LAF).
COSTA RICA. ALAJUELA: between Caiias and Upala, var. luridus. The apparent occurrence of this
Croat 36315 (LAF, MO). Cocos ISLAND:[without local- taxon in the Islas Malvinas, based on S.
ity] Holdridge 5196 (NY). HEREDIA:La Virgen, Finca la malouinensis,is anomalousand is perhapsbased
Tigra, 4 Dec 1982, Young s.n. (LAF, UWSP). LIM6N: on a mislabeledspecimen. The var. berteroanus
Cerro Coronel, Stevens 23744 (LAF, MO). PUNTARENAS:
E of Golfito, Crosby 3711 (LAF, MO). SAN JOSE:La has not been collected in the Malvinas since the
Honduria, Standley 37836 (NY). originalspecimen upon which Muller based the
PANAMA. BOCASDELTORO:Vic. FortunaDam, Allen name malouinensis.
5743 (NY). CHIRIQUI: SW de Campamento de Fortuna,
Salazar Allen et al. (NY). CocLt: La Mesa, Tyson 2472L
21c. Syrrhopodon incompletus var. luridus
(LAF, NY). COL6N:near mouth of Rio Guanche, Crosby
10786 (LAF, MO). DARI?N: between Cerro Pirre and Piji (Paris& Brotherus)Florschiitz,Mosses of Sur-
Vasal, Folsom 6392 (LAF, MO). PANAMA:Cerro Camp- iname 1: 163. 1964. Fig. 37.
ana, Crosby 10910. VERAGUAS:below CerroTute, Crosby
10281 (LAF, MO). Syrrhopodon luridusParis& Brotherus,Rev. Bryol.33:
CUBA. ORIENTE:La Prenda, Hioram & Maurel 2608 56. 1906;Syrrhopodon hobsonii(Greville)Hooker&
(NY). Greville var. luridus (Paris & Brotherus) P. Richards,
JAMAICA. PORTLAND:ca. 4 mi. N of Bath, Crosby Bull. Misc. Inform. 8: 323. 1934. Type. French Guiana.
2970 (LAF, MO). ST. THOMAS: Ca. 4.5 mi. N by W of Michels.n. (holotype, REN; isotypes, L, M, NY, PC).
Bath, Crosby 3282 (LAF, MO).
HAITI. Massif du Nord, Port-de-Paix, Haut Piton, The varietyluridusis closely relatedto variety
Ekman 4863 (NY). berteroanusbut differs in having mostly paired
DOMINICAN REPUBLIC. Samana Peninsula, vic. teeth on the marginsof the upperlamina. Other
of Laguna, Abbott 328 (NY). distinctive featuresinclude the occasional pres-
PUERTO RICO. Caribbean National Forest, Luquillo
Division, El Verde, Reese 14310 (LAF, NY).
ence of elongatecells forminganirregularborder
LEEWARD ISLANDS. SABA: Mount Scenery, van on the marginsof the lower lamina, nearly uni-
Slageren 246M (NY, U). form presence of teniola-like features in the
Calymperaceae 53

36 Io
A' 37

o
c), roo c
oo_0, 00

opO

a
b
c
d

/41 ~ ~

FIGS. 3-8 Syhpdn3.Sino ltsvr o,

ouO'VU C

FIGS. 36-38. Syrrhopodon. 36. S. incompletus var. berteroanus. A. Outline of leaf. B. Portion of upper lamina:
costa at left, margin at right, showing single teeth. C. Marginand cells at upperpartof leaf shoulder. A, C from Schiffner
s.n. (BM), B from Spruce 5 (NY). 37. S. incompletus var. luridus. A. Leaf outline. B. Leaf base. C. Margin and cells
at upper part of leaf shoulder. From Michel s.n. (holotype of Syrrhopodon luridus Par. & Broth., REN). 38. S.
incompletus var. lanceolatus. A. Leaf outlines. B. Margin and cells at leaf shoulder. From Glaziou 9278 (isotype of
Calymperes lanceolatum Hampe, H). Scale bars: a = 1 mm (Figs. 36A, 37A, 37B, 38A); b = 0.1 mm (Fig. 38B); c =
0.1 mm (Figs. 36C, 37C); d = 0.5 mm (Fig. 36B).

lower lamina, very strongly flaring shoulders, FrenchGuiana;Brazil (Amazonas, Roraima).On tree
short cancellinae, ventrally smooth supra- trunksin forests at low elevations;not common.
cancellinarcells, elongatemediananduppercells Selected specimens examined. COLOMBIA. AMA-
of the upperlamina, and dorsally smooth costa. ZONAS:Ca. 7 km norte de Leticia, Churchill et al. 16180,
Sporophytesas in var. incompletus. 16192 (LAF, NY).
Illustrations. Florschiitz (1964, fig. 62 a-g); GUYANA. Essequibo River, Moraballi Creek near
Reese (1977, figs. 5-7). Bartica, Richards 449 (BM, NY).
SURINAME. Sarakreek,above Drie Gebroeders, J. &
Distribution. (Fig. 35B). Endemicto northern P. Florschitz 266 (LAF, U).
South America. Colombia; Guyana; Suriname; FRENCH GUIANA. A 3 km de St. Jean du Maroni,
54 FloraNeotropica

1906, Rey s.n. (NY, PC). ST-LAURENT-DU-MARONI: Can-

teeth to 80 um tall; the spores are green and
ton de Maripasoula, ca. 6 km N of Safil, 200-300 m, Buck
18330,18722 (CAY, NY). granularand average23 um diam.
BRAZIL. AMAZONAS: Reserva Florestal, km 43 along Illustrations. Florschiitz(1964, fig. 46); Reese
BR 174, Bucket al. 1773 (INPA,LAF, NY). RORAIMA:(1977, figs. 14-16).
km328alongBR174,Bucketal.1867(INPA, LAF,NY). Distribution. (Fig. 41B). A rare endemic of
South America.Colombia,Guyana,FrenchGui-
Discussion. Plants of the var. luridusare mac-
ana, Suriname,Brazil (Amazonas, Bahia, Mato
roscopically similar to those of the flexuous- Grosso, Para, Rio de Janeiro, Roraima, Santa
leaved forms of vars. berteroanus and
Catarina,Sao Paulo);on tree trunks.
incompletus. The var. luridus differs from var.
berteroanusin havingdoublerows of teethalong Specimens examined. COLOMBIA. AMAZONAS: Sur
the margins of the upper lamina, ratherthan a estedela pistadepespegue(estede LaPedrera),Churchill
etal. 16103 (LAF,NY).
single row of teeth on each margin, and differs GUYANA.UpperMazaruniRiver,Leng364 (NY).
fromvar. incompletusby a combinationof char- SURINAME. Lawa River, opposite Gransanti,
acters including its oval to rounded-rectangular Geijskes195 (LAF,U).
cells in the upper lamina, uniformly strongly FRENCH GUIANA. ST-LAURENT-DU-MARONI: Can-
tonde Maripasoula,ca. 6 kmN of Saiil,200-400 m, Buck
flaring leaf shoulders, common presence of ten- 18340 (NY), 18843 (CAY,NY), 18955(NY).
iolae-like features,and smooth supracancellinar BRAZIL.PARANA: SerradoPrata,24 Aug1910,Dusen
cells. The distinctions between var. luridusand s.n. (NY). SANTACATARINA: Matado Hoffman, Reitz3148
forms of vars. incompletus and berteroanusare (FH,LAF).
not always easy to distinguish. In a general Discussion. The var. lanceolatusis ratherrare
way, the var. luridus can be thought of as a andpoorly collected. It was originallydescribed
version of the var. berteroanus with its leaf in
Calymperesfrom nonfruitingplants but was
margins doubly toothed;it is confined to north- transferredto Syrrhopodon Reese (1977) be-
ern South America. by
cause of the general similarity of its plants to
21d. Syrrhopodon incompletus var. those of S. incompletusvar. incompletus.Recent
lanceolatus (Hampe)Reese, Bryologist80:13. discoveryof the sporophytesof var. lanceolatus
1977. Fig. 38. (Reese & Buck 1991) clearly demonstrateits
relationshipto the S. incompletus complex of
Calympereslanceolatum Hampe, Vidensk. Meddel. taxa.Underthe microscopeits leaves are gener-
DanskNaturhist. Foren.Kjobenhavn, ser4, 1:78. 1879.
similarto those of the var. incompletus
Type.Brazil.Rio de Janeiro:Corcovado,Glaziou9278 ally very
(holotype,BM;isotypes,H, PC). but differ obviously by the strongly bordered
SyrrhopodonkroneanusC. Miller, Hedwigia39: 265. marginsof the lower laminae.Macroscopically,
1900. Type. Brazil.Sao Paulo:Krones.n. (lectotype, the
here glossy julaceous stems are characteristic.
designated,NY).
21e. Syrrhopodon incompletus var. per-
Thisvarietyis characterizedby havingthemar-
angustifolius Reese, Bryologist 80: 13. 1977.
gins of the lower lamina strongly borderedby Type.Peru.Guayapurina,Spruce5 (NY). Para-
elongate, thick-walled, somewhat sinuose and type. Ecuador. Oriente: Bamboiza, 23 Sep
often pitted cells with pointed ends. The cells of
1910, Allionis. n. (BM). Fig. 39.
the upperlamina are smooth dorsallyand some-
what elongate, the lower lamina flares at the Varietyperangustifoliusdiffers from its close
shoulders, and the upperlaminais lanceolateto relative-var. lanceolatus-in havingspreading,
linear-lanceolatewith its marginscoarsely dou- more or less flexuous leaves with the upperlam-
bly serrateabove and showing few or no stereids ina very narrowandits marginsin section show-
in section. When dry, the plants are characteris- ing several to many stereids,and in the strongly
tically glossy, andratherjulaceousbecauseof the flaringshouldersof the lower lamina.The sporo-
appressedleaves. The sporophytesof the single phyte lacks a peristome.
fertile specimen known (French Guiana, Buck Illustration. Reese (1977, figs. 17-20).
18843, NY) are similar to those of the var. in- Distribution. (Fig. 35B). Ecuador and Peru.
completus, with the peristome of pale, smooth, Very rare; known only from the type material
imperfect,colorless,segmented,connivent,blunt cited above. On tree trunks.
Calymperaceae 55

B
A O

39

C C,.

10o

40 000
9
OOoOo
000
uoooOooO

0c30)

FIGS. 39-40. Syrrhopodon. 39. S. incompletus var. perangustifolius. A. Leaf outlines. B. Portion of leaf base;
intramarginalline indicates width of the band of differentiated teniola-like cells. C. Margin and cells at leaf shoulder.
A (leaf at left), C from Allioni s.n. (paratype of Syrrhopodon incompletus var. perangustifolius, BM), A (leaf at right),
B from Spruce 5 (holotype of Syrrhopodon incompletus var. perangustifolius, NY). 40. S. elatus. A. Leaf outline. B.
Margin and cells at upper part of leaf shoulder. C. Portion of leaf section. From Florschutz & Florschiitz 1813 (U).
Scale bars: a = 1 mm (Figs. 39A, 40A); b = 0.1 mm (Figs. 39C, 40B-C); c = 0.5 mm (Fig. 39B).

22. Syrrhopodon elatus Montagne, Ann. Sci. branched.Leavesnarrowlylanceolateabove flar-

Nat. Bot. 2,3: 198. 1835. Type. French Guiana. ing base, 3-4 mm long, more or less appressed-
Leprieur s.n. (PC-MO). Fig. 40. secund but not otherwise contorted when dry,
ascendingwhen moist; marginsof upper lamina
SyrrhopodonincompletusSchwaegrichen var. elatus with stout,
(Montagne)Florschiitz,Mosses of Suriname1: 163. pairedteeth, showing stereids in sec-
1964. tion, marginsof lower laminaentire,or serrulate-
toothed above shoulders; teniola-like features
Plants a little glossy; rhizoids reddish-brown; lacking; cancellinae rounded distally, ending
stems to 4.5 cm tall but mostly shorter, not much well below shoulders,distalcells often with char-
56 FloraNeotropica

TROPICAL AMERICA FloraNeoropca base map no 1

BI

FI G . - 41 itiuin fAeia yrooo.A .icmltsvr etrau.B

.icmltsvt
lance tus._;L ol a

FIG. 41. Distributions of American Syrrhopodon. A. S. incompletus var. berteroanus. B. S. incompletus var.
lanceolatus.
Calymperaceae57

acteristicbrown pigment;median leaf cells pel- (lectotype,heredesignated,H-BR).Syntypes.Ecuador.

lucid, thick-walled,ca 12 pimdiam., smooth and Gualaquiza,fl. Yumazo,Allionis.n., 1909 (H-BR);V.
Bomboiza-Qualaquiza, Tres Cruces,Allioni 302 (H-
plane dorsallyand ventrally;gemmaeinconspic- BR, PC).
uous, ventral on tips of unmodifiedleaves. Spo-
rophytesunknown. Plants dull, brownish-green, with conspicu-
Illustrations. Florschiitz (1964, fig. 62 h-j);
ously glossy leaf bases, in low to tall tufts; rhi-
Reese (1977, figs. 11-13). zoids brown; stems to 6 cm tall but mostly shorter,
Distribution. (Fig. 27A). Endemic to South not often branched. Leaves dimorphic, vegetative
America:Brazil, FrenchGuiana,Suriname;very involute and mostly contorted when dry, 5-6.5
rare and known from only three collections in mm long, acuminate to ligulate-lanceolate, apex
additionto the type.
narrowly acute; margins thickened, coarsely
toothed with paired or triple teeth; leaf bases
Specimens examined. SURINAME. Parimaribo-
Kabel,bij km 121,J. & P. Florschiitz1813 (LAF,U). golden-brown, glossy, gradually narrowing to
FRENCH GUIANA. ST-LAURENT-DU-MARONI:Can- upper lamina, margins entire or somewhat den-
tondeMaripasoula, ca.6 kmN of Saiil,400m, Buck18935 tate; cancellinae eroded and generally lacking
(CAY,NY). or in part in older leaves, confined to
BRAZIL. PARA: Serra do Cachimbo, km 1223 on entirely
Cuiaba-Santarem road, vie. Serra Maze, Reese 16820 lower 1/3-1/2 of leaf base; median leaf cells
(INPA, LAF, NY). smooth dorsally, bulging ventrally, ca. 9-13 pm
long and 6-8 pm wide; gemmiferous leaves
Discussion. This very rareand highly distinc- rarely produced, narrow, stiffly erect, sometimes
tive moss was treatedby Florschiitz(1964) and reduced nearly to costa, bearing scanty, pale gem-
Reese (1977) as a variety of S. incompletus,but mae on ventral surface of tips. Seta dark red-
restoredto specific rank by Reese (1984), after brown, to 13 mm long; capsule 2 mm long;
discovery of a thirdcollection of the species, far peristome lacking or rudimentary; operculum to
from the collection sites of the first two speci- 1.8 mm long. Spores 16-22 pm, granular to finely
mens. A fourthspecimen was collected recently papillose. Calyptra smooth, 3 mm long.
by W. R. Buck in FrenchGuiana.The plantsmay Illustrations. Bartram (1949, fig. 35: A-C, as
bear a surprising resemblance to those of S. S. bernoullii); Reese (1977, figs. 21-24).
lycopodioides.The peculiarflat cells of theupper Distribution. (Fig. 44A). Endemic to tropical
lamina prevent the leaf from rolling up length- America: Mexico (Oaxaca); Central America,
wise when dry, as it does in many otherspecies West Indies; South America (Brazil, Colombia,
of Syrrhopodon. Ecuador, Peru, Venezuela, Trinidad). On tree
trunks and logs in forests, to 2000 m; more fre-
23. Syrrhopodon circinatus (Bridel) Mitten,J. quent at higher elevations.
Linn. Soc., Bot 12: 122. 1869. Fig. 42.
Selected specimens examined. MEXICO. OAXACA:
DicranumcircinatumBridel,Bryol.univ. 1: 446. 1826, Above Finca La Gloria toward Rio Grande, Sharp 5703
non DicranumcircinatumWilsonnec Dicranumcir- (MICH, NY, TENN).
cinatumR Brownter. in Dixon;Orthothecacircinata GUATEMALA. ALTA VERAPAZ:Cubilquitz,
(Bridel)Bridel,Bryol.univ. 1: 779. 1827.Type.Gua- Tiirckheim6942 (BM, H-BR, NY, PC, S).
deloupe.(B-Bridel). [IninsulaGuadaloupa adtruncos BELIZE. TOLEDO DISTRICT:Between Rancho Chico
arborum .... Clar. de Candolle communicavit.] and Cockscomb, Monkey River, Gentle 4415 (MO).
Syrrhopodonsordidus C. Miiller, Bot. Zeitung (Berlin) 6: NICARAGUA. 3 km NW of El Rosita, 15 Aug 1977,
766. 1848. Type. Venezuela. Carabobo,bei Cumbre de Danin s.n. (MO). BLUEFIELDS: 3.6 km S Cerro San Isidro,
Valencia, Funck & Schlim 617 (lectotype, JE; Proctor et al. 2 7189 (NY).
isolectotypes, BM, G, H-BR, L, NY, PC). COSTA RICA. Cocos ISLAND:Stewart 1411 (NY).
Syrrhopodon pittieri Renauld & Cardot, Bull. Soc. Roy. LIMON:1 mi. from Laguna Penitencia, Steere CR-36
Bot. Belgique 41(1): 54. 1905. Type. Costa Rica. Isla (NY). PUNTARENAS: Near Rinc6n, Koch 5041 (LAF, NY).
Cocos: 1898, Pittiers.n. (holotype, PC; isotype, NY). SANJOSE:vic. El General, Skutch2764 (NY).
Syrrhopodondiversifolius Brotherus, Bot. Jahrb.Syst. 56 PANAMA. COL6N:end of Santa Rita Road, Crosby
(Beibl. 123): 7. 1920. Type. Peru. Sandia: 10473 (MO). DARIEN: Between CerroPirreand Piji Vasal,
Chunchusmayo, Weberbauer1261 (H-BR). Folsom 6401 (LAF, MO). PANAMA: Summit ofCerro Jefe,
Syrrhopodon subrigidus Brotherus, Rev. Bryol. 47: 6. Allen 4940 (MO, NY).
1920. Type. Ecuador.Gualaquiza,1909, Allioni s.n. WINDWARD ISLANDS. GUADELOUPE: Basse Terre,
58 Flora Neotropica

0
42 c ?
A^ A ^~~0
B 0

00

a oO
0

00~~oo0y

FIGS. 4243. Syrrhopodon. 42. S. circinatus. A. Leaf outlines. B. Leaf apex. C. Margin and cells at leaf shoulder.
A from Funck & Schlim 617 (isotype of Syrrhopodon sordidus C. Miill., H), B from Allioni 502 (BM), C from
DeCandolle 6 (B). 43. S. hornschuchii. A. Outlines of vegetative leaves. B. Outline of leaf base. C. outline of
gemmiferous leaf. D. Margin and cells at leaf shoulder. A (leaf at left) from Miquel?s.n. (BM), deaf at right) Woronow
335 b =B1 from
(JE),
42B); mm (Figs. 19 (BM), C cfrom Woronow
42A,
Spruce Fi 335
43 . 2,
(JE', s. Miquel?s.n. (BM). Scale bars: a = 0.1 mm (Fig.
m4)D fromri
Calymperaceae 59

Maisonde la ForetNatl.Park,ProctorB-92663 (COLO, bases folded on the slide, involute-contorted

LAF).DOMINICA:Summitof MomeCouronne,Elliott when of lamina
205a (BM). dry; margins upper irregularly
COLOMBIA. ANTIOQUIA: Mun. CAceres,10-15 km winged and coarsely toothed, margins of lower
SE de Caceres,Callejaset al. 5352 (NY). META:Cordil- lamina irregularlytoothed, at least some of the
lera La Macarena,Schultes11261 (FLAS, LAF, NY). teeth spreading-recurved somewhat as in S.
Cordillera Oriental, NW of Bucaramanga,
SANTANDER:
Steere7634, 7641. rigidus;cancellinaefragile,more or less ephem-
VENEZUELA.AMAZONAS: CainoManteco,Bermtidez eral, small, not clearly differentiatedfrom adja-
PA-00177(herb.Bermidez,LAF). cent cells of lower lamina, confined to base of
TRINIDAD.ST.GEORGE: trailto MorneBleu, Crosby leaf andendingfarbelow shoulders;median leaf
2407(LAF, MO). cells rounded-rectangular,bulging ventrally,
ECUADOR.The types of S. subrigidusBroth. smooth or with low papillae dorsally,mostly ca.
PERU. LORETO: Maynas,80 kmNE of Iquitos,Timme x 9.5-12 pm; gemmiferousleaves stiffly
4837 (herb. Timme, LAF). SANDIA:Chunchusmayo, 7-8.5
Weberbauer1261 (NY). erect, consistingessentially of naked roughened
BRAZIL.PARA: Serrado Cachimbo,1208-1229kmN costa bearingdense clustersofgemmae ventrally
of Cuiabaon roadto Santarem,SerraMaze,Reese16578 at
tips. Seta 6-11 mm long; capsule 1.5-2 mm
(INPA,LAF,MICH,MO,NY).
long;peristomeobsolete or rudimentaryand of a
few erodedsegments,teethsometimesfused
Discussion. Prior to revision of the elimbate andhighly a
forming low, hyalinemembranearoundcap-
species of AmericanSyrrhopodon(Reese, 1977), sulemouth;operculumto2 mm long.Spores19-29
this species was generallyknownas S. bernoullii
C. Mill., a synonym of S. incompletus var. in- pm, granular.
Calyptra3 mm long, smooth.
Illustration. Reese (1977, figs. 25-29).
completus.The coloredleaf bases with theirfrag- Distribution. (Fig. 44B). Endemic to South
ile, erodedcancellinae,and the dimorphicleaves America: Colombia
(Amazonas, Caqueta,
(but gemmiferous leaves are only infrequently Vaupes);Venezuela (Amazonas);
Guyana;Sur-
present) distinguish S. circinatuls easily from S. iname; French Guiana;Ecuador;Peru (Loreto,
incompletus. Syrrhopodonrigidicsis somewhat Madre de
similarbut has stout, often spreadingor recurved Minas Dios); Brazil (Amazonas, Guapore,
Gerais, Para, Rond6nia, Roraima);
teeth on the leaf shoulders and its cancellinae Bolivia.
extend well up into the leaf bases, to the shoul-
ders.Syrrhopodoncircinatusis closely relatedto Selected specimens examined. COLOMBIA.
and very similar to the next species, S. CAQUETA:Rio Orteguesa, Woronow 335 (JE). VAUPES:
hornschuchii; see comments under the latter Soratama,Rio Apaporis, Schultes 12292 (FLAS, LAF).
name for distinctions; also see comments under VENEZUELA. AMAZONAS:
Dpto. Rio Negro, Neblina
Base Camp, 00?50'N, 66?10'W, Sastre-deJesus 325
S. rigidus.
(NY).
GUYANA. EssequiboRiver, MoraballiCreek, near
24. Syrrhopodon hornschuchii Martius, Fl. Bartica, Richards 90 (NY).
brasiliensis 1(2): 6. 1840. Type. Brazil. SURINAME. Via secta Moengo tapoe ad Grote
Martius s.n. (holotype M; isotypes, BM, H- Lanjouw& Lindeman818 (LAF,U).
Zwiebelzwamp,
FRENCH GUIANA. St-Laurent-du-Maroni,Canton
BR). Fig. 43. de Maripasoula, ca. 6 km N of Saiil, 200 m, Buck 18343
(CAY, NY).
SyrrhopodonrufusHomschuch,Fl. brasiliensis1(2): 6. ECUADOR.MORONA:
2?18'S, 78?07'W,Ortega U.
1840,nom.nud.in syn. 423 (LAF, Q). NAPO:Laguna de Cuyobene, Laegarrd
Syrrhopodon miquelianus C. Miller, Syn. musc. frond. 1: 51167 (NY). PASTAZA:
Curaray,Valle de Muerte, Holm-
535. 1849. Type. Suriname.Hb. Miquel.Probablyde- Nielsenet al. 22420 (AAU, NY).
stroyed;see Reese (1977). PERU. LORETO: 80 km NE of Iquitos,Timmie4834
Syrrhopodon surinamensis Dozy & Molkenboer, Prodr. (herb. Timme, LAF).
Fl. bryol.Surinamensis12. 1854.Type.Suriname.Van BRAZIL. ACRE:Cruziero do Sul, Prance etal. 11929
BeekVollenhoven s.n. (holotype,U; isotype,NY). (NY). AMAZONAS:
Rio Negro between Manaus and Sao
Gabriel, Buck 2184 (INPA, LAF, NY). PARA:Serra do
Plants coarse, dull-green; rhizoids dark-red; Cachimbo, km 877 on Cuiaba-Santarem road, Reese
16506 (INPA, LAF, MICH, MO, NY). RONDaNIA: vie.
stems to 2-3 cm tall but mostly less. Leaves first rapids on Rio Pacaas Novos, Reese 13637 (FLAS,
strongly dimorphic;vegetative linear to linear- INPA, LAF, MICH, MO, NY, US).
lanceolate,mostly 5-6 mm long, curvedandwith BOLIVIA. Near Mapiri, Williams 1921 (NY).
60 Flora Neotropica

TROPICAL AMERICA oraNeotroa base map no 1

II~-~~~--- - ~ ' . I___-........-

*~?~v~0qP*1---------
K

/ I
FIG.44~~~.
Dis~~tribu-~ofAA
----tions---- r

A. S. circinatus.B. S. hornschuchii.
FIG. 44. Distributionsof AmericanSyrrhopodon.
Calymperaceae 61

Discussion. See Reese (1977) forhistoryof the Illustrations. Florschiitz(1964, fig. 63); Reese
name Syrrhopodonhornschuchii.This species is (1977, figs. 42-43).
closely related to S. circinatus, from which it Distribution. (Fig. 51A). Endemic to the neo-
differs, among other ways, in its shorter can- tropics. Mexico (Veracruz). Central America:
cellinae and in having somewhat S. rigidus-like Costa Rica (Cocos Island,Puntarenas);Panama
teethon the marginsof the lowerlamina.It is also (Canal Zone, Cocle, Colon, Darien, Panamai,
similarin a generalway to S. rigichds,
withwhichit "Panama-ColombiaFrontier,"Veraguas). West
shares the trait of highly dimorphicleaves, but Indies:Jamaica;PuertoRico;Guadeloupe;Dom-
differsin its very shortephemeralcancellinaeand inica; Martinique;St. Vincent; Grenada.South
in lacking elongate hyaline marginalcells on its America: Colombia (Amazonas, Antioquia,
lowerlaminae. Nariiio, Santander,Valle, Vaupes); Venezuela
(Amazonas,Bolivar, Sucre).Guyana.Suriname.
25. Syrrhopodon rigidus Hooker & Greville, French Guiana. Ecuador. Brazil (Amazonas,
Edinburgh J. Sci. 3: 226. 1825. Type. St. Vin- Parai).Tree trunks,lianas,roots, logs, soil, rock,
cent. Upon trees on Mount St. Andrew, 1012 in moist to mesic forests from near sea level to
feet, Guilding s.n. (holotype, BM; isotypes, above 2100 m, most frequentabove 300 m.
BM-K, NY). Fig. 45. Selected specimens examined. COSTA RICA.
PUNTARENAS: E of Golfito, Crosby 3714 (MO).
Calymperes androgynum Montagne, Ann. Sci. Nat. Bot. PANAMA. DARItN:between Cerro Pirre and Piji
2,3:195. 1835; Syrrhopodonandrogynous (Montagne) Vasal, Folsom 6409 (LAF, MO). PANAMA:summit of
Bescherelle,Ann.Sci. Nat.Bot.6, 3: 197. 1876.Type. Cerro Jefe, Crosby 10053 (LAF, MO). VERAGUAS: N of
FrenchGuiana.sur les arbresvivana, Le Prieurs.n. main peak of Cerro Tute, Crosby 10816 (LAF, MO).
(holotype,PC-MONT;isotypes,BM, L). JAMAICA. [Without data;"Comm. G. Webster"herb.
Syrrhopodon longisetaceus C. Muller, Syn. musc. frond. Dixon ref. no. 506] (BM).
1: 535. 1849,nom. illeg. incl. spec.prior. PUERTO RICO. Sierrade Luquillo, Eslope of El Toro
Range, Steere 5324 (MICH).
Plants green to brownish-green,often glossy, WINDWARD ISLANDS. GUADELOUPE: Abords de
in dense, wiry tufts and cushions, with conspicu- i'Etang Zambi, Stehle 4052 (NY). DOMINICA:St. David,
Emerald Pool, P. & E. Hegewald 9335 (herb. Hegewald,
ous, glossy, golden-reddishleaf bases; rhizoids LAF, U). MARTINIQUE: Calabasse, Duss 81 (MICH). ST.
red, conspicuous; stems mostly ca. 1 cm tall, VINCENT: Valley of S fork of Cumberland River, Morton
forked.Leavesstronglydimorphic,gemmiferous 5816 (NY). GRENADA: Azima Mountainwoods, Apr 1905,
ones narrow,rigidly erect, often overtoppingthe Broadway s.n. (NY).
COLOMBIA. AMAZONAS-VAUPES: Rio Apaporis,
broader flexuous vegetative leaves, the latter 0?15'S, 70?30'W, Schultes & Cabrera 16585 (FH, NY).
mostly 7-11 mm long, linear or wider from ANTIOQUIA:Mun. Guatape, Sastre-de Jesus et al. 1280
scarcely broaderbase, apex blunt or acute with (HUA, LAF, NY). VALLE:Mun. Buenaventura, Churchill
costa sometimes excurrent;margins thickened et al 15255 (CUVC, LAF, NY). VAUPES: Rio Apaporis,
and coarsely toothed above, basal marginsusu- Soratma, Schultes 12292 (FH, NY).
VENEZUELA. AMAZONAS: Dpto. Rio Negro, along
ally borderedby several rows of elongate thick- Rio Mawarinuma, 00?50'N, 66?10'W, Sastre-de Jesis
walled more or less porose cells, and with 321, 327 (NY). BOLIVAR: 45 km N of Santa Elena,
irregularcoarsesharpteeth,at least some spread- Steyermarket al. 112357 (LAF, MO).
TRINIDAD. Mount Tocuche, E. G. Brittonet al. 1394
ing or recurved;cancellinae more or less persis- (NY).
tent, narrow,extending to leaf shoulders,distal GUYANA. Upper MazaruniDistr., E bank of Waruma
cells often colored; median leaf cells usually River, Gradstein 5024 (LAF, U).
incrassate,often with sinuose-porosewalls, cir- SURINAME. In montibus Bakhuis, Florschiitz &
cularto rectangular,bulgingventrally,smoothor Maas 3120 (LAF, U).
FRENCH GUIANA. CAYENNE:Le Prieur 371 (L). ST-
papillose dorsally; gemmiferous leaves com- LAURENT-DU-MARONI: Cantonde Maripasoula,ca. 6 km N
monly present, reduced to naked (or nearly so) of Saiil, 400 m, Buck18832 (CAY, NY), 18922 (NY).
costa with vestigial lamina at base, gemmae ECUADOR. Mt. Guayusa,22 Sep 1910, Allioni s.n. (H).
borne on tips. Seta red-brown,15-20 mm long; BRAZIL. AMAPA:Mun. de Oiapoque, Mori et al.
17230 (NY). AMAZONAS: along Rio Negro between
capsule ovoid, 1.5-2 mm long; peristome lack- Manaus and Sao Gabriel, Buck 2624 (INPA, LAF, NY).
ing; operculum2 mm long. Spores21.5-26.5 pm, PARA:Serra do Cachimbo, km 842 on Cuiabd-Santarem
granular.Calyptra3.5-4 mm long, smooth. road, Reese 16573 (INPA, LAF, MICH, MO, NY).
62 Flora Neotropica

45

a
b
\ ?c
d

B
?o
c

000

0J

00

FIGS. 45-46. Syrrhopodon. 45. S. rigidus. A. Leaf outline. B. Margin and cells at leaf shoulder. From Duss 241
p.p (H). 46. S. rupestris. A. Leaf outline. B. Portionof leaf base. C. Marginand cells at leaf shoulder. A from Lbtzelburg
22404 (JE), B from Spruce 14 (isotype of Syrrhopodon rupestris Mitt., BM), C from Litzelburg 22550 (JE). Scale
bars: a = 0.1 mm (Fig. 46B); b = 1 mm (Fig. 46A); c = 2 mm (Fig. 45A); d = 0.1 mm (Figs. 45B, 46C).
Calymperaceae 63

Discussion. This moss is somewhatsimilarto Selected specimens examined. COLOMBIA. AMA-

ZONAS:Rio Apaporis,Rio Popeyaca,Schultes & Cabera
S. circinatus-also with colored leaf bases-but
15561(FH,FLAS,LAF,NY).
easily distinguishedby its mostly linear leaves, VENEZUELA. AMAZONAS:Cafio Manteco, BermLidez
more persistent cancellinae extending up to the PA-00377 (herb. Bermidez, LAF); along Rio
leaf shoulders,marginsof the lower laminabor- Mawarinuma,Buck 12930 (LAF, NY).
deredby elongate hyaline cells and with coarse, BRAZIL. AMAZONAS: along Rio Negro between Manaus
andSaoGabriel,Rio Marieat Manauna,
Buck2374, 2376
sharp, often recurved teeth, and the common (INPA,LAF,NY);slopesandsummitof SerraCuricuriari,
presenceof the remarkablymodified gemmifer- Buck2449,2533,2559,2562(INPA,LAF,NY).
ous leaves (often lacking in S. circinatus). It is
also similar to S. hornschuchii,which has simi- Discussion. This rareendemic is easily distin-
larly modified gemmiferous leaves, but differs, guishedby its very long and narrowleaves with
among other ways, in its persistentcancellinae spinose tips, shapeof the cancellinae,long, slen-
thatextend much furtherup into the leaf base, to der stems, and habitat. See Reese (1977) for
the leaf shoulders.In S. circinatusthe leaf shoul- furtherdiscussionof this species. Althougha fair
dersareirregularlydentate-serrate butneverhave numberof modemcollectionsof S. rupestrisnow
sharp spreading-recurved teeth, and the can- exist, none is fertile or gemmiferous.
cellinae are always eroded and confined to the Syrrhopodonsteyermarkiiis similar; see com-
lower half of the leaf base. Erectnarrowgemmif- mentsunderthatspecies for distinctions.
erous leaves also occur in S. circinatus but are
neither as common nor as conspicuousas those 27. Syrrhopodon steyermarkii Robinson,Phy-
of S. rigidus. tologia 23: 390. 1972. Type. Venezuela. Ama-
zonas:CerroYapacana,Steyermark& Bunting
26. Syrrhopodon rupestris Mitten, J. Linn. 103154 (US). Fig. 47.
Soc., Bot. 12: 124. 1869. Type. Brazil. Fl. Plantsmoderatelyrobust;rhizoidsveryscanty,
Negro et Uaupes, Spruce 14 (NY). Fig. 46. dark
red; stems 2-8 cm tall, little branched.
Leaveslinear-acuminate,ending in a short,slen-
Plants tall and slender, dicranoidin appear-
der, coarsely toothed tip composed mostly of
ance; rhizoidsdark-red;stems 4-7(-10) cm tall, 6-7 mm long, involute and flexuous-
little branched. Leaves narrowly linear-acumi- costa,
when dry; margins of upper lamina
nate above much broaderbase, 6-9(-11) mm spreading
bearing a double rowof coarsemulticellularteeth
long, ending with excurrent distally coarsely above,thickened,with the
toothed costa, tightly involute and laxly con- thickeningsextending
down into lower laminaas conspicuous teniola-
tortedwhen dry,channeledandwidely spreading
like features, margins of lower lamina entire;
to recurvedwhen moist;marginsof upperlamina
cancellinaesmall, fragile,erodedin older leaves,
thickenedand slightly winged, coarsely toothed
not more than 1/3 lengthof lower lamina,longer
in 2 rows distally with paired teeth, marginsof next to than along costa; median leaf
lower lamina finely and more or less evenly cells margins
serrulate at shoulders; cancellinae fragile and x 13 incrassate, mostly rectangular,mostly ca. 7
pm, smooth dorsally, bulging ventrally.
eroded in older leaves, longer next to margins Gemmaeand
than at costa and forming a V-shapedarea (but sporophytesunknown.
Illustrations. Reese (1977, figs. 33-38); Rob-
this difficult to see unless the lower lamina is inson
(1972, figs. 4-6).
unfolded); median leaf cells squareto rounded- Distribution. (Fig. 27A). Narrowlyendemicin
rectangular,mostly ca. 8-12 x 6-7 pm, smooth northernSouth America. Colombia (Vaup6s);
dorsally, bulging ventrally, more or less porose Venezuela (Amazonas, Bolivar). On trees and
just above cancellinae.Genmae andsporophytes rocks to 1100 m.
unknown.
Illustration. Reese (1977, figs. 30-32). Specimensexamined(in additionto the type). CO-
Distribution. (Fig. 35A). A narrowendemicof LOMBIA. VAUPES: Rio Kananari (affluent of Rio
Cerro Isibukuri, Schultes & Cabrera 13298
northernSouth America.Colombia (Amazonas, Apaporis),
(FLAS,LAF).
Vaupes); Venezuela (Amazonas);Brazil (Ama- VENEZUELA.BOLIVAR: CerroGuaiquinima,5?54'N,
zonas); on bouldersalong streauns. 63?27'W, Sipman 26879 (B, NY).
64 Flora Neotropica

A ? CD D

a ___

A
/ 1/ ln X | ^K DF
9 II

ci E ".0 '0:
5 0Oao~o0
o 0
0 00

FIG. 47. Syrrhopodon steyermarkii. A. Leaf outline. B. Outline of leaf base. C. Leaf tip. D. Portion of leaf at
shoulder; dashed lines indicate intramarginalband of differentiatedcells. E. Marginand cells at leaf shoulder. F. Portion
of section from upper lamina. From Stcyermlark& Bunting 103154 (holotype of Syrrhopodonsteyermarkii Robins.,
US). Scale bars: a = 0.1 mm (C-D); b = 1 mm (A-B); c = 0.1 mm (E-F).

Discussion. This species is only known from from scarcely broader base, reduced to mere
the type and two other specinens, both from wings along costa distally, apex often merely the
north-centralSouth America. It is similar to S. excurrent, spinose-toothed costa; margins
rupestris in a general way but differs, among sharplyserrateat shoulders, irregularlytoothed
above, mostly unistratosebut slightly thickened
other ways, in the teniola-like featuresin its leaf
bases and the entiremarginsof the lower lamina. here and there, especially distally; cancellinae
broad, persistent, extending to or beyond leaf
28. Syrrhopodon theriotii Bartram,Contr.U.S. shoulders;medianleaf cells thick-walled,square
Natl. Herb. 26: 73. 1928. Type. Costa Rica. to rectangular,8.5-13 x 7-8.5 pm, smooth, bulg-
Heredia:On log, Cerro de las Lajas, North of ing orbluntlypapillosedorsally,stronglybulging
San Isidro, 2000-2400 m, 7 Mar 1926, ventrally.Gemmaeandsporophytesunknown.
Standley & Juvenal 51472 (holotype, FH; Illustrations. Bartram(1928, fig. 14); Reese
isotypes, JE,NY). Paratypes.Same localityand (1977, figs. 47-50).
date, Standley & Juvenal 51509b (FH), 51527 Distribution. (Fig. 27A). Endemic to Costa
(FH, NY). Fig. 48. Rica (Heredia,SanJose). Trees,logs, andstumps
at 1200-2400 m.
Plants yellowish-green, slender, to 8 cm tall,
Selected specimens examined. COSTA RICA.
freely forked;rhizoidsinconspicuous,yellowish- HEREDIA:5.5 km N of Vara Blanca, Croat 36064B, 36060
brown. Lea,es fragile, loosely flexuous-spread- (LAF, MO). SANJOSE:Cerros de Zurqui, 17 km NNE city
ing, to 15-20 mm long, long linear-acuminate center of San Jose, Crosby 9679, 10925 (LAF, MO).
Calymperaceae 65

B| i 00 5 g

~
000*00 0
u 0

A ' v aa__J

b C
d
e

/ oDO Wi

FIGS. 48-50. Syrrhopodon. 48. S. theriotii. A. Leaf outline. B. Leaf base. C. Midportion of upper lamina. D.
Margin and cells at leaf shoulder. From Svihla 3102 (FH). 49. S. xanthophyllus. A. Leaf outlines. B. Margin and cells
at leaf shoulder. A from Lintzelburg22324 (leaf at right) (JE), Spruce 16 (leaf at left) (holotype of Syrrhopodon
xanthophyllus Mitt., NY), B from Spruce 16 (holotype, NY). 50. S. cryptocarpos. A. Leaf outline. B. Margin and cells
at leaf shoulder. C. Cells of cancellina. A-B from Simmonds s.n. (CANM); C from Reese 16550 (LAF). Scale bars: a
= 0.1 mm (Fig. 48C); b = 2 mm (Fig. 48A); c = 1 mm (Figs. 48B, 49A); d = 0.1 mm (Figs. 48D, 50B-C); e = 1 mm
(Fig. 50A); f = 0.1 mm (Fig. 49B).
66 Flora Neotropica

Discussion. The long, slender leaves with Sometimeslocally abundant.

mostly unistratosemargins,narrowleaf base,and
often conspicuouslyexcurrentcosta, are distinc- Selectedspecimensexamined.VENEZUELA.AMA-
tive for this rare endemic. Many of the fragile ZONAS:Dpto. Rio Negro, along Rio Mawarinuma,
00?50'N,66?10'W,SastredeJesis 324 (NY).
leaves are brokenin most specimens. BRAZIL. AMAZONAS: N of Manaus at Ponte Negro,
Reese16001 (INPA,LAF, MO, NY). PARA: vicinity of
29. Syrrhopodon xanthophyllus Mitten, J. Reese16862(COLO,INPA,LAF,MICH,MO,
Santar6m,
Linn. Soc., Bot. 12: 115. 1869. Type. Venezu- NY). ROND6NIA:vie. first rapidson Rio PacaasNovos,
Reese13687(INPA,LAF,MO,NY, US).
ela. Javita, Spruce 16 (NY). See Reese (1977) BOLIVIA.BENI:Along Rio Yata, 40 km WSW of
for discussion of the type materialof this spe- Guayaramerin,Reese12900 (INPA,LAF,MO,NY).
cies. Fig. 49.
Discussion. Among otherfeatures,the colored
Heliconemaxanthophyllus(Mitten)Ellis& Eddyin Ellis, leaf
J. Bryol. 15: 728. 1989. Chameleionxanthophyllus bases, lack of peristome,andephemeralcan-
(Mitten)Ellis & Eddy in Eddy, HandbookMalesian cellinae distinguish this little moss from S.
Mosses2: 250. 1990. cryptocarpos;its cancellinarcells also lack the
distinctive thickenings of those of S.
Plants small, slender, glossy; rhizoids dark- cryptocarpos.It seems to be most closely related
purple; stems to 2 cm tall. Leaves ligulate to to S. rigidus, of which it is a miniatureversion
acuminate with mostly broadly mucronateapi- except for its sporophyte.AlthoughEllis (1989)
ces, 2.7-3.5 mm long from reddish-brown, assignedS. xanthophyllusto HeliconemaEllis &
glossy, scarcely broader bases, involute and Eddy, its relationships lie not with S.
curved when dry, ascending and more or less cryptocarpos but with S. rigidus and its allies
tubulose when moist, lying straighton slide but (Reese 1977). See discussion under S.
with base usually folded; marginsof upperlam- cryptocarpos for further comments on S.
ina thickened, irregularly serrate to serrulate, xanthophyllus.
often with low, obscure, paired teeth, lacking
stereids in section, margins of lower lamina 30. Syrrhopodon cryptocarpos Dozy &
nearlyentireto regularlyserratewith sharpteeth, Molkenboer, Prodr. Fl. bryol. surinamensis 14.
often bordered with several to many rows of 1854. Type. Suriname. Van Beek Vollenhoven
elongate cells; cancellinae fragile, quickly s.n. (holotype, U; isotypes L, NY). Fig. 50.
eroded and ephemeral,small and not clearly de-
marcated,obliquely truncatedistally and longer Heliconemacryptocarpos(Dozy & Molkenboer)Ellis &
at leaf margins than at costa; median leaf cells Eddy in Ellis, J. Bryol. 15: 728. 1989. Chameleion
cryptocarpos(Dozy & Molkenboer)Ellis & Eddy in
thick-walled,rounded-squareor elongate,mostly Eddy,HandbookMalesianMosses2: 250. 1990.
ca. 8 pm, nearly smooth or with low papillae Syrrhopodon spruceanusMitten,J. Linn.Soc., Bot. 12:
dorsally, strongly bulging ventrally; gemmae 114. 1869. Type. Brazil. Fl. Negro, ad Sao Gabriel,
scarce, borneventrallyalong costa and leaf mar- Spruce3 (holotype,NY; isotypes,BM, BR, L, MICH,
S).
gins between shouldersand leaf tip, mostly 100- Calymperesleboucherianum Paris& Brotherusin Paris,
200 pm long, clavate, fusiform, or filamentous Rev.Bryol.33: 36. 1906;Syrrhopodon leboucherianus
with thick, dark-browncell walls. Sporophytes (Paris& Brotherus)Reese, Bryologist64: 139. 1961.
completely immersed;seta very short; capsule Type. FrenchGuiana.Mtgnedes Peres, 16 Oct 1906,
1 mm long; peristome lacking; operculum 0.7 Galliots.n.(holotype,REN;isotype,L).
mm long. Spores 17-21 pm, finely granular. Syrrhopodon Crum,Trans.Brit.Bryol.
alatomarginatus
Soc. 3: 72. 1956.Type.Trinidad.St. GeorgeCo.:Top
Calyptra minute, ca. 0.5 mm long, conical, of BlanchisseuseRoad, 25 Sep 1949, Simmondss.n.
extending down only to base of rostrum,papil- (CANM).
lose above.
Illustration. Reese (1977, figs. 39-41). Plants small, slender, dull, green to yellowish-
Distribution. (Fig. 52A). Endemicto northern green, in low, thin, turfs; rhizoids purple; stems
South America. Venezuela (Amazonas);Brazil to 2.5 cm tall but mostly much shorter. Leaves
(Amazonas,Pari, Rond6nia);Bolivia (Beni). On mostly 2.5-3.5 mm long, somewhat contorted
soil, rock,rottedwood,treetunks, andtermitenests. and often secund at stem tips when dry, ligulate
Calymperaceae 67

TROPICAL AMERICA Flora Neotrop,ca base map no 1

--- - - -
_ K---
-, -... ' 1

"'I ""~~`~~tr>
I -i-?---- -..>.. -
O

iN

FIG. 51. Distributions of American Syrrhopodon. A.. . rigidus.B. S. cryptocarpos.

68 FloraNeotropica

to elliptic or oblanceolatefrom scarcely broader ECUADOR.NAPO: Afiangu,75 km E of PuertoFran-

cisco de Orellana,Churchill& Sastre-deJesius13790
base, apex blunt; margins thickened, coarsely
(LAF, NY).
serrate-toothedin rows forming three or more PERU. LORETO:
SW of Iquitos on Rio Nanay, Croat
irregular wings; cancellinae small, narrow, 19021A(LAF,MO).
mostly persistent, reaching to or beyond leaf BRAZIL.ACRE:33 km SW of Rio Branco,Reese&
shoulders,cells of cancellinaeoften more or less McPherson 13261 (INPA, LAF, MO, NY, US). AMAZO-
NAS: Manaus, at INPA, Reese 12668 (INPA, LAF, MO,
eroded,bearingdistinctivefainttransversethick- NY). PARA: Serra do Cachimbo, km 885 on Cuiaba-
enings; median leaf cells circularto rectangular, Santar6mroad, Reese 16544 (INPA, LAF, MICH, MO,
mostly 9-14 pm long and 5.5-8.5 pm wide, NY). ROND6NIA: first rapids on Rio Pacais Novos, Reese
sharply papillose dorsally, mammillose to 13638 (FLAS, INPA, LAF, MICH, MO, NY, US).
RORAIMA: km 328 on Manaus-Venezuela road, Buck et
mammnillose-papillose ventrally;gemmae infre- al. 1821 (INPA, LAF, NY).
quently produced,scarce, in low, dense clusters BOLIVIA.BENI:15-17 km WSW of Guayaramerin,
along ventral surface of costa and occasionally Reese13065(INPA,LAF,MICH,MO,NY, US).
on thickened leaf margins near apex. Sporo-
phytes immersed,inconspicuous;seta to ca. 0.7 Discussion. This distinctivespecies is readily
mm long, completely enclosed by vaginula, identifiedthroughleaf shape, the triply or more
which extends to base of urn;capsule urceolate, winged margins,and the cells of the cancellinae
to 1 mm long; peristome fragile, teeth irregular, with faint transversethickenings.The immersed
transverselystriate-papillose,to ca. 170 pm tall; capsules are also distinctive; only S.
operculumrostrateto conic-rostrate,ca. 0.7 mm xanthophyllusshares the latter feature among
long. Spores 19-24 pm, smooth or granular.Ca- neotropical Syrrhopodon.Ellis (1989) treated
lyptra ca. 0.7 mm long, campanulate-conical, both S. cryptocarpos and S. xanthophyllus in
papillose distally, barelycovering operculum. HeliconemaEllis & Eddy, (nom. illeg. = Cham-
Illustrations. Crum(1956, figs. 1-6, as S. al- eleion Ellis & Eddy in Eddy), primarilybecause
atomarginatusCrum);Florschiitz(1964, fig. 61); of theirimmersedsporophytes.However the two
Reese (1977, figs. 44-46). are only superficially similar, with the latter
Distribution. (Fig. 51B). Africa (Gabon, clearlymoreclosely relatedto the suite of species
Zaire); Central and South America. Panama including S. circinatus, S. hornschuchii, S.
(Colon). Colombia (Amazonas, Antioquia, rigidus, S. rupestris,and S. steyermarkii(Reese
Putumayo, Vaupes). Venezuela (Amazonas). 1977: 27). The immersed-sporophytehabit has
Trinidad. Guyana. Suriname. French Guiana. evidently arisen separatelyin two different lin-
Ecuador(Napo). Peru (Loreto, Madrede Dios). eages of AmericanSyrrhopodon(Reese 1977);
Brazil (Acre, Amazonas, Para, Rond6nia, thusit is a moot questionas to whetheror not the
Roraima). Bolivia (Beni). Widespreadand lo- conditionmeritsgenericrecognition.
cally very common in South America. Tree
trunks,rotted logs, stumps, termitenests, in wet 2. Calymperes Swartzin F. Weber,Tab.Calyptr.
forests to ca. 700 m. oper. musc. frond.gen. 2. 1813. Type species:
C. lonchophyllumSchwaegrichen(designated
Selected specimens examined. PANAMA. COLON:
along path at end of Santa Rita Ridge Road, Crosby 10308
by Williams, Bull. TorreyBot. Club 47: 385.
(LAF,MO). 1920). The name Calymperesis derived from
COLOMBIA. ANTIOQUIA:Mun. Taraza, 210 km N de a composite of two Greek words meaning a
Medellin, Churchill et al. 14960 (HUA, LAF, NY). "covering,"and "to pierce through,"presum-
VENEZUELA. AMAZONAS: Dpto. Rio Negro,
ably in reference to the escape of the spores
00?50'N, 66?10'W, Buck 12965 (NY).
TRINIDAD. Piarco Savanna, S of Arouca, E. G. throughthe perforationsof the calyptra.
Britton et al. 115 (NY).
GUYANA. Basin of Kuyuwini River, Smith 2603a Plants mostly tufted,with erect,mostly simple
(MICH, NY). stems; cells of upper lamina isodiametric or
SURINAME. Kreekbos, Zanderij, J. & P. Florschiitz
788 (LAF, U). transversely elongate; leaf margins generally
FRENCH GUIANA. ST-LAURENT-DU-MARONI: Can- thickenedbut mostly lacking elongate, hyaline
ton de Maripasoula, ca. 6 km N of Saiil, 200-400 m, Buck cells; teniolae, gemma receptacles, and highly
18718 (NY), 18881 (CAY, NY). modifiedgemmiferousleaves common;capsules
Calymperaceae 69

exserted; peristomes lacking; calyptra plicate, cellulae foliorum protruberantes sed

persistent, enclosing the capsule, clasping the epapillosae.
operculumby its rostrumandtwistingaroundthe
seta below, bearingvertical slits above through Type species. Calymperesrubiginosum(Mitt.)
which the spores escape. Reese, Bryologist 78: 92. 1975. Syrrhopodon
Calymperes is world-wide in the tropics and rubiginosusMitt.,J.Linn.Soc., Bot. 12: 125. 1869.
subtropics and particularlywell-developed and
frequent in warm humid low-lying evergreen By virtueof theirsmall stature,frequentpurple
rainforests. The center of distributionfor the color, large bulging epapillose leaf cells, and
genus in the neotropics is northeasternSouth unistratosecrenulatemarginsof the upper lam-
America.Sixteen species of Calymperesoccurin ina, these plantsarehighly distinctiveand with-
the neotropics. Most species of Calymperesare out near relatives anywhere in the world, thus
corticolous but a few grow on rocks and soil; meriting recognition at the subgeneric level.
some species attain modest elevations but the Thereis only one species, endemic to the Amer-
majorityare plantsof warm,humidlowlands. ican tropics.
Threesubgeneraof Calympereshave been for-
mally recognized world wide (see Reese, 1987a When the capsules of Calymperesare dry, the
for review), and a fourthis describedas new in operculum-its rostrumgrippedby the tip of the
this treatment.The subgeneraandtheirAmerican calyptra-is held away from the mouth of the
species are listed alphabeticallybelow. capsule, allowing the spores to disperse
throughthe gaping vertical slits in the calyptra.
SubgenusCalymperes When the capsules are moist the operculum is
CalympereslevyanumBesch., C. lonchophyllum held against the mouthof the capsule-closing
Schwaegr., C. venezuelanum(Mitt.)Pittier it-and the slits in the calyptraare closed. (See
Fig. 72E.)
Subgenus Hyophilina(C. Mull.) Broth. The sole character that distinguishes
Calymperesafzelii Sw., C. bartramiiReese, C. CalymperesfromSyrrhopodonis the remarkable
erosumC. Mull., C. guildingii Hook. & Grev.,C. persistentcalyptra;the two genera intergradein
mitrafugax Florsch., C. nicaraguense Ren. & all other respects. Teniolae, when present, are
Card.,C. othmeriHerzog, C.palisotii Schwaegr., almost diagnostic for Calymperes,but similar
C. platyloma Mitt., C. smithii Bartr.,C. tenerum featuresoccur in the Syrrhopodonincompletus
C. Mull. complexandelsewherein thegenus. Calymperes
is easy to recognizewhensporophytesarepresent
Roth
SubgenusSomphoneurumn or if the leaves bear teniolae. In the absence of
Calymperespallidum Mitt. teniolae, sterile specimens of Calymperesmay
occasionallybe difficultto distinguishfromster-
SubgenusPorphyrophylumReese, subg. nov. ile plants of species of Syrrhopodonwith un-
Folia in margine unistratosa,valde crenulata; borderedleaves.

Key to NeotropicalCalymperes

Upperlaminanarrowlylinear,its cells oftenormostlytransversely

elongate,at leastin part;costain section
usuallyshowing2 or morerowsof guidecells.
2. Leavesstiff, fragile,upperlaIninaconnectedto lowerlaminaby a distinct,petiole-likeconstriction.
............................................. 3. C. venezuelanum.
2. Leavessoft,flexible,lackingpetiole-likeconstriction.
3. Cells of upperlaminasmoothdorsallyandventrally;leavesmostlymorethan8 mm
long . ................... C.. lonchophyllum.
3. Cells of upperlaminapapillosedorsallyandventrally;leavesmostlyless than8 mm
long. .......................................... 2. C. levyanum.
70 Flora Neotropica

1. Upperlaminavariouslylanceolateto ovate,broadlylinear,orsubulate,its cells mostlyisodiametric;costain

sectionmostlyshowinga singlerowof guidecells.
4. Leavesbearing,at leastin shoulderregions,eitherdistinctteniolaeornarrowto broadmar-
ginalbandsof differentiated,
elongatecells.
5. Teniolaepresent(sometimesfaintormoreor less lackingin someleaves,especiallyin
C. nicaraguense and C. othmeri).
6. Distalcells of cancellinaedistinctlymammilloseventrallywithdistally-pointing
mammillae.
7. Costain sectionshowingwell-definedbandsof stereidcells. 6. C. erosum.
7. Costain sectionlackingwell-definedbandsof stereids. . . . 4. C.pallidum.
6. Distalcells of cancellinaeplaneventrally.
8. Teniolaeoftenfaintormoreorless lackingor interrupted.
9. Gemmae borne all aroundtip of excurrentcosta. . . . 7. C. othmeri.
9.
Gemmaeborneonly on ventralsurfaceof costatip.
........................ 15. C. nicaraguense.
8. Teniolaewell-developedandconspicuous.
10. Cancellinaeformingverybroadanglesdistallywithcosta;
upperlaminaoftenas wide as or widerthanlowerlamina;
gemmiferousleavesoftenstronglydifferentiated, the gem-
maebornein hairbrush-like tuftsin a distinctgemma-
receptacle ................... . 8. C.palisotii.
10. Cancellinaemostlyendingin acuteanglesdistallyalong
costa;upperlaminamostlyconspicuouslynarrowerthan
lower;gemmiferousleavesnotusuallymuchdifferentiated,
thegemmaeneitherin hairbrush-like tuftsnorin a distinct
gemma-receptacle.
11. Cellsof teniolaelackingconspicuouslythickened
irregular walls. . . ......... . 5. C. afzelii.
11. Cellsof teniolaeat shoulderswithconspicuously
thickened,oftenpittedandirregularwalls.
12. Marginsof upperlaminastronglyinfoldedwet
anddry. . ......... 13. C. bartramii.
12. Marginsof upperlaminaplaneto erect.
.................. 14. C. guildingii.
5. Teniolaelackingbutleavesborderedatshouldersby distinctbandsof differentiated,
elongatecells.
13. Bordernarrowat shoulders,mostly2-6 cells wide.
. . . . . . . . . . . . . . . . . . . . . . . . . . 10. C. smithii.
13. Border8 or morecells wideat leaf shoulders.
14. Cellsof upperlaminastronglypapillosedorsally,to 16pm
diam.;marginsfinelyserrateabove. 11. C.platyloma.
14. Cellsof upperlaminafinelypapillosedorsally,to
12 pm diameter, margins entire. . . . 12. C. mitrafugax.
4. Leavesbearingneitherteniolaenormarginalbandsof differentiated cells.
15. Plantsgreen;cells atmidleafquadrate,6-8 pm
diam.marginsof upperlaminaentire;gemmae
borneall aroundtipof excurrent costa. 9. C. tenerum.
15. Plantsoftenpurple;cells atmidleafrounded-
subquadrate,to 13 pm diam.;marginsof upper
laminacrenate;gemmaeborneonly on ventral
surfaceof costa. ........ 16. C. rubiginosum.

1. Calymperes lonchophyllum Schwaegrichen, CalympereslonchophylloidesC. Miiller, Gen. Musc.

Frond. 362. 1901. Type. Guatemala.Waldem von
Spec. Muse. frond.suppl. 1: 333. 1816. Type. Chis6r,[collectorunknown],n.v., probablydestroyed.
In arboribusguyanne, Richards.n. (holotype,
PC; isotype, G). Fig. 53. Plantsdarkgreen,in low dull tanglesanddense
stems to 20 mm tall but commonly
Calymperes asperipes Bescherelle, Ann. Sci. Nat. Bot. 8, springymats;
1: 277. 1896. Type. Guadeloupe. Ed. Marie 300 (holo- very short and the plants appearing stemless.
type, BM? n.v.; isotype NY). Leavesnarrowlylinear,to 14 mm or more long,
Calymperaceae 71

TROPICALAMERICA Noo-owa Fioe tbae nap no1

006

. .
B C~~~\
-------------(
?: I `r'~~~~~~~'
I

FI. 2.Ditrbuios f meiciiSyrhpoonan alniers.A.S.xathphlls.B.C.lochphllm

B. C. lonchophyllum.
andCalymperes.A. S. xanthophyllus.
FIG. 52. Distributionsof AmericanSyrrhopodon
72 FloraNeotropica

variouslycontorted-coiledabove whendry,often 500-600 m, E. & P. Hegewald9509 (herb.Hegewald,

LAF). GRENADA: FloridaEstate,Simmonds480 (NY).
transversely undulate; margins heavily thick- GUADELOUPE: Basse-Terre, De Sloover 23525 p.p. (NY).
ened, remotely toothed above; cancellinae MARTINIQUE: L'Alma, W of Route Nationale 3, N of River
sharply distinct; cells of upper lamina smooth, Blanche, 500 m, Crosby 4735 (LAF, MO). ST. LUCIA:E
some or most transversely elongate, often of Le GrandMagasin, Simmonds 246 (NY). ST. VINCENT:
bistratose; costa smooth. Seta 10 mm; capsule Valley of N Forkof CumberlandRiver, Morton5498
1.7-2.5 mm long, cylindric;operculum1.1 mm. (NY).
COLOMBIA. AMAZONAS: Ca. 1-1.5 horas Rio
Spores 17-24 pm, finely papillose. Calyptra4- Caquetaarribade La Pedrera,Churchilletal. 16138 (LAF,
4.7 mm, papillose-roughenedabove. NY). ANTIOQUIA: Mun.Taraza,6.5 km E de El Doce, 200
Selected illustrations. Bartram(1949, fig. 37, m, Sastre-DeJesis et al. 955 (HUA,LAF,NY). VAUPES:
Rio Apaporis,Schultess.n. (FH,FLAS,LAF).
D-G); Florschuitz(1964, fig. 41); Reese (1961, VENEZUELA. AMAZONAS: Dpto. Rio Negro,
figs. 1-6); Reese et al. (1986, figs. 4-7). 00?50'N, 66?01'W, Buck 12867 (NY). BOLIVAR:ca. 45
Distribution. (Fig. 52B). Pantropical.Mostly kmS of Cd.Piar,Purselletal.8045 (LAF,PAC).FALC6N:
corticolous,occasionallyon rockanddeadwood; ParqueNacionalQuebradade la CuevaEl Toro,600 m,
Liesner et al. 7710 (LAF, MO). MIRANDA:
Parque Nac.
forests, to ca. 600 m. Throughoutthe American Guatapo,Nee 17647(NY).
tropics and subtropics. TRINIDAD. ST. ANDREW:
Just E of Valencia, Crosby
2054 (DUKE,LAF).
Selected specimens exanmined.MEXICO. QUINTANA GUYANA. EASTDEMERARA DISTR.:Timehri, Dakara
Roo: 40 km marker above Kuntunil-kin, Sharp & McF- Creek,10m, Gradstein4737(LAF, U).
arland 9380 (LAF, TENN). VERACRUZ:Zona de los Rios SURINAME.Lelygebergte,J.& P. A. Florschiitz4840
Uxpanapa y Solosuichil, Delgadillo M. 3434 (LAF, (LAF,U).
MEXU). FRENCH GUIANA. ST-LAURENT-DU-MARONI:Can-
GUATEMALA. ALTAVERAPAZ: Near Chamd,John- tonde Maripasoula, ca.6 kmN of Saul,200m, Buck18429
son 347a (NY). IZABAL: Montafia del Mico, Steyermark (CAY,NY).
38891 (F). PETEN:Uaxactun, Bartlett 12472 (NY). ECUADOR.NAPO:75 km E of PuertoFranciscodel
BELIZE. Prospecto-Maskall road, Gentle 898-C Orellana,Churchill& Sastre-deJesus 13877 (NY).
(LAF). PERU. LORETO: Maynas,80 km NE of Iquitos,90 m,
HONDURAS. Trumfo Lagoon, Wilson 183 (NY). Timme4816 (herb.Timme,LAF).
NICARAGUA. BLUEFIELDS: 3.6 km SE Cerro San BRAZIL.ACRE: 15km E of Rio Brancoalongroadto
Isidro, Proctor et al. 27095 p.p. (NY). COMARCA DEL Abuna,Reese& McPherson13188 (INPA, LAF, NY).
CABO: Seymour 5878 (MO). ALAGOAS: Mun.Messias,33 km N of Macei6,Boom&
COSTA RICA. LIMON:Vic. Tortuguero,Steere CR-55 Mori1033 (NY). AMAPA: Mun.de Macapa,156km NW
(NY). PUNTARENAS:Rincon de Osa, Liesner 1954b (MO). of PortoGrande,Mori et al. 17533 (NY). AMAZONAS:
PANAMA. CANAL ZONE: Barro Colorado Island, Road to Balbinahydroelectricdam project,Buck2668
SalazarAllen & Chung C. 7856 p.p. (PMA). COLON: End (INPA,LAF,NY); CEPLACagricultural station,30 km
of Santa Anita Ridge Road, 350-450 m, Crosby 10319 N of Manaus,30 m, Griffin,IIIet al. 120 (FLAS,LAF).
(LAF, MO). PANAMA: Vie. Cerro Brewster Brako 8324 BAHIA:Mun. Itanagra,8 kmW of Itanagra,Boom & Mori
(NY). SAN BLAS:Vic. Puerto Obaldia, D'Arcy 13672A 941 (NY). MATOGROSSO: Aripuana, Dardanelos, Lisboa
(NY). VERAGUAS:5.4 km NW of Santa Fe, Crosby 10195 etal. 367(FLAS,INPA,LAF).PARA: Serrado Cachimbo,
(MO, NY). km 877 on Cuiaba-Santarem road,Reese 16517 (INPA,
CUBA. ORIENTE: Valle de Rio Jaguani, Pocs & Reyes LAF,MICH,MO,NY, US). RONDONIA: Abovefirstrap-
9169/AH (NY). ids on Rio PacaasNovos, Reese 13514 (FLAS, INPA,
JAMAICA. John Crow Mtns., 520 m, N. L. Britton LAF,MICH,MO,NY, US).RORAIMA: Bocade Mata,216
4156 (LAF, NY). ST.THOMAS:SW slopes of Winchester km N of Boa Vista,800 m, Bucket al. 2024 (INPA,LAF,
Peak, 1700 ft., Crosby 3325 (DUKE, LAF). TRELAWNY: NY).SAOPAULO: IlhadoCardoso,Vital10342(LAF,SP).
3-5 mi. from Troy heading ca. 300?, ca. 2000 ft., Crosby
3131 (DUKE, LAF). Discussion. Its linearleaves, compact springy
DOMINICAN REPUBLIC. ESPAILLAT: 7.5 km E of
habit, and commonlystemless appearancemake
Gaspar Hernmndez,Buck 19376 (NY).
PUERTO RICO. Caribbean National Forest, Luquillo C. lonchophyllum easy to recognize. Only C.
Divison, 1.9 mi. S of entrance to El Verde Biological levyanumand C. venezuelanumare similar; the
Station, 500 m, Reese 14877 (LAF, NY). papillosecells of the formerandpetiole-like con-
LEEWARD ISLANDS. MONTSERRAT: 5.3 km ESE of strictionabove the leaf base of the latterdifferen-
Plymouth, Mar 1896, Boveys.n. (NY). ST.EUSTATIUS: The tiatethem
Quill, van Slageren 31M (NY, U). ST.KITTS:Mt. Misery, easily. As in C. levyanum,plantsof C.
Wadsworth470 (NY). lonchophyllumcommonlyhave such shortstems
WINDWARD ISLANDS. DOMINICA: St. Andrews: that the leaves appearto arise directly from the
Nach Wet Area Agriculture Station Richtung Marigot, substrate.
Calymperaceae 73

53,o3 ', .C3 o D

0

D O

9
0

ox7
-oo/
/ a a _ _ _ _

O,oO

OOo r:~~~~
FIS 5-4.Clypre.53 .loncpyln A. Lefbs.B efapx .Mrin cll at mila.D
Portios of ectios oflaf. AB fro Broaday 783, C rom M.& H. tehid3972, (topleft)from Boadwa 7843
(scinwihcsa fo .&H.Sek 97,(trgh)fo teemr 46.5.C l-ynm A.
\_Leafoule.B
Marinan cll a lafshuler C. P rtionofla scin.D agi n clsa mde.ACf omRbis6D
from evy sn. (iotypeof Caympers leyanumBeach, NY) Scal bars a =1 mm Figs.53A-B 54A) b = .1 m

FIGS. 53-54. Calymperes. 53. C. lonchophylqlm. A. Leaf base. B. Leaf apex. C. Margin and cells at midleaf. D.
Portions of sections of leaf. A-B from Broadway 7843, C from M. & H. Stehle 3972, D (top left) from Broadway 7843,
(section with costa) from M. & H. StehlM3972, (at right) from Steyermark4465. 54. C. levyanum. A. Leaf outline. B.
Margin and cells at leaf shoulder. C. Portion of leaf section. D. Margin and cells at midleaf. A-C from Robbins 6, D
from Levy s.n. (isotype of Calymperes leyanum Besch., NY). Scale bars: a = 1 mm (Figs. 53A-B, 54A); b = 0.1 mm
(Figs. 53C-D, 54B-D).

2. Calymperes levyanum Bescherelle,Ann.Sci. Stems to ca. 1.5 cm tall, mostly simple, usually
Nat. Bot. 8, 1: 290. 1896. Type. Nicaragua. very short and the plants appearing stemless.
Levys.n. (holotype, BM?, n.v.; isotypes NY, Leaves narrowlylinear, mostly less than 8 mm
PC). Fig. 54. long, contorted-coiledat tips when dry, more or
less straightbelow; margins thickened, toothed
Plants dull-green, in low, springy tangles. or entireat shoulders,serrateabove; cancellinae
74 Flora Neotropica

TROPICAL AMERICA Fora Neotrop.cabase map no 1

-7

,.. .. ; . X o W ............

FIG.5. Dm....." : ,- B-.C. _-_----

FIG. 55. Distributions of American Calymperes. A. C. levyanum. B. C. venezuelanum.

Calymperaceae 75

sharplydisinct, often stoutlybulgingventrallyin Syrrhopodonvenezuelanus Mitten, J. Linn. Soc., Bot. 12:

125. 1869.Type. Venezuela.Schomburgks.n. (holotype,
distal portion;cells of upperlamina usually ob-
NY; isotypes, BM, H).
scure and densely papillose in upperpartof leaf, Calymperesmaguirei Bartram,Bull. Torrey Bot. Club47:
but sometimes papillae small and ratherincon- 393. 1920. Type. Suriname. Tafelberg, Maguire
spicuous, cells sometimes smoothor nearlyso in 24549M (holotype, NY; isotypes, FH, U).
lower part of upper lamina, at least some trans-
Plants glossy, pale green above, blackish in
versely elongate, especially just above can- older parts, in grass-like tufts. Leaves rigidly
cellinae; costa smooth. Seta 6 rmm;capsule 1.8 erect to flexuous, wiry, narrowly linear, mostly
mm long, cylindric;operculumca. 1.7 mm long.
broken off at midleaf, to 25 mm long, lamina
Spores 22-24 pm, finely roughened. Calyptra
interrupted above cancellinae by interval of
roughenedabove. naked costa; margins thickened, remotely
Illustrations: Florschiitz(1964, fig. 43); Mag-
toothed above, entire to serrate or toothed at
ill (1981, fig. 44: 10-18); Reese (1961,figs. 7-10).
Distribution. (Fig. 55A). Mexico, Central shoulders; cancellinae sharply distinct; cells of
America, Cuba, Puerto Rico, northern South upper lamina smooth, transversely elongate,
often bi- tristratose; costa smooth. Sporophytes
America,southernAfrica, southernChina(as C.
unknown.
levyanum var. hainanense Reese & P.-j. Lin). Illustrations. Florschiitz (1964, fig. 42); Reese
Tree trunks,logs; in forests to ca. 350 m.
(1961, figs. 11-14); Reese & Tan (1983, figs.
Selected specimens examined. MEXICO. VERACRUZ: 19-26).
Zona de los Rios Uxpanapa y Solosichil, Delgadillo M. Distribution. (Fig. 55B). Central America
3371 (LAF, MEXU). (Panama); Jamaica; St. Vincent, Guadeloupe;
BELIZE. EL CAYO:Arenal, Mains 3542 (NY), 3543
Venezuela; Trinidad; Guyana; Suriname; French
(FH, LAF). Guiana. Soil and tree trunks, ca. 300-900 m.
NICARAGUA. BLUEFIELDS: 3.6 km SE of Cerro San
Isidro, Proctor et al. 27095 p.p. (NY). Selected specimens examined. PANAMA. PANAMA:
COSTA RICA. PUNTARENAS: 12 km S of Rinc6n de summit of Cerro Jefe, Allen 4927 (LAF, MO).
Osa, Bowers 603-a (TENN, LAF). VENEZUELA. ISLAMARGARITA: Cerro Copey, Sug-
PANAMA. CANAL ZONE:Barro Colorado Island, den 1103 (NY). NUEVAESPARTA: La Sierra, 700 m, Bhat
Crosby 3940 (DUKE, LAF). VERAGUAS: 5.4 km NW of Nf-0010-76 (LAF, TENN). SUCRE:Peninsula de Paria,
Santa Fe, Crosby 10136 (LAF, MO).
Steyertark 91112 (FH, LAF).
CUBA. Sierrade Gavilanes, Leon& Clemrent 6483 (NY). TRINIDAD. Margarita,Kruger s.n. (NY).
PUERTO RICO. Arecibo to Utuado, E. G. Britton333 GUYANA. Upper Mazaruni River region, Boom &
(NY). Gopaul 7387, 7446, 7494, 7507 (NY).
BRAZIL. AMAZONAS: Rio Javari, Estirao do Equador,
16 Nov 1975, Ramos s.n., (INPA, LAF). PARA:Serra doDiscussion. This rare species is easy to recog-
Cachimbo, km 1011 on Cuiabi-Santar6m hwy., 300 m,nize by its grass-like habit, brittle leaves, and
Reese 16741 (COLO, INPA, LAF, NY); km 777 on
Cuiaba-Santarm hwy.,400m, Reese 16080 (INPA, LAF,petiole-like constriction above the leaf base. The
MICH,MO,NY, US). remarkable latter feature is shared with three
unrelated paleotropical members of
Discussion. This is a rarespecies thatis rarely Calymperaceae (Reese & Tan 1983).
collected. It is like a small version of C.
lonchophyllumn, usually appearingdull because 4. Calymperes pallidum Mitten, Philos. Trans.
of its generallyvery papillose leaves. The leaves Royal Soc. London 168: 388. 1879. Type.
of some specinens (such as Bowers 603-a, cited Rodriguez. Balfour s.n. (lectotype, BM).
above) may be less papillose thanthoseof others Fig. 57.
andmore like the leaves of C. lonchophyllum.Its
uleanut Brotherus,Hedwigia 51:124. 1912.
small stature may result in C. levyanumbeing Calymperes
Type. Brazil. Goias: An Palmstammen am Tocantins,
overlooked in the field or being passed over as Oct 1892, Ule 1556 (holotype, H; isotype, NY).
underdevelopedplantsof C. lonlchophyllum. Calymtperescubense Willliams, Bull. Torrey Bot. Club
47: 387. 1920. Type. Cuba. Santa Clara: Ledn & Clem-
ent 6773 (NY).
3. Calymperes venezuelanunm(Mitten) Broth-
Calymperes piovanoi Bizot, Rev. Bryol. Lich6nol. 40:
erusexPittier,Bol. Soc. Venez. Ci. Nat. 3: 360. 116. 1974. Type. Colombia. Leticia, 30 Jan 1969,
1936. Fig. 56. Piovano s.n. (PC).
76 Flora Neotropica

0 \q~00 (VO u", .

S~~ ~~~~~o 00

00 'O

o, O
000
56
O..

A OB
O o' O E
nu0

FIG. 6-5. alyipres5. Cvee'lueanun.A.

owrpiioiioflea. . Lafape. . Mrgnad ell a
mila. .Scto t ila. ,CD rmMaur 259i Bfo Mro 58.57 .plldt .A utieo
veeaielaf .Otieofgmieoslef .Mri adclsa ea hudr . agnadc l tmdef
E.Scto trug osa . olin o ea etin, n soin ea nrgiinscto. ,CD rm Zn letn

6 7 istp f ayiprscbev dlas Y) ,EF frmFtilrs.. clebr: un(is.5AB

57AB) b= . m(Fgs 5C-, 7CF)

FIGS. 56-57. Calymperes. 56. C. venezuelanutn. A. Lower portion of leaf. B. Leaf apex. C. Margin and cells at
midleaf. D. Section at midleaf. A, C-D from Maguire 24549m, B from Morton5987. 57. C. pallidutn. A. Outline of
vegetative leaf. B. Outline of gemmiferous leaf. C. Margin and cells at leaf shoulder. D. Margin and cells at midleaf.
E. Section through costa. F. Portions of leaf sections, one showing leaf margin in section. A, C-D from Leon & Clement
6773 (isotype of Calymperes cubense Williams, NY), B, E-F from Fendler s.n.. Scale bars: a = 1 mm (Figs. 56A-B,
57A-B); b = 0.1 mm (Figs. 56C-D, 57C-F).
Calymperaceae 77

Plants pale, soft, yellowish-green,often tinged Discussion. This species, althoughcommon in

with pink below, to 20 mm tall, leaf bases glossy muchof tropicalAfrica,is very rareelsewhere in
and conspicuous when dry because of broadly its range.Itis easily recognizedby its soft texture,
exposed cancellinae. Leaves folded and uncinate conspicuous cancellinae with ventrally
when dry,2-4 mm long, upperlaminalinear-lan- mammillosedistal cells, and lack of stereid cells
ceolate, aboutas long as or shorterthanthe much in the costa. CalympereserosumC. Mull. is sim-
broaderlower lamina;margins of upperlamina ilarbuthas conspicuousstereidbandsin the costa
slightly thickened,sharplyserrate;teniolaepres- andbearsgemmaeall aroundthe tipsof the costae
ent but sometimes obscuredby involutionof leaf instead of only ventrally.This species was pre-
margins; cancellinae sharply distinct, bulging viously known in the Americas as C. uleanum
ventrally, with conspicuously mammillosecells Broth.,in Africaas C. rabenhorstiiHampe & C.
distally on adaxialsurface;cells of upperlamina Mull., and in Asia as C. gemmiphyllumFleisch.
smooth to papillose dorsally, highly bulging- Calymperessubg. Somphoneurum, of which this
mammillose ventrally,ca. 6-10 pm diam.;costa speciesis the only Americanrepresentative, is de-
mostly highly papillose, lacking distinctbands of fined by the lack of dorsal and ventral fields of
stereid cells as seen in section; gemmniferous well-definedstereidcells in the costa,as viewed in
leaves not much differentiated, bearing gem- section, so that the cell net of the costa section
mae only on ventral surface of tips. Seta ca. 3.5 appearsspongy.
mm long; capsule 2 mm long; operculum 0.7
mm long. Spores 17-24 pm, papillose. Calyp- 5. Calymperes afzelii Swartz,Jahrb.Gewachsk.
tra 3.5-4 mm long, smooth or slightly rough- 1: 3. 1818. Type. Africa. Afzelius s.n. (BM,
ened above. herb.Hampe). Fig. 59.
Selected illustrations. Florschiitz (1964, fig.
51, as C. uleanumBroth.);Magill (1981, fig. 45: Calymperes donnellii Austin, Bot. Gaz. 4: 151. 1879.
Type. U.S.A. Florida: Caloosa, 1878, J. D. Smith & C.
9-16, as C. rabenhorstiiHampe& C. Mull.);Reese F. Austin s.n. (NY).
(1961, figs. 15-21, as C. uleanumBroth.);Reeseet Calymperes chlorosum Hampe, Vidensk. Meddel. Dansk
al. (1986, figs. 29-30, as C. getnmiphyllum Naturhist. Foren. Kj0benhavn 4, 1: 78. 1879. Type.
Brazil. Rio de Janeiro: Glaziou 9229 (holotype, BM?,
Fleisch.).
Distribution. (Fig. 58A). Cuba;Guadeloupe; n.v.; isotypes, BM, NY, PC, REN, S).
Calymperesbrittoniae Bescherelle, Ann. Sci. Nat. Bot. 8,
Colombia; Trinidad;Suriname;FrenchGuiana; 1: 278. 1896. Type. U.S.A. Florida: Polk Co.: Fort
Peru (Loreto); Brazil (Acre, Amazonas, Goias, Meade, Mar 1880, J. D. Smith s.n. (holotype, BM?,
Mato Grosso;Rond6nia);Ecuador;Africa;Mad- n.v., isotype, NY).
rufescens Bescherelle, Ann. Sci. Nat. Bot. 8,
agascar;Java; Sumatra;Sabah. At low altitudes Calymperes
1: 302. 1896. Type. "Antilles." Sur Quassia officinalis,
on tree trunks,particularlyon palms. Richard s.n. (holotype, BM; isotypes, NY, S).
Calymperes lindmanii Brotherus, Bih. Kongl. Svenska
Selected specimens exanined. CUBA. The type of Vetensk-Akad. Handl. 26, 3(7): 15. 1900. Type. Brazil.
CalymperescubenseWilliams. MatoGrosso:Lindman409 (holotype,H; isotypes, NY, S).
WINDWARD ISLANDS. GUADELOUPE: Petit Boury, CalymperesportoricenseRenauld& Cardot,Bull. Soc. Roy.
60 m, Questel892 (FH, LAF,NY). Bot. Belgique41(1): 57. 1902. Type. PuertoRico. East of
COLOMBIA. AMAZONAS: The type of Calymperes Santurce,Heller 4331 (holotype, PC?, n.v.; isotype, NY).
piovanoiBizot. Calymperes guianense Paris & Brotherus in Paris, Rev.
TRINIDAD. ST. ANDREW: Aripo Savanna, just N of Bryol. 33: 35. 1906. Type. French Guiana. Macouria,
Cumuto,100 ft., Crosby2254 (DUKE,LAF). 16 Oct 1904, Galliot s.n. (lectotype, H-BR).
SURINAME. Paramaribo,J. & P. Florschiitz605 Calymperes huallagense Brotherus, Hedwigia 45: 270.
(LAF, U). 1906. Type. Peru. Rio Huallaga . . . im Walde de
PERU. LORETO: Iquitos, 100 m, E. & P. Hegewald Cumbasso, Ule 2363 (holotype, H-BR; isotypes, BM,
6313 (herb. Hegewald). FH, M, MG, PC, S).
BRAZIL. ACRE: 15 km E of Rio Branco, Reese & Calymperes remirense Paris & Brotherus in Paris, Rev.
McPherson 13169 (INPA, LAF, MO, NY, US). AMAZO- Bryol. 33: 36. 1906. Type. French Guiana. Mont-Joly,
NAS: Along Rio Negro near Tapereira, Buck 2202b (NY). 22 Oct 1906, Galliot s.n. (holotype, REN; isotype,
MATO GROSSo: Mpio. de Camapua, 15 km SW of H-BR).
Figueirao, Vital 6400 (FLAS, LAF, SP). ROND6NIA: Just Calymperes reyi Paris & Brotherus in Paris, Rev. Bryol.
below first falls on Rio Pacaas Novos, Reese 13712 33: 36. 1906. Type. French Guiana. Macouria, 16 Oct
(INPA, LAF, NY). 1906, Galliot s.n. (lectotype, REN; isolectotype, PC).
78 Flora Neotropica

TROPICAL AMERICA FlokaNeotrop.a base map no 1

-----------
I~-~-I---T ---

B
I-^ ^ . '; <^' -! ......

_1-~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~i........
.... ..o s c c xe K u

FIG 58 Dstrbuton ofAmricn Clypers.

. .plfdumB.. fzeii

FIG. 58. Distributions of American Calymperes. A. C. palliduit. B. C. afzelii.

Calymperaceae 79

Plants yellowish to dark-greenor brown,dull, DOMINICAN REPUBLIC. SAMANA: 3.5 km E of Las

in low, thin turfsor gregariousandtufted,mostly Terrenas, Buck 8725 p.p. (NY).
PUERTO RICO. CaribbeanNational Forest, Luquillo
to ca. 10 mm tall. Leaves3-6 mm long, not much Division, 1.9 mi S of entrance to El Verde Biological
contortedwhen dry, ligulate to lanceolate from Station, 400-500 m, Reese 14876 (LAF, NY).
broader base; margins thickened, serrate to VIRGIN ISLANDS. ST. JOHN:Vic. Cruz Bay, Reese
14182 (NY). ST. THOMAS:St. Peter Mountain, Reese
coarsely toothed above; teniolae very conspicu- 14171 (NY).
ous; cancellinae distinct, sometimes glossy and WINDWARD ISLANDS. DOMINICA: St. George,
conspicuous when dry, their distal cells plane Trafalgar, E. & P. Hegewald 9357 (herb. Hegewald;
ventrally; cells at midleaf obscure, smooth or LAF). GUADELOUPE: Basse-Terre, De Sloover 23525 p.p.
finely papillose dorsally, bulging papillose ven- (NY). ST.LUCIA:Quilesse, Simmonds 222 p.p. (NY).
COLOMBIA. ANTIOQUIA: 3 km NNE de Caracoli,
trally, 4-5 pm diameter;costa smooth or papil- 700-890 m, Churchill et al. 14345 (HUA, LAF, NY).
lose; gemmiferous leaves often abruptly CHocO: Mun. de Acandi, Churchill & Arbeldez 16205
constrictedat apex and forming a narrow"pro- (NY).
boscis"with marginsrevolutearoundcosta,gem- VENEZUELA. BOLIVAR:45 km S of Cd. Piar, Pursell
mae borne only on ventralsurfaceof costa. Seta 8056 (LAF, PAC). MIRANDA:Parque Nac. Guatopo, Nee
17749 (NY). MONAGAS: 12.5 km NE of San Felix, 460 m,
4-5 mm long; capsule 2-2.5 mm long; opercu- Pursell 8417 (LAF, PAC).
lum 0.75 mm long. Spores 19-23 pm, finely TRINIDAD. ST.GEORGE: S slopes of Cerro del Aripo,
papillose. Calyptra4 mm long. 1400 ft., Crosby 2193 (DUKE, LAF).
Selected illustrations. Bartram(1949, fig. 36 GUYANA. Drainage of Takutu River, Smith 3325a
(NY).
A-E, as C. donnellii Aust.); Crum& Anderson SURINAME. Republiek Geijskes 42 (LAF, U).
(1981, fig. 114 G-L, as C. donnellii Aust.); FRENCH GUIANA. ST-LAURENT-DU-MARONI: Can-
Florschiitz(1964, fig. 52, as C. donnellii Aust.); tondeMaripasoula, ca. 6 km N of Sail, 200 m, Buck 18491
Reese (1961, figs. 77-83, as C. donnelliiAust.); (NY).
Reese et al. (1986, fig. 20). ECUADOR. ESMERALDA: 2-4 km SE of San Lorenzo,
Boom 2574 (LAF, NY). GALAPAGOS ISLANDS: Isla Santa
Distribution. (Fig. 58B). Pantropical.U.S.A. Cruz, 500 ft., WeberB-14402 (COLO, LAF).
(Florida);southernMexico; throughoutCentral PERU. LORETO: 12 km SW of Iquitos airport, Timme
America and the West Indies; northernSouth 4769 (herb. Timme, LAF).
America.Treetrunks,logs, rock,andsoil; mostly BRAZIL. ACRE:33 km SW of Rio Branco, Reese &
McPherson 13237 (INPA, LAF, MO, NY). AMAZONAS:
at low elevations in coastal and interiorforests
Along Rio Negro at cemetery Tapereira, Buck 2202
but collected up to ca. 700 m. (INPA, LAF, NY). BAHIA:Itabuna, Vital 8672 (LAF, SP).
MATOGROSSO:2 km NE of Base Camp, Royal Soci-
Selected specimens exanined. MEXICO. CHIAPAS: ety/Royal Geographic Society Expedition, 12?49'S,
14 mi. SE of Mapastepec, 400 ft., Kofron ID (LAF). 51?46'W, Harley 10805 (BM, LAF). PARA: Serra do
JALISCO:Near Sapotillo, Crum 623 (MICH, NY). Cachimbo, km 1011 along Cuiaba-Santarem hwy., 300 m,
NAYARIT: ca. 3 mi. E of Santa Cruz, Norris & Taranto Reese 16652 (INPA, LAF, NY). PARAIBA: Munic. Areal,
13119 (LAF, NY, TENN). OAXACA: NE side of Chiltepec, Vital 5436 (FLAS, LAF, SP). ROND6NIA: First rapids on
Santos 3852 (NY). VERACRUZ: 1.2 mi. S of Rio Tecolutla, Rio Pacais Novos, Reese 13681 (INPA, LAF, NY).
Reese & Pursell 4600 (LAF, US). RORAIMA: Km 517 along Manaus-Venezuela highway,
GUATEMALA. IZABAL:Rio Tameja, Steyerhnark 100 m, Buck et al. 1913 (INPA, LAF, NY). SANTACATA-
41813a (F). RINA:Porto Belo, Yano2195 (LAF, NY, SP). SAOPAULO:
BELIZE. TOLEDO: Punta Gorda, Whlites.n. (MO). Ilha do Cardoso, Vital 6160 (FLAS, LAF, SP).
NICARAGUA. ZELAYA: Near Rama,Kofron3C (LAF). BOLIVIA. BENI: Cachuela Ezperanza, 47 km NW of
COSTA RICA. Cocos ISLAND: Chatham Bay, Weber Guyaramerin,Reese 12822 (INPA, LAF, MO, NY, US).
4576 (F). LIMON:Along Laguna Penitencia, Steere CR-
121 (NY). Discussion. This moss was formerlyknown in
PANAMA. CANAL ZONE:Barro Colorado Island,
the Americasas C. donnellii Aust. With its small
Crosby 3937 (DUKE, LAF). DARIEN: Punta Guayabo
Grande, Antonio & Hahn 4222A (LAF, MO). obscure cells, prominentteniolae, and gemmae
BAHAMAS. GRANDBAHAMA:6-8 mi. E of Freeport, bore only ventrallyon the often modified leaf
6-15 m, Reese 11783 (LAF). tips, C. afzelii is very easy to identify.
CUBA. Sierrade Escambray,Pocs & Borhidi9004 (NY).
JAMAICA. ST.THOMAS: 1 mi. NNW of Bath, ca. 350
Calympereserosumis similarbut has largerleaf
cells and bearsits gemmae all aroundthe tip of
ft., Crosby 3236 (DUKE, LAF).
HAITI. Dpto. de la Granld'Anse,41 km S of Roseaux, the costa; further,the ventral distal cells of the
Buck 9081 (NY). cancellinaein C. erosum are markedlymammi-
80 Flora Neotropica

A C9

J.. 1 /[ E

I I2o%\ 420

a
b

A B 60o G

FIGS. 59-60. Calymnperes.59. C. afzelii. A. Leaf outline. B. Apex of gemmiferous leaf. C. Cells at midleaf. D.
Portions of leaf sections. E. Margin and cells at leaf shoulder. A from Steere 4962, B-D from Crum 623, E from Smith
& Austin s.n. (holotype of Calymperesdonnellii, NY). 60. C. erosurn.A. Outline of vegetative leaf, from lower portion
of stem. B. Outlines of gemmiferous leaves, from distal portions of stems. C. Margin and cells at leaf shoulder. D.
Margin and cells at midleaf. E. Portions of sections of leaves. F. Cells at midleaf from vegetative leaf (left) and
gemmiferous leaf (right). A from Grout, N. Amer. Musci Perf 322, B (leaf at left) from Simmonds 345a, (leaf at right)
from Simmonds 350a, C from Grout s.n., D-E from Bernoulli & Cario 134 (isotype of Calymperes emersum, NY), F
from Simmonds 345a. Scale bars: a = 1 mm (Figs. 59A-B, 60A-B); b = 0.1 mm (Figs. 59C-E, 60C-F).

lHose(at least in part,sometimes only along distal 1864. Type. Suriname. Paramaribo, Wullschlaegel
1231 (holotype, BR; isotypes, BM, NY).
edges of cancellinae), in contrastto those of C. Calymperes melinoni C. Miller in Bescherelle, Ann. Sci.
afzelii, which are plane distally. Nat. Bot. 8,1: 292. 1896. Type. FrenchGuiana. Melinon
s.n. (holotype, PC; isotype, NY).
6. Calymperes erosum C. Miiller, Linnaea21: Calymperessprucei Bescherelle, Ann. Sci. Nat. Bot. 8, 1:
182. 1848. Type. Suriname. Kegel s.n. (holo- 304. 1896. Type. South America. Amazon, Spruce 20
type, GOET;isotype, PC). Fig. 60. (holotype, BM; isotypes, NY, PC).
CalymrperesertersumrC. Miller, Bull. Herb. Boissier 5:
189. 1897. Type. Guatemala. Mazatenango, Bernoulli
Calymperes wullschlaegelii Lorentz, Moosstud. 160. & Cario 134 (lectotype, NY).
Calymperaceae 81

CalymperesbodeniiC. Miiller,Hedwigia39: 262. 1900. Crosby 10762 (LAF, MO). PANAMA: Along El Llani-Carti
Type. Brazil. Bahia: Boden s.n. (lectotype, NY; road, Crosby 10427 (LAF, MO).
isolectotype,H-BR). CUBA. PINARDELRio: Sierra de Cobra, E. G. Britton
Calymperes mammilliferum Crum & Steere, Bryologist 7293 (NY).
59: 248. 1956. Type. Puerto Rico. Km 102.9 on road to JAMAICA. ST. THOMAS:Bath, E. G. Britton 1000
Maunabo, Steere 4965 (MICH). (NY).
DOMINICAN REPUBLIC. ESPAILLAT: 7 km E de
Plants yellowish-green or darker,sometimes Gaspar Hemindez, Mejia & Pimental 23710A (NY).
PUERTO RICO. Sierra de Luquillo, Steere 6409
tinged below with pink or purple, mostly to 10 (NY).
mm tall,in thinto dense tuftsor turfs,oftenglossy WINDWARD ISLANDS. DOMINICA: St. George,
when dry due to exposed refractivecancellinae. Trafalgar, E. & P. Hegewald 9355 (herb. Hegewald,
Leaves somewhat dimorphic,mostly 3-4.5 mm LAF). GUADELOUPE: Terretear,Questel 3192 (NY). MAR-
TINIQUE:Duss 353 (NY). ST.VINCENT:Gibson Mt. and St.
long, involute and curved when dry, ligulate to Andrews, 500 ft., Elliott 5a (FH-Bartram,LAF).
lanceolatefrombroaderbase;marginsthickened, COLOMBIA. VALLE: Buenaventura, Killip 5993
serrate; teniolae conspicuous; cancellinae dis- (NY).
tinct, often glossy when dry, theirventraldistal VENEZUELA. MONAGAS: 5 km NNE of Maturin,
cells notably mammillose with the mammillae Pursell 9073 (LAF, PAC).
TRINIDAD. ST. ANDREW:Just E of Valencia, Crosby
pointing distally, (but mammillose cells some- 2068 (DUKE, LAF).
times restrictedto distal edges of cancellinae); GUYANA. 1 mi. E of Soesdyke, Pipoly 9291A (NY).
cells at midleaf mostly 5-7 x 7-10 pm, papillose SURINAME. NICKERIE:J. Florschiitz-de Waard &
Zielman 5247 (NY, U).
dorsally, mammillose-papilloseventrally;costa FRENCH GUIANA. ST-LAURENT-DU-MARONI:Can-
mostly smooth dorsally, mammillose-papillose tondeMaripasoula, ca. 6 km N of Sail, 200 m, Buck 18407
ventrally;gemmiferousleaves scarcelydifferen- (CAY, NY).
tiated or often very narrowlyacuminateand re- ECUADOR. MORONA: 2?18'S, 78?7'W, 1800 m, Or-
duced to naked costa distally, bearing often tega U. 425 (LAF, Q). GALAPAGOS ISLANDS: Isla Pinta,
650 m, Sipman M-374 (COLO, LAF, U).
sphericalclustersof gemmae all aroundon tip of BRAZIL. ACRE:170 km E of Rio Branco, Reese 13285
costa. Seta 3-4.5 mm; capsule 2.5-3 mm long;
(INPA, NY). AMAPA:Mun. Calcoene, Mori & Cardoso
operculum0.6-0.8 mm long. Spores finely and 17384 (NY). AMAZONAS: Rio Uatuma at confluence with
densely papillose,20-26 pm diam.Calyptra3.5- Rio Pitinga, Buck3112 (INPA, LAF, NY). MINASGERAIS:
4.5 mm long. ParqueFlorestado Rio Doce, Vital8885 (LAF, SP). PARA:
Serra do Cachimbo, Serra Maze, km 1224 on Cuiabi-
Selected illustrations. Bartram(1949, fig. 36, Santarem highway, Reese 16843 (INPA, LAF, NY).
I-K, as C. emersum C. Mill.); Crum & Anderson ROND6NIA: 2-4 km above first rapids on Rio Pacaas
(1981, fig. 114: A-F); Florschiitz (1964, fig. 50); Novos, Reese 13422 (INPA, LAF, NY). RORAIMA: Boca
Reese (1961, figs. 84-93); Reese et al. (1986, de Mata,216 km N of Boa Vista along Manaus-Venezuela
figs. 27-28). highway, Buck et al. 1967 (INPA, LAF, NY).
BOLIVIA. BENI:Vic. of Guyaramerin, Reese 12918
Distribution. (Fig. 64A). Pantropical.U.S.A. (INPA, LAF, NY).
(Florida); southern Mexico; Central America;
West Indies; northern South America. Tree Discussion. Plants of C. erosur are quite vari-
trunks,logs, rocks, soil, in forests, to ca. 650 m. able, ranging from forms with all the leaves broad
and
vegetative to those with the leaves all nar-
Selected specimens examined. MEXICO. VERACRUZ:
Zona de los Rios Uxpanapa y Solosuchil, Delgadillo M. rowly acuminate and gemmiferous (although the
3394 (LAF, MEXU). gemmae may have fallen). Although the extreme
GUATEMALA. The type of Calymperes emersum C. forms are very different in appearance, they are
Miill. still easy to identify by the ventrally mammillose
BELIZE. TOLEDO:Maya Mtns., Davidse & Brant
distal cells of the cancellinae and the gemmae (or
31884 (MO).
HONDURAS. ATLANTIDA: Vic. San Alejo, Standley their vestiges) borne all around on the tips of the
7925 (F). costae. Calymperespallidum, also with ventrally
NICARAGUA. ZELAYA: 6 mi N or NW of Col6nia San mammillose cancellinar cells, lacks stereids in its
Antonio, Stevens 9033 (MO). costa and bears gemmae only ventrally on the leaf
COSTA RICA. LIMON:Vic. Tortuguero, Steere CR-
10 (NY). PUNTARENAS: Isla de Canlo,Gotmez20001 (MO). tips; C. tenerum, also bearing gemmae all around
PANAMA. CANALZONE:1 a 2 mi. de Fuerte, Barsallo on the tip of the excurrent costa, lacks teniolae;
et al. 6 (LAF, MO). COLON:7 km SW of Portobelo, and C. othneri, also bearing gemmae all around
82 Flora Neotropica

bJ" o (oo
0-OCoO

II S ^ oooo
\ I
K E

\_>7

)O0oOo

FIG. 61. Calymlperesothmteri.A. Leaf outlines, gemmiferous leaf at right. B. Apex of gemmiferous leaf, gemmae
dehisced. C. Margin and cells at leaf shoulder. D. Portions of leaf sections. E. Gemmae. From Reese 16786 (LAF).
Scale bars: a = 0.1 mm (B); b = 1 mmn(A); c = 0.05 mm (C-E).

the costa tip, has plane ventral cells distally on its often robust;rhizoidslight-brown,scanty;stems
cancellinae and weak teniolae. Calymnperes to ca. 3 cm tall. Leaves lanceolate to acuminate
afzelii lacks mammillose cancellinar cells and from broad, often flaring bases, mostly 3.5-4.5
bears gemmae only ventrally on its leaf tips. mm long, involute and loosely curved-contorted
when dry, erect-spreadingand channeled when
7. Calymperes othmeri Herzog, Arch. Bot. Sao moist; margins of upper lamina thickened but
Paulo 1(2): 60. 1925. Type. Venezuela. In dem lacking stereids, entire to irregularly coarsely
Lauf der Caroni, Othmer 443 (holotype, JE, toothed (especially toward apex), margins of
n.v.; isotype, M). Fig. 61. lower lanina entire or serrate; teniolae irregu-
larlypresent,weakandincomplete, 1-2(-3) cells
Syrrhopodonrivularis Herzog, Arch. Bot. Sao Paulo 1(2): wide at shoulderswhen present;cancellinaecon-
59. 1925. Type. Brazil. Espirito Santo: Im salto Lure,
spicuous, ending in broad angles above, distal
Liitzelburgs.n. (JE). cells plane ventrally;median leaf cells obscure,
Calymperes rupicola P. Richards, Bull. Misc. Inform. 8:
323. 1934. Type. Guyana. Essequibo River, first falls, unipapillosedorsally,mammillose-bulgingven-
Richards 357 (holotype, BM; isotype, NY). trally,mostly 6 x 7 pm;gemmaeborneall around
on tip of excurrentcosta, small, reddish, mostly
Plants cespitose, sordid-green to blackish, only 4(-5) cells long. Sporophyteunknown.
Calymperaceae 83

Illustration. Reese (1990, figs. 1-7). Calymperespanamae Bescherelle, Ann. Sci. Nat. Bot. 8,
Distribution. (Fig. 64B). Endemic to South 1: 298. 1896. Type. Panama. 1850, Duchassaing s.n.
(holotype, BM, n.v.; isotype, NY).
America.Columbia;Brazil (Amazonas,Espirito
CalymperesperinvolutumC. Muller, Flora 83: 332. 1897.
Santo,Para,Rond6nia);FrenchGuiana;Guyana; Type. Venezuela. Tovar, Goebel s.n. (lectotype, here
Suriname;Venezuela. Not common but locally designated, NY).
abundanton water-splashedor intermittentlyin- Calymperesbahiense C. Miiller, Hedwigia 39: 261. 1900.
undatedrock along riversin forests at elevations Type. Brazil. Bahia: Boden s.n. (lectotype, here desig-
nated, NY; isolectotype, H).
to ca 450 m; typically along cataracts. Calymperes aberrans Paris, Rev. Bryol. 33: 37. 1906.
Type. French Guiana. Mtgne. des Peres, 22 Oct 1906,
Selected specimens examined. VENEZUELA. Galliot s.n. (holotype, REN; isotype, PC).
BOLIVAR: 6 km from Maniapure,Boom& Grillo 6522 Calymperes brevicaule Paris & Brotherus in Paris, Rev.
(NY). Bryol. 33: 37. 1906. Type. French Guiana. Macouria,
GUYANA.Matope,Graham348 p.p. (NY). 1906, Galliot s.n. (holotype, REN?, n.v.; isotypes, L,
SURINAME. NICKERIE DISTR.:km 117 on road to NY).
Amatopo,50 m, Florschiitz-deWaard& Zielman5613
(LAF,U).
FRENCH GUIANA. RiviereMana,TautPontinent, Plants dark-to yellowish-green,to 10 mm tall
Cremers7621 (FLAS,LAF). but commonly shorter, in low, dense turfs or
BRAZIL.AMAZONAS: AlongtheRioNegro,slopesand clumps. Leaves dimorphic, involute and con-
Buck2429 (INPA,LAF,NY).
summitof SerraCurcuriari, tortedwhen dry,mostly 2-3 mm long, oblong to
PARA:Serrado Cachimbo,Base A6reado Cachimbo,
Reese 16146 (INPA,LAF, NY). ROND6NIA: Firstrapids broadlylanceolate,bases not or only somewhat
of Rio PacaasNovos, Reese13680 (INPA,LAF,NY). broaderthanupperlamina(except in gemmifer-
ous leaves); marginsusually slightly thickened,
Discussion. This species was placed in the entire above, more or less serrateat shoulders;
synonymy of C. nicaraguenseby Reese (1961), teniolae usually conspicuous at shoulders and
but reinstatedby Reese in 1990. It differsfromC. above but sometimes weak or nearly lacking in
nicaraguense in bearing geimnae all aroundat some leaves; cancellinaedistinct, often truncate
the tip of the costa-as in C. erosum and C. distally; cells at midleaf mostly 7-8 pm diam.,
tenerum-while C. nicaraguenlse bears gemmae smooth or slightly papillose dorsally, bulging
only ventrallyon the leaf tip. Plantsof C. othmeri ventrally; costa smooth except towards apex;
are often very similar to those of C. erosum but gemmiferousleaves narroweddistally, often ter-
differ in the weak teniolaeandtheventrallyplane minatingin a distinctgemma-receptaclebearing
distal cells of the cancellinae. gemmae in hairbrush-liketufts on ventral sur-
face, gemmnareceptaclesometimes folded down
8. Calymperes palisotii Schwaegrichen,Spec. adaxially.Seta ca. 3 mm long; capsule to 2 mm
Musc. Suppl. 1(2): 334. 1816. Type. In regno long; operculumca. 0.5 mm long. Sporesmostly
OwareAfricae. Palisots.n. (lectotypeselected 28-35 pm, finely papillose. Calyptra3-3.5 mm
by Edwards 1980, S). Fig. 62. long.
Selected illustrations (all as C. richardii C.
CalymperesrichardiiC. Muller,Syn.musc.frond1: 524. Mill.). Bartram(1949, fig. 36, F-H); Crum &
1849. Calymtperespalisotii Schwaegrichensubsp.
richardii(C. Muller)S. Edwards, J. Bryol. 11: 81. Anderson (1981, fig. 115, A-G); Florschiitz
1980. Type. Brazil.Perambuco: Gardner47 (lecto- (1964, fig. 49); Reese (1961, figs. 35-43).
type,BM;isolectotype,OXF). Distribution. (Fig. 65A). U.S.A. (Florida).
CalymperesbreuteliiBescherelle,Ann.Sci. Nat. Bot. 8, Widespreadin theneotropicsandtropicalAfrica;
1: 278.1896. Type.Antilles.St.Thomas:1841,Breutel westernAsia. Tree trunks,dead wood, rocks, at
s.n. (holotype, BM, n.v.; isotype,NY).
CalymperesguadeloupenseBescherelle,Ann. Sci. Nat.
low elevations;coastal and inlandforests.
Bot. 8, 1: 285. 1896. Type. Guadeloupe.Maries.n.
(holotype,BM, n.v.; isotype,NY). Selected specimens examined. MEXICO. OAXACA:
Calympereshexagonumn Bescherelle,Ann.Sci. Nat.Bot. Along road to dam on Rio Tonto, 50 ft., Reese & Pursell
8,1:286.1896. Type.Antilles.St.Thomas:1782-1789, 4541 (LAF, MEXU, US). PUEBLA: Nora, 10 Sep 1911,
Richards.n. (holotype,BM, n.v.; isotype,S). Nicolas s.n. (NY). QUINTANA Roo: 1 km N de Felipe
CalympereshookeriBescherelle,Ann.Sci. Nat.Bot. 8, 1: CarrilloPuerto, Cardenas 923 (NY). TABASCO: 1 mi from
287. 1896.Type.Antilles.Antigua:[withoutfurtherdata] Teapa, Sharp & McFarland 9117 (LAF, TENN).
(PC). TAMAULIPAS: 11.7 mi S of Mante, Reese & Pursell 5079
84 FloraNeotropica

(LAF, MEXU, US). VERACRUZ: 0.5 mi W of Nautla, Reese sea level, Gradstein5761 (LAF,U).
& Pursell 4651 (LAF, MEXU, US). ECUADOR.GALAPAGOS ISLANDS: Isla Pinta,Sipman
BELIZE. Lime Cay, Spellman & Stoddart B128a M-335 (COLO,LAF,U).
(MO). TOLEDO: Near Punta Gorda, Gentle 7503 (MO). PERU. LORETO: Near Iquitos, 100 m, P. & E.
HONDURAS. COL6N: 4.5 mi NE of Trujillo, Saunders Hegewald6332 (herb.Hegewald,LAF).
913 (MO). BRAZIL.AMAPA: Mun.de Macapa,8 kmSof Macapa,
NICARAGUA. ZELAYA:Corn Island, Stevens 19995 Mori17413(NY).AMAZONAS: InManaus,Griffin021316
(MO). (FLAS, LAF). BAHIA:Mun. EntrRios, Boom & Mori 1000
COSTA RICA. GUANACASTE: Bowers 315-a (LAF, (NY). FERNANDO DENORONHA: Charles Darwin 438 (NY).
TENN). LIM6N: Vic. Tortuguero, Steere CR-200 (NY). PARA: SerradoCachimbo, SerraMaze,km.1208onCuiabi-
PUNTARENAS: 5 Sep 1969, Stone s.n. (LAF). Santaremhwy., Reese 16752 (INPA, LAF, NY). Rio
PANAMA. BOCASDELTORO: Crosby 4083 (LAF, GRANDEDO NORTE: Natal, Vital 5415 (FLAS, LAF, SP).
MO). CANALZONE:Summit, Tyson 1177 (LAF, NY). RONDONIA:PortoVelho,Reese12711(INPA,LAF,NY).
COL6N: Barsallo et al. 20 (LAF, MO). HERRERA:NE of
Paris, McDaniel 8006 (NY). PANAMA:Just outside of Discussion. This species was formerlyknown
Panama City, Allen 4902 (MO, NY).
BAHAMAS. GRANDBAHAMA:
in the Americasas C. richardiiC. Mill., which
Lucaya National Park,
Reese 17996 (LAF, NY, U). LITTLEINAUGUA ISLAND:10 was shown by Edwards(1980) to be synonymous
Apr 1977, Felder s.n. (LAF). NEWPROVIDENCE: E. G. with C. palisotii Schwaegr. Although Edwards
Britton 6625 (NY). recognized the American form as C. palisotii
CUBA. ORIENTE:Prov. Holguin, vic. Cerro Galano,
Pocs & Catastus90233/c (NY). subsp. richardii, the distinctions he noted be-
CAYMAN ISLANDS. LITTLECAYMAN: Vie. W end tween African and American specimens are
of Charles Bight, Proctor 35175 (NY). minor and inconsistentlypresent.This is a very
JAMAICA. ST. MARY: Castleton Gardens, 23 Jun common moss of low tropicalforests and is par-
1968, Griffin, 111s.n. (FLAS, LAF). ticularly common in coastal regions, but much
HAITI. Bayeux, near Port Margot, Nash 160 (NY).
less frequentaway from the seacoast except in
DOMINICAN REPUBLIC. LA ALTAGRACIA: 24 km
SE of Otra Banda, Reese 15565 (LAF, NY). some cities, such as Manaus,Brazil, whereit is a
PUERTO RICO. ISLADECULEBRA: Mt. Resaca, 200 common moss on trees along streets. The broad
ft., 20 Feb 1971, Valentines.n. (LAF). cancellinae,often truncatedistally, and common
VIRGIN ISLANDS. ST. JOHN:Vie. Cruz Bay, Reese
14186 (NY). ST. THOMAS: Estate Dorothea, Reese 14191
presence of hairbrush-liketufts of gemmae on
(LAF, NY). TORTOLA: N. L Britton & Shafer 722 (NY). gemma receptacleson the leaf tips, are distinc-
LEEWARD ISLANDS. ANTIGUA: [without data] Rose tive. The gemma receptaclein many collections
et al. 3687 (NY). MONTSERRAT: Gage's Mt., Shafer 925 tends to be folded down adaxially when dry, as
(NY). SABA:Mary's Point, van Slageren 254M (NY). ST. in thepaleotropicalspecies C. crassinerve(Mitt.)
BARTHELEMY: Grande Saline, Le Gallo 433 (NY). ST.
EUSTATIUS: Gilboa Hill, van Slageren 39M (NY).
Jaeg. and C. motleyiMitt. in Dozy & Molk.
WINDWARD ISLANDS. BARBADOS: St. Peter, 24
Apr 1968, Pearce s.n. (NY). DOMINICA: Roseau, E. & P. 9. Calymperes tenerum C. Miiller,Linnaea37:
Hegewald 9298, (Herb. Hegewald, LAP). GUADELOUPE: 142.1872. Type.India.Im Calcuttum,Kurzs.n.
Capesterre, Le Gallo 1082 (NY). MARTINIQUE: Garden of
(lectotype,here designated,BM). Fig. 63.
Tivoli, M. & H. Stehle 4040 (NY). ST. LUCIA:Cashies,
Simmonds 259 (NY).
COLOMBIA. ANTIOQUIA: Mun. Chigorod6, 7?39'N, CalymperesnashiiWilliams,Bull. TorreyBot. Club47:
391.1920.Type.Haiti.BayeauxnearPortMargot,Nash
76?33'W, Churchill et al. 17088 (LAF, NY). CAQUETA: 51
San Vicente del Caguan, 2?18'N, 74?48'W, Churchill & (NY).
Betancur 17037 (LAF, NY). COoc6: Mun. de Acandi.
Churchill & Arbdelez 16239, 16247(NY). Plants small, mostly 2-3 mm tall, pale-green,
VENEZUELA. APURE:Distr. San Fernando,opposite in loose or dense tuftsand turfs. Leavesinvolute
Isla Peladura, Davidse & Gonzalez 12163 A-2 (LAF, and
MO). MONAGAS: 2 km WNW of Jusepin, Pursell 8338 slightly contortedwhen dry, oblong-lanceo-
(LAF, PAC). SUCRE: late, 2-2.5 mm long, lacking evident shoulders;
Playa Colorado, Pursell 9418 (LAF,
PAC). margins entire, uni- or bistratose above, com-
TRINIDAD. ST. ANDREW:just E of Sangre Grande, posed of broadareasof squaregreen cells exte-
Crosby 2046 (DUKE, LAF). TOBAGO:Scarborough, rior to cancellinaein leaf base; teniolae lacking
Broadway 4742 (NY). or rarely vestigial; cancellinae distinct, small,
GUYANA. Pomeroon River, Bartlett 8676 (NY).
SURINAME. Albina, Marowijne, J. & P. Florschiitz more or less truncatedistally;cells at midleaf 6-8
548 (LAF, U). pm diameter, finely papillose dorsally,
FRENCH GUIANA. CAYENNE: AroundMontabo Mt., mammillose ventrally; costa papillose-rough-
Calymperaceae 85

62

c A
A B

E E

FIGS. 62-63. Calymperes. 62. C. palisotii. A. Outlines of vegetative leaves. B. Outline of gemmiferous leaf (dorsal
view, gemmae lacking). C. Apex of gemmiferous leaf with gemmae, dorsal view. D. Margin and cells at leaf shoulder.
E. Margin and cells at midleaf. F. Portions of leaf sections, one showing the leaf margin in section. A (leaf at left) from
N. L. Britton et al. 10 (NY), (leaf at right) from McFarlin 524 (NY), B from McFarlin 524 (NY), C from N. L. Britton
et al. 10 (NY), D-E from Austin & Smiths.n. (NY), F from E. G. Britton 7292 (NY). 63. C. tenerum. A. Leaf outlines.
B. Outline of leaf base. C. Tips of gemmiferous leaves, one with characteristicspherical cluster of gemmae. D. Margin
and cells at leaf shoulder. E. Margin and cells at midleaf. F. Portionof section of leaf. A-B, F from Phillips 100, C from
Nash 51 (holotype of Calymperes nashii, NY), D from N. L. Britton et al. 1860 (NY), E from Small & Mosier 5158
(NY). Scale bars: a = 1 mm (Figs. 62A-C, 63A-C); b = 0.1 mm (Figs. 62D-F, 63D-F).

ened, especially distally; gemmiferous leaves nashiiWilliams);Reese etal. (1986, figs. 31-32).
with costa excurrent and bearing gemmae all Distribution. (Fig. 64B). Pantropical, but
aroundat apex in conspicuouspale spheres.Spo- very rare in the neotropics. U.S.A. (Florida);
rophytesunknownin the Americas. Mexico (Veracruz); Haiti; Bahamas; Puerto
Illustrations. Crum and Anderson(1981, fig. Rico; Brazil (Sao Paulo). Tree trunks, includ-
115: H-K, as C. nashii Williams);Magill (1981, ing mangroves, in coastal areas, often along
fig. 45: 1-8); Reese (1961, figs. 22-29, as C. ocean beaches.
86 Flora Neotropica

TROPICAL AMERICA Floa Neotipca base map no I

FIG6.iriuonoAercaCayeeACeos C iei(lines)Cteneru(dos)
~~A ~ ~ ii ..
A

...--,'--- .---....
/
9
~,;
,
ir~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

,o,~~~~~~~?I

.--F-------- ~ ~ ~ ~ ~~ ----------------

FIG. 64. Distributions of American Calymtperes.A. C. erosurnt.B. C. othmneri(lines); C. tenerturn(dots).

Calymperaceae 87

Specimens examined. MEXICO. VERACRUZ:

2 mi. SE Trinidad. A rare endemic of northern South
of Boca del Rio, sea level, Reese& Pursell4389 (LAF,
America;knownfromonly a few collections. On
MEXU,US).
BAHAMAS. GRANDBAHAMA: tree trunksand lianas,500-700 m.
Lucaya National Park,
Reese17988 (LAF,NY).
HAITI.The type of C. nashii. Specimens exaniined. TRINIDAD. ST.GEORGE: trail
BRAZIL. SAOPAULO: Ilhado Cardoso,Vital12869, to More Bleu, Crosby 2102 (DUKE, LAF).
Vital& Giancotti13674 (LAF, SP). Mun.de Peruibe, FRENCH GUIANA. ST-LAURENT-DU-MARONI: Can-
Guarau,sea level, Yano& Mello11799 (LAF,SP). tonde Maripasoula,ca.6 kmN of Sauil,400 m Buck18931
(CAY, NY), 18957 (NY).
Discussion: This species was formerlyknown
in the Americasas C. nashii Williams. Although Discussion. The dimorphicleaves with flaring
it is widespreadand common in the paleotropics and stronglyborderedshouldersand gemmae in
it is very rarein the Americas, where it is only distinctreceptaclesare distinctivefor this moss.
known from a few collections. It is likely to turn Calymperes mitrafugaxand C. platyloma are
similarbuthavebroaderbordersat the leaf shoul-
up elsewhere in the neotropics and should be der and papilloseleaf cells.
sought in coastal sites, especially near ports, as
its Americandistributionindicatesthatithasvery
11. Calymperes platyloma Mitten,J. Linn. Soc.,
likely been introducedfromthe paleotropics.The Bot. 12: 128. 1869. Type. Guyana.Appun819
combinationof pale golfball-likespheresof gem-
mae borneall aroundon the tips of the excurrent (holotype,NY; isotype, BM). Fig. 67.
costae, lack of teniolae, small rectangularcan- Plants small,yellowish-green,gregarious;rhi-
cellinae, and broad bands of squaregreen cells
zoids light-brown; stems only to 6 mm tall.
comprisingthe basal leaf marginsis very distinc- Leaves lanceolate to acuminate, ca. 3.5-5 mm
tive.
long, involute and secund when dry, erect and
channeledwhen moist, marginsof upperlamina
10. Calymperes smithii Bartram,Bull. Torrey
Bot. Club 66: 223. 1939. Type. Brazil-Guyana thickened, showing stereids in section, evenly
serrulateby projecting cell angles, margins of
boundary.AkaraiMountains,Smith2985 (ho- lower lamina sharply and irregularly serrate-
lotype, NY; isotypes, BM, FH, U). Fig. 66. toothed; teniolae lacking but leaf shoulders
Plants olive-green, rhizoids reddish-brown; strongly borderedwith to 16 rows of elongate,
thick-walledcells; cancellinae truncatedistally;
stems simple, to 1 cm tall. Leavesmostly ca. 3.5 median leaf cells thick-walled,rounded-rectan-
mm long, curvedwhen dry,dimorphic,vegetative
leaves oblong-elliptic from broaderbase, gem- gular to elliptic, mostly ca. 12 x 14 pm,
miferous leaves reducedto narrowwings along mammillose-papillose dorsally, mammillose
ventrally;gemmae infrequent,ventral on tip of
costa, margins of upper lamina slightly thick- costaof slightlymodifiedleaves. Seta 3 mm long;
ened, rarely unistratose, narrowly winged and capsule ca. 1.8mm long; operculumca. 0.75 mm
coarselyserrate-spinoseabove, sometimesshow- long.Sporesfinelyroughened,18-22 pm. Calyp-
ing stereids in section, marginsof lower lamina tra ca. 3.5 mm long.
mostly entire or somewhat serrateat shoulders; Illustrations.Florschiitz(1964, fig. 47); Reese
teniolae lacking but shouldersborderedby up to
(1961, figs. 44-51).
six (mostly fewer) rows of elongatethick-walled Distribution. (Fig. 68A). Endemic to northern
cells; cancellinaebroadabove; medianleaf cells SouthAmerica.Brazil (Amazonas,Bahia);Guy-
in verticalrows, roundedto oblong, mostly 8-11
x 8 pm, bulging dorsallyand ventrally;gemmae ana;Suriname;FrenchGuiana.
borne in distinct gemma receptacle on ventral Selected specimens examined. GUYANA. Upper
surface of costa. Sporophytesunknown. Demerararegion, Pipoly 8912 (NY).
Illustrations. Florschiitz(1964, fig. 45); Reese SURINAME. Stonemansberg, J. & P. A. Florschiitz
1837(LAF, U).
(1961, figs. 68-74). FRENCH GUIANA. ST-LAURENT-DU-MARONI: Can-
Distribution. (Fig. 65B). Brazil-Guyana ton de Maripasoula,ca. 6 km N of Saiil, 200-300 m, Buck
boundary;Guyana; Suriname;French Guiana; 18765, 18897 (CAY, NY).
88 Flora Neotropica

TROPICALAMERICA FloraNeotrop.cabase rap no 1

-- -- - -- -

A .........

-D
FIG. 6~B5.I, u o i l (ie;.--- .A. CplsiB..

---------..
---------7

FI.'5 Dsriuios fAmrca almers.A C alsti. .C.~nihz lne CIirrmn(ot)

Calymperaceae 89

66 E DoO
C 0000 0000
00

0 0

O F ?

o o 0
0oo

G' OOo 900000

00/

b -

6'7

00

FIGS. 66-67. Catymperes. 66. C smithii. A. Outlines of vegetative leaves. B. Outline of gemmiferous leaf. C.
Leaf base. D. Margin and cells at leaf shoulder. E. Cells at midleaf. F. Section of leaf. From Smith 2985 (FH, isotype
of C. smithii). 67. C. platylomta.A. Leaf outline. B. Outlines of leaf bases. C. Margin and cells at leaf shoulder. D.
Portions of leaf sections. E. Cells at midleaf. From Appun 819 (holotype of Calymperesplatyloma, NY). Scale bars: a
= 1 mm (Figs. 66A-C, 67A-B); b = 0.1 mm (Figs. 66D-F, 67C-E).
90 Flora Neotropica

TROPICAL AMERICA FloraNeotopca base map no 1

FIG. 68. Distributions of American Calymperes. A. C. patylotia (line and dot); C. mitrafugax (triangles). B. C.
guildingii.
Calymperaceae91

BRAZIL. AMAPA:Mun. Calqoene, Mori & Cardoso Discussion. This rarespecies is generallysim-
17293 (NY). AMAZONAS: 62 km W of Humaitai,Fife et al. ilar to C. platyloma but differs, among other
4282 (INPA, LAF, NY); Rio Urubu, km 115 on Manaus-
Caracarai road, Griffin et al. 898 (FLAS, LAF). BAHIA: ways, in its smallerandfinely papilloseleaf cells,
Mun. Una, Maruim, 33 km SW of Olivenca, Boom et al. in contrastto the larger and strongly papillose
818 (NY). cells of C. platyloma.

Discussion. This species is quite rare as it is 13. Calymperes bartramii Reese, Bryologist
known from only a handful of specimens, most 64: 136. 1961. Type. Colombia. Vaupes:
of them collected rather recently. Its broadly bor- Cachivera Palito, Rio Kananari, Schultes &
dered leaf shoulders with jagged margins are Cabrera 13199 (holotype, CANM; isotype,
distinctive. Calymperes smithii and C. FH-BARTRAM). Paratype. Venezuela.
mitrafitgax are somewhat similar. Bolivar, near Rio Tirica, Steyermark 75543
(FH-BARTRAM,NY). Fig. 70.
12. Calymperes mitrafugax Florschiitz, Mosses
of Suriname 1: 129. 1964. Type. Suriname. Plants pale-green; rhizoids reddish-brown;
Along Wane Creek, Lanjouw & Lindeman 521 stems to ca. 9 mm tall. Leavesinvolute-subulate,
(holotype, U; isotype, LAF). Fig. 69. mostly 4.5-5 mm long, strongly inrolled and
contorted when dry; margins of upper lamina
Plants green to yellowish-green, in turfs; rhi- entire,mostlystronglyinfoldedfromapex to just
zoids reddish-brown; stems to 1.5 cm tall. Leaves above shoulders, unistratose above, thickened
acuminate, to 5 mm long, involute and curved below, sometimes showing stereids in section,
when dry, erect-spreading when moist; margins margins of lower lamina sharply serrate-serru-
of upper lamina thickened, mostly entire, mar- late; teniolae usually distinct, of elongate cells
with often thickened,pitted and irregularwalls;
gins of lower lamina sharply serrate-toothed; ten-
iolae lacking but leaf shoulders strongly bordered cancellinaeirregularlyroundedor truncatedis-
by many rows of elongate thick-walled cells; tally; median leaf cells very thick-walled,
cancellinae irregularly truncate distally; median rounded to ellipti9 mostly 10-12 pm, smooth
leaf cells rounded to elliptic, mostly ca. 12 pm dorsally,bulgingto mammilloseventrally;gem-
mae infrequent,on ventraltips of leaves. Seta 3
long, finely papillose dorsally, mammillose ven-
mm long;capsule 1.5 mm long; operculum1 mm
trally; gemmae borne ventrally on tips of undif-
ferentiated leaves. Seta 3 mm long; capsule 1.5 long. Spores smooth, 22 pm. Calyptra 4 mm
mm long. Spores finely granular, 18-23 pm. long.
Illustration. Reese (1961, figs. 106-111).
Calyptra 3.5 mm long.
Distribution. (Fig. 65B). Endemicto northern
Illustration. Florschiitz (1964, fig. 48).
South America. Brazil (Amazonas); Colombia
Distribution. (Fig. 68A). A rare endemic in
northern South America. Brazil; French Guiana, (Vaupes); Venezuela (Bolivar). Very rare; on
trees, to 1000 m elevation.
Peru; Suriname; Venezuela. On tree trunks in
lowland forests.
Specimenexamined.In additionto the type material,
C.bartramiiis knownfromonlyoneadditionalspecimen:
Specimens exaniined. VENEZUELA. BOLivAR:SW- BRAZIL. AMAZONAS: Rio Lages, km 130 along
facing slopes of Chimanta-tepui, Steyermark75429 (NY). Manaus-Caracarai road, Griffin, 11I et al. 765 (FLAS,
SURINAME. The type. LAF).
FRENCH GUIANA. ST-LAURENT-DU-MARONI: Can-
ton de Maripasoula, ca. 6 km N of Saul, 200 m, Buck18469
Discussion. The subulate upper lamina with
(CAY, NY).
PERU. SAN MARTIN: Prov. Lamas, Strasse margins tightly inrolled wet and dry, and C.
Yurimaguas-Tarapoto km 112, Frahm etal. 1746 (NY). guildingii-liketeniolae,are distinctive.
BRAZIL. AMAZONAS: CEPLAC Reserve, km 29 on
Manaus-Itacoatiara road, Prance 20987 (INPA, LAF, 14. Calymperes guildingii Hooker & Greville,
NY). MATAGROSSO:Aripuana, near Humboldt Center,
Lleras & Lima P18265 (INPA, NY). PARA:Serra do EdinburghJ. Sci. 3:223. 1825. Type. Antilles.
Cachimbo, cataracts on Rio Curua, km 877 on BR 163, St. Vincent: Mt. St. Andrews, Guilding s.n.
Reese 16853 (INPA, NY). (holotype,BM-K;isotype, NY). Fig. 71.
92 Flora Neotropica

Ai 9

69 aa _____
b
cz _

B L0
c ____C
ooooo

57Q0v uOO oooo00

70

FIGS. 69-70. Calymperes. 69. C. mitrafugax. A. Leaf outlines. B. Margin and cells at leaf shoulders. C. Cells at
midleaf. From Lanjouw& Lindenan 52 (isotype of Calymperesmitrafugax, LAF). 70. C. bartramii. A. Leaf outlines.
B. Margin and cells at leaf shoulders. C. Cells at midleaf. D. Leaf sections. From Schultes & Cabrera 13199 (holotype
of Calymperes bartramii, CANM). Scale bars: a = 0.05 mm (Fig. 69B); b = 1 mm (Fig. 70A); c = 1 mm (Fig. 69A); d
= 0.05 mm (Figs. 69C, 70B-D).
Calymperaceae 93

Calymperes cruegeri C. Miiller, Syn. musc. frond. 1: 527. Hegewald, LAF). GUADELOUPE: [without locality]
1849. Type. Trinidad. Monte Tocuche, 4 Dec 1847, I'Herminier s.n. (NY). MARTINIQUE:Montagne Pelee,
Criiger s.n. (ectotype, here designated, NY). Crosby & Crosby 4710 (MO, NY). ST. LUCIA:Quilesse,
Syrrhopodon badius Schimper in Bescherelle, Ann. Sci. Simmonds
222 p.p.(NY).
Nat. Bot. 6, 3: 197. 1876; Calymperes badium COLOMBIA. ANTIOQUIA: 210 km N of Medellin, 280
(Schimper in Bescherelle) Brotherus in Engler & m, Churchill etal. 14890 (LAF, NY). NORTEDESANTAN-
Prantl, Nat. Pflanzenfam. 1(3): 379. 1901. Type. An- DER:CordilleraOriental, E of Rio Valegrai,6000-7000 ft,
tilles. Guadeloupe: Vallee de Saint Louis, 1868, Husnot Steere 7403 (LAF, NY).
s.n., PI. des Antilles 195 (holotype, BM, n.v; isotype, VENEZUELA. AMAZONAS: Dpto. Rio Negro,
NY). 00?50'N, 66?10'W, Buck 11336 (NY). ISLAMARGARITA:
Calymperes guadeloupense Brotherus in Urban, Symb. Cerro Copey, Sugden 1098 (NY). NUEVAESPARTA: La
Sierrita,900 m, BermidezNE-00778 (LAF,NY).
antill. 3:423. 1903. Type. Antilles. Guadeloupe: Morne
TRINIDAD.ST.GEORGE:
Hirondelle, Duss 241 (holotype, H-BR; isotype, NY). NorthernRange,2600 ft.,
Crosby2318 (DUKE,LAF).
GUYANA. Basin of EssequiboRiver, Head Falls,
Plants glossy, sordidto light-greenorblackish, Smith2105 (LAF).
in dense, stiff clumps; rhizoids pale- reddish- SURINAME.Tafelberg,Maguire24549M2(NY).
brown, scanty; stems to 2(-5) cm tall. Leaves FRENCH GUIANA. ST-LAURENT-DU-MARONI:
Can-
subulate from oblong, sheathing base, mostly ca.6 kmN of Saiil,400m, Buck18932
tondeMaripasoula,
5-6(-7) mm long, upper auninatightly involute (CAY,NY).
and curled-twistedwhen dry; marginsof upper Slopes and summit of Serra
BRAZIL. AMAZONAS:
Curicuriari,450 m, Buck 2504 (INPA, LAF, NY).
laminaentireor with scatteredserrations,greatly RORAIMA:216km N of BoaVista,800m, Bucketal. 2018
thickened, showing stereids in section, margins (INPA, LAF, NY).
of lower lamina serrate or entire; teniolae 4-6
cells wide at shoulders,of elongate,thick-walled Discussion. This species is related to C.
cells with pitted and somewhat sinuous walls; bartramiiandC. nicaraguense.The infoldedleaf
cancellinae ending in acute angles distally; me- margins of the former, which also has thick-
dian leaf cells square to elliptic or rectangular, walled sinuous cells in its teniolae, easily distin-
thick-walled,mostly ca. 7-14 x 7-9 um,smooth guish it from C. guildingii, and the imperfect
dorsally, mammillose to mammillose-papillose teniolae of C. nicaraguense distinguish it from C.
ventrally;gemmae infrequent,on ventraltips of guildingii. See discussion under C.
leaves. Seta 3-4 mm long; capsule 2.3 mm long; nicaraguense.
operculum 1 mm long. Spores24-35 pm, fmely
roughened.Calyptra4-5 mm long. 15. Calymperes nicaraguense Renauld & Car-
Selected illustrations. Florschiitz(1964, fig. dot, Bull. Soc. Roy. Bot. Belgique 33(2): 117.
54); Reese (1961, figs. 101-105). 1894. Type. Nicaragua. Escondido River,
Distribution. (Fig. 68B). Endemic to the neo- 1892, Richmond s.n. (holotype, PC; isotype,
tropics: Jamaica;Hispaniola;PuertoRico to St. NY). Fig. 72.
Kitts and south throughthe islands to Trinidad;
Brazil (Amazonas, Roraima);Colombia (Anti- CalymperescarionisC. Muller,Bull. Herb.Boissier5:
189. 1897. Type. Guatemala. Bernoulli & Cario 91
oquia, Norte de Santander);Guyana;Suriname; (lectotype,here designated,NY).
French Guiana; Venezuela (Amazonas, Nueva
Calymperes bolivianum Williams, Bull. New York Bot.
Esparta).Growing in stiff tufts, mostly on rock Gard. 3: 115. 1903. Type. Bolivia. Mapiri, Williams
but also on lianas, tree trunks,logs, humus,and 1804 (holotype, NY; isotypes, H-BR, BM-K, MICH).
soil, from ca. 300 to 2000 m. heribaudiiParis& Brotherus,Rev.Bryol.35:
Calymperes
49. 1908. Type. Panama. Chiriqui: In silvula Boquete
Selected specimens examined. JAMAICA. ST. pr. David,Fr. Sep 1906, Helidns.n. (holotype,REN;
THOMAS: WNW Corn Puss Gap, 1700-2200 ft., Crosby isotypes, M, NY).
3255 (DUKE, LAF).
PUERTO RICO. CaribbeanNational Forest, Luquillo Plants light-green, glossy, often matted with
Division, El Yunque Recreation Area, slopes of Mt. rust-colored rhizoids below, in cushions, tufts or
Britton, below 940 m, Reese 14707 (LAF, NY).
LEEWARD ISLANDS. ST. KITTS:[without locality] gregarious; stems 10-20 mm tall. Leaves subu-
1841, Breutels.n.(NY). late, mostly 4-5 mm long from broad base with
WINDWARDISLANDS.DOMINICA: St. David:Em- conspicuously flaring shoulders, involute and
erald Pool, 500-600 m, E. & P. Hegewald9334 (herb. curled-contorted when dry, cancellinae often
94 Flora Neotropica

PonO oO

B 0
000 0 Do
7Z. nD
b~oon eo o

oOcO~oOO

05
~ C

A E

FIGS. 71-72. Calymnperes.71. C. guildingii. A. Leaf outline. B. Margin and cells at leaf shoulder. C. Cells at
midleaf. D. Portion of section of leaf. A from Smith 1455, B from Guilding s.n. (holotype of Calymperesguildingii,
BM-K), C from Steere 4079, D from Husnot, PL des Antilles 135. 72. C. nicaraguense. A. Leaf outline. B. Margin
and cells at leaf shoulder. C. Cells at midleaf. D. Portion of leaf section. E. Dry capsule with operculum enclosed by
calyptra; the operculum is gripped at its rostrum by the calyptra and held away from the mouth of the capsule; spores
escape through the gaping fissures of the dry calyptra. F. Sporophyte with calyptra removed and operculum detached.
A from Gentle 3531, B-F from Richtmonds.n.(isotype of Calymperesnicaraguense, NY). Scale bars: a = 1 mm (Figs.
72E-F); b = 1 mm Figs. 71A, 72A); c = 0.1 mm (Figs. 71B-D, 72B-D).
Calymperaceae 95

conspicuous and glossy when dry; leaf margins with broad, flaring bases, C. nicaraguense is
thickened, finely serrate by projecting cells closely relatedto andmay resembleC. guildingii
above, serrateat shoulders;teniolaevariable,ex- Hook. & Grev. The latter species has strong
tending irregularly downward into shoulders teniolae composed of sinuous, pitted, thick-
from above or lacking;cancellinaedistinct;cells walled cells, very distinctin the leaf shoulders.
at midleaf ca. 8 pm diam.,mammilloseventrally, Calymperesnicaraguenseis apparentlyrare as
smooth or somewhat papillose dorsally; costa relatively few specimens of it have been col-
rough distally on dorsal surface, smooth or lected. It differs from C. erosum, among other
mammillose-papillose ventrally; gemmiferous ways, in lacking ventrally mammillose distal
leaves not much differentiated,contractedat tips cells in its cancellinae,its weakvariableteniolae,
and bearinggemmae on ventralsurfaceof costa, and in bearinggemmaeonly ventrallyat the leaf
which may be shortly excurrent.Seta 4-5 mm; tips;C. afzelii differsin havingsmall obscureleaf
capsule 2-2.5 mm long; operculum ca. 1 mm cells and consistently strong teniolae, among
long. Spores 17-30 pm, finely papillose. Calyp- otherfeatures.
tra rough above.
Illustrations. Bartram(1949, fig. 37: A-C);
Florschiitz (1964, fig. 53); Reese (1961, figs. 16. Calymperes rubiginosum (Mitten) Reese,
94-100) Bryologist78: 92. 1975. Fig. 73.
Distribution. (Fig. 74A). Endemicto the neo-
Syrrhopodon rubiginosusMitten,J. Linn.Soc., Bot. 12:
tropics.SouthernMexico; CentralAmerica;scat- 125. 1869; Thyridiumrubiginosum(Mitten)Jaeger,
teredin West IndiesandnorthernSouthAmerica. Ber. ThiatigkSt. GallischenNaturwiss.Ges. 1877-
Bark, dead wood, rocks, to ca. 1000 m; forests. 1878:415.1880. Type.Venezuela.Maypures,ad.arb.,
Spruces.n. (lectotype,NY; isolectotype,BM).
Selectedspecimensexamined.MEXICO.VERACRUZ: Calymperes rufumHerzog,Beih.Bot.Centralbl.61: 585.
Zona de los rios Uxpanapa y Solosichil, Delgadillo M. 1942.Type.Brazil.Amazonia,Francks.n.(JE).
3430 (LAF, MEXU). Macromitrium shankiiCrum,Bryologist55: 281. 1952.
GUATEMALA. The type of Calymperes carionis C. Type.Honduras. Morazin:Vie. of El Zamorano,Shank
Mull. s.n. (DS).
BELIZE. STANNCREEKDISTRICT: Gentle 3531 (LAF,
NY). ToLEDo:NearMoffredye Creek, Gentle 5303 (MO). Plants small, dark-green or often purplish
NICARAGUA. BLUEFIELDS: 3.6 km SE of Cerro San
(brownin the herbarium),in low, dense or strag-
Isidro, Proctor et al. 2 7020 (MO, NY).
COSTA RICA. Rio Banano, 50 m, Alfaro 109 (LAF, gly mats,stemssometimesrepentwith ascending
NY).
branches.Leaves crispatewhen dry, 1.5-2 mm
PANAMA. BOCASDELTORO:Escudo de Veraguas long, linear to oblong-lanceolate, lacking ex-
Island, McPherson 1143D (MO). CANAL ZONE:Barro panded shoulders;margins unistratose,crenate
Colorado Island, Salazar Allen & Chung C. 7856 p.p.
by protruding cells above cancellinae; can-
(PMA). CHIRIQUt:NE of FortunaCamp site, Salazar A. et cellinae small, distinct; cells of upper lamina
al. 544 (NY, PMA).
PUERTO RICO. Sierrade Luquillo,Steere4005a evenly bulging dorsallyand ventrally, rounded,
(NY). 13-20 pm diam.; costa smooth; gemmiferous
WINDWARD ISLANDS. ST. LUCIA:Basse de l'Isle, leaves not much differentiated,involute above
Sitmmonds 190 (NY). and bearinggemmae only on ventralsurface of
VENEZUELA. BOLIVAR: SW slopes of Chuiauta-
costa at apex.Seta 2-3 mm long; capsule 1.8 mm
tepui, 1000-17000 m, Steyermark75429 (FH, LAF).
TRINIDAD. [without locality] E. G. Brittonetal. 1670 long; operculumca. 1 mm long. Spores ca. 25
(NY). pm, finely papillose. Calyptra2.2-2.5 mm long,
FRENCH GUIANA. Saul, Monts La Fum6e, Boom & roughenedabove.
Mori 153 (NY). Illustrations. Crum (1952, fig. 1-5, as
BRAZIL. AMAZONAS: 30 km E of Manaus, 30 m, Macromitrium shankiiCrum);Florschiitz(1964,
Griffin et al. 021297 (FLAS, LAF). PARA:Serra do as C.
Cachimbo, 350-500 m, Reese 16505 (INPA, LAF, NY). fig. 44, rufum Herz.), Reese (1961, figs.
ROND6NIA: First rapids on Rio PacaaisNovos, ca. 200 m, 30-34, as C. rufumHerz.).
Reese 13491 (INPA, LAF, MICH, MO, NY). Distribution. (Fig. 74B). Central America
(Honduras); northern South America. Tree
Discussion. By its involute-subulate leaves trunksin humidforests at low elevations.
96 Flora Neotropica

b ~ O O

vc 0w Ooo33o

A B

FIG. 73. Calytmperesrubiginosum. A. Outlines of leaves. B. Margin and cells at midleaf. C. Section of leaf. From
Franck s.n. (holotype of Calymperes rufun, JE). Scale bars: a = 1 mm (A); b = 0.1 mm (B-C).

Selected specimens examined. HONDURAS. (the South America were erroneously labeled as to
type of Macromitriumshankii, cited above).
COLOMBIA. VAUPts:Rio Kananari,Cerro Isibukuri, locality. See my 1978 article for furtherdiscus-
Schultes & Cabrera 13304 (FH, NY). sion of this problem.
GUYANA. Upper Demerara/Berbice region, Boom The genus MitthyridiumRobins. (Tlhyridium
7149 (NY). Mitt.,hon. illeg.) has long been consideredto be
FRENCH GUIANA. ST-LAURENT-DU-MARONI:Can-
ton de Maripasoula, ca. 6 km N of Saiil, 200-400 m, Buck representedin the Americas by a single species
18569 (CAY, NY), 18591 (NY), 18830(CAY, NY). collected long ago on the westerncoast of South
PERU. LORETO:Maynas, 100 m, Timme 4841 (herb. America [Mitten 1869, as Syrrhopodon
Timme, LAF). fasciculatus Hook. & Grev.; Nowak 1980, as
BRAZIL. AMAZONAS: 62-67 km W of Humaita,Fife et
aL 4286 (INPA, LAF, NY). PARA: Serrado Cachimbo, km
Mitthyridiumobtlusifolium(Lindb.) Robins.]
780 on Cuiaba-Santar6mroad, Reese 16372 (INPA, LAF,
Mitten(1869) cited one specimen, "Chili,Lobb,"
MICH, MO, NY, US). ROND6NIA:Santa Barbara,8 km S of while Nowak cited anotherspecimen,"Chile:...
MIBRASA office, Fife et aL 4252 (INPA, LAF, NY). 1835, Cumings. n."I have seen the Cumingspec-
imen cited by Nowak, andanotherat BM labeled
Discussion. Calymperes rubiginosum is "Peru,Ulota, Cuming,"but not the Lobb speci-
widely but sparingly distributed in northern mencitedby Mitten.The Cumingspecimenmore
South America; it may occur in abundance lo- or less representsM. obtusifolium.However, be-
cally. It is known otherwise from only one col- cause this species has not been recordedin recent
lection in Central America, the type of time from the Americas,and because both Cum-
Macromnitriumshankii Crum, from central Hon- ing and a W. Lobb collected not only in South
duras. It is readily recognizable by its small size, Americabut also in the paleotropics,it is likely
commonly purple color, and by its leaves with that the reportsof this moss for South America
unistratose, crenate margins and large bulging are based on mislabeled specimens. Thus M.
cells. It has no close relatives. obtusifoliumand the genus Mitthyridiumare ex-
cluded fromthis treatment.
EXCLUDED TAXA
ACKNOWLEDGMENTS
In my revision of the limbate taxa of neotropi-
cal Syrrhopodon (1978), I included S. rigescens I thank the curatorsof the herbariacited for
Schwaegrichen (now treated as a synonym of S. theirunfailingcourtesyin loaning specimens for
involuttus Schwaegrichen, cf. Reese et al. 1986), thisstudy,WilliamR. BuckandS. Rob Gradstein
for reasons explained at the tine. I now conclude, for critical review of the manuscript, Rupert
however, based on study of paleotropical mate- Bareby for correcting the Latin diagnosis of
rial of this species (including field study), that the Calymperessubg. Porphyrophylum,and Emilio
specimens upon which I attributed S. rigescens to Garciafor providingthe Spanish version of the
Calymperaceae 97

TROPICALAMERICA FeoraNeotroprcabase map no 1

F 7 ib n AicanCalymperes.A. C ..... . ..........

C.

FIG. 74. Distributions of AMnericanCalymperes. A. C. nicaraguense. B. C. rubiginosumi.

98 FloraNeotropica

abstract. Steven P. Churchill offered helpful Fleischer,M. 1904.Die MusciderFloravon Buitenzorg.

comments on the manuscript.Documentationof Vol. 1. Leiden.
Florschiitz,P. A. 1964.TheMossesof Suriname.Part1.
ranges of taxa by citation of specimens was Leiden.
greatlyenhancedby supportfromthe WilliamC. Fritsch,B. 1991.Indexto bryophytechromosomecounts.
Steere Fund and The New York Botanical Gar- Bryophytorum Bibliotheca40. Berlin-Stuttgart.
den, arrangedby William R. Buck, which al- Jaeger,A. & F. Sauerbeck.1871-1872[1873].Generaet
lowed me to visit New York to work in the Species Muscorum.III. Verh. [Berichte iiber die
Thatigk.]St. GallischenNaturwiss.Ges. 1871-1872:
bryophyte herbarium.Experience gained as a 309-490.
participanton two collecting tripsto Amazonian Kindberg, N. C. 1897. Genera of European and
South America, supported by Projeto Flora Northamerican Bryineae(Mosses) SynopticallyDis-
throughThe New York Botanical Garden,was posed.G6teborg.
Magill,R. E. 1981.Floraof SouthernAfrica.Bryophyta.
very valuable in that it allowed me to study Part1 mosses.Fascicle1 Sphagnaceae-Grimmiaceae.
Calymperaceaein the field. The University of Pretoria.
Southwestern Louisiana has provided support Mitten,W. 1869.MusciAustro-Americani. J. Linn.Soc.,
throughoutmy study of Calymperaceae.Most of Bot. 12: 1-659.
the illustrationsof Calymperaceaein thistreatment Muller,C. 1849.SynopsisMuscorumFrondosorum. Vol.
1. Berlin.
werepreviouslypublishedin articlesin TheBryol- - 1901.GeneraMuscorumFrondosorum. Leipzig.
ogist and arereprintedherewithpermission. Nowak, H. 1980. Revision der Laubmoosgattung
Mitthyridium (Mitten) Robinson fiir Ozeanien
(Calymperaceae). Bryophyt.Biblioth.20: 1-236.
Orban, S. & W. D. Reese. 1990. Syrrhopodon prolifer
LITERATURECITED (Musci:Calymperaceae): A worldview. Bryologist93:
438-444.
Bartram, E. B. 1928. Costa Ricanmosses collectedby Reese, W. D. 1961.ThegenusCalymperesin the Amer-
PaulC. Standleyin 1924-1926. Contr.U.S. Nat.Herb. icas. Bryologist64: 89-140.
- . 1977.ThegenusSyrrhopodon in the AmericasI.
26(3): 51-114.
. 1949. Mosses of Guatemala.Fieldiana,Bot. 25: Theelimbatespecies.Bryologist80: 2-31.
1-442. -- 1978.ThegenusSyrrhopodon in the AmericasII.
Bescherelle,E. 1896.Essaisurle genreCalymperes. Ann. Thelimbatespecies.Bryologist81: 189-225.
Sci. Nat. Bot. 8, 1:247-308 -- . 1981. Refinementson AmericanSyrrhopodon
Brotherus,V. F. 1901.Musci. In A. Engler& K. Prantl (Musci:Calymperaceae). Bryologist82: 244-248.
(eds.), Die natiirlichenPflanzenfamilien.
Vol. 1. Leip- ---- 1982. Syrrhopodon isthmi (Musci;
zig. Calymperaceae), a new species from Panama.Bryol-
- . 1924. Musci.In A. Engler& K. Prantl(eds.),Die ogist85: 85-87.
natiirlichenPflanzenfamilien,2nd. ed. Vol. 10. Leip- --- 1983a.AmericanCalymperesand Syrrhopodon:
zig. Identification
keyandsummaryof recentnomenclatural
Crum, H. 1952. Mosses from Honduras,II. Bryologist changes.Bryologist86: 23-30.
55: 279- 285. --- . 1983b.Syrrhopodon brasiliensis,a previouslyun-
-- 1956. Syrrhopodon alatomarginatus. A new spe- describedspeciesfromsouthernBrazil.Bryologist86:
cies from Trinidad.Trans.Brit.Bryol.Soc. 3: 72-73. 354-356.
& L. E. Anderson.1981.Mossesof EasternNorth 1. 984. Calymperesand Syrrhopodon(Musci:
America,Vol. 1. New York. Calymperaceae) in the Serrado Cachimbo,Brazil,and
& W. C. Steere. 1957.Themossesof PortoRico Rhacopilopsistrinitensis(C.M.) Britt.& Dix. new to
and the Virgin Islands.The New York Academyof Brazil.Lindbergia10: 123-126.
Sciences,ScientificSurveyof PortoRicoandtheVirgin . 1987a. Calymperes(Musci: Calymperaceae):
Islands7(4): 395-599. Worldranges,implicationsfor patternsof historical
Dixon, H. N. 1930.Classificationof mosses,Pages397- dispersionandspeciation,andcommentsonphylogeny.
412. In F. Verdoorn(ed.) Manualof Bryology.The Brittonia39: 225-237.
Hague. . 1987b.Worldranges,implicationsfor patternsof
Edwards, S. R. 1980. Sporedischargein Calymperes. J. historicaldispersaland speciation,and commentson
Bryol.11: 95-97. phylogeny of Syrrhopodon(Calymperaceae).Mem.
Eddy,A. 1990.A Handbookof MalesianMosses,Volume New YorkBot. Gard.45:426-445.
2. BritishMuseum(NaturalHistory).London. - 1990. Calymperesothmerireinstated.Bryologist
Ellis, L. T. 1988.Typificationof Calymperes laevifolium 93: 37-38.
Mitt.andC.pallidumMitt.J. Bryol.15: 233. . 1992. CalymperesandSyrrhopodon.In B. Allen
--- 1989. A taxonomicrevision of Calymperesin (ed.),A MossFloraof CentralAmerica.Vol. 1. Mono-
southernIndiaandneighbouringislands.J. Bryol.15: graphsin SystematicBotanyfromTheMissouriBotan-
697-732. icalGarden.(Accepted.)
Calymperaceae99

-- & W. R. Buck 1991. A sample of Calymperes and Calymperaceae:A reviewwithanewspecies.Bull.Nat.

Syrrhopodon from French Guiana. Bryologist 94: 298- Sci. Mus.(Tokyo),Ser.B (Bot.)9(1): 23-32.
300. & R. H. Zander. 1988. Comparison of
& D. Griffin, III. 1976[1977]. Syrrhopodon an- CalymperaceaewithPottiaceae.Bryologist91: 18-20.
notinus (Musci: Calymperaceae), a new species from Robinson, H. 1972. Cheilotheca vaginata and
Amazonas, Brazil. Bryologist 79: 518-520. Syrrhopodonsteyermarkii.Two new moss species from
- , T. Koponen & D. H. Norris. 1986. Bryophyteflora Venezuela.Phytologia23: 390-392.
of the Huon Peninsula, Papua New Guinea. XIX. Roth, G. 1912.Ubersichtiiberdie GattungCalymperes.
Calymperes, Syrrhopodon and Mitthyridium Hedwigia51:122-134.
(Calymperaceae,Musci). Acta Bot. Fenn.133:151-202. Tixier, P. 1967.Le genreCalymperopsis(C.M.)Fleisch.
& S. Orban. 1986. Syrrhopodon incompletus (Calymperaceae). RevueBryol.Lich6nol.35:265-302.
Schwaegr. var. incompletus (Musci: Calymperaceae) Williams,R. S. 1920.Calymperaceae of NorthAmerica.
disjunct between the Americas and Africa. Bryologist Bull.TorreyBot.Club47: 367-395. [Reprinted
without
89: 213-214. changeof pagination as"Contr.NewYorkBot.Gard.-
& B. C. Tan. 1983. The "petiolate" No. 223."]

INDEX OF SCIENTIFICNAMES
Anictangium parasiticum (Brid.) Brid. 38 C. leboucherianumPar. & Broth. 66
Brachypodium parasiticum (Brid.) Brid. 38 C. levyanum Besch. 73
Bryum lycopodioides Brid. 46 C. lindmanii Broth. 77
B. parasiticum Brid. 38 C. lonchophylloides C. Mill. 70
Calymperes aberransPar. 83 C. lonchophyllum Schwaegr. 70
C. afzelii Sw. 77 C. maguirei Bartr. 75
C. androgynum Mont. 61 C. mammilliferum Crum & Steere 81
C. armatum Broth. 50 C. melinoni C. Miill. in Besch. 80
C. asperipes Besch. 70 C. meyeri Reese 50
C. badium (Schimp. in Besch.) Broth. 93 C. mitrafugax Florsch. 91
C. bahiense C. Mill. 83 C. nashii Williams 84
C. bartramii Reese 91 C. nicaraguense Ren & Card. 93
C. bodenii C. Mill. 81 C. othmeri Herzog 82
C. bolivianum Williams 93 C. palisotii Schwaegr. 83
C. breutelii Besch. 83 C. palisotii subsp. richardii (C. Miill.) S. Edwards 83
C. brevicaule Par. & Broth. 83 C. pallidum Mitt. 75
C. brittoniae Besch. 77 C. panamae Besch. 83
C. carionis C. Miill. 93 C. perinvolutumC. Miill. 83
C. chlorosum Hampe 77 C. piovanoi Biz. 75
C. crispum (Aust.) Aust. 27 C. platyloma Mitt. 87
C. cruegeri C. Mill. 93 C. portricense Ren. & Card. 77
C. cubense Williams 75 C. remirense Par. & Broth. 77
C. parasiticum (Brid.) Hook. & Grev. 38 C. reyi Par. & Broth.77
C. disciforme C. Miill. 42, 44 C. richardiiC. Miill. 83
C. donnellii Aust. 77 C. rubiginosum (Mitt.) Reese 95
C. emersum C. Miill. 80 C. rufescens Besch. 77
C. erosum C. Miill. 80 C. rufum Herzog 95
C. filigera Mitt. ex Tixier 38 C. rupicola P. Richards 82
C. filigerum Aust. 38 C. semilimbatulum C. Miill. 38
C. gardneri Hook. 47 C. smithii Bartr. 87
C. glaziovii Hampe 50 C. sprucei Besch. 80
C. guadeloupense Broth. in Urb. 93 C. swartzii Amott 38
C. guadeloupense Besch. 83 C. tenerum C. Mill. 84
C. guianense Par. & Broth. 77 C. tenuifolium Sull. 18
C. guildingii Hook & Grev. 91 C. uleanum Broth. 75
C. heribaudii Par. & Broth. 93 C. venezuelanum (Mitt.) Broth. ex Pitt. 75
C. hexagonum Besch. 83 C. wullschlaegelii Lor. 80
C. hobsonii Grev. 48 Calymperopsis disciformis (C. Miill.) Tixier 42
C. hookeri Besch. 83 C. disciformis (Dus.) Fleisch. 42
C. huallagense Broth. 77 C. martinicensis (Broth.) Broth. 38
C. lanceolatum Hampe 54 C. parasitica(Brid.) Broth. 38
100 FloraNeotropica

C. wainioi(Broth.)Fleisch. 38 S. elatusMont. 55
ChameleioncryptocarposEllis& Eddyin Eddy 66 S. elongatusSull. 36
ChameleionxanthophyllusEllis& Eddyin Eddy 66 S. elongatusvar.anomalusTher. 36
Dicranumberteroanum Brid. 51 S. elongatusvar.elongatus 36
DicranumcircinatumBrid. 57 S. elongatusvar.glaziovii(Hampe)Reese 36
Dicranumlycopodioides(Brid.)Sw. 46 S. epapillosusC. Mill. 16
Encalyptaparasitica(Brid.)Sw. 38 S. erubescensBartr. 33
Glyphomitrium parasiticum(Brid.)Brid. 38 S. filigerus(Aust.)Williams 38
Heliconemacryptocarpos(Dozy & Molk.)Ellis& Eddy S. fimbriatusMitt. 38
in Ellis 66 S. flavescensC. Miill. 15
H. xanthophyllus(Mitt.)Ellis & Eddyin Ellis 66 S. flexiareolatusC. Mill. 25
Macromitrium shankiiCrum 95 S. flexifoliusMitt. 39
Mitthyridium obtusifolium(Lindb.)Robins. 96 S. floridanusSull.in Gray 48
Orthothecaberteroana (Brid.)Brid. 51 S. gardneri(Hook) Schwaegr. 47
0. circinata(Brid.)Brid. 57 S. gaudichaudii Mont. 25
0. lycopodioides(Brid.)Brid. 46 S. gaudichaudii var.recurvulus(Mitt.)Thir. 31
Syrrhopodon acanthoneuros C. Miill. 16 S. glazioviiHampe 36
S. aculeociliatusBartr. 31 S. goyazensisBroth. 16
S. africanus(Mitt.)Par. 44 S. gracilescensBroth. 25
S. alatomarginatus Crum 66 S. graminicolaWilliams 42
S. allioniiBroth. 16 S. griffiniiRobins. 29
S. androgynous(Mont.)Besch. 61 S. helicophyllusMitt. 44
S. annotinusReese & Griffin 19 S. hioramiiBiz. & Ther. 50
S. anomalusBroth. 27 S. hobsonii(Grev.)Hook.& Grev. 48
S. arenariusC. Miill. 16 S. hobsoniivar.luridus(Par.& Broth.)P. Richards 52
S. argenteusBroth. 16 S. hobsoniivar.spinulosusHerzog 47
S. argentinicusLor.ex C. Mill. 25 S. hornschuchii Mart. 59
S. assimilisBroth. 11 S. husn6tiiBesch. 19
S. badiusSchimp.in Besch. 93 S. incompletusSchwaegr. 47
S. beroullii C. Miill. 50 S. incompletusvar.berteroanus (Brid.)Reese 51
S. berteroanus var.planifoliusTh6r. 51 S. incompletusvar.elatus(Mont.)Florsch. 55
S. berteroanus var.strictiorHampe 51 S. incompletusvar.incompletus 47
S. berteroanus (Brid.)C. Miill. 51 S. incompletusvar.lanceolatus(Hampe)Reese 54
S. berteroiKindb. 51 S. incompletusvar.luridus(Par.& Broth.)Florsch. 52
S. borinquensisCrum& Steere 33 S. incompletusvar.perangustifolius Reese 54
S. brachystelioides C. Miill. 50 S. inflexusMitt. 25
S. brasiliensisReese 27 S. involutusSchwaegr. 96
S. brevisetusFlorsch. 29 S. iridansMitt. 31
S. brotheriC. Mill. 50 S. isthmiReese 33
S. calochlorusC. Miill. 11 S. juruensisBroth. 50
S. calymperidianus Besch. 15 S. kroneanusC. Miill. 54
S. capillaceusHampe 18 S. laevidorsusBesch. 51
S. carassensisBroth. 16 S. laeviusculusBroth. 11
S. ciliolatusGeh.& Hampein Hampe 16, 47 S. leboucherianus (Par.& Broth.)Reese 66
S. ciliolatusHerzog 16, 47 S. leprieuriiMont. 31
S. cincinnatusHampe 19 S. leptophyllusBroth. 19
S. circinatus(Brid.)Mitt. 57 S. ligulatusMont. 27
S. compactulusC. Miill. 11 S. longifoliusLindb.ex Aongstr. 11
S. crispulusSehnem 25 S. longifoliusvar.alticaulisBroth. 15
S. crispusAust. 27 S. longisetaceusC. Miill. 61
S. cristatusBartr. 31 S. luetzelbergiiHerzog 16
S. cryptocarpos Dozy & Molk. 66 S. luridusPar.& Broth. 52
S. cryptodusMitt. 15 S. lycopodioides(Brid.)C. Miill. 46
S. curvatusC. Miill. 51 S. macrophyllus Broth. 16
S. cylindrothecius C. Miill. 16 S. macrophyllus Broth.in Herzog 15
S. cymbifoliusC. Miill. 44 S. macroprolifer C. Miill. 16
S. decoloransC. Miill. 50 S. malouinensisC. Miill. 51
S. densifoliusHerzog 23 S. martinicensisBroth.in Urban 38
S. disciformisDus. 44 S. martiniiHerzog 31
S. diversifoliusBroth. 57 S. minutusBroth. 11
S. dussiiBroth.in Urban 15 S. miquelianusC. Miill. 59
S. elatiorHampe 16 S. mohriiC. Miill. 50
Calymperaceae 101

S. ochroleucusHerzog 25 S. scaberrimus C. Mill. 31

S. pallidusMitt. 31 S. schiffneriBroth. 42
S. papillosusC. Mill. 15 S. schwaneckeanus C. Mull. 15
S. paraguensisBesch. 27 S. schwaneckeanus fo. majorBesch. 15
S. parasiticus(Brid.)Besch. 38 S. semicompletusSchwaegr. 48
S. parasiticusvar. disciformis(C. Mill.) S. serpentinusC. Mill. 31
Florsch. 42 S. simmondsiiSteere 29
S. parasiticusvar.flexifolius(Mitt.)Reese 39 S. sordidusC. Miill. 57
S. parvulusSchimp.ex C. Mill. 11 S. spininervisLindb.in Aongstr. 25
S. perhorridus C. Mill. 51 S. spruceanusMitt. 66
S. persordidusC. Mill. 11 S. steyermarkii Robins. 63
S. pittieriRen.& Card. 57 S. subdecolorans Broth. 50
S. proliferSchwaegr. 11 S. subintegerLindb.ex Aongstr. 18
S. prolifervar.acanthoneuros (C. Mill.) C. Mull. 16 S. submacrophyllus Broth.in Herzog 15
S. prolifer var. cincinnatus (Hampe) Reese 19 S. subrigidusBroth. 57
S. prolifervar.papillosus(C. Mill.) Reese 15 S. subscaberBroth. 16
S. prolifervar.prolifer 11 S. subviridisBesch. 15
S. prolifervar.scaber(Mitt.)Reese 13 S. surinamensis
S. prolifervar.tenuifolius(Sull.)Reese 18 Dozy & Molk 59
S. sylvaticusMitt. 31
S. pusillusMitt. 31 S. tenuifolius(Sull.)Mitt. 18
S. quintasiiBroth. 47 S. tenuipapillosusTher. 16
S. ramicolaBroth. 44
S. terebellatulusC. Mill. 25
S. recurvulusMitt. 31
S. texanusSull. 23
S. rhizogonioidesC. Miill. 46
S. theriotiiBartr. 64
S. rigescensSchwaegr. 96
S. rigidusHook & Grev. 61 S. tortilisHampe 23
S. rivularisHerzog 82 S. ulei C. Mull. 23
S. rosulatusC. Miill. 23 S. uleivar.percrispaC. Mill. 23
S. rubicundusC. Miill. 16 S. undulatusC. Miill. 25
S. rubiginosusMitt. 95 S. vaginansBroth. 51
S. rufusHornsch. 59 S. venezuelanusMitt. 75
S. rupestrisMitt. 63 S. wainioiBroth. 38
S. rupicolaBroth. 16 S. xanthophyllus Mitt. 66
S. sartoriiC. Mill. 25 ThyridiumafricanumMitt. 44
S. scaberMitt. 15 T. rubiginosum (Mitt.)Jaeg. 95
S. scabervar.breviligulatusC. Miill. 15 Weisiaparasitica(Brid.)Mohr 38

ADDENDUM TO KEY B, PAGE 10

The elimbate SyrrhopodonautotomaiusReese was recently described as a new species (Bryologist 96:
145-148. 1993). Known only from Honduras,in the Departmentsof Olancho and Toledo, S. autotomaius
is remarkablysimilar in most aspects to S. circinatus but differs notably in having caducous leaf tips. The
regularlydehiscent leaf tips are green, filiform, and mostly 2-3.5 mm long by 80 ,xm wide; typical calym-
peraceous foliar gemmae are not formed. This moss also produces sporophytes.Syrrhopodonautotomaius
grows above 1000 m, mostly on palm stems; it is the only member of the Calymperaceaeknown to have
caducous leaf tips.