Organization for Flora Neotropica

Plagiotheciaceae
Author(s): William R. Buck and Robert R. Ireland
Reviewed work(s):
Source: Flora Neotropica, Vol. 50, Plagiotheciaceae (Apr. 20, 1989), pp. 1-20
Published by: New York Botanical Garden Press on behalf of Organization for Flora Neotropica
Stable URL: http://www.jstor.org/stable/4393810 .
Accessed: 17/10/2012 22:21

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .
http://www.jstor.org/page/info/about/policies/terms.jsp

.
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of
content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms
of scholarship. For more information about JSTOR, please contact [email protected].

New York Botanical Garden Press and Organization for Flora Neotropica are collaborating with JSTOR to
digitize, preserve and extend access to Flora Neotropica.

http://www.jstor.org
FLORA NEOTROPIC

MONOGRAPH 50

PLAGIOTHECIACEAE

by
William R. Buck and RobertR. Ireland

TROPiC Of C^NClR
|c~^ r%\',

FLORAt
NEOTROPICA

ROPc of cAP RICORtN t4

Publishedfor
Organizationfor Flora Neotropica
by
The New York BotanicalGarden
New York

Issued 20 April 1989

 Copyright ? 1989
The New York Botanical Garden

Published by
The New York Botanical Garden
Bronx, New York 10458
International Standard Serial Number 0071-5794

Library of Congress Cataloging in Publication Data

Flora neotropica. - Monograph no. 1 - New York: Published
for Organization for Flora Neotropica by the New York
Botanical Garden, 1968-
v.: ill.; 26 cm.
Irregular.
Each issue has distinctive title.
Separatelycatalogedand classifiedin LC before monographno. 40.
ISSN 0071-5794 = Flora neotropica.
1. Botany-Latin America-Classification-Collected works. 2. Botany-
Tropics-Classification-Collected works. 3. Botany-Classification-Col-
lected works. I. Organizationfor Flora Neotropica. II. New York Botanical
Garden.

QK205.F58 581.98'012-dcl9 85-647083

AACR 2 MARC-S
Libraryof Congress [8508]
ISBN 0-89327-342-2
 PLAGIOTHECIACEAE
WILLIAMR. BUCK' AND ROBERTR. IRELAND2

TABLE OF CONTENTS
Abstract ......... .................................................. ...................
Introduction .............................................. ................................ 2
SystematicTreatment........................................................................... 2
GenericDescription . ................................................................ 3
Key to the Species of Plagiothecium.......................................................... 4
Species Descriptions ................................................................ ........ 4
Doubtful Names and ExcludedTaxa.............................................................. 19
Acknowledgments.............................................................................. 19
LiteratureCited .................................................................... ........... 20
Index of ScientificNames . .................................................................... 20

ABSTRACT
Buck, William R. (New York Botanical Garden, Bronx, NY 10458-5126, U.S.A.) and
RobertR. Ireland(BotanyDivision, National Museumof NaturalSciences,P.O. Box 3443,
Station D, Ottawa,OntarioKIP 6P4, Canada).Plagiotheciaceae.Flora Neotrop. Monogr.
50: 1-22. 1989.-Plagiothecium, the only genus of the Plagiotheciaceae,is revised for
all of CentralAmerica,the West Indies, and South America,and six species are recognized.
Plagiotheciumnovo-granatense,with only scant decurrencies,occurs from Venezuela to
Argentinaand in southeasternBrazil.The closelyrelatedP. drepanophyllum has decurrencies
more consistentlypresentand occursfrom Mexico to Colombiaand in the GreaterAntilles.
Plagiotheciumlucidum,recognizedby its slenderleaf apices and narrowleaf cells, occurs
fromColombiato Tierradel Fuegowith disjunctsin southeasternBraziland the Dominican
Republic.Plagiotheciumconostegium,distinguishedby its broad leaf cells and prominent
decurrencies,occursat high elevationsin Mexico, Guatemalaand the DominicanRepublic,
and in the northernAndes and Tierradel Fuego. Plagiotheciumovalifolium,with broadly
acute leaves, is restrictedin South Americato southernmostArgentinaand Chile. Plagio-
theciumfalklandicumis recognizedby julaceousplantsand in South Americais only in the
FalklandIslands. All species are describedand illustrated.Some taxa previouslyincluded
in Plagiotheciumaretransferredto othergenera:Isopterygiumfontigenum (C. Miiller)comb.
nov. and Lepidopilidiumregnellii(Angstr6m)comb. nov.

RESUMEN
Buck, William R. (New York Botanical Garden, Bronx, NY 10458-5126, U.S.A.) and
RobertR. Ireland(BotanyDivision, National Museumof NaturalSciences,P.O. Box 3443,
Station D, Ottawa,OntarioKIP 6P4, Canada).Plagiotheciaceae.Flora Neotrop. Monogr.
50: 1-22. 1989.-Plagiothecium, el finico g6nerode las Plagiotheciaceae,ha sido revisado
paraAm6ricaCentral,las Indias Occidentalesy Am6ricadel Sur, y se han reconocidoseis
especies. Plagiotheciumnovo-granatense,con solamentedecurrenciaslimitadas,ocurrede
Venezuela a Argentina,y en el sudeste de Brasil. Plagiotheciumdrepanophyllum,muy
relacionadocon P. novo-granatense,tiene decurrenciasm/s frecuentesy ocurrede M6xico
a Colombiay en las Antillas Mayores.Plagiotheciumlucidum,reconocidopor presentaren
las hojas los apices delgadosy c6lulasestrechas,ocurrede Colombiaa Tierradel Fuegocon

New York BotanicalGarden,Bronx,New York 10458-5126, U.S.A.

2
Botany Division, National Museum of Natural Sciences, P.O. Box 3443, Station D, Ottawa,OntarioKIP
6P4, Canada.
1
2 Flora Neotropica

disyuncionesen el sudeste de Brasil y en la RepublicaDominicana.Plagiotheciumconos-

tegium,distinguidopor las celulasampliasy decurrenciasprominentes,ocurreen los Andes
n6rticos y Tierradel Fuego y en elevaciones altas de Mexico, Guatemalay la Repfblica
Dominicana. Plagiotheciumovalifolium,con hojas acutas, esta restringidoen la America
del Sur para la parte meridional de Argentinay Chile. Plagiotheciumfalklandicum esta
reconocidopor plantasjulaceasy en Americadel Sur esta solamenteen las Islas Malvinas.
Todas las especies estan descritasy ilustradas.Algunastaxa previamenteincluidasen Pla-
giotheciumhan sido transferidasa otrosgeneros:Isopterygiumfontigenum (C. Muller)comb.
nov. y Lepidopilidiumregnellii(Angstrom)comb. nov.

INTRODUCTION have been used to distinguish species, we have

found them unreliableto the extent previously
The Plagiotheciaceaehave long been a depos- utilized. Some generalcorrelationscan be made
itoryfor a wide assortmentofcomplanate-foliate but leaf charactersare as a rule more reliable
hypnaleanmosses. The family has includedsuch (and always present).
diverse elements as Plagiothecium itself, Iso- Although we are confident of our species de-
pterygium, Isopterygiopsis,Herzogiella, Taxi- limitations, there are problems inherent in a
phyllum,Stereophyllumand Catagonium.How- treatmentof the taxa from an isolated geograph-
ever, we (Buck & Ireland, 1985) have purified ical area. We readily recognize that our south-
the family, dispersingits disparateelements to ernmost taxa are probably circumsubantarctic,
severaldifferentfamilies, and have left it mono- and may have other names, and differentmor-
typic. Here, we treat the many names placed in phologicalvariation,in otherpartsof theirranges.
Plagiotheciumfor Latin America. No such at- Similarly,some of the northernAndeantaxamay
tempt has previously been made. From the be disjunct populations or vicariant taxa of
northernend of the neotropics, Bartram(1949) NorthernHemispherespecies. Nevertheless,for
recognizedfourspeciesof Plagiotheciumin Gua- this study we have been unable to adequately
temala, only two of which are currentlyin the surveythe extralimitalspecies of Plagiothecium.
genus. From the opposite end of the rangeNew- We can take heart,though,in the fact that some
ton (1983b) recognized Plagiotheciumfalklan- of our taxa have the oldest names in the genus,
dicum from the Falkland Islands (Islas Malvi- and we are thus on relatively stable groundno-
nas), and also discussed P. ovalifolium from menclaturally.
southernmostSouth America. However, in the
Andes, where Plagiotheciumreachesits greatest
diversity in South America, no comprehensive SYSTEMATIC TREATMENT
coveragehas been attemptedbeforenow. Rather, PLAGIOTHECIACEAE(Brotherus)Fleischer,
species have been described without critical Nova Guinea8(2, Bot.):748. 1912;Hypnaceae
comparisonto previously describedspecies. As subfam.PlagiothecioideaeBrotherusin Engler
a consequence,many names are placed in syn-
& Prantl,Nat. Pflanzenfam.1(3): 1078. 1908,
onymy here.
We have found severalcharactersof particular "Plagiothecieae."Type. PlagiotheciumBruch,
value when circumscribingtaxa. In a generalway Schimper& Gimbel.
leaf shape, especially at the apex, can be useful Plants dull or more often glossy, somewhat
in a quick sorting of taxa. Three rough classes small to robust, in loose to dense mats. Stems
can be discerned:long-acuminate,short-acumi- prostrate, mostly complanate but sometimes
nate and acute.Cell width correlatesmore or less subjulaceousor erect and julaceous, simple or
directlywith leaf apex shape,with the narrowest sparselyand irregularlybranched;outer layer of
cells in those leaves with the narrowestapices. cortical cells large and thin-walled; rhizoids
Finally, the size of the leaf decurrenciesand the sometimes present just below leaf insertion,
shape of their cells are criticalin sortingout oth- smooth; pseudoparaphylliafoliose, rare.Leaves
erwise similar taxa. Although sporophyticchar- ovate, ovate-lanceolate,oblong-lanceolateor ob-
acters, especially capsule size and orientation, long-ovate, acute, acuminate, rarely piliferous,
SystematicTreatment 3

often concave, erect or spreading, scarcely al- NonVasc.13]:298. 1966.Type.Savicziaobtusissima

tered when dry, often asymmetric, decurrent; A. Abramova& I. Abramov.
(Brotherus)
costae short and double, one branchsometimes
reachingmidleaf, or rarelyecostate, rhizoids at Plants often glossy, in dark green to yellow-
base on dorsal surfacesor adjacentcells, some- green, sometimes whitish green, thin to dense,
times at leaf tips, smooth; uppercells linear,lin- flat mats. Stems 1-10 cm long, usually compla-
ear-flexuose or sometimes linear-rhomboidal, nate-foliate, sometimes subjulaceous to jula-
mostly thin-walled and heavily chlorophyllose, ceous, often stout, simple or sparselyand irreg-
smooth or rarely with minute cuticular rough- ularlybranched;in cross-sectionwith a cortexof
enings; alar cells differentiated,equally distrib- a singlerow of large,thin-walledcells, subtended
uted on either side of the costa, extending into by smaller, thicker-walledcells, internallywith
decurrencies, the walls ? evenly thickened. large,thin-walledcells, centralstrandpresentor
Brood-bodiessometimes present, cylindricalto indistinct;pseudoparaphylliafoliose, rare;axil-
fusiform, uniseriate, clustered in leaf axils or lary hairs with a single, short, brown basal cell
sometimes on abaxial surfaceof leaf. Autoicous and three to four elongate, hyaline apical cells.
or less commonly dioicous. Setae straight,long, Stem and branch leaves similar, ovate, ovate-
smooth, twisted; capsules ceruous or erect, lanceolate, oblong-lanceolate,or oblong-ovate,
straightor arcuate;exothecialcells thin- or thick- oftenconcave,acute,acuminateorpiliferous,soft,
walled; annulus present, often persistent;oper- imbricate to distant, erect or spreading,some-
culum conic to rostrate;peristome double, ex- times secund,little alteredwhen dry, symmetric
ostome teeth on outer surface cross-striolate or oftenasymmetric,smoothor undulate,strongly
below and papillose above, or rarely papillose or scarcelydecurrent;marginsplane or recurved,
throughout,projectingon inner surface;endo- entire or serrulateto serrate at extreme apex;
stome with a medium to high basal membrane, costae shortand double, one branchoften reach-
the segmentsbroador narrow,keeled,not or very ing ?-1/2 the leaf length,rarelyone branchpoorly
narrowlyperforate,shorterthan or almostas long developed and costa appearingsingle, or costa
as the teeth, the cilia well developed to absent. sometimeslacking;uppercells linear,linear-flex-
Calyptraecucullate,smooth, naked. uose or sometimes, when exceptionally broad,
Distribution.As circumscribedhere, the Pla- linear-rhomboidal,often becoming shorternear
giotheciaceaeare a monotypic family, with the leaf apex and towardinsertion,smooth or rarely
single genus Plagiothecium,primarilyoccurring with minute,granular,cuticularroughenings;alar
in temperate latitudes and at higher elevations cells often differentiated;decurrenciestriangular
in the tropics. to auriculate,in one to severalverticalrows,cells
elongate to rounded-quadrate. Brood-bodies
often present (but not in South Americantaxa)
PLAGIOTHECIUM Schimper in Bruch, from a stalked,branchedbase, cylindricalor fu-
Schimper& Giimbel, Bryol.Eur.5:179. 1851; siform, 2-7-celled, uniseriate, clustered in leaf
Stereodon(Bridel)Mitten sect. Plagiothecium axils on stems and branches,sometimes on dor-
(Schimper)Mitten, J. Linn. Soc., Bot. 4: 88. sal leaf surfaces,hyalineto lightgreen.Autoicous
1859. Type. Plagiothecium denticulatum or dioicous. Perichaetiaand perigonianumerous
(Hedwig)Bruch,Schimper& Giimbel. at bases of stems; perigonial bracts short, lan-
PhilosciaBerkeley,Handb.Brit.Moss. 146. 1863.Type. ceolate to ovate, acuminate to filiform-acumi-
Philoscia latebricola(Bruch, Schimper& Giimbel) nate; perichaetial leaves moderately enlarged,
Berkeley. sheathingwith somewhatspreadingapices.Setae
Acrocladiopsis Cardot,Rev. Bryol.41: 9.
(Brotherus) smooth, long, twisted, straight,curved or rarely
1914; CatagoniumC. Miillerex Brotherussect. Ac-
Brotherus
rocladiopsis inEngler&Prantl,Nat.Pflan- circinate,yellow, orangeto red-brown;capsules
zenfam.1(3): 1088. 1908. Type.Acrocladiopsisdray- erect to cernuous, rarelypendulous, straightor
tonii (Sullivant)Cardot. arcuate,yellow, orangeto red-brown,oblong to
PlagiotheciellaFleischerex Brotherusin Engler& Prantl, ovoid, often contractedunder mouth when dry,
Nat. Pflanzenfam.ed. 2, 11: 465. 1925, nom. illeg.
Type.Plagiotheciellalatebricola(Bruch,Schimper& smooth to furrowed;exothecial cells thick- or
Giimbel)Fleischer. thin-walled;annulus differentiatedin 1-3 rows,
SavicziaA. Abramova& I. Abramov,Novit.Syst.P1. deciduous,sometimestardilyso; operculumconic
4 FloraNeotropica
to rostrate,shorterthan the urn;exostome teeth renciessmallbut alwayspresenton
bordered, shouldered, on outer surface cross- every leaf; Mexico to Colombia,
GreaterAntilles. ...............
striolate below, sometimes with overlying pa- ............. 2. P. drepanophyllum.
pillae, coarselypapillose above, rarelypapillose 5. Plants mostly obscurely compla-
throughout, projecting from inner surface;en- nate with closely spaced, ? arched
dostome papillose, with a mostly high basal leaves; decurrencies very small,
often absentor difficultto find;An-
membrane, segments keeled, not or very nar- dean SouthAmerica,SE Brazil. ..
rowly perforate,cilia 1-3, nodulose, as long as ............ 1. P. novo-granatense.
or nearly as long as the segments, rarely rudi-
mentary or lacking. Spores spherical to ovoid, 1. Plagiothecium novo-granatense (Hampe)
smooth or papillose.Calyptraecucullate,smooth,
Mitten, J. Linn. Soc., Bot. 12: 520. 1869.
naked, fugaceous. Fig. 1.
Distribution.Plagiotheciumis a genusof about
80-90 species predominantly occurringin the Hypnum novo-granatenseHampe, Linnaea 31: 531.
1862. Type. Colombia. Cipac6n,Aug 1861, Lindig
temperatezones. Its incursion into the tropical s.n. (lectotype,BM; isolectotypes,BM-3).
latitudes is primarilyat higher elevations. The Hypnum brasilienseHampe, Vidensk.Meddel.Dansk
genusis best recognizedby its mostlycomplanate Naturhist.Foren.Kj6benhavnser.3,9-10:267.1878;
stems with decurrentleaves. The costa is short Hypnum subaequans Hampe, Vidensk. Meddel.
and double and the mostly linear laminal cells Dansk Naturhist.Foren.Kj6benhavnser. 4, 1: 143.
are heavily chlorophyllose. 1879, nom. illeg. incl. sp. prior.;Plagiotheciumbra-
siliense (Hampe)Jaeger,Ber. Titigk. St. Gallischen
Naturwiss. Ges. 1877-1878: 512. 1880; Plagiothe-
cium subaequans(Hampe) Paris, Ind. Bryol. 967.
Key to the Species of 1898, nom. illeg. Type. Brazil.Rio de Janeiro,Gla-
ziou 7363 (holotype,BM, n.v.; isotype, NY).
Plagiothecium HypnumschraderiC. Miiller,Linnaea42: 496. 1879;
1. Plantsjulaceous;leavesoftenobovate,concave, Plagiotheciumschraderi(C. Miiller)Kindberg,Enum.
acute;marginsplane; in South America confined Bryin. Exot. 102. 1891. Type. Venezuela. Fendler
to the FalklandIslands..... 6. P.falklandicum. s.n. (lectotype,NY; isolectotype,H-SOL).
1. Plantscomplanate; leavesovateto ovate-lanceo- PlagiotheciumbellireteC. Muller, Hedwigia36: 130.
1897. Type.Argentina.Tucumfn, 1873,Lorentzs.n.
late,flatto slightlyconcave,acuteto acuminate;
margins oftenerectto recurved; Mexicoto Tierra (lectotype, H-BR).
del Fuego. Plagiotheciumlepidopiladelphus C. Miller, Bull. Herb.
2. Leavesslenderly median leaf Boissier6:119. 1898. Type. Brazil.Serrado Itatiaia,
long-acuminate; 1900m, Mar 1894, Ule 1862 (lectotype,H-BR);2000
cells5-6(-8) umwide;widespread at highel-
evationsandin southernmost SouthAmeri- m, Feb 1894, Ule 1861 (syntype,H-BR).
ca. ........................ 3. P. lucidum. Plants medium-sized to rather large, in mostly
2. Leavesacute to broadlyshort-acuminate;
medianleafcellsmostly8-20 tmwide. dense mats. Stems creeping to ca. 5 cm long, +
3. Leafapicesbroadlyacute;leafcells14-20 turgid, somewhat to distinctly complanate-foli-
,Lmwide;southernArgentina and Chile. ate, usually only obscurely so. Leaves closely
...................... 5. P. ovalifolium. spaced or less often more
3. Leafapicesnarrowly distantly spaced,
acuteto acuminate; ovate to
leafcells8-15 Mmwide;Mexicoto Tierra spreading, broadly oblong-ovate, 1.1-
del Fuego,SEBrazil. 2.0 mm long, short-acuminate; margins plane or
4. Leafcells(9-)11-15 umwide;decur- erect in lower 2/3,entire or with a few small teeth
rencies prominent,large, of short, at extreme apex; cells linear, subflexuose, 115-
roundedcells ...... 4. P. conostegium. 200 x 6-9
4. Leafcells mostly(6-)8-11 um wide; ,im, thin- to firm-walled; alar cells
decurrencies small,inconspicuous. few, gradually and poorly differentiated; decur-
5. Plantsusuallystronglycomplanate rencies very small, often difficult to demonstrate,
with widelyspacedleaves;decur- when present usually only 1-3 rectangular,

FIG. 1. Plagiotheciumnovo-granatense.A. Habit. B. Leaves. C. Apical leaf cells. D. Median leaf cells. E.
Alar cells. F. Operculatecapsule (wet) and inoperculatecapsule (dry).A, F (lower)from Cleef 9516 (CANM);
B (upper),C (upper),D, E from Lindig s.n. (BM, lectotype);B (lower),C (lower),F (upper)from Glaziou 7363
(BM, holotype of P. brasilense).
SystematicTreatment 5

Et

I mm~~~~~~~~~~~~~i X
/F
4 mm K "i
6 FloraNeotropica

scarcelyenlargedcells, cells 17-22 umwide. Au- 2. Plagiothecium drepanophyllumRenauld &

toicous. Perichaetial leaves broadly oblong, Cardot, Bull. Soc. Roy. Bot. Belgique 41(1):
abruptlyandbroadlyshort-acuminate,uppercells 108. 1905. Type. Costa Rica. Cocos Island,
to 12 Am wide. Setae 2-2.5 cm long, reddish, 1902, Pittiers.n. (holotype, PC). Fig. 3
twisted; capsules inclined to cernuous, some-
integrifoliumBartram,Contr.U.S. Natl.
times arcuate,cylindric,2-2.5 mm long, smooth; Isopterygium
Herb.26: 111. 1928.Type.CostaRica.SanJose:
exothecialcellsirregularlyshort-rectangular, firm- Cerrode las Vueltas,Standley& Valerio43764 (ho-
to thick-walled;operculumconic;exostometeeth lotype,US;isotype,NY).
striate-papillosebelow, papillose above; endo- Plagiothecium standleyiBartram,Bryologist49: 121.
1946. Type. Guatemala.Quetzaltenango: Volcan
stome finely papillose, basal membrane high, SantaMaria,3150 m, Standley67646 (holotype,
segmentskeeled, perforate,cilia 2-3. Spores 11- FH;isotypes,F, US).
14 Am in diam., finely papillose.
Distribution(Fig. 2). Andean Colombia, Ven- Plants medium-sizedto ratherrobust,in com-
ezuela, Ecuador,Peru, Argentina; SE Brazil. planate, mostly thin mats. Stems creepingto ca.
6 cm long, strongly complanate-foliate.Leaves
Specimensexamined.COLOMBIA.ANTIOQUIA:Sur widely spaced, wide-spreading, often falcate,
Jardinca. 5-10 km, 2480 m, Churchill& Marulanda ovateto
15704 (HUA, NY), 15721 (HUA, NY, U). BOYACA: narrowlyoblong-ovate,1.4-2.6 mm long,
Paramode la Sarna,3320 m, Cleef9516(CANM,FLAS, graduallyshort-acuminate,not or scarcelycon-
U). CAUCA:"Canaan,"Mt. Purace,ca. 3100 m, Killip cave; margins plane or often erect nearly
6701, 6792 (NY). CUNDINAMARCA:Andes Bogotenses, throughout, entire or more often densely and
Weir s.n. (NY), 129 (H-BR, NY), 236 (H-BR, NY),
430 (NY). sharply serrulateat extreme apex; cells linear,
ECUADOR.AndesQuitenses,Spruce1056 (BM, mostly straight, 100-160 x (6-)8-11(-14) Am,
H-SOL, NY), 1057 p.p.c.fr.(NY). NAPO:Paramo de thin- to firm-walled;alarcells few, graduallyand
Papallacta, 11,600 ft, Steere 27060 (NY). poorlydifferentiated; decurrenciessmallbut con-
NAPO-PASTAZA: Valleyof rio Chingual E of El Pun, sistentlypresent,2(-3) cells wide, 2-5 cells long,
7500-9000 ft, Steere8923 (NY).
PERU. Chaehapaga,s.d., s.c., s.n. (NY). cuzco: Vie. cells rectangular,somewhat enlargedand inflat-
Cuzco, 4000-10,000 ft, Oct 1893, Jay s.n. (NY). ed, those above mostly 1.5:1, to 40 Am wide,
HUANUCO: Tambo de Vaca, 13,000 ft, Bryan 572 (F, those toward base of decurrency3-5:1, ca. 14
MO,NY). mimwide. Autoicous. Perichaetialleaves similar
BOLIVIA. SanMateo-Sunchal, 2800m, Apr1911, to
vegetativeones but with laxerbasal areolation
Herzog4513a (syntypeof P. conostegium;BM, JE).In
der Quebradade Pocona, 2900 m, Apr 1911, Herzog and not decurrent.Setae 1.3-2.5 cm long, red-
5145 (JE). dish, twisted;capsulesusuallyinclined,cylindric,
BRAZIL.Itatiaia,20 Jun 1902, Dusen s.n. (JE), 16 1.5-2.0 mm long, smooth; exothecialcells irreg-
May 1902, Dusen s.n. (H), 2100 m, Dusen 210, 215, ularly short-rectangular,thin- to firm-walled;
926, 986 (H-BR), 2130 m, Schifer-Verwimp& Ver-
wimp 9549 (NY). operculumconic; exostome teeth cross-striolate
below, papillose above; endostome finely papil-
Plagiotheciumnovo-granatenseis a relatively lose, basalmembranehigh,segmentskeeled,nar-
rarespeciesof high elevations.It is distinguished rowly perforate,cilia 2-3. Spores 11-14 ,umin
by relatively robust plants which are only ob- diam., finely papillose.
scurely complanate; the leaves are short-acu- Distribution(Fig. 4). Mexico to Costa Rica,
minate with cells about 8 Am wide. The most Venezuela,Colombia, Jamaica,Dominican Re-
distinctivefeature,though,is the meagerleaf de- public.
currencydevelopment. Many leaves are not de-
currentbut some leaf decurrenciescan be found Specimensexamined.MEXICO.CHIAPAS:Mt. near
on every stem. When present they are only of San Crist6ballas Casas, Arzeni 21b (TENN); Huey-
tepec near Las Casas, 8200 ft, Sharp 3454 (TENN);
one to threerectangularcells.Plagiotheciumnovo- near El Triunfo,6400 ft, Sharp4462 (TENN);N slope
granatenseis only likely to be confused with P. of SierraN of Mapastec,Sharp4617a (FLAS,MICH,
drepanophyllum,which is its more northern MO, NY, TENN); prope San Crist6bal,2200 m, 24
counterpart.In that species, however, every leaf Nov 1907, Minch s.n. (NY). DISTRITO FEDERAL:
siertode los Leones,9600 ft, Sharp1705 (MICH,
De-
has decurrenciesand the plantsare mostly more
TENN);Cafiadade Contreras,3100 m, Rzedowski
complanate with more distantly spaced, often 19481 (MICH,TENN), 19482 (TENN). MEXICO: Pass
more or less falcate-secundleaves. "Portode la Cruces,"3100 m, Duell2/316 (TENN);
Systematic Treatment 7

I
,0 O^________

c
P.
ovalifolium

\Q... P. falklandicum

'i\ '^ oi I /*

90 . X
\
, 5* v1 2 00 30 E.T
LtS .. /

FIG. 2. Distributionof Plagiotheciumin South America:P. novo-granatense(circles,open circle indicates

unknownspecificlocality);P. ovalifolium(squares);P. falklandicum(star).
8 Flora Neotropica

1;:~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~,

N~~~~
50 crm
II I ~/ 50 .mC:

0.5mm
mm' Q0Smm

'~~~~~~~~~

OfE

f If/~~~~~~~~~~

:::~.
?? ;."

i,
I
mm
1mm~F
Systematic Treatment 9

'-7. I/ .I- -''i ^ ^ I i

| I

', ~' .k i

P.
P, drepanophyllum
"
l P. lucidum
* P. conostegium
SCALE 30 0 2'"
200 *00 600 0 0 *00 50 . 00--- _1 -- . 1 lv ->
/ *?/ 0
L
0 t.,o - .

1 0 too 95
asJC 0 sgo 5 0-I

FIG. 4. Distributionof Plagiotheciumin CentralAmericaand the West Indies:P. drepanophyllum(circles);

P. lucidum(squares);P. conostegium(stars,open star indicates unknownspecificlocality).

road to Paso de Cort6z, 3700 m, Sharp et al. 61a COSTA RICA. CARTAGO:DOS Amigos area, SE of
(CANM, TENN). MICHOACAN:8-10 mi NW and WNW El Empalme, 2680 m, Crosby& Crosby5958 (MO).
of Ciudad Hidalgo, 2850-3000 m, McVaugh 9908 HEREDIA: Volcan Barva, 2600 m, Griffinet al. D175,
(CANM, F, MICH, TENN). MORELOS:Sierra de Chi- D255-a (FLAS).
chinautzin,2300 m, Cdrdenas1061 (CANM, MICH). JAMAICA. PORTLAND:Trail from Portland Gap NW
OAXACA:N of Llano de Las Flores, 2400 m, Iwatsuki towardMurdock'sGap, 5700 ft, Crosby3535 (CANM,
&Sharp3176c, 5325 (TENN),Sharpet al. 133a, 2331a, MICH). ST. THOMAS: Along trail from Portland Gap
2349, 2352, 2354 (CANM, TENN); Cerrode San Fe- to BlueMountainPeak,7000 ft, Crosby3478 (CANM,
lipe, 2600 m, Sharpet al. 2557 (CANM, TENN);near FLAS,MICH);lowerW ridgeof Blue Mt. Peak, 1700-
S. Pedro Yolox, 9000 ft, Vitt 17708 (FLAS). PUEBLA: 1950 m, Maxon 9837 (CANM,NY); New Haven Gap,
Campo ExperimentalSan Juan Tetla, 3480 m, May Britton73 (CANM,MICH,NY), 13 May 1914, Harris
Nah 1394 (CANM, FLAS), 1483 (FLAS). VERACRUZ: s.n. (NY), Nichols 175 (MICH, NY); zwischen Whit-
Helechales,15km NE ofHuayacocotla,JudrezG. 1303 field Hall und PortlandGap, 1600-1700 m, Hegewald
(CANM, TENN); Huayacocotla, Judrez G. 1640 & Hegewald7993 (NY).
(CANM);between Cerrode Chiconquiacoand Santa DOMINICAN REPUBLIC. LA VEGA:Trail to Pinar
Rita, Sharp et al. 2980 (CANM, TENN); Los Huer- Parejo,6150 ft, Zanoniet al. 20200K(JBSD,MO, NY,
fanos, 1400 m, Sharp et al. 1845 (CANM, F, NY, U); 18 km W of Valle Nuevo, 2300 m, Norris et al.
TENN), 1964a (CANM, TENN), 3092b (TENN); Pe- 7569, 7588 (NY).
dregalde las Vigas,Sharpet al. 2828a, 2839a (TENN); VENEZUELA. MERIDA: Paramo de Los Conejos,
roadfrom Peroteto Cofre,3900 m, Sharpet al. 7181a 3300 m, Griffin& L6pezPV-577 (CANM, FLAS),PV-
(TENN). 582 (FLAS),PV-592 (CANM,FLAS,H, NY), PV-614
GUATEMALA. JALAPA: 10 km E of Mataquescunt- (CANM, FLAS). TACHIRA: Paramo de El Zumbador,
la, 18 Feb 1971, Dixon s.n. (FLAS). QUEZALTENANGO: 2900 m, GriffinPV-202 (CANM, FLAS, MICH,MO).
Volcan Santa Maria, Steyermark34081, 34115 (F). TRUJILLO:ParamoEl Jab6n,2900-3300 m, GriffinPV-
TOTONICAPAN: E ofTotonicapan, 10,500 ft, Sharp 2607 1344 (FLAS);Paramode Guirigay,3200 m, Griffin&
(MO, NY, TENN). L6pez PV-1480 (F, FLAS, MO).
EL SALVADOR. SANTA ANA: El Jardin, El Man- COLOMBIA.ANTIOQUIA: Ca. 10 km NNW of Santa
zano, near Los Naranjos,Kovar6b (MICH). Rosa de Osos, 2500 m, Churchillet al. 14104 (CANM,

FIG. 3. Plagiotheciumdrepanophyllum.A. Habit. B. Leaves. C. Apical leaf cells. D. Median leaf cells. E.
Alarcells. F. Capsule(dry).A, F from Griffin& LopezPV-592 (CANM);B-E from Standley67646 (US, isotype
of P. standleyi).
10 Flora Neotropica
HUA, NY). CUNDINAMARCA: Paramo de Cruz Verde, Type.Brazil.SantaCatarina:
SerraGeral,Ule1181
2700 m, Schultes 12170 (FLAS); Paramo de Palacio, (holotype,H-BR).
3400 m, Migdefrau 1560 (MICH).
Plants small to medium-sized in complanate,
Plagiothecium drepanophyllumis the most mostly thin mats. Stems creeping to ca. 8 cm,
common species of the genus in Central America. mostly ca. 2 cm long, scarcelybranched,some-
It is distinguished by its small, but consistently what to obscurely complanate-foliate, some-
present leaf decurrencies. In aspect it is mostly times stem apices becoming flagelliform.Leaves
strongly flattened with distantly spaced leaves. closely spaced, erect to erect-spreading,often
It has most commonly gone under the name Pla- archedwith downward-pointingapices, narrow-
giothecium schraderi, but the type of that name ly ovate, (0.7-)0.85-1.5(-2.1) mm long, gradually
has proved to be P. novo-granatense. and slenderlylong-acuminate;marginsplane or
more typicallyerectalmost throughout,entireor
3. Plagiothecium lucidum (Hooker f. & Wilson) rarelywith 1-2 serrulationsat extremeapex;cells
Paris, Ind. Bryol. 962. 1897. Fig. 5. linear, subflexuose,85-140 x 5-6(-8) Am,thin-
to firm-walled;alar cells few in extreme basal
HypnumlucidumHookerf. & Wilson, LondonJ. Bot. angles, quadrateto short-rectangular; decurren-
3: 554. 1844;HypnumlucidulumHookerf. &Wilson cies off with dissection,
in Hooker f., Fl. antarct.2: 418. 1847, ortho. var. occasionally stripping
fide Newton (1983a); Plagiothecium lucidulum narrow, 1-2(-3) cells wide, 3-4 cells long, cells
(Hookerf. & Wilson) Mitten, J. Linn. Soc., Bot. 12: rectangular,often enlargedand inflated,(1.5-)2-
521. 1869, ortho.var. Type. Chile.Magallanes:Her- 3(-4):1, 14-20 im wide. Autoicous. Perichaetial
mite Island, Cape Horn, 1839-1843, Hooker s.n. leaves similar to
vegetative ones but not decur-
(holotype,BM, n.v.; isotypes, H-SOL, NY-3). rent. Setae 9-17 mm
Hypnum andinum Hampe, Ann. Sci. Nat. Bot. V, 5: long, reddish,twisted;cap-
311. 1866;Plagiotheciumandinum(Hampe)Schim- sules erectto horizontal,short-cylindric,0.5-1.5
per ex Jaeger,Ber. Tatigk.St. GallischenNaturwiss. mm long, smooth; exothecial cells subquadrate
Ges. 1876-1877: 448. 1878. Type. Colombia. Te- to short-rectangular, firm-walled;operculumnot
quendama,2500 m, Aug 1863,Lindigs.n.(lectotype, seen; exostome teeth cross-striolatebelow,
BM; isolectotype,BM);Chiquinquira,2700 m, Sep
pa-
1863, Lindigs.n. (syntype,BM). pillose above with a prominentcenterline;endo-
Plagiotheciumalbidens C. Miller, Hedwigia 40: 57. stome with a high, smooth basal membrane,seg-
1901; Isopterygiumalbidens(C. Miller) Brotherus ments keeled, narrowly perforate, smooth or
in Engler& Prantl, Nat. Pflanzenfam.1(3): 1082.
1908.Type.Brazil.SantaCatharina:SerraGeral,Ule 14
finely roughened,cilia 1-2, papillose. Spores 9-
1215 (lectotype,H-BR). Amin diam., papillose.
Plagiotheciummollicaule Williams, Bull. New York Distribution (Figs. 4, 6). Dominican Republic,
Bot. Gard. 6: 256. 1910. Type. Bolivia. Near Peli- Colombiaand Venezuelato Tierradel Fuego;SE
chuco, 2450 m, 30 Apr 1902, Williams2780 (ho- Brazil.
lotype, NY; isotypes, F, MICH).
Plagiotheciummicrosphaerothecium Herzog,Biblioth. Specimensexamined.DOMINICANREPUBLIC.
Bot. 87: 152. 1916. Type. Bolivia. Im Nebelwald LA VEGA: 12.4 km S of Constanza,6200 ft, Shaw 5704
iiberComarapa,ca. 2600 m, Herzog4250 (holotype, (JBSD,NY). PERAVIA:La Nevera, 6700 ft, Buck 5249
JE; isotype, H-BR). (NY), 5266 (JBSD,NY), 8143, 8690 (NY),Steere23070
Plagiothecium submollicaule Brotherus ex Herzog, (NY), Zanoni et al. 19195T (JBSD, MO, NY).
Biblioth.Bot. 88:23. 1920.Type.Bolivia.Estradillas VENEZUELA.BARINAS: Paramo de Santo Cristo,
fiberIncacorral,Herzog 3327 (lectotype,JE; isolec- 3600-3750 m, Griffin& Diaz M. PV-137a, PV-138
totype, H-BR); Nebelwald fiberComarapa,Herzog (CANM, FLAS). DISTRITOFEDERAL:E of Colonia To-
4326 (syntype,JE). var, 2000 m, Steyermark& Wiehler 106540 (US).
PlagiotheciumaraucarietiBrotherus,Denkschr. Kai- MERIDA: Paramo de Mucubaji,3400 m, Griffinet al.
serl.Akad.Wiss.,Math.-Naturwiss. K1.83:338.1926; 1198, 1351(FLAS);SierraNevadade Merida,4000
Lepidopilumaraucarieti C. Miller ex Brotherus, m, Griffinet al. 275 (FLAS).TACHIRA:
Arribade "Las
Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Copas," 2500-2800 m, Steyermark & Dunsterville
K1.83: 338. 1926, "Lepidolum,"nom. nud. in syn. 100785 (US); ParamoEl Batall6n, 3380 m, Griffinet

FIG. 5. Plagiotheciumlucidum.A. Habit. B. Leaves. C. Apical leaf cells. D. Medianleaf cells. E. Alar cells.
F. Capsules(left:dry; right:wet). A, F (right)from Griffin& L6pez PV-1430 (CANM);B (upper),F (left) from
Lindig s.n. (BM, lectotype of P. andinum);B (lower),C-E from Hookers.n. (NY, isotype).
SystematicTreatment 11

2 mm /

,,
.... . mmI , II

,::. '

F I1mm
12 Flora Neotropica
80 70
60 50 40

1~~~~~~~~~~~~~~~~~~~~~~~1

o I ~~ ---- --
i--?e

30
0^

/I

l\p
90 so

MrnSothAerc:|
50

0
I.
P. lucidum-UiE 20

FI. .DitibtonofPagoheim luidm(crle) P onstgum(rinle)

i1- \ X-- - I\Z | X P. conostegium

' S
\ .';?</'?' \ ^~~^~/ ' \CALE /
0 _0
FIG. 6. Distribution of Plagiothecium in South America: P. Iucidum (circles); P. conostegium (triangles).
\ <S^ T^'Tr'^.S--/ ~~~~~~~~~~~~~SINuSOIDAL
PROJECTION

G0
6t
Systematic Treatment 13

al. 597 (FLAS);ParamoLa Negra, 3000 m, Griffinet slightlywider leaf cells. However, in overall leaf
al. 2088 (FLAS). TRUJILLO:Paramo de Guirigay, 3400 shapeand microscopicdetails the two forms are
m, Griffin& L6pez F. PV-1566 (FLAS);Paramo El identicaland frequentintermediatesareencoun-
Jab6n, 2900-3300 m, Griffin & L6pez PV-1430
(CANM, FLAS),PV-1453 (FLAS). tered. We have thus synonymized them. Ad-
COLOMBIA. ANTIOQUIA: Alto de Boquer6n, mittedly, in the extremes the two are quite dif-
Churchill et al. 14233 (HUA, NY). CUNDINAMARCA: ferent in aspect and additional study may show
Paramode Chisaca,3460 m, van der Hammen & Ja- some justification for the recognitionof P. an-
ramillo-M.4351, 4373, 4387 (FLAS,U); paramosentre
Coguay San Cayetano,3700 m, Florschiitz3642 (U);
dinum at some taxonomic level.
Cuchilla El Tablazo, 3260 m, Linares & Bulla 203, Plagiotheciumlucidumseems most relatedto
544a (FLAS);Tequendama,2500 m, Mar 1863, Lindig the northtemperateP. laetum B.S.G. but differs
s.n. (BM, H-BR, H-SOL, NY-2). in a number of features. The former has sym-
ECUADOR. AZUAY: Area Nacional de Recreaci6n
"Cajas," 3950 m, Lewis 78-2424 (F, NY). BO- metric,long-acuminateleaves with planeto erect
LiVAR-CHIMBORAZO: Summit of road from Riobamba margins,leaf apicesentireor rarelywith 1-2 ser-
to Guaranda,4120 m, Lewis 78-2595, 78-2599 (F, rulations and it has never been observed with
NY). COTOPAXI:Parque Nacional Cotopaxi, 4100-4300 brood-bodieson the stems. On the other hand,
m, Balslev et al. 3379 (NY, QCA). IMBABURA: Lago P. laetumhas mostly asymmetric,acuteto short-
SanMarcos,11,200ft, Cazalet&Pennington69 (MICH,
NY). NAPO: S side of Cerro Sumaco, 3700-3800 m,
acuminate leaves with margins narrowly re-
Lojtnant & Molau 12971 (NY). NAPO-PASTAZA:Vicin- curved nearlyto the apex, leaf apices often ser-
ity of La Bonita, nearjct with rio Suci, 5000-6500 ft, rulate and 3-6-celled fusiform brood-bodies
Steere8600 (NY);La Fama,6500-7000 ft, Steere8504 commonlyoccurringon the stems. We have con-
(NY). PICHINCHA:Volcan Cayambe, W shore Laguna sidered the possibility that the two are synony-
SanMarcos,3450 m, Steere27593 (NY);Quito,Jame-
son 71 (MICH, NY). ZAMORA-CHINCHIPE: Km 17 on mous and that these differentiaeare not impor-
Loja-ZamoraRd., 2400 m, Holm-Nielsenet al. 3573 tantenoughto warrantrecognitionof two separate
(US). species. However, because of the great disjunc-
PERU. AMAZONAS:Strasse Chachapoyas-Cajamar-
ca oberhalbLeimebambakm 417, 2800 m, Frahm et tion, plus the fact that P. lucidum is the older
al. 731, 986 (NY, U); Philippi2334 (U). name, we have delayed action on the decision.
BOLIVIA.ZwischenSan Mateo und Sunchal,Apr It is most likely that the two are best considered
1911, Herzog s.n. (JE). Quillapotunea,Mandon 58 vicariantspecies.
(BM).
BRAZIL.Serra do Itatiaia, May 1902, Dusen s.n.
(NY), 2100 m, Dusen 300 (H-BR). 4. PlagiotheciumconostegiumHerzog,Biblioth.
ARGENTINA. TIERRADEL FUEGO: Bahia Buen Su- Bot. 87:154. 1916. Type. Bolivia. Waldgrenze
ceso, Matteri3565, 3615 (BA). iiberTablas, 3400 m, Herzog 2947 (lectotype,
CHILE. MAGALLANES: Tierra del Fuego, Escandallo
Bay, Seno AlmiranteMartinez,30 Jan 1966, Dix s.n. JE);zwischenSan Mateo und Sunchal,Herzog
(US); PuertoGomez, Halle & Skottsberg865 (H-BR). 4513a (syntypes,BM, JE;note: this syntypeis
not P. conostegium,but ratherP. novo-gran-
Plagiothecium lucidum is characterized by
atense);im Nebelwald iiber Comarapa,2600
slenderly long-acuminate leaf apices and leaf cells
m, Herzog 4239 (syntype,n.v.). Fig. 7.
mostly about 6 ,m wide. Although described
from southernmost Chile, it is most common in Plants medium-sized, in complanate, mostly
the northern Andes, and then is disjunct to the thin mats mixed with othermosses. Stems creep-
central highlands of the Dominican Republic. It ing to ca. 4 cm long, complanate-foliate.Leaves
has most often been called P. lucidulum, but closely spaced,spreadingto wide-spreading,1.4-
Newton (1983a) has demonstrated that this name 2.0 mm long, graduallyor ? abruptlyshort-acu-
is in fact an orthographic variant. Because of the minate, slightly concave; marginsplane or nar-
slender size of the plants, this is one of the easiest rowly recurved below, subentire or with a few
species of Plagiothecium to recognize in the field. small teeth at extremeapex;cells linear, straight
The frequent occurrence of flagelliform branches orsubflexuose,110-145 x (9-)11-15 tm, mostly
is also useful in field identification. thin-walled;alar cells few, poorly differentiated,
The type specimens of Plagiothecium andi- rectangular;decurrenciesoften strippingoff with
num and P. araucarieti represent robust forms dissection, 2-3 cells wide, to 8 cells long, cells
of the species. They differ from typical material quadrateto short-rectangular,enlargedand in-
only in the size of the plants and leaves and in flated, 1.5-3:1, 14-23 ,umwide. Autoicous and
14 Flora Neotropica

0.5 mm

B m j
D c I'

D:

,'...\VI:..':i ? . .. ..iC ... _

,.
Systematic Treatment 15

possibly dioicous; perigonia very small in leaf Plagiotheciumconostegiumis distinguishedby

axils. Perichaetial leaves similar to vegetative its relativelybroad leaf cells (11-15 um) and es-
ones but with laxer basal areolation and not de- pecially by its large leaf decurrenciescomposed
current. Setae 1.5-2.5 cm long, reddish, twisted; of short, rounded cells. For these reasons it is
capsules inclined to horizontal, cylindric, ca. 2 easily distinguishedfrom P. novo-granatense,P.
mm long, usually striate or furrowed; exothecial drepanophyllumand P. lucidum.Plagiothecium
cells subquadrate to narrowly rectangular (longer conostegiumis probablymost closely relatedto
in striations), firm- to thick-walled; operculum P. ovalifolium,which also has wide leaf cells and
short-rostrate; exostome teeth striate-papillose large decurrencies.However, that species has
below, papillose above; endostome finely papil- acute leaf apices and leaf cells much shorterand
lose, basal membrane high, segments keeled, not somewhat broader; it is restricted, in South
or very narrowly perforate, cilia 3. Spores 12-15 America, to Tierra del Fuego. In the same lo-
,um in diam., finely papillose. calities,P. conostegiumis a forestspecies,mostly
Distribution (Figs. 4, 6). At high elevations in growingon soil and rottenlogs, whereasP. ovali-
Mexico, Guatemala and the Dominican Repub- folium grows in more exposed areas, often on
lic; northern Andes; Tierra del Fuego. rocks.
Specimens examined. MEXICO. DISTRITO
FEDERAL:
Desierto de los Leones, 10,300 ft, Sharp 63 (CANM,
MICH, NY, TENN). MEXICO: Gap at Nevada de To- 5. PlagiotheciumovalifoliumCardot,Bull. Herb.
luca, 4350 m, Sharpet al. 1536a (TENN). Boissier II, 5: 1010. 1905. Type. Argentina.
GUATEMALA.Withoutlocality or collector(NY).
DOMINICAN REPUBLIC. LA VEGA: 4.7 km S of Tierradel Fuego:In monte supraUshuaia, 530
Constanza,then 8 km toward Pinar Parejo, 6150 ft, m, 6 Mar 1902, Skottsberg177 (lectotype,PC;
Buck 7821, 7844 (NY); 12.4 km S of Constanza,6200 isolectotype, S); in silva ad Bahia Lapataia,9
ft, Shaw 5701A (NY). PERAVIA:
La Nevera, 6700 ft, Oct 1902, Skottsberg178 (syntype,S-2).
Buck 8144 (NY), Steere 23165 (NY), Zanoni et al.
19195K (JBSD, NY). SAN JUAN:Pico Duarte, 3000- Fig. 8.
3150 m, Buck 14268 (NY). Plants medium-sized in complanate, mostly
VENEZUELA. TRUJILLO:
Paramo El Jab6n, 2900-
3300 m, Griffin&L6pezF. PV-1306 (FLAS),PV-1330 thin mats. Stems creepingor + ascendingto ca.
(F, FLAS, NY). 3 cm long, complanate-foliate.Leaves widely
COLOMBIA. BOYACA:
Paramo de la Rusia, 3735 m, spaced, spreading,ovate to oblong-ovate, 1.1-
Cleef 7088 (CANM, FLAS, U); paramos al NW de 1.4 mm long, graduallyand broadlyacute;mar-
Belen, Alto de las Crucesy alrededores,3765 m, Cleef
2288 (CANM, FLAS, U). gins recurved almost throughout, entire; cells
ECUADOR. CARCHI: Paramo above Tifinfo, 11,000- long-hexagonal,85-130 x (9-)14-20 um, thin-
12,000 ft, Steere 9453 (NY). IMBABURA: Otovalo-La- walled; alar cells few, poorly differentiated;de-
guna Mojanadaroad, 12,250 ft, Steere26693 (NY); E currenciesstrippingoff with dissection, broad,
side of CerroImbaburaabove La Esperanza,4040 m, 3-4 cells wide, to 10 cells long, cells rectangular
Lewis 78-2989 (F, FLAS, NY, U), 78-3040 (F). PI-
CHINCHA:Just W of summit along Quito-Baeza road,
to quadrate,sometimes somewhat inflated and
4000 m, Steere E-189 (NY). TUNGURAHUA: Volcan spherical,mostly 2-4:1, 14-20 Amwide. Autoi-
Tunguragua,Spruce1057 p.p.st. (NY), 3690 m, Lewis cous.Perichaetialleavesbroadlyoblongwith cells
78-3406 (F). 6-9 um wide, thin-walled.Setae 2-2.5 cm long,
ARGENTINA. TIERRADEL FUEGO:Monte Olivia,
Matteri 1831 (BA); Paso GaribaldiLaderaEste, 650 reddish,twisted;capsulesinclined, cylindric,ca.
m, Matteri1921 (BA);Caminoal GlaciarMartial,Mat- 2.5 mm long;exothecialcells subquadrate,firm-
teri 1790 (BA);Rutaj, Matteri1829 (BA);BahiaBuen walled;operculumconic;exostome teeth striate-
Suceso, Matteri 3528 (BA); I. Estados, Pt. San S. de papillose below, papillose above; endostome
Sabramento,Matteri1376, 1393 (BA),I. Troffr6,Mat- papillose-spiculose,basal membrane high, seg-
teri 1242 (BA).
CHILI. MAGALLANES: mentskeeled,stronglyperforate,cilia mostly sin-
Depto. Ultiana, Esperanza,
MatteriTBPA-B2754(BA);HermiteIsland,CapeHorn, gle. Spores 9-12 um in diam., finely papillose.
1839-1843, Hookers.n. (NY). Distribution(Fig. 2). SouthernmostArgentina

FIG. 7. Plagiotheciumconostegium.A. Leaves. B. Apical leaf cells. C. Median leaf cells. D. Alar cells. E.
Capsules(dry).A-D from Herzog 2947 (JE, lectotype);E from Cleef7088 (CANM).
16 Flora Neotropica

0.5 m

'''

A~~~~'

., V

50.I.

mmjim

"C NN~~~~~~~~~~~~~~~~~~~~~~~~~~

K R> C~~~~~~~~~~~~~~~~~~~~~~~~~~~~?

.2 >9 22I~\ \ ~
tPD
E~~~~~~~~~~~~~~~?

1mm
Systematic Treatment 17

and Chile; also reported from non-American 6. Plagiothecium falklandicum (Cardot &
subantarcticislands. Brotherus)Newton, Brit. Antarct. Surv. Bull.
66: 86. 1985. Fig. 9.
Specimensexamined.ARGENTINA. WITHOUT DE-
PARTMENT: Patagonia, 1896-1897, Hatcher s.n. Plagiothecium roeseanumSchimperin B.S.G. var.
(CANM), P-118 (NY). SANTA CRUZ:Lago Argentino, Cardot& Brotherus,
falklandicum Kongl.Svenska
Bahia Onelli, Matteri 4532 (BA). TIERRA DEL FUEGO: Vetenskapsakad.Handl.63(10):66. 1923. Type.
RutaComplementaria J, Km 24, Matteri4192 (BA); Falkland Islands.WestFalkland: Mt.Adam,13Dec
ArroyoGrande,Rinc6nde lasSieteCascadas, Matteri 1907,Halle&Skottsberg 872(holotype,PC;isotype,
3232,4371(BA);Vallede TierraMayor,Matteri2506 S).
(BA,CANM);CaminoalGlaciarMartial,300m,Mat-
teri1870(BA);Lapataia, caminoa Laguna Negra,Mat- Plants robust in dense to thin mats. Stems
teri3807(BA);LagoFagnano,Cabecera Lagoat E end julaceous, often ascending, to ca. 8 cm long,
of LagoFagnano,SantessonM706(S);I. Estados,I.
Observatorio, Matteri1231(BA). scarcelybranched.Leavesclosely imbricate,ob-
CHILE.MAGALLANES: Fuegiamedia, LagoLinch, 19 ovate to oblong-ovate,1.4-2.3 mm long (in Falk-
Dec 1928,Roivainen s.n.(CANM,NY, S);PuntaAre- land material, apparentlylargerin extralimital
nas, 31 Dec 1898,Hatchers.n. (NY);Aynardodalen, material),abruptlyacute,concave;marginsplane,
300 m, Halle&Skottsberg 866 (S). entire;cells linear,straightto subflexuose,mostly
Plagiotheciumovalifoliumis characterizedby 75-145 x 9-11 Aum, ? thin-walled;alarcells well
its acute leaves (the only mainlandSouth Amer- developed in 4-5 cell rows across the insertion,
ican species of Plagiotheciumwith this feature), 4-6 cells up the margin,quadrateto short-rect-
its short, broad leaf cells and its largedecurren- angular;decurrenciesusually strippingoff with
cies. In additionto the distinctionsfromP. cono- dissection,broad,3-4 cells wide, to 10 cells long,
stegiumnoted above, there are also several spo- cells short-rectangularto quadrate, often en-
rophytic differentiae: the capsules are never largedand inflated,1-2.5:1, 23-28 ,umwide. Au-
striate,the operculumis shorter,the endostome toicous. [Sporophytesnot known from the Falk-
is much more stronglyornamentedwith the seg- lands; the following description adapted from
ments more perforate,and the cilia are single. Newton (1983b).] Perichaetial leaves broadly
Additionally, the spores are smaller. Although spathulate. Setae (1.2-)1.6-2.2(-2.8) cm long,
tedious to discern, these features can assist in orange;capsuleserectto inclined, cylindric, 1.5-
problematicspecimens. 2.5 mm long;exothecialcells roundedquadrate,
Plagiothecium denticulatum (Hedw.) B.S.G. thick-walled;operculumconic to short-rostrate;
var. obtusifolium(Turn.)Moore, frequentlycon- exostome teeth striate-papillosebelow, papillose
sidered a distinct variety, especially in Europe above; endostome with a high basal membrane,
and Japan(Greene, 1957; Nyholm, 1965; Iwat- segments keeled, narrowlyperforate,cilia rudi-
suki, 1970; Iwatsuki & Noguchi, 1973; Smith, mentary or absent. Spores (11-)12.5-14(-17.5)
1978), is morphologicallyclose to P. ovalifolium. ,umin diam., finely papillose.
The variety is often recognizedby its occasional Distribution(Fig. 2). In SouthAmericaknown
julaceousplants with symmetric,imbricate, con- only from the Falkland Islands; reported by
cave leaves that commonly have obtuse apices. Newton (1983b) from South Georgia.
This contrastswith P. ovalifolium,whose plants
Specimenexamined.Onlythe typeseen.
are always complanatewith symmetric,widely-
spaced,mainly flat leaves with acute apices. The Plagiotheciumfalklandicum is distinct in the
varietyis extremelyvariable,however,with some julaceous,erectplantswith abruptlyacuteleaves.
plants approachingP. ovalifoliumin many fea- The alar cells are better developed than in any
tures. Furthercollectionsand studies of P. ovali- other South Americantaxon. Since it is known
folium may find the two to be synonymous. in our rangeonly from the Falklands(Islas Mal-

FIG. 8. Plagiotheciumovalifolium.A. Leaves. B. Apical leaf cells. C. Median leaf cells. D. Alar cells. E.
Operculatecapsule (dry) and inoperculatecapsule (wet). A-D from Skottsberg177 (PC, lectotype);E from
Roivainens.n. (CANM).
18 Flora Neotropica
,.
.: . ,~-...~.:.. .. .

0.5 mm E.

~~~~~~~~B~~~~~~~~~~~~~~~~~~~~~~~~~~

t~~~~~~~~~~~~~~

50: pm

D
Doubtful Names and Excluded Taxa 19

vinas), and no otherspeciesof Plagiotheciumare Basedon:Brazil.Amazonas:Rio Negro,Spruce

knownfrom that archipelago,there should be no 1058 (BM).
confusion in identification. =IsopterygiumtenerifoliumMitt.
PlagiotheciumorthocarpumMitten,J. Linn.Soc.,
DOUBTFUL NAMES AND Bot. 12: 521. 1869; Isothecium orthocarpum
EXCLUDED TAXA Bridel, Bryol. Univ. 2: 387. 1827, nom. nud.;
HypnumorthocarpumHornschuchex C. Miil-
Plagiotheciumbicolor Warstorf, Hedwigia 57: ler, Syn. Musc. Frond. 2: 253. 1851, hom. il-
116. 1915. Type. Brazil. Mt. Roraima, Ule leg., non Angstr., 1842. Type. Chile. Talca-
2476. We have been unable to locate the type huano, Chamisso 1816. No specimen is at B
or any materialwith this name. Also, we have or NY, althoughone should not expecta spec-
seen no material of any species of Plagiothe- imen at NY since Mitten knew the name only
cium from Mt. Roraimaon the Brazilian/Ven- from the literatureas indicated purely by his
ezuelan/Guyananborder. citation of Miiller'searlierdescription.From
Plagiothecium chrysobasisC. Miiller in Pola- the description it is possible that this is an
kowsky,J. Bot. 15:229. 1877, nom. nud. Based older name for P. ovalifoliumbut without a
on: CostaRica. Cartago:Angostura,Nov 1875, specimen we are not preparedto adopt the
Polakowsky495 (BM). name.
=Pilosium chlorophyllum(Hornsch.)C. Mill. Plagiotheciumproximum C. Miiller, Hedwigia
PlagiotheciumcurtipesSpruce,Cat. Musc.Amaz. 38(Beibl.): 59. 1899, nom. nud.; Plagiothe-
And. 13. 1867, nom. nud. Based on: Brazil. cium restingaeC. Miillervar. tenueC. Miiller,
Amazonas:Rio Negro, Spruce872 (NY). Hedwigia 40: 60. 1901. Type. Brazil. Rio de
=Isopterygium cf. subtenerrimum (C. Mill.) Janeiro: Nova Friburgo,Apr 1895, Ule 233
Par. (lectotype,H-BR; isolectotype, NY).
=Isopterygium tenerum (Sw.) Mitt. (fulvum
Plagiotheciumfallax Herzog, Biblioth. Bot. 87: form)
152. 1916, hom. illeg., non Card. & Ther.,
1902; Plagiothecium herzogii Theriot, Rev. PlagiotheciumregnelliiAngstrom,Ofvers.Forh.
Bryol. Lichenol. 4: 182. 1932. Type. Bolivia. Kongl. Svenska Vetensk.-Akad. 33(4): 44.
Im Bergwalddes Rio Tocorani, ca. 2200 m, 1876. Type. Brazil. Caldas, Regnell 15 (lec-
sehrsparlich,Herzog4064. This specimenwas totype, S-herb.Angstr.;isolectotypes, S-herb.
not locatedat eitherJE or PC. The description Regnell- 2).
suggestsit may be another synonym of P. lu- =Lepidopilidiumregnellii(Angstr6m)Buck &
cidum, if it is even in the genus. The collen- Ireland,comb. nov.
chymatous exothecial cells mentioned in the Plagiotheciumvillae-ricaeBescherelle,Mem. Soc.
protologue would exclude this species from Sci. Nat. Cherbourg21: 271. 1877. Type. Par-
Plagiothecium and leave its correct generic aguay, Guaira:Villa Rica, Balansa 1210 (ho-
placementin doubt. lotype, BM, n.v.; isotype, NY).
C.
Plagiotheciumfontigenum Miiller, Hedwigia =IsopterygiumtenerifoliumMitt.
40:58. 1901. Type. Brazil.SantaCatarina:Ser-
ra do Oratorio, Ule 521 (lectotype,H-BR).
=Isopterygiumfontigenum(C. Miiller)Buck& ACKNOWLEDGMENTS
Ireland, comb. nov.
We thankthe curatorsof the herbariacited for
Plagiothecium inordinatum Mitten ex Spruce, the loan of specimens,without which this study
Cat. Musc. Amaz. And. 16. 1867, nom. nud. would have been impossible. We gratefullyac-

FIG. 9 Plagiotheciumfalklandicum.A. Habit. B. Leaves. C. Apical leaf cells. D. Median leaf cells. E. Alar
cells. A-E from Halle & Skottsberg872 (PC, holotype).
20 Flora Neotropica

knowledgeLinda M. Ley for the preparationof its related genera from Japan and her adjacent
the illustrations. areas,I. J. Hattori Bot. Lab. 33: 331-380.
& A. Noguchi. 1973. Index MuscorumJa-
ponicarum.J. Hattori Bot. Lab. 37: 299-418.
Newton,M. E. 1983a. The identityof Plagiothecium
LITERATURECITED lucidulum(Hook. f. et Wils.) Mitt. J. Bryol. 12:
617.
Bartram,E. B. 1949. Mosses of Guatemala. Fiel- .1983b. A new speciesof the moss genusPla-
diana, Bot. 25: 1-442. giotheciumfromSouthGeorgia.Brit.Antarct.Surv.
Buck,W. R. & R. R. Ireland. 1985. A reclassification Bull. 60: 63-67.
of the Plagiotheciaceae.Nova Hedwigia 41: 89- Nyholm, E. 1965. Illustratedmoss flora of Fenno-
125. scandia. Vol. II. Musci. Fasc. 5: 407-647. Gle-
Greene,S. W. 1957. The British species of the Pla- erups,Lund, Sweden.
giothecium denticulatum-P. silvaticum group. Smith, A. J. E. 1978. The moss floraof Britainand
Trans.Brit. Bryol. Soc. 3: 181-190. Ireland.CambridgeUniversityPress,Cambridge,
Iwatsuki,Z. 1970. A revision of Plagiotheciumand England.706 pp.

INDEX OF SCIENTIFICNAMES
New combinationsare in boldfaceand synonymsare in italics. Page numbersin boldfaceindicate
primarypage references.Pages with an asterisk(*) indicate pages with illustrationsor maps.

Acrocladiopsis3 bicolor19
draytonii3 brasiliense4, 5*
Catagonium2 chrysobasis19
sect. Acrocladiopsis3 conostegium4, 6, 9*, 12*, 13, 14*, 15, 17
Herzogiella2 curtipes19
Hypnaceaesubfam.Plagiothecioideae2 denticulatum3
Hypnum andinum 10 var. obtusifolium 17
brasiliense4 drepanophyllum4, 6, 8*, 9*, 10, 15
lucidulum10 falklandicum2, 4, 7*, 17, 18*, 19
lucidum 10 fallax 19
novo-granatense4 fontigenum 19
orthocarpum19 herzogii 19
schraderi4 inordinatum19
subaequans4 laetum 13
Isopterygiopsis2 lepidopiladelphus 4
Isopterygium2 lucidulum10, 13
albidens 10 lucidum 4, 9*, 10, 11*, 12*, 13, 15, 19
fontigenum19 microsphaerothecium 10
integrifolium6 mollicaule10
subtenerrimum19 novo-granatense4, 5*, 6, 7*, 10, 13, 15
tenerifolium19 orthocarpum19
tenerum 19 ovalifolium 2, 4, 7*, 15, 16*, 17, 19
Isotheciumorthocarpum19 proximum 19
Lepidolum10 regnellii 19
Lepidopilidiumregnellii 19 restingaevar. tenue 19
Lepidopilumaraucarieti10 roeseanumvar.falklandicum 17
Philoscia 3 schraderi4, 10
latebricola3 standleyi6
Pilosium chlorophyllum19 subaequans4
Plagiotheciaceae2 submollicaule10
Plagiotheciella3 villae-ricae19
latebricola3 Saviczia 3
Plagiothecium2, 3, 4, 19 obtusissima3
albidens 10 Stereodonsect. Plagiothecium3
andinum 10, 11*, 13 Stereophyllum2
araucarieti10, 13 Taxiphyllum2
bellirete4