Smithsonian Institution

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smithsonian contributions to botany • number 93

A Chronology
Some Latin American of
Middle
GeneraMissouri
of Hypnaceae Plains
Village
(Musci)Sites

ByRobert
Craig R.
M.Ireland
Johnson
and William R. Buck
with contributions by

Stanley A. Ahler, Herbert Haas, and Georges Bonani

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smithsonian contributions to botany • number 93

Some Latin American

Genera of Hypnaceae
(Musci)

Robert R. Ireland
and William R. Buck

WASHINGTON D.C.
2009
ABSTRACT
Ireland, Robert R. and William R. Buck. Some Latin American Genera of Hypnaceae (Musci). Smithso-
nian Contributions to Botany, number 93, viii + 97 pages, 37 figures, 2009.—A taxonomic revision of
16 of the 21 genera of the large pleurocarpous moss family Hypnaceae is presented for Latin America
(Mexico, Caribbean, Central America, and South America). This study includes the genera Ctenidium
(Schimp.) Mitt., Elmeriobryum Broth., Herzogiella Broth., Homomallium (Schimp.) Loeske, Irelandia
W. R. Buck, Isopterygiopsis Z. Iwats., Isopterygium Mitt., Phyllodon Bruch & Schimp. in Bruch,
Schimp. & W. Gümbel, Platygyriella Cardot, Platygyrium Schimp. in Bruch, Schimp. & W. Gümbel,
Pseudotaxiphyllum Z. Iwats., Puiggariopsis M. Menzel, Pylaisia Bruch & Schimp. in Bruch, Schimp.
& W. Gümbel, Rhacopilopsis Renauld & Cardot, Syringothecium Mitt., and Taxiphyllum M. Fleisch.
A total of 35 species are recognized for the entire 16 genera. All taxa were studied taxonomically with
each genus and its associated species described in detail. Included in this revision are a description of
the family, keys to the genera and species in the Hypnaceae, detailed descriptions of each species with
complete information on the known distribution throughout Latin America, habitat information, and
nomenclature, and discussions on distinguishing features. Illustrations and distribution maps are pro-
vided for many of the species. The following six names are lectotypified: Hypnum micans Sw., Isop-
terygium cylindricarpum Cardot, Isopterygium planissimum Mitt., Plagiothecium bolivianum Broth.,
Plagiothecium restingae var. tenue Müll Hal., and Stereodon distichaceus Mitt.

Cover images, left to right: Figures 1A, 24I, and 32A (rotated 90˚).

Published by Smithsonian Institution Scholarly Press

P.O. Box 37012 / MRC 957
Washington, D.C. 20013-7012
www.scholarlypress.si.edu

Library of Congress Cataloging-in-Publication Data

Ireland, Robert R.
Some Latin American genera of Hypnaceae (Musci) / Robert R. Ireland and William R. Buck.
p. cm. — (Smithsonian contributions to botany ; no. 93)
Includes bibliographical references and index.
1. Mosses—Latin America. I. Buck, William R., 1950– II. Title. III. Series: Smithsonian contribu-
tions to botany ; no. 93.
QK541.8.I74 2009
588'.2—dc22 2009014948

ISSN (print): 0081-024X, ISSN (online): 1938-2812

The paper used in this publication meets the minimum requirements of the American National Standard
for Permanence of Paper for Printed Library Materials Z39.48–1992.
 Dedication

This work is dedicated to the memory of

Dr.Wilfred B. Schofield (1927–2008), Professor Emeritus,
University of British Columbia, Vancouver, Canada—
A great friend, an excellent bryologist,
and an authority of the Hypnaceae.
Contents

LIST OF FIGURES vii

INTRODUCTION 1

SYSTEMATIC TREATMENT 3
Family Hypnaceae Schimp. 3
Key to the Genera 4
1. Ctenidium (Schimp.) Mitt. 5
Ctenidium malacodes 6
2. Elmeriobryum Broth. 7
Elmeriobryum guatemalense 8
3. Herzogiella Broth. 9
Herzogiella cylindricarpa 11
4. Homomallium (Schimp.) Loeske 14
Key to the Species of Homomallium 14
1. Homomallium mexicanum 15
2. Homomallium sharpii 15
5. Irelandia W. R. Buck 16
Irelandia robusticaulis 16
6. Isopterygiopsis Z. Iwats. 18
Key to the Species of Isopterygiopsis 19
1. Isopterygiopsis muelleriana 19
2. Isopterygiopsis pulchella 20
7. Isopterygium Mitt. 23
Key to the Species of Isopterygium 25
1. Isopterygium acutifolium 25
2. Isopterygium affusum 25
3. Isopterygium byssobolax 30
4. Isopterygium jamaicense 32
5. Isopterygium subbrevisetum 34
6. Isopterygium subglobosum 38
7. Isopterygium tenerifolium 39
 8. Isopterygium tenerum 43
Isopterygium Excluded Taxa 51
Isopterygium Nomina Dubia 51
8. Phyllodon Bruch & Schimp. in Bruch, Schimp. & W. Gümbel 52
Phyllodon truncatulus 53
9. Platygyriella Cardot 54
Key to the Species of Platygyriella 55
1. Platygyriella densa 55
2. Platygyriella pringlei 56
10. Platygyrium Schimp. in Bruch, Schimp. & W. Gümbel 57
Platygyrium fuscoluteum 58
11. Pseudotaxiphyllum Z. Iwats. 58
Key to the Species of Pseudotaxiphyllum 59
1. Pseudotaxiphyllum distichaceum 59
2. Pseudotaxiphyllum elegans 62
3. Pseudotaxiphyllum homomallifolium 62
4. Pseudotaxiphyllum richardsii 64
Pseudotaxiphyllum Excluded Taxa 67
12. Puiggariopsis Menzel 67
Puiggariopsis aurifolia 67
13. Pylaisia Schimp. in Bruch, Schimp. & W. Gümbel 68
Key to the Species of Pylaisia 69
1. Pylaisia falcata 69
2. Pylaisia polyantha 70
3. Pylaisia selwynii 71
14. Rhacopilopsis Renauld & Cardot 71
Rhacopilopsis trinitensis 72
15. Syringothecium Mitt. 74
Key to the Species of Syringothecium 74
1. Syringothecium brasiliense 74
2. Syringothecium sprucei 75
16. Taxiphyllum M. Fleisch. 75
Key to the Species of Taxiphyllum 76
1. Taxiphyllum deplanatum 76
2. Taxiphyllum laevifolium 79
3. Taxiphyllum ligulaefolium 79
4. Taxiphyllum taxirameum 82
Taxiphyllum Excluded Taxa 86

ACKNOWLEDGMENTS 87

REFERENCES 89

INDEX OF SCIENTIFIC NAMES 93

Figures

1. Elmeriobryum guatemalense 9
2. Distribution of Elmeriobryum guatemalense in Latin America 10
3. Herzogiella cylindricarpa 12
4. Distribution of Herzogiella cylindricarpa in Latin America 13
5. Irelandia robusticaulis 17
6. Distribution of Irelandia robusticaulis in Latin America 18
7. Isopterygiopsis muelleriana 20
8. Distribution of Isopterygiopsis muelleriana and
I. pulchella in Mexico 21
9. Isopterygiopsis pulchella 22
10. Distribution of Isopterygiopsis pulchella in South America 23
11. Isopterygium acutifolium 26
12. Distribution of Isopterygium affusum, I. byssobolax,
I. subbrevisetum, I. subglobosum, and I. acutifolium
in South America 27
13. Isopterygium affusum 28
14. Distribution of Isopterygium affusum and I. subbrevisetum
in Central America 29
15. Distribution of Isopterygium tenerifolium, I. jamaicense,
I. subbrevisetum, and I. affusum in the Caribbean 30
16. Isopterygium byssobolax 31
17. Isopterygium jamaicense 33
18. Distribution of Isopterygium tenerifolium, I. tenerum,
and I. jamaicense in Mexico 34
19. Isopterygium subbrevisetum 35
20. Isopterygium subglobosum 38
21. Isopterygium tenerifolium 40
22. Distribution of Isopterygium tenerum, I. tenerifolium,
and I. jamaicense in Central America 41
23. Distribution of Isopterygium tenerum and I. tenerifolium
in South America 42
24. Isopterygium tenerum 46
25. Distribution of Isopterygium tenerum in the Caribbean 47
26. Pseudotaxiphyllum distichaceum 60
27. Distribution of Pseudotaxiphyllum distichaceum,
P. homomallifolium, and P. richardsii in Latin America 61
28. Pseudotaxiphyllum elegans 63
29. Distribution of Pseudotaxiphyllum elegans in Latin America 64
30. Pseudotaxiphyllum homomallifolium 65
31. Pseudotaxiphyllum richardsii 66
32. Taxiphyllum deplanatum 77
33. Distribution of Taxiphyllum deplanatum, T. laevifolium,
and T. ligulaefolium in Latin America 78
34. Taxiphyllum laevifolium 80
35. Taxiphyllum ligulaefolium 81
36. Taxiphyllum taxirameum 83
37. Distribution of Taxiphyllum taxirameum in Latin America 84
 Some Latin American Genera
of Hypnaceae (Musci)

INTRODUCTION

The family Hypnaceae is defined by plants with leaves typically homomal-

lous to falcate-secund, with a short, double costa, and alar cells often differ-
entiated. The peristome is typically well developed with the exostome teeth
cross-striolate on the front surface and with prominent trabeculae at back; the
endostome has a high basal membrane, broad segments, and cilia. However,
any of the peristomial features may be reduced when the capsule is erect and the
plants are corticolous.
The family is very heterogeneous and is little more than those Hypnalean
pleurocarps with a short, double costa that have not been placed in other fami-
lies. It may well prove worthwhile, after greater study, to break the family into
more natural groups. On the other hand, though, some of the current subfami-
lies, most notably the Pylaisioideae, are probably conglomerations of conver-
gences on ecological morphotypes and thus polyphyletic.
Recent work has only emphasized the heterogeneity of the Hypnaceae.
Nishimura et al. (1984) tried to define the Hypnaceae on the basis of seven char-
acters: (1) paraphyllia absent; (2) costa double or absent; (3) alar cells mostly
differentiated but nonvesiculate-inflated in a transverse row; (4) capsules in-
clined to horizontal or cernuous, occasionally erect; (5) exothecial cells noncol-
lenchymatous; (6) operculum conic-obtuse to rostrate, mostly not subulate; and
(7) exostome teeth usually with a zigzag center line, well developed, projecting
Robert R. Ireland, Department of Botany, Na- trabeculae, and rather broadly bordered. However, by their strict standards,
tional Museum of Natural History, Smithsonian they included the nonparaphylliate Hylocomiaceae in the Hypnaceae, as well
Institution, Washington, D.C.,USA, irelandr@ as Plagiothecium, Calliergonella, Pseudohygrohypnum, and Campylophyllum.
si.edu; William R. Buck, Institute of System- Few researchers in the last 20 plus years have followed this definition. In an
atic Botany, The New York Botanical Garden, initial look at the Hypnales using molecular data, Buck et al. (2000) found that
Bronx, New York, USA, bbuck@nybg.org. even different species of Hypnum seemed not to be closely related to one an-
Manuscript received 17 April 2008; accepted 9 other and that a number of genera, specifically Isopterygium and Platygyrium,
December 2008.
seem to be more closely related to a noncore group of Sematophyllaceae, includ-
ing Pylaisiadelpha and Taxithelium. This subsequently led Goffinet and Buck
(2004) to describe the Pylaisiadelphaceae for the genera Aptychella (Broth.)
Herzog, Brotherella M. Fleisch., Clastobryopsis M. Fleisch., Clastobryum Dozy
& Molk., Heterophyllium (Schimp.) Kindb., Isocladiella Dixon, Isopterygium
2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Mitt., Mastopoma Cardot, Platygyrium Schimp. in Bruch, wait to be revised until more time can be spent on solving
Schimp. & W. Gümbel, Pterogonidium (Hook.) Müll. the numerous species-level problems within each genus.
Hal. in Broth., Pseudotrismegistia H. Akiy. & Tsubota, This study was based on observation of approximately
Pylaisiadelpha Cardot, Taxitheliella Dixon, Taxithelium 1,900 herbarium specimens, in addition to 195 type speci-
Mitt., Trismegistia (Müll. Hal.) Müll. Hal. and Wijkia mens, belonging to 42 institutions, which are abbreviated
H. A. Crum. Although many of the genera more tradition- as follows for subsequent mention in the text:
ally aligned with the Plagiotheciaceae (e.g., Ireland, 1969)
have subsequently been considered better placed within AAU Herbarium Jutlandicum, Botanical Institute,
the Hypnaceae, some recent work (Pedersen and Hedenäs, University of Aarhus, Aarhus C., Denmark.
2001) has nevertheless continued to argue that genera ALTA Herbarium, Botany Department, University of
such as Pseudotaxiphyllum, Catagonium, Isopterygiopsis, Alberta, Edmonton, Alberta, Canada.
Bardunovia, Platydictya, Myurella, Struckia, Orthothe- B Herbarium, Botanischer Garten und Botanisches
cium, Rhizofabronia, and Herzogiella should be placed in Museum, Berlin-Dahlem, Berlin, Germany.
an expanded Plagiotheciaceae. Like most other workers, BA Herbario, Museo Argentino de Ciencias Naturales,
we have not been convinced of this expanded concept of Bernardino Rivadavia, Buenos Aires, Argentina.
the Plagiotheciaceae. The definition of the Hypnaceae has BIGU Herbario, Universidad de San Carlos de Guate-
become even more confused by the recent work of Ignatov mala, Guatemala City, Guatemala.
and Ignatova (2004), who restricted Hypnum to H. cu- BM Herbarium, Department of Botany, The Natural
pressiforme and its allies, moving the residue into Breidle- History Museum, London, UK.
ria and Stereodon, both in the Pylaisiaceae rather than the C Herbarium, Botanical Museum, University of
Hypnaceae. Within Latin America there are members of Copenhagen, Copenhagen K., Denmark.
the H. cupressiforme complex, but it is unclear to what CANM Herbarium, Bryology Section, Canadian Mu-
other genus (or genera) the other Latin American species seum of Nature, Ottawa, Ontario, Canada.
of Hypnum s.l. might be assigned. Ignatov and Ignatova COL Herbario Nacional Colombiano, Universidad
(2004) also placed Pseudohygrohypnum, Calliergonella, Nacional de Colombia, Bogotá, Colombia.
Callicladium, Ptilium, Homomallium, and Pylaisia in the COLO Herbarium, Museum, University of Colorado,
Pylaisiaceae. Platygyrium was enigmatically moved to the Boulder, Colorado, USA.
Entodontaceae, despite molecular evidence to the contrary CONC Herbario, Departamento de Botánica, Universi-
(Buck et al., 2000). The Ignatov and Ignatova (2004) treat- dad de Concepción, Concepción, Chile.
ment is entirely in Russian, and because of our language DUKE Herbarium, Botany Department, Duke Univer-
inadequacies, their motivation is not entirely clear and is sity, Durham, North Carolina, USA.
made even more confusing by their widely separating the F Herbarium, Botany Department, Field Museum
Hypnaceae from the Pylaisiaceae. Also impacting upon a of Natural History, Chicago, USA.
comprehensive family definition of the Hypnaceae is the FH Farlow Reference Library and Herbarium of
recent transfer by Hedenäs (2003) of Calliergonella, as Cryptogamic Botany, Cambridge, Massachu-
well as Campylophyllum (see Nishimura et al., 1984), into setts, USA.
the Amblystegiaceae. FLAS Herbarium, University of Florida, Gainesville,
We have used a more traditional definition of the Florida, USA.
Hypnaceae in this work, with minimal exception. We G Herbarium, Conservatoire et Jardin botaniques
readily admit that this broad definition of the family may de la Ville de Genève, Genève, Switzerland.
not withstand future molecular work. However, until such H Herbarium, Botanical Museum, University of
data result in a stable classification, this approach will at Helsinki, Helsinki, Finland.
least allow researchers to identify material at hand. H-BR Helsinki: Brotherus Herbarium (part of the Bo-
In this publication, 16 genera of Hypnaceae contain- tanical Museum, University of Helsinki, Finland).
ing 35 species which occur in Latin America are taxonomi- H-SOL Helsinki: Lindberg Herbarium (part of the Botan-
cally treated. The five remaining genera in the family, i.e., ical Museum, University of Helsinki, Finland).
Chryso-hypnum Hampe, Ectropothecium Mitt., Hypnum HIRO Botanical Institute, Hiroshima University, Hiro-
Hedw., Mittenothamnium Henn., and Vesicularia (Müll. shima, Japan.
Hal.) Müll. Hal., most of them containing a large number INPA Herbário, Departamento de Botânica, Instituto
of taxa, will require an enormous amount of time to resolve Nacional de Pesquisas da Amazônica, Manaus,
a number of taxonomic complexes. All five will have to Amazonas, Brazil.
 number 93 • 3

JBSD Herbarium, Jardín Botánico Nacional Dr. Ra- WTU Herbarium, Botany Department, University of
fael M. Moscoso, Santo Domingo, Dominican Washington, Seattle, Washington, USA.
Republic.
Other symbols and abbreviated terms used in this
JE Herbarium Haussknecht, Friedrich-Schiller-
publication are defined as follows:
Universität, Jena, Germany.
L National Herbarium Nederland, Leiden Univer- †
Dagger; when appearing after a name, it indi-
sity branch, Leiden, Netherlands. cates the individual is deceased.
LPB Herbario Nacional de Bolivia, La Paz, Bolivia. ± Plus or minus sign; indicates more or less as
M Herbarium, Botanische Staatssammlung, München, used in the descriptions.
Germany. ~ Approximately; used in “Specimens examined”
MEXU Herbario Nacional, Instituto de Biología, Uni- sections.
versidad Nacional Autónoma de México, Mex- BR National highway in Brazil; used in specimens
ico City, D.F., Mexico. examined.
MG Herbário, Departmento de Botânica, Belém, Cd. Spanish abbreviation for ciudad (“city” in En-
Pará, Brazil. glish); used in specimens examined.
MICH Herbarium, University of Michigan, Ann Arbor, Mpio. Spanish abbreviation for municipality; used in
Michigan, USA. specimens examined.
MO Herbarium, Bryology Section, Missouri Botani- s.l. Latin abbreviation for sensu lato (in the broad
cal Garden, Saint Louis, Missouri, USA. sense).
MSC Herbarium, Botany and Plant Pathology Depart- s.n. Latin abbreviation for sine numero (without a
ment, Michigan State University, East Lansing, number); used in specimens examined.
Michigan, USA. Syn. nov. Latin abbreviation for synonymum novum (new
NY William and Lynda Steere Herbarium, New York synonym); used in specimens examined.
Botanical Garden, Bronx, New York, USA. YPFB Abbreviation for a truck compound in Bolivia;
PC Herbier, Laboratoire de Cryptogamie, Muséum used in specimens examined.
National d’Histoire Naturelle, Paris, France.
Definitions of other italicized Latin abbreviations ap-
PH Herbarium, Botany Department, Academy of
pearing in the family, genus, and species type citations
Natural Sciences, Philidelphia, Pennsylvania, USA.
may be found in van der Wijk et al. (1959).
PMA Herbarium, Departmento de Botánica, Universi-
Nearly all information presented in “Specimens ex-
dad de Panamá, Panamá, Panama.
amined” sections is reproduced exactly as it appeared on
S Herbarium, Botany Departments, Swedish Mu-
actual specimen labels. Minor editorial changes included
seum of Natural History, Stockholm, Sweden.
substitution of “~” for “ca.” and making capitalization and
SP Herbário, Instituto de Botânica, São Paulo, Brazil.
abbreviations consistent. However, Spanish, Portuguese,
TENN Herbarium, Botany Department, University of
German, and other foreign language phrases appearing on
Tennessee, Knoxville, Tennessee, USA.
specimen labels were left untranslated. In addition, conver-
TNS Herbarium, Botany Department, National Sci-
sion into metric of distances given in English units in these
ence Museum, Tokyo, Japan.
sections and in type descriptions is left to the reader.
U Herbarium, Institute of Systematic Botany, State
University of Utrecht, Utrecht, Netherlands.
U at L Utrecht Bryophyte Herbarium; all bryophyte
specimens are now at Leiden (see L). SYSTEMATIC TREATMENT
UBC Herbarium, Botany Department, University of
British Columbia, Vancouver, British Columbia, Family HYPNACEAE Schimp.
Canada.
UPS Botanical Museum (Fytoteket), Uppsala Univer- Family HYPNACEAE Schimp., Coroll. Bryol. Eur. 113. 1855
sity, Uppsala, Sweden. [1856].—Orthotheciaceae Schimp., Coroll. Bryol. Eur.
US United States National Herbarium, Department 103. 1855 [1856].—Pylaisiaceae Schimp., Syn. Musc.
of Botany, Smithsonian Institution, Washington, Eur. 241. 1860, “Pylaisieae”; Hypnaceae subfam. Py-
D.C., USA. laisioideae (Schimp.) M. Fleisch., Musci Buitenzorg 4:
USJ Herbario, Escuela de Biología, Universidad de 1379. 1923, “Pylaisieae.”—Microthamniaceae Kindb.,
Costa Rica, San José, Costa Rica. Bot. Centralbl. 77: 386. 1899, nom. illeg.
4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Plants small to robust, in lax or dense mats, often lus- sometimes present, mostly cylindrical to fusiform, uni-
trous, green to golden, sometimes reddish. Stems mostly seriate, and clustered in leaf axils, or sometimes multi-
creeping, sometimes ascending or rarely erect or pendent, cellular, twisted-vermiform, with acute teeth at apices or
irregularly branched to regularly pinnate, sometimes com- bearing reduced leaves from apex to base of propagula.
planate-foliate, rarely julaceous; rhizoids smooth or pap- Autoicous or dioicous. Perichaetial leaves differentiated.
illose, in leaf axils or abaxial to leaf insertions; stems in Setae elongate, mostly smooth, usually reddish; capsules
cross section sometimes with a unistratose hyalodermis, inclined to horizontal, sometimes erect, mostly asymmet-
subtended by small, firm- or thick-walled cells surround- ric, arcuate or straight, ovoid to long-cylindrical, usually
ing larger, thinner-walled cells, central strand present or smooth, often constricted below the mouth when dry and
absent; paraphyllia usually none, rarely present; pseu- empty; exothecial cells quadrate to rectangular, thin- or
doparaphyllia filamentous or foliose; axillary hairs with thick-walled, mostly not collenchymatous; annulus usu-
1–2(–4) short brown basal cells and 1 to several elon- ally differentiated; operculum conic to rostrate; peristome
gate, hyaline distal cells. Stem and branch leaves mostly typically double, the endostome rarely lacking, usually at-
similar, sometimes differentiated, straight to homomal- tached at the mouth, exostome teeth typically shouldered
lous to falcate-secund, occasionally complanate, linear and bordered, on the front surface cross-striolate below,
to broadly oblong-ovate, but typically lanceolate-ovate, coarsely papillose above, sometimes reduced and papil-
often asymmetric, obtuse to acuminate, often concave, lose throughout or ± smooth, typically strongly trabecu-
sometimes plicate, sometimes decurrent; margins not or late at back; endostome usually free, sometimes fused to
very rarely bordered, entire to serrulate, rarely serrate, the exostome, mostly with a high, or rarely low, basal
plane, recurved or incurved; costa short and double or membrane, endostome broad or rarely narrow, keeled,
absent; cells mostly linear, occasionally shorter, usu- usually perforate, usually as long as the exostome, cilia
ally smooth, rarely prorulose or papillose, mostly thin- mostly in groups of 1–3 or rarely absent. Spores spherical
walled; alar cells usually differentiated, often quadrate, to oval or ovoid, smooth or often papillose. Calyptrae
sometimes enlarged and inflated. Asexual propagula cucullate, usually naked, rarely hairy, usually smooth.

KEY TO THE GENERA

(Only the numbered genera are treated in this publication.)
1. Stems with a hyalodermis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Pseudoparaphyllia present, foliose to broadly filamentous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hypnum p.p.
2. Pseudoparaphyllia lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Leaves entire to minutely serrulate; asexual propagula often present . . . . . . . . . . . . . . . . . . . . . . . . 6. Isopterygiopsis
3. Leaves serrulate to serrate; asexual propagula absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. Herzogiella
1. Stems lacking a hyalodermis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Leaf cells smooth or inconspicuously and irregularly prorulose; branch and stem leaves scarcely differentiated . . . . 5
5. Leaf areolation very lax; pseudoparaphyllia filamentous, cells thin-walled and bulging . . . . . . . . . . Vesicularia
5. Leaf cells narrowly prosenchymatous; pseudoparaphyllia filamentous or foliose, cells firm-walled . . . . . . . . . . 6
6. Leaves dimorphic (lateral and ventral leaves differentiated) . . . . . . . . . . . . . . . . . . . . . . . . 14. Rhacopilopsis
6. Lateral and ventral leaves similar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Leaves falcate-secund . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Capsules erect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12. Pylaisia
8. Capsules inclined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9. Alar cells well developed; pseudoparaphyllia foliose to broadly filamentous . . . . . . . . . . Hypnum p.p.
9. Alar cells scarcely differentiated; pseudoparaphyllia, especially outer ones, filamentous or narrowly
foliose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ectropothecium p.p.
7. Leaves not falcate-secund . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10. Pseudoparaphyllia foliose; alar cells moderate in number . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11. Plants not complanate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12. Capsules erect; branches often terminating in tuft of axillary brood branches . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. Platygyrium
12. Capsules inclined; branches not terminating in tuft of axillary brood branches . . . . . . . 13
13. Costa ¼–½ the leaf length; laminal cells sparsely prorulose . . . . . . . 3. Elmeriobryum
 number 93 • 5

13. Costa ¼ or less the leaf length; laminal cells smooth . . . . . . . . . . . . . . . . . . . . . . . . 14

14. Upper laminal cells ~3:1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. Homomallium
14. Upper laminal cells more than 5:1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15. Branches soft, prostrate, often with hooked apices . . . . . . . . . . . Hypnum p.p.
15. Branches stiff, erect, with straight apices . . . . . . . . . . . . . . . . . . . . 5. Irelandia
11. Plants complanate, often strongly so . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16. Taxiphyllum
10. Pseudoparaphyllia filamentous or absent; alar cells few . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16. Plants mostly without propagula or with small, ovoid ones; pseudoparaphyl-
lia mostly filamentous, rarely absent; plants primarily occurring at low to
moderate elevations, often weedy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17. Plants not complanate; leaves with a single, enlarged cell in the basal
angles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18. Capsules small, ovoid, inclined; annulus differentiated . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ectropothecium p.p.
18. Capsules cylindrical, erect; annulus not differentiated . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. Platygyriella
17. Plants ± complanate; leaves with weakly differentiated alar cells but not
with a single enlarged cell in angles . . . . . . . . . . . . . . . . . . . . . . . . . 19
19. Rhizoids smooth; exostome better developed than endostome
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. Isopterygium
19. Rhizoids smooth to slightly roughened; endostome better devel-
oped than exostome . . . . . . . . . . . . . . . . . 15. Syringothecium
16. Plants with abundant elongate propagula in leaf axils; pseudoparaphyllia
none; plants rare, at low to high elevations . . . . . . 13. Pseudotaxiphyllum
4. Leaf cells conspicuously prorulose; branch and stem leaves ± differentiated . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20. Stems bearing paraphyllia . . . . . . . . . . . . 11. Puiggariopsis
20. Stems without paraphyllia . . . . . . . . . . . . . . . . . . . . . . 21
21. Stem leaves decurrent, with well-developed alar regions
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Ctenidium
21. Stem leaves not decurrent, with poorly developed alar
regions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22. Plants stipitate, with arched stems and slender,
tapering branches; operculum conic-rostrate . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . Mittenothamnium
22. Plants not stipitate, prostrate, with ± blunt branches;
operculum conic to apiculate . . . . . . . . . . . . . . 23
23. Leaves obtuse; laminal cells prorulose and
papillose . . . . . . . . . . . . . . . . . . . 8. Phyllodon
23. Leaves acuminate; laminal cells only prorulose
. . . . . . . . . . . . . . . . . . . . . . . . Chryso-hypnum

1. Ctenidium (Schimp.) Mitt. yellow-brown. Stems creeping to ascending, irregularly

to regularly pinnately branched, the branches sometimes
Ctenidium (Schimp.) Mitt., J. Linn. Soc., Bot. 12: 509. 1869. branched, unequal, not complanate-foliate; stems in cross
section without a hyalodermis, with small thick-walled
Hypnum subg. Ctenidium Schimp., Syn. Musc. Eur. 631. 1860. Stereodon cells surrounding larger thinner-walled cells, central strand
sect. Ctenidium (Schimp.) Mitt., J. Linn. Soc., Bot. 8: 153. 1864. Type: present; rhizoids finely papillose, reddish brown; paraphyl-
Ctenidium molluscum (Hedw.) Mitt. lia none; pseudoparaphyllia foliose, large; axillary hairs
consisting of 1–4 short, hyaline basal cells and 1 to several
Plants small to medium-sized, in often extensive, mostly elongate hyaline distal cells. Asexual reproductive
dense mats, often lustrous, soft, green to yellow-green to bodies none. Stem and branch leaves differentiated, stem
6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

leaves erect-spreading to squarrose-recurved, often se- Ctenidium malacodes

cund, broadly ovate, abruptly acuminate, often plicate, es-
pecially when dry, cordate and broadly decurrent at base; Ctenidium malacodes Mitt., J. Linn. Soc., Bot. 12: 509.
margins serrate to serrulate throughout, plane to erect; 1869. Type: Colombia. Andes Bogotensis, in sylvis su-
costa short and double or none; cells linear to oblong- pra Pacho ad arbores, 6000 ped., J. Weir 243 (lecto-
linear, low-prorulose at upper and sometimes lower ends type: NY!, designated by Nishimura 1985); Ecuador.
on abaxial leaf surface, sometimes smooth, firm-walled, Guarapata, Spruce 1053 (syntype: NY!).
becoming shorter in a few rows across the insertion; alar
cells decurrent, subquadrate to oblong, enlarged. Branch Cupressina anacamptopteris Müll. Hal., Bull. Herb. Boiss. 6: 123. 1898.
leaves smaller than stem leaves, erect-spreading to squar- Ctenidium anacamptopteris (Müll. Hal.) Broth. in Engl. & Prantl, Nat.
rose, obscurely homomallous to falcate-secund, lanceolate Pflanzenfam. 1(3): 1048. 1908. Type: Brazil. Serra Itatiaia, 2100 m, E. Ule
to ovate-lanceolate, gradually acuminate, concave, subpli- 1882 (isotypes: H, US!).
cate, short-decurrent; margins serrate to serrulate above,
subentire to serrulate below, plane to recurved; costa short Meteorium terrestre Müll. Hal., Hedwigia 40: 97. 1901. Ctenidium terrestre
and double or none; cells linear to oblong-linear, conspic- (Müll. Hal.) Broth. in Engl. & Prantl, Nat. Pflanzenfam. 1(3): 1048. 1908.
uously prorulose at upper and sometimes lower ends on Type: Brazil. Santa Catharina: Serra Geral, E. Ule 1206 (isotype: H-BR).
abaxial leaf surface, firm-walled, becoming shorter in a
few rows across the insertion; alar cells subquadrate to Ctenidium muenchii Broth. ex H. A. Crum & Sharp, Bol. Soc. Bot. México
short-rectangular. Dioicous or phyllodioicous. Perichaetial 15: 15. 1953, nom. nud. in syn. Based on: Mexico. Chiapas: distr. San Cris-
leaves erect with spreading apices, oblong to broadly ob- tobal, “los Llanos,” 2500 m, 23 Dec 1907, Münch 7433 (H, NY!).
long, abruptly long-acuminate, concave; margins serrulate
above, serrulate to entire below, plane; costa short and Plants medium-sized, often in extensive mats, lustrous,
double or none; cells oblong-linear to linear-flexuose, usu- soft, yellow-green to golden. Stems to 6 cm long, creep-
ally smooth, becoming laxly rectangular toward the inser- ing to ascending, subpinnately to pinnately branched, the
tion; alar cells not differentiated. Setae elongate, smooth branches often unequal, not complanate-foliate; stems in
or rarely roughened, reddish; capsules inclined to hori- cross section with 3–5 rows of small thick-walled cells,
zontal, arcuate, asymmetric, ovoid to broadly cylindrical; the outermost row sometimes with ± thin outer walls,
exothecial cells ± isodiametric, thick-walled on vertical surrounding larger thinner-walled cells, central strand of
walls, thinner-walled on cross walls; annulus differenti- small thin-walled cells; pseudoparaphyllia foliose, large,
ated; operculum conic, mostly short-rostrate; peristome serrulate; axillary hairs consisting of a single short hyaline
double, exostome teeth shouldered, bordered, on the front basal cell, 1–2 long-rectangular hyaline intercalary cells
surface cross-striolate below, sometimes with overlying and a single elongate hyaline distal cell. Stem and branch
papillae, papillose above, trabeculate at back; endostome leaves differentiated, stem leaves 0.8–1.1 mm long, erect-
with a high basal membrane, segments keeled, perforate, to wide-spreading with flexuose apices, broadly ovate-
nearly as long as the exostome, cilia in groups of 2–3, well triangular, abruptly acuminate, the acumen flexuose,
developed to rudimentary. Spores spherical, finely papil- sometimes twisted, somewhat concave, usually somewhat
lose. Calyptrae cucullate, usually hairy, smooth. plicate, broadly decurrent; margins serrate to sharply serru-
Distribution: Most of the 21 species of Ctenidium late almost to base, plane or occasionally erect; costa short
occur in eastern Asia although the type species, C. mollus- and double, the two forks not united at base; cells linear to
cum, is circum-North Temperate. Only a single species oc- oblong-linear, sparsely prorulose at upper ends on abaxial
curs in Latin America. surface, firm-walled, becoming oblong to rectangular in
Discussion: Ctenidium has differentiated stem and 1–3 rows across the insertion; alar cells extending into the
branch leaves and, in many ways, is like Mittenothamnium decurrencies, subquadrate to short-rectangular in a sizable
but is distinguished by decurrent leaves, especially those of group, surrounding 4–8 larger, oblong, ± inflated cells
the stem, with better alar cell development. Ctenidium is often in a somewhat excavated area. Branch leaves (0.5–)
also characterized by serrate leaves with prorulose cells and 0.7–1.0(–1.1) mm long, wide-spreading, sometimes ± se-
stem leaves that are broadly ovate and abruptly tapered to cund, lanceolate to ovate-lanceolate, gradually acuminate,
an acuminate apex. A relationship to Mittenothamnium is the apex curved to subflexuose and occasionally twisted,
unmistakable not only because of gametophytic characters concave, subplicate, short-decurrent; margins serrate to
but also because of its unequally thickened exothecial cells. serrulate above, subentire to serrulate below, plane above,
 number 93 • 7

usually erect below; costa short and double, or often none; Selected specimens examined: MEXICO. Chiapas:
cells linear to oblong-linear, sparsely to densely prorulose San Cristoval, 2500 m, Münch 7433 (H, NY). México:
at upper and sometimes lower ends at back, firm-walled, Mpio. Ocuilán, Barranca de Mexicapa, 2200–2350 m,
becoming rectangular in 1–2 rows across the insertion; alar Buck 28148 (NY). Michoacán: 4.1 km W of Mil Cum-
cells few, only scarcely extending above the short decur- bres, ~2800 m, McGregor 16574 (NY). Oaxaca: 34.5
rencies, subquadrate to oblong. Asexual propagula none. km above Valle Nacional in Sierra Juárez on Hwy. 175,
Dioicous. Perichaetial leaves 2.0–2.5 mm long, erect, ob- Sharp et al. 4750-b (TENN). Veracruz: Paß an der Straße
long, abruptly slenderly long-acuminate, concave, subpli- Orizaba–Tehuacan, 23 Mar 1979, Frahm s.n. (NY). GUA-
cate; margins serrate in the acumen, coarsely serrate at the TEMALA. Huehuetenango: Sierra de los Cuchumatanes,
shoulder, serrulate to entire below, plane or rarely erect; 2400 m, Steyermark 50030 (NY). San Marcos: Barranco
costa none; cells oblong-linear, smooth, becoming laxly Eminencia, 2500–2700 m, Standley 86365 (NY). Sololá:
rectangular toward the insertion; alar cells not differenti- Volcán Santa Clara, 2100–3000 m, Steyermark 46954
ated. Setae 1.6–2.3 cm long, elongate, smooth, reddish; (NY). Totonicapán: Above Totonicapán, 2800–3100 m,
capsules 1.5–2.0 mm long, inclined to horizontal, arcuate, Standley 84472 (NY). COSTA RICA. Alajuela: La Palma
asymmetric, ovoid to cylindrical; exothecial cells subquad- de San Ramón, Brenes 113 (NY). PANAMA. Chiriquí: Bo-
rate, the vertical walls thick, the cross walls thin, often quete, Helion 812 (H, NY). JAMAICA. Blue Mountains,
± sigmoid; annulus broad, of 2–3 rows of elongate firm- St. Helen’s to Morce’s Gap, Britton 1075 (NY). DOMIN-
walled cells, deciduous; operculum short conic-rostrate; ICAN REPUBLIC. La Estrelleta: Sierra de Neiba, ~1700
exostome teeth shouldered, bordered, on the front surface m, Buck 4790 (NY). La Vega: Vicinity of La Lagunita,
cross-striolate below, with fine, overlying papillae, the ~2600 m, Norris et al. B5738 (H). Peravia: San José de
papillae becoming denser at midtooth and coarser above, Ocoa, La Nevera, ~2100 m, Mejía & Zanoni 8721 (JBSD,
trabeculate at back; endostome with a high, finely pap- NY). COLOMBIA. Boyacá: Mpio. Duitama, Páramo “La
illose basal membrane, segments finely papillose, keeled, Rusia,” 3550 m, 06°N, 73°W, Ireland 23626 (CANM,
perforate, sometimes gaping with age, about as long as the US). Caldas: Cordillera Central, Magana, 3000–3300 m,
teeth, cilia finely papillose, in groups of 2–3, short to ru- Killip & Hazen 12098 (NY); Mpio. Río Sucio, on the
dimentary. Spores 13–17 µm in diameter, spherical, finely Río Sucio, 1900–2080 m, ~05°20'N, 75°15'S, Churchill
papillose. Calyptrae cucullate, appearing naked or with a & Arbeláez A. 15658 (US). Nariño: Mpio. Pasto, be-
few scattered hairs, smooth. tween km 13 and 15, Cocha Lagoon, 3100 m, 01°12'N,
Distribution and ecology: Mexico to northern 77°12'W, Churchill & Arbeláez A. 15960-b (US). Norte
South America; controversially in eastern North America; de Santander: Cordillera Oriental, 2800–3000 m, Kil-
growing typically on rocks but also soil and logs, in humid lip & Smith 19868 (NY). VENEZUELA. Trujillo: Distr.
and often open forests, above 1500 m. Urdaneta, S and above La Mesa de Esnujaque, 3100 m,
Illustrations in publications: Fig. 180G–J in Bar- Griffin III et al. 1377 (FLAS, NY, TNS). GUYANA. Mt.
tram (1949: 408); Pl. 130 in Buck (1998: 330); Fig. 776 in Roraima, Quelch & McConnell 340 (H). ECUADOR.
Nishimura and Ando (1994: 1047). Andes Quitenses, ~1800 m, Spruce 1051 (H-SOL); Tim-
Discussion: Ctenidium malacodes is distinguished giragia. 2700−3050 m, Spruce s.n. (NY). PERU. Conven-
by its broadly ovate-triangular stem leaves that have large ción: Dept. Cuzco, 3200 m, Bües 1468 (NY). BOLIVIA.
decurrencies of differentiated alar cells. The stem leaves are Santa Cruz: Vallegrande, Calaculo, ~26 km S of Valle-
also abruptly acuminate and sharply serrate. The branch grande, road to Khasamonte, 2450 m, 18°38'S, 64°02'W,
leaves are only shortly decurrent and gradually acuminate Churchill & Arroyo P. 21200 (US); Tablas, ~3400 m,
but usually more serrate than the stem leaves. The cells are Herzog 2817 (JE); Río Tocorani, ~2200 m, Herzog
prorulose on the abaxial surface of the leaves. 4082 (H, JE). BRAZIL. Santa Catarina: Serra Geral, Ule
Ctenidium malacodes has been reported by Nishimura 1157 (H).
(1985) as occurring in eastern North America, where collec-
tions of the genus had previously been called C. molluscum 2. Elmeriobryum Broth.
(Hedw.) Mitt. Pedano (1996) suggested that this material is
neither C. molluscum nor C. malacodes but rather a taxon Elmeriobryum Broth., Leafl. Philipp. Bot. 6: 1974. 1913,
intermediate between the two and worthy of species rank. nom. inval. sin. descr. gen., in Engl., Nat. Pflanzenfam.
Buck and Allen (2004) made the appropriate combination ed. 2, 11: 204. 1925. Type: Elmeriobryum philippinense
as C. subrectifolium (Brid.) W. R. Buck & B. H. Allen. Broth.
8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Plants somewhat shiny, golden brown to yellow- Pie de la Cuesta, in upper part of the barranco between
green, in relatively robust, extensive mats. Stems creep- Finca La Lucha and Buena Vista, 2500–2700 m, 6 Feb
ing, irregularly branched, procumbent to ascending; stems 1941, Standley 86466 (holotype: MICH; isotypes: F,
in cross section without a hyalodermis, with small thick- FH, NY!).
walled cells surrounding larger thin-walled cells, central
strand present; pseudoparaphyllia narrowly foliose; ax- Plants relatively robust, in extensive loose mats, shiny,
illary hairs consisting of 1(–2) short brown basal cells golden brown to yellow-green. Stems to ~10 cm, creeping,
and 2–3 elongate hyaline distal cells. Stem and branch elongate, irregularly and distantly 1–2-pinnately branched;
leaves somewhat differentiated, imbricate, erect- to wide- stems in cross section with 2–3 rows of small thick-walled
spreading, ovate to oblong, apiculate to short-acuminate, cells surrounding abruptly larger thin-walled cells, central
strongly concave, somewhat plicate; margins mostly ser- strand of small very thin-walled cells; pseudoparaphyllia
rulate above, entire below, plane to incurved; cells linear, foliose; axillary hairs consisting of 1 short brown basal
smooth or minutely prorulose, thin- to firm-walled; alar cell and 2 elongate hyaline distal cells. Stem and branch
cells not or somewhat differentiated. Asexual propagula leaves somewhat differentiated; stem leaves 1.0–1.8 mm
not seen. Dioicous. Perichaetia large, leaves oblong, long- long, 0.7–1.4 mm wide, erect to erect-spreading when dry,
acuminate, plicate; margins subentire, plane; costa usu- spreading when moist, broadly ovate to oblong, abruptly
ally none; cells linear-flexuose, smooth, becoming laxly apiculate, concave, plicate; margins serrulate above, en-
rectangular toward the insertion; alar cells not differenti- tire below, reflexed at base, plane or broadly inrolled
ated. Setae elongate, smooth, reddish; capsules inclined, above; costa double, the two forks separated at base and
weakly arcuate, ± asymmetric, oblong; exothecial cells ± parallel, extending ¼–½ the leaf length; median cells
subquadrate to irregularly short-rectangular, firm-walled, 25–75 × 4–7 µm, narrowly elliptic to linear, smooth or
not collenchymatous; annulus well- differentiated; opercu- sometimes minutely prorulose, basal cells wider, porose;
lum apiculate; peristome double, exostome teeth yellow- alar cells scarcely differentiated, slightly wider than other
brown, shouldered, bordered, on the front surface with basal cells. Branch leaves 1.0–1.5 mm long, 0.4–0.7 mm
a zigzag median line, not furrowed, cross-striolate below, wide, ovate to elliptic, less plicate than stem leaves. Asex-
coarsely papillose above, trabeculate at back; endostome ual propagula none. Apparently dioicous. Perichaetia un-
with a high basal membrane, segments keeled, narrow, known. Sporophytes unknown.
perforate, about as long as the teeth, cilia in groups of Distribution and ecology: Known only in Central
1–3, nodose. Spores spherical, papillose. Calyptrae small- America from Guatemala, El Salvador, and Costa Rica
cucullate, naked, smooth. (Figure 2). Usually on tree trunks and branches or some-
Discussion: A taxonomic study of the genus Elmer- times on leaves. Also occurring on wet banks and rocks in
iobryum is currently in progress by the junior author. It was moist forests. At moderate to high elevations from 1800 to
recently reviewed by Buck and Tan (2008). Thus far, the 3000 m.
genus is known to have one species in Central America, one Discussion: This little-known species was formerly
in eastern Asia (Philippines and Taiwan), and one in Papua confused with Leptohymenium (Hylocomiaceae), but the
New Guinea. It is sometimes combined with Gollania, but distinction between it and Leptohymenium was resolved by
according to Rohrer (1985), Elmeriobryum differs from Rohrer (1986). Unlike Leptohymenium and the rest of the
that genus by its heteromallous, broadly ovate leaves with Hylocomiaceae, the branching in Elmeriobryum is mono-
acute to apiculate apices and porose basal cells. Gollania podial rather than sympodial, the leaves are not at all de-
is characterized by mostly homomallous, ovate to ovate- current, and the alar cells scarcely if at all differentiated.
lanceolate, acuminate leaves with nonporose to subporose This New World species differs from its Asian counterpart,
basal cells. E. philippinense Broth., by having leaves more strongly pli-
cate, with an apiculate rather than an acute apex, and a
Elmeriobryum guatemalense stronger costa. Also, in E. guatemalense the alar cells are
more poorly differentiated than in E. philippinense.
FIGURE 1 Selected specimens examined: GUATEMALA.
Guatemala: Volcán de Pacaya, Standley 80724 (F, FH,
Elmeriobryum guatemalense J. R. Rohrer, Bryologist 89: MICH, US). Quezaltenango: El Pocito, S of San Martín
29. 1986. Type: Guatemala. San Marcos: Barranco Chile Verde, Standley 85043 (F, MICH, NY). San Mar-
Eminencia, road between San Marcos and San Rafael cos: Barranco Eminencia, Standley 68524 (F, FH). Sololá:
 number 93 • 9

FIGURE 1. Elmeriobryum guatemalense. A. Habit. B. Enlargement of portion of branch. C. Stem leaves. D. Branch leaves. E. Apical leaf cells.
F. Median-marginal stem leaf cells. G. Alar cells of stem leaf. H. Pseudoparaphyllia. I. Portion of stem cross section. All illustrations drawn from
Standley 80724 (US).

Volcán Atitlán, Steyermark 47464 (F, FH, MICH, NY, 3. Herzogiella Broth.
TENN). EL SALVADOR. Santa Ana: Cerro Miramundo
NW of Metapán, Carlson 962b (F, FH). COSTA RICA. Herzogiella Broth. in Engl., Nat. Pflanzenfam. ed. 2,
Heredia: Volcán Barba, Griffin et al. D-268 (CANM, F, 11: 466. 1925. Type: Herzogiella boliviana (Broth.)
MO, NY). M. Fleisch. in Broth.
10 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 2. Distribution of Elmeriobryum guatemalense in Latin America.

Dolichotheca Lindb., Not. Sällsk. Fauna Fl. Fenn. Förh. 13: 417. 1874. illeg. metric, nondecurrent or 1–2 cells decurrent, ovate or lan-
homonym (non Dolichotheca Cass., Dict. Sci. Nat. 51: 476. 1827. =Coreop- ceolate, acuminate; margins serrulate to strongly serrate
sis in Asteraceae). above the middle, serrate to nearly entire below, plane;
costa short and double, rarely lacking; median cells thick-
Isopterygium Mitt. subgen. Dolichotheca (Lindb.) Lindb., Musci Scand. 39. walled, linear-fusiform, smooth, the wall of cells at leaf
1879. base usually pitted, sometimes pits lacking; alar cells usu-
ally clearly differentiated, a few quadrate, rectangular, or
Sharpiella Z. Iwats., J. Hattori Bot. Lab. 28: 202. 1965 (new name for Doli- abruptly inflated and rounded cells present with 1–4 cells
chotheca Lindb.). Type: Plagiothecium repens Lindb. on margins. Autoicous; perigonia and perichaetia near
base of stems, leaves ovate-lanceolate, gradually acumi-
Plants medium-sized, in thin to dense mats, glossy, nate; margins plane. Setae smooth, elongate, straight to
light green to yellow-green. Stems creeping to ascend- curved, often twisted, yellow, brown or red; capsules erect
ing, simple or sparingly and irregularly branched; stems to cernuous, straight or slightly curved, yellow, brown or
in cross section with a hyalodermis, surrounding a few reddish brown, oblong or cylindrical, when dry smooth or
rows of smaller thick-walled cortical cells and large striate, tapering to a sometimes wrinkled neck, often con-
thin-walled cells in center, central strand usually absent; tracted below mouth; operculum conic to short-rostrate;
rhizoids sparse, papillose, in leaf axils of single Latin annulus of 2 or 3 rows of large cells, deciduous; peristome
American species or in leaf axils or abaxial to leaf inser- cross-striolate below, papillose above, bordered, tra-
tions in species from elsewhere, often restricted to base of beculate at back; endostome papillose, consisting of nar-
stems; pseudoparaphyllia lacking; axillary hairs consisting row keeled segments from a high basal membrane, cilia
of 1 short brown basal cell and 2–3 elongate hyaline dis- in groups of 1–3, approximately the length of segments,
tal cells. Asexual reproductive bodies lacking. Stem and sometimes lacking. Spores spherical to ovoid, smooth or
branch leaves similar, rigid, crowded and imbricate to minutely papillose. Calyptra cucullate, smooth, naked.
somewhat remote, spreading to squarrose, sometimes tips Distribution: The eight species of Herzogiella
secund at stem and branch apices, concave, smooth, sym- known in the world occur in terrestrial habitats at low
 number 93 • 11

to high elevations predominately in temperate and boreal Plagiothecium bolivianum Broth. in Herzog, Biblioth. Bot. 87: 153. 1916;
regions. Herzogiella boliviana (Broth.) M. Fleisch. in Broth. in Engl., Nat. Pflanzen-
Discussion: The genus Herzogiella is represented fam. ed. 2, 11: 466. 1925. Type: Bolivia. Waldgrenze über Tablas, Herzog
by five species in North America, with only one, H. cylindri- 2821 (lectotype: H!, here designated); zwischen San Mateo u. Sunchal, Her-
carpa (Cardot) Z. Iwats., present in Latin America. Many zog 4435 (syntype: NY!).
of the taxa presently in Herzogiella, particularly the North
American members, have had a checkered history of name Plants in thin, loose mats, glossy, yellowish green.
changes. Dolichotheca Lindb. was once used as the generic Stems 0.6–1.5 cm long, 1–2 mm wide, ascending, simple
name for many of the eight taxa but it was soon discovered to irregularly branched; rhizoids papillose, in leaf axils.
to be a later homonym of Dolichotheca Cass. (= Coreop- Leaves 1.0–1.5 mm long, 0.4–0.6 mm wide, somewhat
sis) in the Asteraceae. The species were then transferred to rigid, close to distant, loosely complanate-foliate, erect-
Sharpiella Z. Iwats., described in honor of A. J. Sharp, but to wide-spreading, sometimes distinctly secund, smooth
Iwatsuki (1970) later discovered that it was synonymous or weakly plicate, lanceolate, slightly concave, slenderly
with Herzogiella. acuminate; margins sharply serrate above, serrulate below
Herzogiella is characterized by autoicous plants, stems to base, plane; costa short and double or none; median
with a hyalodermis, papillose rhizoids in leaf axils of cells 47–70 × 5–6 µm, smooth, linear; alar regions with a
H. cylindricarpa, pseudoparaphyllia lacking, leaves non- few quadrate, rectangular or rounded cells, often 1–4 cells
decurrent or 1–2 cells decurrent, leaf margins serrate to on margins. Autoicous. Setae 1.0–1.8 cm long, reddish
serrulate throughout, capsules with conic to short-rostrate brown, sometimes yellowish above; capsules 1.5–3.0 mm
opercula, and urns usually striate when dry (except in long, 0.3–0.4 mm wide, yellowish, erect, sometimes
H. cylindricarpa). slightly curved, 1.5–3.0 mm long, 0.3–0.4 mm wide, nar-
A synopsis of the genus Herzogiella and its five North rowly cylindrical, usually contracted below mouth when
American species, including H. cylindricarpa, has recently dry, narrow at the sometimes wrinkled neck; operculum
been published (Ireland, 1991b) showing the distribution conic to obliquely short-rostrate; peristome teeth up to
of the species on the continent. The spores of Herzogi- 0.2 mm long, cilia none. Spores 10–12 µm in diameter,
ella cylindricarpa and six other species in the genus were minutely papillose.
recently studied under the scanning electron microscope Distribution and ecology: Infrequent and thus far
(Ireland, 1990). All seven species had the gemmate type known from Mexico, Guatemala, Costa Rica, and a few lo-
of spore ornamentation. However, two of the species, calities in Colombia, Venezuela, Bolivia, Peru, and Ecuador
H. cylindricarpa and H. striatella (Brid.) Z. Iwats., more (Figure 4); also known from Rwanda (Africa). Usually oc-
frequently had the verrucate type of ornamentation, in- curring on humus on moist, rotting logs, bases of trees and
dicating a closer relationship to each other than to the tree trunks, mainly coniferous, and rarely on rock. Predom-
other five species studied, namely, H. adscendens (Lindb.) inately at high elevations, mostly over 2500 m and known
Z. Iwats. & W. B. Schofield, H. perrobusta (Cardot) up to ~4100 m; recorded as low as 1830 m.
Z. Iwats., H. renitens (Mitt.) Z. Iwats., H. seligeri (Brid.) Discussion: Recognized by the erect- to wide-
Z. Iwats., and H. turfacea (Lindb.) Z. Iwats. The eighth spreading, lanceolate, slightly concave, slenderly acuminate
species in the genus, H. letestui (Dixon & P. de la Varde) leaves with plane margins that are serrate above and ser-
Ando, was not studied. rulate below to the base. The stems have a prominent hya-
lodermis which is evident in cross section. The plants are
Herzogiella cylindricarpa autoicous and the sporophytes are distinctive because of the
very slender, long, erect capsules, narrowed at the neck and
FIGURE 3 contracted below the mouth when dry.
Herzogiella cylindricarpa was originally described as
Herzogiella cylindricarpa (Cardot) Z. Iwats., J. Hattori an Isopterygium by Cardot (1910), but Bartram (1946)
Bot. Lab. 33: 374. 1970. transferred it to Ctenidiadelphus M. Fleisch., a genus then
known to have species only in Java and Borneo. Iwatsuki
Isopterygium cylindricarpum Cardot, Rev. Bryol. 37: 56. 1910; Ctenidiadel- (1970) eventually transferred C. cylindricarpa to Herzogi-
phus cylindricarpus (Cardot) E. B. Bartram, Bryologist 49: 124. 1946. Type: ella when he revised that genus.
Mexico. Amecameca, Pringle, Plantae Mexicanae 10606 (lectotype: MO!, The lectotype (Herzog 2821) of Plagiothecium bolivi-
here designated; isolectotypes: F!, FH!, H!, MICH!, TENN!, US!). anum Broth. in Herzog was designated from a collection
12 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 3. Herzogiella cylindricarpa. A. Habit. B. Enlargement of portion of stem. C. Leaves. D. Apical leaf cells. E. Median-marginal leaf cells.
F. Cells of alar region. G. Portion of stem cross section. H. Capsules (dry). Illustrations A–F drawn from Sharp 2287 (MEXU); G drawn from
Cárdenas 4276 (MEXU); H, one on left from Sharp 2287 (MEXU), two on right from Sharp 2160 (TENN).

in the Brotherus herbarium in Helsinki. The species is (1916: tbl. 7). Another specimen mentioned by Brotherus
here considered synonymous with Herzogiella cylindri- in the protologue of P. bolivianum (syntype: Herzog 4435
carpa although the lectotype has capsules that are much NY) has longer capsules (2.0–2.5 mm), even though they
shorter (1.5–2.0 mm) than those in most of the other are somewhat immature, that are more typical of H. cy-
Latin American collections of H. cylindricarpa. The cap- lindricarpa. The other morphological features of the type
sules from this collection are apparently those illustrated specimens of P. bolivianum are typical of Herzogiella
by Brotherus when he described the species in Herzog cylindricarpa.
 number 93 • 13

FIGURE 4. Distribution of Herzogiella cylindricarpa in Latin America.

Specimens examined: MEXICO. Chiapas: Cerro de Las Flores, Iwatsuki & Sharp 532a (CANM). Tlaxcala:
Mozotal, 29 km NW of Motozintla, 2880 m, Cárdenas Mpio. de Tlaxco, Ex-Hacienda de Tlacotla, 2950 m, Reyna
4252 (MEXU); Cima del Cerro Tzontehuitz, close to Tene- s.n. (MEXU, U). GUATEMALA. Chimaltenango: Cerro
japa, 2870 m, Cárdenas 4276 (MEXU, MO, NY); near de Tecpan, region of Santa Elena, 2400–2700 m, Stand-
Las Casas, 2682 m, Sharp 3481 (CANM, MEXU, MICH, ley 60966 (FH, MICH); slopes of Volcán de Acatenango,
TENN). Districto Federal: Desierto de los Leones, Lyonnet above Las Calderas, 2400–2700 m, Standley 61923 (F, FH).
1623 (MEXU); El Zarco, 3200 m, Sánchez & Herrera 51 Huehuetenango: Sierra de los Cuchumatenes, along road
(MEXU). Hidalgo: 10 km N of Pachuca, Mineral del Monte, beyond La Pradera, km 32, ~3300 m, Standley 81813 (FH);
2810 m, Castillo 33 (MEXU); 2 km N of Presa Jaramillo, El above San Mateo Ixtatán, 3048 m, Sharp 4962a (CANM,
Chico, 2920 m, Alfaro & Castillo 112 (MEXU). México: TENN). Quezaltenango: Volcán Santa María, upper NE
Mpio. de Tlalmanalco, 3 km E of San Rafael, 2750–2900 facing slopes to summit of volcano, 3000–4200 m, Steyer-
m, Cisneros 1666 (CANM); Cañada en la Vertiente NW of mark 34086, 34105 (DUKE, F, FH, MICH), 34121 (F, FH,
Iztaccíhuatl, 2690–2980 m, 19°12'N, 98°43'W, Cárdenas MICH); above Los Vahos, Cerro Quemado, 2900–3000 m,
3973 (NY). Popocatépetl, 11 km E of Amecameca, 3000 Standley 86103 (F, FH, MICH); above Chiquival, 2621 m,
m, Held & van Rhijn HM 11 (CANM). Michoacán: Along Sharp 2160 (TENN); Cerro de Sija, 3048 m, Sharp 5033
Federal Hwy. 15, ~50 km W of Cd. Hidalgo, Sharp et al. (CANM, DUKE, F, FH, MICH, TENN); Getena area N of
2301 (CANM); ~46 km W of Cd. Hidalgo, 2700 m, Sharp Sija, 3002 m, Sharp 2287 (MEXU, TENN). Totonicapán:
et al. 2286b (CANM, TENN). Oaxaca: Above Llano de Pacajá region of Desconsuelo, mountains above Totoni-
las Flores in Sierra Juárez on Hwy. 175 between Tuxtepec capán, 3100–3200 m, Standley 84534 (F, FH, MICH); E of
and Oaxaca, 2400 m, Sharp et al. 132 (MEXU, MICH, Totonicapán, 3200 m, Sharp 2619 (DUKE, FH, TENN).
MO, NY), 140a, 2332 (CANM, TENN), 2341 (CANM, COSTA RICA. San José: ~12 km NE of Canáan, on trail
F, MEXU, TENN); N of Oaxaca, along road N of Llano to Cerro Chirripó, 2940–3120 m, Bowers 914-G, 916-I
14 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

(TENN, USJ), 919-I (TENN); Cerros de las Vueltas, irregularly pinnately branched, not or weakly complanate
2700–3000 m, Standley & Valerio 43761b (FH); south- -foliate; stems in cross section dorsiventrally flattened,
ern slope of Volcán de Turrialba, 2000–2400 m, Standley without a hyalodermis, with medium-sized thick-walled
35013 (FH, NY), 35015, 35159, 35217 (FH). COLOM- cells surrounding larger thinner-walled cells, central strand
BIA. Cundinamarca: Mpio. de Subachoque and Supata, small; pseudoparaphyllia filamentous; axillary hairs with
“El Tablazo,” 3500 m, Ireland 23374 (CANM, COL, a single short brown basal cell and 1–2 elongate hyaline
MO, NY), Lewis 88-1234 (CANM, LPB), Lewis 88-1222 apical cells. Stem and branch leaves similar or somewhat
(LPB). VENEZUELA. Mérida: Páramo de Los Granates, differentiated, not or weakly complanate, erect-spreading
3000–3100 m, Griffin PV-942 (CANM). Trujillo: Above to ± homomallous, concave, not or slightly plicate, ovate-
Mpio. Las Piedras, Páramo de Guirigay, 3200 m, Griffin lanceolate to lanceolate, short- to long-acuminate, short-
& López PV-1536 (CANM); Páramo El Jabón, al SE y ar- decurrent; margins subentire to weakly serrulate above,
riba de Carache, 2900–3300 m, Griffin & López PV-1452 plane or recurved below; costa short and double; cells
(NY). ECUADOR. Imbabura: Lago San Marcos, Cayambe oblong- to linear-rhomboidal, smooth or (extralimitally)
Mountain, 3414 m, Cazalet & Pennington 68 (MICH, prorulose, shorter in the acumen; alar cells relatively nu-
NY). Napo: At Km 45 on road from Salcedo to Napo, merous, subquadrate to oblate, with a single decurrent
3800 m, Laegaard 54158CC (NY); 3600 m, 54159M cell. Asexual propagula not seen. Autoicous. Perichaetial
(NY); N side of Laguna Verdecocha, 2 km E of the peak leaves erect, oblong-lanceolate, short- to long-acuminate,
of Cerro Quilindaña, Cord. Oriental, 4100 m, 78°21'W, not or scarcely plicate; margins entire to serrulate; costa
0°47'S, Løjtnant & Molau 11679 (NY). Pichincha: Road none or short and double. Setae elongate, smooth, reddish;
Olmedo–Laguna San Marcos, W of the pass, 3600 m, Øll- capsules inclined to horizontal, arcuate, asymmetric, cylin-
gaard et al. 34341 (CANM, MO, NY). PERU. Amazo- drical, constricted below the mouth when dry and empty;
nas: Prov. Chachapoyas, Chachapoyas-Cajamarca road, exothecial cells rounded-hexagonal to short-rectangular,
~3 km above Leymebamba, 2700 m, Philippi 2334a (NY). ± thick-walled; annulus differentiated; operculum conic-
BOLIVIA. La Paz: Prov. Saavedra, mountain side directly apiculate to obliquely short-rostrate; peristome double,
N of river, beneath Niño Karine, NNW of Chuma, 3250 exostome teeth pale yellow, shouldered, bordered, on the
m, Lewis 79-1044 (CANM, F, NY), 79-1045A (CANM, front surface with a zigzag median line, not furrowed,
F). Prov. Franz Tamayo, along trail between Pelechuco cross-striolate below, papillose above, trabeculate at back;
and Pata, 3050 m, Lewis 88-1852 d-1 (CANM). Prov. In- endostome with a high basal membrane, segments keeled,
quisivi, Río Glorieta, ~8 km NW of Quime, 3200–3400 perforate, smooth or papillose, cilia in groups of 1–3, nod-
m, Lewis 87570 (MO). ulose. Spores spherical, nearly smooth to finely papillose.
Calyptrae cucullate, naked, smooth.
4. Homomallium (Schimp.) Loeske Discussion: Homomallium is somewhat interme-
diate between the erect-capsuled, epiphytic Hypnaceae
Homomallium (Schimp.) Loeske, Hedwigia 46: 314. genera, like Pylaisia or Platygyrium, and more typical
1907; Hypnum subg. Homomallium Schimp., Syn. 616. terrestrial members of the family. It is characterized by
1860; Stereodon sect. Homomallium (Schimp.) C. E. O. relatively short apical leaf cells, numerous quadrate alar
Jensen, Danm. Moss. 2: 115. 1923. Type: Hypnum in- cells and curved, asymmetric capsules with a perfect peri-
curvatum Brid. stome (Ando, 1964). The genus barely makes it into the
Neotropics, with one species in northern Mexico and ad-
Plants small to medium-sized, in flat mats, shiny, soft, jacent United States and the other a little-collected Mexi-
yellow-green to dark green. Stems creeping, regularly to can endemic.

KEY TO THE SPECIES OF HOMOMALLIUM

1. Branches usually homomallous-foliate; stem and branch leaves scarcely differentiated, relatively short-acuminate; margins
usually recurved below; branch leaf alar cells extending up the margins 20–30 cells . . . . . . . . . . . . . . . 1. H. mexicanum
1. Branches not homomallous-foliate; stem and branch leaves differentiated, relatively long-acuminate; margins erect to nar-
rowly incurved; branch leaf alar cells extending up the margins 8–13 cells . . . . . . . . . . . . . . . . . . . . . . . . . . 2. H. sharpii
 number 93 • 15

1. Homomallium mexicanum Specimens examined: MEXICO. Durango: Along

Hwy. 40, W of Durango, near Km 103, 23°48'N, 105°23'W,
Homomallium mexicanum Cardot, Rev. Bryol. 37: 53. ~2355 m, King & Garvey B311 (MO, NY). Guanajuato:
1910. Type: Mexico. Hidalgo: Cuyamaloya Station, Sierra de Pénjamo, 24 km SW de Cuerámaro, 20°35'N,
17 Sept 1908, Pringle, Plantae Mexicanae 10631 (syn- 101°45'W, Delgadillo 5825 (MEXU, NY). Hidalgo: Only
types: NY-3!, US!), 15250 (syntype: specimen not seen). the type seen.

Plants medium-sized, in flat mats, shiny, soft, pale 2. Homomallium sharpii

green to dark green. Stems to 4 cm long, creeping, elongate,
irregularly pinnately branched; branches erect-ascending, Homomallium sharpii Ando & Higuchi, Bryologist 86:
somewhat curved, 2–7 mm long, rarely flagelliform; in 374. 1983 [1984]. Type: Mexico. Tamaulipas: La Perra
cross section with 1–2 rows of medium-sized, thick-walled (El Porvenir), Rancho del Cielo above Gómez Farias,
cells surrounding abruptly larger firm-walled cells, central 6300 ft, Sharp 5641a (holotype: TENN!; isotypes:
strand of small, very thin-walled cells; pseudoparaphyllia HIRO, MICH!).
filamentous; axillary hairs of 1 short brown basal cell and
1–2 elongate hyaline apical cells. Stem and branch leaves Plants medium-sized, in flat mats, shiny, soft, yellow
similar, 1–1.4 mm long, homomallous, strongly concave, -green to yellow-brown. Stems to 4 cm long, creeping,
ovate-lanceolate, broadly long-acuminate; margins sub- elongate, irregularly pinnately branched; branches erect-
entire to serrulate above, entire below, usually narrowly ascending, somewhat curved, to 1 cm long; stems in cross
recurved below, plane above; costa short and double; cells section with 2–4 rows of small, thick-walled cells sur-
long-hexagonal, flexuose, 40–70 µm long at midleaf, mostly rounding abruptly larger firm-walled cells, central strand
5–6 µm wide, as short as 20 µm long in the acumen, smooth; of small, very thin-walled cells; pseudoparaphyllia fila-
alar cells subquadrate to oblate (especially along the mar- mentous; axillary hairs of 1 short brown basal cell and 1
gins) in 6–8 rows, extending up the margins by 20–30 cells. elongate hyaline apical cell. Stem and branch leaves some-
Perichaetial leaves erect, oblong-lanceolate, convolute, what differentiated; stem leaves 0.9–1.1 × 0.4–0.6 mm,
gradually narrow-acuminate; margins subentire. Setae 0.5– ± homomallous, ± falcate, concave, broadly ovate-lan-
1.5 cm long, reddish; capsules 1.3–1.5 mm long, suberect to ceolate, long-acuminate; branch leaves not homomallous,
inclined, cylindrical; annulus of 1 row of thick-walled cells; 1–1.3 × 0.3–0.5 mm, ovate-lanceolate, long-acuminate;
operculum ~0.4 mm long, mostly obliquely short-rostrate; margins subentire to serrulate above, entire below, usu-
exostome teeth cross-striolate below, coarsely papillose ally erect above, plane below; costa short and double; cells
above, trabeculate at back; endostome yellowish, papillose long-hexagonal, flexuose, (30–)40–60 µm long at midleaf,
throughout, with a high basal membrane, segments keeled, mostly 3–5 µm wide, as short as 25 µm long in the acu-
perforate, cilia mostly in pairs. Spores 15–20 µm in diam- men, smooth; alar cells subquadrate to oblate (especially
eter. Calyptrae cucullate, naked, smooth. along the margins) in 6–8 rows, extending up the margins
Distribution and ecology: Northern Mexico; also in by 8–13 cells. Perichaetial leaves erect, oblong-lanceolate,
southwestern United States; growing on bases of hardwood convolute, gradually narrow-acuminate; margins sub-
trees, logs, and rocks, mostly at (1500–)2000–2700 m. entire. Setae 1–1.8 cm long, reddish; capsules 1.3–2 mm
Illustrations in publications: Fig. 755 in Ando long, suberect to inclined, cylindrical; annulus of 1 row
(1994: 1016). of thick-walled cells; operculum ~0.4 mm long, mostly
Discussion: When sterile, the plants might be con- conic-apiculate; exostome teeth cross-striolate below,
fused with Pylaisia because of the numerous alar cells. coarsely papillose above, trabeculate at back; endostome
However, the shorter apical laminal cells are decidedly dif- yellowish, papillose throughout, with a high basal mem-
ferent than those in that genus. When sporophytes are pres- brane, segments keeled, perforate, cilia in groups of 2–3.
ent (frequent), the inclined, asymmetric capsules separate it Spores 13–15 µm in diameter. Calyptrae cucullate, naked,
from the erect, symmetric capsules of Pylaisia. Homomal- smooth.
lium mexicanum differs from H. adnatum (Hedw.) Broth., Distribution and ecology: Northeast Mexico; en-
of eastern North America, by its larger size (leaves greater demic; growing on limestone, at 1900–2500 m.
than 1 mm long), longer apical laminal cells, and recurved Illustrations in publications: Fig. 756 in Ando
lower leaf margins. (1994: 1018).
16 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Discussion: This species differs from H. mexicanum above, somewhat incurved below; costa short and double
in the nonhomomallous branch leaves with longer acumina or lacking; median cells 47–94 × 5–9 µm, smooth or rarely
and fewer alar cells. The stem and branch leaves are differ- minutely prorulose at upper ends on dorsal surface; alar
entiated much more so than in H. mexicanum. The spores cells strongly differentiated, quadrate to rectangular, in
are smaller and the peristomial cilia are in groups of 2–3. 1 to several rows with 5–15 cells in the margin. Asexual
Specimens examined: MEXICO. San Luis Potosí: propagula none. Dioicous; rarely producing sporophytes.
Just E of San Francisco, 45 km E of San Luis Potosí, at Km Setae 1.2 cm long, brown to reddish, flexuose; capsules
218 along Hwy. 70, 2166 m, Magill 2591a, 2593 (HIRO, 1.5 mm long, inclined, subarcuate, yellowish brown, ob-
TENN); on hillsides along Hwy. 70 near Km 226, 37 km E long-ovoid, somewhat contracted below mouth when
of San Luis Potosí, 2318 m, Magill 2577 (HIRO, TENN). dry; operculum unknown; annulus present, persistent, of
Tamaulipas: Only the type seen. 2 rows of cells; peristome double, exostome teeth cross-
striolate below, papillose above, bordered, trabeculate
5. Irelandia W. R. Buck at back; endostome with a high to low basal membrane,
keeled segments and cilia shorter than or approximately
Irelandia W. R. Buck, Brittonia 36: 179. 1984b. the same length as the segments, in groups of 1–3. Spores
9–19 µm in diameter. Calyptrae not seen.
This is a monospecific genus with the characters of Distribution and ecology: Rare or seldom col-
the species. lected; Mexico, Guatemala, Honduras, Jamaica, and Haiti
(Figure 6); occurring on limestone bluffs, on shaded banks,
Irelandia robusticaulis or on bark of trees in forests, 250–1800 m.
Discussion: This monospecific genus and its dis-
FIGURE 5 tinctive species are characterized by its stiff, dark green
to yellow-green, subjulaceous plants, imbricate or erect-
Irelandia robusticaulis (E. B. Bartram) W. R. Buck, Brit- spreading, concave, ovate-lanceolate, acuminate leaves with
tonia 36: 179. 1984b. margins broadly recurved above, and strongly differentiated
alar regions of 1 to several rows of quadrate to rectangular
Isopterygium robusticaule E. B. Bartram, Bryologist 49: 122. 1946; Taxiphyl- cells with 5–15 differentiated cells along the margin.
lum robusticaule (E. B. Bartram) Ireland, Bryologist 87: 360. 1984 (1985). Buck (1984b), who described this monospecific genus
Type: Guatemala, Dept. Alta Verapaz, Montaña Yxocubvain, 2 ½ mi. W of in honor of the senior author, distinguished it from Taxi-
Cubilgüitz, alt. 300–500 m, Steyermark 44970a (holotype: FH!; isotypes: F!, phyllum primarily on the basis of its stiff plants with erect-
TENN!, US!); Cerro de Agua Tortuga (Sahacoc), vicinity of Cubilgüitz, alt. spreading leaves, noncomplanate foliate branches, and
350–450 m, Steyermark 44584 (paratypes: FH!, MICH!). leaves more or less abruptly long-acuminate with a much
better developed alar region with numerous rectangular
Isopterygium guatemalense E. B. Bartram, Bryologist 49: 123. 1946. Type: cells along the leaf margins. In addition to these charac-
Guatemala, Dept. Chimaltenango, between Chimaltenango and San Martín ters, the plants have conspicuous broadly recurved leaf
Jilotepeque, Standley 80937 (holotype: FH!; isotypes: F!, MICH!), 64364 margins near their apices.
(paratype: FH!). The senior author (Ireland, 1984) placed this species,
which was originally described as an Isopterygium by
Plants in thin, loose mats, glossy, dark green to yellow- Bartram (1946), in the genus Taxiphyllum because it has
green. Stems to 4.5 cm long, 1.5–2.5 mm wide, subjula- foliose pseudoparaphyllia, a julaceous habit, and a large
ceous, simple or irregularly branched; stems in cross section number of quadrate to rectangular alar cells on the leaf
with 1–3 rows of small thick-walled cells surrounding margins. These features also occur in Taxiphyllum cuspidi-
larger somewhat thinner-walled cells, central strand small, folium (Cardot) Z. Iwats. from the southern United States.
consisting of thin-walled cells; pseudoparaphyllia foliose; However, the rigid stems and the strongly recurved distal
axillary hairs with 1–3 short brown basal cells and a single leaf margins of T. robusticaule do not occur in T. cuspida-
elongate hyaline distal cell. Stem and branch leaves similar, tum or any other species of Taxiphyllum and seem to place
1.5–2.0(–2.5) mm long, 0.5–0.8 mm wide, rigid, imbricate the genus Irelandia apart from Taxiphyllum. These distinc-
or erect-spreading, concave, smooth, ovate-lanceolate, tions, however, may not be strong enough to support es-
symmetric, acuminate; margins entire to serrulate or ser- tablishment of a new genus. Future DNA studies may be
rate above, serrulate or entire below, broadly recurved helpful in deciding if Irelandia is truly worth recognizing.
 number 93 • 17

FIGURE 5. Irelandia robusticaulis. A. Habit. B. Enlargement of stem. C. Leaves (leaf above, view of dorsal surface; leaf below, view of ventral
surface). D. Pseudoparaphyllia. E. Apical leaf cells. F. Median leaf cells. G. Marginal leaf cells. H. Alar cells. I. Portion of stem cross section. All
illustrations drawn from Sharp 1378a (TENN).

Buck (1984b) did not include Isopterygium guate- of Taxiphyllum robusticaule, it has been included here in
malense E. B. Bartram in the synonymy of Irelandia at the synonymy of the species until a more detailed study
the time he described the genus. Buck (1998: 323) later of the important characters of the two taxa can be made.
stated that he was unconvinced that it belongs in the genus Specimens examined: MEXICO. Puebla: El Cerro
because it is “a much softer plant that has more or less de Cuhuatepec, 1140 m, Sharp 1378a (TENN, US). GUA-
plicate leaves and shorter acumina.” However, since the TEMALA. Alta Verapaz: Near Río Icvolay, near Haci-
senior author (Ireland, 1984) included it in the synonymy enda Yaxcabnal, 8 km NW of Cubilgüitz, 250–300 m,
18 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 6. Distribution of Irelandia robusticaulis in Latin America.

Steyermark 44702 (F, FH). Baja Verapaz: Patal, 1615 m, Plants small, in thin to dense mats, glossy, light green
Sharp 2827 (MEXU, US). Chimaltenango: Near Río Pix- to yellowish green. Stems sometimes complanate-foliate,
cayó, between Chimaltenango and San Martín Jilotepeque, creeping, simple or sparingly and irregularly branched,
1650–1800 m, Standley 64364 (F, FH). Retalhuleu: Dept. hyalodermis distinct with large cells or somewhat indis-
Quezaltenango, near Chivolandia, along road to San Felipe, tinct with small cells, central strand absent or sometimes
650 m, Standley 87209a (FH). Zacapa: Upper reaches of present and indistinct; rhizoids papillose, in clusters in
Río Sitio Nuevo, between Santa Rosalía and first waterfall, leaf axils; pseudoparaphyllia lacking; axillary hairs con-
1200–1500 m, Steyermark 42208 (F, FH). HONDURAS. sisting of 1 short to slightly long, brown basal cell and
Comayagua: Vicinity of Siguatepeque, ~1050 m, Standley 1–3 elongate, hyaline distal cells. Asexual reproductive
& Chacón 6704 (F). Choluteca: Creek flowing to Río Cho- bodies 30–90 µm long, sometimes present in clusters in
luteca, near La Vesto PM, Olson 83-2 (CANM, MO). Lem- leaf axils on stems and branches, cylindrical or fusiform,
pira: Montana de Celaque, along Río Arquegual, 7.5 km composed of 2–6 smooth cells. Stem and branch leaves
SW of Gracias, 14°34'N, 88°39'W, Allen 11152, 11166 similar, somewhat rigid, crowded and imbricate, erect-
(MO). JAMAICA. Portland: ~3.2 km SW of Ecclesdown, spreading, occasionally secund, smooth, flat or somewhat
686 m, Farr 1358 (CANM). Trelawny: Cockpit Co., just concave, symmetric, nondecurrent, lanceolate or ovate- to
N of Wilson’s Run, 9 km N of bridge at Troy, Buck 5899 oblong-lanceolate, long-acuminate to abruptly acuminate;
(NY), Crosby 13878 (NY). HAITI. Rivière Glace, Morne la margins entire to minutely serrulate, plane to erect; costa
Hotte, 800–1000 m, Holdridge 3090 (CANM, NY). lacking or short and double; median cells often flexuose,
firm-walled, linear-fusiform, smooth or prorulose at upper
6. Isopterygiopsis Z. Iwats. ends on dorsal leaf surface, with walls not pitted; alar cells
not differentiated or sometimes with 1–3 quadrate to
Isopterygiopsis Z. Iwats., J. Hattori Bot. Lab. 33: 379. short-rectangular cells on margins. Autoicous or dioicous;
1970. Type: Plagiothecium muellerianum Schimp., Syn. perigonia and perichaetia clumped at base of stems and
Musc. Eur. 584. 1860. branches, bracts small, lanceolate to ovate, acuminate to
 number 93 • 19

abruptly filiform-acuminate; margins plane. Setae smooth, tic, in boreal and temperate woods, or at high altitudes and
elongate, straight to somewhat curved, twisted, red to red- latitudes in Latin America.
dish brown; capsules erect to cernuous, straight to subar- Discussion: Isopterygiopsis was described by
cuate, yellow to orange-brown, oblong to ovoid, smooth, Iwatsuki (1970) as a segregate of the genus Isopterygium.
contracted below mouth and wrinkled at neck when dry; Seventeen years later, he further refined the generic charac-
operculum conic to obliquely rostrate, shorter than the teristics of Isopterygiopsis while separating another genus,
urn; annulus present, deciduous, of 2–3 rows of cells; peri- Pseudotaxiphyllum, which also lacks pseudoparaphyllia
stome double, exostome teeth cross-striolate below, papil- like Isopterygiopsis, from Isopterygium, which has filamen-
lose above, bordered, trabeculate at back; endostome with tous pseudoparaphyllia (Iwatsuki, 1987). Isopterygiopsis is
a high to low basal membrane, keeled segments, and cilia now distinguished by its small, scarcely branched, glossy
shorter than the segments, in groups of 1–3, sometimes plants; large to small hyalodermis cells; papillose rhizoids in
absent. Spores spherical to ovoid, smooth or minutely the leaf axils; absence of pseudoparaphyllia; nondecurrent,
papillose. Calyptra cucullate, smooth, naked. entire or minutely serrulate leaves; 2–6 celled, smooth, cy-
Distribution: The genus Isopterygiopsis has only lindrical or fusiform propagula in clusters in the leaf axils;
three species in the world, two of which occur in Latin autoicous or dioicous sexual condition; and the presence of
America. The species occupy terrestrial habitats in the Arc- an annulus.

KEY TO THE SPECIES OF ISOPTERYGIOPSIS

1. Stems complanate-foliate, appearing ventrally concave because of curvature of upturned leaves; leaves often abruptly
acuminate, cells smooth or dorsally prorulose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. I. muelleriana
1. Stems not complanate-foliate, leaves erect-spreading, occasionally secund; leaves gradually acuminate, cells smooth . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. I. pulchella

1. Isopterygiopsis muelleriana none or short and double; median cells 50–80 × 2–5 µm,
smooth or prorulose at upper ends on dorsal leaf surface;
FIGURE 7 alar cells not differentiated or sometimes with 1–3 short-
rectangular cells on margins. Sex organs and sporophytes
Isopterygiopsis muelleriana (Schimp.) Z. Iwats., J. Hat- not seen on Latin American plants, known to be dioicous
tori Bot. Lab. 33: 379. 1970. Plagiothecium muelleria- from regions where it is fertile and possesses sporophytes.
num Schimp., Syn. Musc. Eur. 584. 1860; Isopterygium Setae 0.6–1.2 cm long, brown to reddish brown; capsules
muellerianum (Schimp.) A. Jaeger, Ber. Thätigk. St. 0.5–1.5 mm long, erect or rarely ± cernuous, straight, el-
Gallischen Naturwiss. Ges. 1876–77: 441. 1878. Type: lipsoid to ovoid, contracted below the mouth when dry;
Austria?, 30 Sep 1851, J. Müller (holotype: BM!). operculum conic to obliquely rostrate; annulus present, de-
ciduous, of 2–3 rows of cells. Spores 8–12 µm in diameter.
Plants in thin mats, glossy, yellowish green. Stems to Distribution and ecology: Rare; known only from
2 cm long, 0.8–2.0 mm wide, complanate-foliate, appear- a single Mexican collection (Figure 8); occurring on a bluff
ing ventrally concave because of curvature of upturned at 3139 m.
leaves, simple or irregularly branched; hyalodermis of large Discussion: Isopterygiopsis muelleriana is a species
epidermal cells, enclosing 1 to several layers of small, thick- that is rare or rarely collected in Latin America. It is best
walled cortical cells and several large, thin-walled cells in recognized by its plants that are complanate-foliate, appear-
the center. Asexual reproductive bodies not seen on Latin ing concave on the upper surface because of the upturned
American plants, known on plants from elsewhere as ax- leaves, and by the stems that have a hyalodermis with large,
illary clusters of 2–6 celled, smooth, cylindrical or fusi- thin-walled cells (best seen in cross section) with papillose
form, green to yellowish green brood bodies in leaf axils rhizoids clustered in the leaf axils. Other helpful features dis-
on stems and branches. Leaves 0.5–1.0 mm long, 0.2– tinguishing the species, are the ovate- to oblong-lanceolate,
0.3 mm wide, somewhat rigid, crowded and overlapping, nondecurrent leaves, often abruptly acuminate with mar-
erect-spreading and rigidly complanate, smooth, ovate- to gins that are entire or minutely serrulate, the median leaf
oblong-lanceolate, symmetric, acuminate, often abruptly cells that are 50–80 × 2–5 µm, often prorulose at the distal
so; margins plane, entire or minutely serrulate; costa ends on the dorsal leaf surface, the few differentiated alar
20 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 7. Isopterygiopsis muelleriana. A. Habit. B. Enlargement of portion of stem. C. Leaves. D. Apical leaf cells. E. Median-marginal leaf
cells. F. Median leaf cells. G. Alar cells. H. Stem cross section. I. Portion of rhizoid. All illustrations drawn from Sharp 64 (CANM).

cells, with 1–3 short-rectangular ones on the margins. In ad- 2. Isopterygiopsis pulchella
dition, the plants are dioicous, and they sometimes possess
smooth, cylindrical or fusiform brood bodies clustered in FIGURE 9
leaf axils on the stems and branches.
Specimen examined: MEXICO. Distrito Federal: Isopterygiopsis pulchella (Hedw.) Z. Iwats., J. Hattori
Desierto de los Leones near Mexico City, 3139 m, Sharp 64 Bot. Lab. 63: 450. 1987. Leskea pulchella Hedw., Spec.
(CANM, MEXU, MICH, TENN). Musc. Frond. 220. 1801; Isopterygium pulchellum
 number 93 • 21

FIGURE 8. Distribution of Isopterygiopsis muelleriana (▲) and I. pulchella (●) in Mexico.

(Hedw.) A. Jaeger, Ber. Thätigk. St. Gallischen Natur- sometimes distant, erect-spreading, occasionally secund,
wiss. Ges. 1876-77: 441. 1878. Type: Scotland, no col- often appearing distichous, subconcave, smooth, lanceo-
lector cited (holotype: G!). late to slenderly ovate-lanceolate, symmetric, gradually
long-acuminate; margins plane to erect throughout, en-
Hypnum magellanicum Müll. Hal., Flora 68: 425. 1885, hom. illeg.; Isop- tire or sometimes minutely serrulate, often with 1–2 cells
terygium fuegianum Besch., Miss. Sci. Cape Horn, Bot. 5: 301. 1889 (new distinctly serrulate in alar region; costa none or short and
name for Hypnum magellanicum); Plagiothecium magellanicum (Besch.) double; median cells 47–122 × 5–7 µm, smooth; alar cells
Paris, Index Bryol. 963. 1896. Type: Chile, Clarence Is., S. Holl Bay, Hariot differentiated only on margins, 1–3 quadrate to short-
63 (holotype: BM!; isotype?: H!). Syn. nov. rectangular cells, sometimes lacking. Autoicous. Setae
1–2 cm long, red to reddish brown; capsules 0.5–2.5 mm
Plagiothecium leptoplumosum Dusén, Rep. Princeton Univ. Exp. Patagonia, long, subcernuous to cernuous, rarely erect, straight to
Botany 8: 100. 1903; Isopterygium leptoplumosum (Dusén) Cardot, Wiss. subarcuate, oblong to ovoid, contracted below the mouth
Erb. Schwed. Südpolar-Exp. 4(8): 178. 1908. Type. Chile, Patagonia, Río when dry; operculum conic to conic-apiculate; annulus
Chico, Hatcher s.n. (holotype: S!; isotype: NY!). Syn. nov. present, deciduous, of 2–3 rows of cells. Spores 9–14 µm
in diameter.
Plants in thin to dense mats, glossy, light green to Distribution and ecology: Infrequent; Mexico,
yellowish green. Stems to 2 cm long, 1–2 mm wide, Chile, and Argentina (Figures 8, 10); on soil, in crevices of
simple or irregularly branched; hyalodermis present but rock cliffs, and on rotting tree trunks at 150–3650 m and
somewhat indistinct from the 1 to several layers of inner sometimes higher.
thick-walled cortical cells. Asexual reproductive bodies Discussion: Isopterygium fuegianum Besch. and
30–60 µm long, rarely present, cylindrical or fusiform, I. leptoplumosum (Dusén) Cardot are considered here as
clustered in leaf axils on stems and branches, green to new synonyms of I. pulchella. No important differences
yellowish green, composed of 2–5 smooth cells. Leaves could be found for maintaining either species as separate
0.5–1.5 mm long, 0.2–0.4 mm wide, flaccid, close or taxa from I. pulchella. Even Dusén and Bescherelle stated
22 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 9. Isopterygiopsis pulchella. A. Habit. B. Enlargement of portion of stem. C. Leaves. D. Apical leaf cells. E. Median-marginal leaf cells.
F. Median leaf cells. G. Alar cells. H. Stem cross section. I. Portion of rhizoid. J. Capsule (wet). Illustrations A, B drawn from Dusén 22 Feb 1897
(M); C–J from Hatcher, holotype of Plagiothecium leptoplumosum (S).

that their new species were very similar to I. pulchella (47–122 µm) and narrow (5–7 µm), smooth, median leaf
when they described them. cells, the alar regions undifferentiated or sometimes with
Isopterygiopsis pulchella is recognized by the erect- 1–3 short-rectangular to quadrate cells on the margins, the
spreading, sometimes secund, lanceolate to slenderly occasional presence of cylindrical to fusiform, 2–5-celled
ovate-lanceolate, gradually acuminate, nondecurrent brood bodies clustered in the leaf axils and the autoicous
leaves with entire or minutely serrulate margins, the long sexual condition.
 number 93 • 23

FIGURE 10. Distribution of Isopterygiopsis pulchella in South America.

Bryologists may sometimes have difficulty placing Ochyra 6908, Musci Fuegiani Exsiccati 48, (MO); Tierra
I. pulchella into Isopterygiopsis because the stem hyalo- del Fuego australis, Valle del Río Pipo, 300 m, 54°46'S,
dermis cells of I. pulchella are not as distinctly enlarged 68°20'W, Roivainen 1352 (NY); Arroyo Grande, 150 m,
and thin-walled as those of the type species of the genus, 54°47'S, 68°17'W, Roivainen 2095 (MO, NY); E Glaciar
I. muelleriana. However, the other important features dis- Martial, Kuc 107 (CANM); Santa Cruz, in valley above
tinguishing the genus Isopterygiopsis and placing I. pul- Río Santa Cruz, Jan 1905, Dusén s.n. (H); Lago Argentino,
chella within it are present, i.e., lack of pseudoparaphyllia, Dusén 5684 (NY).
presence of cylindrical or fusiform propagula, and papil-
lose rhizoids in the leaf axils. 7. Isopterygium Mitt.
Specimens examined: MEXICO. Mpio. Toluca,
Parque Nacional Nevado de Toluca, NW slopes of Nevado Isopterygium Mitt., J. Linn. Soc., Bot. 12: 21. 1869. Type:
de Toluca volcano, 3650 m, 19°08'N, 99°47'W, Buck Isopterygium tenerum (Sw.) Mitt., J. Linn. Soc., Bot.
28203 (NY); Mt. Orizaba, Sep 1908, Purpus s.n. (FH). 12: 499. 1869. (Lectotype designated by Iwatsuki and
CHILE. Bío-Bío Region: Bío-Bío Prov., National Park Lake Crosby, 1979).
Laja, Los Barros Military Base, 1500 m, 37°27'S, 71°09'W,
Ireland & Bellolio 34124 (CONC, MO, US). Magellanes Plants small to medium-sized, in thin to dense mats,
Region: Antártica Chilena Prov., Communa Cabo de Hor- glossy, light green to yellow-green. Stems complanate-
nos, Parque Nacional Alberto de Agostini, E end of Isla foliate, creeping, simple or sparingly and irregularly
Gordon, S side of Punta Divide on Brazo Sudoeste of Beagle branched, cortical cells small and thick-walled in cross
Channel, 54°58'S, 69°08'W, Buck 47943 (NY); Clarence section, surrounding larger, thinner walled cells, central
Is., Hariot 63 (BM); Patagonia, Río Chico, Hatcher s.n. strand usually absent; rhizoids smooth, on ventral surface
(NY, S); Patagonia, Aysén, Dusén 584 (FH, H, M, NY); of stems and branches just below juncture of leaves; fila-
Río Aysén, 22 Feb 1897, Dusén s.n. (M); Punta Arenas, mentous pseudoparaphyllia present, of 3–6 cells in 1 row
Thaxter 161 (FH), Dusén 63 (H); ~35 km from Puerto Na- or rarely 2 rows near base; axillary hairs consisting of 1
tales, inside Cueva de Milodon, Kuc 117 (CANM). AR- moderately long, brown basal cell and 1–3 elongate, hya-
GENTINA. Dept. Ushuaia, Camino de ascenso al Glaciar line distal cells. Asexual reproductive bodies sometimes
Le Martial, ~900–1200 m, 54°46'S, 68°29'W, Schiavone & present on stems and branches, uniseriate, often branched,
24 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

filamentous, multicellular bodies with papillose cells. Stem 1987; Sainsbury, 1955), placed Isopterygium in the family
and branch leaves similar, rigid or flaccid, crowded and Plagiotheciaceae. While other bryologists (Crum and An-
imbricate to remote, erect-spreading or squarrose, some- derson, 1981; Gangulee, 1978; Vitt, 1984) have placed it
times contorted when dry, smooth, flat or somewhat con- in the Hypnaceae. Buck and Ireland (1985), on the basis of
cave, symmetric or asymmetric, nondecurrent or rarely a morphological evaluation of the Plagiotheciaceae genera,
with 1–2 decurrent cells, ovate or lanceolate, sometimes removed all genera except Plagiothecium from the family.
oblong, acute to acuminate; margins plane to erect, some- Isopterygium is here placed in the Hypnaceae.
times recurved at base, serrulate above the middle, mostly Mitten (1869) originally placed eight species in Isopte-
entire below, sometimes entire throughout; costa short rygium: I. affusum (Müll. Hal.) Mitt., I. curvicolle (Müll.
and double, occasionally lacking; median cells often flex- Hal.) Mitt., I. leucophyllum (Hampe) Mitt., I. tenerum
uose, thin to firm-walled, linear-fusiform, smooth, with Mitt., I. planissimum Mitt., I. tenerifolium Mitt.,
walls not pitted or occasionally those of basal cells pitted; I. brachyneuron (Müll. Hal.) Mitt., and I. chrismarii (Sw.)
alar cells usually clearly differentiated, quadrate to rect- Mitt. However, the genus soon became a repository for
angular, rarely transversely elongate. Autoicous or rarely scores of miscellaneous, complanate-foliate pleurocarpous
dioicous; perigonia scattered along the stems; perichaetia mosses. The genus grew enormously during the next cen-
at base of stems, leaves oblong-lanceolate, gradually acu- tury, eventually comprising about 390 taxa, 92 of which
minate; margins plane. Setae smooth, elongate, straight to have been reported for Latin America (van der Wijk et al.,
curved, usually twisted, yellow, brown, or reddish brown; 1964, 1969).
capsules inclined to cernuous, or sometimes erect, straight Taxonomic studies during the last 20–30 years now
or curved when mature, brown to red-brown, cylindri- make it possible to clearly delimit the genus Isopterygium
cal, ellipsoid or ovoid, smooth or sometimes wrinkled at as it occurs in Latin America and elsewhere. The Japanese
neck and usually contracted below the mouth when dry; bryologist Zennoske Iwatsuki has done more than anyone
operculum conic to short-rostrate, shorter than the urn; to clarify the classification of this complex and disorganized
annulus none; peristome double, exostome teeth cross- genus. His studies of Taxiphyllum M. Fleisch. (Iwatsuki,
striolate below, papillose above, bordered, trabeculate 1963), Herzogiella Broth. (Iwatsuki, 1965, 1970, synonyms
at back; endostome with a high to low basal membrane, Dolichotheca Lindb., Sharpiella Z. Iwats.), Plagiothecium
keeled segments, and cilia shorter than the segments, in (Iwatsuki, 1970), Isopterygiopsis Z. Iwats. (Iwatsuki,
groups of 1–3, sometimes absent. Spores spherical to 1970), and Isopterygium Mitt. and Pseudotaxiphyllum
ovoid, smooth or minutely papillose. Calyptra cucullate, Z. Iwats. (Iwatsuki, 1987) have clarified the taxa in the
smooth, naked. genera closely related to Isopterygium, thus making that
Distribution: Isopterygium is a genus of approxi- genus more clearly defined. Studies of the North American
mately 40–50 species worldwide that in the subtropics and (Ireland, 1969), Mexican (Ireland, 1984), and Latin Ameri-
tropics occur predominantly in terrestrial habitats at low can (Ireland, 1991a, 1992) Isopterygium species also have
elevations. helped to understand the genus to a certain extent.
Discussion: Isopterygium has been considered a We are recognizing eight species of Isopterygium in
taxonomically difficult genus since it was described by Mit- the present treatment. The species are often very similar
ten (1869). The taxonomic characters Mitten used to estab- morphologically and difficult to separate even when spo-
lish Isopterygium were not distinct enough to distinguish it rophytes are present. Some of the species may be only en-
from many other genera, and this has caused a great amount vironmental forms. However, we believe it is prudent to
of confusion. Mitten placed his new genus in the tribe Ste- recognize these eight species at this time until future field
reodonteae, on the basis of its long setae, short, double-leaf and laboratory studies prove otherwise.
costae, and short, dense alar cells. The tribe contained an Isopterygium is distinguished by the complanate-
assemblage of genera, many of which are not considered foliate plants, smooth rhizoids inserted on ventral surface
very closely related today, such as Plagiothecium Bruch of stems and branches just below leaf insertions, filamen-
& Schimp. in Bruch, Schimp. & W. Gümbel, Ctenidium tous, 3–6-celled pseudoparaphyllia, small and thick-walled
(Schimp.) Mitt., Heterocladium Bruch & Schimp. in Bruch, cortical stem cells, usually serrate apical leaf margins, oc-
Schimp. & W. Gümbel, Leucomium Mitt., Ectropothecium casional presence on the stems of simple or branched,
Mitt., Entodon Müll. Hal., Pleurozium Mitt., Acrocladium filamentous, multicellular propagula with papillose cells,
Mitt., and Ptychomnion (Hook. & Wilson) Mitt. Some usually autoicous sexual condition and capsules that lack
bryologists (Brotherus, 1923; Corley et al., 1981; Iwatsuki, an annulus.
 number 93 • 25

KEY TO THE SPECIES OF ISOPTERYGIUM

1. Leaves narrowly lanceolate, mostly more than 3 times as long as broad, usually distant, wide-spreading to squarrose, straight
and symmetric, alar cells on margins mostly elongate- rectangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. I. subbrevisetum
1. Leaves ovate, ovate-lanceolate, oblong or occasionally lanceolate, mostly less than 3 times as long as broad, close or rarely
distant, erect to squarrose, often curved and asymmetric, alar cells on margins mostly quadrate to short-rectangular . . . . 2
2. Leaves with alar regions strongly differentiated, composed of quadrate to rectangular cells in several rows, 4–15 cells on
margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Leaves short, 0.4–0.8 mm in length, acuminate; plants terrestrial . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. I. byssobolax
3. Leaves long, 0.8–1.2 mm in length, acute; plants aquatic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. I. acutifolium
2. Leaves with alar regions weakly differentiated, only a small group of quadrate to short- rectangular cells present, 1–5 cells
on margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Leaves 1–3 mm long; plants aquatic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. I. affusum
4. Leaves 0.7–1.5 mm long; plants terrestrial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Capsules erect or rarely inclined, not or scarcely contracted below mouth when dry . . . . . . . . 4. I. jamaicense
5. Capsules inclined to horizontal, rarely erect, usually strongly contracted below mouth when dry . . . . . . . . . 6
6. Capsules subglobose, about as broad as long, not contracted below mouth when dry . . . . . 6. I. subglobosum
6. Capsules ovoid to ellipsoid, longer than broad, strongly contracted below mouth when dry . . . . . . . . . . . 7
7. Plants large, stems often 2–4 cm long; leaves 1.0–1.5 mm long; setae usually 2–3 cm long; asexual repro-
ductive bodies lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. I. tenerifolium
7. Plants small, stems seldom over 2 cm long; leaves 0.7–1.2 mm long; setae usually 0.5–1.2 cm long;
asexual reproductive bodies sometimes present on stems, filamentous, simple or branched, multicellular,
with papillose cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. I. tenerum

1. Isopterygium acutifolium Discussion: Known only from sterile plants but dis-
tinguished from the other Latin American species of Isopte-
FIGURE 11 rygium by the acute leaves that have prominent alar regions
composed of several rows of quadrate to rectangular cells
Isopterygium acutifolium Ireland, Bryologist 93: 343. with 4–15 cells on the margins. All other Isopterygium species
1990. Type: Venezuela. Bolivar, Meseta de Jáua, NNE have acuminate leaves, and except for Isopterygium bysso-
of Mission of Campamento Sanidad of the Kanarakuni bolax, they have only a few differentiated cells in the alar re-
R., Steyermark 97784 (holotype: US!). gions. Isopterygium byssobolax has about half the number of
quadrate to rectangular alar cells that I. acutifolium possesses.
Plants in thin mats, light green to yellowish green. The aquatic habit is also a clue to the identity of I. acutifo-
Stems up to 2.5 cm long, 1–2 mm wide, simple or sparingly lium since I. affusum is the only other aquatic species in Latin
and irregularly branched. Asexual reproductive bodies America. Isopterygium tenerum sometimes occurs in swamps
lacking. Leaves 0.8–1.2 mm long, 0.4–0.5 mm wide, some- or on river banks but is never found on rocks in rivers.
what rigid, distant, flat to slightly concave, erect-spreading, Specimens examined: VENEZUELA. Known only
smooth, scarcely contorted when dry, ovate-lanceolate or from the type specimen. GUYANA. First falls of Essequibo
oblong, symmetric to somewhat asymmetric, acute; mar- River, Richards 359 (NY).
gins erect, serrulate from apex to leaf middle or just below;
costa none or weak, short and double; median cells 70– 2. Isopterygium affusum
103 × 7–9 µm, smooth; alar cells strongly differentiated,
quadrate to rectangular, in several rows with 4–15 cells on FIGURE 13
margins. Sex organs and sporophytes unknown.
Distribution and ecology: Known only from the Isopterygium affusum Mitt., J. Linn. Soc., Bot. 12: 499.
type locality in Venezuela and from one locality in Guyana 1869. Type: Brazil. Taruma River, Spruce 1059 (lecto-
(Figure 12); attached to rocks near waterfalls and in run- type: NY!, designated by Iwatsuki and Crosby, 1979);
ning water along rapids of rivers, apparently at altitudes isolectotype: BM!); Prov. Ceara, Sierra de Araripe,
of ~400 m. Gardner 106c (synype: BM!).
26 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 11. Isopterygium acutifolium. A. Habit. B. Enlargement of portion of stem. C. Leaves. D. Apical leaf cells. E. Median leaf cells. F. Alar
cells. G. Stem cross section. H, I. Pseudoparaphyllia. All illustrations drawn from Steyermark 97784 (US, holotype).

Isopterygium herminieri Schimp. ex Besch., Ann. Sci. Nat. Bot. sér. 6,3: 256. Nat. Pflanzenfam. 1(3): 1081. 1908. Type: Brazil. Mato Grosso: Macoco
1876. Type: Guadeloupe. In rivers or streams, Herminier s.n. (holotype: River, Lindman 502 (holotype: H!; isotype: BM!).
BM!; isotype: NY!).
Plagiothecium fontigenum Müll. Hal., Hedwigia 40: 58. 1901; Isopterygium
fontigenum (Müll. Hal.) W. R. Buck & Ireland, Fl. Neotrop. 50: 19. 1989.
Ectropothecium submersum Broth., Bih. Kongl. Svenska Vetensk.-Akad. Type: Brazil. Santa Catarina: Serra do Oratorio, Ule 521 (lectotype: H!, des-
Handl. 26 Afd. 3(7): 48. 1900; Isopterygium submersum (Broth.) Broth., ignated by Buck and Ireland, 1989).
 number 93 • 27

FIGURE 12. Distribution of Isopterygium affusum (), I. byssobolax (▲), I. subbrevisetum (●), I. subglobosum (■) and I. acutifolium () in
South America.

Isopterygium aquaticum H. Rob., Bryologist 67: 453. 1964, hom. illeg., -foliate, simple or irregularly branched. Asexual repro-
non Dixon 1922; Isopterygium irelandii H. Rob., Bryologist 70: 43. 1967. ductive bodies lacking. Leaves flaccid, close, complanate,
Type: Honduras. Dept. Francisco Morazán, Río Agua Amarilla, NE slopes erect-spreading, smooth, usually wrinkled and contorted
of Cerro de Uyuca, Morton 7566 (holotype: US!). when dry, 1–3 mm long, 0.4–1.0 mm wide, ovate-
lanceolate to ovate, symmetric or often asymmetric, acu-
Plants in thin, loose masses, yellow-green to green. minate to long-acuminate; margins plane, serrulate above,
Stems up to 10 cm long, 1.5–3.5 mm wide, complanate entire below; costa none or weak, short and double;
28 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 13. Isopterygium affusum. A. Habit. B. Enlargement of portion of stem. C. Leaves. D. Apical leaf cells. E. Median leaf cells. F, G. Alar
cells. H. Operculate capsule (wet). Illustrations A–G drawn from Spruce 1059 (NY, lectotype); H from Stanley & Molina 4511 (F).

median cells smooth, 66–118 × 5–7 µm; alar cells weakly Distribution and ecology: Honduras, Guadeloupe,
differentiated, rectangular or rarely quadrate, in small Venezuela, and Brazil (Figures 12, 14, 15); an aquatic spe-
groups with 1–2 cells on margins. Autoicous or rarely di- cies on rocks in and beside rivers and streams, near water-
oicous. Setae brown to reddish brown, 1.5–2.2 cm long; falls, or occasionally at bases of tufts of grasses and sedges
capsules inclined, 1.0–1.2 mm long, ovoid to ellipsoid, in marshes, occurring at altitudes of 430–1460 m.
somewhat contracted below the mouth when dry; oper- Discussion: The aquatic habit helps to identify
culum obliquely short-rostrate; peristome teeth up to this species since the other Latin American taxa, except
0.2 mm long. Spores only seen immature. Isopterygium acutifolium, are terrestrial. Its distinguishing
 number 93 • 29

FIGURE 14. Distribution of Isopterygium affusum (▲) and I. subbrevisetum (●) in Central America.

morphological features are the large plants, stems to 10 cm 1350 m, Standley & Molina R. 4511 (F). WINDWARD IS-
long, 1.5–3.5 mm wide, leaves ovate-lanceolate to ovate, LANDS. Guadeloupe: Known only from type collection of
symmetric to asymmetric, acuminate, flaccid, close, erect- I. herminieri. VENEZUELA. Bolívar: Morichal cercano al
spreading, strongly wrinkled and contorted when dry, conuco de Odremán, Santa Elena, Gran Sabana, Tamayo
1–3 mm long, alar regions weakly differentiated, 1–2 rectan- 3068 (FH, US). BRAZIL. Amazonas: Taruma Alta, 5 km
gular or rarely quadrate cells on margins, setae 1.5–2.2 cm N of Manaus-Itacoatiara road, near Manaus, Griffin 283-A
long, capsules rarely produced, ovoid to ellipsoid, inclined, (CANM, F, MEXU, MO, NY, SP); between Manaus and Sao
contracted below the mouth when dry, 1.0–1.2 mm long. Gabriel, along shores of Rio Curicuriari and Igarapé Branco
Iwatsuki and Crosby (1979) considered this species (Rio Cariua) from Rio Curicuriari to Cachoeira de Bôto (Ca-
synonymous with I. tenerifolium. However, the aquatic choeira Piraiauara), 0°20'S, 66°55'W, Buck 2549 (CANM,
habitat of I. affusum and its larger leaves, often up to 3 NY). Goiás: Region of the Chapada dos Veadeiros, 4 km N
mm long, and shorter setae, 1.5–2.2 cm long, distinguish of Veadeiros, Dawson 14745 (CANM); region of the south-
these two closely related species. The differences are rather ern Serra Dourada, 20 km E of Formoso, Dawson 14888
minor, and it is possible that additional specimens of the (CANM). Mato Grosso: Serra do Roncador, ~86 km N of
two species may, indeed, prove that they are synonymous. Xavantina, 550 m, Irwin et al. 16370 (NY). Minas Gerais:
Specimens examined: HONDURAS. Francisco Mpio. Santa Bárbara, Serra do Espinhaço, Parque Natural
Morazán: El Quebracho, above El Zamorano, ~950 m, do Caraça, trail from Santuário do Caraça to Pico da Car-
Standley 353 (F); region of Agua Amarilla, above El Zamo- apuça, 1300–1460 m, 20°02'S, 43°30'W, Buck 26686 (NY);
rano, ~780 m, Standley & Williams 13 (F), 900–1100 m, between Serro and Datas, Diamantina, Frahm 1421, 1499
Standley et al. 5047 (BM, F), 5044, 5050 (F); near Joya (MO); ~15 km SE of Datas, Vital 7568 (CANM); National
Grande, on road from El Zamorano to Suyapa, 1200– Park Serra de Itatiaia, 1950 m, Schäfer-Vervimp & Vervimp
30 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 15. Distribution of Isopterygium tenerifolium (), I. jamaicense (▲), I. subbrevisetum (●) and I. affusum (∆) in the Caribbean.

9571 (NY). Pará: Serra do Cachimbo, Base Aérea do tina. Chaco, between Sn. Andrés and Orán, 17 Sep 1873, Lorentz s.n. (lecto-
Cachimbo and vicinity, along bank of Rio Braço Norte, ~20 type: M!, designated by Ireland, 1991a; isolectotypes: H!, JE!).
km N of border of Mato Grosso, 430–480 m, Reese 16169
(MICH, NY, US), 16193 (MICH), 16197, 16418 (MICH, Taxicaulis saprophilus Müll. Hal., Hedwigia 36: 115. 1897; Isopterygium
NY). Santa Catarina: Serra Geral, Ule 1197 (H). São Paulo: saprophilum (Müll. Hal.) Paris, Index Bryol. Suppl. 220. 1900. Type: Ar-
Icem County, Vital DV-1826 (CANM). São Paulo: Mpio. gentina. Campochico, Aug 1873, Lorentz s.n. (lectotype: H!, designated by
Guapiara, Serra Paranapiacaba, Fazenda Intervales, ~800 m, Ireland, 1991a).
~24°16'S, 48°25'W, Vital & Buck 20436 (NY).
Plants in dense mats, light green to yellowish green.
3. Isopterygium byssobolax Stems up to 1.3 cm long, 0.5–1.0 mm wide, simple or ir-
regularly and freely branched. Asexual reproductive bod-
FIGURE 16 ies lacking. Leaves 0.4–0.8 mm long, 0.2–0.3 mm wide,
somewhat rigid, close, concave, erect to slightly spread-
Isopterygium byssobolax (Müll. Hal.) Paris, Index Bryol. ing at tips, smooth, somewhat contorted when dry, ovate
Suppl. 218. 1900. Taxicaulis byssobolax Müll. Hal., to ovate-lanceolate, symmetric, acuminate; margins erect,
Hedwigia 36: 114. 1897. Type: Argentina. Tucuma- entire to minutely serrulate; costa none or weak, short and
nensis, Siambón near Tucumán, Sierra de Tucumán, double; median cells 28–52 × 5–7 µm, smooth; alar cells
1873, Lorentz s.n. (lectotype: JE!, designated by Ireland, strongly differentiated, short-rectangular to quadrate or
1991a); Cuesto de Siambón, Lorentz s.n. (syntype: JE!). transversely elongate, in 2–4 rows with 4–10 cells on mar-
gins. Autoicous. Setae 0.6–1.0 cm long, yellow to reddish
Taxicaulis pyrrhopus Müll. Hal., Hedwigia 36: 116. 1897; Isopterygium brown; capsules 0.5–1.0 mm long, inclined to horizontal,
pyrrhopum (Müll. Hal.) Paris, Index Bryol. Suppl. 220. 1900. Type: Argen- ovoid to ellipsoid, contracted below the mouth when dry;
 number 93 • 31

FIGURE 16. Isopterygium byssobolax. A. Habit. B. Enlargement of portion of stem. C. Leaves. D. Apical leaf cells. E. Median leaf cells. F. Alar
cells. G. Inoperculate capsules (wet on left, dry on right). All illustrations drawn from Lorentz s.n. (JE, lectotype).

operculum short-rostrate; peristome teeth up to 0.2 mm can species (I. subbrevisetum is the other). The small stems
long. Spores 7–9 µm in diameter. reach only 1.3 cm long, 0.5–1.0 mm wide, the minute leaves
Distribution and ecology: Brazil, Bolivia, Para- are ovate to ovate-lanceolate, symmetric, acuminate, rigid,
guay, and Argentina (Figure 12); usually in forests on rot- close, concave, less than 1 mm long, with distinct alar re-
ten wood and tree bases, sometimes on soil and rock, at gions of short-rectangular to quadrate or transversely elon-
30–1180 m. gate cells in 2–4 rows with 4–10 cells on margins, setae
Discussion: Isopterygium byssobolax is recognized 0.6–1.0 cm long, capsules ovoid to ellipsoid, contracted
in part because it is one of the smallest of the Latin Ameri- below the mouth when dry, 0.5–1.0 mm long.
32 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Specimens examined: BRAZIL. Espírito Santo: 4. Isopterygium jamaicense

Domingos Martins, Reserva Florestal Pedra Azul, 1180
m, Schäfer-Verwimp & Verwimp 10078 (CANM). FIGURE 17
Mato Grosso: Palmeiras, Serra do Chapada, Lindman
B413 (BM). Paraná: Mpio. Ponta Grossa, Rio Tibagi, Isopterygium jamaicense (E. B. Bartram) W. R. Buck,
Kummrow 02990 (NY). Rio Grande do Sul: São Fran- Brittonia 36: 88. 1984a. Platygyriella jamaicensis
cisco de Paula, Casuza Ferreira, 800 m, Grazziotin et E. B. Bartram, J. Wash. Acad. Sci. 26: 14. 1936. Type:
al. 3661 (NY); Vila Gonzaga, São Leopoldo, Sehnem 77 Jamaica. Newmarket, Orcutt 7263 (holotype: FH!; iso-
(TENN). Rondônia: Ridges of Serra dos Pacaás Novos, type: CANM!).
along Rio Pacaás Novos, ~400 m, Reese 13550, 13553
(NY). Santa Catarina: Mpio. de Lajes, 27°48'S, 50°21'W, Plants in thin to dense mats, whitish green to
Vital 9397 (CANM, SP). São Paulo: Itapecirica, near Barra yellow-green. Stems up to 2 cm long, 1–2 mm wide,
Mansa, ~1000 m, Schiffner 1572 (BM); Parque Estadual complanate-foliate, simple or irregularly branched. Asex-
das Fontes do Ipiranga, São Paulo city, Yano 269 (MO). ual reproductive bodies lacking. Leaves 0.7–1.5 mm long,
BOLIVIA. Chuquisaca: Prov. Luis Calvo, “Inca Huasi 2,” 0.3–0.5 mm wide, flaccid, close, complanate, erect to erect-
W slope of Serrania Inca Huasi, 7 km NNE of Muyu- spreading, smooth, ovate to ovate-lanceolate, usually sym-
pampa (Vaca Guzman), 19°50'S, 63°43'W, Lewis 84- metric, acuminate; margins plane, entire or serrulate nearly
1050 (LPB). Cochabamba: Prov. Chapare, “Puerto San to the base; costa none or rarely weak, short and double;
Francisco,” between Puerto San Francisco and the alliga- median cells 52–85 × 5–9 µm, smooth; alar cells rectangu-
tor ranch near Río Salbasuma, 16°41'S, 65°10'W, Lewis lar to quadrate, in small groups with 2–4 cells along the
83-1346 (LPB). Santa Cruz: Prov. Chiquitos, “Río Nativi- margins, the cells sometimes inflated. Autoicous. Setae yel-
dad,” 17 km S and 15 km W from San José de Chiquitos, low to reddish brown, 0.5–0.8 cm long; capsules 0.5–1.3
17°53'S, 60°53'W, Lewis 85-1139 (LPB); Prov. Cordil- mm long, erect to somewhat inclined, straight, cylindri-
lera, 28 km SW of Abapo, 19°06'S, 63°32'W, Lewis 84- cal to somewhat ovoid, not or slightly contracted below
1173 (NY). Prov. Florida, Canton Samaipata, Laguna the mouth when dry; operculum short-rostrate; peristome
Volcán, above Bermejo, 1150 m, 18°07'S, 63°38'W, teeth up to 0.3 mm long, endostome segments linear, cilia
Churchill et al. 20893 (MO); Tarija: Prov. Acre, along lacking. Spores 14–24 µm in diameter.
trail from Communidad Salado to Cambari, 3.5 km N of Distribution and ecology: Mexico, Guatemala,
YPFB Campamento Conchas and 53 km N of Bermejo, Panama, and Jamaica (Figures 15, 18, 22); in humid forests,
22°16'S, 64°24'W, Lewis 84-2442, 84-2448 (LPB); ridge on bamboo stems, and on moist soil, reported once on old
N of Tunel, 2 km E of Comunidad Emborozu, 55 km NW thatched roof, at low altitudes from near sea level to 950 m.
of Bermejo, 22°17'S, 64°31'W, Lewis 84-2368 (LPB); Discussion: This species can be distinguished with
headwater area of Quebrada Toro, 1 km SE of YPFB sporophytes but impossible to distinguish from others, like
Campo Toro, 11 km N of Bermejo, 22°38'S, 64°19'W, Isopterygium tenerum or I. tenerifolium, when sterile. It
Lewis 84-2424 (LPB); slope of Cerro Nogal, parallel to is the only Latin American species with a usually cylindri-
and 0.5 km N of Quebrada Nogal, 2–5 km NW of Com- cal, erect to inclined capsule that is not or little contracted
munidad La Mamora, 22°09'S, 64°41'W, Lewis 84-2169 under the mouth when dry. The gametophyte is very similar
(LPB); S 15°W of summit of Cerro La Puñalada on slopes to I. tenerum, but that species has asymmetric leaves and
between Río Bermejo and summit, 6 km SW of Commu- sometimes has asexual reproductive bodies.
nidad La Mamora, 22°12'S, 64°37'W, Lewis 84-2269 The name Isopterygium miradoricum (Müll. Hal.)
(LPB). PARAGUAY. Caaguazú: Coronel Oviedo, Barrio A. Jaeger, described in 1874 from Mirador, Mexico, has
Azucena, 25°25'S, 56°27'W, Bordas 238 (NY). Paraguarí: been used previously for plants that seem to fit the descrip-
Macizo Acahay, 500 m, 25°54'S, 57°09'W, Zardini & tion and type of I. jamaicense (Ireland, 1984). However, the
Cuevas 5397 (NY); Cordillera de Ybytyruzú, Cerro Peró, type of I. miradoricum could not be located, and so it is im-
25°55'S, 56°15'W, Zardini & Velásquez 9377 (NY). AR- possible to be certain of the concept of I. miradoricum. The
GENTINA. Corrientes: Dept. Mercedes: Macrosistema type description states that the capsules are inclined to nod-
Iberá, Estancia Rincón del Diablo, 28°44'S, 58°02'W, ding, which does not fit the capsules in the specimens identi-
Arbo et al. 8008 (NY). Jujuy: Arroyo de los Matos ad fied as I. miradoricum by various bryologists that the senior
Sierra Santa Barbara, Fries 43 (H, MO). Tucumán: Que- author examined. If the type of I. miradoricum is identical to
brada de Lules, Garolera 6971 (FH). I. jamaicense, the former name has priority over the latter.
 number 93 • 33

FIGURE 17. Isopterygium jamaicense. A. Habit. B. Enlargement of portion of stem. C. Leaves. D. Apical leaf cells. E. Median leaf cells. F. Alar
cells. G. Operculate and inoperculate capsules (dry). All illustrations drawn from Santos 3854 (MICH).

Pterogonidium pulchellum (Hook.) Müll. Hal. in those of I. jamaicense, have a single rather than a double
Broth. of the Sematophyllaceae is often confused with peristome like all species of Isopterygium.
Isopterygium jamaicense. The gametophytes of the two Specimens examined: MEXICO. Oaxaca: Tuxte-
are strikingly similar except Pterogonidium is a smaller pec, El Cerro de Palacios, NE side of Chiltepec, Santos
plant with leaves that are seldom spreading as much as 3854 (CANM, F, MICH, TENN). Puebla: Río Tonta,
in I. jamaicense. Fortunately, Pterogonidium usually pro- Paseo Asihualt, 150 m, Sharp 1326a (CANM). Tabasco:
duces sporophytes, and the erect capsules, which are like (no other locality information) Gilly & Hernandez 307
34 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 18. Distribution of Isopterygium tenerifolium (▲), I. tenerum (●), and I. jamaicense () in Mexico.

(CANM, MICH). GUATEMALA. Alta Verapaz: Cu- Isopterygium pusillum Renauld & Cardot, Bull. Soc. Roy. Bot. Belgique
bilquitz, 350 m, Türckheim 6938 (BM, FH, H). Vicinity 41(1): 107. 1905. Type: Puerto Rico. 1900, Heller 4495 (holotype: PC!; iso-
of Puerto Barrios, near sea level, Maxon & Hay 3072, type: NY!).
3076, 3078 (US). PANAMA. Bocas del Toro: Vicinity of
Fortuna Dam, 4.5 km from Chiriquí Grande, 850–950 m, Isopterygium tenerrimum Renauld & Cardot, Bull. Soc. Roy. Bot. Belgique
08°55'N, 82°08'W, Allen 5620 (MO). JAMAICA. Port- 41(1): 107. 1905. nom. nud.
land: Soyo Falls, 1.3 km SW of Reach, 120 m, Proctor
37208 (NY); 11 km NW of Muirton, John Crow Mts., Plants in thin to dense mats, light green to yellowish
350 m, Buck 5662 (M, NY). green. Stems up to 0.8 cm long, 1.0–1.7 mm wide, irreg-
ularly and freely branched. Asexual reproductive bodies
5. Isopterygium subbrevisetum rarely present on stems, uniseriate, filamentous, multi-
cellular, simple or branched, green or brown, cells papil-
FIGURE 19 lose. Leaves 0.5–1.0 mm long, 0.2–0.3 mm wide, somewhat
rigid, distant, somewhat concave, wide-spreading to
Isopterygium subbrevisetum (Hampe) Broth., Nat. Pflan- squarrose, smooth, scarcely changed when dry, lanceolate,
zenfam. 1(3): 1081. 1908. Hypnum subbrevisetum symmetric, acuminate; margins plane or often erect below,
Hampe, Vidensk. Meddel. Dansk Naturhist. Foren. entire to minutely serrulate; costa none or sometimes
Kjøbenhavn ser. 3, 6: 165. 1875. Type: Brazil. Vicinity weak, short and double; median cells 66–99 × 5–7 µm,
of Rio de Janeiro, Glaziou 6356 (holotype: BM!; iso- smooth; alar cells quadrate to short-rectangular, in small
type: H!). groups with 2–4 cells on margins. Autoicous. Setae 0.5–
1.0 cm long, yellow to orange; capsules 0.5–1.0 mm long,
Taxicaulis subtenerrimus Müll. Hal., Hedwigia 37: 253. 1895; Isopterygium inclined, ovoid to ellipsoid, contracted below the mouth
subtenerrimum (Müll. Hal.) Paris, Index Bryol. Suppl. 221. 1900. Type: Ja- when dry; operculum short-rostrate; peristome teeth up to
maica. Bethabara, Wullschlägel s.n. (isotype: NY!). 0.2 mm long. Spores 9–14 µm in diameter.
 number 93 • 35

FIGURE 19. Isopterygium subbrevisetum. A. Habit. B. Enlargement of portion of stem. C. Leaves. D. Apical leaf cells. E. Median leaf cells. F.
Alar cells. G. Cross section of stem. H. Operculate and inoperculate capsules (dry). Illustrations A, B, G, H drawn from Steere 4602 (MO); C–F
from Glaziou 6356 (BM, holotype).

Distribution and ecology: West Indies, Belize, Discussion: The general aspect of this species is
Honduras, Nicaragua, Costa Rica, Panama, Venezuela, Su- striking because of its lanceolate, slightly concave, rigid,
rinam, French Guiana, Ecuador, Peru, and Brazil (Figures distant, wide-spreading to squarrose leaves. The narrow,
14, 15); on tree trunks, stumps, rotten logs, vines, humus, lanceolate leaves and the phyllotaxy separate Isopterygium
earth banks, and occasionally limestone and serpentine subbrevisetum from the other Latin American species. The
rock in humid or wet forests, occurring from 0 to 1940 m. plants are otherwise most similar to I. tenerum, but that
36 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

species has leaves somewhat broader (0.2–0.5 mm wide) Juan, 900 m, Hioram 11812 (MO); Loma San Juan,
and longer (0.7–1.2 mm long). 850 m, Hioram 11618 (MICH). Santiago: Slopes and sum-
The leaf morphology and the phyllotaxy of this spe- mit of El Yunque, near Baracoa, 305–610 m, Pollard &
cies may not seem very important until one considers the Palmer 104 (MO); El Yunque Mt., Baracoa, Underwood
North American plants of Isopterygium tenerum. Among & Earle 1054 (CANM); base of El Yunque Mt., Baracoa,
the hundreds of North American plants of Isopterygium Underwood & Earle 320 (CANM), 1289 (CANM, NY);
that the senior author studied (Ireland, 1969), none ap- 1–2 km W of La Gran Piedra, Shaw 5261 (NY). La Prenda,
proached the distinctive leaf morphology and the phyllo- Guantanamo, Hioram 18173 (MO). JAMAICA. Man-
taxy of I. subbrevisetum. chester: Marshall’s Pen, 4.8 km WNW of Mandeville,
Selected specimens examined: BELIZE. Cayo: 700 m, Crosby 13816 (NY); 4.3 km N of Plowden Hill,
Along Raspaculo Branch above confluence with Monkey 500 m, Crosby 13847 (NY); 1.6 km from Balaclava, Or-
Tail Branch, 480–720 m, Allen 18365 (MO). HONDU- cutt 816 (MICH); 0.8 km NE of Spur Tree, 790 m, Web-
RAS. Olancho: Río de Camote, 3 km N of La Union, 12 ster 5257a (CANM); Shooters Hill, 2.4–3.2 km S of
km below La Muralla, 800 m, 15°02'N, 86°42'W, Allen Mizpah, 430 m, Farr 831, 832 (CANM). Portland: E slope
12890 (MO). NICARAGUA. (Dept. Unknown) Sandy of John Crow Mts., ~2.4 km SW of Ecclesdown, 460 m,
Bay, 11 Dec 1920, Hamilton s.n. (NY). COSTA RICA. Webster 5179 (CANM); John Crow Mts., ~2 km SW of
Alajuela: San Pedro de San Ramon, Brenes 19, 20 (NY), Ecclesdown, 460–610 m, Webster 5588 (CANM); John
99 (FH, NY). Heredia: Finca La Selva, ~11 km S of Puerto Crow Mts., SW of Ecclesdown, 305–610 m, Welch 17542,
Viejo, Koch 4938 (MICH); Selva Biological Station, at 17707 (CANM); between 0.8 and 2.4 km SW of Eccles-
confluence of Río Sarapiquí and Río Puerto Viejo, 50–100 down, Powell & Welch 514 (CANM). St. Ann: Reynolds
m, 10°26'N, 84°01'W, Grayum 9634, 9990, 9999 (MO). Mines area, near Lydford, Howard & Proctor 14032
Limón: Along Río Tortuguero, ~3 km SW of village of (CANM). St. Andrew: Summit of Coopers Hill, 775 m,
Tortuguero, Steere CR-180, CR-195 (NY); 0.8 km SE of Farr 1441 (CANM); near Hermitage Dam, ~610 m, Proc-
Guapiles on Linnea Vieja, Diamantes Rubber Station, tor 17409 (CANM); Second Breakfast Spring, near Mt.
150 m, Svihla 3027a (FH). (Dept. Unknown) Vicinity of Airy, ~915 m, Farr 724 (CANM). St. Thomas: Mountain
La Palma, 1450–1550 m, Maxon 371 (NY). PANAMA. trail between House Hill and Cuna Cuna Gap, 550–725
Coclé: Near El Valle de Anton, ~550 m, Brako 8457 (NY). m, Maxon 8901 (BM); Cuna Cuna area, 610–760 m, Farr
Darién: Cerro de Garagará, Sambu basin, southern Darien, 1566 (CANM); Farm Hill, 1220 m, Proctor 9679 (CANM);
500–974 m, Pittier 5633 (NY). Panamá: Isla de Barro Col- N side of Cuna Cuna Gap, Britton 968 (MICH, NY);
orado, Arrocha 204 (NY), 204a (PMA); Barro Colorado Cuna Cuna Gap, 610 m, Britton 1251 (NY); Bath Foun-
Is., Panama Canal, Gatun Lake, Robbins 8 (CANM, FH); tain, ~150 m, Farr 1320 (CANM); Corn Puss Gap and
Quebrada Culebra, 70–75 m, Dodge & Allen 17087 vicinity, ~14 km N of Bath, on trail to Port Antonio, 550
(MO); El Llano–Carti road, 16 km from Interamerican m, Crosby 13653 (NY); Bath, trail N from Bath Fountain
Hwy., near El Llano, 330 m, Croat 33786 (MO). San Blas: Hotel along Sulphur River, ~90 m, Crosby 13615 (NY).
23 km from Interamerican Hwy. on El Llano–Carti Road, Trelawny: Windsor estate, ~120 m, Powell 335 (CANM);
Allen 4960 (H, MO, NY), 4968 (MEXU, MO); El Llano– Blue Mts., Hampstead, ~1115 m, Jäderholm 118 (H).
Carti road, 19 km from Interamerican Hwy., 350 m, de HAITI. Canape Verte, above Port-au-Prince, Mackaness
Nevers 4108 (MO). CUBA. Habana: Vicinity of Santiago 285 (MICH). Dept. de la Grand’anse, Massif de la Hotte,
de las Vegas, Baker 691 (NY). Las Villas: Santa Clara, along small stream entering Rivière Glace at junction of
S slope of Sierra de los Helechales, Banao Mts., León & road from Beaumont to Camp Perrin, 41 km S of Roseaux,
Clement 5545 (NY); Santa Clara, Sierra de Cantu, León & 710 m, Buck 9060 (NY); Guimbi Galata, Mornes des
Clement 5476 (MICH, US); Santa Clara, Trinidad Mts., Commissaires, 1800 m, Holdridge 1800 (MICH). Nord:
Britton 4879 (CANM), 5380 (CANM, NY). Oriente: Si- Vicinity of Pilate, ~325 m, Leonard 9594 (CANM, FH,
erra Maestra, Loma del Gato, Clemente 14538 (NY); Si- US). Ouest: Above road NW of Forêt des Pins, Imshaug &
erra Maestra, Cordillera de la Gran Piedra, 1250 m, Lopéz Wetmore 22888 (CANM, MSC). Sud’est: Massif de la
68 (CANM); Sierra Maestra, Gran Piedra, near Santiago Selle, 5 km E of Seguin on road to Mare Rouge, 1780 m,
de Cuba, Imshaug 25063 (CANM, MICH); Moa region, Buck 9478 (NY); Osman, Morne la Selle, 1650 m, Hold-
banks of the Río Yagrumaje, SW of Punta Gorda, Webster ridge 3008a, 3009, 3014 (MICH). DOMINICAN RE-
810 (CANM); Sierra de Nipe, foothills at the Río Guaro, PUBLIC. Independencia: Sierra de Baoruco, 30.5 km S of
~10 km SW of Mayari, Webster 873 (CANM); Loma San Puerto Escondito, 1940 m, Buck 14661 (NY). Santo
 number 93 • 37

Domingo: Vicinity of Colonia Ramfis, 400–500 m, Allard (CANM, NY); Gagrs Mtn., 460–760 m, Shafer 825
16290 (NY, US); vicinity of Ciudad Trujillo, 0–25 m, Al- (CANM, NY). Tortola: N side of Tortola, Allen & Harter
lard 16116 (NY). Pedernales: Las Abejas, ~40 km N along 3332 (MO). WINDWARD ISLANDS. Martinique: Bois de
road from Cabo Rojo from junction of road from Oviedo la Capote, Stehle 4002 (NY). Grenada: Grand Etang,
to Pedernales, 1190 m, Reese 14959 (NY); 55 km N of 760 m, Howard 264 (FH). COLOMBIA. Nariño: Mpio.
port of Cabo Rojo on Alcoa road, 1100 m, Steere 22854 Tumaco, Resguardo Indigena de El Hojal, Alrededores,
(NY). La Vega: La Culata, 9 km NNW of Constanza, 245 m, 01°19'N, 78°30'W, Ramírez P. et al. 9808 (MO).
1280 m, Buck 5356 (NY); La Cumbra, road between Con- VENEZUELA. Bolívar: Cerro Sarisariñama, Sima Mayor,
stanza and Jarabacoa, 1220 m, Allard 16508a (NY); Loma 1020 m, Buck & Brewer 15602A, 15619B (NY). TRINI-
Compana, ~1036 m, Allard 18654 (NY); vicinity of Piedra DAD AND TOBAGO. Trinidad: Caura R. Valley, Britton
Blanca, 200–500 m, Allard 18000d (NY); between Loma et al, 1224 (NY). Siparia, 30 m, Simmonds 517 (CANM).
La Sal and Loma La Golondrina, 945 m, Zanoni et al. Tobago: Murray’s Land, Cardiff Road, Broadway 4292
19938A, 19938M (MO). Dajabon: Cañada Tirolis, 1 km (CANM). SURINAM. Brokopondo: Brownsberg, near
S of Villa Anacaona, 400 m, Buck 4837 (NY); along road trail to Witti Creek, ~500 m, Florschütz-de Waard & Ziel-
at Los Amaceyes, 565 m, Imshaug & Wetmore 3401 man 5036A (U); Brownsberg, 450 m, Gradstein 4670 (U).
(MSC, NY). PUERTO RICO. Sierra de Yabucoa, 500 m, Sipaliwini: Blanche Marie Valley, 04°44'N, 56°53'W,
Britton et al. 6313 (NY). Ponce Dist., above Villalba, Doña 100 m, Muñoz 98-26b (MO). FRENCH GUIANA. Com-
Juana Recreation Area, 800–1000 m, Buck 15984 (NY); mune de Saül, Mont Galbao, summit NW peak, ~700 m,
Guaraguao, Pagán 66 (MICH, MO). Mountain between 03°36'N, 53°16'W, Buck 33294, 33306 (NY); NE of Saül,
Guayama and Cayey, 700–900 m, Britton et al. 6590 summit S of Pic Matécho, 13 km N of Saut Nais, 590 m,
(NY). Near Mayagüez, Heller 4495 (F, MICH). Western Cremers 6291 (U). ECUADOR. Galapagos Iss., Isla Santa
end of Laguna Tortuguero, Britton & Brown 6855 (MO, Cruz, around Media Luna, 600–650 m, Gradstein &
NY). Vicinity of Mayagüez, Britton & Marble 610 (NY). Weber M-37 (F); Isabela, S slope of Volcán Alcedo, van
Santa Rosa, Jayuya, Pagán 299 (MICH, MO). La Torre- der Werff 1477 (U). PERU. Santa Elena, upper Amazon,
cilla, Barranquitas, Steere 4546 (FH, MICH, MO), 4588, Tingo Maria, Morrow 9584 (FH). BRAZIL. Acre: Mpio.
4602, 4625, 4648 (MICH, MO), 4664, 4667 (MICH). Sena Madureira, Basin of Rio Purus, Rio Macanã, Coloca-
Ridge above Jajome Alto, Cayey–Guayama Road, Steere ção Castanheira, 09°42'S, 69°07'W, Daly et al. 8134
5115 (MICH, MO). Ridge at Km 15, Maricao-Sabana (NY). Amazonas: Along the Rio Negro, along BR 307,
Grande Road, Steere 5778 (MICH, MO). Just E of Cidra, N from São Gabriel de Cachoeira to Cucuí, at Km 41,
Steere 6347 (MICH), 6379 (MICH, MO). Above Toro Buck 2598 (NY); vicinity of Manaus, along BR 174 at Km
Negro Reservoir, S of Jayuya, Steere 6878 (MICH, MO). 30, Buck et al. 1751 (CANM, NY); along the Rio Uatumã,
Mt. Guilarte, W of Adjuntae, Steere 7191 (MICH, MO). along road to Balbina Dam project from Manaus-Caracarai
Guavate Purchase Unit, Sierra de Cayey, Steere 4820 (FH, Road (BR 174), Buck 2662 (CANM, NY). Bahia: Mpio.
MICH), 4838, 4858 (MICH), 6754 (MICH, MO), 6762 Ilhéus, Centro de Pesquisas do Cacau, Km 22 on BR 415
(MICH). Mt. Cenote, Pagán 1 (MO), 91, 124 (MICH). between Ilhéus and Itabuna, 50 m, ~14°50'S, 39°13'W,
Arroyo de los Corchos, between Adjuntas and Jajuya, Vital & Buck 20190 (NY); Serra da Agua de Rega, ~24 km
800–900 m, Britton 5334a (MICH). Monte Torrecilla, N of Seabra, road to Agua de Rega, 1000 m, Irwin et al.
900–1100 m, Britton et al. 5696 (NY), 5566 (MICH). Río 31095 (CANM). Minas Gerais: Parque Nacional do Ca-
Juan Diego, N slope of Sierra de Luquillo, Steere 6957 paraó, Vale Verde, ~1200 m, 20°30'S, 41°40'W, Buck
(MICH). NW slopes of Sierra de Luquillo, above Verde, 26942 (NY); Vicosa, Mexia 4499-a (FH); Fazenda Bom
Steere 4987 (MICH). Between Cayey and Caguas, Britton Destino, Bandeira 277 (NY). Paraná: Mpio. Foz do Iguaçu,
5704 (NY). Cerro de la Punta, S of Jayuya, Steere 6229 Parque Nacional do Iguaçu, 100–200 m, 25°40'S,
(MICH, NY). Mt. El Toro, trail to Ciánaga Alta, Sierra de 54°25'W, Vital & Buck 11960 (NY). Rio de Janeiro:
Luquillo, Steere 6461 (MICH). Toro Negro Unit, Río Parque Nacional do Itatiaia, vicinity of Pousada do Ele-
Doña Juana, N of Villalba, Steere 6015 (MICH, NY, US). fante, ~22°25'S, 44°36'W, Vital & Buck 20040 (NY); Pet-
22.5 km S of San Juan, Heller 696 (MICH). Río Sabana, ropolis, Pedreira da Quitandinha, Bandeira 191 (NY).
Sierra de Luquillo, S from Luquillo, Steere 5190 (MICH). Roraima: Bonfim, Santiago 14332 (NY). Santa Catarina:
Above upper Río de Marcao, Steere 5666 (MICH). Vi- Without locality, Ule 970 (H); Matta Hoffman, Brusque,
eques Is., Pagán 101 (MICH). LEEWARD ISLANDS. 50 m, Reitz 3144 (NY). São Paulo: Mpio. Guapiara, Serra
Montserrat: Chauers Mtn., 610–760 m, Shafer 806 Paranapiacaba, Fazenda Intervales, slopes of Morro do
38 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Mirante, 800–1000 m, ~24°16'S, 48°25'W, Vital & Buck 6. Isopterygium subglobosum

20495 (NY); Mpio. Santo André, slopes of Paranapiacaba,
~850 m, 23°47'S, 47°17'W, Vital & Buck 20609 (NY);
FIGURE 20
Sierra de São Pedro, São Pedro, 680 m, Schäfer-Verwimp
& Verwimp 9526 (NY); near Alto da Serra, May 1901,
900 m, Schiffner (BM); Mairinque, ~75 km W of São Isopterygium subglobosum Herzog, Biblioth. Bot. 87: 15l.
Paulo, Porta do Sol, Schäfer-Verwimp 8268 (CANM); 1916. Type: Bolivia. Santa Rosa del Chapare, Herzog
Bauru, Vital 11125 (CANM). 2738 (holotype: JE!; isotypes: H!, M!).

FIGURE 20. Isopterygium subglobosum. A. Habit. B. Enlargement of portion of stem. C. Leaves. D. Apical leaf cells. E. Median leaf cells. F.
Alar cells. G. Operculate and inoperculate capsules (dry). All illustrations drawn from Herzog 2738 (JE, holotype).
 number 93 • 39

Plants in thin to dense mats, light green to yellow- Plagiothecium villae-ricae Besch., Mem. Soc. Sci. Nat. Cherbourg 21: 271.
ish green. Stems up to 2.5 cm long, 1.0–1.5 mm wide, 1877. Type: Paraguay. Guaira, Villa Rica, Balansa 1210 (isotype: NY!).
complanate-foliate, simple or irregularly and freely
branched. Asexual reproductive bodies lacking. Leaves Isopterygium longisetum Broth., Bih. Kongl. Svenska Vetensk.-Akad.
0.7–1.0 mm long, 0.3–0.5 mm wide, flaccid, close, con- Handl. 21 Afd. 3(3): 55. 1895; I. mosenii Broth. in Engl. & Prantl., Nat.
cave, wide-spreading to squarrose, smooth, somewhat Pflanzenfam. 1(3): 1082. 1908, nom. illeg. Type: Brazil. Minas Gerais:
contorted when dry, ovate to ovate-lanceolate, often Caldas, Mosén 471 (lectotype: H!, designated by Ireland, 1991a); near
curved and asymmetric, short- to long-acuminate; mar- Capivary, Mosén 439 (syntype: H!); Serra de Caldas, Mosén 469 (syn-
gins erect, serrulate above, entire or nearly so below; costa type: H!); São Paulo, between Campinas and Serra de Caracol, Mosén
none; median cells 52–108 × 7–9 µm, smooth; alar cells 470 (syntype: H!).
rectangular, in small groups with 1–2 cells on margins.
Autoicous. Setae 1.4–2.2 cm long, yellowish red to red- Taxicaulis stigmocarpus Müll. Hal., Nuovo Giorn. Bot. Ital. 4: 150.
dish; capsules 0.5–1.2 mm long, inclined, subglobose, not 1897; Isopterygium stigmocarpum (Müll. Hal.) Paris, Index Bryol.
contracted below the mouth when dry; operculum high- Suppl. 220. 1900. Type: Bolivia. Province Cochambamba, near Choque-
conic to short-rostrate; peristome teeth up to 0.3 mm long. camata, Germain 1173b (lectotype: M!, designated by Ireland, 1991a;
Spores 9–12 µm in diameter. isolectotype: H!).
Distribution and ecology: Known from only the
type locality in Bolivia (Figure 12), occurring on decaying Taxicaulis excelsipes Müll. Hal., Hedwigia 37: 253. 1898; Isopterygium ex-
leaves at margin of creek at ~1700 m. celsipes (Müll. Hal.) Paris, Index Bryol. Suppl. 219. 1900. Type: Jamaica.
Discussion: This species, like the previous one, is Near Kingston, 1897, Hansen s.n. (isotype: H!).
impossible to distinguish from several of the other Latin
American Isopterygium species when capsules are not pres- Plagiothecium meteoriaceum Müll. Hal., Bull. Herb. Boissier 6: 120. 1898;
ent. The subglobose capsules that are not contracted under Isopterygium meteoriaceum (Müll. Hal.) Paris, Index Bryol. Suppl. 220.
the mouth when dry are very distinctive, but this may possi- 1900. Type: Brazil. Serra Itatiaia, Ule 1884 (isotype: H!).
bly represent only an environmental response to light, mois-
ture, etc., with no genetic basis. We have elected to retain Plagiothecium proximum Müll. Hal., Hedwigia 38(Beibl.): 59. 1899.
Isopterygium subglobosum as a separate species until more nom. nud.
collections become available for comparison. The gameto-
phytes of this species are very similar to I. tenerum. Plagiothecium paludigenum Müll. Hal., Hedwigia 40: 58. 1901; Isopteryg-
Specimen examined: BOLIVIA. Only the type ium paludigenum (Müll. Hal.) Broth. in Engl. & Prantl, Nat. Pflanzenfam.
specimens seen. 1(3): 1082. 1908. Type: Brazil. Santa Catarina: Tubarão, Ule 912 (lectotype:
H!, designated by Ireland, 1991a).
7. Isopterygium tenerifolium
Plagiothecium lonchopelmatum Müll. Hal., Hedwigia 40: 59. 1901; Isopte-
FIGURE 21 rygium lonchopelmatum (Müll. Hal.) Broth. in Engl. & Prantl, Nat. Pflan-
zenfam. 1(3): 1082. 1908. Type: Brazil. San Francisco Island, Province Santa
Catarina, Ule 511 (lectotype: H!, designated by Ireland, 1991a); Ule 509,
Isopterygium tenerifolium Mitt., J. Linn. Soc., Bot. 12: 513 (syntypes, H!).
499. 1869. Type: Brazil. Amazon River, Spruce 1060
(holotype: NY!; isotype: BM!). Plagiothecium restingae Müll. Hal., Hedwigia 40: 60. 1901; Isopterygium
restingae (Müll. Hal.) Broth. in Engl. & Prantl , Nat. Pflanzenfam. 1(3):
Plagiothecium inordinatum Mitt. ex Spruce, Cat. Musc. 16. 1867. nom. 1082. 1908. Type: Brazil. Rio de Janeiro: Restinga de Maua, Ule 2144 (lecto-
nud. type: H!, designated by Ireland, 1991a; isolectotypes: FH!, M!).

Ectropothecium longisetum Schimp. ex Besch., Ann. Sci. Nat., Bot. sér. 6, Plagiothecium restingae var. tenue Müll. Hal., Hedwigia 40: 60. 1901. Type:
3: 258. 1876; Isopterygium longisetum (Schimp. ex Besch.) Broth. in Engl. Brazil. Nova Friburgo, in Capoeira, Ule, Bryotheca brasiliensis 233 (lecto-
& Prantl, Nat. Pflanzenfam. 1(3): 1082. 1908, hom. illeg., non Broth. 1895; type: H!, here designated; isolectotypes, BM!, NY!).
Isopterygium altisetum H. A. Crum & Steere, Bryologist 59: 254. 1956,
nom. nov. Type: Guadeloupe. Chemin de la Soufrière, Husnot exsiccata no. Plagiothecium jamaicense Müll. Hal., Hedwigia 40: 61. 1901. Type: Jamaica.
169 (holotype: BM!; isotypes: M!, NY!). 1897, Hansen s.n. (isotype: NY!).
40 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 21. Isopterygium tenerifolium. A. Habit. B. Enlargement of portion of stem. C. Leaves. D. Apical leaf cells. E. Median leaf cells. F. Alar
cells. G. Operculate capsule (wet). H, I. Pseudoparaphyllia. Illustrations A, B, G drawn from Stevens et al. 25048 (MO); C–F, H, I from Spruce
1060 (NY, holotype).

Isopterygium fecundum Renauld & Cardot, Bull. Soc. Roy. Bot. Belgique Isopterygium vagans Herzog, Biblioth. Bot. 87: 152. 1916. Type: Bolivia. An
41(1): 140. 1905. Type: Costa Rica. Potrero del Alto, volcán de Poás, Pittier Wegrändern bei San Miguelito, Herzog 2765 (holotype: JE!; isotype: H!).
& Durand, Plantae costaricenses exsiccatae 5735 (isotypes: FH!, US!).
Plants in thin, loose mats, yellow-green to green. Stems
Isopterygium manaosense Broth., Hedwigia 45: 286. 1906. Type: Brazil. up to 4 cm long, 2–3 mm wide, complanate-foliate, simple
Estado de Amazonas, Manaos, Rio Negro, Ule, Bryotheca brasiliensis 280 or irregularly and freely branched. Asexual reproductive
(lectotype: H!, designated by Ireland, 1991a); Ule 2331 (syntypes, BM!, H!). bodies lacking. Leaves 1.0–1.5 mm long, 0.4–0.7 mm
 number 93 • 41

FIGURE 22. Distribution of Isopterygium tenerum (●), I. tenerifolium (▲), and I. jamaicense () in Central America.

wide, flaccid, close, complanate, wide-spreading to squar- Discussion: Isopterygium tenerifolium is best rec-
rose, smooth when wet, usually wrinkled and contorted ognized by its large plants with stems to 4 cm long, 2–3 mm
when dry, ovate-lanceolate to ovate, often curved and wide, and leaves 1.0–1.5 mm long, ovate-lanceolate to
asymmetric, acuminate; margins plane to erect through- ovate, often curved and asymmetric, acuminate, flaccid,
out, serrate to serrulate above, serrulate below; costa none close, complanate, usually wrinkled and contorted when
or weak, short and double; median cells 71–141 × 5–7 µm, dry, with poorly developed alar cells, 1–2 short-rectangular
smooth; alar cells short-rectangular to quadrate, in small to quadrate cells on the margins, setae, 2–3 cm long, rarely
groups with 1–2 cells on margins. Autoicous. Setae (1.5–) up to 4 cm, and capsules ovoid to ellipsoid, horizontal to
2–3(–4) cm long, brown to reddish brown; capsules pendulous, contracted below the mouth when dry, 1.0–
1.0–1.5 mm long, horizontal to pendulous, rarely erect, 1.5 mm long.
ovoid to ellipsoid, contracted below the mouth when dry; The plant’s large size and especially the long setae,
operculum obliquely short-rostrate; peristome teeth up to longer than any of the other Latin American species of
0.2 mm long. Spores 9–14 µm in diameter. Isopterygium, are obvious features that help identify this
Distribution and ecology: West Indies, southern species.
Mexico to Panama, and from Colombia and Venezuela Selected specimens examined: MEXICO. Chiapas:
to northern Argentina (Figures 15, 18, 22, 23), also from Mpio. Angel Albino Corzo, 9 km S de Finca Liquidambar,
the southeastern United States; on tree trunks, rotten 15°42'N, 92°45'W, Delgadillo M. 4676, 4678, 4679b,
stumps, woody debris and humus in forests, sometimes on 4726, 4728a (MEXU); Cárdenas 4163, 4189 (MEXU).
soil banks and wet cliffs along rivers and near waterfalls, HONDURAS. Cortés: Cusuco Natl. Park, along Río Cu-
rarely in moist grasslands, at low to high elevations from suco, ~22 km W of San Pedro Sula and 16 km S of Cuyamel,
160 to 3000 m. 15°30'N, 88°14'W, 1840–1900 m, Allen 14392 (MO). El
42 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 23. Distribution of Isopterygium tenerum (●) and I. tenerifolium () in South America.

Paraíso: Between Las Mesas and Ojo de Agua, 700 m, Mor- 950 m, Allen 5635 (MO). Canal Area: Barro Colorado Is.,
ton 6981, 6982 (US). Francisco Morazán: Region of Agua Jan–Feb 1938, Willis s.n. (MICH). CUBA. Habana: Bata-
Amarilla, above El Zamorano, 900–1100 m, Standley et bano, Baker & Wilson 2391 (H). Pinar del Río: Herradura,
al. 5050 (FH). COSTA RICA. Alajuela: Volcán Poás, 2500 Ekman 10749 (BM, NY). San Mateo, near Pinar del Río,
m, Valerio 285 (FH). Limón: ~2 km S and 1 km E of Río Wright 160 (CANM, FH, MICH, NY, TENN). JAMAICA.
Colorado, Stevens et al. 25048 (MO). PANAMA. Bocas del Clarendon: Mason River Savanna, 4.4 km NW of Kellits,
Toro: Vicinity of Fortuna Dam, 4.5 km along pipeline road 700 m, Crosby 3118 (NY). DOMINICAN REPUBLIC.
leaving Chiriqui Grande road at Continental Divide, 850– Dajabón: 14.7 km desde el Parque Central del Pueblo de
 number 93 • 43

Loma de Cabrera en el camino a Capotillo y Loma de Alto Grande on São Paulo Railway, 700 m, Schiffner 650 (FH,
de la Paloma, 490 m, Zanoni & Mejía 17989A (NY). Pera- H). BOLIVIA. Santa Cruz: Prov. Chiquitos, “Cerro Tata-
via: La Nevera area, 47 km S of Constanza, 2070–2100 raqui,” 13 km NE of Roboré, 18°16'S, 59°39'W, Lewis
m, Buck 5247 (NY). PUERTO RICO. Vicinity of Utuado, 85-1307 (LPB); Prov. Manuel M. Caballero, Serranía de
Britton 5139a (NY). Cerro de la Punta, Cordillera Cen- Siberia, ~15 km E de Siberia, carretera a Comparapa, 2460
tral, S of Jayuya, Steere 6212 (MICH, MO, NY, US), 6238 m, 17°49'S, 64°40'W, Churchill 22482 (MO, NY). Also
(MICH, MO). South shore of Laguna Tortuguero, near known from type collections of I. stigmocarpum & I. va-
Manati, Steere 6777 (MICH, NY, US). Vega Baha, Heller gans. PARAGUAY. Santo Antonio, Colonia “Elis,” Lind-
1338 (NY). WINDWARD ISLANDS. Guadeloupe: With- man B231 (NY). ARGENTINA. Corrientes: Dept. Santo
out locality, Duss 124 (H, NY), 1045, 1046, 1092, 1260 Tome: Laguna Luna, 28°02'S, 56°49'W, Arbo et al. 8476
(NY). Dominica: Grand Souffriere, on lip of crater, Elliott (NY). Posados, Misiones, Eckman 2096 (FH).
1841, 1843 (BM). Martinique: Without locality, Duss 1
(H, NY), 25, 332 (NY). COLOMBIA. Valle: La Cumbre, 8. Isopterygium tenerum
1800–2200 m, Killip 5572 (NY). Santander: Vicinity of Las
Vegas, 2600–3000 m, Killip & Smith 16026 (NY). VEN- FIGURE 24
EZUELA. Bolívar: Dist. Roscio, 13 km al Noreste de Santa
Elena de Uairén, 900 m, Steyermark & Liesner 127566 Isopterygium tenerum (Sw.) Mitt., J. Linn. Soc., Bot.12:
(MO); vicinity of Canaima and the Río Carrao, A. J. & 499. 1869. Hypnum tenerum Sw., Fl. Ind. Occid. 3:
E. B. Sharp F-7629 (TENN). Tachira: 12 km SW of Punta 1817. 1806; Isothecium tenerum (Sw.) Brid., Bryol.
de Piedra, toward Sacramento, 175 m, Steyermark & Rabe Univ. 2: 385. 1827. Type: Jamaica. Swartz 2719 (holo-
96615 (US). Yaracuy: Dept. Bruzual, Mpio. Campoelias, type: S!; isotypes: BM!, C!).
1300–1550 m, Trujillo 16461 (MO). GUYANA. Without
locality, Parker s.n. (NY). SURINAM. Sabanpassie, Teunis- Hypnum micans Sw., Adnot. Bot. 175. 1829; Rhynchostegium micans
sen & Wildschut 11931 (U). FRENCH GUIANA. Canton (Sw.) Austin, Bot. Gaz. (Crawfordville) 1: 30. 1875; Isopterygium micans
de Approuague-Kaw, ~1.5 km SSW of Pic Matécho, 160 m, (Sw.) Kindb., Enum. Bryin. Exot. 21. 1888; Plagiothecium micans (Sw.)
03°44'N, 53°02'W, Buck 37938 (NY). ECUADOR. Ori- Paris, Index Bryol. 963. 1896. Type: North America [Pennsylvania, Lan-
ente: Without locality, Jun 1910, 1800 m, Allioni s.n. (H). caster Co.?]. Muhlenberg 166 (lectotype, selected here, PH!; no material
BRAZIL. Amazonas: Manaus, Rio Negro, May 1902, Ule found at S).
s.n. (M); Taruma-Fälle, in Quellmulden, ~60 m, 02°53'S,
60°06'W, Schäfer-Verwimp 9803 (NY). Bahia: Mpio. Rio Hypnum splendidulum Hornsch., Fl. Bras. 1(2): 77. 1840; Isopterygium
de Contas, Pico das Almas, 1120 m, 13°00'S, 14°53'W, splendidulum (Hornsch.) Broth. in Engl. & Prantl, Nat. Pflanzenfam. 1(3):
Harley et al. 25965 (NY). Salvador, Bairro of Itapua, vicin- 1081. 1908. Type: Brazil. In Serra dos Orgãos, Feb, Beyrich s.n. (lectotype:
ity of airport, Boom et al. 886, 895 (NY); between Ibirapi- BM!, designated by Ireland, 1991a).
tanga and Ubaitaba, 200 m, Schäfer-Verwimp & Verwimp
8753 (NY). Goiás: Region of southern Serra Dourada, Hypnum fulvum Hook. & Wilson in Drumm., Musci Amer., S. States 110.
Dawson 14888 (FH). Minas Gerais: Serra da Mantiqueira, 1841, hom. illeg.; Plagiothecium fulvum A. Jaeger, Ber. Thätigk. St. Gal-
Delfim Moreira, in Quellmoor, 1330 m, 22°33'S, 45°15'W, lischen Naturwiss. Ges. 1876–77: 450. 1878 (new name); Isopterygium ful-
1330 m, Schäfer-Verwimp & Verwimp 14946 (NY); Serra vum (A. Jaeger) Kindb., Canad. Rec. Sci. 6(2): 72. 1894; Plagiothecium mi-
de Caldas, 15 Sep 1873, Mosén s.n. (H). Parque Nacional cans var. fulvum (A. Jaeger) Paris, Index Bryol. 963. 1896. Type: Louisiana.
do Itatiaia, vicinity of Brajo da Lapa, at Km 8 along entry Drummond, Musci Amer., S. States 110 (isotypes: NY!, WTU!).
road near border of Rio de Janeiro, 2120 m, ~22°22'S,
44°45'W, Vital & Buck 19660 (NY). Pará: Ponta Grossa, Hypnum brachyneuron Müll. Hal., Bot. Zeitung (Berlin) 3: 109. 1845; Isop-
Rio Tapajas, Swallen 6936 (FH, US). Rio Grande do Sul: terygium brachyneuron (Müll. Hal.) Mitt., J. Linn. Soc., Bot. 12: 498. 1869.
Vieira, nearly opposite Rio Grande, Lindman B130 (H, Type: Brazil. Santa Catarina: Sep–Oct 1864, Pabst s.n. (isotype: H!)
NY). Rondônia: Along Rio Pacaás Novos, between its con-
fluence with Rio Mamoré and the first rapids, Reese 13294 Hypnum curvicolle Müll. Hal., Syn. Musc. Frond. 2: 684. 1851, “curvicol-
(NY); Vilhena-Colorado road, 13 km N of Vilhena on BR- lum”; Isopterygium curvicolle (Müll. Hal.) Mitt., J. Linn. Soc., Bot. 12: 498.
364, Nelson 358 (NY). Santa Catarina: Tubarão, Ule 982 1869; Isopterygium curvicolle var. majus Broth. in E. Bauer, Verh. Zool.-Bot.
(H). São Paulo: Near Cerqueira-Cesar, 500 m, Kryptoga- Ges. Wien 55: 576, 578. 1905, Rev. Bryol. 32: 11. 1905. nom. nud. Type:
mae exsiccatae 2898, Schiffner (H, M, US); near Campo Brazil. Santa Catarina: Apr and May 1847, Pabst s.n. (isotypes: H!, JE!).
44 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Hypnum leucophyllum Hampe in Müll. Hal., Syn. Musc. Frond. 2: 280. Hypnum brachystelium Hampe, Vidensk. Meddel. Dansk Naturhist. Fo-
1851; Isopterygium leucophyllum (Hampe) Mitt., J. Linn. Soc., Bot. 12: ren. Kjøbenhavn ser. 4, 1: 139. 1879; Isopterygium brachystelium (Hampe)
499. 1869. Type: Chile. On trees, without specific locality and collector Kindb., Enum. Bryin. Exot. 100. 1891. Type: Brazil. Near Novo Friburgo,
(holotype: BM!). Beyrich s.n. (holotype: BM!).

Hypnum hospitans Hampe, Ann. Sci. Nat., Bot. sér. 5, 5: 313. 1866; Isopte- Hypnum citrinum Hampe, Vidensk. Meddel. Dansk Naturhist. Foren.
rygium hospitans (Hampe) A. Jaeger, Ber. Thätigk. St. Gallischen Naturwiss. Kjøbenhavn ser. 4, 1: 142. 1879; Isopterygium citrinum (Hampe) Kindb.,
Ges. 1876–77: 434. 1878. Type: Colombia. Tequendama, Aug 1863, Lindig Enum. Bryin. Exot. 100. 1891. Type: Brazil. Serra dos Orgãos, collector Bey-
s.n. (holotype: BM!; isotype: M!). rich? (holotype: BM!).

Hypnum cordovense Müll. Hal., Linnaea 38: 652. 1874; Isopterygium cor- Hypnum cacti Müll. Hal., Linnaea 42: 437. 1879; Isopterygium cacti (Müll.
dovense (Müll. Hal.) A. Jaeger, Ber. Thätigk. St. Gallischen Naturwiss. Ges. Hal.) Kindb., Enum. Bryin. Exot. 20. 1888. Type: Argentina. Concepción del
1876–77: 436. 1878. Type: Mexico. Cordova, Sep 1857, Mohr s.n. (isotypes: Uruguar, Quinta del Colegio, Apr 1876, Lorentz s.n. (isotypes: H!, JE!).
NY!, US!).
Hypnum eutrypherum Müll. Hal., Linnaea 42: 497. 1879; Isopterygium eu-
Hypnum leptomiton Müll. Hal., Linnaea 38: 652. 1874; Isopterygium trypherum (Müll. Hal.) Paris, Index Bryol. 707. 1897. Type: Venezuela. Near
leptomiton (Müll. Hal.) A. Jaeger, Ber. Thätigk. St. Gallischen Naturwiss. Colonian Tovar, Fendler 153 (isotypes: BM!, H!).
Ges. 1876–77: 436. 1878. Type: Guatemala. Bernoulli s.n. (isotypes:
BM!, H!). Hypnum restitutum Hampe, Vidensk. Meddel. Dansk Naturhist. Foren.
Kjøbenhavn ser. 4, 1: 141. 1879; Isopterygium restitutum (Hampe) Kindb.,
Hypnum laxum Hampe, Vidensk. Meddel. Dansk Naturhist. Foren. Enum. Bryin. Exot. 100. 1891. Type: Brazil. Vicinity of Rio de Janeiro,
Kjøbenhhavn ser. 3, 6: 166. 1875, hom. illeg. non P. Beauv. 1805; Ectro- Glaziou 7163 (lectotype: BM!, designated by Ireland, 1991a); Glaziou 7455
pothecium laxum A. Jaeger, Ber. Thätigk. St. Gallischen Naturwiss. Ges. (syntype: H!).
1877–78: 266. 1880 (new name); Isopterygium laxum (A. Jaeger) Broth.,
Bih. Kongl. Svenska Vetensk.-Akad. Handl. 21 Afd. 3(3): 56. 1895. Type: Hypnum chlorosum Hampe, Flora 64: 414. 1881; Isopterygium chlorosum
Brazil. Vicinity of Rio de Janeiro, Glaziou 7197 (holotype: BM!; isotypes: (Hampe) Paris, Index Bryol. 706. 1897. Type: Brazil. Near Rio de Janeiro,
H!, JE!). Glaziou 11733 (holotype: BM!; isotype: H!).

Hypnum divaricatulum Müll. Hal. ex Hampe, Vidensk. Meddel. Dansk Isopterygium subtenerum Besch., J. Bot. 5: 348. 1891. Type: Paraguay. Cor-
Naturhist. Foren. Kjøbenhavn ser. 3, 6: 167. 1875; Isopterygium divaricatu- dillère de Péribébui, Balansa 3690 (holotype: BM!; isotype: MICH!).
lum (Hampe) Broth. in Engl. & Prantl, Nat. Pflanzenfam. 1(3): 1081. 1908.
Type: Brazil. Vicinity of Rio de Janeiro, Glaziou 5611 (holotype: BM!; iso- Isopterygium guarapense Besch., J. Bot. 5: 349. 1891. Type: Paraguay.
types: H!, JE!, NY!). Guarapi, Balansa 3619 (possible holotype: BM!; isotype: H!).

Hypnum lamprophyllum Hampe, Vidensk. Meddel. Dansk Naturhist. Fo- Plagiothecium radicisetum Müll. Hal., Malpighia 10: 515. 1896; Isopter-
ren. Kjøbenhavn ser. 3, 6: 167. 1875, hom. illeg. non (Mitt.) Müll. Hal. ygium radicisetum (Müll. Hal.) Broth. in Engl. & Prantl, Nat. Pflanzenfam.
1874; Ectropothecium lamprophyllum A. Jaeger, Ber. Thätigk. St. Gal- 1(3): 1082. 1908. Type. Guyana. Georgetown, near Marshall Falls on the
lischen Naturwiss. Ges. 1877–78: 266. 1880 (new name); Isopterygium Mazaruni River, 1895, Quelch s.n. (isotype: BM!).
lamprophyllum (A. Jaeger) Broth. in Engl. & Prantl, Nat. Pflanzenfam.
l(3): 1081. 1908. Type: Brazil. Vicinity of Rio de Janeiro, Glaziou 6357 Taxicaulis adflatus Müll. Hal., Hedwigia 36: 115. 1897; Isopterygium ad-
(holotype: BM!; isotype: H!). flatum (Müll. Hal.) Paris, Index Bryol. Suppl. 217. 1900. Type: Argentina.
Tucumanensis, near Tucumán, Mar 1872, Lorentz s.n. (isotype: H!).
Ectropothecium clandestinum Ångstr., Öfvers. Förh. Kongl. Svenska
Vetensk.-Akad. 33(4): 43. 1876; Isopterygium clandestinum (Ångstr.) Taxicaulis cylindraceus Müll. Hal., Nuovo Giorn. Bot. Ital. n.s. 4: 151. 1897;
Broth. in Engl. & Prantl, Nat. Pflanzenfam. 1(3): 1081. 1908. Type: Bra- Isopterygium cylindraceum (Müll. Hal.) Paris, Index Bryol. Suppl. 218.
zil. Caldas, Jun 1854, G.A. Lindberg s.n. (isotypes: BM!, H!). 1900. Type: Bolivia. Cochabamba, near Choquecamata, Jun 1889, Germain
s.n. (isotype: H!).
Isopterygium streptopodium Besch., Ann. Sci. Nat., Bot. sér. 6, 3: 257. 1876.
Type: Guadeloupe. Chemin de la Soufrière, Husnot, exsiccata no. 184 (holo- Taxicaulis trichopelma Müll. Hal., Bull. Herb. Boissier 5: 210. 1897; Isopte-
type: BM!). rygium trichopelma (Müll. Hal.) Paris, Index Bryol. Suppl. 221. 1900. Type:
 number 93 • 45

Guatemala. Alta Vera Paz, Pansanala, Dec 1887, von Türckheim s.n. (holo- 1081. 1908. Type: Brazil. Santa Catarina: Campo de Jaguarone, Laguna, Ule
type: BM!). 780 (isotypes: H!, M!).

Taxicaulis weigeltii Müll. Hal., Hedwigia 37: 252. 1898, “weigelti”; Isop- Taxicaulis fabroniiformis Müll. Hal., Hedwigia 40: 69. 1901; Isopterygium
terygium weigeltii (Müll. Hal.) Broth., Nat. Pflanzenfam. 1(3): 1081. 1908. fabroniiforme (Müll. Hal.) Broth. in Engl. & Prantl, Nat. Pflanzenfam. 1(3):
Type: Surinam. Weigelt s.n. (possible isotype: H!). 1083. 1908, “fabroniaeforme.” Type: Brazil. Goyaz, near Mossamedes, Ule
1617 (isotypes: NY!, US!).
Taxicaulis rufisetulus Müll. Hal., Hedwigia 37: 253. 1898; Isopterygium
rufisetulum (Müll. Hal.) Paris, Index Bryol. Suppl. 220. 1900. Type: Cuba. Plagiothecium pseudotenerum Broth. & Kindb. ex Kindb., Rev. Bryol. 28:
Wright, coll. III, no. 96 (possible isotype: H!). 54. 1901; Isopterygium pseudotenerum (Broth. & Kindb.) Broth. in Engl. &
Prantl, Nat. Pflanzenfam. 1(3): 1082. 1908. Type: Colombia. Near Bonda,
Taxicaulis araneosetus Müll. Hal., Hedwigia 37: 255. 1898; Isopterygium vicinity of Santa Marta, Dec 1898, Baker s.n. (holotype: S!; isotype: MO!).
araneosetum (Müll. Hal.) Paris, Index Bryol. Suppl. 218. 1900, “arauco-
setum.” Type: Trinidad. Near Maroval, Jul 1847, Crüger s.n. (possible Isopterygium peruvianum Broth., Bot. Jahrb. Syst. 56 (Beibl. 123): 21. 1920.
isotype: H!). Type: Peru. Sandia Province, Churchusmayo, 7 Jul 1902, Weberbauer s.n.
(holotype: H!).
Isopterygium callochlorum Broth., Bih. Kongl. Svensk. Vetensk.-Akad.
Handl. 26 Afd. 3(7): 46. 1900. Type: Paraguay. Santo Antonio, colonia Isopterygium fernandezianum Broth. in Skottsb., Nat. Hist. Juan Fernan-
“Elisa,” Lindman 231b (holotype: H!). dez 2(12): 440. 1924. Type: Juan Fernandez Islands, Mas a Tierra, C. &
I. Skottsberg 479 (holotype: H!).
Microthamnium angustirete Broth., Bih. Kongl. Svensk. Vetensk.-Akad.
Handl. 26 Afd. 3(7): 48. 1900; Isopterygium angustirete (Broth.) Broth. Isopterygium fernandezianum var. longisetum Broth. in Skottsb., Nat.
in Engl. & Prantl, Nat. Pflanzenfam. 1(3): 1082. 1908. Type: Brazil. Rio Hist. Juan Fernandez 2(12): 440. 1924. Type: Juan Fernandez Islands, Mas
Grande do Sul: S. Leopoldo, Lindman 115 (holotype: H!). a Tierra, Valle Colonial, Quebrada seca, C. & I. Skottsberg 487 (lectotype:
H!, designated by Ireland, 1991a); Cordón Salsipuedes, C. & I. Skottsberg
Plagiothecium bromeliophilum Müll. Hal., Hedwigia 40: 59. 1901; Isopter- 480 (syntype, H!); Valle Anson, near Plazoleta, C. & I. Skottsberg 491
ygium bromeliophilum (Müll. Hal.) Broth. in Engl. & Prantl, Nat. Pflanzen- (syntype: H!).
fam. 1(3): 1082. 1908. Type: Brazil. Santa Catarina: Tubarão, near Concon-
haz, Ule 769 (isotype: H!). Isopterygium brachyneuroides Broth., Akad. Wiss. Wien, Math.-Naturwiss.
Kl., Denkschr. 83: 327. 1926. Type: Brazil. São Paulo: On island “Ilha Com-
Plagiothecium aurantiacum Müll. Hal., Hedwigia 40: 60. 1901; Isopteryg- prida,” near Iguape, Schiffner 362 (holotype: H!; isotype: BM!).
ium aurantiacum (Müll. Hal.) Broth. in Engl. & Prantl, Nat. Pflanzenfam.
1(3): 1082. 1908. Type: Brazil. Rio de Janeiro: Restinga de Maua, Ule 2145 Isopterygium diminutivum E. B. Bartram, J. Wash. Acad. Sci. 18: 581.
(isotypes: FH!, H!). 1928, “dimunitivum.” Type: Mexico. State of Jalisco, Túxpan, Mexía
1028a (holotype: US!; isotype: FH!).
Plagiothecium flaviusculum Müll. Hal., Hedwigia 40: 59. 1901; Isopteryg-
ium flaviusculum (Müll. Hal.) Broth. in Engl. & Prantl, Nat. Pflanzenfam. Plants in thin to dense mats, whitish green to yellow
1(3): 1082. 1908. Type: Brazil. Rio de Janeiro: Restinga de Maua, Ule 1941 -green. Stems 1–2 cm long, rarely 3 cm, 0.5–1.5(–3) mm
(isotypes: BM!, JE!, MO!). wide, often complanate-foliate, simple or irregularly
branched. Asexual reproductive bodies sometimes present
Taxicaulis bahiense Müll. Hal., Hedwigia 40: 59. 1901; Isopterygium ba- on stems, uniseriate, multicellular, simple or branched,
hiense (Müll. Hal.) Broth. in Engl. & Prantl, Nat. Pflanzenfam. 1(3): 1081. often more than 0.5 mm long, green or brown, cells papil-
1908. Type: Brazil. Bahia: Salzmann s.n. (possible isotype: H!). lose. Leaves 0.7–1.2 mm long, 0.2–0.5 mm wide, flaccid,
close, complanate, erect-spreading, often secund at the
Taxicaulis afflictus Müll. Hal., Hedwigia 40: 68. 1901; Isopterygium afflic- tips, smooth, ovate to lanceolate, often asymmetric, acu-
tum (Müll. Hal.) Broth. in Engl. & Prantl, Nat. Pflanzenfam. 1(3): 1083. minate; margins plane, serrate to serrulate above the leaf
1908. Type: Brazil. Goyaz, near Mossamedes, Ule 1616 (isotype: US!). middle, serrulate to entire below, rarely entire through-
out; costa none or weak, short and double; median cells
Taxicaulis ammophilus Müll. Hal., Hedwigia 40: 66. 1901; Isopterygium 38–132 × 5–12 µm, smooth; alar cells short-rectangular
ammophilum (Müll. Hal.) Broth. in Engl. & Prantl, Nat. Pflanzenfam. 1(3): to quadrate or transversely elongate, in small groups
46 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 24. Isopterygium tenerum. A. Habit. B. Enlargement of portion of stem. C, D. Leaves. E. Apical leaf cells. F. Median leaf cells. G,
H. Alar cells. I. Inoperculate and operculate capsules (dry). J. Inoperculate capsule (wet). K. Asexual reproductive body. L. Cells of asexual
reproductive body. Illustrations A, B, J drawn from Mexia 4841 (NY); C, E–G from Schiffner 362 (H, holotype of I. brachyneuroides); D from
Vital 7826 (MO); H from Pabst s.n. (H, isotype of Hypnum curvicollum); I from Mexia 5348a (NY); K, L from Santos 3965 (CANM).

with 1–5 cells on margins. Autoicous. Setae (0.3–)0.5–1.2 to obliquely short-rostrate; peristome teeth up to 0.2 mm
(–2.2) cm long, yellow to reddish brown; capsules 0.5– long. Spores 9–14 µm in diameter.
1.5 mm long, cernuous or rarely erect, curved or sometimes Distribution and ecology: Throughout the West In-
straight, ovoid to ellipsoid, usually strongly contracted dies and from Mexico to northern Argentina and Uruguay
below the mouth when dry; operculum conic-apiculate (Figures 18, 22, 23, 25); also known from eastern North
 number 93 • 47

FIGURE 25. Distribution of Isopterygium tenerum in the Caribbean.

America and recently reported from South Africa (Ochyra spreading, alar cells in small groups of short-rectangular
and Ireland, 2004). Recently, Matteri and Schiavone (2002) to quadrate or transversely elongate cells, asexual repro-
reported the species from the southern part of Argentina ductive bodies sometimes present on stems, filamentous,
in the Tierra del Fuego region. An extremely common spe- multicellular, the cells papillose; setae 0.5–1.2 cm long,
cies often occurring in forests on tree trunks, rotting logs rarely up to 2.2 cm, and capsules 0.5–1.5 mm long, ovoid
and stumps, woody debris, humus, soil banks, and rock, to ellipsoid, inclined to horizontal, usually strongly con-
especially sandstone and limestone; sometimes in swamps, tracted below the mouth when dry.
humid grasslands and on river banks, primarily at low el- Pterogonidium pulchellum (Hook.) Müll. Hal. in
evations from sea level to 500 m but known to occur up to Broth. (Sematophyllaceae) often grows intermixed with
3200 m. I. tenerum. The gametophytes of the two species are su-
Discussion: Isopterygium tenerum is not only the perficially somewhat similar except the leaves of Pterogo-
most common species of Isopterygium in Latin America, as nidium are usually narrowly lanceolate, symmetric, and
it is throughout the world, but it is also the most variable. more erect than the usually wider, ovate to lanceolate,
Its polymorphism undoubtedly explains the large number asymmetric, and spreading leaves of I. tenerum. Fortu-
of taxa that have been included here in its synonymy. Carl nately, Pterogonidium usually produces sporophytes and
Müller alone described 21 of the 45 taxa the senior author the capsules are erect, cylindrical, not contracted under
has recognized as synonyms from various Latin America the mouth when dry and have a single peristome. In con-
localities. trast, the capsules of I. tenerum are inclined to horizontal,
In its typical form, the species is best distinguished by ovoid to ellipsoid, have a double peristome, and are usu-
the usually complanate, medium-sized plants with stems ally strongly contracted under the mouth when dry.
1–2 cm long, 0.5–1.5 mm wide, leaves 0.7–l.2 mm long, We have taken a broad view of Isopterygium tenerum,
ovate-lanceolate, asymmetric, acuminate, close, erect- and some names synonymized here, with more study,
48 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

might be recognized. For example, Buck (1998) recog- Standley 11903 (F, FH). Olancho: Slopes of Montana Pena
nized I. brachyneuron on the basis of its more asymmetric Blanca, 11.5 km S of Gualaco, 980–1000 m, 14°57'N,
leaves, stronger costa, and fewer alar cells. Material iden- 86°08'W, Allen 12905 (MO). NICARAGUA. Sierra de
tified as I. fernandezianum might also be distinct at the Managua, 600–900 m, Garnier 715 (NY). Río Siquia,
species level on the basis of its strongly asymmetric leaves, 6 Jan 1941, LaRue (NY). COSTA RICA. Alajuela: Viento
strongly serrate leaf apices, and fewer alar cells. Fresco, 1600–1900 m, Standley & Torres 47877a (FH);
Selected specimens examined: MEXICO. Chiapas: Volcán Poás, 2500 m, Valerio 285 (US). Limón: Near Car-
Town of Tenejapa, 1768 m, Breedlove 14778 (MICH); men Station on Indiana Branch, ~30 m, Standley & Vale-
Santa Elena, 650 m, 16°53'N, 91°35'W, Delgadillo M. rio 48390 (FH, US). Heredia: Isla Bonita near Vara Blanca,
6308 (MO). Colima: Laguna María, Parque Nacional El ~1675 m, Svihla 3067 (FH). Puntarenas: Coto Brus Can-
Jabalí, Arreguín 790e (MEXU); Alzada, Orcutt 4645 (FH, ton, 1000–1200 m, 08°47'N, 83°02'W, Dauphin 1953
MICH, NY). Hidalgo: Above Tianguistengo on Hwy. 105, (MO); Cocos Is., Chatham Bay, Howell 120 (FH). Vicinity
3 km from Zacualtipan, ~2100 m, Sharp et al. 102 (CANM, of Coliblanco, ~1950 m, Maxon 250 (NY). PANAMA.
MICH, TENN). Jalisco: Known from type locality of I. di- Bocas del Toro: Cerro Colorado, 6.9 km above Chami
minutivum. Nayarit: l4.5 km N of Compostela, Koelz Camp, 1700 m, Allen 5335 (MO). Chiriquí: Boquete,
34043 (MICH). Oaxaca: Sierra Juárez, above Valle Nacio- 1100–1525 m, Svihla 2711 (MICH); El Volcán, Chicker-
nal along road from Tuxtepec to Oaxaca, 600 m, Richards ing 32a, 50 (CANM, NY); Santa Marta, Bugaba, Vinda
& Sharp 7051b (CANM, MICH, TENN). Puebla: At and 1521 (PMA); Cerro Colorado, 5.1 km from Escopeta,
near the summit of El Cerro de Cuhuatepetl, Tehuacan, 800–1000 m, Folsom 4920 (MO). Canal Area: Barro Col-
945–1160 m, Santos 3693 (MICH). Quintana Roo: La- orado Is., Willis 11 (FH, MO), 49 (MO); along Río Ma-
guna Ciega, Isla de Cozumel, 5 m, ~20°36'N, 86°44'W, sambi Grande between Gaillard Hwy., 1.5 km NW of
Delgadillo M. 4401, 4403, Cárdenas 1311, 1312, 1313 Summit Gardens, 70 m, Nee 6935 (MO); summit Experi-
(MEXU), 4402 (MEXU, MICH, MO). San Luis Potosí: W mental Garden, near Gamboa, Welch 20162, 20163,
of Xilitla, 1035 m, Sharp 5935 (CANM, MEXU, MICH, 20164, 20167, 20168 (NY). Darién: Above Cana, leading
TENN). Sinaloa: Along Hwy. 40, ~9.7 km W of Las Pal- to Pirre Massif, 800–900 m, Allen 8982 (MO). BER-
mitas, ~2285 m, Norris et al. 20425 (MICH). Tabasco: MUDA. Paget Marsh, Britton 44 (NY, US), 1855 (MICH,
Along trail between Teapa and Cerro de Azufre, 6 km from MO, NY, TENN). BAHAMAS. Andros Is.: Coppice, near
Teapa, Gilly & Hernandez 308c (MICH); Cerro del Mad- Nicholl’s Town, Small & Carter 8968 (NY, US). Great Ba-
rigal, Zamudio 1342b (MEXU). Tamaulipas: Edge of Casa hama Is.: Britton & Millspaugh 2628 (NY). New Provi-
de Piedra, near Rancho del Cielo above Gómez Farias, dence Is.: Prospect Ridge, Degener 19095 (NY). CUBA.
Sharp 8736 (CANM, F, MEXU, TENN); 32 km SW of Habana: Batabano, Baker & Wilson 2391 (MO). Oriente:
Tuxpan, road to Poza Rica, ~1000 m, Pursell 5138 (MO). Baracoa Region, Altos de Farola, 35 km S of Baracoa, 685
Veracruz: 10 km SE of Campo Experimental de Hule, El m, Webster 894 (CANM). Pinar del Río: Río Guao, Brit-
Palmar, Zongolica, Santos 3965 (CANM, MICH, NY); ton et al. 10142, 10127a (CANM, NY). Santiago: Slopes
~11.2 km NW of Tuxpan on road to Tampico, 15 m, Pur- and summit of El Yunque, near Baracoa, 305–610 m, Pol-
sell 5067 (MO). GUATEMALA. Izabal: ~11 km S of lard & Palmer 104 (NY), 110 (BM, NY, US). Isle of Pines,
Puerto Barrios, 50 m, Croat 41799 (MO). Petén: La Liber- Siguanea, Britton et al. 14373 (MICH, NY), 14932 (NY).
tad, Lundell 3384 (MICH). Quezaltenango: Vicinity of Ingenio Esperanza near Pinar del Río, Wright 206 (CANM,
Fuentes Georginas, slopes of Volcán de Zunil, 2500–2800 MICH, NY, TENN, US). JAMAICA. Clarendon: Ma-
m, Standley 85946 (F, FH, MICH, NY). (Dept. Unknown) son R. Savanna, ~4.4 km NW of Kellits, 700 m, Crosby
Finca El Naranjo, near Chicacoa, Svihla 2883 (FH). BE- 3118 (CANM, MO, NY). Hanover: E slope of Dolphin
LIZE. Cayo: San Augustín, Mains 3999 (MICH), 4084 (F, Head, 365–460 m, Proctor 10434 (CANM). Manchester:
FH, MICH, NY); Retiro, Mains 3698 (MICH), 3706 (FH, 1.6 km from Balaclave, Orcutt 816 (MICH). Portland:
MICH). HONDURAS. Atlántida: Lancetilla Valley, near Murdock’s Gap, 1740 m, Crosby 3536 (CANM, MEXU,
Tela, 20–600 m, Standley 55796 (F, FH, US), 55803 (F, MO, NY). St. Andrew: Cinchona, 1450 m, Farr 949
FH, NY, US); vicinity of San Alejo, base of hills S of San (CANM). St. Ann: 11 km NE of Moneague, 380 m, Farr
Alejo near Río San Alejo, 150–270 m, Standley 7662 (F). 194 (CANM). St. James: 6 km NE of Maroon Town on
Comayagua: Vicinity of Siguatepeque, 1080–1400 m, road to Spring Vale, 150 m, Crosby 13956 (MO). St.
Standley 56053 (F, FH, US). Francisco Morazán: Piedra Thomas: S slope of Winchester Peak, 1.8 km NW of
Herrada, lower slopes of Cerro de Uyuca, 1500–1650 m, Wheelerfield, S end of John Crow Mts., 350 m, Buck 5590
 number 93 • 49

(NY); Whitfield Hall, Hegewald & Hegewald 7960 (MO). tioquia: San José de la Montana, 13 km S of San José, road
Trelawny: Windsor Estate, 120–180 m, Proctor 10522 to la Labores, near Estación La Oculta, ~2600 m, Thiers
(CANM). Westmoreland: 4 km NE of Negril, near sea 4188-b (NY); Mpio. Anorí, Vereda El Carmen, Kms 18–
level, Proctor 35534 (NY). HAITI. Nord: Vicinity of Don- 20.5 N of Anorí, 820–910 m, 07°15'N, 75°12'W, Callejas
don, ~400 m, Leonard 8578 (CANM). Sud’est: Massif de et al. 8831 (NY); Mpio. Domingo, Corregimiento de San-
la Selle, 19 km S of Kenscoff on road from Port-au-Prince, tiago, sector “La Negra,” ~6 km W de Santiago, 1100–
1600 m, Buck 8997 (NY); Bayeux, at mouth of Rivière de 1165 m, ~06°30'N, 75°10'W, Churchill et al. 14706 (NY).
Port Margot, north coast, between Cap Haitien and Le Arauca: Mpio. de Fortul, Vereda Palmarito, 200 m,
Borgne, near sea level, Bartlett 17758a (MICH), 17756 06°44'N, 71°43'W, Churchill et al. 18939 (NY). Cauca:
(CANM, MICH, MO), 17765 (CANM, MICH); Valley of Guapi, Parque Nacional Natural Isla Gorgona, Isla Gorgo-
Rivière de Plaisance from Plaisance to Pilate, Bartlett nilla, cerca al Faro, Rudas & Aguirre 287 (COL); Mpio. de
17749 (MICH). DOMINICAN REPUBLIC. El Seibo: Piendamo, Corregimiento de El Mango, Reserva Natural
19 km N of Hato Mayor on road to El Valle, 300 m, Mejía El Guayabo, Km 93–94, 1640 m, ~02°46'N, 76°33'W,
et al. 10259 (MO, NY). La Vega: Vicinity of Piedra Blanca, Churchill & Franco 16536, 16562 (NY). Magdalena:
200–500 m, Allard 17816 (NY). Pedernales: ~6 km S of El Santa Marta, near Bonda, Baker 6563 (H). Meta: Out-
Aguacate on road to Pedernales, 1525 m, Reese 15286 skirts of Villavicencio, 500 m, 04°09'N, 73°39'W, Schultes
(NY). San Cristobal: Mano Matuey, 12 km from Cambito 11097 (H, MO, TENN, US), 11110 (BM, CANM, COL,
Garabito on road to El Cacao, 550 m, Buck 5166 (NY). La F, MICH, MO, NY). Nariño: Mpio. de Albán, Veredas
Altagracia: 6 km SE of Otra Banda on road to Punta Cana, Bellavista–El Cebadero, 1800–2000 m, 01°26'N, 77°05'W,
~30 m, Buck 5097 (NY). La Estrelleta: Along Río Limpio Ramírez P. 5910 (NY); Mpio. Tumaco, road to La Turbia,
at Colonia Río Limpio, 670 m, Reese 15398 (NY). Puerto 245 m, 01°18'N, 78°30'W, Ramírez P. et al. 9901 (MO).
Plata: Pico El Murazo, 1083 m, Buck 4839 (NY). Samaná: Putumayo: 15 km NW of Puerto Asís, 305 m, King &
Loma Pan de Azúcar, 6.8 km N of road to El Valle, 60 m, Guevara C-1122 (US). Santander: Mesa de los Santos,
Zanoni et al. 20758L (U). PUERTO RICO. Along Río la 1500 m, Killip & Smith 15329 (NY). Vaupés: Río Vaupés,
Coca, Sierra de Luquillo, S of Mameyes, Steere 7048 Mitú and surroundings, ~250 m, Schultes & Cabrera
(MICH). Río Sabana, Sierra de Luquillo, S from Luquillo, 13953 (CANM). Valle: Costa del Pacífico, Río Cajambre,
Steere 5172 (MICH, MO). Mpio. de Maricao, in moun- 5–80 m, Cuatrecasas 17383-B (US). VENEZUELA. Ama-
tains ~l.6 km outside Maricao, Stimson 3299 (MO, NY). zonas: Dept. Río Negro, along Río Mawarinuma, just out-
Villalba, Pagán 55 (MO), 54 (FH, MICH, MO), 60, 65 side Cañon Grande, vicinity of Neblina base camp, ~140 m,
(MICH, MO). Los Picachos de Jayuya, 915 m, Pagán 328 Buck 11227, 11352 (NY). Aragua: Parque Nacional Henry
(MO). E of Cidra, Masters 442 (M), 456 (M, MO). La Pittier, Las Locas, ~2000 m, Sharp C-7610A (TENN). Ba-
Torrecilla, Barranquitas, Steere 4689 (MICH, MO). Gu- rinas: Along Hwy. 5, 2 km W of Río Acequia, SW of junc-
vate Purchase Unit, Sierra de Cayey, Steere 4747 (FH, tion of road to Ciudad Bolivia, 175 m, Steyermark & Rabe
MICH, MO), 4804, 4849 (MICH, MO), 4858 (MO). Dist. 96510 (US). Bolívar: Dist. Roscio, Río Uairén, 9 km NE of
de Arecibo, Mpio. de Utuado, Stimson 3582 (NY). Mount Santa Elena de Uairén, 900 m, Steyermark & Liesner
Morales, near Utuado, Britton & Marblef 478, 1175 (NY), 127350 (MO). Carabobo: Between Los Tanques and La
1187 (MO, NY). La Juanita, near Las Marias, Britton Toma, at the head of Río San Gián, 750 m, Steyermark &
3999 (NY). Cerro de Las Pinas, near Las Cruces, 600–720 Steyermark 95588 (MO, US). Delta Amacuro: Dept. Ped-
m, Britton 6905 (NY). VIRGIN ISLANDS. St. Thomas: ernales, boundary with Dept. Tucupita, Caño Simoina, W
Slopes of St. Peter Mt., just off Crown Mt. Rd., 425 m, of Isla Cocuina, S of Barra de Cocuina, 50 m, 09°45'N,
Buck 3280 (NY). WINDWARD ISLANDS. Guadeloupe: 61°53'W, Steyermark et al. 114365 (MO). Distrito Fed-
Rivière Pérou, Le Gallo 811, 814, 815, 819, 820, 823 eral: Around Caracas, 800–1000 m, Pittier 11108 (FH,
(CANM). Dominica: Morne Micotrin, Elliott 1129c (BM). NY, US). Lara: Dist. Palavecino, 20 km from Cabudare,
Martinique: Rivière de Tivoli, 320 m, Stehlé 4039 (NY). 1200–1300 m, Steyermark et al. 103335 (MO, US).
St. Lucia: Between Castries and Dennery, Hegewald 9607 Mérida: Foot of Sierra Nevada, near Mérida, 1850 m,
(NY). St. Vincent: Morn Garn, 460 m, Elliott 56b (BM). Alston 6759a (BM). Miranda: Providencia, 1200 m, Me-
Grenada: St. George’s, 26 Jun 1906, Broadway (NY). dina 981, 991 (CANM). Monagas: Bajo Guarapiche, Ma-
NETHERLANDS ANTILLES. Saba: Mt. Scenery, trail turin, Medina 986 (CANM). TRINIDAD AND TOBAGO.
from parking area at SW end of mountain, 530–814 m, Trinidad: Vicinity of Tabaquite, Britton 2615 (FH, NY,
17°37'N, 63°14'W, Buck 50633 (NY). COLOMBIA. An- US); North Range, Welch 21272 (NY). Tobago: Arnos
50 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Vale Bay, Arnos Vale Hotel, 11°13'N, 60°45'W, Djan- 31332 (CANM, F, NY). Distrito Federal: Rio Sobradinho,
Chékar 95-13 (NY). GUYANA. Demerara-Mahaica Re- immediately W of Sobradinho, 1100 m, Irwin et al. 33235
gion: Along Linden-Soesdyke Hwy., ~1.6 km E of Soesdyke, (CANM, NY). Goiás: Mpio. de Formosa, Vital 8275
1–50 m, Pipoly 9286A, 9295 (NY). SURINAM. Broko- (MO); Mpio. de Caldas Novas, Vital 8501 (CANM, SP);
pondo: Phedra, along Surinam R., 67 km SE of Paramar- Mineiros, Galeriewald nahe der Grenze zu Mato Grosso,
ibo, 20 m, 05°20'N, 55°05'W, Allen 19342 (MO). 680 m, Schäfer-Verwimp & Verwimp 8554 (NY). Minas
Nickerie: Kabalebo Dam project, 0–50 m, Florschütz-de Gerais: Vicosa Agricultural College lands, Horticultural
Waard & Zielman 5496, 5687, 5712 (U). Paramaribo: Hill, 670 m, Mexia 4841 (BM, FH, F, MICH, MO, NY,
Cultuurtuin, Florschütz & Florschütz 607, 611 (U). Sipali- U); Vicosa Agricultural College lands, E boundary to reser-
wini: Blanche Marie valley, 100 m, 04°44'N, 56°53'W, voir, Mexia 5348a (BM, FH, MO, NY); Corrigo Riberro,
Muñoz 98-26b (MO); Talfelberg Natl. Park, vicinity of 675 m, Mexia 4884 (BM, F, FH, MICH, MO, NY, U, US),
Rudy Kappel airstrip, 325 m, 03°48'N, 56°08'W, Allen 4888 (F, FH, MO, US). Mato Grosso: Palmeiras, Lindman
20306, 20320 (MO); White Sand Savanna, 32 km NE of 413 (H). Pará: Belém, Reserva Mocambo, Lisbôa et al. 706
Apura, 55 m, 05°10'N, 57°00'W, Allen 19287 (MO). (MG); Serra do Cachimbo, Base Aérea do Cachimbo and
Brownsberg, trail to Leo Falls, 29 Jan 1976, 500 m, Sang vicinity, Km 780–820 on Cuiabá–Santaré Hwy. (BR 163),
& van der Wiel s.n. (U). Marowijne River, Armina Falls, ~20 km N of border of Mato Grosso, along Rio Braço
29 Sep 1952, Geijskes s.n. (MEXU, U). Blanche-Marieval, Norte, 430–480 m, Reese 16169, 16418 (MO), 16170,
vicinity of waterfall between Van Edenval and Graniet- 16209 (MICH, MO), 16142, 16171, 16175, 16178,
soela, 21–24 Aug 1982, Kuc s.n. (CANM). FRENCH 16185, 16190, 16192, 16195, 16210, 16218, 16237,
GUIANA. Approuague-Kaw: Pic Matécho, summit of wa- 16238, 16239, 16256, 16428 (NY), 16223 (MICH, MO,
terfall on N side of peak, ~500 m, 03°45'N, 53°02'W, US). Paraná: Mpio. Foz do Iguaçu, Parque Nacional do
Buck 37830 (NY); Commune de Régina, Kaw Mts. Trail Iguaçu, along Rio Iguaçu, 100–200 m, Vital & Buck 12097
up Montagne Favard from Savane de Kaw, 0–100 m, (NY). Rio Grande do Sul: S. Francisco de Paula, 900 m,
04°30'N, 52°03'W, Buck 32939 (MO); St.-Laurent-du- Sehnem 4645 (FH). Rio de Janeiro: Mpio. de Resende, Ita-
Maroni: Canton de Maripasoula, Commune de Saül, tiaia Natl. Park, S face of Mt. Itatiaia, 1780 m, Eiten &
~6 km N or Saül along road to Bélizon, vicinity of Eaux Eiten 7654, 7656 (US). Rondônia: Ridges of Serra dos Pa-
Claires, ~200 m, 03°37'N, 53°12'W, Buck 18690 (NY). caás Novos along the river, ~400 m, Reese 13320, 13556,
ECUADOR. Azuay: Gualaquiza, Allioni 8300, 8362, 8379 13616, 13623 (NY); Guajará-Mirim, estrada do Bom Sos-
(H). Cotopaxi: Las Juntas on road El Corazón–Quevedo, sego, Km 25, Lisbôa et al. 491 (MG); Guajará-Mirim,
200 m, Anderson 1010 (U). El Oro: ~17 km above Zara- 6 km NE of Guajara Mirim, 2 km E of gravel road, 175 m,
cay on road to Catacocha, 700 m, Anderson 1005 (U). 10°43'S, 65°15'W, Nee 34688 (NY). Roraima: Rio Urari-
Loja: Road Celica-Alamor, Km 11–16, 1600–1700 m, An- coeara, vicinity of Uaicá airstrip, Prance et al. 19953,
derson 944 (U). Pichincha: ~18 km NW of San Miguel de 19954, 19956 (CANM, NY). Santa Catarina: Tubarão,
los Bancos, 800 m, Buck 10326 (NY). Santiago-Zamora: Ule 87 (FH, H, NY, SP, US), 91 (BM, F, FH, H, NY, US).
Méndez, 700 m, Harling 2217 (CANM). Galapagos Is., São Paulo: Mpio. Guapiara, Serra Paranapiaacaba, Fa-
Isla Santa Cruz, trail from Bella Vista to Media Luna, zenda Intervales, ~800–1000 m, ~24°16'S, 48°25'W, Vital
550 m, Sipman M-234 (U). PERU. Junín: E of Quimirí & Buck 20430, 20449, 20458, 20475 (NY); Mpio. Te-
Bridge, near La Merced, 800–1300 m, Killip & Smith odoro Sampaio, Reserva Florestal de Teodoro Sampaio,
24038 (CANM, NY). Loreto: Maynas Prov., ~135 km NE Vital 7826 (CANM, MO); Mpio. Rancharia, along Ra-
of Iquitos, ~2 km N of Río Sucasari, 03°30'S, 72°55'W, poso Tavares road, between Presidente Prudente to Assis,
Timme 14349, 14351, 14352, 14359 (MO); Iquitos, Vital 2584 (MO). BOLIVIA. Beni: Prov. Ballivian, Beni
~100 m, Killip & Smith 26942 (CANM, NY); Middle and Biological Station, near Porvenir, ~60 km E of San Borja,
upper part of Río Maniti, E of Iquitos, 20 Jan 1981, Kuc 200–250 m, ~14°30'S, 66°30'W, Lewis 89-018 d-3 (MO);
s.n. (CANM). BRAZIL. Acre: 15 km E of Rio Branco vicinity of Guayaramerín, Reese 12739, 12753, 12775,
along road to Abuna, Reese & McPherson 13195 (CANM, 12794, 12798, 12803 (CANM, NY). Cochabamba: Prov.
NY). Amazonas: Ilha Acarabu, along the Rio Negro be- Carrasco, Municipio Pojo, Estación Experimental Sacta,
tween Manaus and São Gabriel at the mouth of the Rio 250 m, 17°05'S, 64°46'W, Churchill & Arroyo P. 21634
Marié, Buck 2347 (CANM, NY); estrada Manaus-Itacoa- (US); Prov. Chaparé, road to Corani, 3200 m, Hermann
tiara, Km 26, reserva florestal Ducke, Lisbôa 57, 63, 70 24671 (US). La Paz: Prov. Nor Yungas, “Alto Coro Coro,”
(MG). Bahia: ~10 km W of Barreiras, 500 m, Irwin et al. 7 km N of Caranavi, 15°46'S, 67°32'W, Lewis 84-066
 number 93 • 51

(LPB); Prov. Inquisivi, “Río Pichincha Grande,” along Río isotypes: BM!, US!). = Wijkia alstonii (E. B. Bartram)
Pichincha Grande from its mouth at the Río Cañamina to Ireland, Caldasia 16(76): 267. 1991.
its headwaters above Rancho Pichincha, ~7 km SE of
Miguillas, 16°35'S, 67°17'W, Lewis 86-1462 (LPB). Santa Isopterygium semicostatum Renauld & Cardot, Bull. Soc.
Cruz: Prov. Nuflo de Chavez, “Doloresa,” along San Roy. Bot. Belgique 41(1): 143. 1904 [1905]. Type: Costa
Javier–Concepción road, ~1 km past turnoff for Esstancia Rica: Forêts de General, Feb 1891, Pittier & Tonduz
Doloresa, approximately half way between San Javier and 5817 in Pittier & Durand, Plantae costaricenses exsicca-
Concepción, 16°14'S, 62°25'W, Lewis 85-925A (LPB); tae (isotype: US!). = Lepyrodon tomentosus (Hook.) Mitt.
Prov. Chiquitos, “Río Roboré,” base of Serrania Santiago according to H. Robinson in The Mosses of Juan Fernan-
in canyon along the Río Roboré, 18°18'S, 59°44'W, Lewis dez Islands, Smithsonian Contr. Bot. 27: 52. 1975.
85-1212 (LPB); Prov. Velasco, Natl. Park Noel Kempff
Mercado, 250 m, 14°32'S, 60°48'W, Churchill & Churchill Isopterygium subtrichopelma Renauld & Cardot, Bull.
20505 (MO); near Aten, Williams 2070 (F, FH, MICH, Soc. Roy. Bot. Belgique 41(1): 142. 1904 [1905]. Type:
NY, TENN, US). Tumupasa, Williams 2059 (F, FH, Costa Rica: Forêts du Barba, 31 Aug. 1889, Pittier &
MICH, TENN). Apolo, Williams 2091 (BM, F, FH, MICH, Tonduz 5790 (isotype: FH!). = Mittenothamnium sp.
NY, US). New Brazil, Williams 2094 (MICH, NY, US).
CHILE. Juan Fernandez Iss., Mas a Tierra, Valle Colonial, Isopterygium Nomina Dubia
435 m, Skottsberg & Skottsberg 484 (NY); trail to Porte-
zuelo de Villagra, 427–549 m, Hatcher & Engel 14 (US); The status of the following Latin American taxa must re-
Valle Anson, 400–500 m, Skottsberg M40 (H). PARA- main unknown until their types can be found and examined.
GUAY. Amambay: Parque Nacional Cerro Corá, trail up
Cerro Muralla, ~300 m, ~22°40'S, 56°00'W, Buck 12518 Isopterygium brevisetum (Hornsch.) Broth., Nat. Pflanzen-
(NY). Caaguazú: Coronel Oviedo, Barrio Azucena, Bordas fam. 1(3): 1081. 1908; Hypnum brevisetum Hornsch., Fl.
114A (NY). Canendiyú: 2.5 km E of Río Corrientes, road Bras. 1(2): 78. 1840. Type: Brazil. Near Rio de Janeiro,
from Salto del Guairá, ~200 m, ~24°35'S, 55°55'W, Buck Jul and Aug, Merkel s.n. Type apparently not in BM.
12446A (NY). Central: San Lorenzo, Ciudad Universita-
ria, 25°30'S, 57°30'W, Bordas 61B (NY). Cordillera: Cho- Isopterygium chrismarii (Müll. Hal.) Mitt., J. Linn. Soc.,
loló, Km 80 S of Piribebuy, 100–200 m, 25°33'S, 57°02'W, Bot. 12: 500. 1869; Hypnum chrismarii Müll. Hal., Syn.
Churchill 20248 (MO, NY). Guairá: Villarrica, 25°45'S, Musc. Frond. 2: 682. 1851. Type: Mexico. Michoacán,
56°26'W, Bordas 65 (NY). Itapúa: Centro de Desarrollo Cerro San Andres, Mar 1849, Chrismar s.n.
Forestal (CEDEFO) at Km 428 on Ruta 6, just W of Ar-
royo Pirapó, ~100 m, ~26°55'S, 55°29'W, Buck 12171 Isopterygium exiguum Kindb., Enum. Bryin. Exot. 100.
(NY). Paraguarí: Parque Nacional Ybycuí, trail along Ar- 1891 (new name); Hypnum exiguum Geh. & Hampe
royo Mina, ~200 m, ~26°05'S, 56°53'W, Buck 12107, in Hampe, Vidensk. Meddel. Dansk Naturhist. Foren.
12141 (NY); El Chaco, Rosario, Lindman 324 (H); Ma- Kjøbenhavn ser. 4, 1: 140. 1879, hom. illeg. Type: Bra-
cizo Acahay, 500 m, 25°54'S, 57°09'W, Zardini & Cuevas zil. Near Apiahy, Jun 1877, Puiggari s.n. Type appar-
5397 (MO). ARGENTINA. Lago Encondido, Matteri 799 ently not in BM.
(BA). Buenos Aires, Santa Catalina, 7 Aug 1973, Oleksy
s.n. (BA). San Miguel de Tucuman, Lorentz s.n. (FH). Isopterygium exile (Müll. Hal.) Paris, Index Bryol. Suppl.
Parque Nacional, Iguazú, 4 Jan 1984, Kuc s.n. (CANM). 219. 1900; Taxicaulis exilis Müll. Hal., Hedwigia 36:
URUGUAY. Canelones, Banado del Negro, cerca de Ai- 116. 1897. Type: Paraguay. Balansa 3619.
royo Pando, Castellanos 8582 (FH). Curticeiras Rivera,
Ruta 5, Zorrón 2098 (US). Prov. Rocha, Parque Nacional Isopterygium fruticola (Müll. Hal.) Paris, Index Bryol.
Sta. Teresa, 5 Mar 1984, Kuc s.n. (CANM). Suppl. 219. 1900; Taxicaulis fruticola Müll. Hal., Hed-
wigia 37: 252. 1898, “fruticolus.” Type: Surinam. Near
Isopterygium Excluded Taxa Paramaribo, Aug 1844, Kegel s.n.

Isopterygium alstonii E. B. Bartram, Bryologist 56: 168. Isopterygium gracillimum (Hornsch.) Broth. in Engl. &
1953. Type: Venezuela: Federal Dist., Pico de El Avila, Prantl, Nat. Pflanzenfam. 1(3): 1081. 1908; Hypnum
alt. 2000 m, 17 Dec 1938, Alston 5574c (holotype: FH!; gracillimum Hornsch., Fl. Bras. 1(2): 78. 1840. Type:
52 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Brazil. Province Minarum, collector unknown. Type ap- 8. Phyllodon Bruch & Schimp. in Bruch,
parently not in BM. Schimp. & W. Gümbel

Isopterygium hookeriophilum (Müll. Hal.) Broth. in Engl. Phyllodon Bruch & Schimp. in Bruch, Schimp. & W. Güm-
& Prantl, Nat. Pflanzenfam. 1(3): 1083. 1908; Taxicau- bel, Bryol. Eur. 5 (fasc. 46–47, Monogr. Hookeria 4): 60.
lis hookeriophilus Müll. Hal., Hedwigia 40: 67. 1901. 1851. Type: Hookeria retusa Wilson, nom. nud. = Phyl-
Type: Brazil. São Paulo, 1891, Wohltmann s.n. lodon truncatulus (Müll. Hal.) W. R. Buck.

Isopterygium microplumosum (Müll. Hal.) Broth. in Glossadelphus M. Fleisch., Musci Buitenzorg 4: 1351. 1923. Lectotype:
Engl. & Prantl, Nat. Pflanzenfam. 1(3): 1083. 1908; Hypnum truncatulum Müll. Hal. (designated by Robinson, 1974) ≡ Phyl-
Taxicaulis microplumosus Müll. Hal., Hedwigia 40: lodon truncatulus (Müll. Hal.) W. R. Buck.
68. 1901. Type: Brazil. Minas Gerais: Serra Ouro
Preto, Ule 1486. Plants small to medium-sized, in dull, soft to ± stiff,
flat mats, green to yellow-green. Stems creeping, subpin-
Isopterygium miradoricum (Müll. Hal.) A. Jaeger, Ber. nately branched, complanate-foliate; stems in cross sec-
Thätigk. St. Gallischen Naturwiss. Ges. 1876–77: tion without a hyalodermis, with small, thick-walled cells
436. 1878; Hypnum miradoricum Müll. Hal., Linnaea surrounding larger thin-walled cells, central strand pres-
38: 650. 1874. Type: Mexico. Mirador, Apr 1873, ent; pseudoparaphyllia foliose; axillary hairs consisting of
Sartorius s.n. a single short brown basal cell and 1–4 elongate hyaline
distal cells. Stem and branch leaves similar but dorsal and
Isopterygium pseudosubulatum (Müll. Hal.) Paris, Index ventral leaves differentiated from lateral leaves, compla-
Bryol. Suppl. 220. 1900; Taxicaulis pseudosubula- nate, erect- to wide-spreading, often asymmetric, oblong
tus Müll. Hal., Flora 83: 340. 1897. Type: Venezuela. to ligulate, dorsal and ventral leaves narrower than lateral
Goebel s.n. leaves, obtuse to acute, lateral leaves very concave; mar-
gins serrate above, the teeth sometimes bifid, plane; costa
Isopterygium pygmaeocarpum (Müll. Hal.) Broth. in short and double or absent; cells linear-flexuose, prorulose
Engl. & Prantl, Nat. Pflanzenfam. 1(3): 1081. 1908; and/or seriately papillose, thin- to firm-walled; alar cells
Taxicaulis pygmaeocarpus Müll. Hal., Hedwigia 40: not or scarcely differentiated. Asexual propagula not seen.
66. 1901. Type: Brazil. Santa Catarina: Tubarão, Dioicous, autoicous, or synoicous. Perichaetial leaves
Ule 973. often crispate, lanceolate to ovate, acuminate to acute or
obtuse; margins serrate, plane; costa usually none; cells
Isopterygium sapricola (Müll. Hal.) Broth. in Engl. & linear-flexuose, prorulose and/or papillose, becoming
Prantl, Nat. Pflanzenfam. 1(3): 1083. 1908; Taxicau- laxly rectangular and smooth toward the insertion; alar
lis sapricola Müll. Hal., Hedwigia 40: 67. 1901. Type: cells not differentiated. Setae elongate, smooth or rough-
Brazil. Rio de Janeiro, Monte Corcovado, Ule 165. ened above, reddish; capsules inclined, arcuate, asymmet-
ric, ovoid; exothecial cells quadrate to short-rectangular,
Taxicaulis sapricola var. minor Müll. Hal., Hedwigia 40: firm-walled, not collenchymatous but sometimes with the
68. 1901. Type: Brazil. Rio de Janeiro, Monte Corco- horizontal walls thinner than the vertical walls; annulus
vado, Ule 1722. differentiated; operculum high-conic; peristome double,
exostome teeth yellow-brown, shouldered, bordered, on
Isopterygium subcurvicolle (Müll. Hal.) Paris, Index Bryol. the front surface with a zigzag median line, not furrowed,
ed. 2, 3: 125. 1905, “subcurvicollum”; Taxicaulis sub- cross-striolate below, coarsely papillose above, trabecu-
curvicollis Müll. Hal., Hedwigia 40: 65. 1901. Types: late at back; endostome with a high basal membrane, seg-
Brazil. Santa Catarina, Apr 1888, Ule s.n.; Tubarao, ments keeled, perforate, nearly as long as the exostome,
Ule 640, 783; Serra Geral, Ule 1202. cilia mostly single. Spores spherical, papillose. Calyptrae
cucullate, naked, smooth, falling early.
Isopterygium subsplendidulum (Müll. Hal.) Paris, Index Discussion: We have one species of Phyllodon in
Bryol. Suppl. 221. 1900; Taxicaulis subsplendidulus the New World; the remainder of the species are Asian
Müll. Hal., Bull. Herb. Boissier 5: 210. 1897. Types: and African. The genus is characterized by ligulate leaves,
Guatemala. Mazatenango, Bernoulli & Cario 75, 84. often conspicuously toothed at the apex, with laminal cells
 number 93 • 53

strongly prorulose and/or seriately papillose. The alar cells or absent; cells linear-flexuose, prorulose and/or seriately
are scarcely differentiated. The genus probably has a rela- papillose, the papillae lower in basal half of leaf, thin-
tionship with Taxiphyllum rather than to the Sematophyl- walled; alar cells scarcely differentiated, with a few sub-
laceae, where Phyllodon (as Glossadelphus) has often been quadrate, concolorous cells in basal angles and across the
placed. Indeed, species like Taxiphyllum laevifolium and insertion. Asexual propagula none. Synoicous. Perichaetial
T. ligulaefolium seem to bridge the gap between the two leaves 0.6–1.25 mm long, ovate to oblong-ovate, obtuse to
genera. Tentatively, though, we are not merging the gen- short-acuminate; margins singly to doubly serrate above,
era (with Phyllodon as the older name) but keeping them entire below, plane; costa none; cells linear-flexuose, spicu-
separate on the basis of cells sometimes papillose over the lose, the “papillae” to 30 µm tall, thick-walled, shorter and
lumina and the almost total lack of alar differentiation in smooth toward the insertion; alar cells not differentiated.
Phyllodon. This relationship was first hinted at by Buck Setae 1.5–2.5 cm long, elongate, smooth, reddish; capsules
(1987) when he suggested the use of the name Phyllodon ~1.25 mm long, inclined, arcuate, asymmetric, ovoid; exo-
rather than Glossadelphus. thecial cells oval to short-rectangular, firm-walled, not col-
lenchymatous; annulus of 1–2 rows of thick-walled cells;
Phyllodon truncatulus operculum ~0.8 mm long, high-conic; exostome teeth
yellow-brown, shouldered, bordered, on the front surface
Phyllodon truncatulus (Müll. Hal.) W. R. Buck, Mem. cross-striolate below, coarsely papillose above, trabeculate
New York Bot. Gard. 45: 521. 1987. Hypnum truncatu- at back; endostome yellowish, with a high, smooth basal
lum Müll. Hal., Syn. Musc. Frond. 2: 263. 1851; Ectro- membrane, segments finely papillose, keeled, perforate,
pothecium truncatulum (Müll. Hal.) Mitt., J. Linn. Soc., nearly as long as the exostome, cilia finely papillose, single,
Bot. 12: 517. 1869; Trichosteleum truncatulum (Müll. broad, 2–3-seriate below, sometimes split in two above.
Hal.) A. Jaeger, Ber. Thätigk. St. Gallischen Naturwiss. Spores 13–17 µm in diameter, spherical, finely papillose.
Ges. 1876–77: 415. 1878; Glossadelphus truncatulus Calyptrae cucullate, naked, smooth, falling early.
(Müll. Hal.) M. Fleisch., Musci Buitenzorg 4: 1352. Distribution and ecology: Costa Rica, Panama,
1923. Type: Peru. Pöppig (BM!). Colombia, Ecuador, Peru, Brazil, French Guiana, Jamaica,
Hispaniola (Haiti and Dominican Republic), Puerto Rico,
Hookeriopsis cocoensis R. S. Williams, Bryologist 27: 40. 1924; Glossa- and St. Vincent; growing mostly on moist rocks but also on
delphus cocoensis (R. S. Williams) E. B. Bartram, Proc. Calif. Acad. Sci. tree trunks, in humid forests, at 200–600(–1200) m.
IV, 21: 86. 1933. Type: Cocos Island, 1905–1906, A. Stewart 1432a (holo- Illustrations in publications: Fig. 174 in Florschütz-
type: NY!). de-Waard and Veling (1996: 455); Pl. 129 in Buck (1998:
327); Fig. 142 in Buck (2003: 160).
Glossadelphus longisetus E. B. Bartram, Contr. U.S. Natl. Herb. 26(3): 109. Discussion: Phyllodon truncatulus is an extraordi-
1928. Type: Costa Rica. Limón: vicinity of Guapiles, 300–500 m, 12–13 nary species not likely to be confused with any other due to
Mar 1924, Standley 37280 (holotype: FH!; isotype: US!). the ligulate leaves with broadly rounded apices and strongly
prorulose and/or papillose cells. The marginal bifid serra-
Plants somewhat small, in dull, ± stiff, flat mats, green tions are particularly striking. The species seems close to
to yellow-green. Stems to ~3 cm, creeping, elongate, sub- the Angolan P. truncatus (Welw. & Duby) W. R. Buck but
pinnately branched, the branches numerous, elongate, differs, among other features, by the smooth rather than
complanate-foliate; stems in cross section with 2–3 rows strongly roughened seta.
of small thick-walled cells surrounding abruptly larger Selected specimens examined: COSTA RICA.
thin-walled cells, central strand of small, very thin-walled Cartago: Vicinity of Pejivalle, 900 m, Standley & Valerio
cells; pseudoparaphyllia foliose; axillary hairs consisting 47141 (NY). Guanacaste: Slopes of Miravalles, above Bi-
of 1 short, brown basal cell and 2 elongate, hyaline distal jaagua, 1500 m, Gomez et al. 19105 (MO, NY). Puntar-
cells. Stem and branch leaves similar but dorsal and ven- enas: Near Monteverde, 1373 m, 20 Feb 1972, W. James
tral leaves differentiated from lateral leaves, 0.75–1.0 mm s.n. (NY). PANAMA. Panamá: 5.5 km N of Cerro Azul,
long, complanate, erect- to wide-spreading, often asym- 763 m, Tyson et al. 2483 (NY). JAMAICA. Portland Par-
metric, broadly ligulate, dorsal and ventral leaves narrower ish: Hardwar Gap Falls, ~1220 m, Welch 17594A (NY). St.
than lateral leaves, broadly rounded to emarginate, lateral Andrew Parish: Trail from Hardwar Gap to Mt. Caledonia
leaves very concave; margins sharply serrate above, the (Old England Trail),1.4–2.8 km NNW of Hardwar Gap,
teeth bifid, serrulate below, plane; costa short and double 18°06'N, 76°44'W, 1342–1403 m, Crosby 3210 (DUKE,
54 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

NY). St. Thomas Parish: Corn Puss Gap and vicinity, ~12.4 with relatively short branches; stems in section without a
km N of Bath on trail to Port Antonio, 18°00'N, 76°21'W, hyalodermis, with 2–4 outer layers of small, thick-walled
550 m, Buck 5542, 5586 (NY). DOMINICAN REPUB- cells surrounding ± thin-walled, large cells, central strand
LIC. La Vega: Vicinity of Piedra Blanca, near Goodrich absent; pseudoparaphyllia filamentous; axillary hairs with
Rubber grove, 200–500 m, Allard 18812 (NY). ST. VIN- 1–2 short, brown basal cells and 3–5 elongate, hyaline
CENT. Below 305 m, Smith 1490a (NY). BRAZIL. Ama- apical cells. Stem and branch leaves only slightly differen-
zonas: Along the Rio Negro, slopes and summit of Serra tiated, the stem leaves somewhat larger and occasionally
Curicuriari, from Igarapé Arabú of the Rio Curicuriari to with recurved margins and laxer cells across the insertion.
the summit, 0°20'S, 66°50'W, 450 m, Buck 2477 (INPA, Branch leaves erect-spreading to imbricate, lanceolate to
NY). Mato Grosso: Chapada dos Guimaraes, Wasserfall ovate, acute to narrowly acuminate, slightly or (in broader
Véu da Neuva, 500 m, Schäfer-Verwimp 8626 (NY). Minas leaves) distinctly concave, short-decurrent; margins plane
Gerais: Mpio. São Roque de Minas, Parque Nacional da or reflexed, entire to serrulate near the apex or com-
Serra da Canastra, Caachoeira Casca D’Anta, 20°18'S, monly almost to the base, the marginal cells somewhat
46°31'W, 1000 m, Vital & Buck 11936 (NY, SP). Rio de broader and forming an indistinct border; costa short
Janeiro: Parque Nacional do Itatiaia, along trail to Véu and double, rarely absent; upper cells linear-rhomboidal,
de Noiva and Itaporani, ~13.5 km N of town of Itatiaia, smooth, straight to indistinctly flexuose; basal cells lax,
22°25'S, 44°36'W, 1100 m, Vital & Buck 19976 (NY, SP). thin-walled and porose; alar cells quadrate to oblate in
Santa Catarina: Blumenau, Park Floresta Negra, 200 m, sizable triangular groups usually not reaching the costa,
Schäfer-Verwimp 8240 (NY). São Paulo: São Paulo, Pico the walls distinctly thicker than those of the upper cells,
do Jaragua, 810 m, Schäfer-Verwimp & Verwimp 9026 decurrent in 1–3 rows with the cells in the decurrencies
(NY). COLOMBIA. Antioquia: Mpio. Santo Domingo, conspicuously enlarged. Dioicous or autoicous. Perichae-
Corregimiento de Santiago, sector “La Negra,” ~6 km tial leaves oblong-lanceolate, acuminate, serrulate above,
O [W] de Santiago, 06°30'N, 75°10'W, 1100–1165 m, ecostate; upper cells linear and lower cells ± rhomboidal-
Churchill et al. 14851 (NY). Caquetá: Transecto Neiva– rectangular, large, lax. Setae reddish, slightly twisted,
San Vicente del Caguán, Quebrada Chorreras, entre Guay- smooth; capsules erect, symmetric, cylindrical, red-brown;
abel y Los Andes, 02°30–33'N, 74°30'–75'00W, 1300 m, exothecial cells thin- to thick-walled, oblong to square;
Churchill & Betancur 16864-B (NY). Magdalena: Parque annulus none; operculum conic to short-rostrate, straight
Nacional de la Sierra Nevada de Santa Marta, Vertiente ori- to oblique, the cells usually thick-walled; peristome at-
ental del Río Buritaca, 700 m, Griffin et al. 1137 (FLAS, tached slightly below the mouth, double; exostome teeth
NY). ECUADOR. Napo: Añangu, S shore of Río Napo, lanceolate, strongly bordered, on the outer surface with a
~75 km E of Puerto Francisco de Orellana, “Coca,” 0°32'S, zigzag median line and ± broad plates, irregularly striolate-
76°23'W, 245–325 m, Churchill & Sastre-De-Jesús 13820 papillose to cross-striolate below (or sometimes the basal
(AAU, NY). Pastaza: ~5 km SE of Puyo on road to Macas, plates smooth and with strongly thickened walls), be-
01°31'S, 77°55'W, 1000 m, Churchill & Sastre-De-Jesús coming papillose to obliquely striolate at the middle and
13688 (AAU, NY). FRENCH GUIANA. St-Laurent-du- ± smooth above, on the inner surface with ± narrow, non-
Maroni: Canton de Maripasoula, Commune de Saül, ~6 km projecting trabeculae, smooth or coarsely papillose, oc-
N of Saül along road to Bélizon, vicinity of Eaux Claires, casionally perforate; endostome with a fairly high basal
03°37'N, 53°12'W, 200 m, Buck 18508 (NY). PERU. San membrane, papillose to granulose, segments narrow, flat
Martín: Prov. Lamas, Strasse Yurimaguas–Tarapoto, Km to slightly keeled, not perforate, papillose below, smooth
112, 900 m, Frahm et al. 1472 (B, NY). above, nearly as long as the teeth, cilia none or rarely ru-
dimentary. Spores finely papillose, 12–35 µm in diameter.
9. Platygyriella Cardot Calyptrae cucullate, naked, smooth.
Discussion: Platygyriella (and its synonym, Bryo-
Platygyriella Cardot, Rev. Bryol. 37: 9. 1910. Type P. he- sedgwickia [Buck, 1984a]) was traditionally placed near
licodontioides Cardot Bryosedgwickia Cardot & Dixon the Entodontaceae, especially Erythrodontium, because
in Dixon, J. Bot. 50: 154. 1912. Type: B. kirtikarii Car- of the numerous alar cells and the erect capsules (Buck,
dot & Dixon in Dixon. 1980). However, the relationships are more with the Hyp-
naceae because of the exostome ornamentation. The genus
Plants in slender to medium-sized mats, mostly is characterized by its lanceolate to ovate leaves that are
golden, shiny or dull. Stems creeping, irregularly pinnate mostly acuminate, a short, double costa, and numerous
 number 93 • 55

oblate to quadrate alar cells. Gametophytically, the best is erect and symmetric, with a reduced endostome having
character by which to recognize the genus is the enlarged a relatively high basal membrane, narrow, mostly flat seg-
cells of the decurrencies that usually remain attached to ments, and no cilia or only rudimentary ones. In the two
the stem on dissection. In this respect it is like Ectropo- American species the basal membrane is almost always
thecium, but in that genus there is but a single enlarged coarsely granular. In addition, there are two African and
cell in the decurrencies, rather than several. The capsule three Asian species of Platygyriella.

KEY TO THE SPECIES OF PLATYGYRIELLA

1. Leaves erect-spreading, lanceolate to lanceolate-ovate, gradually acuminate, 0.35–0.8 mm long . . . . . . . . . . . . 1. P. densa
1. Leaves imbricate, broadly ovate, abruptly acuminate, 0.9–1.2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. P. pringlei

1. Platygyriella densa brown basal cell(s) and 3–5 elongate, hyaline apical cells.
Asexual reproduction sometimes by flagellate branches.
Platygyriella densa (Hook. in Kunth) W. R. Buck, Brit- Leaves 0.35–0.8 mm long, erect-spreading wet or dry, lan-
tonia 36: 86. 1984a; Leskea densa Hook. in Kunth, ceolate to lanceolate-ovate, concave, gradually acuminate;
Syn. Pl. 1: 61. 1822; Entodon densus (Hook. in Kunth) margins entire or more often minutely serrulate almost
Mitt., J. Linn. Soc., Bot. 12: 531. 1869; Erythrodon- throughout, plane or sometimes reflexed; costa short and
tium densum (Hook. in Kunth) Paris, Index Bryol. ed. 2, double; cells linear, rarely shorter at the apex; alar cells
2: 158. 1904; Bryosedgwickia densa (Hook. in Kunth) quadrate to oblate, usually extending to the costa, the de-
Bizot & P. de la Varde, Rev. Bryol. Lichénol. 20: 246. currencies usually of 1–2 greatly enlarged cells. Dioicous.
1951. Type: Mexico. [Michoacán:] Between Pazcuaro Setae 1.2–1.5 cm long; capsules ~2 mm long; operculum
and Ario, 1828–2111 m, Humboldt & Bonpland s.n. short- to high-conic; exostome teeth orange, striolate to
(holotype: BM!; isotype: NY!). papillose below, smooth or faintly papillose above; endo-
stome with a moderately high basal membrane, with walls
Platygyrium guatemaliense Schimp. ex A. Jaeger, Ber. Thätigk. St. Gallischen sometimes very thickened, often coarsely papillose, and
Naturwiss. Ges. 1876–77: 276. 1878, nom. nud. narrow, flat, finely to coarsely papillose segments, cilia
none. Spores 13–20 µm in diameter.
Entodon pallidissimus Müll. Hal., Linnaea 42: 494. 1879; Erythodontium Distribution and ecology: Mexico (where it is
pallidissimus (Müll. Hal.) Paris, Index Bryol. 437. 1896; Bryosedgwickia common and weedy), Costa Rica, Venezuela, Bolivia, Bra-
pallidissima (Müll. Hal.) Bizot & P. de la Varde, Rev. Bryol. Lichénol. 21: zil, Paraguay, and Peru; mistakenly reported from Haiti
10. 1952. Type: Venezuela. Prope Coloniam Tovar, 1854–5, A. Fendler 96 (Buck and Steere, 1983); commonly on bark of deciduous
(lectotype: NY!, designated by Buck, 1984a). tree trunks, sometimes on rocks, in mesic to dry forests, at
(200–)400–1700(–2400) m.
Platygyriella helicodontioides Cardot, Rev. Bryol. 37: 9. 1910. Type: Mex- Illustrations in publications: Fig. 161F–H in Bar-
ico. México: Amecameca, 1908, Pringle 15256 (lectotype: PC!; isolectotypes: tram (1949: 363); Pl. 757 in Buck (1994: 1020).
NY!, US!, both types designated by Buck, 1984a). Discussion: This moss is often found in mixtures
with other mosses, often on tree trunks. When by itself, it
Erythrodontium subdensum Broth. & Thér. in Thér., Recueil Publ. Soc. is frequently small, sterile, and weatherworn. It is distinc-
Havraise Études Diverses 88: 313. 1921; Bryosedgwickia subdensa (Broth. tive by its small size, ovate-lanceolate, gradually acumi-
& Thér.) Bizot & P. de la Varde, Rev. Bryol. Lichénol. 21: 10. 1952. Type: nate leaves, short-double costa, and one or two greatly
Costa Rica. Alajuela, 900 m, Jan 1910, O. Jiménez s.n. (holotype: H!; iso- enlarged cells of the decurrencies (that typically remain
type: PC!). attached to the stems upon dissection). The moderately
high basal membrane of the endostome, which is coarsely
Plants small and slender, in mats, dull green to golden. papillose and often has strikingly thick walls, and narrow,
Stems to ~4 cm long, mostly shorter, irregularly branched, flat segments are unmistakable. There is little chance of
the branches prostrate, short; stems in cross section with confusion with P. pringlei, which has much larger, imbri-
2–4 rows of small, thick-walled cells surrounding abruptly cate, and more or less abruptly tapered leaves and usually
larger, thinner walled cells, central strand lacking; pseu- more than two enlarged cells in the decurrencies, as well
doparaphyllia filamentous; axillary hairs with 1(–2) short, as a rupestral habit.
56 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Specimens examined: MEXICO. Chiapas: Mpio. Valle, Tyson 2477 (MO); Los Santos: Trail between Jobero
Angel Albino Corzo, 9 km S of Finca Liquidambar, 15°42'N, and headwaters of Río Pedregal, 500–780 m, Croat 34535
92°45'W, 2000–2150 m, Delgadillo 4707 (MEXU, NY). (MO). BOLIVIA. Tajira: Prov. Arce, slope of Cerro Nogal,
Districto Federal: La Venta, San Miguel, 3469–3965 0.5 km N of Quebrada Nogal, 2–5 km NW of Commu-
m, 17 Oct 1926, Antipovich s.n. (MO, NY). Guerrero: nidad La Mamora, 22°09'S, 64°41'W, 1230 m, Lewis
Mpio. Chilpancingo de los Bravos, Ixtamalco, 18 km W 84-2157 (F, NY). BRAZIL. Goiás: Alto Paraíso, Chapada
of Chilpancingo on road to Olmitemi, 1500 m, Thomas dos Veadeiros, am Rio São Miguel westlich des National-
& Contreras 3728 (NY). Jalisco: Etzaltan, Pringle, Plantae parks, 800 m, Schäfer-Verwimp 9857 (NY). PARAGUAY.
Mexicanae 10617 (US); Río Blanco, Guadalajara, Pringle Cordillera: Cerro Ybitú Silla, 1 km S of Tobatí, 25°12'S,
15235 (US); N slope of La Cumbre, 16 km SW of Autlán 57°07'W, 297 m, Zardini 6024 (MO, NY). PERU. Above
on La Resolana road, 1464 km, Crum 481 (MICH, NY). San Miguel, 1830 m, Foote 25 (NY).
México: Barranca de Malinaltenango, 18°46'N, 99°42'W,
1450–1600 m, Buck 18174 (NY); Santa Fé, Pringle 15653 2. Platygyriella pringlei
(US); Canada, Pringle 15113 (US). Michoacán: Vicinity of
Morelia, Cerro Azul, 2300 m, Arsène 4534, 4546, 4551, Platygyriella pringlei (Cardot) W. R. Buck, Brittonia 36:
4795, 4796 (US); Jesús del Monte, 2000 m, Arsène 7614 87. 1984a; Erythrodontium pringlei Cardot, Rev. Bryol.
(US); Campanario, 2200 m, Arsène 7450, 7525 (US); on 37: 11. 1910; Bryosedgwickia pringlei (Cardot) Bizot
Federal Hwy. 15, 41 km W of Cd. Hidalgo, Sharp et al. & P. de la Varde, Rev. Bryol. Lichénol. 21: 10. 1952.
2191 (NY, TENN). Morelos: At Mirador, near Km 67 on Type: Mexico. Michoacán: Coru, 1907, Pringle 10438
Cuernavaca–Mexico City toll road, Hwy. 95D, 1952 km, p.p. (lectotype: PC!; isolectotypes: CANM!, FH!, US!,
Magill 2510 (NY, TENN). Nayarit: Mpio. Acaponeta, La both types designated by Buck, 1984a).
Ciénaga, ~16 km NW of Mesa del Nayar, 2593 km, Nor-
ris & Taranto 14421a (NY, TENN). Oaxaca: Dist. Putla, Erythrodontium densum var. brevifolium Cardot, Rev. Bryol. 37: 12. 1910;
~19 km S of Putla de Guerrero on road to Pinotepa Na- Bryosedgwickia densa var. brevifolia (Cardot) Bizot & P. de la Varde, Rev.
cional, Norris & Taranto 15876 (NY, TENN). Puebla: Bryol. Lichénol. 21: 10. 1952. Type: Mexico. Michoacán: Coru, 1907,
Vicinity of Puebla, Hacienda Santa Bárbara, 2160 m, Ar- Pringle 10438 p.p. (holotype: PC!; isotypes CANM!, FH!, NY!, US!).
sène 4516 (US). Durango, el Oro, Rangel & Arsène 4881
(US). Sinaloa: Along Hwy. 40, ~10 km W of Las Palmi- Erythrodontium imbricatifolium R. S. Williams ex Cardot, Rev. Bryol. 37:
tas, 2288 m, Norris 20448 (NY, TENN). GUATEMALA. 12. 1926; Platygyriella imbricatifolia (Cardot) Thér., Smithsonian Misc.
Baja Verapaz: Near Patal, 1585 m, Sharp 2906 (US); Chi- Collect. 78(2): 27. 1926. Type: Mexico. Jalisco: Barranca de Guadalajara,
maltenango: Along Río Guacalate, SE of Chimaltenango, Pringle 709 (holotype: PC; isotypes: FH!, NY!).
~1700 m, Standley 80038 (NY); Quezaltenango: Region of
Las Nubes, S of San Martín, Chile Verde, ~2250 m, Stand- Plants moderately robust in mostly dense mats,
ley 83644 (NY); Suchitepéquez: S slope of Volcán Santa brownish, rather shiny. Stems to ~10 cm but usually about
Clara, 1219 m, July 1948, Holdridge s.n. (NY). HONDU- half that, irregularly branched; stems in cross section with
RAS. Lempira: Montana de Celaque, along Río Arquegual, (2–)3–5 rows of small, thick-walled cells surrounding
7.5 km SW of Gracias, 14° 34'N, 88°39'W, 1400 m, Allen abruptly larger, thinner-walled cells, central strand lack-
11160 (MO, NY). EL SALVADOR. La Liberted: Cafetale ing; branches prostrate, slightly curved, terete; pseudo-
de Miguel Eduardo Araujo, 13°39'N, 89°22'W, 1000 m, paraphyllia filamentous; axillary hairs with 1(–2) short,
Monro et al. 2320 (MO); Santa Ana: Cerro Verde Na- brown basal cell(s) and 4–5 elongate, hyaline apical cells.
tional Park, 13°29'N, 88°32'W, 2365 m, Sidwell et al. 455 Asexual reproduction rarely by flagellate branches with
(MO). NICARAGUA. Jinotega: ~3.2 km NW of Jinotega, greatly reduced leaves. Leaves 0.9–1.2 mm long, imbri-
road to San Rafael del Norte, 1000 m, Croat 43029 (MO); cate, concave, ovate, abruptly acuminate; margins entire
Madriz: Cerro Quisuca, 13°31'N, 86°31'W, 1100–1250 to minutely serrulate, erect; costa short and double; cells
m, Stevens 16139 (MO); Managua: Sierra de Managua, linear, not shorter in the apex; alar cells quadrate to ob-
600–900 m, Garnier 707 (US). COSTA RICA. Alajuela: late, rarely reaching the costa; the decurrencies of mostly
Carrillos de Poas, cerca de Río Poas, Brenes 17401 (NY); 3–6 somewhat enlarged cells in 1–2 rows. Dioicous. Setae
San José: Santa María de Dota, 1600 m, Standley 43172 ~2 cm long, red; capsules 3–4 mm long; operculum short-
(NY). PANAMA. Chiriquí: Cerro Colorado, Chami Camp, rostrate; cells at mouth of urn rarely tardily deciduous
8°35'N, 81°44'W, 1050 m, Allen 5006A (MO); Coclé: El (not forming a true annulus); exostome teeth light yellow,
 number 93 • 57

cross-striolate below, smooth to faintly papillose at the of Volcán Zunil, 14°45'N, 91°29'W, 2442 m, Quedensley
extreme apex; endostome with a moderately high basal 2140 (BIGU, NY).
membrane; segments narrow, ± keeled, smooth to lightly
papillose, cilia none. Spores 14–19 µm in diameter. 10. Platygyrium Schimp. in Bruch,
Distribution and ecology: Mexico (where it is rela- Schimp. & W. Gümbel
tively common) and Guatemala; usually occurring on sunny
rocks, less often on trunks of deciduous trees, commonly in Platygyrium Schimp. in Bruch, Schimp. & W. Gümbel,
oak-pine forests, above 2000 m. Bryol. Eur. 5(46–47): 95.1851, nom. cons. Type: P. re-
Illustrations in publications: Fig. 162A, B in Bar- pens (Brid.) Schimp.
tram (1949: 365); Pl. 758 in Buck (1994: 1021).
Discussion: This is a very attractive moss, with a Plants small to medium-sized, in creeping mats, glossy,
beautiful brownish coloration. It can be distinguished from yellowish green to brownish green or darker. Stems subpin-
P. densa by its rupestral habitat, greater stature, imbricate nately branched; branches short to elongate, erect to pros-
and abruptly tapered leaves, and larger decurrencies with trate, straight to curved at tips, loosely foliate to julaceous,
less markedly differentiated cells. There are peristomial fea- in cross section without a hyalodermis, central strand lack-
tures separating the two as well, but neither is commonly ing; pseudoparaphyllia filamentous to narrowly foliose; ax-
found fertile. Confusion is more likely with Erythrodon- illary hairs composed of 1–2 short, brown basal cells and
tium longisetum (Hook.) Paris. However, in Erythrodon- 1–5 elongate, hyaline apical cells. Stem and branch leaves
tium longisetum the leaves are oblong rather than ovate, similar, ovate, imbricate to homomallous, acute to acumi-
the decurrencies are quite broad, the inflorescences are au- nate, flat to somewhat plicate, concave; margins mostly
toicous, and the setae are yellow rather than red. Eryth- entire, rarely subserrulate apically, somewhat to strongly
rodontium densum var. brevifolia, characterized only by recurved, especially below; costa short and double to ab-
flagellate branches, is a rare and unimportant expression sent; cells elongate, smooth; alar cells numerous, quadrate,
of the species. thick-walled. Asexual reproduction by flagellate branch-
Selected specimens examined: MEXICO. Chihua- lets at branch apices. Dioicous, but often fertile. Perichae-
hua: N of Barranco de Batopilas between Divisaero and tia small, inconspicuous. Setae straight, elongate, smooth,
Quirire, Bye 6346d (COLO, NY). Durango: Below Salto, twisted when dry. Capsules erect, long-exserted, cylindrical
2532 m, Sharp 1826 (NY, TENN). Guerrero: Near Taxaco, to subcylindrical, symmetric or slightly asymmetric; exo-
Welch 20364 (MICH, NY). Jalisco: Guadalajara, Pringle, thecial cells oblong-rectangular, stomata phaneroporous,
Plantae Mexicanae 15074 (US); 14 km al W de Cuautla, few, confined to base of urn; annulus differentiated in 1–3
por la brecha Ayutla–Mascota, Guzmán 912 (MEXU, rows of cells; operculum obliquely rostrate; peristome dou-
NY). México: Mpio. Ocuilán, Barranca de Mexicapa, Km ble, exostome pale, yellow-orange, teeth finely papillose to
14 on road from Sta. Mónica de Ocuilán to Cuernavaca, papillose-striolate on outer surface, bordered, smooth at
18°58'N, 99°20'W, 2200–2350 m, Buck 28137 (NY). Mi- tips, weakly trabeculate at back; endostome segments nearly
choacán: 6 km NE of Zinapécuaro, 19°54'N, 100°47'W, as long as teeth, brown, narrowly linear, perforate or not,
2020 m, Delgadillo 4980 (MEXU, NY); vicinity of More- sometimes adherent to teeth, cilia rudimentary to lacking.
lia, Jesús del Monte, 2000 m, Arsène 7602, 7603, 7606, Spores green, spherical. Calyptra cucullate, naked.
7618, 7624 (US); Carríndapaz, 2100 m, Arsène 7702, Discussion: Platygyrium is a genus of about eight
7953 (US); Cerro San Miguel, 2200 m, Arsène 5101 (US); species, mostly distributed in Asia. In North America there
Bosque San Pedro, 1950 m, Arsène 4584 (US). Morelos: is the common and widespread P. repens (Brid.) Schimp.,
oberh. D. neuen Autobahn zw. Cuernavaca u. Mexico City, but it does not reach into our flora area. Only a single spe-
2050–2100 m, Düll 199 (MICH, NY). Oaxaca: 22.4 km cies is in our region. The genus is similar to and often con-
N on Hwy. 175 from jct. with Hwy. 190, near La Punte, fused with Homomallium and Pylaisia. Fertile specimens of
NE of Oaxaca, 17°12'N, 96°05'W, 2532 m, Vitt 17624 Homomallium are easily distinguished because they have
(ALTA, NY). Puebla: Honey Station, Pringle 15128 (US); arcuate, asymmetric capsules, whereas Platygyrium and
Zacatlán, Orcutt 1262 (NY). San Luis Potosí: Above Xil- Pylaisia have erect, symmetric capsules. Pylaisia can be
itl, 763 m, Sharp 5906 (NY,TENN). Sinaloa: Along Hwy. distinguished from Platygyrium by the smooth exostome
40, ~3 km E of Las Palmitas, 2288 m, Norris 20598 (NY, teeth and the shorter operculum. In Platygyrium the long,
TENN). GUATEMALA. Quetzaltenango: Mpio. Zunil, obliquely rostrate operculum is in striking contrast to that
road to Fuentes Georginas, 4 km S of Zunil, NW slopes of Pylaisia, which has a conic operculum with only a short,
58 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

straight rostrum. These three genera have more or less ho- with subquadrate alar cells, make the species distinctive.
momallous leaves and numerous alar cells. The epiphytic The axillary brood branchlets are less conspicuous and
habitat and axillary brood branchlets distinguish Platygy- less common than in the more northerly P. repens, but
rium from Pylaisia, as well as from Homomallium. they are usually present.
Homomallium mexicanum Cardot is similar to P. fus-
Platygyrium fuscoluteum coluteum. As mentioned above, the capsules (if the plants
are fertile) in Homomallium are curved. Vegetatively, the
Platygyrium fuscoluteum Cardot, Rev. Bryol. 37: 49. two may be separated by the brood branchlets in Platygy-
1910. Type: Mexico. Chihuahua: environs de Madera, rium and by the homomallous leaves in Homomallium.
1908, Palmer 445 p.p. (holotype: NY!). From Pylaisia, Platygyrium fuscoluteum can be separated
by the straight branches and straight to mildly homomal-
Regmatodon fusco-luteus Schimp. ex Besch., Mém. Soc. Sci. Nat. Cherbourg lous leaves, in addition to the brood branchlets; some
16: 232. 1872; Platygyrium fuscoluteum (Besch.) Cardot, Rev. Bryol. 38: Pylaisia species have curved branches and strongly ho-
40. 1911, hom. illeg., non Cardot, 1910. Type: Mexico. Orizaba, F. Müller momallous to falcate leaves.
(holotype: BM!). Specimens examined: MEXICO. Coahuila: Made-
ras del Carmen, 2500 m, Riskind 1990a (NY). Michoacan:
Plants in relatively robust mats, yellowish brown. Stems Between Chilchota and Zacapu, 2135 m, Weber & Cha-
to ~5 cm long, irregularly to regularly 1–2-pinnately branched, rette B-9854 (NY). Veracruz: Just S of the city of Xalapa,
in cross section without a hyalodermis, central strand lack- Rancho Guadeloupe, 1300 m, Buck 35323 (NY). Zacate-
ing; branches 3–10 mm long, rarely longer, straight to cas: Sierra de los Morones, 37 km W de Jalpa, Cárdenas
curved; axillary brood branchlets occasionally near branch 845 (MEXU, NY).
tips. Leaves mostly 1.3–1.6 mm long, 0.3–0.4 mm wide, ob-
long-lanceolate, acuminate, usually homomallous but some- 11. Pseudotaxiphyllum Z. Iwats.
times straight, slightly plicate; margins entire, conspicuously
recurved in lower half, sometimes almost to leaf apex; me- Pseudotaxiphyllum Z. Iwats., J. Hattori Bot. Lab. 63:
dian cells mostly 50–75 × 4–6 µm, slenderly elongate; alar 448. 1987. Type: Isothecium elegans Brid., Bryol. Univ.
cells 5–12 up the margins, subquadrate, sometimes colored. 2: 356. 1827.
Perichaetial leaves oblong-lanceolate, acuminate, ± plicate;
margins serrulate above. Setae 8–20 mm long, reddish. Cap- Plants in medium-sized, often complanate-foliate,
sules 2–2.5 mm long, pale yellow-brown, cylindrical, nearly thin to dense mats, dull to glossy, light green to yellow-
symmetric; operculum slenderly and mostly obliquely long- ish green. Stems creeping, simple or often sparingly and
rostrate. Annulus of 1–2 rows of cells. Exostome teeth cross- irregularly branched, cortical cells small, thick-walled,
striolate below, smooth to lightly roughened above, strongly central strand sometimes present; rhizoids smooth, in
trabeculate at back; endostome segments brownish, smooth; clusters below leaf insertions; pseudoparaphyllia lacking;
cilia rudimentary, single. Spores 15–20 µm in diameter, finely axillary hairs with a single short-rectangular, light brown
granular. Calyptra 2.0–2.5 mm long. basal cell and 2–3, elongate, hyaline distal cells. Asexual
Distribution and ecology: Mexico; also in the reproductive bodies sometimes present in clusters at stem
southwestern United States; occurring on tree trunks and apices or below in leaf axils, 0.1–0.5 mm long, elongate,
logs, rarely on soil in coniferous and deciduous forests at multicellular, smooth-celled, twisted-vermiform, with 1–5
altitudes of 2200–2800 m. acute teeth, or resembling the parent plant but smaller,
Illustrations in publications: Pl. 751 in Ando (1994: 0.8–1.2 mm long, bearing reduced leaves from apex to
1011). base of propagula. Stem and branch leaves similar, some-
Discussion: This species is confined to Mexico what rigid, crowded and imbricate, erect-spreading to
and the southwestern United States (Arizona, New Mex- squarrose, sometimes upturned-homomallous, occasion-
ico, and Texas). It mostly occurs at higher elevations on ally complanate, smooth or undulate, flat or concave,
tree trunks and logs in hardwood and mixed conifer-hard- symmetric to asymmetric, nondecurrent, lanceolate, ovate,
wood forests at 1300–2500 m. The glossy yellowish brown ovate-lanceolate or oblong-lanceolate, acute, sometimes
plants with homomallous leaves and long, slender apices abruptly so, to acuminate; margins plane to erect, serrate
on elongate branches are characteristic. The strongly re- to serrulate; costa lacking or short and double; cells often
curved leaf margins and somewhat plicate leaves, coupled flexuose, firm-walled, linear-fusiform, smooth or apical
 number 93 • 59

cells sometimes prorulose at upper ends on dorsal surface, groups of 1–3. Spores spherical to ovoid, minutely papil-
walls not pitted; alar regions not or distinctly differenti- lose. Calyptra cucullate, smooth, naked.
ated with numerous quadrate to short-rectangular cells. Distribution: Pseudotaxiphyllum is a genus of 11
Dioicous or sometimes autoicous, usually sterile; perigo- species occurring in terrestrial habitats in temperate, sub-
nia along stems; perichaetia at base of stems, bracts small, tropical, and tropical regions.
lanceolate to ovate, acuminate to abruptly filiform-acumi- Discussion: Pseudotaxiphyllum is a segregate of
nate; margins plane. Setae smooth, elongate, straight or the genus Isopterygium described by Iwatsuki (1987). The
sometimes curved, twisted, red to reddish brown; capsules genus is distinguished by the medium-sized, simple or spar-
cernuous to pendulous, straight to subarcuate, dark brown ingly branched, dull to glossy plants, the small, thick-walled
to dark red, smooth to wrinkled, contracted below mouth cortical stem cells, the smooth rhizoids below the leaf in-
and wrinkled at neck when dry; operculum conic to short- sertions, the absence of pseudoparaphyllia, the nondecur-
rostrate, shorter than urn; annulus present, deciduous, of rent, serrate to serrulate leaves, the occasional presence of
2–3 rows of cells; peristome double, exostome teeth cross- asexual reproductive bodies, the dioicous or sometimes au-
striolate below, papillose above, bordered, trabeculate toicous sexual condition, and the presence of an annulus.
at back; endostome with a high to low basal membrane, Sex organs and sporophytes have never been seen on plants
keeled segments, and cilia shorter than the segments, in of three of the four species present in Latin America.

KEY TO THE SPECIES OF PSEUDOTAXIPHYLLUM

1. Median leaf cells broad, 5–9 µm wide; propagula lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Leaves long-acuminate, upturned-homomallous; leaf margins plane when dry; alar regions poorly differentiated, a few
short-rectangular cells present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. P. homomallifolium
2. Leaves acute to short-acuminate, not upturned-homomallous; leaf margins involute when dry; alar regions distinctly dif-
ferentiated, numerous quadrate cells present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. P. richardsii
1. Median leaf cells narrow, usually 4–7 µm wide; propagula usually present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Leaves asymmetric, often cultriform; propagula clustered at stem apices and in upper leaf axils, elongate, twisted-
vermiform, with 1–5 acute teeth at apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. P. distichaceum
3. Leaves symmetric, never cultriform; propagula usually clustered in leaf axils below stem apices, resembling parent plant
but smaller, bearing reduced leaves from apex to base of propagula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. P. elegans

1. Pseudotaxiphyllum distichaceum Taxiphyllum howellianum H. A. Crum & L. E. Anderson, J. Elisha

Mitchell Sci. Soc. 74: 38. 1958. Type: North Carolina, Jackson Co., W
FIGURE 26 of Cashiers, below Cowee Gap, Howell Tract, 1158 m, Anderson 11071
(isotype: CANM!).
Pseudotaxiphyllum distichaceum (Mitt.) Z. Iwats., J. Hat-
tori Bot. Lab. 63: 449. 1987. Stereodon distichaceus Mitt., Plants in thin to loose mats, glossy, light green to
J. Proc. Linn. Soc., Bot., Suppl. 1: 105. 1859; Isopterygium yellowish green. Stems to 2 cm long, 1.0–3.5 mm wide,
distichaceum (Mitt.) A. Jaeger, Ber. Thätigk. St. Gallischen complanate-foliate, simple or irregularly branched. Asex-
Naturwiss. Ges. 1876–77: 439. 1878. Type: Nepal, India, ual reproductive bodies often present, 0.1–0.5 mm long,
Wallich s.n. (lectotype: NY!, here designated). clustered in leaf axils at or near stem apices, yellowish
green, elongate, twisted-vermiform, composed of 2–4
Plagiothecium subfalcatum Austin, Musci Appalach. 366. 1870; Isopteryg- layers of smooth cells, with 1–5 acute, erect teeth at apex.
ium subfalcatum (Austin) A. Jaeger, Ber. Thätigk. St. Gallischen Naturwiss. Leaves 1.0–1.8 mm long, 0.3–0.6 mm wide, semiflaccid to
Ges. 1876–77: 438. 1878. Type: Mountains of New Jersey and New York, rigid, distant, squarrose, complanate, smooth, ovate- or
Austin’s, Musci Appalach. 366 (isotypes: CANM!, DUKE!, NY!, US!). oblong-lanceolate, often cultriform, asymmetric, acumi-
nate; margins plane or narrowly recurved at base, serrate
Isopterygium poasense Renauld & Cardot, Bull. Soc. Roy. Bot. Belgique to serrulate above, serrulate to entire below; costa usually
41(1): 141. 1904 [1905]. Type: Costa Rica, Potrero del Alto, volcán de Poás, strong, short and double; median cells 56–120 × 4–7 µm,
2463 m, Pittier 5791 (isotypes: FH!, US!). smooth; apical cells sometimes prorulose at upper ends
60 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 26. Pseudotaxiphyllum distichaceum. A. Habit. B. Enlargement of stem apex showing asexual reproductive bodies. C. Leaves. D. Api-
cal leaf cells. E. Median leaf cells. F. Marginal leaf cells. G. Cells of alar region. H. Asexual reproductive bodies. All illustrations drawn from
Juárez G. 1630 (CANM).

on dorsal surface; alar cells undifferentiated or 1–3 mar- drical with a distinct neck; operculum conic. Crum and
ginal cells quadrate to rectangular. Sex organs and sporo- Anderson (1981) report spores 8–10 µm in diameter, mi-
phytes not seen on Latin American plants. Reported from nutely roughened.
India (Buck, 1998) to be dioicous? (no perigonia seen) Distribution and ecology: Rare or seldom col-
or monoicous (Gangulee,1980). Buck also reported setae lected; known from Mexico, Guatemala, Honduras, Costa
~1.5 cm long, reddish, curved just below the urn; capsules Rica, Dominican Republic, Colombia, Venezuela, Ecua-
~1.1–1.5 mm long, horizontal, symmetric, broadly cylin- dor and Brazil (Figure 27); also known from southeastern
 number 93 • 61

FIGURE 27. Distribution of Pseudotaxiphyllum distichaceum (●), P. homomallifolium (◆), and P. richardsii (▲) in Latin America.

Canada, eastern United States, Asia, and Australia; occur- gula are present, their location on the stems as well as their
ring on shaded, moist to somewhat dry soil or soil over morphology are distinctly different in the two species.
rocks and boulders, limestone cliffs, often on humid, steep Pseudotaxiphyllum distichaceum has propagula that are
ravines, 0–4040 m. elongate, twisted-vermiform with 1–5 acute teeth at the
Discussion: Pseudotaxiphyllum distichaceum is apices, and they occur in the leaf axils at or near the stem
best recognized by the distant, squarrose, complanate, apices. On the other hand, P. elegans has propagula in the
ovate- or oblong-lanceolate, often cultriform, asymmetric, leaf axils that always occur below the stem apices and they
acuminate leaves with serrate to serrulate margins, the nar- resemble the parent plant, being much smaller, possessing
row median leaf cells, the poorly differentiated alar cells, small leaf-like structures along their length.
sometimes 1–3 quadrate to rectangular cells on the mar- Specimens examined: MEXICO. Jalisco: Nevado
gins, and the usual presence of clusters of elongate, twisted- de Colima, 3940–3970 m, De Luna 509a (CANM). Ve-
vermiform propagula with 1–5 acute teeth at apices in the racruz: 10 km NE of outskirts of Huayacocotla, Juárez
upper leaf axils. The Mexican plants studied often lacked G. 1630 (CANM). Zacatecas: Cerro de la Bufa, 2700
the characteristic propagula that occur clustered in the m, Cárdenas 339 (CANM, MEXU). GUATEMALA.
upper leaf axils and, when present, make this one of the Quezaltenango: Canyon above Baños Georginas, 8 km
easiest plants to recognize. Most of the plants seen else- above Zunil, ~2470 m, ~14°46'N, 91°28'W, Gereau &
where in Latin America, however, have propagula. Martin 1876 (NY). HONDURAS. Lempira: Montana de
Pseudotaxiphyllum distichaceum is easily confused Celaque, Cerro Mojon, 13 km SW of Gracias, 14°32'N,
with P. elegans, especially when propagula are not pres- 88°41'W, Allen 12271 (MO). COSTA RICA. Cartago:
ent. The asymmetric, often cultriform leaves of P. dis- ~97 km S of Cartago, behind Hotel La Georgina on Intera-
tichaceum will distinguish it from P. elegans, which has merican Hwy., 2900 m, Bowers 830-1A (TENN). San José:
symmetric leaves that are never cultriform. When propa- Cerro de Zurqui, N of San Luis Norte, 17 km NNE of San
62 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

José, 10°03'N, 84°01'W, Crosby 9686 (MO). Parque Na- not seen on Latin American plants. North American plants
cional Volcán Poás, Morales 538 (CANM). DOMINICAN reported (Ireland, 1969) to be dioicous. Setae 1.0–2.5 cm
REPUBLIC. La Vega: Vicinity of pyramids, 13.8 km S of long, dark red; capsules 1–2 mm long, cernuous to pendu-
Valle Nuevo, 44.7 km S of Constanza, 2256 m, Shaw 5642 lous, straight or subarcuate, oblong-ovoid to ovoid, wrin-
(NY). San Juan–Santiago: Parque Nacional J. Armando kled and contracted below mouth when dry; operculum
Bermúdez, on the Loma La Pelona, 3100–3200 m, Zanoni conic to short-rostrate, 0.4–0.7 mm long; annulus present.
et al. 42050 (NY). COLOMBIA. Boyacá: Mpio. de Du- Crum and Anderson (1981) reported spores 7–10 µm in
itama, around Páramo “La Rusia,” 2620 m, Ireland 23602 diameter, minutely roughened.
(CANM, COL). Cundinamarca: Mpio. de Subachoque and Distribution and ecology: Very rare; known from
Supata, N of Bogota, “El Tablazo,” 3500 m, Ireland 23422 one locality on the Falkland Islands, Argentina, where it oc-
(CANM); Páramo de Palacio, between Mina de Cal and curred on rocks, and from one in Chile, where it occurred
Río Chuza, Páramo Seco, Florschütz 3883 (U). Santander: on a limestone cliff, both localities apparently at sea level
La Sierra, along road between Párama de la Rusia and Vi- (Figure 29); known elsewhere from Canada, United States,
rolin, ~2600 m, Lewis 88-1350 (CANM, LPB). VENEZU- Europe, and Iceland; reported from Asia and Africa (van
ELA. Mérida: Páramo de los Conejos, Sierra del Norte, der Wijk et al., 1964).
3300 m, Griffin & Lopez PV-611 (CANM). ECUADOR. Discussion: This species is recognized by the close
Imbabura: E side of Cerro Imbabura above La Esperanza, to somewhat distant, erect-spreading, lanceolate, ovate- or
ENE of Otavalo, 4040 m, Lewis 78-2969B (F). BRAZIL. oblong-lanceolate, asymmetric, acuminate leaves with ser-
Paraná: Ponta Grossa, Park Vila Velha, 900 m, Schäfer- rulate to strongly serrate margins, the narrow median leaf
Verwimp & Verwimp 9201 (CANM). cells, the poorly differentiated alar cells, 1–3 marginal cells
quadrate to rectangular, and the usual presence of clusters
2. Pseudotaxiphyllum elegans of propagula in the leaf axils below stem apices, the propa-
gula resembling the parent plant but smaller, bearing re-
FIGURE 28 duced leaves from apex to base. Pseudotaxiphyllum elegans
is morphologically close to P. distichaceum. For distinctions
Pseudotaxiphyllum elegans (Brid.) Z. Iwats., J. Hattori see the discussion under that species.
Bot. Lab. 63: 449. 1987. Hypnum elegans Hook., Musci The Halle and Skottsberg 873 (UPS) specimen from
Exot. 1: Pl. 9. 1818, hom. illeg.; Isothecium elegans the Falkland Islands has been reported as Isopterygium
Brid., Bryol. Univ. 2: 356. 1827; Isopterygium elegans elegans var. nanum (Jur. in Milde) A. Jaeger (Cardot and
(Brid.) Lindb., Not. Sällsk. Fauna Fl. Fenn. Förh. 13: Brotherus, 1923; Matteri, 1986; Matteri and Schiavone,
416. 1874; Plagiothecium elegans (Brid.) Schimp., Syn. 2002), but an examination of the specimen fits our con-
Musc. Eur. ed. 2, 698. 1876. Type: Canada, British Co- cept of P. elegans var. elegans. The type of the var. nanum
lumbia, Vancouver Is., Nootka, Menzies 35 (holotype: has not been examined, but judging from Juratzka’s
BM!; isotype: NY!). (1864) description of the type (as Plagiothecium nanum
Jur. in Milde), it falls within the range of variation that we
Plants in thin to dense mats, glossy, dark green to yel- associate with P. elegans var. elegans. Therefore, we are
lowish green. Stems to 1 cm long, 1–2 mm wide, simple doubtful that the variety is worth taxonomic recognition,
or irregularly branched. Asexual reproductive bodies but the final decision will have to wait until the type of the
usually present, clustered in leaf axils below stem apices, var. nanum is examined.
0.8–1.2 mm long, yellow to green, smooth-celled, resem- Specimens examined: CHILE: Bío-Bío: Arauco
bling the parent plant but smaller, bearing reduced leaves Prov., 2 km N of Casa de Piedra Beach, a.s.l.[at sea level],
from apex to base of propagula. Leaves 0.5–1.2 mm long, 38°28'S, 73°31'W, Ireland & Bellolio 33252 (CONC, MO).
0.2–0.6 mm wide, semiflaccid to rigid, close to somewhat ARGENTINA: Falkland Islands: West Falkland, Warrah
distant, erect-spreading, somewhat concave, smooth, lan- River, 17 Dec 1907, Halle & Skottsberg 873 (UPS).
ceolate, ovate- or oblong-lanceolate, symmetric, acumi-
nate; margins plane, serrulate to strongly serrate above, 3. Pseudotaxiphyllum homomallifolium
serrulate to entire below; costa usually strong, short and
double; median cells 48–100 × 4–7 µm, smooth; apical FIGURE 30
cells sometimes minutely prorulose at upper ends on dor-
sal surface; alar cells undifferentiated or 1–3 quadrate to Pseudotaxiphyllum homomallifolium (Redf.) Ireland, Cal-
rectangular cells on margins. Sex organs and sporophytes dasia 16(78): 267. 1991. Isopterygium homomallifolium
 number 93 • 63

FIGURE 28. Pseudotaxiphyllum elegans. A. Habit. B. Enlargement of portion of stem showing asexual reproductive bodies. C. Leaves. D. Api-
cal leaf cells. E. Median leaf cells. F. Marginal leaf cells. G. Cells of alar region. H. Asexual reproductive bodies. All illustrations drawn from
Halle & Skottsberg 873 (UPS).

Redf., Bryologist 76: 440. 1973. Type: Texas, Kimble 1.2 mm long, 0.3–0.5 mm wide, semiflaccid to rigid, dis-
Co., along North Llano River, ~½ mi. S of Roosevelt, tant to close and overlapping, erect-spreading, sometimes
Redfearn 27170 (holotype: NY!; isotype: MICH!). upturned-homomallous, occasionally complanate, smooth,
lanceolate to ovate-lanceolate, symmetric, long-acuminate;
Plants in thin to dense mats, glossy, yellowish green. margins plane, serrulate nearly to base; costa weak, short
Stems to 1.5 cm long, 1–2 mm wide, simple or irregularly and double or lacking; median cells 60–120 × 5–9 µm,
branched. Asexual reproductive bodies lacking. Leaves 0.8– smooth; apical cells smooth; alar cells poorly differentiated,
64 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 29. Distribution of Pseudotaxiphyllum elegans in Latin America.

a few short-rectangular cells often present. Sporophytes not Specimens examined: MEXICO. Sonora: Loop of
seen on Latin American plants. North American plants re- the Río de Bavispe, cerro del Capulin, NW of Aribabi, 1875
ported (Ireland, In press a) to be autoicous. Setae 0.8–1.6 cm m, Harvey 1704 (MICH); Cañon de Huépari, N of Aribabi,
long, yellow to reddish; capsules 1.4–1.7 mm long, erect to 1350 m, Harvey 1747 (MICH).
horizontal, slightly cernuous, ellipsoid, contracted below
the mouth when dry; operculum high-conic, 0.4–0.6 mm 4. Pseudotaxiphyllum richardsii
long, high-conic to short-rostrate; annulus present, decidu-
ous, of 2–3 rows of cells. Spores 9–16 µm in diameter. FIGURE 31
Distribution and ecology: Rare; known in Latin
America only from the state of Sonora, Mexico (Figure 27); Pseudotaxiphyllum richardsii (E. B. Bartram) Ireland,
occurring on rocks and under rock ledges, 1350–1875 m; Caldasia 16(78): 267. 1991. Isopterygium richardsii
also known to occur in the southwestern United States (Ari- E. B. Bartram, Bryologist 44: 64. 1941. Type: Mexico,
zona, New Mexico, and Texas). Sonora, Imuris, along Río de los Alisos, Richards &
Discussion: Pseudotaxiphyllum homomallifolium Drouet 757 (holotype: F!; isotypes: DUKE!, F!, FH!,
is distinctive because of its long-acuminate, upturned- MICH!, NY!).
homomallous leaves with serrulate margins, its broad
median leaf cells, its few differentiated cells in the alar re- Plants in thin mats, somewhat glossy to dull, light
gions, and its autoicous sexual condition. green to yellowish green. Stems to 1 cm long, 1–2 mm
Although foliose pseudoparaphyllia were attributed wide, simple or irregularly branched. Asexual reproduc-
to P. homomallifolium when Redfearn (1973) described tive bodies lacking. Leaves 0.5–1.0 mm long, 0.3–0.5 mm
the species, we do not believe they should be classified as wide, flaccid, close, erect, indistinctly complanate when
such structures. The multicellular structures rarely found dry, more noticeably so when moist, concave, smooth,
on the stems seem to be part of a developing branch pri- ovate to ovate-lanceolate, symmetric, acute to short-
mordium. They do not appear to be distinctly separated acuminate; margins plane or strongly involute when dry,
from the rest of the primordium, sometimes not even evi- serrate to serrulate nearly to base; costa weak, short and
dent on the stems at the bases of mature branches like double; median cells 47–75 × 5–9 µm, smooth; apical
typical pseudoparaphyllia. cells smooth; alar cells distinctly differentiated, numerous
 number 93 • 65

FIGURE 30. Pseudotaxiphyllum homomallifolium. A. Habit. B. Leaves. C, D. Apical leaf cells. E. Marginal leaf cells. F. Median leaf cells. G.
Cells of alar region. All illustrations drawn from Harvey 1704 (MICH).

quadrate to short-rectangular cells present. Autoicous. Sonora (Figure 27); occurring on soil over rock in ravines
Setae 0.6–0.7 cm long, reddish; capsules ~1 mm long, in- and near rivers, ~1300 m.
clined, nearly straight, ellipsoid, not or scarcely contracted Discussion: This species is recognized by the
below the mouth when dry; operculum high-conic; annu- combination of small (0.5–1.0 mm long), ovate to ovate-
lus not seen. Spores and calyptra not seen. lanceolate, acute to short-acuminate leaves with serrate
Distribution and ecology: Rare; endemic to Mex- to serrulate margins that are strongly involute when dry;
ico, where it is known only from the states of Nayarit and broad median leaf cells; distinctly differentiated alar regions
66 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 31. Pseudotaxiphyllum richardsii. A. Habit. B. Enlargement of apical portion of stem (wet). C. Enlargement of apical portion of stem
(dry). D. Leaves. E. Apical leaf cells. F. Median leaf cells. G. Marginal leaf cells. H. Cells of alar region. I. Cross section of stem. A, D–I from
Richards & Drouet 757 (F, holotype); B, C from Harvey 1669a (MICH).

composed of numerous quadrate to short-rectangular cells; Richards, Drouet & Lockhart 722 (F, FH, MICH, NY,
and autoicous sexual condition. US); on side of Cerro Agua Caliente, along Río de los Al-
Specimens examined: MEXICO. Nayarit: Mpio. isos, near Imuris, Richards & Drouet 749 (F, FH, MICH,
Acaponeta, near Labra along trail W of Jesús Maria, Nor- NY, TENN, US); on side of mountain just W of Ala-
ris & Taranto 14132 (MICH, TENN). Sonora: 15.2 km mos, Richards & Drouet 780 (F, MICH, US); Cañon de
NE of Baviácora, Richards, Drouet & Lockhart 676 (F, Huépari, N of Aribabi, 1310 m, Harvey 1669a (MICH);
FH, MICH, TENN, US); canyon 24.8 km SW of Cumpas, Cajón de la Higuera, near Moctezuma, White 349, 362,
 number 93 • 67

366a, 367, 376, 390 (MICH); Río Moctezuma, Mocte- Puiggariopsis aurifolia
zuma, White 328, 329, 331, 348, 349 (MICH); Arroyo el
Sance, near Moctezuma, White 318 (MICH). Puiggariopsis aurifolia (Mitt.) Menzel, J. Hattori Bot.
Lab. 71: 239. 1992; Ctenidium aurifolium Mitt., J. Linn.
Pseudotaxiphyllum Excluded Taxa Soc., Bot. 12: 509. 1869; Puiggariella aurifolia (Mitt.)
Broth. in Engl. & Prantl, Nat. Pflanzenfam. 1(3): 1047.
Pseudotaxiphyllum machrisianum (H. A. Crum) Ireland, Cal- 1908. Type: Brasilia tropica, Burchell 2275 (holotype:
dasia 16(78): 267. 1991. Taxiphyllum machrisianum H. A. NY!; isotype: BM!).
Crum, Los Angeles County Mus. Contr. Sci. 18: 5. 1957;
Isopterygium machrisianum (H. A. Crum) Ireland, Bry- Pilotrichum disciflorum Geh. & Hampe, Flora 64: 403. 1881; Puiggariella
ologist 87: 356. 1985; Symphyodon machrisianus (H. A. disciflora (Geh. & Hampe) Broth. in Engl. & Prantl, Nat. Pflanzenfam. 1(3):
Crum) W. R. Buck & Ireland, Bryologist 95: 433. 1992. 1047. 1908. Type: Brazil. Apiahy, Apr 1879, Puiggari 618 (holotype: BM!;
Type. Brazil, 4 km N of Veadeiros, Chapada dos Veadeiros, isotype: H!).
Goiás, Dawson 14743a (isotypes: CANM!, FH!).
Papillaria rutenbergiacea Müll. Hal., Hedwigia 40: 90. 1901. Type: Brazil.
This species was sterile when it was described from Rio de Janeiro, ad truncos arborum montis Tijuca, Nov 1893, Ule 1697 (iso-
a Brazilian collection as a Taxiphyllum by Crum (1957), types: H!, US!).
who at that time expressed some doubts about its place-
ment in the genus. All subsequent specimens examined by Plants in medium-sized to robust mats, yellowish
the authors from Mexico, Costa Rica, Nicaragua, Colom- green. Stems 2–3(–5) cm long, ascending to arching, rarely
bia, Venezuela, and Brazil were also sterile. However, the creeping, subpinnately branched, sometimes becom-
junior author discovered that the gametophytes of T. ma- ing flagelliform; stems in cross section lacking a hyalo-
chrisianum and Symphyodon americanus were similar. dermis, with a few rows of small, thick-walled cells and
Steere (1982) described S. americanus from plants with large, thin-walled cells in center, central strand present;
sporophytes from Colombia and sterile plants from Ecua- rhizoids finely papillose, reddish brown, on ventral side
dor. The sporophytes of this species have capsules covered of stem; paraphyllia present, narrowly lanceolate; pseu-
with single-celled spines and setae that are strongly mam- doparaphyllia foliose; axillary hairs reddish brown, con-
millose or papillose a short distance below the capsules. sisting of 2–4, short basal cells and (1–)2–3(–5) elongate,
These features are both typical of species of Symphyodon distal cells. Asexual reproductive bodies lacking. Branch
but not of the smooth capsules and setae possessed by and stem leaves differentiated; stem leaves 1.9–2.2 mm
Taxiphyllum, Pseudotaxiphyllum, or Isopterygium, where long, 0.9–1.3 mm wide, imbricate, julaceous, erect to
T. machrisianum had been placed at one time or another spreading, weakly secund, ovate, gradually acuminate
(see list of synonyms above). Because the specific name with the acumen twisted, plicate, shortly decurrent; mar-
of the Taxiphyllum antedated that of the Symphyodon, it gins plane, serrulate throughout; costa double, separated
necessitated a new combination, S. machrisianus (H. A. at base, to ¼ the leaf length; median cells (30–)45–55
Crum) W. R. Buck & Ireland (Buck and Ireland, 1992). (–65) × ~2.5 µm (lumen; ~5 µm including walls), linear,
However, only two years later, Churchill (1994) discov- thick-walled, porose, upper cells prorulose at upper ends
ered that the species was synonymous with S. imbricat- on abaxial surface, becoming shorter, broader, thicker-
ifolius (Mitt.) S. P. Churchill from the West Indies and walled and smooth toward the insertion; alar cells deeply
Central and South America. excavate, weakly differentiated, rectangular, thick-walled,
porose. Branch leaves 1.3–1.9 mm long, 0.6–0.8 mm wide,
12. Puiggariopsis Menzel smaller and narrower than stem leaves. Dioicous. Perichae-
tial leaves erect, oblong-obovate, abruptly acuminate, not
Puiggariopsis Menzel, J. Hattori Bot. Lab. 71: 239. 1992, plicate; margins weakly serrulate above. Setae 15–18 mm
nom. nov.; Puiggariella Broth. in Engl. & Prantl, Nat. long, smooth, reddish brown; capsules 1.0–1.5 mm
Pflanzenfam. 1(3): 1046. 1908, hom. illeg., non Speg., long, 0.8–1.0 mm wide, inclined to horizontal, ovoid,
Anales Soc. Ci. Argent. 12: 99. 1881 (= Strigula Fr., strongly constricted below the mouth when dry; opercu-
Strigulaceae). Type: Puiggariella aurifolia (Mitt.) Broth. lum ~1.2 mm long, obliquely conic-rostrate; exostome
teeth yellowish, with a zigzag center line, on front surface
This is a monospecific genus with the characters of cross-striolate with overlying papillae below, irregularly
the species. striolate with overlying papillae toward middle, papillose
68 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

above; endostome almost smooth, with a low basal mem- Thedenia Schimp. in Bruch, Schimp. & W. Gümbel, Bryol. Eur. 5: 83.
brane, nearly as high as exostome teeth, keeled, narrowly 1852, hom. illeg. Type: Thedenia suecica Schimp. in Bruch, Schimp. &
perforate, cilia 1–2, rudimentary or absent. Spores too few W. Gümbel.
to accurately determine size, appearing spherical, finely
papillose. Calyptrae cucullate, densely hairy. Giraldiella Müll. Hal., Nuovo Giorn. Bot. Ital. n.s. 5: 192. 1898. Type:
Distribution and ecology: Known from Mexico to Giraldiella levieri Müll. Hal.
Nicaragua, southern Brazil, Venezuela, and Peru; usually
growing on tree trunks and branches, less often on rocks or Plants small to large, in often extensive mats, yellow-
soil, at (200–)700–1000(–1800) m. ish green to dark green. Stems creeping, regularly or irreg-
Illustrations in publications: Fig. 780 (as Puiggari- ularly pinnately branched; stems in cross section lacking a
ella aurifolia (Mitt.) Broth.) in Nishimura and Ando (1994: hyalodermis, with small, thick-walled cells around large,
1052); Fig. 182E–H (as Puiggariella aurifolia (Mitt.) Broth.) thin-walled cells, central strand small; rhizoids smooth,
in Bartram (1949: 412). reddish, emerging from dorsal surface of stem in rows at
Discussion: This distinctive moss is recognized by its leaf bases; paraphyllia none; pseudoparaphyllia foliose;
differentiated branch and stem leaves with prorulose laminal axillary hairs with 1 short, brown basal cell and (1–)2–
cells. The leaves are plicate and short-decurrent, with strongly 3 elongate, hyaline apical cell(s). Asexual reproductive
excavate but otherwise weakly differentiated alar cells. The bodies only known from protonematal buds. Stem and
genus is unique among the Latin American Hypnaceae by branch leaves differentiated or not; stem leaves straight to
the presence of paraphyllia. The densely hairy calyptra is falcate, narrowly oblong-lanceolate to ovate, acuminate,
also distinctive. Although Nishimura et al. (1984) defined not or somewhat plicate, not decurrent; margins plane
the Hypnaceae as lacking paraphyllia, he (Nishimura, 1989) or recurved, subentire to serrulate above; costa short
appears to make an exception for this genus. and double or none; cells linear or linear-rhomboidal,
The description of the spore shape and ornamenta- smooth, firm-walled; alar cells well-differentiated, mostly
tion was taken from Nishimura (1989), who saw only 2–3 numerous, subquadrate to short-rectangular. Autoicous.
spores in a scanning electron microscope study of the peri- Perichaetial leaves erect, oblong-lanceolate, acuminate,
stome. However, the few spores that he observed were not margins serrulate at shoulders. Setae smooth, twisted;
enough to determine their size (N. Nishimura, Okayama capsules erect, ovoid to oblong-cylindrical, not constricted
University of Science, Japan, personal communication). below the mouth when dry; annulus differentiated or lack-
Selected specimens examined: MEXICO. Oaxaca: ing; operculum conic to obliquely rostrate; exostome erect
59 km NE de Llano de las Flores, Delgadillo 854 (MEXU, when moist, incurved when dry, on front surface smooth
NY). Puebla: El Cerro de Cuhuatepetl, Tehuacan, Santos below, sometimes papillose above, scarcely trabeculate at
3329 (FH, MICH, NY). Veracruz: El Esquilón, 1300 m, back; endostome free or fused to exostome teeth, mostly
Buck 35534 (NY). GUATEMALA. Alta Verapaz: Near papillose, with a high to low basal membrane, segments
Chirriacté, Standley 91604 (FH). Baja Verapaz: Civija, keeled, perforate or not, cilia rudimentary. Spores spheri-
Sharp 5193 (FH, MICH, NICH). NICARAGUA. Without cal, papillose. Calyptrae cucullate, smooth, naked.
locality, Garnier 5053 (FH, MICH). BRAZIL. Paraná: Ban- Distribution: Pylaisia is a genus of 15 species,
hado, Dusén 12177 (BM, FH, MICH, NY). Rio de Janeiro: primarily distributed in eastern Asia, but with a handful
Type of Papillaria rutenbergiacea. Rio Grande do Sul: Bom of species being circumboreal. Two of these extend into
Jesús, Sehnem 211 (FH, NY). Santa Catarina: Nova Vene- Mexico. Pylaisia falcata is the only species extending into
zia, E. Ule 1186 (H). São Paulo: Mpio. Eldorado Paulista, the Southern Hemisphere. The species are almost all ex-
Vital & Buck 12543 (NY, SP). clusively tree trunk epiphytes, although they may persist
on fallen trees.
13. Pylaisia Schimp. in Bruch, Schimp. Discussion: Pylaisia has long been known in North
& W. Gümbel America under the name Pylaisiella because of parahom-
onymy of the name. However, Pylaisia was recently con-
Pylaisia Schimp. in Bruch, Schimp. & W. Gümbel, Bryol. served. The genus is characterized by epiphytic pleurocarps
Eur. 5: 87. 1851, nom. cons.; Isothecium subg. Pylaisia with erect capsules. The branches are often curved when
(Schimp.) Boulay, Musc. France 146. 1884; Pylaisiella dry, and the leaves are somewhat falcate with often well
Kindb. ex Grout, Bull. Torrey Bot. Club 23: 228. 1896. developed alar regions. Pylaisia is uniquely characterized
Type: Pylaisia polyantha (Hedw.) Schimp. among its near relatives, Platygyrium and Platygyriella, by
 number 93 • 69

the autoicous sexual condition, foliose pseudoparaphyllia, Arikawa (2004) recently revised Pylaisia for the
and exostome teeth smooth on the front surface. world.

KEY TO THE SPECIES OF PYLAISIA

1. Alar regions extending up the margins by more than 15 cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. P. selwynii
1. Alar regions extending up the margins by less than 15 cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Endostome segments perforate; spores 25–40 µm in diameter; alar regions extending up the margins by only 2–6 cells . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. P. falcata
2. Endostome segments not perforate; spores 11–14 µm in diameter; alar regions extending up the margins by 8–10 cells. . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. P. polyantha

1. Pylaisia falcata Plants large, in mats, glossy, yellow-green. Stems to

5 cm long, creeping, regularly to irregularly pinnately
Pylaisia falcata Schimp. in Bruch, Schimp. & W. Gümbel, branched, the branches 5–10 mm long, usually creeping,
Bryol. Eur. 5: 89. 1851; Stereodon falcatus (Schimp.) rarely ascending and curved. Leaves falcate-secund, ovate
M. Fleisch. ex Broth. in Engl., Nat. Pflanzenfam. ed. 2, to narrowly oblong-ovate, ± gradually long-acuminate,
11: 452. 1925, hom. illeg., non Dozy & Molk.; Pylaisi- stem leaves 1.6–2.2 mm long, 0.6–0.9 mm wide, branch
ella falcata (Schimp.) Ando, Phyta 1: 14. 1978. Type: leaves 1.6–1.9 mm long, 0.4–0.6 mm wide, concave, not
Mexico. Orizaba, Liebmann (holotype: BM!; isotypes: plicate; margins plane, subentire to serrulate above; costa
H, PC). short and double or none; median cells 50–70 × 3–4 µm,
linear; alar cells subquadrate, extending up the margins by
Pylaisia subfalcata Schimp. in Bruch, Schimp. & W. Gümbel, Bryol. Eur. 2–6 cells, in 6–10 rows across the insertion. Perichaetial
5: 89. 1851; Stereodon subfalcatus (Schimp.) M. Fleisch. ex Broth. in Engl., leaves to 2.3 mm long, oblong-ovate, convolute, gradu-
Nat. Pflanzenfam. ed. 2, 11: 452. 1925; Stereodon falcatus var. subfalcatus ally long-acuminate; margins serrulate at shoulders. Setae
(Schimp.) Thér., Rev. Bryol. Lichénol. 5: 110. 1933. Type: Mexico. Lieb- 15–25 mm long, red-brown, sinistrorsely twisted in lower
mann 9/42 (lectotype: BM, designated by Arikawa, 2004). 2/3 and dextrorsely twisted in upper 1/3 when dry; cap-
sules 1.5–2.5 mm long, cylindrical, yellowish to reddish
Stereodon hamatus Mitt., J. Linn. Soc., Bot. 12: 533. 1869; Hypnum hama- brown; operculum obliquely short-rostrate; exostome teeth
tum (Mitt.) A. Jaeger, Ber. Thätigk. St. Gallischen Naturwiss. Ges. 1877–78: ~250 µm long, smooth below, finely papillose above; end-
312. 1880, hom. illeg., non Dozy & Molk.; Pylaisia hamata (Mitt.) Cardot, ostome free or somewhat adherent to the exostome teeth
Rev. Bryol. Lichénol. 38: 103. 1911; Hypnum mittenohamatum B. C. Tan, below, with a low, smooth, basal membrane, endostome
Mem. New York Bot. Gard. 68: 5. 1992. Type: Ecuador. Andes Quitenses, longer than the exostome teeth, densely papillose-spiculose,
in montibus Chimborazo, Jameson (lectotype: NY!, designated by Arikawa, keeled, perforate, sometimes bifid, cilia rudimentary. Spores
2004); Colombia. Andes Bogotenses, ad viam inter Tipaquira et Pacho, Weir 25–40 µm in diameter, papillose.
265 (syntype: NY!); Ecuador. Pichincha, Spruce 1046 (syntype: NY!). Distribution and ecology: Known from Mexico to
Bolivia; also in eastern Asia (China and Nepal); growing on
Pylaisia macrotis Cardot, Rev. Bryol. 37: 10. 1910. Type: Mexico. Las tree trunks and logs at 2000–4000 m.
Chipans, district de San Cristobal, 23 Dec 1907, Münch s.n. (holotype: PC). Illustrations in publications: Pl. 753 (as Pylaisiella
falcata (Bruch, Schimp. & W. Gümbel) Ando) in Ando
Pylaisia panduraefolia Herzog, Biblioth. Bot. 87: 126. 1916; Stereodon pan- (1994: 1014); Fig. 25 in Arikawa (2004: 110); Fig. 175H–J
durifolius (Herzog) Broth. in Engl., Nat. Pflanzenfam. ed. 2, 11: 452. 1925. (as Stereodon falcatus (Schimp.) M. Fleisch.) in Bartram
Type: Bolivia. In Hochland von Totora, Herzog 5123 (holotype: JE; isotypes: (1949: 397).
L, M, S). Discussion: Pylaisia falcata is the most common
Latin American species of the genus and also the one with
Pylaisia falcata var. intermedia Thér., Smithsonian Misc. Collect. 78(2): 28. the largest plants. It is characterized by its large size, falcate-
1926; Stereodon falcatus var. intermedius (Thér.) Thér., Rev. Bryol. Liché- secund leaves, narrow laminal cells, relatively few quadrate
nol. 5: 110. 1933. Type: Mexico. Vicinity of Puebla, Esperanza, Arsène 4667 alar cells, papillose-spiculose endostomial segments, and
(lectotype: PC, designated by Arikawa, 2004). large spores.
70 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Selected specimens examined: MEXICO. Chiapas: 2. Pylaisia polyantha

Cima del Cerro Mozotal, 30 km NW of Motzintla, Delga-
dillo M. 4771 (MEXU, NY). Distrito Federal: Cima, Prin- Pylaisia polyantha (Hedw.) Schimp. in Bruch, Schimp.
gle 10521 (MO, NY). Durango: 30 km W of El Salto, Sharp & W. Gümbel, Bryol. Eur. 5: 89. 1851; Leskea poly-
1846 (NY). Hidalgo: Cerro Xihuingo, al N de Tepeapulco, antha Hedw., Sp. Musc. Frond. 229. 1801; Hypnum
2750 m, Cárdenas S. 1730 (MEXU, NY); above Real de polyanthum (Hedw.) Dicks., Fasc. Pl. Crypt. Brit. 4: 17.
Monte, 2852 m, Sharp 506 (US). Jalisco: El Cuartón, above 1801; Pterogonium polyanthos (Hedw.) Muhl., Cat.
Manantlán, Crum 1199 (MICH, NY). México: Mpio. Pl. Amer. Septentrionalils 98. 1813; Isothecium poly-
San Martín de las Pirámidas, Cerro Gordo, Buck 21486 anthum (Hedw.) Spruce, Musci Pyren. 79. 1847; Ste-
(NY). Michoacán: Hills of Patzcuaro, Pringle 749b (NY). reodon polyanthos (Hedw.) Mitt., J. Proc. Linn. Soc.,
Oaxaca: Summit of Cerro Yucuyacua, ~12 km S of Tlaxi- Bot., Suppl. 1: 94. 1859; Pylaisiella polyantha (Hedw.)
aco, Hallberg 1085 (NY). Puebla: Esperanza, Purpus 4290 Grout, Bull. Torrey Bot. Club 23: 229. 1896. Type: Ad
(MO, NY). Veracruz: Slope of Mt. Orizaba, 3050 m, Smith truncos arborum annosarum, maxime salicum remotius
15 p.p. (NY). GUATEMALA. Huehuetanango: Sierra de ab earum basi in Germania, Anglia (lectotype: G, desig-
los Cuchumatanes, along road beyond La Pradera, Km nated by Geissler in Margadant and Geissler, 1995).
32, Standley 81795 (NY). Sololá: Sierra Madre Moun-
tains, Cerro María Tecum, Williams et al. 41755B (MO). See Arikawa (2004) for additional synonymy based on
Totonicapán: Along road between San Francisco El Alto non–Latin American types.
and Momostenango, Standley 84028 (NY). COSTA RICA.
Cartago: S slope of Volcán Irazu, on highway about 5 km Plants rather small, in low mats, glossy, yellow-green to
NE of Finca Robert, Holm & Iltis 1176 (G). San José: bright green. Stems to 2 cm long, prostrate, regularly pin-
Cerro de la Muerte, ~4.1 km NW of Villa Mills, Koch 5083 nately branched, the branches to 1 cm long, usually creep-
(NY). PANAMA. Chiriquí: Along road from gatehouse to ing, rarely ascending and curved. Leaves mostly straight,
summit of Volcán Baru, Allen 9149 (MO, NY). COLOM- ovate-lanceolate, gradually long-acuminate, stem leaves
BIA. Antioquia: Mpio. de Sonsón, 10.7 km E de Sonsón 1.3–2.0 mm long, 0.4–0.5 mm wide, branch leaves 1.1–
hacia Nariño/Argelia, Churchill & Sastre-De Jesús 13013 1.4 mm long, 0.3–0.5 mm wide, concave, slightly plicate;
(NY). Boyacá: Mpio. de Duitama, al lado de páramo Rusia, margins plane or slightly involute above, serrulate above;
Churchill et al. 19061 (NY). Caldas: Mpio. de Vállamaria, costa short and double; median cells 50–80 × 4–5 µm, lin-
cerca de Km 214, carretera a Nevado del Ruíz, Churchill & ear; alar cells subquadrate to short-rectangular, extending
Betancur 17934-B (NY). Cauca: Mpio. de Coconuco, ~18 up the margins by 8–10 cells, in 5–8 rows across the inser-
km E de Puracé, Churchill & Rengifo M. 17287 (NY). Cun- tion. Perichaetial leaves to 1.4 mm long, oblong-lanceolate,
dinamarca: Along trail NE from Bogotá (via La Calera) to abruptly short-acuminate; margins serrulate at shoulders.
Mundo Nuevo, 2593–3050 m, Steere 7725 (NY). Nariño: Setae 10–18 mm long, yellow- to red-brown, sinistrorsely
Mpio. de Pasto, 3 km E de Dolores, Ramírez P. 3612 twisted in lower 3/4 to 4/5 and dextrorsely twisted in upper
(NY). VENEZUELA. Mérida: Páramo de Los Granates, al 1/4 to 1/5 when dry; capsules 1.6–2.2 mm long, long-
rededor de El Chimborazo, un sector del páramo do Los cylindrical, reddish brown; operculum obliquely conic;
Granates, Griffin III & López F. PV-1092 (NY); Distr. exostome teeth ~200 µm long, smooth below, finely papil-
Rivas Davila, páramo La Negra, above town of Bailadores, lose above; endostome free, with a well-developed, smooth,
Griffin III et al. 1992 (NY). ECUADOR. Azuay: 0–2 km basal membrane, endostome longer than the exostome
W of Laguna Llaviucu, ~10 km WNW of Sayausí, 3200 m, teeth, papillose, keeled, not perforate, cilia single, rudimen-
Løjtnant et al. 14761 (NY). Carchi: Tulcán–El Angel road, tary. Spores 11–14 µm in diameter, finely papillose.
34 km S of Las Juntas, 3550 m, Balslev 901 (NY). Chim- Distribution and ecology: Pan North Temperate,
borazo: 11 km E of Alao, 3526 m, Dorr & Barnett 6224 extending into eastern Mexico (Nuevo Léon, San Luis Po-
(NY). Imbabura: Cotocachi–Apuela road, 3111 m, Steere tosí, Veracruz); occurring throughout North America, also
26636 (NY). Napo: 6 km E of Papallacta, 2880 m, Buck in Europe, northern and central Asia, China, Japan, and
9968 (NY). Pichincha: Road Chillogallo–Chiriboga, 3000 Africa; growing on tree trunks at 2500–3600 m.
m, Holm-Nielsen 18658 (NY). PERU. Cajamaraca: Prov. Illustrations in publications: Pl. 754 (as Pylaisiella
Celendin, östlich des Passes zwischen Celendin und Balas, polyantha (Hedw.) Grout) in Ando (1994: 1015); Fig. 16 in
Hegewald & Hegewald 6659 (NY). BOLIVIA. See type of Arikawa (2004: 94); Fig. 560A–G in Crum and Anderson
Pylaisia panduraefolia Herzog. (1981: 1137); Pl. 343 in Ireland (1982: 600).
 number 93 • 71

Discussion: This North Temperate species just in Crum and Anderson (1981: 1134); Pl. 341 in Ireland
barely makes it into the Neotropics. It is characterized by (1982: 598).
a moderate number of alar cells (intermediate number be- Discussion: This North Temperate species creeps
tween P. selwynii and P. falcata), a free endostome with into the Neotropics in western Mexico. It is characterized by
nonperforate segments, and small spores. Its branches are ascending and curved branches, relatively short-acuminate
not ascending and curved as in P. selwynii, and the leaves leaves, relatively short laminal cells, numerous alar cells, an
are not falcate as in P. falcata. endostome adherent to the exostome teeth, and spores in-
Specimen examined: Mexico. Nuevo León: Main termediate in size between P. falcata and P. polyantha.
road up Cerro Potosi, just below summit, ~17 km NW of Specimens examined: Mexico. Sonora: E of El
Galeana, 3600 m, 26 Jul 1985, Whittemore 2721 (MO). Tigre, Cañon de Santa Rosa, White 559 (UBC); Río de
Bavispe, La Nacha, White 4013 (S). Sinaloa: ~2.8 km E of
3. Pylaisia selwynii Las Palmitas, along Hwy. 40, Norris 20579 (UBC).

Pylaisia selwynii Kindb., Ottawa Naturalist 2: 156. 1889; 14. Rhacopilopsis Renauld & Cardot
Pylaisiella selwynii (Kindb.) H. A. Crum, Steere & L. E.
Anderson, Bryologist 67: 164. 1964. Type: Canada. Rhacopilopsis Renauld & Cardot, Rev. Bryol. 27: 47.
Ottawa, Richmond Road, 15 May 1885, Macoun s.n. 1900. Type: Rhacopilopsis dupuisii (Renauld & Car-
(holotype: S!; isotype: CANM!). dot) Renauld & Cardot.

Plants small, in mats, glossy, yellow-green to dark green. Hypnum sect. Dimorphella Müll. Hal., Flora 69: 523. 1886; Dimorphella
Stems to 2 cm long, creeping, regularly pinnately branched, (Müll. Hal.) Renauld & Cardot, Bull. Soc. Roy. Bot. Belgique 41(1): 101.
branches to 3 mm long, ascending and curved when dry. 1904 [1905]. Type: Hypnum pechuelii Müll. Hal.
Leaves straight to secund, ovate, abruptly short-acuminate,
stem leaves 0.9–1.2 mm long, 0.5–0.6 mm wide, branch Acanthocladiella M. Fleisch., Musci Buitenzorg 4: 1381. 1923. Type: Acan-
leaves 0.7–0.9 mm long, 0.25–0.3 mm wide, not concave, not thocladiella flexilis (Renauld & Cardot) M. Fleisch.
plicate; margins plane or slightly incurved above, serrulate
above; costa short and double; median cells 25–45 × 4.3– Plants in thin mats, often lustrous, soft, green to pale
7 µm, linear; alar cells oblate to short-rectangular, extending green. Stems creeping, irregularly to regularly pinnate, the
up the margins by 20–25 cells, in 6–10 rows across the inser- branches complanate-foliate; stems in cross section without
tion. Perichaetial leaves to 1.6 mm long, oblong-lanceolate, a hyalodermis, with small thick-walled cells surrounding
gradually long-acuminate; margins subserrulate above. Setae larger thinner walled cells, central strand none; para-
10–15 mm long, yellow- to red-brown, sinistrorsely twisted phyllia none; pseudoparaphyllia filamentous, but often
in lower ½ and dextrorsely twisted in upper ½ when dry; 2–5-seriate at base, or rarely foliose; axillary hairs consist-
capsules 1.0–1.6 mm long, cylindrical, reddish brown; oper- ing of 1–2 short, brown basal cell(s) and 2–3 elongate,
culum obliquely conic; exostome teeth ~150–200 µm long, hyaline distal cells. Asexual reproductive bodies lacking.
smooth below, finely papillose above; endostome adherent Stem and branch leaves somewhat differentiated, ventral,
to exostome teeth below, free above, with a well developed, dorsal and lateral stem leaves sometimes differentiated,
smooth, basal membrane, segments extending beyond the when differentiated the ventral ones symmetric, oblong-
exostome teeth, papillose, keeled, bifid with ½ of segment ovate, with a wide insertion, gradually long-acuminate,
plus cilium united and adherent below to exostome teeth. the dorsal ones somewhat asymmetric, ovate, gradually
Spores 16–25 µm in diameter, finely papillose. acuminate, concave, the lateral ones strongly asymmetric,
Distribution and ecology: Reaching its southern ovate-oblong, ± abruptly acuminate, concave; stem leaf
limit in western and southern Mexico (Chiapas, Chihua- margins subentire to distantly subserrulate throughout,
hua, and Oaxaca in addition to those cited below); com- plane to recurved; costa short and double; cells long-hex-
monly occurring in eastern North America; and also known agonal to linear, smooth but with thickened cell end walls
from eastern Asia, extending into central European Russia; giving the impression of prorulae, firm-walled, not porose;
growing on tree trunks. alar cells differentiated in extreme basal angles, not reach-
Illustrations in publications: Pl. 752 (as Pylaisiella ing the costa, 2–4 cells enlarged and inflated, ± oblong,
selwynii (Kindb.) Crum, Steere & L. E. Anderson) in Ando sometimes colored, with a small group of subquadrate to
(1994: 1013); Fig. 31 in Arikawa (2004: 120); Fig. 558 short-rectangular cells directly above them, in lateral leaves
72 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

the enlarged cells sometimes only on one side of the costa. graph (Watling and O’Shea, 2000) has reduced the num-
Branch leaves with dorsal and lateral leaves sometimes ber of species to two.
differentiated, when differentiated, the dorsal ones erect-
spreading, symmetric, lanceolate, gradually acuminate, the Rhacopilopsis trinitensis
lateral ones wide-spreading, asymmetric, oblong-ovate,
gradually or ± abruptly acuminate, somewhat concave; all Rhacopilopsis trinitensis (Müll. Hal.) E. Britton & Dixon
branch leaves with margins subentire to serrate through- in Dixon, J. Bot. 60: 88. 1922. Hypnum trinitense
out, the lateral leaves more strongly toothed, plane to nar- Müll. Hal., Syn. Musc. Frond. 2: 284. 1851; Ectropo-
rowly recurved, especially above; costa short and double; thecium trinitense (Müll. Hal.) Mitt., J. Linn. Soc., Bot.
cells linear to linear-flexuose, smooth, firm-walled, not po- 12: 514. 1869. Type: Trinidad. Ad Maraccas, Arima
rose, sometimes becoming shorter in the acumen; alar cells et montem Touche, Aug et Dec 1844 et 1847, Crüger
differentiated in extreme basal angles, only rarely reaching (syntype: BM!).
the costa, colored, 2–4 cells enlarged with a small group of
rhomboidal to subquadrate cells directly above them. Di- Dimorphella brasiliensis Broth. & Herzog in Herzog, Arch. Bot. Est. São
oicous. Perichaetial leaves erect, lanceolate-ovate, abruptly Paulo 1: 86. 1925. Type: Brazil. Minas Gerais: Serra da Caraça, 22 Jan 1921,
acuminate, margins sometimes strongly toothed above. Hoehne 8283 (173) (isotype: BM!).
Setae smooth, red or orange; capsules inclined to horizon-
tal, ovoid to short-cylindrical, constricted below the mouth See Watling and O’Shea (2000) for additional synonymy based on African
when dry; operculum conic, mammillate to short-rostrate; types.
exostome teeth on front surface cross-striolate below, pap-
illose above; endostome finely papillose, about as long as Plants slender to medium-sized, in ± thin mats, lustrous,
the teeth, with a high basal membrane, segments keeled, soft, mostly pale green. Stems to 10 cm long, long-creeping,
perforate, cilia in pairs, about as long as the teeth. Spores often regularly pinnate, the branches complanate-foliate,
spherical, finely papillose. Calyptrae unknown. less than 1 cm long; stems in cross section with 2–4 rows of
Distribution: Of the two species of Rhacopilopsis small thick-walled cells surrounding larger thinner walled
(Watling and O’Shea, 2000), one occurs in Latin America cells, central strand none; pseudoparaphyllia filamentous;
and Africa, while the other is strictly African; usually at low axillary hairs consisting of 1 relatively long or 2 short, brown
elevations. basal cell(s) and 2–3 elongate, hyaline distal cells. Stem and
Discussion: Although not obvious, once the differ- branch leaves somewhat differentiated, ventral, dorsal and
entiated dorsal and lateral branch leaves are noticed, Rha- lateral stem leaves strongly differentiated, the ventral ones
copilopsis is unlikely to be confused with few genera. The 0.65–0.95 mm long, symmetric, oblong-ovate, gradually
often whitish green plants with complanate-foliate branches long-acuminate, the dorsal ones 1.0–1.3 mm long, some-
are also helpful characters for recognition. However, this what asymmetric, ovate, gradually acuminate, concave, the
generic definition only holds up for the single Latin Ameri- lateral ones 0.85–1.2 mm long, strongly asymmetric, often
can one, and not the African endemic R. variegatum (Welw. cultriform, ovate-oblong, ± abruptly acuminate, concave; all
& Duby) Watling & O’Shea. stem leaves with margins distantly subserrulate throughout,
We have seen no sporophytes, and the descriptions plane; costa short and double; median cells long-hexagonal
provided here are adapted from that provided by Watling to linear, 8–20:1, smooth but with thickened cell end walls,
and O’Shea (2000). Bartram (1949: 414) described the giving the impression of prorulae, firm-walled, becoming
setae as 1–2 cm long, the capsules small (~1 mm long) thicker-walled and somewhat porose toward the insertion;
and nodding, the endostomial segments arising from a alar cells mostly yellow, 2–4 cells enlarged in extreme basal
high basal membrane, and the spores 7–8 µm in diam- angles, oblong, to 55 µm long, sometimes absent on one
eter. Crum (1994b) reported the spores to be ~17 µm in side of lateral leaves, with 5–12 rhomboidal to subquad-
Mexican plants. rate cells directly above the enlarged ones. Branch leaves
According to Dixon (1922), the genus is monospecific with dorsal and lateral leaves differentiated, the dorsal ones
and the American and African material is inseparable. 0.8–1.1 mm long, half as many as the lateral ones, sym-
However, Buck (1993) merged Rhacopilopsis with the Af- metric, erect-spreading, lanceolate, gradually acuminate,
rican Acanthocladiella, resulting in a more speciose genus. the lateral ones 1.1–1.3 mm long, somewhat asymmetric,
The nonfalcate, mostly toothed leaves with a few inflated wide-spreading, sometimes subfalcate, oblong-ovate, grad-
alar cells also hold the genus together. A recent mono- ually or ± abruptly acuminate, somewhat concave; branch
 number 93 • 73

leaf margins subentire to serrulate throughout, the lateral Darién: Pirre Massif just W of Cana, Allen 8889 (MO).
leaves more strongly toothed, plane or narrowly recurved Panamá: 6.5 km by road N of Lago Cerro Azul, Nee 9325
above; costa short and double, rarely lacking; cells linear to (MO, NY). Veraguas: 0.3 km beyond fork in road at Es-
linear-flexuose, 10–20:1, smooth, firm-walled, sometimes cuela Agricola Alto Piedra, Croat & Folsom 33939 (MO).
becoming shorter in the acumen, becoming thicker walled Isla Bahia Honda: Without locality, Taylor 1110 (NY).
and somewhat porose toward the insertion; alar cells yel- DOMINICA. Syndicate Estate, ~8 km S of Portsmouth,
low, 3–5 cells enlarged in extreme basal angles, oblong, to 605 m, Townsend 91/296 (NY). GUADELOUPE. Vicin-
55 µm long, rarely absent on one side of lateral leaves, with ity of Grand Etang, 400 m, Crosby & Crosby 4800 (MO,
4–12 rhomboidal to subquadrate cells directly above the NY). MARTINIQUE. Without locality, 1899, Duss 86
enlarged ones. Asexual propagula not seen. Dioicous. Pe- (NY). TRINIDAD AND TOBAGO. Trinidad: Morne Bleu,
richaetial leaves long-attenuate; margins strongly toothed. Britton 2301 (NY). COLOMBIA. Arauca: Mpio. de Tame,
Setae 17–30 mm long; capsules 1.8–2.5 mm long. S de Tame, entre San Lope, 485 m, Churchill et al. 18941
Distribution and ecology: Guatemala to Panama, (NY). Boyacá: Mpio. de Cubara, ~5 km E de Cubara, 300
northern South America, Brazil, Trinidad; elsewhere in m, Churchill et al. 18933 (NY); Casanare, Tauramena,
central Africa and Madagascar; growing on tree trunks, es- 600 m, Uribe 4292(US). Caquetá: Transecto Neiva–San
pecially at the bases, occasionally on logs, mostly in moist Vicentge del Caguán, 1200–1300 m, Churchill & Betan-
forests, at 400–900 m. cur 16897 (NY). Meta: Cordillera La Macarena, Schultes
Illustrations in publications: Fig. 779 in Crum 11157 (NY, US); 20 km SE of Villavicencio, 480 m, Alston
(1994b: 1051); Fig. 175 in Florschütz-de Waard and Veling 7581 (US). Putumayo: ~40 km NW of Puerto Asís, ~305 m,
(1996: 457); Pl. 121 in Buck (1998: 309); Fig. 143 in Buck King & Guevara C-1136 (US). Santander: E of Hacienda El
(2003: 161); Fig. 183A–D in Bartram (1949: 415). Playón, NW of Bucaramanga, 1220–1373 m, Steere 7638
Discussion: The heterophyllous leaves with yel- (NY). VENEZUELA. Amazonas: Along Río Mawarinuma,
lowed insertions are distinctive. The species is decidedly just outside Cañon Grande of Cerro de la Neblina, 140 m,
rare in the West Indies, although it is not uncommon in Buck 12933 (NY). Apure: Reserva Forestal San Camilo,
Trinidad, Central America, the Guianas, and northern An- 250–280 m, Steyermark et al. 101723 (NY,US), 101730,
dean South America. 101850 (US); Quebrada La Azulita, 280 m, Steyermark
Although we have not critically evaluated Cardot’s et al. 101607 (US); 4.5–5 km N of San Camilo, 280 m,
(1909: 50) two varieties—Dimorphella pechuelii (“i”) Steyermark et al. 101789 (US). Barinas: Distr. Pedraza,
(Müll. Hal.) Renauld & Cardot var. acuminata Cardot trail from Mesa de Canagua to Cerro de Filón, 600–880 m,
and D. pechuelii (“i”) (Müll. Hal.) Renauld & Cardot Dorr et al. 7842 (NY). Bolivar: Gran Sabana, San Igna-
var. gracilis Cardot—that were described from Africa cio de Yuruaní, 850 m, Liesner 24230 (MO, NY). Sucre:
but which have also been reported from the New World Peninsula de Paria, entre Manacal y Los Pocitos de Santa
(Dixon, 1922), we are unconvinced for this treatment that Isabel, 700–900 m, Dumont et al. VE-7496 (NY); Cerro de
they deserve taxonomic recognition. Río Arriba, vecindad de “Los Positos,” 700 m, Steyermark
Selected specimens examined: GUATEMALA. & Rabe 96298 (US). Tachira: Oeste de Ayari, 200 m, Steyer-
Izabal: Valley of tributary of Río San Francisco del Mar, mark & Rabe 96644 (US). Yaracuy: Quebrada Honda,
3.2 km NE of Hopi, 19.3 km E of Entre Ríos, 10-20 m, 17.3 km del pueblo de Aroa, 1100 m, Steyermark 105405
Steyermark 39779 (F). HONDURAS. Yoro: Cordillera (NY, US). Zulia: Sierra de Perija, valley of the Río Socuy,
Nombre de Dios, N slope, ridge between Río Guán Guán 2400–2800 m, Griffin III 161 (NY). Hato La Vergareña,
and Quebrada Aguacatal, Allen 13599 (MO, NY). COSTA ~75 km S of Cd. Piar, Pursell 8050 (US). GUYANA. Upper
RICA. Alajuela: Río Jesús (Santiago) de San Ramón, Brenes Mazaruni Dist.: Right bank of Mazaruni River, 05°52'N,
20420 (NY). Limón: Up to 4 km inland from Green Turtle 60°34'W, 473–641 m, Robinson 85-0081 (US). Kamak-
Station and village of Tortuguero, Steere CR-5 (NY). San usa, along the upper Mazaruni River, Leng 4 (NY); NW
José: Vicinity of El General, Skutch 2694 (MO, NY). Pun- portion of Kanuku Mountains, Mt. Iramaikpang, 850 m,
tarenas: Cocos Island, without locality, Holdridge 5194 Smith 3618 (NY, US); Essequibo River, Moraballi Creek,
(NY). PANAMA. Canal Area: Pipeline road leading NW near Bartica, near sea level, Richards 241 (NY). Potaro-
from Gamboa, Crosby 10800 (MO). Chiriquí: Cerro Colo- Siparuni Region: Kaieteur Natl. Park, 400 m, Pipoly 9904
rado, Chami Camp vicinity Cerro Colorado Mine, Allen (NY). SURINAM. Sipaliwini: Tafelberg, Maguire 24339
5000 (MO, NY). Colón: Along Santa Rita Ridge, ~12.4 km M3 (NY), Allen 20549 (MO, NY). FRENCH GUIANA.
NE from Boyd–Roosevelt Hwy., Crosby 4468 (MO). Approuague-Kaw: Pic Matécho, 590 m, Buck 37969 (NY),
74 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

l’Inini Region: Mont Atachi Bacca, 03°33N, 53°55'W, 670 erect-spreading, lanceolate to ovate, acuminate, concave,
m, Cremers et al. 10444 (US). Maripasoula: Commune de nondecurrent; margins plane or erect, sometimes recurved
Saül, vicinity of Eaux Claires, 200 m, Buck 18354 (NY). below, subentire to serrulate above, subentire below; costa
Roura: Kaw Mountains, Trésor Réserve Naturelle, 300 short and double; cells linear, smooth, thin- to firm-walled,
m, Buck 37769 (NY). ECUADOR. Napo: 6 km along Río not porose; alar cells abruptly differentiated, few, quad-
Pano, 0°58'S, 77°52'W, 600 m, Holm-Nielsen & Jeppesen rate, extending up the margin by a few cells. Autoicous.
702a (US). Pastaza: Lorocachi, 2 km del Río Curaray, 200 Perichaetial leaves erect, triangular-lanceolate, acuminate;
m, Jaramillo et al. 30715 (NY). Pichincha: ~18 km NW of margins plane. Setae smooth, straight, sometimes twisted,
San Miguel de los Bancos, forest reserve of ENDESA, 800 orange to reddish; capsules erect to suberect, cylindrical, not
m, Buck 10365 (NY). PERU. Madre de Dios: Los Amigos at all contracted below mouth; annulus poorly developed
Biological Station, 250–320 m, Majestyk 4169 (NY). Near or not differentiated; operculum high-conic, bluntly ros-
Pucallpa, Aug 1962, Worthley s.n. (US). BOLIVIA. La trate; exostome teeth erect when dry, incurved when moist,
Paz: Franz Tamayo, Parque Nacional Madidi, 700–900 m, bordered, on front surface at extreme base cross-striolate,
Fuentes et al. 4441 (MO, NY). BRAZIL. Amapá: Mpio. de finely papillose-spiculose above, scarcely projecting at back;
Oiapoque, BR156 between Calçoene and Oiapoque, Mori endostome densely and finely papillose throughout from a
& Cardoso 17108 (NY). Mato Grosso: Chapada dos Gui- medium high basal membrane, segments keeled, not or nar-
marães, Wasserfall Veu da Noiva, 500 m, Schäfer-Verwimp rowly perforate, sometimes as tall but not taller than teeth,
& Verwimp 8628 (NY). Pará: Serra do Cachimbo, Rio Ja- cilia none to single. Spores spherical, finely papillose or
manxím and vicinity, 300 m, Reese 16677 (NY, US); Serra roughened. Calyptrae cucullate, smooth, naked.
Maze and vicinity, ~05°55'S, 55°40'W, ~100–200 m, Reese Distribution: The two species of Syringothecium
16826 (US). Roraima: Pacaraima, 226 km N of Boa Vista, occur in terrestrial or epiphytic habitats at moderate eleva-
1000 m, Buck et al. 1944 (NY). tions in South America.
Discussion: Syringothecium is distinguished by the
15. Syringothecium Mitt. endostome being as well developed or more so than the exo-
stome. The endostome segments are very long, straight, erect
Syringothecium Mitt., J. Linn. Soc., Bot. 12: 497. 1869. and extend beyond the exostome teeth. In all the Latin Ameri-
Type: Syringothecium sprucei Mitt. can Isopterygium spp. the endostome segments are scarcely
evident. The rhizoids are smooth to sometimes slightly rough-
Plants small to medium-sized, in lax to dense mats, ened compared to the consistently smooth rhizoids of Isopter-
yellow-green. Stems creeping, irregularly but freely ygium. Although traditionally placed in the Sematophyllaceae,
branched; stems in cross section lacking a hyalodermis, with the genus seems to be a segregate from Isopterygium. In fact,
a couple of rows of small, thick-walled cells and large, thin- Buck (1989) synonymized the genus with Isopterygium. Al-
walled cells in center, central strand absent; rhizoids smooth though we suspect that the genus truly is derived from within
to slightly roughened, in clusters just below the leaf inser- Isopterygium (and thus making that genus paraphyletic), we
tions; paraphyllia absent; pseudoparaphyllia filamentous; are tentatively recognizing it here. Both species seem to be
axillary hairs consisting of 1 short, brown basal cell and rare, and recent collections are unavailable for DNA sequenc-
2 elongate, hyaline distal cells. Asexual reproductive bod- ing. It seems to differ only from Isopterygium by the peristo-
ies lacking. Stem and branch leaves similar, soft, imbricate, mial features upon which it was described.

KEY TO THE SPECIES OF SYRINGOTHECIUM

1. Leaves lanceolate, gradually long-acuminate, with erect or narrowly recurved margins; spores 8–10 µm in diameter . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. S. brasiliense
1. Leaves lanceolate-ovate to ovate, short-acuminate, with plane margins; spores 14–17 µm in diameter . . . . . . . 2. S. sprucei

1. Syringothecium brasiliense rango, ~800 m, in culmis Chusquearum, 17 Feb 1904,

Dusén 3895 (lectotype: H!, designated by Buck, 1989;
Syringothecium brasiliense Broth., Denkschr. Kaiserl. isolectotype: H!); Dusén 3894 (syntype: H!).
Akad. Wiss., Math.-Naturwiss. Kl. 83: 338. 1926; Isop-
terygium brasiliense (Broth.) W. R. Buck, Bryologist 92: Plants in lax mats, yellowish green. Stems ~2 cm
531. 1989. Type: Brazil. Paraná: Serra do Mar, Ypi- long, creeping, irregularly but freely branched, somewhat
 number 93 • 75

complanate-foliate; stems in cross section with 1–2 rows spreading, little altered when dry, lanceolate-ovate to
of epidermal cells. Leaves 1.1–1.4 mm long, spreading to ovate, short-acuminate, concave; margins plane, subentire
wide-spreading, somewhat contorted when dry, lanceo- to serrulate above, subentire below; costa short and double,
late, long-acuminate, concave; margins erect, sometimes sometimes to ¼ the leaf length; cells 4–6 µm wide, linear,
narrowly recurved below, entire or subserrulate above, becoming shorter in the acumen and across the sometimes
subentire below; costa short and double, rarely to 1/5 the yellowed insertion; alar cells few, quadrate, extending up
leaf length; cells 5.5–6.5 µm wide, linear, mostly thin- the margin by 2–3 cells. Autoicous. Perichaetial leaves
walled, somewhat broader, to 9 µm in the acumen but not 1.4–1.9 mm long, erect, triangular-lanceolate, gradually
much shorter, shorter and broader in a single row across long-acuminate; margins plane to erect, entire or laxly ser-
the insertion; alar cells few, rectangular to subquadrate, rulate at apex; costa none; cells linear, firm-walled, be-
extending up the margin by 3–4 cells. Autoicous. Perichae- coming shorter and broader across the yellowed insertion.
tial leaves 1.4–2.0 mm long, erect, triangular-lanceolate, Setae 11–19 mm long, reddish; capsules ~1.5 mm long,
gradually long-acuminate; margins mostly erect, subentire erect to suberect, cylindrical, not at all contracted below
to serrulate above. Setae 10–13 mm long, orange, straight, mouth; annulus poorly developed, of 1–2 rows of small,
twisted; capsules 1–1.3 mm long, erect to suberect, cy- quadrate cells, sometimes adhering to operculum; opercu-
lindrical, not at all contracted below mouth; annulus ap- lum ~0.8 mm long, high-conic, bluntly rostrate; exostome
parently not differentiated; operculum ~0.75 mm long, teeth ~430 µm high, reddish but becoming yellowish
high-conic, bluntly rostrate; exostome teeth ~600 µm tall, soon after operculum falls, lanceolate, strongly bordered,
orange, lanceolate, strongly bordered, not shouldered, not shouldered, on front surface at extreme base cross-
on front surface at extreme base cross-striolate, directly striolate, finely papillose-spiculose above, trabeculate at
above that papillose-striolate, often in irregular patterns, back; endostome with basal membrane of medium height,
finely papillose-spiculose above, scarcely trabeculate segments not or narrowly perforate only toward base,
at back; endostome with a basal membrane of medium usually extending more than ~20 µm above the teeth, cilia
height, segments not perforate, extending to the height of none. Spores 14–17 µm in diameter, finely papillose.
the teeth but appearing taller in dry material, cilia usually Distribution and ecology: Rare and so far known
single and short, sometimes rudimentary. Spores 8–10 µm only from Colombia, Ecuador, and Bolivia; usually grow-
in diameter, finely roughened. ing on twigs and rotten wood at 1000–2100 m.
Distribution and ecology: Known only from the Illustrations in publications: Figs. 1–8 in Buck
type specimens in southeastern Brazil; growing on culms (1989: 530).
of bamboo. Specimens examined: COLOMBIA. Antioquia:
Illustrations in publications: Figs. 9–15 in Buck Mpio. Guatapé, Finca Montepinar, ~15 km NNE de Gua-
(1989: 530). tapé, 06°18'N, 75°09'W, 1810–1840 m, Churchill 16405
Specimen examined: BRAZIL. Paraná. Seen only (NY); Mpio. Frontino, corregimiento Nutlbara, coenca alta
from the type collection. del Río Cuevas, Finca “La Peña de la India,” 1880, Sánchez
et al. 698 (NY). Nariño: Mpio. Barbacoas, Corregimiento
2. Syringothecium sprucei Altaquer, Vereda El Barro, Reserva Natural Río Ñambí,
01°18'N, 78°08'W, Giraldo et al. 1973 (NY). ECUADOR.
Syringothecium sprucei Mitt., J. Linn. Soc., Bot. 12: 497. Napo: Summit of Sierra de Guayamacas, new road between
1869; Isopterygium sprucei (Mitt.) W. R. Buck, Bryolo- Tena and Baeza, 2100 m, on rotting log along road, 12 Jan
gist 92: 529. 1989. Type: Ecuador. Andes Quitenses, 1982, Steere E-131 (NY). BOLIVIA. Known only from the
Montaña de Canelos, 3000 ft., in forest, Spruce 1517 type locality of S. nemodontium.
(holotype: NY!; isotypes: NY–2!).
16. Taxiphyllum M. Fleisch.
Syringothecium nemodontium Herzog, Hedwigia 67: 267. 1927. Type: Bo-
livia. Hacienda Simaco am Weg nach Tipuani, ~1400 m, Oct 1920, Buchtien Taxiphyllum M. Fleisch., Musci Buitenzorg 4: 1434. 1922.
121 (holotype: JE; isotypes: H!, BR!, US!). Type. Hypnum deplanatum Schimp. ex Müll. Hal., Syn.
Musc. Frond. 2: 260. 1851.
Plants in dense mats, yellowish green. Stems ~3 cm
long, creeping, irregularly but freely branched, somewhat Plants medium- to large-sized, in thin to dense mats,
complanate-foliate; stems in cross section with 1–2(–3) glossy, light green to dark green or yellow-green. Stems
rows of epidermal cells. Leaves 0.8–1.2 mm long, erect- complanate-foliate or sometimes terete, creeping, simple
76 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

or sparingly and irregularly branched, cortical cells small, teeth cross-striolate below, papillose above, bordered,
thick-walled, central strand sometimes present; rhizoids trabeculate at back; endostome with a high to low basal
smooth, in clusters below leaf insertion usually on ventral membrane, keeled segments and cilia shorter than or ap-
surface of stems; pseudoparaphyllia large to small, foliose; proximately the same length as the segments, in groups of
axillary hairs consisting of 1–3 short, brown basal cell(s) 1–3. Spores 7–28 µm in diameter, globose to ovoid, smooth
and 1 elongate, hyaline distal cell. Asexual reproductive to minutely papillose. Calyptra cucullate, smooth, naked.
bodies unknown. Stem and branch leaves similar, rigid Distribution: Taxiphyllum is a genus of approxi-
or occasionally somewhat flaccid, crowded and imbricate mately 30–35 species occurring in terrestrial habitats, pri-
to distant, erect, erect-spreading or wide-spreading, often marily on calcareous substrates, in temperate, subtropical,
complanate, rarely secund at the subjulaceous, tips, some- or tropical regions.
times smooth or occasionally plicate, plane to concave, sym- Discussion: This is an easy genus of hypnaceous
metric or nearly so, nondecurrent, ovate, ovate-lanceolate, mosses to recognize because of the distinctive fish scale ap-
oblong-lanceolate, oblong-ovate or linear-oblong, acute pearance of the leaves, especially evident at magnifications
to subobtuse; margins plane or recurved, serrate to serru- of ~25–50×. Other more important distinguishing char-
late above, sometimes the serrations are bifid at the leaf acters are the medium- to large-sized, simple or sparingly
apices (only in T. laevifolium), serrulate to entire below; branched, complanate or subjulaceous, glossy plants, the
costa lacking or short and double; cells firm-walled, linear- small, thick-walled cortical stem cells, the smooth rhi-
flexuose, rhomboidal near leaf apices, smooth or prorulose zoids on the stems below the leaf insertions, the presence
at upper ends on dorsal leaf surface, with walls not pit- of foliose pseudoparaphyllia, the nondecurrent, serrate to
ted; alar regions distinctly differentiated with 1 to several serrulate leaves, the lack of asexual reproductive bodies,
rows of quadrate to rectangular cells, or not to nearly un- the dioicous or rarely synoicous sexual condition, and the
differentiated. Dioicous or rarely synoicous, usually sterile; presence of an annulus.
perigonia and perichaetia at base of stems and branches; Iwatsuki (1963) clarified the taxonomy of Taxiphyl-
perigonial bracts small, ovate; perichaetial bracts lanceo- lum, while Ireland (1969) revised the genus in Canada and
late to ovate, slenderly acuminate, spreading from an erect the United States and later published a synopsis (Ireland,
base; margins plane. Setae smooth, elongate, straight or 1986) of the North American taxa. Buck (1998) published
somewhat flexuose, dark red to brown; capsules inclined to a treatment of the genus for the West Indies and provided
cernuous, straight to subarcuate, yellowish brown to red- excellent illustrations for the four species that occur there.
dish brown, smooth, oblong-ovoid, somewhat contracted However, one species that he places in the genus, namely,
below mouth and wrinkled at neck when dry; operculum T. scapellifolium (Müll. Hal.) Broth. in Engl., the senior au-
obliquely rostrate, often shorter than urn; annulus present, thor considers better placed in the genus Mittenothamnium
persistent, of 2 rows of cells; peristome double, exostome (see Excluded Taxa).

KEY TO THE SPECIES OF TAXIPHYLLUM

1. Alar regions strongly differentiated with a large region of quadrate to short-rectangular cells, 3–9 differentiated cells extend-
ing up margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. T. deplanatum
1. Alar regions undifferentiated or weakly differentiated with only a few quadrate to short-rectangular cells, 1–5 differentiated
cells extending up margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Leaves usually remote, wide-spreading to squarrose, acute to acuminate, rarely subobtuse; margins narrowly and indis-
tinctly recurved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. T. taxirameum
2. Leaves usually close, erect-spreading, acute to obtuse, margins plane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Leaves ovate- to oblong-lanceolate, apices broad, often irregularly notched with cells ending at different levels, 0.8–1.5 mm
long, 0.4–0.7 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. T. laevifolium
3. Leaves linear-oblong, apices narrow, ending in a single or a few terminal cells, 0.9–2.5 mm long, 0.2–0.3 mm wide . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. T. ligulaefolium

1. Taxiphyllum deplanatum deplanatum Schimp. ex Müll. Hal., Syn. Musc. Frond.

2: 260. 1851; Isopterygium deplanatum (Schimp. ex
FIGURE 32
Müll. Hal.) Mitt., J. Linn. Soc., Bot. 12: 498. 1869; Pla-
Taxiphyllum deplanatum (Schimp. ex Müll. Hal.) giothecium deplanatum (Schimp. ex Müll. Hal.) Spruce,
M. Fleisch., Musci Buitenzorg 4:1435. 1923. Hypnum J. Bot. 18: 289. 1880. Type: United States, Sullivant,
 number 93 • 77

FIGURE 32. Taxiphyllum deplanatum. A. Habit. B. Enlargement of stem. C. Leaves. D. Pseudoparaphyllium. E. Apical leaf cells. F. Median leaf
cells. G. Marginal leaf cells. H. Alar cells. Illustrations A, B drawn from Sharp 3939 (TENN); C–H from Sharp 8728b (TENN).

Musci Allegh. 50 (lectotype: NY!, designated by Ire- per Sabino Canyon, 2743 m, Bartram, Mosses S. Arizona 161 (isotypes:
land, 1969); isolectotype: US!). CANM!, DUKE!, NY!, US!, WTU!).

Isopterygium planissimum var. laxirete Thér., Smithsonian Misc. Collect. 85 Plants in thin to dense mats, glossy, light green to
(4): 43. 1931. Type: Mexico, Michoacán, vicinity of Morelia, Loma Santa yellow-green. Stems to 2.5 cm long, 1–3 mm wide, com-
María, 2000 m, Arséne 4877 (isotypes: FH!, US!). planate-foliate, simple or irregularly branched. Leaves 1.0–
2.5 mm long, 0.4–1.0 mm wide, flaccid, usually crowded,
Plagiothecium deplanatum f. ovatum Grout, Moss Fl. N. Amer. 3(3): 161. imbricate, erect- to wide-spreading, somewhat concave
1932. Type: Arizona, Pima Co., Santa Catalina Mts., Mt. Lemmon, Up- or flat, smooth, ovate to ovate-lanceolate, symmetric to
78 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

somewhat asymmetric, acute or acuminate, somewhat foliate plants, the imbricate, erect to wide-spreading,
contorted when dry; margins plane, serrate to serrulate slightly concave or flat, ovate to ovate-lanceolate, acute or
above, serrulate below; costa very short and double or acuminate leaves with plane margins, and the noticeably
lacking; median cells 47–136 × 5–9 µm, smooth; alar cells differentiated alar regions of 1 to several rows of quadrate
strongly differentiated, quadrate to short-rectangular, in 1 to short-rectangular cells with 3–9 differentiated cells in
to several rows with 3–9 cells in the marginal row. Sex or- the marginal rows.
gans and sporophytes not seen on Latin American plants. The Latin American plants of T. deplanatum are notice-
North American plants reported (Ireland, In press b) to be ably different from their North American counterparts in
dioicous; perigonia unknown; perichaetia small, numer- that the leaves are more distant, wide-spreading and they are
ous, the bracts lanceolate to ovate-lanceolate, acuminate. somewhat contorted when dry. Microscopically, the plants
Setae 0.7–1.0 cm long, yellowish brown to red, flexuose; were observed to be similar to the North American ones.
capsules 0.8–1.5 mm long, cernuous, straight or arcuate, Specimens examined: MEXICO. Durango: ~14.5
yellowish brown to light brown, oblong or ovoid, con- km W of La Ciudad, 2637 m, Bowers et al. 5083c (CANM,
tracted below mouth and tapered to a wrinkled neck when TENN). Hidalgo: Above Tianguistengo, 3.2 km from
dry; operculum 0.3–0.5 mm, obliquely rostrate; annulus Zacualtipan, 2100 m, Sharp et al. 1825a (CANM, TENN),
present, persistent. Spores 11–13 µm in diameter. 1837 (CANM, F, MEXU, TENN). Morelos: ~11 km above
Distribution and ecology: Rare or seldom col- Cuernavaca, 2200 m, Düll 2/276 (MICH). San Luis Potosí:
lected; Mexico and Honduras (Figure 33); also in eastern Just E of San Francisco, ~45 km E of San Luis Potosí at
and southwestern North America; on shaded soil or lime- K[m]-218 along Hwy. 70, 4260 m, Magill 2609 (MO, NY).
stone boulders and cliffs, sometimes on bases of trees and Tamaulipas: Edge of Casa de Piedra, near Rancho del Cielo
exposed tree roots, at 1000–2600 m. above Gómez Farias, Sharp 8728b (TENN); trail to Indian
Discussion: Taxiphyllum deplanatum is recog- Spring, Rancho del Cielo above Gómez Farias, 1100–1200
nized by the flaccid, light to yellow-green, complanate- m, Iwatsuki & Sharp 4845 (TENN). Veracruz: Txolo Falls,

FIGURE 33. Distribution of Taxiphyllum deplanatum (∆), T. laevifolium (▲), and T. ligulaefolium (◆) in Latin America. Colombia locality of
T. laevifolium uncertain.
 number 93 • 79

near Jalapa, 1006 m, Sharp 3939 (MICH, TENN). HON- The leaf margins in this species sometimes have bifid
DURAS. Comayagua: Vicinity of Siguatepeque, ~1050 m, serrations on the apical leaf cells which is very unusual
Standley & Chacón 6151 (F, FH). and it was only observed in the single collection from Ve-
racruz, Mexico. This morphological feature was noted by
2. Taxiphyllum laevifolium H. A. Crum (1994a), who placed the species in the genus
Glossadelphus. However, the illustration in Crum (1994a:
FIGURE 34 fig. 747e) does not show the bifid teeth; they are instead
shown for G. ligulaefolius (Crum, 1994a: fig. 747b). This
Taxiphyllum laevifolium (Mitt.) W. R. Buck, Mem. New is undoubtedly an error in the illustration of the species
York Bot. Gard. 45: 521. 1987. Ectropothecium laevifo- since the description states that G. laevifolius has “ser-
lium Mitt., J. Linn. Soc., Bot. 12: 517. 1869. Type: Co- rations often minutely bifid.” However, the Mexican
lombia, Andes Bogotenses, Weir 425 (holotype: NY!). plants may have much larger bifid serrations than shown
in the illustration (see collection from Veracruz, Mexico,
Glossadelphus laevifolius (Mitt.) E. B. Bartram, Bull. Brit. Mus. (Nat. Hist.), at TENN). No other species of Taxiphyllum is known
Bot. 2: 49. 1955. to have bifid serrations from Latin America or, as far as
we know, from anywhere else in the world. This could
Plants in thin to dense mats, glossy, light green to yellow- be a reason for placing this species in another genus, but
green. Stems usually ~2 cm long, rarely to 7 cm, 1.5–2.0 mm further study is necessary to decide whether it belongs in
wide, complanate-foliate, usually irregularly branched. Glossadelphus or another genus. Also, although Crum in-
Leaves 0.8–1.5 mm long, 0.4–0.7 mm wide, rigid, imbricate, dicates that G. laevifolius does not have prorulae on cells
erect to erect-spreading, slightly concave, smooth, ovate- to on the dorsal surface of the leaf, we have found that they
oblong-lanceolate, symmetric, or somewhat asymmetric, are often present.
acute to obtuse, the apices broad, irregularly notched with Although the type collection (NY) from Colombia
the cells ending at different levels; margins plane, serrate to was described as monoicous, the senior author was unable
serrulate above, sometimes the serrations bifid at leaf apices, to find any antheridia on plants with sporophytes when
serrulate to entire below; costa lacking or weak, short and the specimen was examined. The only Jamaican collection
double; median cells 38–80 × 5–6 µm, smooth or often pro- contained plants with synoicous inflorescences but with no
rulose at upper ends on dorsal surface; alar cells poorly dif- sporophytes. Synoicous pleurocarpous mosses are usually
ferentiated, long- to short-rectangular or rarely quadrate, in rare, with the exception of members of the Hookeriales,
a few rows with 1–4 cells in the marginal rows. Synoicous. and only a very small number of species in other orders are
Setae 1.2–1.5 cm long, red, flexuose; capsules 0.7–1.0 mm known with this sexual condition.
long, inclined, straight or nearly so, ovoid, often contracted Specimens examined: MEXICO. Veracruz: Road
below mouth when dry; operculum unknown; annulus pres- to Bastonal, above Tebanca, 700 m, Richards, A.J. & E.B.
ent, persistent. Spores 23–28 µm in diameter. Sharp 5863b (TENN). PANAMA. Bocas del Toro: Vicinity
Distribution and ecology: Rare or seldom col- of Fortuna Dam, along ridge below Continental Divide on
lected; Mexico, Panama, Jamaica and Colombia (Figure Chiriqui Grande road, 800 m, 08°55'N, 82°08'W, Allen
33); occurring on lava boulders, rocks in woods, beside 5831 (MO). JAMAICA. Portland: Forest Reserve Area W
streams and near waterfalls or on trunks of fallen trees, at of Hardwar Gap, 1219–1295 m, Crosby 2923 (DUKE).
~700–1300 m. Also reported from Costa Rica, Galápagos COLOMBIA. Known only from the type locality.
Iss., Cocos Iss., and Dominica (Crum, 1994a, as Glossadel-
phus laevifolius). 3. Taxiphyllum ligulaefolium
Discussion: This species is distinguished by the rigid,
light to yellow-green, complanate-foliate plants, the imbri- FIGURE 35
cate leaves that are erect to erect-spreading, slightly concave,
ovate- to oblong-lanceolate with plane margins, acute to ob- Taxiphyllum ligulaefolium (E. B. Bartram) W. R. Buck,
tuse apices that are often irregularly notched with the cells Trop. Bryol. 2: 42. 1990. Glossadelphus ligulaefolius
ending at different levels, the serrate leaf margins at apices E. B. Bartram, Bryologist 49: 123. 1946. Type: Guate-
sometimes bifid and the poorly differentiated alar regions of mala, Dept. Izabal, jungle between Escobas and water-
a few rows of short-rectangular or rarely quadrate cells with fall, across bay from Puerto Barrios, 20–25 m, Steyer-
1–4 differentiated cells in the marginal rows. mark 39846 (holotype: FH!).
80 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 34. Taxiphyllum laevifolium. A. Habit. B. Enlargement of stem. C. Leaves. D. Pseudoparaphyllia. E. Apical leaf cells. F. Marginal leaf
cells. G. Median leaf cells. H. Alar cells. Illustrations A–D drawn from Weir 425 (NY); E–H from Crosby 2923 (DUKE).

Taxiphyllum gallorum W. R. Buck, Brittonia 36 (2): 178. 1984b. Type: Ja- rigid, close, erect-spreading, complanate, slightly con-
maica, Trelawny Parish, Cockpit Co., just N of Wilson’s Run, 5.8 mi. N of cave at base, smooth, linear-oblong, symmetric, abruptly
bridge at Troy, 500 m, 18°16'N, 77°40'W, Buck 5888 (holotype: NY!). acute to subobtuse, ending in a single or a few terminal
cells; margins plane, sometimes erect below, serrate to ser-
Plants in thin mats, glossy, yellowish green. Stems to rulate above, serrulate to subentire below; costa usually
1 cm long, 2–3 mm wide, complanate, simple or irregularly weak, short and double, sometimes lacking; median cells
branched. Leaves 0.9–2.5 mm long, 0.2–0.3 mm wide, 47–94 × 4–5 µm, smooth or prorulose at upper ends on
 number 93 • 81

FIGURE 35. Taxiphyllum ligulaefolium. A. Habit. B. Enlargement of stem. C. Leaves. D. Apical leaf cells. E. Marginal leaf cells. F. Median leaf
cells. G. Alar cells. All drawn from Crosby 13875 (NY).

dorsal surface; alar cells usually differentiated, quadrate to Discussion: Taxiphyllum ligulaefolium is a very
short-rectangular, 2–4 extending up margins. Sex organs distinctive species that is easily recognized by the leaves
and sporophytes unknown. that are close, erect-spreading, linear-oblong, with abruptly
Distribution and ecology: Very rare or seldom col- acute to subobtuse apices usually ending in a single termi-
lected; known only from Mexico, Guatemala, Belize, and nal cell, the serrate to serrulate margins, the narrow median
Jamaica (Figure 33); occurring on limestone in humid for- leaf cells, and the small region of differentiated alar cells,
ests, sometimes near waterfalls, up to 1158 m. 2–4 quadrate to short-rectangular cells on margins.
82 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

It is of interest to note that the serrations on the mar- 0.3–0.6 mm wide, rigid, usually distant, wide-spreading,
gins near the leaf apex of the only Mexican plant (Puebla, flat or somewhat concave, smooth or plicate, ovate- to
Santos 3694 MICH) known from the country thus far oblong-lanceolate, symmetric or nearly so, acuminate or
are described as simple in the Glossadelphus key (Crum, abruptly narrowed to an acute or rarely subobtuse apex;
1994a: 1002) and description (Crum, 1994a: 1003), which margins very narrowly recurved almost to apex, serrulate
we consider a synonym of this species (as G. ligulaefolius to serrate throughout; costa lacking or short and double;
E. B. Bartram), but illustrated as minutely bifid (Crum, median cells 66–120 × 3–7 µm, smooth or prorulose at
1994a: 1004, fig. 747b). However, no such minute bifid upper ends on dorsal leaf surface; alar cells poorly dif-
serrations were observed in any of the Mexican plants ferentiated, long- to short-rectangular or rarely quadrate,
from the same collection (seen at FH but not at MICH) or in 1–3 rows with 1–5 cells in the marginal rows. Sex or-
any of the plants from elsewhere that the authors studied. gans and sporophytes not seen on Latin American plants.
This species can be confused with a Pseudotaxiphyl- Reported and observed to be dioicous; perigonia small,
lum because of the few foliose pseudoparaphyllia present bracts ovate, acute; perichaetia small, bracts lanceolate to
on its stems. ovate-lanceolate, acuminate. Setae 0.7–1.2 cm long; cap-
Specimens examined: MEXICO. Puebla: Near sum- sules 1.0–1.5 mm long, reddish brown. Spores 11–13 µm
mit of El Cerro de Cuhuatepetl, Tehuacan, 945–1158 m, in diameter.
Santos 3694 (FH). GUATEMALA. Known only from type Distribution and ecology: Common; West Indies,
locality. BELIZE. Toledo, Columbia River Forest Reserve, Mexico to Costa Rica, Colombia, Surinam, Ecuador, Bra-
Union Camp, 680 m, 16°23'N, 89°08'W, Allen 18552 zil, and Bolivia (Figure 37); also known from eastern North
(MO). JAMAICA. Same locality as type of Taxiphyllum America and eastern Asia; usually on moist, shaded lime-
gallorum, Crosby 13875 (NY). stone rocks and cliffs in woods, sometimes on soil banks be-
side streams and along roadsides, rarely on logs and bases
4. Taxiphyllum taxirameum of trees, at 40–2500 m.
Discussion: This is a distinctive species recognized
FIGURE 36 by the dark green to yellow-green, rigid, complanate-foliate
plants, the distant, wide-spreading, flat or somewhat con-
Taxiphyllum taxirameum (Mitt.) M. Fleisch., Musci Bu- cave, ovate- to oblong-lanceolate, acuminate or acute to
itenzorg 4: 1435. 1923. Stereodon taxirameus Mitt., rarely subobtuse, sometimes plicate leaves with very nar-
J. Linn. Soc., Bot., Suppl. 1: 105. 1859; Isopterygium rowly recurved margins, and the poorly differentiated alar
taxirameum (Mitt.) A. Jaeger, Ber. Thätigk. St. Gal- regions of 1–3 rows of quadrate to rectangular cells with
lischen Naturwiss. Ges. 1876–77: 439. 1878. Type: 1–5 cells in the marginal rows. It is the most common spe-
In[dia], Himalayae reg. temp., Simla et Kumaon, Thom- cies of Taxiphyllum in Latin America, and it is especially
son 1023 (lectotype: NY!, designated by Iwatsuki and prevalent in parts of Mexico and the West Indies.
Crosby (1979: 393); 1023b cited by Mitten in the type Selected specimens examined: MEXICO. Chiapas:
description is apparently an error; Ceylon, Gardner s.n. Rancho Monte Armenio, Ejido Chintul, Tecpatán, 300 m,
(syntype: NY!). Zamudio & Rodríguez 1499 (MEXU); Santa Elena, 650 m,
16°53'N, 91°35'S, Delgadillo M. 6324 (MO); above Paval
Isopterygium planissimum Mitt., J. Linn. Soc., Bot. 12: 498. 1869; Taxiphyl- N of Mapastepec, 1524 m, Sharp 4443 (MICH, TENN).
lum planissimum (Mitt.) Broth. in Engl., Nat. Pflanzenfam. ed. 2, 11: 463. Chihuahua: At bridge Puerto de Río Urique, 61 km S of
1925; Plagiothecium planissimum (Mitt.) E. B. Bartram, Bryologist 49(4): Creel, 1707 m, Bowers et al. 5356 (CANM, MEXU,
122. 1946. Type. Jamaica, Wilson 675 (lectotype: NY!, here designated). TENN), 5330-b, 5332-c, 5362b (TENN). Durango: Near
Palos Colorados, W of Durango, 2499 m, Sharp 1499c
Isopterygium cavernicola Cardot, Rev. Bryol. 37: 56. 1910; Taxiphyllum (TENN). Hidalgo: 82.2 km SW of Tempoal on Molango
cavernicola (Cardot) Thér., Rev. Bryol. Lichénol. 5: 109. 1932. Type: Mex- Hwy., Spessard 696a (TENN); ~26 km S of Tamazunchale,
ico, Morelos, near Cuernavaca, Pringle, Plantae Mexicanae 15179 (isotypes: Bowers et al. 5049-a (TENN). Jalisco: Puente San Pedro,
H!, NY!). 8 km SW of Tecalitlán, ~1200 m, 22 Sep 1958, McVaugh
s.n. (MICH, NY); Arroyo de La Resolana, near Sapotillo,
Plants in thin to dense mats, glossy, dark green to yel- 305 m, Crum 643 (MICH, NY, TENN); near summit of La
low-green. Stems to 5 cm long, 2–4 mm wide, compla- Cumbre, 16 km SW of Autlán, 1524 m, Crum 1345 (MICH,
nate, simple or irregularly branched. Leaves 1–2 mm long, TENN, US); 27 km W of Ameca on road to Los Volcanes,
 number 93 • 83

FIGURE 36. Taxiphyllum taxirameum. A. Habit. B. Enlargement of stem. C. Leaves. D. Pseudoparaphyllia. E. Apical leaf cells. F. Median leaf
cells (smooth). G. Median leaf cells (prorulose). H. Marginal leaf cells. I. Alar cells. Illustrations A, B drawn from Magill 2609 (TENN); C (two
on left), D (left), E (top), F, H, I from Sharp 4240 (CANM); C (one on right), D (right), E (bottom), G from Smith & Sharp 114 (CANM).

1500 m, 20°34'N, 104°15'W, Whittemore 2107 (MO). (CANM, TENN); ~3 km W of Compostela on road to
México: Hacienda Lanzarote, município de Villa Nicolás Puerto Vallarta, Norris & Taranto 15161(CANM, TENN);
Romero, 2350 m, Cárdenas 3820a (MEXU). Michoacán: 50 km E of Ruiz, 10.7 km E of San Pedro Ixcatán on road
Morelos Natl. Park, near Morelia, Norris & Taranto 15528 to Mesa de Nayar, 109 m, 22°02'N, 104°50'W, Whitte-
(CANM, TENN); Cascade de Caradacuan in Río Cupatzio, more 3227 (MO). Nuevo Léon: Cañon de San Francisco
Patrick 615 (FH). Nayarit: ~6 km from Compostela on near Monterrey, Spessard 95 (CANM, TENN); Río Valle
road to San Pedro Jagunilla, Norris & Taranto 15079 Alto, Spessard 436, 441 (TENN); Chipinque, Sierra Madre
84 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 37. Distribution of Taxiphyllum taxirameum in Latin America.

Oriental, ~14 km NW of Monterrey, 1158 m, Hermann of Villa Aldama, Pursell 5458 (MO); South Mt., near Ran-
26039 (CANM), 26039b (US); Sierra Madre, near Monter- cho del Cielo, Gómez Farias, 1037 m, 23°15'N, 99°12'W,
rey, Pringle 15103 (NY). Oaxaca: Sierra Juárez above Río Manuel 396 (MO); roadside of Hwy. 101, 37 km SW of
Valle Nacional, 20 Dec 1962, Robinson s.n. (MEXU, NY, Victoria, 1098 m, Bowers et al. 2776 (NY); Paso de las
US); below Tamazulapam Mixe, ~70 km E of Oaxaca, Calabazas, along Río Sabinas, near Gómez Farias, Sharp
Smith et al. 2994 (CANM, MEXU, TENN), 3001 (CANM, 8763 (CANM); just E of San Francisco, 45 km E of San
TENN). Puebla: Below Zacapoaxtla, near Xalacapa, Luis Potosí, 2164 m, Magill 2609 (TENN). Tlaxcala: Near
1524 m, Sharp 4240 (CANM, MICH, TENN); vicinity of falls of Río Zuahuapan, near Amaxac, Sharp 401 (MEXU,
Puebla, 2200 m, Arsène 584 (US). San Luis Potosí: W of TENN). Veracruz: Sierra de Chiconquiaco, between Santa
Xilitia, 838 m, Sharp 5915 (MEXU, MICH, TENN, US); Rita and Bella Luz, ~40 km NW of Xalapa, 1750 m, Her-
above Xilitla, 762 m, Sharp 5856 (MEXU, MICH, TENN, mann 28844 (CANM, TENN); above Poza Rica on Hwy.
US); just E of San Francisco, 45 km E of San Luis Potosí, 130, 1000 m, Sharp et al. 1280 (CANM, TENN), 4858
2164 m, Magill 2609 (TENN); 49 km S of Antiguo More- (CANM, F, MEXU, TENN); El Esquilón, ~1300 m,
los, road to Huizache, Pursell 5367 (MO). Sinaloa: 45 km 19°37'N, 96°56'W, Buck 35518 (NY); La Concha, along
NE of Mazatlan, 152 m, Bowers et al. 5127 (CANM, F, Carretera La Concha a San Antonio, ~1000 m, 19°36'N,
MEXU, TENN). Tamaulipas: 14 km above San Antonio on 96°53'W, Buck 35406, 35416 (NY); just S of city of Xa-
Hwy. 101, 1773 m, Sharp et al. 1002-b (TENN); ~16 km lapa on road to Coatepec, Instituto de Ecología, Rancho
from Victoria on road to San Luis Potosí, 914 m, Smith & Guadeloupe, ~1300 m, 19°31'N, 96°56'W, Buck 35380
Sharp 112, 114 (CANM, TENN), 116 (TENN); ~16 km N (NY). Zacatacas: 7 km S of Laguna Grande, N of Monte
 number 93 • 85

Escobedo, 2030 m, Cárdenas 3057b (MEXU), 3028 Yara, Pócs 9073c (NY). Trinidad Mtns., Santa Clara, León
(MEXU, NY); 8 km E of Monte Escobedo, 2030 m, Cárde- 14149 (NY). JAMAICA. Manchester: Marshall’s Pen, 5 km
nas 3070 (MEXU, NY). GUATEMALA. Alta Verapaz: WNN of Mandeville, 701 m, Crosby 13825 (NY); 4 km N
Cerro Chinajá, between Finca Yalpemech and Chinajá, of Plowden Hill, 500 m, Crosby 13848 (MO, NY). St. An-
150–700 m, Steyermark 45564 (F, FH); 3 km N of Coban, drew: Newcastle road, between 16 & 17 mile posts, 762–
1341 m, Sharp 2994 (FH). Baja Verapaz: Patal, 1615 m, 1069 m, Farr 104 (CANM). St. Catherine: Byndloss Gully,
Sharp 2827 (FH, TENN). Chimaltenango: Quisaché, 1.6 km N of Linstead, 91 m, Farr 106 (CANM). St. Thomas:
1800 m, Standley 62310 (F, FH). Quezaltenango: Above Corn Puss Gap and vicinity, ~14 km N of Bath, 550 m,
Santa María de Jesús, ~1650 m, Standley 67279 (F, FH). Buck 5561 (NY); Bath, along Sulphur River, 91 m, Crosby
Retalhuleu: Near Chivolandia (Dept. Quezaltenango) along 13631 (MO, NY). Trelawny: Vicinity of Windsor Cave,
road to San Felipe, ~650 m, Standley 87209a (F), 87213 140 m, Buck 5932 (NY); Cockpit Country, just N of Wil-
(DUKE, F, FH, MICH, NY). Sacatepéquez: Above Pastores, son’s Run, 9 km N of bridge at Troy, 500 m, Crosby 13879
~1680 m, Standley 60768 (F, FH, MICH, NY, US). Zacapa: (MO, NY). HAITI. Vicinity of St. Louis du Nord, Morne
Trail between Río Hondo and waterfall, 250–400 m, Stey- Colombot, Leonard & Leonard 14249 (NY, US), 14319
ermark 29413 (F, FH, MICH). BELIZE. Cayo: S side of (F, FH, NY, US). Vicinity of Port au Prince, Pétionville,
Rapaculo Branch, 16°49'N, 88°56'W, Allen 18458, 18474 Leonard & Leonard 15792 (FH, NY, US). Nord: vicinity of
(MO). Toledo: Solomon Camp, vicinity of junction of Rich- Marmelade, 800 m, Leonard 8139 (CANM, FH, NY, US).
ardson creek and Bladen Branch, foothills of the Maya Sud’est: Massif de la Selle, 19 km S of Kenscoff, 1600 m,
Mts., 80–420 m, Davidse & Brant 32073B (MO); Upper Buck 9015 (NY). Vicinity of Bombardopolis, 610 m, Leon-
Bladen Branch, Bladen Nature Reserve, 16°54'N, 88°55'W, ard & Leonard 13560 (FH, US). Les Trois Sources, between
Whittemore 5800 (MO, NY); Gabriel Camp-Edwards Cen- Chambellan and Cayaba, 442 m, Bartlett 17792 (DUKE,
tral Camp, Boutin & Schlosser 5155 (MO); Columbia MICH). DOMINICAN REPUBLIC. La Altagracia: Be-
River Forest Reserve, 16°22'N, 89°07'W, Allen 18678 tween Higüey and Miches, ~20 km N of Otra Banda, Jones
(MO). HONDURAS. Comayagua: Vicinity of Siguate- & Norris 1403A, 1405A (NY). Dajabón: Cañada Tirolis,
qeque, ~1050 m, Standley & Chacón 6599 (F); Quebrada 1 km S of Villa Anacaona, 396 m, Reese 15386, 15390
Agua Helado, above San José del los Planes, ~15 km E of (NY). La Estrelleta: Sierra de Neiba, from 8 to 15 km N of
Lago Yojoa, cerro Meámbar, 14°47'N, 87°51'W, Allen Angel Féliz, 1402–1829 m, Buck 4605 (NY). La Vega: Vi-
13717 (MO). Cortés: Along Río Lindo, N of Lake Yojoa, cinity of Piedra Blanca, 200–500 m, Allard 16886, 16941d,
500–600 m, Morton 7875, 7876 (US); near Jaral, 600 m, 16962 (NY, US). Pedernales: 4 km E of Mencia, 457 m,
Standley 7021 (F); Nacimiento del Río Lindo, near Jaral, Reese 15073 (NY). Samaná: Parque Nacional Los Haitises,
~600 m, Standley 7075 (F). El Paraíso: Vicinity of Danlí, S of Naranjo, Smith 10416 (NY); Hacienda Nydia, 3.5 km
700–800 m, Standley 16394 (F). Olancho: Slopes of Mon- E of Las Terrenas, 122 m, Steere 23193 (NY). San Cris-
tana Pena Blanca, 11.5 km S of Gualaco on road from San tóbal: Mano Matuey, 12 km from Cambito Garabito, 549
Francisco de La Paz to Gualaco, 980–1000 m, 14°57'N, m, Buck 5159, 5197, Reese 15608, 15622, 15628 (NY).
86°08'W, Allen 12405, 12426 (MO). EL SALVADOR. Sánchez Ramírez: Loma La Guacara, N of Comedor Ar-
Ahuachapán, near Ataco, 1200–1400 m, Standley & Pa- riba, ~5 km S of Fantino, 100–240 m, Zanoni & Peláez
dilla 2689, 2694, 2701 (F). NICARAGUA. Jinotega: Be- 16185 (MO, NY, U). PUERTO RICO. Arecibo, between
tween Las Camelias and La Salvadora, along tributary of Manati and Morovis, ~300 m, Buck 16149 (DUKE, MO,
Río Jiguina, ~13°05'–06'N, 85°53'–54'W, Stevens & Gri- NY). ~6 km SSW of Florida, just S of junction with Río
jalva 15359 (MO). COSTA RICA. Cartago: Río Macho, Yúnes, Reese 14434 (NY). Cordillera Central, 13 km S of
1100 m, Morales et al. 963 (USJ). Puntarenas: ~2 km SE of Utuado, 6 km N of Adjuntas, Reese 14511, Buck 3759,
Monteverde, 1500–1550 m, Gentry & Burger 2755D 3763 (NY). Cordillera Central, above Villalba, Dona Juana
(MO). Aserri, 1400 m, Alfaro 70b (FH). BERMUDA. Recreation Area, 800-900 m, Allen 6390 (MO). Between
Tucker’s Town, Church Cave, Britton 418 (NY). CUBA. Arecibo and Utuado, Britton 2077 (F, FH, MICH, NY).
Guantánamo: Loma Lagunato, ESE of El Ramón, 300–400 LEEWARD ISLANDS. Nevis: St. George Gingerland Parish,
m, Reyes 9193B (NY). Oriente: Loma del Gato, Hioram Golden Rock Estate, Golden Rock Nature Trail, ~300 m,
12044 (F). Pinar del Río: Sierra la Buira, Leon 4710 (NY). 17°08'N, 62°34'W, Buck 29538 (NY). WINDWARD
Santo Domingo: Barrabos, 400 m, 1887, von Eggers s.n. ISLANDS. Guadeloupe: Rivière Beaugendre, LeGallo 651
(H). Santiago de Cuba: La Gran Piedra, 1000 m, Buck (CANM), 705 (MICH). COLOMBIA. Antioquia: Mpio.
7609, Shaw 5098, 5153 (NY). Seibo: Azui, Taylor 278b Liborina, N of Liborina en la carretera a Sabanalarga, 910–
(NY). Sierra Maestra: “El Peru,” in the valley of the Río 1030 m, Churchill et al. 14653 (NY); Mpio. Dabeiba,
86 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

2–3 km NE of Dabeiba, 460 m, Churchill et al. 17048 The senior author agrees with Crum (1968) that this
(CANM, NY). Caldas: Mpio. of Río Sucio, on the Río species belongs in the genus Mittenothamnium, as M. sca-
Sucio, Vereda el Tabor, 1900–2080 m, Churchill & Arbe- pellifolium (Müll. Hal.) H. A. Crum, primarily because
láez 15646 (NY). Huila: Transecto Neiva, San Vicente del of the different type of branching pattern that has never
Caguán, Río Las Ceibas, 680 m, Churchill & Betancur been seen to occur in other taxa of the genus Taxiphyllum.
16767 (CANM, NY). Nariño: Mpio. Alban, Campo The main stems are creeping, wiry, with small, spreading,
Bello, 01°28'N, 77°04'W, Ramírez P. 5868 (MO); Mpio. ovate-lanceolate, acuminate leaves that differ strikingly
de Barbacoas, km 1 El Mirador–Junín, 1500 m, 01°08'N, from the complanate, oblong to ovate-oblong, abruptly
78°09'W Ramírez P. 3631 (NY); Mpio. de Sandoná, short-acuminate to mucronate leaves of the secondary
1900 m, Ramírez 5678 (NY). SURINAM. Without local- stems. This type of branching pattern with dimorphic
ity, Korthals s.n. (H). ECUADOR. Azuay: Gualaquiza, leaves does not occur in the genus Taxiphyllum. Although
Jun 1909, Allioni 8343 (H). BRAZIL. Paraná: Mpio. Foz the complanate secondary leaves are unlike those of any
do Iguaçu, Parque Nacional do Iguaçu, forest along Rio species known in the genus Mittenothamnium, the wiry
Iguaçu, 100–200 m, ~25°40'S, 54°25'W, Vital & Buck creeping stems giving rise to secondary stems do occur in
12074 (NY). São Paulo: Serra de Paranapiacaba, Eldo- the genus. Crum placed it in the genus Mittenothamnium
rado, Caverna do Diabo, 500 m, Schäfer-Verwimp & because of the stipitate growth habit. He also mentions the
Verwimp 11082 (CANM). BOLIVIA. Santa Cruz: Aneres prorulose leaf cells as a reason for placing it in the genus,
Ibanes, Espejillas, 20 km W of San José, 17°56'S, although prorulose leaf cells are also known in species of
63°26'W, Fuentes 140A (MO). Taxiphyllum. In spite of this the senior author considers
further studies to be necessary before it is prudent to place
Taxiphyllum Excluded Taxa the species in Taxiphyllum. Buck (1998: 326) is “not com-
pletely convinced of a relationship within Taxiphyllum,”
Taxiphyllum scapellifolium (Müll. Hal.) Broth. in Engl., but he has left it in the genus for lack of a better one in
Nat. Pflanzenfam. ed. 2, 11: 462. 1925. which to place it.
Acknowledgments

W
e are very grateful to the following curators for the loan of speci-
mens: J. Aguirre C. (COL); B. H. Allen, M. R. Crosby, and R. E.
Magill (MO); H. A. Crum† (MICH); C. Delgadillo M. (MEXU);
A. Eddy† and A. Newton (BM); J. J. Engel (F); T.-B. Engelmark
(S); P. Geisler (G); M. F. Goncalves-Carralves and G. G. Hassel de Menéndez
†

(BA); S. R. Gradstein (U); H. Hertel (M); H. A. Imshaug (MSC); P. Isoviita (H);

E. Lawton† (WTU); M. Lewis (LPB); R.C. L. Lisbôa (MG); F. K. Meyer (JE);
B. D. Mishler (DUKE); R. Moberg (UPS); G. S. Mogensen (C); M. I. Morales
(USJ); D. H. Pfister (FH); H. E. Robinson (US); N. Salazar A. (PMA); A. J.
Sharp† and D. K. Smith (TENN); and D. M. Vital (SP). We also thank M. Kuc,
Ottawa, Canada, for allowing us to examine his South American collections
and A. Schäfer-Verwimp, Museum für Naturkunde Stuttgart, Germany, for the
many Brazilian specimens he sent. We thank Bruce Allen (MO) for his valuable
time in correcting mistakes and adding comments to the manuscript, as well as
aiding in preparing the illustrations and maps. We appreciate the assistance of
Linda M. Ley (CANM) for her comments on the manuscript and for her techni-
cal expertise, particularly for the tedious and time-consuming job of preparing
the illustrations as well as for her help with the maps. Halina Bednarek-Ochyra
(Institute of Botany, Polish Academy of Sciences, Kraków, Poland) did the il-
lustrations of Elmeriobryum for the junior author, and we are very grateful for
her excellent work. We thank Marjorie Knowles, formerly a Research Assistant
at the Smithsonian Institution, for her assistance with revising the maps with
the computer, for correcting mistakes in the manuscript, and for her computer
skills in various other aspects of preparing this paper. We are indebted to Joe
Ireland for revising the maps and illustrations for publication and for his help
with numerous computer problems. Wilf Schofield† (UBC) contributed his time
in correcting mistakes in the manuscript, for which we are grateful. Also, to
Harold Robinson, Smithsonian Institution, the senior author owes a debt of
gratitude for providing space, equipment, and assistance to finish the studies for
this paper. We thank Paul Peterson of the Smithsonian Institution for his help
with the review process of the manuscript; and we also acknowledge the excel-
lent work by the Smithsonian Scholarly Press, especially Meredith McQuoid
88 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

and Tshawna Byerly, for seeing this paper through to of Nature (Ottawa, Canada) as well as Ruth Schallert,
publication. Finally, we are grateful to A. Stewart, J.-G. Smithsonian Botany Library, for their help in obtaining
Brisson, and other library staff of the Canadian Museum many literature references for this study.
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Index of Scientific Names

Accepted names of taxa are in regular type and synonyms are in italics. A bold page
number indicates the location of the main entry for the taxon; figure numbers are in
parentheses.

Acanthocladiella M. Fleisch., 71, 72

flexilis (Renauld & Cardot) M. Fleisch., 71
Aptychella (Broth.) Herzog, 1

Brotherella M. Fleisch., 1
Bryosedgwickia Cardot & Dixon in Dixon, 54
densa (Hook. in Kunth) Bizot & P. de la Varde, 55
densa var. brevifolia (Cardot) Bizot & P. de la Varde, 56
kirtikarii Cardot & Dixon in Dixon, 54
pallidissima (Müll. Hal.) Bizot & P. de la Varde, 55
pringlei (Cardot) Bizot & P. de la Varde, 56
subdensa (Broth. & Thér.) Bizot & P. de la Varde, 55

Clastobryopsis M. Fleisch., 1
Clastobryum Dozy & Molk., 1
Ctenidiadelphus 11
cylindricarpus (Cardot) E. B. Bartram, 11
Ctenidium (Schimp.) Mitt., 5, 6
anacamptopteris (Müll. Hal.) Broth. in Engl. & Prantl, 6
aurifolium Mitt., 67
malacodes Mitt., 6, 7
muenchii Broth. ex H. A. Crum & Sharp, 6
subrectifolium (Brid.) W. R. Buck & B. H. Allen, 7
terrestre (Müll. Hal.) Broth. in Engl. & Prantl, 6
Cupressina anacamptopteris Müll. Hal., 6

Dimorphella (Müll. Hal.) Renauld & Cardot, 71

brasiliensis Broth. & Herzog in Herzog, 72
pechuelii (“i”) (Müll. Hal.) Renauld & Cardot var. acuminata Cardot, 73
pechuelii (“i”) (Müll. Hal.) Renauld & Cardot var. gracilis Cardot, 73
Dolichotheca Lindb., 10, 11, 24
94 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Ectropothecium Mitt., 2, 4, 5, 24, 55 eutrypherum Müll. Hal., 44

clandestinum Ångstr., 44 exiguum Geh. & Hampe in Hampe, 51
laevifolium Mitt., 79 fulvum Hook. & Wilson in Drumm., 43
lamprophyllum A. Jaeger, 44 gracillimum Hornsch., 51
laxum A. Jaeger, 44 hamatum (Mitt.) A. Jaeger, 69
longisetum Schimp. ex Besch., 39 hospitans Hampe, 44
submersum Broth., 26 incurvatum Brid., 14
trinitense (Müll. Hal.) Mitt., 72 lamprophyllum Hampe, 44
truncatulum (Müll. Hal.) Mitt., 53 laxum Hampe, 44
Elmeriobryum Broth., 4, 7, 8 leptomiton Müll. Hal., 44
guatemalense J. R. Rohrer, 8, 9 (Fig. 1), 10 (Fig. 2) leucophyllum Hampe in Müll. Hal., 44
philippinense Broth., 7 magellanicum Müll. Hal., 21
Entodon Müll. Hal., 24 micans Sw., 43
densus (Hook. in Kunth) Mitt., 55 miradoricum Müll. Hal., 52
pallidissimus Müll. Hal., 55 mittenohamatum B. C. Tan, 69
Erythrodontium Hampe, 54 pechuelii Müll. Hal., 71
densum (Hook. in Kunth) Paris, 55 polyanthum (Hedw.) Dicks., 70
densum var. brevifolium Cardot, 56, 57 restitutum Hampe, 44
imbricatifolium R. S. Williams ex Cardot, 56 sect. Dimorphella Müll. Hal., 71
longisetum (Hook.) Paris, 57 splendidulum Hornsch., 43
pallidissimus (Müll. Hal.) Paris, 55 subbrevisetum Hampe, 34
pringlei Cardot, 56 subg. Ctenidium Schimp., 5
subdensum Broth. & Thér. in Thér., 56 subg. Homomallium Schimp., 14
tenerum Sw., 43
Giraldiella Müll. Hal., 68 trinitense Müll. Hal., 72
levieri Müll. Hal., 68 truncatulum Müll. Hal., 52, 53
Glossadelphus M. Fleisch., 52, 53, 79, 82
cocoensis (R. S. Williams) E. B. Bartram, 53 Irelandia W. R. Buck, 5, 16
laevifolius (Mitt.) E. B. Bartram, 79 robusticaulis (E. B. Bartram) W. R. Buck, 16, 17 (Fig. 5), 18
ligulaefolius E. B. Bartram, 79 (Fig. 6)
longisetus E. B. Bartram, 53 Isocladiella Dixon, 1
truncatulus (Müll. Hal.) M. Fleisch., 53 Isopterygiopsis Z. Iwats., 2, 4, 18, 19, 24
muelleriana (Schimp.) Z. Iwats., 19, 20 (Fig. 7), 21 (Fig. 8)
Herzogiella Broth., 2, 4, 9, 10, 11, 24 pulchella (Hedw.) Z. Iwats., 20, 21 (Fig. 8), 22 (Fig. 9), 23
boliviana (Broth.) M. Fleisch. in Broth., 9, 11 (Fig. 10)
cylindricarpa (Cardot) Z. Iwats., 11, 12 (Fig. 3), 13 (Fig. 4) Isopterygium Mitt., 1, 5, 10, 11, 16, 19, 23, 24, 25, 33, 36, 39,
Heterophyllium (Schimp.) Kindb., 1 41, 47, 51, 59, 67, 74
Homomallium (Schimp.) Loeske, 2, 5, 14, 57, 58 acutifolium Ireland, 25, 26 (Fig. 11), 27 (Fig. 12), 28
adnatum, 15 adflatum (Müll. Hal.) Paris, 44
mexicanum Cardot, 14, 15, 16, 58 afflictum (Müll. Hal.) Broth., 45
sharpii Ando & Higuchi, 14, 15 affusum Mitt., 24, 25, 27 (Fig. 12), 28 (Fig. 13), 29 (Fig. 14),
Hookeria retusa Wilson, 52 30 (Fig. 15)
Hookeriales, 79 alstonii E. B. Bartram, 51
Hookeriopsis cocoensis R. S. Williams, 53 altisetum H. A. Crum & Steere, 39
Hypnaceae Schimp., 1, 2, 3, 14, 24, 54, 68 ammophilum (Müll. Hal.) Broth., 45
subfam. Pylaisoideae (Schimp.) M. Fleisch., 3 angustirete (Broth.) Broth., 45
Hypnum, 1, 2, 4, 5 aquaticum H. Rob., 27
brachyneuron Müll. Hal., 43 araneosetum (Müll. Hal.) Paris, 45
brachystelium Hampe, 44 aurantiacum (Mull. Hal.) Broth., 45
brevisetum Hornsch., 44, 51 bahiense (Müll. Hal.) Broth., 45
cacti Müll. Hal., 44 brachyneuroides Broth., 45
chlorosum Hampe, 44 brachyneuron (Müll. Hal.) Mitt., 24, 43
chrismarii Müll. Hal., 51 brachystelium (Hampe) Kindb., 44
citrinum Hampe, 44 brasiliense (Broth.) W. R. Buck, 74
cordovense Müll. Hal., 44 brevisetum (Hornsch.) Broth., 51
curvicolle Müll. Hal., 43, 46 bromeliophilum (Müll. Hal.) Broth., 45
deplanatum Schimp. ex Müll. Hal., 75, 76 byssobolax (Müll. Hal.) Paris, 25, 27 (Fig. 12), 30, 31 (Fig. 16)
divaricatulum Müll. Hal. ex Hampe, 44 cacti (Müll. Hal.) Kindb., 44
elegans Hook., 62 callochlorum Broth., 45
 number 93 • 95

cavernicola Cardot, 82 poasense Renauld & Cardot, 59

chlorosum (Hampe) Paris, 44 pseudosubulatum (Müll. Hal.) Paris, 52
chrismarii (Müll. Hal.) Mitt., 24, 51 pseudotenerum (Broth. & Kindb.) Broth., 45
citrinum (Hampe) Kindb., 44 pulchellum (Hedw.) A. Jaeger, 20
clandestinum (Ångstr.) Broth., 44 pusillum Renauld & Cardot, 34
cordovense (Müll. Hal.) A. Jaeger, 44 pygmaeocarpum (Müll. Hal.) Broth., 52
curvicolle (Müll. Hal.) Mitt., 24, 43 pyrrhopum (Müll. Hal.) Paris, 30
curvicolle var. majus Broth. in E. Bauer, 43 radicisetum (Müll. Hal.) Broth., 44
cylindraceum (Müll. Hal.) Paris, 44 restingae (Müll. Hal.) Broth., 39
cylindricarpum Cardot, 11 restitutum (Hampe) Kindb., 44
deplanatum (Schimp. ex Müll. Hal.) Mitt., 76 richardsii E. B. Bartram, 64
diminutivum E. B. Bartram, 45 robusticaule E. B. Bartram, 16
distichaceum (Mitt.) A. Jaeger, 59 rufisetulum (Müll. Hal.) Paris, 45
divaricatulum (Hampe) Broth., 44 sapricola (Müll. Hal.) Broth., 52
elegans (Brid.) Lindb., 62 saprophilum (Müll. Hal.) Paris, 30
elegans var. nanum (Jur. in Milde) A. Jaeger, 62 semicostatum Renauld & Cardot, 51
eutrypherum (Müll. Hal.) Paris, 44 splendidulum (Hornsch.) Broth., 43
excelsipes (Müll. Hal.) Paris, 39 sprucei (Mitt.) W. R. Buck, 75
exiguum Kindb., 51 stigmocarpum (Müll. Hal.) Paris, 39
exile (Müll. Hal.) Paris, 51 streptopodium Besch., 44
fabroniiforme (Müll. Hal.) Broth., 45 subbrevisetum (Hampe) Broth., 25, 27 (Fig. 12), 29 (Fig. 14),
fecundum Renauld & Cardot, 40 30 (Fig. 15), 34, 35 (Fig. 19), 36
fernandezianum Broth. in Skottsb., 45 subcurvicolle (Müll. Hal.) Paris, 52
fernandezianum var. longisetum Broth. in Skottsb., 45 subfalcatum (Austin) A. Jaeger, 59
flaviusculum (Müll. Hal.) Broth., 45 subglobosum Herzog, 25, 27 (Fig. 12), 38 (Fig. 20), 39
fontigenum (Müll. Hal.) W. R. Buck & Ireland, 26 submersum (Broth.) Broth., 26
fruticola (Müll. Hal.) Paris, 51 subsplendidulum (Müll. Hal.) Paris, 52
fuegianum Besch., 21 subtenerrimum (Müll. Hal.) Paris, 34
fulvum (A. Jaeger) Kindb., 43 subtenerum Besch., 44
gracillimum (Hornsch.) Broth., 51 subtrichopelma Renauld & Cardot, 51
guarapense Besch., 44 taxirameum (Mitt.) A. Jaeger, 82
guatemalense E. B. Bartram, 16, 17 tenerifolium Mitt, 24, 25, 30 (Fig. 15), 32, 34 (Fig. 18), 39,
herminieri Schimp. ex Besch., 26 40 (Fig. 21), 41 (Fig. 22), 42 (Fig. 23)
homomallifolium Redf., 62 tenerrimum Renauld & Cardot, 34
hookeriophilum (Müll. Hal.) Broth., 52 tenerum (Sw.) Mitt., 23, 24, 25, 32, 34 (Fig. 18), 36, 41
hospitans (Hampe) A. Jaeger, 44 (Fig. 22), 42 (Fig. 23), 43, 46 (Fig. 24), 47 (Fig. 25)
irelandii H. Rob., 27 trichopelma (Müll. Hal.) Paris, 44
jamaicense (E. B. Bartram) W. R. Buck, 25, 30 (Fig. 15), 32, vagans Herzog, 40
33 (Fig. 17), 34 (Fig. 18), 41 (Fig. 22) weigeltii (Müll. Hal.) Broth., 45
lamprophyllum (A. Jaeger) Broth., 44 Isothecium
laxum (A. Jaeger) Broth., 44 elegans Brid., 58, 62
leptomiton (Müll. Hal.) A. Jaeger, 44 polyanthum (Hedw.) Spruce, 70
leptoplumosum (Dusén) Cardot, 21 subg. Pylaisia (Schimp.) Boulay, 68
leucophyllum (Hampe) Mitt., 24, 44 tenerum (Sw.) Brid., 43
lonchopelmatum (Müll. Hal.) Broth., 39
longisetum Broth., 39 Lepyrodon tomentosus (Hook.) Mitt., 51
longisetum (Schimp. ex Besch.) Broth., 39 Leskea
machrisianum (H. A. Crum) Ireland, 67 densa Hook. in Kunth, 55
manaosense Broth., 40 polyantha Hedw., 70
meteoriaceum (Müll. Hal.) Paris, 39 pulchella Hedw., 20
micans (Sw.) Kindb., 43
microplumosum (Müll. Hal.) Broth., 52 Mastopoma Cardot, 2
miradoricum (Müll. Hal.) A. Jaeger, 32, 52 Meteorium terrestre Müll. Hal., 6
mosenii Broth. in Engl. & Prantl, 39 Microthamniaceae Kindb., 3
muellerianum (Schimp.) A. Jaeger, 19 Microthamnium angustirete Broth., 45
paludigenum (Müll. Hal.) Broth., 39 Mittenothamnium Henn., 2, 5, 6, 51, 76, 86
peruvianum Broth., 45 scapellifolium (Müll. Hal.) H. A. Crum, 86
planissimum Mitt., 24, 82
planissimum var. laxirete Thér., 77 Orthotheciaceae Schimp., 3
96 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Papillaria rutenbergiacea Müll. Hal., 67, 68 Pterogonidium (Hook.) Müll. Hal. in Broth., 2, 33, 47
Phyllodon Bruch & Schimp. in Bruch, Schimp. & W. Gümbel, pulchellum (Hook.) Müll. Hal. in Broth., 33, 47
5, 52, 53 Pterogonium polyanthos (Hedw.) Muhl., 70
truncatulus (Müll. Hal.) W. R. Buck, 52, 53 Puiggariella Broth. in Engl. & Prantl, 67
truncatus (Welw. & Duby) W. R. Buck, 53 aurifolia (Mitt.) Broth. in Engl. & Prantl, 67, 68
Pilotrichum disciflorum Geh. & Hampe, 67 disciflora (Geh. & Hampe) Broth. in Engl. & Prantl, 67
Plagiotheciaceae, 2, 24 Puiggariopsis Menzel, 5, 67
Plagiothecium Bruch & Schimp. in Bruch, Schimp. & W. Güm- aurifolia (Mitt.) Menzel, 67
bel, 1, 24 Pylaisia Schimp. in Bruch, Schimp. & W. Gümbel, 2, 4, 14, 15,
aurantiacum Müll. Hal., 45 57, 58, 68, 69
bolivianum Broth. in Herzog, 11 falcata Schimp. in Bruch, Schimp. & W. Gümbel, 68, 69, 71
bromeliophilum Müll. Hal., 45 falcata var. intermedia Thér., 69
deplanatum (Schimp. ex Müll. Hal.) Spruce, 76 hamata (Mitt.) Cardot, 69
deplanatum f. ovatum Grout, 77 macrotis Cardot, 69
elegans (Brid.) Schimp., 62 panduraefolia Herzog, 69, 70
flaviusculum Müll. Hal., 45 polyantha (Hedw.) Schimp., 68
fontigenum Müll. Hal., 26 polyantha (Hedw.) Schimp. in Bruch, Schimp. & W. Gümbel,
fulvum Hook. & Wilson in Drumm., 43 69, 70
inordinatum Mitt. ex Spruce, 39 selwynii Kindb., 69, 71
jamaicense Müll. Hal., 39 subfalcata Schimp. in Bruch, Schimp. & W. Gümbel, 69
leptoplumosum Dusén, 21, 22 Pylaisiaceae Schimp., 2, 3
lonchopelmatum Müll. Hal., 39 Pylaisiadelpha Cardot, 1, 2
magellanicum (Besch.) Paris, 21 Pylaisiella Kindb. ex Grout, 68
meteoriaceum Müll. Hal., 39 falcata (Bruch, Schimp. & W. Gümbel) Ando, 69
micans (Sw.) Paris, 43 polyantha (Hedw.) Grout, 70
micans var. fulvum (A. Jaeger) Paris, 43 selwynii (Kindb.) H. A. Crum, Steere & L. E. Anderson, 71
muellerianum Schimp., 18, 19 Pylaisioideae (Schimp.) M. Fleisch., 1, 3
paludigenum Müll. Hal., 39
planissimum (Mitt.) E. B. Bartram, 82 Regmatodon fusco-luteus Schimp. ex Besch, 58
proximum Müll. Hal., 39 Rhacopilopsis Renauld & Cardot, 4, 71, 72
pseudotenerum Broth. & Kindb. ex Kindb., 45 dupuisii (Renauld & Cardot) Renauld & Cardot, 71
radicisetum Müll. Hal., 44 trinitensis (Müll. Hal.) E. Britton & Dixon in Dixon, 72
repens Lindb., 10 variegatum (Welw. & Duby) Watling & O’Shea, 72
restingae Müll. Hal., 39 Rhynchostegium micans (Sw.) Austin, 43
restingae var. tenue Müll. Hal., 39
subfalcatum Austin, 59 Sharpiella Z. Iwats., 10, 11, 24
villae-ricae Besch., 39 Stereodon Mitt., 2
Platygyriella Cardot, 5, 54, 55, 68 distichaceus Mitt., 59
densa (Hook. in Kunth) W. R. Buck, 55 falcatus (Schimp.) M. Fleisch. ex Broth. in Engl., 69
helicodontioides Cardot, 54, 55 falcatus var. intermedius (Thér.) Thér., 69
imbricatifolia (Cardot) Thér., 56 falcatus var. subfalcatus (Schimp.) Thér., 69
jamaicensis E. B. Bartram, 32 hamatus Mitt., 69
pringlei (Cardot) W. R. Buck, 55, 56 pandurifolius (Herzog) Broth. in Engl., 69
Platygyrium Schimp. in Bruch, Schimp. & W. Gümbel, 1, 2, 4, polyanthos (Hedw.) Mitt., 70
14, 57, 58, 68 sect. Ctenidium (Schimp.) Mitt., 5
fuscoluteum (Besch.) Cardot, 58 sect. Homomallium (Schimp.) C.E.O. Jensen, 14
fuscoluteum Cardot, 58 Stereodonteae Mitt., 24
guatemaliense Schimp. ex A. Jaeger, 54 subfalcatus (Schimp.) M. Fleisch. ex Broth. in Engl., 69
repens (Brid.) Schimp., 57 taxirameum Mitt., 82
Pseudohygrohypnum Kanda, 1, 2 Strigula Fr., 67
Pseudotaxiphyllum Z. Iwats., 2, 5, 19, 24, 58, 59, 67, 82 Strigulaceae, 67
distichaceum (Mitt.) Z. Iwats., 59, 60 (Fig. 26), 61 (Fig. 27) Symphyodon
elegans (Brid.) Z. Iwats., 63, 63 (Fig. 28), 64 (Fig. 29) americanus Steere, 67
homomallifolium (Redf.) Ireland, 61 (Fig. 27), 62, 65 imbricatifolius (Mitt.) S. P. Churchill, 67
(Fig. 30) machrisianus (H. A. Crum) W. R. Buck & Ireland, 67
machrisianum (H. A. Crum) Ireland, 67 Syringothecium Mitt., 5, 74
richardsii (E. B. Bartram) Ireland, 61 (Fig. 27), 64, 66 brasiliense Broth., 74
(Fig. 31) nemodontium Herzog, 75
Pseudotrismegistia H. Akiy. & Tsubota, 2 sprucei Mitt., 74, 75
 number 93 • 97

Taxicaulis Taxiphyllum M. Fleisch., 5, 16, 24, 53, 67, 75, 76, 79, 82, 86
adflatus Müll. Hal., 44 cavernicola (Cardot) Thér., 82
afflictus Müll. Hal., 45 cuspidifolium (Cardot) Z. Iwats., 16
ammophilus Müll. Hal., 45 deplanatum (Schimp. ex Müll. Hal.) M. Fleisch., 76, 77
araneosetus Müll. Hal., 45 (Fig. 32), 78 (Fig. 33)
bahiense Müll. Hal., 45 gallorum W. R. Buck, 80, 82
byssobolax Müll. Hal., 30 howellianum H. A. Crum & L. E. Anderson, 59
cylindraceus Müll. Hal., 44 laevifolium (Mitt.) W. R. Buck, 76, 78 (Fig. 33), 79, 89
excelsipes Müll. Hal., 39 (Fig. 34)
exilis Müll. Hal., 51 ligulaefolium (E. B. Bartram) W. R. Buck, 53, 76, 78
fabroniiformis Müll. Hal., 45 (Fig. 33), 79, 81 (Fig. 35)
fruticola Müll. Hal., 51 machrisianum H. A. Crum, 67
hookeriophilus Müll. Hal., 52 planissimum (Mitt.) Broth., 82
microplumosus Müll. Hal., 52 robusticaule (E. B. Bartram) Ireland, 16, 17
pseudosubulatus Müll. Hal., 52 scapellifolium (Müll. Hal.) Broth. in Engl., 86
pygmaeocarpus Müll. Hal., 52 taxirameum (Mitt.) M. Fleisch., 76, 82, 83 (Fig. 36), 84
pyrrhopus Müll. Hal., 30 (Fig. 37)
rufisetulus Müll. Hal., 45 Taxitheliella Dixon, 2
sapricola Müll. Hal., 52 Taxithelium Mitt., 1, 2
sapricola var. minor Müll. Hal., 52 Thedenia Schimp. in Bruch, Schimp. & W. Gümbel, 68
saprophilus Müll. Hal., 30 suecica Schimp. in Bruch, Schimp. & W. Gümbel, 68
stigmocarpus Müll. Hal., 39 Trichosteleum truncatulum (Müll. Hal.) A. Jaeger, 53
subcurvicollis Müll. Hal., 52 Trismegistia (Müll. Hal.) Müll. Hal., 2
subsplendidulus Müll. Hal., 52
subtenerrimus Müll. Hal., 34 Wijkia H.A. Crum, 2
trichopelma Müll. Hal., 44 alstonii (E. B. Bartram) Ireland, 51
weigeltii Müll. Hal., 45
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