SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 80 

A Taxonomic Revision of the 

Moss Families Hookeriaceae and 

Hypopterygiaceae in Malaya 

Haji Mohamed and Harold Robinson 

SMITHSONIAN INSTITUTION PRESS 

Washington, D.C. 

1991 
ABSTRACT 

Mohamed, Haji, and Harold Robinson. A Taxonomic Revision of the Moss Families 

Hookeriaceae and Hypopterygiaceae in Malaya. Smithsonian Contributions to Botany, number 

80,44 pages, 168 figures, 1 map, 1991.-Keys, descriptions, and some figures are provided for 

the 9 genera and 28 species of Hookeriaceae (including Chaetomitriopsis and Chaetomitrium) 

and the 3 genera and 7 species of Hypopterygiaceae recognised in Malaya. Eight species are 

reported as new to Malaya: Actinodontium rhaphidostegum, Daltonia angustijolia, D. armata, 

Distichophyllum brevicuspes, D. jungermannioides, D. maibarae, Cyathophorella hookeriana, 

and C. spinosa. Chaetomitrium perakense Brotherus ex Dixon and Distichophyllum ulukaliense 

Damanhuri & Mohamed are reduced to the synonymy of C. orthorrhynchum and D. 

jungermannioides respectively. 

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is 

recorded in the Institution’s annual report, Smithsonian Year. SERIES COVER DESIGN: Leaf 

clearing from the katsura tree Cercidiphyllum japonicum Siebold and Zuccarini. 

Library of Congress Cataloging-in-Publication Data 

Mohamed, Haji 

A taxonomic revision of the moss families Hookeriaceae and Hypopterygiaceae in Malaya / Haji Mohamed and Harold 

Robinson. 

p. 

Includes bibliographic references and index. 

1. Hookeriaceae-Malaysia-Malaya-Classification. 

2. Hypopterygiaceae-Malaysia-Malaya-Classification. 

I. Robinson, HaroldEmesk 1932- . 11. Title. 111. Series. 

QKlS2747 no. 80 [QK539.H64] 581 s-dc20 [588’.2] 91-4223 

cm. - (Smithsonian contributions to botany : no. 80) 

@ The paper used in this publication meets the minimum requirements of the American 

National Standard for Permanence of Paper for Printed Library Materials 239.48-1984. 
Contents 

Page 

Introduction ........................................ 1 

Abbreviations .................................... 1 

Acknowledgments ................................. 3 

Family HOOKERIACEAE ................................. 3 

Key to the Genera of Malayan Hookeriaceae ..................... 3 

Genus Actinodontium ................................. 4 

Key to the Malayan Species of Actinodontium ................... 4 

Actinodontium ascendens .............................. 4 

Actinodontium rhaphidostegum ........................... 4 

Genus Callicostella .................................. 4 

Key to the Malayan Species of Callicostella .................... 6 

Callicostella papillata ............................... 6 

Callicostella prabaktiana .............................. 6 

Excluded Species .................................. 6 

Genus Calyptrochaeta ................................. 9 

Calyptrochaeta remotifolia ............................. 9 

Genus Chaetomitriopsis ................................ 9 

Chaetomitriopsis glaucocarpa ........................... 9 

Genus Chaetomitrium ................................. 9 

Key to the Malayan Species of Chaetomitrium .................. 10 

Chaetomitrium borneense .............................. 10 

Chaetomitrium leptopoma ............................. 10 

Chaetomitrium orthorrhynchum .......................... 11 

Chaetomitrium papillifolium ............................ 11 

Chaetomitrium setosum ............................... 11 

Excluded Species .................................. 11 

Genus Cyclodictyon .................................. 12 

Cyclodictyon blumeanum .............................. 12 

Genus Daltonia .................................... 12 

Key to the Malayan Species of Daltonia ...................... 12 

Daltonia angustifolia ................................ 13 

Daltonia armata .................................. 13 

Daltonia contorta .................................. 13 

Excluded Species .................................. 13 

Genus Distichophyllidium ............................... 17 

Distichophyllidium nymanianum .......................... 17 

Genus Distichophyllum ................................ 17 

Distichophyllum brevicuspes ............................ 19 

Distichophyllum cirratum .............................. 19 

Distichophyllum cuspidaturn ............................ 19 

Distichophyllum jungermannioides ......................... 22 

Distichophyllum maibarae ............................. 22 

Distichophyllum malayense ............................. 25 

Distichophyllum mittenii .............................. 25 

Distichophyllum nigricaule ............................. 25 

Key to the Malayan Species of Distichophyllum .................. 17 

... 

111 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Distichophyllum osterwaldii ............................ 26 

Distichophyllum schmidtii ............................. 26 

Distichophyllum spathulatum ............................ 29 

Distichophyllum tortile ............................... 29 

Excluded Species .................................. 29 

Family HYPOPTERYGIACEAE ............................... 29 

Key to the Genera of Malayan Hypopterygiaceae .................. 32 

Genus Cyathophorella ................................. 32 

Key to the Malayan Species of Cyathophorella .................. 32 

Cyathophorella burkillii .............................. 32 

Cyathophorella hookeriana ............................. 32 

Cyathophorella spinosa ............................... 35 

Excluded Species .................................. 35 

Genus Hypopterygium ................................. 35 

Key to the Malayan Species of Hypopterygium .................. 35 

Hypopterygium aristatum .............................. 37 

Hypopterygium tenellum .............................. 37 

Genus Lopidium .................................... 37 

Key to the Malayan Species of Lopidium ..................... 37 

Lopidium struthiopteris ............................... 40 

Lopidium trichocladon ............................... 40 

Literature Cited ...................................... 43 

Index ........................................... 44 
A Taxonomic Revision of the 

Moss Families Hookeriaceae and 

Hypopterygiaceae in Malaya 

Haji Mohamed and Harold Robinson 

Introduction 

The present study revises taxonomically the species of the 

families Hookeriaceae and Hypopterygiaceae known in Malaya 

(Map 1). The circumscription of these two families has been a 

matter of controversy (Miller, 1971; Crosby, 1974; Buck, 1987; 

Robinson, 1971, 1986, and Tan & Robinson, 1990). In this 

study, we have followed the concept of Fleischer (1908) and 

Bartram (1939) in circumscribing the two families. However, 

we believe that Chaetomitriopsis and Chaetomitrium, though 

treated here, in the Hookeriaceae, do not really belong to that 

family but occupy a position closer to the Hypnaceae. Not 

included here is Sclerohypnum riparium Dixon which has been 

known only from the type specimen from Malaya. It has been 

reexamined by Harrington and Miller (1979). They have 

suggested that if peristome structure is used as the determining 

factor, it should provisionally be assigned to the Hookeriaceae. 

Although we have seen no material of the species, the scanning 

electron micrographs of the exostome shown by Harrington 

and Miller do not show the wide median furrow similar to that 

which is seen on many of the Hookeriaceae. Although a region 

of depression is present in the middle, it is quite different from 

the wide furrow usually found in the Hookeriaceae. In addition, 

the presence of somewhat inflated, subquadrate alar cells on the 

leaves suggest that it would be more appropriate to place 

Sclerohypnum in the Hypnaceae. 

A total of 28 species in 9 genera (including 1 species in 

Chaetomitriopsis and five in Chaetomitrium) are recognized in 

the Hookeriaceae while 7 species in 3 genera are recognized in 

the Hypopterygiaceae. Eight species: Actinodontium 

rhaphidostegum, Daltonia angustifolia, D. armata, Distichophyllum brevicuspes, D. jungermannioides, D. maibarae, 

Cyathophorella hookeriana, and C. spinosa are recorded as 

Haji Mohamed, Botany Department, University of Malaya, 59100, 

Kuala Lumpur, Malaysia. Harold Robinson, Department of Botany, National Museum of Natural History, Smithsonian Institution, 

Washington, D.C. 20560. 

new to Malaya. Chaetomitrium perakense Brotherus ex Dixon 

and Distichophyllum ulukaliense Damanhuri & Mohamed are 

reduced to synonymy of C. orthorrhynchum and D. jungermannioides respectively. Previous claims of Callicostella beccariana, Chaetomitrium ciliatum, C. elongatum, C. lanceolatum, C. 

muricatum, C. nematosum, C. torquescens, Cyathophorella 

tenera, Daltonia aristifolia, and Distichophyllum undulatum in 

Malaya have been found to be based on misidentified 

specimens. Distichophyllum malayense is the only endemic 

species of the Hookeriaceae in Malaya. 

The specimens cited in the text are limited to types studied 

and representative specimens which show the distribution of 

the species within Malaya. All the specimens cited are from the 

Herbarium of the Botany Department, University Malaya, 

Kuala Lumpur, Malaysia (KLW, unless denoted otherwise. 

ABBREVIATIONS.-The following abbreviations are used to 

indicate the herbaria in which the specimens studied are 

located. 

B Botanisches Museum, Berlin-Dahlern, Germany 

BM British Museum (Natural History) London, 

England 

BO Herbarium Bogoriense, Bogor, Indonesia 

FH Farlow Herbarium at Harvard University, 

Cambridge, Massachusetts (includes the Bartram and Fleischer Herbaria) 

Conservatoire de Jardin botaniques, Geneva, 

Switzerland 

Botanical Museum, University of Helsinki, 

Finland 

Helsinki: Brotherus Herbarium (part of the 

Botanical Museum, University of Helsinki, 

Finland) 

New York Botanical Garden, Bronx, New 

York 

G 

H 

H-BR 

L Rijksherbarium, Leiden, Netherlands 

NY 

1 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Scale 1: 3 125 000 

0 150 km 

AILAND International boundary 

----- State boundary 

UMATRA 

INDONESIA ) 

MAP l.-Map of peninsular Malaysia (Malaya) showing the divisions of states and three well collected 

mountains. 
NUMBER 80 3 

PC 

SING Singapore Botanic Gardens, Singapore 

UKMB Herbarium of the Botany Department, 

Universiti Kebangsaan Malaysia, Bangi, Malaysia 

Botanical Museum and Herbarium, Utrecht, 

Netherlands 

United States National Herbarium, Smithsonian Institution, Washington D.C. 

Ranges given for the species in text are mostly from the 

literature. Literature on the mosses of Malaya used in the study 

in addition to that cited in text includes Damanhuri and 

Mohamed (1986), Dixon (1924, 1931), Johnson (1980), 

Mohamed (1985), and Mohamed and Mohamed (1986). 

Further references used covering other areas include Bartram 

(1939), Dozy and Molkenboer (1845-1854), Gangulee (1977), 

and Townsend (1982). 

ACKN0WLEDGMExTS.-The first author would like to 

acknowledge the research fellowship grant awarded by the 

Smithsonian Institution which enabled him to carry out 

research at the National Herbarium (US) at Washington D.C. 

Together, we thank Halimatul Sadiah Abdullah (JSLU), Ahmad 

Damanhuri Mohamed (UKMB), B.C. Tan and D. Pfister 0, 

P. Isoviita 0, A. Harrington (BM), H. Bischler (PC), W. Buck 

Laboratoire de Cryptogamie, Museum d’Histoire Naturelle, Paris, France 

U 

us 

my), M. Shah (SING), P. Tixier, and C.C. Townsend for the 

loan of specimens. We thank A. Damanhuri Mohamed for 

drawing the figures of Cyathophorella. We also thank Sherry 

Pittam, R.M. King, and other members of the Botany 

Department of the Smithsonian Institution who helped in one 

way or another in the completion of the research and the 

preparation of the manuscript. Special thanks are offerred to 

B.C. Tan and Bruce Allen for their detailed reviews of the 

manuscript. 

Family HOOKERIACEAE 

HCOKERIACEAE Schimper, Corollarium Bryologiae Europeae, 101, 1855. 

Plants small to robust, frequently complanate; primary stems 

usually branched and prostrate; secondary stems prostrate, 

suberect, or pendent; leaves variable in shape, with or without 

border, the dorsal leaves erect-spreading to spreading; costa 

single, double, or absent; cells smooth or papillose; alar cells 

not differentiated. Synoicous, autoicous, or dioicous. Calyptra 

generally mitriform, smooth, scabrous, or pilose, the base 

usually lobed or fringed-ciliate. Seta short or elongate, smooth, 

papillose, or ciliate; capsule inclined or horizontal, occasionally erect; peristome double, exostome teeth papillose or striate, 

often with a wide median furrow; cilia lacking, rarely 

rudimentary. 

Key to the Genera of Malayan Hookeriaceae 

1. Costa single. .................................... 2 

Costa double or none. .............................. .3 

2. Stems not complanate-foliate; leaves isomorphous, oblong-lanceolate to lanceolatelinear; exostome without a median furrow, papillose .......... Daltonia 

Stems complanate-foliate; leaves dimorphous, ovate-oblong to spathulate; exostome 

deeply furrowed, suiolate .................... Distichophyllum 

Costae strong, reaching at least midleaf. ..................... .4 

Costae weak, short or none. ........................... .6 

3. 

4. Leaves distinctly bordered; leaf cells lax, more than 30 pm wide .......... .................................... Cyclodictyon 

Leaves not or only weakly bordered; leaf cells firm, less than 30 pm wide .... 5 

5. Exostome outer surface papillose, without median furrow; stems ascending; leaf cells 

more than twice as long as wide; costae not protruding abaxially ......... ................................... Actinodontium 

Exostome outer surface striolate, with median furrow; stems prostrate; leaf cells 

mostly isodiametric; costae protruding abaxially .......... Callicostella 

6. Leaves bordered; leaf cells smooth ......................... 7 

7. Leaves large, more than 2 mm long; margin distinctly toothed above; seta densely 

spinuliferous ............................ Calyptrochaeta 

Leaves not bordered; leaf cells often prorulose ................... 8 

Leaves small, less than 2 mm long; margin entire; seta smooth ........... ................................. Distichophyllidium 

Leaves strongly divergent or deflexed; operculum conic ..... Chaetomitriopsis 

Leaves imbricate to erect-spreading; operculum rostrate ...... Chaetomitrium 

8. 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Genus Actinodontium midleaf; cells elongate, smooth. Heteroicous. Calyptra naked, 

Actinodontium Schwaegrichen, Spec. Musc. Suppl., 2:75, 1826. [Type: A. laciniate at base. Capsule erect, subcylindric; operculum 

ascendens Schwaegrichen.] rostrate; exostome teeth papillose with zig-zag median line. 

Plants small, green, growing in tufts; stems ascending, The plants of this genus are mainly epiphytic, in Malaya 

radiculose below, simple, densely radially foliate; leaves often occumng on tea plants. The leaves are bicostate as in 

ovate-lanceolate, acuminate; costae double, extending beyond Callicostella but the costae do not protrude abaxially. 

Key to the Malayan Species of Actinodontium 

Leaves ovate to oblong-elliptic; apices acute to short-acuminate; capsules up to 2.5 mm 

long .................................... A. ascendens 

.................................... A. rhaphidostegum 

Leaves oblong-lanceolate; apices long-acuminate; capsules up to 3 mm long ....... 

Actinodontium ascendens 

FIGURES 5-8 

Acfinodontiwn ascendens Schwaegrichen, Spec. Musc. Suppl., 2(2):75, 1826. 

[Type: Java, sine loco, Reinwardt s.n. (G?), not seen.] 

Plants to 1 cm high, in tufts: stems simple, rarely branched: 

leaves crowded, erect and slightly shrunken or crisped when 

dry, erect-spreading when moist, ovate to oblong-elliptic, 

2.1-2.8 mm long, 0.6-0.9 mm wide, widest at midleaf or 

below, apices acute to short-acuminate: margin entire above, 

plane or slightly recurved below; border scarcely differentiated, 

consisting of one row of weakly differentiated cells near apex; 

costae unequal, extending to 3/4 of leaf; median cells 

thin-walled, rhomboid to elongate-hexagonal, 60- 120 pn 

long, 15-20 prn wide; basal cells 90-150 pm long, 12-17 ym 

wide, elongate-hexagonal to somewhat linear; basal cells at 

attachment quadrate to short-rectangular. Calyptra often stuck 

on operculum even after capsule matures, mitrate, laciniate and 

whitish below, smooth and brownish above. Seta smooth, to 7 

mm long; capsule including operculum 2.5 mm long, suberect; 

operculum rostrate. 

SELECTED SPECIMENS EXAMINED.-Pahang: Cameron 

Highlands, Ringlet, Boh Tea plantation estate, on twigs of tea 

plants, 1540 m, Abdullah & Zamzuri 1127; Cameron Highlands, Tanah Rata, Parit Waterfalls, 1100 m, Tixier 5219 (PC). 

WORLI) DIS’~iUf3UTION.-Sri Lanka, Thailand, Java, and 

Malaya. 

FERTILITY .-Both packets examined have sporophytes. 

REMARKS.-Actinodontium ascendens is distinguished from 

A. rhaphidostegum on the basis of leaf shape, leaf size, and 

capsule length. Actinodontium ascendens has elliptic to ovate 

leaves with acute to short-acuminate apices and capsules rarely 

exceeding 2.5 mm in length, while A. rhaphidostegum has 

oblong-lanceolate leaves with long-acuminate apices and 

capsules measuring up to 3 mm long. The leaves of A. 

ascendens are smaller, measuring 2.1-2.8 mm long 0.6-0.9 

mm wide, while A. rhaphidostegum has leaves measuring 

3.5-4.5 mm long, 0.8-1.2 mm wide. Despite the differences, 

there are specimens in which the characters intergrade. In 

Malaya, these two species are found in the same habitats and 

localities. 

Actinodontium rhaphidostegum 

FIGURES 1-4 

Actinodonlium rhaphidostegum (C. Muller) Bosch & Lacoste, Bryol. Jav., 

Hookeria rhaphidostega C. Muller, Syn., 2:677, 1857. [Type: Java, Blwne s.n 

2:37, 1862. 

(B, destroyed).] 

Plants to 1.5 cm high; leaves crowded, slightly shrunken and 

flexuose when dry, erect-spreading when moist, oblonglanceolate, gradually long-acuminate, 3.5-4.5 mm long, 

0.8-1.2 mm wide; margins entire above, involute for 2/3 of 

length below; costae unequal, parallel, extending beyond 

midleaf; leaf cells smooth, thin-walled, elongate-rhomboid to 

elongate-hexagonal, 80-130 pm long, 16-20 pm wide. 

Calyptra mitrate, laciniate at base, covering only top of capsule. 

Seta smooth, erect, to 6 mm long; capsule erect or slightly 

inclined, ovate-cylindrical with a short apophysis, to 3 mm 

long (including operculum); operculum rostrate. 

SELECTED SPECIMENS EXAMINED.-Pahang: Cameron 

Highlands, Tanah Rata, MARDI Research Station, 1440 m, on 

tea plants, H. Mohamed & Zamzuri 1052a; Baki Bakar 258. 

WORLD DIsTruBun0N.-Malaya, Java, Celebes, Borneo, 

and Philippines. 

FERTILITY.-Both collections had sporophytes. 

REMARKS.-The differences between this species and A. 

ascendens are discussed under the latter. 

Genus Callicostella 

Callicosrella (C. Muller) Mitten, nom. cons., J. Linn. Soc. Bot. Suppl., 1:136, 

1859. [Type: C. papillata (Montagne) Mitten.] Crosby (1975) showed that 

CaNicoslella was a later synonym for Schizornitrium Bruch, Schimper, and 

Giimbel. However, Koponen and Isoviita (1984) proposed that the name 

Callicostella which had been used for a long time, be. conserved. The 

proposal was approved at the 1987 Berlin Congress. 
NUMBER 80 5 

1 

A 1-3,5-7 

_____I 

1.0 mm 

FIGURES l-S.-Actinodontium. 1-4, A. rhaphidostegum (Pahang: H. Mohamed & Zamzwi IOMa): 1-3, leaves; 

4, cells at leaf apex. 5-8, A. ascendens (Pahang: Abdullah & Zumzwi 1127): 5-7, leaves; 8, cells at leaf apex. 
6 SMITHSONTAN CONTRIBUTIONS TO BOTANY 

Plants slender, yellowish to dark green, in mats; stems 

creeping, irregularly branched, laxly complanate-foliate; leaves 

oblong, short-acute to short-acuminate, not bordered, closely 

serrate above; costae double, thick, reaching beyond midleaf, 

cnding in abrupt apical protrusion; cells firm, hexagonal, or 

rhomboidal, nearly isodiametric, smooth or with central 

papilla. Synoicous, autoicous, or dioicous. Calyptra naked, 

scabrous distally, weakly plicate and laciniate at base. Seta 

elongate, often partly or completely roughened; capsule 

inclined to horizontal, subcylindric; operculum rostrate; exostome teeth outer surface striolate, with median furrow. 

This genus is characterised by the presence of two distinct 

leaf costae which protrude abaxially just below the apex. It 

resembles Cycfodicryon in habit and in having double costae, 

but the leaves are not bordered and the leaf cells are 

firm-walled. 

Key to the Malayan Species of Callicostella 

Leaf cells papillose ............................... C. papillata 

Leaf cells smooth .............................. C. prabaktiana 

Callicostella papillata 

FIGURES 9-15 

Callicostella pupillafa (Montagne) Mitten, J. Linn. SOC. Bot. Suppl., 1:136, 

1859. 

above 1000 m, the highest at 1560 m. It is a polymorphic 

species; the only reliable specific character is its papillose cells. 

Callicostella prabaktiana 

Hookeria prabakfianu C. Miiller. Syn., 2:678, 1851. [Type: Java, Mt. Prabakti, 

Zollinger 3503 (B, destroyed).] 

Plants in flat mats, yellow to dark green; stems creeping, 

prostrate, irregularly branched; leaves complanate, somewhat 

curved and plicate when dry, oblong-lingulate, 1-1.8 mm long 

0.55-0.65 mm wide, broadly acute to abruptly acuminate; 

margins not bordered above, a single row of elongate cells 

below, sermlate to dentate in upper half, often with bigeminate 

teeth, lower half entire; costae ceasing just below apex in 

abaxial protrusion, smooth below, toothed above; median cells 

irregularly 6-7 sided, hexagonal to oblong, 6-14 pm long, 

6-9 pm wide, thick-walled, with a single papilla over the 

lumen, basal cells elongate and smooth, 30-50 pm long, 8-10 

pm wide; perichaetial leaves lanceolate, smaller than vegetative leaves, margin serrate. Synoicous. Seta smooth below, 

weakly papillose above, 0.8-1.5 cm long; capsules ovoidcylindric. 

SELECTED SPECIMENS EXAMINED.-Johore: Gunung Panti, 

H. Mohamed 9369. Pahang: Fraser Hill, H. Mohamed & 

Zamzuri 945a; Cameron Highlands, Robinson Falls, Manuel 

3328. Perak: Gunung Hijau Trail, Manuel 3240 & 3267a. 

Selangor: 3 mile mark on old Genting Highlands road, Manuel 

28066. 

WORLD DISTRIBUTION.-Madagascar, South India, Andaman Islands, Sri Lanka, Thailand, Vietnam, Malaya, Sumatra, Java, Borneo, Philippines, Taiwan, Ryukyus, New Guinea, 

New Caledonia, Fiji, Tahiti, and Samoa. 

FERTILITY.-Twelve of the 16 packets examined had 

sporophytes. 

REMARKS.-Callicostella uauillata has been collected on 

This species is similar in all aspects to C. papillata except 

that the leaf cells are smooth and the apical margins are less 

dentate. 

SELECTED SPECIMENS EXAMINED.-Johore: Mersing, Clear 

1394, 1449; Foothill of Gunung Ledang, A. Mohamed 125. 

Kelantan: Jeram Paw, Aishah 124. Negeri Sembilan: Perhentian Tinggi, Ridley 571 (SING). Pahang: Taman Negara, Lata 

Berkoh area, on Sungei Tahan, Manuel 2551; Cameron 

Highlands, Robinson's Waterfall, H. Mohamed & Zamzuri 

1123~. Perak: on rock near waterfall, H. Mohamed et al. 8355. 

Penang: Botanic Gardens, H. Mohamed 8414. Selangor: Ulu 

Gombak, Zarina Mat 4, 10, & 31. 

WORLD DISTRIBUTION.-Vietnam, Thailand, Malaya, Java, 

Borneo, Philippines, and New Guinea. 

FERTILITY.-Fourteen of the 25 specimens examined had 

sporophytes. 

REMARKS.-This species has been collected on rocks, logs, 

and soil on banks of streams and rivers in lowland forests in the 

shade. It is sometimes found on rocks which are occasionally 

inundated. Of the 25 specimens examined, 23 were collected 

between 50 m and 400 m. Specimen H. Mohamed & Zamzuri 

1123a and specimen H. Mohamed & Damanhuri 1054 were 

collected at 1460 m and 1580 m respectively. 

Excluded Species .. logs and rocks in and near streams in mountain forests in partly 

shaded areas. This is essentially a medium to high altitude 

species unlike C. prabaktiana which has been collected mostly 

below 400 m. Fourteen of the packets examined were collected 

Callicostella beccariana (Hampe) Jaeger was reported from 

Malaya by Mohamed and Tan (1988) based on Burkill 6566 

collected from Selangor, Telok Reserve (SING). This specimen 

is in fact C. prabaktiana. 

Callicostella prabaktiana (C. Muller) Bosch & Lacoste, Bryol. Jav., 2:40,

1862.
NUMBER 80 7 

0.05 mm 

15 

FIGURES 9-15.4aNicosiellapapillata (Johore: H. Mohamed 9369): 9, plant, wet; 10.11. leaves; 12, cells at leaf 

apex; 13, cells at base of leaf; 14, cells at margin near midleaf; 15, median cells of leaf. 
8 SMITHSO" CONTRIBUTIONS TO BOTANY 

FIGURES 16-22.-Calyptrochaeta remtifolia (Pahang: H. Mohamed 7089): 16, 17, dorsal leaves; 18, 19 lateral 

leaves; 20, ells at upper margin of leaf; 21, cells at leaf apex; 22, median cells of leaf. 
NUMBER 80 9 

Genus Calyptrochaeta 

Calyptrochaeta Desvaux, MCm. Soc. EM. Pans, 3:226, 1825. [Type: Leskea 

crirt~t~ Hedwig.] 

Plants medium-sized to robust, brownish-green, in tufts; 

stems wiry, prostrate or ascending, mostly simple, sometimes 

forked, flat, laxly cornplanate-foliate, usually with clusters of 

brown, filiform propagules on stems among upper leaves; 

lateral leaves spreading, dorsal and ventral rows smaller and 

erect, ovate, short-acuminate, bordered, serrate above; costa 

short and forked; cells lax, hexagonal or rhomboidal, smooth. 

Autoicous or dioicous. Calyptra naked or pilose, fringed. Seta 

strongly spinuliferous; capsule small, pendulous; exostome 

teeth transversely striolate with a median furrow. 

Eriopus, an earlier generic name in common use, has been 

found to be illegitimate (Margadant 1959; Crosby 1974), and 

the name Calyptrochaeta must be used. A large number of 

collections from Malaya have been examined, and since no 

consistent differences in characters have been found, only one 

species is recognized here. Sporophytes are produced infrequently in the genus and thus sporophyte characters are not 

useful in keys to the species. 

Calyptrochaeta remotifolia 

FIGURES 16-22 

Calyptrocheta remotifolia (C. Muller) Iwatsuki, Tan, & Touw, 5. Hattori Bot. 

Eriupw remotifoliw C. Muller, Bot. Zeit., 5:828. 1847. [Type: Java, Mt Gede,

Lab., 44:150, 1978. 

Reinwardt s.n. (B, destroyed).] 

Plants large, pale green to dark green, secondary stems 

ascending, to 5 cm high, simple or branched, blunt at tips, 

complanate-foliate, to 8 mm wide with leaves: numerous 

clusters of brown, filiform gemmae present near leaf bases; 

lateral leaves oblong-ovate, short-acuminate, to 5 mm long, 2 

mm wide; margins coarsely serrate in upper half with long 

slender spinose teeth; distinctly bordered all around with 3-4 

rows of elongate cells, with strongly pitted walls; costa short, 

forked, weakly defincd; median cells rhomboidal with firm 

walls, 50-75 pm long, 20-25 pm wide, basal cells laxer; 

perichaetial leaves with round to shortly acute apices. Dioicous. 

Calyptra hairy, laciniate at base. Seta to 6 mm long, 

spinuliferous; capsule erect or inclined, ovoid. 

SELECTED SPECIMENS EXAMINED.-Pahang: Gunung Ulu 

Kali, corticolous, 1500 m, H. Mohamed 7089; Cameron 

Highlands, Hutan Simpan Huh Bertam, 1620 m, H. Mohamed 

& Zamzuri 111.5~. Perak: Maxwell Hill, 1330 m, H. Mohamed 

8079. 

WORLD DIsTRIBUn0N.-India (Manipur), Malaya, Sumatra, 

Java, Borneo, Philippines, and New Guinea. 

REMARKS.-This species has been collected between 1200 

m and 2000 m. Two different types of habitats seem to be 

favored by the Malayan plants of this species. One habitat is the 

base of shrubs and trees in relatively dry areas while the other 

habitat is on partially immersed rocks in swift-flowing streams. 

Those plants growing on bases of shrubs are mostly light-green 

in color and more strongly toothed in the upper margins while 

those such as H. Mohamed & Zamzuri 1115c, growing on 

partially immersed rocks, are dark to dark green in color and 

have upper margins that are not strongly toothed. 

Genus Chaetomitriopsis 

Chaetornitriopsir Fleischer, Musd Fl. Buitenz, 4:1371, 1923. [Lectotype: C. 

glaucocarpa (Schwaegrichen) Fleischer] 

Plants slender, wiry, yellowish-green glossy, in compact, 

intricate tufts; stems creeping, densely pinnate to bipinnate; 

branch leaves squarrose-spreading with decurved points, 

ovate-acuminate; border not differentiated; costae double, 

short; cells elongate, prorulose at apical angles. Calyptra 

cucullate, sparingly pilose. Seta long, smooth; capsule pendulous; exostome teeth striolate, without median furrow; operculum conical, short. 

Chaetomitriopsis glaucocarpa 

Chaetomitriopsir glaucocarpa (Schwaegrichen) Fleischer, Musd Fl. Buitenz., 

4:1372, 1923. 

Hypnwn glaucocarpon Reinwardt ex Schwaegrichen, Spec. Musc. Suppl., 

3(1):228a, 1828. [Type: In Insulae Javae monte Mulawar: Pr. Reinwardt 

(G?), not seen.] 

Plants yellowish-green, in dense tufts or mats; stems more 

than 10 cm long, with pinnate to bipinnate secondary branches; 

ultimate branches erect with densely inserted squarrose leaves 

on all sides; leaves cordate-ovate, to 0.8 mm long, 0.4 mm 

wide, short-acuminate; margins serrate to base; costa short, 

unequal; leaf cells irregularly rhomboid, 20-30 pm long, 4-6 

pm wide, prorulose. Autoicous. Seta 2.5 cm long; capsule 

ovoid-cylindric, 0.6 mm long. 

SELECTED SPECIMENS EXAMINED.-Pahang: Cameron 

Highlands, Brinchang Rose Garden, on stem of tree fern, 1540 

m, H. Mohamed 9182. 

WORLD DIsTRIBUn0N.-Nepal, Sikkim, Assam, Laos, 

Vietnam, Malaya, Java, Celebes, New Guinea, Philippines, and 

Taiwan. 

REMARKS.-Chaetomitriopsis glaucocarpa has been collected on branches and twigs of trees at altitudes above 1400 m. 

This species is easily recognized by the densely branched stems 

and the crowded, squarrose, deflexed, cordate-ovate branch 

leaves. 

Genus Chaetomitrium 

Chetomitrium Dozy & Molkenboer, Musci Frond. Ined. Archip. Indid, 2:117, 

1847. [Lectotype: C. elongatwn (Dozy & Molkenboer) Dozy & Molkenboer.) 

Plants small to medium sized; stems long, creeping, usually 

regularly pinnate, sometimes with clusters of filamentous 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

gemmae among leaves on ultimate branches; leaves erectspreading, widely spreading or appressed, ovate, concave, 

acuminate to rounded; margins usually dentate, without 

differentiated border; costae short and double or none; cells 

linear, shorter near apex, generally prorulose; perichaetial 

leaves larger than vegetative leaves. Autoicous or dioicous. 

Calyptra mitrate or cucullate, always strongly hispid or 

spinose-ciliate, usually fringed at the base. Seta elongate, 

papillose or setose; capsule more or less inclined; operculum 

rostrate: exostome teeth striolate, not furrowed. 

Key to the Malayan Species of Chaetomitrium 

1. Branch leaves arranged in spirally seriate rows .................. .2 

Branch leaves evenly spaced, not in spirally seriate rows ............. 3 

2. Upper leaf margins sermlate; setae papillose above .......... C. borneense 

Upper leaf margins spinose-serrate; setae setulose above ........ C. setosum 

3. Branch leaves homomallous; leaf margins only weakly toothed above ....... ................................... C. papillifolium 

Branch leaves not homomallous; leaf margins strongly toothed above ...... .4 

4. Leaves less than 0.7 mm long, leaf apices not constricted below tips; setae papillose 

above, ................................ C. leptoporna 

Leaves more than 1 mm long, leaf apices constricted below apices; setae setulose 

above ....................................... 5 

5. Leaves to 1.2 mm long, ovate-lanceolate, margins serrate; papillae on sera to 50 pm 

high .............................. C. orthorrhynchum 

Leaves up to 1.7 mm long, ovate, upper margin spinose-serrate; papillae on seta up 

to 90 pm high ............................. C. setosum 

Chaetomitrium borneense foliation is indistinct. It is possible that C. elongatum from Java 

may be synonymous with this species. A study of the type 

specimens of both species shows no appreciable differences in 

the sporophytic and gametophytic characters except for the 

indistinct spirally seriate foliation and the more acuminate leaf 

apices in C. elongaturn. The nomenclatural problems surrounding this species have been dealt with by Mohamed (1985). 

Chefomitrim borneense Mitten, Rep. Sci. Res. Voyage Challenger Bot., 

Chuetomitrim seriatum Brotherus, Bot. Jahrb., 49:350, 1913. [Isotype: 

1:213, 1885. [Type: Borneo, Motley 4 (NY!).] 

Southern Borneo, Hayup. H. Winkler 2522 (L!).] 

1889, L. WruyJr. 3550 & 3636 (BM!).] 

Pilotrichella perukensis Brotherus in Dixon, J. EM. SOC., Bot., 43:314, 1916. 

nom. nud. in synon. [Based on: Malaya, Perak, Upper Perak, 300 ft, June 

Plants golden green; primary stems creeping, up to 16 cm 

long; branches erect, to 1.5 cm long; branch leaves spirally 

seriate, ovate-oblong, cucullate above, abruptly acute to 

rounded with a short apiculus, constricted below apex, 0.8-1.3 

mm long, -0.55 mm wide, widest below midleaf, concave, 

plicate below: margins serrulate above: costae short and weak; 

cells linear, weakly prorulose; perichaetial leaves longacuminate, with spinose-serrate marginal teeth. Dioicous. 

Calyptra mitriform, hirsute with branched hairs, fringed below 

with long hairs. Seta up to 1 cm, papillose above, the papillae 

up to 25 pm high, smooth below; capsule cylindrical, 

operculum rostrate. 

SELECTED SPECIMENS EXAMINED.-Pahang: Temerloh, epiphytic on tree, Poore B1128. Trengganu: Dungun, Kampong 

Pasir Raja, Sungei Pertang, 60 m, on tree, H. Mohamed 9523. 

WORLD DISTRIBUTION.-Malaya and Borneo. 

REMARKS.-This species is easily recognized by the spirally 

seriate arrangement of the leaves on the stem similar to the 

arrangement in some South American species of Pilotrichella 

and Orthostichopsis. However, there are collections such as H. 

Mohamed 9523 and A. Damanhuri 2143 in which the seriate 

Chaetomitrium leptopoma 

Chetomitrium leptopoma (Schwaegrichen) Bosch & Lacoste, Blyol. Jav., 

Hypnwn leptopoma Schwaegrichen, Spec. Musc. Suppl., 4:323b, 1842. [Type: 

252. 1862. 

Java, Reinwardt & Blm (G), not seen, (FH!).] 

Plants small, densely branched in compact tufts; leaves 

erect-spreading, ovate with an acute apex, 0.3-0.7 mm long, 

0.08-0.3 mm wide, widest just below midleaf, concave; 

margins serrulate in upper 2/3 of leaf length; costae short and 

weak: median cells linear, distinctly prorulose; perichaetial 

leaves often plicate, to 1.8 mm long, 0.35 mm wide, 

oblong-lanceolate with a long flexuose apex, spinose-serrate on 

margins in upper 2/3. Dioicous. Calyptra mitriform, spinosehispid on surface, fringed with short hairs. Seta to 1 cm high, 

papillose above, smooth below: operculum rostrate. 

SELECTED SPECIMENS EXAMINED.-Perak: Birch’s Hill, 

4200 ft, on a branch, Burkiff (BM). Selangor: Ginting Bidai, 

1896, Ridley 405 (SING). 

WORLD DISTRIBUm0N.-Java, Malaya, Borneo, and Philippines. 
NUMBER 80 

REMARKS.-This species is the smallest of all the Chaetom’- 

trium species in Malaya. It is recognized by its dense 

branching, small ovate leaves with serrulate margins and 

papillose seta. 

11 

Chaetomitrium orthorrhynchum 

Chaetomitrium orlhorrhynchum (Dozy & Molkenboer) Bosch & Lacoste, 

Hookeria orfhorrhynchwn Dozy & Molkenboer, AM. Sci. Nat. Bot., ser. 3, 

Chaelomilriwn perakense Brotherus ex Dixon, Bull. Torrey Bot. Club, 51:237, 

Bryol. Jav., 2:45, 1862. 

2:305, 1844. [Type: Java, Zippe1iu.v (L), not seen, (FH!).] 

1924. [Type: Perak, Bidor, Tapah, HN. RidZey 159 (BM!).] syn. nov. 

Plants small, in golden-green mats; stems pinnate to 

bipinnate; leaves ovate-lanceolate, acuminate, - 1.2 mm long, 

to 0.4 mm wide, widest below midleaf; ultimate branch leaves 

smaller, concave, constricted below the apex; margin sharply 

serrate in upper 2/3 of leaf, entire below; costae short, faint; cells 

linear, prorulose, especially on the back of leaf; perichaetial 

leaves ciliate-dentate. Dioicous. Calyptra mitriform, spinosehispid above, fringed with long hairs. Seta 6-10 mm long, 

setulose above, with setulae to 50 pm long, smooth below; 

capsule cylindrical; operculum rostrate. 

SELECTED SPECIMENS EXAMINED.-Johore: Sungei Pelepah, Md. Nur 19138 (SING). Pahang: Taman Negara, on leaf, 

Damanhuri s.n. (UKMB). Perak: Bidor, Tapah, Ridley 164 

(SING). Selangor: Gua Batu, Ridley 474 (SING). Trengganu: 

Ulu Trengganu District, Sungei Berua, off Tapah Road, H. 

Mohamed 9653. 

WORLD DIsTRIBunoN.-Malaya, Indonesia, and Philippines. 

REMARKS.-This species in Malaya is known only from 

below 150 m on rotten logs, twigs, and trees in moist places. 

The Malayan specimens have larger leaves and more pronounced serrations on the margin when compared to the Javan 

type specimen. 

Chaetomitrium papillifolium 

Chaeiomitrium papilfijofium Bosch & Lacoste, Bryol. Jav.. 2:50, 1862. [Type: 

Java, Von Teysmann, ex Herb. Sande Lac. (FH!).] 

Primary stems creeping; branches to 0.8 cm long, homomallous; branch leaves usually ovate-lanceolate to oblonglanceolate, acute to acuminate, slightly constricted below apex, 

1.0-1.4 mm long, 0.4-0.5 mm wide, widest near midleaf, 

margin weakly toothed to denticulate above, minutely toothed 

below; costa short; cells linear, prorulose at apical angles, 

shorter at apex; perichaetial leaves ciliate-dentate. Dioicous. 

Calyptra mitriform, with a few scattered, appressed long hairs 

on surface, fringed with long ciliate hairs below. Seta 0.6-1.4 

cm long, smooth below, papillose above, the papillae up to 15 

pm high; capsule cylindrical, 1.4 mm long, operculum rostrate. 

SELECTED SPECIMENS EXAMINED.-Johore: Mersing, on 

slender twig 9” above ground, Clear 1452. Penang: Western 

Hill, 2500 ft, wet logs, Spare 3181 (BM). Perak: Dindings, 

Ridley 373 (BM). Selangor: Ulu Gombak forest Reserve, on 

twig 6’ from ground in forest, Burkill 9964 (SING). Pahang: 

Janda Baik, 400 m, on tree, H. Mohamed 8883. Trengganu: 

Dungun district, Kampong Pasir Raja, 60 m, on tree along 

Sungei Pertang, H. Mohamed 9660. 

WORLD DISTRIBUTION.-Sri Lanka, Indochina, Malaya, 

Java, Borneo, and Philippines. 

REMARKS.-Chaetomitrium papillifolium is distinguished 

by the homomallous arrangement of branch leaves, ovatelanceolate to oblong-lanceolate leaf shape, weakly toothed leaf 

margins and a seta which is weakly papillose above. 

Chaetomirrium orthorrhynchum is distinct from the present 

species in the more coarsely serrate leaves and the setulose seta. 

Chaetomitrium setosum 

Chaetomitrium setoswn Brotherus ex Dixon, Bull. Torrey Bot. Club, 52:237, 

1924. [Type: Perak, Tapah, Nov. 1908, Ridley 169 (BM!).] 

Primary stems creeping, secondary stems simple or 

branched, to 1.5 cm high; leaves arranged in more or less 

seriate rows; somewhat tumid, erect-spreading, ovate with an 

acuminate apex, concave, often cucullate and constricted near 

the tip, 1.3-1.7 mm long, 0.6-0.7 mm wide, widest near 

midleaf; upper margins spinose-serrate, with teeth less prominent below; costa weak, short; leaf cells linear, prorulose; basal 

region of attachment to stem yellowish; perichaetial leaves to 2 

mm long, to 0.55 mm wide, long-acuminate, teeth on upper 

margin large, spinose. Autoicous. Caly-ptra mitriform, densely 

hirsute, fringed with long hairs. Seta to 1 cm long, setulose 

above, with setulae to 90 pm long, papillose or smooth below; 

capsule cylindrical; operculum rostrate. 

SELECTED SPECIMENS EXAMINED.-Kedah: Choong Meng, 

on stem of shrub, 100 ft, spare 2804 (BM). Sarawak: Gunung 

Dulit, 300 m, twigs and leaves of small tree in rain forest near 

torrent, Oxford Expedition 1932, No. 1144 (BM). 

WORLD DISTRIBUTION.-Malaya and Borneo. 

REMARKS.-This species is distinct in having a combination 

of leaves constricted below the apices, leaf margins with 

spinose-serrate teeth, and setae setulose. The leaves are larger, 

more ovate, the margins more sharply serrate and the setae 

much more setulose than those of C. orthorrhynchum. 

Excluded Species 

Chaetomitrium ciliatum Bosch & Lacoste was reported from 

Malaya by Tixier (1980) based on Tixier 5234 collected in 

Pahang, Cameron Highlands, Parit Waterfalls, on twigs in 

forest, Dec. 25, 1970 (PC). This has proven to be C. 

orthorrhynchum. 

Chaetomitrium elongatum Dozy & Molkenboer was reported from Malaya by Dixon (1926). The report was based on 

two collections, Ridley 373 from Perak, Dindings, 1897 (BM, 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

SING) and Burkill 16569 from Pahang, Sungei Perting, 

Bentong (BM). In both specimens the leaves are much smaller 

and the apices are much more acute than in C. elongatum. The 

specimens are C. papillifolium. 

Chaetomitrium lanceolatum Bosch & Lacoste was reported 

by Tixier (1980) based on Tixier 5234. The specimen has the 

same number, same locality, and same station as that given for 

C. ciliatum above, and it also proves to be C. orthorrhynchum. 

Chaetomitrium muricatum Bosch & Lacoste was reported by 

Dixon (1926) based on Ridley 405 and 411 from Selangor, 

Ginting Bidai, 1896 (BM). These specimens are C. leptopoma. 

Chaetomitrium nematosum Brotherus in Dixon was reported 

by Dixon (1926) based on Ridley 205 from Perak, Kuala 

Kenering (BM). Despite the peculiar brood-filaments and the 

color of the plant, it is certainly C. orthorrhynchum. 

Chaetomitrium torquescens Bosch & Lacoste was reported 

by Tixier (1971) based on Tixier 4809 collected from Pahang, 

Fraser's Hill, Dec. 27, 1969 (PC). The specimen is in fact C. 

papillifolium. 

Genus Cyclodictyon 

Cyclodictyon Mitten, J. Linn. Soc., Bot., 7:163, 1864. [Type: C. laefevirens 

(Hooker & Taylor) Mitten.] 

Plants medium-sized, soft, delicate, in mats; stems prostrate, 

branched, complanate-foliate; leaves very laxly areolate, 

oblong-ovate, acute, narrowly bordered, weakly toothed above; 

costa double, extending beyond midleaf; cells large, roundhexagonal, smooth. Synoicous, autoicous or dioicous. Calyptra 

naked. Seta elongate, smooth; capsule horizontal; exostome 

transversely striolate with a wide median furrow; operculum 

rostrate. 

Cyclodictyon blumeanum 

Cyclodicfyon blwneanum (C. Muller) 0. Kuntze, Revis. Gen. Pl., 2:835,1891. 

Hookeriana blumeana C. Muller, Syn., 2:676. 1851. [Type: Java, Blwne s.n. 

(B, destroyed).] 

Plants, light-green, irregularly branched, branches 2 cm 

long, to 2 mm wide with leaves; leaves weakly complanate, 

slightly distorted when dry, ovate to oblong-lingulate, 1.2-1.8 

mm long, 0.6-0.9 mm wide, broadly acute to shortly 

acuminate, the apiculus 50-80 pm long; margins weakly 

toothed above near apex, plane below; border distinctly 

differentiated, with two rows of linear cells; costa reaching 4/5 

the length of leaf, with several dorsal spines towards apex; cells 

large, 33-50 pm wide, irregularly hexagonal above, becoming 

rectangular towards base. Synoicous or autoicous. Calyptra 

mitrate, covering operculum and capsule mouth when mature, 

smooth, yellowish, laciniate at base. Seta up to 1.6 cm high, 

reddish brown, smooth; capsule dark brown, suberect to 

cemuous, ovoid-cylindrical; urn 1.5 mm; mouth enlarged, 

mamillose. 

SELECTED SPECIMENS EXAMINED.-Pahang: Cameron 

Highlands, Hutan Simpan Hulu Bertam, 1620 m, H. Mohamed 

& Zamzuri 1207b; Genting Highlands, near water tank, - 1500 

m, H. Mohamed 4558. 

WORLD DISTRIBUTION.-S~~ Lanka, Sumatra, Java, Bali, 

Malaya, Borneo, Philippines, Taiwan, New Guinea, and Fiji. 

FERmrrY.-Three out of the 5 packets examined had 

sporophytes. 

REMARKS.-Cyclodictyon blumeanum is recognized by its 

bicostate leaves, and the lax areolation formed by its large leaf 

cells. It may be confused locally only with the genus 

Callicostella which is also bicostate. However, the smaller firm 

leaf cells which may or may not be papillose and the 

unbordered leaf apices of Callicostella separate it from 

Cyclodictyon blumeanum. This species has been collected on 

wood and rocks near moist, shady places such as waterfalls and 

streams. 

Genus Daltonia 

Dalfonia Hooker & Taylor, Musci Brit., 80, 1818, nom. cons. [Type: D. 

splachnoides (J.E. Smith) Hooker & Taylor.] The nomenclatural history of 

Daltonia leading to cmservation is given by Crosby (1968). 

Plants small to medium-sized, often tufted, more or less 

glossy; stems laxly erect, often simple, rarely forked, densely 

radially foliate; leaves ovate or lanceolate, acuminate, bordered, carinate along costa below; costae single, ending some 

distance below apex; upper cells oval-rhomboidal, linear at 

margins in a distinct border. Synoicous, autoicous, or dioicous. 

Calyptra deeply fringed at the base. Seta scabrous above; 

capsule suberect; exostome teeth papillose, without median 

furrow. 

The plants of this genus in Malaya are generally epiphytic, 

growing in small tufts on bark of trees and bases of small 

shrubs. 

Key to the Malayan Species of Daltonia 

1. Leaves more than 3 mm long, ending in a long arista; costa ending below middle or 

distal third of leaf. ........................... D. armata 

Leaves less than 3 mm long, acuminate; costa reaching 4/s of leaf length ..... 2 

2. Leaves strongly twisted or contorted when dry, oblong-lanceolate; margins plane, the 

apices abruptly short acuminate .................... D. contorta 

Leaves flexuose when dry, narrowly lanceolate to linear; margins recurved above, 

apices gradually acuminate .................... D. angustifolia 
NUMBER 80 13 

Daltonia angustifolia 

FIGURES 23-29 

Daltonia angurtifolia Dozy & Molkenbcer, Ann. Sci. Nat. Bot., ser. 3,2:302, 

1844. [Type: Borneo, sine loco, Korthals s.n. (L), not seen.] 

Plants small, yellowish-green to green, forming cushions; 

stems simple or rarely branched, 5-8 mm high; leaves 

crowded, erect and flexuose when dry, erect-spreading when 

moist, 2.0-2.6 mm long, 0.3-0.4 mm wide, widest below 

midleaf, narrowly lanceolate to linear, apex gradually acuminate; margins entire, strongly recurved above but often only on 

one side below; border well differentiated, with 1-2 rows of 

linear cells above, to six rows of linear cells below; costa 

reaching 4/5 of leaf length; cells oval to narrowly rhomboidal, 

20-33 pm long, 5-8 pm wide, with walls slightly thickened; 

basal cells rectangular, lax, 25-42 pm long, 6-12 pm wide; 

perichaetial leaves shorter and narrower than vegetative leaves. 

Synoicous. Calyptra fringed with long hairs at base, covering 

only operculum. Seta 5-8 mm long, scabrous in upper half, 

smooth below; capsules ovoid, erect to suberect, 1 mm long; 

exostome teeth reflexed when wet. 

SELECTED SPECIMENS EXAMINED.-Pahang: Cameron 

Highlands, Gunung Brinchang, 1900 m, H. Mohamed 9139; 

Genting Highlands, 1690 m, on moist wood, H. Mohamed 

7662; Gunung Ulu Kali, 1770 m, on rotten log, A. Damanhuri 

3809. 

WORLD DISTRIBUTION.-Central Africa, Madagascar, Sri 

Lanka, Nepal, Vietnam, Malaya, Philippines, Java, Borneo, 

New Guinea, and New Zealand. 

REMARKS.-Daltonia angustifolia occurs in extensive mats 

on moist trunks and branches of trees at altitudes from 

1900-2000 m. This is the most common and smallest species 

of the genus in Malaya. It is distinct in having linear-lanceolate 

leaves and distinctly recurved leaf margins. Daltonia contorta 

has ovate-lanceolate leaves which are strongly contorted when 

dry whereas leaves of D. angustifolia are hardly altered when 

by. 

Daltonia armata 

FIGURFS 37-42 

Daltonia armta Banram, Farlowia, 1508, 1944. [Type: Philippines, Mindanao, Sacred Mountain, Lanao, Zwickey 638 (FH!).] 

Plants moderately robust, yellowish green, in lax cushions; 

stems rarely branched, to 1.5 cm high; leaves crowded, erect to 

erect-spreading, flexuous, 3.0-3.8 mm long, 0.7-0.9 mm 

wide, widest at or above midleaf, broadly ovate-oblong to 

obovate, acuminate with long spirally contorted arista 200-300 

pm long; margins entire, with 1(-2) rows of distinct elongate 

cells, often involute below; costa weak, less than l/2 to 2/3 leaf 

length, hidden in strong basal carina; median leaf cells 50-84 

pm long, 15-20 p-n wide, narrowly hexagonal to oval, with 

thin walls; basal cells irregularly hexagonal, 80-120 pm long, 

18-22 pm wide, orange; propagulae numerous, fusiform, 

140-170 pm long, of 6-8 cells; perichaetial leaves smaller, 

ovate-lanceolate, long-acuminate, ecostate. Dioicous. Seta 5-7 

mm long, strongly scabrous above; capsules 1 mm long. 

SELECTED SPECIMEN EXAMINED.-Pahang: Cameron Highlands, Strawberry Park Road, on base of tree, 1500 m, H. 

Mohamed 9307 (KLU, US). 

WORLD DIsTRIBUn0N.-Malaya, Sumatra, and Philippines. 

FERTILITY.-The Malayan specimen had no sporophytes. 

REMARKS.-Daltonia armuta is distinct in possessing large 

ovate-oblong to obovate leaves which are particularly strongly 

carinate, a weakly defined costa often not reaching to midleaf, 

a weak border of 1(-2) rows of cells and a long arista. The 

weak costa, which is unusual in Daltonia, is hidden in the 

carina of the leaf. Philippine material of the species has been 

described as having a stronger costa reaching 2/3 the leaf length 

(Tan & Robinson 1990). 

Daltonia contorta 

FIGURES 30-36 

Daltonia contorta C. Muller, Syn., 2:660, 1851. [Type: Java, sine loco, Blwne 

s.n. (B, destroyed).] 

Medium-sized yellow to brownish plants in dense tufts; 

stems to 1.5 cm high, rarely branched; leaves crowded, erect 

and strongly twisted and contorted when dry, erect-spreading 

when moist, ovate-lanceolate, 2.2-2.7 mm long, 0.4-0.6 mm 

wide, widest below midleaf, abruptly short-acuminate; margins 

entire and plane; border well differentiated, with 1-2 rows of 

cells above, to 7 rows of linear cells below; costa strong, 

extending 4/s of leaf length; median cells oval-rhomboidal with 

firm pale walls, rather opaque, 9-15 pm long, 7-9 pm wide; 

cells at base rectangular, 20-40 pm long, 8-12 p wide. 

Calyptra deeply fringed. Seta 6-8 mm long, scabrous in upper 

half only, smooth below; capsule erect to suberect, ovoid, 1.2 

mm long. 

SELECTED SPECIMENS EXAMINED.-Pahang: Cameron 

Highlands, Gunung Brinchang, 1890 m, H. Mohamed 9146; 

Genting Highlands, 1600 m, Sadiah & Zumzuri 1590 & 1563. 

WORLD DISTRIBUTION.-^^^ Lanka, Malaya, Java, Philippines, Fiji, and Hawaii. 

FERTILITY.-A~~ 4 packets examined had sporophytes. 

REMARKS.-This species has been collected only on the 

bases of shrubs at altitudes between 1400 m and 1900 m. The 

combination of distinctly twisted and contorted ovatelanceolate leaves, plane leaf margins, wide borders below, and 

short-acuminate non-aristate apex will differentiate this species 

from other Malayan members of the genus. 

Excluded Species 

Daltonia aristifolia Renault & Cardot was reported from 

Malaya by Tixier (1980) based on Tixier 5220 collected from 

Pahang, Cameron Highlands, Parit Waterfalls, on twigs, 1400 

m, Oct. 26, 1970 PC). The specimen has been found to be 

Distichophyllum cuspidatum. 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

23 

FIGURES 23-29.-Daltonia angurtifolia (Pahang: H. Mohamed 9139): 23, plant, wet; 24, capsule, wet; 25, 26, 

leaves; 27, cells at leaf apex; 28, cells at margin near midleaf, 29, cells at base of leaf. 
0 

€0 

E 

cv, 

0 m 
16 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGLKES 37-42.-Daltonia armfa (Pahang: H. Mohamed 90 9037): 37, plant, wet; 38, 39, leaves showing 

carma, costa not shown; 40, cells at margin near midleaf; 41. cells at leaf apex; 42, cells at base of leaf. 
NUMBER 80 17 

Genus Distichophyllidium 

Distichophyllidium Fleischer, Musci Fl. Buitenz., 3:966, 1908. 

Plants small, stems branched, densely foliate; leaves erect to 

erect-spreading, ovate, acuminate with long flexuose arista; 

margin entire above, recurved below: leaf border narrow; 

ecostate: median cells hexagonal, homogeneous. Autoicous. 

Calyptra smooth, laciniate at base. Seta smooth; capsules erect 

to suberect, ovoid; peristome double, exostome teeth papillose, 

without median furrow, segments not perforated; operculum 

rostrate. 

Distichophyllidium nymanianum 

Distichophyllidium nymanianum Fleischer, Musci Fl. Buitenz., 3:967, 1908. 

[Type: West-Java vor Tjibodas am Gedeh, 1200 m, Nymn s.n. (FH), not 

seen.] 

Plants small, in light green mats, to 1 cm; stems often 

branched; leaves erect when dry, erect-spreading when wet, 

slightly crispate when dry, ovate, 1.1-1.4 mm long, 0.4-0.5 

mm wide, widest at midleaf or just below, concave, base 

distinctly carinate forming a basal pocket, apex acute to obtuse, 

ending in a long flexuose arista, arista to 0.5 mm long; leaf 

border well differentiated, with 1 row of linear cells from base 

to apex; margin entire above, recurved below; median cells 

hexagonal, 15-22 pm wide, thin-walled, more or less 

homogeneous, cells near base elongate-hexagonal. Calyptra 

smooth, fringed at base. Seta smooth, 3-4.5 mm high: capsules 

erect to suberect, ovoid; urn expanded at mouth, 0.4 mm long. 

SELECTED SPECIMENS EXAMINED.-Pahang: Cameron 

Highlands, 1500 m, H. Mohamed & AD.Mohamad 1118. 

WORLD DISTRIBUTION.-Malaya, Java, and New Guinea. 

FERTILITY.-AU 3 packets examined had sporophytes. 

Rmms.-Distichophyllidium nymanianum grows in large 

mats on the bark of trees in shaded moist areas between 1500 m 

and 1750 m. The absence of a costa, the presence of a long 

flexuose arista, a strongly carinate leaf base and a papillose 

peristome will differentiate this species from species of 

Distichophyllum. 

Genus Distichophyllum 

Distichophyllum Dozy & Molkenboer, Musci Frond. Ined. Archip. Indici, 4:99, 

1846. 

Plants small to robust; stems sparingly branched; leaves 

complanate, weakly dimorphous, dorsal and ventral rows 

usually weakly differentiated, crowded, ovate or spathulate, 

usually entire, usually with a border of elongated cells: costa 

single, extending from midleaf to leaf apex; upper cells 

isodiametric, smooth, more lax and elongated at base. Calyptra 

mitrifonn, fringed at base. Seta smooth or papillose; capsule 

erect to pendulous; peristome double; exostome teeth transversely striolate, with a wide median furrow; endostome with 

high basal membrane, segments sometimes perforated; operculum conic-rostrate. 

Distichophyllum is the genus of Hookeriaceae with the most 

species in Malaya. The members of this genus generally grow 

on moist, shady substrates such as rocks, soil, humus, and 

rotten logs at high elevations. Only Distichophyllum schmidtii 

has been collected at low elevations. They also occasionally 

occur as epiphytes. 

1. 

2. 

3. 

4. 

5. 

6. 

7. 

Key to Malayan Species of Distichophyllum 

I. With Sporophytes 

(excepting D. mluyense) 

Calyptra with long hairs on the surface. ............... D. maibarae 

Calyptra scabrous or smooth but without long hairs on surface, long hairs only in 

basal fringe ................................... .2 

Seta scabrous throughout or in part. ....................... 3 

Seta smooth. ................................... .8 

Seta scabrous throughout ............................. .4 

Seta scabrous only near capsule, smooth below .......... D. brevicuspes 

Leaf apices not bordered by elongated cells ............. D. osterwaldii 

Leaf apices bordered by at least one row of elongated cells ............ 5 

Submarginal cells of leaf distinctly smaller than paracostal cells; leaf border narrow, 

consisting of 1-2 rows of elongated cells .................... 6 

Leaf cells more or less homogeneous: leaf border wider, consisting of 2-3(-4) rows 

of elongated cells ................................. 7 

Plants dioicous ........................... D. spathulatum 

Plants autoicous ............................. D. mittenii 

Median leaf cells 14-16(-20) pm wide; leaf apiculus shorter, 30-50(-60) pm long 

..................................... D. schmidtii 

Median leaf cells 40-55 pm wide: leaf apiculus 50-100 pn long ... D. tortile 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

8. Leaf contracted to a long cuspidate apiculus more than 200 pm long ....... ................................... D. cuspidatum 

Leaf apiculus less than 100 pm long ....................... .9 

................................ D. jungermannioides 

9. Leaves mostly less than 1.5 mm long; costa reaching midleaf, rarely longer .... 

Leaves mostly longer than 1.5 mm; costa reaching 2/3 of leaf length ...... 10 

10. Leaves distinctly crispate and contorted when dry; submarginal cells in upper half 

of leaf much smaller than juxtacostal cells .............. D. cirratum 

Leaves scarcely crispate and contorted when dry; cells in upper half of leaf 

homogeneous ............................ D. nigricaule 

1. 

2. 

6. 

7. 

8. 

9. 

10. 

11. 

12. 

Key to the Malayan Species of Distichophyllum 

11. Based on Gametophytic Material 

Leaf apices with marginal cells quadrate to hexagonal ....... D. osterwaldii 

Leaf apices bordered by elongated cells ...................... 2 

Leaves contracted to a long cuspidate point, the leaf acumen more than 200 pm . ........................................... 3 

Leaf apices round or obtuse, muticous or shortly mucronate, the acumen if present, 

less than 100 pm long. ............................ .4 

Costa less than l/2 of leaf length ................... D. malayense 

Costa ending just below apex .................... D. cuspidatum 

Leaves more than 3 mm long. ......................... .5 

Leaves 3 mm long or shorter ............................ 8 

Submarginal cells of leaf distinctly smaller than paracostal cells; leaf border narrow, 

consisting of 1-2 rows of cells; leaf apiculus less than 20 pm long ...... 6 

Submarginal cells and paracostal cells more or less similar in size; leaf border 

wider, consisting of 2-3(-4) rows of elongated cells .............. 7 

Plants dioicous ........................... D. spathulaturn 

Plants autoicous ............................. D. mittenii 

Median leaf cells 40-55 pm wide; leaf apiculus 50-100 pm long . . , D. tortile 

Median leaf cells 14-16(-20) pm wide: leaf apiculus 30-50(-60) pm long ... ..................................... D. schmidtii 

Median leaf cells less than 14 pm wide ................ D. maibarae 

Median leaf cells more than 14 pm wide ..................... 9 

Leaves strongly crispate and contorted when dry; submarginal cells much smaller 

than paracostal cells .......................... D. cirratum 

Leaves hardly contorted and crisped when dry; cells in upper half of leaf more or 

less homogeneous, ............................... 10 

Leaves often carinate along costa near base; leaf cells collenchymatous; leaf apices 

acute to obtuse. .......................... D. brevicuspes 

Leaves not carinate along costa; leaf cells not collenchymatous; leaf apices obtuse 

to round ..................................... 11 

Plants with stems less than 8 mm high; costa reaching midleaf, rarely longer ... ................................ D. jungermannioides 

.......................................... 12 

Leaves ovate to broadly elliptic; median cells 16-23 pm wide ........... .................................... D. nigricaule 

Leaves oblong-ovate to obovate; median cells 14-16(-20) pm wide ....... ..................................... D. schmidtii 

Plants with stems to 2.0-3.5 cm long; costa reaching at least 2/3 of leaf length . . 
NUMBER 80 19 

Distichophyllum brevicuspes 

Dislichophyllum brevicuspes Fleischer, Musci EL. Buitenz., 3:979, 1908. 

[Type: Java, am Poentjak Pass, FIeischer s.n. (kctotype, FH!).] 

Plants small to medium sized, mat forming: stems to 1.4 cm 

long, 2.5 mm wide with leaves, often simple, sometimes 

branched; leaves yellowish green to green, slightly crisped, 

ovate to oblong-elliptic, rarely lingulate, often carinate along 

costa near the base, 1.3-2.3 mm long, 0.7-10 mm wide, acute 

to obtuse, cuspidate, the apiculus (25-)50-82 pm long, not 

twisted: margin entire above, weakly revolute below; border 

well differentiated, with 1-2 rows of linear cells near apex, 

2(-3) rows below; median cells round to hexagonal, collenchymatous, 13-17 pm wide, more or less homogeneous in upper 

half of leaf, cells at base hexagonal to rectangular, 40-90 pm 

long, 18-25 pm wide. Dioicous. Calyptra dark brown and 

smooth above, yellowish and fringed below. Seta to 6 mm high, 

scabrous just below neck, smooth below; capsule ovoid 

cylindrical, 1.5 mm long, mamillose; operculum rostrate. 

SELECTED SPECIMENS EXAMINED.-Pahang: Cameron 

Highlands, Gunung Brinchang, alt. 1890 m on moist Cyathea 

and wood in partly shaded area, H. Mohamed 9071 & 9091. 

WORLD DIsTRIBun0N.-Java, Philippines, and Malaya. 

FERTILITY,-B~~~ specimens examined had sporophytes. 

REMARKS.-Distichophyhn brevicuspes is a new record 

for Malaya. Fleischer (1908) compared it to D. montagneanum 

sensu Fleischer (= D. maibarae). Although it resembles D. 

maibarae in areolation and leaf apex, it differs in a number of 

characters. Distichophyllum brevicuspes has a carinate leaf 

base, median leaf cells 13-17 pm wide, setae scabrous just 

below the capsule and smooth calyptrae, while D. maibarae has 

a plane leaf base, median cells 9-15 pm wide, smooth setae 

and hairy calyptrae. Distichophyllum brevicuspes may also be 

confused with D. schmidtii. It differs in having smaller leaves, 

shorter leaf apiculi, and setae which are Scabrous only just 

below the neck. In D. schmidtii the leaves are larger, the apiculi 

longer, and the setae are scabrous throughout. In addition, D. 

brevicuspes has been collected only above 1000 m, while D. 

schmidtii has only been collected below 800 m. 

Distichophyllum cirratum 

FIGURES 50,52,53,56 

Distichophyllum cirratum Renauld & Cardot, Rev. Bryol., 23:104, 1896. 

[Type: Java, Tjibodas, Massart 1397 (isotype, FH!).] 

Dktichophyllum nigricaule var. cirratum (Renauld & Cardot) Fleischer, Musci 

Fl Buitenz., 3:982, 1908. 

Durrchophylfurn elmerr Brotherus, hallets Philipp. BOL, 2:656, 1909. [Type: 

Luzon, Baguio, Benguet Prov., Efmer 10453 (holotype, H-BR!).] 

Distichophyflum sinuosulwn Dixon, J. Siam SOC., Nat. Hist., Suppl. 10 (1):lS. 

1935. [Type: iMalaysia, Perak, Birch's Hill, Burkill 12606 (holotype, BM!).] 

Dishhophyllum nigricaule var. elmeri (Brothem) Tan & Robinson, Smithsonian Contributions to Botany, 75:20, 1990. syn. nov. 

Small plants in lax tufts, dull yellowish green; stems 

prostrate, 6-10 mm high, up to 3 mm wide with leaves, dark 

brown or black, laxly foliate, usually simple; leaves strongly 

crisped and contorted when dry, 1.8-2.5 mm long, 1.0-1.4 mm 

wide, widest at midleaf or slightly above, ovate to obovate, 

base often narrowed, apically rounded and apiculate, the 

apiculus (10-)30-50(-75) pn long; ventral and dorsal leaves 

smaller; margin entire, somewhat undulate; leaf border well 

differentiated with 2-3 rows of linear cells near the apex, 4-6 

rows of linear cells near the base: costa length variable, 3/5 to 4/5 

of leaf length, sometimes percurrent; paracostal cells irregularly hexagonal, 22-35 pm wide, gradually decreasing in size 

towards margins, collenchymatous; marginal cells hexagonal 

to round, 12-16 pm wide, cell walls slightly thickened, weakly 

collenchymatous; basal cells hexagonal to short rectangular, 

50-75 pm long, 15-25 pm wide. Dioicous. Calyptra scabrous 

above, fringed with long cilia. Seta 7-8 mm long, smooth: 

capsule horizontal to inclined; urn 1 mm long, mamillose; 

operculum rostrate. 

SELECTED SPECIMENS EXAMINED.-Perak: Maxwell Hill, 

1180 m, H. Mohamed et al. 8107. Pahang: Cameron Highlands, 

1560 m, H. Mohamed 9132 & 9172; Tanah Rata, 1440 m, H. 

Mohamed & Zamzuri 1059a. 

WORLD DISTRIBUTION.-Nepal, Thailand, Ryukyu Islands, 

Taiwan, Malaya, Sumatra, Java, Borneo, Philippines, and New 

Guinea. 

FERnLm.-Of the 16 packets examined, 10 had sporophytes. 

REMARKS-Distichophyllum cirratum has been collected 

between the altitudes of 1180 m and 1880 m on the mainland 

and at 800 m on the island of Tioman. Thirteen of the packets 

were collected on moist soil while three packets were collected 

on moist rocks. 

Distichophyllum cirratum is characterised by dark stems, 

strongly crisped and contorted leaves when dry, wide borders, 

'and submarginal cells which are smaller than paracostal cells. 

In addition, the median cells of the Malayan specimens are 

collenchymatous. Differences between this species and D. 

nigricaule are discussed under the latter. 

Distichophyllum cuspidatum 

FIGURB 43-49 

Distichophyllum curpidatum (Dozy & Molkenboer) Dozy & Molkenboer, 

Musci Frond. Ined. Archip. Indici, 4:101, 1846. 

Hookeria curpidata Dozy & Molkenboer, Ann. Sci. Nat. Bot., ser. 3, 2:305. 

1844. [Types: Java, prope Limbang, Zippeliur s.n., Korthals s.n. (L), not 

seen; Sumatra, Batang-Bessie, Korfhals s.n. (L), not seen.] 

Plants dark green above, yellow below, prostrate, in tufts; 

stems to 2 cm long: leaves Compressed on the stem in a 

dorsi-ventral arrangement, 3-4.6 mm long, 0.9-1.3 mm wide, 

elliptic-oblong to oblanceolate, abruptly contracted to a long 

cuspidate point to 650 pm in length: margins entire, slightly 
20 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

48 

43 

5.0 mm 

44,45 

1.0 mm 

46-49 

0.1 mm 

FIGURFS 43-49.4irtichophyllwn curpidatum (Pahang: M.G. Manuel 2729): 43. plant, wet; 44,45, leaves; 46, 

cells at leaf apex; 47, median cells of leaf, 48, cells at margin near midleaf; 49, cells at base of leaf. 
FIGURES 50-57.-Dktichophyllumm; 50,52,53,56, D. clrratum (Pahang: H. Mohamed 9132): 50, plant, dry; 52, 

53, leaves; 56, cells at leaf apex; 51,54,55,57, D. nigricaule (Pahang: H. Mohamed & Zomzuri 922%): 51, plant, 

dry; 54.55, leaves; 57, cells at leaf apex. 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

revolute at leaf base: borders well developed, with 2-3 rows of 

thick walled elongated cells; costa extending to near apex; 

median cells round, (6-)9-15(-17) pm, thick walled, collenchymatous; basal cells lax, rhomboidal, 40-60 pm long, 

18-24 pm wide. Synoicous and autoicous. Calyptra naked, 

fringed at base. Seta smooth, up to 5 mm long: capsule erect to 

inclined, 1 mm long. 

Kali, 5000 ft., Manuel 2729, 3417; Cameron Highlands, 

Gunung Chantek, Sadiah and Zumzuri 1114. Perak: Maxwell 

Hill, trail to Microwave Station, H. Mohamed et al. 8085. 

Penang: Moniot Road, on branches near ground, Burkill 2584 

(SING) . 

WORLD DISTR~BUTION.-S~~ Lanka, Sumatra, Thailand, 

Malaya, Java, Borneo, Philippines, Taiwan, Japan, New 

Guinea, New Caledonia, and Society Islands. 

FERTILITY .-All 8 specimens examined had sporophytes. 

REMARKS.-A~~ the specimens examined were collected on 

wood or trees in moderately shaded places between 1200 m and 

1800 m. This is one of the few epiphytic Distichophyllum 

species in Malaya. It is not likely to be confused with any other 

species of the genus in Malaya by virtue of its long cuspidate 

arista, collenchymatous leaf cells, and costa ending just below 

the apex. 

SELECTED SPECIMENS EXAMINED.-Pahang: Gunung UlU 

Distichophyllum jungermannioides 

FIGURES 58-64 

Dirfichophy22urnjwlgermannioides (C. Muller) Bosch & Lacoste, Bryol. Jav., 

Mniadelphus jungermnnnioides C. Muller, Syn., 2:660, 1851. [Type: Java, 

Mniadelphus nanus (Dozy & Mokenboer) Jaeger, Ber. St. Gall. Naturw. Ges., 

Disfichophyllurn nunurn Dozy & Molkenboer in Zolhger, Syst. Verzeichn., 

Dirrichophyllurn ulukalienre Damanhuri & Mohamed, I. Bryol., 14:329, 1986. 

2:22, 1861. 

Elume s.n. (B, destroyed).] 

1875-1876:324, 1877 (Ad. 2:228). 

32, 1855. [Type: Java, Herb. van den Bosch (L!).] 

[Type: Pahang, Gunung Ulu Kali, Dumonhuri 264.5 (KLU!).] syn. nov. 

Plants small, in mats, light green: stems suberect, to 8 mm 

long, 2.5 mm wide with leaves when wet, simple or branched; 

leaves crispate and slightly shrunken when dry, ovate to 

obovate, 1.1-1.5(-1.9) mm long, 0.7-1 mm wide, widest 

between midleaf and 3/4 from base of leaf, obtuse to rounded, 

apiculate, the apiculus (50-)70-90 pm long, often twisted; 

margins entire, plane: borders well differentiated, with 1-2 

rows of linear cells near apex, 2(-3) rows towards base; costa 

rarely extending beyond midleaf; median cells rounded to 

hexagonal, thin walled, weakly collenchymatous, 20-25 pm 

long, 16-21 pm wide, more or less homogeneous in upper half 

of leaf; basal cells hexagonal to rectangular, 40-50 pm long, 

18-22 pm wide. Dioicous. Calyptra smooth, brown above, 

colorless and fringed below. Seta to 4 mm, smooth: capsule 

ovoid; urn 0.7 mm long: opcrculum rostrate. 

SELECTED SPECIMENS ExAM1SED.-Pahang: Pulau Tioman, 

Gunung Kajang, 840 m, Zumzuri 162; Cameron Highlands, 

1560 m, H. Mohamed 9117~; Gunung Ulu Kali, 5000 ft, 

Manuel 27266. 

WORLD DISTRIBUTION.-Java, Malaya, Sumbawa, Botel 

Tobago, and Taiwan. 

FERTILITY.-Eight of the 10 packets examined had sporophytes. 

REMARKS.-Distichophyllum jungermannioides is a new 

record for Malaya. It has been collected on the lower trunk of 

trees above 1400 m on the mainland while on the island of 

Tioman it has been collected at 800 m. This is a highly variable 

species. The type specimen has apiculi measuring 84-108 pm 

long whereas the Malayan specimens rarely have apiculi 

exceeding 90 pm. Zumzuri 162 is a robust specimen in which 

the median cells measure 15-20 pm wide and the costa 

sometimes extends to 3/4 of leaf length. The leaves are also 

larger, measuring 1.5-1.9 mm long, 0.7-1 mm wide. This 

species is easily recognised by its small size, short costa, and a 

distinct apiculus. The type specimen of Distichophyllum 

ulukaliense Damanhuri & Mohamed differs from the type 

specimen of D. jungermannioides in having a shorter apiculus, 

40-60(-80) pm long, which is not twisted, and in having 

smaller leaves and leaf cells than those of D. jungermannioides, 

but they are similar in all other aspects. Distichophyllum 

brevicuspes differs from the present species in having much 

larger leaves which are distinctly carinate along the costa and 

median cells which are collenchymatous. 

Distichophyllum maibarae 

Dkfichophyllum maibarae Bescherelle, J. Bot. (Morot), 13:40. 1899. [Type: 

Central Japan, Maibara, Faurie 11130 (holotype, BM!).] 

Plants small to medium sized, in loose tufts: stems creeping, 

to 8 mm long, 3 mm wide with leaves when wet, simple, rarely 

branched: leaves slightly shrunken and crisped when dry, 

ovate-oblong to obovate, 1.3-2 mm long, 0.6-10 mm wide, 

widest between midleaf and 3/4 from base, slightly asymmetrical, rounded to obtuse with a stout apiculus, the apiculus 

(30-)45-80 pm long: margin entire and plane: borders well 

differentiated, with 1-2 rows of linear cells near the apex, 2-3 

rows towards the base; costa flexuose above, extending 3/s to 4/5 

of leaf length: median cells small, round to hexagonal, 9-14 

pm in width, thin-walled, not collenchymatous, more or less 

homogeneous in upper half of leaf and in submarginal region 

towards base: paracostal cells in lower half of leaf larger than 

submarginal cells, hexagonal to rectangular, 35-60 pm long, 

18-25 pm wide. Autoicous. Calyptra with several hairs on the 

surface, fringed at base. Seta smooth, to 7 mm long: capsule 

inclined, ovoid, 1 mm long; exothecial cells mamillose, 

collenchymatous; operculum rostrate. 

SELECTED SPECIMEN EXAMINED.-Pahang: Cameron Highlands, Gunung Brinchang, 1760 m, lower montane forest, on 

tree base facing fast flowing stream, H. Mohamed & Zamzuri 

1186 (mixed with Distichophyllum cuspidatum). 
NUMBER 80 23 

63 

58 

62 

5.0 mm 

59-60 

1.0 mm 

6 1-64 

64 

0.1 mm 

FIGURES 58-64.4istichophyllwn jungermonnioides (Pahang: H. Mohamed 9117~): 58, plant, wet; 59, 60, 

leaves; 61, cells at leaf apex; 62, median cells of leaf; 63, cells at margin of midleaf; 64, paracostal cells at base 

of leaf. 

WORLD DISTRIBUTION.-Japan, China, Taiwan, Malaya, 

and Philippines. 

REMARKS.-Distichophyllum maibarae is newly recorded 

for Peninsular Malaysia. This is the only species of the genus in 

Malaya which possesses hairy calyptrae. Apart from this 

character, the homogeneous nature of the cells in the upper half 

of leaf, the small size of the cells, and the distinct apiculus will 

set it apart from other species. Although Bescherelle (1899) 

compared it to D. nigricaule, Noguchi (1956) and Tan & 

Robinson (1990) believed that it is more closely related to D. 

mntagneanum (C. Muller) Bosch & Lacoste. To study the 

relationships between these two species, the type specimens of 

both the species as well as non-types were examined. The type 

specimen of D. maibarae specifically differs from the type of 

D. montagneanum in having a longer, stouter apiculus on the 

leaf. The type of D. montagneanum does not possess 

sporophytes. However, a number of specimens of this species 

from Sri Lanka and India have sporophytes which have smooth 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

e 

24 

72 

65 

5.0 mm 

66,67 

1.0 mm 

68-72 

0.1 mm 

FIGURES 65-72.-Dis1ichophyllum mittenii (Pahang: H. Mohamed & Damanhuri 5057): 65, plant, wet; 66,61, 

leaves; 68, paracostal cells near midleaf; 69, cells at leaf apex; 70, cells at base of leaf; 71, cells at margin of 

midleaf; 72, paracostal cells at tip of costa. 
NUMBER 80 25 

calyptra. In his keys to the genus Distichophyllum in Indonesia, 

Fleischer (1908) described D. montagneanum as having a hairy 

calyptra (. . .haube mit aufrechten Cilien besetzt.). By courtesy 

of the staff at FH, we were able to examine the three specimens 

from Java cited by Fleischer: (1) bei Tjibodas am Gedeh 1450 

m, (2) bei Huis Ten Bosch am Gedeh 2100 m, (3) 

Tjibodas-Trawas Pandjang, 1900 m. All three specimens have 

sporophytes with hairy calyptrae. The fourth specimen cited by 

Fleischer, which is from Nilghiris, India, could not be located. 

However, another specimen labelled by Fleischer as D. 

montagneanum has sporophytes which have have smooth 

calyptrae. Because D. montagneanum and D. maibarae are 

similar in leaf shape, areolation, and in the size of the leaf cells, 

Fleischer confused the two species. Distichophyllum montagneanum evidently is restricted to India and Sri Lanka, and the 

specimens from Java under the name D. montagneanum with 

hairy calyptrae are D. maibarae. 

Distichophyllum malayense 

Distichophyiium maloyenre Damanhuri & Mohamed, J. Bryol., 14:327, 1986. 

[Type: Pahang, Genting Highlands, Damanhuri 2723 (KLU!).] 

Plants in lax tufts, pale green; stems to 6 mm high and 2.5 

mm wide with leaves, brown to reddish brown, mostly simple, 

rarely branched; leaves slightly contorted and undulate when 

dry, elliptic to ovate-lanceolate, 1.8-2.5 mm long, 0.5-0.7 mm 

wide, acuminate with a long subulate point, usually twisted 

when dry, the apiculus 250-450 pm long; margins entire; 

border distinctly differentiated, consists of 1-2 rows of linear 

thin-walled cells; costa weakly defined, reaching up to 2/5 of 

leaf length, rarely longer; median leaf cells more or less 

uniform in size, lax, thin-walled, large, hexagonal, 38-77 pm 

long, 30-57 pm wide; basal cells lax, rectangular, 64-123 pm 

long, 25-40 pm wide; perichaetial leaves without costa, 

oblong-lanceolate, 0.9-1.2 mm long, 0.2-0.3 mm wide; apex 

subulate. Dioicous. Sporophyte unknown. 

SELECTED SPECIMENS EXAMINED.-Pahang: Genting Highlands, 960 m, Damanhuri 2723; 750 m, Damanhuri 3028 

(KLU, UKMB). 

WORLD DISTRIBUTION.-Endemic to Malaya. 

REMARKS.-Distichophyllum malayense has been collected 

on rotten logs in shade near streams between 750 m and 1000 

m. It is distinct in having leaves with a long subulate point and 

a weakly developed costa extending less than l/2 the leaf length. 

It may be confused with D. cuspidatum, but the distinct costa 

extending to 3/4 of the leaf length and the narrower leaf cells in 

the latter will set it apart. 

Distichophyllum mittenii 

FIGURES 65-12 

Distichophyllum miffenii Bosch & Lacoste, Bryol. Jav., 2:25, 1861. [Types: 

Java, sine loco, Teysmann s.n. (L). not seen; Holle s.n. (L). not seen; prop 

Toegoe, pr. Bogor, Amann s.n. (L). not seen.] 

Plants robust, forming mats; stems simple or branched, 

prostrate, rarely ascending, complanate, to 9 cm long, 1 cm 

wide with leaves; leaves densely set on stem, contorted and 

moderately crisped when dry, 3.4-6.0 mm long, 1.4-2.2 mm 

wide, widest above mid-leaf, spathulate to obovate, broadly 

rounded above with one or two tiny apiculi, apiculus often less 

than 20 pm long; costa strong, reaching 3/4 to 4/5 of leaf length; 

margins entire; borders narrow, with 1 row of linear cells at the 

apex and 2 rows below; submarginal cells much smaller than 

paracostal cells, quadrate to oval, 10 to 20 pm wide, forming 

several rows near margin above; paracostal cells hexagonal to 

polygonal, 30 to 60 pm long, 25 to 35 pm wide, thick-walled; 

cells at base rectangular or long-hexagonal, 70 to 100 pm long, 

20-35 pm wide. Autoicous. Calyptra scabrous, base fringed 

with long cilia. Seta up to 8 mm, scabrous throughout, the 

projecting papillae 25 to 35 pm long; capsules inclined to 

horizontal, ovoid to ovoid-oblong, exothecial cells mamillose; 

operculum conic-rostrate. 

SELECTED SPECIMENS EXAMINED.-Pahang: H. Mohamed 

& Damanhuri 5057; Cameron Highlands, Gunung Brinchang, 

H. Mohamed & Zamzuri 1169; Gunung Ulu Kali, Stone 13378. 

Perak: Maxwell Hill, Gunung Hijau Trail, Manuel 2341. 

WORLD DISTRIBUTION.-Sri Lanka, Thailand, Malaya, Java, 

Sumatra, Borneo, Philippines, Taiwan, Japan, New Guinea, 

New Caledonia, and Vanuatu. 

FERTILITY.-Eight of the 12 packets examined had sporophytes. 

REMARKS.-This species has been collected above 1000 m 

on rotten logs, humus, and soil in moist shady places. 

Distichophyllurn nigricaule 

FIGURES 51,54,55,57 

Dislichophyllum nigricoule Mitten ex Bosch & Lacoste, Bryol. Jav., 2:20, 

1861. [Types: Java, Mt. Pangerango, Motley s.n. (lectotype, NY!); sine loco, 

Teysmann s.n. (syntypes. BO, L), not seen).] 

Distichophyllum grucilicaule Fleischer, Musci F1. Buitenz., 3:983, 1908. 

[Types: West Java, um Tjibodas am Gedeh. Flekcher s.n. (lectotype, FH!); 

am Poentjak, Flekcher s.n. (syntype, FH!).] 

Plants in medium-sized mats, light green; stems with leaves 

to 2 cm long, to 3.5 mm wide when wet, simple or branched; 

leaves distantly placed on stem, scarcely shrunken and only 

slightly crispate when dry, slightly concave, ovate to broadly 

elliptic, rarely obovate, 1.5-2.2 mm long, 0.95-1.1 mm wide, 

widest at midleaf, obtuse to rounded with a distinct apiculus, 

the apiculus (25-)30-60 pm long; margin entire and plane; 

borders well differentiated, with 1-2 rows of linear cells near 

apex, 3-4 rows below; costa flexuose above, extending from 

3/4 to 4/5 of leaf length; median cells hexagonal to rounded, 

16-23 pm wide, moderately thick-walled, weakly collenchymatous, more or less homogeneous in upper half of leaf and 

submarginal region towards base; paracostal cells towards base 

larger, hexagonal to rectangular, 45-90 pm long, 20-27 pm 

wide. Dioicous. Calyptra smooth, fringed, and yellowish 
26 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

below, dark brown above. Seta smooth, to 7 mm long; capsule 

too young to be examined. 

SELECTED SPECIMENS EXAMINED.-Pahang: Fraser Hill, 

980 m, H. Mohamed & Zamzuri 9225~ & 9226; Genting 

Highlands, Damanhuri 2556, 2568, 2570 & 2574; Pulau 

Tioman, Gunung Kajang, 920 m, Zamzuri 170. 

JAVA: Batavia, Gunung Pasir Angin, near Gadok, regio 

calida, 500 m, Schiffner 12732 (US). 

WORLD DIsTFuBun0N.-Java, Borneo, Malaya, and Thailand. 

REMARKS.-Tan & Robinson (1990) noted that when 

Fleischer (1908) described D. gracilicaule, he had been 

misguided by atypical specimens of D. nigricaule. Fleischer 

stated that the costa of D. nigricaule is often percurrent and that 

the submarginal cells are distinctly smaller than the inner ones. 

The lectotype of D. nigricaule from NY does not show such 

distinctions. Evidently as a result of Fleischer’s misconception 

of D. nigricaule, many later authors such as Bartram (1939) 

have mistakenly cited D. cirratum as a synonym of D. 

nigricauie. 

Distichophyllum nigricaule differs from D. cirratum in the 

wider leaf border near the apex and homogeneous size of the 

leaf cells in the upper half of the leaf. In addition, D. nigricaule 

differs from D. cirratum by its leaves which are only slightly 

crispate when dry. In D. cirratum the leaf cells are often lax and 

collenchymatous whereas in D. nigricaule the cells are often 

firm and only slightly collenchymatous. In Malaya, D. cirratum 

has been collected only above 1000 m, whereas D. nigricaule 

has only been collected below 1000 m. 

Distichophyllum osterwaldii 

FIGURES 73-79 

Disrichophyllurn osfenualdii Fleischer. Musci Fl. Buitenz., 3:994,1908. [Type: 

West Java, Gedehgebirge bei Tjibodas, Fleircher s.n. (FH!).] 

Plants robust, laxly tufted, green to dark green; stems 

ascending from a rhizomatous base, to 4 cm high, often 

branched, laxly foliate, flat, to 6 mm wide with leaves; leaves 

weakly undulate-crispate when dry, lateral rows widely 

spreading, lingulate, 4-6 mm long, 1.7-2.5 mm wide, rounded 

above and muticous; margins sometimes undulate, minutely 

crenulate above where unbordered; not bordered at apex, 

bordered below by 2-3 rows of linear pellucid cells for 3/4 of 

leaf length; costa extending to 4/5 of leaf length; median cells 

isodiametric to irregularly hexagonal, 15-22 pm wide near 

apical margin, quadrate to hexagonal, 25-40 pm wide near 

apical portion of costa; cells near base laxer, rectangular to 

elongate-hexagonal, 50-1 15 pm long, 20-27 pm wide. 

Dioicous. Calyptra fringed at base and scabrous above. Seta 

0.8- 1.2 cm long, flexuose, distinctly papillose throughout, 

papillae to 23 pm high; capsule inclined, 1 mm long. 

SELECTED SPECIMENS ExAM1sED.-Pahang: Cameron 

Highlands, Gunung Brinchang, H. Mohamed 6; Damanhuri 

5059b & 5074; Damanhuri 2014; Gunung Ulu Kali, area 

between microwave station and radar station, Manuel 2457. 

WORLD DISTRIBUTION.-Malaya, Borneo, Java, Philippines, 

Taiwan, Mainland China, and Japan. 

FERTILITY.-Only 5 of the 8 specimens examined had 

sporophytes. 

REMARKS.-Distichophyhm osterwaldii has been collected 

on shaded moist humus and wood in montane ericaceous forest. 

All 8 specimens examined were collected between 1600 m and 

2100 m. The Malayan specimens of D. osterwaldii differ in a 

number of ways from the type from Java. The type specimen is 

extremely large, measuring to 8 cm long, while the Malayan 

specimens rarely measure above 4 cm. The leaves in the type 

are distantly arranged on the stem and are distinctly undulate 

whereas the local specimens are densely set on the stem and 

only slightly undulate. Further, the leaves in the type are 

unbordered in the upper half of the leaf while the Malayan 

specimens are unbordered only in the upper quarter of the leaf. 

Distichophyllum ostenvaldii is closely related to D. deniiculatum Dixon from Borneo and D. obtusifolium ThCriot from 

Japan. However, the latter two species differ in being much 

smaller in size and in having at least one row of differentiated 

cells at the apex, Distichophyllum denticulatum has a distinctly 

denticulate margin at the apex whereas D. osterwaldii has at 

most a crenulate margin. Distichophyllum obtusifolium differs 

from the present species in having wide, homogeneous median 

cells in the upper half of the leaf. 

Distichophyllum schmidtii 

FIGURES 80-86 

Disfichophyllwn schrnidrii Brotherus, Bot. Tidskr., 24: 122, 1901. [Holotype: 

Siam, Ins. Koh Chang, ad rupos irror in silvos, 1900, J. Schmidt (H-BR!).] 

Plants medium- to large-sized, in mats; stems prostrate, to 

3.5 cm long, 5 mm wide with leaves when wet, simple or 

branched; leaves scarcely shrunken and slightly crisped when 

dry, oblong-ovate to obovate, 2-3 mm long, 0.9-1.1 mm wide, 

widest between 3/5 to 4/5 from base of leaf, round to obtuse with 

a small mucro, the mucro 30-50(-60) pm long; margins entire 

and plane; borders well differentiated, with 1-2 rows of linear 

cells near the apex, 2-3(-4) rows of linear cells near the base; 

costa flexuose on top, extending from 3/4 to 4/s of leaf length; 

median and upper cells irregularly hexagonal, 20-35 pm long, 

14-16(-20) pm wide, thin-walled, more or less homogeneous 

in upper quarter of leaf and submarginal region to midleaf, 

paracostal cells much larger below, lax, hexagonal to rectangular, 50-95 pm long, 15-28 pm wide. Dioicous. Calyptra 

smooth, fringed at base. Setae up to 8 mm high, scabrous 

throughout, the projecting papillae 8- 10 pm long; operculum 

rostrate. 

SELECTED SPECIMENS EXAMINED.-Selangor: Templer 

Park, Manuel 2692,2823; Sadiah & Zumzuri 1565,1568. 

SINGAPORE: Bukit Timah, Holttum 19328. 
NUMBER 80 27 

78 79 

5.0 mm 

7475 

1.0 mm 

76-79 

0.1 mm 

FIGURES 73-79.4ktichophylIum ostenvaldii (Pahang: Damanhuri 2014): 73, plant, wet; 74, IS, leaves; 76, 

cells at apex of leaf; 77, basal cells of leaf; 78, cells at margin at midleaf; 79, paracostal cells at tip of costa. 
28 

83 

85 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

80 

5.0 mm 

8 1-82 

1.0 mm 

83-86 

0.1 mm 

FIGURFS 80-86.-Distichophyllwn schmidtii (Pahang: Manuel 2692): 80, plant, wet: 81, 82, leaves; 83. cells at 

apex of leaf; 84, cells at margin near midleaf; 85, paracostal cells below midleaf; 86, cells at base of leaf. 
NUMBER 80 29 

THAILAND: Payap, -30 km S of Khun Sam, Touw 11323. 

WORLD DISTRIBUTION.-Singapore, Malaya, Thailand, and 

FERTILITY .-Of the 6 packets examined, 5 had sporophytes. 

REMARKS.-unlike other species of this genus in Malaya, D. 

schmidtii grows at very low altitudes. The type was collected 

near Lem Dan which has an altitude of around 100 m. In 

Malaya and Singapore, it has been collected below 100 m while 

Touw 11323 (L) was collected at 550 m. It grows on moist 

rocks and bases of trees near streams. Distichophyllum 

schmidtii superficially resembles D. mittenii. However, D. 

schmidtii differs from the latter in the smaller size of the plants 

and leaves, homogeneous areolation in the upper quarter of the 

leaf, broader leaf border, longer apiculi, and smooth calyptrae. 

In addition, D. mittenii is a plant of higher altitudes (above 

1000 m). The present species seems to be closely related to 

Distichophyllum lorzanum Fleischer from New Guinea in 

having leaves with similar morphology, homogeneous leaf 

areolation, short apiculi, and a narrow border. More specimens 

of the latter must be studied to ascertain the relationship to the 

present species. Although D. schmidtii resembles D. tortile in 

possessing homogeneous upper leaf cells, it has narrower leaf 

cells 14-16(-20) pm compared to cells wider than 30 pm in 

the latter. 

Bangladesh. 

Distichophyllum spathulatum 

FIGURES 87-93 

Distichophyllum spathulatum (Dozy & Molkenboer) Dozy & Molkenboer, 

Hookeria spathulata Dozy & Molkenboer, Ann. Sc. Nat. Bot., ser. 3, 2:305, 

Musci Frond. Ined. Archip. Indici, 4:103, 1846. 

1844. [Type: West-Java, Buitenzorg? Miquel (L), not seen.] 

This species is similar in all aspects to D. mittenii except that 

it has a dioicous inflorescence. 

SELECTED SPECIMENS EXAMINED.-Pahang: H. Mohamed 

& Damanhuri 5015; Cameron Highlands, Holttum 23369 

(SING); Fraser’s Hill, Burkill 2088 (SING). Perak: Maxwell 

Hill, Burkill 13197 (SING). 

WORLD DISTRIBUTION.-Thailand, Malaya, Java, Sumatra, 

Taiwan, New Caledonia, and Tahiti. 

FERTILITY.-Thirty-one of the 42 specimens examined had 

sporophytes. 

REMARKS.-This species is the largest and most common of 

the genus in Malaya, and forms extensive mats on moist shaded 

humus and rotting logs in montane ericaceous forests. Plants 

are rarely found growing on soil and rocks. Thirty nine of the 

specimens were collected between 1500 m and 2100 m and 

three of them were collected between 1200 m and 1400 m. 

Bosch & Lacoste (in Dozy & Molkenboer, 1861) distinguished 

this species from D. miftenii on the basis of sexuality; D. 

spathulatum being dioicous and D. mittenii being autoicous. 

Such a distinction is present in Malayan plants although there 

are no differences in other gametophytic characters or in the 

sporophytic characters. Both species are found in the same 

locality but D. spathulatum is much more common than D. 

mittenii. 

Distichophyllum tortile 

FIGURES 94-98 

Dktichophyllum tortile Dozy & Molkenboer ex Bosch & Lacoste, Bryol. Jav.. 

2:27, 1862. [Types: Java, Gedeh. Teysmnn s.n. (L), not seen: Banca, sine 

loco, Amnn s.n. (L). not seen.] 

Plants robust, green to dark green; stems to 4 cm long, to 6 

mm wide with leaves; leaves loosely inserted on stem, crisped 

when dry, oblong-lingulate to slightly obovate, 3.3 to 4.4 mm 

long, 1.0-1.4 mm wide, widest at distal l/4, obtuse above, 

apiculate, the apiculus 50-100 pm long; margins entire, plane; 

borders distinct, with 2 rows of linear cells near the apex, 3(-4) 

rows below; costa reaching 3/4 to 4/5 of leaf length; median leaf 

cells large, homogeneous in the upper part of leaf, hexagonal, 

40 to 55 pm wide, thin-walled; basal cells rectangular to 

long-hexagonal, 70-1 10 pm long, 30-40 um wide. Dioicous. 

Calyptra smooth above, fringed below. Seta 5-7 mm long, 

papillose, the projecting papillae to 15 pm high. 

SELECTED SPECIMENS EXAMINED.-Pahang: Cameron 

Highlands, H. Mohamed, et al. 9181e, 9185a, 9207,9224 and 

9235; Fraser Hill, trail to Pine Tree Hill, Manuel 2766. Perak: 

Gunung Hijau trail, Manuel 3248,3256. 

WORLD DISTRIBUTION.-Thailand, Malaya, Indochina, 

Java, Banka, Borneo, and Philippines. 

FERTILITY.-Only 2 of the 9 specimens examined had 

sporophytes. 

REMARKS.-Distichophyllum tortile has invariably been 

collected on rocks and humus in constantly moist and shady 

habitats in moderately high altitudes between 950 m and 1640 

m. This species is very distinct in the large median cells which 

are homogeneous in the upper part of the leaf. The wide, 

distinct borders also set this species apart from other large 

Distichophyllum species in Malaya. 

Excluded Species 

Distichophyllum undulatum Dozy & Molkenboer ex Bosch 

& Lacoste was reported from Malaya by Dixon (1926) based on 

Wray 945 collected from Perak, Gunung Batu Puteh (BM, 

SING). The species was also reported by Noguchi (1973) based 

on the specimens Inoue 10432, 16345, 16356, and 16364 

collected in Pahang, Cameron Highlands (NICH). All the 

specimens are D. mittenii. 

Family HYPOPTERYGIACEAE 

HYFQPTERYGIACEAE Kindberg, Hedwigia. 40:277,1901. 

Plants medium-sized to robust, primary stems elongate or 

creeping, densely tomentose; secondary stems simple, 
30 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 87-93.-Distichophyllum spathulatwn (Pahang: H. Mohamed & Damanhuri 5015): 87, plant, wet; 88, 

89, leaves; 90, cells at leaf apex; 91, cells at margin near midleaf; 92, cells at base of leaf, 93, median cells of leaf. 
NUMBER 80 31 

94 

5.0 mm 

95,96 

1.0 mm 

97,98 

0.1 mm 

FIGURES 94-98.4i.stichophyllm tortile (Pahang: H. Mohamed et al. 9181e): 94, plant, wet; 95,96, leaves; 91, 

cells at margin near midleaf; 98, cells at leaf apex. 

plumose, or dendroid, often with clusters of brood filaments on 

stems among upper leaves; leaves dimorphous, lateral rows 

complanate, ovate, asymmetrical, serrate, narrowly bordered; 

costae single, ending below tip or excurrent; cells hexagonal, 

smooth; ventral leaves (amphigastria) smaller, symmetrical, 

appressed to stem, acuminate. Calyptra cucullate or conical, 

naked. Capsules exserted, erect or pendulous; peristome 

double, teeth papillose or striolate, with a zig-zag median line; 

operculum rostrate. 

The relationship of this family to Hookeriaceae has been 

discussed in detail by various authors (Miller, 1971; Buck, 

1987; Whittemore & Allen, 1989; Tan & Robinson, 1990). In 

this treatment, the traditional placement of Cyathophorella, 

Lopidium, and Hypopterygium in the Hypopterygiaceae is 

followed. Most members of this family inhabit forests at 

moderately high to high altitudes. Hypopterygium aristatum 

occurs on limestone at low altitudes, but it also grows in 

non-limestone habitats at high altitudes. 
32 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Key to the Genera of Malayan Hypoptergiaceae 

1. Secondary stems simple, caudate at tips; exostome teeth papillose; costae less than l/2 

leaf length ............................. Cyath oph ore lla 

Secondary stems branched, not caudate at tips; pinnately branched or dendroid; 

peristome teeth striolate; costae more than l/2 leaf length ............ 2 

2. Plants often dendroid; stem with central strand; costae ending just above 1/2-3/4 leaf 

length in lateral leaves ...................... Hypopterygium 

Plants pinnately branched, often plumose; stem without central strand; costae 

excurrent in lateral leaves ........................ Lopidium 

Genus Cyathophorella 

Cyathophorella (Brotherus) Fleischer, Musci Fl. Buitenz., 3:1088, 1908. 

Cyathophorwn Sect. Cyathophorella Brotherus, Nat. Pfl., 1(3):965, 1907. 

Large, dull green plants in lax tufts; primary stems creeping, 

rhizomatous, tomentose; stems with central strand; secondary 

stems simple, rarely forked, distantly foliate, tips caudate and 

often with abundant brood filaments; lateral leaves in two rows, 

asymmetrical, widely spreading, smaller toward base and tip, 

ovate, acuminate, unbordered or weakly bordered, more or less 

toothed; costae short, single or forked; cells oval-hexagonal, 

smooth; amphigastria smaller, symmetrical, in one row, with 

axis parallel with stem. Dioicous. Calyptra conic, covering only 

rostrum of operculum, mostly naked, not fringed. Setae short, 

smooth; capsule erect; exostome teeth papillose; cilia lacking; 

operculum rostrate. 

The members of this genus are easily distinguished within 

the family by the simple, usually unbranched secondary stems. 

In Malaya, the genus is only found at high altitudes on tree 

bark. 

Key to the Malayan Species of Cyathophorella 

1. 

2. 

Leaf margins spinose-serrate ....................... C. spinosa 

Leaf margins weakly toothed .......................... .2 

Amphigastria ovate .......................... C. hookeriana 

Amphigastria broadly ovate to orbicular ................ C. burkillii 

Cyathophorella burkillii 

FIGURFS 99-109 

Cyathophorella burkil1ii (Dixon) Brotherus, Nat. Pfl., ed. 2, 11:278, 1925. 

Cyathophorwn burkillii Dixon, Rec. Bot. Sum. India, 6:67, 1914. [Type: Abor, 

2800 ft, on tree trunk, Burkill 37737 (SING!).] 

Plants robust, dull green; secondary stems to 5 cm high, to 8 

mm wide with leaves, densely foliate, caudate at tips, 

tomentose at base; leaves asymmetrical, ovate, 4.6-5.4 mm 

long 1.9-2.4 wide, ovate-lanceolate, short-acuminate with a 

long arista, the arista to 400 pm long; margins weakly toothed 

above; border well differentiated, with 2-4 rows of elongate 

cells; costae short, single or forked; cells oval-hexagonal; 

amphigastria symmetrical, broadly ovate to orbicular, obtuse to 

rounded with a long apiculus to 650 pm long; margins serrulate 

above; border weak or lacking; costae absent or only weakly 

developed: perichaetial leaves ovate, apex long-acuminate. 

Dioicous. Seta to 6 mm long; capsule horizontal to pendulous, 

2.6 mm long. 

SELECTED SPECIMENS EXAMINED.-Pahang: Cameron 

Highlands, 1400 m, Brinchang Road, on tree in forest, H. 

Mohamed et al. 9224,9229d. 

WORLD DISTRIBUTION.-India and Malaya. 

REMARKS.-This species is distinguished from C. hookeriana by its broadly ovate to orbicular amphigastrial leaves. The 

amphigastrial leaves of C. hookeriana are simply ovate. 

Cyathop horella hoo keriana 

FIGURES 110-121 

Cyathophorella hoobrianu (Griffith) Fleischer, Musci Fl. Buitenz., 3:1094, 

1908. 

Necbra hoobriana Griffith, Notul. P1. As., 2:464, 1849 (plate legend) and 

Icon P1. As., 2: pl. 84, fig. 2A, 1849. [Type: Khasia Hills, Griffith 187 

(BM!).] 

Plants robust, dull green; secondary stems to 5 cm high, to 8 

mm wide with leaves, densely foliate, caudate at tips, 

tomentose at base; leaves asymmetrical, gradually decreasing 

in size towards tips, 4.4-5.0 mm long, 1.4-2 mm wide, 

ovate-lanceolate, narrowed to an aristate point at apex, arista up 

to 400 pm long; margins distantly and irregularly toothed 

above, sometimes almost entire; border weakly differentiated, 

with 1-3 rows of elongate cells, often on one side only; costae 

short, ending below midleaf, forked; cells oval-hexagonal, 

52-84 pm long, 14-22 pm wide, larger and laxer towards 

base. Amphigastria symmetrical, ovate, 2.6-3.4 mm long, 

1.6-1.9 mm wide, short-acuminate with a long arista to 600 

pm; margin weakly toothed above, entire below; border weak 
NUMBER 80 33 

FIGURES 99-109.4yathophorella burkillii (Pahang: H. Mohamed et 01.9224): 99. plant, wet; 100-102, lateral 

leaves; 103-105, amphigastrialleaves; 106, cells at leaf apex; 107, median cells of leaf; 108, cells at uppermargin 

of leaf; 109, cells at base of leaf. 
34 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 80 35 

or lacking; costae variable, absent or ending below midleaf. 

Dioicous. Calyptra scabrous above. Seta to 1 cm long; capsule 

cylindrical, to 3 mm long. 

SELECTED SPECIMENS EXAMINED.-Pahang: Cameron 

Highlands, Gunung Brinchang, 1890 m, on rotten log and tree, 

semi-shaded area, H. Moharned 9083,9181,9182,9293~. 

WORLD DISTRIBUTION.-India, Sikkim, Vietnam, Malaya, 

Philippines, China, and Japan. 

REMARKS.-Cyathophorella hookeriana is the most common of the three species of the genus in Malaya. The plants are 

found growing on rotten logs, twigs and bases of trees in moist, 

shaded areas between 1700 m and 2000 m. The distinguishing 

characters of C. hookeriana are the ovate-lanceolate leaves 

with margins distantly irregularly toothed above and the strictly 

ovate amphigastrial leaves. The type specimen has almost 

entire margins, however, there are Malayan specimens that 

have leaves with almost entire margins as well as weakly 

toothed margins in the same plant. The type of C. anisodon 

Dixon & Herzog in BM (Darjeeling, Sikkim, Himalaya, 

Kerstan 13c) has leaves of similar shape and size and is 

irregularly and distantly toothed above as in Malayan plants, 

but it differs from the Malayan plants in possessing caudate 

branch tips. Cyathophorella burkillii is very similar in all 

aspects except in the shape of the amphigastrial leaves. The 

amphigastrial leaves in C. hookeriana are ovate whereas the 

amphigastria of C. burkillii are broadly-ovate to orbicular. The 

plants of C. hookeriana from Japan are much smaller than the 

Malayan specimens. 

Cyathophorella spinosa 

FIGURES 122-130 

Cyalhoporella spinosa (C. Muller) Fleischer, Musci Fl. Buitenz, 3:1O91,1908. 

Hookeria spinosa C. Miiller, Syn., 2677. 1851. [Type: Java, Blwne. Hb. A1 

Braun (B), not seen.] 

Plants robust, dull green; secondary stems to 5 cm long, tips 

caudate, to 1 cm wide with leaves, frequently with conspicuous 

clusters of orange-red brood filaments among leaves; leaves 

widely spreading, asymmetrical, broadly ovate, shortacuminate with an arista to 200 pm long, 4.5-6.0 mm long, 

2.4-2.8 mm wide, widest below midleaf; margin spinoseserrate in upper third with long multicellular teeth; border not 

or weakly differentiated; costae short, single or forked; cells 

oval-hexagonal, 77-95 pm long, 25-36 pm wide, thin-walled; 

amphigastria broad, rounded ovate with an apiculus to 250 pm 

long, distantly spinose above; costae short, unequally forked; 

perichaetial leaves smaller than lateral leaves; ovate with a long 

arista. Dioicous. Sporophytes not seen. 

SELECTED SPECIMEN EXAMINED.-Pahang: Cameron Highlands, Gunung Jasar, 1440 m, on base of Cyathea sp., H. 

Mo hamed 9322. 

WORLD DISTRIBUTION.-Thailand, Malaya, Java, Borneo, 

Moluccas, Philippines, Vanatua, and New Guinea. 

REMARKS.-This species is recognized by the spinoseserrate margins in the upper part of the leaves. Closest 

relationship is to two other species with spinose-serrate 

margins which are found in adjacent regions, C. adiantum 

(Griffith) Fleischer (Himalayas, Assam, Thailand, Philippines, 

and Borneo) and C. tonkinensis (Brotherus & Paris) Brotherus 

(Japan, Ryukyus, Taiwan, Vietnam, Thailand, and Borneo). 

Cyathophorella tonkinensis is distinct in having more densely 

foliate stems, mostly unicellular marginal teeth, and smaller 

amphigastria. Cyathophorella adiantum differs from C. spinosa in the less robust plants, the narrowly ovate leaves with 

longer acuminate apices, and the less toothed ovate amphigastria. 

Excluded Species 

Cyathophorella tenera (Bosch & Lacoste) Fleischer was 

reported from Malaya by Tixier (1971) based on Tixier 4827 

collected from Pahang, Fraser’s Hill (PC). The specimen is 

actually C. hookeriana. 

Genus Hypopterygium 

Hypopterygium Bridel, Bryol. Univ., 2:709, 1827. 

Medium-sized yellowish green plants with primary stems 

creeping, radiculose; stems with central strand; secondary 

stems erect, with a fan-shaped cluster of tertiary stems arising 

in upper part; leaves dimorphous, lateral leaves asymmetrically 

inserted, complanate, usually bordered by 1-3 rows of elongate 

cells; costae single, reaching from midleaf to 3/4 of leaf length; 

ventral row of leaves (amphigastria) symmetrically inserted, 

appressed to the stem; bordered by 1-2 rows of elongate cells; 

costae single, weakly defined to excurrent. Autoicous, dioicous, or heteroicous. Calyptra cucullate or conical, naked, not 

fringed. Seta elongate, smooth; Capsule inclined or pendulous; 

exostome teeth transversely striolate; cilia 2-3; operculum 

rostrate. 

Key to the Species of Malayan Hypopterygium 

Lateral leaves subulate acuminate, costae reaching to 3/4 of leaf length; costae of 

amphigastria distinct and excurrent . . . . . . . . . . . . . . . . . . . . H. aristutum 

Lateral leaves short-acuminate, costae reaching to 3/5 of total leaf length; costae of 

amphigastria weakly defined, ending well below apex. . . . . . . . . . . H. tenellurn 
36 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

12 

127 

FIGURES 122-130.-Cyuthophorellu spinosu (Pahang: H. Mohumd 9322): 122, plant, wet; 123, 124, lateral 

leaves; 125, 126, amphigastrial leaves; 127, multicellular tzeth on upper margin of lateral leaf; 128, cells at leaf 

apex; 129, median cells of leaf; 130, cells at base of leaf. 
NUMBER 80 37 

Hypopterygiu m arista tu m 

FIGURES 131-140 

Hypopterygium arisfutum Bosch & Lacoste, Bryol. Jav., 2:12, pl. 141, 1861. 

[Type: West-Java, am Gedeh und Pangerango 2090-3000 m, Motley (L), not 

seen.] 

Plants dendroid, to 2.5 cm in height, dull green; lateral leaves 

pinnately spreading, ovate, 1.2-1.5 mm long, 0.6-0.9 mm 

wide, widest below midleaf; apex subulate-acuminate with a 

distinct arista, to 200 pm long; margin toothed in upper part on 

one side of leaf, entire below; bordered all around by 1(-2) 

rows of elongate cells; costae reaching 3/4 of leaf length; 

median cells hexagonal, 20-35 pm long, 15-20 pm wide; 

amphigastria orbicular, 0.8-1.1 mm long, 0.6-0.8 mm wide; 

margin entire below, weakly and distantly toothed above; 

bordered all around by 1(-2) rows of elongate cells; costae 

distinct, excurrent, ending in a long arista to 350 pm long; 

median cells hexagonal, 20-35 pm in long, 13-20 pm wide. 

Synoicous. Seta smooth, to 1.4 cm long; capsule suberect to 

horizontal; urn oval-cylindrical, to 2 mm long; operculum to 2 

mm long. 

SELECTED SPECIMENS EXAMINED.-Pahang: Cameron 

Highlands, Robinson's Waterfall, H. Mohamed & Zumzuri 

1439; Taman Negara, Gua caves (limestone), Manuel 2619, 

2608. 

WORLD DISTRIBU'I?ON.-Phifippines, Malaya, and Java. 

REMARKS.-Hypopterygium aristatum grows mostly on 

rocks and sometimes on logs at altitudes above 1000 m. 

Specimens have been collected at 200 m but at such low 

altitudes the species invariably grows only on limestone. The 

species fruits abundantly throughout the year. Hypopterygium 

aristatum is generally much larger than H. tenellum and 

dendroid in habit. Other differences are discussed under H. 

tenellum. 

Hypopterygium tenellum 

FIGURES 141-150 

Hypopterygium tenellum C. Muller, Bot. Zeit., 12:557, 1854. [Type: India, In 

cortice arborum una cum Hypopt. struthiopteris circa Ootacamund a Perrotet 

(B), not seen.] 

Plants to 2.0 cm in height, dull green; lateral leaves pinnately 

spreading, 0.8-1.4 mm long, 0.6-0.8 mm wide, widest below 

midleaf, ovate with a short-acuminate apex, apiculate, the 

apiculus to 50 pm long; margin toothed in upper part, entire 

below, bordered all around by 1(-2) rows of linear cells; costae 

reaching 3/5 of leaf length; median cells hexagonal, 25-50 pm 

long, 12-15 pm wide; amphigastria 0.6-0.8 mm long, 0.5-0.7 

mm wide, orbicular with a sharp apiculus to 150 pm long; 

margins crenulate above, entire below, weakly bordered all 

around with 1(-2) rows of cells; costae weakly defined, 

extending up to '/2 of leaf length; median cells hexagonal, 

25-35 pm in length, 16-20 pm in width. Synoicous. Seta 

smooth, to 1 cm in length; capsule horizontal to cernuous; urn 

oval-cylindrical, 1 mm long. 

SELECTED SPECIMENS EXAMINED.-Pahang: Cameron 

Highlands, Robinson Falls, Manuel 3344, 3344a, 3362; Pulau 

Tioman, Gunung Kajang, Zumzuri 216,226. 

WORLD DISTRIBUTION.-Sri Lanka, Malaya, Sumatra, Java, 

Borneo, Philippines, and New Guinea. 

REMARKS.-Hypopterygium tenellum has been collected on 

the island of Tioman, at 600 m, but on the mainland it has been 

collected only above 1200 m. It has been collected on rocks and 

trees. This species is distinguished from Hypopterygium 

aristatum by its smaller size, amphigasuia that have weakly 

defined costae which seldom reach midleaf, and a shorter 

apiculus (to 150 pm). Hypopterygium aristatum is a larger 

plant, has an excurrent costa on the amphigastria, and a longer 

arista (to 350 pm). 

Genus Lopidium 

bpidium Hooker f. & Wilson, Fl. Nov. Zel., 2:119, 1854. [Lectotype: L. 

concinnum (Hooker) Wilson cf. Dixon, New Zealand Inst. Bull., 3(5):293, 

1927.1 

Plants moderately robust, light green to green, with creeping 

stoloniform shoots; stems without central strand; fronds erect, 

elongate, more or less regularly pinnate, with simple or slightly 

pinnate branches; leaves complanate, distinctly dimorphic; 

lateral leaves spreading, narrow, oblong-lingulate, acute with a 

distinct apiculus; margin denticulate to entire; costae single, 

excurrent; cells small, rounded, smooth, incrassate; amphigastria smaller than lateral leaves, ovate with a broad base; margin 

denticulate; bordered on both sides; unicostate, percurrent to 

excurrent. Autoicous or dioicous. Calyptra cucullate, short, 

naked, not fringed. Seta short, variably papillose; capsule 

suberect; exostome teeth mnversely striolate; cilia lacking; 

operculum long-rostrate. 

Key to the Malayan Species of Lopidium 

Lateral leaves bordered on both sides of the lamina . . . . . . . . . . . L. struthiopteris 

Lateral leaves bordered only on one side of the lamina . . . . . . . . . . L. trichocladon 
38 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

136 / i 

\ 

\ \ 

132 

2.0 mm 

131 

10 mm 

133- 136 

0.5 mrn 

139 137- 140 

0.1 mm 

FIGURFS 131-140.-Hypopterygiumarirtatum (Pahang: Manuel 2619): 131, plant, wet; 132, lateral branch, wet; 

133, 134, amphigastrial leaves; 135, 136, lateral leaves; 137, cells at apex of lateral leaf; 138, median paracostal 

cells of leaf; 139, cells at apex of amphigastrial leaf; 140, cells at margin near midleaf. 
NUMBER 80 39 

148 

1 

142 

2.0 mm 

141 

10.0 mm 

143- 146 

0.5 mm 

147- 150 - 

0.1 mm 

FIGURES 141-150.-Hypopterygiwntene[[um (Pahang: zOmzuri216): 141, plant, wet; 142, a lateral branch; 143. 

144, amphigastrial leaves; 145, 146, lateral leaves; 147, cells at leaf apex of lateral leaf; 148, median paracostal 

cells of lateral leaf; 149, cells of margin at middle of lateral leaf; 150, cells at apex of amphigastrial leaf. 
40 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Lopidiu m stru th iopte ris 

FIGURES 151-158 

Lopidrum siruihiopteris (Bridel) Fleischer, Musci F1. Buitenz., 3: 1073, 1908. 

Hypnwn struthiopteris Bridel, Spec. Musc., 237, 1812. [Type: In Insula 

Primary stems creeping, to 8 cm long; secondary stems erect, 

to 3.5 cm high, yellowish green, dull, densely bipinnate in an 

elongate plumose frond, branched nearly to base; stem leaves 

widely spreading, slightly arched with deflexed points when 

dry, ovate-lingulate from a broad, slightly clasping base, 

acuminate, 1.8-2.4 mm long, 0.8-1.1 mm wide, widest near 

the base; margins distantly denticulate in upper half; bordered 

all around with 1-2(-3) rows of linear cells (except at the 

base); costae excurrent in an arista to 150 ym long; basal 

portion of lamina often folded on one side; median cells 

rounded, 8-12 pm long, 6-10 pm wide, thick-walled, 

distinctly collenchymatous; amphigastria ovate-lanceolate with 

a wide base, subulate-acuminate, 0.7-1.2 mm long, 0.9-1.1 

mm wide, widest near base; distinctly bordered by 1-2 rows of 

linear cells except near the base; costae long-excurrent in an 

arista to 200 p high. Sporophytes not seen. 

SELECTED SPECIMEN EXAMINED.-Pahang: Cameron Highlands, Brinchang Rose Garden, 1540 m, on tree, H. Mohamed 

et al. 9185d. 

WORLD DISTRIBUTTON.- Reunion Island, India (Nilghiris), 

Sri Lanka, Malaya, Thailand, Sumatra, Java, Philippines, New 

Guinea, and New Caledonia. 

REMARKS.-bpidium struthiopteris has been collected 

between 1300 m and 1620 m on trees in partly shaded areas. 

The long plumose fronds of this genus are easily distinguished 

rrom the fanlike fronds of the genus Hypopterygium. Apart 

from the presence of the border on both sides of the lamina on 

Bourbonis, unde Commerson (Bridel herb. B), not seen.] 

the lateral leaves, there are no other clear-cut differences 

between this species and L. trichocladon. 

Lopidium trichocladon 

FIGURES 159-168 

Lopidim trichocladon (Bosch & Lacoste) Fleischer, Musci F1. Buitenz., 

Hypopterygium trichocladon Bosch & Lacoste, Bryol. Jav., 2:9, 1861. [Type: 

3:1069, 1908. 

West-Java, am Berg Salak, 600-800 m, A~M (L), not seen.] 

himary stems creeping, to 6 cm long, secondary stems erect, 

to 3 cm high, generally smaller than L. struthiopteris; stem 

leaves oblong-lanceolate, with a slightly broader base, 1.5-1.8 

mm long, 0.5-0.6 mm wide, short-acuminate; margins entire to 

slightly crenulate, bordered on only one side by 1(-2) rows of 

linear cells; costae excurrent in an arista, the arista up to 85 pm 

long; median cells round to hexagonal, thick-walled, distinctly 

collenchymatous, 8- 13 pm wide; amphigastria broadly ovate, 

long-acuminate, 0.9- 1.2 mm long, 0.6-0.8 mm wide, bordered 

on both sides of lamina by 1(-2) rows of linear cells, with 

border ceasing just above the base; margins weakly toothed 

above; costae percurrent to excurrent. Sporophytes not seen. 

SELECTED SPECIMENS EXAMINED.-Pahang: Cameron 

Highlands, Gunung Brinchang, 1760 m, on Cyarhea sp., H. 

Mohamed & Zamzuri 1160c, 1113b; Gunung Ulu Kali, 3500 ft, 

Manuel 3427; Pulau Tioman, Gunung Kajang, 480 m, on rock, 

Zumzuri 128. Perak: Maxwell Hill, 1280 m, on bark, H. 

Mohamed et al. 8091. 

WORLD DISTRIBUTION.-Thailand, Indochina, Malaya, 

Java, Sumatra, Borneo, Moluccas, Philippines, and Taiwan. 

REMARKS.-Lopidium trichocladon has been collected on 

living trees between 1000 m and 1700 m on the mainland but 

on Tioman Island, it has been collected on rocks at 480 m, and 

on trees at 840 m. 
NUMBER 80 41 

152 ~ 

153 ~ 

54 ~ 

151-154 

1.0 mm 

1 55- 1 58 

0.05 mm 

157 

FIGURES 151-158.-bpidium struthiopteris (Pahang: H. Mohamed et al. 918Sd): 151. 152, amphigastrial 

leaves; 153, 154, lateral leaves; 155, cells at leaf apex; 156, median cells of lateral leaf; 157, cells of margin at 

middle of lateral leaf; 158, cells at base of lateral leaf. 
42 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURFS 159-168.--Lopidium trichocladon (Pahang: H. Mohamed & hmzuri 1160~): 159, plant, dry; 160, 

lateral branch; 161, median cells of lateral leaf: 162, cells of margin at midleaf; 163, 164, lateral leaves; 165, 166. 

amphigastrialleaves; 167, cells at leaf apex; 168. cells at base of lateral leaf. 
Literature Cited 

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1939. 

Bescherelle, E. 

1898-1 899. 

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Bryologiae japonicae supplementum I. Journal de Botanique, 

12:280-300; 13:37-45. 

Buck, W.R. 

1987. Taxonomic and Nomenclatural Rearrangment in the Hookeriales 

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Damanhuri, A,, and M.A.H. Mohamed 

1986. 

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Dixon, H.N. 

1924. 

1926. 

193 1. 

1845-1854. Musci Frondosi Inediti Archipelagi Indici. Leiden. 

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New Genera of Asiatic Mosses. Journal of Botany, 69:l-7. 

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Dozy, F., and J.H. Molkenboer (completed by R.B. van den Bosch, C.M. van 

der Sande Lacoste) 

1855-1870. 

1908. 

1917. Mosses OJ Eastern India and Adjacent Regions. Fascicle 6 

Bryologiu Javunicu. 2 volumes. Leiden. 

Fleischer, M. 

Gangulee, H.C. 

Die Musci der Flora von Buitenzorg. Volume 3. Leiden. 

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The Affinities of Sclerohypnum Dix. Journal of Bryology, 10:277- 

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Mosses of Singapore and Malaysia. Singapore: University Press. 

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Harrington. A.J., and H.A. Miller 

1979. 

Johnson, A. 

1980. 

Koponen, T., and P. Isoviita 

1984. 

Hookeriaceae). Taxon, 33:112-113. 

Typification and Conservation of Generic Names in Musci. Acta 

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1959. 

Miller, H.A. 

1971. 

LMohamed, M.A.H. 

1985. 

11:80-82. 

Mohamed, M.A.H., and A. Damanhuri Mohamed 

1986. Four Genera of Mosses New to Peninsular Malaysia. Journal of 

B~ology, 14~333-338. 

Mohamed, M. A,H., and B.C. Tan 

1988. 

Noguchi, A. 

1956. 

1973. 

A Checklist of Mosses of Peninsular Malaysia and Singapore. The 

BryOlOgirt, 91~24-44. 

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A Revised Classification for the Orders and Families of Mosses. 

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1971. 

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35:2-3. 

Tan, B.C.. and H. Robinson 

1990. 

Tixier, P. 

1971. 

1980. 

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43 
Index 

(Synonyms and page numbers of principal accounts in italics) 

Actincdontium, 3,4 

ascendens, 4, figs. 5-8 

rhaphidostegum, 1,4, figs. 1-4 

Callicostella, 3,4, 6, 12 

beccariana. 1.6 

papillata, 4,6. figs 9-15 

prabaktiana, 6 

Calyptrochaeta, 3, 9 

remotifolia, 9, figs. 16-22 

Chaetomitriopsis, 1, 3, 9 

glaucocarpa, 9 

Chaetomitriurn, 1, 3, 9, 10, 11 

bomeense, 10 

ciliatum, 1, 11, 12 

elongatum, 1,9, 10, 11, 12 

lanceolaturn, 1. 12 

leptopoma, 10, 12 

rnuricatum, 1, 12 

nernatosurn. 1. 12 

orthorrhynchurn. 1, 10,II. 12 

papillifoliurn, 10.11, 12 

peraketue, 1, 11 

seriatum, 10 

setosum, 10,11 

torquescens, 1, 12 

adiantum, 35 

anisdon, 35 

burkiUii. 32, 35, figs. 99-109 

hookeriana, 1,32,35, figs. 110-121 

spinosa, 1, 32, 35, figs. 122-130 

tenera, 1, 35 

tonkinensis, 35 

Cyathophorum, 32 

burkiiiii, 32 

Cyathophorella, 3,31,32 

Sect. Cyothophoreila, 32 

blumeanum 12 

Daltonia, 3,12,13 

angustifolia, 1, 12.13, figs. 23-29 

aristifolia, 1, 13 

armata, 1, 12, 13, figs. 37-42 

contorta, 12, 13, figs. 30-36 

splachnoides, 12 

nymanianum, 17 

brevicuspes, 1, 17, 18, 19, 22 

cirratum, 18.19, 26, figs. 50. 52, 53, 56 

cuspidaturn, 13, 18, 19,22, 25, figs. 43-49 

denticulatum, 26 

eimeri, 19 

gracilicaufe, 25.26 

jungermannioides. 1, 18.22, figs. 58-64 

lorianum, 29 

maibarae, 1, 17, 18, 19,22, 23,25 

malayense, 1, 17. 18, 25 

mittenii, 17, 18,25,29, figs. 65-72 

montagneanum, 19,23,25 

nigricaule, 18, 19,23,25,26. figs. 51,54,55,57 

nigricaule var. cirralum, 19 

nigricaule var. eimeri. 19 

obtusifolium, 26 

ostenvaldii, 17, 18,26, figs. 73-79 

schmidtii, 17, 18, 19,26,29, figs. 80-86 

sinuosuium, 19 

spathulaturn. 17, 18,29, figs. 87-93 

tottile, 17, 18,29, figs. 94-98 

ulukalietue. 1, 22 

undulatum, 1,29 

Cyclodictyon, 3,6,12 

Dis tichophyllidium, 3.17 

Distichophyllum, 3.17, 18.22.25, 29 

MnWtl, 22 

Eriopus, 9 

remotifolius, 9 

Hookeria,4,6, 11, 12, 19,29, 35 

blwneana, 12 

curpidata, 19 

orthorrhynchum, 11 

ppiiiara. 6 

prabaktiana. 6 

rhaphidostega, 4 

spathuia fa, 29 

spinosa, 35 

giaucocarpon, 9 

ieptopoma, 10 

struthiopteris, 40 

aristaturn, 31, 35, 37, figs. 131-140 

tenellurn, 35,37, figs. 141-150 

trichocladon, 40 

Lopidium, 3 1, 32,37 

Hypnum, 9 

Hypopterygiurn, 3 1,32,35,40 

stmthiopteris, 3740, figs. 151-158 

trichocladon. 37,40, figs. 159-168 

Mniadeiphus, 22 

jung erma nn ioides ,22 

MRU, 22 

Neckera, 32 

hookeriana, 32 

Orhostichopsis. 10 

Pilotrichella, 10 

peraketuis, 10 

Schizomilriwn, 4 

Sclerohypnurn riparium, 1 

44