SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 75 

A Review of Philippine 

Hookeriaceous Taxa 

(Musci) 

Benito Tan and Harold Robinson 

SMITHSONIAN INSTITUTION PRESS 

Washington, D.C. 

1990 
ABSTRACT 

Tan, Benito, and Harold Robinson. A Review of Philippine Hookeriaceous Taxa (Musci). 

Smithsonian Contributions to Botany, number 7.541 pages, 63 figures, 1990.--The limits of the 

Hookeriaceae are discussed with the exclusion of some recently added elements, Cyathophorum, Cyathophorella, and Dendrocyathophorum. Of the genera traditionally placed in the 

family, CIraetomitrium is also excluded, but briefly reviewed. Segregate families based on 

peristome structure, such as the Daltoniaceae, are rejected. Papillose peristomes show at least 

three basically distinct types in the Hookeriaceae and they probably evolved at least three 

separate times. Keys are furnished to the 11 genera and 36 species of Hookeriaceae recognized 

in the Philippines and to the 17 species of Chaetomitrium in the country. Eleven species are 

newly reported from the country including four in Chaetomitrium, four in Distichophyllum, two 

in Leskeodon, and one in Calyptrochaeta. Bryobrothera is reported from the country. Ten new 

synonyms are cited. Five new combinations, Calyptrochaeta microblasta, C. ramosa, 

D istichophyllum nigricaule var. elmeri,Daltonia robbinsii, and Leskeodon brevicuspidatus, and 

one new name, Chaetomitrium schofieldii, are proposed. Lectotypes are selected for 

Calyptrochaeta ramosa (Fleischer) Tan & Robinson, Distichophyllum brevicuspes Fleischer, D. 

nigricaule Mitten ex Bosch & Lacoste, D. gracilicaule Fleischer, Lepidopihm sumatranum 

Bosch & Lacoste, and Hookeriopsis wichurae Fleischer. New records reported from other 

countries include Daltonia armata for Sumatra and Achrophyllum dentatum and Distichophyllum subnigricaule for New Guinea. 

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

recorded in the htitution’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 

A review of Philippine HodtCriaceous taxa (Musci) / Benito Tan and Harold Robinson. 

p. 

Includes bibliographical references. 

Supt of Docs. no.: SI 1.2975 

1. HookeriaccabPhilippilassi~cation. I. Robinson, Harold Ernest, 1932- . 11. Title. 111. Series, 

[QK539HH64] 581 Jdc20 [588’.2] 90-9530 

Tan, BcNto Chhg, 1948- 

cm.+Smithsonian contributions to botany ; no. 75) 
Con tents 

Page 

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

Abbreviations .................................... 3 

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

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

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

Note on Pterygophyllum novae-guineae ...................... 4 

Genus Actinodontium .................................. 5 

Key to the Philippine Species of Actinodontium .................. 5 

Genus Bryobrothera .................................. 6 

Bryobrothera crenulata ............................... 6 

Genus Callicostella ................................... 6 

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

Callicostella papillata ................................ 7 

Callicostella prabaktiana .............................. 7 

Genus Calyptrochaeta ................................. 7 

Key to the Philippine Species of Calyptrochaeta .................. 9 

Calyptrochaeta microblasta (Brotherus) Tan & Robinson. new combination ... 9 

Calyptrochaeta parviretis .............................. 9 

Calyptrochaeta ramosa (Fleischer) Tan & Robinson. new combination ..... 10 

Calyptrochaeta remotifolia ............................. 10 

Genus Cyclodictyon .................................. 11 

Cyclodictyon blumeanum .............................. 11 

Genus Daltonia .................................... 11 

Key to the Philippine Species of Daltonia ..................... 11 

Daltonia angustifolia ................................ 12 

Daltonia aristifolia ................................. 13 

Daltonia armuta .................................. 13 

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

Genus Distichophyllum ................................ 13 

Key to the Philippine Species and Varieties of Distichophyllum ......... 15 

Distichophyllum brevicuspes ............................ 17 

Distichophyllum catinifolium ........................... 17 

Distichophyllum collenchymatosum ........................ 11 

Distichophyllum cucullatum ............................ 18 

Distichophyllum cuspidatum ............................ 18 

Distichophyllum kinabaluenre ........................... 18 

Distichophyllum maibarae ............................. 19 

Distichophyllum nigricaule ............................ 20 

Distichophyllum nigricaule var . nigricaule .................... 20 

Distichophyllum nigricaule var . cirratum ..................... 20 

combination ................................... 22 

GenusAchrophyllum .................................. 4 

Achrophyllum javense ................................. 4 

Actinodontium ascendens .............................. 5 

Actinodontium rhaphidostegum ........................... 5 

Distichophyllum mittenii .............................. 19 

Distichophyllum nigricaule var . elmeri (Brotherus) Tan & Robinson. new 

iii 
iv SMITHSONIAN CONTRTBUTIONS TO BOTANY 

Distichophyllum obtusifolium ........................... 22 

Distichophyllum ostenvaldii ............................ 23 

Distichophyllum santosii .............................. 23 

Distichophyllum subcuspidatum .......................... 23 

Distichophyllum subnigricaule ........................... 23 

Distichophyllum tortile ............................... 24 

Genus Hookeriopsis .................................. 25 

Key to the Philippine Species of Hookeriopsis .................. 25 

Hookeriopsis utacamundiana ........................... 25 

Hookeriopsis wichurae ............................... 26 

Key to the Philippine Species of Leskeodon .................... 27 

Leskeodon acum’natur ............................... 27 

bination ..................................... 27 

Leskodon philippinensis .............................. 29 

Notes on Philippine Chaetom‘trium ......................... 29 

Key to the Philippine Species of Chaetomitrium ................. 30 

Chaetom’trium schofieldii Tan & Robinson. new name .............. 31 

Accepted Names of Philippine Chaetom’trium .................. 31 

Literature Cited ..................................... 32 

Genus Hookria .................................... 24 

Hookeria acutifolia ................................. 24 

Genus Leskeodon ................................... 26 

Leskeodon brevicuspidatus (Bartram) Tan & Robinson. new comFigures 35-63 ...................................... 34 

Index ........................................... 41 
A Review of Philippine 

Hookeriaceous Taxa 

(Musci) 

Benito Tan and Harold Robinson 

Introduction 

The present study has a primary aim of reviewing the 

members of the family Hookeriaceae in the Philippines. For 

this purpose, keys and descriptions are provided for species 

known from that country with accompanying illustrations of 

leaf structure (Figures 1-34) for some critical groups. In 

addition, observations are made on the limits of the family and 

possible subdivisions within the family. The interpretations of 

peristome characters are based on SEM studies of their detailed 

structure (Figures 35-63). The family limits and potential 

subdivisions of the family are considered fist, and these are 

followed by the systematic review of the family in the 

Philippines. 

The family Hookeriaceae has been repeatedly revised in 

recent years by Miller (1971),Crosby (1974),and Buck (1987). 

Its relationship with other pleurocarpous families was discussed by Robinson (1971,1986) who suggested a previously 

unrecognized closeness to the Sematophyllaceae and Hypnaceae. 

Buck (1987), believing in the heterogeneous nature of the 

family delineation and following the leads given by his 

predecesors (Miller, 1971; Crum, 1984; Allen, 1987a), proposed a formal split into five groups: Hookeriaceae (sensu 

stricto), Daltoniaceae, Callicostaceae, Leucomiaceae, and Adelotheciaceae. Although Buck (1987) based his new familial 

treatment on a combined consideration of both the sporophytic 

and gametophytic characters, the placement of some genera, 

in our opinion, is still unsatisfactory. For example, the 

circumscription of the redefied Hookeriaceae fide Buck 

(1987) is not convincing. His placement of Cyathophorum, 

Cyathophorella, and Dendrocyathophorum, traditionally of the 

Benito Tan, Imtitute of Biological Sciences and Museum of Natural 

History, University of the Philippines at Los Bahs, College, Laguna. 

The Philippines. Harold Robinson, Department of Botany, National 

Museum of Natural History, Smithsonian Institution, Washington, 

D.C. 20560. 

Hypopterygiaceae, in the vicinity of Hookeria is, at least, 

controversial. 

Buck (1987) justified his transfer of three cyathophoroid 

genera from the Hypopterygiaceae to the Hookeriaceae on the 

following shared character combination: little or un-branched 

stems, unbordered leaves, double costae, and hexagonal and 

porose leaf cells. The ranked foliation of the cyathophomid 

genera is thought to have evolved separately from that of 

Hypopterygium. However, a close re-examination of these 

similarities between the cyathophoroids and the hookerioids 

suggests to us that they are more likely the result of convergent 

evolution. The double costae of the Cyathophorum group are 

short, and the ranking of the leaves is unlike anything in the 

Hookeriaceae. Furthermore, leaves with hexagonal cells with 

or without borders are a rather widespread specialization. None 

of these characters seems particularly indicative of a close 

relation between the Cyathophorum group and the Hookeriaceae including Hookeria. 

One distinctive gametophytic difference between the cyathophoroid and hookerioid ma is their stem anatomy. In the 

former group, the stem anatomy shows a layer of thick-walled 

epidermis surrounding the cortex and a large central strand 

consisting of numerous small, hexagonal, and thin-walled 

hydroid cells. There is cellular differentiation within the strand. 

This is true in most members of Hypopterygiaceae (cf. Frey and 

Richter, 1982). In contrast, most hookerioid genera do not have 

a differentiated central strand in the stem. In a few genera like 

Hookeria, Calyptrochaeta, and Achrophyllum where central 

differentiation occurs in the stem, the central strand is a weakly 

defined group of parenchyma cells. 

Also, the leaf cell anatomy varies between the two groups. In 

Cyathophorella, Cyathophorum, Dendrocyathophorum, and 

Hypopterygium, the leaf cells have well developed pits on 

lateral walls, a feature seen comparatively rarely in members of 

Hookeriaceae. Furthermore, Cyathophorella hookeriana (Griffith) Fleischer and Dendrocyathophorum paradoxum (Brothem) Dixon have distinct foliose pseudoparaphyllia which are 

1 
2 SMITHSO” CONTRIBUTIONS TO BOTANY 

not matched in the Hookeriaceae. The structures that look like 

small pseudopmphyllia in a few Hookeriaceae such as 

Callicostella and Achrophyllum have a different aspect and 

should be examined more carefully. 

As to sporophytic characters, Noguchi (1952) has emphasized and illustrated their similarity in Hypopterygiaceae. For 

example, Cyathophorum and Dendrocyathophorum have a 

cucullate calyptra and a cross-striated, unfurrowed exostome 

with cilia present on the endostome, characters allegedly 

incongruent in Hookeriaceae. The exception is Cyathophorella 

which appears to have a papillose exostome with a median 

zigzag line, reminiscent of the daltoniaceous exostome. In spite 

of the fact that the cyathophorelloid exostome is generally 

papillose, careful microscopic examination shows the dense 

papillae are sometimes borne on low ridges or folds formed on 

the outer surface of the exostome. We tend to interpret this as 

a modified hypopterygioid peristome. 

Furthermore, the outer and inner layers of the exostome in 

Cyathophorum and Cyathophorella are more or less equal in 

thickness, unlike the daltoniaceous or hookeriaceous type of 

exostome described by Allen and Crosby (1986) and Buck and 

Vitt (1986). This is reflected in the fact that the exostomial teeth 

neither bend outward nor curve inward when wet. 

Considering all the evidence at hand, we prefer to keep the 

three cyathophoroid genera in or near the Hypopterygiaceae. 

We also examined under the SEM the papillose exostomes of 

Actinodontium, Daltonia, Distichophyllidium, and Leskeodon 

in an effort to understand their inter-relationship. Much to our 

surprise, the details of their peristomial designs are different 

and can be described in three types based on the size and 

thickening of each of the dorsal and ventral exostomial plates, 

the presence or absence of high trabeculae on both sides of the 

plate, and the alignment and composite nature of surface 

papillae (Figures 35-63). These three types are: the daltonioid, 

actinodontoid (lepidopiloid?), and distichophyllidioid. The 

exostome of Asiatic Leskeodon can be included under the 

daltonioid type. Clearly, this reenforces the view of Buck 

(1987) that it is phylogenetically incorrect to place all 

hookeriaceous ma with papillose peristomes in a single 

separate family. We consider this as evidence that the character 

of exostomial papillosity has arisen independently at least three 

times in the family Hookeriaceae, perhaps more. Future work 

will probably show that papillose peristomes have also arisen at 

least twice in the related family Sematophyllaceae. 

On the other hand, hookeriaceous taxa like Calyptrochaeta, 

Callicostella, and Distichophyllum, with cross-striolate exostomid teeth and with a distinct median gap, are relatively 

uniform in the details of their peristomial designs. The reported 

“striae” are actually distinctive plate-like extensions from the 

outer or dorsal exostomial layer forming a pile or stack between 

the well developed dorsal trabeculae (Figures 35,38). They are 

termed “lamellulae” here for want of a technical term. They 

may or may not be studded with papillae. 

Because the scope of our study is restricted to the Philippine 

species and related taxa, the results do not provide us with an 

adequate basis for a new interpretation of the family organization. We are accepting, therefore, a broadly defined family in 

this treatment. We suspect that the tendency to subdivide the 

family results more from the richness of striking characteristics 

than from excessive phyletic diversity in the family. 

For those who seek a homogenous or natural familial 

classification with a narrower family concept, we suggest that 

a narrower Hookeriaceae may contain only Hookeria. The 

segregated Callicostaceae of Crum (1984) and Buck (1987), at 

its best, would represent those elements with two costae 

distinct from the base, including Bryodusenia H. Robinson. 

The Daltoniaceae, at its best, represents the forms with a single 

costa or a costa branched above the base, including Achrophyllum, Adelothecium, and Bryobrothera. The present authors, do not accept the implication of other studies that the 

forms with single costae are primitive in the group. As 

suggested by Robinson (1975, 1986), all the strongly costate 

forms, both the single and double, are derived from basically 

short and doubly costate forms of the type seen in the related 

families Ptychomniaceae, Leucomiaceae, and Sematophyllaceae. Some of the genera with single costae such as 

Calyptrochaeta and Achrophyllum betray traces of their 

double-costate ancestry by the branching of their costae. 

Most of the genera with the specialized isodiametric median 

leaf cells seem to fall within the daltoniaceous group of the 

family. Nevertheless, the characteristic, in combination with a 

contrasting border of narrow cells, is found in such callicostaceous genera as Cyclodictyon and in scattered genera elsewhere 

among the mosses, including some of the acrocarps like Mnium 

and Tortula mniifolia (Sullivant) Mitten. The character alone 

cannot be considered as evidence for or against close 

relationship. The highly specialized nature of the leaf cells in 

the daltoniaceous series, however, supports the specialized 

nature of the group within the Hookeriales. 

Our revisionary study of Philippine taxa falls within the old 

familial limit circumscribed by Fleischer (1908) and Bartram 

(1939), excluding Chaetomitrium Dozy & Molkenboer and 

Chaetomitriopsis Fleischer. Chaetomitrium, Chaetomitriopsis, 

and Dimorphocladon Dixon (Dixon, 1922) are three related 

genera in the Hookeriaceae, sensu lato, which occupy a 

relatively isolated position. Their discordant features are the 

narrowly oblong to vermiform, often prorulate leaf cells, much 

enlarged perichaetial leaves, and a hypnaceous peristome. We 

agree with Buck (1987) in transfemng these three genera to 

Hypnaceae sensu Nishimura et al. (1984), or at least into some 

family closer to the Hypnaceae. However, in order to account 

for all the taxa treated in the Hookeriaceae by Bartram (1939), 

we include a tentative key to the revised Philippine species of 

Chaetomitrium. 

In 1939, Bartram described some 30 species and 10 genera of 

Philippine Hookeriaceae. In the over 50 years since, additional 

collections made by local and foreign botanists have increased 

the number of known taxa significantly. The present revision 
NUMBER 75 3 

treats a total of 53 species distributed in 12 genera, including 

Chaetomim'um. Eleven new records of Philippine mosses, ten 

new synonyms, five new combinations, and one new name, 

Chaetom'trium schofieldii, are reported. Two ma, Chaetomb 

trium seriatum Brotherus ex Bartram and Eriopus microblastus 

Brotherus, are resurrected from synonymy, and Distichophyllum elmeri Brotherus is resurrected as a variety of D. 

nigricaule. Lectotypes are selected for Calyptrochaeta ramosa, 

Distichophyllum brevicuspes, D. nigricaule, and Hookeriopsis 

wichurae, and for two of the names in synonymy, Distichophyllum gracilicaule and Lepidopilum sumatranum. In 

addition, Daltonia armata is reported new to Sumatra, and, 

Achrophyllum dentatum and Distichophyllum subnigricaule as 

new records for New Guinea. 

Excluding Chaetomitrium, only three species, or about 5%, 

of the Philippine species of Hookeriaceae are locally endemic. 

They are Calyptrochaeta microblasta (Brotherus) Tan & 

Robinson, Distichophyllum santosii Bartram, and Leskeodon 

philippinensis Brotherus. 

The specimens cited in the text are limited to (1) types 

studied, (2) any changes in the published identification of 

specimens, and (3) recent collections not listed in Bartram 

(1939) and which represent new locality records for the 

country. In the case of foreign collection citations, they 

represent significant range extensions in the region. Type 

specimens of several species from adjacent areas were studied 

for possible new synonyms. 

ABBREVIATIONS.-Abbreviations used to indicate the location of specimens examined in this study are as follows: 

B 

BISH 

Botanisches Museum, Berlin-Dahlem, Germany 

Bernice P. Bishoo Museum. Honolulu. Hawaii 

BM British Museum (Natural History) . 

BO Herbarium Bogoriense, Bogor. Indonesia 

BR 

CAHP 

Jardin Botanique de I'Etat, Bruxelles, Belgium 

University of the Phdippines at Los Banos 

FH Farlow Herbarium at Haward University, Cambridge, 

Massachusetts (includes the Bartram and Fleisher HerConservatoire de Jardin botaniques. Geneva, Switzerland 

Botanical Museum, University of Helsinki, Finland 

bariums) 

G 

H 

H-BR Helsinki: Brotherus Herbarium @art of the Botanical 

HIRO Botanical Institute, Hiroshima University, Hiroshima, 

Museum, University of Helsinki, Finland) 

L 

M ICH 

NICH 

NY 

Pc 

PNOC 

U 

us 

Japan 

Rijksherbarium, Leiden, Netherlands 

University of Michigan, Ann Arbor, Michigan 

Hattori Botanical Labratory.Nichinan, Japan 

New Yd Botanical Garden, N.Y., New York 

Labratoire de Cryptogamie. Museum National d'Histoire 

Herbarium of the Philippine National Oil Corporation 

Botanical Museum and Herbarium, Utrecht, Netherlands 

United States National Herbarium, Smithsonian Institution, 

Naturelle, Paris, France 

Washington, D.C. 

ACKNOWLEDGMENTS.-"he first author Would like to 

acknowledge the research fellowship grant given by the 

Smithsonian Institution which enabled him to conduct the 

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

Together, we thank the following curators for the loans of 

specimens: W. Buck (NY), S. Hattori (NICH), A. Touw (L), D. 

pfister 0, P. Isoviita (H-BR), A. Harrington (BM), and W. 

Schultze-Motel (B). We thank Bruce Allen and William Buck 

for the many helpful comments on the manuscript, and we 

appreciate the help of Dan Norris on Bryobrothera. We also 

express appreciation to B. Kahn for helping with the SEM 

photography, B. Eastwood for the printing of negatives, and A. 

Tangerini for the line drawings. E. Judziewicz,E. Farr,D. Bell, 

and other staff members of the Botany Department of the 

Smithsonian Institution helped in one way or another in the 

completion of the research and the preparation of the 

manuscript. 

Family HOOKERIACEAE 

HOOKERJACEAE Schimper, Corollarim Bryologiae Europaeae, 101,1855. 

Key to the Genera of Philippine Hookeriaceae 

1. 

2. 

3. Leaves scarcely complanate, isomorphous, oblong-lanceolate, lanceolate to 

lanceolate-linear ............................. Daltonia 

Leaves complanate, usually dimorphous, ovate-oblong, lingulate, obovate, or 

spathula ....................................... 4 

4. Exostome teeth without median furrow, papillose ........... Leskeudon 

Exostome teeth with median furrow, cross-striolate ................ 5 

5. Leaves without differentiated marginal cells, cells of lamina with strongly 

incrassate walls, sometimes mamillose ............... Bryobrotheru 

Leaves usually with elongate marginal cells, cells of lamina with thin or scarcely 

thickened walls, not mamillose ................. Distichophyllum 

Leaf margins strongly erose or irregularly dentate ......... Achrophyllum 

Leaf margins not strongly emse or dentate .................... 2 

Costa single, reaching at least half the leaf length ................. 3 

Costae double, branched above the base, short, or lacking ............ 6 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

6. 

7. 

8. 

9. 

10. 

11. 

12. 

Costae strong, reaching at least half the leaf length ................ 7 

Costae weak, short,or none. ........................... 10 

Leaf borders distinctly differentiated; leaf cells round to hexagonal, large, more than 

30 pm in diameter. ......................... Cyclodictyon 

Leaf borders not or weakly differentiated; leaf cells oval, rhomboidal to elongate, 

longer than wide, less than 30 pm wide. .................... 8 

Exostome teeth papillose ...................... Actinodontium 

Exostome teeth cross-striolate ........................... 9 

Costae reaching near the apex, strongly protruding abaxially ..... Callicostella 

Costae reaching only half the leaf length or less, not or weakly protruding abaxially 

.................................... Hookerwpsis 

Leaf cells broadly oval-rhomboidal, large, more than 18 p wide, very thin-walled 

...................................... Hookeria 

Leaf cells narrowly rhomboidal, oblong, fusiform, elongate, or vermiform, less than 

18 pm wide, firm-walled ............................ 11 

Leaf borders distinctly differentiated; setae strongly spinose or hirsute ...... 

Leaf borders not or weakly differentiated; setae smooth or papillose, at most 

scabrous ..................................... 12 

Leaves strongly divergent or deflexed ............. (Chaetomitriopsis) 

Leaves imbricate, complanate, at most erect-spreading ..... (Chaetomifrium) 

................................... Calyptrochaeta 

Genus Achrophyllum 

Achrophyllwn Via & Crosby, Bryologist, 75174, 1972. [w: Hookeria 

quodrqaria J.E. Smith (= Achrophyllwn quadruariwn (J.E. Smith) Vitt & 

Crosby).] 

The name, Pterygophyllum Bridel, is invalid having originally included the type species of the genus Hookeria. The 

name has been replaced by Achrophyllum (Crosby, 1972; Vitt 

and Crosby, 1972). 

The main distribution centers of the genus are in the 

Southern Hemisphere with two species reaching Malesia: A. 

javense is widespread, while A. dentatum (Hooker f. & Wilson) 

Vitt & Crosby is reported here for New Guinea. 

NOTE ON Pterygophyllum novae-guineae 

Bartram (1961) described Pterygophyllum novae-guineae as 

a much more robust plant than P. dentatum, having longer 

leaves that are coarsely toothed above with multicellular teeth. 

However, the plant size and the leaf dentation of Achrophyllum 

dentatum are variable features noted by Sainsbury (1955) in 

New Zealand populations. In addition, our own examination of 

herbarium specimens of Achrophyllum dentatum from New 

Zealand, Australia, and Chile further shows that the length of 

the leaf costa and the shape of the leaves are equally variable. 

We think that the two species are best treated as synonyms. 

Thus, A. dentatum is reported here as new to New Guinea (= 

Pterygophyllum novae-guineae, syn. nov.). 

Achrophyllumjavense is also known from New Guinea. It 

differs from A. dentatum in having an acute to acuminate leaf 

apex and a much more strongly erose leaf margin. In A. 

dentatum, the laminal cells are 45-60 pm in diameter, the leaf 

apices are mostly obtuse or rounded and the cells of the 

marginal teeth are 30-36(-60) p in diameter. 

SPECIMENS STUDIED.-PAPUA NEW GUINEA. Morobe District: Mt. Kaindi, vicinity of Wau, Weber & McVean B-34871 

(US, as Pterygophyllum novae-guineae). Western Highland: 

Wabag area, Sugarloaf Mt., Robbins 2828 (holotype of 

Pterygophyllum novae-guineae, FH); ibid, Hoogland & 

Schodde 7225 0. 

Achrophyllum javense 

Achrophyllwn javense (Dixon ex Froehlich) Iwatsuki, Tan, & Touw, J. Hattori 

Bot. Lab, 44149,1978. 

Pferygophyliwn javense Dixm ex Frcehlich. Ann. Natuh. Mus. Wien, 5999, 

1953. [Type: Java, im Urwalde Oben Tjibodas, leg. Schifier (Clyptogam. 

Exsicc. Museo Hist. Natur. Vindobonensi 4372, isotype, US I).] 

Plants tufted, sparingly branched. Stems to 6 cm tall and -3 

mm wide including the leaves. Stem cross-section with a layer 

of hyalodermis surrounding the thickened outer cortical cells, 

and a small central strand. Leaves moderately cornplanate, 

strongly contorted to crisped when dry, variable in size and 

shape. Lateral leaves mostly oblanceolate, occasionally ovate 

and lanceolate. Dorsal and ventral leaves ovate to oblong. Leaf 

apices acute to acuminate; unbordered leaf margins strongly 

and irregularly toothed, with several cells forming each of the 

tooth segments measuring to 75 wm long. Costa single, short, 

often branched above the base. Leaf cells round-hexagonal, 

36-40 pm in diameter, smooth, thin-walled, slightly thickened 

at corners, becoming rectangular and elongate at base. 

Rhizoidal propagules formed from the leaf cells. Sporophytes 

not seen. 
NUMBER 75 5 

SPECIMENS STUDIED.-PHILIPPINES. Luzon: Mt. Pulog, 

Jacob B 1% (B, L, NY); Mt. Data, Nishimuru 6725 (HIRO, 

US). Mindoro: Mt. Halcon, Tan 87-095, 87-185 (CAHP). 

Mindanao: Mt. Apo, Tun 82-232, 82-330 (CAHP); Mt. 

Hilong-Hilong, Cabadbaran, Tan 84403,84436 (CAHP). 

REMARKS-T~~S is a species of humid and humic habitats in 

mossy forests. It was first reported for the country by Iwatsuki 

and Sharp (1968) from Mt. Data in northern Luzon. It is easily 

identified by its filmy leaves that have large leaf cells and an 

erose marginal dentation. 

Genus Actinodontium 

Actinodontiwn Schwaegrichen, Spec. Musc. Suppl, 2:75, 1826. "&pe: A. 

ascendens Schwaegrichen.] 

This is a genus of small, epiphytic plants, very much like 

Dultonia in general habit. The leaves are silky yellow-green or 

dark-green, and densely cover the stem. They are doubly 

costate, but the costae do not protrude abaxially. The stem 

cross-section consists of a weakly differentiated epidermis 

enclosing a group of parenchyma cells. Often, a cluster of 

brown, thin-walled stem epidermal cells grows outward from 

the leaf insertion forming a pseudo-alar structure. The calyptra 

is mitrate and the base is laciniate instead of fiinged with cilia. 

The erect capsule is without collenchymatous exothecial cells. 

Two species have been reported from the Philippines; but 

one,A. ascendens, lacks a definitive herbarium record. 

Key to the Philippine Species of Actinodontium: 

Leaves ovate to lanceolate, the apices acute to short acuminate; capsules ovoid-oblong, 

less than 2 mm long. ........................... A. ascendens 

Leaves narrowly lanceolate, the apices long acuminate; capsules oblong-cylindric, more 

than 2 mm long ........................... A. rhaphidostegum 

Actinodonlium ascendens 

FIGURES 47-50 

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

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

REMARKS.-Tixier (1978) reported this taxon as new to the 

Philippines based on a collection he ma& from the Benguet 

Highlands in northern Luzon. In the same article, he disputed 

Bartram's (1939) Philippine report of A. rhuphidosfegum by 

remarking that the latter is a very small species found in shady 

forest habitats. All of the herbarium specimens of Actinodontium from the Benguet Highlands seen by us are A. 

raphidostegum, which grows as an epiphyte on the trunks of 

various vascular plant species in semi-open to openly disturbed 

environments such as roadsides and lawns. Efforts to relocate 

Tixier's collection have so far failed. An earlier record of A. 

ascendens in the Philippines (Williams, 1914) was corrected to 

A. rhaphidostegum by Barttam (1939). 

Based on our field records and examination of herbarium 

specimens, the two species are actually of the same size range 

and both grow epiphytically on tree trunks in both primary and 

disturbed forests. 

Actinodontium ascendens can be distinguished by its broadly 

acute leaf apices in wet condition, and also by the ovoid 

capsules. A good illustration of it can be found in Fleischer 

(1908,fig. 176). It is not uncommon in the Malesian region and 

is also known from Sri Lanka, Thailand, and Java. It is 

probably present in the Philippines, but because we have not 

Seen a Philippine specimen, we remain uncertain about its 

presence. 

Actinodontium rhaphidostegum 

Actinodontiwn rhaphidosiegwn (C. Muller) Bosch & Lacoste, Bryol. Jav., 

Hookeria raphiabsiega C. Muller, Syn., 2:677, 1851. [Type: Java, Blme s.n. 

237,1862. 

(B, destroyed).] 

Plants small, less than 2 cm tall, forming scanty tufts on 

trunks of trees, Pundunus, and bamboo culms. Stems simple or 

branched, densely covered by somewhat flexuose leaves, often 

with a purplish tinge. Leaves 2-2.5 mm long, shiny green, 

slightly shrunken when dry, erect-spreading when wet, oblonglanceolate, gradually acuminate into a narrow point, the 

margins recurved, weakly semlate near apex, otherwise entire. 

Costae double, long, more than half the length of the lamina. 

haf cells elongate-rhomboidal, (45-)51-135 pm long and 

12-21 pn wide, thin-walled, becoming short-oval to hexagonal toward apex, narrower near margin, more rectangular and 

laxer at base. Perichaetial leaves small, lanceolate, acute to 

obtuse, ecostate. Perigonial leaves similar. Calyptra mitrate, 

laciniate at base. Setae short, about 5 mm long. Operculum 

rostrate. Capsules oblong-cylindric, to 3 mm long. Exothecial 

cells not collenchymatous, but thickened along the vertical 

walls. Exostome teeth papillose, with a distinct median zigzag 

line, opening outward when wet. Spores green, papillose. 

SPECIMENS STUDIED.--~~ILIPPINES. Luzon: Mt. Santo 

Tomas, Tun s.n., 29-30 Nov, 1986 (CAHP); Baguio, MSAT 

campus, Onruedt 84.P.10783 (CAHP); Imugan, Nueva Viscaya, Tun 84-13 (CAHP); Mt. Banahao, Iwutsuki & Sharp 

16453, (NICH, CAHP). Mindoro: Mt. Halcon, Tun 87-119 

(CAHP). Mindanao: Mt. Apo, Tun 82-261 (CAHP). 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

REMARKS.-This species is variable in the length of its leaf 

cells, leaf apices and capsules. Indeed, in some herbarium 

specimens the leaf apices are intermediate between acute and 

acuminate. The illustration of A. uscendens in Dozy and 

Molkenboer's Bryologiu Juvunicu (1855-70) clearly shows a 

variation of leaf apices, ranging from broadly acute to short 

acuminate, like that of A. rhuphidostegum. The two taxa may 

eventually prove to be synonymous. 

Actinodontium rhuphidostegum is widespread in the Malesian region. 

Genus Bryobrothera 

Btyobroihera Theriot,Rev. Bryol., 47:26,1920. [Type: B. crenulata (Brothems 

& Paris) Theriot.] 

The position of this monotypic genus has been shown to be 

with the Hookeriaceae by Norris and Robinson (1979). The 

latter authors have shown the geographical range to include 

Queensland, Australia, Fiji, the Solomon Islands, and the type 

locality, New Caledonia. The Philippine record is reported by 

Tan and Koponen (1989). 

Bryobrothera crenulata 

Bryobrothera crenulata (Brothems & Pans) Thbriot, Rev. Bryol.. 47:26,1920. 

Mesochaeie crendaia Brothems & Pans, Oefv. Finsk. Vet.-&. Foerh., 51:19, 

1908. [m: New Caledonia, Mt. Dzumac, Le Rat s.n. (H), not seen.] 

Calomnion dixonii Bartram,Bishop Mus. Occ. Pap., 11:9,1936. [Types: Fiji, 

Viti Lvu, near summit of Mt. Victoria, Gillespie 4114 (holotype, BISH; 

isotype. FH). not seen.] 

Plants small, mat-forming. Stems prostrate, complanatefoliate. Lateral leaves elliptical to oblong, 0.5-1.5 mm long, 

somewhat asymmetrical. Median and ventral leaves smaller, 

broadly oblong to obovate, symmetrical. Leaf apices broadly 

acute to obtuse; margins crenulate to weakly toothed above, 

entire below, without elongate cells. Costa single, percurrent or 

ending below the apex. Leaf cells mostly isodiametric, 10-12 

pm, with thick walls, smooth to slightly mamillose. Plants 

dioicous. Calyptra mitrate, plicate toward base, sparingly 

pilose. Setae 3-4 mm long, smooth. Operculum rostrate. 

Capsule horizontal to pendent, 0.8-1.2 mm long. Exothecial 

cells collenchymatous. Exostome teeth cross-striate with a 

distinct longitudinal median furrow. 

SPECIMENS STUDIED.-PHILIPPINES. Mindanao: Mt. HilongHilong, Cabadbaran, Agusan Norte Province, Tun & Nuvurez 

84-520 (CAW, H). 

Rmms.-The complanate and matted habit is rather 

distinctive for this species. In addition, the strong single costa 

and the unbordered leaves distinguish the species from most 

other members of the family. Because of the latter combination 

of characters, the species was placed in other families such as 

the Calomniaceae and Rhizogoniaceae before sporophytes 

were discovered (Noms and Robinson, 1979). Sporophytes are 

still known only from the Noms Queensland collection. 

Recently, Akiyama (1988) reported Culomniom complunutum (Hooker f. & Wilson) Lindberg, a New Zealand 

endemic, from the island of Seram (Ceram) in Indonesia. Both 

the illustrations by Akiyama and the geography favor identification of the material as Bryobrotheru. Bryobrotheru has been 

mistaken for Culomnion before, and the two are similar in the 

strong costae and unbordered leaves. Nevertheless, the leaves 

of C. complunutum are markedly 3-ranked, its lateral leaves are 

oblong-lanceolate with sharply acute to shortly acuminate 

apices, and sporophytes, when present, are terminal. The leaves 

of B. crenulutu, on the other hand, are not 3-ranked, the lateral 

leaves are oblong-elliptical, never lanceolate, its leaf apices are 

mostly obtuse, and sporophytes, when present, are lateral. 

Genus Callicostella 

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

1859. [w: C. papillata (Montagne) Mitten.] 

The generic name Cullicostellu has been shown to be a later 

synonym by Crosby (1975) who replaced it with an earlier 

name, Schizomirrium Bruch, Schimper, & Gumbel. However, 

the name Cullicostellu, which has long been in use, was 

proposed for conservation by Koponen and Isoviita (1984). The 

proposal was approved at the 1987 Berlin Congress. 

The genus is characterized by the presence of two strong and 

abaxially protruding leaf costae. The calyptra is notable in 

being large, covering the entire capsule, strongly plicate and 

laciniate at base, but without a hairy basal fringe. The calyptra 

proper is only sparsely hairy, being scabrous near the tip. The 

exostome teeth are typically hookeriaceous with a cross-striate 

pattern and a distinct median furrow. 

There are two known Philippine species of Cullicostellu in 

this largely neotropical genus. Buck (1987) placed Cullicostellu with Cullicostu and other Hookeriaceae having a double 

costa in the family Cullicostuceue of Crum (1984). 

Key to the Philippine Species of Callicostella 

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

Leaf cells smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. prabaktiana 
NUMBER 75 7 

Callicostella papillata 

Callicostella papillata (Montagne) Mitten, J. Linn. Soc. Bot., Suppl. 1:136, 

Hookeria papillata Montagne, hdon J. BOL, 3532,1844. [Type: Java, Prov. 

1859. 

Buitenzorg, Miquel s.n. (U), not seen.] 

Plants mat-forming, light yellow to dark green in color. Main 

stems long, creeping, pinnately to subpinnately branched, the 

secondary branches 2-5 cm long and less than 1 mm wide 

including leaves. Stem cross-section without a central strand. 

Leaves complanate, somewhat flexuose, two strong costae 

forming two plicae on the leaf blade when dry, ovate to 

oblong-lanceolate, 1 .O- 1.5 mm long, broadly acute to abruptly 

acuminate. Leaf costae reaching near the apex, protruding 

abaxially, the costal surfaces smooth to strongly toothed near 

the distal end, at times lamellate. Leaf margins plain, without a 

clearly differentiated border, irregularly toothed to lower half, 

at times serrulate. Leaf cells hexagonal-oval, shortly oblong to 

rhomboidal, 5-15 pm long and 6-9 pn wide, angular, thin to 

thick-walled, but not pitted, mostly with a single papilla over 

the lumen, cells elongate and smooth near the base; the 

marginal leaf cells shorter, projecting apically. Perichaetial 

leaves mostly lanceolate, slightly smaller than the vegetative 

leaves; the margins serrate. Synoicous or autoicous. Setae 

smooth below, weakly papillose above, 1-2 cm long. Capsules 

ovoidcylindric, horizontal to inclined. Exothecial cells collenchymatous. Exostomes cross-striate with a large median 

furrow. 

SPECIMENS STUDIED.-PHILIPPINES. Luzon: Lama0 River, 

Bataan Prov., Williams 852 (US); National Botanic Garden, 

Real, Quezon Prov., Tan 81-455 (CAHP); Tignaong, Tayabas 

Prov., Leiberg 1230 (US). Panay: Jamindan, Capiz Prov., 

Rams & Edano (Bur. Sci.) 31226, 31090 (US). Mindanao: 

District of Zamboanga, Merrill8359 (US); TIMEX logging area 

west of Ipil, Mt. Silligain, Zamboanga Prov., Hale & Banaag 

25306 (US). 

Rmms.-Although this species is noted for its polymorphism, few have described the observed variations in detail. In 

general, the ventral leaves, which are in direct contact with the 

moist substrate, tend to be more lanceolate and long-acuminate 

than the dorsal and lateral leaves. Also, the cells of the ventral 

leaves become more rhomboidal-oblong in outline with only 

thin walls. Often the single papilla over the cell lumen is 

weakly developed and may be absent from the cells of many 

ventral leaves. Plants having only a few papillose leaf cells are 

often incorrectly named C. prabaktiana. The marginal teeth of 

the leaves are equally variable. Each tooth segment may be 

formed by one to many cells, often becoming bigeminate to 

trigeminate apically. The abaxial surface of the leaf costa can 

vary from smooth to spinose or even lamellate. These variable 

characters appear to be unreliable bases for specific as well as 

infra-specific classification. 

As presently interpreted, C. papillata is widespread in the 

paleotropics. It is very common in Philippine lowland 

rainforests. 

Callicostella prabaktiana 

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

1862. 

Hookerio prabakiaM C. Muller, Syn., 2678,1851. [Type: Java, Mt. Prabakti, 

Zollinger 3503 (B), destroyed.] 

Plants like C. papillata in general habit. The leaf cells more 

oval-oblong, smooth or slightly mamillose, with thick walls. 

SPECIMENS STUDIED.-PHILIPPINES. Luzon: Bicol National 

Park, Camarines Sur Prov., Tan 74-413 (US, CAHP). Polillo: 

Leiberg 1224 (US). 

REMARKS.-Contrary to the claim of Bartram (1939), this is 

actually a rare or uncommon species in the Philippines. We 

have seen only three undoubted collections from the region. 

Except for the smooth leaf cells, it resembles the preceding 

species in most details. As in C. papillafa, the leaf shapes, 

marginal dentation of the leaf, and the abaxial surface 

ornamentation of leaf costae of C. prabaktiana are variable 

characters. 

One Philippine specimen of C. prabaktiana with lamellate 

leaf costae (Leiberg 1224, US) was named C. beccariana 

(Hampe) Jaeger by Brotherus. However, this plant has some 

leaves with smooth to spinose costae. 

Most specimens of C. prabaktiana have been collected from 

very wet environments, a condition that appears to cause the 

partial disappearance of leaf papillae in C. papillata. When 

more collections of these two species are studied and analyzed 

in the light of a better understanding of the environmental 

effects on the morphogenesis of moss plant organs, specially 

the leaf papilla formation, the two may prove to be conspecific. 

Callicostella prabaktiana is known also from Java, Borneo, 

peninsular Malaya, and Indochina. 

Genus Calyptrochaeta 

Calyptrochaeta Desvaux, Mem. Soc Linn. Paris, 3:226,1825. [?).pe: Lskea 

cristata Hedwig (= Calyptrochaeta cristata (Hedwig) Desvaux).] 

The generic name Eriopus Bride1 has been found to be 

illegitimate (Margadant 1959; Crosby 1974), and has been 

replaced by Calyptrochaeta. The genus is characterized by a 

complanate and heterophyllous foliation, often with attenuated 

branches bearing numerous filiform propagules in the leaf 

axils, and a spinose or hirsute seta. 

In the Philippines, species concepts have traditionally been 

based on variation in a few gametophytic characters such as the 

leaf border, leaf cell morphology, and the leaf outline. As the 

plants infrequently produce sporophytes, reproduction usually 

occurs by means of the asexual propagules on the stems. It is 

possible that the various species treated here may represent 

clonally perpetuated microspecies with a fixed suite of 

characters. Until breeding studies and transplanting experiments are conducted for the genus, we are accepting the 

narrowly defined species concepts proposed by various 

authors. 
8 SMlTHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 1-10.4alyprrochaeta. 1,2, C. parvireh (Java: Gedeh, Flehcher, type, FH): 1, upper leaf cells; 2. 

whole leaf. 3,4, C. rams (Philippines: Luwn, Mt. Banahao, Merrill7523, type of EriopusfIoccidus, FH): 3, 

upper leaf cells; 4, whole leaf. 5.6. C. remot&dia (Java: Tjibumum,Fleischer, FH): 5, upper leaf cells; 6, whole 

leaf. 7-10, C. microMarta (philippines:Sibuyan Is., Mt. Giting-Giting. Tan & Hernaez 87-447, US): 7, upper leaf 

cells; 8, whole leaf; 9, plant, dry; 10. plant, wet. 
NUMBER 75 

Key to the Philippine Species of Calyptrochaeta 

1. Plants large, more than 4 cm long; lateral leaves oblong-lanceolate, flaccid and 

Plants small to medium size, less than 4 cm long; lateral leaves ovate-elliptic, firm or 

slightly wrinkled when dry. .......................... .4 

2. Upper leaf cells mostly oblong, elliptic to fusiform, ratio of length to width (3-5:1), 

frequently thin-walled; leaf borders, especially near the apex, consisting of (2)3-4 

rows of elongate-linear cells; perichaetial leaves with blunt to short acute apices . ................................... C. remotifolia 

Upper leaf cells mostly oval, rhomboidal, or hexagonal, ratio of length to width 

(1-3:1), thin to moderately thick-walled with thickened corners; leaf borders, 

especially near apex, consisting of 2(-3) rows of elongate-linear cells; perichaetial 

leaves with acuminate or abruptly mucronate apices .............. 3 

3. Upper leaf cells mostly hexagonal to oval, often collenchymatous; marginal teeth of 

leaf weak to moderately strong .................... C. parviretis 

Upper leaf cells mostly rhomboidal, not collenchymatous; marginal teeth of leaf 

strong to very strong. ......................... C. ramosa 

4. Lateral leaves ovate to broadly elliptic; upper leaf cells collenchymatous; the upper 

leaf borders at most with 2 rows of elongated cells; filiform propagules borne in 

leafaxils ............................... C. parviretis 

Lateral leaves mostly obovate-elliptic to broadly oblanceolate; upper leaf cells not 

collenchymatous; upper leaf border with (3-)4 rows of linear cells; filiform 

propagules borne terminally on attenuate branches with small, modified leaves . ................................... C. microblasta 

strongly wrinkled when dry. ......................... .2 

9 

Calyptrochaeta microblasta (Brotherus) Tan & Robinson, 

new combination 

FIGURES 7-10 

Enopus microblastus Brotherus, Philip. J. Sci., 8C (Bot.):82,1913, basionym. 

[Ws: Philippines, Luwn, Mt. Banahao, Robinson Bw. Sci. 9813 

(holotype, H-BR!; isotypes. FH!, NY!).] 

Plants small to medium size, tuft forming. Stems mostly 

branched at base, less than 4 cm tall, laxly foliate. Attenuate 

propaguliferous branches often naked or with very small 

leaves. Leaves cornplanate, slightly wrinkled when dry. Lateral 

leaves obovate to elliptic, slightly asymmetrical at base, 

strongly cuspidate, 2-3 mm long and 1-1.5 mm wide, the 

margins serrate to serrulate, with 3-4 rows of linear cells all 

around. Costae double, short. Laminal cells broadly rhomboidal to hexagonal, becoming laxer and rectangular near base. 

Dorsal and ventral leaves variable in shape, ovate to lanceolate, 

smaller in size; areolation and marginal serration similar to 

lateral leaves. Perichaetial leaves much reduced in size, with 

long-acuminate apices. Setae mostly appearing terminal on 

branches, 4-6 mm long, very spinose. Operculum conic. 

Capsules ovoid, 1 mm long. 

SPECIMENS STUDIED.-PHILIPPINES. Luzon: Mt Babatuging, Cagayan Prov., Edano (Bur. Sci.) 3335, det. by Dixon as 

Eriopuspropagulifer, in scheda (US,FH). Sibuyan Island: Mt. 

Giting-Giting, Tan & Hernaer 87-447 (CAHP, US). Panay 

Island Antique Prov., Mt. Madyaas, M. Price s.n., Jan 15-16, 

1987 (CAHP). Negros: Cuernos Mt., Palmore 2843 0; ibid, 

Magdamo 23 0. Mindanao: Mt. Hilong-Hilong, Cabadbaran, Agusan Prov., Tan & Navaret 84-471, proparte (CAHP). 

REMms.-Calyptrochaeta microblasta was considered a 

synonym of C. parviretis by Bartram (1939). The species, 

however, can be distinguished from all other Philippine species 

by the following characters: (1) plant size small, with a lax 

foliation and a firm leaf texture that gives the plant the aspect 

of a species of Epipterygium; (2) smng leaf borders, especially 

near the apex, consisting of 3-4 rows of linear cells; (3) short 

upper leaf cells as in C. parviretis; (4) leaf apices to 450 p 

long, consistently longer than all other species of Philippine 

Calyptrochaeta. 

In addition, the presence of a nearly naked mass of filiform 

propagules borne terminally on attenuated branches is unusual 

and is known only from a few other taxa in the family (Allen, 

1987b). 

Calyptrochaeta microblasta is a Philippine endemic (Brothem, 1913). It has been collected from several islands in the 

country, mostly from limestone areas. 

Calyptrochaeta parviretis 

FIGURES 1,2 

Cafyptrochuetu parvirerir (Fleischer) Iwatsuki. Tan & Touw, J. Hattori Bot. 

Eriopus purviretir Fleischer, Musci Fl. Buitenmrg, 3:1008, 1908. [Type: West 

Lab.,44:150, 1978. 
10 SMlTHSONIAN CONTRIBUTIONS M BOTANY 

Java, Pangerango, Kandang Badak, Fleircher s.n. (FH).] 

Plants large, tuft-forming. Main stem long, often creeping, 

secondary stems complanately foliate, 4-5 cm tall, erect or 

inclined, branched, often attentuate at tip with bundles of 

brown, fiiform propagules borne in the leaf axils. Stem 

cross-section with weakly differentiated central strand. Lateral 

leaves wrinkled when dry, at times flat, broadly ovate to 

elliptic, asymmetrical, 3 mm long and 2 mm wide, cuspidate or 

shortly acuminate. Costae double, very short. Leaf border 

distinct, toothed near apex, consisting of mostly 2 rows of 

elongate cells. Upper leaf cells mostly oval to short rhomboidal, 30-45(75) pn long and 12-18 pn wide, thickened at 

corners, becoming laxer and rectangular near base. Dorsal and 

ventral leaves smaller, mostly broadly ovate, the leaf border 

less sermlate. Dioicous. Perichaetial leaves small, lanceolate, 

about 1 mm long, acuminate, ecostate. Calyptra hairy, laciniate 

at base. Seta 5-6 mm long, spinose. Capsule erect or inclined, 

ovoid. Exostome teeth cross-striate, with median furrow. 

SPECIMENS STUDIED.-PHILIPPINES. Mindoro: Mt. Halcon, 

Tan 87-447 (CAHP). INDONESIA. Celebes: Soputan Mt.,AZston 

15921 (FH). 

REMMS.-This species is often confused with C. remotifoZia. The latter is a bigger plant with leafy stems more than 4 mm 

wide. The upper leaf cells of C. parviretis are shorter, often 

collenchymatous, and the leaf border consists of 2(3) rows of 

elongated cells. Calyptrochaeta parviretis can also be difficult 

to separate from C. ramosa. The latter, however, is a large plant 

similar in size to C. remotijioiia. 

Calyptrochaeta parviretis is a widespread species in the 

Malesian region. 

Calyptrochaeta ramosa (Fleischer) Tan & Robinson, new 

combination 

FIGURES 3.4.35-38 

Erwpus ramosw Fleischer, Musci F1. Buitenzorg. 3:1007, 1908, basionym. 

[Types: West Java, Tjibeureum, am Gedeh, Fleischer m. (lectotype, with 

spowphyte. FH!; isolectotype H-BR!); am Kandang-Badak, Fleircher s.n. 

(syntype. without sporophyte, FH!).] 

Cyathophorum Iimbatnlum Renauld & Cardot, Rev. Bryol., 23:108,1896, syn. 

nw. [?Lpe: Java, Tjiboda,Massart s.n., 1856 (isotyp. FHI).] 

EriopuspOccidus Brother, philipp. I. Sci., 13C (Bot.):213, 1918, syn. nov. 

[Types: Philippines, Luzon, Mt. Banahao, Merrill7523 (holotype, H-BR!; 

isotypes, NY!, FH!).] 

Plants as large or larger than C. remotifolia, some secondary 

stems to 10 cm tall and 4-5 mm wide. Leaf shape and 

areolation like C. remotifolia, except the border with only 

2(-3) rows of elongate cells and the perichaetial leaves with 

acuminate apices. Perigonial leaves like the perichaetial leaves, 

but with the marginal borders less defined. Sporophyte as in C. 

parviretis. 

SPECIMENS STUDIED.-PHILIPPINES. Luzon: Mt. Pulog, Tan 

82-149 (CAHP); Mt. Banahao,Merrill 7523 0. 

REMARKS.-T~~S species apparently differs from C. remotifolia in having narrower leaf borders and acuminate perichaetial leaves. The upper leaf cells, measuring 36-90 pn long and 

15-24 pn wide, are intermediate between those of C. 

parviretis and C. remotifolia. In the protologue, Fleischer 

(1908) compared C. ramosa to C. remotifolia. However, it is 

more similar to C. parviretis, both in terms of leaf areolation 

and the acuminate perichaetial leaf morphology. 

The illustration and type description of Eriopus spinosus 

(Noguchi, 1937) from Taiwan is probably a juvenile form of C. 

ramosa. 

Eriopusflaccidus was described by Brotherus (1918) as a 

more slender plant with flaccid leaves and shortly spinose setae 

in comparison with C. ramosa. However, Bartram (1939) 

compared it to C. remotifolia and stated that the differences 

between the two taxa are hardly characters of taxonomic 

importance for the genus. We concur with his observation that 

the name is a synonym, but we consider it to be a synonym of 

C. ramosa. Eriopus flaccidus and Calyptrochaeta ramosa both 

have similar narrow leaf borders consisting of 2(-3) rows of 

elongate cells, and both have acuminate perichaetial leaves. 

The type specimen of Cyathophorum limbatulum was 

studied. It is a juvenile plant of Calyptrochaeta ramosa. The 

characters used by Fleischer (1908) in maintaining the species 

are untenable. 

This formerly Javan species is here reported as new to the 

Philippines. 

Calyptrochaeta remot(fo1ia 

Row S,6 

Calyptrochaeta remotifolia (C. Miiller) Iwatsuki, Tan, & Touw, J. Hattori Bot. 

Erwpus remotifoliw C. Miiller, BOL &it.. 5:828,1847. [l)pe: Java, Mt. GM6, 

Lab., 44:150,1978. 

Reinward s.n. (B, destroyed; L?).] 

Plants large, as in C. ramosa, secondary stems to 5-6 mm 

tall. LateraI leaves oblong to oblanceolate, asymmetrical, 

wrinkled when dry. Leaf margins strongly toothed, with (3)4 

rows of linear cells near the apex. Costae double, short. Upper 

leaf cells rhomboidal to fusiform, (90)105-12q150) pm long 

and 21-30 pm wide. Dorsal and ventral leaves variable, mostly 

lanceolate or elliptic, the marginal teeth less strong. Perichaetial leaves with blunt or round apices, occasionally shortly 

acute. Calyptra and capsule as in C. parviretis. 

SPECIMENS STUDIED.-PHILIPPINES. Luzon: Mt. Pulog, Tan 

75-154 (CAHP); Mt Banahao, Tan 84-99 (CAHP). Negros: Mt. 

Canlaon, Iwatsuki & Sharp (NICH 263395). INDONESIA. 

Sumatra: Brash@, Holttum 15488 (US). JAVA. Tjiburnum, 

Fleischer 0. 

REMARKs.-Among its local congeners, this species and C. 

parviretis are commonly seen in mossy forests from mid- to 

high altitudes (-500-2500 m). Past circumscriptions of the two 

species have emphasized the size and shape of the upper leaf 

cells, which vary greatly in lateral, dorsal, and ventral leaves 

from the same plant. The use of the leaf cell size should be 

confined only to mature lateral leaves. We found the width of 

the leaf border also a reliable diagnostic character. The 
NUMBER 75 11 

perichaetial leaf apices of both taxa which were nicely 

illustrated by Fleischer (1908), are another useful character to 

distinguish the two taxa. 

Calyptrochaeta remotifolia may prove difficult to separate 

from C. ramosa. The latter has, in general, shorter leaf cells in 

the upper half of the leaf and narrower leaf borders. Also, the 

perichaetial leaves of C. ramosa have acuminate apices, which 

are unlike the blunt perichaetial leaves of C. remotifolia. 

Genus Cyclodictyon 

Cyclodictyon Mitten, J. Linn. Soc. Bot., 7:163, 1864. [w: C. hctevirens 

(Hooker & Taylor) Mitten.] 

This genus is distinctive in having strong, double costae, a 

clearly differentiated leaf border, and lax leaf cells that are 

large, round to hexagonal, and smooth. It is largely a 

neotropical taxon with only one species represented in the 

Philippines. 

Cyclodictyon blumeanum 

Cyclodutyon blwneanwn (C. Muller) 0. Kuntze,Rev. Gen. R., 2835,1891. 

Hookeria bhmana C. Muller, Syn.. 2:676,1851. [Type: Java, Blwne s.n. (B, 

destroyed).] 

Plants mat-forming, stems creeping, irregularly branched, 

branches to 1.5 cm long and 1.5-2 mm wide including the 

leaves, light green in color. Leaves complanately arranged, 

slightly flexuose when dry, ovate to oblong-lanceolate, 1-1.5 

mm long and about 1 mm wide, broadly acute to acuminate. 

The leaf margins plain, entire below, slightly toothed above 

near the apex. The leaf borders narrow, with 2 rows of linear 

cells. Leaf cells large, (36-)45-60 pm wide, smooth, 

round-hexagonal above, becoming rectangular below near 

base. Autoicous. Calyptra naked, mitrate, laciniate at base. 

Setae to 2 cm long. Capsules ovoid-oblong, horizontal. 

Exostome teeth transversely striolate, with a wide median gap. 

SPECIMENS STUDIED.-PHILIPPINES. Lumn: Mt. Data,Nishimura 6721 (HIRO, US); Mt. Makiling, Nishimura 6967 

(HIRO, US). 

Rmms.-Cyclodictyon blumeanum is a widespread species in the Philippines prefemng humid and shaded places such 

as creek banks and wet cliff faces. Under an ordinary lox field 

lens, the flexuose leaves (in dry state) exhibit the soft texture 

and also conspicuous net-like leaf cell areolation also found in 

Hookeria and Leskeodon. The distinctive double costae 

forming two deep plicae on the leaf blade will distinguish this 

taxon from Hookeria or Leskeodon. A beautiful illustration of 

this species can be found in Mohamed and DamanhuriMohamed (1986). 

This species is widespread in Southeast Asia and the Pacific 

Islands. 

Genus Daltonia 

Daltonia Hooker & Taylor, Musd Brit., 80, 1818, nom. cons. [?Lpe: D. 

splachnoides (J.E. Smith) Hooker & Taylor.] 

Daltonia is a conserved generic name with a complicated 

nomenclatural'history (Crosby, 1%8). Having bordered leaves 

with a single costa, it bears a resemblance to Distichophyllum. 

However, the two genera are different in the degree of 

homogeneity of leaf cells as seen from the apex to the base. In 

Daltonia, only the proximal basal leaf cells are different in 

shape and larger in size than the rest of the laminal cells, while 

in Distichophyllum, the lower half of the leaf consists of 

enlarged rectangular-elongate cells that are morphologically 

different from the round-hexagonal upper leaf cells. Overall, 

the leaves of Daltonia also are narrower and more erectspreading than the leaves of Distichophyllum. 

In addition, the exostome teeth on the outer surface of 

Daltonia are papillose while those of Distichophyllum are 

strongly cross-striated. 

Daltonia, like Actinodontiwn, develops a bulging group of 

colored, thin-walled, pseudo-alar cells at the leaf insertion that 

are of stem origin. This feature is not well developed in 

Distichophyllum. 

The New World members of the genus have been revised by 

Bartram (1931) who considered the perigoniaVperichaetial 

leaves, the peristome, the operculum, and the calyptra to be 

fairly stable characters,not useful for species delineation. In the 

Philippines, this also is true and the species are separated 

mainly on vegetative leaf characters. 

There are four epiphytic species of Daltonia known from the 

Philippines. Their small size and tendency to grow in sparse 

patches probably contributes to the small number of collections 

deposited in various herbaria. 

Key to the Philippine Species of Daltonia 

1. Leaf apices aristate or abruptly mucronate ..................... 2 

Leaf apices acute to acuminate ........................... 3 

2. Upper leaf cells large, more than 30 pm long; setae strongly tuberculate-spinose 

above. ................................. D. armata 

Upper leaf cells small, less than 30 pm long; seta scabrous above . . D. ariscifolia 

3. Leaves strongly twisted or contorted when dry, oblong-lanceolate, the margins plain, 

the apices acute ............................ D. contorta 

Leaves slightly flexuose when dry, narrowly lanceolate to linear, the margins 

recurved, the apices acuminate .................. D. anguswolia 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

14 

FIGURES 11-14.4oltonio upper leaf cells: 11, D. ornuat0 (Phihpphes: Zamboanga. Dansalan, Zwickey 638, 

FH); 12, D. contort0 (Philippines: Baguio, Will- 1666, US); 13, D. orbtifolio (New Guinea: Robbins 27970, 

B); 14, D. ongustifolia (Philippines: Mt Canlaon, MerriN6813, US). 

Daltonia angusCCfolia 

FIGURE 14 

Ddtonia angustifolio Dozy & Molkenboer, Ann. Sci. Nat. Bot., ser. 3,2:302, 

1844. [Type: Bomeo,sine lom,Korthafs s.n. &),not seen.] 

Ddtonia revolvta Brotherus, Philipp. J. Sci., 5C (Bot.):156, 1910. D.pe: 

Philippines, Luwn, Mt. Pulog, Curron, Mcrritt & ZFchokke Bur. Sci. 16405 

(isotype, NY!).] 

Plants small, yellowish green, tuft forming. Stems simple or 

branched, without central strand, 5-10 mm tall. Leaves 

crowded, flexuose when dry, erecto-patent when moist, 2-3 

mm long, narrowly lanceolate to linear, with a welldifferentiated border of 2-3(4) rows of linear cells, the margins 

entire, strongly recurved on both sides. Costa single, strong, 

ending some distance below the apex. Leaf cells oval to 

narrowly rhomboidal and short fusiform, 21-30 pm long and 

4-6 pm wide, slightly thick-walled, becoming rectangular and 

lax only at base. Synoicous. Perichaetial leaves shorter and 

narrower. Calyptra strongly fringed with long hairs at base, 

covering only the operculum. Setae 5-10 mm long, scabrous in 

the upper half. Capsules ovoid, erect to inclined, 1 mm long; 

the exothecial cells collenchymatous. Exostome teeth reflexed 

when wet, papillose throughout; endostomial segments narrowly elongate, papillose throughout. 

SPECIMENS STUDIED.-PHILIPPINES. Luzon: Mt. Pulog, Tun 

82-200 (CAHP); Mt. Data,Nishimura 6788,67% (HIR0,US); 

Baguio, Williams 166.5 (US). Mindoro Is.: Mt. Halcon, Tan 

87-181 (CAHP). Neps: Mt. Canlaon, Merrill 6813 (US). 

Mindanao: Mt. Apo, Tun 82-270 (CAHP). 

Rmm.-Widespread in Southeast Asia and Australasia, 

this species is best distinguished by its long-acuminate leaf 

apices and recurved leaf margins. The margins of D. contortu 

are plain. The broad basal leaf border of D. ungustifoliu is not 

as abruptly differentiated as in D. contortu. In the former, the 

basal leaf border merges imperceptibly into the adjacent leaf 

cells. Several varieties have been treated by Fleischer (1908), 

Bartram (1939) and Sainsbury (1955). Considering the great 

variation of vegetative characters Seen in the few collections 

available for study, we have refrained from making comments 

on the infraspecific taxa. 
NUMBER 75 13 

Daltonia arisrjfolicr 

FIGURE 13 

Daltonia aristifolia Renauld & Cardot,Rev. Bryol., 23:105,1896. [”p: Java, 

Tjibodas, Massart inter 1395 (BR?),not seen.] 

Plants nearly of the same size as in D. angustifolia. Leaves 

also narrowly lanceolate, but with long, aristate apices. Costae 

reaching 2/s-1/2 of the leaf length. Leaf borders clearly 

differentiated, reaching the apex, entire, the margins plane or 

recurved. Leaf cells of somewhat irregular shapes, mostly 

oval-hexagonal, mixed with short oblong, rhomboidal to 

fusiform ones, (9-)15-30(-60) p long and 6-9 p wide, 

thin-walled, becoming more or less rectangular at leaf base. 

Autoicous. Sporophytes as in D. angustifolia, but exothecial 

cells thin-walled, slightly collenchymatous. 

SPECIMENS STUDIED.-PHILIPPINES. Mindanao: Mt. Apo, 

Tan & Aguila 82-230 (CAHP). NEW GUINEA: Robbins 2797a 

(B). SN LAM<A:Hubschmannma~ s.n. (B). 

REMARKS.--T~~S species is distinctive in having leaves with 

long hairpoints and comparatively small cells. Some forms of 

D. angustifolia develop unusually long-acuminate leaf apices 

and may key to D. aristifolia. The arista of D. aristifolia, 

however, is made up of several thick-walled,pitted, linear cells, 

similar to the leaf border cells. In D. angustifoliu, the long leaf 

apex consists of cells that are not different from other upper leaf 

cells. 

Daltonia baldwinii Brotherus from Hawaii has similar long 

aristate leaf tips, but the leaf cells are longer than those of D. 

aristifolia. The leaf costa of the former also reaches */3 of the 

leaf length. 

Daltonia aristifolia is a widespread species in tropical Asia. 

Its presence in the Philippines was fit reported by Tan and 

Iwatsuki (1983). 

DaUonia armata 

FIOURE 11 

Ddtonia armat0 Bartram. Farlowia, 1508, 1944. [?Lpe: Philippines. Mindanao. Sacre-d Mountain. Lanao,Zwickey 638 (holotype, FH!).] 

Plants small, as in D. angustifolia. Stems to 10 mm tall. 

Leaves flexuous when dry, narrowly lanceolate, gradually 

becoming aristate, or oblong-lingulate, 4 mm long, blades 

slightly folded at the lower half. Leaf costa reaching 2/3 the leaf 

length. Leaf borders weakly differentiated, with 1-2 rows of 

linear cells, the margins plane above, entire, and narrowly 

recurved below. Leaf cells oblong-fusiform above, 54-75 p 

long and 12-16 p wide, thin-walled, becoming laxly 

elongate below. Setae 5-7 mm long, strongly scabrous above. 

Capsules ovoid, 1 mm long. 

SPECIMENS STUDIED.-PHILIPPINES. Zamboanga, Dansalan, 

Zwickey 638 (FH). INDONESIA. Sumatra: Batang-Karo, 

Westkriods, no collector, ex Herb. Borg., sub Daltonia 

spiniseta Fleischer, nom. nud. 0. 

Rmms.--ACCording to Bartram (1944). this Philippine 

species resembles D. aristifolia in its aristate leaf apices, but 

differs from the latter in having longer leaf cells (Figure 1 1) and 

a strongly tuberculate seta. 

Daltonia urmata is hereby reported as new to Sumatra based 

on a specimen at FH named D. spiniseta in scheda by Fleischer. 

Apparently, Fleischer too was impressed by the strongly 

scabrous to nearly spinose setae of the present species. 

DaUonia contorta 

FIGURES 12,43-46 

Ddtonia contor& C. Muller. Syn.. 2:660,1851. I?Lpe: Java, sine loco, BIume 

s.n. (B, destroyed).] 

Plants small to medium size. Stems to 1-3 cm long. Leaves 

strongly contorted or twisted when dry, lanceolate, longacuminate. Leaf borders strongly differentiated, consisting of 

4-5 rows of linear cells at base; leaf margins not recurved. Leaf 

cells oval to short-rhomboidal, 15-36 pm long and 6-9 pm 

wide, thick-walled, becoming rectangular at base. Calyptra 

strongly fringed at base. Setae 5-6 mm long, nearly smooth or 

weakly papillose throughout. Operculum conic. Capsules 

variable in shape,ovoid to cylindric, 1-1.5 mm long. 

SPECIMENS TUDIED.-PHILIPPINES. Luzon: Mt. Data, Tan 

& Aguilar 81 -92 (CAHP); ibid, Nishimura 6528,6758 (HIRO, 

US); Baguio, Williams 1666 (US). 

REMARKs.--The leaves of D. contorta are strongly twisted 

when dry, befitting the species epithet. The acute to acuminate 

leaf apices with plane leaf margins are also diagnostic; 

although the last mentioned character is inconspicuous in the 

dry state. 

Daltonia contorta is also known from Sri Lanka, Java, Fiji, 

and Hawaii. Ail Philippine collections, thus far, come from the 

Island of Luzon (Bartram, 1939). 

Genus DistichophyUum 

Distichophyllwn Dozy & Molkenboer,Musci Frond. Ined. Archip. Jndici.4:99, 

1846. 

This is one of the large genera in the Hookeriaceae with 

about 65 species in tropical Asia and Oceania. It is also the 

largest genus in the family in the Philippines. 

The genus is characterized by somewhat complanate 

foliation, bordered leaves with rounded to hexagonal upper leaf 

cells, a single costa, short setae, and capsules with a typical 

hookericeous exostome. 

Like Actinodontium and Daltonia, the stem cross-section 

shows little or no internal differentiation. A central strand is 

absent in all of the Philippine species studied. However, many 

stem parenchyma cells develop a “sieve plate’’ structure with 

elliptic pits on their transverse walls, a peculiar feature 

illustrated by Fleischer (1908, fig. 168) and Noguchi (1956). 

Hookeria is another genus in the family with a similar sieve 
14 SMITHSO" CONTRIBUTIONS TO BOTANY 

FTOUW 15-22.-Dbichophyllwn: 15.16, D. snbcuspiaiatwn (Philippines: Sibuyan Is.. Mt. Giting-Giting, Tan 

& Hernaez 87-468, CAHP): 15, upper leaf cells; 16. whole leaves. 17,18, D. brevicwpes (Philippines: Sewer 

s.n., L): 17. whole leaves; 18, upper leaf cells. 19-22, D. kinabalmwe (Philippines: Sibuyan Is., Mt. 

Giting-Giting, Tan & Hernaez 87-446, CAHP): 19, plant, wec 20, plant, dry: 21, whole leaves; 22, upper leaf 

Cells. 
NUMBER 75 15 

plate development in the stem cell walls, but the stem 

cross-section of Hookeria shows a weakly differentiated central 

strand. 

The degree of heterophylly and leaf density along the stem 

axis is variable in Distichophyllum. Compared to the lateral 

rows of leaves, the two alternate rows of median leaves are 

variable both in their leaf morphology and leaf cell areolation. 

Thus, the foliar taxonomic characters described below in the 

species treatment refer only to the mature, lateral leaves. 

Furthermore, the leaf habit in dry state, the degree of 

homogeneity of cell size in the upper half of the leaf, the length 

of leaf costa, and the width of the leaf border are all variable to 

some extent. Still, within a limited range of variation, they can 

be used effectively for separation of related species. 

The perichaetial and perigonial leaves, surprisingly, are 

rather uniform in many species. They are often small (-l/3-% 

the size of lateral leaves), ovate-lanceolate, acute-acuminate, 

weakly bordered, and ecostate. Consequently, they do not have 

taxonomic significance. In contrast., the length and the 

papillosity of the setae are important diagnostic characters 

which correlate well with the sectional division of the genus. In 

general, section Mniadelphus Mitten includes large species 

with leaves more than 3 mm long and scabrous to papillose 

setae. The small species belonging to section Distichophyllum 

have leaves less than 3 mm long and have smooth, short setae. 

Also, the calyptra, which covers half of the capsule, is rather 

uniformly mitrate and fringed with a row of long hairs at its 

base. Many species differ in having a scabrous or papillose, 

hairy or smooth calyptral surface; but, these would have limited 

practical systematic value since collections often lack calyptrae 

even when parts of the sporophyte are present. Capsules, when 

present, are generally ovoid-cylindric with collenchymatous 

exothecial cells. In the following treatment, these sporophytic 

characters are not mentioned unless the Philippine specimens 

bear them. 

The majority of the Philippine species of Distichophyllum 

are dioicous, or at least reportedly so. Perichaetia are 

commonly seen in collections, but perigonia and sporophytes 

are infrequently encountered. Compared to Calyptrochaeta and 

Leskeodon, species of Distichophyllum usually do not produce 

filiform brood bodies in the leaf ails or on the adaxial leaf 

surfaces. , 

Philippine Distichophylla are commonly found in wet, 

humid, and shaded sites such as creek banks and dripping wet 

cliffs inside rainforests. A few species have become epiphytic 

on tree trunks and branches in humid, montane forests. Perhaps 

as an adaptation to this environment, the leaves of epiphytic 

taxa tend to be less complanate and the plants more tufted. 

In 1939,Bartram treated seven species of Distichophyllum in 

the Philippine archipelago. The number increased to nine in the 

checklist of Iwatsuki and Tan (1979). Our present study, which 

includes three as new synonyms, shows a total of 14 species, 

five of which are new additions to the local moss flora. 

Distichophyllum jungermannioides and D. nigricaule var. 

nigricaule have not been found in the Philippines, however, 

they are included in the key inside parentheses since they are 

likely to occur in the country. 

1. 

2. 

3. 

4. 

5. 

6. 

Key to the Philippine Species and Varieties of Distichophyllum 

Leaf apices coarsely toothed, denticulate or serrulate ............... 2 

Leaf apices entire. ................................ .3 

Leaf apices coarsely toothed. ...................... D. suntosii 

Leaf apices finely denticulate ................... D. obtusifolium 

Plants small, stems less than 25 mm long and not more than 3 mm wide including 

the lateral leaves. ............................... .4 

Plants large, stems more than 25 mm long and 3 mm wide including the lateral 

leaves ...................................... 16 

Leaf apices long-acuminate to aristate or cuspidate, the acumen more than 100 p 

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

less than 100 p long. ............................ .7 

Leaf cells small, less than 15 p wide, weakly or not collenchymatous ...... ................................... D. cuspidaturn 

Leaf cells large, more than 15 p wide, strongly collenchymatous ....... 6 

Leaves oblongelliptic, the apices broadly cuspidate; upper leaf cells mostly 15-21 

pm wide, pitted. ........................ D. subcuspidaturn 

Leaves narrowly lanceolate, the apices gradually acuminate to aristate; upper leaf 

cells mostly 21-33 p wide, not clearly pitted. ........ D. kinubuluense 

long ........................................ 5 
16 SMITHSONIAN CONTIUBUTIONS TO BOTANY 

7. 

8. 

9. 

10. 

11. 

12. 

13. 

14. 

15. 

16. 

17. 

18. 

19. 

20. 

21. 

Leaf apices round to obtuse, muticous ....................... 8 

Leaf apices apiculate or short-mucronate ..................... 9 

Leaves less than 2 mm long, broadly obovate from a short narrow base to nearly 

orbicular, the apices cucullate ................... D. catinifolium 

Leaves more than 2 mm long, spathulate to oblanceolate from a long narrow base, 

the apices not cucullate ......................... D. mittenii 

Leaf borders wide, of 3-4(-5) rows of linear cells, especially in the lower half of 

thele ........................................ 10 

Leaf borders narrow, of 1-2(-3) rows of linear cells in the lower half of the leaf . .......................................... 11 

Upper submarginal leaf cells small, 9-15 p wide, clearly smaller than the 

paracostal ones; leaf borders more than 30 p wide; leaf apices shortly mucronate, 

mucros less than 30 p long .............. D. nigricaule var. elmeri 

Upper submarginal leaf cells large, 21-30 pn wide, not or slightly smaller than the 

paracostal ones; leaf borders less than 30 p wide; leaf apices stoutly apiculate, 

acumina 30-60 p long ............. (0. nigricaule var. nigricaule) 

Leaf apices with stout acumen, more than 70 pm long ............. 12 

Leaf apices with short apiculus, less than 70 pn long .............. 13 

Leaves more than 2 mm long, not carinate at base along the costa; the submarginal 

leaf cells much smaller than the inner ones, slightly collenchymatous, not pitted . ............................... D. collenchymatosum 

Leaves less than 2 mm long, often carinate at base along the costa; the submarginal 

leaf cells not distinctly smaller than the inner ones, clearly collenchymatous and 

pitted. ............................... D. brevicuspes 

Leaves more than 2 mm long, narrowly oblanceolate to spathulate . . D. mittenii 

Leaves less than 2 mm long, oblong-ovate to obovate and broadly lingulate . . 14 

Upper leaf cells mostly less than 15 pm wide, thick-walled; costae well developed, 

reaching 2/3 of the leaf length .................... D. maibarae 

Upper leaf cells mostly more than 15 pm wide, thin-walled; costae weak, reaching 

only midleaf. .................................. 15 

Leaf apices apiculate, acumina less than 20 p long; upper leaf cells more or less 

homogenous from margin to costa ............... D. subnigricaule 

Leaf apices short-mucronate, acumina more than 20 pm long; upper submarginal 

leaf cells smaller than the inner ones ........... (0. jungermannioides) 

Leaves more than 2.5 mm long, oblanceolate, lingulate to spathulate ...... 17 

Leaves less than 2.5 mm long, ovate-oblong to short-obovate .......... 21 

Upper leaf cells more than 20 pm wide, more or less homogenous in size from 

margin to costa. ................................ 18 

Upper submarginal leaf cells less than 20 pm wide, distinctly smaller than the 

paracostal ones ................................. 19 

Leaf apices round or with short mucro, cucullate .......... D. cucullatum 

Leaf apices broadly obtuse, apiculate, not cucullate ........... D. tortile 

Leaves indistinctly or not bordered in the upper half. ....... D. osterwaldii 

Leaves distinctly bordered all around. ...................... 20 

Leaf borders with 1-2 rows of linear cells; leaf apices round, with a tiny mucro . D. mittenii 

Leaf borders with 2-3 rows of linear cells; leaf apices pointed, with distinct 

apiculus ........................... D. collenchymatosum 

Leaf borders with 2-3 rows of linear cells, not clearly differentiated at leaf base . ............................... D. collenchymatosum 

Leaf borders with 3-4(-5) rows of linear cells, strongly differentiated in the lower 

half of leaf. .................................. .22 

..................................... 
NUMBER 75 17 

22. 

23. 

Upper submarginal leaf cells more than 18 p wide, not distinctly smaller than the 

paracostal cells; leaf borders narrow, less than 30 pm wide ............ ........................... (0. nigricaule var. nigricaule) 

paracostal cells; leaf borders strong, more than 30 pm wide .......... 23 

Upper submarginal leaf cells less than 18 pm wide, distinctly smaller than the 

Leaf apices with well developed apiculus; the costae reaching more than 2/3 the leaf 

length, at times percurrent. ............. D. nigricaule var. cirraturn 

Leaf apices round, with a tiny mucro; the costae at most reaching 2/3 of the leaf 

length. ......................... D. nigricaule var. elrneri 

Distichophyllurn brevicuspes 

FIGLJREs 17.18 

Dktichophyllwn brevicuspes Fleischer. Musci Fl. Buitenzorg, 3:979, 1908. 

[Types: West Java, am Poentjak Pass, FIeischer s.n. (lectotype, FH!); West 

Java, bei Tjibodas, Flekcher s.n. (syntype, FH!).] 

Plants small, mat-forming. Stems to 1.5 cm long, 1.5 mm 

wide with leaves. Leaves oblong-elliptic to lingulate with a 

narrow, short, somewhat carinate leaf base, 1-2 mm long, 

cuspidate, the acumina (60-)75-90 p long, the borders 

narrow, with 1-2 rows of linear cells. Leaf cells hexagonalpolygonal, (18-)21-27 pm, thin-walled, collenchymatous, 

homogenous throughout the upper half. Gametangia and 

sporophytes not seen. 

SPECIMENS STUDIED.-PHILIPPINES: Semper s.n. (I,!); 

Luzon: Mt. Tabayog,Jacob B426 (L!). 

REhlARKs.-Known previously from Java, this species is a 

new addition to the Philippine flora. It is best recognized by its 

rather homogenous upper leaf cells, cuspidate leaf apices, and 

a somewhat carinate or folded leaf base. Distichophyllum 

nigricaule differs in having larger leaves (more than 2 mm 

long), larger paracostal leaf cells, more broadly cuspidate leaf 

apices, and stronger costae (Figures 23-28). The leaf borders 

of D. nigricaule are also stronger, consisting often of 3-4 rows 

of linear cells as compared to 1-2 rows for D. brevicuspes. 

Distichophyllum subcuspidatum differs from the present 

species in having a concave, more expanded leaf blade in the 

upper half, and a leaf base that is not carinate. 

Distichophyllum subcarinatum Noguchi & Iwatsuki from 

Borneo (Noguchi and Iwatsuki, 1972) was characterized as 

having a distinctly carinate leaf and a short leaf acumen. 

Although the cells of D. subcarinatum were described as 

non-collenchymatous, it was illustrated in the original publication with collenchymatous leaf cells. It appears likely that D. 

subcarinatum may be a synonym of D. brevicuspes. 

In the Fleischer Herbarium 0, both syntypes are present. 

The specimen from Poentjak Pass is selected as the lectotype 

because of its abundance of material. 

Distichophyllurn catinifoliurn 

Distichophyllum catinifolium Froehlich, Ann. Naturh. Mus. Wien,59:98,1953. 

['I).pe: Sumatra, Mt. Singgalang, Meijer 6157 (isotype. L!).] 

Dktichophyllwn hattorii Noguchi & Iwatsuki, J. Hattori Bot. Lab., 36~472, 

1972, syn. nov. [Type: North Borneo: Mt. Kinabalu,Z. Iwatsuki NICH 679a 

(holotype, NICH!).] 

Plants small, forming dense mats. Stems prostrate, to 2 cm 

long, 2 mm wide with leaves. Leaves contorted when dry, 

imbricate when wet, shortly obovate to orbicular, to about 1 

mm long, the apices obtuse to round, at times with small 

apiculus, cucullate. Costae short but clearly developed, 

reaching 2/3 of the leaf length. Leaf borders narrow, with 1-2 

rows of linear cells, the margins entire, somewhat incurved. 

Leaf cells lax, hexagonal-rounded, 12-16 pm, thin-walled, 

more or less homogenous throughout upper half of leaf, 

Perichaetial leaves small, broadly ovate, round to obtuse. Setae 

5-6 mm long, smooth. Capsules not seen. 

Rmm.-The type of Distichophyllum catinifolium was 

compared with D. hattorii and found to be identical. The round, 

cucullate leaf apices coupled with the lax and thin-walled leaf 

areolation are unmistakable. The other Philippine species 

having similarly cucullate leaf apices is D. cucullutum, but the 

latter is a large plant with spathulate leaves. Unfortunately the 

better known binomial, D. hattorii, postdates D. catinifolium 

by 19 years. 

The present species was first reported for the Philippines 

from Mt. Data by Iwatsuki and Tan (1980). We add here a 

second locality: the Philippine National Botanic Garden, 

Quezon Province,Luzon (Tan 81-456, CAHP). It is also known 

from several localities in Borneo and Sumatra. 

Distichophyllurn collenchyrnatosurn 

Dktichophyllwn collenchymatoswn Cardot, Bull. Soc. Bot. Genbe, s6r. 2, 

3:278,1911. [Type: Japan, Yakushima, Faurie 1156 in prt (holotype, PC), 

not seen.] 

Plants small to medium size. Stems slender, to 4 cm long and 

3 mm wide with lateral leaves. Leaves contorted when dry, 

broadly oblong, elliptic to lingulate, 2-3 mm long and 1.5 mm 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

wide, with well developed acumina (60-)75-90(- 100) pm 

long. Leaf borders with 2-3 rows of linear cells, entire, slightly 

undulate. Costae strong, reaching beyond 2/3 of the leaf length. 

Leaf cells hexagonal to round, (18-)21-30(-33) pm wide, 

slightly collenchymatous, markedly decreasing in size toward 

the margins in the upper half of the leaf, Calyptra smooth 

above. Setae 15-20 mm long, smooth. 

SPECIMEN STUDIED.-PHILIPPINES. Negros: Mt. Talinis 

(Cuernos Mt.),A. de Jesus, s.n. Oct-Nov. 1982 (CAHF'). 

REMARKs.-Distichophyllum collenchymatosum is a subtropical species in East Asia with its southern limit in the 

central Philippines (Tan and Iwatsuki, 1983). It can be 

confused with D. nigricaule, from which it differs in having a 

narrower leaf border. In spite of its species epithet, this species 

does not develop leaf cells that are as strongly collenchymatous 

as in some other local species. 

Distichophyllum cucullatum 

Distichophyllum cucuUatum Bartram, Lloydia, 5279, 1942. [?Lpe: West 

Irian, Lake Habbena,Brass 95W (holotype, FHI).] 

Plants large, stems to 5 c m long and 4 mm wide including the 

leaves. Leaves long-spathulate to obovate, 2-3 mm long and 

1.25-2 mm wide, the apices round, with very small apiculus, 

mostly cucullate. Leaf borders narrow, of 1-2 rows of linear 

cells. Costae weak, reaching l/2 of the leaf length. Leaf cells 

large, hexagonal to polygonal, (18-)24-36(-45) pm, thinwalled, laxly arranged in the expanded orbicular upper part of 

the leaf, the submarginal cells slightly smaller than the medial 

ones. Setae strongly hirsute, 5-7 mm long. Capsules cylindric, 

to 1 mm long, neck elongated. 

SPECIMENS STUDIED.-PHILIPPINES. Mindanao: Muralong 

Mt., near Kabasalan,Zamboanga,Ebalo 731 (MICH, CAHF'). 

REMARKS.-This species is easily recognized by its large 

size and spathulate leaves that are cucullate at the apex. The 

Philippine specimen compares well with the type of D. 

cucullatum, differing only in the smaller plant size. Because of 

its large size, Distichophyllum cucullatum is unlikely to be 

confused with the much smaller species, D. catinifolium, which 

also has similarly cucullate leaf apices. According to Bartram 

(1942), the setae of D. cucullatum are papillose, while those of 

D. catinifolium are smooth. 

Distichophylum cucullatum was known previously only 

from New Guinea. Its extension to the island of Mindanao in 

the southern Philippines is a disjunctive distribution pattern 

shared by a few other mosses such as Plagiotheciopsis oblonga 

(Brotherus) Brotherus ex Dixon (Tan, 1987). 

Distichophyllum cuspidatum 

FIGURES 29,30 

Dhtichophyllwn cwpidatwn (Dozy & Molkenboer) Dozy & Molkenboer, 

Hwkeria cwpi&ta Dozy & Molkenboer, Ann. Sc. Nat. Bot.. ser. 3,2305, 

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

1844. [Types: Java, pope Limbang, Zippeliw s.n., Korthals s.n. (L); 

Sumatra, Batang-BessieJorthds s.n. (L).] 

Plants forming aggregate tufts. Stems to 2 cm long. Leaves 

twisted when dry, not complanate, 1-1.5 mm long and 1-1.25 

mm wide, mostly elliptic-oblong to oblanceolate, rarely 

narrowly lingulate, at times concave, long-cuspidate, weakly 

carinate at leaf base, the leaf acumina variable in length, 

150-300(-360) pm long, the leaf borders well developed, 

consisting of 2-3 rows of thick-walled, elongate cells. Leaf 

margins entire, slightly revolute at leaf base; the costae 

reaching more than half the leaf length. Leaf cells small, round 

to hexagonal, (6-)9-15(-18) pm, thin to moderately thickwalled, weakly collenchymatous toward the apex. Setae 

smooth below, slightly to strongly papillose above, 3-5 mm 

long. Capsules short-ovoid to ovoid+blong, erect or inclined. 

SPECIMENS STUDIED.-PHILIPPINES. Sibuyan Is.: Mt. GitingGiting, Tan & Hernaez 87-528 (CAW, US). INDONESIA. East 

Ceram: G. Selagor, near Kp. Selagor,Buwalda 5731 @I). 

REMARKs.-Although D. cuspidatum was well illustrated by 

Dozy and Molkenboer (1846, pl. 33) and Fleischer (1908, fig. 

169), this reportedly widespread species in Malesia and 

Oceania has not been well understood. 

Recent authors (Bartram, 1939; Noguchi, 1956) have 

expanded its concept to include a number of species with 

similarly long-cuspidate leaf apices. The only Philippine 

specimen (Mt. Apo, Williams 2672) cited by Bartram (1939) as 

D. cuspidatum and the Taiwan specimens illustrated by 

Noguchi (1956) show a totally different leaf shape: ligulate 

with a narrowly triangular leaf apex terminating in a long arista 

(Figures 20, 21). The leaf cells of these specimens are also 

collenchymatous, a feature not typically seen in D. cuspidatum. 

In fact, Williams 2672 is D. kinabaluense, a species Noguchi 

and Iwatsuki (1972) segregated from D. cuspidatum on the 

basis of their Bornean collections. 

Furthermore,in D. cuspidatum, the broadest part of the leaf 

is just below the apex and is about 1 mm wide, while in the leaf 

of D. kinabaluense the broadest part is near the middle and is 

less than 1 mm wide (Figures 21,30). These two taxa may 

prove to be conspecific, but for the present, we believe that they 

are distinct species. 

Distichophyllum kinabaluense 

FIGURES 19-22 

Dktichophyllwn kinabaluense Noguchi & Iwatsuki, J. Hattori BOL Lab., 

36:474, 1972. [Types: Sabah, Mt. Silam, alt. 900-1000 m, Iwcl~~uki NICH 

5512 (holotype, NICH!; isotyp, Herb. Noguchi!).] 

Plants slightly larger than D. cuspidatum. Leaves contorted 

when dry, narrowly lanceolate, oblong-lingulate to shortligulate, about 2 mm long, the upper half gradually narrowed to 

long aristate tips, the acumina 200-480 pm long, and slightly 

subcarinate at leaf base. Leaf costae distinct, reaching 2/3 of the 

leaf length or shorter. Leaf margins entire, plane above, 
NUMBER 75 19 

revolute at leaf base. Leaf cells round to polygonal, (15-)18- 

27(-33) m, thick-walled, subporose, collenchymatous, becoming more lax and rectangular below, the marginal rows of 

cells slightly smaller than the medial ones. Rhizoidal filiform 

brood bodies present in some leaf mils. Setae to 5 mm long, 

smooth throughout. Capsules not seen. 

SPECIMENS STUDIED.-PHILIPPINES. Sibuyan Is.: Mt. GitingGiting, Tan & Hernaez 87-446 (CAHP), 87-452 (US). 

Mindanao: Mt. Apo, Williams 2672, sub D. cuspidutum (US); 

Mt. Malindang, Misamis Occidental Prov., Reyes 15, sub 

Distichophyllidium philippinense Bartram, nom. nud. (NY). 

Rmms.-The above species description is based on the 

Philippine materials determined by us on the basis of the 

original description and illustration in Noguchi and Iwatsuki 

(1972). The identification has since been confirmed by 

examination of type material. 

In Distichophyllum kinabaluense the length of the leaf 

acumina and costae can vary greatly in leaves taken from the 

same plant. For the distinctions from D. cuspidutum see the 

discussion under that species. 

Distichophyllum iwatsukii Noguchi from Mt. Kinabalu, 

Borneo, was described as having a similar leaf cell areolation 

but the plants are larger with leafy stems up to 2.5 cm long and 

4 mm wide (Noguchi and Iwatsuki, 1972). The leaf acumina of 

D. iwatsukii have been illustrated as shorter than those of D. 

kinabaluense. The relationship between these two taxa deserves 

more attention. 

Distichophyllum maibarae 

Dirtichophyllwn maibarae Bescherelle, J. Bot (Morot), 1340, 1899. [?Lpe: 

Central Japan, Maibara, Faurie 11130 (holotype, BM), not seen.] 

Plants small to medium size, in dense mats. Stems creeping, 

branched, to 1.5 cm long, less than 3 mm wide including the 

leaves. Leaves crisp and undulate when dry, somewhat 

imbricate when wet, oblong-lanceolate to slightly oblanceolate, 

1-1.25 mm long and 0.5-0.75 mm wide, apiculate to shortly 

mucronate, the acumina (21-)30-50(-60) pm long. Leaf 

borders narrow, of about 2 rows of linear cells, becoming 

weakly differentiated near the apex and toward the base. Costae 

flexuose above, reaching 2/3 of the leaf length. Leaf cells small, 

round to hexagonal, 9- 15(-18) pm, not collenchymatous and 

more or less homogenous in the upper half of the leaf in fresh 

state. Calyptra rather hairy. Setae short, less than 10 mm long, 

smooth. Capsules small, ovoid, 1 mm long, the exothecial cells 

slightly mamillose, collenchymatous. 

SPECIMENS STUDIED.-PHILIPPINES. Batanes Is.: Mt. Iraya, 

Tan 84-163 (CAX-IP). Luzon: Mt. Tabayog, Jacob B 485 (sub 

D. nigricaule), B 314 (L, CAHP); Mt. Data, Nishimura 6731 

(HIRO). 

REMARKS.-T~~S Sino-Japanese species was first reported 

for the Philippines by Shin (1965), from the Batan Islands 

located at the far northern tip of the country. Since then, several 

collections have been made from the high mountains of 

northern Luzon where the local flora has strong Himalayan and 

temperate Asian affiiities. 

Among its local congeners, D. maibarae is rather 

characteristic in its oblong-lingulate leaf shape with a short 

mucro consisting of a few short, oblong to rhomboidal cells. 

These cells contrast strongly with the linear border cells, but are 

similar to the adjacent lamina1 cells. Occasionally, specimens 

of D. nigricaule may be found with a similar leaf apiculus, but 

the leaf borders of D. nigricaule are always broader or more 

strongly developed than those of D. maibarae. 

The species is reported from mainland China, Taiwan, Japan, 

and the Philippines. In Indochina and the Himalayas the species 

may be present under the name, D. sinuosulum Dixon. The 

description and illustration of D. sinuosulum provided by 

Gangulee.( 1977) matches D. maibarae. mica1 D. sinuosulum 

from the Malay Peninsula is not D. maibarae but is a synonym 

of D. nigricaule var. elmeri. 

The species may be in herbaria from other parts of Malesia 

under the name D. montagneanum (C. Muller) Bosch & 

Lacoste. If this turns out to be so, then D. montagneanum, 

which was recently redescribed by Townsend (1982) from the 

type deposited at Paris, is the oldest available name. 

Distichophyllum mittenii 

Dirtichophyllwn mittenii Bosch & Lacoste, Bryol. Jav., 225, 1861. [?Lpes: 

Java, sine loco, Teysmann s.n. (L); Holle s.n. (L); prope Toegoe, pr. Bogor, 

Amann s.n. &),not seen.] 

Plants medium to large size, mat-forming, laxly foliate, 

complanate. Stems simple or branched, prostrate or ascending, 

to 4 cm long and 4 mm wide including the leaves. Leaves 

slightly wrinkled to moderately crisped when dry, spathulate to 

obovate from a narrow leaf base, 2-3 mm long and 1.5 rnm 

wide above midleaf, the apices broadly rounded above with one 

or two tiny apicula measuring less than 15 pm long. Costae 

strong, reaching near the apex. Leaf borders narrow, with 1(-2) 

rows of linear cells all around, the margins entire, undulate 

even when wet. Leaf cells hexagonal to polygonal, thickwalled, pitted, not collenchymatous, the submarginal cells in 

upper half of the leaf forming several rows of small (9-15 pm) 

cells; paracostal cells distinctly larger, 18-27 pm. Calyptra 

scabrous above. Setae scabrous, 7-8 mm long. Capsules ovoid 

to ovoid-oblong, the exothecial cells strongly mamillose. 

SPECIMENS STUDIED,-PHILIPPINES. Luzon: Mt. Makiling, 

Pancho 4913 (CAHP); Mt. Banahao, Pancho 1087 (CAHP); 

boundary of Albay and Sorsogon provinces, A. de Jesus, s.n. 

(PNOC). Negros: Dumaguete area, Mack 1302 p.p. (NY). 

MALAYSIA. Malay Peninsula: Pahang, Fraser Hill, Burkill & 

Holttum 8714 0; Selangor, Templer Park, Manuel 2692, sub 

D. tortile (NY). 

REMARKS.-This is the most common species of Distichophyllum found in the Philippine rainforests. The most 

obvious characters are the very narrow leaf border in the upper 

half of the leaf with usually one row of linear cells, the minutely 
20 SMlTHSONIAN CONTRIBUTIONS TO BOTANY 

mucronate leaf apices, and the several rows of submarginal 

cells distinctly smaller in size than the medial paracostal cells. 

Distichophyllum nigricaule var. cirratum has similarly 

heterogeneous upper leaf cells, but its leaves are smaller and 

have broad, strongly developed borders consisting of 3-4 rows 

of linear cells. Furthermore, the leaf shapes of the two taxa are 

different, spathulate from a narrow base in D. mittenii versus 

oblong-elliptic to oblanceolate or lingulate in D. nigricaule var. 

cirratum. The Philippine specimen of D. nigricaule from Mt. 

Makiling (Hadden 137a) reported by Bartram (1939) consists 

of juvenile plants of D. mittenii. 

Distichophyllum undulatum Dozy & Molkenboer ex Bosch 

& Lacoste has a similar leaf shape, leaf border, and leaf cell 

areolation, but differs from the present species in having an 

irregularly denticulate upper leaf border. 

Distichophyllum mittenii is widely distributed in tropical 

Asia and Oceania, reaching as far south as New Caledonia and 

Vanuatu. 

Distichophyllum nigricaule 

FIGURES 23-28 

Distichophyllwn nigricaule Mitten ex Bosch & Lacoste, Bryol. Jav., 2:20, 

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

Eysmn s.n. (syntypes, BO, L).] 

REMARKS.-T~~S is a variable species and several varieties 

have been described in the literature. ?3pically the infraspecific 

taxa are quite distinctive, but many intermediate forms can be 

found. One common feature present in all of the taxa closely 

related to D. nigricaule is a dark brown stem. Filiform brood 

bodies have been observed in Philippine specimens of D. 

nigricaule. 

Distichophyllum nigricaule has been reported from Indochina to MaIesia. Based on our examination of Philippine 

material, we recognize three varieties. 

DistichophyUum nigricaule var. nigricaule 

FIOURES n, 28 

Distichophyllwn gracilicaule Fleischer, Musci Fl. Buitenz.org, 3:983, 1908, 

syn. nov. [Types: West Java, um Tjibodas am Gedeh, Fleischer s.n. 

(lectotype, FH!); am Poentjak,Fleischer s.n. (syntype, FH!).] 

Plant small to medium size, dull yellow-green, mat-forming. 

Stems dark brown, prostrate to procumbent, slender, 2-6 cm 

long and -2.5-3 mm wide including the leaves, laxly foliate, 

Leaves strongly crisped when dry, 2-2.5 mm long and 1-1.25 

mm wide, short-ovate to obovate, at times broadly oblong, the 

apices rounded to broadly obtuse, shortly apiculate, the 

acumina 30-60 pm long. Leaf borders broad, well differentiated at leaf bases, with 2-3(-4) rows of linear cells and 

measuring 15-21(-30) pm wide. Costae moderately strong, to 

2/3 of the leaf length or shorter. Leaf cells round to hexagonal, 

(18-)21-30(-39) pm, thin-walled, weakly collenchymatous 

near the apex, slightly decreasing in size toward the margins 

and becoming short-rectangular at base. Perichaetial leaves 

very small. 

REMARKS.-There are two syntypes of D. nigricaule 

mentioned in the original publication. Since the species was 

attributed to Mitten by the authors of Bryologia Javanica (Dozy 

and Molkenboer, 1855-1870), the specimen at the New York 

Botanical Garden was selected as the lectotype. 

In Distichophyllum nigricaule, the leaves tend to be more 

oblong and narrowly obovate, although broadly oblong ones 

are not rare. On the other hand, the leaves of D. gracilicaule are 

mostly broadly elliptic to broadly oblong, as illustrated by 

Fleischer (1908, fig. 168). These differences appear to be 

insignificant for the group, and the two taxa should be united. 

When Fleischer (1908) described D. gracilicaule, he appears 

to have been misguided by his examination of atypical 

specimens of D. nigricaule. In his treatment of the two ma, 

Fleischer stated that the costa of D. nigricaule is often 

percurrent and also that the submarginal cells are distinctly 

smaller than the inner ones. These distinctions are not found in 

the type specimen of D. nigricaule. As a result, many 

specimens of D. nigricaule, including those cited by Fleischer 

(1908) and by Bartram (1939), are misidentified. They are 

instead D. nigricaule var. cirratum, which has leaves with 

strong and undulate borders, a nearly percurrent costa, and 

several rows of distinctly smaller submarginal leaf cells. The 

leaf borders of var. cirratum are broader (about 30-42 pm 

wide) and have 3-4(-5) rows of linear cells,especially near the 

leaf bases. In var. nigricaule, the leaf border measures only 

15-21(-30) pm wide and often consists of 2-3(-4) rows of 

linear cells near the base. 

Although we have not seen any Philippine material of var. 

nigricaule, we include a species description based on our 

examination of the lectotype at NY for purposes of comparison. 

All previous reports of D. nigricaule from the Philippines, have 

proven to be either var. cirratum or var. elmeri. 

Distichophyllum nigricaule var. cirratum 

FIGURES B,24 

Distichophyllum nigricaule var. cirratwn (Renauld & Cardot) Fleischer, Musci 

Fl. Buitenzolg,3:982,1908. 

Distichophyllwn cirratwn Renauld & Cardot, Rev. Bryol., 23:104, 18%. 

[%: Java, Tjbodas, Massart 1397 (isotype, FHI).] 

Distichophyllwn prwddatwn Dixm, J. Linn. Soc. BOL, 50106,1935, syn. 

nov. [Type: North Borneo: Sarawak, Ulu Koyan, Oxford Exped. no. 1863 

(holotype, BMU.1 

Plants small to medium size, mat-forming. Stems dark 

brown, slender, 1.5-3 cm long and 2-2.5 mm wide including 

the leaves. Leaves strongly contorted when dry, 1-2.5 mm 

long and about 1 mm wide, oblong-elliptic, oblanceolate to 

short-lingulate, the margins entire, undulate, with strong 
NUMBER 75 21 

FI~URES 23-28.-Disrichophyllwn nigrkade varieties: 23,24, D. n. var. cirrarwn (Java: Tjibodss, Massart s.n., 

type of D. cirratum, FH): 23, upper leaf cells; 24, whole leaves. 2526, D. n. var. elmeri (Philippines: Baguio, 

Elmer 8716, FH): 25, upper leaf cells; 26, whole leaves. 2728, D. n. var. nigricaule (Java: Pangemgo, Morley, 

type of D. nigricaule, NY): 27,upper leaf cells; 28, whole leaves. 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

apiculus, the leaf acumina (54-)60- 105 p long. Leaf borders 

strong, 30-42 p wide, with 3-4(-5) rows of thick-walled 

linear cells well differentiated at leaf bases. Costae strong, 

reaching near apex, at times percurrent. Leaf cells hexagonalpolygonal, moderately thick-walled, forming 6-8 rows of 

small submarginal cells, 9-18 p, and large paracostal cells, 

21-33 pm. Sporophytes not seen. 

SPECIMENS STUDIED.-PHILIPPINES. Luzon: Mt. Makiling, 

Tan 84-66 (CAHP). INDONESIA. Java: Tjibodas, Massart s.n. 

0. 

REMms.-When D. cirratum was fiist described by 

Renauld and Cardot (1 896), it was compared to D. cuspidatum 

which has a much longer leaf acumen. Fleischer (1908) 

correctly reduced it to a variety of D. nigricaule. 

The var. cirratum has not been clearly distinguished, in the 

past, from the typical form of D. nigricaule. The differences 

between the two taxa are discussed above under D. nigricaule 

var. nigricaule. In addition, the leaf acumina of var. cirratum, 

measuring 60-105 p long, are longer than those of var. 

nigricaule, which measure about 30-60 p long. The 

description of D. nigricaule in Fleischer (1908) better fits the 

present variety. 

Distichophyllum perundulatum (Dixon, 1935) from Borneo 

is identical with this variety. 

Distichophyllum nigricaule var. elmeri (Brotherus) Tan & 

Robinson, new combination 

FI~URES 25.26 

Dirtichophyllwn elmeri Brotherus, Leaflets Philipp. Bot., 2656, 1909, 

basicmym. [Type: Philippines, Lum, Baguio. Benguet Prov., Elmer 10453 

Dbtichophyllwn sinuaruiwn &on, I. Siam Soc.,Nat. Hist., Suppl. 10 (1):15, 

1935, syn. nov. [Type: Malaysia, Perak, Binch’s Hill, Burkill 12606 

(holoty~e, H-BRI).] 

(holotype, BW.1 

Plants similar to var. cirratum in habit, size and leaf shape. 

Like var. cirratum, the submarginal cells are 9-15 p in 

diameter, and are markedly smaller than the 21-27 p 

paracostal cells; the leaf borders are as broad, consisting of 3-4 

rows of linear cells (Figure 25). However, in var. elmeri, the 

leaf apices are muticous or very shortly mucronate (6-9 pm) 

and the costae never reach the leaf tips (Figure 26). 

SPECIMENS S TUDIED.-PHILIPPIN ES . Luzon: Baguio, Elmer 

8716 0; Mt. Makiling, Hadden 137 (FH), Tan 82-52 

(CAHP). Basilan: UP Land Grant, Zamboanga Prov., Ebalo 

964 0. 

REMARKS.-Brotherus (1909) distinguished D. elmeri from 

D. nigricaule by its different leaf form and its broader leaf 

border. Later, this species was reduced to a synonym of D. 

nigricaule by Bartram (1939), who did not examine the original 

material. The species was described by Brotherus (1909:656) 

as having leaves narrowly oblong, about 2.5 mm long and 0.9 

mm wide, with undulate margins consisting of strong borders 

of 3-4 rows of cells and marginal leaf cells much smaller than 

the inner ones, and this compares well with D. nigricaule var. 

cirratum. Our examination of the type of D. elmeri confirms 

their close relationship. 

Typical plants of var. elmeri are easily separated from var. 

cirratum and var. nigricaule. This led Noguchi and Iwatsuki 

(1972) to resurrect D. elmeri. The presence, however, of 

intermediate collections, e.g., Touw 11 709 from Khao Luang, 

Thailand (NY, L), indicate to us that the taxon should be 

recognized at the varietal level. 

Distichophyllum obtusifolium 

Distichophyllwn obtusifoliwn Theriot, l.e Monde des mantes, ser. 2, 9~22, 

1907. [Type: Japan, Amami-oshima, Naze, J. Ferrie s.n., Nov. 1899 

(holotype, PC), not seen.] 

Plants small to medium size, in thin mats. Stems procumbent, branched, often becoming ascending, to 3 cm long and 4 

mm wide including the leaves. Leaves crisp when dry, 

narrowly oblanceolate to broadly obovate and spathulate. Leaf 

borders narrow, consisting of 1-2 rows of linear cells, weakly 

differentiated into one narrow row of short rectangular cells 

near the round apex, the margins plane, finely denticulate by 

the prominent cell angles near the apex. Costae reaching l/2-2/3 

of the leaf length. Leaf cells mostly large, round-hexagonal, 

thin-walled, not collenchymatous, with several rows of 

submarginal cells, about 12-24 p wide, clearly smaller than 

the 30-48 p inner paracostal cells. Sporophytes not seen. 

Rmms.-Like D. maibarae, this species was first 

collected in the Philippines from the Batan Islands (Shin, 

1965). Its leaves are distinctive in having a broadly obovate to 

spathulate outline with rounded, denticulate apices. The 

peculiarly toothed leaf apices were accurately illustrated by 

Noguchi (1956). It has yet to be re-collected in the Philippines. 

The Malayan taxon, D. denticulatum Dixon, is closely 

related to the present species in having similarly rounded and 

toothed leaf apices. The two taxa differ from each other in the 

more narrowly obovate to oblanceolate leaves with fewer rows 

of smaller submarginal cells in D. denticulatum. Perhaps more 

collections of both taxa in tropical Southeast Asia will bridge 

this observed morphological gap and prove their synonymy. 

Distichophyllum undulatum Bosch & Lacoste from Java and 

Borneo is also similar in having weakly toothed leaf apices. 

Nevertheless, D. undulatum has an apiculate leaf tip, the apical 

marginal teeth of the leaf are fewer and irregularly positioned, 

and the leaf cells are more or less homogenously small (12-21 

p) in the upper half of the leaf. Another pronounced 

difference between the two species is in the width of the median 

leaf cells. In D. undulatum, the paracostal leaf cells are 18-27 

p wide, whereas those of D. obtusifolium are 30-48 p 

wide. Lastly, the leaf borders of D. undulatum, like those of D. 

m’ttenii, are made up of a single row of linear cells in the upper 

half of the leaf. 

Compared to D. obtusifolium, D. osterwaldii can be 

distinguished by its total lack of a differentiated border in the 
NUMBER 75 23 

leaf apex. Distichophyllum ostenvaldii is also a larger plant 

with entire leaf margins. These differences are best seen in 

fresh specimens where the leaf border cells are still fully turgid. 

In rehydrated specimens of D. osterwaldii, some leaf border 

cells may stay collapsed, thereby creating a false appearance of 

having a faintly mthed leaf margin. 

Dislichophyllum osterwaldu 

Distichophyllwn ostenvaldii Fleischer, Musci Fl. Buitenzorg, 3~994, 1908. 

[?Lpe: West Java, Gedehgebirge bei Tjibodas, Flcischer s.n. (FH),not sea] 

Plants large, loosely tufted. Stems ascending, to 8 cm long, 

laxly foliate, flat,4 cm long and 8 mm wide with leaves. Leaves 

strongly undulate-crispate when dry, lingulate to spathulate 

from a narrow leaf base, 4 mm long and 3 mm wide, broadly 

rounded above and muticous. Leaf borders distinct only in the 

lower half, narrow, 1-2 rows of linear cells. Costae slender, 

reaching more than 3/4 up the leaf length. Leaf cells small, 

round-hexagonal, 6-10 rows of submarginal cells, 12-21 pm 

in diameter, distinctly smaller than the 24-36 pm paracostal 

cells. Setae 5-7 mm long, papillose. Capsules ovoid-oblong, 1 

mm long. 

SPECIMENS STUDIED.-PHILIPPINES. Luzon: Mt. Wing, 

Tan 81-426 (CAHP), Nishimura 6989 (HIRO); National 

Botanic Garden, Real, Quezon Prov., Tan 81-452 (CAHP). 

Leyte: Mt. Pangasugan, Baybay, Tan & Na;arez 84-253 

(CAHP). 

REMARKs.-This is the largest and one of the more 

commonly seen species of Distichophyllum in the Philippines. 

The large, spathulate leaves with an undifferentiated upper leaf 

border are unique among the Philippine species. 

Distichophyllum tortile has comparably large leaves, but its 

leaf cells are large and relatively homogenous in the upper half 

of the leaf. Some forms of D. obtusijolium with weakly 

differentiated apical leaf borders may be confused with D. 

osterwaldii, but the former is a small plant about half the size of 

the present species. Also, in D. obtusijolium the leaf apices 

always have some trace of a differentiated border. 

Distichophyllum ostenvaldii is also known from Java, 

Borneo, Taiwan, mainland China, and Japan. 

Distichophyllum santosu 

Distichophyllwn santosii Bartram. Philipp. J. Sci, 68:259, 1939. [Type: 

Plants small, in dense mats. Stems creeping, much branched, 

reaching 2 cm long. Leaves not contorted when dry, very 

cornplanate, narrowly spathulate-lingulate from a long narrow 

base, 2-3 mm long, rounded at apex, bluntly toothed by the 

protruding apical marginal cells. Leaf borders narrow and 

indistinct, both at the apex and base. Costae faint, short, 

reaching only midleaf. Izaf cells very lax, hexagonal, nearly 

homogenous throughout. Setae 5-6 mm long, papillose. 

Capsules ovoid-oblong, the exothecial cells mamillose and 

Philippines, Luzon, Baler.Tayabas Pmv., Santos 231 (holotype, FH!).] 

collench ymatous. 

SPECIMENS STUDIED.-PHILIPPINES. Samar: Bagacay, Tun 

s.n., June 29,1975 (CAHP). 

REMms.-This is the only endemic Distichophyllum in the 

country. Its strongly toothed apical leaf margins have no match 

among local congeners, although its leaf shape and leaf border 

differentiation suggest a relationship with D. obtusjfolium. In 

the Philippines, D. santosii seems to occupy only the wet 

eastern part of the archipelago that faces the Pacific Ocean. 

The report of D. aff. santosii from Seram Island (Akiyama, 

1988) is based on a specimen (Akiyama 16268, CAHP) that 

proves to be D. denticulatum Dixon. 

Dhtichophyllum subcuspidatum 

15.16 

Distuhophyllwn subcaupidatwn Noguchi & Iwatsuki. J. Hattori Bot. Lab., 

36:476, 1972. [?Lpe: North Borneo, Mt. Kinabalu, below Paka Cave, 

Iwatsuki NICH 838 (holotype, NICH).] 

Plants small, like D. cuspidatum. Leaves slightly contorted 

when dry, oblong to oblanceolate, slightly concave in some, 

weakly plicate in others, broadly cuspidate, the acumina strong, 

210-240 pm long. Leaf margins plane, entire, the borders 

strong, with 3-4 rows of very thick-walled, linear cells. Costae 

reaching 2/3 of the leaf length. Leaf cells hexagonal to round, 

(9-)12-21(-24) pm, thick-walled, strongly collenchymatous, 

distinctly prose, more or less homogenous in the upper half. 

Sporophytes not seen. 

SPECIMENS STUDIED.-PHILIPPINES. Sibuyan Is.: Mt. GitingGiting, Tan & Hernaer 87-508 (US), 87-468,87-518 (CAHP). 

REMARKS.-Distichophyllum subcuspidatum, which is 

newly reported from the Philippines, appears to be closely 

related to D. cuspidatum. Both taxa develop long, apical leaf 

acumina and strong leaf borders. However, the leaf cells of D. 

subcuspidatum have distinctly pitted cell walls and collenchymatous corners. These distinctions may prove erroneous when 

more collections of the two species are made and compared. 

Noguchi and Iwatsuki (1972) reported fusiform and falcate 

brood bodies for the species from Bornean plants. 

The Philippine plants, like the Bornean specimens, came 

from basic substrates at an elevation between 3500-4000 ft. 

Distichophyllum subnigricaule 

Distuhophyllwn subnigrkaule Brotherus, philipp. J. Sd.. 31:289, 1926. 

"Qpes: Philippines, Mindanao, Zambcanga, hierrill 8324 (holotype, HI; 

Distichophyllum laxirete Noguchi & Iwatsuki, J. Hattori Bot. Lab., 36:425. 

1972, syn. nov. [?Lpe: North Borneo, Mt Kinabalu, near Ulu Liwagu. 

Mizvtani 3604~ (holotype. NICH!).] 

DiFtichophyllwn rotdtwn Noguchi & Iwatsuki, J. Hattori Bot. Lab., 36~476, 

1972 [?Lpe: Sabah: Mt Kinabalu, between Hot Spring, Poring, and Royal 

Soc. Bungalow,Iwatssuki 17110 (holotype.NICH).] 

Plants small to medium size, pale green, in dense mats. 

isotypes. NY!, FH!).] 
24 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Stems dark brown, 1-3 cm long and 2-2.5 mm wide including 

the leaves. Leaves slightly contorted when dry, 1-2 mm long, 

broadly lingulate to obovate, at times nearly orbicular, the 

apices rounded to broadly obtuse, weakly apiculate in some 

leaves, the apiculus, if present, 9-15 pn long. Leaf borders 

very narrow, with one row of linear cells near apex. Costae 

short, often less than half the leaf length. Leaf cells large, 

hexagonal, 27-40(-45) pn wide, very thin-walled, laxly 

arranged. Setae strongly spinose, 6-8 mm long. 

SPECIMENSTUDIED.-PHILIPPINES. Luzon: Mt. Wing, 

Tan 84-62 (CAHP). Leyte: Mt. Pangasugan, Baybay, Tun 

REMms.-This species is distinctive in its very lax leaf 

cell areolation. The distinction between this species and D. 

cutinifolium has been clatified by Noguchi and Iwatsuki 

(1972). 

Iwatsuki and Tan (1980) reported D. subnigricaule for 

Borneo under the synonym D. rotundaturn Noguchi & 

Iwatsuki. In the same publication, they reported D. luxirere as 

a new addition to the Philippine moss flora. According to 

Noguchi and Iwatsuki (1972) and Iwatsuki and Tan (1980), D. 

laxirete differs from D. subnigricaule in leaf shape and costa 

length. As more Philippine specimens of D. subnigricaule have 

been studied, these differences have become less clear. It seems 

best to recognize only one species. 

Distichophyllum jungermannioides (C. Muller) Bosch & 

Lacoste, from Borneo and Java, is closely related to the present 

species. The two share a similarly lax leaf cell areolation, but 

the leaf costae of D. jungermannwides are shorter, often less 

than l/3 the leaf length, and the leaf apiculus stouter, measuring 

60-75 prn long. The upper leaf cells of D. jungermannwides 

also appear to be smaller, 12-21 p wide. Fleischer (1908,fig. 

168) elegantly illustrated D. jungermannwides showing the 

characteristically concave, broadly obovate to somewhat 

rhomboidal leaves with a short base. 

Distichophyllum nigricaule differs from the present species 

in having smaller leaf cells, several rows of submarginal leaf 

cells distinctly smaller than the paracostal cells, and a well 

developed leaf costa reaching at least to midleaf. A New 

Guinean specimen (Koponen 36044, FH) distributed as D. 

gracilicaule is instead D. subnigricaule, a new record for New 

Guinea. 

The short leaf costae of D. subnigricaule are reminiscent of 

Distichophyllidium Fleischer. The latter genus is defined as 

having, among other characters, a papillose peristome. Since 

the sporophytes of D. subnigricaule are unknown, we have 

maintained it in Distichophyllum. 

84-252b (CAHP). 

Distichophyllum torrile 

Distichophyllum tortile Dozy & Molkenboer ex Bosch & Lacoste, Btyol. Jav., 

2:21, 1862. [Qpes: Java, Gedeh, Eysmam s.n.; Banca, sine loco. Amann 

s.n. (L). not seen.] 

Plants large, as in D. osterwaldii. Stems to 4 cm long and 

6-10 mm wide including the leaves. Leaves strongly crisped 

when dry, oblong-lingulate to slightly obovate, 4-5 mm long 

and 1-1.5 mm wide, roundly obtuse above, apiculate to 

mucronate, the acumina 27-60 pn long. Leaf borders strong, 

with 2-3(-4) rows of linear cells, 30-42 pn wide, somewhat 

wavy even in fresh state. Costae slender,reaching 2/3 of the leaf 

length or shorter. Leaf cells large, 45-60 pn wide, lax, 

thin-walled. Calyptra smooth above, fringed below. Setae 

papillose, less than 1 cm long. 

SPECIMEN STUDIED.-PHILIPPINES. Luzon: Bataan Prov., 

Upper Lama0 River, Williams 851 (US). 

REMw.-This distinctive species has large plants and 

leaves with large, thin-walled cells in the upper half. It is rather 

uncommon in the Philippine rainforests, and is also known 

from Indochina, the Malay Peninsula, Java, and Borneo. It most 

often occurs on wet rocks in deeply shaded sites. 

Genus Hookeria 

Hookeria S.E. Smith. Trans. J..inn. SOC. hdon. 9:275, 1808, nan. ms. 

[Qpe: H. lucens (Hedwig) Smith.] 

This is the type genus of the family Hookeriaceae. It is easy 

to recognize in the field by its thin, flat, somewhat transparent 

leaves and its large, net-like areolation. Only one species is 

known from the Philippines. 

Hookeria acutffolia 

Hookeria acu~ifolia Hooker & Greville. Edinburgh J. Sci., 2:225,1825. [?Lpe: 

Nepal. Hooker (K).] 

Plants mat-forming. Stems medium size, 2-3 cm long, pale 

green, simple or branched, mostly prostrate on ground. Leaves 

complanate, ovate to lanceolate, broadly acuminate, not 

bordered, slightly wrinkled when dry. Costa none. Leaf cells 

oval to rhomboidal, 42-75 pn long and 18-24 pn wide, 

thin-walled, smooth, becoming short-rectangular on the margin 

and laxer at base. Leaf apex with a cluster of short oval cells 

(27-30 prn long) often producing rhizoidal gemmae. Perichaetial leaves small, half the size of the vegetative leaves. Calyptra 

mitrate, covering the rostrate operculum, slightly tom at base. 

Seta lateral, smooth, 1-2 cm long. Capsules ovoid-oblong, 

inclined. Peristome teeth with median fmw on the exostome, 

cilia on the endostome absent or occasionally with a few cells. 

SPECIMENS STUDIED.-PHILIPPINES. Batanes Is.: Mt. Iraya, 

Tan 84-162 (CAHP). Luzon: Mt. Pulog, Tun 82-212 (CAIB); 

Bicol Region, A. de Jesus s.n. (PNOC Herb.). Negros: Mt. 

Cuernos, Tun 81-142 (CAHP). Mindanao: Mt. Apo, Tun 

82-246 (CAHP); Mt. Hilong-Hilong, Cabadbaran, Tan 84-510 

(CAHP). 

Rmms.-In the Philippines, this temperate and pantropical species is widespread in both lowland and montane 

rainforests. Its discovery dates only from 1%5, when it was 

reported by Shin from Batan Island at the northern end of the 

archipelago. 
NUMBER 75 25 

Genus Hookeriopsis 

Hookerwpir (Bescherelle) Jaeger,Ber. S. Gall. Naturw. Ges., 1875-1876358, 

1877. 

The genus Hookerwpsis is in need of a world monograph. 

Over the years, its species composition has grown too large and 

too diverse to represent a namal grouping. 

The history of the genus was recently outlined by Buck 

(1987) who redefined the taxon to include only the two original 

Lesser Antillean species first proposed by Bescherelle as a 

section of Hookeria. The remaining species have been 

reassigned by Buck (1987) to three genera: Thamniopsis 

(Mitten) Fleischer, Trachyxiphiwn Buck, and Brymela Crosby 

& Allen. In our opinion, Brymela is a worthwhile segregate. 

The separation of Hookeriopsis from Thamniopsis, however, is 

not convincing, nor is the segregation of the new genus, 

Trac hyxiphium. 

Because the genus consists of only two species in the study 

area and a few more in the tropical Asiatic region, we are 

content to stay with the traditional treatment in the absence of 

a satisfactory body of evidence to do otherwise. For those who 

wish to follow the generic revision of Buck (1987), the two 

Philippine species should be placed in Thamniopsis. 

Key to the Philippine Species of Hookeriopsis 

Apices of mature, lateral leaves long-acuminate; the upper leaf cells mostly elongate, with 

a ratio of length to width 6-8:l ...................... H. wichurae 

Apices of mature, lateral leaves broadly acute or short-acuminate; the upper leaf cells 

mostly narrowly rhomboidal to oblong, with a ratio of length to width 3-5:l ..... .................................... H. utacamundiana 

Hookeriopsis utacamundiana 

Hookerwpis utace (Montagne) BrotheNs,Nat W., 1(3):942,1907. 

Hookeria ukacamwuiiana Montagne. Ann. Sc. Nat. Bot., ser. 2,17:247,1842. 

[Type: India, Months Nilgheriensibus. Prope Ootacamund. Perrottet s.n., 

Hookeria secwrdo Griffith,Calcuaa J. Nat. Hist.3:280.1843, syn. nov. [1spe: 

India, Mumbree, Gr@th s.n. (isotype, NYI).] 

Hookeria purpwea C. Muller, Flora. 82:459, 1896, fide Buck (1987). W: 

Hawaii, sine loco spec.. Hillebrand s.n. (B, destroycd),not seen.] 

Hookeriopir geminidens B&N.S. Philipp. J. Sci., 5C (BoL): 156.1910, syn. 

nov. ['l)ps: Philippines. Luzon. Benguet, Eacani s.n., For. Bur. 15929 

(holotype., HI; isotypes, NYI, FHI).] 

Lpidopilwn swnatranwn Bosch & Lacoste, Bryd. Jav.,242,1862,fide &on 

(1916). [w: Indonesia. Sumatra. monte Simpai, Kwfhah, Herb. Ludg. 

Bat. 898.20-113-17 (Iectotype,L).] 

Plants mat-forming, silky green, with deep purplish tinge. 

Primary stems prostrate, complanate; the secondary stems and 

branches creeping or erect, to 10 mm tall. Leaves flatly 

arranged, slightly flexuose when dry, concave when wet. 

Lateral leaves ovate-oblong, 2-3 mm long, broadly acute; 

marginal borders not well differentiated, except for the strongly 

bifid, unicellular teeth along leaf margins near the apex. Costae 

double, strong, asymmetrically placed at leaf base, reaching 

mid-leaf, spinose abaxially. Dorsal and ventral leaves like the 

lateral ones but smaller, the apices sometimes acuminate. Leaf 

cells narrowly rhomboidal, (36-)45-60(-90) pn long and 

12-15 pm wide, smooth or at times slightly prorulose, 

becoming oval near apex and rectangular at base, thin to 

thick-walled. All collections studied are sterile. 

According to Fleischer (1908) and Gangulee (1977), the 

calyptrae are mitrate and hairy, the setae smooth, the capsules 

(isotype. NYI).l 

ovoid-oblong, horizontal, and the exostome teeth crossstriolate at base, papillose above, with distinct median furrow. 

SPECIMENS STUDIED.-PHILIPPINES. Mindanao: Mt. Kitanglad, Tan & Nuvarez 84-379 (CAHP, US). Mindoro: Mt. 

Halcon, Tan 87-048 (CAHP). SRI LANKA. Hanasgiriya, Alston 

1564, ex herb. Peradeniya (US). NEW GUINEA. Port Moresby, 

Carr 14185 (US). 

RmARKs.-Hookeriopsis utacamundiana is a widespread 

species in tropical Asia, reaching Hawaii in the Pacific 

(Bartram, 1933 as H. purpurea). The leaf marginal teeth, the 

length of the costae, as well as the leaf areolation can be highly 

variable. Consistent characters include the presence of at least 

some deep purplish coloration, the broadly acute to obtuse leaf 

apices, and the bifid marginal teeth near the leaf apex. Dorsal 

and ventral leaves often assume a variety of shapes and in some 

cases may develop weak marginal dentation in combination 

with acuminate apices. Such atypical forms may be confused 

with H. wichurae. 

In comparing the species to H. geminidens, Brotherus (1910) 

described H. utacamundiana as having only weak dentation on 

the leaf margins and a dioicous sexuality. However, Fleischer 

(1908) and Dixon (1916) reported that Javan and Indian 

populations of the latter are monoicous or autoicous. In 

addition, the leaf marginal dentation in the type of H. 

geminidens was observed by us to vary significantly. In the 

case of Hookeriopsis sumatrana and H. utacamundiana, the 

two differ only in the extent of their purplish coloration, angle 

of leaf insertion, and sexuality, characters not good for 

diagnostic purposes. 

Likewise, the differences between the Indian taxon, H. 
26 SMITHSON" CONTRIBUTIONS TO BOTANY 

secundu, and the widespread H. utacamundiana, as illustrated 

in Gangulee (1977),and Iwatsuki and Mizutani (1972), involve 

the above mentioned variable characters. Buck (1987) recently 

suggested the synonymy of H. geminidens and H. secundu, but 

does not consider them synonymous with H. utacamundianu. 

On the other hand, Dixon (19 16) proposed that H. sumatrana 

be treated as a synonym of H. utacamundiana. We consider all 

these taxa synonymous. 

Although we were not able to study the Hawaiian type of H. 

purpurea, we have seen many collections from Hawaii and 

agree with Buck (1987) that it is a synonym of H. 

utacamundiana. 

Hookerwpsis wichurae 

Hookorbpsis wichurae Fleischer, Musci Fl. Buitenzorg. 3: 1032.1908. "Islpes: 

Indonesia, West Java, am Berg Malabar bei Bandoeng, Wichwu s.n.(lectotype, FHI); am Gedeh, obehalb TjiMas bei Huis, Fleirchr s.n. (syntype, 

FH!).] 

Plants mat-forming, often deeply purplish. Stems 1-2.5 cm 

long, laxly branched. Leaves shrunken when dry, slightly 

plicate, ovate-lanceolate, long-acuminate, 1-1.75 mm long and 

less than 1 mm wide. Leaf margins weakly bordered with a row 

of shorter cells, toothed above, with bigeminate ends. Costae 

double, strong, asymmetrically placed at leaf base, reaching 

midleaf or beyond, spinose abaxially. Leaf cells narrowly 

fusiform to elongate, thick-walled, slightly pitted. Calyptrae 

mitrate, covering only the upper portion of the capsule, 

laciniate at base. Setae to 2 cm long. Operculum rostrate. 

REMARKS.-T~~ Malesian Hookzriopsis wichurae is very 

similar to H. utacamundianu, except that the leaf apices are 

long-acuminate, the leaf cells more elongate, and the leaf 

marginal teeth weakly developed and rarely bifid. Thus far, the 

differences between the two species appear to be consistent in 

the few specimens examined by us. Fleischer (1908) has 

illustrated H. wichurae. 

The first and only Philippine record of this species was 

reported by Tan and Iwatsuki (1983) from Mt. Cuernos (= Mt. 

Talinis) on Negros Island in Central Philippines. 

Genus Leskeodon 

hkeodon Brotherus,Nat. Pfl., 1(3):925,1907. [Type: not selected. See below 

IE L. a~atur (c. MUk) Brothetus.] 

The genus shows a combination of characters that calls into 

question the recent redelimitations of the family Hookeriaceae. 

Its leafy gametophyte closely resembles Distichophylhm but 

the sporophytic characters such as the non-collenchymatous 

exothwial cells and the papillose exostome without a median 

furrow are definitely not distichophylloid. Without capsules, 

specimens of Leskeodon can be difficult to place generically. 

The enigma of Distichophyllum and Leskodon is a good 

example of the discordant evolution of the gametophyte and 

sporophyte. A parallel situation exists in another generic pair, 

Distichophyllum and Distichophyllidium. As mentioned by 

Norris and Koponen (1985), these related generic pairs, which 

have almost identical gametophytes but radically different 

sporophytic designs, cast doubts on the appropriateness of the 

sporophyte oriented philosophy expressed by Crosby (1974) 

and Men et al. (1985) in their familial classification of 

hookeriaceous genera. 

Recently, Noms and Koponen (1985) described a species 

from Papua New Guinea, Distichophyllum leskodontoides, 

that has leaves similar to those of Leskeodon acuminatus, but 

has a distichophylloid peristome. The specimen may indicate a 

plausible intergeneric hybridization between Distichophyllum 

and Leskodon, resulting in the odd combination of gametophytic and sporophytic characters. Already there are a few 

intergeneric hybrids reported in literature (Andrews and 

Hermann, 1959; Rushing and Snider, 1985). In all cases, the 

structure of the diploid sporophytic body is the one affected by 

the cross-fertilization process. Of course, theoretically, the 

characters of the gametophytes could also be recombined in the 

hybrid sporophyte. Indeed, this is one area of cytogenetics and 

developmental morphology in bryophytes that has not been 

critically investigated. 

In Southeast Asia, species of Leskeodon tend to be small to 

medium-size plants growing to 2 cm long. The somewhat 

complanately arranged leaves nearly always have a strong and 

firm border above. When dry, the leaves have a stiff appearance 

with a wavy marginal contortion. The upper laminal cells of 

Leskeodon are mostly hexagonal to subhexagonal-rhomboidal 

with sharp angles. They are longer than wide, moderately 

thick-walled, and arranged in a vertically oriented pattern along 

the leaf costa. They are never round-hexagonal and thin-walled 

as in most Distichophyllum species. In our opinion, the leaf cell 

areolation of Asiatic Leskodon is more similar to that of 

Calyptrochaeta than Distichophyllum. 

However, when the New World species of Leskodon are 

studied and compared, two basic patterns of leaf cell areolation 

emerge. One of which, represented by L. andicola (Spruce) 

Brotherus and L. mariei (Bescherelle) Brotherus, approximates 

the leaf cell areolation of the Old World taxa. The other group, 

L. longipilus (Bescherelle) Bartram and L. cubensis (Mitten) 

Thbriot, has basically large, round, thin-walled cells with a 

weak costa, much more reminiscent of the members of 

Distichophyllidium from Southeast Asia. More strikingly, the 

weakly collenchymatous exothecial cells and the fragile, 

papillose peristomes seen in L. longipilus (Figures 55-60) also 

closely resemble those of Distichophyllidium, especially when 

seen under the SEM (Figures 61-63). We have not been able to 

examine mature capsules of L. andicola and its related taxa to 

decide on the proper disposition of these neotropical taxa in 

relation to Distichophyllidium and the Old World taxa of 

Leskodon. It appears that a proper typification of the genus 

will be of critical importance in the light of the above described 

variations of leaf cell patterns. The species emphasized in the 

original description of the genus (Brotherus, 1907), L. auratus 
NUMBER 75 27 

(C. Muller) Brotherus from Trinidad (also reported from 

Dominica, Bartram, 1955) is unfortunately not well known. In 

any case, the genus, as defined today, is doubtless a 

heterogenous group. 

Bartram (1961) described L. robbinsii from New Guinea 

with a rather atypical leaf cell areolation for the genus. The 

isotype (New Guinea, Wabag area, Robbins 31 91, B!) has laxly 

imbricate, ovate-lanceolate leaves with a short apiculate apex. 

The leaf cells are more or less homogenously rhomboidal to 

subrectangular and thin-walled. The leaf costa is percurrent and 

the leaf margins are revolute, and have a narrowly differentiated border with only two rows of linear cells. We think that the 

species is best placed in Daltonia, Daltonia robbinsii (Bartram) 

Tan & Robinson, comb. nov. (Basionym: Leskeodon robbinsii 

Bartram,Rev. Bryol. Lichenol., 30200,1961.) 

There are three species of Leskeodon known from the 

Philippines. Two of these, Leskeodon acuminatus and L. 

brevicwpidatus, are new additions to the local moss flora. 

Key to the Philippine Species of Leskeodon 

1. Leaf borders narrow, less than 20 pm wide, with 1-2 rows of linear cells; leaf cells 

large, mostly more than 30 pm in diameter; leaf apices gradually long acuminate .................................. L. philippinensis 

Leaf borders wide, more than 20 pm wide, with 3-5 rows of linear cells: leaf cells 

smaller, mostly less than 25 pm in diameter: leaf apices abruptly narrowed into a 

long acumen ................................... 2 

2. Leaf costae percurrent or ending 15 or less cells below the apex; leaf borders strong, 

especially near the base, 30-60 pm wide; leaf acumina stout, 160-280 pm long 

and 48-72 pm wide at base .................... L. acuminatus 

Leaf costae not percurrent, ending well below the apex; leaf borders narrower, not 

well differentiated at base, 21-33 pm wide; leaf acumina slender, 240-480 pm 

long and 32-56 pm wide at base ............... L. brevicuspidatus 

Leskeodon acuminatus 

Leskeodon acwninafus (Bosch & Lacoste) Fleischer, Musci Fl. BUitenz.org, 

Dktichophyllwn acwninatwn Bosch & Lacoste, Bryol. Jav., 2:26,1862. [Type: 

3:971,1908. 

Java,Mt. Gedeh,Teymann s.n. (L),not seen.] 

Plants large, in dense mats, at times tufted. Stems 2-3 cm 

long and 4-6 mm wide including the leaves. Leaves 

complanate and slightly crisped when dry, 3-4 mm long and 

about 1.5 mm wide, long-acuminate to cuspidate at apex, the 

acumina stout, semi-twisted, 160-280 pm long and 48-72 pm 

wide basally. Leaf borders strong throughout, to 60 pm wide, 

with 2-4(-5) rows of thick-walled linear cells. Costa single, 

reaching mostly */3 of the leaf length, at times percurrent. Leaf 

cells sharply subhexagonal, (15-)18-24(-30) pm long, 

slightly longer than wide, becoming isodiametrically hexagonal toward margin and rectangular near base. Perichaetial 

leaves small, 1 mm long, acute-acuminate, weakly bordered, 

and shortly costate. Calyptra deeply fringed at base. Setae to 2 

mm long, smooth below and scabrous above, Capsules ovoid, 

to 1 mm long, exothecial cells non-collenchymatous. Operculum conic-rostrate. Exostome teeth papillose. 

SPECIMENS STUDIED.-PHILIPPINES. Mindanao: Mt. HilongHilong, Agusan Prov., Tan & Navarer 84-399 (CAHP, US). 

MALAYSIA. Borneo: Mt. Kinabalu, Bryotropica 4937 (B). 

REMARKS.-Leskeodon acuminatus is a new addition to the 

Philippine moss flora. Filiform brood bodies were observed by 

us on Bornean specimens, but not on the Philippine plants. The 

species is presently known from all the major islands of 

Malesia. 

Leskeodon brevicuspidatus (Bartram) Tan & Robinson, new 

combination 

ROWS 3 1-34.51-54 

Distichophyllum brevicuspidatwn Bartram, Brittonia, 9:48, 1957. basimym. 

[Type: New Guinea, Mt. Dayman, epiphyllous, Brass 23069 (holotype, 

FH!).] 

Plants large, stems to 2 cm long, densely foliate. Leaves 

somewhat twisted when dry, erect-spreading when wet, 

oblong-obovate to broadly spathulate from a short base, 

2.5-4.0 mm long and 1-2 mm wide, rounded obtuse above, 

cuspidate, the acumina slender, often twisted, 240-480 pm 

long and basally 32-56 pm wide. Leaf borders entire, strong in 

the upper half, less markedly differentiated near base, with 

2-3(-4) rows of thin-walled linear cells, 21-33(-45) pm 

wide. Costae ending well below the apex. Leaf cells sharply 

subhexagonal, slightly longer than wide, (12-)15-21(-36) pm 

long, becoming smaller near the margin and rectangular toward 

the base. Calyptra fringed at base. Setae 2 mm long, smooth. 

Capsules erect, ovoid, less than 1 mm long; exothecial cells 
28 SMITHSO" CONTRIBUTIONS TO BOTANY 

FIOUW 29-34.4istichophyllum and Lcskodon: 29,30, Disiichophyllwn cwpida~wn (Philippines: Sihyan 

Is., Mt. GitingGiting. Tan & Hernaez 87-528, US): 29, upper leaf cells; 30, whole leaves. 31-34, Lcskcodon 

brevicurpi&arur (Philippines: Mindanno, Mt. Hilag-Hilong, Tan & Navarea 84-471, US): 31, plant, wet; 32. 

plant, dry; 33, whole leaves; 34, upper leaf cells. 
NUMBER 75 29 

quadrate, thin-walled, non-collenchymatous. 

SPECIMENS STUDIED.-PHILIPPINES. Mindanao: Mt. HilongHilong, Agusan Prov., Tan & Navarez 84-471 (CAHP, US). 

PAPUA NEW GUINEA. Timin River, Sau Valley, Robbins 2873, 

sub Distichophyllum brevicuspidatum (B). 

RmARKs.-Leskeodon brevicuspidatus was described as a 

Distichophyllum by Bartram (1957). In the protologue, the 

author mentioned the capsule but failed to notice the nature of 

the exostomial ornamentation. The holotype specimen at FH 

has two capsules showing the papillose exostome. A second 

collection from New Guinea (Robbins 2873) bears mature 

capsules having a papillose exostome without a median furrow. 

The differences between the present species and L. acuminatus are outlined in our diagnostic key. The Philippine 

specimen collected from Mindanao compares well with the 

type at FH, except that the leaves are smaller and have longer 

leaf acumens ranging 153-300 pm long. 

Leskeodon brevicuspidatus is a new addition to the 

Philippine flora. It is known from Papua New Guinea and 

Mindanao, Philippines. 

Leskeodon philippinensis 

Leskeodon philippinensis Brotherus, Philipp. I. Sd.. 13 C (Bot.):213, 1918. 

[Type: Southern Philippines, Biliran Island, Mudregor 18468 (holotype, 

H!).] 

Plants large, light green in color, in flat, dense mats. Stems to 

2.5 cm long, densely foliate, about 5 mm wide including the 

leaves. Lateral leaves elliptic-spathulate, entire, 4-5 mm long 

and 1.0-1.5 mm wide, gradually long-acuminate at apex, the 

acumina 90-120 p long, often twisted. Costae long, nearly 

percurrent. Leaf borders very narrow for the genus, consisting 

of 1-2 rows of linear cells, not well differentiated at leaf base. 

Leaf cells hexagonal, mostly large, 25-30 p in diameter, 

thin-walled, with only a few smaller submarginal cells. 

Calyptra deeply fringed at base. Setae short, 1-2 mm long, 

smooth below, papillose above. Capsules small, ovoid, about 1 

mm long. Peristome teeth papillose. 

REMARKs.-Brotherus (1918), when describing Leskeodon 

philippinensis compared the species to L. acuminatus and 

considered them to differ in “foliorum forma cellulisque 

laxioribus.” Indeed, the leaf cells of L. philippinensis are much 

larger and thinner-walled. Later, Brotherus (1925) reported that 

L. philippinensis was synoicous and L. acuminatus was 

heteroicous or autoicous. This difference in sexuality does not 

hold true, since we have seen a synoicous specimen of L. 

acuminatus from Mt. Kinabalu, Borneo. We think the two taxa 

are best separated by leaf shape. In L. philippinensis, the leaves 

are elliptic-spathulate with both ends gradually becoming 

narrower. Bartram (1939) provided a good sketch of this 

peculiar leaf form. The leaves of L. acuminatus are typically 

spathulate with a greatly expanded upper half that becomes 

abruptly cuspidate. 

This Philippine endemic is known only from the type 

collection in Helsinki. The specimen labelled L. philippinensis 

at the Philippine National Herbarium in Manila is cited from 

Biliran Island, but no collector is given, and the specimen is 

apparently not an isotype. The packet contains material of L. 

acuminatus. 

Notes on Philippine Chaetomitrium 

The genus Chaetom’trium is unique among the hookeriaceous 

taxa in the following combined gametophytic features: unbordered leaves with short double costae, narrowly oblong to 

elongate, thick-walled and prorate leaf cells, and an enlarged 

perichaetium with often erose or sharply toothed leaf margins. 

Each of the marginal teeth of the perichaetial leaves consists of 

a long cell with a bigeminate tip, often projecting perpendicularly from the leaf margin. 

The exostome structure of Chaetomkium is also atypical for 

the Hookeriaceae in which it has traditionally been placed. 

They appear cross-striolate under the light microscope, but the 

ornamentation of the outer exostomial plate actually consists of 

irregularly and incompletely formed folds or ridges studded 

with papillae Figures 39-42 ). This type of ornament is more 

commonly seen among Hypnalean taxa (Higuchi, 1985). 

Dixon (1935) stated that Borneo (13 species) and New 

Guinea (14 species) represent the two “headquarters” of this 

remarkable genus. The Philippines, with 17 species, would 

seem to be a third center of local distribution. 

Among the local species, Chaetom‘trium seriatum Brotherus 

ex Bartram had been considered by Mohamed (1985) to be a 

synonym of the Bornean C. borneense Mitten. Our study of the 

types of these two taxa has led us to conclude otherwise. The 

Philippine specimens differ consistently from C. borneense in 

lacking an acute to shortly acuminate leaf apex. Perichaetial 

leaves of Philippine specimens are weakly toothed whereas the 

perichaetial leaves in the Bornean species are strongly toothed. 

The calyptra of C. seriatum is only sparsely fringed at the base, 

quite unlike that of C. borneense. The seta of C. seriatum is 

also longer than that of C. borneense. Apparently, Mohamed 

(1985) was overly influenced by the spirally ranked pilotrichelloid foliation shared by the two species. 
30 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Key to the Philippine Species of Chaetomitrium 

In the key, which is based as much as possible on the gametophytic characters, a few muplets lead to two 

species. This implies that the taxa involved may prove to be synonymous when their respective types are 

examined. 

1. 

2. 

3. 

4. 

5. 

6. 

7. 

8. 

9. 

10. 

11. 

12. 

13. 

14. 

15. 

Leaf cells strongly tuberculate or sharply spiculose ................ 2 

Leaf cells smooth, prorulate, or at most prorate. ................. 5 

Papillae of leaf cells tuberculate or knob-like; setae more than 1.5 cm in length . , 

...................................... C. elmeri 

Leaf apices broadly rounded, at times with a small apiculus; calyptra cucullate . . Papillae of leaf cells spiculose; setae less than 1.5 cm in length ......... 3 

................................... C. perarmatum 

Leaf apices acute to acuminate; calyptra mitrate ................. 4 

Leaves ovate to short-lanceolate, the apices acute, slightly constricted near the tip; 

setae papillose ............................ C. leptopoma 

Leaves lanceolate, the apices acuminate, often strongly constricted near the tip; 

setae setose ................... C. beccariilC. orthorrhynchum 

Leaf apices mostly truncate, broadly rounded or bluntly obtuse, at times abruptly 

terminated in a small apiculus .......................... 6 

Leaf apices acute to acuminate .......................... .9 

Branch leaves spirally seriate in arrangement. .................. 7 

Leaf apices short apiculate or bluntly obtuse, not recurved ..... C. schojieldii 

Branch leaves not seriately arranged. ....................... 8 

Leaf apices rounded to broadly obtuse, if apiculate, the apiculus very small, 

recurved ............................... C. warburgii 

Upper half of the leaf margins strongly wavy, entire, or serrate; leaf apices nearly 

........................ C. warburgiilc. pseudo-elongatum 

Upper half of the leaf margins undulate, or constricted, forming a throat-like apex; 

leaf apices acuminate, strongly armed marginally with long, unicellular, uncinate 

teeth.. ................................ C. everettii 

Branches complanate ............................... 10 

Branches tumid, not complanate ......................... 11 

Leaves concave when wet, with apices constricted, forming a somewhat flaring 

throat-like region ............................ C. weberi 

Leaves flat or slightly concave when wet, without any apical constriction ..... ...................................... C. elegans 

Upper half of leaf margins undulate, strongly toothed with long, sharp, and uncinate 

teeth .................................. C. everettii 

Upper half of leaf margins plane, or slightly rugose, entire, sermlate or serrate . 12 

orbicular, often recurved, muticous or apiculate .................. 

Leaf apices without a constriction, or a flaring throat-like region ........ 13 

Leaf apices constricted, forming a throat-like region ............... 14 

margins entire or weakly sermlate; calyptrae mitrate ...... C. jimbriatulum 

Plants small, lateral branches to 5 mm long and 1 mm wide; leaf apices acute; leaf 

Plants larger, lateral branches more than 5 mm long and wider than 1 mm; leaf 

apices long-acuminate; leaf margins slightly irregularly serrate; calyptrae 

cucullate .............................. C. lanceolatum 

Calyptra cucullate, not or scarcely fringed at base ......... C. lanceolaturn 

Calyptra mitrate, fringed at base ......................... 15 

Leaf apices mostly acuminate to long-acuminate ................ 16 

Leaf apices mostly acute to short-acuminate ................... 17 
NUMBER 75 31 

16. Leaf margins irregularly serrate; leaf cells sharply prorate; sew setose above ... ................................. C. orthotrhynchum 

Leaf margins regularly sedate; leaf cells prorulose; setae papillose above .... ................................... C. papillvolium 

17. Leaves oblong-lanceolate, widest at midleaf or below, slightly rugose; leaf cells 

distinctly prorate. ............................... .18 

Leaves oblong-oblanceolate, widest above midleaf, strongly rugose; leaf cells 

sm ooth...................................... 19 

18. Setae short, less than 10 mm, strongly papillose above ..... C. philippinense 

Sew long, more than 10 mm, slightly papillose above ......... C. weberi 

19. Leaf margins regularly toothed, setae papillose above ........ C. elongatum 

Leaf margins irregularly toothed, with larger teeth near the throat-like constricted 

region; setae spinose-hispid above ................. C. laevifolium 

Chaetomitrium schojieldu Tan & Robinson, new name 

FTOURES 39-42 

Chaeiomiirium seriaium Brotherus ex Bertram, Pwp. J. .%., 68:271,1939. 

hom. illeg. non C. serkium Brotherus,Ba Jahrb. 1913. 

Because the name C. seriatum as applied to the Philippine 

species is an illegitimate homonym, we propose to replace it 

with C. schofieldii, in honor of Prof. W.B. Schofield, an 

eminent Canadian bryologist and former professor of the first 

author at the University of British Columbia. 

ACCEPTED NAMES OF PHILIPPINE Chaetom’frium 

An asterisk (*)indicates a new record for Philippine moss flora. 

C. beccarii Dixon 

* C. elegans Geheeb 

C. elmeri Brotherus 

* C. elongatwn (Dozy & Molkenboer) Dozy & Molkenboer 

* C. everettii Mitten ex Dixon 

* C. fimbriatulum (Dozy & Molkenboer) Bosch & Lacoste 

C. laevifolium Dixon ex Bartram 

C. lanceolatwn Bosch & Lacoste 

C. leptopoma (Schwaegrichen) Bosch & Lacoste 

C. orthorrhynchum (Dozy & Molkenboer) Bosch & Lacoste 

C. papillifolium Bosch & Lacoste 

C. perarmatum Brotherus 

C. philippinense (Montagne) Bosch & Lacoste 

C. pseudo-elongatum Brotherus 

C. schofieldii Tan & Robinson, nom. nov. 

C. wurburgii Brotherus 

C. weberi Brotherus 
Literature Cited 

Akiyama, H. 

1988. 

Men, B.H. 

Bryophytes. In M. Kato, editor, Taronomic Studies of tk Plonts of 

Serum Island, pages 13-59. Tokyo: University of Tokyo. 

1987% A Revision of the Genus Lcucomium (Leucomiaceae). Memoirs of 

1987b. HooAeriopsis crosbyi (Musu. Hookeriaceae): A New Species h 

tk New York Botanical Gorden, 45661-677. 

Western Panama. The Bryologirt, 90:45-47. 

A Revision of the Genera Pilotrichidium and Diplonewon (Musck 

Hookeriaceae). JOWMI of the Hottori Botanical Laboratory, 

Allen, B.H., and M.R. Cmby 

1986. 

61 :45-64. 

Men,B.H., M.R. Cmby. and R.E. Magill 

Andrews, A. LeRoy, and F.J. Hennann 

Bartram,E.B. 

1985. A Review of the Genus Stedictyon. Lindbergio, 11:149-156. 

1959. A Natural Hybrid in the Ditrichaceae. The Bryologist, 62:119-122. 

1931. 

1933. 

1939. 

1942. 

1944. 

1955. 

1957. 

1%1. 

A Review of the American Species of Doltonio. Bulletin of he 

Twrsy Botanical Club, 48:31-48. 

Manual of Hawaiian Mosses. Bernice P. Bishop Museum Bulletin, 

Mosses of the Philippines. Philippine JOWMI of Science, 68:l-423. 

Third Archbold Expedition, Mosses from the Snow Mountains. 

Netherlands New Guinea. Llo$io, 5245-292 

New and Noteworthy Philippine Mosses. Forlowio, 1503-513. 

Mosses of Dominica. British West Indies. BulIetin of tk British 

Museum (Notwol History) Botany, 2(2):37-49. 

Mosses of Eastern Papua, New Guinea. Brittonio, 9:32-56. 

Mosses of the Western Highlands of Eastern New Guinea. Revue 

Bryologque et Licheologque, 30:185-207. 

101:l-275. 

Brotherus, V.F. 

1907. 

1909. 

1910. 

1913. 

1918. 

1925. 

Hookeriaceae. In A. Engler and K. htl, Die naturlichen

Pflanzenfomilien, pages 227-229. Leipzig. 

Musci Novi Philippinenses I. Lcojlee of Philippine Botany, 

Contributions to the Bryological Flora of the Philippines. IIL 

Philippine JOWMI of Science, SC (Botany):137-162. 

Conntributions to the Bryological Flora of the Philippines, IV. 

Philippine JOWMI of Science, 8C (Botmy):65-98. 

Contributions to the Bryological Flora of the Philippines. V. 

Philippine JOWMI of Science, 1X (Botany):201-222. 

Hodreriales. In A. Engler and K. Prantl, Die Mtiirlichen Pflanzenfomilien. 2nd edition, Volwne 11,poges 214-278. kipzig. 

2:65 1-658. 

Buck, W.R. 

1987. Taxonomic and Nomenclatural Rearrangement in the Hookeriales 

with Notes on West Indian Taxa. Brittonio, 39:210-224. 

1986. Suggestions for a New Familial Classification of Pleurocarpous 

Buck, W.R., and D.H. Via 

Mosses. TOXO~, 35:21-60. 

Cmsby. M.R. 

1968. Proposal for the Conservation of the Name Doltonio Brid. 

(Musci-Hderiaceae). Taon, 17:443,444. 

1972. PterygophylIwn Brid., Nmen Rejiciendum et Illegitimum. T'n, 

21 :205-212. 

1974. Toward a Revised Classification of the Hookeriaceae (Musci). 

JOWMI of tk HottoriBotonicalL4borotory, 38:129-141. 

1975. Lectotypification of Schizomitrium B.S.G. (Musci, Hookeriaceae). 

Tun, 24~353-355. 

Cnrm. H.A. 

1984. Notes on Tropical American Mosses. The Bryologist, 87:203-216. 

Dixon, H.N. 

1916. Miscellanea Bryologica, V. JOWMI of Botany, 54:352-359. 

1922. Some New Genera of Mosses. Jownol of Boiany, 60:101-110. 

1935. A Gmtributiim to the Moss Flora of Borneo. Jownol of Linneon 

Society, Botany, 5057-140. 

Dozy. F.. and J.H. Molkenboer 

Dozy, F., and J.H. Molkenboer (completed by R.B. van den Bod, and C.M. 

1845-1854. Musci Frondosi Inediti Archipelogi Indici. Leiden. 

van der Sande Lacoste) 

1855-1870. Bryologia Jovonica. 2 volumes. Leiden. 

1908. Die Murci &r Floro von Buitenzorg. volume 3. Leiden. 

1982. Perforierte Hydroiden bei Laubmoosen. Jowd of the Hottori 

Botanical Laboratory, 51:51-60. 

Fleischer, M. 

Frey. W., and U. Richter 

Gangulee, H.C. 

1977. Mmses of hiern India ond Mjocent Regwns. Fascicle 6 

(Hookeriales). Calcutta, India. 

Higuchi, M. 

1985. A Taxonomic Revision of the Genus Gdlonio Broth. (Musci). 

Jownol of the Hottori Botanical Laborotory, 59:l-77. 

Colored IIIustrolions of Bryophytes of Jopon. Japan: Hoikusha 

Publishing Co. 

1968. Some Interesting Mosses from the Philipphes. JOWM~ of the 

Hotiori Botanical Laboratory, 31 :50-54. 

1979. Checklist of Philippine Mosses. Kolikason, Philippine JOWMI of 

1980. Noteworthy Philippine Mosses at the Hattori Botanical Laboratory 

(NICH), Japan. Kalihon, Philippine JOWMI of Biology, 9267- 

282. 

Koponen T, and P. Isoviita 

1984. Proposal to Ccnserve Collicostello against Schizomitriwn (Musci, 

Hookeriaceae). Taron, 33:112,113. 

Miller. H.A. 

1971. An Overview of the Hookeriales. Phytologio, 21943-252. 

Margadant, W.D. 

1959. Typification and Conservation of Generic Names in Musci. Act0 

Boianico Neerlondica, 8:271-276. 

On the Identity of Chactomiiriwn borneense Mitt., C. seriotum 

Broth., C. seriotum Broth. ex Bartr. and C. elmeri Broth. Lindbergio, 

Iwatsuki, 2, and M. Mizutani 

1972. 

Iwatsuki, Z, and AJ. Sharp 

Iwatsuki, Z, and B.C. Tan 

Biology, 8~179-210. 

Moham4,M.A. Haji 

1985. 

11:80-82. 

Mohamed, M.A. Haji, and A. Damanhuri Mohamed 

1986. Four Genera of Mosses New to Peninsular Malaysia. JOWMI of 

Bryology, 14:333-338. 

Nishimura,N., M. Higuchi, T Seki, and H. Ando 

1984. Delimitation and Subdivision of the Moss Family Hypaceae. 

Jownol of the Hotiori Botanical Laborotory, 55~227-234. 

32 
NUMBER 75 33 

Noguchi, A. 

1937. Studies on the Japanese Mosses of the Orders Isobryales and 

Hookeriales, 11. Journal of Science of the Hiroshima University, 

series B, division 2 (Botany) 3:37-56. 

1952. Musd Japonici. I: Hypopterygiaceae. JOWMI of the Hattori 

Botanical Laboratory, 7: 1-22. 

1956. Musd Japonici, V: The genus Distichophyllwn. Journal of the 

Hattori Botanical Laboratory, 17:19-31. 

1972. Mosses of North Bomeo, I. JourmaI of the Hattori Botanical 

Laboratory, 36:455-486. 

Bryophytes from Frieda River, East and West Sepik Provinces. 

Papa New Guinea, L AnnaIes Botanici Fennici, 22:367-396. 

The Systematic Position of Bryobrothera crenulata. The Bryologh, 

Noguchi, A., and 2 Iwatsuki 

Norris, D.H., and T Koponen 

1985. 

Norris, D.H., and H. Robinson 

1979. 

82:305-309. 

Renauld, F., and J. Cardot 

1896. Mousses newltees BaJava par M.J. Massart. Revue Bryologique, 

23 :7-108. 

Robinson, H. 

1971. A Revised Classification for the Orders and Families of Mosses. 

1975. The Mosses of Juan Femandez Islands. Smitkonian Contributions 

1986. On the Relationships of the Hookeriaceae. The Bryological Times, 

35:2,3. 

A Natural Hybrid between Bruchia microspora Nog. and TrenvltoPhytologia, 21 ~289-293. 

to Botany, 27~1-88. 

Rushing, A.E., and J.A. Snider 

1985. 

don longicollis Michx. Monographs in Systenvltic BOranyfrom the 

Missouri Botanical Garden, 11:121-132. 

A Handbook of the New Zealand Mosses. Roycrl Society of New 

Zcaland Bulletin, 5:l-490. 

A Small Collection of Mosses from Isl. Batan, the Philippines. 

Sainsbury,G.O.K. 

1955. 

shin. T 

1965. 

Tan, B.C. 

1987. New Records of Philippine Mosses. Memoirs of the New York 

Botanical Garden, 45:6454. 

1983. Nineteen New Records of Philippine Mosses. Kaliknsan, Philippine 

Journal of Biology, 12328-350. 

Additions and Corrections for the Philippine Moss Flora. Crypogamie, Bryologie Lichendogie, 10:235-245. 

Musci Igoroti (Mosses of the Mountain Provinces. Luzon, Philippines). Kalikasan, Philippine Jour~I qfBWlogy, 7:187-210. 

1982. Notes on Mosses from Ceylon and India, 6 An Illustrated Key to 

Distichophyllwn in Ceylon and S. India. Journal of Bryology, 

1215-21. 

Hikobia, 4263-271. 

Tan, B.C., and Z. Iwatsuki 

Tan, B.C., and T Koponen 

1989. 

Tixier, P. 

1978. 

Townsend, C.C. 

Via, D.H., and M.R. Crosby 

1972. Achrophyllwn-A New Name for a Genus of Mosses. The 

Bryologist, 75 174,175. 

Philippine Mosses. Bulletin of the New York Botanical Garden, 

Williams, R.S. 

1914. 

8~33 1-377. 
34 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 35-38.4EM views of peristome of Calypirochaeia ramosa (Java: SchiJkr, Exsicc. Museo Hist. 

Natur. Vindobonensis 3885, US): 35, surface view of an exostome tooth showing the well developed dorsal (or 

outer) lamellae and the median furrow; 36, upper portion of exostome teeth showing the papillose Ornamentation; 

37, lateral view of the exostane tooth showing the equally well developed dorsal and ventral (or inner) lamellae 

at base, and longer ventral lamellae distally; 38, cross-section of an exostome tooth showing the dorsal lamellae 

and the stacks of lamellulae described as "striae"under the light microscope. 
NUMBER 75 35 

RouREs 39-42.fEM views of peristome of Chaeromirriwn schofieldii (Philippines: Butunn, Webcr 1496, (ype 

of C. serkiwn Brotherus ex Bartram): 39. surface view of exostane showing the well developed dard Lmcllw 

and the vertical median partitions; 40,lated view of exostome tooth showing strongly developed ventral lamellae 

from base to tip; 41, portion of the exostane tooth showing the dorsal lamellae and the incompletely formed striae 

or folds with papillae; 42, details of interlamellate folds or striae. 
36 SMITHSONTAN CONTRIBUTIONS TO BOTANY 

FIGURES 43-46.SEh4 views of penstane of Daltonia contorta (Philippines: Baguio, Williams 1666, US): 43, 

surface view of the exostome showing papillae clustered primarily almg the margins of the cell plates; 44. lateral 

view of exosme tooth showing the strong and papillose ventral or inner lamellae; 45. ventral or inner view of 

the exosme tooth; 46. details of lamellar papillae showing the composite and lugose surface. 
NU.M3ER 75 37 

FIGURES 47-50.4EM views of peristome of Actinodonriwn ascendens (Thailand: Nakhon Sawan Doi Musae 

Banana Plantation, Tow 8006, US): 47, peristome showing the partly intact annulus; 48, portion of exostOme 

tooth showing large papillae clustered along the margins of the cell plates; 49, ventral or inner view of the 

exostome tooth showing the thickened portion along the median partition line; 50, lateral view of the exostOme 

tooth showing the lack of well developed ventral lamellae. 
38 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

RffUm 51-54.-SEM views of peristome of Leskodon brevicurpidattcr (Phdippines: Mt. Hilong-Hilong. Tan 

& Navarez 84-471, US): 51,52, surface views of exostome teeth showing the papillae clustered along the margins 

of the cell plates; 53. details of lamellar papillae showing the complex organization; 54, lateral view of the 

exostome tooth showing the low ventral lamellae. 
NUMBER 75 39 

FIGURES 55-58.PEM views of peristome of Leskeodon Iongipilus (Dominica: St. George Parish, H. Robimon, 

22 Feb. 1964, US): 55. surface view of exostome tooth showing constrictions between cell plates and the dense 

surface papillae; 56, lateral view of exostome tooth showing the presence of ventral lamellae and absence of 

dorsal lamellae; 57, segment of exostome tooth showing the vertically aligned rows of large, coalesced papillae; 

58, details of the simple type of exostomial papillae arranged in horizontal rows. 
40 SMITHSON" CONTRIBUTIONS TO BOTANY 

FIGURES 59-63.fEM views of peristomes of Leskeodon and Distichophyllidiwn: 59.60, Leskeodon longipilus 

(Dominica: St. George Parish, H. Robinvon, 22 Feb. 1964, US): 59. details of the coalesced type of exostomial 

papillae near the edge of the exostome tooth; 60, cross-section of the exostome tooth showing the ventral 

lamellae; 61-63, Distichophyllidiwn nymanianwn Fleischer (Malaysia: Pahang, Mohamed & Zamzuri 11 10, US): 

61, surface view of exostome teeth showing the constrictions between the cell plates: 62, portion of exostome 

tooth showing the folds m the cell plates. and showing the small surface papillae; 63, lateral view of the exostome 

tooth at base showing the well developed ventral lamellae. 
Index 

(Synonyms and page numbers of principal accounts in italics) 

Achrqhyllum, 1.2,3 

dentatum, 3,4 

javense, 4 

quadrifarium, 4 

Actinodcntium,2,4,5, 11,13 

ascendens,5,6, figs. 47-50 

rhaph1dostegum,5,6 

Adelothecium, 2 

Brymela, 25 

Bryobrothera, 2,3,6 

crenulata, 6 

Bryodusenia, 2 

Callimsta, 6 

Calliwstella, 2,4,6 

beccariana, 7 

papillata, 6,7 

prabaktiana,6,7 

Calomnion complanatwn, 6 

Calomnion dironii, 6 

Calyptrochaeta, 1,2,3,4,7,15,26, 

figs. 1-10 

cristata, 7 

microblasta,3,9, figs. 7-10 

parviretis.9, 10,figs. 1.2 

ramosa, 3,9,10, 11,figs. 3,4,35-38 

remotifolia, 9,10, 11, figs. 5.6 

Chaetomitriopsis, 2,4 

Chaexomitrium. 2.3,4,29, 30 

beccarii,30,31 

bomeense, 29 

elegans,30,31 

elmeri, 30.31 

elOngaNm, 3 1 

everettii, 30.3 1 

fimbriatulum, 30,3 1 

laevifohm, 3 1 

lanceolatum, 30.3 1 

leptopoma, 30.3 1 

orthorrhynchum. 30.3 1 

papillifolium, 3 1 

perarmatum.30.31 

philippinense, 3 1 

pseudo-elongatum, 30.31 

schofieldii, 3,30,31, figs. 39-42 

seriatwn, 3.29 

warburgii, 30,3 1 

webe-ri. 30.3 1 

Cyathophorella, 1,2 

hcokeriana, 1 

Cyathophomm, 1,2 

Iimbatdwn, 10 

Cyclodictyon, 2.4,l I 

blumeanum. 11 

laetevirens, 11 

Daltonia.2.3,5,11, 13,figs. 11-14 

angustifolia, 11,12,13,fig. 12 

aristifolia, 11,13, fig. 12 

armata,3.11,13, fig. 11 

baldwinii, 13 

ccnmrta, 11,12,13, figs. 12.43-46 

revoiuia, 12 

robbinsii,27 

spiniseta, 13 

splachnoides, 11 

Dendrocyathophorum, 1,2 

linbatulwn, 10 

paradoxum, 1 

Dimorphocladon.2 

Distichophyllidium, 2,26 

Distichophyllum 2,3,11,13, 14,15,19, 

nymanianum,figs. 61-63 

23,24,26, figs. 15-22.29-34 

acwninatwn, 27 

brevicuspes,3,16,17 figs. 17.18 

brevicuspidatus,27,29 

catinjfolium, 16,17, 18.24 

cirratum, 20,22 

collenchymatosum, 16,17, 18 

cucullatum, 16,17,18 

cuspidaturn, 15,18,19,20,23, figs. 29,30 

denticulatum, 22,23 

elmeri, 3,22 

gracilicade, 3,20, 24 

haitorii, 17 

iwatsukii. 19 

jungermannioides. 15,16,24 

kinabaluense, 15,18, 19, figs. 19-22 

kaxirete, 23, 24 

leskeodcntoides, 26 

maibarae, 16,19, 22 

mittenii, 16,19,20,22 

monfagneanum, 19,20 

nigricaule,3,15,17.18,19,20,22,24, 

nigricaule var. drratum. 17,20,22, figs. 23,24 

nigricaule var. elmeri, 16,17,20,22, figs. 25.26 

nigricaule var. nigricaule, 16,17,20, 22, 

obtusifolium, 15,22,23 

ostenvaldii, 16.22.23, 24 

perundulatwn, 20, 22 

philippinense, 19 

rotundatum,23,24 

santosii, 3,15,23 

sinuosulwn, 19,22 

SubcarinaNm, 17 

subcuspidaturn, 15,17,23, fig. 15, 16 

subnigricaule,3,16.23,24 

tortile,16,19,23,24 

undulatum, 20.22 

figs. 23-28 

figs. 27.28 

Distichophyllum 

sect. Mniadelphus, 15 

Epipterygium, 9 

Eriopus, 7 

floccidus, 10, fig. 4 

microblastus, 3.9 

ramosus, 10 

remotifolius, 10 

prviretir, 9, 10 

Hookeria, 1,2,4,11,13, 14,24 

acutifoLa,24 

blumeanum, 1 I 

curpidata, 18 

ppillata, 7 

prabakiana, 7 

purpwea,25,26 

quodrvariwn, 4 

raphidastega, 5 

see&, 25,26 

utacamundiana, 25 

Hookeria sect. Hookeriopsis, 25 

Hookeriopsis, 4,25 

gem’nidens, 25,26 

purpwea, 25 

sumatrana, 25,26 

wichurae, 3,25,26 

utamundiana,25,26 

Hypoperygium, 1 

Lepidophilwn swnatranwn, 3,25 

Leskea cristata, 7 

Leskeodon,2,3,11,15,26, figs. 29-34 

acuminatus, 26.27 29 

auratus, 26 

andicola, 26 

brevicuspidatus,27,29, figs. 31-34,51-54 

cubensis. 26 

longipilus, 26, figs. 55-58,59,60 

mariei, 26 

philippinensis, 3.27.29 

robbinsii, 27 

Mesochaete crendata, 6 

Mnium, 2 

Plagiotheciopsis oblmga, 18 

Pterygophyllum, 4 

dentaturn, 4 

javense, 4 

novae-guineae, 4 

Schizomitriwn, 6 

Thamniopsir, 25 

Tormla mniifolia, 2 

Trachyxiphium, 25 

41