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A taxonomic revision of the moss genera Rhegmatodon and Macrohymenium

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A taxonomic revision of the moss genera Rhegmatodon and Macrohymenium
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Eakin, David Alvin, 1945-
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English
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vii, 122 leaves : ill. ; 28 cm.

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Mosses -- Classification ( lcsh )
Botany thesis Ph. D
Dissertations, Academic -- Botany -- UF

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Thesis:
Thesis--University of Florida.
Bibliography:
Bibliography: leaves 116-121.
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by David Alvin Eakin.

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Full Text
A TAXONOMIC REVISION OF THE MOSS GENERA
RHEGMATODON AND MACROHYMENIUM
By
DAVID ALVIN EAKIN
A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF
THE UNIVERSITY OF FLORIDA
IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE
DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1976




ACKNOWLEDGMENTS
I would like to express my gratitude to Dr. Dana G. Griffin III for his guidance as Chairman of my Supervisory Committee. His example of professionalism in the field of bryological taxonomy, and botany in general, has been an inspiration to me. I would also like to express my love and appreciation to my wife, Darlene, for her patience, understanding, and assistance during the years of my graduate work, and especially during the period of this study.
My thanks also go to Drs. James W. Kimbrough, Indra K. Vasil, Richard C. Smith, and Jonathan Reiskind for their careful reading of the dissertation, and helpful comments.
Specimens in the following herbaria were studied and I express my appreciation to the curators. The abbreviations are those recommended by the International Bureau of Plant Taxonomy and Nomenclature in Index Herbariorum, Part 1, 1974. NICH, FH, MO, MICH, NY, MPU, Z, HBG, B, LD, PE, 0, S, L, GL, MANCH, BM, C, HIRO, H, BR, TENN, PC, FI, FLA, and JE. In several instances H-BROTH was used to indicate that the specimen was from the Brotherus collection housed at Helsinki.
Of the many friends and relatives who have helped and encouraged me, I would especially like to thank my father, Mr. John F. Eakin; my wife's parents, Mr. and Mrs. James E. Walker; and Mike and Diane McCarry. Special thanks go to Ann Ellis and Robert Pentecost for their assistance with my photographic plates, and to Norma Donovan and Phyllis Young for typing the manuscript.
ii




TABLE OF CONTENTS
Page
ACKNOWLEDGMENTS ................................................. ii
TABLE OF CONTENTS ............................................... iii
LIST OF TABLES .................................................. v
LIST OF FIGURES ................................................. vi
ABSTRACT ........................................................ vii
STATEMENT OF PROBLEM ............................................ 1
HISTORY OF RHEGMATODON AND MACROHYMENIUM ........................ 4
FAMILIAL AFFINITIES OF RHEGMATODON AND MACROHYMENIUM............. 14
INFRAGENERIC RELATIONSHIPS IN RHEGMATODON ....................... 16
DESCRIPTION OF THE GENUS RHEGMATODON ............................ 20
KEY TO THE SPECIES OF RHEGMATODON ............................... 23
DISTRIBUTION AND HABITAT OF THE GENUS RHEGMATODON................ 36
DISPOSITION OF NAMES OTHER THAN SYNONYMS IN RHEGMATODON.......... 37 INFRAGENERIC RELATIONSHIPS IN MACROHYMENIUM ..................... 39
DESCRIPTION OF THE GENUS MACROHYMENIUM .......................... 42
KEY TO THE SPECIES OF MACROHYMENIUM ............................. 46
DISTRIBUTION AND HABITAT OF THE GENUS MACROHYMENIUM ............. 57
DISPOSITION OF NAMES OTHER THAN SYNONYMS IN MACROHYMENIUM....... 58 LEGENDS FOR THE FIGURES ......................................... 60
APPENDIX I HISTORICAL PERSPECTIVES IN TAXONOMIC BRYOLOGY...... 93
iii




Page
APPENDIX II PREVIOUS KEYS TO THE SPECIES OF RHEGMATODON
AND MACROHYMENIUM BY BROTHERUS AND KIAER .................. 110
LITBRATURE CITED ................................................ 116
BIOGRAPHICAL SKETCH ............................................. 122
iv




LIST OF TABLES
Page
Table 1. A chronology of important dates and publications
in the history of the genera Rhegmatodon and
Macrohymenium including significant dates in the
history of taxonomic bryology............................. 11
Table 2. Data on three traditional leaf characters formerly
used to differentiate smooth-setaed Rhegmatodons......... .35
v




LIST OF FIGURES
Page
Figure 1. Rhegmatodon spp ....................................... 62
Figure 2. Rhegmatodon spp ....................................... 64
Figure 3. Rhegmatodon spp ....................................... 66
Figure 4. Rhegmatodon declinatus.................................. 68
Figure 5. Rhegmatodon polycarpus.................................. 70
Figure 6. Macrohymenium spp ..................................... 72
Figure 7. Macrohymenium spp ..................................... 74
Figure 8. Macrohymenium rufum..................................... 76
Figure 9. Macrohymenium acidodon.................................. 78
Figure 10. Macrohymenium mdelleri.................................. 80
Figure 11. Macrohymenium strictum.................................. 82
Figure 12. Distribution of Rhegmatodon spp. ....................... 84
Figure 13. Distribution of Macrohymenium spp. ..................... 86
Figure 14. Distribution of Macrohymenium spp ..................... 88
Figure 15. Previous Illustrations of Rhegmatodon.................. 90
Figure 16. Previous Illustrations of Macrohymenium................ 92
vi




Abstract of Dissertation Presented to the Graduate Council
of the University of Florida in Partial Fulfillment of the Requirements
for the Degree of Doctor of Philosophy
A TAXONOMIC REVISION OF THE MOSS GENERA RHEGMATODON AND MACROHYMENIUM
By
David Alvin Eakin
August, 1976
Chairman: Dana G. Griffin, III
Major Department: Botany
The moss genera Rhegmatodon (Leskeaceae) and Macrohymenium
(Sematophyllaceae) are revised on a worldwide basis. Two species,
Rhegmatodon declinatus (Hook.) Bridel emend. Eakin and Rh. polycarpus (Griff.) Mitten emend. Eakin, are recognized for the genus Rhegmatodon.
Four species, Macrohymenium acidodon (Mont.) Doz. and Molk. emend.
Eakin, M. mDlleri Doz. and Molk., M. rufum (Reinw. and Hornsch.)
C. Muell. emend. Eakin, and M. strictum Bosch. and Lac. emend. Eakin, are recognized for the genus Macrohymenium. Keys to the species are included. Presented with each taxon is a complete list of synonomy, a taxonomic description and discussions on morphology and distribution. Standard notation of herbarium disposition is supplied for all
specimens examined, and distributions of the taxa are based only on
these specimens. Where available, information is given on the ecology
of each taxon.
vii




STATEMENT OF PROBLEM
The purpose of this investigation is to present a revisionI of the moss genera Rhegmadodon and Macrohymenium. In the field of bryology today, there is a critical need for taxonomic revisions of many of the currently recognized genera. The relatively short history of modern bryology (less than 200 years) has produced more than a twentyfold multiplication of genera. However, only a very few have been subjected to the scrutiny of a major revision and, of these few, most were revised before the 1930's. In spite of the many fine investigations since the turn of the century, bryological taxonomy remains in the 'alpha' state. Revisional bryology is well characterized by the idea, "the fields are white unto harvest, but the laborers are few." The need for revision appears even more critical when we realize that many of the current genera were established during a period when the philosophical concept of genus in Musci was undergoing tremendous changes in perspective.
Rhegmatodon and Macrohymenium were typical of the many genera needing revision, but at the same time offered features that made them particularly attractive for dissertation research. Both included a "manageable" number of species and there existed a good possibility of procuring nomenclatorial type specimens, so indispensable to revisional taxonomy. The decision to consider two genera, which are
See discussion about the definitions of revision and monograph in Appendix I (page 93).
1




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currently assigned to two different families, was based on the fact that their histories are greatly intertwined. As late as 1898, Paris (Index Bryologicus) included all previously described species of the two genera under the name Rhegmatodon. This was fascinating to me, since even a beginning student of bryology would place these genera in two different groups on the basis of current generic and familial concepts.
The objectives of this study were
1. to collect, translate, and reevaluate all of the previous
taxonomic literature with reference to these genera.
2. to provide a history of these genera with special reference
to the changing concept of the genus within Musci as a whole.
3. to collect and critically reexamine all available type
material on which previous taxonomic concepts were based.
4. to provide a revised taxonomic treatment of these genera
based on type material and the greatest possible number of
other collections.
This treatment includes complete descriptions of the genera and keys to their component species. The types are designated for all species, and their location noted. Collection data and herbaria citations have been given for all specimens examined. Discussions of previous taxonomic treatments, nomenclatorial changes, and synonomy are included. Distributions are presented for each species as recognized by the author. To date, no cytological, genetic, or ecological studies have been completed on these rare genera. None were attempted by this author due to the lack of fresh, living material.




3
There are two areas of background necessary for a proper understanding of the results and conclusions of this dissertation. The first involves an understanding of those basic taxonomic concepts revealing the underlying philosophy and assumptions of modern taxonomic research in bryology. It is important to specifically relate these ideas to the modern generic concept in Musci. Secondly, one must recognize that the thread of history relating Rhegnatodon and Macrohymenium is most meaningfully seen in the context of the overall historical fabric of taxonomic bryology.




HISTORY OF RHEGMATODON AND MACROHYMENIUM
The original specimen of Rhegmatodon was described as part of
a collection of Nepalese mosses sent by Dr. Francis Buchanan Hamilton1 to William J. Hooker. Hooker (1808) based his "Musci Nepalenses" on this collection. In error, he classified his specimen on the basis of a single row of peristome teeth; designating it as Pterogonium declinatum sp. n.,a
Pterogonium caule repente, foliis imbricatus ovatus
integerrimis mediotenus uninervibus, capsule declinata,
operculo incurvato (Hooker, 1808, p. 309).
Had he seen the exostome, and following the generic concepts of his day, he would either have designated it a new species of Hypnum, or perhaps erected a new genus. He was clearly aware of its Hypnaceous appearance.
I have named this plant from its most obvious character,
the drooping capsule, in which it differs from all the other species of this genus described by Hedwig or Bridel, and has,
in consequence of it, so completely the appearance of a Hypnum,
that no one would suspect its real family, without first
examining it (Hooker, 1808, p. 310).
It seems strange that Hooker should have missed the exostome. According to Margadant (1968), Hooker was elected a Fellow of the Linnaean Society of London (1806) at 21 years of age on the basis of his discovery and
IH. C. Gangulee in his Mosses of Eastern India and Adjacent Regions, gives some interesting historical anecdotes about those who have collected in eastern India. The collections of 1802-03, mostly from the Kathmandu valley of Nepal, were the first on the Indian subcontinent and earned for Hamilton the distinction of being the pioneer collector of bryophytes for this area.
4




5
description of Bauxhaumia aphylla in the British Isles. This moss also has a double row of peristome teeth which are not nearly as distinct as those of Rhegmatodon. In addition, W. J. Hooker was an accomplished illustrator and engraver. Nor was it a case of having seen poor material for the original specimen has both endostome and exostome. My only explanation is that he first viewed the specimen dry, in which case, the external teeth would not have been seen at low magnification. In Rhegmatodon the exostome teeth alternate with those of endostome and, when dry, are inflexed between them. This possibility is seen by comparing Hooker's illustration (Fig. 15:7-9) with a low magnification photograph of a dry capsule (cf. Fig. 1:2). We should recall that microscopic characters were not commonly used at this time and microscopes were generally poor. Nevertheless, he did draw the endostome at a higher magnification and it is still puzzling to me how he missed picking off a few exostome teeth.
With reference to the endostome he noted:
There will also be found under the microscope a strong
peculiarity in the circumstance of the teeth of the peristomium being cleft in a very curious manner (Hooker, 1808,
p. 310).
This peculiarity is in fact a central pointin circumscribing the genus Rhegmatodon. His description is as follows:
Peristomium dentibus sedecim, suberectis, e capsulae
membrane interiore reticulate ortis, lineari-subulatis, luteis, transverse striatis, a basi ad medium longitudinaliter fissis (Hooker, 1808, p. 310).
S. E. Bridel, in Bryologia Universa (1827), established the new genus Regmatodon based primarily on this peculiarity of the peristome. The
iNote that the original spelling of the name is Regmatodon and not Rhegmatodon. These two spellings are orthographic variants. However, while some weight should be given to the spelling of the original




6
name itself reflects the character, being formed from the two Greek words pifyp fissure, and oSo~ of the teeth. F. C. Kiaer (1882, p. 41) notes that Bridel "who only knew the plant from the Hooker description, also designated the peristome as simple." Thus, while Bridel did segregate the specimen as a new genus, he perpetuated Hooker's error.
The genus Macrohymenium with one species, M. rufum, was established by Carl Muller in 1847, based on three specimens previously described as Leskea? rufa Reinwardt and Hornschuch (1826), Leskia mitrata Dozy and Molkenboer (1844), and Leskea acidodon Montagne (1845). Muller was apparently strongly influenced in this decision by various characters of the leaves; as indicated by the first of his "essential characters."
The leaves ecostate formed from narrow prosenchymatous
cells (ellipsoidal areolation), strongly concave, subplicate, at the base to both sides provided with several large yellow
lateral cells, densely overlapping, subsecund. With the
capsule mouth constricted. The peristome double; the outer:
the 16 teeth from the oblong base provided with striae in the
middle, traveculate, suddenly long cuspidate, bent inwards; the interior: the teeth just as many, much overlapping the
very longest outer teeth, widest, cuspidate, membrane lacking,
put together from quadrate large yellow cells, placed on the
reddish membrane, scarcely canaliculate, tessellated, not
perforated (Muller, 1847, col. 825).
Just three years later, however, when treating these same specimens in Synopsis Muscorum Frondosorum, he included his new genus under the concept of Rhegmatodon. He dealt with only three species; Rh. declinatus (Hook.) Brid., Rh. orthostegius Mont., and Rh. rufus C. Muller. The former two are Rhegmatodon in the modern sense, but rufus is a Macrohymenium. It may first be puzzling that Muller would dissolve a author, I feel that Bridel's spelling is in error. The Greek word should have a rough breathing symbol over the letter rho indicating the presence of 'h' when the word is transliterated to English. The correct spelling is therefore Rhegmatodon (cf. Stearn, William, 1967, Botanical Latin, p. 263).




7
concept which just three years earlier had been so clear in his mind. Had he overlooked some important characters, or perhaps new and better specimens had come to light? Neither was the case. In fact, in expressing his concept of Rhegmatodon in this latter work, he did not even consider gametophytic characters. Note the following description from Synopsis Muscorum Frondosorum:
The calyptra dimidiate. The peristome double. The
external teeth very short, trabeculate, hypnaceous; the internal upon a short, scarcely keeled, tessellated, as if perforated membrane, turning reddish, very long, for a long time rising above the external, scarcely keeled,
with the longitudinal line gaping to undivided, furrowed,
remotely articulated (Muller, 1850, p. 29).
There is no mention of the once predominant leaf characters in this latter description. The only gametophytic character used in this later classification is the nature of the costa.
This actually represents a shift in Muller's views regarding the relative importance of sporophytic versus gametophytic characters in the delimitation of genera. This example well illustrates the problem facing bryologists as a result of the dual nature of mosses. The difficulty of systematizing a double organism, i.e., one representing two distinct generations of the life cycle is discussed in Appendix I. This problem is in fact central in the history of the changes which have taken place in the concept of genus in mosses. This disciplinewide vacillation with regard to the selection of essential characters is reflected throughout the history of Rhegmatodon and Macrohymenium. Muller was obviously struggling with the problem in his treatment of the two genera. Hedwigian tradition emphasized the importance of the sporophyte. Muller at first breaks with this tradition by emphasizing various gametophytic characters and establishing the new genus




8
Macrohymenium. He then conservatively shifts his emphasis back to sporophytic characters which, in the case of these taxa, will unite them under the same generic concept. This is a characteristic trend in the classification of the two genera. Whenever the gametophytic leaf characters were emphasized, Macrohymenium and Rhegmatodon were separated. Emphasis on the sporophyte, especially the peristome characters, united the specimens. Dozy and Molkenboer (1848) in Musci frondosi inediti archepelagi indici . ., attempted to refine the genus concept for Macrohymenium on the basis of two new species, M. mulleri and M. serrulatus. Again we find the major emphasis given to the shape and structure of the peristome; especially to the ratio of length between the endostome and exostome teeth. Some insight into their view of the limited importance of gametophytic characters is shown by the following:
Quadrate areoles to both sides at the wing of the
leaves are not uniformly evident in all species, and
therefore, they are seen to us not to be of great value,
at least in explanation of the essential characters (Dozy
and Molkenboer, 1848, p. 166).
Emphasis on the similarities of the peristome caused them to ignore the obvious lack of inflated alars in serrulatum. This further illustrates the principle stated above. However, in this case, Dozy and Molkenboer were dealing with two species of Macrohymenium and only one of Rhegmatodon; thus including both genera under the name Macrohymenium. In 1864, van der Bosch and van der Sande Lacoste continued Dozy and Molkenboer's work in Bryologia Javanica. They added one new species, M. strictum, and reclassified M. serrulatum as Rhegmatodon serrulatus. While they did not specify the reason for this move, a look at their




9
description of Rh. serrulatum implies greater stress on gametophytic differences.
A major revision of the two genera was undertaken by F. Kiaer
in 1882, which essentially established the modern distinctions between Rhegmatodon and Macrohymenium. There has been very little change in the circumscription of these taxa since Kiaer. Later taxonomic changes revolved around the question of closeness of relationship between the two genera. Kiaer used mostly gametophytic characters to distinguish the genera but placed them in the same subfamily, Macrohymenieae, on the basis of their proportionately long endostome. He lists: Rh. schotheimioides, Rh. filiformis, Rh. crizabanus, Rh. orthostegius, Rh. polycarpus, Rh. brasiliensis, Rh. secundus, Rh. declinatus, and Rh. serrulatus. Under Macrohymenium he includes: M. rufum, M. strictum, M. acidodon, M. nietneri, M. laeve, M. mulleri, and M. gracillimum.
All of these species distinguish themselves (as a
group-DAE) by means of the great inner peristome with
the processes which are 2-4 times longer than the teeth
(Kiaer, 1882, p. 10).
With the publication of Philibert's Ptudes sur le piristome (1884-90) the peristome again was emphasized as the essential character in taxonomic bryology. The impact of Philibert on the classification of these genera is most clearly reflected by the treatment of these species by Paris (1898) in Index Bryologicus. All species of Macrohymenium were synonomized under the name Rhegmatodon. Twenty-one species are listed, apparently on the basis of this similarity in peristome structure. Gametophytic distinctions must necessarily have been entirely abandoned as generic criteria.
When Brotherus (1909), in Engler and Prantl's Die Naturlichen
Pflanzenfamilien (1st edition), established the family Sematophyllaceae




10
on the basis of Mitten's (1869) Tribus 16, Sematophylleae, the stage was set for separating Rhegmatodon and Macrohymenium. Brotherus' family concept was later emended by Fleischer (1923), Die Musci der Flora von Buitenzorg, into four subfamilies: Clastobryoideae, Heterophylloideae, Sematophylloideae, and Macrohymenioideae. Brotherus (1925) followed this treatment in the second edition of Engler and Prantl.
Moving away from the overwhelming emphasis on the sporophyte, the essential characteristics of the family Sematophyllaceae (sensu Brotherus and Fleischer) are
1. costa absent, or short and bifurcated
2. alar cells enlarged
3. operculum long-rostrate.
Both of these authors gave little regard to the peristome and thus the sporophytic characteristics within the Sematophyllaceae are widely variable. This was the last major treatment of these taxa. Currently Macrohymenium is found as a subfamily of the Sematophyllaceae, and Rhegmatodon holds a similar position in the Leskeaceae. Subsequently, five new species have been described and I have found a great number of names ex herbario, which apparently have not been validly published.




Table 1. A Chronology of important dates and
publications in the history of the genera Rhegmatodon and Macrohymenium including significant dates in the
history of taxonomic bryology.
1741 Dillenius Historia Muscorum
1753 Linnaeus Species Plantarum
1781 Hedwig Fundamentum Hist. natural
1801 Hedwig Species Muscorum
1808 Hooker, W. J. First specimen of Rhegmatodon;
described as Pterogonium declinatum 1819 Bridel, Mantissa Muscorum
1826 Bridel, Bryologia Universa the establishment of
the new genus Rhegmatodon based on Hooker's Pterogonium
declinatum
1826-27 Leskea? rufa Reinwardt & Hornschuch
1836 Bryologia Europaea begun
1838 Anhymenium polycarpon Griff.
1839 Leskea parvula Hampe
1842 Rh. orthostegius Mont.
1844 Rh. parvulus Hamp.
1844 Leskia mitrata Dz. & Molk.; in Muscorum FrondosorumNovae Species ex Archipelago Indico et Japanio 1845 Leskea acidodon Mont.
1847 Macrohymenium rufum MiUller; the establishment of the
new genus Macrohymenium
1848 M. mUlleri Dz. & Molk; M. serrulatum Dz. & Molk; in
Musci frondosi inediti Archipelagi indici sive descriptio
et adumbratio Muscorum Frondosorum in insulis Java,
Borneo, Sumatra, Celebes, Amboina.




12
1849 Anhymenium polysetum Griff.
1848-51 (50) MUller, Synopsis Muscorum Frondosorum M. rufum changed
to Rh. rufum
1854 Bryologia Europaea, Bruch, Schimper, & Guembel; completed
1856 Clasmatodon parvulus Hook. & Wils.
1859 Mitten's Musci Indiae Orient
1862 Rh. brasiliensis MUller
1855-70 (64) Bryologia Javanica, M. strictum Dz. & Molk. 1868 Rh. kunzii M ller in schedule
1869 Mitten's Musci Austro-Americana; Rh. schlotheirnioides
Spruce; Tribus Sematophylleae established
Hypnum minutum Mitten
Rh. nietneri MUller
1871 Bescherelle, Prodromus Bryclogiae Mexicanae
Rh. filiformis Besch.
Rh- fusco-luteus Schimp
Rh. hypnoides Besch. 1872 Rh. chryseus Schimp.
1872 (73?) M. laeve Thw. & Mitten 1876-77 Adumbratio florae muscorum, Jaeger & Sauerbeck
First reference to Rh. orizabanus
1880 M. acidodon (part of rufum moved by Bescherelle)
Florule Bryologique de la Reunion M. acidodon var. acutissima Besch.
1881 Rh. filiformis forma major (collection date of in schedule
specimen of Bruch)
1882 Rh. madagassus Geheeb
Kiaer, revision of Rhegmatodon and Macrohymenium
Rh. secundus Kiaer
Rh. census Schimp. in schedule
M. gracillimum M 11. in litt.
Rh. orizabensis Besch. in schedule M. cuspidatum Mitt. sedis incertis
1884-90 Philibert, H. 9tudes sur le pe'ristome
1891 Rh. feanus MUller




13
1895 Rh. palustris Broth. in schedula
Rh. palustris var. minor in schedula
Rh. schlotheimioides var. minor 1897 Rh. newtonii Broth.
1898 Paris, Index Bryoloicus (all species of Rhegmatodon and
Macrohymenium under Rhegmatodon)
1904-05 Rh. bornmulleri Broth. in schedula
1905 Rh. fissidens Theriot in schedula
1907 Rh. filirameus Broth. in schedula
1909 Rh. sinense Theriot
1909 Engler & Prantl 1st edition made the family
Sematophyllaceae from bitten's Tribe Sematophylleae 1910 Rh. pringlei Cardot
1910-11? Rh. mulleri forma pubea Dixon in schedula
1910 Rh. crassirameus Cardot
1910 Rh. cameruniae Broth. (collection date)
1923 Fleisher, Die Musci der Flora von Buitenzorg (4 vol.)
Macrohymenioideae one of four subfamilies of Sematophyllaceae
M. mitratum
M. wichurae Broth. in schedula
M. densirete Broth.
1924-25 Musci, Brotherus, in Engler and Prantl, Die naturlichen
Pflanzenfamilien 2nd edition 1924 Rh. handelii Broth.
1929 Rh. declinatus var. minor Broth.
Rh. novo-guinense Reimers
1932 Manual of Bryology, Verdoorn. Contains Dixon's
classification
1947 Rh. brevicuspes P. de la Varde and Leroy (collected 1936)
1955 Rh. schwabei Herzog




FAMILIAL AFFINITIES OF RHEGMATODON AND MACROHYMENIUM
There have been some comments in the literature suggesting that the genus Rhegmatodon be transferred from the Leskeaceae to its own monotypic family, the Rhegmatodontaceae (Crum, 1951, 1973). The apparent motivation for the suggestion is the unique peristome of Rhegmatodon. Brotherus, in the second edition of Engler and Prantl (1925), segregates Rhegmatodon in its own subfamily, the Rhegmatodontoideae, on the basis of the
Peristom weit unter der Mundung inseriert, das
aussere veil kurzer.
Macrohymenium holds a parallel position with respect to the Sematophyllaceae. Brotherus separates this genus into its own subfamily, the Macrohymenioideae, on the basis of
Fortsatze 2-3 mal so lang als die Peristomzahne.
In both cases, the remaining genera of these families have peristomes in which the endostome is either the same length as the exostome or perhaps a little shorter or absent. These two genera stand alone in their respective families by virtue of an endostome 2-3 times longer than the exostome. However, this peculiarity of the peristome is not a sufficiently strong character to warrant such a transfer. With reference to some of the more recent discussions of the Leskeaceae and related families (Watanabe, 1972; Noguchi, 1972; Crum, 1973), all of the Rhegmatodon generic characters fit well into the family concept, with the exception of the ratio of endostome to exostome length. A
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similar situation exists with respect to the relationship of Macrohymenium to the Sematophyllaceae. Although Seki (1968) did not specifically deal with Macrohymenium in his revision of the Sematophyllaceae of Japan, this genus fits very well into his revised concept of the family; again with the exception of the endostome to exostome ratio. Both in the case of Rhegmatodon and Macrohymenium, transfer to a monotypic family would be unnecessary and place too great an emphasis on this one peculiarity of the peristome. It is recommended that Rhegmatodon remain in the subfamily Rhegmatodontoideae of the Leskeaceae, and that Macrohymenium retain a similar position in the subfamily Macrohymenioideae of the Sematophyllaceae. Much more revisional work should be done with reference to the genera in these families before new families are proposed.




INFRAGENERIC RELATIONSHIPS IN RHEGMATODON
As previously noted (cf. p. 9), Kiaer accurately circumscribed the genus Rhegmatodon with reference to both gametophytic and sporophytic characters. Within Rhegmatodon he erected two sections, Laeviseta and Scabroseta, which correspond to the two species of the present treatment. In the wake of Kiaer's revision, however, a proliferation of new species occurred on the basis of certain gametophytic characters, i.e., leaf shapo, average leaf ratio and perichaetial leaves. In the present work each of these characters was tested, both in the type specimens and later collections. I attempted to use both gametophytic and sporophytic characters to delimit species and gave careful consideration to those characters used by previous workers. However, the traditional gametophytic characters were found either to be continuously variable or to represent ecotypic variants, and were thus set aside.
The morphological structures of taxonomic importance in Rhegmatodon are completely sporophytic. The only sure way to distinguish the two species is by examination of the endostome teeth, the seta and the walls of the exothecial cells. These three characters will be found in one of two discrete combinations, each characterizing one of the two species of Rhegmatodon. Rh. declinatus can be recognized by the combination 1) endostome teeth lacking dotting (Figs. 2:5, 4:12), 2) seta clearly rough from top to bottom (Fig. 4:3), and 3) cell walls
16




17
of the exothecium irregularly thickened, presenting a collenchymatous appearance (Fig. 4:13). Rh. polycarpus is characterized by 1) dense dotting of the endostome teeth (Fig. 5:1, 2), 2) a smooth seta (Fig. 5:3), and 3) the walls of the exothecial cells regularly thickened parallel to the central axis of the capsule, with the transverse walls appearing thinner (Fig. 5:23).
The taxonomic treatment of Rhegmatodon by Brotherus (in Engler and Prantl, 1925) is based on F. C. Kiaer's revision of 1832, Genera muscorum Macrohymenium et Rhegmatodon revisa specieque nova aucta exposuit.1
Within Kiaer's and Brotherus' Scabroseta, species were differentiated by serrations on the leaf tips. This character proved to be a continuum, being highly variable, even within a single specimen. From specimen to specimen an examination of leaves from the branch tips revealed a continuum from those completely entire, to a few somewhat serrated, to those distinctly serrate (Fig. 4:5-1). In all cases, the leaves at the base of the branch were entire. To date no specimens of Rh. polycarpus with serrate leaves have been seen, though a slight tendency toward serration was noted (Fig. 5:9). Caution should be exercised in using this difference alone to distinguish species, as leaf serrations have been shown to be ecotypically variable (Briggs, 1965).
In section Laeviseta, Kiaer and Brotherus used leaf ratios (2-2.5:1 vs. 1.8:1) and leaf shapes (oval vs. oblong-oval) to distinguish species. Table 2 presents data taken primarily from type specimens for all the
For the purpose of comparison, translations of both these keys are presented in Appendix I.




18
species with smooth setae. This clearly shows a continuum of leaf ratios. The averages in Table 2 are based on an examination of a minimum of 20 individual leaves per specimen. No correlation exists between leaf length and ratio or ratio and shape. Figure 5:11-22 well illustrates the range of the continuous variables of size, shape, ratios, and secundity. The extent and degree of secundity varied even within the same specimen. For example, the three leaf tips shown in Fig. 5:7-9 are from the same specimen. Secundity cannot be used in Rhegmatodon as a discrete species marker. Developmental studies will probably show this character to be an ecotypic variable as has been shown for other mosses (Streeter, 1970). Similarly, the degree of filiformity in branches was found to be highly variable within single specimens, and was excluded as a marked species character. This character has also been shown to be an ecotypic variant of mosses in very moist microhabitats (Richards, 1967).
Several pictures of the exostome teeth have been included which clearly show two external plates backed by a single set of highly trabeculate plates (Fig. 2:1-3, 6-12). This was considered necessary since the well-known illustration by Dozy and Molkenboer (Fig. 15:4) is incorrect on this point. Their illustration represents only the interior set of trabeculate plates. The error has been widely circulated in that Brotherus chose this plate as his illustration of Rhegmatodon in Die natUrlichen Pflanzenfamilien (1925). In this same drawing, the relative height of the endostomal and exostomal membranes is misrepresented. In the genus Rhegmatodon, the point of attachment of the endostome and exostome to their respective membranes is the same, i.e., the point of attachment for both the endostome and exostome




19
is an equal distance below the rim of the capsule. Dozy and Molkenboer drew this as if Rhegmatodon serrulatus was in fact a Macrohymenium, as they supposed. Both parts of the peristome are inserted below the rim of the capsule in Rhegmatodon.
The presence of paraphyllia in Rhegmatodon was first noted by Fleischer (1923) and subsequently by Brotherus (1925). However, no description accompanied these notations. These structures actually fit the concept of pseudoparaphyllia as presented by Ireland (1971). A comparison of Ireland's illustrations of pseudoparaphyllia with the pictures in Fig. 2:13-22 will document the similarity of structure. Furthermore, they are relatively sparse in Rhegmatodon by virtue of being restricted to branch and reproductive primordia on the branches.
One other character used by Kiaer and Brotherus was the presence
of teeth on the margins of the inner perichaetial leaves (cf. Fig. 1:17). The consistency of this character was tested in several specimens by the examination of many perichaetia. The presence and number of teeth was not at all consistent, whether in the same specimen or in different specimens, supposedly of the same species. This character was also excluded.




DESCRIPTION OF THE GENUS RHEGMATODON
Regmatodon (Rhegmatodon) Bridel, Bryol. Univ. 2:294. 1827.
Pterogonium declinatum Hook., Trans. Linn. Soc. IX, t. 26, f. 3.
1808. TYPE: Dr. Francis Buchanan Hamilton, Nepal, 1802-03
(holotype, NY; isotypes, S, H, BM).
Plants slender to fairly robust, forming loose to dense mats,
mostly dull or somewhat shiny, rigid, dark green or brownish. Primary stems prostrate, creeping, yellow to brownish, irregularly subpinnately and densely branched, rooting by means of radicles on the underside. Stem leaves small, usually absent, short, ovate-acute, entire, alars inconspicuous, marginal cells quadrate, the rest elliptical. Branches ascending to upright, terete, straight to curved, rigid, varying from short, thick and clavate to long, slender and of uniform diameter. Pseudoparaphyllia present but sparce. Branches densely foliose, leaves appressed and densely imbricate when dry, erect-spreading when wet, 0.5-1.7 mm long. Leaves broadly ovate, oblong-ovate to ovatelanceolate, short acuminate, concave, completely entire to strongly serrate at the apex, margins weakly to strongly reflexed to base of short acumen. Costa single from a wide base, narrowing rapidly, disappearing at or somewhat above midleaf. Laminal cells oblong, arranged in oblique and longitudinal series, crowded, becoming subquadrate toward the margins and base of the leaf. Indistinctly bordered by a single
20




21
row of quadrate marginal cells with transverse walls at right angles to the margin. Alars undifferentiated. Goniautoicous. Inner perchaetial leaves erect, lanceolate, gradually acuminate, with either completely entire or crenately denticulate margins (Fig. 1:12-17). Perichaetium inserted near the base of the branch, budlike, somewhat open at the apex, archegonia shortly stipitate, cylindric from an oval base, with numerous filiform paraphyses, a little longer than the archegonia (Fig. 1:8). Perigonium axillary, budlike, ovate, leaves somewhat round ovate, suddenly constricted into a short, wide, moderately obtuse acumen, concave, completely entire, ecostate, the cells loose, hexagonal-rhomboidal (Fig. 1:6, 9). Paraphyses filiform and numerous (Fig. 1:9). Seta either smooth or rough, 4.5-13.5 mm in length. Capsule erect to inclined, unsymmetric, straight to somewhat curved, slightly constricted below the mouth, especially when dry, oblong cylindric to long cylindric, 1.5-3.1 mm long, smooth, brown to reddish brown, sometimes substrumose at base, exannulate. Exothecial cells either collenchymatous or with regularly thickened longitudinal walls and thinner transverse walls. Operculum short, conic, acumen short, obtuse, 1/4-1/2 the length of the deoperculate capsule (cells regularly hexagonal). Peristome double, the sixteen teeth of the exostome lanceolate, obtuse, inflexed in the dry state, composed of a double series of cross-striolate external plates, and a single series of inner plates, trabeculate on the inner face (Fig. 2:1-3, 6-12). The sixteen processes of the endostome, well-developed, 2X longer than the exostome teeth, translucent, with or without dotting, split on midline from base to middle with one to three perforations in segments immediately above, appearing faintly to strongly bordered (Fig. 2:5). No cilia.




22
Spores yellow-green to olive, globose, papillose, 21-30 pm. Calyptra cucullate, smooth, somewhat longer than the operculate capsule.




KEY TO THE SPECIES OF RHEGMATODON
. Seta rough from top to bottom; endostome teeth translucent and not dotted; the walls of the exothecial cells irregularly thickened, appearing collenchymatous. .... Rh. declinatus Seta smooth from top to bottom; endostome teeth translucent and densely dotted; the walls of the exothecial cells regularly thickened longitudinally, the transverse walls appearing thinner Rh. polycarpus
23




24
1. RHEGMATODON DECLINATUS (Hook.) Brid., Bryol. Univ. 2:204-205. 1827.
(Figure 4)
Pterogonium declinatum Hooker, Trans. Linn. Soc. Lond. 9:309-310,
t. 26, f. 3. 1808. TYPE: Nepal, without precise locality,
Francis Buchanan Hamilton, s.n. (holotype, NY; isotypes,
S, H, BM).
Macrohymenium serrulatum Doz. & Molk., Musci Frond. Ined. Archip.
Indici 6:170, t. 56. 1848. TYPE: Java, Mt. Patoeha,
Korthals, s.n. (holotype, L; isotypes, S, H, BM, O, L, PC).
New synonomy.
Rhegmatodon serrulatus (Doz. & Molk.) Bosch. & Lac., Bryol. Jav.
2:111-112. 1864. New synonomy.
Rhegmatodon feanus C. Miell., Nuov. Giorn. Bot. Ital. 23:601. ]891.
nom. nud. TYPE: Burma, Bhamo, Feanus 25 (holotype, FI).
New synonomy.
Rhegmatodon declinatus var. minor Broth., Symb. Sin. 4:94. 1929.
TYPE: China, in Kwangtsaoba, S. W. Kweitschou, HandelMazzetti 10395 (holotype, H-BROTH; isotype, S). New synonomy.
Rhegmatodon schwabei Herz., J. Hatt. Bot. Lab. 14:66, f. 22. 1955.
TYPE: Formosa, western Middletaiwan, "Tigerkopf," Schwabe,
s.n. (holotype, JE). New synonomy.
Branch leaves 0.6-1.7 mm long, 0.3-0.7 mm wide, the average length to width ratios ranging from 2.2-3.0. Leaf margins range from completely entire to coarsely serrate, from midleaf to tip. Seta from 4.5 to 10.5 mm in length, 187-260 pm wide, conspicuously rough from top to bottom. Vaginule 1.0-1.9 mm long. Capsule 1.5-2.4 mm long. Exothecial cell




25
walls with irregular thickenings giving a collenchymatous appearance. Endostome teeth translucent and lacking dotting. Endostome teeth are twice the length of the exostome teeth, ranging from 450-600 pm, while the exostome range from 200-300 pm.
Habitat: Usually on bark of fallen logs or trees, also on moist
shady rocks. Mostly 500-1700 meters in altitude (rarely to 3000 meters).
Distribution: Formosa, China, Burma, India, Nepal, Borneo, Sri Lanka, Java, Thailand, and Malakka (Malay Peninsula), Sikkim, Assam (Fig. 6:2).
FORMOSA. Mt. Lu-Chang-Ta, Nao-liao to Kuei-shan, Wu-feng Hsiang, Hsinchu, on bark of log on roadside in hardwood forest, 1800 m, Wang 1562; upstream area of Luming Chi, Hung-yey Tsun, Yen-ping Hsiang, Taitung, on hardwood log lying on roadside in hardwood forest, 100 m, Wang 0903; upstream area of A-li-pu-tang Chi, Wang-mei Tsun Sin-yi Hsiang, Nantow, on large branch of broad-leaved tree in hardwood forest, 1200 m, Wang 1449 (NICH); Hsueh Shan Shan Mo, on fallen tree in hardwood forests just below Anma-shan, c. 2100 m, Iwatsuki & Sharp 3140; Hsueh Shan Shan Mo, 17 km, above Anma-shan at edge of logging area near Chunghsueh-shan in a wet, n-facing ravine on a decaying log, c. 2500 m, Iwatsuki & Sharp 732a (NICH, TENN). Western Middletaiwan, Tigerkopf, Schwabe, s.n.. (JE).
CHINA. Fukien Province, Buong Kang, Yenping, on mossy bark covering the whole base of a tree, 700 m, Chung B34 (NICH, FH, MICH, O, PE, S); on a mushroom log, 1000 m, Chung B117 (NICH, FH, MICH, O, BM); on a mushroom log in a bamboo forest, 3000 m, Chung B112 (FH, MICH, PE); on mossy rock, 700 m, Chung B48b, B52 (FH); on branch of madiellus, Chung B90 (FH, MICH); on rock, Chung B65, B68, B67a (FH, MICH); Chung B74a




26
(FH, MICH, O); B90a (MICH); Fukien Province, Kushan near Foochow, on a shady rock, 500 m, Chung B173 (FH); on a moist shady rock, Chung B245a (FH, HBG, MICH). Province Kweitschou, austro-occid, ad viam TschenningKuang-tsaoba, Yunnan in silva ad vic. Djitschangring pr. opp. Muyusse, substr. truncis viv. arb. frondos, c. 1050 m, Handel-Mazzetti 10395 (S, H-BROTH). Province Kweitshou orient, prope oppidum Liping in silva mixta Nandjing-schan, ad arbores, c. 750 m, Handel-Mazzetti 10981 (H). Province Kwangsi, Laoshan, Ling Yuin Hsine, Cheo 1963 CFH). Yunnan, frontier of Burma, no collector cited, 1898 (H).
BURMA. Yunnan frontier, Niebolitz, 1911 (L, JE, PC). Bhamo, 3000', Feanus 25 (FI).
SIKKIM. Yoksam, 1700 m, U. Tokyo Bot. Exp. to E. Ind. 200244
(NICH, NY, L); 20046 (H); Himalaya, prope Kurseong, 1372 m, P. Decoly & Schaul 2356 (BM, H).
ASSAM. Khasia, Myrung, Griffith 212, Herb. E. India Co. 515 (BM).
INDIA. Dehra Dhoon, Simla 717 (Griffith) (NY, BM).
NEPAL. Without precise locality, Buchanan, s.n. (NY, S, H, BM). Without precise locality, Wallich, s.n. (BM).
THAILAND. Payap, limestone massive Doi (Mt.) Chiegdao, hill evergreen forest, on fallen tree trunks, 980 55' E 190 25' N, Touw 9063 (L).
EAST INDIES. Without precise locality, Buchanan 1316 (S, BM).
BORNEO. Between Sosopodon and S. Kelinggen, foot of Mt. Kinabalu, on fallen trunk, 1350-1400 m, Iwatsuki 1355 (NICH).
SRI LANKA. Central Province, Muller 248 (H, PC).
JAVA. Mt. Patoeha, Korthals, s.n. (S, H, L). Mt. Gede, Gerker, s.n.; Mt. Patoeha, Korthals, s.n. (L). Lignes i Habitus mest den fra Java in ag pedicelli Lenghe, Monte Patoeha, Korthals (0). Buitenzorg




27
an Baum, Schiffner 11016 (S). Without precise locality, Korthals, s.n. L, BM, S, PC). Without precise locality, no collector cited (II, S, BM, C). Natural Reserve, Tjibogo, rainforest, tree trunk in shade, 1480 m, Soekar, 1949 (L). Tjibodas, Pentjuran mas on bark, van der Wijk 1001
(L). Without precise locality, Junghuhn, s.n. (L).
MALAKKA. Bergland der Sakai, Werner, 1913 (S, JE).
Rhegmatodon declinatus is clearly distinguished from Rh. polycarpus by its rough seta, the absence of dotting on the endostome teeth, and the collenchymatous appearance of the exothecial cell wall thickenings. It is unfortunate that no morphological characters of the gametophyte can be used to separate sterile specimens of the genus.. Although no specimens of Rh. polycarpus having serrated leaf tips have been collected, I would caution against using this character alone to identify a specimen as Rh. declinatus. Although branch leaf characters fall within a continuum, the branch leaves of Rh. declinatus are, on the average, longer and more narrow than those of Rh. polycarpus. Similarly, the range of length for both the capsule and the seta is somewhat shorter in Rh. declinatus.
The holotype for Rh. schwabei Herz. was sterile. As only Rh.
declinatus has been collected in Formosa, Rh. schwabei was placed under it.
While Rh. polycarpus is widely distributed in the New World,
no collections of Rh. declinatus have yet been made from this part of the world. Rh. declinatus seems then to be restricted to the Old World tropics and subtropics.




28
2. RHEGMATODON POLYCARPUS (Griff.) Mitt., J. Linn. Soc. Bot. Suppl.
1:127. 1859. TYPE: Khasia, Hooker and Thomson 775 (lectotype,
BM).
(Figure 5)
Anhymenium polycarpon Griff., Griff. muscolog. itin. Assamici,
1838 (Calcutta J. of Nat. Hist. 3:275, t. 16. 1843), Not.
p. 471, Griff Icones plant. Asiat. 2: t. 97, f. 1. 1849.
TYPE: Assam, Mumbree, Griffith, 1835 (not seen).
Rhegmatodon orthostegius Mont., Ann. Sc. Nat. Bot. ser. 2, 17:248.
1842. TYPE: India, Nilghiris, Perrottet, s.n. (lectotype,
O; syntypes, BM, PC, NY, O). New synonomy.
Anhymenium polysetum Griff., Not. p. 472, 1849, Griff. Icones
plant. Asiat. 2: t. 97, f. 2., 1849. nom. inval. TYPE:
Assam, without precise locality, Griffith, s.n. (lectotype,
NY).
Rhegmatodon brasiliensis Lindb. ex C. MUller, Bot. Zeit. 20:374.
1862. TYPE: Brazil, prov. Sao Paulo, Santos, Lindberg,
1854 (holotype, S; syntypes, O, BM, H). New synonomy.
Rhegmatodon schlotheimioides Spruce ex Mitt., J. Linn. Soc. Bot.
12:566. 1869. TYPE: Ecuador, Andes Quitenses, near
Antombos of the river Pastasa, Spruce 1437 (holotype, NYMITT; isotypes, MPU, O, S, BM, H, BR, PC). New synonomy.
Rhegmatodon filiformis Schimp. ex Besch., Mem. Soc. Sc. Nat.
Cherbourg 16:231. 1872, et Besch. in Fourn., Mex. Pl. 1
(Crypt.): 43. 1872. TYPE: Mexico, Chinantla, Liebmann.
1841 (lectotype, C; syntypes, 0, BM, H). New synonomy.




29
Rhegmatodon orizabanus Hamp. in Jaeg., Ber. S. Gall. Naturw. Ges.
1877-78:477. 1880 (Ad. 2:741) nom. nud. TYPE: Mexico,
Mt. Orizaba, no collector cited, s.n. (lectotype, NY; syntype, BM).
Rhegmatodon filiformis f. major Bruch, in schedula, (holotype,
BM). New synonomy.
Rhegmatodon secundus Kiaer, Forh. Vid. Selsk. Christiania 1882
(24): 38, 2 f. 5-7, 3 f. 1-5. 1883. TYPE: Malagasy Rep.,
Mt. Ankaratra, Borgen, 1887 (holotype, 0; isotypes, S, NY,
L, BM, C, H, BR, PC). New synonomy.
Rhegmatodon orizabensis Besch. in Kiaer, Forh, Vid. Selsk.
Christiania 1882 (24): 37. 1883. nom. nud. in synon.
Type not seen.
Rhegmatodon densus Schimp ex Kiaer, Forh. Vid. Selsk. Christiania
1882 (24): 37. 1883. TYPE: Mexico, Orizaba, Mohr, 1857
(lectotype, BM). New synonomy.
Rhegmatodon palustris Broth., Bih, K. Svensk. Vet. Ak. Handl. 21
Afd. 3(3): 64. 1895 nom. nud. in synon. TYPE: Brazil,
prov. Minas Geraes, Caldas, Mosen 365 (holotype, H; isotypes,
Z, FH, S, BM). New synonomy.
Rhegmatodon schlotheimioides var. minor Broth., Bih. K. Svensk.
Vet. Ak. Handl. 21 afd. 3(3): 64. 1895. TYPE: Brazil,
without precise locality, Cardot 38 (holotype, H). New
synonomy.
Rhegmatodon palustris var. minor Broth., in schedula. TYPE:
Brazil, without precise locality, Binot, s.n. (holotype, BR). Rhegmatodon newtonii Broth., Bot. Jahrb. 24:281. 1897. TYPE:




30
Fernando Po, Pic Clarence, Newton, 1894 (holotype,.H-BROTH;
isotype, S). New synonomy.
Rhegmatodon filirameus Broth., in schedula, 1907. TYPE: Guatemala,
Alta Verapaz, Coban, TUrckheim 6744 (holotype, H-BROTH;
isotypes, FH, NY, LD, S, BM, PC, FI). New synonomy.
Rhegmatodon pringlei Card., Rev. Bryol. 37:58. 1910. TYPE:
Mexico, Puebla, Tezuitlan, Pringle 15285 (holotype, H; isotypes, FH, NY, LD, O, S, L, MANCH, BM, C, PC). New
synonomy.
Rhegmatodon crassirameus Card., Rev. Bryol. 37:58. 1910. TYPE:
Mexico, Morelos, Cuernavaca, Pringle 15283 (lectotype, H;
cotype 15310, NY). New synonomy.
Rhegmatodon cameruniae Broth., in schedula, 1910 (TYPE: Cameroon,
Kamerunberg, Musalse, Bushwald, Hintz, 1910 (holotype,
H-BROTH). New synonomy.
Rhegmatodon handelii Broth., Sitzungsber. Ak. Wiss. Wien Math.
Nat. Kl. Abt. 1, 133:578. 1924. TYPE: China, Yunnan,
Handel-Mazzetti 260 (holotype, H-BROTH; isotype, S). New
synonomy.
Rhegmatodon brevicuspis P. Vard et Leroy, Bull. Jard. Bot. Bruxelles
18:182. 16. 1947. hom. illegit. TYPE: Africa, Tschibinda,
Leroy 227 (holotype, BR). New synonomy.
Branch leaves 0.5-1.3 mm long, 0.2-0.6 mm wide, the average length to width ratios range from 1.85-2.83. Leaf margins are completely entire with no serrations, weakly to strongly reflexed to base of the acumen. Seta from 5.5-13.5 mm in length, smooth from top to bottom. Vaginule 1.0-1.9 mm long. Capsule 1.9-3.1 mm long. Exothecial cells




31
with regularly thickened longitudinal walls; transverse walls appearing thinner. Endostome teeth translucent, densely punctate. Endostome teeth ranged from 470-620 Pm and exostome teeth ranged from 200-350 Jim.
Habitat: Usually on the bark of trees or fallen logs, occasionally on wet ground and rocks. Ranging from 1000-3048 m in altitude.
Distribution: Sikkim, Assam, India, China, Sri Lanka, Thailand, Malagasy Rep., Malawi, Guinea, Fernando Po, Cameroon, Mexico, Panama, Guatemala, Brazil, Bolivia, Burma (Fig. 6:1).
SIKKIM. Without precise locality, 2-4000', no collector cited, s.n. (NY). Punkabari, 2-4000' Kurz 2453 (H, BM). Himalaya, 2-4000', Kurz 276/5 (0). Without precise locality, 5000', King 114b (BM). Without precise locality 2-8000', King, 1894 (BR).
ASSAM. Moosai, Griffith, s.n. (O, H). Khasia, regio sub trop., 4000', JD Hooker and Thomson 775 (BM).
INDIA. Kumaon, NW Himalaya, Malkarzun, Askote district, 1388 m, Khan, Broth 1940 (NY, S, H, BM); Rolam River, 3048 m, Khan, Broth 1935 (O, H). Nilghiris, Perrottet 1644 (BM, NY); Perrottet, s.n. (NY, BM, O, PC); Montagne, s.n. (NY, BM, BR, L, O); prope Ootacamund, Weir 276/21 (NY, BM); Ootacamund, 7000', Gambleu 17260 (H); Coonor, trees, 5000', Gambleu 13013 (H). Nilghirebirge bei Coonor in Simsparkjungle an Baumen, 1950 m, Fleischer 511 (B, H, L). Eastern Nilghiris, Kotagiri, tree, 6500', Sedwick 757 (BM). Nilghiri Hills, Srinivasan 277 (HIRO). Palni (Pulney or Palm) Hills, Kodaikanal, Foreau, 1927 (FH); in groves along the Kodai road about 29th mile, c. 5900', Foreau 477 (FH); Kodai road, Foreau 199 (BM); Togaivarai Shola, 4500', Foreau 262/26 (F, BM); Perumalmalai Shola, along torrent, 4500', Foreau 141/26 (BM); near Shembaganur, Pragasafuram, 6400', Foreau, 1957 (BR); 6700', Foreau, 1958 (TENN). Kodaikanal, on trees, 2400 m, Foreau and Roine 71 (MANCH,




32
PC, FH, O, H); Madura, Foreau, 1911 (BR); en Madure dans les Gathes, 7000'+, Andre 35 (PC); Andre, 1909 (BM).
BURMA. Yunnan frontier, Niebolitz, 1911 (JE).
CHINA. Yunnan, Pe yen tsin, auf Baumrinde, Ten, 1921 (FH, JE,
S); Ten 9 (S); Ten, 1924 (H). Prope urbem Yunnanfu, in regionis calide temperatae, truncis viv. Quercuum ad templ. Helungtang, c. 1950 m, Handel-Mazzetti 260 (H, JE, S).
SRI LANKA. Without precise locality, Thwaites, s.n. (S); Thwaites 248 (S, BM, H). An Heinen in der Bathschlucht bei Hakgala, 1300 m, Herzog 139 (H, L); 138 (JE).
THAILAND. Payap, granitic massive Doi (Mt.) Inthanon, on branches of Quercuus in clearing, 980 30' E, 180 35' N, 1750 m, Touw 9845 (L). Payap, limestone massive Doi (Mt.) Chiengdao, deciduous forest on NNE slope, on tree trunks, 980 55' E, 190 25' N, 1000 m, Touw 8903 (L).
MALAGASY REP. Mt. Tsaratanana, 1200-2400 m, Bathie, 1924 CFH, S, JE, H, BR). Mt. Ankaratra, Borgen, 1877 (NY, O, BR, PC); Borgen, s.n. (NY, O, L, BM, C, H, S).
MALAWI. Vernoy Nyasaland, Luchenya Plateau, Mlange Mt., Mlange dist., on trunks of trees in dense forest shade, 1890 m, Brass 16537
(NY).
GUINEA. In jugo Danguina, 1100 m, Robegrin, s.n. (O, S, L, GL, H, BR, PC); 1800 m, Robegrin, 1843 (H).
FERNANDO PO. Pic Clarence, 2000 m, Newton, 1894 (S, H).
CAMEROON. Bamenda, Lake Bambaluwe, on tree trunks in montane
forest, Richards R5250 (GL, L, BM). Victoria div., below Likonge, on stem in montane forest, 6500-7000', Richards R4264 (BM, L). Cameroon Mt., on trunks of fallen trees, 5000', Dunlap, 1926 (BM); on trunks of




33
trees near timberline, 7000', Dunlap, s.n. (BM).
MEXICO. Oaxaca, vor dem Pass oberh. Teotitlan in der Sierra, in epiphytenreichen Bergwald an Barke, 2100 m, Dull 92 (MICH). Puebla, Tezuitlan, on wet banks, 7000', Pringle 15285 (H, FH, NY, LD, O, S, L, MANCH, BM, C, PC); Tulancingo Road west of Huauchinango, bark of alder, 5000', Sharp 887 (TENN); on Carpinus, 5000', Sharp 913a, 911B (TENN). Chinantla, Liebmann, s.n. (O, BM, C, H). Michoacan, vicinity of Morelia, Cerro Azul, 2300 m, Arsene 4543 (FH, 5). Morelos, near Cuernavaca, Pringle 15283 (H), 15671 (S, MICH, PC), 15310 (NY). Chiapas, ditio San Cristobal, "Los Llanos" in silva quercuum, 2200 m, Munch 7464 (NY, FH, H, BM); prope San Cristobal, "Los Llanos," in praeruptis umbrosis, 2500 m, Munch 7462 (NY, BM, H); in montibus supra S. Cristobal, 2100 m, Munch 7282 (H, PC); Cerro Hueytepec, near Las Casas, 8000', Sharp 3283, bark of oak, Sharp 3283a, 3270; between Las Casas and San Gregorio, on oak, 7500', Sharp 4683, 4682a (TENN); SW Jalisco slopes of La Ferreria, above Manantlan, on rotted log, oak forest, 6500', Crum 1041 (MICH, S, TENN, NY). Guerrero, gorge below Ormiltemi, 30 km west of Chilpancingo, 6300', Sharp 1149a (TENN); Rancho del Cielo above Gomez Farias, Tamps., at the mine, fallen log, meso, partial shade, Sharp 3618 (TENN). Orizaba, no collector cited, 1857 (NY); No. 68 (BM), Borgeau, 1866 (PC). In planitic Orizabac in sylving ad 6500', Mohr, 1856 (0). Without precise locality, no collector cited, 1865, (NY); without precise locality, Liebmann, s.n. (BM); Totutle, ad hunc ono pertinet fruches perfecti et caespes flaveferis, Liebmann 8467 (C).
PANAMA. Chiriqui, El Volcan, 6500', Llano, 1952 (HIRO); vicinity of El Boquete, 1000-1300 m, on tree trunk, Maxon 4985 (NY).
GUATEMALA. Quetzaltenango, on oak, 7800', Sharp 2063 (MO, TENN);




34
7700', Sharp 2086 (TENN). Chimaltenango, on oak above Tecpam, 8300', Sharp 2571 (TENN). Coban, Alta Verapaz, in arbore vetusta, 1310 m, Turckheim, Broth 6744 (NY, FH, LD, S, BM, H, PC, FI).
BRAZIL. Caldis, Lindberg, 1854 (NY). Prov. Sao Paulo, rariss ad truncos arb. pu. silv. primae. propr opp. Santos, Lindberg, 1854 (O, S, BM, H). Prov. Minas Geraes, Serra de Caldos ad radices arb in suargine paludis, Mosen 365 (NY, S, H, Z, FH).
BOLIVIA. Below Pelichuco, on rock, 3000 m, Williams 2851 (NY, H, BM).
ECUADOR. Andes Quitenses, Antombos, Pastasa River, 5000', Spruce 1441, 1437 (NY, MPU, O, S, BM, H, BR, PC).
Rhegmatodon polycarpus is readily distinguished by 1) highly dotted endostome teeth, 2) a smooth seta, and 3) exothecial cell walls regularly thickened longitudinally, with the transverse walls appearing thinner. The margins of the branch leaves are completely entire. The perichaetial leaf margins with or without teeth. The shape of the branch leaves is highly variable. The branches range from thick and clavate to filiform.
This species has a pantropical and subtropical distribution.




Table 2. Data on three traditional leaf characters formerly used to differentiate smooth-setaed Rhegmatodons.
O 0 0 4 -P 4
Rh. newtonii (H) 1.85 1.52 2.05 .985 mm .728 mm 1.140 mm .532 mm .416 mm .624 mm
.H .i.it)- 4 4
-I4J 4~J 4 :1 01
Rh. schlotheimioides (H) 1.89 1.50 2.35 .690 mm .478 mm .832 mm .364 mm .249 mm .457 mm
var. minor
Rh. brasiliensis (H) 2.04 1.80 2.36 1.027 mm .624 mm 1.206 mm .503 mm .353 mm .561 mm
rqM d rd rz (a (v rzS~
tpa 4 4-4 *H'- -4iLW-44- -H 44 *H4~ 44-H 4-H 4
4d rdr a)r > )) > > a
Rh. brevicuspis (B) 2.05 1.5279 2.0538 1.112985 mm .72998 mm 1.14260 mm .53240 mm .436 mm .624 mm
Rh. schfilirameus imiodes (H) 2.08 1.5079 2.3542 .69750 mm .47665 mm .936 mm .36459 mm .249 mm .457 mm
var. minor
Rh. brasiliensius (0) 2.08 1.78 2.3645 1.112 mm .62873 mm 1.20689 mm .534 mm .35374 mm .603 mm
Rh. orizabanusis (BR) 2.0512 1.95 2.33 .877112 mm .99728 mm .936 mm .413 mm .312 mm .6478 mm
Rh. fischlotheimioideseus (N) 2.18 1.7981 2.4269 .75896 mm .7695 mm 1.040936 mm .35420 mm .374 mm .478 mm
Rh. cammeruniaes (H) 2.0819 1.7891 2.46 .827 mm .72873 mm .956 mm .376 mm .37412 mm .41603 mm
Rh. crassirameizabanus (NY) 2.1230 1.85 2.3364 .87967 mm .832 mm 1.144936 mm .41320 mm .291 mm .47561 mm
Rh. schypnothimoides (N) 2.1833 2.0681 2.6947 .89757 mm .624 mm .894 mm .32420 mm .37249 mm .47816 mm
Rh. crizabanusniae (B) 2.1934 1.95 2.64 .902 mm .72869 mm .936 mm .37691 mm .332 mm .41678 mm
Rh. dncrassiraeus (NYBM) 2.37 2.13.85 2.6471 .688 mm .83520 mm 7.900144 mm .291 mm .29128 mm .56312 mm
Rh. hypnfiiformides (C) 2.38 2.0611 2.4786 1.02757 mm .832 mm 1.1854 mm .434 mm .3129 mm .5406 mm
Rh. orizabthostangius (BR) 2.38 2.195 2.65 1.114902 mm .76928 mm 1.248936 mm .39468 mm .312 mm .47561 mm
Rh. denspolycarpus () 2.38 2.17 2.64 1.15488 mm .52936 mm 1.31900 mm .29484 mm .22353 mm .6032 mm
Rh. fipolysetuiformis (N) 2.3842 2.13 2.76 1.025 mm .83294 mm 1.18564 mm .43422 mm .332 mm .499 mm
Rh. crassirameius (BR(H) 2.3845 2.1522 2.75 1.11052 mm .832 mm 1.144 mm .428 mm .31274 mm .56120 mm
Rh. palustrips (H) 2.3846 2.18 2.6493 .925 mm .7696 mm 1.31040 mm .48376 mm .35312 mm .60457 mm
Rh. handpolysetusii (HN) 2.48 2.18 2.7681 .859 mm .89665 mm 1.164 mm .3452 mm .3270 mm .49916 mm
Rh. brasiliensis (S) 2.48 2.2217 2.7588 1.092 mm 1.019832 mm 1.164 mm .438 mm .375 mm .52478 mm
Rh. palustris (H) 2.46 2.18 2.93 .925 mm .769 mm 1.040 mm .376 mm .312 mm .457 mm
Rh. handelli (H) 2.48 2.18 2.81 .859 mm .665 mm .956 mm .345 mm .270 mm .416 mm
Rh. brasiliensis (S) 2.48 2.17 2.88 1.092 mm 1.019 mm 1.164 mm .438 mm .375 mm .478 mm
Rh. polycarpus (NY) 2.53 2.33 2.84 1.110 mm 1.081 mm 1.248 mm .438 mm .353 mm .499 mm
Rh. pringlei (H) 2.55 2.25 2.84 .834 mm .707 mm .936 mm .326 mm .270 mm .374 mm
Rh. orthostegius (L) 2.58 2.14 3.05 .875 mm .624 mm 1.081 mm .339 mm .270 mm .395 mm
Rh. filiformis f. major (BM) 2.83 2.18 3.14 1.027 mm .915 mm 1.144 mm .361 mm .291 mm .457 mm




DISTRIBUTION AND HABITAT OF THE GENUS RHEGMATODON
Rhegmatodon declinatus has only been collected from the tropics and subtropics of the Old World (Fig. 12:2). In contrast, Rh. polycarpus (Fig. 12:1) has a pantropical distribution ranging from Central and South America to Africa, Madagascar, India,and China. Unlike Rh. declinatus, Rh. polycarpus has not been collected in Indonesia and Malaysia. The ranges of these two species overlap in China (Yunnan), Sikkim (Himalaya), Assam (Khasia), Ceylon, and Thailand (Mt. Chiengdao). The areas of collection are characterized by altitudes of 650-1400 m.
The disjunctive distribution of Rh. polycarpus is similar to that
of other moss genera such as Pilotrichella, Squamidium, Lindigia, Braunia, and Dimerodontium. Herzog (1926, 1932) uses the continental disjunctions of certain mainly austral families of vascular plants, with similar patterns of distribution in support of Wegener's (1924) theory of continental drift. The disjunctive distribution of Rh. polycarpus is very similar to that of Leptodontium viticulosoides and vars. (Zander, 1972).
Rhegmatodon is most often found at the middle and high altitudes
in the mountainous regions of the tropics and subtropics. Rh. declinatus grows at lower altitudes than Rh. polycarpus. Rh. declinatus has been found on the bark of fallen logs or trees, and occasionally on moist shady rocks, most often between 500-1700 m. Rh. polycarpus, usually found between the altitudes of 1400-2500 m is also corticolous, but may be collected on wet ground and rocks.
36




DISPOSITION OF NAMES OTHER THAN SYNONYMS IN RHEGMATODON
The following list includes the names of in schedula specimens, as well as validly published specimens, where examination of the types showed them to belong to some other taxa than Rhegmatodon. Most of these specimens were collected in areas for which adequate keys are not available, and only a limited attempt was made to assign these taxa to genera. For the purpose of completeness, previous exclusions and transfers are included in this list.
Rhegmatodon bornmulleri Broth. in schedula, 1904. TYPE: Brazil,
Cruz Alta, Colonia Nova Wurtemburg, Elsenau, in arboribus,
450 m., Bornmuller 6156 (holotype, JE: isotypes, B, S). This
differed from Rhegmatodon by having no exostome and highly
papillose, bifid endostome teeth. Most likely this is
Dunerodontium mendozense Mitt.
Rhegmatodon chryseus Schimp nom. nud. in synon. E Rozea chrysea
Besch. in Mem. Soc. Sc. Nat. Cher. 16:242. 1872. And Besch. in Fourn., Mex. Pl. 1 (Crypt.):48. 1872. TYPE:
Mexico, no collector cited, s.n.
Rhegmatodon fissidens Thdr., in schedula, 1905. TYPE: China,
Prov. Kong Tiheau, Carabrie 3629 (holotype, H). Laminal
cells all subquadrate. This is probably a Leskea sp.
Rhegmatodon fusco-luteus Besch., Mem. Soc. Sc. Nat. Cherbourg
16:232. TYPE: Mexico, Orizaba, Muller, s.n., in herb.
37




38
Schimper (not seen) = Platygyrium tuscoluteum Card. cf.
Card, Rev. Bryol. 38:40. 1911.
Rhegmatodon minutus (Mitt.) Par., Ind. Bryol. 1109. 1898. TYPE:
Andes, Jameson, s.n., (not seen) : Helicodontium minutum
(Mitt.) Jaeg., Ber. S. Gall. Naturw. Ges. 1876-77:225.
1878 (Ad. 2:291).
Rhegmatodon nietneri C. Muell., Linnaea 36:20. 1869. TYPE:
Ceylon, Nietner, s.n. (isotype, 0). The areolation of the lamina was not like Rhegmatodon. I was unable to identify
this to genus.
Rhegmatodon parvulus Hamp. Icon. Musc. 2:14. 1844. TYPE: Georgia,
Eleenezar near Savannah River, no collector cited, s.n.,
(not seen) E Clasmatodon parvulus (Hamp) Sull. in Gray,
Man. Bot. N.U. States ed. 2:660-5. 1856.




INFRAGENERIC RELATIONSHIPS IN MACROHYMENIUM
The taxonomic treatment of Macrohymenium by Brotherus (in Engler and Prantl, 1925) is based on F. C. Kiaer's revision of 1882, Genera muscorum Macrohymenium et Rhegmatodon revisa specieque nova aucta exposuit. Kiaer based his key totally on gametophytic characters in order to include M. sinense. This specimen has subsequently been placed in the genus Giraldiella. Thus Brotherus also separated the various specimens of Macrohymenium on the basis of gametophytic characters.
In this present study, many of these characters are excluded.
The following were found to be variable, varying even within the same specimen: denticulations on the margins of the perichaetial leaves, loose to densely imbricated leaves, and leaves heteromallous to homomallous.
Some gametophytic characters are useful in determining species of Macrohymenium, e.g., leaf size, shape, and areolation. M. rufum leaves are the smallest (.79-1.41 mm long) and widest (2.27) ovate and abruptedly constricted to a short acumen (Fig. 8:13-18). In M. acidodon, branch leaves are small (.89-1.45 mm) (Fig. 9:10-13) and more narrow than M. rufum (2.77). M. acidodon can be differentiated from the other species by virtue of the uniform size of the leaf cells (Fig. 9:2,3,7, 9). Each of the other species has leaf cells which are much longer at
For the purpose of comparison, translations of both the keys from both works are presented in Appendix II.
39




40
the base and gradually shorter toward the tip. M. mulleri is the largest of the species, the leaves the longest (1.9-2.5 mm) and the most narrow (4.0) (Fig. 10:11-14). It also has more enlarged alar cells (12-20) (Fig. 10:19). The other species have from 6-9. M. strictum has slightly larger leaves (1.0-1.43 mm) than M. acidodon and can be distinguished by a sudden constriction to a long acumen (Fig. 11:2-3).
Brotherus did include one sporophytic character in his key,
separating his six species into two equal groups on the basis of the seta plainly rough versus the seta set above with very low, broad warts. While some setae may appear rough when dry (Fig. 6:1), moistened setae of all species are smooth at the base and towards the top have a few low, broad projections (Fig. 6:11-12). Neither Kiaer nor Brotherus used the presence or absence of perforations in the endostome teeth as key characters. Probably, this was due to Kiaer's statement that he had found both perforate and imperforate endostome teeth on the same specimen. In every specimen examined during this study this character was discrete. The endostome teeth were either perforate or imperforate, and never differed within the same endostome.
Of the four species recognized in the present work, two have
perforated endostomes and two are imperforate. The two species which have perforations are also quite different in the expression of the character. M. mulleri has very regular perforations in nearly every segment, producing a scalloped effect. M. mulleri also has the shortest peristome, the endostome ranging from 250-300 pm and the exostome from 190-200 pm (Fig. 10:1-3). The seta is moderately long (811 mm) and quite thin (130-150 pm). On the other hand, M. acidodon has irregular perforations in the upper portion of the teeth, sometimes




41
nearly continuous (Fig. 9:4-5). The peristome is moderate in size, the endostome ranging from 350-500 Pm and the exostome from 200-250 Pm. The seta is 9-11 mm long and 115-180 pm wide. The endostomes of M. rufum (Fig. 8:4-5) and M. strictum (Fig. 11-4-5) are imperforate. The peristome of M. strictum is much larger (endostome, 550-750 urm; exostome, 300-410 pm) than that of M. rufum (endostome, 440-600 pm; exostome, 170300 pm). The seta of M. strictum also is much longer (8-21 mm) and wider (200-374 Pm) than that of M. rufum (6-6.5 mm long and 160-220 Pm wide).




DESCRIPTION OF THE GENUS MACROHYMENIUM
Plants lustrous, yellow-brown to reddish green, in moderately compact, low sods, copiously fruiting. Pleurocarpous mosses, prostrate and rooting. The sporophytes lateral at base of the branches. Goniautoicous. Primary stems thin, stolon-like, zigzag, creeping, lacking a central strand, copiously branched. Branches typically simple, ascending, short, thick and round, straight to curved, densely foliose; leaves imbricate, heteromallous to somewhat homomallous. Paraphyllia or pseudoparaphyllia lacking. Branch leaves ecostate, concave, ovate-lanceolate, acuminate, with the leaf margins entire, more or less recurved. Laminal cells elongate, oval, walls incrassate osteoform with thin connecting areas between cells. Alar cells rectangular and quadrate, colored and somewhat inflated. Leaves .792.53 mm long, and .29-.66 mm wide. Inner perichaetial leaves large ecostate or faintly hinting of a costa, upright, sheathing, ovatelanceolate, acuminate, or somewhat blunt. Perichaetium rooting. Vaginule small, oblong cylindric. Perigonium axillary, hidden between the stem and branch leaves, budlike, very small, 1/3 length of leaves. Perigonial leaves broadly ovate, acuminate, ecostate, entire. Antheridia many, pedicellate, short stipitate, oblong-cylindric, with small paraphyses sparce or lacking. Perichaetium axillary, inserted in the axils of the branchlets, budlike, many-leaved, and obscured between the leaves. The archegonia short, moderately robust, with very few short, filiform 42




43
paraphyses. Setae flexuous or twisted in the dry state, short to long, 6-21 mm, smooth at base, but with low hyaline protuberances below the capsule. Capsule, erect-inclined, constricted below the mouth, oblong, reddish brown. Exothecial cell walls irregularly thickened, appearing collenchymatous. Operculum conic rostrate, nearly equalling the capsule in length. Peristome double; exostome of 16, equidistant, erect-incurved teeth, inflexed in the dry state, broad at the base with two striated dorsal plates separated by a zigzag line, suddenly constricted to a long ciliate tip, the zigzag line continuing to the tip. Dorsal plates backed by a single row of trabeculate plates. Endostome of 16 teeth, from a low basal membrane, without cilia, the processes about 2X longer than the exostome teeth, keeled, entire, either perforate or imperforate on keeled midline. Membrane subplicate, tesselated. Spores globose and papillose, 15-27 pm in diameter. Calyptra cucullate.
Margadant (1959) posed a nomenclatorial problem with reference to the genus Macrohymenium. Fleischer (1923) synonomized Leskea? rufa Reinwardt and Hornschuch with Acroporium braunii (C. Mull.) Fleisch. Because Leskea rufa was an earlier collection than the type for A. braunii the name was changed to Acroporium rufum (Hornsch. and Reinw.) Fleischer. Fleischer said he had examined the type for Leskea rufa in the Berlin Herbarium. Believing this specimen to be the basionym for Macrohymenium rufum, the type species of the genus Macrohymenium, Margadant felt that this transfer threatened the nomenclatural standing of Macrohymenium. He therefore proposed that Macrohymenium acidodon (Mont.) Doz. and Molk. (1848) be designated the lectotype and the name Macrohymenium be conserved. All of this is based on the assumption that Carl Muller used Leskea rufa Reinwardt and Hornschuch as the type specimen for his new genus. There is good reason to believe that this was




44
not the case.
The original specimen of Leskea rufa was described in Nov. Act. Caes. Leop. 14: Suppl. 2: pp. 716-17. 1826. It was collected by Reinwardt on Mt. Malabar (Malabaria), Java. In 1847, when erecting the new genus, Macrohymenium, Muller cited three specimens in synonomy. The first was Leskea rufa Reinwardt and Blume, Java. This same citation is found in Synopsis Muscorum Frondosorum 2:30, 1850, when Muller dissolved Macrohymenium and moved M. rufum into the genus Rhegmatodon as Rh. rufus. In neither of these citations does Muller ever mention Malabaria. This suggests that his type for M. rufum was in fact based on another specimen. There are several facts which support this idea.
The original description of Leskea rufa Reinwardt and Hornschuch, states that the peristome was destroyed. Yet MUller's genus description contains a detailed description of both the endostome and exostome; neither of which could be mistaken for Acroporium.
Further support issues from the synonomy of Dozy and Molkenboer (1848) in which they list two specimens from Java; Blume and Reinwardt, Java, and a separate Reinwardt, Mt. Malabar.
Java: collegerunt Blume et Reinwardt, monente Mllero:
Malabaria: collegit Reinwardt (cf. Nov. Acta Acad. Caes.
Leop. 1. cit.). (Dozy & Molk., 1848, p. 167.)
Note that the first specimen was "collected by Blume and Reinwardt, with advice by Muller," while the Malabar listing only mentions Reinwardt as the collector, and by the accompanying citation gives clear indication that this second specimen is the Leskea rufa of 1826. Whenever the Mt. Malabar specimen is cited, Reinwardt's name is used alone, never in conjunction with Blume.
There is sufficient evidence to support the existence of a separate specimen from Java, collected by Blume & Reinwardt, on which the type




45
description of M. rufum is based. Fleischer, recognizing that the specimen from Mt. Malabar was actually an Acroporium, placed it in synonomy with Acroporium braunii without jeopardizing the nomenclatorial standing of Macrohymenium.
One other possibility exists. Muller may have described his new genus using a combination of characters from the three specimens in his synonomy. This would assume the Mt. Malabar collection to be the same as the Blume and Reinwardt collection. If so, his description of the peristome could only be that of L. mitrata since no perforations of the endostome are noted. L. acidodonI has definite perforations, for which reason it was taken out of synonomy and designated the type for Macrohymenium acidodon (1848). I recently received a collection of Macrohymenium from Jena (JE) which contained a specimen with the following data:
Rhegmatodon rufus mihi!
Java: Blume
Vereinigte Herbarien Karl Schliephaeke, Osterfeld und
Hermann Winter, Gotha.
This specimen is clearly a M. rufum and has two sporophytes in excellent condition. While there is no date, I believe this is the missing specimen of Blume and I am designating it as the lectotype for M. rufum. If futher evidence negates my argument, I recommend that Leskia mitrata Doz. and Molk. be designated the lectotype for Macrohymenium rufum. For reasons stated above, Leskia mitrata Doz. and Molk., Musci Frond. ex Archip. Indici. 1844, p. 15; Ann. des Sc. Nat. 1844, p. 511, is the earliest described specimen of Macrohymenium rufum C. Muller.2
1Assuming Muller saw the 1845 specimen of Doz. and Molk.
2Granting the above argument, this would be the proper citation for M. rufum.




KEY TO THE SPECIES OF MACROHYMENIUM
. Endostome teeth with perforations in the keeled midline,
at least from mid-tooth to tip ...... ............. 2
1. Endostome teeth with no perforations of the keeled midline
............... .......................3
2. Perforations even, uniform, giving a scalloped appearance from the base to the tip; large branch
leaves (1.9-2.5 mm long) with long basal cells (12:1),
the cells gradually shortening to the tip (3:1)
......... .................... ...M. miilleri
2. Perforations uneven, irregular, restricted mostly to
the upper half of the teeth, sometimes appearing bifid
at the tip; smaller branch leaves (0.9-1.5 mm long),
the cells of uniform size from the base to the tip of
the leaf (5:1 3:1) ... .......... M. acidodon
3. Branch leaves lanceolate, gradually acuminate; perichaetial leaves abruptly constricted to acuminate tip;
seta ranging from 8-21 mm in length. . M. strictum
3. Branch leaves from a squarish base, widely ovate-lanceolate,
abruptly acuminate; some of the perichaetial leaves blunt;
seta ranging from 6-7 mm in length. . M. rufum
46




47
1. MACROHYMENIUM RUFUM C. Muell., Bot. Zeit. 5:825. 1847. TYPE:
Java, without precise locality, Blume and Reinwardt, s.n. (lectotype, JE).
(Figure 8)
Leskea tufa Reinw. and Hornsch., Nov. Act. Acad. Caes. Leop. Carol.
14, p. 2 suppl. 1826, p. 716-717. TYPE: Java, Mt. Malabar,
Reinwardt s.n. (not seen).
Leskea mitrata Doz. & Molk., Ann. Sc. Nat. 3 ser. 2. p. 311. 1844.
TYPE: Sumatra, Korthals, s.n. (holotype, L). New synonomy.
Rhegmatodon rufus (Reinw. and Hornsch.) C. Muell., Syn. 2:30. 1850.
TYPE: Java, Blume and Reinwardt, s.n.
Macrohymenium curvirostrum, Braun, in schedula. TYPE: Java,
Braun, s.n. (holotype, BM). New synonomy.
Plants small, the leaves .79-1.41 mm long and .31-62 mm wide, with
an average length to width ratio of 2.27. The leaves ovate and abruptedly constricted into the acumen. The leaf cells longer at the base of the leaf (10-12:1), gradually shortening towards the tip (2-3:1). Six to nine inflated quadrate colored alar cells. Usually, one or two of the inner perichaetial leaves with a very blunt tip. Seta from 6-6.5 mm in length and 160-200 pm in width. The endostome teeth lacking perforations on the midline. The endostome teeth ranging from 440-600 pm, and the exostome teeth ranging from 170-300 pm.
Habitat: On tree trunks in forests,epiphytic, usually found between 1500-2000 m, but occasionally up to 3030 m.
Distribution: Java, Sumatra, Borneo, New Guinea, Australia, Sri Lanka, Malagasy Rep.




48
JAVA. Tjibodas, mountain garden, Meijer, 1953 (L); Nurta, 1953
(L). Between Tjibodas and Tjihoerang, in Eupaternia pallescens plain at Koebang tree trunk in the sun, 1380 m, Soekar 2247 (L). Namtrong Djimter, Jeger Bentang, rainforest tree trunk in the sun, 1450 m, Soekar 3187 (L). G. Sindoro, rainforest, 2400 m, Leeuwen 8889 (L). Without precise locality, Gerker, s.n. CL); Braun, s.n. CL). Without precise locality, no collector cited, s.n. (BR). Without precise locality, Blume, s.n. (JE)
SUMATRA. Mt. Sogo, near summit, mossy forest, 2000 m, Meijer 6126
(L). Korthals, s.n. (L, H, O, S). West coast, Mt. Merapi, west side, Prim. forest on branch, Waalkes 2236 (L); 1700 m, Belukar, on trunk, Waalkes 2270 CL). Aek na Vli, near Toba Lake, in mixed forest on bark, 1400 m, Wijk 1870 (L). Without precise locality, no collector cited, s.n. (S, C).
BORNEO. E. Kutai, peak of B. papan, terr. Beul, on Magnoliaceae, 600-700 m, Meijer B1968 (L). N. Borneo, west coast res, Mt. Kinabalu, near Paka Cave, c. 9700', Meijer Bll-903 (L).
NEW GUINEA. Star Mts. Mt. Antares, on rotten trunk of tree in
rain forest, 1500 m, van Zanten 385b (BM). Eastern Highlands District, Arau, low on a tree in Castanopsis, oak forest, 1400 m, Brass 31976a
(L). Sepik District, Ambunti Subdistrict, summit of Sumset, Mt. Hunstein, in elfin woodland on summit plateau, epiphytic on upper branches of small tree, c. 5000', Hoagland and Craven 10985 (L). Western Highlands District, Wabag area, Ranges south of Wabag, Nothofagus lower montane rain forest, corticous, 8000', Robbins 2859 CL).
AUSTRALIA. Queensland, Ravenshoe, Watts 660 (H).
SRI LANKA. Beim Stausee von Kandy, c. 55 m, Herzog, 1906 (BM).




49
MALAGASY REP. "Certainly from Madagascar," no collector cited, s.n. (BM).
M. rufum has the smallest and widest leaves of any of the species. The widely ovate leaves, with a short abrupt acumen, are easily recognized. The imperforate endostome is similar to that of M. strictum but smaller, and the seta is the shortest of all Macrohymenium species. M. rufum is the only species in which some of the inner perichaetial leaves are blunt.




50
2. MACROHYMENIUM ACIDODON (Mont.) Dzy et Molk., Musci Frond. Ined.
Archip. Indici 6:168. 1848.
(Figure 9)
Leskea acidodon Mont., in Ann. sc. nat. 1845, p. 96, t. 5, f. 4
and Syll. p. 19. 1845. TYPE: Rdunion, no collector cited
s.n. (holotype, L).
Macrohymenium laeve Thwait. and Mitt., J. Linn. Soc. Bot. 13:317.
1873. TYPE: Ceylon, Thwaites 236 (holotype, NY-MITT; isotypes, O, H-BROTH, PC, BM). New synonomy.
Macrohymenium acidodon var. acutissima Besch. ex Par., Ind. Bryol.
1108. 1898 nom. inval. in synon. err. pro M. a. fo. Besch., Ann. Sc. Nat. Bot. ser. 6, 10:292. 1880. TYPE: Madagascar,
Perville, s.n. (holotype, L; isotype, L, H-BROTH). New
synonomy.
Rhegmatodon madagassus C. MUell. ex Geh., Abh. Naturw. Ver. Bremen
7:211. 1882. TYPE: Madagascar. forest of Ambatondrazaka,
Rutenber, s.n. (holotype, 0).
Rhegmatodon acidodon (Mont.) Par., Ind. Bryol. 1108. 1898.
Rhegmatodon laevis (Thwait. and Mitt.) Par., Ind. Bryol. 1109.
1898.
Rhegmatodon acidodon var. acutissima (Besch.) Par., Ind. Bryol.
1108. 1898.
Plants with leaves, .89-1.45 mm long, .29-.58 mm wide, with an average length to width ratio of 2.77. Leaf cells uniform in length from the base to the tip of the leaf. Six to eight golden-colored inflated quadrate alars. Perichaetial leaves slightly serrate, ecostate or with a faint trace of a costa on some of the large inner leaves.




51
Seta from 9-11 mm in length, 115-180 pm in width, slightly twisted, quite smooth at the bottom with hyaline protuberances toward top. Endostome teeth perforated on their midline with uneven irregular perforations. The perforations are mostly in the top half of the tooth, occasionally producing a bifid tip. Endostome teeth ranged from 350-500 pm and the exostome teeth ranged from 200-250 pm.
Habitat: On branches and rocks, 1800-2500 m.
Distribution: Malagasy Rep., Reunion, Seychelles, Sri Lanka, N. Borneo.
MALAGASY REPUBLIC. Forest of Ambatondrazaka, Rutenberg, s.n. (0); Hildebrandt, 1880. Without precise locality, no collector cited, s.n. (0, L); Hildebrandt, 1880 (L); Perville, 1879 (H, 0).
REUNION. Without precise locality, Lepervanche, 1879 (0, BM, H,
PC); no collector cited (O, L); Richards (BM); De L'Isle (PC); Rodriguez
(PC); Plain au Cafres, De L'Isle (BM).
SEYCHELLES. Without precise locality, De L'Isle (PC).
SRI LANKA. An baumen beim Rambotta pass, c. 2000 m, Herzog 115 (H, JE, L). Central Province, Thwaites 236 (T, PC, NY, O, BM). An Baumen beim Tec von Nuwara Eliya, 1800 m, Herzog, s.n. (PC, S). Neuwara Eliya, rotting stump in jungle, Binstead 183 (BM).
BORNEO. N. Borneo, West Coast Res. Mt. Tambuyokon, c. 15 miles
NE of Kinabalu Peak, summit area on branches and rocks, c. 8000', Meijer Bll-378 (L).
M. acidodon differs from all other species by the uniform size of the leaf cells from base to tip. The endostome of M. acidodon is also uniquely perforated with irregular, uneven slits, mostly in the upper half of the precesses. This sometimes produces a bifid appearance.




52
3. MACROHYMENIUM MULLERI Doz. and Molk., Musci Frond. Ined. Archip.
6:168. 55. 1848. TYPE: Java, Mt. Gede and Talaga-Bodas,
Korthals, s.n. (cotypes, O, C, H, S, L). Borneo, Mt. Sakoembang, Korthals, s.n. (lectotype, L). Sumatra, Batang-Bessie, Korthals,
s.n.
(Figure 10)
Rhegmatodon mulleri (Doz. and Molk.) C. Muell., Linnaea 36:21.
1869.
Macrohymenium mulleri forma pubea Dix., in schedula, 1909. TYPE:
Java, Mt. Pangerango, Palmer & Bryant 982a (holotype, BM).
New synonomy.
Plants large, the branches thick and curved, leaves 1.9-2.53 mm long and .42-.66 mm wide, with an average length to width leaf ratio of 4.0. Leaf cells at base much longer (10:1) than cells at the tip (3:1). Twelve to twenty inflated, colored, quadrate alar cells. Seta from 811 mm in length, 130-150 pm in width. The endostome teeth perforated on the midline nearly the whole length with very uniform and regular perforations giving a scalloped appearance to the midline. Occasionally the tip of a tooth is bifid. Endostome teeth ranging from 250-300 pm, the exostome teeth from 190-200 pm.
Habitat: On trees and ravine in very moist forest, usually found at elevations from 1200-1800 m, but occasionally up to 3100 m.
Distribution: Java, Borneo, Malay Peninsula, and Malagasy Rep.
JAVA. Mt. Pangerango, 4-6000', Motley, 1906 (NY); 7-10,000',
Motley, 1906 (BM); Palmer & Bryant 982a (BM). Prov. Preanger, In decliv. austral. montis Pangerango, in horto montano Tjibodas, Regio nubium,




53
1420 m, Schiffner 12729 (S); in Cinchoneto, Daradjat prope Garut ad truncos putridos, Regio nubium, 1730 m, Schiffner 12127 (L). Mt. Salak, Binnendijk (S, L). Rawa gajonggong Tjibeureum, plain of gajongong, tree trunk in the sun, 1700 m, Soekar, 1949 (L). West Java, Tjibodas, Mt. Gede, Batu gonggang, Nature reserve, in rain forest, on Vernonia arborea base of trunk, shadowed, 1600 m, VanOostatroom 14235 (L). Without precise locality, Schroted, 1927 (H).
BORNEO. In Sakoembang, Korthals, s.n. (L). Kinabalu, Low, 1859
(BM); Below Kamborangah, ravine in wet montane forest, 6000', Richards R5758 (L).
MALAY PENINSULA. Gunong Tahan, Pahang, Ridley 1035 (BM).
MALAGASY REPUBLIC. Without precise locality, no collector cited, s.n. (S).
M. mUlleri is distinguished from the other species of Macrohymenium by its large size, and its long, narrow leaves with many alar cells. The endostome of M. mUlleri is differentiated by its peculiar pattern of perforations, and by its generally smaller peristome. Although M. mUlleri is the most robust of all the species, it is marked by the smallest peristome and a very thin seta.




54
4. MACROHYMENIUM STRICTUM Bosch. and Lac., in Doz. & Molk., Bryol.
Jav. 2:114. 1865. TYPE: Borneo, Mt. Kinabalu, Low, s.n. ex
herb. Hook (holotype, NY; isotypes, O, S, L).
(Figure 11)
Rhegmatodon strictus (Bosch. and Lac.). C. Muell., Linnaea 36:21.
1869.
Macrohymenium nova-guineense Reim., Hedwigia 69:129, 1929. TYPE:
Nova Guinea neerlandica, Mt. Doorman, Lam 1701 (not seen).
New synonomy.
Macrohymenium acidodon (Mont.) Doz. et Molk. fo. longisetum, in
schedula. TYPE: Madagascar, Massif de L'Anjanaharibe,
Humbert, Capuron and Cours, 1951 (holotype, PC). New
synonomy.
Plants moderate to large, the leaves 1.0-1.43 mm long, and .37-.55 mm wide, with an average length to width ratio of 2.7. The leaves ovatelanceolate with a long acumen. The leaf cells generally uniform in size over most of the leaf (3:1), much longer at the base (10-12:1). Eight to twelve inflated, quadrate, colored alars. All of the perichaetial leaves with acuminate tips. Seta from 8-21 mm in length and 200-372 pm in width. Endostome twice as long as the exostome; endostome teeth not perforated. Endostome teeth ranging from 550-750 pm; exostome teeth ranging from 300-410 pm.
Habitat: On trees and ground in subalpine forests and open thickets, occasionally found 1000-2000 m, but usually above 3000 m and up to 4370 m.
Distribution: Malagasy Rep., Seychelles, Sri Lanka, Borneo, Sarawak, Sumatra, New Guinea, Australia, and the Philippines.




55
MALAGASY REP. Massif de L'Anjanaharibe (Pentes and Sommet Nord),
a L'ouest D'Andapa Haute AndraMonta, Bassin de la Lokoho, nord-est, foret ombrophile sur gneiss et grantie, 1000 m, Humbert, Capuron and Cours, 1951 (PC).
SEYCHELLES. Pres la baie Sainte Anne, no collector cited, s.n.
(PC).
SRI LANKA. Nuewara Eliya, foot of tree by road, Binstead 181 (BM).
BORNEO. Mt. Kinabalu, Low, s.n. (O, S, L, NY). North Borneo,
West Coast Res., Mt. Tambuyokon, c. 15 miles NE of Kinabalu, Meijer B11-473; ultrabasic area, 9000', Meijer B10-242 (L). North Borneo, Kota Belud distr., one mile n. of Kambarangan camp on path to Paka Cave, Mt. Kinabalu, common on ground in turf under an open, scrubby growth, averaging 12' high, dominated by Ericaceae, Dacrydium and Phyllocladus,in fairly pure, abundantly fruiting tufts up to 9" diameter, moist and shady, deep humus over shallow, grey loam, 2750 m, Wood 1539 (L).
SARAWAK. Without precise locality, Oxford Exped. (1932) 1631 (BM).
SUMATRA. Sumatra occid, ad declif. occid. montis ignivomi Merapi, regio nubium, 1800-2050 m, Schiffner 12724 (L, PC).
NEW GUINEA. Mt. Albert Edward, common on forest trees, 3680 m, N. Guinea Exped. of Am. Mus. of Nat. Hist. 4431, 4438, 4439, 4435 (NY). Western Highlands, Kubor Range, Mt. Kinkain, on tree trunk, alpine shrubbery, 3580 m, Vink 16178 (L). Western Highlands, Wabog Area, Sugarloaf Mt. area, montane rain forest, epixylic, 10,000', Robbins 2805 (L). Eastern Highlands, Bismarck Ranges, Mt. Wilheim, on trees in grove at base of Bogonota Ridge, SE of Lake Aunde of edge of valley floor, 11,000', Weber and McVean B-32129 (LD, GL); Pindaude, Mt. Wilheim trail, epiphytic on Coprosma sp. in alpine tussock grassland, locally common, 14,000',




56
Weber and McVean B-33522 (TENN); on dead wood in edge of subalpine forest 3700 m, Brass 29994 (L). NE Wilhelmina, top, epiphyte, 3650 m, Brass and Myer-Drees 9775 (TENN). Lake Habbema, enveloping branch in open thickets, 3225 m, Brass 9453 (L, TENN).
AUSTRALIA. Molanda, Queensland, Watts 623, 664 (H). Ravenshoe, Queensland, Watts 662, 657 (H).
PHILIPPINES. SE Mindanao, Todaya, Mt. Apo, on tree, 1220 m, Williams 2675 (NY, H).
M. strictum is distinguished by its larger leaves with a long,
suddenly constricted acumen. It also has the largest peristome of any species. Though the imperforate endostome closely resembles that of M. rufum, it is differentiated by its much larger size. The seta is also much longer and more robust than that of M. rufum, and the perichaetial leaves more acuminate. M. strictum is a species growing at very high altitudes, usually above 3000 m, while M. rufum is found at lower altitudes, from 1700 m down to 600 m.




DISTRIBUTION AND HABITAT OF THE GENUS MACROHYMENIUM
Species of Macrohymenium have been collected only in the Old World tropics (Fig. 13:1-2; Fig. 14:1-2). The genus is essentially insular, occurring from the Malagasy Republic through Sri Lanka and Indonesia to northeastern Australia. Species of Macrohymenium are very similar in their distribution. Every species has been found from both Borneo and the Malagasy Republic, and all but one, M. nileri, from Sri Lanka.
Although the species of Macrohymenium occupy areas very similar in latitude, the altitudinal ranges of each vary widely. M. strictum has been collected primarily above 3000 m. M. acidodon is found at relatively lower altitudes, 1800-2500 m, while M. mrlleri is, for the most part, encountered between 1400-1700 m, but has been reported from as low as 600 m. Macrohymenium is primarily a corticolous genus, although a few specimens have been collected on rocks and soil. Collection data for most specimens are so scarce that it is difficult to be more precise in describing the habitat of the genus.
57




DISPOSITION OF NAMES OTHER THAN SYNONYMS IN MACROHYMENIUM
The following list includes the names of in schedula specimens, as well as validly published specimens, where examination of the types showed them to belong to some other taxa than Macrohymenium. Most of these specimens were collected in areas for which adequate keys are not available, and only limited attempt was made to assign these taxa to genera. For the purpose of completeness, previous exclusions and transfers are included in this list.
Macrohymenium cuspidatum Mitt. in Kiaer, Forh. Vid. Selsk.
Christiania 1882 (24): 45. 1883. TYPE: Assam, Mt. Khasia,
Griffith (No. 212, Hb. Kew. No. 516 intermixed) (isotype, 0). Index Muscorum incorrectly attributes a new species,
Sematophyllum cuspidatum, to Kiaer on the basis of this
specimen. Actually, he notes that the peristome was
destroyed by age and unknown to him, and that it perhaps
belonged to the genus Sematophyllum. Having seen the specimen, I would agree.
Macrohymenium densirete Broth. in Fleisch., Musci Fl. Buitenzorg 4: 1192. 1923 nom. nud. in synon. TYPE: not given
E Clastobryum conspicuum Fleisch.
Macrohymenium gracillimum C. Mull. ex Kiaer, Forh. Vid. Selsk.
Christiania 1882 (24): 19. 1883 nom. inval. TYPE: not
given.
58




59
Macrohymenium kunzii C. Mull., in schedula, 1868. TYPE: India,
Pegu, Yomah, Kunz 2939 (holotype, H). Broad bodies were
present on this specimen. It is clearly in the Sematophyllaceae and probably in the Clastobryoideae.
Macrohymenium sinense Ther., Bull. Ac. Int. Geogr. Bot. 19:20.
1909. TYPE: China, Prov. Kweitschou, Lou-mong-touan,
Fortunat, 1903 (holotype, S) = Giraldiella levieri C. Mull.
fid. Reim., Hedwigia 71: 62. 1931. I have seen this specimen and agree.
Macrohymenium wichurae Broth. in Fleisch., Musci Fl. Buitenzorg
4: 1364. 1923 nom. nud. in synon. TYPE: not given.




LEGENDS FOR THE FIGURES
To facilitate reference and comparison of the characters presented in Figures 1-11, the following standardized magnifications have been used:
Habits X8 Spores X466
Calyptrae X30 Leaf tips X164
Opercula X30 Alar cells X164
Endostome teeth X125 Basal cells X164
Exostome teeth X125 Juxtacostal cells X164
Exothecial cells X164 Antheridia X117
Setae X117 Archegonia X117
Perichaetia X30 Portion of branch X30
Vaginules X30 Perichaetial leaves X30
Pseudoparaphyllia X164 Branch leaves X30
Perichaetial teeth X164
The maps in Figures 12-14 show the distribution of the various species of Rhegmatodon and Macrohymenium on a world-wide basis. Only the locations of those specimens examined and verified by the author have been plotted. The base map is a modified Denoyer Semi-elliptical Projection.
Figures 15 and 16 contain the major published illustrations of these genera.
60




Figure 1: Pictures 1-18. Rhegmatodon spp. 1-3. Habit. 4-5. Operculae. 6. Perigonium. 7. Perichaetial Leaf Cells. 8. Archegonia and Paraphyses. 9. Antheridium and Paraphyses. 10. Capsule and Operculum. 11. Calyptra. 12-16. Perchaetial Leaves. 17. Perchaetial Leaf Tip with Tooth. 18. Exothecial Cells.




62 AIL ul
IT,
12 13 14 15 16 17 le




Figure 2: Pictures 1-22. Rhegmatodon spp. 1-2. Exostome Teeth on Capsule. 3. Exostome from Top. 4. Spores. 5. Endostome Teeth. 6-12. Exostome Teeth. 13-22. Pseudoparaphyllia.




64
5
4
6 7 10 11 12
13 14 15 16
17 18 19 20 21 22




Figure 3: Pictures 1-7. Rhegmatodon spp. 1-7. Leaf Cells.




cc
LO DUO CA




Figure 4: Pictures 1-13. Rhegmatodon declinatus. 1-2. Endostome Teeth. 3. Rough Seta. 4. Terminal Portion of Branch. 5-7. Leaf Tips. 8-12. Branch Leaves. 13. Exothecial Cells.




68
7
. ic 10 12 13 LAI




Figure 5: Pictures 1-23. Rhegmatodon polycarpus. 1-2. Endostome Teeth. 3. Smooth Seta. 4. Terminal Portion of Branch. 5-9. Leaf Tips. 10-22. Branch Leaves. 23. Exothecial Cells.




70
S3 4
2
5 6 7 8 9
10 11 12 13 14 1* 5 6
17 18 19 20 21 22 23




Figure 6: Pictures 1-14. Macrohymenium spp. 1-3. Habit. 4. Antheridia. 5-6. Alar Cells. 7. Perchaetium. 8. Perigonial Leaf. 9-10. Basal Leaf Cells. 11. Seta with Warty Projections. 12. Vaginule and Seta. 13-14. Basal Perichaetial Leaf Cells with Pores.




72
21
4
88
56
9 10
12 13 14




Figure 7: Pictures 1-15. Macrohymenium spp. 1-2. Endostome and Exostome Teeth. 3. Complete Endostome. 4. Spores. 5-8. Exostome Teeth. 9. Complete Exostome. 10-11. Calytrae. 12-14. Operculae. 15. Exothecial Cells.




74
OIL
As
Jt
10 12 13 14 15




Figure 8: Pictures 1-19. Macrohymenium rufum. 1. Portion of Branch. 2-3. Leaf Tips. 4-5. Endostome Teeth. 6-7. Cells of Midleaf. 8-10.
Perchaetial Leaves. 11-12. Cells of Leaf Base. 13-18. Branch Leaves. 19. Alar Cells.




76
V ] V 1A, op I
12 10
13 14 15 16 17 18 19




Figure 9: Pictures 1-14. Macrohymenium acidodon. 1. Portion of Branch. 2-3. Leaf Tips. 4-5. Endostome Teeth. 6. Perchaetial Leaf. 7. Cells of Midleaf. 8, 14. Alar Cells. 9. Cells of Leaf Base. 10-13. Branch Leaves.




78
10 11 12 13 14




Figure 10: Pictures 1-16. Macrohymenium muelleri. 1-3. Endostome Teeth. 4-5. Leaf Tips. 6-7. Cells of Midleaf. 8-9. Cells of Leaf Base. 10. Perchaetial Leaf. 11-14. Branch Leaves. 15-16. Alar Cells.




80
41
pi
10
12 13 14 15 16




Figure 11: Pictures 1-14. Macrohymenium strictum. 1. Portion of Branch. 2-3. Leaf Tips. 4-5. Endostome Teeth. 6. Perchaetial Leaf. 7. Cells of Midleaf. 8-12. Branch Leaves. 13-14. Alar Cells.




82
13 9 10 11 12 14




Figure 12: 1-2. Distribution of Rhegmatodon spp. 1. Distribution of Rhegnatodon polycarpus.
2. Distribution of Rheqmatodon declinatus.




VID
r O-f
77
Rhegmatodon potycarpus
Al
Rhegmatodon declinatus
2
00 X,




Figure 13: 1-2. Distribution of Macrohymenium spp. 1. Distribution of Macrohymenium rufum. 2. Distribution of Macrohymenium acidodon.




86 log
C340
Coll,
Macrohymenium rufum
Macrohymenium acidodon
2




Figure 14: 1-2. Distribution of Macrohymenium spp. 1. Distribution of Macrohymenium muelleri. 2. Distribution of Macrohymenium strictum.




88 CIO ID
0
Macrohymenium mueZZeri
CIO N M
C:;,w
Macrohymenium strictum
2




Figure 15: Pictures 1-9. Previous Illustrations of Rhegmatodon.
1. Anhymenium polycarpon Griff., Calcutta Jour. Nat. Hist. t. 16. 1843. 2. Anhymenium polysetum Griff., Icones Plant. Asiat. t. 97, f. 2. 1849.
3. Macrohymenium serrulatum Doz. & Molk. Musci Frond. Ined. Archip. Indici t. 56. 1848. 4. Rhegmatodon serrulatus (Doz. & Molk.) Bosch. & Lac., in A. Engler & K. Prantl, "Die naturlichen Pflanzenfamilien," Band 11, p. 298, f. 648. 1925. 5. Regmatodon declinatus (Hook.) Brid., Bryol. Univ. 2, t. 9. 6. Regmatodon declinatus (Hook.) Brid., Schwaegrichen, Species Muscorum Frondosorum, t. 204. 1827. 7. Pterogonium declinatum Hook., Trans. Linn. Soc. Lond. 9: t. 26, f. 3. 1808.
8. Enlargement of part of no. 7. 9. Original pencil drawings by Hooker found with the holotype (NY).




90
alp,




Figure 16: Pictures 1-6. Previous Illustrations of Macrohymenium.
1. Macrohymenium mulleri Doz. & Molk., in A. Engler & K. Prantl, "Die naturlichen Pflanzenfamilien," Band 11, p. 445, f. 745. 1925.
2. Macrohymenium mitratum (Doz. & Molk.) Flsch., in Fleischer, Die Musci der Flora von Buitenzorg, 4: 1365, f. 221. 1923. 3. Macrohymenium miller Doz. & Molk., Musc. fr. ined. Archip. ind. pp. 167-68, t. 55. 1848.
4. Macrohymenium rufum Doz. & Molk, Musci fr. ined. Archip. ind. pp. 16869, t. 54. 1848. 5. Leskea acidodon Mont., Ann. sc. nat. p. 96, t. 5, f. 4. 1845. 6. Original pencil drawings found with the holotype (L).




92
. ... .............................




APPENDIX I
HISTORICAL PERSPECTIVES IN TAXONOMIC BRYOLOGY
Modern classification is the end-product of a long series of decisions regarding the relative importance of certain characters as indicators of some "natural" relationship. Naturally, whatever biases or assumptions are used to determine these characters will be reflected in our final classification. This appendix is provided to supply some background with regard to the historical development of taxonomic bryology. The basic points to be covered are as follows
1. Bryology is a relatively new discipline, having begun in the modern sense with Hedwig only two centuries ago. This, and the fact that there are so few workers in the field, had slowed the growth of knowledge so that bryology is still in the exploratory and descriptive stages of taxonomy.
2. The current classification of mosses (Dixon) is the product of a long series of individual opinions and tradition with its roots deep in the folk science of various civilizations. A brief history of the ebb and flow of taxonomic opinion in bryology, especially with reference to the concept of the genus, suggests that each taxon is an abstraction in which we have assembled certain plants for the purpose of understanding and separated them from other groups.
3. Classification of mosses, like other plants, has been traditionally based on morphological or anatomical characters which are
93




Full Text

PAGE 1

A TAXONOMIC REVISION OF THE MOSS GENERA RHEGMATODON AND MACROHYMENIUM By DAVID ALVIN EAKIN A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 1976

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ACKNOWLEDGMENTS I would like to express my gratitude to Dr. Dana G. Griffin III for his guidance as Chairman of my Supervisory Committee. His example of professionalism in the field of bryological taxonomy, and botany in general, has been an inspiration to me. I would also like to express my love and appreciation to my wife, Darlene, for her patience, understanding, and assistance during the years of my graduate work, and especially during the period of this study. My thanks also go to Drs. James W. Kimbrough, Indra K. Vasil, Richard c. Smith, and Jonathan Reiskind for their careful reading of the dissertation, and helpful comments. Specimens in the following herbaria were studied and I express my appreciation to the curators. The abbreviations are those recom mended by the International Bureau of Plant Taxonomy and Nomenclature in Index Herbariorum, Part 1, 1974. NICH, FH, MO, MICH, NY, MPU, Z, HBG, B, LD, PE, 0, S, L, GL, MANCH, BM, C, HIRO, H, BR, TENN, PC, FI, FLA, and JE. In several instances H-BROTH was used to indicate that the specimen was from the Brotherus collection housed at Helsinki. Of the many friends and relatives who have helped and encouraged me, I would especially like to thank my father, Mr. John F. Eakin; my wife's parents, Mr. and Mrs. James E. Walker; and Mike and Diane Mccarry. Special thanks go to Ann Ellis and Robert Pentecost for their assistance with my photographic plates, and to Norma Donovan and Phyllis Young for typing the manuscript. ii

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TABLE OF CONTENTS ACKNOWLEDGMENTS ................................................ TABLE OF CONTENTS .............................................. LIST OF TABLES ................................................. LIST OF FIGURES ................................... ............. ABSTRACT ....................................................... S TATEMENT OF PROBLEM ........................................... HISTORY OF RHEGMATODON AND MACROHYMENIUM ....................... FAMILIAL AFFINITIES OF RHEGMATODON AND MACROHYMENIUM ........... INFRAGENERIC RELATIONSHIPS IN RHEGMATODON ...................... DESCRIPTION OF THE GENUS RHEGMATODON ........................... KEY TO THE SPECIES OF RHEGMATODON ............................. DISTRIBUTION AND HABITAT OF THE GENUS RHEGMATODON .............. DISPOSITION OF NAMES OTHER THAN SYNONYMS IN RHEGMATOIXJN .... .... INFRAGENERIC RELATION S HIPS IN MACROHYMENIUM .................... DE S CRIPTION OF THE GENUS MACROHYMENIUM ................... ...... KEY TO THE SPECIES OF MACROHYMENIUM ........................... DISTRIBUTION AND HABITAT OF THE GENUS MACROHYMENIUM ............ DISPOSITION OF NAMES OTHER THAN SYNONYMS IN MACROHYMENIUM ...... LEGENDS FOR THE FIGURES ........................................ APPENDIX I HISTORICAL PERSPECTIVES IN TAXONOMIC BRYOLOGY ..... iii Page ii iii V vi vii 1 4 14 16 20 23 36 37 39 42 46 57 58 60 93

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Page APPENDIX II PREVIOUS KEYS TO THE SPECIES OF RHEGMATODON AND MACROHYMENIUM BY BROTHERUS AND KIAER .................. 110 LITERATURE CITED .......................................... ..... 116 BIOGRAPHICAL SKETCH ............................................. 122 iv

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LIST OF TABLES Table 1. A chronology of important dates and publications in the history of the genera Rhegmatodon and Macrohymenium including significant dates in the Page history of taxonomic bryology ..................... . 11 Table 2. Data on three traditional leaf characters formerly used to differentiate smooth-setaed Rhegmatodons ......... 35 V

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Figure 1. Figure 2. Figure 3. Figure 4. Figure 5. Figure 6. Figure 7. Figure 8. Figure 9. Figure 10. Figure 11. Figure 12. Figure 13. Figure 14. LIST OF FIGURES Rhegmatodon spp. Rhegmatodon spp. Rhegmatodon spp. Page 62 64 66 Rhegmatodon declinatus................................. 68 Rhegmatodon polycarpus................................. 70 Macrohymenium spp. Macrohymenium spp. 72 74 Macrohymenium rufum.................................... 76 Macrohymenium acidodon................................. 78 Macrohymenium mi.ielleri ... .............................. 80 Macrohymenium strictum ................................. 82 Distribution of Rhegmatodon spp. ...................... 84 Distribution of Macrohymenium spp. ................... 86 Distribution of Macrohymenium spp. ............... ... 88 Figure 15. Previous Illustrations of Rhegmatodon .................. 90 Figure 16. Previous Illustrations of Macrohymenium ................ 92 vi

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Abstract of Dissertation Pr e sented to the Graduate Council of the University of Florida in Partial Fulfillment of the Requirem e nts for the Degree of Doctor of Philosophy A TAXONOMIC REVISION OF THE MOSS GENERA RHEGMATODON AND MACROHYMENIUM By David Alvin Eakin August, 1976 Chairman: Dana G. Griffin, III Major Department: Botany The moss genera Rhegmatodon (Leskeaceae) and Macrohymenium (Sematophyllaceae) are revised on a worldwide basis. Two species, Rhegmatodon declinatus (Hook.) Bridel emend. Eakin and Rh. polycarpus (Griff.) Mitten emend. Eakin, are recognized for the genus Rhegmatodon. Four species, Macrohymenium acidodon (Mont.) Doz. and Molk. emend. Eakin, M. m~lleri Doz. and Molk., M. rufum (Reinw. and Hornsch.) C. Muell. emend. Eakin, and M. strictum Bosch. and Lac. emend. Eakin, are recognized for the genus Macrohymenium. Keys to the species are included. Presented with each taxon is a complete list of synonomy, a taxonomic description and discussions on morphology and distribution. Standard notation of herbarium disposition is supplied for all specimens examined, and distributions of the taxa are based only on these specimens. Where available, information is given on the ecology of each taxon. vii

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STATEMENT OF PROBLEM The purpose of this investigation is to present a revision 1 of the moss genera Rhegmadodon and Macrohymenium. In the field of bryology today, there is a critical need for taxonomic revisions of many of the currently recognized genera. The relatively short history of modern bryology (less than 200 years) has produced more than a twentyfold multiplication of genera. However, only a very few have been subjected to the scrutiny of a major revision and, of these few, most were revised before the 1930's. In spite of the many fine investigations since the turn of the century, bryological taxonomy remains in the 'alpha' state. Revisional bryology is well characterized by the idea, "the fields are white unto harvest, but the laborers are few." The need for revision appears even more critical when we realize that many of the current genera were established during a period when the philosophical concept of genus in Musci was undergoing tremendous changes in perspective. Rhegmatodon and Macrohymenium were typical of the many genera needing revision, but at the same time offered features that made them particularly attractive for dissertation research. Both included a "manageable" number of species and there existed a good possibility of procuring nomenclatorial type specimens, so indispensable to revisional taxonomy. The decision to consider two genera, which are See discussion about the definitions of revision and monograph in Appendix I (page 93). 1

PAGE 9

2 currently assigned to two different families, was based on tl).e fact that their histories are greatly intertwined. As late as 1898, Paris (Index Bryologicus) included all previously described species of the two genera under the name Rhegmatodon. This was fascinating to me, since even a beginning student of bryology would place these genera in two different groups on the basis of current generic and familial concepts. The objectives of this study were 1. to collect, translate, and reevaluate all of the previous taxonomic literature with reference to these genera. 2. to provide a history of these genera with special reference to the changing concept of the genus within Musci as a whole. 3. to collect and critically reexamine all available type material on which previous taxonomic concepts were based. 4. to provide a revised taxonomic treatment of these genera based on type material and the greatest possible number of other collections. This treatment includes complete descriptions of the genera and keys to their component species. The types are designated for all species, and their location noted. Collection data and herbaria citations have been given for all specimens examined. Discussions of previous taxonomic treatments, nomenclatorial changes, and synonomy are included. Distributions are presented for each species as recog nized by the author. To date, no cytological, genetic, or ecological studies have been completed on these rare genera. None were attempted by this author due to the lack of fresh, living material.

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3 There are two areas of bac k ground necessary for a prope r under standing of the results and conclusions of this dissertation. The first involves an understanding of those basic taxonomic concepts re vealing the underlying philosophy and assumptions of modern taxonomic research in bryology. It is important to specifically relate these ideas to the modern generic concept in Musci. Secondly, one must recognize that the thread of history relating Rhegmatodon and Nacro hymenium is most meaningfully seen in the context of the overall historical fabric of taxonomic bryology.

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HISTORY OF RHEGMATODON AND MACROHYMENIUM The original specimen of Rhegmatodon was described as part of a collection of Nepalese mosses sent by Dr. Francis Buchanan Hamilton 1 to William J. Hooker. Hooker (1808) based his "Musci Nepalenses" on this collection. In error, he classified his specimen on the basis of a single row of peristome teeth; designating it as Pte rogonium declinatum sp. n., a Pterogonium caule repente, foliis imbricatus ovatus integerrimis mediotenus uninervibus, capsule declinata, operculo incurvato (Hooker, 1808, p. 309). Had he seen the exostome, and following th e generic concepts of his day, he would either have designated it a new species of Hypnum, or perhaps erected a new genus. He was clearly aware of its Hypnaceous appearance. I have named this plant from its most obvious character, the drooping capsule, in which it diff ers from all the other species of this genus described by Hedwig or Bridel, and has, in consequence of it, so completely the appearance of a Hypnum, that no one would suspect its real family, without first examining it (Hooker, 1808, p. 310). It seems strange that Hooker should have missed the exostome. According to Margadant (1968), Hooker was elected a Fellow of the Linnaean Society of London (1806) at 21 years of age on the basis of his discovery and H. C. Gangulee in his Mosses of Easte~_-ri India and Adjacent Regions, gives some interesting historical anecdotes about those who have col lected in eastern India. The collections of 1802-03, mostly from the Kathmandu valley of Nepal, were the first on the Indian subcontinent and earned for Hamilton the distinction of being the pioneer collector of bryophytes for this area. 4

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5 description of Bauxbaumia aphylla in the British Isles. This moss also has a double row of peristome teeth which are not nearly as distinct as those of Rhegmatodon. In addition, W. J. Hooker was an accomplished illustrator and engraver. Nor was it a case of having seen poor material for the original specimen has both endostome and exostome. My only explanation is that he first viewed the specimen dry, in which case, the external teeth would not have been seen at low magnification. In Rhegmatodon the exostome teeth alternate with those of endostome and, when dry, are inflexed between them. This possibility is seen by comparing Hooker's illustration (Fig. 15:7-9) with a low magnification photograph of a dry capsule (cf. Fig. 1:2). We should recall that microscopic characters were not commonly used at this time and microscopes were generally poor. Nevertheless, he did draw the endostome at a higher magnification and it is still puzzling to me how he missed picking off a few exostome teeth. With reference to the endostome he noted: There will also be found under the microscope a strong peculiarity in the circumstance of the teeth of the peri stomium being cleft in a very curious manner (Hooker, 1808, p. 310). This peculiarity is in fact a central point:in circumscribing the genus Rhegmatodon. His description is as follows: Peristomium dentibus sedecim, suberectis, e capsulae membrana interiore reticulata ortis, lineari-subulatis, luteis, transverse striatis, a basi ad medium longitud inaliter fissis (Hooker, 1808, p. 310). S. E. Bridel, in Bryologia Universa (1827), established the new genus Regmatodon 1 based primarily on this peculiarity of the peristome. The Note that the original spelling of the name is Regmatodon and not Rhegmatodon. These two spellings are orthographic variants. However, while some weight should be given to the spelling of the original

PAGE 13

6 name itself reflects the character, being formed from the two Greek words pnya fissure, and OOOVs of the teeth. F. c. Kiaer (1882, p. 41) notes that Bridel "who only knew the plant from the Hooker description, also designated the peristome as simple." Thus, while Bridel did segregate the specimen as a new genus, he perpetuated Hooker's error. The genus Macrohymenium with one species, M. rufum, was estab lished by Carl Muller in 1847, based on three specimens previously described as Leskea? rufa Reinwardt and Hornschuch (1826), Leskia mitrata Dozy and Molkenboer (1844), and Leskea acidodon Montagne (1845). Muller was apparently strongly influenced in this decision by various characters of the leaves; as indicated by the fir3t of his "essential characters." The leaves ecostate formed from narrow prosenchymatous cells (ellipsoidal areolation), strongly concave, subplicate, at the base to both sides provided with several large yellow lateral cells, densely overlapping, subsecund. With the capsule mouth constricted. The peristome double; the outer: the 16 teeth from the oblong base provided with striae in the middle, traveculate, suddenly long cuspidate, bent inwards; the interior: the teeth just as many, much overlapping the very longest outer teeth, widest, cuspidate, membrane lacking, put together from quadrate large yellow cells, placed on the reddish membrane, scarcely canaliculate, tessellated, not perforated (Muller, 1847, col. 825). Just three years later, however, when treating these same specimens in Synopsis Muscorum Frondosorum, he included his new genus under the concept of Rhegmatodon. He dealt with only three species; Rh. declinatus (Hook.) Brid., Rh. orthostegius Mont., and Rh. rufus C. Muller. The former two are Rhegmatodon in the modern sense, but rufus is a Macro hymenium. It may first be puzzling that Muller would dissolve a author, I feel that Bridel's spelling is in error. The Greek word should have a rough breathing symbol over the letter rho indicating the presence of 'h' when the word is transliterated to English. The correct spelling is therefore Rhegmatodon (cf. Stearn, William, 1967, Botanical Latin, p. 263).

PAGE 14

7 concept which just three years earlier had been so clear in his mind. Had he overlooked some important characters, or perhaps new and better specimens had come to light? Neither was the case. In fact, in expressing his concept of Rhegmatodon in this latter work, he did not even consider gametophytic characters. Note the following description from Synopsis Muscorum Frondosorum: The calyptra dimidiate. The peristome double. The external teeth very short, trabeculate, hypnaceous; the internal upon a short, scarcely keeled, tessellated, as if perforated membrane, turning reddish, very long, for a long time rising above the external, scarcely keeled, with the longitudinal line gaping to undivided, furrowed, remotely articulated (Muller, 1850, p. 29). There is no mention of the once predominant leaf characters in this latter description. The only gametophytic character used in this later classification is the nature of the costa. This actually represents a shift in Muller's views regarding the relative importance of sporophytic versus gametophytic characters in the delimitation of genera. This example well illustrates the problem facing bryologists as a result of the dual nature of mosses. The difficulty of systematizing a double organism, i.e., one representing two distinct generations of the life cycle is discussed in Appendix I. This problem is in fact central in the history of the changes which have taken place in the concept of genus in mosses. This discipline wide vacillation with regard to the selection of essential characters is reflected throughout the history of Rhegmatodon and Macrohymenium. Muller was obviously struggling with the problem in his treatment of the two genera. Hedwigian tradition emphasized the importance of the sporophyte. Muller at first breaks with this tradition by emphasizing various gametophytic characters and establishing the new genus

PAGE 15

8 Macrohymenium. He then conservatively shifts his emphasis back to sporophytic characters which, in the case of these taxa, will unite them under the same generic concept. This is a characteristic trend in the classification of the two genera. Whenever the gametophytic leaf characters were emphasized, Macrohymenium and Rhegmatodon were separated. Emphasis on the sporophyte, especially the peristome characters, united the specimens. Dozy and Molkenboer (1848) in Musci frondosi inediti archepelagi indici ... attempted to refine the genus concept for Macrohymenium on the basis of two new species, M. mulleri and M. serrulatus. Again we find the major emphasis given to the shape and structure of the peristome; especially to the ratio of length between the endostome and exostome teeth. Some insight into their view of the limited importance of gametophytic characters is shown by the following: Quadrate areoles to both sides at the wing of the leaves are not uniformly evident in all species, and therefore, they are seen to us not to be of great value, at least in explanation of the essential characters (Dozy and Molkenboer, 1848, p. 166). Emphasis on the similarities of the peristome caused them to ignore the obvious lack of inflated alars in s errulatwa. This further illustrates the principle stated above. However, in this case, Dozy and Molkenboer were dealing with two species of Macrohymenium and only one of Rhegmatodon; thus including both genera under the name Macrohymenium. In 1864, van der Bosch and van der Sande Lacoste continued Dozy and Molkenboer's work in Bryologia Javanica. They added one new species, M. strictum, and reclassified M. serrulatum as Rh e gmatodon serrulatus. While they did not specify the reason for this move, a look at their

PAGE 16

9 description of Rh. serrulatum implies greater stress on gametophytic differences. A major revision of the two genera was undertaken by F. Kiaer in 1882, which essentially established the modern distinctions between Rhegmatodon and Macrohymenium. There has been very little change in the circumscription of these taxa since Kiaer. Later taxonomic changes revolved around the question of closeness of relationship between the two genera. Kiaer used mostly gametophytic characters to distinguish the genera but placed them in the same subfamily, Macrohymenieae, on the basis of their proportionately long endostome. He lists: Rh. schotheimioides, Rh. filiformis, Rh. crizabanus, Rh. orthostegius, Rh. polycarpus, Rh. brasiliensis, Rh. secundus, Rh. declinatus, and Rh. serrulatus. Under Macrohymenium he includes: M. rufum, M. strictum, M. acidodon, M. nietneri, M. laeve, M. mulleri, and M. gracillimum. All of these species distinguish th e ms e lv es (as a group-DAE) by means of the great inner peristomc with the processes which are 2-4 times longer than the t ee th (Kiaer, 1882, p. 10). With the publication of Philibert's Etudes sur le peristome (1884-90) the peristome again was emphasized as the essential character in taxonomic bryology. The impact of Philibert on the classification of these genera is most clearly reflected by the treatment of these species by Paris (1898) in Index Bryologicus. All species of Macrohymenium were synonomized under the name Rhegmatodon. Twenty-one species are listed, apparently on the basis of this similarity in peristome structure. Gametophytic distinctions must necessarily have been entirely abandoned as generic criteria. When Brotherus (1909), in Engler and Prantl's Die Naturlichen Pflanzenfamilien (1 st edition), established the family Sematophyllaceae

PAGE 17

10 on the basis of Mitten's (1869) Tribus 16, Sematophylleae, the stage was set for separating Rhegmatodon and Macrohymenium. Brotherus' family concept was later emended by Fleischer (1923), Die Musci der Flora van Buitenzorg, into four subfamilies: Clastobryoideae, Heterophylloideae, Sematophylloideae, and Macrohymenioideae. Brotherus (1925) followed this treatment in the second edition of Engler and Prantl. Moving away from the overwhelming emphasis on the sporophyte, the essential characteristics of the family Sematophyllaceae (sensu Brotherus and Fleischer) are 1. costa absent, or short and bifurcated 2. alar cells enlarged 3. operculum long-rostrate. Both of these authors gave little regard to the peristome and thus the sporophytic characteristics within the Sematophyllaceae are widely variable. This was the last Qajor treatment of these taxa. Currently Macrohymenium is found as a subfamily of the Sematophyllaceae, and Rhegmatodon holds a similar position in the Leskeaceae. Subsequently, five new species have been described and I have found a great number of names ex herbario, which apparently have not been validly published.

PAGE 18

1741 1753 1781 1801 1808 1819 1826 Table 1. A Chronology of important dates and publications in the history of the genera Rhegmatodon and Macrohymenium including significant dates in the history of taxonomic bryology. Dillenius Historia Muscorum Linnaeus Species Plantarum Hedwig Fundamentum Hist. natural Hedwig Species Muscorum Hooker, W. J. First specimen of Rhegmatodon; described as Pterogonium declinatum Bridel, Mantissa Muscorum Bridel, Bryologia Universa the establishment of 11 the new genus Rhegmatodon based on Hooker's Pterogonium declinatum 1826-27 Leskea? rufa Reinwardt & Hornschuch 1836 1838 1839 1842 1844 1844 1845 1847 1848 Bryologia Europaea begun Anhymenium polycarpon Griff. Leskea parvula Hampe Rh. orthostegius Mont. Rh. parvulus Hamp. Leskia mitrata Dz. & Molk.; in Muscorum Frondosorum Novae Species ex Archipelago Indico et Japanio Leskea acidodon Mont. Macrohymenium rufum Muller; the establishment of the new genus Macrohymenium M. mulleri Dz. & Molk; M. serrulatum Dz. & Molk; in Musci frondosi inediti Archipelagi indici sive descriptio et adumbratio Muscorum Frondosorum in insulis Java, Borneo, Sumatra, Celebes, Amboina.

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12 1849 Anhymenium polysetum Griff. 1848-51 (50) Muller, Synopsis Muscorum Frondosorum M. rufum changed to Rh. rufum 1854 Bryologia Europaea, Bruch, Schimper, & Guembel; completed 1856 Clasmatodon parvulus Hook. & Wils. 1859 Mitt e n's Musci Indiae Orient 1862 Rh. brasiliensis Muller 1855-70 (64) Bryologia Javanica, M. strictum Dz. & Molk. 1868 Rh. kunzii Muller in schedula 1869 Mitten's Musci Austro-Americana; Rh. schlotheimioides Spruce; Tribus Sematophylleae established Hypnum minutum Mitten Rh. nietneri Muller 1871 Besch e r e lle, Prodromus Bryologiae Mexicana e Rh. filiformis Besch. Rh. fusco-luteus Schimp Rh. hypnoides Besch. 1872 Rh. chryseus Schimp. 1872 (73?) M. laeve Thw. & Mitten 1876-77 Adumbratio florae muscorum, Jaeger & Sauerbeck First reference to Rh. orizabanus 1880 M. acidodon (part of rufum mov e d by Be s cherell e ) Florule Bryologiqu e d e la Reunion M. acidodon var. acutissima B e sch. 1881 Rh. filiformis forma major (collection date of in schedula specimen of Bruch) 1882 1884-90 1891 Rh. madagassus Geheeb Kiaer, revision of Rhegmatodon and Macrohymenium Rh. secundus Kiaer Rh. densus Schirnp. in schedula M. gracillimum Mull. in litt. Rh. orizabensis Besch. in schedula M. cuspidatum Mitt. sedis incertis Philibert, H. Etudes sur le peristome Rh. feanus Muller

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1895 1897 1898 1904-05 1905 1907 1909 1909 1910 1910-11? 1910 1910 i923 1924-25 1924 1929 1932 1947 1955 Rh. palustris BrJth. in schedula Rh. palustris var. minor in schedula Rh. schlotheimioides var. minor Rh. newtonii Broth. 13 Paris, Index Bryoloicus (all species of Rhegmatodon and Macrohymenium under Rhegmatodon) Rh. bornmulleri Broth. in schedula Rh. fissidens Theriot in schedula Rh. filirameus Broth. in schedula Rh. sinense Theriot Engler & Prantl 1 st edition made the family Sematophyllaceae from hitten's Tribe Sernatophylleae Rh. pringlei Cardot Rh. mulleri forma pubea Dixon in schedula Rh. crassirameus Cardot Rh. cameruniae Broth. (collection date) Fleish e r, Die Musci der Flora van Buitenzorg (4 vol.) Macrohymenioideae one of four subfamilies of Sematophyllaceae M. mitratum M. wichurae Broth. in schedula M. densirete Broth. Musci, Brotherus, in Engler and Prantl, Die naturlichen Pflanzenfamilien 2 nd edition Rh. handelii Broth. Rh. declinatus var. minor Broth. Rh. novo-guinense Reimers Manual of Bryology, Verdoorn. Contains Dixon's classification Rh. brevicuspes P. de la Varde and Leroy (collected 1936) Rh. schwabei Herzog

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FAMILIAL AFFINITIES OF RHEGMATODON AND MACROHYMENIUM There have been some comments in the literature suggesting that the genus Rhegmatodon be transferred from the Leskeaceae to its own monotypic family, the Rhegmatodontaceae (Crum, 1951, 1973). The apparent motivation for the suggestion is the unique peristome of Rhegmatodon. Brotherus, in the second edition of Engler and Prantl (1925), segregates Rhegmatodon in its own subfamily, the Rhegmatodon~ toideae, on the basis of the F erL;tom wei t unter der Mundung inseriert I das dussere veil kUrzer. Macrohymenium holds a parallel position with respect to the Sernatophyllaceae. Brotherus separates this genus into its own swfamily, the Macrohymenioideae, on the basis of Fortsatze 2-3 mal so lang als die Peristomzahne. In both cases, the remaining genera of these families have peristomes in which the endostome is either the same length as the exostome or perhaps a little shorter or absent. These two genera stand alone in their respective families by virtue of an endostome 2-3 times longer than the exostome. However, this peculiarity of the peristome is not a .3ufficiently strong character to warrant such a transfer. With reference to some of the more recent discussions of the Leskeaceae and related families (Watanabe, 1972; Noguchi, 1972; Crum, 1973), all of the Rhegmatodon generic characters fit well into the family concept, with the exception of the ratio of endostome to exostome length. A 14

PAGE 22

15 similar situation exists with re spe ct to the relationship of Macro hymenium to the Sematophyllaceae Although Seki (1968) did not speci fically deal with Macrohymenium in his revision of the Sematophyllaceae of Japan, this genus fits very well into his revised concept of the family; again with the exception of the endostome to exostome ratio. Both in the case of Rhegmatodon and Macrohymenium, transfer to a mono typic family would be unnecessary and place too great an emphasis on this one peculiarity of the peristome. It is recommended that Rhegma todon remain in the subfamily Rh e gmatodontoid e ae of the Leskeacea e and that Macrohymenium retain a similar position in th e subfamily Macro hymenioideae of the Sematophyllaceae. Much more revisional work should be done with reference to the genera in these families before new families are proposed.

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INFRAGENERIC RELATIONSHIPS IN RHEGMATODON As previously noted (cf. p. 9), Kiaer accurately circumscribed the genus Rhegmatodon with reference to both gametophytic and sporo phytic characters. Within Rhegmatodon he erected two sections, Laeviseta and Scabros e ta, which correspond to the two species of the present treatment. In the wake of Kiaer's revision, however, a proliferation of new species occurred on the basis of certain gameto phytic characters, i.e., leaf shapo, average leaf ratio and perichaetial leaves. In the present work each of these characters was tested, both in the type specimens and later collections. I attempted to use both gametophytic and sporophytic characters to delimit species and gave careful consideration to those characters used by previous workers. However, the traditional gametophytic characters were found either to be continuously variable or to represent ecotypic variants, and were thus set aside. The morphological structures of taxonomic importance in Rh e gmatodon are completely sporophytic. The only sure way to distinguish the two species is by examination of the endostome teeth, the seta and the walls of the exothecial cells These three characters will be found in one of two discrete combinations, each characterizing one of the two species of Rhegmatodon. Rh. declinatus can be recognized by the combination 1) endostome teeth lacking dotting (Figs. 2:5, 4:12), 2) seta clearly rough from top to bottom (Fig. 4:3), and 3) c e ll walls 16

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17 of the exothecium irregularly thickened, presenting a collenchymatous appearance (Fig. 4:13). Rh. polycarpus is characterized by 1) dense dotting of the endostome teeth (Fig. 5:1, 2), 2) a smooth seta (Fig. 5:3), and 3) the walls of the exothecial cells regularly thickened parallel to the central axis of the capsule, with the transverse walls appearing thinner (Fig. 5:23). The taxonomic treatment of Rhegmatodon by Brotherus (in Engler and Prantl, 1925) is based on F. C. Kiaer's revision of 1832, Genera muscorum Macrohym e nium e t Rhegmatodon revisa sp e cieque nova aucta t 1 exposu1. Within Kiaer's and Brotherus' Scabroseta, species were differentiated by serrations on the leaf tips. This character proved to be a continuum, being highly variable, even within a single specimen. From specimen to specimen an examination of leaves from the branch tips revealed a continuum from those completely entire, to a few somewhat serrated, to those distinctly serrate (Fig. 4:5-1). In all cases, the leaves at the base of the branch were entire. To date no specimens of Rh. polycarpus with serrate l e aves have been seen, though a slight tend e ncy toward serration was noted (Fig. 5:9). Caution should be exercised in using this difference alone to distinguish species, as leaf serrations have been shown to be ecotypically variable (Briggs, 1965). In section Laeviseta, Kiaer and Brotherus used leaf ratios (2~2.5:l vs. 1.8:1) and leaf shapes (oval vs. oblong-o v al) to distinguish species. Table 2 presents data taken primarily from type specimens for all the For the purpose of comparison, translations of both these keys are presented in Appendix I.

PAGE 25

species with smooth setae. This c learly shows a continuum of l e af ratios. The averages in Table 2 are based on an examination of a minimum of 20 individual leaves per specimen. No correlation exists between leaf length and ratio or ratio and shape. Figure 5:11-22 well illustrates the range of the continuous variables of size, shape, ratios, and secundity. The extent and degree of secundity varied even within the same specimen. For example, the three leaf tips shown in Fig. 5:7-9 are from the same specimen. Secundity cannot be used in Rhegmatodon as a discrete species marker. D e velopmental studies will probably show this character to be an ecotypic variable as has been shown for other mosses (Streeter, 1970). Similarly, the degree of filiformity in branches was found to be highly variable within single specimens, and was excluded as a marked species character. This character has also been shown to be an ecotypic variant of mosses in very moist microhabitats (Richards, 1967). Several pictures of the exostome teeth have been included which clearly show two external plates backed by a single set of highly trabeculate plates (Fig. 2:1-3, 6-12). This was considered necessary since the well-known illustration by Dozy and Molkenboer (Fig. 15:4) 18 is incorrect on this point. Their illustration represents only the interior set of trabeculate plates. The error has been widely circu lated in that Brotherus chose this plate as his illustration of Rhegmatodon in Die naturlichen Pflanzenfamilien (1925). In this same drawing, the relative height of the endostomal and exostomal membranes is misrepresented. In the genus Rhegmatodon, the point of attachment of the endostome and exostome to their respective membranes is the same, i.e., the point of attachment for both the endostome and exostome

PAGE 26

19 is an equal distance below the rim of the capsule. Dozy and Molkenboer drew this as if Rhegmatodon serrulatus was in fact a Macrohymenium, as they supposed. Both parts of the peristome are inserted below the rim of the capsule in Rhegmatodon. The presence of paraphyllia in Rhegmatodon was first noted by Fleischer (1923) and subsequently by Brotherus (1925). How~ver, no description accompanied these notations. These structures actua1ly fit the concept of pseudoparaphyllia as presented by Ireland (1971). A comparison of Ireland's illustrations of pseudoparaphyllia with the pictures in Fig. 2:13-22 will document the similarity of structure. Furthermore, they are relatively sparse in Rhegmatodon by virtue of being restricted to branch and reproductive primordia on the branches. One other character used by Kiaer and Brotherus was the presence of teeth on the margins of the inner perichaetial leaves (cf. Fig. 1:17). The consistency of this character was tested in several specimens by the examination of many perichaetia. The presence and number of teeth was not at all consistent, whether in the same specimen or in different specimens, supposedly of the same species. This character was also excluded.

PAGE 27

DESCRIPTION OF THE GENUS RHEGMATOIXJN Regmatodon (Rhegmatodon) Bridel, Bryol. Univ. 2:294. 1827. Pterogonium declinatum Hook., Trans. Linn. Soc. IX, t. 26, f. 3. 1808. TYPE: Dr. Francis Buchanan Hamilton, Nepal, 1802-03 (holotype, NY; isotypes, s, H, BM). Plants slender to fairly robust, forming loose to dense mats, mostly dull or somewhat shiny, rigid, dark green or brownish. Primary stems prostrate, creeping, yellow to brownish, irregularly subpinnately and densely branched, rooting by means of radicles on the underside. Stem leaves small, usually absent, short, ovate-acute, entire, alars inconspicuous, marginal cells quadrate, the rest elliptical. Branches ascending to upright, terete, straight to curved, rigid, varying from short, thick and clavate to long, slender and of uniform diameter. Pseudoparaphyllla present but sparce. Branches densely foliose, leaves appressed and densely imbricate when dry, erect-spreading when wet, 0.5-1.7 mm long. Leaves broadly ovate, oblong-ovate to ovate lanceolate, short acuminate, concave, completely entire to strongly serrate at the apex, margins weakly to strongly reflexed to base of short acumen. Costa single from a wide base, narrowing rapidly, dis appearing at or somewhat above midleaf. Laminal cells oblong, arranged in oblique and longitudinal series, crowded, becoming subquadrate toward the margins and base of the leaf. Indistinctly bordered by a single 20

PAGE 28

21 row of quadrate marginal cells with transverse walls at right angles to the margin. Alars undifferentiated. Goniautoicous. Inner per chaetial leaves erect, lanceolate, gradually acuminate, with either completely entire or crenately denticul~te margins (Fig. 1:12-17). Perichaetium inserted near the base of the branch, budlike, somewhat open at the apex, archegonia shortly stipitate, cylindric from an oval base, with numerous filiform paraphyses, a little longer than the archegonia (Fig. 1:8). Perigonium axillary, budlike, ovate, leaves somewhat round ovate, suddenly constricted into a short, wide, moderately obtuse acumen, concave, completely entire, ecostate, the cells loose, hexagonal-rhomboidal (Fig. 1:6, 9). Paraphyses filiform and numerous (Fig. 1:9). Seta either smooth or rough, 4.5-13.5 mm in length. Capsule erect to inclined, unsymmetric, straight to somewhat curved, slightly constricted below the mouth, especially when dry, oblong cylindric to long cylindric, 1.5-3.1 mm long, smooth, brown to reddish brown, sometimes substrumose at base, exannulate. Exothecial cells either collenchymatous or with regularly thickened longitudinal walls and thinner transverse walls. Operculum short, conic, acumen short, obtuse, 1/4-1/2 the length of the deoperculate capsule (cells regularly hexagonal). Peristome double, the sixteen teeth of the exo stome lanceolate, obtuse, inflexed in the dry state, composed of a double series of cross-striolate external plates, and a single series of inner plates, trabeculate on the inner face (Fig. 2:1-3, 6-12). The sixteen processes of the endostome, well-developed, 2X longer than the exostome teeth, translucent, with or without dotting, split on midline from base to middle with one to three perforations in se~nents immediately above, appearing faintly to strongly bordered (Fig. 2:5). No cilia.

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22 Spores yellow-green to olive, globose, papillose, 21-30 m. Calyptra cucullate, smooth, somewhat long e r than the operculate capsul e

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KEY TO THE SPECIES OF RHEGMATODON 1. Seta rough from top to bottom; endostome teeth translucent and not dotted; the walls of the exothecial cells irregularly thickened, appearing collenchymatous ..... Rh. declinatus 1. Seta smooth from top to bottom; endostome teeth translucent and densely dotted; the walls of the exothecial cells regularly thickened longitudinally, the transverse walls appearing thinner. . . . Rh. polycarpus 23

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24 1. RHEGMATODOM DECLINATUS (Hook.) Brid., B:ryol. Univ. 2:204~205. 1 8 27. (Figure 4) Pterogonium declinatum Hooker, Trans. Linn. Soc. Lond. 9:309-310, t. 26, f. 3. 1808. TYPE: Nepal, without precise locality, Francis Buchanan Hamilton, s.n. (holotype, NY; isotypes, S, H, BM) Macrohymenium serrulatum Doz. & Molk., Musci Frond. Ined. Archip. Indici 6:170, t. 56. 1848. TYPE: Java, Mt. Patoeha, Korthals, s.n. (holotype, L; isotypes, S, H, BM, O, L, PC). New synonomy. Rhegmatodon serrulatus (Doz. & Molk.) Bosch. & Lac., Bryol. Jav. 2:111-112. 1864. New synonomy. Rhegmatodon feanus c. Muell., Nuov. Giorn. Bot. Ital. 23:601. ]891. nom. nud. TYPE: Burma, Bhamo, Feanus 25 (holotype, FI). New synonomy. Rhegmatodon declinatus var. minor Broth., Symb. Sin. 4:94. 1929. TYPE: China, in Kwangtsaoba, S. W. Kweitschou, Handel Mazzetti 10395 (holotype, H-BROTH; isotype, S). New synonomy. Rhegmatodon schwabei Herz., J. Hatt. Bot. Lab. 14:66, f. 22. 1955. TYPE: Formosa, western Middletaiwan, "Tigerkopf," Schwabe, s .n. (holotype, JE) New synonomy. Branch leaves 0.6-1.7 mm long, 0.3-0.7 mm wide, the average length to width ratios ranging from 2.2-3.0. Leaf margins range from completely entire to coarsely serrate, from midleaf to tip. Seta from 4.5 to 10.5 mm in length, 187-260 m wide, conspicuously rough from top to bottom. Vaginule 1.0-1.9 mm long. Capsule 1.5-2.4 mm long. Exothecial cell

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25 walls with irregular thickenings giving a collenchymatous appearance. Endostome teeth translucent and lacking dotting. Endostome teeth are twice the length of the exostome teeth, ranging from 450-600 m, while the exostome range from 200-300 m. Habitat: Usually on bark of fallen logs or trees, also on moist shady rocks. Mostly 500-1700 meters in altitude (rarely to 3000 meters). Distribution: Formosa, China, Burma, India, Nepal, Borneo, Sri Lanka, Java, Thailand, and Malakka (Malay Peninsula), Sikkim, Assam (Fig. 6: 2) FORMOSA. Mt. Lu-Chang-Ta, Nao-liao to Kuei-shan, Wu-feng Hsiang, Hsinchu, on bark of log on roadside in hardwood forest, 1800 m, Wang 1562; upstream area of Luming Chi, Hung-yey Tsun, Yen-ping Hsiang, Taitung, on hardwood log lying on roadside in hardwood forest, 100 m, Wang 0903; upstream area of A-li-pu-tang Chi, Wang-mei Tsun Sin-yi Hsiang, Nantow, on large branch of broad-leaved tree in hardwood forest, 1200 m, Wang 1449 (NICH); Hsueh Shan Shan Mo, on fallen tree in hard wood forests just below Anma-shan, c. 2100 m, Iwatsuki & Sharp 3140; Hsueh Shan Shan Mo, 17 km, above Anma-shan at edge of logging area near Chunghsueh-shan in a wet, n-facing ravine on a decaying log, c. 2500 m, Iwatsuki & Sharp 732a (NICH, TENN). Western Middletaiwan, Tigerkopf, Schwabe, s.n. (JE). CHINA. Fukien Province, Buong Kang, Yenping, on mossy bark cover ing the whole base of a tree, 700 m, Chung B34 (NICH, FH, MICH, 0, PE, S); on a mushroom log, 1000 m, Chung Bll7 (NICH, FH, MICH, O, BM); on a mushroom log in a bamboo forest, 3000 m, Chung B112 (FH, MICH, PE); on mossy rock, 700 m, Chung B48b, B52 (FH); on branch of madiellus, Chung B90 (FH, MICH); on rock, Chung B65, B68, B67a (FH, MICH); Chung B74a

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26 (FH, MICH, 0); B90a (MICH); Fuki e n Province, Kushan near Foochow, on a shady rock, 500 m, Chung Bl73 (FH); on a moist shady rock, Chung B245a (FH, HBG, MICH). Province Kweitschou, austro-occid, ad viam Tschenning Kuang-tsaoba, Yunnan in silva ad vie. Djitschangring pr. opp. Muyusse, substr. truncis viv. arb. frondos, c. 1050 m, Hand e l-Mazz e tti 10395 (S, H-BROTH). Province Kweitshou orient, prope oppidum Liping in silva mixta Nandjing-schan, ad arbores, c. 750 m, Handel-Mazzetti 10981 (H). Province Kwangsi, Laoshan, Ling Yuin Hsine, Chea 1963 (FH). Yunnan, frontier of Burma, no collector cited, 1898 (H). BURMA. Yunnan frontier, Niebolitz, 1911 (L, JE, PC). Bhamo, 3000', Feanus 25 (FI). SIKKIM. Yoksam, 1700 m, U. Tokyo Bot. Exp. to E. Ind. 200244 (NICH, NY, L); 20046 (H); Himalaya, prope Kurseong, 1372 m, P. Decoly & Schaul 2356 (BM, H). ASSAM. Khasia, Myrung, Griffith 212, Herb. E. India Co. 515 (BM). INDIA. Dehra Dhoon, Simla 717 (Griffith) (NY, BM). NEPAL. Without precise locality, Buchanan, s.n. (NY, S, H, BM). Without precise locality, Wallich, s.n. (BM). THAILAND. Payap, limestone massive Doi (Mt.) Chiegdao, hill ever green forest, on fallen tree trunks, 98 55' E 19 25' N, Touw 9063 (L). EAST INDIES. Without precise locality, Buchanan 1316 (S, BM). BORNEO. Between Sosopodon and S. Kelinggen, foot of Mt. Kinabalu, on fallen trunk, 1350-1400 m, Iwatsuki 1355 (NICH). SRI LANKA. Central Province, Muller 248 (H, PC). JAVA. Mt. Patoeha, Korthals, s.n. (S, H, L) Mt. Gede, Gerker, s.n.; Mt. Patoeha, Korthals, s.n. (L). Lignes i Habitus mest den fra Java in ag pedicelli Lenghe, Monte Patoeha, Korthals (0). Buitenzorg

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27 an Baum, Schiffner 11016 (S). Without precise locality, Korthals, s.n. L, BM, S, PC). Without precise locality, no collector cited (II, s, BM, C). Natural Reserve, Tjibogo, rainforest, tree trunk in shade, 1480 m, Soekar, 1949 (L). Tjibodas, Pentjuran mas on bark, van der Wijk 1001 (L). Without precise locality, Junghuhn, s.n. (L). MALAKKA. Bergland der Sakai, Werner, 1913 (S, JE). Rhegmatodon declinatus is clearly distinguished from Rh. poly carpus by its rough seta, the absence of dotting on the endostome teeth, and the collenchymatous appearance of the exothecial cell wall thicken ings. It is unfortunate that no morphological characters of the gameto phyte can be used to separate sterile specimens of the genus Although no specimens of Rh. polycarpus having serrated leaf tips have been col lected, I would caution against using this character alone to identify a specimen as Rh. declinatus. Although branch leaf characters fall within a continuum, the branch leaves of Rh. declinatus are, on the average, longer and more narrow than those of Rh. polycarpus. Similarly, the range of length for both the capsule and the seta is somewhat shorter in Rh. declinatus. The holotype for Hi1. schwabei Herz. was sterile. As only Rh. declinatus has been collected in Formosa, Rh. schwabei was placed under it. While Rh. polycarpus is widely distributed in the New World, no collections of Rh. declinatus have yet been made from this part of the world. Rh. declinatus seems then to be restricted to the Old World tropics and subtropics.

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28 2. RHEGMATODON POLYCARPUS (Griff.) Mitt., J. Linn. Soc. Bot. Suppl. 1:127. 1859. TYPE: Khasia, Hooker and Thomson 775 (lectotype, BM). (Figure 5) Anhymenium polycarpon Griff., Griff. muscolog. itin. Assarnici, 1838 (Calcutta J. of Nat. Hist. 3:275, t. 16. 1843), Not. p. 471, Griff Icones plant. Asiat. 2: t. 97, f. 1. 1849. TYPE: Assam, Murnbree, Griffith, 1835 (not seen). Rhegmatodon orthostegius Mont., Ann. Sc. Nat. Bot. ser. 2, 17:248. 1842. TYPE: India, Nilghiris, Perrottet, s.n. (lectotype, 0; syntypes, BM, PC, NY, 0). New synonomy. Anhymenium polysetum Griff., Not. p. 472, 1849, Griff. Icones plant. Asiat. 2: t. 97, f. 2., 1849. nom. inval. TYPE: Assam, without precise locality, Griffith, s.n. (lectotype, NY). Rhegmatodon brasiliensis Lindb. ex C. Muller, Bot. Zeit. 20:374. 1862. TYPE: Brazil, prov. Sao Paulo, Santos, Lindberg, 1854 (holotype, S; syntypes, 0, BM, H). New synonomy. Rhegmatodon schlotheimioides Spruce ex Mitt., J. Linn. Soc. Bot. 12:566. 1869. TYPE: Ecuador, Andes Quitenses, near Antornbos of the river Pastasa, Spruce 1437 (holotype, NY MITT; isotypes, MPU, O, S, BM, H, BR, PC). New synonomy. Rhegmatodon filiformis Schimp. ex Besch., Mem. Soc. Sc. Nat. Cherbourg 16:231. 1872, et Besch. in Fourn., Mex. Pl. 1 (Crypt.): 43. 1872. TYPE: Mexico, Chinantla, Liebmann. 1841 (lectotype, C; syntypes, O, BM, H). New synonomy.

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29 Rhegmatodon orizabanus Hamp. in Jaeg., Ber. S. Gall. Naturw. Ges. 1877-78:477. 1880 (Ad. 2:741) nom. nud. TYPE: Mexico, Mt. Orizaba, no collector cited, s.n. (lectotype, NY; syn type, BM). Rhegmatodon filiformis f. major Bruch, in schedula, (holotype, BM). New synonomy. Rhegmatodon secundus Kiaer, Forh. Vid. Selsk. Christiania 1882 (24): 38, 2 f. 5-7, 3 f. 1-5. 1883. TYPE: Malagasy Rep., Mt. Ankaratra, Borgen, 1887 (holotype, O; isotypes, S, NY, L, BM, C, H, BR, PC). New synonomy. Rhegmatodon orizabensis Besch. in Kiaer, Forh, Vid. Selsk. Christiania 1882 (24): 37. 1883. nom. nud. in synon. Type not seen. Rhegmatodon densus Schimp ex Kiaer, Forh. Vid. Selsk. Christiania 1882 (24): 37. 1883. TYPE: Mexico, Orizaba, Mohr, 1857 (lectotype, BM). New synonomy. Rhegmatodon palustris Broth., Bih, K. Svensk. Vet. Ak. Handl. 21 Afd. 3(3): 64. 1895 nom. nud. in synon. TYPE: Brazil, prov. Minas Geraes, Caldas, Mosen 365 (holotype, H; isotypes, Z, FH, S, BM) New synonomy. Rhegmatodon schlotheimioides var. minor Broth., Bih. K. Svensk. Vet. Ak. Handl. 21 afd. 3(3): 64. 1895. TYPE: Brazil, without precise locality, cardot 38 (holotype, H). New synonomy. Rhegmatodon palustris var. minor Broth., in schedula. TYPE: Brazil, without precise locality, Binot, s.n. (holotype, BR). Rhegmatodon newtonii Broth., Bot. Jahrb. 24:281. 1897. TYPE:

PAGE 37

30 Fernando Po, Pie Clarence, Newton, 1894 (holotype, H-BROTH; isotype, S). New synonomy. Rhegmatodon filirameus Broth., in schedula, 1907. TYPE: Guatemala, Alta Verapaz, Coban, Turckheim 6744 (holotype, H-BROTH; isotypes, FH, NY, LD, S, BM, PC, FI). New synonomy. Rhegmatodon pringlei Card., Rev. Bryol. 37:58. 1910. TYPE: Mexico, Puebla, Tezuitlan, Pringle 15285 (holotype, H; isotypes, FH, NY, LD, O, S, L, MANCH, BM, C, PC). New synonomy. Rhegmatodon crassirameus Card., Rev. Bryol. 37:58. 1910. TYPE: Mexico, Morelos, Cuernavaca, Pringle 15283 (lectotype, H; cotype 15310, NY). New synonomy. Rhegmatodon cameruniae Broth., in schedula, 1910 (TYPE: Cameroon, Kamerunberg, Musalse, Bushwald, Hintz, 1910 (holotype, H-BROTH). New synonomy. Rhegmatodon handelii Broth., Sitzungsber. Ak. Wiss. Wien Math. Nat. Kl. Abt. 1, 133:578. 1924. TYPE: China, Yunnan, Handel-Mazzetti 260 (holotype, H-BROTH; isotype, S). New synonomy. Rhegmatodon brevicuspis P. Vard et Leroy, Bull. Jard. Bot Bruxelles 18:182. 16. 1947. ham. illegit. TYPE: Africa, Tschibinda, Leroy 227 (holotype, BR). New synonomy. Branch leaves 0.5-1.3 mm long, 0.2-0.6 mm wide, the average length to width ratios range from 1.85-2.83. Leaf margins are completely entire with no serrations, weakly to strongly reflexed to base of the acumen. Seta from 5.5-13.5 mm in length, smooth from top to bottom. Vaginule 1.0-1.9 mm long. Capsule 1.9-3.1 mm long. Exothecial cells

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31 with regularly thickened longitudinal walls; transverse walls appearing thinner. Endostome teeth translu c ent, densely punctate. Endostome teeth ranged from 470-620 m and exostome teeth ranged from 200-350 pm. Habitat: Usually on the bark of trees or fallen logs, occasionally on wet ground and rocks. Ranging from 1000-3048 min altitude. Distribution: Sikkim, Assam, India, China, Sri Lanka, Thailand, Malagasy Rep., Malawi, Guinea, Fernando Po, Cameroon, Mexico, Panama, Guatemala, Brazil, Bolivia, Burma (Fig. 6:1). SIKKIM. Without precise locality, 2-4000', no collector cited, s.n. (NY). Punkabari, 2-4000' Kurz 2453 (H, BM). Himalaya, 2-4000', Kurz 276/5 (0). Without precise locality, 5000', King Il4b (BM). With out precise locality 2-8000 1 King, 1894 (BR). ASSAM. Moosai, Griffith, s.n. (0, H). Khasia, regio sub trop., 4000', JD Hooker and Thomson 775 (BM). INDIA. Kumaon, NW Himalaya, Malkarzun, Askot e district, 1388 m, Khan, Broth 1940 (NY, S, H, BM); Rolam River, 3048 m, Khan, Broth 1935 (0, H). Nilghiris, Perrott e t 1644 (BM, NY); P e rrottet, s.n. (NY, BM, 0, PC); Montagne, s.n. (NY, BM, BR, L, 0); prope Ootacamund, Weir 276/21 (NY, BM); Ootacamund, 7000', Gambleu 17260 (H); Coonor, trees, 5000', Gambleu 13013 (H). Nilghirebirge bei Coonor in Simsparkjungle an Baumen, 1950 m, Fleischer 511 (B, H, L). Eastern Nilghiris, Kotagiri, tree, 6500', Sedwick 757 (BM). Nilghiri Hills, Srinivasan 277 (HIRO). Palni (Pulney or Palm) Hills, Kodaikanal, Foreau, 1927 (FH); in groves along the Kodai road about 29th mile, c. 5900', Foreau 477 (FH); Kodai road, Foreau 199 (BM); Togaivarai Shala, 4500', Foreau 262/26 (F, BM); Perumalmalai Shala, along torrent, 4500', Foreau 141/26 (BM); near Shembaganur, Pragasafuram, 6400', Foreau, 1957 (BR); 6700', Foreau, 1958 (TENN). Kodaikanal, on trees, 2400 m, Foreau and Raine 71 (MANCH,

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PC, FH, O, H); Madura, Foreau, 1 9 11 (BR); en Madure dans les Gathes, 7000'+, Andre 35 (PC); Andre, 1909 (BM). BURMA. Yunnan frontier, Niebolitz, 1911 (JE). 32 CHINA. Yunnan, Pe yen tsin, auf Baumrinde, Ten, 1921 (FH, JE, S); Ten 9 (S); Ten, 1924 (H). Prope urbem Yunnanfu, in regionis calide temperatae, truncis viv. Quercuum ad templ. Helungtang, c. 1950 m, Handel-Mazzetti 260 (H, JE, S). SRI LANKA. Without precise locality, Thwaites, s.n. (S); Thwaites 248 (S, BM, H). An Heinen in der Bathschlucht bei Hakgala, 1300 m, Herzog 139 (H, L); 138 (JE). THAILAND. Payap, granitic massive Doi (Mt.) Inthanon, on branches of Quercuus in clearing, 98 30' E, 18 35' N, 1750 m, Touw 9845 (L). Payap, limestone massive Doi (Mt.) Chiengdao, decid110us forest on NNE slope, on tree trunks, 98 55' E, 19 25' N, 1000 m, Touw 8903 (L). MALAGASY REP. Mt. Tsaratanana, 1200-2400 m, Bathie, 1924 (FH, S, JE, H, BR). Mt. Ankaratra, Borgen, 1877 (NY, O, BR, PC); Borgen, s.n. ( NY 0 L BM C H S ) MALAWI. Vernoy Nyasaland, Luchenya Plat e au, Mlange Mt., Mlange dist., on trunks of trees in dense forest shad e 1890 m, Brass 16537 (NY) GUINEA. In jugo Danguina, 1100 m, Rob e grin, s.n. (0, S, L, GL, H, BR, PC); 1800 m, Robegrin, 1843 (H). FERNANDO PO. Pie Clarence, 2000 m, Newton, 1894 (S, H). CAMEROON. Bamenda, Lake Bambaluwe, on tree trunks in montane forest, Richards R5250 (GL, L, BM). Victoria div., below Likonge, on stem in montane forest, 6500-7000', Richards R4264 (BM, L). Cameroon Mt., on trunks of fallen trees, 5000', Dunlap, 1926 (BM); on trunks of

PAGE 40

33 trees near timberline, 7000', Dunlap, s.n. (BM). MEXICO. Oaxaca, vor dem Pass oberh. Teotitlan in der Sierra, in epiphytenreichen Bergwald an Barke, 2100 m, Dull 92 (MICH). Puebla, Tezuitlan, on wet banks, 7000', Pringle 15285 (H, FH, NY, LD, o, s, L, MANCH, BM, C, PC); Tulancingo Road west of Huauchinango, bark of alder, 5000', Sharp 887 (TENN); on Carpinus, 5000', Sharp 913a, 911B (TENN). Chinantla, Liebmann, s.n. (0, BM, C, H). Michoacan, vicinity of Morelia, Cerro Azul, 2300 m, Arsene 4543 (FH, S). Morelos, near cuernavaca, Pringle 15283 (H), 15671 (S, MICH, PC), 15310 (NY). Chiapas, ditio San Cristobalr "Los Llanos" in silva quercuum, 2200 m, Munch 7464 (NY, FH, H, BM); prope San Cristobal, "Los Llanos," in praeruptis umbrosis, 2500 m, Munch 7462 (NY, BM, H); in montibus supra S. Cristobal, 2100 m, Munch 7282 (H, PC); Cerro Hueytepec, near Las Casas, 8000', Sharp 3283, bark of oak, Sharp 3283a, 3270; between Las Casas and San Gregorio, on oak, 7500', Sharp 4683, 4682a (TENN); SW Jalisco slopes of La Ferreria, above Manantlan, on rotted log, oak forest, 6500', Crum 1041 (MICH, S, TENN, NY). Guerrero, gorge below Ormiltemi, 30 km west of Chilpancingo, 6300', Sharp 1149a (TENN); Rancho del Cielo above Gomez Farias, Tamps., at the mine, fallen log, meso, partial shade, Sharp 3618 (TENN). Orizaba, no collector cited, 1857 (NY); No. 68 (BM), Borgeau, 1866 (PC). In planitic Orizabac in sylving ad 6500', Mohr, 1856 (0). Without precise locality, no collector cited, 1865, (NY); without precise locality, Liebmann, s.n. (BM); Totutle, ad hunc ono pertinet fruches perfecti et caespes flaveferis, Liebmann 8467 (C). PANAMA. Chiriqui, El Volcan, 6500', Llano, 1952 (HIRO); vicinity of El Boquete, 1000-1300 m, on tree trunk, Maxon 4985 (NY). GUATEMALA. Quetzaltenango, on oak, 7800', Sharp 2063 (MO, TENN);

PAGE 41

34 7700', Sharp 2086 (TENN). Chimaltenango, on oak above Tecpam, 8300', Sharp 2571 (TENN). Coban, Alta Verapaz, in arbore vetusta, 1310 m, Turckheim, Broth 6744 (NY, FH, LD, S, BM, H, PC, FI). BRAZIL. Caldis, Lindberg, 1854 (NY). Prov. Sao Paulo, rariss ad truncos arb. pu. silv. primae. propr opp. Santos, Lindberg, 1854 (O, S, BM, H). Prov. Minas Geraes, Serra de Caldas ad radices arb in suargine paludis, Mosen 365 (NY, S, H, Z, FH). BOLIVIA. Below Pelichuco, on rock, 3000 m, Williams 2851 (NY, H, BM). ECUAIXJR. Andes Qui tenses, Antombos, Pastasa River, 5000', Spruce 1441, 1437 (NY, MPU, 0, S, BM, H, BR, PC). Rhegmatodon polycarpus is readily distinguished by 1) highly dotted endostome teeth, 2) a smooth seta, and 3) exothecial cell walls regularly thickened longitudinally, with the transverse walls appearing thinner. The margins of the branch leaves are completely entire. The perichaetial leaf margins with or without teeth. The shape of the branch leaves is highly variable. The branches range from thick and clavate to filiform. This species has a pantropical and subtropical distribution.

PAGE 43

DISTRIBUTION AND HABITAT OF THE GENUS RHEGMATODON Rhegmatodon declinatus has only been collected from the tropics and subtropics of the Old World (Fig. 12:2). In contrast, Rh. poly carpus (Fig. 12:1) has a pantropical distributton ranging from Central and South America to Africa, Madagascar, Indi~ and China. Unlike Rl1. declinatus, Rh. polycarpus has not been collected in Indonesia and Malaysia. The ranges of these two species overlap in China (Yunnan), Sikkim (Himalaya), Assam (Khasia), Ceylon, and Thailand (Mt. Chiengdao). The areas of collection are characterized by altitudes of 650-1400 m. The disjunctive distribution of Rh. polycarpus is similar to that of other moss genera such as Pilotrichella, Squamidium, Lindigia, Braunia, and Dimerodontium. Herzog (1926, 1932) uses the continental disjunctions of certain mainly austral families of vascular plants, with similar pat terns of distribution in support of Wegener's (1924) theory of continental drift. The disjunctive distribution of Rh. polycarpus is very similar to that of Leptodontium viticulosoides and vars. (Zander, 1972). Rhegmatodon is most often found at the middle and high altitudes in the mountainous regions of the tropics and subtropics. Rh. declinatus grows at lower altitudes than Rh. polycarpus. Rh. declinatus has been found on the bark of fallen logs or trees, and occasionally on moist shady rocks, most often between 500-1700 m. Rh. polycarpus, usually found between the altitudes of 1400-2500 mis also corticolous, but may be collected on wet ground and rocks. 36

PAGE 44

DISPOSITION OF NAMES OTHER THAN SYNONYMS IN RHEGMATODON The following list includes the names of in schedula specimens, as w e ll as validly published specimens, where examination of the types showed them to belong to some other taxa than Rhegmatodon. Most of these specimens were collected in areas for which adequate keys are not available, and onl y a lirriit.ed attempt was made to assign these taxa to genera. For the purpose of completeness, previous exclusions and transfers are included in this list. Rhegmatodon bornmulleri Broth. in schedula, 1904. TYPE: Brazil, Cruz Alta, Colonia Nova Wurterriburg, Elsenau, in arboribus, 450 m., Bornmuller 6156 (holotype, JE: isotypes, B, S). This differed from Rhegmatodon by having no exostome and highly papillose, bifid endostome teeth. Most likely this is Dunerodontium mendozense Mitt. Rhegmatodon chryseus Schimp nom. nud. in synon. Roz e a chrys e a Besch. in Mem. Soc. Sc. Nat. Cher. 16:242. 1872. And Besch. in Fourn., Mex. Pl. 1 (Crypt.) :48. 1872. TYPE: Mexico, no collector cited, s.n. Rhegmatodon fissidens Ther., in schedula, 1905. TYPE: China, Prov. Kong Tiheau, Carabrie 3629 (holotype, H). Laminal cells all subquadrate. This is probably a Leskea sp. Rhegmatodon fusco-luteus Besch., Mem. Soc. Sc. Nat. Cherbourg 16:232. TYPE: Mexico, Orizaba, Muller, s.n., in herb. 37

PAGE 45

Schimper {not seen) = Platygyrium r.uscolut e um Card. cf Card, Rev. Bryol. 38:40. 1911. 38 Rhegma todon minutus (Mitt.) Par. Ind. Bryol. 110 9 1 89 8. TYPE: Andes, Jameson, s.n., (not seen) = Helicodontium minutum (Mitt.) Jaeg., Ber. S. Gall. Naturw. Ges. 1876-77:225. 1878 (Ad. 2: 291) Rhegmatodon nietneri C. Muell., Linnaea 36:20. 1869. TYPE: Ceylon, Nietner, s.n. (isotype, 0). The areolation of the lamina was not like Rhegmatodon. I was unable to id e ntify this to g e nus. Rhegmatodon parvulus Hamp. Icon. Muse. 2:14. 1 8 44. TYPE: Georgia, Eleenezar near Savannah River, no collector cited, s.n., (not s ee n) = Clasmatodon parvulus (Hamp) Sull. in Gray, Man. Bot. N.U. States ed. 2:660-5. 1856.

PAGE 46

INFRAGENERIC RELATIONSHIPS IN MACROHYMENIUM The taxonomic treatment of Macrohymenium by Brotherus (in Engler and Prantl, 1925) is based on F. C. Kiaer's revision of 1882, Genera muscorum Macrohymenium et Rhegmatodon revisa specieque nova aucta exposuit. 1 Kiaer based his key totally on gametophytic characters in order to include M. sinense. This specimen has subsequently been placed in the genus Giraldiella. Thus Brotherus also separated the various specimens of Macrohymenium on the basis of gametophytic characters. In this present study, many of these characters are excluded. The following were found to be variable, varying even within the same specimen: denticulations on the margins of the perichaetial leaves, loose to densely imbricated leaves, and leaves heteromallous to homomallous. Some gametophytic characters are useful in determining species of Macrohymenium, e.g., leaf size, shape, and areolation. N. rufum leaves are the smallest (.79-1.41 mm long) and widest (2.27) ovate and abruptedly constricted to a short acumen (Fig. 8:13-18). In M. acido don, branch leaves are small (.89-1.45 mm) (Fig. 9:10-13) and more narrow than M rufum (2.77). M. acidodon can be differentiated from the other species by virtue of the uniform size of the leaf cells (Fig. 9:2,3,7, 9). Each of the other species has leaf cells which are much longer at For the purpose of comparison, translations of both the keys from both works are presented in Appendix II. 39

PAGE 47

40 the base and gradually shorter toward the tip. M. mulleri is the largest of the species, the leav e s the longest (1.9-2.5 mm) and the most narrow (4.0) (Fig. 10:11-14). It also has more enlarged alar cells (12-20) (Fig. 10:19). The other species have from 6-9. M. strictum has slightly larger leaves (1.0-1.43 mm) than M. acidodon and can be distinguished by a sudden constriction to a long acumen (Fig. 11:2-3). Brotherus did include one sporophytic character in his key, separating his six species into two equal groups on the basis of the seta plainly rough versus the seta set above with very low, broad warts. While some setae may appear rough when dry (Fig. 6:1), moistened setae of all species are smooth at the base and towards the top have a few low, broad projections (Fig. 6:11-12). Neither Kiaer nor Brotherus used the presence or absence of perforations in the endostome teeth as key characters. Probably, this was due to Kiaer's statement that he had found both perforate and imperforate endostome teeth on th e same specimen. In every specimen examined during this study this character was discrete. The endostome teeth were either perforate or imperforate, and never differed within the same endostome. Of the four species recognized in the present work, two have perforated endostomes and two are imperforate. The two species which have perforations are also quite different in the expression of the character. M. mulleri has very regular perforations in nearly every segment, producing a scalloped effect. M. mulleri also has the shortest peristome, the endostome ranging from 250-300 m and the exo stome from 190-200 m (Fig. 10:1-3). The seta is moderately long (811 mm) and quite thin (130-150 m). On the other hand, M. acidodon h~s irregular perforations in the upper portion of the teeth, sometimes

PAGE 48

41 nearly continuous (Fig. 9:4-5). The peristome is moderate in size, the endostome ranging from 350-500 m and the exostome from 200-250 m. The seta is 9-11 mm long and 115-180 m wide. The endostomes of M. rufum (Fig. 8:4-5) and M. strictum (Fig. 11-4-5) are imperforate. The peristome of M. strictum is much larger (endostome, 550-750 m; exostome, 300-410 m) than that of M. rufum (endostome, 440-600 m; exostome, 170300 1Jm). The seta of M. strictum also is much longer (8-21 mm) and wider (200-374 lJm} than that of M. rufum (6-6.5 mm long and 160-220 lJm wide).

PAGE 49

DESCRIPTION OF THE GENUS MACROHYMENIUM Plants lustrous, yellow-brown to reddish green, in moderately com pact, low sods, copiously fruiting. Pleurocarpous mosses, prostrate and rooting. The sporophytes lateral at base of the branches. Goniautoicous. Primary stems thin, stolon-like, zigzag, creeping, lacking a central strand, copiously branched. Branches typically simple, ascending, short, thick and round, straight to curved, densely foliose; leaves imbricate, heteromallous to somewhat homomallous. Paraphyllia or pseudoparaphyllia lacking. Branch leaves ecostate, concave, ovate-lanceolate, acuminate, with the leaf margins entire, more or less recurved. Lamina] cells elongate, oval, walls incrassate osteoform with thin connecting areas between cells. Alar cells rec tangular and quadrate, colored and somewhat inflated. Leaves .792.53 mm long, and .29-.66 mm wide. Inner perichaetial leaves large ecostate or faintly hinting of a costa, upright, sheathing, ovate lanceolate, acuminate, or somewhat blunt. Perichaetium rooting. Vaginule small, oblong cylindric. Perigonium axillary, hidden betw~en the stem and branch leaves, budlike, very small, 1/3 length of leaves. Perigonial leaves broadly ovate, acuminate, ecostate, entire. Antheridia many, pedicellate, short stipitate, oblong-cylindric, with small para physes sparce or lacking. Perichaetium axillary, inserted in the axils of the branchlets, budlike, many-leaved, and obscured between the leaves. The archegonia short, moderately robust, with very few short, filiform 42

PAGE 50

43 paraphyses. Setae flexuous or twisted in the dry state, short to long, 6-21 mm, smooth at base, but with low hyaline protuberances below the capsule. Capsule, erect-inclined, constricted below the mouth, oblong, reddish brown. Exothecial cell walls irregularly thickened, appearing collenchymatous. Operculum conic rostrate, nearly equalling the capsule in length. Peristome double; exostome of 16, equidistant, erect-incurved teeth, inflexed in the dry state, broad at the base with two striated dorsal plates separated by a zigzag line, suddenly constricted to a long ciliate tip, the zigzag line continuing to the tip. Dorsal plates backed by a single row of trabeculate plates. Endostome of 16 teeth, from a low basal membrane, without cilia, the processes about 2X longer than the exostome teeth, keeled, entire, either perforate or imperforate on keeled midline. Membrane subplicate, tesselated. Spores globose and papillose, 15-27 min diameter. Calyptra cucullate. Margadant (1959) posed a nomenclatorial problem with reference to the genus Macrohymenium. Fleischer (1923) synonomized Leskea? rufa Reinwardt and Hornschuch with Acroporium braunii (C. Mull.) Fleisch. Because Leskea rufa was an earlier collection than the type for A. braunii the name was changed to Acroporium rufum (Hornsch. and Reinw.) Fleischer. Fleischer said he had examined the type for Leskea rufa in the Berlin Herbarium. Believing this specimen to be the basionym for Macrohymenium rufum, the type species of the genus Macrohymenium, Margadant felt that this transfer threatened the nomenclatural standing of Macrohymenium. He therefore proposed that Macrohymenium acidodon (Mont.) Doz. and Molk. (1848) be designated the lectotype and the name Macrohymenium be conserved. All of this is based on the assumption that Carl Muller used Leskea rufa Reinwardt and Hornschuch as the type speci men for his new genus. There is good reason to believe that this was

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44 not the case. The original specimen of L e skea rufa was described in Nov. Act. Caes. Leop. 14: Suppl. 2: pp. 716-17. 1826. It was collected by Reinwardt on Mt. Malabar (Malabaria), Java. In 1847, when erecting the new genus, Macrohymenium, Muller cited three specimens in synonomy. The first was Leskea rufa Reinwardt and Blume, Java. This same citation is found in Synopsis Muscorum Frondosorum 2:30, 1850, when Muller dis solved Macrohymenium and moved M. rufum into the genus Rhegmatodon as Rh. rufus. In neither of these citations does Muller ever mention Malabaria. This suggests that his type for M. rufum was in fact based on another specimen. There are several facts which support this idea. The original description of Leskea rufa Reinwardt and Hornschuch, states that the peristome was destroyed. Yet Muller's genus description contains a detailed description of both the endostome and exostome; neither of which could be mistaken for Acroporium. Further support issues from the synonomy of Dozy and Molkenboer (1848) in which they list two specimens from Java; Blume and Reinwardt, Java, and a separate Reinwardt, Mt. Malabar. Java: collegerunt Blume et Reinwardt, monente Mlillero: Malabaria: collegit Reinwardt (cf. Nov. Acta Acad. Caes. Leop. 1. cit.). (Dozy & Molk., 1848, p. 167.) Note that the first specimen was "collected by Blume and Reinwardt, with advice by Muller," while the Malabar listing only mentions Reinwardt as the collector, and by the accompanying citation gives clear indication that this second specimen is the Leskea rufa of 1826. Whenever the Mt. Malabar specimen is cited, Reinwardt's name is used alone, never in conjunction with Blume. There is sufficient evidence to support the e xist e nce of a separate specimen from Java, collected by Blume & Reinwardt, on which the type

PAGE 52

45 description of M. rufum is based. Fleischer, recognizing that the specimen from Mt. Malabar was actually an Acroporium, placed it in synonomy with Acroporium braunii without jeopardizing the nomenclatorial standing of Macrohymenium. One other possibility exists. Muller may have described his new genus using a combination of characters from the three specimens in his synonomy. This would assume the Mt. Malabar collection to be the same as the Blume and Reinwardt collection. If so, his description of the peristome could only be that of L. mitrata since no perforations of the endostome are noted. L. acidodon 1 has definite perforations, for which reason it was taken out of synonomy and designated the type for Macro hymenium acidodon (1848). I recently received a collection of Macro hymenium from Jena (JE) which contained a specimen with the following data: Rhegmatodon rufus mihi! Java: Blume Vereinigte Herbarien Karl Schliephaeke, Osterfeld und Hermann Winter, Gotha. This specimen is clearly a M. rufum and has two sporophytes in excellent condition. While there is no date, I believe this is the missing specimen of Blume and I am designating it as the lectotype for M. rufum. If futher evidence negates my argument, I recommend that Leskia mitrata Doz. and Molk. be designated the lectotype for Macrohymenium rufum. For reasons stated above, Leskia mitrata Doz. and Molk., Musci Frond. ex Archip. Indici. 1844, p. 15; Ann. des Sc. Nat. 1844, p. 511, is the 2 earliest described specimen of Macrohymenium rufum C. Muller. 1 Assuming Muller saw the 1845 specimen of Doz. and Molk. 2 Granting the above argument, this would be the proper citation for M. rufum.

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KEY TO THE SPECIES OF MACROHYMENIUM 1. Endostome teeth with perforations in the keeled midline, at least from mid-tooth to tip .... 2 1. Endostome teeth with no perforations of the keeled midline 3 2. Perforations even, uniform, giving a scalloped ap pearance from the base to the tip; large branch leaves (1.9-2.5 mm long) with long basal cells (12:1), the cells gradually shortening to the tip (3:1) M. m~lleri 2. Perforations uneven, irregular, restricted mostly to the upper half of the teeth, sometimes appearing bifid at the tip; smaller branch leaves (0.9-1.5 mm long), the cells of uniform size from the base to the tip of the leaf (5:1 3:1) ... M. acidodon 3. Branch leaves lanceolate, gradually acuminate; peri chaetial leaves abruptly constricted to acuminate tip; seta ranging from 8-21 mm in length ... M. strictum 3. Branch leaves from a squarish base, widely ovate-lanceolate, abruptly acuminate; some of the perichaetial leaves blunt; seta ranging from 6-7 mm in length ... M. rufum 46

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1. MACROHYMENIUM RUFUM C. Muell., Bot. Zeit. 5:825. 1847. TYPE: Java, without precise locality, Blume and Reinwardt, s.n. (lec totype, JE) 47 (Figure 8) Leskea rufa Reinw. and Hornsch., Nov. Act. Acad. Caes. Leap. Carol. 14, p. 2 suppl. 1826, p. 716-717. TYPE: Java, Mt. Malabar, Reinwardt s.n. (not seen). Leskea mitrata Doz. & Molk., Ann. Sc. Nat. 3 ser. 2. p. 311. 1844. TYPE: Sumatra, Korthals, s.n. (holotype, L). New synonomy. Rhegmatodon rufus (Reinw. and Hornsch.) C. Muell., Syn. 2:30. 1850. TYPE: Java, Blume and P.einwardt, s.n. Macrohym e nium curvirostrum, Braun, in schedula. TYPE: Java, Braun, s.n. (holotype, BM). New synonomy. Plants small, the leaves .79-1.41 mm long and .31-62 mm wide, with an average length to width ratio of 2.27. The leaves ovate and abruptedly constricted into the acumen. The leaf c e lls longer at the base of the leaf (10-12:1), gradually shortening towards the tip (2-3:1). Six to nine inflated quadrate colored alar cells. Usually, one or two of the inner perichaetial leaves with a very blunt tip. Seta from 6-6.5 mm in length and 160-200 min width. The endostome teeth lacking perfora tions on the midline. The endostome teeth ranging from 440-600 m, and the exostome teeth ranging from 170-300 m. Habitat: On tree trunks in forests, epiphytic, usually found be tween 1500-2000 m, but occasionally up to 3030 m. Distribution: Java, Sumatra, Borneo, New Guin e a, Australia, Sri Lanka, Malagasy Rep.

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48 JAVA. Tjibodas, mountain garden, Meijer, 1953 (L); Nurta, 1953 (L). Between Tjibodas and Tjihoerang, in Eupaternia pallescens plain at Koebang tree trunk in the sun, 1380 m, Soekar 2247 (L). Namtrong Djimter, Jeger Bentang, rainforest tree trunk in the sun, 1450 m, Soekar 3187 (L). G. Sindoro, rainforest, 2400 m, Leeuwen 8889 (L). Without precise locality, Gerker, s.n. (L); Braun, s.n. (L). Without precise locality, no collector cited, s.n. (BR). Without precise locality, Blume, s.n. (JE) SUMATRA. Mt. Soga, near summit, mossy forest, 2000 m, Meijer 6126 (L). Korthals, s.n. (L, H, O, S). West coast, Mt. Merapi, west side, Prim. forest on branch, Waalkes 2236 (L); 1700 m, Belukar, on trunk, Waalkes 2270 (L). Aek na Vli, near Toba Lake, in mixed forest on bark, 1400 m, Wijk 1870 (L). Without precise locality, no collector cited, s.n. (S, C). BORNEO. E. Kutai, peak of B. papan, terr. Beul, on Magnoliaceae, 600-700 m, Meijer B1968 (L). N. Borneo, west coast res, Mt. Kinabalu, near Paka Cave, c. 9700', Meijer Bll-903 (L). NEW GUINEA. Star Mts. Mt. Antares, on rotten trunk of tree in rain forest, 1500 m, van Zanten 385b (BM). Eastern Highlands District, Arau, low on a tree in Castanopsis, oak forest, 1400 m, Brass 31976a (L). Sepik District, Ambunti Subdistrict, summit of Sumset, Mt. Hunstein, in elfin woodland on summit plateau, epiphytic on upper branches of small tree, c. 5000', Hoagland and Craven 10985 (L). Western Highlands Dis trict, Wabag area, Ranges south of Wabag, Nothofagus lower montane rain forest, corticous, 8000', Robbins 2859 (L). AUSTRALIA. Queensland, Ravenshoe, Watts 660 (H). SRI LANKA. Beim Stausee van Kandy, c. 55 m, Herzog, 1906 (BM).

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49 MALAGASY REP. "Certainly f rom Madagascar," no collector cited, s.n. (BM). M. rufum has the smallest and widest leaves of any of the species. The widely ovate leaves, with a short abrupt acumen, are easily recog nized. The imperforate endostome is similar to that of M. strictum but smaller, and the seta is the shortest of all Macrohymenium species. M. rufum is the only species in which som e of th e inner perichaetial leaves are blunt.

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50 2. MACROHYMENIUM ACIDODON (Mont.) Dzy et Molk., Musci Frond~ Ined. Archip. Indici 6:168. 1848. (Figure 9) Leskea acidodon Mont., in Ann. sc. nat. 1845, p. 96, t. 5, f. 4 and Syll. p. 19. 1845. TYPE: Reunion, no collector cited s.n. (holotype, L). Macrohymenium laeve Thwait. and Mitt., J. Linn. Soc. Bot. 13:317. 1873. TYPE: Ceylon, Thwaites 236 (holotype, NY-MITT; iso types, 0, H-BROTH, PC, BM). New synonomy. Macrohymenium acidodon var. acutissima Besch. ex Par., Ind. Bryol. 1108. 1898 nom. inval. in synon. err. pro M. a. fo. Besch., Ann. Sc. Nat. Bot. ser. 6, 10:292. 1880. TYPE: Madagascar, Perville, s.n. (holotype, L; isotype, L, H-BROTH). New synonomy. Rhegmatodon madagassus C. Muell. ex Geh., Abh. Naturw. Ver. Bremen 7:211. 1882. TYPE: Madagascar. forest of Ambatondrazaka, Rutenber, s.n. (holotype, O). Rhegmatodon acidodon (Mont.) Par., Ind. Bryol. 1108. 1898. Rhegmatodon la e vis (Thwait. and Mitt.) Par., Ind. Bryol. 1109. 1898. Rhegmatodon acidodon var. acutissima (Besch.) Par., Ind. Bryol. 1108. 1898. Plants with leaves, .89-1.45 mm long, .29-.58 mm wide, with an average length to width ratio of 2.77. Leaf cells uniform in length from the base to the tip of the leaf. Six to eight golden-colored in flated quadrate alars. Perichaetial leaves slightly serrate, ecostate or with a faint trace of a costa on some of the large inner leaves.

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51 Seta from 9-11 mm in length, 115-180 min width, slightly twisted, quite smooth at the bottom with hyaline protuberanc e s toward top. En dostome teeth perforated on their midline with un e ven irregular p e rfora tions. The perforations are mostly in the top half of the tooth, oc casionally producing a bifid tip. Endostome teeth ranged from 350-500 m and the exostome teeth ranged from 200-250 m. Habitat: On branches and rocks, 1800-2500 m. Distribution: Malagasy Rep Reunion, Seychelles, Sri Lanka, N. Born e o. MALAGASY REPUBLIC. Forest of Ambatondrazaka, Rutenberg, s.n. (O); Hildebrandt, 1880. Without precise locality, no collector cited, s.n. (0, L); Hildebrandt, 1880 (L); Perville, 1879 (H, 0). R&UNION. Without precise locality, Lepervanche, 1879 (0, BM, H, PC); no collector cited (0, L); Richards (BM); De ",'Isle (PC); Rodriguez (PC); Plain au Cafres, De L'Isle (BM). SEYCHELLES. Without precise locality, D e L'Isle (PC). SRI LANKA. An baumen beim Rambotta pass, c. 2000 m, H e rzog 115 (H, JE, L). Central Province, Thwaites 236 (T, PC, NY, 0, BM). An Baumen beim Tee von Nuwara Eliya, 1800 m, H e rzog, s.n. (PC, S). Neuwara Eliya, rotting stump in jungle, Binstead 183 (BM). BORNEO. N. Borneo, West Coast Res. Mt. Tambuyokon, c. 15 miles NE of Kinabalu Peak, summit area on branches and rocks, c. 8 000', Meijer Bll-378 (L). M. acidodon differs from all other species by the uniform size of the leaf cells from base to tip. The endostome of M. acidodon is also uniquely perforated with irregular, uneven slits, mostly in the upper half of the precesses. This sometimes produces a bifid appearance.

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52 3. MACROHYMENIUM MULLERI Doz. and Molk. Musci Frond. Ined. Archip. 6:168. 55. 1848. TYPE: Java, Mt. Gede and Talaga-Bodas, Korthals, s.n. (cotypes, 0, C, H, S, L). Borneo, Mt. Sakoembang, Korthals, s.n. (lectotype, L). Sumatra, Batang-Bessie, Korthals, s.n. (Figure 10) Rhegmatodon mulleri (Doz. and Molk.) C. M~ell., Linnaea 36:21. 1869. Macrohymenium mulleri forma pubea Dix., in schedula, 1909. TYPE: Java, Mt. Pangerango, Palmer & Bryant 982a (holotype, BM). New synonomy. Plants large, the branches thick and curved, leaves 1.9-2.53 mm long and .42-.66 mm wide, with an average length to width leaf ratio of 4.0. Leaf cells at base much longer (10:1) than cells at the tip (3:1). Twelve to twenty inflated, colored, quadrate alar cells. Seta from 811 mm in length, 130-150 min width. The endostome teeth perforated on the midline nearly the whole length with very uniform and regular perforations giving a scalloped appearance to the midline. Occasionally the tip of a tooth is bifid. Endostome teeth ranging from 250-300 m, the exostome teeth from 190-200 m. Habitat: On trees and ravine in very moist forest, usually found at elevations from 1200-1800 m, but occasionally up to 3100 m. Distribution: Java, Borneo, Malay Peninsula, and Malagasy Rep. JAVA. Mt. Pangerango, 4-6000', Motley, 1906 (NY); 7-10,000', Motley, 1906 (BM); Palmer & Bryant 982a (BM). Prov. Preanger, In decliv. austral. mantis Pangerango, in horto montano Tjibodas, Regio nubium,

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53 1420 m, Schiffner 12729 (S); in Cinchoneto, Daradjat prope Garut ad truncos putridos, Regio nubium, 1730 m, Schiffner 12127 (L). Mt. Salak, Binnendijk (S, L). Rawa gajonggong Tjibeureum, plain of gajongong, tree trunk in the sun, 1700 m, Soekar, 1949 (L). West Java, Tjibodas, Mt. Gede, Batu gonggang, Nature reserve, in rain forest, on Vernonia arborea base of trunk, shadowed, 1600 m, VanOostatroom 14235 (L). Without precise locality, Schroted, 1927 (H). BORNEO. In Sakoembang, Korthals, s.n. (L). Kinabalu, Low, 1859 (BM); Below Kamborangah, ravine in wet montane forest, 6000', Richards R5758 (L). MALAY PENINSULA. Gunong Tahan, Pahang, Ridley 1035 (BM). MALAGASY REPUBLIC. Without precise locality, no collector cited, s.n. (S). M. mulleri is distinguished from the other species of Macrohymenium by its large size, and its long, narrow leaves with many alar cells. The endostome of M. mulleri is differentiated by its peculiar pattern of perforations, and by its generally smaller peristome. Although M. mlilleri is the most robust of all the species, it is marked by the smallest peristome and a very thin seta.

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4. MACROHYMENIUM STRICTUM Bosch. and Lac., in Doz. & Molk., Bryol. Jav. 2:114. 1865. TYPE: Borneo, Mt. Kinabalu, Low, s.n. ex herb. Hook (holotype, NY; isotypes, o, S, L). 54 (Figure 11) Rhegmatodon strictus (Bosch. and Lac.). C. Muell., Linnaea 36:21. 1869. Macrohymenium nova-guineense Reim., Hedwigia 69:129, 1929. TYPE: Nova Guinea neerlandica, Mt. Doorman, Lam 1701 (not seen). New synonomy. Macrohymenium acidodon (Mont.) Doz. et Molk. fo. longisetum, in schedula. TYPE: Madagascar, Massif de L'Anjanaharibe, Humbert, Capuron and Cours, 1951 (holotype, PC). New synonomy. Plants moderate to large, the leaves 1.0-1.43 mm long, and .37-.55 mm wide, with an average length to width ratio of 2.7. The leaves ovate lanceolate with a long acumen. The leaf cells generally uniform in size over most of the leaf (3:1), much longer at the base (10-12:1). Eight to twelve inflated, quadrate, colored alars. All of the perichaetial leaves with acuminate tips. Seta from 8-21 mm in length and 200-372 m in width. Endostome twice as long as the exostome; endostome teeth not perforated. Endostome teeth ranging from 550-750 m; exostome teeth ranging from 300-410 m. Habitat: On trees and ground in subalpine forests and open thickets, occasionally found 1000-2000 m, but usually above 3000 m and up to 4370 m. Distribution: Malagasy Rep., Seychelles, Sri Lanka, Borneo, Sarawak, Sumatra, New Guinea, Australia, and the Philippines.

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55 MALAGASY REP. Massif de L'Anjanaharibe (Pentes and Sammet Nord), a L'ouest D'Andapa Haute AndraMonta, Bassin de la Lokoho, nord-est, foret ombrophile sur gneiss et grantie, 1000 m, Humbert, Capuron and Cours, 1951 (PC). (PC) SEYCHELLES. Pres la baie Sainte Anne, no collector cited, s.n. SRI LANKA. Nuewara Eliya, foot of tree by road, Binstead 181 (BM). BORNEO. Mt. Kinabalu, Low, s.n. (0, S, L, NY). North Borneo, West Coast Res., Mt. Tambuyokon, c. 15 miles NE of Kinabalu, Meijer Bll-473; ultrabasic area, 9000', Meijer Bl0-242 (L). North Borneo, Kota Belud distr., one mile n. of Kambarangan camp on path to Paka Cave, Mt. Kinabalu, common on ground in turf under an open, scrubby growth, aver aging 12' high, dominated by Ericaceae, Dacrydium and Phyllocladus, in fairly pure, abundantly fruiting tufts up to 9" diameter, moist and shady, deep humus over shallow, grey loam, 2750 m, Wood 1539 (L). SARAWAK. Without precise locality, Oxford Exped. (1932) 1631 (BM). SUMATRA. Sumatra occid, ad declif. occid. mantis ignivomi Merapi, regio nubium, 1800-2050 m, Schiffner 12724 (L, PC). NEW GUINEA. Mt. Albert Edward, common on forest trees, 3680 m, N. Guinea Exped. of Am. Mus. of Nat. Hist. 4431, 4438, 4439, 4435 (NY). Western Highlands, Kubor Range, Mt. Kinkain, on tree trunk, alpine shrub bery, 3580 m, Vink 16178 (L). Western Highlands, Wabog Area, Sugarloaf Mt. area, montane rain forest, epixylic, 10,000', Robbins 2805 (L). Eastern Highlands, Bismarck Ranges, Mt. Wilheim, on trees in grove at base of Bogonota Ridge, SE of Lake Aunde of edge of valley floor, 11,000', Weber and Mcvean B-32129 (LD, GL); Pindaude, Mt. Wilheim trail, epiphytic on Coprosma sp. in alpine tussock grassland, locally common, 14,000',

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56 Weber and Mcvean B-33522 (TENN); on dead wood in edge of subalpine forest 3700 m, Brass 29994 (L). NE Wilhelmina, top, epiphyte, 3650 m, Brass and Myer-Drees 9775 (TENN). Lake Habbema, enveloping branch in open thickets, 3225 m, Brass 9453 (L, TENN). AUSTRALIA. Molanda, Queensland, Watts 623, 664 (H). Ravenshoe, Queensland, Watts 662, 657 (H). PHILIPPINES. SE Mindanao, Todaya, Mt. Apo, on tree, 1220 m, Williams 2675 (NY, H). M. strictum is distinguished by its larger leaves with a long, suddenly constricted acumen. It also has the largest peristome of any species. Though the imperforate endostome closely resembles that of M. rufum, it is differentiated by its much larger size. The seta is also much longer and more robust than that of M. rufum, and the peri chaetial leaves more acuminate. M. strictum is a species growing at very high altitudes, usually above 3000 m, while M. rufum is found at lower altitudes, from 1700 m down to 600 m.

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DISTRIBUTION AND HABITAT OF THE GENUS MACROHYMENIUM Species of Macrohymenium have been collected only in the Old world tropics (Fig. 13:1-2; Fig. 14:1-2). The genus is essentially insular, occurring from the Malagasy Republic through Sri Lanka and Indonesia to northeastern Australia. Species of Macrohymenium are very similar in their distribution. Every speci e s has been found from both Borneo and the Malagasy Republic, and all but one, M. miill e ri, f:r:om Sri Lanka. Although the species of Macrohymenium occupy areas very similar in latitude, the altitudinal rang e s of ea c h vary widely. M. strictum has been collected primarily above 3000 m. M. acidodon is found at relatively lower altitudes, 1800-2500 m, while M. miilleri is, for the most part, encountered between 1400-1700 m, but has been reported from as low as 600 m. Macrohymenium is primarily a corticolous genus, al though a few specimens have been collected on rocks and soil. Collec tion data for most specimens are so scarce that it is difficult to be more precise in describing the habitat of the genus. 57

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DISPOSITION OF NAMES OTHER THAN SYNONYMS IN MACROHYMENIUM The following list includes the names of in schedula specimens, as well as validly published specimens, where examination of the types showed them to belong to some other taxa than Macrohymenium. Most of these specimens were collected in areas for which adequate keys are not available, and only limited attempt was made to assign these taxa to genera. For the purpose of completeness, previous exclusions and transfers are included in this list. Macrohymenium cuspidatum Mitt. in Kiaer, Forh. Vid. Selsk. Christiania 1882 (24): 45. 1883. TYPE: Assam, Mt. Khasia, Griffith (No. 212, Hb. Kew. No. 516 intermixed) (isotype, 0). Index Muscorum incorrectly attributes a new species, Sematophyllum cuspidatum, to Kiaer on the basis of this specimen. Actually, he notes that the peristome was destroyed by age and unknown to him, and that it perhaps belonged to the genus Sematophyllum. Having seen the speci men, I would agree. Macrohymenium densirete Broth. in Fleisch., Musci Fl. Buiten zorg 4: 1192. 1923 nom. nud. in synon. TYPE: not given = Clastobryum conspicuum Fleisch. Macrohymenium gracillimum C. Mull. ex Kiaer, Forh. Vid. Selsk. Chri3tiania 1882 (24): 19. 1883 nom. inval. TYPE: not given. 58

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59 Macrohymenium kunzii C. Mull., in schedula, 1868. TYPE: India, Pegu, Yomah, Kunz 2939 (holotype, H). Broad b o dies were present on this specimen. It is clearly in th e Sematophylla ceae and probably in the Clastobryoideae. Macrohymenium sinense Ther., Bull. nc. Int. Geogr. Bot. 19:20. 1909. TYPE: China, Prov. Kweitschou, Lou-mong-touan, Fortunat, 1903 (holotype, S) = Giralcliella levieri C. Mull. fid. Reim., Hedwigia 71: 62. 1931. I have s e en this specimen and agree. Macrohymenium wichurae Broth. in Fleisch., Musci Fl. Buitenzorg 4: 1364. 1923 nom. nud. in synon. TYPE: not given.

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LEGENDS FOR THE FIGURES To facilitate reference and comparison of the characters pre sented in Figures 1-11, the following standardized magnifications have been used: Habits XB Spores X466 Calyptrae X30 Leaf tips Xl64 Opercula X30 Alar cells Xl64 Endostome teeth Xl25 Basal celL:.; Xl64 Exosto:rae teeth Xl25 J uxtacostal cell3 Xl64 Exothecial cells Xl64 Antheridia Xll7 Setae Xll7 Archegonia Xll7 Perichaetia X30 Portion of branch X30 Vaginules X30 Perichaetial leavE.s X30 eseudoparaphyllia Xl64 Branch leaves X30 Perichaetial teeth Xl64 The maps in Figures 12-14 show the distribution of the various species of Rhegmatodon and Macrohymenium on a world-wide basis. Only the locations of those specimens examined and verified by the author have been plotted. The base map is a modified Denoyer Semi-elliptical Projection. Figures 15 and 16 contain the major published illustrations of these genera. 60

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Figure 1: Pictures 1-18. Rhegmatodon spp. 1-3. Habit. 4-5. Operculae. 6. Perigonium. 7. Perichaetial Leaf Cells. 8. Archegonia and Paraphyses. 9. Antheridium and Paraphyses. 10. Capsule and Operculum. 11. Calyptra. 12-16. Perchaetial Leaves. 17. Perchaetial Leaf Tip with Tooth. 18. Exothecial Cells.

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62 2 i 5 6

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Figure 2: Pictures 1-22. Rhegmatodon spp. 1-2. Exostome Teeth on Capsule. 3. Exostome from Top. 4. Spores. 5. Endostome Teeth. 6-12. Exostome Teeth. 13-22. Pseudoparaphyllia.

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i I 3 6 7 8 17 2 ~e v 4 ,. I I 9 10 20 64 11 12 16 21

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Figure 3: Pictures 1-7. Rhegmatodon spp. 1-7. Leaf Cells.

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66 1 2 4 5 6 7

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Figure 4: Pictures 1-13. Rhegmatodon declinatus. 1-2. Teeth. 3. Rough Seta. 4. Terminal Portion of Branch. 8-12. Branch Leaves. 13. Exothecial Cells. Endostome 5-7. Leaf Tips.

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68 2 3 4 5 6 7 ,. -. 8 9 10 11 12 13

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Figure 5: Pictures 1-23. Rhegmatodon polycarpus. 1-2. Teeth. 3. Smooth Seta. 4. Terminal Portion of Branch. Tips. 10-22. Branch Leaves. 23. Exothecial Cells. Endostome 5-9. L e af

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70 3 4 17 18 19

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Figure 6: Pictures 1-14. Macrohymenium spp. 1-3. Habit. 4. Antheridia. 5-6. Alar Cells. 7. Perchaetiurn. 8. P e rigonial Leaf. 9-10. Basal Leaf Cells. 11. Seta with Warty Proj e ctions. 12. Vaginule and Seta. 13-14. Basal Perichaetial Leaf Cells with Pores.

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72 2 3 1 4 8 7 11 12 13 14

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Figure 7: Pictures 1-15. Macrohymenium spp. 1-2. Endostom e and Exostome Teeth. 3. Complete Endostome. 4. Spores. 5-8. Exostome Teeth. 9. Complete Exostome. 10-11. Calytrae. 12-14. Operculae. 15. Exothecial Cells.

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74 3 1 2 4 r ) l -' El .:._ ~ Tig: I 9 5 6 7 8 .... 10 11 12 13 14 15

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Figure 8: Pictures 1-19. Macrohymenium rufum. 2-3. Leaf Tips. 4-5. Endostorne Teeth. 6-7. Perchaetial Leaves. 11-12. Cells of Leaf Base. 19. Alar Cells. 1. Portion of Branch. Cells of Midleaf. 8-10. 13-18. Branch Leaves.

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76 8 ___ 9 11 12 13 14 15 16 17 19

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Figure 9: Pictures 1-14. Macrohymenium acidodon. 1. Portion 2-3. Leaf Tips. 4-5. Endostome Teeth. 6. Perchaetial Leaf. Cells of Midleaf. 8, 14. Alar Cells. 9. Cells of Leaf Base. Branch Leaves. of Branch. 7. 10-13.

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78 2 4 5 8 9 10 11 12 13 14

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Figure 10: Pictures 1-16. Macrohymenium muelleri. 1-3. Endostome Teeth. 4-5. Leaf Tips. 6-7. Cells of Midleaf. 8-9. Cells of Leaf Base. 10. Perchaetial Leaf. 11-14. Branch Leaves. 15-16. Alar Cells.

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1 ; /j. I I .' It ~ n 2 10 11 8 0 8 9 I ,_. 12 13 14 15 16

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Portion of Perchaetial 13-14. Alar Figure 11: Pictures 1-14. Macrohymenium strictum. 1. Branch. 2-3. Leaf Tips. 4-5. Endostome Teeth. 6. Leaf. 7. Cells of Midleaf. 8-12. Branch Leaves. Cells.

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82 1 4 5 6 13 8 9 10 11 12 14

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Figure 12: 1-2. Distribution of Rhegmatodon spp. 1. Distribution of Rhegmatodon polycarpus. 2. Distribution of Rhegmatodon declinatus.

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do n po7,ycarpus Rhegmato 1 d declinatus Rhegmato on 2

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Figure 13: 1-2. Distribution of Macrohymenium spp. 1. Distribution of Macrohymenium rufum. 2. Distribution of Macrohymenium acidodon.

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8 6 Maarohymeniwn rufwn Maarohymeniwn aaidodon 2

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Figure 14: 1-2. Distribution of Macrohymenium spp. 1. Distribution of Macrohymenium muelleri. 2. Distribution of Macrohym e nium strictum.

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88 ... Maarohymenium mueZZeri 1 .... '-~ '{" l:-:-. )------------------7'1--7' Maarohymenium strictum 2

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Figure 15: Pictures 1-9. Previous Illustrations of Rh eg matodon. 1. Anhymenium polycarpon Griff., Calcutta Jour. Nat. Hist. t. 16. 1843. 2. Anhymenium polysetum Griff., Icones Plant. Asiat. t. 97, f. 2. 1849. 3. Macrohymenium serrulatum Doz. & Molk. Musci Frond. Ined. Archip. Indici t. 56. 1848. 4. Rhegmatodon serrulatus (Doz. & Molk.) Bosch. & Lac., in A. Engler & K. Prantl, "Die naturlichen Pflanzenfamilien," Band 11, p. 298, f. 648. 1925. 5. Regmatodon declinatus (Hook.) Brid., Bryol. Univ. 2, t. 9. 6. Regmatodon declinatus (Hook.) Brid., Schwaegrichen, Species Muscorum Frondosorum, t. 204. 1827. 7. Ptero gonium declinatum Hook., Trans. Linn. Soc. Lond. 9: t. 26, f. 3. 1808. 8. Enlargement of part of no. 7. 9. Original pencil drawings by Hooker found with the holotype (NY).

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1 4 b flA ?.~*i 7 8 2 5 J 3 6 9 I I 1\ 11 90

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Figure 16: Pictures 1-6. Previous Illustrations of Macrohymenium. 1. Macrohymenium mulleri Doz. & Molk., in A. Engler & K. Prantl, "Die naturlichen Pflanzenfamilien," Band 11, p. 445, f. 745. 1925. 2. Macrohymenium mitratum (Doz. & Molk.) Flsch., in Fleischer, Die Musci der Flora von Buitenzorg, 4: 1365, f. 221. 1923. 3. Macrohymenium mulleri Doz. & Molk., Muse. fr. ined. Archip. ind. pp. 167-68, t. 55. 1848. 4. Macrohymenium rufum Doz. & Molk, Musci fr. ined. Archip. ind. pp. 16869, t. 54. 1848. 5. Leskea acidodon Mont., Ann. sc. nat. p. 96, t. 5, f. 4. 1845. 6. Original pencil drawings found with the holotype (L).

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1 3 I .. / ~ 1: ) ,\ ; rr, /Y. r A 1 I i ., I,' ) ~: I\ 1 ~ '' \ I '-, / J, ? .\~1 ~-), t ._,. .. ( I i \ h C=u .. \ i \; ~ e:. L ,, ,,. I .~ -~ c:".. "'' I ,. ,. JY : .L,,,.-J,i:t ,,,.,;1,,,,;,11 U 5 92 2 4 6

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APPENDIX I HISTORICAL PERSPECTIVES IN TAXONOMIC BRYOLOGY Modern classification is the end-product of a long series of decisions regarding the relative importance of certain characters as indicators of some "natural" relationship. Naturally, whatever biases or assumptions are used to determine these characters will be reflected in our final classification. This appendix is provided to supply some background with regard to the historical development of taxonomic bryo logy The basic points to be covered are as follows 1. Bryology is a relatively new discipline, having begun in the modern sense with Hedwig only two centuries ago. This, and the fact that there are so few workers in the field, had slowed the growth of knowledge so that bryology is still in the exploratory and descriptive stages of taxonomy. 2. The current classification of mosses (Dixon) is the product of a long series of individual opinions and tradition with its roots deep in the folk science of various civilizations. A brief history of the ebb and flow of taxonomic opinion in bryology, especially with reference to the concept of the genus, suggests that each taxon is an abstraction in which we have assembled certain plants for the purpose of understanding and separated them from other groups. 3. Classification of mosses, like other plants, has been tradi tionally based on morphological or anatomical characters which are 93

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94 readily observable by ordinary methods. Characters which require great time to discern are impractical for the purpose of putting a nam e on a plant. 4. The choice of critical characters in bryology is arbitrary, based on speculative logic of individual workers, because there is no fossil evidence to suggest what characters should be considered primitive or advanced. Origins of the Generic Concept From the beginning of time man has expressed a need to name and classify the various aspects of his sensory world. Even the dim light of prehistory reveals that, within the folk science of all civilizations, those botanical aspects of man's environment which were of greatest importance to his survival were distinguished and named. An obvious example of this would be man's ability to discover and retain information about edible and non-edible plants. With increasing experience, finer distinctions would be required for plants previously included under the same name. Bartlett (1940) suggests that the original name then attained generic significance, as a basis for specific names. He gives "grass" as an example of this analytical pro cess. "Many kinds of grass are so similar that we can hardly b e lieve that the concept 'grass' was not more ancient than the distinction of particular kinds" (p. 349). On the other hand, some groupings undoubtedly arose when people noted previously unrecognized similarities and combined various plants which are superficially very dissimilar. Bartlett believed this to be the case for the generic concept of "fern." On the contrary, the generic concept 'fern' is a technical one, depending upon close observation, so wh e n we find a people of relatively low culture, such as the

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Batak of Sumatra, defining extremely diverse plants as 'fern' pretty much as the modern botanist does, on the basis of a relatively obscure characteristic, namely, the leaf-borne sporangia, we feel sure that a genus has been set up by synthesis of things superficially very unlike. (Bartlett, 1940, pp. 349-350) 95 Thus, delimitation of various groups or taxa came about by one or both of these processes of analysis and synthesis. This reasoning applies equally well to the description, naming, and classification of mosses. For instance, one would expect very inclusive grouping for those plants which were inconsequential in the day to day existence of a people. This is true for the Musci. Bartlett notes, from personal experience with the Batak of Sumatra, that lumut 1 suffices as an inclusive generic name for almost all moss-like plants. He further states that Classification of lumut is hardly attempted, but the conspicuous Leucobryum has a generic name. The condition of their moss nomenclature is after all not so very dif ferent from that in scientific systematic botany before Dillenius. (Bartlett, 1940, p. 354) The role of utility in the development of plant nomenclature is certainly not restricted to the Batak. The bryophytes have always been of limited value to man and therefore were not studied until fairly recently. An exception to this is Sphagnum whose utility has been recognized almost everywhere it is found. For this reason, it was so sufficiently defined in pre-Linnaean folk science that modern workers 2 have agreed to designate the beginning of this modern genus concept with Linnaeus' 1753 publication of Species Plantarum, starting point for the nomenclature of higher plants. Even today, ... bryophytes are almost totally free from economic involvements. Excepting the modest peat moss industry, the 1 Pardembanan dialect (a sub-Toba dialect) of Asahan, Sumatra. 2 Third International Botanical Congress, Vienna, 1905.

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small favor they find in a few Japanese gardens and th e ir limited usefulness as packing materials for shipping live plants, mosses, and liverworts have no economic worth. (Anderson,1974, p. 56) 96 Due to the purely academic nature of bryological studies, the starting point for the nomenclature of bryophytes, except for Sphagnum and the Jungermanniales, was designated 1 with the much later publication of Hedwig's Species Muscorum (1801). Closer examination permitted inves tigators of moss-like plants to segregate originally large inclusive genera into smaller genera and species. Steere (1947) indicates the extensiveness of this process by noting that th e original 35 g e n e ra of Hedwig had multiplied to more than 700 genera by 1947. Thus, d e tailed analysis has been the most important process in the development of moss taxonomy. History of the Concept of the Genus in Musci In presenting a historical perspective of the chang e s which have taken place in the concept of genera of Musci, I have r e li e d h ea vily on the classic presentations of Steere (1947), Florschutz (19 6 0), Margadant (1968), and Dixon (1932). The first book dealing explicitly with the mosses and liv e rworts was Historia Muscorum by Dillenius (1741). Tw e nty gen e ra w e r e d es cribed and, following the general concepts of his day, h e includ e d e v e ry lowgrowing plant which was not clearly a flowering plant, fern or a fungus" (Margadant, 1968, p. I). In addition to h e patics, lycopods, lichens, and algae Dillenius was acquaint e d with six m o d e rn ge n e ra of mosses: Sphagnum, Mnium, Fontinalis, Hypnum, Bryum, and Polytrichum. Except for Sphagnum, these genera, while including some presently 1 Fourth International Botanical Congress, Brussels, 1910.

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97 recognized species, were very broad and inclusive. Bryum, for example, included most of the mosses which grow in tufts and produce their sporophytes terminally (i.e., the acrocarpous mosses) while Hypnum was in clusive of those mosses which produce the sporophyte laterally and are generally mat-forming and prostrate (i.e., the pleurocarpous mosses). Dillenius made no attempt to show intergeneric relationships or to suggest a systematic arrangement. In Species Plantarum, Linnaeus added two new moss genera, Phascum and Splachum, to the six of Dillenius. At the same time he restricted the term Musci to mosses more nearly in the modern sense. He did include, however, the hepatic genus Porella, some other hepatics under Mnium, and the fern ally Lycopodium. We should remember that Linna e us was as significant to bryological nomenclature as he was to the rest of taxonomy, being the first to establish the universal us e of binomials in nomenclature. Linnaeus also did not attempt to show a systematic arrangement of intergeneric relationships. The next landmark in taxonomic bryology is the work of Johannes Hedwig. Perhaps the clearest indicator of his importance to bryology is the fact that his magnum opus, Species Muscorum, publi s h e d po s humously by Schwaegrichen (1801) was chosen as the starting point for all legitimate nomenclature of Musci. This decision reflect e d a recognition of his careful observation and meticulous description of characters previously Wlknown or unappreciated. Florschutz has stat e d, Hedwig was one of the best observ e rs of his day. he used and improved the microscope which even at that time was sometimes considered as being unreliable. He d e v e lop e d a special preparation-technique and a t e chnique for dr a win g magnifications by microscope. He was th e first to publish very accurately the form and structure of the small es t organs of cryptogams. Several of these organs w e re alr e ady

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known, but nobody had perceived them so exactly as Hedwig did. (Florschiitz, 1960, p. ix) 98 In Hedwigii Fundamenta Historiae Naturalis Muscorum Frondosorum (1782) he included, for the first time in bryological literature, a key to genera. The key clearly shows Hedwig's concept of the relationships between these genera. The presence or absence of the peristome, the form and number of the peristome teeth, and the form of the male "flower" (perigonium), were the critical generic characters. In his final concepts he stressed the position of the male flower on the stem, rather than its form. Species Muscorum includes a glossary of terms, k e y to g e n e ra (35), descriptions of many species, and 77 plates published originally as hand-engraved colored masterpieces. A storm of controversy surrounded Hedwig's choice of characters, especially those microscopic details; viz, the male "flower." Steere (1947) records an interesting criticism of Hedwig's 'Stirpium Crypto gamicarum" (1787-1797) by one A. Menzies (1798). The laborious works of the c e l e brat e d and pers e rv e ing Hedwig have, of late years, thrown much light upon th e subjects of this natural order; but the general complaints agairist his new arrangement of it ar e that his genera are too artificial, and that their characters are taken from parts so minute and difficult to examine, that they rather tend to perplex and discourage a young beginner in his investigations, than aid his pursuits in acquiring a scientific knowledge of this intricate tribe. While it can therefore be avoided, no gen e ric or specific charac ters ought ever to be adopted, that cannot easily and distinctly be seen by the assistance of a single lens magnifier, such as botanists commonly carry in their pockets. (Steere, 1947, pp. 248-249) This quote is particularly interesting because it reveals that most botanists or bryologists did not at this time make regular us e of the microscope. The "minimal accessory equipme nt" us e d to accompany a

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99 bryologist's sharp eyes was a hand magnifier. Only when the microscope became readily available were Hedwig's characters generally accepted. This criticism is perhaps more clearly appreciated when we recall that Hedwig was the first to discover and describe the male "flower." His system was a vast improvement over earlier treatments of Musci. It is currently felt that Hedwig's system was too artificial in that it gave almost exclusive weight to the sporophyte (i.e., peristome charac ters) while largely ignoring gametophytic characters. This resulted in grouping under the same generic name many mosses with comparable peristomes, but completely dissimilar gametophytes. This does, in fact, represent one extreme facet of the systematic problem presented by plants which are composed of both the haploid and diploid generations, and where the sporophyte is persistently attached and at least somewhat dependent on the gametophyte. Nevertheless, The main characters Hedwig used are still the very important characters for our present classification of the mosses. Although our delimitations of genera changed, the discovery of the importance of these characters came from the brilliant thoughts of this eminent bryologist. (Florschutz, 1960, p. xx) In 1819 Sam Bridel published his Mantissa Muscorum as an apparent supplement to his earlier works on mosses. Although adopting Hedwigian classification, he did introduce two new characters; the position of the female flower (perichaetium) and various characters of the cillyptra. This was considered a great improvement over characters of the male flower since both capsule position and the form of the calyptra are features which can be seen with the unaided eye. In this work, the terms acrocarpi and pleurocarpi were introduced for the first time as the two most important classes of his Section I, Olocarpi. These char acters became the pivotal points of classification in Bridel's famous

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100 two volume work, Bryologia Universa (1826). On the basis of variation in calyptra characters, Bridel proposed several new genera. During the period 1811-1842, there appeared three supplements to "Species Muscorum." These were the work of Christian Friedrich Schwaegrichen, whose respect for his teacher is demonstrated by the fact that all three supplements appeared under the name of Hedwig. Schwaegrichen also proposed new genera. The first bryologist to reintroduce the potential use of gameto phytic characters in the classification of mosses was Carl Muller in his two volume, Synopsis Muscorum Frondosorum, 1848-1851. Mosses were divided into two classes; Cleistocarpi and Stegocarpi. The former included those mosses with sessile sporophytes, sphaerical to sub sphaerical capsules with no definable mouth or peristome. The Stegocarpi included those forms with the sporophyte on a seta, elongate, with an operculum, and a well-defined mouth and peristome. This latter class was divided into the Acrocarpi and Pleurocarpi, using leaf arrangement to delimit the still smaller groups of taxa. The sporophytic characters were then used to define genera. The use of leaf arrangement was not generally accepted by other workers, and thus his system was not adopted. One important contribution of this work should be noted. The various comprehensive treatments of mosses during the nineteenth century lacked a world-wide perspective, their taxonomic treatment reflecting a "local-flora" or regional outlook. Muller did attempt to compile in his work all of the known, described species of mosses to date. Even the monumental Bryologia Europaea (Bruch, Schimper, and Guembel, 1836-1854) did not present world-wide coverage of the Musci. Bryologia Europaea did establish a significant proportion of the generic names now in use in Europe and North America, many genera appearing for the first time

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101 as segregates of some of Hedwig's mammoth genera. Such well-known generic names as Thuidium, Pseudoleskea, Bracythecium, and Amblyst eg ium, all cleaved from Hedwig's Hypnum, date from this time. The tremendous importance of this work is suggested by the fact that most authors during the following 40 years adopted Schimper's classification in toto. The descriptions of species and detailed illustrations are among the very finest in bryological literature. In 1855, Schimper attached his Corollarium Bryologiae Europaea, in which he classified all of the European species of Musci. Here Schimper elevated the terms acrocarpi and pleurocarpi, coined by Bridel (1819), by designating his two main orders Musci acrocarpi and Musci pleurocarpi. Jaeger and Sauerbeck (1869-1878), in their Adumbratio, attempted a bibliographic treatment of all of the world's mosses, but simply inter calated the non-European genera into Schimper's system. Two important anatomical studies were published within th e l as t quarter of the nineteenth century which greatly influenced further innovations in the classification of Musci. Grundlinien zu einer vergleichenden Anatomie der Laubmoose, authored by Lorentz (1867-1868), aroused new interest in the importance of anatomical characters. He specially called attention to the usefulne s s of microscopic detail s in leaf cells. While leaf characters had been used prior to this tim e both in Bryologia Europaea and many of Muller's publications, many look to Lorentz for the most persuasive argument of the correctn e ss of this view. For example, Limpricht (1895) in his Die Laubmoose incorporates Lorentz's ideas within the older acrocarpous-pleurocarpous concept. (Die Laubmoose dealt only with European genera.) The second important anatomical influence on the classifi c ation of mosses was Philibert's Etudes sur la peristome which app e ar e d a s

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102 many separate papers in Revue Bryologique between the years 1886 and 1890. This work represents a magnificant investigation into the de velopment and structure of the moss peristome. Philibert was able to identify certain basic kinds of peristomes within Musci, greatly in creasing the feeling that the peristome reflected the phylogeny of the moss more clearly than the gametophyte. The occurrence of a similar kind of peristome in two taxa persuaded Philibert that they must be closely related. He could not conceive of such a complex structure as the periostome having evolved in two separate groups as the result of convergent evolution. The feeling that peristome types are basic and dependable indica tors of phylogenetic relationships is exemplified by the following statement from Dixon: Certain basic principles at any rate may be recognized and utilized for taxonomic purposes. For one thing, it is quite clear that the main types of peristome among the Byrales are of great phylogenetic importance, and are more or less primitive, viz. the Nematodonteae and the Arthrodonteae, the latter divided into Haplolepideae and Diplolepideae. No part of our classification based on gametophyte characters must cut across these broad lines. (Dixon, 1932, pp. 402-403) It would be an oversight to fail to indicate the great influ e nce the theory of evolution had on moss taxonomy. The following statement by Dixon will suffice to indicate this point. The gradual acceptance of the doctrine of evolution has placed all taxonomy on a new plane. Resemblances and differences between organisms mean something entirely different from what they meant before. It is the business of the taxonomist not only to tabulate and classify differences and resemblances, but so to classify them that the relation ships between the organisms shall thereby be brought out. Resemblances may indicate relationship, but th ey may not. Wide apparent differences may S\lggest wide divergence of origin, but they may be only apparent, not fundamental ones. The taxonomist has to lay down a system which shall, so far

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103 as possible, indicate the actual phylogenetic relationship of the plants with which he is dealing. (Dixon, 1932, p. 398) During the period from 1900 to 1923, Max Fl e isch e r publish e d his four volumes on Die Musci der Flora van Buitenzorg. This work presented a system of classification which has been hailed as the most cr e ative scheme of the twentieth century. It incorporates Philibert's peristome groupings, and the location of the sporophyte and various gametophytic characters. This is regarded as the first important treatment of the mosses from a non-European viewpoint and included a great numb e r of n e w families, orders, and g e nera. With few alterations, this s y s t e m is u se d among present-day bryologists. It was adopted by Brotherus in his treatment of the Musci in Engler and Prantl, Die naturlich e n Pflanzen familien (2 nd edition, 1925), and also was used as a basic outline of Dixon's classification, as presented in Verdoorn's Manual of Bryology (1932). It is the system which has been used as a basis for the present work. Modern Taxonomic Bryology In theory, modern taxonomists are "to lay down a system which shall, so far as possible, indicate the actual phylogen e tic relationship of the plants with which he is dealing" (Dixon, 1932, p. 398). This is not possible in the case of mosses for sev e ral reasons. In vi e w of th e general acceptance of the theory of evolution, charact e rs c hos e n to group taxa are to be selected because they are thought to b e st r e pr e s e nt phylogenetic relationships, i.e., primitive vs. advanced charact e rs. selection of a character as primitive, however, must result from th e exam1nation of the only direct evidence available, i.e., th e geologic The record. In the case of mosses, this source of information is virtually

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non-existent. The only bryophyte fossils, primarily hepatics, are nearly identical to extant forms. Anderson remarked that, Steere and Inoue (1972) (JHBL 35) ... summed up the status of present-day taxonomic and evolutionary knowledge of bryophytes in a devastatingly frank statement. "Although there are many published papers discussing the species con cept of bryophytes," they stated, "we still know little about the nature of species, and even the concept of taxo nomic rank is not clearly understood." To this pessimistic appraisal, they might well have added that we know even less about the age of bryophytes, their evolutionary origins directions and rates. (Anderson, 1974b, p. 8) 104 Thus, any attempted phylogenetic groupings must at this time be recog nized as purely speculative. A further problem arises from the fact that each moss consists of two different plant generations; the gametophyte and sporophyte. Mosses are unusually difficult to systematize because they present a double organism a sporophyte plant that lives at least in part parasitically on a green gametophytic plant ... If undue weight is placed upon characters derived from either generation alone, an artificial system like that of Hedwig's will be created. (Steere, 1947, p. 253) Hedwig's system exemplifies the extreme of ass1gning almost complete weight to sporophytic characters. While no single system of classification can be noted to exemplify the oth e r extreme it is clear that grouping on the basis of gametophytic characters alone would be equally artificial. The third possibility is to classify mosses on the basis of some combination of gametophytic and sporophytic characters. Our current classification systems reflect this philosophy. But the selec tion of "critical" characters, representing both generations, is equally arbitrary in view of the absence of fossils. The classification of bryophytes, then, will always be open to the charge of artificiality unless definitive fossils are found.

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Nearly all bryologists and vascular plant systematists agree that bryophytes are not ancestral to the vascular plants or their presumed fossil progenitors. It is also generally agreed that none of the three bryophyte classes recognized today are ancestral to any of the others. But popular and current as these views may b e th e y ar e ba se d on speculative logic and not facts. There is no fossil evidence to relate bryophytes to either the green algae or the tracheophytes .... unless additional fossil ma terial comes to light or some other fresh evidence is forth coming, the subject is likely to remain, as Richards (1959) puts it, "a fascinating subject for speculation." (Anderson, 1974a, p. 78) 105 Delimitation of species is considered less arbitrary than the higher taxa, because the criterion of degree of interbreeding is add e d. Theoretically, species will be separated by nearl y complete gaps in variability, and by "a complete or nearly complete barrier to interbreeding" (Cronquist et al., 1972, p. 6). The disability to freely interbreed will maintain differences between populations and allow further divergence. Unfortunately, present knowledge is scant r e garding the causes of variation, what factors produce barriers to int e rbreeding, and the role of asexual reproduction, etc., in r e ference to speciation. The nature of many bryophyte species is poorly understood, prin cipally because the causes of variation are not known. Thus, whil e th e "species is ordinarily considered by taxonomists to be a r e al entity, existing in nature, which we recognize and d e scribe but do n o t c r e at e ," we can see why some workers "have considered it to be a mere cr e at i on of the mind, simply an aid to understanding or cataloging natur a l d 11 ( t t 1 1972 6) The taxonomic c ateg o ri es 1vers1ty Cronquis e a ., p. above the rank of species have generally be e n r e cogniz e d as conv e ni e nt devices to show our concept of the degre e of r e lationship a mong s p ec i es

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All taxonomic groups are abstractions, in which we assemble, for purposes of understanding, certain individuals or groups of individuals, and separate these collectively from other groups. These groups are founded on the ensemble of.simi~arities and differences, our efforts at analysis being aided by the fact that diversity among living things is not continuous but exhibits gaps of varying size ... The basic difficulty with taxonomic groups above the rank of a species is that the precise rank of a group is not inherent but is determined by individual opinion and by custom, which is the sum of a series of individual opinions ... Where custom itself is divided ... one chooses the course one prefers, and custom remains divided. (Cronquist et al., 1972, p. 5) As Anderson has said generic criteria are evasive in any group and especially in bryophytes ... Bryophyte genera rest heavily on tradition, which is not always good science ... (Anderson, 1974, p. 10) l Ob Traditionally, the fundamental characters used in taxonomic classification are features of the external morphology of the organism. A predictable sequence of characters has been used since man first began to group plants. The most obvious characters of the plant, e.g., habit, flower color,leaf arrangement, etc., were first used for classification. Even today there is a feeling that genera of plants should be recognizable with little or no aid from magnifiers. Treating mosses in this manner would create very large, inclusive groupings. Sometimes severe criticism is directed at taxonomists who only sparingly use cytological or chemical characters. Howev e r, one of the foremost concerns of taxonomy has been to provide a practical way for scientists and laymen alike to put a name on a specimen. Characters which are not easily observed are impractical for this purpose. External morphology furnishes an amazingly larg e number of differences which can be readily obs e rved and has consequently been the chief source of the character s by which taxonomic groups are recognized and d e fin e d. Anatomical, physiological, biochemical, and cytologic a l characters are at least theoretically of equal importanc e

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107 to the morphological ones. Indeed, all morphological as well as other differences arise eventually from biochemical ones. On the other hand, the acquisition of sufficient data f~om these other sources so that one can be sure he is dealing with features of groups rather than of mere individuals is so time-consuming as to be often impractical. Data from th e s e sources tend to be concordant with data from morpholog y In some cases in which morphology does not provide conclusiv e evidence of affinities, information from these other sourc e s may tip the scales decisively to one side or the other. Not infrequently, however, when morphology is inconclusive, other data are equally ambiguous. (Cronquist et al., 1972, p. 5) With reference to vascular plants, Cronquist says that specific rank should not be applied to "plants of ordinary siz e which cannot b e di s tinguished by sharp eyes with only minimal accessory equipm e nt. On e should not have to do an Arbeit to put a nam e on hi s sp e cim e n." One further advance should be mentioned. Because of th e ir small size, mosses were relatively unknown until the development of light microscopy. We must remember that the microscopes of Hedwig's day were not capable of the magnifications and resolution of modern opti c al instruments. In the introduction to Hedwig's "Sp e cies Muscorum," Florschutz (1960) comments that the microscope which Hedwig obta i n e d from Schreber was a simple "Rheinthaler" with a magnification of SO X With this microscope he apparently did all of his investigations, al though he was later able to boost the magnification to 170-290 X M a gni fication, however, does not tell the whole story. Th e most critical feature of a lens system is its ability to r e solv e C e rtainly th e resolution of Hedwig's lenses was considerably less than would be found in modern lenses providing equivalent magnification. Th e nin e t ee nth century saw improvements in the grinding of lens e s which e v e ntually mad e it possible to resolve objects at some 2000 diameters (Gardn e r, 1 96 5). This is the limit set by the wavelength of light itself. Th e firS t achromatic lenses to correct for chromatic ab e rration w e r e n o t a vail able until 1830, and the sub-stage condenser was not d e sign ed until 1870

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10 8 The significance of this to bryological taxonomy of co urse, was the extension of our ability to view a new world of morphological char acters which could be used to define and separate taxa. And as micro scopes became "backpocket" tools of the biologist, microscopic char acters were accepted more and more as critical taxonomic features of mosses. Another important factor in the development of bryology and of the study of cryptogams in general, was that more and better microscopes became available. Microscopic details could be studied with greater precision and ease, and they began to play a more important role in the delimitation of species and other taxa. Life history studies became more accurate. This motivated an intensification of the work be gun by Hedwig. (Margadant, 1968, p. II) Today a similar situation exists with respect to the development of the electron microscope; both for transmission and scanning EM. Again the range of observable characters has been incr e as ed Biochemical and cytogenetic techniques have added even further possible characterizations. However, these characters are not universally available, many times requiring days in preparation. The presence or absence of characters which can only be seen with a scanning electron microscope are at this time impractical for general use in keys. Perhaps if this new technology becomes as commonplace as light microscopy, th ese char acters may supplant some of the larger traditional morpholo gica l char acters. Summary The history of the generic concepts for Rhegmatodon and Macro hymenium can be fully appreciated only in the context of th ese overall changes. The taxonomic histories of Rhegmatodon and Macrohym e nium span almost the entire history of bryology. Taxonomic concepts and opinions were continually changing as new and different specimens and characteriS t ics

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109 came to light. Corresponding changes were evidenced in th e concepts of these two genera, as the influence of these new characteristics grew in the general taxonomy of bryophytes. I believe that the history of these genera presents a nearly perfect "case in point" to show the practical implications of these influences in the classification of mosses.

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APPENDIX II PREVIOU S KE Y S TO THE SPECIES O F RHEGMATODON AND MACROHYMENIUM BY BROTHERUS AND KIAER Key to Species of Rhegmatodon s e nsu Kia e r (1882) A. Laeviseta The seta smooth, the operculum equalling 1/4-1/3 of th e d e op e rculat e capsule, the longitudinal walls of the e x ternal c e lls of the ca p sul e thickened throughout, the processes distinctly punctulate ...... 1 B. Scabroseta The seta scabrous, the operculum equalling n e arly 1/ 2 o f th e deop e rculate capsule, the walls of th e e xt e rnal cells of th e capsul e interruptedly thickened, the process e s smooth ............ 14 1. 2. 3. The branch leaves more narrow (2.00-2.58) ........... 2 The branch leaves wide (1.82-1. 8 5) .............. 11 With the maximum width of the l e af a littl e a b o v e th e base ... 3 With the maximum width of the l e af a littl e b e l o w th e mid d l e .. 6 The branch leaves more loosely imbricat e d, th e p e ri c ha e ti a l leaves with the teeth solitary or v e ry f e w coars e ly s e r r at e distinctly costate .. . . 4 Rh. schlotheimioides The branch leaves densely appr es s ed th e pe ri c h aet i a l l eaves completely entire obsoletely co s tat e .... Rh. filiformis llO 5

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6. 8. 11. 14. 111 With the margin of the perichaetial leaves provided with teeth one to each. 7 Rh. orizabanus The perichaetial leaves completely entire or the margin erosedenticulate .......................... 8 The branches at the apex attenuate, the branch leaves (apical) homomallous, the costa extended to the middle ..... 9 Rh. orthostegius The branches subclavate at the apex, thickened, the leaves heteromallous, the costa extended above the middle ...... 10 Rh. polycarpus The branches nearly straight, the branch leaves heteromallous, the processes indistinctly bordered .............. 12 Rh. brasiliensis The branches curved, the branch leaves homomallous, the processes distinctly hyaline-bordered. . . 13 Rh. secundus The branch leaves completely entire .............. 15 Rh. declinatus The branch leaves at the apex finely serrate .... 16 Rh. serrulatus

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11 2 Key to Species of Rhegmatodon sensu Broth e rus (1925)1 I. (A) Seta smooth A. (a) Branch leaves 2-2.5:1 1. (Aaa) Branch leaves oval a. (AaaI) Branch leaves laxly d appresse ; inn e r involucral leaves with serrated teeth Rh. schlotheimioides Spruce b. (AaaII) Branch leaves tightly appressed; inn e r involucral leaves with entire margins; obs c ur e ly costate Rh. filiformis Schimper Rh. pringlei Card. 2. (AaS) Branch leaves oblong oval a. (AaSI) Inner involucral leaves with indi v idual littl e teeth Rh. densus Schimp. b. (AaSII) Inner involucral leaves with e ntir e m argi ns or crenately denticulate 1) (AaSIIl) Branches acuminate; th e up pe r b ranc h l eave s bent to one side (homomallous); costa st opp in g at midleaf Rh. orthostegius Mont. 2) (AaSII2) Bran c h e s c lub-sha pe d, thi c k ened; bra n c h l e av e s not bent to on e s id e t co st a co n t inu ing pas t midleaf Rh. polycarpus (Griff.) Mitt. B. (Ab) Branch leaves about 1.8-1 1. (Aba) Branches nearly straight; bran c h l e av es n o t b e nt to one side Rh. crassirameus C ard. The charact e rs in parenth e s e s w e r e originally u sed by Brot h e rus in A. Engler and K. Prantl, "Die naturlich e n Pfl a nz e nfa m il ie n." Ed 2 1 925 11:298-99. I have supplied my own outlin e f o rm in t h e in terest of c l ar i ty

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Rh. brasiliensis Lindb. 2. (Ab8) Branches cu r ved; branch l e ave s b e nt to on e sid e Rh. secundus Kiaer Rh. newtoni Broth. II. (B) Seta rough A. (Ba) Branch leaves with entire margin Rh. declinatus (Hook.) Brid B. (Bb) Branch leaves serrate to the tip Rh. serrulatus (Doz. et Molk.) 113

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114 Key to Species of Macrohymenium sensu Kiaer (1882) Macrohymenium gracillimum (C. Mull. in correspondence 1882), of Tahiti, with the cells of the leaves provided with alars round-quadrate, inflated, yellow, the rest is not know to me. 1. 3. 5. 7. 9. The Other Six Species The branch leaves wider (2.19) smaller ............ 2 M. rufum The branch leaves narrower (above 2.87) larger ......... 3 The branch leaves very long (3.91) ............... 4 M. strictum The branch leaves shorter (2.88-3.23) ............. 5 The perichaetial leaves denticulate ............. 6 M. acidodon The perichaetial leaves completely e ntire ........... 7 The branch leaves loosely imbricated, outspread ........ 8 M. nietneri The branch leaves densely imbricated erect-spreading or spreading. . . . 9 The branch leaves heteromallous ......... .10 M. laeve The branch leaves homomallous ........... .11 M. mulleri

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llS Key to Species of Macrohymenium sensu Brotherus (1925) Section I. Leidontium Broth. n. s e ct. slend e r p lant s P e ri s t o m e teeth smooth. The processes of the inner peristome broad, finely papillose, transparent. (6 species) A. The seta above set with very low, broad warts. Aa. The inner perichaetial leaves entire margined. Aaa. Slender plants; the branch leaves nearly 2:1 M. mitratum (Doz. and Molk.) Fleisch. [M. rufum (Reinw. and Hornsch.) C. M~ll.J Sumatra, Java, Queensl. Aab. Less slender plants; the branch leaves v e ry long, about 4:1 M. strictum Bryol. Jav., Born e o, Mindinao. Ab. The inner perichaetial leaves denticulat e : M. acidodon (Mont.) Doz. and Molk., Bourbon, Madagascar.; f. acutissima Besch. (Rhegmatodon madagassus Geh. nach Kiaer), Madagascar, Seychelles. B. The seta plainly rough; the inner perichaetial leaves entire margined. Ba. The dry branch leaves loosely appress e d, wh e n moist spreading: M. nietneri (C. Mull.) Mitt., C e ylon. Bb. The dry branch leaves densely appr e ss e d, wh e n w e t ere ct spreading. Section II. Bba. The branch leaves spreading on all sides: M. la e ve Thwait. and Mitten., Ceylon. Bbb. The branch leaves unilateral: M, mull e ri Doz. and Molk. (Fig. 745), Sumatra, Java, Born e o. Trachydontium Broth. n. teeth papillose abov e narrow, through a thick species) s e ct. Robust plants. Th e p e ri s tom e Th e proc e ss e s of th e inn er p e ri s t o m e framework of papilla e opaqu e (1 M. sinense Th~r., Kweitschou.

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Anderson, L. E. opinion. 1954. ogists. LITERATURE CITED 1940. The generic concept. II. A survey of modern Bull. Torr. Bot. Club 67:363-369. Hoyer's solution as a rapid mounting medium for bryol The Bryologist 57:242-244. 1963. Modern species concepts: mosses. The Bryologist 66: 107-109. 1974a. Bryology 1947-1972. In 25 years of botany. Ann. Missouri Bot. Gard. 61:56-85. 1974b. Taxonomy and evolution of bryophytes. J. Hattori. Bot. Lab. 38:1-11. Bartlett, H. H. 1940. History of the generic concept in botany. Bull. Torr. Bot. Club 67:349-362. Bescherelle, E. 1871. Prodromus bryologiae mexicana e OU e num e ration d~s mousses du Mexique avec description des esp~ces nouvelles. Mem. Soc. Nat. Sci. Nat. Cherbourg 16:145-256. 1872. Musci. In E. Fournier, Mexicanas plantas nuper a collectoribus expeditionis scientificae allatis aut longis ab annis in herbario musei parisiensis depositas. 1:7-58. Paris. 1880. Florule bryologique de la Reunion et des autr e s il e s Austro-Africaines de l'Oc~an Indien 2. Ann. Sc. Nat. Bot. VI, 10:233-332. Bosch, R. B. van den, & van der Sande LaCoste. 1865. In Dozy and Molkenboer. Bryologia Javanica 2:114. Bridel, S. E. 1819. Muscologia recentiorum supplem e ntum. IV. (Mantissa Muscorum). 1826-27. Bryologia Universa, seu systematica ad novum m e thodum dispositio, historia et descriptio omnium muscorum frondosorum hucusque cognitorum cum synonymia ex auctoribus probatissimi s I. 1-860. Leipzig. Briggs, D. 1965. Experimental taxonomy of som e British sp e ci e s o f the genus Dicranum. New Phytol. 64:366-386. 116

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117 Brotherus, V. F. 1895. Laflore bryologique du Br e 's1.. B h 1. ang Till K. Sv. Vet.-Akad. Handl. Band 21. Afd. 3(3):64. 1897. Musci Africani. II. Bot. Jahrb. 24:281-282. 1924. Musci novi sinenses. Sitz b k ungs er. A. Wis s Wi e n. Math. Nat. Kl. Abt. 1, 133:578-579. 1924-25. Musici. In Engler and Prantl. Die naturlich e n pflanzenfamilien. 2 ed. Leipzig. 1929. Symbol. Sin. 4:94. Bruch, P., Schimper, W. P., & T. Gumbel. 1836-1855. Bryologia Europa e a seu genera muscorum Europaeorum monographice illustrata, 6 vols Stuttgart. Cardot, J. 1910. Diagnoses preliminaires de mouss e s M e xicaines. R e vu e Bryologique et Lichenologique. Caen; Paris 37:49-59. Cronquist, A., Holmgren, Arthur & Noel, & J. Rev e al. 1972. Int e mountain flora: vascular plants of the intermountain W e st. Vol. 1. New York Botanical Gardens. Crum, H. 1951. The Appalachian-Ozarkian el e ment in the moss flora of Mexico with a check-list of all known Mexican moss e s. Diss e rta tion. University of Michigan. 1973. Mosses of the Great Lak e s for e st. C ontribution s fro m the University of Michigan Herbarium, 10:1-104. Dillenius, J. J. 1741. Historia Muscorum. Oxford. Dixon, H. N. 1932. Classification of mosses. In V e rdoorn, Manual o f bryology. Chapter 14, pp. 397-412. Dozy, F., & J. H. Molkenboer. 1844. Muscorum frondos o rum no v a e spec i es ex archipelago indico et japonia. Ann. Sc. Nat. III, se r II 1845-1854. Mus c i frondosi in e diti ar c hip e l ag i In d i c i s i ve d e scriptio et adumbratio muscorum frondo so rum in in s uli s Java Born e o, Sumatra, Celeb e s, Amboin e a ... LugduniBa t a v or um, I L W Hazenberg & Soc. 1861-1870. Descriptio muscorum frondosorum ar c hi pe l a gi indi c i. Bryologia Javanica 2. 1 K P tl 1909 Die naturlichen pflanz e nf a mili e n. Eng er, A., & ran Vol. I & II. Leipzig. 1925. Ib. Bands 10 & 11. Fleischer, M. 1904-1923. Die Musci d e r flora van Buit e nz o r g 4 vo l s. Duncker & Humblot.

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118 Florschutz, P., Introduction to Hedwig's 'Speci es Muscorum', reprint of 1960 by H. R. Engelmann (J. Cramer) and Wheldon & Wesley, LTD. New York. Hafner Pub. Gangulee, H. C. Fasc. 1. 1969. Mosses of eastern India and adjacent regions. Calcutta, by the author. Gardner, Eldon. 1965. History of biology. 2 ed B Pub urgess Co., Minneapolis, Minn. Geheeb, A. 1882. Geh. Reliquiis Rutenbergianis III. Bot. Laubmoose. Abhandl. v. Naturw. Ver. zu Brem.:211. Griffith, W. 1838. Muscologia itineris Assamici. In Calcutta J. Nat. Hist. 3:275-276. 1843. 1849. Posthumous papers bequeathed to the honorable East India Co., and printed by the order of the governm e nt of Bengal. Icones Plantarum Asiaticarum. Arranged by J. McClelland. Part 2, on the higher cryptogamous plants. Calcutta. Hampe, E. 1839. Relation uber die vaon dem Reisenden C. Beyrich auf seiner letzten Reise in Nordamerika gesammelten Laubmoose. In Linnaea 13:39-48. 1844. Icones muscorum novorum vel minus cognitorum ... Part 2, Sumptibus Henry & Cohen, Bonnae. Hedwig, J. 1787-97. Descriptio et adumbratio microscopico-analytica muscorum frondosorum (Stirpium Crytogamicarum). 4 vols. Leipzig. 1782. Fundamenta historiae naturalis muscorum frondosorum. 1801. Species muscorum frondosorum. 1-352. Leipzig. Herzog, T. 1926. Geographie der Moose. Jena. 1932. Geographie. In Verdoorn, Manual of bryology. Chapter 10, pp. 273-296. & A. Noguchi. 1955. Beitrag zur k enntnis bryophytenflora von ----FormOsa und den Benachbarten inseln Bat e l Tobago und Kwashyoto. J. Hatt. Bot. Lab 14:29-70. H 1 P K & W K k 19 74. Index H e rbariorum: Part I o mgren, .. eu en. The herbaria of the world. Oosthoek, Scheltema & Holkema, Emmalaan 27, Utrecht, Netherlands. Hooker, W. J. 1808. Musci Nepalenses; or descriptions of several new mosses from Nepal. Trans. Linn. Soc. Land. 9:307-322. & W. Wilson. 1856. In Gray, Man. Bot. N. U. States, 2 ed --Ireland, R. 1971. Moss pseudoparaphyllia. The Bryologist 74:31 2 330

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Jaeger, A., & F. Sauerbeck. systematice disposita orbis terrarum. Ber. 18 7 0-80. G e nera e t species muscorum seu adumbratio florae mus cor um totius Th~tigk: St. Gallischen Naturw. Ges. 119 Kiaer, F. C. 1882. Genera muscorum Macrohym e nium et Rhegmatodon revisa specieque nova aucta exposuit. Christiana Vid e n sk-Selsk Forhandl. 1882(24) :l-53. Limpricht, K. G. 1890-1904. Die Laubmoose Deutschlands, Oesterreichs und Schweiz. 2 Abt. 2:740. Linnaeus, C. 1753. Species Plantarum 2. Lorentz, P. G. 1867-1868. Grundlinien zu einer verglochend e n Anatomie der Laubmoose. Jahrb. Wiss. Bot. 6:363-466. Margadant, W. D. in Musci. 1959. Typification and conservation of generic nam es Act. Bot. Neerl. 8:275. 1968. Early bryological literatur e. Hunt Botanical Library, Pittsburg, Penn. Mitten, W. 1859. Musci Indiae Orientalis. J. Proc. Linn. Soc. Bot. Suppl. 1:1-171. 1868. A list of mosses collected by the Rev. Thomas Powell in the Samoa or Navigator's Islands. J. Linn. Soc. Bot. 10:166-195. 1869. Musci Austro-Americani. Enumeratio muscorum omnium austro-americanorum auctori husque cognitorum. J. Linn. Soc. Bot. 12:1-650. 1873. New species of Musci collected in Ceylon by Dr. Thw a it es. J. Linn. Soc. Bot. 13:293-326. Montagne, c. 1842. Cryptogamae Nilgherienses. Ann. Sc. Nat. Bot. ser. 2, 17:248-249. 1845. Cinquieme et sixieme centuries de plantes cellulaires exotiques nouvelles. In Ann. Sc. Nat. Bot. 4:96. Muller, C. 1847. De muscis nonnullis novis vel minus cognitis exoticis. Bot. Zeit. 5: col. 801-806, 825-830. 1849-1851. Synopsis muscorum frondosorum omnium hucusque cognitorum. 2 vols. Berlin. 1862. Bot. Zeit. 20:374. 1869. Linnaea 36:21. 1891. Nuov. Giorn. Bot. Ital. 23:601.

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120 Noguchi, A. 1972. Musci Japonici IX. The Leskeaceae. Lab 36:499-529 J. Ilatt. Bot. Paris, E. G. 1894-1900. Index bryologicaus sive enumeratio muscorum hucusque cognitorum. 2 ed., 5 vols. Paris. Philibert, H. 1898. Etudes sur le peristome. Le peristome interne; ses variations. Rev. Bryol. 15:56-60, 65-69. Reimers, H. 1929. Beitrage zur bryophytenflora Neuguineas. Hedwigia 69:129-130. 1931. Ib. 71:62. Reinwardt, C. G. C., & C. F. Hornschuch. 1826. Musci frondosi Javanici. Nov. Act. Acad. Caes. Leop. XIV, Suppl., pp. 716-717. Richards, P. W. 1967. Ecology. In Verdoorn, Manual of bryology. Chapter 13, pp. 367-395. Schimper, W. P. 1855. Corollarium bryologiae Europaeae, conspectum diagnosticam familiarum, generum et specierum, adnotationes novas atque emandations complectens. 1-140. 1872. In Bescherelle, Mem. Soc. Sc. Nat. Cherbourg 16. Schwaegrichen, C. F. 1828. Species Muscorum supplementum. III. 1. Seki, T. 1968. A revision of the family Sematophyllaceae of Japan with special reference to a statistical demarcation of the family. J. Sc. Hiro. U. Series B, 2 (Botany) 12:1-80. Stearn, W. T. 1967. Botanical Latin. 566 pp. London. Thomas Nelson Lts. Steere,W. C. 1947. A consideration of the concept of genus in Musci. The Bryologist 50:247-258. Streeter, D. T. 1970. Bryophyte ecology Sc. Prog. Oxf. 58:419-434. _, Theriot, I. 1909. Bull. Acad. Georg. Bot. 19:20. Verdoorn, F. 193 2 Manual of bryology. The Hague. Martinus Nijhoff. Reprinted by A. Asher & Co., Amsterdam, 1967. d d l L 194 7. Bull. Jard. Bot. Bruxelles 18:182-183. Var, P. e a, & eroy. Watanabe, R. 1972. adjacent areas. A revision of the family Thuidiaceae in Japan and J. Hatt. Bot. Lab 36:171-320. Sk 1) 1924. The origins of con Wegener, A. (Translated by J. G. A. er tinents and oceans. London.

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121 Wijk, R. van der, W. D. Margadant, & P.A. Florschutz. 19 6 7. Ind ex Muscorum. 5 vols. Utrecht. Zander, R.H. 1972. Revision of the genus Leptodontium (Musci) in the New World. The Bryologist 75:213280

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BIOGRAPHICAL SKETCH David Alvin Eakin was born February 21, 1945, in Bridgeport, Connecticut. He was graduated from Seneca High School, Louisville, Kentucky in June, 1962. He received the Bachelor of Arts with a major in Biology from the University of Louisville in Louisville, Kentucky, in January, 1967. He worked as a graduate teaching assist ant at the University of Louisville, receiving his Master of Science in Biology in December, 1972. From September, 1971, until the present time he has pursued his work toward the degree of Doctor of Philosophy. David Alvin Eakin is married to the former Lois Darlene Walker, and is the father of a daughter, Heather Rebecca, and a son, Jonath a n David. He is a member of the American Bryological and Lichenological Society of America and the Association of Southeastern Biologists. 122

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I certify that I have read this study and that 1 n m y opinion it conforms to a~ceptable standards of scholarly presentation and is fully adequate, _in scope and quality, as a dissertation for the de ree of Doctor of Philosophy. g I certify that I have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality, as a dissertation for the degree of Doctor of Philosophy. ames W. Kimbrough rofessor of Botany I certify that I have read this study and that in my opinion it conforms to acceptable standards of scholarly pr e sentation and is fully adequate, in scope and quality, as a dissertation for the degree of Doctor of Philosophy. Indra K. Vasil Professor of Botany I certify that I have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality, as a dissertation for the degree of Doctor of Philosophy. athan Reiskind Associate Professor of Zoology

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I certify that I have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality, as a dissertation for the degree of Doctor of Philosophy. Associate ProfessOJ'." of Botany This dissertation was submitted to the Graduate Faculty of the College of Agriculture and to the Graduate Council, and was accepted as partial fulfillment of the requirements for the degree of Doctor of Philosophy. August, 1976 College of Agri u ture Dean, Graduate School