A REVISION of the MOSS FAMILY METEORIACEAE in AUSTRALIA with more detailed studies

on the genus Papillaria (C. Muell.) Lor.

by HEINAR STREIMANN

M. Sc. Thesis 1988

UNIVERSITY

of

NEW SOUTH WALES STATEMENT

ACCORDING TO ARTICLE 29 OF THE INTERNATIONAL CODE OF BOTANICAL NOMENCLATURE, THIS THESIS DOES NOT QUALIFY AS AN EFFECTIVE AND VALID PUBLICATION. THEREFORE, THE DESCRIPTION OF THE NEW TAXON, THE NEW COMBINATION AND THE NEW SYNONYMIES CONTAINED HEREIN ARE NOT YET VALIDLY PUBLISHED. ii

DECLARATION

I HEREBY DECLARE THAT THIS SUBMISSION IS MY OWN WORK AND THAT TO THE BEST OF MY KNOWLEDGE AND BELIEF, IT CONTAINS NO MATERIAL PREVIOUSLY PUBLISHED OR WRITTEN BY ANOTHER PERSON NOR MATERIAL WHICH TO A SUBSTANTIAL EXTENT HAS BEEN ACCEPTED FOR THE AWARD OF ANY OTHER DEGREE OR DIPLOMA OF A UNIVERSITY OR ANY OTHER INSTITUTE OF HIGHER LEARNING, EXCEPT WHERE DUE ACKNOWLEDGEMENT IS MADE IN THE TEXT.

Heinar Streimann

2 December 1988 iii

CONTENTS

STATEMENT 1 DECLARATION 11 LIST OF TABLES VI LIST OF FIGURES Vll ABSTRACT 1 AIMS OF THE STUDY 2 MATERIALS AND METHODS 5

SECTION I

INTRODUCTION TO THE FAMILY METEORIACEAE 12 Prior taxonomic concepts 12 Cytology 17 Ecology 17 Distribution and phytogeography 19

SECTION II

STUDIES ON THE GENUS PAPILLARIA 24 Historical outline 24 DISfRIBUTION OF p APILIARIA 24 HISfORY OF THE GENUS 25 rnxoN's 'sruDrns' 1927 27 AUSfRALIAN SPECIES 28 NORFOLK ISIAND SPECIES 35 EARLY COMPARISON OF AUSfRALIAN TAXA 36 Taxonomic relationships in Australia 40 CIADISTIC ANALYSIS 40 REIATIONSHIP BEIWEEN PAPILIARIA AND AfETEORIUM 43 Morphological studies 45 GAMETOPHYTE 45 SPOROPHYTE 64

CYTOLOGY 66 ASEXUAL REPRODUCTION 66 Contents iv

Ecology 68 Distribution 72

SECTION III

STUDIES ON OTHER GENERA 76 Introduction 76 History and distribution 76 AEROBRYOPSIS 76 BARBELIA 78 FLORIBUNDARIA 82 MEIEORIOPSIS 84 MEIEORIUM 86 WEYMOUTI-IIA 88

SECTION IV

TAXONOMIC TREATMENT OF THE FAMILY METEORIACEAE 92 Introduction 92 The family Meteoriaceae 92 TIIE GENUSPAPILLARIA 96 1 - P. amblyacis 100 2-P. crocea 114 3 - P. flavolimbata 131 4 - P. jlexicaulis 146 5 - P. intricata -158 6 - P. leuconeura 165 7 - P. zelojlexicaulis 173 UNKNOWN p APILIARIA SPECIES 182 TIIE GENUsAEROBRYOPSIS 183 1 - A. wal/ichii 185 nrn GENusBARBELLA 191 1 - B. cubensis 193 2-B. nitens 199 Contents V

TIIE GENUS FLORIBUNDARIA 207 1 - F. floribunda 209 2 - F. pseudofloribunda 213 3 - F. walkeri 216 TIIE GENusMETEORIOPSIS 220 1 - M. rec/inata 222 2 - M. robustu/a 226 TIIE GENusMETEORIUM 230 1 - M. buchananii 232 TIIE GENUS WEYMOUTH/A 238 1 - W. cochlearifolia 240 2 - W. mollis 248

EXCLUDED SPECIES 253

CONCLUSION 255

ADDENDUM 257

ACKNOWLEDGEMENTS 258

REFERENCES 260

TAXONOMIC INDEX 271 List of tables vi

LIST OF TABLES

1 Australian taxa of Meteoriaceae reported prior to this revision 7 2 Genera and their distribution prior to the present study 14 3 Reported chromosome numbers in Meteoriaceae 16 4 Meteoriaceae species and their reported habitats 18 5 Number of endemic species of Meteoriaceae m regions recognized in Wijk et al. (1959) 20 6 Worldwide distribution of Papi/laria species before this study in the regions recognized by Wijk et al. (1959) 24 7 Cladistic character set data 41 8 Meteoriaceae taxa reported for Australia prior to this revision and the present status 256 List of Figures vii

LIST OF FIGURES

1 Leaf dimensions and location of leaf sections illustrated 7 2 Distribution of genera and species prior to this revision 13 3 Rainfall and rain forest types in eastern Australia 22 4 Distribution of Papillaria species prior to this revision 25 5 Cladogram of Australian Papillaria and outgroup species 42 6 Branch cross-sections of Papillaria 47 7 Stem cross-sections of Papillaria 48 8 Branch leaf length/width in Papillaria amblyacis and P. crocea 49 9 Branch leaf length/width in Papillaria zeloflexicaulis sp. nov., P. flexicaulis and P. flavolimbata 51 10 Auricular cells of Papillaria flexicaulis and P. crocea 54 11 Cross-sections of Papillaria branch leaves 56 12 Rhizoids of Papillaria 57 13 Antheridia of Papillaria 60 14 Archegonia of Papillaria 63 15 Calyptrae and operculae of Papillaria 64 16 Distribution of Aerobryopsis species before this revision 77 17 Distribution of Barbella species before this revision 79 18 Distribution of Floribundaria species before this revision 83 19 Distribution of Meteoriopsis species before this revision 85 20 Distribution of Meteorium species before this revision 87 21 Distribution of Weymouthia species before this revision 90 22 Papillaria amblyacis branch leaves 101 23 Papillaria amblyacis branch leaf cells 103 24 Papil/aria amblyacis branches, sporophytes, leaf surface and archegonia 104 25 Papillaria amblyacis on tree branches 107 26 Distribution of Papil/aria amblyacis 108 27 Upper leaf cells near mid-leaf, Papillaria amblyacis and P. leuconeura 111 28 Papillaria crocea branch leaves and perichaetial leaf 116 29 Papil/aria crocea branch leaf cells 117 30 Papil/aria crocea branches, leaf surface, sporophytes and peristome 118 List of Figures viii

31 Papillaria crocea in cool temperate rain forest 120 32 Distribution of Papillaria crocea 122 33 Distribution of Papillaria crocea on Norfolk Island 124 34 Papillaria f/avolimbata branch leaves 133 35 Papillaria flavolimbata branch leaf cells and apices of young branch leaves 135 36 Papillaria f/avolimbata branches, leaf surface, sporophytes, peristome and spore 136 37 Papillaria flavolimbata on shrub branches 138 38 Distribution of Papillaria flavolimbata 140 39 Papillaria flexicaulis branch leaves, perigonial leaf and leaf cells 149 40 Papillaria f/exicau/is branches with perigonia, leaf cells and antheridium 150 41 Distribution of Papillaria flexicaulis 153 42 Papillaria intricata branches 158 43 Papillaria intricata leaves and leaf cells 159 44 Distribution of Papillaria intricata 162 45 Papillaria leuconeura branch and branch tip 165 46 Papillaria /euconeura leaves and leaf cells 167 47 Distribution of Papillaria /euconeura 170 48 Papillaria ze/of/exicaulis branches and capsules 173 49 Papillaria zeloflexicaulis leaves and leaf cells 175 so Papillaria ze/oflexicaulis branch, base of branch, leaf, capsule surfaces, capsule, peristome, archegonia and spore 176 51 Distribution of Papillaria zeloflexicaulis 179 52 Distribution of Aerob,yopsis wallichii 188 53 Distribution of Barbella cubensis 196 54 Distribution of Barbella nitens 203 SS Distribution of F/oribundaria floribunda 211 56 Distribution of F/oribundaria pseudofloribunda 215 57 Distribution of F/oribundaria wa/keri 217 58 Distribution of Meteoriopsis reclinata 225 59 Distribution of Meteoriopsis robustu/a 228 60 Distribution of Meteorium buchananii 234 61 Distribution of Weymouthia cochlearifolia 243 62 Distribution of Weymouthia mollis 251 1

ABSTRACT

A taxonomic revision undertaken of the family Meteoriaceae in Australia recognizes 7 genera with 17 species from the previously accepted 9 genera and 26 species. A total of 7 species from the family are placed into synonymy. Chrysocladium phaeum (Mitt.) Fleisch. and Pilotrichella conferta Ren. & Card. do not belong to the Australian moss flora while Barbella perpinnata (Broth.) Broth. is referred to Brachythecium in the Brachytheciaceae. Papillaria ze/oflexicaulis is described as a new species while P. intricata (Mitt.) C. Muell. & Broth. and P. leuconeura (C. Muell.) Jaeg. are now recognized as occurring in Australia. P. flexicaulis (Wils.) Jaeg. is added to the Norfolk Island moss flora while P. flavolimbata (C. Muell. & Hpe) Jaeg. is deleted and P. amblyacis (C. Muell.) Jaeg. is newly reported for Lord Howe Island. F/oribundaria robustu/a Broth. & Watts is transferred to Meteoriopsis robustula (Broth. & Watts) Streim., but it may belong in the highly variable M. reclinata (C. Muell.) Fleisch. ex Broth.

Papillaria species are treated in greater detail and illustrated because no recent revisions are available. Delineation of some species was uncertain as they were described on inadequate material. A cladistic analysis of Papillaria is presented which shows good resolution for five species.

Keys for the identification of genera and for species within the genera are provided. Lectotypes for six Papillaria and one Barbella species were chosen. For each accepted species synonymy, description, diagnostic characters, ecology, habit, distribution, notes and selected specimens examined are given.

Problems still remain in F/oribundaria and Meteoriopsis. Also the position of Barbella nitens (Hook.f. & Wils.) Nog. needs further investigation. A global treatment with peristome and cladistic studies could resolve these problems. Aims of the Study 2

AIMS OF THE STUDY

This study was undertaken to carry out a taxonomic revision of the family

Meteoriaceae Kindb. and in particular the genus Papillaria ( C. Muell.) Lor. in Australia and the Australian external territory of Norfolk Island. Very few taxonomic revisions have been made on Australian mosses and the family Meteoriaceae is no exception. Scott & Stone (1976) briefly outlined the family for Australia and enumerated 23 species in 9 genera. The greatest concentration of species is in Queensland (Table 1). Only one species is mentioned for Floribundaria although Wijk et al. (1967) credits Australia with three, and Tan (1984) added another one. However, there are numerous reports in which the authors discuss identified collections of Meteoriaceae including Papillaria and other genera together with their distribution, but little if any, detail concerning the taxonomy or particular problems is given. The major difficulty has been that the morphological characters generally used are vegetative features, because sporophytes are reported as rare. Scott & Stone (1976) suggested that capsules are rare because 'the whole sporophyte tends to drop out of the vaginula when old'.

The present study has some limitations since some of the taxa studied have close relatives in localities such as New Guinea, New Caledonia and other South Pacific Islands that are geographically close to Australia. It has not been possible to examine all of these here. Some of the taxa recognized here may eventually require additional synonymy when a wider geographical study becomes possible. In the absence of any published worldwide familial revision, a limited revision for a local area can, however, contribute considerably to the assessment of the family. Aims of the Study 3

Taxon Australian Distribution

Aerobryopsis wallichii (Brid.) OLD Fleisch. Barbella cubensis (Mitt.) Broth. OLD, NSW, LHI B. nitens (Hook.f. & Wils.) Nog. OLD, NSW, VIC, TAS * B. perpinnata (Broth.) Broth. OLD Chrysocladium phaeum (Mitt.) AUST Fleisch. Floribundaria floribunda (Dozy & OLD Molk.) Fleisch. F. pseudofloribunda Fleisch. OLD * F. robustula Broth. & Watts OLD F. walkeri (Ren. & Card.) Broth. OLD Meteoriopsis reclinata (C. Muell.) OLD Fleisch. ex Broth. * Meteorium baileyi (Broth.) Broth. OLD M. buchananii (Brid.) Broth. OLD * M. compressum Mitt. OLD Papillaria amblyacis (C. Muell.) OLD,NSW Jaeg. P. crocea (Hpe) Jaeg. OLD, NSW, VIC, TAS, NI * P. eavesiana (Hpe) Jaeg. OLD P. flavolimbata (C. Muell. & Hpe) OLD, NSW, VIC, TAS, NI Jaeg. P. flexicaulis (Wils.) Jaeg. OLD, NSW, TAS * P. fulva (Mitt.) Jaeg. VIC * P. reginae (Hpe) Jaeg. OLD P. squamata (Hpe) Hpe E.AUST Pilotrichella conferta Ren. & Card. AUST * P. dimorpha (C. Muell.) Jaeg. NSW * P. recurvula C. Muell. AUST Weymouthia coch/earifolia VIC, TAS (Schwaegr.) Dix. W. c. var. billardieri (Hpe) Dix. VIC, TAS W. mollis (Hedw.) Broth. LHI, VIC, TAS

* Endemic

Table 1. Australian taxa of Meteoriaceae reported prior to this revision. Aims of the Study 4

Taxonomic problems requiring detailed attention in this investigation are the number of published names that may refer to the same taxon. Previously 8 species of Papillaria have been reported together with 8 other genera comprising 18 species in the Meteoriaceae in Australia. It is envisaged that it will be necessary to reduce to synonymy a number of species thought to be endemic, and some species may need to be transferred to other genera. As many names were based on inadequate collections, or because some European bryologists, especially C. Mueller, regarded most Australian collections as new to science and described them as such, the number of endemic species has been inflated. Amongst the large number of unnamed collections from tropical Australia it is anticipated that there will be some interesting new records. Apart from Papillaria, no members of this family have been reported from Norfolk Island. Published records for Lord Howe Island (Ramsay 1984) list two genera,

Barbella and Weymouthia, each with one species. Materials and Methods 5

MATERIALS AND METHODS

The study was carried out at the Australian National Botanic Gardens and at the University of New South Wales. Loans of specimens including types were obtained from Australian and overseas herbaria. A total of 2559 specimens have been examined in detail.

The taxonomic decisions made · in this revision were based on the following numbers of Australian collections studied for each of the species accepted:

Aerob,yopsis wallichii 143 Barbella cubensis 118 B. nitens 203 Floribundariafloribunda 5 F. pseudofloribunda 5 F. walkeri 2 Meteoriopsis reclinata 36 M. robustu/a• 4 Meteorium buchananii 130 Papillaria amblyacis 409 P. crocea 647 P. flavolimbata 353 P. flexicaulis 256 P. intricata 2 P. leuconeura 14 P. zeloflexicaulis•• 47 Weymouthia cochlearifolia 85 W. mollis 101 * New combination made in these studies. * * New species made in these studies. These collections were obtained on loan from the following herbaria: ALTA, BM, BRI, C, CANB, COLO, FH, GRO, H, HO, JE, L, LD, MEL, MELU, Materials and Methods 6

NSW, NY, UBC, UNSW, W, WELT as well as the CBG collections. These acronyms for herbaria are based on Vitt et al. (1985).

Full label data were recorded on cards for every specimen examined. Permanent slides (including types) were made of leaves, sporophytes etc. for a representative selection of all the Australian taxa using Hoyer's solution (Anderson 1954). With the aid of a compound microscope (Nikon) these slides were sorted into groups with apparently similar leaf structure. Any problems, or unusual collections, were set aside for further study. Once the groups were delineated problem slides were re-examined and placed into the appropriate groups or set aside as new taxa.

While the slides were being made an information sheet was filled out noting the features of the moss both dry and wet. A detailed study of morphological features from each slide enabled additional data to be entered onto the information sheet. The length and width measurements were taken as indicated in Fig. 1 together with the following additional data:

Length and width of cells near the apex and in the median section of the leaf;

Maximum length of cells at the base near the costa Guxtacostal);

Shape and size of cells in the auricles;

Costa length and characteristics;

Denticulation of the margin.

Any other unusual features were noted, including thickness of cell walls, cell shape and alignment, papillosity etc.

Sporophytic characters, obtained from specimens and permanent slides, were checked against measurements taken from temporary aqueous slides. Materials and Methods 7

Hand sections of the stems and branches of Papillaria species were made using a razor blade. For leaf and seta, sections 14um thick were prepared using a freezing microtome.

Fig.1. Leaf dimensions and location of leaf sections illustrated (see p. 9).

The literature search for each taxon provided the historical background for the taxonomic concepts for each species accepted at present. In addition a profile of general morphology, colour, leaf (stem and branch) shape, size, margin, cells, costa etc., rhizoids, perigonia, perichaetia, sporophytes, chromosomes, derivation of botanical names and illustrations was developed. The information was generally patchy and incomplete. Comments made by authors concerning problems relating to Australian taxa and other statements were checked for accuracy against Australian collections. Information on Australian distribution and ecology were obtained from label data and from observations during field studies. Extra Australian distribution for the species was mainly from Noguchi (1976 & 1985). Materials and Methods 8

Scanning electron microscopy was carried out at the University of New South Wales in Sydney (Cambridge Stereoscan 54-10) and at the Australian National University in Canberra (Cambridge Stereoscan 360), using Ilford FP4- 125 ASA film. Specimens were prepared for scanning electron microscopy by the following techniques. Those photographed at the University of New South Wales were: 1) rehydrated in distilled water for about 30 minutes, 2) fixed in 2% of aqueous osmium tetroxide for two hours at room at room temperature and then thoroughly rinsed in distilled water, 3) dehydrated in ethanol series 10-100% in 10% steps, 4) followed by 3 series of ethanol acetate allowing 15 minutes per step and then critical point dried. The material was then mounted on aluminium SEM stubs using double tape and polaron and sputter coated with a thin layer of gold. Those at the Australian National University were set in nail polish on SEM stubs which were then placed in a Dynovac evaporative coater which gave a gold coating of 100 Aongstroem. The majority of the SEM illustrations were by the latter method. Black and white photographs of the leaf and leaf cells were taken from permanent slides with a Leitz Orthoplan and compound microscopes using phase contrast, often with a blue filter. Kodak Technical Pan-2415-50 ASA and Agfa Pan 25 ASA films were used.

All Papi/laria type collections were photographed in black and white or as colour slides at the Australian National Botanic Gardens.

Interspecific relationships in Papillaria were studied with the aid of the cladistics program PAUP, version 2.4.1, Illinois Natural History Survey. This program relies on the method in which branch and bound algorithms are used to find all parsimonious trees.

Most illustrations were made from permanent slides with the aid of a Nikon compound microscope with a Nikon drawing tube attachment. Some Materials and Methods 9

illustrations were traced from black and white prints taken through a light microscope. Collections cited for illustrations are deposited in the following herbaria: BM - Cruttwell; CANB - Adams; CBG - Curnow, HS, Thor; MELU - Stone; NSW - Boorman, Constable, Forsyth, Goldsmid, Watts; NY - Norris. Fig. 1 shows the locations on the leaf of the sections illustrated (1 - tip, 2 - upper middle, 3 - median margin, 4 - median middle, 5 - auricle). Data analysis of leaf lenght/width was carried out and plotted as graphs but no statistical tests were carried out.

Generic descriptions and sporophytic characters for all species ( except Papillaria, Barbella nitens and Weymouthia) are modified from those given by Noguchi (1976), with slight modifications based on my observations. Additional data on spores were obtained from Fleischer (1908).

Final species descriptions are based both on herbarium collections and fresh material when available. Field studies paid special attention to recording accurately their habit and habitat.

Workable keys to genera and species are included. All synonymy relevant to Australian taxa are included with the descriptions, except for excluded species which are listed separately. The terms perichaetium and perigonium are used, as defined by Crum & Anderson (1981:1294), to refer to the female and male inflorescences respectively. The term incrassate is used to define the cell wall thickness.

Citations of sources for original descriptions were obtained from Wijk et al. (1967) and then checked against the protologue. Any citation errors detected were corrected. Abbreviations for periodicals are those r~commended by Lawrence et al. (1968). Materials and Methods 10

Type specimens of all Papillaria species (including Barbella nitens) were examined unless stated to the contrary. However for the remaining genera, only types of Australian taxa were examined. The basionym for each valid species is given. Papillaria was studied in greater detail because it has the most species and they had not been reviewed elsewhere. The remaining genera are widespread outside Australia, are better known, and have been studied by Noguchi (1976), and to a lesser extent by Manuel (1977a, 1977b), Norris & Koponen (1985), Tan (1984) and Wu (1985a,b, 1986a,b). The opinion of these authors was accepted for non-Australian types. The types of all Australian taxa were studied and their status is indicated. For the Australian taxa only synonyms relevant to Australia were cited. For full synonymy see Noguchi 1976.

A listing of all illustrations of the species occurring in Australia was aimed at. For Papillaria species it is hopefully complete and nearly complete for the remaining genera for the Asian-Pacific region. As some non-Papillaria genera are far ranging it was decided that a complete listing was not feasible for this study without time consuming searches in literature only available overseas.

Geissler & Greene (1982) bring together numerous papers on generic and specific concepts which illustrate the diversity of ideas. Horton believes in the relevance of geographic isolation as a factor in speciation, but states that populations which are functionally genetically isolated because of geographic separation cannot be presumed to be species, since they may interbreed if brought into contact. As moss genetics is poorly understood other characters have to be used to reflect the biology of mosses. lwatsuki reviews 'new' characters which are under genetic control and thus less variable than the traditional morphological characters which he lists. Zander takes the view that species should 'reflect repeatability in identification' with which I agree. He then poses several questions with comments on the practical aspects of dealing Materials and Methods 10a

with scientific names and species concepts. Further papers are presented by Zander, Koponen, Longton and others who review various characters and fields

as an aid for taxonomy. Therefore I feel a fixed species concept is too inflexible at present and a concept should be developed as one studies the various moss groups. I am in agreement with Horton's (1982) statement 'that the fundamental aim of the taxonomist should be nomenclatural stability'. Because of this and the wide variations found in the species of this family which can be due to the great geographical range, I have tended to be cautious in describing new species. After studying considerable numbers of Australian (and overseas) collections, I feel that a reasonably broad variability in leaf and cell shape and size can be tolerated within most species. Because judgements in taxonomy can be considered subjective and arbitrary, I have tried to base my judgements on characters which I considered to be obvious and constant and not on narrow vague differences. Diagnostic characters are in italics.

As sporophytes are not common in Papillaria, Weymouthia nor Barbella nitens, and were not observed at all for the remaining taxa, characters based on that structure are not used in the keys.

Distribution maps, with collections up to the end of 1987, were prepared with the aid of the Australia 1:250,000 map series gazetteer, Australian Publishing Service, Canberra, 1975. Where discrepancies in spelling occurred this publication was followed. Only localities with precise data were plotted on the distribution maps. The standard Australian National Mapping use of Mt and Mount was followed. The contraction Mt is used for a geographical feature while Mount designates a settlement. In the citation of locality data for specimens studied the following abbreviations or contractions were used: Materials and Methods 10b

Ck Creek N.P. National Park I. Island R. River L Lake Ra. Range LHI - Lord Howe Island Rd Road NI Norfolk Island S.F. State Forest Full surnames of collectors are recorded except for H. Streimann, which is cited asHS. 11

SECTION I

INTRODUCTION TO THE FAMILY METEORIACEAE 12

SECTION I INTRODUCTION TO THE FAMILY METEORIACEAE

PRIOR TAXONOMIC CONCEPTS

The family Meteoriaceae was raised to family rank from a sub-family of the Neckeraceae by Kindberg in 1897 (Crosby & Magill 1977). The first references to this as a family were by Fleischer (1908) and Brotherus (1925) after which the family became generally accepted. Further information on the classification prior to the creation of Meteoriaceae as a family is given under the genera.

Most familial classifications beginning with Fleischer have emphasized peristome features. These classifications have varied slightly in detail but Meteoriaceae were usually considered to be closely related to Pterobryaceae in the sub-order Leucodontineae between Fontinalineae and Neckerineae.

Buck & Vitt (1986) proposed a new system of familial classification for the pleurocarpous mosses based on both gametophytic and sporophytic characters. Peristome characters were still considered, but leaf cell shape was added to the analysis. They proposed that cell shapes evolved from hexagonal through rhombic to linear-rhombic. Their data analyses were influenced by cladistic theory but they did not follow it in the true Henningian sense, rather they constructed a phylogenetic tree. They termed this a 'pseudocladistic' approach.

In this new classification (Buck & Vitt 1986) the sub-order Leucodontineae is placed between the new sub-order Pterobryineae and Neckerineae while Fontinalineae is transferred to the order Hypnales. In their system the family Meteoriaceae is considered closely related to the families, Introduction to the family Meterioriaceae 13

Trachypodaceae and Lepyrodontopsidaceae. This is only a slight re- arrangement of the families close to Meteoriaceae as outlined in the Fleischer system.

, .... Oc. 7/26

Oc.

Aust.2 3/7

Ant.

Fig. 2. Distribution of genera (above slash) and species (below slash) prior to this revision. (Present Australian distribution is 7/17.)

No fossil remains have been reported for this family. Reliable data are not available to suggest the age of the family or that of any other moss genera (Schofield 1985). In general fossil moss capsules are rare, only leaves or sporophyte parts being preserved. Parihar (1965) states that as most moss (Bryopsida) remains date from the Pleistocene period (1 million years ago) they can be considered only as sub-fossils. Introduction to the family Meterioriaceae 14

The Meteoriaceae 1s a family of pendulous mosses distributed predominantly in tropical and sub-tropical forests, but also extending into temperate forests. The family is composed of 18 genera and 293 species (Wijk et al. 1959-1969; Crosby 1977, 1979; Crosby & Bauer 1981, 1983, Tan 1984, Bauer & Crosby 1986, Noguchi 1986) with the greatest diversity appearing in the tropical regions, especially in the Americas (Fig. 2 & Table 2).

No. of Genus species Region of Main Occurrence

Aerobryidium 5 Asia Aerobryopsis 17 Asia, Africa Aerobryum* 2 Asia Ancistrodes 1 Chile Barbella 28 Asia Chrysoc/adium 3 Asia Cryphaeophilum 1 Patagonia, Chile Floribundaria 21 Africa, Asia Lindigia 8 South America Meteoridium 1 Central & South America Meteoriopsis 5 Asia Meteorium 28 Asia, Central & South America Papillaria 69 South America Pi/otriche/la 63 Africa Pseudobarbe/la 6 Taiwan Squamidium 26 South America Weymouthia 3 South America, Australasia Zelometeorium 5 South America

*Noguchi (1976:349) doubts whether this genus belongs in Meteoriaceae.

Table 2. Genera and their distribution prior to the present study (refer also to Fig. 2). Introduction to the family Meterioriaceae 15

Sainsbury (1955a) recognized two genera and seven species m New

Zealand. One of the Papillaria species, P. nitens was transferred from Papillaria to Barbella nitens by Noguchi (1985). All New Zealand species are also common in eastern Australia. In Noguchi's (1976) treatment of the Asian taxa in this family he dealt with seven genera and ten species which occur in Australia. The following ten species were considered as Australian endemics:

Barbella perpinnata (Broth.) Broth. Floribundaria robustu/a Broth. & Watts Meteorium baileyi (Broth.) Broth. M. compresswn Mitt. Papillaria eavesiana (Hpe) Jaeg. P. fulva (Mitt.) Jaeg. P. reginae (Hpe) Jaeg. P. squamata (Hpe) Hpe ex F. Muell. Pi/otrichel/a dimorpha (C. Muell.) Jaeg. P. recurvula C. Muell.

A further taxon, Pilotrichella conferta (Table 1), is listed by Wijk et al. (1969) as occurring in Australia and Southern Africa (Afr. 4).

In Gangulee's (1976) treatment of Meteoriaceae for eastern India, he stated that the major problems were the species Barbella enervis (Thwait. & Mitt.) Fleisch. and Meteorium miquelianum (C. Muell.) Fleisch. He regarded the former as a synonym of Barbella cubensis which occurs in Central America and the Caribbean and the latter to be only a form of the widespread Meteorium buchananii. He made the combination of M. buchananii fo. polytrichum (Dozy & Molk.) Gang. because the designated forma had two years priority over the specific name M. miquelianum.

Norris & Koponen (1985) have accepted Gangulee's arguments on these two problems, and the New Guinea M. miquelianum was regarded as M. buchananii without formalizing this by citing synonyms. They preferred to await the expected revision of the neo-tropical Meteoriaceae by M. Menzel of Berlin.

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(Broth.) (Broth.)

Muell.) Muell.)

(Thwait. (Thwait.

P. P.

Nog. Nog.

(Mitt.) (Mitt.)

Muell.) Muell.)

(Brid.) (Brid.)

Molk.) Molk.)

Nog. Nog.

[as [as

Jaeg. Jaeg.

(C. (C.

(C. (C.

Card.) Card.)

(Broth.) (Broth.)

& &

(Mitt.) (Mitt.)

& &

nipponica nipponica

chromosome chromosome

Muell. Muell.

(Card.) (Card.)

(Besch.) (Besch.)

Card.) Card.)

Card.) Card.)

& &

F. F.

Jaeg. Jaeg.

(C. (C.

attenuata attenuata

& &

retrorsum retrorsum

aurea aurea

[as [as

(Hook.) (Hook.)

(Dozy (Dozy

subdivergens subdivergens

reclinata reclinata

Horik. Horik.

(Ren. (Ren.

bucha11a11ii bucha11a11ii

amblyacis amblyacis

(Hpe) (Hpe)

Reported Reported

(Ren. (Ren.

Nog. Nog.

flagellifera flagellifera

helminthocladum helminthocladum

helminthocladulwn helminthocladulwn

kiushiuensis kiushiuensis

asperifolia asperifolia

3. 3.

P. P.

M. M.

M. M.

B. B.

subpolytrichum subpolytrichum

undulata undulata

levieri levieri

crocea crocea

flavolimbata flavolimbata

/uscescens /uscescens

stevensii stevensii

floribunda floribunda

Species Species

[as [as

[as [as

[as [as

(Nog.) (Nog.)

[as [as

Table Table

P. P.

Pseudobarbe/la Pseudobarbe/la

P. P.

P. P.

Papillaria Papillaria

P. P.

M. M.

Meteorium Meteorium

M. M.

Meteoriopsis Meteoriopsis

F. F.

Chrysocladium Chrysocladium

Floribundaria Floribundaria

B. B.

Barbella Barbella Aerobryopsis Aerobryopsis Introduction to the family Meterioriaceae 17

CYTOLOGY

Very little is known about the cytology of many of the genera in Meteoriaceae (Table 3). The majority of chromosome numbers reported are by Inoue (1965) and Ramsay (1967, 1974) as shown in Table 3, the only Australian counts being those of Ramsay (1967) for 3 species of Papillaria.

ECOLOGY

In Australia the Meteoriaceae are also regarded as a predominantly tropical family as is demonstrated by its pendant and ramicolous growth habit.

The marked exception being Weymoutlzia, which prefers the Nothofagus and cool temperate forests. Predominantly rain forest mosses of the tropics are

Aerob,yopsis, Floribundaria, Meteoriopsis, Meteorium, Papillaria intricata and P. leuconeura. Those confined to tropical and sub-tropical regions are Barbella cubensis and the new species reported here Papillaria zeloflexicaulis sp. nov. The remaining species extend from the tropical and sub-tropical rain forests to the cool temperate and Nothofagus forests (Table 4).

The altitudinal range varies, with most of the ram forest species occurring from sea level to 1220m. The exceptions being Meteoriopsis which very rarely has been reported below 360m, Papillan·a zeloflexicaulis which has not been found above 200m while the only collection of P. intricata was from the lowlands. At lower altitudes these mosses are found in the forest but at the higher altitudes, where more constant moisture is available, they survive in more exposed situations. lntroductio11 to the family Meterioriaceae 18

...., ...... , ., GI ., GI GI .. GI .c .. C 0 .. ID 0 0 "'C ... 0 ... C ...... , C ...... GI ID ... ,a ...... GI .. ., > ...... >, ... 0 ID ,a >, GI l .., .c ID ... GI ., .c Q, .. .. Q, "'GI ... 0 GI 1111 .0 ID ., I 0 > ., .. Q, ::,j ::, ::,1 .. GI GI E D'li .. .c.., 4,lj GI GI .. ... GI ID I . u ~ ... C 0 u .., ...., 1/) > >,I u ...... , OI 0 ... , ., .. ., C ... .Cl .. .. ID I GI C ... .., 0 ..,, 0 UI >, > > ID GI 0 01 0 "'GI ::,1 ...... 0 SPECIES a: 3: u ZI ll,, a: !all 0 a: E-

Aerobryopsis wallichii X X X X

Barbella cubensis X X I B. nitens X X X X I Floribundaria floribunda X

F. pseudofloribunda X

F. walkeri X Meteoriopsis reclinata X X Meteorium buchananii X X X X X X Papillaria amblyacis X X x· X X X X J -P. crocea X X x, Jx l X X I -P. flavolimbata X X X X ! X P. flexicaulis X l X X X X - l I J I I -P. intricata I X l l ! J -P. leuconeura X I 1 -P. zeloflexicaulis* X X I We~outhia I cochlearifolia J xl X l I I l J J l w. mollis I X XJ I I X I l - f I ·I I I *New species reported in these studies.

Table 4. Meteoriaceae species and their reported habitats.

Papillaria species are treated in greater detail under Ecology in section

1 1 SfUDIES ON PAPILLARIA • Introduction to the family Meterioriaceae 19

DISTRIBUTION AND PHYTOGEOGRAPHY

The Meteoriaceae is represented by genera in all continents ( except Antarctica) with only one endemic species, Barbella strongylensis Bott., in Europe. There are three monotypic genera (Ancistrodes, Cryphaeophilum and Meteoridium) all endemic to Central and South America.

Brazil, Paraguay, Guiana, Trinidad and Tobago, which are designated by Wijk et al. (1959) as 'America 5' (Am. 5) have the highest world concentration of species (Fig. 2 p. 13) and endemics (Table 5) with 42 species of which 22 belong to Papillaria. This is followed by central Africa and St Helena Island ('Africa 2' - Afr. 2) where 33 endemic species occur, with the genus Pilotrichella accounting for 20 species. The area of 'America 4' (Am. 4), which includes Columbia, Ecuador, Venezuela, Peru and Bolivia, has 22 endemic species, with the genus Squamidium containing 7 species.

In Australia (Austr. 1) 10 endemic species have been reported of which four belong to Papillaria. There are no endemic species in New Zealand, Norfolk Island or Lord Howe Island.

Thus the tropical regions show a high rate of endemism, while the non­ tropical areas have low levels of endemism (Table 5).

Species of the genera Chrysocladium, Meteoridium and Zelometeorium are widely distributed and are not endemic anywhere in any of the regions designated by Wijk et al. (1959). ~

~

s·

~

~

1·

~

s.

s

g·

~

~

~

;ii'

~

'is­

2

7

1

2

2

14

OC.

6

4

2

3

1

1

1

AUSTR.

3

1

4 1

2

1

4

12

1

1

1

1

6

2

3

13

AS.

3

3

1

2

1

2

2

1

15

2

1

4

9

5

1

1

1

16

AFR.

(1959).

al. 2

3

1 1

5

3

l

2 3

33

20

et

Wijk

7

1

1

1

2

1

1

1

6

NONE

in

NONE

5

8

3

7

5

42

22

7

4 3

3

2 1

4

22

AM.

regions recognized

in

6

3

l

3

3

3

5

3

1

2

16

Meteoriaceae

of

species

1 1 1 1

------NONE

EUR.

endemic

of

endemi~I

Number

5.

Table

Zelometeorium

Total:

Sguamidium

Weymouthia

Pilotrichella Pseudobarbella

Papillaria

Meteorium

Meteoriopsis

Lindigia Meteoridium

Floribundaria

Cryphaeophilum

Barbella

Chrysocladium

Ancistrodes

GENUS

Aerobryum

Aerobryopsis

Aerobryidium Introduction to the family Meterioriaceae 21

Genera most widely represented in South America are Papillaria, Meteorium, Lindigia, Squamidium and Zelometeorium. Pseudobarbella from southeast Asia has its greatest representation in Taiwan with 7 species of which

3 are endemic. Aeorobryopsis, Aerobryidium, Aerobryum, Barbella and Floribundaria show the greatest diversity in tropical Asia, although endemism in the latter genus is equally well developed in this region and in central Africa. In

Africa Pilotrichella is the largest genus together with Floribundaria and to a lesser extent Aerobryopsis. The small non-tropical genus of Weymouthia 1s widely distributed in Australasia and southern South America.

In Australia this family is restricted to eastern Australia where all genera occur east of the Great Dividing Range because of ample rainfall and the resulting rain forest (Fig. 3). In Tasmania three genera are represented while on Norfolk Island (Table 1 p. 7) only Papillaria occurs. The genera Aerobryopsis, Floribundaria, Meteoriopsis and Meteorium are restricted to Queensland. Weymouthia is rare in New South Wales, but quite common in Tasmania, as is Papillaria flavolimbata. Surprisingly Weymouthia cochlearifolia has been reported from western Victoria (Portland) where it was common in a town park when collected in September 1956. Of the two Barbella species, B. cubensis is restricted to Queensland and New South Wales including Lord Howe Island where it is common on the higher peaks, while B. nitens (previously Papillaria) ranges from Queensland to Tasmania. With the exception of Weymouthia this family has not been reported farther west than the Otway Ranges (143°26'E) in Victoria.

A detailed study of the distribution of Papillaria is discussed under

1 Bryogeography in the section 'sruoie.s oN PAPILLARIA • Introduction to the family Meterioriaceae 22

D Rain forests

-- Lsohyets (mm)

700 I I I I I I I NT I I I I I I I QUEENSLAND I I ------,I I I I I SOUTH AUSTRALIA /------I' I I Subtropical rain forest I I NEW SOUTH WALES I ,I___ \ I \..__ l I '\.. . I Temperate ram I forest.J- I VICTORIA ----()()0

1000 700 0 100 200 JOO 400 ~m

Fig. 3. Rainfall and rain forest types in eastern Australia. 23

SECTION II STUDIES ON THE GENUS PAPILLARIA 24

SECTION II STUDIES ON THE GENUS PAPILLARIA

HISTORICAL OUTLINE

Distribution of Papillaria

The genus Papillaria with 69 species and 11 varieties (Wijk et al. 1967, 1969, Noguchi 1976, Crosby 1979, Bauer & Crosby 1986) is the largest genus in the Meteoriaceae with its greatest centre of distribution around the Pacific basin. The greatest concentration of species is in the northeast of South America (Am. 5). The worldwide distribution of species is summarised in Table 6 and Fig. 4.

North America, excl. Mexico 1 Central America, incl. Mexico 11 South America 33 Europe 0 Africa 13 Asia (Soviet Asia - 0) 11 Australia 8 New Zealand 4 Oceania 10

Table 6. Worldwide distribution of Papillaria species before this study in the regions recognized by Wijk et al. (1959). Studies on the genus Papil/aria 2S

...... Oc. 7/10

Oc.

Aust.2 0/4

Fig. 4. Distribution of Papillaria species prior to this revision (number of endemics above slash, total number of species below; present Australian distribution 1/7.)

History of the genus

Problems with the nomenclature relevant to Papillaria has been confused. Initially C. Mueller (1850) published Papillaria as a sub-section of the section Pseudopilotrichum in the genus Neckera Hedw., characterized by the 'leaves arranged in indistinct series' and having 'papillose cells'.

Recently Isoviita (1985) suggested that whilst Aongstroem (1876) first raised the sub-section to generic rank, attributing the change to C. Mueller, he did not actually validate the generic name, either as a new name or by elevating the supposed basionym to generic rank (Isoviita 1985). Historically the name Papillaria as a genus, had been used twice prior to 1876. Firstly Kickx used the name for a lichen-forming fungus in 1835, based on P. ventricosa J. Kickx, an Studies on the genus Papillaria 26

illegitimate name. In his 'Moosstudien' Lorentz (1864) validly raised Neckera subsect. Papillaria to generic rank. The name Papillaria C. Muell. was conserved in 1956 against Papillaria Dulac 1867 (a monocotyledonous marsh genus now named Scheuchzeria) and Tricholepis Kindb. (a moss genus) in 1899. Lanjouw et al. (1961) corrected this to Papillaria (C. Muell.) C. Muell. deleting Tricholepis. Isoviita (1985) proposed that the conserved name be cited as Papillaria (C. Muell.) Lorentz and that P. nigrescens (Sw. ex Hedw.) Jaeg. be adopted as the type species of the genus. He also suggested that the citation of Papillaria Dulac be dropped from Appendix III of the Code since Papillaria ( C. Muell.) Lorentz antedated it, and that the citation of Papillaria J. Kida f. nom. rej. be added to Appendix III.

The earliest recognized taxon now placed in Papillaria was Hypnum nigrescens Sw. ex Hedw. (1801) a species of the Americas and north Africa. Subsequently Pterigynandrum illecebrum Brid. and Neckera fuscescens Hook. were described in 1819. The former taxon occurs in the Caribbean and central America, and is now considered to be a variety of Papillaria nigrescens (Sw. ex Hedw.) Jaeg., while the latter, now treated as Papillaria fuscescens (Hook.) Jaeg., comes from southeast Asia. Thereafter species now included in Papillaria were described in the following sequence (Wijk et al. 1967, 1969):

1819-1829 2 species 1830-1859 11 1860-1889 22 1890-1919 30 Since 1920 six species have been described with the latest in 1983, but subsequently Noguchi (1976) transferred two of them into Floribundaria. Their occurrences are - northwest South America, Easter Island, Natal and China. Studies on the genus Papillaria 27

In his monograph 'A taxonomic revision of the family Meteoriaceae of Asia' Noguchi (1976) is of the opinion that 'the genus Papillaria seems to be a natural one'. But he believes that the distinction between the Asian P. jlexicaulis and P. crocea is not always clear. He notes that the taxon several authors called P. cuspidifera (Wils.) Jaeg. ( = P. crocea) should in fact be considered as P. flexicaulis. These two taxa are still being confused.

Dixon's 'Studies' 1927

New Zealand mosses have been studied in some detail and the problems that have been encountered there could reasonably apply to their Australian counterparts. The most critical work on Papillaria in New Zealand was that of Dixon (1927) in his 'Studies in the bryology of New Zealand' hereafter referred to as 'Studies'. Dixon was an English schoolmaster, whose studies on Australian and New Zealand taxa were based entirely on herbarium collections.

Before Dixon's 'Studies' the following morphological characters were used to segregate species in Papillaria:

leaf shape presence or absence of a leaf border denticulation of the auricles length of the costa. Dixon believed, however, that most of these characters were 'ill defined and elusive' and possibly inconstant and misleading. He considered the following character to be significant and based his key on:

'the position and appearance of leaves when dry, especially on the more robust branches, - the stem leaves and the leaves of the more slender flagelliform branches are apt to be less characteristic.' Studies on the genus Papillaria 28

In his key Dixon (p. 258) gave four characteristic leaf patterns for the six species then recognized for New Zealand - P. filipendula (Hook. f. & Wils.) Jaeg., P. crocea, P. flexicaulis, P. flavolimbata, P. nitidiuscula Broth. and P. amblyacis. However, leaf characters had to be used in conjunction with other features because they showed considerable variability.

Australian species

The earliest recorded Australian species of Papillaria [Pilotrichum nigrescens Hornsch. = Papillaria crocea] was placed into a new subtribe

Neckeraceae of the large tribe Hypnoideae by C. Mueller (1850). When W.J. Wilson published his 'Flora Novae Zealandiae' (1854) and 'Flora Tasmaniae' ( 1859) he placed Papillaria in the tribe Pilotricheae. In 1856 the family of

Neckerae (tribe Neckeraceae) was created by Schimper (Crosby & Magill 1977), and Jaeger (in Jaeger & Sauerbeck 1877) used this family for his new combinations in Papillaria. Subsequently, Hampe (1881), Brotherus (1906), Radway (1914), Brotherus & Watts (1918) and Dixon (1927) referred Papillaria to the family Neckeraceae.

The family Meteoriaceae was published by Kindberg in 1897. Fleischer (1908) was the first bryologist to use this name at family level for a publication where Australian species of Papillaria were mentioned. He was followed subsequently by Brotherus (1925) after which this family became generally accepted. It was quite common, however, for early authors not to assign families when describing taxa; for example Pilotrichum croceum Hpe (1853) and Neckera flavolimbata C. Muell. & Hpe (1853) were not assigned to any family. Studies on the genus Papillaria 29

The earliest Australian specimen of Papillaria was collected by Robert

Brown (NY) between 1801-1803 at an unknown locality when he accompanied Matthew Flinders on the 'Investigator' voyage. This was followed in 1823 by a collection made in New South Wales (near Sydney) by F.W. Sieber while R.W. Lawrence sent mosses (including Papillaria) from Tasmania to London between 1830-1833. It is surprising that more collections of Papillaria were not made earlier. For instance J.J. Labillardiere collected no Papillaria while in Van Diemens Land (Tasmania). It seems likely that Papillaria would have been common in the original vegetation of the Sydney area, based on the fact that F.v. Mueller collected it at Parramatta (1855) and H.L.K. Whitelegge at Homebush in 1885.

a) Papillaria crocea

C. Mueller (1850) cited Pilotrichum nigrescens for Australia based on a Sieber collection. The name P. nigrescens, however, is a nomen nudum now included in Papillaria crocea. In 1881 Hampe cited Neckera nigrescens Homsch. ex Hpe ( = Papillaria nigrescens) an apparent lapse when Pilotrichum nigrescens ( = Papil/aria crocea) was intended.

Hampe (1853) reported Pilotrichum croceum, based on C. Stuart's collection from Tasmania, the first species actually named subsequently as Papillaria for Australia. Later C. Mueller ( 1857) placed this taxon in the genus Neckera. J.D. Hooker and W. Wilson (in Wilson 1854) named Meteorium cuspidiferum Wils. based on a collection by A. Sinclair from the Bay of Islands, New Zealand. Mitten (1860) placed this species in Trachypus homschuchii Mitt. Another species, Neckera kermadecensis C. Muell., was described by C. Mueller (1857) based on a collection by Cuming from the Kermadec Islands. Studies on the genus Papillaria 30

Subsequently Jaeger (in Jaeger & Sauerbeck 1877) made the following combinations for these three names: Pilotrichum croceum = Papillaria crocea,

Neckera cuspidifera Tayl. (nom. nud. in syn.) = Papillaria cuspidifera and

Neckera kennadecensis = Papillaria kennadecensis (C. Muell.) Jaeg.

After studying the respective types, Dixon (1927) transferred P. cuspidifera, P. kennadecensis and Trachypus hornschuchii into synonymy under

Papillaria crocea. He felt that the confusion requiring the extensive synonymy had resulted from the variation encountered within the wide geographical range.

He stated that the 'Indian specimens of P. crocea are weakly plicate and sometimes hardly so, while the Australasian plants are markedly plicate' (p. 260). But he had noted some Indian collections that were plicate and one New

Zealand collection that was not plicate. As a consequence his original key was not satisfactory so he provided two characters to help distinguish between P. crocea and P. flavolimbata, and another two characters to distinguish P. crocea from P. flexicaulis, P. filipendula and P. amblyacis. Distinguishing characters cited for P. crocea by Dixon (1927) are:

1) leaves may be erect,.± rigidly appressed 2) plant is rigid, not flexuose nor undulate 3) stout costa shows a clear furrow on each side (plicate) 4) apex is not recurved. Studies on the genus Papil/aria 31

b) Papillaria flavolimbata

Neckera flavolimbata was described by C. Mueller and E. Hampe (1853) based on an Australian collection from Sealers Cove in Victoria (BM). However, a collection in the Melbourne Herbarium (MEL) with no data, is also annotated as a type, together with one from Sealers Cove. Meteorium cerinum

Hook.f. & Wils. was described by J.D. Hooker and W. Wilson (in Wilson 1859) based on Tasmanian collections by R.C. Gunn (1605) and W. Archer (on trees, in creek) cited as types. Later Mitten (1860) placed this species in Trachypus giving Neckera cerina (Hook.f. & Wils.) Tayl. and N. luteola Tayl. as synonyms. This taxon was then relegated to a variety in Meteorium cuspidiferum by J.D. Hooker (1867) and eventually Paris (1896) made the combination Papillaria cerina (Hook.f. & Wils.) Par. Jaeger (in Jaeger & Sauerbeck 1877) made the combinations: P. flavolimbata and P. cuspidifera var. cerina (Hook.f. & Wils.) Jaeg. Both P. cuspidifera var. cerina and P. cerina were placed into synonymy with P. flavolimbata by Wijk et al. (1967:6,7) where they referred to Mitten 1882:82. In this latter reference Mitten reports Meteorium cerinum as a synonym of M. limbatum Mitt., but the latter is a nom. illeg. because Neckera flavolimbata had priority.

In his discussion Dixon (1927), states that Papillaria flavolimbata is the most robust of the New Zealand species and frequently has long filiform, flagellate branches. At times the colour of this moss can be orange-yellow.

Other characters of this species are:

1) costa prominent on the back of the leaf ( also in P. crocea) 2) leaves undulate and flexuose 3) leaf margins often recurved 4) pale border, more marked than in other species 5) upper cells often obscured by papillae Studies on the genus Papillaria 32

6) youngest leaves terminate with a forked tip ( communicated by Sainsbury).

c) Papillaria flexicaulis

The next species to be described was Meteorium flexicaule Wils. Wilson (1854) based this on several New Zealand collections, one by Dr Stanger from the North Island and another by W. Colenso from Hawke's Bay. Mitten (1860) transferred this species to the genus Trachypus. J.D. Hooker and W. Wilson (in

Wilson 1859) then named Meteorium filipendulum, based on Tasmanian collections by R.C. Gunn (26 & 1606) and R.W. Lawrence, while collections from Ash Island in the Hunter River made by H. Scott and from the Brisbane

River (lectum mis Cl. G. Reichenbach) were cited as types of Neckera scottiae C. Muell. by Mueller (1868). In 1877, Jaeger transferred these taxa to Papillaria ftexicaulis, P. filipendula and P. scottiae (C. Muell.) Jaeg. respectively. Fleischer (1914) also cited Cryphaea novae-valesiae C. Muell. as a synonym of Papillaria filipendula, but this epithet is a nomen nudum.

Sainsbury ( 1935) later placed both P. filipendula and P. scottiae into

synonymy with P. flexicaulis.

Dixon (1927) admitted that P. flexicaulis and P. filipendula are very closely allied and that the characters separating these two species are unreliable. He rejected Rodway's (1914) opinion that these two species were conspecific. Dixon felt that the distinctions should be based primarily on habitat and partly on leaf form. The characters he used to separate the species were: Studies on the genus Papillaria 33

P. flexicaulis P. filipendula

plant moderately slender plant extremely slender plant stouter, more rigid plant subpiliform leaves wider leaves very small leaf tip acute leaf tip acuminate not piliferous often piliferous yellow-green dull olive-green

He considered P. flexicaulis to be intermediate between P. amblyacis and P. filipendula in leaf characters, with the upper lamina cells to be nearly identical to those of P. crocea.

He claimed this species to be the most slender of all the New Zealand

Papillarias, but noted that some attenuate forms of P. crocea might be nearly as slender.

Sainsbury (1955b) mentioned the problems elaborated by Dixon (1927). Contrary to Dixon's opinion, Sainsbury accepted Rodway's conclusion (1914) that P. flexicaulis and P. filipendula were conspecific and gave the former priority. He considered P. filipendula to be only a slender form of P. flexicaulis.

d) Papillaria fulva

A collection made by F.v. Mueller during 1853 at Tarwin in Gippsland (Victoria) was described by Mitten (1860) as Meteorium fulvum Mitt. This

species was transferred to Papillaria by Jaeger (in Jaeger & Sauerbeck 1877), but Scott & Stone (1976) suggested that it could well be P. nitens ( = Barbella nitens). Studies on the genus Papillaria 34

e) Papillaria squamata

Hampe ( 1870) described Neckera squamata Hpe basing it on an Australian collection with no collection data, but in 1881 he transferred it to Papillaria.

f) Papillaria amblyacis

Neckera amblyacis C. Muell. was named by C. Mueller (in Hampe 1870) who designated a Leichhardt collection as the type. Jaeger (in Jaeger & Sauerbeck 1877) made the combination Papillaria amblyacis.

Dixon (1927) claimed that of the four species occurring in both New Zealand and Australia, P. amblyacis was the easiest to recognise, using the following characteristics:

1) branches terete and julaceous when dry 2) branch leaves - very broad, smooth and convex at back - very closely imbricate - exhibited a very short and abrupt pale recurved mucro or cuspidate point - with a costa longer than any other species. Studies on the genus Papillaria 35

g) Papillaria eavesiana and P. reginae

Hampe (1876) described Neckera eavesiana Hpe and N. reginae Hpe sequentially in the same publication. The former was based on a collection by Eaves from tropical eastern Australia [Queensland], while the latter was based on a Hartmann collection from south Queensland. At present both taxa appear under Papillaria where they were placed by Jaeger (in Jaeger & Sauerbeck

1877). Papillaria eavesii Kindb. (Enum. Bryin. Exot. 27.1888) is an illegitimate name; the epithet eavesiana has priority (Wijk et al. 4:7.1976). Kindberg's worldwide listing of species introduced 'simplified' spellings for several other

Australian moss species as well.

After 1877 some Papillaria species (P. amblyacis, P. eaveszana, P. filipendula and P. scottiae) were placed into Pilotrichella by Hampe ( 1881) and

Meteorium (Papillaria amblyacis) by Mitten (1882), but Jaeger's combinations

are still recognized today.

Norfolk Island species

The first report of Papillaria from Norfolk Island was Meteorium

cuspidiferum ( = P. crocea) by W.J. Wilson in 1854, while in 1867 J.D. Hooker reported M. cuspidiferum var. cerinum (Hook.f. & Wils.) Hook.f. & Wils. ( = P. flavolimbata ). Strangely, Maiden ( 1904) did not mention Papillaria amongst the

mosses for the island. No discussion of the Norfolk Island specimens was found

in literature. Studies on the genus Papillaria 36

Early comparisons of Australian taxa

Bryologically-rich Tasmania was the first Australian colony to have the mosses investigated by a resident bryologist, namely by R.A. Bastow ( 1887). He mentioned four Meteorium species (now in Papillaria) giving details of the species' characteristics:

M. cerinum ( = Papillaria flavolimbata) M. cuspidiferum ( = P. crocea) M. filipendulum ( = P. flexicau/is) M. flexicau/e ( = P. flexicau/is)

Rodway (1914) followed Bastow and was the first to publish a key to Tasmanian Papillaria where he listed the following species:

P. filipendu/a (syn. Meteorium flexicau/e) P. flavolimbata P. intricata P. kennadecensis

In the Tasmanian flora he accepted P. kermadecensis rather than P. crocea and included P. intricata (Mitt.) C. Muell., which was originally described from the Pacific Islands (Samoa). Sainsbury (1955b:50,51) subsequently inferred that this latter name could be a misdetermination for Barbella nitens. I have since located this collection amongst material from HO (Coward as Weymouth 2111c) and have confirmed that this is indeed so. Many W.W. Watts collections were determined initially by V.F. Brotherus (H, NSW) as P. intricata, but fortunately they were never published under this name. Studies on the genus Papillaria 37

Brotherus (1906,1925) reported the following Papillaria species for Australia:

P. amblyacis P. flavolimbata P. crocea P. kennadecensis P. eavesiana P. reginae P. filipendula P. squamata

He reported P. flexicaulis for New Zealand only, while in his 1925 work neither

P. fulva nor P. nitidiuscula were listed for Australia.

In his 'Studies' Dixon (1927) discussed at length the problems relating to P. cuspidifera and P. kermadecensis (both now considered synonyms of P. crocea). He stated that when C. Mueller (1857) described Neckera kermadecense no comparison was made with Papillaria cuspidifera, hence he believed that two tax.a must be involved. However, in 1868 C. Mueller did recognize the affinities between the specimens and listed Meteorium cuspidiferum as a synonym of Neckera kermadecense. This transfer has been accepted for the purposes of this revision.

Despite Dixon's (1927) extensive writings he came to few conclusions.

For instance, although he claimed that the reported differences between P. flexicaulis and P. filipendula were quite unreliable he did not synonymize these species, unlike Rodway (1914) who came to a similar conclusion and listed P. filipendula as a synonym of P. flexicaulis.

Papillaria was not included in Watts and Whitelegge (1902,1906) 'Census Muscorum Australiensium' since this work only covered acrocarpous species. Burges (1932,1935) who completed this 'Census' after the death of Watts, recorded seven species of Papillaria in New South Wales but was misleading in claiming that P. crocea occurred 'throughout the state'. He included the name Studies on the genus Papillaria 38

'P. filicaule Tayl. ex Burg.' in the 'Census' (1935), apparently a typographical error referring, in fact, to P. flexicaulis.

The Tasmanian representatives of Papillaria were studied by Sainsbury (1955b) and the current view of this genus in Australia was consolidated by him. He was not surprised by the morphological variation observed in P. crocea given its extensive range from India to the Pacific Islands. He considered that P. filipendula was simply a slender form of P. flexicaulis, but that P. flavolimbata was a characteristic species.

Several new combinations in Papillaria were proposed by Hilbrands (1969) in his unpublished studies and herbarium collections were thus annotated.

Scott and Stone (1976) provided descriptions and a key to five common Australian species. They suggested that P. fulva might be identical to Barbella nitens but did not elaborate. They also listed, but did not discuss the three Australian mainland endemics - Papillaria eavesiana, P. reginae and P. squamata. In fact, no one had critically examined these three species to ascertain their taxonomic relationships. The Papillaria species recognized by Scott & Stone (1976) were:

P. nitens (Hook.f. & Wils.) Sainsb. P. amblyacis (C. Muell.) Jaeg. P. crocea (Hpe) Jaeg. P. flavolimbata (C. Muell. & Hpe) Jaeg. P. flexicaulis (Wils.) Jaeg. P. eavesiana (Hpe) Jaeg. P. reginae (Hpe) Jaeg. P. squamata (Hpe) Hpe P. fulva (Mitt.) Jaeg. Studies on the genus Papillaria 39

In addition to P. crocea, Noguchi (1985:99) listed P. leuconeura for Australia and New Zealand. Although he had not seen the type of P. leuconeura, he believed P. amblyacis to be conspecific with it (pers. comm. 1986). P. leuconeura has been reported from southeast Asia and New Caledonia.

Although Papillaria seems now to be a natural genus the early confusion on specific limits seems surprising. For example the confusion in the syntypes for P. flexicaulis where one syntype was P. amblyacis. Or the fact that most collections of P. eavesiana ( = P. amblyacis) from the Mitten Herbarium (NY) were annotated as P. crocea. In both cases the distinction between the species is quite marked. Studies on the genus Papillaria 40

TAXONOMIC RELATIONSHIPS IN AUSTRALIA

Cladistic Analysis

An attempt was made to analyse the phylogenetic relationships within the Australian species of Papillaria. In addition to the Australian species, four non-Australian species (P. semitorta (C. Muell.) Jaeg., P. subaongstroemiana Fleisch., P. chrysoclada (C. Muell.) Jaeg. and P. fuscescens) were included with the last two species being initially considered the outgroup. In the outgroup method a genus or species is selected which is considered to have a close relationship to the genus or species being studied (Maddison et al. 1984). Well corroborated hypotheses of cladistic relationships are rare in mosses (Rohrer 1985) and the two outgroup species were chosen only because they differed in having immersed capsules. It is not known whether this character was derived. With the outgroup method the polarities were determined with respect to the outgroup as the ancestral states were not known. From the apomorphy lists produced by the program the supposed ancestral states for the characters could be deduced.

Traditional, unambiguous characters were chosen which were also available in Noguchi (1976) for the non-Australian species. Cladistic characters (Table 7) were analysed by the computer program PAUP version 2.4, which was developed by D.L. Swofford, with Penny's branch and bound algorithm using parsimony analysis (Farris 1983). This was run on an IBM XT personal computer. Early attempts suggested that the two outgroup species were not suitable and that P. semitorta may be more suitable because it tended to constantly separate out on its own. This proved to be so, resulting in only one parsimonious tree being found (Fig. 5) with a consistency index of 0.47. In the homoplasy parallelisms predominated with characters 5, 15, 19, 21, 23, 24 being s. ~ ;:: .., ~· ~ tl ..., § ~- ::- ;::

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feuc ambl flav flex zelo suba croc intr fuse chry semi 2 5 10 7 19 15 10 23 21 19 24 6 25 4 5 8 18 12 14 22 14 19 19 20 9 21 21 23 24

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Fig. 5. Cladogram of Australian Papi/laria and outgroup species. Parallelisms are represented by double bars, reversals by arrows. The numbers refer to the characters given in Table 7.

particularly unstable. In further studies these characters need to be looked at closely with a view to redefining or omitting them. The number of synapomorphic characters was six. Using the same character set but having P. Studies on the genus Papillaria 43

intricata as the outgroup the synapomorphic characters remained at six, but the general relationship did not alter significantly. The length of the tree (53) and the consistency index did not alter.

The cladogram (Fig. 5) did confirm the suspected close relationship between P. flexicaulis and P. zeloflexicaulis. However, surprisingly P. crocea was not close to P. flexicaulis with which it was often confused. Also a closer relationship was expected between P. amblyacis and P. leuconeura.

In the program Meteorium was used as the outgroup but while Meteorium is closely related to Papillaria it is not close enough to be considered part of the genus. While the method clearly distinguished between Meteorium and Papillaria as genera, the program did not separate the individual Papillaria species.

For a more confident resolution of the relationships a greater range of characters should be used, e.g. from the perigonium, perichaetium and the exostome. Ideally for this type of analysis all Papillaria species worldwide ( 69) would have to be considered against an outgroup of another genus.

Relationship between Papillaria and Meteorium

While discussing Papillaria nitens, Sainsbury (1955b) remarked on the weakly defined differences between the genera Papillaria and Meteorium. The differences are not clear cut because the two genera are based on gametophytic characters which tend to intergrade, but they appear to be closely related

(Noguchi 1976:235,236). W.R. Buck (pers. comm.) claimed that these two genera are often hard to separate in collections from the Neotropics. In both genera the basal angles of the leaves develop into auricles, and peristome Studies on the genus Papillaria 44

structure is similar with exostome teeth and segments irregular in outline. These characters set the two genera apart from the rest of the family. The only characteristic difference between the genera is that Papillaria is pluri-papillose while Meteorium is predominantly uni-papillose with resultant clearer cells. Because of their close relationship some species originally described as Papillaria were subsequently transferred to Meteorium.

In 1888 V.F. Brotherus published Papillaria baileyi Broth. based on a Bailey collection from the Brisbane River, but he later transferred this to Meteorium (Brotherus 1906). Mitten (1882) described M compressum, again on a Bailey collection from the same localilty, but although this was subsequently (Enum. Bryin. Exot. 102.1891) transferred to Papil/aria by Kindberg, this change has not been accepted.

In the present study seven species of Papil/aria are recognized for Australia. One of these, P. zeloflexicaulis, is described as a new species; the remaining species are - P. amblyacis, P. crocea, P. flavolimbata, P. flexicaulis, P. intricata, and P. leuconeura. For Meteorium only M. buchananii is recognized as occurring in Australia. The taxonomic conclusions were based on extensive morphological studies. Studies on the genus Papillaria 45

MORPHOLOGICAL STUDIES

Gametophyte

Constant gametophytic characters for species differentiation are difficult to find in Papillaria. Dixon (1927) discussed the problems he encountered when he used the leaf shape, presence or absence of a clear border, denticulation of the auricles and the length of the costa; he concluded that these characters are ill defined and often lead to confusion. He preferred to use the appearance and position of the dry leaves. This feature is more obvious and constant in the robust branch leaves than in the stem leaves or on the long, slender trailing forms. This seems to be the most reliable approach in the light of present knowledge and the species key has been constructed accordingly. The stem leaves are often damaged or worn down by abrasion with grit and dirt and it may be difficult to obtain intact representative leaves. The leaf border was found to be quite variable or absent and it proved to be an unreliable character except in P. flavolimbata. Many previous misdeterminations could have been due to too much reliance being placed on leaf border as a character.

a) General morphology

Papillaria species are perennial plants and often form extensive, interwoven pendulous mats which commonly trail from tree branches, tree trunks and rock faces. The primary and secondary stems of these mosses are clearly differentiated. When pendant, the branches may be slender and flagelliform. Such morphotypes are very common in the case of P. f/avolimbata but are rarer in P. crocea. In the past these slender forms have often been Studies on the genus Papi//aria 46

collected frequently without the main branches and this had led to several new species being described erroneously.

b) Colour

Papillaria is always green but can vary from a lighter or yellowish-green in P. crocea to a dull, darker green in other species. The colour is also dependent on the age of the moss, thus the young growing stems and branches are light green while the older parts may be reddish to brownish. The latter colour is common in P. intricata. Old herbarium specimens generally change to golden-green or to golden-brown.

c) Stems

The stems of Papillaria species are rounded to oval in cross-section and up to 400um thick. They are usually grey-green, but may darken with age. The structure of the cross-section of the branches (Fig. 6) usually resembles that of the main stem (Fig. 7). The epidermis plus outer cortex are 4 or 5(6) cells thick and of a distinct golden colour. In transverse section the epidermal cells cannot be distinguished from the outer cortical cells. Cells near the perimeter are small (1.0-2.0um) and rounded with very thick walls, the cell size increases towards the inner cortex. Sometimes the cells can be narrowly elongated and pointed towards the central strand. Studies on the genus Papi/laria 47

2

3

5 6

~ 0 ..0

Fig. 6. Branch cross-sections of Papillaria. 1. P. amblyacis (HS 27933); 2. P. crocea (HS 27944); 3. P. flavolimbata (HS 7798); 4. P. flexicaulis (Watts M13006); 5. P. leuconeura (HS 29717); 6. P. intricata (Dietrich-H); 7. P. zeloflexicaulis sp. nov. (Watts 5737). Studies 011 the genus Papillaria 48

In the transparent inner cortex the cells are large, up to 30um in the centre, with very thin walls. However, P. amblyacis can often have very irregular, even triangular cells and the cell walls are thick. A central strand of 2 or 3 very small cells is present in the stems of all species, but are least developed in P. crocea. A central strand was noted only in the branches of P. flavolimbata and P. flexicaulis.

1 2 3

Fig. 7. Stem cross-sections of Papillaria: 1. P. flexicaulis (x215 - HS 31651); 2,3. P. amblyacis (x430 - HS 27933).

d) Leaves

The leaves are generally arranged spirally on the stems and branches, being unistratose except near the costa. In P. crocea and often in P. zeloflexicaulis the leaves are arranged in 5 longitudinal ranks along the stem being quite pronounced when the leaves are wet and spread at 45° from the stem. In the julaceous species (P. amblyacis, P. leuconeura and P. flexicaulis) the Studies on the genus Papillaria 49

moist leaves do not usually spread widely. In P. flavolimbata the leaves become more regular when moist with the bent back margin straightening. Generally the branch leaves are smaller than the stem-leaves, the variation being between 11-33% in the length and 5-30% in the width. This was not always the case in P.

amblyacis where branch and stem leaves were similar. Figs. 8 & 9 show the

I P,pill1ri1 amblyacis I I I 2 n•201 I I I ' I I I ' ;;;- I I I .. I I I I I O! 3 2 I 4 2 3 3 1 8 I 2 I I I I I 2 I 2 I 3 t I I ~ I I 11 4 I I l24 II I 21 2 e I I ' 3 I I 2 3 2 41 2 I 111 .§. 11 I 2 2 .&: 2 I 3 I I I I I I .. 2 2 I 4 3 2 I :2 I 11 2 3 I 3 I I I I I I I I 22 I I I I 2 I I ] 2 2

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Fig. 8. Branch leaf length/width in Papillaria amblyacis (C. Muell.) Jaeg. and P. crocea (Hpe) Jaeg.

relationship between leaf length and width of P. amblyacis, P. crocea, P. jlavolimbata, P. flexicaulis and P. zeloflexicaulis. It is apparent that the majority Studies on the genus Papillaria 50

of the branch leaves of the latter are smaller than that of P. flexicau/is with which it was previously confused. P. flavolimbata is the species with the largest leaves. The cluster of large leaves for P. flavolimbata (Fig. 9) was from a

Berggren collection from Fernshaw (between Lilydale and Black Spur) in

Victoria. Two other duplicated collections (H, NY) had leaves within the normal ranges. From Fig. 8 it can also be noted that an increase in leaf length did not lead to an obvious increase in width for P. amblyacis. Because of insufficient collections, the leaves of P. intricata and P. /euconeura were not studied. The width to length ratio of branch leaves was essentially the same for all species (0.49-0.54) and for stem leaves 4-18% higher, otherwise the morphological differences between species were small. In this study leaves from the central part of the branch were used for comparison because the stem leaves were often dead or broken from abrasion.

Papillaria species exhibit a whorl of small rounded, sparsely papillose, ecostate leaves at the base of each branch. Above this whorl (Fig. 50 p. 176) the leaves gradually change to normal branch leaves with the costa length increasing, so caution must be exercised when choosing which leaves to examine.

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e) Leaf shape

The leaves of Papillaria are ovate to broadly lanceolate and up to 3mm long. In the majority of Australian species the apex is acute but in P. amblyacis it is generally apiculate. Furthermore the leaves are normally asymmetrical, especially towards the base. This is most marked in P. flavolimbata but not so marked in P. flexicaulis.

The leaf base is often clasping, being generally more prominent in the stem leaves, and is particularly obvious in P. flexicaulis.

Auriculate leaf bases were best developed in the stem leaves, but are also prominent on the branch leaves. P. flavolimbata showed this feature best, while P. intricata has this feature least developed.

f) Leaf margin

The margin of the leaves in Papillaria is usually weakly, and only rarely strongly denticulate or toothed in the auricles (Fig. 10). Sometimes the denticulation can continue up the leaf margin, becoming successively weaker towards the apex or it may be absent. All the Australian species show this variation except P. flexicaulis and P. zeloflexicaulis which appear to have a nearly entire margin. Only under the highest magnification are slight denticulations apparent. Studies on the genus Papillaria 53

g) Costa

In all Australian species, except P. intricata, the costa reaches at least 3 / 4 of the leaf length, but in some species it approaches the leaf apex. Width of the costa at the base is between 30-SOum for the majority of species but in P. jlexicaulis it is only 20-30um wide. The costa is quite inconspicuous in P. jlexicaulis and P. amblyacis, being barely visible even under 16-fold magnification under a stereomicroscope or hand lens. This may result, in part, from a lack of colour differentiation between the leaf cells and the costa. When the leaf is a dark green the dark costa is not easily visible, but on a light coloured leaf the reddish or reddish-brown costa is quite visible. The costa of P. intricata can be quite faint. The costa cells are homogeneous in cross section (Fig. 11).

h) Lamina cells

The lamina cells of Australian Papillaria species are elliptical to rhomboidal. The median cell walls are moderately thick, while cells towards the apex are often longer and more incrassate. In P. leuconeura all these cells are incrassate and sigmoid. Cells in the auricles in P. jlexicaulis and P. zelojlexicaulis are uniform and similar to the median cells, with cells intersecting the margin at right angles (Fig. 10.1). In all other species the cells in the auricles diverge and align parallel to the margin of the auricles (Fig. 10.2). Parallel auricular cells in P. crocea sometimes are not so prominent, but small cells nearer the base of the auricle do show this tendency. Cells in the leaf base nearer the costa are porose only in P. amblyacis. In all species there are larger cells in the basal corners, Studies 011 the genus Papillaria

often 1 or 2 rows being golden or golden-brown. The juxtacostal cells at the base are long with thin walls.

1 2

Fig. 10. Auricular cells of Papillaria (x600). 1. P. flexicaulis (Watts 2129); 2. P. crocea (Boorman as Watts 1544).

The leaves of all species bear papillae which obscure the cells to a varying degree (Fig. 11 ). In P. flavolimbata (Fig. 11.4) they are sufficiently dense to make it exceedingly difficult to see the cells. In most species this is true for both stem and branch leaves, but in P. amblyacis the stem leaves have very few papillae and these are often difficult to detect. Papillae are usually densest on the mid-leaf cells and juxtacostal cells are transparent in the auricles with papillosity increasing towards the margins. The papillae appeared to be two per cell, in P. amblyacis rounded and up to lum high, in P. flavolimbata a large 'mushroom' shaped papilla 2um high was present over each cell with a small definite gap between the cells, and P. crocea and P. flexicaulis have one or two very small papillae over each cell while P. intricata had one papilla per cell, but not all cells having papillae. The more incrassate the cell the bigger the gap Studies on the genus Papillaria 55

between the papillae. In some species papillae may be lacking in the leaf tips and in the 2-5 rows of marginal cells.

i) Leaf cross-section

Cross-sections of leaves were very similar in all seven species (Fig. 11 ). The same variation in cell shape and size was present in all, but there was variation in the types of cells of the costa. The thickness of the costa in cross­

section was between 20-55um, with P. flavolimbata (Fig. 11.4) having the most prominent costa. Leaf cells in most cases were ovate to rounded with P. crocea tending to quadrate, with cell size between 4-lOum. Leaf thickness varied from 10-25um with P. flavolimbata being the thickest and with the largest cells. Wall

thickness between cells was 1.0-2.5um, but in P. intricata (Fig. 11.5) walls tended

to be thinner. Cell numbers in the costa varied: P. flexicaulis had about 8; P.

zeloflexicaulis 9-11; P. flavolimbata up to 20, while in the other species it was between 13-16. Costa cells were often in three irregular rows and were generally smaller than those of the leaf, their size and shape being quite variable. Very rarely was anything approaching a definite row of larger guide cells noted. Studies on the genus Papillaria 56

Fig. 11. Cross-sections of Papi//aria branch leaves near mid leaf. 1. P. amblyacis (HS 27933); 2. P. crocea (near base - HS 27944); 3. P. crocea (HS 27944); 4. P. flavolimbata (HS 7798); 5. P. flexicaulis (HS 38465); 6. P. intricata (Dietrich-H); 7. P. leuconeura (HS 2~717); 8. P. zeloflexicaulis (Holotype). St11dies 011 the ge11us Papillaria 57

j) Rhizoids

In all species the rhizoids were reddish, smooth and similar in form (Fig.

12). The width was generally 12um, but in P. jlexicaulis (Fig. 12.1) and P. intricata the rhizoids were 17um and 18um wide respectively. The cells varied in length from lOOum (P. jlexicaulis) to 200um (P. jlavolimbata). Generally the cells are prosenchymatous but in P. intricata and P. leuconeura (Fig. 12.3) the cell ends are rounded and overlap. Sometimes the cell boundaries were not distinct. In P. crocea and P. jlexicaulis the rhizoids were borne in dense clusters, and in the latter species were quite common, especially near the branch tips.

' ~c.=====....""""_==-_==-_==-.,,.==-_==_-_ -- -_-t!,.2~::::::::::1e~:::.-- z ::: s

3 == *

tOOpm

Fig. 12. Rhizoids of Papillaria. 1. P. flexicaulis (Watts M13006); 2. P. flavolimbata (Forsyth M12991); 3. P. leuconeura (HS 28688).

Sexuality

The genus is dioicous with male and female plants similar in size.

Fertile material was uncommon in Papillaria and often quite rare for some species (e.g. P. jlexicaulis). Perigonia and perichaetia were even rarer, in fact no perigonia or perichaetia were noted in Australian collections of P. Studies on the genus Papil/aria 58

intricata or perigonia in P. leuconeura. However this may be due to the small numbers of collections available for study. Features dependent on the sexual parts and the sporophyte cannot be used, therefore, in the specific key because the information is from a limited number of samples only.

Perigonia and perichaetia occurred on trailing branches in the species studied.

Perigonia

Rounded perigonia were borne singly on the branches and resembled leaf buds but were often distinguished from the latter by having slightly protruding, acuminate leaf tips. Leaf buds however had smaller leaves with rounded apices.

Inner perigonial leaves were longest and in all species tended to be ovate, oblong or intermediates, except for P. intricata, in which they were lanceolate. The inner perigonial leaf length could be classified into two groups:

0.9-1.lmm - P. amblyacis, P. flavolimbata, P. flexicaulis, P. leuconeura 0.5- 0.6mm - P. crocea, P. intricata, P. zeloflexicaulis

Leaf tips were acuminate, margin entire or distantly irregular. Narrower elongated marginal cells were noted in P. amblyacis and P. crocea, while in P. flavolimbata this was more prominent in the basal section. It appears that as the perigonial leaves mature this margin of rectangular cells becomes less prominent. No costa was noted in P. amblyacis or P. jlexicaulis, while in P. crocea, P. jlavolimbata and P. zelojlexicaulis a weak costa to 3/4 of leaf length Studies on the genus Papi/laria 59

was noted. In the last two species the costa was not always present. This feature again depended on the stage of perigonial leaf development. Weak papillae were noted on all perigonial leaves.

Upper perigonial leaf cells were hexagonal or rhomboidal, being incrassate in P. crocea, P. flexicau/is and P. intricata while the latter species had very narrow (Zurn) cells. Longest cells (37urn) were in P. flavolimbata and P. amblyacis while P. zeloflexicau/is had the shortest (12urn). Median cell shapes were similar to upper cells, but longer, with P. intricata (62um) and P. zeloflexicau/is (12-23urn) representing extremes.

Antheridia were club shaped and reddish when mature with an irregular pattern of quadrate or shortly rectangular cells (Fig. 13 ). The number of antheridia per perigoniurn varied from about 20 in P. flavolimbata to about 5 or 6 in P. crocea, P. flexicau/is and P. zeloflexicau/is. The largest antheridia were in

P. flavolimbata ( 400x120um - Fig. 13.4 & 13.5) while the smallest were in P. zeloflexicaulis (280x120urn - Fig. 13.7).

Paraphyses were clear, thick walled, uniseriate, multicellular and generally as long and as many as the antheridia. The thickness of the paraphyses varied from 15um in P. zeloflexicau/is to 25um in P. amblyacis, longest cells being in P. amblyacis (75urn) and shortest in P. zeloflexicau/is ( 40urn). Immature paraphyses appeared catenulate with weak cell outlines. Studies on the genus Pupil/aria 60

Fig. 13. Antheridia of Papi/laria. 1,2. P. amblyacis (Watts 1688); 3,4. P. crocea (Watts 9470); 5,6. P. flavolimbata (Watts 10361); 7. P. flexicaulis (Schofield 90576); 8. P. zeloflexicaulis (Bailey-H).

Perichaetia

Perichaetia were borne singly on the branches (Fig. 40.1 & 40.2 p. 150) and were as rare as perigonia. In fact in P. amblyacis unfertilized perichaetia were extremely rare. The buds were rounded and similar to perigonia, but elongated following fertilization. Studies on the genus Papillaria 61

In most species inner perichaetial leaves were variously ovate to obovate with long acuminate apices. Perichaetial leaves ranged from 850-1600um, except in P. zeloflexicaulis (550um). Perichaetial leaves were entire in most cases except in P. amblyacis and P. leuconeura which were denticulate in upper half, and P. flavolimbata which had a serrate tip. Marginal cells were narrow elongate in P. crocea, in the upper half only of P. leuconeura and P. zeloflexicaulis, while in the latter species the margin had quadrate cells in the lower half. In P. amblyacis the margin was weakly developed with rectangular cells or absent. P. intricata and P. leuconeura were ecostate while P. amblyacis sometimes had a very faint costa. However, a faint costa reaching 3/4 leaf length was noted in P. crocea, P. flavolimbata and P. zeloflexicaulis, while it was only 2/3 leaf length in P. flexicaulis . Weak papillae were noted on all species except P. crocea and P. flavolimbata. During the early stages of perichaetial leaf development marginal cells were rectangular and the costa lacking.

Upper leaf cells were variously elongated in all species, being incrassate in P. crocea and P. flavolimbata. Cells were longest in P. intricata (37um), varying from 25 to 30um in the remaining species with P. intricata and P. zeloflexicaulis having the narrowest cells (Zurn). Median cells were variously rhombic or hexagonal in all species excepting P. intricata where they were elongate. Cells were longest in P. intricata and P. leuconeura (63um) but shortest in P. zeloflexicaulis (20um).

Archegonia (Fig. 14) were reddish when mature, with a pattern of quadrate and rectangular cells, while in P. amblyacis (Fig. 14.1) they were nearly quadrate. The neck of P. crocea (Fig. 14.2) appears most distinct and more multicellular than expected. The number of archegonia varied from about 12 in P. flavolimbata to about 5 or 6 in P. amblyacis, P. crocea, P. flexicaulis and P. Studies on the genus Papillaria 62

zeloflexicaulis. Archegonial necks were longest in P. flavolimbata (410um) and P. leuconeura ( 450um) and shortest in P. intricata (280um).

Paraphyses were clear, thick-walled, uniseriate, multicellular and mostly as long, and as many, as the archegonia. Thickness of paraphyses varied from 12um in P. intricata to 20um in P. amblyacis. Cells were between 50-63um long except in P. zeloflexicaulis (37um) and in P. crocea ( 40um). Immature paraphyses were similar to those in perigonia.

After fertilization, as the young sporophyte develops, the perichaetial leaves and paraphyses also appear to grow and the latter increase in number. Paraphyses were biseriate in the lower half, while at the base a very short reddish-orange section of small quadrate cells (3 cells wide) was present.

The perichaetial leaves at the base of the capsule did not seem to have any characters that were taxonomically significant excepting those of P. flavolimbata which were plicate. In all species the length of the inner perichaetial leaves varied from 1.5-6.0mm, being narrowly triangular to oblong­ lanceolate with parallel sides. In one collection of P. crocea (Watts 9908) conspicuous alar cells were noted.

The vaginula was generally only lightly attached and the sporophyte broke off readily. There were no significant size or shape differences in the foot of the various species.

Numerous paraphyses were present at the base of the capsules with cells varying in length, lO0um being the longest observed. Cells near the centre of the paraphyses were longest. Paraphyses were up to 3 cells wide. The clustered paraphyses were slightly fused near the base at insertion. No specific variation was observed in the paraphyses of the species examined. Studies on the genus Papillaria 63

2 4

100 Jm

•

Fig. 14. Archegonia of Papillaria. 1. P. amblyacis (HS 9336); 2. P. crocea (HS 38467); 3. P. flavolimbata (Boorman ?as Watts 1519); 4,5. P. flexicau/is (Forsyth as Watts 1109); 6. P. /euconeura (Norris 34566); 7. P. intricata (Fleischer B1249 from Samoa); 8. P. zeloflexicaulis (Watts 5673). Studies on the genus Papillaria 64

Sporophyte

a) Calyptrae

The calyptrae were cucullate (Fig. 15) and hairy. P. zeloflexicaulis had the smallest calyptra while the largest was noted in P. flavolimbata.

d2 :% ~ " ~ ~ ! \~ 1· t~ ,_ /{21 2i -~g

Fig. 15. Calyptrae and opercula of Papi//aria. 1-4. P. amblyacis (Watts 3806): 1. Calyptra; 2-4. Operculae. 5-10. P. crocea: 5. Calyptra (Porter as Watts 2284); 6- 10. Operculae (Tindale & Constable M12981). 11-17. P. flavolimbata: 11. Calyptra (Watts 6176); 12-17. Operculae (12-15,17. HS 15369, 16. Curnow 1287). 18-21. P. zeloflexicaulis: 18. Calyptra (Watts 1728); 19-21. Operculae (19. Holotype, 20,21 Watts 1728). 22. P. flexicaulis: operculum (HS 38465).

b) Setae

Sporophytes were lateral on the branches, usually one per perichaetium but two have sometimes been noted (P. crocea - Watts 9468). Studies on the genus Papillaria 65

The length of the seta varied between species being 2.8-4.Smm long in P. crocea and up to 12.6mm long in P. flavolimbata. Normally there was no noticeable twisting of the seta, but occasionally it was slightly twisted to the right. The specific significance of this character could not be readily assessed since fruiting material was so uncommon, being particularly rare in P. flexicaulis.

c) Capsules

Capsules of Papillaria were erect and symmetrical with an ovoid urn (Fig. 36 p. 136). The exothecial cells, though irregular, were generally square to rounded and about 25um in width. The cells of inner layers of the capsule jacket were usually similar, but more difficult to observe clearly. In all species the thin spore sac tended to have cells similar in size and shape to the exothecial cells. Near the base of the capsule the cells became elongated and tended to be rectangular, resembling those on the seta. No stomata were found at the base of the capsule.

The peristome is diplolepideous, well developed with both exostome and endostome present and it is Isobryalean in structure. There were sixteen exostome teeth, linear-lanceolate in shape and varying in length from 350um (P. flexicaulis) to 650um (P. crocea). In all species the teeth were variously papillose and their structure appeared similar (Fig. 36.7 p. 136). Alternating with exostome teeth were long thin and fragile endostome segments, one cell wide (P. crocea and P. amblyacis) with papillae.

The operculum was conical with a long curved rostrum (Figs 14, 30 pp. 111, 118) sometimes as long as the capsule. The region of the annulus needs further study when more fertile material becomes available. Studies on the genus Papillaria 66

d) Spores

Spores were typically rounded (12-30um diameter) and papillose (Fig. 36.9 p. 136) with no significant differences noted among the species.

Cytology

The only chromosome numbers reported for Australian species of Papillaria were by Ramsay (1967, 1974):

P. amblyacis n=ll,n=22

P. flavolimbata n = 11 P. flexicau/is n = 11

These chromosome numbers, from Ramsay's publications, have been cited by

Gangulee (1976), Scott & Stone (1976) and Fritsch (1982).

Voucher specimens on which Ramsay (1967) based her chromosome reports were re-examined and the collection annotated as P. jlexicaulis was in fact P. crocea.

Asexual Reproduction

Capsules of Papillaria species were not common, especially those of P. jlexicaulis, yet the genus is common and widespread in suitable habitats. Evidence likely to indicate asexual reproduction was the abundance of rhizoids on tips of branches in some species, e.g. in P. jlexicaulis numerous and dense patches of rhizoids were noted near the ends of the branches. A fragment Studies on the genus Papillaria 67

dislodged or transported by birds or wind could conceivably grow if deposited in a suitable habitat or substratum.

I have noted that Papillaria, because of its trailing habit, was favoured by some bird species [eastern yellow rob bin - Eopsaltria australis (White)] for nests. This method of transporting stems and branches could be a suitable method for the establishment of new colonies. Studies on the genus Papillaria 68

ECOLOGY

In the tropical and sub-tropical ram forests in Australia the most common species of Papillaria to be found are P. amblyacis, P. crocea and P. flexicaulis. Rarer tropical species are P. intricata, P. leuconeura and P. zeloflexicaulis. However, P. flavolimbata prefers the temperate and Nothofagus forests where it can develop into extensive, pure, trailing colonies. Papillaria amblyacis is the only other species extending to the Nothofagus forest on the Australian mainland. In all vegetation types the preferred substrata are tree branches and trunks, shrub branches and stems.

Papillaria species are often found outside these forests, on the margin of the wet sclerophyll forest. P. amblyacis and P. crocea have been reported from scrubby vegetation, from reasonably exposed areas and on rocks, but then generally near water courses or swamps.

In localities with a suitable combination of humidity, mist, cloudiness and ram, Papillaria species can occur in large colonies in relatively exposed situations. Suitable moisture conditions in the tropical and sub-tropical forests allow for the development of sizeable colonies on upper tree trunks and in tree crowns. In the cool temperate forests on the north central coast of New South

Wales P. amblyacis is the most common species where it can form large colonies on the upper trunks, but this species has rarely been reported from tree crowns. Papillaria species prefer some light and in the denser darker forests this genus is quite rare on the lower tree trunks and shrubs, being confined to the more exposed branches in the tree crown. However, on exotic street trees at Mount Wilson in New South Wales, trunks and branches are draped with trailing, often mixed, masses of P. amblyacis, P. crocea and P. flexicaulis. In the township of Eungella (Queensland) P. amblyacis was collected from a planted palm near a Studies on the genus Papillaria 69

residence. Luxuriant, pure colonies of P. crocea have been found on the sparse remnant vegetation between the road and paddocks on the Atherton Tableland in Queensland. On Norfolk Island P. crocea is found only on the highest ridges where there is ample moisture enabling large colonies to develop.

P. crocea, which is the most common and widespread species, shows a preference for tree trunks although in some localities dense colonies can also develop on surrounding vines, shrubs and large rock outcrops (Misty Mountain Road and Bimberamala Creek, Currowan State Forest, southern New South

Wales). It appears that this is the most successful of the Papillaria species in establishing new colonies in variable environments. One collection was obtained from a ten year old Ficus dammaropsis Diels in an Atherton orchard.

Besides occurring in rain forest, Papillaria flexicaulis, can be found in dry sclerophyll forest and dry vine thickets, but rarely in open woodland. So far P. flexicaulis and P. crocea are the only species reported from limestone outcrops, but all species can occur on various other rock types. P. flavolimbata has been collected rarely from rocks, but near Greenwich Park (32km NE of Goulburn, New South Wales) it is found only on shaded conglomerate. The surrounding vegetation is dry sclerophyll forest which is affected by bushfires. It appears that the reduction of rain forests and the resulting drier conditions has forced some

Papillaria species to extend into sclerophyll forests. This may indicate a slow evolutionary change to cope with more xerophytic conditions.

P. flexicaulis has seldom been reported from tree roots, vines or road cuttings which the other three species can colonize. Ferns seldom act as hosts for Papillaria species, but P. amblyacis has been collected off Lastreopsis in Queensland (Blake 20417A) and Dryopteris in New South Wales (NSW M6), Studies on the genus Papillaria 70

while Papillaria flavolimbata was found on Dicksonia stems in Victoria (HS 35529) and in New South Wales (HS 31672).

From available data it appears that Papillaria species do not exhibit preferences for particular woody shrub or tree species. P. amblyacis and P. crocea have been reported on Araucaria cunninghamii (conifer) at Atherton in northern Queensland, while the latter moss species is common on A. heterophylla on Norfolk Island. As conifers are not common in Australian rain forest or wet sclerophyll forest, and most collection data are poor, no reliable conclusions can be reached. I have noted extremely few bryophytes on Agathis in Australia and Papua New Guinea because the bark is so unstable. Thus it is most unlikely that Papillaria species will colonize that genus.

The altitudinal range of Papillaria species is related to the latitude, with the altitude being higher in low latitudes. In Victoria the highest locality recorded for P. crocea is 270m above sea level while in northern New South Wales it has been reported from as high as 1200m. A similar variation is exhibited by P. flavolimbata which has been reported up to 800m above sea level in Victoria and Tasmania, but in New South Wales it reaches 1500m. P. flexicaulis is more commonly found at lower altitudes, the upper recorded limit being 800m. The only reported localilty for P. intricata was from the lowlands. Similarly P. zeloflexicaulis occurs in lowland forest and has not been reported above 200m. Poorer or more open rain forest is the habitat for P. leuconeura. It is pendant from twigs, vines and has also been collected off a fallen tree branch. The altitudinal range is 550-1140m.

Very often several Papillaria species grow intermixed in the larger colonies, especially in the moister areas where competition for space is greatest. Studies on the genus Papillaria 71

Other Meteoriaceae such as Barbella nitens also often grow mixed with Papillaria species.

Altitudinal, habit and habitat details are often missing on labels of both early and more recent collections. This unfortunately leaves a large gap in available data making it difficult to determine accurately habit and habitat preferences. This in turn makes it extremely difficult to summarize, with certainty, the ecological preferences or to make comparisons with data from more recent collections. It is thus almost impossible to formulate conservation strategies. Label data are important and collectors should be encouraged to include more than collector, plant name and a vague locality.

Rev. W.W. Watts in his ministry in northern New South Wales, regularly . visited the scattered timber settlements of the Richmond River Valley (Ramsay 1980). In his journals W.W. Watts mentions that he collected regularly from the crowns of felled trees (pers. comm. H.P. Ramsay). Unfortunately, as he seldom recorded detailed information on his labels, it is impossible to tell from which part of a shrub, tree or tree crown any particular collection originated. Studies on the genus Papillaria 72

DISTRIBUTION

Most of the tropical Australian moss species occur on the eastern side of the Great Dividing Range because of the higher rainfall giving rise to rain forests (Fig. 3). The distribution of Papillaria is therefore generally on the coastal side of the Divide on the mainland and extends into Tasmania ( see section on Ecology). Reports of Papi/laria west of the Divide are few, most occurring close to the Divide. Examples include collections of P. crocea and P. jlavolimbata from the upper Ovens Valley of Victoria. The former has not been recollected since the original collection was made by Mrs McCann during the last century while there is only one recent collection of the latter. These two species together with P. jlexicaulis, have also been collected from the Nandewar Range (Mts Kaputar and Lindsay) in northern New South Wales. The most westerly, but still on the coastal side, report of Papillaria (P. jlavolimbata) is from the upper Calder River in the Otway Ranges of Victoria.

The present distribution of Papillaria species reflects largely the areas intensively collected or where collectors lived, often in bryologically rich areas. Thus common localities recorded in early literature were those of the prolific collector Rev. W.W. Watts who combed the Cambewarra, Blue Mountains and Richmond River regions of New South Wales. Similarly in Queensland the Atherton Tableland and Lamington National Park on the McPherson Range were popular collecting localities as is borne out by the frequency of reports, while central coastal Queensland is still significantly under-collected, although fewer suitable habitats are available.

P. crocea is the most common species at low to mid altitudes in New South Wales (including Lord Howe Island) and Queensland, being found from

Tasmania to just north of Cairns (Fig. 32 p. 122). P. jlexicaulis has a similar Studies on the genus Papillaria 73

distribution, but surprisingly has not yet been reported from Victoria (Fig. 41 p. 153). This species is not as common as P. crocea, nor does it form such large colonies. P. flexicau/is can survive in drier forests and vine thickets and therefore extends further inland than the other species as was shown by collections from near Emerald, the Blackdown Tableland and Mt Surprise in

Queensland. Norris & Koponen (1985) commented on the fact that this species was found only in the western and drier parts of the Huon Peninsula of Papua New Guinea. My (1981) Papua New Guinea collections confirmed this observation.

As P. flavolimbata prefers cool-temperate forest it is therefore the most common species in Tasmania (Fig. 38 p. 140). This is also the most common species found at higher altitudes where it can withstand frost and snow ( Great Dividing Range, E of Captains Flat, alt. 1220m, New South Wales). Its northern limit is reached in the Bunya Mountains of Queensland.

A more tropical species is P. leuconeura which occurs from just north of Cairns (Mt Lewis t6°35'S) to The Great Dividing Range west of Cardwell. However, two collections have been reported from the McPherson Range on the Queensland-New South Wales border. P. amblyacis, a very similar species, extends from Mt Lewis in northeast Queensland to Mt Dromedary in southern New South Wales (Fig. 26 p. 108). Both P. amblyacis and P. /euconeura are not common in north Queensland but the latter predominates north of Ingham. The majority of P. amblyacis collections studied originated from Batemans Bay to just north of Brisbane.

In Australia P. intricata, P. /euconeura and P. zelojlexicaulis are predominantly tropical species. The only known collection of P. intricata was collected from near Rockhampton in 1866. Papillaria zeloflexicaulis extends Studies on the genus Papillaria 74

from Eumundi in southern Queensland to Wauchope in New South Wales. One collection from further south was annotated as being doubtfully collected from Wyong (Watts 9505). The majority of the reports are from the Watts' collections from northern New South Wales.

The pressures for logging and clearing ram forests in the past have destroyed or damaged many Papillaria habitats. This could, for instance, be the reason why:

1) P. flavolimbata is not represented m recent collections from southeastern Tasmania; 2) P. amblyacis, crocea and P. flexicaulis have not been reported for Tasmania this century; 3) there are no reports of P. flexicaulis from Victoria; 4) there is only one recent collection of Papillaria from the upper Ovens River of Victoria; 5) there are only two recent collections of P. zeloflexicau/is from New South Wales and none from Queensland; 6) only one collection of P. intricata 1s known from Australia (Queensland).

As P. flexicaulis, P. intricata and P. zeloflexicaulis tend to inhabit lower altitudes their rarety could well be the result of land clearing. Touw (1982:476) makes a strong case for exploring the tropical lowland vegetation in Malesia which is under greater threat than any other zone. In Australia a similar priority should be given to studies on the remaining tropical lowland forests. 75

SECTION III

STUDIES ON OTHER GENERA 76

SECTION III STUDIES ON OTHER GENERA

INTRODUCTION

This section deals briefly with the history of each genus and its species in the family Meteoriaceae that are represented in Australia. World distribution of each genus and species is presented for each genus on a map subdivided into regions used by Wijk et al. (1959).

HISTORY AND DISTRIBUTION

1 Aerobryopsis Fleisch.

Fleischer (1905:304) created the genus Aerobryopsis while studying the mosses of the Dutch East Indies (now Indonesia), but he did not assign it to any family. He based the genus on the section of Eriocladium of the genus Aerobryum which had been published by C. Mueller (1876).

Aerobryopsis contains 16 species (Wijk et al. 1959, Noguchi 1976) and is well represented in southern Africa as well as extending from India to the

Pacific (Fig. 16). Only the widespread southeast Asian species, A. wallichii, extends into northeastern Queensland where preferred habitats are rain forests, but it can also be found on the margins of grasslands and Eucalyptus woodlands, if sufficient moisture is present. It has been commonly reported from tree trunks, shrubs and rocks from creek sides. Studies on other genera 77

The first collection of this genus appears to be that by C.J. Wild from near Cairns in 1890. Also one undated F.M. Bailey pre 1892 collection was noted .

..... Qc. 2/4

Oc.

Aust.2

Fig. 16. Distribution of Aerobryopsis species before this revision (number of endemics above slash, total number of species below; present Australian distribution 0/1).

However, the first report of Aerobryopsis for Australia was by Fleischer (1905) where he cites A. bauerae Fleisch. from a C. Mueller manuscript name of Eriocladium bauerae C. Muell. He cited no collections or descriptions and hence both names are nomina nuda. This report was also cited by Brotherus (1906, 1925). Studies on other genera 78

In their paper on 'Mosses of north Queensland' Brotherus & Watts (1918) reported Aerobryopsis sigmatophylla Broth. & Watts claiming it to be quite common. Again no collections were cited nor was a description given of this species and therefore this is also a nomen nudum.

An article on north Queensland mosses by Dixon (1942) led to the first

Australian report of A. longissima (Dozy & Molk.) Fleisch. Noguchi (1976) found this species to be a synonym of A. wallichii, but he did not report its occurrence in Australia. This synonymy was confirmed by Norris & Koponen (1985) who did report its occurrence for Australia.

Thus in Australia only P. wallichii is now recognized.

2 Barbella Fleisch.

The genus Barbella was created by Brotherus (1906), being based on

Pilotrichella subsection Barbella, which had been published by C. Mueller in 1896.

Barbella is a tropical genus which has its greatest diversity in southeast Asia (Fig. 17). The genus comprises 28 species (Crosby 1979, Noguchi 1976); none are found in Africa, while B. strongylensis is the only member of this family to reach Europe. The Australian species are rain forest mosses, although B. nitens also occurs in wet sclerophyll, Nothofagus and temperate forests reaching south to Tasmania.

The first Australian specimen of Barbella was described by C. Mueller

(1874) as Neckera trichophoroides Hpe ex C. Muell., based on a collection from Mt Gower on Lord Howe Island. The collection date and collector are Studies on other genera 79

unknown. Although Fleischer (1908) placed this species into synonymy with Barbella enervis, Brotherus & Watts (1918) still reported it for Queensland under its original name.

C. Mueller (in Geheeb 1876) published Meteorium dicladioides from the

Sydney area where it was collected by Dr Kayser, a Bavarian (German) visitor. This name was attributed to C. Mueller, but it is a nomen nudum. M dicladioides was placed into Pi/otrichella trichophoroides (C. Muell.) Jaeg. by Geheeb (1897) but Fleischer (1908) recognized this name as another synonym of Barbella enervis .

...... Qc. 2/3

Oc. .;

Aust.2 0/1

Fig. 17. Distribution of Barbella species before this revision (number of endemics above slash, total number of species below; present Australian distribution 0/2). Studies on other genera 80

B. enervis was reported first for Australia by Fleischer (1908) and then by Brotherus & Watts (1915) for a collection from Mt Gower on Lord Howe Island. This name was recognized until 1976 when Gangulee pointed out that this widespread species was in fact the same as the earlier named Caribbean and central American moss B. cubensis. This synonymy was confirmed by Norris & Koponen (1985).

J.D. Hooker & W. Wilson (in Wilson 1854) described Meteorium nitens naming a Sinclair collection from the North Island of New Zealand as a type. This species was transferred to Papillaria in 1952 by Sainsbury. The first Australian collection of this species was by F.v. Mueller at Rockingham Bay in 1865 and one in the following year at Rockhampton by A. Dietrich.

Brotherus (1916) described P. nitidiuscula, citing 19 of Watts' collections from northern New South Wales as syntypes. In his 'Studies' Dixon (1927) described P. nitidiuscula, on the assumption that the Brotherus taxon was only a manuscript name. Dixon cited a Watts collection from Ballina as the type. P. nitidiuscula was later recognized as being synonymous with P. nitens by Sainsbury (1955b ). In retrospect it is surprising that Dixon did not recognise that these two species were the same taxon.

More recently Noguchi (1985) concluded that P. nitens has greater affinities with Barbella because of the following characteristics:

1) sporophyte has greater similarities to B. pendula (Sull.) Fleisch. than to any Papillaria; 2) areolation of leaves and especially the papillae on the leaf cells are also similar to Barbella pendula; 3) basal angles of the leaves in B. nitens are shortly decurrent, neither rounded nor auriculate as they are in Papillaria species. Studies on other genera 81

He has admitted (pers. comm. 1986) however, that he examined few Australian collections.

A moss found by F.v. Mueller at Tarwin in Victoria was named by Mitten (1860) as Meteorium fulvum. Jaeger (in Jaeger & Sauerbeck 1877) transferred this to Papillaria and Hampe (1881) to Pilotrichella. An isotype from the Melbourne Herbarium (MEL) had a small annotation by J.H. Willis (1952) where he correctly recognized this species as a synonym of Barbella nitens, but his conclusion was never published. It remained recognized as a Papillaria, but

Scott & Stone (1976) suggested that this moss might be P. nitens. This study confirmed their suspicion.

An undated collection from the Blue Mountains by an unknown collector, was named by C. Mueller (in Hampe 1872) as Neckera dimorpha.

Jaeger (in Jaeger & Sauerbeck 1879) placed this in Pilotrichella while Mitten

(1882) considered this moss to be aMeteorium, but this change was not followed and it is now considered to be a synonym of Barbella nitens.

Brotherus (1900) described Pilotrichella perpinnata which he based on a 1897 collection made by F. Whitteron in Queensland. Brotherus (1906) then transferred this species to Barbella.

Thus in Australia the following species of Barbella were accepted before this revision: B. ene,vis; B. nitens; and B. perpinnata. Studies on other genera 82

3 Floribundaria Fleisch.

Fleischer (1905) created this genus, basing it on Mueller's Papillaria section Floribundaria which was a nomen nudum (Wijk et al. 1962:323).

Presently this tropical genus has 21 species (Noguchi 1976, Crosby 1977, Wijk et al. 1962, Tan 1984) the majority of which occur in central Africa and from India to the Pacific (Fig. 18). All Australian species are rare rain forest dwellers from north Queensland. In fact F. robustula is so far known only from the type and one other collection.

The first species of the genus Floribundaria was named by Dozy & Molkenboer in 1844 based on Dutch East Indies (Indonesian) material, as

Leskea floribunda Dozy & Molk. The same authors transferred this species to Meteorium in 1848 (Wijk et al. 1964:353). The most recently described species,

Floribundaria torquata Wang & Lin was named from a Taiwanese collection in 1975 (Crosby 1977). The first Australian report of F. floribunda was from north

Queensland by Brotherus & Watts (1918) being based on four Watts' collections. The next report was by van Zanten (1964). This and subsequent citations were all secondary reports.

Fleischer (1908) recognized F. pseudofloribunda as a distinct species close to F. floribunda. The first Australian report was by Brotherus & Watts (1918) based on five Watts' collections. There have been no subsequent primary reports. Studies on other genera 83

Oc. 0/4

Oc. .;

Aust.2

Fig 18. Distribution of Floribundaria species before this revision (number of endemics above slash, total number of species below; present Australian distribution 0/3).

The third species, F. robustula, was published by Brotherus & Watts (1918) based on a Watts' collection from Malanda in Queensland, gathered during July 1913. This species was found mixed among numerous other mosses, some of which were Meteoriaceae. There has been only one other report of this species.

F. walkeri was reported by Tan (1984), who redetermined two of Watts' north Queensland F. jloribunda collections.

Thus four species were recognized in Australia before this revision: F. jloribunda; F. pseudojloribunda; F. robustula; and F. walkeri. Studies on other genera 84

4 Meteoriopsis Fleisch. ex Broth.

The first species treated now as Meteoriopsis was Pilotrichum reclinatum described by C. Mueller in 1854 (Wijk et al. 1967). The present genus was circumscribed by Brotherus (1906). He based this on a name published by Fleischer in 1902 (Musci Frond. Arch. Ind. ser. 5:236) without a description and therefore a nomen invalidum.

Meteoriopsis with five species (Fig. 19), is best represented in southeast Asia (Manuel 1977b) by two widespread species, M. squarrosa (Hook.) Fleisch. ex Broth. and M reclinata, the latter species also extending to southern Japan (Kyushu), Australia and Vanuatu (New Hebrides). The remaining three species are endemics: M. undulata Horik. et Nog. confined to Japan; M. conanensis

Goa described for Tibet in 1979 (Crosby & Bauer 1981); and M. novoguineensis Nog. restricted to Papua New Guinea (Noguchi 1986).

Manuel (1977a) created the genus Zelometeorium to accommodate the species from Meteoriopsis section Squarridium C. Muell. ex Broth. which have smooth leaf cells and completely papillose exostome teeth. Manuel transferred four species to his new genus while also describing one new species. All of these species are restricted to South America with one other in central Africa. Studies on other genera H5

Fig. 19. Distribution of Meteoriopsis species before this revision (number of endemics above slash, total number of species below; present Australian distribution 1/2).

The Australian reports of Meteoriopsis reclinata are quite recent, commencing with Brotherus & Watts (1918) where they cited Watts' 1913 collections. This was followed Dixon (1942) who cited a Sherrin collection of 1922. No subsequent primary reports were noted. This species appears to be uncommon and is restricted to the Atherton Plateau of north Queensland or its fringes.

Thus M. reclinata was the only species reported for Australia prior to these studies. Studies on other genera 86

5 Meteorium Dozy & Molk.

The genus Meteorium was created by Dozy and Molkenboer in 1854 (Wijk et al. 1964:349) based on Pilotrichum, a Bridel-Brideri sub-genus of 1827. The first species in the present day genus was named by Bridel-Brideri (1827) as Isothecium buchananii. The earliest Australian records of Meteorium were those of Wilson (1854, 1859) where he described species that are now placed mainly in Papillaria and in Barbella nitens. The most recently described species was Meteorium filifonne Nog. (Noguchi 1986) for Papua New Guinea.

Meteorium is a moss genus of the tropics with 27 species (Bauer & Crosby 1986, Crosby 1979, Crosby & Bauer 1983, Noguchi 1976, 1986) which are best represented in tropical America and southeast Asia (Fig. 20). In Australia the widespread southeast Asian species of M. buchananii is common in tropical north Queensland where it prefers the rain forest or its moister fringes. Single collections have been seen from Rockingham Bay, Brisbane River and the McPherson Range near the New South Wales border.

A Bailey collection from Brisbane River, Queensland, was named by Mitten (1882) as Meteorium compressum. Kindberg (1891) transferred this species to Papillaria, but this change was not accepted by subsequent bryologists.

Meteorium cinclidotus, a C. Mueller manuscript name, was cited by Bailey (1883) without further information, therefore this tax.on is a nomen nudum. Studies on other genera 87

...... , Qc, 0/2

Oc.

Aust.2

Ant.

Fig. 20. Distribution of Meteorium species before this revision (number of endemics above the slash, total number of species below; present Australian distribution 0/1).

The first Australian report of the present M buchananii was by Brotherus (1888) when he described Papillaria baileyi based on an undated collection gathered by F.M. Bailey beside the Brisbane River. Also in the same year Bailey included Meteorium baileyi Broth. ex Bailey in a supplement to his 1883 work 'A synopsis of the Queensland flora'. This name was based on an undated collection made by K. Broadbent from Rockingham Bay and is considered a nomen nudum. Brotherus transferred Papillaria baileyi to Meteorium baileyi in 1906. M. miquelianum was first reported for Australia by Brotherus & Watts (1918) where they also reported M. baileyi. They claimed Studies on other genera 88

both species to be common on the Atherton Plateau and it seems surprising that Brotherus did not recognize the similarities between the species. Both species were reported by several authors subsequently but none based their reports on examined collections. Noguchi (1976) split this species into two sub-species with the new sub-species restricted to China, Taiwan and Japan.

Thus three species were recognized in Australia before this revision: M. baileyi; M. buchananii; and M. compressum.

6 Weymouthia Broth.

Brotherus (1906) created this genus which he named in honour of William Anderson Weymouth (1842-1928), a prominent Tasmanian bryologist.

Weymouthia is a genus of mosses found mostly in temperate forests, thus differing from the rest of the genera in this family which are predominantly tropical mosses. It also differs from the other genera in the presence of conspicuous alar cells, the absence of papillae and the costa being inconspicuous or absent. This small genus has only three species and one variety, with two species distributed in southeast Australia, New Zealand and South America, while the third species is endemic to South America (Fig. 21).

The first species of Weymouthia was described by Hedwig as Leskea mollis in 1801 based on New Zealand collections, but not being assigned to any family. Schwaegrichen, in 1816, named Hypnum cochlearifolium which was collected by J.J. Labillardiere during his voyage to Tasmania (Apr. 1792 & Jan. 1793 at Storm Bay). Again no family was designated. Robert Brown also collected both taxa from unknown locations in his travels with Matthew Flinders between 1801-1803. Studies on other genera 89

C. Mueller (1850) transferred Leskea mollis to Neckera mollis in the tribe Hypnoideae, but in 1854 Wilson renamed this as Meteorium molle and in 1859 added the variety majus which he placed in the tribe Pilotricheae. In the same work Wilson assigned Hypnum cochlearifolium also to the tribe Hypnoideae. Both species had been transferred to Isothecium by Mitten (1856), but in 1860 he considered these were best placed in the genus Stereodon in the family Hypnaceae. In the same year Hampe named Neckera billardieri which he based on a F.v. Mueller collection from Apollo Bay in Victoria. Reichardt (1870) made the combination Coelidium cochlearifolium. Stereodon mollis and Neckera billardieri were later transferred to Pilotrichella in the family Pilotricheae by

Jaeger (in Jaeger & Sauerbeck 1887). However, he followed Reichardt by retaining Coelidium cochlearifolium in the family Hypnaceae. Mitten (1882) made the combinations of Porotrichum cochlearifolium and Meteorium billardierii placing them in the Neckeraceae. In 1896 Lindberg (in Paris 1897) made the combination Lembophyllum cochlearifolium while Brotherus (1906) transferred Pilotrichella mollis and Meteorium billardieri to his new genus Weymouthia.

The next person to examine these species was Dixon (1927) who made the combination W. cochlearifolia to which he added W. cochlearifolia var. billardieri after a lengthy discussion. The only other taxonomic comments were those made by Sainsbury (1955a, 1955b) where he considered that the variety billardierii was too weak to be retained. Scott & Stone (1976) described both species and explained how to separate them from mosses with a similar appearance (Lembophyllum divulsum (Hook.f. & Wils.) Lindb. and Acrocladium chlamydophyllum C. Muell. & Broth.). Studies on other genera 90

Thus in Australia two species of Weymouthia were recognized: W. cochlearifolia; and W. mollis.

, .... Oc.

Oc. ..

Aust.2 0/2

Ant.

Fig. 21. Distribution of Weymouthia species before this revision (number of endemics above the slash, total number of species below; present Australian distribution 0/2). 91

SECTION IV

TAXONOMIC TREATMENT OF THE FAMILY METEORIACEAE 92

SECTION IV TAXONOMIC TREATMENT OF THE FAMILY METEORIACEAE

INTRODUCTION

This section deals with the taxonomic features of the family Meteoriaceae; all the genera and species in Australia are described in full, with keys provided to genera and species. Distribution maps are included based on all known collections available for this study. Illustrations, including some scanning electron micrographs, are presented for all Papillaria species. Comments are made on the distribution, ecology and known substrata for each species. Type collections are discussed, as well as unusual features or variations within each species and similarities to other species. A comprehensive listing of all published illustrations is included.

THE FAMILY METEORIACEAE Kindb. Eur. N. Amer. Bryin. 7:1897

Type Genus: Meteorium (Brid.) Dozy & Molk., Musci Frond. Ined. Archip. Ind. 157.1854. Pilotrichum subg. Meteorium Brid., Bryol. Univ. 2:264.1827.

Pleurocarpus plants, slender to robust, usually pendant from trees. Primary stems with or without very small central strand, creeping, filiform; secondary stems distantly and irregularly branched, elongate, flexuose, pendant and densely foliose. Older leaves often dark brown to black. Leaves generally papillose, ovate-lanceolate, acuminate, often with a cordate base, margin generally denticulate or dentate, costa slender usually to 1/2-2/3 of the leaf length, seldom more or absent, leaf cells prosenchymatose. Alar regions rarely Taxonomic treatment of the Meterioriaceae 93

well developed and inferior cells hyaline. Dioicous, seldom autoicous. Perigonial and perichaetial leaves differentiated and often with paraphyses. Calyptra cucullate or mitrate, long often hairy. Capsules smooth, erect, rarely sub-erect, mostly ovate to oblong, often exerted on a short, smooth seta. Operculum high, conical or rostrate and generally curved. Peristome diplolepideous, double, exostome teeth 16, linear-lanceolate, densely papillose or only so in the upper half. Endostome segments alternating with exostome, papillose, linear to filiform, mostly perforated and as long as the exostome teeth. Basal membrane high or low. Spores spherical to ovate, 12-35um and papillose.

Chromosome numbers: n = 6, 9, 10, 11, 22 (see also Table 3 p. 16).

This family is usually classified into two sub-families:

la Leaves oval, cucullate, rarely hair pointed, rarely papillose. Costa short, double or none, alar cells.± prominent ...... PILOTRICHELLOIDEAE

lb Leaves narrow, oval to laceolate often with a hair point and a broad base, generally papillose. Costa single, alar cells hardly differentiated ...... METEORIOIDEAE

The sub-family Pilotrichelloideae is represented m Australia by the single genus Weymouthia with two species.

The sub-family Meteorioideae is more common in Australia, being represented by fifteen species in the following genera: Aerobryopsis, Barbella, Floribundaria, Meteoriopsis, Meteorium and Papillaria. Taxonomic treatment of the Meterioriaceae 94

KEY to AUSTRALIAN METEORIACEAE

la Leaves concave, orbicular to broadly ovate, apex.± obtuse ...... (7) WEYMOUTHIA lb Leaves ovate, ovate-oblong to linear, acuminate ...... 2

2a Cells in.± auriculate leaf base in divergent rows ...... 3 2b Cells in vertical rows, leaf base without auricles ...... 5

3a Leaves generally 3-plicate, cells unipapillose ...... (6) METEORIUM 3b Leaves if plicate, with 1 or 2 only, cells multipapillose ...... 4

4a Cells weakly papillose, plants light or golden-green ...... :...... (3b) BARBELLA NITENS 4b Cells densely papillose, plants darker green ...... (1) PAPILLARIA

5a Leaves narrow, lanceolate, apex filiform or flexuose-capillaceous, seldom acuminate ...... 6 5b Leaves broader, ovate, always acuminate ...... 7

6a Leaves widely spreading, multipapillose, cell outline.± obscure ...... ( 4) FLORIBUNDARIA 6b Leaves.± appressed, scarcely papillose, cell outline clear ...... (3a) BARBELLA CUBENSIS

7a Leaves appressed, apex transversely undulate ...... (2) AEROBRYOPSIS 7b Leaves spreading, apex straight ...... (5) METEORIOPSIS Taxonomic treatment of the Meterioriaceae 95

Fertile collections are not common in this family, in fact no fertile Australian collections were found for Barbella cubensis, Floribundaria, Aerobryopsis,

Meteoriopsis and Meteorium. The lack of fertile specimens from tropical Australia is surpnsmg. A study into the reasons for this could provide a valuable and interesting insight into the reproductive strategies of tropical mosses.

The Meteoriaceae has considerable structural diversity as pointed out by Buck & Vitt (1986:52) who suggested that this 'diversity may indicate that the family needs to be subdivided into more homogenous families'. This was further evidenced by Barbella comes (Griff.) Broth. being transferred to lsotheciopsis (Noguchi 1970) in the Lembophyllaceae. Buck (1980), however considered this genus more appropriate for Meteoriaceae together with Dolichomitriopsis. However, no formal transfer of species has taken place. Noguchi (1976:349)

also expressed doubts about the familial position of Aerobryum which he felt was inappropriate for Meteoriaceae.

Weymouthia, especially W. cochlearifolia is very similar to, and has been misdetermined as Lembophyllum. Perhaps the position of Weymouthia, which is not tropical, should be kept in mind when the Lembophyllaceae is revised.

The sub-family Pilotrichelloideae has a suite of good characters which could well merit it being raised to family level. A scanning electron microscope study of the sporophytic characters in Meteoriaceae might assist in the subdivision of the family and delineation of the genera. The genus Papillaria 96

I. The genus PAPILLARIA (C. Muell.) Lor., Moosstudien 165.1864

Type: Hypnum nigrescens Sw. ex Hedw., Spee. Muse; 250.65,f.1-4.1801 [= P. nigrescens (Sw. ex Hedw.) Jaeg.] Neckera subsect. Papillaria C. Muell., Syn. 2: 134.1851. Syn: Papillaria (C. Muell.) C. Muell., Oefvers. Forh. Kongl. Svenska Vetensk.-Akad. 33(4 ):34.1876.

Slender or robust plants, more slender or filiform when pendant, without lustre, dull, green or dark greenish, mostly corticulous on trunks or branches, often pendant in long intricate masses but also on rocks. Branching generally at right angles to the stem, branch leaves loosely or densely appressed. Leaves spirally arranged, ovate, oblong-lanceolate or triangular, often strongly auriculate, insertion narrow, amplexicaul, often with a cordate base and an accuminate tip. Stem leaves slightly narrower and longer. Margin variable, inconspicuously dentate near base.

Smooth, single, red to reddish-brown costa, extending in leaf from 1/2 way to leaf-tip. Lamina cells incrassate, multi-papillose, papillae often obscuring the cells. Median cells elliptic or oblong-rhomboidal, upper and marginal cells very similar. Basal juxtacostal cells or those at base not papillose, but hyaline, longer, rectangular and thick walled. Cells in auricles rhomboidal or linear-rhomboidal, arranged in divergent or arcuate rows.

Rhizoids smooth, reddish, fused at base, margins thickened.

Dioicous. Perigonia rounded, perichaetia rounded or elongated, both generally on branches. Inner perigonial leaves entire or distantly irregular, ovate, oblong to lanceolate, apex acuminate or blunt, papillae weak. Costa weak to 3 / 4 of leaf length or ecostate. Antheridia 5-20 per perigonium, 220- 17ie genus Papillaria 97

400um long, club shaped, surface with irregular quadrate and rectangular cells, dark reddish when mature. Paraphyses as long and as many as antheridia, transparent, uniseriate, multicellular with thick walls. Inner perichaetial leaves entire or denticulate in upper section, apex acuminate, marginal cells often distinct, papillae weak. Costa weak up to 3/ 4 leaf length or ecostate. Archegonia 5-12 per perichaetia, 280-4 lOum long, widest at venter, surface with irregularly quadrate, rectangular or both types of cells, dark reddish when mature, egg and neck canal blackish. Paraphyses as long and as numerous as the archegonia, similar to those in perigonia.

Capsules not common, ovoid or oblong, smooth, exserted on short, erect and smooth seta. Calyptra pilose, split along one side, operculum highly conical, mostly curved. Peristome double, exostome teeth linear-lanceolate, irregular in outline, densely papillose; endostome segments linear to filiform, papillose, same length as exostome, irregular in outline, perforated. Basal membrane low. Spores rounded, medium sized, 12-25um, papillose.

Chromosome numbers: n = 10 (India, Kumar 1973); 11, 22 (Australia, Ramsay 1967, 1974).

E1YMOLOGY: from the Latin, papilla, referring to the nipple like protuberances on the leaf cells.

The genus is divided into two sections. The main differences being (Noguchi 1976):

la Capsules on short seta, inner perichaetial leaves short ...... Sect. Papillaria

lb Capsules immersed, inner perichaetial leaves long ...... Sect. Penicillatae The genus Papillaria 98

Only sect. Papillaria occurs m Australia, Norfolk Island and New Zealand.

Fertile material is not common, especially for P. flexicaulis and P. leuconeura. Papillaria does not fruit prolifically in Australia and the few sporophytes that may be present are easily disguised on short setae on trailing branches.

Perigonia and perichaetia were found on plants collected between April and November. Representative data were obtained for the perigonia of P. amblyacis and P. flexicaulis and for the perichaetia of P. amblyacis and P. zeloflexicaulis. For P. amblyacis, 20 collections with perigonia were noted, of these 9 were collected during September. Six collections with perichaetia were collected between late August and early October. Perichaetia were difficult to find in this species and those that were found were expanding having been fertilised, but for the other species this was not the case. Very immature sporophytes of P. amblyacis were noted in several collections made in November. In all species, except P. zeloflexicaulis, perigonia were more commonly present.

Development data for sporophytes are not available and therefore the growth rates are unknown. However, it was noted that a range from very young developing sporophytes to remnants, where the sporophytes had dropped out, could be found in the same collections of several species. So perhaps sporophyte development is slow and may take up to a year. Most data are available for P. amblyacis and P. crocea in which immature or developing sporophytes were most commonly found during August and September.

Mature capsules generally occurred between April and October while in some species they were also present in November, December and February. As The genus Papillaria 99

most of the Papillaria collections are from New South Wales, the effect of latitude and altitude could not be investigated. Fertile collections outside New South Wales were rare.

KEY to AUSTRALIAN SPECIES of PAPILLARIA

la Costa up to 1/2 leaf length, leaves twisted ...... (5) P. INTRICATA lb Costa more than 2/3 leaf length, leaves straight ...... 2

2a Cells in auricle meeting border at right angles, (Fig. 10 p. 54) margin entire, cells.± uniform ...... 3 2b Cells in auricle parallel to border (Fig. 10 p. 54), margin generally denticulate, cell size variable ...... 4

3a Plants very slender, leaves v-shaped in cross section, less than 0.5mm wide, costa strong and slightly twisted ...... (7) P. ZELOFLEXICAULIS 3b Plants slender, leaves shallowly rounded, leaves wider than 0.5mm, costa straight and thin ...... (4) P. FLEXICAULIS

4a Margin recurved, auricles with a broad border of long narrow cells ...... (3) P. FLAVOLIMBATA 4b Margin straight, without a broad border ...... 5

5a Upper and median cells at least lOx as long as wide, very incrassate,.± sigmoid ...... (6) P. LEUCONEURA 5b Upper and median cells less than 6x as long as wide, slightly incrassate, straight ...... 6

6a Leaf smoothly curved when dry, costa usually straight, if twisted then forked ...... (1) P. AMBLYACIS 6b Leaf undulate and long grooved, costa twisted, without a fork ...... (2) P. CROCEA The genus Papillaria 100

1 PAPILLARIA AMBLYACIS (C. Muell.) Jaeg., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1875-76:267.1877 (Ad. 2:171).

Neckera amblyacis C. Muell., Linnaea 36:521.1870 Type: 'Leichhardt, New South Wales'. LECTOTYPE: BM! Pilotrichella amblyacis (C. Muell.) Hpe, Fragm. Phytogr. Austral. Suppl. 11:50.1881 ('-acris'). Meteorium amblyacis (C. Muell.) Mitt., Trans. & Proc. Roy. Soc. Victoria 19:82.1882. Neckera squamata Hpe, Linnaea 36:522.1870, syn. nov. Type: no data, ex Herb. Com. Solms. HOLOTYPE: BM! Papillaria squamata (Hpe) Hpe, Fragm. Phytogr. Austral. Suppl. 11:112.1881, syn. nov. Neckera eavesiana Hpe, Linnaea 40:319.1876, syn. nov. Type: 'Eaves, subtropical eastern Australia'. HOLOTYPE: BM!; ISOTYPES: MEL (32239)!, H.! Papillaria eavesiana (Hpe) Jaeg., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1875-76:266.1877 (Ad. 2:170), syn. nov. Pilotrichella eavesiana (Hpe) Hpe, Fragm. Phytogr. Austral. Suppl. 11:50.1881, syn. nov. Meteorium eavesianum (Hpe) Mitt., Trans. & Proc. Roy. Soc. Victoria 19:82.1882, syn. nov. Papillaria eavesii Kindb., Enum. Bryin. Exot. 27: 1888, nom. illeg. incl. spec. prior.

Robust plants, yellowish to brownish-green. Primary stems to 2cm long, secondary stems terete, densely leaved, up to 30cm long with pinnate or irregular branching. Branches flexuose, terete and julaceus or at times cuspidate, especially on new growth, attenuate at apex. Leaves erect, densely imbricated, only slightly erecto-patent when wet. Stems a very light greenish colour which darken with age to a brownish-black. The genus Papillaria 101

3

...· . ·. ·. : ~

...... ' .- ...... ___. .. .. ~-·-

I00,11111

Fig. 22. Papillaria amblyacis (C. Muell.) Jaeg. Branch leaves: 1,4. Constable 5995; 2,5,6. Adams 1917A; 3,7. HS & Curnow 37703.

Branch leaves smooth and convex at back, ovate-oblong to 'fiddle­ shaped' (panduriform), often with nearly parallel sides. Leaves (0.9) 1.1- 1.5( 1.9)x0.5-0. 7(0.8)mm, .± cucullate with a short recwved mucro, base not The genus Papillaria 102

cordate, but broadly auriculate. Leaves sometimes longer with an acuminate tip. Margin variably denticulate in the auricles. Ratio of leaf width to length ea 0.52. Costa generally reaching at least 3/4 leaf length, rarely to tip, at base 30- 40(50)um wide, becoming thinner towards tip. At times costa obscure, often with some forking above mid-leaf and then costa slightly twisted. Costa cells rectangular 33-55x6um. Costa more prominent on older leaves.

Cells sometimes irregular at tip of leaf, incrassate and lacking papillae, but then gradually merging into the normal densely papillose laminal cells with 3-5 small rounded papillae per cell which did not obscure the cell outlines. Cells in upper leaf elliptic to rhomboidal 10.0-15.0x5.0-7.5 um, with strong and irregular thickenings (incrassate) and often sinuose. Median cells very similar, but with a few larger cells. Juxtacostal cells above the base smooth and transparent, porose, elongated, up to 65um long and prosenchymatous, but nearer the base slightly shorter, more rectangular and parenchymatous. In auricles, elliptical cells in divergent rows and more incrassate than those in upper leaves. Auricle cells smooth, with a margin comprising slightly longer and narrower smooth cells that generally reach 3 / 4 leaf length.

Stem leaves noticeably clasping stem, less cucullate, with leaf narrowing to a short acute point. Stem leaves weakly papillose with hardly a visible border and the cells of similar shape and size to those of branch leaves. At base of branches, smaller (0.7mm) circular leaves with few papillae, often one papilla per cell, ecostate and the lower half denticulate. Cells in upper leaf 15x4um, irregularly rhomboidal, incrassate. Basal juxtacostal cells 30x4um covering a larger area than in branch leaves. Auricles with elongated cells 10x4um. Basal leaves not prominent tending to be obscured by the bigger rounded leaves above them and by those on the stem. 171e genus Papi/laria 103

Fig. 23. Papillaria amblyacis (C. Muell.) Jaeg. Branch leaf cells (Watts 10837): 1. Apex; 2. Near mid-leaf; 3. Median marginal cells; 4. Median near mid-leaf; 5. Auricle.

Stems rounded to ovate in cross-section up to 220x170um. Epidermal and outer cortical cells are small 2.5-5.0um varying from rounded to triangular, incrassate forming a band 5 cells wide. Inner cortical cells thin walled and cell size increasing towards the centre where they reach 20.0x12.5um, with the shape quite variable from quadrate, hexagonal to roughly rounded with some very irregular. Smaller cells often scattered among larger cells. Central strand a group of small (2-4um) cells. Inner cortex up to 160xl 10um thick. Branches rounded to ovate in cross-section up to 200xl 70um thick. Epidermal and outer cortical cells slightly larger and less incrassate than in stem. Inner cortex The genus Papillaria 104

1

,.

4 5 6

7 8

Fig. 24. Papi/laria amblyacis (C. Muell.) Jaeg. - Watts 2912: 1. Branch (x27); 2. Branch and sporophyte (x8); 3. Papillose leaf surface (x529); 4. Archegonia (x130); 5. Peristome teeth (x167); 6. Capsule surface (x98); 7. Capsule (x18); 8. Upper capsule with peristome (x64). The genus Papillaria 105

130xl00um thick, with cells nearly the same as those of stem cortex, but not so irregular. Central strand absent. Boundary between golden-brownish outer cortex and clear inner cortex distinct in both the stem and branch. Both the stem and branch sections slightly denticulate.

Rhizoids not common, 12.Sum wide with prosenchymatous cells up to 120um long, with the cell boundaries often difficult to discriminate.

Perigonia covered by branch leaves, not conspicuous. Inner perigonial leaves oblong up to 1000x500um, tip acuminate, ecostate, weak papillae scattered on upper and median cells. Margin entire with smaller rectangular cells. Upper cells hexagonal rarely elongate sigmoid, 37x5um, median cells elongate-rhomboidal, up to 50x7um, basal cells similar but rectangular. Outer perigonial leaves ovate, tip cells hexagonal, 25x7um, ecostate, not papillose. Antherida ..±. 12 per bud, up to 480x160um. Paraphyses up to 500um long and 25um wide, with rectangular cells up to 75um long. Perichaetia small elongate. Inner perichaetial leaves ovate-obovate 910(1100)x450(550)um, ecostate or faint, papillae lacking or very weakly developed. Margin denticulate in upper half, ..±. elongate rectangular marginal cells better developed in the smaller rounded outer perichaetial leaves with denticulate tips. Upper cells elongate, up to 30x5um, median cells elongate-hexagonal 37(105)x5um, quite variable, oblique, hexagonal at base with a row of rectangular cells. Archegonia ..±. 6 per bud, up to 350x70um, neck up to 60um wide, with quadrate cells (15x15um). Neck of growing archegonia appears fragile. Paraphyses as many and as long as the archegonia, up to 20um thick when one cell wide, cells up to 50um long. Lower half of paraphyses two cells wide, nearer base often three cells wide, but then with very small, quadrate and reddish cells. At base of capsules inner perichaetial leaves lanceolate with nearly parallel sides, up to 2.6mm long, apex acuminate, costa nearly to apex in weakly denticulate leaves. Cells in upper and The genus Papillaria 106

mid-leaf 37-50x3um, incrassate, with irregular sides and very few faint papillae. Basal cells rectangular longer, up to 75um, and more regular than the cells above and at times appearing as a border along the margin. Base of the perichaetial leaves with small rounded leaves with elongated rectangular cells_± quadrate along the margin, ecostate with upper margin serrulate. Paraphyses numerous, up to 25um (3 cells) wide, mid-cells up to lO0um long, parenchymatous, at apex incrassate.

Calyptra ea 3 mm long. Capsules ovoid, 1.6-2.0x0.5-1.0mm, setae 2-6mm long. Peristome teeth lanceolate, endostome segments linear, 350-450um long. Operculum not seen. Exothecial cells generally rounded to squarish from 25.0um square to 25.0x7.5um, or irregular in shape. Cells on inner capsule wall similar. Cells on both sufaces difficult to discriminate. Spores ( 12.5) 17 .0- 20.0(25 .0)um.

Chromosome numbers: n = 11, 22 (Ramsay 1967, 1974).

DISTRIBUTION and ECOLOGY: Eastern Australia - Queensland, New South Wales and Tasmania (Fig. 26) including Lord Howe Island; extending also to New Zealand.

Habitats include rain forest, Nothofagus forest and the edges of wet sclerophyll forest. It has also been reported from a swamp, a Tristania dominated river side, on willows beside a creek, windswept bushland and is common on exotic street trees at Mount Wilson (Blue Mountains, New South Wales).

Papillaria amblyacis is an epiphyte on tree trunks and branches, shrub stems and branches, often forming pendulous, pure colonies. It has also been reported in tree crowns, on logs, rocks, road cuttings, vines and ferns The genus Papillaria 107

(Lastreopsis silvestris D.A. Sm. ex Tindale and Dryopteris albovillosa Watts). This moss has also been found epiphytic on the following trees and shrubs:

Acronychia, Araucaria, Argyrodendron, Bursaria, Citriobatus, Ceratopetalum, Cryptocarya, Daphnandra, Doryphora, Drimys, Endiandra, Flindersia, Hedycarya,

Nothofagus, Prostanthera, Quintinia, Orites, Salix and Tristania. Often it is found mixed with other Papillaria species, Barbella species, Meteorium, lichens and liverworts. Papillaria amblyacis has been collected from near sea level to 1300m in New South Wales and to 1200m in Queensland.

Fig. 25. Papillaria amblyacis (C. Muell.) Jaeg., on tree branches on edge of tropical forest, Credition State Forest, west of Mackay, Queensland (HS 37702 - Photo: J. Curnow). Thegenus Papillaria 108

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Fig. 26. Distribution of Papi/laria amblyacis (C. Muell.)Jaeg. The genus Papillaria 109

ETYMOLOGY: from the Greek, ambly = blunt and acis = pointed, alluding to the small pointed leaf tips.

ILLUSTRATIONS: Brotherus 1906:316, 1925:163.

TYPE MATERIAL

The types of Papillaria eavesiana and P. squamata are merely morphological extremes met within the range observed for P. amblyacis. C. Mueller described the leaves of P. amblyacis as being hooded, fiddle-shaped and terminated by a.± short apiculus, but noted that the delicate forms which are more loosely imbricate have longer and larger leaves with tips which are gradually apiculate. The type of P. squamata has leaves that are not hooded, not fiddle-shaped and tips are acute. Similarly the type of P. eavesiana has leaves which are not prominently hooded, but are slightly longer and narrower than those of P. squamata and P. amblyacis, although here the leaves are weakly fiddle-shaped. More significantly the cells and costa are similar in all three specimens. Thus both synonyms (P. squamata and P. eavesiana) which were described by Hampe (1870, 1876) are here considered to be forms encompassed within the range mentioned by C. Mueller.

Minor differences were noted in the branch cross-sections. P. squamata was similar to P. amblyacis except that the cortex was more yellow and the cells were quite irregular. In P. eavesiana there were more thin, narrow cells in the epidermal region which pointed towards the centre, the cells near the branch edge in the epidermis were slightly larger and there was a gradual increase to the bigger cortical cells which again were quite irregular. The genus Papillaria 110

NOTES

There appears to be no correlation of leaf shape with distribution and geography. When the plants were trailing from tree or shrub branches, both the stem and branch leaves were generally longer. However in temperate populations the short recurved mucro predominated. In some cases the leaf apices were long, narrow and curled (HS 38363, Watts 4094) then however the branch leaves were often not julaceous. Branch leaves were noted up to 1.7x0.9mm. Plants with acuminate tips had a few rows of cells at the tip that were shorter and thicker, other cells being long and narrow,.± sigmoid, with the cell walls up to twice the thickness of the cells. As the leaf tip becomes acute the cells tended to be shorter, thicker and appeared more like the normal leaf tip cells. On plants from tree trunks, logs or rocks, the leaf was normally shorter with an apiculus.

Some collections with longer leaves had older irregular leaves resembling those of P. crocea. However, in this species such long leaves were rare and a search of young branches in specimens of this kind revealed the typical smooth curved leaf surface of P. amblyacis. At the base of the branches of P. amblyacis small leaves remained rounded and were conspicuous, while in P. crocea these leaves were not prominent.

Prominent serrations were noted at leaf base in one collection (MEL 1045364).

The costa was faint in some collections (Watts 6360, Whitelegge Ml3015) but its width at base and length remained unaltered.

This species is very similar in gross appearance to P. leuconeura and only

an examination of the laminal cells will separate these two species. In P. 17ie genus Papillaria 111

amblyacis the upper and median leaf cells are elliptic to rhomboidal, up to

15.0x7.5um and slightly incrassate, while in P. leuconeura they are irregularly sigmoid, up to 25x3um and prominently incrassate (Fig. 27). P. leuconeura, very rarely possessed the short recurved mucro of P. amblyacis, which sometimes had an acuminate leaf tip similar to P. leuconeura. The short recurved mucro type leaf of P. amblyacis was smoothly rounded and concave on the back of the leaf while the acuminately tipped leaves of both species were not so rounded or smooth.

1 2

Fig. 27. Upper leaf cells near mid-leaf (x600) . 1. Papillaria amblyacis (Watts 10837); 2. Papillaria leuconeura (Stone 15767).

Hilbrands (1969) suggested thatP. amblyacis was best treated as a variety of P. leuconeura. He claimed that the proposed variety was a robust moss with concave leaves having a mucro. In contrast P. leuconeura was a fine to robust moss with the leaf hardly ever concave and only gradually forming an acuminate The genus Papillaria 112

apex. As the last two sets of characters tend to be variable and intergrade, they are not suitable for segregating the two taxa. Hilbrands (1969) also admitted that there were many intermediate forms which made classification difficult.

Fertile collections were not common, but collections with mature capsules were collected between April and November.

SPECIMENS STUDIED: Australia 409 (47 cited), New Zealand 22.

QUEENSLAND: Mt Lewis, Stone 15767 ( + P. flexicaulis - MELU), 3 Aug. 1979; Windsor Tableland, 1140m, 38km NW of Mossman, rain forest with Agathis robusta, trailing from treelet branches, HS 29717 (CBG, DUKE, H, HO, L, NICH, NY, RNG), 26 Jun. 1984; Rockingham Bay, Wild (BRI 331587), no date; Yuccabine Ck, Kirrama Rd, Cardwell Ra., 27km WNW of Cardwell, 550m, poor rain forest, trailing from shrub branches, scattered but never common, HS 28696 ( + Barbella nitens - B, CBG, H, HSC, L, NICH, NY, PRE), 20 Jun. 1986; Ridge to Mt Dryander, 15km N of Proserpine, .± 600m, closed tropical forest, on rocks & trees, Henderson H2198 (BRI), 21 Jui. 1974; Eungella (Dalrymple Heights), 9.5km W of Finch Hatton, 720m, on planted palm in town, HS & Curnow 37703 (CBG, NY), 1 Jui. 1986; Credition S.F., 19km SW of Finch Hatton, 720m, roadside with Lantana in rain forest, on dead treelet branches, HS 37702 (B, CBG, COLO, DUKE, HSC, NAM, NY, RNG), 1 Jui. 1896; Cania, Bartlett (as Stone 21039 - MELU), 6 Aug. 1983; Mary Cairncross Park, 4km S of Montville, rain forest, epiphyte on shrubs, Henderson & Sharpe H3042 (BRI), 3 Nov. 1983; Balfour Ra., Brisbane Valley Highway, 29km SE of Nanango, 400m, dry semi-tropical rain forest with hoop pine (Araucaria cunninghamii), on tree branches, HS 9336 (ALTA, B, BISH, CBG, CHR, COLO, FH, H, L, MO, NICH, NY, UBC), 29 Aug. 1979; Mt Perry, Kalliwa Ck, Goodnight Scrub S.F., 80m, Araucaria vine forest, common, hanging from branches, Forster 802 (BRI), 2 Jui. 1981; Eumundi, rain forest, Fleischer B2696 ( + P. flexicaulis - ALTA, CANB, LO, NY), 25 Mar. 1903; Maryborough, ground moss, Clemens (L), Oct. 1948; Bunya Mountains N.P., on Bunya Mountain Rd, ea 14.4km SW of intersection with Bunya Highway, 760m, dense forest with Araucaria, hanging from a liana, Churchill & Kantak 12737 (CBG, NY), 8 May 1984; Brisbane, Bailey 815 (BM), no date; Pimpamae Ck, Wild (MEL 32146), Aug. 1887; Mt Mistake, Bailey 211 (BRI), no date; Sylvester Lookout, Cunninghams Gap N.P., tree branch, Stone & Curtis 20843 (MELU), 26 Feb. 1983; Mt Roberts, Lamington N.P., 885m, on branches of Argyrodendron, Blake 22919 (BRI), 8 Jun. 1968. NEW SOUTH WALES: Brindle Ck, Wiangaree S.F., 27km NNE of Kyogle, 750m, Cunoniaceae-Lauraceae dominated forest, trailing The genus Papil/aria 113

from bush branches, HS 6060 ( + P. crocea - CBG, H, L), 3 Oct. 1978; Whian Whian S.F., rain forest, pendant from branches, Robbins 2610 (CANB), 24 Nov. 1959; Cox Rd, Toonumbar S.F., 29km NW of Kyogle, 450m, on rock outcrop at edge of dry hoop pine (Araucaria) forest, HS 6960 (CBG, H, L), 18 Oct. 1978; Teven, Richmond R., on rocks, Watts (BM), Jui. 1897; Alstonville Rd, Ballina, on tree beside canal, Watts 3443 (NSW), Nov. 1899; Tintenbar, Richmond R., Bauerlin (as Watts 2402 - H, NSW), Aug. 1895; Gibraltar Ra., NE of Glen Innes, rain forest, Crawford (CBG, L, MEL), Dec. 1%3; Woolgoolga Ck, on shrubs, Schofield et al. 80749 (UBC), Jui. 1983; Chaelundi Mountain, 37km N of Ebor, 1376m, cool temperate forest, common trailer on shrub branches at forest edge, HS 6731 (ALTA, BA, CBG, CIBIMA, H, L, MO, NFLD, NICH, UBC), 14 Oct. 1978; Briggsvale, Wild Cattle Ck S.F. 12km NNE of Darrigo, 700m, Cunoniaceae dominated forest, canopy 20m with young emergent Araucaria cunninghamii, on tree trunk, HS 6635 (CBG, L), 13 Oct. 1978; Darrigo N.P., 4km SSE of Darrigo, 640m, rain forest, on the ground fallen from tree crown, Curnow 651 (B, CBG, DUKE, FH, HO, HSC, MICH, PRE, RNG), 23 Feb. 1986; Wilson's Ck Rd, Mt Boss S.F. 37km NNW of Wauchope, 940m, cool temperate rain forest, in crown of large tree, HS 7107 (CBG), 20 Oct. 1978; off Doyle's R. Rd, from Oxley Highway, ea 48 miles [77km] W of Wauchope, rain forest, on Dryopteris a/bovillosa, Garden (NSW M6), 4 Oct. 1951; Gloucester R., below Gloucester Tops, WSW of Gloucester, 1080m, pendulous on trees and logs in dense rain forest, Constable 5995 (NSW), 17 Jui. 1%5; Lister Park, Upper Allyn S.F. (road to Barrington Tops), on branches, Verdon 108B (CBG, L, NICH), 14 Aug. 1%9; The Bulga, Wingham, Watts 10752 (NSW), 18 Apr. 1915; Mt Coricudgy, E of Kandos, 360m, Eucalyptus-Doryphora rain forest, pendant from trees in dense garlands, van Steenis 22273 (C, CBG, H, L), 30 Oct. 1973; Mount Wilson, 23km NNE of Katoomba, 980m, on exotic street trees (elms), HS 31540 ( + P. crocea & P. flexicaulis - CBG, HO, NICH, NY), 1 Nov. 1984; near Homebush on Parramatta R., on trees, Whitelegge 275 (NSW), Jui. 1885; Jenolan Caves, Blakely 879 (H, NSW), Jun. 1899; Royal N.P., on boulder, Schofield 51422 (UBC), 17 Aug. 1972; near Moss Vale, on trees, Whitelegge (NSW Ml3015), 8 Nov. 1884; Vandenbergh Rd, off Jamberoo Rd, 4km E of Robertson, 580m, remnant temperate rain forest, on semi-exposed road cutting amongst grasses, HS 10491 (ALTA, B, BISH, BRI, CBG, COLO, DUKE, FH, H, HO, L, MICH, MO, NICH, NY, PRE, RNG, UBC), 17 Sep. 1980; Cambewarra Mountain, south side, rain forest, on trees, McVean 26457 (CBG), Jan. 1964; Vines Ck, Budawang Ra., 19km NW of Ulladulla, 660m, temperate Cunoniaceae dominated forest, on rotting log, HS 5463 ( + P. flavolimbata - AD, BA, CBG, CIBIMA, H, L, MO, NFLD, NICH, NY, UBC), 26 Apr. 1973; Mt Dromedary, 13km SW of Narooma, 870m, disturbed Eucryphia and Eucalyptus forest on moderate slope, trailing from lower tree trunk, shrub stem and branches, HS 27933 (ALTA, B, BA, BISH, CBG, DUKE, FH, H, HO, L, MICH, NAM, NFLD, NICH, NY, RNG). LORD HOWE ISLAND: Rocky Point to Boat Harbour, coastal rain forest, Ramsay s.m. (UNSW), 10 Nov. 1981. TASMANIA: Van Diemens Land, Lawrence (BM), no date. The genus Papillaria 114

2 PAPILLARIA C~OCEA (Hpe) Ja~g., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1875-76.267.1877 (Ad. 2.171).

Pilotrichum croceum Hpe, Linnaea 25:715.1853. Type: 'Van Diemens Land (Stuart)' HOLOTYPE: BM! Neckera crocea (Hpe) C. Muell., Bot. Zeitung (Berlin) 15:779.1857. Pi?otrichum nigrescens Homsch., Syn. 2: 135.1850, nom. nud. m syn. Neckera nigrescens Hornsch. ex Hpe, Fragm. Phytogr. Austral. Suppl. 11:50.1881, nom. inval. in syn. e". pro Pilotrichum nigrescens Hornsch. Meteorium cuspidiferum Wils., FI. Nov. Zel. 2: 101.1854. Type: 'Sinclair, J.D.H., Bay of Islands' HOLOTYPE: BM! Papillaria cuspidifera (Wils.) Jaeg., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1875-76:272.1877 (Ad. 2: 176). Neckera kermadecensis C. Muell., Bot. Zeitung (Berlin) 15:779.1857. Type: 'Cuming July 1854, Kermadec Is.' Papillaria kermadecensis (C. Muell.) Jaeg., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1875-76:265.1877 (Ad. 2:169). Pilotrichella kermadecensis (C. Muell.) Hpe, Fragm. Phytogr. Austral. Suppl. 11:50.1881 ('-doc-'). Meteorium kermadecense (C. Muell.) Mitt., Trans. & Proc. Roy. Soc. Victoria 19:82.1882. Trachypus homschuchii Mitt., J. Linn. Soc., Bot. 4:90.1860 (Meteorium cuspidiferum Wils., 1854 ), Trachycarpus homschuchii Mitt. ex Burg., Proc. Linn. Soc. New South Wales 60:86.1935, e". pro Trachypus homschuchii Mitt. Neckera reginae Hpe, Linnaea 40:319.1876, syn. nov. Type: 'Hartmann, Queensland' HOLOTYPE: BM!; ISOTYPE: LD! Papillaria reginae (Hpe) Jaeg., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1875-76:272.1877 (Ad. 2:176), syn. nov. Pilotrichella reginae (Hpe) Hpe, Fragm. Phytogr. Austral. Suppl. 11:50.1881, syn. nov. The genus Papi/laria 115

Papillaria acrocea (Hpe) Jaeg. ex Sainsb., Victorian Naturalist 49:259.1932, err. pro P. crocea (Hpe) Jaeg.

Slender or rarely robust plants, yellow-brown to dark green; primary stems up to 20cm long, often trailing, rarely branched. Secondary stems up to 10cm, distantly branched, branches up to 2cm, attenuate at apex. Leaves dense, .± imbricated, erect and appressed when dry, erecto-patent in 5 whorls at an angle of 45°(80°) when wet. Herbarium collections retain yellow-brown colour.

Branch leaves from broadly cordate, ovate-triangular to broadly lanceolate (0.8)1.0-1.5(2.0)x0.4-0.8(1.0)mm, tip acute or rarely acuminate, base broadly acuminate or rarely slightly winged. Margin slightly undulate, entire or with slight denticulations on the auricles. Ratio of leaf width to length ea 0.50. Costa prominent on dry leaves, nearly reaching to tip becoming slightly twisted. Leaf plicate on each side of costa, this being quite pronounced as a ridge in the upper part of the leaf, width at base (20)30-40(50)um, cells rectangular 50.0x2.5um in central part.

Cells in leaf tip longer and narrower, incrassate, with no papillae, sometimes sigmoid. Cells in the upper lamina rounded-rhomboidal, oval or rhomboid (2.5)7.5-12.5(15.0)x2.0-5.0(7.0)um, median cells of similar size and shape to those above, very rarely longer. Cells thin walled, densely papillose with papillae often obscuring cell outline. Marginal cells (1 or 2 rows) slightly smaller, quadrate or more rounded, barely or not papillose, .± forming a border. Juxtacostal cells above base up to 30-35(60)um long, smooth, prosenchymatous, getting shorter above and gradually merging into papillose median cells. Long juxtacostal cells are arranged in divergent rows and in auricle margin reach 10.0x2.5-5.0um. These cells are parallel to margin, with few or no papillae. Towards base, cells are slightly smaller and parenchymatous, quadrate cells occasionally observed on basal corners of auricles. n1e genus Papil/aria 116

Fig. 28. Papillaria crocea (Hpe) Jaeg. 1-9. Branch leaves: 1,4. HS 5504; 2. Goldsmid 876; 3,7,9. HS 28682; 5,6,8. HS 6524. 10. Perichaetial leaf: HS 38467.

Stem leaves slightly longer and narrower, but often leaf border not well developed.

Smaller (0.9mm) rounded to ovate leaves with few papillae occur at base of branches. Costa and denticulations in the base are barely visible. Cells in upper leaf are 10.0x3.0um and similar to branch leaves, juxtacostal cells near base up to 50um long, border lacking. Large central portions of leaves appear clear with only the edges darker. 171e genus Papillaria 117

--20-

Fig. 29. Papi/laria crocea (Hpe) Jaeg. Branch leaf cells: 1,2. Apex; 3. Upper margin; 4. Median margin; 5. Median near mid-leaf; 6. Auricle. (1-4. HS 6528; 5,6. Boorman M1544).

Stems rounded to oval in cross-section 230-400x160-310um. Epidermal cells quite similar to outer cortical cells, hexagonal to roughly rounded, irregular in shape 10.0-12.5x2.5-3.0um, incrassate. Inner cortical cells are quite distinct with size increasing towards centre where they are narrow hexagonal to rounded 28.0x12.5(21.5)um. A very small central strand was noted in some sections. Inner cortical cells up to 300x160um. Branch cross-section rounded to oval 180- 300x160-220um. Epidermal cells similar to outer cortical cells, irregular in shape 10.0-12.5x2.5-3.0um, cell walls equally wide, but slightly thicker than those of the stem. Inner cortex up to 180x100um thick and clear. In this, and the stem, outer cortex is about 5 cells wide. Boundary between golden-brownish outer cortex and clear inner cortex is quite distinct in both stem and branch sections. The genus Papillaria 118

1 2 3

4 5 6

7

Fig. 30. Papillaria crocea (Hpe) Jaeg. 1. Branch (x53); 2. Branch and sporophyte (xll); 3. Capsule and peristome (x53); 4. Peristome teeth (x207); 5. Papillose leaf surface (x533); 6. Capsule with peristome (x64); 7. Operculum (x29). (1-3,5,7. Watts 9468; 4,6. Tindale & Constable M12981). The genus Papillaria 119

Rhizoids not common, short, 12.Sum wide, cells prosenchymatous, 30- 40um long.

Inner perigonial leaves ovate-obovate, 600x400um, tip acuminate, costa weak, 3/4 leaf length, weakly multi-papillose, margin entire, bordered by narrow rectangular cells. Upper cells hexagonal 16-20(35)x7um, incrassate, median cells hexagonal up to 37x7um, nearer base up to 50um long. Antheridia .± 6 per bud, up to 380x200um. Paraphyses as long (5-6 cells) as the antheridia, up to 17um wide, cells up to 63um long. Perichaetia rounded or slightly elongated. Inner perichaetial leaves ovate, up to 850x350um, costa faint, 3 / 4 leaf length. Margin distantly irregular, marginal cells rectangular and hexagonal. Tip and upper cells hexagonal or.± sigmoid, ea 25x5um, incrassate, median juxtacostal cells similar, 37x2um, shorter towards the margin and with 1- 3 papillae per cell, towards base irregularly rectangular ea 37x5um, along base short rectangular cells. Outer perichaetial leaves ovate, ecostate, not papillose. Archegonia .± 6 per bud, up to 330x70um, neck 50um wide, with irregular short rectangular cells. Paraphyses few, as long as the antheridia or slightly longer, up to 14um wide, cells rectangular up to 40x7um. At base of capsules inner perichaetial leaves lanceolate with parallel sides 0.9-1. 7x0.6mm, cos ta extending nearly to leaf tip, faint, margins entire. Upper and median cells ea 12.5x2.5um, incrassate, papillae up to 3 per cell. Cells slightly sigmoid in apex. Lower cells longer, (up to 80um) but similar to cells of branch leaves. Basal margin cells rectangular, long and narrow (55x2um) but nearer tip often with a prominent margin of hexagonal cells. Alar cells rarely present. Smaller rounded leaves at base of perichaetial leaves slightly papillose. Paraphyses numerous, up to 25um (2-3 cells) wide, mid-leaf cells up to lO0um long. Die genus Papillaria 120

Fig. 31. Papillaria crocea (Hpe) Jaeg., in cool temperate rain forest, Mt Dromedary, southern New South Wales (HS 38220 - Photo: R. Hotchkiss). 17ie genus Papillaria 121

Calyptra 2.3-2.Smm long. Capsules ovoid l.3-l.6x(0.4 )0.6-0.9mm, setae (2.8)4.0-4.5(5.0)mm long. Peristome teeth 650um long and two cells wide, endostome segments linear, 500um long and narrow, one cell wide with narrow long papillae. Operculum (0.9mm) with long curved beak. Exothecial cells mostly irregularly rounded to quadrate (from 25um), some 25.0x7.5um with irregular outline. Inner capsule wall cells similar to exothecial but more obscure. Spore sac with similar cell pattern. Spores 23-30um.

Chromosome number: n = 11 (Ramsay 1967, 1974; erroneously reported as P. f/exicaulis.)

DISTRIBUTION and ECOLOGY: Eastern Australia - Queensland, New South Wales, Lord Howe Island, Victoria and Tasmania (Fig. 32); extending also to Norfolk Island (Fig. 33) and New Zealand. Other reports are for India, Sri Lanka, Malaya, Philippines, Vietnam, Java, Papua New Guinea, Kermadec Is., New Caledonia, Fiji, Samoa, Raratonga and Tahiti.

Commonly inhabits moist areas - rain forest, wet sclerophyll forest, scrubby vegetation or in more exposed areas beside streams. Most commonly found growing on lower tree trunks but if the forest is very dense, then it is found higher on the trunks and in the tree crowns. P. crocea has also been collected on rocks and this occurrence could be more common than the collections indicate. This moss also occurs on vines, rotting logs and tree roots. In suitable areas it may occur on any of the above substrata and is often mixed with other Papillaria species, Barbella nitens, Macromitrium spp., liverworts and lichens. Papillaria crocea has been found epiphytic on Acacia, Acmena, Araucaria, Argyrodendron, Bursaria, Cardwe/lia, Chrysophyllum, Cinnamomum, Citrus, Elaeodendron, Endiandra, Eucalyptus, Ficus, Hymenanthera, Lantana, Thegenus Papillaria 122

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Fig. 32. Distribution of Papillaria crocea (Hpe) Jaeg. The genus Papillaria 123

Melaleuca, Notelaea, Polyscias, Pomaderris, Salix, Telopea and Tristania. Near Batemans Bay (New South Wales) it was also found growing on Casuarina cunninghamiana Miq. on the edge of an Avicennia and Aegiceras dominated swamp.

It is distributed from sea level to 1200m, the higher altitudes being observed in the more northerly locations of its range in Australia. In Victoria the highest altitude is 270m, and no data are available for Tasmania. The species is far more common in New South Wales and Queensland.

On Norfolk Island (Fig. 33), at the higher elevations, (130-300m) P. crocea is a very common moss, especially on the ridges where there is ample moisture due to low clouds and mist. The preferred vegetation type is the native forest which is now generally heavily infested with Psidium guajava L.

(guava) and Lantana camara L. It occurs on all parts of trees, on dead Cyathea

(tree ferns), on vines (Freycinetia) and on rock outcrops. P. crocea was also recorded on the following - Araucaria, Citrus limon (L.) Burm.f. ( domestic lemon), Cordyline (or Dracaena), Elaeodendron, Lagunaria and Syzygium.

Associated with the extensive masses of Papillaria crocea were the following: lichens - Coccocarpia, Leptogium and Parmelia (s.1.); liverworts - Frullania,

Metzgeria and numerous other unidentified genera; mosses - Macromitrium and

Orthorrhynchium.

ETYMOLOGY: from the Latin, croceus = saffron yellow, alluding to the plant's colour.

ILLUSTRATIONS: Noguchi 1976:243; Noguchi & Bartlett 1985:266; Sainsbury

1955a:355. The genus Papi//aria 124

29° 04'

i]Nepean Is

167° 55'E 167° 59'

Fig. 33. Distribution of Papillaria crocea (Hpe) Jaeg. on Norfolk Island. The genus Papillaria 125

TYPE MATERIAL

The type of Papillaria regi,nae is very close to P. cuspidiferum, but leaves of the type of P. crocea are larger. The plicate leaf and the twisting of the costa are present in all three types and all other features are similar. The branch cross-section of P. regi,nae is similar to P. crocea, but the yellow colour extends to the inner cortex where the cells are also less regular.

NOTES

Long acuminate curled leaf tips may be found in the species. The most prominent (Watts 3649) having rounded or quadrate cells in single rows at the apex. Lower down where the apex widens the cells are narrow, very incrassate and in several rows.

The leaves of P. crocea can be long and narrow (Topic - H, JE) with length of the costa not related to the leaf length. Plants with these leaf types may have a superficial resemblance to P. flexicaulis, but this latter species has leaves which are always much more smoothly curved abaxially and cell arrangement in the auricles differs. The collection by Topic: also exhibited very prominent long, narrow and incrassate laminal cells which were not noted in other collections. Two collections by Hartmann (JE) from southern Queensland have the normal appressed leaves but they are slightly arched to give the moss a slightly 'puffed out' appearance. The twisted costa is quite conspicuous on the back of the leaf under 16x magnification. Sometimes stem leaves can have a more solid, darker red costa.

Often some perigonial leaves have very thin sigmoid cells in the tip and upper part of the leaf. Perigonia have a similar appearance to branch buds. The genus Papillaria 126

Mature collections seem to be common during October and to a lesser extent December and May.

In Norfolk Island collections the costa appears straighter. There are long, narrow leaves on the shoots and stem, branch leaves generally being 0.9- 1.7x0.5-0.7mm. Cells in upper 5/7 leaf are papillose while near the base a clear straight line division between papillose and non-papillose sections is present. This is not the case in Australian populations. Some long, fine branches have the general appearance of P. jlexicaulis.

In Asian species the upper part of the costa is rarely twisted as prominently as it is in Australian species. One Australian collection (Stone 15785) had a straight costa and no plication of the leaf.

Differences between P. crocea and P. jlexicaulis in Asia have not been satisfactorily elucidated because of lack of collections (Noguchi 1976). The same was true for the Australian species. One obvious difference is in the way the leaves are held on the branches. In P. crocea the leaves have a prominently raised costa and the leaves are not tightly wrapped around the branch. However, in P. jlexicaulis the costa is not prominent nor raised, the leaves being smoothly and tightly wrapped around the stem. In the auricles of P. crocea the divergent cells tend to turn parallel to the margin but in P. jlexicaulis the cells intersect the margin at right angles and the auricular cells are more uniform (Fig. 10 p. 54 ). Also in P. crocea the basal margin can be denticulate while in P. jlexicaulis it is only finely crenulate.

Only a limited number of Asian species were studied but it seems that confusion between these two could have arisen because some P. crocea collections were as slender as P. jlexicaulis, the costa was not ridged and the leaves were held more tightly to the branches. However, the more 17ie genus Papillaria 127

characteristic 'crocea' type leaves can be found if specimens are carefully examined.

Fleischer (1908) notes the size of spores as 12-15um and irregularly rounded for the Java species, that is smaller than in Australian specimens.

Norris & Koponen (1985) doubted the occurrence of P. crocea, based on a Brass collection in Papua New Guinea. Because Noguchi (1976) had found the differences between it and P. jlexicaulis not ~!ways clear, Norris & Koponen suggested that the Brass collection could be the latter species.

The Brass (22664) collection was obtained on loan from Farlow

Herbarium (FH), and Hilbrands (1968) had redetermined the collection as P. leuconeura with which I agree. This specimen therefore represents the first New Guinea record for P. leuconeura. However, amongst Papua New Guinea collections of mine and another (determined as P. flexicaulis) from Helsinki, P. crocea was found. Both collections showed characters typical of Australian P. crocea.

SPECIMENS STUDIED: Australia 647 (66 cited), Norfolk Island 42 (3 cited), New Zealand 45, Sri Lanka 3, India 3, Vietnam 1 ( cited), Indonesia 2, Philippines 1, Papua New Guinea 3 (cited), Vanuatu 3, Kermadec Is. 1, Raratonga 2, Fiji 3, Tahiti 7.

EXSICCATI: C. Mueller, Kryptogamae Exs. 99 ( + P. flexicau/is).

QUEENSLAND: Windsor Tableland, 38km NW of Mossman, 1140m, rain forest with Agathis robusta, trailing from treelet branches, HS 29718 (CBG, NY), 26 Jui. 1984; Mossman R. Gorge, 4 miles [6km] W of Mossman, lowland rain forest, on branch, Weber B-31812 (ALTA, COLO), 19 Apr. 1968; Mt Lewis, 1040m, on Citrus branches, clearing in forest, Brass 33938 (FH), 3 Nov. 1968; Davies Ck, SE of Mareeba, virgin upland rain forest, on blown down tree, Weber B-32408 (COLO), 21 Apr. 1968; Yungaburra Rd, 2km SE of Atherton, 850m, exotic fruit orchard, on Ficus dammaropsis stem, HS 16851 (CBG, L, NICH), 27 Jan. 1983; Maalan, 11km E of Ravenshoe, The genus Papillaria 128

980m, remnant scrub between road and paddock, on a shrub, HS 30606 (CBG, NICH, NY), 30 Jun. 1984; Babinda, Watts Q327 (NSW), Jui. 1913; Yuccabine Ck, Kirrama Rd, Cardwell Ra., 27km WNW of Cardwell, 550m, rain forest, on a vine, HS 28682 (CBG), 20 Jun. 1984; Mt Spee S.F., 6km W of Paluma, 920m, Lauraceae-Syzygium dominated forest, on treelet stem, HS 36988 (CBG), 18 Jun. 1986; Credition S.F., 20km SSW of Finch Hatton, 840m, Argyrodendron-Syzygium dominated forest, on upper tree trunk, HS 37635 (B, CBG, DUKE, NAM, NY, RNG), 1 Jui. 1986; Rockhampton, Dallachy (MEL 31235), 1863; Keppel Bay, Mueller (BM), no date; Cania Gorge N.P., Stone 22702 (MELU), 6 Aug. 1984; Hurdle Gully, 14km WSW of Monto, 300m, dry semi-tropical scrub forest on creek flats, on shrub branch, common, HS 9896 (CBG, L), 16 Jan. 1980; Mapleton Falls, earth bank, Schofield 51612 (UBC), 23 Aug. 1972; Bunya Mountains N.P., on Bunya Mountain Rd, 760m, dense forest with Araucaria, on tree trunk, Churchill & Kantak 12734 (CBG, NY), 8 May 1984; Balfour Ra., 29km SE of Nanango, 400m, dry semi-tropical rain forest with hoop pine (Araucaria), on tree trunk, HS 9339 (CBG, L, NICH), 29 Aug. 1979; Caboolture, Shirley 29.5, no date; Enogerra, Broadbent (MEL 32251), no date; Brisbane, Bailey (C), no date; Mt Glorious, Maiala S.F., 635m, sub-tropical rain forest, on tree bark, Sawyer & Ross 165 (BRI), 13 May 1985; Toowoomba, ad arboreus, Hartmann (BM, LD), Dec. 1876; Tamborine Mountain, Longman & White (BRI 331567), Feb. 1916; Mt Roberts, McPherson Ra., rain forest, bark of Eucalyptus saligna, Blake 23000 (BRI), 19 Aug. 1968; Mt Cordeaux, Cunninghams N.P., 1000m, open rain forest, on shaded tree trunk, Norris 36118 (CBG, H, NY), 14 Mar. 1974. NEW SOUTH WALES: Murwillumbah, on trees on river flats, Goldsmid 876 (H, NSW), Aug. 1899; Cox Rd, 29km NW of Kyogle, 450m, rock outcrop with numerous shrubs on edge of dry hoop pine (Araucaria) forest, on fallen hoop branch, HS 6957 (CBG, L), 18 Oct. 1978; Tintenbar, Richmond R., on trees, Watts (BM), Dec. 1897; Grafton Common, Watts 146 (NSW), 14 Feb. 1896; Hyland S.F., Hyland Rd, head of Perrotts Ck, 30km NW of Darrigo, 700m, temperate rain forest, on Meliaceae trunk, HS 6691 (CBG, FH, H, L, MO, NICH, UBC), 14 Oct. 1978; Mt Lindsay, Forsyth (FH), Sep. 1900; Bruxner Park Flora Reserve, 6km NW of Coffs Harbour, 200m, lowland rain forest, on Polyscias, HS 6524 (CBG, L), 12 Oct. 1978; Forbes R., Mt Boss S.F., 51km NW of Wauchope, 1060m, Eucalyptus forest, on Eucalyptus trunk, HS 7166 (CBG), 20 Oct. 1978; Gannons Ck, 11km SW of Wauchope, 150m, edge of cool temperate and Eucalyptus forest, on a rotting log, HS 5947 (CBG, L), 28 Aug. 1978; Manning R., Cross (MEL 32218), Dec. 1882; O'Sullivans Gap, 9km N of Bulahdelah, tropical rain forest, Ramsay 2619 (UNSW), Aug. 1986; Gloucester R., 28km WSW of Gloucester, 400m, Eucalyptus and Casuarina forest, on tree trunk, HS 6455 (CBG, L), 11 Oct. 1978; The Bulga via Wingham, Watts 10799 (NSW), May 1915; Kingwell, Wyong, Watts 9468 ( + Barbella nitens - NSW), Nov. 1907; Cessnock­ Wollombi Rd, on shaded perpendicular outcrop, Schofield et al. 80993 (UBC), 15 Jui. 1983; Mount Wilson, 23km NNE of Katoomba, 980m, on tree (elm) trunk, HS 31547 (CBG, H, HO, NICH, NY), 1 Nov. 1984; Megalong Valley, on tree trunk in forest, Schofield et al. 79605 (UBC), The genus Papillaria 129

19 Jun. 1983; Stanwell Park, Watts 8231 (NSW), May 1905; Parramatta, Mueller (BM), no date; Royal N.P., branch and bark of tree, Ramsay 2504 (UNSW), 16 Jui. 1972; Macquarie Pass, 25km SW of Wollongong, 100m, rain forest, on branches of small tree, HS 4770 (CBG, L, NICH), 12 Aug. 1977; Vandenbergh Rd, 4km E of Robertson, 480m, remnant temperate rain forest with Acacia, Elaeocarpus & Cunoniaceae, on tree trunk, HS 10494 (CBG, H, L), 17 Sep. 1980; Butmaro Ck, 24km NW of Braidwood, 860m, grasslands, on exotic tree trunk, HS 4655 (AD, CBG, H, L, MO, NFLD, NICH, NY), 29 May 1977; Buckenbowra R., 7.5km WNW of Batemans Bay, 2m, Casuarina cunningltamiana dominated swampy area, on Casuarina stem, HS & Curnow 35822 (B, CBG, NY), 2 Nov. 1985; Long Beach, 5km E of Batemans Bay, 10m, vegetation on cliff, on boulder, HS 27709 (ALTA, B, BA, BISH, CBG, COLO, DUKE, FH, GRO, H, L, MICH, MO, NAM, NFLD, NICH, NY, RNG), 28 May 1983; Head of Oulla Ck, 18km SW of Araluen, 700m, cool temperate forest, on branches of big tree, HS 7339 (AD, ALTA, B, BA, CBG, CHR, COLO, DUKE, FH, H, HSC, HO, L, MICH, MO, NAM, NFLD, NICH, PRE, RNG, UBC), 16 Nov. 1978; Mt Dromedary, 13km SW of Narooma, 870m, disturbed Euc,yphia forest with Eucalyptus, on tree trunk, HS 27944 (ALTA, CBG, H, HO, L, NICH, NY), 27 Oct. 1983. LORD HOWE ISLAND: Whitelegge s.n. (MEL), Aug./Sep. 1887; Mt Lidgbird, Vitt 28365 (ALTA), 16 Nov. 1981. VICTORIA: upper Ovens R., McCann (MEL 32197), no date; Martin Ck, Bonang Highway, 34km NNE of Orbost, 270m, regrowth beside stream in wet sclerophyll forest, on Telopea stem, HS 35476A (B, CBG, NY), 26 Oct. 1985; Masons Falls, Kinglake N.P., on damp Acacia melanoxylon trunk at foot of falls, Willis (MEL 1046599), 14 Oct. 1945; Mt Drummer, on a tree in the forest, Melville 2832C (MEL), 12 Jan. 1953; Croajingolong N.P., The Rapids, W side of Wingan R., Beauglehole 31945 (MEL), 21 Nov. 1969; Iguana Ck, Bairnsdale District, Vincent (MEL 1023998), 2 Jan. 1972; Deadcock Ck Gorge, 17km NW of Bairnsdale, on jungle tree trunks, Willis (MEL 1334366), 26 Jun. 1946; Sherbrooke Forest, Dandenong Ra., on dead tree branches, Hand (MEL 32199), 14 Feb. 1953; Loch, on stones in creek, Murdock (MEL 32203), Dec 1913. TASMANIA: Lawrence (BM), Jun. 1857; Glengary, McLeod (MEL 32183), no date; Trevallyn Ck, South Esk R., Launceston, on a tree, Weymouth 2055 (H, HO), 20 Jan. 1888. NORFOLK ISLAND: Mt Pitt 300m, poor forest, on shaded rock outcrop, HS 34837 (CBG), 10 Dec. 1984; between Palm Glen and Red Rd, 180m, upper parts of tree, Chinnock 5974 (AD, CBG), 15 Jan. 1984; Filmy Fern Trail off Selwyn Pine Rd, Mt Pitt Reserve, 130m, dense forest beside creek, on dead Cyathea, HS 32077 (CBG, H, NICH, NY), 3 Dec. 1984. PAPUA NEW GUINEA. MO ROBE PROVINCE: E of Wantoat R., 8km NNW of Wantoat, 1500- 1700m, partially disturbed montane forest with occasional areas of cultivation, Norris 60445 (H), 29 May 1981; Manki, ea 3 miles [5km] SW of Bulolo, in an almost pure stand of Castanopsis which has regenerated on area formerly covered by kunai (lmperata cylindrica) grass, on branch of fallen Castanopsis tree, Shea 6494 (CBG) 16 Nov. 1974; Upper Nawata Banda, 10km S of Bulolo, 1500m, Castanopsis dominated ridge, HS 33969 (CBG, H, LAE, NY), 10 Feb. 1983. The genus Papillaria 130

VIETNAM: Hanoi, on tree trunk, Balansa (H), 1888. The genus Papillaria 131

3 PAPILLARIA FLAVOLIMBATA (C. Muell. & Hpe) Jaeg., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1875-76:267.1877 (Ad. 2:171).

Neckera jlavolimbata C. Muell. & Hpe, Linnaea 26:502.1853. Type: 'Australia felici sine loco indic' LECTOTYPE: BM!, ISOLECTOTYPES: BM!, MEL(32174)! Pilotrichella jlavolimbata ( C. Muell. & Hpe) Par., Ind. Bryol. 944.1897. Meteorium cerinum Hook.f. & Wils., FI. Tasm. 2:203.175.f.4.1859. Type: 'On trees. (Barren specimens). (Gunn, 1605). On trees, in creek, Archer'. LECTOTYPE: Archer - BM!, ISOLECTOTYPE: NY! SYNTYPE: Gunn (1605) BM! Trachypus cerinus (Hook.f. & Wils.) Mitt., J. Linn. Soc., Bot. 4:91.1860. Neckera cerina (Hook.f. & Wils.) Tayl., J. Linn. Soc., Bot. 4:91.1860. Pilotrichel/a cerina (Hook.f. & Wils.) Hpe, Fragm. Phytogr. Austral. Suppl. 11:50.1881, nom. illeg. incl. spec. prior. Papil/aria cerina (Hook.f. & Wils.) Par., Ind. Bryol. 902.1896. Neckera luteola Tayl., J. Linn. Soc., Bot. 4:91.1860, nom nud. zn syn. Meteorium cuspidiferum Wils. var. cerinum (Hook.f. & Wils.) Hook.f. & Wils., Handb. New Zealand FI. 459.1867. Papil/aria cuspidifera (Wils.) Jaeg. var. cerina (Hook.f. & Wils.) Jaeg., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1875-76:272.1877 (Ad. 2: 176). Meteorium limbatum Mitt., Trans. & Proc. Roy. Soc. Victoria 19:82.1882, nom. illeg. incl. spec. prior.

Plants robust, yellowish, light to dark green; primary stems branched or unbranched, flexuose up to 30cm long. Secondary stems creeping or trailing to 10cm long and 1.5-2.0cm thick with irregular pinnate branching which is generally at right angles. Branches attenuate at apex. Leaves on very slender trailing shoots terminate in a slender forked point, that can be multiply branched, The genus Papillaria 132

or club shaped, spikey and papillate. Some of these features are also developed on some longer, narrower leaves. In the field moist plants are dark green in colour but a lighter green towards apices, while dried plants are yellow-green as are stems on herbarium collections. Stem and branch leaves imbricated, turgid, erect to erecto-patent, particularly when moist.

Branch leaves broadly lanceolate-triangular up to (1.1)1.9-2.2(3.0)x0.6- 1.6mm with an acute apex and auricled clasping base. Young leaves can also be very narrow. Margin undulate, entire with slight denticulations on the auricles rarely reaching apex, margin strongly bent back abaxially in mid-leaf. Ratio of leaf width to length ea 0.49. Costa extending to (1/2)3/4 length of lamina (rarely to the tip), 30-50um wide at base, cells up to 50um long, poorly differentiated on both sides of lamina.

Upper lamina! cells (7) 18-20( 42)x5-7um, rhomboidal to narrowly oval, slightly longer and narrower towards tip and thicker walled, with marginal cells to 25um long. Median cells (7) 12-20x5-7um, same shape as above with marginal cells to 37um long, more rectangular and incrassate. Cells nearer costa parallel to it, cells further out tend to align obliquely to costa, rarely vertically. In lower leaf, cells gradually merge into basal cells. Papillae dense, often obscuring cells, irregularly shaped up to 5.0x2.5um, fewer towards base where papillae smaller and more regular. Rarely are cells near the margin papillose, thus forming a conspicuous border. Basal cells smooth or only slightly papillose as in lower sections of auricles. Basal juxtacostal cells (25)50-65x5-7um. A group of quadrate, apparently alar cells, sometimes with a red-yellow tinge, often occur in basal corners of auricles. Cells tend to align parallel to margin, away from costa in auricles, becoming thinner, elliptic with rounded ends (22-50um along the margin). This band of smooth cells was quite broad in auricles forming a border 1-4 rows wide in upper lamina. 11te genus Papi/laria 133

Fig. 34. Papil/aria flavolimbata (C. Branch leaves: 1-3. Muell. & Hpe) Jaeg. HS 9535; 4,5. HS 10495. The genus Papillaria 134

A whorl of rounded ecostate leaves (1.0mm long) with slightly cucullate tips occur at the base of each branch. No papillae observed in cells at base, or along margin, but in upper part of lamina up to 3 papillae per cell. Upper cells 5.0-7.0xl.0-2.0um and slightly wider (2.Sum) at tip. Cells 25x1(2)um and uniform except at base. At basal corners a group of quadrate (lOum) or slightly rectangular alar like cells with an orange tinge are present. All the cells were aligned vertically.

Stem leaves 2.3-2.5xl.l-1.3mm, auricles wider and better developed than on branch leaves.

Stems ovate to rounded in cross-section up to 330x240um thick. Epidermal cells 4 or 5 cells wide were similar to cells of outer cortex which were small and rounded (3-Sum), incrassate near margin but with cell size increasing towards centre. Inner cortical cells slightly irregular 25.0x12.5um, more angular, with central strand. Inner cortex can be up to 200x150um. Branch stems were also rounded, up to 300x250um thick. The epidermal cells were rounded 2.Sum, incrassate while inner cortical cells were up to 27.0xl 7.Sum, of variable shapes ranging from oval to rounded. Central strand present. Inner cortex was up to 200x150um thick. Boundary between the golden-brownish outer cortex and the clear inner cortical cells was distinct. The stem and branch sections were not noticeably denticulate.

Rhizoids not common, scattered along stems in very sparse clusters, 10( 12.S)um wide, cells prosenchymatous and generally up to 200um long. The genus Papillaria 135

---... 20.um

Fig. 35. Papillaria f/avo/imbata (C. Muell. & Hpe) Jaeg. Branch leaf cells: 1. Leaf apex; 2. Upper cells in mid-leaf; 3. Median cells at mid-leaf; (1-3. Forsyth M12991); 4. Cells at median margin (HS 35570); 5. Cells in auricle (Forsyth M12991); 6-11. Apices of young branch leaves (HS 36702). The genus Papillaria 136

Fig. 36. Papi//aria flavo/imbata (C. Muell. & Hpe) Jaeg. 1. Branch and sporophyte (x4); 2. Capsule (x20); 3. Median marginal cells (x400); 4. Auricular cells (x750); 5. Branch (xll); 6. Operculum (x31); 7. Peristome teeth (x207); 8. Papillose leaf surface (x533); 9. Spore (x960). (1,2,6. HS 15369; 3. Adams 2251; 4. Forsyth M12991; 5,8,9. Curnow 1287). The genus Papillaria 137

Inner perigonial leaves oblong up to l. lx0.8mm, tip acuminate, margin distantly irregular, ecostate or weak, 3/4 of leaf length, rarely papillose, but then only in upper leaf with 3 or 4 papillae per cell, rarely weakly developed. Margin distantly irregular, narrow rectangular marginal cells best developed in lower half. Upper cells rhomboidal 37x7um, median cells rhomboidal­ hexagonal, 50x7um, basal juxtacostal cells similar but up to 58um long, _± rectangular. Antheridia _± 20 per bud, up to 570x200um. Paraphyses up to 500um long and 17um wide, cells rectangular up to 67(112)um long. Perichaetia rounded, not conspicuous, often partly covered by leaf. Inner perichaetial leaves ovate 1100x450um, with_± parallel sides, serrate, costa faint, 3/4 of leaf length, papillae prominent in upper half. No rectangular marginal cells, rarely so in lower half. Upper cells linear-elongated, 30x2-4um, incrassate, median cells hexagonal, 37x5um, nearer base elongated but irregularly shaped, base with band of rectangular cells, 25x10um. Archegonia _± 12 per bud, up to 460x60um, neck 40um wide. Paraphyses as long and as numerous as the archegonia, up to 15um wide, cells_± 40( 60)um long. Thinner paraphyses often have the longest cells. At base of capsules inner perichaetial leaves plicate, narrow to narrowly triangular, variable l.1-2.7x0.3-0.7mm with very few papillae. Cells variable throughout up to 115x2um, juxtacostal cells at base 63x5um with a row at leaf insertion shorter and becoming yellow. All cells were aligned vertically. Costa very faint extending to 3/4 leaf length. Smaller rounded leaves at base ecostate, or costa barely visible, apex rounded, serrulate. Paraphyses numerous and colourless, up to 20um (2 rarely 3 cells) wide, cells incrassate, up to lO0um long, with granular content.

Calyptra 3.5mm long with cells up to 70um long. Capsules ovoid (1.6)2.0-2.Sx0.8-l.0mm, setae 7.5-12.6mm long. No operculum seen. Peristome The genus Papillaria 138

Fig. 37. Papillaria flavolimbata (C. Muell. & Hpe) Jaeg., on shrub branches at Monga State Forest, New South Wales (Curnow 1287 & HS 37807 - Photos J. Curnow). The genus Papillaria 139

teeth 600um, endostome segments 400um. Exostome cells 25um, rounded, slightly larger nearer base. Sporophytes emerge at irregular intervals on branches and may contain immature, mature, overmature or broken setae. Spores (15)25(35)um.

Chromosome number: n = 11 (Ramsay 1967, 1974).

DISTRIBUTION and ECOLOGY: Eastern Australia - Queensland, New South Wales, Victoria and Tasmania (Fig. 38). Occurring outside Australia only in New Zealand.

This species is generally found in rain forest, Nothofagus dominated forest and wet sclerophyll forest that fringes denser, moister forest. Common on tree trunks and in tree crowns, trailing from the branches of trees or shrubs. Often in very dense, pure masses. Plants continue to grow on dead fallen tree branches and this could have led to reports of it occurring on logs. Several collections were from rocks, but this is not common for this species. However, near Greenwich Park (32km NE of Goulburn, New South Wales) Papillaria flavolimbata was found scattered on Megalong Conglomerates in dry sclerophyll forest which suffers from regular burning. Here it occurs on the steep sides of the deep shaded rocky gullies where direct sunlight is rare; it grows in pure, small to medium sized colonies, often surrounded by numerous other mosses. It also grew entwined amongst roots of Dendrobium striolatum Reichb.f. and Py"osia rupestris (R.Br.) Ching. This species has been collected on a large range of phorophytes including: Acacia, Atherosperma, Bedfordia, Bursaria, Ceratopetalum, Cyathea, Dicksonia, Elaeocarpus, Eucryphia, Leptospermum, Notelaea, Notlzofagus, Pomaderris, Plzyllocladus, Prostantlzera and Telopea. 1 ne genus rapmana 140

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... Cooktown I <7 0 \ I • \,, Ca1tns L I ; I o ,,/ I r ,'-Z. I I G'I I \ I \ NT I \ I \, 20' I ' Mackay I \ I \ I QUEENSLAND \ I \ I \ ' 25 I \ I \ ------, \ • .'S I '-----� I �- I z � � BRISBANE I

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0 100 200 JOO 4«11,m Simple projectionconiC

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Fig. 38. Distribution of Papil/ariaflavolimbata (C. Muell. & Hpe) Jaeg. The genus Papillaria 141

Papillaria jlavolimbata shows an altitudinal range of 60- lS00m being most common at high altitudes, the highest altitude observed in Tasmania and Victoria being 800m but in New South Wales up to 1500m. The Queensland collections are without detail, but were approximately 1000m above sea level. In fact there are only ten collections (mainly recent collections by I.G. Stone) known from Queensland mostly from the ranges near the New South Wales border and extending north to the Bunya Mountains. The earliest Queensland collection was referred to as P. semitorta (MEL 32268 ex Herb. Bailey).

E1YMOLOGY: from the Latin, jlavus = yellow, limbatus = bordered, alluding to the lighter colour of the lamina margin.

ILLUSTRATIONS: Allison & Child 1971:109; Sainsbury 1955a:355; Wilson 1859:fig.175 (Meteorium cerinum).

TYPE MATERIAL

The BM collections were chosen as lecto and isolectotypes respectively since they were from Hampe's herbarium and annotated 'Neckera jlavo-limbata' in his handwriting. The annotation 'Sealers Cove' had been added later, but the writing is similar even though it is smaller.

A collection from MEL (32176) is annotated as type but this came from Sonder's collection (with his handwriting). Consequently it cannot be designated as such with certainty. However, a second MEL (32174) collection can be assumed to be an isolectotype as it is annotated in Hampe's handwriting, even though he renamed it as a Meteorium.

For the name M. cerinum the Archer collection (BM) was chosen as the lectotype. A duplicate exists in NY and it is of equal quality to that of Gunn ' The genus Papi/laria 142

1605. Both specimens were cited in the original diagnosis by J.D. Hooker & W. Wilson.

Paris (1897:904) gives Pi/otrichella jlavolimbata Jaeg. as a synonym of

Papillaria jlavolimbata. However, in the stated reference (Jaeger Ad. II, p. 171) no such entry was found therefore this Paris synonym can be regarded as an error.

NOTES

In some forms dense pendulous mats have trailing filiform stems and branches. The filiform branches are very similar to those of Barbella nitens because the leaf margins are strongly bent abaxially in both species. However,

B. nitens is a golden colour and appears delicate, while Papillaria jlavolimbata is a very light green and has a firmer texture. These two species often grow intermixed.

When the auricles are not well developed then the leaves can quite easily be mistaken for those of P. crocea. However, just above the base there is often a 'pinch' that is noticeable in fixed slides. This is due to irregularities caused by the undulate auricles. Another distinguishing feature is the very wide margin of long smooth cells in the auricles.

Papillaria jlavolimbata tends to show alar cell development more often than the other Papi/laria species (Campbell BRI 3316616, Bate MEL 32182, Weymouth 2404). The margin of clear cells is rarely absent in the upper leaf (HS 10492). Very seldom is the costa forked in mid-leaf (Watts 10333).

Some perichaetial leaves had a forked tip with prominent denticulations on the apex (Curnow 1287). The genus Papillaria 143

No reliable pattern for fertile material could be detected from the limited data. However it appears that mature capsules may be expected during February-April and also June-October. From my own observations at Monga (New South Wales) during October 1986 numerous colonies were noted which had immature, mature and over mature capsules together with perichaetial leaves and paraphyses where the capsule had dropped out.

Sainsbury mentioned that New Zealand collections were quite often a colourful orange-yellow, but this was not noted in the Australian collections.

A Victorian collection identified as P. flavolimbata (Merrah - MEL 32177) is possibly another distinct species. Dr Noguchi wrote that he also found a distinct species from New Zealand (McMahon 356 - WELT), which on examination appeared similar to the Australian collection in leaf aerolation and leaf shape, but differed in leaf sizes though not significantly. A trip to Mt Ellery (eastern Victoria) failed to locate any material. Before this species can be described further attempts should be made to obtain additional collections from Mt Ellery and all the New Zealand collections of P. flavolimbata should be examined.

This distinct species differs from P. flavolimbata by its longer linear lamina! cells and the very incrassate cell walls. Because both species are densely papillose the cell outline is often obscured. Only when the leaf is cleared do the differences become apparent.

No P. flavolimbata was found among Norfolk Island collections.

SPECIMENS STUDIED: Australia 353 (43 cited), New Zealand 52.

QUEENSLAND: Mt Kingarow, Bunya N.P., on trunk of grass trees, Schofield 90647 (UBC), 16 Sep. 1987; Mt Mistake N.P., Stone 20890 (MELU), 27 Feb. 1983; Lamington N.P., Stone & Curtis 20582 (MELU), 16 Feb. 1983; Granite N.P., Wallangarra, Clemens (FH), Nov. 1944. NEW The genus Papillaria 144

SOUTH WALES: Wiangaree S.F., on trees, Schofield et al. 80674 (UBC), 10 Jui. 1983; Gibraltar Ra., 15 mile [24km] E of Glen Innes, coachwood rain forest, Perich B-5320 (COLO), Dec. 1960; Chaelundi Mountain Rd, Marengo S.F., 30km NNE of Ebor, 1300m, cool temperate rain forest, on tree trunk, HS 6766 (CBG, H, L, MO, NICH), 14 Oct. 1978; Mt Kaputar, Nandewar Ra., 44km NNE of Boggabri, 1500m, exposed rocky summit, on boulders, HS 9291 (CBG, H, L, MO, NICH, NY, UBC), 26 Aug. 1979; Forbes R., Mt Boss S.F., 51km NW of Wauchope, 1060m, Eucalyptus dominated grasslands, on Eucalyptus trunk, HS 7165 (CBG), 20 Oct. 1978; Ellenborough Falls, Watts 10864 (NSW), Apr. 1915; Gloucester Tops, 34km WSW of Gloucester, 1260m, Nothofagus forest, trailing from treelet, HS 6421 (BA, CBG, CIBIMA, H, L, MO, NFLD, NICH, UBC), 10 Oct. 1978; Mt Alum, Bulahdelah, Boorman (NSW Ml2989), 16 Aug. 1911; Barrington Tops, ea 40 miles [64km] N of Singleton, epiphyte on high branches of Nothofagus, Pullen 3795A (CANB, L, NSW), 4 Feb. 1963; Mount Wilson-Mount Irvine Rd, 25km NNE of Katoomba, 720m, cool temperate forest (Cunoniaceae dominated), on a vine, HS 31595 (B, CBG, H, MICH, NICH, NY, RNG), 1 Nov. 1984; Lawson, in gully, Watts 5986 (NSW), 4 Apr. 1903; Parramatta, Mueller (MEL 32181), no date; Kangaroo R., 22km N of Nowra, 150m, Eucalyptus forest, on Leptospennum, HS 4968 (CBG, H, L, MO, NICH), 26 Aug. 1977; Wollondilly R. via Longreach, cliffs beyond river, Watts 9035 (NSW), 23 Feb. 1909; Moreton N.P., Yurnga Lookout, 35km NE Braidwood, 740m, temperate rain forest, on rock face, Curnow 72 (B, CBG, NICH, NY), 28 Oct. 1984; Higgins Ck, 19km NE of Batemans Bay, 60m, remnant cool temperate forest, upper branches of large tree, HS 5754 (CBG, H, L, NICH), 9 Aug. 1978; Great Dividing Ra., 5km SE of Captains Flat, 1220m, Eucalyptus forest on ridge dominated by shrubs, on shrub branches, HS 31346 (B, CBG), 8 Aug. 1984; Mt Dromedary, 13km SW of Narooma, 870m, disturbed Eucryphia-Eucalyptus forest, on Eucryphia trunk, HS 27943 (CBG, H, HO, L, NICH, NY), 27 Sep. 1983; Brown Mountain, 50km SE of Cooma, wet sclerophyll forest, on Dicksonia antarctica, Doing M81 (L), 20 Nov. 1963. VICTORIA: upper Ovens R., McCann (MEL 32175), no date; Eurobin Falls, Mt Buffalo, Stone 1316 (MELU), 11 Apr. 1969; Bonang R., 95km NE of Orbost, 720m, wet sclerophyll forest, tree trunk, HS 10047 (ALTA, B, BISH, CBG, COLO, FH, H, HO, L, MO, NFLD, NICH, NY, PRE, UBC), 4 Apr. 1980; Mt Ellery, E. Gippsland, Merrah (MEL 32173), 1887; Croajingolong N.P., Howe Ra., near Harrisons Ck, Beauglehole 31415 (MELU), 1 Nov. 1969; Myrtle Ck, Healesville­ Donna Buang Rd, 10km SE of Healesville, 800m, wet sclerophyll forest beside stream, fallen from Acacia, HS 36294 (B, CBG, HO, HSC, MICH), 3 Jan. 1986; Bemm R., Princes Highway, 8km SSW of Club Terrace, 100m, remnant cool temperate forest with Acmena and tree ferns, on dead Cyathea, HS 35529 (CBG), 27 Sep. 1986; Boggy Ck, Nowa Nowa, Clark 4 (MEL), 24 Aug. 1952; Powelltown off Pioneer Ck Rd at Astelia Ck, 400m, wet sclerophyll forest, van Zanten 82.03.1040 (GRO), 3 Mar. 1982; Kallista, Stone 536 (MELU), Jui. 1966; Bulga N.P., epiphytic in forest, Lewinsky 1806 (C), 13 Dec. 1979; Arkins Ck, Beauglehole 74273 (MEL), 3 Oct. 1966; Melba Gully State Park, near Lavers Hill, Otway Ranges, on fallen tree trunk near stream, Beauglehole & The genus Papillaria 145

Willis 5217 (MEL), 10 Mar. 1962. TASMANIA: Sumac R., logged rain forest, Brasell H332 (CBG, HO), 10 Nov. 1979; Ulverstone, Gerard (as Bastow 810 - HO, MEL), Jui. 1882; Blue Tiers, 7km ENE of Weldborough, 710m, Jarman 1783 (CBG, HO), 16 Dec. 1982; Ruby Flat Forest near Ringarooma, 300m, on tree limb, Weymouth 2726 (HO), 11 Nov. 1913; Murchison Highway, 4km S of Tullah, 310m, Jarman 1240 (CBG, HO), 18 Jan. 1983; Henty R., west coast, on celery top pine, Weymouth 633 (CBG, HO), 7 Feb. 1891; Gordon R., on trees, Weymouth 2759 (HO), 1 Jan. 1914; Newmans Ck, 'Fernleigh', Tasman Peninsula, on wood, Weymouth 692 (HO), 19 Mar. 1891. The genus Papi//aria 146

4 PAPILLARIA FLEXICAULIS (Wils.) Jaeg., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1875-76:271.1877 (Ad. 2:175).

Meteorium flexicaule Wils., FI. Nov. Zel. 2:101.1854. Type: 'Colenso, Hawke's Bay, Dr Stanger, Northern Island, New Zealand'. LECTOTYPE: Colenso - BM!; ISOLECTOTYPE: BM! SYNTYPE: Dr Stanger, BM! ( = P. amblyacis). Trachypus flexicaulis (Wils.) Mitt., J. Linn. Soc., Bot. 4:91.1860. Leskeaflexicaulis Tayl., Handb. New Zealand FI. 460.1867, nom. nud. in syn. Meteorium filipendulum Hook.f. & Wils., Fl. Tasm. 2:203.1859. Type: '(Gunn 26, 1606), (Lawrence)'. LECTOTYPE: Lawrence - BM (H 2113)! ISOLECTOTYPE: Lawrence - BM (H 2212)! Papillariafilipendula (Hook.f. & Wils.) Jaeg., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1875-76:271.1877 (Ad. 2:175). Pilotrichella filipendula (Hook.f. & Wils.) Hpe, Fragm. Phytogr. Austral. Suppl. 11:50.1881. Neckera scottiae C. Muell., Linnaea 35:621.1868. Type: 'Miss Helen Scott, Ash Island in Hunter River, eastern Australia; Brisbane River lectum mis. Cl. G. Reichenbach'. LECTOTYPE: Helen Forde - NY! Pilotrichella scottiae (C. Muell.) Hpe, Fragm. Phytogr. Austral. Suppl. 11:50.1881. Meteorium filum C. Muell. ex Bailey, Syn. Queensland FI. Suppl. 3:98.1890, nom. nud. Papillaria scottiae (C. Muell.) Jaeg., Ind. Bryol. 908.1897. Cryphaea novae-valesiae C. Muell., Hedwigia 55:284.1914, nom. nud. Pilotrichum pendulinum Tayl. in Dix., Notes Roy. Bot. Gard. Edinburgh 20:95.1950, nom. nud. in syn. Papillaria filicaule Tayl. ex Burg., Proc. Linn. Soc. New South Wales 60:85.1935, nom. nud. e". pro P. flexicaulis. The genus Papillaria 147

Cryphaea papillarioides Broth., Proc. Linn. Soc. New South Wales 24:375.1899, nom. nud.

Plants slender, very dull, dark green or dark green. Primary stems distantly pinnate. Secondary stems up to 30cm long, flexuose. Branches short (1cm), flexuose, pinnate or sub-pinnate, generally nearly at right angles, often attenuate at tips, especially on slender trailing forms. Ends of secondary stems or branches at times clavate, where leaves at first tend to spread out slightly. Stems and branches terete, leaves imbricated and strongly appressed or erect and erecto-patent when wet. Secondary stems up to 0.5mm thick, including leaves, while the branches are up to 0.8mm thick. Stems and leaves on herbarium collections pale green to brownish-green with age.

Branch leaves 0.8-1.7(2.0)x0.4-l.2mm, triangular ovate concave, not plicate, tip acute, base widely cordate. Younger branch leaves with a long tip. Margin appearing slightly crenulate due to papillae and slightly projecting marginal cells. Width to length ratio of leaves 0.54. Costa extends to 3/ 4 lamina length, sometimes faint and hard to see, width at base 20-30( 60)um, costa cells (up to 45um long) all similar, but often hard to differentiate.

Leaf tip when acute with apical cells generally similar to those of upper leaf. When tip acuminate, cells tend to be more variable, often longer and narrower ( ea 25x4um), incrassate, with irregular sides. Cells in upper leaf (7.5)10.0-15.0x2.5-5.0um, rhomboidal with rounded corners or rarely oval, being generally of uniform shape and size, papillose. Juxtacostal cells are slightly bigger and parallel to it. Median cells similar to upper cells, more uniform in shape and size, at times appearing to be oblique to costa, papillae dense extending to margin. Leaves often with 1-4 rows of cells at margin not so densely papillose, but rarely without papillae, never with an appearance of a The genus Papillaria 148

border. Juxtacostal cells near the base smooth, long and narrow, 26-50x5um; away from costa intersecting margin at right angles and similar to rest of laminal cells. Papillae increase gradually towards margin, but are not dense. Cell size gradually decreases upward from base while density of papillae increases. On lowest part of lamina distant from the costa, cell shape and size variable, and cells smooth.

Stem leaves are slightly narrower and longer, but otherwise similar.

Hidden at base of branches are smaller,..± rounded, 380-500x300-400um, leaves, scarcely papillose, ecostate with denticulations similar to those of branch leaves. Cells in upper part of leaf 15-20um long in middle of base to 40um long. Area with basal juxtacostal cells small, cells variously elongated up to 50x5um. Cells at basal margin are more rectangular and quadrate meeting margin at right angles.

Stems more rounded in cross-section in this species, up to 230x210um. Epidermal and outer cortical cells small, 2um, incrassate near the margin but cell size increasing towards centre. Outer cortex 5 cells wide. Inner cortical cells more irregular towards the centre, still rounded-ovate, but with some quadrate cells. Cells up to 20.0x12.5um. Inner cortex up to 150x120um wide with a small central strand. Branches similar but more rounded, up to 170x160um with inner cortex 120um in diameter. Epidermal and outer cortical cells rounded, but larger than those in stem, inner cortical cells rounded, 12um in diameter with larger and smaller cells mixed. Boundary between outer cortex and inner cortex distinct. A small central strand sometimes present. Stem and branch margins weakly denticulate.

Rhizoids occur as thick tufts between leaves, up to 17um wide, prosenchymatous cells up to lO0um long, often hard to see. The genus Papillaria 149

1 2 3

5

7

50 JJm

Fig. 39. Papillaria flexicaulis (Wils.) Jaeg. 1-4. Branch leaves (Archer - HO 7453, Cruttwell, ibid, Watts 2129); 5. Perigonial leaf (Watts 4041); 6. Leaf apex; 7. Upper cells near mid-leaf; 8. Median cells at mid-leaf; 9. Median marginal cells; 10. Cells in auricle; (6-10. Archer - BM). The genus Papil/aria 150

6

Fig. 40. Papil/aria flexicaulis (Wils.) Jaeg. 1,2. Branch with perigonia (x22, x27); 3. Median cells near costa (x600); 4. Median leaf margin (x600); 5. Leaf base (x600); 6. Antheridium (x850); (1,2. HS 38465; 3,5. Watts M13006; 4. Archer­ BM; 6. Watts 4041).

Inner perigonial leaves ovate to obovate up to 900x350um, ecostate, tip acuminate. Margin entire, border lacking or not very prominent. Upper cells and apex incrassate, rhomboidal rarely sigmoid, .± 25x2um, median cells hexagonal or rhomboidal 25-40x2-5um, basal juxtacostal cells.± rectangular up The genus Papillaria 151

to 63x10um. Upper and median cells weakly papillose with up to 3 papillae per cell. Antheridia .± 5 per bud, up to 400x120um. Paraphyses few, as long as the antheridia, up to 20um wide, cells up to 44um long. Perichaetia bud like. Inner perichaetial leaves entire, ovate up to 910x420um, ecostate or costa faint to 2/3 leaf length, weakly papillose in upper half, margin entire, border not prominent. Upper and median cells elongated 25x5um, juxtacostal cells nearer base rectangular 37x10um, all cells vetically aligned. Archegonia .± 6 per bud, up to 330x60um, neck 40um thick. Paraphyses as long as the archegonia and about twice the number, up to 17um wide, cells up to 37um long. At base of capsules inner perichaetial leaves variable 1.5-2.6x0.2-0.4mm narrowly triangular, tip acuminate, base not cordate. Margin with 2 or 3 rows of rectangular cells, rarely quadrate, with few papillae, up to 3 per cell. Costa faint, extending 3/4 leaf length and sometimes nearly to apex. Apical cells 33x5um, incrassate, median cells 42x5um and juxtacostal cells at the base quite variable 65-75x5um. All cells are aligned vertically. Paraphyses numerous, colourless, generally 30um (2 cells) wide. Cells at tip 12.Sum long while in mid-section up to lO0um long, parenchymatous.

Calyptra hairy, 1.8mm long, cucullate, smooth on the inside. Capsules 1.20x0.85mm, ovoid; setae smooth up to 8.7mm long. Operculum not seen. Peristome teeth lanceolate, 350um with small papillae. Exothecial cells obscure. Spores 7-20um rounded or slightly irregular.

Chromosome number: no reports. Previous reports (Ramsay 1967, 1974) are referrable to P. crocea.

DISTRIBUTION and ECOLOGY: Eastern Australia - Queensland, New South Wales, Victoria and Tasmania (Fig. 41). This species occurs widely outside Australia, in Tanzania (Ochyra & P6cs 1985), Sri Lanka, India, Indonesia, The genus Papi//aria 152

Philippines, Formosa, Papua New Guinea, Norfolk Island, New Zealand, Juan Fernandez Island and southern Chile. Habitats include rain forest, dry sclerophyll forest and rarely open Eucalyptus woodland. This moss is generally found trailing in intricate masses from shrub or treelet branches, on tree trunks, or more rarely on tree roots, vines or road cuttings. On one occasion it was found mixed with other Papillaria species on a fallen branch from tree crown in rain forest. It has also been collected on rocks with the most common report being from limestone. Occasional associates on rocks include Orthotrichum cupulatum Brid, Encalypta vulgaris Hedw., Hypnum cupressiforme Hedw., Thuidium and Racopilum cuspidigerum (Schwaegr.) Aongstr. Papillaria flexicaulis has been found to be epiphytic on Acmena, Brachychiton, Bursaria, Ceratopetalum, Chrysophyllum, Eucalyptus, Ficus, Polyscias, Pomaderris, Sterculia and Strychnos.

Reported altitudinal range is been between 100-800m. This species could have occured at lower altitudes as Whitelegge collected this at Georges River, Botany Bay (New South Wales) in April 1885. Collections from Pimpamae and Burpengary in Queensland are also from near sea level.

ETYMOLOGY: from the Latin, flexi = to bend and caulis = stalk, referring to the pliant stems.

ILLUSTRATIONS: Noguchi 1976:241; Norris & Koponen 1985:29; Wilson 1859 fig. 175 (Meteoriumfilipendulum); Wu & Lin 1985a:16. Thegenus Papillaria 153

.,,1f

Cuokto111111

l'J - I 0 I I I I xr I ' I ' NT \ ' I \ I 20" I I I I QUEENSLAND I • • 2s• I I I •• • ;• ------, • •• 2 I I I

I <( ''----____ ,,,,,-- ...... ' _ 0 _ 3 I ____ SOUTH I AUSTRALIA I I 30' I

I ' - � NEW SOUTH WALES I"\ ---·• I ,,....' -. . I � ...... \ 'I:""/ I, __ '-- Q ,' ADELAIDE f I ·-, I '\." I ,_, -____ '\ 35• f VICTORIA --� � \ I ----- c, �----�,'

40"S

JO'S

0 100 200 3()() 400"m S,mple con,c proj«tion ,so• ,s:.·

Fig. 41. Distribution of Papillariaflexicaulis (Wils.) Jaeg. The genus Papillaria 154

TYPE MATERIAL

On examination of the syntypes of Meteorium flexicaule (BM) it was found that the two collections did not represent the same taxa. Dr Stanger's collection (H 2109) is in fact Papillaria amblyacis and clearly shows the characters of that species. Further the illustration on the sheet clearly shows the folding of the leaf that develops due to the auricles and the cucullate leaf so characteristic of that species.

The Colenso collection (1431b) is a slender form with female 'parts' and annotated with a varietal name. The illustration on the sheet is clearly that of the present species P. flexicaulis. Consequently the Colenso (1431b) collection has been chosen as the lectotype for P. flexicaulis.

For Meteorium filipendulum the Lawrence collection in BM (H 2113 in pencil, 48 in ink) was chosen as the lectotype because it is a rich well-formed collection. This and both isolectotypes (in pencil: H 2212 & 138) have the stamp 'Herbarium Hookerianum'. Both have the same annotations in a similar hand. The Gunn 26 & 1606 collections remain syntypes. All exhibit the typical characteristics of P. flexicaulis.

A collection of Neckera scottiae, collected by Helen Scott under her married name (Forde) was found amongst the NY loan. This was designated the lectotype of this taxon. The other collection from the Brisbane River could not be located. Dr F.K. Meyer (JE) suggested it could be at Hamburg University but Walther & Martienssen (1976) do not list it as being in that institution. The genus Papillaria 155

NOTES

This species is quite easily recognized and it does not exhibit the range of variations that were observed in Papillaria crocea or in P. flavolimbata. One variation found, which was not common, was in the leaf tip which tended to be longer and finer. In fact one collection (Watts 4188) could be described as having long hair points. Another collection (Watts 205) had leaves of nearly equal length and width. The distribution of papillae was quite regular but one collection with long thin leaves (Schofield 51131) had numerous papillae in the leaf base. When wet the leaves spread 60-80° from the stems and appeared to be weakly arranged in 5 rows.

Hilbrands (1969) considered this species to be close to P. crocea and thus proposed that it be considered as a variety of it. The differences were that P. jlexicaulis had shortly triangular leaves with a straight weak costa reaching .± 1/2 leaf length while P. crocea had elongate triangular leaves, the costa was bent, strong and reaching nearly to the tip. However, the leaf shape in P. crocea is quite variable and therefore not a reliable character. No mention was made of the plicate leaf nor the leaf cells which are different in the two species. Therefore these two taxa should remain as two species.

No collections with mature capsules were noted.

For comments on confusion between P. flexicaulis and P. crocea refer to notes under the latter species (p. 126).

SPECIMENS STUDIED: Australia 256 (39 cited), Norfolk I. 1 (cited), New Zealand 28, Taiwan 1, Philippines 2, Indonesia 7, Papua New Guinea 10 (3 cited). The genus Papillaria 156

QUEENSLAND: Millstream Falls, Ravenshoe, basalt cliff, W.B. & M.I. Schofield 80081 (UBC), 30 Jui. 1983; 40 Mile Scrub (basalt), Stone 17921 (MELU), 21 Oct. 1981; Undara Crater, microphyll vine forest on inside of crater, epiphyte on Strychnos lucida on Pleiogynium timorense, Goodwin (as Stone 23051 - MELU), 29 Nov. 1984; Ashton Ck, Wallaman Falls Rd, 28km W of Ingham, 300m, poor rain forest beside rocky stream, on boulder, HS 28509 (CBG), 19 Jun. 1984; Tabletop, Hervey Ra., 39km WSW of Townsville, 350m, dry sclerophyll forest, on boulder, HS 37193 (B, CBG, DUKE, NY, PRE), 20 Jun. 1986; Eungella (Dalrymple Heights), 720m, on planted palm in town, HS & Curnow 37704 (B, CBG, NY), 1 Jui. 1986; Port Curtis, Bailey 396 (BRI), no date; between Springsure and Emerald, on trees in vine thicket, McVean 26353 (CBG), Nov. 1963; 81km N of Injune, bottle tree scrub, Stone 20486 (MELU), 28 Jui. 1982; Hurdle Gully, 14km WSW of Monto, 300m, semi-tropical scrub on creek flats, common on shrub branches, HS 9895 (CBG, H, L, MO, NICH, NY), 16 Jan. 1980; Lockaber Ck near Eidsvold, Sherrin 45 (BM), Apr. 1922; Balfour Ra., 29km SE Nanango, 400m, dry semi-tropical rain forest, on tree trunk, HS 9333 (CBG, H, L, MO, NICH, UBC), 29 Aug. 1979; Wescott Track, Bunya Mountains, Stone 17446 ( + P. amblyacis - MELU), 23 Oct. 1980; Caboolture, Wild (BRI 331554), Sep. 1888; Newmarket, Brisbane, Wild (BRI 331595), Jui. 1888; Knoll N.P., Tambourine Mountain, Stone 20686 (MELU), 20 Feb. 1983; Cunninghams Gap, Bailey (BRI 331581), no date; Burleigh Heads N.P., Stone 22950 (MELU), 13 Aug. 1984; Binna Burra, Caves Circuit, Stone 12762 (MELU), 11 Jun. 1975; Brown Falls, 4km SE of Killarney, remnant forest, on a rock, HS 230A (CBG), 26 Sep. 1973. NEW SOUTH WALES: Sharps Scrub, Richmond R., on tree, Watts 377 (H, NSW), 15 May 1896; Drake, on stone, Porter 1969 (NSW), Jun. 1898; Grafton Common, Watts 145 (H), 14 Feb. 1896; Hillgrove near Armidale, in gorge, Watts (NSW M12998), Nov. 1903; Bruxner Flora Reserve, 6km NW of Coffs Harbour, 200m, lowland rain forest, on trunk of Chrysophyllum, HS 6534 ( + P. crocea - CBG, H, L, NICH), 12 Oct. 1978; Briggsvale, Wild Cattle Ck S.F., 12km NNE of Dorrigo, 700m, Cunoniaceae andAraucaria dominated forest, HS 6630 (CBG, L), 13 Oct. 1978; Cambden Haven S.F., 28km SW of Port Macquarie, 480m, rain forest on very rocky southerly slope, on treelet stem, HS 38465 (B, NY), 29 Aug. 1987; Barrington, N of Dungog, Stone 11772 (MELU), 29 Aug. 1976; Mount Wilson-Mount Irvine Rd, 25km NNE of Katoomba, 750m, cool temperate forest Monimiaceae dominated, on a vine, HS 31567 (CBG, H), 1 Nov. 1984; Jenolan Caves, scrub forest, Vitt 27727 (ALTA), 18 Oct. 1981; Hurstville, on Sterculia diversifolia, Forsyth s.n. (MEL), 18 Jui. 1901; Georges R., Botany Bay, Whitelegge 264 (H), Apr. 1885; Bungonia Gorge, 28km ESE of Goulburn, 120m, Acmena smithii & Casuarina dominated creek bank, on Acmena branch, HS 6185 (CBG, H, L), 14 Sep. 1978; Misty Mountain Forest Rd, Currowan S.F., 23km NW of Batemans Bay, sassafras and tree fern dominated forest, on tree (Eucryphia) trunk, HS 37874 (CBG, NY), 2 Dec. 1986; Wadbilliga R., 44km ESE of Cooma, 350m, on rocks amongst shrubbery beside river, HS 3760 (CBG, H, L, MO, NICH), 13 Jui. 1976. TASMANIA: Scottsdale­ St Helens Highway, 1.5 mile [2km], E of Parr Rivulet, 300m, Eucalyptus-Atherospenna forest, on The genus Papillaria 157

Pomaderris, Norris 31895 (H, NY), 11 Nov. 1974; Westbury, on trees, Roper 2053 (H, HO, NSW), 8 Sep. 1888; Cheshunt, Archer (HO 74753), no date. NORFOLK ISLAND: Wools s.n. (amongstP. crocea - BM), no date. PAPUA NEW GUINEA. SIMBU PROVINCE: Dirima Village, 3km NW of Gumine, 6°11'S, l44°55'E, 1680m, on tree trunk beside river in grasslands, Toia 140 (ALTA, B, BA, BISH, CBG, COLO, DUKE, FH, H, LAE, NICH, RNG), 17 Dec.1982. MOROBE PROVINCE: Middle Ck­ Watut R. Divide, 8km NW of Bulolo, 7°09'S, 146°53'E, 950m, disturbed lower montane forest on northerly facing slope, on fallen tree branch, HS 14294 (CBG, H, LAE), 2 Feb. 1981; Forestry College, Bulolo, 7°10'S, 146°40'E, 730m, maintained garden in flat exposed area, on cultivated Pandanus, HS 14142 (CBG), 31 Jan. 1981. The genus Papillaria 158

5 PAPILLARIA INTRICATA (Mitt.) C. Muell. & Broth., Abh. Naturwiss. Vereine Bremen 16:505.1900.

Meteorium intricatum Mitt., J. Linn. Soc., Bot. 10. l 71.f.5a.1868. Type: 1 Tutuila, on living trees in the shady forests' (20-lO00ft) No. 5. HOLOTYPE: NY!; ISOTYPES: NY! MEL!

Slender plants, dark green, darkening to deep dark reddish-brown with age, secondary stems to 20cm long, laxly pinnate, densely leaved, not terete, apex acute or attenuate. Leaves erecto-patent when dry, when wet only slightly more so (up to 45°). Stems on herbarium collections same colour as dried leaves which are a golden-green or brownish golden-green.

1 2

Fig. 42. Papillaria intricata (Mitt.) C. Muell. & Broth. Dietrich-H: 1. Branch (x27); 2. Branch with emerging secondary branch (x12).

Branch leaves lanceolate to ovate l.2-1.4x0.3-0.5mm, apex acuminate or shortly attenuate and twisted, base cordate. Margin.± recurved, denticulate at base, distantly and less prominent further up the leaf. Ratio of leaf width to 17ie genus Papillaria 159

length not calculated. Costa, reaching to 1/2 of leaf length, very weak or indistinct, width at base 20um.

]:

. _:.r .. :. · .. ·.

20.....

Fig. 43. Papi//aria intricata (Mitt.) C. Muell. & Broth. Dietrich-H: 1-3. Branch leaves; 4. Leaf apex; 5. Upper cells near mid-leaf; 6. Median marginal cells; 7. Median cells near mid-leaf; 8. Cells in auricle.

Apical cells up to 25x2um, thin walled, not papillose, upper and median cells linear, up to 40x3um with 4-6 small, not prominent, papillae in a row over each lumen, thin walled, with flexuose, threadlike thickening on the inner surface The genus Papillaria 160

of the walls. Marginal cells similar to median cells but slightly shorter, thinner and less papillose. Cells in auricles irregularly rhomboid, size variable (5)13(17)x3-5um, not papillose, but upper part may be slightly so. Alar cells of variable size, irregularly quadrate, cells at base of auricle rectangular. Basal juxtacostal cells rectangular ( 15)30x( 4) 12um in one or two rows, not coloured.

Stem leaves 1.3-1.8x0.6mm, lanceolate tip acute, not so twisted, auricles more prominent, papillae reaching further down into the auricles.

Base of branches with no conspicuous whorl of smaller leaves, but very few leaves broadly ovate .± 0.6x0.4mm, tip rounded, ecostate, margin and papillae similar to branch leaves, with prominent group of alar cells ea 12x3um.

Branches ovate in cross-section up to 125x105um. Epidermal cells similar to outer cortical cells which are .± 2 layers wide, cells irregular 5x2um very incrassate. Inner cortical cells .± 6 layers wide, cells clear, irregular up to 17x10um, incrassate. No clear division between outer and inner cortical cells.

Rhizoids not common, up to 0.7mm long, but generally less and 18um wide, but narrowed at insertion, cells up to 125um long, ends mostly rounded and overlapping, rarely prosenchymatous.

Perigonial leaves lanceolate 0.6x0.2mm, tip acuminate, base rounded. Cells linear to linear-hexagonal in median section. Apical cells 30x2um, incrassate, median cells 62x7um, thin walled; irregular roughly hexagonal and obscure at base. Antheridia 8-11 up to 340um long. Perichaetia small elongated buds. Inner perichaetial leaves entire, lanceolate-ovate 1.0x0.3mm, tip long acuminate, ecostate, papillae none or very weak. Margin entire with inconspicuous narrow rectangular cells. Upper cells linear up to 37x2um, incrassate, median cells very weak, linear-rhomboidal up to 63x4um, band of The genus Papillaria 161

various sized rectangular cells along base. Archegonia 6-12 per bud, 320x30um, neck 20um, reddish with venter blackish-red, neck cells rectangular up to 25x12um. Paraphyses few (up to 4) as long as the archegonia, 12um wide, cells up to 50um long with thick walls. At base of capsules perichaetial leaves acuminate spreading.

Calyptra with a few hairs. Capsule oblong-cylindrical and asymmetrical. Setae moderately thick and nearly smooth with paraphyses. Peristome teeth elongate, basal membrane lacking. Operculum rostrate.

Chromosome number: no reports.

DISTRIBUTION and ECOLOGY: Northeastern Australia - Queensland (Fig. 44). This species has also been reported from Guam, Marianas, New Caledonia, Fiji, Samoa, Raratonga and Tonga.

The Australian collections which were not fertile, had no habit or habitat data, but it may be assumed the collections were found in lowland forest. Generally this species is found on tree and treelet trunks and branches.

ETYMOLOGY: from the Latin, intricatus = entangled, referring to the dense mats this plant forms.

ILLUSTRATIONS: Mitten 1868:fig. SA; Noguchi 1985:98; Noguchi & Bartlett 1985:266. 771egenus Papillaria 162

..--��-�---�·--·--,.----r------r------.----,

' ' Cooktown I ' ,�· I 0 � , I I I o ,, (r-Z- : 1 \ G" I \ I \ I \, NT I ',, I \

I \ I QUEENSLAND \ 5' I \ 2 I \

---I ---- \I I \ 2s· II '--·-----� I.!)', .. , I z ' ( I ! BRISBANE <( \ 'L__ � I __ ,,,,-- ...... ' - __ 0 _ _ 3 __ __ , I \ I ,- 30" I , ... ,/ SOUTH I AUSTRALIA I ' .,. I ': I _ ' r--, NEW SOUTH WALES ' ()---- I '-- () ,- ADELAIDE I 1 ·-, I ·'\." �--- ,J- ..... - '\• c A'NBERRA 1 , 35' I --� , I ---- c,}----�,-' VICTORIA � \ t '

40"S 0 i 4(/"S

o roo 200 JOO 400/

HOBART Simple conic prqection TASMANIA 140-E ,,s•V ,so• ,ss-

Fig. 44. Distribution of Papillariaintricata (Mitt.) C. Muell. & Broth. The genus Papillaria 163

NOTES

This species is a surprising, although not an unexpected, addition to the Australian moss flora. Only two collections were noted from a Helsinki loan, although one could be a duplicate. Perhaps because of the loss of lowland forests this species has not been collected again. However, as the central coastal areas of Queensland have rarely been collected, this moss could perhaps be found again after patient searching.

Noguchi (1985) mentions the development of flagellate branches that have small lanceolate leaves, but this was not noted in the Australian collections.

No adequate descriptions were found of the sporophytes, excepting the original brief description by Mitten (1868). However, it appears that perigonia are slightly less common.

Dixon & Greenwood (1930:284) claimed that Papillaria intricata and P. helictoplzylla (Mont.) Broth. have similar leaf characteristics and could only be separated when fertile. The latter species has immersed capsules (Brotherus 1925:814 Fig. 608) while P. intricata has exerted capsules. However, Hilbrands (1969) did find the following differences between the two species:

1) P. intricata cells.± 35um, P. he/ictophylla cells less than 30um long; 2) P. intricata has alar cells while P. he/ictophylla has none; 3) P. intricata has up to 7 or 8 papillae per cell while P. helictophylla has 4 or 5 papillae per cell; 4) P. intricata has sinuous cells, P. he/ictophylla does not. The genus Papillaria 164

Noguchi (1985) comments that this species has a close resemblance to Barbella nitens in outward appearance. In fact the earliest collections of B. nitens by Watts were determined as Papillaria intricata by Brotherus. However the two taxa are quite distinct with the twisted branch leaves of P. intricata separating it from Barbella nitens. This latter taxon also has characteristic recurving of the leaf and a distinctive golden colour. Noguchi (1985) also considers that Papillaria intricata is close to the Asian species P. semitorta. On outward appearance P. fuscescens resembles P. intricata as both species have a twisted leaf tip and many stem and older branch leaves are dark reddish-brown. Description of the perichaetia was from a Samoan collection (Fleischer BH 1249) where only two very small paraphyses were noted.

SPECIMENS STUDIED: Australia 2 (1 cited), Samoa 7, Fiji 1, Pacific Islands 1. QUEENSLAND: Rockhampton, Dietrich (H), 1866. The genus Papillaria 165

6 PAPILLARIA LEUCONEURA (C. Muell.) Jaeg., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1875-76:267.1877 (Ad. 2:171).

Neckera leuconeura C. Muell., Syn. 2:670.1851. Type: 'Java: Blume'.

Slender plants, pale green to green-brown, not glossy. Primary stems short, secondary stem up to 30cm long, flexuose densely leaved with pinnate or irregular branching. Branches widely spreading, irregular in length up to 1.5mm long and 0.7mm thick, densely leaved, terete, attenuate at apex. Leaves appressed, densely imbricated, only slightly erecto-patent when wet. Stems and branches a very light greenish colour, older stems darkening to a dark brownish­ black.

Branch leaves ovate-lanceolate often weakly panduriform (1.1)1.4-

1.5x(0.4 )0.5-0.7mm, tip acuminate, base cordate and denticulate, margin erect or slightly undulate above mid-leaf. Ratio of leaf width to length not calculated.

Costa pellucid reaching nearly to leaf tip and 35-50um wide at base.

1 2

Fig. 45. Papillaria leuconeura (C. Muell.) Jaeg. HS 29717: 1. Branch (x23); 2. Branch tip (x27). The genus Papillaria 166

Apical cells variable 15-25x2um, in extreme tip quadrate, upper laminal cells 15-23x2um, median cells 10-13(23)x3um. All cells narrowly elliptic, sigmoid and very incrassate with numerous papillae. Juxtacostal cells near base elliptic up to 63x4um, thin walled or slightly incrassate, at base rectangular 25x7um. Divergent cells in mid-section of auricle nearly uniform 10x3um, with margin of 1 or 2 rows with linear-elliptic cells up to 37x2-3um. Larninal margin with up to 4 rows of non-papillose thicker straight cells. Cells at basal margin of auricles..± quadrate, on decurrent strip narrowly elliptic. Apparent alar cells distant from margin, irregularly quadrate to rectangular 10x7um, slightly brownish.

Stem leaves similar to branch leaves but ovate, sometimes panduriform, 1.7-2.lx0.7-1.0mm, tip narrowly acurninate, base auriculate. Cells in stem leaves, longer except in tip, upper cells 30x3um, median cells 15-25x2-4um, divergent cells in mid-section of auricle slightly bigger. Cells not so prominently sigmoid and slightly less incrassate.

Base of branch with smaller rounded leaves, ecostate, denticulate in lower half. Upper laminal cells up to 25x4um, irregularly elongated, 2 or 3 papillae per cell. Median cells up to 37x5um irregularly elongated or rhomboidal, papillose. Basal juxtacostal cells rectangular, up to 63x6um, smaller near base. Small mainly rectangular..± alar cells 10x5-7um. Lower 1/3 leaf near centre with no papillae, 1 or 2 papillae towards median section and margin.

Branch cross-section ovate to circular up to 175x142um. Epidermal cells similar to outer cortical cells, irregular, 3x2um incrassate forming a band ..± 4 cells wide (1/3 of radius), cell size increasing towards centre. Inner cortical cells up to 13x7um, very irregular, thin walled. Central strand of slightly smaller cells present. The genus Papillaria 167

Fig. 46. Papillaria leuconeura (C. Muell.) Jaeg. 1-6. Branch leaves (HS 28696, 29717, 29699, Norris 34472, ibid, HS 28696); 7,8. Leaf apex (HS 29717, Stone 15767); 9,10. Upper cells near mid-leaf (Stone 15767, HS 28696); 11,13. Median cells near mid-leaf (Stone 15767, HS 28696); 12. Median margin cells (HS 29717); 14. Lower basal cells (Thor 4953); 15. Basal cells (HS 29717). The genus Papillaria 168

Rhizoids in dense clusters, slightly longer than leaves and 12um wide, cells up to 125um long with rounded overlapping ends.

Perichaetia bud like, covered by branch leaves. Inner perichaetial leaves lanceolate-ovate, up to 900x400um, tip acuminate, ecostate, slightly papillose in median section. Margin entire or slightly denticulate near tip, pronounced in the shorter rounded perichaetial leaves, short rectangular marginal cells in upper half. Leaf tip cells variable, elongated-hexagonal, up to 30.0x7.5um, median cells elongated up to 63x6um, towards the base shorter and rectangular. Archegonia ..±. 9 per perichaetium, 500x50um with irregular sides, neck up to 30um wide, cell pattern hard to see but irregularly quadrate and rectangular. Paraphyses very few, as long as the archegonia, 17um thick, cells rectangular up to 50um long with thick walls. At base of sporophyte perichaetial leaves oblong­ lanceolate up to 1.6x0.5mm, tip slightly acuminate, costa 1/2 leaf length with long rectangular marginal cells. Paraphyses numerous, elongate reaching above the perichaetium, 3 cells (20um) wide.

Capsules erect, ovoid ea 1.7xl.0mm, setae smooth 4-Smm long. Exostome teeth linear-lanceolate, densely papillose, pellucid, yellowish. Basal membrane of endostome high, segments linear, almost as long as the exostome teeth, perforated and densely papillose. Spores 15-24um, papillose.

Chromosome number: no reports.

DISTRIBUTION and ECOLOGY: Eastern Australia - Queensland (Fig. 47). This species has also been reported from India, Vietnam, Indonesia, Papua New Guinea (Streimann 1988) and New Caledonia.

The limited data available indicate that this species prefers the rain forest or poor rain forest. Generally it is found trailing from tree or shrub The genus Papillaria 169

branches, but it has been collected from a vine and a fallen tree branch. Some collections were mixed with Barbella nitens. Papillaria leuconeura has only been observed in sparse, scattered colonies. The altitudinal range is between 550- 1140m.

E1YMOLOGY: from the Greek, leukos = white or pale, and neuron = nerve, referring to the paler nerve.

ILLUSTRATIONS: Dozy & Molkenboer 1864:pl.209.

TYPE MATERIAL

The type of this species could not be located despite extensive correspondence with numerous European herbaria. Dr Noguchi (pers. comm.) also was unsuccessful in locating the type. No lectotype has been nominated because a search of European herbaria would be required to locate Blume's

Java collections which were annotated by C. Mueller. Most of Blume's collections are no longer at Leiden (pers. comm. A. Touw).

NOTES

Papillaria leuconeura collections were borrowed from Land NY, as none were available in Australian herbaria, and they all showed the characteristic cell shape. This species is very close to P. amblyacis and some Australian collections had been named as P. leuconeura in the belief that they were synonymous because the latter name has priority by 19 years. Characters separating the species proved to be constant with no intermediate types. For a discussion of the differences see P. amblyacis (p. 110). 17zegenus Papi/laria 170

' ,,,rE. 'J-,' 150" 10•

.... I I' I 0 I I I / I 0 / I ,-z.:r .... I G' II I I I I '' I 20· ,, NT If I I I I I I QUEENSLAND 2s• I I ------, 25• I I I I < 0:: --- - '' ----- ____ .,,--...... 30• I I ' ,-' I I I <, , __ ; 30-- SOUTH I AUSTRALIA I I I NEW SOUTH WALES ' I, ....-, �.... o / ADELAIDE I I ';• I '-, '---- I \ .J-- - '\ • �A'NBERRA 35• --,� I �---� ,, I VICTORIA ---�---'c. --,,\ :

40"S 0 \) 40"S

0 100 200 JOO '40llm HOBART Simple conic projection TASMANIA 1"1"E 145•V ,so• ,s�·

Fig. 47. Distribution of Papillaria /euconeura (C. Muell.) Jaeg. The genus Papillaria 171

The sigmoid appearance of the cells is due to irregular thickenings of the cells and rarely are they smoothly S shaped. Sometimes the sigmoid appearance was not present in cells of the lower median section. The divergence of the auricular cells often carried into the juxtacostal cells in the median section. In Australian collections the costa was less prominent and appeared thinner than in P. amblyacis. Specimens of P. leuconeura both from Australia and overseas exhibited twisting where there was a slight fork in the costa similar to that in the acuminately tipped P. amblyacis. P. leuconeura does not appear to have porous cells, the leaf is not smoothly convex on the back and in extreme cases it can be confused with P. crocea, but this latter species is plicate on both sides of the costa.

One mature sporophyte was noted in a July collection, while a very immature sporophyte was noted for February.

The small basal branch leaves of these species were also different. In P. leuconeura the smooth elongated cells were only in the lower 1/3 of leaf with smaller cells towards the margin and median sections, while in P. amblyacis these elongated cells covered 1/2-2/3 of lamina with very little differentiation at the margins.

SPECIMENS STUDIED: Australia 14 (10 cited), Indonesia 4, Papua New Guinea 2, New Caledonia 2.

QUEENSLAND: Windsor Tableland, Stone 16039 (MELU), 21 Aug. 1979; ditto, 38km NW of Mossman, 1140m, rain forest with Agathis robusta, on fallen tree branch, HS 29699 (CBG), 26 Jun. 1984; ditto, trailing from tree branches, HS 29717 (CBG, DUKE, H, HO, L, NICH, NY, RNG), 26 Jun. 1984; Mossman R. Gorge, ea 80km N of Cairns, rain forest edge, hanging from branches of shrubs, van Zanten 68981 (CBG, GRO), 19 Jui. 1968; Mt Lewis, Stone 15767 (MELU), 3 Aug. 1979; Juccabine Ck, Kirrama Rd, Cardwell Range, 27km WNW of Cardwell, 550m, poor rain forest beside stream, on a vine, HS 28688 ( + Barbella nitens - CBG), 20 Jun. 1984; ditto, trailing from shrub branches, HS 28696 ( + Barbella nitens - B, CBG, H, HSC, L, The genus Papi//aria 172

NICH, NY, PRE), 20 Jun. 1984; Binna Burra, 900m, rain forest, on boulder, Norris 34472 (NY), 22 Feb. 1974; Yargabullang Lookout, south of Binna Burra, 1000m, open windswept bushland, pendant on twigs, Norris 34566 (NY), 23 Feb. 1974. The genus Papillaria 173

7 PAPILLARIA ZELOFLEXICAULIS sp. nov.

Ad P. flexicaulem similaris sed a hac specie caulibus gracilioribus, foliis brevioribus et angustioribus et costis prominentioribus differt.

I Typus: 1 Richmond River, New South Wales, Australia, C. Moore. HOLOTYPUS: NY!; ISOTYPUS: BM!

Very slender plants, light green to dark green in colour, becoming brownish-green with age. Primary stems up to 30cm long, secondary stems to 20cm with branches up to 2cm long at irregular intervals, acute at apex. Stems and branches terete and julaceous, on older branches tips cuspidate while on growing branches tips thickened, with slightly spreading leaves. Leaves erect and densely imbricated, slightly erecto-patent (± 45°) when wet and in 5 apparent rows.

Fig. 48. Papillaria zeloflexicaulis Streim. (x4 - Holotype: NY; Photo: H. Lepp). The genus Papillaria 174

Branch leaves, concave, cordate to lanceolate (0.3)0.5-0.9(1.l)x0.Smm, not plicate, tip acute, base cordate, leaves more slender on trailing branches. Margin entire, rarely denticulate at base. Ratio of leaf width to length ea 0.49. Costa 2/3 leaf length, reddish-brown, often slightly twisted above mid-leaf, thick, 4-6 cells(± 20um) wide at base, cell size variable up to 40um long.

Apical cells (7.5)10.0-12.5x5.0um, median cells similar. Marginal cells quadrate (7.5um), juxtacostal cells becoming elongated-elliptic with rounded ends. Basal juxtacostal cells up to 38um long, diverging towards margin which they intersect at right angles. At base, cells.± rectangular, smooth. Basal comer cells elliptic 5-7um long, at extreme ends often .± quadrate. Papillae absent from apical cells and 1 or 2 rows near margin but with no noticeable margin. Cells equally and slightly incrassate throughout, papillae not obscuring cell outlines.

Stem leaves similar to branch leaves except lanceolate-ovate l.1-1.5x0.5- 0.6um, tip acute, longer than on branch leaves, base cordate. Papillae rarely hard to see, cells not obscured, margin rarely weakly denticulate. Apical cells irregular often club shaped, 12.5-20.0xS.Oum, incrassate, no papillae. Upper cells elongated, slightly irregular in outline 12.5-17.5x5.0um, median cells similar, juxtacostal cells nearer base up to 25x5um, diverging to margin, cells at leaf base rectangular 25.0x7.5um. Cell walls of uniform thickness.

Branch cross-section ovate to circular up to 100x75um. Epidermal cells similar to outer cortical cells which are.± 2 layers wide, cells 3x2um, incrassate, size increasing towards center. Inner cortical cells up to 10x5um, incrassate. No clear division between outer and inner cortical cells. The genus Papillaria 175

0 0 in

1 4

Fig. 49. Papillaria zeloflexicaulis Streim. 1-6. Branch leaves; 7,9. Leaf apex; (1-7,9. Holotype); 8. Upper cells; 10. Median cells; 11. Cells in auricle; (8,10,11. Isotype). The genus Papillaria 176

Fig. 50. Papi/laria zeloflexicau/is Streim. 1. Branch (x44); 2. Base of branch (x44); 3. Papillose leaf surface (x533); 4. Spore (x960); 5. Branch and sporophyte (xll); 6. Archegonia and perichaetial leaves (x137); 7. Branch leaves (x120); 8. Peristome teeth (x207); 9. Archegonia (x270); 10. Capsule (x37); 11. Capsule surface (x98); (1-3,5,8,10,11. Watts 1728; 4. Watts 1687; 6,7,9. Watts 5673). The genus Papillaria 177

No conspicuous whorl of smaller, rounded leaves at base of branch. Leaves rounded, small (± 350um) sparsely scattered, cells papillose but ecostate.

Rhizoids not common; small, in short clusters, 10.0-12.Sum wide, cells prosenchymatous up to 180um long, outlines hard to see.

Perigonia opposite or single on branches. Inner perigonial leaves fragile, broadly ovate, 500-600x250um, tip short and rounded, ecostate or very faint up to 3 / 4 leaf length. Leaf tip cells .± incrassate, irregularly quadrate to rhombic, shorter than the cells in upper part that are 12x5um, hexagonal, smooth. Median cells 12-23x6-10um, rhomboidal with one papilla on middle of cell, basal cells near centre up to 53x5um, becoming rectangular at base. Margin entire, rectangular marginal cells, if present, then weakly developed in basal region. Antheridia .± 5 per bud, up to 330x130um. Paraphyses few, as long as the antheridia, 15(20)um wide, cells up to 40um long with thick walls. Young perichaetia bud like, elongated. Inner perichaetial leaves ovate, often shouldered, 550x250um, tip acuminate, costa faint, .± 3 / 4 of leaf length, very weakly papillose. Margin entire, upper margin with thin elongated cells, in lower half quadrate to shortly rectangular 12x10um, often several rows wide. Upper cells variable, elongate 25x2um, median cells 20x7um rhomboidal, similar towards base. Archegonia .± 5 per bud, up to 280um long, 50um wide at venter, width of neck 25um, cells shortly rectangular. Paraphyses long, twisted, 2 cells wide (17um), cells up to 63um long with thick walls, very small quadrate cells at base. Single cell paraphyses not noted, young archegonia without paraphyses. At base of capsule inner perichaetial leaves obovate-lanceolate, up to 1.7x0.4mm, entire, costa 3/4 leaf length or to apex, cells weakly papillose especially near the costa, base generally not papillose. Apical cells variable, elliptic, up to 50um long; upper and median cells 40(87)x5um, at mid-leaf The genus Papil/aria 178

juxtacostal cells tend to be rectangular; nearer margin, cells are rhomboidal 25x5um; basal cells rectangular up to 80x15um. Margin of smaller quadrate or shortly rectangular cells. Paraphyses clear up to 25um (2 cells) wide in lower half, upper half one cell wide, cells quadrate or predominantly rectangular up to 65(88)um long, sometimes bulging at nodes.

Calyptra ea 1.8mm long. Capsules erect, ovoid (1.0)1.3(1.5)x0.5-0.8mm, setae (2.0)2.5-2.7(3.0)mm. Exostome teeth papillose, linear-lanceolate up to 420um, width at base 75um; endostome segments papillose, linear 370x10um. Operculum 0.6mm long with a long narrow curved beak. Exothecial cells rectangular 30x10um near top, irregularly quadrate to rounded(± 30um) in mid section. Endothecial cells similar but .± obscure. Spores 12-18(23)um, papillose.

Chromosome number: no reports.

DISTRIBUTION and ECOLOGY: Eastern Australia - Queensland and New South Wales (Fig. 51). This species is endemic to Australia. Habitats are lowland rain forest, fringing rain forest and Araucaria vine thickets. Most commonly it is reported as trailing from branches and twigs; once from a fallen tree branch and less specifically on 'tree' and 'sapling'. It has been found mixed with Papillaria amblyacis, P. crocea. P. flexicaulis, Barbella cubensis and B. nitens. Recorded altitudinal range was up to 200m.

ETYMOLOGY: from the Greek, zelo = emulate, envy or rival, referring to the plant having the appearance of Papillaria flexicaulis. 77,egenus Papillaria 179

.,,11 1')1}

1().

Cooktow11

0 I 0 '\ CJ1rns I ' I 0 ,,// I ,;--:i., I : G" I I I I I NT I I I 20' I Mackay I I I QUEENSLAND I I 25' I I ------, 25' I I I z I < I - ' ------Q: ---- -.,, -...... ,,-.,-- I --- '-,/"•;, • SOUTH I AUSTRALIA I I I I I NEW SOUTH WALES I � \ ' I, __ �- o / ADELAIDE I I �---·:· , ', \ - '\ ·�ANBERRA ' ..1 -.... :w I VICTORIA --,�� \ : I �,---"':,. / --J·�-- -,,

40'S

40' S

0 100 200 300 ,oo•m s;mp1e con.: p,OJKlion

140'E 150' 15:,·

Fig. 51. Distribution of Papillaria zeloflexicau/is Streim. in Australia. TTie genus Papillaria 180

NOTES

Most specimens were previously determined as P. flexicaulis to which this species bears a superficial resemblance in its thin trailing sterns and branches. However, the branch leaves are always smaller (P. flexicaulis 0.5-l.7x0.4-l.2rnrn, P. zeloflexicaulis 0.4-0.9x0.3-0.5mm) and generally a lighter green. The leaf cells

are also similar and intersect the margin at right angles as they do in P. flexicaulis, but the papillae reach lower down the margin in the latter species. However, the costa appears conspicuously thick and quite often exhibits a twist

above the middle as in P. crocea. Around the margin, cells are quadrate especially nearer the base of the smaller leaves (Camara 37), then it appears quite unlike P. flexicaulis.

In Porter (as Watts 2281) there were several groups of abnormal branches. These branches were shorter and thicker than normal with several branches emerging around the main branch. These branches again sub-divided into smaller thicker branches. This produced a 'gall-like' appearance.

Capsules appear to mature during September and October.

Comprehensive habit, habitat and altitudinal data are lacking because the majority of the collections were at least 80 years old. However, it appears that this species may have been present in the lowland forests of northern New South Wales and southern Queensland most of which have been cleared for agriculture.

SPECIMENS STUDIED: Australia 47 (10 cited).

QUEENSLAND: Eumundi, Staer (NSW), 28 Mar. 1910; Mt Perry, Kalliwa Ck, Goodnight Scrub S.F. 80m, Araucaria vine forest, hanging from branches, common, Foster 801 (NSW), 2 Jui. 1981; Brisbane, Bailey (MEL 32156), 1886; Mudgerabah, Wild (BRI 331599), 29 Mar. 1889. NEW SOUTH WALES: Tintenbar, Richmond R, on trees in scrub, Watts 752a (H, NSW), Mar. 1896; The genus Papillaria 181

O'Donnell Ck, Toonumbar S.F., 10 miles [16km] W of Wiangaree, dense rain forest on basalt, frequent on shrub branches, Constable 4882 (NSW), 26 May 1964; Tenterfield-Casino, Porter (as Watts 2281 - NSW), Oct. 1898; Upper Thora Rd, 28km W of Bellingen, 100m, fringe of rain forest, on a twig, Barnsley 178 (CBG), 16 May 1978; Wilson R, 200m, on fallen tree branch, Schofield et al. 80881 (UBC), 13 Jui. 1983; Wyong, Watts 9501 (NSW), no date. The genus Papil/aria 182

UNKNOWN PAPILLARIA SPECIES

The following collections could not be placed into presently recognized Australian species, nor those described by Noguchi (1976) for Asia, nor were similar plants seen in the available Pacific Islands collections:

Stone 15785 ( + P. amblyacis - MELU ): has an appearance of P. crocea but with a long wavy apex. Branch leaf cells however were not similar to P. crocea, especially in the apex where the cells were long and narrow. Material inadequate. Mt Lewis N.P., Queensland, 3 Aug. 1979.

Stone 20880 (MELU): leaves similar to P. crocea but cells like thicker P. /euconeura cells. Papillae small, dense, extending to auricles. Material inadequate. Mt Mistake N.P., Queensland, 26 Feb. 1983.

Merrah (MEL 32177): has the appearance of P. flavolimbata, but with long narrow incrassate cells, this feature is often masked by dense papillae. Mt Ellery, East Gippsland, Victoria, 1887. The genus Aerobryopsis 183

II. The genus AEROBRYOPSIS Fleisch. Hedwigia 44:304.1905

Type: non designatus.

Plants medium-sized to large. Primary stems elongate, laxly or densely branched and leaved. Secondary stems elongate, with several branches, laxly or densely leaved; branches short, obtuse at apex. Stem-leaves appressed or complanately spreading, ovate to oblong, gradually or rapidly attenuate to a long, straight, or flexuose, or crisped acumen, cordate at base, concave .± undulate above; margin erect or somewhat undulate in median region, serrulate or serrate throughout; costa slender, extending beyond 2/3 leaf-length. Laminal cells rhomboidal to elongate-rhomboidal, with a distinct papilla over centre of lumen, walls thick, subporose, middle lamellae visible; apical and basal cells larger without papillae; cells in alar region rectangular or subquadrate but not forming a distinct group. Branch-leaves similar to those of stem, but wide­ spreading.

Rhizoids smooth, red to reddish-brown.

Dioicous. Perigonia and perichaetia on branches. Inner perigonial leaves ovate, cymbiform. Inner perichaetial leaves oblong-sheathing, acumen pointed, short or long, costa weak; paraphyses short. Calyptrae cucullate, smooth. Capsules suberect or erect, oblong-cylindrical, with a distinct apophysis, often asymmetrical, smooth on the surface. Setae elongate, much longer than the capsule, slightly scabrous. Operculum conical with a long, somewhat oblique beak. Annulus present only 1 per capsule. Peristome double, exostome teeth long, linear-lanceolate, papillose throughout; segments mostly linear, slender as long as the exostome teeth, often perforated along the The genus Aerob,yopsis 184

keel; cilia none; basal membrane rather low. Spores rounded, medium sized, 12-20um, papillose.

Chromosome number: n = 11 (Japan, Inoue & Momii 1971).

E'IYMOLOGY: from the Greek, aeros = aerial and the Greek, bryo = relating to mosses, referring to trailing habit. The genus Aerobryopsis 185

AEROBRYOPSIS WALLICHII (Brid.) Fleisch, Musci FI. Buitenzorg 3:789.1908.

Hypnum wallichii Brid., Bryol. Univ. 2:416.1827. Type: In regione Nepal Indiae Orientalis, Wallich 1825. HOLOTYPE: FH. Meteorium sigmatophyllum C. Muell. ex Bailey, Syn. Queensland FI. Suppl. 3:98.1890, nom. nud. Aerobryopsis longissima (Dozy & Molk.) Fleisch., Hedwigia 44:305.1905. A. sigmatophylla Broth. & Watts, Proc. Linn. Soc. New South Wales 43:559.1918, nom. nud., syn. nov.

Plants robust, slightly glossy, yellow-green. Primary stems long creeping, secondary stems pendulous, long flexuose up to 50cm, both appressed or complanately leaved; distantly branched, branches den~ely appressed and sometimes complanately leaved, narrowing towards tip. Creeping stems usually with denser branches and leaves than those hanging. Complanate stems and branches with fewer and smaller leaves.

Branch leaves ovate-lanceolate, (1.5)1.9-2.7(3.3)x(0.5)0.7-l.0mm, gradually tapering to an elongated acumen, often flexuose, if so then leaf is also undulate in upper median section. Margin se"ate in upper section, less so and distantly in lower section but often hard to see. Costa light brown, thin, or weak, reaching 1/2(3/4) leaf length, at times papillose on both sides, but more prominent on the underside. Cells often smooth at extreme tip and near the leaf base. Towards base the papillae became smaller and harder to see.

Cells in leaf tip of variable shape and size, up to 40x7um, rarely incrassate. Cells in upper leaf linear or linear-rhomboidal, wavy or.± sigmoid, up to 35x7um, median cells similar, often not so wavy or sigmoid. Juxtacostal cells near base up to 50(75)um long, ..±. porose, becoming rectangular, at basal The genus Aerobryopsis 186

corners cells are roughly rounded to squarish 8-lOum, cell walls slightly thicker and cells.± with granular content.

Stem leaves similar in shape but larger, 2.9(3.l)x0.5-0.7-0.9(1.l)mm, tip elongated, rarely papillose, costa with few low papillae on both sides, cells at base with granular content.

Rhizoids in clusters, up to 1.1mm long and 10.0-12.Sum wide, cells up to 112um long, ends rounded or often overlapping.

Perigonia and perichaetia on branches. Inner perigonial leaves ovate 1.2x0.5mm, tip acuminate, ecostate, no papillae, margin entire with long rectangular cells in upper half. Upper and median cells linear-hexagonal, size variable ea 50x5um, basal cells .± rectangular 50x10um. Antheridia .± 18 per bud, 450x150um, reddish-brown with irregularly quadrate and rectangular cells. Paraphyses as many as, and slightly longer than, the antheridia, up to 17um wide, cells thick walled, rectangular up to 80um long. Inner perichaetial leaves acute concave up to 1.5x0.5mm, margin denticulate throughout, mostly ecostate or if costa present visible only to mid-leaf; paraphyses numerous.± 1mm long.

Capsules suberect, .± curved and asymmetric, up to 2.Sxl.0mm, brownish. Setae to 2cm long, scabrous above, nearly smooth below. Exostome teeth finely attenuate, to .± 0.6mm in length, minutely papillose throughout, papillae on the basal part arranged somewhat vertically, segments widely perforated along keel. Spores 12-18(25)um.

Chromosome number: no reports.

DISTRIBUTION and ECOLOGY: Eastern Australia - Queensland (Fig. 52). Extends from Sri Lanka, southeast Asia to Japan and throughout the Pacific to central America and the Caribbean. The genus Aerobryopsis 187

Aerobryopsis wallichii is reported from rain forest, rain forest fringes,

Araucaria plantation, edge of advanced regrowth, remnant poor scrub and

Eucalyptus woodland. Outside the rain forest it is found in areas with a suitable moisture regime, being generally reported near streams. The altitudinal range is 20-1180m.

Aerobryopsis wallichii is commonly an epiphyte on tree trunks, tree crowns, small shrubs, tree roots, rocks and boulders, rarely on vines, tree leaves and rotting logs. Trees on which it has been reported are Araucaria, Endiandra and Ficus.

E1YMOLOGY: named after a Danish doctor N. Wallich ( = N. Wolff), 1786-

1854, who worked and collected mosses in India.

ILLUSTRATIONS: Bartram 1933:170, 1939:pl. 16, figs 273,274 (A. philippinensis Bartr.); Brotherus 1906:819, 1925:166; Dozy & Molkenboer

1848:159, pl. 48 (Meteorium longissimum (Dozy & Molk.) Dozy & Molk.), 1864:pl. 202 (M. longissimum var. tenue Bosch & Lac.); Fleischer 1905:305 (var. tenue), 1908:781; Gangulee 1976:1319,1320 (Aerobryopsis wallichii); Johnson

1980:8; Noguchi 1976:296,297 (A. wallichii); Norris & Koponen 1985:37; Sakurai

1943:254 (A. pemitens Sak.); Whittier 1976:146. All as Aerobryopsis longissima except where stated otherwise. n,e ge1111s Aerob1)1opsis 188

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40"S � TASMANIA "' 0 100 200 300 ,ooi.m A Simple conie p,Ofe<:lion 140"£ V 1:5. :so· ,s,;·

Fig. 52. Distribution of Aerobryopsis wal/ichii (Brid.) Fleisch. 17ie genus Aerobryopsis 189

NOTES

This species can be as variable in Australia as Bartram (1939) found it to be in the Philippines where he found the robust form uncommon whereas in Australia the robust form is common. Previously A. longissima was considered to be a robust plant with squarrose leaves while A. wallichii was a slender plant with leaves nearly complanate. The former widespread species was considered by Noguchi (1976) to be a synonym of A. wallichii as he interpreted the many variations as only ecological modifications. In Australian collections both slender and robust forms were often found together. Robust forms tended to exhibit transverse undulations more so than slender plants. Leaf areolation was found to be the same in both forms. While slender forms could perhaps be mistaken for Barbella cubensis this species never exhibits the transverse undulations seen in the upper part of the leaf of Aerobryopsis wallichii.

The black tinge was not common in Australian plants, and the leaf length of the Australian plants (3.5mm) was shorter than that reported by Bartram (1939) for the Philippines (4.0mm) and by Norris & Koponen (1985) for Papua New Guinea (4.5mm).

No collections with perigonia, perichaetia or sporophytes were found amongst the Australian material. It is interesting to note that Norris & Koponen (1985) did not find fertile material amongst their Papua New Guinea collections.

Townsend (1978) gives a lengthy outline on the naming of this species and he concluded that this species is best placed in Aerob,yidium and made the new combination accordingly. However, Norris & Koponen (1985) have preferred to retain this species in Aerob,yopsis. 17ie genus Aerob,yopsis 190

Whilst many mosses are known to accumulate metals, e.g. Racomitrium lanuginosum (Hedw.) Brid. - lO00Oug/g zinc, 6000ug/g copper, 4900ug/g

uranium, Campylium polyganum (B.S.G.) C. Jens. - 6000ug/g nickel (Lee et al. 1977), but chromium has not been one of these. However, Lee et al. ( 1977) reported unusually high concentrations, 7500ug/ g chromium, for this moss in New Caledonia.

SPECIMENS STUDIED: 143 (21 cited).

QUEENSLAND: Big Tableland, 26km S of Cooktown, 590m, edge of stunted rain forest and grasslands, base of treelet, HS 30849 (CBG, H, NICH, NY), 4 Jul. 1984; Cedar Bay Rd, Stone 15848 (MELU), 6 Aug. 1979; Windsor Tableland, 1080m, rain forest beside stream, on a vine and shrub branches, HS 29676 (CBG), 26 Jun. 1984; Mossman Gorge, outcrop and on trees, W.B. & M.I. Schofield 79901 (UBC), 27 Jun. 1983; Mary Ck, Main Coast Ra., 15km NNW of Mount Molly, 1080m, rain forest, on a large boulder, HS 30398 (B, CBG, NICH, NY), 30 Jun. 1984; Mt Lewis, on leaf, Stone 15773 (MELU), 3 Aug. 1979; Black Mountain, 25km NW of Kuranda, 500m, 'dry' rain forest, on treelet stem, HS 31109 (CBG), 7 Jui. 1984; Davies Ck Rd, on fallen tree, Stone 12252 (MELU), 31 May 1975; Cairns, Wild (BRI), Aug. 1890; Coastal Escarpment, Gillies Rd, between Gordonvale and Atherton, grassy Eucalyptus forest, along a cascade (Huntsbrook Ck), on a rock, Weber B-31893 (COLO), 17 Apr. 1968; Harvey Ck, 22km SE of Gordonvale, 20m, lowland rain forest, buttress & trunk of large tree, HS 27320 (CBG, H, L), 3 Mar. 1983; Mt Baldy, 4km SW of Atherton, 1080m, Araucaria cunninghamii plantation with regrowth rain forest, on Araucaria trunk, HS 29265 (B, CBG, H, L, NICH, NY), 25 Jun. 1984; L. Eacham, 16km E of Atherton, 800m, poor lowland rain forest, on Ficus stem, HS 16856 (CBG, L), 28 Feb. 1983; Boulder Reserve, Babinda, 80m, lowland forest alongside creek, on tree trunk, Kantak & Churchill 924 (CBG, NY), 25 Apr. 1984; Bonjie S.F., 22km SE Yungaburra, 600m, logged rain forest on flats, on Endiandra muelleri branch, HS 27545 (CBG, H, L, NICH, NY), 3 Mar. 1983; Cooroo logging area, 16km WNW of Innisfail, 100m, rain forest, on tree trunk, HS 29986 (CBG, L), 27 Jun. 1984; Maalan, 11km E of Ravenshoe, 980m, remnant scrub between road and paddock, on a shrub, HS 30598 (B, BA, BISH, CBG, DUKE, H, L, NAM, NICH, NY, RNG), 30 Jun. 1984; Walter Hill Ra., 25km SE of Ravenshoe, 780m, rain forest, on tree buttress, HS 30526 (CBG, NY), 2 Jui. 1984; Koombooloomba Dam Rd, 23km SE of Ravenshoe, 780m, rain forest, on treelet stem, HS 28895 (CBG, H, L, NICH), 23 Jun. 1984; Yuccabine Ck, Kirrama Rd. Cardwell Ra., 27km WNW of Cardwell, 550m, poor rain forest beside stream, on tree trunk, HS 28700 (CBG, H, L, NICH, NY), 20 Jun. 1984; Old Mill Rd, Blue Water Ck, 39km WSW of Ingham, 600m, rain forest, on shaded rock beside stream, HS 28408 (CBG), 19 Jun. 1984. The genus Barbella 191

III. The genus BARBELLA Fleisch. Nat. Pfl. 1(3):823.1906 Type: non designatus.

Plants medium sized. Primary stems filiform, elongate, laxly leaved, subpinnate or pinnate, obtuse, apical part of both primary and secondary stems often pendant, with few branches. Secondary stems with complanate and spreading leaves on basal part; filiform part of branches with appressed leaves. Stem-leaves appressed or laxly spreading both dry and moist, lanceolate, ovate­ oblong, ovate, or oblong, attenuate to an elongate, subulate or piliferous apex, base hastate to cordate; margin erect, often ..±. undulate at median portion, serrulate or entire; costa slender, reaching to mid-leaf, indistinct or none. Lamina! cells very translucent, linear to oblong-rhomboidal, smooth or with one or more, often inconspicuous, papillae in center of lumen, often with a thread­ like, plasmolized protoplast, thin-walled, walls often with localized thickenings; cells of leaf-insertion wider than median cells, rectangular, with thickened walls; alar cells lax, rectangular or subquadrate, forming a well-defined alar group, or sublinear, not forming a specialized alar group. Basal cells divergent or not.

Branch-leaves complanately spreading when dry or moist, rarely appressed, similar to those of stem in outline, but usually with sharper marginal teeth. Laminal cells similar to those of stem-leaves, but papillae more distinct and alar cells usually less differentiated.

Rhizoids short, smooth and reddish, margins thickened.

Dioicous, rarely autoicous. Perigonia on branches. Inner leaves widely ovate, shortly acuminate, cymbiform, accompanied by a few or no paraphyses. Perichaetia on branches or stems. Inner perichaetial leaves with or without paraphyses. Calyptra mitriform and lobate at base, or cucullate, smooth or The genus Barbella 192

hairy. Capsule erect, oblong to oblong-cylindrical, smooth. Seta short, almost as long as the capsule or longer, often curved, smooth or scabrous. Operculum rostrate. Exostome teeth usually linear-lanceolate, fairly elongate, usually papillose throughout, sometimes papillae arranged in diverse rows in lower portion, lamellae high; basal membrane moderately high; segments long, almost as long as exostome teeth or a little shorter, linear to linear-lanceolate, usually perforated along the keel, minutely papillose. Spores rounded, medium sized to large, 12-35um, papillose.

Chromosome numbers: n = 9, 10 (India, Sing & Gangulee 1969; Japan, Inoue 1965).

E1YMOLOGY: from the Latin, barbatus = bearded or long tufts of weak hairs, referring to the long slender trailing habit.

KEY to AUSTRALIAN SPECIES of BARBELLA

la Branch leaves concave, closely imbricate, base without auricles, costa absent or very weakly developed ...... (1) B. CUBENSIS

lb Branch leaves with margins bent back abaxially, weakly imbricate or spreading, base auricled, costa 1/2 leaf length ...... (2) B. NITENS The genus Barbella 193

1 BARBELLA CUBENSIS (Mitt.) Broth., Nat. Pfl. 1(3):824.1906.

Meteorium cubense Mitt., J. Linn. Soc., Bot. 12:435.1869. Type: 'Ins. Cuba, Wright, n. 82'. HOLOTYPE: BM. Meteorium enerve Thwait. & Mitt., J. Linn. Soc., Bot. 13:317.1873. Type: 'Thwaites 208', without definite locality. HOLOTYPE: NY. Barbella enervis (Thwait. & Mitt.) Fleisch., Nat. Pfl. 1(3):824.1906. Neckera trichophoroides Hpe ex C. Muell., Linnaea 38:668.1874. Type: 'Lord Rowe's Island, Summit of Mount Gower'. HOLOTYPE: BM!; ISOTYPE: MEL (32292)! Pilotrichella trichophoroides (Hpe ex C. Muell.) Jaeg., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1875-76:259 (Ad. 2:163). Meteorium trichophoroides (Hpe ex C. Muell.) Mitt., Trans. & Proc. Roy. Soc. Victoria 19:82.1882. Barbella trichophoroides (Hpe ex C. Muell.) Broth., Nat. Pfl. 1(3):824.1906. Meteorium dicladioides, C. Muell. ex Geh., Rev. Bryol. 3:4.1876, nom. nud. M cinclidotus C. Muell. ex Bailey, Syn. Queensland FI. 733.1883, nom. nud. M. compressum Mitt., Trans. & Proc. Roy. Soc. Victoria 19:83.1882, syn. nov. Type: 'Brisbane River, Bailey'. HOLOTYPE: ?NY. Papillaria compressum (Mitt.) Kindb., Enum. Bryin. Exot. 102.1891, syn. nov.

Plants slender, glossy light green to yellow-green. Primary stems up to

20cm long, sparse or densely irregularly branching, laxly appressed or spreading, apex filiform with leaves appressed on pendant sections. Secondary stems with The genus Barbella 194

few branches, widely and complanately leaved. Branches short, up to 2cm long, obtuse or flagelliform, at times complanate.

Primary stem leaves lanceolate, 2.8-4.0x0.7-0.9mm, apex acuminate to filiform, concave, broadest at hastate base. Margin erect or incurved from apex to median section, variously denticulate at base, sometimes with weak denticulations extending to leaf apex. Costa absent or weakly developed. Secondary stem leaves similar except that a weak costa may be present. Branch leaves similar to those on the stem, rarely piliferous, 2.5-3.4x0.3-0.5mm. Leaves at pendant branch tips 2.5-3.lx0.3-0.4mm, 1/2 of leaf being the narrow tip, which often has prominent denticulations.

Branch leaf cells pellucid, weakly unipapillose and thin walled. Upper leaf cells prosenchymatous, linear to sub-linear (45)75-100(160)x7-10um, outline often hard to see. Median cells 100-130x7um; juxtacostal basal cells .± lOOxlOum, cell walls thicker(± Sum); alar cells irregularly angular, well defined extending nearly to middle of leaf base with thick to thin cell walls. All basal cells seem to have darker granular deposits, these being most prominent in older cells.

At base of branches small, inconspicuous, widely ovate to rounded oblong leaves, 500x350um, ecostate, cells smooth and margin denticulate. Cells in upper leaf linear, rarely irregularly rhombic, 37-50x7um, median cells 37- 67x10um cells at basal angles rectangular (± 25um long) and square (± 17um) with a golden-brown colour.

Rhizoids matted in rounded 'pads' 0.5mm across, cells up to 600um long, ends rounded often overlapping. The genus Barbella 195

Perigonia on secondary stems and branches, inner leaves.± 1.5mm long, paraphyses few. Calyptra up to 1.2mm long. Capsules oblong 1.5-2.0x0.9mm, seta up to 2mm long. Opercula up to 1.2mm long. Exostome teeth yellowish, linear-lanceolate .± 0.65mm long, finely attenuate and pellucid at apex.

Horizontally and densely papillose-striate in lower half with large papillae above, segments.± 0.45mm long. Rudimentary cilia sometimes present. Spores 20-35um.

Chromosome number: po reports.

DISTRIBUTION and ECOLOGY: Eastern Australia - Queensland, northern New South Wales (Fig. 53) and Lord Howe Island. Outside Australia it has been reported from India, Sri Lanka, Thailand, Japan, China, Formosa, Philippines, Indonesia, New Guinea, Central America and the Caribbean Islands. Menzel & Schultze-Motel (1987) reported it as a rare species for Peru. Barbella cubensis is a rain forest moss which has been reported previously only as a pendant epiphyte from shrub, treelet and tree branches, but it also occurs on vines, in tree crowns and was collected once on rotting wood. This moss is common in the montane forest on Lord Howe Island. In Australia it has been collected from near sea level to 1080m.

E'IYMOLOGY: named after Cuba from where the type originated.

ILLUSTRATIONS: Bartram 1939:pl. 17, fig. 283, 1949.271; Brotherus 1929:pl. 2 (Barbellopsis sinensis Broth.); Gangulee 1976:1327; Noguchi 1948:76 (Barbella determesii (Ren. & Card.) Fleisch. ex Broth.), 1965:148, 1968:315; Norris & Koponen 1985:43; Sakurai 1938:471 (Aptychopsis uii Sak.). 77,egenus Barbella 196

j,l'f t� 'J·, ·�· \

G') ::0 "" 1"....

IS° 0 I 0 I I o , • I ,''1-r • I , G' N I \ \ NT I � I \ I ',, I \ I \ I QUEENSLAND \ I \ I \ 25• I \ I \ ------, \ 2s• I , _____ ,,u) I c..,,,', I z \ I �

0

40"S � TASMANIA " o ,oo 200 JOO '""'"" A Simple conic p,ojec:IIOfl ,�s.V ,so- ,s;-

Fig. 53. Distribution of Barbella cubensis (Mitt.) Broth. The genus Barbella 197

TYPE MATERIAL

The type of Meteorium compressum Mitt. could not be located at NY, but from Mitten's (1882:83) description it is clearly Barbella cubensis. Several Bailey collections (from BM) annotated only as Brisbane were studied.

NOTES

Barbella cubensis can be quite variable depending on whether it has a creeping stem with short branches where the leaves are broader and shorter or has pendant, pendulous, flagelliform branches. On these branches the leaves are more appressed and narrower and 1/2 leaf length consists of a fine point with uniseriate cells up to 15 cells long. Some pendant branches (Stone 18497) had distant spreading leaves which gave the appearance of B. nitens. The difference between the leaves on the pendant and creeping sections is generally conspicuous. Denticulation is variable being formed by protruding leaf cells. Papillae are low and often very few and hard to see, being best developed on the median juxtacostal cells. On small leaves at the branch base, denticulation is variable in size and can even be different on both sides of leaf. Blackish leaves were observed rarely (Stone 16043). This species and B. macroblasta Broth., a Philippine endemic, are the only Barbella species without a costa. No perigonia, perichaetia or sporophytes were noted on Australian collections.

SPECIMENS STUDIED: Australia 118 (21 cited), New Caledonia 1.

QUEENSLAND: Windsor Tableland, 39km NW of Mossman, 1080m, rain forest beside small stream, HS 29677 (CBG, NICH, NY), 26 Jun. 1984; Mossman Gorge, Stone 15879 (MELU), 7 Aug. 1979; Mt Lewis, on road from Julatten, 910m, pendulous from upland rain forest trees, Weber B-31770 (COLO, NY), 18 Apr. 1968; Davies Ck Rd, rain forest, on fallen tree, Stone 12271 (MELU), 31 May 1975; Palm Camp, Bellenden Ker, Bailey 631 (BRI), 1889; Wright Ck, L. Eacham, 16km E of Atherton, 800m, poor lowland rain forest, on fallen tree branch, HS 16903 (CBG, H, L, NICH, NY), 28 Feb. 1983; Cooroo logging area, 16km WNW of Innisfail, 100m, rain The genus Barbella 198

forest, in crown of large tree, HS 29964 (CBG), 27 Jun. 1984; Ravenshoe-Millaa Millaa Rd, festooning shrubs, W.B. & I.M. Schofield 80145 (UBC), 30 Jun. 1983; Kirrama, Stone 16469 (MELU), 16 Aug. 1980; Mt Spee S.F., 6km W of Paluma, 920m, Lauraceae-Syzygium dominated forest, on rotting wood, HS 36979 (CBG), 18 Jun. 1986; Gympie, Musson 384 (MEL), no date; Blackall Ra., Whitteron (MEL 32298), Apr. 1898; Pine R., Whitteron (MEL 32296), Mar. 1885; McPherson Ra, Nothofagus forest, pendulous on branches, Blake 22800 (BRI), 23 Aug. 1967. NEW SOUTH WALES:· Wiangaree S.F. 27km NNE of Kyogle, 750m, Cunoniaceae-Lauraceae dominated forest, on tree trunk, HS 6047 (CBG, L, NICH), 3 Oct. 1978; Myocum, Richmond R. scrub, Watts (BM, JE, MEL), Feb. 1900; Bellinger and McLeay Rivers, Rudder (NY), no date; Mt Boss S.F., tropical rain forest, Vitt 27528 (ALTA), 10 Oct. 1981; Allyn Forest Park, festooning shrubs, Schofield et al. 80962 (UBC), 14 Jui. 1983; Hunter's R., Rudd (MEL 32286), 1890. LORD HOWE ISLAND: Mt Gower Summit, low closed forest, pendant from branches, common, Crisp 4560 (CBG, L), 23 Oct. 1978; track Erskine Valley to Mt Gower Summit, 100-875m, Vitt 28535 (ALTA, CBG), 17 Nov. 1981. The genus Barbella 199

2 BARBELLA NITENS (Hook.f. & Wils.) Nog., J. Hattori Bot. Lab. 58:100.1985.

Meteorium nitens Hook.f. & Wils., FI. Nov. Zel. 2:101.87.f.7.1854. Type: 'Northern Island, Sinclair'. LECTOTYPE: BM!; ISOLECTOTYPE: BM!. Papillaria nitens (Hook.f. & Wils.) Sainsb., Rev. Bryol. Lichenol. 21:221.1952. Meteorium fulvum Mitt., J. Linn. Soc., Bot. 4:85.1860, syn. nov. Type: 'Dr. F. Mueller, Tarwin, Victoria Australia'. HOLOTYPE: NY!; ISOTYPES: MEL (29182, 32242)! Papillaria fulva (Mitt.) Jaeg., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1875-76:267.1877 (Ad. 2:171), syn. nov. Pilotrichella fulva (Mitt.) Hpe, Fragm. Phytogr. Austral. Suppl. 11:50.1881, syn. nov. Neckera dimorpha C. Muell., Linnaea 37:517.1872, syn. nov. Type: 'Blue Mountains, New South Wales' HOLOTYPE: BM!; ISOTYPE: MEL (32249)! Pilotrichella dimorpha (C. Muell.) Jaeg., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1877-78:508.1879 (Ad. 2:772), syn. nov. Meteorium dimorphum (C. Muell.) Mitt., Trans. & Proc. Roy. Soc. Victoria 19:82.1882, syn. nov. Papillaria nitidiuscula Broth., Proc. Linn. Soc. New South Wales 41:593.1916. Type: 'N.S.W.: Richmond River, Alstonville Road (Watts, n. 4032, 4043, 4062); E. Ballina (Watts, n. 207, 3401, 4003, 4110, 4351); Tintenbar (Watts, n. 4363); Pimlico (Watts, n. 4882); Uralba Road (Watts, n. 3711, 3715); Pearce's Creek (Watts, n. 441); Brooklet (Watts, n. 4793); Teven Creek (Watts, n. 4093, 4396, 4418); North Creek (Watts, n. 4755); Brunswick River, Myocum (Watts, n. 4337)'. LECTOTYPE: Watts 4363. On bush close to ground, Tintenbar, Richmond River, July 1900- H!; ISOLECTOTYPE: NSW! P. nitidiuscula Broth. ex Dix., New Zealand Inst. Bull. 3(5):261.1927, nom. illeg. incl. spec. prior. Type: 'On trees in scrub, Ballina, N.S.W., July 1900, W.W. Watts'. HOLOTYPE: BM!; ISOTYPES: H!, NSW! The genus Barbella 200

Plants slender, flexuose, soft, light green to light golden-green, when dry characteristically golden, often slightly glossy. Primary stems to 15cm long, pendant or creeping. Branches at base of primary stem regularly spaced, very short, robust, curved and densely foliate, leaves appressed. Pendulous branches more slender, pinnately and distally branched, not imbricated.

Leaves lanceolate-subulate (1.5)1.7-1.9x0.6-0.7mm, slightly plicate, upper part contracted to a long filiform subula, base broadly triangular with inflexed auricles. On stems and older branches leaves are more laxly plicate, being the same when wet or dry. Flexuose subula is channelled with margins bending back abaxially in mid-leaf forming a 'tube'. Margin denticulate in auricles, diminishing further up lamina, costa thin, sometimes not clearly visible, 1/2 length of lamina, sometimes 1/3-3/4 length of lamina, 20um wide at base, costa cells similar to surrounding cells. In young leaves costa .± absent. Apical cells often up to 57um long, incrassate, cells in upper leaf smooth 25-37x2-3um, linear-rhomboid and some papillae developing further down. Median cells similar to upper cells but with 1-3 small papillae per cell to leaf margin, sometimes papillae hard to see. Juxtacostal cells near base up to 37um long but still similar to cells in upper part of leaf, prosenchymatous. Below this, are smaller and thicker parenchymatous incrassate cells. In inner corners of auricles a small area of quadrate cells may be present. Marginal cells not differentiated.

Rhizoids.± 600um long, 13um wide, cells to l0Oum long.

No pengoma seen. Perichaetia elongated bud like, scattered on branches. Inner perichaetial leaves ovate-lanceolate, .± shoulders, up to 930x300um, tip acuminate, ecostate, very few weak papillae in lower median section, or none. Margin entire, slightly .± denticulate above, inconspicuous The genus Barbella 201

border of rectangular cells. Upper and median cells elliptic, up to 87x6um, cells in basal corners rectangular, 37x10um, across base.± quadrate. Archegonia reddish, .± 10 per bud up to 340x40um, neck slightly thinner, with shortly rectangular cells. Paraphyses very few, one cell wide. At base of capsule inner perichaetial leaves lanceolate-acuminate 1. 7-1.8x0.4-0.5mm, ecostate and smooth.

Basal cells up to 120um long, median cells up to 90um long with slightly larger or thicker cells along the leaf margin. Paraphyses 2.2mm long, 2 or 3 cells (25um) wide, mid-cells up to 130um long, not clear and bulging at nodes.

Calyptra not seen. Capsules symmetrical, round-oblong 1.5x0.5mm when dry, when wet the width can double, setae up to 5mm, smooth. Operculum not seen. Exostome teeth 400um long and yellowish with small papillae best developed along the cell margins. Exothecial cells isodiametric, 4 to 5 sided to rounded, thin walled, becoming quadrate and rectangular nearer base, up to 50x8um. Thin spore sac has similar cells to capsule with very small papillae. Spores (immature) 12-17um.

Chromosome number: no reports.

DISTRIBUTION and ECOLOGY: Eastern Australia - Queensland, New South Wales, Victoria and Tasmania (Fig. 54 ). It also occurs in New Zealand and New Caledonia.

Commonly found in ram forest and wet sclerophyll forest, but also reported from Nothofagus forest, Cunoniaceae-Lauraceae dominated forest, Casuarina-Eucalyptus forest and a Casuarina swamp. On one occassion collected in dry sclerophyll forest beside a creek and from a heath near Ballina. Commonly on tree trunks, tree crowns or treelet/shrub branches, but also noted The genus Barbella 202

on palm leaves, vines (Ca/amus, Freycinetia and Marinda) and on a rock face. Barbella nitens has been collected from the following trees or treelets - Bosistoa, Endiandra, Eucalyptus, Ficus and Notelaea. Barbella nitens is very often found associated with Papillaria spp. and numerous other mosses.

Reported from 40m to 1370m altitude in New South Wales and up to 1220m in Queensland. In Tasmania and Victoria it occurs at lower altitudes, but is not common.

E1YMOLOGY: from the Latin, nitens = shining or lustrous, referring to the golden-yellow colour.

ILLUSTRATIONS: Dixon 1927:373 (P. nitidiuscula and Meteorium nitens); Noguchi 1985:101,102.

TYPE MATERIAL

The material of Meteorium nitens at BM could hardly be called a scrap as described in the protologue, even though it was barren. The lectotype and isolectotype chosen were 11cm and 7cm in length respectively. Both are pieces from a creeping section of a plant as evidenced by the short, dense secondary branching. The illustrations on the herbarium sheet also fit the brief description.

The lectotype of Papillaria nitidiuscula, Watts 4363, was chosen as it is an ample collection with a duplicate. This collection also shows well the different parts of the plant from the short, falcate, densely leaved branches to the long, fine, trailing branches. None of the syntypes nor the lectotype had sporophytes. 771egenus Barbella 203

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Fig. 54. Distribution of Barbella11ite11s (Hook.f. & Wils.) Nog. The genus Barbella 204

For Dixon's P. nitidiuscula it seemed reasonable to assume that the type was Watts 4351. The BM collection is without number and annotated as 'near Ballina, on tree in scrub'. The collections from NSW and Hare annotated as 'Ballina, on tree in pine scrub'. Both have the same date - July 1900. This collection was cited as a syntype by Brotherus in his description of this species.

NOTES

Barbella nitens is readily recognized by the golden colour and the 'rolling' of the leaf which is similar to Papillaria flavolimbata, but Barbella nitens has a more delicate appearance. Some plants have only short, thick, falcate branches and give this species an atypical appearance. This species is more frequently mixed with any of the Papillaria species rather than found in pure colonies. One Barbella specimen collected (HS 28372) in north Queensland had been made into a small, bird's nest.

Only two fertile collections were noted (HS 6062a & Watts 1764) from northern New South Wales. Both had capsules maturing in December.

All Watts' collections from northern New South Wales were determined originally by V.F. Brotherus as Papillaria intricata, the name being subsequently used by Watts for his earlier collections.

It is unfortunate that this species could not be retained in Papillaria with which it often shares the same general appearance, an auriculate leaf base and the basal cells arranged in divergent rows. The costa is weak and often not discernible, but this is also found in other species (e.g. P. intricata). However, the lack of numerous prominent papillae is not typical of Papillaria. When Noguchi (1985) transferred this moss to Barbella he was influenced especially by sporophyte characters such as the peristome as well as the leaf areolation and The genus Barbella 205

papillae. However, he concluded that the basal angles of the leaves were more akin to Papillaria.

On examining the leaf of Barbella pendula ( of which I had only limited collections) I do not believe that they appear to be similar to B. nitens. The former species has obscure cells with plasmolysed cytoplasm while B. nitens has clear cells. Few papillae and a weak short costa are shared by both species. In

my opinion B. nitens is closer to Papillaria than to Barbella.

When additional fertile collections become available an examination of the sporophytes, perhaps by scanning electron microscopy, might more satisfactorily settle the generic position of this taxon.

SPECIMENS STUDIED: Australia 203 (35 cited).

QUEENSLAND: Windsor Tableland, Stone 16011 (MELU), 23 Aug. 1979; Mossman Gorge, Stone 15869 (MELU), 7 Aug. 1979; Mt Lewis, Stone 15798 (MELU), 3 Aug. 1978; Davies Ck, on fallen tree, Stone 12297 (MELU), 31 May 1978; L. Eacham, 16km E of Atherton, 8()()m, poor lowland rain forest, on Ficus stem, HS 16868 (CBG), 28 Feb. 1983; Boonjie S.F., 22km SE of Yungaburra, 600m, rain forest, on Endiandra mue//eri branch, HS 27555 (CBG, L), 3 Mar. 1983; Cooroo logging area, 16km WNW of Innisfail, 100m, rain forest, crown of large tree, HS 29976 (CBG, L), 27 Jun. 1984; Maalan, 11km E of Ravenshoe, 980m, remnant shrubs and treelets between road and paddock, on a shrub, HS 30590 (CBG, NY), 2 Jui. 1984; Rockingham Bay, Mueller (FH ex 8456, MEL 32167), 1855; Blencoe Ck, 48km NW of Cardwell, 740m, Lauraceae­ Syzygium dominated rain forest, on a vine (Freycinetia), HS 36888 (CBG), 17 Jun. 1986; Wallaman Falls Rd, 30km W of Ingham, 500m, disturbed rain forest, trailing from a vine, HS 28493 (CBG, L), 19 Jun. 1984; Mt Spee S.F., 6km W of Paluma, 920m, Lauraceae-Syzygium dominated forest, on shrub twigs, HS 37036 (B, CBG), 18 Jun. 1986; Credition S.F., 16km SW of Finch Hatton, 700m, Eucalyptus grandis dominated woodlands, base of Eucalyptus, HS 37701 (CBG, NY), 1 Jui. 1986; Rockhampton, Dietrich (NY), 1866; 33 k Builan, Stone 22841 ( + Papillaria amblyacis - MELU), 8 Aug. 1984; Blackall Ra., on tree, Whitteron 211 (H, MEL 32248), Apr. 1901; Mudgerabah, ex Herb. Wild s.n. (MEL), 29 Mar. 1889; McPherson Ra., 1130m, on Nothofagus moorei (F. Muell.) Kasser, Helmsing (BM), 28 Dec. 1934. NEW SOUTH WALES: Brindle Ck, Wiangaree S.F., 750m, Cunoniaceae-Lauraceae dominated forest, on fallen tree branch, HS 6062A (CBG), 3 Sep. 1978; Alstonville Rd, Ballina, on sappling, Watts 207 (H, NSW, syntype The genus Barbella 206

Papillaria nitidiuscula), 13 Apr. 1896; Chaelundi Mountain, 37km N of Ebor, 1376m, cool temperate rain forest, branches of big Lauraceae, HS 6741 (CBG, L), 14 Oct. 1978; Wilson R., Mt Boss S.F., 15 miles [24km] N of Wauchope, rain forest, on Bosistoa branches, Constable (NSW M11046), 31 Oct. 1956; The Bulga via Wingham, Watts 10850 (NSW), Apr. 1915; Gloucester R., 28km WSW of Gloucester, 400m, Eucalyptus-Casuarina forest, on tree trunk, HS 6457 (CBG, DUKE, H, HSC, L, MO, NAM, NICH, NY, PRE, RNG, UBC), 10 Oct. 1978; Allyn Forest Park, festooning shrubs, Schofield et al. 80957 (UBC), 14 Jui. 1983; Kingwell, Wyong, on fallen tree, Watts 9539 (NSW), 16 Aug. 1907; Megalong Valley, festooning branches of shrubs, Schofield et al. 79610 (UBC), 19 Jun. 1983; Fitzroy Falls, on fern, W.B. & M.I. Schofield 79043 (UBC), 5 Jun. 1983; Mayfield, Nowra, swamp, on she-oak, Watts 6556 (NSW), 2 Jun 1903; Barretts Ck, Araluen­ Moruya Rd, 10km NW of Moruya, 40m, dry sclerophyll forest, on Note/aea stem, HS 15419 (CBG, H, L), 8 Apr. 1981; Rutherford Ck, 11km SE of Nimmitabel, 850m, cool temperate forest, trailing from shrub branches, HS 16750 (CBG, L), 19 Feb. 1982. VICTORIA: Bemm R., Princes Highway, 8km SSW of Club Terrace, remnant cool temperate forest, on treelet branches, HS 35538 (CBG), 27 Sep. 1985; Dianah Ck, near Mt Cann, on jungle vine (Morinda jasminoides), Youl (MEL 1011166), Feb. 1972. TASMANIA: Denison Gorge, on wood, Weymouth 1900 (HO, MEL), 3 Feb. 1894; Mt Direction, East Tamar, on trees, Weymouth 252 (HO), 8 Feb. 1892. The genus Floribundaria 207

IV. The genus FLORIBUNDARIA Fleisch. Hedwigia 44:301.1905 Type: non designatus.

Plants not glossy. Stems elongate, complanately leaved, laxly or pinnately branched. Branches short, simple, some taxa complanately leaved, obtuse at apex. Stem-leaves gradually tapering to an elongate, linear-lanceolate or flexuose-capillaceous apex from a cordate, ovate, or oblong-ovate base; margin serrulate or crenate excluding the acumen; costa reaching mid-leaf. Branch-leaves similar to those of stem, but smaller. Lamina! cells hyaline or obscure, sublinear to rhomboidal, multi- or unipapillose, walls thin..±. sinuose; inferior cells lax, not papillose, alar cells slightly differentiated.

Dioicous. Inner perigonial leaves widely ovate. Perichaetia with or without paraphyses. Calyptra cucullate or mitriform, lobed at base, with or without hairs. Capsules oblong on short, ..±. smooth, setae often curved. Operculum conic, with a short or long, curved beak. Exostome teeth linear­ lanceolate, densely striolate in lower half, densely papillose in the upper half; segments of endostome as long as the exostome teeth, perforated along keel, papillose; cilia none; basal membrane usually high. Spores round, medium­ sized, 15-25um, papillose.

Chromosome numbers: n = 6, 11 (Japan, Inoue 1965, Inoue & Momii, 1971).

ETYMOLOGY: from the Latin,floribundus = profusely flowering.

No Australian collections with rhizoids, perigonia, perichaetia or sporophytes were seen. ThegenusF/oribundaria 208

KEY to AUSTRALIAN SPECIES of FLORIBUNDARIA

la Laminal cells with 3-8 papillae ...... 2 lb Laminal cells with 1-3 papillae ...... (2) F. PSEUDOFLORIBUNDA

2a Papillae of the laminal cells in a single row, leaf apex clear ...... (1) F. FLORIBUNDA 2b Papillae of the laminal cells in two rows, leaf apex papillose ...... (3) F. WALKERI ThegenusF/oribundaria 209

1 FLORIBUNDARIA FLORIBUNDA (Dozy & Molk.) Fleisch., Hedwigia 44:302.1905.

Leskeaff.oribunda Dozy & Molk., Ann. Sc. Nat. Bot. Ser. 3,2:310.1844. Type: 'Java. Sumatra'. HOLOTYPE: L.

Slender, feathery diffuse plants, pale green. Stems up to 20cm, flexuose, creeping or pendant, apex often filiform. Stem leaves appressed to loosely ascending. Branches short, simple obtuse at apex, complanately leaved.

Branch leaves widely complanately spreading when wet or dry. Leaves linear-lanceolate, commonly 1.5x0.3-0.4mm, but quite variable, apex acuminate, base ovate, cordate. Margin very weakly denticulate, more pronounced at tip. Costa very faint, reaching to (1/3) 1/2 leaf length. Cells sub-linear to hexagonal; cells in acumen up to 20x2um, longer at apex , upper cells up to50x5um, .± sigmoid, in mid-leaf similar. Juxtacostal cells near base to 50x6um. Cells pellucid, outlines easily confused with lines of plasmolysed cytoplasm within very obscure cells. Cells with 3-8 papillae in a regular row over cell, extending nearly to leaf base and acumen. Basal cells hyaline, few corner cells thicker-walled, 20x10um, juxtacostal cells near base generally linear 50x5(10)um, nearer margins similar but gently curved, no alar cells.

Stem leaves similar except margin is entire, acumen is longer and filiform, median cells are slightly wider and cell margins are not so obscure.

Perigonia on branches. Inner perigonial leaves wide-ovate, cymbiform, ea 0.7mm long, entire, ecostate; paraphyses numerous. Perichaetia on branches with perichaetial leaves up to 1.5mm long, subulate-acuminate, papillose, paraphyses longer. ThegenusFloribundaria 210

Calyptra cucullate ± 1.5mm with several hairs. Capsule ovate-oblong, 1.8x0.8mm, setae blackish-brown 2-2.5(4)mm long, often curved,_± rugose at apophysis. Opercula ± 1mm long, conic-rostrate with a long curved beak. Exostome and endostome teeth ea 0.5mm long, yellow in lower half, segments wide, keeled. Spores (12)15-22(25)um.

Chromosome number: n = 11 (Japan, Inoue 1965).

DISTRIBUTION and ECOLOGY: Eastern Australia - north Queensland (Fig. 55). This widespread species is found in Africa, Madagascar, throughout southern and southeast Asia, Japan, Taiwan, New Guinea, the Pacific Islands and in Brazil. This species is rare and nothing is known of its ecological preferences in Australia. However, it can be safely assumed that this moss occurs in the rain forest where it prefers tree trunks or is trailing in diffuse mats from shrub stems and branches.

The few Australian collections available lacked altitudinal data, but it appears that the range would be between 400-1200m.

ETYMOLOGY: from the Latin,jloribundus = profusely flowering.

ILLUSTRATIONS: Bartram 1933:173, 1939:pl. 16, fig. 276; Brotherus

1906:822, 1925:170; Dozy & Molkenboer 1848:162, pl. 53 (Meteorium jloribundum); Fleischer 1905:303, 1908:817; Johnson 1980:86; Norris & Koponen 1985:32; Sehnem 1980:78; Sim 1926:393; Wu & Lin 1985b:17. 77ie genus Floribundaria 211

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TASMANIAV 0 ,oo 200 J(X) J()Olun HOBART Simple conic proiectlOf'I r,5" ,so• ,s:;-

Fig. 55. Distribution of F/oribundaria f/oribtmda (Dozy & Molk.) Fleisch. ThegenusF/oribundaria 212

NOTES

Noguchi (1976) stated that the seta is smooth in this genus while Norris & Koponen (1985:35) mentioned that the seta of Floribundaria floribunda is densely papillose. On examination of a F. floribunda (Koponen 32913) collection from Papua New Guinea the seta appeared weakly papillose­ denticulate with projecting cell tips.

Fleischer (1905:302) and Tan (1984:8) considered F. floribunda to be an extremely polymorphic species. F. walkeri is very similar to this species and the Australian collections are hard to separate. See notes after that species.

SPECIMENS STUDIED: Australia 5.

QUEENSLAND: Mt Lewis, Stone 15740 (MELU) 3 Aug. 1979; Malanda, Watts O494B (H, NSW), 0619 (H, NSW), 29 Jul. 1913; near Millaa Millaa, on shrub, W.B. & M.I. Schofield 80175 (UBC), 30 Jun. 1983; Palmerston N.P., Schofield 80315 (UBC), 1 Jul. 1983. ThegenusF/oribundaria 213

2 FLORIBUNDARIA PSEUDOFLORIBUNDA Fleisch., Musci Fl. Buitenzorg 3:822.1908.

Type: 'Exsiccata: M. Fleischer, Muse. Archip. Ind. et Polynes. No. 435'. HOLOTYPE: FH.

Plants fine and feathery, pale green, slightly glossy, yellowish-green in herbarium collections. Secondary stems filiform, elongate up to 4cm long. Branches simple 1.0-1.Smm long, regularly complanately leaved. Branch and stem leaves all spreading at an angle.

Branch leaves lanceolate, generally 1.6x0.4mm, rarely ovate-oblong. Apex shortly acuminate or narrowly acute, base ovate and cordate, margin denticulate from base to apex. Costa faint, light brown, rarely reaching 1/2 leaf length.

Lamina! cells hyaline, but outline obscure, sub-linear to linear­ rhomboidal with one, rarely two, papillae per lumen, Upper lamina! cells sigmoid 37(50)x3-7um, median cells sigmoid up to 37x4um, juxtacostal cells nearer base hexagonal to rectangular up to 50x5um, 2 or 3 rows of lowermost cells variable, rectangular ea 20x2um, shorter & wider, clearly visible papillae, thin walls. Juxtacostal cells at base rectangular 20x12um to quadrate, sometimes one large yellowish alar cell, elliptic to ovate 25x20um.

Stem leaves average size 1.8x0.6mm, costa fainter than in branch leaves, generally two papillae per cell, but not so prominent, alar cells more prominent.

Perigonia on branches. Perigonial leaves widely ovate, all similar, 0.98x0.32mm, apex acuminate, ecostate, slight denticulation in apex. Cells mostly hexagonal, 90x80um, clear with two faint papillae per lumen. Basal cells rectangular 20xl0um. Cells at apex obscure in vegetative plant leaves. 11ie genus Floribundaria 214

Paraphyses few (1 or 2) up to 12um wide. Perichaetia on branches. Inner perichaetial leaves ovate-oblong with a long, recurved acumen, paraphyses long, numerous,.± wavy, bistratose.

Calyptra cucullate .± 1.5mm with several hairs. Capsules oblong 1.5- 2.0x0.8-1.2mm, brown, becoming black when older, exostome teeth and endostome segments 0.40-0.45mm long. Setae smooth, 1.5-1.8mm long, 0.22- 0.25mm thick. No spores seen.

Chromosome number: no reports.

DISTRIBUTION and ECOLOGY: Eastern Australia - north Queensland (Fig. 56). This species also occurs from India to southeast Asia, Taiwan, New Guinea and Vanuatu. Habit and habitat notes are lacking on Australian collections, but it is reasonable to assume that this moss occurs in the rain forest, on tree and treelet trunks, branches, twigs or on leaves. The altitudinal range is about 700- 840m.

E1YMOLOGY: from the Greek, pseudo = false or resembling, referring to the similarity in appearance to F. floribunda.

ILLUSTRATIONS: Bartram 1933:173, 1939:pl. 16, fig. 277; Noguchi 1976:276;

Norris & Koponen 1985:37; Wu & Lin 1985b:19.

SPECIMENS STUDIED. Australia 5.

QUEENSLAND: Mossman Gorge, Stone 23166 (MELU), 14 May 1985; Watts Q582 (H, NSW), 2 Aug. 1913, 0652 (H, NSW), Aug. 1913; near Ravenshoe, Watts 0624 (H, NSW), 3 Aug. 1913; Malanda, Watts 0655 (H, NSW), 29 Jui. 1913. 17ie genus Floribundaria 215

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TASMANIAV 0 IOO 200 JOO 4Cl»m HOBART Simple conic pn,jeetion 145• ,so• ,s:,·

Fig. 56. Distribution of Floribundaria pseudofloribunda Fleisch. The genus F/oribundaria 216

3 FLORIBUNDARIA WALKERI (Ren. & Card.) Broth., Nat. Pfl. 1(3):822.1906.

Papillaria walkeri Ren. & Card., Bull. Soc. Roy. Bot. Belgique 34(2):70.1896. Type: 'Kurseong (Edentale Plantation) - Darjeeling, Walker, Nov. 1893'. HOLOTYPE: PC.

Plants delicate and slender, yellow-green, older parts..± brownish. Stems creeping, laxly and complanately leaved, up to 4cm long, branches few. Branches simple, laxly and complanately leaved, up to 1.5mm wide, the upper part slender, capillaceous, upper branch leaves appressed.

Branch leaves narrowly lanceolate (0.9)1.0-1.3x0.2-0.4(0.6)mm, papillose, tip narrowly attenuate, widest near ovate base. Margin plane and serrulate with projecting cells, costa weak, extending to mid-leaf. Apical, upper and median lamina! cells..± obscure, elongate-rhomboidal or sub-linear with thin walls and papillae. Upper cells (40)50x2-4(7)um, median cells 37-50x2-4um, juxtacostal cells near base hexagonal 50-75x3-5(12)um, thin walls and papillae more prominent, cells at base shorter, wider and with no papillae. No alar cells.

Stem leaves ovate-lanceolate, to 1.5x0.5mm, tip long acuminate, base cordate.

No sporophytic data available.

Chromosome number: no reports. 17ze genus Floribwzdaria 217

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Fig. 57. Distribution of F/oribwzdaria walkeri (Ren. & Card.) Broth. The genus F/oribundaria 218

DISTRIBUTION and ECOLOGY: Eastern Australia - north Queensland. (Fig. 57). This species has also been reported from South Africa, Nepal, India, Laos, Philippines, Sarawak, Java and Samoa. While no data are available on the Australian habit, habitat and altitudinal range of this species, it is estimated that the altitude of the collections is between 720-850m.

ETYMOLOGY: named after the collector.

ILLUSTRATIONS: Gangulee 1976:1306; Noguchi 1976:274; Wu & Lin 1985a:18.

NOTES

Tan ( 1984 ), in his investigation of Meteoriaceae, reports that the accepted distinctions between Floribundaria floribunda and F. walkeri are less obvious in some collections from distant Pacific islands. I studied the two

Australian collections of F. walkeri he cited and found the stated characters hard to see, especially the biseriate papillae. At the base of the leaf up to 4 papillae per cell were commonly observed. On most Australian F. floribunda specimens the leaf tip is generally filiform while in F. walkeri it is shorter and attenuate. However the interpretation of what constitutes a leaf tip will have a bearing. I accept Tan's determinations but feel that the differences, as far as Australian material is concerned, are hard to observe and differentiate. Final judgement has to be postponed until more Australian collections are available

and a worldwide revision of Floribundaria is undertaken. Noguchi (1976) suggested that this genus is unnatural with further study required on the leaf papillae as well as the calyptra. The genus Floribundaria 219

SPECIMENS STUDIED: Australia 2.

QUEENSLAND: Yungaburra, Watts 0668 (H, NSW), 30 Jul. 1913; track to Major Falls, Ravenshoe, Watts 0579 (H, NSW), Aug. 1913. The genus Meteoriopsis 220

V. The genus METEORIOPSIS Fleisch. ex Broth. Nat. Pfl. 1(3):825.1906.

Lectotype: Meteoriopsis squan-osa (Hook.) Fleisch. ex Broth., Nat. Pfl. 1(3):826.1906; Manuel 1977:587. Stems creeping, subpinnately branched, laxly leaved. Secondary stems. elongate, prostrate or pendant, subpinnately or laxly branched, densely leaved. Branches short or long, often reflexed, densely leaved, obtuse or acute at apex.

Stem leaves squarrose or widely spreading, ovate to ovate-oblong, gradually tapering to a long or short, canaliculate apex, variously cordate and mostly clasping at base; margin erect, serrate to serrulate; costa single, light brown, slender, usually extending to mid-leaf. Laminal cells pellucid or .± obscure, linear-rhomboidal, thin-walled, usually with 1, rarely 2 or 3 papillae, with plasmolized cytoplasm; inferior cells larger, not papillose, alar cells slightly delimited, shorter, ovate, subquadrate or rectangular, not papillose.

Rhizoids smooth reddish with thickened margins.

Branch leaves similar to those of stem but usually broader, more compact, more recurved.

Dioicous. Perichaetia on branches. Inner perichaetial leaves narrowly oblong, long-subulate, paraphyses numerous. Calyptrae mitriform, lobed at base, or cucullate, long-hairy or naked. Capsules erect, ovate to oblong­ cylindrical, with an indistinct apophysis. Setae short, smooth or .± scabrous. Operculum rounded-conical, with an erect, subulate beak. Exostome teeth linear-lanceolate, papillose above, densely striolate below, lamellae high; basal membrane low; segments linear, attenuate, keeled. Spores medium sized, 20- 30um, papillose. The genus Meteoriopsis 221

Chromosome number: n = 11 (Japan, Inoue & Momii 1971).

ETYMOLOGY: from the Greek, meteoron = things in the air, referring to the aerial trailing habit.

KEY to AUSTRALIAN SPECIES of METEORIOPSIS

la Cells clear, unipapillose ...... (1) M. RECLINATA lb Cells_± obscure, 1-3 papillae per cell ...... (2) M. ROBUSTULA The genus Meteoriopsis 222

1 METEORIOPSIS RECLINATA (C. Muell.) Fleisch. ex Broth., Nat. Pfl. 1:826.1906. Pi/otrichum reclinatum C. Muell., Bot. Zeit. 12:572.1854. Type: 'Mont. Neigherrenoses [Nilghiri Mts, Madras, India], Schmid s.n.' LECTOTYPE: JE.

Plants small to medium sized, yellow-green to brown-yellow, slightly glossy. Secondary stems to 15cm long, short sparsely irregularly branching, densely leaved. Leaves squarrose-spreading from an erect base, usually with points neatly and uniformly deflexed.

Branch leaves ovate-lanceolate (1.6)1.9 x 0.7mm, gradually contracted to a long, rarely short, and usually filiform twisted acumen broadest near a somewhat clasping base; squarrose with the leaf tips mostly at right angles to the stem. Margin plane, weakly serrulate becoming serrate in the acumen with large teeth at right angles to margin and pointing downward. Costa light brown, extending to 1/2-2/3 leaf length, rarely beyond, width at base 30um. Cells in leaf apex up to 50x6um, incrassate but quite variable, upper leaf cells 45x4um, median cells linear-rhomboidal (25)35-50x5um. Juxtacostal cells towards base hyaline, sub-linear up to 60x7um, at base cells.± rectangular and lOum wide, juxtacostal cells 25x13um with localized thickening. Cells pellucid with one high central papilla, rarely two. Cells clear at extreme apex and near base. Alar cells elliptical, not prominent, larger, rectangular or subquadrate.

Stem leaves similar except costa is weaker and serrations on leaf margin are slightly more prominent.

Rhizoids up to 12um wide, cells up to 175um long, ends overlapping.

Perigonia on branches, inner perigonial leaves ovate to obovate with an acumen, cymbiform, up to 1.3x0.5mrn. Margin entire, ecostate, no papillae, cells The genus Meteoriopsis 223

narrow elliptic up to 65x4um, porose. Cells shorter in the acumen, slightly shorter and rectangular along the margin. Antheridia elongated 0.45x0.13mm, cells shortly rectangular 25x20um or quadrate, but with considerable variation in length, cells with low papillae. Paraphyses 20nm (1 cell) wide with rectangular cells up to 90nm long, margins and nodes thickened. Apices of paraphyses pointed. Perichaetia on branches. Inner leaves narrowly oblong up to 1. 7mm long, apex subulate, paraphyses numerous exerted well beyond leaves.

Capsules oblong-cylindrical to ea 2.5xl.3mm, brown but becoming black with age. Calyptrae ea 1.5mm long. Setae ea 2mm long, scabrous or nearly smooth. Opercula ea 1.5mm long. Exostome teeth ea 0.7mm long, endostome segments laxly papillose. Spores not known.

Chromosome number: n = 11 (Japan, Inoue & Momii 1971).

DISTRIBUTION and ECOLOGY: Eastern Australia - north Queensland (Fig. 58), where the species is restricted to the Atherton Tablelands. Meteoriopsis reclinata is widespread in southeast Asia extending from India and Sri Lanka to Taiwan, Japan, New Guinea and Vanuatu. The species inhabits rain forest, but has been found to occur in an old Araucaria cunninghamii plantation with advanced regrowth and in Eucalyptus grandis dominated grassland. Meteoriopsis reclinata has been collected on a variety of substrates, with not one substrate predominating. It occurs mainly on tree trunks, upper tree trunks and fallen tree branches, but has also been collected from a log, a leaf, and a shaded boulder. Grows on various tree and treelet species but Eucalyptus grandis is the only one specified. The altitudinal range is 360-1200m.

ETYMOLOGY: from the Latin, reclinis = to lean back, referring to the leaf being bent or turned down. The genus Meteoriopsis 224

ILLUSTRATIONS: Bartram 1939:pl. 17, fig. 286; Dozy & Molkenboer 1864:pl. 201 (Meteorium phaeum auct. non Mitt.); Gangulee 1976:1354; Manuel

1977b:589; Noguchi 1976:336,337,339,340; Norris & Koponen 1985:45; Wu & Lin 1986a:15.

NOTES

Noguchi {1976) recognized 5 varieties for M. reclinata, but recently (pers. comm. 1988) he expressed his doubts about whether these varieties should be recognized taxonomically. Dr Noguchi thought that one Australian collection

(HS 29588) might be M. reclinata var. ancistrodes (Ren. & Card.) Nog. However, having observed a wide variation in leaf shape and size in the Australian species, I am convinced that they also are highly variable. See

comments in M. robustula notes.

Some collections had partly black leaves on the older stems, but rarely on branch leaves.

No fertile Australian collections were noted.

SPECIMENS STUDIED: Australia 36 (10 cited).

QUEENSLAND: Windsor Tableland, 45km NW of Mossman, 1200m, open rain forest, on tree trunk, HS 29588 (CBG), 26 Jun. 1987; Davies Ck Rd, on fallen tree, Stone 12299 (MELU), 31 May 1975; Mt Baldy, 4km SW of Atherton, 1080m, Araucaria cunninghamii plantation with regrowth, on tree stem, HS 2'n97 (CBG), 25 Jun. 1984; Tinaroo L. area, road to Mt Haig, rain forest, on moist shaded boulders, Norris 41583 (NY), 10 May 1974; L. Eacham, 16km E of Atherton, 800m, poor lowland rain forest, on fallen tree branch, HS 16898 (CBG, L), 28 Feb. 1983; Hugh Nelson Ra., 15km S of Atherton, 1090m, Eucalyptus grandis dominated grasslands on a ridge, trunk of large E. grandis, HS 29499 (ALTA, BA, BISH, CBG, COLO, DUKE, FH, H, HSC, NICH, NY, RNG), 25 Jun. 1984; Boonjie S.F., 22km SE of Yungaburra, 600m, rain forest, on fallen tree branch, HS 27573 (CBG, L, NICH), 4 Mar. 1983; Millaa Millaa Falls N.P., 900m, complex notophyll vine forest, Thiers 2256 (NY), 6 Jui. 1984; Bellenden Ker, Bailey 633 (BRI), 1889; Palmerston N.P., rain forest, Thiers 2372 (NY), 8 Jui. 1984. The gmus Meteoriopsis 225

f,0-f 150• ,o•

,s• I 0 I I o,,, I ,r~ I I G'I ~ I \ NT I \ I \ I \ I ',\ I \ I \ I QUEENSLAND \ I I· \ I \ I \ I ------, \\ I '-·-~- .. ~ I 'et, ...... I ~ I z ! BRISBANE C( \ I Q: I ',------.,--... ,.,,,r~)-- I , SOUTH ,-' I ' AUSTRALIA I <., ...', JO' I ~ ., I I NEW SOUTH WALES ,I... -, I '-- () ,, SYDNEY ADELAIDE I , •• ) . _,....,____ I ', ~ I \ - '\ • 00,, B RRA ..J ---- ~ I VICTORIA ~ \ i I - --"I·r. !'----~_,· ~,

,rs 0 ~ 411'S

TASMANIAV 0 100 200 lllll - .. HOl!IART Sonv• - .,,..._ r«rf 14Y ,,o.. 15:, ·

I'll- 58. Distribution of Meteoriopsis r«linata (C. Muell.) Acisch. ex Broth.. The genus Meteoriopsis 226

2 METEORIOPSIS ROBUSTULA (Broth. & Watts) Streim., comb. nov.

Basionym: Floribundaria robustula Broth. & Watts, Proc. Linn. Soc. New South Wales 43:560.1918. Type: 'N. Queensland: Malanda, on tree-trunks, [Watts] n. 618a'. HOLOTYPE: H!; ISOTYPES: BM!, NSW!.

Robust plants, green to pale green. Primary stems creeping, secondary stems to 6cm long, branches irregularly scattered up to 2cm long, apex acute rarely obtuse. Stem and branch leaves obliquely spreading, often slightly twisted, not squarrose.

Branch leaves ovate-oblong 2.1-2.Sx0.5-l.lmm, tapering to an elongated apex, broadest near a somewhat clasping base. Margin plane, weakly serrulate or distantly serrate, becoming serrate in the acumen with medium sized teeth, rarely at right angles to the margin .. Costa extending 1/2 (3/4) leaf length, width at base 25-30um.

Cells in leaf apex 35-50x5um, slightly sigmoid, variable, upper cells of similar size, median cells up to 50x4um. Juxtacostal cells towards base up to 75x7um, basal cells.± rectangular, up to lOum wide. Cells .± obscure with 1-3 low papillae. Stem leaves sparser, 2.8-3. lxl.3mm, long fine apex with teeth at right angles. Upper cells unipapillose, median cells with 2 or 3 papillae which decrease towards base. Leaves on small buds ovate .± 850um long, ecostate, cells obscure except near apex, 2 (3) papillae per cell and serrations more prominent.

Rhizoids up to 15um wide, cells up to 150um long, ends overlapping, rarely prosenchymatous. The genus Meteoriopsis 227

No sporophyte data available.

Chromosome number: no reports.

DISTRIBUTION and ECOLOGY: Eastern Australia - north Queensland (Fig. 59). A rare species restricted to the rain forest on Atherton Tableland where it prefers tree trunks. Often mixed with Meteoriopsis reclinata, Py"hobryum paramattense (C. Muell.) Manuel (Watts Q681c), Papillaria crocea, Muellerobryum whiteleggei (Broth.) Fleisch., Barbella nitens (Watts Q681e.p.), Macromitrium and with lichens Leptogium and Pseudocyphellaria.

The altitudinal range is 800-850m.

ETYMOLOGY: from the Latin, robustus = firm, robust, referring to its robust appearance.

ILLUSTRATIONS: none.

NOTES

This species clearly belongs to Meteoriopsis and not Floribundaria. It is a robust moss having widely spreading leaves with densely leaved branches and the appearance is unmistakably that of Meteoriopsis. The serrate leaf apex and incrassate cells are not characteristic of Floribundaria (Noguchi 1976:270), but the somewhat obscure cells with several papillae per cell are more characteristic of Floribundaria. I have no hesitation, however, in placing this taxon in Meteoriopsis. The genus Meteoriopsis 228

140-E 145" ,so• 70'

t"' -p,.-,\\ .. __ 'i

\ Cooktown I 15' \ I o ' \ Cairns I L I I -o/~ I ,'"1-r 1 I ~ 20' I \ NT I '. I \ I \ I ' I \ I \ I QUEENSLAND \ I \ I \ 25' I \ I \ ------, \ 25• I , .... ___ .... u.1 I --~ .. ' z ...... ,, I ~ BRISBANE ~ \

'------____ ,,, 'Q: .,.,,r--I I' ~-~ I Ju' ,/ ', SOUTH I ~ I :- AUSTRALIA c, I I -> I ~ ', I I NEW SOUTH WALES I ~ ,.... -, ' I "' <:> ,, AOELAtOE f 1 .... ~;, .--....- I ', ~ I ,_,-_ '\ •cA'NBERRA 35' I VICTORIA --~\~ ( I ---~--~----~-,, _,,

WS 0 ~

40'$

TASMANIAV 0 700 200 JOO 400lom ~?BART Simplo tonic prqoct,on

lfO'E 745' -6 '-' 750' 155'

Fta. 59. Distribution of Meteoriopsis robustu/a (Broth. & Watts) Streim. The genus Meteoriopsis 229

In appearance this species strongly resembles M. reclinata and finding reliable constant characters that separate the two species was extremely difficult. The characters of M. robustula that appear to be somewhat constant are the .± obscured cells with lines of plasmolysed cytoplasm, 1-3 papillae per cell and the more rounded rectangular basal cells. In M. reclinata the rectangular cells are more angular with the horizontal sides obliquely angled. Alar cells of the former species tend to be rounded while in M. reclinata they appear to be more rectangular, subquadrate or quadrate, but most of these features tend to intergrade.

I am reasonably confident that M. robustula, with.± obscure cells may be only an extreme form of M. reclinata. One collection of the latter species (Watts Q629, det.? Brotherus) has cells only slightly obscured while most Australian collections have clear cells. Some Asian collections had .± obscure cells between the type of M. robustula and the common Australian M. reclinata, in fact similar to Watts Q629. Thus in Australian collections an intergrading from clear to.± obscure cells occurs.

Final taxonomic judgement on M. robustula will have to wait until more collections, especially fertile material, become available.

SPECIMENS STUDIED: Australia 4.

QUEENSLAND: Barrine L., Atherton, upper tree trunk, W.B. & I.M. Schofield 80057 (UBC), 29 Jun. 1983; Malanda, Watts 0629 (NSW), 29 Jui. 1913; Wopen Ck, Stone 18166 (MELU), 16 Aug. 1979; Palmerston N.P., on branch within spray of falls, W.B. & 1.M. Schofield 80338 (UBC), 2 Feb.1983. The genus Meteorium 230

VI. The Genus METEORIUM Dozy & Molk. Musci Archip. Ind. Ined. 157.1854

Type: non designatus.

Stems creeping, flexuose, densely leaved, terete, densely or laxly branched. Branches often long, flexuose and pendant, densely leaved, turgid, obtuse at apex. Stem leaves erect, imbricated, ovate-oblong to ovate, amplexicaul, not decurrent, generally cucullate at apex, sometimes plicate, the insertion narrow, basal corners rounded and often auriculate, undulate, apex rounded-obtuse with a short or long flexuose acumen; margin incurved; basal corners often undulate, generally entire throughout; costa single, reaching 1/2- 2/3 leaf length. Branch leaves similar to those of stem, but often narrower and usually more concave. Laminal cells hyaline, mostly with one large papilla over lumen, thick-walled; median laminal cells elliptic, rhomboidal, elongate­ rhomboidal, to sub-linear, the walls incrassate at cell-corners, sometimes becoming shorter towards apices and margins, apical cells longer, without papillae. Juxtacostal basal cells longer, linear-rectangular, walls thicker, porose; alar cells not defined, but cells at basal corners elliptic, rectangular, subquadrate, sub-linear, arranged in divergent or arcuate rows, scarcely papillose.

Rhizoids reddish.

Dioicous. Perigonia on stems or branches, with many perigonial leaves; inner perigonial leaves ovate, cymbiform, costa weak, without paraphyses. Perichaetia on branches. Inner perichaetial leaves oblong-lanceolate, shortly acuminate; paraphyses numerous, extending well beyond the leaves. Calyptrae cucullate, long-hairy. Capsules erect, ovate to ovate-oblong, smooth, The genus Meteorium 231

macrostomous. Setae much longer than capsules, densely papillose. Opercula conical with a long, oblique beak. Exostome teeth linear-lanceolate, irregular in outline, densely papillose throughout, obscure, white when dry, lamellae rather high; endostome densely papillose, basal membrane low; segments linear, as long as the exostome teeth, irregular in outline,.± flexuose, narrowly perforated along the keel. Spores spherical, medium-sized, 15-30um, papillose.

Chromosome number: n = 10 (Japan, Inoue 1965).

ETYMOLOGY: from the Greek, meteoron = things in the air, referring to its aerial trailing habit. 11ie genus Meteorium 232

METEORIUM BUCHANANII (Brid.) Broth., Nat. Pfl. 1(3):818.1906.

Isothecium buchananii Brid., Bryol. Univ. 2:363.1827. Type: 1n regione Nepal Indiae, Buchanan'. HOLOTYPE: G. Meteorium bailey Broth. ex Bail., Syn. Queensland FI. Suppl. 2:71.1888, nom. nud. M. miquelianum (C. Muell.) Fleisch., Nat. Pfl. 1(3):818.1906. Papillaria baileyi Broth., Bot. Centralb. 36:86.1888, syn. nov. Type: 'Australia orientalis, Queensland, ubi ad Brisbane River leg. F.M. Bailey'. HOLOTYPE: H!; ISOTYPE: MEL (MEL 32284)! Meteorium baileyi (Broth.) Broth., Nat. Pfl. 1(3):818.1906, syn. nov.

Plants large, rigid, golden to dark green, not glossy, becoming black with age. Stems up to 30cm long and 1.5mm thick; branches short, up to 2.5mm thick. Stem leaves dense, erect and conspicuously appressed when dry but not always so on the branches.

Leaves ovate to oblong-ovate, obtuse or .± rounded obtuse, up to 3.0xl.2mm, with a long erect or.± flexuose or a linear-lanceolate acumen, basal angles rounded, auriculate, variably concave, prominently .3-plicate. Margin crenulate or rarely entire, incurved above, slightly undulate at base, costa 2/3 leaf length. Unipapillose, but often smooth in the acumen, auricles weakly papillose and base generally smooth.

Branch leaves similar but more concave, narrower 2.0-3.lx0.7-1.2mm, rounded-obtuse, with a slender, shorter acumen, or gradually attenuating to a long point, base cordate, often asymetrical. Cells in acumen linear elongate to narrow rhomboidal, 17.0-25.0xS.0-7.Sum, .± incrassate, upper leaf cells rhomboidal 25-45x5-7um, median cells 20-4 lxS-7um, often slightly sigmoid, The genus Meteorium 233

juxtacostal cells near base up to 45um long, rarely obscure. Auricles with divergent cells. Cells in basal corners 17x8um, more rectangular and thicker than surrounding cells, sometimes a golden-brown.

Leaves at base of branches .± rounded, hooded 530x45um, upper cells 12.5x3.0-5.0um, variably shaped, in mid-leaf 37x5um, irregular to sigmoid, leaf tip obtuse with irregular outline. Unipapillose, but ecostate.

Rhizoids clustered, up to 1mm long, 13um wide, edges thickened, cells up to 175um long.

Perichaetia with long paraphyses. Capsules oblong up to l.5xl.0mm; exostome teeth to 0.5mm. Setae up to 7mm long, scabrous except for the basal portion. Spores 15-30um, irregularly big and dissimilar. No fertile Australian collections were found.

Chromosome number: n = 10 [Japan, Inoue 1965 for M. helminthocladulum (Card.) Broth. = M. buchananii var. helminthocladulum (Card) Nog.].

DISTRIBUTION and ECOLOGY: Eastern Australia - Queensland (Fig. 60). A widespread moss ranging from India and Sri Lanka to Japan and Korea, and in the Pacific as far as Fiji.

A very common moss of the rain forest, wet sclerophyll forest, remnant forest left as road verges, moister regrowth areas and collected once from a Eucalyptus savannah. It has been reported on tree trunks reaching up to the tree crown branches; trailing from shrubs or from vines. Often found in large pure colonies on larger tree trunks. Rarely have other bryophytes been found mixed with it. Meteorium buchananii has been collected from the following trees or shrubs: Acacia, Alphitonia, Araucaria, Carallia, Endiandra, Eucalyptus, Helicia and Sloanea. The genus Meteorium 23~

150' I(}'

0 I I I I I I I NT I I /U' I I I QUEENSLAND I I I 2s• I I ------, 25' I I I z I < I ______'- .,.--- ~ 30' ! SOUTH I ,-' I AUSTRALIA C, ',' 30• I , .. , I ~ \ I NEW I SOUTH WALES I ,__ \ ' I ~- o / ADELAIDE I I ·-, I ', c· I \ -'------,\1-'\ • ,'CA°NBERRA 35' I VICTORIA ~ \ : I ---~,',._ ~,,--"'l- -,,.·'

40'S

"1'S

0 IOll 200 JOO 400ltm Simple conic pro,ection

1'0'E ,so• ,s~-

Fig. 60. Distribution of Meteorium buchananii (Brid.) Broth. The genus Meteorium 235

This species ranges from 100-1080m in altitude, but has most commonly been collected between 600-900m.

ETYMOLOGY: named after the Scottish doctor F. Buchanan Hamilton who worked on, and collected plants in India.

ILLUSTRATIONS: Bartram 1939:pl. 16 fig. 271 (Meteorium miquelianum); Cardot 1904:25 (Papillaria helminthocladula); Dozy & Molkenboer 1848:131,pl. 51,52 (Meteorium polytrichum Dozy & Molk.); Gangulee 1976:1294,1296 (M. buchananii fo. polytrichum); Fleischer 1908:774 (M. miquelianum); Johnson 1980:86 (M miquelianum); Noguchi 1948:59 (M. helminthocladulum var. cuspidatum (Okam.) Nog.), 1976:253; Norris & Koponen 1985:32; Sakurai 1911:144 (M. cuspidatum Okam.); Wu & Lin 1986b:13 (M miquelianum ssp. atrovariegatum (Card. & Tuer.) Nog.).

TYPE MATERIAL

The specimen on which the name M baileyi (MEL 32284) was based, but never validly described, was extremely plicate with the leaf tip not rounded but gradually narrowed into a large apex. This would place it close to and just a variation of M. miquelianum which is now in synonymy with M. buchananii.

The holotype (annotated M. Menzel Dec. 1986) gives only Queensland as the locality, while the isotype has Brisbane River included. Both collections are in Brotherus' handwriting with the isotype also having 'corn Brotherus' on it.

NOTES

This species has quite variable leaves. Gangulee (1976) mentioned that M. miquelianum and M. buchananii show great variation and it is hardly possible The genus Meteorium 236

to distinguish between the two species. Fleischer (1908) distinguished M. buchananii from M miquelianum as having:

'leaves not plicate, a narrower base, leaf tips rounded with suddenly pointed tips'.

Noguchi (1976) mentions that M. buchananii is not plicate while Gangulee (1976) claims it is plicate. The Australian collections studied showed all degrees of variation between these characters with those of M miquelianum predominating. But in one collection (HS 29372) the tip was rounded and then suddenly narrowed and both leaf apex types were present with a significant variation between the two types. Plication was not always consistent in the Australian collections. Some specimens (HS 37411, Watts Q574, Q648) showed very little, if any, plication and the leaf tip gradually narrowed to a subula. Many collections were 4-plicate. Generally the growing stems and branches are not plicate and when the leaf tip is piliferous then this species can be mistaken for Aerobryopsis wallichii.

Thus it is evident that the characters used by Fleischer to separate the two species are as variable in Australian specimens as Gangulee (1976) found them to be in Indian ones. This is further evidence that Meteorium buchananii is a very variable species.

SPECIMENS STUDIED: Australia 130 (15 cited).

QUEENSLAND: Big Tableland, 26km S of Cooktown, 550m, edge of Eucalyptus dominated grasslands and rain forest, on tree trunk, HS 30893 (CBG, NY), 4 Jui. 1984; Windsor Tableland, 39km NW of Mossman, 1080m, rain forest, trailing from treelet branches, HS 2%80 (CBG, H, NICH, NY), 26 Jun. 1984; Main Coast Ra., 11km NW of Mount Molloy, 940m, clearing with Acacia, on Acacia branch, HS 30445 (CBG), 30 Jun. 1984; Cairns, on trees, Wild (BRI), 1890; Bellenden Ker Ra., Mulgrave R., SW of Gordonvale, 120m, rain forest, on a vine, Kantak & Churchill 981 (CBG, NY), 26 Apr. 1984; Mt Baldy, 4km SW of Atherton, 1080m, old Araucaria cunninghamii plantation with regrowth, on Araucaria trunk, HS 31324 (CBG), 25 Jun. 1984; Barron S.F., 11km SSW of Atherton, 1050m, Eucalyptus savannah, on Eucalyptus trunk, HS 27271 The genus Meteorium 237

(CBG, L, NICH), 2 Mar. 1983; Boonjie S.F., 22km SE of Yungaburra, 600m, logged rain forest, on Endiandra muelleri branch, HS 27536 (CBG, GRO, H, L, NFLD, NICH, NY), 3 Mar. 1983; Cooroo logging area, 16km WNW of Innisfail, 100m, rain forest, in crown of large tree, HS 29967 (CBG, NY), 27 Jun. 1984; Mill Stream, Kennedy Highway, 38km S of Atherton, disturbed highland rain forest with Eucalyptus, on tree limb, Kantak & Churchill 795 (NY), 2 Apr. 1984; Johnstone R., Shirley (BRI), no date; Koombooloomba Dam Rd, 23km SE of Ravenshoe, 780m, rain forest, on shrub, HS 28880 (CBG, L), 23 Jun. 1984; Kirrama Rd, 23km WNW of Cardwell, 730m, disturbed rain forest, on stone, HS 31246 (CBG), 8 Jui. 1984; Wallaman Falls, on shrub, W.B. & 1.M. Schofield 80354 (UBC), 3 Jui. 1983; Conway S.F. 16km ENE of Proserpine, 160m, lowland rain forest, on Carallia brachiata branch, HS 37411 (CBG), 28 Jun. 1986; Lamington N.P., Ramsay 208 (UNSW), 7 Nov. 1981. The genus Weymouthia 238

VII. The genus WEYMOUTHIA Broth. Nat. Pfl. 1(3):811.1906.

Type: non designatus.

Plants medium sized to large. Primary stems creeping, secondary stems numerous, pendulous, often with tapering to slender tips. Branches short, distant and almost at right angles to stem. Stem leaves appressed or laxly spreading when both dry and moist, branch leaves similar. Leaves oblong­ spathulate to sub-orbicular, apex obtuse to rounded or cucullate, base broad or subcordate; margin plane entire, apex often slightly crenulate. Costa none or double and faint. Lamina! cells smooth, long sigmoid or fusciform, superior cells shorter, inferior cells shorter and wider, alar cells sub-quadrate, oval or rounded-oblong, very incrassate, darker or yellowish, auricles distinct.

Rhizoids reddish-brown with thickened margin.

Dioicous. Perichaetium sheathing, inner perichaetial leaves bluntly pointed. Calyptra cucullate, smooth. Capsules suberect, oval, setae short or long, slender, reddish. Operculum conical or conico-rostrate. Peristome double, inserted slightly below rim. Teeth lanceolate-subulate, hyaline bordered, transversely striolate with zig zag median line, papillose above. Ventral lamellae rather high. Basal membrane of endostome about 1/3 length of teeth; processes slightly papillose, lanceolate, perforated on the median line. Cilia two or absent. Spores medium sized, 20-30um, papillose.

Chromosome number: no reports.

ETYMOLOGY: named after W.A. Weymouth, a Tasmanian bryologist in the 19th century. The genus Weymouthia 239

KEY to AUSTRALIAN SPECIES of WEYMOUTH/A (after Scott & Stone 1976)

la Leaves sub-orbicular, very concave, wrinkled when dry. Plants firm ...... (1) W. COCHLEARIFOLIA lb Leaves oblong-spathulate, predominantly U channelled, not...... wrinkled. Plants soft ...... (2) W. MOLLIS The genus Weymouthia 240

1 WEYMOUTHIA COCHLEARIFOLIA (Schwaegr.) Dix., New Zealand Inst. Bull. 3:255.1927.

Hypnum cochlearifolium Schwaegr., Spee. Muse. Suppl. 1(2):221.88.1816. Type: 1n Australasia lectum communicavit cl. La Billardiere '. HOLOTYPE: G. Isothecium cochlearifolium (Schwaegr.) Mitt., Hooker's J. Bot. Kew Gard. Misc. 8:264.1856. Stereodon cochlearifolia (Schwaegr.) Mitt., J. Linn. Soc., Bot. 4:88.1860. Coelidium cochlearifolium (Schwaegr.) Rchdt, Reise Oesterr. Freg. Novara Bot. 1(3):191.1870. Porotrichum cochlearifolium (Schwaegr.) Mitt., Trans. & Proc. Roy. Soc. Victoria 19:84.1882. Lembophyllum cochlearifolium (Schwaegr.) Lindb., Ind. Bryol. 718.1897. Meteorium molle Wils. var. majus Wils., FI. Tasm. 2:203.1859. Pilotrichella mollis (Hedw.) Jaeg. var. major Jaeg., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1875-76:260.1877 (Ad. 2:164). Neckera billardieri Hpe, Linnaea 30:637.1860, syn. nov. Type: 'Apollo Bay, F. Mueller 1857'. HOLOTYPE: BM!; ISOTYPES: BM!, MEL (32257)! Pilotrichella billardieri (Hpe) Jaeg., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1875-76:259.1877 (Ad. 2:163). Meteorium billardieri (Hpe) Mitt., Trans. & Proc. Roy. Soc. Victoria 19:82.1882, syn. nov. Weymouthia billardieri (Hpe) Broth., Nat. Pfl. 1(3):812.1906. W. cochlearifolia (Schwaegr.) Dix. var. bi/Lardieri (Hpe) Dix., New Zealand Inst. Bull. 3:255.1927, syn. nov.

Plants generally robust, rarely slender, loosely tufted, creeping or pendulous, yellow-brown, yellow-green or pale green, glossy. Collections often age to a golden-reddish colour. Stems flexuose and robust, rarely filiform. The genus Weymouthia 241

Branches very variable in length and type of branching, leaves tightly imbricated, tumid, usually wrinkled or terete. Branches often pinnate, rigid and freely branching with long flexuose branches or may also be short. Branch tips with spreading leaves, leaf size decreasing up the branches giving a flagellate appearance.

Branch leaves sub-erect when dry or wet, sub-orbicular, cochleariform, very concave up to 1.3.x2.0mm, apex cucullate with a hooded appearance, base wide. Margin entire, but generally weakly denticulate or crenulate at apex, costa lacking or very faint, cells smooth.

Cells at leaf tip up to 35x5um, but variable, incrassate, median cells linear-sigmoid with pointed ends (25)30-80x4-6um. Cell size decreasing and width increasing nearer base, incrassate and porose. Alar groups small but conspicuous generally with a reddish or yellow tinge or clear. Cells quadrate or rectangular up to 25x12um, with granular content and very incrassate. Marginal cells smaller and narrower.

Stem leaves up to 2.lxl.8mm and often wider than long. Denticulation at tip very weak or more. Cells similar to branch leaves.

Rhizoids 13um wide, cells up to 175um long, ends sometimes overlapping.

Perigonia not seen. Perichaetia elongated, bud like, scattered on branches. Inner perichaetial leaves ovate up to 900x450um, tip shortly acuminate, margin entire, ecostate, no papillae. Upper and median cells up to 50x3um, incrassate, .± sigmoid, basal cells variable rectangular (± 25x7um), porose, reddish at base. Archegonia .± 16 per bud, with irregularly rectangular cells, reddish-brown, up to 450x40um, neck up to 20um wide. Paraphyses up to The genus Weymouthia 242

900um long, 13um (1 cell) wide, cells rectangular, thick walled, up to 63um long. Sporophytes at bases of branches. At base of capsule inner perichaetial leaves similar but lanceolate-ovate up to 2.7xl.0mm, tip acuminate, upper and median cells up to 63x4um, more incrassate. Paraphyses similar to those surrounding archegonia except 15um wide, cells up to lOOum long.

Capsules brown oblong to oval, rarely asymetrical up to 1. 7x0.8mm, setae very long up to 15.6mm long with a slight anti-clockwise twist. Operculum acutely or obtusely conical, about half length of capsule. Peristome yellow­ brown, teeth up to 400um long, transversely striolate up to 240um from base. Margin of striolate section with semi-circular indentation corresponding with each zig zag. Processes keeled. Cilia 2, nodulose, shorter than the processes. Spores 20-25um.

Chromosome number: no reports.

DISTRIBUTION and ECOLOGY: Southeast Australia - New South Wales, Victoria and Tasmania (Fig. 61). It also occurs in New Zealand and Chile.

Habit, habitat and altitudinal data are lacking for most collections, but it appears that Weymouthia cochlearifolia is found in temperate and Nothofagus forests where the preferred substratum is tree trunks (Nothofagus and

Lagarstrobus). This species has also been reported on logs and wood. The altitudinal range appears to be from sea-level to 460m in Tasmania, in Victoria it has been reported from the lowlands to 1150m in eastern Gippsland and in New South Wales at 60m above sea level.

E1YMOLOGY: from the Latin, cochlea = spoon shaped and foliatus = leaves, referring to the rounded cucullate leaves. The genus Weymouthia 243

'IWf 1',tJ·· ; ft ,o•

Cl 'lD

"'1P ~ ~ Cookfown I ,.,. I o ' -. I .c:,. '\ Cairns I I o ,61,,/ I I ,-z.~ l G" I \ I \ .,, NT I \ I \ I \, I \ I I \ I QUEENSLAND \ I \ I 25• \ I \ I \ ------, \ \ 25• ' '---,,~ I ~, I ',, I z ~ BRISBANE I < \ ,------' ____,, ---'Q: ,1r-: - 30" I ~ ~ SOUTH I AUSTRALIA :- 1 ~ , __~ , JO' I ~ ', I ' •• ' NEW SOUTH WALES Q-/ I I r--,I '~' ~-,__., I '-- _-..___ ADELAIDE I ''-, I ,J--.... '\ I VICTORIA I -~r~,,~."" ,, -Sv ~,

411'S

411"S

0 100 200 300 4(XJl,.m Simple conic projection r«rf ,so• ,,,,.

Fig. 61. Distribution of Weymouthia cochlearifolia (Schwaegr.) Dix. The genus Weymouthia 244

ILLUSTRATIONS: Allison & Child 1971:108 & pl.25; Brotherus 1906:866, 1925:206 (Lembophyllum cochlearifolium); Dixon 1927:373; Schwaegrichen 1816 fig. 88.

TYPE MATERIAL

The type of Weymouthia cochlearifolia var. billardieri (Neck.era billardieri - BM) was examined and proved to be within the acceptable range for variation of Weymouthia cochlearifolia. Sainsbury (1955a) found that the characters Dixon (1927) used to delineate this variety were unsatisfactory: longer, more flexuose stems; shorter setae; and almost globose capsules. These characters showed considerable intergrading throughout a single plant. Dixon (1927) showed very clearly the weaknesses between the species and its variety and it is surprising that he decided to keep them separate. Australian collections studied here also exhibited this same degree of variation. Perpetuation of this confusion could perhaps be explained by the rarity of fertile material and the insufficient number of collections available from which to observe all the variations.

The type for Weymouthia cochlearifolia could not be located. From

Wilson's (1859) description of the new variety, Meteorium molle var. majus, it was not clear whether the collections cited were for the new variety or for the species generally. Therefore as this variety is in synonymy, no type was designated.

NOTES

Specimens of Weymouthia cochlearifolia have often been misdetermined as Lembophyllum divulsum causing considerable problems. The main differences are compared below. The genus Weymouthia 245

Weymouthia cochlearifolia Lembophyllum divulsum

1) Cells long 40-80x6um Cells short 20x6um with tapering sigmoid rhomboidal & variable ends 2) Costa absent, faint Costa commonly wide at or double base, forked, generally 1/2 of leaf length 3) Leaves very obtuse, Leaves narrowed at wide insertion forming a waist 4) Porose at cell ends Porose only at base, rarely at tip 5) Alar cells - short, Alar cells - rounded square or oblong 6) Leaf tip not pouch Leaf tip at times like pouch-like 7) Various sized leaves Even sized leaves on branch, often throughout flagelliform 8) On tree trunks On rocks, soil, only sometimes on trees

In Scott & Stone (1976) descriptions for these two genera leaf tips are described as different: Weymouthia - 'entire or at most slightly crenulate'; Lembophyllum - 'crenulate-denticulate'. This was found to be variable between the two genera and not a reliable character.

Weymouthia cochlearifolia has also been confused with Acrocladium chlamydophyllum but this latter species has pointed shoots, flatter leaves and big inflated empty hyaline cells in the auricles. Both Lembophyllum and Acrocladium usually occur at the base of trees or on the ground while Weymouthia is found generally on tree trunks. The genus Weymouthia 246

W. cochlearifolia can develop filiform tips to branches where the leaves are very small and widely spaced (Mueller-MEL 32264 & Willis-MEL 1034367). Bigger leaves on these branches have a similar appearance to W. mollis leaves.

Perigonia and perichaetia were extremely hard to find because the rounded branch leaves completely cover these organs. It appears that sporophytes mature between August and October.

Wijk et al. (1969) list W. cochlearifolia as occurrmg m Asia (not confirmed by Noguchi 1976), and for the Pacific - Oceania (not substantiated by Miller et al. 1976). Brotherus (1906, 1925) does not list W. mollis for Hawaii, but Sainsbury (1955a) stated that the genus occurs in the Sandwich Islands (Hawaii). No doubt he based this on the works of Brotherus, where W. desmoclada (C. Muell.) Broth., which is now considered conspecific with the Hawaiian endemic Pilotrichella maaiensis (Sull.) Jaeg., is reported for Hawaii.

SPECIMENS STUDIED: Australia 85 (23 cited), New Zealand 57.

NEW SOUTH WALES: Higgins Ck, 19km NE of Batemans Bay, 60m, remnant cool temperate rain forest along creek, on trunk of small rotting shrub, HS 5759 (CBG, H, L, MO, NICH, UBC), 9 Aug. 1978. VICTORIA: Rooty Break Track, Goonmirk Rocks Rd, Scenic Reserve, 1150m, dense mountain forest, epiphyte, Thies FN 1471 M (MEL), 28 Mar. 1986; Croajingolong N.P., Camp Ck, S of Cicada Trail, Beauglehole 31993 (MEL), 22 Nov. 1%9; Mt Ellery, Merrah (BM, NY), 1887; Mt Drummer, Melville & Wakefield 28320 (MEL, NSW), 12 Jan. 1953; Ettas Glen, Black Spur, near Healesville, Watts V436 (NSW), 21 Dec. 1886; Cement Ck, Warburton, Hansen (MEL 38462), Jan. 1947; Powelltown, Willis (MEL 1034367), Dec. 1929; Skenes Ck, Apollo Bay, Mott (MEL 32258), 29 Oct. 1891; West Portland Reserve, abundant-spreading, Beauglehole 73113 (MEL), 16 Oct. 1956; Boundary Ck, Yeodene, SE of Colac, Beauglehole 1%3 (MEL), Nov. 1949; Turtons Pass, 13km E of Beech Forest, 430m, temperate rain forest, on tree trunk, HS 2428 (CBG, H, L, MO, NICH), 24 Aug. 1975; Tarwin, Mueller 122 (BM, NY), no date; Sealers Cove, Mueller 112 (BM, NY), no date. TASMANIA: Ulverstone, Lodder (MEL 32279), Mar. 1886; Denison Gorge, Scottsdale railway line, Weymouth 1898 (NY), 8 Feb. 1894; Hellyer R., Ramsay 1860 (UNSW), 8 Dec. 1981; Goulds Country, Robinson (as Bastow 385 - MEL, NSW), Aug. 1886; Lyell Highway, shaded log, wet sclerophyll forest, Norris 33430 (COLO, NY), 31 Jan. 1974; Philip The genus Weymouthia 247

Is., Macquarie Harbour, dense forest, Anon. 1315 (NY), 1 Nov. 1946; Gordon R., forest, Milligan 785 (NY), no date; Mt Field N.P., Nothofagus forest, Vitt 29129 (ALTA), 6 Dec. 1981; Sandy Bay, Weymouth 2363 (NY), 22 Oct.1910. The genus Weymouthia 248

2 WEYMOUTHIA MOLLIS (Hedw.) Broth., Nat. Pfl., 1(3):812.1906.

Leskea mollis Hedw., Spee. Muse. 234.1801. Type: 'Habitat in nova Seelandia'. HOLOTYPE: G. Neckera mollis (Hedw.) C. Muell., Syn. 2:131.1850. Meteorium mo/le Wils., Fl. Nov. Zel. 2:100.1854. Isothecium mo/le (Hedw.) Mitt., Kew J. Bot. 8:263.1856. Stereodon mollis (Hedw.) Mitt., J. Linn. Soc., Bot. 4:88.1860. Pilotrichella mollis (Hedw.) Jaeg., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1875-76:260.1877 (Ad. 2:164). P. weymouthii C. Muell., Hedwigia 41:130.1902. Type: 'Tasmania, Nelson, Gally back Mt., Nov. 1890: W. A. Weymouth in Hb. 0. Burchard 1891'.

Slender plants with soft texture, frequently in extensive masses, green or pale yellow-brown. Secondary stems slender forming filiform pendulous masses up to 20cm long. Branches simple, flexuose, short up to 2cm long, bluntly cuspidate. Leaves loosely imbricated. Branch leaves oblong-spathulate, cucullate, up to 1.5x0.9mm, apex rounded or obtuse, base contracted to insertion on branch, often sub-cordate at base. Margin plane, entire, generally slightly denticulate at apex, costa absent, or rarely very faint and double. Cells smooth. Apical cells up to 35(63)x4um, sinuose, incrassate, median cells up to 60x5um, sinuose with thinner walls, basal juxtacostal cells up to 67x5um, often incrassate. Alar cells conspicuous, brownish-red, with granular content, with cells both lOum quadrate and rectangular 20x10um, incrassate, often forming auricles.

Stem leaves longer, lanceolate, narrower, generally not cucullate 2.0x0.8mm. Alar region more pronounced.

Rhizoids long (± 3mm), 15-17um wide, cells up to 150um long. The genus Weymouthia 249

Perigonia bud like, elongated, scattered on branches. Inner perigonial leaves ovate up to 900x400um, tip shortly acuminate, ecostate, very few papillae near base, margin entire. Upper cells elongate up to 37x4um, incrassate, median cells elongate up to 50x4um, less incrassate, cells towards base becoming rectangular; margin with inconspicuous border of narrow rectangular cells. Antheridia .± 7 per bud, club shaped up to 410x140um, reddish-brown with quadrate and shortly rectangular cells. Paraphyses clear up to 500x20um, margin thickened, rectangular cells up to lOOum long. Perichaetia not noted. Archegonia at base of capsules, club shaped, up to 400x30um, neck not markedly differentiated, reddish-brown, with quadrate and short rectangular cells. Inner perichaetial leaves at base of capsules, very long sheathing, linear­ lanceolate, curved, up to 3.2x0.5mm, tip shortly acuminate, margin entire, ecostate, no alar cells, very few papillae on basal cells. Upper cells linear, up to 65x2um, incrassate, median cells linear up to 65x4um, incrassate, .± sigmoid, near base variably rectangular (± 30x20um). Paraphyses 12um (1 cell) wide, cells rectangular up to 75um long. Sporophytes lateral, placed near the bases of the branches.

Calyptra hairy (?). Capsules 1.0-1.5x0.6-1.0mm, ovoid, brown. Seta 4.0- 5.Smm long, slender and curved. Operculum conico-rostrate, with a long curved beak. Annulus weakly developed. Peristome yellow-brown, processes keeled. No cilia. Spores 20-30um.

Chromosome number: no reports.

DISTRIBUTION and ECOLOGY: Southeastern Australia - New South Wales, Lord Howe Island, Victoria and Tasmania (Fig. 62). Also occurring in New Zealand, Juan Fernandez Is. and Chile. The genus Weymouthia 250

Weymouthia mollis prefers moister, semi-shaded habits and has commonly been collected in wet sclerophyll, cool temperate and Nothofagus (myrtle or beech) forests. The most common substrate is tree or shrub branches where it often forms dense, pendulous masses. Rarely has it been noted on cliff faces, rocks, ground, rotting logs or on tree roots. This species has been collected from Acradenia, Atherosperma, Bedfordia, Nothofagus, Pomaderris and Telopea.

The altitudinal range of Weymouthia mollis in Tasmania is from sea-level up to 800m, in Victoria between 120-lO00m but in New South Wales it is unknown.

ETYMOLOGY: from the Latin mollis = soft, pliant, referring to the long delicate trailing stems and branches.

ILLUSTRATIONS: Allison & Child 1971:pl. 24; Brotherus 1906:812, 1925:159.

TYPE MATERIAL

The type of Pilotrichella weymouthii was not amongst the collections received on loan.

NOTES

This is quite a distinctive moss once recognized. However it may be mistaken for Camptochaete gracilis (Hook.f. & Wils.) Par., but this species has much shorter (up to 30um) median cells. Rarely should it be mistaken for Acrocladium chlamydophyllum which has much wider leaves, pointed shoots and a more shiny green colour. The genus Weymouthia 251

.,. ·t 'J·1 ,,;()" t }\',~ '! G\ >l I"',...... Cooktown 15" I 0 I ..::. :

I O I I I ,r;_ ' I {\ G' I \ NT I \ I '\°' 20• I ', I \ I \ I QUEENSLAND \ I \ I \ 25" I \ I \ 25• ------,I \, _____ \>)

I' --13,,, I z ! BRISBANE < \

'------______,,,,,,,~- ...... ,Q:: ..,.~J---I ' 30" 1 ,/ ', SOUTH I ~ I : AUSTRALIA I <-' __ :-, I I ' ~ __ • l NEW SOUTH WALES (?-/ I ~' ~-- , .... -, ' ? __ .. I '-- <:> > ADELAIDE I I •-, I '\.' ,_--- I \J--- '\ •5=A'NBERRA 35• ' I VICTORIA --,"'~ \ : I <;I-_,--~ / ---~-- ...~',

40"S

,o-s

0 100 200 JOO "'°""' Simple conic p,ojection

140-E 150' 15:,'

Fig. 62. Distribution of Weymouthia mollis (Hedw.) Broth. The genus Weymouthia 252

Perigonia were hard to find, and in fact no perichaetia were observed. The leaf shape and arrangement would not hide these organs as in Weymouthia

cochlearifolia. Sporophytes appear to mature in November.

SPECIMENS STUDIED: Australia 101 (22 cited), New Zealand 25.

NEW SOUTH WALES: Blue Mountains, Anon., (BM, MEL 32272), no date. LORD HOWE ISLAND: Summit of Mt Gower, Watts (NSW 183270), Aug. 1911. VICTORIA: Errinundra R. West Branch, 20km S of Bendoc, 960m, Atherospenna moschatum and tree fern dominated forest, HS 36584 (B, CBG, CHR, COLO, DUKE, FH, HO, HSC, NY, PRE, RNG), 9 Apr. 1986; Martin Ck, Bonang Bighway, 34km NNE of Orbost, 270m, wet sclerophyll forest, on Telopea stem, HS 35479 (B, CBG, CHR, COLO, DUKE, MICH, NY, PRE, RNG), 26 Sep. 1985; Club Terrace­ Combienbar Rd, 6km N of Club Terrace, 120m, regrowth with Pittosporum, Acacia and Bedfordia, shrub branch, HS 35571 (B, CBG, NY), 27 Sep. 1985; Ettas Glen, Healesville, Watts V302 (NSW), 21 Dec. 1906; Upper Yarra Park, Powelltown, on trees along gully, Beauglehole 15390 (MEL), 23 Jan. 1966; Fern Tree Gully, Sassafras Ck, 40km ESE of Melbourne, 330m, Cyathea dominated wet sclerophyll forest, on tree trunk, Curnow 196 (CBG), 21 May 1985; Head of Bunyip R., festooning sassafras branches, Willis (MEL 1034368), 2 Sep. 1935; Korumburra, Martin (MEL 32276), 1891; Cape Otway N.P., Mails Rest, Beauglehole 16317 (MEL), 19 Nov. 1966; Tarwin, Mueller (BM, MEL 32271), 1855. TASMANIA: Strzelecki Peaks, Furneaux Group (Flinders I.) 750m, Gabriel (MEL 1040657), 23 Jan. 1893; Circular Head, Weymouth 1047 (HO), no date; Hellyers Gorge, between Burnie and Waratah, wet Nothofagus forest-riparian woodland, Weber & McVean B-33285 (COLO), 23 Feb. 1968; Poatina, 800m, Eucalyptus forest, cliff face, Norris 32424 (HO, NY), 13 Jan. 1974; Maria I., Perrin 666 (MEL), Oct. 1886; Wanderer R., 6km upstream from mouth, riverine forest, Buchanan 6235 (HO, NY, TNS), 8 Mar. 1985; Newmans Ck, Tasman Peninsula, on trees, Weymouth 658 (HO), 16 Mar. 1891; Earl Point, facing Port Davey, myrtle forest on seashore, Davies 1310 (MEL), 14 Mar. 1954; Hobart, McRobies Gully, on sassafras, Weymouth 647 (HO), Jan. 1888. Excluded species 253

EXCLUDED SPECIES

BARBELLA PERPINNATA (Broth.) Broth., Nat. Pfl. 1(3):825.1906. Pilotrichella perpinnata Broth., Oefvers. Forh. Finska Vetensk. -Soc. 42:108.1900. The type of this taxa could not be located, but two collections (MEL 32300, MEL 1050657 and a duplicate of latter at NSW - 183263) were studied. These collections were from Brown Falls near Killarney in southern Queensland. The substratum was a rock near a waterfall.

It was concluded (no sporophytes available) that this taxon is best placed in Brachytheciaceae, being close to Brachythecium salebrosum (Web. & Mohr) B.S.G. This relationship was supported by the original annotation Eurhynchium remotifolium (Grev.) Jaeg. (on MEL 1050657) which was later changed to Hypnum. It is most unusual, if not impossible, to find Australian Barbella species on rocks, but this is quite a common substratum for Brachythecium.

CHRYSOCLADIUM PHAEUM (Mitt.) Fleisch., Musci Fl. Buitenzorg 3:830.1908.

An extensive literature search yielded no Australian references nor were any Australian collections noted during this study. Dr Margadant (in litt. Feb. 1988) was of the opinion that the reference for Australia is in error (Wijk et al. 1969:484). Excluded species 254

PILOTRICHELLA CONFERTA Ren. & Card., Bull. Soc. Roy. Bot. Belgique 38(1):24.1900. This is an incorrect reference (Wijk et al. 1967:67) to this tax.on in Australia (in litt. Dr Margadant, Feb. 1988).

PILOTRICHELLA RECURVULA C. Muell., Hedwigia 41:129.1902.

This species is Calyptothecium recurvulum (Broth.) Broth. m the Pterobryaceae. The type, from Helsinki, was studied. It had previously been annotated as C. recurvulum.

When C. Mueller published this species he referred to Trachyloma recurvulum in Hb Brothen. In fact Brotherus (1893:53) published T. recurvulum giving credit to C. Mueller. Bailey (1890:98) listed this as Pterob,yon

(Trachyloma) recurvulum C. Muell. which is a nomen nudum. Apparently C. Mueller was not aware that Brotherus had validated this tax.on and thus published it as a new species in 1902. The location data given by the authors were:-

Bailey: Bellenden Ker Brotherus: Queensland, Bellenden Ker Range, ubi anno 1889 legit F.M. Bailey. Mueller: Queensland, Kev Range, Ballandea 1889. Conclusion 255

CONCLUSION

The Meteoriaceae in Australia now consists of 7 genera with 17 species (Table 8), a reduction of 2 genera and 9 species. Amongst the Papillaria studied a further 3 species may be involved. However, no taxonomic judgements could be made because of inadequate collections. With more systematic collecting in the Queensland rain forest, taxa other than Papillaria will no doubt be discovered. Conclusion 256

Previously recognized taxa Present taxa and status

Aerobryopsis wal/ichii (Brid.) Fleisch. Barbella cubensis (Mitt.) Broth. B. nitens (Hook.f. & Wils.) Nog. B. perpinnata (Broth.) Broth. **Brachythecium ?salebrosum (Web. & Mohr) B.S.G. Chrysoc/adium phaeum (Mitt.) Fleisch. Not Australian Floribundaria floribunda (Dozy & Molle.) Fleisch. F. pseudofloribunda Fleisch. F. robustu/a Broth. & Watts *Meteoriopsis robustula (Broth. & Watts) Streim. F. walkeri (Ren. & Card.) Broth. Meteoriopsis reclinata (C. Muell.) Fleisch. ex Broth. Meteorium baileyi (Broth.) Broth. M. buchananii M. buchananii (Brid.) Broth. M. compressum Mitt. Barbella cubensis Papillaria amblyacis (C. Muell.) Jaeg. P. crocea (Hpe) Jaeg. P. eavesiana (Hpe) Jaeg. P. amblyacis P.flavolimbata (C. Muell. & Hpe) Jaeg. P. flexicaulis (Wils.) J aeg. P. fulva (Mitt.) J aeg. Barbella nitens Papil/aria intricata (Mitt.) C. Muell. & Broth. P. leuconeura (C. Muell.) Jaeg. P. reginae (Hpe) Jaeg. P. crocea P. squamata (Hpe) Hpe P. amblyacis *P. zeloflexicaulis Streim. Pi/otrichel/a conferta Ren. & Card. Not Australian P. dimorpha (C. Muell.) Jaeg. Barbella nitens P. recurvula C. Muell. **Calyptothecium recurvulum (Broth.) Broth. Weymouthia cochlearifolia (Schwaegr.) Dix. W. c. var. billardieri (Hpe) Dix. W. cochlearifolia W. mollis (Hedw.) Broth.

* Endemic ** Not Meteoriaceae

Table 8. Meteoriaceae taxa reported for Australia prior to this revision and the present status. Addendum 257

ADDENDUM

A recent Australian National Botanic Gardens field trip to Mitchell River National Park, near Bairnsdale in Gippsland led by F.E. Davies found a bird's nest on the ground. The nest was rimmed with Parmotrema reticulatum (Tayl.) Choisy with the lower sides and base being of fine mosses - Isopterygium, Papillaria flavolimbata and P. flexicaulis. This is the first report of the latter species from Victoria. The apparent absence of this species from Victoria seemed unusual as this moss had previously been reported from southern New South Wales and northern Tasmania. It must be rare as recent trips to Gippsland had failed to locate it. The nest will be kept for display. Field work will be planned to this locality to search for this species. Acknowledgements 258

ACKNOWLEDGEMENTS

My deep gratitude is extended to Dr H.P. Ramsay for all the encouragement and assistance she has given me in Australian bryology. Also I am grateful to Dr G. Argent who initially flamed my interest in mosses while we were both roaming the forests in Papua New Guinea.

As literature was often limited in Canberra, and to a lesser extent in Australia, the willing help provided in locating old and obscure works by I. Gleadhill, librarian at the Australian National Botanic Gardens, was greatly appreciated. Often I had to seek many works overseas and here Drs W.R. Buck (NY), T. Koponen (H), R. Hoogland (PC) and B. van Zanten (GRO) were always ready to assist promptly.

D. Verdon cheerfully assisted with the Latin translations and shared his knowledge of and helped with the freezing microtome techniques. Further assistance with microtome sections was kindly provided by N. Plovanic of the Australian National University, Canberra. M. Jayachandran of the University of New South Wales, Sydney, and N.D. Mackie of the Australian National University also assisted with the preparation of the samples for the Scanning Electron Microscope (SEM) and in the use of the instrument.

Interesting discussions, helpful advice and assistance with cladistics was provided by J. Armstrong and Dr M.D. Crisp of the Australian National Botanic Gardens. Greatly appreciated was the advice and discussions with Dr J.A. Elix of the Australian National University about selection of types, taxonomic points and numerous problems as they arose. Dr A. Noguchi of the Kumamoto University was always ready to share his vast knowledge of Meteoriaceae with me which was greatly appreciated. Acknowledgements 259

I wish to thank the curators of the herbaria mentioned on pages 5 and 6 for their assistance in promptly providing loans of collections.

Initial typing was performed by my daughter, Arlene and by J.A. Curnow who had to put up with my bad handwriting. J.A Curnow also put the work onto disk and kept it up to date through numerous revisions and assisted in various ways. J.R. Croft of the Australian National Botanic Gardens assisted with computer-formatting of the text. Mapping was skilfully carried out by K. Cowan of the Australian National University; he also advised on various effective ways to present data. References 260

REFERENCES

ALLISON, K.W. & J. CHILD 1971. The mosses of New Zealand. Univ. Otago Press, Dunedin.

ANDERSON, L.E. 1954. Hoyer's solution as a rapid permanent mounting medium for bryophytes. Bryologist 57:242-244.

AONGSTROEM, J. 1876. Primae lineae muscorum cognoscendorum, qui ad Caldas Brasiliae sunt collecti. Oefvers. Forh. Kongl. Svenska Vetensk. Akad. 33:3-50.

BAILEY, F.M. 1883. A synopsis of the Queensland flora; containing both the phanerogamous and cryptogamous plants. Gov. Print., Brisbane.

BAILEY, F.M. 1888. A synopsis of the Queensland flora, suppl. 2. Gov. Print., Brisbane.

BAILEY, F.M. 1890. A synopsis of the Queensland flora, suppl. 3. Gov. Print., Brisbane.

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INDEX to BOTANICAL NAMES

The names of sections and tax.a above genera are omitted as are generic names that repeatedly occur in discussions. Accepted tax.a are in Roman type; treated tax.a are in bold type; synonyms are in italics; pages for detailed descriptions of tax.a are in bold print; pages with illustrations are in italics.

Acrocladium chlamydophyllum ...... 89, 245, 250 Aerobryidium ...... 14, 20, 21, 189 Aerobryopsis ...... 14, 20, 76, 93, 94, 95, 183-184 A. bauerae ...... 77 A. longissima ...... 78, 185, 187, 189 A. pemitens ...... 187 A. philippinensis ...... 187 A. sigmatophyl/a ...... 78, 185 A. subdivergens ...... 16 A. wallichii...... 3, 5, 18, 76, 78, 185-190, 236, 256 Aerobryum ...... 14, 20, 21, 76, 95 Ancistrodes ...... 14, 19, 20 Aptychopsis uii ...... 195 Araucaria cunninghamii ...... 223 Barbella ...... 14, 20, 78, 93, 191-192 B. asperifolia ...... 16 B. comes ...... 95 B. cubensis ...... 3, 5, 15, 17, 18, 21, 80, 94, 95, 178, 189, 192, 193-198, 256 B. detennesii ...... 195 B. enervis ...... 15, 79, 80, 81, 193 B. flagellifera ...... 16 B. macroblasta ...... 197 B. nitens ...... 1, 3, 5, 8, 9, 10, 15, 18, 21, 33, 36, 38, 70, 78, 80, 81, 86, 94,121,142,164,169,178,192,197, 199-206, 227,256 B. pendula ...... 80, 205 B. perpinnata ...... 1, 3, 15, 81, 253, 256 Index to botanical names 272

B. stevensii ...... 16 B. strongylensis ...... 19, 78 B. trichophoroides ...... 193 Barbellopsis sinensis ...... 195 Brachythecium ...... 1 B. salebrosum ...... 253, 256 Calyptothecium recurvulum ...... 254, 256 Camptochaete gracilis ...... 250 Campylium polyganum ...... 190 Casuarina cunninghamiana ...... 123 Chrysocladium ...... 14, 19, 20 C. phaeum ...... 1, 3, 253, 256 C. retrorsum ...... 16 Citrus limon...... 123 Coelidium cochlearifolium ...... 89, 240 Cryphaea novae-valesiae ...... 32, 146 C. papillarioides ...... 147 Cryphaeophilum ...... 14, 19, 20 Dendrobium striolatum ...... 139 Dolichomitriopsis ...... 95 Dryopteris albovillosa ...... 107 Encalypta vulgaris ...... 152 Eriocladium bauerae ...... 77 Eucalyptus grandis ...... 223 Eurhynchium remotifolium ...... 253 Ficus dammaropsis ...... 69 Floribundaria ...... 14, 20, 82, 93, 94, 95, 207-208, 227 F. aurea ssp. nipponica ...... 16 F. floribunda ...... 3, 5, 16, 18, 82, 83,208, 209-212, 214,218,256 F. ltipponica ...... 16 F. pseudofloribunda ...... 3, 5, 18, 82, 83, 208, 213-215, 256 F. robustula ...... 1, 3, 15, 82, 83, 256 F. torquata ...... 82 F. walkeri...... 3, 5, 18, 83, 208, 212, 216-219, 256 Hypnum coch/earifolium ...... 88, 89, 240 H. cupressiforme ...... 152 H. nigrescens ...... 26, 96 Index to botanical names 273

H. wal/ichii ...... 185 Isopterygium ...... 257 Isotheciopsis ...... 95 Isothecium ...... 89 I. buchananii ...... 86, 232 I. coch/earifolium ...... 240 I. molle ...... 248 Lantana camara ...... 123 Lastreopsis silvestris ...... 107 Lembophyllum ...... 95 L. cochlearifolium ...... 89, 240, 244 L. divulsum ...... 89, 244, 245 Leskea flexicaulis ...... 146 L. floribunda ...... 82, 209 L. mollis ...... 88, 89, 248 Lindigia ...... 14, 20, 21 Meteoridium ...... 14, 19, 20 Meteoriopsis ...... 14, 20, 84, 93, 94, 95, 220-221 M. conanensis ...... 84 M. novoguineensis ...... 84 M. reclinata ...... 1, 3, 5, 16, 18, 84, 85, 221, 222-225, 227, 229, 256 M. reclinata var. ancistrodes ...... 224 M. robustula ...... 1, 5, 221, 224, 226-229, 256 M. squarrosa ...... 84, 220 M. undulata ...... 16, 84 Meteorium ...... 14, 20, 35, 43, 44, 81, 82, 86, 92, 93, 94, 95, 230-231 M. amblyacis ...... 100 M. baileyi ...... 3, 15, 87, 88, 89,232,235,256 M. bil/ardieri ...... 89, 240, M. buchananii ...... 3, 5, 15, 16, 18, 44, 86, 87, 88, 232-237, 256 M. buchananii var. helminthocladulum ...... 233 M. buchananii fo. polytrichum ...... 15, 235 M. cerinum ...... 31, 36, 131, 141 M. cinclidotus ...... 86, 193 M. compressum ...... 3, 15, 44, 86, 88, 193, 195, 256 M. cubense ...... 193 M. cuspidatum ...... 235 Index to botanical names 274

M. cuspidiferum ...... 29, 31, 35, 36, 37, 114 M. cuspidiferum var. cerinum ...... 35, 131 M. dic/adioides ...... 79, 193 M. dimorphum ...... 199 M. eavesianum ...... 100 M. enerve ...... 193 M. filifonne ...... 86 M. filipendulum ...... 32, 36, 146, 152, 154 M. filum ...... 146 M. flexicaule ...... 32, 36, 146, 154 M. floribundum ...... 210 M. fulvum ...... 33, 81, 199 M. helminthoc/adulum ...... 16, 233 M. helminthocladulum var. cuspidatum ...... 235 M. helminthoc/adum ...... 16 M. intricatum ...... 158 M. kennadecense ...... 114 M. limbatum ...... 31, 131 M. longissimum ...... 187 M. longissimum var. tenue ...... 187 M. molle ...... 89, 248 M. mo//e var. majus ...... 89, 240,244 M. mique/ianum ...... 15, 87, 232, 235, 236 M. miquelianum var. atrovariegatwn ...... 235 M. nitens ...... 80, 199, 202 M. phaeum ...... 224 M. polytrichum ...... 235 M. signiatophyllum ...... 185 M. subpolytrichum ...... 16 M. trichophoroides ...... 193 M uellerobryum whiteleggei ...... 227 Neckera ...... 25, 26, 29, 96 N. amb/yacis ...... 33, 100 N. billardieri ...... 89, 240, 244 N. cerina ...... 31, 131 N. crocea ...... 114 N. cuspidifera ...... 30 Index to botanical names 275

N. dimorpha ...... 81, 199 N. eavesiana ...... 35, 100 N. flavolimbata ...... 28, 31, 131, 141 N. fuscescens ...... 26 N. kennadecensis ...... 29, 30, 37, 114 N. leuconeura ...... 165 N. luteola ...... 31, 131 N. mollis ...... 89, 248 N. nigrescens ...... 29, 114 N. reginae ...... 35, 114 N. scottiae ...... 32, 146, 154 N. squamata ...... 34, 100 N. trichophoroides ...... 78, 193 Orthotrichum cupulatum ...... 152 Papillaria ...... 14, 20, 94, 96-99 P. acrocea ...... 115 P. amblyacis ...... 1, 3, 5, 16, 18, 28, 30, 33, 34, 35, 37, 38, 39, 41, 44, 47, 48,48,49, 50, 52, 53, 54, 56, 58, 59, 60, 60, 61, 62, 6~ 64,65,66,68,69, 73, 74,98,99, 100-113, lOL 103, JO~ 107,111,146,154,169,171,178,182,256 P. baileyi ...... 44, 87, 232 P. cerina ...... 31, 131 P. chrysoclada ...... 40 P. compressum ...... 193 P. crocea ...... 3, 5, 16, 18, 27, 28, 29, 30, 31, 33, 35, 36, 37, 38, 39, 41, 44, 45, 46, 47, 48, 49, 53, 54, 54, 55, 56, 57, 58, 59, 60, 61,62,6~ 64,65,66, 68,69, 70, 72, 73, 74, 98,99, 110, 114-130, 116, 117, 118, 120, 142, 151, 155, 171, 178, 180,182,227,256 P. cuspidifera ...... 27, 30, 37, 114, 125 P. cuspidifera var. cerina ...... 31, 131 P. eavesiana ...... 3, 15, 35, 37, 38, 39, 100, 109, 256 P. eavesii ...... 35, 100 P. filicaule ...... 38, 146 P. filipendula ...... 28, 30, 32, 33, 35, 36, 37, 38, 146 Index to botanical names 276

P. flavolimbata ...... 1, 3, 5, 16, 18, 21, 28, 30, 31, 35, 36, 37, 38, 44, 45, 47, 48, 48, 49, 50, 51, 52, 54, 55, 56, 57, 57, 58, 59, 60, 61, 62,63,64,64,65,66,68,69, 70, 72, 73, 74,99, 131-145, 133,135,136,138,155,204,256,257 P. flexicaulis ...... 1, 3, 5, 16, 18, 27, 28, 30, 32, 33, 36, 37, 38, 39, 41, 44, 47,48, 49, 50, 51, 52, 53, 54,54, 55, 56, 57, 57, 58, 59, 60,61,63,64,65,66,68,69, 70, 72, 73, 74,98,99, 121, 125,126, 127,146-157,149,150, 173,178,180,256,257 P. fulva ...... 3, 15, 33, 37, 38, 199, 256 P. fuscescens ...... 16, 26, 40, 164 P. helictophylla ...... 163 P. helminthocladu/a ...... 235 P. intricata ...... 1, 5, 17, 18, 36, 42, 44, 46, 47, 50, 52, 53, 54, 55, 56, 57, 58, 59, 61, 62, 63, 68, 70, 73, 74, 99, 158, 158-164, 159, 204,256 P. kennadecensis ...... 30, 36, 37, 114 P. leuconeura ...... 1, 5, 17, 18, 39, 41, 44, 47, 48, 50, 53, 56, 57, 57, 58, 61, 62, 63, 68, 70, 73, 98, 99, 110, 111, 111, 127, 165, 165- 172, 167,182,256 P. nigrescens ...... 26, 29, 96 P. nitens ...... 15, 33, 38, 43, 80, 81, 199 P. nitidiuscu/a ...... 28, 37, 80, 199, 202, 204 P. reginae ...... 3, 15, 35, 37, 38, 114, 125, 256 P. scottiae ...... 32, 35, 146 P. semitorta ...... 40, 141, 164 P. squamata ...... 3, 15, 34, 37, 38, 100, 109, 256 P. subaongstroemiana ...... 40 P. ventricosa ...... 26 P. wa/keri ...... 216 P. zeloflexicaulis ...... 1, 5, 17, 18, 41, 44, 47, 48, 49, 51, 52, 53, 55, 56, 58, 59, 60, 61, 62, 63, 64, 64, 68, 70, 73, 74, 98, 99, 173, 173- 181, 175, 176, 256 Parmotrema reticulatum ...... 257 Pilotrichella ...... 14, 20, 21, 35, 78, 81, 89 P. amblyacis ...... 100 P. bi//ardieri ...... 240 P. cerina ...... 131 Index to botanical names 277

P. conferta ...... 1, 3, 15, 254, 256 P. dimorpha ...... 3, 15,199,256 P. eavesiana ...... 100 P. filipendu/a ...... 146 P. flavolimbata ...... 131, 142 P. fulva ...... 199 P. kennadecensis ...... 114 P. mauiensis ...... 246 P. mollis ...... 89, 248 P. mollis var. major...... 240 P. perpinnata ...... 81, 253 P. recurvula ...... 3, 15, 254, 256 P. reginae ...... 114 P. scottiae ...... 146 P. trichophoroides ...... 79, 193 P. weymouthii ...... 248, 250 Pilotrichum ...... 92 P. croceu,n ...... 28, 29, 30, 114 P. nigrescens ...... 28, 29, 114 P. pendulinun1 ...... 146 P. reclinatum ...... 84, 222 Porotrichum cochlearifolium ...... 89, 240 Pseudobarbella ...... 14, 20, 21 P. attenuata ...... 16 P. kiushiuensis ...... 16 P. levieri ...... 16 Psidium guajava ...... 123 Pterigynandrnm illecebrnm ...... 26 Pterobryon recurvulum ...... 254 Pyrrhobryum paramattense ...... 227 Pyrrosia rupestris ...... 139 Racomitrium lanuginosum ...... 190 Racopilum cuspidigerum ...... 152 Scheuchzeria ...... 26 Squamidium ...... 14, 19, 20, 21 Stereodon ...... 89 S. cochlearifolia ...... 240 Index to botanical names 278

S. mollis ...... 89, 248 Trachycarpus homschuchii ...... 114 Trachyloma recurvulum ...... 254 Trachypus ...... 31, 32 T. cerinus ...... 131 T. flexicaulis ...... 146 T. homschuchii ...... 29, 30 Tricholepis ...... 26 Weymouthia ...... 14, 20, 88, 93, 94, 114, 238-239 W. billardieri ...... 240 W. cochlearifolia ...... 3, 5, 18, 21, 89, 90, 95,239, 240-247, 252,256 W. cochlearifolia var. billardieri ...... 3, 89, 240, 244, 256 W. desmoc/ada ...... 246 W. mollis ...... 3, 5, 18, 90, 239, 246, 248-252, 256 Zelometeorium ...... 14, 19, 20, 21, 84