Habitat, distribution and the phytogeographical affinities of mosses in the Wet Tropics bioregion, north–east Queensland, Australia.

Andi  Cairns

A total of 56 bryophyte taxa associated with biological soil crusts was collected from 282 sites in semi-arid and arid eastern Australia. The 41 moss taxa and 15 liverwort taxa found were dominated by the families Pottiaceae and Ricciaceae respectively. Bryophytes were present at 224 of the 282 sites, and their distribution varied widely according to different landscapes. The mean number of taxa per site ranged from 8.9 on plains with red earths to 2.1 on active floodplains. The most common bryophytes found in the survey area were Bryum pachytheca, Didymodon torquatus, Gigaspermum repens, Goniomitrium enerve, Desmatodon convolutus, Stonea oleaginosa, Crossidium davidai, Riccia limbata and Riccia lamellosa. In this paper, the distribution of bryophytes in relation to landscape types, and some strategies for surviving in arid environments and their roles in biodiversity and rangeland condition assessment are discussed. Their distribution is compared with other published and unpublishe...

Abstract Bryophytes (mosses, liverworts, hornworts) are an abundant and conspicuous component of the World Heritage Gondwana Rainforests of Australia which comprise several discontinuous areas of subtropical forests and woodland along the Great Escarpment of north-eastern New South Wales and south-eastern Queensland. The Gondwana Rainforests are considered to be of exceptionally high conservation value, with more than 200 rare or threatened plant and animal species but, surprisingly, no mention is made of bryophytes in the statement of outstanding universal values. Recent studies in Werrikimbe and Willi Willi National Parks using bryophytes as fine-scale indicators of rainforest condition, have identified a number of unusual and interesting bryophytes including the moss Rosulabryum epiphyticum (Bryaceae), Leptodontium viticulosoides, (Pottiaceae) and several endemic species of Macromitrium (Orthotrichaceae) The moss Fissidens thorsbornei (synonym: Nanobryum thorsbornei) (Fissidentac...

The aim of this study was to determine the factors that affect bryophyte richness and species composition in wet eucalypt forest, including old growth and forest disturbed by wildfire or silvicultural practice. Approximately one third of the total bryophyte flora for Tasmania was recorded in old growth mixed forest, with more liverwort than moss species found. Bryophyte species composition was significantly different between groups of sites of forest from the northwest, central and southern areas of the state. Mean annual temperature, altitude, rainfall of the driest month and aspect were most significant in predicting variation in bryophyte species composition. The use of vascular plants as surrogates for the conservation of bryophyte species was examined. Vascular plant and fern species richness were significant but poor predictors of bryophyte species richness. A minimum set of 31 sites reserved all vascular species and a large percentage (82.9%) of bryophyte species at least onc...

Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland 371 Habitat, distribution and the phytogeographical affinities of mosses in the Wet Tropics bioregion, north–east Queensland, Australia. Helen P. Ramsay1 and Andi Cairns2 1 National Herbarium of New South Wales, Royal Botanic Gardens, Sydney NSW 2000, AUSTRALIA. 2School of Tropical Biology, James Cook University, Townsville, Queensland 4811, AUSTRALIA. Abstract: A checklist of the mosses (Bryophyta) of the Wet Tropics bioregion, north-east Queensland is presented. Included is an update on the taxonomy of species, listing a total of 408 taxa. The habitat and distribution patterns of species within the area and in Australia, together with information on the phytogeographical affinities of these taxa in related areas beyond Australia, are discussed. Cunninghamia (2004) 8(3): 371–408 Dedication The authors present this work as a tribute to the memory of the late Ilma Stone (1913–2001) and Heinar Streimann (1938–2001), whose work in the area formed the basis for these studies. The work began in the 1980s, between 1984 and 1998 with Ilma Stone, whose taxonomic studies and data from collections made in the area over many years were immeasurable. Heinar Streimann assisted later in the 1990s, with various taxonomic contributions and data from many collections. Without their assistance and knowledge, the work would not have been written. Their deaths in January 2001 and August 2001 respectively were a serious and tragic blow to Australian bryology. Introduction Mosses and liverworts, the major groups of bryophytes, are a significant component of the biodiversity in the Australian wet tropics, in north-east Queensland, occurring in all ecosystems as colonisers of soil, rocks, fallen logs, and as epiphytes and epiphylls. While much attention has been paid to the vascular plants, vegetation types and ecology of tropical north-east Queensland (Tracey & Webb 1975, Kershaw 1978, Tracey 1982), R.M. Schuster (in Keto & Scott 1986) emphasised the importance of bryophytes in the region and criticised the paucity of studies. Many mosses from north-east Queensland were described and published at the end of the 19th and beginning of the 20th century (Bailey 1893, 1913; Watts & Whitelegge 1902, 1906; Brotherus & Watts 1918; Dixon 1938, 1942; Bartram 1952). Ramsay et al. (1987) estimated the moss flora in the ‘wet tropical region of north Australia’ (approximately equivalent to the area under discussion) at about 325 species, representing one quarter of the total species for Australia. There were about 114 Australian endemics present, of which 50–60 were thought to be endemic to the region (Ramsay et al. 1987). The areas of greatest species richness were in the rainforests on the high mountain peaks, and on the Atherton Tableland. Windolf (1987) estimated there were approximately 173 liverwort species in the region, with 20% of these being endemic (Hicks 1986). Recent estimates put the liverwort figure at over 300 taxa (D. Meagher pers. comm.). Bryophytes in rainforests For bryophytes, rainforests provide niches largely absent in other communities (Pócs 1982, Richards 1984, Gradstein 1992) including soil, earth banks, rocks, fallen trees and living roots on the forest floor, and further upwards, the bases, buttress roots and trunks of rainforest trees. Habitats in the canopy include lianes, twigs and branches for epiphytic species, while leaf surfaces provide habitats for epiphyllous species. Bryophytes on rotting logs maintain a moist regime that enhances decomposition and recycling through the activities of fungi and other microorganisms. They play an important role in ecological succession by protecting soil from erosion and providing moist beds for seeds to germinate. In all rainforest environments, bryophytes provide microhabitats and refugia for invertebrates and microorganisms, and sites for egg-laying and nurseries for insect larvae. Growth forms such as mats, cushions or pendents provide surfaces important for harvesting water from cloud and storing minute quantities of nutrients from exudates and droppings of insect larvae that are released slowly as leachates over time. Bryophytes control and reduce runoff by gradually releasing moisture during dry periods, maintaining humidity in the forest. They also contribute to humus accumulation, all important to the maintenance of the rainforest as an ecosystem. 372 Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland History of rainforests in north-east Queensland The history of rainforest bryoflora is closely associated with the history of Australian rainforests, and in north-east Queensland there is evidence of a long ancestry (Webb et al. 1986, Hill 1994, Webb & Tracey 1994, Martin 1998). Original Gondwanan-derived primitive angiosperm species survive in the rainforests of tropical Queensland. In addition, some of the oldest living forms of our present-day dry-adapted sclerophyllous vegetation and their related ecosystems also survive there. Barlow and Hyland (1988) summarised the important paleogeographic and paleoclimatic events that have led to ‘an intricate pattern of humid forest communities which show significant differences in composition, age, endemism, relictuality and richness’. Martin (1998) provided additional evidence, based on palynological studies, of the Tertiary climate in Australia and stated that the precipitation in northeast Australia remained above critical levels for rainforest throughout the Tertiary and most of the Quaternary, enabling the region to be a refuge for many rainforest taxa. Australian rainforests reached their maximum development in the mid Tertiary, thereafter, increasing aridity resulted in reduction of rainforest areas and their retreat to areas of higher rainfall (Frakes 1999). Rainforests in Australia In Australia, ‘rainforest’ is a generic term and includes evergreen forests along the eastern coast (including Tasmania) and seasonally deciduous forests of the north (Lynch & Neldner 2000). Descriptors of rainforests, such as ‘tropical’, ‘sub-tropical’, ‘monsoonal’ and ‘temperate’ that relate to climate (latitude), and ‘montane’ and ‘sub-montane’ that apply to altitude (Beadle & Costin 1952), have application in other regions of the world but do not adequately describe the range of rainforest types found in Australia. Webb (1959) recognised twenty rainforest structural types and classified Australian rainforests on the basis of structure, based on leaf size, seasonality of leaf fall, presence of other life-forms etc., which he later correlated with environmental factors (climate, soil nutrient status) (Webb 1968). Type descriptions of ‘vine forests’ and ‘vine thickets’ (depending on canopy height) are based on leaf sizes (microphyll, notophyll and mesophyll), deciduousness, and structural complexity. The floristic composition of each forest type varies from one locality to another as a result of past rainforest expansions and contractions due to palaeoclimatic events (Martin 1994). Typically, boundaries between structural types are not clear, although distinctions between rainforest and sclerophyll (eucalyptus) communities are generally abrupt, controlled by fire (Adam 1994). The Wet Tropics bioregion The Wet Tropics bioregion (Satler & Williams 1999) is situated along the coast of north-east Queensland, extending south from near Cooktown (15° 35' 46"S, 145°15’20"E) to just north of Townsville (19°23’51"S, 146°29' 28"E), a distance of over 500 km along the Australian coast (Goosem et al. 1999). It lies to the east of the Great Dividing Range and consists of a coastal plain dissected by ridges of the Eastern Escarpment and associated coastal ranges. Along its western flank, it borders the Einasleigh Uplands bioregion, and at its southern extremity, the Townsville Plains province of the Northern Brigalow Belt (Sattler & Williams 1999). To the north, the Wet Tropics links to the remote Cape York Peninsula bioregion. The Wet Tropics bioregion covers approximately 1% of Queensland, with an area of 1 849 725 ha (Goosem et al. 1999). A map of the region is shown in Figure 1. The Wet Tropics is dominated by forested mountains and ranges, with areas of high plateau. Inland from Daintree (lat.16°S) lies the Mt Windsor Tableland (800–1360 m), and extending westwards from the Bellenden Ker Range, the Atherton Tableland (700–1200 m). Transects (see Fig.1, Fig.2 A–C) across Mt Windsor Tableland to the coast, across the Atherton Tableland and the Bellenden Ker Range to the coast, and through Cardwell Range to Hinchinbrook Island, and the south-north profile (Fig.2 D) running through the major peaks and rivers, illustrate the diverse topography of the region (Figs 2 A–D).The Wet Tropics bioregion includes the highest mountains in Queensland — Mt Bartle Frere (1622 m) and Mt Bellenden Ker (1582 m) on the Bellenden Ker Range; Thornton Peak (1374 m), Mt Lewis (1224 m), and Mt Finnigan (1148 m) (Figs. 1 and mountainous Hinchinbrook Island, with Mt Bowen (1142 m) and Mt Diamantina (955 m), is also part of this assemblage. The Cooktown–Ingham massif containing these high peaks covers an area of approximately 360 × 80 km (Webb & Tracey 1981). Physiography The coastal massif and associated ranges form a more or less continuous belt along the eastern Queensland coast, and are igneous in origin. Rocky outcrops are common on mountains, and soils of the massif are primarily granitic. The underlying geology comprises mainly marine Silurian, Devonian and Carboniferous sediments of the Hodgkinson Formation (Arnold & Fawckner 1980). Barron River Metamorphics are exposed in valleys around Cairns, and south to Tully, and sandstones predominate near Cooktown (Wilmott & Stephenson 1989). The scoria cones, maars, obvious basalt flows and fertile basaltic soils of the Atherton Tableland represent volcanism dating from the late Tertiary and Quaternary and overlay a mainly metamorphic base. In some areas, basalt flows followed river systems and reached the coastal plain (Stephenson et al. 1980), although coastal lowland soils are mainly of alluvial and colluvial deposits, largely accumulated during the late Quaternary (Nott et al. 2001). Repeated coastal transgressions and retreat during the Quaternary led to the deposition of coastal and estuarine sediments in some areas, with the attainment of the present sea level approximately 6000 years ago. Large rivers — the Daintree, Bloomfield, Barron, Russell, Mulgrave, North and South Johnston, Tully and Herbert drain the region into the Coral Sea, with higher western areas draining south into the Burdekin River, and north to the Mitchell. Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland 373 Mt Finnigan Fig. 2B. Transect across latitude 17 15' 45"S from Great Dividing Range to the east coast (70 km) (Division of National Mapping Topographic Map 1:100000 Series R631, Edition 2-AAS: Sheet 7963 Atherton and Sheet 8063 Bartle Frere). Fig. 2C. Transect across latitude 18°21’S from Gorge Range to Hinchinbrook Island (75 km) (Division of National Mapping Topographic Map 1:100 000 Series R631, Edition 2-AAS: Sheet 8061 Kirrama and Sheet 8161 Cardwell.) Fig. 2A. Transect across latitude 16°15' S from Mount Windsor Tableland to east coast (55 km) (Division of National Mapping Topographic Map 1:100000 Series R631, Edition 1-AAS: Sheet 7865 South Palmer River and Sheet 7965 Mossman.) Fig. 2D. South-north profile through the Wet Tropics bioregion, showing major peaks and rivers. (Note the x axis is approximate.) Mt Finnigan Fig. 1. The Wet Tropics bioregion, north-east Queensland. Dotted line represents the 1250 mm rainfall isohyet; shaded areas represent extant rainforest. 374 Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland The Wet Tropics bioregion is subdivided into nine provinces (Fig. 3), reflecting differences within the landscape, generally associated with geology, geomorphology and climate. Goosem et al. (1999) provide descriptions of provinces (geology, landform, soils and vegetation) and associated regional ecosystems. Much of the bioregion incorporates the Wet Tropics of Queensland World Heritage Area (WTQWHA) (Satler & Williams 1999), recognised by the international community for the richness and diversity of its flora and fauna. Bloomfield Daintree Cairns Climate The climate generally results in two seasons, a separate wet season (December–April), with summer monsoons and occasional tropical cyclones, and an almost dry season (May– November) with occasional showers. Mean annual temperatures vary with topography, from 30°C in the tropical lowland to less than 10°C in montane areas (Webb 1968). Annual rainfall over the region is highly variable, strongly influenced by local topography, and declines towards the north and the south (Adam 1994). There are three locations on and around the high peaks where the rainfall can exceed an average of 3750 mm per annum: • • • Thornton Peak, Mt Finnigan, Cape Tribulation (north of the Daintree River), and Mt Lewis (north west of Cairns); Mt Bartle Frere, Bellenden Ker and Babinda between the Mulgrave and Russell Rivers; lowland areas between the Johnston and Tully Rivers. Mt Bellenden Ker Centre Peak is the wettest meteorological station in Australia; annual rainfall averages over 8000 mm, peaking at 12 461 mm in 2000 (Bureau of Meteorology 2000). West of the Babinda–Tully area, coastal cloud and rainfall influence extends much further inland, but drops to less than 1300 mm per annum west of the Atherton Tableland (Adam 1994). The approximate position of the 1250 mm rainfall isohyet is indicated on Figure 1. Tropical cyclones are a signficant feature of the region’s climate and these are recognised as being a major factor in shaping the structural and floristic differentiation of the vegetation, particularly in the coastal lowlands (Adam 1994). Vegetation The vegetation of the humid tropics (lat. 15–19°S, long. 145– 146°30’E) (approximating the Wet Tropics bioregion) was described by Tracey (1982), based on Webb (1959, 1968, 1978) and Tracey and Webb (1975) for rainforests, and Specht (1970) for other vegetation communities. Vegetation descriptions are currently under review (Stanton & Stanton 2000, 2001a, 2001b). Thirteen rainforest structural types (Webb 1968) are recognised within the Wet Tropics bioregion (Tracey 1982). The Wet Tropics encompasses the most diverse and the largest contiguous area of tropical rainforest in Australia. Innisfail Tully Cardwell Hinchinbrook Island Fig. 3. Provinces of the Wet Tropics bioregion (redrawn from Satler and Williams 1999) showing approximate boundaries. Numbers correspond to the list of provinces in Appendix 1. Key: 1 = Herbert, 2 = Tully, 3 = Innisfail, 4 = Atherton, 5 = Paluma– Seaview, 6a = Cardwell and Kirrama ranges, 6b = Hinchinbrook Island, 7 = Bellenden Ker-Lamb Range, 8 = Macalister, 9 = Daintree–Bloomfield. Notophyll vine forest is the major vegetation type on granite uplands across the region, whereas simple microphyll vinefern forest/thicket predominates on cloudy wet uplands and highlands (Tracey 1982). Rocky granite outcrops on drier areas in lowlands and foothills support deciduous microphyll vine thicket. The best-developed rainforest (complex mesophyll vine forest) that generally occurs in the foothills of the highest mountains and on basaltic soils in upland areas of the Atherton Tableland, has largely been lost by clearing for grazing and agriculture. The few areas of mesophyll rainforest which do remain, including those on cloudy uplands, are mostly preserved in National Parks e.g Mossman Gorge, Daintree, Wooroonooran, Crater Lakes, Tully Gorge, Russell River, Thornton Peak. These areas, and the forest around Cape Tribulation, are extremely rich in bryophytes. Pockets of wet sclerophyll forests (the tall open-forest of Specht 1970) and eucalypt woodlands occur in the drier areas (1500–2000 mm per annum rainfall) and on western fringes of the Atherton and Windsor Tablelands (Tracey & Webb 1975, Tracey 1982, Kitching 1987). Plant species diversity is particularly high. Vascular plant species in the Wet Tropics number 4 500, of which approximately 3 000 are recorded from the Wet Tropics World Heritage Area (WTMA 2002). Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland 375 Brachymenium nepalense, epiphytic in pioneer rainforest at edge of cleared area, Longlands Gap, near Atherton (Province 4). Leaves 0.9–1.6 × 2.4–4.8 mm. North Johnstone River flowing through mesophyll vine forest near Malanda, Atherton Tableland (Province 4). 376 Cunninghamia 8(3): 2004 Dawsonia polytrichoides growing on tree stump in wet sclerophyll forest dominated by Syncarpia glomulifera, Paluma Range, near Ingham (Province 5). Leaves linear, 6–10 mm long. Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Dawsonia polytrichoides, Paluma Range, near Ingham (Province 5). Male plants; leaves linear, 6–10 mm long. Birthday Creek, simple notophyll vine forest, alt. 800 m, Paluma Range, near Ingham (Province 5). Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland 377 Epiphytes at Wallaman Falls, near Ingham, Girringun National Park (previously known as Lumholtz National Park) (Province5). Distichophyllum mittenii growing on filmy fern (Cephalomanes brassii) adjacent to creek in simple notophyll vine forest, Paluma Range, near Ingham (Province 5). Leaves 2.0–2.8 × 1.0–1.2 mm. Macromitrium involutifolium, epiphytic on Alphitonia petrei, Paluma village, near Ingham (Province 5). Leucobryum candidum, an epiphyte in simple notophyll vine forest, Paluma Range. Shoots 1–5 cm long. 378 Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Lopidium struthiopteris growing on a sapling in simple notophyll vine forest, alt. 900 m, Paluma Range, near Ingham. Surrounded by eucalypt vegetation, Wallaman Falls in Girringun National Park is the longest single-drop waterfall in Australia, falling 305 m to a large pool (Province 5). Curran Creek, Kirrama Range, near Cardwell, flowing through simple notophyll vine forest, alt. 650 m. (Province 6a). Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland 379 Leucobryum aduncum, Kirrama Range, near Cardwell. Leaves 2–4 mm long. Mesochaete taxiforme growing adjacent to creek, alt. 650 m, Kirrama Range, near Cardwell. (Province 6a) Leaves 3.4–6.0 × 1.5–2.7 mm. Hinchinbrook Island from the mainland. (Province 6b). 380 Cunninghamia 8(3): 2004 Creek on the summit of Mt Bellenden Ker, in Wooroonooran National Park, near Babinda, alt. 1550 m. (Province 7). Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Dracophyllum sayeri (Epacridaceae) understory in simple microphyll stunted vine-fern thicket dominated by Leptospermum wooroonooran, on summit of Mt Bellenden Ker, near Babinda. (Province 7). Ectropothecium zollingeri and Fissidens crispulus growing on rock, upper Mulgrave River, west of Mt Bartle Frere. (Province 7). Mossy rocks and Helmhotzia sp. (Philidraceae), alt. 1550 m, summit of Mt Bellenden Ker, near Babinda. (Province 7). Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Epiphytes at Tchupalla Falls, Wooroonooran National Park, near Innisfail. (Province 7). Powellia involutifolia. (Province 7). Leaves 1.5–2 mm × 0.7–1 mm. 381 382 Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Taxithelium merrillii carpeting mangrove mud, Russell River estuary. (Province 7). Upper Mulgrave River flowing through mesophyll vine forest. (Province 7). Tchupalla Falls surrounded by mesophyll vine forest, Wooroonooran National Park, near Innisfail. (Province 7). Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Aims This study was undertaken to provide an up to date checklist of the mosses of the Wet Tropics bioregion and to determine the distribution of species within the individual provinces of the region. In addition, the distribution of each species listed was compared with its phytogeographical affinities in areas close to Australia such as SE Asia (particularly Malesia, Papua New Guinea), New Zealand, New Caledonia, Norfolk Island etc. A comprehensive listing of published literature relevant to tropical Australian mosses was compiled. 383 1970; Shaw 1985; Spence 1996, 2004; Spence & Ramsay 1996, 2002, 2005 (in press, Flora of Australia for Bryaceae); Stone 1975–1997 [see reference list]; Stone & Catcheside 1993; Stone & Scott 1973; Streimann 1991a,b,c, 1993, 1997, 1999, 2000a, 2001; Syed 1973; Tan et al.1996, 1998; Tangney 1996,1997; Touw 1971, 2001a,b; Touw & Falter-van den Haak 1989; Vitt 1984, 1989; Vitt & Ramsay 1985a,b. Additional literature of use includes Buck et al. (2002), Crosby et al. (1992), Crosby & Magill (1978), Ramsay (1987), Ramsay and Schofield (1987), Ramsay et al. (1987), and Ramsay & Seur (1994). Methods This present study looks at Wet Tropics mosses in detail, including an update of species taxonomy incorporating data from revisions, new species records, and reports of additional species not previously recorded from the region. The data presented here have been obtained from literature, early collections in Herbaria including BRI, CANB (includes CBG), NSW, as well as the more recent collections made by I.G. Stone (MELU now at MEL), H. Streimann (CANB), D.H. Vitt (ALTA & NSW), H.P. Ramsay (NSW), W.B. Schofield (UBC & NSW), J.R. Spence (NSW), and others made as part of field studies during revisions undertaken for the Flora of Australia (Bryophytes). In addition, some collections made in the 1960s and 1970s e.g. those of B.O. van Zanten (GRO, NSW made in 1968), W.A. Weber (CO made in 1968) and D.H. Norris (HSC made in 1970s) have also provided useful information. The moss checklist (Appendix 1) contains names in current use. The updated Catalogue of Australian Mosses (Streimann & Klazenga 2002) has been particularly useful for checking synonyms and for tracing literature. The Catalogue also includes important information on the distribution of the various species in Australian States other than Queensland for comparison and is not repeated here. The checklist (Appendix 1) includes data on location of species within provinces of the Wet Tropics bioregion in Australia, indicates endemics (WT) within the bioregion and those more widespread in Australia (AU), and contains distribution relationships and affinities to areas adjacent to Australia. During the last thirty years many new species have been described from north-east Queensland and additional species recorded as a result of intensive collecting and revisions. Publications include Allen 1981, 1987; Ando 1972, 1977; Buck 1979, 1980, 1982, 1990; Buck & Crum 1978; Buck & Tan 1989; Catcheside & Frahm 1985; Catcheside & Stone 1988; Crum 1971, 1986, 1991; During 1977; Ellis 1985,1991; Enroth 1991, 1995, 1996; Frahm 1987a,b, 1988, 1990, 1991, 1992, 1993, 1994; Hedenäs 2002; Hyvönen 1989a; Ireland 1992; Isoviita 1986; Klazenga 2003; Koponen 1980, 1982, 1988, 1990; Kruijer 2002; Lai 1992; Lewinsky 1984, 1990; Miller & Manuel 1982; Mohamed 1979; Norris & Robinson 1979; Nowak 1980; Ochi 1970, 1973, 1980, 1982; Ochyra 1986; Ramsay et al. 2002a,b, 2004; Reese1989, 1992; Reese & Stone 1987, 1995; Salazar Allen 1993; Schultze-Motel Results Moss Diversity Of the 1074 accepted species of mosses in Australia, 527 are found in Queensland (Streimann & Klazenga 2002). In the Wet Tropics bioregion the present estimate of diversity is about 408 taxa (398 species, 2 subspecies, and 8 varieties) in 159 genera and 62 families. Data presented here have increased the previous estimate (Ramsay et al. 1987) of the number of species by 83 and added 20 additional families and 29 genera. The checklist (Appendix 1) includes 21 new records for the Wet Tropics and 8 new species for Australia. Lists of genera and their families, and families and their genera are presented in Appendices 2 and 3. Families with the highest species richness in the area include Bryaceae, Calymperaceae, Dicranaceae (Table 1). Table 1: Families of mosses with the highest species richness in the Wet Tropics. Family Bryaceae Calymperaceae Dicranaceae Meteoriaceae (incl. Papillaria) Sematophyllaceae Number of genera Number of species 5 7 7 8 23 47 29 13 16 32 (incl. 2 varieties) A few genera have many species in the area e.g Fissidens, Macromitrium, Syrrhopodon (Table 2), but many genera (see Appendix 1) are represented by only one to three species. Table 2: Genera of mosses with the largest species representation in the Wet Tropics Genus Number of species Fissidens Macromitrium Syrrhopodon Calymperes Campylopus Mitthyridium Leucobryum 35 17 16 15 12 9 8 (incl. 2 varieties) (incl. 2 subspecies) (incl. 2 varieties) (incl. 1 variety) 384 Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Creek at Mt Lewis (Province 9), complex notophyll vine forest, alt. 800 m. Ephemeropsis tjibodensis, an epiphyllous moss, under high power (× 400). (Province 9). Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Discussion Distribution of mosses in the Wet Tropics Topography and rainfall patterns combine in the region to support a variety of vegetation communities, including mangroves, woodlands and rainforests of different structural types (Tracey 1982) which provide specialised habitats for bryophytes. Mangrove communities: Mangroves have a disjunct occurrence along the east coast in intertidal areas from Torres Strait to Victoria but reach their maximum diversity in northern Australia (Specht & Specht 1999). Mosses are not common in mangroves but a few terrestrial and epiphytic species have adapted to this habitat. There are no studies of bryophytes, which include mosses, for the mangrove systems of north-east Queensland but Windolf (1989) made a study of bryophytes in an area in south-east Queensland. He recorded three epiphytic mosses and 15 leafy liverworts. Of the mosses, only Calymperes was recorded on mangroves, the other two species being on trees at the edges of the mangrove community. A study in Thailand by Thaithong (1984) recorded five moss species (including two species of Calymperes) and 21 leafy liverworts. It is clear that mosses are rare in such communities. One notable exception is Taxithelium merrillii, a terrestrial species which is found as extensive mats on mud and exposed mangrove roots, particularly in mangroves that have a high freshwater input e.g. Noah Head NP, Cape Tribulation, and Russell River (Ramsay et al. 2002a). This moss is also recorded in similar communities in the Philippines (Bartram 1939, Iwatsuki & Tan 1979; Tan & Iwatsuki 1991) and Southeast Asia (Tan & Iwatsuki 1993). On Hinchinbrook Island, T. merrillii carpets sandbars in a heavily shaded intertidal region of Mulligan’s Creek, associated with Melaleuca quinquenervia forest (R. Lovatt pers. comm.). Unlike T. merrillii, species such as Calymperes spp., Syrrhopodon spp., Macromitrium aurescens, Leucophanes sp., and Octoblepharum albidum, epiphytic on mangrove trees, are not subject to frequent tidal inundation and are not restricted to this habitat. An ephemeral species with persistent protonema, Archidium minutissimum was recorded from sandy soil at the edge of mangroves south of Cooktown where it was submerged at exceptionally high tides (Stone 1985b). Three other collections in the area were not associated with a mangrove habitat. The species has not been seen since 1984 and is worth further investigation in season. Freshwater habitats: Aquatic mosses are not abundant in streams in the region. Obligate aquatics are more common in cooler climates (Vitt & Glime 1984, Suren 1993); however, many ground-dwelling moss species may also be found growing submerged in freshwater habitats (Scott 1994). In most stream systems, variable current velocities, water levels and humidities contribute to broad environmental gradients from aquatic to terrestrial habitats. In streams at higher elevations where water temperatures are generally lower, some species of genera such as Bryum, Philonotis, 385 Distichophyllum, Hypnodendron and Fissidens are common both above water and immersed, particularly in areas of low flow. The morphology of certain species appears to be influenced by environmental conditions e.g. fronds of Hypnodendron vitiense subsp. australe extend much further when growing submerged (A. Touw pers. comm.). The occurrence of mosses in freshwater habitats in the Wet Tropics may be more common than previously recorded. An unpublished study on the Atherton Tableland (average elevation 700–800 m) showed there is often extensive growth of submerged mosses. In Mazlin Creek, near Atherton, the cosmopolitan moss Leptodictyum riparium flourished in a fast-flowing stream where nutrient levels were high. Biggs (1996) determined that bryophytes dominate in steeper headwater streams with stable substrates that promote fastflowing, turbulent waters, often with high flow variability. However, Taxiphyllum taxirameum was also recorded near Atherton, growing abundantly in a permanent spring-fed pool with minimal current velocity, with a constant water temperature of around 22°C. The flow environment of stream bryophytes in the Wet Tropics warrants further attention. More intensive surveys are clearly needed. The Wet Tropics endemic Touwia laticostata is known from only one stream (altitude 300 m) on Mt Bellenden Ker (Ochyra 1986). The aquatic moss Platyhypnidium muelleri is also reported from that area (Scott 1994), as is Platyhypnidium austrinum. In his overview of the Brachytheciaceae in Australia, Hedenäs (2002) concluded that Australian specimens of P. muelleri are more likely to be P. austrinum; however, both species have been confirmed in the Wet Tropics. A recent survey (Cairns & Werren 2002, unpublished) of the upper East Mulgrave River revealed extensive submerged beds of Ectropothecium zollingeri, previously unrecorded in the Mulgrave catchment, in a shaded riffle. The only record of Sphagnum in the area is Sphagnum perichaetiale from a Melaleuca swamp near Bramston Beach (Stone 1990c). Terrestrial mosses: Terrestrial bryophytes act as colonisers of soils or in drier areas of south-western Queensland they form an important component of soil crusts (Eldridge et al. 2000). Bryophytes are not abundant on the earth floor of tropical rainforests, perhaps because the continual build-up of leaf litter restricts their development. Nevertheless, in wetter areas in the Wet Tropics, the soil-binding capacity of some terrestrial mosses is significant and they are able to quickly colonise surface soil of newly disturbed steep earth banks and reduce erosion. In the Wet Tropics, Pogonatum and Dawsonia colonise large areas of the more exposed road cuttings, and Dicranella and more rarely Garckea, the more protected cuttings. Many species of Fissidens are prominent as colonisers of shaded earth banks or forest tracks, some in coastal areas e.g. F. crispulus, F. perobtusus, and others at higher altitudes e.g. F. dietrichiae, F. pallidus. Tiny earth mosses such as Archidium, Erpodium, and Gigaspermum are often present but may be hard to find. The families Bryaceae (e.g. Rosulabryum spp., Gemmabryum spp.), Dicranaceae (e.g. Dicranella, Campylopus, Dicranoloma), Funariaceae 386 Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland (e.g. Funaria hygrometrica) and Pottiaceae (e.g. Barbula) (Zander 1993) include many taxa of importance in colonising bare soil or shallow soil over rock surfaces. Epiphytic and epiphyllous mosses: The rainforest presents a range of microhabitats and substrates for bryophytes to colonise, depending on the diversity of microclimates and their individual ecological tolerances. Pócs (1982) observed the zonation of epiphytes in rainforests in Africa, both vertically and horizontally, and differentiated four broad zones on the phorophyte – basal trunk, main trunk, branches, and terminal twigs. The spatial distribution of corticolous bryophytes is therefore determined by a network of fluctuating variables e.g. light intensity, temperature, relative humidity, air currents (Smith 1982, Franks & Bergstrom 2000). Epiphytic bryophytes contribute significantly to the diversity of tropical rainforests worldwide. In lowland forests in French Guiana, Montfort and Ek (1990) found 154 species of bryophytes, (42% were mosses), on 28 trees representing 22 species. Along an altitudinal gradient in the Columbian Andes, Wolf (1993) found that mosses comprised 36.6% of the 295 epiphytic bryophyte taxa identified. In south-east Queensland, Franks (2000) and Franks and Bergstrom (2000) found that epiphytic bryophytes formed a significant component of microphyll fern forests. Sampling on Nothofagus moorei trunks was carried out from ground level to 2 m on 25 individual trees and showed that the composition and community structure altered with both height above ground level and the direction of exposure (Franks & Bergstrom 2000). Patterns of distribution were thought to reflect changes in moisture availability and degree of desiccation tolerance of the various taxa. Of forty-three bryophyte taxa, 37% were mosses. These represented a range of families and genera including Calymperaceae: Syrrhopodon armatus; Dicnemonaceae: Eucamptodon muelleri; Dicranaceae: Dicranoloma spp.; Hookeriaceae: Cyathophorum bulbosum; Hypnaceae: Hypnum cupressiforme; Hypopterygiaceae: Lopidium concinnum; Lembophyllaceae: Lembophyllum divulsum; Leucobryaceae: Leucobryum candidum; Orthotrichaceae: Macromitrium exsertum; Ptychomniaceae: Hampeella pallens; Rhizogoniaceae: Pyrrhobryum paramattense; Sematophyllaceae: Wijkia extenuata. No such studies have yet been undertaken for the Wet Tropics, although observations indicate that the species composition of rainforest epiphytes would include some of the species recorded by Franks and Bergstrom (2000), but are also likely to include a range of taxa of tropical origin not found in the south-east forests. Examples include various genera and species in families including Calymperaceae, Sematophyllaceae, Orthotrichaceae and Meteoriaceae. Epiphyllous mosses occur in the wetter forests and moist rainforest gullies of the Wet Tropics e.g. Mt Bellenden Ker summit, Mt Lewis. Again, no studies have been reported. Worldwide, epiphyllous mosses are uncommon compared with tiny leafy liverworts (Lejeuneaceae), and in tropical America, Africa and Asia are mostly members of the Hookeriaceae (Gradstein 1997, Bates 2000). In the Wet Tropics this does not appear to be the case, although Distichophyllum mittenii has been found growing on the leaves of filmy ferns (Cephalomanes brassii) (unpublished data). Obligate epiphyllous bryophytes tend to be minute plants, growing appressed to the leaf surface, anchored by bundles of rhizoids (Gradstein 1997). Few mosses fit this description. One notable exception is Ephemeropsis tjibodensis, which consists of a protonemal gametophyte and forms dense algal-like patches on the surface of leaves. Most mosses growing on leaves are facultative epiphylls (Gradstein 1997), spreading onto angiosperm leaves from adjacent twigs e.g. members of the Meteoriaceae: Barbellopsis trichophora, Aerobryopsis longissima, Meteorium polytrichum. Richards (1984) noted that different types of forest structure affect levels of light and humidity, providing a range of microclimates within the forest. The bryological diversity of the Wet Tropics bioregion may be a reflection, in part, of differences between rainforest types and the variety of microhabitats they offer. However, to date, few bryologists have listed the rainforest habitat of moss collections according to these classifications (one notable exception is Reese & Stone 1995), and there have been no published analyses of bryophyte communities in relation to rainforest types. Mosses of other vegetation communities: ‘Dry’ rainforest (Gillison 1987), Casuarina dominated riparian vegetation, and other non-sclerophyll communities have been highlighted as worthy of investigation (Streimann 1994, 2000b). Patches of these vegetation types occur throughout the Wet Tropics, particularly in areas of lower rainfall; nevertheless, an understanding of the bryophyte flora of these habitats is lacking. A survey of the moss flora of dry rainforests to the west of the Wet Tropics revealed a unique assemblage, comprising species from wetter areas and species from drier environments (Fensham & Streimann 1997). Of these habitats, Streimann (1994) commented: ‘The species numbers may not be so great, nor the colonies as spectacular, but they do contain an interesting and poorly studied bryophyte flora which may be bryogeographically exciting and significant’. Mosses with a restricted distribution Species with restricted distributions include those limited to high mountain peaks and lowland forests. In addition, many Southeast Asian mosses (Appendix 1) are at the limits of their range in the Wet Tropics (Reese & Stone 1995) e.g. Calymperes porrectum recorded only from Cape Tribulation, Macromitrium incurvifolium recorded from Big Tableland south of Cooktown, Mt Lewis, and the Russell River. Others have restricted altitudinal ranges e.g. Syrrhopodon prolifer which has one variety, var. prolifer, found above 1200 m, and the other variety, var. mossmanensis found below 500 m (Fig. 4c). While this species is pantropical, the varieties are each Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland 387 represented in Queensland from a single locality, the type variety S. prolifer var. prolifer from Thornton Peak and the other S. prolifer var. mossmanensis from Mossman (Reese & Stone 1995). Mosses restricted to high mountain peaks: Mountain peaks receive high levels of rainfall and high altitude forests (‘cloud’ forests) may also capture considerable amounts of water from passing clouds (Foster 2001). They therefore offer unique habitats not found elsewhere in the tropics. In tropical regions worldwide, the mass and diversity of both mosses and liverworts increases with altitude (Buck & Thiers 1989). The Wet Tropics is no exception, and bryophytes are particularly abundant in the cloud belt (1200–1600 m) in simple microphyll vine-fern thicket (Webb 1968). Epiphytic mosses are widespread, but some are known only from one or two sites e.g. Calyptrochaeta rotundifolia, Clastobryum dimorphum, Daltonia contorta, Macromitrium funiforme, Macromitrium dielsii (Mt Bellenden Ker and/or Mt Bartle Frere), Meiotheciella papillosa (Mt Spec near Townsville). Some mosses which are common in lower altitudes in southern Australia and are also in New Zealand e.g. Calyptrochaeta apiculata, Cyathophorum bulbosum, Daltonia splachnoides, Dicranoloma menziesii and Warburgiella leucocytus, are present only on the highest peaks in the Wet Tropics, where cooler temperatures prevail and there is constant mist and cloud. Streimann (2000b) nominated isolated peaks and ranges in the Wet Tropics as ‘hot spots’ both for endemics, and species with affinities to Southeast Asia and Malesia (see Tan & Iwatsuki 1999). Fig. 4A. Distribution of Calymperes species by altitude (Reese and Stone 1995). Contemporary climate change has been nominated by Williams et al. (2003) as a significant threat to the long-term preservation of the biota in the tropical rainforests of the Wet Tropics. Complex notophyll vine forests, simple notophyll and simple microphyll forests and thickets in the wet uplands and highlands have been identified as particularly sensitive to even moderate increases in temperature (Hilbert et al. 2001). On the highest mountains, the cloud base is expected to increase in altitude and consequently reduce mist (Foster 2001, Hilbert et al. 2001). Extinction risks for bryophytes endemic to these unique habitats are high. Fig. 4B. Distribution of Mitthyridium species by altitude (Reese and Stone 1995). Mosses of the lowland coastal belt: Included here, apart from terrestrial mosses, are many epiphytic species in high rainfall areas at lower altitudes: e.g. Chaetomitrium tahitense, Leucophanes spp., Octoblepharum albidum, Leucobryum spp. Most taxa in the family Calymperaceae (e.g. Calymperes, Mitthyridium and Syrrhopodon) are well represented at lower altitudes, generally below 500 m (Fig. 4 A–C) (Reese & Stone 1995) with very few restricted to higher altitudes. The family Sematophyllaceae includes species such as Acanthorrhyncium papillatum, Radulina hamata, Taxithelium instratum, T. nepalense and Trismegistia lancifolia, that are well represented in lowland regions such as Cape Tribulation, Mossman Gorge, Daintree National Park. (Ramsay et al. 2002a). Fig. 4C. Distribution of Syrrhopodon species by altitude (Reese and Stone 1995). 388 Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Bryophytes are generally thought to be few in lowland tropical forests compared with forests at higher altitudes. Streimann (1994) suggested that bryophyte diversity in these habitats is, in fact, much greater than it appears; many species are restricted to tree canopies and are not easily accessible. Species widely distributed throughout the area: Some species have wide-ranging geographic distributions throughout the area, at various altitudes. Species such as Bryobrothera crenulata and Callicostella papillata var. papillata are found from Mt Finnigan in the north, to Eungella south of the area (Streimann 2001). Hookeriopsis utacamundiana ranges from Mt Bellenden Ker and Mt Bartle Frere to Eungella, some 800 km south. Others that are more widely dispersed in the tropics e.g. Calymperes erosum, distributed across northern Australia (QLD, WA & NT), occur as far north as Cape York and extend south to Proserpine (Reese & Stone 1995). Phytogeographical affinities of mosses in the Wet Tropics Affinities of the bryoflora of this tropical part of Australia range from endemic species, to species of Gondwanan origin, present primarily in Australia and New Zealand but also in South America and southern Africa. Gondwanan taxa occur frequently in ancient rainforests or montane ecosystems. Many have palaeotropical affinities with species in Malesia and Southeast Asia. Others have relationships to the flora of New Caledonia and/or Oceania and the Pacific. Some cosmopolitan species are also represented e.g. Funaria hygrometrica. Species reported as endemic to area The Wet Tropics bioregion is well known for the ancient nature and high degree of endemism of vascular plants (Keto & Scott 1986), and has been nominated as a significant centre for endemism (Boden & Given 1995, Crisp et al. 2001). Surprisingly, present studies have found that the number of moss species endemic to the Wet Tropics bioregion is much lower than expected, representing less than 7% of the species present (Appendix 1 as WT). This disparity can perhaps be explained by the comparative ease of dispersal of bryophytes by spores or asexual propagules (Laaka-Lindberg 2003). Similar rates of endemism have been estimated for tropical regions elsewhere e.g. Galapagos Islands 10%, Cuba 12%, Mt Kilimanjaro 6% (Frahm 2003). Taxa formerly considered to be endemic may now be in synonymy under another name or, particularly in the case of small earth mosses, may yet be found in other countries. Others may possibly be synonymous with Southeast Asian mosses at the extremity of their range. Endemic species of Fissidens are associated with moist lowland areas. Isolated mountain peaks are particularly important habitats for some taxa e.g. Mt Finnigan (Calyptrochaeta brassii), Mt Bellenden Ker, Mt Bartle Frere and Mt Lewis (Clastobrum dimorphum) (Streimann 2000b). Endemic species come from a wide range of families and genera e.g. Brachytheciaceae: Rhynchostegium nanopennatum Buxbaumiaceae: Buxbaumia thorsborneae; Fissidentaceae: Fissidens flabellulus. var. eachamensis, F. gymnocarpus, F. henryae; Hypnodendraceae: Hypnodendron comatulum; Neckeraceae: Touwia laticostata; Orthotrichaceae: Macromitrium dielsii, M. funiforme. More widespread Australian endemics in the area Many Australian endemic species (AU in Appendix 1) that are found in the Wet Tropics have a pattern of distribution along the east divide, often in pockets of remnant forest e.g species belonging to Dicranoloma, Eucamptodon, Fissidens, Macromitrium, Papillaria, Schlotheimia. Although this study has added about 83 additional taxa to the total previously recorded from the area, the numbers of Australian endemic species (AU), including those that occur exclusively in the Wet Tropics (WT), has been reduced in this study compared to the earlier estimates made by Ramsay et al. (1987), to about 25% species. The genus with the largest numbers of endemic species is Fissidens 14 spp. (including 7 in WT), with Macromitrium 12 spp. (2 in WT), Archidium 6 spp. (1 in WT); and 12 genera are each represented by a single endemic species. Moss species occurring across northern tropical Australia A number of species from the Wet Tropics bioregion also occur throughout northern tropical Australia including north Queensland, the Northern Territory and/or northern Western Australia (Table 3). Little is known about the wetter parts of the Northern Territory, particularly the moist monsoon vegetation of Arnhemland, an area highlighted by Streimann (2000b) as worthy of further exploration. Families well represented across the northern tropics include Archidiacae, Bryaceae, Calymperaceae, Fissidentaceae and Sematophyllaceae. Many of these are also distributed in the palaeotropical regions of Southeast Asia, particularly Malesia. In the Calymperaceae for example, while 42 taxa in three genera occur in Australia (Reese & Stone 1995), the majority are present in the warm humid tropics of the Northern Territory and north-east Queensland. Across this region, Syrrhopodon has 18 taxa and Calymperes 14, while two species (C. tenerum and C. erosum) also occur in northern Western Australia. The other genus in the Calymperaceae, Mitthyridium, occurs as a single species, M. flavum, in the Northern Territory, with the nine Australian representatives all in the Wet Tropics. In the Sematophyllaceae, species such as Taxithelium spp. and Radulina hamata are present in northeast Queensland and the Northern Territory, while Papillidiopsis ramulina has been recorded from north west Western Australia and the Northern Territory but not Queensland (Catcheside & Stone 1988, Stoneburner et al. 1993, Ramsay et al. 2002b, 2004). Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Table 3. Representative moss species occurring across northern tropical Australia Archidium ohioense Archidium rothii Brachymenium indicum Bryum argenteum Calymperes afzelii Calymperes erosum Calymperes graeffeanum Calymperes motleyi Calymperes porrectum Calymperes tenerum Campylopus laxitextus Eccremidium minutum Erpodium coronatum var. australiense Fissidens ceylonensis Fissidens curvatus Fissidens gymnocarpus Fissidens holstii Fissidens perobtusus Garckea flexuosa Gemmabryum apiculatum Gemmabryum indicum Hyophila involuta Leucobryum aduncum Leucobryum aduncum var. scalare Leucobryum chlorophyllosum Leucophanes glaucum Mitthyridium flavum Octoblepharum albidum Pelekium investe Radulina hamata Taxithelium instratum Taxithelium kerianum Taxithelium nepalense Taxithelium planum Taxiphyllum minutirameum Trachyphyllum inflexum On the Cape York Peninsula, north of the Wet Tropics, Crisp et al. (2001) found significant floristic links between the eastern coastal strip and Trans-Fly and Port Moresby ‘dry’ areas in Papua New Guinea. Their recent nomination of the Iron Range-McIlraith Range region to the north of the Wet Tropics as an important centre of endemism for vascular plants suggests that this area should be further investigated by bryologists. Patches of vine thicket on Cape York Pensinsula have revealed several previously unreported genera and species (Streimann 2001a); however, bryological surveys of the Cape are few, possibly due to the difficulties of access. Similarly, islands of the Torres Strait, between Australia and New Guinea, are bryologically unknown, although Streimann (2001) suggests that there is no reason why, for example, Hookeriaceae should not occur there. 389 Quaternary climatic oscillations (Williams et al. 1993), would be a mechanism for increased diversity through random processes. Extrapolation of these conclusions to include all plants should, however, be approached with caution; bryophytes employ significantly different dispersal mechanisms (e.g. spores, fragments, gemmae) compared with flowering plants. Tan and Pócs (2000) emphasised that a regional bryoflora is developed as a result of the complex processes of long-distance dispersal, vicariance events, extinction, and the vicissitudes of climate during the Tertiary and Quaternary. Affinities with the mosses of Malesia and Southeast Asia Relationships exist between Australian bryophytes and those of the Palaeotropics, particularly Papua New Guinea, the Philippines, Borneo and other parts of Southeast Asia. About 45% of the moss species in the Wet Tropics have affinities with those regions (Appendix 1). Within Malesia, the Malay Peninsula, Sumatra, and Borneo in the west, and New Guinea in the east, form cores of ever-wet climate. These border a mosaic of wet and seasonal areas (the Philippines, Sulawesi, the Moluccas, Java, and the Lesser Sunda Islands), which form a corridor connecting the seasonally dry areas of tropical Southeast Asia and Australia (Touw 1992b). The Moss Flora of Malesiana (Eddy 1988, 1990, 1996) lists many species common to both areas. By comparison affinities of Wet Tropics mosses with those of temperate rainforests of New Zealand is only 15% (Appendix 1). Afffinities with the mosses of Southeast Asia and New Caledonia Klazenga (1999) reported that the genus Dicranoloma shows a Malesian-Australasian-Pacific distribution extending marginally into Southeast Asia. The genus D. braunii is widespread in Malesia but in Australia occurs only in north-east Queensland, while D. menziesii occurs in Papua New Guinea and is found throughout eastern Australia, New Zealand and New Caledonia (Klazenga 2003). Floristic affinities: Strong affinities exist beween rainforest angiosperms in the Wet Tropics, Malesia and New Caledonia (Morat 1986). Based on cladistic biogeographical analyses, Crisp et al. (1999) reviewed relationships between distribution patterns of angiosperm taxa. They described a northern element of the ‘South Pacific track’ which appeared to link New Guinea, Queensland, Northern Territory, New Caledonia and Fiji, and concluded that ‘the flora of Australia is most closely related to that of its Gondwanan neighbours’, noting that there are biogeographical anomalies yet to be resolved. Acknowledgement of tropical flora as ‘palaeoautochthonous, derived from Gondwanan stock’ (Webb et al. 1986) is now widely accepted (Specht & Specht 1999). Walker (1990) agreed that the specific interrelationships of some groups of plants [and animals] in rainforests imply a long evolutionary association i.e. the occurrence of species together is more than just chance and is indicative of the maintenance of rainforest assemblages through time (Webb et al. 1986). Re-assembly of tropical forests by expansion from refugia (Webb & Tracey 1981), as has occurred as a result of Between Australia and the Lesser Sunda Islands (South Malesia), species such as Sclerodontium pallidum and Thuidiopsis sparsa are seasonal drought-tolerant plants from more or less exposed habitats (Touw 1992, 2001a). These distribution patterns might be explained by exchange of biota between Malesia and Australasia long before the collision of Gondwana and Laurasia; however, Touw (1992) considered these patterns to be the result of post-glacial establishment during the Miocene. Similarly, Mittenia plumula, reported from Mt Spec in the southern Wet Tropics bioregion but more common in eastern Australia, Tasmania, Papua New Guinea and New Zealand, also occurs in Western Malesia, probably as the result of post-glacial colonisation (Tan 1998). Nevertheless, the Australian element is far smaller in the Lesser Sunda Islands than one would expect from their proximity (Touw 1992). The family Thuidiaceae includes species such as Pelekium gratum and P. synoicum with affinities to Southeast Asia. One species, P. velatum, so far reported only from central coastal Queensland by one record (Touw 2001b) but common in Malesia, could well be found in the Wet Tropics. 390 Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland The same is true of affinities between New Guinea and Australia (Piipo and Koponen 2003). Montane forests in New Guinea and South Malesia include only 12 species of mosses with Australian affinity (Ramsay et al. 1986). In the genus Macromitrium, well represented in both New Guinea (29 spp.) (Vitt et al. 1995) and Australia (22 spp.) Vitt and Ramsay 1985a, b), only two species are common to both areas. Examples of species with restricted distribution between north-east Australia and New Guinea include Acroporium lamprophyllum var. percaudatum, Gemmabryum australe, Groutiella tomentosa, Macromitrium incurvifolium, Orthomnium elimbatum, Pogonatum neesii, Sorapilla papuana (Norris and Koponen 1989a, b; van Zanten and Pócs 1981, Vitt et al. 1995, Tan et al. 1996, Ramsay et al. 1995 Spence & Ramsay 2005, Flora of Australia vol 51 in press). In an analysis of the biogeography of the Bryaceae (subfamily Bryoideae) and its relatives in tropical regions of north-east Queensland, Spence and Ramsay (1996) found that the strongest affinities are with Malesia (including New Guinea). The family Sematophyllaceae has strong affinities with Malesia with many species of the genera found in Australia such as Acroporium, Acanthorrhynchium, Clastobryum, Meiothecium, Taxithelium, Trichosteleum, Trismegistia, Warburgiella, and Radulina having species common to these areas (Ramsay & Schofield 1987; Ramsay et al. 2002a, b, 2004; Tan et al. 1992). Gondwanan distributions in temperate rainforest zones of Australasia and South America, as well as in the tropical montane forests of Malesia and Sri Lanka, suggest fragmentation of a continuous area forming a mosaic of wetter and drier environments. Gondwanan families such as Dawsoniaceae (van Zanten 1973), Hypnodendraceae (Touw 1971), Racopilaceae (van Zanten & Pócs 1981, van Zanten & Hofman 1995) and Leptostomataceae (Crum 1989) also occur in New Guinea and the western Pacific. Genera such as Campylopus have distribution patterns that include Africa (Frahm 1992) as well as Southeast Asia. Distribution of the genus Papillaria (Meteoriaceae) includes both Malesia/Southeast Asia and the Pacific, with greatest diversity in New Caledonia (Streimann 1992). In the Daltoniaceae [Hookeriaceae], Calyptrochaeta apiculata has a disjunct distribution, occurring in the Wet Tropics on Mt Bellenden Ker and in southern Australia, but also reported from Chile, Argentina, the Falkland Islands, and New Zealand (Streimann 2000a). Distunctive patterns of distribution may be difficult to resolve. In the Neckeraceae, Caduciella marei, known in Australia from one specimen collected near the Daintree River in 1882 (Enroth 1991), has also been reported from Tanzania, the Comoro Islands, India (Assam), SW China, Indochina, Malesia and Oceania. Tan (1998) considered this widespread distribution to represent either fragmented parts of a former continuous Gondwanan range or the result of chance dispersal between extant populations on three continents. Commenting on disjunctive patterns of bryophyte distribution between South Malesia/Phillippines and Australasia/Oceania, Tan (1998) attributed the presence of Gondwanan taxa in Malesia e.g. Bescherellia elegantissima and Dawsonia superba (also present in the Wet Tropics), to their arrival following the collision of SE Asia and the Australian margin in the mid-Miocene; reciprocal invasions into northern Australia could also have occurred during this period. Affinities of moss species across tropical Australia and Malesia/Southeast Asia demonstrate that there are strong phytogeographical relationships between the palaeotropical areas and north-east Queensland. This may be at the family, generic or species level. Taxa come from a wide range of families including Archidiaceae, Bryaceae, Calymperaceae, Daltoniaceae, Fissidentaceae, Meteoriaceae, Orthotrichaceae, Thuidiaceae, and Sematophyllaceae. Species in the Wet Tropics with affinities to the mosses in Southeast Asia (particularly Malesia) are indicated in Appendix 1. Publications pertaining to the bryoflora of southeast Asia that may also have relevance to the Wet Tropics bioregion include: Akiyama et al. (1991); Enroth (1990); Koponen and Norris (1983); Koponen et al. (1986); Mohamed (1998); Norris and Koponen (1987, 1990a, b); Reese et al. (1986a, b); Tan (1994); Yamaguchi (1993). Index Muscorum (Wijk et al. 1959–1969) has been useful for checking distributions. Affinities with New Caledonia: All moss families occurring in New Caledonia are also present in Australia. The strong Southeast Asian element in north-east Queensland also extends to the Pacific; further to the west, the diversity of this element decreases. Previously it was thought that relationships between the species in New Caledonia and Australia were not close but recent studies have increased the number of species common to both and reduced the number of endemics in both countries (see Appendix 1). It is expected that this trend will continue. Knowledge of distributions between the two is still limited for many genera. Pursell and Reese (1982) provided a list of species reported for New Caledonia. Of the 45 names listed in the genus Macromitrium, six occur in Australia (Vitt & Ramsay 1985a), in the Meteoriaceae six of the 16 species are found here (Streimann 1991a, b, c; 1992, 1993), in the Bryaceae six of 22 species including Rosulabryum subfasciculatum (Pursell & Reese 1982, Spence & Ramsay 1996), and in the Sematophyllaceae 11 of the 63 species reported for New Caledonia occur in Australia (Tan et al. 2002). Yamaguchi and Iwatsuki (1987) reduced the previous figure of 15 species of Leucobryum, including seven endemics, to four species with no endemics. Three of these species occur in Australia, including north-east Queensland. Pursell and Reese (1982) listed 39 taxa (also 50 nom. nud.) for Fissidens, of which 33 were endemics. A revision of the Fissidentaceae by Iwatsuki (1982) reduced the number of species to 23 including 13 endemics. Additional studies by Iwatsuki and Suzuki (1989) recorded 27 species and further reduced the number of endemics to three. Nanobryum Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland thorsbornei (Stone 1990b) [= Fissidens thorsbornei], previously known only from Australia, was added to the list. In a discussion on the origins of the New Caledonian species, Iwatsuki (1990) reported six species of Fissidens with an Australian-New Caledonian distribution, but predicted an increase when northern Australian species were investigated. An Australian revision (Stone 1994a) has increased this number to about 23 Fissidens species now known to occur in both countries, including F. rupicola. Two additional Australian species have recently been added to the total for New Caledonia by Müller et al. (2003). Ephemerum fimbriatum, previously known only from Australia, has been reported from New Caledonia by Matsui and Iwatsuki (1991). Most, if not all, of the small cleistocarpous mosses (in Pottiaceae, Ditrichaceae, Ephemeraceae and Archidiaceae) present in New Caledonia also occur in Australia. In addition Viridivellus pulchellum has also been recorded for New Caledonia (Iwatsuki pers. comm.) It is evident that a considerable affinity exists between the terrestrial mosses of the north-east region of Australia and those of New Caledonia. While epiphytic pleurocarpous species appear at present to have less affinity in the two areas, further taxonomic studies may increase the relationships. Affinities with Norfolk Island: Recently a survey of the mosses of Norfolk Island (Streimann 2002) has compared the distribution of species with those in Australia, New Zealand, and New Caledonia. Norfolk Island lies off the coast of Queensland approximately midway between New Caledonia and New Zealand. Affinities of mosses are strongest with Australia (91%) and New Zealand (66%), with only 29% of taxa in common with New Caledonia (Streimann 2002). Sixty nine species of mosses were listed for Norfolk Island, of which 38 (55%) have been recorded in the Wet Tropics bioregion. Just three of these taxa are restricted to Australia and Norfolk Island: Erpodium hodgkinsoniae, Fissidens dietrichiae, and F. oblongifolius var. hyophilus, and within Australia none are limited to Queensland (Streimann & Klazenga 2002). Conclusions In recent years there has been renewed interest in bryophytes; moss species have been described and their distributions in the Wet Tropics reported, generally associated with revisions of families or genera (e.g. Reese & Stone 1995; Eddy 1988, 1990, 1996, Streimann 1997, 1999, 2000a; Klazenga 2003). The majority of these have been published in dedicated bryological journals outside Australia (exceptions are e.g., Buck 1990, Klazenga 2003, Ramsay et al. 2002a), not easily accessible to general botanists and those involved in vegetation surveys. It is therefore not surprising that bryophytes have been frequently omitted from surveys of vegetation, perhaps because few researchers are familiar with their identification. This study summarises the available data on the mosses of the Wet Tropics bioregion and provides a useful and current checklist of moss species, identifying the 391 areas in which the various species are distributed. The Wet Tropics is one of the smallest bioregions in Queensland (Satler & Williams 1999), yet the mosses present represent 77.4% of all Queensland mosses. While the greatest number of collections has been made in the rainforests of the Wet Tropics bioregion, the area is still relatively unexplored, particularly in areas of lowland forest (Streimann 2001). Hopefully, publication of these studies will encourage and facilitate more useful investigation of this area. Other important and interesting regions of Queensland adjacent to the area in the north, south and west of the present studies with different rainfall, vegetation and soil types should eventually provide much of interest to bryologists. Acknowledgements We are most grateful to all those who have given us ready access to collections in both public and private Herbaria as listed in the text. I.G. Stone collected and studied mosses from Queensland over many years with some financial support for collecting at various times provided by ARGC, ABRS. Assistance at various times from ABRS and ARGC for H.P. Ramsay is gratefully acknowledged. Early studies by H.P. Ramsay were carried out while she was on the staff of School of Botany, University of New South Wales. The authors wish to thank the Queensland Parks and Wildlife Service and Queensland Forestry Department for permission to collect. I.G. Stone was particularly grateful to The School of Botany, University of Melbourne for providing facilities and to Alan Stone, A & M. Thorsborne, W.T. Travers, M. Godwin, J. Clarkson and H.S. Curtis, for kind cooperation in many ways. H. Streimann appreciated support and provision of finance from ABRS and CANB ANBG for many collecting trips to north-east Queensland over 15 or more years. We also thank those preparing revisions for the Flora of Australia who have given us access to unpublished data including B.O. van Zanten, T. Pócs, N. Klazenga, S. Gilmore. Our thanks to Judith Curnow (CANB ANBG) for supplying distribution data. The support and encouragement of the Australian Centre for Tropical Freshwater Research (ACTFR) is much appreciated. Assistance in the production of the map (Fig. 1) by Emily Bolitho of Tropical Biology, James Cook University, is gratefully acknowledged. We wish to particularly thank an unknown reviewer who provided useful and detailed sugestions for setting out and presenting the data as well as meticulously checking the species authorities. References Adam, P. (1994) Australian Rainforests. Oxford Biogeography Series No.6. Oxford University Press, Oxford. 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Distribution: showing relationships in Australia and affinities to areas adjacent to Australia. Distributions in other Australian states and territories are available in Streimann & Klazenga (2002). WT= Wet Tropics endemics AU = Australian endemics NS = New species for Australia NR = New records for the Wet Tropics S.E.Asia = species present particularly in parts of Malesia, some in Papua New Guinea. NZ = New Zealand OC = Oceania New Caled. = New Caledonia Location: within the Wet Tropics bioregion The Wet Tropics bioregion is subdivided into a series of provinces (Sattler & Williams 1999). The numbers used here relate to the list of provinces according to Goosem et al. (1999). Province 1. Herbert: Low rainfall, delta of the Herbert River, Quaternary alluvium; alluvial plains with relic stream channels, low stream levees and prior streams associated with the coastal escarpment between the Cardwell Range and Bluewater Creek; open forests and woodlands. Province 2. Tully: High rainfall, Quaternary alluvium; alluvial plains, channels, levees, lagoons and piedmont fans; mesophyll rainforest. Province 3. Innisfail: High rainfall, Quaternary alluvium, some basalt and metasediments; undulating low hills, alluvial plains, channels, levees, lagoons; mesophyll rainforest, fan palm and feather palm rainforests, woodlands. Province 4. Atherton: Moderately high rainfall, 700–1000 m altitude; tableland area dominated by basalt plains, subtropical climate; mainly fragmented mesophyll rainforest. Province 5. Paluma-Seaview: Relatively low rainfall, 800–900+ m elevation; granitic southern ranges, separated from the northern provinces by the Herbert River Gorge; simple notophyll rainforest Province 6. Kirrama-Hinchinbrook: Moderately high rainfall. Steep environmental gradients associated with steep slopes, waterfalls, shallow soils and complex vegetation including simple and complex notophyll rainforest, mesophyll vine forest, tall open forest and woodland. Subdivided for the purposes of this study into: 6a. Cardwell and Kirrama Ranges: Rainfall moderate 6b. Hinchinbrook Island: Rainfall variable, to 3500 mm per annum Province 7. Bellenden Ker-Lamb Range: Very high rainfall, high altitudes to 1600+ m; wet and cloudy upland granitic massifs; mostly rainforest; a major centre of endemism in the bioregion. Province 8. Macalister (north and west of Cairns): Moderate rainfall; undulating tableland bounded by a steep dissected escarpment falling to a narrow coastal plain; includes the Black Mountain corridor; vegetation mixed. Province 9. Daintree–Bloomfield: Variable rainfall; a complex province, likely to be subdivided in the future- includes Mt Carbine, Windsor and Big Tablelands, Mount Finnigan, Thornton Peak, Mt Lewis. Species Location Distribution Acanthorrhynchium papillatum (Harv.) M.Fleisch. 1, 2, 6b S.E.Asia Achrophyllum dentatum (Hook. f. & Wilson) Vitt & Crosby 4 S.E.Asia, PNG, NZ Acroporium lamprophyllum Mitt. var. percaudatum (E.B.Bartram) B.C.Tan, H.P.Ramsay & W.B.Schofield 2, 5, 6a, 9 S.E. Asia, NS Acroporium microcladon (Dozy & Molk.) B.C.Tan var. rhizogemmae B.C.Tan, W.B.Schofield & H.P.Ramsay (includes specimens identified as C. conspicuum M. Fleisch.) 2, 4, 5, 6a, 7, 9 S.E.Asia NS Acroporium stramineum (Reinw. & Hornsch.) M.Fleisch. = A. erythropodium (Hampe) Broth. (AU); Rhynchostegium erythropodium (Hampe) Mitt.; Sematophyllum erythropodium (Hampe) A. Jaeger 4, 5 6a, 7, 9 S.E.Asia 398 Cunninghamia 8(3): 2004 Acroporium strepsiphyllum (Mont.) B.C.Tan Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland 5, 6a, 9 S.E.Asia, NS Aerobryopsis longissima (Dozy & Molk.) M.Fleisch. 1, 2, 3, 4, 5, 6a, 7, 9 S.E. Asia Anoectangium aestivum (Hedw.) Mitt. 3 Asia Archidium brevinerve P. de la Varde 2 Equatorial Africa Archidium capense Hornsch. 2 Southern Africa Archidium clarksonianum I.G.Stone 5 AU Archidium elatum Dixon & Sainsbury 2, 6 NR Archidium microthecium Dixon & P. de la Varde 2 Southern Africa & India Archidium minutissum I.G. Stone 9 WT Archidium ohioense Schimp. ex Müll.Hal. 4 Cosmopolitan, NS Archidium rothii Watts ex G.Roth 1, 2, 3, 8, 9 AU Archidium species A 2 AU Archidium species B 3 AU Archidium species C 2, 3, 8, 9 AU Arthrocormus schimperi (Dozy & Molk.) Dozy & Molk. 2, 3, 7, 9 S.E.Asia Barbellopsis trichophora (Mont.) W.R Buck 1, 2, 3, 4, 5, 6a, 7, 8, 9 S.E.Asia Barbula subcalycina Mull Hal. 4 AU Bartramia mossmaniana Müll.Hal. 7 South America Bescherellia elegantissima Duby 2, 4, 5, 6a, 9 New Caled. Brachymenium nepalense Hook. 2, 3, 4, 5 6a, 7, 9 Paleotrop. S.E. Asia, OC Brachythecium salebrosum (F.Weber & D.Mohr) Schimp. 4 Southern Africa, NZ, Braithwaitea sulcata (Hook.) A.Jaeger & Sauerb. 4 NZ Breutelia affinis (Hook.) Mitt. 6a NZ Bryobrothera crenulata (Broth. & Paris) Thér. 5, 6a, 6b, 7, 9 S.E.Asia, Melanesia Bryum argenteum Hedw. 1, 2, 3, 5, 6a, 7, 9 S.E.Asia Bryum auratum Mitt. Syn: Anomobryum auratum 4, 5 S.E.Asia Bryum lanatum (P.Beauv.) Brid. Syn: Anomobryum lanatum 1, 2, 3,4, 5, 6a, 7, 9 S.E.Asia Buxbaumia colyerae Burges 7 AU Buxbaumia thorsorneae I. G. Stone 7 WT Caduciella mariei (Besch.) Enroth 9 S.E.Asia Callicostella papillata (Mont.) Mitt. var. papillata 9 S.E.Asia Callicostella papillata (Mont.) Mitt. var. prabaktiana (Müll.Hal.) Streimann 3, 8, 9 S.E.Asia Calymperes afzelii Sw. 1, 2, 3, 5, 6a, 9 S.E.Asia, Pantropical Calymperes boulayi Besch. 8 NR, S.E.Asia Calymperes couguiense Besch. 2, 3, 7, 9 S.E.Asia Calymperes crassinerve (Mitt.) A. Jaeger 2, 3, 9 S.E.Asia Calymperes erosum Müll.Hal. 1, 3, 4, 6b, 7, 9 S.E.Asia, Pantropical Calymperes graeffeanum Müll.Hal. 1, 2, 3, 4, 6a, 6b, 7, 8, 9 S.E.Asia Calymperes lonchophyllum Schwägr. 2, 9 S.E.Asia Calymperes moluccense Schwägr. 2, 3, 7, 9 S.E.Asia Calymperes motleyi Mitt. 1, 2, 5, 9 S.E.Asia Calymperes porrectum Mitt. 9 S.E.Asia Calymperes serratum A. Braun ex Müll.Hal. 3, 4, 7, 9 S.E.Asia Calymperes strictifolium (Mitt.) G. Roth 3 S.E.Asia Calymperes subintegrum Broth. 2, 6b, 7, 9 S.E.Asia Calymperes taitense (Sull.) Mitt. 2, 3, 5, 6b, 9 S.E.Asia Calymperes tenerum Müll.Hal. 1, 2, 3, 4, 5, 7, 9 S.E.Asia Calyptothecium acutum (Mitt.) Broth. 1, 2 AU Calyptothecium australinum (Mitt.) Paris 4 AU, NS Calyptothecium humile (Mitt.) Broth. 1, 9 South America Calyptothecium recurvulum (Broth.) Broth. 7 Melanesia Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland 399 Calyptothecium subecostatum Dixon 3 WT Calyptrochaeta apiculata (Hook. f. & Wilson) Vitt 7 S. America, NZ, Antarctic Islands Calyptrochaeta brassii (E. B. Bartram) Streimann 4, 9 WT Calyptrochaeta rotundifolia (Nog. & Z. Iwats.) Touw 7 S.E.Asia, NR Camptochaete curvata Tangney 4 (rare) AU Camptochaete deflexa (Wilson) A. Jaeger 4, 9 NZ Camptochaete excavata (Taylor) A. Jaeger 2, 3, 4, 5, 6b, 7, 9 AU Campylopus catarractilis (Müll.Hal.) Paris 1, 2, 4, 5, 6a Southern Africa, NZ Campylopus clemensiae E.B. Bartram 4 S.E.Asia Campylopus comosus (Schwägr.) Bosch & Sande Lac. 3, 4, 5, 8 S.E.Asia Campylopus ericoides (Griff.) A. Jaeger 8 S.E.Asia Campylopus flexuosus (Hedw.) Brid. 2, 3, 4, 7, 9 S.E.Asia, NZ -India Campylopus flindersii Catches. & J.-P. Frahm 6a AU Campylopus introflexus (Hedw.) Brid. 4, 7, 9 Southern Hemisphere Campylopus laxitextus Sande Lac. 2, 4, 9 S.E.Asia, New Caled NZ, subtropics, N.Caled. Campylopus pyriformis (Schultz) Brid. 4, 7 Campylopus robillardei Besch. 4, 6a, 7, 9 Tropical Africa Campylopus sinensis (Müll.Hal.) J.-P. Frahm 3, 4, 9 S.E.Asia Campylopus umbellatus (Schwägr. & Gaudichaud ex Arn.) Paris 1, 4, 5, 6a, 7, 9 S.E.Asia Chaetomitrium tahitense (Sull.) Mitt. 3, 4, 9 S.E.Asia Claopodium assurgens (Sull. & Lesq.) Cardot 1, 3 S.E.Asia Clastobryum dimorphum (I. G. Stone) B. C. Tan, Z. Iwats. & D. H. Norris 7, 9 AU Clastobryum epiphyllum (Renauld & Cardot) B. C. Tan &Touw 5, 6a S.E.Asia Cryphaea tenella (Schwägr.) Hornsch. ex Müll.Hal. 4 Tropical Pacific, NZ Cryptogonium phyllogonioides (Sull.) Isov. 3 S.E.Asia Cyathophorum bulbosum (Hedw.) Müll.Hal. 7 NZ. Cyclodictyon blumeanum (Müll.Hal.) O.Kuntze 4, 7, 9 S.E.Asia , Pacific Cyptodon muelleri (Hampe) M. Fleisch. 3, 4, 8 AU Daltonia contorta Müll.Hal. 4, 7 S.E.Asia Daltonia splachnoides (Sm.) Hook. & Taylor 7 NZ Dawsonia longiseta Hampe? 4, 9 AU Dawsonia polytrichoides R. Br. 4, 5, 9 AU Dawsonia superba Grev. var. pulchra Zanten 4, 9 AU Dicnemon calycinum (Hook.) Schwägr. 8 NZ Dicranella dietrichiae (Müll.Hal.) A. Jaeger 4, 9 NZ Dicranella euryphylla Dixon 4 WT Dicranella pycnoglossa (Broth.) Kindb. 3, 4, 7, 8, 9 AU Dicranoloma austroscoparium (Müll.Hal ex Broth.) Watts & Whitel. 4, 7, 9 WT Dicranoloma braunii (Müll.Hal. Ex Bosch & Sande Lac.) Paris 9 S.E.Asia Dicranoloma daymannianum E.B.Bartram 4, 5, 6a, 7 S.E.Asia Dicranoloma dicarpum (Nees) Paris 4 S.E.Asia, NZ Dicranoloma leichhardtii (Hampe) Watts & Whitel. 4, 5 AU Dicranoloma menziesii (Taylor) Renauld 4, 7, 9 NZ, New Caled. Dicranoloma robustum (Hook.f. & Wilson) Paris 7 NZ, sub-Antarctic Islands Dicranoloma wattsii Broth. 4 AU Diphyscium mucronifolium Mitt. 7, 9 S.E.Asia, NR Distichophyllum crispulum (Hook. f. & Wilson) Mitt. 4, 5, 7, 9 NZ Distichophyllum cuspidatum (Dozy & Molk.) Dozy & Molk. 3 S.E.Asia Distichophyllum mittenii Bosch & Sande Lac. 4, 5, 6a S.E.Asia Ditrichum difficile (Duby) M. Fleisch. 2, 4, 5, 7 S.E.Asia, NZ Eccremidium brisbanicum (Broth.) I. G. Stone & G. A. M. Scott 2 S.E.Asia, New Caled. Eccremidium minutum (Mitt.) I. G. Stone & G. A. M. Scott 3 NZ, New Caled. Eccremidium pulchellum (Hook. & Wilson) Müll.Hal. 2 NZ Ectropothecium moritzii A. Jaeger 7, 8 S.E.Asia 400 Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Ectropothecium riparioides E. B. Bartram 4 AU OC Ectropothecium umbilicatum var. umbilicatum (Müll.Hal.) Paris 4,7, 8 Ectropothecium zollingeri (Müll.Hal.) A.Jaeger 4, 7,8 S.E.Asia Entodon mackaviensis Müll.Hal. 4, 5 AU Entodon plicatus Müll.Hal. 4, 6a, 9 S.E.Asia Ephemeropsis tjibodensis K. J. Goebel 2, 4, 7, 9 S.E.Asia, NZ Ephemerum fimbriatum Müll.Hal. 2, 6a New Caled. Erpodium coronatum (Hook. f. & Wilson) Mitt. var. australiense (I G. Stone) I.G. Stone 5 AU Erpodium hodgkinsoniae (Hampe & Müll.Hal.) 1 Norfolk Island Erpodium solmsiellaceum (Müll.Hal. & Broth.) I.G.Stone 2, 3 New Caled. Eucamptodon muelleri var. muelleri Hampe & Müll.Hal. 2, 4, 5, 6a, 6b, 7, 8, 9 AU Eucamptodon scalarirete (Dixon) B.C. Tan, H.P. Ramsay & W.B. Schofield 4, 5, 7, 9 AU Euptychium setigerum (Sull.) Broth. subsp. setigerum 3 S.E.Asia Eurhynchium laevisetum Geh. 4 AU Exostratum blumei (Nees ex Hampe) L.T. Ellis 2, 7, 9 S.E.Asia Fabronia australis Hook. 4, 9 NZ Fallaciella gracilis (Hook. f. & Wilson) H.A.Crum 3, 4 NZ, S.E.Asia, Fissidens altisetus Dix. [= F. bogoriensis according to Streimann & Klazenga (2002) but F. bogoriensis not known in Australia (pers. comm. I. Stone)] 9 AU ? Fissidens asplenioides Hedw. 4 Asia Fissidens autoicus Thér. & Broth. 4, 7 S.E.Asia Fissidens badyinbarus I. G. Stone & Catches. 6b Fissidens bryoides Hedw. var. schmidii (Mull. Hal.) R.S. Chopra & S.S. Kumar 4 WT Asia Fissidens cambewarrae Dixon 2 AU Fissidens ceylonensis Dozy & Molk. 7 S.E.Asia, NZ Fissidens crassinervis Sande Lac. 9 Asia Fissidens crenulatus Mitt. 1, 6a Indo-Burmese Fissidens crispulus Brid. 1, 3, 4, 6a, 7, 9 Southern Africa, Japan Fissidens curvatus Hornsch. 4 S.E.Asia Fissidens dietrichiae Müll.Hal. 4 New Caled. Fissidens flabellulus Thwaites & Mitt. 2, 3, 4, 6a S.E.Asia, New Caled. Fissidens flabellulus Thwaites & Mitt. var. eachamensis I. G. Stone 4 WT Fissidens gardneri Mitt. 1, 9 S.E.Asia Fissidens gymnocarpus I. G. Stone 9 AU Fissidens henryae I. G. Stone 2 WT Fissidens hollianus Dozy & Molk. 9 S.E.Asia Fissidens holstii Broth. 8 S.E.Asia Fissidens hyalinus Hook. & Wilson 4 S.E.Asia Fissidens leptocladus Müll. Hall ex Rodway 2 NZ New Caled. Fissidens linearis Brid. var. obscurirete (Broth.) I. G. Stone 9 Fissidens oblatus I. G. Stone & Catches. 4 WT Fissidens oblongifolius Hook. f. & Wilson 4 WT Fissidens oblongifolius Hook. f. & Wilson var. hyophilus (Mitt.) Beever & I. G. Stone 2, 4, 7 AU Fissidens pallidus Hook. f. & Wilson var. pallidus 4 NZ Fissidens patulifolius Dixon 2 AU Fissidens pellucidus Hornsch. 1, 4 Asia Fissidens perobtusus Dixon (on termite mounds) 2, 9 AU Fissidens polypodioides Hedw. 5 NR Fissidens pseudopallidus I. G. Stone 8 WT Fissidens punctulatus Sande Lac. 6a, 9 S.E.Asia Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Fissidens rupicola Paris & Broth. 4, 6a, 9 OC Fissidens serratus Müll.Hal. 9 S.E.Asia, New Caled. Fissidens sufflatus I. G. Stone 7 WT Fissidens tenellus Hook. f. & Wilson var. australiensis (A.Jaeger) Beever & I.G.Stone 4 AU, NR Fissidens zollingeri Mont. 2, 3 S.E.Asia, OC Floribundaria floribunda (Dozy & Molk.) M. Fleisch. 2, 4, 7, 9 S.E.Asia Floribundaria pseudofloribunda M. Fleisch. 4, 9 S.E.Asia Floribundaria walkeri (Renauld & Cardot) Broth. 4 Southern Africa, S.E.Asia Funaria hygrometrica Hedw. 5, 6a, 9 Cosmopolitan Garckea flexuosa (Griff.) Margad. & Nork. 7, 9, 8 S.E.Asia Garovaglia elegans (Dozy & Molk.) Bosch & Sande Lac. subsp. dietrichiae (Müll.Hal.) During 1, 3, 4, 5, 6a, 7, 8, 9 S.E.Asia, OC Gemmabryum acuminatum (Harv. In Hook.) J.R. Spence and H.P. Ramsay 4, Pantropical, subtropical Gemmabryum apiculatum (Schwägr.)J.R. Spence & H.P. Ramsay = Bryum apiculatum Schwägr. 1, 2, 3, 4, 6a, 7, 8, 9 S.E.Asia, OC, N.Z. Gemmabryum australe (Hampe) J.R. Spence & H.P. Ramsay = Bryum australe Hampe Gemmabryum chrysoneuron (M üll Hal.) J.R. Spence & H.P. Ramsay = Bryum chrysoneuron Müll Hal. Gemmabryum clavatum (Schimp.) J.R. Spence & H.P. Ramsay = Bryum clavatum (Schimp.) Müll.Hal. 1,9 NZ, S.E.Asia 3,4,6,7 New Caled., Oc, NZ 4, 7, 8 S.E.Asia, N.Z., OC Gemmabryum coronatum ( Schwägr.) J.R. Spence & H.P. Ramsay = Bryum coronatum Schwägr.; = B. subatropurpureum Müll.Hal. 1, 4, 5 New Caled., NZ Gemmabryum dichotomum (Hedw.) J.R. Spence & H.P. Ramsay = Bryum dichotomum Hedw; = B. pimpamae Müll.Hal. 4, 6a, 8 NZ, Subantarctic Gemmabryum exile (Dozy & Molk.) J.R. Spence & H.P. Ramsay 4,7, 9 S.E.Asia, OC Gemmabryum indicum (Dozy & Molk.) J.R. Spence & H.P. Ramsay 2, 3, 8, 9 S.E.Asia, OC Gemmabryum pachythecum (Müll.Hal.) J.R. Spence & H.P. Ramsay =Bryum pachytheca Müll.Hal. 9 S.E.Asia, NZ Gemmabryum preissianum (Hampe) J.R.Spence & H.P. Ramsay 2, 6, 9 AU Gigaspermum repens (Hook.) Lindb. 4 Southern Africa, NZ ?Glossadelphus hermaphroditus M. Fleisch. [=Chaetomitrium entodontoides Broth. & Watts Glossadelphus = Phyllodon (see Buck 1987) 3 S.E.Asia Grimmia pulvinata (Hedw.) Sm var. africana (Hedw.) Hook. f. & Wilson 8 NZ Groutiella tomentosa (Hornsch.) Wijk & Margad. 4, 6a, 9 S.E.Asia Hampeella concavifolia Hattaway & Norris (in prep.) 7, 9 WT, NR Hampeella pallens (Sande Lac.) M. Fleisch. 4, 5, 6b, 7, 9 S.E.Asia Herpetineuron toccoae (Sull. & Lesq.) Cardot 4, 7 Cosmopolitan Himantocladium cyclophyllum (Müll.Hal.) M. Fleisch. 7, 9 S.E.Asia Holomitrium perichaetiale (Hook.) Brid. 1, 2, 3, 4, 5, 6a, 7, 8, 9 NZ Homaliodendron exiguum (Bosch & Sande Lac.) M. Fleisch. 1, 2, 4, 9 S.E.Asia Homaliodendron flabellatum (Sm.) M. Fleisch. 4, 9 S.E.Asia Hookeriopsis utacammundiana (Mont.) Broth. 5, 7, 9 S.E.Asia Hyophila involuta (Hook.) A. Jaeger 2, 3, 4, 9 S.E.Asia Hypnodendron comatulum (Geh. ex Broth.) Touw 2, 3, 4, 5, 6a, 7 WT Hypnodendron spininervium (Hook.) A.Jaeger & Sauerb. subsp. archeri (Mitt.) Touw 4, 7, 8 AU Hypnodendron vitiense Mitt. subsp. australe Touw 4, 5, 9 AU Hypnodendron vitiense Mitt. subsp. vitiense 4, 7 S.E.Asia, OC Hypnum cupressiforme Hedw. 2, 3, 4, 5, 8, 9 Cosmopolitan Hypnum subchrysogaster (Broth.) Paris 7 AU Hypopterygium tamarisci (Sw.) Brid. ex Müll.Hal. 4, 5, 6b NZ Isocladiella wattsii (Broth.) B.C. Tan, H.P. Ramsay & W.B. Schofield 7 WT 401 402 Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Isopterygium acuminatum Bosw. 4, 7, 8 AU, NR Isopterygium albescens (Hook.) A. Jaeger 2, 4 S.E.Asia, OC, NZ Isopterygium minutirameum (Müll.Hal.) A. Jaeger var. brevifolium (M.Fleisch.) E.B.Bartram 7 S.E.Asia, OC Isopterygium minutirameum (Müll.Hal.) A. Jaeger var. minutirameum 2,3,6b,9 NZ Isopterygium novae-valesiae Broth. 2,4,7 AU Lembophyllum divulsum (Hook. f. & Wilson) var. clandestinum (Hook. f. & Wilson) Wijk & Margad. 7, 9 NZ Leptobryum pyriforme (Hedw.) Wilson 5 NZ Leptodictyum riparium (Hedw.) Warnst. 4 NR Leptostomum erectum R. Br. 1, 2, 4 AU Leptotrichella tenax (Müll.Hal.) Ochyra var. longipes (Müll.Hal.) Ochyra 9 AU Leucobryum aduncum Dozy & Molk. var. aduncum. 2, 3, 4, 6a, 8, 9 S.E.Asia Leucobryum aduncum Dozy & Molk. var. scalare (Müll.Hal.ex M.Fleisch) A.Eddy 1, 2, 4, 6a, 9 S.E.Asia, New Caled. Leucobryum ballinense Broth. 3, 4, 5 AU Leucobryum candidum (Brid. ex P. Beauv.) Wilson 2, 3, 4, 5, 6a, 6b, 7, 8, 9 Asia, New Caled., NZ, Leucobryum chlorophyllosum Müll.Hal. 1, 4, 9 S.E.Asia, New Caled. NR Leucobryum sanctum (Brid.) Hampe 2, 3, 4, 6a, 7, 8, 9 S.E.Asia Leucobryum subchlorophyllosum Hampe 4, 6a, 7, 9 NR Leucobryum wattsii Broth. 1, 2, 6a, 6b, 9 AU Leucoloma [various undetermined species] 4, 6a, 7, 9 WT Leucoloma molle (Müll.Hal.) Mitt. 4, 7 AU Leucomium strumosum (Hornsch.) Mitt. 3, 4 S. Amererica, NR Leucophanes angustifolium Renauld & Cardot 1, 3, 5, 7 Madagascar Leucophanes candidum (Schwägr.) Lindb. 9 S.E.Asia, OC Leucophanes glaucum (Schwägr.) Mitt. 2, 3, 7, 8, 9 S.E.Asia, OC Leucophanes octoblepharoides Brid. 3, 4, 6b, 7, 8, 9 S.E.Asia, OC Lopidium concinnum (Hook.) Wilson 2, 6b, 7, 9 South America, NZ Lopidium struthiopteris (Brid.) M. Fleisch. 4, 5, 7 S.E.Asia, OC Macgregorella indica (Broth.) W.R. Buck 4 S.E.Asia Macrohymenium mitratum (Dozy & Molk.) M. Fleisch. 6b S.E.Asia Macromitrium archeri Mitt. 4, 7, 9 AU Macromitrium aurescens Hampe 1, 4, 6a AU Macromitrium caloblastoides Müll.Hal. 1, 2, 4 AU Macromitrium diaphanum Müll.Hal. 4 AU Macromitrium dielsii Broth. ex Vitt & H.P. Ramsay 7 WT Macromitrium exsertum Broth. 4, 7, 9 AU Macromitrium funiforme Dixon 7, 9 WT Macromitrium hemitrichodes Schwägr. 4, 9 AU Macromitrium hortoniae Vitt & H.P. Ramsay 5 AU, NR Macromitrium incurvifolium (Hook. & Grev.) Schwägr. 3, 9 OC Macromitrium involutifolium (Hook. & Grev.) Schwägr. subsp. involutifolium 4, 5, 7 New Caled. 4, 5, 6a, 7, 9 OC Macromitrium involutifolium (Hook. & Grev.) Schwägr. subsp. ptychomitrioides (Besch.) Vitt & H.P. Ramsay Macromitrium leratii Broth. & Paris 4, 6a, 6b, 7, 9 AU Macromitrium ligulaefolium Broth. 2, 3, 4, 5, 6a, 7, 8, 9 NZ, New Caled., OC. Macromitrium microstomum (Hook. & Grev.) Schwägr. 1, 2, 3, 4, 7, 9 NZ, OC Macromitrium repandum Müll.Hal. 1, 4, 5, 6a, 7, 8, 9 AU Macromitrium stoneae Vitt & H.P. Ramsay 4 AU Meiotheciella papillosa (Broth.) B.C. Tan, H.P. Ramsay & W.B. Schofield 5 S.E.Asia, New Caled. NS Meiothecium secundifolium Dixon 2 WT, NS Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Meiothecium microcarpum (Hook.) Mitt. = Meiothecium wattsii (Broth.) Broth. 2, 6b, 9 [Australian specimens of Meiothecium jagorii (Müll.Hal.) Broth are this, earlier missidentified] 403 S.E.Asia, New Caled., Oc. Meiothecium tenellum Broth. & Paris 3, 4, 8 S.E.Asia Mesochaete taxiforme (Hampe) Watts & Whitel. 4, 5, 9 AU Mesochaete undulata Lindb. 1, 5 AU Mesonodon flavescens (Hook.) W.R. Buck 4 S.E.Asia Meteoriopsis reclinata (Müll.Hal.) M. Fleisch. ex Broth. 3, 4, 7, 8, 9 S.E.Asia Meteorium polytrichum Dozy & Molk 3, 4, 5, 6a, 7, 9 S.E.Asia Mittenia plumula (Mitt.) Lindb. 5 NZ, PNG, NR Mitthyridium constrictum (Sull.) H.Rob. 3, 7, 9 S.E.Asia Mitthyridium crassum (Broth.) H.Rob. 2, 9 S.E.Asia Mitthyridium fasciculatum (Hook. & Grev.) H.Rob. 2, 3, 4, 7, 9 S.E.Asia Mitthyridium flavum (Müll.Hal.) H.Rob. 1, 2, 3, 6a, 7, 8, 9 S.E.Asia Mitthyridium leucoloma (Müll.Hal.) H.Rob. 3 S.E.Asia Mitthyridium papuanum (Broth.) H.Rob. 2, 3, 7, 9 S.E.Asia Mitthyridium perundulatum (Broth.) H.Rob. 3, 7, 9 S.E.Asia S.E.Asia Mitthyridium repens (Harv.) H.Rob. 2, 3, 7, 9 Mitthyridium subluteum (Müll.Hal.) H.K. Nowak 1, 2, 3, 5, 9 S.E.Asia Muellerobryum whiteleggei (Broth.) M. Fleisch. 1, 2, 3, 4, 5, 6b, 7, 9 AU Myurium rufescens (Reinw. & Hornsch.) M. Fleisch. subsp. purpuratum (Mitt.) Maschke [Oedicladium — no combination available] 4, 7, 9 S.E.Asia, Oc Nanobryum thorsbornei I. G. Stone = Fissidens thorsbornei (I. G. Stone) Brugg.-Nann. 3, 5, 6a, 9 New Caled. Neckeropsis lepineana (Mont.) M. Fleisch. 1, 4, 9 S.E.Asia Neckeropsis nanodisticha (Geh.) M. Fleisch. 1, 2 S.E.Asia . Neolindbergia vitiensis (E.B.Bartram) Enroth 9 S.E.Asia Notoligotrichum australe (Hook. f. & Wilson) G.L.Sm. 4 NZ Octoblepharum albidum Hedw. 1, 2, 4, 6a, 6b, 9 S.E.Asia Oedicladium rufescens (Reinw. & Hornsch.) M. Fleisch. subsp. rufescens [see Myurium also] 4, 6b, 9 S.E.Asia, OC Orthomnion elimbatum (Nog.) T. J. Kop. 3, 4, 9 PNG Orthorrhynchium elegans (Hook. f. & Wilson) Reichhardt subsp. cymbifolioides (Müll.Hal.) Lin. 7 S.E.Asia, NZ S.E.Asia, NZ Papillaria crocea (Hampe) A. Jaeger 4, 5, 6a, 8, 9 Papillaria flexicaulis (Wilson) A. Jaeger 4, 5 S.E.Asia, New Caled. Papillaria leuconeura (Müll.Hal.) A. Jaeger 6a, 9 S.E.Asia Papillaria nitens (Hook. f. & Wilson) Sainsbury 3, 4, 5, 6a, 6b, 7, 9 NZ, New Cal. Pelekium gratum (P.Beauv.) Touw 1, 6b, 7 S.E.Asia Pelekium investe (Mitt.) Touw 4 S.E.Asia, NR Pelekium synoicum (Touw) Touw 4,7,8 S.E.Asia Philonotis hastata (Duby) Wijk & Margad. 3, 4, 8, 9 S.E.Asia, NZ AU [Philonotis pseudomollis (Müll.Hal.) A. Jaeger (? P. tenuis)] 4, 7 Philonotis tenuis (Taylor) Reichardt 4, 7 NZ Philonotis thwaitesii Mitt. 9 Asia, S.E.Asia, S.America, NR Pinnatella alopecuroides (Mitt.) M. Fleisch. 7, 9 S.E.Asia Pinnatella kuehliana (Bosch & Sande Lac.) M. Fleisch. 2, 4, 6a S.E.Asia, OC Platyhypnidium austrinum (Hook. f. & Wilson) M. Fleisch. 7 NZ Platyhypnidium muelleri (A. Jaeger) M. Fleisch. 7 PNG, Hawaii, NR Pogonatum neesii (Müll.Hal.) Dozy 4 S.E.Asia, OC Pogonatum tubulosum Dixon 9 S.E.Asia Polytrichum juniperinum Hedw. 4 S.E.Asia Powellia breviseta (E. B. Bartram) Zanten 1, 2 (in Flora) New Caled., OC, NR Powellia involutifolia Mitt. 2, 5, 7 S.E.Asia, OC 404 Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Pseudohypnella verrucosa (Dozy & Molk.) M. Fleisch. 3, 7 S.E.Asia Pseudospiridentopsis horrida (Mitt. ex Cardot) M. Fleisch. 7 WT Pseudosymblepharis bombayensis (Müll.Hal.) P. Sollman 9 Asia, S.E.Asia, NR Pterobryella breviacuminata Besch. 4, 9 OC Pterobryidium australe Broth. & Watts 4 AU Ptychomitrium australe (Hampe) A. Jaeger 4 NZ Ptychomnion aciculare (Brid.) Mitt. 7 NZ, NR Pyrrhobryum latifolium (Bosch. & Sande Lac.) Mitt. 4, 5, 6a, 9 S.E. Asia Pyrrhobryum medium (Besch.) Manuel 4, 5, 7 S.E.Asia, OC Pyrrhobryum paramattense (Müll.Hal.) Manuel ?doubtful species (Frahm 2003) 1, 3, 4, 6b, 7, 9 S.E.Asia, NZ Pyrrhobryum spiniforme (Hedw.) Mitt. 1, 4, 6a, 7, 9 S.E.Asia Racopilum cuspidigerum (Schwägr.) Ångstr. var. convolutaceum (Müll.Hal.) Zanten & Dijkstra 3, 7 OC, NZ S.E.Asia, OC Racopilum cuspidigerum (Schwägr.) Ångstr. var. cuspidigerum 7 Radulina hamata (Dozy & Molk.) W.R.Buck & B.C. Tan 1, 3, 5, 8, 9 S.E.Asia, OC Rhaphidorrhynchium amoenum (Hedw.) M. Fleisch. var. amoenum 9 NZ Rhizogonium graeffeanum (Müll.Hal.) A. Jaeger 4, 7, 9 S.E.Asia Rhodobryum aubertii (Schwägr.) Thér. 4, 6a, 7, 9 S.E.Asia Rhynchostegium distratum (Hampe) A. Jaeger 4 AU Rhynchostegium nanopennatum (Broth.) Kindb. 7 WT Rhynchostegium tenuifolium (Hedw.) Reichhardt var. tenuifolium 4 S.E.Asia, OC, NZ Rosulabryum albolimbatum (Hampe) J.R. Spence 2, 4, AU Rosulabryum billarderi (Schwägr.) J.R. Spence 1, 4, 5, 6a, 7, 9 OC Rosulabryum capillare (Hedw.) J.R. Spence 9 Cosmopolitan Rosulabryum epiphyticum J.R. Spence & H.P. Ramsay 2 AU, NS Rosulabryum lamingtonicum J.R. Spence & H.P. Ramsay 3 AU Rosulabryum leptothrix (Mull. Hal.) J.R. Spence 2, 9 AU Rosulabryum subfasciculatum (Hampe) J.R. Spence 1, 2, 3, 4, 5, 6a, 8, 9 New Caled. Rosulabryum subtomentosum (Hampe) J.R. Spence 2, 4 NZ Rosulabryum torquescens (Bruch ex De Not) J.R.Spence 1 Cosmopolitan Rosulabryum tuberosum (Mohamed & Damanhuri) J.R. Spence 3 S.E.Asia Rosulabryum wightii (Mitt.) J.R. Spence 1, 2, 4, 5, 6a, 7, 8, 9 India Schlotheimia brownii Schwägr. 4, 6a, 7, 9 AU Schlotheimia funiformis Taylor ex Dixon 4, 6a AU Schoenobryum concavifolium (Griff.) Gangulee 4 S.E.Asia Sclerodontium clavinerve (Müll.Hal.) H.A.Crum 3, 4 AU Sclerodontium pallidum (Hook.) Schwägr. subsp. pallidum 4, 9 NZ Sematophyllum homomallum (Hampe) Broth. 2 (rare) S.E.Asia, OC Sematophyllum subhumile (Müll.Hal.) M. Fleisch. var. subhumile 2, 4, 5, 6, 7, 8, 9 S.E.Asia, NZ, OC Sematophyllum subhumile (Müll.Hal.) M. Fleisch. var. contiguum (Mitt.) B.C. Tan, W.B. Schofield & H.P. Ramsay 6,7 OC, NZ S.E.Asia, OC Sematophyllum subpinnatum (Brid.) E. Britton 1, 2, 3, 4, 5, 6, 9 Sorapilla papuana Broth. & Geh. 9 S.E.Asia Sphagnum perichaetiale Hampe 3 S.E.Asia, NZ Stereophyllum radiculosum (Hook.) Mitt. 4, 5 AU Syrrhopodon albovaginatus Schwägr. 7, 9 S.E.Asia Syrrhopodon aristifolius Mitt. 2, 9 S.E.Asia S.E.Asia Syrrhopodon armatus Mitt. 2, 3, 4, 6a, 7, 8, 9 Syrrhopodon ciliatus (Hook.) Schwägr. ‘rare’ (Reese & Stone 1995) S.E.Asia, OC Syrrhopodon confertus Sande Lac. 3, 7, 9 S.E.Asia Syrrhopodon croceus Mitt. 3, 7, 9 S.E.Asia Syrrhopodon cyrtacanthos Reese 9 WT Syrrhopodon involutus Schwägr. 2, 3, 6a S.E.Asia Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Syrrhopodon muelleri (Dozy & Molk.) Sande Lac. 3, 4, 7, 8, 9 405 S.E.Asia Syrrhopodon parasiticus (Brid.) Besch. 6a, 7, 9 S.E.Asia Syrrhopodon platycerii Mitt. 2, 4, 9 AU Syrrhopodon prolifer Schwägr. var. mossmanensis Reese 9 WT Syrrhopodon prolifer Schwägr. var. prolifer 9 S.E.Asia WT Syrrhopodon stoneae Reese 6b, 7 Syrrhopodon trachyphyllus Mont. 1, 6a, 6b, 8, 9 S.E.Asia Syrrhopodon tristichus Nees ex Schwägr. 7, 9 S.E.Asia Taxiphyllum taxirameum (Mitt.) M. Fleisch. 4 S.E.Asia, OC Taxithelium instratum (Brid.) Broth. 1,5, 7, 9 S.E.Asia Taxithelium kerianum (Broth.) Broth. 3, 6a, 7, 9 S.E.Asia Taxithelium merillii Broth. 3, 6b, 7, 8, 9 S.E.Asia Taxithelium muscicola (Broth.) B.C.Tan, H.P.Ramsay & W.B.Schofield 2, 7, 9, AU Taxithelium nepalense (Schwägr.) Broth. 4, 5, 7 S.E.Asia, tropical Africa Taxithelium planum (Brid.) Mitt. 4, 5 S.E.Asia, tropical Africa Thamnobryum ellipticum (Bosch & Sande Lac.) W. Schultze-Motel 3 S.E.Asia Thamnobryum pandum (Hook. f. & Wilson) I. G. Stone & G.A.M. Scott 9 NZ Thamnobryum pumilum (Hook.f. & Wilson) Nieuwl. 7 NZ, Thuidiopsis sparsa (Hook. f. & Wilson) Broth. 4, 9 NZ, Thuidium cymbifolium (Dozy & Molk.) Dozy & Molk. 1, 3, 4, 6b, 9 New Caled., OC,NZ Touwia laticostata Ochyra 7 WT Trachycarpidium brisbanicum (Mull. Hal.) I.G. Stone 1, 2 AU Trachyloma diversinerve Hampe 4, 9 NZ Trachyloma indicum Mitt. 4 S.E.Asia NZ Trachyloma planifolium (Hedw.) Brid. 4, 5, 9 Trachyphyllum inflexum (Harv.) A. Gepp. 1, 8 S.E.Asia Trachypus humilis Lindb. 4 S.E.Asia, OC Trachythecium verrucosum (A.Jaeger) M.Fleisch. 4 S.E.Asia, OC Trematodon baileyi Broth. 9 AU Trematodon longescens Müll.Hal. 7, 8 AU Trematodon longicollis Michx. 8 NR Trematodon suberectus Mitt. 4 NZ Trichosteleum boschii (Dozy & Molk.) A. Jaeger 7 S.E.Asia, O Trichosteleum ruficaule (Thwaites & Mitt.) B.C. Tan 2, 4, 7, 9, S.E.Asia, OC Trichosteleum subfalcatulum (Broth. & Watts) B.C. Tan, W.B. Schofield & H.P. Ramsay 1, 2, 4, 6a, AU Trichosteleum wattsii (Paris) B.C. Tan, W.B. Schofield & H.P. Ramsay 2, 6b AU S.E.Asia Trichostomum brachydontium Bruch. 4 Trismegistia rigida (Mitt.) Broth. 9 S.E.Asia Vesicularia rivalis Broth. 5, 9 AU Viridivellus pulchellum I. G. Stone 3, 5, 7 AU Warburgiella leptorhynchoides (Mitt.) M.Fleisch. 7, 9 S.E.Asia Warburgiella leucocytus (Müll.Hal.) B.C. Tan, W.B. Schofield & H.P. Ramsay 1, 4, 7 NZ Weissia balansae (Müll.Hal.) R.H.Zander 1, 2 New Caled. Weissia controversa Hedw. 4 Cosmopolitan Weissia platystegia (Dixon ) A. Eddy = Astomum platystegium reported by Norris & Koponen (1989) 5 S.E.Asia Wijkia extenuata (Brid.) H.A.Crum 1, 2, 4, 6a 7, 9 New Caled. NZ Wilsoniella karsteniana Müll.Hal. 8 WT Zygodon intermedius Bruch & Schimp. 4 NZ Note: For one taxon, Gemmabryum, we are using the name to be published in the Flora of Australia treatment in volume 51 (in press 2005). 406 Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Appendix 2. Alphabetic list of genera with families Acanthorrhynchium Achrophyllum Acroporium Aerobryopsis Anoectangium Archidium Arthrocormus Barbellopsis Barbula Bartramia Bescherellia Brachymenium Brachythecium Braithwaitea Breutelia Bryobrothera Bryum Buxbaumia Caduciella Callicostella Calymperes Calyptothecium Calyptrochaeta Camptochaete Campylopus Chaetomitrium Claopodium Clastobryum Cryphaea Cryptogonium Cyathophorum Cyclodictyon Cyptodon Daltonia Dawsonia Dicnemon Dicranella Dicranoloma Diphyscium Distichophyllum Ditrichum Eccremidium Ectropothecium Entodon Ephemeropsis Ephemerum Erpodium Eucamptodon Euptychium Eurhynchium Fabronia Fallaciella Fissidens Floribundaria Funaria Garckea Garovaglia Gemmabryum* Gigaspermum Glossadelphus Grimmia Groutiella Hampeella Herpetineuron Himantocladium Holomitrium Homaliodendron Sematophyllaceae Hookeriaceae Sematophyllaceae Meteoriaceae Pottiaceae Archidiaceae Calymperaceae Meteoriaceae Pottiaceae Bartramiaceae Cyrtopodaceae Bryaceae Brachytheciaceae Trachylomataceae Bartramiaceae Adelotheciaceae Bryaceae Buxbaumiaceae Neckeraceae Pilotrichaceae Calymperaceae Pterobryaceae Daltoniaceae Lembophyllaceae Dicranaceae Symphyodontaceae Leskeaceae Sematophyllaceae Cryphaeaceae Pterobryaceae Hookeriaceae Pilotrichaceae Cryphaeaceae Daltoniaceae Polytrichaceae Dicnemonaceae Dicranaceae Dicranaceae Diphysciaceae Daltoniaceae Ditrichaceae Ditrichaceae Hypnaceae Entodontaceae Daltoniaceae Ephemeraceae Erpodiaceae Dicnemonaceae Garovagliaceae Brachytheciaceae Fabroniaceae Lembophyllaceae Fissidentaceae Meteoriaceae Funariaceae Ditrichaceae Garovagliaceae Bryaceae Gigaspermaceae Hypnaceae Grimmiaceae Orthotrichaceae Ptychomniaceae Anomodontaceae Neckeraceae Dicranaceae Neckeraceae Hookeriopsis Hyophila Hypnodendron Hypnum Hypopterygium Isocladiella Isopterygium Lembophyllum Leptobryum Leptodictyum Leptostomum Leptotrichella Leucobryum Leucoloma Leucomium Leucophanes Lopidium Macgregorella Macrohymenium Macromitrium Meiotheciella Meiothecium Mesochaete Mesonodon Meteoriopsis Meteorium Mittenia Mitthyridium Muellerobryum Myurium Nanobryum Neckeropsis Neolindbergia Notologotrichum Octoblepharum Oedicladium Orthomnion Orthorrhynchium Papillaria Pelekium Philonotis Pinnatella Platyhypnidium Pogonatum Polytrichum Powellia Pseudohypnella Pseudospiridentopsis Pseudosymblepharis Pterobryella Pterobryidium Ptychomitrium Ptychomnion Pyrrhobryum Racopilum Radulina Rhaphidorrhynchium Rhizogonium Rhodobryum Rhynchostegium Rosulabryum Schlotheimia Schoenobryum Sclerodontium Sematophyllum Sorapilla Sphagnum Stereophyllum Syrrhopodon Pilotrichaceae Pottiaceae Hypnodendraceae Hypnaceae Hypopterygiaceae Sematophyllaceae Hypnaceae Lembophyllaceae Meesiaceae Amblystegiaceae Leptostomataceae Dicranaceae Leucobryaceae Dicranaceae Leucomiaceae Calymperaceae Hypopterygiaceae Myriniaceae Sematophyllaceae Orthotrichaceae Sematophyllaceae Sematophyllaceae Rhizogoniaceae Entodontaceae Meteoricaeae Meteoriaceae Mitteniaceae Calymperaceae Pterobryaceae Myuriaceae Nanobryaceae Neckeraceae Pterobryaceae Polytrichaceae Calymperaceae Myuriaceae Mniaceae Orthorrhynchiaceae Meteoriaceae Thuidiaceae Bartramiaceae Neckeraceae Brachytheciaceae Polytrichaceae Polytrichaceae Racopilaceae Sematophyllaceae Meteoriaceae Pottiaceae Pterobryellaceae Pterobryaceae Ptychomitriaceae Ptychomniaceae Rhizogoniaceae Racopilaceae Sematophyllaceae Sematophyllaceae Rhizogoniaceae Bryaceae Brachytheciaceae Bryaceae Orthotrichaceae Cryphaeaceae Dicranaceae Sematophyllaceae Sorapillaceae Sphagnaceae Stereophyllaceae Calymperaceae Cunninghamia 8(3): 2004 Taxiphyllum Taxithelium Thamnobryum Thuidiopsis Thuidium Touwia Trachycarpidium Trachyloma Trachyphyllum Trachypus Trachythecium Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Hypnaceae Sematophyllaceae Neckeraceae Thuidiaceae Thuidiaceae Neckeraceae Pottiaceae Trachylomataceae Pterigynandraceae Meteoriaceae Hypnaceae Trematodon Trichosteleum Trichostomum Trismegistia Vescicularia Viridivellus Warburgiella Weissia Wijkia Wilsoniella Zygodon Bruchiaceae Sematophyllaceae Pottiaceae Sematophyllaceae Hypnaceae Viridivelleraceae Sematophyllaceae Pottiaceae Sematophyllaceae Ditrichaceae Orthotrichaceae Appendix 3. Index to genera and families (Shaw & Goffinet 2000, earlier classification in square brackets. * = new taxonomy). Adelotheciaceae [Hookeriaceae] Bryobrothera Amblystegiaceae Leptodictyum Anomodontaceae Herpetineuron Archidiaceae Archidium Bartramiaceae Bartramia Breutelia Philonotis Brachytheciaceae Brachythecium Eurhynchium Platyhypnidium Rhynchostegium Bruchiaceae Trematodon Bryaceae Brachymenium Bryum Gemmabryum* Rhodobryum Rosulabryum Ditrichaceae Ditrichum Eccremidium Garckea Wilsoniella Entodontaceae Entodon Mesonodon Ephemeraceae Ephemerum Erpodiaceae Erpodium Fabroniaceae Fabronia Fissidentaceae Fissidens Funariaceae Funaria Garovagliaceae Euptychium Garovaglia Gigaspermaceae Gigaspermum Grimmiaceae Grimmia Hookeriaceae Achrophyllum Cyathophorum Buxbaumiaceae Buxbaumia Calymperaceae Arthrocormus Calymperes Exostratum Leucophanes Mitthyridium Octoblepharum Syrrhopodon Hypnaceae Ectropothecium Glossadelphus Hypnum Isopterygium Taxiphyllum Trachythecium Vescicularia Cryphaeaceae Cryphaea Cyptodon Schoenobryum Hypnodendraceae Hypnodendron Hypopterygiaceae Hypopterygium Cyrtopodaceae Bescherellia Lembophyllaceae Daltoniaceae [Hookeriaceae] [Hookeriaceae] Calyptrochaeta Daltonia Distichophyllum Ephemeropsis Camptochaete Fallaciella Lembophyllum Leptostomataceae Leptostomum Leucobryaceae Leucobryum Leucomiaceae Leucomium Meesiaceae Leptobryum Meteoriaceae Aerobryopsis Barbellopsis Floribundaria Meteoriopsis Meteorium Papillaria Pseudospiridentopsis Trachypus Dicnemonaceae Dicranaceae Diphysciaceae Dicnemon Eucamptodon Campylopus Dicranella Dicranoloma Holomitrium Leptotrichella Leucoloma Sclerodontium Diphyscium Lopidium 407 408 Cunninghamia 8(3): 2004 Ramsay & Cairns, Mosses in the Wet Tropics bioregion NE Queensland Mitteniaceae Mittenia Pterobryellaceae Pterobryella Mniaceae Orthomnion Ptychomitriaceae Ptychomitrium Myriniaceae Macgregorella Ptychomniaceae Hampeella Myuriaceae Myurium Oedicladium Racopilaceae Powellia Racopilum Rhizogoniaceae Mesochaete Pyrrhobryum Rhizogonium Sematophyllaceae Acanthorrhynchium Acroporium Clastobryum Isocladiella Macrohymenium Meiotheciella Meiothecium Pseudohypnella Radulina Rhaphidorrhynchium Sematophyllum Taxithelium Trichosteleum Trismegistia Warburgiella Wijkia Sorapillaceae Sorapilla Sphagnaceae Sphagnum Stereophyllaceae Stereophyllum Nanobryaceae [Fissidentaceae] Nanobryum Neckeraceae Caduciella Himantocladium Homaliodendron Neckeropsis Pinnatella Thamnobryum Touwia Orthorrhynchiaceae Orthorrhynchium Orthotrichaceae Groutiella Macromitrium Schlotheimia Zygodon Pilotrichaceae [Hookeriaceae] Callicostella Cyclodictyon Hookeriopsis Polytrichaceae Dawsonia Notologotrichum Pogonatum Polytrichum Pottiaceae Anoectangium Barbula Hyophila Pseudosymblepharis Trachycarpidium Trichostomum Weissia Pterigynandraceae Trachyphyllum Pterobryaceae Calyptothecium Cryptogonium Muellerobryum Neolindbergia Pterobryidium Ptychomnion Symphyodontaceae [Ho okeriaceae] Chaetomitrium Thuidiaceae Pelekium Thuidiopsis Thuidium Trachylomataceae Braithwaitea Trachyloma Viridivelleraceae Viridivellus

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Habitat, distribution and the phytogeographical affinities of mosses in the Wet Tropics bioregion, north–east Queensland, Australia.

Habitat, distribution and the phytogeographical affinities of mosses in the Wet Tropics bioregion, north–east Queensland, Australia.

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