J. Hattori Bot. Lab. No. 82: 105- 121(July 1997)

THE MOSSES OF THE YARRANGOBILLY CAVES DISTRICT, , A REVIEW OF THE MOSSES COLLECTED BY THE REVEREND W W WATTS IN 1906

1 1 ALISON J. DOWNINa1, PATRICIA M. SELKIRK AND RONALD J. 0LDFIELD

ABSTRACT. In 1906, Reverend W. W. Watts collected mosses in the vicinity of Yarrangobilly Caves, a limestone area in the Southern Alps of New South Wales and in 1912, with Brotherus, published a list of these species. This study updates the taxa recorded by Brotherus and Watts and compares the species collected by Watts in 1906 with our own recent (1993) collections. Most of the mosses collected by Watts are still present in the area. The few species no longer found in the immediate vicinity of Yarrangobilly are known from other locations in the Southern Alps of eastern Australia. Certain bryophyte taxa were not collected by Watts in 1906 but are now conspicuous components of the bryofiora at Yarrangobilly. Changes to the moss assemblage at Yarrangobilly are attributed to a range of environmental changes resulting from European settlement and development. KEYWORDS. Australia, bryophytes, calcareous substrates, limestone, mosses, Watts, Yarrangobilly.

INTRODUCTION In the northern hemisphere it has been possible to document changes in bryophyte as­ semblages at particular locations by comparisons of studies at various times (e.g. Jones 1991 ; Bates 1995). This has not been the case in Australia, where, until recently, there have been few published species lists. In January, 1906, Reverend W. Walter Watts collected mosses at Yarrangobilly Caves in the Southern Alps of New South Wales. The published species list from his collections was one of the earliest Australian bryological studies for a given location and certainly the first record of mosses on limestone in Australia (Brotherus & Watts 1912; Ramsay 1980). Although a detailed description of the geology of the Yarrangobilly area was included in Watts' paper, it is not possible from the information given to determine the nature of the substrate rock, whether calcareous or non-calcareous, from which each species was collected. We have re-examined the bryoftora of Yarrangobil­ ly and adjacent limestone areas and compared our recent collections with those of Watts of 1906. The Yarrangobilly Caves are located in a belt of massive Silurian limestone (Yarran­ gobilly Limestone) 12 km long and 1.5 km wide (from 35°38'S, 148°28'E to 35°44'S, 148°29'E) in a montaii.e to sub-alpine area of the Southern Alps of New South Wales. The limestone occurs on the western side of a plateau (average 1100 m altitude) through which the Yarrangobilly cuts a spectacular gorge so that the river bed is 200 m below the top of the plateau. Came & Jones (1919) have assessed the limestones as 98% calcium carbon­ ate. Bordering the limestone to the east are the Silurian Gooberaganda Volcanics of por-

1 School of Biological Sciences, Macquarie University, N. S. W. 2109, Australia. 106 J. Hattori Bot. Lab. No. 82 I 9 9 7 phyritic dacite and rhyodacite (Wyborn et al. 1990). Karst, the characteristic landscape where rock has been d.issolved by water rather than eroded, is well developed in the lime­ stone with numerous sink holes, caves and underground streams (Lishmund et al., 1986). At Yarrangobilly _C.aves, tall, open Eucalyptus forest dominates the vegetation includ­ ing Eucalyptus delegatensis R. Baker, E. viminalis Labill., E. dalrympleana Maiden, E. ru­ bida Deane & Maiden, E. paucifiora Sieber ex Sprengel and E. stellulata Sieber ex DC. with Leptospermum lanigerum (Aiton) Smith as an understory. Pinus sylvestris L. (Scots pine) has become naturalised in some areas. Numerous exotic trees and shrubs have been planted in the vicfo.ity of Caves House. At Yarrangobilly village, approximately 8 km north of the Caves, pine forest was es­ tablished in the 1920s and 1930s. The timber is now being cut and the area revegetated with local native species. The last residents of the village left in 1970 (New South Wales National Parks & Wildlife Service 1990). Some native grassland and Eucalyptus woodland occurs on limestone in the vicinity of the village. Yarrangobilly enjoys cool summers (mean summer temperature approximately 18°C and cold winters (mean approximately 5°C), with numerous frosts and some snow in win­ ter (Bureau of Meteorology, 1989). Rainfall is high, averaging 1150 mm per annum and is distributed relatively evenly throughout the year (Rainfall records maintained at Yarrangob­ illy Caves, unpublished). Aboriginal people are known to have occupied the areas surrounding Yarrangobilly Caves although it is believed that the caves were too cold and wet for habitation (New South Wales National Parks & Wildlife Service l 99la). The first European record of Yarrangobilly Caves is their discovery in 1834 by John Bowman while searching for cattle. Development of the area for tourists began in 1879 and over the next 70 years, e;ictensive building and road and path construction took place. Tourists came to Yarrangobilly, not only to visit the spectacular caves, but also to bathe in the thermal pool. This was con­ structed in 1897 around a natural hot spring which was held to have therapeutic qualities. Grazing probably began on the treeless, high altitude plains to the east ofYarrangobil­ ly in the 1840s. Stock was moved here from as far away as the western areas of New South Wales. Stock was excluded from alpine areas in 1944 and from other areas of the National Park by 1969 because of damage caused by grazing and potential contamination of dam catchment areas (New South Wales National Parks & Wildlife Service 1990). Mosses were also collected from Devonian Lickhole Limestone at Ravine (43°25'S, 146°53'E), lOkm south-west of Yarrangobilly Caves. Here the limestone is well-bedded limestone with interbedded calcareous shale (Lishmund et al., 1986). Along the road to Ravine, loose, nodular limestones form conspicuous terraces down the west facing slopes of Round Top Mountain, 996 m alt. down to 600 m alt. at Ravine village. Eucalypts are ab­ sent from most of the hillside terraces of nodular limestone at Ravine, although Eucalyptus woodlands and forests dominate adjacent non-calcareous substrates. The calcareous ter­ races are almost devoid of natural vegetation and have been invaded by introduced weed species, such as Hypericum sp., Rosa sp., Rubus spp. and Verbascum thapsus L. A. DOWNING et al.: The mosses of the Yarrangobilly Caves district, Australia 107

METHODS In order to compare our records with those of Watts, we determined synonyms for the taxa recorded by Brotherus and Watts by reference to Streimann & Curnow ( 1989). A study of Watts' collections at the National Herbariurn of New South Wales (NSW) was also undertaken, and Watts' specimens checked against modern keys such as Beever et al. (1992), Catcheside (1980), Scott & Stone (1976) and recent taxonomic publications includ­ ing Lewinsky (1984), Spence (1996), Spence & Ramsay (1996) and Sollman (1996). After some consideration of Zander (1993), we have decided to retain certain genera, such as Tortu/a rather than Syntrichia and Barbu/a rather than Pseudocrossidium for their relevant species in order to simplify comparisons. We considered some specimens be more appro­ priately assigned to taxa other than those used by Brotherus & Watts. Authorities are in­ cluded in Table 1 or in the text ifthe taxa are not included in the table. In December 1993, we collected mosses from the vicinity of Yarrangobilly Caves at locations specifically mentioned by Watts. The nature of the substrate from which each species was collected, whether calcareous, non-calcareous or epiphytic, was recorded. Ad­ ditional collections were made from calcareous and non-calcareous substrates at Ravine, and also from pine plantations, established on limestone north and east of Yarrangobilly village. These collections were returned to the laboratory for determination. We did not at­ tempt to locate two species recorded by Watts from the Turnut River, Brachythecium rivu­ lare and Eurhynchium cucul/atum, as we do not consider this to be part of the Yarrangobil­ ly district. We were unable to locate one of Watts' specimens, Funaria (Entosthodon) sp. (Watts 8910). Reference was made to recent collections of H. Streimann in the Australian National Herbarium in Canberra (CANB) to determine whether taxa collected by Watts in 1906, but not by us in 1993, are present in the Southern Alps of New South Wales and the Australian Capital Territory.

RESULTS AND DISCUSSION Our examination of Watts' collections and its terminology showed it to contain 95 taxa from 64 genera, somewhat Jess than his original 118 species from 61 genera (Table 1) . Some taxa proved particularly difficult to resolve. For example we have tentatively included his species of Campylium and Sciaromium in Cratoneuropsis relaxa, and we have lumped together numerous species of Philonotis as P. tenuis. These groups are in urgent need of re­ vision and our allocations of specimens to these taxa should not be seen to have taxonomic significance. Although most of the species collected by Watts at Yarrangobilly are not uncommon in south-eastern New South Wales (Ramsay l 984a), his collections included some unex­ pected and unusual taxa. Two species of Tortu/a, T. anderssonii and T. phaea are rare in Australia, and until recent collections from the Southern Alps were determined by Sollman ( 1996), T. phaea (as Syntrichia phaea (Hook. f. & Wils.) Zand.) was considered to be a New Zealand endemic (Fife 1995). Although Watts listed only 3 species of Orthotrichum in his species list, his collections included 4 species, 0. cupu/atum var. cupu/atum, 0. hort­ ense, 0. rupestre and 0. tasmanicum var. tasmanicum. In addition to these species, a fifth species, 0. assimile, is now not uncommon on the bark of some eucalypts and exotic tree 108 J. Hattori Bot. Lab. No. 82 I 9 9 7 species. Distichium capillaceum is known from calcareous substrates in the northern hemi­ sphere (Watson 1981 ), but we have not previously collected it from other south eastern Australian limestones. Almost every species collected by Watts was found to be still present in the immediate vicinity of Yarrangobilly Caves. However the current absence of some species previously (Brotherus & Watts 1912) collected appears to be related to environmental changes result­ ing from local landuse. The swamp where Watts was "rewarded with many treasures" (Brotherus & Watts 1912, p. 364), has disappeared with the construction and improvements to the thermal pool, and with it have gone some of Watts' "treasures" such as Drepan­ ocladus brachiatus and Campyliadelphus polygamus. Similarly, we presume that road con­ struction may have contributed to the demise of Sphagnum cristatum (Brotherus & Watts' S. subbicolor) and Meesia muelleri. However, recent collections by Streimann show that these species are not uncommon throughout the Southern Alps. Some species appear to have become more abundant following development. Poly­ trichum commune and P juniperinum are each represented by only one Watts' collection but both species are common and abundant on siliceous substrates, in particular the broad, frequently graded roadsides of the Snowy Mountain Highway. Prior to the planting of numerous exotic tree species, there were few trees suitable for epiphytic mosses since the local tree species shed bark annually. Watts' collections includ­ ed only 12 epiphytes and we found that most of these, species of Hypnaceae and Brachytheciaceae, grow only on the base of rough barked eucalypts. Orthotrichum hort­ ense, 0. tasmanicum var. tasmanicum, Zygodon intermedius and Z. hookeri were collected from the hard bark of some native shrubs. Our collections added an additional 36 species to Watt's list, giving a total of 131 species. Some of these species, such as Goniobryum subbasilare, Aulacomnion pa/ustre, Amblystegium riparium and Andreaea mutabi/is were collected from areas probably not ac­ cessible to Watts. A significant number of species collected by us but not present in Watts' collections have probably been introduced as a result of human activities. Species such as Barbu/a un­ guiculata, Brachythecium albicans and Pottia truncata are common in disturbed areas, near paths, roads, buildings, gardens and picnic areas. Bryum argenteum is a surprising omis­ sion from Watts' list, together with B. capillare, B. dichotomum, Tortu/a muralis, T. pago­ rum and T. papillosa. All are species common on concrete walls and pavements (Schofield 1980). Tortu/a pagorum and T. papillosa also grow on the bark of some eucalypts and on exotic trees in the vicinity of the caves. Both species may have been introduced on the bark of young trees brought to Yarrangobilly for horticultural purposes. Some species included by Watts, such as Ceratodon purpureus, Funaria hygrometrica and Leptobryum pyriforme also flourish in these disturbed areas. Grazing by sheep and cattle appears to contribute to change in bryophyte assemblages on calcareous substrates, often leading to an increase in the number of introduced and cos­ mopolitan species (Downing 1992) and may have influenced bryophyte assemblages at Yarrangobilly. From the 1830s until the end of grazing leases in the National Park in 1972, sheep and cattle were moved to high altitude treeless plains of the during A. DOWNING et al. : The mosses of the Yarrangobilly Caves district, Australia 109

the summer. In the 1954-55 season, it is estimated that 1500 head of cattle, and 40,000 sheep were moved from the lowlands via Yarrangobilly to the summer grazing leases (New South Wales National Parks & Wildlife Service. 199lb). We would suggest that Aloina aloides var. ambigua, Bryum argenteum (in part) and Trichostomum brachydontium are in­ troduced or cosmopolitan species which may have been introduced to the Yarrangobilly area by stock movements. In previous studies of eastern Australian limestones, we have found that the number of species collected from calcareous substrates is significantly greater than the number col­ lected from non-calcareous substrates. The bryoflora is typically dominated by acrocarpous mosses from two families, Pottiaceae and Bryaceae. Certain key species, Gigaspermum repens, Fissidens vittatus, Didymodon torquatus and Bryum dichotomum and/or Bryum pachytheca are always present. Not only does this assemblage typify eastern Australian limestones, it is also typical of the bryoflora of arid and semi-arid calcareous soils in south central Australia (Downing 1992; Downing 1993; Downing & Selkirk 1993 ). The moss assemblage Watts collected on limestone at Yarrangobilly in 1906, was, to a certain extent, representative of assemblages found on eastern Australian limestones now. Many more species of acrocarpous mosses (38) than of pleurocarpous mosses (24) were collected from limestones. Pottiaceae (9 species) and Bryaceae (10 species) dominated the bryoflora, and Didymodon vinealis (syn. D. torquatus) was present. However, other key species, such as Gigaspermum repens, Fissidens vittatus, Bryum dichotomum and B. pachytheca were not present. In 1993, numerous additional taxa, including 22 acrocarps and 5 pleurocarps, were collected from the Yarrangobilly limestones, producing a moss assemblage on limestone much more typical of those of other eastern Australian limestones. There are now twice as many species of acrocarpous mosses (60) as pleurocarpous mosses (29), including 13 addi­ tional species of Pottiaceae (total 22) and 6 species of Bryaceae (total 16). Didymodon vinealis is still present, but, surprisingly, all the remaining key species, Gigaspermum repens, Fissidens vittatus, Bryum dichotomum and B. pachytheca are now present. There is now not a great difference in the number of species which occur on calcareous substrates (89 including 46 exclusives) compared with the number on non-calcareous substrates (72 including 33 exclusives). Pleurocarpous mosses now present include some taxa which have not previously been recorded by us from eastern Australian limestones, such as Disti­ chophyl/um crispulum, D. microcarpum, Acrocladium chlamydophyllum, Cratoneuropsis relaxa and numerous species of Brachytheciaceae. In other studies of Australian limestone bryoflora, Downing (1993) and Downing & Selkirk (1993) proposed that the ratio ofacro­ carpous to pleurocarpous mosses appeared to be related to rainfall, in that the number of pleurocarpous mosses increased with increasing rainfall. Eldridge & Tozer (1997) in stud­ ies of microbiological crusts in southern Australia have found that increase in rainfall re­ duces both the pH and the calcium content of arid and semi-arid soils. At Yarrangobilly, it seems probable that high rainfall (1150mm per annum) and reduced evaporation due to cool temperatures lead to a lowering of pH and reduction in the effect of calcium, allowing calcifuge species to grow on calcareous substrates. This may also account for the presence of three acrocarps, Breutelia affinis, B. pendula and Ceratodon purpureus, which are con- 110 J. Hattori Bot. Lab. No. 82 1 9 9 7 sidered to be calcifuges in south eastern Australia, yet they are not uncommon on calcare­ ous substrates at Yarrangobilly. Some of the additional tax.a now present at Yarrangobilly are those of microphytic soil crusts characteristic of calcareous soils of semi-arid and arid areas of western New South Wales, Victoria and South Australia. These species are Alaina aloides var. ambigua, Didy­ modon vinea/is, Barbu/a ca/ycina, B. crinita, B. hornschuchiana, Bryum argenteum, B. pachytheca, B. campylothecium, Crossidium geheebii, Desmatodon convolutus, D. recurva­ tus, Tortu/a antarctica, Trichostomum brachydontium and two epiphytes, Tortu/a pagorum and T. papillosa (Downing & Selkirk 1993; Eldridge & Tozer 1997). We consider that there are three elements which have contributed substantially to the presence of these arid and semi-arid area species at Yarrangobilly, a suitable substrate for colonisation, a source of propagules, and a means of dispersal. Development of roads, paths, buildings, farms and gardens at Yarrangobilly, begin­ ning in earnest about the tum of the century and still continuing to the present day, has pro­ vided a constantly changing environment which has regularly provided newly disturbed soil or rock surfaces suitable for bryophyte colonisation. Microphytic soil crusts on calcareous soils in western New South Wales, Victoria and South Australia are the probable source of propagules and we consider dust storms to be the most likely means of dispersal. Although dust storms have been occurring in Australia for the last 2 million years, there is evidence to suggest that changing land use following European settlement has significantly acceler­ ated the loss of soils from arid and semi-arid areas during dust storms (McTainsh 1989; McTainsh & Leys 1993). Severe droughts in south-eastern Australia from the late 1800s until 1940s were often followed by severe dust storms (Yu et al. 1993) and as recently as 1983, a massive dust storm which began in South Australia carried dust to the east coast of Australia (Bureau of Meteorology 1989). In glasshouse trials, moss propagules present in calcareous soils from semi-arid Australia "germinate" and grow rapidly, forming a com­ plete cover within 8 weeks (Downing 1993). The speed with which mosses are able to colonise bare ground has probably been significant in their ability to colonise eastern Aus­ tralian limestones which experience much higher rainfall than their place of origin. The arrival of the epiphytes Tortu/a pagorum and T. papillosa at Yarrangobilly since Watts' visit may also be attributed to dust storms. Both species are common on the bark of black box (Eucalyptus largiftorens F. Muell.) in western New South Wales, and propagules may well have been dispersed by dust to Yarrangobilly. Exotic trees planted at Yarrangobil­ ly in association with tourist development many have provided a suitable substrate on which propagules of these arid zone species could lodge although propagules may have been present on the bark of young exotic trees prior to their planting at Yarrangobilly.

CONCLUSION Ramsay ( l 984b) commented on the vital contribution made by early collectors whose specimens may be the only record of the bryoflora of an area subsequently altered by Euro­ pean settlement. Although many undisturbed locations in the vicinity of Yarrangobilly, on both calcareous and non-calcareous substrates, have bryophyte assemblages similar to those collected by Watts in 1906, there are significant differences in areas which have been A. DOWNING et al.: The mosses of the Yarrangobilly Caves district, Australia Ill intensively utilised.

ACKNOWLEDGMENTS We thank the National Parks and Wildlife Service of New South Wales for their sup­ port and permission to work within the Snowy Mountains National Park. We thank the Commonwealth Key Centre for Biodiversity and Bioresources at Macquarie University for financial support for this study. We thank the National Herbarium of New South Wales for access to Watts' collections and Helen Ramsay, Elizabeth Brown and Robert Coveny for their assistance. Heinar Streimann, Helen Ramsay, R. Ochyra, Hans Kruiger, Rod Seppelt and Graham Bell assisted us with the identification of some difficult species, and we thank Heinar Streimann for access to information from the collections of the Australian National Botanic Gardens (CANB). Peter Mitchell assisted with information on dust storms. We thank Don Adamson whose knowledge of the Southern Alps, Yarrangobilly, Kiandra and Ravine areas proved invaluable.

REFERENCES CITED Bates, J. W. 1995. A bryophyte flora of Berkshire. J. Bryology 18 (3): 503--620. Beever, J. , K. W. Allison & J. Child 1992. The Mosses of New Zealand. University of Otago Press, Dunedin. Brotherus, V. F. & W. W Watts. 1912. The mosses of the Yarrangobilly Caves District, N. S. W. Proc. Linn. Soc. New South Wales 37 (2): 363-382. Bureau of Meteorology. 1989. Climate of Australia. Australian Government Publishing Service, Can­ berra. Came, J.E. and L. J. Jones 1919. The Limestone Deposits of New South Wales. Mineral Resources. No. 25. Government Printer, Sydney. Catcheside, D. G. 1980. Mosses of South Australia. Government Printer, South Australia. Downing, A. J. 1992. Distribution of bryophytes on limestones in eastern Australia. Bryologist 95 (1): 5-14. Downing, A. J. 1993. Distribution of bryophytes on calcareous substrates in South Eastern Australia. M. Sc. thesis, Macquarie University, unpublished. Downing, A. J. & P. M. Selkirk 1993. Bryophytes on the calcareous soils of Mungo National Park - an arid area of southern central Australia. Great Basin Naturalist 53 (1): 13-24. Eldridge, D. J. and M. E. Tozer 1997. Environmental factors controlling the distribution ofterricolous bryophytes and lichens in semi-arid eastern Australia. Bryologist: in press. Fife, A. 1995. Check.list of the mosses ofNew Zealand. Bryologist 98 (3): 313- 337. Jones, E.W. 1991. The changing bryophyte flora of Oxfordshire. J. Bryology 16: 513- 549. Lewinsky, J. 1984. The genus Orthotrichum Hedw. (Musci) in Australasia. A Taxonomic Revision. J. Hattori Bot. Lab. 56: 369-460. Lishmund, S. R., A. D. Dawood and W. V. Langley 1986. The Limestone Deposits of New South Wales. Geological Survey of New South Wales. Mineral Resources 25, 2nd Edition. NSW De­ partment of Mineral Resources. N. S. W. Government Printer, Sydney. McTainsh, G. H. 1989. Quaternary aeolian dust processes and sediments in the Australian region. Queensland Sci. Revue 8: 235- 253. McTainsh, G. H. & J. F. Leys 1993. Soil erosion by wind. Jn : G. H. McTainsh & W. C. Boughton (Ed­ itors). Land Degradation Processes in Australia. Longman Cheshire, Melbourne. 112 1. Hattori Bot. Lab. No. 82 I 9 9 7

New South Wales National Parks & Wildlife Service. 1990. The South End. Kosciusko National Park. NSW National Parks & Wildlife Service, Hurstville. New South Wales National Parks & Wildlife Service. 199la. Yarrangobilly Caves. Limestone Caves and Short Walks in the Yarrangobilly Valley. NSW National Parks & Wildlife Service, Hurstville. New South Wales National Parks & Wildlife Service. 199lb. The North End. Kosciusko National Park. NSW National Parks & Wildlife Service, Hurstville. Ramsay, H. P. 1980. Contributions of Rev. W.W. Watts F. L. S. to Australian botany. Taxon 29 (4): 455-469. Ramsay, H.P. 1984a. Census of New South Wales Mosses. Cunninghamia 2 (5): 455- 534. Ramsay, H.P. 1984b. Phytogeography of the mosses of New South Wales. Cunninghamia 2 (5): 535- 548. Scott, G. A.M. & I. G. Stone 1976. The Mosses of Southern Australia. Academic Press, London Schofield, W. B. 1980. The phytogeography of the mosses of North America (North of Mexico). In : Taylor, R. J. & Leviton, A. E. (editors). The Mosses of North America. Pacific Division, Ameri­ can Association for the Advancement of Science. San Francisco. Sollman, P. 1996. Studies on Australian Pottiaceous mosses. Lindbergia 20: 144-146. Spence, J. R. 1996. Rosulabryum genus novum (Bryaceae). Bryologist 99 (2): 221-225. Spence, J. R. & H.P. Ramsay, 1996. New and interesting species of the family Bryaceae (Bryopsida) from Australia. J. Adelaide Bot. Gard. 17: 107_c_l 18. Streimann, H. & J. Curnow 1989. Catalogue of Mosses of Australia and its External Territories. Aus­ tralian Flora and Fauna Series, No. 10. Australian Government Publishing Service, Canberra. Watson, E. V. 1981. British Mosses and Liverworts. Cambridge University Press, Cambridge. Wyborn, D., M. Owen & L. Wyborn 1990. Geology of the Kosciusko National Park (1: 250,000 scale map). Bureau of Mineral Resources, Canberra. Yu, B., P. P. Hesse & D. T. Neil 1993. The relationship between antecedent regional rainfall condi­ tions and the occurrence of dust events at Mildura, Australia. J. Arid Environments 24: 109-124. Zander, R. H. 1993. Genera of the Pottiaceae: Mosses of Harsh Environments. Bull. Buffalo Soc. Nat. Sci. Vol. 32. Buffalo, New York. Table I. Mosses collected at Yarrangobilly Caves. W= collected by Watts, 1906; P= collected in present studies, 1993; S= taxa not collected in present studies but in CANB collected by Streimann in Australian Southern Alps; C=calcareous substrate; N=non-calcareous substrate; E=epiphytic

Tan present w p s Names used by Brotherus & Watts for Notes c N E Watts 1906 collections ?>' 0 Sphqnaceae w s Sphagnum subbicolor Hpe Not common N Sphagnum cristatum Hampe ~z Ditrlcbaceae w p Ceratodon purpureus (L.) Brid. Common on both calcareous and non-calcareous c N Cl ~ Ceratodon purpureus (Hedw.) Brid. Ceratodon purpureus f. nervo excedente substrates. Particularly common in disturbed areas. ·~ Distichium capillaceum (Hedw.) B.S.G. w p Distichium capillaceum (Sw.) Br.Eur. Common on limestone, especially damp soil in c sheltered areas. ~ g Ditrichum difficile (Dub.) Reisch. w p Ditrichum difficile (Dub.) Reisch. Common and abundant on siliceous substrates. N "'~ Pleuridium nervosum (Hook.) Miu. p Pleuridium gracilentum (Miu.) Common and abundant on siliceous substrates, w N ....,"'0 frequently on disturbed roadside banks. e- Ditrichum cylindricarpum (C .Muell.) " F.Muell. p Occasional on non-calcareous substrates. N g § Dic:nmaceae (included in Ditrichaceae) Campy/opus introflexus (Hedw.) Brid. w p Campy/opus introflexus (Hedw.) Mitt. Common and abundant on non-calcareous substrates N ~ .z Di crane Ila cardotii (R. Br. ter.) Dix. w p Dichodontium wattsii Broth. sp. nov. Uncommon on calcareous substrates c Dicranel/a dietrichiae (C.Muell.) Jaeg. w p Dicranella sp. (?) (8875) In the absence of sporophytes it is not possible to N ~ or D. jamesonii (Mitt.) Broth. Dicranel/a sp. (?) (8754) determine Watts' collections. Tuey arc most e: probably D. dietrichiae. j"' · Dicranoloma billardieri (Brid.) Par. w p Dicranoloma subpungens (Hpe) On siliceous rock in wet gullies N Dicranoloma sullivani (C.M.) Sclerodontium pallidum (Hook.) Schwaegr. w p Leucoloma Sieberi Hsch. On siliceous rock. N i;::., (syn. Dicnemoloma pallidum (Hook.) Wijk & Marg.)

Campy/opus pyriformis (Schultz) Brid. p Occasional on non-calcareous substrates. N

..., Table I. (continued) ~

Flssldentaceae Fissidens /eptocladus C.Muell. ex Rodw. w p Fissidens leptocladus C.M. Common on moist, calcareous soil c Probably Fissidens taylorii C .Muell. w p Fissidens lilliputano-incurvus C.M. F. taylorii occurs occasionally on limestone. c Fissidens rigidulus Hook. f. & Wils. w p Fissidens rigidulus H.f.W. Occasional in or near streams on both calcareous c N ssp. rigidulus and siliceous substrates. Fissidens vittatus Hook. f. & Wils. p Common on calcareous soil in exposed situations. c Abundant on nodular oolitic limestone at Ravine.

Pottlaceae Barbu/a chlorotricha (Broth. et Gch.) Par. Common and abundant on calcareous substrates, Barbu/a crinita Schuliz w p Barbu/a pseudo-pilifera Hpe. et C.M. occasional on siliceous substrates. c N

Barbu/a luteola (Mitt.) Par. w p Barbu/a pseudo-pilifera var. scabrinervis Occasional on limestone c ~ ::i:: Didymodon vinealis (Brid.) Zand. (syn. D. w p Barbu/a torquata Tayl. Common and abundant on limestone, occasional on c N g torqUDtus (Tayl.) Catches.) siliceous substrates. Exposed situations. :i. t.ll Gymll()stomum calcareum Nees & Hornsch. p Gymnostomum calcareum Br. Germ. Occasional on limestone, common near the caves. w c ~ Torrella daldnii Willis w p Torte/la knightii (Mitt.) Peristome teeth are no longer present on Watts' c ~ specimens which are probably T. dakinii. z Aongstr. p Tortu/a (Syntrichia) brunnea Broth. Common on limestone throughout the area. !=' Tortu/a anderssonii w c 00 et Watts, sp.nov. ...., Tortu/a antarctica (Hampe) Wils. in Hook.f. w p Tortu/a panduraefolia (Hpe. et C.M.) Common and abundant on calcareous substrates, c N Tortu/a princeps De Not. occasional on siliceous substrates. Tortu/a phaea (Hook.f. & Wils.) Dix. w s Tortu/a (Syntrichia) subbrunnea Broth. et Uncommon on siliceous rock. N Watts, sp. nov. Tridontium tasmanicum Hook. f. w p Tridontium tasmanicum Hook. f. Common on stream banks in limestone areas. c Triquetrella papillata (Hook.f & Wils.) w p Triquetrella papillata (H.f.W.) Common and abundant on calcareous and siliceous c N Broth. Triquetrella albicuspes Broth. substrates

Aloina aloides (Schultt) Kindb. var. ambigua (B .S.G.) Craig. p On calcareous soil in exposed situation at Ravine. c Barbu/a calycina Schwaegr. p Occasional on non-<:alcareous soils. N Barbu/a homschuchiana Schul!Z p Occasional in exposed situations on calcareous soils. c '° '°-..l Barbu/a unguicu/ata Hedw. p Occasional on calcareous substrates, especially in c disturbed areas.

Crossidium geheebii (Broth.) Broth. p On calcareous soil at Ravine). c Desmatodon convolutus (Brid.) Grout. p On calcareous soil in exposed situation at Ravine. c ?>' t:I Desmatodon recurvatus (Hook.) Miu. p On calcareous soil in exposed situation at Ravine. c

Pottia truncata (Hedw.) Br. & Schimp. p Common on calcareous substrates in disturbed c N ICl areas. Occasional on non-alcareous substrates. ~ ~ Tortu/a muralis Hedw. p Common on mortar in limestone walls. c ;l Tortu/a pagorum (Milde) de Not. p Occasional on limestone, common on exotic trees. c E <> Tortu/a papillosa Wils. p Occasional on limestone, occasional on exotic treeS. c E p Uncommon, on limestone. I Tortu/a ruralis (Hedw.) Gaertn, Meyer & c 0 Scherb. -. ~ Trichostomum brachydontium Bruch. p Common on calcareous soil, especially in disturbed c areas. g § Weissia controversa Hedw. p Common on calcareous soil, especially disturbed c N ~ areas. Occasional on non-alcareous substrates. [ -< Encalyptaaae Occasional on limestone rock, uncommon on non- Encalypta vulgaris Hedw. w p Encalypta tasmanica Hpe. et C.M. calcareous substrates. c N ? "'c:i.. Grimmiaceae !a" Grimmia laevigata (Brid.) Brid. w p Grimmia campestris Burch. Occasional on non-alcareous substrates N :2 . rn Grimmia pulvinata (Hedw.) Sm. var. w p Grimmia pulvinata (L.) Sm. , var. obtusa Common and abundant on limestone rock, c N africana (Hedw.) Hook.f. & Wils. (Brid.) Br.Eur. occasional on non-alcareous rock. ~ [ Racomitrium crispulum (Hook.f. & Wils.) w p Rhacomitrium pseudo-patens (C.M .) Par. Common and abundant on siliceous substrates. N ; · Hook.f. & Wils. Rhacomitrium symphyodon Mitt.

Schistidium apocarpum (Hedw.) B.S.G. w p Grimmia apocarpa (L.) Hedw. Common and abundant on calcareous substrates, c N Grimmia apocarpa f. submutica occasional on siliceous substrates. Grimmia mutica Hpe.

Grimmia trichophyl/a Grev. p Occasional on non-alcareous substrates N u. Table I. (continued) ;; Ortbotrlchaceae Amphidium eyathicarpum (Mont.) Broth. w s Amphidium eyathicarpum (Mont.) Not collected by us. N Orthotrichum cupulatum Hoffm. ex Brid. var. w p Orthotrichum Sul/ivani C.M. (Watts 8949) Common and abundant on limestone rock c cupulatum.

Orthotrichum hortense Bosw. w p Orthotrichum acroblepharis C. M. Epiphyric E

Orthotrichum rupestre Schwaegr. w p Orthotrichum Sul/ivani C.M. (Watts 8827) Occasional on non-calcareous rock. N

Orthotrichum tasmanicum Hook.f. & Wils. w p Orthotrichum laterale Hpe. Epiphyric on exotic trees in the vicinity of the c E var. tasmanicum limestone caves, occasional on limestone. Zygodon inurmedius B.S.G. w p Zygodon Brownii Schwgr. Common and abundant on limestone rock, c N E Zygodon intermedius Br.Eur. occasional on non-calcareous rock, also epiphytic Zygodon hookeri Hampe. w p Zygodon hookeri Hpe Occasional on limestone rock, also epiphytic c !--< :c Orthotrichum assimile C. Muell. p Epiphyric. E g ::i. Zygodan menziesii (Schwaegr.) Amott p On limestone rock, in a moist, sheltered and shaded c situation. ?-°' Funarlaceae Funaria (Entosthodon) sp. Specimen not located. ~ ? w (Watts 8910) z 9 Funaria hygrometrica Hedw. w p Funaria hygrometrica (L.) Common on disturbed soil. c N 00 N Entosthodon muehlenbergii {Turn.) Fife (syn. Funaria tasmanica Hpe. et C.M. Common on calcareous soil c Funaria glabra TayI. ) w p Funaria aristata Broth. Bryaceae Bryum amblyodon C. Muell. (det. J. Spence). w p Bryum altisetum C.M. Occasional on calcareous and siliceous substrates c N Bryum blandum Hook. f. & Wils. w p Bryum blandum H.F.W. Common on streams banks in limestone c Bryum laevigatum H. f. W.

Bryum caespiticium Hedw. w p Bryum caespiticium L. Collected from calcareous and non-calcareous c N substrates.

B. su/livanii C. Muell. w Bryum Sullivani C.M . Uncommon on calcareous substrates c Bryum pseudotriquetrum (Hedw.) G.M.S. w p Bryum bimum Schreb. Occasional on calcareous and siliceous substrates c N Bryum ventricosum Dic.ks. \0 \0 Leptobryum pyriforme (Hedw.) Wils. w p Leptobryum pyriforme (L.) Wils. Occasional on limestone (Eagles Nest) c __, Poh/ia cruda (Hedw.) Lindb. w Poh/ia cruda (L.) Lindb. Not collected by us. c N Pohlia nutans (Hcdw.) Lindb. w Pohlia nutans (Schreb.) Lindb. Not collected by us. c N Poh/ia wahlenbergii (Web. & Mohr) w p Mniobryum tasmanicum Broth. Occasional on non

Rosulabryum billardierei Schwaegr. var. w p Bryum abruptinervium C.M. Common and abundant on a wide range of c N > billardierei Bryum pohllaeopsis C.M. substrates. Rosulabryum torquescens Bruch. ex de Not. w p Bryum pyrothecium Hpe. et C .M. Common and abundant on calcareous substrates. c N ~ Occasional on siliceous substrates. ~ ~ Schizymenium bryoides Hov. ex Hook. w p Mie/ichhoferia australis Hpe Common on non

Aulacomnion palustre (Hedw.) Schwaegr. p Occasional in swampy ground on non

Bartramiaceae Bartramia erecta (Hpe) Broth. (collected Common on non-Ollcareous substrates. N Bartramia hampei (Mitt.) Catches. w p W. Forsythe 1901) Bartramia ithyphyl/a Brid. w p llartramia papil/ata H. f. W . Common on non-Ollcareous substrates, especially N OD damp soil.

Bartramia mossmaniana C. Muell. (syn. B. w p Bartramia mossmaniana C.M. Not uncommon in swampy, non~careous areas N hal/eriana)

Breutelia qffinis (Hook.) Mitt. w p Breutelia ajfinis (Hook.) Mitt. Common and abundant on siliceous substrates, c N Breutelia commutata (Hpe.) Par. occasional on calcareous substrates.

Breutelia pendu/a (Sm.) Mitt. w p Breutelia pendu/a (Hook.) Jaeg. Occasional on siliceous substrates, less common on N Breutelia divaricarta Mitt. calcareous substrates

Philonotis scabrifolia (Hook. f. & Wils.) w p Philonotis scabrifolia (H.f.W.) Uncommon on siliceous substrates in moist, N !- Braithw. sheltered locations. g::c: Philonotis tenuis (Tay I.) Reichdt. w p Philonotis sp. (?) 8778 We have experienced great difficulty in attempting N :1 . Philonotis sp. (?) 8817 to resolve the taxonomy of these species. It appears t:tl Philonotis sp. (?) 8868 likely that all the specimens examined could be ~ Philonotis sp. Broth. et Watts, sp. nov. attributed to the species commonly known in specimen lost. Australia as P. tenuis. ~ Philonotis austro{alcara Broth. et Watts. z ? Common, panicularly on wet rock or soil. sp.nov. ...,00 Philonotis dicranel/acea C.M. Philonotis fontanoides Broth. et Watts, sp. nov. Philonotis tenuis Tay!. Polytrichaceae Common on disturbed ground on siliceous Polytrichum commune Hedw. w p Polytrichum commune L. substrates, especially roadside embankments. N Polytrichum juniperinum Hedw. w p Polytrichum juniperinum Willd. Common on disturbed ground on siliceous N substrates, especially roadside embankments.

Hedwigiaceae Hedwigia ciliata (Hedw.) Beauv. w p Hedwigia albicans (Web.) Lindb. Occasional on non-calcareous substrates N Hedwigidium integrifolium (Beauv.) Dix. w p Hedwigidium imberbe B. & S. Occasional on non-Ollcareous substrates N Ned

Thuldlaceae Thuidiumfutfurosum (H.f.W.) Jaeg. Usually in sheltered, moist situations e.g. in gullies. c N Thuidium futfurosum Reichdt. w p Thuidium hastatum C.M. On calcareous and siliceous substrates. Thuidium unguiculatum H.f.W. Thuidium sparsum (Hook. f. & Wits.) Jaeg. w p Thuidium sp. (Watts Ml2102) Common in wide range of situations on calcareous c N Thuidium suberecrum (Hpe.) Jaeg. and siliceous substrates. Often in exposed situations. "' Table 1. (continued) N 0

Anomodon tasmanicus Broth. p Occasional on limestone, occasional on quanz c N porphyry. Amblystepaceae w p Acrocladium ch/amydophyllum (H.f.W.) Common and abundant on limestone, especially in c E Acroc/adium chlamydophyllum (Hook.f. & Broth. moist, sheltered locations. Also epiphytic, climbing Wils.) C.Muell. & Broth. over and through tree srumps and suckers.

CampyUadelphus polygamus (B.S.G.) Kanda w s Drepanocladus strictifolius Broth. et Probably disappeared at some stage during work on Watts, sp. nov. the thermal pool. Cratoneuropsis relaxa (Hook.f. & Wils.) w p Campylium decussatum (H.f.W.) Common on limestone, especially in sheltered, c N Fleisch. Campylium relaxum (H.f.W.) moist environments such as cave enfiances and sink Campylium subrelaxum Broth. holes. Less common on non-calcareous substrates Sciaromium elimbatum Broth. et Watts, where it is always associated with wet, sheltered sp. nov. environments. Sciaromium forsythii Broth., sp. nov. ~ Drepanocladus brachiatus (Mitt.) Dix. w s Drepanocladus brachiatus (Mitt.) Not collected ,by us, and probably disappeared f Drepanoc/adus fluitans (Dill.) var. during construction of the thermal pool. Not ;::! . l:P falcatus Sch. uncommon in the Southern Alps. ~ Amblystegium riparium (Hedw.) B.S.G. (syn. p In swampy ground on non-calcarous substrates. N Kiandra goldtields. ~ Leptodictyum riparium) z Hypnaceae Ectropothecium condensatum Broth. et 9 E 00 Ectropothecium condensatum Broth. et Watts w Watts N

Hypnum cupressiforme Hedw. var. w p Stereodon cupressiformis (L.) Brid. (8795) Occasional on a wide range of substrates. c N E cupressiforme Stereodon walterianus (Hpe.)

Hypnum cupressiforme Hedw. var. /acunosum w p Stereodon cupressiformis (L.) Brid. (8508) Common and abundant on limestone c Brid. Hypnum cupressiforme Hedw. var. w p Stereodon Mossmannianus (C.M.) Occasional on calcareous and siliceous substrates c N mossmanianum (C.Muell.) Ando.

Hypnum cupressiforme Hedw. var. filiforme p Brid. Uncommon c N Sematophyllaceae Rhaphidorrhynchium cal/idioides (C.Muell.) w p Rhaphidostegium callidioides (C.M.) Jaeg . Uncommon, epiphytic. E Broth. 'C Wijkia extenuata (Brid.) Crum w p Acanthoc/adium extenuatum (Brid.) Mitt. Occasional, mostly on non-calcareous substrates. N E 'C ..... Plagiotheciaceae w s lsopterygium amblyocarpum (Hpe.) Material collected by Watts appears to fit the lsopterygium limatum (Hook.f. & Wils.) description of I. limatum in Scou & Stone (1976) . Broth.

Brachytheciaceae ?>­ p Brachythecium paradoxum (Hook.f. & Wils.) w Brachythecium paradoxum (H .f.W.) Jaeg. Common and abundant on a wide range of c N E Cl Jacg. substrates ~ Brachythecium rivulare B. S. G. w Brachythecium rivulare Br. Eur. (Tumut We were unable to locate the original specimen. River) ~ ~ Brachythecium rutabulum (Hcdw.) B.S.G. w p Brachythecium plumosum (Sw.) Common and abundant, especially in grassy areas, c N E ~ Brachythecium rutabulum (L.) Br.Eur. on a wide range of substrates ~ Brachythecium salebrosum (Web. & Mohr) w p Brachythecium salebrosum (Hoffm.) Occasional, especially in grassy areas, on a wide c N E g B.S.G. Br.Eur. range of substrates. ~ p "' Eurhynchium austrinum (Hook.f. & Wils.) w Oxyrhynchium austrinum (H.f,W.) Broth. Occasional in a wide range of substrates c N E "'0...., Jaeg. (syn. Platyhypnidium austrinum (sic) (Hook.f. & Wils.) Fleisch. g- ~ Eurhynchium cucullatum (Miu.) Stone & w Rhynchostegiella subconvolutifolia Broth. Not collected E Scoll et Wal!S, sp. nov . () ~ Rhynchostegiella muriculata (Hook. f. & w p Amb/ystegium Novae-Valesiae Broth. et Occasional at Yarrangobilly Village. c [ Wils.) Broth. Watts, sp. nov. (Originally included by -< Brotherus & Watts in Hypnaceac) ~ Rhynchostegium tenuifolium (Hedw.) Reichdt. w p Rhynchostegium collatum (H.f.W.) Occasional on a range of substrates. c N "' Rhynchostegium tenuifolium (Hedw.) Jaeg. e: ~. Brachythecium albicans (Hedw.) B.S.G. p Common on calcareous substrates, especially in c •!:i. disturbed areas in the vicinity of Caves House. > Andreaeaceae p Andreaea mutabilis (Hook.f. & Wils.) Uncommon on siliceous rock. N io; ·

Gi&aspennaceae Occasional on calcareous soil, usually exposed c Gigaspermum repens (Hook.) Lindb. p situations.

N