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Bryogeographical Relationships of the Mosses of Sri Lanka Brian 1. O'sheai

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Bryogeographical Relationships of the Mosses of Sri Lanka Brian 1. O'sheai J Hattori Bot. Lab. No. 93: 293-304 (Jan. 2003) BRYOGEOGRAPHICAL RELATIONSHIPS OF THE MOSSES OF SRI LANKA BRIAN 1. O'SHEAI ABSTRACT. The moss flora of Sri Lanka is comparatively rich (561 taxa) with a high level (11%) of endemism. This paper looks at the origin of the flora and relationships with neighbouring areas. De­ spite its Gondwanan origin, and proximity to India, there is a very strong link with Indochina and Malesia as well as with the Indian sub-continent. INTRODUCTION The objective of this paper and an associated checklist (O'Shea 2002) is to act as an introduction to the moss flora of the island of Sri Lanka, and to quantify the relationship of the moss flora with neighbouring areas: India, Indochina, Malesia and sub-Saharan Africa. Sri Lanka is a beautiful country with a diverse landscape and climate and a remark­ ably rich bryoflora, but is only 430 km from north to south and 224 km wide. The island is of Gondwanan origin, having started migration in the Cretaceous period about 135 million years ago with peninsular India (as part of the Deccan Plate) from an origin next to Mada­ gascar (Frahm 1994; Davis, Heywood & Hamilton 1995). The plate reached Asia about 40 million years ago, during the early Tertiary. It is likely that the climatic conditions at this time were warm temperate to sub-tropical, and climate similar to that of the present also might have prevailed during the drift of the Deccan plate (Ashton & Gunatilleke 1987). Sri Lanka only became an island in the post-Pleistocene period, as the glaciers melted, in about 5900 BP, although it had also been an island in earlier times, e.g. 30000 BP (Deraniyagala 1993). Data from Sri Lanka, India and Africa shows that 5900 BP was a time of heavy rainfall in all these areas, which continued to about 3700 BP, before the climate started to become drier and more seasonal (Roberts 1993), after which the present era of settled culti­ vation is likely to have begun (Ashton & Gunatilleke 1987). Sri Lanka also has a long his­ tory of human habitation: Homo sapiens activity in the area started at least 34000 BP, with the mesolithic period of micro lithic tools starting in around 28000 BP, at about the same time as this was happening in Eastern and Southern Africa (Deraniyagala 1993), with pos­ sible evidence of early agriculture in the area by 15000 BP (Flenley 1979). Much of the grassland in Sri Lanka is thought to be due to human forest clearance (Flenley 1979). All of the island receives at least 0.5 m of rain a year, with up to 5 m per year on the western slopes of the highlands (Somasekaram 1997). The wettest zone (2-5 m of rain per year) is in the SW and coincides with the most mountainous region, with a maximum ele­ vation of 2524 m, and an extensive area over 500 m. This allows areas of lowland and mon­ tane rainforest to exist in the Sw, but because of the monsoons, lowland forest exists throughout the island, although scarcer in the west (Collins, Sayer & Whitrnore 1991). The levels of endemism on the island are high (Collins, Sayer & Whitmore 1991): 1141 Fawnbrake Avenue, London SE24 OBG, u.K. (e-mail: [email protected]). 294 J. Hattori Bot. Lab. No. 93 2 0 0 3 over 50% of reptiles and amphibians, 14% of mammals, 8% of the birds and 28% of the flowering plants (most of the latter being in the rain forest). Prof. A. B. Abeywickrama produced a literature-based guide to the genera of Sri Lankan mosses (1960) and (with Jansen) a checklist (1978), but no specimen-based list has so far been produced .. BRYOLOGICAL EXPLORATION OF THE ISLAND Sri Lanka became a British colony in 1798, following a treaty signed with the King of Kandy, after almost 200 years of occupation by first the Portuguese and then the Dutch (Desmond 1992). The Royal Botanic Garden at Peradeniya was established in 1821, but it was not until 1844 when George Gardner became superintendent of the gardens that bryol­ ogy was included within its scope, and after his death aged 37 in 1849, Dr G. H. K. Thwaites continued the tradition until his retirement aged 68 in 1880. The specimens of Gardner & Thwaites provided the main impetus for Mitten's Musci Indiae Orientalis (1859), the first major contribution to our knowledge of the Sri Lankan mosses. The herbarium remains in Peradeniya, although it is understood that there are at present no resi­ dent bryologists on the island, and most recent collections have been lodged outside of Sri Lanka (see Table. I). There are many unpublished Sri Lankan collections in The Natural History Museum, London (BM), and it would be a major exercise to extract this data, but this would be nec­ essary to complete a collection-based list of Sri Lankan bryophyte distribution. A. H. G. Alston commenced such a collection-based list, probably in the early 1930s, but this was never completed and is unpublished. He also produced a rather more complete hepatics list. Both manuscripts are held in BM. MATERIALS AND METHODS Data quality The data used to produce the Sri Lanka checklist is held on a computer database (O'Shea 1993) that relates taxa to geographic areas and to the literature supporting that ge­ ographic distribution. It can thus be used not only to construct the checklist, but also to look at endemism, and taxa shared (or not shared) between geographic areas. The results of course depend on the quality of the data, and not all published data is of good quality, and some data derived from older papers will contain large numbers of taxa no longer accepted, other than as synonyms. Lists produced from such a database thus need a great deal of checking. The data for each geographic area considered in this paper has been 'cleaned' as much as possible, by eliminating nomina nuda, identifying and removing both homotypic and heterotypic synonyms (paying particular attention to recent revisions), checking against Index Muscorum and Index of Mosses, and validating against other checklists and relevant literature. Although the resulting quality is not perfect, for Sri Lanka it is quite good, and for other areas it is at least comparable with that of data used in other comparisons between palaeotropical areas, for instance with a recent study of the western Melanesian moss flora B. 1. O' SH EA: Bryogeographical relationships the mosses of Sri lanka 295 Table l. Collectors of bryophytes in Sri Lanka. Collector Year Published Collections* Moon, A. 1816-1825 BM Walker, A 1825- 1827 Mitten 1859 NY Wight, R. 1836 BM Maxwell(, E.?) « 1839?) Mitten 1859 NY Gardner, G. 1844-1849 Mitten 1859 BM, NY, PDA Thwaites, G. H. K. 1849- 1880 Mitten 1859, 1873 ; BM, NY,PDA Dixon 1930 Nietner, 1. < 1869 Muller 1869 B? Beccari, o. D. 1865 Hampe 1872 BM Beckett, T. W. N. < 1873, 1882- 1883 Mitten 1873; Fleischer B?, BM, BO?, NY 1915-1922 Hance, H. F. < 1886 BM Stone, A. B. 1891 BM Schiffner, V. 1893 Froehlich 1953 S,W Andersson ? BM Fleischer, M. 1898 Musci F1. Ind. Arch. BM Darrell, 1. H. 1898 Dixon 1914 BM Herzog, T. 1906 Herzog 1910[1911], 1926 BM Queste, Fr. 1911 Dixon 1914 BM Binstead, C. H. 1913 Dixon 1915, 1919 BM Alston, A. H. G. 1925- 1930 BM,PDA Inoue, H. 1966 Noguchi 1973 TNS Townsend, C. C. 1973 Townsend 1978, 1982, Hb. Townsend, BM 1983, 1991 Tixier, P. 1973 Tixier 1975 PC Ruinard, C. 1978 Townsend 1982, Hb. Townsend, PC? Tixier 1983 Onraedt, M. 1976, 1977, 1981 Onraedt 1986 Hb. Onraedt, BR Schaefer-Verwimp, A. 1984 BM, Hb. Schaefer- Verwimp * Locality of collections: B=Berlin, BM=London, BO=Bogor, BR=Brusse1s, PC=Paris, PDA= Peradeniya, S=Stockholm, TNS=Tokyo, W=Vienna. (Piippo & Koponen 1997), which is discussed later. It should be noted that both collecting and taxonomic activity is continuing, which may change the specific figures given here, but such changes are likely to be minor, and to affect floras of each geographic area in a similar way, so the overall results are unlikely to be significantly affected over time. Comments on datasets Sri Lanka: There is an existing checklist (Abeywickrama & Jansen 1978), now rather old, and compiled from previous lists. Main problems: errors from previous lists perpetuat- 296 1. Hattori Bot. Lab. No. 93 2 0 0 3 ed; using multiple source lists allowed several taxa to get on the list under more than one name; original publications not used; earlier synonyms not usually listed. A new checklist has therefore been produced (O'Shea 2002), and used for this exercise. Indochina: Recent checklist (Tan & Iwatsuki 1993) compiled from source literature. Main problem: many old and not recently used nomina nuda were included. Indochina in­ cludes Burma (Myanmar), Thailand, Cambodia, Laos and Vietnam. Malesia: No existing checklist, although the first three volumes of the late Alan Ed­ dy's 'Malesian Mosses' have been published, and a great deal of information has been pub­ lished in the 'Huon Peninsula' series of papers from Koponen et al. in Helsinki. Checklists exist for certain areas, such as the Malay peninsula (Dixon 1926), Philippines (Tan & Iwatsuki 1991), Lesser Sunda Isles (Touw 1992) and Borneo (Touw 1978), and Fleischer (1915- 1922) contains an overview of most taxa in the area. The amount of data held on the database used for this paper is by no means complete, and although it is hoped that the pro­ portion of endemic and non-endemic taxa will be about right, overall numbers are likely to be significantly understated.
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