J Hattori Bot. Lab. No. 93: 293-304 (Jan. 2003)

BRYOGEOGRAPHICAL RELATIONSHIPS OF THE OF

BRIAN 1. O'SHEAI

ABSTRACT. The 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 , 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 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- period, as the glaciers melted, in about 5900 BP, although it had also been an island in earlier times, e.g. 30000 BP ( 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 (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 , 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 (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. Malesia includes Indonesia, Malaysia, Papua and Philippines. India: India is probably the largest area of land in the world without a published checklist. There have been lists published in the past (e.g. Mitten 1859, Briihl 1931) but these were either not complete or not critical, are difficult to match to current geographical boundaries and are now considerably out of date. Chopra (1975) covered a rather larger area (e.g. including the non-Indian ) and Gangulee (1969-80) only the east. As an example of the problems associated with this older data, the original list of endemics was reduced from 598 to 288, mainly by removing homotypic and heterotypic synonyms. Although mosses are not respecters of political boundaries, legislators are, and it is impor­ tant to be able to provide lists for political units, for instance for purposes of conservation legislation. The database of Indian mosses used for this exercise is gathered from many sources but is certainly not complete, but other authors should remedy this. The country is too large and too geographically diverse to be really meaningful in biogeographic terms, and a list for just peninsular India (or even just the southern part of the peninsula) would have been useful for comparison with Sri Lanka. Hopefully this will be possible when dis­ tribution data by State is also available. Sub-Saharan Africa: This area was included to measure the influence of the Gond­ wanan origins of the Sri Lanka flora versus more recent influences. There is an existing checklist for the area which is regularly updated, which also includes the list of 72 coun­ tries and islands included in the totals (O'Shea 1999); the number of en demi cs is also pub­ lished (O'Shea 1997).

RESULTS The results focus on two areas: the levels of endemism in the area, and the relation­ ships between the moss floras of the different geographic areas.

Endemism Table 2 shows the level of endemicity for each geographic unit and relates this to the B. 1. O'SHEA: 8 ryogeographical relationships the mosses of Srilanka 297

Table 2. Endemic taxa.

No. of No. of % Area in Endemicsl Taxa! Country taxa endemics endemics km2 IOKkm2 IOKkm2

Sri Lanka 561 63 11% 65610 9.6 85 .51 India 1594 288 18% 3166830 0.91 5.03 Indochina 1008 201 20% 1939320 1.04 5.2 Malesia (Asia-4) 1454 483 33% 3015250 1.6 4.82 Sub-Saharan Africa 2863 2061 72% 23284250 0.89 1.23

Table 3. Shared taxa (percentage).

Sri Lanka India Indochina Malesia Sub-So Africa

Sri Lanka 390 24% 327 32% 342 24% 141 5% India 390 69% 508 51% 453 31% 315 11% Indochina 327 58% 508 32% 556 38% 153 5% Malesia 342 61 % 453 28% 556 55% 214 7% S. S. Africa 141 25% 315 20% 153 15% 214 15%

size of the country. The larger the area, the higher the probability is of an increased number of endemics (a very small area is unlikely to have any, the whole world has 100%), and this is borne out in the '% endemics' column. However, comparing unit areas shows that is­ lands such as Sri Lanka (along with other islands (O'Shea 1997)), despite their small size, can score very highly in both relative diversity and endemism. It should be noted that the number of endemics in Africa is inflated by a large number of taxa in need of revision. The figures for numbers of taxa and endemics are based where possible on exist­ ing checklists, but in the case of Malesia and India are derived from other published litera­ ture. Those more knowledgeable about the flora of these areas may produce different fig­ ures, but it is likely that the figures will not be significantly different, and that any correc­ tions to these figures will affect the numbers of both taxa and endemics, which will not have much effect on the overall percentage.

Relationships between geographic areas The number of taxa shared between geographic units was analysed, and expressed as a percentage of the flora of each country, shown in Table 3. For instance, the number of taxa shared between Sri Lanka and India is 390, which represents 69% of Sri Lanka's flora, but only 24% ofIndia's much larger flora. To express the data graphically, the percentages were processed in a spreadsheet using Kroeber's percentage of similarity, which allows the commonality of floras to be expressed in a standard way. This method was used recently by Piippo & Koponen (1997) to study the western Melanesian moss flora, but it also included Sri Lanka as one of the elements and 298 J. Hattori Bot. Lab. No. 93 200 3

Sri Lanka-Indochina as one of the relationships, so it is interesting to compare results. Their results showed 519 taxa for Sri Lanka and 1003 for lndochina, so their Sri Lanka total was a little low, but they only found 216 taxa in common. The most likely explanation is that the synonyms in the Sri Lanka list had not been recognised and were being treated as 'endemics' and were not matching up with the same taxa (under different names) in In­ dochina. Quality of data, as discussed above, is clearly a significant factor in influencing the results. As a result, Kroeber's percentage of similarity was calculated as 31.6, much lower than 41 .1 shown in Table 4. These figures are presented graphically in Fig. 1. See Pi­ ippo & Koponen (1997) for the method of calculation.

DISCUSSION Tennent (1859) says "Gardner commented that Ceylon exhibits more of the Malaya flora and that of the Eastern Archipelago than any portion of India to the West of it", and although this may be broadly true, it is important to quantify and explain the thesis. A possible hypothesis to predict the relationships of the floras of the study area would be based on the influence of a number of factors: I. Sri Lanka's origin in , from which we might expect the flora to contain elements associated with Africa, although climatic factors as well as time may have limited

Table 4. Shared taxa (Kroeber's percentage of similarity).

Sri Lanka India Indochina Malesia Sub-Sah. Africa (SRL) (IND) (lNC) (AS4) (SUB)

Sri Lanka 46.7 45.2 42.0 14.9 India 41.1 29.7 15.4 Indochina 46.7 10.3 Malesia 1 I.1 Sub-Saharan Africa

Sri Lanka moss flora relationships

50 CII 45 +-...... -:-c-I t---"""""'--.....---l g' 40 ~ 35 I oSri Lanka l CII ~ 30 1.lndia 8. 25 , olndochina -~ 20 IOMalesia II 15 CII Sub-Saharan Africa e 10 I. lie:: 5 o ~...... ---,...... l....L.-:.r. SRL IND INe AS4 SUB Fig. 1. Graphical presentation of Kroeber's percentage of similarity. B. 1. O'SHEA: Bryogeographical relationships the mosses of Srilanka 299 this element; 2. its proximity to India, from which a strong relationship with peninsular India might be expected; and 3. an element of radiation from neighbouring areas, considering the availability of similar characteristics of geography and vegetation, and the prevailing winds. The latter element may have importance because of the post-glacial origin of much of the current vegetation. The results from the analysis are as follows: Sri Lanka shows strong relationships with India, Indochina and Malesia, as might be predicted from its geological history and current geography, but a much lower affinity with Africa. India shows a much stronger link with Sri Lanka than Indochina, and a much weaker link with Malesia. This is a surprising result, and might suggest either that the data is weak in the area of Himalayan taxa, or alternatively that the vegetation types of India, for in­ stance the general lack of evergreen rainforest, may mean that Sri Lanka, despite its small size, has a lot more in common with Indochina and Malesia than does India. Indochina has its strongest link with Malesia, as might be expected from geographic factors, but it also has a very strong link with Sri Lanka, and slightly less so with India .. Malesia (,Asia-4' of Index Muscorum), despite its very strong link with Indochina, also shows a strong link with Sri Lanka but rather less so with India. Common geographic factors may be involved, but also the prevalent winds from the east may ensure that a steady stream of spores and propagules are blown towards the , and that the climate and are more conducive to propagation than those of the rest of eastern India. The only area in southern India that might have similar qualities is on the south west coast of India, which will be in a wind shadow. Malesia is likely to have maintained a moist climate for the last 100 million years (Ashton & Gunatilleke 1987), and so will act as a reservoir of taxa favouring this climate. Sub-Saharan Africa appears to be of little influence on the Sri Lankan flora. To test this further, the Sri Lankan flora was compared with an area with which it is not likely to have any particular relationship: the United Kingdom. The Kroeber percentage of similari­ ty in this case was 5.6, around half that of Sri Lanka's similarity with Africa. Looking at the list of species Africa and the UK have in common with Sri Lanka, it is quite clear where the difference lies. The 64 species in common with the UK are almost entirely com­ mon boreal or cosmopolitan weeds-species such as Bryum argenteum and Hypnum cu­ pressiforme-whilst those species shared between Sri Lanka and Africa (apart from the weeds) are predominantly tropical, and include many interesting disjunctions. Pocs (1992) looked at the correlation between the tropical African and Asian bryofloras and concluded that the African link with Asia was much stronger than the link to America, despite the rel­ atively recent split of America from Africa (ca. 80 million years ago). For Pocs, this vali­ dated the concept of the palaeotropics as distinct from the neotropics, but it also emphasis­ es the post-Gondwanan nature of that relationship. 300 1. Hattori Bot. Lab. No. 93 200 3

• •

Fig. 2. Distribution of Campy/opus controversus (after Frahm (1994».

Fig. 3. Bryocrumia vivicolor (Broth. & Dixon) W.R.Buck (from O'Shea & Buck 2001).

Distribution of some Sri Lankan species The relationship with Africa is thus significant and still relevant, but nevertheless, what seems clear from these figures is that the primary influence on the moss flora of Sri Lanka is modern- the result of post-glacial dispersal- and that geological origin is of less influence. However, at the species level, particularly relating to non-forest taxa, it is still possible to trace the Gondwanan link (see Fig. 2). Looking at other types of distribution featuring Sri Lanka: The small moss Bryocrumia vivicolor (Broth. & Dixon) W.R.Buck (Fig. 3) of the is found in waterfalls, rocks in streams, and wet rocks in forest, but has been found only a few times (and given three different names in three different genera). The dis­ tribution seems to pick out favourable habitats at altitude (1000- 2680 m in the tropics, 400- 500 m in eastern USA) rather than representing any particular geographic or tectonic B. 1. Q ' SHEA: 8ryogeographical relationships the mosses of Sri lanka 301

c. bogoricum

Fig. 4. Clastobryophilum M.Fleisch. (after O'Shea 2000).

Fig. 5. Radulina WR.Buck & B.C.Tan (Sematophyllaceae). history. Particularly for the Indian sub-continent it has been found (usually in some quanti­ ty) in forested areas that enjoy a particularly long wet season, and it might therefore be ex­ pected in the Meghalaya area of India, and possibly elsewhere in Indochina and Malesia. Clastobryophilum bogoricum (Bosch & Sande Lac.) M.Fleisch. (Fig. 4) is a classic example of the western radiation of taxa from Malesia to Sri Lanka, and onwards to Sey­ chelles and . Radulina (Fig. 5) is a palaeotropic forest genus, often growing on rotten wood, again pinpointing moist, forested areas. However, the African population seems to be spreading rapidly, so it may be a relatively recent arrival from Asia.

How is Sri Lanka different from the rest of the Indian sub-continent? Looking at all this data in another way, we can look at differences as well as similari­ ties. Sri Lanka has 113 mosses-that is 20%-that it does not share with the rest of Asia-3 . This includes 63 endemic taxa and 4 nomina nuda, but if we exclude these, the remaining 46 taxa are shared as in Table 5. (NB. If a taxon was shared with say Europe and America, then both counts would be incre­ mented.) This emphasises just how strong the link is with Malesia, and how Sri Lanka above 302 1. Hattori Bot. Lab. No. 93 2 0 0 3

Table 5. Taxa shared with other areas.

No. of taxa Shared with

2 Europe 5 America 7 China & Japan 7 Africa 8 Oceania 8 Australasia 34 Malesia the rest of the sub-continent is in the path of radiation of taxa from this area.

CONCLUSION Looking at the results, it is clear that the moss flora of Sri Lanka has close relation­ ships with all of India, Indochina and Malesia, but much less so with Africa. Rather sur­ prisingly, Sri Lanka shows a stronger relationship with Indochina than does India: there are a number of taxa that prefer higher altitudes that are distributed across the Himalayas and into Indochina, and India shares quite a long border with Burma (Myanmar), whereas Sri Lanka and Indochina are 1600 km away from each other. The fact that Sri Lanka shares many species with Malesia that are not present in the adjacent countries suggest that Sri Lanka has both the opportunity and the environment to host Malesian species. However, the close proximity of Indochina to Malesia will also make it such a recipient, which might explain why so much is shared between Indochina and Sri Lanka. The poor state of the of many groups of tropical mosses means that analy­ ses are likely be distorted, and the number and interrelationships of the factors involved in determining the nature of the flora mean that generalisations from a limited data set are looking at only one facet of a complex set of circumstances. Nevertheless it provides a hy­ pothesis to be tested. Sri Lanka has a long history of human habitation and this has had a significant effect on the vegetation, and the increasing human popUlation will no doubt continue to degrade some of the existing areas of natural vegetation. However, all the remaining rainforest is protected, and so the wet, montane areas will survive, where the greatest diversity of bryophytes are. There is a strong tradition of conservation in the country, and the many vis­ itors to the island, who come to view the scenery, wildlife and historical buildings as well as to sit on the beach, give a great boost to the economy-so we hope the bryophytes are safe, even if at present still requiring much more study.

ACKNOWLEDGEM ENT Thanks to the anonymous reviewer who provided the reference to Ashton & Gu­ natilleke (1987). B. 1. O'SHEA: 8ryogeographical relationships the mosses of Srilanka 303

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