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The Genus Agalmyla (Gesneriaceae-Cyrtandroideae)

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The Genus Agalmyla (Gesneriaceae-Cyrtandroideae) EDINBURGH JOURNAL OF BOTANY 59 (1): 1–210 (2002) 1 DOI: 10.10M/S096042860200001X THE GENUS AGALMYLA (GESNERIACEAE-CYRTANDROIDEAE) O. M. H* & B. L. B* Agalmyla Blume (including Dichrotrichum de Vriese and Tetradema Schltr.) (Gesneriaceae-Cyrtandroideae) is revised. Ninety-seven species are recognized, 62 of them newly described, and these are divided into three sections with distinct geographical ranges. Sect. Agalmyla (24 species) is restricted to the Malay Peninsula, Sumatra, Java, Borneo and Palawan; sect. Exannularia (19 species) is the only section on Sulawesi and has a few species nearby; sect. Dichrotrichum (54 species) is represented in the Philippine Islands (excluding Palawan) by 13 species, in the Moluccas by two species, and in New Guinea by 39 species. No species is found in more than one of these areas and the key to sect. Dichrotrichum is therefore split between the Philippine species and those on the Moluccas and New Guinea. There are thus four keys altogether. The affinity and geographical distribution, both of the whole genus and of the sections, are discussed, and special attention is given to distribution of species in relation to the geological terranes of New Guinea. Points of interest and taxonomic importance in the morphological features are noted in a separate section. Keywords. Malesia, phytogeography, taxonomy. C Introduction –––––––––––––––––––––––––––––––––––––––––––––––––– 2 The taxonomic history of Agalmyla –––––––––––––––––––––––––––––– 4 The affinity of Agalmyla –––––––––––––––––––––––––––––––––––––––– 6 Morphological characters –––––––––––––––––––––––––––––––––––––– 9 Phytogeography –––––––––––––––––––––––––––––––––––––––––––––– 15 Taxonomy –––––––––––––––––––––––––––––––––––––––––––––––––– 24 The genus and its subdivision –––––––––––––––––––––––––––––––– 25 Key to sections –––––––––––––––––––––––––––––––––––––––––––– 26 Sect. Agalmyla ––––––––––––––––––––––––––––––––––––––––––––– 26 Key to species in Sundaland ––––––––––––––––––––––––––––––– 27 Sect. Exannularia ––––––––––––––––––––––––––––––––––––––––––– 69 Key to species in Lesser Sunda Islands, Moluccas, Sulawesi and West New Guinea –––––––––––––––––––––––––––––––––––––– 70 Sect. Dichrotrichum –––––––––––––––––––––––––––––––––––––––– 100 Key to species in the Philippines and Moluccas –––––––––––––– 101 Key to species in New Guinea –––––––––––––––––––––––––––– 128 (cont.) * Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh EH3 5LR, UK. Downloaded from https://www.cambridge.org/core. Laurentian University Library, on 23 Dec 2019 at 19:00:58, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S096042860200001X 2 O. M. HILLIARD & B. L. BURTT Addendum ––––––––––––––––––––––––––––––––––––––––––––––– 204 Species excludendae –––––––––––––––––––––––––––––––––––––––– 205 Acknowledgements –––––––––––––––––––––––––––––––––––––––––– 206 References ––––––––––––––––––––––––––––––––––––––––––––––––– 206 Index –––––––––––––––––––––––––––––––––––––––––––––––––––––– 208 I Agalmyla is a genus, now comprising nearly 100 species, that is found in rain forest areas from the Malay Peninsula and Sumatra eastwards to the Louisiade Archipelago at the eastern end of New Guinea. It very neatly maps out Malesia, the area covered by Flora Malesiana. The genus is treated here in the broad concept, including Dichrotrichum and Tetradema, established in earlier work (Burtt, 1968). Nothing has been discovered in the present study to suggest that either of these two genera should be restored to independent status, and the coherence of the enlarged Agalmyla has been strengthened by biochemical data ( Kvist & Pedersen, 1986 – see below under ‘The affinity of Agamyla’, p. 6, for further details). This paper is not a monograph of Agalmyla: it is an analysis and a tentative interpretation of the material to which we have had access. It aims to show the present state of knowledge and to provide a groundwork for future research. There can be little doubt that future field-work, especially in the vast, little-explored areas east of Borneo, will uncover further new species and will extend the ranges of those already recognized; field-work, however, is also needed over the whole range of the genus to observe pollination and pollinators, and to discover more about the ecology of the plants. Sometimes the seed certainly germinates on the ground and the plants sprawl unless they come into contact with a trunk suitable for climbing by their internodal roots; but does epiphytic germination also take place? Such questions cannot be answered in the herbarium. The descriptions of the species may, at first sight, seem unduly long; however, material of most species (and particularly of type specimens) is scanty, in both quantity and quality, and it is highly undesirable that anyone who studies Agalmyla should feel the need to re-dissect it. Thus the descriptions presented here (uniformly carried out by O.M.H., whose original sketches and measurements are preserved at the Royal Botanic Garden Edinburgh) serve an important conservational function. With field-work in mind we have paid special attention to geographical patterns, for it is in one or other of the areas where Agalmyla is richly developed that field studies must be made. Thus we give four separate, geographically based keys. These will almost certainly be found more useful than a single key. Fortunately the sections we have recognized have largely distinct geographies. ‘Wallace’s Line’ is mentioned several times in this paper, and it is necessary to clarify how we have used this term. A.R. Wallace (1863) recognized, in the course of his extensive travels and collecting activities in the Malay archipelago, that in the west there was a predominance of animals and plants of Indian affinity, but that as Downloaded from https://www.cambridge.org/core. Laurentian University Library, on 23 Dec 2019 at 19:00:58, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S096042860200001X THE GENUS AGALMYLA 3 he moved eastwards the numbers that belonged to groups having closer relationships with Australia increased. He came to the conclusion that the main division lay between Borneo and Sulawesi, the deep trough of the Makassar strait forming a partial barrier. At its southern end this hypothetical dividing line passed between Bali and Lombok: north of Sulawesi it swept away north-eastwards, leaving the whole of the Philippines to the west (Fig. 2). Very soon afterwards T.H. Huxley (1868) wrote on the classification and distri- bution of a group of gallinaceous birds. He recognized two major subgroups and drew a world-wide line separating the northern group from the southern. He pointed out that in SE Asia ‘the boundary in question would coincide with what may be called ‘‘Wallace’s Line’’ between the Indian and Papuan divisions of the Malay archipelago. But it would run northwards as far as the Philippines, and, passing between them and Formosa, would tend southward and eastward to the Samoan archipelago’. Huxley’s map (op. cit. p. 294) shows that he was proposing exactly the same modification of Wallace’s Line as that suggested by Dickerson & Merrill (Merrill, 1923a, 1926) based on plant distributions, particularly those of Dipterocarpaceae. Huxley left Palawan, the south-westernmost of the Philippines, west of his version of the line, as did Dickerson & Merrill in theirs. We refer to this as the modified F. 2. Map of SE Asia showing Wallace’s Line, modified Wallace’s Line and Lydekker’s Line. Downloaded from https://www.cambridge.org/core. Laurentian University Library, on 23 Dec 2019 at 19:00:58, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S096042860200001X 4 O. M. HILLIARD & B. L. BURTT Wallace’s Line (Fig. 2), and that is the form of it used throughout this paper. It has been chosen simply because it provides the best fit with the classification of Agalmyla. A recent study in Cyrtandra (another genus of Gesneriaceae) was designed to test the validity of Huxley’s modification using molecular methods (Atkins et al., 2001). It was found that only two of the nine species known from Palawan belonged to the clade including species from Borneo, the remaining seven belonging to a distinct clade of Philippine species. In Cyrtandra, therefore, it seems that the modification is not justified (always remembering that the samples of Bornean and Philippine species were small and random). In Epithema, a much smaller genus of Gesneriaceae that we have studied morphologically, the Palawan material seen belongs to a species from N Borneo, whereas elsewhere in the Philippines one of the species is E. benthamii whose distribution includes N Sulawesi and the Vogelkop Peninsula of westernmost New Guinea. It seems likely that north of the Makassar strait the position of a ‘line’ will be found to vary from group to group. Wallace himself may have come to the same view, for in his later work Island Life (we quote from the second edition, Wallace, 1892: 387–388) he uses a list of Philippine mammals from which ‘the Palawan group has been omitted as these islands contain so many Bornean species that if included they obscure the special features of the [Philippine] fauna’. Despite this treatment for mammals, Wallace may not have been convinced of the general validity of Huxley’s modification, for he made no change in the position of the line in his map of the zoological regions
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