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Patterns of Distribution of Malesian Vascular Plants

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Patterns of Distribution of Malesian Vascular Plants Malesian plant distributions 243 Patterns of distribution of Malesian vascular plants W J Baker1, M J E Coode, J Dransfield, S Dransfield, M M Harley, P Hoffmann and R J Johns The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, UK 1Department of Botany, Plant Science Laboratories, University of Reading, Whiteknights, Reading, Berkshire, RG6 6AS, UK Key words: biogeography, phytogeography, palynology, SE Asia, Malesia, Palmae, Gramineae, Euphorbiaceae, Elaeocarpaceae, Antidesma, Elaeocarpus, Nypa, Spinizonocolpites Abstract analytical phase Biogeographical work con- cerned with the analytical phase has appeared A miscellaneous selection of Malesian plant distributions is increasingly in the systematic literature and it is presented, including examples from the Palmae, here that modern methods are most evident Gramineae, Euphorbiaceae, Elaeocarpaceae, and various fern genera Hypotheses of the tectonic evolution of the Previously, most classifications have been based area may be required to explain many of the observed pat- on intuition and overall similarity which, though terns that are described Two major distribution types are they may stand the test of time, are nevertheless identified repeatedly, the first displaying a strongly Sundaic subjective Despite the introduction of statistical bias and the second focusing on E Malesia Patterns involv- techniques which aimed to make similarity- ing New Guinea are complex as they tend to include a vari- able combination of other islands such as Sulawesi, Maluku, based or phenetic classifications more testable, the Bismarck archipelago and the islands of the W Pacific A theoretical objections have led to the decline of number of striking disjunctions exist, some of which have phenetics in favour of cladistic methods relatively narrow overall ranges, such as those of the palm Cladistics uses character patterns of extant or- genera Cyrtostachys and Rhopaloblaste which occur across ganisms to construct a hypothesis of their evolu- Malesia excluding central parts of the region In other exam- tionary or phylogenetic relationships These pat- ples, however, the separation is more profound including disjunctions between parts of Malesia and Madagascar in terns are generally visualised in the form of Orania (Palmae) and Nastus (Gramineae), and between branching diagrams or cladograms which can be Fiji, Vanuatu and Palawan in Veitchia (Palmae) At this interpreted appropriately stage, the significance of these distributions for the under- This paper is concerned mainly with the nec- standing of the geological history of SE Asia remains un- essary precursors to the analytical phase, the clear It is noted that distributions of species from the genus Antidesma (Euphorbiaceae) are more easily explained in descriptive and narrative phases of biogeogra- terms of dispersal and environmental factors Formal phy In selecting taxa for discussion we have cladistic biogeographic analyses of these and other groups chosen only those which we believe will prove should aid interpretation of the regions history to be sound when subjected to the rigours of cladistics The only groups which are acceptable in the cladists eyes are those described as Introduction monophyletic A monophyletic (natural) group contains all the descendants of a common an- Over the last four decades, there has been a cestor and is defined by shared characters, or radical change in the methods used by systema- synapomorphies, that support a specific node tists to classify organisms This change has had a on a cladogram A group which contains some, concurrent effect on biogeographical methods but not all descendants of a common ancestor Ball (1975) describes three phases in biogeo- may appear to be coherent, but cannot be de- graphical studies which he calls the descriptive fined in the same way and, indeed, it can be or empirical phase, the narrative phase and the argued that it cannot be defined at all These Biogeography and Geological Evolution of SE Asia, pp! 243-258 Edited by Robert Hall and Jeremy D! Holloway © 1998 Backhuys Publishers, Leiden, The Netherlands 244 W- J- Baker et al- paraphyletic groups are a partial and arbitrary represented in the region except for the expression of the product of common ancestry Phytelephantoideae and make no sense in an evolutionary context Palm distributions in the region have been The inclusion of non-monophyletic groups in discussed previously in an anecdotal fashion biogeographical studies is flawed for these rea- (Dransfield, 1981, 1987; Uhl and Dransfield, sons For basic introductions to cladistic theory, 1987) but these discussions lack a firm cladistic general overviews may be found in Patterson basis However, with the explicit analytical (1982), Wiley et al- (1991) and Scotland (1992) framework that we are developing, we have There is a substantial literature describing been able to select a number of genera or plant distributions in Malesia which we do not groups of genera that we believe to be mono- summarise here Instead we present distribu- phyletic and display distribution patterns of con- tions from a miscellany of vascular plant fami- siderable biogeographic interest lies Firstly, two monocotyledonous families are We present examples from three subfamilies: discussed; from the palm family, generic distri- Nypoideae, Calamoideae and Arecoideae As butions within the three subfamilies Nypoideae, mentioned above, the last subfamily is probably Calamoideae and Arecoideae, and from the paraphyletic but the examples we have chosen grass family, distributions within the two bam- from within it are, we believe, monophyletic boo genera, Dinochloa and Nastus Secondly, in the dicotyledons, the genera Antidesma (Eu- phorbiaceae) and Elaeocarpus (Elaeocarpaceae) Nypoideae are considered The fern genus Christensenia (Marattiaceae) and a variety of other fern and This is the only subfamily in the Palmae repre- angiosperm examples from the diverse flora of sented by a single extant species Nypa fruticans New Guinea complete the assortment The term is distinguished from all other palms by a Malesia is used in this paper to identify the area number of unique features including the pros- bounded to the north by the Thai-Malaysian trate dichotomously branched stem, the erect border and to the south by the Torres Straits inflorescence bearing a terminal head of pistil- (Steenis, 1950; Johns, 1995) It includes the fol- late flowers, the lateral spikes of staminate flow- lowing political entities: Brunei Darussalam, In- ers and unusual features of the flowers them- donesia, Malaysia, Papua New Guinea, Philip- selves (Uhl and Dransfield, 1987) Furthermore, pines and Singapore It is an area of exceptional the spiny zonosulcate pollen grains are highly biological diversity distinctive With the exception of similar but much smaller and incompletely zonosulcate pollen produced by many species of the genus Palmae Salacca (Calamoideae), such pollen grains are not encountered elsewhere in the family, and The Palmae (Arecaceae) are a diverse, largely are unknown outside the Palmae The pollen tropical and subtropical family of about 200 gen- grains are notably uniform in their morphology era and 2700 species In the latest classification throughout the geographic range of the genus of the family (Uhl and Dransfield, 1987) six sub- In a recent phylogenetic study of the whole fam- families are recognised The delimitation of ily, Nypa is resolved as sister taxon to all other these subfamilies is currently under rigorous palms (Uhl et al, 1995) phylogenetic analysis, and preliminary evidence Nypa fruticans is a mangrove palm, often suggests that four of the six (Coryphoideae, Ca- growing in vast natural stands in a variety of es- lamoideae, Nypoideae and Phytelephantoideae) tuarine conditions Its present day distribution are monophyletic whereas two, Ceroxyloideae extends throughout Malesia and also includes and Arecoideae, may be paraphyletic (Uhl et al-, Sri Lanka, the Ganges delta, Indochina, NW Aus- 1995) However, the relationships of many of tralia, the Solomon Islands and Ryukyu Islands the tribes, subtribes and genera within the sub- (Uhl and Dransfield, 1987) It has also been in- families need further study troduced to W Africa and to Panama where it is The Malesian and W Pacific region represents now well established one of the richest areas of palm diversity in the Unlike its modern counterpart, fossil Nypa world An estimated 1000 species in 93 genera pollen, Spinizonocolpites (Muller, 1968), has are found in the area from Indochina to Australia been shown to possess a range of spine lengths and the W Pacific islands of Fiji and New Cal- and spine distributions This variation has been edonia All subfamilies mentioned above are demonstrated by a number of authors to be very Malesian plant distributions 245 4 Ma 15 Ma 45 Ma Fig!1 Distribution maps for Spinizonocolpites generated using Atlas palaeomapping program (Cambridge Paleomap Services, 1992), incorporated into Plant Fossil Record 22 (Lhotak and Boulter, 1995) 246 W- J- Baker et al- Calamus Calospatha Ceratolobus Daemonorops Myrialepis Plectocomia Plectocomiopsis Pogonotium Retispatha Fig!2 Distributions of genera in subfamily Calamoideae (Palmae), rattans localised, often occurring in quite small matrix grains from offshore cores in the West Java Sea samples (e-g-, Harley et al-, 1991) The distribu- (R J Morley, pers comm, 1997) By the end of tion of Spinizonocolpites in the fossil record is the Miocene (Fig1A), the distribution did not
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