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A review of dipterocarps - Center for International Forestry Research

A review of dipterocarps - Center for International Forestry Research

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Biogeography and Evolutionary Systematics <strong>of</strong> Dipterocarpaceae<br />

Asian <strong>dipterocarps</strong> deeply imprint the <strong>for</strong>est ecology<br />

and economy <strong>of</strong> the places where they grow. They<br />

constitute prominent elements <strong>of</strong> the lowland rain <strong>for</strong>est<br />

(Whitmore 1988) and are also well represented in the<br />

understorey. As a family they dominate the emergent<br />

stratum. Most belong to the mature phase <strong>of</strong> primary<br />

<strong>for</strong>est, which contains most <strong>of</strong> the entire genetic stock<br />

(Jacobs 1988). All species can colonise secondary<br />

<strong>for</strong>ests during the succession phases provided there is a<br />

seed source; seed dispersal is limited, except among<br />

water dispersed species. However, none seems presently<br />

confined to secondary <strong>for</strong>mations. Certain <strong>dipterocarps</strong><br />

<strong>of</strong> the seasonal regions dominate the fire-climax<br />

deciduous <strong>for</strong>ests <strong>of</strong> northeast India and Indo-Burma.<br />

In Asia, <strong>dipterocarps</strong> occupy a large variety <strong>of</strong> habitats<br />

(Symington 1943, Wyatt-Smith 1963) from coastal to<br />

inland, riverine to swampy and to dry land, undulating to<br />

level terrain, ridges, slopes, valley bottoms, soils deeply<br />

weathered to shallow, well-drained to poorly drained, and<br />

rich to poor in nutrients. In Peninsular Malaysia the<br />

altitudinal zonation <strong>of</strong> their main habitat types ranges<br />

from 0-300 m (low-undulating dipterocarp <strong>for</strong>est), 300-<br />

750 m (hill dipterocarp <strong>for</strong>est), and 750-1200 m (upper<br />

dipterocarp <strong>for</strong>est). Zonation however, differs in Borneo<br />

and Sri Lanka. The freshwater swamps, especially in drier<br />

parts, are rich in species (Corner 1978, in Jacobs 1988)<br />

while true peat-swamp is relatively poor. The dipterocarp<br />

flora is also poor on limestone and riverine fringes.<br />

Asian <strong>dipterocarps</strong> are limited altitudinally<br />

(Symington 1943) by climatic conditions, and the<br />

conjunction <strong>of</strong> altitude and other natural barriers, such<br />

as large rivers and watersheds, have obstructed the<br />

distribution <strong>of</strong> species in Borneo. For example, the<br />

northwest and northeast <strong>of</strong> Kalimantan, Sarawak, Brunei<br />

and Sabah are much richer in species than the rest <strong>of</strong><br />

Kalimantan. The everwet areas are also richer in species<br />

than the seasonal ones as shown in Sri Lanka by the<br />

concentration <strong>of</strong> species in the southwest quarter, or in<br />

the Thai-Malaysian transition belt, or from Java to the<br />

Lesser Sundas (Jacobs 1988).<br />

Distribution <strong>of</strong> Dipterocarps and<br />

Related Taxa<br />

The present distribution patterns <strong>of</strong> <strong>dipterocarps</strong> are<br />

thought to reflect routes <strong>of</strong> colonisation and past climatic<br />

conditions (Fig. 1). Living <strong>dipterocarps</strong> sensu lato are<br />

spread over the tropical belt <strong>of</strong> three continents <strong>of</strong> Asia,<br />

Africa and South America. They occupy several<br />

12<br />

phytogeographical zones that mainly con<strong>for</strong>m to climatic<br />

and ecological factors. However, in southeast Asia,<br />

Wallace’s line where it runs east <strong>of</strong> the Philippines and<br />

between Borneo and Celebes, is a major phytogeographic<br />

boundary <strong>for</strong> <strong>dipterocarps</strong>. It cannot be explained in terms<br />

<strong>of</strong> climatic differences but requires the intervention <strong>of</strong><br />

continental shelf drift.<br />

Phytogeographical Regions <strong>of</strong> Living Taxa<br />

The South American region (Fig. 1, Table 4) corresponds<br />

to Guyana, Venezuela and the part <strong>of</strong> Colombian Amazon<br />

which overlies the Guyana shield.<br />

The African region (Fig. 1, Table 4) includes a<br />

continental area and an insular part in Madagascar. The<br />

<strong>for</strong>mer is in two disjunct areas (Aubreville 1976): a) a<br />

narrow strip in the northern hemisphere from Mali on<br />

the west, to Sudan on the east, neither reaching the<br />

Atlantic nor the Indian Ocean; and b) in the southern<br />

hemisphere the Monotes-Marquesia area covers a semidry<br />

region between the two oceans, south <strong>of</strong> the<br />

Congolese rain <strong>for</strong>est, most <strong>of</strong> which is essentially<br />

central and does not reach the Atlantic or Indian Oceans.<br />

The Asian region (Fig. 1, Table 4) corresponds to the<br />

Indo-Malesian area, which concentrates a high number<br />

<strong>of</strong> genera and species in the equatorial <strong>for</strong>ests. This area<br />

is limited northward by the Himalayan foothills, then<br />

approximately by the borders <strong>of</strong> Assam, Arunachal<br />

Pradesh (India), Burma, Laos and Vietnam, and<br />

penetrating into south China including Hainan Island. On<br />

the extreme southwest the large belt <strong>of</strong> Asian<br />

<strong>dipterocarps</strong> reaches the Seychelles (1 sp. Vateriopsis<br />

seychellarum), and covers India and Sri Lanka. Its eastern<br />

border corresponds to New Guinea. The Sundalands<br />

delimit the most southern part. No dipterocarp species<br />

is found in Australia.<br />

Five main phytogeographical regions are classically<br />

recognised within this distribution area: 1) Malesia:<br />

Peninsular Malaysia, Sumatra, Java, Lesser Sunda Islands,<br />

Borneo, the Philippines, Celebes, the Moluccas, New<br />

Guinea and the Bismarks. The northern frontier <strong>of</strong><br />

Peninsular Malaysia delimits this part; 2) Mainland<br />

southeast Asia: Burma, Thailand, Cambodia, Laos,<br />

Vietnam and south China (Smitinand 1980, Smitinand et<br />

al. 1980, 1990); 3) south Asia: India, Andaman islands,<br />

Bangladesh, Nepal; 4) Sri-Lanka; and 5) Seychelles. In<br />

these Asian phytogeographical areas each dipterocarp<br />

group manifests a more or less distinctive pattern <strong>of</strong><br />

variation at the species level (Ashton 1982).

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