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A review of dipterocarps - Center for International Forestry Research

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

Tertiary dipterocarp fossils have been reported from<br />

the African Miocene <strong>of</strong> Ethiopia (Beauchamp et al.<br />

1973, Lemoigne 1978, Laveine et al. 1987) and from<br />

the putative (but probably earlier) Plio-Pleistocene <strong>of</strong><br />

Somalia (Chiarugi 1933). That is in a continent where<br />

not a single living dipterocarpoid has been found, and<br />

where present monotoids are living. Lemoigne (1978,<br />

p.123) specifies ‘c’est avec des bois de la famille des<br />

Dipterocarpaceae, notamment ceux du genre Monotes<br />

que notre echantillon parait avoir le plus d’affinites...<br />

Certes les affinites avec la famille des Lauraceae sont<br />

aussi remarquables’. In spite <strong>of</strong> its name<br />

Dipterocarpoxylon, this fossil is thus <strong>of</strong> a Monotoid<br />

type (not Dipterocarpoid). In this case, is the African<br />

rain <strong>for</strong>est species Marquesia excelsa derived from a<br />

common ancestor with Monotes and adapted to a more<br />

humid climate, or is it the only surviving species <strong>of</strong> some<br />

Dipterocarpoid ancestor which could have fossilised in<br />

Ethiopia and Somalia (and Egypt?)? A study <strong>of</strong> the pollen<br />

exine structure in Marquesia is needed to clarify this<br />

genus situation, as well as a critical re-examination <strong>of</strong><br />

all dipterocarpoid fossils (doubtful or not).<br />

Numerous accepted fossils from Asia (Awasthi 1971)<br />

testify that the present great species richness <strong>of</strong> the Asian<br />

flora (Table 6) probably existed since the Miocene and<br />

persists through the Pliocene and Pleistocene, up to the<br />

Quaternary (Anisopteroxylon, Dipterocarpoxylon,<br />

Dryobalanoxylon, Hopenium, Shoreoxylon,<br />

Vaticoxylon, Vaterioxylon).<br />

These fossils demonstrate a reduction <strong>of</strong> dipterocarp<br />

distribution area both in Africa (Fig. 1, Tables 5 and 6)<br />

and Asia (extinction <strong>of</strong> Anisoptera and Dryobalanops<br />

in India, and <strong>of</strong> the latter in Indo-China), and total<br />

extinction in Europe and North America (doubtful<br />

fossils?). Could thus the Tertiary distribution area <strong>of</strong><br />

<strong>dipterocarps</strong> sensu stricto include Africa and Asia (and<br />

Europe?)?<br />

Hypotheses on the Geographical Origin <strong>of</strong><br />

Dipterocarpaceae<br />

If Monotoideae and Pakaraimaea are to be connected<br />

to Asian <strong>dipterocarps</strong> (by a single family or into different<br />

families), a common ancestor and its migration path have<br />

to be found. During the transition from the later<br />

Cretaceous to the very early Eocene the paleogeographic<br />

changes, in combination with other effects, could have<br />

produced the present geography. Thus dipterocarp<br />

ancestors should have been present when land<br />

17<br />

connections still existed between South America, Africa<br />

and India and between them and southeast Asia (and<br />

probably with the European and north American Laurasia<br />

block with its intermittent ‘Grande coupure’). This<br />

situation occurred (Figs. 2 (A, B), 3) in the Permo-<br />

Triassic period. Later, parts <strong>of</strong> the northeastern Gondwana<br />

land detached from the Gondwana shelf, crossed the<br />

Tethys and joined southeast Laurasia (just as India would<br />

do later). These changes would have happened during the<br />

Permo-Triassic, Jurassic and Cretaceous times according<br />

to recent works on Cathaysian floras (Kovino 1963,<br />

1966, 1968, all in Vozenin-Serra and Salard-Cheboldaeff<br />

1994, Lemoigne 1978, Jaeger et al. 1983, Vozenin-Serra<br />

1984, Vozenin-Serra and Taugourdeau-Lantz 1985,<br />

Laveine et al. 1987, Taugourdeau and Vozenin-Serra<br />

1987, Renous 1989, Scotese and McKerrow 1990 in<br />

Vozenin-Serra and Salard-Cheboldaeff 1994), and on the<br />

Tethys Sea (Dercourt et al. 1992).<br />

Croizat (1964, 1952 in Aubréville 1976) and Ashton<br />

(1969) expressed the view <strong>of</strong> a dipterocarp Gondwanan<br />

origin <strong>of</strong> the present distribution area and to a further<br />

migration towards Indo-Malesia. Aubréville (1976)<br />

considered that Dipterocarpaceae probably occupied two<br />

main areas be<strong>for</strong>e the Cretaceous general drift <strong>of</strong><br />

Gondwanan shelves: one in Asia and one in the joined<br />

Africa-India-Seychelles-Sri Lanka complex. He believed<br />

that the origin was in Europe from where ancestors <strong>of</strong><br />

monotoids would have migrated towards Africa and<br />

further from there to India. He suggested two Tertiary<br />

centres <strong>of</strong> <strong>dipterocarps</strong>: one Indo-Malesian from<br />

Laurasian origin; and one Africano-Indian from a<br />

Gondwanan origin, on both sides <strong>of</strong> the Tethys Sea. More<br />

recent studies (Renous 1989, Dercourt et al. 1992,<br />

Vozenin-Serra and Salard-Cheboldaeff 1994) identify<br />

direct land connections between southeast Asia and<br />

Laurasia lands. These authors consider a possible series<br />

<strong>of</strong> small blocks detached from the northeastern part <strong>of</strong><br />

Gondwana, moving through the Tethys, and <strong>for</strong>ming an<br />

archipelago (Fig. 3: Tazrim-Sino-Korean block (1), north<br />

China block (2), north Tibet block (3), Khorat-Kontum<br />

block: east Thailand and most parts <strong>of</strong> Laos, Cambodia<br />

and Vietnam (4), south Tibet block (5), Kashmir block<br />

(6), Iran-Afghanistan block (7), Turkey block (8), Spain<br />

block (9)) together with the different southeast Asian<br />

plates. This putative archipelago would have served as a<br />

relay, perhaps owing to volcanism which would open<br />

routes <strong>for</strong> floral migration. Be<strong>for</strong>e the rise <strong>of</strong> the<br />

Himalayas, floristic exchanges would also have been<br />

possible around the Tethys (Dercourt et al. 1992).

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