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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 />

These features group together the living and fossil<br />

<strong>dipterocarps</strong> which have lived together in the same<br />

phytogeographical areas (Table 5). They also correspond<br />

to the subsequent hypothesis concerning their potential<br />

<strong>for</strong> differentiation (see above). They also suggest an<br />

eventual remote relation from Anthoshorea to<br />

Marquesia and then to Monotes.<br />

In the Valvate division the Dipterocarpinae group<br />

underlines the relations between the 2 subgroups: 1)<br />

Dryobalanops, and 2) Dipterocarpus with Anisoptera.<br />

Similarly the Vaticinae group emphasises the<br />

existence <strong>of</strong> 5 subgroups:<br />

1. Upuna alone but near subgroup 2;<br />

2. Cotylelobium close to Sunaptea;<br />

3. Vatica pro-parte intermediate between groups 1 and<br />

2, and 4 and 5;<br />

4. Stemonoporus position between subgroups 2 and 5;<br />

5. Vateria, and Vateriopsis with particular similarities<br />

(cotyledon position and shape, germination type) with<br />

Vatica and also Anisoptera, Stemonoporus and<br />

Cotylelobium.<br />

Vatica genus (excluding Sunaptea) stands somewhat<br />

isolated within the family Dipterocarpaceae by the pollen<br />

characters principally, and much less so by some aspects<br />

<strong>of</strong> embryo shape and structure (particularly seedling<br />

vascular structure). As mentioned, the tilioid surface <strong>of</strong><br />

the pollen exine could suggest a proximity with<br />

monotoid taxa, however, its structure is definitely<br />

different (Maury et al. 1975a, b). New investigations are<br />

needed on a great number <strong>of</strong> species <strong>of</strong> Vatica genus<br />

sensu lato (including Sunaptea and Pachynocarpus) to<br />

permit a clearer view on infra-generic variation <strong>of</strong> these<br />

characters.<br />

Main lines <strong>of</strong> Kostermans’ classification (See also<br />

Tables 1, 2)<br />

Kostermans has mainly considered the Sri Lankan taxa<br />

so that, as in Meijer’s work, only the Asian genera<br />

represented in this geographical area were analysed in<br />

detail. Contrary to Maury-Lechon’s work he <strong>for</strong>mally<br />

described the Monotaceae family, as well as genera<br />

Doona and Sunaptea (the latter including<br />

Cotylelobium). No publication remains on his views<br />

concerning the affinities <strong>of</strong> the genera inside his<br />

Dipterocarpaceae family, nor in eventual sections within<br />

the Shorea genus which includes Pentacme (Kostermans<br />

1992). In Stemonoporus he suggests 2 sub-divisions<br />

based on the pericarp aperture at germination (character<br />

used in Maury 1978 and Maury-Lechon 1979a, b).<br />

Taxonomical levels<br />

Family (2):<br />

Monotaceae (3 genera): Pakaraimaea,<br />

Marquesia, Monotes<br />

Dipterocarpaceae (15 genera): Hopea,<br />

Neobalanocarpus, Balanocarpus, Shorea,<br />

Doona, Parashorea, Dipterocarpus,<br />

Anisoptera, Dryobalanops, Upuna,<br />

Sunaptea (Cotylelobium included), Vatica,<br />

Stemonoporus, Vateria, Vateriopsis<br />

33<br />

Main Recent Taxonomic Changes:<br />

They successively concerned the:<br />

1. establishment <strong>of</strong> subgenera Shorea, Anthoshorea,<br />

Richetia and Rubroshorea (Meijer 1963, Meijer and<br />

Wood 1964);<br />

2. establishment <strong>of</strong> 11 sections in genus Shorea including<br />

the previous genera Doona and Pentacme (Ashton<br />

1964, 1968, 1980, 1982);<br />

3. proposition to re-install some ancient genera such<br />

as Doona Thw., Anthoshorea Heim and Richetia<br />

Heim outside genus Shorea Gaertn., Sunaptea Griff.<br />

outside genus Vatica and Vateriopsis Heim out <strong>of</strong><br />

genus Vateria L. (Maury 1978, Maury-Lechon<br />

1979a, b);<br />

4. acceptance <strong>of</strong> the re-establishment <strong>of</strong> Vateriopsis<br />

genus by Ashton (1982);<br />

5. announcement <strong>of</strong> the discovery and description <strong>of</strong><br />

Pakaraimaea (Maguire 1979, Maguire and Ashton<br />

1980, Maguire and Steyermark 1981);<br />

6. <strong>for</strong>mal re-establishment <strong>of</strong> genus Doona Thw. outside<br />

Shorea (Kostermans 1984), genus<br />

Banalocarpus Beddome outside Hopea and independent<br />

from Neobalanocarpus (Kostermans<br />

1981a) genus Sunaptea outside genus Vatica but including<br />

genus Cotylelobium (Kostermans 1987); and<br />

7. discovery and description <strong>of</strong> Pseudomonotes<br />

tropenbosii (Londoño et al. 1995) which is included<br />

in Monotoideae sensu Maguire et al. (1977) close<br />

to the African Monotes and Marquesia.<br />

Discussion and Conclusions<br />

Morphology, as well as anatomy and ecophysiology,<br />

shows many characters tightly related to their biological<br />

functions, and these functions are connected to both the<br />

biotic associations and the climatic environmental<br />

features which influence flower pollination, seed

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