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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 />

perhaps allow rapid spread <strong>of</strong> favourable ecotypes in<br />

heterogeneous terrain. However apomixis should<br />

probably lead to lowered genetic variability within<br />

populations, and thus could still result in increased<br />

probability <strong>of</strong> extinction when unadapted to the<br />

environmental changes. Constant differences in habit,<br />

morphology and reproductive biology exist between<br />

emergent and understorey trees (Richards 1952, Hallé<br />

et al. 1978, Yap 1982 in Ashton 1984). Ontogenesis thus<br />

follows predictable patterns in the larger trees and<br />

selection acts on these characteristics, which include tree<br />

habit and leaf shape, irrespective <strong>of</strong> their systematic<br />

relationships. These species present an ecological<br />

complementarity. It is however, not certain that species<br />

sharing a common habitat and geography will identically<br />

respond from flower-bud initiation to sensitivity to light,<br />

mycorrhizal invasion, water stress, seed predation, or<br />

share the same pollinators and seed vectors (M.S. Ashton<br />

1992, 1995). Ecological criteria such as seed water<br />

content and seed resistance to desiccation are<br />

expressions <strong>of</strong> the species’ biological plasticity and the<br />

possible complementarity between species (Maury-<br />

Lechon 1993). Certain species are abundant and others<br />

rare. Detailed systematic and biosystematic comparisons<br />

between rare and abundant congeners should bring some<br />

answers (Ashton 1992). We need to confirm whether<br />

the means exist <strong>for</strong> gene flow, and then to directly<br />

measure the level and pattern <strong>of</strong> genetic variability.<br />

Amenable source <strong>of</strong> evidence could also be tested <strong>for</strong><br />

the presence or absence <strong>of</strong> associations between<br />

population distributions in space, and demographic and<br />

population genetic studies are also required on the basis<br />

<strong>of</strong> repeated observations. These observations and the use<br />

<strong>of</strong> species’ complementarity would rein<strong>for</strong>ce the<br />

understanding <strong>of</strong> systematics and help the <strong>for</strong>est<br />

managers to make decisions in rehabilitation and<br />

conservation programmes (Maury-Lechon 1991, 1993,<br />

1996).<br />

Bearing in mind the economic value and the present<br />

status <strong>of</strong> Dipterocarpaceae, it is also urgent to relate<br />

phylogeny to comparative ecology within genera and<br />

sections by a combination <strong>of</strong>:<br />

1. molecular phylogenic studies, concentrating on genera<br />

and below;<br />

2. comparative demographic studies <strong>of</strong> groups <strong>of</strong> related<br />

species, especially those which co-occur:<br />

a) during the reproductive phase (bud to recruit, and<br />

including fecundity),<br />

b) during stand development and trough to mortality,<br />

35<br />

c) population genetics <strong>of</strong> selected species under selected<br />

conditions;<br />

3. comparative ecophysiological experiments on seeds<br />

and seedlings; and<br />

4. competition experiments.<br />

The teams exist within the frame <strong>of</strong> the <strong>International</strong><br />

Working Group on Dipterocarps (IWGD-IUFRO S.07-<br />

17 Working Party) and contacts have already been taken<br />

between the authors <strong>of</strong> this paper and their direct<br />

commentators <strong>for</strong> this purpose. Field and laboratory<br />

works will be organised on the base <strong>of</strong> complementarity.<br />

Overall, a cooperative, integrated and detailed reassessment<br />

is needed <strong>for</strong> the whole family<br />

Dipterocarpaceae sensu lato, with the establishment <strong>of</strong><br />

an evolutionary classification based on a general<br />

consensus. Several groups around the world are currently<br />

working on projects that can lead to such a solution. They<br />

include: Forest <strong>Research</strong> Institute Malaysia, Kepong and<br />

Unité Mixte de Recherche 5558 du Centre National de<br />

la Recherche Scientifique with Lyon University (France),<br />

in association with Harvard University (USA), who are<br />

working on the genetic analysis <strong>of</strong> DNA sequences;<br />

Massachusetts University (USA) on genetic and breeding<br />

system studies (Murawski and Bawa 1994, Murawski et<br />

al. 1994); and Cambridge-Edinburgh (U.K.) on computer<br />

identification keys (Newman et al. 1995). Several more<br />

complementary works are needed in palynology (pollen<br />

and exine), stamen architecture and shape (the flower<br />

being observed from the pollination point <strong>of</strong> view),<br />

ontogenesis (structure and morphology <strong>of</strong> fruit-embryogermination,<br />

anatomy <strong>of</strong> cotyledonary node and petiole,<br />

epidermis <strong>of</strong> primordial leaves), wood anatomy,<br />

chemotaxonomy and architecture <strong>of</strong> juvenile stage. These<br />

complementary works will require broad cooperation<br />

with colleagues and institutions from the Asian, African<br />

and South American zones. More ef<strong>for</strong>t is needed <strong>for</strong><br />

the Asian <strong>dipterocarps</strong> from China, Burma and Indo-<br />

China. Likewise, there is a need <strong>for</strong> contact with African<br />

colleagues<br />

Acknowledgements<br />

We express our sincere thanks to the <strong>Center</strong> <strong>for</strong><br />

<strong>International</strong> <strong>Forestry</strong> <strong>Research</strong> and most particularly to<br />

C. Cossalter <strong>for</strong> having initiated and supported the<br />

realisation <strong>of</strong> a book which provides the first broad<br />

synthesis <strong>of</strong> the present status <strong>of</strong> knowledge on<br />

<strong>dipterocarps</strong> that complements the proceedings <strong>of</strong> the

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