A review of dipterocarps - Center for International Forestry Research

Biogeography and Evolutionary Systematics of Dipterocarpaceae
them for several hundred metres and sometimes about
one kilometre. In the open dry deciduous forest of the
seasonal Asian regions the dispersal of dipterocarp
winged fruits becomes much more efficient. However,
occasional dispersal over long distances has probably
succeeded, judging from the present high diversity of
Asian dipterocarp habitats in aseasonal regions.
Classification
The full set of taxonomic and systematic works available
has been reported and discussed in the more recent flora
books, monographs, theses and other publications of
Ashton (1962, 1963, 1964, 1967, 1969, 1972, 1977,
1979b, c, d, 1980, 1982), Bancroft (1933, 1934, 1935a,
b, c, d, e, 1936a, b, 1939), Bisset et al. (1966, 1967,
1971), Damstra (1986), Diaz and Ourisson (1966), Diaz
et al. (1966), Gottwald and Parameswaran (1964, 1966,
1968), Jacobs (1981), Jong and Lethbridge (1967), Jong
and Kaur (1979), Jong (1976, 1978, 1979, 1980, 1982),
Kochummen (1962), Kochummen and Whitmore
(1973), Kostermans (1978, 1980, 1981a, b, c, 1982a,
b, 1983, 1984, 1985, 1987, 1988, 1989, 1992), Maguire
(1979), Maguire et al. (1977), Maguire and Ashton
(1980), Maguire and Steyermark (1981), Maury et al.
(1975a,b), Maury 1978, Maury-Lechon (1979a, b, 1982
in Ashton: p. 265-266 ripe embryo, germinating
seedlings and 268 palynology, 1985 in FAO: palynology,
Maury-Lechon and Ponge (1979), Meijer (1963, 1968,
1972, 1973, 1974a, b, 1979), Meijer and Wood (1964,
1976), Ourisson (1979), Parameswaran (1979a, b),
Parameswaran and Gottwald (1979), Parameswaran and
Zamuco (1979, 1985 in FAO), Rojo (1976, 1977, 1979,
1987), Rojo et al. (1992), Sidiyasa et al. (1986, 1989a,
b), Smitinand (1958a, b, 1979, 1980), Smitinand and
Santisuk (1981), Smitinand et al. (1980, 1990), Sukwong
(1981), Sukwong et al. (1975), Tao and Tong (1982),
Tao and Zhang (1983), Tao and Dunaiqiu (1984), Tewary
(1984), Tewary and Sarkar (1987a, b), Verdcourt (1989),
Vidal (1960, 62, 79), Whitmore (1962, 1963, 1976,
1979, 1988), Wildeman (1927), and Wildeman and
Staner (1933).
Affinities of the Dipterocarpaceae Family
Hutchinson (1959, 1969 in Maury 1978) put the family
in the order Ochnales and later suggested a phylogenetic
location of the order at the end of a series whose
progressive steps were ordered as follows: Magnoliales,
Dilleniales, Bixales, Theales and Ochnales.
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Cronquist (1968) moved Dipterocarpaceae into the
order Theales and Takhtajan (1969) regrouped Ochnales,
Theales and Guttiferales in Theales and placed
Dipterocarpaceae under it near Lophiraceae and
Ancistrocladaceae. Dalgren (1975) placed the
dipterocarp family in the order Malvales under
superorder Dilleniflorae.
After the description and detailed study of
Pakaraimaea dipterocarpacea and its inclusion in
Dipterocarpaceae, Ashton removed the family from
Guttiferales and considered the imbricate calyx of
Dipterocarpaceae and Sarcolaenaceae (Maguire et al.
1977) as an isolated and derived character among
otherwise Malvalian features, and hence a justification
for inclusion in Theales. From the study of Pakaraimaea,
a closer affinity was underlined between the
Dipterocarpaceae and Sarcolaenaceae families. Indeed,
the affinities between these two families within the
Malvales could be regarded as greater than with the
Tiliaceae (Maguire and Ashton in Maguire et al. 1977:
p. 359-361). The Tiliaceae relation is stronger with the
African taxa while for the American Pakaraimaea the
strong affinities are to be found with the Sarcolaenaceae
(De Zeeuw, in Maguire et al. 1977). The Monotoideae
could be a link between Tiliaceae and Dipterocarpaceae.
Previously Blume (1825), who first described the
family Dipterocarpaceae, Pierre (1889-1891 in Maury
1978), Heim (1892) and Hallier (1912 in Ashton 1982)
mentioned the close affinity of Monotes and Tiliaceae.
Heim and Hallier had concluded that Monotes did not
belong to Dipterocarpaceae (Kostermans 1985). The
tilioid structure of the pollen exine (Maury et al. 1975a,
b) in Asian and African taxa again called attention to these
possible affinities with Tiliaceae.
Kostermans (1978) excluded Monotoideae and
Pakaraimoideae from Asian Dipterocarpaceae and
formally described the family Monotaceae (suggested
by Maury 1978, Maury-Lechon 1979a, b, Maury-Lechon
and Ponge 1979) and recognised close relations between
Monotaceae and Tiliaceae. By the structure of the pollen
exine the Monotoideae strongly differ from Asian
Dipterocarpaceae in spite of the similarities of the tilioid
aspect of the exine surface.
Later Kostermans (1985) concluded that ‘it is very
difficult to make a decision of alliance of the real Asiatic
Dipterocarpaceae. They are not much allied to Guttiferae
or Ternstroemiaceae’ (=Theaceae) ‘and apparently
represent an ancient family, in which nowadays links with