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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 />

Table 1. Recent content <strong>of</strong> Dipterocarpaceae family.<br />

Families Sub families Genera<br />

Maguire et al. 1977, Maguire and Ashton 1980<br />

Dipterocarpaceae Monotoideae Monotes<br />

Marquesia<br />

Pakaraimoideae Pakaraimaea<br />

Dipterocarpoideae see table 2<br />

Maury 1978, Maury-Lechon 1979a, b*<br />

Monotaceae Monotoideae Monotes<br />

Marquesia<br />

Pakaraimoideae Pakaraimaea<br />

Dipterocarpaceae see table 2 see table 2<br />

Kostermans 1978, 1985, 1989<br />

Monotaceae Monotes<br />

Marquesia<br />

Pakaraimaea<br />

Dipterocarpaceae see table 2 see table 2<br />

Londoño et al. 1995<br />

Monotoideae Pseudomonotes<br />

Monotes<br />

Marquesia<br />

* presented 1977, no <strong>for</strong>mal status <strong>for</strong> taxonomic ranks, emphasis on<br />

greater affinities among taxa.<br />

the data now available, and the collaboration <strong>of</strong> still active<br />

workers, to define a solution acceptable to all in the<br />

laboratory, herbaria and field and the timber markets.<br />

First, however, more collections are needed <strong>of</strong> what<br />

appear to be key characters, in order to test their validity,<br />

particularly among species currently difficult to assign<br />

to supraspecific groupings.<br />

Botany<br />

Pakaraimaea are relatively small trees or sometimes<br />

even shrubs with alternate leaves (Table 3), conduplicate<br />

in aestivation, triangular stipules tomentulose outside<br />

and glabrous within, early fugaceous, glabrescent<br />

petioles, inflorescences axillary, racemi-paniculate,<br />

flowers 5-merous, petals shorter than sepals, neither<br />

connate at the base nor <strong>for</strong>ming a cup and not winged at<br />

all, all 5 sepals become ampliate and none alate, calyx<br />

persistent, anthers deeply basi-versatile, connective<br />

conspicuously projected as an apical appendage, pollen<br />

grains tricolporate, exine 4-layered, ovary 5-locular<br />

(rarely 4), each loculus 2-ovulate (rarely 4), fruit with 5<br />

ali<strong>for</strong>m short sepals, capsule at length dehiscent or<br />

splitting along dorsal line <strong>of</strong> carpel, wood, leaves and<br />

ovary devoid <strong>of</strong> resin or secretory canals, wood rays<br />

dominantly biseriate. No economic use is known<br />

(Maguire et al. 1977, Maguire and Steyermark 1981).<br />

Monotoideae are <strong>of</strong> three genera, Monotes,<br />

Pseudomonotes and Marquesia, and are trees or shrubs<br />

(Table 3). They have alternate leaves presenting an extrafloral<br />

nectary at the base <strong>of</strong> the midrib above (Verdcourt<br />

1989), small caducous stipules papyraceous,<br />

inflorescences in simple panicles, flowers 5-merous, 5<br />

sepals equally accrescent, petals longer than sepals and<br />

variously pubescent, calyx persistent, anthers basiversatile<br />

with apical connective-appendage scarcely to<br />

somewhat developed, pollen grains tricolporate, exine<br />

4-layered; ovary 1 to 3 locular (rarely 2, 4 or 5) with<br />

generally 2 ovules in each locule (rarely 4) except in<br />

Pseudomonotes (1 only), wood, ovary and commonly<br />

leaves without resin ducts, fruit sepals ali<strong>for</strong>m and neither<br />

connate at the base nor <strong>for</strong>ming a cupule, wood rays<br />

dominantly uniseriate.<br />

In Marquesia, trees are tall to medium-sized and<br />

buttressed, leaves evergreen and acuminate, nerves<br />

prominent with tertiary venation densely reticulate,<br />

indumentum <strong>of</strong> simple hairs and minute spherical glands<br />

on nerves and venation; flowers are small in terminal and<br />

axillary panicles; ovary 3-locular becoming 1-locular<br />

above parietal placentation, 6 ovules; fruit is ovoid with<br />

5 wings derived from the accrescent calyx, <strong>of</strong>ten 1seeded<br />

and apically 2, 3 or 4-dehiscent.<br />

Monotes are shrubs to medium-sized trees without<br />

buttresses, with leaves mostly rounded or retuse at apex,<br />

rarely acuminate, with more or less rounded extra-floral<br />

nectary at the base <strong>of</strong> the midrib above and sometimes<br />

additional ones in lower nerve-axils, with very varied<br />

indumentum and small spherical glands sparse or dense<br />

on both surfaces which <strong>of</strong>ten make the blades viscid,<br />

flowers in axillary small or compound panicles, ovary<br />

ovoid and hairy completely divided in 1, 2 or 3<br />

(sometimes 4: Maury 1970b, or 5: Verdcourt 1989)<br />

locules with 2 ovules in each locule, fruit subglobose<br />

presenting 5 equal minutely hairy wings derived from<br />

accrescent calyx, fruit normally 1-seeded and<br />

indehiscent (<strong>of</strong>ten 2, sometimes 3 or 4, rarely 5; in Maury<br />

1970b).<br />

Pseudomonotes trees are 25-30 m tall with a 70-80<br />

cm diameter, with poorly developed buttresses. This<br />

species <strong>for</strong>ms entire alternate leaves conduplicate in<br />

7

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