J. Terborgh et al., PREDATION BY VERTEBRATES AND INVERTEBRATES ON THE SEEDS OF 5 CANOPY TREE SPECIES OF AN AMAZONIAN FOREST, Vegetatio, 108, 1993, pp. 375-386
We studied the pre-germination loss of seeds to invertebrate and verte
brate seed predators of 5 species of Amazonian trees (Astrocaryum macr
ocalyx - Palmae; Bertholletia excelsa - Lecithydaceae; Calatola venezu
elana - Icacinaceae; Dipteryx micrantha - Leguminosae (Papilionoidae);
Hymenaea courbaril -Leguminosae (Caesalpinoidae)). These five species
were selected from a large tree flora on several criteria. All posses
s large (3-10 cm) well-protected seeds that might plausibly be attract
ive to mammalian seed predators. The reproductive biology of three of
the species, or close congeners, had been studied elsewhere in the Neo
tropics (Astrocaryum, Dipteryx, Hymenaea); one is important to the eco
nomy of southeastern Peru (Bertholletia); and one, despite large and a
pparently edible seeds, appeared to suffer no pre-germination loss to
predators (Calatola). We conducted the research in mature forests in t
he Manu National Park of southeastern Peru where mammal densities are
unperturbed by human activities. Densities of adult trees of the five
species in our area range from very high (> 30 per ha: Astrocaryum) to
very low (much less than 1 per ha: Hymenaea). Loss of seeds to all ca
uses, and to mammalian seed predators in particular, was determined fo
r seeds placed in 2-square meter mammal exclosures and in open control
s located at 10 m (near) and 50 m (far) from a large mature individual
of the target species (with minor variations in the design for Astroc
aryum and Calatola). The exclosures were of two types: impermeable - d
esigned to exclude all mammals, but not invertebrate seed predators, a
nd semipermeable - designed to admit small (< 500 g), but not large ma
mmals. Experimental and control plots were stocked with apparently via
ble seeds during the dry-wet transition period (October-November) and
scored one year later. A significant distance effect (higher predation
near vs far from a large conspecific adult) was found in only one of
the species (Astrocaryum), the only one to be attacked with high frequ
ency by invertebrate seed predators. The absence of any detectable dis
tance effect attributable to mammals suggests that mammals, over the c
ourse of a year, thoroughly search the forest floor for seeds. Inverte
brates may thus be responsible for most pre-germination distance (dens
ity) effects. With respect to the treatments, we found three qualitati
vely distinct results: seeds of three species (Astrocaryum, Berthollet
ia, Dipteryx) were significantly protected by the impermeable, but not
semipermeable exclosures, implicating small mammals in seed loss; the
seeds of one species (Hymenaea) were significantly protected by exclo
sures of both types, implicating large mammals; and the seeds of one s
pecies (Calatola) exhibited 100% survival, whether or not protected by
exclosures. The importance of large mammals as seed predators is gene
rally underestimated in these experiments because semipermeable exclos
ures may serve as foraging reserves for small mammals. Finally, we not
ed no relationship between the intensity of mammalian seed predation (
as suggested by the survival of unprotected seeds) and the abundance o
f adults of the five species in the environment. The diversity of resu
lts obtained for the five species reveals that large-seeded tropical t
rees may display a wide range of demographic patterns, and points to t
he likely importance of post-germination bottlenecks in the population
biology of many species, even those that may experience severe pre-ge
rmination seed loss.