unaware of any representatives of these in Monte-
verde. Fruit-eating fish and large herbivorous lizards
are absent from the area, probably due to topographic
and climatic factors.
The distribution of dispersal syndromes of Monteverde
plants. Dispersal syndromes of Monteverde plants
are summarized in Figure 8.3 (Wheelwright et al.
1984, Bronstein and Hoffman 1987, Sargent 1994,
K. G. Murray, unpubl. data, Appendix 1). In many
cases, inferences are based on plant characteris-
tics, rather than on direct observation of dispersal
events. Dispersal agents for many species need
investigation.
Comparison of the Monteverde flora with those at
other neotropical sites is difficult because most stud-
ies provide data for only a subset of plants, usually
trees (Howe and Smallwood 1982). Dispersal by ani-
mals is nearly as common among trees at Monteverde
as at other neotropical wet forest sites. More than 81%
of the tree species in Figure 8.3 appear to be adapted
for consumption and dispersal by vertebrates, com-
pared to 89% at Alto Yunda, Colombia (Hilty 1980),
92% at La Selva, Costa Rica (Frankie et al. 1974), and
92% at Rio Palenque, Ecuador (Gentry 1982). As at
the other sites, adaptation for dispersal by birds is
Figure 8.3. Frequencies (% of species) of dispersal
systems of Monteverde flowering plants, by growth form.
The graph includes native plant species for which
Appendix 1 identifies or predicts the dispersal system.
Category abbreviations: BD = bird; BDBT = bird/bat; BT
= bat; TM = terrestrial mammals; AM = aboreal
mammals; AE = animal exteriors; ANT = ant; EX =
ballistic (explosive); WD = wind; and GR = gravity.
especially common, accounting for 80% of the animal-
dispersed tree species at Monteverde and distributed
widely among plant taxa. Adaptation for dispersal by
bats is far less common among Monteverde plants,
and is concentrated in a small number of taxa and
growth forms, primarily shrubs (Piperaceae and
Solanaceae) and vines (Araceae and Cyclanthaceae).
We include few species in the "bird/bat" category
because most fleshy-fruited plants fit into one or the
other of the syndromes; studies focused on both birds
and bats are needed (Fig. 8.3). Plants with obvious
adaptations for dispersal by ants are uncommon in
Monteverde, perhaps because the cool wet condi-
tions are unfavorable for many of the seed-harvesting
ant taxa responsible for dispersal at other sites (see
Chap. 4, Insects and Spiders). This category does not
include species with small seeds that are secondarily
dispersed by ants.
Animal-dispersed species do not dominate all
growth forms. A majority of epiphytes (66%) and
herbs (73%) are adapted for dispersal by abiotic
means, especially wind (Appendix 1). The domina-
tion of the epiphyte growth form by a single family,
the Orchidaceae (>350 epiphytic species, all wind dis-
persed), however, obscures the importance of bird
dispersal among epiphytes. Lianas and shrubs are
intermediate between trees and herbs, with approxi-
mately 35-45% of species abiotically dispersed. That
35% of terrestrial herbs bear adaptations for wind
dispersal is surprising, since wind dispersal is thought
to predominate among plants whose fruits occur in
the upper strata of the forest (Howe and Smallwood
1982). Most wind-dispersed terrestrial herbs at Monte-
verde belong to the Asteraceae, and many are weedy
plants more characteristic of roadsides and river
courses than of closed-canopy forest.Fruit-handling techniques of Monteverde frugivores.
Guilds of fruit-eating animals obtain and eat fruits in
particular ways. Birds and bats may eat fruits while
perched in the plant where they are produced, or they
may carry the fruit some distance away before eating
it. The latter behavior is especially common among
bats. In Santa Rosa National Park, for example, Carol-
lia perspicillata typically consume fruits at night
roosts 30-40 m from their source trees (Heithaus and
Fleming 1978). Birds also use different feeding tech-
niques. Flycatchers, cotingas, thrushes, and Resplen-
dent Quetzals typically take fruits on the wing via
"sallying," whereas most tanagers and finches pluck
fruits while perched. "Mashers" typically crush fruits
in the bill after plucking them and then discard the
fruit skin and most or all of the seeds before swal-
lowing the remaining pulp; "gulpers" swallow fruits
whole, ingesting the skin and all of the seeds (Levey258 Plant-Animal Interactions