the MCL for the Children's International Rainforest,
in which a major impetus for purchasing land in the
San Bosco area was the discovery of nonbreeding
quetzals from Monteverde. Using a charismatic sym-
bol such as the quetzal and maps of its migration
routes can create a successful fund-raising campaign,
but does this method go beyond quetzals to protect
the forest and its fauna? The answer depends on the
degree to which the movement patterns of other taxa
coincide with those of the quetzal. Species such as the
Black-faced Solitaire (which breed in the same habi-
tat and migrate to the same areas as the quetzal; Fig.
6.4) will likely survive and benefit. Others, such as
the Plain-capped Starthroat (which migrates up and
down the Pacific slope) will gain little benefit from
a reserve designed to protect quetzals. The target-
species approach is not therefore the complete solu-
tion for protecting Monteverde's bird community.
Conservation efforts must be balanced with human
needs and financial limitations; the quetzal-oriented
approach in Monteverde has helped to protect a siz-
able portion of forest that is probably large enough for
viable populations of a large number of species, in-
cluding some that are migratory. Because financial re-
sources available for land acquisition and protection
are limited, a better strategy to protect Monteverde's
birds may not exist.
6.9. ConclusionsResearch in Monteverde has greatly contributed to our
understanding of birds in general and neotropical
ornithology in particular. We have learned a number
of major lessons in 25 years of research:
(1) The fauna of Monteverde is a mix of spe-
cies of North American, Mesoamerican, and
South American origin including both resi-
dents and migrants.
(2) Seasonal movements are a regular feature of
the biology of many species, especially nec-
tarivores and frugivores.
(3) Species join mixed-species foraging flocks
for different reasons, including predator de-
fense, foraging efficiency, and territory de-
fense.
(4) Understanding behaviors of long-lived birds
such as manakins and jays requires consid-
eration of the long-term benefits of those
behaviors.
(5) The demographic characteristics of montane
tropical birds (small clutch size and long
adult survivorship) are similar to lowland
tropical birds.
(6) Interactions between hummingbirds and the
plants they pollinate and between frugivoresand the plants they disperse are similar in
that seasonal availability of plant resources
causes seasonal movements of the birds;
they differ in that nectarivores tend to have
more specialized diets than frugivores.
(7) Conservation efforts must account for the
seasonal movements of threatened species.By many measures, Monteverde is similar to other
montane sites in the neotropics, so these lessons are
likely to be valid for bird communities elsewhere in
the tropics.
What directions would be fruitful for future re-
search? Communitywide studies of birds are lacking,
so our understanding of how birds distribute them-
selves across the forest landscape is incomplete (Ter-
borgh et al. 1990). Without this information, we can-
not estimate bird abundance in Monteverde and
cannot assess the importance of birds in the ecosys-
tems they inhabit. We also do not know why conge-
neric species separate across elevational gradients.
Studies similar to Hairston's (1980) studies of sala-
mander distributions in North America would be
valuable for understanding the processes causing this
widespread pattern.
We have no existing monitoring program to docu-
ment changes in communities over time. Aside from
species-specific studies that show that population
sizes and ranges ofMyiobius Redstarts and Brown Jays
are changing (Shopland 1985, Williams et al. 1994),
the only communitywide monitoring data available
are a 10-16-year series of annual surveys for the pres-
ence/absence of breeding species in selected life zones
(Pounds et al. 1997). With so much emphasis on the
possible decline of North American migrants and the
conservation of tropical resident species (e.g., Myers
1988, Terborgh 1990, Martin and Finch 1995, Stotz
et al. 1996), more detailed monitoring data will be
very useful for identifying population trends. Most
research continues to take place in Zones 2 and 3 near
the community of Monteverde; the mid-elevation
Caribbean slope (Zones 5 and 6) remains among the
most poorly studied habitats in Costa Rica.
We emphasize the success of the autecological
method in understanding bird biology. The majority
of the studies at Monteverde have focused on one or
a few species of birds. By focusing attention on indi-
vidual species, biologists in Monteverde have discov-
ered subtleties about the natural history of tropical
birds that broader studies may have missed. Migra-
tion patterns in quetzals and bellbirds, causes of co-
operation in manakins, costs and benefits of flocking,
and demographic patterns in several species would
not have come to light without in-depth studies of
individual species. Future work in Monteverde should203 Birds