Tropical Tritrophic Interactions 287
the leaf biomass monitored for an appropriate
period of time (Holt 2000). If a species cascade
is demonstrated for the majority of species, it
suggests that top-down forces of natural enemies
are likely to result in community-wide effects. In
addition, this approach provides a theoretical link
to the issues about specialization discussed ear-
lier in this chapter. Tropical communities could
be modeled as parallel chains of species cas-
cades with coevolved specialized consumers and
their host plants. These chains could be con-
nected by generalist consumers and detrital webs.
Such a modeling approach would provide a frame-
work for connecting these two bodies of tritrophic
research.
CONCLUSION
Specialization and trophic cascades hypotheses
should continue to provide guidance to empiri-
cal studies in tropical communities. To test these
hypotheses, tropical ecologists must utilize solid
phylogenetic data combined with creative exper-
imental, correlational, observational, and mod-
eling approaches. A concerted effort by tropical
research programs that utilize these approaches
to study focal communities or other model sys-
tems (e.g., Schoener and Spiller 1999, Schoener
et al. 2001, Janzen and Hallwachs 2002, Dyer
and Palmer 2004) will allow for fruitful syn-
thesis and development of a useful theoretical
framework for tropical specialization and trophic
cascades. Such a synthetic approach would be
an improvement over the existing cacophony of
experiments, observations, and phylogenetic work
across the geographic and taxonomic landscape of
thetropics.Tritrophicinteractionsintropicalcom-
munitiesareusuallypartof aconvolutedwebwith
highly variable interaction strengths, yet with
the right approaches and study systems we can
determine which interactions are the strongest for
particular taxa and ecosystems.
ACKNOWLEDGMENTS
I thank W. Carson for inviting me to put these ideas
into writing. Research reported in this chapter
was made possible by funding from the National
Science Foundation (DEB-0344250 and DEB-
0346729), Department of Energy (Southcentral
Regional Center of NIGEC), Tulane University
(Lathro pFund), and Earthwatch Institute. Ideas,
text, and data that helped me write this chapter
were provided by D. Letourneau, R. Matlock,
G. Gentry, J. Stireman, D. Janzen, W. Hallwachs,
B. Braker, R. Marquis, P. Barbosa, and P. Coley.
Parts of the manuscript were improved by com-
ments from G. Gentry, A. Smilanich, T. Massad,
R. Matlock, T. Floyd, O. Schmitz, W. Carson, and
J. Cronin.
REFERENCES
Anderson,R.S.(1993)Weevilsandplants–phylogenetic
versus ecological mediation of evolution of host plant
associations in Curculioninae (Coleoptera, Curculion-
idae).Memoirs of the Entomological Society of Canada
165, 197–232.
Barbosa, P. (1988) Natural enemies and herbivore–plant
interactions: influence of plant allelochemicals and
host specificity. In P. Barbosa and D.K. Letourneau
(eds),Novel Aspects of Insect–Plant Interactions. John
Wiley and Sons, NewYork, pp. 201–229.
Barbosa, P. and Caldas, A. (2004) Patterns of
parasitoid-host associations in differentially para-
sitized macrolepidopteran assemblages on black wil-
lowSalix nigra(Marsh) and box elderAcer negundoL.
Basic and Applied Ecology5, 75–85.
Barbosa, P., Gross, P., and Kemper, J. (1991) Influence
of plant allelochemicals on the tobacco hornworm
and its parasitoid,Cotesia congregata.Ecology72,
1567–1575.
Barbosa, P., Saunders, J.A., Kemper, J., Trumbule, R.,
Olechno, J., and Martinat, P. (1986) Plant allelo-
chemicals and insect parasitoids: effects of nico-
tine onCotesia congregata(Hymenoptera: Braconidae)
and Hyposter annulipes (Cresson) (Hymenoptera:
Ichneumonidae).Journal of Chemical Ecology 12,
1319–1328.
Becerra, J.X. (2003) Synchronous coadaptation in an
ancient case of herbivory.Proceedings of the National
Academy of Sciences of the United States of America100,
12804–12807.
Becerra, J.X. and Venable, D.L. (1999) Macroevolution
of insect–plant associations: The relevance of host
biogeography to host affiliation.Proceedings of the
National Academy of Sciences of the United States of
America96, 12626–12631.
Bernays, E.A. and Cornelius, M.L. (1989) Generalist
caterpillar prey are more palatable than specialists for