of the dry season and overwinter in the coffee plantations, feeding on fruits,
nectar, and insects. Shaded coffee plantations are the most widely used of all
agricultural habitats in the tropics, and the abundance of migratory birds in
shade coffee sometimes is even higher than that in primary forest (Greenberg
et al. 1997). Among the common migrants that visit coffee plantations are fly-
catchers, wood warblers, tanagers, and orioles. The migrant birds appear to
fare well in shaded coffee plantations (and better than resident bird popula-
tions), with survival rates of many migrant bird species comparable to those in
natural habitats (Wunderle and Latta 2000). This ability of migrant birds to
survive in shade coffee plantations is thought to reflect their more flexible
habitat needs compared with residents who breed in the area (Perfecto et al.
1996).
Arthropods in Coffee Ecosystems
The arthropod fauna in shaded coffee plantations can also be large, with the
abundance of leaf litter, fallen twigs, trees, and weeds in coffee plantations pro-
viding habitat for both ground-dwelling and arboreal species (Nestel et al.
1993, 1994). A total of 609 morphospecies of arthropods was found in shaded
coffee systems in Mexico (Ibarra-Nuñez 1990), 78 families of arthropods
(mostly Diptera, Coleoptera, Hymenoptera, and Collembola) were collected
from coffee plantations in Chiapas (Moron and Lopez-Mendez 1985), 322
insect species were found in a Costa Rican coffee plantation (Gonzalez
1999b), more than 30 ant species were registered in a traditional, shaded cof-
fee agroecosystem in Costa Rica (Perfecto and Vandemeer 1994; Barbera
2001), 50 species of Pimplinae wasps have been recorded in Salvadorian cof-
fee plantations (The Natural History Museum 2002), and 168 butterfly
species were found in a shaded coffee plantation in Colombia (Botero and
Baker 2001). A recent study reported 130 species of Auchenorrhyncha
(Homoptera), mostly Cicadellidae (82 species), in very simple coffee–Eryth-
rina poeppigianaor coffee–E. poeppigiana–Cordia alliodorasystems in Turri-
alba, Costa Rica (Rojas et al. 2001a, 2001b). Perhaps most surprisingly, 30 ant
species, 103 other Hymenopterans, and 126 beetle species were collected from
the canopy of a single E. poeppigianatree in a coffee plantation in Costa Rica.
In another E. poeppigianatree, located less than 200 m away, 30 ant species,
103 other Hymenopteran species, and 126 beetle species were found. Species
overlap between these two trees was only 14 percent for beetles and 18 per-
cent for ants, suggesting that shaded coffee plantations may have very high
levels of arthropod diversity; however, the species composition often is distinct
from that of natural forest (Perfecto et al. 1996).
Coffee agronomists have documented the varied arrays of natural enemies,
notably several species of Hymenoptera, Diptera, and Fungi, that control the
population of every known insect pest of coffee. Borers are notorious pests in
- Biodiversity Conservation in Neotropical Coffee Plantations 213