Figure 4.11. Life cycle for Stmtegusjugurtha: (a) egg;
(b-d) first-, second-, and third-instar larva; (e) pupa;
(f) adult.the "weedy" species are trapped out and disappear
(Janzen 1983, B. Ratcliffe, pers. obs.).
Acknowledgments I thank Bob Law, John and Doris
Campbell, Pedro Belmar, the Monteverde Conserva-
tion League, and the Tropical Science Center for logis-
tical support. Mary Liz Jameson (University of
Nebraska), Norman Penny (California Academy of
Sciences), and Alex Reifschneider (Sierra Madre,
Calif.) assisted with collecting and field observations.
Fernando Mejia (INBio) assisted with specimen data
management for the project. Mark Marcuson (Uni-
versity of Nebraska State Museum) provided artwork
(Fig. 4.10). Research was supported by grants from the
University of Nebraska Foundation and the National
Science Foundation (DEB9200760).
4.4.4. Cleridae of Central American
Cloud Forests
Jacques Rifkind
Cleridae is a medium-sized family of predaceous
beetles that attains its maximum diversity in the trop-
ics. Sometimes called "checkered beetles," clerids areoften colorful and many are mimics of insects that
show warning coloration, such as velvet ants (Mutil-
lidae). Members of this family are diverse in body
shape and size. There are about 425 described spe-
cies of Cleridae from Mexico and Central America
(Barr 1975), which is an underestimate because
Panama and Costa Rica each contain at least as many
undescribed as described species. Although clerid
diversity is high in lowland rain forests, these beetles
become abundant only at 700 m and above.
Some cloud forest clerids have a broad altitudinal
tolerance. Priocera clavipes and the nocturnal Cymato-
dera prolixa are found in the lowland rain forests of
Belize and in the cloud forest of Monteverde. Colyphus
cylindricus has a more disjunct distribution, but it
seems not to be narrowly limited by phytogeographic
or climatic factors. It is found in relatively xeric oak
and pine forests in Guatemala, in tropical deciduous
forests at middle elevations in southern Mexico, and
in Honduras in the cloud forest of La Tigra (1800 m).
Other clerids occur only in upland rain forests and
cloud forests. Colyphus irazu, for example, is found
in Monteverde and 110 km to the southeast on the
high slopes of isolated volcanoes surrounding the
central valley of Costa Rica. Some species are re-
stricted to a single isolated cloud forest. Whether these
represent relict species, local speciation events, or
insufficient collecting is unknown. Some narrowly
endemic cloud forest clerids exhibit odd morpholo-
gies and apparently lack close living relatives.
Costa Rica, the Central American country with the
most extensively studied cloud forests, contains ap-
proximately 25 genera of Cleridae. Of these, only
Callotillus, Monophylla and Lecontella—genera that
appear to be adapted to xeric thorn forests—probably
do not contain species that inhabit cloud forests. Al-
though some genera are restricted to higher elevations,
none is limited to cloud forests.
Clerid beetles occur on flowers, in dead and dying
wood, in fungi, and on vegetation (Clausen 1940). In
their larval state, they feed on the larvae of wood-
boring beetles, although some genera (e.g., Lecontella)
are predaceous on hymenopteran larvae in their nests,
and others specialize on grasshopper eggs or carrion.
As adults, clerids will eat most insects that they can
overpower, including "distasteful" species such as
coccinellid beetles.
In Monteverde, several clerid species are common
on the foliage of shrubs and small trees at forest edges
and along paths, the rough bark of larger trees, and
at blacklight and Malaise traps. Perilypus species
are very common in shaded areas. With their reddish
pronota (upper surface of the first body segment) and
bluish black striped or concolorous elytra, they
belong to a mimicry complex including similarly col-113 Insects and Spiders