enlarged hind legs that are used during mating to fend
off other males (Eberhard and Marin 1996). Larvae are
unknown in most species but some South American
species are stem-borers.
Members of the subfamily Criocerinae are recog-
nizable by a constricted pronotum (the dorsal plate
of the first thoracic segment) and a shiny body sur-
face. Larvae are free-living, and some construct pro-
tective shelters from their own feces. Lema, the most
abundant genus in this subfamily, exceeds 1300 m
elevation only occasionally. In contrast, Metepoceris
has been regularly collected in Monteverde and other
cloud forests; they are large, iridescent green with two
pairs of purple patches on the elytra, and feed on
Solanaceae (Flowers and Janzen 1997).
Eumolpinae is a large group, most of which have a
robust, compact body and shiny or metallic colora-
tion. Larvae are root feeders, and very few have been
studied. In Costa Rica, the majority of Eumolpinae live
at lower elevations (Flowers 1996), but species have
been collected above 1300 m, three of which are true
high-altitude species. Alethaxius and Dryadomolpus
are small dark-colored beetles that have been col-
lected in Monteverde and on the peaks of Poas and
Barva volcanoes and from the Andes in South
America. Colaspoides bates! shows an interesting al-
titudinal separation from its almost identical relative,
Colaspoides unicolor: C. batesi, which is bright me-
tallic green, has only been collected at Monteverde
at elevations higher than 1040 m; C. unicolor, which
is dark shiny blue, is abundant in the Cordillera de
Guanacaste from 600 to 1100 m.
Chrysomelinae includes the largest and most col-
orful of Costa Rica's leaf beetles. Their larvae are
free-living. Plagiodera bistripunctata and Stilodes
retifera appear to be restricted to cloud forest ele-
vations, and most specimens of Doryphora paykulii
(the largest chrysomelid found in Costa Rica) come
from above 1300 m in the Talamanca Mountains.
Several species of Calligrapha are found higher than
1300 m. Platyphora, the most diverse chrysomeline
genus, only occasionally strays into cloud forest
elevations.
The tribe Galerucinae-Galerucini is abundant in
tropical regions, and some species of Diabrotica and
related genera are important crop pests. Larvae of many
species are root feeders but some genera (e.g., Cecro-
pia-feeding Monocesta and Ceratoma) have free-living
larvae. As in the Eumolpinae, most species occur at low
elevations with relatively few species in cloud for-
ests or above. Four species are apparently restricted
to higher elevations: Cochabamba impressipennis,
Cochabamba sp., Diabrotica porracea, and Paranapia-
caba dorsoplagiata. Diabrotica and Paranapiacaba are
very abundant at low elevations; Cochabamba appears
to be an exclusively high-altitude genus.
Because of their enlarged hind femora and jump-
ing abilities, the tribe Galerucinae-Alticini is called
"flea beetles." Adults can be located by finding pin-
holed leaves caused by their feeding. Many larvae are
root feeders, but some are free-living. Alticini is the
largest tribe of Chrysomelidae and has been the most
successful in colonizing cloud forests. We know of
only two cases where all specimens come from above
1300 m (Acrocyum dorsalis and Omophoeta albo-
fasciata), whereas other cloud forest taxa can also be
found at lower elevations. Examples include Macro-
haltica, which are frequent defoliators of Gunnera
(Gunneraceae); flightless Longitarsus, which occur in
forest litter in Atlantic rain forests to paramo; and
Marcapatica and Neothona, which occur in the Tala-
manca Mountains. The two Omophoeta represent an-
other case of altitudinal separation; O. aequinoctialis
is ubiquitous in the lowlands but is almost absent above
1300 m, whereas O. albofasciata has not been col-
lected below this altitude.
Hispinae-Hispini is relatively well-known taxo-
nomically. Larvae of some species of Hispini are
leaf miners; species of Cephaloleia live in rolled-up
emerging leaves of Heliconia (Heliconiaceae) and
other monocots. This large and abundant tribe de-
creases in species richness with increasing altitude.
Acentroptera nevermanni appears to be one of the few
exclusively high-altitude species (based on two speci-
mens). More than 60 species of Cephaloleia occur in
Costa Rica and of these, around 20 are recorded from
Monteverde. Their larval host plants are poorly known
(Staines 1996). Alurnus salvini from the Talamanca
Mountains is closely related to the very large and
colorful A. ornata, which is a palm-feeding beetle lim-
ited to lowland rain forests.
Tortoise beetles (tribe Hispinae-Cassidini) have the
edges of the pronotum and elytra expanded and flat-
tened, and many are brilliant gold or green, although
these colors fade soon after death. Larvae are free-
living, some carrying their cast skins on modified
projections that they hold over their back to hide.
Relatively few tortoise beetles are found in cloud for-
ests: Stolas costaricensis, Stolas sp., and Tapinaspis
atroannulus feed on composites (Asteraceae); Chari-
dotella emarginata probably feeds on one of the morn-
ing glories (Convolvulaceae) (Windsor et al. 1992).
Adult females of Stolas stay with their young, shelter-
ing them from predators with their greatly expanded
elytra.
For 72% of the leaf beetle species, we do not even
know the identity of their host plants, and most of the
rest are species of the north temperate zone (Jolivet116 Insects and Spiders