ored Cantharidae, Lycidae, Chrysomelidae, and Oede-
meridae. Another Monteverde resident is the enopliine
Muisca octonotata, an obovate flattened species with
a distinct pattern of dark elytral spots on a yellowish
background. This clerid is remarkably similar to two
species of Chrysomelidae: Neobrotica sp. and Mala-
corhinus sp. Although it presumably gains protection
by mimicking these leaf beetles, Muisca also tends to
curl itself under when disturbed, which may facili-
tate its fall from the plant when being pursued by a
bird or lizard. Many species of clerids, distributed
among several subfamilies, are excellent mimics of
Hymenoptera, particularly ants and mutillid wasps.
A small, shiny black Phyllobaenus common in Monte-
verde is nearly indistinguishable from the ant Campo-
notus canescens, which runs on the same waxy-leafed
vegetation as the beetle.
4.4.5. Long-horned Beetles (Cerambycidae
and Disteniidae) of Monteverde
Frank T. Hovore
Members of the families Cerambycidae and Disteniidae
are wood-borers. They are known as long-horned
beetles because most have relatively long antennae.
They have elongate bodies (3-80+ mm in length).
Diurnal species are often colorful; nocturnal species
tend to be more drab. In the Monteverde area, Schwar-
zerion holochlorum (Tempisque Borer) is bright me-
tallic green with reddish legs and long black anten-
nae and is most active on warm sunny days. However,
Psalidognathus modestus, which also flies in bright
sunlight, is shining black and is recognized by its huge
size, spiny head, and enormous, curving mandibles.
Examples of nocturnal species include the tan and
pale green Chlorida cincta and dark brown Dero-
brachus longicornis. The best known long-horned
beetle in Monteverde is the ornately patterned Har-
lequin Beetle, Acrocinus longimanus, a species that
is distinctive for the extremely long front legs of the
male.
The family Cerambycidae is divided into seven
New World subfamilies, five of which occur in Costa
Rica, plus the closely related family Disteniidae.
Parandrinae and Prioninae are medium- to large-
sized beetles, brown to black in color. The former is
distinguished by their relatively short antennae, with
beadlike segments and large curved mandibles in the
males. Cerambycinae and Lamiinae, the two most
diverse subfamilies, are variable in size and color.
Lepturinae are usually brightly colored and diurnal.
Disteniidae are nocturnal, often dark metallic col-
ored, and have long hairs laid flat along the inside
of the antennae.
There are an estimated 35,000 described species
worldwide. Costa Rica has as many as 950 species.
Within Monteverde, approximately 260 species, in-
cluding about 40 undescribed taxa, have been re-
corded since the 1960s, based on about 4,500 total
specimens from the area. As of 1996, 31 species have
been found only in the MCFP, 153 have been taken
in the greater Monteverde area but not in the MCFP,
and 31 occur in both areas.
Adult long-horned beetles may or may not feed.
Those that feed generally consume pollen, foliage,
bark, plant sap, or fungi. Many species can be found
in flowers of Croton (Euphorbiaceae). Cephalodina
crassiceps (Lamiinae-Hemilophini) feeds on leaves
of figs (Ficus, Moraceae). Captive Harlequin Beetles
(Acrocinus longimanus, Lamiinae-Acrocinini) have
been kept alive for six months on a diet of soft fruits
such as mango, cantaloupe, and avocado. Most long-
horns are strong fliers, although in several species
females are incapable of flying, and both sexes of
Phrynidius echinus (Lamiinae-Apomecynini) are
wingless.
Long-horn beetles exhibit an array of defense
mechanisms. Diurnal species often mimic noxious or
stinging insects; others are distasteful and exhibit
aposematic (warning) coloration. In Monteverde, many
flower-inhabiting species of Clytini (Cerambycinae)
and Lepturinae appear to mimic wasps and bees,
being brightly marked with bands of yellow and black.
Erana fulveola (Lamiinae-Hemilophini) and species of
Rhinotragini (Cerambycinae) resemble ichneumonid
wasps when flying; many Tillomorphini (Cerambyci-
nae) and Acestrilla minima (Lamiinae-Apomecynini)
are ant mimics. The bright red and black Oedudes
bifasciata (Lamiinae-Hemilophini) chews notches in
living stems of urticating plants (e.g., Urera elata,
Urticaceae), apparently feeding on their sap and ac-
quiring protective chemicals. Most nocturnal long-
horns are somber or cryptically colored or have their
dorsal surfaces ornately sculptured or clothed with
hairs, often closely matching substrates on which
they rest. Most long-horned beetles emit an alarm-
ing, scraping sound when handled, and some species
can inflict a painful bite.
In many Cerambycinae, copulation takes place
shortly after the adults emerge from their pupal
cells, but in Lamiinae mating usually does not occur
until after a period of feeding. Except for flower-
visiting species, mating takes place on the host plant.
Many long-horned beetles emit a strong-smelling sex
pheromone (attractant) that is detectable by human
observers. Male beetles with well-developed man-
dibles often compete violently for females or host
resources.114 Insects and Spiders