Characteristics
Size 12.0-52.0 mm from apex of clypeus to apex of pygidium. Form elongate
oval, some species with greatly enlarged hind legs in the male; labrum horizontally
produced with respect to the clypeus; clypeus and mentum lacking apicomedial
projections; mandible with apical tooth recurved; frontoclypeal suture complete;
antennae with 10 segments; protibiae tridentate, inner protibial spur apical;
foretarsomeres cylindrical in males and females; elytra without membranous
margin; terminal spiracle positioned in pleural suture.
Generic Profiles (click on the
genus name)
Elcarmeniella | Macropoides | Parisoleoides |
Heterosternus | Mesosternus | Plesiosternus |
Homoiosternus | Paraheterosternus | Promacropoides |
Macropoidelimus | Parisolea |
Distribution
The subtribe Heterosternina includes 11 genera, 18 species, and 1 subspecies.
Species in the subtribe inhabit semideciduous and cloud forests in the Neotropics.
They occur from northern Mexico to Ecuador.
Ecology
Heterosternine beetles inhabit semideciduous and cloud forests (sea level
to 3000 m). Host plant records include Persea, Apeiba, and Quercus. Adults
can be collected at lights at night from February to December. Female heterosternines
lay their eggs in rotting logs. Adult males in some species possess enlarged
hind legs. Observations of these beetles do not support the idea that legs
are used to grasp females during copulation.
Classification Status
Horn (1866) was first to create the group, the "Macropni," which
he stated was intermediate between the "Pelidnotae" and "Areodae"
in the "true Rutelides." Horn placed Macropnus crassipes
Horn in the Macropni, and he distinguished this group from the "Pelidnotae"
and "Areodae" based on morphological characters. Bates (1888) made
reference to Horn's distinction of this group, but he referred to the group
as the Heterosterninae. Bates included in the Heterosterninae three genera:
Macropoides, Heterosternus, and Parisolea. Because the family
group name Macropnina has not been used since Horn (1866), we consider the
name a nomen oblitum.
Phylogeny
Sister-Group Relationships: In the classification of the
Rutelinae, Ohaus (1918) indicated that the Areodina or Acrobolbiina (transferred
to Dynastinae: Cylcocephalini by Jameson, Ratcliffe, and Maly 2001) was closely
related to the Heterosternina. Morón (1983) stated that the Areodina
was closely related to the Heterosternina, and he hypothesized that the group
evolved recently (11,000 years before present) and rapidly. Jameson's (1998)
analysis of relationships in the tribe Rutelini (based on cladistic methods
and morphological characters) hypothesized that the Rutelarcha clade is basal
to the Heterosternina. This clade is composed of three Old World genera and
is part of the "Parastasiina grade."
Monophyly: Morón (1983) did not discuss synapomorphic
characters for the Heterosternina. Preliminary phylogenetic analyses by Team
Scarab demonstrated that the subtribe is monophyletic and is supported by
several morphological characters.
In-group Relationships: Morón (1983) conducted an
in-group analysis of the Heterosternina using distance measures to and an
in-group cladistic analysis using Phylip hypothesize relationships. Preliminary
analyses conducted by Team Scarab demonstrated much homoplasy in the Heterosternina.
Larvae
Three genera and 4 species of heterosternine larvae are described: Macropoides
crassipes (Morón 1983), M. nietii (Morón 1983),
Paraheterosternus luedeckei (Morón 2000), and Heterosternus
buprestoides (Morón 1983). Larvae live in rotten logs (including Liquidambar,
Alnus, Brosimum, Pinus, and Quercus) and have been found
in combination with larvae of Macraspis and Chrysina (Scarabaeidae:
Rutelinae: Rutelini). Larvae have a 1 to 2 year life cycle.
References
DELGADO, L., and J. BLACKALLER-BAGES. 1997. A new Mexican
species of Homoiosternus (Coleoptera: Melolonthidae; Rutelinae).
Journal of the New York Entomological Society 105:170-179.
MAES, J.-M., and B. C. RATCLIFFE. 1996. Scarabaeidae nuevos
para la fauna de Nicaragua. Revista Nicaragüense de Entomologia 34:17-18.
MORON R., M. A. 1983. A revision of the subtribe Heterosternina
(Coleoptera, Melolonthidae, Rutelinae). Folia Entomol—gica Mexicana
55:31-101.
MORON, M. A. 1987. Adiciones a los Heterosternina (Coleoptera:
Melolonthidae, Rutelinae). Fol. Ent. Mexicana 73:69-87.
MORON, M. A. and H. F. HOWDEN. 1992. A second species of
Plesiosternus Morón with notes on other Heterosternina (Coleoptera:
Scarabaeidae: Rutelinae). Coleopterists Bulletin 46:15-19.
MORON, M.-A. 1994. La diversidad generica de los Coleopteros
Melolonthidae en Mexico. Acta Zoologica Mexicana (n.s.) 61:7-19.
MORON, M.-A.,
C. DELOYA, A. RAMIREZ-CAMPOS, and S. HERNANDEZ-RODRIGUEZ. 1998. Fauna
de Coleoptera Lamellicornia de la region de Tepic, Nayarit, Mexico. Acta Zoological
Mexicana 75:73-116.
MORON, M.-A., and G. NOGUEIRA. 2000. Third stage larva and pupa of
Paraheterosternus luedeckei (Becker) (Coleoptera: Melolonthidae;
Rutelinae). Journal of the Kansas Entomological Society 73:62-67.
OHAUS, F. 1905. Beitrage zur Kenntniss der amerikanischen
Ruteliden. Stett. Entomol. Zeit. 1905:283-329.
OHAUS, F. 1912. Beitrage zur Kennitis der Ruteliden. X. Stett.
Ent. Zeit. 73:273-319.
OHAUS, F. 1934. XXVII. Beitrage zur Kenntnis der Ruteliden
(Col. Scarabaeidae). Mitteilungen der Deutschen Entomologischen Gesellschaft
1934:9-15.
SIGWALT, B. 1987. Promacropoides bertrandi: nouveau
genre et nouvelle espece de Rutelidae Heterosternina d'Equateur (Coleoptera,
Scarabaeoidea). Revue Fr. Ent. (N.S.) 9:13-15.