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.