Characteristics
Form elongate oval. Labrum produced weakly beyond apex of clypeus (except
in Anomalacra [Anomalini]). Antenna with 9 or 10 segments, antennal
club with 3 segments. Scutellum exposed. Anterior coxae transverse. Mesotibial
apex with 2 spurs; spurs mesad, adjacent (not separated by basal metatarsal
segment). Tarsal claws on all legs independently movable, claws unequal in
length or size and frequently weakly split at apex, 1 claw of each pair greatly
reduced (1 claw lacking on all legs in Leptohoplia [Anomalini]).
Onychium laterally flattened. Pygidium exposed beyond apices of elytra. References:
Ohaus 1934, Machatschke 1957, Machatschke 1965.
Classification Status
MacLeay (1819) proposed the family Rutelidae for taxa previously included
in the genus Scarabaeus Linneaus. Since Blanchard (1851) the group has been
referred to as a subfamily of Scarabaeidae. Some workers regard the group
as a family or as a subfamily of Melolonthidae. We follow Machatschke (1972)
and Lawrence and Newton (1995) and regard the Rutelinae as a subfamily of
the Scarabaeidae. Tribal classification of the subfamily is stable, but some
tribes may be paraphyletic (i.e., Rutelini [see Jameson 1998]). The ruteline
tribe Anomalini is occasionally regarded as a subfamily of Scarabaeidae (i.e.,
Potts 1974, 1977a, 1977b) or a subfamily of Melolonthidae (i.e., Sabatinelli
1991). In addition, Hoplia and its congeners have occasionally been
included as a tribe (Hopliini) and included the Rutelinae (i.e., Baraud 1985).
Subtribal classification of the subfamily is, in many cases, based on diverse
assemblages of taxa, and many subtribes are not monophyletic. Phylogenies
for the Scarabaeoidea hypothesize three differing views for the relationship
of the Rutelinae: 1) that the subfamily Dynastinae is ancestral to the Rutelinae,
2) that the Dynastinae and Rutelinae are sister groups, or 3) that the subfamilies
are possible sister groups and that their relationships remain unresolved
(Endrödi 1966; Howden 1982; Iablokoff-Khnzorian 1977; Meinecke 1975;
Scholtz and Chown 1995).
Distribution
The subfamily Rutelinae is composed of approximately 200 genera and 4,100
species that are distributed worldwide (Machatschke 1972), although many taxa
remain to be described. The subfamily is divided into six tribes, five of
which occur in the New World. The tribe Spodochlamyini is found only in Central
and South America; the tribe Anoplognathini occurs in the Australia and western
Central and South America; the tribe Geniatini is distributed in Central and
South America; the tribe Rutelini is widely distributed but is most speciose
in the Neotropics; the tribe Anomalini is widely distributed and is most speciose
in the Old World. The tribe Adoretini is exclusively distributed in the Old
World. In the New World, the subfamily includes about 95 genera. Keys to genera
and species: Casey 1915; Cooper 1983; Jameson 1990, 1997. Keys to larvae:
Ritcher 1966; Jameson et al. 1994, Jameson 1997. United States catalog: Hardy
1991. Mexico, Central and South America catalog: Blackwelder 1944. World catalog:
Machatschke 1972.
New World Tribes
Anomalini
Anoplognathini
Geniatini
Rutelini
Spodochlamyini
TRIBE
ANOMALINI
Characteristics: Labrum horizontally produced with respect
to the clypeus. Antennae with 9 segments. Protibiae bidentate (rarely unidentate
or tridentate), inner protibial spur subapical (lacking in Leptohoplia);
foretarsomeres not enlarged or densely setose ventrally. Elytra with membranous
border at lateral margin. Terminal spiracle not positioned in pleural suture.
The tribe Anomalini includes one of the largest genera in the Animal Kingdom:
the genus Anomala, which includes approximately 1,000 species worldwide. Adult
anomalines feed primarily on flowers and floral parts. Larvae feed primarily
on plant roots. One introduced member of the tribe, Popillia japonica
Newman, causes economic damage to agricultural crops and ornamental plants.
Despite the agricultural importance of some members, the tribe is poorly known
taxonomically and in desperate need of revisionary studies. The tribe includes
11 genera in the New World. Keys: Cooper 1983; Potts 1974, 1977a, 1977b. Key
to larvae: Ritcher 1966.
TRIBE
ANOPLOGNATHINI
Characteristics: Labrum vertically produced with respect
to the clypeus, with apicomedial projection. Mentum lacking apical tooth or
projection. Antennae with 9 or 10 segments. Terminal spiracle positioned in
pleural suture.
The tribe Anoplognathini is distributed in the southern hemisphere. Members
are found in Australia (some members are commonly called Christmas beetles)
and from central Mexico to southern South America. In the New World, the tribe
includes five genera (Aulacopalpus, Brachysternus, Hylamorpha,
Phalangogonia, and Platycoelia). Keys: Smith 2003.
TRIBE
GENIATINI
Characteristics: Labrum vertically produced with respect
to the clypeus, with apicomedial projection. Antennae with 10 segments. Protibiae
tridentate, inner protibial spur apical; foretarsomeres usually enlarged in
males and/or females and densely setose ventrally. Elytral margin chitinous
or membranous. Terminal spiracle positioned in pleural suture.
The tribe Geniatini is exclusively a Neotropical group of beetles. Little
is known of the natural history of the group. Members inhabit deciduous forests
and cloud forests, and some adults are attracted to lights at night. In the
new World, the tribe includes about 12 poorly studied genera. Keys: Machatschke
1965; Villatoro and Jameson 2001; Jameson
and Hawkins 2005.
TRIBE RUTELINI
Characteristics: Labrum horizontally produced with respect
to the clypeus. Antennae with 10 segments (8 or 9 in Parachrysina). Protibiae
tridentate, inner protibial spur apical; foretarsomeres not enlarged or densely
setose ventrally. Elytral margin entirely chitinous. Terminal spiracle positioned
in pleural suture.
The tribe Rutelini is distributed worldwide but is most speciose in the Neotropics.
A wide array of morphological forms is exhibited by members of the tribe including
taxa with enlarged, horn-like mandibles (Fruhstorferia from Asia),
backward-projecting thoracic horns (Peperonota from Asia), enlarged
hind femora (Heterosternus and Chrysina from the New World),
and strikingly-colored, metallic silver and gold beetles (Plusiotis
from the New World). Exemplar genera in the tribe include Pelidnota,
Macraspis (both of which are most diverse in Central and South America),
and Parastasia (most diverse in the Old World with one species in
the New World). The tribe includes over 70 genera in the New World. Keys:
Cooper 1983; Jameson 1990; Jameson in prep. Keys to larvae: Ritcher 1966;
Jameson et al. 1994; Jameson 1997. Catalog
to Pelidnota and related genera. Overview to Heterosternina.
TRIBE SPODOCHLAMYINI
Characteristics: Labrum vertically produced with respect
to the clypeus, lacking apicomedial projection. Antennae with 10 segments.
Protibiae tridentate, inner protibial spur apical; foretarsomeres not enlarged
or densely setose ventrally. Elytral margin entirely chitinous. Terminal spiracle
positioned in pleural suture.
The tribe Spodochlamyini is a small, Neotropical group that includes four
genera. Members of the group are distributed primarily in cloud forest habitats
in northernern South America. Adults are attracted to lights at night. Keys:
Machatschke 1965.
Ecology
Adult rutelines are phytophagous and feed on leaves, flowers, or flower parts.
Larvae feed on roots, compost, and decaying vegetation. Some taxa, such as
Popillia japonica Newman and Anomala species (both Anomalini),
are agricultural pests. The common name of the subfamily, the shining leaf
chafers, reflects the fact that many members of the subfamily are brightly
colored, beautifully patterned, and often brilliantly metallic leaf-feeding
beetles. Others in the subfamily, such as the genus Anomala, are
small, obscure beetles. Adults may aid in pollination of plants.
Larvae
Despite the potential agricultural importance of some rutelines, few ruteline
larvae have been described. Larvae typically feed on decaying wood, compost,
or roots. Ritcher (1966) characterized the North American ruteline larvae
as follows: Mandibles with stridulatory area consisting of transverse ridges.
Maxilla with row of anteriorly directed, sharp, pointed, stridulatory teeth;
lacinia with 1-3 unci. Epipharynx with haptomerum, with or without plegmata,
without proplegmata, two nesia usually present; haptomerum with two or more
prominent heli (beak-like or mound-like) and with 15 or more prominent spine-like
setae. Dorsal surface of abdominal segments 9 and 10 never fused. Raster with
or without palidia; anal slit transverse, slightly curved. Claws each bearing
2 setae. Keys to genera: Jameson et al. 1994.
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