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Shocker Open Access Repository Evolution and Phylogeny of The Wichita State University Libraries SOAR: Shocker Open Access Repository Mary Liz Jameson Biological Sciences Evolution and phylogeny of the scarab subtribe Anisopliina (Coleoptera: Scarabaeidae: Rutelinae: Anomalini) Mary Liz Jameson University of Nebraska State Museum, [email protected] Estefania Micó Universidad de Alicante, Alicante, Spain Eduardo Galante Universidad de Alicante, Alicante, Spain __________________________________________________________________ Recommended citation Jameson, Mary Liz., Micó, Estefania and Eduardo Galante. 2007. Evolution and phylogeny of the scarab subtribe Anisopliina (Coleoptera: Scarabaeidae: Rutelinae: Anomalini). Systematic Entomology 32(3): 429-449. This paper is posted in Shocker Open Access Repository http://soar.wichita.edu/dspace/handle/10057/3382 Published in Systematic Entomology 32:3 (July 2007), pp. 429–449; doi:10.1111/j.1365-3113.2006.00380.x Copyright © 2007 Mary Liz Jameson, Estefania Micó, and Eduardo Galante. Journal compilation copyright © 2007 The Royal Entomological Society. Published by Blackwell Publishing. Used by permission. Accepted September 18, 2006; published online April 20, 2007 Evolution and phylogeny of the scarab subtribe Anisopliina (Coleoptera: Scarabaeidae: Rutelinae: Anomalini) Mary Liz Jameson1, Estefania Micó2, and Eduardo Galante2 1 University of Nebraska State Museum, Lincoln, Nebraska, U.S.A. 2 Centro Iberoamericano de la Biodiversidad (CIBIO), Universidad de Alicante, Alicante, Spain * Correspondence — Mary Liz Jameson, W436 Nebraska Hall, Division of Entomology, University of Nebraska State Museum, Lincoln, NE 68588-0514, U.S.A. Email: [email protected] Abstract The subtribe Anisopliina (Scarabaeidae: Rutelinae: Anomalini) is associated with grasses, and its spe- cies are distributed in the Palaearctic, Oriental, Ethiopian, Nearctic and Neotropical biogeographi- cal regions. Phylogenetic analysis of adult morphological characters was conducted to examine the monophyly and classification of the group, as well as to examine characters associated with grass pol- linivory and graminivory. We review the biology, phylogeny and classification of the Anisopliina and provide an overview of each genus. The analysis of ninety-one morphological characters using parsi- mony does not support the monophyly of the subtribe Anisopliina. Instead, the results provide support for a group referred to here as the anisopliine clade, a circum-Mediterranean group, forming an inter- nal clade within the well-supported tribe Anomalini. Sister group relationships are discussed, possi- bly being associated with a New World anomaline taxon. Character states associated with grass herbiv- ory, including mouthpart and leg characters, are discussed based on the phylogenetic analysis. Within the Anomalini, an evolutionary shift from generalized leaf feeding to grass associations and grass pol- len feeding is supported. Introduction isopliina and the evolution of grass–anisopliine associations. The objectives of our research were three-fold: (1) to test the The subtribe Anisopliina (Scarabaeidae: Rutelinae: Anom- monophyly of the Anisopliina as currently composed; (2) to alini) comprises approximately 100 species and nine gen- test the monophyly of Baraud’s (1991, 1992) subgeneric and era (Figure 1) that are distributed in the Palaearctic, Oriental, group classification of Anisoplia (s.l.); and (3) to determine Ethiopian, Nearctic and Neotropical biogeographical regions. whether character states associated with pollen and grass feed- Anisopliines are associated with cultivated and wild grasses, ing evolved one time or many times within the Anisopliina. feeding on grass pollen or immature grass seeds as adults and On the basis of the results of our analyses, we discuss the evo- grass roots as larvae. Members of the subtribe are character- lutionary associations of the anisopliine clade and grasses, as ized by an elongated and recurved clypeal apex (for example, well as the classification of the Anisopliina. Figure 7A, B, see later), a trait that enables adults to extract and consume the pollen-loaded grass anthers (Micó, 2001). Taxonomic history The group includes the wheat grain beetle or bread beetle, An- isoplia (Autanisoplia) austriaca (Herbst), and other species On the basis of current classifications (Machatschke, 1972; that are occasional pests of cultivated grasses, such as rye, Potts, 1974; Baraud, 1992), the subtribe Anisopliina in- corn and wheat (Hurpin, 1962). cludes nine genera and approximately 100 species distrib- On the basis of phylogenetic analyses using adult morpho- uted in the New and Old World: Anisoplia Schönherr, An- logical characters, we examined the monophyly of the An- thoplia Medvedev, Anomalacra Casey, Brancoplia Baraud, 429 430 JAMESON, MICÓ, & GALANTE IN SYSTEMATIC ENTOMOLOGY 32 (2007) Figure 1. Exemplar species of Anisopliina: A, Anisoplia agricola; B, Anomalacra clypealis; C, Anthoplia floricola; D, Brancoplia leucaspis; E, Callirhinus metallescens; F, Chaetopteroplia segetum; G, Hemichaetoplia gossypiata (Fairmaire); H, Rhinyptia indica; I, Tropiorhynchus podagricus. Callirhinus Blanchard, Chaetopteroplia Medvedev, Hemi- nomic studies of anomaline scarabs in the U.S.A, Potts (1974) chaetoplia Baraud, Rhinyptia Burmeister and Tropiorhynchus placed the genus Anomalacra in the Anisopliina. Blanchard. The circum-Mediterranean genera Anisoplia, Anthoplia, As a higher level taxon, the generic composition of the Brancoplia, Chaetopteroplia and Hemichaetoplia [referred to Anisopliina has varied over time. Burmeister (1844) first de- here as Anisoplia (s.l.)] are the most species-rich groups in the scribed the “Anisopliadae” in which he included, amongst subtribe and have been the subject of much European study. other genera, Anisoplia and Rhinyptia. Shortly thereafter, Baraud (1986) characterized this group based on the recurved Burmeister (1855) included Tropiorhynchus and Callirhinus. clypeal apex, form of the parameres and internal sac, and ex- Ohaus (1918) established the worldwide classification for the ternal characters such as placement and kind of setae. Within Anomalini, dividing the tribe into four subtribes, including the Anisoplia (s.l.), the wide variation in colour, pattern and se- Anisopliina, Anomalina, Popilliina and Isopliina. In the sub- tae has led to descriptions of new genera, subgenera, species tribe Anisopliina, Ohaus (1918) included four genera: Aniso- and species groups, and a large body of literature containing plia, Rhinyptia, Tropiorhynchus and Callirhinus. This classi- several classifications (for example, Mulsant, 1842, 1871; Er- fication was used by Machatschke in the Genera Insectorum ichson, 1847; Kraatz, 1883; Reitter, 1903; Medvedev, 1949; (Machatschke, 1957) and in the Coleopterorum Catalogus Machatschke, 1972; Baraud, 1986; Zorn, 2006) (see Table 1). (Machatschke, 1972). Most recently, on the basis of taxo- Reitter (1903) divided the genus Anisoplia into three groups EVOLUTION AND PHYLOGENY OF THE SCARAB SUBTRIBE ANISOPLIINA 431 based on the form of the setae on the elytral epipleuron (se- the “agricola group,” “zwickii group,” “lodosi group,” “deser- tae spiniform or not) and form of the setae elsewhere on the ticola group,” “tempestiva group,” “signata group,” “montic- body (setae decumbent or not decumbent, long or short, dense ola group,” and “villosa group.” No revisions have been con- or not). Medvedev (1949), in revising the scarabs of Rus- ducted subsequently on the group. sia, proposed five new subgenera of Anisoplia based partially As with many subtribes in the Rutelinae, character-based on the species groups of Reitter (1903). He elevated Group circumscriptions for the Anisopliina are lacking. Machatschke 1 of Reitter (species with spiniform setae on the elytral epi- (1957) did not include an overview of the Anisopliina. Baraud pleuron) to the subgenus Chaetopteroplia; Group II became (1986) characterized Anisoplia (s.l.) and discussed taxa that the subgenera Lasioplia Medvedev and Anthoplia (species share some of these characters, but he did not characterize the with long, dense setae above and below); and Group III be- Anisopliina. Potts (1974) characterized the group based on its came the subgenera Ammanisoplia Medvedev, Autanisoplia “thinned” clypeus and “reduced” labrum. Traditionally, au- Medvedev and Anisoplia (species with setae decumbent, often thors have relied primarily on the recurved form of the clypeal short or glabrous). Machatschke (1957) synonymized Medve- apex to circumscribe the group (Figures 6; 7, see later), but dev’s subgenera within the genus Anisoplia and, instead, rec- this singular character varies greatly. The form of the clypeal ognized three groups that corresponded directly with Reitter’s apex in the New World genus Anomalorhina Jameson, Pau- (1903) classification: the “segetum group” (Reitter’s Group car-Cabrera, & Solís is similar to that of other members of the I), the “villosa group” (Reitter’s Group II), and the “austri- Anisopliina (recurved and attenuated at apex), but the taxon aca group” (Reitter’s Group III). In 1972, Machatschke re- was considered to belong to the Anomalina (Anomalini) vised his classification (Machatschke, 1972), further subdi- (Jameson et al., 2003). Baraud (1986) discussed the affinity of viding the “segetum group” into a total of three groups: the Dicranoplia deserticola (Lucas) (Anomalini: Popilliina) with “leucaspis group,” “lanata group,” and “segetum group.” The other Anisopliina based on the form of the clypeus, but he dis-
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