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alone to divide up the rutelines (as was later done by Ohaus). He felt that anoplognathines, geniatines, and anomalines were more closely related than previously thought. Ohaus (1918) was the first to divide the subfamily Rutelinae into two groups based on the form of the labrum: the Rutelinae homalochilidae (Anomalini and Rutelini) with a labrum parallel to the clypeus, and the Rutelinae orthochilidae (Adoretini, Anoplognathini, Geniatini, and Spodochlamyini) with the labrum perpendicular to the clypeus. Ohaus (1904a, 1904b, 1905, 1918) also did a major taxonomic overhaul on the Anoplognathini, dividing the tribe into the five currently recognized subtribes: Anoplognathina (Australia and New Guinea), Schizognathina (Australia), Brachysternina (Chile, Argentina), Phalangogoniina (México and Central America), and Platycoeliina (Neotropics). Early authors (Burmeister 1844; Lacordaire 1856; Sharp 1878; Ohaus 1904a) commented on the similarities between some of the Australian and Neotropical genera. However, when Ohaus (1918) classified the Australian genera into two subtribes and Neotropical genera into three separate subtribes, the similarities between the Australian and Neotropical groups were obscured and have not been studied since. Machatschke (1965) briefly discussed the evolution and biogeography of the Anoplognathini. He thought that the Neotropical Anoplognathini and Australian Anoplognathini were both monophyletic groups, but stopped short of suggesting that the two groups were sister taxa. The Australian subtribes of Anoplognathini were reviewed by Carne (1954, 1955, 1956, 1957, 1958). Carne (1958) mentioned some problems with Ohaus’ classification scheme in the context of Australian Rutelinae. Some Australian genera share several key character states with Geniatini, but he hypothesized that this is due to convergent evolution. Carne (1958) did not make any changes to Rutelinae classification above the generic level when reviewing the Australian fauna. I think it is most unfortunate that Carne did not critically examine at least the subtribes of Anoplognathini, especially since there are relatively few genera in the rest of MONOGRAPHIC REVISION OF PLATYCOELIA 5 the world. Machatschke (1965, 1972) mimicked Ohaus’ classification scheme in the most recent world catalogs of the subfamily Rutelinae, but Machatschke (1965) was critical of Ohaus’ division of Rutelinae tribes into multiple subtribes because many of these groups were based on a single character. For the same reason, Machatschke was critical of the division of Rutelinae into two groups (Ohaus’ Rutelinae homalochilidae and Rutelinae orthochilidae). He argued that this classification system (and the classification of Scarabaeidae in general) “is based on completely outdated opinions” (developed by Burmeister [1844] before evolutionary theory was elucidated), and that classification systems such as this will not endure when they “are still based on single characters.” Many recent authors have echoed Machatschke’s frustration with the antiquated scarab classification system. It seems inevitable that modern phylogenetic analysis will topple many of the ancient and untested ideas about relationships within the family Scarabaeidae! Jameson (1998) demonstrated problems with Ohaus’ tribal-level classification scheme of the Rutelinae, but she did not include any Anoplognathini in her analysis. Up to then, there had been no published attempt to verify the monophyly of Ohaus’ subtribes within the Anoplognathini or of the Anoplognathini with respect to the rest of the Rutelinae using modern phylogenetic methods. All three of the Neotropical subtribes have recently been revised (Brachysternina by Jameson and Smith [2002], Ratcliffe and Ocampo [2002], and Smith [2002a]; Phalangogoniina by Smith and Morón [2003]; and Platycoeliina by Smith [this paper]). My phylogenetic analysis (Smith 2002a) supported the monophyly of the subtribe Brachysternina, and I hypothesized that the split between the Australian and Neotropical taxa occurred during the breakup of the Gondwana supercontinent during the late Cretaceous or early Tertiary. The tribe Anoplognathini is, therefore, quite old, having arisen soon after the adaptive radiation of the phytophagous scarabs (Melolonthinae, Rutelinae, Dynastinae, Cetoniinae and other minor subfamilies) during the Cretaceous, along with the adaptive
6 BULLETIN OF THE UNIVERSITYOF NEBRASKA STATE MUSEUM radiation and diversification of the angiosperm plants on which they feed. Phylogenetic analyses have cast doubt on the validity of some of Ohaus’ ruteline subtribes. Jameson (1998) demonstrated the paraphyly of most of the subtribes within the tribe Rutelini and, as a result, synonymized most of the names. My preliminary phylogenetic research on the tribe Anoplognathini has supported the monophyly of the Neotropical subtribes of Anoplognathini (Brachysternina, Phalangogoniina, Platycoeliina), but not the Australian subtribes (Anoplognathina, Schizognathina). Preliminary molecular and biogeographic analyses also shed some doubt on the monophyly of the tribe Anoplognathini as well. Rutelini and Anoplognathini are paraphyletic based on these preliminary data. The analysis suggests that the Anoplognathini are basal lineages of Rutelinae and the Rutelini are the more apical lineages (having multiple origins within the Anoplognathini). This question, along with phylogenetic analyses of the Rutelinae + Dynastinae clades (preliminarily examined by Jameson [1998]), must be thoroughly addressed before a stable and enduring new Rutelinae classification system can be proposed to replace Ohaus’ classification system. It is obvious that modifications are needed. Subtribe Platycoeliina The subtribe Platycoeliina contains one genus, Platycoelia, and is endemic to the Neotropics. The subtribe Platycoeliina was first erected by Burmeister (1844) (as Platycoeliidae) to accommodate the genera Platycoelia and Phalangogonia Burmeister. Ohaus (1904b, 1905) later removed Phalangogonia and added Callichloris Burmeister and Leucopelaea Bates (now junior synonyms of Platycoelia) to the taxon. Ohaus (1918) was also the first to use Platycoeliina in its current sense and usage as a subtribe of the tribe Anoplognathini. The subtribe became monogeneric when Machatschke (1965) synonymized all of the generic names in the subtribe under Platycoelia. As discussed in the phylogeny section, the subtribe Platycoeliina (genus Platycoelia) is a monophyletic group. How- ever, the preservation of this subtribe may render other subtribes paraphyletic. The monophyly and validity of the subtribes within the Anoplognathini are topics of ongoing research on which I will report in the future. TAXONOMIC HISTORY OF THE GENUS PLATYCOELIA During their historic journey to the Spanish colonies in South America from 1799- 1803, Alexander Humboldt and Aimé Bonpland collected many plants and animals for scientific description. In South America, they collected material in Venezuela, Colombia, Ecuador, and Perú. Upon returning to Europe, Humboldt contracted Pierre André Latreille to describe some of the insects collected during the voyage. These descriptions were in the insect section of Voyage de Humboldt et Bonpland: Observations de Zoologie et d’Anatomie Comparée, which was published from 1805-1832. A specimen (or specimens) of Platycoelia was collected by Humboldt and Bonpland, and Latreille (1813) described it as Melolontha flavostriata Latreille. Blanchard [1851], Ohaus [1918], Machatschke [1965, 1972] and others have erroneously listed the date of this work as 1833, but it was published in 1813 (Sherborn 1899). At the time, there were very few generic names used for scarab beetles and most non-metallic rutelines were placed in Melolontha Fabricius. Soon afterward, authors such as Dejean, MacLeay, LePeletier and Serville, Kirby, and Hope began splitting up the old genera and creating many new generic names to accommodate the tremendous diversity of the group. Dejean published a series of catalogs listing the species in his personal collection (see Madge [1988] for more details). According to Arrow (1899), Latreille gave Dejean a specimen of P. flavostriata (which eventually ended up at BMNH). This specimen is now the lectotype of P. flavostriata. Dejean (1833, 1836) then listed the species under the new generic name Platycoelia in his catalogs. The dates of the Dejean catalogs have been the source of much discussion in the literature with Madge (1988)
Page 2 and 3: Bulletin of the University of Nebra
Page 4 and 5: C O N T E N T S Abstract...........
Page 6 and 7: LIST OF FIGURES Fig. 1. Platycoelia
Page 8 and 9: Bulletin of the University of Nebra
Page 10 and 11: Members of the genus Platycoelia De
Page 14 and 15: eing the most recent authority on t
Page 16 and 17: els Amongst the Great Andes of the
Page 18 and 19: tion. Zischka was a German who live
Page 20 and 21: Puncture size: large = 0.17 mm or l
Page 22 and 23: 26. Pronotal surface sparsely punct
Page 24 and 25: Fig. 11. Platycoelia alternans male
Page 26 and 27: MONOGRAPHIC REVISION OF PLATYCOELIA
Page 28 and 29: Fig. 33. Platycoelia rufosignata me
Page 30 and 31: CNCI Canadian National Collection o
Page 32 and 33: subspecies AND varieties in the sam
Page 34 and 35: PHYLOGENETIC PATTERNS AND BIOGEOGRA
Page 36 and 37: metaunguitractor plate usually with
Page 40 and 41: larvae, and Paucar-Cabrera and Smit
Page 50 and 51: Erichson (1847) did not state how m
Page 52 and 53: species]; Ohaus 1918:178 [catalog l
Page 58 and 59: Platycoelia marginata and Platycoel
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5. Platycoelia meridensis new speci
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in apical half. Protarsomere 4 with
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MONOGRAPHIC REVISION OF PLATYCOELIA
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7. Platycoelia prasina Erichson, 18
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written and typeface), e) “PLATYC
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mentum. Apex of mentum with triangu
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1972:302 [catalog listing as subspe
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green to yellow; parameres with the
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duced tooth. Apex of mentum with tr
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Mnisnech,” “Dr Ohaus Vidit 1903
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VARIATION. Male (n=19). Length 19.2
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“Platycoelia forcipalis cotype Oh
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males with internomedial stridulato
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tures moderate with dark coloration
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gins. Ventrally with dark brown met
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19. Platycoelia flavostriata (Latre
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20. Platycoelia nigrosternalis Ohau
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LOCALITY DATA. 66 specimens examine
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Calloodes nitidissimus Lea, 1919 (C
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