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First Phylogenetic Analysis of the Subfamily Pachydeminae (Coleoptera, Scarabaeoidea, Melolonthidae): the Palearctic Pachydeminae*

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First Phylogenetic Analysis of the Subfamily Pachydeminae (Coleoptera, Scarabaeoidea, Melolonthidae): the Palearctic Pachydeminae* J. Zool. Syst. Evol. Research 41 (2003) 2–46 Received on 24 October 2000 Ó 2003 Blackwell Verlag, Berlin ISSN 0947–5745 1Department of Systematic Zoology, Evolutionary Biology Centre, Uppsala University, Uppsala, Sweden (Present address); 2Departamento de Biodiversidad y Biologı´a Evolutiva (Entomologı´a), Museo Nacional Ciencias Naturales, Madrid, C.S.I.C., Spain First phylogenetic analysis of the subfamily Pachydeminae (Coleoptera, Scarabaeoidea, Melolonthidae): the Palearctic Pachydeminae* 1,2 2 I. SANMARTI´ N and F. MARTI´ N-PIERA Abstract This paper presents the first phylogeneticanalysis of Pachydeminae Reitter, 1902; one of the least known subfamilies of Melolonthidae, ‘leaf- chafers’ (Scarabaeoidea, Coleoptera). Some species of Pachydeminae have recently become agricultural pests in southern Spain. We analysed the phylogenetic relationships among 49 species belonging to 16 genera in the Palearctic region, based on a set of 63 morphological characters from the adult external morphology, wing anatomy, mouthparts and male and female genitalia. The last three sets of characters are described here for the first time. The phylogeny shows that the Palearctic Pachydeminae are monophyletic within the subfamily. Mouthparts and male and female genitalia provide the best synapomorphies for intergeneric relationships. In contrast, most of the external morphological characters used in the taxonomy of Pachydeminae are highly homoplastic. The phylogeny shows a basal split between the genera Hemictenius Reitter, 1897; Pachydema Castelnau, 1832, and the monospecific Peritryssus Reitter, 1918; and a second clade including the rest of genera. The remarkable Peritryssus is confirmed as a Pachydeminae, being the sister group to the monophyletic Hemictenius. Except for the position of P. rubripennis (Lucas, 1848) and P. zhora Normand, 1951, the phylogeny supports the monophyly of Pachydema but rejects the traditional division into species groups and the monophyly of the endemic Canarian species. In contrast, Tanyproctus Faldermann, 1835, must be rejected as polyphyletic. Otoclinius Brenske, 1896, is also probably polyphyletic(two new speciessynonymies), whereas Leptochristina Baraud and Branco, 1991, is either mono- or paraphyletic. The two Mediterranean genera Ceramida Baraud, 1897, and Elaphocera Gene´ , 1836, form a monophyleticgroup, this cladebeing the best supported by the data set. Ceramida is clearly monophyletic, whereas Elaphocera is probably monophyleticexceptfor E. barbara Rambur, 1843, which shares with Ceramida the character state for numerous mouthpart and genitalic characters. The phylogeny questions the genericstatus of the small and monospecificgenera of Pachydeminae. The monotypic Alaia Petrovitz, 1980, and Brenskiella Berg, 1898, are merged with Europtron Marseul, 1867, into one clade, whereas Atanyproctus Petrovitz, 1954, is grouped with some species of Tanyproctus, and the monotypic Pachydemocera Reitter, 1902, is proposed as a junior synonym of Elaphocera. Key words: Pachydeminae – Melolonthidae – Coleoptera – leaf-chafers – Palearctic – phylogeny – mouthparts – wing anatomy – internal genitalia Introduction disjunct. In the Holarctic region, they are present in western The Scarabaeoidea is one of the largest and best-studied North America and the southern Palearctic, with an east–west superfamilies of Coleoptera. It has traditionally been classified Mediterranean disjunction well known in other groups of into two ecological groups: the Laparosticti (dung beetles) and animals (Oosterbroek and Arntzen 1992). In the Neotropics, the Pleurosticti (leaf-feeding beetles or leaf chafers). The the Pachydeminae are restricted to the temperate region phylogeny of the Laparosticti is relatively well known (Bolivia, Peru, Uruguay but mainly Argentina and Chile). (Holloway 1960; Howden 1982; Scholtz 1986; Scholtz et al. They are also more abundant (both in number of species and 1987, 1988; Zunino 1984a), whereas the Pleurosticti, especially genera) in the south-east part of the Afrotropical region the most derived families, are far from resolved. Melolonthi- (Lacroix 2000). These disjunct distributions, together with the dae is an especially poorly defined family, comprising numer- reduced geographic ranges of most species (females are usually ous subfamilies of uncertain status (Scholtz 1990; Browne and flightless), make the subfamily Pachydeminae especially inter- Scholtz 1998). However, the family is economically important esting for biogeographicanalysis. because the larvae (‘white grubs’) are pests of crops and The Palearctic Pachydeminae comprise about 20 genera and underground stems of plants (Ritcher 1966). 280 species, although there is no definitive inventory (Baraud Pachydeminae Reitter, 1902, is one of the least known 1992; Lacroix 1993; Piattella and Sabatinelli 1996; Lacroix subfamilies of Melolonthidae. Its species were not considered 2000). The Palearctic region is the best-represented in terms of important for agriculture (Balachowsky 1962) until 1995, when number of species, whereas the Afrotropical region leads in Ceramida Baraud, 1987 was reported as a new pest of olive number of genera. Five out of the eight genera of Pachydemi- trees in southern Spain (Alvarado et al. 1996; Serrano et al. nae that include more than 10 species are Palearctic (Ceramida, 1996; Sanmartı´ n and Martı´ n-Piera, in prep.). The Pachydemi- Elaphocera Gene´ , 1836; Hemictenius Reitter, 1897; Pachydema nae comprise about 115 genera and 530 species worldwide, Castelnau, 1832;, Tanyproctus Faldermann, 1835). The Pale- although there is not a definitive catalogue (Lacroix 2000). arctic Pachydeminae are distributed across southern Eurasia They are distributed in all major zoogeographical regions from the Canary Islands to China, including southern Europe except India and Australia but their distribution is very (except France and Italy), North Africa, Near East, Asia Minor, Iran, Afghanistan, Caucasus, and Central Asia (see Appendix). Most species are distributed in the south-west *The second author of this paper, my thesis advisor and my good mentor, Palearctic (west of the Ural Mountains) with the eastern Fermı´ n Martı´ n-Piera, died shortly after this paper was accepted. This paper Palearctic represented by only 10 species in China and one in is dedicated to his memory (I. Sanmartı´n). Burma (Lacroix 2000). U.S. Copyright Clearance Center Code Statement: 0947–5745/03/4101–0002$15.00/0 www.blackwell.de/synergy Phylogeny of the Palearctic Pachydeminae 3 The taxonomy of the Palearctic Pachydeminae is partic- structures, wing anatomy, and male and female genitalia. We ularly difficult. The paucity of specimens in entomological compared the value as putative synapomorphies of these new collections has hampered the establishment of character characters with the male external morphologic characters variability, and has led to the description of new taxa on currently employed in the taxonomy of Pachydeminae. A specimen or interpopulation differences (Sanmartı´ n and second objective was to test the monophyly of the Palearctic Martı´ n-Piera 1999a). In addition, identification rests exclu- genera which might help to advance the reconstruction of a sively on characters of the male external morphology because phylogeneticsystem for the entire subfamily. females are difficult to collect and have only been described for a few species of the largest genera. However, most species of Pachydeminae present an accentuated sexual dimorphism. Materials and methods Females are characterized by a massive abdomen, reduced legs Prior to the cladistic analysis, we carried out a comparative study on and antennae, and variable wing reduction. These traits are the adult morphology of Pachydeminae, including external morphol- probably adaptations to fossorial life because, after emerging ogy, mouthparts, wing anatomy, and male and female genitalia (Sanmartı´ n 1998; Sanmartı´ n and Martı´ n-Piera 1999a). We examined in the autumn, females dig into the ground for oviposition in 125 species (more than 2000 specimens) belonging to 14 Palearctic the spring (Sanmartı´ n and Martı´ n-Piera 1997; in prep.). Most genera: Alaia, Atanyproctus, Brenskiella, Ceramida, Elaphocera, of the characters used in the identification of genera (related to Europtron, Hemictenius, Leptochristina, Otoclinius, Pachydema, Pachy- the male external morphology) are not good candidates for democera, Peritryssus, Tanyproctoides and Tanyproctus (see Appendix synapomorphies because either they vary within the genus for a complete list of the species studied and their distribution). This (polymorphic) or are also present in other genera (symplesi- represents approximately 45% of the total number of species in the Palearctic region and an average of 78.5% species studied per genus, omorphies). This is especially so in the larger and more ranging from 100% (e.g. Ceramida and Elaphocera) to 21% in geographically widespread genera Tanyproctus and Pachydema. Tanyproctus (Appendix). One consequence of this lack of reliable characters is the For the study of anatomical structures, specimens were relaxed in tendency in the taxonomy of Pachydeminae to create new boiling distilled water. Mouthparts were studied under immersion in genera that are either monotypic or contain a few species. 70% ethanol and fixed to a label pinned to the specimen. The study of About 62% of the genera of Pachydeminae contain only one wing anatomy followed Browne and Scholtz (1994): the left wing was species (Lacroix 2000). Most of the monotypic
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