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Amphimallon Berthold, 1827, in the Mediterranean Basin (Coleoptera, Scarabaeidae, Melolonthinae, Rhizotrogini) ⁎ Olivier Montreuil

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Amphimallon Berthold, 1827, in the Mediterranean Basin (Coleoptera, Scarabaeidae, Melolonthinae, Rhizotrogini) ⁎ Olivier Montreuil Available online at www.sciencedirect.com Palaeogeography, Palaeoclimatology, Palaeoecology 259 (2008) 436–452 www.elsevier.com/locate/palaeo Biogeographic hypothesis explaining the diversity of the genus Amphimallon Berthold, 1827, in the Mediterranean Basin (Coleoptera, Scarabaeidae, Melolonthinae, Rhizotrogini) ⁎ Olivier Montreuil Département Systématique et Evolution, Muséum national d'Histoire naturelle, CP50, Entomologie, 45 rue Buffon, 75005 Paris, France Received 23 October 2007; accepted 23 October 2007 Abstract In this paper, the analysis of the current distribution of species, or species-groups, of the Western–Palaearctic genus Amphi- mallon Berthold, 1827, based on a phylogenetic hypothesis recently provided and relevant paleogeographical events that occurred in the Mediterranean Basin from Late Paleogene (35 Myr BP) to the present, supports a biogeographical scenario to explain species diversity within this genus. The center of origin of this genus is hypothesized to be in the Iberian Peninsula. Microplate fragments drifting from the Iberian Peninsula, successive landmass suture events between the Eastern and Western Mediterranean Basins, separation of Tethys and Paratethys Oceans, re-establishment of marine connections, uplift of the Alps, and drying out of the Mediterranean are hypothesized as the principal events allowing dispersal of populations throughout Mediterranean Basin and biogeographical vicariances within the genus Amphimallon. © 2007 Elsevier B.V. All rights reserved. Keywords: Biogeography; Coleoptera; Melolonthinae; Rhizotrogini; Mediterranean Basin; Neogene 1. Introduction lands and are crop pests (Hurpin, 1962; Britton, 1978). The most severely affected plants occur in the tropical With some 10,000 described species, and many more regions, but severe damage to agriculture also has been still undescribed species, the Melolonthinae are an reported in many regions. In Western Europe, damage is example of an extremely species rich phytophagous mainly caused by species of the genera Melolontha beetle lineage. They constitute an ecologically and Fabricius, 1775 (Régnier, 1950; Hurpin, 1962) and taxonomically diverse group within the Scarabaeoidea Amphimallon Berthold, 1827 (Régnier, 1939, 1940, (Scarabs, Chafers, June Beetles etc…). The Melolonthi- 1950; Gambrell, 1946; Hurpin, 1962). nae are characterized, among others, by white grub Amphimallon belongs to a large tribe, the Rhizo- larvae that feed on the roots of grasses, palms and trogini, which groups together about 1400 species from various groups of trees. Many species occur in grass- Palaearctic, Nearctic, Neotropical and Oriental regions (Sabatinelli and Pontuale, 1998). Based on a revision in ⁎ Fax: +33 1 4079 3699. progress, this genus is composed of approximately 60 E-mail address: [email protected]. species inhabiting the Western–Palaearctic region. It is 0031-0182/$ - see front matter © 2007 Elsevier B.V. All rights reserved. doi:10.1016/j.palaeo.2007.10.030 O. Montreuil / Palaeogeography, Palaeoclimatology, Palaeoecology 259 (2008) 436–452 437 Table 1 geological events which occurred in the Mediterranean Geological timetable Basin. I follow the summaries of Mediterranean Basin 0.01 Myr BP Holocene Neogene palaegeography given by Rögl and Steininger (1983) 1.8 Myr BP Pleistocene and Robertson and Grasso (1995). Additionally, the 5.5 Myr BP Pliocene works of Palmer and Cambefort (2000) provided an 23.5 Myr BP L Miocene M important paleogeographical context from Late Paleo- E gene (35 MYBP) to the present (see Table 1 for 34 Myr BP Oligocene Paleogene geological timetable). 2.2. Phylogeny of the genus Amphimallon distributed across Europe (except in the northern regions), in the Middle East, from Central Asia to A phylogenetic hypothesis based on morphology Siberia, and in Northern Africa (Morocco, Northern recently has been proposed for the genus Amphimallon Algeria). Despite recent phylogenetic and taxonomic (Montreuil, 2000, Fig. 1). Nine Amphimallon species- work focusing on the Rhizotrogini (Medvedev, 1951; groups have been recognized within this genus and placed Baraud, 1977, 1985, 1992; Coca Abia, 1995; Montreuil, in three principal supergroups. The first supergroup, the 2000), our understanding of the diversity and evolution A. solstitiale lineage, consists of the A. pini-group of this group is very limited. In this study, a (seven species), the A. vernale-group (five species), the biogeographic scenario is developed based upon a A. solstitiale-group (seven species), the A. arianae- phylogenetic analysis. In the context of this analysis of group (two species), and the A. peropacum-group (one Amphimallon, there are certain assumptions regarding species). The second supergroup, the A. fuscum lineage, is the geological evolution of the Mediterranean Basin, comprised of the A. fuscum-group (eleven species) and which are related to the origin, the evolution and the the A. naceyroi-group (seven species). The third super- mode of diversification of this taxon. group, the A. ruficorne lineage, contains the A. majale group (five species) and the A. lusitanicum-group (six 2. Materials and methods species). A phylogenetic hypothesis, performed on these groups, shows sister-group relationships between the 2.1. Methods A. ruficorne lineage and A. fuscum+ A. solstitiale lineages. Seven species, namely A. amphibolum Peyer- In order to explain the current distribution of the imhoff, 1949 and six species forming the A. scutellaris- species of the genus Amphimallon in the Mediterranean group, previously placed in Amphimallon have been Basin, I implemented a phylogenetic hypothesis, as excluded from this genus and are considered species in- recommended in Humphries and Parenti (1999), and certae sedis. The above lineages and associated hypoth- followed the methodology provided by Soulier-Perkins esis of phylogeny will be used in this paper to support a (1997, 2000). The phylogenetic pattern of taxon hypothesis for historical biogeography of the genus distribution areas is inferred from the phylogenetic Amphimallon. hypothesis. This phylogenetic pattern is compared to the palaeogeographic pattern of area evolution (continental 2.3. General distribution of the genus Amphimallon breakup, seal level variation, orogeny, etc,…). When there is an agreement between both patterns, the A general distribution map of Amphimallon has been explanation of the phylogenetic pattern is thought to drawn based on a compilation of all species distributions result from the palaeogeographic evolution. (Fig. 2). Amphimallon is widely distributed around the Species distributions were treated as an attribute, and Mediterranean Basin, but species have not been were supplied for each taxon for the phylogeny, discovered in the southeastern portion of the region, previously provided for the studied group by Montreuil including Tunisia, Libya, and Egypt. Only one species, (2000), and optimized (Matile, 1990; Soulier-Perkins, A. solstitiale (Linné), has been reported from Lebanon 1997; Montreuil, 1998; Soulier-Perkins, 2000) with and Israel (Chikatunov and Pavlicek, 1997). In addi- MacCLADE (Maddison and Maddison, 1993). The tion, the presence of Amphimallon sensu stricto among most parsimonious hypothesis was retained. The the islands of the Western Mediterranean Basin is circum-Mediterranean distribution of the genus Amphi- questionable. Indeed, Amphimallon seems to be absent mallon then was analyzed, together with the results of from Balearic Islands; the sole species cited from this the performed optimization and consideration of the region, as Amphimallon, in fact belongs to the genus 438 O. Montreuil / Palaeogeography, Palaeoclimatology, Palaeoecology 259 (2008) 436–452 Fig. 1. Phylogenetic hypothesis of the genus Amphimallon based on Montreuil (2000). Rhizotrogus Berthold, 1827, as established by Montreuil recently collected Corsican specimens of A. majale (1997). Concerning the Corsican–Sardinian massif, we can be added (Michard and Michard leg.), and a sole are unaware of any citation of Amphimallon from species, A. atrum, has been cited by Baraud (1992) from Sardinia and none of the species that have been cited Piana, apparently from a single male, preserved in the from Corsica in classical works dealing with this fauna collection of Baraud (MNHN). Consequently, there are are attributable to this genus (Sainte-Claire Deville, too few specimens known from this region to confirm 1914, 1920, 1921, 1926; Schaefer, 1964; Paulian, 1977). that these species truly belong to the Corsican fauna. Three specimens preserved in old collections and Another hypothesis is that the presence of these belonging to three other species have been studied specimens in Corsica could simply be the result of from Corsica, without precise locality data. Four casual introductions from the continent. Nevertheless, O. Montreuil / Palaeogeography, Palaeoclimatology, Palaeoecology 259 (2008) 436–452 439 Fig. 2. Global distribution of the genus Amphimallon according to Medvedev (1951) and Baraud (1985, 1992). examined specimens that belong to Amphimallon were defined (Table 2, Fig. 3). These geographic areas species-groups are well-represented in Iberian Penin- approximate, for the most part, those designated by sula, France, and Italy, and therefore could have been Oosterbroek and Arntzen (1992) and de Jong (1998). present in the Corsican-Sardinian Massif (see below). None of the examined species in the genus Amphi- Collection of new material will be necessary to resolve mallon is distributed in an
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