Dispersal vs. vicariance in the Mediterranean: historical biogeography of the Palearctic Pachydeminae (Coleoptera, Scarabaeoidea)

Aim The geological evolution of the Mediterranean region is largely the result of the Tertiary collision of the African and Eurasian Plates, but also a mosaic of migrating island arcs, fragmenting tectonic belts, and extending back‐arc basins. Such complex paleogeography has resulted in a ‘reticulate’ biogeographical history, in which Mediterranean biotas repeatedly fragmented and merged as dispersal barriers appeared and disappeared through time. In this study, dispersal‐vicariance analysis (DIVA) is used to assess the relative role played by dispersal and vicariance in shaping distribution patterns in the beetle subfamily Pachydeminae Reitter, 1902 (Scarabaeoidea), an example of east–west Mediterranean disjunction. Location The Mediterranean region, including North Africa, the western Mediterranean, Balkans–Anatolia, Middle East, Caucasus, the Iranian Plateau, and Central Asia. Methods A phylogenetic hypothesis of the Palearctic genera of Pachydeminae in conjunction with distributional data was analysed using DIVA. This method reconstructs the ancestral distribution in a given phylogeny based on the vicariance model, while allowing dispersal and extinction to occur. Unlike other methods, DIVA does not enforce area relationships to conform to a hierarchical ‘area cladogram’, so it can be used to reconstruct ‘reticulate’ biogeographical scenarios. Results Optimal reconstructions, requiring 23 dispersal events, suggest that the ancestor of Pachydeminae was originally present in the south‐east Mediterranean region. Basal splitting within the subfamily was caused by vicariance events related to the late Tertiary collision of the African microplates Apulia and Arabia with Eurasia, and the resultant arise of successive dispersal barriers (e.g. the Red Sea, the Zagros Mountains). Subsequent diversification in Pachydeminae involved multiple speciation events within the Middle East and Iran–Afghanistan regions, which gave rise to the least speciose genera of Pachydeminae (e.g. Otoclinius Brenske, 1896). Finally, the presence of Pachydeminae in the western Mediterranean region seems to be the result of a recent dispersal event. The ancestor of the Iberian genera Ceramida Baraud, 1987 and Elaphocera Gené, 1836 probably dispersed from the Middle East to the Iberian Peninsula across North Africa and the Gibraltar Strait during the ‘Messinian salinity crisis’ at the end of the Miocene. Main conclusions Although the basal diversification of Pachydeminae around the Mediterranean appears to be related to vicariance events linked to the geological formation of the Mediterranean Basin, dispersal has also played a very important role. Nearly 38% of the speciation events in the phylogeny resulted from dispersal to a new area followed by allopatric speciation between lineages. Relationships between western and eastern Mediterranean disjuncts are usually explained by dispersal through Central Europe. The biogeographical history of the Pachydeminae corroborates other biogeographical studies that consider North Africa to be an alternative dispersal route by which Mediterranean taxa could have achieved circum‐Mediterranean distributions.     

五种鳃金龟幼虫描述（鞘翅目）

对日胸突鳃金龟Hoplosternus japonicus Harold,坦狭肋鳃金龟Holotrichia tonkinensis Moser,白云鳃金龟替代亚种Polyphylla alba vicaria Semenov,大皱鳃金龟Trematodes grandis Semenov和二色希鳃金龟Hilyotrogus bicoloreus(Heyden)5种幼虫首次进行了形态描述,绘制了特征图,并编制了幼虫分类检索表。     

First phylogenetic analysis of the subfamily Pachydeminae (Coleoptera,Scarabaeoidea, Melolonthidae): the Palearctic Pachydeminae*

This paper presents the first phylogenetic analysis 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 species synonymies), whereas Leptochristina Baraud and Branco, 1991 , is either mono‐ or paraphyletic. The two Mediterranean genera Ceramida Baraud, 1897, and Elaphocera Gené, 1836, form a monophyletic group, this clade being the best supported by the data set. Ceramida is clearly monophyletic, whereas Elaphocera is probably monophyletic except for E. barbara Rambur, 1843, which shares with Ceramida the character state for numerous mouthpart and genitalic characters. The phylogeny questions the generic status of the small and monospecific genera 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 .     

索鳃金龟属四新种记述（鞘翅目：鳃金龟科）

本文记述了索鳃金龟属ＳｏｐｈｒｏｐｓＦａｉｒｍａｉｒｅ４新种,无距索鳃金龟Ｓ．ａｃｄｌｃａｒｉｕｍ,细体索鳃金龟Ｓ．ｓｔｅｎｏｃｏｒｐｕｓ,粉臀索鳃金龟,Ｓ．ｐｒｕｉｎｏｓｉｐｙｇａ和漾濞索鳃金龟Ｓ．ｙａｎｇｂｉｅｎｓｉｓ。模式标本保存在沈阳农业大学植保系昆虫标本室。     

鳃金龟科两属属征修订及3新种记述（鞘翅目：金龟总科）

本文对鳃金龟科的索鳃金龟属Sophtops和黄鳃金龟属Metabolus的属性作了修订,并记述3个新种。     

四种鳃金龟对不同植物挥发物的触角电位反应

华北大黑鳃金龟Holotrichia oblita Falderman、暗黑鳃金龟Holotrichia parallela Motschulsky、小黄鳃金龟Metabolus flavescens Brenske和福婆鳃金龟Brahmina faldermanni Kraatz有不同的寄主范围.本研究采用溶剂提取法...     

Revision of the rhinoceros beetle genus Oryctophileurus Kolbe with description of a new species,the male of O. varicosus Prell,and notes on biogeography (Scarabaeoidea,Dynastinae, Phileurini)

The genus Oryctophileurus is reviewed and its validity is supported by a combination of the following apomorphic characters: a single cephalic horn with lateral carina, pronotal cavity with ocellate punctures and two teeth or tubercles close behind the anterior pronotal margin. The male of Oryctophileurus varicosus Prell, 1934, is described for the first time. A new species, Oryctophileurus guerrai Perger & Grossi sp. n., from subhumid Tucuman-Bolivian forest in the Southern Bolivian Andes is described. The new species is distinguished from its closest relative, O. armicollis Prell, 1911, by a narrower distance between the inner teeth of the dorsal pronotal protuberances and a reduced area of weakly developed ocellate punctures above the posterolateral pronotal margin. The occurrence of Oryctophileurus species in areas of endemism along the eastern slope of the tropical Andes suggests that these populations represent biogeographic “relicts”, and the discovery of Oryctophileurus guerrai sp. n. in the southern Bolivian Andes suggests that this area is underrated with respect to insect diversity and endemism.     

东北与华北大黑鳃金龟分类地位的研究（鞘翅目：鳃角金龟科）

采用RAPD技术和杂交方法对东北大黑鳃金龟（Holotrichia diomphalia Bates1888)和华北大黑鳃金龟（H.oblita(Faldermann1935)的分类地位进行了研究。作者应用40条引物,采用UPDGA法对华北大黑的南京和文登种群。东北大黑的沈阳种群。以及其外群矮臀鳃金龟（H.ernesti Reitter)进行了RAPD分析。所建立的系统发育树显示,东北大黑具有与华北大黑两个地理种群相同的分类地位。同时,作者对两种金龟甲进行了正反交的杂交试验。试验共得到21头正交F1代成虫,3头反交F1代成虫,F1代生活正常,采用东北大黑与F1代回交的方法研究F1代的可育性,2001年6月10日的检查结果显示,四个处理均得到大量回交杂种一龄幼虫和少量二龄幼虫,F1代的雌雄虫均具有正常的生殖力。以上结果表明;两种金龟的次异名。为无效名。     

Does the Trans‐mexican Volcanic Belt represent a natural biogeographical unit? An analysis of the distributional patterns of Coleoptera

Aim We analysed the geographical distribution of beetle species of the families Buprestidae, Cerambycidae, Dryophthoridae, Melolonthidae, Passalidae and Staphylinidae from the Trans‐mexican Volcanic Belt (TVB) through a track analysis and a parsimony analysis of endemicity (PAE), in order to test its naturalness and determine its affinities. Location The area analysed corresponds to the TVB, which is a biogeographical province of the Mexican Transition Zone. Methods The panbiogeographical analysis was based on the comparison of the individual tracks of 299 species of Buprestidae, Cerambycidae, Dryophthoridae, Melolonthidae, Passalidae and Staphylinidae (Coleoptera). The TVB was divided into 1^o × 1^o grid cells and we also included in the analysis the remaining Mexican biogeographical provinces. Parsimony analysis of endemicity with progressive character elimination (PAE‐PCE) was applied to classify areas by their shared taxa according to the most parsimonious cladograms. The nested sets of areas were represented as generalized tracks. Results Three generalized tracks were obtained: (1) grid cells 9C, 9D, 10D, 10E, Sierra Madre Oriental, Chiapas, Mexican Gulf and the Sierra Madre del Sur; (2) grid cells 3B, 3C, 4B, 4C, 5C, 6C, 7C, Sierra Madre Occidental, Sierra Madre del Sur, Balsas Basin and the Mexican Pacific Coast, and (3) grid cells 8D, 9C, 9D, 10D, 10E, Yucatán Peninsula, Chiapas, Sierra Madre Oriental and the Mexican Gulf. Main conclusions We conclude that the TVB does not represent a natural biogeographical unit because it shows different relationships with other biogeographical provinces, being clearly transitional between the Nearctic and Neotropical provinces. Some parts of the TVB are related to Neotropical provinces (Chiapas, Mexican Gulf and Mexican Pacific Coast) and others to the remaining provinces of the Mexican Transition Zone (Sierra Madre Oriental, Sierra Madre del Sur, Sierra Madre Occidental and Balsas Basin).