Beutel Rolf G, Balke Michael, Steiner Warren E
Institut für Spezielle Zoologie und Evolutionsbiologie, FSU Jena, 07743 Jena, Germany.
Zoologische Staatssammlung, Münchhausenstrasse 21, 81247 München, Germany and Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK.
Cladistics. 2006 Apr;22(2):102-131. doi: 10.1111/j.1096-0031.2006.00092.x.
A phylogenetic analysis of Adephaga is presented. It is based on 148 morphological characters of adults and larvae and focussed on a placement of the recently described Meruidae, and the genus-level phylogeny of the smaller aquatic families Gyrinidae, Haliplidae and Noteridae. We found a sister group relationship between Gyrinidae and the remaining adephagan families, as was found in previous studies using morphology. Haliplidae are either the sister group of Dytiscoidea or the sister group of a clade comprising Geadephaga and the dytiscoid families. Trachypachidae was placed as the sister group of the rhysodid-carabid clade or of Dytiscoidea. The monophyly of Dytiscoidea including Meru is well supported. Autapomorphies are the extensive metathoracic intercoxal septum, the origin of the metafurca from this structure, the loss of Mm. furcacoxalis anterior and posterior, and possibly the presence of an elongated subcubital setal binding patch. Meruidae was placed as sister group of the Noteridae. Synapomorphies are the absence of the transverse ridge of the metaventrite, the fusion of abdominal segments III and IV, the shape of the strongly asymmetric parameres, and the enlargement of antennomeres 5, 7 and 9. The Meru-noterid clade is the sister group of the remaining Dytiscoidea. The exact position of Aspidytes within this clade remains ambiguous: it is either the sister group of Amphizoidae or the sister group of a clade comprising this family and Hygrobiidae + Dytiscidae. The sister group relationship between Spanglerogyrinae and Gyrininae was strongly supported. The two included genera of Gyrinini form a clade, and Enhydrini are the sister group of a monophylum comprising the remaining Enhydrini and Orectochilini. A branching pattern (Peltodytes + (Brychius + Haliplus)) within Haliplidae was confirmed. Algophilus, Apteraliplus and the Haliplus-subgenus Liaphlus form a clade. The generic status of the two former taxa is unjustified. The Phreatodytinae are the sister group of Noterinae, and Notomicrus (+ Speonoterus), Hydrocoptus, and Pronoterus branch off successively within this subfamily. The search for the larvae of Meru and a combined analysis of morphological and molecular data should have high priority.
本文对肉食亚目进行了系统发育分析。该分析基于成虫和幼虫的148个形态特征，重点关注新描述的沼甲科的分类位置，以及小型水生科豉甲科、水龟虫科和小粒龙虱科的属级系统发育。我们发现，豉甲科与其余肉食亚目科之间存在姐妹群关系，这与之前使用形态学的研究结果一致。水龟虫科要么是龙虱总科的姐妹群，要么是一个包含步甲总科和龙虱总科各科的进化枝的姐妹群。沼梭甲科被置于隐翅甲-步甲进化枝或龙虱总科的姐妹群位置。包括沼甲属在内的龙虱总科的单系性得到了有力支持。其自有衍征包括广泛的后胸基节间中隔、后叉骨源于该结构、前后叉骨基节肌缺失，以及可能存在的延长的肘下刚毛结合斑。沼甲科被置于小粒龙虱科的姐妹群位置。其共同衍征包括后胸腹板横脊的缺失、腹部第三和第四节的融合、强烈不对称的阳茎侧叶的形状，以及触角第5、7和9节的增大。沼甲-小粒龙虱进化枝是其余龙虱总科的姐妹群。隐肢龙虱属在该进化枝中的具体位置仍不明确：它要么是两栖甲科的姐妹群，要么是一个包含该科以及湿水龟科+龙虱科的进化枝的姐妹群。斯氏豉甲亚科和豉甲亚科之间的姐妹群关系得到了有力支持。豉甲族包含的两个属形成一个进化枝，而水栖豉甲族是一个包含其余水栖豉甲族和尖腹豉甲族的单系群的姐妹群。水龟虫科内的分支模式（佩氏水龟虫 +（布氏水龟虫 + 水龟虫属））得到了确认。嗜水龙虱属、无翅水龟虫属和水龟虫属的利亚水龟虫亚属形成一个进化枝。前两个分类单元的属级地位不合理。穴水龟亚科是小粒龙虱亚科的姐妹群，而穴粒龙虱属（+ 斯氏粒龙虱属）、穴水龟属和前粒龙虱属在该亚科内依次分支。寻找沼甲的幼虫以及对形态学和分子数据进行综合分析应具有高度优先权。
