Revision and analysis of Pseudosaldula Cobben (Insecta, Hemiptera, Saldidae) : a group with a classic Andean distribution. (Bulletin of the American Museum of Natural History, no. 323)
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Abstract
The genus Pseudosaldula Cobben, which is restricted to the Andean Subregion of South America, is revised. Fourteen valid species are recognized, nine of them being described as new and 10 previously published names are treated as junior synonyms based on the examination of approximately 3500 specimens from Colombia, Ecuador, Peru, Bolivia, Argentina, and Chile. All taxa are described or redescribed. A key to the species is provided. Color habitus illustrations, distributional maps, and detailed measurements are provided for all species. Scanning electron micrographs of the vestiture, parameres, parandria, face, and pretarsus are provided for representative species, as are color views of the face and the nymphs. The concept of a postclypeus in the Saldidae is questioned and the term transverse swelling, as coined by Parsons, is applied in discussing distinctive aspects of facial morphology in Pseudosaldula. A previously unreported, presumably glandular, pore is documented on the parameres in the Saldinae in the form of a cavernous pit with internal digitiform processes. A phylogenetic analysis based on morphological character data documents the monophyly of Pseudosaldula. Characters treated as synapomorphic for Pseudosaldula are five cells in the membrane of the forewing, the incomplete connection of the transverse swelling across the posterior margin of the clypeus, and the straight connection across the posterior margin of the parandria; nymphal coloration is also distinctive, although treated as ambiguous because this character was not scored for all species. DNA sequence data from the 16S rDNA region of the mitochondrion and H3 nuclear region were acquired for 13 Pseudosaldula spp. and five outgroup taxa. The combined analysis of morphological and sequence data consistently treated Pseudosaldula as paraphyletic. These results are interpreted as the result of inadequate sampling of both taxa and gene regions, in light of the fact that the patterns of distribution become transpacific, as opposed to a monophyletic group in the Andean Region. Not unexpectedly, several morphological characters documenting the monophyly of Pseudosaldula show greater homoplasy in the combined analysis than when analyzing morphological data alone. Therefore, the results of the morphological cladistic analysis are further used to examine distributional patterns in the group. Five areas of endemism are recognized: northern Andes, northern Peru, Puna, central Chile, and subantarctic; the boundaries of these areas show substantial correspondence with those proposed for other groups of insects.