The coral bugs, genus Halovelia Bergroth (Hemiptera, Veliidae). II. Taxonomy of the H. malaya-group, cladistics, ecology, biology, and biogeography

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Abstract
Marine bugs of the genus Halovelia Bergroth inhabit intertidal coral reefs and rocky coasts along the continents and islands bordering the Red Sea, Indian Ocean, and western Pacific Ocean as well as island groups and atolls in these areas. In Part I of this work, the genus Halovelia was redescribed together with five previously known species; fifteen new species were described. In the present part, two previously known species are redescribed and eight species described as new, all belonging to the H. malaya Esaki-group: H. sulawesi sp.n. (Sulawesi); H. abdominalis sp.n. (Java, West Malaysia); H. nicobarensis sp.n. (Nicobar Islands); H. convexa sp.n. (Maldive Islands); H. poissoni sp.n. (Kenya, Tanzania); H. seychellensis sp.n. (Seychelles, Madagascar); H. depressa sp.n. (Madagascar); and H. mauricensis sp.n. (Mauritius). A key to the species of the H. malaya-group is included. Using the computer programs PAUP and Hennig86, a cladistic analysis of relationships between the species of Halovelia was performed. Other genera of Haloveliinae were used as outgroup taxa. 46 characters (each with 2-4 states) are listed. The cladistic analysis of the character state matrix yields 18 equally parsimonious cladograms, each 155 steps long. The preferred cladogram is evaluated both by characters and by clades. An account is given of the ecology and biology of the coral bugs, chiefly based upon original observations by the author. The distributions of each of the 30 species of Halovelia are mapped and discussed. The historic biogeography of the species is analysed using two different methods of cladistic (or vicariance) biogeography: component and parsimony analysis. Reduced area cladograms are produced for most species-groups as well as a summary cladogram for these groups. The biogeographic history of Halovelia is discussed in the light of these results and compared with the biogeography of other marine Haloveliinae, the marine Gerridae, and other groups of Indo-Pacific animals.

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