Systematics and biogeography of the family Scorpionidae (Chelicerata : Scorpiones), with a discussion on phylogenetic methods
A cladistic analysis of relationships among the genera of Scorpionidae Latreille, 1802—Heterometrus Ehrenberg, 1828; Opistophthalmus C. L. Koch, 1837; Pandinus Thorell, 1876; and Scorpio Linnaeus, 1758—based on morphology and DNA sequence data from loci of three genes in the mitochondrial genome (12S ribosomal DNA (rDNA), 16S rDNA and cytochrome oxidase I) and one gene in the nuclear genome (28S rDNA) is presented. The analysis makes use of exemplar species, specifically selected to test the monophyly of the genera, rather than supraspecific terminal taxa. Other methods used in the analysis are justified in the context of a discussion of current methods for phylogenetic reconstruction. Relationships among the scorpionid genera are demonstrated to be as follows: (Opistophthalmus (Scorpio (Heterometrus + Pandinus))). This reconstruction identifies Opistophthalmus as the basal lineage of the Scorpionidae, rather than the sister-group of Scorpio. Revised descriptions, diagnoses and a key to identification of the four scorpionid genera are provided, together with a summary of what is known about their ecology, distribution and conservation status.