Phylogeny of theNorthAmerican vaejovid scorpion subfamilySyntropinae Kraepelin, 1905, based on morphology, mitochondrial and nuclearDNA

Abstract

The first rigorous analysis of the phylogeny of the North American vaejovid scorpion subfamily Syntropinae is presented. The analysis is based on 250 morphological characters and 4221 aligned DNA nucleotides from three mitochondrial and two nuclear gene markers, for 145 terminal taxa, representing 47 species in 11 ingroup genera, and 15 species in eight outgroup genera. The monophyly and composition of Syntropinae and its component genera, as proposed by Soleglad and Fet, are tested. The following taxa are demonstrated to be para-or polyphyletic: Smeringurinae; Syntropinae; Vaejovinae; Stahnkeini; Syntropini; Syntropina; Thorelliina; Hoffmannius; Kochius; and Thorellius. The spinose (hooked or toothed) margin of the distal barb of the sclerotized hemi-mating plug is demonstrated to be a unique, unambiguous synapomorphy for Syntropinae, uniting taxa previously assigned to different subfamilies. Results of the analysis demonstrate a novel phylogenetic relationship for the subfamily, comprising six major clades and 11 genera, justify the establishment of six new genera, and they offer new insights about the systematics and historical biogeography of the subfamily, and the information content of morphological character systems.

Supplemental Material for 'Systematic revision of the North American syntropine vaejovid scorpion genera Maaykuyak, Syntropis, and Vizcaino, with description of the adults of Syntropis williamsi. (Bulletin of the American Museum of Natural History, 405)'

“…Although Hendrixson (2001) considered V. pequeno the closest relative of M. waueri, this species was found to be more closely related to Serradigitus Stahnke, 1974, andStahnkeus Soleglad andFet, 2006, than to the members of subfamily Syntropinae in the phylogenetic analyses presented by González-Santillán and Prendini (2015a Diagnosis: Syntropis differs from other Syntropinae in the carination of metasomal segments I-IV, where the ventral submedian carinae are absent (indicated by pairs of macrosetae only, no difference in ornamentation evident from the adjacent intercarinal surfaces), and the ventral median carinae distinct (protruding above the adjacent intercarinal surfaces), smooth to finely granular, and unpigmented (figs. 7D-F).…”

“…Although both Vaejovis viscainensis Williams, 1970, andVaejovis vittatus Williams, 1970, were initially assigned to the eusthenura group, Williams (1970a: 412), noted the distinctiveness of V. viscainensis, in particular, noting that it "is of systematic interest because it appears to represent a distinct branch from the eusthe nura group of Vaejovis." Williams (1970b) compared V. viscainensis to two species from the Baja California Peninsula, Vaejovis diazi Williams, 1970, andVaejovis schwenkmeyeri Williams, 1970, from which it INTRODUCTION Four distinct genera, forming two monophyletic groups, were placed basal in the phylogeny of the North American vaejovid scorpion subfamily Syntropinae Kraepelin, 1905, presented by González-Santillán and Prendini (2015a): Kone tontli Maaykuyak González-Santillán and Prendini, 2013;and Vizcaino González-Santillán and Prendini, 2013. The 16 species accommodated within these genera, distributed across the Baja California Peninsula (Syn tropis, Vizcaino, and one species each of Konetontli and Maaykuyak) and the Mexican mainland (nine species of Konetontli and one species of Maaykuyak), represent a morphologically and ecologically diverse assemblage, including three of the four ecomorphotypes observed in the subfamily (figs.…”

“…(Williams, 1970), and H. waueri (Gertsch and Soleglad, 1972). The classification presented by Soleglad and Fet (2008) was not based on quantitative phylogenetic analysis, however, and most of the new taxa proposed therein were demonstrably paraphyletic or polyphyletic when tested by an analysis based on morphological characters and DNA sequences from the nuclear and mitochondrial genomes (González-Santillán and Prendini, 2015a). Whereas the species assigned to Hoffmannius, i.e., the former eusthenura group of Vaejovis, were consistently polyphyletic in these analyses, those assigned to the four basal genera defined by were consistently monophyletic and formed two monophyletic groups: (Konetontli + Maaykuyak) and (Syntropis + Vizcaino).…”

“…When Soleglad and Fet (2008) Kochius Soleglad and, were devised. As mentioned above, the classification presented by these authors was not based on quantitative phylogenetic analysis and most of the new taxa proposed therein proved to be paraphyletic or polyphyletic when tested by González-Santillán and Prendini (2015a). Among other findings, the type species of Hoffmannius and Paravaejovis formed a monophyletic group, to the exception of other species of Hoffmannius and the exemplar species of Paruroctonus and Smeringu rus, rendering Hoffmannius, Smeringurinae, and Syntropinae paraphyletic and requiring the redefinition and relimitation of Syntropinae and its component genera presented by .…”

Systematic Revision of the North American Syntropine Vaejovid Scorpions With a Subaculear Tubercle,KonetontliGonzález-Santillán and Prendini, 2013

“…The subaculear tubercle was suggested as a potential diagnostic synapomorphy for these species (Armas and Martín-Frías, 2001). However, phylogenetic analysis (Gonzá lez-Santillán and Prendini, 2015) demonstrated that they do not form a monophyletic group and the subaculear tubercle evolved on multiple independent occasions in Vaejovidae Thorell, 1876. The monophyletic group of five species assigned to Konetontli by Gonzá lez-Santillá n and Prendini (2013) was not monophyletic with other vaejovids possessing a subaculear tubercle, i.e., the exemplar species of the mexicanus group of Vaejovis C.L.…”