X-ray microtomography of the Late Carboniferous whip scorpions (Arachnida, Thelyphonida) Geralinura britannica and Proschizomus petrunkevitchi

Whip scorpions (Thelyphonida) and schizomids (Schizomida) are closely related arachnid orders, whose low species diversity compared to other arachnid groups is reflected in a limited fossil record. Here we investigate two key fossil whip scorpions from the British Middle Coal Measures of Coseley, Staffordshire (Late Carboniferous, ca. 315 Ma), UK using X-ray microtomography, and incorporate the taxa into an updated arachnid phylogeny. Geralinura brittanica Pocock, 1911 is an unequivocal whip scorpion, whose pedipalps are closer to the modern condition than previously assumed. The trochanter bears a dorsal flange with prodorsal teeth, and the whole pedipalp is rendered subchelate by the presence of at least a patellar apophysis. These results suggest a high degree of stasis in whip scorpions since the Late Carboniferous, and the genus Geralinula Scudder, 1884 is within the crown-group; i.e. the extant family Thelyphonidae. Proschizomus petrunkevitchi Dunlop & Horrocks, 1996 lacks median eyes, a character shared with Schizomida, but unlike schizomids the pedipalp also has patellar and tibial apophyses; albeit with the appendage articulating at an angle of ~45° rather than articulating horizontally as in living whip scorpions, or vertically as in schizomids. Our phylogeny refutes a previous hypothesis that P. petrunkevitchi is a stem-schizomid, and places it among the whip scorpions instead. However, the species may have branched early from the whip scorpion lineage, and thus its morphology might reflect a plesiomorphic arrangement in which the pedipalps were transitioning from a more leg-like to a fully raptorial and claw-like set of prey-catching appendages.