The herbivorous Chalicotheriidae from Oligocene to Pleistocene are unique among perissodactyls in possessing claws rather than hooves (Coombs, 1983). The ungual phalanges of early chalicotheres, however, were almost unknown previously due to a sparse fossil record. Lucas and Schoch (1989) briefly mentioned that the Eocene chalicothere Grangeria canina possesses deep, narrow, and shallow fissured claws, but did not provide any more descriptions or illustrations. Recently, new specimens of the early Eocene chalicothere Litolophus gobiensis, including unguals and other parts of multiple skeletons, were unearthed from the Arshanto Formation in the Erlian Basin, Inner Mongolia, China (Meng et al., 2007;Bai et al., 2010;Wang et al., 2010). The Litolophus assemblage was caused by a catastrophic event, and L. gobiensis, represented by at least 24 individuals, was the most abundant larger mammal in the quarry (Bai et al., 2011). The species retains a tetradactyl hand with functional phalanges from digit II to digit V, which is a plesiomorphic character among perissodactyls. Phylogenetic analysis shows that L. gobiensis was a primitive member of the clade and basal to all other known chalicotheres apart from earliest Eocene forms . These new materials shed light on the ungual evolution of chalicotheres.
MATERIALS AND METHODSThirty isolated ungual phalanges are preserved in the assemblage, but most of them are immature and susceptible to breakage (IVPP V 16187.1-21). The material includes two mature unguals of digit III (V 16187.1-2), 12 mature and/or immature unguals of digit II (V 16187.3-14), five unguals of digit IV (V 16187.15-19), two immature unguals of manus digit V (V 16187.20-21), and nine unguals indeterminate to digit (V 16187.22-30). An immature right articulated pes of L. gobiensis preserves the ungual phalanges of digits II through IV (V 16200).Institutional Abbreviation-IVPP, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing.Anatomical Abbreviations-da, distal angle; dp, dorsal process; lf, lateral foramen; mf, medial foramen; sn, sagittal notch; sp, subungual process; vf, volar foramen; ph2f, phalanx 2 facet.
MORPHOLOGICAL DESCRIPTION
Ungual Phalanx of Digit IIIAn ungual phalanx of digit III (V 16187.1) referred to a mature pes is well preserved. The proximal facet for phalanx 2 is rectangular in outline, wider than high, concave dorsoventrally, and divided by a rather weak median ridge into two nearly equal portions (Fig. 1A).The proximal border of the dorsal surface is slightly arched (Fig. 1B), and most of the dorsal surface is rough and porous. The distally placed sagittal notch is short and shallow, similar to * Corresponding author. a trough cutting the dorsodistal portion of the bone. A large dorsal foramen is located on either side of the sagittal notch, near the distal margin. The distal angles are prominent and pointed at the ends, whereas proximal angles are absent.The dorsal part of the distal end is covered by irregular foramina or grooves, except...