The Phylogeny of Living and Extinct Pangolins (Mammalia, Pholidota) and Associated Taxa: A Morphology Based Analysis

Skeletal Anatomy of the North American Pangolin Patriomanis americana (Mammalia, Pholidota) from the Latest Eocene of Wyoming (USA)

“…We describe all the known material of Patriomanis and compare its anatomy to that of the other fossil pangolins. As has been noted elsewhere (e.g., Gaudin et al, 2006; Gaudin, 2010), the fossil record of Pholidota is poor, with very few extinct taxa (McKenna and Bell, 1997;Gaudin et al, 2009). That said, the taxa that do exist are reasonably well known, including Eomanis (middle Eocene of Germany; Storch, 1978Storch, , 2003, Euromanis (middle Eocene of Germany; Storch and Martin, 1994;Gaudin et al, 2009), Crypto manis (late Eocene of China; Gaudin et al, 2006), and Necromanis (Oligocene-Miocene of central Europe; Koenigswald and Martin, 1990; Koenigswald, 1999; see also Storch, 2003;Gaudin et al, 2009;Hoffmann et al, 2009;1999).…”

“…As has been noted elsewhere (e.g., Gaudin et al, 2006; Gaudin, 2010), the fossil record of Pholidota is poor, with very few extinct taxa (McKenna and Bell, 1997;Gaudin et al, 2009). That said, the taxa that do exist are reasonably well known, including Eomanis (middle Eocene of Germany; Storch, 1978Storch, , 2003, Euromanis (middle Eocene of Germany; Storch and Martin, 1994;Gaudin et al, 2009), Crypto manis (late Eocene of China; Gaudin et al, 2006), and Necromanis (Oligocene-Miocene of central Europe; Koenigswald and Martin, 1990; Koenigswald, 1999; see also Storch, 2003;Gaudin et al, 2009;Hoffmann et al, 2009;1999). Given their slow movements, alongside the low diversity and abundance at which their populations exist, and their sluggish reproductive rate (litter size typically one, occasionally two or three in Asiatic species, birthed once a year in all but African tree pangolins, which may breed two or three times in a year; Kingdon, 1974Kingdon, , 1997Heath, 1992a Heath, , 1992b Heath, , 1995Nowak, 1999), they do not seem like a group that would be likely to spread quickly or to have a broad, global distribution.…”

“…All three are basal apomorphies of Pholidota (Gaudin et al, 2009). Additional preparation of the posterior reaches of the palate failed to reveal any remains of the palatine bones, so the location and form of both the maxillary-palatine suture and the palatine foramina cannot be ascertained.…”

“…The pangolins themselves, relegated to their own order, the Pholidota, are an extraordinary group of placental mammals. Rare, secretive, solitary, and largely nocturnal, the animals in this group comprise only eight living species distributed across sub-Saharan Africa, the Indian subcontinent, Southeast Asia, Indonesia, and the westernmost Philippines (Corbet and Hill, 1991;Nowak, 1999;Gaubert and Antunes, 2005, Schlitter, 2005;Gaudin et al, 2009). Thus, they are relatively poorly known even in the parts of the globe they frequent and virtually unheard of outside those areas.…”

“…In the only other fossil pangolin that preserves the premaxilla, Eomanis waldi, the leading edge of the bone is roughly vertical and distinctly concave anteriorly, giving the whole dorsal process a C shape in lateral view (Storch, 2003;. This morphology is also present in the palaeanodont Metachei romys and in the enigmatic edentate mammal Eurotaman dua, making it a synapomorphy of Pholidotamorpha (as defined by Gaudin et al, 2009) that is subsequently modified in patriomanids and manids. Emry (2004:131) described the incomplete posteromedian process on the palatal portion of the premaxilla as "essentially like that of M.…”