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White-Bellied Pangolin Phataginus Tricuspis (Rafinesque, 1820) Raymond Jansen1,2, Olufemi Sodeinde3, Durojaye Soewu4, Darren W

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White-Bellied Pangolin Phataginus Tricuspis (Rafinesque, 1820) Raymond Jansen1,2, Olufemi Sodeinde3, Durojaye Soewu4, Darren W CHAPTER 9 White-bellied pangolin Phataginus tricuspis (Rafinesque, 1820) Raymond Jansen1,2, Olufemi Sodeinde3, Durojaye Soewu4, Darren W. Pietersen5,6, Daniel Alempijevic7 and Daniel J. Ingram8 1Department of Environmental, Water and Earth Sciences, Tshwane University of Technology, Pretoria, South Africa 2African Pangolin Working Group, Pretoria, South Africa 3Department of Biological Sciences, New York City College of Technology, City University of New York, Brooklyn, NY, United States 4Department of Fisheries and Wildlife Management, College of Agriculture, Ejigbo Campus, Osun State University, Osogbo, Nigeria 5Mammal Research Institute, Department of Zoology and Entomology, University of Pretoria, Hatfield, South Africa 6IUCN SSC Pangolin Specialist Group, N Zoological Society of London, Regent’s Park, London, United Kingdom 7Integrative Biology, Florida Atlantic University, Boca Raton, FL, United States 8African Forest Ecology Group, Biological and Environmental Sciences, University of Stirling, Stirling, United Kingdom OUTLINE Taxonomy 140 Ontogeny and reproduction 150 Description 140 Population 151 Distribution 145 Status 152 Habitat 146 Threats 152 Ecology 147 References 153 Behavior 148 Pangolins DOI: https://doi.org/10.1016/B978-0-12-815507-3.00009-5 139 © 2020 Elsevier Inc. All rights reserved. 140 9. White-bellied pangolin Phataginus tricuspis Taxonomy Etymology: Phataginus (see Chapter 8); the species name refers to the three (tri-) points or Previously included in the genus Manis cusps (-cuspis) on the scales (Gotch, 1979). (Meester, 1972; Schlitter, 2005), the species is here included in Phataginus based on both morphological (Gaudin et al., 2009) and Description genetic (du Toit et al., 2017; Gaubert et al., 2018) evidence. Hatt (1934) reported that the The white-bellied pangolin (Phataginus tri- species exhibits slight morphological variation cuspis) is a small, semi-arboreal African pango- across its range, including in scale pigmenta- lin with a body weight of 1À3 kg and a total tion and hair length (e.g., on the ventrum). length of about 100 cm (Table 9.1). It is the Allen and Loveridge (1942) and Meester (1972) lightest pangolin species, marginally lighter proposed two distinct subspecies based on mor- than the broadly sympatric black-bellied pan- phological analyses, P. tricuspis tricuspis (range golin (P. tetradactyla). The species is not sexu- except Uganda) and P. t. mabirae (Uganda). ally dimorphic but males are reported to be These subspecies are not considered valid here slightly longer and heavier than females following Kingdon and Hoffmann (2013). (Page`s, 1968). Up to 60% of the total body The species is included in the subfamily length comprises the long, prehensile tail Phatagininae to designate small African pan- (35À60 cm); head-body length ranges between golins based on mitogenomic distances 25 and 38 cm, potentially longer (Table 9.1). between the genera Manis, Smutsia, and The tail is flat ventrally and rounded dorsally, Phataginus, and morphological traits unique to and has a naked cutaneous pad at the tip Phataginus (see Chapter 2; Gaubert et al., 2018). (which is truncated) on the ventral side that Gaubert et al. (2016) identified six geographic replaces two median and two lateral scales lineages within the white-bellied pangolin that (Pocock, 1924). The sensory tail pad contains were identified as Evolutionarily Significant many mechanoreceptor nerve endings that aid Units (ESUs) and may warrant species or sub- sensitivity in terms of touch and grip (Doran and species status; this requires further research. Allbrook, 1973), and the tail functions as a fifth These lineages were partitioned into Central limb when the species is climbing (see Behavior). Africa, Gabon, Dahomey Gap, Ghana, western There are 41 caudal vertebrae (Table 9.1). Africa, and western Central Africa (see The body is covered in small, overlapping Chapter 2). Hassanin et al. (2015) discovered a scales that grow from the epidermis in a high genetic nucleotide divergence between a lattice-like arrangement and are tricuspid white-bellied pangolin from Gabon and speci- (Jentink, 1882; Rahm, 1956). The scales cover mens from Cameroon, Ghana and Nigeria, and the dorsal and lateral surfaces of the body, the the potential for a new taxon in Gabon fore- (except the upper portions) and hin- requires further research. dlimbs, the tail (dorsally and ventrally), and The type locality is given as “Guine´e”, the neck and head, but are absent from the West Africa (Rafinesque, 1820). Chromosome anterior of the face, the ventral body surface number is not known. Synonyms: Manis multi- and inner-side of the limbs. On the upper por- scutata (Gray, 1843), Manis tridentata (Focillon, tion of the forelimbs (elbow to wrist) dense 1850), P. t. mabirae (Allen and Loveridge, (sometimes long) brown hair replaces the 1942). scales (Pocock, 1924). There are 18À22 I. What is a Pangolin? Description 141 TABLE 9.1 White-bellied pangolin morphometrics. Measurement Country Source(s) Weight Weight (R)a 1.67 (1.2À2.3) kg, n 5 8 Southwest Nigeria FMNHb 2.36 (1.74À2.86) kg, n 5 4 Southeast Nigeria Kingdon and Hoffmann, 2013 Weight (Q) 1.71 (1.2À2.2) kg, n 5 8 Southwest Nigeria FMNHa 2.6 (1.94À2.88) kg, n 5 11 Southeast Nigeria Kingdon and Hoffmann, 2013 Body Total length (R) 793.2 (617À1027) mm, n 5 25 Democratic Republic of Hatt, 1934 the Congo Total length (Q) 768.4 (630À920) mm, n 5 25 Democratic Republic of Hatt, 1934 the Congo Head-body length (R) 350 (330À380) mm, n 5 7Coˆte d’Ivoire, Democratic FMNHb Republic the of Congo, Liberia 319 (254À375) mm, n 5 17 Southeast Nigeria Kingdon and Hoffmann, 2013 Head-body length (Q) 333 (308À367) mm, n 5 8Coˆte d’Ivoire, Democratic FMNHb Republic of the Congo, Ghana, Liberia 310 (265À351) mm, n 5 14 Southeast Nigeria Kingdon and Hoffmann, 2013 Tail length (R) 469.6 (360À607) mm, n 5 25 Democratic Republic of Hatt, 1934 the Congo Tail length (Q) 460.4 (350À590) mm, n 5 25 Democratic Republic of Hatt, 1934 the Congo Vertebrae Total number 69 Frechkop, 1931; Jentink, of vertebrae 1882 Cervical 7 Jentink, 1882; Mohr, 1961 Thoracic 13 Jentink, 1882; Mohr, 1961 Lumbar 6 Jentink, 1882; Mohr, 1961 Sacral 2 Jentink, 1882; Mohr, 1961 Caudal 41 Jentink, 1882; Mohr, 1961 Skull Length (R) 72.8 (63.8À80.8) mm, n 5 22 Democratic Republic of Hatt, 1934 the Congo Length (Q) 68.7 (58.5À79.2) mm, n 5 20 Democratic Republic of Hatt, 1934 the Congo (Continued) I. What is a Pangolin? 142 9. White-bellied pangolin Phataginus tricuspis TABLE 9.1 (Continued) Measurement Country Source(s) Breadth across 27.3 (22.7À32) mm, n 5 22 Democratic Republic of Hatt, 1934 zygomatic the Congo processes (R) Breadth across 25.4 (20.2À29.3) mm, n 5 20 Democratic Republic of Hatt, 1934 zygomatic the Congo processes (Q) Scales Total number 935 (794À1141), n 5 25 Cameroon, Nigeria, Sierra Ullmann et al., 2019; of scales Leone, Uganda, unknown D. Soewu, unpubl. data No. of scale rows 18À22 Democratic Republic of Frechkop, 1931; Hatt, 1934 (transversal, body) the Congo No. of scale rows 19À25 Democratic Republic of Frechkop, 1931; Hatt, (longitudinal, body) the Congo 1934; Jentink, 1882 No. of scales on outer 35À40 Democratic Republic of Frechkop, 1931; Hatt, margins of tail the Congo 1934; Jentink, 1882 No. of scales 30À33 Democratic Republic of Frechkop, 1931; Hatt, 1934 on median the Congo row of tail Scales (wet) as 0.15À0.20 African Wildlife proportion Foundation, unpubl. data of body weight Scales (dry) as No data proportion of body weight aMales are reported to be slightly heavier than females; these data are based on a small sample size. bFMNH=Florida Museum of Natural History. transversal and 19À25 longitudinal scale rows Jansen, pers. obs.). The scales grow in length on the body, and 35À40 rows of scales on the more rapidly than in width: scales from the outer margins of the tail that are kite-shaped mid-dorsal region grow from being approxi- (Frechkop, 1931; Hatt, 1934; Jentink, 1882). The mately as long as wide, to four times length medio-dorsal row of scales on the tail is inter- versus width. The scales are laterally striated rupted towards the tail tip and is replaced by (Fig. 9.2) and keeled on the flanks and limbs, two rows of 3À6 scales (Jentink, 1882). Total and are less robust (i.e., are flimsy) compared scales number between B790 and 1140 to scales on the black-bellied pangolin and (Table 9.1). The largest scales are on the dorsal other pangolin species. The striations and surface and measure up to 47 3 26 mm; single cusps wear over time, sometimes completely, scales weigh up to 0.66 g (O. Sodeinde, and the cusps may break off with age (Hatt, unpubl. data). The scales are uniform in color 1934; R. Jansen, pers. obs.). Unlike Asian pan- but vary substantially from brownish-gray to golins, hairs do not project between the scales. reddish- or yellowish-brown, and the distal The skull demonstrates morphological fea- edge may color faint yellow, especially in older tures related to adaption to a myrmecophagous individuals (Figs. 9.1À9.3; Rahm, 1956;R. diet (Chapter 1; see Hatt, 1934; Heath, 2013). I. What is a Pangolin? Description 143 FIGURE 9.1 White-bellied pangolin foraging for prey in Central African Republic. The cusps on the scales can clearly be seen on this individual. Photo credit: Alex Ley. As in the black-bellied pangolin, the lacrimal and, as in other pangolin species, is attached to bone is present À it is absent in other pangolin the caudal end of the xiphoid process (xiphi- species (Emry, 1970). The head is conical- sternum) in the abdomen (Doran and Allbrook, shaped and the muzzle is broader than that of 1973). In this species, the proximal region of the black-bellied pangolin (Hatt, 1934).
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