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Hemidactylus nzingae CERÍACO, AGARWAL, MARQUES & BAUER<br />BAUER, 2020

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Higher TaxaGekkonidae, Gekkota, Sauria, Squamata (lizards: geckos) 
Subspecies 
Common NamesE: Queen Nzinga’s Tropical Gecko
Portuguese: Osga da Rainha Nzinga 
SynonymHemidactylus nzingae CERÍACO, AGARWAL, MARQUES & BAUER
BAUER 2020: 48
Hemidactylus muriceus — PETERS 1881: 147
Hemidactylus muriceus — BOCAGE 1895: 13
Hemidactylus longicephalus — SCHMIDT 1933: 4 [part]
Hemidactylus longicephalus — MANAÇAS 1963: 227
Hemidactylus brookii angulatus — LAURENT 1964: 29
Hemidactylus cf. muriceus — CERÍACO et al. 2016: 59
Hemidactylus cf. muriceus — CERÍACO et al. 2018: 422
Hemidactylus cf. muriceus — MARQUES et al. 2018: 186
Hemidactylus murecius [sic] — BRANCH et al. 2019: 315 
DistributionAngola (Malanje, Uíge, Bíe, Huambo, Lunda Sul, Lunda Norte)

Type locality: Cangandala National Park (-9.872667°, 16.700889°, 1092 m), Malanje Province, Republic of Angola  
Reproduction 
TypesHolotype. CAS 258418 (field number Jvv 9224; Fig. 24; Table 5), adult male, collected by Luis Ceríaco, Mariana Marques, Suzana Bandeira, Edward Stanley and Jens Vindum on 17 September 2015.
Paratypes. CAS 258422, adult male, Cangandala National Park (-9.81011°, 16.65564°, 1086 m), Malanje Province, collected by Luis M. P. Ceríaco, Mariana P. Marques, Suzana Bandeira, Edward L. Stanley and Jens vindum on 26 September 2015; MHNCUP/Rep-464 (field number Jvv 9473), adult male, Cangandala National Park (-9.872667°, 16.700889°, 1092 m), Malanje Province, collected by Luis M. P. Ceríaco, Mariana P. Marques, Suzana Bandeira, Edward L. Stanley and Jens vindum on 10 October 2015; AMNH 47786, adult female, Huambo (-12.776111°, 15.739167°, 1700 m), Huambo Province, collected Herbert Lang and W. Rudyerd Boulton in 1925; AMNH 47782, adult male, Dande (-11.166667°, 17.166667°, 1126 m), Malanje Province, collected by Herbert Lang and W. Rudyerd Boulton in 1925; AMNH 47774, adult male, Chitau (-11.424927°, 17.154407°, 1506 m), Bié Province, collected by Herbert Lang and W. Rudyerd Boulton in August 1925; ANSP 32189, male adult, Pavalange (-10.969765°, 17.612431°, 1127 m), Malanje Province, collected by Harold T. Green in October 1930; CM 5724, male adult, Gaúca (-11.18333°, 17.45°, 1243 m) Bié Province, collected by W. Rudyerd Boulton and Laura Boulton on 10 January 1931; FMNH 18476, male adult, with the same collecting details as the previous paratype; IICT 4-1957 and 5-1957, two female adults, both from Fazenda OtíliaEncoge (-7.55°, 15.03333°, 765 m), Uíge Province, collected by Fernando Frade on 26 August 1957; MD 4008, adult female, Alto-Cuílo (-10.049015°, 19.515477°, 1259 m), Luanda Sul Province, collected by António Barros Machado in June 1954. 
DiagnosisDiagnosis. A medium sized Hemidactylus, maximum snout-vent length 51.5 mm (Fig. 23). Dorsal pholidosis heterogeneous, with 17–21 irregularly arranged longitudinal rows of subtrihedral, striated, strongly keeled tubercles at midbody. Two well developed pairs of postmentals, the inner pair longer than the outer pair, but about the same size as the mental, and in broad contact behind the mental. ventrolateral folds distinct, about 23–27 scale rows across venter. Five or six divided scansors beneath first digit of both manus and pes, 6–8 beneath fourth digit of manus, nine (rarely eight) beneath the fourth digit of pes. Males with a continuous series of 6–8 precloacal pores. Original tail roughly cyclotetragonal, strongly spinose; scales on the tail slightly larger than dorsals of trunk, striated, imbricate, with a longitudinal series of two enlarged, strongly keeled, striated, pointed tubercles on either side of the median dorsal furrow. Subcaudal scales small and imbricate, about one fifth of tail width. Body dorsum with a series of dark-brown “W-shaped” transverse markings from occiput to sacrum, bordered by a cream-colored stripe extending from behind the eyes to the insertion of the legs; tail with distinct alternating light and dark bands.

Comparison with West and Central African congeners. Hemidactylus nzingae sp. nov.is readily distinguished from H. kamdemtohami and H. richardsonii by the lack of basal digital webbing and from H. matschiei by having spiny tubercles on the dorsum and small subcaudal scales. It may be distinguished from H. steindachneri by lacking a longitudinal row of keeled tubercles on the the ventrolateral border of flanks, from H. hecqui by having a a stereotypic color pattern featuring paired dark markings along the entire length of the body dorsum (absent in H. hecqui, which instead bears a series of thin pale crossbands; Fig. 12). Hemidactylus nzingae sp. nov. can be distinguished from H. longicephalus sensu stricto by having a higher number of tubercle rows on the dorsum (18–21 versus 17–18), fewer granular scales between the dorsal tubercles (2–3 versus 3–6), fewer rows of ventral scales (22–27 versus 30–33), by having strongly keeled and pointed dorsal and caudal tubercles (versus feebly keeled in H. longicephalus); and by its distinctive dark W-shaped markings on the dorsum and well-marked dorsolateral light stripes (dorsal markings incomplete or inconspicuous, and dorsolateral striping absent or not evident in H. longicephalus). It may be distinguished from H. benguellensis by a much lower number of precloacal-femoral pores in males (6–8 versus 23–33), more tubercle rows on the dorsum (18–21 versus 13–18), fewer granular scales between the dorsal tubercles (2–3 versus 4–9), and by its distinctive dark W-shaped marking on the dorsum and well-marked dorsolateral light stripes (marking less evident or inconspicuous, dorsolateral striping absent, dorsolateral row of tubercles orange in H. benguellensis). Hemidactylus benguellensis also has a row of pointed tubercles separating the venter and the dorsum and the flanks are without tubercles, whereas in H. nzingae sp. nov. this ventrolateral row of tubercles is lacking, and tubercles extend onto the flanks. The new species can be distinguished from H. bayonii by its larger size (maximum SvL 51.5 mm versus 36.2 mm), by having higher number of tubercle rows on the dorsum (18–21 versus 14–16), fewer granular scales between the dorsal tubercles (2–3 versus 4–7), fewer rows of ventral scales (22–27 versus 28–33), by having strongly keeled and pointed dorsal and caudal tubercles (versus feebly keeled in bayonii), and by its very distinctive, dark W-shaped markings on the dorsum, dorsolateral light stripes starting posteriorly to the eyes and gradual coloration transition from lateral side to venter (marking constituting pairs of marks across dorsum, dorsolateral striping starting on the snout, ventral coloration strongly contrasting with flank coloration in H. bayonii). Hemidactylus nzingae sp. nov. is easily distinguishable from H. mabouia by its smaller size (maximum SvL 51.5 mm versus 67.4 mm), lower number of precloacal-femoral pores in males (6–8 versus 28–39), higher number of tubercle rows on the dorsum (18–21 versus 12–19), fewer granular scales between the dorsal tubercles (2–3 versus 5–10), lower number of rows of ventral scales (22–27 versus 34–37), by having strongly keeled and pointed dorsal and caudal tubercles (versus feebly keeled in H. mabouia), and by its very distinctive, dark W-shaped markings on the dorsum and well-marked dorsolateral light stripes (ill-defined or complete lack of markings in H. mabouia). The median subcaudal scales form transversely enlarged scutes in H. mabouia, about half of tail width in extent, whereas in the new species the subcaudals are small, approximately one fourth of tail width.
Hemidactylus nzingae sp. nov. can be distinguished from H. muriceus sensu stricto by a higher number of tubercle rows on the dorsum (17–21 versus 7–12), fewer granular scales between the dorsal tubercles (2–3 versus 5–10); and fewer nasals surrounding the nostril (2 versus 3). The new species can be distinguished from H. pseudomuriceus by having the median subcaudal scales small and similar to those of the ventrolateral aspect of the tail (median subcaudals enlarged and hexagonal in H. pseudomuriceus), fewer precloacal-femoral pores in males (6–8 versus 14–17), a higher number of tubercle rows on the dorsum (17–21 versus 6–14), fewer granular scales between the dorsal tubercles (2–3 versus 5–10); and fewer nasals surrounding the nostril (2 versus 3). The newly described species can be distinguished from H. echinus by lacking the two rows of scattered spiny tubercles on the tail venter (a character that differentiates H. echinus from all other known Hemidactylus), having a higher number of tubercle rows on the dorsum (17–21 versus 13–15), and by being smaller (max SvL 51.5 mm versus 68.0 mm). Hemidactylus nzingae sp. nov. differs from H. ansorgii by a higher number of tubercle rows on the dorsum (17–21 versus 6–12), a lower number of precloacal-femoral pores (6–8 versus 10–11), fewer nasals surrounding the nostril (2 versus 3), a stockier body (H. ansorgii has a slender body and narrow head with a short interorbital distance). Hemidactylus ansorgii also has a row of pointed tubercles separating the venter and the flanks, which are completely devoid of any tubercles, (see figure in Tornier 1902 and Perret 1975), whereas in H. nzingae sp. nov. this ventrolateral row of tubercles is lacking, and tubercles extend across the flanks and dorsum. 
Comment 
EtymologyThe species is named after Ngola Nzinga Mbande (1583–1663), also known by the Portuguese Catholic name of Ana de Sousa, queen of the kingdoms of Ndongo and Matamba and one of the most important Angolan chieftains, whose kingdoms spanned approximately the same region as the known distribution of the new species. 
References
  • Bocage,J.V. du B. 1895. Herpétologie d'Angola et du Congo. Lisbon: Imprimerie Nationale, i-xx, 203 pp.
  • Branch, William R.; Pedro Vaz Pinto, Ninda Baptista, and Werner Conradie 2019. The Reptiles of Angola: History, Diversity, Endemism and Hotspots. Chapter 13 in: B. J. Huntley et al. (eds.), Biodiversity of Angola. Springer Verlag, pp. 283-334 - get paper here
  • Ceríaco, L M P; Marques, M P; Bandeira, S; Blackburn, D C & Bauer, A M 2018. Herpetological survey of Cangandala National Park, with a synoptic list of the amphibians and reptiles of Malanje Province, Central Angola. Herpetological Review 49 (3): 408-431 - get paper here
  • Ceríaco, L.M.P., Marques, M.P., Bandeira, S.A. 2016. Anfíbios e Répteis do Parque Nacional da Cangandala. Instituto Nacional da Biodiversidade e Áreas de Conservação & Museu Nacional de História Natural e da Ciência, Luanda & Lisboa, 96 pp
  • CERÍACO, LUIS M. P.; ISHAN AGARWAL, MARIANA P. MARQUES, AARON M. BAUER 2020. A review of the genus Hemidactylus Goldfuss, 1820 (Squamata: Gekkonidae) from Angola, with the description of two new species. Zootaxa 4746 (1): 1-71. - get paper here
  • Laurent, R.F. 1964. Reptiles et batraciens de l'Angola (troisième contribution). Companhia de Di amantes de Angola (Diamang), Serviços Culturais, Museu do Dundo (Angola), No. 67, 165 pp.
  • Manacas,S. 1963. Saurios de Angola. Mem. Junta Invest. Ultram. 2nd ser. 43: 223-240
  • Marques, Mariana P.; Luis M. P. Ceríaco , David C. Blackburn , and Aaron M. Bauer 2018. Diversity and Distribution of the Amphibians and Terrestrial Reptiles of Angola -- Atlas of Historical and Bibliographic Records (1840–2017). Proc. Cal. Acad. Sci. (Ser. 4) 65: 1-501 (Supplement II)
  • Peters, Wilhem Carl Hartwig 1881. Zwei neue von Herrn Major von Mechow während seiner letzten Expedition nach West-Afrika entdeckte Schlangen und eine Übersicht der von ihm mitgebrachten herpetologischen Sammlung. Sitzungsber. Ges. naturf. Freunde Berlin 1881 (9): 147-150 - get paper here
  • Schmidt, Karl Patterson 1933. The reptiles of the Pulitzer Angola Expedition. Annals of the Carnegie Museum 22 (1): 1-15
 
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