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Naja peroescobari CERÍACO, MARQUES, SCHMITZ & BAUER, 2017

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Higher TaxaElapidae, Colubroidea, Alethinophidia, Serpentes, Squamata (snakes) 
Subspecies 
Common NamesPortuguese: Cobra-preta
E: São Tomé cobra 
SynonymNaja (Boulengerina) peroescobari CERÍACO, MARQUES, SCHMITZ & BAUER 2017
Naja haje var. nigra — BOCAGE 1866: 51
Naja haje var. nigra — BOCAGE 1879: 87Naja haje — GREEF 1884: 47
Naja haje — BOCAGE 1886: 69
Naja haje — BOCAGE 1895: 24
Naja haje var. nigra — VIEIRA 1886: 237
Naja haje var. nigra — BOCAGE 1889: 34
Naia melanoleuca — FERREIRA 1902: 133
Naja melanoleuca — BOCAGE 1905: 94
Naia melanoleuca — BOULENGER 1906: 215
Naja melanoleuca — THEMIDO 1941: 6
Naja melanoleuca melanoleuca — MANAÇAS 1958: 190
Naja melanoleuca melanoleuca — CAPOCACCIA 1961: 299
Naja melanoleuca — MANAÇAS 1973: 228
Naja haje var. melanoleuca — BEDRIAGA 1982: 299, 440
Naja melanoleuca — SCHÄTTI & LOUMONT 1992: 31
Naja melanoleuca — NILL 1993: 71;
Naja melanoleuca — HOFER 2002: 78, 89 
DistributionSão Tomé Island

Type locality: vicinity of Praia Inhame (0.028636° N, 6.523203° E, WGS-84; 17 m above sea level), São Tomé Island, Republic of São Tomé e Príncipe  
Reproduction 
TypesHolotype: MUHNAC/MB03–001065 (Fig. 4), adult male collected, by Luis Ceríaco, Mariana Marques and Ana Carolina Sousa on 24 February 2016. The specimen was chopped in half by a local man with a machete, and collected minutes after this event. The choice of this particular specimen as holotype is justified as it is the source of the comparative molecular data.
Paratypes. All specimens from the Island of São Tomé, Republic of São Tomé and Príncipe. Eight specimens: IICT 18-1972, adult male collected in Ribeira Peixe (0.090278o N, 6.615278o E, this and all following coordinates use map datum WGS-84; 17 m above sea level) by an unknown collector on 24 December 1972; IICT 20-1967, adult female collected in Santa Josefina (0.247882o N, 6.738315o E; 69 m above sea level) by an unknown collector on October 1967; IICT 2-1966, adult female, without specific locality (São Tomé island) by an unknown collector on 20 February 1966; ZMH R10526, unsexed adult, without specific locality (São Tomé island) collected by Carl Weiss in the late 1840s; ZMH R10527, unsexed adult, without specific locality (São Tomé island) collected by Richard Greef in 1879/1880; MHNG 2518.41 and MHNG 2518.42, both unsexed adults collected in Rio Angra Toldo (0.156881 N, 6.668792o E; 75 m above sea level) by Tilman Nill in 3 April 1991; MHNG 2462.43, adult male collected in the environs of Neves (0.353192 N, 6.634972o E; 281 m above sea level) by Heft and Fahr on 8 August 1989. 
DiagnosisDiagnosis. Naja peroescobari is placed in the subgenus Boulengerina (as defined by Wallach et al. 2009) on the basis of having 2–4 maxillary teeth, the penultimate supralabial height reaching eye level the combination of one preocular and one anterior temporal, rostral much broader than deep, internasals shorter than the prefrontals, dorsal scales smooth and fangs not modified for spitting. Morphologically, N. peroescobari differs from other members of the subgenus Boulengerina by a combination of coloration and scalation patterns. It differs from N. (Boulengerina) chrysti in having 19 dorsal scale rows (17 in chrysti), three infralabials contacting the chinshields (four in chrysti), and by its homogeneous dark coloration (brownish in chrysti). Naja (Boulengerina) chrysti is considered to be strictly aquatic (Chippaux 2006), whereas N. peroescobari is mainly terrestrial. It also differs from N. (Boulengerina) annulata by the lower number of dorsal scales (19 in peroescobari versus 21 to 25 in annulata), and by its coloration, as annulata has between 21–23 dark bands (the first 3–5 simple, subsequent bands double) over a yellow or brown dorsum (contrasting with the homogeneous black of peroescobari), and the ventral coloration (uniformly dark brown in annulata, versus white banded throat, followed by homogeneous black of peroescobari). Naja peroescobari differs from N. (Boulengerina) multifasciata in its size (260 cm maximum in peroescobari versus maximum total length reported for multifasciata 76 cm; Chippaux 2006), by having two postoculars (three in multifasciata), in having 19 dorsal scale rows (15–17 in multifasciata), a higher number of subcaudals (30 to 39 in multifasciata versus 52 to 70 in peroescobari), a higher number of ventral (153 to 175 in multifasciata versus 208 to 215 in peroescobari), and by its coloration (multifasciata is uniformly light yellow on the ventral side, and the dorsal scales are yellow in the anterior part and black in the posterior, with a dark head, separated by the rest of the body by a white collar; Chippaux 2006).
Naja peroescobari differs from the more closely related West African N. melanoleuca, with which it was formerly confused, having a combination of white and black bands, but the white bands are always confined to the first 22 ventral scales, whereas in N. melanoleuca these white bands extend throughout the first two thirds of the venter, usually between the 80 to the 100 scale (max. 125 scale, min. 50 scale; in the type series the last white ventral scale of syntype ANSP 6878 corresponds to the 81st scale, the other having it on the 83th (ANSP6876), 86 th
(ANSP6875) and 90th (ANSP6879).This is even more evident when comparing the new species with N. subvulva, which has whitish scales mottled with dark spots extending all across the venter. Also the dorsal coloration of N. peroescobari is uniformly shiny black, never exhibiting the whithish/yellow markings of the dorsal neck area common in N. melanoleuca. In terms of scalation, the most striking difference between N. peroescobari and N. melanoleuca is the contact of the posterior sublinguals. In N. peroescobari the posterior chin shields are mostly never in contact due to the intrusion of a large scale between them, whereas in N. melanoleuca these are always in contact, either fully across their extent or at least in the anterior part (Meirte 1992 considered this character [sous- linguales postérieures se touchent] as the way to differentiate N. melanoleuca in the key to African snakes). These differences were already mentioned by Capocaccia (1961) and are here confirmed with a larger sample of both taxa.

Diagnosis. In the original description, Naja peroescobari was diagnosed from the other members of the N. melanoleuca group through a lack of white ventrals posterior to ventral 22, a lack of lighter markings on the dorsum, and the separation of the posterior chin shields. Our sample does not fully support the diagnostic value of these characters: at least one specimen (BMNH 1906.3.30.80) has the posterior chin shields in contact, and discrete dark bands separated by lighter bands (although often suffused with dark brown pigment) extend as far back as ventrals 45 and 55, respectively, in BMNH 1906.3.30.80 and MBL 1954. Naja peroescobari is distinct from N. subfulva in never having a brown forebody or a light posterior venter and in lacking dark speckling or spotting on the forebody. It displays greatly reduced ventral banding compared to N. melanoleuca and N. savannula, and, unlike N. guineensis, never has 17 midbody dorsal scale rows.
The species is also diagnosable through unique mitochondrial haplotypes (Ceríaco et al., 2017; cyt b: GenBank MH337634; ND4: MH337440) and a unique PRLR haplotype (MH337499) [Wüster et al. 2018: 86] 
CommentSynonymy: after Ceriaco et al. 2017 (who do not list all references cited in the synonymy). 
EtymologyThe species is named after Pêro (also known as Pedro) Escobar (?–?), one of the 15 -century Portuguese navigators who discovered São Tomé Island on December 21, 1471, Annobon Island on January 1, 1472, and Príncipe island on January 17, 1472, and participated in the discovery of the ocean route from Portugal to India under the command of Vasco da Gama (circa 1460–1524) in 1497, as well as in the expedition of Pedro Álvares Cabral’s (circa 1467–circa 1520) that “discovered” Brazil in 1500.  
References
  • Bocage, J. V. Barboza du. 1886. Reptis e amphibios de S. Thomé. Jornal de Sciencias Mathematicas, Physicas e Naturaes, Lisboa, 11:65-70.
  • Bocage, J.V.B. 1879. Subsidios para a Fauna das possessões portuguezas d’Africa occidental. Jornal das Sciencias Mathematicas, Physicas e Naturaes, 7 (26), 85–96
  • Bocage, J.V.B. 1889. Breves considerações sobre a fauna de S. Thomé. Jornal das Sciencias Mathematicas, Physicas e Naturaes. Segunda Série, 1 (1), 33–36
  • Bocage, J.V.B. 1895. A doninha de São Tomé. Jornal das Sciencias Mathematicas, Physicas e Naturaes, Segunda Série, 4 (13), 24–27
  • Bocage, J.V.B. du 1905. Contribution à la faune des quatre îles du Golfe de Guinée, IV- Île de St. Thomé. Jorn. Acad. Sci. Lisboa, VII (2): 65-96
  • Bocage,J.V. du B. 1866. Lista dos reptis das possessões portuguezas d'Africa occidental que existem no Museu Lisboa. Jorn. Sci. Math. Phys. Nat. Lisboa 1: 37-56
  • Boulenger, G.A. 1906. Report on the reptiles collected by the late L. Fea in West Africa. Ann. Mus. civ. Stor. Nat. Genova (3) 2: 196-216 - get paper here
  • Capocaccia, L. 1961. Contributo allo studio dei serpenti delle isole del Golfo di Guinea. Ann. Mus. civ. Stor. nat. Genova, 72: 285-309. - get paper here
  • CERÍACO, LUIS M. P.; MARIANA P. MARQUES, ANDREAS SCHMITZ, AARON M. BAUER 2017. The “Cobra-preta” of São Tomé Island, Gulf of Guinea, is a new species of Naja Laurenti, 1768 (Squamata: Elapidae) Zootaxa 4324 (1): 121–141 - get paper here
  • Ferreira, J.B. 1902. Lista dos Reptis e Batrachios da colecção do Sr. Newton. Jornal das Sciencias Mathematicas, Physicas e Naturaes, Segunda Série, 6, 281–284
  • Greef, R. 1885. Die Fauna der Guinea-Inseln S. Thomé und Rolas. Sitzungsberichte der Gesellschaft zur Beförderung der Gesammten Naturwissenschaften zu Marburg, (2) :41—79 [1884]
  • Hofer, D. 2002. The São Tomé and Principé Handbook. D. Hofer Verlag, Bern, 152 pp.
  • Manacas, S. 1958. Anfíbios e Répteis das ilhas de São Tomé e do Príncipe e do Ilhéo das Rolas. Conferencia Intern. dos Africanistas Ocidentais, Comunicacoes Zool. e Biol. Animal, 6. Sess., 4: 179-192.
  • Manaças, S. 1973. Alguns dos anfíbios e répteis da província de S. Tomé e Príncipe. In: Livro de Homenagem ao Prof. Fernando Frade. Junta Investigação do Ultramar, Lisboa, pp. 219–230
  • Nill, T. 1993. Die Schlangen der Insel São Tomé (Golf von Guinea) (Reptilia: Serpentes). Faunistische Abhandlungen Staatliches Museum für Tierkunde Dresden, 19 (7), 71–73
  • Schatti, B., & Loumont, C. 1992. [A contribution to the herpetofauna of Sao Tome (Gulf of Guinea) (Amphibia & Reptilia)]. Zool. Abhandl. Staatl. Mus. für Tierkunde, Dresden, 47: 23-36
  • Themido, A.A. 1941. Répteis e batráquios das colónias portuguesas. Memórias e Estudos do Museu Zoológico da Universidade de Coimbra, 119, 1–31
  • Vieira, L. 1886. Noticia ácerca dos productos zoologicos trazidos da ilha de S. Thomé para o Muzeu Zoologico da Universidade de Coimbra pelo Sr. Adolpho F. Moller, em 1885. O Instituto, 34: 235–241
  • WÜSTER, WOLFGANG; LAURENT CHIRIO, JEAN-FRANÇOIS TRAPE, IVAN INEICH, KATE JACKSON, ELI GREENBAUM, CESAR BARRON, CHIFUNDERA KUSAMBA, ZOLTÁN T. NAGY, RICHARD STOREY, CARA HALL, CATHARINE E. WÜSTER, AXEL BARLOW, DONALD G. BROADLEY 2018. Integration of nuclear and mitochondrial gene sequences and morphology reveals unexpected diversity in the forest cobra (Naja melanoleuca) species complex in Central and West Africa (Serpentes: Elapidae). Zootaxa 4455 (1): 068–098 - get paper here
 
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