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Naja (Boulengerina) melanoleuca Hallowell, 1857 Naia haie var. melanoleuca Hallowell, 1857, Proc. Acad. Nat. Sci. Philadelphia: 61. Type locality: Gabon, syntypes ANSP 6875���76, 6878���79. Naja haje var. leucosticta Fischer, 1885, Jahr. Hamburg. Wiss. Anst. 2: 115, pl. v, fig. 11. Type locality: Cameroon and Ogoou�� River, Gabon, syntypes ZMH 4280, 7048, 7299���7302. Naia melanoleuca; Boulenger, 1896: 376 (part, vars. A [a,b] & B [a,b]) Naja melanoleuca melanoleuca; Laurent, 1956: 290, pl. xxvi, fig. 2. Naja (Boulengerina) melanoleuca ; Wallach et al., 2009. Boulengerina melanoleuca; Wallach et al., 2014: 122. Aspidelaps bocagei Sauvage, 1884: 204 (type locality: Gabon and Majumba; holotype MNHN 1884.0015) has been listed as a synonym of N. melanoleuca (e.g., Broadley, 1983; Wallach et al., 2014; Cer��aco et al., 2017), but is in fact a synonym of Naja annulata, as is evident from Sauvage���s description, which notes approximately 20 dark double bands along the entire body length, and 21 mid-dorsal scale rows (see also Schmidt, 1923). Diagnosis: DOrsum blAck, OftEN with 1���3 sEmidividEd yEllOw crOssbANds ON thE NEck, thE first mAy bE AN OcEllus; vENtEr yEllOw with 4 tO 6 blAck bANds iN thE first 100 vENtrAls, thErEAftEr uNifOrm blAck. Variation: DOrsAl scAlE rOws ON NEck 19���27, At midbOdy 19 (vEry rArEly 17 Or 21); vENtrAls 209���230; subcAudAls 59���74 (TAblE 8). SuprAlAbiAls 7, thE third ANd fOurth ENtEriNg thE Orbit; iNfrAlAbiAls 8, thE first fOur iN cONtAct with thE ANtEriOr subliNguAls, NO cuNEAtE; prEOculAr 1; pOstOculArs 3 (vEry rArEly 2 Or 4); tEmpOrAls 1+2 Or 1+3; NuchAls bOrdEriNg tEmpOrAls 5���9, usuAlly 7. Colouration: HEAd brOwN, thE suprAlAbiAls bArrEd blAck ANd yEllOw, chiN yEllOw. BlAck AbOvE, sOmEtimEs A yEllOw mONOcEllAtE mArkiNg ON thE hOOd, Or 1���3 smAll yEllOw blOtchEs, thE dOrsAl scAlEs mAy bE tippEd with whitE iN juvENilEs (Fig. 8). YEllOw Or whitE bElOw with 4 tO 6 blAck bANds ON thE first 100 vENtrAls, usuAlly uNifOrm blAck thErEAftEr. Largest recorded: 2250+420 = 2670 mm, frOm MONiyA, IbAdAN, NigEriA (ButlEr, 1982: 110). Suggested common name: CENtrAl AfricAN fOrEst cObrA. Distribution: CENtErEd ON thE CONgO BAsiN, wEst tO sOuthwEstErN NigEriA ANd pOssibly sOuthErN BENiN, sOuth tO NOrthErN ANgOlA, NOt ExtENdiNg EAst Of thE AlbErtiNE Rift VAllEy, whErE it is rEplAcEd by N. subfulva. ThE wEstErN rANgE limits ArE pOOrly uNdErstOOd. POpulAtiONs frOm sOuthEAstErN NigEriA ArE clEArly AssigNAblE tO this fOrm. A fEw spEcimENs frOm LAmtA, sOuthErN BENiN (IRD 12.B, IRD 54.B, IRD 60.B), ANd GhANA (MNHN 1983.0663���64; NO furthEr lOcAlity iNfOrmAtiON) AlsO AppEAr tO bE AssigNAblE tO N. melanoleuca., Published as part of W��ster, Wolfgang, Chirio, Laurent, Trape, Jean-Fran��ois, Ineich, Ivan, Jackson, Kate, Greenbaum, Eli, Barron, Cesar, Kusamba, Chifundera, Nagy, Zolt��n T., Storey, Richard, Hall, Cara, W��ster, Catharine E., Barlow, Axel & Broadley, Donald G., 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) in Zootaxa 4455 (1), DOI: 10.11646/zootaxa.4455.1.3, http://zenodo.org/record/1456971, {"references":["Hallowell, E. (1857) Notes of a collection of reptiles from the Gaboon country, West Africa, recently presented to the Academy of Natural Sciences of Philadelphia, by Dr. Henry A. Ford. Proceedings of Academy of Natural Sciences, Philadelphia, 9, 48 - 72.","Fischer, J. G. (1885) Ichthyologische und herpetologische Bemerkungen. V. Herpetologische Bemerkungen. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, 2, 82 - 121.","Boulenger, G. A. (1896) Catalogue of the Snakes in the British Museum (Natural History) III. British Museum (Natural History), London, 727 pp. https: // doi. org / 10.5962 / bhl. title. 54273","Laurent, R. F. (1956) Contribution a l'herpetologie de la region des Grands Lacs de l'Afrique centrale. I. Generalites. II. Cheloniens. III. Ophidiens. Annales du Musee royal du Congo Belge, 48, 1 - 390.","Wallach, V., Wuster, W. & Broadley, D. G. (2009) In praise of subgenera: taxonomic status of cobras of the genus Naja Laurenti (Serpentes: Elapidae). Zootaxa, 2236, 26 - 36.","Wallach, V., Williams, K. L. & Boundy, J. (2014) Snakes of the World. A Catalogue of Living and Extinct Species. CRC Press, Boca Raton, 1227 pp. https: // doi. org / 10.1201 / b 16901","Sauvage, H. E. (1884) Notice sur une collection de reptiles et de poissons recueillie a Majumba, Congo. Bulletin de la Societe Zoologique de France, 9, 199 - 204.","Broadley, D. G. (1983) FitzSimons' Snakes of Southern Africa. Delta Books, Johannesburg, 376 pp.","Ceriaco, L. M. P., Marques, M. P., Schmitz, A. & Bauer, A. M. (2017) The \" Cobra-preta \" of Sao Tome Island, Gulf of Guinea, is a new species of Naja Laurenti, 1768 (Squamata: Elapidae). Zootaxa, 4324 (1), 121 - 141. https: // doi. org / 10.11646 / zootaxa. 4324.1","Schmidt, K. P. (1923) Contributions to the herpetology of the Belgian Congo based on the collection of the American Museum Congo Expedition, 1909 - 1915. Part II. Snakes. Bulletin of the American Museum of Natural History, 49, 1 - 146.","Butler, J. A. (1982) Capture au Nigeria d'un Naja noir et blanc de grande taille et notes sur les autres Najas de l'Ouest Africain. Notes Africaines, 176, 110 - 111."]}

Naja (Boulengerina) savannula sp. nov. Broadley, Trape, Chirio & W��ster Naia melanoleuca (not Hallowell) Boulenger, 1896: 376 (part, var. D). Naja ���banded form��� Hughes, 2013: 128. Naja sp. 1 cf. melanoleuca Hallowell, 1857 (yellow banded dorsum) Trape & Bald��, 2014:318. Naja sp. 1 cf. melanoleuca (banded savanna form) Trape & Bald��, 2014: 336. Holotype: MNHN 2018.0002, an adult male collected by Laurent Chirio on 31/03/ 2007 in Ni��ni��, W Biosphere Reserve, Benin, 11.05920 ��N, 2.20488 ��E, elevation 272 m (Fig. 7). Paratype: BMNH 1975.657, a male specimen collected on the grounds of Amadou Bello University Hospital, Zaria, Nigeria, by D. A. Warrell. Diagnosis: DistiNguishAblE frOm OthEr spEciEs Of thE N. melanoleuca cOmplEx by thE prEsENcE Of 3 tO 8 sEmidividEd yEllOwish Or whitish bANds ON thE ANtEriOr dOrsAl fOrEbOdy, bEcOmiNg uNifOrm blAck cAudAd; vENtEr yEllOw with 2 tO 8 blAck bANds. GENErAlly highEr mEAN subcAudAl scAlE cOuNts thAN thE OthEr spEciEs. GENEticAlly diAgNOsAblE thrOugh pOssEssiON Of uNiquE mitOchONdriAl hAplOtypEs (cytOchrOmE b: GENBANk MH337597 ���602; ND4: MH337403 ���408) ANd uNiquE PRLR ANd UBN1 hAplOtypEs (PRLR: MH337501 ���504; UBN1: MH337532 ��� 535). Description of holotype. DimENsiONs: A mAlE spEcimEN, sNOut���vENt lENgth 1815 mm, tAil lENgth 376 mm, tOtAl lENgth 2191 mm, rAtiO tOtAl lENgth: tAil lENgth 5.8. BOdy scAlAtiON: 25 scAlE rOws ArOuNd hOOd, 19 ArOuNd midbOdy, 15 ONE hEAd lENgth AhEAd Of thE vENt, All smOOth ANd ObliquE. DOrsAl scAlEs smOOth, ObliquE. VErtEbrAl rOw NOt ENlArgEd. 213 vENtrAls (ANd ONE iNcOmplEtE vENtrAl bEfOrE thE ANAl), 69 subcAudAls, All dividEd, ANAl siNglE. HEAd scAlAtiON: 7/7 suprAlAbiAls, 3rd & 4th cONtAct Orbit, 6th lArgEst; 8/8 iNfrAlAbiAls, first 4 cONtAct ANtEriOr chiN shiElds; pOstEriOr chiN shiElds iN NArrOw cONtAct At thEir ANtEriOr ENd, sEpArAtEd by mEdiAN scAlE AlONg mOst Of thEir lENgth; 1/1 prEOculAr, twicE As lONg As high; 3/3 pOstOculArs; 1/1 ANtEriOr tEmpOrAl; 3/3 pOstEriOr tEmpOrAls; rOstrAl brOAdEr thAN high, clEArly visiblE frOm AbOvE; 7 tEmpOrAls ANd NuchAls cONtActiNg thE pAriEtAls. PAttErN: uppEr sidE Of hEAd brOwN, lips ANd vENtrAl sidE Of thE hEAd pAlE crEAmy yEllOwish with EAch lAbiAl scAlE lAtErAlly bOrdErEd with blAck. NEck brOwN, dOrsum bEcOmiNg grAduAlly blAck tOwArds rEAr. TwO vEry distiNct pAlE dOrsAl crOssbANds At thE lEvEl Of vENtrAls (V) 24 tO 28, twO OthEr At lEvEl Of V 40 ��� 45, thrEE At lEvEl Of V 58 ��� 64, thrEE lEss distiNct At lEvEl Of V 77 ��� 83, ANd thrEE slightly mArkEd At lEvEl Of V 98 ��� 104. VENtrAls 1 tO 11, 13���19, 23��� 31, 38���47 ANd 57���66 pAlE crEAmy yEllOwish, V 20���22 ANd 32 ��� 37 blAck; V 12 ANd 48���56 yEllOwish spOttEd with blAck spOts. FrOm vENtrAl 67 tO thE vENt, vENtrAls spOttEd with blAck grAduAlly bEcOmE cOmplEtEly blAck. SubcAudAls blAck. Variation: DOrsAl scAlE rOws ON NEck 19���25, At midbOdy 19, bEfOrE vENt 12���15; vENtrAls 211���233, subcAudAls 63���77 (TAblE 8). DOrsAl sEmi-dividEd yEllOw bANds 3���8; vENtrAl priNcipAl blAck bANds 2���8 (Fig. 7). Etymology: ThE NAmE is dErivEd frOm thE cONtrActiON Of its sAvANNA hAbitAt ANd ANNulAtEd cOlOur pAttErN ANd wAs cOiNEd by BArry HughEs iN AN uNpublishEd 1968 mANuscript. WE hAvE rEtAiNEd this NAmE At thE rEquEst Of Our cOllEAguE BArry HughEs. Largest recorded: 1825+405 = 2230 mm, frOm MEdiNA DjikOyE, SENEgAl (IRD 6155.S). Suggested common name: WEst AfricAN bANdEd cObrA. Distribution: SENEgAl ANd GAmbiA EAst tO NOrthErN CAmErOON (Fig. 6). Naja savannula AppEArs tO bE rEstrictEd tO gAllEry fOrEst ArEAs iN AfricA iN GuiNEAN FOrEst/SAvANNA MOsAic, ExtENdiNg NOrthwArds iNtO WEst SudANiAN SAvANNA (ChiriO, 2003, 2013; MONAstEriO et al., 2016). Our rEcOrds ArE mOstly frOm lAtitudEs 10���14��N, ExcEpt iN thE DAhOmEy GAp iN EAstErN GhANA ANd BENiN, whErE thE spEciEs ApprOAchEs thE Gulf Of GuiNEA COAst. ThE EAstErN ExtENt Of thE rANgE is pOOrly uNdErstOOd. A spEcimEN frOm MArgui WANdAlA district, NOrthErN CAmErOON (ApprOx. 10.5��N 13.6��E; MNHN 1962.0022) AppEArs tO bE AssigNAblE tO this spEciEs, ANd ONE Of us (JFT) rEcENtly cOllEctEd A spEcimEN frOm MbOurA, 20 km SW BA��bOkOum, LOgONE OriENtAl PrOviNcE, ChAd (7.598��N, 15.596��E; IRD 2281.N), ANd thErE is A rEcENt rEcOrd frOm NEAr NiAmEy, NigEr (LC, uNpublishEd dAtA). It sEEms likEly thAt thE spEciEs hAs A widEr distributiON iN NOrthErN CAmErOON, ExtrEmE sOuthErN ChAd ANd pOssibly EvEN ExtrEmE NOrthwEstErN CENtrAl AfricAN REpublic (CAR)., Published as part of W��ster, Wolfgang, Chirio, Laurent, Trape, Jean-Fran��ois, Ineich, Ivan, Jackson, Kate, Greenbaum, Eli, Barron, Cesar, Kusamba, Chifundera, Nagy, Zolt��n T., Storey, Richard, Hall, Cara, W��ster, Catharine E., Barlow, Axel & Broadley, Donald G., 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) in Zootaxa 4455 (1), DOI: 10.11646/zootaxa.4455.1.3, http://zenodo.org/record/1456971, {"references":["Boulenger, G. A. (1896) Catalogue of the Snakes in the British Museum (Natural History) III. British Museum (Natural History), London, 727 pp. https: // doi. org / 10.5962 / bhl. title. 54273","Hughes, B. (2013) Snakes of Benin, West Africa. Bulletin de la Societe Herpetologique de France, 144, 101 - 159.","Hallowell, E. (1857) Notes of a collection of reptiles from the Gaboon country, West Africa, recently presented to the Academy of Natural Sciences of Philadelphia, by Dr. Henry A. Ford. Proceedings of Academy of Natural Sciences, Philadelphia, 9, 48 - 72.","Trape, J. - F. & Balde, C. (2014) A checklist of the snake fauna of Guinea, with taxonomic changes in the genera Philothamnus and Dipsadoboa (Colubridae) and a comparison with the snake fauna of some other West African countries. Zootaxa, 3900 (3), 301 - 338. https: // doi. org / 10.11646 / zootaxa. 3900.3.1","Chirio, L. (2003) Inventaire des reptiles de la region de la Reserve de Biosphere Transfrontaliere du W (Niger / Benin / Burkina Faso: Afrique de l'Ouest). Bulletin de la Societe Herpetologique de France, 132, 13 - 41.","Chirio, L. (2013) Inventaire des reptiles de la region de Sangaredi (Guinee maritime). Bulletin de la Societe Herpetologique de France, 144, 67 - 100."]}

Naja (Boulengerina) subfulva Laurent, 1955 Naia melanoleuca (not Hallowell); Boulenger, 1896: 376 (part, vars A [c] & E). Naja melanoleuca subfulva Laurent, 1955, Revue Zool.��� Bot. Afr. 51: 132. Type locality: Lwiro, 1850 m, Kabare, Kivu, Belgian Congo [= Democratic Republic of Congo], holotype MRAC 17514. Naja melanoleuca aurata Stucki-Stirn, 1979, Snake Report 721: 617���620. Type locality: Bamenda, Cameroon. No type designated. Naja subfulva; Chirio & Ineich, 2006: 54. Naja melanoleuca subfulva; Chirio & LeBreton, 2007: 584. Naja (Boulengerina) melanoleuca subfulva ; Broadley & Blaylock, 2013: 139. Naja (Boulengerina) melanoleuca; Conradie et al., 2016: 28. Diagnosis. MidbOdy scAlE rOws 19, ExcEpt AlONg cOAstAl rEgiONs Of EAst AfricA (KENyA, TANzANiA), whErE mOst spEcimENs hAvE 17 rOws. PAttErN highly vAriAblE. Adults Of mOst pOpulAtiONs distiNguishAblE iN hAviNg A brOwN fOrEbOdy, OftEN with spOts, gENErAlly bEcOmiNg dArkEr Or blAckish pOstEriOrly. LAbiAl pAttErN mAy bE AttENuAtEd iN mANy Adults. VENtEr with sEvErAl blAck, dArk brOwN Or grEyish crOssbANds ON thE first 50 vENtrAls, grAduAlly bEcOmiNg uNifOrm blAck cAudAd iN sOmE pOpulAtiONs, but OftEN rEmAiNiNg ENtirEly light, OftEN with ExtENsivE dArkEr spOttiNg Or spEckliNg. WhErE prEsENt, thE light fOrEbOdy ANd/Or light pOstEriOr vENtEr ArE diAgNOstic fOr this spEciEs. GENErAlly fEwEr vENtrAl bANds ANd vENtrAl scAlEs thAN N. melanoleuca Or N. savannula ANd fEwEr subcAudAls thAN N. savannula (TAblE 8). GENEticAlly diAgNOsAblE thrOugh pOssEssiON Of uNiquE mitOchONdriAl hAplOtypEs (cyt b: GENBANk MH337603 ���633; ND4: MH337409 ���439) ANd uNiquE PRLR ANd UBN1 hAplOtypEs (PRLR: MH337441 ���471; UBN1: MH337531, MH337536 ���562, MH337564 ���566). Variation. DOrsAl scAlE rOws ON NEck 19���27, At midbOdy 19 (vEry rArEly 17 Or 21); vENtrAls 196���226; ANAl ENtirE; subcAudAls 55���71 (TAblE 8). SuprAlAbiAls 7 (vEry rArEly 5 Or 6), thE third ANd fOurth ENtEriNg Orbit; iNfrAlAbiAls 8 (rArEly 7), thE first fOur (rArEly thrEE) iN cONtAct with thE ANtEriOr subliNguAls, NO cuNEAtE (vEry rArEly ONE); prEOculAr 1; pOstOculArs 3; tEmpOrAl 1+2 Or 1+3; tEmpOrAl bOrdEriNg pAriEtAls 5���9, usuAlly 7. Colouration. HEAd brOwN, thE suprAlAbiAls usuAlly bArrEd blAck ANd yEllOw, chiN yEllOw. DOrsum uNifOrm blAck, Or with fAiNt whitE crOssbArs Or whitE-tippEd dOrsAl scAlEs, rEsultiNg iN A stippliNg EffEct, iN All juvENilEs. Adults frOm thE pEriphEry Of LAkE VictOriA ANd pArts Of thE CONgO BAsiN (Fig. 9) hAvE A uNifOrmly blAck dOrsum As Adults, whErEAs iN mOst Of thE rANgE, Adults bEcOmE brOwN ANtEriOrly. ThErE mAy bE 2 Or 3 fAiNt yEllOw dividEd crOssbArs ON thE NEck. VENtrAl bANdiNg OftEN fAiNt iN Adults, with grEy Or pAlE brOwN vENtrAl bANds thAt mAy bE vEry iNcONspicuOus, Or lAckiNg AltOgEthEr. NArrOw AccEssOry vENtrAl bANds usuAlly AbsENt. Largest recorded. 2016+380 = 2396 mm (tAil truNcAtEd) [JPT 1856 ��� ZiNAvE, MOzAmbiquE], but NMZB-UM 17594 frOm INyANgANi TEA EstAtEs, ZimbAbwE, hAd A tOtAl lENgth Of 2690 mm (ONly hEAd ANd NEck prEsErvEd). Suggested common name. BrOwN fOrEst cObrA. This suggEstiON rEflEcts thE fAct thAt iN mOst pOpulAtiONs, with thE ExcEptiON Of thE surrOuNdiNgs Of LAkE VictOriA ANd sOmE lOcAtiONs iN thE CONgO BAsiN, thE ANtEriOr bOdy Of Adults is brOwN rAthEr thAN blAck. WE NOtE thAt thE distributiON Of this spEciEs iNcludEs mANy NON-fOrEstEd hAbitAts, iNcludiNg sAvANNAs ANd OpEN wEtlANds (SpAwls et al., 2018). HOwEvEr, rEtENtiON Of thE wOrd ���fOrEst��� iN thE cOmmON NAmE cONvEys thE phylOgENEtic iNfOrmAtiON thAt this spEciEs is pArt Of thE Naja melanoleuca cOmplEx. Distribution. FOrEst/sAvANNA mOsAic, ENcircliNg thE CONgO BAsiN, frOm thE grAssfiElds Of wEstErN CAmErOON NOrthwArd, rEAchiNg 13.45��N At BOl, ChAd, ON thE NOrthErN shOrE Of LAkE ChAd (IRD 2802.N), EAst thrOugh thE CAR ANd SOuth SudAN tO wEstErN EthiOpiA, sOuth thrOugh UgANdA, wEstErN KENyA, EAstErN DRC, RwANdA, BuruNdi ANd wEstErN TANzANiA tO NOrthErN MAlAwi, wEst thrOugh ZAmbiA ANd KAtANgA tO ANgOlA ANd thE LOwEr CONgO rEgiON (Fig. 6). ThE distributiON ExtENds disjuNctly EAst ANd sOuth tO thE EAst AfricAN cOAst frOm KENyA tO NOrthErN KwAZulu-NAtAl (SOuth AfricA), iNcludiNg iNlANd lOcAtiONs iN wEstErN ZimbAbwE. ThE prEcisE distributiON Of this tAxON iN thE CONgO rEgiON rEmAiNs tO bE AscErtAiNEd: wE hAvE hErE prOvidEd mOlEculAr EvidENcE Of multiplE spEcimENs Of this spEciEs frOm thE EvErgrEEN fOrEst zONE Of NOrthErN REpublic Of CONgO (LikOuAlA FOrEst ��� JAcksON et al., 2007) ANd thE NOrthEAstErN DRC (NOrthwEstErN OriENtAlE PrOviNcE AlONg thE CONgO RivEr). SOmE Of thEsE spEcimENs ANd pOpulAtiONs ArE difficult tO diffErENtiAtE frOm sympAtric N. melanoleuca, NEcEssitAtiNg thE usE Of DNA bArcOdiNg ApprOAchEs (HEbErt et al., 2003; ClAusE et al., 2016) tO ENsurE rEliAblE idENtificAtiON. TwO spEcimENs frOm sOuthEAstErN NigEriA with lOw vENtrAl scAlE cOuNts ANd lAckiNg vENtrAl bANdiNg Or displAyiNg A siNglE fAdEd bANd mAy AlsO bE AssigNAblE tO this spEciEs (DEgEmA, RivErs StAtE: BMNH 1902.11.10.9, ANd ���Oil RivEr ���: BMNH 88.8.29.20), suggEstiNg A widEr distributiON iN EAstErN NigEriA. Comment. This spEciEs displAys cONsidErAblE gEOgrAphic vAriAtiON iN pAttErN ANd scAlAtiON, As wEll As iNdicAtiONs Of phylOgEOgrAphic structurE. ThE prEsENt study fOcusEd ON thE rElAtiONships AmONg thE fivE mitOchONdriAlly dEfiNEd cANdidAtE spEciEs, ANd did NOt ENcOmpAss thE full rANgE Of mOrphOlOgicAl vAriAtiON iN N. subfulva. It is pOssiblE thAt futurE wOrk mAy idENtify AdditiONAl cryptic divErsity withiN this widE-rANgiNg ANd vAriAblE spEciEs., Published as part of W��ster, Wolfgang, Chirio, Laurent, Trape, Jean-Fran��ois, Ineich, Ivan, Jackson, Kate, Greenbaum, Eli, Barron, Cesar, Kusamba, Chifundera, Nagy, Zolt��n T., Storey, Richard, Hall, Cara, W��ster, Catharine E., Barlow, Axel & Broadley, Donald G., 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) in Zootaxa 4455 (1), DOI: 10.11646/zootaxa.4455.1.3, http://zenodo.org/record/1456971, {"references":["Laurent, R. F. (1955) Diagnoses preliminaires de quelques serpents venimeux. Revue de Zoologie et Botanique Africaine, 51, 127 - 139.","Boulenger, G. A. (1896) Catalogue of the Snakes in the British Museum (Natural History) III. British Museum (Natural History), London, 727 pp. https: // doi. org / 10.5962 / bhl. title. 54273","Stucki-Stirn, M. C. (1979) Snake Report 721. Herpeto-Verlag, Teuffenthal, 650 pp.","Chirio, L. & Ineich, I. (2006) Biogeography of the reptiles of the Central African Republic. African Journal of Herpetology, 55, 23 - 59. https: // doi. org / 10.1080 / 21564574.2006.9635538","Chirio, L. & LeBreton, M. (2007) Atlas des reptiles du Cameroun. Museum National d'Histoire Naturelle, Paris, 686 pp.","Broadley, D. G. & Blaylock, R. (2013) The Snakes of Zimbabwe and Botswana. Edition Chimaira, Frankfurt am Main, 387 pp.","Conradie, W., Bills, R. & Branch, W. R. (2016) The herpetofauna of the Cubango, Cuito, and lower Cuando river catchments of south-eastern Angola. Amphibian & Reptile Conservation, 10, 6 - 36."]}

Naja (Boulengerina) guineensis sp. nov. Broadley, Trape, Chirio, Ineich & W��ster Naia melanoleuca (not Hallowell) Boulenger, 1896: 376 (part, var. B [c,d], C). Naja sp. 2 cf. melanoleuca Hallowell, 1857 (blackish dorsum) Trape & Bald��, 2014: 318. Naja sp. 2 cf. melanoleuca (forest form). Trape & Bald��, 2014: 336. Holotype: MNHN 1921.0485, a male from N���Z��b��la, Macenta Prefecture, Nz��r��kor�� region of forested southeastern Guinea (8�� 05���N, 9�� 05���W), elev. 490 m, Coll. Paul Chabanaud (1876���1959) between 1919���1920 (Chabanaud, 1921: 471) (Fig. 5). Paratype: BMNH 1960.1.3.72, a male from Njala, Kori, Sierra Leone, Coll. C.T. Pyne. Diagnosis. Naja guineensis cAN bE distiNguishEd frOm thE pArtly sympAtric N. savannula sp. NOv. by lAckiNg ExtENdEd dOrsAl bANdiNg, OftEN hAviNg 17 rAthEr thAN 19 dOrsAl scAlE rOws At midbOdy, A gENErAlly lOwEr subcAudAl scAlE cOuNt, fEwEr vENtrAl bANds, A lEssEr pOstEriOr ExtENt Of thE vENtrAl bANdiNg, ANd A strONg tENdENcy tOwArds mElANism iN Adults. SpEcimENs with 19 midbOdy dOrsAl scAlE rOws cAN bE distiNguishEd frOm N. melanoleuca thrOugh thE rEducEd NumbEr Of vENtrAl bANds, lEssEr pOstEriOr ExtENt Of bANdiNg ANd tENdENcy Of ONtOgENEtic mElANism frOm N. subfulva iN lAckiNg A lightEr ANtEriOr dOrsum ANd thrOugh ONtOgENEtic mElANism, ANd frOm N. peroescobari iN hAviNg thE pOstEriOr chiN shiElds iN cONtAct. Description of holotype. DimENsiONs: A mAlE spEcimEN with dissEctEd tAil bAsE, sNOut���vENt lENgth 1850 mm, tAil lENgth 370 mm, tOtAl lENgth 2220 mm, rAtiO tOtAl lENgth: tAil lENgth 6:1. BOdy scAlAtiON: 23 scAlE rOws ArOuNd hOOd, 17 ArOuNd midbOdy, 13 ONE hEAd lENgth AhEAd Of thE vENt, All smOOth ANd ObliquE; 208 vENtrAls, 62 subcAudAls, All dividEd, ANAl siNglE. HEAd scAlAtiON: 7/7 suprAlAbiAls, 3rd & 4th cONtAct Orbit, 6th lArgEst; 8/8 iNfrAlAbiAls, first 4 cONtAct ANtEriOr chiN shiElds; ANtEriOr pAir Of chiN shiElds ANd iNfrAlAbiAls 3���4 with AN ANOmAly ON bOth sidEs, thEy ArE wArty ANd cOvErEd with circumvOlutiONs; 1/1 prEOculAr, twicE As lONg As high; 3/3 pOstOculArs; 1/1 ANtEriOr tEmpOrAl; pOstEriOr tEmpOrAls 3/3; rOstrAl brOAdEr thAN high ANd slightly dAmAgEd ANtEriOrly, clEArly visiblE frOm AbOvE; 7 tEmpOrAls ANd NuchAls cONtActiNg bOth pAriEtAls. PAttErN: uppEr sidE Of hEAd ANd uppEr tEmpOrAl rEgiON dArk brOwN iNcludiNg pAriEtAls ANd tEmpOrAls but blAck bEhiNd thEm, lips ANd vENtrAl sidE Of hEAd crEAmy yEllOwish withOut dArk mOttliNg ON chiN ANd ANtEriOr thrOAt; EAch pOstEriOr suprAlAbiAl EdgE brOAdly OutliNEd iN blAck frOm EyE lEvEl tO mOuth cOrNEr; tEmpOrAls uNifOrmly brOwNish. DOrsum uNifOrm brOwNish-blAck thrOughOut. VENtEr: ANtEriOr first 32 cm (14% Of tOtAl lENgth) Of vENtEr with AltErNAtiNg yEllOw ANd blAckish irrEgulAr bANds, thE pOstEriOr yEllOw bANds iNcrEAsiNgly suffusEd with dArk mOttliNg. ONly vENtrAl 1 is uNifOrm clEAr yEllOw, vENtrAls 2���10, 19, 28���29, 32���35 yEllOw but AlwAys iNcrEAsiNgly suffusEd with dArk mOttliNg, rEmAiNiNg vENtrAls ANd uNdErsidE Of tAil blAck (AftEr vENtrAl 35). Variation. MidbOdy dOrsAl scAlE rOw cOuNts Of 17 ANd 19 ArE ApprOximAtEly EquAlly cOmmON iN this spEciEs. IN lArgE Adults, light pAttErN ElEmENts ON thE hEAd ANd thrOAt OftEN bEcOmE hEAvily suffusEd with blAck pigmENt, lEAdiNg tO A virtuAlly ENtirEly mElANistic sNAkE. SOmE spEcimENs hAvE 1���4 gENErAlly fAiNt Or pOOrly dEfiNEd light bANds AcrOss thE NEck, ANd OccAsiONAlly AN OcEllAtE hOOd mArkiNg. Largest recorded: 1818+437 = 2255 mm, frOm BAllAssOu, GuiNEA (IRD 4213.G), but lArgEr spEcimENs hAvE bEEN ObsErvEd. BAsEd ON lOcAlity, MENziEs��� (1966) rEpOrt Of A spEcimEN mEAsuriNg ���Eight fEEt, Eight iNchEs��� (264 cm) frOm BO, SiErrA LEONE, is likEly tO rEfEr tO this spEciEs. Etymology. ThE spEcific EpithEt guineensis mEANs ���frOm GuiNEA ��� ANd is chOsEN tO rEflEct thE distributiON Of thE spEciEs iN thE UppEr GuiNEA fOrEsts Of WEst AfricA, pArt Of thE WEst AfricAN FOrEsts biOdivErsity hOtspOt (MyErs et al., 2000). Suggested common name. BlAck fOrEst cObrA. Distribution. ThE distributiON Of Naja guineensis AppEArs tO bE rEstrictEd tO thE UppEr GuiNEA FOrEsts Of wEstErN AfricA, frOm wEstErN TOgO tO LibEriA ANd GuiNEA (TrApE & BAld��, 2014) (Fig. 6). ThErE is A siNglE rEcOrd frOm CONtubOEl, GuiNEA BissAu (MBL 535). All OthEr rEcOrds 10��N Or lOwEr., Published as part of W��ster, Wolfgang, Chirio, Laurent, Trape, Jean-Fran��ois, Ineich, Ivan, Jackson, Kate, Greenbaum, Eli, Barron, Cesar, Kusamba, Chifundera, Nagy, Zolt��n T., Storey, Richard, Hall, Cara, W��ster, Catharine E., Barlow, Axel & Broadley, Donald G., 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) in Zootaxa 4455 (1), DOI: 10.11646/zootaxa.4455.1.3, http://zenodo.org/record/1456971, {"references":["Boulenger, G. A. (1896) Catalogue of the Snakes in the British Museum (Natural History) III. British Museum (Natural History), London, 727 pp. https: // doi. org / 10.5962 / bhl. title. 54273","Hallowell, E. (1857) Notes of a collection of reptiles from the Gaboon country, West Africa, recently presented to the Academy of Natural Sciences of Philadelphia, by Dr. Henry A. Ford. Proceedings of Academy of Natural Sciences, Philadelphia, 9, 48 - 72.","Trape, J. - F. & Balde, C. (2014) A checklist of the snake fauna of Guinea, with taxonomic changes in the genera Philothamnus and Dipsadoboa (Colubridae) and a comparison with the snake fauna of some other West African countries. Zootaxa, 3900 (3), 301 - 338. https: // doi. org / 10.11646 / zootaxa. 3900.3.1"]}

Naja (Boulengerina) savannula sp. nov. Broadley, Trape, Chirio & Wüster Naia melanoleuca (not Hallowell) Boulenger, 1896: 376 (part, var. D). Naja “banded form” Hughes, 2013: 128. Naja sp. 1 cf. melanoleuca Hallowell, 1857 (yellow banded dorsum) Trape & Baldé, 2014:318. Naja sp. 1 cf. melanoleuca (banded savanna form) Trape & Baldé, 2014: 336. Holotype: MNHN 2018.0002, an adult male collected by Laurent Chirio on 31/03/ 2007 in Niénié, W Biosphere Reserve, Benin, 11.05920 °N, 2.20488 °E, elevation 272 m (Fig. 7). Paratype: BMNH 1975.657, a male specimen collected on the grounds of Amadou Bello University Hospital, Zaria, Nigeria, by D. A. Warrell. Diagnosis: DistiNguishAblE frOm OthEr spEciEs Of thE N. melanoleuca cOmplEx by thE prEsENcE Of 3 tO 8 sEmidividEd yEllOwish Or whitish bANds ON thE ANtEriOr dOrsAl fOrEbOdy, bEcOmiNg uNifOrm blAck cAudAd; vENtEr yEllOw with 2 tO 8 blAck bANds. GENErAlly highEr mEAN subcAudAl scAlE cOuNts thAN thE OthEr spEciEs. GENEticAlly diAgNOsAblE thrOugh pOssEssiON Of uNiquE mitOchONdriAl hAplOtypEs (cytOchrOmE b: GENBANk MH337597 –602; ND4: MH337403 –408) ANd uNiquE PRLR ANd UBN1 hAplOtypEs (PRLR: MH337501 –504; UBN1: MH337532 – 535). Description of holotype. DimENsiONs: A mAlE spEcimEN, sNOut–vENt lENgth 1815 mm, tAil lENgth 376 mm, tOtAl lENgth 2191 mm, rAtiO tOtAl lENgth: tAil lENgth 5.8. BOdy scAlAtiON: 25 scAlE rOws ArOuNd hOOd, 19 ArOuNd midbOdy, 15 ONE hEAd lENgth AhEAd Of thE vENt, All smOOth ANd ObliquE. DOrsAl scAlEs smOOth, ObliquE. VErtEbrAl rOw NOt ENlArgEd. 213 vENtrAls (ANd ONE iNcOmplEtE vENtrAl bEfOrE thE ANAl), 69 subcAudAls, All dividEd, ANAl siNglE. HEAd scAlAtiON: 7/7 suprAlAbiAls, 3rd & 4th cONtAct Orbit, 6th lArgEst; 8/8 iNfrAlAbiAls, first 4 cONtAct ANtEriOr chiN shiElds; pOstEriOr chiN shiElds iN NArrOw cONtAct At thEir ANtEriOr ENd, sEpArAtEd by mEdiAN scAlE AlONg mOst Of thEir lENgth; 1/1 prEOculAr, twicE As lONg As high; 3/3 pOstOculArs; 1/1 ANtEriOr tEmpOrAl; 3/3 pOstEriOr tEmpOrAls; rOstrAl brOAdEr thAN high, clEArly visiblE frOm AbOvE; 7 tEmpOrAls ANd NuchAls cONtActiNg thE pAriEtAls. PAttErN: uppEr sidE Of hEAd brOwN, lips ANd vENtrAl sidE Of thE hEAd pAlE crEAmy yEllOwish with EAch lAbiAl scAlE lAtErAlly bOrdErEd with blAck. NEck brOwN, dOrsum bEcOmiNg grAduAlly blAck tOwArds rEAr. TwO vEry distiNct pAlE dOrsAl crOssbANds At thE lEvEl Of vENtrAls (V) 24 tO 28, twO OthEr At lEvEl Of V 40 – 45, thrEE At lEvEl Of V 58 – 64, thrEE lEss distiNct At lEvEl Of V 77 – 83, ANd thrEE slightly mArkEd At lEvEl Of V 98 – 104. VENtrAls 1 tO 11, 13–19, 23– 31, 38–47 ANd 57–66 pAlE crEAmy yEllOwish, V 20–22 ANd 32 – 37 blAck; V 12 ANd 48–56 yEllOwish spOttEd with blAck spOts. FrOm vENtrAl 67 tO thE vENt, vENtrAls spOttEd with blAck grAduAlly bEcOmE cOmplEtEly blAck. SubcAudAls blAck. Variation: DOrsAl scAlE rOws ON NEck 19–25, At midbOdy 19, bEfOrE vENt 12–15; vENtrAls 211–233, subcAudAls 63–77 (TAblE 8). DOrsAl sEmi-dividEd yEllOw bANds 3–8; vENtrAl priNcipAl blAck bANds 2–8 (Fig. 7). Etymology: ThE NAmE is dErivEd frOm thE cONtrActiON Of its sAvANNA hAbitAt ANd ANNulAtEd cOlOur pAttErN ANd wAs cOiNEd by BArry HughEs iN AN uNpublishEd 1968 mANuscript. WE hAvE rEtAiNEd this NAmE At thE rEquEst Of Our cOllEAguE BArry HughEs. Largest recorded: 1825+405 = 2230 mm, frOm MEdiNA DjikOyE, SENEgAl (IRD 6155.S). Suggested common name: WEst AfricAN bANdEd cObrA. Distribution: SENEgAl ANd GAmbiA EAst tO NOrthErN CAmErOON (Fig. 6). Naja savannula AppEArs tO bE rEstrictEd tO gAllEry fOrEst ArEAs iN AfricA iN GuiNEAN FOrEst/SAvANNA MOsAic, ExtENdiNg NOrthwArds iNtO WEst SudANiAN SAvANNA (ChiriO, 2003, 2013; MONAstEriO et al., 2016). Our rEcOrds ArE mOstly frOm lAtitudEs 10–14°N, ExcEpt iN thE DAhOmEy GAp iN EAstErN GhANA ANd BENiN, whErE thE spEciEs ApprOAchEs thE Gulf Of GuiNEA COAst. ThE EAstErN ExtENt Of thE rANgE is pOOrly uNdErstOOd. A spEcimEN frOm MArgui WANdAlA district, NOrthErN CAmErOON (ApprOx. 10.5°N 13.6°E; MNHN 1962.0022) AppEArs tO bE AssigNAblE tO this spEciEs, ANd ONE Of us (JFT) rEcENtly cOllEctEd A spEcimEN frOm MbOurA, 20 km SW BAïbOkOum, LOgONE OriENtAl PrOviNcE, ChAd (7.598°N, 15.596°E; IRD 2281.N), ANd thErE is A rEcENt rEcOrd frOm NEAr NiAmEy, NigEr (LC, uNpublishEd dAtA). It sEEms likEly thAt thE spEciEs hAs A widEr distributiON iN NOrthErN CAmErOON, ExtrEmE sOuthErN ChAd ANd pOssibly EvEN ExtrEmE NOrthwEstErN CENtrAl AfricAN REpublic (CAR).

Naja senegalensis sp. nov. Trape, Chirio and Wüster Figs. 4–8 Holotype: MNHN 2008.0074 (previously IRD S- 8549), collected in September 2008 near Dielmo (13 ° 43 ’N, 16 ° 25 ’W) by Mr. Babacar N’Dao, veterinary agent at Keur Lahim Fatim, who sent it to the first author (Fig. 4–6). Paratypes: 31 specimens, all from Sénégal: MNHN 2008.0075 (previously IRD S- 409), MNHN 2008.0076 (previously IRD S- 1027) MNHN 2008.0077 (previously IRD S- 1028), MNHN 2008.0078 (previously IRD S- 1578), MNHN 2008.0079 (previously IRD S- 1589): Keur Lahine Fatim (13 ° 44 ’N, 16 ° 23 ’N), Sine Saloum; MNHN 2008.0080 (previously IRD S- 443), MNHN 2008.0081 (previously IRD S- 2302), MNHN 2008.0082 (previously IRD S- 2306): Keur Bakar Mané (13 ° 37 ’N, 16 ° 17 ’W), Sine Saloum; MNHN 2008.0083 (previously IRD S- 762), MNHN 2008.0084 (previously IRD S- 5283): Keur Seny Gueye (13 ° 36 ’N, 16 ° 19 ’W), Sine Saloum; MNHN 2008.0085 (previously IRD S- 855), MNHN 2008.0086 (previously IRD S- 858), MNHN 2008.0087 (previously IRD S- 1640): Keur Gadji (13 ° 38 ’N, 16 ° 19 ’W), Sine Saloum; MNHN 2008.0088 (previously IRD S- 1283): Keur Santhiou (13 ° 39 ’N, 16 ° 21 ’W), Sine Saloum; IRSNB 2654 (previously IRD S- 605), IRSNB 2655 (previously IRD S- 1429), IRSNB 2656 (previously IRD S- 1435), IRD S- 1439, IRD S- 1440, IRD S- 1442, IRD S- 1461, IRD S- 3429, IRD S- 3430, IRD S- 5692: Dielmo (13 ° 43 ’N, 16 ° 25 ’N), Sine Saloum; IRD S- 2113: Landieni (12 ° 33 ’N, 12 ° 22 ’W), eastern Senegal; IRD S- 5344: Saroudia (12 ° 32 ’N, 11 ° 35 ’W), eastern Senegal; IRD S- 5427: Sambarabougou (13 °06’N, 11 ° 51 ’W), eastern Senegal; IRD S- 5849, IRD S- 5854: Guénoto (13 ° 33 ’N, 13 ° 50 ’W), eastern Senegal; IRD S- 6204: Keur Lamine Diamé (13 ° 37 ’N, 16 ° 16 ’W), Sine Saloum; IRD S- 6461: Touba Baria (13 ° 38 ’N, 16 ° 14 ’W), Sine Saloum. Other specimens examined. 86 specimens: Senegal (46 specimens): IRD S- 343, IRD S- 3431: Senegal; IRD S- 462: Keur Ayip Kâ (13 ° 39 ’N, 16 ° 19 ’W), Sine Saloum; IRD S- 604, IRD S- 606, IRD S- 664: Keur Bakar Mané (13 ° 37 ’N, 16 ° 17 ’W), Sine Saloum; IRD S- 856, IRD S- 1634: Keur Gadji (13 ° 38 ’N, 16 ° 19 ’W), Sine Saloum; IRD S- 1279, S- 1280, S- 1281, S- 1292, S- 1293: Keur Santhiou (13 ° 39 ’N, 16 ° 21 ’W), Sine Saloum; S- 1411, S- 1472, S- 1482: Dielmo (13 ° 43 ’N, 16 ° 25 ’W), Sine Saloum; IRD S- 1588: Keur Lahim Fatim (13 ° 44 ’N, 16 ° 23 ’N), Sine Saloum; IRD S- 3849: Badiara (13 ° 13 ’N, 14 ° 12 ’W), Haute Casamance; IRD S- 3952: Goundaga (12 ° 51 ’N, 14 °05’W), Haute Casamance; IRD S- 4806: Tialé (15 ° 14 ’N, 16 ° 49 ’W), Cayor; IRD S- 5085: Oubadji (12 ° 40 ’N, 13 °03’W), Sénégal oriental; IRD S- 5307: Saroudia (12 ° 32 ’N, 11 ° 35 ’W), Sénégal oriental; IRD S- 5795, IRD S- 6090: Keur Momat Souna (13 ° 38 ’N, 16 ° 17 ’W), Sine Saloum; IRD S- 5851, IRD S- 5853: Guénoto (13 ° 33 ’N, 13 ° 50 ’W), eastern Senegal; IRD S- 5862: Médina Djikoye (13 ° 37 ’N, 16 ° 18 ’W), Sine Saloum; IRD S- 6239: Touba Baria (13 ° 38 ’N, 16 ° 14 ’W), Sine Saloum; IRD S- 6656: Takoudialla (12 ° 50 ’N, 14 °04’N), Haute Casamance; IRD S- 6680: Ségoto (13 ° 18 ’N, 11 ° 49 ’W), Sénégal oriental; IRD S- 7200: Touba Ndiaye (15 °09’N, 16 ° 52 ’W), Cayor; IFAN 55 - 4-13: Cambérène (14 ° 45 'N, 17 ° 25 'W); IFAN 52 - 11-90, IFAN 53 - 11-143: Dakar (14 ° 42 'N, 17 ° 27 'W); IFAN 47 - 1-10, IFAN 47 - 1-15, IFAN 50 - 9-149, IFAN 51 - 12-53, IFAN 52 - 3-23, IFAN 52 - 7-47, IFAN 56 - 5-50: Hann (14 ° 43 'N, 17 ° 26 'W), IFAN 82 - 1-2: Keur Massar (14 ° 47 'N, 17 ° 18 'W); IFAN 52 - 1-8: Malika (14 ° 47 'N, 17 ° 20 'W); IFAN 53 - 3-20: Ouakam (14 ° 43 'N, 17 ° 29 'W); IFAN 44 - 1-3: Popenguine (14 ° 33 'N, 17 °07'W). Mali (32 specimens): IRD 2353 -M, IRD 2354 -M: Ballabougou (12 ° 52 ’N, 06° 52 ’W); IRD 238 -M: Bangaya (13 ° 14 ’N, 10 ° 43 ’W); IRD 1179 -M, IRD 1181 -M: Djinagué (12 ° 59 ’N, 09° 52 ’W); IRD 103 -M: Doussoudiana (11 °09’N, 07° 48 ’W); IRD 2368 -M: Koundian (13 ° 10 ’N, 10 ° 40 ’W); IRD 805 -M: Laminina (11 ° 12 ’N, 07° 47 ’W); IRD 1977 -M, 2003 -M, 2017 -M: Mamoroubougou (11 ° 13 ’N, 05° 28 ’W); IRD 1796 -M, IRD 3419 -M: Npiébougou (11 ° 59 ’N, 08°00’W); IRD 2352 -M: Sadjouroubougou (12 ° 35 ’N, 07° 44 ’W); IRD 1591 -M: Sare-Soma (14 ° 45 ’N, 03° 55 ’W); IRD 878 -M, 957 -M: Sebekourani (12 ° 12 ’N, 08° 42 ’W); IRD 184 -M, IRD 2102 -M, IRD 2109 -M, IRD 2118 -M, IRD 2145 - M, IRD 2186 -M, IRD 3606 -M, IRD 3617 -M, IRD 3618 -M, IRD 3683 -M: Titiéna (11 ° 27 ’N, 06° 33 ’W); IRD 156 -M, IRD 581 -M, IRD 2349 -M, IRD 2350 -M, IRD 2351 -M (13 °09’N, 07° 57 ’W). Niger (2 specimens): IRD 201 -N: Karosofua (13 ° 37 ’N, 06° 37 ’E); IRD 1504 -N: Téla (12 °08’N, 03° 28 ’E). Burkina Faso (2 specimen): LC 6531: Kondio (11 ° 37 ’N, 02°01’E); IFAN 48 - 2-9: Dano près Diébougou (11 °09'N, 03°04'W); USNM 237088 8km S of Dana (NW 1202 C 1). Bénin (1 spécimen): LC 7109: Niénié (11 ° 22 ’N, 02° 12 ’E). Guinée (1 specimen): IFAN 52 - 6-34: Niandan-Banie (approximatively 10 ° 20 'N, 09° 50 'W). Nigeria (1 specimen): CM 92607 Shagunu, west bank of Kainji Lake (10 º 20 ’N, 04º 28 ’E). Diagnosis. Naja senegalensis resembles all other members of the N. haje complex and differs from all other Naja in having a row of subocular scales separating the orbit from the supralabial scales. Naja senegalensis can be distinguished from other species of the N. haje complex through a combination of scale counts and the coloration of juveniles and adults. Comparative scale counts are given in Table 3. Naja senegalensis is distinguishable from N. haje through its higher neck scale row count: N. senegalensis normally has 25 dorsal scale rows around the neck, although some specimens have 23 or 27. By contrast, W. African N. haje have fewer neck scale rows (19–21 in five specimens from Niger, 21–23 in three specimens from Nigeria, 21 in one specimen from Tombouctou, Mali). In other parts of Africa, the majority of specimens also have 21 or fewer scale rows around the neck (Table 3). A cobra specimen from Djibouti, with 27 scale rows around the neck and 23 at midbody, tentatively assigned to the N. haje complex by Ineich (2001), appears to be a spitting cobra. Other scalation characters do not distinguish N. senegalensis from N. haje, although the new species tends to occupy the upper end of the spectrum of ventral scale counts in the complex (Table 3). Another diagnostic feature of N. senegalensis is the juvenile pattern: the great majority of juveniles and subadult specimens have a highly contrasting white blotch on the neck, within the dark collar encircling the neck (Fig. 7). This pale patch is present in 37 out of 39 small and medium-sized specimens, but barely discernible or absent in almost all larger adults. However, one of us (LC) recently photographed a large captive adult (approximately 200 cm total length) from the area of Bamako, Mali, that retained a very conspicuous, heart-shaped nuchal mark (Fig. 8). We have never observed this patch in N. haje, and therefore consider its presence to be a diagnostic character for N. senegalensis. Naja senegalensis differs strongly in coloration from sympatric or parapatric West African N. haje: adults of N. senegalensis are almost invariably uniformly dark brown dorsally, whereas juveniles are greyish dorsally and yellowish ventrally, with a dark collar (Fig. 7) around the neck and usually a white neck blotch. Small adult specimens tend to be dark brown with paler speckles and their ventral side is yellowish. Small adult N. senegalensis from W National Park, in the Niger – Burkina Faso – Benin border region, have a brown dorsal coloration with small reddish dots of one scale each. The entire head, and in particular the supralabial region, are normally uniformly dark brown. In Niger and Nigeria, where both N. senegalensis and N. haje are found (the former in the Sudan savanna, the latter in the Sahelian zone, but with possible areas of sympatry), both adults and juveniles of N. haje display quite different colours: the body of adults is yellow to dark brown dorsally (often mostly yellow, especially in Niger), often with scattered individual dark scales, but the venter is at least partly cream-coloured, or with contrasting light and dark bands or blotches. The sides of the head, and in particular the supralabial region, normally display contrasting areas of pale and dark pigmentation, and, most noticeably, a dark spot under the eye, reminiscent of the “teardrop” marking present in N. nubiae (Wüster & Broadley, 2003), and a more or less distinct dark greyish neck band, approximately ten scales wide. Juveniles lack the pale neck patch present in young N. senegalensis. An adult N. haje from Niger is illustrated in Trape & Mané (2006, p. 195). Elsewhere in Africa, N. haje shows great variation in colour pattern, but differs consistently from that shown in N. senegalensis as follows: - The Moroccan/Western Saharan population extends into the northern parts of Western Sahara, and specimens have been illustrated by Bons & Geniez (1996, p. 251), Geniez et al. (2004, p. 169, 171) and Dobiey & Vogel (2007, p. 67). Adults are usually uniform black except for a yellowish gular area. Some may be dark brown above and grey below. - N. haje extends across northern Algeria south of the Atlas Mountains (Schleich et al., 1996), but no voucher specimens have been examined. LC did not encounter the species during two years that he spent in the region; local people knew it, but reported that it was very rare. Renker (1966) reported encountering a uniformly “sandy brown” specimen at Ghardaia and both black and brown specimens at Bir Ghellalia, Msila Province - Nine Tunisian specimens showed great variation, but most are yellowish or mottled brown, with head and neck blackish, the venter may be dark mesially or suffused with brown. MNHN 8797 from south Tunisia is red-brown above and purple below. Only a 446 mm male from Sfax (FMNH 83646) has a distinct throat band covering ventrals 8–18. Specimens from Libya are similar (see photo in Schleich et al., 1996, plate 49) but a 1370 mm male from Kouf National Park (FMNH 214914) is brown with yellow flecks above and shows faint banding, while a 610 mm male from near Misurata (FMNH 83058) has black throat bands covering V 3– 6 and 11–25. Kramer & Schnurrenberger (1963) reported that Libyan juveniles are cream with dark dorsal crossbands, black head and neck and a broad black throat band. - Egyptian specimens are yellow to brown above, often mottled, and the head often darker, with faint darker edges on the head scales and an indication of a “teardrop” marking. FMNH 171897 from Hahîg, Matruh, has 3 yellow bands on the posterior body and 3 on the tail (3 + 3), and FMNH 75232 from northwest of Cairo has 4 + 2 similar bands. Most Egyptian cobras have a single dark throat band covering ventrals ca. 15–25. See photos in Saleh (1997, p. 175), Baha El Din (2006, fig. 109) and Dobiey & Vogel (2007, p. 66). - No material has been examined from northern Sudan, but most specimens from the south resemble those from Egypt. However, FMNH 190325, a female from Kassala, has a dark brown dorsum with pale streaks and 9 + 2 yellow bands on body and tail, these extend ventrally. FMNH 58468, a 412 mm female from Torit, has two black neck bands on V 12–13 and 15–29, while NMK 3231, a 1880 mm male from Sennar, has brown bands on V 1–9 and 13–27. - Ethiopian specimens are usually brown with numerous scattered patches of yellow scales, sometimes with a divided yellow band on the neck, but AAU H. 664, a male from north of Gondar, has yellow blotches coalescing to form bands caudad, while the venter is blackish with 9 + 2 distinct yellow bands. Dobiey & Vogel (2007, p. 65) illustrate a specimen from Keren, NW Eritrea, which displays a striking pattern of dark brown or black marbling on a creamy-white background. It is unclear whether this is an individual aberrance or a characteristic of the local population. - Ugandan specimens are yellow to grey-brown, the head and neck often darker and frequently with a faint yellow band on the neck. MUZM (un-numbered), a 900 mm female from Soroti, is black above with 7 + 2 yellow bands, and two juveniles (NMZB-UM 5236 – 7) from this area show faint banding on the dorsum. A black throat band usually covers ca. v 12–24. - Kenyan specimens are usually mottled brown and yellow above, with contrasting facial and supralabial markings, sometimes with a yellow band on the neck, rarely other bands caudad. Usually a dark throat band covers ca. V 16–25, rest of venter yellow with brown blotches. Tanzanian specimens are similar, but KMH 3184, a 1063mm female from Mangola, is grey-brown above, with one yellow band on the nape, three on the posterior body and two on the tail. - There are few records of N. haje from the north-eastern Democratic Republic of Congo (DRC) and northern Central African Republic (CAR). One of us (LC) collected four specimens in northern CAR, which do not differ from those of northern Cameroon, where all the specimens collected by one of us (LC) are very dark, grey or black but not brown, with a pale throat, and superficially similar to Moroccan specimens (photograph in Chirio & LeBreton, 2007, p. 579). Naja senegalensis differs from N. anchietae, N. annulifera and N. arabica in having consistently higher scale row numbers around the neck (23 or more vs. 21 or fewer in the three other species). Additionally, N. anchietae and N. annulifera differ in having a pointed, enlarged rostral scale, fewer ventral scales (males: maximum 201; females: maximum 206, vs. minimally 205 and 219, respectively, in N. senegalensis) and, with few exceptions, 19 or fewer midbody dorsal scale rows. Naja arabica also differs from N. senegalensis in colour pattern, which is highly variable (Gasperetti, 1988; Egan, 2007). Specimens from south-west Saudi Arabia and Yemen may be blackish-brown above and below, or with the head and neck black, the rest of the body yellow, the venter often dark mesially. Gasperetti (1988) noted that some individuals were dull black, copper coloured, or various shades of brown or yellow, with blackish top of head and tail, and Egan (2007) additionally noted entirely orange specimens with yellow heads. An adult female from Dhofar, Oman (BMNH 1976.1487) is yellow-brown, but with a black head and neck and becoming black caudad and with a black venter. A 418mm male from the same region (BMNH 1977.1198) has a brown head, yellow-brown dorsum and yellow venter, and van der Kooij (2001, p. 59) illustrates a largely black specimen with coppery lower sides and described a “copper coloured ventral surface”, although this is not evident in the photo. Description of holotype (Fig. 4–6). The holotype (MNHN 2008.0074, previously IRD S- 8549) is an adult male of the following dimensions: total length 1430 mm, snout-vent length 1175 mm, tail length 255 mm, ratio total length: tail length 5.3. Head broad and short, weakly distinct from the neck, which is partly dilated. Snout rounded. Eye small, pupil round. Rostral as broad as high, clearly visible from above. The nostril is large and entirely divides the nasal. Two internasals, two prefrontals. The frontal is slightly longer than the prefrontals and internasals, their greatest width is similar. Loreal absent. A single rectangular preocular, twice as long as wide, between eye and nasal. Two postoculars on the left, three on the right. Two suboculars on the left and three on the right entirely separate the eye from the supralabials. 1 + 2 temporals on right, 1 + 3 on left. Seven supralabials, sixth is largest. Eight infralabials, the first four contact the anterior chin shields. No cuneates. The posterior chin shields are as long as but narrower than the anterior ones. Dorsal scales smooth and oblique, in 25 rows around the neck, 21 around midbody and 15 ahead of the vent. Vertebral row not enlarged. 211 ventrals, anal single. 65 subcaudals, all divided except the second to the ninth, which are single. Stomach content: one Bufo xeros. Upper side of head, body and tail entirely grey-brown. Lower flanks lighter on first two dorsal scale rows, except at anterior and posterior end of body, where they are of the same colour as the dorsum. Lower side of head is grey-brown, similar to the upper side. Underside includes a dark grey area extending from the fifth to the 30 th ventral scale, excluding the 12 th and 17 th ventrals, which are partially light. From the 31 st ventral, the dominant colour of the ventrals is yellowish, with dark spots that become fainter towards the posterior part of the body. Subcaudals entirely yellowish, except on the terminal third of the tail, where they become progressively darker. Description of paratypes. The 31 paratypes include 17 males and 14 females. The largest male (IRD S- 3429) measured 2065 mm, the largest female (IRD S- 1640) 2315 mm in total length. Mean length of males was 1035 mm (SD = 699 mm), of females 1110 mm (SD = 654 mm). The total length: tail length ratio ranged from 5.7 to 6.6 in males (mean: 6.2; SD: 0.2) and from 6.1 to 6.8 in females (mean 6.4, SD 0.3). Midbody dorsal scale rows 21 in males and 21 (N = 13) or 23 (N = 1) in females. The number of scale rows around the neck is 23 (3 males, 1 female), 24 (1 female), 25 (13 males and 9 females), 26 (1 female) or 27 (2 males, 2 females). Ventrals 205–216 (mean 211.7, SD 2.7) in males, 219–225 (mean 222.3, SD 1.6) in females. Subcaudals 59–65 in males (mean 61.5, SD 1.5) and 56–64 in females (mean 59.9, SD 2.2), all or mostly divided. Nasal always fully divided, loreal always absent. Preocular always single, elongate and rectangular. 1–3 postoculars, 1–3 suboculars, the total number of scales around the eye varying from 5 to 7. Supralabials always 7, except in one specimen with 8 on one side. Temporals 1 + 2 (N = 5), 1 + 3 (N = 18), or a combination of the above (N = 9

Naja incertae sedis Three of the four currently recognized extinct cobra species, † Naja antiqua Rage, 1976, † Naja iberica Szyndlar, 1985, and † Naja robusta Meylan, 1987, cannot be allocated to any subgenus with confidence, as identifying synapomorphies are lacking (Rage, 1976; Szyndlar, 1985; Meylan, 1987). They are therefore placed in Naja incertae sedis., Published as part of Wallach, Van, Wüster, Wolfgang & Broadley, Donald G., 2009, In praise of subgenera: taxonomic status of cobras of the genus Naja Laurenti (Serpentes: Elapidae), pp. 26-36 in Zootaxa 2236 on page 32, DOI: 10.5281/zenodo.190422, {"references":["Rage, J. - C. (1976) Les squamates du Miocene de Beni Mellal. Geologie Mediterraneenne, 3 (2), 57 - 69.","Szyndlar, Z. (1985) Ophidian fauna (Reptilia, Serpentes) from the uppermost Miocene of Algora (Spain). Estudios Geologicos, 41, 447 - 465.","Meylan, P. A. (1987) Reptiles and birds. Fossil snakes from Laetoli. In: Leakey, M. D. & Harris, J. M. (Eds.), The Pliocene site of Laetoli, northern Tanzania, Oxford University Press, Oxford, pp. 78 - 82."]}

Subgenus Naja Laurenti, 1768 Naia Merrem, 1820: 147 (unjustified emendation of Naja Laurenti, 1768) Aspis Wagler, 1830: 173 (not Laurenti) (type species Coluber naja Linnaeus, 1758) Tomyris Eichwald, 1831: 171 (type species Tomyris oxiana Eichwald, 1831) Naga Nicholson, 1874: 104 (unjustified emendation of Naja Laurenti, 1768) † Palaeonaja Hoffstetter, 1939: 57 (type species † Palaeonaja romani Hoffstetter, 1939) Type species: Naja lutescens Laurenti, 1768 (= Coluber naja Linnaeus, 1758), by subsequent designation (Leviton, 1968). Gender: feminine. Etymology: derived from the Sinhala Naya, cobra. Distribution: southern and south-eastern Asia and the East Indies, from Transcaspia to the Philippines and the Lesser Sunda Islands. Content: eleven species: Naja (Naja) atra Cantor, 1842: 482 Naja (Naja) kaouthia Lesson, 1831: 122 Naja (Naja) mandalayensis Slowinski & Wüster, 2000: 260 Naja (Naja) naja (Linnaeus, 1758: 221) Naja (Naja) oxiana (Eichwald, 1831: 171) Naja (Naja) philippinensis Taylor, 1922: 265 Naja (Naja) sagittifera Wall, 1913: 247 Naja (Naja) samarensis Peters, 1861: 690 Naja (Naja) siamensis Laurenti, 1768: 91 Naja (Naja) sputatrix Boie, 1827: 557 Naja (Naja) sumatrana Müller, 1890: 277 Diagnosis: Extracranial (ventral) anterior Vidian canal position, 0–1 solid maxillary teeth in all species (Wüster, 1990 —only 6 out of 650 specimens examined in that study had 2 solid maxillary teeth), seven supralabials with penultimate (sixth) shield low, combination of single preocular and two (occasionally three) anterior temporals, rostral broader than deep; internasals shorter than prefrontals; fang structure variable, all species except N. naja and N. oxiana have some degree of adaptation to spitting (Wüster & Thorpe, 1992 b). We tentatively include the extinct † Naja (Naja) romani (Hofstetter, 1939) in this subgenus based on the shared derived condition of the basisphenoid morphology and the vestibular window, despite the possession of two solid maxillary teeth (Szyndlar & Rage, 1990). Comments: this is a morphologically relatively conserved, but ecologically highly adaptable subgenus that appears to be the result of a single colonization event of Asia from an African origin (Slowinski & Wüster, 2000; Wüster et al., 2007; Wüster, unpublished data). The issue of the type species of Naja has a complex background. Laurenti’s (1768) Naja was based upon six species from Seba (1734 –1735), all of which Linnaeus (1758) included in his synonymy of Coluber naja (in addition to 1735: pl. 85, fig. 1 and 1735: pl. 94, fig. 1): N. brasiliensis (1735: pl. 89, fig. 4) = Naja naja, N. fasciata (1735: pl. 89, fig. 3) = Naja naja, N. lutescens (1734: pl. 44, fig. 1) = Naja naja, N. maculata (1735: pl. 90, fig. 2) = Naja naja, N. non Naja (1735: pl. 90, fig. 1) = Naja kaouthia, and N. siamensis (1735: pl. 89, figs. 1–2) = Naja siamensis. Naja naja (Linnaeus) has been considered the type species of Naja by tautonymy, monotypy, and subsequent designation of Stejneger (1936: 140), M. Smith (1943: 426), Oshima (1944: 204), and others. However, David & Vogel (1996: 146) suggested that those assumptions were incorrect and that the only valid type species designation was that of Williams & Wallach (1989: 97), who selected Naja lutescens. However, Leviton (1968: 547) designated Naja lutescens Laurenti (= Coluber naja Linnaeus) as the type species of Naja Laurenti and several earlier nomenclatural acts precede this action. Cantor (1847: 1038) could be considered the first revisor as he synonymized Naja lutescens with Coluber naja Linnaeus. Deraniyagala (1945: 108–109) then restricted the name Naja lutescens to the race of cobras inhabiting India south of 20 ° N Latitude. The type locality of Seba’s (1735) pl. 44, fig. 1 was given as “ India Orientali.” Deraniyagala (1945) recognized Naja naja lutescens as a subspecies and designated the type locality as Madras, with N. fasciata and N. maculata as synonyms., Published as part of Wallach, Van, Wüster, Wolfgang & Broadley, Donald G., 2009, In praise of subgenera: taxonomic status of cobras of the genus Naja Laurenti (Serpentes: Elapidae), pp. 26-36 in Zootaxa 2236 on pages 29-30, DOI: 10.5281/zenodo.190422, {"references":["Laurenti, J. N. (1768) Austriaci Viennensis Specimen Medicum, Exhibens Synopsin Reptilium Emendatam Cum Experimentis Circa Venena et Antidota Reptilium Austriacorum. J. T. De Trattern, Viennae, 214 pp.","Merrem, B. (1820) Versuch eines Systems der Amphibien. Johann Christian Krieger, Marburg, xv + 191 pp., 1 Pl.","Wagler, J. (1830) Naturliches System der Amphibien, mir Vorangehender Classification der Saugethiere und Vogel. Ein Beitrag zur Vergleichenden Zoologie. J. G. Cotta, Munchen, 354 pp.","Linnaeus, C. von. (1758) Systema Naturae per Regna Tria Naturae secundum Classes, Ordines, Genera, Species cum Characteribus Differentiis, Synonymis, Locis. Stockholm, Laurentii Salvii, 824 pp.","Eichwald, E. (1831) Zoologia specialis quam expositis animalibus tum vivis, tum fossilibus potissimum Rossiae in universum, et Poloniae in species, in usum lectionum publicarum in Universitate Caesareae Vilnensi habendarum. Part 3. Josephi Zawadski, Vilnius, 396 pp.","Nicholson, E. (1874) Indian snakes. An elementary treatise on ophiology with a descriptive catalogue of the snakes found in India and the adjoining countries. Second edition. Higginbotham and Company, Madras, 188 pp., 20 Pls.","Hoffstetter, R. (1939) Contribution a l'etude des Elapidae actuels et fossiles et de l'osteologie des ophidiens. Archives du Museum d'Histoire Naturelle de Lyon, 15, 1 - 78, 2 Pls.","Leviton, A. E. (1968) The venomous terrestrial snakes of East Asia, India, Malaya, and Indonesia. In: Bucherl, W., Buckley, E. E. & Deulofeu, V. (Eds.), Venomous animals and their venoms. Volume I. Venomous vertebrates. Academic Press, New York, pp. 529 - 576.","Cantor, T. E. (1842) General features of Chusan, with remarks on the flora and fauna of that island. Annals and Magazine of Natural History, 1, 265 - 277.","Lesson, R. P. (1831) Catalogue des reptiles qui font partie d'une collection zoologique recueillie dans l'Inde continentale ou en Afrique et rapportee en France par M. Lamare-Piquot. Bulletin des Sciences Naturelles et de Geologie, Paris, 25, 119 - 123.","Slowinski, J. B. & Wuster, W. (2000) A new cobra (Elapidae: Naja) from Myanmar (Burma). Herpetologica, 56, 257 - 270.","Taylor, E. H. (1922) The Snakes of the Philippine Islands. Bureau of Printing, Manila, 312 pp.","Wall, F. (1913) A popular treatise on common Indian snakes. Naia tripudians (Merrem). The cobra. Journal of the Bombay Natural History Society, 22, 243 - 259.","Peters, W. K. H. (1861) Eine zweite Ubersicht (vergl. Monatsberichte 1859 p. 269) der von Hrn. F. Jagor auf Malacca, Java, Borneo und den Philippinen gesammelten und dem Kgl. Zoologischen Museum ubersandten Schlangen. Monatsberichte der Koniglichen Preussischen Akademie der Wissenschaften zu Berlin, 1861, 683 - 691.","Boie, F. (1827) Bemerkungen uber Merrem's Versuch eines Systems der Amphibien. Erste Lieferung: Ophidier. Isis von Oken, 20, col. 508 - 566.","Wuster, W. (1990) Population evolution of the Asiatic cobra (Naja naja) species complex. PhD Thesis, University of Aberdeen.","Wuster, W. & Thorpe, R. S. (1992 b) Dentitional phenomena in cobras revisited: fang structure and spitting in the Asiatic species of Naja (Serpentes: Elapidae). Herpetologica, 48, 424 - 434.","Szyndlar, Z. & Rage, J. - C. (1990) West Palearctic cobras of the genus Naja (Serpentes: Elapidae): interrelationships among extinct and extant species. Amphibia-Reptilia, 11, 385 - 400.","Seba, A. (1734 - 1735) Locupletissimi rerum naturalium thesauri accurata descriptio, et iconibus artificiosissimis expressio, per universam physices historiam. Tomus I-II. Janssonio-Waesbergios, J. Wetstenium & Gul. Smith, Amstelaedami, 178 pp., 111 Pls. & 154 pp., 114 Pls.","Stejneger, L. (1936) Types of the amphibian and reptilian genera proposed by Laurenti in 1768. Copeia, 1936 (3), 133 - 141.","Smith, M. A. (1943) The fauna of British India, Ceylon and Burma, including the whole of the Indo-Chinese sub-region. Reptilia and Amphibia. Vol. III. - Serpentes. Taylor & Francis, London, 583 pp.","Oshima, M. (1944) \" Poisonous snakes in East and South Asia. \" Hokuryukan, Tokyo, 330 + 19 pp., 18 Pls. [in Chinese] Peters, W. K. H. (1854) Diagnosen neuer Batrachier, welche zusammen mit der fruher (24. Juli und 17. August) gegebenen Ubersicht der Schlangen und Eidechsen mitgetheilt werden. Bericht uber zur Bekanntmachung geeignete Verhandlungen der Koniglichen Preussischen Akademie der Wissenschaften zu Berlin, 1854, 614 - 628.","David, P. & Vogel, G. (1996) The snakes of Sumatra: an annotated checklist and key with natural history notes. Edition Chimaira, Frankfurt am Main, 260 pp., 8 Pls.","Williams, K. L. & Wallach, V. (1989) Snakes of the world. Volume I. Synopsis of snake generic names. Krieger Pubishing Company, Malabar, 234 pp.","Cantor, T. E. (1847) Catalogue of reptiles inhabiting the Malayan Peninsula and islands, collected or observed by Theodore Cantor. Venomous serpents. Journal of the Asiatic Society of Bengal, 16, 1026 - 1078.","Deraniyagala, P. E. P. (1945) Some new races of the python, Chrysopelea, binocellate cobra, and tith-polonga inhabiting Ceylon and India. Spolia Zeylanica, 24 (2), 103 - 112."]}

Naja senegalensis sp. nov. Trape, Chirio and W��ster Figs. 4���8 Holotype: MNHN 2008.0074 (previously IRD S- 8549), collected in September 2008 near Dielmo (13 �� 43 ���N, 16 �� 25 ���W) by Mr. Babacar N���Dao, veterinary agent at Keur Lahim Fatim, who sent it to the first author (Fig. 4���6). Paratypes: 31 specimens, all from S��n��gal: MNHN 2008.0075 (previously IRD S- 409), MNHN 2008.0076 (previously IRD S- 1027) MNHN 2008.0077 (previously IRD S- 1028), MNHN 2008.0078 (previously IRD S- 1578), MNHN 2008.0079 (previously IRD S- 1589): Keur Lahine Fatim (13 �� 44 ���N, 16 �� 23 ���N), Sine Saloum; MNHN 2008.0080 (previously IRD S- 443), MNHN 2008.0081 (previously IRD S- 2302), MNHN 2008.0082 (previously IRD S- 2306): Keur Bakar Man�� (13 �� 37 ���N, 16 �� 17 ���W), Sine Saloum; MNHN 2008.0083 (previously IRD S- 762), MNHN 2008.0084 (previously IRD S- 5283): Keur Seny Gueye (13 �� 36 ���N, 16 �� 19 ���W), Sine Saloum; MNHN 2008.0085 (previously IRD S- 855), MNHN 2008.0086 (previously IRD S- 858), MNHN 2008.0087 (previously IRD S- 1640): Keur Gadji (13 �� 38 ���N, 16 �� 19 ���W), Sine Saloum; MNHN 2008.0088 (previously IRD S- 1283): Keur Santhiou (13 �� 39 ���N, 16 �� 21 ���W), Sine Saloum; IRSNB 2654 (previously IRD S- 605), IRSNB 2655 (previously IRD S- 1429), IRSNB 2656 (previously IRD S- 1435), IRD S- 1439, IRD S- 1440, IRD S- 1442, IRD S- 1461, IRD S- 3429, IRD S- 3430, IRD S- 5692: Dielmo (13 �� 43 ���N, 16 �� 25 ���N), Sine Saloum; IRD S- 2113: Landieni (12 �� 33 ���N, 12 �� 22 ���W), eastern Senegal; IRD S- 5344: Saroudia (12 �� 32 ���N, 11 �� 35 ���W), eastern Senegal; IRD S- 5427: Sambarabougou (13 ��06���N, 11 �� 51 ���W), eastern Senegal; IRD S- 5849, IRD S- 5854: Gu��noto (13 �� 33 ���N, 13 �� 50 ���W), eastern Senegal; IRD S- 6204: Keur Lamine Diam�� (13 �� 37 ���N, 16 �� 16 ���W), Sine Saloum; IRD S- 6461: Touba Baria (13 �� 38 ���N, 16 �� 14 ���W), Sine Saloum. Other specimens examined. 86 specimens: Senegal (46 specimens): IRD S- 343, IRD S- 3431: Senegal; IRD S- 462: Keur Ayip K�� (13 �� 39 ���N, 16 �� 19 ���W), Sine Saloum; IRD S- 604, IRD S- 606, IRD S- 664: Keur Bakar Man�� (13 �� 37 ���N, 16 �� 17 ���W), Sine Saloum; IRD S- 856, IRD S- 1634: Keur Gadji (13 �� 38 ���N, 16 �� 19 ���W), Sine Saloum; IRD S- 1279, S- 1280, S- 1281, S- 1292, S- 1293: Keur Santhiou (13 �� 39 ���N, 16 �� 21 ���W), Sine Saloum; S- 1411, S- 1472, S- 1482: Dielmo (13 �� 43 ���N, 16 �� 25 ���W), Sine Saloum; IRD S- 1588: Keur Lahim Fatim (13 �� 44 ���N, 16 �� 23 ���N), Sine Saloum; IRD S- 3849: Badiara (13 �� 13 ���N, 14 �� 12 ���W), Haute Casamance; IRD S- 3952: Goundaga (12 �� 51 ���N, 14 ��05���W), Haute Casamance; IRD S- 4806: Tial�� (15 �� 14 ���N, 16 �� 49 ���W), Cayor; IRD S- 5085: Oubadji (12 �� 40 ���N, 13 ��03���W), S��n��gal oriental; IRD S- 5307: Saroudia (12 �� 32 ���N, 11 �� 35 ���W), S��n��gal oriental; IRD S- 5795, IRD S- 6090: Keur Momat Souna (13 �� 38 ���N, 16 �� 17 ���W), Sine Saloum; IRD S- 5851, IRD S- 5853: Gu��noto (13 �� 33 ���N, 13 �� 50 ���W), eastern Senegal; IRD S- 5862: M��dina Djikoye (13 �� 37 ���N, 16 �� 18 ���W), Sine Saloum; IRD S- 6239: Touba Baria (13 �� 38 ���N, 16 �� 14 ���W), Sine Saloum; IRD S- 6656: Takoudialla (12 �� 50 ���N, 14 ��04���N), Haute Casamance; IRD S- 6680: S��goto (13 �� 18 ���N, 11 �� 49 ���W), S��n��gal oriental; IRD S- 7200: Touba Ndiaye (15 ��09���N, 16 �� 52 ���W), Cayor; IFAN 55 - 4-13: Camb��r��ne (14 �� 45 'N, 17 �� 25 'W); IFAN 52 - 11-90, IFAN 53 - 11-143: Dakar (14 �� 42 'N, 17 �� 27 'W); IFAN 47 - 1-10, IFAN 47 - 1-15, IFAN 50 - 9-149, IFAN 51 - 12-53, IFAN 52 - 3-23, IFAN 52 - 7-47, IFAN 56 - 5-50: Hann (14 �� 43 'N, 17 �� 26 'W), IFAN 82 - 1-2: Keur Massar (14 �� 47 'N, 17 �� 18 'W); IFAN 52 - 1-8: Malika (14 �� 47 'N, 17 �� 20 'W); IFAN 53 - 3-20: Ouakam (14 �� 43 'N, 17 �� 29 'W); IFAN 44 - 1-3: Popenguine (14 �� 33 'N, 17 ��07'W). Mali (32 specimens): IRD 2353 -M, IRD 2354 -M: Ballabougou (12 �� 52 ���N, 06�� 52 ���W); IRD 238 -M: Bangaya (13 �� 14 ���N, 10 �� 43 ���W); IRD 1179 -M, IRD 1181 -M: Djinagu�� (12 �� 59 ���N, 09�� 52 ���W); IRD 103 -M: Doussoudiana (11 ��09���N, 07�� 48 ���W); IRD 2368 -M: Koundian (13 �� 10 ���N, 10 �� 40 ���W); IRD 805 -M: Laminina (11 �� 12 ���N, 07�� 47 ���W); IRD 1977 -M, 2003 -M, 2017 -M: Mamoroubougou (11 �� 13 ���N, 05�� 28 ���W); IRD 1796 -M, IRD 3419 -M: Npi��bougou (11 �� 59 ���N, 08��00���W); IRD 2352 -M: Sadjouroubougou (12 �� 35 ���N, 07�� 44 ���W); IRD 1591 -M: Sare-Soma (14 �� 45 ���N, 03�� 55 ���W); IRD 878 -M, 957 -M: Sebekourani (12 �� 12 ���N, 08�� 42 ���W); IRD 184 -M, IRD 2102 -M, IRD 2109 -M, IRD 2118 -M, IRD 2145 - M, IRD 2186 -M, IRD 3606 -M, IRD 3617 -M, IRD 3618 -M, IRD 3683 -M: Titi��na (11 �� 27 ���N, 06�� 33 ���W); IRD 156 -M, IRD 581 -M, IRD 2349 -M, IRD 2350 -M, IRD 2351 -M (13 ��09���N, 07�� 57 ���W). Niger (2 specimens): IRD 201 -N: Karosofua (13 �� 37 ���N, 06�� 37 ���E); IRD 1504 -N: T��la (12 ��08���N, 03�� 28 ���E). Burkina Faso (2 specimen): LC 6531: Kondio (11 �� 37 ���N, 02��01���E); IFAN 48 - 2-9: Dano pr��s Di��bougou (11 ��09'N, 03��04'W); USNM 237088 8km S of Dana (NW 1202 C 1). B��nin (1 sp��cimen): LC 7109: Ni��ni�� (11 �� 22 ���N, 02�� 12 ���E). Guin��e (1 specimen): IFAN 52 - 6-34: Niandan-Banie (approximatively 10 �� 20 'N, 09�� 50 'W). Nigeria (1 specimen): CM 92607 Shagunu, west bank of Kainji Lake (10 �� 20 ���N, 04�� 28 ���E). Diagnosis. Naja senegalensis resembles all other members of the N. haje complex and differs from all other Naja in having a row of subocular scales separating the orbit from the supralabial scales. Naja senegalensis can be distinguished from other species of the N. haje complex through a combination of scale counts and the coloration of juveniles and adults. Comparative scale counts are given in Table 3. Naja senegalensis is distinguishable from N. haje through its higher neck scale row count: N. senegalensis normally has 25 dorsal scale rows around the neck, although some specimens have 23 or 27. By contrast, W. African N. haje have fewer neck scale rows (19���21 in five specimens from Niger, 21���23 in three specimens from Nigeria, 21 in one specimen from Tombouctou, Mali). In other parts of Africa, the majority of specimens also have 21 or fewer scale rows around the neck (Table 3). A cobra specimen from Djibouti, with 27 scale rows around the neck and 23 at midbody, tentatively assigned to the N. haje complex by Ineich (2001), appears to be a spitting cobra. Other scalation characters do not distinguish N. senegalensis from N. haje, although the new species tends to occupy the upper end of the spectrum of ventral scale counts in the complex (Table 3). Another diagnostic feature of N. senegalensis is the juvenile pattern: the great majority of juveniles and subadult specimens have a highly contrasting white blotch on the neck, within the dark collar encircling the neck (Fig. 7). This pale patch is present in 37 out of 39 small and medium-sized specimens, but barely discernible or absent in almost all larger adults. However, one of us (LC) recently photographed a large captive adult (approximately 200 cm total length) from the area of Bamako, Mali, that retained a very conspicuous, heart-shaped nuchal mark (Fig. 8). We have never observed this patch in N. haje, and therefore consider its presence to be a diagnostic character for N. senegalensis. Naja senegalensis differs strongly in coloration from sympatric or parapatric West African N. haje: adults of N. senegalensis are almost invariably uniformly dark brown dorsally, whereas juveniles are greyish dorsally and yellowish ventrally, with a dark collar (Fig. 7) around the neck and usually a white neck blotch. Small adult specimens tend to be dark brown with paler speckles and their ventral side is yellowish. Small adult N. senegalensis from W National Park, in the Niger ��� Burkina Faso ��� Benin border region, have a brown dorsal coloration with small reddish dots of one scale each. The entire head, and in particular the supralabial region, are normally uniformly dark brown. In Niger and Nigeria, where both N. senegalensis and N. haje are found (the former in the Sudan savanna, the latter in the Sahelian zone, but with possible areas of sympatry), both adults and juveniles of N. haje display quite different colours: the body of adults is yellow to dark brown dorsally (often mostly yellow, especially in Niger), often with scattered individual dark scales, but the venter is at least partly cream-coloured, or with contrasting light and dark bands or blotches. The sides of the head, and in particular the supralabial region, normally display contrasting areas of pale and dark pigmentation, and, most noticeably, a dark spot under the eye, reminiscent of the ���teardrop��� marking present in N. nubiae (W��ster & Broadley, 2003), and a more or less distinct dark greyish neck band, approximately ten scales wide. Juveniles lack the pale neck patch present in young N. senegalensis. An adult N. haje from Niger is illustrated in Trape & Man�� (2006, p. 195). Elsewhere in Africa, N. haje shows great variation in colour pattern, but differs consistently from that shown in N. senegalensis as follows: - The Moroccan/Western Saharan population extends into the northern parts of Western Sahara, and specimens have been illustrated by Bons & Geniez (1996, p. 251), Geniez et al. (2004, p. 169, 171) and Dobiey & Vogel (2007, p. 67). Adults are usually uniform black except for a yellowish gular area. Some may be dark brown above and grey below. - N. haje extends across northern Algeria south of the Atlas Mountains (Schleich et al., 1996), but no voucher specimens have been examined. LC did not encounter the species during two years that he spent in the region; local people knew it, but reported that it was very rare. Renker (1966) reported encountering a uniformly ���sandy brown��� specimen at Ghardaia and both black and brown specimens at Bir Ghellalia, Msila Province - Nine Tunisian specimens showed great variation, but most are yellowish or mottled brown, with head and neck blackish, the venter may be dark mesially or suffused with brown. MNHN 8797 from south Tunisia is red-brown above and purple below. Only a 446 mm male from Sfax (FMNH 83646) has a distinct throat band covering ventrals 8���18. Specimens from Libya are similar (see photo in Schleich et al., 1996, plate 49) but a 1370 mm male from Kouf National Park (FMNH 214914) is brown with yellow flecks above and shows faint banding, while a 610 mm male from near Misurata (FMNH 83058) has black throat bands covering V 3��� 6 and 11���25. Kramer & Schnurrenberger (1963) reported that Libyan juveniles are cream with dark dorsal crossbands, black head and neck and a broad black throat band. - Egyptian specimens are yellow to brown above, often mottled, and the head often darker, with faint darker edges on the head scales and an indication of a ���teardrop��� marking. FMNH 171897 from Hah��g, Matruh, has 3 yellow bands on the posterior body and 3 on the tail (3 + 3), and FMNH 75232 from northwest of Cairo has 4 + 2 similar bands. Most Egyptian cobras have a single dark throat band covering ventrals ca. 15���25. See photos in Saleh (1997, p. 175), Baha El Din (2006, fig. 109) and Dobiey & Vogel (2007, p. 66). - No material has been examined from northern Sudan, but most specimens from the south resemble those from Egypt. However, FMNH 190325, a female from Kassala, has a dark brown dorsum with pale streaks and 9 + 2 yellow bands on body and tail, these extend ventrally. FMNH 58468, a 412 mm female from Torit, has two black neck bands on V 12���13 and 15���29, while NMK 3231, a 1880 mm male from Sennar, has brown bands on V 1���9 and 13���27. - Ethiopian specimens are usually brown with numerous scattered patches of yellow scales, sometimes with a divided yellow band on the neck, but AAU H. 664, a male from north of Gondar, has yellow blotches coalescing to form bands caudad, while the venter is blackish with 9 + 2 distinct yellow bands. Dobiey & Vogel (2007, p. 65) illustrate a specimen from Keren, NW Eritrea, which displays a striking pattern of dark brown or black marbling on a creamy-white background. It is unclear whether this is an individual aberrance or a characteristic of the local population. - Ugandan specimens are yellow to grey-brown, the head and neck often darker and frequently with a faint yellow band on the neck. MUZM (un-numbered), a 900 mm female from Soroti, is black above with 7 + 2 yellow bands, and two juveniles (NMZB-UM 5236 ��� 7) from this area show faint banding on the dorsum. A black throat band usually covers ca. v 12���24. - Kenyan specimens are usually mottled brown and yellow above, with contrasting facial and supralabial markings, sometimes with a yellow band on the neck, rarely other bands caudad. Usually a dark throat band covers ca. V 16���25, rest of venter yellow with brown blotches. Tanzanian specimens are similar, but KMH 3184, a 1063mm female from Mangola, is grey-brown above, with one yellow band on the nape, three on the posterior body and two on the tail. - There are few records of N. haje from the north-eastern Democratic Republic of Congo (DRC) and northern Central African Republic (CAR). One of us (LC) collected four specimens in northern CAR, which do not differ from those of northern Cameroon, where all the specimens collected by one of us (LC) are very dark, grey or black but not brown, with a pale throat, and superficially similar to Moroccan specimens (photograph in Chirio & LeBreton, 2007, p. 579). Naja senegalensis differs from N. anchietae, N. annulifera and N. arabica in having consistently higher scale row numbers around the neck (23 or more vs. 21 or fewer in the three other species). Additionally, N. anchietae and N. annulifera differ in having a pointed, enlarged rostral scale, fewer ventral scales (males: maximum 201; females: maximum 206, vs. minimally 205 and 219, respectively, in N. senegalensis) and, with few exceptions, 19 or fewer midbody dorsal scale rows. Naja arabica also differs from N. senegalensis in colour pattern, which is highly variable (Gasperetti, 1988; Egan, 2007). Specimens from south-west Saudi Arabia and Yemen may be blackish-brown above and below, or with the head and neck black, the rest of the body yellow, the venter often dark mesially. Gasperetti (1988) noted that some individuals were dull black, copper coloured, or various shades of brown or yellow, with blackish top of head and tail, and Egan (2007) additionally noted entirely orange specimens with yellow heads. An adult female from Dhofar, Oman (BMNH 1976.1487) is yellow-brown, but with a black head and neck and becoming black caudad and with a black venter. A 418mm male from the same region (BMNH 1977.1198) has a brown head, yellow-brown dorsum and yellow venter, and van der Kooij (2001, p. 59) illustrates a largely black specimen with coppery lower sides and described a ���copper coloured ventral surface���, although this is not evident in the photo. Description of holotype (Fig. 4���6). The holotype (MNHN 2008.0074, previously IRD S- 8549) is an adult male of the following dimensions: total length 1430 mm, snout-vent length 1175 mm, tail length 255 mm, ratio total length: tail length 5.3. Head broad and short, weakly distinct from the neck, which is partly dilated. Snout rounded. Eye small, pupil round. Rostral as broad as high, clearly visible from above. The nostril is large and entirely divides the nasal. Two internasals, two prefrontals. The frontal is slightly longer than the prefrontals and internasals, their greatest width is similar. Loreal absent. A single rectangular preocular, twice as long as wide, between eye and nasal. Two postoculars on the left, three on the right. Two suboculars on the left and three on the right entirely separate the eye from the supralabials. 1 + 2 temporals on right, 1 + 3 on left. Seven supralabials, sixth is largest. Eight infralabials, the first four contact the anterior chin shields. No cuneates. The posterior chin shields are as long as but narrower than the anterior ones. Dorsal scales smooth and oblique, in 25 rows around the neck, 21 around midbody and 15 ahead of the vent. Vertebral row not enlarged. 211 ventrals, anal single. 65 subcaudals, all divided except the second to the ninth, which are single. Stomach content: one Bufo xeros. Upper side of head, body and tail entirely grey-brown. Lower flanks lighter on first two dorsal scale rows, except at anterior and posterior end of body, where they are of the same colour as the dorsum. Lower side of head is grey-brown, similar to the upper side. Underside includes a dark grey area extending from the fifth to the 30 th ventral scale, excluding the 12 th and 17 th ventrals, which are partially light. From the 31 st ventral, the dominant colour of the ventrals is yellowish, with dark spots that become fainter towards the posterior part of the body. Subcaudals entirely yellowish, except on the terminal third of the tail, where they become progressively darker. Description of paratypes. The 31 paratypes include 17 males and 14 females. The largest male (IRD S- 3429) measured 2065 mm, the largest female (IRD S- 1640) 2315 mm in total length. Mean length of males was 1035 mm (SD = 699 mm), of females 1110 mm (SD = 654 mm). The total length: tail length ratio ranged from 5.7 to 6.6 in males (mean: 6.2; SD: 0.2) and from 6.1 to 6.8 in females (mean 6.4, SD 0.3). Midbody dorsal scale rows 21 in males and 21 (N = 13) or 23 (N = 1) in females. The number of scale rows around the neck is 23 (3 males, 1 female), 24 (1 female), 25 (13 males and 9 females), 26 (1 female) or 27 (2 males, 2 females). Ventrals 205���216 (mean 211.7, SD 2.7) in males, 219���225 (mean 222.3, SD 1.6) in females. Subcaudals 59���65 in males (mean 61.5, SD 1.5) and 56���64 in females (mean 59.9, SD 2.2), all or mostly divided. Nasal always fully divided, loreal always absent. Preocular always single, elongate and rectangular. 1���3 postoculars, 1���3 suboculars, the total number of scales around the eye varying from 5 to 7. Supralabials always 7, except in one specimen with 8 on one side. Temporals 1 + 2 (N = 5), 1 + 3 (N = 18), or a combination of the above (N = 9, Published as part of Trape, Jean-Fran��ois, Chirio, Laurent, Broadley, Donald G. & W��ster, Wolfgang, 2009, Phylogeography and systematic revision of the Egyptian cobra (Serpentes: Elapidae: Naja haje) species complex, with the description of a new species from West Africa, pp. 1-25 in Zootaxa 2236 on pages 12-21, DOI: 10.5281/zenodo.190424, {"references":["Ineich, I. (2001) Reptiles et Amphibiens de la Republique de Djibouti. Bilan des Conaissances a l'Aube du Troisieme Millenaire. Rapport sur la mission herpetologique de Ivan Ineich, Avril 1999. Museum National d'Histoire Naturelle, Paris.","Wuster, W. & Broadley, D. G. 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Key to the species of the Naja haje species complex 1. West African savannas; 25 (rarely 23 or 27) scale rows around the neck; head and supralabial region dark, without pattern in adults; juveniles normally with a strongly contrasting white blotch on the dark neck .......... N. senegalensis - Elsewhere; 23 or fewer scale rows around neck; head and supralabial region often contrastingly patterned; juveniles without white blotch on neck....................................................................................................................................... 2 2. Africa south of Tanzania and Congo forests; rostral enlarged, snout pointed; normally 19 or fewer midbody dorsal scale rows...................................................................................................................................................................... 3 - Africa from Tanzania northward, Arabian Peninsula; rostral not enlarged; often 21 midbody dorsal scale rows....... 4 3. Western southern Africa; 17 midbody dorsal scale rows, 15 dorsal scale rows around neck .................... N. anchietae - Eastern southern Africa; 19 midbody dorsal scale rows, 19 dorsal scale rows around neck.................... N. annulifera 4. Arabian Peninsula; normally over 65 subcaudals; often 19 scale rows around neck, supralabials usually unpatterned ....................................................................................................................................................................... N. arabica - Africa from Tanzania northward; normally 65 or fewer subcaudals, rarely fewer than 21 scale rows around neck, supralabials often contrastingly patterned with ���teardrop��� marking, except in very dark animals ................... N. haje, Published as part of Trape, Jean-Fran��ois, Chirio, Laurent, Broadley, Donald G. & W��ster, Wolfgang, 2009, Phylogeography and systematic revision of the Egyptian cobra (Serpentes: Elapidae: Naja haje) species complex, with the description of a new species from West Africa, pp. 1-25 in Zootaxa 2236 on page 21, DOI: 10.5281/zenodo.190424

Naja ashei sp. nov. ��� Ashe���s spitting cobra Holotype. National Museums of Kenya NMK S/ 3993, a female specimen from Watamu, Kenya (3 �� 21 ���S: 40 �� 01���E), coll. Royjan Taylor, maintained in captivity at Bio-Ken Snake farm until 29 /09/ 2004 with reference number BK 10030 (Fig. 4,5). Paratypes (three males and two females): BMNH 1955.1.12.4a and 4 b (Kilifi, Kenya) BMNH 1963.456 (Kiboko, Kenya); BMNH 2005.1604 (Baringo, Kenya); NMZB 3349 (Ex USNM 40954) (Guaso Nyiro [=Ewaso Ng���iro], Kenya). Diagnosis. Naja ashei differs from all other African spitting cobras in possessing a unique clade of mtDNA haplotypes. From the data presented here, we identified 12 fixed nucleotide differences that differentiate N. ashei from the other eastern African spitting Naja. These correspond to positions 105, 169 and 315 of the ND 4 sequence of the holotype (DQ 897706), and to positions 60, 108, 153, 201, 348, 381, 507, 630 and 676 of the cytochrome b sequence of the same specimen (DQ 897749), the diagnostic bases at these positions being C, T, G, C, G, T, T, T, A, C, T and A, respectively. Morphologically, N. ashei differs from East African N. nigricollis in a number of characters relating to adult colour pattern and scalation. In particular, its midbody and posterior ventral colour is predominantly light, with dark pigment encroaching mostly from the sides of the body (venter normally largely or entirely dark in N. nigricollis), it lacks any red, orange or pink pigment under the throat (usually pronounced in N. nigricollis), and the head is the same olive-brown colour as the rest of the body (often black above and below in East African N. nigricollis). Scalation does not provide any absolutely diagnostic characters for N. ashei, but mean scale counts and the range differ clearly from those of East African N. nigricollis (Table 4). In particular, N. ashei can be distinguished from most eastern African N. nigricollis by the combination of high ventral scale and dorsal scale row counts. Most N. ashei have over 195 ventrals and at least 21 and typically more scale rows around the neck, whereas most N. nigricollis with 195 or more ventrals have at most 21, and usually 19 or fewer scale rows around the neck, whereas higher scale row counts around the neck tend to be found in specimens with fewer ventral scales. Naja ashei differs from the more closely related N. mossambica in lacking any dark edges on the labial scales and ventral scales, in having a less complex ventral banding pattern, and in having higher average ventral scale counts, but lower dorsal scale row counts. Naja pallida and N. nubiae differ in having higher midbody dorsal scale row counts (usually 25, compared to 21���23 in N. ashei). In addition, N. pallida differs from N. ashei in having a single, very clearly defined and clean-edged throat band (which very obviously crosses the neck except in older, darker specimens), in usually having higher ventral scale counts, and in the frequent presence of a single preocular and seven supralabials. Naja nubiae also has a cleaner, neater throat pattern, and two dark bands across the neck and two or three across the throat; a characteristic black tear-drop marking (consisting of dark edges to the supralabial suture below the eye) is almost invariably present; moreover, N. nubiae has almost consistently higher ventral scale counts, and often has seven supralabials and/or a single preocular (see W��ster & Broadley, 2003). Naja katiensis has consistently lower ventral and subcaudal scale counts (Table 4), a much smaller adult size, and lacks cuneate scales. Among the non-spitting cobras, N. ashei is most likely to be confused with N. haje, on account of its drab brownish coloration and large size. However, N. haje differs in having a single preocular, a row of suboculars separating the eyes from the supralabials, a greatly enlarged sixth supralabial, a single anterior temporal, and in lacking spitting adaptations to the fangs (Bogert, 1943), and thus being incapable of spitting venom. Naja melanoleuca similarly differs from N. ashei in having a single preocular, no suboculars, an enlarged sixth supralabial and a single anterior temporal. Description of holotype. Body dimensions: Snout-vent length 1268 mm, tail length 239 mm, dorsal head length (snout to end of parietal suture) 33.3 mm, lateral head length (snout to posterior end of lower jaw articulation) 51.7 mm. Head width across supraoculars 19.7 mm, maximum overall width of head 39.7 mm. Head broad, heart-shaped from above. Eye small to moderate, diameter much less than distance from mouth or from nostril. Body scalation: 197 ventrals, 55 subcaudals, all paired except for the first, the intact tail terminates in a spine. Dorsal scale rows: 23 on neck, 21 at midbody, 15 one head length ahead of vent. Scale rows at midbody Subcaudals - Females Dorsal scale row reduction formula: 25 5 + 6 (2) 24 7 + 8 (4) 23 4 + 5 (12 / 13) 21 4 + 5 / 5 + 6 (21) 19 + 6 (30 / 30) 21 4 + 5 / 5 + 6 (122) 19 4 + 5 / 5 + 6 (131) 17 4 + 5 (151 / 154) 15 3 + 4 (186) 14 + 4 (187) 15 4 + 5 (191 / 194) 13 + 3 (195 / 195) 15 Caudal scale reduction formula: 11 2 + 3 (2) 10 2 + 3 (3 / 3) 8 4 + 5 (5) 7 3 + 4 (6) 6 2 + 3 (16 / 17) 4 Head scalation: Preoculars 2 / 2, postoculars 2 / 2, supralabials 6 / 6, third enters eye, infralabials 8 / 9, first four contact anterior chin shields. On the left, infralabials 5 and 6 are homologous to the cuneate scales of Asiatic cobras (W��ster, 1998), whereas on the right hand side, infralabials five and seven are cuneates (Fig. 5). Anterior temporals 2 / 2, posterior temporals 5 / 5. Seven temporals and nuchals contact the lateral and posterior edges of the parietals. Rostral 1.5 times wider than high, visible from above. Posterior chin shields separated by two rows of smaller, elongate scales. Nasal scale entirely divided into a prenasal and a postnasal scale by the large, vertically elongate nostril. Frontal longer than wide (9.0 x 7.1 mm), slightly shorter than distance from rostral (10.3 mm), shorter than supraoculars (12.0 mm), widest along anterior edge; shape pentagonal, anterior edge straight, posterior edge ends in obtuse angle, border with supraoculars slightly concave. Colour and pattern in life: Head uniformly brownish olive on top, paler and greyer in supralabial region and around eye. Underside of head very finely dusted with brownish grey pigment, scale bases cream, overall impression light brownish grey. Dorsal colour generally olive-brown. Neck immediately posterior to head darker than top of head or the remainder of the dorsum. Otherwise, overall appearance largely uniform. Most dorsal scales with a slightly lighter lower basal edge. Interstitial skin mostly dark grey, with indistinct lighter variegations, visible especially when exposed by inflation of the body. Dorsal scales within lighter variegations have more pronounced light bases, giving an indistinct mottled appearance. Throat and ventral pattern (Fig. 5): first seven ventrals heavily mottled with greyish brown, scale bases creamy-white, light area sharply demarcated from darker pigment. Ventrals 8���10 similarly patterned, but with a slightly darker, more saturated brown pigment, covering 85���90 % of each scale except the base near the middle of the scale. Ventrals 11���13 as ventrals 1���7. Ventrals 14���20 almost entirely covered with pigment of intermediate density, with only a few lighter flecks on some scale bases. The remainder of the ventral and subcaudal scales are creamish with isolated blotches of greyish-brown pigment. Distal lateral tips of the ventrals also covered in greyish brown pigment, which forms a continuation of the colour of the lower dorsal scale rows. There are no dark scale bases or edges on the ventral surface. Variation for all material examined. Variation in scale counts in N. ashei and other African spitting cobras is given in Table 4. In addition to the characters listed there, N. ashei is notable for frequently having only two postocular scales, rather than three. Among the specimens included in our principal components analysis, eight out of fifteen N. ashei had two postoculars on at least one side, compared to one out of twentynine N. nigricollis. Variation in colour and pattern concerns especially the ventral pattern. The first ventrals may be largely light or more or less heavily suffused with dark pigment, but the transition from these to the main dark throat band normally remains distinguishable. Juveniles have a lighter dorsal ground colour, often with a faint ���herring-bone��� pattern, but the top and upper sides of the head and the neck are dark greyish brown (Fig. 6). The darker colour on the neck is more intense on the sides (where it merges into the dark throat band), and gradually merges into the dorsal body colour, without there being a clearly defined band. Size. This appears to be the largest spitting cobra, at least in terms of average size. Largest male examined (NMK / O 2401 ���Nguni, Kitui District, Kenya) 1750 + 360 = 2110 mm; largest female (NMK, unnumbered, ��� Kenya ���) 1800 + 350 = 2150 mm. However, giant specimens are generally underrepresented in collections. Specimens measuring 2 metres are not rare along the Kenyan coast, and a number of specimens of well over 2 metres in total length have been recorded. Pitman (1974) records males with total lengths of 2743 and 2311 mm from the Baringo region of Kenya, which are almost certainly referable to N. ashei. However, a record specimen measuring 2819 mm (Seronera, Serengeti National Park, Tanzania���Pitman, 1974) cannot confidently be attributed to N. ashei, as there are no records of the species from the park, and some northern Tanzanian N. nigricollis also reach very large sizes (W.W., pers. obs.). Etymology. We dedicate this species to the memory of the late James Ashe (1925���2004), in recognition of his contributions to East African herpetology, of the inspiration he gave to others working on the herpetofauna of this part of the world (see Spawls, 2004), of his early recognition of the distinctiveness of the species that now bears his name, and in gratitude for his support for this work. Distribution. Naja ashei appears to be sympatric with N. pallida over much of its range, i.e. dry lowland regions of northern and coastal Kenya, extending south along the coast to at least Diani Beach and north into southern Somalia and south-eastern Ethiopia. It occurs in northeast Uganda at Amudat in Karamoja District (BMNH 1954.1. 12.46, 1974.5145 ��� 7). It probably also occurs in the far north and/or northeast of Tanzania, but there appear to be no confirmed records. It should be looked for in the Serengeti National Park and the northernmost parts of the Tanzanian coast. The brown-headed and often very large spitting cobras from the Usambara Mountains and the central coastal region of Tanzania are referable to N. nigricollis, as demonstrated by our molecular analyses here. The northern and western distributional limits of N. ashei remain somewhat unclear. Some specimens of N. nigricollis from southern Sudan, northern Uganda and north-eastern Congo are also brownish above, but differ from N. ashei as highlighted in the diagnosis. However, the precise distributions of these forms require further investigation. The isolated population of spitting cobras assigned to N. nigricollis by W��ster & Broadley (2003), from Jebel Marra, Darfur Province, Sudan, where it occurs sympatrically with N. nubiae, also superficially resembles N. ashei due to its colour pattern, but clusters with N. nigricollis in our multivariate analyses. Further genetic studies are required to ascertain the status of this form. Medical relevance. As always, the discovery of a new species of venomous snake raises the question of whether existing antivenoms provide adequate protection (W��ster & McCarthy, 1996; Fry et al., 2003). The question is particularly relevant as large Naja ashei can secrete prodigious quantities of venom. A large specimen milked at the Bio-Ken snake farm in Watamu, Kenya, produced 6.2 ml of liquid venom, weighing 7.1 g (Fig. 7). Dry weight was not recorded, but if the ratio of 34.6���41.3 % solids by weight obtained by Mirtschin et al. (2006) from a selection of four species of Naja applies to N. ashei, then this suggests venom yields of up to 3 grams of dry venom, a record-breaking yield emphasising the potential danger of this species. Case histories have not been documented specifically for N. ashei, but bites by African spitting Naja typically result in severe necrosis (Warrell et al., 1976; Tilbury, 1982), but often limited systemic symptoms., Published as part of W��ster, Wolfgang & Broadley, Donald G., 2007, Get an eyeful of this: a new species of giant spitting cobra from eastern and north-eastern Africa (Squamata: Serpentes: Elapidae: Naja), pp. 51-68 in Zootaxa 1532 on pages 58-64, DOI: 10.5281/zenodo.177729, {"references":["Wuster, W. & Broadley, D. G. (2003) A new species of spitting cobra from north-eastern Africa (Serpentes: Elapidae: Naja). Journal of Zoology, London, 259, 345 - 359.","Bogert, C. M. (1943) Dentitional phenomena in cobras and other elapids, with notes on adaptive modification of fangs. Bulletin of the American Museum of Natural History, 81, 285 - 360.","Wuster, W. (1998) The cobras of the genus Naja in India. Hamadryad, 23, 15 - 32.","Pitman, C. R. S. (1974). A Guide to the Snakes of Uganda. Revised Edition. Wheldon & Wesley, Codicote, 290 pp.","Spawls, S. (2004) James Ashe: 1925 - 2004. African Journal of Herpetology, 53, 203 - 204.","Wuster, W. & McCarthy, C. J. (1996) Venomous snake systematics: implications for snakebite treatment and toxinology. In C. Bon & M. Goyffon (Eds.), Envenomings and their Treatments. Fondation Merieux, Lyon, pp. 13 - 23.","Fry, B. G., Winkel, K. D., Wickramaratna, J. C., Hodgson, W. C. & Wuster, W. (2003) Effectiveness of snake antivenom: species and regional venom variation and its clinical impact. Journal of Toxicology - Toxin Reviews, 22, 23 - 34.","Mirtschin, P. J., Dunstan, N., Hough, B., Hamilton, E., Klein, S., Lucas, J., Millar, D., Madaras, F. & Nias, T. (2006) Venom yields from Australian and some other species of snakes. Ecotoxicology, 15, 531 - 538","Warrell, D. A., B. M. Greenwood, N. McD. Davidson, L. D. Ormerod & C. R. M. Prentice (1976) Necrosis, haemorrhage and complement depletion following bites by the spitting cobra (Naja nigricollis). Quarterly Journal of Medicine, New Series, 45, 1 - 22.","Tilbury, C. R. (1982) Observations on the bite of the Mozambique spitting cobra (Naja mossambica mossambica). South African Medical Journal, 61, 308 - 313."]}

Naja ashei sp. nov. — Ashe’s spitting cobra Holotype. National Museums of Kenya NMK S/ 3993, a female specimen from Watamu, Kenya (3 º 21 ’S: 40 º 01’E), coll. Royjan Taylor, maintained in captivity at Bio-Ken Snake farm until 29 /09/ 2004 with reference number BK 10030 (Fig. 4,5). Paratypes (three males and two females): BMNH 1955.1.12.4a and 4 b (Kilifi, Kenya) BMNH 1963.456 (Kiboko, Kenya); BMNH 2005.1604 (Baringo, Kenya); NMZB 3349 (Ex USNM 40954) (Guaso Nyiro [=Ewaso Ng’iro], Kenya). Diagnosis. Naja ashei differs from all other African spitting cobras in possessing a unique clade of mtDNA haplotypes. From the data presented here, we identified 12 fixed nucleotide differences that differentiate N. ashei from the other eastern African spitting Naja. These correspond to positions 105, 169 and 315 of the ND 4 sequence of the holotype (DQ 897706), and to positions 60, 108, 153, 201, 348, 381, 507, 630 and 676 of the cytochrome b sequence of the same specimen (DQ 897749), the diagnostic bases at these positions being C, T, G, C, G, T, T, T, A, C, T and A, respectively. Morphologically, N. ashei differs from East African N. nigricollis in a number of characters relating to adult colour pattern and scalation. In particular, its midbody and posterior ventral colour is predominantly light, with dark pigment encroaching mostly from the sides of the body (venter normally largely or entirely dark in N. nigricollis), it lacks any red, orange or pink pigment under the throat (usually pronounced in N. nigricollis), and the head is the same olive-brown colour as the rest of the body (often black above and below in East African N. nigricollis). Scalation does not provide any absolutely diagnostic characters for N. ashei, but mean scale counts and the range differ clearly from those of East African N. nigricollis (Table 4). In particular, N. ashei can be distinguished from most eastern African N. nigricollis by the combination of high ventral scale and dorsal scale row counts. Most N. ashei have over 195 ventrals and at least 21 and typically more scale rows around the neck, whereas most N. nigricollis with 195 or more ventrals have at most 21, and usually 19 or fewer scale rows around the neck, whereas higher scale row counts around the neck tend to be found in specimens with fewer ventral scales. Naja ashei differs from the more closely related N. mossambica in lacking any dark edges on the labial scales and ventral scales, in having a less complex ventral banding pattern, and in having higher average ventral scale counts, but lower dorsal scale row counts. Naja pallida and N. nubiae differ in having higher midbody dorsal scale row counts (usually 25, compared to 21–23 in N. ashei). In addition, N. pallida differs from N. ashei in having a single, very clearly defined and clean-edged throat band (which very obviously crosses the neck except in older, darker specimens), in usually having higher ventral scale counts, and in the frequent presence of a single preocular and seven supralabials. Naja nubiae also has a cleaner, neater throat pattern, and two dark bands across the neck and two or three across the throat; a characteristic black tear-drop marking (consisting of dark edges to the supralabial suture below the eye) is almost invariably present; moreover, N. nubiae has almost consistently higher ventral scale counts, and often has seven supralabials and/or a single preocular (see Wüster & Broadley, 2003). Naja katiensis has consistently lower ventral and subcaudal scale counts (Table 4), a much smaller adult size, and lacks cuneate scales. Among the non-spitting cobras, N. ashei is most likely to be confused with N. haje, on account of its drab brownish coloration and large size. However, N. haje differs in having a single preocular, a row of suboculars separating the eyes from the supralabials, a greatly enlarged sixth supralabial, a single anterior temporal, and in lacking spitting adaptations to the fangs (Bogert, 1943), and thus being incapable of spitting venom. Naja melanoleuca similarly differs from N. ashei in having a single preocular, no suboculars, an enlarged sixth supralabial and a single anterior temporal. Description of holotype. Body dimensions: Snout-vent length 1268 mm, tail length 239 mm, dorsal head length (snout to end of parietal suture) 33.3 mm, lateral head length (snout to posterior end of lower jaw articulation) 51.7 mm. Head width across supraoculars 19.7 mm, maximum overall width of head 39.7 mm. Head broad, heart-shaped from above. Eye small to moderate, diameter much less than distance from mouth or from nostril. Body scalation: 197 ventrals, 55 subcaudals, all paired except for the first, the intact tail terminates in a spine. Dorsal scale rows: 23 on neck, 21 at midbody, 15 one head length ahead of vent. Scale rows at midbody Subcaudals - Females Dorsal scale row reduction formula: 25 5 + 6 (2) 24 7 + 8 (4) 23 4 + 5 (12 / 13) 21 4 + 5 / 5 + 6 (21) 19 + 6 (30 / 30) 21 4 + 5 / 5 + 6 (122) 19 4 + 5 / 5 + 6 (131) 17 4 + 5 (151 / 154) 15 3 + 4 (186) 14 + 4 (187) 15 4 + 5 (191 / 194) 13 + 3 (195 / 195) 15 Caudal scale reduction formula: 11 2 + 3 (2) 10 2 + 3 (3 / 3) 8 4 + 5 (5) 7 3 + 4 (6) 6 2 + 3 (16 / 17) 4 Head scalation: Preoculars 2 / 2, postoculars 2 / 2, supralabials 6 / 6, third enters eye, infralabials 8 / 9, first four contact anterior chin shields. On the left, infralabials 5 and 6 are homologous to the cuneate scales of Asiatic cobras (Wüster, 1998), whereas on the right hand side, infralabials five and seven are cuneates (Fig. 5). Anterior temporals 2 / 2, posterior temporals 5 / 5. Seven temporals and nuchals contact the lateral and posterior edges of the parietals. Rostral 1.5 times wider than high, visible from above. Posterior chin shields separated by two rows of smaller, elongate scales. Nasal scale entirely divided into a prenasal and a postnasal scale by the large, vertically elongate nostril. Frontal longer than wide (9.0 x 7.1 mm), slightly shorter than distance from rostral (10.3 mm), shorter than supraoculars (12.0 mm), widest along anterior edge; shape pentagonal, anterior edge straight, posterior edge ends in obtuse angle, border with supraoculars slightly concave. Colour and pattern in life: Head uniformly brownish olive on top, paler and greyer in supralabial region and around eye. Underside of head very finely dusted with brownish grey pigment, scale bases cream, overall impression light brownish grey. Dorsal colour generally olive-brown. Neck immediately posterior to head darker than top of head or the remainder of the dorsum. Otherwise, overall appearance largely uniform. Most dorsal scales with a slightly lighter lower basal edge. Interstitial skin mostly dark grey, with indistinct lighter variegations, visible especially when exposed by inflation of the body. Dorsal scales within lighter variegations have more pronounced light bases, giving an indistinct mottled appearance. Throat and ventral pattern (Fig. 5): first seven ventrals heavily mottled with greyish brown, scale bases creamy-white, light area sharply demarcated from darker pigment. Ventrals 8–10 similarly patterned, but with a slightly darker, more saturated brown pigment, covering 85–90 % of each scale except the base near the middle of the scale. Ventrals 11–13 as ventrals 1–7. Ventrals 14–20 almost entirely covered with pigment of intermediate density, with only a few lighter flecks on some scale bases. The remainder of the ventral and subcaudal scales are creamish with isolated blotches of greyish-brown pigment. Distal lateral tips of the ventrals also covered in greyish brown pigment, which forms a continuation of the colour of the lower dorsal scale rows. There are no dark scale bases or edges on the ventral surface. Variation for all material examined. Variation in scale counts in N. ashei and other African spitting cobras is given in Table 4. In addition to the characters listed there, N. ashei is notable for frequently having only two postocular scales, rather than three. Among the specimens included in our principal components analysis, eight out of fifteen N. ashei had two postoculars on at least one side, compared to one out of twentynine N. nigricollis. Variation in colour and pattern concerns especially the ventral pattern. The first ventrals may be largely light or more or less heavily suffused with dark pigment, but the transition from these to the main dark throat band normally remains distinguishable. Juveniles have a lighter dorsal ground colour, often with a faint “herring-bone” pattern, but the top and upper sides of the head and the neck are dark greyish brown (Fig. 6). The darker colour on the neck is more intense on the sides (where it merges into the dark throat band), and gradually merges into the dorsal body colour, without there being a clearly defined band. Size. This appears to be the largest spitting cobra, at least in terms of average size. Largest male examined (NMK / O 2401 —Nguni, Kitui District, Kenya) 1750 + 360 = 2110 mm; largest female (NMK, unnumbered, “ Kenya ”) 1800 + 350 = 2150 mm. However, giant specimens are generally underrepresented in collections. Specimens measuring 2 metres are not rare along the Kenyan coast, and a number of specimens of well over 2 metres in total length have been recorded. Pitman (1974) records males with total lengths of 2743 and 2311 mm from the Baringo region of Kenya, which are almost certainly referable to N. ashei. However, a record specimen measuring 2819 mm (Seronera, Serengeti National Park, Tanzania—Pitman, 1974) cannot confidently be attributed to N. ashei, as there are no records of the species from the park, and some northern Tanzanian N. nigricollis also reach very large sizes (W.W., pers. obs.). Etymology. We dedicate this species to the memory of the late James Ashe (1925–2004), in recognition of his contributions to East African herpetology, of the inspiration he gave to others working on the herpetofauna of this part of the world (see Spawls, 2004), of his early recognition of the distinctiveness of the species that now bears his name, and in gratitude for his support for this work. Distribution. Naja ashei appears to be sympatric with N. pallida over much of its range, i.e. dry lowland regions of northern and coastal Kenya, extending south along the coast to at least Diani Beach and north into southern Somalia and south-eastern Ethiopia. It occurs in northeast Uganda at Amudat in Karamoja District (BMNH 1954.1. 12.46, 1974.5145 – 7). It probably also occurs in the far north and/or northeast of Tanzania, but there appear to be no confirmed records. It should be looked for in the Serengeti National Park and the northernmost parts of the Tanzanian coast. The brown-headed and often very large spitting cobras from the Usambara Mountains and the central coastal region of Tanzania are referable to N. nigricollis, as demonstrated by our molecular analyses here. The northern and western distributional limits of N. ashei remain somewhat unclear. Some specimens of N. nigricollis from southern Sudan, northern Uganda and north-eastern Congo are also brownish above, but differ from N. ashei as highlighted in the diagnosis. However, the precise distributions of these forms require further investigation. The isolated population of spitting cobras assigned to N. nigricollis by Wüster & Broadley (2003), from Jebel Marra, Darfur Province, Sudan, where it occurs sympatrically with N. nubiae, also superficially resembles N. ashei due to its colour pattern, but clusters with N. nigricollis in our multivariate analyses. Further genetic studies are required to ascertain the status of this form. Medical relevance. As always, the discovery of a new species of venomous snake raises the question of whether existing antivenoms provide adequate protection (Wüster & McCarthy, 1996; Fry et al., 2003). The question is particularly relevant as large Naja ashei can secrete prodigious quantities of venom. A large specimen milked at the Bio-Ken snake farm in Watamu, Kenya, produced 6.2 ml of liquid venom, weighing 7.1 g (Fig. 7). Dry weight was not recorded, but if the ratio of 34.6–41.3 % solids by weight obtained by Mirtschin et al. (2006) from a selection of four species of Naja applies to N. ashei, then this suggests venom yields of up to 3 grams of dry venom, a record-breaking yield emphasising the potential danger of this species. Case histories have not been documented specifically for N. ashei, but bites by African spitting Naja typically result in severe necrosis (Warrell et al., 1976; Tilbury, 1982), but often limited systemic symptoms.