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Based on molecular, morphological and field data, the status and zoogeography of the taxa of the Psammophissibilans group north of 12°S are reviewed. Molecular data including sequences from 20 of the 22 described species known to occur north of 12°S suggest that P. sibilans distribution is restricted to northeastern Africa, from Egypt to Ethiopia. Po-pulations from West Africa are described as a new species, P. afroccidentalis sp. nov., and those from Chad, Cameroon and Central African Republic are assigned to P. rukwae which is also distributed from Tanzania to Ethiopia. Molecular data indicate the occurrence within this complex of three additional cryptic species in the Horn of Africa. Populations previous-ly assigned to P. phillipsi in Central Africa north, south and east of the Congo forest block are assigned to P. mossambicus and the status of P. occidentalis is discussed. P. phillipsi is restricted to West Africa, with P. irregularis as junior synonym.

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).

Key to the species of Pyxicephalus 1.a. Odontoids wider than long.......................................................... P. angusticeps New Status 1 b. Odontoids longer than wide............................................................................. 2 2.a. Upper jaw free of barring in adults.............................................................. P. adspersus 2 b. Upper jaw barred in adults.............................................................................. 3 3.a. Tympanum hypertrophied (width greater than eye diameter), and positioned less than one half eye diameter in distance from eye; toes with moderate webbing................................................................. P. obbianus 3 b. Tympanum not hypertrophied (width less than or equal to eye diameter), and positioned more than one half eye diameter in distance from eye; toes with rudimentary webbing..................................................... P. edulis, Published as part of Scott, Elizabeth, Visser, John D., Yetman, Caroline A., Oliver, Lauren & Broadley, Donald G., 2013, Revalidation of Pyxicephalus angusticeps Parry, 1982 (Anura: Natatanura: Pyxicephalidae), a bullfrog endemic to the lowlands of eastern Africa, pp. 201-228 in Zootaxa 3599 (3) on page 204, DOI: 10.11646/zootaxa.3599.3.1, http://zenodo.org/record/217094

Scott, Elizabeth, Visser, John D., Yetman, Caroline A., Oliver, Lauren, Broadley, Donald G. (2013): Revalidation of Pyxicephalus angusticeps Parry, 1982 (Anura: Natatanura: Pyxicephalidae), a bullfrog endemic to the lowlands of eastern Africa. Zootaxa 3599 (3): 201-228, DOI: http://dx.doi.org/10.11646/zootaxa.3599.3.1

Pyxicephalus angusticeps Parry, 1982 New Status Figs 1���4, 6, 10 G���H, 11 A���C, 12 B, 13 E. Pyxicephalus adspersus angusticeps Parry 1982: 281 ���292, figs 1, 4, 5, 7, table 1. Poynton & Broadley 1985: 123 ���124. Pyxicephalus edulis: Channing, Du Preez & Passmore 1994: 141 ���148, fig. 1 b, plate 1 b, synonymized P. adspersus angusticeps. Channing 2001: 349 ���351, figs 22.5, 281. Channing & Howell 2006: 322 ���323, fig. 22.6. Pyxicephalus edulis: Pickersgill 2007: 105 ���108 (part), figs 45, 46. Type material. Holotype: Adult 3, NMSA 1992 (UNP 3099, NMSA Type number 2581), from Beira, Mozambique (Fig. 2). Paratypes: One adult 3, NMSA 1991 (UNP 3098), and two subadults, NMSA 1990 (UNP 3063), NMSA 1993 (UNP 3100) with same data as the holotype; twelve subadult paratypes NMZB-UM 6451 (two specimens), NMZB-UM 7516 (five specimens), NMZB-UM 19774 (four specimens; no. A shown in Figs 11 A���C) and NMZB-UM 23374, from Beira, Mozambique. Diagnosis. This species is assigned to the genus Pyxicephalus by the combination of presence of mandibular odontoids (outgrowths of the dentary), femoral glands, cranial exostosis on the frontoparietals and nasals, hypertrophied inner metatarsal tubercle, and supraorbital flanges on the frontoparietals (see Scott 2005 for comparisons of these characters in other genera of ranoid frogs). Pyxicephalus angusticeps can be distinguished from P. obbianus (Figs 8, 10 C���D, 11 D���F) by the: (1) rudimentary webbing between the toes (more extensive in P. obbianus; Fig. 8 F); (2) tympanum width less than eye diameter (more than one and one half times eye diameter in adult male P. obbianus; Figs 8 A, 8 C), positioned approximately one tympanum width or slightly less from eye (less than half the tympanum width in P. obbianus; Fig. 8 C); and (3) vertebral stripe running most of the length of body (restricted to the snout in adult P. obbianus; Figs 8 A, 8 D). Pyxicephalus angusticeps can be distinguished from P. edulis (Figs 5, 7, 10 E���F, 12 C���D) and P. adspersus (Figs 9, 10 A���B, 12 A) by: (1) nuptial pads present only on innermost finger (Digit II) of breeding males (as in P. obbianus, but present on Digits II���IV in P. adspersus and P. e d u l i s); (2) head of breeding males less proportionately enlarged (see Fig. 10), particularly in width, than in P. adspersus and P. e d u l i s; (3) distinct subadult gular colouration (Fig. 11 C), which is darkly marbled and more persistent in the adult female (Fig. 3 F) than in P. adspersus and P. e d u l i s; (4) small, triangular, pointed odontoids, not longer than wide (Fig. 12 B in P. angusticeps vs. Figs 12 C���D in P. e d u l i s; Fig. 8 D in P. obbianus), whereas odontoids are large and well-developed in all other species of Pyxicephalus; (5) presence of at most poorly-developed dorsal ridges, usually the most distinct running from behind eyes (dorsum of type material covered only by weakly or strongly developed oval or round warts), whereas these are thick and highly developed in P. adspersus (Fig. 10 A); and narrower, finer and often continuous in P. e d u l i s (Figs 5 A, 10 E); (6) vertebral stripe wide, pale (coppery-orange in preservation) and diffuse, formed by pale background colour visible between densely-packed, dark dorsal blotches, which are absent medially, with fine, dark stippling present elsewhere between dark dorsal blotches (Figs 3, 4), whereas the vertebral stripe is absent in adults of P. adspersus, and variably (but usually) present, thin and pale green-yellow in P. e d u l i s (Figs 7 A, 10); (7) femoral glands well-developed and relatively larger and more conspicuous than in all other species of Pyxicephalus. Redescription of the holotype. Adult male, NMSA 1992 (field number UNP 3099, NMSA Type Number 2581, Fig. 2), collected at Beira, the swamp beyond Estoril [19 �� 50 ' 37 "S, 34 �� 50 ' 20 "E]: Jan. 26, 1959, Nyasaland- Mozambique Expedition [of the University of Natal, Pietermaritzburg]. Head. Tympanum visible, subcircular (oval) in shape, slanted slightly anteriorly. Tympanum width smaller than that of eye. Tympanum positioned approximately its own width in distance from eye (i.e. less than one times width of eye away from eye). White spot or marking absent from right tympanum, but minute white spot present on left tympanum (Fig. 2 C). Upper lip with thin rim of light colouration. Upper jaw with four pale coppery-orange brown bars on each side; posterior two broader than anterior two. Two anterior-most bars on upper jaw on each side of face meet anterior to eyes. Gular region pale with diffuse brown colouration laterally. Light irregular spots (not a complete crossbar) across head above eyes. Head narrow, not as wide as body at level of sacrum, not disproportionately wide. Odontoids in lower jaw weakly-developed, triangular. Tongue notched, typically ranid, no mid-lingual papilla. Nostrils large, situated slightly closer to eyes than tip of snout. Secondary sexual characteristics. Nuptial pads visible only on Digit II (inner finger) of hand, covering most of dorsal and lateral surfaces of these digits, including side of thenar tubercle. Forearms only slightly thickened, indicating youth. Testes large, elongated, dark yellow in preservation with blackened mesenteric tissue above. Limbs. A gracile, slender-bodied animal. Toes slightly webbed, web extending to half length of longest toe (approximately to level of proximal subarticular tubercle), such that three phalanges of longest toe free of webbing; remaining length of toes with slight paler lateral flanges. Inner metatarsal tubercle hypertrophied, spade-like; outer metatarsal tubercle absent. Digits of hand elongated, tapering, with slight paler lateral flanges. Ventral surface of hands lighter, dorsal side brown. Digit II (inner finger) longer than Digit III (second finger) and Digit V (fourth finger), Digit IV (third finger) longest. Subdigital tubercles present, large and rounded. Subarticular tubercles present on Digit IV, inconspicuous on other fingers. Thenar tubercle large and bilobed (not divided completely), palmar tubercle present. Palms of hands otherwise smooth. Digit tips unexpanded, rather narrowing distally, with a paler hardened tip. Dorsal and dorso-lateral surfaces of thighs, calves and along margin of tarsus with small white asperites, arranged loosely into lines on weak skin ridges on dorsal surfaces of calves. White asperites present on posterior third of dorsum, set on small raised tubercles, most densely arranged around cloaca. White asperites absent from ventral surfaces of limbs. Femoral granules present around cloaca, and on posterior-dorsal surface to half length of thighs. Ventral colouration. Colouration of dorsal surface of calves extending onto ventral surface of calves. Flanks and underside of limbs lightly stippled with diffuse brown pigment. Abdomen off-white, granular. Pectoral region with two medially directed triangular ���waistcoat��� scars 1 on each side, interior of which is coloured with diffuse light brown stippling. Glands. Pectoral glands present, small, elongate, triangular and inconspicuous. Femoral glands large, elongate, oval, brown, inconspicuous, set within pigment horizon of ventral thighs, which is indistinct, grading into ventral colouration. Femoral glands positioned closer to knee than cloaca, confined to distal two-fifths of length of thighs. 1. These triangular ���waistcoat��� patterns are commonly observed in Pyxicephalus, and may be relictual scars from the emergence of forelimbs. Dorsal colouration. Dorsum dark brownish-grey in preservation. Small rounded darker dorsal blotches visible anteriorly, indistinct posteriorly. No symmetrically arranged (opposite or alternate) blotches visible over urostyle region. Vertebral stripe broad, coppery-orange brown, the edges formed by irregularly-positioned, dark rounded dorsal blotches; vertebral stripe extending to tip of snout, but narrowing anteriorly from approximate level of eyes, and indistinct from approximate level of sacrum posteriorly. Small patches of mottling and stippling apparent anteriorly against pale base colour of dorsum, becoming less conspicuous posteriorly. Dorsal ridges and warts. Primary Dorsal Ridge Series 2 broken and only partially visible on right side, not extending full length of dorsum. Primary Dorsal Ridge Series 1 and 3 are greatly reduced to few weakly-developed warts. Lateral ridge absent on left, weakly present on right. Numerous conspicuous but narrow, raised oval warts present on flanks. Dorsum otherwise relatively smooth, with few slightly raised warts. Weak supratympanic ridge of warts apparent. Variation. Adult material from the coastal lowlands of Kenya and Tanzania (Fig. 4, Appendix 1) differs from typical Mozambican material of P. angusticeps (Figs 2, 3) in the following respects. The vertebral stripe is narrower, better defined and a darker orange-red colour in preservation. The dorsum may present more, but poorlydefined and wider broken ridges, and have more distinct dark dorsal blotches. Barring is occasionally absent between the upper lip and the eye. Adults are larger than those from Beira. The asperites on the dorsal surface of the tibia are more irregularly arranged. Metamorphs and subadults. The metamorphs of P. angusticeps can be distinguished from both P. adspersus and P. edulis on at least three external characters. The adult pattern of fewer ridges and conspicuous oval warts is also evident in metamorphs and subadults of P. angusticeps (Figs 11 A���C). There are only a few broken ridges in place of Primary Dorsal Ridge Series 2, and the dorsum is covered in conspicuous oval warts, usually more pronounced laterally, in metamorph material (with tail vestiges) from NMZB that we assigned to P. angusticeps based on occurrence at the same locality as the distinctive subadult paratypes. These evenly-spaced warts in metamorphs and sub-adults of P. angusticeps are absent from the dorsum in metamorphs of P. adspersus (Figs 11 G���I), P. e d u l i s and P. obbianus (Figs 11 D���F), and conform in appearance to those observed in the warty adult individual illustrated by Channing (2001, fig. 22.5), Channing et al. (1994, plate 1 b), and Channing & Howell (2006, plate 22.6). They resemble dorsal warts observed in the pyxicephaline genus Poyntonia Channing & Boycott, 1989. The metamorphs of P. obbianus also differ markedly in colouration from their congeners, being finely vermiculated over the entire dorsum, with larger blotches present only on the limbs, and in having immaculate abdomens. A primary series of six ridges develop in late-stage P. adspersus and P. edulis larvae, and are pronounced by the time metamorphosis is complete (Figs 11 G���I); these ridges are lacking in metamorphs of P. obbianus (Figs 11 D���F). Larvae. Tadpoles listed in the account of Pickersgill (2007 fig. 47, top, viz. MP 2155) from Tica, Mozambique, are typical of P. e d u l i s (according to information presented in Du Preez & Carruthers 2009: 414���417), in having the labial tooth row formula 5 (3-5)/ 3. The tadpole of P. angusticeps is unknown. Distribution. Parry (1982) suggested that P. angusticeps was probably isolated from the western populations of P. edulis by the old course of the Zambezi River, which reached the sea south of Beira. He predicted that the range of P. angusticeps would include the area north to the Zambezi River and west to the Urema Trough. Although the southern limit of the known distribution of P. angusticeps appears to be the type locality, Beira in Mozambique, the species probably extends northwards through the lowland plains of East Africa, as far as Kakuyuni in Kenya (Appendix 1). A distribution map of P. angusticeps in East Africa is presented in Fig. 1, and is similar to that presented for P. edulis in fig. 252 of Channing & Howell (2006). There is, however, a substantial disjunction between known records from Mozambique and those from Tanzania and Kenya, which may be due to poor collecting effort in the intervening lowlands. Few herpetological surveys have been undertaken in Mozambique, and even those conducted in areas where Pyxicephalus should occur (e.g. Jacobsen et al. 2010) may not detect these frogs, due to their prolonged periods of inactivity. We have used the distinctive characteristics of the metamorphs (discussed above) to assign some metamorphs without associated adult material from two additional localities to P. angusticeps. Jorge (1933 DD) is on the Buzi River, about 120 km upstream from its confluence with the Pungwe River estuary. Alvez de Lima (2034 AB) was a safari camp ca. 50 km south-west of Beira, far from any major rivers. According to unpublished ecological modeling results (C.A. Yetman), areas in northern Mozambique with potentially suitable habitat for P. angusticeps include parts of north-central Manica Province, south-west of Quelimane on the coast in Zambezia Province, and along the Limpopo River in central Gaza Province. Conservation status. Additional survey work is required to accurately assess the range and conservation status of P. angusticeps, which may be more widespread in East Africa. Currently, it is best considered as ���Data Deficient��� (IUCN 2011)., Published as part of Scott, Elizabeth, Visser, John D., Yetman, Caroline A., Oliver, Lauren & Broadley, Donald G., 2013, Revalidation of Pyxicephalus angusticeps Parry, 1982 (Anura: Natatanura: Pyxicephalidae), a bullfrog endemic to the lowlands of eastern Africa, pp. 201-228 in Zootaxa 3599 (3) on pages 204-217, DOI: 10.11646/zootaxa.3599.3.1, http://zenodo.org/record/217094, {"references":["Parry, C. R. (1982) A revision of southern African Pyxicephalus Tschudi (Anura: Ranidae). Annals of the Natal Museum, 25 (1), 281 - 292.","Poynton, J. C. & Broadley, D. G. (1985) Amphibia Zambesiaca 2. Ranidae. Annals of the Natal Museum, 27 (1), 115 - 181.","Channing, A., Du Preez, L. H. & Passmore, N. I. (1994) Status, vocalization and breeding biology of two species of African bullfrogs (Ranidae: Pyxicephalus). Journal of Zoology, London, 234, 141 - 148. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1994. tb 06061. x","Channing, A. (2001) Amphibians of Central and Southern Africa. Protea Book House, Pretoria, 470 pp.","Channing, A. & Howell, K. M. (2006) Amphibians of East Africa. Comstock Publishing Associates, Cornell University Press, Ithaca, NY, 418 pp.","Pickersgill, M. (2007) Frog Search. Results of Expeditions to Southern and Eastern Africa from 1993 - 1999. Frankfurt Contributions to Natural History Volume 28. Edition Chimaira, Frankfurt am Main, 574 pp.","Scott, E. (2005) A phylogeny of ranid frogs (Anura: Ranoidea: Ranidae), based on a simultaneous analysis of morphological and molecular data. Cladistics, 21 (6), 507 - 574. http: // dx. doi. org / 10.1111 / j. 1096 - 0031.2005.00079. x","Du Preez, L. & Carruthers, V. C. (2009) A Complete Guide to the Frogs of Southern Africa. Struik Publishers, Cape Town, 488 pp. + CD.","Jacobsen, N. H. G., Pietersen, E. W. & Pietersen, D. W. (2010). A preliminary herpetological survey of the Vilanculos Coastal Wildlife Sanctuary on the San Sebastian Peninsula, Vilankulo, Mozambique. Herpetology Notes 3, 181 - 193 (published online on 31 May 2010)."]}

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."]}

Subgenus Boulengerina Dollo, 1886 Limnonaja Schmidt, 1923: 124 (type species Boulengerina christyi Boulenger, 1904) Paranaja Loveridge, 1944: 231 (type species Naja anomala Sternfeld = Naia multifasciata Werner, 1902) Type species: Boulengerina stormsi Dollo, 1886 (= Naja annulata Buchholz & Peters in Peters, 1876), by monotypy. Gender: feminine. Etymology: dedicated to George Albert Boulenger, famous Belgian herpetologist, and curator of reptiles at the British Museum (Natural History) in London from 1881 to 1920. Distribution: forested regions of western, central, eastern and south-eastern Africa. Content: four species: Naja (Boulengerina) annulata Buchholz & Peters in Peters, 1876: 119 Naja (Boulengerina) christyi (Boulenger, 1904: 14) Naja (Boulengerina) melanoleuca Hallowell, 1857: 61 Naja (Boulengerina) multifasciata Werner, 1902: 347 Diagnosis: Intracranial (dorsal) anterior Vidian canal position, 2–4 solid maxillary teeth, penultimate (sixth) supralabial high, combination of one preocular and one anterior temporal (except N. christyi, which sometimes has two anterior temporals), rostral much broader than deep, internasals shorter than prefrontals, dorsal scales highly polished, fangs not modified for spitting. Comments: This subgenus is morphologically and ecologically highly diverse, including species at both the large (Naja melanoleuca) and small (Naja multifasciata) extremes of the size spectrum of the cobra clade, and semi-fossorial, terrestrial and aquatic forms. However, they are united by their restriction to forest and forest edge habitats, and their distribution is centred on the central African forests, with only N. melanoleuca extending significantly into West and East Africa., 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 31, DOI: 10.5281/zenodo.190422, {"references":["Dollo, L. (1886) Note sur les reptiles et batraciens recueillis par M. le Capitaine Em. Storms dans la region du Lac Tanganyika. Bulletin du Musee Royal d'Histoire Naturelle de Belgique, 4, 151 - 160.","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, with field notes by Herbert Lang and James P. Chapin. Bulletin of the American Museum of Natural History, 49, 1 - 146.","Boulenger, G. A. (1904) Descriptions of two new elapine snakes from the Congo. Annals and Magazine of Natural History, 7, 453 - 456.","Loveridge, A. (1944) Further revisions of African snake genera. Bulletin of the Museum of Comparative Zoology, 95 (2), 121 - 247.","Werner, F. (1902) Uber westafrikanische Reptilien. Verhandlungen der Zoologischen und Botanischen Gesellschaft in Wien, 52, 332 - 348.","Peters, W. K. H. (1876) Eine zweite Mittheilung uber die von Hrn. Professor D. R. Buchholz in Westafrika gesammelten Amphibien. Monatsberichte der Koniglichen Preussischen Akademie der Wissenschaften zu Berlin, 1875, 117 - 123.","Hallowell, E. (1857) Notice on a collection of reptiles from the Gaboon country, West Africa, recently presented to the Academy of Natural Sciences in Philadelphia by Dr. Henry A. Ford. Proceedings of the Academy of Natural Sciences of Philadelphia, 9, 48 - 72."]}

Subgenus Uraeus Wagler, 1830 Type species: Coluber haje Linnaeus, 1758, by monotypy. Gender: masculine. Etymology: derived from the Egyptian uraeus, a symbol of the goddess Wadjet in the shape of a cobra with a spread hood, which formed part of the head-dress of Egyptian divinities and kings. Distribution: open formations in most of Africa and southern Arabia. Content: six species: Naja (Uraeus) anchietae Bocage, 1879: 89 Naja (Uraeus) annulifera Peters, 1854: 624 Naja (Uraeus) arabica Scortecci, 1932: 47 Naja (Uraeus) senegalensis Trape, Chirio & Wüster in Trape et al, 2009: xxx Naja (Uraeus) haje (Linnaeus, 1758: 225) Naja (Uraeus) nivea (Linnaeus, 1758: 223) Diagnosis: Intracranial (dorsal) anterior Vidian canal position, almost always 2–3 solid maxillary teeth (one specimen out of 21 examined by Bogert [1943] and Szyndlar & Rage [1990] had one solid maxillary tooth on each side), seven supralabials with penultimate (sixth) shield high, combination of one preocular and one anterior temporal, rostral as broad as deep, internasals as long as prefrontals, dorsal scales matt or moderately shiny, and fangs not adapted for spitting. Comments: This is a morphologically relatively conserved lineage that occupies open formations in much of Africa and the southern Arabian Peninsula. The most basal cladogenic split separates Naja (Uraeus) nivea, a species largely restricted to southern African areas with winter rainfall, from the remaining species, which occupy primarily tropical or subtropical formations (Wüster et al., 2007)., 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 31, DOI: 10.5281/zenodo.190422, {"references":["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.","Bocage, J. V. B. Du (1879) Subsidios para a fauna das possessoes portuguezas d'Africa occidental. Jornal de sciencias mathematicas, physicas, e naturaes da Academia Real das Sciencas de Lisboa, 7, 85 - 96.","Scortecci, G. (1932) Rettili dello Yemen. Atti della Societa Italiana di Scienze Naturali, 71, 39 - 49.","Trape, J. - F., Chirio, L., Broadley, D. G. & Wuster, W. (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. Zootaxa, in press.","Bogert, C. M. (1943) Dentitional phenomena in cobras and other elapids with notes on adaptive modifications of fangs. Bulletin of the American Museum of Natural History, 81, 285 - 360.","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."]}

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. (2003) A new species of spitting cobra from northeastern Africa (Serpentes: Elapidae: Naja). Journal of Zoology, London, 259, 345 - 359.","Trape, J. F. & Mane, Y. (2006) Guide des Serpents d'Afrique Occidentale. Savane et Desert. Paris, IRD Editions, 226 pp.","Bons, J. & Geniez, P. (1996) Amphibiens et Reptiles du Maroc (Sahara Occidental compris). Atlas biogeographique. Asociacion Herpetologica Espanola, Barcelona, 320 pp.","Geniez, P., Mateo, J. A., Geniez M., & Pether, J. (2004) The Amphibians and Reptiles of Western Sahara. Chimaira, Frankfurt am Main, 229 pp.","Dobiey, M. & Vogel, G. (2007) Venomous Snakes of Africa. Chimaira, Frankfurt a. M.","Schleich, H. H., Kastle, W. & K. Kabisch. (1996) Amphibians and Reptiles of North Africa. Koeltz Scientific Books, Koenigstein, 630 pp.","Renker, G. (1966) Verkanntes Schlangenvolk. Adolf Sponholtz Verlag, Hameln, 160 pp.","Kramer, E. & Schnurrenberger, H. (1963) Systematik, Verbreitung und Okologie der Libyschen Schlangen. Revue Suisse de Zoologie 70, 453 - 568.","Saleh, M. A. (1997) Amphibians and reptiles of Egypt. National Biodiversity Unit, Cairo, 234 pp.","Baha El Din, S. (2006) A Guide to the Reptiles and Amphibians of Egypt. The American University in Cairo Press, Cairo, 359 pp.","Chirio, L. & LeBreton, M. (2007) Atlas des Reptiles du Cameroun. Museum National d'Histoire Naturelle, IRD, Paris, 688 pp.","Gasperetti, J. (1988) Snakes of Arabia. Fauna of Saudi Arabia, 9, 169 - 450.","Egan, D. (2007) Snakes of Arabia. A Field Guide to the Snakes of the Arabian Peninsula and its Shores. Motivate Publishing, Dubai.","van der Kooij, J. (2001) The herpetofauna of the Sultanate of Oman. Part 4: the terrestrial snakes. Podarcis, 2, 54 - 64.","Roman, B. (1973) Viperides et Elapides de Haute-Volta. Notes et Documents Voltaiques, 6, 1 - 49.","Roman, B. (1980) Serpents de Haute-Volta. C. N. R. S. T., Ouagadougou, 312 pp.","Chirio, L. & Ineich, I. (2006) Biogeography of the reptiles of the Central African Republic. African Journal of Herpetology, 55, 23 - 59.","Cerling, T. E., Harris, J. M., MacFadden, B. J., Leakey, M. G., Quade, J., Eisenmann, V. & Ehleringer, J. R. (2006) Global vegetation change through the Miocene / Pliocene boundary. Nature, 389, 153 - 158.","Kelly, C. M. R., Barker, N. P., Villet, M. H., Broadley, D. G. & Branch, W. R. (2008) The snake family Psammophiidae (Reptilia: Serpentes): Phylogenetics and species delimitation in the African sand snakes (Psammophis Boie, 1825) and allied genera. Molecular Phylogenetics and Evolution, 47, 1045 - 1060.","Bosworth, W., Huchon, P., McClay, K. (2005) The Red Sea and Gulf of Aden Basins. Journal of African Earth Science, 43, 334 - 378.","Fernandes, C. A., Rohling, E. J. & Siddall, M. (2006) Absence of post-Miocene Red Sea land bridges: biogeographic implications. Journal of Biogeography, 33, 961 - 966.","Amer, S. A. M. & Kumazawa, Y. (2005) Mitochondrial DNA sequences of the Afro-Arabian spiny-tailed lizards (genus Uromastyx; family Agamidae): phylogenetic analyses and evolution of gene arrangements. Biological Journal of the Linnean Society, 85, 247 - 260.","Winney, B. J., Hammond, R. L., Macasero, W., Flores, B., Boug, A., Biquand, V. & Bruford, M. W. (2004) Crossing the Red Sea: phylogeography of the hamadryas baboon, Papio hamadryas hamadryas. Molecular Ecology, 13, 2819 - 2827."]}

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

Key to the species of Megatyphlops in eastern and southern Africa 1a. Rostral circular in dorsal view and not strongly projecting; short nasal suture arising from the rostral; dorsum light red-brown, pigment confined to posterior half of each scale M. anomalus 1b. Rostral unguiform, oval or cuneiform in dorsal view, strongly projecting; long nasal suture arising from first or second supralabial; dorsum lineolate, blotched, mottled or uniform black or grey-brown............................................ 2 2a. Rostral unguiform in dorsal view; midbody scale rows usually 34–44; dorsum mottled with black or entirely black ................................................................................................................................................................. M. schlegelii 2b. Rostral oval or cuneiform in dorsal view; midbody scale rows usually 30–36; dorsum lineolate, blotched with black or uniform grey-brown...................................................................................................................................... 3 3a. Rostral oval in dorsal view, convex in profile; rostral length/width ratio usually less than 1.25; lineolate or blotched, some large adults uniform grey-brown..................................................................................... M. mucruso 3b. Rostral cuneiform in dorsal view, depressed in profile; rostral length/width ratio usually more than 1.25; juveniles lineolate, adults uniform grey-brown ............................................................................................................ M. brevis, Published as part of Broadley, Donald G. & Wallach, Van, 2009, 2255, pp. 1-100 in Zootaxa 2255 on page 77

Key to the species of Afrotyphlops in eastern and southern Africa (note: due to variation in most diagnostic characters, some species appear twice in this key) 1a. A subocular scale usually separates preocular from second and third labials............................................................ 2 1b. No subocular scale separating preocular from second and third labials..................................................................... 3 2a. A blue-grey lineolate dorsal pattern, with or without superimposed black blotches; midbody scale rows 26–28; middorsals 404–463; Usambara Mt ............................................................................................................. A. gierrai 2b. Dorsum uniform black or blotched black and white; midbody scale rows 30–34; middorsals 464–542; Udzungwa, Ukaguru, and Uluguru Mts. ............................................................................................................... A. nigrocandidus 3a. Middorsals less than 310; less than 200 mm in total length....................................................................................... 4 3b. Middorsals more than 310; usually more than 200 mm in total length...................................................................... 5 4a. Midbody scale rows 22–26; middorsals 232–286; uniform grey-brown to black above and below, yellow patches below head and posterior venter .............................................................................................................. A. fornasinii 4b. Midbody scale rows 30; middorsals 287–291; dorsum lineolate, venter immaculate...................... A. nanus sp. nov. 5a. Preocular not contacting supralabials; dorsal scales in 28 rows throughout (no reductions caudally); inferior nasal suture arising from between first and second infralabials ....................................................................... A. kaimosae 5b. Preocular contacting supralabials; dorsal scale rows with some reductions caudally; inferior nasal suture arising from first supralabial and/or rostral........................................................................................................................... 6 6a. Supraocular with lateral apex between ocular and preocular..................................................................................... 7 6b. Supraocular with lateral apex between preocular and nasal.................................................................................... 12 7a. Ocular overlapped by second supralabial (T-0) ........................................................................................... A. bibronii 7b. Ocular not overlapped by second supralabial (T-X).................................................................................................. 8 8a. Preocular usually overlapped by second supralabial................................................................................................. 9 8b. Preocular not overlapped by second supralabial...................................................................................................... 11 9a. Eye beneath ocular................................................................................................................................................... 10 9b. Eye beneath preocular or ocular/preocular border .................................................................................. A. angolensis 10a. Midbody scale rows 28–32; eye beneath ocular/preocular border; Ethiopian highlands ......................... A. blanfordii 10b. Midbody scale rows 26–28; eye beneath preocular; Congo Basin forests............................................. A. steinhausi 11a. Dorsum heavily blotched with black, venter immaculate; midbody scale rows 26–30; supraoculars transverse....... .................................................................................................................................................................. A. congestus 11b. Dorsum with lineate or punctate pattern, venter blotched or spotted like dorsum; midbody scale rows 30–34; supraoculars oblique ................................................................................................................................. A. punctatus 12a. Preocular overlapped by second supralabial............................................................................................................. 13 12b. Preocular not overlapped by second supralabial...................................................................................................... 15 13a. Midbody scale rows 30–34; supraocular wedged between ocular and preocular; venter pigmented like dorsum...... .................................................................................................................................................................. A. punctatus 13b. Midbody scale rows 22–26; supraocular wedged between preocular and nasal; venter immaculate..................... 14 14a. Supraocular larger than frontal.............................................................................................................. A. rondoensis 14b. Frontal larger than supraocular ................................................................................................................... A. schmidti 15a. Ocular overlapped by second supralabial (T-II)....................................................................................................... 16 15b. Ocular not overlapped by second supralabial (T-X)................................................................................................ 17 16a. Midbody scale rows 26–32 [eye beneath ocular/preocular border; supraocular transverse] .................... A. lineolatus 16b. Midbody scale rows 22–24; [eye beneath ocular shield; supraocular oblique] .................................. A. tanganicanus 17a. Dorsum heavily blotched with black; venter immaculate; midbody scale rows 26–28 ...................... A. usambaricus 17b. Dorsum, and often venter, usually lineolate or spotted........................................................................................... 18 18a. Midbody scale rows 26–32 ....................................................................................................................... A. lineolatus 18b. Midbody scale rows 22–26....................................................................................................................................... 19 19a. Ocular overlapped (or nearly so) by second supralabial; SE Tanzania.............................................. A. tanganicanus 19b. Preocular overlapped (or nearly so) by second supralabial; E Angola, SE Democratic Republic of the Congo, N Zambia ........................................................................................................................................................ A. schmidti, Published as part of Broadley, Donald G. & Wallach, Van, 2009, 2255, pp. 1-100 in Zootaxa 2255 on pages 76-77

Key to the species of Typhlops in eastern Africa 1a. Preocular overlaps second supralabial (SIP T-III); snout rounded in profile; superior nasal suture present; middorsals 400���425; 24 scale rows; body slender (L/W ratio> 50) ............................................................ T. platyrhynchus 1b. Second supralabial overlaps preocular (SIP T-V); snout wedge-shaped in profile; superior nasal suture absent; middorsals 216���302; scale rows usually 22 (24); body short and robust (L/W ratio ................................................................................................................................................................ T. cuneirostris 2b. Middorsals 257 or more; relatively longer snout (snout/head width ratio> 0.85); eastern Ethiopia, northern Somalia, and northeastern Kenya ...................................................................................................................... T. calabresii, Published as part of Broadley, Donald G. & Wallach, Van, 2009, 2255, pp. 1-100 in Zootaxa 2255 on page 76

Afrotyphlops kaimosae (Loveridge) (Fig. 4C) Kakamega blind-snake Typhlops kaimosae Loveridge, 1935, Bull. Mus. Comp. Zool. 79(1): 5. Type locality: “Kaimosi Forest, near Friends’ Africa Mission, Kakamega District, Nyanza Province, Kenya Colony” [= Kenya], holotype MCZ 40060, collected by A. Loveridge, 7 March 1934; Loveridge, 1936: 226, 1957b: 243, 1960: 267; Pitman, 1938: 44; Uthmöller, 1941a: 40; Barbour & Loveridge, 1946: 209; Roux-Estève, 1974: 47, 1975b: 56; Spawls, 1978: 1; Hahn, 1980: 50; McDiarmid et al., 1999: 90. Typhlops angolensis — Roux-Estève, 1974: 47 (part), 1975b: 56 (part); Spawls, 1978: 2. Description. Snout rounded, prominent. Rostral oval dorsally, moderate ventrally; frontal hexagonal; supraocular transverse, its lateral apex between preocular and ocular; eye visible beneath ocular adjacent to preocular border; nasal sulcus arising from first (right) or second (left) labial; nasal in contact with ocular inferiorly, excluding preocular from lip; SIP T-X (N-N-O-O). MSR 28 with no scale row reductions; MD 390; vertebrae 215; MD/V ratio 1.81; L/D ratio 43. Dark brown above, scales lighter basally, not forming a lineate or punctate pattern, slightly lighter below. Length originally 215 mm, now 208 mm. Habitat. Lowland rainforest. Distribution. Known only from the holotype (Fig. 12). Locality. KENYA. Kaimosi MCZ 40060 (holotype of Typhlops kaimosae Loveridge)., Published as part of Broadley, Donald G. & Wallach, Van, 2009, 2255, pp. 1-100 in Zootaxa 2255 on page 32, {"references":["Loveridge, A. (1935) Scientific results of an expedition to rain forest regions in eastern Africa. I. New reptiles and amphibians from East Africa. Bulletin of the Museum of Comparative Zoology at Harvard College (Cambridge), 79 (1), 1 - 19.","Pitman, C. R. S. (1938) A guide to the snakes of Uganda. The Uganda Society, Kampala, viii + 338 pp., Pls. 1 - 20, A - W.","Uthmoller, W. (1941 a) Die Schlangen Ostafrikas in ihrem Lebenstraum unter besonderer Berucksichtigung der Vulkane Kilimandjaro, Meru, Hanang, der Grossen Ostafrikanischen Bruchstufe und des Usambara-Gebirges. Akademische Verlagsgesellschaft Becker & Erler, Leipzig, 70 pp. [Reprinted in Archiv fur Naturgescichte (Leipzig), (Ser. 2) 10 (1), 1 - 70, 1941].","Barbour, T. & Loveridge, A. (1946) First supplement to typical reptiles and amphibians. Bulletin of the Museum of Comparative Zoology at Harvard College (Cambridge), 96 (2), 57 - 214.","Roux-Esteve, R. (1974) Revision systematique des Typhlopidae d'Afrique: Reptilia - Serpentes. Memoirs du Museum National d'Histoire Naturelle (Paris), (A, Zoologie), (Ser. 2) 87, 1 - 313, Pls. 1 - 12.","Roux-Esteve, R. (1975 b) Serpents: Typhlopidae et Leptotyphlopidae. In: Saeger, H. de, Witte, G. - F. de & Roux-Esteve, R. (Eds.), Exploration du Parc National des Virunga, Fondation pour Favoriser les Recherches Scientifiques en Afrique, Bruxelles, (Ser. 2) (24), 56 - 61.","Spawls, S. (1978) A checklist of the snakes of Kenya. The Journal of the East Africa Natural History Society and National Museum (Nairobi), 31 (167), 1 - 18.","Hahn, D. E. (1980) Liste der rezenten Amphibien und Reptilien: Anomalepididae, Leptotyphlopidae, Typhlopidae. Das Tierreich (Berlin), 101, i - xii, 1 - 93.","McDiarmid, R. W., Campbell, J. A. & Toure, T. A. (1999) Snake species of the world. A taxonomic and geographic reference. Volume 1. The Herpetologists' League, Washington, 511 pp."]}

Megatyphlops gen. nov. Giant blind-snakes Type species. — Onychocephalus mucruso Peters, 1854, Ber. Bekanntmach. Geeignet. Verhandl. Königl.- Preuss. Akad. Wiss. Berlin, 1854(11): 621. Diagnosis. —An endemic African genus characterized by the following combination of characters: snout sharply angular in profile, with a keratinized horizontal edge in adults, supralabial imbrication pattern T-II or T-X, nasal shield incompletely divided and lacking posterior concavity, nostrils directed ventrally, inferior nasal suture contacting first or second supralabial or rostral (Megatyphlops anomalus), 3–7 postoculars, dorsal rostral broad (greater than 1/2 interocular head width), ventral rostral broad (greater than 1/2 internarial snout width), well-developed eye with discernible pupil, dorsum pigmented (often with lineate, mottled or blotched pattern), and vestigial left lung present. Other typical characters include 30–44 midbody scale rows, robust body (length/width 17–58), and large size (maximum length 350–950 mm). Distribution. —Endemic to eastern and southern Africa. Content. —Four species., Published as part of Broadley, Donald G. & Wallach, Van, 2009, 2255, pp. 1-100 in Zootaxa 2255 on page 47, {"references":["Peters, W. C. 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 Bekanntmachunggeeigneten Verhandlungen der Koniglich-Preussischen Akademie der Wissenschaften zu Berlin, 1854 (11), 614 - 628."]}

Key to the species of Rhinotyphlops in southern Africa 1a. Snout distinctly hooked in lateral profile, dorsal pattern of small dark, irregular crossbars on a ground color of pink; SIP T-II ......................................................................................................................................................... R. schinzi 1b. Snout not hooked in profile, dorsal pattern uniformly brown or grey; SIP T-X......................................................... 2 2a. Snout obtusely angled .................................................................................................................................... R. boylei 2b. Snout acutely pointed ................................................................................................................................. R. lalandei, Published as part of Broadley, Donald G. & Wallach, Van, 2009, 2255, pp. 1-100 in Zootaxa 2255 on page 78

Afrotyphlops nigrocandidus (Broadley & Wallach) Bicoloured blind-snake Typhlops punctatus gierrai — (not Mocquard) Barbour & Loveridge, 1928: 106 (part, Bagilo); Loveridge, 1957b: 242 (part, Uluguru Mts). Typhlops gierrai — (not Mocquard) Roux-Estève, 1974: 133 (part, Bagilo); Broadley & Howell, 1991: 21 (part, Ukaguru and Uluguru Mts); Spawls et al., 2002: 286 (part), fig., 2006: 89 (part), fig. Rhinotyphlops nigrocandidus Broadley & Wallach, 2000, Afr. J. Herpetol., 49(2): 166. Type locality: “Kiranzi-Kitungulu Forest Reserve, Udzungwa Mountains, Iringa District, Tanzania ” (08°09’S, 36°05’E, elevation 1500 m), holotype ZMUC 52223, collected by P. Gravlund & M. Andersen, 17 January 1996; Brogard, 2005: 113; Broadley, 2008: 172. Typhlops sp. nov. — Burgess et al., 2002: 145. Description. Snout prominent, rounded. Rostral broad dorsally (two-thirds width of head), oval, but truncated posteriorly and barely reaching the level of the eyes, ventrally it is constricted at the level of the nostrils and has parallel sides. The semidivided nasals extend just beyond the rostral. The oblique supraoculars contact the nasals and the subequal hexagonal frontal, which is larger than the median scales caudad, they overlap the preoculars. A small intercalary scale (first preocular of Roux-Estève, 1974) separates the large preocular from the second and third labials. The intercalary scale is fused with the preocular on both sides of ZMUC 52221 and on the left side of NMZB 7484. The eyes are visible beneath the preocular/ocular sulcus, bordered above by the supraocular. The first labials contact the rostrals and both segments of the nasals, the second labials contact the posterior nasal and overlap the intercalary scales, the third labials contact the intercalary scales and the oculars, the fourth labials border the oculars, SIP T-II (N2, N2/LP, O, O); MSR 28–34; MD 464–542 (mean 509.3, n = 7); vertebrae 242 (n = 1); MD/V ratio 1.91 (n = 1); L/D 38–52 (mean 45.8, n = 6). Largest specimen (NMZB 17548—Uluguru South Forest Reserve) has a total length of 573 mm. Everted hemipenis 4.3 mm long, simple; sulcus undivided and terminating in an apical depression in the central dome of the distal cap; sulcal folds raised but unadorned. Ornamentation: nude, a prominent distal cap with a central nipple; short basal region unadorned. The anterior half of the rostral (nearly all in NMZB 7484) and most of the nasals, preoculars and oculars are yellowish, thereafter the median 11 to 13 dorsal scale rows are black, uniform except for a white transverse mark at the base of each scale, irregular at the lateral margins, venter yellow. In the subadult (ZMUC 52222) the black dorsal colouration covers only 7 scale rows and in NMZB 9914 it covers 9–11 scale rows. NMZB 7484 is irregularly blotched in black and white in subequal proportions, covering a maximum of 15 dorsal scale rows. The first specimen from the Uluguru Mountains, MCZ 23084, is reported to be uniformly blue-black above, except for a minute white spot at the base of each scale (Barbour & Loveridge, 1928: 107). However, NMZB 17548 is white, the dorsum heavily blotched with black. Habitat. MCZ 23084 was hoed up in a shamba bordering rain forest (Barbour & Loveridge, 1928: 108). The other specimens were probably taken inside montane forest (NMZB 17548 at 1515m). Distribution. The southern Eastern Arc mountains in Tanzania, 1450–1750 m (Fig.12). So far recorded from the Ukaguru, Uluguru and Udzungwa Mountains, Afrotyphlops nigrocandidus is also likely to be found in the Nguru Mountains. Localities. TANZANIA. Udzungwa Mts—Kawambwa Forest Reserve ZMUC 52222; Kilanzi-Kitungulu Forest Reserve ZMUC 52223 (holotype of Rhinotyphlops nigrocandidus Broadley & Wallach); Lulanda Forest NMZB 9914; Udzungwa Scarp Forest ZMUC 52222; Ukaguru Mts NMZB 7484; Uluguru Mts—Bagilo MCZ 23084; Uluguru South Forest Reserve NMZB 17548., Published as part of Broadley, Donald G. & Wallach, Van, 2009, 2255, pp. 1-100 in Zootaxa 2255 on pages 38-39, {"references":["Barbour, T. & Loveridge, A. (1928) A comparative study of the herpetological faunae of the Uluguru and Usambara Mountains, Tanganyika Territory, with descriptions of new species. Memoirs of the Museum of Comparative Zoology at Harvard College (Cambridge), 50 (2), 87 - 265.","Loveridge, A. (1957 b) Check list of the reptiles and amphibians of East Africa (Uganda; Kenya; Tanganyika; Zanzibar). Bulletin of the Museum of Comparative Zoology at Harvard College (Cambridge), 117 (2), 151 - 362 + i - xxxvi.","Roux-Esteve, R. (1974) Revision systematique des Typhlopidae d'Afrique: Reptilia - Serpentes. Memoirs du Museum National d'Histoire Naturelle (Paris), (A, Zoologie), (Ser. 2) 87, 1 - 313, Pls. 1 - 12.","Broadley, D. G. & Howell, K. M. (1991) A checklist of the reptiles of Tanzania, with synoptic keys. Syntarsus (Bulawayo), (1), 1 - 70.","Spawls, S., Howell, K. M., Drewes, R. C. & Ashe, J. (2002) A field guide to the reptiles of East Africa: Kenya, Tanzania, Uganda, Rwanda and Burundi. Academic Press, San Diego, 543 pp.","Broadley, D. G. & Wallach, V. (2000) A new blind snake (Serpentes: Typhlopidae) from montane forests of the Eastern Arc Mountains in Tanzania. African Journal of Herpetology (Johannesburg), 49 (2), 165 - 168.","Brogard, J. (2005) Zoogeographical checklist of reptiles. Volume 1. Afrotropical and Palearctic realms. Editions du Point Veterinaire, Maisons-Alfort, 301 pp.","Broadley, D. G. (2008) Book review: \" Pocket guide to the reptiles and amphibians of East Africa. By Stephen Spawls, Kim M. Howell and Robert C. Drewes. 2006. \" African Journal of Herpetology (Matieland), 55 (2), 171 - 173.","Burgess, N. D., Doggart, N. & Lovett, J. C. (2002) The Uluguru Mountains of eastern Tanzania: the effect of forest loss on biodiversity. Oryx (London), 36 (2), 140 - 152."]}

Typhlops platyrhynchus Sternfeld (Fig. 3C) Tanga blind-snake Typhlops platyrhynchus Sternfeld, 1910b, Mitt. Zool. Mus. Berlin, 5(1): 69. Type locality: “ Tanga, Deutsch-Ostafrika ” [= Tanzania], syntypes MCZ 39798, ZMB 20729, ZMB 63442 (formerly 20729A), ZMB 63443 (formerly 20729B), collected by Martienssen; Sternfeld, 1910c: 11; Boulenger, 1915b: 615; Werner, 1921: 299; Schmidt, 1923: 33; Loveridge, 1924: 3, 1957b: 243; Uthmöller, 1941a: 40; Barbour & Loveridge, 1946: 210; Roux-Estève, 1974: 45, figs. 13.8 & 26, 1975a: 490, fig. 30; Hahn, 1977: 37, 1980: 67; Welch, 1982: 129, 1994: 73; Broadley & Howell, 1991: 21; Meirte, 1992: 23; Frank & Ramus, 1995: 254; Wallach, 1998a: 183, 1998b: 422; Mattison, 1999: 148; McDiarmid et al., 1999: 113; Bauer et al., 2002: 165; Spawls et al., 2002: 285, 2006: 92; Murphy, 2003: 552; Brogard, 2005: 115. Description. Snout rounded, prominent. Rostral oval; frontal a laterally truncated crescent; supraocular oblique, its lateral apex between preocular and ocular; eye distinct, under ocular; nasal sulcus arising from the second labial and extending forward towards the rostral, so that the nasal is almost divided; SIP T-III-P (N1, P, P/O, O); MSR 24; MD 400–425 (mean 414.7, n = 4); vertebrae 218–227 (mean 222.7, n = 3); MD/V ratio 1.83–1.87 (mean 1.86, n = 3); L/D ratio 50–60 (mean 55.0, n = 4). Pale reddish-yellow, paler below. Largest specimen (MCZ 39798 — Tanga, Tanzania) has a total length of 273 mm. Distribution. Northeastern Tanzania, NSL (Fig. 11). Known only from the four syntypes. Localities. Tanzania: Tanga. MCZ 39798; ZMB 20729, 20729 A–B (syntypes of Typhlops platyrhynchus Sternfeld)., Published as part of Broadley, Donald G. & Wallach, Van, 2009, 2255, pp. 1-100 in Zootaxa 2255 on pages 25-26, {"references":["Sternfeld, R. (1910 b) Neue Schlangen aus Kamerun, Abessynien u. Deutsch-Ostafrika. Mitteilungen aus dem Zoologischen Museum in Berlin, 5 (1), 69 - 70.","Sternfeld, R. (1910 c) Die Fauna der deutschen Kolonien Herausgegeben mit Unterstutzung des Reichs-Kolonialamtes vom Zoologischen Museum in Berlin. Reihe III: Deutsch-Ostafrika. Heft 2. Die Schlangen Deutsch-Ostafrikas. R. Friedlander & Sohn, Berlin, 47 pp.","Boulenger, G. A. (1915 b) A list of the snakes of East Africa, north of the Zambesi and south of the Soudan and Somaliland, and of Nyassaland. Proceedings of the Zoological Society of London, 1915 (4), 611 - 640.","Werner, F. (1921) Synopsis der Schlangenfamilie der Boiden und Typhlopiden auf Grund des Boulengers'schen Schlangenkatalogs (1893 - 1896). Archiv fur Naturgeschichte (Berlin), (Ser. A) 87 (7), 266 - 338.","Schmidt, K. P. (1923) Contributions to the herpetology of the Belgian Congo based on the collections of the American Museum Congo Expedition, 1909 - 1915. Part II. - Snakes. Bulletin of the American Museum of Natural History (New York), 49, 1 - 146, Pls. 1 - 22.","Loveridge, A. (1924) Check list of the Reptilia recorded from the British Territories in East Africa. The Journal of the East Africa and Uganda Natural History Society (Nairobi), Special Supplement (3), 1 - 16.","Loveridge, A. (1957 b) Check list of the reptiles and amphibians of East Africa (Uganda; Kenya; Tanganyika; Zanzibar). Bulletin of the Museum of Comparative Zoology at Harvard College (Cambridge), 117 (2), 151 - 362 + i - xxxvi.","Uthmoller, W. (1941 a) Die Schlangen Ostafrikas in ihrem Lebenstraum unter besonderer Berucksichtigung der Vulkane Kilimandjaro, Meru, Hanang, der Grossen Ostafrikanischen Bruchstufe und des Usambara-Gebirges. Akademische Verlagsgesellschaft Becker & Erler, Leipzig, 70 pp. [Reprinted in Archiv fur Naturgescichte (Leipzig), (Ser. 2) 10 (1), 1 - 70, 1941].","Barbour, T. & Loveridge, A. (1946) First supplement to typical reptiles and amphibians. Bulletin of the Museum of Comparative Zoology at Harvard College (Cambridge), 96 (2), 57 - 214.","Roux-Esteve, R. (1974) Revision systematique des Typhlopidae d'Afrique: Reptilia - Serpentes. Memoirs du Museum National d'Histoire Naturelle (Paris), (A, Zoologie), (Ser. 2) 87, 1 - 313, Pls. 1 - 12.","Hahn, D. E. (1977) Book review: \" Revision systematique des Typhlopidae d'Afrique, Reptilia - Serpentes. By Rolande Roux-Esteve. 1974. \" Journal of the Herpetological Association of Africa (Umtali), (15), 34 - 39.","Hahn, D. E. (1980) Liste der rezenten Amphibien und Reptilien: Anomalepididae, Leptotyphlopidae, Typhlopidae. Das Tierreich (Berlin), 101, i - xii, 1 - 93.","Welch, K. R. G. (1982) Herpetology of Africa: a checklist and bibliography of the orders Amphisbaenia, Sauria and Serpentes. Robert E. Krieger Publishing Company, Malabar, 293 pp.","Welch, K. R. G. (1994) Snakes of the world: a checklist. 2. Boas, pythons, shield-tails and worm snakes. R & A Research and Information Limited & KCM Books, Somerset, 89 pp.","Broadley, D. G. & Howell, K. M. (1991) A checklist of the reptiles of Tanzania, with synoptic keys. Syntarsus (Bulawayo), (1), 1 - 70.","Meirte, D. (1992) Cles de determination des serpents d'Afrique. Musee Royal de l'Afrique Centrale (Tervuren), (Sciences Zoologiques), 267, 1 - 152.","Frank, N. & Ramus, E. (1995) A complete guide to scientific and common names of reptiles and amphibians of the world. N. G. Publishing, Pottsville, 377 pp.","Wallach, V. (1998 a) The lungs of snakes. In: Gans, C. & Gaunt, A. S. (Eds.), Biology of the Reptilia. Volume 19 (Morphology G). Visceral organs. Society for the Study of Amphibians and Reptiles, Ithaca, pp. 93 - 295.","Wallach, V. (1998 b) The visceral anatomy of blindsnakes and wormsnakes and its systematic implications (Serpentes: Anomalepididae, Typhlopidae, Leptotyphlopidae). Ph. D. Dissertation, Northeastern University, Boston, xxvi + 611 pp.","Mattison, C. (1999) Snake. Dorling Kindersley Publishing, London, 192 pp.","McDiarmid, R. W., Campbell, J. A. & Toure, T. A. (1999) Snake species of the world. A taxonomic and geographic reference. Volume 1. The Herpetologists' League, Washington, 511 pp.","Spawls, S., Howell, K. M., Drewes, R. C. & Ashe, J. (2002) A field guide to the reptiles of East Africa: Kenya, Tanzania, Uganda, Rwanda and Burundi. Academic Press, San Diego, 543 pp.","Spawls, S., Howell, K. M. & Drewes, R. C. (2006) Reptiles and amphibians of East Africa. Princeton University Press, Princeton, 240 pp.","Murphy, J. B. (Ed.) (2003) Grzimek's Animal life encyclopedia. Volume 7. Reptiles. Second edition. The Gale Group, Farmington Hills, 593 pp.","Brogard, J. (2005) Zoogeographical checklist of reptiles. Volume 1. Afrotropical and Palearctic realms. Editions du Point Veterinaire, Maisons-Alfort, 301 pp."]}

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.