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) Author Wüster, Wolfgang Author Chirio, Laurent Author Trape, Jean-François Author Ineich, Ivan Author Jackson, Kate Author Greenbaum, Eli Author Barron, Cesar Author Kusamba, Chifundera Author Nagy, Zoltán T. Author Storey, Richard Author Hall, Cara Author Wüster, Catharine E. Author Barlow, Axel Author Broadley, Donald G. text Zootaxa 2018 2018-08-01 4455 1 journal volume 29115 10.11646/zootaxa.4455.1.3 0d838ca1-52f1-4278-895d-68fc481b7558 1175-5326 1456971 2EA2EE4A-2BA4-438A-A1CB-D9C54F37DC78 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 ). FIGURE 7. Naja ( Boulengerina ) savannula sp. nov. Top row and bottom left: holotype, MNHN 2018.0002. Bottom right: live specimen from Kindia, Guinea, showing conspicuous, broad dorsal bands and ventral banding, including narrow accessory bands (not vouchered). Photos J.-F. Trape. 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 ).