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 ) guineensis sp. nov. Broadley, Trape, Chirio, Ineich & Wüster Naia melanoleuca (not Hallowell) Boulenger, 1896 : 376 (part, var. B [c,d], C). Naja sp. 2 cf. melanoleuca Hallowell, 1857 (blackish dorsum) Trape & Baldé, 2014 : 318 . Naja sp. 2 cf. melanoleuca (forest form). Trape & Baldé, 2014 : 336 . Holotype : MNHN 1921.0485, a male from N’Zébéla, Macenta Prefecture, Nzérékoré 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. FIGURE 5. Naja ( Boulengerina ) guineensis sp. nov. Left and top right: holotype, MNHN 1921.0485, dorsal and ventral view and side view of head. Note extensive mottling of throat and anterior ventral side and limited posterior extent of lighter ventral markings. Bottom right: live adult specimen measuring approximately 200 cm total length, from Sekondi-Takoradi, Western Region, Ghana, displaying dark suffusion of throat and anterior venter (not preserved; photo L. Chirio). 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 & BAldé, 2014 ) ( Fig. 6 ). ThErE is A siNglE rEcOrd frOm CONtubOEl, GuiNEA BissAu (MBL 535). All OthEr rEcOrds 10°N Or lOwEr.