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 ) 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 Ogooué 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 ; Ceríaco 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. FIGURE 8 . Naja ( Boulengerina ) melanoleuca . Adult specimens from Yaoundé, Cameroon (left—photo J.-F. Trape) and Tsibilé, Gabon (right—photo L. Chirio). Note the diffuse but distinct hood mark that is often present in this species, and the combination of broad main bands and narrow accessory bands on the ventral side. TABLE 8. Variation in meristic characters in the species of the N. melanoleuca complex.

Naja guineensis N =71	Naja melanoleuca N =307		Naja savannula N =24		Naja subfulva (excl. eastern and southern African pops.) N =202		Naja peroescobari N = 8
Mean ± 1SD	Range	Mean ± 1SD	Range	Mean ± 1SD	Range	Mean ± 1SD	Range	Mean ± 1SD	Range
Midbody dorsal scale rows	17.0 ± 1.0	17–19	19.0 ± 0.3	17–21	19.0 ± 0	19	19.0 ± 0.2	18–21	19.0 ± 0.0	19
Ventrals	211.8 ± 4.2	203–221	217.8 ± 4.4	206–232	216.9 ± 4.2	211–226	210.0 ± 4.8	197–228	209.4 ± 3.4	204–215
Subcaudals	65.1 ± 2.3	60–70	65.5 ± 2.7	57–74	70.0 ± 1.6	67–73	62.8 ± 2.8	57–72	66.2 ± 4.6	57–70
First dark ventral of first main band	18.0 ± 2.3	10–23	16.3 ± 1.7	11–21	16.4 ± 2.2	11–21	14.2 ± 1.7	10–22	18.2 ± 3.4	14–22
Last dark ventral of first main band	22.2 ± 2.3	16.26	19.9 ± 1.9	15–25	19.4 ± 2.4	14–25	18.2 ± 2.1	13–26	22.63 ± 6.4	15–29
Width of first main band (in ventral scales)	5.2 ± 1.4	2–9	4.5 ± 1.0	2–9	4.0 ± 0.8	3–5	5.0 ± 1.0	2–10	5.4 ± 3.1	2–9
Number of main bands	1.5 ± 0.7	1–4	3.7 ± 0.9	1–7	3.6 ± 1.3	1–7	2.5 ± 0.8	1–5	1.7 ± 0.7	1–3
Number of accessory bands	0.8 ± 1.2	0–5	1.4 ± 1.7	0–9	1.4 ± 1.9	0–6	0.3 ± 0.79	0–4	0.7 ± 0.9	0–2
Total number of discrete dark bands	2.3 ± 1.4	1–6	5.1 ± 1.7	2–12	5.0 ± 2.5	2–12	2.8 ± 1.1	1–8	2.4 ± 1.2	1–5
Last ventral of last discrete dark band	28.5 ± 9.2	17–52	68.6 ± 15.4	24–114	59.4 ± 19.1	19–103	36.8 ± 10.0	13–82	27.6 ± 15.4	15–55

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 .