Taxonomic and morphological revision of butterfly rays of the Gymnura micrura (Bloch & Schneider 1801) species complex, with the description of two new species (Myliobatiformes: Gymnuridae) Author Carvalho, Marcelo Rodrigues De text Zootaxa 2017 2017-10-12 4332 1 1 74 journal article 31854 10.11646/zootaxa.4332.1.1 c8e7f4a7-d89e-4d9b-9fe7-e4938f445e75 1175-5326 1052041 C7D6FF64-5813-4AAB-AE40-C24ABB74433F Gymnura sereti , sp. nov. ( Figs. 22–31 , 34c , 36c, d , 37c , 38c , 42c , 43c, d , 44c , 46c – 48c ; Tabs. 5 , 6 ) Pteroplatea micrura : Ehrenbaum, 1914 : (listed, Cameroon); Ehrenbaum, 1915 : 73 (listed, Cameroon);? Barnard, 1925 : 80 (description, reported from Agulhas Bank);? Monod, 1927 : 650 (W. Africa, not seen). Gymnura micrura : Fischer et al. 1981 : 223 –224 (identification guide, Eastern Central Atlantic); Cappetta, 1984 : 634 , pl. 1, fig. 9 (teeth morphology, Ivory Coast); McEachran & Séret, 1990 : 65 (checklist, Eastern Tropical Atlantic); Bianchi, 1986 : 117 (identification guide, Angola); Séret & Opic, 1997 : 68 –69 (identification guide, Tropical West Africa); Albaret et al. 2004 : 41 (listed, Gambia estuary). Holotype . USNM 193741 ( 383 mm DW, adult male), Liberia , off St. Paul River , 7-13 m , 0 6 Jan 1953 , accession num. 202912 ( Fig. 22 ). Paratypes. USNM 193896 ( 338 mm DW, adult male), Liberia, Bushrod Island Beach, collector: G. Miller, 30 Oct 1952 , field # 1791, accession num. 202912, date cat. 0 9 Jul 1963 ( Fig. 23 ); USNM 279558 ( 289 mm DW, female), Sierra Leone , St. Anne Banana Islands, collector: G. Naylor , Feb 1986 , field # 27, accession num 369276, remarks: from Sierra Fishing Co. (free town, Sierra Leone ) ( Fig. 24 ) ; MNHN 1989-1216 ( 500 mm DW, adult male), Senegal, Bandiala, 15o00’00”N, 18o00’00”W, estuary, 14m , sector 3423, collectors: Cauri, Séret, 0 7 May 1983 , n. station 02; MNHN 1989-1225 (gravid female, 665 mm DW with 3 embryos: female, 79 mm DW; male, 81 mm DW and male, 85 mm DW), Senegal, Saloum, Missira, 15o00’00”N, 18o00’00”W, estuary, 4–10m , sector 3423, collectors: Cauri, Séret, 0 7 May 1983 , trawl, n. station 4; MNHN 1989-1229 (juvenile female, 324 mm DW), Ivory Coast , 0 5o 04’59”N, 0 3o 48’00”W, marine, 15m , sector 3434, donation: Campagne Chalci , n. 291, collectors: Andre Nizery , Camp. Chalci , Mar 1979 , trawl, n. station 88 ; MNHN 1989-1238 (juvenile male, 290 mm DW), Senegal, Saloum, Bandiala, 15o00’00”N, 18o00’00”W, estuary, 7–12m , sector 3423, collectors: Cauri, Séret, 0 7 May 1983 , trawl, n. station 0 1. Diagnosis . A medium size butterfly ray occurring in the eastern central Atlantic. Gymnura sereti , sp. nov. is distinguished from other species of butterfly rays (except G. micrura and G. lessae ) by the combination of the following characters: absence of spiracular tentacle, caudal stings and dorsal fin; tail relatively short (mean postcloacal length 24% DW), and prominently banded, presenting 3 to 6 (often 4) black bands that may be less distinct in large adults. Gymnura sereti , sp. nov. is distinguished from G. micrura and G. lessae by the following characters: clasper of mature males stouter and shorter, Lclasper 6.8–9.2% DW ( vs . clasper slender and longer, Lclasper 9.3– 11% DW in G. micrura and 9.4–14.3% DW in G. lessae ); cranial fontanelle keyhole-shaped ( vs . U-shaped); mesopterygium divided into two solid elements ( vs . mesopterygium divided into one anterior solid element and 5– 7 smaller fragments); distance between anteroventral and posteroventral fenestrae of scapulocoracoid about 20% of scapulocoracoid length ( vs . distance between AVF and PVF 30–40% of scapulocoracoid length); lateral projection of synarcual base starting at its half length ( vs . lateral projection of the base of synarcual starting at its anterior third); higher number of diplospondylous vertebrae (mean 110 vs . 98 and 97 in G. micrura and G. lessae , respectively), and lower number of radials in the posterior mesopterygial element compared to the mesopterygial fragments in G. micrura and G. lessae (range 10–15 vs . 15–17 and 14–19 in G. micrura and G. lessae , respectively). Gymnura sereti , sp. nov. is further distinguished from G. micrura by its dorsal side brownish with a vermiculate background ( vs . dorsal side usually uniformly brown or gray, without any vermiculate pattern), and basipterygium representing <50% of pelvic propterygial length ( vs . basipterygium representing>60% of pelvic propterygial length). Gymnura sereti , sp. nov. is further distinguished from G. lessae by the dorsal contour of its hyomandibula with two conspicuous protuberances ( vs . dorsal contour of the hyomandibula with a conspicuous proximal protuberance followed by a inconspicuous distal one) and lower number of pectoral radials (range 109–115 vs . 118–127). Other differences discussed in the text. Description. Measurements are presented in Table 5 ; meristic data in Table 6 . The following description is based on all specimens examined. Throughout the text the proportions are presented as: minimum value–maximum value (mean). See Figs. 22–28 , 29a , 30 and 31 for external morphology and color, Fig. 34c for ventral lateral line pattern and Figs. 29b , 36c, d , 37c , 38c , 42c , 43c, d , 44c and 46c – 48c for skeletal morphology. TABLE 5. Measurements taken on specimens of Gymnura sereti sp. nov. Holotype: USNM 193741. Ranges, means and standard deviations (SD) were calculated from all specimens (N). Postcloacal clasper length was obtained only from adult specimens.

Holotype	N	Range	Mean	SD
mm %DW	mm	%DW	mm	%DW	mm	%DW
Disc width	383 100	51	182–748	100–100	379.4	100	150.4	–
Disc length	226 59	51	101–424	52.9–61.3	215.2	56.5	87.3	1.9
Total length	301 78.6	51	136–547	68.6–82.9	283.1	74.8	110.5	2.9
Snout to max. width	241 62.9	51	111–450	57.8–66.6	230.7	61	89.9	1.6
Preorbital snout length	40 10.4	51	13–61	7.1–11	33.6	8.8	14	0.9
Orbit diameter	10 2.6	51	5–17	1.7–3.2	8.5	2.3	2.6	0.3
Eye diameter	6 1.6	51	3–8.5	0.9–2.2	5.2	1.5	1.4	0.3
Interorbital width	39 10.2	51	17.5–68	8.4–10.7	35.7	9.4	14.2	0.4
Spiracle length	12 3.1	51	7–29	2.3–4.2	12.8	3.4	5.7	0.4
Interspiracular width	36 9.4	51	17–61	8–9.7	33.5	8.9	12.3	0.4
Precloacal length	193 50.4	51	85–368	44.8–52.4	184.3	48.3	76.4	2
Postcloacal tail	96 25.1	47	40–166	15.8–27.7	89.1	23.8	31.8	2
Cloaca length	– –	41	5–34	1.9–4.7	12.7	3.1	7.1	0.5
Head length	92.5 24.2	51	41–166	20.9–26.1	88.1	23.3	34.5	1.2
Preoral snout length	46 12	51	16–89	8.7–13.3	41.6	10.8	18.8	1
Mouth width	33 8.6	51	17–62	7.6–9.9	32.8	8.7	12.3	0.5
Nasal curtain width	31 8.1	50	13.5–51	6.1–8.5	28.2	7.5	10.3	0.5
Nasal curtain length	10.5 2.7	49	4–17	1.7–2.9	8.9	2.4	3.3	0.3
Nasal curtain base	25 6.5	50	12–42	5.3–7	26.6	6.3	8.3	0.4
Nostril length	11 2.9	50	4–24	2.7–4.3	12.5	3.4	4.8	0.3
Internasal width	8 2.1	51	3.5–17	1.6–2.7	7.5	2	3	0.3
Dist. btw. 1 st gill slits	66.5 17.4	50	34–126	16–19.1	65	17.2	25	0.7
Dist. btw. 5th gill slits	43 11.2	50	22–87	10.3–13	43	11.4	16.8	0.5
Preorbital snout width	157 41	51	72–300	33.6–44	148.3	39.1	60.1	1.9
Postspiracle snout width	198 51.7	50	95–417	47.8–57.1	197.9	52.1	80.1	2.3
Pectoral-fin height	192 50.3	51	90–377	47.4–56	192.3	50.8	75.9	1.4
Clasper (postcloacal)	29 7.6	14	26.5 - 45	6.8–9.2	35.7	8	6.2	0.6

FIGURE 22. Dorsal (A) and ventral (B) views of holotype of Gymnura sereti , sp. nov. (USNM 193741, adult male, 383 mm DW). TABLE 6. Meristic data for specimens of Gymnura sereti sp. nov. Abbreviation: SD, standard deviation.

N	Range	Mode	Mean	SD
Vertebral counts
to prepelvic process	6	31–34	34	32.8	1.3
to posterior tip of basipterygium	6	45–50	50	48	2.3
Monospondylous	6	31–36	36	33.5	2.3
Diplospondylous	5	99–121	–	110.2	9.2
Total vertebrae	5	135–152	–	143.2	7.5
Radial elements
Propterygial	8	34–38	36	36	1.3
Mesopterygial
- first element	8	6–10	10	8.7	1.2
- second element or fragments	8	10–15	11	12.1	1.5
- total	8	19–22	21	20.9	0.9
Metapterygial	6	52–58	54	54.4	1.7
Total pectoral radials	6	109–115	110	111.1	1.7
Pelvic fin radials	6	11–18	14	13.9	2.4

External morphology. Disc lozenge shaped, 1.63–1.89 (1.77) times broader than long [1.64–1.76 (1.72) for adult males; 1.72–1.82 (1.79) for adult females]. Trunk strongly flattened, slightly raised above scapular region and posterior head. Snout relatively short and obtuse with a subtle lobe at snout tip. Although adult males present proportionally longer snouts, sexual dimorphism is not so evident in disc shape, and adult males do not possess a triangular anterior half of disc as observed in G. micrura and G. lessae ( Fig. 25 ). Preoral snout length 8.7–13 (11)% DW [9–12 (10)% DW in adult females; 11–13 (12)% DW in adult males], preorbital snout length 7.1–11 (9)% DW [8–9 (8)% DW in adult females; 9–11 (10)% DW in adult males], preorbital snout width 34–44 (39)% DW, postspiracle snout width 48–57 (52)% DW. Anterior margins of disc with a medial concavity, becoming weakly convex towards extremities; pectoral-fin apices acutely angular; posterior margins convex; free rear tip broadly rounded; axis of greatest width positioned posterior to the half length of disc. Pelvic fin single lobed, rectangular, with rounded corners ( Fig. 26 ). Skin entirely smooth, without denticles on the dorsal and ventral surfaces of the disc. Interorbital space broad, interorbital width 8.4–11 (9.4)% DW. Eyes dorsolateral, small, oval, and protruding slightly ( Fig. 27 ); eye diameter 0.9–2.2 (1.5)% DW, representing 9–23 (15)% of interorbital width; orbit diameter 1.7–3.2 (2.3)% DW, representing 41–94 (71)% of spiracle length. Eyes more protruded and relatively larger in embryos. Spiracles posterior to eyes, relatively large, lozenge-shaped, contour of the inner margin rectilinear to slightly concave ( Fig. 27 ); spiracular length 2.3–4.2 (3.4)% DW, 1.6–4.1 (2.4) times eye diameter; its inner posterior margin without tentacles. Ventral head length 21–26 (23)% DW. Nostril narrowly oval, diagonally directed, only the circular distal margin not covered by nasal curtain, posterior lateral margin with lobe; nostril length 2.7–4.3 (3.4)% DW, 1.0–2.2 (1.7) times internasal width; internasal width 1.6–2.7 (2.0)% DW. Anterior nasal flaps medially expanded and fused into a broad, skirt-shaped, posteriorly expanded nasal curtain that covers internasal space and reaches mouth ( Fig. 28a ). Nasal curtain skirt-shaped, weakly to moderately bilobed, wide, its width 2.5–4.1 (3.2) times its length; lateral margin concave; posterolateral apices rounded; posterior margin straight to weakly concave, a small medial gap may be present. Mouth relatively wide, its width 7.6–9.9 (8.7)% DW, 31–42 (38)% head length, 1.0–1.4 (1.2) times nasal curtain width, without papillae on floor or labial folds, although some striations may originate radially from mouth corners; lower lip arched rearward toward corners, medial region moderately ( Fig. 28b ) to strongly concave ( Fig. 28c ); when strongly concave, medial region limited by angular projections which may form prominent rounded lumps or knobs ( Fig. 28d ). A patch of corrugate skin forms a half circle below mouth, ending at mouth corners. Small, numerous and closely crowded teeth in bands; teeth with one medial, pointed cusp directed towards inside of mouth; base somewhat broad, not twisted, practically straight; labial face of teeth T-shaped ( Fig. 28e ); teeth similar between jaws and sexes, their number increasing with growth. FIGURE 23. Dorsal (A) and ventral (B) views of Gymnura sereti , sp. nov. (paratype, USNM 193896, adult male, 338 mm DW). FIGURE 24. Dorsal (A) and ventral (B) views of Gymnura sereti , sp. nov. (paratype, USNM 279558, female, 289 mm DW). FIGURE 25. Specimens of Gymnura sereti , sp. nov. , in dorsal view. A) Adult male, 432 mm DW (MNHN 1989-1240); B) Paratype, gravid female, 665 mm DW (MNHN 1989-1225); C) Juvenile male, 290 mm DW (FLMNH 29993); D) Juvenile female, 267 mm DW (ZMB 8868). FIGURE 26. Ventral view of pelvic fin of Gymnura sereti , sp. nov. A) Male (MNHN 1989-1240, 432 mm DW); B) Female (paratype, MNHN 1989-1225, 665 mm DW). FIGURE 27. Dorsal view of head of Gymnura sereti , sp. nov. (paratype, MNHN 1989-1216, 500 mm DW). Gill slits moderately S-shaped, first four gill slits markedly larger than fifth ( Fig. 28a ). Distance between gill slits decreasing posteriorly; distance between inner ends of fifth pair 58–74 (66)% distance between inner ends of first pair; distance between inner ends of first pair 16–19 (17)% DW, 67–84 (74)% ventral head length, 6.3–11.0 (8.7) times internasal width; distance between inner ends of fifth pair 10–13 (11)% DW, 42–54 (49)% ventral head length, 4.1–7.2 (5.7) times internasal width. Tail slender and short, whip-like, cross-banded (see color section), tapering toward tip, without caudal stings or dorsal fin ( Fig. 31 ); postcloacal tail 16–28 (24)% DW, 28–49 (42)% DL, 32–59 (50)% precloacal length; embryos with proportionally longer tails, postcloacal tail 29–35 (31)%; low longitudinal skin folds present on dorsal and ventral surfaces, skin folds not reaching tail tip. Clasper of mature males cylindrical, somewhat depressed, relatively stout and shorter (compared to G. micrura and G. lessae ) ( Fig. 29 ); distance from the posterior margin of cloaca to clasper tip 6.8–9.2 (8)% DW; tip somewhat depressed, rounded to bluntly pointed, not calcified; non-calcified portion relatively long when compared to G. micrura and G. lessae ; apopyle (on the anterior dorsal surface) connected to hypopyle by a long, dorsomedial, posteriorly curved clasper groove; spermatic duct ending well before reaching clasper tip; rhipidion and pseudorhipidion absent; a long ventral pseudosiphon laterodistally situated to hypopyle; a well-developed dorsal pseudosiphon on inner margin of clasper, the dorsal pseudosiphon more anteriorly located when compared to G. micrura and G. lessae ; ventral surface of clasper entirely smooth. Coloration. Freshly caught specimens not analyzed, description based on preserved specimens from collections. Dorsal side brownish, vermiculated (similar to G. lessae ). The vermiculate pattern varying from less conspicuous with lighter and darker shades to a conspicuous pattern where the lighter areas become larger and more remarkable ( Figs. 30a–f ). Light bands may be regularly distributed on outer margins of disc ( Fig. 30g ) and round white spots may be present on disk ( Fig. 30h ). Like its congeners, this species probably has the ability to vary its shade according to the bottom. Ventral surface whitish to brownish, creamy; generally darkening toward edges. Tail prominently cross-banded, with 3–6 black bands, often 4 ( Fig. 31a ); a black dot may be present between the black bands ( Fig. 31b ); bands less distinct in large specimens ( Fig. 31c ). FIGURE 28. Morphological details of sereti , sp. nov. A) Ventral view of head (paratype, USNM 193896, 338 mm DW); B) Detail showing lower lip with medial region moderately concave (MNHN 1985-217, 297 mm DW); C) Detail showing lower lip with medial region strongly concave (MB 2297, 490 mm DW); D) Detail showing lower lip with prominent knobs (AMNH 9897, 560 mm DE); E) Lower tooth, labial face (modified from Cappetta, 1984, pl. 1, fig. 9). Size . One litter presents embryos 183–199 mm DW without external yolk sac, but with the intestine full of yolk, while a term embryo 212 mm DW was found. The smallest juvenile analyzed measured 182 mm DW. Birth size is estimated around 180–220 mm DW. The largest juvenile male analyzed was 450 mm DW, while the smallest adult male was 338 mm DW. A 362 mm DW male was maturing, and all males larger than 450 mm DW were mature. The size at maturity of males should be around 300–450 mm DW. The largest male examined measured 560 mm DW (AMNH 9897). The smallest gravid female examined was 665 mm DW (MNHN 1989-1225), and the largest female examined (also pregnant) was 748 mm DW (MNHN 1989-1247). This species should reach about 1000–1100 mm DW. Geographical distribution and habitat. According to material examined, G. sereti , sp. nov. , inhabits the West African coast from Senegal to Angola (south to Baía Farta, Benguela Province) ( Fig. 32 ). A butterfly ray supposedly without spiracular tentacles, identified as Pteroplatea micrura , was recorded from Agulhas Bank, South Africa, by Barnard (1925) . This may be the southernmost record of G. sereti , sp. nov. However, we can not confirm this since in the description of the genus provided in the same study, the author mentions that its tail has a sting; also, G. natalensis (commonly occurring in this area) may have extremely reduced spiracular tentacles, potentially a source of confusion with G. sereti , sp. nov. This species inhabits sandy and muddy bottoms in inshore waters, also entering estuaries. FIGURE 29. Clasper of Gymnura sereti , sp. nov. A) Illustration in dorsal view, right clasper, showing external morphology; B) Radiograph dorsoventral view (holotype, USNM 193741), showing skeleton. Abbreviations: ap , apopyle; at , accessory terminal cartilage; ax , axial cartilage; B , beta cartilage; b1 , first basal segment; b2 , second basal segment; cg , clasper groove; dp , dorsal pseudosiphon; DT1 , dorsal terminal 1 cartilage; DT2 , dorsal terminal 2 cartilage; hpp , hypopyle; pf , pelvic fin; VM , ventral marginal cartilage; vp , ventral pseudosiphon; VT , ventral terminal cartilage. FIGURE 30. Dorsal color variation in Gymnura sereti , sp. nov. A–F) Variation in vermiculate color pattern; B, D and F are amplifications of the white squares in A, C and E (A. SMF 3992, 284 mm DW; C. MNHN 1989-1215, 468 mm DW; E. MNHN 1989-1240, 432 mm DW); G) Dorsal view of anterolateral disc region showing light bands on outer anterior margin of disc (FMNH 188133, 368 mm DW); H) Specimen with round white spots on back (paratype, MNHN 1989-1229, 324 mm DW). FIGURE 31. Tail color in Gymnura sereti , sp. nov. , dorsal view A) Tail cross-banded (FLMNH 29993, 512 mm DW); B) Tail cross-banded, presenting a black dot between the black bands (MNHN 1989-1232, 730 mm DW); C) Tail of a larger specimen, with the bands less distinct (MNHN 1989-1217, 535 mm DW). Etymology. This species is a tribute to Dr. Bernard Séret, in honor of his important contributions to the systematics of elasmobranchs and to our knowledge of West African fishes. Bernard has contributed substantially to the collections of the MNHN, including thousands of specimens from West Africa, and is an esteemed friend of the second author. Material examined (55 specimens). Senegal ( 8 specimens ): FLMNH 176854 (female, 541 mm DW), Djifère , Djifère Senegal market ; MNHN 1989-1214 (juvenile male, 220 mm DW), estuary of Saloum river , 15o00’00”N , 18o00’00”W ; MNHN 1989-1215 (adult male, 468 mm DW), estuary of Saloum river , 15o00’00”N , 18o00’00”W ; MNHN 1989-1217 (adult male, 535 mm DW), estuary of Saloum river , 15o00’00”N , 18o00’00”W ; MNHN 1989- 1218 (male, 510 mm DW), estuary of Saloum river , 15o00’00”N , 18o00’00”W ; MNHN 1989-1220 (juvenile male, 207 mm DW), 12o58’01”N, 16o52’59”W; MNHN 1989-1223 (adult male, 451 mm DW), estuary of Saloum river , 15o00’00”N , 18o00’00”W ; MNHN 1989-1224 (adult male, 404 mm DW), estuary of Saloum river , 15o00’00”N , 18o00’00”W . Guinea-Bissau ( 5 specimens ): MNHN 1989-1232 (gravid female, 730 mm DW with 3 embryos), Cacheu , 12o12’00”N , 16o 10’ 0 1 ” W ; MNHN - 1235 (juvenile male, 246.5 mm DW), Cacheu , 12 o 12 ’ 0 0 ” N , 16o10’01”W . Sierra Leone ( 2 specimens ): FLMNH 29993 ( 2 specimens : juvenile male, 290 mm DW; female, 512 mm DW), upper Sierra Leone River , 0 8o 34’30”N, 13o06’00”W . Liberia ( 3 specimens ): BMNH 1911.6 .30.6 (juvenile male, 200 mm DW), Axim , Gold Coast ; USNM 222623 ( 2 specimens : female, 289 mm DW; female, 311 mm DW), 0 6o 18’N, 10o49’W . Ivory Coast ( 5 specimens ): MNHN 1989-1219 (juvenile male, 249 mm DW), 0 5o 04’59”N, 0 3o 48’00”W ; MNHN 1989-1226 (female, 476 mm DW), 0 5o12’00”N, 0 4o21’00”W; MNHN 1989- 1227 (adult male, 397 mm DW), 0 5o12’00”N, 0 3o49’01”W; MNHN 1989-1240 (adult male, 432 mm DW), 0 5o12’00”N, 0 4o21’00”W; MNHN 1989-1242 (juvenile male, 213 mm DW), 0 5o04’59”N, 0 3o48’00”W. Ghana ( 3 specimens ): BMNH 1930.3 .24.4 (juvenile male, 236 mm DW), Accra , Gold Coast ; BMNH 1930.8 .26.9 (juvenile female, 184 mm DW), Accra , Gold Coast ; USNM 222600 (juvenile female, 182 mm DW), Tema , Teshie Bay . Togo ( 13 specimens ): MNHN 1989-1244 (female, 660 mm DW), 0 6o 13’01”N, 0 1o 37’59”E ; MNHN 1989-1245 (gravid female, 692 mm DW with 4 embryos: male, 183 mm DW; male, 186 mm DW; male, 199 mm DW and male, 212 mm DW), 0 6o13’01”N, 0 1o37’59”E; MNHN 1989-1247 (gravid female, 748 mm DW with 4 embryos: female, 134 mm DW; female, 140 mm DW; female, 144 mm DW and male, 145 mm DW), 0 6o13’01”N, 0 1o37’59”E; SMF 3992 (female, 284 mm DW); SMF 3993 (juvenile male, 283 mm DW). Republic of Benin ( 4 specimens ): MNHN 1967-0737 (juvenile male, 207 mm DW), 0 6o 19’01”N, 0 2o 24’00”E ; MNHN 1969-0211 (3 specimens: juvenile male, 250 mm DW; juvenile male, 306 mm DW; juvenile male, 319 mm DW), 0 6o21’00”N, 0 2o54’00”E. Nigeria ( 2 specimens ): MNHN 1985-0217 (female, 297 mm DW), 0 3o 49’59”N, 0 6o 13’59”E ; USNM 198011 (female, 299 mm DW), vicinity of Lagos . Cameroon ( 1 specimen ): ZMH 10333 (juvenile male, 261 mm DW), Duala . Gabon ( 1 specimen ): MNHN 1985-0237 (juvenile male, 284 mm DW), 0 0o 00’00”N, 0 5o 00’00”E . Angola (6 specimens): AMNH 9897 (adult male, 560 mm DW); FMNH 188133 (juvenile male, 368 mm DW); MB 2106 (subadult male, 362 mm DW), enseada Mucula ; MB 2279 (adult male, 489 mm DW), Baía da Caota , Baía Farta ; MB 2297 (adult male, 490 mm DW); MB 2364 (female, 514 mm DW), Catumbo Dande , Bengo . West Africa ( 2 specimens ): ZMB 8868 (female, 267 mm DW), Loango Coast ; BMNH 1963.8.9.3 (female, 337 mm DW).