Siluriformes, Amphiliidae) with the Descriptions of New Species from the Upper Sanaga River and Nyong River Basins

Home , Doumea, Microsynodontis, Phractura, Sanaga River

Species of the Doumea chappuisi Complex (Siluriformes, Amphiliidae) with the Descriptions of New Species from the Upper Sanaga River and Nyong River Basins

Carl J. Ferraris, Jr.1, Paul Skelton2, and Richard P. Vari3

Copeia 2010, No. 4, 705–715

Species of the Doumea chappuisi Complex (Siluriformes, Amphiliidae) with the Descriptions of New Species from the Upper Sanaga River and Nyong River Basins

Carl J. Ferraris, Jr.1, Paul Skelton2, and Richard P. Vari3

The Doumea chappuisi complex within the catfish family Amphiliidae is diagnosed on the form of the dorsolateral and ventrolateral processes of the vertebrae along the posterior portion of the body. Three species are recognized in the complex: Doumea chappuisi of the West African coastal river basins in Guinea, Guinea-Bissau, Cote d’Ivoire, and Liberia; D. reidi, new species, described herein from a portion of the upper Sanaga River in Nigeria; and D. stilicauda, new species, described herein from the Nyong River basin in Cameroon. Members of the complex are distinguished from each other on the basis of the overall body form, the caudal-peduncle length, the predorsal length, the head length, the degree of development of the pelvic fin in larger specimens, the anterior extent of the exposed vertebral processes along the ventral surface of the body, and details of the pigmentation pattern of the unbranched rays of the pectoral and pelvic fins.

ATFISHES of the amphiliid genus Doumea range interesting specimen from the Nigerian portion of the upper across a major portion of Africa from Angola Sanaga River. While closely resembling other species of C through the Congo River basin and the Lower Doumea in most features, the vertebral ossifications of the Guinea region to Guinea-Bissau. The seven species now posterior portion of the body in that specimen extend recognized in the genus (Ferraris, 2007; Skelton, 2007a, through the dermis, albeit not expanding distally into bony 2007b) are all elongate streamlined dwellers of flowing water plates comparable to those present in Andersonia, Belonogla- habitats with enlarged pectoral and pelvic fins and ventrally nis, Phractura, and Trachyglanis. At approximately the same positioned mouths used for epibenthic grazing. Species of time, the WorldFish Centre was engaged in a fishery Doumea are characterized by a number of unusual internal investigation of the ichthyologically poorly known middle modifications presumably correlated with this distinctive Nyong River in Cameroon (Brummett et al., 2010). Those life style, most notably the expansion of the neural and efforts yielded specimens of a species of Doumea that hemal spines and the development of bilateral dorsolateral differed from Doumea gracila, which had been described and ventrolateral processes (Harry, 1953:fig. 11e). Doumea previously from the Nyong River (Skelton, 2007a). That had been distinguished within the Doumeinae by the species also exhibited extensions of the dorsolateral and absence of two series of superficial osseous plates that ventrolateral vertebral processes of the posterior portion of extend dorsolaterally from lateral of the base of the dorsal the body similar in form to those of the specimen from the fin to the posterior end of the caudal peduncle and upper Sanaga River. The Nyong and Sanaga River specimens, ventrolaterally from posterior of the pelvic-fin origin to nonetheless, clearly represented different species. the end of the caudal peduncle in the other genera of the This intermediate degree of development of the exten- subfamily (Andersonia, Belonoglanis, Phractura, and Trachy- sions of the vertebrae (penetrating the skin but not glanis; Poll and Gosse, 1995:pl. 58, figs. 343–351). These expanding into surface plates as in Andersonia, Belonoglanis, ossifications, which are sometimes referred to as bony Phractura, and Trachyglanis) initially appeared unique to scutes, are distal expansions of dorsolateral and ventrolat- those two species within the Amphiliidae. Close examina- eral extensions of the caudal vertebrae, which are charac- tion of Doumea chappuisi, however, revealed that it possessed teristic of doumeins (Harry, 1953; Skelton, 2007a). Species of a similar condition. This moderate-sized and, until relatively Doumea were reported to have the vertebral processes recently, rare species was not reported to exhibit superficial extending distally to the dermis, but were not known to ossifications of the dorsal and ventral surfaces of the penetrate to the skin surface (Skelton, 2007a:58). Harry posterior portion of the body. However, examination of (1953:220) commented, however, that ‘‘scutal elements are specimens of a range of body sizes revealed that the vertebral evident without dissection only in the smaller specimens’’ processes extend to the skin surface in that region in all of D. typica. Our examination of individuals of that species specimens. Externally, the ossifications in D. chappuisi and across a range of sizes revealed that the vertebral processes the species from the Nyong and Sanaga rivers appear as a are superficially evident through the relatively thin, some- narrow, slightly irregular series of bony processes that in what transparent skin of juveniles, but nonetheless fail to combination form distinct ridges, rather than the plate-like reach the body surface. scutes of Andersonia, Belonoglanis, Phractura, and Trachygla- Recent research has revealed the degree of development of nis. the vertebral processes is more variable than previously Herein, we report on the discovery of the two new species suspected. In the course of a survey of specimens of Doumea, and provide a much needed redescription of Doumea conducted as the initial stage of a comprehensive study of chappuisi. Based on the shared and apparently derived the Doumeinae, our attention quickly focused on an presence of the extension of the caudal peduncular ridges

1 2944 N.E. Couch Street, Portland, Oregon 97232; E-mail: [email protected]. 2 South African Institute for Aquatic Biodiversity, Pvt Bag 1015, Grahamstown, 6140, South Africa; E-mail: [email protected]. 3 Division of Fishes, Smithsonian Institution, P.O. Box 37012, National Museum of Natural History, WG-14, MRC 159, Washington, D.C. 20013-7012; E-mail: [email protected]. Send reprint requests to this address. Submitted: 30 March 2010. Accepted: 9 August 2010. Associate Editor: D. Buth. F 2010 by the American Society of Ichthyologists and Herpetologists DOI: 10.1643/CI-10-050 706 Copeia 2010, No. 4 onto the surface of the skin, we treat these three species as a Doumea chappuisi Pellegrin, 1933 subgroup of Doumea that we term the D. chappuisi complex. Figures 1, 2; Table 1 This morphology of the vertebral processes is intermediate between the absence of surface ossifications, which is Doumea chappuisi Pellegrin, 1933:114, fig. on p. 115 [Type characteristic of the other species of Doumea, and the locality: Danane´, Coˆte d’Ivoire; syntypes MNHN 1932- possession of superficial bony plates in Andersonia, Belono- 0302, 1; NMBA 4281–4287, 7].—Daget, 1948:39 [Guineé glanis, Phractura, and Trachyglanis. We modify the current Fse, Mt. Nimba, Nzoˆ, 500 m].—Bertin and Este`ve, 1950:38 diagnosis of Doumea to accommodate the three species with [Type depository].—Daget, 1952:325, fig. 14 [Guinea, such restricted surface ossifications. Reserve naturelle inte´grale do Mont Nimba].—Lambert, 1958:49 [Guineé Français, Rivie`re Gbin].—Daget, 1962:109, fig. 36 [Kororo River, tributary of Komba River; MATERIALS AND METHODS description; comments on distribution].—Daget, Counts and measurements follow the methods outlined by 1963:585 [Guinea, Reserve naturelle inte´grale du Mont Skelton (1984, 2007a), with fin-ray counts taken both from Nimba].—Daget and Iltis, 1965:165, fig. on 165, map whole specimens and radiographs. Abbreviations in the text [Coˆte d’Ivoire; morphometrics and meristics].—Daget and are standard length (SL) and head length (HL). Institutional de Rham, 1970:803 [southern Coˆte d’Ivoire; Tabou].— abbreviations are as listed at http://www.asih.org/codons. Skelton and Teugels, 1986:61 [Summary synonymy; pdf, with the addition of SAIAB for the South African Guinea, Liberia, and Ivory Coast; maximum size].— Institute for Aquatic Biodiversity, Grahamstown. Kottelat and Sutter, 1988:54 [type depository].—Teugels et al., 1988:225, 229 [Cavally, Nipoue´, and Tabou river basins].—Burgess, 1989:114 [in listing of members of RESULTS Doumea].—Le´veˆque et al., 1989:110, 119 [Guineé Bissau, Corobal River].—Paugy et al., 1990:334, 339 [Liberia, Cess Doumea Sauvage, 1879 River].—Skelton, 1992:462, fig. 26.11 [Cavally and Gbin rivers in St. John basin of Liberia; Corubal basin in Doumea Species of are firm-bodied, streamlined, and tapered Guineé-Bissau).—Paugy et al., 1994:57 [freshwater, Cor- with pointed heads, long, slender, and depressed caudal ubal River basin].—Diogo, 2003:431 [within Doumea].— peduncles and large, falcate pectoral and pelvic fins Ferraris, 2007:25 [listing of species; type depositories; (Skelton, 2007a:58). The mouth is generally small with distribution].—Seegers, 2008:150 [comments on compar- thick papillose lips and tapered papillose barbels. The teeth ative features and distribution]. are either minute, vestigial, or absent. Vertebral elaborations Doumea chapuisi.—Harry, 1953:195 [based on original in Doumea lack distal expansions in the form of bony plates description of species by Pellegrin, 1933]. on the body surface contra the condition in Andersonia, Belonoglanis, Phractura, and Trachyglanis. In those genera, a Diagnosis.—Doumea chappuisi is distinguished from all series of plates run from lateral of the base of the dorsal fin species of Doumea, except D. reidi and D. stilicauda,by to the end of the caudal peduncle and from the area slightly having the dorsolateral and ventrolateral vertebral processes posterior of the origin of the pelvic fin, or even further extending through the skin to form longitudinal bony anteriorly, to the end of the caudal peduncle (Poll and ridges from the region of the base of adipose fin posteriorly Gosse, 1995:pl. 58, figs. 343–351). Nonetheless, the distal to the caudal-fin base and from slightly posterior of the portions of the vertebral processes of D. chappuisi, D. reidi, pelvic-fin origin to the caudal-fin base. Exposed processes new species, and D. stilicauda, new species, do penetrate are in the shape of longitudinal bars with slightly corrugated through the dermis to the body surface as a series of narrow surfaces that form irregular longitudinal ridges. Doumea ossifications that together form an irregular margin of an chappuisi differs from D. reidi in caudal-peduncle length (22– osseous ridge and exhibit a pattern similar to, but with the 29% SL, versus 31%, respectively), the posterior extent of the exposed distal portions of the vertebral processes distinctly adpressed pelvic fin in specimens over 90 mm SL (falling narrower than, Phractura, as illustrated in Poll and Gosse short of the anterior limit of the anal fin, versus extending (1995:pl. 58, fig. 345). The form of the vertebral extensions distinctly beyond that point, respectively), the anterior to the surface of the body in the species of the Doumea extent of the exposed vertebral processes along the ventral chappuisi complex, penetrating as they do to the body surface of the body (extending to slightly posterior of pelvic- surface, confronts us with two options: the creation of a new fin origin, versus extending to the area lateral to the anal-fin genus for these three species, or the modification of the base, respectively), and the pigmentation pattern of the definition of Doumea to include the presence of narrow dorsal surface of the unbranched rays of the pectoral and ossifications on the surface of the caudal peduncle. In- pelvic fins (uniformly pigmented, versus with series of asmuch as the generic level classification of the subfamily irregular dark spots against a lighter background, respec- Doumeinae has yet to be critically analyzed phylogenetical- tively). Doumea chappuisi differs from D. stilicauda in ly, it is quite possible that such a future study would predorsal length (32–37% SL, versus 27–30%, respectively), necessitate new generic level groupings with diagnoses that head length (18% SL, versus 15–17%, respectively), caudal- differ from those currently in use within the subfamily peduncle length (22–29% SL, versus 35–41%, respectively), (Harry, 1953; Poll and Gosse, 1995). As such, we defer from and caudal-peduncle depth (7.3–14.5 times in caudal- proposing a new generic name until one is shown to be peduncle length, versus 19.6–25.8 times, respectively). required by a phylogenetic analysis. In the interim, we modify the current diagnosis of Doumea to include all Description.—Body elongate and progressively tapering pos- species of the Doumeinae that lack distal expansions of the teriorly. Greatest body depth located slightly anterior of dorsolateral and ventrolateral processes of the caudal dorsal-fin origin and greatest width at anterior limit of vertebrae in the form of superficial interlocking plates. pectoral-fin insertion. Body smooth skinned and firm. Ferraris et al.—Doumea chappuisi complex 707

Fig. 1. Doumea chappuisi, 58.8 mm SL, MRAC 89-031-P-0028-0056, Cote d’Ivoire, Danane´, tributary to Cavally River on road between Danane´and Sipitou. Dorsal, lateral, and ventral views.

Middorsal ridge present between terminus of dorsal-fin base Head profile broadly triangular from dorsal view with and adipose-fin origin. Dorsal margins of neural arches lateral margin slightly convex and snout tip rounded. Head apparent through skin in lightly colored individuals, but not pointed and depressed from lateral view. Snout long, extending to body surface. Abdominal region transversely relatively slender, and tapering anteriorly. Eye situated flattened. Dorsal and ventral profiles of body progressively entirely within posterior half of head. Diameter of eye converge from anal-fin origin to posterior end of caudal approximately one-half of interorbital distance. Distance peduncle. Lateral line complete, running along midlateral from posterior naris to anterior margin of eye in smaller surface of body. Short, variably spaced and grouped, dorsal individuals less than distance between anterior naris and tip and ventral branches extending off main portion of lateral of snout; distances approximately equal in larger specimens. line. Exposed tips of dorsolateral vertebral processes apparent Posterior naris closer to anterior naris than to eye. Distance on skin surface from region of adipose-fin base posteriorly to between nares of each side slightly less than distance of each caudal-fin base. Processes on ventral surface of body extend to contralateral naris. Mouth subterminal, small, ovoid from slightly posterior of pelvic-fin origin to caudal-fin base. when open, with fleshy tuberculate upper lip and smooth Exposed processes in shape of longitudinal bars with slightly firm lower lip, flanked to each side by tuberculate sections. corrugated surfaces, forming irregular longitudinal ridge. Barbels short, tuberculate, and distally tapered. Maxillary Caudal peduncle relatively elongate, slender, and dorsoven- barbel longest, reaching approximately to vertical through trally depressed over much of its length, but approximately as middle of snout length. Inner mandibular barbel shortest, wide as high at caudal-fin base. Least depth of caudal with base situated slightly lateral of midline and of lateral peduncle approximately equal to length of eye. margin of smooth portion of lower lip. Outer mandibular 708 Copeia 2010, No. 4

fin base; fin small with length of base approximately one- third distance between posterior terminus of dorsal-fin base and adipose-fin origin; fin adnate, with posterior limit of fin situated posterior of vertical through anal-fin base, but not past posterior of vertical through tip of adpressed anal fin. Pectoral fin large and falcate with outer ray curved, pectinate, broad, and with fleshy pad covering ventral surface. Innermost pectoral-fin rays short, posterodorsally orientated, and adpressed to body wall. Tip of adpressed pectoral fin falling distinctly short of pelvic-fin origin and reaching to vertical through posterior terminus of dorsal-fin base. Pelvic fin large, slightly shorter than pectoral fin. Outer pelvic-fin ray pectinate with ventral surface covered by fleshy pad. Pelvic-fin origin located distinctly posterior of vertical through posterior end of dorsal-fin base. Tip of adpressed pelvic fin extending past anal-fin origin in smaller individuals, but falling short of that point in larger examined specimens. Anal fin large, with distal margin straight. Caudal fin forked, nearly symmetrical but with Fig. 2. Map of western central Africa, showing the distribution of ventral lobe slightly longer than dorsal lobe. Fin lobes small Doumea chappuisi (dots), D. reidi (triangle), and D. stilicauda (filled and acutely pointed. Middle rays of caudal fin slightly more square); some symbols represent more than one collecting locality than one-half length of longest rays. Dorsal-fin rays i,6,i, or or sample. i,7; pectoral-fin rays i,9, i,9,i, or i,10; pelvic-fin rays i,5; anal- fin rays iii (rarely iv),5–7,i; principal caudal-fin rays i,6,7,i barbel arises from angle of mouth and extends posteriorly (rarely i,7,6,i). approximately one-half length of branchiostegal membrane. Branchiostegal membrane continuous medially, with ven- Coloration in alcohol.—Body coloration of examined speci- tral portion of posterior margin straight or slightly concave. mens variable; ranging from light tan overall with scattered Gill slit extends posterodorsally to above pectoral-fin origin. small dark chromatophores through darker with distinct Humeral process located above origin of gill opening, short. pigment patterns on overall dark brown body. Pigmentation Supraoccipital process narrow, peg-like, separated by gap in all conditions darker dorsally with brown coloration from small trilobed nuchal shield. extending ventral of lateral line on abdomen and caudal Dorsal fin falcate, unbranched ray longest. Tip of peduncle, more so in darker individuals. Boundary between adpressed dorsal fin extends to vertical through middle of brown pigmentation of dorsal and dorsolateral regions and pelvic fin. Adipose-fin origin situated above middle of anal- lighter coloration of ventrolateral and/or ventral regions

Table 1. Proportional Measurements for the Species of the Doumea chappuisi Complex. Doumea chappuisi based on MRAC 89-031-P-0028-0056 and MNHN 1979-0121 (n = 13) and D. stilicauda on holotype and 12 paratypes. Values 1 to 10 are percentages of standard length; values 12 to 16 percentages of head length; and values 17 to 19 are cited ratios.

D. chappuisi D. reidi D. stilicauda Min–max Mean Holotype Min–max Mean Holotype Standard length (mm) 42–116 93 108–175 158 1 Predorsal length 32–37 35.1 33 27–30 28.3 28 2 Head length 19–23 20.9 18 15–17 16.7 16 3 Body depth at dorsal-fin origin 9–13 12.2 11 9–14 10.6 11 4 Body width at dorsal-fin origin 12–15 13.6 12 10–15 12.2 12 5 Caudal-peduncle length 22–29 26.3 31 35–41 37.9 37 6 Caudal-peduncle depth 2–3 2.9 2 1–2 1.9 2 7 Dorsal-fin length 19–24 20.6 20 17–20 18.3 19 8 Pectoral-fin length 21–27 24.1 23 19–23 20.4 19 9 Pelvic-fin length 16–23 20.1 20 16–20 18.0 19 10 Posterior limit of insertion pelvic-fin base to anus 5–9 7.7 8 6–11 7.8 9 11 Head depth 44–58 49.9 49 47–55 51.3 55 12 Head width 69–83 75.2 75 75–84 78.4 77 13 Snout length 52–63 57.8 64 59–65 62.1 62 14 Orbit diameter 12–16 14.6 12 12–16 13.8 15 15 Interorbital width 23–30 26.6 26 20–24 22.1 24 16 Postorbital length 27–33 29.4 31 29–34 33.0 30 17 Postorbital length/Snout length 0.45–0.61 0.51 0.49 0.48–0.57 0.53 0.48 18 Caudal-peduncle length/Caudal-peduncle depth 7.3–14.5 8.8 11.8 19.6–25.8 22.4 22.5 19 Body width/Body depth 1.0–1.4 1.1 1.1 1.0–1.4 1.2 1.1 Ferraris et al.—Doumea chappuisi complex 709 irregular albeit relatively distinct in specimens with well- Doumea reidi, new species developed darker coloration. Darker individuals with more Figures 2, 3; Table 1 intense pigmentation extending further ventrally and then Doumea sp. cf. thysi.—Reid, 1995:4, 17, 33, color plates 11, gradually becoming somewhat lighter. Specimens with 12 [Nigeria, Gashaka Gumti National Park, Mayo Melafa, distinct pigmentation patterns having irregular midlateral stream near Mayo Dande Village; Sanaga River basin; dark pigmentation continuous dorsally with ventral por- habitat and water conditions]. tions of four distinct saddles of dark pigmentation extending ventrally from middorsal line. First saddle extends Holotype.—USNM 337914, 93 mm SL, Nigeria, Gashaka longitudinally from supraoccipital-spine tip to beyond Gumti National Park, Mayo Dande Village, tributary of origin of dorsal fin and angles posteroventrally; second Banyo River, of Mbam River basin [Sanaga River drainage] saddle begins posterior of dorsal-fin base and terminates near border with Cameroon, 6u599000N, 11u399300E, approx- distinctly anterior of adipose-fin origin; third saddle origi- imately 1525 m asl, 13 April 1994, G. M. Reid. nates slightly anterior of anterior limit of adipose-fin base and extends posteriorly along two-thirds of fin base; fourth Diagnosis.—Doumea reidi is distinguished from all species of saddle begins somewhat posterior of adipose-fin base and Doumea, except D. chappuisi and D. stilicauda, by having the continues posteriorly for distance about equal to three dorsolateral and ventrolateral vertebral processes extending orbital diameters and continuous posteriorly with irregular through the dermis to form longitudinal bony ridges from dark middorsal stripe. Combination of dark brown mid- the base of the adipose fin posteriorly to the caudal-fin base lateral stripe and dark brown saddles enclose more lightly and from slightly posterior of the anal-fin origin to the colored saddles. Ventrolateral portion of abdomen and caudal-fin base. The exposed processes are in the shape of ventral part of caudal peduncle with scattered dark chro- longitudinal bars with slightly corrugated surfaces that matophores, but chromatophores absent on ventral portion together form an irregular longitudinal ridge. Doumea reidi of abdomen even in darker examined specimens. Lateral- differs from D. chappuisi in caudal-peduncle length (31% SL, line canal variably paler than background coloration, more versus 22–29%, respectively), the posterior extent of the obviously so posteriorly in many specimens. adpressed pelvic fin in specimens over 90 mm SL (pelvic fin Dorsal and lateral portions of head brown, with scattered extending distinctly beyond anterior limit of the anal fin, darker regions throughout. Ventral part of head tan with versus falling short of that point), the anterior extent of the fine darker pigmentation ventrolaterally in many individu- exposed vertebral processes along the ventral surface of the als. Maxillary barbel dusky, especially dorsally. Mandibular body (extending to the area lateral to the anal-fin base, barbels unpigmented. versus extending to slightly posterior of pelvic-fin origin, Dorsal surfaces of leading unbranched pectoral- and pelvic- respectively), and the pigmentation pattern of the dorsal fin rays ranging from tan to brown, without any distinct surface of the unbranched rays of the pectoral and pelvic coloration pattern. Dorsal surface of both fins darker brown fins (with a series of irregular dark spots against a lighter basally and with medially narrowing band of dark brown background, versus uniformly pigmented, respectively). pigmentation on distal half of rays, but pigmentation falls Doumea reidi differs from D. stilicauda in predorsal length short of distal margin. Ventral surfaces of paired fins, (33% SL, versus 27–30%, respectively), head length (18% SL, particularly fleshy basal pad, more lightly pigmented, but versus 15–17%, respectively), caudal-peduncle length (31% with darker pigmentation of dorsal surface of rays apparent to SL, versus 35–41%, respectively), and caudal-peduncle depth varying degrees. Dorsal-fin rays pale to dusky, and with broad, (11.8 times in caudal-peduncle length, versus 19.6–25.8 dark patch on distal portions of anterior rays. Some individuals times, respectively). with dark brown band on basal portions of posterior fin rays. Description.—Body elongate and progressively tapering pos- Distribution.—Examined specimens of Doumea chappuisi teriorly. Greatest body depth located slightly anterior of originated in the coastal river basins of western Africa (Cote dorsal-fin origin and greatest width at anterior limit of d’Ivoire, Guinea, Liberia; Fig. 2). Le´veˆque et al. (1989:110, pectoral-fin insertion. Head and body with numerous small 119) reported the species from the portion of the Corubal papillae; papillae distributed over lateral and dorsal portions River in Guinea Bissau. The range of the species includes, of head. Ventral surface of head with close-set papillae but probably is not limited to, the Goˆ, Cavally, St. John’s, arranged in irregular rows. Rows diverging laterally from and Corubal river basins. Doumea chappuisi was the only region somewhat posterior of lower lip. Papillae also present species of Doumea found among the examined specimens of on lateral and dorsal surfaces of anterior portion of body. the genus from this region. Middorsal ridge present between terminus of dorsal-fin base and adipose-fin origin. Dorsal margins of neural arches Material examined.—FMNH 50281, 2, 51–60 mm SL, Liberia, apparent through skin, but not extending to body surface. stream near Mount Nimba. USNM 193950, 3, 33–43 mm SL, Abdominal region transversely flattened. Dorsal and ventral Liberia, Gbarnga District, St. John’s River tributary. MNHN profiles of body progressively converge from anal-fin origin 1970-0015, 1 (of 2), 53 mm SL, Cote d’Ivoire, Tabou. MNHN to posterior end of caudal peduncle. Exposed tips of 1979-0121, 3 (of 6), 82–116 mm SL, Cote d’Ivoire, Troko- dorsolateral vertebral processes apparent on skin surface limpleu, Cavally River basin. MRAC 89-031-P-0028–0036, from region of base of adipose fin posteriorly to caudal-fin 10, 42–61 mm SL. Cote d’Ivoire, Danane´, tributary to base. Processes on ventral surface of body extend from Cavally River. MRAC 77-44-P-131–132, 2, 50–51 mm SL, slightly posterior of anal-fin origin to caudal-fin base. Guinea, Guineé forestıère, Gbin River. MRAC 119500, 1, Exposed processes in shape of longitudinal bars with slightly 36 mm SL, Guinea, Guineé forestıère, Gbin River. MNHN corrugated surfaces that jointly form irregular longitudinal 1982-1191, 2 (of 4), 62–64 mm SL, Guinea, Gaoual, upper ridge. Lateral line complete and runs along midlateral Corubal River basin, Tomine River. surface of body. Short, variably spaced and grouped dorsal 710 Copeia 2010, No. 4

Fig. 3. Doumea reidi, new species, holotype, 93 mm SL, USNM 337914, Nigeria, Mayo Dande Village, Mayo Melafa, tributary of Banyo River, Mbam River basin [Sanaga River drainage] near border with Cameroon, 6u599000N, 11u399300E. Dorsal, lateral, and ventral views. and ventral branches extend off from main portion of lateral Distance between nares of each side slightly less than line. Caudal peduncle relatively elongate, slender, and distance of each to contralateral naris. Mouth subterminal, dorsoventrally depressed over much of its length, but small, ovoid when open, with fleshy tuberculate upper lip approximately as wide as high at caudal-fin base. Least and smooth firm lower lip, flanked to each side by depth of caudal peduncle approximately equal to two-thirds tuberculate sections. Barbels short, tuberculate and distally length of eye. tapered. Maxillary barbel longest, reaching approximately to Head broadly triangular from dorsal view with lateral vertical through middle of snout length. Inner mandibular margin very slightly convex and snout tip rounded. Head barbel shortest, with base located slightly lateral of midline pointed and depressed from lateral view. Snout long, and of lateral margin of smooth portion of lower lip. Outer relatively slender, and anteriorly tapering. Eye situated mandibular barbel arises from angle of mouth and extends entirely within posterior half of head. Diameter of eye posteriorly approximately one-half length of branchiostegal approximately one-half of interorbital distance. Distance membrane. Branchiostegal membrane continuous medially from posterior naris to anterior margin of eye approximately with ventral portion of posterior margin slightly concave. equal to distance between anterior naris and tip of snout. Gill slit extends posterodorsally to above pectoral-fin origin. Posterior naris distinctly closer to anterior naris than to eye. Humeral process located above origin of gill opening, short. Ferraris et al.—Doumea chappuisi complex 711

Supraoccipital process narrow, peg-like, separated by gap with darker pigmentation of dorsal surface of rays apparent from small trilobed nuchal shield. to limited degrees, more so in case of pelvic fin. Dorsal-fin Dorsal fin falcate, unbranched ray longest. Tip of rays pale with some darker brown areas on distal portions of adpressed dorsal fin extends to vertical through middle of anterior rays. Adipose fin with dusky, posterodorsally angled pelvic fin. Adipose-fin origin situated above middle of anal- band on central portion bordered by paler regions anteriorly fin base; fin small with length of base approximately one- and posteriorly. Caudal fin with scattered dark spots, more third of distance between posterior terminus of dorsal-fin so on lower lobe. Fleshy pad overlying base of caudal-fin rays base and adipose-fin origin; fin adnate, with posterior extent with dark spots. Anal-fin rays dusky, with scattered darker of fin extending past vertical through posterior limit of anal- spots, particularly on distal one-third of anterior rays. fin base, but not posterior of vertical through tip of adpressed anal fin. Pectoral fin large and falcate with outer Coloration in life.—Color plates in Reid (1995:pls. 11, 12) ray curved, pectinate, broad, and with fleshy pad covering show that coloration is overall comparable to that of ventral surface. Innermost pectoral-fin rays short, poster- preserved specimen other than that lighter areas are more odorsally orientated, and adpressed to body wall. Tip of cream colored in life. adpressed pectoral fin falling distinctly short of pelvic-fin origin and reaching to vertical through posterior terminus Distribution and reproduction.—Doumea reidi is known only of dorsal-fin base. Pelvic fin large, slightly shorter than from a single specimen from the type locality in eastern pectoral fin. Outer pelvic-fin ray pectinate with ventral Nigeria, not far from the border with Cameroon (Fig. 2). The surface covered by fleshy pad. Pelvic-fin origin located type locality is a stream close to the Village of Mayo Dande distinctly posterior of vertical through posterior end of and drains to the Banyo River and then the Mbam River of dorsal-fin base. Tip of adpressed pelvic fin extending well the Sanaga River basin (Reid, 1995:33). The holotype has an past anal-fin origin. Anal fin large, with distal margin extensive series of papillae on the head and anterior portion straight. Caudal fin forked and nearly symmetrical but with of the body comparable in form to those present in one of ventral lobe slightly longer than dorsal lobe. Lobes small the paratypes of D. stilicauda (see comments under species and sharply pointed with middle rays of caudal fin account). The specimen of D. stilicauda was a ripening approximately one-half length of longest rays. Dorsal-fin female and the presence of the papillae in the holotype of D. rays i,6; pectoral-fin rays i,10; pelvic-fin rays i,5; anal-fin rays reidi may indicate that it was entering into the breeding iii,8; principal caudal-fin rays i,6,7,i. season when collected.

Coloration in alcohol.—Body with tan ground coloration, Habitat.—The type locality of Doumea reidi is a cool, briskly overlain with brown dorsally. Brown coloration extends flowing, upland stream less than 4 m wide and less than somewhat ventral of lateral line across abdomen, but limited 1.5 m maximum depth (Reid, 1995:34). The substratum was to region dorsal of lateral line on caudal peduncle. Boundary reported to consist of large, granitic boulders interspersed between brown and tan pigmentation distinct. Brown among deposits of coarse sand, mud, and leaf litter. The pigmentation overlain with irregularly sized and placed type locality was the only location in the Gashaka Gumti darker spots of varying sizes and shapes. Midlateral portion National Park where the genus was recorded; however, of body with irregular stripe of dark pigmentation; stripe most fish collections made during the ichthyological more obvious anteriorly. Dorsal and dorsolateral portions of survey of the park were from the Benue River system, body with four distinct, dark brown middorsal saddles which flows west through Nigeria before emptying into the continuous ventrally with midlateral stripe. Anterior saddle Niger River. extends from tip of supraoccipital-spine to middle of dorsal fin; second saddle extends from slightly posterior of dorsal- Etymology.—The species name, reidi, honors Dr. Gordon fin base about three-quarters of distance to adipose-fin McGregor Reid, of the North of England Zoological Society, origin, but only reaches midlateral stripe along its posterior who collected the holotype and who has dedicated a large region; third saddle begins somewhat anterior to adipose-fin portion of his career helping to protect, and improve our origin and extends along much of base of fin and angles understanding of, wildlife and freshwater fishes worldwide. anteroventrally to midlateral stripe; small fourth saddle starts slightly posterior of adipose-fin base. Ventrolateral Remarks.—In addition to Doumea reidi, one other species of portion of abdomen and ventral part of caudal peduncle the genus is known from the Sanaga River, that being D. covered with scattered dark chromatophores, but with sanaga, an endemic of the portions of that river system in chromatophores absent ventrally on anterior part of abdo- Cameroon (Skelton, 2007a, 2007b). Doumea reidi differs men. Lateral line canal pale for entire length. from D. sanaga in the degree of extension of the dorsolateral Dorsal and lateral portions of head brown, with scattered and ventrolateral projections of the vertebrae on the middle slightly darker spots throughout. Ventral part of head tan and posterior portions of the body (reaching through skin overall, with fine darker pigmentation ventrolaterally but and forming bony ridges along surface, versus covered by lacking chromatophores ventrally. Maxillary barbel dusky, skin), in the overall head and body form (compare lateral more so dorsally. Mandibular barbels unpigmented. view of Fig. 3 with Skelton, 2007a:fig. 11), the length of the Dorsal surface of pectoral and pelvic fins with scattered predorsal portion of the body (33% SL, versus 34–37%, dark spots on dorsal surface of unbranched rays that form respectively), the head depth (49% HL, versus 51–60%, distinct, but irregular pattern. Dorsal surface of branched respectively), the orbital diameter (12% SL, versus 13–18%, rays of both fins darker basally and with dark pigmentation respectively), and the pigmentation pattern of the dorsal in form of medially narrowing band on distal half of rays, surface of the unbranched rays of the pectoral and pelvic but with band falling short of distal margin. Ventral surface fins (with distinct dark marks against a light background, of fins, particularly fleshy basal pad, lightly pigmented, but versus uniformly pigmented, respectively). 712 Copeia 2010, No. 4

Fig. 4. Doumea stilicauda, new species, holotype, 158 mm SL, SAIAB 80182, Cameroon, So’o River, tributary of Nyong River, at Pont So’o, 3u199N, 11u299E. Dorsal, lateral, and ventral views.

Doumea stilicauda, new species 153 mm SL, March 2008. SAIAB 80798, 1, 144 mm SL, May Figures 2, 4; Table 1 2008.

Diagnosis.—Doumea stilicauda is distinguished from all Holotype.—SAIAB 80182, 158 mm SL, Cameroon, Sud species of Doumea, except D. chappuisi and D. reidi,by Region, Pont So’o, So’o River, Nyong River system, from having the dorsolateral and ventrolateral vertebral processes indigenous fish trap along river channel, 3u199N, 11u299E, extending through the dermis to form longitudinal bony April 2007, R. Etoundi Owona and R. E. Brummett. ridges from slightly anterior to the base of the adipose fin posteriorly to the caudal-fin base and from somewhat Paratypes.—All collected at type locality by same collectors anterior of the anal-fin origin to the caudal-fin base. and method: SAIAB 85547 (formerly SAIAB 80182, in Exposed processes are in shape of longitudinal bars with part), 1, 155 mm SL, collected with holotype. SAIAB slightly corrugated surfaces that together form irregular 80791, 6, 107.5–162 mm SL; SAIAB 80792, 2, 119–175 mm longitudinal ridges. Doumea stilicauda is readily distinguish- SL; USNM 396012 (formerly SAIAB 80791, in part), 2, 138– able from D. chappuisi in predorsal length (27–30% SL, Ferraris et al.—Doumea chappuisi complex 713 versus 32–37%, respectively), head length (15–17% SL, approximately to vertical through middle of snout length. versus 19–23%, respectively), caudal-peduncle length (35– Inner mandibular barbel shortest, with base located slightly 41% SL, versus 22–29%, respectively), and caudal-peduncle lateral of midline and of lateral margin of smooth portion of depth (19.6–25.8 times in caudal-peduncle length, versus lower lip. Outer mandibular barbel arises from angle of 7.3–14.5 times, respectively). Doumea stilicauda is readily mouth and extends posteriorly approximately one-half distinguishable from D. reidi in predorsal length (27–30% SL, length of branchiostegal membrane. Branchiostegal mem- versus 33%, respectively), head length (15–17% SL, versus brane continuous medially with central region of posterior 18%, respectively), caudal-peduncle length (35–41% SL, margin slightly concave. Gill slit extends posterodorsally to versus 22–29%, respectively), and caudal-peduncle depth above pectoral-fin origin. Humeral process above origin of (19.6–25.8 times in caudal-peduncle length, versus 11.8 gill opening, short. Supraoccipital process narrow, peg-like, times, respectively). separated by gap from small trilobed nuchal shield. Dorsal fin falcate, unbranched ray longest. Tip of Description.—Body elongate and progressively tapering pos- adpressed dorsal fin extends to vertical through middle of teriorly. Greatest body depth located slightly anterior of pelvic fin. Adipose-fin origin situated above middle of anal- dorsal-fin origin other than in specimens with extended fin base; fin small with length of base approximately one- abdomen, in which greatest depth at middle of abdomen. third of distance between posterior terminus of dorsal-fin Greatest body width at anterior limit of pectoral-fin base and adipose-fin origin; fin adnate, with posterior extent insertion other than in specimens with extended abdomen, of fin extending past vertical through posterior limit of anal- in which greatest width at middle of abdomen. Head and fin base, but not posterior of vertical through tip of body of holotype covered with numerous very small adpressed anal fin. Pectoral fin large and falcate with outer papillae. Papillae distributed over lateral and dorsal portions ray curved, pectinate, broad, and with fleshy pad covering of head. Ventral surface of head with small, apparently ventral surface; innermost pectoral-fin rays short, poster- developing, close set papillae arranged in irregular rows odorsally orientated, and adpressed to body wall other than diverging from region somewhat posterior of lower lip; when fin fully extended. Tip of adpressed pectoral fin falling papillae in this region most obvious posteriorly. Small distinctly short of pelvic-fin origin and reaching nearly to or papillae also present on lateral and dorsal surfaces of slightly beyond vertical through posterior terminus of anterior portion of body. Middorsal ridge present between dorsal-fin base. Pelvic fin large, slightly shorter than pectoral terminus of dorsal-fin base and adipose-fin origin. Dorsal fin, outer ray pectinate with ventral surface with fleshy pad. margins of neural arches apparent through skin, but not Pelvic-fin origin located distinctly posterior of vertical extending to body surface. Abdominal region transversely through posterior end of dorsal-fin base. Tip of adpressed flattened other than in specimens with distended abdomens pelvic fin extending slightly beyond anal-fin origin. Anal fin in which abdomen somewhat transversely convex. Dorsal large, with distal margin straight. Caudal fin forked and and ventral profiles of body progressively converge from nearly symmetrical but with ventral lobe slightly longer anal-fin origin to posterior end of caudal peduncle. Exposed than dorsal lobe. Lobes small and sharply pointed with tips of dorsolateral vertebral processes apparent on skin middle rays of caudal fin slightly less than one-half length of surface from region slightly anterior of base of adipose fin longest rays. Dorsal-fin rays i,6; pectoral-fin rays i,10 posteriorly to caudal-fin base. Processes on ventrolateral (including holotype) or i,11; pelvic-fin rays i,5; anal-fin rays surface of body extend from somewhat anterior of anal-fin iii (rarely ii or iv),6; principal caudal-fin rays i,7,6,i. origin to caudal-fin base. Exposed processes in shape of longitudinal bars with slightly corrugated surfaces that form Coloration in alcohol.—Body with tan to light brown ground irregular longitudinal ridge. Lateral line complete, running coloration and distinctly brown dorsally and dorsolaterally. along midlateral surface with short, variably spaced and Brown coloration extends somewhat ventral of lateral line grouped dorsal and ventral branches extending from main across abdomen. Boundary between brown and tan pigmen- portion of system. Caudal peduncle relatively elongate, tation distinct. Brown pigmentation overlain with irregu- slender, and dorsoventrally depressed over much of its larly sized and placed darker spots of varying sizes and length, but approximately one and one-half times wide as shapes with enclosed variably present light regions. Some high at caudal-fin base. Least depth of caudal peduncle specimens with irregular stripe of dark pigmentation along approximately equal to two-thirds length of eye. midlateral portion of body; stripe more obvious anteriorly. Head broadly triangular from dorsal view with lateral Dorsal and dorsolateral portions of body with four, variably margin very slightly concave to anterior margin of opercle obvious, distinct, dark brown saddles extending ventrally and then slightly convex to posterior limit of opercle and from middorsal region. Anterior saddle extends from tip of snout tip rounded. Head pointed and depressed from lateral supraoccipital spine to middle of dorsal fin; second saddle view. Snout long, relatively slender, and anteriorly tapering. extends from slightly posterior of dorsal-fin base about Eye situated entirely within posterior half of head. Diameter three-quarters of distance to adipose-fin origin; third saddle of eye slightly less than one-half of interorbital distance. begins somewhat anterior to adipose-fin origin and extends Distance from posterior naris to anterior margin of eye along much of base of; small fourth saddle starts slightly distinctly less than distance between anterior naris and tip posterior of adipose-fin base. Ventrolateral portion of of snout. Posterior naris distinctly closer to anterior naris abdomen with scattered dark chromatophores in some than to eye. Distance between nares of each side slightly less specimens, but with chromatophores absent ventrally on than distance of each to contralateral naris. Mouth subter- anterior part of abdomen. Lateral line canal irregularly more minal, small, ovoid when open, with fleshy tuberculate lightly pigmented than surrounding regions in some upper lip and smooth firm lower lip, flanked to each side by specimens. Distinct, narrow, dark stripe extending from tuberculate sections. Barbels short, tuberculate, and distally somewhat posterior of terminus of pelvic-fin base posterior tapered. Maxillary barbel longest, reaching posteriorly to above base of anal fin. 714 Copeia 2010, No. 4

Fig. 5. Type locality of Doumea stilicauda. Indigenous fishing fence on So’o River, Nyong River basin, Cameroon, in high water conditions. Photograph by P. Skelton, June 2008.

Dorsal and lateral portions of head brown, with scattered tributary of the Nyong River, local fishers constructed a slightly darker spots throughout. Ventral part of head tan wooden palisade barrier dam with a chute basket traps overall, with some fine darker pigmentation ventrolaterally known as an ‘‘alam’’ (Fig. 5). This indigenous trap was but lacking chromatophores ventrally. Maxillary barbel effective in catching a wide variety of fish species, some of dusky, especially dorsally. Mandibular barbels pale, without which were previously unknown from the Nyong River dark chromatophores. system. Included among these were the samples of Doumea Dorsal surface of pectoral fin dark brown with variably stilicauda that form the basis of the species description. developed lighter band extending from midsection of anterior rays to distal portion of posterior rays. Distal Distribution.—Doumea stilicauda is known only from the So’o margin of fin hyaline. Ventral surface of fin with dusky River, a tributary of the Nyong River, Cameroon (Fig. 2). band extending across fin approximately three-quarters of distance from base, but with distal portion of fin hyaline. Etymology.—The species name, stilicauda, from the Latin Dorsal surface of pelvic fin dusky other than for distal stilus, a stake, and cauda, tail, refers to the stake-like caudal margin and basal portion of middle rays. Ventral surface of peduncle; treated as a noun. fin with dusky band extending across fin approximately three-quarters of distance from base, but with distal portion ACKNOWLEDGMENTS of fin hyaline. Dorsal-fin rays dusky overall, more so on Support for this project to CJF was provided by the All Catfish unbranched ray and on rays and membranes distally. Species Inventory, a program funded by the National Science Adipose and caudal fins dusky. Fleshy pad overlying base Foundation (DEB-0315963); to PHS by M. Stiassny for support of caudal-fin rays darkly pigmented. Anal-fin rays dusky of visits to AMNH to investigate the amphiliids of West- over distal one-third of anterior rays, but with margin Central Africa; and to RPV by the Herbert R. and Evelyn hyaline. Some specimens fixed in alcohol show bluish gray Axelrod Chair in Systematic Ichthyology, in the Division of color with spots evident on dorsal head and body. Fishes, National Museum of Natural History of the Smithso- nian Institution. R. Brummett, WorldFish Centre, Yaounde´, Sexual dimorphism and reproduction.—Anus and genital pore Cameroon, generously made available the specimens that in males of the species located midway between base of served as the basis for the description of Doumea stilicauda pelvic fins and anterior edge of pelvic-fin origin, but with and hosted PHS at the holotype locality. We thank M. Rogers females having openings slightly closer to pelvic-fin base. A (FMNH), P. Pruvost (MNHN), and J. Snoeks (MRAC) for the single specimen (SAIAB 80798) collected in May 2008, later loan of specimens. S. Raredon (USNM) provided radiographs in the yearly cycle than the other samples of the species, is a of specimens and other assistance during the study. Figures 1, female with ripening ovaries and tubercles covering the 3, and 4 were prepared by T. Griswold; Figure 2 was prepared dorsal surface of the head and body indicating that the by W. Coetzer (SAIAB). reproductive season follows at some later time period. LITERATURE CITED Habitat.—The Nyong River is a ‘‘blackwater’’ rainforest system with low pH waters (mean pH 6.2; Brummett et al., Bertin, L., and R. Este`ve. 1950. Catalogue des Types de 2010). At one study site along the So’o River, a large forest Poissons du Muse´um National d’Histoire Naturelle; 5e Ferraris et al.—Doumea chappuisi complex 715

partie: Ostariophysaires (Siluriformes). Impress Nationale, Liberia: annotated checklist and distribution. Revue Paris. d’Hydrobiologie Tropicale 23:329–350. Brummet, R. E., D. Nguenga, F. Tiotsop, and J.-C. Abina. Paugy, D., K. Traore´, and P. S. Diouf. 1994. Faune 2010. The commercial fishery of the middle Nyong River, ichtyologique des eaux douces d’Afrique de l’Ouest, Cameroon: productivity and environmental threats. p. 35–66. In: Biological Diversity of African Fresh- and Smithiana Bulletin 11:3–16. Brackish Water Fishes. G. G. Teugels, J. F. Gue´gan, and J. J. Burgess, W. E. 1989. An Atlas of Freshwater and Marine Albaret (eds.). Annales, Sciences Zoologiques, Musee´Royal Catfishes. A Preliminary Survey of the Siluriformes. T.F.H. de l’Afrique Centrale 275:1–177. Publications, Neptune City, New Jersey. Pellegrin, J. 1933. Voyage de Ch. Alluaud et P. A. Chappuis Daget, J. 1948. La collection de poissons d’eau douce de en Afrique occidentale Française (Dec. 1930–Mars 1931). l’I.F.A.N. Catalogues de Institute Français d’Afrique Noire IV.—Poissons. Archiv fu¨r Hydrobiologie 26:101–120. 3:1–59. Poll, M., and J.-P. Gosse. 1995. Genera des poissons d’eau Daget, J. 1952. La re´serve naturelle inte´grale du Mont douce de l’Afrique. Me´moire de la Classe des Sciences, Nimba. Poissons. Me´moires de Institute Français d’Afrique troisieme se´rie 9:1–324. Noire 19:311–334. Reid, G. M. 1995. The Fishes of the Gashaka Gumti National Daget, J. 1962. Les poissons du Fouta Dialon et de la Basse Park, Nigeria: Taxonomy, Ecology, Resource Utilisation Guineé. Me´moires de Institute Français d’Afrique Noire and Conservation. Nigerian Conservation Foundation, 65:1–210. World Wide Fund for Nature, U.K. Daget, J. 1963. La re´serve naturelle inte´grale du Mont Seegers, L. 2008. Die Welse Afrikas. Ein Handbuch fu¨r Nimba. XXVII. Poissons (Deuxie`me note). Me´moires de Bestimmung und Pflege/The Catfishes of Africa, a Hand- Institute Français d’Afrique Noire 66:573–600. book for Identification and Maintenance (English edi- tion). Tetra Verlag, Berlin-Velten. Daget, J., and P. de Rham. 1970. Sur quelques poissons du Skelton, P. H. 1984. A systematic revision of species of the Sud de la Coˆte d’Ivoire. Revue Suisse de Zoologie catfish genus Amphilius (Siluroidei, Amphiliidae) from east 77:801–806. and southern Africa. Annals of the Cape Provincial Daget, J., and A. Iltis. 1965. Poissons de Coˆte d’Ivoire (eaux Museums (Natural History) 16:41–71. douces et saumaˆtres). Me´moires de Institute Français Skelton, P. H. 1992. Amphiliidae, p. 451–467. In: Faune des d’Afrique Noire 74:1–385. Poissons d’Eaux Douces et Saumaˆtres de l’Afrique de Diogo, R. 2003. Anatomy, phylogeny, and taxonomy of l’Ouest, Vol. 2. C. Le´veˆque, D. Paugy, and G. G. Teugels Amphiliidae, p. 401–438. In: Catfishes. G. Arratia, B. G. (eds.). Museé Royal de l’Afrique Central, Tervuren, and Kapoor, M. Chardon, and R. Diogo (eds.). Science Editions de l’ORSTOM, Paris. Publishers, Enfield, New Hampshire. Skelton, P. H. 2007a. New species of amphiliid catfish Ferraris, C. J., Jr. 2007. Checklist of catfishes, Recent and genera Amphilius, Doumea and Phractura and the taxono- fossil (Osteichthyes: Siluriformes), and catalogue of siluri- my of Paramphilius from West Africa (Siluriformes, form primary types. Zootaxa 1418:1–628. Amphiliidae). Zootaxa 1578:41–68. Harry, R. R. 1953. A contribution to the classification of the Skelton, P. H. 2007b. Amphiliidae, p. 753–789. In: Poissons African catfishes of the family Amphiliidae, with descrip- d’Eaux Douces et Saumaˆtres de Basse Guineé, Oueste de tion of collections from Cameroon. Revue de Zoologie et l’Afrique Centrale. The Fresh and Brackish Water Fishes of de Botanique Africaines 47:177–232. Lower Guinea, West-Central Africa. [Collection Faune et Kottelat, M., and E. Sutter. 1988. Catalogue des types de Flore Tropicales 42], Vol. 1. M. L. J. Stiassny, G. G. Teugels, poissons du Museé d’histoire naturelle de Baˆle (Naturhis- and C. D. Hopkins (eds.). Institut de Recherche´ pour le torisches Museum Basel). Verhandlungen der Natur- De´veloppement, Paris, Museum National d’Histoire Nat- forschenden Gesellschaft in Basel 98:51–57. urelle, Paris, and Museé Royal de l’Afrique Centrale, Lambert, J. G. 1958. Poissons Siluriformes et Cyprinodonti- Tervuren. formes rećolte´s en Guineé Française, avec la description Skelton, P. H., and G. G. Teugels. 1986. Amphiliidae, d’une nouvelle espe`ce de Microsynodontis. Revue de p. 54–65. In: Check-list of the Freshwater Fishes of Africa, Zoologie et de Botanique Africaines 57:39–56. Volume 2. J. Daget, J.-P. Gosse, and D. F. E. Thys van den Le´veˆque, C., D. Paugy, G. G. Teugels, and R. Romand. Audenaerde (eds.). ISNB, Bruxelles; MRAC, Tervuren; and 1989. Inventaire taxonomique et distribution des poisons ORSTOM, Paris. d’eau douce des bassins coˆtiers de Guineé et de Guineé Teugels, G. G., C. Le´veˆque, D. Paugy, and K. Traore´. 1988. Bissau. Revue d’Hydrobiologie Tropicale 22:107–127. E´tat des connaissances sur la faune ichtyologique des Paugy, D., C. Le´veˆque, G. G. Teugels, R. Bigorne, and R. basins coˆtiers de Coˆte d’Ivoire et de l’Ouest du Ghana. Romand. 1990. Freshwater fishes of Sierra Leone and Revue d’Hydrobiologie Tropicale 22:221–237.