25. Neoperla camerunensis (Enderlein, 1909)

(Figs. 112, 126, 131–132, 143–149)

Ochthopetina camerunensis Enderlein, 1909b: 343, figs 21, 23.

not Ochthopetina camerunensis — Navás 1912: 403, fig. 2; see N. africana !

Neoperla camerunensis — Zwick 1973a: 502, figs. 70, 71; new combination, lectotype designation, redescription.

Ochthopetina haugi Navás, 1916: 51, fig. 29; new synonymy.

Type material studied: ♀ lectotype O. camerunensis: Kamerun Barombi Conradt \ Type [red] \ Neoperla camerunensis Enderl . ♀ Type [in Enderlein’s hand] det. Dr. Enderlein [print] \ Lectotypus design. P.Zwick 1971 \ Mus. Zool. Polonicum Warszawa 12/45 \ Mus. Zool. Polonicum Warszawa Typus No. 1295; 2♀ paralectotypes, Type numbers 1296 and 1300, respectively with the same data (all in MIZ; pinned genitalia and egg slides on Zelluloid, on the specimen pin).Additional paralectotypes (No. 1296–1299, labelled “Kamerun, Barombi, Conradt”) are misassociated or too damaged for identification, see Notes below .

Holotypus N. haugi: Gabonese Republic: Bas Ogooué entre Lambaréné et la mèr E. HAUG 1901 \ Ochthopetina haugi ♂ Nav. [in Navás’ hand] Navás S.J. det. [green print] \ Type (MNHN; pinned, genitalia on pinned microslide).

Additional material. Republic of Angola: 1♀, Angola,Dundo, Piège lumineux VIII.53 Carvalho (ex coll. Brinck; slide Z18.04). Republic of Cameroon: 1♀: Kamerun, Mukonje 28.3.–4.4.38 Buhr S.G. (MfNB). 2♀, 6°8’26.25’’N, 10°6’3-68’’E, NW Reg., Bamenda, Mentchum River contributor 15 km NNW Bafut, 630m asl, 9 Nov.2011, MV-lamp,A.Zwick (SMNS; NEOP 127, NEOP 128); 1♂: Cameroon, Ekok [5.81N, 8.85E] 24 mi E Tekmo, 650m, 12.X.1966, Ross & Lorenzen (CASENT 8413093; Z18.52). 1♀, Adamana (Camerun) [red ink] \ africana Klapàlek [print] \ det. P. Zwick 1988 Neoperla sp. 9 [handwriting on photoprint] (genital and egg preparation on glass slide; NMCZ; old egg slide in SMNS). 2♂, Makak, 14–26-XI-1972 filtered black light, J.A.Gruwell (USNM _ 82A, 86); Libamba [3.55°N, 11.02°E], 10km E of Makak (filtered) black light, J.A.Gruwell: 1♂, 15–17-IX.1973; 1♂, 6–10-XII-1973; 2♂, 10–11-I-1974; 1♂, 2♀, 1–2.iii.1974; 1♂, 19–22-IV-1974 (USNM, in ETOH, genitalia in microvial or on slide, duplicates also SMNS). 1♂, en route Douda Yaounde FrCam W.Afr. 36 \ Van Zwaluvenburg & McGough (USNM _27). Democratic Republic of the Congo: 1♀, Coll. Mus. Congo Ruwenzori W. Mutwanga [29.40E, 0.20N] IV 1937 Lisfranc (MRAC); 2♂, Congo belge: P.N.U. [Upemba] Kaswabilenga (700m) 17-X-1947 Mis. G.F.de Witte, 845a \ Coll. Mus. Congo (ex coll. I.P.N.C.B.) \ Neoperla spio (Newman) Dr. N.Hynes det., 1951 (SMNS, ex MRAC). 2♂, B. Congo, 39 km S of Walikale [1.47S, 28.07E], 700m, XII-25-57 E.S.Ross & R.E.Leech (CASENT 8413090 & 91; Z18.48 & 49); 1♂: B.Congo, 23 mi E of Kama [3.52S, 27.12E], 16.8.57, Ross & Leech (CASENT 8413094, Z18.53); 6♂: B. Congo, 12 ml. N of Matadi [5.85N, 13.46E] VII-28-1957 E.S.Ross & R.E.Leech (CASENT 8413095; Z18.54). 1♀: Coll. Mus. Tervuren Tschela III-1964 (Fain) coll. H. Schouteden; 1♂: Musée du Congo Boma [-5.850, 13.071] 1933 (Dr. Hallet) (both MRAC). 1♂: B.Congo, 50km S of Tshela [5.01S, 12.96E], 26.7.57, Ross & Leech (CAS). 1♀: Congo 149 Lámpázáz 11. Feb. 1988. Manyara National Park, camp site (slide Z19.78; HMNH). 1, Congo 149 Lámpázáz 11. Feb. 1988. Manyara National Park, camp site (slide Z19.78); 1, Republic of Kenya, Lake Turkana, Sibiloi National Park [3.92N, 36.18], Koobi Fora Base Camp 14.3.1988 (Slide Z19.76, both HNHM). Federal Democratic Republic of Ethiopia: 9 ♀, Ghibe River 13-14/5-61 215 km from Adis Ababa wooden bridge: road: track to The Crown Prince’s Saw Mill, A.Tj. coll (at light). 3 ♀, Ghibe River (260 km) (at light) 6/5-61 S. Chojnacki leg.; 4♀, Cojeb River [7.059N, 37.366E] 9-11/4-61 at light S. Chojnacki leg. (gift P. Brinck). 2♂ (Z75/2, NEOP 124; + Z16.182, NEOP 123), 1♂ „old slide“, 1♀ (Z16.161 & egg-SEM) 15km NW of Chora, Illubabor prov., 1600m, VII.1973, G. de Rougemont (SMNS). Gabonese Republic: 1♀, Gabon Moyen-Ogooué Rég., Najolé, Auberge St. Jean, Ogooué R., 0°11’03’’S, 10°46’00’’S 16.12.1993 C.C. Jongkind (slide GAB.01\2; ZMBN). Republic of Ghana: 1♂, Western region, Ankasa game prod. Reserve [5.44N, 2.07W, 47m] 10–16.12.93 St. 12 J. Kjaerendsen & T. Andersen light trap; NUFU-project (ZMBN, NEOP 125). Republic of Guinea: 2♂, petit affl. du Milo à Kousankoro [near Conakry, 9.53800, -13.67700] 21.10.1984 (gift J.M.Gibon, slide Z16/176). 1♂, no details (Fig. 17.3; slide 18.104). 1♂, Guinea, Riv. Niandan (bassin du Niger) à Bambaya 24.10.1984 J.M.Gibon (SMNS; slide). République de Côte d’Ivoire: 2♂, Elfenbeinküste, Boa, ca. 10 above entrance into Sassandra [township Sassandra: 5.563N, 6.579W], no date (SMNS, gift B. Statzner; Afr.86/17); 1♂: Ivory Coast, Cavally R. (blackwater) [township Cavally, Republic of Liberia, near border with Ivory Coast; 6.064N, 7.890W] nr Tai 02.07.1977 (SMNS, gift B. Statzner; Afr.86/19); 1♀, Côte d’Ivoire, Danané [7.25°N, 8.15°E] \ Muséum Paris 12.1930- IV.1931 CH. Alluaud & P.A.Chappuis (MNHN, egg slide in SMNS). LIBERIA: 4♂: Liberia, 8 mi NW Belefuanai [7.26N, 9.43W], S fork S.Paul R., 11.8.66, Ross & Lorenzen (CASENT 8413092; Z18.51). Republic of Sierra Leone: 1♂, Pampana près de Magburaka [8.67°N, 11.96°W], 5.2.89 (SMNS, gift J.M.Élouard; Z16.80, NEOP 126).

Habitus. WL 10–15mm, Rs with 3–4 terminal branches ( camerunensis lectotype: 12mm, three branches). Yellowish to ochre, a brownish heart-shaped spot between ocelli, sometimes traces of brownish pigment along occipital suture. Flagellum brown, cercus pale. Legs light, or a little infuscated around knees. Wings turbid.

Male. Tergites are similar to N. africana . T 7 in rear half with a triangular sclerite ending in a short narrow process, its raised tip with a few SB. Bottom of vertical rear face with a few SB. T8 with Y-shaped flat sclerite. A transverse line of SB at base of sclerite, some more on the narrow band-like caudal part. T9 unmodified, lateral humps pilose, median furrow distinct, SB mainly on humps. Mediobasal callus of HT10 slender, truncate, tip of HT short, not reaching beyond T9, fairly stout, gently curved outward. Epiproct and sternites unmodified.

Penis (Figs 143–149) tubular, sclerotised, apex curving ventrad, with shallow bare dorsal and ventral notches (Fig. 145). Endophallus twice as long as tube or longer and much thinner, forming a long curved hose (Fig. 143). Near its base the endophallus is often annulate and tends to curl. Two basolateral groups of small slender spinules unite dorsally and form a single row of erect spines which stand at considerable distances from one another (e.g., Fig. 146). Ventral face and sides bare.

Female (Figs. 131–132). S8 with three brown maculae that may merge and leave a pale central area (Fig. 131, top) or surround a variably distinct anchor pattern. Vagina slender, calyx-shaped, without spines or lateral sclerites. SSt slender, flexible, basal section with longitudinal folds on concave and a narrow spine band on convex side long, distally the entire SSt is scaly. Length of coil very variable, from 1.5 (top of Fig. 132) to up to 6 rings (Fig. 126). In relatively short coils there may be an angular projection of the distally widening spine band, in long coils the change is gradual.

Egg (Figs. 112, 131). Slender, 410*213µm, with few (9 in the camerunensis lectotype, Fig. 131) straight ridges, entire surface densely irregularly punctate, punctures small (Fig. 112). Anchor pole narrow, the elongate fused collar with two rings of cells with high walls. A step in the egg profile marks the end of ridges, the conical operculum is highly domed (Fig. 131). Micropyles are exposed, orifice on minute bare swelling (Fig. 112), the micropylar canal directed towards the operculum.

Variation. The SSt varies in length, is flexible, and takes variable positions in the female body. It is distinctly longer than the SSt of N. africana and N. panafricana . The lectotype of N. camerunensis has a long spermathecal coil (Fig. 131), similar specimens are known from several localities, from Ethiopia and Manyara N.P. in the east to near the Gaboon coast (Moyen Ogooué) and West Ghana. Several individuals from Cameroon and the ♀ from Danané (Ivory Coast) have exceptionally long coils (Fig. 126).

DNA (Figs. 491–492, 495). A total of six specimens from Sierra Leone, Ghana, Cameroon, and Ethiopia were sequenced, representing most of the geographic spread of this species and providing very strong support (99.4/100/100) for the monophyly of this species. Both sexes are represented.

While monophyletic based on mitochondrial DNA sequence data and morphology, the recognition of N. camerunensis (Enderlein) as a distinct species renders the concept of the above N. panafricana n. sp. paraphyletic (see above). They form a species complex that cannot be resolved with current molecular characters and should be investigated further with a much broader range of nuclear markers than just the mitochondrial COX1. The distinct N. camerunensis is not conspecific with the male holotype of N. panafricana n. sp. from the Republic of South Africa, but future revisions of the species complex will probably require the delineation of additional species.

Notes. Enderlein’s original material is mixed. By the lectotype designation (Zwick 1973a), former syntypes became paralectotypes (ICZN, Recommendation 74F). Only two of the surviving paralectotypes were correctly associated. Three more are misassociated: N. burgeoni Navás, N. dolium n. sp., and N. africana . The fourth is too damaged to be identified. The drawing of an egg from the N. camerunensis lectotype (Zwick 1973a, his fig. 71) is too stout (see Fig. 131).

The endophallus structure of N. camerunensis is apomorphic. It is much longer and narrower than in the related species, near the base only single spines stand dorsally, at wide distances from each other which is by transparency also visible in contracted penes (e.g., the N. haugi type, Figs. 144–145). Distal spines are smaller, stouter, and near the end there are several rows. Spermathecal coils and endophalli are exceptionally long in N. camerunensis; only N. sjostedti is similar in this respect. In contrast, the endophallus of N. panafricana is barely longer than the penis tube and the base bears dorsally a wide band of many spines (Figs. 138–142).

Only limited fresh material of N. camerunensis was available. DNA-analyses of many old samples failed. Therefore, the molecular analyses (Figs. 491–492, 495) include only few specimens, but the species is widespread and apparently not rare.