Marcusenius angolensis ( Boulenger, 1905 )

Kramer, Bernd & Wink, Michael, 2013, East-west differentiation in the Marcusenius macrolepidotus species complex in Southern Africa: the description of a new species for the lower Cunene River, Namibia (Teleostei: Mormyridae), Journal of Natural History (J. Nat. Hist.) 47 (35 - 36), pp. 2327-2362 : 2338

Marcusenius angolensis ( Boulenger, 1905)

( Figure 4B )

Gnathonemus angolensis Boulenger, 1905, p. 458 . Origin, Quanza River, Angola.

Gnathonemus macrolepidotus angolensis: Poll and Gosse 1963, p. 93 .

Marcusenius angolensis: Taverne 1971a, p. 103 ; Taverne 1971 b, p. 134; Taverne 1972, p. 166.

Marcusenius macrolepidotus angolensis: Gosse 1984, p. 86 . “Distribution: Angola ( Zaïre basin and Upper Zambezi), Zaïre, (Upper basin).” In consequence, Skelton et al. (1985) state that Marcusenius macrolepidotus angolensis applies to the Upper Zambezi form.

Marcusenius angolensis: Kramer et al. 2007, p. 680 –681.

Type specimens

Holotype (unique): BMNH 1905.5 .29.64, examined.

Non-types, examined:

– USNM 042332 Marcusenius macrolepidotus angolensis , 4 specimens, “pond near Cunga, Angola, Africa”, coll: Brown, W.H., 25 December 1889, assuming this location refers to the one Cunga that is associated with the Quanza, near Malanga in the Lower Quanza region , at 9 ◦ 25’29” S, 16 ◦ 21’19” E. ( There are at least four more Cungas in Angola that are associated with the Cunene (in the southwest), the Cuito or the Kwando (southeast), a tributary of the Lower Congo (northwest), and an independent coastal stream discharging into the Atlantic south of the Congo mouth, northwest ).

– USNM 042357 Marcusenius macrolepidotus angolensis , three specimens, “Quanza R., Angola ”, coll: Brown, W.H., “no tag in jar, no entry in date field of ledger, if locality is correct, date probably Dec. 1889 ” .

– ZSM 20948–949 , Gnathonemus angolensis Blgr 1905 , two specimens, “ Cuanza / Angola, SW-Afrika, III.1957, leg. Schoenfeldt, det. Terofal ” .

– BMNH 1907.6.29.231–233, Gnathonemus angolensis Blgr 1905 , three specimens, “Cunene, Mossamedes”, purchased Dr W. Ansorge. Mossamedes refers to the district of Mossamedes, then Angola’s southernmost district bordering Namibia (Stieler’s Hand-Atlas, 1910) .

Type locality

Angola: Quanza River (no. 2, Figure 1 ).

The former subspecies M. macrolepidotus angolensis ( Poll and Gosse, 1963) refers to the single specimen of G. angolensis Boulenger 1905 from the Quanza River in Angola (no more specific origin than the Lower Quanza is indicated by Boulenger 1910). The fishes (i.e., fish species) of the upper Quanza appear to be related to the fishes of the Zambezi basin and to the Cunene ( Trewavas 1973; P. Skelton 1994). Headwaters of both river systems arise just northeast of the city of Huambo on the Plateau of Bié, less than 20 km apart (the Cutato for the Quanza; Operational Navigation Chart, ONC N-3, 1: 1,000,000). Huambo is at an altitude of 1700 m, and both rivers flow in opposite directions. The type specimen, whose origin Boulenger (1905) gives only as “Quanza River, Angola ”, originates from the distant and tropical Lower Quanza, as concluded from the geographical detail given by J.W. Ansorge in Boulenger (1910). “The Lower Quanza ecoregion is considered its own distinct bioregion because the Zambezian fauna is absent or poorly represented, and a number of endemic species have been described” ( Thieme et al. 2005, p. 303). Seven specimens from USNM are also given as originating from the Quanza River, however, it is not clear from which section of this river, of which by far the longest section lies in the mountains, with many rapids and waterfalls. Two specimens from ZSM are listed as originating from the Quanza River (without further detail), however, this information could be incorrect (D. Neumann, personal communication). From the report by Hellmich (1957) and additional information at ZSM these specimens could originate from close to Ganda (13 ◦ 00 ′ 30 ′′ S, 14 ◦ 37 ′ 25 ′′ E), which lies on a small coastal system between Huambo and Benguela (the Jamba, a tributary of the Cubal). For comparison, the anatomical measures of these nine specimens are also given in Table A1 (in Appendix A) but cannot be used for taxonomic decisions unless their origin is known with greater precision.

Figures

Figure 1. Map of sampling locations in southern Africa. (A) Type locality for Marcusenius macrolepidotus (Peters, 1852), Tete on the Lower Zambezi River; (B) origin of M. macrolepidotus, SAIAB 60847, from Marromeu, Lower Zambezi delta (Figure 4A); (C) origin of M. macrolepidotus from the Buzi System, same location as SAIAB 67369, presently alive in aquarium; (D) likely type region for Marcusenius angolensis (Boulenger, 1905), the Lower Quanza; (E) Marcusenius altisambesi Kramer et al., 2007, type locality on the Upper Zambezi River; (F) M. altisambesi from the Kwando River; (G) M. altisambesi from the Okavango delta; (H, I) Marcusenius multisquamatus sp. nov., stretching from below the Ruacana Falls to above the Epupa Falls (I), the type locality; (J) specimens from the Cunene River Mouth.

Figure 4. Photographs of bulldog fish from various origins. (A) Marcusenius macrolepidotus (Peters, 1852), SAIAB 60847, coll. R. Bills 1 August 1999, Lower Zambezi; (B) Marcusenius angolensis (Boulenger, 1905), BMNH 1905.5.29.64 (holotype); (C) Marcusenius altisambesi, coll. F. H. van der Bank and B. Kramer, 21 August 1999, Upper Zambezi, Kalimbeza, live specimen of 16.5 cm SL photographed 3 July 2003 in aquarium; (D) M. altisambesi, coll. F.H. van der Bank and B. Kramer, 11/12 August 2004, Okavango River, live fish of SL 13 cm photographed 20 April 2006 in aquarium; (E) Marcusenius multisquamatus sp. nov., coll. B. Kramer and Ernst Swartz, coll. and photographed on 19 August 2006, Cunene River, below Ruacana Falls, specimen RUAC01, SL 15.4 cm.