Marcusenius altisambesi Kramer et al., 2007

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 : 2335-2337

Marcusenius altisambesi Kramer et al., 2007

( Figure 4C, D )

Gnathonemus okavangensis Pappenheim, 1907 . “Appeared as a form of Gnathonemus macrolepidotus Peters from the Okavango R., Damaraland, Africa; regarded as infrasubspecific and not available” (Eschmeyer 2013). “Nomen dubium” according to Gosse (1984) and Seegers (1996, p. 73).

Gnathonemus macrolepidotus: Gilchrist and Thompson 1913, pp. 330–331 .

Marcusenius altisambesi Kramer et al. (2007) , pp. 681–684.

Type specimens

Holotype: SAIAB 79135 (specimen L39isi), Namibia: Caprivi Strip: Lisikili on Upper Zambezi River . Paratypes: SAIAB 79136 (6) , SAIAB 79137 (3) , ZSM 35086 (5) , ZSM 35085 (2) , ZSM 35097 (1) , ZSM 35082 (2) ; all examined.

– Non-types, examined. One hundred and four specimens from the Upper Zambezi River System , East Caprivi, Namibia, some specimens from Kalimbeza presently alive in Aquarium :

– SMF 28264 (22 specimens), from the Zambezi River, Lisikili backwater, 17 ◦ 33 ′ S, 24 ◦ 29 ′ E (type locality), coll.: F.H. van der Bank and B. Kramer, 5–7 March 1994 ,

– SMF 28264 (45 specimens) , ZSM 35084 (1) , from the Kwando River , Nakatwa, 18 ◦ 06 ′ S, 23 ◦ 23 ′ E, in Mudumu National Park, coll.: B. Kramer, 9–15 March 1994, locality 6 on Figure 1 ,

– SMF 28264 (two specimens), from Kwando River , Nkasa Island (18 ◦ 27 ′ S, 23 ◦ 42 ′ E) in Mamili National Park, close to locality 6 on Figure 1 , coll.: F.H. van der Bank and B. Kramer, 9–10 September 1993 ,

– 31 specimens, about 500 m from opposite Kalimbeza fishing camp, at downstream tip of small island between Lisikili side channel and main channel, coll.: F.H. van der Bank and B. Kramer, caught 21 August 1999, water conductivity and temperature, 84 µS cm, 22 ◦ C, size range 7.2–13.3 cm SL, arrival live in Regensburg 2 September 1999, EOD recording 28 September to 7 October 1999 at 100 µS cm−1 water conductivity and 21 ◦ C ( EOD recording in Germany for quicker transport in Africa), presently alive,

−1

– ZSM 35083 (1), from Kwando River , Kongola Bridge, 17 ◦ 47 ′ 26.7 ′′ S, 23 ◦ 20 ′ 40.0 ′′ E, 24 January 2001, coll.: F.H. van der Bank and B. Kramer ,

– Non-types (63 specimens)from the Okavango River , Botswana, totalling at least 10 males, male size range 110–181 mm SL, juvenile/female size range 54–169 mm SL, examined :

– SAIAB 79140 (9) , ZSM 35079 (1) , ZSM 35080 (3) , ZSM 35081 (6) from the Okavango River, Makwena Lodge, near the township of Etsha no. 6, 19 ◦ 07 ′ 30 ′′ S, 22 ◦ 22 ′ E, coll.: F.H. van der Bank, J. Engelbrecht and B. Kramer, 20–22 January 2001, locality 7 on Figure 1 ,

– SAIAB 79143 (6), ZSM 35096 (5), and 24 specimens presently alive in aquarium, from the Okavango River at Guma Lagoon , 18 ◦ 57 ′ 46.6 ′′ S, 22 ◦ 22 ′ 25.3 ′′ E, coll.: F.H. van der Bank and B. Kramer, 10–12 August 2004, close to locality 7 on Figure 1 ,

– SAIAB 79141 (1), ZSM 35095 (1), and seven specimens presently alive in aquarium, details as in preceding paragraph, except for locality at Makwena, 19 ◦ 03 ′ 13.85 ′′ S, 22 ◦ 22 ′ 42.6 ′′ E, 12 August 2004.

Samples examined for genetics. DNA samples are stored at Institute of Pharmacy and Molecular Biotechnology, Heidelberg University ( IPMB).

– IPMB 44903–44905 , Namibia: Upper Zambezi: Kalimbeza , 17 ◦ 32 ′ 27.3 ′′ S, 24 ◦ 31 ′ 26.2 ′′ E, coll. F.H. van der Bank and B. Kramer, 21 August 1999 ;

– IPMB 44638–44640 Botswana: Okavango : Guma Lagoon, 18 ◦ 57 ′ 46.6 ′′ S, 22 ◦ 22 ′ 25.3 ′′ E, coll. F.H. van der Bank and B. Kramer, 10 August 2004 ; IPMB 44641 , 44642 , as before, but 10–12 August 2004; GenBank accession numbers: ( KC 202230 – KC 202237 ) .

Type locality

Upper Zambezi River in East Caprivi ( Namibia); specifically Upper Zambezi River between Lisikili and Kalimbeza (or Kalambesa, 17 ◦ 33 ′ S, 24 ◦ 29 ′ E to 17 ◦ 32 ′ 27.3 ′′ S, 24 ◦ 31 ′ 26.2 ′′ E; 22–26 km straight line downstream from Katima Mulilo; Figure 1 , no. 5) .

The first record of G. macrolepidotus for the Upper Zambezi is that of Gilchrist and Thompson (1917, p. 562), specifying Lialui, Barotseland as origin. For a description, the authors refer to Gilchrist and Thompson (1913, p. 330), a description of South African specimens that Kramer et al. (2007) have referred to M. pongolensis (Fowler, 1934) . The presence of G. macrolepidotus in the Upper Zambezi System was confirmed by Jubb (1958). Another possible synonym would be G. okavangensis if it were available (this name should be dropped from a list of synonyms, as suggested by Kramer et al. 2007). Upper Zambezi and Okavango specimens were recognized as representing a new species, M. altisambesi , that is well differentiated from M. macrolepidotus ( Peters, 1852) by Kramer et al. (2007).

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.