Marcusenius altisambesi, Kramer & Skelton & Bank & Wink, 2007

Kramer, Bernd, Skelton, Paul, Bank, Herman Van Der & Wink, Michael, 2007, Allopatric differentiation in the Marcusenius macrolepidotus species complex in southern and eastern Africa: the resurrection of M. pongolensis and M. angolensis, and the description of two new species (Mormyridae, Teleostei), Journal of Natural History 41 (9 - 12), pp. 647-708 : 695-697

Marcusenius altisambesi , sp.n.

Holotype. SAIAB 79135 (specimen L39isi), SL 15.3 cm, total length 17.5 cm, Namibia: Caprivi Strip: Lisikili on side arm of Upper Zambezi River, 17 u 339S, 24 u 299E, 7 March 1994, male, coll.: F.H. van der Bank and B. Kramer, collected within 2 m depth from surface.

Paratypes. SAIAB 79136 (6) [L01isi, L06isi, L10isi, L19isi, L20isi, L26isi], ZSM 35085 (2) [L41isi, L40isi], ZSM 35086 (5) [L02isi, L04isi, L05isi, L12isi, L16isi], 5–7 March 1994, same origin and collectors as holotype, SAIAB 79137 (3) [B5, ID62, ID69], ZSM 35097 (1) [ID30], also from a site at Lisikili, 17 u 32931.90S, 24 u 26917.70E, coll.: F.H. van der Bank and B. Kramer, 5–6 August 2004, water conductivity and temperature, 75– 76 MS cm 21, 18.2–20.0 u C, ZSM 35082 (2) [ID86, ID87)], paratypes, two specimens as in preceding paragraph but from nearby locality Kalimbeza, 17 u 32927.30S, 24 u 31926.20E, 7 August 2004, similar water conductivity and temperature ,

- 104 specimens from the Upper Zambezi River System , East Caprivi, Namibia, size range 35–195 mm SL, smallest gravid female 113 mm SL (juveniles could not be sexed), some specimens from Kalimbeza presently alive in Aquarium :

- SMF 28264 (22 specimens, voucher references: LXXisi, XX ranging from 01 to 50) from the Zambezi River, Lisikili backwater, 17 u 339S, 24 u 299E (type locality), coll.: F.H. van der Bank and B. Kramer, 5–7 March 1994, water conductivity and temperature, 56.1 MS cm 21, 26.8 u C, 11 gravid females ,

- SMF 28264 (45 specimens, voucher references: NXXka or NXXXk (XX or XXX ranging from 54 to 126), ZSM 35084 (1) (specimen N123ka), from the Kwando River , Nakatwa, 18 u 069S, 23 u 239E, in Mudumu National Park, coll.: B. Kramer, 9–15 March 1994, water conductivity and temperature, 130 MS cm 21, 24.9 u C, eight gravid females, locality 12 on Figure 4 ,

- SMF 28264 (two specimens, voucher references: ‘‘7fish’’, 195 mm SL, female, and ‘‘11fish’’, 73 mm SL), from Kwando River, Nkasa Island (18 u 279S, 23 u 429E) in Mamili National Park, close to locality 12 on Figure 4 , coll.: F.H. van der Bank and B. Kramer, 9– 10 September 1993, water conductivity and temperature, 108 MS cm 21, 18–19 u C, not gravid,

- 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 MS cm 21 ,22 u C,size range 7.2– 13.3 cm SL, arrival live in Regensburg 2 September 1999, EOD recording 28 September–7 October 1999 at 100 MS cm 21 water conductivity and 21 u C (EOD recording in Germany for quicker transport in Africa ), Figure 1K , presently alive ,

- 1 specimen from Kwando River , Kongola Bridge, 17 u 47926.70S, 23 u 20940.00E, 24 January 2001, coll.: F.H. van der Bank & B. Kramer, ZSM 35083 (1) (specimen Kon08g) ,

- 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:

- 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 u 079300S 22 u 229E, coll.: F.H. van der Bank, J. Engelbrecht and B. Kramer, 20–22 January 2001, water conductivity and temperature, 37 MS cm 21, 29.9–30.6 u C, voucher references: OkaXXv (XX integer numbers ranging from 04 to 52), at least nine males as determined by the presence of a kink in anal fin, locality 13 on Figure 4 ,

- SAIAB 79143 (6), ZSM 35096 (5), and 24 specimens presently alive in aquarium, from the Okavango River at Guma Lagoon, 18 u 57946.60S, 22 u 22925.30E, coll.: F.H. van der Bank and B. Kramer, 10–12 August 2004, water conductivity and temperature 38 MS cm 21 and 21.4 u C, voucher references: Guma00XX (X for integer numbers ranging from 4 to 64), one male, close to locality 13 on map of Figure 4 , Figure 1J ,

- SAIAB 79141 (1), ZSM 35095 (1), and seven specimens presently alive in aquarium, details as in preceding paragraph, except for locality at Makwena, 19 u 03913.850S,22 u 22942.60E, 12 August 2004, voucher references Makw00XX (X ranging from 0 to 28).

Figures

Figure 1. All species and forms of the Marcusenius macrolepidotus species complex, as studied in the present paper. (A) M. macrolepidotus (Peters, 1852), ZMB 3678 (lectotype L. Seegers; photo: L. Seegers). (B) Gnathonemus angolensis Boulenger 1905, BMNH 1905.5.29.64 (holotype). (C) Gnathonemus moeruensis Boulenger 1915, MRAC 14137 (holotype). (D) Gnathonemus pongolensis Fowler 1934, ANSP 54950 (holotype). (E) M. macrolepidotus (Peters 1852), SAIAB 060847, coll. R. Bills 1 Aug.1999, Lower Zambezi. (F) M. macrolepidotus (Peters 1852), SAIAB 060947, coll. R. Bills 15 Aug.1999, lower Pungwe River System. (G) M. devosi sp. n., coll. L. de Vos and B. Kramer 3/6 Sept. 2001, Lower Tana River/Kenya. (H) M. macrolepidotus (Peters, 1852), SAIAB 73790 (largest specimen), coll. R. Bills 14 Aug. 2003, Rovuma System. (I) M. macrolepidotus (Peters 1852), SAIAB 055874, coll. R. Bills 20 July 1997, Mulela River/Mozambique. (J) M. altisambesi sp. n., 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. (K) M. altisambesi sp. nov, 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. (L) M. macrolepidotus (Peters 1852), sampled together with SAIAB 67369, coll. R. Bills 29 Sept. 2002, Buzi River System, specimen of 13 cm SL photographed alive 22 February 2005. (M) M. pongolensis (Fowler, 1934), resurrected species, specimen of 11 cm SL reared in captivity from parents caught in Crocodile River, Incomati System, in February 1997, photographed alive 15 March 2005.

Figure 4. Partial map of southern Africa showing the sampling localities for Marcusenius sp. Only the larger rivers are shown. Numbers code for origins of samples: (1) Tete on the Lower Zambezi (M. macrolepidotus, as for types); (2) Quanza River (G. angolensis, type; exact sampling site unknown); (3) Lake Mweru (G. moeruensis, type); (4) Pongola River near Paulpietersburg (G. pongolensis, type); (5) Sabie River; (6) Crocodile River; (7) Groot Letaba River (Limpopo System); (8) Blyde River (Limpopo System; (9) KwaMaZulu stream, Mhlatuze System (not shown); (10) Pongola River, more downstream compared to (4); (11) Lisikili, Upper Zambezi; (12) Kwando River, (13) Okavango System; (14) Tana River; (15) Buzi River System; (16) Pungwe River; (17) Lower Zambezi, delta region; (18) Mulela River; (19) Rovuma River System.