Marcusenius pongolensis (Fowler, 1934)

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 : 697-699

Marcusenius pongolensis (Fowler, 1934)

Material examined, all from South Africa.

- Gnathonemus pongolensis Fowler, 1934 . Holotype (unique): ANSP 54950. Origin , Pongola River, Paulpietersburg (locality no. 4, Figures 4 and 8 ; Figure 1D ),

- SAIAB 54445–54448: 30 specimens from the Sabie River System in or close to the Kruger National Park, totalling at least nine males, at least 14 females, size range 49 – 171 mm SL, largest male 171 mm SL, largest female 150 mm SL, no gravid material, coll.: P. Skelton and B. Kramer, 23–25 September 1993 ,

- SAIAB 54445, one specimen from the Sand River (tributary of the Sabie) at Londolozi, site 14, 24 u 479310S, 31 u 319320E, standing pool, muddy water of 250 MS cm 21 and 26 u C temperature, male, SL 87 mm,

- SAIAB 54446, 11 specimens from the Sabie River above Hazy View, at least seven of which are male, two female, size range 49–171 mm SL 25 u 029S, 31 u 009E, water conductivity and temperature, 120 MS cm 21, 21.1 u C ,

- SAIAB 54447, six specimens from the Sabie River above Skukuza just below the weir, all female, size range 74–150 mm SL, site 8, 24 u 589350S, 31 u 359050E, water conductivity and temperature, 160 MS cm 21, 26 u C,

- SAIAB 54448, 12 specimens at least one of which male, six female, size range 49–118 mm SL, Sabie River immediately above Hazy View (municipal picnic site), site 5, 25 u 019480S, 31 u 019210E, water temperature and conductivity, 125 MS cm 21, 24.4 u C,

- 14 specimens from the Sabie River, bridge near Lower Sabie tourist camp (25 u 079S, 31 u 559E), water conductivity and temperature, 139 MS cm, 25.1 u C, coll.: F.H. van der Bank and B. Kramer, 29–30 March 1996, size range 103–173 mm SL, largest male 173 mm SL, largest female 133 mm SL, no gravid material, imported live,

21

- SAIAB 79144 (1), one specimen from Sabie River, Sekurekwane (24 u 599S, 31 u 119E), water conductivity and temperature, 106 MS cm 21, 23.3 u C, coll.: F.H. van der Bank and B. Kramer, 27 March 1996, 82 mm SL,

- SAIAB 79145 (1), one specimen from the anastomosing reaches of the Sabie/Sand confluence at 24 u 57922.70S, 31 u 42938.70E, water conductivity and temperature, 129 MS cm 21 and 24.3 u C, coll.: F.H. van der Bank and B. Kramer, 28 March 1996, 8.6 cm SL, all of the above near locality 5 on Figures 4 and 8 ,

- 29 specimens (23 for EOD) from the Crocodile River (Incomati System), Stentor Estates (town: Kaapmuiden , Province Mpumalanga), 25 u 309350S, 31 u 119580E, altitude 432 m, coll.: F.H. van der Bank and J. Engelbrecht, 14 February 1997, water conductivity and temperature, 230 MS cm 21 at 25 u C , 23 males, six females, size range 148–275 mm SL, largest male 275 mm SL, largest female 200 mm SL (in the Crocodile River region it had rained a lot, the river was flowing very strong and the water was not clear), locality 6 on Figures 4 and 8 , Figure 1M , imported live ,

- ZSM 35089 (1), one specimen from the Groot Letaba River just below Tzaneen Dam (Northern Province), 23 u 499000S, 30 u 109000E, coll.: W. Vlok and B. Kramer, 22 September 1998, water conductivity and temperature, 114 MS cm, 21.4 u C, 69 mm SL, locality 7 on Figures 4 and 8 , field no. Lim03 ,

21

- SAIAB 79147 (6), ZSM 35088 (4), 10 specimens from Blyde River just below Blyderivierspoort Dam (Mpumalanga), 24 u 329110S, 30 u 479520E, coll.: J. Engelbrecht, W. Vlok and B. Kramer, 25/ 26 September 1998, water conductivity and temperature, 154 MS cm 21, 16.7 u C, size range 40–140mm SL (both the Blyde and Letaba rivers are tributaries of the Olifants River that forms part of the Limpopo System ), locality 8 on Figures 4 and 8 ,

- SAIAB 79149 (4), ZSM 35090 (3), seven specimens from the KwaMaZulu stream close to where it flows into Goedertrouw Dam, part of the Mhlatuze System in KwaZulu-Natal (approx. 28 u 259 300S, 31 u 19 300E), water conductivity and temperature, 134 MS cm 21 and 17.1 u C, coll.: J. Engelbrecht and B. Kramer, 12 August 1999, locality 9 on Figures 4 and 8 ,

- SAIAB 79148 (5), ZSM 35087 (5), 10 specimens from the Pongola River in KwaZulu- Natal (27 u 019 150S, 32 u 189E), bridge at road connecting Ndumo with Kosibay, coll.: J. Engelbrecht and B. Kramer 14 August 1999, water conductivity and temperature, 600 MS cm, 22 u C, locality 10 on Figures 4 and 8 ,

21

- SAIAB 77296 (2), two specimens from the Nswananzi River: Nkanini stream in KwaZulu-Natal, Kosibay, below bridge just outside Manguzi town on road to Mozambique, 26 u 569240S 032 u 449390E, coll: O. Gon, H. Larson, V. Mthombeni and S. Kyle, 13 November 2005, fresh and clear water, sandy to silt bottom with rocks on the banks (no. 20 on Figure 8 ).

Further downstream the Mhlatuze, mormyrids (only possibility: bulldogs) were also demonstrated (but not caught) using an electronic fishfinder (making EODs audible through headphones) at Lake Mzingazi near Richards Bay, in water of 496 MS cm 21 and 21.8 u C on August 12, 1999. The Mhlatuze System is the southernmost record for bulldogs and mormyrids in general (J. Engelbrecht, pers. comm.).

Additional material studied for gonad histology. Of the above specimens from the Crocodile River, five males (178, 183, 245, 250 and 252 mm SL), two females (148 and 153 mm SL), plus six additional fish from the Crocodile River (same collection dates), size range 93– 141 mm SL, one male of 137 mm SL, five females, largest female 141 mm SL.

Marcusenius devosi , sp. n.

Thirty specimens

Holotype. SAIAB 79138 (specimen Ta13na), SL 10.2 cm, total length 11.6 cm, Kenya: Tana River: Tana Primate Research Centre, 1 u 52938.10S, 40 u 08922.50E, south of village Wenje , east of road B8, 48 m above sea level, locality 14 on Figure 4 , 3 September 2001, female, coll. by L. De Vos and B. Kramer at ǩ 1.5 m depth, water conductivity and temperature, 185 MS cm 21 and 26 u C.

Paratypes. SAIAB 79139 (14), ZSM 35091 (3), ZSM 35092 (1), ZSM 35093 (4), ZSM 35094 (7) (specimens Ta10na, TA11na, Ta14na, Ta18na, Ta20na, Ta21na, Ta22na,

Ta25na, Ta27na, Ta28na, Ta29na, Ta30na, Ta31na, Ta32na, Ta33na, Ta34na, Ta47na, SinEOD1, SinEOD4, SinEOD7, ID1–ID9), same origin as holotype, 3–6 September 2001, size range 79–121cm SL.

Material examined for DNA

Four specimens of M. macrolepidotus from the Lower Zambezi River, Mozambique (macrolepidotus 22688, 12 August 1999, 18 u 179S 35 u 579E, near village Marromeu; macrolepidotus 22689, 22690, 13 August 1999, 18 u 199S35 u 559E, SE from Marromeu; macrolepidotus 22686, Marromeu fish market, 11 August 1999, 18 u 179S 35 u 579E; numbers are institutional accession numbers, University of Heidelberg, M. Wink) [GenBank accession numbers, M.macrolepidotus .MOZ DQ863663 , M.macrolepidotus .MOZ DQ863664 ], five specimens of M. macrolepidotus from the middle Buzi River System (macrolepidotus 22681, 22682, 22683, 22684, 22685; see Material examined); [GenBank accession numbers, M.macrolepidotus .MOZ DQ863662 , M.macrolepidotus .MOZ DQ863665 , M.macrolepidotus .MOZ DQ863669 , M.macrolepidotus .MOZ DQ863671 ], two specimens of M. macrolepidotus from the Rovuma River (origin and date as given in Material examined) [GenBank, M.macrolepidotus .Rov DQ863673 , M.macrolepidotus .Rov DQ863674 ]. Five specimens of M. pongolensis from the Crocodile River, Stentor Estates, Kaapmuiden, Mpumalanga, South Africa (25 u 309350S, 31 u 119580E), 14 February 1997, ecological data as given above, two female (112 and 141 mm SL), two male (134 and 137 mm SL), one juvenile (109 mm SL) [GenBank, M.pongolensisSA14.C DQ863650 , M.pongolensisSA15.C DQ863651 , M.pongolensisSA9.Cr DQ863657 , M.pongolensisSA12.C DQ863658 , M.pongolensisSA13.C DQ863659 ]; two specimens from the Pongola River, South Africa, 14 August 1999, SL 15, 16.9 cm [ M. pongolensis .9127, M. pongolensis .9129]; three specimens from the Mhlatuze System, KwaMazulu stream, South Africa, 12 August 1999, SL 7.6, 8.1 and 12.9 cm, the latter of which a male [GenBank, M.pongolensis .Mhlat DQ863647 , M.pongolensis .Mhlat DQ863648 , M.pongolensis .Mhlat DQ863649 ]. Five specimens of M. altisambesi from the Upper Zambezi System, East Caprivi, Namibia: NAM1 (18 u 049530S, 24 u 029350E) from the Linyanti River, 22 February 1997, NAM3 and NAM5 (18 u 099240S, 23 u 239090E) from the Kwando River, NAM2 (23 February 1997; 125 mm SL, 24 g, SAIAB 74735) and NAM148 (7 April 1996, 81 mm SL, juvenile) from the Upper Zambezi River (17 u 329240S, 24 u 319250E), altisambesi.145 from Upper Zambezi System (Upper Zambezi River at same location, or lower Kwando River) [GenBank, M.altisambesi .NAM2.U DQ863652 , M.altisambesi .NAM5.K DQ863653 , M.altisambesi .NAM1.L DQ863654 , M.altisambesi .NAM148 DQ863655 , M.altisambesi .NAM3.K DQ863656 ]. Two specimens of M. altisambesi, BOT 22670 and BOT 22671, from the Okavango, Thoage River, Makwena, Botswana (19 u 079300S, 22 u 229E), coll. F.H. van der Bank, 21 January 2001 [GenBank M.altisambesi .BOT226 DQ863660 , M.altisambesi .BOT226 DQ863661 ]. Included as a confamilial outgroup are two Pollimyrus marianne Kramer et al. 2003 from the Upper Zambezi System, East Caprivi, Namibia, Linyanti River, 18 u 049530S, 24 u 029350E, 22 February 1997 [GenBank: AY234099 ], and one specimen each of Hippopotamyrus ansorgii (Kwando River, Kongola bridge, 24 January 2001) [GenBank: AY236995 ] and H. szaboi (Upper Zambezi River, Wenela at Katima Mulilo, 27 August 1999) [GenBank: AY236983 ]. As a more distant outgroup ( Siluriformes ) served one Schilbe intermedius Rüppell, 1832 (SAIAB 056106) sampled from the Upper Zambezi River, Katima Mulilo, Namibia (17 u 329S, 24 u 319E), 8 May 1997 [GenBank: DQ863646 ].

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 3. Oscillograms of the waveforms of electric organ discharges (EODs) in the M. macrolepidotus species complex. (A) specimens from the Buzi River (Mozambique), (B) from the Okavango River (Botswana), (C) from the Upper Zambezi and Kwando Rivers (Namibia) as indicated, (D) from the Incomati System (South Africa), (E) from the Tana River (Kenya). J, juveniles; F, females; M, males, all with standard length in mm. (C) Superimposed baseline shown in male discharge of long duration. Head-positivity is upwards. Abscissa, see time bar (identical for all). All waveforms normalized to 25uC. (A, lower right) Head-negative pre-potential present in specimens from the Upper Zambezi/Okavango Systems, not from the Buzi River nor any other origin (amplified 63).

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.

Figure 8. Map of sampling locations in South Africa. Numbers explained in Figure 3, except no. (20) which represents two samples from the Nswananzi River: Nkanini stream, KwaZulu-Natal.