Abstract
Recent morphological and genetic studies have revealed two new species of snoutfish in South Africa, Marcusenius caudisquamatus and M. krameri, which had been confused with M. pongolensis, the South African bulldog fish. All known mormyriform fish are nocturnal and emit electric organ discharges (EODs) for communication that are characteristic for their species. This paper examines whether or not the EODs of these three closely-related South African species can be differentiated from each other. An EOD pulse of a bulldog fish consists of a head-positive phase P, followed by a head-negative phase N of short duration. We measured and compared six variables of the EOD pulse waveform for South African samples for the three species from different locations using MANOVA, ANOVA and Discriminant Analysis, with M. devosi from Kenya as an outgroup. The EOD waveforms, normalized to the same P-phase amplitude, varied significantly from each other in four variables, most strongly in the amplitude of the N phase and the duration of the P phase. In two species, M. devosi and M. krameri, there was no evidence of difference between sexes, in contrast to M. pongolensis and M. caudisquamatus whose male pulses were of longer duration. M. devosi and M. krameri were statistically significantly independent of each other and of any other group studied. By contrast, the M. pongolensis specimens from different locations showed a high degree of variability amongst each other, including significant separation, and overlap with M. caudisquamatus.
References
Bullock, T.H., Hopkins, C.D., Popper, A.N. & Fay, R.R. (Eds.), (2005) Electroreception. Springer, New York, 467 pp.
Crass, R.S. (1960) Notes on the freshwater fishes of Natal with descriptions of four new species. Annals of the Natal Museum, 14, 405–458.
Fowler, H.W. (1934) Fishes obtained by Mr. H. W. Bell-Marley chiefly in Natal and Zululand in 1929 to 1932. Proceedings of the Academy of Natural Sciences of Philadelphia, 86, 405–514.
Graff, C. & Kramer, B. (1992) Trained weakly-electric fishes Pollimyrus isidori and Gnathonemus petersii (Mormyridae, Teleostei) discriminate between waveforms of electric pulse discharges. Ethology, 90, 279–292.
http://dx.doi.org/10.1111/j.1439-0310.1992.tb00839.xHanika, S. & Kramer, B. (1999) Electric organ discharges of mormyrid fish as a possible cue for predatory catfish. Naturwissenschaften, 86, 286–288.
http://dx.doi.org/10.1007/s001140050616Hanika, S. & Kramer, B. (2000) Electrosensory prey detection in the African sharptooth catfish, Clarias gariepinus (Clariidae), of a weakly electric mormyrid fish, the bulldog (Marcusenius macrolepidotus). Behavioral Ecology and Sociobiology, 48, 218–228.
http://dx.doi.org/10.1007/s002650000232Hanika, S. & Kramer, B. (2005) Intra-male variability of its communication signal in the weakly electric fish, Marcusenius macrolepidotus (South African form), and possible functions. Behaviour, 142, 145–166.
http://dx.doi.org/10.1163/1568539053627677Hanika, S. & Kramer, B. (2008) Plasticity of electric organ discharge waveform in the South African bulldog fish, Marcusenius pongolensis: tradeoff between male attractiveness and predator avoidance? Frontiers in Zoology, 5, 7–7.
http://dx.doi.org/10.1186/1742-9994-5-7Kramer, B. (1997) A field study of African elephantfish (Mormyridae, Teleostei): electric organ discharges in Marcusenius macrolepidotus (Peters, 1852) and Petrocephalus catostoma (Günther, 1866) as related to sex. Journal of African Zoology, 111, 313–341.
Kramer, B. (2009a) Electric communication and electrolocation. In: Binder, M.D., Hirokawa, N. & Windhorst, U. (Eds.), Encyclopedia of Neuroscience. Springer-Verlag, Berlin Heidelberg, pp. 1038–1045.
http://dx.doi.org/10.1007/978-3-540-29678-2_2913Kramer, B. (2009b) Electric fish. In: Binder, M.D., Hirokawa, N. & Windhorst, U. (Eds.), Encyclopedia of Neuroscience. Springer Verlag, Berlin Heidelberg, pp. 1045–1050.
http://dx.doi.org/10.1007/978-3-540-29678-2_2915Kramer, B. (2009c) Electric organ discharge. In: Binder, M.D., Hirokawa, N. & Windhorst, U. (Eds.), Encyclopedia of Neuroscience. Springer Verlag, Berlin Heidelberg, pp. 1050–1056.
http://dx.doi.org/10.1007/978-3-540-29678-2_2917Kramer, B. (2013) A morphological study on species of African Mormyrus (Teleostei: Mormyridae) and their electric organ discharges. African Journal of Aquatic Science, 38, 1–19.
http://dx.doi.org/10.2989/16085914.2012.745806Kramer, B., Bills, R., Skelton, P. & Wink, M. (2012) A critical revision of the churchill snoutfish, genus Petrocephalus Marcusen, 1854 (Actinopterygii: Teleostei: Mormyridae), from southern and eastern Africa, with the recognition of Petrocephalus tanensis, and the description of five new species. Journal of Natural History, 46, 2179–2258.
http://dx.doi.org/10.1080/00222933.2012.708452Kramer, B., Skelton, P., Van der Bank, H. & Wink, M. (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, 647–708.
http://dx.doi.org/10.1080/00222930701250987Kramer, B. & Van der Bank, F.H. (2000) The southern churchill, Petrocephalus wesselsi, a new species of mormyrid from South Africa defined by electric organ discharges, genetics, and morphology. Environmental Biology of Fishes, 59, 393–413.
http://dx.doi.org/10.1023/A:1026562827022Kramer, L.B., Van der Bank, F.H. & Wink, M. (2016). Marcusenius desertus sp. nov. (Teleostei: Mormyridae), a mormyrid fish from the Namib Desert. African Journal of Aquatic Science, 41, 1–18.
http://dx.doi.org/10.2989/16085914.2015.1137855Kramer, B. & Westby, G.W.M. (1985). No sex difference in the waveform of the pulse type electric fish, Gnathonemus petersii (Mormyridae). Experientia, 41, 1530–1531.
http://dx.doi.org/10.1007/BF01964790Lamml, M. & Kramer, B. (2007) Allopatric differentiation in the acoustic communication of a weakly electric fish from southern Africa, Marcusenius macrolepidotus (Mormyridae, Teleostei). Behavioral Ecology and Sociobiology, 61, 385–399.
http://dx.doi.org/10.1007/s00265-006-0267-0Lavoué, S., Sullivan, J.P. & Arnegard, M. (2010) African weakly electric fishes of the genus Petrocephalus (Osteoglossomorpha: Mormyridae) of Ozdala National Park, Republic of the Congo (Lékoli River, Congo River basin) with description of five new species. Zootaxa, 2600, 1–52.
Lowe-McConnell, R.H. (1987). Ecological studies in tropical fish communities. Cambridge University Press, Cambridge, 382 pp.
http://dx.doi.org/10.1017/cbo9780511721892Maake, P.A., Gon, O. & Swartz, E.R. (2014) Description of three new species of Marcusenius Gill 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa, 3780 (3), 455–480.
http://dx.doi.org/10.11646/zootaxa.3780.3.2Machnik, P. & Kramer, B. (2008a) A male's playback signal turns female Marcusenius pongolensis receivers on or off depending on his behavioural state. Communicative & Integrative Biology, 1, 128–131.
http://dx.doi.org/10.4161/cib.1.2.6860Machnik, P. & Kramer, B. (2008b) Female choice by electric pulse duration: attractiveness of the males' communication signal assessed by female bulldog fish, Marcusenius pongolensis (Mormyridae, Teleostei). The Journal of Experimental Biology, 211, 1969–1977.
http://dx.doi.org/10.1242/jeb.016949Machnik, P. & Kramer, B. (2011) Novel electrosensory advertising during diurnal resting period in male snoutfish, Marcusenius altisambesi (Mormyridae, Teleostei). Journal of Ethology, 29, 131–142.
http://dx.doi.org/10.1007/s10164-010-0235-zMachnik, P., Markowski, B. & Kramer, B. (2010) Intra- versus intersexual selection in the dimorphic electric organ discharges of the snoutfish Marcusenius altisambesi (Mormyriformes, Teleostei). Behaviour, 147, 677–704.
http://dx.doi.org/10.1163/000579510X489895Moller, P. (1995) Electric fishes: History and behavior. Chapman & Hall, London.
Paintner, S. & Kramer, B. (2003) Electrosensory basis for individual recognition in a weakly electric, mormyrid fish, Pollimyrus adspersus (Günther, 1866). Behavioral Ecology and Sociobiology, 55, 197–208.
http://dx.doi.org/10.1007/s00265-003-0690-4Peters, R.C. (2009) Electroreceptor organs. In: Binder, M.D., Hirokawa, N. & Windhorst. U. (Eds.), Encyclopedia of Neuroscience. Berlin Heidelberg: Springer Verlag, Berlin Heidelberg, pp. 1078–1083.
http://dx.doi.org/10.1007/978-3-540-29678-2_2952Peters, W.C.H. (1852) Einige neue Säugethiere und Flussfische aus Mossambique. Monatsber. Königl. Preuss. Akademie der Wissenschaften zu Berlin, 1852, 273–276.
Werneyer, M. & Kramer, B. (2002) Intraspecific agonistic interactions in freely swimming mormyrid fish, Marcusenius macrolepidotus (South African form). Journal of Ethology, 20, 107–121.
http://dx.doi.org/10.1007/s10164-002-0062-yWerneyer, M. & Kramer, B. (2005) Electric signaling and reproductive behaviour in a mormyrid fish, the bulldog Marcusenius macrolepidotus (South African form). Journal of Ethology, 23, 113–125.
http://dx.doi.org/10.1007/s10164-004-0136-0Werneyer, M. & Kramer, B. (2006) Ontogenetic development of electric organ discharges in a mormyrid fish, the bulldog Marcusenius macrolepidotus (South African form). Journal of Fish Biology, 69, 1–12.
http://dx.doi.org/10.1111/j.1095-8649.2006.01195.xWiltschko, W. & Kramer, B. (2009) Magnetic and electric senses. In: Binder, M.D., Hirokawa, N. & Windhorst, U. (Eds.), Encyclopedia of Neuroscience. Springer Verlag, Berlin Heidelberg, pp. 2198–2206.
http://dx.doi.org/10.1007/978-3-540-29678-2_3268