taxonID	type	description	language	source
3F5BF765FF86FFF3FF31FA67FA64C1B6.taxon	materials_examined	Holotype. SAIAB 88684, 166 mm SL, male, Mhlatuze River, under the bridge on road R 102, 28 ° 50 ' 44.2 " S, 31 ° 51 ' 59.6 " E, South Africa, P. Maake, B. Kramer and B. Mackenzie, 8 November 2009. Hologenetype COI (GenBank number KJ 174311), Hologenetype cyt b (GenBank number KJ 174301). Paratypes. SAIAB 191225, 6: 87.6 – 175 mm SL, Paragenetypes COI (GenBank KJ 174309 and KJ 174310), Paragenetypes cyt b (GenBank KJ 174299 and KJ 174300), same collection data as holotype, and MRAC, B 3 - 16 - P- 3, 113.07 mm SL, Paragenetype COI (GenBank KJ 174312), paragenetype cyt b (GenBank KJ 174302), same collection data as holotype. SAIAB 79149, 4: 60.3 – 114.4 mm SL, KwaMaZulu stream close to where it flows into Goedertrouw Dam, part of the Mhlatuze River system in KwaZulu-Natal, 28 ° 25 ' 30 '' S, 31 ° 1 ' 30 '' E, J. Engelbrecht and B. Kramer, 12 August 1999. SAIAB 96607, 2: 200 – 218 mm SL, Nseleni River - Nsezi Lake, 28 ° 44.766 ' S, 31 ° 58.728 ' E, O. Weyl, R. Karsing and B. Ellender, 27 May 2010. BMNH 2013 9.4. 74, 215 mm SL, same collection data as previous lot. SAIAB 88692, 2: 242 – 255 mm SL, Nseleni River, 28 ° 42 ' 57.8 " S, 31 ° 59 ' 34.4 " E, P. Maake and O. Weyl, 10 November 2009. SAIAB 96543, 2: 192 – 193 mm SL, Nseleni River - Nature Reserve - Richards Bay, 28 ° 41.965 ' S, 32 ° 00.077 ' E, O. Weyl, R. Karsing, R. Jones and B. Ellender, 23 May 2010. USNM 410775, 184 mm SL, same collection data as previous lot. SAIAB 96603, 196 mm SL, Lower Nseleni River opposite Davidson's Farm, 28 ° 41.965 ' S, 32 ° 00.077 ' E, O. Weyl, R. Karsing, R. Jones and B. Ellender, 27 May 2010.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF86FFF3FF31FA67FA64C1B6.taxon	diagnosis	Diagnosis. Middle body depth 23.8 – 25.9 % SL; dorsal fin length 15.4 – 19.2 % SL; anal fin length 23.0 – 24.9 % SL; pre-dorsal length 61.7 – 66.7 % SL; pre-anal length 60.7 – 62.8 % SL; caudal peduncle depth 40.4 – 51.7 % of its length (greater than and non-overlapping with M. krameri and M. lucombesi); dorsal fin rays 20 – 22; anal fin rays 27 – 28; scales around caudal peduncle 19; lateral line scales 72 – 73; anterior GR (4) + (5 – 6) = (9 – 10); posterior GR (6) + (9) = 15; conical teeth on upper / lower jaws 5 – 5.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF86FFF3FF31FA67FA64C1B6.taxon	description	Description. Measurements and meristic counts are given in Table 5. Head with terminal mouth well in front of eye; mental lobe on lower jaw protruding in front of upper jaw; snout rounded and blunt; pre-anal distance shorter than pre-dorsal distance; distance from origin of anal fin to origin of dorsal fin greater than middle body depth; pre-pelvic distance twice as long as distance between pelvic and anal fin; dorsal and anal fins set well back on the body, dorsal fin situated about three fourths of standard length from snout and opposite anal fin; dorsal fin shorter, originating on vertical at 4 th or 5 th anal fin ray and ending before anal fin base; distal margin of dorsal and anal fins obliquely orientated, with rays becoming gradually shorter posteriorly; dorsal fin rays 20 (20 – 22), its anterior rays highest, with distal margin rounded and slightly concave; anal fin rays 27 (27 – 28); males at sexual maturity have kink in the base of the anal fin distinctly curving inward; anterior anal fin rays of sexually mature males strong and sometimes darker, 3 rd – 5 th rays longer than the first two rays, crescentic and rounded or curved backwards, but are anteriorly sharp or pointy in females and juveniles; pectoral fin rays 10; pectoral fin length 73.4 – 96.9 % of head length; middle body depth 25.1 % (23.7 – 25.9 %) of SL; caudal peduncle thicker and stronger, subcylindrical across its entire length, 19.7 % (18.6 – 21.6 %) in SL; scales along the caudal peduncle circumference 19 (N = 15); caudal peduncle depth 45.2 % (40.4 – 51.7 %) into its length; lateral line scales 70 (72 – 73); lateral line scales cycloid with reticulate striae; jaws with 5 – 5 conical teeth; 44 vertebrae (excluding urostyle); anterior GRt = 9 (9 – 10); on the first gill arch (Table 6), gill rakers of the anterior row on first arch conicals, thin and more developed (longer and smooth); posterior GRt = 15 on the first gill arch; rakers on the posterior row shorter, thicker and with flattened tips. Character Holotype N Mean / Median Min Max Std. Dev SL, mm 166 mm 15 157.19 84.64 250 49.23 HL, mm 35.34 mm 15 32.77 19.13 51.3 8.89 Ratio of SL HL 0.2129 15 0.2114 0.1868 0.2273 0.0117 vBD 0.2404 15 0.2399 0.2305 0.2481 0.0050 mBD 0.2512 15 0.2495 0.2376 0.2588 0.0065 dAD 0.2519 15 0.2556 0.2443 0.2727 0.0081 CPL 0.1966 15 0.2017 0.1864 0.2168 0.0083 PDL 0.6527 15 0.6414 0.6171 0.6668 0.0138 PAL 0.6165 15 0.6186 0.6078 0.6285 0.0052 pD 0.4047 15 0.4056 0.4001 0.4092 0.0028 * MP pA 0.4313 15 0.4339 0.4301 0.4379 0.0026 * MP pPL 0.2301 15 0.2365 0.2301 0.2464 0.0056 pVL 0.4119 15 0.4089 0.3912 0.4234 0.0090 dVA 0.2134 15 0.2127 0.1927 0.2336 0.0110 NB: MP = M. pongolensis, ML = M. lucombesi, MK = M. krameri Electric organ discharge. Kramer et al. (2007) concluded that the EODs of seven individuals from Mhlatuze River (SAIAB 79149, 4; ZSM 35090, 3) cannot be differentiated from those of M. pongolensis collected from the Incomati and Pongola river systems.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF86FFF3FF31FA67FA64C1B6.taxon	discussion	Remarks. Marcusenius caudisquamatus can easily be distinguished by its shorter and thicker caudal peduncle, (40.4 – 51.37 % of peduncle length as opposed to 26.0 – 40.6 % in M. krameri and 28.7 – 45.66 % in M. pongolensis). The deeper caudal peduncle has more circumpeduncular scales than in other species (19 vs 16 in M. krameri and 16 – 17 for M. pongolensis, Table 7). The dorsal profile of the head of M. caudisquamatus specimens from the Mhlatuze River has a steep, nearly vertical hump in front of the eye. It is slightly concave behind the eye (Fig. 2 c) as compared to being convex in M. pongolensis and M. lucombesi (Figs. 2 a and 2 d, respectively), and straight in M. krameri (Fig. 2 b). The number of GR on the posterior side of the first arch is constant and a reliable character to distinguish this species from both M. pongolensis and M. krameri (Table 6). Compared to M. krameri and M. pongolensis in which the scales series across their body are easily visible, M. caudisquamatus appears smooth and the scales are almost fully embedded in the skin (observed both in live and preserved specimens). The species is genetically closely related to M. krameri (Fig. 4). Sequence divergences between M. krameri and M. caudisquamatus is 0.9 – 2.4 %, and that is much larger that within the latter species (0.0 – 0.5 %). Gill arch, anterior series Gill arch, posterior series Upper limb Lower limb Upper Limb Lower limb Number of GR 4 5 6 4 5 6 6 7 8 7 8 9 M. caudisquamatus 10 * 9 * 1 10 10 M. krameri 13 * 2 15 * 15 * 15 * M. lucombesi 4 * 1 4 * 1 5 * 5 * M. pongolensis 17 4 21 18 3 18 3 Total Gill Rakers Gill arch, anterior series Gill arch, posterior series Number of GR 8 9 10 11 13 14 15 16 17 M. caudisquamatus 9 1 10 * - M. krameri 13 * 2 - 15 * M. lucombesi 3 * 2 5 * - M. pongolensis 17 4 18 3 - Dorsal rays Anal rays Circumpeduncular scales 20 21 22 23 24 25 26 27 28 29 30 31 14 15 16 17 18 19 M. caudisquamatus 6 * 8 1 7 * 8 - 19 * M. krameri 4 26 * 3 2 20 11 * 3 1 35 * - M. lucombesi 1 7 * 5 4 9 * 5 * 5 3 - M. pongolensis 5 15 * 25 5 4 33 * 13 25 * 25 - Coloration. In preservation, specimens are grey to blackish. In 70 % alcohol the upper back and belly are covered with a milky-grey mucus layer, especially on the head where opaque ‘ mormyrid skin’ (electroreceptor organ area) is present.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF86FFF3FF31FA67FA64C1B6.taxon	distribution	Distribution and ecology. Currently this species is known only from the Nseleni and the Mhlatuze rivers in KwaZulu-Natal, South Africa (Fig. 1). The Nseleni River was generally deep with strong currents and all the specimens were collected with gill nets. The Mhlatuze River was wide and shallow, about 1.5 m deep, with a sandy or rocky bottom with dense reed beds on the periphery. Water current was moderate to fast in sections where M. caudisquamatus was collected. The water conductivity was relatively high (503 µS / cm to 707 µS / cm) in summer (November 2009) as compared to the 134 µS / cm recorded by Kramer et al. (2007) in winter (12 August 1999). The water temperature at noon was around 25 – 26 ° C in November 2009, the water was clear and specimens were caught at various depths depending on the sampling station. The Mhlatuze River system has the southernmost record of mormyrids in Africa (Kramer et al. 2007). The biology and ecology of the species is unknown, but the characteristic anal fin notch of sexually mature males was observed from about 96.7 mm SL. The largest male was 250 mm and the largest female was 215 mm SL.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF86FFF3FF31FA67FA64C1B6.taxon	etymology	Etymology. A combination of the Latin words cauda, for tail, and squama for scale. This name refers to the high number of circumpeduncular scales in this species relative to the other members of the genus in South Africa.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF83FFEDFF31F957FEE0C2C4.taxon	materials_examined	Holotype. SAIAB 188295, 116.21 mm SL, male, Mogalakwena River (also known as the Nyl), Modimolle town, Pretoria Street, above a drift, 24 ° 42 ' 04.2 " S, 28 ° 24 ' 40.8 " E, Limpopo River system, South Africa, P. Maake and O. Gon, 14 October 2010. Hologenetype COI (GenBank KJ 174307), Hologenetype cyt b (GenBank KJ 174297). Paratypes. SAIAB 191224, 5: 92 – 117 mm SL, Paragenetype COI (GenBank KJ 174305 and KJ 174306), Paragenetypes cyt b (GenBank KJ 174295 and KJ 174296), same collection data as holotype. MRAC B 3 - 16 - P- 1 - 2, 2: 100 – 107.77 mm SL, Paragenetype COI (GenBank KJ 174308), Paragenetype cyt b (GenBank KJ 174298), same collection data as holotype. SAIAB 188294, 11: 50 – 147 mm SL, Mokolo River, above dam wall (opposite bank) of dam at intersection of roads R 518 and R 510, 23 ° 22 ' 01.7 " S, 27 ° 41 ' 09.6 " E, P. Maake and O. Gon, 12 October 2010. USNM 410774, 2: 119.73 – 130.8 mm SL, same collection data as previous lot. SAIAB 188290, 20: 57 – 197 mm SL, Tshinane River, R 523 below bridge at Ngudza, 22 ° 53 ' 56.3 " S, 30 ° 31 ' 28.7 " E, P. Maake and O. Gon, 10 October 2010. BMNH 2013.9.4.72 - 73, 2: 110 – 142 mm SL, same collection data as previous lot. Non-type specimens. SAIAB 79147, 6: 37.2 – 122 mm SL, Blyde River. SAIAB 188301, 2: 62.2 – 140.92 mm SL, Cross River, Danda village, below bridge, tributary from Nwanedi Dam. SAIAB 84510: 22 mm SL, Damani Dam. SAIAB 84485, 5: 35 – 48 mm SL, Damani Dam. SAIAB 84439, 2: 52 – 79 mm SL, Sambandou Junction. SAIAB 84527, 7: 32 – 56 mm SL, Mbwedi Pumphouse.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF83FFEDFF31F957FEE0C2C4.taxon	diagnosis	Diagnosis. Middle body depth 21.9 – 28.1 % SL; distance from dorsal fin origin to end of caudal peduncle 39.9 – 41.7 % SL, distance from anal fin origin to end of caudal peduncle 42.8 – 44.9 % SL; dorsal fin length 15.5 % – 20.0 % SL; anal fin length 23.2 – 25.5 % SL; depth of caudal peduncle 26.0 – 40.6 % of its length; scales around caudal peduncle 16; total vertebrae 44; anterior GR (5 – 6) + (5) = (10 – 11); posterior GR (8) + (9) = 17; dorsal fin rays 22 – 25; anal fin 28 – 31 rays; scales along lateral line 69 – 72. Conical teeth on upper / lower jaws 4 – 6.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF83FFEDFF31F957FEE0C2C4.taxon	description	Description. Measurements and meristic counts are given in Tables 8. Head with terminal mouth well in front of eye; mental lobe on lower jaw protruding in front of upper jaw; head and body compressed; snout rounded and blunt; pre-anal distance shorter than pre-dorsal distance; distance from origin of anal fin to origin of dorsal fin slightly greater or equal to middle body depth; pre-pelvic distance twice as long as the distance between pelvic and anal fins; dorsal and anal fins set well back on the body, situated about two thirds of standard length from the snout and opposite each other; dorsal fin shorter, originating on vertical at 4 th or 5 th anal-fin ray and ending before anal-fin base; distal margin of dorsal and anal fins obliquely orientated, with rays becoming gradually shorter posteriorly; dorsal fin rays 23 (22 – 25), its anterior rays highest; anal fin rays 29 (28 – 31); males at sexual maturity have kink in anal fin base, distinctly curving inward; anterior anal fin rays of sexually mature males strong and sometimes darker, 3 rd – 5 th rays longer than first two rays, crescentic and rounded or curved backwards, but anteriorly sharp or pointy in females and juveniles; pectoral fin rays 10, fin length 53.7 – 82.3 % of head length; body depth 26.6 % (21.9 – 28.1 %) SL; caudal peduncle slender, subcylindrical across its entire length, 19.2 % (17.6 – 24.5 %) SL; 16 scales (N = 35) around the caudal peduncle; caudal peduncle depth 34.9 % (26.0 – 40.6 %) of its length; lateral line scales 70 (69 – 72), cycloid and with reticulate striae; jaws with 4 – 6 (usually 4 – 5, N = 27) conical teeth; 44 vertebrae (excluding urostyle); gill rakers on anterior side more developed (longer and smooth, i. e. no spines), anterior GRt = 10 (10 – 11) on the first gill arch (Table 6), gill raker of anterior side conicals with thin tips; posterior GRt 17 on the posterior side of first gill arch; gill rakers on posterior side shorter, thicker and with flattened tips. Coloration. Immediately after death specimens are medium brown, speckled with darker, irregular blotches on the sides, fins yellowish (Fig. 2 b). In 70 % alcohol, the head (including eye), fins and upper back sometimes covered by milky-grey mucus layer. Specimens from Tshinane River are similar to those from type locality, but clearly darker from belly to back, including the fins. Electric organ discharge: According to Kramer et al. (2007), the EODs of ten individuals (of which six were examined morphologically in the present study: SAIAB 79147) of this species (referred to as Marcusenius Limpopo population) were distinct from those of M. pongolensis, M. macrolepidotus, M. altisambesi and M.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF83FFEDFF31F957FEE0C2C4.taxon	discussion	Remarks. Compared with M. pongolensis, M. krameri specimens are best distinguished by their moderately short, laterally compressed deep body. Marcusenius krameri also has the greater distance from dorsal fin origin to end of caudal peduncle (39.9 – 41.7 % vs 37.1 – 39.8 % SL) and from anal fin origin to end of caudal peduncle (42.8 – 44.9 % vs 40.8 – 42.1 % SL), and its anal-fin base is longer (23.2 – 25.5 % vs 19.5 – 21.9 % of SL), all of which do not overlap with M. pongolensis. Marcusenius krameri differed clearly from M. macrolepidotus with the dorsal and anal fin length, caudal peduncle depth and body depth, all of which are less and caudal peduncle length and number of scales around caudal peduncle which are more in the latter species. Marcusenius krameri also has the highest number of posterior gill rakers and anal-fin rays among the South African species (Tables 6 and 7, respectively). Caudal peduncle depth of M. krameri increased significantly with size as compared to other species, resulting in the unusually large range of this character. Marcusenius krameri is closely related to M. caudisquamatus, but the latter species has a deeper caudal peduncle into its length (40.4 – 51.7 % vs 26.0 – 40.5) and 19 circumpeduncular scales (see also Tables 6, 7). Marcusenius krameri species can also be distinguished based on the phylogenetic relationships forming a sister species to M. caudisquamatus and its genetic divergence in the in the cyt b dataset (0.9 – 2.4 %) (Fig. 4).	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF83FFEDFF31F957FEE0C2C4.taxon	distribution	Distribution and ecology. Marcusenius krameri occurs in the mainstem of the Limpopo River and several of its tributaries, ranging from the Matlabas River in the west to the Olifants River in the east. Individuals of this species were found in groups. During the day they were commonly encountered below undercut river banks, especially in dense networks of tree roots or reed beds along the stream margins. Places with shallow water and slow-flowing reaches of rivers and streams are especially attractive to them. Large numbers of small juveniles were found in bushes of aquatic reeds and floating grass on the periphery of the river or in side channels. The conductivity in most rivers where it occurred ranged from 55 to 106 µS / cm (type locality), and a pH from 7.41 to 8.5 units in October 2010 (pH = 7.86, in type locality). In summer, water temperature was 24 – 27 ° C between 10 am to 6 pm. Kramer et al. (2007) reported a higher water conductivity of 154 µS / cm and a much colder temperature, 16.7 ° C, between the 25 th and 26 th September 1998. The turbidity where specimens were collected was moderate to high at a depth not exceeding 1.5 m. The biology of the species is unknown, but males mature from an estimated length of 84 mm SL (the characteristic notched anal fin of males was observed at this size), but females probably mature at a larger size. The largest specimens were a 207 mm SL female and a 192 mm SL male.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF83FFEDFF31F957FEE0C2C4.taxon	etymology	Etymology. Marcusenius krameri is named for Prof. Bernd Kramer from the Zoological Institute of the University of Regensburg, Germany, in recognition of his contribution to the systematics of southern African mormyrids.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF9DFFEFFF31FB8AFD23C2C7.taxon	materials_examined	Holotype. SAIAB 73884, 74.0 mm SL, male, Lucombe River, Lucombe stream pools, Nyati Road, Niassa Game Reserve, 12 ° 5 ' 15 '' S, 37 ° 33 ' 38 '' E, Ruvuma River system, R. Bills, 22 August 2003 Hologenetype COI (GenBank KJ 174304), Hologenetype cyt b (GenBank KJ 174293). Paratypes. SAIAB 191226, 6: 52 – 72 mm SL, Paragenetype COI (GenBank KJ 174303), paragenetypes cyt b (GenBank KJ 174294), same collection as the holotype. MRAC, B 3 - 16 - P- 4, 61.85 mm SL, same collection data as holotype. SAIAB 73885, 58.3 mm SL, same collection data as holotype. SAIAB 73886, 2: 48.8 – 55.5 mm SL, upper Lucombe River, isolated swamp pools in forest, Mbatamila-Nyati road, 12 ° 5 ' 2 '' S, 37 ° 33 ' 43 '' E, R. Bills, 26 August 2003. SAIAB 73816, 73 mm SL, Lucombe River, fourth stream, pools in bed, Mbatamila-Matondovela Road, 12 ° 7 ' 49 '' S, 37 ° 26 ' 10 '' E, R. Bills, 26 August 2003. SAIAB 73790, 4: 73.0 – 121.5 mm SL, Nakanjambo River, Nakajambo stream, pools in bed, Mbatamila-Matondovela Road, 12 ° 7 ' 45 '' S, 37 ° 21 ' 41 '' E, R. Bills, 14 August 2003. BMNH 2013.9.4.75, 71 mm SL, same collection data as previous lot.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF9DFFEFFF31FB8AFD23C2C7.taxon	diagnosis	Diagnosis. Distance dorsal fin origin to end of caudal peduncle 40.0 – 43.2 % of SL; distance anal fin origin to end of caudal peduncle 43.6 – 45.9 % of SL; shortest pre-anal length 56.6 – 59.2 % of SL; pre-dorsal length 59.5 – 63.3 % of SL; low caudal peduncle depth 25.0 – 30.8 % into its length; circumpeduncular scales 14 – 16; anal fin rays 28 – 29; dorsal fin rays 22 – 24; total vertebrae 44; lateral line scales 55 – 56; anterior GR (4 – 5) + (4 – 5) = (8 – 10); posterior GR (6) + (7) = 13; jaws with 5 – 5 conical teeth.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF9DFFEFFF31FB8AFD23C2C7.taxon	description	Description. Measurements and meristic counts are given in Table 9. Head with terminal mouth well in front of eye, mental lobe on lower jaw protruding in front of upper jaw; head and body compressed; snout very pointy; pre-anal distance shorter than pre-dorsal distance; distance from origin of anal fin to origin of dorsal fin equal to the middle body depth; pre-pelvic distance twice as long as the distance between pelvic and anal fins; dorsal and anal fins set well back on the body, situated about two thirds of standard length from the snout and opposite each other; dorsal fin shorter, originating on vertical at 3 rd anal fin ray and ending before anal fin base; distal margins of dorsal and anal fins obliquely orientated, with rays becoming gradually shorter posteriorly; dorsal fin rays 23 (22 – 24), its anterior rays highest, with distal margins rounded and slightly concave; anal fin rays 29 (28 – 29); males at sexual maturity have kink in the base of the anal fin distinctly curving in ward; anterior anal fin rays of sexually mature males longer, appear stronger, are crescentic and rounded, but are anteriorly sharp or pointy in females and juveniles; pectoral fins rays 10; pectoral fins distinctly very long, reaching origin of pelvic fin (when pressed against body); middle body depth 28.4 % (25.0 – 28.4 %) of SL; caudal peduncle thinner, sub-cylindrical across its entire length 20.3 % (19.7 – 21.7 %) in SL; 14 (14 – 16) scales along the caudal peduncle circumference; caudal peduncle depth 26.7 % (25.0 – 30.8 %) into its length; jaws with 5 - 5 conical teeth; lateral line scales 55 (55 – 56); lateral line scales cycloid with reticulate striae; 44 vertebrae (excluding urostyle). gill rakers on anterior side covered with many minute spines, anterior GRt = 8 (8 – 9) on the first gill arch (Table 6); posterior GRt 13 on the posterior side of first gill arch; gill rakers on posterior side shorter, thicker and with covered by minute spines. NB: MP = M. pongolensis, ML = M. krameri, MK = M. caudisquamatus Electric organ discharge. No data. Coloration. All specimens light brown when preserved. Homogeneous coloration without any blotches, increasingly lighter on the belly.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF9DFFEFFF31FB8AFD23C2C7.taxon	distribution	Distribution. This species is currently known only from the Lucombe River and its small tributary streams, as well as the Nakajambo River, both tributaries of the Ruvuma River system within the Niassa Game Reserve, Mozambique. It was collected with Petrocephalus catostoma (Günther, 1866) in aquatic weed beds, marginal vegetation and root-stocks of bank vegetation in headwater streams through to floodplain margins (Kramer et al. 2007; R. Bills, pers. comm.). Water conductivity was 104 – 268 ΜS / cm in August 2003, probably reflecting human impact on this aquatic system (Kramer et al. 2012).	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF9DFFEFFF31FB8AFD23C2C7.taxon	discussion	Remarks. Marcusenius lucombesi may be the smallest species of Marcusenius in southern Africa, reaching sexual maturity at only 56 mm SL, with none of the males at the SAIAB collection exceeding 75 mm SL. The largest female was 120.5 mm SL. In comparison to other species examined in this study, M. lucombesi has the lowest number of scales along the lateral line (55 – 56), and the anal fin has a more anterior origin than the dorsal fin by only three rays as compared to 4 – 5 rays in M. krameri and M. caudisquamatus. It should be emphasised that M. lucombesi also differs significantly from M. macrolepidotus of the Lower Zambezi in several characters, i. e. distance from dorsal fin origin to end of caudal peduncle, caudal peduncle depth, pre-anal and dorsal lengths, all of which were greater in M. macrolepidotus (Kramer et al. 2007). Marcusenius lucombesi is also genetically divergent, and morphologically distinct in having significantly smaller caudal peduncle depth, longer dorsal fin and head lengths, and one anal and dorsal fin ray more than in M. devosi (Krame et al. 2007). Another bulldog species, Marcusenius livingstonii, occurs in the Ruvuma but it differs from M. lucombesi in having a dark uniform bar on the body below the origin of the dorsal fin and bicuspid teeth. These two characters also distinguish M. livingstonii from all other known species of Marcusenius in southern and eastern African rivers. Marcusenius lucombesi is therefore distinct from M. livingstonii and M. macrolepidotus.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF9DFFEFFF31FB8AFD23C2C7.taxon	etymology	Etymology. The specific name refers to the type locality of this species, Lucombe River, a tributary of the Ruvuma River system in the Niassa Game Reserve.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF9FFFE9FF31FB87FBF4C35F.taxon	materials_examined	Holotype. ANSP 54950, 172.5 mm SL; Pongola River at Paulpietersburg district, South Africa, KwaZulu-Natal; not examined. Material examined. Limpopo River system: SAIAB 188291, 11: 111 – 193 mm SL, Olifants River, below iron bridge, after Mogologolo village. SAIAB 188303, 4: 115 – 147 mm SL, Olifants River, above Loskop Dam in Aventuur, bridge on dirt road. SAIAB 188302, 20: 55 – 150 mm SL, Groot Letaba River, Broederstroom drift about 10 km from (south) of Tzaneen. SAIAB 188289, 24: 59 – 183 mm SL, Matlabas River, on road R 510 bridge towards Thabazimbi. Incomati River system: SAIAB 188297, 13: 46 – 136 mm SL, Crocodile River, below N 4 bridge at Matsulu village. SAIAB 188300, 4: 129 – 189 mm SL, Omnia farm dam, Kopmeiden. SAIAB 54448, 12: 49 – 118 mm SL, Sabie River, above Hazyview. SAIAB 188280, 2: 77 – 107 mm SL, Sabie River at Hoxani area below dam. SAIAB 188279, 5: 61 – 158 mm SL, Sabie River, Sabana tributary at the back of the Sabie Adventures. SAIAB 188293, 2: 52 – 53 mm SL, Sabie River, Marite. SAIAB 188304, 11: 82 – 136 mm SL, Sand River. Nkomati River system: SAIAB 69893, 2: 65.8 – 78.8 mm SL, Phophonyane River, Ntfonjeni Road. SAIAB 69907, 3: 113 – 252 mm SL, Mlumati River, Driekoppies Dam. SAIAB 69949, 7: 39.0 – 73.5 mm SL, Nyonyane River, Sisa Ranch. Pongola River system: SAIAB 79148, 5: 115.6 – 176.5 mm SL, Pongola River, bridge at road connecting Ndumo with Kosi Bay. SAIAB 188277, 2: 129 – 140 mm SL, Sitilo River, below bridge on N 2 road to Jozini. SAIAB 188296, 16: 79 – 165 mm SL, Pongola River on R 33 road to Paulpietersburg. Kosi River system: SAIAB 88637, 2, Nkanini stream, below bridge just outside Manguzi on road to Mozambique. SAIAB 88633, 11: 98.8 – 147.7 mm SL, Mahlampane River, at Manguze village below bridge. SAIAB 88615, 11: 61.7 – 178.3 mm SL, Siyadla River, below the bridge just about one metre down and above the bridge.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF9FFFE9FF31FB87FBF4C35F.taxon	discussion	Remarks. The measurements and meristic data of Kramer et al. (2007) ’ s M. pongolensis were not always in agreement with our data (Table 10), probably due to the inclusion of the Mhlatuze and Nseleni rivers as a southern population of M. pongolensis. Since these populations are now recognised as a new species, M. caudisquamatus, a new diagnosis for M. pongolensis is necessary.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF9FFFE9FF31FB87FBF4C35F.taxon	diagnosis	Diagnosis and comparison. Marcusenius pongolensis is distinguished from other species in its more slender body (elongated, smaller body depth), with pD (37.1 – 39.8 % SL), pA (40.8 – 42.2 % SL) and LA (19.5 – 21.9 % SL) (Table 10), all shorter and non-overlapping with M. krameri (Table 8), M. caudisquamatus (Table 5) and M. lucombesi (Table 9). Total vertebrae ranges between 42 – 43 vs 44 in other species; posterior total gill rakers (GRt) 13 – 14 vs 15 and 17 in M. caudisquamtus and M. krameri, respectively (Table 6). Lateral line scales 70 – 73. Table 7 provides the range for counts of dorsal and anal fin rays, and circum-peduncular scales in M. pongolensis, M. krameri, M. caudisquamatus and M. lucombesi. Dorsal fin situated about three fourths of standard length from snout. Colour in life of M. pongolensis ranges from a homogeneous medium brown to grey-brown with yellowish to golden sheen, underside lighter, body has darker blotches on sides with yellowish fins (Fig. 2 a). These diagnostic characters agree with those mentioned in Kramer et al. (2007) for the holotype of M. pongolensis. Marcusenius pongolensis is considered closest to M. macrolepidotus based on phylogenetic analyses of the mitochondrial cyt b dataset (Fig. 4), and significant divergence from M. krameri, M. caudisquamatus and M. lucombesi (Table 2). Electric organ discharge. Detailed in Kramer et al. (2007).	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF9FFFE9FF31FB87FBF4C35F.taxon	distribution	Distribution. Kramer et al. (2007) presumed that M. pongolensis was absent from the Limpopo River system. However, M. pongolensis was found in small numbers in the Olifants, Groot Letaba, and Matlabas rivers (Fig. 1). Marcusenius pongolensis therefore occurs from the tropical sections of the Limpopo River system in the north to the Kosi River system in the south.	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
3F5BF765FF9FFFE9FF31FB87FBF4C35F.taxon	biology_ecology	Biology and ecology. The biology and ecology of M. pongolensis is not well understood, but populations from an oligotrophic impoundment in Swaziland matured within the first year, at an estimated minimum length of 134 and 119 mm SL for males and females, respectively (Booth & Khumalo 2010).	en	Maake, Pholoshi A., Gon, Ofer, Swartz, Ernst R. (2014): Descriptions of three new species of Marcusenius Gill, 1862 (Teleostei: Mormyridae) from South Africa and Mozambique. Zootaxa 3780 (3): 455-480, DOI: 10.11646/zootaxa.3780.3.2
