taxonID	type	description	language	source
F54D87B96754FF92FC79FF7C4210588D.taxon	description	(FIGS 8 A, 19; TABLES 1, 7, 8) Labeobarbus sp. ‘ Inkisi’ (Wamuini Lunkayilakio, 2010: 131)	en	Vreven, Emmanuel J. W. M. N., Musschoot, Tobias, Decru, Eva, Lunkayilakio, Soleil Wamuini, Obiero, Kevin, Cerwenka, Alexander F., Schliewen, Ulrich K. (2019): The complex origins of mouth polymorphism in the Labeobarbus (Cypriniformes: Cyprinidae) of the Inkisi River basin (Lower Congo, DRC, Africa): insights from an integrative approach. Zoological Journal of the Linnean Society 186: 414-482
F54D87B96754FF92FC79FF7C4210588D.taxon	materials_examined	Holotype: MRAC A 7 - 009 - P- 0815, Inkisi River basin, Bongolo River, Kinsende village, Lower Congo (5 ° 23 ’ 10.7 ’ S – 15 ° 15 ’ 25.2 ’ E) Coll. S. Wamuini Lunkayilakio, 30 August 2006 (172.6 mm SL) (DNA tag n ° 713). Paratypes: MRAC A 6 - 007 - P- 0519, Inkisi River basin, Kisantuvillage, LowerCongo (5 ° 08 ’ 02.6 ’ S- 15 ° 03 ’ 51.5 ’ E) Coll. Exp. Bas-Congo 2005, 6 October 2005 (123.8 mm SL) (DNA tag n ° 567) – MRAC A 6 - 007 - P- 0535, Inkisi River basin, Ngufu River, at the bridge between Luangu and Kavuaya village, Lower Congo (5 ° 04 ’ 44.2 ’ S- 15 ° 07 ’ 51.0 ’ E) Coll. Exp. Bas-Congo 9 October 2005 (202.9 mm SL) (DNA tag n ° 618) – MRAC A 7 - 009 - P- 0818, Inkisi River basin, Fidi River, Kiyenga village, Lower Congo (5 ° 31 ’ 33.8 ’ S- 15 ° 16 ’ 41.0 ’ E) Coll. S. Wamuini Lunkayilakio, 4 September 2006 (177.2 mm SL) (DNA tag n ° 735) – MRAC A 7 - 009 - P- 0821, Inkisi River basin, Nua River, Mawunzi village, Lower Congo (05 ° 18 ’ 48.0 ’ S- 15 ° 16 ’ 31.9 ’ E) Coll. S. Wamuini Lunkayilakio, 14 February 2007 (139.7 mm SL) (DNA tag n ° 780) – MRAC A 7 - 009 - P- 0822, Inkisi River basin, Kinsende village, Lower Congo (5 ° 23 ’ 33.8 ’ S- 15 ° 15 ’ 13.1 ’ E) Coll. S. Wamuini Lunkayilakio, 31 August 2006 (204.1 mm SL) (DNA tag n ° 730) – MRAC A 9 - 0014 - P- 0375, Inkisi River basin, Nsanga village, downstream of the falls, Lower Congo (4 ° 50 ’ 39.4 ’ S- 14 ° 57 ’ 27.5 ’ E) Coll. S. Wamuini Lunkayilakio, 13 October 2008 (117.7 mm SL) (DNA tag n ° 1076). Differential diagnosis: Within the Congo basin L. nzadimalawu can be distinguished from L. altipinnis, L. ansorgii, L. batesii, L. brauni, L. cardozoi, L. caudovittatus, L. dartevellei, L. fasolt, L. habereri, L. humphri, L. iphthimostoma, L. iturii, L. jubbi, L. longidorsalis, L. longifilis, L. lufupensis, L. macroceps, L. macrolepidotus / cf. macrolepidotus, L. macrolepis, L. mawambi, L. mawambiensis, L. mirabilis, L. nanningsi, L. oxyrhynchus, L. paucisquamatus, L. stappersii, L. trachypterus, L. upembensis and L. wittei by its high number of lateral line scales, i. e. 35 – 41 (vs. less than 34); from L. leleupanus by its low number of lateral line scales, i. e. 37 – 41 (vs. 45 – 47); from L. tropidolepis and L. platyrhinus by its low number of scales between the lateral line and the dorsal and ventral midline, i. e. 4.5 – 6.5 and 5.5 – 6.5 (vs. 7.5 – 8.5 and 7.5 – 9.5 in L. tropidolepis and 6.5 – 7.5 and 6.5 – 8.5 in L. platyrhinus) and from the latter by its low number of circumpeduncular scales as well, i. e. 12 – 16 (vs. 16 – 18); from L. robertsi by the absence of papillae on the anterior edge of the lower jaw (vs. with numerous well identifiable papillae); from L. pellegrini by the presence of two pair of well-developed barbels (vs. a single pair of minute posterior barbels in L. pellegrini); from L. progenys by its non-prognathous lower jaw (vs. prognathous); from L. altianalis and L. gestetneri by the last unbranched dorsal-fin ray not being transformed into a well-developed spine, but instead being clearly segmented for about half of its length, i. e. 42.8 – 57.7 % dorsal-fin height [vs. transformed into a spine, clearly segmented only at its most distal end, i. e. less than 30.0 % dorsal-fin height (data missing for the holotype of L. somereni as the segmented part of the dorsal spine is broken off)]; and from L. somereni, by its high total number of gill rakers on the first gill arch, i. e. 18 – 22 (vs. 11) and a, positively allometric, narrow mouth width, i. e. 16.1 – 26.5 % HL (vs. 31.3 % HL). Further, L. nzadimalawu can be distinguished from both the other members of the Inkisi complex, L. nzadinkisi and the intermediate / hybrid specimens by the presence of a free mental lobe (vs. no mental lobe but instead a cornified Varicorhinus real cutting edge on the outer edge of the lower jaw in L. nzadinkisi and no or only a rudimentary or attached mental lobe in hybrid specimens). In addition, L. nzadimalawu can be distinguished from L. nzadinkisi by its narrow mouth width, 16.1 – 26.5 % HL (vs. 26.8 – 50.5 % HL); long head length, 23.0 – 26.4 % SL (vs. 20.1 – 22.1 % HL); short dorsal-fin base length, 12.1 – 16.0 % SL (vs. 14.4 – 17.9 % SL); and long prepectoral distance, 22.6 – 26.0 % SL (vs. 20.0 – 22.1 % SL) (Figs 13 A, B, 14 A, B). Finally, L. nzadimalawu can be distinguished from Acapoeta tanganicae by its low number of lateral line scales, i. e. 35 – 41 (vs. 57 – 67). Within the adjacent Lower Guinea ichthyofaunal province L. nzadimalawu can be distinguished from L. axelrodi, L. batesii, L. brevispinis, L. cardozoi, L. caudovittatus, L. compiniei, L. habereri, L. fimbriatus, L. jaegeri, L. malacanthus, L. mariae, L. mbami, L. micronema, L. mungoensis, L. roylii, L. sandersi, L. semireticulatus, L. steindachneri, L. tornieri, L. versluysii and L. werneri by its higher number of lateral line scales, i. e. 35 – 41 (vs. less than 34); from L. aspius, L. lucius and L. progenys by its non-prognathous lower jaw (vs. lower jaw clearly prognathous); and from L. rocadasi by its last unbranched dorsal-fin ray not being transformed into a well-developed spine, but instead being clearly segmented over approximately half its length, i. e. 42.8 – 57.7 % dorsal-fin height (vs. last unbranched dorsal-fin ray transformed into a spine, clearly segmented only at its most distal end). Finally, L. nzadimalawu can be distinguished from Sanagia velifera by its high number of lateral line scales, i. e. 35 – 41 (vs. 22 – 24). Within the adjacent Quanza ichthyofaunal province, L. nzadimalawu can be distinguished from L. ansorgii, L. gulielmi, L. jubbi, L. nanningsi, L. rhinophorus, L. rosae and L. roylii by its high number of lateral line scales, i. e. 35 – 41 (vs. less than 34); from L. clarkeae, L. ensifer and L. varicostoma by the absence of papillae on the anterior edge of the lower jaw (vs. with well identifiable papillae); from L. lucius and L. progenys by its non-prognathous lower jaw (vs. lower jaw clearly prognathous); and from L. boulengeri [previously L. latirostris (Boulenger, 1910)], L. ensis, L. girardi, L. steindachneri, L. stenostoma and L. rocadasi by its last unbranched dorsal-fin ray not being transformed into a well-developed spine, but instead being clearly segmented over approximately half its length, i. e. 42.8 – 57.7 % dorsal-fin height (vs. last unbranched dorsal-fin ray transformed into a spine, clearly segmented only at its most distal end). Description: The holotype is illustrated in Figure 19 and Figure 8 A for its mouth phenotype. Meristics and measurements are given in Tables 7 and 8, respectively. Labeobarbus nzadimalawu has a rather shallow body depth, a low dorsal-fin height, and a shallow and elongated caudal peduncle. It is a relatively small-sized Labeobarbus species with a maximum observed size of ± 213 mm SL. Labeobarbus nzadimalawu has a typical Lab. - mouth phenotype (Table 1) characterized by the presence of a free mental lobe of variable size (Figs 8 A, B, 19). Anterior and posterior pair of barbels always present and well-developed (Figs 8 A, B, 19). Never papillae on jaws, neither on upper nor on lower jaw. Some specimens have a nose appendage (Fig. 8 B). Lower jaw always clearly shorter than upper jaw. Tubercles (see Fig. 20) are present on all (≥ 117.7 SL) except the smallest of the examined specimens (62.9 mm SL) and, as such, are most likely present in males and females (not dissected to identify sex). They are apparently present all over the seasons, as one specimen collected in February (small dry season) and several ones collected from August to October (end of major dry season and beginning of the wet season) (see Wamuini Lunkayilakio, 2010) all have them. Usually, tubercles are present all over the snout, i. e. on the dorsum of snout and lateral sides of snout between angle of fleshy lips and eye, and all over the dorsum of head up to nape, i. e. between the posterior edge of the head and the anterior edge of the predorsal area. A few larger specimens (139.7 and about ≥ 200.0 mm SL) have a small number of tubercles below eyes, level with the infraorbital bones up to the preopercular / opercular skin fold. Both specimens with a well-developed nose were largely devoid of tubercles on the dorsum of the snout, i. e. from behind the nose up to the level of the anterior edge of the nostrils. Finally, remainder of body without tubercles. Tubercles are easiest observable in a specimen with a large mental lobe and in another one with a nose appendage [MRAC A 9 - 0014 - P- 0372: 212.5 mm SL (19 / 08 / 2008); and MRAC A 6 - 07 - P- 535: 202.9 mm SL (09 / 10 / 2005)]. Lateral line scales of the examined specimen (Fig. 21) with sinuous and parallel, although sometimes somewhat converging, striae. Number of striae on the posterior edge of these scales between about 34 and 43. The pharyngeal teeth number of the only dissected specimen is 2.3.5. (left) – 5.3.2. (right) with the first tooth of the inner row being absent on the right pharyngeal bone (Fig. 22). Note that on the left pharyngeal bone there are two additional smaller teeth present median of the inner row; those are not firmly attached to the bone but rather loosely embedded in the surrounding soft tissue; one of them already well-developed, the other one is more like a dome-like cusp. Coloration: Live specimens silvery to white-grey on head, lateral side and dorsal midline. Most proximal part of the scales, i. e. the scale pockets, dark grey to blackish and this especially for the scales above the lateral line. The remaining, more distal part of the surface of the scales, silvery, white-grey. Fins darker, greyish-blue, with their distal margins whitish or even translucent. Ventral side and lower lateral side of head and body, below the lateral line, often a yellowish hue. Base of pectoral and pelvic fins sometimes also yellowish. In alcohol, proximal surface of scales generally blackish and this especially for the scales above the lateral line. Absent in some of the larger sized specimens examined (MRAC A 7 - 009 - P- 1141: 210 mm SL; MRAC A 9 - 14 - P- 0372: 213 mm SL), while persistent in others (MRAC A 6 - 07 - P- 0535: 203 mm SL; MRAC A 7 - 009 - P- 0822: 204 mm SL). Fins generally light brown-whitish, although some parts might be blackish. The fin rays themselves yellowish-white. With increasing size, fins more and more blackish, although the rays remain light brown yellowish-white. Distribution: Species endemic to the Inkisi River, Lower Congo River basin above the Zongo Falls (Fig. 23). Currently only known from the DRC part of the basin. Etymology: Before the Christian missionaries arrived, the Inkisi River was locally referred to as the ‘ Nzadi malawu ’ in Kikongo (Kintandu / Kindibu dialects), which means ‘ the river that brings good luck’ (i. e. the porte-bonheur River in French). The part of the river towards the northern border of Angola still bears this name. Species name referring to this ancient name of the river basin to which it seems endemic. Species name to be treated as a noun in apposition (ICZN, 1999: Articles 31.2.1. and 34.2.1), making its gender ending unchangeable. Ecology: A small-scale ecological study of ten fishing stations on the Inkisi basin (Wamuini Lunkayilakio, 2010: tables 4.2, 4.3 4.5, 4.8; Wamuini Lunkayilakio et al., 2010: figs 1, 3) revealed L. nzadimalawu, the Lab. - mouth phenotype species, to be most abundant at Kinsendi, downstream and upstream of the Sanga Dam, where the Inkisi River is wide. In contrast, L. nzadinkisi, the Var. - mouth phenotype species, was less abundant or even absent at those stations. For more details on L. nzadinkisi, see below. Although preliminary, these data seem to reveal important differences in habitat preferences between both species but, unfortunately, no data are available for the intermediate-mouth phenotype specimens or hybrids, which were not included in these analyses. Other specimens examined: MRAC A 7 - 009 - P- 1141, Inkisi River basin, Nsanga village, downstream of dam, Lower Congo (4 ° 50 ’ 39.4 ’ S- 14 ° 57 ’ 27.5 ’ E) Coll. S. Wamuini Lunkayilakio, 23 September 2006 (210.4 mm SL) (DNA tag n ° 891). – MRAC A 9 - 0014 - P- 0372, Inkisi River basin, Boko village, Sawu dia Boko, Bas Congo (5 ° 19 ’ 13.9 ’ S- 15 ° 12 ’ 14.0 ’ E) Coll. S. Wamuini Lunkayilakio, 19 August 2008 (212.5 mm SL) (DNA tag n ° 930). – MRAC A 9 - 0014 - P- 0376 - 0377., Inkisi River basin, Nsanga village, slope of the dam, Lower Congo (4 ° 50 ’ 33.8 ’ S- 14 ° 57 ’ 34.9 ’ E) Coll. S. Wamuini Lunkayilakio, 16 October 2008 (62.9 – 130.2 mm SL) (DNA tag n ° 1113 – 1114).	en	Vreven, Emmanuel J. W. M. N., Musschoot, Tobias, Decru, Eva, Lunkayilakio, Soleil Wamuini, Obiero, Kevin, Cerwenka, Alexander F., Schliewen, Ulrich K. (2019): The complex origins of mouth polymorphism in the Labeobarbus (Cypriniformes: Cyprinidae) of the Inkisi River basin (Lower Congo, DRC, Africa): insights from an integrative approach. Zoological Journal of the Linnean Society 186: 414-482
F54D87B96751FF9CFF03F952420D5879.taxon	description	(FIGS 8 F, 24; TABLES 1, 9, 10) Varicorhinus latirostris non Boulenger, 1910 (Wamuini Lunkayilakio, 2010: 137)	en	Vreven, Emmanuel J. W. M. N., Musschoot, Tobias, Decru, Eva, Lunkayilakio, Soleil Wamuini, Obiero, Kevin, Cerwenka, Alexander F., Schliewen, Ulrich K. (2019): The complex origins of mouth polymorphism in the Labeobarbus (Cypriniformes: Cyprinidae) of the Inkisi River basin (Lower Congo, DRC, Africa): insights from an integrative approach. Zoological Journal of the Linnean Society 186: 414-482
F54D87B96751FF9CFF03F952420D5879.taxon	materials_examined	Holotype: MRAC A 7 - 009 - P- 1180, Inkisi River basin, Bongolo River, Kinsende village, Lower Congo (DRC) (5 ° 23 ’ 10.7 ’ S- 15 ° 15 ’ 25.2 ’ E) Coll. S. Wamuini Lunkayilakio, 30 August 2006 (205.2 mm SL) (DNA tag n ° 721). Paratypes: MRAC A 7 - 009 - P- 1361, Inkisi River, Kinsende village, Lower Congo (DRC) (5 ° 23 ’ 10.0 ’ S- 15 ° 15 ’ 17.1 ’ E) Coll. S. Wamuini Lunkayilakio, 31 August 2006 (182.7 mm SL) (DNA tag n ° 729) – MRAC A 7 - 009 - P- 1377, Inkisi River basin, N’Soni River, Yanama village, Lower Congo (DRC) (5 ° 24 ’ 08.2 ’ S- 15 ° 10 ’ 16.6 ’ E) Coll. S. Wamuini Lunkayilakio, 17 July 2006 (101.2 mm SL) (DNA tag n ° 861) – MRAC A 7 - 009 - P- 1386, Inkisi River basin, Luidi River, Kinua-Nsudi village, Lower Congo (DRC) (5 ° 28 ’ 22.8 ’ S- 15 ° 13 ’ 11.1 ’ E) Coll. S. Wamuini Lunkayilakio, 20 July 2006 (95.7 mm SL) (DNA tag n ° 877) – MRAC A 9 - 014 - P- 0381, Inkisi River basin, Vini River, Ngomina village, Mafukusa, Lower Congo (DRC) (4 ° 47 ’ 42.1 ’ S- 14 ° 56 ’ 06.0 ’ E) Coll. S. Wamuini Lunkayilakio, 11 / 10 / 2008 (203.6 mm SL) (DNA tag n ° 1047). Differential diagnosis: Within the Congo basin L. nzadinkisi can be distinguished from L. altipinnis, L. ansorgii, L. batesii, L. brauni, L. cardozoi, L. caudovittatus, L. dartevellei, L. fasolt, L. habereri, L. humphri, L. iphthimostoma, L. iturii, L. jubbi, L. longidorsalis, L. longifilis, L. lufupensis, L. macroceps, L. macrolepidotus / cf. macrolepidotus, L. macrolepis, L. mawambi, L. mawambiensis, L. mirabilis, L. nanningsi, L. oxyrhynchus, L. paucisquamatus, L. stappersii, L. trachypterus, L. upembensis and L. wittei by its high number of lateral line scales, i. e. 35 – 41 (vs. less than 34); from L. leleupanus by its low number of lateral line scales, i. e. 35 – 41 (vs. 45 – 47); from L. tropidolepis and L. platyrhinus by its low number of scales between the lateral line and the dorsal and ventral midline, i. e. 4.5 – 5.5 and 5.5 (vs. 7.5 – 8.5 and 7.5 – 9.5 in L. tropidolepis and 6.5 – 7.5 and 6.5 – 8.5 in L. platyrhinus) and from the latter by its low number of circumpeduncular scales as well, i. e. 12 – 14 (vs. 16 – 18); from L. robertsi by the absence of papillae on the anterior edge of the lower jaw (vs. with numerous well identifiable papillae); from L. progenys by its non-prognathous lower jaw (vs. prognathous); from L. altianalis, L. gestetneri and L. somereni by its lack of both pairs of barbels (vs. two pair of well-developed barbels); and from L. pellegrini by its short prepelvic length, i. e. 46.5 – 48.5 % SL (vs. 50.6 % SL in L. pellegrini); its short pelvic length, i. e. 17.9 – 21.0 % SL (vs. 21.8 % SL in L. pellegrini); and its large eye, i. e. 29.1 – 34.6 % HL (vs. 27.1 % HL, in L. pellegrini). Further, L. nzadinkisi can be distinguished from the other members of the Inkisi complex, L. nzadimalawu and the intermediate / hybrid specimens by the presence of a cornified Varicorhinus real cutting edge on the outer edge of the lower jaw in combination with (1) the absence of barbels and (2) poorly developed fleshy lips on the lateral side of the lower jaw [vs. never with a cutting edge but instead always with a free mental lobe in combination with (1) two pairs of well-developed barbels and (2) well-developed fleshy lips in L. nzadimalawu. Although a cornified Varicorhinus real cutting edge can be found in some specimens, this most often in combination with (1) at least a single pair of well-developed barbels and (2) well-developed fleshy lips in the hybrid specimens (for details see Table 1)]. In addition, L. nzadinkisi can be distinguished from L. nzadimalawu by its broad mouth width, 26.8 – 50.5 % HL (vs. 16.1 – 26.5 % HL); short head length, 20.1 – 22.1 % SL (vs. 23.0 – 26.4 % SL); long dorsal-fin base length, 14.4 – 17.9 % SL (vs. 12.1 – 16.0 % SL); and short prepectoral distance, 20.0 – 22.1 % SL (vs. 22.6 – 26.0 % SL) (Figs 13 A, B, 14 A, B). Finally, L. nzadinkisi can be distinguished from Acapoeta tanganicae by its low number of lateral line scales, i. e. 35 – 41 (vs. 57 – 67). Within the adjacent Lower Guinea ichthyofaunal province, L. nzadinkisi can be distinguished from L. axelrodi, L. batesii, L. brevispinis, L. cardozoi, L. caudovittatus, L. compiniei, L. habereri, L. fimbriatus, L. jaegeri, L. malacanthus, L. mariae, L. mbami, L. micronema, L. mungoensis, L. roylii, L. sandersi, L. semireticulatus, L. steindachneri, L. tornieri, L. versluysii and L. werneri by its higher number of lateral line scales, 35 – 41 vs. less than 34; from L. aspius, L. lucius and L. progenys by its non-prognathous lower jaw (vs. lower jaw clearly prognathous); and from L. rocadasi by its last unbranched dorsal-fin ray not being transformed into a well-developed spine, but instead being clearly segmented over approximately half its length, or 43.0 – 50.1 % dorsal-fin height (vs. last unbranched dorsal-fin ray transformed into a spine, clearly segmented only at its most distal end). Finally, L. nzadinkisi can be distinguished from Sanagia velifera by its high number of lateral line scales, 35 – 41 (vs. 22 – 24). Within the adjacent Quanza ichthyofaunal province, L. nzadinkisi can be distinguished from L. ansorgii, L. gulielmi, L. jubbi, L. nanningsi, L. rhinophorus, L. rosae and L. roylii by its high number of lateral line scales, 35 – 41 (vs. less than 34); from L. clarkeae, L. ensifer and L. varicostoma by the absence of papillae on the anterior edge of the lower jaw (vs. with well identifiable papillae); from L. lucius and L. progenys by its non-prognathous lower jaw (vs. lower jaw clearly prognathous); and from L. boulengeri [previously L. latirostris (Boulenger, 1910)], L. ensis, L. girardi, L. steindachneri, L. stenostoma and L. rocadasi by its last unbranched dorsal-fin ray not being transformed into a well-developed spine, but instead being clearly segmented over approximately half its length, or 43.0 – 50.1 % dorsal-fin height (vs. last unbranched dorsal-fin ray transformed into a spine, clearly segmented only at its most distal end). Description: The holotype is illustrated in Figure 8 F and Figure 24 for its mouth phenotype. Meristics and measurements are given in Tables 9 and 10, respectively. Labeobarbus nzadinkisi has a rather shallow body depth, low dorsal-fin height, and shallow and elongated caudal peduncle. A relatively small-sized Labeobarbus species with a maximum observed size of ± 205 mm SL. Labeobarbus nzadinkisi has a typical Var. - mouth phenotype (Table 1) characterized by the presence of a cornified cutting edge on the anterior edge of the lower jaw (Fig. 8 F). Anterior and posterior pair of barbels generally absent (Fig. 8 F), but a minute pair of posterior barbels might be present. Lower jaw always clearly shorter than upper jaw. Tubercles (Fig. 20) are present on all specimens (≥ 99.8 SL) except for the smallest examined one (95.7 mm SL) and, as such, are likely to be present in males and females (not dissected to identify sex). Specimens with tubercles were collected only in the major dry season and the beginning of the wet season (June and October) (see Wamuini Lunkayilakio, 2010). In these specimens, tubercles are present at least on the snout and often on the dorsum of the head too, although in a different degree. Large specimens (about ≥ 180.0 mm SL) with conical tubercles over mid-ventral and lateral sides of the snout and the lateral sides of head below the eye reaching posteriorly up to area of the preopercular / opercular skin fold. A few larger tubercles are also present between nostril and eye. Further, numerous and tiny tubercles are scattered over dorsal regions of snout, head and nape, between the posterior edge of the head and the anterior edge of the predorsal area. Finally, remainder of body without tubercles. Tubercles are easiest observable in one of the largest specimens examined [MRAC A 9 - 014 - P- 0381: 203.6 mm SL (11 / 10 / 2006)]. Lateral line scales of the examined specimen (Fig. 21) with less sinuous and parallel striae towards their horizontal midlines but slightly radiating striae towards their upper and lower edge. Number of striae on the posterior edge of these scales between about 46 – 48. The pharyngeal teeth number of the only dissected specimen is 2.3.4. (left) – 5.3.2. (right), with the first tooth of the inner row being absent on the left pharyngeal bone (Fig. 22). Note that on both the left and right pharyngeal bone there are two additional smaller teeth present median of the inner row; these are not firmly attached to the bone but rather are loosely embedded in the surrounding soft tissue; one of them well-developed, the other one is more like a dome-like cusp. Coloration: Live specimens silvery grey on lateral sides of head and body. Dorsal midline of head blackish. Dorsal midline of body, in front, along and behind the dorsal fin, with a yellowish-green silver hue. Proximal part of scales, i. e. scale pockets, darker in colour, especially for the scales above the lateral line. Fins of comparable blackish overall colour with their distal margins whitish translucent. Fin rays whitish. Ventral side of head and body yellowish-white. In alcohol, small-size specimens (± ≤ 130 mm SL) uniformly brownish, becoming darker towards the dorsal midline of head and body. Scale pockets, especially those above the lateral line, typically darker brown due to more numerous melanin dots. Anteriormost ventral part of snout and jaws, ventral side of head and belly yellowish light brown. Fins whitish translucent and even transparent with sparsely set minute dark brown or blackish melanin dots. Large-size specimens (± ≥ 183 mm SL) uniformly dark brown, becoming even darker above the lateral line towards the dorsal midline of the head and body. No distinct scale pocket coloration at first sight, although the proximal part of the scales is dark brown while the overall distal border is translucent or even transparent. Anteriormost ventral part of snout and jaws, ventral side of head and belly yellowish light brown. Head generally somewhat lighter of colour compared to body. Fin tissues faintly brownish or even more blackish. Fin rays often more white-yellowish. Distal edge of fins whitish translucent and sometimes even transparent. Distribution: Species endemic to the Inkisi River, Lower Congo River basin above the Zongo Falls (Wamuini, 2010) (Fig. 23). Currently only known from the DRC part of the basin. Etymology: The current name of the Inkisi River is derived from its local appellation ‘ Nzadi i nkisi ’ (i. e. the river of the ‘ nkisi ’) in Kikongo (Kintandu / Kindibu dialects), referring to the missionaries who threw the ‘ mi - nkisi ’ (‘ fetish-es’) [‘ fetish’ object containing a certain nkisi spirit – the oldest ancestor spirit, which has an entirely spiritual existence until it has been caught and incorporated in a man-made object, a ‘ fetish’, which is referred to as a nkisi (Jackobson-Widding, 1979: 131 – 132)] in the river in their effort to convert the local populations to Christianity. Species name referring to this new name of the river basin to which it appears endemic. In addition, by its reference to the nkisi - objects, indirectly referring to the enigmatic hybridization complex of which this species is a parental species. Species name to be treated as a noun in apposition (ICZN, 1999: Articles 31.2.1. and 34.2.1), making its gender ending unchangeable. Ecology: A small-scale ecological study of ten fishing stations on the Inkisi basin (Wamuini Lunkayilakio, 2010: tables 4.2, 4.3 4.5, 4.8; Wamuini Lunkayilakio et al., 2010: figs 1, 3) revealed that L. nzadinkisi, the Var. - mouth phenotype species, was most abundant in the stations characterized by grass banks, i. e. at Luidi, Bongo, Wungu and Nua, all affluents of the Inkisi. Instead, L. nzadimalawu, the Lab. - mouth phenotype species, was far less abundant at those stations. For more details on L. nzadimalawu, see above. Although preliminary, these data seem to reveal important differences in habitat preferences between both species. Unfortunately, no data are available for the intermediate-mouth phenotype specimens, i. e. hybrids, which were not included in these analyses. Other specimens examined: MRAC A 7 - 009 - P- 1195, Inkisi River basin, Luidi River, Kinua-Nsudi village, Lower Congo (DRC) (5 ° 28 ’ 22.8 ’ S- 15 ° 13 ’ 11.1 ’ E) Coll. S. Wamuini Lunkayilakio, 21 July 2006 (99.8 mm SL) (DNA tag n ° 878) – MRAC A 9 - 014 - P- 0382, Inkisi River basin, Vini River, Ngomina village, Mafukusa, Lower Congo (DRC) (4 ° 47 ’ 42.1 ’ S- 14 ° 56 ’ 06.0 ’ E) Coll. S. Wamuini Lunkayilakio, 11 / 10 / 2008 (122.2 mm SL) (DNA tag n ° 1054) – MRAC A 9 - 014 - P- 0380, Inkisi River basin, Vini River, Kizinga Village, Kindawula, Lower Congo (DRC) (4 ° 46 ’ 58.6 ’ S- 14 ° 56 ’ 47.8 ’ E) Coll. S. Wamuini Lunkayilakio, 11 October 2008 (129.9 mm SL) (DNA tag n ° 1040). INTERSPECIFIC HYBRIDS:	en	Vreven, Emmanuel J. W. M. N., Musschoot, Tobias, Decru, Eva, Lunkayilakio, Soleil Wamuini, Obiero, Kevin, Cerwenka, Alexander F., Schliewen, Ulrich K. (2019): The complex origins of mouth polymorphism in the Labeobarbus (Cypriniformes: Cyprinidae) of the Inkisi River basin (Lower Congo, DRC, Africa): insights from an integrative approach. Zoological Journal of the Linnean Society 186: 414-482
F54D87B9675FFF98FF67F9CC41DC5948.taxon	description	(FIG. 8 C – E; TABLES 11, 12) In general, within the Inkisi basin, the hybrids can be identified by the following unique combination of mouth phenotype related characters (Fig. 8 C – E): (1) the absence of a free mental lobe, although an attached or rudimentary lobe can sometimes be present, (2) most often at least a single, well-developed pair of barbels (i. e. the posterior one) and (3) the presence of quite well-developed fleshy lips on the lateral side of the lower jaw (for details see Table 1). Measurements and counts are provided in Tables 11 and 12. The Inkisi hybrids, as for the parental species, are rather small. Maximum observed size, ± 207 mm SL. Within the Congo basin, the L. nzadimalawu × L. nzadinkisi hybrids can be distinguished from all Labeobarbus species as follows: from L. altipinnis, L. ansorgii, L. batesii, L. brauni, L. cardozoi, L. caudovittatus, L. dartevellei, L. fasolt, L. habereri, L. humphri, L. iphthimostoma, L. iturii, L. jubbi, L. longidorsalis, L. longifilis, L. lufupensis, L. macroceps, L. macrolepidotus / cf. macrolepidotus, L. macrolepis, L. mawambi, L. mawambiensis, L. mirabilis, L. nanningsi, L. oxyrhynchus, L. paucisquamatus, L. stappersii, L. trachypterus, L. upembensis and L. wittei by their high number of lateral line scales, i. e. 35 – 41 (vs. less than 34); from L. leleupanus by their low lumber of lateral line scales, i. e. 35 – 41 (vs. 45 – 47); from L. tropidolepis and L. platyrhinus by their low number of scales between the lateral line and the dorsal and ventral midline, 4.5 – 5.5 and 4.5 – 5.5 (vs. 7.5 – 8.5 and 7.5 – 9.5 in L. tropidolepis and 6.5 – 7.5 and 6.5 – 8.5 in L. platyrhinus) and from the latter by their low number of circumpeduncular scales, i. e. 12 – 14 (vs. 16 – 18); from L. robertsi by the absence of papillae on the anterior edge of the lower jaw (vs. with well identifiable papillae); from L. progenys by their non-prognathous lower jaw (vs. lower jaw clearly prognathous); from L. altianalis and L. gestetneri by the last unbranched dorsal-fin ray not being transformed into a well-developed spine, but instead being clearly segmented over about half its length, i. e. 37.5 – 53.5 % SL [vs. last unbranched dorsal-fin ray transformed into a spine, clearly segmented only at its most distal end, i. e. less than 6.0 % SL (data missing for holotype of L. somereni as the segmented part of the dorsal spine is broken off)]; from, L. pellegrini by their short prepelvic length, i. e. 46.6 – 48.5 % SL (vs. 50.6 % SL in L. pellegrini); their short pelvic length, i. e. 17.9 – 21.0 % SL (vs. 21.8 % SL in L. pellegrini); and their large eye, i. e. 29.1 – 34.6 % HL (vs. 27.1 % HL, in L. pellegrini); and from L. somereni, by their high total number of gill rakers on the first gill arch, i. e. 18 – 22 (vs. 11) and a, positively allometric, narrow mouth width, i. e. 16.1 – 26.5 % HL (vs. 31.3 % HL). Further, the intermediate / hybrid specimens can be distinguished from both the other members of the Inkisi complex, L. nzadimalawu and L. nzadinkisi sp. nov., by the lack of a free mental lobe and, in most cases, the lack of a cornified Varicorhinus real cutting edge on the outer edge of the lower jaw as well (vs. always with a free mental lobe in L. nzadimalawu, and always with a cornified Varicorhinus real cutting edge on the outer edge of the lower jaw and with poorly developed fleshy lips on the lateral sides of the lower jaw in L. nzadinkisi). In those cases where the intermediate / hybrid specimens bear a cornified Varicorhinus real cutting edge, they also have at least one pair of well-developed barbels (vs. no barbels in L. nzadinkisi). Finally, the intermediate / hybrid specimens can be distinguished from Acapoeta tanganicae by its low number of lateral line scales, i. e. 35 – 41 (vs. 57 – 67). Within the adjacent Lower Guinea ichthyofaunal province, the L. nzadimalawu × L. nzadinkisi hybrids can be distinguished from L. axelrodi, L. batesii, L. brevispinis, L. cardozoi, L. caudovittatus, L. compiniei, L. habereri, L. fimbriatus, L. jaegeri, L. malacanthus, L. mariae, L. mbami, L. micronema, L. mungoensis, L. roylii, L. sandersi, L. semireticulatus, L. steindachneri, L. tornieri, L. versluysii and L. werneri by their higher number of lateral line scales, i. e. 37 – 41 (vs. less than 34 scales); from L. aspius, L. lucius and L. progenys by their non-prognathous lower jaw (vs. lower jaw clearly prognathous); and from L. rocadasi by their last unbranched dorsal-fin ray not being transformed into a well-developed spine, but instead being clearly segmented over approximately half its length, i. e. 37.5 – 53.5 % SL (vs. last unbranched dorsal-fin ray transformed into a spine, clearly segmented only at its most distal end). Finally, the intermediate / hybrid specimens can be distinguished from Sanagia velifera by their high number of lateral line scales, 35 – 41 (vs. 22 – 24). Within the adjacent Quanza ichthyofaunal province, the L. nzadimalawu × L. nzadinkisi hybrids can be distinguished from L. ansorgii, L. gulielmi L. jubbi, L. nanningsi, L. rhinophorus, L. rosae and L. roylii by their high number of lateral line scales, i. e. 35 – 41 (vs. less than 34); from L. clarkeae, L. ensifer and L. varicostoma by the absence of papillae on the anterior edge of the lower jaw (vs. with well identifiable papillae); from L. lucius and L. progenys by their non-prognathous lower jaw (vs. lower jaw clearly prognathous); and from L. ensis, L. girardi, L. boulengeri [previously L. latirostris (Boulenger, 1910)], L. steindachneri, L. stenostoma and L. rocadasi by their last unbranched dorsal-fin ray not being transformed into a well-developed spine, but instead being clearly segmented over approximately half its length, i. e. 37.5 – 53.5 % SL (vs. last unbranched dorsal-fin ray transformed into a spine, clearly segmented only at its most distal end). Tubercles (Fig. 20) are present on all examined specimens (105.6 – 206.5 mm SL) and, as such, are most likely present in males and females (not dissected to identify sex). They are apparently present all over the year as one specimen collected in February (small dry season) and several ones collected from August to October (the second half of the major dry season and the beginning of the wet season) (see Wamuini Lunkayilakio, 2010) all have them. Usually, tubercles are present all over the snout, i. e. from the dorsal region of the snout and lateral snout sides between angle of fleshy lips and eye and all over the dorsum of head up to nape, i. e. between the posterior edge of the head and the anterior edge of the predorsal area. Some specimens had a small number of tubercles below eyes, i. e. at the level of the infraorbital bones. A few specimens had tubercles on the cheek and two specimens, of 117.7 and 196.4 mm SL, respectively, up to the area of the preopercular / opercular skin fold. Finally, remainder of body without tubercles. Tubercles are easiest observable in one of the largest specimens examined [MRAC A 6 - 007 - P- 0459 - 0460: 161.1 (09 / 10 / 2005)]. Lateral line scales of the examined specimen (Fig. 21) with less sinuous but parallel, and even somewhat converging, striae. Number of striae on the posterior edge of these scales between about 28 – 33. The pharyngeal teeth number of the only dissected specimen is 2.3.5. (left) – 4.3.2. (right), with the first tooth of the inner row absent on the right pharyngeal bone (Fig. 22). Note that on the left pharyngeal bone there is an additional small tooth present median of the inner row, which is not firmly attached to the bone but is rather loosely embedded in the surrounding soft tissue. Live colour pattern most comparable to that of L. nzadinkisi, i. e. silvery grey on lateral sides of head and body. Dorsal midline of head blackish. Dorsal midline of body, in front, along and behind the dorsal fin, with a yellowish-green silver hue. Proximal part of scales, i. e. scale pockets, darker in colour and this especially for the scales above the lateral line. Fins of comparable blackish overall colour with their distal margins whitish translucent. Fin rays whitish. Ventral side of head and body yellowish-white. Preserved specimens with a uniformly brownish overall body colour, becoming darker towards the dorsal midline of head and body. Scale pockets, especially those above the lateral line, typically darker brown. Anteriormost ventral part of snout and jaws, ventral side of head and belly yellowish light brown. Fins whitish translucent and even transparent with sparsely set minute blackish melanin dots. In at least one of the large-size specimens, the marked distinct scale pocket coloration is absent (MRAC A 6 - 007 - P- 0538) (207 mm SL). Head generally somewhat lighter of colour compared to body. Fin tissue dark brown, blackish due to the presence of numerous minute dark brown, blackish melanin dots. Fins rays generally more white-yellowish except, sometimes, for the longest upper and lower rays of the caudal fin, which are also dark brown or blackish. Distal edge of fins still whitish translucent and sometimes even transparent. Hybrids, as for both parental species, seem to be endemic to the Inkisi River basin above the Zongo Falls (Fig. 23). Currently only known from the DRC part of the basin. Specimens examined: MRAC A 6 - 007 - P- 0459 - 0460, Inkisi River basin, Ngufu River, at bridge village Luangu & Kavuaya, Lower Congo (DRC) (5 ° 04 ’ 44.2 ’ S- 15 ° 07 ’ 51.0 ’ E) Coll. Lower Congo Expedition 2005, 09 / 10 / 2005 (140.3 – 161.1 mm SL) (DNA tag n ° 621 – 622) – MRAC A 6 - 007 - P- 0536 - 0537, Inkisi River basin, Ngufu River, at bridge village Luangu & Kavuaya, Lower Congo (DRC) (5 ° 04 ’ 44.2 ’ S- 15 ° 07 ’ 51.0 ’ E) Coll. Lower Congo Expedition 2005, 09 / 10 / 2005 (146.1 – 157.3 mm SL) (DNA tag n ° 619 – 620) – MRAC A 6 - 007 - P- 0538, Inkisi River basin, Ngufu River, at bridge village Luangu village Kavuaya, Lower Congo (DRC) (5 ° 04 ’ 44.2 ’ S- 15 ° 07 ’ 51.0 ’ E) Coll. Lower Congo Expedition 2005, 09 / 10 / 2005 (206.8 mm SL) (DNA tag n ° 615) – MRAC A 7 - 009 - P- 0851 - 0852, Inkisi River basin, Ngeba / Ngufu River, village Ngeba, Lower Congo (DRC) (5 ° 11 ’ 01.5 ’ S- 15 ° 12 ’ 23.1 ’ E) Coll. S. Wamuini Lunkayilakio, 25 / 08 / 2006 (125.6 – 130.3 mm SL) (DNA tag n ° 685 & 688) – MRAC A 7 - 009 - P- 0853, Inkisi River basin, Nua River, village Mawunzi, Lower Congo (DRC) (5 ° 18 ’ 48.0 ’ S- 15 ° 16 ’ 31.9 ’ E) Coll. S. Wamuini Lunkayilakio, 14 / 02 / 2007 (129.5 mm SL) (DNA tag n ° 782) – MRACA 7 - 009 - P- 0856 - 0857, Inkisi River basin, River Lukusu, village Ngeba / Mboma, Lower Congo (DRC) (5 ° 13 ’ 36.9 ’ S- 15 ° 13 ’ 02.0 ’ E) Coll. S. Wamuini Lunkayilakio, 23 August 2006 (117.7 – 196.4 mm SL) (DNA tag n ° 662 – 663) – MRAC A 7 - 009 - P- 1173, Inkisi River basin, Nsanga village, downstream of dam, Lower Congo (DRC) (4 ° 50 ’ 39.4 ’ S- 14 ° 57 ’ 27.5 ’ E) Coll. S. Wamuini Lunkayilakio, 23 September 2006 (145.0 mm SL) (DNA tag n ° 896) – MRAC A 7 - 009 - P- 1372, Inkisi River basin, riv. Muala, village Muala-Kinsende, Lower Congo (DRC) (5 ° 16 ’ 11.4 ’ S- 14 ° 57 ’ 42.1 ’ E) Coll. S. Wamuini Lunkayilakio, 11 July 2006 (114.8 mm SL) (DNA tag n ° 813) – MRAC A 7 - 009 - P- 1373, same data (114.4 mm SL) (DNA tag n ° 814) – MRAC A 7 - 009 - P- 1376, Inkisi River basin, riv. Mingididi, village Mawunzi, Lower Congo (DRC) (5 ° 19 ’ 20.0 ’ S- 15 ° 16 ’ 49.9 ’ E) Coll. S. Wamuini Lunkayilakio, 14 July 2006 (105.6 mm SL) (DNA tag n ° 835) – MRAC A 7 - 009 - P- 1378, Inkisi River basin, Luguga River, village Kiyanika, Lower Congo (DRC) (5 ° 16 ’ 32.7 ’ S- 15 ° 12 ’ 36.3 ’ E) Coll. S. Wamuini Lunkayilakio, 28 August 2006 (122.4 mm SL) (DNA tag n ° 706) – MRAC A 7 - 009 - P- 1382, Inkisi River basin, Nua River, village Mawunzi, Lower Congo (DRC) (5 ° 18 ’ 48.0 ’ S- 15 ° 16 ’ 31.9 ’ E) Coll. S. Wamuini Lunkayilakio, 15 July 2006 (170.7 mm SL) (DNA tag n ° 838) – MRAC A 9 - 014 - P- 0004, Inkisi River basin, village Zongo, upstream of dam, Lower Congo (DRC) (4 ° 47 ’ 22.4 ’ S- 14 ° 54 ’ 35.5 ’ E) Coll. S. Wamuini Lunkayilakio, 9 October 2008 (188.6 mm SL) (DNA tag n ° 1035) – MRAC A 9 - 014 - P- 0007, Inkisi River basin, Bongolo River, village Kinsende, Ndimba Kitadi, Lower Congo (DRC) (5 ° 23 ’ 10.7 ’ S- 15 ° 15 ’ 28.2 ’ E) Coll. S. Wamuini Lunkayilakio, 23 August 2008 (133.1 mm SL) (DNA tag n ° 938).	en	Vreven, Emmanuel J. W. M. N., Musschoot, Tobias, Decru, Eva, Lunkayilakio, Soleil Wamuini, Obiero, Kevin, Cerwenka, Alexander F., Schliewen, Ulrich K. (2019): The complex origins of mouth polymorphism in the Labeobarbus (Cypriniformes: Cyprinidae) of the Inkisi River basin (Lower Congo, DRC, Africa): insights from an integrative approach. Zoological Journal of the Linnean Society 186: 414-482
