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Haplochromis squamipinnis Regan, 1921 Figs 1–2, 38–40; Table 1 Haplochromis squamipinnis Regan, 1921: 636. Haplochromis squamipinnis – Trewavas 1933: 338 (redescription). — Greenwood 1973: 204, fig. 31 (redescription). Harpagochromis squamipinnis – Greenwood 1980: 13. Differential diagnosis Species with a piscivorous morphology; body rather deep [BD 32.4–39.3 (mean 35.7) % SL]; oral jaws very long [LJL 47.8–58.6 (52.7) % HL], narrow [LJW 32.6–44.7 (37.2) % LJL], and steep (gape inclination 30–45°); outer oral teeth many and small [UOT 39–79 (median 58)]; dominant males slate blue. Amongst piscivorous species from the Lake Edward system, H. squamipinnis differs from all by presence vs absence of minute scales on proximal part of dorsal fin (rarely few scales present in H. quasimodo sp. nov.). It further differs from H. latifrons sp. nov. and H. mentatus by the combination of small vs large outer oral teeth, a larger number of outer upper jaw teeth [UOT 39–79 (58) vs 22–47 (27–36)], a steeper gape (30–45° vs 15–30°), and a deeper body [BD 32.4–39.3 (35.7) vs 27.2–32.3 (28.6–31.2) % SL]; from H. mentatus by dominant males uniformly slate blue vs yellow-green with a red anterior part of flank. It further differs from H. rex sp. nov., H. simba sp. nov., H. glaucus sp. nov., and H. aquila sp. nov. by the combination of small vs large outer oral teeth, a larger number of outer upper jaw teeth [UOT 39–79 (58) vs 22–47 (27–36)], and dominant males uniformly slate blue vs cream-coloured with an orange operculum, yellow with an orange anterior part of flank, light blue with a dusky to black head, or light grey with a black head, respectively; further from H. rex sp. nov., H. simba sp. nov., and H. glaucus sp. nov. by a steeper gape (30–45° vs 15–30°); further from H. aquila sp. nov. by a smaller eye [ED 23.1–29.7 (26.6) vs 30.0–31.5 (30.6) % HL]. It further differs from H. kimondo sp. nov. by a concave to straight vs convex dorsal outline of head, a gentler snout inclination (30–40° vs 40–50°), and dominant males slate blue vs grey dorsally and yellow ventrally; further from H. falcatus sp. nov. by a shorter head [HL 35.1–36.9 (36.0) vs 36.6– 39.6 (38.2) % SL] and dominant males slate blue vs olive-green with an orange-red anterior part of flank; further from H. curvidens sp. nov. and H. pardus sp. nov. by a deeper cheek [ChD 24.9–36.0 (29.0) vs 20.8–24.9 (22.5–23.2) % HL]; further from H. pardus sp. nov. by a larger adult size (max. 211 vs 96 mm SL) and colour pattern of small specimens (H. quasimodo sp. nov. by the combination of a broader interorbital area [IOW 48.6–55.6 (51.9) vs 40.5–48.7 (43.9) % HW], a longer lower jaw [LJL 47.8–58.6 (52.5) vs 44.2–49.6 (47.1) % HL], a steeper gape inclination (30–45° vs 20–35°), and dominant males slate blue vs light grey dorsally and blue-black ventrally. Etymology Specific name not explained in original description, from the Latin ‘ squamus ’ for ‘scale’, and ‘ pinnis ’ for ‘fin’; probably referring to minute scales on basal parts of dorsal and anal fins. Material examined Holotype DEMOCRATIC REPUBLIC OF THE CONGO (most likely) • 1 ♀, 136.9 mm SL; Lake Edward; 1907– 1908; H. Schubotz leg.; NHMUK 1914.4.8.32. Other material DEMOCRATIC REPUBLIC OF THE CONGO • 1 ♀, 75.9 mm SL; “Lac Edouard: Bugazia” [Lake Edward: Bugazia]; 0°23′40.8″ S, 29°23′02.0″ E (inferred); 16 May 1935; IRSNB 12939 • 1 ♂, 168.4 mm SL; “Lac Edouard: au large de la riv. Talia” [Lake Edward: offshore Talia River]; 0°31′05″ S, 29°20′26″ E (inferred); 23 Apr. 1953; KEA exped. leg.; IRSNB 13475 • 1 ♀, 167.7 mm SL; “Lac Edouard: au large de la riv. Kigera” [Lake Edward: offshore of the Kigera River]; 0°29′42″ S, 29°38′14″ E (inferred); 25 May 1953; KEA exped. leg.; IRSNB 13477 • 1 ♀, 210.5 mm SL; “Lac Edouard: 2–3 km à l’Ouest de Kiavinionge” [Lake Edward: 2–3 km west of Kiavinionge]; 0°11′39″ S, 29°32′31″ E (inferred); 1 Jun. 1953; KEA exped. leg.; IRSNB 13482. UGANDA – Lake Edward • 2 ♀♀, 110.9, 117.4 mm SL; 0°12′00.0″ S, 29°47′38.4″ E; 23 Oct. 2016; HIPE1 exped. leg.; deep catch, open water ± 20 m deep; RMCA 2016.035.P.0235, 0238 • 2 ♂♂, 182.0, 211.4 mm SL; 0°24′16.0″ S, 29°46′24.8″ E; 9 Nov. 2016; HIPE1 exped. leg.; bought at Rwenshama landing site; RMCA 2016.035.P.0251, 0254 • 1 ♂, 2 ♀♀, 148.5–177.3 mm SL; mouth of Kazinga Channel, hard substrate; 0°12′14.4″S, 29°52′37.2″ E; 23 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0375, 0377, 0379 • 1 ♂, 105.4 mm SL; Rwenshama rocky shore; 0°24′05.7″ S, 29°46′35.1″ E; 25 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0385 • 1 ♀, 113.2 mm SL; Kayanja, offshore; 0°05′31.2″ S, 29°45′30.3″ E; 21 Jan. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0368. – Kazinga Channel • 1 ♂, 111.3 mm SL; near Queen Elisabeth Bush Lodge; 0°08′09.6″ S, 30°02′27.6″ E; 28 Oct. 2016; HIPE1 exped. leg.; RMCA 2016.035.P.0244. – Lake George • 1 ♂, 80.7 mm SL; Akika Island; 0°01′26.7″ S, 30°09′38.2″ E; 28 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0387 • 1 ♀ (90.9 mm SL); Akika Island; 0°01′26.7″ S, 30°09′38.2″ E; 29 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0398 • 1 ♂, 180.1 mm SL; Kashaka bay, north of inlet; 0°04′52.2″ S, 30°10′47.3″ E; 2 Feb. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0369 • 2 ♀♀, 77.6, 107.4 mm SL; Kashaka bay, north of inlet; 0°04′52.2″ S, 30°10′47.3″ E; 2 Feb. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0371 to 0372. Description Based on 20 specimens (75.9–211.4 mm SL); body average in depth in comparison to generalised H. elegans (but deep for a piscivorous species; Table 1) and oval to rhomboid (Fig. 38). Head long, narrow, and with a straight to concave dorsal outline; eye small; interorbital area narrow; cheek and lacrimal deep. Snout long, acute, and slopes very gently at 30–35°; premaxillary pedicel very long and prominent. Jaws isognathous to strongly prognathous, slim, very narrow, and rounded in dorsal view; upper jaw long and lower jaw very long; gape large and slopes steeply at 30–45°; maxilla extends to between verticals through anterior margins of orbit and pupil. Lower jaw shallow and with a straight ventral outline in lateral view, mental prominence weakly or strongly developed, and lower jaw side steep with an inclination of 35° to horizontal in anterior view. Upper jaw expanded slightly anteriorly and ventrally. Lips and oral mucosa thin. Neurocranium average in depth, ethmo-vomerine block horizontally inclined, preorbital region shallow (19–25% NL), orbital region average in depth (28–32% NL), and supraoccipital crest deep and pyramidical or weakly wedge-shaped (Fig. 39b). Outer oral teeth numerous, unicuspid, and small. Necks stout, conical, and straight; crowns weakly recurved in lower jaw, recurved in upper jaw, and acutely pointed. Dental arcades rounded. Outer teeth closely and regularly set with neck-distances of ½–1 neck-width. In upper jaw, 1–3 posteriormost teeth sometimes slightly enlarged. Inner teeth small, weakly recurved, unicuspid in large specimens (> 120 mm SL), tri- to rarely unicuspid in upper jaw and uni- to weakly tricuspid in lower jaw of small specimens (H. nigripinnis Regan, 1921 followed by H. angustifrons Boulenger, 1914 (Moreau et al. 1993); insects and to a lesser degree small fishes and plant fragments contribute to diet in small specimens (et al. 1973). Sexual dimorphism in size, as observed by Greenwood (1973), seems absent; both sexes reach> 210 mm SL., Published as part of Vranken, Nathan, Steenberge, Maarten Van, Heylen, Annelies, Decru, Eva & Snoeks, Jos, 2022, From a pair to a dozen: the piscivorous species of Haplochromis (Cichlidae) from the Lake Edward system, pp. 1-94 in European Journal of Taxonomy 815 on pages 68-73, DOI: 10.5852/ejt.2022.815.1749, http://zenodo.org/record/6484153, {"references":["Regan C. T. 1921. The Cichlid fishes of Lakes Albert Edward and Kivu. Annals and Magazine of Natural History Series 9: 632 - 639. https: // doi. org / 10.1080 / 00222932108632633","Trewavas E. 1933. Scientific results of the Cambridge expedition to the East African Lakes, 1930 - 1. The cichlid fishes. Zoological Journal of the Linnean Society 38: 309 - 341. https: // doi. org / 10.1111 / j. 1096 - 3642.1933. tb 00062. x","Greenwood P. H. 1973. A revision of the Haplochromis and related species (Pisces: Cichlidae) from Lake George, Uganda. Bulletin of the British Museum (Natural History) Zoology 25: 139 - 242.","Greenwood P. H. 1980. Towards a phyletic classification of the ' genus' Haplochromis (Pisces, Cichlidae) and related taxa. Part II. Bulletin of the British Museum (Natural History) Zoology 39: 1 - 101. https: // doi. org / 10.5962 / bhl. part. 13268","Schraml E. 2004. Die artenvielfalt der fische in Ugandas gewassern: beitrage zur kenntnis der fischfauna Ugandas. DCG-Informationen Sonderheft 3: 2 - 48.","Moreau J., Christensen V. & Pauly D. 1993. A trophic ecosystem model of Lake George, Uganda. In: Christensen V. & Pauly D. (eds) Trophic Models of Aquatic Ecosystems. ICLARM Conference Proceedings 26: 124 - 129.","Moriarty D. J. W., Darlington J. P. E. C., Dunn I. G., Moriarty C. M. & Tevlin M. P. 1973. Feeding and grazing in Lake George, Uganda. Proceedings of the Royal Society B: Biological Sciences 184: 299 - 319."]}

Haplochromis aquila sp. nov. urn:lsid:zoobank.org:act: 18BC8C85-5A02-48F0-8D90-7B069122A09D Figs 1–2, 20–22; Table 1 Differential diagnosis Species with a piscivorous morphology; eye large [ED 30.0–31.5 (mean 30.6) % HL]; outer oral teeth few and large [UOT 25–37 (median 31)]; dominant males light grey with a black head and a bright red anal fin. Amongst piscivorous species from the Lake Edward system, H. aquila sp. nov. differs from H. latifrons sp. nov., H. mentatus, H. rex sp. nov., H. simba sp. nov., and H. glaucus sp. nov. by a larger eye [ED 30.0–31.5 (30.6) vs 22.2–29.9 (24.4–28.3) % HL]. It further differs from H. latifrons sp. nov. and H. mentatus by a shorter caudal peduncle [CPL 14.6–15.4 (15.0) vs 15.7–18.0 (16.6–17.0) % SL]; further from H. latifrons sp. nov. by the absence vs presence of a well-defined mid-lateral band; further from H. mentatus by dominant males uniformly light grey vs yellow-green with a red anterior part of flank. It further differs from H. rex sp. nov. by a shallower lacrimal [LaD 17.0–19.1 (18.3) vs 18.9–22.5 (20.8) % HL] and dominant males light grey with black operculum and snout vs cream-coloured with an orange operculum and a light blue snout. It further differs from H. simba sp. nov. by a larger number of scales between first anal-fin spine and upper lateral line (ULL-A 12–15, rarely 11 vs 9–11), an absent vs strongly developed mental prominence, and dominant males light grey with a black head vs uniformly yellow with an orange anterior part of flank. It further differs from H. glaucus sp. nov. by a broader head [HW 40.1–43.7 (42.0) vs 38.9–40.9 (39.7) % HL] and dominant males light grey with bright red anal and caudal fins vs light blue with crimson anal and caudal fins. It differs from H. kimondo sp. nov., H. falcatus sp. nov., H. curvidens sp. nov., H. pardus sp. nov., H. quasimodo sp. nov., and H. squamipinnis by the combination of large vs small outer oral teeth and smaller number of outer upper jaw teeth [UOT 25–37 (31) vs 39–79 (45–58)]. It further differs from H. kimondo sp. nov., H. falcatus sp. nov. and H. quasimodo sp. nov. by absence vs mostly presence of a well-defined mid-lateral band and dominant males light grey with a black head vs grey dorsally and yellow ventrally, olive-green with an orange-red anterior part of flank, or light grey dorsally and blue-black ventrally; further from H. kimondo sp. nov. by a narrower head [HW 40.1–43.7 (42.0) vs 42.9–48.0 (45.1) % HL]. It further differs from H. curvidens sp. nov. and H. pardus sp. nov. by a deeper cheek [ChD 26.8–30.8 (28.3) vs 20.8–24.9 (22.5–23.2) % HL]; further from H. pardus sp. nov. by dominant males light grey vs speckled to uniformly black. It further differs from H. squamipinnis by larger eyes [ED 30.0–31.5 (30.6) vs 23.1–29.7 (26.6) % HL] and dominant males light grey vs slate blue. Etymology Specific name from the Latin ‘ aquila ’ for ‘eagle’; referring to predatory morphology and large eyes. Material examined Holotype UGANDA • ♂, 113.6 mm SL; Lake Edward, Kayanja offshore; 0°05′31.2″ S, 29°45′30.3″ E; 20 Jan. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0355. Paratypes UGANDA – Lake Edward • 1 ♀, 2 ♂♂, 83.9–117.7 mm SL; islands near Katwe; 0°10′04.9″ S, 29°52′27.4″ E; 18 Jan. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0349 to 0351 • 1 ♀, 1 ♂, 108.7, 117.6 mm SL; islands near Katwe; 0°10′04.9″ S, 29°52′27.4″ E; 19 Jan. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0352 to 0353 • 1 ♂, 85.6 mm SL; mouth of Kazinga Channel; 0°12′34.8″ S, 29°53′01.5″ E; 19 Jan. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0354 • 1 ♂, 122.9 mm SL; 0°24′16.0″ S, 29°46′24.8″ E; 24 Jan. 2018; HIPE3 exped. leg.; bought at Rwenshama landing site; RMCA 2018.008.P.0356. Description Based on 8 specimens (83.9–122.9 mm SL); body shallow (Table 1) and oval (Fig. 20). Head long, narrow, and with a straight to gently convex dorsal outline; interorbital area narrow; eye average in size in comparison to generalised H. elegans (but large for a piscivorous species); cheek deep; lacrimal average in depth. Snout average in length, rounded in dorsal view, narrow, relatively blunt, and slopes gently at 35–45°; premaxillary pedicel long and (strongly) prominent. Jaws isognathous to weakly prognathous, long, narrow, and rounded in dorsal view; gape large and slopes at 25–35°; maxilla extends to vertical through anterior margin of pupil. Lower jaw relatively stout and with straight ventral outline in lateral view, mental prominence absent or weakly developed, and lower jaw side nearly flat with an inclination of 25–35° to horizontal in anterior view. Upper jaw expanded anteriorly. Lips and oral mucosa relatively large. Neurocranium average in depth, ethmo-vomerine block decurved, preorbital region shallow (18–24% NL), orbital region average in depth (30–33% NL), and supraoccipital crest shallow and wedge-shaped (Fig. 21b). Outer oral teeth few, unicuspid, and very large. Necks stout, conical, and straight; crowns straight to weakly recurved and acutely pointed. Dental arcades with anterior half expanded laterally. Outer teeth widely and irregularly set with neck-distances of 1–4 neck-widths. No enlarged teeth posterior in upper jaw. Inner teeth small, recurved, a mixture of unicuspid and weakly tricuspid, weakly tricuspid rare in large specimens (> 100 mm SL), and acutely pointed in all specimens. Tooth bands very slender crescent-shaped with 1–2 rows of inner teeth, and narrow posteriorly until only outer row remains past ¾ length of tooth band in upper jaw, past ⅔ length of tooth band in lower jaw. Inner teeth closely and regularly set on 1 outer neck-width from outer row; implantation recumbent; size uniform throughout tooth band. Lower pharyngeal bone long, very narrow, slim, and shallow over whole length (Fig. 22). Pharyngeal teeth relatively large and slender; major cusps acutely pointed; cusp gaps concave; minor cusps and cusp protuberances very small. Teeth in two median longitudinal rows equal in size and form to lateral teeth, 12 in each row. Posterior transverse row with 16–17 teeth, implanted erectly with a lateral inclination; major cusps nearly straight, bluntly pointed, and laterally compressed; minor cusps mostly absent. Chest scales small; transition to larger flank scales gradual. Minute scales on proximal half of caudal fin. Caudal fin emarginate; dorsal and anal fins reach to between verticals through caudal-fin base and two scales anterior to this vertical. Pectoral fin reaches to between genital opening and second anal-fin spine; pelvic fin reaches to genital opening in females, to between first and second anal-fin spines in males; first pelvic fin slightly elongated in all specimens. Ceratobranchial gill rakers in outer row of first gill arch short, stout, and simple; posteriormost rakers anvil-shaped or bifid. Epibranchial gill rakers slender and simple. Colouration in life Dominant males: body metallic grey; dorsum greyish; belly speckled black; chest, cheek, snout, lower jaw, and lips black; eye with (dark) grey outer ring and golden inner ring (Fig. 21c). Nostril, interorbital, and lacrimal stripes very faint. Pectoral fin hyaline, pelvic fin black; dorsal fin sooty and with black lappets; anal and caudal fins bright red and with dusky bases; anal fin with dusky posterior part and 1–2 large orange egg-spots with dusky rings. Females and juveniles: body and dorsal part of head silver with yellowish sheen; belly, chest, operculum, cheek, lower jaw, and lips white; snout dusky; eye with (dark) grey outer ring and silver inner ring (Fig. 21d). Nostril and interorbital stripes faint. Pectoral fin hyaline, pelvic fin white; dorsal and caudal fins dusky; dorsal fin with black lappets; caudal fin with a yellowish base and a faint red distal part; anal fin yellow with a red sheen. Preserved colouration Body brown; dorsum dark-brown; ventral part of body and operculum yellowish; chest and lower jaw black in males; cheek yellowish in females, dark-brown in males; snout dusky (Fig. 21a). Nostril and interorbital stripes faint; lacrimal stripe narrow and well-defined; mental blotch present; vertical preopercular stripe well-defined in males. Pectoral fin hyaline; pelvic fin yellowish with blackish first rays in females, black in males; dorsal fin dusky and with black lappets and maculated posterior part; anal fin yellowish with a dusky overlay and, in males, with black lappets and 1–2 large egg spots; caudal fin with a dusky base, a hyaline distal part, and, in males, whole ventral half hyaline. Distribution and ecology Only known from Lake Edward, found in inshore areas over muddy substrates. Based on its morphology, most probably a piscivorous species., Published as part of Vranken, Nathan, Steenberge, Maarten Van, Heylen, Annelies, Decru, Eva & Snoeks, Jos, 2022, From a pair to a dozen: the piscivorous species of Haplochromis (Cichlidae) from the Lake Edward system, pp. 1-94 in European Journal of Taxonomy 815 on pages 39-43, DOI: 10.5852/ejt.2022.815.1749, http://zenodo.org/record/6484153

Haplochromis simba sp. nov. urn:lsid:zoobank.org:act: 266E8987-E220-4EC4-AD72-0A46BE085EC8 Figs 1–2, 14–16; Table 1 Differential diagnosis Species with a piscivorous morphology; premaxillary pedicel and mental prominences strongly developed; outer oral teeth few and large [UOT 22–31 (median 27)]; dominant males yellow with an orange anterior part of flank. Amongst piscivorous species from the Lake Edward system, H. simba sp. nov. differs from H. latifrons sp. nov., H. mentatus, and H. glaucus sp. nov. by the combination of a narrower interorbital area [IOW 45.5–50.4 (mean 48.1) vs 50.9–63.3 (53.8–60.0) % HW] and a strongly developed vs absent or weakly developed mental prominence. It further differs from H. latifrons sp. nov. by a longer anal fin base [AFB 17.3–19.0 (18.0) vs 14.7–17.3 (15.7) % SL] and absence vs presence of a well-defined mid-lateral band; further from H. mentatus by a smaller number of outer upper jaw teeth [UOT 22–31 (27) vs 28–46 (36)] and dominant males yellow with an orange anterior part of flank vs green-yellow with a red anterior part of flank; further from H. glaucus sp. nov. by a narrower lower pharyngeal element [LPW 83.8–87.9 vs 93.3–95.1% LPL] and dominant males yellow with an orange anterior part of flank vs uniformly lightblue. It differs from H. rex sp. nov. and H. aquila sp. nov. by the combination of a smaller number of scales between first anal-fin spine and upper lateral line (ULL-A 9–11 vs 12–16, rarely 11) and a strongly developed vs absent or weakly developed mental prominence. It further differs from H. rex sp. nov. by a larger number of longitudinal line scales (LongL 32–33 vs 34–38) and dominant males uniformly yellow with an orange anterior part of flank vs cream-coloured with an orange operculum and a light blue snout; further from H. aquila sp. nov. by a smaller eye [ED 26.7–29.5 (28.3) vs 30.0–31.5 (30.6) % HL] and dominant males yellow with an orange anterior part of flank and a yellow head vs uniformly light grey with a black head. It differs from H. kimondo sp. nov., H. falcatus sp. nov., H. curvidens sp. nov., H. pardus sp. nov., H. quasimodo sp. nov., and H. squamipinnis by the combination of large vs small outer oral teeth and a smaller number of outer upper jaw teeth [UOT 22–31 (27) vs 39–79 (45–58)]. It further differs from H. falcatus sp. nov., H. curvidens sp. nov., and H. pardus sp. nov. by a deeper lacrimal [LaD 18.7–20.5 (19.5) vs 16.0–18.8 (16.7–18.0) % HL]; further from H. kimondo sp. nov. and H. quasimodo sp. nov. by a narrower head [HW 39.5–41.5 (40.8) vs 42.0–48.1 (45.1–45.3) % HL]; further from H. squamipinnis by absence vs presence of minute scales on proximal part of dorsal fin. Etymology Specific name from Swahili ‘simba’ for ‘lion’; referring to yellow body, orange cheeks that resemble manes, and predatory morphology. Material examined Holotype UGANDA • ♂, 105.8 mm SL; Lake Edward; 0°24′16.0″ S, 29°46′24.8″ E; 9 Nov. 2016; HIPE1 exped. leg.; bought at Rwenshama landing site; RMCA 2016.035.P.0225. Paratypes DEMOCRATIC REPUBLIC OF THE CONGO • 1 ♂, 98.4 mm SL; “Lac Edouard: Vitshumbi (au Nord)” [Lake Edward: north of Vitshumbi]; 0°40′50.6″ S, 29°23′22.6″ E (inferred); 2 Jul. 1953; KEA exped. leg.; IRSNB 13488. UGANDA – Lake Edward • 2 ♂♂, 1 ♀, 97.9–105.5 mm SL; 0°24′16.0″ S, 29°46′24.8″ E; 9 Nov. 2016; HIPE1 exped. leg.; bought at Rwenshama landing site; RMCA 2016.035.P.0222 to 0224 • 1 ♀, 87.1 mm SL; Rwenshama, rocky shore; 0°24′05.7″ S, 29°46′35.1″ E; 25 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0356 • 3 ♂♂, 90.0– 100.3 mm SL; same collection data as for preceding; RMCA 2017.006.P.0357 to 0359 • 1 ♀, 109.0 mm SL; 0°24′16.0″ S, 29°46′24.8″ E; 24 Jan. 2018; HIPE3 exped. leg.; bought at Rwenshama landing site; RMCA 2018.008.P.0348. Description Based on 10 specimens (87.1–109.0 mm SL); body shallow (Table 1) and oval to slightly pyriform; caudal peduncle shallow (Fig. 14). Head long, narrow, and with a straight to weakly convex dorsal outline and a weak concavity above eye; eye small and high on head; interorbital area narrow; cheek and lacrimal deep. Snout long, rounded in dorsal outline, acute in lateral view, and slopes gently at 30–45°; premaxillary pedicel average in length and strongly prominent. Jaws isognathous to prognathous, long, slim, very narrow, and rounded in dorsal view; gape large and slopes gently at 20–25°; maxilla extends to between verticals through anterior point of orbit and just past this point. Lower jaw relatively shallow and with a straight ventral outline in lateral view, mental prominence strongly developed, and lower jaw side steep with an inclination of 35–40° to horizontal in anterior view. Upper jaw expanded anteriorly and ventrally. Lips and oral mucosa thin. Neurocranium shallow, ethmo-vomerine block decurved, preorbital region very shallow (18–20% NL), orbital region shallow (26–28% NL), and supraoccipital crest shallow and wedge-shaped (Fig. 15b). Outer oral teeth few, unicuspid, and very large. Necks stout, conical, and straight; crowns straight to weakly recurved, and acutely pointed. Dental arcades rounded. Outer teeth widely and irregularly set with neck-distances of 1–4 neck-widths. In upper jaw, 1–2 posteriormost teeth enlarged. Inner teeth small, recurved, unicuspid, and acutely pointed. Tooth bands very slender crescent-shaped with 1–2 rows of inner teeth, and narrow posteriorly until only outer row remains past ⅔ length of tooth band. Inner teeth widely and irregularly set on 1–2 outer tooth neck-widths from outer row; implantation erect in first row and recumbent in subsequent rows; in upper jaw, size decreases slightly buccally and posteriorly. Lower pharyngeal bone long, very narrow, slim, and shallow over whole length (Fig. 16). Pharyngeal teeth relatively large and slender; major cusps acutely pointed; cusp gaps concave; minor cusps and cusp protuberances very small. Teeth in two median longitudinal rows equal in size and form to lateral teeth, 12 in each row. Posterior transverse row with 16–18 teeth, implanted recumbently with a lateral inclination; major cusps weakly recurved, bluntly pointed, and laterally compressed; minor cusps mostly absent. Chest scales small; transition to larger flank scales gradual. Minute scales on proximal half of caudal fin. Caudal fin emarginate; dorsal and anal fins reach to between verticals through two scales anterior to and one scale posterior to caudal-fin base. Pectoral fin reaches to between anal opening and second analfin spine; pelvic fin reaches to between first and second anal-fin spines; first branched pelvic-fin ray elongated. Ceratobranchial gill rakers in outer row of first gill arch short, stout, and simple; posteriormost rakers anvil-shaped to weakly bifid. Epibranchial gill rakers slender and simple. Colouration in life Dominant males: body yellow; operculum, cheek, and anterior part of flank orange; belly and chest white (Fig. 15c). Snout dusky; lower jaw and lips pink; eye with (dark) grey outer ring and silver to golden inner ring. Nostril and lacrimal stripes faint; mental blotch present. pectoral fin yellow; pelvic fin black; dorsal fin hyaline to dusky and with black lappets and a dusky posterior part; anal fin faint crimson and with 2–3 very large orange egg-spots with hyaline rings; caudal fin dusky and with a faint crimson ventral part and a maculated dorsal part. Non-dominant males: anterior part of flank, operculum, and cheek with pink sheen; anal fin with crimson anterior part and hyaline posterior part; caudal fin uniformly dusky. Females and juveniles: body, operculum, and cheek yellow; belly and chest white; lacrimal and lower jaw pink; snout dusky; eye with (dark) grey outer ring and silver to golden inner ring (Fig. 15d). Nostril and interorbital stripes faint; mental blotch present. Pectoral, anal, and caudal fins yellow; anal fin with a white base; caudal fin with a dusky base; dorsal fin hyaline and with black lappets and a dusky distal part. Preserved colouration Body whitish; dorsum and cheek brown; belly, chest, and operculum white; snout dusky (Fig. 15a). Nostril stripe well-defined; lacrimal stripe and mental blotch present; interorbital stripe and nape band faint. Pectoral fin dusky; pelvic fin yellowish in females, black in males; dorsal fin dusky and with black lappets and sooty posterior part; anal fin yellowish and with 1–2 large egg-spots; caudal fin dusky and with a maculated dorsal part. Distribution and ecology Only known from Lake Edward, found in inshore areas over hard substrates. Based on its morphology, most probably a piscivorous species.

Haplochromis mentatus Regan, 1925 Figs 1–2, 8–10; Table 1 Haplochromis mentatus Regan, 1925: 188, pl. 10. Haplochromis mentatus – Greenwood 1973: 204. Harpagochromis mentatus – Greenwood 1980: index. Differential diagnosis Species with a piscivorous morphology; body shallow [BD 29.0–32.3 (mean 31.2) % SL]; snout very acute in dorsal and lateral views; outer oral teeth few and large [UOT 28–46 (median 36)]; vertebrae many (Va+Vc 30–32); dominant males yellow-green with a red anterior part of flank. Amongst piscivorous species from the Lake Edward system, H. mentatus differs from H. latifrons sp. nov. by the combination of a longer dorsal fin base [DFB 50.3–54.2 (52.3) vs 47.2–50.1 (49.0) % SL], a weakly vs strongly prominent premaxillary pedicel, a steeper sloping lower jaw side (30–45° vs 25– 30°), and absence vs presence of a well-defined mid-lateral band. It differs from H. rex sp. nov. and H. aquila sp. nov. by the combination of a longer caudal peduncle [CPL 15.7–17.5 (16.6) vs 13.5–16.2 (14.8–15.0) % SL], a gentler sloping snout (30–35° vs 35–50°), and dominant males yellow-green with a red anterior part of flank vs cream-coloured with an orange operculum, or light grey with a black head, respectively; further from H. rex sp. nov. by a larger number of infraorbital cheek scales [ChSi 3–4 vs 5–6 (rarely 4 or 7)]; further from H. aquila sp. nov. by smaller eyes [ED 25.4–29.9 (27.2) vs 30.0–31.5 (30.6) % HL]. It differs from H. simba sp. nov. by the combination of a broader interorbital area [IOW 51.3–61.0 (55.5) vs 45.5–50.4 (48.1) % HW], a larger number of outer upper jaw teeth [UOT 28–46 (36) vs 22–31 (27)], absent to weakly prominent vs strongly prominent premaxillary pedicel and mentum, and dominant males yellow-green with a red anterior part of flank vs yellow with an orange anterior part of flank. It differs from H. glaucus sp. nov. by the combination of a longer caudal peduncle [CPL 15.7–17.5 (16.6) vs 13.4–16.1 (14.8) % SL], a narrower lower pharyngeal bone [LPW 83.6–85.7 vs 93.3–95.1% LPL], a slightly shorter pre-pectoral distance [PrP 33.1–38.2 (36.0) vs 36.4–39.4 (38.1) % SL], and dominant males yellow-green with a red anterior part of flank vs uniformly blue. It differs from H. kimondo sp. nov. and H. quasimodo sp. nov. by the combination of a narrower head [HW 39.4–42.3 (40.8) vs 42.0–48.1 (45.1–45.3) % HL] and dominant males yellow-green with a red anterior part of flank vs grey dorsally and yellow or blue-black ventrally; further from H. kimondo sp. nov. by a very acute vs blunt snout, and a gentler sloping snout (30–35° vs 40–50°); further from H. quasimodo sp. nov. by a shallower body [BD 27.2–30.1 (28.6) vs 33.5–41.7 (37.4) % SL]. It differs from H. falcatus sp. nov. by the combination of a shorter predorsal distance [PrD 33.3–36.4 (35.3) vs 36.9–41.1 (39.5) % SL], a shorter head [HL 33.4–37.0 (35.1) vs 36.6–39.6 (38.2) % SL], straight to weakly recurved vs strongly recurved outer oral teeth, a steeper lower jaw side (30–45° vs 15–25°), absence vs presence of well-defined mid-lateral and dorsal-lateral bands. It differs from H. curvidens sp. nov. and H. pardus sp. nov. by the combination of a deeper lacrimal [LaD 18.1–20.9 (19.7) vs 16.0–18.3 (16.7–17.3) % HL] and smaller eyes [ED 25.4–29.9 (27.2) vs 29.4–34.1 (30.4–31.9) % HL]; further from H. pardus sp. nov. by dominant males yellow-green with a red anterior part of flank vs speckled to uniformly black. It differs from H. squamipinnis by the combination of large vs small outer oral teeth and a smaller number of outer upper jaw teeth [UOT 28–46 (36) vs 46–71 (58)], a shallower body [BD 29.0–32.3 (31.2) vs 32.4–39.3 (35.7) % SL], absence vs presence of minute scales on proximal part of dorsal fin, and dominant males yellow-green with a red anterior part of flank vs uniformly slate blue. Etymology Specific name not explained in original description, probably derived from the Latin ‘ mentum ’for ‘chin’; probably referring to the protruding lower jaw (i.e., projecting lower jaw sensu Regan 1925). Material examined Holotype UGANDA • ♀, 93.1 mm SL; Lake Edward; 1924; J.C. Phillips leg.; MCZ 31523. Other material UGANDA – Lake Edward • 1 ♂, 94.4 mm SL; ‘Coral Reef’, hard substrate at mouth of Nyamugasani River; 0°10′08.4″ S, 29°49′37.2″ E; 21 Oct. 2016; HIPE1 exped. leg.; RMCA 2016.035.P.0255 • 1 ♂, 2 ♀♀, 111.3–128.1 mm SL; Rwenshama, rocky shore; 0°24′05.7″ S, 29°46′35.1″ E; 25 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0408 to 0410 • 3 ♂♂, 111.5–118.5 mm SL; Kayanja offshore; 0°05′34.8″ S, 29°45′28.8″ E; 30 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0411 to 0413 • 1 ♂, 117.7 mm SL; same collection data as for preceding; RMCA 2017.006.P.0414 • 1 ♀, 1 ♂, 87.2–133.2 mm SL; islands near Katwe; 0°10′04.9″ S, 29°52′27.4″ E; 18 Jan. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0380 to 0381 • 1 ♀, 1 ♂, 87.8–95.5 mm SL; same collection data as for preceding; RMCA 2018.008.P.0383 to 0384 • 2 ♀♀, 108.3–137.2 mm SL; 0°24′16.0″ S, 29°46′24.8″ E; 24 Jan. 2018; HIPE3 exped. leg.; bought at Rwenshama landing site; RMCA 2018.008.P.0387 to 0388 • 1 ♂, 137.1 mm SL; 0°24′16.0″ S, 29°46′24.8″ E; 1 Feb. 2018; HIPE3 exped. leg.; bought at Rwenshama landing site; RMCA 2018.008.P.0389. – Lake George • 1 ♂, 107.9 mm SL; ‘Bivalve Site’, north of Kankurunga Island; 0°00′39.6″ N, 30°08′13.2″ E; 29 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0424 • 1 ♀, 92.1 mm SL; Kashaka bay, north of inlet; 0°04′52.2″ S, 30°10′47.3″ E; 2 Feb. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0394 • 2 ♀♀, 94.0, 120.0 mm SL; same collection data as for preceding; RMCA 2018.008.P.0395 to 0396. Description Based on 20 specimens (87.2–137.2 mm SL); body shallow (Table 1) and oval; caudal peduncle long (Fig. 8). Head very narrow and with a straight dorsal outline and a slightly convex nape; cheek average in depth; lacrimal deep; eye small; interorbital area narrow in comparison to generalised H. elegans (but relatively broad for a piscivorous species). Snout long, very acute in dorsal and lateral views, and slopes very gently at 30–35°; premaxillary pedicel long and weakly prominent. Jaws long, very narrow, acute in dorsal view, relatively slim, and isognathous; gape large and slopes gently at 20–30°; posterior tip of maxilla reaches a vertical just past anterior margin of orbit. Lower jaw slim and with a straight to weakly concave ventral outline in lateral view, mental prominence absent or weakly developed, and lower jaw side steep with an inclination of 30–45° to horizontal in anterior view. Upper jaw expanded anteriorly and weakly ventrally. Lips and oral mucosa large. Neurocranium shallow, ethmo-vomerine block decurved, preorbital region very shallow (18–22% NL), orbital region shallow (28–31% NL), and supraoccipital crest shallow and wedge-shaped (Fig. 9b). Outer oral teeth few, unicuspid, and large. Necks stout, conical, and straight; crowns slightly recurved and acutely pointed. Dental arcades acute. Outer teeth widely and irregularly set with neck-distances of ½–2 neck-widths. In upper jaw, 1–3 posteriormost teeth slightly enlarged. Inner teeth small, strongly recurved, unicuspid, and acutely pointed. Tooth bands very slender crescent-shaped with 1–2 rows of inner teeth, and narrow posteriorly until only outer row remains past ⅔ length of tooth band in upper jaw, past ½ length of tooth band in lower jaw. Inner teeth closely and regularly set on 1–2 outer neck-widths from outer row; implantation recumbent; size uniform. Lower pharyngeal bone long, narrow, slim, and shallow over whole length (Fig. 10). Pharyngeal teeth relatively large and slender; major cusps acutely pointed; cusp gaps concave; minor cusps and cusp protuberances mostly absent. Teeth in two median longitudinal rows equal in size and form to lateral teeth, 11 in each row. Posterior transverse row with 14–21 teeth, implanted recumbently with a lateral inclination; major cusps nearly straight, bluntly pointed, and laterally compressed; minor cusps mostly absent. Chest scales small; transition to larger flank scales gradual. In some specimens, basal parts of membranes between anal-fin spines covered by few (1–3) minute, ellipsoid scales (nearly invisible to naked eye); remaining part of anal and dorsal fins scale-less. Minute scales on proximal half of caudal fin. Caudal fin emarginate; dorsal and anal fins reach to between vertical through caudal-fin base and two scales anterior to this vertical. Pectoral and pelvic fins reach to between genital opening and first anal-fin spine; pelvic fin reaches to third anal-fin spine in males; first branched pelvic-fin ray slightly elongated in all specimens. Ceratobranchial gill rakers in outer row of first gill arch short, stout, and simple; posteriormost rakers mostly anvil-shaped or weakly bifid. Epibranchial rakers slender and simple. Colouration in life Dominant males: body, cheek, and lower jaw yellow-green with blue sheen; flank, dorsal part of head, and operculum bright red; belly and chest speckled black (Fig. 9c). Snout and lips dusky; branchiostegal membrane black; eye with (dark) grey outer ring and yellow inner ring. Flank with very faint midlateral, dorsal-lateral, and 5–7 vertical stripes; nostril stripe faint. Dorsal and anal fins dusky and with black lappets; anal fin with a crimson distal part and 2 small orange egg spots with dusky rings. Caudal fin dusky with a crimson distal part. Pectoral fin dusky yellow and pelvic fin black. Females and juveniles: dorsal parts of body and operculum green-yellow; gradual transition to white ventral parts of body and operculum, cheek, and lower jaw; flank with a blue sheen (Fig. 9d). Lacrimals, snout, and lips dusky green; lacrimals with a blue sheen; eye with (dark) grey outer ring and yellow inner ring. Nostril stripe faint. Dorsal fin dusky and with black lappets; anal fin white-yellow and with 1–2 small spots resembling egg-spots; caudal fin dusky. Pectoral fin yellowish; pelvic fin white-yellow. Preserved colouration In dominant males, body uniformly brown, pectoral fin dusky, pelvic fin black, and anal fin dusky and with 1–2 small egg-spots (Fig. 9a). In females, dorsal part of body yellowish, gradual transition to white ventral part of body, cheek light yellow, pectoral fin hyaline, pelvic fin yellowish, and anal fin with a white base and a dusky distal part. In all specimens, snout dusky and nostril, interorbital, and lacrimal stripes faint. Dorsal fin dusky and with black lappets; caudal fin dusky. Distribution and ecology Endemic to Lake Edward system, found in offshore areas, mostly in shallow waters. Based on its morphology, most probably a piscivorous species. Systematic comment Both Trewavas (1933) and Greenwood (1973) noted that the holotype of H. mentatus strongly resembled H. squamipinnis. Greenwood recognised that this specimen deviated from H. squamipinnis in that it lacked minute scales on the basal parts of the dorsal- and anal-fin membranes, but he was uncertain about the relevance of this difference. We found this trait to be present in all H. squamipinnis specimens. We discovered that abdominal vertebrae 6–9 of the holotype of H. mentatus are broken and the vertebral column kinked (Fig. 9b). This resulted in the holotype of H. mentatus having an aberrant rhomboidshaped instead of an oval body as in other specimens of its species. Because of this deformation, the specimen indeed resembles H. squamipinnis in overall habitus., Published as part of Vranken, Nathan, Steenberge, Maarten Van, Heylen, Annelies, Decru, Eva & Snoeks, Jos, 2022, From a pair to a dozen: the piscivorous species of Haplochromis (Cichlidae) from the Lake Edward system, pp. 1-94 in European Journal of Taxonomy 815 on pages 18-24, DOI: 10.5852/ejt.2022.815.1749, http://zenodo.org/record/6484153, {"references":["Regan C. T. 1925. Three new cichlid fishes of the genus Haplochromis from Lake Edward, Central Africa. Occasional Papers of the Boston Society of Natural History 5: 187 - 188.","Greenwood P. H. 1973. A revision of the Haplochromis and related species (Pisces: Cichlidae) from Lake George, Uganda. Bulletin of the British Museum (Natural History) Zoology 25: 139 - 242.","Greenwood P. H. 1980. Towards a phyletic classification of the ' genus' Haplochromis (Pisces, Cichlidae) and related taxa. Part II. Bulletin of the British Museum (Natural History) Zoology 39: 1 - 101. https: // doi. org / 10.5962 / bhl. part. 13268","Trewavas E. 1933. Scientific results of the Cambridge expedition to the East African Lakes, 1930 - 1. The cichlid fishes. Zoological Journal of the Linnean Society 38: 309 - 341. https: // doi. org / 10.1111 / j. 1096 - 3642.1933. tb 00062. x"]}

Haplochromis glaucus sp. nov. urn:lsid:zoobank.org:act: E5948349-E3F9-49A1-81CD-1C7D332E2F00 Figs 1–2, 17–19; Table 1 Differential diagnosis Species with a piscivorous morphology; snout acute in lateral view; outer oral teeth few and large [UOT 25–47 (median 30)]; males grey with light blue flank and a dusky to black head; female colour pattern similar to males. Amongst piscivorous species from the Lake Edward system, H. glaucus sp. nov. differs from H. latifrons sp. nov. and H. mentatus by a shorter caudal peduncle [CPL 13.4–16.1 (mean 14.8) vs 15.7–18.0 (16.6– 17.0) % SL]; further from H. latifrons sp. nov. by a longer anal fin base [AFB 17.3–20.3 (18.6) vs 14.7– 17.3 (15.7) % SL] and absence vs presence of a well-defined mid-lateral band; further from H. mentatus by a broader lower pharyngeal bone [LPW 93.3–95.1 vs 83.6–85.7% LPL], a slightly longer pre-pectoral distance [PrP 36.4–39.4 (38.1) vs 33.1–38.2 (36.0) % SL], and dominant males uniformly light blue vs yellow-green with a red anterior part of flank. It differs from H. rex sp. nov. and H. simba sp. nov. by a broader interorbital area [IOW 50.9–57.1 (53.8) vs 44.9–52.7 (48.1–48.9) % HW]; further from H. rex sp. nov. by a gentler sloping snout (30–40° vs 40–50°), acute vs rounded oral jaws in dorsal view, and dominant males light blue with a blackish operculum and a dusky snout vs cream-coloured with an orange operculum and a light blue snout; further from H. simba sp. nov. by a broader lower pharyngeal bone [LPW 93.3–95.1 vs 83.8–87.9% LPL], absent or weakly developed vs strongly developed mental prominence, and dominant males uniformly light-blue vs yellow with an orange anterior part of flank. It differs from H. aquila sp. nov. by the combination of a smaller eye [ED 23.2–28.7 (26.8) vs 30.0–31.5 (30.6) % HL], a narrower head [HW 38.9–40.9 (39.7) vs 40.1–43.7 (42.0) % HL], and dominant males light blue with crimson anal and caudal fins vs light grey with bright red anal and caudal fins. It differs from H. kimondo sp. nov., H. curvidens sp. nov., and H. quasimodo sp. nov. by the combination of a narrower head [HW 38.9–40.9 (39.7) vs 42.0–48.1 (43.4–45.3) % HL], large vs small outer oral teeth, and a smaller number of outer upper jaw teeth [UOT 25–47 (30) vs 43–71 (49–58)]; further from H. kimondo sp. nov. and H. quasimodo sp. nov. by dominant males light blue vs grey dorsally and yellow or blue-black ventrally. It differs from H. falcatus sp. nov. by the combination of a shorter pre-dorsal distance [PrD 35.4–37.0 (36.1) vs 36.9–41.1 (39.5) % SL], a steeper lower jaw side (35–45° vs 15–25°), weakly recurved vs strongly recurved outer oral teeth, and dominant males uniformly light blue vs olive-green with an orange-red anterior part of flank. It differs from H. pardus sp. nov. by the combination of deeper lacrimal [LaD 18.0–22.7 (19.8) vs 16.0–18.3 (17.3) % HL], a broader interorbital area [IOW 50.9–57.1 (53.8) vs 39.3–48.4 (44.6) % HW], and dominant males light blue vs speckled to uniformly black. It further differs from H. squamipinnis by the combination of a gentler gape inclination (20–30° vs 30–45°), absence vs presence of minute scales on proximal parts of dorsal and anal fins, and dominant males light blue vs slate blue. Etymology Specific name from the Latin ‘ glaucus ’ for ‘greyish blue’; referring to grey and light-blue colour pattern of all adult specimens. Material examined Holotype UGANDA • ♂, 106.2 mm SL; Lake Edward, Kayanja offshore; 0°05′34.8″ S, 29°45′28.8″ E; 21 Mar. 2019; HIPE4 exped. leg.; RMCA 2019.002.P.0016. Paratypes DEMOCRATIC REPUBLIC OF THE CONGO • 1 ♂, 158.3 mm SL; “Lac Edouard: au large de la riv. Kigera” [Lake Edward: offshore Kigera river]; 0°29′42″ S, 29°38′14″ E (inferred); 25 May 1953; KEA exped. leg.; IRSNB 13477 • 1 ♂, 150.9 mm SL; “Lac Edouard: 2–3 km ± 500 m au large à l’Ouest de Kiavinionge” [Lake Edward: 2–3 km ± 500 m offshore west of Kiavinionge]; 0°11′39″ S, 29°32′31″ E (inferred); 1 Jun. 1953; KEA exped. leg.; IRSNB 13480. UGANDA – Lake Edward • 1 ♀, 1 ♂, 90.7–93.6 mm SL; Kagoro fishing ground; 0°12′50.1″ S, 29°49′19.7″E; 4 Feb. 2018; HIPE3 exped. leg.; open water; RMCA 2018.008.P.0365 to 0366 • 1 ♀, 92.6 mm SL; same collection data as for preceding; RMCA 2018.008.P.0367 • 1 ♀, 1 ♂, 102.1– 104.7 mm SL; Kayanja offshore; 0°05′34.8″ S, 29°45′28.8″ E; 21 Mar. 2019; HIPE4 exped. leg.; RMCA 2019.002.P.0017 to 0018 • 1 ♂, 2 ♀♀, 107.7–122.3 mm SL; same collection data as for preceding; RMCA 2019.002.P.0019 to 0021. Description Based on 11 specimens (90.7–158.3 mm SL); body shallow (Table 1) and oval (Fig. 17). Head long, very narrow, and with a straight dorsal outline; eye small; interorbital area narrow; lacrimal and cheek deep. Snout long, rounded in dorsal view, very narrow, acute, and slopes gently at 30–40°; premaxillary pedicel long and weakly prominent. Jaws long, relatively stout, narrow, isognathous, and acute in dorsal view; gape large and slopes gently at 20–30°; maxilla extends (almost) to vertical through anterior margin of orbit. Lower jaw relatively stout and with a straight to slightly convex ventral outline in lateral view, mental prominence absent or weakly developed, and lower jaw side steep with an inclination of 35–45° to horizontal in anterior view. Upper jaw expanded anteriorly and weakly ventrally. Lips and oral mucosa large. Neurocranium shallow, ethmo-vomerine block decurved, preorbital region shallow (20–23% NL), orbital region shallow (28–31% NL), and supraoccipital crest shallow and wedge-shaped (Fig. 18b). Outer oral teeth few, unicuspid, and very large. Necks conical, stout, and straight; crowns recurved and acutely pointed. Dental arcades rounded. Outer teeth widely and irregularly set with neck-distances of 1–4 neck-widths. In upper jaw, 1–3 posteriormost teeth slightly enlarged. Inner teeth small, strongly recurved, unicuspid, and acutely pointed. Tooth bands very slender crescent-shaped with 1–3 rows of inner teeth, and narrow posteriorly until only outer row remains past ¾ length of tooth band. Inner teeth closely and regularly set on 1–2 outer neck-widths from outer row; implantation recumbent; size decreases slightly buccally and posteriorly. Lower pharyngeal bone average in length, triangular, slim, and shallow over whole length (Fig. 19). Pharyngeal teeth relatively large and slender; major cusps acutely pointed; cusp gaps straight; minor cusps and cusp protuberances mostly absent. Teeth in two median longitudinal rows equal in size and form to lateral teeth, 10–12 in each row. Posterior transverse row with 15–16 teeth, implanted recumbently with a lateral inclination; major cusps recurved, bluntly pointed, and laterally compressed; minor cusps small. Chest scales small; transition to larger flank scales gradual. Minute scales on proximal half of caudal fin. Caudal fin emarginate; dorsal and anal fins reach to between verticals through one scale anterior to and two scales posterior to caudal-fin base. Pectoral and pelvic fins reach to between anal opening and first anal-fin spine; first branched pelvic-fin ray not elongated. Ceratobranchial gill rakers in outer row of first gill arch short, stout, and simple; posteriormost rakers anvil-shaped to rarely weakly trifid. Epibranchial gill rakers slender and simple. Colouration in life Dominant males: flank and caudal peduncle light-blue; dorsum grey with a yellow sheen; belly, chest, and operculum blackish with a yellow sheen; flank with 5–7 faint vertical stripes (Fig. 18c). Preoperculum black; cheek black with blue sheen; lacrimal and snout dusky; lower jaw blackish; branchiostegal membrane black; eye with dark grey outer ring and silver inner ring. Nostril, interorbital, supraorbital, and lacrimal stripes well-defined; nape band, vertical preopercular stripe, and mental blotch present; operculum with black posterior border. Pectoral and dorsal fins hyaline; dorsal fin with black lappets, dusky base, and dusky and maculated crimson posterior part. Anal and caudal fins crimson; anal fin with dusky posterior part and 1–2 large orange egg-spots with dusky rings; caudal fin with dusky base, hyaline posterior part, and crimson maculae. Females: strikingly similar to dominant males. Body, dorsum, and caudal peduncle grey with a yellow sheen; flanks light blue and with 5–7 faint vertical stripes; belly and chest blackish with a yellow sheen (Fig. 18d). Operculum, preoperculum, cheek, lacrimal, and snout grey; operculum with blue sheen; cheek with black sheen; lower jaw blackish; branchiostegal membrane black; eye with dark grey outer ring and silver inner ring. Nostril, interorbital, supraorbital, and lacrimal stripes well-defined; nape band, vertical preopercular stripe, and mental blotch faint; operculum with faint black posterior border. Pectoral and dorsal fins hyaline; dorsal fin with black lappets, dusky base, and dusky posterior part. Anal and caudal fins yellow with a red sheen; anal fin with hyaline posterior part and 1–2 spots resembling egg-spots; caudal fin with dusky base, hyaline posterior part, and dusky maculae. Juveniles: dorsum greyish, belly, chest, operculum, and cheek white; transition gradual. Nostril, interorbital, supraorbital, and lacrimal stripes, nape band, and mental blotch faint. Pectoral, pelvic, dorsal, and anal fins hyaline; dorsal fin with black lappets and dusky base and posterior part; anal fin with a yellow sheen and 1–2 spots resembling egg-spots. Caudal fin dusky and with black maculae and a yellow ventral part. Preserved colouration Dorsum brown; belly and chest speckled black; flank dark yellowish and with 6–8 faint vertical stripes. Cheek yellow and speckled black; snout dusky; lower jaw and pre- and subopercula black (Fig. 18a). Nostril, interorbital, and vertical preopercular stripes well-defined; lacrimal stripe broad and welldefined; supraorbital stripe and nape band faint. Pectoral and anal fins hyaline; anal fin with black lappets, a dusky posterior margin, and 1–2 large egg-spots; pelvic fin black; dorsal fin dusky and with black lappets and a maculated posterior part; caudal fin dorsally dusky, ventrally hyaline, and with a dusky base. Distribution and ecology Only known from Lake Edward, found over sandy substrates. Based on its morphology, most probably a piscivorous species.

Haplochromis kimondo sp. nov. urn:lsid:zoobank.org:act: 24F01142-A596-458D-9887-D5BC8E68A275 Figs 1–2, 23–25; Table 1 Differential diagnosis Species with a piscivorous morphology; head blunt and with convex dorsal outline; cheek deep [ChD 27.1–35.2 (mean 30.9) % HL]; outer oral teeth many and small [UOT 43–70 (median 56)]; dominant males grey dorsally and yellow ventrally. Amongst piscivorous species from the Lake Edward system, H. kimondo sp. nov. differs from H. latifrons sp. nov., H. mentatus, H. rex sp. nov., H. simba sp. nov., H. glaucus sp. nov., and H. aquila sp. nov. by the combination of a broader head [HW 42.9–48.0 (45.1) vs 36.8–43.7 (39.2–42.0) % HL], small vs large outer oral teeth, and a larger number of outer upper jaw teeth [UOT 43–70 (56) vs 22–47 (27–36)]. It differs from H. falcatus sp. nov., H. curvidens sp. nov., H. pardus sp. nov., H. quasimodo sp. nov., and H. squamipinnis by the combination of body pyriform vs oval to rhomboid, and snout blunt vs (very) acute in dorsal view. It further differs from H. falcatus sp. nov. and H. curvidens sp. nov. by the combination of deeper cheeks [ChD 27.1–35.2 (30.9) vs 22.4–28.0 (23.2–26.0) % HL], broader jaws [LJW 44.7–53.3 (49.3) vs 38.5–45.5 (40.8–42.5) % LJL], and a slightly broader head [HW 42.9–48.0 (45.1) vs 39.9–44.4 (42.6– 43.4) % HL]; further from H. falcatus sp. nov. by dominant males grey dorsally and yellow ventrally vs olive-green with an orange-red anterior part of flank; further from H. curvidens sp. nov. by presence vs absence of a well-defined mid-lateral band in all specimens. It further differs from H. pardus sp. nov., H. quasimodo sp. nov., and H. squamipinnis by a steeper sloping snout (40–50° vs 30–40°) and dominant males grey dorsally and yellow ventrally vs speckled to uniformly black, light grey dorsally and blue-black ventrally, or slate blue, respectively; further from H. pardus sp. nov. and H. quasimodo sp. nov. by a broader interorbital area [IOW 49.2–58.5 (52.8) vs 39.3–48.7 (43.9–44.6) % HW]; further from H. squamipinnis by absence vs presence of minute scales on proximal part of dorsal and anal fins. Etymology Specific name from the Swahili ‘kimondo’ for ‘meteor’; referring to blunt head, pyriform body with mid-lateral band, and yellow colouration of ventral part of body. Material examined Holotype UGANDA • ♂, 152.4 mm SL; Lake Edward, Kayanja offshore; 0°05′34.8″ S, 29°45′28.8″ E; 21 Mar. 2019; HIPE4 exped. leg.; RMCA 2019.002.P.0015. Paratypes DEMOCRATIC REPUBLIC OF THE CONGO • 1 ♀, 158.2 mm SL; “Lac Edouard: au large de la riv. Kigera” [Lake Edward: offshore of the Kigera River]; 0°29′42″ S, 29°38′14″ E (inferred); 25 May 1953; KEA exped. leg.; IRSBN 13477 • 1 ♂, 1 ♀, 149.9, 171.3 mm SL; “Lac Edouard: 2–3 km à l’Ouest de Kiavinionge” [Lake Edward: 2–3 km west of Kiavinionge]; 0°11′39″ S, 29°32′31″ E (inferred); 1 Jun. 1953; KEA exped. leg.; IRSBN 13482. UGANDA – Lake Edward • 1 ♀, 146.4 mm SL; Rwenshama; 0°24′05.7″ S, 29°46′35.1″ E; 8 Nov. 2016; HIPE1 exped. leg.; rocky shore; RMCA 2016.035.P.0226 • 3 ♀♀, 103.8–124.4 mm SL; Rwenshama,; 0°24′05.7″ S, 29°46′35.1″ E; 26 Mar. 2017; HIPE2 exped. leg.; rocky shore; RMCA 2017.006.P.0360 to 0362 • 1 ♀, 1 ♂, 106.4–131.1 mm SL; Kayanja offshore; 0°05′34.8″ S, 29°45′28.8″ E; 31 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0363 to 0364 • 2 ♂♂, 1 ♀, 81.6–118.4 mm SL; same collection data as for preceding; RMCA 2017.006.P.0365 to 0367 • 1 ♂, 2 ♀♀, 89.6–115.9 mm SL; Kayanja offshore; 0°05′31.2″ S, 29°45′30.3″ E; 20 Jan. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0358 to 0360 • 2 ♂♂, 2 ♀♀, 128.1–146.8 mm SL; 0°24′16.0″ S, 29°46′24.8″ E; 24 Jan. 2018; HIPE3 exped. leg.; bought at Rwenshama landing site; RMCA 2018.008.P.0361 to 0364 • 1 ♀, 142.9 mm SL; Rwenshama; 0°24′05.7″ S, 29°46′35.1″ E; 24 Jan. 2018; HIPE3 exped. leg.; rocky shore; RMCA 2018.008.P.0357. Description Based on 21 specimens (81.6–171.3 mm SL); body shallow (Table 1) and pyriform (Fig. 23). Head long, stout, average in width in comparison to generalised H. elegans (but broad for a piscivorous species), and with a convex dorsal outline; eye small; interorbital area average in width; cheek and lacrimal deep. Snout long, blunt, and slopes at 40–50°; premaxillary pedicel long and slightly prominent. Jaws isognathous to slightly prognathous, long, relatively stout, rounded in dorsal view, and narrow; gape large and slopes at 25–30°; maxilla extends to vertical through pupil. Lower jaw with a straight ventral outline in lateral view, mental prominence absent or weakly developed, and lower jaw side nearly flat with an inclination of 15–30° to horizontal in anterior view. Upper jaw weakly expanded anteriorly. Lips and oral mucosa large. Neurocranium average in depth, ethmo-vomerine block decurved, preorbital region average in depth (23–25% NL), orbital region average in depth (31–33% NL), and supraoccipital crest shallow and wedge-shaped (Fig. 24b). Outer oral teeth numerous and very small. Necks stout, conical, and straight; crowns recurved and unicuspid in large specimens (> 85 mm SL), bicuspid in small specimens ( 100 mm SL), weakly tricuspid in small specimens (, Published as part of Vranken, Nathan, Steenberge, Maarten Van, Heylen, Annelies, Decru, Eva & Snoeks, Jos, 2022, From a pair to a dozen: the piscivorous species of Haplochromis (Cichlidae) from the Lake Edward system, pp. 1-94 in European Journal of Taxonomy 815 on pages 44-48, DOI: 10.5852/ejt.2022.815.1749, http://zenodo.org/record/6484153

Haplochromis pardus sp. nov. urn:lsid:zoobank.org:act: 8DDACAC6-C993-4750-BE1B-1BF4C0312DB7 Figs 1–2, 32–34; Table 1 Differential diagnosis Species with a piscivorous morphology; adult size small (max. known size 96 mm SL); outer oral teeth many and small [UOT 39–56 (median 58)]; dominant males speckled to uniformly black. Amongst piscivorous species from the Lake Edward system, H. pardus sp. nov. differs from all by the combination of a smaller adult size (max. 96 vs 109–211 mm SL) and colour pattern of small specimens ( It further differs from H. latifrons sp. nov., H. mentatus, H. glaucus sp. nov., H. kimondo sp. nov., and H. squamipinnis by the combination of a shallower cheek [ChD 20.8–24.4 (mean 22.5) vs 23.8–36.0 (26.4– 30.9) % HL] and a narrower interorbital area [IOW 39.3–48.4 (44.6) vs 48.6–63.3 (51.9–60.0) % HW]. It further differs from H. rex sp. nov., H. simba sp. nov., and H. aquila sp. nov. by the combination of a shallower cheek [ChD 20.8–24.4 (22.5) vs 26.8–33.5 (28.3–31.1) % HL], smaller outer oral teeth, and a larger number of outer upper jaw teeth [UOT 39–51 (45) vs 22–37 (27–31)]. It further differs from H. falcatus sp. nov. by the combination of by weakly recurved vs strongly recurved outer jaw teeth, a shallower cheek [ChD 20.8–24.4 (22.5) vs 25.1–28.0 (exceptionally 23.3 in one specimen) (mean 26.0) % HL], and a shorter pre-dorsal distance [PrD 34.1–37.8 (36.0) vs 38.2–41.1 (exceptionally 36.9 in one specimen) (mean 39.5) % SL]. It further differs from H. curvidens sp. nov. by the combination of a longer anal-fin base [AFB 19.2–22.2 (20.5) vs 17.9–18.6 (18.3) % SL] and a slightly narrower interorbital area [IOW 39.3–48.4 (44.6) vs 46.4–52.5 (49.1) % HW]. It further differs from H. quasimodo sp. nov. by the combination of a shallower cheek [ChD 20.8–24.4 (22.5) vs 23.7–32.9 (27.5) % HL] and a smaller number of caudal peduncle scales (CPS 16, rarely 17 vs 17–20, rarely 16). Etymology Specific name from the Latin ‘ pardus ’ for ‘leopard’; referring to nearly uniform black to yellow-pink flanks with clear black blotches, i.e., interrupted horizontal and vertical stripes. Material examined Holotype UGANDA • ♂, 89.2 mm SL; Lake Edward, Mukutu Kihinga, rocky offshore of Mweya; 0°11′31.2″ S, 29°52′26.4″ E; 23 Oct. 2016; HIPE1 exped. leg.; RMCA 2016.035.P.0202. Paratypes UGANDA – Lake Edward • 1 ♀, 84.9 mm SL; Mukutu Kihinga, rocky offshore of Mweya; 0°11′31.2″ S, 29°52′26.4″ E; 23 Oct. 2016; HIPE1 exped. leg.; RMCA 2016.035.P.0203 • 2 ♂♂, 2 ♀♀, 83.6–96.1 mm SL; mouth of Kazinga Channel; 0°12′32.4″ S, 29°53′06.0″ E; 24 Oct. 2016; HIPE1 exped. leg.; RMCA 2016.035.P.0204 to 0207 • 1 ♀, 1 ♂, 75.9, 84.7 mm SL; Rwenshama rocky shore; 0°24′05.7″ S, 29°46′35.1″ E; 26 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0342 to 0343 • 3 ♀♀, 67.4– 70.5 mm SL; Kayanja offshore; 0°05′34.8″ S, 29°45′28.8″ E; 30 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0346 to 0348 • 1 ♀, 1 ♂, 67.7, 78.1 mm SL; Kayanja offshore; 0°05′34.8″ S, 29°45′28.8″ E; 30 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0344 to 0345 • 1 ♀, 92.4 mm SL; islands near Katwe; 0°10′04.9″ S, 29°52′27.4″ E; 19 Jan. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0331 • 1 ♀, 1 ♂, 72.3, 81.7 mm SL; Rwenshama rocky shore; 0°24′05.7″ S, 29°46′35.1″ E; 24 Jan. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0332 to 0333 • 2 ♀♀, 2 ♂♂, 71.0– 84.9 mm SL; Rwenshama rocky shore; 0°24′05.7″ S, 29°46′35.1″ E; 26 Mar. 2017; HIPE2 exped. leg.; IRSNB 920 to 922. Description Based on 20 specimens (67.4–96.1 mm SL); body shallow (Table 1) and oval to slightly rhomboid (Fig. 32). Head narrow, shallow, and with a straight dorsal outline with a concavity above eye; eye average in size in comparison to generalised H. elegans (but large for a piscivorous species); interorbital area very narrow; cheek shallow; lacrimal average in depth. Snout average in length, very acute, and slopes gently at 30–40°; premaxillary pedicel long and prominent. Jaws iso- to slightly prognathous, average in length, narrow, and rounded in dorsal view; gape large and slopes gently at 15–25°; maxilla extends to between verticals through anterior margins of orbit and pupil. Lower jaw shallow and with straight ventral outline in lateral view, mental prominence absent, and lower jaw side nearly flat with an inclination of 15–30° to horizontal in anterior view. Upper jaw weakly expanded anteriorly and ventrally. Lips and oral mucosa large. Neurocranium average in depth, ethmo-vomerine block decurved, preorbital region very shallow (18–22% NL), orbital region average in depth (30–33% NL), and supraoccipital crest shallow and wedge-shaped (Fig. 33b). Outer oral teeth numerous, small, and weakly embedded in oral mucosa. Necks stout, cylindrical, and straight; crowns weakly recurved, unicuspid in large specimens ( 70 mm), and acutely pointed in all specimens. Dental arcades rounded and with anterior half weakly expanded laterally. Outer teeth closely and regularly set with neck-distances of ½ neck-width; lateral outer teeth implanted slightly labially. In upper jaw, 1–3 posteriormost teeth enlarged. Inner teeth small, recurved, unicuspid in large specimens (> 80 mm), tricuspid in small specimens ( Lower pharyngeal bone average in length, narrow, slim, and shallow over entire length (Fig. 34). Pharyngeal teeth small and slender; major cusps acutely pointed; cusp gaps nearly straight; minor cusps and cusp protuberances small. Teeth in two median longitudinal rows equal in size and form to lateral teeth, 11–12 in each row. Posterior transverse row with 22 teeth, implanted erectly with a slight lateral inclination; major cusps weakly recurved, bluntly pointed, and laterally compressed; minor cusps mostly present. Chest scales small; transition to larger flank scales gradual. Minute scales on proximal half of caudal fin. Caudal fin emarginate; dorsal and anal fins reach to between verticals through one scale anterior and one scale posterior to caudal-fin base. Pectoral and pelvic fins reach to genital opening; pelvic fin reaches to first anal-fin spine in males; first branched pelvic-fin ray not elongated. Ceratobranchial gill rakers in outer row of first gill arch short, stout, and simple; posteriormost rakers mostly weakly bifid. Epibranchial gill rakers relatively slender and simple. Colouration in life Dominant males: body speckled to uniformly black with a blue sheen; dorsum and dorsal part of head pink to speckled black; chest white; belly, caudal peduncle, operculum, lower jaw, and lips black; cheek, lacrimal, and snout speckled to uniformly black, cheek with yellow sheen; eye with grey to dark outer ring and silver to dark inner ring (Fig. 33c). Flank with dorsal-lateral and mid-lateral bands and 5–6 vertical stripes, all black, broad, well-defined, but interrupted, hereby body seemingly blotched. Nostril, interorbital, supraorbital, lacrimal, vertical preopercular stripes, nape band, mental blotch, and black posterior margin of operculum well-defined. Pectoral, dorsal, and anal fins dusky; lappets, bases, and posterior parts of anal and dorsal fins black; anal fin with faint crimson flush and 1–3 small orange eggspots with dusky rings. Pelvic and caudal fins black; caudal fin with dusky distal part and a faint crimson flush. Females and juveniles: body yellowish with a dusky sheen; dorsum and dorsal part of head pink; belly, chest, operculum, and cheek, white; lacrimal speckled black; snout dusky to black; eye with grey to dark outer ring and silver to dark inner ring (Fig. 33d). Flank with dorsal-lateral and mid-lateral bands and 5–6 vertical stripes, all faint and interrupted. Nostril and interorbital stripes faint; supraorbital, lacrimal, vertical preopercular stripes, mental blotch, and nape band well-defined. Pectoral, pelvic, and dorsal fins dusky; anal and caudal fins dusky with yellow sheen; dorsal and anal fins with black lappets and posterodistal part; caudal fin with black base and 1–2 small spots resembling egg-spots. Preserved colouration In all specimens, dorsal part of body dark brown to speckled black; ventral part of body dark brown to uniformly black; belly black; chest whitish (Fig. 33a). Flank with faint to well-defined, broad, but interrupted mid-lateral, dorsal-lateral, and dorsal-medial bands and 5–6 vertical stripes. Snout dusky; lips dusky with well-defined black spots; cheek yellowish to black; operculum black. Nostril, interorbital, supraorbital, lacrimal, vertical preopercular stripes, nape band, mental blotch, and black posterior margin of operculum well-defined. Pectoral, dorsal, and anal fins dusky; lappets, bases, and posterior parts of anal and dorsal fins black; anal fin with 1–3 egg-spots. Pelvic and caudal fins dusky to black; caudal fin with blackish distal part. Distribution and ecology Only known from Lake Edward; found in inshore areas. Based on its morphology, most probably a piscivorous species.

Key to the piscivorous species of Haplochromis from the Lake Edward System This identification key is intended as a practical guide and first step towards the identification of the piscivorous species from the Lake Edward system; in case of doubt, the differential diagnoses in the species’ descriptions should be consulted. When identifying live dominant males, the colour patterns are highly diagnostic and should be checked. A simple dichotomous key cannot be compiled as overlap in character states and in the range of values is omnipresent in species of Haplochromis, especially when they have a similar ecology (Greenwood 1973; Snoeks 1994). However, the key allows for the identification of most specimens. Within the key, terminology is used in reference to species with a piscivorous morphology (instead of the generalised Haplochromis morphology as in the rest of the manuscript). 1. Adult specimens H. pardus sp. nov. – Adult specimens> 100 mm SL; specimens H. latifrons sp. nov. – Body deep [BD 28.4–41.7 (30.8–37.4) % SL]; anal-fin base long [AFB 16.7–21.9 (18.0–19.9) % SL]; interorbital area narrow [IOW 39.3–61.0 (43.9–55.5) % HW]; 8–11 anal fin branched rays... 3 3. Body deep [BD 32.4–39.3 (35.7) % SL]; interorbital area broad [IOW 48.6–55.6 (51.9) % HW]; lower jaw long [LJL 47.8–58.6 (52.7) % HL]; gape steep (30–45°); rows of minute scales on basal part of membranes of both dorsal and anal fins (nearly invisible to naked eye) (Fig. 38); dominant males slate blue (Fig. 39c)........................................................................................ H. squamipinnis – Body deep [BD 33.5–41.7 (37.4) % SL]; interorbital area narrow [IOW 40.5–48.7 (43.9) % HW]; lower jaw short [LJL 44.2–49.6 (47.1) % HL]; gape gentle (20–35°); rows of minute scales on basal part of membrane of anal fin in some specimens, rarely few isolated scales on dorsal fin (nearly invisible to naked eye) (Fig. 35); dominant males light grey dorsally and blue-black ventrally (Fig. 36c).......................................................................................................... H. quasimodo sp. nov. – Body shallow [BD 28.4–36.0 (30.8–33.2) % SL]; interorbital area broad [IOW 39.3–61.0 (44.6– 55.5) % HW]; lower jaw short [LJL 42.4–53.1 (44.7–49.5) % HL]; gape gentle (15–35°); minute scales on dorsal or anal fins absent.................................................................................................... 4 4. Head broad [HW 40.1–43.7 (42.0) % HL]; eye large [ED 30.0–31.5 (30.6) % HL]; 25–37 (median 31) outer upper jaw teeth; dominant males light grey with black head and bright red anal fin (Fig. 21c).................................................................................................................. H. aquila sp. nov. – Head narrow [HW 36.8–42.3 (39.2–40.8) % HL]; eye small [ED 22.2–29.9 (24.6–28.3) % HL]; 22–47 (medians 27–36) outer upper jaw teeth.................................................................................. 5 – Head broad [HW 39.9–48.0 (42.9–45.1) % HL]; eye variable [ED 24.6–31.5 (27.5–30.4) % HL]; 39–70 (medians 45–56) outer upper jaw teeth.................................................................................. 8 5. Pre-dorsal distance short [PrD 33.3–37.0 (35.3–36.1) % SL]; interorbital area broad [IOW 50.9– 61.0 (53.8–55.5) % SL]; 3–4, rarely 5 infraorbital cheek scales; 25–47 (medians 30–36) outer upper jaw teeth............................................................................................................................................. 6 – Pre-dorsal distance long [PrD 36.1–39.2 (37.3–38.0) % SL]; interorbital area narrow [IOW 44.9– 52.7 (48.1–48.9) % SL]; 4–7 infraorbital cheek scales; 22–36 (medians 27–29) outer upper jaw teeth.................................................................................................................................................... 7 6. Pre-pectoral distance short [PrP 33.1–38.2 (36.0) % SL]; caudal peduncle long [CPL 15.7–17.5 (16.6) % SL]; head short [HL 33.4–37.0 (35.1) % SL]; dominant males yellow-green with a bright red anterior part of flank (Fig. 9c)................................................................................... H. mentatus – Pre-pectoral distance long [PrP 36.4–39.4 (38.1) % SL]; caudal peduncle short [CPL 13.4–16.1 (14.8) % SL]; head long [HL 35.9–37.9 (36.9) % SL]; dominant males (and females) light blue with black cheek (Fig. 18c)........................................................................................... H. glaucus sp. nov. 7. Cheek deep [ChD 27.6–33.5 (31.1) % SL]; eye small [ED 22.2–28.3 (24.6) % SL]; 22–27 upper lateral line scales; 12–16 scales between upper lateral line and first anal spine; dominant males cream-coloured with orange operculum and dorsal part of head and light blue snout (Fig. 12c)....................................................................................................................... H. rex sp. nov. – Cheek shallow [ChD 27.0–29.6 (28.3) % SL]; eye large [ED 26.7–29.5 (28.3) % SL]; 19–21 upper lateral line scales; 9–11 scales between upper lateral line and first anal spine; dominant males yellow with an orange anterior part of flank (Fig. 15c)....................................................... H. simba sp. nov. 8. Body pyriform; head convex and broad [HW 42.9–48.0 (45.1) % HL]; cheek deep [ChD 27.1–35.2 (30.9) % HL]; lower jaw broad [LJW 44.7–53.3 (49.3) % LJL]; dominant males grey dorsally and yellow ventrally (Fig. 24c).................................................................................. H. kimondo sp. nov. – Body oval; head straight or slightly convex and narrow [HW 39.9–44.4 (42.6–43.4) % HL]; cheek shallow [ChD 22.4–28.0 (23.2–26.0) % HL]; lower jaw narrow [LJW 38.5–45.5 (40.8–42.5) % LJL]............................................................................................................................................... 9 9. Anal-fin base long [AFB 18.4–20.3 (19.3) % SL]; cheek deep [ChD 23.3–28.0 (26.0) % HL]; dominant males olive-green with an orange-red anterior part of flank and well-defined mid-lateral and dorsal-lateral bands (Fig. 27c)....................................................................... H. falcatus sp. nov. – Anal-fin base short [AFB 17.9–18.6 (18.3) % SL], cheek shallow [ChD 22.4–24.9 (23.2) % HL]; non-dominant males dusky greenish with 5–7 vertical stripes (Fig. 30c)........ H. curvidens sp. nov., Published as part of Vranken, Nathan, Steenberge, Maarten Van, Heylen, Annelies, Decru, Eva & Snoeks, Jos, 2022, From a pair to a dozen: the piscivorous species of Haplochromis (Cichlidae) from the Lake Edward system, pp. 1-94 in European Journal of Taxonomy 815 on pages 73-75, DOI: 10.5852/ejt.2022.815.1749, http://zenodo.org/record/6484153, {"references":["Greenwood P. H. 1973. A revision of the Haplochromis and related species (Pisces: Cichlidae) from Lake George, Uganda. Bulletin of the British Museum (Natural History) Zoology 25: 139 - 242.","Snoeks J. 1994. The Haplochromine Fishes (Teleostei, Cichlidae) of Lake Kivu, East Africa: a Taxonomic Revision with Notes on their Ecology. Royal Museum for Central Africa, Tervuren."]}

Haplochromis rex sp. nov. urn:lsid:zoobank.org:act: EF8B7189-BCA4-4A72-8387-498689D76CEB Figs 1–2, 11–13; Table 1 Differential diagnosis Species with a piscivorous morphology; head narrow [HW 36.8–41.6 (mean 39.2) % HL]; cheek deep [ChD 27.6–33.5 (31.1) % HL]; eye small [ED 22.2–28.3 (24.6) % HL]; outer oral teeth few and large [UOT 24–36 (median 29)]; dominant males cream-coloured with an orange operculum and a light blue snout. Amongst piscivorous species from the Lake Edward system, H. rex sp. nov. differs from H. latifrons sp. nov. and H. mentatus by the combination of a shorter caudal peduncle [CPL 13.5–16.2 (14.8) vs 15.7–18.0 (16.6–17.0) % SL] and a narrower interorbital area [IOW 44.9–52.7 (48.9) vs 51.3–63.3 (55.5–60.0) % HW]; further from H. latifrons sp. nov. by absence vs presence of a well-defined midlateral band; further from H. mentatus by dominant males cream-coloured with an orange operculum vs yellow-green with a red anterior part of flank. It differs from H. simba sp. nov. by the combination of a larger number of longitudinal line scales (LongL 34–38 vs 32–33), a larger number of scales between first anal-fin spine and upper lateral line (ULL-A 12–16 vs 9–11), absent or weakly developed vs strongly developed mental prominence, and dominant males cream-coloured with an orange operculum and a light blue snout vs yellow with an orange anterior part of flank and a yellow snout. It differs from H. glaucus sp. nov. by the combination of a narrower interorbital area [IOW 44.9–52.7 (48.9) vs 50.9–57.1 (53.8) % HW], a steeper snout (40–50° vs 30–40°), rounded vs acute oral jaws in dorsal view, and dominant males cream-coloured with an orange operculum and a light blue snout vs uniformly light blue with a dusky snout. It differs from H. aquila sp. nov. by the combination of a deeper lacrimal [LaD 18.9–22.5 (20.8) vs 17.0–19.1 (18.3) % HL], smaller eye [ED 22.2–28.3 (24.6) vs 30.0–31.5 (30.6) % HL], and dominant males cream-coloured with an orange operculum vs light grey with a black head. It differs from H. kimondo sp. nov., H. falcatus sp. nov., H. curvidens sp. nov., H. pardus sp. nov., H. quasimodo sp. nov., and H. squamipinnis by the combination of stout vs slim oral jaws, large vs small outer oral teeth, and a smaller number of outer upper jaw teeth [UOT 24–36 (29) vs 39–79 (45–58)]. It further differs from H. kimondo sp. nov., H. curvidens sp. nov., and H. quasimodo sp. nov. by a narrower head [HW 36.8–41.6 (39.2) vs 42.0–48.1 (43.4–45.3) % HL]; further from H. kimondo sp. nov. and H. quasimodo sp. nov. by dominant males cream-coloured with an orange operculum vs grey dorsally and yellow or blue-black ventrally. It further differs from H. falcatus sp. nov., H. pardus sp. nov., and H. squamipinnis by a larger number of longitudinal line scales (LongL 34–38 vs 29–33, rarely 34) and dominant males cream-coloured with an orange operculum vs uniformly olive-green with an orange-red anterior part of flank, speckled to uniformly black, or uniformly slate blue, respectively. Etymology Specific name from the Latin ‘ rex ’ for ‘king’ (one that holds a preeminent position); referring to very small eyes, deep cheeks, and strong jaws set with large and acute teeth indicating this piscivore has most specialised morphology among all piscivores from the Lake Edward system to hunt on large prey (Barel et al. 1977). Material examined Holotype UGANDA • ♂, 148.5 mm SL; Lake Edward; 0°24′16.0″ S, 29°46′24.8″ E; 24 Jan. 2018; HIPE3 exped. leg.; bought at Rwenshama landing site; RMCA 2018.008.P.0345. Paratypes DEMOCRATIC REPUBLIC OF THE CONGO • 1 ♂, 131.5 mm SL; “Lac Edouard: Vitshumbi (contre le pier)” [Lake Edward: Vitshumbi (against the pier)]; 0°40′50.6″ S, 29°23′22.6″ E (inferred); 26 Mar. 1953; KEA exped. leg.; IRSNB 13474 • 1 ♂, 154.2 mm SL; “Lac Edouard: 2–3 km ± 500 m au large à l’Ouest de Kiavinionge” [Lake Edward: 2–3 km ± 500 m offshore west of Kiavinionge]; 0°11′39″ S, 29°32′31″ E (inferred); 1 Jun. 1953; KEA exped. leg.; IRSNB 13480 • 1 ♀, 154.8 mm SL; “Lac Edouard: 2–3 km à l’Ouest de Kiavinionge” [Lake Edward: 2–3 km west of Kiavinionge]; 0°11′39″ S, 29°32′31″ E (inferred); 1 Jun. 1953; KEA exped. leg.; IRSNB 13485. UGANDA – Lake Edward • 1 ♂, 114.4 mm SL; 0°12′00.0″ S, 29°47′38.4″ E; 23 Oct. 2016; HIPE1 exped. leg.; deep catch, open water ± 20 m deep; RMCA 2016.035.P.0220 • 1 ♂, 154.5 mm SL; 0°08′51.9″ S, 29°52′59.6″ E; 24 Oct. 2016; HIPE1 exped. leg.; bought at Katwe landing site; RMCA 2016.035.P.0221 • 1 ♂, 114.7 mm SL; Rwenshama, rocky shore; 0°24′05.7″ S, 29°46′35.1″ E; 26 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0352 • 1 ♂, 2 ♀♀, 101.0–140.0 mm SL; Kayanja offshore; 0°05′34.8″ S, 29°45′28.8″ E; 31 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0353 to 0355 • 1 ♀, 80.5 mm SL; Kayanja offshore; 0°05′31.2″ S, 29°45′30.3″ E; 20 Jan. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0342 • 1 ♂, 1 ♀, 134.6, 147.7 mm SL; 0°24′16.0″ S, 29°46′24.8″ E; 24 Jan. 2018; HIPE3 exped. leg.; bought at Rwenshama landing site; RMCA 2018.008.P.0343 to 0344 • 1 ♀, 84.2 mm SL; Rwenshama, offshore; 0°24′14.4″ S, 29°45′57.0″ E; 24 Jan. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0346 • 1 ♂, 128.3 mm SL; Kagoro fishing ground; 0°12′50.1″ S, 29°49′19.7″ E; 4 Feb. 2018; HIPE3 exped. leg.; open water; RMCA 2018.008.P.0347 • 1 ♀, 134.2 mm SL; Kayanja offshore; 0°05′34.8″ S, 29°45′28.8″ E; 21 Mar. 2019; HIPE4 exped. leg.; RMCA 2019.002.P.0014. Description Based on 16 specimens (80.5–154.8 mm SL); body shallow (Table 1) and oval to rectangular (Fig. 11). Head long, very narrow, and with a (weakly) convex dorsal outline; cheek deep; lacrimal very deep; eye very small and very high on head; interorbital area very narrow. Snout long, rounded in dorsal view, blunt, and slopes at 40–50°; premaxillary pedicel long and strongly prominent. Jaws isognathous to weakly prognathous, long, very narrow, stout, and rounded in dorsal view; jaws expand slightly laterally, hereby often broader than snout. Gape large and slopes gently at 15–25°; maxilla extends to vertical through anterior margin of orbit. Lower jaw relatively deep and with a straight ventral outline in lateral view, mental prominence absent or weakly developed, and lower jaw side with an inclination of 30–35° to horizontal in anterior view. Upper jaw strongly expanded anteriorly. Lips and oral mucosa large. Neurocranium shallow, ethmo-vomerine block decurved, preorbital region very shallow (18–20% NL), orbital region shallow (26–29% NL), and supraoccipital crest shallow and wedge-shaped (Fig. 12b). Outer oral teeth few, unicuspid, and large. Necks very stout, conical, and straight; crowns weakly recurved, rarely straight, and acutely pointed. Dental arcades rounded. Outer teeth widely and regularly set with neck-distances of 1–2 neck-widths. No enlarged teeth posterior in upper jaw. Inner teeth straight to recurved, unicuspid, and relatively small in small specimens ( 125 mm SL). Tooth bands very slender crescent-shaped with 1–3 rows of inner teeth, and narrow posteriorly until only outer row remains past ⅔ length of tooth band in upper jaw, past ½ length of tooth band in lower jaw. Inner teeth widely and regularly set on 1–2 outer neck-widths from outer row; implantation erect in first row and recumbent in subsequent rows; size decreases slightly buccally and posteriorly. Lower pharyngeal bone average in length, narrow, slim, and shallow over whole length (Fig. 13). Pharyngeal teeth relatively large and slender; major cusps acutely pointed; cusp gaps concave; minor cusps and cusp protuberances mostly absent. Teeth in two median longitudinal rows equal in size and form to lateral teeth, 11 in each row. posterior transverse row with 15–16 teeth, implanted recumbently with a lateral inclination; major cusps weakly recurved, bluntly pointed, and laterally compressed; minor cusps mostly absent. All scales relatively small, many scale rows between anal fin and upper lateral line (12–16), infraorbital scales on cheek (4–7), and scales around caudal peduncle (18–20); many scales in longitudinal line (34–38); chest scales small; transition to larger flank scales gradual. Minute scales on proximal half of caudal fin. Caudal fin emarginate; dorsal and anal fins reach to between verticals through two scales anterior to and one scale posterior to caudal-fin base. Pectoral fin reaches to between anal opening and three scales anterior to this point; pelvic fin reaches to anal opening in females, to first anal-fin spine in males; first branched pelvic-fin ray not elongated. Ceratobranchial gill rakers in outer row of first gill arch short, stout, and simple; posteriormost rakers anvil-shaped to weakly trifid. Epibranchial gill rakers relatively slender and simple. Colouration in life Dominant males: body cream-coloured with faint orange sheen; dorsum greyish; belly and chest black; caudal peduncle with blue sheen (Fig. 12c). Operculum, dorsal part of head, and interorbital area orange; snout, lower jaw, and lips light blue; cheek white and speckled black; eye with (dark) grey outer ring and silver inner ring. Nostril, interorbital, supraorbital, and lacrimal stripes faint; mental blotch present. Pectoral and dorsal fin hyaline; dorsal fin with a dusky base, a dusky and maculated posterior part, and orange lappets, except for black anteriormost lappets. Pelvic fin black; anal and caudal fins orange-red and with dusky bases; anal fin with dusky posterior part and 1–3 small orange egg-spots with dusky rings; caudal fin dorsally maculated orange. Females and juveniles: body cream-coloured with yellow sheen; dorsum and dorsal part of head greyish; belly, chest, operculum, cheek, and lower jaw white; snout dusky and faint light blue; eye with (dark) grey outer ring and silver inner ring (Fig. 12d). Nostril, interorbital, supraorbital, and lacrimal stripes slender and well-defined; mental blotch present. Pectoral, pelvic, anal, and caudal fins yellowish; anal fin with 2 spots resembling egg-spots; caudal fin with dusky base and dorsal part; dorsal fin dusky and with black lappets. Preserved colouration Body, belly, operculum, and cheek yellowish; dorsum brown; chest yellowish in females, black in males (Fig. 12a). Flank rarely with 7–8 very faint narrow vertical stripes. Snout dusky; in males, lower jaw black. In females, all head stripes faint and lacrimal stripe reduced to blotch below eye; in males, head stripes as described here. Nostril and interorbital stripes narrow, well-defined, and horizontally inclined in lateral view, while positioned anterior of nostrils and orbits, respectively, in dorsal view; lacrimal stripe distinct, broad (may cover whole lacrimal), and vaguely delineated; vertical preopercular stripe welldefined; supraorbital stripe present; nape band and mental blotch faint. Pectoral and anal fins yellowish; anal fin with dusky base and posterior margin and 2 small egg-spots in males; pelvic fin yellowish with a black first ray in females, black in males. Dorsal and caudal fins dusky and with yellowish posterior parts; dorsal fin with black lappets and maculated posterior parts. Distribution and ecology Only known from Lake Edward, found over sandy substrates. Based on its morphology, most probably a piscivorous species., Published as part of Vranken, Nathan, Steenberge, Maarten Van, Heylen, Annelies, Decru, Eva & Snoeks, Jos, 2022, From a pair to a dozen: the piscivorous species of Haplochromis (Cichlidae) from the Lake Edward system, pp. 1-94 in European Journal of Taxonomy 815 on pages 24-29, DOI: 10.5852/ejt.2022.815.1749, http://zenodo.org/record/6484153, {"references":["Barel C. D. N., van Oijen M. J. P., Witte F. & Witte-Maas E. L. M. 1977. An introduction to the taxonomy and morphology of the haplochromine Cichlidae from Lake Victoria. Netherlands Journal of Zoology 27: 333 - 380."]}

Haplochromis latifrons sp. nov. urn:lsid:zoobank.org:act: F6BB9E11-AF52-476E-87CE-5C2602859D1B Figs 1–2, 5–7; Table 1 Differential diagnosis Species with a piscivorous morphology; body very shallow [BD 27.2–30.1 (mean 28.6) % SL]; interorbital area flat and broad [IOW 57.4–63.3 (60.0) % HW]; outer oral teeth few and large [UOT 24–42 (median 31)]; females green dorsally, white ventrally, and with a well-defined mid-lateral band; dominant male colour pattern unknown. Amongst piscivorous species from the Lake Edward system, H. latifrons sp. nov. differs from all, except H. mentatus and H. kimondo sp. nov., by the combination of a broader interorbital area [IOW 57.4–63.3 (60.0) vs 39.3–57.1 (43.9–53.8) % HW], a shorter anal fin base [AFB 14.7–17.3 (15.7) vs 17.1–22.2 (18.0–20.5) % SL], and a smaller number of branched anal-fin rays [AFRr 7–8 vs 9–11, rarely 8]. It differs from H. mentatus by the combination of a shorter dorsal-fin base [DFB 47.2–50.1 (49.0) vs 50.3–54.2 (52.3) % SL], a strongly vs weakly prominent premaxillary pedicel, a gentler sloping lower jaw side (25–30° vs 30–45°), juveniles and females green dorsally and white ventrally vs uniformly yellow-green, and presence vs absence of a well-defined mid-lateral band. It differs from H. kimondo sp. nov. by the combination of large vs small outer oral teeth, a smaller number of outer upper jaw teeth [UOT 24–42 (31) vs 43–70 (56)], and a shorter anal fin base [AFB 14.7–17.3 (15.7) vs 17.0–19.2 (18.0) % SL]. Etymology Specific name from Latin ‘ latus ’ for ‘wide’ and ‘ frons ’ for ‘forehead’; referring to very broad interorbital area for a piscivorous species. Material examined Holotype UGANDA • ♀, 158.2 mm SL; Lake Edward; 0°24′16.0″ S, 29°46′24.8″ E; 24 Jan. 2018; HIPE3 exped. leg.; bought at Rwenshama landing site; RMCA 2018.008.P.0330. Paratypes UGANDA – Lake Edward • 3 ♀♀, 122.8–134.9 mm SL; 0°12′00.0″ S, 29°47′38.4″ E; deep catch, open water ± 20 m deep; 23 Oct. 2016; HIPE1 exped. leg.; RMCA 2016.035.P.0199 to 0201 • 1 ♀, 105.4 mm SL; Rwenshama, rocky shore; 0°24′05.7″ S, 29°46′35.1″ E; 25 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0340 • 1 ♂, 82.2 mm SL; Kayanja offshore; 0°05′34.8″ S, 29°45′28.8″ E; 31 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0341 • 1 ♂, 75.3 mm SL; Kayanja offshore; 0°05′31.2″ S, 29°45′30.3″ E; 20 Jan. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0328 • 1 ♀, 155.7 mm SL; 0°24′16.0″ S, 29°46′24.8″ E; 24 Jan. 2018; HIPE3 exped. leg.; bought at Rwenshama landing site; RMCA 2018.008.P.0329. Description Based on 8 specimens (75.3–158.2 mm SL); body very shallow (Table 1) and oval to pyriform; caudal peduncle long (Fig. 5). Head narrow, dorsally flat, and with straight dorsal outline; interorbital area flat and average in width in comparison to generalised H. elegans (but very broad for a piscivorous species); eye very small and high on head; cheek and lacrimal deep. Snout very long, acute, and slopes gently at 30–40°; premaxillary pedicel long and strongly prominent. Jaws isognathous, long, stout, very narrow, and rounded in dorsal view; gape large and slopes gently at 15–25°; maxilla (almost) extends to vertical through anterior margin of orbit. Lower jaw relatively deep and with a weakly convex ventral outline in lateral view, mental prominence absent or weakly developed, and lower jaw side nearly flat with an inclination of 25–30° to horizontal in anterior view. Upper jaw expanded anteriorly. Lips and oral mucosa large. Neurocranium shallow, ethmo-vomerine block decurved, preorbital region very shallow (19–21% NL), orbital region shallow (27–32% NL), and supraoccipital crest very shallow and pyramidical or wedge-shaped (Fig. 6b). Outer oral teeth few, unicuspid, and large. Necks stout, conical, and straight; crowns recurved and acutely pointed. Dental arcades rounded. Outer teeth widely and very irregularly set with neck-distances of 1–5 neck-widths. In upper jaw, 2–3 posteriormost teeth enlarged. Inner teeth relatively large, recurved, unicuspid in large specimens (> 75 mm SL), tricuspid in small specimens ( Lower pharyngeal bone long, narrow, slim, and shallow over entire length (Fig. 7). Pharyngeal teeth relatively large and slender; major cusps acutely pointed; cusp gaps concave; minor cusps and cusp protuberances very small. Teeth in two median longitudinal rows equal in size and form to lateral teeth, about 10 in each row. Posterior transverse row with 15–16 teeth, implanted recumbently with lateral inclination; major cusps recurved, bluntly pointed, and laterally compressed; minor cusps small. Chest scales small; transition to larger flank scales gradual. Minute scales on proximal half of caudal fin. Caudal fin emarginate; dorsal and anal fins reach to vertical through 2–4 scales anterior to caudal-fin base. Pectoral and pelvic fins reach to just anterior to genital opening in females, unknown in males; first branched pelvic-fin ray slightly elongated in all specimens. Ceratobranchial gill rakers in outer row of first gill arch short, stout, and simple; posteriormost rakers simple to trifid or anvil-shaped. Epibranchial gill rakers slender and simple. Colouration in life Dominant males: colour pattern unknown. Females and juveniles: dorsal parts of body and operculum olive-green; ventral parts of body and operculum white; abrupt transition at height of lower lateral line (Fig. 6c–d). Flank with a well-defined black mid-lateral band from posterior margin of eye to caudal-fin base. Dorsum with a blue sheen; cheek and lower jaw white; lacrimal, snout, and lips dusky olive-green; eye with dark grey outer ring and golden inner ring. Nostril and interorbital stripes faint; lacrimal blotch large anteroventrally of eye. Pectoral fin yellowish; pelvic fin hyaline with a greenish sheen; dorsal and caudal fins uniformly dusky, dorsal fin with black lappets (i.e., distal extensions of membrane between spines); anal fin with a hyaline-white base, a dusky-yellow distal part, and 1–2 small spots resembling egg-spots. Preserved colouration Females and juveniles: dorsal part of body brown; ventral part of body, cheek, and lower jaw white; lacrimal, snout and lips dusky (Fig. 6a). Flank with a mid-lateral band from posterior margin of eye to caudal-fin base. Nostril and interorbital stripes faint, lacrimal blotch present. Pectoral fin hyaline; pelvic fin white; dorsal and caudal fins uniformly dusky, dorsal fin with black lappets; anal fin with a white base and dusky distally. Distribution and ecology Only known from Lake Edward, found in offshore areas. Based on its morphology, most probably a piscivorous species.

Haplochromis falcatus sp. nov. urn:lsid:zoobank.org:act: B780DC12-8C55-4B57-98F1-5799201CD438 Figs 1–2, 26–28; Table 1 Differential diagnosis Species with a piscivorous morphology; outer oral teeth many, small, and strongly recurved [UOT 39– 51 (median 45)]; dominant males olive-green with an orange-red anterior part of flank and well-defined mid-lateral and dorsal-lateral bands. Amongst piscivorous species from the Lake Edward system, H. falcatus sp. nov. differs from all except H. curvidens sp. nov. by strongly recurved vs straight to weakly recurved outer jaw teeth. It further differs from H. latifrons sp. nov., H. rex sp. nov., H. simba sp. nov., and H. aquila sp. nov. by the combination of smaller outer oral teeth and a larger number of outer upper jaw teeth [UOT 39–51 (45) vs 22–42 (27–31)]; further from H. latifrons sp. nov., H. rex sp. nov., and H. simba sp. nov. by a shallower lacrimal [LaD 16.1–18.8 (mean 18.0) vs 18.7–23.0 (19.5–20.8) % HL]; further from H. rex sp. nov., H. simba sp. nov., and H. aquila sp. nov. by presence vs absence of well-defined mid-lateral and dorsal-lateral bands, and dominant males uniformly olive-green with an orange-red anterior part of flank vs cream-coloured with an orange operculum and light blue snout, uniformly yellow with an orange anterior part of flank, or light grey with a black head, respectively. It further differs from H. mentatus and H. glaucus sp. nov. by the combination of a longer pre-dorsal distance [PrD 36.9–41.1 (39.5) vs 33.3–37.0 (35.3–36.1) % SL], a gentler lower jaw side (15–25° vs 30–45°), and presence vs absence of well-defined mid-lateral and dorsal-lateral bands; further from H. mentatus by a longer head [HL 36.6–39.6 (38.2) vs 33.4–37.0 (35.1) % SL]; further from H. glaucus sp. nov. by dominant males olive-green with an orange-red anterior part of flank vs uniformly light blue. It further differs from H. kimondo sp. nov. by the combination of an oval vs pyriform body, a straight vs convex dorsal outline of head, shallower cheeks [ChD 23.3–27.4 (exceptionally 28.0 in one specimen) (mean 26.0) vs 27.1–35.2 (30.9) % HL], narrower jaws [LJW 40.2–45.6 (42.5) vs 44.7–53.3 (49.3) % LJL], and dominant males olive-green with an orange-red anterior part of flank vs grey dorsally and yellow ventrally. It differs from H. curvidens sp. nov. and further differs from H. pardus sp. nov. by the combination of a deeper cheek [ChD 25.1–28.0 (exceptionally 23.3 in one specimen) (mean 26.0) vs 20.8–24.9 (22.5– 23.2) % HL] and a longer pre-dorsal distance [PrD 38.2–41.1 (exceptionally 36.9 in one specimen) (mean 39.5) vs 34.1–37.9 (36.0–36.3) % SL]; further from H. curvidens sp. nov. by presence vs absence of well-defined mid-lateral and dorsal-lateral bands; further from H. pardus sp. nov. by larger adult size (max. 137 vs 96 mm SL) and colour pattern of small specimens (H. quasimodo sp. nov. and H. squamipinnis by the combination of a longer head [HL 36.6–39.6 (38.2) vs 33.9–37.2 (35.5–36.0) % SL], a shorter pelvic fin [VL 21.6–25.7 (23.5) vs 25.2–35.4 (28.8–29.4) % SL], and dominant males olive-green with an orange-red anterior part of flank vs light grey dorsally and blue-black ventrally or slate blue, respectively; further from H. squamipinnis by absence vs presence of minute scales on proximal parts of dorsal and anal fin. Etymology Specific name from the Latin ‘ falcatus ’ for ‘sickle-shaped’; referring to acutely pointed sickle-like outer oral teeth. Material examined Holotype UGANDA • ♂, 101.3 mm SL; Lake Edward, Kayanja offshore; 0°05′31.2″ S, 29°45′30.3″ E; 21 Jan. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0401. Paratypes DEMOCRATIC REPUBLIC OF THE CONGO • 1 ♀, 136.8 mm SL; “Lac Edouard: riv. Luniasenke” [Lake Edward: Luniasenke River]; 0°27′19.2″ S, 29°22′08.7″ E (inferred); 04 Jan. 1953; KEA exped. leg.; IRSNB 13469 • 1 ♀, 137.1 mm SL; “Lac Edouard: ½h à l’Ouest d’Ishango” [Lake Edward: ½ hour west of Ishango]; 0°08′14″ S, 29°38′23″ E (inferred); 27 Mar. 1953; KEA exped. leg.; IRSNB 13473. UGANDA – Lake Edward • 1 ♂, 2 ♀♀, 98.6–112.8 mm SL; 1 km east of Nyamugasani River; 0°10′22.8″ S, 219°50′13.2″ E; 22 Oct. 2016; HIPE1 exped. leg.; sand substrate; RMCA 2016.035.P.0256 to 0258 • 1 ♀, 110.8 mm SL; mouth of Kazinga Channel; 0°12′14.4″ S, 29°52′37.2″ E; 23 Mar. 2017; HIPE2 exped. leg.; hard substrate; RMCA 2017.006.P.0415 • 1 ♂, 1 ♀, 119.1–120.7 mm SL, no morphometrics taken; 0°24′16.0″ S, 29°46′24.8″ E; 25 Mar. 2017; HIPE2 exped. leg.; bought at Rwenshama landing site; RMCA 2017.006.P.0416 to 0417 • 1 ♀, 81.3 mm SL; Kayanja offshore; 0°05′34.8″ S, 29°45′28.8″ E; 30 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0418 • 2 ♀♀, 81.6, 93.8 mm SL; Kayanja offshore; 0°05′34.8″ S, 29°45′28.8″ E; 30 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0419 to 0420 • 3 ♀♀, 75.0– 109.6 mm SL; Kayanja offshore; 0°05′34.8″ S, 29°45′28.8″ E; 31 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0421 to 0423 • 1 ♂, 1 ♀, 88.5–103.3 mm SL; Kayanja, offshore; 0°05′31.2″ S, 29°45′30.3″ E; 20 Jan.2018; HIPE3 exped. leg.; RMCA 2018.008.P.0397 to 0398 • 2 ♂♂, 102.1–110.4 mm SL; Kayanja, offshore; 0°05′31.2″ S, 29°45′30.3″ E; 21 Jan. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0399 to 0400 • 1 ♀, 93.5 mm SL; same collection data as for preceding; RMCA 2018.008.P.0402 • 2 ♀♀, 110.9–114.2 mm SL; 0°24′16.0″ S, 29°46′24.8″ E; 24 Jan. 2018; HIPE3 exped. leg.; bought at Rwenshama landing site; RMCA 2018.008.P.0403 to 0404. Description Based on 22 specimens (75.0– 137.1 mm SL); body shallow (Table 1) and oval (Fig. 26). Head very long, narrow, and with a straight dorsal outline; eye small; interorbital area narrow; cheek and lacrimal average in depth. Snout long, acute, and slopes gently at 35–45°; premaxillary pedicel long and strongly prominent. Jaws isognathous to slightly prognathous, long, slim, narrow, and rounded in dorsal view; gape large and slopes gently at 20–30°; maxilla extends to vertical through pupil. Lower jaw shallow and with a straight ventral outline in lateral view, mental prominence absent, and lower jaw side nearly flat with an inclination of 15–25° to horizontal in anterior view; lower jaw expands slightly laterally halfway its length. Upper jaw weakly expanded anteriorly. Lips and oral mucosa large. Neurocranium shallow, ethmo-vomerine block decurved, preorbital region very shallow (18–22% NL), orbital region shallow (28–30% NL), and supraoccipital crest average in depth and wedge-shaped (Fig. 27b). Outer oral teeth numerous, unicuspid, and relatively small. Necks stout, conical, and recurved; crowns recurved to strongly recurved, and acutely pointed. Dental arcades rounded, and with anterior half expanded laterally. Outer teeth closely and regularly set with neck-distances of ½–1 neck-width. No enlarged teeth posterior in upper jaw. Inner teeth small, strongly recurved, unicuspid, and acutely pointed. Tooth bands very slender crescent-shaped with 1–2 rows of inner teeth, and narrow posteriorly until only outer row remains past ⅔ lengths of tooth bands. Inner teeth closely and regularly set on ½–1 neck-width from outer row in lower jaw, on 1–2 neck-widths from outer row in upper jaw; implantation recumbent; size uniform throughout tooth band. Lower pharyngeal bone average in length, triangular, slim, and shallow with a slightly deeper keel (Fig. 28). Pharyngeal teeth relatively large and slender; major cusps acutely pointed; cusp gaps straight; minor cusps and cusp protuberances very small. Teeth in two median longitudinal rows equal in size and form to lateral teeth, 10 in each row. Posterior transverse row with 16–21 teeth, implanted erectly with a lateral inclination; major cusps weakly recurved, bluntly pointed, and laterally compressed; minor cusps mostly absent. Chest scales small; transition to larger flank scales gradual. Minute scales on proximal half of caudal fin. Caudal fin subtruncate; dorsal and anal fins reach to vertical through caudal-fin base. Pectoral fin reaches to between genital opening and first anal-fin spine; pelvic fin reaches to between genital opening and first anal-fin spine in females, to first anal fin branched ray in males; first branched pelvic-fin ray elongated in dominant males. Ceratobranchial gill rakers in outer row of first gill arch very short, stout, and simple; posteriormost rakers weakly anvil-shaped. Epibranchial gill rakers relatively slender and simple. Colouration in life Dominant males: body olive-green with yellow sheen; anterior part of flank and operculum bright orange-red; belly and chest speckled black (Fig. 27c). Cheek olive-green with red sheen; snout and lips dusky; lower lip with blue sheen; branchiostegal membrane black; eye with dark silver outer ring and silver to golden inner ring. Flank with well-defined mid-lateral and dorsal-lateral bands; dorsum with 5–7 faint vertical stripes between dorsal-fin base and dorsal-lateral band. Lacrimal, nostril, and interorbital stripes and a mental blotch well defined; supraorbital stripe and nape band faint. Pectoral fin hyaline; pelvic fin black; dorsal fin dusky and with black lappets; anal fin faint orange-red and with dusky base and posterior part, and 3 small yellow egg spots with hyaline rings. Caudal fin with dusky base and hyaline distal part. Non-dominant males: similar to dominant males except for white belly and chest and faint orange-red antero-dorsal part of flank above mid-lateral band. Females and juveniles: body, operculum, cheek, and lacrimal olive-green yellow; belly, chest, and lower jaw white; eye with dark silver outer ring and silver to golden inner ring (Fig. 27d). Flank with welldefined mid-lateral and dorsal-lateral bands; dorsum with 5–7 faint vertical stripes between dorsal-fin base and dorsal-lateral band. Lacrimal, nostril, and interorbital stripes and a mental blotch well defined; supraorbital stripe and nape band faint. Pectoral and pelvic fins yellowish; dorsal fin dusky and with black lappets; anal fin yellow and with 2–3 small spots resembling egg spots; caudal fin dusky. Preserved colouration Dorsal part of body brown; ventral part of body yellowish to white in females; chest and belly speckled black in dominant males (Fig. 27a). Cheek yellowish and snout dusky. Flank with well-defined midlateral and dorsal-lateral bands; dorsum with 5–7 faint vertical stripes between dorsal-fin base and dorsallateral band. Lacrimal, nostril, and interorbital stripes and a mental blotch well defined; supraorbital stripe and nape band faint. Pectoral fin dusky; pelvic fin dusky in females, black in dominant males; dorsal fin dusky and with black lappets; anal fin with dusky base and posterior part, yellowish distal part, and 1–3 small egg spots; caudal fin dusky and faintly maculated in dorsal part. Distribution and ecology Only known from Lake Edward, found over sandy substrates. Based on its morphology, most probably a piscivorous species., Published as part of Vranken, Nathan, Steenberge, Maarten Van, Heylen, Annelies, Decru, Eva & Snoeks, Jos, 2022, From a pair to a dozen: the piscivorous species of Haplochromis (Cichlidae) from the Lake Edward system, pp. 1-94 in European Journal of Taxonomy 815 on pages 48-53, DOI: 10.5852/ejt.2022.815.1749, http://zenodo.org/record/6484153

Haplochromis curvidens sp. nov. urn:lsid:zoobank.org:act: E11E39DC-0085-4CB5-9E4C-E3EA0C11216F Figs 1–2, 29–31; Table 1 Differential diagnosis Species with a piscivorous morphology; outer oral teeth many, small, and (strongly) recurved [UOT 45–60 (median 49)]; non-dominant males dusky green with a blue sheen and 5–7 (faint) vertical stripes. Amongst piscivorous species from the Lake Edward system, H. curvidens sp. nov. differs from H. latifrons sp. nov., H. mentatus, H. rex sp. nov., H. simba sp. nov., H. glaucus sp. nov., and H. aquila sp. nov. by small vs large outer oral teeth and a larger number of outer upper jaw teeth [UOT 45–60 (49) vs 22–47 (27–36)]; further from H. latifrons sp. nov., H. mentatus, H. rex sp. nov., H. simba sp. nov., and H. glaucus sp. nov. by a shallower lacrimal [LaD 16.0–17.8 (mean 16.7) vs 18.0–23.0 (19.5–20.8) % HL]; further from H. rex sp. nov., H. simba sp. nov., H. glaucus sp. nov., and H. aquila sp. nov. by a shallower cheek [ChD 22.4–24.9 (23.2) vs 26.2–33.5 (28.3–31.1) % HL]. It differs from H. kimondo sp. nov., H. falcatus sp. nov., and H. quasimodo sp. nov. by the combination of a shallower cheek [ChD 22.4–24.9 (23.2) vs 24.8–35.2 (exceptionally 23.3 and 23.7 in one specimen of H. falcatus sp. nov. and H. quasimodo sp. nov., respectively) (means 26.0–30.9) % HL] and absence vs presence of a well-defined mid-lateral band; further from H. kimondo sp. nov. by narrower jaws [LJW 38.5–43.2 (40.8) vs 44.7–53.3 (49.3) % LJL]; further from H. falcatus sp. nov. by a shorter pre-dorsal distance [PrD 34.5–37.9 (36.3) vs 38.2–41.1 (exceptionally 36.9 in one specimen) (mean 39.5) % SL]; further from H. quasimodo sp. nov. by a shallower body [BD 29.0–32.0 (30.8) vs 33.5–41.7 (37.4) % SL]. It differs from H. pardus sp. nov. by the combination of a shorter anal fin base [AFB 17.9–18.6 (18.3) vs 19.2–22.2 (20.5) % SL], a slightly broader interorbital area [IOW 46.4–52.5 (49.1) vs 39.3–48.4 (43.9) % HW], and all specimens faint yellow to dusky green vs speckled to uniformly black. It differs from H. squamipinnis sp. nov. by the combination of a shallower body [BD 29.0–32.0 (30.8) vs 32.4–39.3 (35.7) % SL], a shallower cheek [ChD 22.4–24.9 (23.2) vs 24.9–36.0 (29.0) % HL], and absence vs presence of minute scales on proximal parts of dorsal and anal fins. Etymology Specific name from the Latin ‘ curvus ’ for ‘curvature’, and ‘ dentatus ’ for ‘tooth’; referring to strongly recurved oral teeth. Material examined Holotype UGANDA • ♂, 112.0 mm SL; Lake Edward; 0°24′16.0″ S, 29°46′24.8″ E; 9 Nov. 2016; HIPE1 exped. leg.; bought at Rwenshama landing site; RMCA 2016.035.P.0219. Paratypes UGANDA – Lake Edward • 1 ♂, 94.8 mm SL; ‘Coral Reef’, hard substrate at mouth of Nyamugasani river; 0°10′08.4″ S, 29°49′37.2″ E; 21 Oct. 2016; HIPE1 exped. leg.; RMCA 2016.035.P.0215 • 2 ♂♂, 1 ♀, 91.2–102.5 mm SL; Mukutu Kihinga, rocky offshore of Mweya; 0°11′31.2″ S, 29°52′26.4″ E; 23 Oct. 2016; HIPE1 exped. leg.; RMCA 2016.035.P.0216 to 0218 • 1 ♀, 90.2 mm SL; islands near Katwe; 0°10′04.9″ S, 29°52′27.4″ E; 18 Jan. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0340 • 1 ♂, 101.1 mm SL; 0°24′16.0″ S, 29°46′24.8″ E; 24 Jan. 2018; HIPE3 exped. leg.; bought at Rwenshama landing site; RMCA 2018.008.P.0341 • 1 ♀, 92.3 mm SL, no morphometrics taken; islands near Katwe; 0°10′04.9″ S, 29°52′27.4″ E; 18 Jan. 2018; HIPE3 exped. leg.; IRSNB 919. Description Based on 8 specimens (90.2–112.0 mm SL); body shallow (Table 1) and oval (Fig. 29). Head long, narrow, and with a straight to very gently convex dorsal outline; eye and interorbital area average in width; cheek and lacrimal average in depth. Snout average in length, narrow, acute, and slopes gently at 35–45°; premaxillary pedicel long and strongly prominent. Jaws isognathous to weakly prognathous, long, slim, very narrow, and rounded in dorsal view; gape large and slopes gently at 15–30°; maxilla (almost) extends to vertical through anterior point of pupil. Lower jaw shallow and with a straight ventral outline in lateral view, mental prominence absent, and lower jaw side nearly flat with an inclination of 20–30° to horizontal in anterior view. Upper jaw weakly expanded anteriorly. Lips and oral mucosa thin. Neurocranium shallow, ethmo-vomerine block decurved, preorbital region very shallow (19–21% NL), orbital region shallow (28–31% NL), and supraoccipital crest shallow and wedge-shaped (Fig. 30b). Outer oral teeth numerous, unicuspid, and small. Necks slender, cylindrical to conical, and weakly recurved; crowns recurved and acutely pointed. Dental arcades rounded. Outer teeth closely and regularly set with neck-distances of ½ neck-width. In upper jaw, 1–3 posteriormost teeth enlarged. Inner teeth small, strongly recurved, unicuspid, and acutely pointed. Tooth bands very slender crescent-shaped with 1–2 rows of inner teeth, and narrow posteriorly until only outer row remains past ⅔ lengths of tooth bands. Inner teeth closely and regularly set on ½–1 neck-width from outer row in lower jaw, on 1–2 neck-widths from outer row in upper jaw; implantation recumbent; size uniform throughout tooth band. Lower pharyngeal bone average in length, narrow, slim, and shallow with a slightly deeper keel (Fig. 31). Pharyngeal teeth relatively large and slender; major cusps acutely pointed; cusp gaps straight; minor cusps and cusp protuberances very small. Teeth in two median longitudinal rows equal in size and form to lateral teeth, 10–11 in each row. Posterior transverse row with 20–22 teeth, implanted erectly with a lateral inclination; major cusps weakly recurved, bluntly pointed, and laterally compressed; minor cusps mostly absent. Chest scales small; transition to larger flank scales gradual. Minute scales on proximal half of caudal fin. Caudal fin emarginate; dorsal and anal fins reach to between verticals through caudal-fin base and two scales anterior to this vertical. Pectoral and pelvic fins reach to anal opening, pelvic fin reaches to first anal-fin spine in males; first branched pelvic-fin ray elongated in all specimens. Ceratobranchial gill rakers in outer row of first gill arch short, stout, and simple; posteriormost rakers anvil-shaped, bi-, or trifid. Epibranchial gill rakers slender and simple. Colouration in life Dominant males: colour pattern unknown. Non-dominant males: based on pictures of two recently deceased specimens (Fig. 30c). Dorsal half of body dusky greenish; ventral half of body faint yellow; transition gradual; flank with blue sheen and 5–7 vertical stripes; belly, chest, and cheek whitish; operculum yellowish; snout dusky; eye with (dark) grey outer ring and silver inner ring. Nostril and lacrimal stripes and nape band present; interorbital and supraorbital stripes faint. Pectoral fin yellowish; pelvic fin black; dorsal fin dusky and with black lappets; anal fin faint orange and with 1–3 relatively large, yellow egg-spots with dusky rings; caudal fin dusky. Females and juveniles: body faint yellow with a greenish dorsum; belly and chest white; snout dusky; eye with (dark) grey outer ring and silver inner ring (Fig. 30d). Nostril, interorbital, supraorbital, and lacrimal stripes and mental blotch faint. Pectoral and pelvic fin yellowish; dorsal fin dusky and with black lappets; anal fin yellowish and with 2 spots resembling egg-spots; caudal fin dusky. Preserved colouration Body brown; dorsum dark brown; chest and belly white; cheek and operculum yellowish; snout dusky (Fig. 30a). Flank with 5–7 (faint) vertical stripes. Nostril and lacrimal stripes and mental blotch present; interorbital and supraorbital stripes faint. Pectoral fin hyaline; pelvic fin yellowish with black first rays in females, black in males; dorsal and caudal fins dusky; dorsal fin with black lappets and posterior sooty part; caudal fin with maculated dorsal part; anal fin yellowish and with dusky distal margin and 1–3 relatively large egg-spots. Distribution and ecology Only known from Lake Edward, found over muddy substrates. Based on its morphology, most probably a piscivorous species.

Haplochromis quasimodo sp. nov. urn:lsid:zoobank.org:act: 6AE722D9-3BF6-4DD5-8637-55B8400EDD11 Figs 1–2, 35–37; Table 1 Differential diagnosis Species with a piscivorous morphology; body rather deep [BD 33.5–41.7 (mean 37.4) % SL]; interorbital area narrow [IOW 40.5–48.7 (43.9) % HL]; outer oral teeth many and small [UOT 46–71 (median 58)]; dominant males light grey dorsally and blue-black ventrally. Amongst piscivorous species from the Lake Edward system, H. quasimodo sp. nov. differs from H. latifrons sp. nov., H. mentatus, H. rex sp. nov., H. simba sp. nov., H. glaucus sp. nov., and H. aquila sp. nov. by the combination of small vs large outer oral teeth and a larger number of outer upper jaw teeth [UOT 46–71 (58) vs 22–47 (27–36)]; further from H. mentatus, H. rex sp. nov., H. simba sp. nov., H. glaucus sp. nov., and H. aquila sp. nov. by presence vs absence of a well-defined mid-lateral band. It further differs from H. latifrons sp. nov. and H. mentatus by a deeper body [BD 33.5–41.7 (37.4) vs 27.2–32.3 (28.6–31.2) % SL]; further from H. rex sp. nov., H. simba sp. nov., and H. glaucus sp. nov. by a broader head [HW 42.0–48.1 (45.3) vs 36.8–41.6 (39.2–40.8) % HL]. It differs from H. kimondo sp. nov. and H. squamipinnis by a narrower interorbital area [IOW 40.5– 48.7 (43.9) vs 48.6–58.5 (51.9–52.8) % HW]; further from H. kimondo sp. nov. by the combination of a rhomboid vs pyriform body, a concave to weakly convex vs convex dorsal outline of head, a gentler sloping snout (30–40° vs 40–50°), and dominant males light grey dorsally and blue-black vs grey dorsally and yellow ventrally; further from H. squamipinnis by a gentler gape inclination (20–35° vs 30–45°), a shorter lower jaw [LJL 44.2–49.6 (47.1) vs 47.8–58.6 (52.5) % HL], mostly absence vs presence of minute scales on proximal part of dorsal fin (rarely few rows of 1–4 scales present on dorsal fin in H. quasimodo sp. nov.), and dominant males light grey dorsally and blue-black ventrally vs slate blue. It differs from H. falcatus sp. nov. by the combination of a shorter head [HL 33.9–37.2 (35.5) vs 36.6– 39.6 (38.2) % SL], a longer pelvic fin [VL 26.2–33.7 (29.4) vs 21.6–25.7 (23.5) % SL], weakly recurved vs strongly recurved outer oral teeth, and dominant males light grey dorsally and blue-black ventrally vs olive-green with an orange-red anterior part of flank. It differs from H. curvidens sp. nov. and H. pardus sp. nov. by a deeper cheek [ChD 24.8–32.9 (exceptionally 23.7 in one specimen) (mean 27.5) vs 20.8–24.9 (22.5–23.2) % HL]; further from H. curvidens sp. nov. by a deeper body [BD 33.5–41.7 (37.4) vs 29.0–32.0 (30.8) % SL] and presence vs absence of a well-defined mid-lateral band; further from H. pardus sp. nov. by a larger number of caudal peduncle scales (CPS 17–20, rarely 16 vs 16, rarely 17), a larger adult size (max. 165 vs 96 mm SL), and colour pattern of small specimens (H. schubotziellus Greenwood, 1973 in overall habitus and colour pattern. It differs from the holotype of H. schubotziellus (NHMUK 1972.6.2.351; ♀, 75.0 mm SL; Lake George, Kankurunga Island) by a smaller eye [ED 26.2–31.8 (29.2) vs 33.7% HL], a longer lower jaw [LJL 44.2–49.6 (47.1) vs 42.6% HL], a deeper cheek [ChD 23.7–32.9 (27.5) vs 24.0% HL], outer oral teeth with no to a small minor cusp vs a well-defined minor cusp, and inner oral teeth set in 1–2 weakly defined rows vs 2 well-defined rows in both jaws. Etymology Specific name from Quasimodo, hunchbacked character in Victor Hugo’s novel ‘Notre-Dame de Paris’ (1831); referring to rather shallow head and deep and rhomboid bodies of large specimens. Material examined Holotype UGANDA • ♂, 120.4 mm SL; Lake Edward; 0°21’31.7″ S, 29°43’17.7″ E; deep catch, open water ± 30 m deep; 1 Feb. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0336. Paratypes DEMOCRATIC REPUBLIC OF THE CONGO • 1 ♀, 147.8 mm SL; “Lac Edouard: 2–3 km ± 500 m au large à l’Ouest de Kiavinionge” [Lake Edward: 2–3 km ± 500 m offshore west of Kiavinionge]; 0°11′39″ S, 29°32′31″ E (inferred); 1 Jun. 1953; KEA exped. leg.; IRSNB 13480 • 1 ♀, 164.9 mm SL; “Lac Edouard: Kaniatzi (partie N. du lac)” [Lake Edward: Kaniatzi (northern part of Lake)]; 1 Jun. 1953; KEA exped. leg.; IRSNB 13481 • 1 ♂, 147.6 mm SL; “Lac Edouard: 2–3 km à l’Ouest de Kiavinionge” [Lake Edward: 2–3 km west of Kiavinionge]; 0°11′39″ S, 29°32′31″ E (inferred); 1 Jun. 1953; KEA exped. leg.; IRSNB 13485 • 2 ♀♀, 141.5 mm SL; “Lac Edouard: Vitshumbi (au Nord)” [Lake Edward: north of Vitshumbi]; 0°40′50.6″ S, 29°23′22.6″ E (inferred); 02 Jul. 1953; KEA exped. leg.; IRSNB 13488. UGANDA – Lake Edward • 2 ♀♀, 2 ♂♂, 92.7–114.3 mm SL; Katoko breeding ground, soft substrate offshore of Katwe; 0°09′43.2″ S, 29°53′16.8″ E; 20 Oct. 2016; HIPE1 exped. leg.; RMCA 2016.035.P.0208 to 0211 • 2 ♀♀, 83.4, 93.1 mm SL; Mukutu Kihinga, rocky offshore of Mweya; 0°11′31.2″ S, 29°52′26.4″ E; 23 Oct. 2016; HIPE1 exped. leg.; RMCA 2016.035.P.0213 to 0214 • 1 ♂, 78.9 mm SL; Mukutu Kihinga, rocky offshore of Mweya; 0°11′31.2″ S, 29°52′26.4″ E; 23 Oct. 2016; HIPE1 exped. leg.; RMCA 2016.035.P.0212 • 3 ♀♀, 79.4–110.1 mm SL; Rwenshama, rocky shore; 0°24′05.7″ S, 29°46′35.1″ E; 26 Mar. 2017; HIPE2 exped. leg.; RMCA 2017.006.P.0349 to 0351 • 1 ♂, 89.1 mm SL; islands near Katwe; 0°10′04.9″ S, 29°52′27.4″ E; 18 Jan. 2018; HIPE3 exped. leg.; RMCA 2018.008.P.0334 • 1 ♀, 130.7 mm SL; 0°24′16.0″ S, 29°46′24.8″ E; 24 Jan. 2018; HIPE3 exped. leg.; bought at Rwenshama landing site; RMCA 2018.008.P.0335 • 1 ♀, 148.8 mm SL; 0°21′31.7″ S, 29°43′17.7″ E; 1 Feb. 2018; HIPE3 exped. leg.; deep catch, open water ± 30 m deep; RMCA 2018.008.P.0339 • 1 ♀, 1 ♂, 117.9, 126.2 mm SL; 0°21′31.7″ S, 29°43′17.7″ E; 1 Feb. 2018; HIPE3 exped. leg.; deep catch, open water ± 30 m deep; RMCA 2018.008.P.0337 to 0338. Description Based on 21 specimens (78.9–164.9 mm SL); body average in depth in comparison to generalised H. elegans (but deep for a piscivorous species; Table 1) and rhomboid (Fig. 35). Head average in width and with a concave to weakly convex dorsal outline; eye small; interorbital area very narrow; cheek and lacrimal deep; lacrimal somewhat convex and hereby protrudes somewhat laterally. Snout average in length, acute, and slopes gently at 30–40°; premaxillary pedicel long and prominent. Jaws isognathous, long, slim, narrow, and rounded in dorsal view; gape large and slopes gently at 20–35°; maxilla extends to between verticals through anterior margins of orbit and pupil. Lower jaw shallow and with a straight ventral outline in lateral view, mental prominence absent or weakly developed, and lower jaw side nearly flat with an inclination of 15–25° to horizontal in anterior view. Upper jaw not expanded. Lips and oral mucosa thin. Neurocranium average in depth, ethmo-vomerine block decurved to horizontally inclined, preorbital region shallow (21–24% NL), orbital region average in depth (30–32% NL), and supraoccipital crest deep and pyramidical or weakly wedge-shaped (Fig. 36b). Outer oral teeth numerous and very small. Necks stout, conical, and straight; crowns weakly recurved, unicuspid in large specimens (> 80 mm SL), anteriorly unicuspid and posteriorly mostly bi-, weakly bi-, and weakly tricuspid in small specimens (, Published as part of Vranken, Nathan, Steenberge, Maarten Van, Heylen, Annelies, Decru, Eva & Snoeks, Jos, 2022, From a pair to a dozen: the piscivorous species of Haplochromis (Cichlidae) from the Lake Edward system, pp. 1-94 in European Journal of Taxonomy 815 on pages 62-68, DOI: 10.5852/ejt.2022.815.1749, http://zenodo.org/record/6484153, {"references":["Greenwood P. H. 1973. A revision of the Haplochromis and related species (Pisces: Cichlidae) from Lake George, Uganda. Bulletin of the British Museum (Natural History) Zoology 25: 139 - 242."]}

Genus Enteromius Cope, 1867 The results confirmed the occurrence of two syntopic species of Enteromius with a smooth dorsal fin ray in the Lake Edward system. Specimens of group A were conspecific with the types of E. alberti, except for one type specimen, which was assigned to group B, and with the holotype and paratypes of E. cercops. Enteromius alberti used to be considered a synonym of E. stigmatopygus. Based on our results, and following the principle of priority for species names, we propose to revalidate E. alberti with E. cercops as its junior synonym. A redescription of E. alberti is provided based on the lectotype (here designated) (Fig. 7a) and one paralectotype of E. alberti, the type series of E. cercops, here synonymised with E. alberti, and the additional specimens from the Lake Edward system examined in this study and identified as E. alberti (n = 37). The other paralectotype of E. alberti, which was revealed not to be conspecific with E. alberti, was not included in the redescription. The measurements and meristics can be found in Table 2. An illustration of a fresh specimen of E. alberti, captured during the HIPE expeditions is provided in Fig. 7b. The specimens of group B were most similar to E. mimus, though the PCA on log-transformed measurements (Fig. 6) suggested some differences. Therefore, we decided to assign the specimens of group B to E. cf. mimus with ���cf��� indicating specimens that are comparable to E. mimus but whose identification is uncertain (Decru et al. 2016). Because of the unsuitable original description of E. mimus for modern taxonomic purposes, we decided to make a redescription based on the lectotype (here designed) (Fig. 8) and eight of the nine paralectotypes of E. mimus. We found one paralectotype to have a serrated dorsal fin ray; therefore, it was considered not to be conspecific with the other types and excluded from the redescription. Specimens from the Lake Edward system identified as E. cf. mimus, the specimens from Tshambi and the aberrant lectotype of E. alberti were not included in the redescription, but their measurements and meristics are given in Table 2. A picture of a fresh specimen of E. cf. mimus is given in Fig. 9. For both redescribed species, a lectotype was chosen from the syntypes. For E. alberti, this is the specimen illustrated in the original description by Poll (1939), which is also the largest specimen. The lectotype of E. mimus is the specimen that resembles most the original figure of E. mimus in the Catalogue of the Fresh-water Fishes of Africa in the British Museum (Natural History), Volume IV, by Boulenger (1916). The sampling sites of the two species in the Lake Edward system and the type localities of E. mimus, E. alberti and E. cercops are illustrated in Fig. 10. The specimens from Tshambi, initially identified as E. alberti, are indicated as E. cf. mimus on the map, as found in the present study., Published as part of Maetens, Heleen, Steenberge, Maarten Van, Snoeks, Jos & Decru, Eva, 2020, Revalidation of Enteromius alberti and presence of Enteromius cf. mimus (Cypriniformes: Cyprinidae) in the Lake Edward system, East Africa, pp. 1-28 in European Journal of Taxonomy 700 on page 12, DOI: 10.5852/ejt.2020.700, http://zenodo.org/record/3981069, {"references":["Decru E., Moelants T., De Gelas K., Vreven E., Verheyen E. & Snoeks J. 2016. Taxonomic challenges in freshwater fishes: a mismatch between morphology and DNA barcoding in fish of the north-eastern part of the Congo Basin. Molecular Ecology Resources 16: 342 - 352. https: // doi. org / 10.1111 / 1755 - 0998.12445","Boulenger G. A. 1916. Catalogue of the Fresh-Water Fishes of Africa in the British museum (Natural History), Vol. IV. British Museum (Natural History), London, UK. https: // doi. org / 10.5962 / bhl. title. 8869"]}

Enteromius mimus (Boulenger, 1912) Fig. 8, Table 2 Diagnosis Enteromius mimus (Fig. 8) belongs to the group of species of Enteromius with a flexible last unbranched dorsal fin ray that lacks serrations along its posterior edge. A comparison was made to other species of Enteromius from this group from the East Coast and Nilo-Sudan ichthyofaunal provinces (list made based on information available in, e.g., Greenwood 1962; Okaronon et al. 1997; Seegers et al. 2003; Schmidt et al. 2017; Froese & Pauly 2018). Enteromius mimus can easily be distinguished from the other species of this group from the East Coast and Nilo-Sudan ichthyofaunal regions by the following combinations of characteristics (data from original description, unless otherwise stated): a complete lateral line vs an incomplete lateral line in E. atkinsoni, E. pumilus, E. serengetiensis, E. tongaensis and E. toppini; two pairs of barbels vs one pair in E. pseudotoppini, and no barbels in E. anema and E. profundus; one to three dark spots on the flanks, which sometimes fuse into a mid-lateral line in preserved specimens, starting posterior to the operculum vs a dark line running from the tip of the snout to the caudal fin base in E. bifrenatus and E. yongei, and a thin dark line from the beginning of the operculum to the caudal fin base in E. viviparus; 11���12 scales around the caudal peduncle vs 8 in E. leonensis, 9���10 in E. venustus, and 10 in E. magdalenae and E. yeiensis; 24���27 lateral line scales (median 25) vs 30 in E. lineomaculatus, and 31 E. innocens; 3.5���4.5 scales between the dorsal fin base and the lateral line vs 5.5 in E. unitaeniatus, and 6 in E. usambarae; 3���4 scales between the lateral line and the pelvic fin (median 3) vs 2 in E. trispilopleura, and 2���2.5 in E. neglectus (Boulenger 1907); 7���8 branched dorsal fin rays vs 9 in E. quadripunctatus; a body depth which is larger than the head length vs a body depth which is equal to the head length in E. nigeriensis; a smaller snout length [4.1���7.1% SL (average 6.0) vs 7.1���10.1] in E. radiatus (Greenwood 1970). Enteromius mimus differs from E. perince by a combination of a lower number of lateral line scales [24���27 (median 25) vs 29���30 (30)], a lower number of scales between the dorsal fin base and lateral line [3.5���4.5 (median 3.5) vs 5.5], a lower number of scales around the caudal peduncle [11���12 (median 12) vs 14], a smaller interorbital width [6.3���8.3% SL (average 7.8) vs 9.8���10.2 (10.0)], a larger pre-pectoral distance [26.0���29.0% SL (average 27.4) vs 23.1���24.5 (23.8)], a lower minimum caudal peduncle depth [11.1���14.0% SL (average 12.5) vs 15.0���16.4 (15.5)], a lower maximum caudal peduncle depth [12.9���16.5% SL (average 14.6) vs 17.7���19.4 (18.8)], and a smaller anal fin base length [7.3���8.7% SL (average 8.0) vs 8.5���9.6 (9.1)]. Enteromius mimus differs from E. stigmatopygus by a combination of a smaller pre-pectoral distance [26.0���29.0 (average 27.4) vs 29.1���33.1 (30.8)], and a larger anal fin length [18.0���19.7 (average 19.0) vs 12.6���17.4 (14.8)]. Enteromius mimus differs from E. alberti by a lower number of lateral line scales [24���27 (median 25) vs 27���34 (31)], a larger pre-anal distance [68.4���74.8% SL (average 72.4) vs 65.1��� 73.0 (69.3)], a smaller post-anal distance [16.3���19.3% SL (average 17.8) vs 16.3���24.2 (21.1)], a larger body depth [26.5���34.3% SL (average 30.1) vs 21.7���31.2 (26.9)], a larger head depth [17.8���21.8% SL (average 20.2) vs 15.1���19.0 (17.3)] and a larger pelvic fin length [18.2���22.4% SL (average 20.4) vs 15.0���20.9 (18.0)] (Table 2). Specimens from E. mimus differ from the specimens of the population of E. cf. mimus from the Lake Edward system by a lower number of scales between the origin of the dorsal fin and the lateral line [3.5���4.5 (median 3.5) vs 3.5���4.5 (4.5)], a larger pelvic fin length [18.2���22.4% SL (average 20.4) vs 14.4���21.3 (18.5)], a smaller snout length [4.1���7.1% SL (average 6.0) vs 4.6���8.3 (7.0)], a smaller interorbital width [6.3���8.3% SL (average 7.8) vs 6.9���9.9 (8.7)], a larger post-dorsal distance [34.8���40.4% SL (average 38.1) vs 31.0���39.9 (35.9)] and a larger anal fin length [18.0���19.7% SL (average 19.0) vs 14.4���21.8 (17.7)] (Table 2). Etymology The specific epithet ��� mimus ��� means ���imitator���, a reference to the ���striking resemblance��� to E. neglectus (Boulenger 1912; Scharpf & Lazara 2019). Material examined Lectotype (here designated) KENYA ��� Euasso Nyiro, bellow falls, British EastAfrica; BMNH1912.3.22.99 (Supplementary file SM.01: Fiche-185). Paralectotypes KENYA ��� 8 specs [of 9 in lot]; same collection data as for lectotype; BMNH 1912.3.22.90 to 1912.3.22.98 (Supplementary file SM.01: Fiche-184, Fiche-186 to Fiche-188, Fiche-B to Fiche-E). Description The redescription of E. mimus is based on the specimens of the type series only (the lectotype and eight paralectotypes).The lectotype is illustrated in Fig. 8. Maximum observed size: 45.6 mm SL. Morphometric and meristic data are given in Table 2. Body fusiform, largest depth anterior to dorsal fin. Dorsal profile from tip of snout to origin of dorsal fin slightly convex, slightly tapering to base of caudal fin. Ventral profile from operculum to origin of pelvic fin slightly convex, slightly tapering to posterior end of anal fin base, then slightly concave to caudal fin. Head small. Eye large and round, located laterally, closer to tip of snout than distal margin of operculum, interorbital profile slightly convex. Snout rounded. Mouth subterminal. Two pairs of barbels, anterior barbel shorter than posterior with anterior one usually reaching up to vertical through the middle of the eye, while posterior one can reach the vertical trough posterior margin of pre-operculum. Dorsal fin with 3 unbranched and 7 to 8 branched rays, distal margin slightly concave, origin located on vertical through pelvic fin insertion or slightly anterior (1 lateral line scale). First branched dorsal fin ray longest, posterior rays decreasing progressively in size. Pectoral fin with 1 unbranched and 12 to 14 branched rays, distal profile straight to slightly convex, not reaching anterior base of pelvic fin. Pelvic fin with 1 unbranched and 7 branched rays, distal margin convex. Anal fin with 3 unbranched and 5 to 6 branched fin rays, distal margin concave. Caudal fin forked with outer rays twice as long as median ones and both lobes rounded and of similar size. Anus and urogenital opening situated immediately in front of anal fin base. Scales cycloid rounded and radially striate. Lateral line completely pored with many lateral line scales (total) [24���27 (median 25)], gently curved downwards over abdomen but running straight along middle of caudal peduncle and ending at base of caudal fin. Colour pattern No sexual dimorphism is observed. No information of in vivo colour pattern is available. In preserved specimens, overall background colour of body brown, darker dorsally. One to three dark spots on flanks on thick silvery midlateral band; spots fused into mid-lateral line in some specimens, overlying silvery band and starting posterior to operculum. First spot situated anterior to dorsal fin origin, median one below or somewhat anterior to last ray of dorsal fin and last one at caudal fin base. Distribution Enteromius mimus is known from its type locality, the Euasso Nyiro River below the falls, east of Lake Baringo, Kenya, and from the Tana River system (Seegers et al. 2003). For the time being, the specimens of the Lake Edward system are referred to as E. cf. mimus and are not included in the distribution of E. mimus., Published as part of Maetens, Heleen, Steenberge, Maarten Van, Snoeks, Jos & Decru, Eva, 2020, Revalidation of Enteromius alberti and presence of Enteromius cf. mimus (Cypriniformes: Cyprinidae) in the Lake Edward system, East Africa, pp. 1-28 in European Journal of Taxonomy 700 on pages 18-21, DOI: 10.5852/ejt.2020.700, http://zenodo.org/record/3981069, {"references":["Boulenger G. A. 1912. On a collection of Fishes made by Mr. A. Blayney Percival in British East Africa to the East of Lake Baringo. Proceedings of the Zoological Society of London 82 (3): 672 - 676. https: // doi. org / 10.1111 / j. 1469 - 7998.1912. tb 07548. x","Greenwood P. H. 1962. A revision of certain Barbus (Pisces, Cyprinidae) from East, Central and South Africa. Bulletin of the Natural History Museum, Zoology series 8 (4): 151 - 208. Available from https: // www. biodiversitylibrary. org / part / 118628 / summary [accessed 21 Nov. 2018].","Okaronon J. O., Katunzi E. F. B. & Asila A. A. 1997. Fish Species Identification Guide for Lake Victoria, Enumerator's Field Guide. Lake Victoria Fisheries Research Project Phase II, Technical Document 1, Jinja, Uganda. Available from http: // aquaticcommons. org / 4150 / [accessed 21 Nov. 2018].","Seegers L., De Vos L. & Okeyo D. O. 2003. Annotated checklist of the freshwater fishes of Kenya (excluding the lacustrine haplochromines from Lake Victoria). Journal of East African Natural History 92: 11 - 47. https: // doi. org / 10.2982 / 0012 - 8317 (2003) 92 % 5 B 11: ACOTFF % 5 D 2.0. CO; 2","Schmidt R. C., Bart Jr. H. L. & Nyingi W. D. 2017. Multi-locus phylogeny reveals instances of mitochondrial introgression and unrecognized diversity in Kenyan barbs (Cyprininae: Smiliogastrini). Molecular Phylogenetics and Evolution 111: 35 - 43. https: // doi. org / 10.1016 / j. ympev. 2017.03.015","Froese R. & Pauly D. (eds) 2018. FishBase. Available from www. fishbase. org [accessed 21 Nov. 2018].","Boulenger G. A. 1907. Zoology of Egypt: the Fishes of the Nile. Published for the Egyptian Government by Hugh Rees, London. https: // doi. org / 10.5962 / bhl. title. 51710","Greenwood P. H. 1970. A revision of the cyprinid species Barbus (Enteromius) radiatus Peters, 1853, with a note on the synonymy of the subgenera Beirabarbus and Enteromius. Revue de Zoologie et de Botanique africaines 82: 1 - 13.","Scharpf C. & Lazara K. J. 2019. The ETYFish Project. Available from www. etyfish. org [accessed 17 Sep. 2019]."]}

Enteromius alberti (Poll, 1939) Fig. 7 a���b, Table 2 Enteromius cercops (Whitehead, 1960) (here synonymised). syn. nov. Diagnosis Enteromius alberti (Fig. 7 a���b) belongs to the group of species of Enteromius with a flexible last unbranched dorsal fin ray that lacks serrations along its posterior edge. A comparison was made with other species of Enteromius from this group from the East Coast and Nilo-Sudan ichthyofaunal provinces (list made based on information available in, e.g., Greenwood 1962; Okaronon et al. 1997; Seegers et al. 2003; Schmidt et al. 2017; Froese & Pauly 2018). Enteromius alberti can easily be distinguished from the other species of this group from the East Coast and Nilo-Sudan ichthyofaunal regions by the following combination of characteristics (data from original description, unless otherwise stated): a complete lateral line vs an incomplete lateral line in E. atkinsoni (Bailey, 1969), E. pumilus (Boulenger, 1901), E. serengetiensis (Farm, 2000), E. tongaensis (Rendahl, 1935) and E. toppini (Boulenger, 1916); two pairs of barbels vs one pair in E. pseudotoppini (Seegers, 1996), and no barbels in E. anema (Boulenger, 1903) and E. profundus (Greenwood, 1970); one to three dark spots on the flanks, which sometimes fuse into a mid-lateral line in preserved specimens, starting posterior to the operculum vs a dark line running from the tip of the snout to the caudal fin base in E. bifrenatus (Fowler, 1935) and E. yongei (Whitehead, 1960), and a thin dark line from the beginning of the operculum to the caudal fin base in E. viviparus (Weber, 1897); 12 scales around the caudal peduncle, with one aberrant specimen in the paratypes of E. cercops (16), vs 8 in E. leonensis (Boulenger, 1915), 9���10 in E. venustus (Bailey, 1980), and 10 in E. magdalenae (Boulenger, 1906) and E. yeiensis (Johnsen, 1926); 4.5 scales between the dorsal fin base and the lateral line vs 3.5 in E. radiatus (Peters, 1853) (Greenwood 1970), 5.5 in E. unitaeniatus (G��nther, 1867), and 6 in E. usambarae (L��nnberg, 1907); a dorsal fin length which is larger than the head length vs a dorsal fin length equal to the head length in E. innocens (Pfeffer, 1896); a body depth which is larger than the head length vs a body depth which is equal to the head length in E. nigeriensis (Boulenger, 1903) and E. trispilopleura (Boulenger, 1902); a pectoral fin length which is ��������� of the head length vs �� in E. lineomaculatus (Boulenger, 1903), and �� to �� in E. neglectus (Boulenger, 1903) (Boulenger 1907); a maximum caudal peduncle depth which is �������� of the head length vs ����� ��� in E. quadripunctatus (Pfeffer, 1896). Enteromius alberti differs from E. perince by a combination of a smaller body depth [21.7���31.2% SL (mean 26.9) vs 33.8���37.5 (35.0)], a smaller minimum caudal peduncle depth [10.0���13.6% SL (mean 12.0) vs 15.0���16.4 (15.5)], and a smaller maximum caudal peduncle depth [11.8���15.5% SL (mean 13.6) vs 17.7���19.4 (18.8)]. Enteromius alberti differs from E. stigmatopygus by a combination of a higher number of lateral line scales [27���34 (median 31) vs 20���25 (21)], a smaller pre-dorsal distance [45.7���51.3% SL (mean 49.0) vs 51.9���55.8 (53.8)] and a smaller pre-occipital distance [16.6-21.8% SL (mean 18.6) vs 22.1���27.3 (24.0)]. Enteromius alberti differs from E. mimus by a higher number of lateral line scales [27���34 (median 31) vs 24���27 (25)], a smaller pre-anal distance [65.1���73.0% SL (average 69.3) vs 68.4���74.8 (72.4)], a larger post-anal distance [16.3���24.2% SL (average 21.1) vs 16.3���19.3 (17.8)], a smaller body depth [21.7���31.2% SL (average 26.9) vs 26.5���34.3 (30.1)], a smaller head depth [15.1���19.0% SL (average 17.3) vs 17.8���21.8 (20.2)] and a smaller pelvic fin length [15.0���20.9% SL (average 18.0) vs 18.2���22.4 (20.4)] (Table 2). Specimens of E. alberti differ from the population of E. cf. mimus from the Lake Edward system by a smaller head depth. In general, a specimen with a head depth smaller than 19% of the standard length can be assigned to E. alberti (only one specimen had a higher value). Specimens with a head depth larger than 19% of the standard length can be assigned to E. cf. mimus (only one specimen had a lower value). Other characters to distinguish E. alberti from E. cf. mimus are the higher number of lateral line scales [27���34 (median 31) vs 21���31 (26)], the smaller interorbital width [5.8���8.3% SL (mean 7.3) vs 6.9���9.9 (8.7)], the smaller pre-pelvic distance [44.8���53.3% SL (mean 49,2) vs 48.4���55.4 (52.1)], the smaller body depth [21.7���31.2% SL (mean 26.9) vs 23.5���35.8 (31.0)], the smaller maximum [11.8��� 15.5% SL (mean 13.6) vs 8.7���19.0 (16)] and minimum [10.0���13.6% SL (mean 12) vs 7.1���15.9 (13.7)] caudal peduncle depth, and the smaller head width [10.4���14.2% SL (mean 12) vs 11.6���14.5 (13.2)] (Table 2, Fig. 3). Etymology Etymology not explained in the original description. Probably, the species epithet alberti is derived from the name of the former ���Parc National Albert��� (now Virunga National Park) in which the type locality is located. Material examined Lectotype (here designated) DEMOCRATIC REPUBLIC OF THE CONGO ��� Rutshuru River, May-Ya-Moto; 5���11 Nov. 1934; de Witte leg.; MRAC 64723 (Supplementary file SM.01: Fiche-209). Paralectotype DEMOCRATIC REPUBLIC OF THE CONGO ��� 1 spec.; same collection data as for lectotype; MRAC 64722 (Supplementary file SM.01: Fiche-210). Other material examined KENYA ��� Lake Victoria basin ��� 1 spec., holotype of E. cercops; Luambwa, Nzoia River, Nyanza Province; 1960; BMNH 1960.6.7.37 ��� 7 specs [of 13 in lot], paratypes of E. cercops; same collection data as for holotype; BMNH 1960.6.7.38 to 1960.6.7.50 (Supplementary file SM.01: Fiche-177 to Fiche-183). UGANDA ��� Lake Edward ��� 1 spec. [of 2 in lot]; Nchwera River, Rwenshama-Ishasha road; 0��27��30.8�� S, 29��48��07.0�� E; 8 Nov. 2016; HIPE exped.; GenBank: MT251114; RMCA 2016.035.P.0148 to 2016.035.P.0149 (Supplementary file SM.01: E31).��� 1 spec. [of 2 in lot]; same collection data as for preceding; RMCA 2016.035.P.0171 to 2016.035.P.0172 (Supplementary file SM.01: HP1028).��� 1 spec.; Kayanja, offshore, Lake Edward; 0��05��31.2�� S, 29��45��30.3�� E; 20 Jan. 2018; HIPE exped.; GenBank: MT251133; RMCA 2018.008.P.0247 (Supplementary file SM.01: HP2599).��� 2 specs.; Nyamweru River, mouth, Kisenji; 0��18���42.5��� S, 29��51���36.0��� E; 31 Jan. 2018; HIPE exped.; RBINS 25585 to 25586 (Supplementary file SM.01: Fiche-118, Fiche-119) ��� Lake George ��� 5 specs; River 2 km east of Kitongore; 0��05��45.6�� S, 30��40��51.6�� E; 31 Oct. 2016; HIPE exped.; GenBank: MT251105, MT251120, MT251121, MT251106, MT251107; RMCA 2016.035.P.0159 to 2016.035.P.0163 (Supplementary file SM.01: HP592, HP595, HP596, HP600, HP602) ��� 2 specs; Mpanga River, mouth, Lake George; 0��02��20.4�� N, 30��17��24.0�� E; 31 Oct. 2016; HIPE exped.; GenBank: MT251119, MT251104; RMCA 2016.035.P.0164 to 2016.035.P.0165 (Supplementary file SM.01: HP575, HP579) ��� 1 spec.; Mahoma River, Rwimi-Nyabani road; 0��21��00.2�� N, 30��16��41.0�� E; 1 Apr. 2017; HIPE exped.; GenBank: MT251110; RMCA 2017.006.P.0244 (Supplementary file SM.01: HP1878) ��� 1 spec.; Lake George, Kashaka Bay, south of inlet; 0��05��04.6�� S, 30��10��45.6�� E; 2 Feb. 2018; HIPE exped.; GenBank: MT251111; RMCA 2018.008.P.0248 (Supplementary file SM.01: HP3246) ��� Kazinga Channel ��� 4 specs; Kyambura River, mouth, Kazinga Channel; 0��07��25.8�� S, 30��03��07.5�� E; 30 Oct. 2016; HIPE exped.; GenBank: MT251100, MT251101, MT251102, MT251103; RMCA 2016.035.P.0155 to 2016.035.P.0158 (Supplementary file SM.01: HP562(2), HP563 to HP565) ��� 7 specs; same locality; 4 Feb. 2018; HIPE exped.; RBINS 25588 to 25594 (Supplementary file SM.01: Fiche-143 to Fiche-149) ��� 4 specs; same collection data as for preceding; GenBank: MT251099, MT251112; RMCA 2018.008.P.0249 to 2018.008.P.0252 (Supplementary file SM.01: HP3312, HP3314, HP3317, HP3319) ��� 1 spec.; Kyambura River, middle course, north of Lake Katinda; 0��12��23.7�� S, 30��06��24.3�� E; 2 Feb. 2018; HIPE exped.; RBINS 25587 (Supplementary file SM.01: Fiche-128) ��� 5 specs; Kazinga Channel, near Queen Elisabeth Bush Lodge; 0��08��09.6�� S, 30��02��27.6�� E; 4 Nov. 2016; HIPE exped.; GenBank: MT251123, MT251124, MT251125, MT251108, MT251109; RMCA 2016.035.P.0166 to 2016.035.P.0170 (Supplementary file SM.01: HP719 to HP721, HP723, HP724) ��� 2 specs; Kazinga Channel, opposite of mouth Kyambura River; 0��07��17.5�� S, 30��02��59.0�� E; 4 Feb. 2018; HIPE exped.; RBINS 25595 to 25596 (Supplementary file SM.01: Fiche-121, Fiche-123). Description Based on the lectotype and one paralectotype of E. alberti, the type series of E. cercops, which is synonymised here as E. alberti, and 37 additional specimens from the Lake Edward system, identified as E. alberti. The lectotype is illustrated in Fig. 7a. Maximum observed size: 74.3 mm SL. Morphometric and meristic data are given in Table 2. Body fusiform, largest depth anterior to dorsal fin. Dorsal profile from tip of snout to origin of dorsal fin slightly convex, slightly tapering to base of caudal fin. Ventral profile from operculum to origin of pelvic fin slightly convex, slightly tapering to posterior end of anal fin base, then slightly concave to caudal fin. Head small. Eye large and round, located dorso-laterally, closer to tip of snout than to distal margin of operculum, interorbital profile slightly convex. Snout rounded and mouth subterminal. Two pairs of barbels, anterior barbel usually slightly shorter than posterior with anterior one usually reaching up to vertical through posterior margin of eye, while posterior one usually up to vertical trough posterior margin of pre-operculum. Dorsal fin with 3 unbranched and 7 to 9 branched rays, distal margin straight to slightly concave, origin located slightly anterior [1 or 2 lateral line scale(s)] to vertical through pelvic fin insertion. First branched dorsal fin ray longest, posterior rays decreasing progressively in size. Pectoral fin with 1 unbranched and 13 to 16 branched rays, distal profile straight to slightly convex, not reaching anterior base of pelvic fin. Pelvic fin with 1 unbranched and 6 to 8 branched rays, distal margin slightly convex. Anal fin with 3 unbranched and 5 to 6 branched fin rays, distal margin slightly concave. Caudal fin forked with outer rays twice as long as median ones and both lobes rounded and of similar size. Anus and urogenital opening situated immediately in front of anal fin base. Scales cycloid, rounded and radially striate. Lateral line completely pored with many lateral line scales [27���34 (median 31)], and gently curved downwards over abdomen but running straight along middle of caudal peduncle and ending at base of caudal fin. Lateral-line scales smaller on caudal peduncle than below dorsal fin. Colour pattern No sexual dimorphism observed. In life, overall background colour of body silvery, darker toward dorsum and lighter towards belly (Fig. 7b). Thick yellowish midlateral band, often with one to three darker spots. All fins translucent. In preserved specimens, overall background colour of body yellowish, greyish dorsally (Fig. 7a). One to three dark spots on flanks on thick silvery midlateral band; spots fused into mid-lateral line in some specimens, overlying silvery band and starting posterior to operculum. First spot situated anterior to dorsal fin origin, median one below last ray of dorsal fin and last one at caudal fin base. Specimens with more than three spots are rare (1 specimen with 4 spots and another with 6 spots). Distribution (Fig. 10) Enteromius alberti occurs in river systems throughout the Lake Edward basin. For the Congolese part of the basin, it is known from its type locality, the Rutshuru River, May-Ya-Moto (Poll 1939). Although the Congolese part of the system was not sampled during the recent expeditions, we identified specimens from the collection at the RMCA (86-01-P-479 to 500 and 86-09-P-70 to 71) from Vitshumbi, located in the southern Congolese part of the Lake Edward system, as E. alberti. In the Ugandan part, we collected the species in the Mahoma, Mpanga, Nchwera, Nyamweru and Kyambura (an affluent of the Rusangwe) Rivers, in Lake Edward offshore at Kayanja, and in Lake George at Kashaka Bay. For the Lake Victoria basin, E. alberti is known from the Nzoia River, the Malawa River and the Middle Akagera system (Whitehead 1960; Greenwood 1966; De Vos & Thys van den Audenaerde 1990; Seegers et al. 2003)., Published as part of Maetens, Heleen, Steenberge, Maarten Van, Snoeks, Jos & Decru, Eva, 2020, Revalidation of Enteromius alberti and presence of Enteromius cf. mimus (Cypriniformes: Cyprinidae) in the Lake Edward system, East Africa, pp. 1-28 in European Journal of Taxonomy 700 on pages 15-18, DOI: 10.5852/ejt.2020.700, http://zenodo.org/record/3981069, {"references":["Whitehead P. J. P. 1960. Three new cyprinid fishes of the genus Barbus from the Lake Victoria basin. Revue de Zoologie et de Botanique africaines 62 (1 - 2): 106 - 119.","Greenwood P. H. 1962. A revision of certain Barbus (Pisces, Cyprinidae) from East, Central and South Africa. Bulletin of the Natural History Museum, Zoology series 8 (4): 151 - 208. Available from https: // www. biodiversitylibrary. org / part / 118628 / summary [accessed 21 Nov. 2018].","Okaronon J. O., Katunzi E. F. B. & Asila A. A. 1997. Fish Species Identification Guide for Lake Victoria, Enumerator's Field Guide. Lake Victoria Fisheries Research Project Phase II, Technical Document 1, Jinja, Uganda. Available from http: // aquaticcommons. org / 4150 / [accessed 21 Nov. 2018].","Seegers L., De Vos L. & Okeyo D. O. 2003. Annotated checklist of the freshwater fishes of Kenya (excluding the lacustrine haplochromines from Lake Victoria). Journal of East African Natural History 92: 11 - 47. https: // doi. org / 10.2982 / 0012 - 8317 (2003) 92 % 5 B 11: ACOTFF % 5 D 2.0. CO; 2","Schmidt R. C., Bart Jr. H. L. & Nyingi W. D. 2017. Multi-locus phylogeny reveals instances of mitochondrial introgression and unrecognized diversity in Kenyan barbs (Cyprininae: Smiliogastrini). Molecular Phylogenetics and Evolution 111: 35 - 43. https: // doi. org / 10.1016 / j. ympev. 2017.03.015","Froese R. & Pauly D. (eds) 2018. FishBase. Available from www. fishbase. org [accessed 21 Nov. 2018].","Boulenger G. A. 1916. Catalogue of the Fresh-Water Fishes of Africa in the British museum (Natural History), Vol. IV. British Museum (Natural History), London, UK. https: // doi. org / 10.5962 / bhl. title. 8869","Boulenger G. A. 1903. Diagnosis of four new species of Barbus from the Nile. Annals and Magazine of Natural History, Series 7 12: 532 - 533. https: // doi. org / 10.1080 / 00222930308678890","Greenwood P. H. 1970. A revision of the cyprinid species Barbus (Enteromius) radiatus Peters, 1853, with a note on the synonymy of the subgenera Beirabarbus and Enteromius. Revue de Zoologie et de Botanique africaines 82: 1 - 13.","Boulenger G. A. 1907. Zoology of Egypt: the Fishes of the Nile. Published for the Egyptian Government by Hugh Rees, London. https: // doi. org / 10.5962 / bhl. title. 51710","Greenwood P. H. 1966. The Fishes of Uganda. The Uganda Society, Kampala.","De Vos L. & Thys van den Audenaerde D. F. E. 1990. Petit Barbus (Pisces, Cyprinidae) du Rwanda. Revue d'Hydrobiologie tropicale 23 (2): 141 - 159."]}

Density plot of DAPC. Densities of individuals on the first discriminant function of the DAPC shown in Fig. 4. (PDF 28 kb)

Individual-based latent fixed mixed model (LFMM) analysis. Distribution of adjusted p-values, corrected with the genomic inflation factor. Made with the LEA package in R. (PDF 4 kb)

Sequencing quality and information on missing data per individual. Table shows the sampling site, individual ID, number of SNPs, number of missing SNPs, frequency of missing SNPs, mean read depth and number of raw reads per individual. (PDF 370 kb)

Outlier analysis based on BayeScan v2.1. A. FST-Log10 posterior probability for two levels (sampling site and subbasin) for each of the three replicates (R1, R2, R3). B. Q value for grouping according sampling site and subbasin with each of the three replicates (R1, R2, R3). (PDF 169 kb)

AFLP-based Neighbour Joining tree of 108 southern specimens of Tropheus. Three T. duboisi specimens were used as outgroup, statistical support was estimated by performing 1000 bootstrap replicates. (PDF 8 kb)

Morphological variation vs. geographical distance along the shoreline for the individual meristics and measurements. Measurements were expressed as percentages of head (for measurements taken on the head) or standard length, raw data were shown for meristics and for SL; the distance was taken along the shoreline (in km) starting from the northwesternmost locality. LOESS curves were calculated with a fixed smoothening parameter of 0.2. As this resulted in a straight line for ASp, 0.1 was used instead (*). (PDF 4487 kb)

Output of cluster analysis performed in Structure. Groups are indicated as they occur along the shoreline with 1: T. sp. ‘maculatus’, 2: T. sp. ‘red’, 3: T. moorii ‘yellow’, 4: T. moorii ‘South’, 5: T. moorii ‘Southeast’, 6: T. brichardi ‘Kipili’, 7: T. sp. ‘Mpimbwe’. The northernmost specimen of T. sp. ‘red’ (Kikoti, loc. 4) is represented by the first sample of group 2, even though it appears within the distribution range of group 1. (PDF 317 kb)

Tables (.docx with: 1) list of material and sampling sites, 2) morphological parameters studied, 3) results of the PCA on the merictic and 4) on the measurement data. (DOCX 50 kb)

Estimates of delta K and L (K) given by Structure. (PDF 2935 kb)