aqua International Journal of

Vol. 16 (3), 15 July 2010

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A new species of , Haplochromis katonga n. sp. (: Cichlidae) from the Katonga River, Uganda

Erwin Schraml1 and Herbert Tichy2

1) Haferstrasse 18c, D-86179 Augsburg, Germany. E-mail: [email protected] 2) Max-Planck-Institut für Biologie, D72076 Tübingen, Germany. E-mail: [email protected]

Received: 04 February 2010 – Accepted: 29 March 2010

Abstract LS, collectés dans la rivière Katonga, Uganda. Elle diffère A new species of the Haplochromis, H. katonga, is des autres espèces du genre par sa taille relativement described from nine specimen, ranging from 37.1 to 68.7 réduite, la combinaison de caractéristiques (comme l’œil mm SL, collected from the Katonga River, Uganda. It dif- relativement grand, un nombre réduit d’écailles sur la linge fers from other species of the genus by its relatively small latérale centrale (<31 vs. >30), une hauteur de corps moins size, the combination of morphological characters (such as prononcée que pour les autres espèces fluviatiles (32,7 vs. the relatively large eye, a low number of scales in the mid- 35-40 % LS chez Astatotilapia), et par sa coloration (les lateral line (<31 vs. >30), a smaller body depth than in mâles: bleuâtre à gris-brun foncé sur le dos, flancs other fluviatile species (32.7 vs. 35-40 % SL in Astatoti- verdâtres, parties ventrales jaunâtres, avec des zones lapia), and its coloration (in males: bluish to dark grey- rougeâtres dans les nageoires impaires). C’est la seule brown on dorsum, flanks greenish, ventral parts yellowish, espèce d’Haplochromis connue de la rivière Katonga qui with reddish areas in unpaired fins). It is the only known relie les bassins du lac Victoria et du lac George. Elle mon- Haplochromis species from the Katonga River, which con- tre des affinités avec les genres Astatotilapia et Ente- nects the Lake Victoria and Lake George basins. It shows rochromis (comme définis par Greenwood 1979, 1980). affinities to the genera Astatotilapia and Enterochromis (as defined by Greenwood 1979, 1980). Sommario Una nuova specie del genere Haplochromis, H. katonga, è Zusammenfassung descritta sulla base di nove esemplari, di dimenzioni variabili Eine neue Art der Gattung Haplochromis: H. katonga, wird dai 37.1 ai 68.7 mm SL, raccolti nel fiume Katonga, nach neun Exemplaren mit 37,1 bis 68,7 mm SL Uganda. Essa differisce dalle altre specie del genere per la sue beschrieben, die im Katonga-Fluss in Uganda gefangen wur- dimensioni relativamenente ridotte, per una combinazione den. Von den anderen Arten der Gattung unterscheiden sie di caratteri morfologici, quali l’occhio relativamente grande, sich durch relativ geringe Größe, die Kombination be- un numero ridotto di scaglie sulla linea laterale mediana stimmter morphologischer Merkmale (beispielsweise ein rel- (<31 vs. >30), una minore altezza del corpo rispetto alle altre ativ großes Auge, eine geringe Zahl von Schuppen auf der specie fluviatili (32.7 vs. 35-40 % SL in Astatotilapia) e per seitlichen Mittellinie (<31 vs. >30), eine geringere Körper- la colorazione dei maschi che appare da bluastra a grigio- tiefe als bei anderen flusslebenden Arten (32,7 vs. 35-40 % marrone scuro sul dorso, con fianchi verdastri e parti ven- der Standardlänge bei Astatotilapia) sowie durch die Farbge- trali giallastre e con aree rossastre sulle pinne impari. è la sola bung (bei Männchen: bläulich bis dunkel graubraun am specie di Haplochromis rinvenuta nel Katonga, un fiume che Rücken, grünlich die Flanken, im Bauchbereich gelblich, congiunge i bacini del lago Victoria e del lago George. rötliche Bereiche an den unpaaren Flossen). Es handelt sich Mostra somiglianze con i generi Astatotilapia ed Ente- um die bisher einzige Haplochromis-Art vom Katonga-Fluss, rochromis (come definiti da Greenwood 1979, 1980). der den Victoriasee mit dem Georgsee verbindet. Verbindende Merkmale ließen sich zu den Gattungen Asta- INTRODUCTION totilapia und Enterochromis (nach den Definitionen von Greenwood 1979, 1980) feststellen. Describing new Haplochromis species is a difficult undertaking. Genera are poorly defined, phyloge- Résumé netic relationships are poorly known and there are Une nouvelle espèce du genre Haplochromis, H. katonga, hundreds of closely related species, with a very est décrit sur base de neuf spécimens, de 37.1 à 68,7 mm large number of undescribed species. Many of

81 aqua vol. 16 no. 3 - 15 July 2010 A new species of Haplochromis, Haplochromis katonga n. sp. (Perciformes: Cichlidae) from the Katonga River, Uganda them seem to have originated in the recent past, with Haplochromis katonga n. sp. the formation of hybrid swarms partially generating (Figs 1-10; Table I) the species flocks of the Lake Victoria region. Many species are extremely difficult to identify through Holotype: ♂, 68.7 mm SL, ZSM 30362 [8683] autapomorphies other than coloration. Morphologi- (Figs 2, 4), Katonga River at the village of Kaba go - cal differences between the species are small and le, Uganda, (0°11’53.74”N; 30°53’33.98”E); H. intraspecific variability is relatively high. Nevertheless Tichy & E. Schraml, 26 October 1998. the of Lake Victoria and adjacent Paratypes: ZSM 30363 (Figs 3, 4), [8676], ♀, lakes have been regarded as of special interest since 40.0 mm SL; [8677], ♀, 37.1 mm SL; [8678], ♂, Greenwood (1964, 1969, 1981) introduced them as 47.7 mm SL; [8679], ♀, 38.1 mm SL; [8680], ♀, an example of rapid evolution of a species flock. 42.7 mm SL; [8681], ♀, 37.1 mm SL; [8684], ♀, Greenwood (1973) stated that the headwater of the 40.9 mm SL; [8685], ♀, 48.8 mm SL; 25/26 river Katonga, which connects Lake Victoria with October 1998, same data as for the holotype. Lake George, is apparently impenetrable to all but Diagnosis: A relatively small species with a mod- air-breathing fishes. Today the Katonga and Mpanga erate body depth, laterally compressed, with gener- rivers occupy the riverbed of the ancient Katonga alised body shape (for genus and family), a termi- which originally drained the Lake Victoria area west- nal, slightly acute, symmetric mouth, lower jaw wards (Said 1993). The drifting apart of the Nubian slightly protruding, and relatively large eyes (33.8 and Somalian tectonic plates is most probably % HL vs (i. e.) 23.7 in H. nubilus, 29.4 in H. responsible for the unstable geology of the interven- akika, 30.5 in H. commutabilis). It shares with ing area across which this river formerly flowed other East African fluviatile species, especially the (Stamps et al. 2008). Geological movements of these so-called Astatotilapia, the smaller longitudinal plates finally split it into the modern river Katonga, scale count (<31) compared to lacustrine species which thereafter flowed eastwards into Lake Victoria, (>30), with exceptions H. akika Lippitsch, 2003, with the Mpanga continuing to flow westwards to H. ampullarostratus Schraml, 2004, H. nubilus Lake George. Both rivers have a common source in a (Boulenger, 1906), from which (except H. huge swampy area that forms the watershed between ampullarostratus) it is distinguished by its lesser the two rivers (Doornkamp & Temple 1966) (Fig. body depth (<35 vs. >35 % SL). From H. 1). The Katonga river resembles a swamp, to a large ampullarostratus it differs in, for instance, greater extent choked by papyrus and other vegetation. The head length (34.7-37.5 vs. 32.5-33.5 % SL) and authors collected a species of the genus Haplo chro mis, shorter snout length (21.5-28.7 vs. 29.1-31.4 % which will be described hereinafter as H. ka ton ga, at HL). Additionally it differs from H. nubilus in Kabagole, a village approximately half-way between head shape (dorsal head profile straight vs con- lakes Victoria and George. cave), and nuptial coloration (solid black in H. nubilus vs colourful: see description below), and MATERIAL AND METHODS colour of anal fin which is plain red in nuptial- General terminology, counts and measurements coloured males of H. katonga n. sp. vs. only mar- follow Barel et al. (1977); for terminology regard- ginally reddish or different-coloured in all other ing scale features see Lippitsch (1990, 1993). Mea- fluviatile species. The overall nuptial coloration of surements were taken with dial callipers, up to 0.5 males is also a diagnostic feature: live coloration of mm for body measurements and up to 0.1 mm for males bluish to dark grey-brown on dorsum, flanks head measurements. Head measurements were greenish, ventral parts yellowish, with reddish areas taken through a binocular microscope. in unpaired fins; of females greyish blue to greyish Institutional abbreviations are: ZSM (Zoologische yellow with a faint metallic blue hue dorsally. Staatssammlung München, Abteilung Vertebrata, Description: The following description is based Sektion Ichthyologie, München, Germany), on nine individuals, the holotype (adult) and eight BMNH (Natural History Museum, London, U.K.). paratypes, all adult and sub-adult specimens rang- Numbers such as [8683] represent collection ing from 37.1 to 68.7 mm SL. numbers in the private collection of Herbert Tichy, Morphology. Relatively small species with gener- in which specimens were stored before transfer to alised body shape. Body moderately deep, com- ZSM, and were used in primer publications, as pressed laterally. Head moderately large with rela- well as for GenBank. tively large eye and small mouth. Cheek depth aqua vol. 16 no. 3 - 15 July 2010 82 Erwin Schraml and Herbert Tichy comparatively small, eye depth greater than cheek premaxillary pedicel not prominent. Maxillary depth. Snout pointed, isognathous (lower jaw posterior extension reaching vertical line through slightly protruding), upper and lower profiles iris. Lips not thickened, maxilla not bullate. Lower equally angled. Dorsal head profile almost straight, (horizontal) border of preoperculum convex,

Fig. 1. Type locality of Haplochromis katonga. The short yellow arrows indicate the approximate watershed and direction of river flow. (Picture from JPL-SRL2/SIR-C).

Fig. 2. Haplochromis katonga n. sp., male, holotype ZSM 30362, photographed shortly after capture. Photo by E. Schraml.

83 aqua vol. 16 no. 3 - 15 July 2010 A new species of Haplochromis, Haplochromis katonga n. sp. (Perciformes: Cichlidae) from the Katonga River, Uganda rounded, curvature extending upward to maxilla; extending past vertical line through base of caudal upper (vertical) border extending downward to fin, but not in females, where tips hardly reach that meet this curvature. Eye almost circular, relatively line. Caudal fin subtruncate. large, but not reaching dorsal outline of head. Scales. partially scaled. Suboperculum Pupil slightly drop-shaped, with perimeter tapering fully covered with cycloid scales; interoperculum caudally. Caudal peduncle longer than deep. fully covered by one row of cycloid scales; cheeks Fins. Pectoral insertion located on imaginary line fully scaled; postorbital squamation with two rows running posteriorly downwards from base of dorsal of different-sized scales; size of occipital scales not fin to base of ventral at angle of about 10° from significantly smaller than dorsal scales; predorsal vertical; pectoral not extending to vertical line squamation pattern irregular; cycloid and weakly through origin of anal fin; ventrals extending to ctenoid scales on dorsum rostrally, more strongly this line. In adult males tips of dorsal and anal fins ctenoid scales increasing caudally; flanks with

Fig. 3. Haplochromis katonga n. sp, holotype, preserved (ZSM 30362). Photo by E. Schraml.

Fig. 4. Haplochromis katonga n. sp, female, paratype ZSM 30363, photographed shortly after capture. Photo by E. Schraml. aqua vol. 16 no. 3 - 15 July 2010 84 Erwin Schraml and Herbert Tichy

Table I. Morphometrics of nine specimens of Haplochromis katonga n. sp. ranging from 37.1 to 68.7 mm SL.

Haplochromis katonga Holotype Paratypes (n=8) mean Standard length (SL) in mm 68.7 37.1-48.8 Percentages of standard length Head length (HL) 34.2 34.7-37.5 35.9 Body depth 34.1 30.4-34.6 32.7 Caudal peduncle length 15.6 14.8-18.0 16.1 Caudal peduncle depth 11.4 10.2-11.7 10.9 Anal fin base length 22.9 20.4-24.2 22.1 base length 56.5 51.4-57.4 54.3 Predorsal length 37.0 34.3-38.9 36.0 Pre-anal length 67.1 64.9-67.1 65.8 Prepectoral length 36.6 35.7-40.3 38.5 Prepelvic length 42.2 40.4-45.2 42.6

Percentages of head length Head width 47.2 44.4-50.0 47.2 Snout length 30.2 21.5-28.7 25.9 Snout width 32.3 24.7-32.4 28.7 Interorbital width 29.8 27.7-34.5 30.0 Preorbital depth 13.6 7.4-14.9 11.7 Eye length 31.1 33.1-36.5 33.8 Eye depth 29.9 30.5-34.4 32.5 Cheek depth 23.0 16.3-24.7 19.2 Lower jaw length 40.4 34.0-39.9 36.7 Lower jaw width 18.7 16.3-24.1 19.4 Premaxillary pedicel length 28.1 18.9-26.3 24.5

Ratios Eye depth/length 1.0 0.9-1.0 1.0 Lower jaw length/width 2.2 1.5-2.3 1.9

Counts Lateral line scales 20+10 20+10; 20+9; 19+12; 19+11; 18+10; 18+7; not usable (2) Lateral line - dorsal fin (scale rows) 4 4 (8) Pectoral - bases (scale rows) 4 4 (6), 3 (2) Cheek (vertical scale rows) 4 4 (7), 3 (1) Dorsal fin XVI/8 XVI/9 (2), XVI/8 (1), XV/10 (1), XV/9 (3), XV/8 (1) Anal fin III/8 III/10 (1), III/9 (6), III/8 (1) Number of teeth (upper jaw/lower jaw) 43/20 33/25, 33/21, 33/20, 31/21, 30/26, 30/20, 29/22, 29/20.

85 aqua vol. 16 no. 3 - 15 July 2010 A new species of Haplochromis, Haplochromis katonga n. sp. (Perciformes: Cichlidae) from the Katonga River, Uganda strongly ctenoid scales (Fig. 6); granulation of margins short or even extremely short, but well flank scales covering area reminiscent in shape of a developed at middle axis; first interradial circuli Viking axe, with relatively slight broadening hori- concave or straight, but with convex ridges extend- zontally and vertically at focus, but with extent ing onto tongue (Fig. 7); scales immediately larger than 150° at caudal margin; only caudal part behind pectoral origin cycloid, but becoming more of focus, for short distance to absolute centre of strongly ctenoid posteriorly; scales on caudal scale, strongly covered with granulae, close to cen- peduncle strongly ctenoid; scales on chest laterally tre having shape of tubercles, partly conjoined, and ventrally strongly ctenoid; large interpelvic closer to margin with shape reminiscent of short scale present; scales on belly weakly ctenoid, about peanut-shells, and then gradually becoming longer 1/3 smaller than flank scales and imbricating; towards margin and finally intergrading into large, scales on anal-genital region of similar size to flank conical denticles, only weakly curved or hooked; scales, weakly to strongly ctenoid; scales on caudal remaining part of focus of flank scale either cov- cycloid, arranged in single row between and along ered only weakly with slightly thickened tubercles rays; two scale rows between upper and lower lat- along circular structure or covered completely with eral line, rostrally and caudally; four scale rows grains (see Fig. 6); radii simple, except for last radii between upper lateral line and base of dorsal; four all of primary type; tongues on upper and lower scale rows between bases of pectoral and ventral

a b

c d

e f

Fig. 5. Paratypes of Haplochromis katonga n. sp. in ZSM 30363: a. [8676], ♀, 40.0 mm SL; b. [8678], ♂, 47.7 mm SL; c. [8679], ♀, 38.1 mm SL; d. [8680], ♀, 42.7 mm SL; e. [8681], ♀, 37.1 mm SL; f. [8684], ♀, 40.9 mm SL). Photos by E. Schraml. aqua vol. 16 no. 3 - 15 July 2010 86 Erwin Schraml and Herbert Tichy

Fig. 6. Two lateral scales from the holotype, removed on the right side from the second and third rows beneath the upper right-hand lateral line below the anterior part of the dorsal fin. Photos by E. Schraml.

Fig. 7. Radii (left) and ctenii (right) o