aqua, International Journal of Ichthyology

Nothobranchius nubaensis (: ) a new annual from Sudan and Ethiopia

Stefano Valdesalici 1, Marc Bellemans 2, Kiril Kardashev 3, Alexander Golubtsov 4

1) Via Cà Bertacchi 5, 42030 Viano (RE), Italy. E-mail: [email protected] and [email protected] 2) Mortselsesteenweg 138, 2540 Hove, Belgium 3) Stramna 4, 2600 Dupnitsa, Bulgaria 4) Laboratory of Lower Vertebrate Ecology, A.N. Severtsov Institute of Ecology & Evolution, Russian Academy of Sciences, Leninskii Pr. 33, Moscow119071, Russia

Received: 19 March 2009 – Accepted: 04 July 2009

Abstract Résumé A new of annual killifish, nubaensis , Une nouvelle espèce de killy annuel, Nothobranchius is described based on specimens collected from ephemeral nubaensis, est décrite sur base de spécimens collectés dans water bodies in central Sudan and south -west Ethiopia. The des poches d’eau temporaires du centre du Soudan et du new species is distinguished from the other members of the sud-ouest de l’Ethiopie. La nouvelle espèce se distingue des Nothobranchius ugandensis species group by the following autres membres du groupe Nothobranchius ugandensis par combination of characters : 17-19 dorsal fin rays; 17-19 anal la combinaison des caractéristiques suivantes: 17 à 19 fin rays; 29-30 scales in median lateral series; dorsal and anal rayons dorsaux; 17 à 19 rayons à l’anale, 29 à 30 écailles fins in males with short filamentous rays; pelvic fins short, dans les rangées latérales médianes; dorsale et anale tips reaching the anus; orange red head with three distinct des mâles avec de courts rayons filamenteux; pelviennes oblique bars on the operculum ; dorsal fin light blue with a courtes, les extrémités touchant l’anus; tête rouge orange pattern of irregular orange -red spots and elongated yellow avec trois barres obliques nettes sur l’opercule; dorsale spots in distal areas; anal fin yellow, light blue at base, with a bleu clair avec un patron irrégulier de taches rouge-oran- pattern of orange -red spots; pelvic fins yellow with a pattern ge; pelviennes jaunes avec un patron de taches rouge of orange -red spots; pectoral fins orange -red with a light blue orange; pectorales rouge orange avec un liseré bleu clair; margin; caudal fin orange -red with short pale or dark red caudale rouge orange avec de petites lignes rouges pâles ou lines , extending from fin base onto fin rays. foncées s’étendant de la base de la nageoire jusqu’aux rayons. Zusammenfassung Eine neue Art der einjährigen Killifische: Nothobranchius Sommario nubaensi, wird auf der Basis von Exemplaren beschrieben, Una nuova specie di killifish annuale, Nothobranchius die in kurzzeitigen Wasserlöchern im Zentralsudan und in nubaensis, è descritta sulla base di esemplari raccolti in corpi SW-Äthiopien gesammelt wurden. Sie unterscheidet sich idrici temporanei del Sudan centrale e dell’Etiopia sudocci - von anderen Angehörigen der Nothobranchius-ugandensis- dentale. La nuova specie si distingue dagli altri membri del Artengruppe durch die folgende Merkmale: 17-19 Rück - complesso di specie Nothobranchius ugandensis per la se- enflossenstrahlen; 17-19 Afterflossenstrahlen; 29-30 guente combinazione di caratteri: 17-19 raggi dorsali; 17-19 Schuppen in median-lateralen Reihen; die dorsalen und raggi anali; 29-30 scaglie lungo la serie mediana laterale; analen Flossen bei Männchen mit kurzen fadenförmigen pinna dorsale e anale nei maschi con brevi raggi filamentosi; Flossenstrahlen; Bauchflossen kurz, die Spitzen reichen bis pinne pelviche brevi, con le punte che arrivano fino all’ano; zum After; orangeroter Kopf mit drei deutlichen schrägen capo rosso-arancio con tre distinte barre oblique sull’oper - Streifen auf dem Kiemendeckel; Rückenflosse hellblau mit colo; pinna dorsale blu chiaro con un motivo di macchioline einem unregelmäßigen Muster orangeroter Flecken und irregolari rosso-arancio e macchie allungate gialle nelle aree länglichen gelben Flecken im distalen Bereich; Afterflosse distali; pinna anale gialla, blu chiaro alla base, con un gelb, hellblau am Grund, mit einem Muster orangeroter motivo di macchioline rosso-arancio; pinne pelviche gialle Flecken; Brustflossen gelb mit einem Muster orangeroter con un motivo di macchioline rosso-arancio; pinne pettorali Flecken; Brustflossen orangerot mit hellblauem Rand; rosso-arancio con un margine blu chiaro; pinna caudale Schwanzflosse orangerot mit kurzen blassen oder dunkel - rosso-arancio con brevi linee pallide o rosso scuro che si roten Linien, die sich von der Flossenbasis bis zu den estendono dalla base della pinna fino ai raggi. Flossenstrahlen erstrecken.

143 aqua vol. 15 no. 3 - 20 July 2009 Nothobranchius nubaensis (Cyprinodontiformes: Nothobranchiidae) a new annual killifish from Sudan and Ethiopia

INTRODUCTION peduncle length , which is measured from the poste - The freshwater fauna of Sudan is poorly rior end of the anal fin base to the posterior end of known. It was studied by Sandon (1950a, 1950b, the hypural plate ; eye diameter , which is measured 1957) and more recently by Bailey & Hickley between anterior and posterior orbital walls ; and (1986 ), Hickley & Bailey (1987) , Lévêque et al. snout length , which is measured from the anterior (1991), and Bailey (1994). The freshwater fish fauna tip of the lower jaw to the anterior edge of the of Ethiopia is better known (Tedla 1973, Tedla & orbital wall. Measurements were taken with digital Meskel 1981, Dgebuadze et al. 1994, Golubtsov et callipers to the nearest 0.1 mm, partly under a dis - al. 1995, de Graaf et al. 2000). secting microscope. All measurements and counts Nothobranchius virgatus Chambers, 1984 is the were taken from the left side. All visible rays of the only species of Nothobranchius , which has been dorsal and anal fins were counted. The counts of described from Sudan. It was collected in October scales on the median longitudinal series is the num - 2008. From Ethiopia it is known from Gambella, ber of scales between the superior junction of the Obela Stream , near the road from Gambela to Gog opercular membrane and the hypural plate . The (J. Friel , pers. comm.). scales on the base of the caudal fin were counted sep - On 5 December 2002, live specimens of a red arately. All measurements are presented as percent - species of Nothobranchius were collected by M. Belle - ages of standard length (SL); except of eye diameter mans in Khor Angarko (which is part of the Khor and snout length, which are given as percentage of Abu Habl system ), South Kordofan , about 21 km head length (HL). Terminology for the cephalic south of Dilling and 100 km north of Kadugli (Belle - neuromast series follows Scheel (1968) , the frontal mans 2003). This species was provisionally described squamation following Hoedeman (1958). as N. nubaensis (Bellemans 2003), but this name is Type specimens and additional material is not available, for the following reasons: (1) the men - deposited in the following institutions:, Musée Royal tion of a “provisional description ” may questionably de l’Afrique Centrale , Tervuren, Belgium (MRAC ); be considered to violate ICZN Article 16.1; (2) no Natural History Museum, London, UK (BMNH ); type has been designated in the paper, the author Muséum National d’Histoire Naturelle, Paris, France rather states “types will be designated …”. Article (MNHN ); Zoological Museum, Moscow State Uni - 16.4.1 of the ICZN clearly requires an “explicit fixa - versity, Moscow, Russia (ZMM ); South African tion of a holotype, or syntypes ”, even if not immedi - Institute for Aquatic Biodiversity, Grahamstown, ately deposited. The paper mentions several speci - South Africa (SAIAB ); and, Museo Civico di Storia mens without giving numbers. The name N. nubaen - Naturale “G. Doria ”, Genova, Italy (MSNG ). sis is not a “nomen nudum ” as a description is given, Comparative material is listed in Blache & Miton but an “anoplonym ” according to Dubois (2000). (1960), Chambers (1984, Rahad el Fula population), In November 2005 and November 2006, K. Kar - and Wildekamp (1995) . Further collections exami - dashev and others collected specimens of red nated include: Stefano Valdesalici private collection: coloured Nothobranchius in Khor Maada Mulaha , Nothobranchius robustus , 4 males, 31,7-36,3 mm SL, belonging to the Wadi Al Ghallah system (Karda - Uganda, Kalisizo 0°33’S 31°36’E ; N. rubroreticula - shev 2007 a , b). In November 1986, November tus , 2 males, 34,3-36,9 mm SL, Tchad, Zakouma 1994 and December 2006, A. Golubtsov collected National Park 10°50’N 19°38’E ; N. ugandensis , 1 specimens of red coloured Nothobranchius near male, 47,3 mm SL, Uganda, Butiaba 01 °53’N Abobo , Ethiopia. They occured in seasonal ponds of 31°24’E; 1 male, 43,7 mm SL , Uganda, Busesa the Alvero River drainage , White Nile basin. Speci - 00°30’N 33°31 E. mens of these three populations from Sudan and Ethiopia are considered to belong to the same species on the basis of the coloration of males and Nothobranchius nubaensis n. sp. morphological characters . Here below this annual (Figs 1-6, Table I) fish is described as a new species , Nothobranchius nubaensis . Nothobranchius nubaensis Bellemans, 2003 (anoplonym ) MATERIAL AND METHODS Measurements and counts were taken according to Holotype: MRAC 2008-05-P-1, male, 42,9 mm Amiet (1987) ; except for the addition of the caudal SL; Sudan: Angarko, 21 km south of Dilling and aqua vol. 15 no. 3 - 20 July 2009 144 Stefano Valdesalici , Marc Bellemans , Kiril Kardashev , Alexander Golubtsov

100 km north of Kadugli, 11°53’ 05.23”N base, with a pattern of orange -red spots; pelvic fins 29°42’31.82”E, Marc Bellemans, 5 December yellow with a pattern of orange -red spots; pectoral 2002. Collected with hand net and preserved after fins orange -red with a light blue margin; caudal fin living for about six months in aquaria . orange -red , with short pale or dark red lines extend - Paratypes (collected with holotype ): MRAC 2008- ing from fin base onto fin rays. 05-P-2, 1 male, 43,9 mm SL ; MRAC 2008-05-P-3- Description: Morphometric measurements are 4, 2 females, 28,1-33,2 mm SL ; MSNG 54286, 1 given in Table I. Robust, deep bodied Notho - male, 40,8 mm SL ; MNHN 2008-1140–1141- branchius with rounded snout ; terminal mouth 1179, 3 males, 38,4-45,8 mm SL ; SAIAB 80410, 1 directed upward; dorsal fin rays 17-19; anal fin rays male, 36,5 mm SL ; BMNH 2008.4.2.11-12, 1 17-19; scales in median lateral series 29-30 + 2-3 on male, 45,1 mm SL, 1 female, 33,1 mm SL . caudal fin base; cephalic squamation pattern vari - None-type material: SAIAB 80411, 1 male, able; anterior neuromast series of ‘open’ type; central 49 mm SL, Sudan: Maada Mulaha, 11°09.405’N series in shallow groove with low lobes and with 3 40° 19.441’ E, Kiril Kardashev and Plamen Ivanov, neuromasts; posterior cephalic neuromast series 22 November 2005 , collected with large hand nets curved with 3 neuromasts; preopercular neuromast and preserved in the field ; SAIAB 80412, 1 female, system in open groove, distal ridge overlaping oper - 40,8 mm SL , BMNH 2008.4.2.13-14, 1 male, cle slightly; one neuromast on each scale of median 47,1 mm SL, 1 female, 39,8 mm SL, same data as longitudinal series. SAIAB 80411 ; ZMMGU P22009, 3 males, 38,4- Maximum observed length 49.9 mm SL in males; 49,9 mm SL , Ethiopia: Alvero River drainage, 7 km body deep and moderately compressed ; dorsal pro - S of the southern end of the Alvero Dam, 7° 47’N file slightly concave to nearly straight on head, con - 34° 25’E, altitude about 500 m, Alexander Gol - vex from nape to end of dorsal fin base ; ventral pro - ubtsov, 4 December 2006 , collected with large nets file convex, slightly concave on caudal peduncle pos - and preserved in the field. terior to dorsal and anal fins; maximum body height Diagnosis: Nothobranchius nubaensis can be distin - occurring at the base of the pelvic fins; dorsal and guished from all other species of the by the anal fins shallow and rounded, tips with short fila - following combination of characters: 17-19 dorsal mentous rays; dorsal and anal fins with papillate fin rays; 17-19 anal fin rays; 29-30 scales in median contact organs , distributed over fin rays; pelvic fins lateral series; dorsal and anal fin with short filamen - shorts, tips reaching the anus; caudal fin subtrun - tous rays in males; pelvic fins short, tips reaching the cate ; branchiostegal membrane projecting from anus; orange -red head with 3 distinct oblique bars opercle, distal edge slightly wrinkled. on the operculum ; dorsal fin light blue with a pat - Female smaller than male, maximum observed size tern of irregular orange -red spots and elongated yel - 40.8 mm SL; body less compressed and less deep low spots in distal areas; anal fin yellow, light blue at than in males; dorsal fin rounded; anal fin triangu -

Fig. 1. Adult male of Nothobranchius nubaensis (not preserved) from Angarko. Photo by S. Valdesalici.

145 aqua vol. 15 no. 3 - 20 July 2009 Nothobranchius nubaensis (Cyprinodontiformes: Nothobranchiidae) a new annual killifish from Sudan and Ethiopia

Table I. Morphometric measurements of Nothobranchius nubaensis (including type series and additional specimens).

Holotype Males (n=12) Females (n=5) SL (mm) 42.9 36.5-49.9 33.1-40.8 Depth at the pelvic fins 31.4 25.9-34.0 21.4-30.4 Preanal length 60.3 56.5-65.7 58.7-68.8 Predorsal length In percents of standard length (SL) 56.8 54.4-61.8 57.8-61.4 Prepelvic length 47.5 44.0-54.9 49.7-56.0 Caudal peduncle lenght 20.2 18.0-22.0 14.9-22.3 Caudal peduncle depth 12.8 11.7-15.4 9.3-12.9 Head length 29.3 26.3-33.6 25.6-33.9 Snouth length 23.8 21.6-32.3 20.7-26.8 In percents of head length (HL) Eye diameter 23.8 20.7-29.8 23.7-29.1 lar with rounded tip; anal fin positioned more pos - Female (Fig. 4): body pale olive-brown, lighter terior ; short snout and narrower caudal peduncle brown to silvery ventrally; unpaired and paired fins than in males; opercular membrane not projecting hyaline; iris golden, with faint black vertical bar from opercle. through center of eye. Color in life : Male (Figs 1-3): body and head Color in alcohol: Male (Fig. 5): body scales scales light blue with broad orange -red margin, cre - light brown to whitish, almost all scales with distinct ating a reticulated pattern on body ; almost com - red margin; dorsal and anal fins light brown to pletely orange -red on head; lips, snout, throat, whitish with a pattern red spots; caudal fin of a light frontal and superior portion of head all orange -red; pale red; pelvic fins light brown to whitish with red opercular region with three distinct oblique bars on spots near base; pectoral fins of a light pale red; iris the operculum, extending from the eye to the upper bluish. part of the head; branchiostegal membrane orange - Female (Fig. 6): body light brown to whitish; oper - red with light blue rim; dorsal fin light blue with cular and ventral area yellowish; unpaired and paired broad dark red reticulation, forming approximately fins whitish; iris bluish. horizontal arch-like stripes, broad border with inter - Distribution (Fig. 7) : Populations have been found rupted red pattern; some elongated yellow spots in scattered over a large area. Two groups of localities distal areas; anal fin yellow, light blue at base, with can be distinguished: (1) Khor Angarko (in South pattern of large orange -red spots ; spots in the mid - Kordofan around the foothills of the Nuba Moun - dle of the fin dark red, distal spots orange -red; cau - tains) and the area near Khor Maada Mulaha (in the dal fin orange -red with pale or dark red lines extend - plains of West Kordofan) in Sudan ; (2) near Abobo ing from base onto fin rays , some specimens having (Gambella Region) in Ethiopia. a faint dark red marginal band; pelvic fins yellow Habitat (Figs 8-10) : The specimens of Notho - with orange -red spots, spots near base dark red; pec - branchius nubaensis described here were found in toral fins orange -red with light blue margin, some residual pools of seasonal rivers (called “Khor”) and specimens with yellow spots and short dark or pale rainwater filled depressions (Bellemans 2003, Kar - red lines extending onto fin rays; iris golden, with dashev 2007 a , b). All populations of Notho - faint black vertical bar through center of eye. branchius in central Sudan and south -west Ethiopia undergo important annual fluctuations in abun - dance and extent of distribution area. In Khor Angarko , N. nubaensis was the only fish species pre - sent in 2002. During successive surveys at the same collection site during the post-rain period in 2005 and 2006 no specimens of N. nubaensis were found, not even in the mud collected from the residual pools (M. B.). Fig. 2. Adult male of Nothobranchius nubaensis (not pre - In Khor Maada Mulaha Nothobranchius virgatus served) from Khor Maada Mulaha. Photo by K. Kardashev. and N. aff. rubroreticulatus were found in in the aqua vol. 15 no. 3 - 20 July 2009 146 Stefano Valdesalici , Marc Bellemans , Kiril Kardashev , Alexander Golubtsov same locality in 1998 and 1999 (Bellemans 2000) there are two seasonal ponds near Abobo where and in 2004 (J. Bulterman pers. comm.). In 2005 Nothobranchius nubaensis is known to occur, both and 2006 N. virgatus , N. nubaensis and Protopterus belonging to the Alvero river drainage (White Nile sp. (probably Protopterus annectens Owen, 1939) basin). were found in the same locality, but not a single In late November 1994 , two males and three specimen of N. aff . rubroreticulatus . In this locality females of Nothobranchius were collected from a all males Nothobranchius nubaensis were collected at swamp-like pond at the land divide between the the surface near rocks. Almost all females were col - Alvero and Gilo rivers (the latter river seems to be a lected in about 2 to 2.5 m deep water. No aquatic tributary of the Pibor River, affluent of Sobat, afflu - vegetation was present, with only dry grass along the ent of White Nile). The pond was muddy and cov - shores. The water was brown, slightly turbid, with a ered with dense macrophyte vegetation. The Pro - pH of 7.8- 8.2. At 12:30 h the temperature was topterus sp. larvae were sampled simultaneously with 22°C, at 1 m below the water surface. The air tem - the Nothobranchius specimens. Based on the anec - perature was 42°C. The bottom substrate of these dotal information from the local Anuak people, the habitats was brown-black clay (K.K.). In Ethiopia , pond is never connected to any river system; it is

Fig. 3. Freshly collected adult male of Nothobranchius nubaensis, ZMMGU P22009, from Akobo. Photo by A. Golubtsov.

Fig. 4. Adult female of Nothobranchius nubaensis (not preserved) from Angarko. Photo by S. Valdesalici.

147 aqua vol. 15 no. 3 - 20 July 2009 Nothobranchius nubaensis (Cyprinodontiformes: Nothobranchiidae) a new annual killifish from Sudan and Ethiopia filled with rain water. The same pond was re-sam - elephants (judging from foot prints). The pond was pled on 4 December 2006. No Protopterus were never re-sampled until present (A. G.). found. New villages and cultivated lands appeared in Etymology: The new species is named after type the vicinity of the pond, which became a source of locality, which lies in the Nuba mountains. drinking water for the nearest settlement located Discussion: Based on the known distribution, on about 2.5 km to the north . the coloration of males and on morphology, Notho - In the other pond, in late November 1986, about branchius nubaensis can be grouped in the Notho - 25 km W of Abobo (now 15 km W of a dam) branchius ugandensis species group, which is defined 7° 54’ N 34° 20’ E, altitude about 430 m, only one here . This clade includes N. ugandensis Wildekamp, female of Nothobranchius was sampled from the 1995, an undescribed species of Nothobranchius pan-like pond in the flood plain of the Alvero River ”” (see comments in Wildekamp 1990: (a tributary of the Baro River, affluent of Sobat, 199, 1996: 147, and pictures in Seegers 1997: 82- affluent of White Nile). The pond had a light clay 83) and N. nubaensis , which is described here. bed and no dense macrophyte vegetation. Judging Synapomorphies to define this group are: male col - from the fish species composition (occurring oration consisting of light blue scales with a large together with the only Nothobranchius specimen), irregularly reticulated pattern on body , forming i.e. small Barbus sp ., Ctenopoma sp ., Neolebias sp ., oblique bars in vivid red color, vivid red coloration Aplocheilichthys (? , possibly Micropanchax ), the pond on the head and in dorsal areas, yellow to yellowish was connected to the river during the rainy season. anal fin, large vivid red spot pattern on dorsal and When we tried to re-sample the pond in early Janu - anal fins, red caudal fin of subtruncated shape; rela - ary 1987, no fish was found there; the water tem - tively deep body (up to 38% of SL) ; moderately perature was about 43°C; it was a watering place of compressed, and rounded head ; dorsal profile of

Fig. 5. Nothobranchius nubaensis , MRAC 2008-05-P-1, holotype, 42 .9 mm SL, male. Photo by S. Valdesalici.

Fig. 6. Nothobranchius nubaensis , MRAC 2008-05-P-3, paratype, 33.2 mm SL, female. Photo by S. Valdesalici. aqua vol. 15 no. 3 - 20 July 2009 148 Stefano Valdesalici , Marc Bellemans , Kiril Kardashev , Alexander Golubtsov head slightly concave to nearly straight, convex from spots pattern (broad and arranged in horizontal row nape to end of dorsal fin base ; variable cephalic vs. small spots , sometime arranged in thin lines squamation. extending onto fin rays) ; anal fin spot pattern (broad Male N. nubaensis differ from a similarly red male vs. absent or if present, few and arranged in lines morph of N. ugandensis by coloration of the whole extending to fin rays ) ; pectoral fin color (red vs. hya - body (orange -red broad scales margin vs. narrow red line). Males and females with relatively a reduced scale margins) and head (almost completely orange - number of scales in the longitudinal series (29-30 vs. red vs. red on lips and superior part only ); dorsal fin 28-33), modally less deep body in males (25.9-34. 0

Fig. 7. Geographical distribution of Nothobranchius nubaensis (asterisk; type locality: open asterisk ), Nothobranchius virgatus (circle) and N. aff . rubroreticulatus (orange squares) (black squares are cities with their names). Drawing by S. Valdesalici.

149 aqua vol. 15 no. 3 - 20 July 2009 Nothobranchius nubaensis (Cyprinodontiformes: Nothobranchiidae) a new annual killifish from Sudan and Ethiopia vs. 25.0-37.3% of SL) and females (21.4-30. 4 vs. line) ; caudal fin coloration (red vs . light blue with a 22.4-34. 0% SL) ; dorsal fin positioned more anterior black margin followed by a submarginal white band in males (54 .4-61.8 vs. 54.5-65.2% of SL) and and a faint black vertical bar in the inner region ); females (57 .8-61.4 vs. 57.0-66.5% of SL) ; origin of and females color pattern (hyaline without particu - pelvic fins relatively more anterior in males (44 .0- lar pattern vs. with faint oblique bars) ; more dorsal - 54.9 vs. 47.2-57.0% of SL) ; shorter head in males fin rays in males and in females (17-19 vs. 13-16); (26 .3-33.6 vs. 26.6-36.8% of SL) and females (25 .6- more anal -fin rays in males and females (17-19 vs. 33.9 vs. 27.6-37.8% of SL) ; body scales (without 14-16) ; modally less deep caudal peduncle in males ctenii vs. with ctenii), ctenii structures on operculum (11 .7-15.4 vs. 14.3-16. 4% SL) ; dorsal fin positioned (absent vs. present). more anterior in males (54 .4-61.8 vs. 59.0-62.8% of Among other species of Nothobranchius occurring SL) ; shorter head in males (26 .3-33.6 vs. 31.4- in the area, male N. nubaensis differ from male N. 35.7% of SL) ; pelvic fin shorter (tips reaching the virgatus (found in the same habitat in Khor Maada anus vs. tips reaching anal fin). Mulaha) by the coloration pattern of the whole Male N. nubaensis differ from male N. robustus by body (broad orange -red scale margins vs. thin red coloration of the whole body (light blue with broad scale margins , forming about 10-16 oblique bars) orange -red scale margins vs. completely red to red - and head (almost completely orange red vs. light brown) ; dorsal and anal fin pattern (spotted vs. plain blue with narrow red scale margins) ; anal fin spot red) ; pectoral fin coloration (red vs. hyaline) ; males pattern and coloration (yellow, light blue at base, and females with more dorsal -fin rays (17-19 vs. 14- with pattern of large orange -red spots, spots in the 16); shorter head in males (26 .3-33.,6 vs. 29.2- middle of fin dark red, distal spots orange -red vs. 38.8% of SL); different shape of head and snout light blue with red spots forming 6-8 curved lines (rounded vs. pointed). and a broad margin) ; pectoral fin color (red vs. hya - Male N nubaensis differ from male N. rubroreticu - latus by coloration of the whole body (broad orange - red scale margins vs. red broad scale margins form - ing a reticulated irregular pattern) and head (almost completely orange -red vs. light blue with narrow red scale margins); dorsal fin spot pattern (broad and arranged in horizontal rows vs. spots near the fin base , arranged in an irregular horizontal line, spots in the median part arranged in broad lines extend - ing onto fin rays, light blue submarginal band and broad black margin); anal fin coloration and spot pattern (yellow, light blue at base, with pattern of large orange -red spots, spots in the median part dark red, distal spots orange red vs. light blue with a series of red spots near the base forming a broad irregular horizontal line, spots in the median part arranged in broad red lines extending onto fin rays, light blue submarginal band and broad black margin) ; pectoral fin color (red vs. hyaline) ; caudal fin coloration (red vs. light blue with a pattern of broad red lines extending onto fin rays, light blue submarginal band and broad black margin ; deeper body in males (26 .3-34.0 vs. 25.4-29.7% of SL) ; anal fin posi - tioned more posteriorly in males (56 .5-65.7 vs. 55.1-59.3% of SL) ; shorter head in female (25 .6- 33.9 vs. 32.4-37.2% of SL) ; different shape of head and snout (rounded vs. pointed). The present disjunct distribution pattern of N. Fig. 8. Sudan: Angarko; type locality of Nothobranchius nubaensis , similar to that of N. virgatus can be nubaensis . Photo by M. Bellemans. explained by the dynamics of the changing rain pat - aqua vol. 15 no. 3 - 20 July 2009 150 Stefano Valdesalici , Marc Bellemans , Kiril Kardashev , Alexander Golubtsov tern and climatic conditions over the vast plains of in Chambers (1984), Hoelzmann et al. (2000) and central and southern Sudan and SW Ethiopia (see Leblanc et al. (2006). also Lévêque 1990). The higher water levels of the A comparison of the samples did not reveal any Nile in the past and the presence of a fossil embank - apparent differences in morphometry, color pattern ment surrounding a pre-historic lake south of Khar - or meristics among populations. Determining the toum, suggest a more continuous and permanent full extent of the distribution of N. nubaensis will connection between the presently known popula - require additional collections. tions of N. nubaensis (and N. virgatus), as reported

Fig. 9. Sudan: Khor Maada Mulaha. Photo by K. Kardashev.

Fig. 10. Ethiopia: Alvero River drainage, 7 km S of the southern end of the Alvero Dam (pond after intensive sampling). Photo by A. Golubtsov.

151 aqua vol. 15 no. 3 - 20 July 2009 Nothobranchius nubaensis (Cyprinodontiformes: Nothobranchiidae) a new annual killifish from Sudan and Ethiopia

ACKNOWLEDGEMENTS HOEDEMAN , J. 1958. The frontal scalation pattern in some The first author wishes to thank Francesca Fontana, groups of tooth carps (Pisces: Cyprinodontiformes). Daniele Cavazzoni, and Ruud Wildekamp for their Bulletin of Aquatic Biology 1: 23-28. assistance with the first draft on the manuscript; HOELZMANN , P., KRUSE , H.-J. & ROTTINGER , F. 2000. Pre - cipitation estimates of the E-Saharan Palaeomonsoon: sed - Alain Dubois for suggestions regarding nomencla - imentary geochemistry; satellite image remote sensing and ture; Maurice Kottelat for suggesting the etymology; a regional water balance model of the West-Nubia-Lake. Dirk Neumann for help with references; C. E. Global and Planetary Change 26 : 105-120. Cherenkov for the “clipping path” of Ethiopian fish GOLUBTSOV , A. S., DARKOV , A. A., DGEBUADZE , Y U. Y U. & picture; and John Friel for information on N. virga - MINA , M. V. 1995. An artificial key to fish species of the tus in Ethiopia . We thank two anonymous reviewers Gambella region (the White Nile basin in the limits of for their their critical comments on the manuscript. Ethiopia). Joint Ethio-Russian Biological Expedition. We would also like to express our gratitude to Miguël Addis Ababa, Ethiopia, 84 pages. Parrent (MRAC); James Maclaine (BMNH); Patrice KARDASHEV , K. 2007 a. Sudan 2005-2006. AIK Notizie Killi 3: 5-9. Pruvost and Romain Causse (MNHN); Roger Bills, KARDASHEV , K. 2007 b. Catturare Notho nella Terra Dei Paul Skelton and Bernard MacKenzie (SAIAB) and Neri - Sudan 2005. AIK Notizie Killi 5: 6-14. Giuliano Doria (MSNG) for access to collections LEBLANC , M. J., LEDUC , C., STAGNITTI , F., VAN OEVELEN , under their care. P. J., JONES , C., MOFOR , L. A., RAZACK , M. & FAVREAU , G. 2006. Evidence for Megalake Chad, north-central REFERENCES Africa,during the late Quaternary from satellite data. AMIET , J. 1987. Faune du Cameroun. Fauna of Cameroon. 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