Garra dembecha, Stiassny & Getahun, 2007

Stiassny, Melanie L. J. & Getahun, Abebe, 2007, An overview of labeonin relationships and the phylogenetic placement of the Afro-Asian genus Garra Hamilton, 1922 (Teleostei: Cyprinidae), with the description of five new species of Garra from Ethiopia, and a key to all African species, Zoological Journal of the Linnean Society 150 (1), pp. 41-83 : 62-79

GARRA DEMBECHA SP. NOV. ( FIG. 15 ; TABLE 6)

Garra dembecha Getahun, 2000 , nomen nudum, (p. 45 unpublished PhD thesis)

Gobio quadrimaculatus (non-Rüppell, 1836): Boulenger, 1903, Proc. Zool. Soc. Lond., p. 330; Boulenger, 1907, The fishes of the Nile, p. 186; Boulenger, 1909, Catalogue of the freshwater fishes of Africa, Vol. 1, p. 351

Discognathus quadrimaculatus (non-Rüppell, 1836): Boulenger, 1903, Proc. Zool. Soc. Lond., p. 330; Boulenger, 1907, The fishes of the Nile, p. 186; Boulenger, 1909, Catalogue of the freshwater fishes of Africa, Vol. 1, p. 351.

Crossocheilus quadrimaculatus: Boulenger, 1902 ; Ann. Mag. N.H. (7), p. 422; Boulenger, 1907, The fishes of the Nile, p. 186; Boulenger, 1909, Catalogue of the freshwater fishes of Africa, Vol. 1, p. 351

Diagnosis: Distinguished from African congeners by the following combination of features: weakly developed disc; predorsal region partially scaled anterior to dorsal fin; chest, belly, and postpelvic region asquamate; moderately elongate intestine (SL 40.0–56.0% Int.L); large gas bladder (posterior chamber 21.7– 22.4% SL); and vent situated distant from anal fin (vent distance 24.3–43.2, m. 32.0%).

Description: Morphometric and meristic data for holotype and 27 paratypes given in Table 6. Maximum size: 129.7 mm SL (MSNM 2162). Body relatively gracile (depth 16.7–23.0, m. 19.5% SL), greatest body depth located midway between dorsal-fin origin and snout. Head gracile and depressed (depth 50.0–65.2, m. 56.9% HL), without tubercles on snout. Eyes positioned medially on head. Dorsal head profile rises steeply over snout, dorsum of head more-or-less flattened, dorsal body profile weakly convex, almost flat to dorsal-fin origin in most specimens, although in some larger individuals dorsal body profile rises steeply behind head and is smoothly covex to dorsal fin. Disc weakly developed (type A), but often with narrow free posterior edge in large specimens, and usually wider than long. Rostral cap well developed, extending ventrally over upper jaw and with invecked ventral margin. Two pairs of barbels: maxillary barbels usually about same length as rostral pair. Either 37 or 38 scales along lateral line; either five or six predorsal scales anterior to dorsal fin; 3.5 scales from lateral line to origin of pelvic fins; 5.5 scales from lateral line to dorsal-fin origin. Chest, belly, and postpelvic region asquamate. Dorsal-fin rays iv, seven, inserted well in advance of pelvic fin, with a slightly rounded tip, first and second unbranched rays longest, but not extending to last ray; posterior margin straight, posterior rays not extending to pelvic-fin tip. Predorsal length 45.3–43.2% SL. Anal-fin rays iv, five, with subacuminate tip; posterior margin concave. Pectoral rays iv, 12; pectoral length 16.6–22.3% SL, with somewhat pointed tips. Vent distant from anal fin (vent distance 24.3–43.2, m. 32.0%). Intestine moderate in length (SL 40.0–56.0% Int.L). Gas bladder well developed and large (posterior chamber 21.7–22.4% SL).

Colour in preservation: Body brown dorsally, slightly lighter brown below lateral line and ventrally. Trace of midlateral band of variable width often present on flanks and caudal peduncle. Distinct bar spanning depth of caudal peduncle, just anterior to caudal fin. Dorsum of head, snout, and most of opercle: dark brown. Lips, gular region, posterior margin of opercle, and branchiostegal membranes: creamy yellowbrown. Entire cheek often pale creamy brown, with pale coloration extending around entire eye. Characteristically well-developed, small black spot present just behind opercle, in region of first lateral line scale. Dorsal-fin membrane pale cream distally, with strongly pigmented, wide black submarginal band in membrane between rays, and narrow pale cream band proximally. Either four or five elongate black spots extend dorsally between rays in basal membrane of dorsal fin, often reaching into basal third of dorsal fin. Anal fin uniformly pale creamy brown. Caudal fin pale dusky brown, with median rays a slightly darker brown. Pelvic and pectoral fins uniformly pale creamy brown.

Distribution and habitat: Recorded from river basins in Eritrea, Ethiopia, and Kenya. Type locality, Bula River, is in the Abbay drainage basin in northern Ethiopia.

Remarks: G. dembecha includes many specimens previously identified as G. quadrimaculata (non-Rüppell) following the descriptions of Boulenger (1903, 1907, 1909); however, Boulenger’s D. quadrimaculatus differs trenchantly from G. quadrimaculatus ( Rüppell, 1836) . Notable differences include the presence of a weakly developed type-A disc in G. dembecha (vs. a well-developed disc in G. quadrimaculata ) and an asquamate belly and chest (vs. scaled in G. quadrimaculata ).

Etymology: Dembecha , in reference to the town near which the type series was collected.

Material examined – type material: Holotype of G. dembecha, AMNH 228483, 78.8 mm SL, Bula River, crossing southern part of Dembecha, Gojam , Ethiopia; paratypes, AMNH 223693, 27 ex., three ex. C & S, 23.5–82.1 mm SL, Bula River, crossing southern part of Dembecha, Gojam , Ethiopia.

Material examined – non-type material: BMNH 1908.1.20:59–68, ten ex., 46.0– 108.3 mm SL, Lake Zurui, Kenya (misidentified as D. quadrimaculatus ); BMNH 1902.12.13.424–433/392–399, 18 ex., 29.7– 86.4 mm SL, Lake Tana, Ethiopia (misidentified as D. quadrimaculatus ); BMNH 1933.9.30.1–3, three ex., 67.1–77.6 mm SL, small Abbay River, Lake Tsana (misidentified as D. quadrimaculatus ); BMNH 1948.1.14.153–156, four ex., 27.3–31.8 mm SL, Mai Toccor, Eritrea (misidentified as G. quadrimaculatus ); BMNH 1908.1.20.69, one ex., 59.6 mm SL, Akaki River, Ethiopia (misidentified as G. quadrimaculata ); BMNH 1937.4.20.14–21, eight ex., 52.6–85.5 mm SL, Lake Ziway, Ethiopia (misidentified as G. quadrimaculata ); BMNH 1902.12.13.400–409, ten ex., 23.9– 95.2 mm SL, Chaffe-Donsa high plateau, Abyssinia (misidentified as G. quadrimaculata ); BMNH 1903.11.16.13.14, two ex., 45.28–67.3 mm SL, Suksuk River, Lake Ziway, Ethiopia (misidentified as D. quadrimaculatus ); MNHN 85-93-54-7, four ex., 41.9–56.8 mm SL, Lake Tana, Ethiopia (misidentified as G. quadrimaculatus ); MNHN 85-93-54-2, three ex., 36.2–62.1 mm SL, Afrique orientale (misidentified as G. quadrimaculatus ); MNHN 85.93.54.3, seven ex., 22.2–30.6 mm SL, Abyssinia (misidentified as G. quadrimaculatus ); MNHN 85-93-54-6, one ex., 18.9 mm SL, Abyssinia (misidentified as G. quadrimaculatus ); MNHN 85-93-54-1, three ex., 25.5– 35.6 mm SL, Lake Victoria, Nyanza (misidentified as G. quadrimaculatus ); MNHN 85-93-54-5, one ex., 25.6 mm SL, Abyssinia (misidentified as G. quadrimaculatus ); MHNG 1446.99, one ex., 90.9 mm SL, affluent of River Athi, environs of Nairobi (near Masai Lodge), Kenya (misidentified as G. dembeensis ); MHNG 2030.23, one ex., 88.9 mm SL, Kilimanjaro, Tanzania (misidentified as G. dembeensis ); MSNM 2138 (ex 4322), one ex., 69.40 mm SL, Lake Tana, Bahirdar , Ethiopia (misidentified as D. quadrimaculatus ); MSNM 2162 (ex 4314), six ex., 84.7–129.7 mm SL, ‘Fiume Caha’, Gondar, Ethiopia (misidentified as D. quadrimaculatus ); AMNH 223702, four ex., 34.0– 64.1 mm SL, Lah River, south of Finote Selam town, Gojam, Ethiopia (10°50′N, 37°15′E); AMNH 223665, six ex., 44.7–84.6 mm SL, Mesha River, north of Butajira town, Rift Valley, Ethiopia (8°10′N, 38°30′E); AMNH 223689, one ex., 100.5 mm SL, Gedeb River, about 20 km north of Debremarkos town, Gojam, Ethiopia (10°30′N, 37°40′E); AMNH 223700, 12 ex., 55.7– 97.3 mm SL, Kechen River, 30 km south of Finote Selam town, Gojam, Ethiopia (10°45′N, 37°20′E); AMNH 223724, five ex., 112.7–124 mm SL, Gumara River, northern end of Maksegnit town, about 30 km south of Gondar town, Ethiopia (12°40′N, 37°40′E); AMNH 223735, three ex., 54.2–64.0 mm SL, Dirma River, in Kola Diba town, about 80 km south of Gondar town, on the way to Gorgora, Gondar, Ethiopia (12°25′N, 37°20′E); AMNH 223796, 19 ex., 17.9– 81.5 mm SL, Wenchet creek, on Gondar-Bahirdar road, about 5 km south of Wereta village, Ethiopia (11°45′N, 37°30′E); AMNH 223805, nine ex., 18.4– 61.2 mm SL, Idiyemo creek, c. 20 km north of Bahir Dar on Gondar road, Ethiopia (11°40′N, 37°30′E); AMNH 223699, eight ex., 58.7–95.1 mm SL, Kechen river, about 30 km south of Finote Selam town, Gojam, Ethiopia (10°45′N, 37°20′E); AMNH 223745, seven ex., 55.4–69.7 mm SL, Elala River, crossing northern part of Makale town, Tigrai, Ethiopia (13°30′N, 39°30′E); AMNH 223677, 33 ex., 21.9–32.8 mm SL, Angordgood creek, 4 km north-east of Bahirdar town, Gojam, Ethiopia (11°35′N, 37°30′E); AMNH 223833, 76 ex., 31.0– 91.2 mm SL, Kela River, 1 km east of Welenkomi town, on the way from Addis Ababa to Nekempte, Shoa, Ethiopia (9°5′N, 38°10′E); AMNH 227303, five ex., 40.9–88.6 mm SL, Arer River, c. 30 km north-east of Jima town, on the road to Addis Ababa, Kefa, Ethiopia (7°50′N, 37°10′E); AMNH 227261, 42 ex., 40.2–101.6 mm SL, Dabena River, 2 km south-west of Bedele town, Illubabor, Ethiopia (8°20′N, 36°15′E); AMNH 223694, two ex., 63.1– 84.6 mm SL, Chereka River, c. 10 km north of Dembecha town, Gojam, Ethiopia (10°40′N, 37°25′E); AMNH 223773, one ex., 42.38 mm SL, Worka River, Wondo Genet, Small stream near the hot spring, Shoa, Ethiopia (7°10′N, 38°40′E); AMNH 223702, four ex., 33.9–64.1 mm SL, Lah River, south of Finote Selam town, Gojam, Ethiopia (10°50′N, 37°15′E); AMNH 223704, 14 ex., 40.7–102.0 mm SL, Kakist River, near Tilili town, c. 50 km north of Finote Selam town, Gojam, Ethiopia (10°55′N, 37°5′E); AMNH 227694, three ex., 90.9–140.2 mm SL, Lake Tana, Gerima , c. 25 meters off shore (11°35′N, 37°24′E); AMNH 236097, three ex., 96.4–105.7 mm SL, Bula River, crossing southern part of Dembecha town, Gojam, Ethiopia.

GARRA QUADRIMACULATA ( RÜPPELL, 1836)

( FIG. 16 A, B; TABLE 7)

Gobio quadrimaculatus Rüppell, 1836 , Mus . Senckenberg. Abhandl. Beschr. Naturg. Vol. 2, p. 22, Plate 3 (original description, type locality: Ethiopia)

Gobio hirticeps Rüppell, 1836 , Mus . Senckenberg. Abhandl. Beschr. Naturg. Vol. 2, p. 23, Plates 3 and 4 (original description, type locality: Ethiopia)

Garra tibanica Trewavas, 1941 , British Museum (Natural History) expedition to South-west Arabia 1937–38, 3. Freshwater fishes, Vol. 1, p. 8. (original description, type locality: Pool at Usaifira, 1 mile north of Ta’izz, Yemen)

Garra brittoni Trewavas, 1941 , British Museum (Natural History) expedition to South-west Arabia 1937– 38, 3. Freshwater fishes, Vol. 1, p. 11. (original description, type locality: Migyal al Alaf, in a cistern 8 miles south of San’a, Yemen)

Diagnosis: Distinguished from African congeners by the following combination of features: well developed papillate disc; fully scaled predorsum, chest, belly, and postpelvic regions; vent positioned very close to anal fin (vent distance 8.6–19.1, mean 12.3%); and long intestine (SL 20.9–24.2% Int.L).

Description: Morphometric and meristic data presented in Table 7. Maximum size: 100.8 mm SL (BMNH 1940.8.15:192, type of Garra britoni ). Body gracile (depth 16.0–20.7, m. 18.4% SL), greatest body depth located at level of pelvic-fin tips. Head somewhat depressed, with characteristic tuberculation consisting of large rounded tubercles concentrated in rows around nostrils. Eyes positioned medially on head. Dorsal head profile rises steeply over snout; dorsum of head more-or-less contiguous with dorsal body profile and smoothly convex to dorsal-fin origin. Disc well-developed (type C), with a wide free posterior margin, and abundant papillae on lower lip and around disc periphery. Disc length 22.8–26.8% HL, wider than long (width 34.4–47.3% HL). Rostral cap well developed, extending ventrally over upper jaw and with invecked ventral margin. Two pairs of barbels: maxillary barbels usually slightly longer than rostral pair. Between 33 and 35 scales along lateral line; predorsal region fully scaled; 3.5 scales from lateral line to origin of pelvic fin; 3.5–4 scales from lateral line to dorsal-fin origin. Belly, chest, and postpelvic regions fully scaled; scales often small and deeply embedded. Dorsal-fin rays iii, seven, inserted well anterior to pelvic fin, with subacuminate tip, first and second unbranched rays longest, extending slightly beyond last ray; posterior margin concave, posterior rays extending almost to pelvic-fin tip. Predorsal length 47.4–49.7% SL. Anal-fin rays iii, five, blunt tip; posterior margin straight. Pectoral rays iv, 11; pectoral length 19.0–19.6% SL, with pointed tips. Vent located very close to anal fin (vent distance 8.6–19.1, mean 12.3%). Intestine long (SL 20.9–24.2% Int.L). Gas bladder well-developed (posterior chamber 19.0– 30.0% SL).

Distribution: Found mainly on the Arabian Peninsula ( Saudi Arabia and Yemen) but present also in southeastern Eritrea, south-eastern Ethiopia, and possibly also Somalia. The Somalian specimens mentioned by Menon (1964) cannot be located, and we have been unable to confirm the presence of G. quadrimaculata in that country.

Remarks: Considerable confusion exists in the literature regarding the identity of G. quadrimaculata , and much of this dates to Boulenger’s (1903) comment that Discognathus (i.e. Garra ) quadrimaculatus have ‘a small mental disc, feebly marked and a little longer than broad’. Apparently Boulenger had not examined Rüppell’s type specimen and instead based his remarks on Rüppell’s abbreviated description, and on specimens he received from Ethiopia (and erroneously assumed to be D. quadrimaculatus Rüppell ). It appears that most subsequent workers followed Boulenger’s description of D. quadrimaculata as having a weakly developed disc, and this has resulted in numerous misidentifications of this species in museum holdings.

Based on examination of type materials, two Arabian species, Garra ghorensis Krupp, 1982 and Garra sahilia Krupp, 1983 appear very similar to G. quadrimaculata , but study of additional material is necessary before their synonomy can be justified.

Material examined – type material: Holotype of G. quadrimaculatus, SMF 5396, 28.9 mm SL, Abyssinia ; Paralectotype of G. hirticeps, SMF 5403, 39.9 mm SL, Abyssinia ; Syntypes of G. tibanica, BMNH 1940.2.15:2–11, ten ex., 70.7–97.9 mm SL, Pond at Usaifira, one mile north of Taizza, Yemen ; Type of G. brittoni, BMNH 1940.8.15: 192, 100.8 mm SL, Migyal al Alaf, cistern 8 miles south of Sana, Yemen .

GARRA AETHIOPICA ( PELLEGRIN, 1927)

( FIG. 16C, D; TABLE 8)

Discognathus aethiopicus Pellegrin, 1927 , Bull. Soc. Zool. Fr., Vol. 52, p. 232 [original description, type locality: Ethiopia, Kabana River , tributary of Aouche River (Awash River basin)]

Garra aethiopicus Menon, 1964 , Memoires of the Indian Museum, Vol. 14, p. 193 (as synonym of G. quadrimaculata ; should have been mentioned as Discognathus aethiopicus ), Daget et al., 1984, CLOFFA, Vol. 1, p. 304 (as synonym of G. quadrimaculata ; should have been mentioned as D. aethiopicus )

Diagnosis: Distinguished from African congeners by the following combination of features: intermediate (type B) disc; scaled predorsal, belly, and postpelvic regions; either smooth or weakly invecked ventral margin to rostral cap; and pale ash-grey markings in both fresh and preserved specimens.

Description: Morphometric and meristic data presented in Table 8. Maximum size: 114.3 mm SL (AMNH 223672). Most individuals are relatively robust (depth 18.9–25.2, m. 22.2% SL); greatest body depth well in advance of dorsal-fin origin, at level of pectoral-fin tips. Head somewhat depressed, no tubercles on snout. Eyes positioned anteriorly on head. Dorsal head profile rises steeply over snout, then more-or-less smoothly convex and contiguous with dorsal body profile to dorsal-fin origin. Disc of intermediate development (type B), with narrow free posterior and lateral margins but usually not enlarged and without abundant papillation around posterior edge. Rostral cap well developed, extending ventrally over upper jaw. Ventral margin either smooth or only weakly invecked, and often incurved so that margin is obscured. Two pairs of barbels of similar length. Between 33 and 36 scales along lateral line; predorsal region fully scaled to dorsal fin; 3.5 scales from lateral line to origin of pelvic fin; 4–4.5 scales from lateral line to dorsal-fin origin. Entire dorsum, belly, and postpelvic regions scaled, although scales often deeply embedded; chest asquamate. Dorsal-fin rays iii–iv, seven, inserted well anterior to pelvic fin, with subacuminate tip, first and second unbranched rays longest, extending beyond last ray; posterior margin weakly concave, posterior rays not extending to pelvic-fin tip. Predorsal length: 43.3–51.1% SL. Anal-fin rays iii–iv, five, with subacuminate tip; posterior margin concave. Pectoral rays iv, 11; pectoral length 17.9– 23.0% SL, with somewhat pointed tips. Vent medially located (vent distance 19.2–29.5, m. 25.9%). Intestine of medium length (SL 47.0–76.0% Int.L). Gas bladder well-developed (posterior chamber 18.3– 21.1% SL).

Colour in preservation: Well-defined dark streak over dorsum of head and body, and broad dark ash-grey midlateral band to base of caudal peduncle usually present. Grey midlateral band somewhat expanded and wedge-shaped at caudal base. Head and body between dorsal streak and midlateral band pale creamy brownish-grey, body uniformly pale ventrally. Well-defined, small, black spot present just behind opercle, in region of first lateral line scale. Lips, gular region, posterior margin of opercle, and branchiostegal membranes: creamy yellow-brown. Dorsal fin more-or-less uniformly dusky creamy-yellow, with no obvious dark submarginal band. Either four or five small black spots between rays in basal membrane of dorsal fin. Anal fin uniformly pale creamy yellow. Caudal fin pale dusky brown, with median rays a darker brown. Pelvic and pectoral fins uniformly pale creamy yellow.

Distribution: Currently known only from the Awash River basin of Ethiopia.

Material examined – type material: Lectotype, MNHN 1927-125 , 88.4 mm SL, Kabana River , tributary of Awash , in Addis Ababa city at an altitude of 2400 m, Ethiopia; paralectotype, MNHN 1927-126 , 84.4 mm SL, Kabana River, tributary of Awash, in Addis Ababa city at an altitude of 2400 m, Ethiopia.

Material examined – non-type material: BMNH 1985.7.16.113–117, five ex., 63.7–79.7 mm SL, Abo River near Addis Ababa, Ethiopia (misidentified as G. dembeensis ) ; BMNH 1985.7.16:111–112, two ex., 26.8–110.9 mm SL, Awash River , Ethiopia (misidentified as G. dembeensis ) ; BMNH 1984.9.7:48, one ex., 44.9 mm SL, Mojo River , Ethiopia (misidentified as G. dembeensis ) ; AMNH 223672 , eight ex., 45.4– 114.3 mm SL, Kabana River , Addis Ababa, Ethiopia (9°N, 38°45′E) ; AMNH 223649 , nine ex., 41.1– 81.5 mm SL, Meki River , Near Meki Town, Ethiopia (8°15′N, 38°50′E) ; AMNH 223674 , 17 ex., 38.8– 78.5 mm SL, Akaki River , Near Addis Ababa (9°N, 38°45′E) ; AMNH 223676 , one ex., 85.8 mm SL, Mekanisa River , Near Addis Ababa, Ethiopia (9°N, 38°45′E) ; AMNH 227315 , 66 ex., 28.8–105.4 mm SL, Megecha (Gubre) River , in Gubre town , about 10 km south of Welkite town on the road to Hosaina, Ethiopia (8°5 N, 37°55E) ; AMNH 223836 , 86 ex., 21.3–99.1 mm SL, Awash River , 1 km east of Ginchi town , 80 km from Addis Ababa on the way to Nekempte, Shoa, Ethiopia (9°5 N, 38°5E) .

GARRA DEMBEENSIS ( RÜPPELL, 1836)

( FIG. 17 A, B; TABLE 9)

Chondrostoma dembeensis Rüppell, 1836 , Mus . Senckenberg Abhandl. Beschr. Naturg. Vol. 2, p. 16, plate 2, fig. 4 (original description, type locality: ‘Dembea’ Lake at Goraza, Abyssinia, type unavailable) Discognathus chiarinii Vinciguerra, 1883 , Ann. Mus . Genova, Vol. 18, p. 696 (original description, type locality: Lake Arsadé, Adda, Ethiopia), Daget et al., 1984, CLOFFA, Vol. 1, p. 302 (as synonym of G. dembeensis ) Discognathus johnstonii Boulenger, 1901 ), Proc. Zool. Soc. Lond., Vol. 2, p. 159 (original description, type locality: Victoria Nyanza), Daget et al., 1984, CLOFFA, Vol. 1, p. 302 (as synonym of G. dembeensis ), Menon 1964, Memoirs of the Indian Museum, Vol. 14, p. 199 (as synonym of G. dembeensis )

Diagnosis: Distinguished from African congeners by the following combination of features: well-developed disc; between zero and four predorsal scales anterior to dorsal fin; asquamate chest, belly, and postpelvic region; intestine of medium length (SL 39.0–42.0% Int.L); and moderately sized gas bladder (posterior chamber 20.9–21.6% SL).

Description: Morphometric and meristic data presented in Table 9. Maximum size: 146.1 mm SL (BMNH 1937.4.20:27). Body relatively gracile (depth 15.6–23.9, mean 20.4% SL), greatest body depth located at level of pelvic-fin tips. Head gracile and moderately depressed, often with small circular tubercles in front of nostrils, becoming tubular beneath eyes. Eyes positioned medially on head. Dorsal head profile rises steeply over snout, then smoothly rounded, more-or-less contiguous with dorsal body profile and smoothly convex to dorsal-fin origin. Disc usually well-developed (type C) with a wide free posterior margin and abundant papillae on lower lip and over disc periphery. However, disc development displays considerable variation with some individuals bearing an almost intermediate (type B) disc with little papillation over disc periphery. In these individuals the disc has free lateral, as well as posterior, margins and is never as weakly developed as in type- A discs. Rostral cap well developed, extending ventrally over upper jaw and with weakly invecked ventral margin. Two pairs of barbels: maxillary barbels usually slightly longer than rostral pair. Between 36 and 38 (modally 37) scales along lateral line; between zero and four predorsal scales anterior to dorsal fin; 3.5–4.5 scales from lateral line to origin of pelvic fin; 3.5–5.5 scales from lateral line to dorsal-fin origin. Belly and chest asquamate; postpelvic region usually asquamate, although occasionally with either one or two deeply embedded scales present. Dorsal-fin rays iii–iv, seven, inserted well anterior to pelvic fin, with short subacuminate tip, first and second unbranched rays longest, extending to last ray; posterior margin either weakly concave or straight, posterior rays may or may not extend to pelvic-fin tip. Predorsal length 44.0–52.0% SL. Anal-fin rays iv, five, with subacuminate tip; posterior margin concave. Pectoral rays iv, 11–12; pectoral length 15.1–23.9% SL, with somewhat pointed tips. Vent located relatively distant from anal fin (vent distance 18.2–38.2, m. 26.0%). Intestine of medium length (SL 39.0–42.0% Int.L). Gas bladder of moderate size (posterior chamber 20.9–21.9% SL).

Colour in preservation: Variable, but most specimens are from medium to dark brown over most of the head and body, with some slight lightening ventrally. Ventral surface of head and body usually a deep creamy yellow-brown. Dorsal-fin pigmentation variable, either with or without a well-marked submarginal black band. Between two and four large black spots between rays in basal membrane of dorsal fin. Anal fin usually with broad submarginal band of dusky brown pigment. Caudal fin pale dusky greybrown; median rays often a slightly darker brownish grey. Pelvic and pectoral fins dark brownish-grey with pale leading edges.

Distribution: As currently recognized, this is one of the most widespread of African Garra , present in 75% of localities surveyed in Ethiopia ( Getahun & Stiassny, 1998), as well as in Cameroon, Nigeria, Tchad, Egypt, Kenya, and Tanzania. Further analysis, particularly of material from western regions, is necessary to confirm the conspecificity of specimens from those areas.

Remarks: Getahun (2000) established that no type specimen for C. dembeensis Rüppell, 1836 was ever deposited at the Senckenberg Museum, and considered that Rüppell’s description was probably based upon information from field observations. Because Rüppell’s original description is inadequate for unambiguous identification, and because considerable taxonomic confusion persists regarding the identity of this widespread species and its synonyms, Getahun (2000) proposed the designation of a neotype to taxonomically fix the name, and we concur. A specimen from the same locality as Rüppell’s type (Lake Tana, Ethiopia) has been selected as a neotype and is illustrated here ( Fig. 17A, B ).

Material examined – type material: Neotype of G. dembeensis, AMNH 223731, 73.8 mm SL, Lake Tana at Gorgora, Ethiopia (12°15′N, 37°20′E); paratypes of D. chiarinii, BMNH 1903.8.10:14–16, three ex., 27.4– 29.9 mm SL, Lake Arsade, ‘Adda Gallaland’, Abyssinia; type of D. johnstonii, BMNH 1901.6.24:94, 98.4 mm SL, Lake Victoria (Victoria Nyanza).

Material examined – non-type material: BMNH 1902.12.13:421–423, three ex., 53.0– 84.7 mm SL, Lake Tana , Ethiopia ; BMNH 1908.1.20:50, one ex., 132.9 mm SL, Didessa River , Western Ethiopia ; BMNH 1937.4.20:24–26, three ex., 95.4–107.9 mm SL, Ghibe River , south-western Ethiopia ; BMNH 1902.11.8:10, one ex., 75.3 mm SL, Nairobi River , Kilimanjaro at 1980 m, Kenya ; BMNH 1937.4.20:28, one ex., 116.5 mm SL, Didessa River , western Ethiopia ; BMNH 1937.4.20:27 and 1900.1.20:51–52, three ex., 123.1–146.1 mm SL, Juju River , Ethiopia ; BMNH 1912.3.22:40–49, nine ex., 53.8–84.4 mm SL, Saya River , Ethiopia ; BMNH 1975.10.7:69–70, two ex., 24.9–58.8 mm SL, Kikuletwa River entering Nyumba Ya Munga dam, Upper Pangani, Tanzania ; BMNH 1985.7.16:107–110, four ex., 22.6–39.8 mm SL, Bilate River entering into Lake Abaya, Ethiopia ; BMNH 1981.4.9:363–366, one ex., 52.4 mm SL, Athi River at Kithimani (Yatta), Kenya ; BMNH 1971.8.12:15–17, three ex., 35.4–38.0 mm SL, tributary of Sanya-Pangani River , Arush National Park, Tanzania ; BMNH 1981.4.9:340, one ex., 61.8 mm SL, Athi River above Lugardo falls, Kenya ; BMNH 1984.9.7:49–52, four ex., 10.7–41.6 mm SL, Muger River , Ethiopia ; BMNH 1971.3.22:13–24, 12 ex., 28.6–35.0 mm SL, Savo River , Ethiopia ; BMNH 1915.12.2:1, one ex., 47.4 mm SL, Makindu and Isavo Rivers, Achi system, Kenya ; BMNH 1969.2.11:215– 230, 16 ex., 21.4–53.8 mm SL, Soni River , trib. of Pangani, Tanzania ; BMNH 1981.4.9:337–339, three ex., 102.1–121.0 mm SL, Ewsso Nyiro , Kenya ; BMNH 1936.12.22:42–47, six ex., 13.6–69.5 mm SL, Migaboti River , Athi system ; BMNH 1937.6.4:24–27, four ex., 42.2–54.7 mm SL, Athi River , Kenya ; BMNH 1971.3.22:25–41, 17 ex., 36.2–67.1 mm SL, Tsavo River , Kenya ; BMNH 1908.1.20:53–58, six ex., 93.9– 115.5 mm SL, Sibe River ; BMNH 1908.1.20:45–48, four ex., 121.6–125.5 mm SL, Sibe River ; MNHN 85- 93 - 18-4 , two ex., 34.7–74.6 mm SL, Tchad ; MNHN 85- 93 - 18-5 , two ex., 63.4–86.7 mm SL, Cameroon ; MNHN 85-93 - 10-4 , two ex., 84.8–100.6 mm SL, Kabana River , Abyssinia ; MNHN 85-93 - 10-5 , 22 ex., 30.3–95.3 mm SL, Ethiopie meridionale ; MNHN 85- 93 - 10-1 , two ex., 26.4–30.5 mm SL, Abyssinia ; MNHN 85-93 - 27-2 , one ex., 74.1 mm SL, Courema River , Affluent of Awash, Arsi, Ethiopia ; MRAC 94-074 -p- 1111–1120, 10 ex., 36.7–69.0 mm SL, Cameroon ; MRAC 91-051 -P-11–15, 10 ex., 29.8–50.2 mm SL, Kotta creek , Ethiopia (misidentified as G. quadrimaculata ) ; MHNG 2317.81 , nine ex., 23.3–48.5 mm SL, Athi River , 14 Fall pres de Thika, Kenya ; MHNG 1446.100 , one ex., 72.6 mm SL, Prov. Simien, Balaghes Valley , Tsion Mariam, 2500 m altitude, Ethiopia ; MHNG 1525,58 – 61 , four ex., 48.9–57.6 mm SL, de l’Ouest du lac Chamo, Rift Valley , ambouchure d’un petit affluent, III, Ethiopia ; MHNG 1523.10 – 12 , three ex., 33.4–39.5 mm SL, Athi River , Falls pres Thika, Kenya ; SMF 1446.100 , one ex., 72.9 mm SL, Prov. Simien, Balaghes Valley , Tihon Mariam, 2500 m altitude, Ethiopia ; SMF 1525.58 – 61 , four ex., 45.0– 55.6 mm SL, de l’Ouest du lac Chamo, Rift valley , ambouchoure d’un petit affluent, III, Ethiopia ; CAS 63736, ten ex., 17.9–51.9 mm SL, Muheza district , Tanga region , Zigi River at corn mill, in the Usambara mountains, 70 km south-east of Amani, Tanzania ; USNM 339720 , three ex., 40.9–62.3 mm SL, Mayo Sabere , a stream draining to Mayo Gashaka , which eventually drains to the River Taraba that joins the River Benue, Nigeria (07°0′20″N and 011°34′30″E) ; USNM 339707 , three ex., 31.4–61.8 mm SL, Hot spring at Matashrip, which eventually drains to the River Taraba which joins the river Benue, Nigeria (07°04′00″N, 011°26′00″E) ; USNM 339699 , three ex., 16.3–24.3 mm SL, Warm spring, tributary of River Jiagum, which eventually drains to the River Taraba that joins the River Benue, Nigeria (07°23′40″N, 011°31′50″E) ; USNM 338339 , three ex., 32.0– 50.1 mm SL, Mayo Dundere , the Upper reaches of the Mayo Gashaka / Mayo Korngal, which eventually drains to the River Taraba which joins the Benue, Nigeria (07°01′50″N, 011°34′00″E) ; AMNH 223771 , 223772 , 60 ex., 27.6–99.0 mm SL, Ardaba River 9 km north of Arsi Negele on route to Wendo Genet, Shoa, Ethiopia (7°30′N, 38°40′E) ; AMNH 223653 , 223654 , 223775 , 50 ex., 29.6–71.1 mm SL, Worka River , Wondo Genet, small stream along the hot spring, Shoa, Ethiopia (7°10′N, 38°40′E) ; AMNH 223788 , one ex., 58.9 mm SL, Chufa stream, c. 4 kilometers south of Abura village, Arsi, Ethiopia (7°55′N, 39°5′E) ; AMNH 223657 , 27 ex., 26.7–68.5 mm SL, Wondo Genet River , crossing southern part of Wondo Genet town, Ethiopia (7°10′N, 38°40′E) ; AMNH 223668 , six ex., 60.8–70.7 mm SL, Acamaja River , about 10 km north of Butajira town on the way to Addis Ababa, Ethiopia (8°10′N, 38°35′E) ; AMNH 223671 , 29 ex., 32.9–91.1 mm SL, Lebu River , about 20 km north of Butajira on the way to Addis Ababa, Ethiopia (8°30′N, 38°40′E) ; AMNH 223683 , 223684 , 36 ex., 27.1–73.1 mm SL, Gurunz River , near Debre Libanos, c. 100 km north of Addis Ababa, Ethiopia (9°40′N, 38°50′E) ; AMNH 223690 , three ex., 28.8– 62.9 mm SL, Gedeb River , c. 20 km north of Debre Markos town, Gojam, Ethiopia (10°30′N, 37°40′E) ; AMNH 223679 , eight ex., 35.8–75.8 mm SL, Duberkela River , 70 km north of Addis Ababa, Shoa, Ethiopia (9°30′N, 38°50′E) ; AMNH 223782 , five ex., 39.5– 67.5 mm SL, Katar River , c. 30 km west of Abura village, Arsi, Ethiopia (7°55′N, 39°E) ; AMNH 223673 , 17 ex., 32.9–70.5 mm SL, Akaki River , crossing southeastern part of Addis Ababa, Addis Ababa, Ethiopia (9°N, 38°45′E) ; AMNH 223758 , six ex., 22.9–109.6 mm SL, Borkena River , at southern end of Kombolcha town, Wollo, Ethiopia (11°10′N, 39°50′E) ; AMNH 223705 , 44 ex., 34.5–45.2 mm SL, Kakist River , near Tillili town c. 50 km north of Finote Selam town, Gojam, Ethiopia (10°55′N, 37°5′E) ; AMNH 223650 , 16 ex., 41.6–87.4 mm SL, Meki River , crossing southern part of Meki town, Rift Valley, Ethiopia (8°15′N, 38°50′E) ; AMNH 223681 , 45 ex., 36.0– 65.4 mm SL, Duberkela River , 70 km north of Addis Ababa, Shoa, Ethiopia (9°30′N, 38°50′E) ; AMNH 227265 , 15 ex., 21.0– 33.7 mm SL, Dogi River , 2 km west of Kumbabe town on the way to Metu, Illubabor, Ethiopia (8°10′N, 36°50′E) ; AMNH 227272 , nine ex., 30.0– 73.6 mm SL, Sore River , at the eastern end of Metu town, Illubabor, Ethiopia (8°15′N, 35°40′E) ; AMNH 227326 , one ex., 99.0 mm SL, Borkena River , southern end of Kombolcha town, Wollo, Ethiopia (11°10′N, 39°50′E) ; AMNH 227296 , 65 ex., 24.5–87.7 mm SL, Meni River , c. 75 km east of Tepi town on the way to Bonga town , Sueda / Gaweta area, Kefa, Ethiopia (7°10′N, 35°50′E) ; AMNH 227315 , 66 ex., 28.8–105.4 mm SL, Megecha / Gubre River , Gubre town , c. 10 km south of Welkite town , on the way to Hosaina, Shoa, Ethiopia (8°5′N, 37°55′E) ; AMNH 223832 , 34 ex., 26.9–91.7 mm SL, Berga River , 2 km west of Addis Alem town , on the way from Addis Ababa to Nekempte, Shoa, Ethiopia (9°5′N, 38°20′E) ; AMNH 227289 , nine ex., 48.6– 116.7 mm SL, Yebeg Wuha River ( Boko river ), at the eastern end of Tepi town, Kefa, Ethiopia (7°10′N, 35°10′E) ; AMNH 227303 , five ex., 40.9–88.6 mm SL, Arer River , c. 30 km north-east of Jima town, on the road to Addis Ababa, Kefa, Ethiopia (7°50′N, 37°10′E) ; AMNH 223840 , four ex., 27.9–61.8 mm SL, Melka Gufu River , 50 km east of Bako town , 200 km west of Addis Ababa, travelling from Addis Ababa to Nekempte, Welega, Ethiopia (9°5′N, 37°15′E) ; AMNH 227293 , 15 ex., 17.7–85.4 mm SL, Minjibat River , c. 30 km east of Tepi town , on the way to Bonga town, Kefa, Ethiopia (7°N, 35°35′E) ; AMNH 223838 , 60 ex., 28.2–97.4 mm SL, Guder River , at the western end of Guder town, Shoa, Ethiopia (9°N, 37°50′E) . AMNH 227248 , two ex., 42.2–70.3 mm SL, Gibe River , western end of Bako town , 255 km west of Addis Ababa on the way to Nekempte, Welega, Ethiopia (9°10′N, 37°5′E) ; AMNH 227268 , three ex., 25.3–58.8 mm SL, Geyi River , c. 5 km east of Metu town, Illubabor, Ethiopia (8°20′N, 35°45′E) ; AMNH 227260 , nine ex., 21.3– 94.2 mm SL, Dabena River , 2 km south-west of Bedelle town, Illubabor, Ethiopia (8°20′N, 36°15′E) ; AMNH 227274 , 22 ex., 24.3–59.0 mm SL, Wangus River , c. 15 km west of Gore town on the way to Gambella , near Uka town, Illubabor, Ethiopia (8°10′N, 35°20′E) ; AMNH 227301 , four ex., 54.2–79.1 mm SL, Bulbul River , c. 10 km north-east of Jima town on the way to Addis Ababa, Kefa, Ethiopia (7°45′N, 36°50′E) ; AMNH 227298 , nine ex. 29.4–69.5 mm SL, Weshi River , c. 10 km south-west of Bonga town, Kefa, Ethiopia (7°20′N, 36°10′E) ; AMNH 227287 , six ex., 48.3– 90.4 mm SL, Sasa River , 12 km south of Gore town, Illubabor, Ethiopia (8°20′N, 35°35′E) ; AMNH 223692 , six ex., 29.09–42.9 mm SL, Bula River , crossing southern part of Dembecha town, Gojam, Ethiopia (10°35′N, 37°30′E) ; AMNH 223783 , 11 ex., 37.05–47.2 mm SL, Katar River , c. 30 km west of Abura village, Arsi, Ethiopia (7°55′N, 39°E) .

GARRA BLANFORDII BOULENGER, 1901

( FIG. 17C, D; TABLE 10)

Discognathus blanfordii Boulenger, 1901 , Proc. Zool. Soc. Lond., p. 160 (original description, type locality: Erythrea or Ethiopia), Daget et al., 1984, CLOFFA, Vol. 1, p. 304 (as synonym of G. quadrimaculata ), Menon, 1964, Mem. Indian Museum, Vol. 14, p. 192 (as synonym of G. quadrimaculata )

Discognathus vinciguerrae Boulenger, 1901 , Proc. Zool. Soc. Lond., p. 160 (original description, type locality: Nile River, north of Kermeh, Sudan), Daget et al., 1984, CLOFFA, Vol. 1, p. 304 (as synonym of G. quadrimaculata ), Menon, 1964, Mem. Indian Museum, Vol. 14, p. 199 (as synonym of G. quadrimaculata )

Discognathus giarrabensis Gianferrari, 1932 , Boll. Zool. Napoli, Vol. 1, p. 2 (original description, type locality: Giarraba, Eritrea), Daget et al., 1984, CLOFFA, Vol. 1, p. 302 (as synonym of G. dembeensis ). Menon, 1964, Mem. Indian Museum, Vol. 14, p. 200 (as synonym of G. dembeensis )

Discognathus blanfordii cimmaruta Gianferrari, 1936 ; Atti. Soc. Ital. Milano, Vol. 75, p. 295 (original description, type locality: Gherungura Stream, Adua, Ethiopia), Daget et al., 1984, CLOFFA, Vol. 1, p. 304 (as synonym of G. quadrimaculata )

Diagnosis: Unique among African Garra in a uniformly either cream or pale yellow-brown coloration and with the absence of distinct markings on head, body, or fins. Further distinguished from African congeners by the following combination of features: well developed disc; either smooth, or only very weakly incised ventral margin to rostral cap; fully scaled predorsal region; asquamate chest, belly, and postpelvic region; and relatively large gas bladder (posterior chamber 21.7–24.4% SL).

Description: Morphometric and meristic data presented in Table 10. Maximum size: 91.7 mm SL (AMNH 223832). Relatively robust and deep-bodied (depth 20.0–25.1, m. 22.8% SL); greatest body depth well in advance of dorsal-fin origin, at level of pectoralfin tips. Head small, snout blunt, and without tubercles. Eyes positioned anteriorly on head. Dorsal head profile rises steeply over snout, then rises more-or-less smoothly convex and contiguously with dorsal body profile to dorsal-fin origin. Disc development, somewhat variable, usually well developed, type-C disc, with abundant papillae on lower lip and around disc periphery. Disc papillation reduced in some specimens, but wide free posterior and lateral margins always present. Rostral cap well developed, extending ventrally over upper jaw. Ventral margin smooth, occasionally very weakly incised. Two small pairs of barbels: rostral barbels usually slightly longer than maxillary pair. Between 34 and 38 scales along lateral line; predorsal region fully scaled to dorsal fin, scales often deeply embedded; 3.5 scales from lateral line to origin of pelvic fin; 4–4.5 scales from lateral line to dorsal-fin origin. Chest, belly, and postpelvic regions asquamate; region between vent and anal fin also asquamate. Dorsal-fin rays iv, seven, inserted well anterior to pelvic fin, with short rounded tip, first and second unbranched rays longest, not extending beyond last ray; posterior margin straight, posterior rays not extending to pelvic-fin tip. Vent distance (20.6–39.5, m. 28.5%). Intestine of moderate length (SL 40.0– 50.0% Int.L). Gas bladder well-developed (posterior chamber 21.7–24.4% SL).

Colour in preservation: Noteworthy in lacking all dark pigmentation and markings on head, body, and fins. Head and body more-or-less uniformly either cream or pale yellow-brown, a few individuals with some slight lightening ventrally. Dorsal fin pale hyaline, with no black spots in basal membranes. All other fins similarly uniformly pale hyaline with no markings.

Distribution: Recorded from Eritrea, Sudan, and Ethiopia (where it is abundant in the Abbay River basin).

Remarks: Getahun (2000) notes that sympatric individuals of G. dembecha are often depigmented and may superficially resemble G. blandfordii in coloration. The two species are readily distinguished by disc morphology (well developed in G. blandfordii vs. vestigial in G. dembecha ).

Material examined – type material: Lectotype of D. blanfordii, BMNH 1869.11.4:39, 56.6 mm SL. Stream at Suru and small pool at Amba, about 25 miles north of Massawa, Eritrea ; Paralectotypes of D. blanfordii, BMNH 1869.2.8:4–11, eight ex., 36.2–51.3 mm SL, Abyssinia ; Paralectotype of D. blandfordii, BMNH 1872.12.18:3, 41.6 mm SL, Abyssinia ; Syntype of D. vincuguerrae, BMNH 1907.12.2:1175–1179, one ex., 35.4 mm SL, near Kermeh , 3rd Cataract (Nile River), Abyssinia ; holotype of D. giarrabensis, MSNM 22 (ex. 4290), 59.2 mm SL, Giarraba , Eritrea ; Holotype of G. blanfordii cimmaruta, MSNM 21 (ex. 4844), 72.9 mm SL, Gherungura , Adua, Abyssinia .

Material examined – non-type material: BMNH 1962.10.31:18–32, 13 ex., 37.3–76.3 mm SL, Blue Nile , 25 miles north of Debre Marcos, Ethiopia (misidentified as G. quadrimaculata ) ; BMNH 1962.10.31:1–17, 18 ex., 37.2–68.9 mm SL, Blue Nile , 25 miles north of Debre Marcos (misidentified as G. quadrimaculata ) ; BMNH 1902.12.13:410–419, ten ex., 46.3–85.5 mm SL, Jerrer River , near Harrar, Ethiopia (misidentified as D. quadrimaculatus ) ; BMNH 1903.11.16:15–17, three ex., 26.4–61.7 mm SL, Gadshimboda River , Ethiopia (misidentified as D. quadrimaculatus ) ; MNHN 85-93 - 10-2 , three ex., 53.4–68.0 mm SL, Afrique Orientale ; MNHN 85-93 - 10-3 , two ex., 53.4–67.3 mm SL, Abyssinia; MSNM 2108 (ex. 4424), one ex., 44.4 mm SL, Abyssinia; MSNM 20 (ex. 4321), one ex., 45.6 mm SL, Massawa , Abyssinia ; AMNH 223686 , 45 ex., 24.4– 62.9 mm SL, Chimuga River , 3 km south of Debremarkos town, Gojam, Ethiopia (37°45E, 10°15 N) ; AMNH 223748 , 35 ex., 27.1–48.5 mm SL, Geba River , 2 km south of Hagere Selam town, Tigrai, Ethiopia (13°35 N, 39°25E) ; AMNH 223832 , 34 ex., 26.9– 91.7 mm SL, Berga River , 2 km west of Addis Alem town on the way to Nekempte from Addis Ababa, Shoa, Ethiopia (9°5 N, 38°20′E) ; AMNH 223685 , 18 ex., 26.4–79.0 mm SL, Chimuga River , 3 km south of Debremarkos town, Gojam, Ethiopia (10°15′N, 37°45′E) .

GARRA IGNESTII ( GIANFERRARI, 1925)

( FIG. 18 A, B; TABLE 11)

Discognathus ignestii Gianferrari, 1925 , Atti. Soc. Ital. Milano, Vol. 64, p. 185 (original description, type locality: Torrent Kahha & Angrab, Ethiopia), Eschmeyer, 1998, Catalogue of fishes, Vol. 1, p. 759 (year of description incorrectly quoted as 1926)

Diagnosis: Distinguished from African congeners by the following combination of features: extremely well-developed, large papillate disc; scaled predorsal and postpelvic regions; chest and belly asquamate; vent located close to anal fin (vent distance 19.4– 25.0, m. 18.2%); and intestine long (SL 20.9–24.2% Int.L).

Description: Morphometric and meristic data presented in Table 11. Maximum size: 103.1 mm SL (AMNH 223709). Robust and deep-bodied (depth 15.3–27.3, m. 21.6% SL); greatest body depth in advance of dorsal-fin origin at level of pectoral-fin tips. Head robust and only slightly depressed, snout prominent with large horny tubercles in both sexes. Eyes positioned medially on head. Dorsal head profile rises steeply over snout, dorsum of head smoothly rounded and contiguous with dorsal body profile, which is steeply convex to dorsal-fin origin in most specimens. Disc well-developed, large, type C, with wide free posterior and lateral margins. Numerous batteries of papillae over lower lip and around disc periphery. Disc wider than long. Rostral fold very well developed and ventrally downgrown to lower lip, with invecked ventral margin. Two pairs of barbels: rostral barbels usually well developed and longer than maxillary pair. Between 34 and 36 scales along lateral line: predorsum scaled (10–12 scales) anterior to dorsal fin; 3.5 scales from lateral line to origin of pelvic fins; 4.5 scales from lateral line to dorsal-fin origin. Chest and belly asquamate. Usually a few embedded scales on postpelvic region. Dorsal-fin rays iv–v, seven, inserted well in advance of pelvic fin, with a slightly rounded tip, first and second unbranched rays longest, extending slightly beyond last ray; posterior margin concave, posterior rays not approaching pelvic-fin tip. Predorsal length 30.6–49.0% SL. Anal-fin rays iv, five, with pointed tip; posterior margin straight. Pectoral-fin rays iv, 11–12; pectoral length 12.4–23.3% SL, large and paddle-shaped with somewhat pointed tips. Vent moderate distance from anal fin (vent distance 19.4– 25.0, m. 18.2%). Intestine is long (SL 20.9–24.2% Int.L). Gas bladder well-developed (posterior chamber 19.0–30.0% SL).

Colour in preservation: Either dark brown or brownish-black; more-or-less uniformly so over head and body, somewhat paler along lower third of body, and ventral surface uniformly either pale creamy-yellow or orange. Faint traces of a darker midlateral band sometimes discernable on posterior flanks. Dorsal fin usually uniformly dusky grey-brown with no clearly demarcated black submarginal band. Either three or four black spots between rays in basal membrane of dorsal fin. Anal fin usually uniformly pale creamy brown without broad submarginal band of dusky brown pigment. Caudal fin uniformly pale dusky greybrown. Pelvic and pectoral fins pale brownish-grey with pale cream leading edges.

Distribution: Known only from Ethiopia, where it occurs in the Tekezze and Abbay River drainages in northern Ethiopia.

Material examined – type material: Syntypes of D. ignestii, MSNM 24 (ex. 4315), 25 ex., 39.1–89.4 mm SL, Kahha and Angereb Rivers, Abyssinia .

Material examined – non-type material: MSNM 2155 (ex. 4316), 14 ex., 39.4–67.7 mm SL, Kahha River, Abyssinia; SMF 1447.1 – 2 , two ex., 80.2–84.5 mm SL, Tihou Mariam , 2500 m, Balaghes valley, Simien province, Ethiopia ; BMNH 1984.9.7:50–60, 11 ex., 31.3– 83.2 mm SL, Tributary of Awash on Dessie road, Ethiopia (misidentified as G. dembeensis ) ; MHNG 1447.1 – 2 , two ex., 79.9–83.4 mm SL, Balaghes valley , Simien prov., Tihou Mariam (2500 m), Ethiopia ; MHNG 1525.49 – 57 , nine ex., 28.3–80.4 mm SL, ‘embouchure d’un petit affluent l’Quest du Lac Chamo’ , Ethiopia ; AMNH 223738 , 223739 , 29 ex., 45.0– 100.4 mm SL, Angereb River , 1 km north of Gondar town, Gondar, Ethiopia (12°50′N, 37°35′E) ; AMNH 223744 , 223746 , 22 ex., 38.9–80.4 mm SL, Elala River , crossing northern part of Makale town, Tigrai, Ethiopia (13°30′N, 39°30′E) ; AMNH 223749 , 223750 , 12 ex., 27.5–80.5 mm SL, Kechen Abeba River , c. 20 km south of Lalibela town on the way to Woldiya town, Wollo, Ethiopia (11°55′N, 39°10′E) ; AMNH 223752 , 223753 , 12 ex., 19.7–64.8 mm SL, Tekezze River , c. 50 km south of Lalibela town, Wollo, Ethiopia (11°50′N, 39°15′E) ; AMNH 227275 , 101 ex., 23.1–100.3 mm SL, Uka River , c. 20 km west of Gore town on the way to Gambella , near Uka town, Illubabor, Ethiopia (8°15′N, 35°10′E) ; AMNH 223759 , seven ex., 45.3–89.3 mm SL, Borkena River , southern end of Kombolcha town, Ethiopia (11°10′N, 39°50′E) ; AMNH 223678 , 223680 , 29 ex., 31.5–77.8 mm SL, Duberkela River , 70 km north of Addis Ababa, Ethiopia (9°30′N, 38°50′E) ; AMNH 223691 , eight ex., 33.3–69.0 mm SL, Gedeb River , c. 20 km north of Debremarkos town, Gojam, Ethiopia (10°30′N, 37°40′E) ; AMNH 223695 , 223696 , 15 ex., 25.0– 75.5 mm SL, Chereka River , Ethiopia (10°40′N, 37°25′E) ; AMNH 223697 , 223698 , 9 ex., 30.7–70.3 mm SL, Kechen River , 30 km south of Finote Selam town, Gojam, Ethiopia (10°45′N, 37°20′E) ; AMNH 223701 , four ex., 31.8–43.4 mm, Lah River , south of Finote Selam town, Ethiopia (10°50′N, 37°15′E) ; AMNH 223709 , 223710 , four ex., 42.5– 103.1 mm SL, Muat River , north of Durbete town, Gojam, Ethiopia (11°10′N, 37°E) ; AMNH 223712 , 223713 , six ex., 32.8–35.0 mm SL, Ernu River , near Tara Gedam, c. 100 km south of Gondar town, Ethiopia (12°20′N, 37°40′E) ; AMNH 223718 , two ex., 33.1– 49.8 mm SL, Wenbaha River , c. 80 km south of Gondar town, Ethiopia (12°20′N, 37°40′E) ; AMNH 223719 , five ex., 25.8–37.8 mm SL, Garno River , southern end of Enfraz town, c. 50 km south of Gondar town, Ethiopia (12°40′N, 37°40′E) ; AMNH 223722 , 223723 , 223725 , 34 ex., 29.6–66.0 mm SL, Gumara (Maksegnit) River , northern end of Maksegnit town, Ethiopia (12°40′N, 37°40′E) ; AMNH 223726 , three ex., 32.4– 39.2 mm SL, Megech River , Ethiopia ; AMNH 223742 , three ex., 24.6–31.5 mm SL, Zarima River , within Tekezze valley, c. 150 km north of Gondar town, Ethiopia (13°25′N, 37°55′E) ; AMNH 223840 , four ex., 28.0– 61.8 mm SL, Melka-Gufu River , 50 km east of Bako town , 200 km west of Addis Ababa, traveling from Addis Ababa to Nekempte, Wellega, Ethiopia (9°5′N, 37°15′E) ; AMNH 223734 , 223737 , 22 ex., 30.2– 50.0 mm SL, Dirma River , about 80 km south of Gondar on the way to Gorgora, in Koladiba town, Ethiopia (37°20′E, 12°25′N) ; AMNH 223791 , 223792 , 34 ex., 20.4–66.9 mm SL, Angordgood creek 4 km northeast of Bahirdar town, Gojam, Ethiopia (11°35′N, 37°30′E) ; AMNH 223800 , 223801 , 53 ex., 18.3– 60.1 mm SL, Gelder River , c. 30 km north of Bahirdar town, Gondar, Ethiopia (11°40′N, 37°30′E) ; AMNH 223804 , three ex., 30.1–34.2 mm SL, Idiyemo creek , c. 20 km north of Bahirdar on Gondar road, Gojam, Ethiopia (11°40′N, 37°30′E) ; AMNH 227253 , three ex., 70.5–95.9 mm SL, Loko River , 60 km south-west of Nekempte , near Didessa River, Welega, Ethiopia (8°45′N, 36°25′E) ; AMNH 227263 , two ex., 51.6– 63.0 mm SL, Tipecha River , 5 km south-west of Bedele town on the way to Metu, Illubabor, Ethiopia (8°15′N, 36°10′E) .

GARRA MAKIENSIS ( BOULENGER, 1903)

( FIG. 18C, D; TABLE 12)

Discognathus makiensis Boulenger, 1903 , Proc. Zool. Soc. Lond., Vol. 2, p. 330, plate 21, fig. 1 (original description, type locality: Maki River at Lake Zwai, Ethiopia), Eschemeyer, 1998, Catalogue of fishes, Vol. 2, p. 1006 and Vol. 3, p. 2220 (date incorrectly quoted as 1904)

Discognathus rothschildi Pellegrin, 1905 , Bull. Mus . Hist. Nat. Paris, p. 291 (original description, type locality: Gotta River, Ethiopia), Menon, 1964, Memoirs of the Indian Museum, Vol. 14, p. 198 (as synonym of G. makiensis ), Daget et al., 1984, CLOFFA, Vol. 1, p. 303 (as synonym of G. makiensis ), Eschemeyer, 1998, Catalogue of fishes, Vol. 2, p. 1474 and Vol. 3, p. 2226 (as synonym of G. makiensis )

Diagnosis: Distinguished from African congeners by the following combination of features: well-developed, large papillate disc; scaled predorsal and postpelvic regions; chest and belly with a few non-overlapping scales over chest and belly; vent located very close to anal fin (vent distance 10.3–16.9, m. 13.6%); and intestine very long (SL 15.9–18.1% Int.L).

Description: Morphometric and meristic data presented in Table 12. Maximum size: 135.3 mm SL (MNHN 1905-0246-0247, syntype of D. rothschildi ). Robust and deep-bodied (depth 17.5–27.8, m. 24.7% SL); greatest body depth well in advance of dorsal-fin origin at mid-level of pectoral fin. Head robust, moderately depressed, snout large and prominent, studded with variously sized tubercles in both sexes. Eyes positioned either medially or slightly posteriorly in large specimens. Dorsal head profile rises steeply over snout, dorsum of head smoothly rounded and contiguous with dorsal body profile, which is steeply convex to dorsal-fin origin. Disc fleshy and welldeveloped, large type C, with wide free posterior and lateral margins. Numerous batteries of papillae over lower lip and around disc periphery. Disc slightly wider than long. Wide free posterior margin studded with large and conspicuous papillae. Rostral fold very well developed and ventrally downgrown to lower lip, with invecked ventral margin. Two pairs of barbels: rostral barbels well developed and longer than maxillary pair. Between 35 and 37 scales along lateral line; predorsum scaled (14–15 scales) anterior to dorsal fin; either three or four scales from lateral line to origin of pelvic fins; 3.5–4.5 scales from lateral line to dorsal-fin origin. Predorsal and postpelvic regions scaled; belly with some large, non-overlapping scales embedded in skin. Chest asquamate. Dorsal-fin rays iv, seven, inserted well in advance of pelvic fin, with short subacuminate tip, first and second unbranched rays longest extending to last ray; posterior margin slightly concave, posterior rays not approaching pelvic-fin tip. Predorsal length 43.6–50.6% SL. Anal-fin rays iv, either four or five, with pointed tip; posterior margin straight. Pectoral-fin rays iv, 11; pectoral length 15.3– 24.3% SL, large with somewhat pointed tips. Vent close to anal fin (vent distance 10.3–16.9, m. 13.6%). Intestine very long (SL 15.9–18.1% Int.L). Gas blad- der well developed (posterior chamber 19.0–22.0% SL).

Colour in preservation: Either dark brown or brownish-black; somewhat paler along lower third of body and ventral surface either uniformly pale creamy yellow or orange. Faint traces of a darker midlateral band often discernable on flanks and caudal peduncle. Dorsal fin usually uniformly pale dusky grey-brown, usually with no clearly demarcated black submarginal band (submarginal present in a few specimens). Between three and five small black spots between rays in basal membrane of dorsal fin. Anal fin usually uniformly pale creamy brown, sometimes with faint submarginal band of dusky brown pigment. Caudalfin: uniformly pale dusky grey-brown. Pelvic and pectoral fins pale brownish-grey with pale cream leading edges.

Distribution: Known only from Ethiopia, where it occurs in Lake Langano and the rivers that drain this and other rift valley lakes (Ziwai and Abaya), also found in the Awash and Ghibe-Omo basins in southern Ethiopia.

Material examined – type material: Lectotype of D. makiensis, BMNH 1905.7.25:87, 69.4 mm SL, Maki River running into Lake Ziwai, Ethiopia; paralectotype of D. makiensis, BMNH 1905.7.25:88, 48.6 mm SL, Maki River running into Lake Ziwai, Ethiopia; syntypes of D. rothschildi, MNHN 1905 -0246-0247, two ex., 108.7–135.3 mm SL, Gotta River, Awash basin, Ethiopia.

Material examined – non-type material: BMNH 1970.10.28:57, one ex., 121.8 mm SL, Elgo River , Lake Abaya basin, Ethiopia (misidentified as G. dembeensis ) ; BMNH 1968.7.24:49–53, five ex., 37.6–66.6 mm SL, Ethiopia N.O.O (undefined acronym) (misidentified as G. quadrimaculata ) ; BMNH 1984.9.7:61–62, two ex., 31.9–56.0 mm SL, Didessa River , Ethiopia (misidentified as G. quadrimaculata ) ; MRAC 91.051 - p-65–74, five ex., 48.9–70.5 mm SL, Gota River , Harar province, Ethiopia (misidentified as G. quadrimaculata ) ; AMNH 223655 , three ex., 51.2–60.4 mm SL, Wondo Genet River , crossing southern part of Wondo Genet town, Ethiopia (7°10′N, 38°40′E) ; AMNH 223774 , four ex., 70.2–103.7 mm SL, Worka River , Wendo Genet, small stream along hot spring, Ethiopia (7°10′N, 38°40′E) ; AMNH 223652 , three ex., 30.8– 80.9 mm SL, Worka River , adjacent to the hot spring of the Wabi-Shebelle hotel, Wondo Genet, Ethiopia (7°10′N, 38°40′E) ; AMNH 223666 , 31 ex., 32.5– 87.5 mm SL, Mesha River , north of Butajira town, in the rift valley, Ethiopia (8°10′N, 38°30′E) ; AMNH 223667 , three ex., 90.1–98.6 mm SL, Acamaja River , about 10 km north of Butajira town on the way to Addis Ababa, in the rift valley, Ethiopia (8°20′N, 38°35′E) ; AMNH 223669 , 14 ex., 72.5–105.6 mm SL, Acamaja River , about 10 km north of Butajira town on the way to Addis Ababa, in the rift valley, Ethiopia (8°20′N, 38°35′E) ; AMNH 223656 , four ex., 32.3– 50.8 mm SL, Wondo Genet Rriver , crossing southern part of Wondo Genet town, Ethiopia (7°10′N, 38°40′E) ; AMNH 223670 , two ex., 56.8–91.7 mm SL, Lebu River , about 20 km north of Butajira on the way to Addis Ababa, Ethiopia (8°30′N, 38°40′E) ; AMNH 223762 , three ex., 42.7–61.9 mm SL, Lake Langano , about 150 km south of Addis Ababa, Ethiopia (7°35′N, 38°45′E) ; AMNH 223763 , 15 ex., 31.2–47.2 mm SL, Lake Langano , about 150 km south of Addis Ababa, Ethiopia (7°35′N, 38°45′E) ; AMNH 223789 , 31 ex., 26.1–59.3 mm SL, Chufa stream, about 4 km south of Abura village, Arsi, Ethiopia (7°55′N, 39°5′E) ; AMNH 223770 , four ex., 38.7–52.3 mm SL, Katar River , about 30 km west of Abura village, Arsi, Ethiopia (7°55′N, 39°E) ; AMNH 227289 , nine ex., 48.6–116.7 mm SL, Yebeg Wuha River , at the eastern end of Tepi town, Kefa, Ethiopia (7°10′N, 35°10′E) ; AMNH 223723 , 14 ex., 72.6–130.8 mm SL, Errer Gota River , Eastern side of Errer town, pools near main road, Hararge, Ethiopia (09°30′N, 41°15′E) ; AMNH 227290 , 19 ex., 25.8– 114.0 mm SL, Bitin River , 2 km east of Tepi town , on the way to Bonga town, Kefa, Ethiopia (7°10′N, 35°15′E) ; AMNH 227296 , 65 ex., 24.5–87.7 mm SL, Meni River , 75 km east of Tepi town on the way to Bonga town , Sheda / Gawete area , Kefa, South-west of Ethiopia (7°10 N, 35°50E) ; AMNH 227275 , 101 ex., 23.1–100.3 mm SL, Uka River near Uka town about 20 km west of Gore town on the way to Gambella, Illubabor, Ethiopia (8°15′N, 35°10′E) ; AMNH 223664 , three ex., 45.9–100.8 mm SL, Mesha River , north of Butajira town, Rift Valley, Ethiopia (8°10′N, 38°30′E) ; AMNH 223655 . three ex., 51.1–60.4 mm SL, Wondo Genet River , crossing southern part of Wondo Genet town, Shoa, Ethiopia (7°10′N, 38°40′E) .

Figures

Figure 15. Garra dembecha, holotype, AMNH 228483, 78.8 mm standard length.

Figure 16. (A) and (B) Garra quadrimaculata (type of Garra tibanica modified after Trewavas, 1941). (C) and (D) Garra aethiopicus, AMNH 223649, 84.8 mm standard length.

Figure 17. (A) and (B) Garra dembeensis, neotype, AMNH 2233731, 73.8 mm standard length. (C) and (D) Garra blanfordii, AMNH 223686, 65.8 mm standard length.

Figure 18. (A) and (B) Garra ignestii, AMNH 223750, 81.4 mm standard length. (C) and (D) Garra makiensis, AMNH 223667, 107.2 mm standard length.