A STUDY OF THE LOACHES OF THE GENERA COBITIS AND SABANEJEWIA (PISCES, COBITIDAE) OF GREECE, WITH DESCRIPTION OF SIX NEW TAXA

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1 Trav. Mus. nad. Hist. nat. (~Grigore Antipan Vol. XXXVI pp A STUDY OF THE LOACHES OF THE GENERA COBITIS AND SABANEJEWIA (PISCES, COBITIDAE) OF GREECE, WITH DESCRIPTION OF SIX NEW TAXA PANOS S. ECONOMIDIS, TEODOR T. NALBANT Les CobitidCs de Grtce des genres Cobitis et Sabanejewia ont au total neuf esptces, dont huit appartiennent au premier genre et une au second. Quatre espkces nouvelles de Cobitis sont dccrites (stephanidisi, hellenica, arachthoser~sis et punctilineata), ainsi que deux sous-esptces nouvelles de Sabanejevia arrrata (doiranica et thrakica). L'article contient aussi une ample discution sur I'origine de la faune gr&que de CobitidCs. INTRODUCTION The systematics and phylogeny of the loaches of the Balkan Peninsula, which belong to the genera Cobitis and Sabanejewia, represent one the most complex and interesting of problems. Therefore, the main aim of this contribution is to clarify, as accurately as possible, the position of each species. This seems to be a difficult task, especially because Karaman (1924, 1928, 1929, 1955) has described many taxa of Cobitis very briefly and without figures, excepting ohridana and strumicae. Our difficulties consisted in ascribing our specimens to one or to another of Karaman's taxa. Only for strumicae (see Karaman, 1955: , Fig. 4) did he gave more details and quite a good figure but no information about the structure of the lamina circularis (Canestrini's scale), an extremely important feature for defining Cobitis species. On the other hand, neither the book of Vukovic & Ivanovic (1971), dealing with the freshwater fishes of Yugoslavia, nor the contribution of Grupce & Dimovski (1976), dedicated to the genus Cobitis in the southern part of this country, give any solution in this problem. Moreover, there is no any mention of Karaman's type material. As far the situation of Cobitis and Sabanejewia in Greece is concerned, the information is very poor. Stephanidis (1939) refers to Cobitis taenia (now Cobitis hellenica sp. n.) from, the river Louros (western Greece) and more recently (Stephanidis, 1974) he gives a compiete

2 296 P.S. ECONOMIDIS. T.T. NALBANT description of a new species, Cobitis (Bicanestrinia) trichonica, from lakes Trichonis, Lyssimachia and Ozeros of the western Greek mainland. On the other hand Economidis (1974) noted all morphological variability of various populations of Cobitis taenia (now Cobitis stumicae ) and Cobitis aurata (= Sabanejewia aurata) in their area of distribution, i.e. the aquatic system from the river Evros to the river Strymon in the Greek parts of Thrace and eastern Macedonia. Later, Economidis et al. (1981) and Economidis & Voyadjis (1985) listed Cobitis taenia (Cobitis vardarensis) and Sabanejewia aurata from the Aliakmon and Axios and Gallikos rivers respectively, and Economidis & Sinis (1982) gave a short description of Cobitis taenia (now Cobitis strumicae) from lakes Koronia and Volvi system. Therefore, another aim of this paper is to present a synthesis of all species of Cobitis and Sabanejewia recorded from Greece, on basis of a great number of samples collected at many localities in the country. Besides this, a rich collection of loaches in the Institute of Biology of Bucharest (Romania) is used (see below) as comparative material. Six (4 species and 2 subspecies) taxa are here described as new from different basins of Greece. For more information about the aquatic drainge of Greece see Fig. 1, in which are also presented all Balkan drainges south of the river Danube. Cobitis ohridana Karaman, 1928, may be present also in the Greek freshwater catchment, since some Cobitis specimens from river Aoos near Konitsa, agree quite well with Karaman's figure (1928: 163, Fig. 5). However, the original description of ohridana is extremely brief and cannot be used for its identification. Therefore, based on our observation of specimens from river Aoos, which have a single lamina circularis, we refer ohridana rather to Cobitis s. str. than to Bicanestrinia. MATERIAL AND METHODS Collections. All specimens collected in Greece are mentioned under "material" for each species. Paratypes of the new taxa are also deposited to the following Museums: BMNH, CAS, ISBB, MINB, MNHN, MZUT, NMW, SMF, USNM, ZSM (see abbreviations). Compara five material. We also used much material from different institutions as follow: ISBB, MINB, MSNT, NMC, RMNH, SMF, ZMB. In order to understand the value of each charactrer and to discern the relationships between the species, we have used as comparative material the following species (measurements represent the standard lenght in milimeters): Cobits taenia Linnaeus, 1758: RMNH. AAA , (7) Sweden, no other indications; RMNH. SBM , (1) 80.0, Sweden, Tumba, Uttran; RMNH, AAA , (1) 76.0, Sweden without other data; RMNH. AAA , (2) 52.7 and 61.0,

3 COBITIDS OF GREECE 297 Sweden, without other data; RMNH. A , (1) 47.8 Sweden, Varmdon, Sagsjon; ISBB 2751 (9) , Holland (The Netherlands) without other data. Cobitis danubialis Biicescu, 1993: ISBB uncat. (12) , Romania, Transilvania, river Murev, Stinceni, Cobitis megaspila Nalbant, 1993: ISBB uncat. (6) , Romania, Danube Delta, Caraorman, paratypes. Cobitis maroccana Pellegrin, 1929: ISBB uncat., (3) , Portugal, Tajo drainage, Rio Latas at Coruche; ISBB 3981, (5) , Spain, Guadiana drainage, Rio Quejigares, Fontanosas near Ciudad Real. SMF 3231, (2) 65.0 and 66.0, Spain, Silla at Valencia, Aqua Comunero at 1.5 km from Albufera, paratypes of Cobitis haasi Klausewitz, Cobitis bilineata Canestrini, 1866: ISBB 3089, (9) , Italy, Po drainage, 'Torente Meletta near Carmagnola; ISBB 3090, (15) , Italy, Po drainage, Canale Moneta at Carmagnola. Cobitis narentana Karaman, 1929: ISBB 4577 (3) , river Neretva, Croatia, no other data. Cobitis zanandreai Caviccioli, 1965; ISBB uncat., (1) 63.0, Italy, Lago di Fondi near the Thyrenian Sea, no other data. Cobitis turcica Hanko, 1925: ISBB uncat., (1) 63.0, Turkey, Anatolia, Iqikli golu; ISBB uncat. (3) , Turkey, Kizil Dere, Bagilli-Koyu, 1993; MZUT uncat. (1) 54.0, bearing same data as previous specimens. Cobitis simplicispina Hanko, 1925; ISBB 3566, (2) 55.2 and 76,2, Turkey, Anatolia, Aci golu, Karahiiyuk; MINB uncat., (1) 66.3, Turkey, Golhisar, Cobitis elazigensis Coad & Sarieiyupoglu, 1988: NMC uncat., (1) 108.0, Turkey, Anatolia, Elazig stream near Elazig. Cobitis linea Heckel, 1840: ISBB uncat., (3) , Iran, river Kor near Persepolis, 1800 m altitude, Cobitis elongata Heckel & Kner, 1858, ISBB uncat., (I) Romania, south Banat, river Nera at Sasca Montana. Cobitis calderoni Biicescu, 1962: ISBB 2744, (2) 54.0 and 64.2, Spain, Zamora province, Lago de Sandin de Carbaleda. Cobitis bilseli Battalgil, 1942: MSNT 3948, (1) 195.0, Turkey, Anatolia, Beyshehir golu, Beyshehir; ISBB uncat. (1) 155.0, Beyshehir golu, Beyshehir. Cobitis macrostigma Dabry, 1872: ISBB uncat., (2) and 135.0, China, Hupeh, lake Liang Tse-hu, Yangste drainage. Cobitis melanoleuca Nichols, 1925: ISBB 3230, (4) China, Shansi, Kwei-hwa, Third Asiatic Expedition; ISBB 2290, (9) , Mongolia, Ugij nor, river Nargin Gol, Selenga drainage, Arhangaj Aimak, paratypes of Cobitisgranoei olivai Nalbant, Holcik & Pivnicka, 1970.

4 298 P.S. ECONOMIDIS. T.T. NALBANT Cobitis dolichorhynchus Nichols, 1918: ISBB uncat., (1) 79.9, China, Yangtse basin, no other data. Cobitis sinensis Sauvage & Dabry, 1874: ISBB uncat., (4) , South Korea,river Yung San. Cobitis lutheri Rendahl, 1935: ISBB uncat., (4) , South Korea, river Yung San. Cobitis biwae Jordan and Snyder, 1901: ISBB uncat., (4) , Japan, Honsu, river Kamo at Kyoto. Cobitis striata Okada, 1936: ISBB uncat., (5) , Japan, Honsu, Okayama-ken, river Yoshi. Iksookimia koreensis (Kim, 1976): ISBB uncat., (5) , South Korea, Gyonggy-do, river South Han. Iksookimia longicorpus (Kim, Choi and Nalbant, 1976): ISBB uncat., (4) , South Korea, river Seom Jin. Iksookimia hugowolfeldi Nalbant, 1993: ISBB 4495 (1) holotype 64.5 and ISBB 4496 (2) paratypes , South Korea, river Yung San. Definitions. In defining a species of the family Cobitidae besides meristic and metric values it very important to take into consideration the following characters. Scales: shape, size and extent over the body. Rendahl (1943, 1944, 1945, 1948) gave special attention to the shape and size of subdorsal scales. These scales are situated halfway between the dorsal fin and the lateral line or lateral midline of the body, and were used by him for measurements. The shape of these scales is also characteristic for each species. Therefore every mention in the present contribution (or drawing of a scale) refers only to the subdorsal scales. Colour pattern. In Cobitis species there is a peculiar arrangement of longitudinal rows of spots or bands, already pointed out by Gambetta (1934), and called Gambetta's zones of pigmentation (see Fig. 3 a). However, we found at least six main arrangements and shape of these rows as variations of the initial type (Fig. 3 a-f). On the other hand, at the upper part of base of caudal fin there is a black spot which in different species can be faded to complete absence or, in a few cases, there can be two or even three such spots at the base of caudal. In the closely related genera Sabanejewia and Iksookimia (Nalbant, 1993, 1994) this arrangement of spots is lacking. However, one or two rows of small spots (particularly a more of less lengthy band of vermiculate speckles) are present between the dorsal row and lateral row of large spots.no jet black spot at the upper part of base of caudal is present in Sabanejewia but always there are two dark gray spots sometimes united, at the base of the caudal fin. The pigmentation of the head is also important. In Cobitinae there is always a dark gray, sometimes blackish, band from the anterior rim of the

5 COBITIDS OF GREECE 299 orbit to the tip of snout (Fig. 2a, b, c), but in different species (see Fig. 2 a, b) other dark gray bands, posterior or inferior the eye, may be present. Sexual dimorphism. When present in Cobitinae loaches, this is one of the most important characters. The genus Cobitis has peculiar sexual dimorphism: most adult males have in pectoral fins at the base of second ray, sometimes in both first and second rays, a special plate-like process, so called lamina circularis or Canestrini's scale. We follow Rendahl (1930, 1933 a, 1933 b, 1943, 1944, 1945, 1948) in calling this process the lamina circularis instead of Canestrini's scale since the latter term may be confused with a body scale. Although the shape of this process is very variable, in most cases it is round, and we can thus utilize the term for defining this thin osseous plate-like process. This process is characteristic in shape and size for different species (Fig. 4 a, b, c). In the adult males of Sabanejewia this kind of sexual dimporphism is lacking but the males have on the both sides of body two swellings, the larger being before of the dorsal fin and the smaller, behind this fin. Suborbital spine. This is the movable lateral ethmoid bone which in nearly all Botiinae and Cobitinae is placed on the each side of the head. The different parts of the suborbital spines are treated by Nalbant (1963). Generally the shape and size of the suborbital are characteristic in many species. Barbels and mental lobes. The structure of the lip in Cobitinae is one of the most important characters in defining species. Rendahl(1933 a, 1944, 1945, 1948) described in detail for each species the structure of mouth and the arrangments of barbels and mental lobes around it. We follow him in this way. Different parts of the mouth structure in Cobitinae were already discussed by Nalbant (1963) and BHnZirescu & Nalbant (1964). Lateral line. In species of Cobitinae this can be very reduced to a small portion above the pectoral fin or can be completely absent; only in the genus Somileptes from India is the lateral line complete to the base of caudal fin. However, pairs of neuromast pits are present along the sides of the body (Zanandrea et al., 1965). For terminology of different morphological aspects (mouth and barbels, scales, color pattern, suborbital spine, gas-bladder capsule) see Nalbant (1963). Counts and measurements. These were made according to BgnZirescu et al. (1972), by the same person in order to avoid individual differences in estimates. All measurements are expressed in percentage of standard length and presented in the tables. Drawings and maps. Those of species and anatomical parts were made by TTN from whole specimen or after dissections, while the maps were designed by PSE in a Macintosh PC using the Canvas 3.0 program.

6 300 P.S. ECONOMIDIS. T.T. NALBANT KEY TO THE SPECIES OF COBITINAE OF GREECE l(6) Pigmentation of body not disposed in four Gambetta's zones. Caudal fin with 12 branched rays. Sexual dimorphism: adult males without lamina circularis at base of first pectoral rays but with a conspicuous swelling on each side of body before dorsal fin (genus Sabanejewia) (5) Lateral spots generaly large and square-like. Only in rivers (4) lateral spots, mostly Sabanejewia aurata thrakica ssp. nov. 4(3) 7-14 lateral spots, mostly Sabanejewia aurata balcanica 5(2) Lateral spots mostly small and round. Only in lake (Doirani) Sabanejewia aurata doiranica ssp. nov. 6(1) Pigmentation of body disposed in four Gambetta's zones. Caudal fin with 14 branched rays. Sexual dimorphism: adult males with lamina circularis at base of first two pectoral ray or only at second one (genus Cobitis) (10) Adult males with one plate-like lamina circularis at base of second pectoral ray (subgenus Cobitis s. str.) Fig. 4c (9) The spots of the fourth Gambetta's zone well separated (as in Fig. 3 a)... Cobitis vardarensis 9(8) The spots of the fourth Gambetta's zone as a continuous band, rarely interrupted (as in Fig. 3 b, c and 6 a)... Cobitis stephanidisi sp. nov. lo(7) Adult males with two plate-like lamina circularis at base of first and second pectoral rays (subgenus Bicanestrinia) Fig. 4 d ll(12) Body relatively elongate (11) Body high and stout (16) Lateral spots of fourth Gambetta's zone generally normal in size, somewhat square-like and not disposed on a black stripe (as in Fig. 3 f) (15) In young specimens the lateral spots of fourth Gambetta's zone are rare or very rare remote enough one another (as in Fig. 12 a -f) Cobitis arachthosensis sp. nov. 15(14) In young specimens the lateral spots of fourth Gambetta's zone are normal in number close enough one another (as in Fig. 10 a, b and 12 g-i)... Cobitis hellenica sp. nov. 16(13) Lateral spots of fourth Gambetta's zone reduced in size, rounded, generally contiguous or very close to one another and on a black stripe. (as in Fig. 11 a and 12 j-1)... Cobitis punctilineata sp. nov. 17(18) The general pigmentation of body is cloudy, the spots being not well delimited (as in Fig. 8 a)... Cobitis meridionalis 18(17) The general pigmentation of body and spots are well defined... 19

7 COBITIDS OF GREECE (20) Lateral spots of fourth Gambetta's zone very large and well individualized, on the caudal peduncle being mostly vertical (as in Cobifis trjchonica Fig. 9 a)... 20(19) Lateral spots of fourth Gambetta's zone normal or reduced in size, sometimes, close together, having same shape on the whole length of body (as in Fig. 7 a, b)... Cobitis sfrumicae RESULTS In the systematic account all species are presented as follows: Genus COBITIS Linnaeus, 1758 Subgenus Cobitis s. str. Cobitis vardarensis Karaman, 1928 (Fig. 5 a-g) Cobitis taenia. Apostolidis (partim): 27 (description: an arched black spot in the base of the upper part of caudal fin: distribution: in many rivers and streams of Thessaly). 1907: 23 (Thessaly): Stephanidis. 1950: 207 (river Pinios in Thessaly), Dimovski & Grupce, 1971: 27 (river Axios drainage: tributary BrCgalnitsa): Economidis, 1973 (partim): 471 (from literature: lake Doirani), 559 (from collections: lake Doirani and rivers & streams Axios, Gallikos, Anthemous): Economidis, Kattoulas & Stephanidis, 1981: 93 (list, lower part of rivers Aliakmon. Edesseos and Itamos): Economidis & Voyadjis, 1985: (list Axios, Doirani, Gallikos). Cobitis taenia taenia. Karaman, 1924 (partim): 74 (river Axios near Skopje, lake Doirani). Cobitis taenia vardarensis Karaman. 1928: (short description as new subspecies from river Axios or Vardar): 1929: 173 (short description again); Taler, 1953 (partim): 447 (river Axios or Vardar, lake Doirani): Grupce & Dimovski, 1973: 89 (river Axios system); 1976: 32 (characters, comparisons with other species of South Yugoslavia). Material: MZUT (28) , river Axios catchment, channels near Koufalia, Macedonia, Greece, 16 July 1971, Economidis coll.; MZUT (7) , Gallikos stream, ditch near village Mandres Kilkis, Macedonia, Greece, 27 July 1971, Economidis coll.; MZUT (10) , Gallikos stream near village Argyroupoli Kilkis, Macedonia, Greece, 29 July 1971, Economidis coll.; MZUT (7) , Gallikos stream, near village Petroto Kilkis, Macedonia, Greece, 15 November 1977, Economidis coll.; MZUT (19) , river Loudias catchment, near Skydra, Macedonia, Greece, 21 September 1977, Economidis coll.; ISBB uncat., (15) , bearing same data as previous; MZUT (13) , river Loudias catchment, near village Exaplatanos Aridea, Macedonia, Greece, 10 November 1987, Economidis coll.; MZUT (8) , river Aliakmon, down bridge of old national road Thessaloniki-Katerini, Macedonia, Greece,

8 302 P.S. ECONOMIDIS. T.T. NALBANT 22 September 1971, Stephanidis coll.; MZUT (3) , river Pinios catchment, channels near Hasabali, Thessaly, Greece, 10 August 1986, Economidis coll.; MZUT (5) , river Pinos catchment, tributary Lithaios within the town Trikala, Thessaly, Greece, 18 October 1989, Economidis and Banarescu coll.; ISBB 4575 (10) , bearing same data as previous; ISBB uncat., (12) , river Pinios catchment, tributary Xerias at Mati Tyrnavou, Thessaly, Greece, 15 November 1977, Economidis coll.; ISBB uncat., (2) , river Axios catchment, at village Gynekokastro Kilkis, Macedonia, Greece, 8 November 1977, Economidis coll.; ISBB 4573, (15) , river Loudias drainage, at Aridea, Macedonia, Greece, 11 November 1987, Economidis coll. Karaman (1928) gave only a short description of the species, and thus a more detailed one needs to be provided here. Description: Fin ray formula. Generally the number of fin rays is constant, with D I11 7, A , V I 6-1 6, P I 9-1 9, Cn n, but in a few individuals we found 5 branched rays in both ventrals, 8 branched rays in both pectorals and or branched rays in caudal. Morphology. The body is moderately elongate as in Cobitis taenia with a short caudalpeduncle (Fig. 5 a, b, c). A very reduced keel above and below of the peduncle, near the base of caudal fin. Ventrals are insered under the second or third dorsal branched rays but in a few specimens the insertion of ventrals is on the same vertical line as the dorsal origin. The head is relatively short and stout, with a blunt snout. All three pairs of barbels are short. The mental lobes (Fig. 5 d) have folded zone well developed but the median lobes are relatively reduced. The suborbital spine (Fig. 5 f) is generally straight, with laterocaudal process reduced or very reduced. A relatively small spine is observed in the largest specimens from the rivers Loudias, Gallikos and Pinios. The scale (Fig. 5 e) are generally circular, wider at base with a small eccentric focal zone. They are embedded and cover the whole body except the head. Sexual dimorphism: In males there is a plate-like lamina circularis at the base of the second pectoral ray (Fig. 5). In males from the rivers Pinios and Gallikos, the distal margin of the lamina circularis is more rounded than in the males from river Loudias, in which this'process is somewhat more acute. The colourpattern in life is very similar to that of Cobitis taenia. The whole body is yellowish with olive gray spots and dots. The arrangement and shape of spots and dots is similar in all population from Pinios, Gallikos, Axios and Loudias (Fig. 5a, b, c). The head is pigmented with small speckles. from the antrior part of eye to the tip of the snout a dark stripe. Two other stripes run from the near part of the orbit to the middle of

9 COBITIDS OF GREECE 303 the opercular margin and another obliquely below the eye down towards the mouth. The median row of dorsal spots is prsent in all specimens.these spots are large enough before the dorsal fin and large and square-like in the post-dorsal space. All four Gambetta's pigmentary zones are well developed. The median lateral row of spots (Gambetta's fourth zone) is formed by a large number of round spots (13-24). The iris of the eye is golden-yellow. A jet black spot in the superior part of caudal base is normally well marked in all populations but may be lacking in a few individuals. Dorsal and caudal fins with 4-5 rows of dots, the rest of the fin being pale. The intramuscular septum along the lateral midline is visible as a longitudinal black stripe. In all preserved specimens (ethanol 70%) the ground colour becomes bright yellowish-brown or yellowish-white with gray or grayish-brown spots. Habitat: The species is typically a still water fish, preferring better death branch of river with more or less deep water (more than cm) where it burrows into a muddy or silty bottom, especially among the roots of plants. However, it also can be observed in shallows and, rarely, in slowly running water. Range (Fig ): The area of this species is confined to the following water systems: Pinios (Thessaly), lower Aliakmon, Loudias, Axios and Gallikos (central Macedonia). Remarks: This species differs enough from Cobitis taenia in its combination of characters: a relatively high body, numerous and small lateral spots and by the presence of a large jet black spot at superior base of caudal fin. Cobitis stephanidisi sp. nov. (Fig. 6 a -e) Cobitis taenia, Apostolidis, 1892 (partim): 27 (description: an arched black spot in the base of the upper part of the caudal fin, and a series of 12 to 18 black along the side of the body; distribution: karstic spring Kefalovriso in Velestino village): Panagiotopoulos, 1916: 576 (list: lake Karla). Material: Holotype: MZUT , 56.9, lake Karla catchment, spring within Velestino village, Thessaly, Greece, 27 August 1992, Economidis coll. Paratypes: MZUT (9) , lake Karla catchment, spring within the village Velestino, Thessaly, Greece, 14 April 1988, Economidis coll.; MZUT (56) , same locality as previous, 2 May 1989, Economidis coll.; MZUT (26) , same locality as previous, 27 August 1989, Economidis coll.; MZUT

10 304 P.S. ECONOMIDIS. T.T. NALBANT (23) , same locality as previous, 27 August 1992, Economidis coll.; USNM (I), same locality as previous, 27 August 1992, Economidis coll.; SMF (4), bearing same data as previous; MNHN (2), bearing same data as previous; NMW (1 & 2), bearing same dataas previous. Diagnosis: A long bodied species of Cobitis with a continuous black or dark gray, fourth Gambetta's zone, along the lateral midline of the body, a conspicuous jet black dot at the upper base of the caudal fin. Description: Fin ray formula. D 111 7, A , V I 6-1 6, P I 9-1 9, Cn n. There is some variation in the number of dorsal rays, in few specimens there are 6 branched rays and in a single individual 5 branched rays. In the caudal, fin, generally there is some variation in number of branched rays; two individuals had branched rays, in one and another 7 + 8, but in this last case. the uppermost branched ray was possibly divided to the base. Morphology. Body long, slightly compressed (Fig. 6 a). Head normally large with a pointed snout. Eyes moderately large situated somewhat near the tip of the snout, rather than the near margin of the operculum. The fins moderately large. The dorsal is inserted a little in the second half of the body. Ventrals insertion is situated on the same line of dorsal or a slightly in advance. Caudal peduncle long enough and well compressed laterally. Mouth (Fig. 6 b) well arched. Upper (anterior) lip moderately thick laterally, and is thickened in its anterior part. Fine transverse grooves on the surface of this lip can be observed. Lower (posterior) lip fleshy, well folded with conspicuous mental lobes having pointed tips. Barbels small. Suborbital spine (Fig. 6 d) is straight but with well curved caudal processes (thorns). The thorns are not divergent, the smaller being about half the length of the longer. Scales (Fig. 6 c) nearly rectangular with rounded corners having a very small eccentric focal zone. They cover whole body except the head. Sexual dimorphism. In stephanidisi the males have a large rounded lamina circularis (Fig. 6 e) at the base of the pectoral ray, which is more thickened than the first. This is covered by soft tissue and skin. The part which connects the ray with the plate-like process, is large enough. Such a part is lacking in the species of Bicanestrinia or somewhat reduced in other species of Cobitis s. str. as in vardarensis (compare the Figures of lamina circularis of vardarensis and stephanidisi). CoJour pattern (based on preserved specimens). General ground colour is yellowish-white and the spots and dots dark-gray or bluish-dark gray. On the back there are 4-7, usually 5-6, relatively large predorsal spots, while under the dorsal there are only two. Postdorsal spots generally 6-7, rarely fewer or more. All four Gambetta's zones are prksent. First and

11 COBITIDS OF GREECE 305 third zones, composed of small dots, have a normal extension. The second zone is formed by long lines. The fourth zone is also formed by lines, or very long spots or a nearly continuous band. In the last quarter of the body the first, second and third zones mix and form a network of dots and reticulations. Head with numerous small dots and a dark-gray band between the anterior rim of orbit and tip of snout. Dorsal and caudal fin with numerous gray dots. Other fins pale each having one row of minute dots. On the upper part of the base of the caudal fin a large jet black spot. Habitat: It is a typical still water living species, frequently all depths from very shallow to deeper places, especially where aquatic plants are present. From the general aspect of its type locality (Velestino spring) it can be assumed that the species does not avoid clean and slow running water. However, silty bottoms, into which it normally burrows, are the more favorite habitats. Range (Fig ): To this date the species has been found only in a very limited area, in the Kefalovriso karstic spring at Velestino (Feres) village in eastern Thessaly. This spring belongs to drainage of lake Karla (now dried up), from which the species was listed by Panagiotopoulos (1916) as Cobitis taenia. The species may still survive in the canals passing through the basin of the former lake. Remarks: This is an unusual species in its combination of characters of shape of body, squamation, suborbital spine, lamina circularis and especially in pigmentation. It is noteworthy that some specimens with somewhat similar colour pattern have been observed in a few Romanian rivers (see Bgngrescu & Nalbant, 1957). Etymology: The name of this species is given in memory of Dr. Alexander I. Stephanidis ( ), the first modern explorer of the Greek freshwater fish fauna. Subgenus Bicanestrinia Bscescu, 1962 Cobitis strumicae Karaman, 1955 (Fig. 7 a-f) Cobitis taenia, Kovatchev, 1921a: 88 (river Strymon), 1921 b. 93 (river Evros); Drensky. 1928: 167. fig (description, distribution: rivers Strymon. Nestos, Evros), 1930 (partim): 679 (list: rivers Strymon, Nestos, Evros), 1951 (partim): 124, fig. 80 (description; distribution: river basins of Mediterranean); Berg, 1932 (partim): (partim): 455 (from literature: rivers Evros to Strymon); Michajlowa. 1865a: 66 (Strymon). 1965b: 280 (Evros); Russev, 1966: 268 (Evros); Economidis, 1973 (partim): 471 (from literature: rivers Strymon, Nestos, Evros). 559 (from collections: lakes Koronia-Volvi and Vistonis, and rivers Strymon-Aggitis, Filiouris, Loutros and Evros), 1974: 91, map 25, fig. 27 (description, variation. distribution: all waters from Strymon to Evros. except stream Marmaras); Economidis & Sinis. 1982: 305 (lakes Koronia and Volvi). Cobitis taenia taenia. Drensky, 1929 (partim): 231 (list: rivers Evros, Nestos. Strymon).

12 306 P.S. ECONOMIDIS. T.T. NALBANT Cobitis taenia vardarensis, Taler, 1953 (partim): 447 (river Strymon drainge: tributary Strumica). Cobitis taenia strumicae Karaman, 1955: 190, fig. 4 (description as new sub-species, distribution: tributary Strumica, river Strymon drainage); Grupce & Dimovski, 1976: 34 (characters. comparisons with other species of South Yugoslavia). Cobitis pechevi Siskov & Dobrobolov, 1984: 1673, fig. 1-3 (description as new species. isoelectric focusing of muscle myogens; may be a subspecies of strumicae). Material: MZUT (23) , river Strymon, downstream of national roads's bridge near village Strymonikon, Macedonia, Greece, 31 August 1971, Economidis coll.; MZUT (11) , lake Volvi drainage, stream Rihios (Redina), Macedonia, Greece, 28 March 1978, Economidis coll.; MZUT (9) , river Nestos catchment, marsh near village Avramilia Kavalas, Macedonia, Greece, 6 July 1972, Economidis coll.; MZUT (18) , lake Vistonis, at shore near village Dialambi, Thrace, Greece, 5 July 1972, Economidis coll.; MZUT (29) , stream Filiouris, near village Karydia, Thrace, Greece, 4 July 1972, Economidis coll.; MZUT (241) , stream Bospos near Komotini town, Thrace, Greece, 20 August 1986, Economidis coll.; MZUT (15) , Stream near village Loutros Alexandroupolis, Thrace, Greece, 5 November 1971, Economidis coll.; MZUT (285) , river Evros catchment, tributary Erythropotamos near village Mikro Dereio, Thrace, Greece, 4 July 1972, Economidis coll.; ISBB 3971, (1) 94.0, same data as previous.; ISBB 3972, (20) , river Evros catchment, tributary near village Ardanio, Thrace, Greece, 5 October 1977, Economidis coll.; ISBB 3973, (14) , lake Volvi, Macedonia, Greece, 20 August 1971, Economidis coll.: ISBB 4579 (30) , stream Bospos, near Komotini town, Thrace, Greece, 20 August 1986, Economidis coll. Description: Fin ray formula. Number of rays generally constant in all fins: D I11 7, A , VI 6-1 6, P I 9-1 9, C n n. In few specimens there are 6 or 8 branched rays in the dorsal fin or 7 or 8 branched rays in pectorals. Karaman (1955) described the species without providing complete information on its morphology. Morphology. Body long, generally isomorphic (Fig. 7 a, b). Head moderately long, with snout relatively longer than postorbital space. Eyes normal in size, a little close to the hind margin of the head. The insertion of the dorsal fin is equidistant between the tip of snout and base of caudal fin. However there are specimens in which the predorsal distance is a little greater. Ventrals generally inserted on the same line of the dorsal fin but, in some specimens, these are inserted a little behind, under the second or third branched ray of dorsal. Caudal fin long enough. Mouth (Fig. 7 c) generally well arched. Upper (or anterior) lip moderately thick, unfurrowed although fine grooves can be seen

13 COBITIDS OF GREECE 307 sometimes. Lower (posterior) lip fleshy, wide enough and well folded. Mental lobes are shorter than the rest of the lip but moderately large and pointed. Barbels short. The suborbital spine (Fig. 7 e) is straight, with the longer thorn curved and the smaller about one third of the longer. The subdorsal scales (Fig. 7 d) are generally rounded somewhat wider at base and with a small eccentric focal zone. The lateral line is reduced, and does not exceed the length of pectoral fins. As in all species of Cobitinae the lateral line is superimposed in each side of the body on a small swelling which is probably in connection with the gas-bladder capsule. Sexual dimorphism (Fig. 7 f). Males with two laminae circulares, at base of first (unbranched) and second (branched) pectoral rays. Colourpattern (of live specimens or freshly preserved ones). Ground colour yellowish-white, laterally with an golden-green iridescent band from the tip of the snout to the caudal base. The dorsal part is light olive with 9-15 dark gray blotches. The four Gambetta's zones are complete and usually as in Fig. 3 b, e, rarely as in Fig. 3 a. In strumicae from the Evros basin, from the river Nestos and from the stream Bospos, the lateral spots or blotches are elongated or sometimes two round spots are united to form one elongate blotch. Generally the specimens of strumicae from lake Volvi and from river Strymon system have well defined spots along the sides, round or square-like. The upper part of base of caudal fin without a jet black dot or this dot is gray and reduced to a small point. Dorsal and caudal fins with small dark brown or gray dots. Other fins pale. Habitat: Over its very wide area of distribution this species presents a remarkable adaptation to different type of habitats with soft bottom. It can be observed from still water of lakes and river's death branch, with more or less deep water and rich vegetation, to slowly running and shallow water of streams. In these places the bottom change from muddy to silty or to sandy respectively, the vegetation being scarce or laking in the last case. In all these biotopes, the typical behavior of the fish, i.e. burrowing in the bottom, is always apparent, especially during summer when streams and death branch retain little water in shallows. Range: (Fig. 17-7): In Greece this species is dispersed in eastern Macedonia: river Strymon basin, lakes Volvi and Koronia, systems of the Thracian rivers Nestos, Bospos, Filiouris and Evros. Outside Greece Cobitis strumicae is present in the Strymon system (Yugoslavia and Bulgaria) and in the river Evros drainage of Bulgaria (as Cobitis pechevi) and probably in Turkey. It is also present in a few rivers in Bulgaria, tributaries of the Black Sea, but until now has not been found in the basin of the river Danube. Remarks: Obviously Cobitis strumicae in its elongated body and colour pattern is close to trichonica-meridionalis group, rather than to the

14 308 P.S. ECONOMIDIS. T.T. NALBANT helenica-arachthosensis-punctilineata complex. However, strumicae has the scales with a reduced focal zone, and is therefore quite different from the western species meridionalis and trichonica. Cobitis meridionalis Karaman, 1924 (Fig. 8 a-e) Cobitis taenia var. meridionalis Karaman, 1924: 75 (short description as new var., distribution: lake Prespa); Stephanidis, 1950: 208 (Vrigiis or Prespa lakes system). Cobitis taenia meridionalis. Taler. 1953: 447 (lake Prespa); Poljakov et a]., 1958: 103 (lake Prespa): Grupce & Dimovski, 1976: 30 (characters, comparison with other species of South Yugoslavia). Cobitis meridionalis, Economidis. 1993: 23 (full species rank). Material: MZUT (16) , lake Megali (Great) Prespa at village Psarades, Macedonia, Greece, 9 September 1988, Economidis coll.; ISBB 4568 (16) , bearing same data. Description: Fin ray formula. D I11 7, A I1 5, V I 6-1 6, P I 9-1 9, C n n, in a few specimens the number of dorsal branched rays is 6 and pectoral rays 8. Morphology. Body elongate, head long (Fig. 8 a). Mouth arched with the lips a little furrowed, especially the inferior lip which is folded (Fig. 8 b). Mental lobes well marked, in some specimens pointed. Suborbital spine straight with latero-caudal process very reduced (Fig. 8 d). Scales are subcircular (Fig. 8 c) with a large focal area. They are embedded and cover the whole body except the head. Sexual dimorphism: In males expresed as a thickened process (Fig. 8 e) at the base of the first an second pectorl rays, both being covered by skin. Generally the pectoral fins are longer in males than in females. The colour pattern is very characteristic in meridionalis (Fig. 8 a). The whole body is yellowish-white, with cloudy dark gray or olive blotches and dots. Back with 7-12 spots, generally rectangular. The second Gambetta's zone is formed by isolated spots, from the upper rear part of the operculum to the superior part of the caudal base. First and third Gambetta's zone is formed each by one row of speckles. The fourth Gambetta's zone is formed by rounded, relatively small spots. Lateral median intramuscular septum blackish, very visible. Upper half of head with small dots, without any dusky band. The dot in upper part of caudal base is very reduced and grayish. Dorsal fin with 3-4 rows of gray dots. Caudal fin with 4-5 irregular rows of graysh dots. Sometimes on the anal, ventral and pectoral fins there is one row of grayish dots. Iris golden. Habitat: The species is a typical still water fish, frequenting mainly lakes. It has also been observed in some short, stream-like still or slowly

15 COBITIDS OF GREECE 309 running water bodies, which over the year retain water intermittently, especially near to the mouths. However, as in the other close related species, this fish prefers habitats near the lake shore where muddy to silty bottoms and some vegetation are present. Range (Fig. 17-5): To date, Cobitis meridionalis is known only as endemic to lake Prespa basin. Remarks: Due to its pigmentation Cobitis meridionalis is well separated from the other members of Bicanestrinia. However, in its elongate body, lack of black spot at caudal base, presence of the black stripe on the intramuscular septum along the lateral midline of the body, this species appeas quite similar to Cobitis trichonica, a Bicanestrinia species with a western range in Balkan Peninsula too (see below). Both species may be holophyletic. Cobitis trichonica Stephanidis, 1974 (Fig. 9 a-3) Cobitis (Bicanestrinia) trichonica Stephanidis, 1974: 227, figs 1-3 (description as new species. distribution: lakes Trichonis and Lyssimachia). Material: MZUT , (10) , lake Trichonis catchment, tributary at village Panetolio near Malamatas fish farm, Etolia, Greece, 3 August 1988, Economidis coll.; ISBB 4578 (12) , bearing same data as previous; MZUT , (10) , lake Trichonis catchement, tributary at village Myrtia, Etolia, Greece, 4 August 1988, Economidis coll.; MZUT , (11) , lake Trichonis, Etolia, Greece, 15 September 1988, Economidis coll.; ISBB uncat., (13) , lake Ozeros catchment, channel to river Acheloos, Akarnania, Greece, 24 Sept. 1986, Economidis coll.; MZUT , (1) 28.6, river Acheloos catchment, spring at village Lesini, Etolia, Greece, 22 April 1992, Economidis coll.; MZUT , (3) , river Acheloos catchment, spring Agios Dimitrios Lesini, Etolia, Greece, 22 April 1992, Economidis coll. Stephanidis (1974) gave a good description of the species. In the following are added some details. Description: Fin ray formula. On most specimens it is as follows: D , A I1 5, V I 6-1 6, P , C n 7+7 n but a few individuals there are 8 branched rays in the dorsal and 7 or 9 branched rays in the pectoral. Morphology. Body relatively long, but in a few specimens, both males and females, it is short and stout. Head relatively short with an arched upper profile (Fig. 9 a). Eye moderately small and placed about midway between tip of snout and hind margin of operculum. Dorsal and ventrals generally equidistant between tip of snot and base of caudal.

16 310 P.S. ECONOMIDIS. T.T. NALBANT Mouth (Fig. 9 b) is arched enough with upper lip less furrowed but lower (or posterior) lip well folded with developed mental lobes, their tips being rounded rather than pointed. Barbels short. Suborbital spine (Fig. 9 d) is straight with well diverging caudal processes (thorms) but these being reduced enough. Scales (Fig. 9 c) are somewhat trapezoidal in shape with a large and eccentric focal zone. They cover entirely the body except the head. The lateral line, as in all Cobitinae loaches, is very reduced, not longer than the pectoral fins. Sexual dimorphism. In all males there are two laminae circulares at the base of the first (unbranched) and second (branched) pectoral rays, both processes being covered by a soft tissue and skin. In males pectcrals and ventrals are longer than in females. Colour pattern, in live specimens: the ground colour is yellowishwhite, sides of head and body having an iridescent or golden band. Dorsum olive with brown squareshaped spots. The fourth Gambetta's zones normally expressed as in Fig. 3 a. The colour of the dots and spots is dark brown or dark gray. In many specimens the fourth Gambetta's zone is rectangular and disposed vertically at least on caudal peduncle. The head is covered by dark gray or dark brown speckles. Generally the pigmentation of the head is similar to that shown in Fig. 2 c. At the base of the caudal fin there are two dark-gray spots. The inferior part of the caudal peduncle is sometimes pigmented with dark brown or grayish brown speckles. Dorsal and caudal fins with irregularly rows of dark brown dots, the rest of fins being whitish with generally one row of dark-brown dots. In many specimens the ground colour of the caudal fin is dull orange even reddish. Habitat: The species present a rather clear preference for still water, given that in the area of its distribution, i.e. the aquatic system of river Acheloos, it is more frequent in lakes. However, it has also been observed in slow to moderately fast running water, as in the Panetolion stream, flowing into lake Trichonis, and the canal uniting lake Ozeros and river Acheloos. In all cases the habitats are characterized by clean water of different depths, a more or less rich aquatic vegetation and a muddy to silty bottom in which the fish burrows. Range (Fig. 17-6): lake Trichonis and the adjacent lakes, Lysimachia, Ozeros, Amvrakia and also in the river Acheloos in Akarnania and Etolia, western Greece. Stephanidis (1974), did not found the species in lake Amvrakia. Nevertheless, the species has been observed once by one of us (PSE) near the shore of the lake, swimming among aquatic plants. Remarks: This species is very distinct in its pigmentary features. However, apparently it is close to strumicae and possibly meridionalis in its relatively long body and colour pattern.

17 COBITIDS OF GREECE 311 Cobitis hellenica sp. nov. (Fig. 10 a-f and 12 g-i) Cobitis taenia. Stephanidis, 1939: 32 (river Louros); Schmidt-Ries, 1943: 325 (from literature. river Louros). Material: Holotype: MZUT , (1) 69.3, river Louros catchment at Barbanakos spring near village Stephani, Epirus, Greece, 7 September 1989, Economidis coll. Paratypes: MZUT (22) , river Louros catchment at Barbanakos spring near village Stephani, Epirus, Greece, 1 August 1963, Economidis coll.; ISBB 3967, (13) , same locality as previous, 17 August 1963, Economidis coll.; MZUT (5) , same locality as previous, 19 October 1977, Economidis coll.; MZUT (25) , same data as previous; RMNH, uncat, (4) , same locality as previous, 25 August 1987, Economidis coll.; ZSM, uncat. (2) , same data as previous; CAS, uncat. (6) , same data as previous; ISBB 4577, (33) , same locality as previous, 23 September 1985, Economidis coll.; ISBB 4570, (11) same locality as previous, 28 August 1987, Economidis coll.; USNM , (2) same locality as previous, 25 August 1987, Economidis coll.; SMF (2), bearing same data as previous; MNHN , bearing same data as previous; NMW (1 & 2), bearing same data as previous; MZUT (34) , same locality as previous, 7 September 1989, Economidis coll.; MZUT (29) , river Louros catchment near village Philippiada, Epirus, Greece, 13 November 1979, Economidis coll.; ISBB 4569, (10) , river Louros catchment, near Hanopoulo village, Epirus, Greece, 7 August 1988, Economidis coll.; MZUT (15) , same locality as previous, 4 August 1988; MZUT (4) , river Louros at springs Agios Georgios, Epirus, Greece, 8 Sept. 1989, Economidis coll.; SMF (5) and MINB uncat., (2) all with same data as previous. Diagnosis: A stout bodied species, laterally compressed, and a high caudal peduncle. Colour based on five pigmentary zones. Description: Fin ray formula. D I11 7, A , V I 6-1 6, P I 9-1 9, C n n. On a few specimens we found D and even D (ISBB specimens from Barbanakos spring, possibly mutilated). A reduced number of rays in the pectoral, 8 and 7, are also present in some specimens all of which are males. Morphology. Body stout, very high in comparison to other species of Cobitis and well compressed (Fig. 10 a, b). Generally the body profile is isomorphic and are not taper from head to tail. The caudal peduncle is strongly laterally compressed. Head stout with moderate eyes which are sitdated generally close to snout than to the hind margin of head. Only in a

18 312 P.S. ECONOMIDIS. T.T. NALBANT few specimens are they situated in the middle of the head. Dorsal fin inserted a little behind the midpoint from tip of snout to caudal base. Ventrals generally inserted under second or third dorsal branched ray. Caudal enough, with rounded margin. Paired fins are short in adults. Mouth (Fig. 10 c) arched, with moderately thick and unfurrowed upper lip. Sometimes a few fine grooves can be seen on the surface of this lip. Lower lip fleshy, folded with well developed mental lobes. Barbels long enough. Suborbital spine (Fig. 10 e) relatively straight, the two thorns (caudal processes) being curved, the smaller about half the length of the other. Scales (Fig. 10 d) round, with a large focal zone, cover whole body excepting the head. The lateral line, being very reduced, does not exceed the length of the pectorals. Sexual dimorphism (Fig. 10 f). Males with two laminae circulares, one at base of first (unbranched) pectoral ray and another at the base of the second (branched) pectoral ray. Colour pattern. In preserved specimens, very characteristic of this species, the ground colour is yellow-brown or orange-brown. On the dorsum there are 4-6 large dark brown spots before the dorsal fin. In juvenile specimens these spots are smaller. Under the dorsal insertion there are 2, rarely 3, dark brown spots. In post dorsal there are 4-6 large dark brown spots rarely less or more. The first zone of Gambetta is formed by small and irregular brown speckles. The second row is formed by round brown dots. The third row, like the first, is also formed by brown spekles. All three rows of dots and speckles in the last fourth part of the body mix and form an area with reticulations or vermiculations. The fourth zone consists of dark brown spots, generally square-like or triangular. The upper part of this row is superimposed on a black or a deep bluish stripe. In most specimens, under the third row there is another row of very small brown dots, generally interrupted in the mid part of the body. Head with a reticulation of gray-brown spekles or minute dots. The dark band before the anterior rim of orbit is hardly discernible. Dorsal and caudal with irregular rows of dark brown spots. Other fins usually pale, but sometimes these are pigmented with small brown dots. Upper base of caudal with a very small dark gray dots. Habitat: As with many other loaches this species prefers still to moderate running water, better those of death branch of river, large ditches and springs. It frequents more or less deep water, where normally it burrows into the muddy to silty bottom. However, it can also be observed in shallow water but near aquatic plants, as in the Barbanakos spring. Range: (Fig. 17-1). This species is found only in western Greece in the river Louros tributary in Amvrakikos Gulf, to the Ionian Sea.

19 COBITIDS OF GREECE 313 Remarks: Of course, hellenica and closely related species, possessing this strange combination of characters (a Bicanestrinia group with deep and compressed body and well pigmented) represent an interesting presence in the freshwater of Greece. It is very difficult to refer the hellenica complex to another group of species in southern Europe, especially from the Balkan Peninsula. In the circum-mediterranean region there are another four deepbodied species of Cobitis, vettonica and maroccana from the waters of Spain and Morocco, zanandreai from southern half of Italy and turcica, in the Central part of Turkey.The first three species have only one lamina circularis, i.e. belong to Cobitis s. str. But turcica deserves special attention. The examination of many specimens from central Turkey (see above in Material and Methods) revealed that these series agrees with the equivocal original description by Hanko (1925: 154, fig. 8). But he does not give some inlormation about sexual dimorphism (if he had some males). The two turcica types from "Eregli", were lost by fire in October 1956 in the National Museum of Natural History in Budapest. However, Hanko gave interesting information about colour pattern of his turcica, concerning the presence of a supplementary row of speckles under the fourth row of Gambetta. He states that: "under the lateral rows of spots, between the pectorals and ventrals, on the yellow ground colour 5, there are some dark fine row of dots" (in original: "Unter den Seitenstreifen zwischen den Pectoralen und Ventralen auf gelber Grundfarbe 5, aus feinen und weningen dunklen Punkten bestehende Nebelflecke"). If turcica, from Eregli belongs to the subgenus Bicanestrinia, it is obvious that it represent the closest related species to the hellenica complex. Etymology: The name hellenica is derived from the name of the country (Hellas). Cobifis arachthosensis sp. nov. (Fig. 12 a-f) Material: Holotype: MZUT , 41.9, river Arachthos death branch near Akropotamia village Epirus, Greece, 4 August 1988, Economidis coll. Paratypes: MZUT , (2) , river Arachthos at Arta, Epirus, Greece, 20 October 1977, Economidis coll.; ISBB 4571, (2) , bearing same data as previous; MZUT , (5) , river Arachthos death branch near Akropotamia village Epirus, Greece, 4 August 1988, Economidis coll.; MZUT , (46) , bearing same data as previous; ISBB 4574, (10) , bearing same as previous; USNM , (1) bearing same data as previous; SMF (I), bearing same data as previous; MNHN , (2) bearing same data as previous; NMW (I), bearing same data as previous.

20 314 P.S. ECONOMIDIS. T.T. NALBANT Diagnosis: Similar in many respects to C. hellenica excepting young specimens in which the fourth zone of Gambetta is formed by round deep brown spots very remote from one another. The row with small dots of third zone of Gambetta is lacking. Description: Fin ray formula. D I11 7, A I11 5, V I 6-1 6, P I 9-1 9, C n n. Same variation in the number of dorsal and pectoral fin rays as in C. hellenica was observed. Morphology. The shape of the body and other characters as mouth, suborbital spine. scales, sexual dimorphism are exactly as in hellenica. Howevcr. in many adults the lateral dark brown spots (fourth Gambetta's zone) are well spaced and round. In all juveniles of arachthosensis the lateral spots are few (9-15, mainly 11-12) and remote from one another. Thc third Gambetta's zone is completely lacking. Habitat: As in the other loaches this is a typically benthic fish, presenting a remarkable adaption from still to moderately running water, and from clean to slightly turbid. It has been observed in the main bed of the river Arachthos, in a place with a more or less fast current, and in some of its death branches with a rather still water. Obviously it prefers biotopes with rich aquatic vegetation, but it may also found out of these. it also shows obvious variation of substrate preference, normally frequenting silty to muddy bottoms but also sandy or even gravely ones. The same variation has been observed in water depth, the fish living well in very shallow to deeper water. Range (Fig. 17-2): The area of this taxon covers the Arachthos basin, a flowing also as Louros, to the Amvrakikos Gulf and then to Ionian sea. Remarks: The main difference between arachthosensis and hellenica is in the pigmentation of juvenile specimens (compare Fig. 12 a-f with 12 g-i). This character is extremely important and can separate as good species arachthosensis from hellenica. The reduction of pigmentation represent an apomorphy. On the colour pattern this loach somewhat resembles another species of the subgenus Bicanestrinia, Cobitis linea, from the systems of rivers Kor and Kul, basin of the Arabian Sea, Iran (Bianco & Nalbant, 1980: 904, fig. 1 A). A recently discovered population from the river Thyamis (about 100 km northwestward), which is now under scrutiny, seems to agree well enough with those from river Arachthos. Initially both populations are regarded as conspecific. Etymology: The name was given after the river Arachthos in which this loach was found. Cobitis punctilineata sp. nov. (Fig. 11, Cobitis taenia. Economidis (partim): 91 (river Strymon drainage: Tributary Aggitis. color pattern and other features).

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