ArticlePDF Available

A revision of the Rutilus complex from Mediterranean Europe with description of a new genus, Sarmarutilus, and a new species, Rutilus stoumboudae (Teleostei: Cyprinidae)

Authors:

Abstract and Figures

Abstract By combining morphology, ecology, biology, and biogeography with the available molecular (sequence variation of the entire mitochondrial cytochrome b gene; cyt-b) and karyology data, the taxonomy of several species of the Rutilus complex inhabiting southern Europe is revised. Rutilus stoumboudae, new species, is described from Lake Volvi, Greece. It differs from Rutilus rutilus in possessing more total GR and less branched rays in both dorsal and anal fins and in its placement in the cyt-b based phylogeny of the genus. The resurrected genus Leucos Heckel, 1843 (type species Leucos aula, Bonaparte, 1841), which according to molecular data diverged from Rutilus more than 5 million years ago, during the Messinian salinity crisis, includes five species of small size, without spinous tubercles on scales and head in reproductive males, pharyngeal teeth formula 5-5, and all show a preference for still waters. Leucos aula is the Italian species endemic in the Padany-Venetian district: L. basak is widespread in Croatia, Albania, Montenegro and former Yugoslav Republic of Macedonia (FYROM); L. albus, recently described from Lake Skadar, Montenegro, is also found in rivers Moraca and Zeta (Montenegro). L. albus differs from L. basak, its closest relative, in having more scales on the LL and less anal-fin rays; L. panosi is endemic to the western-Greece district, and L. ylikiensis is endemic to lakes Yliki and Paralimni in eastern Greece (introduced in Lake Volvi). Among the nominal species examined, Rutilus karamani, R. ohridanus, R. prespensis and R. prespensis vukovici are all junior synonyms of Leucos basak. Rutilus vegariticus is definitively regarded as junior synonym for R. rutilus. Sarmarutilus n.gen. is a monotypic genus, with Sarmarutilus rubilio as the type species. According to phylogenetic data, Sarmarutilus rubilio is basal to a cluster of species that includes Leucos basak, L. albus, L. aula, L. panosi and L. ylikiensis. Sarmarutilus possibly evolved in pre-Messinian time, in the Lago Mare, entered the Mediterranean area during the Messinian Lago Mare phase of the Mediterranean Sea and survived only in the Tuscany-Latium district. This genus differs from Leucos in having large pearl organs on the central part of head and body scales in mature males and for the habitat preference, being a riverine-adapted species. It differs from Rutilus in pharyngeal teeth formula (5-5 in Sarmarutilus and 6-5 in Rutilus), size (small in Sarmarutilus and large in Rutilus) and for the preferential habitat (riverine vs. still water). Finally, lectotypes for Leucos basak, Leucos aula, and Sarmarutilus rubilio are designated. Key words: Freshwater fish, Mediterranean Europe, Cyprinidae, genera Rutilus, Leucos, Sarmarutilus new genus, new species
A) Lectotype of Leucos aula, 74 mm SL (ANSP 6434), Padua Province, northern Italy; B) Lectotype of Leucos basak, 117 mm SL (NMW 50723-1), Vergoraz and Lake Drusino near Imotsky, Croatia; C) Lectotype of Sarmarutilus rubilio, 93 mm SL (ANSP 6509), Lake Nemi, Central Italy. Examined materials. IZA 83120, 33, Croatia, R. Krupa (R. Neretva basin), P.G. Bianco, 28 May 1983.—IZA 048, 5, Croatia, L. Bačinska (R. Neretva basin), M. Mrakovic, 10 Aprile 1997.—IZA 00164, 5, FYROM, L. Ohrid, R. Gruptché, 10 April 1970.—IZA 0429, 30 (out of 122), FYROM; L. Ohrid, P.G. Bianco, 28–29 August 1987.—IZA 0430, 10, FYROM, L. Ohrid, R. Gruptché, 13 March 1969 .-IZA 85481, 30 (out of 232), Montenegro, L. Skadar, P.G. Bianco and B. Knezevic, 24–25 July 1984.—IZA 0441, 4, Montenegro, L. Skadar, P.G. Bianco and D. Marić, 9 November 1999.—IZA 0055, 5, FYROM: L. Prespa, P. Banarescu, 29 November 1975.—IZA 0435, 10, FYROM, L. Prespa, R. Gruptché, 15 May 1968.—IZA 0422, 12, Greece, lakes Prespa and Micraprespa, P.G. Bianco, 22 August 1998. – MNHN 1977.281, 3, Albania (syntypes of Rutilus karamanni Vladikov & Petit, 1930). Diagnosis. A species of Leucos characterized by moderate size, usually 120–140 mm SL; absence of a midlateral band; body uniformly silvery in life; peritoneal membrane black.; usually 36–38 pored scales on LL, as opposed to 38–43 in the other Leucos species. It may be distinguished from Leucos aula for the absence of lateral band and the color of the peritoneal membrane, black in L. basak, and silvery in L. aula. It differs from L. panosi and L. ylikiensis mainly by the number of GR, which are usually 9–10 in L. basak and respectively, 18–20 in L. ylikiensis and 13–14 in L. panosi. The most closely related species, according to molecular analyses (Fig. 1B) is L. albus. The two species can be identified on the basis of the number of LL scales, usually 41–42 in L. albus and 36–38 in L. basak, the number of D branched rays, 9 in L. basak and 8 in L. albus, and apparently for the color of the peritoneal membrane, nearly silvery in L. albus, and blackish in L. basak. Description. A small-medium sized species. In Croatia may reach 220 mm TL and 180 g of weight, but usually less than 150 mm TL (Mačrovcić et al., 2006). Body uniformly silvery without longitudinal band; color of
… 
Content may be subject to copyright.
Accepted by L. Page: 26 Jun. 2014; published: 28 Jul. 2014
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2014 Magnolia Press
Zootaxa 3841 (3): 379
402
www.mapress.com
/
zootaxa
/
Article
379
http://dx.doi.org/10.11646/zootaxa.3841.3.4
http://zoobank.org/urn:lsid:zoobank.org:pub:E52D2F6B-631D-45FE-97E0-DD26137757FE
A revision of the Rutilus complex from Mediterranean Europe with description
of a new genus, Sarmarutilus, and a new species, Rutilus stoumboudae
(Teleostei: Cyprinidae)
PIER GIORGIO BIANCO
1
& VALERIO KETMAIER
2,3
1
Department of Biological Sciences, Zoological Section, Via Mezzocannone, 8, University of Naples “Federico II”, I-80134 Napoli,
Italy. E-mail: gibianco@unina.it
2
Unit of Evolutionary Biology/Systematic Zoology, Institute of Biochemistry and Biology, University of Potsdam, Karl-Liebknecht-
Strasse 24-25, D-14476, Potsdam, Germany;
3
Department of Biology and Biotechnology “Charles Darwin”, University of Rome “Sapienza”, V.le dell’Universita’ 32, 00185,
Rome, Italy. E-mail:ketmaier@uni-potsdam.de
Abstract
By combining morphology, ecology, biology, and biogeography with the available molecular (sequence variation of the
entire mitochondrial cytochrome b gene; cyt-b) and karyology data, the taxonomy of several species of the Rutilus com-
plex inhabiting southern Europe is revised. Rutilus stoumboudae, new species, is described from Lake Volvi, Greece. It
differs from Rutilus rutilus in possessing more total GR and less branched rays in both dorsal and anal fins and in its place-
ment in the cyt-b based phylogeny of the genus. The resurrected genus Leucos Heckel, 1843 (type species Leucos aula,
Bonaparte, 1841), which according to molecular data diverged from Rutilus more than 5 million years ago, during the
Messinian salinity crisis, includes five species of small size, without spinous tubercles on scales and head in reproductive
males, pharyngeal teeth formula 5-5, and all show a preference for still waters. Leucos aula is the Italian species endemic
in the Padany-Venetian district: L. basak is widespread in Croatia, Albania, Montenegro and former Yugoslav Republic
of Macedonia (FYROM); L. albus, recently described from Lake Skadar, Montenegro, is also found in rivers Moraca and
Zeta (Montenegro). L. albus differs from L. basak, its closest relative, in having more scales on the LL and less anal-fin
rays; L. panosi is endemic to the western-Greece district, and L. ylikiensis is endemic to lakes Yliki and Paralimni in east-
ern Greece (introduced in Lake Volvi). Among the nominal species examined, Rutilus karamani, R. ohridanus, R. pre-
spensis and R. prespensis vukovici are all junior synonyms of Leucos basak. Rutilus vegariticus is definitively regarded
as junior synonym for R. rutilus. Sarmarutilus n.gen. is a monotypic genus, with Sarmarutilus rubilio as the type species.
According to phylogenetic data, Sarmarutilus rubilio is basal to a cluster of species that includes Leucos basak, L. albus,
L. aula, L. panosi and L. ylikiensis. Sarmarutilus possibly evolved in pre-Messinian time, in the Lago Mare, entered the
Mediterranean area during the Messinian Lago Mare phase of the Mediterranean Sea and survived only in the Tuscany-
Latium district. This genus differs from Leucos in having large pearl organs on the central part of head and body scales in
mature males and for the habitat preference, being a riverine-adapted species. It differs from Rutilus in pharyngeal teeth
formula (5-5 in Sarmarutilus and 6-5 in Rutilus), size (small in Sarmarutilus and large in Rutilus) and for the preferential
habitat (riverine vs. still water). Finally, lectotypes for Leucos basak, Leucos aula, and Sarmarutilus rubilio are designat-
ed.
Key words: Freshwater fish, Mediterranean Europe, Cyprinidae, genera Rutilus, Leucos, Sarmarutilus new genus, new
species
Introduction
Since the work of Kottelat (1997), which presented a general overview of the systematics of European cyprinids as
well as of the many still open questions concerning their taxonomy, a wealth of molecular studies has been
published on different genera to shed light on the evolution of this speciose family in the area (Squalius: Durand et
al., 1999; Te le st es and Scardinius: Ketmaier et al., 2004; Barbus: Tsigenopoulos & Berrebi, 2000, Bianco, 1998).
Bianco & Taraborelli (1985) and Bogutskaya & Illiadou (2006) morphologically analyzed a number of European
BIANCO & KETMAIER
380
·
Zootaxa 3841 (3) © 2014 Magnolia Press
species of the genus Rutilus. In particular, Bogutskaya & Illiadou (2006) split the genus into two subgenera; Rutilus
s.str., which includes species of southern European origin and Pararutilus, limited to central and eastern Europe.
Kottelat & Freyof (2007) did not consider this arrangement as valid and proposed a new classification. Rutilus,
widespread in Europe but absent in the Iberian Peninsula, would include 16 species (one not described) from three
distinct areas: five species from central Europe and from drainages in the Black and Caspian Sea regions; three
from the Italian peninsula and seven species from the Balkan peninsula. Two species, Rutilus rutilus (Linnaeus,
1758), and R. frisi (Nordmann, 1840) were reported for more than one area. The situation is particularly complex
for the three largest lakes of the Balkan peninsula (Ohrid, Prespa and Skadar) where alternatively one, R. ohridanus
(Karaman, 1924), or three, R. karamani Valdikov & Petit, 1930, R. ohridanus and R. prespensis (Karaman, 1924)
species have been described. Recently, Marić (2010) described an additional new species, R. albus Marić, 2010
from Lake Skadar (Montenegro). Ketmaier et al. (2008) analyzed sequence variation in the mitochondrial DNA
(mtDNA) region coding for the cytochrome b (cyt-b) gene to assess phylogenetic relationships among 12 of 14
species at that time ascribed to the genus and reported in FishBase (http://www.fishbase.org). Molecular data were
in agreement with Kottelat & Freyof (2007) in rejecting the subdivision of Bogutskaya & Illiadou (2006) in two
subgenera but did not support a specific distinction for Rutilus basak/R.prespensis or for R.vegariticus/R.rutilus as
proposed by Kottelat and Frehyof (2007).
Here, we take advantage of the extensive availability of Rutilus complex specimens in the collection of the first
author to re-analyze the morphology of several species with special attention to those from the western Balkan
area. Morphological results are then discussed in light of the existing cyt-b phylogenetic hypothesis (Ketmaier et
al., 2008) to derive a reliable taxonomic arrangement. Three distinct groups of Rutilus complex species occur in the
study area. The first group, well differentiated at both molecular and morphological levels, includes the resurrected
genus Leucos with five species: Leucos aula (the type species), L. basak, L. panosi, L. albus and L. ylikiensis. The
biogeography of this genus mirrors that of the genus Teles t e s (Ketmaier et al., 2004). We show how both
morphology and molecules confirm Leucos albus as a valid taxon and identify a new species of Rutilus s.str. from
Lake Volvi (Greece). Indeed, according to Economidis & Sinis (1981), the Rutilus rutilus reported for Lake Volvi
and some other eastern Greek basins would represent a distinct taxon. Our molecular and morphological data agree
with this opinion. In examining the material at our disposal we found two species of the Rutilus complex co-
occurring in Lake Volvi. One shows a remarkable genetic distance from the other allegedly conspecific populations
of R. rutilus (Ketmaier et al., 2008); this entity is described below as a new species. The second species is Leucos
ylikiensis, which was very likely introduced in the basin from lakes Yliki and Paralimni as reported for Scardinius
graecus Stephanidis, 1957 (Bianco & Kottelat, 2005). It should be noted that prior to translocations, Leucos
ylikiensis and Scardinius graecus were endemic to lakes Yliki and Paralimni, respectively (Bianco & Kottelat,
2005). Finally we describe the new, monospecific genus Sarmarutilus to include S. rubilio. According to
phylogenetic data, Sarmarutilus rubilio is basal to a cluster of species that includes Leucos basak, L. albus, L. aula,
L. panosi and L. ylikiensis. This genus probably diverged from its central European progenitor after the Messinian
salinity crisis.
Further, given the overall instability of the nomenclature of the genus, we designate lectotypes for Leucos
basak (Heckel, 1843), Leucos aula (Bonaparte, 1841) and Sarmarutilus rubilio (Bonaparte, 1838).
Material and methods
Localities of material examined from Italy, Croatia, Montenegro, Greece, and FYROM are shown in Fig. 1A. Our
sampling covers the Danubian (for comparisons), Padany-Venetian, Tuscany-Latium, Dalmatian, Albanian,
western Greek and Aegeo-Macedo-Anatolian ichthyogeographic districts defined in Bianco (1990).
Most of the individuals examined in the study are housed in the first author’s collection at the Department of
Biological Sciences, University of Naples “Federico II”, Naples, Italy (IZA). Abbreviations for additional
institutions where the examined specimens are deposited are: ANSP, Academy of Natural Sciences of Philadelphia,
USA; MNHNP, Muséum National d’Histoire Naturelle, Paris, France; MSNM, Museo Civico di Storia Naturale di
Milano, Italy; MCCI, Museo Civico di Scienze Naturali di Carmagnola, Torino, Italy; NMW, Naturhistorisches
Museum,Vienna, Austria. Abbreviations for the examined morphological characters are the following: LL, lateral
line; D, dorsal fin; A, anal fin; P1, pectoral fin; P2, pelvic fin; GR, gill rakers; SL, standard length; TL, total length.
Zootaxa 3841 (3) © 2014 Magnolia Press
·
381
NEW GENUS AND SPECIES FROM MEDITERREAN EUROPE
Counts and measurements follow Bianco & Kottelat (2005). Part of the material included in this study had been
analyzed molecularly in Ketmaier et al. (2008). Further details on the examined materials are given in each species
description. Among the examined material and for every species of small size, proportional measurements include
only specimens of comparable standard lengths as they are subjected to environmental influence and growth
allometries. From the extensive collection of samples of Sarmarutilus rubilio from Italy and Leucos basak from
lakes Skadar and Ohrid at our disposal, only a part has been used for comparisons with others species. The fishes
were dissected and examined internally to check the color of peritoneal membrane and maturity of gonads. The age
has been determined by a scalimetric technique (Bianco & Taraborelli, 1985). The synonyms of Rutilus rutilus
discussed are for nominal taxa of incertae sedis. For a complete list of synonyms of this species see Kottelat
(1997). The complete list of synonyms for Leucos aula, Sarmarutilus rubilio and Rutilus pigus are reported in
Bianco & Taraborelli (1985).
FIGURE 1. A) Collecting localities (L.=lakes, R.=rivers) of Rutilus complex species. 1, R. Thames; 2, R. Danube (Slovakia);
3, R. Danube (Romania); 4, L. Como; 5, L. Maggiore; 6, R. Arno and R. Serchio; 7, R. Albegna and R. Cecina; 8, R. Ombrone
and R. Fiora; 9, R. Tiber and R. Mignone; 10, L. Trasimeno; 11, R. Liri and R. Garigliano; 12, R. Volturno and R. Sele; 13, R.
Pescara and R. Tavo; 14, R. Trigno; 15, R. Tronto; 16, R. Livenza; 17, L. Pernica; 18, R. Sava; 19, L. Drusino; 20, R. Krupa (R.
Neretva basin); 21, L. Besanka (R. Neretva basin); 22, R. Moraca and R. Zeta; 23, L. Skadar; 24, L. Dojran; 25, L. Horid; 26, L.
Prespa and L. Micraprespa; 27, L. Vegoritis; 28, L. Volvi; 29, L. Joannina; 30, L. Yliki and L. Paralimni; 31, L. Ambrakia; 32,
L. Trichonis. Bold numbers are the different peri-Mediterranean ichthyogeographic districts described in Bianco (1990). B) A
schematic temporal reconstruction of the diversification of the genus in southern Mediterranean Europe based on cyt-b is
shown (modified from Ketmaier et al., 2008). For each taxon the corresponding number of the district of origin is shown;
triangles indicate that multiple populations were analyzed. Numbers at nodes are divergence times in million years; all labeled
nodes were robustly supported in the phylogenetic reconstructions (see Ketmaier et al., 2008 for further details).
BIANCO & KETMAIER
382
·
Zootaxa 3841 (3) © 2014 Magnolia Press
Systematic accounts
Among the species of the Rutilus complex listed in Kottelat & Frehyof (2007), several do not deserve further
discussion as they can be unequivocally distinguished. This is not the case for several nominal taxa described from
the western Balkans. According to the molecular data of Ketmaier et al. (2008), there is a group of five species
native to the area representing distinct evolutionary lineages possibly evolved during the Lago Mare phase of the
Mediterranean Sea. These species are placed in the resurrected genus Leucos Heckel, 1843. One species, R. rubilio,
does not cluster with any other lineage and is placed in a distinct new genus: Sarmarutilus n.g. Rutilus pigus,
endemic to the Padany-Venetian district, clearly belongs to the lineage of Rutilus s.str. when compared with R.
rutilus from Danubian region and specimens introduced in the study area. According to Economidis & Sinis
(1981), the R. rutilus distributed in Lake Volvi and adjacent eastern Greek basins should be regarded as a distinct
taxon. Genetically, the population from Lake Volvi is clearly differentiated from the other allegedly conspecific
populations (Ketmaier et al., 2008). The morphological data presented here further support this view (Tables 4, 5),
and a formal description is given below. From the rivers Moraca and Zeta (Montenegro), there is a recently
described species that, according to molecular and morphological data would belong to the Leucos lineage: L.
albus (Marić, 2010), close relative of L. basak Heckel, 1843.
Genus Leucos Heckel, 1843
Type species: Squalius aula Bonaparte, 1841: Fauna It., fasc. XXX, Tav. 116, fig. 3: type locality; Padua Province;
north-eastern Italy.
Leucos cisalpinus Heckel, 1843: 1081 (Lake Garda) was the subsequent type species designation by Jordan &
Gilbert (1883). But it was a nomen nudum. Type species by present designation: Squalius aula Bonaparte, 1841:
type locality; Italy, Padua Province, north-eastern Italy. It is the only species from Lake Garda, common in every
lake of northern Italy, which corresponds with the short description below of the genus. A short diagnosis
(translated from Latin) of this genus was given by Heckel, (1843:1081): pharyngeal teeth, 5-5; mouth small; lips
smooth; barbels absent; dorsal and anal fins short; origin of D at level of insertion of pelvic fins. Others species
placed by Heckel (1843) in this genus: Leucos rubella Heckel, 1843 (nomen nudum); Leucos basak Heckel, 1843;
Leucos (now placed in Delminichthys) adspersus Heckel, 1843, Leuciscus selysii Selys-Longchamps, 1841 (junior
synonym of Rutilus rutilus); and Leuciscus rutiloides Selys-Longchamps, 1842 (junior synonym of R. rutilus). The
genus Leucos was used by several authors until it was synonymized with Rutilus (Howes, 1981).
Diagnosis. Distinguished from others genera of European cyprinids by a combination of the following
characters: pharyngeal teeth 5-5 (rarely 4-5 or 5-4 or 6-5) hooked and slightly serrated. Mouth small, terminal or
slightly upturned or downturned; origin of D nearly at same level of origin of P2; free margin of D and A concave
or slight concave; caudal fin from moderately to deeply forked; peritoneal membranae from silvery to blackened by
fused melanophores; body without longitudinal stripes (excepted Leucos aula). GR short. Usually, 8–9 branched
rays in the D and in the A; 8 branched rays in the P2; 12–14 circum-peduncular scales; absence of large pearl organ
(nuptial tubercles) on head and central part of scales in reproductive males; small size, did not exceed 180 mm SL;
mostly a still water species. Sequence variation of the entire mitochondrial cytochrome b gene shows a strictly
relationship among five species, which separated from a Leuciscinae ancestor about 4-5 milion years ago, during
the Lago Mare phase of the Mediterranean (Bianco, 1990), probably following the same evolutionary trajectories
of the genus Tele s t es, which occurs in the same geographic area (Ketmaier et al., 2004).
Leucos aula (Bonaparte, 1841)
(Fig. 2A, 3A)
Squalius aula Bonaparte, 1841: Fauna It., fasc. XXX, Tav. 116, fig. 3: type locality; Padua Province; north-eastern Italy
Examined material: All from Italy: ANSP 6434-6445, 12 (types of Squalius aula Bp), ANSP 6467-6468, 2 (types
of S. elatus Bp), NE Italy.—IZA 8361, 2, R. Sile at Jesolo, G. Delmastro, 10 August 1980.—IZA 8361, 14, small
Zootaxa 3841 (3) © 2014 Magnolia Press
·
383
NEW GENUS AND SPECIES FROM MEDITERREAN EUROPE
canal near Jesolo, G. Delmastro, 10 August 1980.—IZA 83138, 12, R. Tartaro near Lazise, C. Oppi, 20 May 1975.-
IZA 83139, 6, R. Tartaro near Isola della Scala, C. Oppi, 20 November 1978.—IZA 83140, 4, Le Porte del
Menago, Verona, G. Togni, 1 May 1978.—IZA 8412, 25, R. Tartaro, Isola della Scala, C. Oppi, May 1978.—IZA
8345, 14, L. Major, P.G. Bianco, 18 April 1982.—IZA 8365,10, R. Po near Cuneo, G. Delmastro, 8 August
1980.—IZA 8368, Canal Moneta (R. Po basin), 10, G. Delmastro, 5 October 1977.—IZA 8377, 8, Rio Stellone (R.
Po basin), G. Delmastro, 22 May 1980.—IZA 8349, 10, L. Piediluco near Terni, P.G. Bianco, 1 May 1983
(introduced).—IZA 8372, 9, L. Monticchio, Basilicata, P.G. Bianco, 18 June 1977 (introduced).—IZA 0198, 2, L.
Massaciuccoli near Pisa, E. Baldaccini, 7 April 2001(introduced).—IZA 8837, 14, R. Foglia, P.G. Bianco, 22 June
1988.—IZA 0425, 7, R. Bacchiglione, 4 April 1996, E. Marconato.
FIGURE 2. Native species of the Rutilus complex in southern Mediterranean Europe: A) Leucos aula, 115 mm SL, R.
Bacchiglione, northern Italy, April 1996 (IZA 0425); B) Leucos basak, 95 mm SL, Lake Ohrid, FYROM, August 1987, (IZA
0429); C) Leucos ylikiensis, 120 mm SL, Lake Yliki, April 1987 (IZA 8733); D) Leucos panosi, 130 mm SL, canal to Lake
Trichonis, May 1987 (IZA 8766); E) Leucos albus, 98 mm SL, River Moraca, Montenegro, November 1999 (IZA 0421); F)
Sarmarutilus rubilio, 100 mm SL, River Ofanto, southern Italy, February 2002 (IZA 02183); G) Rutilus pigus, 230 mm SL,
Lake Major, northern Italy, (IZA 0474); H) Holotype of Rutilus stoumboudae new species, 148 mm SL, October 1998, Lake
Volvi, Greece (IZA 02107).
Diagnosis. The species can be distinguished from all others species of Leucos for the presence of a middle
lateral band, and a smaller head length, which in fish of comparable size is less than 4.5 times in the SL, except L.
panosi, and more than 4.0 times in the others species. Additionally, L. aula differs from in the modal number of
circum-peduncular scales, which is 12 as opposed to 14 in other species.
Description. General appearance in Fig. 2A. A small-medium sized, not exceeding 180 mm, usually 80–120
mm SL. A species of Leucos characterized by a quite marked mid-lateral band, extending from the operculum
(sometimes from the tip of the snout) to the end of the caudal peduncle: on caudal peduncle the band is sometimes
BIANCO & KETMAIER
384
·
Zootaxa 3841 (3) © 2014 Magnolia Press
enlarged to form an oval spot; body moderately elongate, dorsal profile convex; sometimes a small hump in largest
specimens; posterior mouth corner placed beyond the vertical crossing the anterior margin of orbital cavity; snout
blunt; mouth opening oblique and terminal to slightly sub-terminal; lips smooth; horizontal diameter of eye, less or
equal than pre-orbital length; head small, its length about 3.8–4.2 times in the SL; body quite deep, its depth about
2.6–3.0 times in the SL; origin of the D at same level or slightly below the insertion of the P2; free margin of D and
A, slightly concave; caudal fin moderately forked; LL complete, with 35–42 pored scales, placed below the middle
of the body, slightly concave and extending from the upper margin of opercular membranae, to the end of caudal
peduncle; modally 3 un-branched and 9 branched rays both in the D and A; P1 with 1 un-branched and 14–15
branched rays; P2 with 1 un-branched ray and constantly 8 branched rays; about 8–10 total GR; color of eye in life
from yellowish to a brilliant red; fins yellowish; peritoneal membranae silvery sometimes with few scattered
melanophores. Adult reproductive males may have minute, granular tubercles on head, check, upper side of paired
fins and fan-shaped on the free border of scales of the body. For additional description and shape, see Fig. 2A and
Tables 1, 2.
Distribution. The species was endemic in the Padany-Venetian ichthyogeographic district, including all the
river basins tributary of the upper Adriatic Sea, from Croatia, in basins near the town of Zara (Mačrovcić et al.,
2006), Slovenia and down, in Italy, to the River Reno in the Marche Region. It was introduced in several basins of
central and southern Italy: at least lakes of Monticchio, Massaciuccoli, Bracciano, Piediluco and rivers Tiber, Arno,
Mignone (Bianco & Taraborelli, 1985; Bianco & Santoro, 2011; P.G. Bianco, pers observ).
Lectotype designation for Leucos aula (Bonaparte, 1841). The syntypes series of Leucos aula (Bomparte’s
original catalogue number 438) in ANSP includes two species: 11 individuals are L. aula (ANSP 6434-6444,
55–101 mm SL), and 1 is Scardinius hesperidicus (ANSP 6445, 43.0 mm SL). Among the 11 syntypes, the
specimen ANSP 6434, 74 mm SL, in very good condition, is designated as lectotype (Fig. 3A). It has 39 scales on
the LL; 9 branched rays in D and 9 in A; 7.5 rows of scales above LL and 3.5 below LL; total GR, 9; 14 circum-
peduncular scales; pharyngeal teeth formula, 5-5. The 10 paralectotypes, ANSP 6435-6444, 43–101 mm SL, show
the following ranges of meristic counts: LL, 38–40; 7.5–8.5 rows of scales above and 2.5–3.5 below LL; 9–11 total
GR. D constantly with 9 branched rays; A with 9–11 branched rays (modal value, 9); 12–14 circum-peduncular
scales; 5-5, 5-4 pharyngeal teeth. The single specimen of Scardinius hesperidicus has 38 scales on LL; 9 branched
rays in D, 11 in A, 14 circum-peduncular scales and 14 total GR. Pharyngeal teeth formula: 5.3-3.5.
Remark on the Bonaparte fish collection. Several question were raised about the value of the syntypes of the
Bonaparte collection housed in ANSP and catalogued by Böhlke (1984) by Dr. Maurice Kottelat (pers. comm.).
Kottelat’s point of views were followed by several ichthyologists and, among them, one of the reviewers of the first
version of this contribution, originally submitted to Zootaxa. The study was rejected mainly on the grounds that
there are no proof that syntypes of Bonaparte housed in ANSP were used for the original description. But this
controversy was recently solved and the collection of Bonaparte’s new cyprinid species are confirmed to be
syntypes (Bianco, 2014a).
Remarks on ecology, biology and conservation. Preferential habitats: lakes and still waters of rivers. Not
thriving in moderately to fast flowing rivers. This species strongly competes and tends to eliminate Sarmarutilus
rubilio in still waters. It is responsible for the extinction of S. rubilio in central and southern Italian lakes and rivers
(Bianco & Ketmaier, 2001). The reproductive season extends from April to August, peaking in May–June. Age at
first reproduction is 1+ or 2+ for males and 2+ for females. Maximum age observed, 7+ (Bianco & Taraborelli,
1985). Its conservation status is of “Low Concern” according to the IUCN (2013) red list.
Leucos basak Heckel, 1843
(Figs. 2B, 3B)
Leucos basak Heckel, 1843: 1006 (Type locality: Vergoraz and Lake Drusino, near Imotschi, Croatia).
Rutilus aula karamani Vladikov & Petit, 1930: 391 (type locality, Lake Ohrid, Albania). The authorship of this species is
reported as Fowler, 1977 in Kottelat (1997) because Kottelat assumed that the original description was infrasubspecific
(Rutilus aula natio karamani), which should not be made available (ICZN, 1999; art. 45.5). It should be noted, however,
that Vladikov & Petit (1930) in the text of their original description used the name Rutilus aula karamani, without the use
of “natio”, making clear their intention to describe a subspecific taxon (ICZN, 1999: art. 45.6.4.1).
Leucos aula var. ohridana Karaman, 1924: 56 (type locality, Lake Ohrid, FYROM)
Zootaxa 3841 (3) © 2014 Magnolia Press
·
385
NEW GENUS AND SPECIES FROM MEDITERREAN EUROPE
Leucos aula var. prespensis Karaman, 1924: 57 (type locality, Lake Prespa, FYROM)
Rutilus prespensis vukovici Marić, 1989: 65 (type locality, Lake Skadar, Montenegro)
FIGURE 3. A) Lectotype of Leucos aula, 74 mm SL (ANSP 6434), Padua Province, northern Italy; B) Lectotype of Leucos
basak, 117 mm SL (NMW 50723-1), Vergoraz and Lake Drusino near Imotsky, Croatia; C) Lectotype of Sarmarutilus rubilio,
93 mm SL (ANSP 6509), Lake Nemi, Central Italy.
Examined materials. IZA 83120, 33, Croatia, R. Krupa (R. Neretva basin), P.G. Bianco, 28 May 1983.—IZA 048,
5, Croatia, L. Bačinska (R. Neretva basin), M. Mrakovic, 10 Aprile 1997.—IZA 00164, 5, FYROM, L. Ohrid, R.
Gruptché, 10 April 1970.—IZA 0429, 30 (out of 122), FYROM; L. Ohrid, P.G. Bianco, 28–29 August 1987.—IZA
0430, 10, FYROM, L. Ohrid, R. Gruptché, 13 March 1969 .- IZA 85481, 30 (out of 232), Montenegro, L. Skadar,
P.G. Bianco and B. Knezevic, 24–25 July 1984.—IZA 0441, 4, Montenegro, L. Skadar, P.G. Bianco and D. Marić, 9
November 1999.—IZA 0055, 5, FYROM: L. Prespa, P. Banarescu, 29 November 1975.—IZA 0435, 10, FYROM,
L. Prespa, R. Gruptché, 15 May 1968.—IZA 0422, 12, Greece, lakes Prespa and Micraprespa, P.G. Bianco, 22
August 1998. – MNHN 1977.281, 3, Albania (syntypes of Rutilus karamanni Vladikov & Petit, 1930).
Diagnosis. A species of Leucos characterized by moderate size, usually 120–140 mm SL; absence of a mid-
lateral band; body uniformly silvery in life; peritoneal membrane black.; usually 36–38 pored scales on LL, as
opposed to 38–43 in the other Leucos species. It may be distinguished from Leucos aula for the absence of lateral
band and the color of the peritoneal membrane, black in L. basak, and silvery in L. aula. It differs from L. panosi
and L. ylikiensis mainly by the number of GR, which are usually 9–10 in L. basak and respectively, 18–20 in L.
ylikiensis and 13–14 in L. panosi. The most closely related species, according to molecular analyses (Fig. 1B) is L.
albus. The two species can be identified on the basis of the number of LL scales, usually 41–42 in L. albus and
36–38 in L. basak, the number of D branched rays, 9 in L. basak and 8 in L. albus, and apparently for the color of
the peritoneal membrane, nearly silvery in L. albus, and blackish in L. basak.
Description. A small-medium sized species. In Croatia may reach 220 mm TL and 180 g of weight, but
usually less than 150 mm TL (Mačrovcić et al., 2006). Body uniformly silvery without longitudinal band; color of
BIANCO & KETMAIER
386
·
Zootaxa 3841 (3) © 2014 Magnolia Press
the eye in living animals is yellowish; fins yellowish or pale grayish in preserved specimens; snout pointed, pre-
orbital distance near equal to the horizontal diameter of the eye; lips smooth; mouth opening oblique, the corner of
maxillae at the same level of the vertical crossing the anterior border of the orbit; mouth terminal or slightly
inferior; profile of dorsum convex sometimes humped in large specimens; paired and un-paired fins yellowish with
few scattered melanophores, free margin of D and A concave; P1 longer in males, were may reach the origin of P2;
caudal fin forked; peritoneal membrane blackened by several melanophores addensed and fused; head length about
3.6–3.8 times in the SL; body deep about 3.5–3.7 times in the SL; origin of the D at same level of the insertion of
the P2; LL complete, and extending from the margin of opercular membrane, to the end of caudal peduncle, with
35–41 pored scales, usually 36–38; 7.5 above and 3.5 below the LL; constantly 3 un-branched rays followed by 9
branched rays in the D; constantly 3 un-branched and usually 8 branched rays in the anal fin; P1 with 1 un-
branched and 14–15 branched rays; P2 with 1 un-branched ray and constantly 8 branched rays; usually 9–10 total
GR; in adult males pearl organ absent. For additional description and shape, see Fig. 2B and Tables 1, 2, 3.
Distribution. The species distribution includes lakes and mostly still water of rivers of the Adriatic drainages,
from Croatia, till to Albany, and Montenegro. In Lake Skadar the species is sympatric with Leucos albus. In
Croatia its distribution is limited to basins of south-eastern part: lakes Crven, Modro, Bačinska and rivers Matica
and Neretva (Mracovčić et al., 2006).
Remarks on synonyms. According to mtDNA (Ketmaier et al., 2008) and to the morphological analyses
presented here on Leucos basak (Table 3) from the Dalmatian, Albanian, and Aegean ichthyogeographic districts
(sensu Bianco 1990), the populations from lakes Ohrid, Prespa and Skadar and River Krupa (River Neretva
drainage) cannot be unequivocally distinguished from one another and should be regarded as a single species. This
would be partially at odds with the results of Milošević et al. (2011). These authors, based on mitochondrial (cyt-b)
and nuclear (five polymorphic microsatellites) markers coupled with morphology proposed the existence of three
species in the area (Rutilus ohridanus, R. prespensis and R. albus). It should be noted, however, that divergence at
the mtDNA level was shallow (up to a maximum of five substitutions) and that the lack of gene flow at the nuclear
loci could reflect the geographical isolation of these lakes rather than a specific status of the taxa. The authors
themselves claimed that “R. prespensis should be further evaluated to determine if the taxon is synonymous with
other Rutilus from the Adriatic basin”.
Rutilus ohridanus is a problematic species as the diagnosis of meristic characters given in Karaman’s (1924)
original description (D with 10–11 branched rays, A with 11 branched rays and 44 LL scales) overlaps with that
given for R. pygus virgo for Lake Dojran (FYROM, Greece) in the very same paper. The overlapping in biometric
characters with R. pygus virgo is very likely due to a printing error; also the maximum size reported Karaman
(1924), (about 150 mm TL) would classify the species as a small sized Rutilus (Ivanović, 1973; Soric, 1983; Marić,
1988), (see Tab 2). In our extensive survey of samples of R. basak (see above) we found neither 10/11 branched
rays in D nor 10 in A. R. ohridanus should be then placed as synonym for R. basak. R. prespensis, R. p. vukovici
and R. karamanni, cannot be separated from R. basak neither molecularly nor morphologically and are here
regarded as junior synonyms for R. basak. Thus all the populations of the synonymized nominal taxa represent a
single species; this would also reflect the hydrography of the Albanian district as these lakes are connected to one
another: the lakes Prespa and Ohrid through a subterranean karstic channel, lakes Ohrid and Skadar through the
Bojana-Drina river system. Lake Ohrid is considered to be the oldest continuously existing lake in Europe with a
likely age of three to five million years. An extraordinarily high degree of endemism, including more than 210
described endemic species, renders the lake a unique aquatic ecosystem of worldwide importance (Albrecht &
Wilke, 2008), and the center of origins of native fish and biodiversity in the Albanian district. Lake Skadar, on the
contrary, is relatively young and originated through flood of a karstic field occupying a crypto-depression of recent
origin (Lasca et al., 1981); it is thus reasonable to hypothesize that its fish fauna originated through multiple
colonization events from surrounding water bodies.
Lectotype designation. Due to the high diversity of the genus Rutilus complex in the Balkan Peninsula (see
Introduction and Ketmaier et al., 2008), a lectotype is here selected among the four syntypes housed in NMW
50723, and 50725. The specimen, 117 mm SL (151 mm TL), bearing the catalogue number NMW 50723-1 is
designed as the lectotype. A picture of that specimen was published in Bianco & Taraborelli (1985: fig. 6, p.149).
According to ICZN (1999), other things being equal, preference for selection of a lectotype should be given to a
syntype that has been illustrated in a publication (art 74. recommendation 74A). The lectotype is in good condition
(Fig. 3B). It has 40 pored scales on LL; 9 above and 4 below LL; 9 branched dorsal rays and 8 branched anal rays;
Zootaxa 3841 (3) © 2014 Magnolia Press
·
387
NEW GENUS AND SPECIES FROM MEDITERREAN EUROPE
pharyngeal teeth formula, 5-5. Head length about 3.8 times in SL; head depth 4.8 times in SL; body depth about 4.0
times in SL, least body depth about 2.3 times in body depth; snout length about 3.4 times in head length. Dorsal and
anal fins slightly concave; caudal fin forked; origin of D at same level as origin of P2; mouth terminal and slightly
oblique; dorsum quite convex.
Remarks on ecology, biology and conservation. It is a still-water adapted species, invasive in lakes Skadar,
Ohrid and lakes of Adriatic drainages of Croatia, and low course of rivers. Spawning takes place between April and
May (Mačrovcić et al., 2006). Its conservation status is of “Low Concern” according to IUCN (2013) red list.
Leucos ylikiensis (Economidis, 1991)
(Fig. 2C)
Rutilus aula rubella var. ylikiensis Stephanidis, 1939 (infrasubspecific name not available: locality, Lake Yliki)
Rutilus ylikiensis Stephanidis, 1991: 269. (Type localities: lakes Yliki and Paralimni and River Kifissos). Available by
reference to Stephanidis (1991). See Kottelat (1997: 82)
Examined material. IZA 8733, 8, Greece, lakes Yliki and Paralimni, P.G. Bianco, 25 April 1987.—IZA 049, 20,
Greece, L. Yliki, P.G. Bianco, 11–16 August 1998.—IZA 0479, Greece, 4, L. Volvi, P.G. Bianco, 21 October 1998.
Diagnosis. A species of Leucos characterized by moderate size, not exceeding 120–130 mm SL; absence of a
mid-lateral band; peritoneal membrane blackened by dense melanophores; usually 42–43 pored scales on LL, as
against 36–38 in Leucos basak. It may be distinguished from Leucos aula for the absence of lateral band and the
color of the peritoneal membrane, black in L. ylikiensis and silvery in L. aula. It differs from all others species of
Leucos for the high number of total GR, usually 18–20 in L. ylikiensis and 8–16 in all others. The most closely
related species, also for the geographical position and according to molecular analyses (Fig. 1B) is L. panosi from
western Greece. Four specimens from Lake Volvi with 17–18 total GR and 41–42 scales on LL are identified as L.
ylikiensis, introduced in this lake similarly to what happened for another endemic species from Lake Yliki,
Scardinius acarnanicus (Bianco & Kottelat, 2005). Bogutskaya & Iliadou (2006) did not report the species as
occurring in Lake Volvi.
Description. Body silvery; lateral band absent; color of eye in living fish, yellowish; fins yellowish or pale
grayish in preserved specimens; snout pointed; pre-orbital distance near equal to the horizontal diameter of the eye;
lips smooth; mouth opening oblique, the corner of maxillae at the same level or a little below; the anteriormost
margin of the eye; head length about 3.8–4.2 times in standard length; body depth about 3.4–3.8 in standard length;
mouth terminal or slightly inferior; profile of dorsum convex, slightly humped in several cases; paired and un-
paired fins yellowish with few scattered melanophores; free margin of dorsal and anal fin concave; caudal fin
forked; peritoneal membrane blackened by several melanophores dense and fused; origin of the D at same level of
the insertion of the P2; LL complete, and extending from the margin of opercular membrane, to the end of caudal
peduncle; 39–45 pored scales, usually 42–43; usually 7.5 row of scales above and 3.5 below the LL; constantly 3
un-branched rays followed by 9 branched rays in the D; constantly 3 un-branched and usually 8 branched rays in
the anal fin; P1s with 1 un-branched and 14–15 branched rays; P2 with 1 un-branched ray and constantly 8
branched rays; 16–21 total GR; in adult reproductive males collected in April, there are granular tubercles on the
free margin of scales of the body and on the head . For additional description and shape, see Fig. 2C and Tables 1,
2.
Distribution. Endemic to the two small and adjacent connected lakes Yliki and Paralimni, and River Ksifissos
in western Greece. Introduced in Lake Volvi (present data).
Remarks on ecology, biology and conservation. A preferentially still water species. There are no data
available on the species life history. According to local fisherman may reach up to 200 mm TL. The maximum
length observed in our materials, collected on April, was for a female of about 150 mm TL, aged 3+, with eggs of
about 1.0–1.2 mm of diameter, all of equal size (a sign of a probable single spawning reproductive habit). Adult
males, of 120 mm TL, aged, 2+, have fan-shaped minute tubercles on the border of scales, of the body and on the
upper part of paired fins. The probable spawning season extends from March to May. Among the IUCN categories,
the species is classified as “Endangered” (IUCN, 2013).
BIANCO & KETMAIER
388
·
Zootaxa 3841 (3) © 2014 Magnolia Press
Zootaxa 3841 (3) © 2014 Magnolia Press
·
389
NEW GENUS AND SPECIES FROM MEDITERREAN EUROPE
TABLE 2. Meristic counts and several general features in five species of Leucos. CV: most common values; N= sample size; L. = Lake; R. = River.
Characters
Leucos basak Leucos aula Leucos ylikiensis Leucos panosi Leucos albus
R. Krupa, L. Besanka,
L. Prespa Micraprespa,
L.Skadar, L.Ohrid
Several basins
Northen Italy
L. Yliki, L. Paralimni,
L.Volvi (introduced) eastern
Greece
L. Ambrakia, L. Trichonis,
L. Joannina (introduced)
Western Greece
R. Moraca & Zeta, L. Skadar
N=130 N= 152 N=32 N=90 N=11
Range
CV
Range
CV
Range
CV
Range
CV
Range
CV
SL (mm) 62–16 89–180 75–116 62–166 58–111
Scales
LL 35–43 36–38 35–42 38–39 39–45 42–43 40–45 42–43 41–44 41–42
Above LL 7½–8½ 6½–8½ 7 ½ 7½–8½ 7½–8½ 7½–8
Below LL 3–4 2½–3½ 3½–4 4 3–4 3–3½ 3
Circumpeduncular 14 14 12–14 12 14 14 14 14 14 14
GR
Total 8–12 9–10 8–11 9 16–21 18–20 12–16 13–14 8–9 8
Branched rays
Dorsal fin 8–9 9 8–10 9 8–9 9 8–10 9 8 8
Anal fin 7-8 8 8-9 9 9
10 9 8–9 9 8 8
Pelvic fin 7–8 8 7–8 8 8 8 7–8 8 8 8
Pharyngeal teeth 5-5/5-4 5-5 5-5/5-4 5-5 6-5/5-5 6-5/5-5 6-5/5-5 6-5/5-5 5-5 5-5
Tubercles in males no Rarely granular
Rarely granular
no no
Lateral stripe no yes no no no
Preferential habitat Lacustrine Lacustrine Lacustrine Lacustrine Lacustrine
Maximum size reached
l
Small Small Small Small Small
BIANCO & KETMAIER
390
·
Zootaxa 3841 (3) © 2014 Magnolia Press
Zootaxa 3841 (3) © 2014 Magnolia Press
·
391
NEW GENUS AND SPECIES FROM MEDITERREAN EUROPE
Leucos panosi (Bogutskaya & Iliadou, 2006)
(Fig. 2D)
Rutilus panosi Bogutskaya & Iliadou, 2006: 283 (type localities rivers Achelos and Louros, western Greece).
Examined materials. IZA 0416, 7, Greece, L. Trichonis; P.G. Bianco, 25 April 1987.—IZA 8766, Greece, canal
emissary of L. Trichonis, P.G. Bianco, 1 May 1987.—IZA 0417, Greece, L. Trichonis, P.G. Bianco, 10 August
1998.—IZA 85513, 28, Greece, L. Ambrakia, P.G. Bianco, 30 April 1984.—IZA 83108B, 27, Greece, L. Joannina
(introduced), P.G. Bianco, 23 February 1976.
Diagnosis. A species of Leucos characterized by moderate size, not exceeding 160–170 mm SL, usually
120–140 mm SL; absence of a mid-lateral band; peritoneal membrane blackened by dense melanophores. Leucos
panosi is quite similar to L. ylikiensis from which it differs mostly for the number of total GR, usually 18–20 in L.
ylikiensis and 13–14 in L. panosi. It may be distinguished from L. basak for the number of LL scales, usually 42–43
as against 36–38 in L. basak. It may be distinguished from L. aula for the absence of lateral band and the color of
the peritoneal membrane, black in L. panosi and silvery in L. aula. It differs from all other species of Leucos for the
high number of total GR, usually 13–14 in L. panosi and 8–10 in all others, excepted L. ylikiensis that has a higher
number of total GR, usually 18–20. The closest species, both on geographical distribution and molecular grounds
(Fig. 1B) is L. ylikiensis from western Greece.
Description. Body uniformly silvery without longitudinal band; color of eye in living fish, yellowish; fins
yellowish or pale grayish in preserved specimens; a triangular pale, crescent spots at insertion of each scale of the
body; snout pointed; pre-orbital distance near equal to the horizontal diameter of the eye; lips smooth; mouth
terminal, or slightly inferior, its opening oblique; the corner of maxillae at the same level or slightly overpass the
vertical crossing the anterior border of the orbit; profile of dorsum convex, slightly humped; paired and unpaired
fins yellowish with few scattered melanophores; free margin of dorsal and anal fin concave; caudal fin moderately
forked; peritoneal membrane blackened by several dense and fused melanophores; head length about 4.2–4.6 times
in SL; body deep about 2.8–3.2 times in the SL; origin of the D placed slightly above the vertical crossing the
insertion of the P2; LL complete, and extending from the margin of opercular membrane, to the end of caudal
peduncle; 39–45 pored scales, usually 42–43; 7.5 above and 3.5 below the LL; 14 circum-peduncular scales;
constantly 3 un-branched rays followed by 9 branched rays in the D; constantly 3 un-branched and usually 8
branched rays in the anal fin; P1 with 1 un-branched and 14–16 branched rays; P2 with 1 un-branched ray and
constantly 8 branched rays; 12–16 total GR; fine granular tubercles on head and checks has been observed in adult
reproductive males. For additional description and shape, see Fig. 2D and Tables 1, 2.
Distribution. The species was reported for the Acheloos, and Louros river-systems in western Greece
(Bogutskaya & Iliadou, 2006). The species was found also in lakes Trichonis and Ambrakia. Leucos panosi was
introduced in Lake Joannina, (Leonardos et al., 2008), where it coexists with the introduced Rutilus rutilus.
Remarks on ecology, biology and conservation. A still-water adapted species. In fish communities, Leucos
panosi tends to become dominant and invasive in lakes while it is quite rare in the flowing part of rivers Louros and
Acheloos, (P.G. Bianco, pers. obs.). Spawners have been collected in February suggesting that the reproductive
season may start at the end of January and extend possibly until March–April. The species is classified as of “Low
Concern” among the IUCN (2013) categories.
Leucos albus Marić, 2010
(Fig. 2E)
Rutilus albus Marić, 2010: 153 (type locality, Lake Skadar, Montenegro).
Examined materials. IZA 0421, 2, Montenegro, R. Moraca at the confluence with R. Zeta, D. Maríc and P.G.
Bianco, 11 November 1999 (specimens also tested molecularly by Ketmaier et al. (2008).- IZA 0469, 7, 58,
Republic of Montenegro, R. Zeta, 13 November 1999, P.G. Bianco and D. Marić.
Diagnosis. This species was recently described by Marić (2010). Here we provide a short diagnosis according
to our materials from the Moraca-Zeta river system, which flows into the Lake Skadar; the species is not reported
for that rivers (Marić, 2010). Leucos albus can be distinguished from L. basak, its closest relative genetically (see
BIANCO & KETMAIER
392
·
Zootaxa 3841 (3) © 2014 Magnolia Press
Fig. 1B), for the number of LL pored scales (41–44 in Leucos albus 37–39 in L. basak) and for the modal number
of branched rays in the D (9 in L. basak, 8 in L. albus). Flanks are silvery, without lateral bands as observed in L.
aula.
Description. A small-medium sized species, not exceeding 190 mm SL, usually 80–120 mm SL. Body
uniformly silvery without longitudinal band; color of eye in living fish, yellowish; fins yellowish or pale grayish in
preserved specimens; snout pointed, pre-orbital distance near equal to the horizontal diameter of the eye; lips
smooth; mouth opening oblique, the corner of maxillae slightly protruded and at level with the vertical crossing of
the anterior border of the orbit; mouth slightly inferior; profile of dorsum convex; paired and unpaired fins
yellowish; free margin of dorsal and anal fins concave; caudal fin forked; peritoneal membrane blackened by
several melanophores addensed and fused; head length more than 3.8–4.0 times in the SL; body deep about 3.6–3.8
times in the SL; origin of the D at same level of the insertion of the P2; LL complete, and extending from the
margin of opercular membrane, to the end of caudal peduncle; 41–44 pored scales, usually 41–42; 7.5 above and
3.5 below the LL; constantly 3 un-branched rays followed by 8 branched rays in the D; constantly 3 un-branched
and usually 8 branched rays in the anal fin; P1s with 1 un-branched and 14–15 branched rays; P2 with 1 un-
branched ray and constantly 8 branched rays; usually 8–9 total GR; in adult males prominent or vestigial tubercles
absent. The specimens from River Moraca differ from those from Lake Skadar for a more pointed snout and
slender body. For additional description and shape, see Fig. 2E and Tables 1, 2.
Distribution. L. albus is originally described from Lake Skadar (Montenegro): it is quite rare in the lake, but
seasonally frequent in submerged springs (Marić, 2010). We also found the species in Zeta and Moraca river
system, both flowing into Lake Skadar. The species was found together with Salmo sp, Telestes montenegrinus and
Phoxinus phoxinus, all cold-water adapted species. The species, which seems riverine adapted, is steadily declining
both in rivers Moraca and Zeta mainly due to illegal fishing of trout (Drago Marić, pers. comm.). The scarce
material we obtained (nine specimens) is the result of extensive sampling efforts by electro fishing between 9 and
13 November 1999.
Remarks on ecology, biology and conservation. According to Maríc (2010), the preferential habitats are the
sub-lacustrine springs of Lake Skadar, where large numbers of fish gather during the winter period. We found also
a riverine sample. Leuco albus coexists with L. basak in Lake Skadar. The proportion between Leucos albus and L.
basak is 20:1 in favor of the second. Spawning season starts in January but in February females have spent gonads.
Leucos albus spawns earlier in the season than L. basak. Still not evaluated by IUCN International commission, but
according to the IUCN (2012) category, it satisfy to the point B of the “Endangered Category”, as the area of
occupancy is less than 500 km
2
and the species is known only from one basin (Lake Skadar and the river flowing
into it), which continues to decline as a result of habitat alteration (Marić, 2010).
Sarmarutilus n. gen.
Type species: Leuciscus rubilio Bonaparte, 1837: Fauna It: fasc. XIX, Tav III (Type locality; voulcanic lakes of Nemi and
Bracciano, central Italy).
Diagnosis. The species, previously classified as Rutilus rubilio, cannot be placed in the genus Rutilus, because of
the pharyngeal teeth formulae, 5-5 in R. rubilio and 6-6 or 6-5 in Rutilus, of the small size and the riverine
preference as opposed to still waters and large size in Rutilus. Similarly, it cannot be placed in the genus Leucos for
the presence of prominent tubercles at the center of each scale of the body and on the head in reproductive males
(Fig. 4), a character unknown in the genus Leucos, and for the preferentially riverine habits as opposed to still
waters in the five species of Leucos. In addition, it shows a marked lateral stripe, quite different from that of L.
aula, masked by heavy pigmentation on flanks formed by well marked crescent triangular spot on scales (Fig. 2F),
of a shape not found in any of all other species of Leucos and Rutilus in the examined area. Finally, it is well
separated from the Leucos species on karyology grounds (Bianco et al., 2004) and from Rutilus and Leucos at the
mtDNA level (Ketmaier at al., 2008).
Etymology. The generic name derives from the Sarmatic Sea, or Lago Mare, an ancient central European inner
freshwater sea where this monotypic genus probably has its evolutionary roots.
Origins. Sequence variation of the entire mitochondrial cytochrome b gene shows a strict relationship among
Zootaxa 3841 (3) © 2014 Magnolia Press
·
393
NEW GENUS AND SPECIES FROM MEDITERREAN EUROPE
five species, which diverged from a Leuciscinae ancestor about 4–5 million years ago, during the Lago Mare phase
of the Mediterranean Sea (Bianco, 1990). Similar biogeographic and temporal patterns have been observed in the
genus Tel est e s , which occupies the same geographic area (Ketmaier et al., 2004). According to karyology (Bianco
et al., 2004) Sarmarutilus rubilio displays few, if any, elements with centromeric heterochromatin and many with
solid telomeric bands; this evidence differentiates S. rubilio from two representative species of the genus Leucos
(L. aula and L. panosi). The three examined species should be considered as of distinct origins and, according to
molecular data, probably separated since the Lago Mare Phase of the Mediterranean Sea in the Miocene Messinian
(5 MYA).
FIGURE 4. Anterior part of body of male Sarmarutilus rubilio showing prominent tubercles at centers of scales (from Bianco
& Taraborelli, 1985).
Sarmarutilus rubilio (Bonaparte, 1837)
(Figs. 2F, 3C)
Leuciscus rubilio Bonaparte, 1837: Fauna It: fasc. XIX, Tav III (Type locality; voulcanic lakes of Nemi and Bracciano, central
Italy).
Examined material. samples from the following rivers were analyzed for the study. All the examined material is
from Italy and, since the species was introduced in several basins, only samples from the native distribution has
been included in the description and following comparisons. IZA 8344, 6, R. Elsa (R. Albegna basin), Tuscany, G.
Delmastro, 20 May 1981.—IZA 8355, 13, R. Cecina, Tuscany, G. Delmastro, 18 May 1981.—IZA 8358, 11, R.
Serchio, Tuscany, 18 May 1981.—IZA 8396, 11, Fosso della Lena (R. Ombrone basin), Tuscany, P.G. Bianco, 10
April 1982.—IZA 8367, 8, R. Trasubbie (R. Arno basin), Tuscany, G. Delmastro, 19 May 1981.—IZA8374, 5, R.
Fiora, Tuscany, P.G. Bianco, April, 1975.—IZA 8363, 22, R. Mignone, Latium, P.G. Bianco, June 1974.—IZA
8390, 12, R. Tronto, Marche, C. Albertini, 22 November 1982.—IZA 8381, 20 (out of 43), R. Tavo, Abruzzo, P.G.
Bianco, 12 April 1979.—IZA 8373, 14, L. Provvidenza (R. Vomano basin), P.G. Bianco, 10–15 April 1983.—IZA
BIANCO & KETMAIER
394
·
Zootaxa 3841 (3) © 2014 Magnolia Press
8376, 16, R. Pescara, Abruzzo, F. Recchia, May 1982.—IZA 8364, 5, R. Trigno, Molise, P.G. Bianco, 10 July
1977.—IZA 8381, 12, R. Volturno, Campania, P.G. Bianco, 8 August 1981.—IZA 0252, 20 (out of 54) , R. Calore,
Campania, P.G. Bianco & V. Frezza, 31 October 2001.—IZA 00236, 20 (out of 84) R. Sele, Campania, P.G. Bianco,
24 April 2001.—IZA 00408, 6, R. Nestore (R. Tiber basin), P.G. Bianco & M. Lorenzoni, 11 April, 1996.—IZA
02122, 20, R. Merse (R. Ombrone basin), Tuscany, P.G. Bianco , 2 July, 1997.—IZA 02183, 20, (out of 75), R.
Ofanto, Campania (probably introduced), P.G. Bianco & V. Frezza, 12 February 2002.—IZA Uncatalogued, 7, L.
Trasimeno, Umbria, P. Calderoni, 1984.—IZA 87146, 20 (out of 87), R. Liri, Lazio, P.G. Bianco, 7 July 1987.
Diagnosis. Corresponds to that of the genus.
Description. A small-medium sized species, not exceeding 160 mm SL, usually 80–120 mm SL. Scales of the
body marked by evident crescent triangular spot on the origins, sometimes this heavy pigmentation masks a more
or less evident longitudinal stripe; color of eye in living samples, yellowish; fins yellowish or pale grayish in
preserved specimens; in reproductive adults are reddish; snout moderately blunt; pre-orbital distance nearly equal
to the horizontal diameter of the eye; lips smooth; mouth opening oblique, the corner of maxillae placed anteriorly
to the vertical crossing of the anterior border of the orbit; mouth slightly inferior; profile of dorsum convex ; free
margin of D and A concave; P1 and P2 longer in males, where they may reach the origin of P2 fin; caudal fin
forked; peritoneal membrane blackened by several melanophores dense and fused; head length about 4.0–4.3 times
the SL or more; body depth about 2.8–3.1 times the SL; origin of the D at same level or slightly below the insertion
of the P2; LL complete and extending from the margin of opercular membrane to the end of caudal peduncle;
37–42 pored scales on LL; 7.5 row of scales above and 3.5 below the LL; constantly 3 un-branched rays followed
by modally 8 branched rays in the D; constantly 3 un-branched rays followed by modally 9 branched rays in the
anal fin; P1 with 1 un-branched and 14–15 branched rays; P2 with 1 un-branched ray and modally 8 branched rays;
usually 10–11 total GR; in adult males prominent or vestigial tubercles present. For additional description and
shape, see Fig. 2F, Tables 4,5, and Bianco & Taraborelli (1985).
Distribution: The species was endemic to the Tuscany-Latium ichthyogepgraphic district, including an area
limited on the Tyrrhenian side between River Magra to the north and River Sele to the south. The species range on
the Adriatic side extends from River Tronto (north) to River Trigno (south) (Bianco & Taraborelli, 1985). The
species was introduced in most of the southern Italian basins, from where we have the following documented
materials: IZA 83127, 75, L. Campotosto, Abruzzo, L. G. Albertino, June 1982.—IZA 8347, 26, R. Bussento,
Campania, P.G. Bianco, 24 June 1977.—IZA 8384, 54, R. Mingardo, Campania, P.G. Bianco, 25 June 1978.- IZA
8393, 17, R. Cavone, Basilicata, P.G. Bianco, 31 July 1978.—IZA 83104, 7, R. Bradano, Basilicata, P.G. Bianco, 11
August 1981.- IZA 83107, 30, R. Crati, Calabria P.G. Bianco, 1 August 1978.- IZA 83141, 2, R. Neto, Calabria,
July 1982, A. Marconato.- IZA 87156, 122, R. Basento, Basilicata, P.G. Bianco, 14 July 1987.- IZA 00244, 70, R.
Bussento, Campania, P.G. Bianco, 25 march 2001.- R. Agri, IZA 87130, 761, Basilicata, P.G. Bianco, 13 August
1987.—IZA 87151, 49, R. Sinni, Basilicata, P.G. Bianco 1987.—IZA 87153, 131, R. Lao, Calabria, P.G. Bianco, 10
August 1987.—IZA 00225, 109, R. Mingardo, Campania, 12 June 1989.—IZA 87164, 17, R. Cavone, Basilicata,
P.G. Bianco, 13 August 1987.—IZA 87172, 23, R. Noce, Calabria, P.G. Bianco, 9 August 1987. In Sicily it was
introduced in 1985 (Tigano & Ferrito, 1986).
Lectotype designation. The type series of this species includes 11 syntypes, 66–121 mm SL, ANSP 6509-
6519. The Bonaparte description and illustration are based on a single specimen, of “6 inches and 9 lines” (about
17 cm TL), from Lake Nemi (the type locality) which cannot be recognized from the series. Hence, a lectotype is
here selected: the specimen ANSP 6509, 93 mm SL (Fig. 3C), owing to its very good condition, is designated as
lectotype. It has 38 scales on LL; 8 branched rays in D and 9 in A; 7.5 rows of scales above and 3 below LL; total
GR, 10; 14 circum-peduncular scales; 5-5 pharyngeal teeth. The 10 paralectotypes, ANSP 6510-6519, 66–121 mm
SL: LL, 36–40; 7.5–8.5 rows of scales above and 2.5–3.5 below LL. D constantly with 8 branched rays, A with 9
(10 in one case) and 9–11 total GR. Pharyngeal teeth formula is 5-5.
Remarks on ecology biology and conservation. Preferential habitats: running waters. Not well adapted in
still waters where it is easily excluded by the introduced Leucos aula, Rutilus rutilus and possibly other introduced,
still water-adapted species such as rudds and bleaks. In southern Italian rivers, however, where it was introduced,
detrimentally affected or eliminated the populations of the less riverine well-adapted species as the endemic
Alburnus albidus (Bianco & Ketmaier, 2001).
The reproductive season extends from March to June with sporadic case in February and July. Sexual maturity
is reached at age 1+ or 2+ in males and 2+ in females. Maximum age observed, 6 years for females and 5 for males.
Zootaxa 3841 (3) © 2014 Magnolia Press
·
395
NEW GENUS AND SPECIES FROM MEDITERREAN EUROPE
Maximum length reached, 160 mm SL a female of age 5+, and weight of 45 g. (Bianco & Taraborelli, 1985; Bianco
& Santoro, 2004). According to Bianco et al. (2013), and IUCN (2013), Sarmarutilus rubilio, throughout its native
range, is placed in the category of “Nearly Threatened”, while wherever it is introduced it has become invasive.
Rutilus rutilus (Linnaeus, 1758)
(introduced in the study area)
Cyprinus rutilus Linnaeus, 1758: 234 (type locality: Europa)
Rutilus rutilus vegariticus Stephanidis, 1950: 203 (type locality, Lake Vegoritis, northern Greece).
Examined material. IZA 83108A, 107, Greece, Epirus, L. Joannina, P.G. Bianco, 23 February 1976.—IZA 88124,
6, England, R. Thames, A. Wheeler, 18 June 1986.—IZA 78115, 11, Romania, R. Danube, P. Banarescu, 23
September 1975.—IZA 0475, 15, Greece, L. Vegoritis (type locality of R. vegariticus), P.G. Bianco, 21 August
1998.—IZA 0452, 5, FYROM, L. Dojran, R. Gruptché, 14 January 1971.—0477, 7, Italy L. Como, O. Mangione,
April 2004.—IZA 0478, 5, Slovenia, L. Pernica (R. Danube basin) at Mari-Bor, M. Povz, 29 March 1999.
Diagnosis. Rutilus rutilus compared to R. pigus has less scales on LL (39–44 vs 46–51), and higher number of
total GR (12–16 vs. 9–12). Compared to Rutilus stoumboudae n.sp., R. rutilus has less total GR (12–15 vs. 15–18)
and more branched rays both in the dorsal and anal fins (9–11 vs. 8–9). Proportional measurements and meristic
counts are reported in Tables 4, 5.
Description. A species of Rutilus of medium-large size, characterized by the absence of a mid lateral band;
body slightly compressed; and moderately elongate, dorsal profile convex; sometimes humped in larger specimens;
posterior mouth corner placed above the vertical crossing the anterior margin of orbital cavity; mouth terminal,
opening small, slightly downturned; lips smooth; horizontal diameter of the eye more than or equal to the pre-
orbital length; head small, with a length about 4.2–4.6 times the SL; body depth, about 3.2–3.6 times the SL; origin
of the D slightly below the insertion of the P2; free margin of dorsal and anal fins slightly concave; caudal fin
deeply forked; LL complete, with 39–44 pored scales, placed below the middle of the body, slightly concave and
extending from the upper margin of the opercular membrane to the end of caudal peduncle; modally 3 un-branched
and modally 10 branched rays both in the dorsal and anal fins; P1 with 1 un-branched and 15–17 branched rays; P2
with 1 un-branched ray and constantly 8 branched rays; usually 12–14 total GR; color in life: eye from yellowish to
a brilliant red; fins yellowish with some red; dorsal and caudal fins dark brown; peritoneal membrane silvery
sometimes with few scattered melanophores. Additional description in Tables 4, 5.
Remark on synonym. The status of Rutilus vegariticus was still confused and inquirenda. It was considered as
a valid species (Economidis & Banarescu, 1991), or a subspecies of Rutilus (Economidis, 1995). Kottelat (1997)
still considered questionable its validity as distinct taxon. Following its original description, no recent diagnosis
and description of this nominal taxon were performed. According to molecular data (Ketmaier et al., 2008) and to
the morphological analyses presented here (Tables 4, 5), Rutilus vegariticus cannot be consistently separated from
R.rutilus as all morphometric and meristic characters overlap (Tables 4, 5). Rutilus vegariticus (Lake Vegoritis ,
eastern Greece) is genetically very close to a number of central European and introduced northern Italian
populations of R. rutilus (Ketmaier et al., 2008). Larmuseau et al. (2009) claimed that this population should be
regarded as a distinct Rutilus rutilus lineage, possibly a junior synonym for the Ponto-Caspian R. heckelii. This
seems not realistic biogeographically (Bianco, 1990), since the Ponto-Caspian region is well differentiated from
the circum-Mediterranean one; we believe that the most parsimonious explanation for the occurrence of Rutilus
rutilus in the area is introduction by humans and, therefore, we consider this taxon as junior synonym of R. rutilus.
Distribution. Native to central Europe, the species has been introduced in several circum-Mediterranean
ichthyogeographical districts. In northern Italy, the species has been introduced in the Po basin and in many
subalpine lakes where it compete with the native Rutilus pigus and Leucos aula; in central Italy introductions are
known for the Tiber and Arno rivers and for lakes Montedoglio and Corbara, where it caused the disappearance of
the endemic Sarmarutilus rubilio.
BIANCO & KETMAIER
396
·
Zootaxa 3841 (3) © 2014 Magnolia Press
Zootaxa 3841 (3) © 2014 Magnolia Press
·
397
NEW GENUS AND SPECIES FROM MEDITERREAN EUROPE
BIANCO & KETMAIER
398
·
Zootaxa 3841 (3) © 2014 Magnolia Press
Rutilus pigus (La Cepède, 1804)
(Fig. 2G)
Cyprinus pigus La Cepède, 1803: 503, 607 (type localities: lakes Major and Como, northern Italy).
Examined material. IZA 0471, 12, Italy, L. Como, O. Mangione, April 2004.— IZA 0472, 3, Italy, R. Livenza, M.
Povz, 22 May 1996.—MSNM (uncataloged), 33, Italy, L. Maggiore, A. Mojetta, June 1988.—IZA 843,6, Italy, R.
Po, G. Delmastro,13 March 1983.- IZA 842, 6, Italy, L. Major, April 1983, G. Giussani.
Diagnosis. Rutilus pigus, was only recently separated from R.virgo (Kottelat & Frehyof, 2007; Ketmaier et al.,
2008). R. pigus, is endemic of Padany-Venetian district, and differs from R. virgo from central Europe, in
squamation as it has 46–51 scales on LL, and 8.5 above and 3.5 below the LL, as against 40–46 scales on LL and
7.5 above and 3.5 below the LL in R. virgo. The two species show also a different position of nuptial tubercles on
head (Kottelat & Freyhof, 2007) and also phylogenetically are very distant (Ketamaier et al., 2008). Compared
with R. rutilus, has more scales on LL scales, 46–51 in R pigus, and 39–44 in R. rutilus; and less number of total
GR, 9–12 in R. pigus and 12–16 in R. rutilus. Compared to R. stoumboudae n.sp., R. pigus has less total GR (12–15
vs. 15–18) and more branched rays both in the dorsal and anal fins (9–11 vs. 8–9).
Description. A species of Rutilus characterized by large size and the absence of mid lateral band. In
reproductive males, well developed and large prominent nuptial tubercles on head, checks, and at center of the
scales of the body. Absence, in reproductive males, of fan-shaped granular tubercles, on the free border of each
scales of the body. Body laterally compressed, moderately elongate, dorsal profile convex; sometimes a small
hump in largest specimens; snout pointed; eye diameter equal to the pre-orbital length; posterior mouth corner
placed beyond the vertical crossing the anterior margin of orbital cavity; mouth opening oblique and terminal to
slightly sub-terminal; lips smooth; eye small, its horizontal diameter less than or equal to the pre-orbital length;
head length about 4.3–4.6 times in the SL; body quite deep, its depth about 3.2–3.4 times in the SL; origin of the D
placed at same level of the insertion of the P2; free margin of dorsal and anal fins, from straight to slightly concave;
caudal fin, when enlarged, moderately forked; LL complete, with 46–51 pored scales extending from the upper
margin of opercular membrane, to the end of caudal peduncle; modally 3 un-branched and 10 branched rays in the
dorsal and 11 in the anal fins; P1 with 1 un-branched and 15–17 branched rays; P2 with 1 un-branched ray and
constantly 8 branched rays; about 8–12 total GR; color of the eye white; the color of all fins is from yellowish to
dark grays; peritoneal membrane blackened by dense melanophores. For additional description and shape, see Fig.
2G, and Tables 4, 5.
Distribution. Padano-Venetian district, from River Po to River Isonzo (upper Adriatic Sea drainages). Its
survival is strongly affected by the introduction of two alien species: the European roach, Rutilus rutilus and the
European nase, Chondrostoma nasus (Linnaeus, 1758).
Remarks on ecology, biology and conservation. A mostly lacustrine adapted species, which may reach 500
mm TL and 2 kg of weight, and at least 9 years of age. Sexual maturity is late and is reached at age 3+ or 4+ in
males and 5+ in females at about 280–330 mm SL. Reproduction takes place between April and May, longevity is
up to 9 years (Bianco & Taraborelli, 1985; Puzzi et al., 2007). Although the species was placed in the “Low
Concern” category by (IUCN, 2013), according to the recent assessment made by the Italian National IUCN red list
commission, R. pigus was placed in the “Critical Endangered” category (Bianco et al., 2013). This species is
progressively disappearing throughout its native range, mainly as a result of competition with the invasive
introduced R. rutilus and Chondrostoma nasus (Puzzi et al., 2007; Volta & Jepsen, 2008).
Rutilus stoumboudae Bianco & Ketmaier, new species
(Fig. 2H)
Type materials. Holotype (Fig. 2H) IZA 02107A, 128 mm SL, Greece, Lake Volvi, P.G. Bianco, 21 October 1998
(specimen also used for molecular analysis by Ketmaier et al., 2008).
Paratypes: IZA 02107B, 11, 64–156 mm SL, Greece, Lake Volvi; P.G. Bianco, 10 October 1998.
Diagnosis. Rutilus stoumboudae differs from R. rutilus, its closest relative genetically, both in the number of
total GR (16–17 vs. 13–14) and number of branched rays of the D and A (modally 9 and 9 in R. stoumboudae and
Zootaxa 3841 (3) © 2014 Magnolia Press
·
399
NEW GENUS AND SPECIES FROM MEDITERREAN EUROPE
10 and 10 in R. rutilus). It differs from R. pigus in apparently lacking nuptial tubercles, in the numbers of LL scales
(39–43 vs. 46–51) and of branched rays in the anal fin (8–9 vs. 9–11).
Description. Apparently a small to medium sized species. Maximum SL observed, 156 mm and 197 mm of TL
in our materials. Body moderately compressed, its depth about 3.1–3.3 times in SL. Head length about 3.8–4.0
times in SL. Mouth, small, terminal and slight up-turned. Pre-orbital distance less than eye diameter. Eye diameter
about 2.7–3.3 times in head length. Origin of D slight behind the origin of P2. Free margins of D and A concave;
dorsal profile convex, slightly humped in the holotype; pre-dorsal distance slightly longer than pre-ventral
distance. Pectoral-pelvic distance nearly equal to pelvic-anal distance. Caudal peduncle depth about 1.1–1.3 times
in caudal peduncle length. Free margin of pectoral and pelvic fins rounded, caudal fin, when extended, moderately
forked. D with 3 non-branched and 9 (in one case 8) branched rays. A with 3 non-branched and 9 branched rays.
Fourteen circum-peduncular scales; 38–41 pored scales on the LL. 7.5 or 8.5 rows of scales above and 3.5 or 4.0
below the LL; 8–10 total GR; pharyngeal teeth on both sides, 6-5; Peritoneal membrane darkened by dense
melanophores; apparently absence of tubercles. Ground color silvery; back plain dark brown. Top of head plain
dark brown; sides light; ventral surface of head, belly and caudal peduncle yellowish. All fins yellow-brownish.
General shape of holotype in Fig. 2H. Morphometric and meristic data of holotype and 11 paratypes in Tables 4, 5.
Remarks. According to Larmuseau et al. (2009), the species from Lake Volvi examined by Ketmaier et al.
(2008), should be identified as Rutilus heckeli. But these authors specifically stated in their study that the
identification of the alleged Rutilus heckeli material they used was uncertain: the authors did not deposite the
corresponding sequences in GenBank and we are thus unable to include their data in our phylogenetic analyses.
According to Larmuseau et al. (2009), Rutilus heckeli seems the sister species of R. rutilus. Extensive diagnosis of
this species is found in Berg (1949), and Banarescu (1964): it may reach the size of about 500 mm TL (size
unknown in Lake Volvi, since Rutilus stoumboudae belongs to a group of species of smaller size). Adult males of
Rutilus heckeli show well developed nuptial tubercles all along the dorsal area of the body (apparently not found in
R. stoumboudae); pharyngeal teeth formula is modally 6-6 in R. heckeli, while in our new specie is modally 6-5;
the number branched rays of dorsal and anal fins are modally 10 in R. heckeli, and 9 in R. stoumboudae. Based on
these differences we cannot identify samples of the new species as Rutilus heckeli. Another geographically close
species, described for the River Maritza in Bulgaria (Evros in Greece), Rutilus maritza, Drenski, 1926, is
considered since a long time as a junior synonym of R. rutilus (Kottelat, 1997). In addition, the Bulgarian
freshwater fish fauna is of Danubian origin, with the exception of a single endemic species, Alburnus mandrensis
(Drensky, 1943).
Color pattern. In preserved specimens, dorsum brownish, sides yellowish without a longitudinal band. Scales
of flanks show a reticulate pigmentation. Peritoneal membrane with more or less concentrated or scattered
melanophores. Pectoral, dorsal, and caudal fins black; pelvic and anal fins yellowish.
Distribution: There are several doubts about the native origin of this species in Lake Volvi. Kottelat & Frehyof
(2007) reported a Rutilus sp. from River Sperchios (south-eastern Greece). According to the count methodology
proposed in Bianco & Kottelat (2005) the species shows 9 branched rays in the anal and in the dorsal fins. Among
the Rutilus species revised in this contribution, only the new species possess 9 branched rays in the dorsal and anal
fins. The general shape of the specimens from the River Sperchios described by Kottelat (2007: p 249) is very
similar to that of the holotype of R. stoumboudae.
Translocations of fishes by fishermen from southern Greece basins to the shore market of Lake Volvi for
selling seems quite frequent. Leucos ylikiensis and Scardinius acarnanicus, endemic to Lake Yliki, have been
transplanted in Lake Volvi following this practice.
Etymology. The species is named after the Greek ichthyologist, colleague and friend, Maria Stoumboudi, in
honor of her research activity on the ecology and conservation of freshwater fishes of Greece.
Remarks on ecology biology and conservation: a still water adapted species. No data available on its
biology. According to the IUCN (2012) category, it satisfies point B and potentially should be placed in the
“Endangered category” as the area of occupancy is less than 500 km
2
and the species keeps declining as a result of
habitat reduction.
BIANCO & KETMAIER
400
·
Zootaxa 3841 (3) © 2014 Magnolia Press
Conclusion
In this study, we updated the taxonomy of the Rutilus complex in southern Europe and presented an arrangement
different from that proposed by Kottelat & Freyhof (2007). In our opinion, this new taxonomic scheme better
mirrors the effects that the paleogeography and paleoecology of the area had on the diversification of this group of
cyprinids (Fig. 1B) (Bianco, 1990; Ketmaier et al., 2008). Based on phylogenetics, karyologic, morphological and
ecological considerations, we resurrected the genus Leucos Heckel, 1854, which includes five species. Among
them, the Balkanic roach (L. basak) shows the broadest geographical range, possibly because it took advantage of
the multiple river connections in the area during the Würmian Ice Age. L. aula is endemic to northern Italy; L.
ylikiensis and L. panosi are both endemic to Greece. Leucos albus is endemic to Albany, where it coexists with L.
basak. In central Italy the monotypic genus Sarmarutilus is described. The only representative of the genus
probably originated in the Sarmatic area in Middle Miocene, reached the Mediterranean area during the Lago Mare
phase, and survived only in the Tuscany-Latium district of Italy. In the genus Rutilus, R. rutilus is of alien origin in
the study area. Rutilus pigus is endemic to the Padano-Venetian district. Rutilus stoumboudae n.sp. is endemic to
Lake Volvi (Greece) and is well differentiated both morphologically and molecularly from the other two species of
Rutilus. We also emphasize that the conservation status of several species are affected by alien introductions. Today
Rutilus stoumboudae has probably gone extinct in Lake Volvi (Greece) due to the dramatic reduction in water level
that the basin has witnessed in recent years and probably also as a result of the introduction of L. ylikiensis. Rutilus
pigus in Italy is currently threatened by the introduction of R. rutilus, which has become invasive throughout the
whole peninsula. It is worth noting that the joint introduction of R. rutilus and Chondrostoma nasus is affecting
also other still water species such as Leucos aula and Chondrostoma soetta, that have nearly vanished from
northern Italian lakes. Leucos basak and L. panosi seem to be in a relatively good conservation status across their
respective ranges. R. ylikiensis is affected by habitat reduction as Lake Yliki and Paralimny nearly dry up during
the summer season (P.G. Bianco, pers. obs.). According to several private Italian pond owners, L. basak has been
stocked with forage predators such as zanders and black basses (Bianco, 2014b). Sarmarutilus rubilio, endemic to
the Tuscano-Latium district, central Italy, west of the Apennines, is locally threatened as a result of the introduction
of species such as the European Rutilus rutilus, and Leucos aula, Alburnus arborella, Chondrostoma genei and
Scardinius hesperidicus, all species introduced from the Padano-Venetian district and that are apparently more
competitive and able to displace it (Bianco & Ketmaier, 2001). Unfortunately, what we just described is not an
isolated case but is rather a general trend in the Italian fish fauna, which has been severely altered by introductions
and translocations of species in the last hundred years, mostly for stocking purposes (Bianco & Ketmaier, 2001;
2005). Something similar is happening in Greece, where introduction of non-native L. ylikiensis in Lake Volvi, L.
panosi and Rutilus rutilus in Lake Joannina have affected native species (Leonardos et al., 2008).
Acknowledgements
We wish to thank the following colleagues for their support during the collecting trips, for lending Rutilus samples
or giving access to the collection of their institutions. Draco Marić (Podgorica University, Montenegro), Mrako
Mrakovic (Zagreb, Croatia), Meta Povz, (Ljubljana, Slovenia), Giovanni Delmastro (Carmagnola, Italy), William
Smith-Vanitz (Philadelphia, USA), Olindo Mangione (Milano, Italy).
References
Albrecht, C. & Wilke, T. (2008) Ancient Lake Ohrid: biodiversity and evolution. Hydrobiologia, 615, 103–140.
http://dx.doi.org/10.1007/s10750-008-9558-y
Banarescu, P. (1964) Fauna Republicii populare Romine. Pisces, Osteichthyies. Vol. 13. Bucuresti, 959 pp.
Berg, L. (1949) Freshwater fish of the USSR and adjacent countries. Israel Program for Scientific Translation (1964). Vol. 2.
Jerusalem, 496 pp.
Bianco, P.G. (1990) Potential role of the palaeohistory of the Mediterranean and Parathetys basin on the early dispersal of
Europe-Mediterranean freshwater fishes. Ichthyological Exploration of Freshwaters, 1, 167–184.
Bianco, P.G. (1998) Diversity of Barbinae fishes in southern Europe with description of a new genus and a new species. Italian
Journal of Zoology, 65 (Supplement), 125–136.
Zootaxa 3841 (3) © 2014 Magnolia Press
·
401
NEW GENUS AND SPECIES FROM MEDITERREAN EUROPE
http://dx.doi.org/10.1080/11250009809386804
Bianco, P.G. (2014a) Lectotype designation for Squalius squalus (Bonaparte, 1837) (Pisces:Cyprinidae). Proceedings of the
Academy of Natural Sciences of Philadelphia, 163, 91–93.
http://dx.doi.org/10.1635/053.163.0103
Bianco, P.G. (2014b) An update of the status of the native and exotic freshwater fish of Italy. Journal of Applied Ichthyology,
30, 62–77.
Bianco, P.G., Aprea, G., Balletto, E., Capriglione, T., Fulgione, M. & Odierna, G. (2004) The karyology of the cyprinid genera
Scardinius and Rutilus in southern Europe. Ichthyological Research, 51, 274–278.
http://dx.doi.org/10.1007/s10228-004-0221-y
Bianco, P.G., Caputo, V., Ferrito, V., Lorenzoni, M., Nonnis Marzano, F., Stefani, F., Sabatini, A. & Tancioni, L. (2013) Pesci
d’acqua dolce. In: Rondinini, C., Battistoni, A., Peronace, V. & Teofili, C. (Compilers.), Lista Rossa IUCN dei Vertebrati
Italiani. Comitato Italiano IUCN e Ministero dell’Ambiente e della Tutela del Territorio e del Mare, Roma, pp. 54.
Bianco, P.G. & Ketmaier, V. (2001) Anthropogenic changes in the freshwater fish fauna of Italy, with reference to the central
region and Barbus graellsii, a newly established alien species of Iberian origin. Journal of Fish Biology, 59, 190–208.
http://dx.doi.org/10.1111/j.1095-8649.2001.tb01386.x
Bianco, P.G. & Kottelat, M. (2005) Scardinius knezevici, a new species of rudd from Lake Skadar, Montenegro (Teleostei:
Cyprinidae) Ichthyological Exploration of Freshwaters, 16, 231–238.
Bianco, P.G. & Santoro, E. (2004) Biologia della rovella, del cavedano e del barbo nei bacini del Parco Nazionale del Cilento e
Vallo di Diano (Cyprinidae). Biologia Ambientale, 18, 85–91.
Bianco, P.G. & Santoro, E. (2011) I pesci e i decapodi d’acqua dolce della Riserva Naturale Monterano: alterazioni prodotte,
status degli autoctoni e indicazioni gestionali. In: Bianco, P.G. & de Filippo, G. (Eds.), Pesci parchi vol 3. Contributo alla
conoscenza della fauna ittica d’acqua dolce in aree protette d’Italia. Researches on Wildlife Conservation. Publisher:
Lulu.com USA IGF Publ., ISBN: 978-1-4709-2573-4, pp. 1–24.
Bianco, P.G. & Taraborelli, T. (1985) Contributo alla conoscenza del genere Rutilus R. in Italia e nei balcani occidentali (Pisces,
Cyprinidae). Bollettino del Museo Regionale di Scienze naturali di Torino, 3, 131–172.
Bogutskaya, N.G. & Iliadou, K. (2006) Rutilus panosi, a new roach from western Greece (Teleostei: Cyprinidae).
Zoosystematica Rossica, 14, 293–298.
Böhlke, E.B. (1984) Catalogue of type specimen in the ichthyological collection of the Academy of Natural Sciences of
Philadelphia. Academy of Natural Sciences, Special Publication, 14, 1–246.
Durand, J.D., Guinand, B. & Bouvet, V. (1999) Local and global multivariate analysis of geographical mitochondrial DNA
variation in Leuciscus cephalus L. 1758 (Pisces: Cyprinidae) in the Balkan Peninsula. Biological Journal of the Linnean
Society, 67, 19–42.
http://dx.doi.org/10.1111/j.1095-8312.1999.tb01927.x
Economidis, P.S. (1991) Check list of freshwater fish of Greece. Recent status of threats and protection. Hellenic Society for the
Protection of Nature, Athens, 48 pp.
Economidis, P.S. (1995) Endangered freshwater fishes of Greece. Biological conservation, 72, 201–211.
http://dx.doi.org/10.1016/0006-3207(94)00083-3
Economidis, P.S. & Banarescu, P. (1991) The distribution and origins of freshwater fish in the Balkan peninsula, especially in
Greece. Internationale Revue gesamten Hydrobiologie, 76, 257–283.
http://dx.doi.org/10.1002/iroh.19910760209
Economidis, P.S. & Sinis, A.I. (1982) Les poissons du système des lacs Koronia et Volvi (Macédonie, Grèce). Biologia Gallo-
Hellenica, 9, 291–317.
Heckel, J.J. (1943) Ichthyologie. In: Russenger, J. (Ed.), Reisen in Europa, Asien und Afrika mit besonder Rücksicht auf die
naturwissenschaftlichen Verhältnisse der betreffenden Länder, unternommen in den Jharen 1835 bis 1841. Erster band.
Reise in Griechenland, Unteregypten im nördlichen Syrien und südöstlinchen Klein Asien. Zweiter Theil. Stuttgart
(Schweizerbart), pp. 991–1099.
Howes, G.J. (1981) Anatomy and phylogeny of the Chinese major carp Ctenopharyngodon Steind. 1866 and
Hypophthalmyichtys Blk. 1860. Bulletin of the British Museum of Natural History (Zool.), 41, 1–52.
ICZN (1999) International Commission on Zoological Nomenclature. International Code of zoological nomenclature. 4
th
Edition. The International Trust for Zoological Nomenclature, London, 306 pp.
IUCN (2012) IUCN Red List categories and criteria: version 3.1 IUCN, Species Survival Commission. Gland, Switzerland and
Cambridge, UK, 32 pp.
IUCN (2013) IUCN Red List of Threatened Species. Version 2013.2. Available from: www.iucnredlist.org (accessed 20 May
2014)
Ivanović, B.M. (1973) Ichthyofauna of Skadar Lake. Montenegro, Titograd, 146 pp.
Jordan, D.S. & Gilbert, C.H. (1883) Synopsis of fishes of North America. Bulletin of the United States National Museum, 16,
1–1018.
Karaman, S.L. (1924) Pisces macedoniae. Hrvatska Stamparija, Split, 90 pp.
Ketmaier, V., Bianco, P.G., Krivokapic, M., Cobolli, M. & De Matthaeis, E. (2004) Molecular phylogeny of two lineages of
Leuciscinae cyprinids (Te l e s t e s and Scardinius) from the peri-Mediterranean area based on cytochrome b data. Molecular
Phylogenetics and Evolution, 32, 1061–1071.
BIANCO & KETMAIER
402
·
Zootaxa 3841 (3) © 2014 Magnolia Press
http://dx.doi.org/10.1016/j.ympev.2004.04.008
Ketmaier, V., Bianco, P.G. & Durand, J.D. (2008) Molecular systematics, phylogeny and biogeography of roaches (Rutilus,
Teleostei, Cyprinidae). Molecular Phylogenetics and Evolution, 49, 362–367.
http://dx.doi.org/10.1016/j.ympev.2008.07.012
Kottelat, M. (1997) European freshwater fish. Biologia, 52 (Supplement), 1–271.
Kottelat, M. & Freyhof, J. (2007) Handbook of European Freshwater Fishes. Kottelat, Cornol, Switzerland and Frehyof,
Berlin, Germany, 646 pp.
Leonardos, D., Kagalou, I.I., Tsoumani, M. & Economidis, P.S. (2008) Fish fauna in a protected Greek lake: biodiversity,
introduced fish species over a 80-year period and their impacts on the ecosystem. Ecology of Freshwater Fish, 17,
165–173.
http://dx.doi.org/10.1111/j.1600-0633.2007.00268.x
Larmuseau, M.H.D., Frehyof, J., Volkaert, F.A.M. & Houdt, J.K.J. van (2009) Matrilinear phylogeography and demographical
patterns of Rutilus rutilus: implications for taxonomy and conservation. Journal of Fish Biology, 75, 332–353.
http://dx.doi.org/10.1111/j.1095-8649.2009.02322.x
Lasca, N.P., Radulovic,V., Ristic, R.J. & Cherkauer, D.S. (1981) Geology, hydrology, climate and bathymetry of Lake Skadar.
In: Karaman, G.S. (Ed.), The biota and Limnology of Lake Skadar. Smithsonian Institution D.C. USA, printed in
Yugoslavia, Titograd, pp. 17–38.
Mačrovcić, M., Brigić, A., Buj, I., Ćaleta, M., Mustafić, P. & Zanella, D. (2006) Red book of freshwater fishes of Croatia.
Ministry of Culture, State Institute for Nature Protection, Zagreb, 253 pp.
Marić, D. (1988) The species revision of genus Rutilus Raf. from western part of Balkan peninsula. Prirode Prirodnjackog
Museja Titograd, 21, 55–79.
Marić, D. (2010) Rutilus albus sp. n. (Teleostei: Cyprinidae) from Lake Skadar. Periodicum Biologorum, 112, 153–158.
Milošević, D., Winkler, K.A., Marić, D. & Weiss, S. (2011) Genotypic and phenotypic evaluation of Rutilus spp. from Skadar,
Ohrid and Prespa Lakes supports revision of endemic as well as taxonomic status of several taxa. Journal of Fish Biology,
79, 1094–1100.
http://dx.doi.org/10.1111/j.1095-8649.2011.03090.x
Puzzi, C.M., Trasforini, S., Bardazzi, M.A., Polisciano, N., Montonati, S. & Maggio, A. (2007) Carta Provinciale Delle
Vocazioni Ittiche della Provincia Di Milano. Servizio Gestione Attività Venatoria e Piscatoria, Milano, 447 pp.
Soric, V.M. (1983) Rutilus rubilio (Cyprinidae, Pisces) U Ohrid-Drim-Skadar sistemu. Acta Biologica Jugoslavica,
Biosistematika, 9, 61–70.
Tigano, C. & Ferrito, V. (1986) Sulla presenza di Rutilus rubilio (Bp. 1837) in Sicilia (Pisces,Cyprinidae). Animalia, 13,
109–124.
Tsigenopoulos, C.S. & Berrebi, P. (2000) Molecular phylogeny of north Mediterranean freshwater barbs (Genus Barbus:
Cyprinidae) inferred from cytochrome b sequences: biogeographic and systematic implications. Molecular Phylogenetics
and Evolution, 14, 165–179.
http://dx.doi.org/10.1006/mpev.1999.0702
Vladikov, V.D. & Petit, G. (1930) Sur quelques poisons d’eau douce d’Albanie. Bulletin de la Societé Zoologique de France,
55, 383–409.
Volta, P. & Jepsen, N. (2008) The recent invasion of Rutilus rutilus (L.), (Pisces: Cyprinidae) in a large South-Alpine lake:
Lago Maggiore. Journal of Limnology, 67, 163–170.
http://dx.doi.org/10.4081/jlimnol.2008.163
... Among these, more than 60 fish species are listed, including bony fishes and cyclostomes. Focusing the attention on the species recorded in the "Coastal Lagoon" habitats of the Mediterranean Sea, on the basis of data provided by Member States [3], it was possible to fill the list in Table 3 [3, [10][11][12][13][14], further completed with the information on the Conservation status under the Habitat Directive and the red list assessments compiled by the International Union for the Conservation of Nature (IUCN) [13]. This list includes 26 species that could be grouped in functional guilds, according to the classification proposed by Potter et al. [9]: anadromous fishes (five species), freshwater fishes (17 species) and lagoon residents (four species). ...
... Table 3. List of fish species in Habitat Directive's Annex II and their presence in the habitats 1150* Coastal lagoons in the Mediterranean Sea basin. The first column included the species as listed in HD, and the second column the currently accepted nomenclature, verified by consulting [10][11][12]. IUCN category abbreviations as follow: DD-Data deficient; LC-Least concern; NT-Near Threatened; VU-Vulnerable; EN-Endangered; CR-Critically Endangered; NA-Not Assessed. Conservation status abbreviations as follow: FV-Favourable; U1-Unfavourable/Inadequate; U2-Unfavourable/Bad; XX-unknown. ...
Article
Full-text available
Transitional waters are fragile ecosystems with high ecological, social and economic values, that undergo numerous threats. According to the information provided by European Member States in the framework of the European Directive 92/43/EEC (Habitat Directive), the main threat to these ecosystems is represented by morphological and hydrological changes. The present work focuses on six lagoon fish species included in the Habitat Directive annex II (species requiring conservation measures: Aphanius fasciatus, A. iberus, Knipowitschia panizzae, Ninnigobius canestrinii, Valencia hispanica and V. letourneuxi) that spend their entire life cycle in the Mediterranean priority habitat 1150* “Coastal lagoons”. The overview of the current scientific literature allowed us to highlight how the presence and abundance of these species may provide important indications on the conservation status of coastal lagoon habitats. In fact, their occurrence, distribution and biology depend on the presence of peculiar structures, such as salt marshes, small channels, isolated pools and oligohaline areas. Coastal lagoon fragmentation and habitat loss have led to a significant reduction in genetic diversity or local population extinction. Although Aphanius and gobies have been shown to survive in eutrophic environments, it is clear that they cannot complete their life cycle without salt marshes (mainly Aphanius) and wetland areas (mainly gobies).
... Distribuzione ed ecologia in provincia di Siena Nei corsi d'acqua del versante tirrenico della Toscana il triotto è alloctono (transfaunato), introdotto agli inizi degli anni '90 del secolo scorso (Bianco, 1994;Nocita, 2002) con i ripopolamenti effettuati a scopi alieutici con materiali provenienti da impianti di acquacoltura della Pianura Padana (Bianco, 1994;Nocita, 2002)Bianco, 1994;Bianco, 1995b;Kottelat & Freyhof, 2007;Fortini, 2011;IUCN Comitato Italiano, 2015).Tassonomia Confusa per molto tempo con il triotto (cf.Tortonese, 1970), solo negli anni '80 è stata riconosciuta come entità distinta (Gandolfi et al., 1991). Recentemente è stato proposto di collocarla nel genere monotipico Sarmarutilus (Bianco & Ketmaier, 2014).Gandolfi et al., 1991;Bianco, 1995b;Kottelat & Freyhof, 2007;Fortini, 2011;Froese & Pauly, 2015(Bianco, 1994;Bobbio & Sala, 2007;Kottelat & Freyhof, 2007;Fortini, 2011;Gallo et al., 2012;Froese & Pauly, 2015).Gandolfi et al., 1991;Balestrieri et al., 2006;Kottelat & Freyhof, 2007;Fortini, 2011;Froese & Pauly, 2015).Kottelat & Freyhof, 2007;Ketmaier et al., 2004;Bianco, 2014).Gandolfi et al., 1991;Kottelat & Freyhof, 2007;Fortini, 2011;Froese & Pauly, 2015Kottelat & Freyhof, 2007;Froese & Pauly, 2015;IUCN Comitato Italiano, 2015).Kottelat & Freyhof, 2007;Bruslé & Quignard, 2013;Froese & Pauly, 2015).Kottelat & Freyhof, 2007;Bianco & Delmastro, 2011;Bianco, 2014a), 1970;Gandolfi et al., 1991;Zerunian, 2002), è oggi ritenuta distinta tanto da un punto di vista morfologico che genetico (Kottelat & Freyhof, 2007;Ketmaier et al., 2004;Bianco, 2014a). In via preliminare, si attribuiscono a questa specie le popolazioni del Torrente Foenna in Val di Chiana, sulla base dei loro caratteri morfologici. ...
... Distribuzione ed ecologia in provincia di Siena Nei corsi d'acqua del versante tirrenico della Toscana il triotto è alloctono (transfaunato), introdotto agli inizi degli anni '90 del secolo scorso (Bianco, 1994;Nocita, 2002) con i ripopolamenti effettuati a scopi alieutici con materiali provenienti da impianti di acquacoltura della Pianura Padana (Bianco, 1994;Nocita, 2002)Bianco, 1994;Bianco, 1995b;Kottelat & Freyhof, 2007;Fortini, 2011;IUCN Comitato Italiano, 2015).Tassonomia Confusa per molto tempo con il triotto (cf.Tortonese, 1970), solo negli anni '80 è stata riconosciuta come entità distinta (Gandolfi et al., 1991). Recentemente è stato proposto di collocarla nel genere monotipico Sarmarutilus (Bianco & Ketmaier, 2014).Gandolfi et al., 1991;Bianco, 1995b;Kottelat & Freyhof, 2007;Fortini, 2011;Froese & Pauly, 2015(Bianco, 1994;Bobbio & Sala, 2007;Kottelat & Freyhof, 2007;Fortini, 2011;Gallo et al., 2012;Froese & Pauly, 2015).Gandolfi et al., 1991;Balestrieri et al., 2006;Kottelat & Freyhof, 2007;Fortini, 2011;Froese & Pauly, 2015).Kottelat & Freyhof, 2007;Ketmaier et al., 2004;Bianco, 2014).Gandolfi et al., 1991;Kottelat & Freyhof, 2007;Fortini, 2011;Froese & Pauly, 2015Kottelat & Freyhof, 2007;Froese & Pauly, 2015;IUCN Comitato Italiano, 2015).Kottelat & Freyhof, 2007;Bruslé & Quignard, 2013;Froese & Pauly, 2015).Kottelat & Freyhof, 2007;Bianco & Delmastro, 2011;Bianco, 2014a), 1970;Gandolfi et al., 1991;Zerunian, 2002), è oggi ritenuta distinta tanto da un punto di vista morfologico che genetico (Kottelat & Freyhof, 2007;Ketmaier et al., 2004;Bianco, 2014a). In via preliminare, si attribuiscono a questa specie le popolazioni del Torrente Foenna in Val di Chiana, sulla base dei loro caratteri morfologici. ...
... Valid species names used were in accordance with Barbieri et al. (2015), for consistency with recent inventory and monitoring studies. In some instances, we retained the original genus of taxa for which taxonomic revisions have been recently proposed, pending further acceptance (e.g., we retained the genus Rutilus that was resurrected to the genus Leucos by Bianco & Ketmaier, 2014). Throughout the text, we use the terms taxa or species interchangeably, depending on the targeted analysis. ...
Article
Full-text available
This study provides an annotated checklist of the freshwater fish species recorded in lentic ecosystems of Greece. Species distributional data were derived from an extensive review of published and unpublished sources and were evaluated for their consistency, adequacy and reliability. Overall, 650 freshwater lentic ecosystems (149 natural and 501 artificial) were investigated from which, 480 were fishless or fish data were unavailable. In the remaining 170 ecosystems, 122 taxa were recorded (belonging to 22 families and 57 genera). Some of the records concerned species with extirpated populations, species of doubtful occurrences and taxa of uncertain taxonomic status. The highest species richness (52 species) was recorded in the Ionian ecoregion while endemicity was highest in the Macedonia-Thessaly ecoregion (43.75%). The Eastern Aegean ecoregion presented the highest level of introduced species (75%). Only eight ecosystems hosted 20 or more species, while 92 ecosystems had over 50% of introduced species. The most widespread native species (Anguilla anguilla and Luciobarbus albanicus) were two highly migratory species, highlighting the importance of connectivity of aquatic ecosystems. The most widespread translocated species was Cyprinus carpio due to its high commercial and recreational value, whereas Gambusia holbrooki confirmed its high dispersal ability being the most widespread alien species. More than one-third of the species recorded (37.23 %) are threatened based on the IUCN Red List. The ultimate contribution of this study is to support the coordination and dissemination of baseline information on the freshwater fishes of Greek lentic ecosystems, required by a range of users. This knowledge will enable further understanding of the regional assemblage structure and other biogeographical patterns of the ichthyofauna of Greece and will contribute to species conservation by prioritising areas hosting species in need of protection. Finally, this study reveals the high knowledge gap regarding species composition in many Greek lentic ecosystems and highlights the need for relevant primary research including more systematic and standardised samplings.
... The cactus roach is rheophilic, requiring fast-flowing gravel substrates for spawning; found in medium to large-sized rivers . Previously grouped with R. pigus, found in Italy and Switzerland (Bianco & Ketmaier 2014). ...
Research
Full-text available
We reviewed the potential impact of large-scale hydropower expansion on the conservation status and extinction threat of 113 freshwater fish species on the Balkan Peninsula. Each of these species is listed in one of three IUCN threat categories and/or listed in one or more annexes of the European Habitats Directive or Bern Convention. For 81 of these species, GIS-based distribution maps have been overlaid with distributions of existing hydropower facilities (ca. 1,000), those in construction (ca. 180), or those planned (ca. 2,800) to explicitly demonstrate the spatial dimension of potential habitat loss. Based on these plans, we predict that up to 49 freshwater fish species are faced with either the threat of extinction or loss of between 50 and 100% of their Balkan distribution. Of these, eleven endemic species are threatened with extinction, seven will become critically endangered, and the number of endangered species will double to twenty-four. For 68 of 69 endemic species, habitat losses are estimated between 30 and 100%, resulting in increased levels of endangerment for essentially the entire endemic fauna. Additionally, the four migratory sturgeon species would essentially lose their potential for rehabilitation in the lower Danube if additional dams were constructed there. Three exemplary hotspots of biodiversity and threat were further described: The Neretva basin in Bosnia-Herzegovina and Croatia with fourteen listed species, the Morača/Skadar system in Montenegro and Albania with eleven listed species, and the Tara/upper Drina system in Montenegro and Bosnia-Herzegovina with nearly 200 kilometers of free-flowing riverine habitat. An additional six rivers of strategic conservation interest (the Sava in Slovenia and Croatia; the Kolpa on the Croatian-Slovenian border; the Una on the Croatian-Bosnian-Herzegovinian border, the Lim in Montenegro; and the Sana in Bosnia-Herzegovina) are described with respect to their length and value in conserving self-sustaining populations of key species, such as the endangered huchen. Several smaller rivers (Cetina River, Croatia; Kalamas River, Greece; Treska River, Republic of Macedonia) are discussed as critical habitat for species, whose global existence is endangered.
... The cactus roach is rheophilic, requiring fast-flowing gravel substrates for spawning; found in medium to large-sized rivers . Previously grouped with R. pigus, found in Italy and Switzerland (Bianco & Ketmaier 2014). ...
Technical Report
Full-text available
We reviewed the potential impact of large-scale hydropower expansion on the conservation status and extinction threat of 113 freshwater fish species on the Balkan Peninsula. Each of these species is listed in one of three IUCN threat categories and/or listed in one or more annexes of the European Habitats Directive or Bern Convention. For 81 of these species, GIS-based distribution maps have been overlaid with distributions of existing hydropower facilities (ca. 1,000), those in construction (ca. 180), or those planned (ca. 2,800) to explicitly demonstrate the spatial dimension of potential habitat loss. Based on these plans, we predict that up to 49 freshwater fish species are faced with either the threat of extinction or loss of between 50 and 100% of their Balkan distribution. Of these, eleven endemic species are threatened with extinction, seven will become critically endangered, and the number of endangered species will double to twenty-four. For 68 of 69 endemic species, habitat losses are estimated between 30 and 100%, resulting in increased levels of endangerment for essentially the entire endemic fauna. Additionally, the four migratory sturgeon species would essentially lose their potential for rehabilitation in the lower Danube if additional dams were constructed there. Three exemplary hotspots of biodiversity and threat were further described: The Neretva basin in Bosnia-Herzegovina and Croatia with fourteen listed species, the Morača/Skadar system in Montenegro and Albania with eleven listed species, and the Tara/upper Drina system in Montenegro and Bosnia-Herzegovina with nearly 200 kilometers of free-flowing riverine habitat. An additional six rivers of strategic conservation interest (the Sava in Slovenia and Croatia; the Kolpa on the Croatian-Slovenian border; the Una on the Croatian-Bosnian-Herzegovinian border, the Lim in Montenegro; and the Sana in Bosnia-Herzegovina) are described with respect to their length and value in conserving self-sustaining populations of key species, such as the endangered huchen. Several smaller rivers (Cetina River, Croatia; Kalamas River, Greece; Treska River, Republic of Macedonia) are discussed as critical habitat for species, whose global existence is endangered.
... The cactus roach is rheophilic, requiring fast-flowing gravel substrates for spawning; found in medium to large-sized rivers . Previously grouped with R. pigus, found in Italy and Switzerland (Bianco & Ketmaier 2014). ...
Technical Report
Full-text available
We reviewed the potential impact of large-scale hydropower expansion on the conservation status and extinction threat of 113 freshwater fish species on the Balkan Peninsula. Each of these species is listed in one of three IUCN threat categories and/or listed in one or more annexes of the European Habitats Directive or Bern Convention. For 81 of these species, GIS-based distribution maps have been overlaid with distributions of existing hydropower facilities (ca. 1,000), those in construction (ca. 180), or those planned (ca. 2,800) to explicitly demonstrate the spatial dimension of potential habitat loss. Based on these plans, we predict that up to 49 freshwater fish species are faced with either the threat of extinction or loss of between 50 and 100% of their Balkan distribution. Of these, eleven endemic species are threatened with extinction, seven will become critically endangered, and the number of endangered species will double to twenty-four. For 68 of 69 endemic species, habitat losses are estimated between 30 and 100%, resulting in increased levels of endangerment for essentially the entire endemic fauna. Additionally, the four migratory sturgeon species would essentially lose their potential for rehabilitation in the lower Danube if additional dams were constructed there. Three exemplary hotspots of biodiversity and threat were further described: The Neretva basin in Bosnia-Herzegovina and Croatia with fourteen listed species, the Morača/Skadar system in Montenegro and Albania with eleven listed species, and the Tara/upper Drina system in Montenegro and Bosnia-Herzegovina with nearly 200 kilometers of free-flowing riverine habitat. An additional six rivers of strategic conservation interest (the Sava in Slovenia and Croatia; the Kolpa on the Croatian-Slovenian border; the Una on the Croatian-BosnianHerzegovinian border, the Lim in Montenegro; and the Sana in Bosnia-Herzegovina) are described with respect to their length and value in conserving self-sustaining populations of key species, such as the endangered huchen. Several smaller rivers (Cetina River, Croatia; Kalamas River, Greece; Treska River, Republic of Macedonia) are discussed as critical habitat for species, whose global existence is endangered.
... There are several species found in Adriatic area which today's distribution belongs to western and south-western Europe and can be defined as former paleo Po distribution during glacial conditions. Some of those species are Rana latasteiBoulenger, 1879 (Gasc et al. 2004, Kuljerić 2011, two fish species, Barbus plebejus Bonaparte, 1839, Leucos aula (Bonaparte, 1841), Romanogobio benacensis (Pollini, 1816)(Kottelat & Freyhof 2007, Bianco & Ketmaier 2014, Jelić et al. 2016) and two historical data of Vipera aspis francisciredi today's rivers of north Adriatic basin in Croatia were tributary rivers of Po River(Maselli et al. 2011) and these populations maintained in Croatia are possible remains of those historical populations which are now isolated by Adriatic Sea. ...
Article
Full-text available
Based on known literature data and data collected during field research in 2014 and 2015, we present updated distribution map of the common spadefoot toad (Pelobates fuscus) in Western Balkans (Bosnia and Herzegovina and Croatia). Pelobates fuscus is listed as least concern on the global IUCN Red List, data deficient in Croatian Red Book and the populations are in constant decline. Until year 2014 this species was only suspected to inhabit Bosnia and Herzegovina. Today we have confirmed localities in Posavina region (western, central and eastern part) presented with precise coordinates and elevation. In this paper we also present first findings of tadpoles, more precisely, the reproductive sites of the common spadefoot toad in the Bosnia and Herzegovina. In Croatia, this species is found along the rivers Mura, Drava and Sava, including most lowland areas up to 300 m above sea level. Historical records of Pelobates fuscus in Adriatic region are discussed and compared with distribution area of Italian isolated population.
... All the environmental variables, except the average current speed, were significantly correlated with this axis, which effectively described the changes that occurred table ii. -Updated list of species found in the study area according to recent revisions of Bianco (2014) and Bianco and Ketmaier (2014), and their origin. For native species is also reported the threatened status according to iUCn reD List criteria for italy (rondinini et al., 2013), and year of introduction for alien species in the tiber basin and reference, when available. ...
Article
The purpose of the research was to analyse the distribution, abundance and growth of Carassius spp., a taxon which includes non-native species introduced into the water bodies of the Umbrian portion of the Tiber River basin during the late 1980s. Additionally, the study estimated the relative weight for Barbus tyberinus, Squalius lucumonis and Sarmarutilus rubilio to investigate the presence of competitive interactions between Carassius spp. and these native species that characterize the middle and lowland reaches of the Umbrian watercourses. Currently, Carassius spp. is widely distributed in both the stagnant and the slow-running waters of the entire basin. The presence of these species was recorded in 34.81% of the total sampling sites; the results showed that the presence of Carassius spp. was associated with the presence of other non-native species and with poor water quality. In total, 4520 specimens of Carassius spp. were collected and the age composition ranged from 0+ to 9+. The equation for the length-weight relationship (TL-W) was determined as W = 0.010TL3.180. For the Etruscan chub Squalius lucumonis, an endemic species of central Italy, the mean value of the relative weight was significantly higher when Carassius spp. was absent. The results of the research suggest the need to establish effective management conservation strategies for this endemic species.
Article
Full-text available
A checklist of the freshwater fish fauna of Croatia is presented for the first time. It is based on 1360 publications of historical and recent data in the literature. According to the literature review, there were 137 fish species in 30 families and 75 genera recorded in Croatia. The checklist is systematically arranged and provides distributional data of the freshwater fish fauna as well as whether the species is endemic, introduced or translocated.
Article
Full-text available
Two species from the Adriatic drainage basin of western part of Balkan Peninsula: Rutilus basak (Heckel) and Rutilus prespensis (Karaman) have been described in this paper. There were recorded an d described two subspecies of each species: Rutilus basak basak (Heckel) from Hertzegovina and Rutilus basak ohridanus (Karaman) from Skadar Lake, Rutilus prespensis prespensis (Karaman) from Prespa Lake and rutilus prespensis vukovici ssp.n, from Skadar and Šaso Lakes. The most significant morphological properties of the subspecies have been presented.
Article
Full-text available
Rutilus panosi sp. n. is described from Acheloos River drainage, Western Greece. It is distinguished from the other species of Rutilus by the following combination of characters: (40)41-44(45) lateral line scales; (11)12-14(15) gill rakers; pharyngeal teeth formula 6-5; commonly 9½ branched anal rays and 9½ branched dorsal rays; snout pointed, triangular; maximum body depth 25-27% SL; dorsal fin depth 19-22% SL; anal fin depth 12-14% SL; snout length 26-32% head length; eye diameter 20-24% head length; 40-41total vertebrae (common formulae 23+17, 23+18 and 22+18), absence of any lateral stripe. A comparison with species of Rutilus s. str. from the Balkan Peninsula is given.
Article
Discusses efforts to protect some endangered fishes of western North America by placing representative samples in controlled environments until recovery actions reduce or eliminate threats to their survival. Efforts for western fishes have been under way for several decades, and especially since 1974 at Dexter National Fish Hatchery, New Mexico. The contributions of the Dexter hatchery, acting as a refuge and research center as well as a production facility for reintroducing imperiled native fishes back into nature, have been substantial. It is nonetheless only an interim solution to immediate problems of species disappearing in nature. -from Editors