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A taxonomic study on the genus Phoxinus (Acthinopterigy, Cyprinidae) from Italy and western Balkans with description of four new species: P. ketmaieri, P. karsticus, P. apollonicus and P. likai

Authors:
  • Regional Environmental Protection Agency Lazio Italy

Abstract and Figures

The genus Phoxinus from Italy and western Balkans was analyzed. Five species are recognised: P. Phoxinus, distributed in the rivers of the Alpine slope of the Padano-Venetian, district; P. ketmaieri n.sp, described from Krk island and River Zrmanje, but whose distribution probably includes also the River Cetina and others rivers of the Dalmatian district, the rivers Krka and Neretva. P. karsticus described from the Endorheic-system of the River Trebinje, Bosnia-Herzegovina is another representative of the Dalmatian-Croatian district; P.apollonicus described from the Moraca-Zeta River system, Lake Skadar drainage, is the representative species of the Albanian district. Finally, P. likai should be regarded as an endemic species of the extensive endorheic river system of the Lika region and underwater connected rivers of the Bosnia Erzegovina-Croatian district.
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Pier Giorgio Bianco
Gabriele de Filippo
Editors
Researches on Wildlife Conservation
Volume 4
A taxonomic study on the genus A taxonomic study on the genus Phoxinus (Acthinopterigy, Cyprinidae)
from Italy and western Balkans with description of four new species:
P. ketmaieri, P. karsticus, P. apollonicus and P. likai
Pier Giorgio Bianco and Salvatore De Bonis
IGF Publishing
P. likai
Pier Giorgio Bianco
Gabriele de Filippo
Editors
Researches on Wildlife Conservation
volume 4
IGF Publishing
Reference
Bianco P.G. e de Filippo G. (eds.) 2015. Res.Wildl.Conserv. 4. IGF Publ., USA.
Published by Lulu.com
© 2015 Istituto di Gestione della Fauna onlus
info@gestionefauna.com
www.gestionefauna.com
Content
A taxonomic study on the genus Phoxinus (Acthinopterigy,
Cyprinidae) from Italy and western Balkans with description
of four new species: P. ketmaieri, P. karsticus, P. apollonicus
and P. likai
Page 1
P. G. Bianco and S. De Bonis
Researches on Wildlife Conservation, vol. 4, 2015, IGF publ.
A taxonomic study on the genus Phoxinus
(Acthinopterigy, Cyprinidae) from
Italy and western Balkans with description of four new species: P. ketmaieri,
P. karsticus, P. apollonicus and P. likai
Pier Giorgio Bianco* e Salvatore De Bonis
Ichthyological laboratory, via Paparelle al Pendino, 5, 80134 Naples (Italy)
*Correspondence author : gibianco@unina.it
Abstract
The genus Phoxinus from Italy and western Balkans was analyzed. Five species are recognised
: P.
Phoxinus, distributed in the rivers of the Alpine slope of the Padano-Venetian, district; P. ketmaieri
n.sp, described from Krk island and River Zrmanje, but whose distribution probably includes also
the River Cetina and others rivers of the Dalmatian district, the rivers Krka and Neretva. P.
karsticus
described from the Endorheic-system of the River Trebinje, Bosnia-Herzegovina is
another representative of the Dalmatian-Croatian district; P.apollonicus described from the Moraca-
Zeta River system, Lake Skadar drainage, is the representative species of the Albanian district.
Finally, P. likai shuold be regarded as an endemic species of the
extensive endorheic river system
of the Lika region and underwater connected rivers of the Bosnia Erzegovina-Croatian district.
Studio tassonomico del genere Phoxinus in Italia e nei Balcani occidentali e descrizione di
quattro nuove specie P. ketmaieri, P. karsticus, P. apollonicus and P.likai.
Il genere Phoxinus
da Italia e Balcani occidentali è stato analizzato. Cinque specie sono state
identificate: P. phoxinus, distribuito nei fiumi del versante alpino del distretto Padano-Veneto; P.
ketmaieri
n.sp, descritto per l’ isola di Krk in Croazia, e il fiume Zrmanje, ma la sua distribuzione
potrebbe includere il fiumi Cetina Krka, e Neretva, del distretto della Dalmazia-Croazia; P. karsticus
descritto per il sistema carsico del fiume Trebinje in Bosnia-Herzegovina è un altro rappresentante
del distretto della Dalmazia-Croazia; P.apollonicus n.sp. descritto per il fiume Moraca, pricipale
affluente del lago di Scutari, è la specie rappresentative del distretto Albanese. Infine, P.likai n.sp
viene considerato un endemismo del vasto sistema fluviale endoreico della regione della Lika e
connessioni carsiche sotterranee dei fiumi della Bosnia Erzegovina e della Croazia adriatica.
Étude taxonomique du genre Phoxinus
en Italie et dans les Balkans occidentaux et la
description de quatre nouvelles espèces P. ketmaieri, P. karsticus, P. apollonicus et P. likai.
Le genre Phoxinus
de l'Italie et les Balkans occidentaux a été analysé. Cinq espèces ont été
identifiées: P. Phoxinus, distribué dans les rivières de la pente de la région montagneuse Padano-
Veneto; P. ketmaieri décrit pour la 'île de Krk en Croatie, et la rivière Zrmanje, mais la distribution
peut comprendre les rivières Cetina Krka et Neretva, dans le district de la Dalmatie et la Croatie; P.
karsticus
décrit de la endoréiques-système de la rivière Trebinje, Bosnie-Herzégovine est un autre
représentant de le district de Dalmatie et la Croatie; P. apollonicus
n.sp. décrit pour la rivière Moraca,
principal affluent du lac de Skadar, est les espèces représentatives du district albanais. Enfin, P. likai n.sp est
considéré comme une espèce endémique de le réseau fluvial de la région de steppe de Lika et connexions
rivières karstiques souterrains de la Bosnie-Herzégovine et de l'Adriatique de la Croatie.
Key words: Taxonomy, Cyprinidae, Phoxinini, Phoxinus, Italy and western Balkans, new species
Introduction
Phoxinus
Leuciscinae. The North-American representatives of the clade received considerable attention since they
include nearly the whole cyprinid fish fauna of the Nearctic region, with about 300 species (Simons, et al.,
2003; Strange & Mayden, 2009; Mayden & Allen, 2015).
Among the European Phoxinini, the genus Phoxinus
exception of extreme southern area of the peri-Mediterranea peninsular regions and nearly all islands. At
peculiarity of this genus is the presence, in reproductive males, of patches of breasts scales between the
pectoral fins, a character shared with genera Margariscus, Couesius, and Clinostomus. But the question of the
affinities of Phoxinus with these genera is still debated (Hing-Yu Chen, 1996).
The taxonomy of this genus in Europe has been relatively little studied in the recent past when compared to
others Palearctic genera such as Rutilus, Barbus, Chondrostoma, and Telestes
Europe, reporting seven species, three of which new to science, reporting for Italy and western Balkans, a
single species Phoxinus lumaieurl Schinz, 1849, but suggesting the potential presence of at least a new species
from the Lake Skadar system. The recent molecular analyses of Palandačić
that the genus Phoxinus in the Western Balkans and Italy entails a complex of species with at about seven
characterized by distinct ichthyogeographic districts, each of them characterized by a complex of localized
analyzed the material sampled in these districts available in the collection of the Department of Biology and
Zoological Museum of the Naples University, Italy (IZA). The collection is not exhaustive but – on a
had a heavy impact on the original geographic distribution of Phoxinus species (Kottelat, 2007).
Material and Methods
ethylic alcohol 60%. All the samples included in the current analyses are preserved in the collection of the
Department of Biology and Zoological Museum of Naples University, Italy (acronym: IZA).
Figure 1 – Map showing the approximate locations of examined materials in Italy and Western Balkans
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Counts, and measurements and the description of the position of patch of breasts scales follow Kottelat
(2007), except for the standard length (SL), which was measured from the tip of snout to the upper
insertion of the caudal fin. Due to the reduced size of fish, most counts and measurements were taken with
the aid of a millimeter digital caliper and a stereoscopic microscope; all measurements were taken from the
left side of the body. Pharyngeal bones were dissected mainly from the left side, but in several cases also
from the right side. Laboratory illustration has been made with a scanner, Epson perfection 3170 photo,
using sensibility from 350 up to 1200 DPI. The sampling localities of the examined specimens are reported
in Fig. 1
.
Systematic accounts
Phoxinus phoxinus (Linnaeus, 1758)
(Fig. 2)
Figure 2 – Female, 49 mm SL, above, and Male, 53 mm sl below of P. phoxinus from the Po river basin.
Synonym. Phoxinus lumaireul Schinz, 1849: 331 (type locality, Lake Major).
Examined material.
IZA 0427, 9 specimens, 47-54 mm SL, Vicenza, River Bacchiglone, 4 April 1996, P. G.
Bianco coll. – IZA 00299, 6 specimens, 38-54 mm SL, stream Chiamogna, western tributary of the Po
basin, 2 May 1980, P. G. Bianco coll. - IZA 83134, 9 specimens, 36-53 mm SL, Fosso Pinzano, River
Tagliamento, Friuli, Italy, 15 April 1982, P. G. Bianco coll) – IZA uncat., 5 es, 41-58 mm SL, United
Kingdom, River Frame, East Stoke, Dorset, 22 April 1982, A. Wheeler coll.
According to Kottelat (2007), the species living in Italy and Western Balkans is Phoxinus lumaireul. A detailed
comparison of my samples from upper Adriatic rivers with the extensive neotype description given by
Kottelat (2007) for Phoxinus phoxinus and with a sample of P. phoxinus from England shows P. lumaireul
be a
synonym of P. phoxinus
; this conclusion – already reported in Bianco (2014) – is also supported by the
molecular data of Palandačić (2015). The Phoxinus
inhabiting water bodies from Croatia to Albany and
Montenegro represents a distinct complex of yet undescribed species.
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In the following re-description of P. phoxinus
the pharyngeal teeth formulae, were not reported by Kottelat (2007) but we found that the combination of
them are diagnostic especially for the new species described below.
Re-description. In Phoxinus phoxinus from Italy the patches of breast scale are separated (Fig 3, 4) (the
base (Fig 2) (lateral line usually reaching beyond anal base); Depth of caudal peduncle 11-12 % SL (8-11
%SL); 2.4-2-8 in its length (2.6-3.1 times), 1.9-2.3 times in body depth (2.1-2.6 times body depth), patched
of breast scales separated by unscaled area (the same). Lateral line from incomplete to nearly complete
(neotype has a nearly complete pored lateral line scales). In addition, P. phoxinus
in longitudinal series and 30-34 circumpeduncular scales. The pharyngeal teeth formula is Italy is usually 5.2
on both sides, as in P. phoxinus from England (Fig 5).
Figure 3 – 4 . Ventral region of a male 54 mm SL from River Bacchiglione showing well separated patches of breast scale
(left side) - Ventral view of anterior part of body of a male 53 mm SL from River Tagliamento, showing separated
patches of breast scales (right side).
Figure 5 – A: Left pharyngeal bone, of about 3 mm of length, of a fish 54 mm SL from river Po.
B: Left pharyngeal bone of about 3 mm from a fish 53 mm SL from England.
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Total gill rakers, 6-8; branched rays of dorsal, anal and pelvic fin, 7; free margin of dorsal and anal fins
straight to slightly convex (Fig 6); dorsal profile of the body, slightly convex; longitudinal stripe extending
from the tip of snout to the end of caudal peduncle were, at base of caudal fin, is enlarged in a roundish
spot; about 14-16 vertical blotches accompanying 1-2 myomere of the trunk and of caudal peduncle,
sometimes covering the distinctness of the longitudinal stripe.
According to this re-description, the Italian populations cannot be distinguished from Phoxinus phoxinus as
described by Kottelat (2007).
Radmler et al., (2015), stated that according to their geometric morphometrics analyses, population of
Phoxinus from Italy and western Balkan show a deeper body and a deeper caudal peduncle when compared
with population of central European origins and stated that their results may confirm the validity of P.
lumaieurl. This seems incongruent with present data as depth of body and of caudal peduncle are highly
variable in the species here described (Table I), and body depth, according to ours results, seems deeper in
P. phoxinus from transalpine countries, in contrast with the results of Radmler (2015). In addition, Collin &
Fumagalli (2011), according to molecular and geometric morphometrics analyses, founds that shape of
body (but also several others phenotypic and genotypic characters) of P. phoxinus is habitat dependant and
lacustrine populations show a deeper body and caudal peduncle when compared with the riverine forms. In
conclusion, the depth of body and caudal peduncle are adaptive characters in P. phoxinus, and have little
value in taxonomy.
Figure 6 – Female, 53 mm SL, from River Tagliamento Friuli Region showing convex free margin of dorsal and anal fins.
Sexual dimorphisms: males in April have nuptial tubercles on the head and fan shaped on the free margin
of scales of the body and especially the ventral region of the caudal peduncle (Fig. 7). Pectoral fins are
longer than in female and thickened also, on dorsal faces, by minute nuptial tubercles, or “unculi” aligned
along the first six seven branched rays. Anal fin has tubercles, but with the first three rays thickened. All
tubercles disappear or just leave some vestigial remains out of spawning season. Small tubercles, fan shaped
are on the free margin of the patch of scales between pectoral fins. The scales are embricated and well
separated in males in spawning condition, but in females and in non-reproductive fishes, the patches are
apparently lacking or deeply hidden in the skin: this features of sexual dimorphisms in males Phoxinus were
described in details also by Howes (1985).
Colour pattern, In preserved specimens, the colour patterns consist in a blackish, longitudinal stripe
extending from the tip of the snout till caudal peduncle, ending in a roundish black spot on the base of
caudal fin. Vertical blotches can only be accentuated. All fins are colorless, the dorsal and the caudal fins,
with several scattered melanophores. Peritoneal membrane silvery with few scattered melanophores.
5
Figure 7 – Lateral view of the caudal peduncle of a male 49 mm SL, Phoxinus phoxinus showing minute fan shaped
tubercles on the free margin of each scale , and the thickened first 3 rays of the anal fin
In alive specimens, according to a colour slide taken on a freshly collected fish in River Astico on October
1995, but not preserved, the ventral part appears silver from the throat and the operculum to the far end of
the caudal peduncle, up to the insertion the caudal fin. The longitudinal band is obscured by thick vertical
stripes of a dark brown. The paired fins are reddish at their base, the unpaired and caudal, grayish.
Phoxinus ketmaieri, new species
(Fig 8)
Figure 8 – Holotype of Phoxinus ketmaieri, male, 43 mm SL, IZA 8823-1, Isla of Krk
Holotype: IZA 8823-1, Male 43 mm SL, brook on south-eastern slope of Krk island, Baš
ka brook at
Baška draga , Croatia, April 1982, R. Hacker coll
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Researches on Wildlife Conservation, vol. 4, 2015, IGF publ.
Paratypes: IZA 8823, 2 specimens, 31-33 mm SL, brook on south-eastern slope of Krk island, Baška
brook at Baš
ka draga, Croatia, April 1982, R. Hacker col.-l IZA 8792, 9 specimens, 37-46 mm SL, River
Zrmanje, Obrovac, Croatia, 22-24 August 1987, P. G. Bianco coll.
Diagnosis: Phoxinus ketmaieri is distinguished from other Phoxinus species from Italy and Western Balkans
by the low number of scales in longitudinal series 58-61, and number of circumpeduncular scales, 24-26.
Holotype, a male, has a depigmented body but with melanophores traces of the former pigmentation. In a
paratype from Zrmanje (Fig 9) the longitudinal stripe extends from the tip of snout to the end of caudal
peduncle where it ends in a roundish spot. The holotype has breast blotches of scales separated (Fig 10).
The pharyngeal teeth formula is usually 4.1 or 4.2. and meristics are reported in Table I and II. Largest
specimens analyzed was a female 46 mm SL from River Zrmanie. Body moderately elongate. Dorsal profile
convex sometimes with a slightly pronounced hump, dorsal profile of the head slightly more convex than
ventral, snout blunt, mouth slightly subterminal, chin rounded.
Figure 9 - Paratype of Phoxinus ketmaieri, 44 mm SL, female, River Zrmanje
Description. The outline of the species is shown in figs 6 and 7. Detailed morphometrics and meristics are
reported in Table I and II. Largest specimens analyzed was a female 46 mm SL from River Zrmanie. Body
moderately elongate. Dorsal profile convex sometimes with a slightly pronounced hump, dorsal profile of
the head slightly more convex than ventral, snout blunt, mouth slightly subterminal, chin rounded. Dorsal
fin with 3 simple and 7 branched rays; free margin of dorsal and anal fin straight to slightly convex; anal fin
with one simple followed by 6-7 branched rays; pectoral fin rounded with one simple and 14-15 branched
rays; in males may nearly reach and overpass the insertion of pelvic fin, in females reaches 2/3 of the
distance between the insertion of pelvic and ventral fin; pelvic fin with 3 simple and 7-8 branched rays free
margin of anal fin from straight to moderately convex; origin of anal fin from slightly below or at the same
level of vertical through posterior insertion of dorsal fin. About 58-64 scales in longitudinal series, lateral
line incomplete with 12-24 pored scales; in small specimens, less of 30 mm SL, the scales are incipient and
only 2-3 are pored, scales embedded but distinct, about 24-26 circumpeduncular scales. Tubercles in males,
fan shaped on the free margin of most scales of the body especially on the lower part of caudal peduncle.
Depth of caudal peduncle 1.9-2.4 times in it length; head depth 1.3-1.6 times in its length; eye diameter less
than the preorbital distance; body depth about 4.3-4.7 times in SL.
Colour pattern. The general color pattern in preserved specimens is yellowish with more or less evident
stripes or vertical bands. In most specimens, especially from River Zrmanje, the longitudinal stripe is well
marked or confused by 14-16 vertical reduced stripe which follow 1-2 myomere of the trunk and of the
caudal peduncle. Peritoneal membrane, silvery with few scattered melanophores. All fins are almost
colourless, with few scattered melanophores.
7
Figure 10 – Ventral view of anterior part of the body of the holotype of Phoxinus ketmaieri showing separated breast
patch of scales.
Distribution and habitat. Species of genus Phoxinus
are usually considered as coldwater adapted species
living in the trout region (Bianco & Ketmaier, 2015). Nevertheless the Phoxinus living in the Zrmanje should
be considered as a warm water adapted populations as, at time of collection, the temperature was 22° C,
and the fish communities were represented by typical warm water adapted species, namely chubs, bleaks,
gobies, river blennies, cobitids, and eels. On the Krk island the species was observed in a small brook not far
from the sea: Baška brook at Baš
ka draga. This island is, apparently, the only island around the
Mediterranean to host a Phoxinus
species. Its presence was firstly recorded by Heckel on 1850. Later on the
species was recorded several times and collected always from the same brook and preserved at
Naturhistorisches Museum Wien. The current conservation status of this unique population is unknown.
The distribution would probably include also the River Cetina, R. Zrmanje and probably others rivers of
the Dalmatian Ichthyogeographic district, namely, the River Krka, and Neretva.
Etymology
. From Valerio Ketmaier, a molecular biologist and friend with a long collaborative history with
the senior author.
Phoxinus karsticus, new species
(Fig, 11)
Figure 11 – Holotype of Phoxinus karsticus, female, 58 mm SL, IZA 00314-1 , Endorheic River Trebinje, Bosnia-
Herzegovina.
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Researches on Wildlife Conservation, vol. 4, 2015, IGF publ.
Holotype
. IZA 00314-1, 73 mm SL, Endorheic River Trebinje, near Jasenica Village, Bosnia Herzegovina,
24 October 1989, B. Elvira coll.
Paratypes
. IZA 00314, 4 specimens, 48-68 mm SL, endorheic river Trebinje, near Jasenica Village, Bosnia
Herzegovina, 24 October 1989, B. Elvira coll.
Diagnosis. Phoxinus karsticus distinguished from others Phoxinus
from Italy and Western Balkans by the
high number of scales in longitudinal series 81-86, and number of circumpeduncular scales, 40-42, but also
for the size reached of about 91 mm of TL. Holotype has a longitudinal band discolored or reduced.
Holotype, and paratypes has patches of breast scales well developed and united in their third front ( Fig 12).
Dorsal has three simple rays followed by 7-8 branched rays; 8 branched rays only seldom were found in all
others sample here analyzed. The pharyngeal teeth formula from left bone is 5.2 or 4.2.
Figure 12 - Ventral view of anterior part of the body of the paratype 49 mm SL, of Phoxinus karsticus showing united
patch of breast scales
One other character useful to distinguish the species, but not unique are, the position, the anterior origin of
anal fin placed in advance of vertical through posterior insertion of dorsal fin, reaching the level of the fifth
or sixth branched rays of the dorsal fin.
Description
. The outline of the holotype, a female, is shown in Fig. 11. Detailed morphometric and
meristic data are reported in Table I and II. Largest specimens analyzed was the holotype, a female 73 mm
SL with immature eggs 0.4-0.6 mm of diameter. Body quite elongate when compared with others species
here analyzed; dorsal profile, straight or moderately convex; dorsal profile of the head and snout slightly
convex; snout pointed its dorsal profile is in continuum with that of the head and back; mouth slightly sub-
terminal; eye diameter less than prerbital distance. Dorsal fin with 3 simple and 7-8 branched rays; free
margin of dorsal fin straight to slightly convex. Anal fin with three simple and 7 branched rays; free margin
of anal fin straight or slightly convex. Apparently the length of anal fin equal in both sex. Pectoral fin
rounded with one simple and 14-15 branched rays; pelvic fin with one simple followed by 7 branched rays.
Caudal peduncle 2.3-2.6 times longer than deep; head depth 1.3-1.5 times in its length; eye diameter less to
the preorbital distance; body depth about 4.4-5.0 times in SL.
9
About 81-86 scales in longitudinal series, lateral line incomplete with 23-63 pored scales; about 40-42
circumpeduncular scales. Tubercles in males incipient (fish were collected in October, outside the
reproductive season) on dorsal surface of pectoral fin and scales of body and caudal peduncle; females
carry egg of 0.2-0.6 mm of diameter.
Colour pattern. The general colour pattern in preserved specimen, is yellowish with more or less evident
stripe or vertical bands. In most specimens, especially from Krk Island, the longitudinal stripe is faintly
marked or confused by 14-16 vertical darker bars. All fins are almost colourless, with few scattered
melanophores.
Distribution and habitat. P. karsticus
is probably endemic to the Karstic Popovo Polje-Trebinje endorheic
river system.
Derivatio nominis: from the karstic area of Popovo Polje placed in westernmost area of Bosnia-
Erzegovina region.
Phoximus apollonicus, new species
(Fig. 13)
Figure 13 - Holotype of Phoxinus apollonicus, male 58 mm SL IZA 87103-1, River Moraca, Montenegro.
Holotype
. IZA 87103-1, 58 mm SL, upper River Moraca tributary to Lake Skadar, Montenegro, 26 august
1987, P.G.Bianco coll.
Paratypes. IZA 87103, 42 specimens, 23-68 mm SL, upper River Moraca tributary to Lake Skadar,
Montenegro, 26 august 1987, P.G.Bianco coll.
Diagnosis. Phoxinus apollonicus
is unequivocally
distinguished from all others Phoxinus
species by
the reduced number of teeth on pharyngeal bones.
The species usually has four teeth on external row
(Fig 14), but also 3 or 2, and 1 or 0 in the inner
row. Others characters which, combined, can help
to distinguish this species from the others: the
holotype has patches of breast scales connected
anteriorly (Fig 15); the lateral line is nearly
completed, posteriorly reaching the vertical line
crossing the upper and the lower insertion of the
caudal fin; flank of body with a marked mid-lateral
longitudinal stripe extending from tip of snout to
the base of caudal fin; a distinct spot at base of the
caudal fin; a marked black spot on the opercular
bone; depth of caudal peduncle, 2.2-2.8 times in its
length and 1.8-2.5 in body depth; eye diameter less
than the preorbital distance.
10
Figure 14 – Left pharyngeal bone of the holotype Phoxinus
apollonicus (length 2.5 mm), showing 4 teeth on external
and 0 on inner rows).
Researches on Wildlife Conservation, vol. 4, 2015, IGF publ.
Description: patches of breast scales absent in specimens less of 30 mm SL, where the scales are still
incipient. In largest specimens, breast scales are very difficult to check because deeply hidden into the skin.
Apparently, most of the paratypes do not show breast scales, only in few specimens, are connected or
separated by unscaled area. The longitudinal strip is well developed in specimens of about 25-30 mm SL,
which are nearly scale-less. In specimens of 12-18 mm SL, is still unformed. In preserved specimens, the
broad longitudinal band extend from tip of snout till the caudal peduncle, where, at the base of the caudal
fin, is enlarged in a roundish spot. 14-18 vertical stripe of melanophores, more or less expanded, follow
most of myomere of the trunk and the caudal peduncle. Pharyngeal teeth usually 4.1, but also 4.0 and 3.0;
in P. apollonicus the principal formula is 4.1-1.4, with several case of teeth placed on a single outer row (4-4;
4.1-4) (Fig 15); the lateral line is nearly complete, the pored section include about 58-64 scales, while there
are overall about 71 to 79 scales in longitudinal series. 24-26 circumpeduncular scales; 11.5-13.5 above and
10.5-12.5 below the lateral line. Dorsal fin with three simple rays followed by 7 branched rays (rarely 8).
Anal fin with constantly three simple rays followed by 7 branched rays. Pelvic fins with one simple followed
constantly by 7 branched rays; free margin of dorsal fin from straight to slightly convex; free margin of anal
fin from straight to slightly concave.
Eye roundish, iris whitish. Gill rakers on the first gill arch, left side, are incipient o very reduced with 5-7
total gill rakers.
Figure 15 - Ventral view of anterior part of the body of the holotype, of P. apollonicus showing
united patch of breast scales
Colour pattern. In preserved specimens all fins are whitish with few scattered melanophores; peritoneum
silvery with few scattered melanophores; in living specimens (from a slide taken from a specimens of River
Zeta, collected on November 1999, but not preserved) the longitudinal band is masked by very marked and
extensive vertical blotches, which, clearly, accompany one or two myomere of the trunk and of the caudal
peduncle. All fins are golden yellow; the belly and the lower part of head and the body is brilliant silvery.
11
Distribution. River Moraca and its tributary Zeta, both belonging to the Lake Skadar drainage. Possibly P.
apollonicus represents the endemic species of the Albanian ichthyogeographic district.
Derivatio nominis. From Apollonia, an ancient name to delimitate Albany.
Phoxinus likai, new species
(Fig 16)
Figure 16 – Holotype of Phoxinus likai female 47 mm SL, IZA 02263-1, River Oruca system, Bosnia-Erzegovina.
A small sample of five specimens collected in the Karstic area of Lika Valley of Croatia, in a brook
belonging to the endorheic river Oruca near Gračac, show characters that cannot be applied to any of the
four taxa described above. Indeed, this species has the patches of breast scales united at the centre. This is a
very valuable character for species identification for Phoxinus
(Kottelat, 2007). In the four species here
described, only one other species has patches of breast scales united, Phoxinus karsticus. But this species have
a nearly incomplete pored lateral line and a higher number of scales either on longitudinal series or around
the caudal peduncle. Based on this evidence, we described the small sample as a new species in the hope
to stimulate conservationists and taxonomists to undertake more researches in Croatia on this still poorly
investigated genus in Europe.
Holotype. IZA 02263-1, 47 mm SL, Croatia, River Oruca system, near the village of Gračac, 8 December
1999, P.G. Bianco coll.
Paratypes. IZA 02263, 4 specimens, 43-49 mm SL, Croatia, River Oruca system, near the village of Gračac,
8 December 1999, P.G. Bianco coll.
Diagnosis. Phoxinus likai is distinguished from Phoxinus phoxinus and the species here described by having
patches of breast scales united and continuous across breast ( Fig 17) except P. karsticus, where they are
united only on anterior part. It may be distinguished from P. karsticus also for the number of scales in
longitudinal series which are 82-87 in P. karsticus and 72-79 in P. likai and for circumpeduncular scales, 40-42
in P. karsticus and 30-32 in P. likai. Others characters useful for determination, but not unique to this
species are the pharyngeal teeth formula, 5.2 or 4.2, shared with P. poxinus but not P. apollonicus
; pored
lateral line nearly complete, reaching almost the end of the caudal peduncle, in P. ketmaieri and P. phoxinus
do not reach more than half of the length of caudal peduncle; eye large, its diameter more than the
preorbital distance, but less in the other species; snout short and blunt; longitudinal band from tip of snout
to caudal peduncle where it forms a roundish spot; the band is more evident in the second half of the
body; the flanks have 12-14 mid-lateral row of vertical elliptic blocks, more evident on the second half of
body; anterior origin of anal fin placed at same level of vertical through posterior insertion of dorsal fin.
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Figure 17- Ventral region of a male of P. likai 49 mm SL showing the united patches of breast scales.
Description: General outline of the holotype is shown in Fig 16 and morphometric and meristic data are
given in Table I and II. The largest specimen examined was 61 mm of total and 49 mm of SL. Body
moderately elongate, dorsal profile more convex than the ventral, dorsal profile of the head straight, snout
short and blunt; mouth moderately downturned; chin rounded. Dorsal fin with 3 simple and 7 branched
rays, free margin convex; anal fin with three simple and 7 branched rays, its free margin from straight to
slightly convex; when depressed it reaches about one half of caudal peduncle length; pectoral fins oval, with
one simple and 13-15 branched rays, when depressed on the flanks it may reach or overpass the insertion of
the pelvic fins, in males, in females may reach 2/3 of the pectoral-pelvic fins distance; depth of caudal
peduncle about 1.7-2.0 times its length; head depth 1.2-1.4 times its length; eye diameter 1.2-1.2 times the
preorbital distance: body depth about 4.3-4.6 times SL. Latera
l line incomplete with 23-65 pored scales;
73-79 scales on longitudinal series; 30-32 circumpeduncular scales; pharyngeal teeth hooked at the tip and
serrated; about 5.2 or 4.2 teeth on pharyngeal bones. Tubercles are present in males on the head and on
the dorsal side of the pectoral fins, which are thickened also by the presence of minute nuptial tubercles
aligned along the first six seven branched rays.
Colour pattern. In preserved specimens the lateral band extending from the tip of snout to the caudal
peduncle is more evident on the second half of the body; vertical blotches are more enlarged on sides of
caudal peduncle; a large blackish spot on the opercular bone; ventral region of body and caudal peduncle,
which extend about 1/3 of body depth, yellowish; all fins yellowish with few scattered melanophores;
peritoneal membrane silvery with few scattered melanophores.
Distribution. Probably endemic of the extensive endorheic river system of the Lika region and underwater
connected rivers of the Croatian Adriatic slope.
Derivatio nominis: from the Lika-Dinaric karstic region of Croatia.
Discussion
This contribution seems to be complementary to that of Kottelat (2007), who re-analyzed most of the
Phoxinus species from transalpine western Europe, but left somehow unsolved the taxonomy of the genus in
the western Balkan area, reporting just a single species, Phoxinus lumaieurl
, now placed in synonym with
Phoxinus phoxinus
, distributed in the Padano-Venetian district. In other district of the Western Balkans we
found a Phoxinus species complex formed by at least 4 new species; our conclusions are in partial agreement
with those based on molecular data of Palandačić
et al. (2015). We found, also, that most important
diagnostic characters are the meristic counts, principally squamation, the breast patches of scales and the
pharyngeal teeth formulae. Several proportional measurements seems to be habitat-dependant and, for
13
riverine ones (Collin & Fumagalli, 2011). Regarding the teeth on pharyngeal bones, while in Phoxinus
phoxinus from the Po basin, and others rivers of the upper Adriatic drainage and England the pharyngeal
direction with the tendency to lose one or two teeth on outer and one or both in the inner rows.
In the karstic area of Bosnia-Herzegovina, a distinct species was described from River Trebinje, an
we described the new species Phoxinus apollonicus, mainly according to the dentition of pharyngeal bone and
Palandačić et al. (2015) that in Italy and Western Balkan a complex of Phoxinus
of Palandačić et al. (2015) reporting seven different clades.
Regarding the biogeographic history of the group, we can postulate that Phoxinus phoxinus probably reached
karstic Alpine-Dinaric system of Eastern Italy and Western Slovenia and Croatia. An underground dispersal
has been already reported in cyprinids (Palandačić et al., 2012). The reason why P. phoxinus did not colonize
exclusion by either competition or predation (or a combination of both). As a matter of fact, Phoxinus
absent in all the Adriatic Italian rivers south of the Po river. This distributional pattern implies that Phoxinus
adapted species such as Salmo farioides, Cottus gobio, Telestes muticellus and the eurythermal Squalius squalus did.
These species used this phase of the Po river to reach several Adriatic drainages and the River Zrmanje in
Croatia (Bianco, 2014, Bravničar et al., 2015) and, following the orogenesis of the Apennines, to cross the
species in the head waters of the main rivers of the Tuscany-Latium district: Arno, Serchio, Ombrone. The
species Cottus gobio, including a variety of moderately cold water adapted species, did used these routes to
reach its current distribution, which encompasses all the afore-mentioned districts (Bianco, 1995).
The Phoxinus species flock found in the Western Balkans probably is the result of a combination of
penetration through underwater connections in the Karstic area (P. karsticus
opposite drainage systems during the orogenesis of the Dinaric mountains (P. apollonicus).
Acknowledgement
We must thank a secret referee for improving the final version of the manuscript.
This article was printed on 15 November 2015 in 30 copies. A copy was sent to each of the following official institutions: Natural
Natural History, Corso Venezia 55, Milan; Regional Museum of Natural Sciences, Via Giolitti, 36, Turin; Civic Museum of Natural
History, Palace Lomellini Carmagnola (To); Civic Museum of Natural History, Via Brigata Liguria 9, Genoa.
be purchased from the bookseller at site Lulu.com
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Researches on Wildlife Conservation, vol. 4, 2015, IGF publ.
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... However, genetic evidence from Palandačić et al. (2015Palandačić et al. ( , 2017 suggested several new species in the Western Balkans. Also, morphological examination of specimens collected in Italy, Croatia, Bosnia and Herzegovina, Montenegro and Albania led to description of Phoxinus ketmaieri Bianco and De Bonis, 2015, Phoxinus likai Bianco and De Bonis, 2015, Phoxinus apollonicus Bianco and De Bonis, 2015, and Phoxinus karsticus Bianco and De Bonis, 2015 Although Croatian and Bosnian karstic area is known for its high diversity and endemicity of freshwater ichthyofauna (Jelić et al., 2016), three of four species described by Bianco and De Bonis (2015) cannot be validated using molecular species delimitation analyses . Two of them (P. ...
... However, genetic evidence from Palandačić et al. (2015Palandačić et al. ( , 2017 suggested several new species in the Western Balkans. Also, morphological examination of specimens collected in Italy, Croatia, Bosnia and Herzegovina, Montenegro and Albania led to description of Phoxinus ketmaieri Bianco and De Bonis, 2015, Phoxinus likai Bianco and De Bonis, 2015, Phoxinus apollonicus Bianco and De Bonis, 2015, and Phoxinus karsticus Bianco and De Bonis, 2015 Although Croatian and Bosnian karstic area is known for its high diversity and endemicity of freshwater ichthyofauna (Jelić et al., 2016), three of four species described by Bianco and De Bonis (2015) cannot be validated using molecular species delimitation analyses . Two of them (P. ...
... ketmaieri and P. likai) were synonymised with P. lumaireul, whereas P. apollonicus was synonymised with P. karsticus . Also, the 8% genetic divergence between P. lumaireul and P. phoxinus on cytochrome c oxidase subunit 1 (COI) supports revalidation of P. lumaireul by Kottelat (2007) and rejects synonymization of P. lumaireul and P. phoxinus made by Bianco and De Bonis (2015). ...
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... Thus, lack of obvious morphological characters and further diversity in the genus Phoxinus revealed by molecular studies [17,18] point to the existence of cryptic lineages in the genus. At present, eleven Phoxinus species are suggested for European drainages (Table 1), with P. phoxinus having a very broad distribution that includes the north-eastern Atlantic, North Sea, Baltic, and Black Sea basins [12,19]. Kottelat [13] and Kottelat & Freyhof [12] mentioned the Danube minnow as a lineage different from P. phoxinus, though they gave no morphological characteristics enabling such discrimination. ...
... Additionally, Knebelsberger et al. [17] found four different lineages populating the area of the P. phoxinus type locality that became obvious to the authors only after molecular analysis. Bianco and De Bonis [19] based three out of four species descriptions on only one or two populations per species, represented by 5-12 specimens, excluding possible variability range of the characters used. Therefore, morphologically defined species were evaluated with molecular data to form secondary species hypotheses. ...
... Finally, the utility of morphological characters for species delimitation in Phoxinus should not be excluded, for example, Ramler et al. [16] suggested the inclusion of all body planes for finding morphologically distinguishing features. However, the problem of hybridisation events in Phoxinus possibly extends to morphology and could be the reason why several studies have not been able to find stable characters for species delimitation [12,13,19]. Thus, in species complexes such as Phoxinus, with closely related species, limited morphological information and numerous hybrid zones, only most modern approaches combined with integrative taxonomy will possibly enable species delimitation. ...
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... In Zrmanja River, P. lumaireul is distributed from the source spring down to Obrovac (Bianco & De Bonis, 2015). However, it remains unclear why it is absent from most of the middle and lower reaches of the Zrmanja River as well as from its main tributary Krupa River. ...
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... In the Palearctic region, the genus is distributed from northern Spain to northern China. Traditionally, morphological traits have been used to identify and describe Phoxinus species (Bianco & De Bonis, 2015;Bogutskaya & Naseka, 2004;Kottelat, 2007). However, morphological analyses alone have poorly resolved species status and relationships within Phoxinus . ...
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... A recent molecular study of P. phoxinus also revealed evidence of a multispecies complex in the western Balkan Peninsula (Palandacic et al., 2015). Morphological analyses of Phoxinus Rafinesque 1820 from Italy and the western Balkans described four additional species (Bianco & De Bonis, 2015), though some incongruencies were found in subsequent molecular analyses (Palandacic et al., 2017). Studies of widely distributed Nearctic leuciscids, including species of Campostoma Agassiz 1855 (Blum et al., 2008;Schönhuth et al., 2011), Cyprinella Girard 1856 (Glotzbecker et al., 2016) and Dionda Girard 1856 (Schönhuth et al., 2012), have uncovered similar evidence of unrecognized diversity, leading to the resurrection of previously described species and the discovery of new lineages that probably warrant species recognition. ...
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An updated, corrected checklist of both native and established alien freshwater fish species in Italy is given based on molecular, morphological and biogeographical data. Some 12 native species, reported as conspecific with transalpine species in official Italian ichthylogical literature, are in fact endemics. Previous taxonomic confusion has resulted in the introduction of several alien species, either with official stockings or mixed in as impurities. Rehabilitated species include the cyprinids Scardinius hesperidicus, Scardinius scardafa and Telestes savigny from northern Italy, as well as Squalius ruffoi and the Telestes comes from southern Italy. Squalius albus is a junior synonym of S. squalus. The endemic gudgeon, previously assigned to the genus Romanogobio, is returned to the genus Gobio (G. benacensis). Phoxinus lumaireul is a junior synonym of P. phoxinus. Among the Salmonidae, Salmo cenerinus is a junior synonym of S. marmoratus, while Salmo farioides represents the trout species of the Adriatic lineage for which a neotype is designated. Thymallus aeliani represents the endemic lineage of grayling of the Adriatic populations. The esocid Esox cisalpinus is an endemic pike species and Esox flaviae is a junior synonym; the extensive exportation as well as the presence of this species throughout Europe is possibly due to humans. Among sculpins, Cottus scaturigo and C. ferrugineus are junior synonyms of C. gobio. The International Union for Conservation of Nature (IUCN) categories for native species of Italy are updated. At present, 52 native freshwater fish species are listed: 2 are extinct (Acipenser sturio and Huso huso), 12 are critically endangered, 7 endangered, 10 vulnerable, 3 near-threatened, 15 low concern and 3 data-deficient; 35 species are the result of human transfers. Among the 51 introduced species, 6 are recently established (Leuciscus leuciscus, Oreochromis niloticus, Poecilia reticulata, Xiphophorus helleri, Amatitlania nigrofasciatus, Hemichromis sp.), 37 are already established, 5 are probably established and 3 are non-established Chinese carp, maintained in the wild by intensive stockings. The family most involved is the Cyprinidae, with 22 alien and 20 native species.
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European bullhead sensus lato is a species complex, in which lineage determination based on morphology has proven to be extremely difficult, probably due to low interspecific and high intraspeciefic morphological variation. Therefore, major evolutionary lineages across Europe have been determined based on molecular data, which also to some extent corresponds to known morphological variation. Cottus populations on the Balkan Peninsula have not yet been thoroughly studied. Thus we used molecular genetic approach (mtDNA-CR and nu-LSU) to evaluate evolutionary distinctive units of Cottus in the Western Balkans and compare them to other European lineages. We found presence of five distinct lineages. Populations corresponding to Cottus gobio clade inhabit tributaries of Danube River in southern Serbia, rivers of Adriatic drainage and Drava River. Interestingly, we found three additional Cottus linages in Sava River and its tributaries, differing considerably from “main” C. gobio. First, we confirmed morphologically determined Cottus meate from Upper Sava, with species range from the source of Sava River, including tributaries, to the inflow of Kolpa River. Second, we found a new lineage inhabiting the Kolpa and Una river systems. Third, lineage was found inhabiting Bosna and Vrbas rivers. High density of evolutionary distinctive lineages in such a small area is according to our knowledge exceptional for European bullhead and lines up with the general notion of the Balkans as major refuge and present day biodiversity hotspot. Our results correspond with two wave colonization theory of Europe by Cottus genus probosed by Volckaert, first possibly being in Pliocene and second during the Pleistocene, with Balkan Peninsula as starting point for both of them.
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