Table Of ContentZOOSYSTEMATICA ROSSICA, 18(1): 126-173 3 JULY 2009
A review of vertebral and fin-ray counts in the genus Alburnoides
(Teleostei: Cyprinidae) with a description of six new species
N.G. BOGUTSKAYA & B.W. COAD
N.G. Bogutskaya, Zoological Institute, Russian Academy of Sciences, Universitetskaya nab. 1, St. Petersburg
199034, Russia. E-mail: [email protected]
B.W. Coad, Canadian Museum of Nature, Ottawa, Ontario K1P 6P4, Canada. E-mail: [email protected]
Variations in the different vertebral patterns and unpaired fin counts in the genus Alburnoides
are discussed based on an examination of over 1100 specimens from a wide area of distribution
of the genus. Though it was commonly considered that the morphological differences between
subspecies and local forms of A. bipunctatus auctorum appear slight, our study shows that
there are pronounced differences between many of them in vertebral and dorsal and anal fin
counts. To estimate reliability of differences, statistical tests were used and a cluster analysis
(UPGMA – Average Linkage clustering method) was performed based on calculated values,
and tree diagrams are presented and analyzed in their taxonomic context. The differences in
fin ray and vertebral counts in combination with some other morphological characters, dis-
tinguished some of the former subspecies or local forms as distinct species. Along with five
nominal taxa re-established to a species level (A. rossicus, A. kubanicus, A. maculatus, A. eich-
waldii, A. fasciatus), we describe six new species: A. gmelini (type locality Sunzha River, Terek
River drainage, Eastern Ciscaucasia, Russia), A. varentsovi (type locality Ashkhabadka River,
northern slope of the Kopetdag, Turkmenistan), A. petrubanarescui (type locality Qasemlou
Chay, Orumiyeh Lake basin, Iran), A. namaki (type locality qanat at Taveh, Namak Lake basin,
Iran), A. nicolausi (type locality Simareh River in Karkheh River system, Tigris River drain-
age, Iran), and A. idignensis (type locality Bid Sorkh River, Gav Masiab River system, Tigris
River drainage, Iran).
Key words: freshwater fishes, Cyprinidae, Alburnini, Alburnoides, new species, morphology,
axial skeleton, unpaired fins
INTRODUCTION Anaecypris Collares-Pereira, 1983 (Gilles
et al., 2001; Cunha et al., 2002; Durand et
Over 30 species, subspecies and local
al., 2002; Freyhof et al., 2005). The latter
forms of the leuciscine cyprinid fishes were
genus was also shown to be morphologi-
considered by Bogutskaya (1990a, 1990b,
cally close to the Alburnini (Bogutskaya
1997b). They represent a monophyletic & Collares-Pereira, 1997).
taxon, the tribe Alburnini, characterized The uncertainty of the generic defini-
by a set of derived characters, some of them tions of Alburnus, Chalcalburnus and Al-
unique, in external morphology, sensory burnoides was discussed earlier (Bianco,
canals and skeleton. Alburnus Rafinesque, 1980; Krupp, 1985) and an opinion was
1820, Chalcalburnus Berg, 1932, Alburn- proposed that they may be synonyms
oides Jeitteles, 1861, Tropidophoxinellus (Coad, 1991). Later, Alburnus and Chal-
Stephanidis, 1974 and Leucaspius Heckel calburnus were synonymized by Boguts-
& Kner, 1858 were included in the tribe. kaya (1997a) since the distinguishing
Later, a number of molecular studies re- characters of Chalcalburnus according
vealed that the group does form a clade to Berg (1932b, 1949) – a partly scaled
and includes, if representatives geneti- ventral keel, numerous long gill rakers
cally examined are considered, at least and slightly serrated or unserrated pha-
Alburnus, Chalcalburnus, Leucaspius and ryngeal teeth – have no diagnostic value
© 2009 Zoological Institute, Russian Academy of Scienсes
N.G. BOGUTSKAYA & B.W. COAD. A REVIEW OF ALBURNOIDES 127
being variably present in most Alburnus, considered a complex species with a number
Alburnoides and Chalcalburnus species. On of subspecies found from France through
the other hand, according to Berg (1949), Europe north of the Alps eastwards to the
the only difference between Alburnus and Black, Caspian and Aral Sea basins (e.g.
Alburnoides is unserrated pharyngeal Berg, 1949; Bogutskaya & Naseka, 2004;
teeth. Synonymization of Alburnus and Coad, 2009). Alburnoides ohridanus (Kara-
Chalcalburnus was supported by molecu- man, 1928) and A. prespensis (Karaman,
lar studies (e.g. Durand et al., 2002) and 1924) were recently given a rank of species
is widely accepted (e.g. Bogutskaya & (Kottelat & Freyhof 2007) as well as A.
Naseka, 2004; Kottelat & Freyhof, 2007). eichwaldii (De Filippi, 1863) (Fricke et al.,
In contrast, Alburnoides stands apart from 2007), and a new species from the Pulvar R.
Alburnus in molecular trees (Hänfling & system in southern Iran will be described in
Brandl, 2000; Gilles et al., 2001; Cunha et a paper by Coad & Bogutskaya (2009).
al., 2002; Freyhof et al., 2005). However, The following nominal taxa, which have
morphological definitions of the two gen- appeared in the literature as subspecies or
era appear to be quite obscure. They both distinct forms, have been historically synon-
include species which are characterized ymized with A. bipunctatus: Alburnoides bi-
by a more or less pronounced ventral keel punctatus armeniensis Dadikyan, 1972 (type
(from scaled to completely scaleless), a locality: rivers Arpa, Vorotan, Vedi, Marma-
slightly to considerably elongated anal fin, rik, Kasakh with tributaries of the Aras R.
pharyngeal teeth commonly 2.5-5.2 or 2.5- system in the Kura R. drainage, Caspian
4.2, an absolutely and relatively elongated Sea basin, Armenia), Alburnus eichwaldii De
predorsal vertebral subregion (to 15-17 Filippi, 1863 (Kura R. at Tiflis, Caspian Sea
vertebrae which is commonly over 38% of basin, now Georgia), Aspius fasciatus Nor-
total and 70% of abdominal vs. 33-37% and dmann, 1840 (rivers of the western coast of
60-66%, respectively, in most other groups the Black Sea eastward to Mingrelia, now
of the Leuciscinae), a tendency to equal- Russia and Georgia), Alburnoides bipunc-
ity of the numbers of vertebrae in abdomi- tatus rossicus natio kubanicus Berg, 1932
nal and caudal regions (modal difference (unavailable name; Kuban’ R., Sea of Azov
between abdominal and caudal numbers basin, Russia), Alburnus maculatus Kessler,
decreases from 4 or 3 to 1, 0, and [-1] vs. 1859 (Salgir R., Sea of Azov basin, Crimea
modes 4 to 6 in the most Leuciscinae), and Peninsula, now Ukraine), Alburnus bipunc-
a large orbit with respective configuration tatus ohridanus Karaman, 1928 (Ohrid L.,
of all the cranial elements it is formed from Adriatic Sea basin, now the Former Yugo-
(Bogutskaya, 1990a, 1990b; Bogutskaya slav Republic of (FYRO) Macedonia), Al-
et al., 2000). Actually, the only character burnus bipunctatus var. prespensis Karaman,
easy to use for distinguishing Alburnus and 1924 (Prespa L. and its tributaries, no di-
Alburnoides is the pattern of pigmenta- rect link to any basin, now FYRO Macedo-
tion. In Alburnoides, small black spots are nia), Alburnoides bipunctatus rossicus Berg,
located on each side of the lateral line ca- 1924 (Dnieper, Black Sea basin, and Volga,
nal pore outlining the canal at least along Caspian Sea basin, now Ukraine and Rus-
its anterior portion, and a dark stripe goes sia), Alburnoides bipunctatus var. smyrnae
from behind the eye to the caudal fin base Pellegrin, 1927 (Mélèl stream near Smyrna,
though both characters may be variably Aegean Sea basin, Turkey), Alburnoides
developed in live fish or are absent. bipunctatus strymonicus Chichkoff, 1940
Besides Alburnoides oblongus Bulgakov, (Struma R. drainage, Bulgaria), Alburnoi-
1923 and Alburnoides taeniatus (Kessler, des bipunctatus tzanevi Chichkoff, 1933
1874) from the Aral Sea basin, Alburnoides (Rezova R., Black Sea basin, Bulgaria and
bipunctatus (Bloch, 1782) has long been Turkey), and Alburnoides bipunctatus subsp.
© 2009 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 18(1): 126-173
128 N.G. BOGUTSKAYA & B.W. COAD. A REVIEW OF ALBURNOIDES
(Berg, 1932a, 1932b) (Kuma, Terek, Sulak with regard to the dispersion, we calculated
rivers, Caspian Sea basin, Russia). Kul’bak’s Divergence that has a meaning
The most comprehensive review of A. bi- of average information measure of the dif-
punctatus s.l. is still that by Berg (1949) who ference between two empiric distributions
successfully used the number of pharyngeal (Kulbak, 1967) and a cluster analysis (UP-
teeth and the number of branched anal-fin GMA – Average Linkage clustering meth-
rays for distinguishing six subspecies and od), were performed based on calculated
infrasubspecies forms (one unnamed) with- values. Similarity Indices (r) (Zhivotovs-
in the species. Thought it was later con- kiy, 1991) were calculated for each charac-
sidered that the morphological differences ter separately, for five vertebral characters
between subspecies and local forms of A. and for all seven characters under consid-
bipunctatus auctorum appear slight (Kotte- eration, and tree diagrams (UPGMA – Av-
lat & Freyhof, 2007), our study shows that erage Linkage clustering method) are pre-
there are pronounced differences between sented and analyzed. Abbreviations used: n,
many of them in vertebral and dorsal and number of specimens; min, minimum value;
anal fin counts, and these differences be- max, maximum value; avg, average (mean);
tween the different vertebral patterns are m, standard error of the mean; std, standard
discussed below. In combination with some deviation; Me – median; Mo – mode.
other morphological characters this gave Below we provide two lateral line scale
reason to distinguish some of them as dis- counts, the total lateral line scale count
tinct species. Along with five nominal taxa which includes all pierced scales, from the
resurrected to a species level, we describe first one just behind the supracleithrum to
six new species from the eastern part of the the very posteriormost one, and the later-
area of distribution of the genus. al line scale count which includes pierced
scales from the first one just behind the su-
MATERIAL AND METHODS pracleithrum caudad to the scale (inclusive)
at the base of the caudal fin rays (i.e. poste-
Counts and measurements follow Hubbs rior margin of hypurals); the second count
& Lagler (1958). Measurements are to thus excludes 1, 2 or 3 scales located on the
the nearest 0.1 mm. Head length and in- bases of the caudal fin rays. Osteological
terorbital width were measured to their characters are examined in cleared-and-
bony margins. Fin ray counts separate un- stained specimens and from radiographs of
branched and branched rays. The last two 918 specimens from over 1100 listed below.
branched rays articulated on a single ptery- We examined representatives of Alburn-
giophore in dorsal and anal fins are noted as oides s.l. from most of its range; however,
“1½”. All statistical calculations were done we did not discuss western Anatolian and
without “½”. For morphometric data, t-tests Aegean (Greek and Bulgarian) forms for
were used to compare males and females for we have not had enough material for com-
morphometric characters after testing for parisons of, e.g. A. bipunctatus var. smyrnae,
heteroscedascity; a non-parametric Mann- A. bipunctatus strymonicus and A. bipuncta-
Whitney U-test was used if t-tests were not tus tzanevi. Vertebrae counts are given ac-
applicable. For fin ray and vertebrae counts, cording to Naseka (1996) as discussed for
we calculated the most common statistical the tribe Alburninae in Bogutskaya et al.
values and criteria. To estimate reliability of (2000). Cephalic sensory canal terminol-
differences between means (averages), Stu- ogy follows Illick (1956) and is discussed
dent’s t-tests were performed and the com- in Bogutskaya (1991): CIO, infraorbital ca-
puted t-scores are presented. Besides, for nal; CPM, preopercular-mandibular canal;
getting additional information on degree CSO, supraorbital canal; CST, supratem-
of similarity of means between the samples poral canal. Other abbreviations used: SL,
© 2009 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 18(1): 126-173
N.G. BOGUTSKAYA & B.W. COAD. A REVIEW OF ALBURNOIDES 129
standard length; HL, head length; institu- 80 specimens from Crimea, Movchan &
tions and collections: BMNH, Natural His- Smirnov (1983) found no single specimen
tory Museum, London; CMNFI, Canadian with 5 teeth in the main row on the right
Museum of Nature, Ottawa; DUM, Zoolog- 5th ceratobranchial among 80 examined
ical Museum of Sciences and Art Faculty, specimens. According to Berg (1949), A.
Dicle University, Diyarbakir; MKC, pri- fasciatus invariably possess 2.5-4.2. In 27
vate collection of Maurice Kottelat; MBL, specimens of A. fasciatus examined by us
Museu Bocage, Lisbon; MRSNT, Museo we found 2.5-4.2 (24), 2.4-4.2 (1) and 2.5-
Regionale di Scienze Naturali, Zoologia, 4.1 (2). Variants with 5 teeth in the lon-
Torino; NMW, Naturhistorisches Museum, ger row on the right ceratobranchial are
Wien; SMF, Senckenberg Museum, Frank- more frequent in A. eichwaldii, and Berg
furt a. Main; VPC, collection of V. Poznyak (1949) considered this to be the main di-
in Elista State University; ZISP, Zoological agnostic feature of A. bipunctatus eichwal-
Institute, Russian Academy of Sciences, St. dii in his understanding. In a sample from
Petersburg; ZMH, Zoologisches Museum Chaldyr Lake (Kura drainage), we found
und Institut, Universität Hamburg. C&S 2.5-5.2 teeth in 5 specimens along with
indicates cleared and stained specimens. 25 fishes possessing 2.5-4.2. As shown in
the descriptions below, in all species of the
RESULTS AND DISCUSSION former A. bipunctatus eichwaldii complex
the formula 2.5-4.2 is the most common. It
Pharyngeal teeth. Teeth are hooked at is worth mentioning that the formula 2.5-
tip, with a grinding area below lacking evi- 5.2, which should be considered a charac-
dent striations. We found no differences in ter state close to a condition plesiomor-
the shape of the teeth that could serve for phic for the Leuciscinae as being widely
species diagnostics. The number of pharyn- presented in Leuciscus and other less spe-
geal teeth was traditionally used for dis- cialized genera, is present among Alburn-
tinguishing subspecies among A. bipunc- oides in species that should be considered
tatus s.l. The groups of species within the the most derived if anal fin and vertebral
former A. bipunctatus can be seen in Tables counts are concerned (see below).
1-3; the nominotypical subspecies, or “typ- Branched dorsal-fin rays. The number
ical” A. bipunctatus, is almost exclusively of branched dorsal-fin rays is commonly
characterized by 2.5-4.2 teeth along with 8½. However, in A. eichwaldii (Kura-Aras
A. ohridanus (e.g. Berg, 1949). Alburnoides & Lenkoran) 7½ are rarely present (found
rossicus and A. kubanicus (forming to- in 6% of specimens) while in A. idignen-
gether A. bipunctatus rossicus auctorum) sis it was found in 29% of specimens. In
are different by commonly having 2.5- A. petrubanarescui sp. n. and especially A.
5.2 and other variants with 5 teeth in the nicolausi sp. n. the number of 7½ clearly
main row on the right 5th ceratobranchi- predominates, 67% and 91%, respectively
al. However, Movchan & Smirnov (1983) (Table 1).
found that the formula 2.5-4.2 is also of- Branched anal-fin rays. The number
ten met – they found 4 teeth in the main of branched anal-fin rays is widely dis-
row on the right 5th ceratobranchial in cussed below (see also Tables 1 and 4) for
30% of 108 specimens examined by them. it has been long considered as one of the
In 15 specimens of A. kubanicus examined main diagnostic characters of Alburnoides
by us we found only 2.5-5.2. Alburnoides species. The lowest counts are discovered
maculatus represents the north-western in two Iranian species – A. petrubanares-
form of the former A. bipunctatus fascia- cui sp. n. from the Urmia basin (a mode is
tus s.l. which is characterized by the pre- 9½) and A. nicolausi sp. n. from the Tigris
dominance of the 2.5-4.2 formula. Among drainage (a mode is 10½). Increase of the
© 2009 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 18(1): 126-173
130 N.G. BOGUTSKAYA & B.W. COAD. A REVIEW OF ALBURNOIDES
17 4
16 3 11 5 5
15 12 5 33 4 13 1
14 21 1 5 5 26 29 12 8 4
).
ed ys
s shad al finhed ra 13 21 2 2 16 20 6 45 18 18 4
count Anbranc 12 5 15 15 53 4 62 4 14 24 16 5 18 1
al
d
mo 11 2 7 9 38 5 17 9 23 22 30 4
al
m
maxi 10 1 3 1 11 3 23 2 3 5 1
d
n
a
al 9 16 8 1
m
ni
mi
( 8 3 2
s
y
a
r
d
he 9 2 2 1 4
c
n
a
d anal fin br Dorsal finanched rays 78 591 18 21 45 24 83 54 53 14810 301 411 2010 462 384 3510 263 49 6
an br
al
s
or 6 1
d
of
r
e
b
m
u
n
e
able 1. Frequency of occurrence of variants of th Western group, pharyngeal teeth 2.5-4.2 A. bipunctatus (Danube), n=62 A. ohridanus, n=18 North-central group, pharyngeal teeth 2.5-5.2 A. rossicus, n=21 A. kubanicus, n=45 South-eastern group, pharyngeal teeth 2.5-4.2 A. maculatus: Chiornaya R., n=24 A. maculatus: Salgir, Alma, Kacha rivers, n=83 A. fasciatus, n=54 A. gmelini sp. n., n=53 A. eichwaldii: Kura-Aras & Lenkoran, n=160 A. cf. eichwaldii: Safid Rud, n=31 A. varentsovi sp. n., n=42 A. petrubanarescui sp. n., n=30 A. namaki sp. n., n=48 A. nicolausi sp. n., n=42 A. idignensis sp. n., n=46 Alburnoides sp. Pulvar R., n=30 A. taeniatus, n=53 A. oblongus, n=6
T
© 2009 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 18(1): 126-173
N.G. BOGUTSKAYA & B.W. COAD. A REVIEW OF ALBURNOIDES 131
Fig. 1. Tree diagram (UPGMA) computed from means of the number of branched anal-fin rays. See
also Tables 1, 4 and 7.
branched anal-fin rays number is observed Cephalic lateral line canals. The gen-
in all groups delimited by the number of eral topography of cephalic sensory canals
pharyngeal teeth – in the western 2.5-4.2 and numbers of pores is rather similar in all
group (in A. bipunctatus from the Danube examined Alburnoides species and is close in
with the modes of 12½ and 13½), in the the main features to the typical pattern de-
north-central 2.5-5.2 group (modes 15½ scribed by Bogutskaya (1988). The supraor-
and 16½ in, respectively, A. rossicus and A. bital canal is not lengthened in its posterior
kubanicus), and in the south-eastern 2.5- section and has 7-11, commonly 8-10 pores,
4.2 group (a mode of 14½ in A. gmelini sp. with 2-4 and 5-7 canal openings on the nasal
n. and A. fasciatus). As seen in the Table 7, and frontal bones, respectively. The infraor-
statistically reliable differences are found bital canal has 10-16 pores (commonly 12-
in the number of branched anal-fin rays 14) with 4 (commonly) or 5 canal openings
between most taxa under consideration. on the first infraorbital. The preopercular-
Thus, total counts differ significantly be- mandibular canal is complete, with 11-17,
tween geographically close A. rossicus and modally 13-16, pores with (3)4-6 and 7-10
A. kubanicus, between true A. eichwaldii canal openings on the dentary and preoper-
(Kura & Lenkoran’) and A. cf. eichwaldii culum, respectively. The supratemporal ca-
from Safid River, between true A. eichwal- nal is complete, with commonly 5-7 pores.
dii and A. namaki, etc. Tree diagrams which We do not discuss below the counts of sen-
represent the anal-fin rays data clustering sory pores and the canal pattern for they are
are given in Figs 1 and 2. not diagnostic for the species. However, it
© 2009 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 18(1): 126-173
132 N.G. BOGUTSKAYA & B.W. COAD. A REVIEW OF ALBURNOIDES
Fig. 2. Tree diagram (UPGMA) computed from similarity indices of the number of branched anal-
fin rays.
may be useful for further comparisons with erably varies in terms of both absolute and
Alburnus and other genera. relative values. For example, A. bipuncta-
Vertebrae. As it is seen from the Table tus from the Danube and A. taeniatus pos-
1, there is some correlation between the sess equal means of this counts (13.6) while
branched anal-fin rays count and the num- they have 41.2 and 37.9 mean total verte-
ber of total vertebrae: the highest values of brae indicating that the relative length of
total vertebrae are found in A. rossicus and the predorsal vertebral region comprises
A. kubanicus which are characterized by the 33% and 36% of the total vertebral column,
highest number of anal-fin rays (commonly respectively. The lowest predorsal count
41-43). The lowest number is in A. taeniatus (mean 12.2) is found in A. varentsovi sp. n.,
(a mode of 38). As it is seen from the Table 8, A. namaki sp. n., and A. idignensis sp. n. Dif-
statistically reliable differences are found in ferences in the predorsal count give reasons
the number of total vertebrae between most for distinguishing some samples as distinct
taxa under consideration. Thus, total counts taxa. For example, true A. eichwaldii from
differ significantly between geographically the Kura drainage and Lenkoran’ Province
close A. rossicus and A. kubanicus, between significantly differs from a sample from Sa-
true A. eichwaldii (Kura & Lenkoran’) and fid Rud (fish from this river is commonly
A. cf. eichwaldii from Safid Rud, between identified as A. bipunctatus eichwaldii) (13.7
true A. eichwaldii and A. namaki sp.np, etc. vs. 12.5, Tables 5 and 9).
It is worth mentioning that the number of We paid special attention not only to ab-
predorsal vertebrae that reflects the length solute values of vertebrae in the abdominal
of the predorsal distance in general consid- and caudal regions (for differences between
© 2009 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 18(1): 126-173
N.G. BOGUTSKAYA & B.W. COAD. A REVIEW OF ALBURNOIDES 133
16 2
15 3 3 4 10 10 4 1
e
a
ebr 14 34 2 17 27 10 20 30 40 86 1 12 1 1 6 25 5
t
r
e
v
al
ors 13 25 13 1 18 14 60 20 1 62 15 7 18 10 24 9 24 24
d
e
r
P
12 3 3 2 16 32 35 18 34
).
d
ade 11 2 2 2
h
s
s
ount 43 1 10 6 9
c
al
d
mo 42 16 11 18 4 9 12 6 52 1 1 1
al
m
xi
ma 41 41 1 18 14 50 30 20 73 9 3 14 3 14 3
d
n
a
al
m e
mini ebra 40 4 2 3 6 24 10 26 23 21 28 14 27 5 8 16 2
( rt
rtebrae Total ve 39 13 2 1 2 1 10 1 18 28 30 12
e
v
al
ors 38 2 1 9 7 24
d
e
r
p
d
an 37 1 16
al
ot
r of t 36 1
e
b
m
u
n
able 2. Frequency of occurrence of variants of the Western group, pharyngeal teeth 2.5-4.2 A. bipunctatus (Danube), n=62 A. ohridanus, n=18 North-central group, pharyngeal teeth 2.5-5.2 A. rossicus, n=21 A. kubanicus, n=45 South-eastern group, pharyngeal teeth 2.5-4.2 A. maculatus: Chiornaya R., n=24 A. maculatus: Salgir, Alma, Kacha rivers, n=83 A. fasciatus, n=54 A. gmelini sp. n., n=53 A. eichwaldii: Kura-Aras & Lenkoran, n=160 A. cf. eichwaldii: Safid Rud, n=31 A. varentsovi sp. n., n=42 A. petrubanarescui sp. n., n=30 A. namaki sp. n., n=48 A. nicolausi sp. n., n=42 A. idignensis sp. n., n=46 Alburnoides sp. Pulvar R., n=30 A. taeniatus, n=53 A. oblongus, n=6
T
© 2009 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 18(1): 126-173
134 N.G. BOGUTSKAYA & B.W. COAD. A REVIEW OF ALBURNOIDES
Fig. 3. Tree diagram (UPGMA) computed from similarity indices of the difference between the
numbers of abdominal and caudal vertebrae.
samples see Tables 10 and 11) but also to punctatus (Danube), and A. cf. eichwaldii
their relative length. In most leuciscine from Safid Rud (Tables 3, 6 and 12). Data
cyprinids the abdominal region is 3-4 ver- clustering based on similarity indices (r)
tebrae longer than the caudal region and it displays 6 major clusters which reflect the
may be considered as a probable primitive main types of vertebral pattern in terms of
pattern while in the Alburnini (Bogutskaya relative length of the abdominal and caudal
et al., 2000) the caudal region is relatively region (Fig. 3, Table 3).
elongate and is often longer than the ab- Data clustering which summarize all
dominal region. A comparatively primitive five vertebral characters, both by means
character state (difference between the ab- and by frequency arrays, are presented in
dominal and caudal count is positive, from Figs 4 and 5. The tree diagrams given in
+3 to +1, or the counts are equal), is found these figures are rather similar for the main
in the Orumiyeh [Urmia] Lake basin (A. clusters are identical on the both diagrams.
petrubanarescui sp. n.) and in the Crimea However, the second diagram seems to be
Peninsula (A. maculatus) (Tables 3 and 6). a more appropriate one since it represents
A relatively long abdominal region is also frequency arrays – frequency of distinct
characteristic for A. oblongus, A. ohridanus, phenotype occurrence – rather than mean
A. taeniatus, and A. nicolausi sp. n. The most values. It is evident that the most main clus-
specialized pattern (difference between the ters are formed from geographically distant
abdominal and caudal count is negative, species. This may be explained by, first, the
from -1 to -3) is found in A. rossicus, A. bi- mosaic distribution of the primitive pattern
© 2009 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 18(1): 126-173
N.G. BOGUTSKAYA & B.W. COAD. A REVIEW OF ALBURNOIDES 135
d
n
al a -3 3 1
m
mini al -2 15 8 6 2 5 11 7 3 1
r ( min
mbe bdobers -1 34 1 9 18 12 11 33 8 13 9 14 12 5
u am
ominal and caudal n Difference between and caudal nu +2+10 55 1322 3 615 8122 482310 2020 30210 62949 9911 225 12133 2228 1877 1714 162 221114 31
d
b
a
the +3 2 2 2 2 1 1 2
n
e
e
tw 23 1 1
e
b
e
c
en 22 15 17 12 2 9
r
e
he diff brae 21 39 3 27 2 5 22 16 76 20 10 1 6 12
brae, anal t audal verte 20 7 4 6 14 58 28 35 66 9 28 14 32 13 23 18 4 2
rte C 19 12 8 20 2 2 9 2 4 15 10 22 22 23 4
e
v
udal 18 2 7 1 24
a
c
nd 17 2
a
al
min 22 4 7 8 2 3
o e
mber of abd nal vertebra 2021 508 161 129 2124 20 715 2826 3418 10346 191 272 253 332 282 24 282 19 3
nu mi
of the Abdo 19 4 1 1 2 11 13 13 12 21 33
ts 18 1 1 1
n
a
ri
a
Table 3. Frequency of occurrence of vmaximal modal counts shaded). Western group, 2.5-4.2 A. bipunctatus (Danube), n=62 A. ohridanus, n=18 North-central group, 2.5-5.2 A. rossicus, n=21 A. kubanicus, n=45 South-eastern group, 2.5-4.2 A. maculatus: Chiornaya R., n=24 A. maculatus: Salgir, Alma,Kacha rivers, n=83 A. fasciatus, n=54 A. gmelini sp. n., n=53 A. eichwaldii: Kura-Aras &Lenkoran, n=160 A. cf. eichwaldii: Safid Rud, n=31 A. varentsovi sp. n., n=42 A. petrubanarescui sp. n., n=30 A. namaki sp. n., n=48 A. nicolausi sp. n., n=42 A. idignensis sp. n., n=46 Alburnoides sp. Pulvar R., n=30 A. taeniatus, n=53 A. oblongus, n=6
© 2009 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 18(1): 126-173
