Iran. J. Ichthyol.

(September 2015), 2(3): 155–164 Received: June 10, 2015

© 2015 Iranian Society of Ichthyology Accepted: August 27, 2015
P-ISSN: 2383-1561; E-ISSN: 2383-0964 doi:
http://www.ijichthyol.org

Osteological characteristics of Paraschistura nielseni (Nalbant & Bianco,

1998) (Cypriniformes: Nemacheilidae)
Hoda AZIMI1, Hamed MOUSAVI-SABET1*, Soheil EAGDERI2
1
Department of Fisheries, Faculty of Natural Resources, University of Guilan, Sowmeh Sara, Iran.
2
Department of Fisheries, Faculty of Natural Resources, University of Tehran, Karaj, Iran.
*Email: mousavi-sabet@guilan.ac.ir

Abstract: Paraschistura Prokofiev, 2009 is a newly described genus based on

osteological characters, but detailed information for all species of the genus is not
available. To describe osteological characteristics of the endemic species of
Paraschistura nielseni, twelve specimens were collected from the Shapur River of
Persis basin which drains to the Persian Gulf and their osteological characteristics
were examined. According to the results, P. nielseni is characterised by bearing a
square-shaped prevomer, four basibranchials, a semi-ossified sesamoid, a non-
alveolar bony swim bladder capsule, and separated pelvic bone bases. Based on these
features, P. nielseni could be distinguished from other loaches species.
Keywords: Osteology, Fish skeleton, Neurocranium, Loach, Persis basin.

Introduction ascribed to it or related genera (Coad 2015). Recently

The members of the family Nemacheilidae, with
about 72 genera and 651 species, are found across
Eurasia with one species in northeast Africa (Nelson
2006; Eschmeyer & Fong 2015; Mafakheri et al.
2015). This family has a great diversity in Iranian
interior waters (Coad 2015). Classification of these
taxa are complex and researches are trying to
determine their phylogenic status (Prokofiev 2010;
Mafakheri et al. 2015). Recent classifications of
members of the family Nemacheilidae have been
merely based on external morphology and to a small
extent on anatomy, osteology and molecular data
(Bănărescu & Nalbant 1995; Freyhof et al. 2015).
The first osteological work on this group of
fishes was performed by Regan (1911), who
separated the subfamily Nemacheilinae from
Cobitidae. The only comprehensive phylogenetic
study based on the osteology of nemacheilid fishes
was carried out by Prokofiev (2010). Among the
members of nemacheilids, Paraschistura Prokofiev,
2009 is a newly described genus, and therefore, not
all of its species have been fully examined and
155
Vatandoust and Eagderi (2015) described
P. ilamensis from Tigris River drainage as first
species of the genus of Paraschistura from Iranian
part of this basin. Freyhof et al. (2015) reviewed the
genus Paraschistura from Iran and described six new
species, including P. abdolii (from the Sirjan basin
and the western tributaries of the Hamun-e Jaz
Murian basin), P. aredvii (from the Zohreh drainage),
P. hormuzensis (from the Minab drainage),
P. naumanni (from the Kol and Mond drainages and
Lake Maharlo basin), P. pasatigris (from the Karun
and Karkheh drainages) and P. susiani (from the
Jarahi drainage) based on the morphological and
molecular (the mtDNA COI barcode region) data set.
Freyhof et al. (2015) provided the diagnostic
characters for all eleven recognized species including
Paraschistura nielseni and treated Metaschistura
Prokofiev, 2009 as a synonym of Paraschistura
Prokofiev, 2009.
Many members of this genus were previously
described in the genus Schistura (McClelland 1838).
Species of the genus Paraschistura are small and
 Iranian Journal of Ichthyology (September 2015), 2(3): 155-164
Fig.1. Paraschistura nielseni, 55.7 mm SL, collected from Shapur River, in the Persis basin, southern Iran.
found in inland waters of Turkmenistan, east of
Baluchistan in Iran, and the Indus River in Pakistan
and Afghanistan (Coad 2015; Freyhof et al. 2015).
Paraschistura nielseni (Nalbant & Bianco 1998) (Fig.
1) is endemic to Iranian inland waters commonly
called sagmahi-ye Nielseni (Coad 2015). In 1966,
this species was described by Bărănescu and Nalbant
as Nemacheilus bampurensis (Coad 2015; Freyhof et
al. 2015). Paraschistura nielseni is known from the
Helleh and Mond River drainages (Persis basin)
which drain to the northern Persian Gulf (Freyhof et
al. 2015). Due to difficulties in using external
morphology of the members of this genus for
studying their taxonomy and limited information on
these species, the present study was conducted to
provide a detailed description of osteological
characteristic of P. nielseni. Since, osteological
features are important in the taxonomy of the family
Nemacheilidae; the results of this study can be used
as a reference to more accurately compare and
distinguish different species of the genus.
Materials and Methods
Twelve specimens of P. nielseni (38-65 mm in
standard length) were collected by electrofishing
from the Shapur River (Helleh drainage, Persis basin,
Fars Province, Iran) (51°33'06"E, 29°45'44"N). The
collected specimens were anesthetized in 1% clove
solution and then fixed in 5% buffered formalin. For
osteological examination, specimens were cleared
156
and stained using alcian blue and alizarin red based
on Taylor & Van Dyke (1985). The skeletal
structures were dissected and photographed using a
scanner (Epson v600) equipped with a glycerol bath.
The skeletal structure of cleared and stained
specimens were observed and studied by an
MS5Leica stereomicroscope. The skeletal elements
were drawn based on digital pictures using
CorelDrawX6 software. Nomenclature and
abbreviation of the skeletal elements were based on
Prokofiev (2009).
Results
Neurocranium: The anterior part of the neurocranium
is narrower and its posterior part is approximately
oval-shaped. The maximum width of the skull is
formed at the level of the pterotic. The ethmoid
region comprises of the paired lateral ethmoid and,
unpaired prevomer and supraethmoid-ethmoid (Fig.
2a). The supraethmoid-ethmoid is a stretched bone
that is vertically fused to the prevomer and firmly
connected to the frontal by a zigzag gap posteriorly.
In the anterior part of the neurocranium, the paired L-
shaped lateral ethmoids are present. The lateral
ethmoid possesses a well-developed anterior process
and also possesses a rod-like processes posteriorly.
The prevomer is almost square-shaped and connected
to the orbitosphenoid and parasphenoid posteriorly
(Fig. 2b).
Several small and free bones are connected to the
 Azimi et al.-Osteological characteristics of Paraschistura nielseni

of Cobitis avicennae

Fig.2. Neurocranium of Paraschistura nielseni (from the dorsal (a), lateral (b), and ventral (c) sides): pr-Bo: basioccipital
process; Bo: basioccipital; Epo: epiotic; Exo: exoccipital ; fon: fontanelle; Fr: frontal; fr-Exo: foramen exoccipital; Let:
lateral ethmoid; Orb: orbitosphenoid; Pa: parietal; Pe: prevomer; Pro: prootic; Ps: parasphenoid; Pto: pterotic; Pts:
pterosphenoid; Se: supraethmoid-ethmoid; Soc: supraoccipital; Spo: sphenotic.
ethmoid part of the neurocranium, including the connecting to the prepalatine laterally (Fig. 3a). Also,
unpaired kinethmoid, paired preethmoids-II and this bone is connected to the anterior edge of the
prepalatines (Fig. 3). The preethmoid-II is rod-like prevomer posteriorly and to the maxilla anteriorly.

157
 Iranian Journal of Ichthyology (September 2015), 2(3): 155-164
Fig.3. The lateral view of the preethmoid-II,
kinethmoid and prepalatine bones in Paraschistura
nielseni. Peth-II: preethmoid-II (a); Ke: kinethmoid
(b); Ppl: prepalatine (c).
The kinethmoid is a small and free bone that it is
positioned between the two maxilla bones (Fig. 3b).
The prepalatine is a small bone connecting to the
preethmoid-II dorsally and maxilla anteriorly (Fig.
3c). The paired sesamoids are semi-ossified in the
ethmoid region.
The orbital region is composed of the frontal,
orbitosphenoid, pterosphenoid, parasphenoid, and
sclerotic bones. The paired frontals are the largest
bones of the skull roof and connected to the
orbitosphenoid, pterosphenoid and sphenotic
laterally, and parietal posteriorly (Figs. 2a‒2b).
These bones include about half of the length of the
neurocranium which are separated by the fontanel
posteriorly. The orbitosphenoid is connected to the
parasphenoid ventrally and pterosphenoid postero-
dorsally (Fig. 2b). The pterosphenoid is connected to
the frontal dorsally and sphenoid postero-laterally.
The posterior margin of the pterosphenoid is curved,
creating a cavity with the prootic and parasphenoid
(Fig. 2c). The parasphenoid is the longest bone in the
base of the neurocranium, and extended from the
prevomer to the basioccipital. This bone is wider in
the middle part and bifurcated at the two ends (Fig.
2c).
The otic region comprises the parietal,
sphenotic, pterotic, prootic and epiotic (Fig. 2). The
posterior margin of the parietal is connected to the
supraoccipital and epiotic, and its lateral margin is
connected to the pterotic and sphenotic. Also, these
paired bones are anteriorly connected to the frontal
158
Fig.4. The lateral view of the upper (a) and lower (b)
jaws in Paraschistura nielseni. Art: articular; Cm:
coronomeckelian; Den: dental; Mx: Maxilla; Pmx:
Premaxilla; Rar: retroarticular.
and separated from each other by the fontanel (Fig.
2a). The pterotic is quarter-circle in shape and
connected to the epiotic and sphenotic posteriorly
and to the prootic and exoccipital ventrally. The
sphenotic forms part of the lateral wall of the skull
and connected to the pterotic ventrally and to the
parietal postero-dorsally (Fig. 2b). The prootic is the
largest bone of the skull base. Its anterior part
contacts the parasphenoid, its upper edge contacts the
sphenotic and its posterior part contacts the pterotic,
exoccipital and basioccipital (Fig. 2c). There is a
foramen in the antero-lateral part of the prootic (Fig.
2c). The epiotic is the posterior most element of the
otic region, and positioned between the occipital
region and pterotic (Fig. 2c).
The occipital region comprises of the
exoccipital, supraoccipital, and basioccipital. The
supraoccipital is pentagon-shaped and its anterior
margin is contributed in the formation of the fontanel.
In addition, this bone is connected to the exoccipital
dorsally and fontanel anteriorly. The exoccipital
bones bear exoccipitalis foramen. The basioccipital
is positioned between the two exoccipitals and
connected to the prootic anteriorly (Fig. 2c). Also,
this bone has a ring-like process posteriorly (process
basioccipital) (Fig. 2b). The neurocranium has two
facets for articulation with the heads of the
hyomandibular. The anterior facet is formed by the
 Azimi et al.-Osteological characteristics of Paraschistura nielseni
of Cobitis avicennae
Fig.5. Lateral view of suspensorium in Paraschistura
nielseni. Apl: autopalatine; Ect: ectopterygoid; End:
endopterygoid; Hm: hyomandibular; Io: interopercle;
Mtp: metapterygoid; Op: opercle; Po: preopercle; Q:
quadrate; So: subopercle; Sym: symplectic.
pterosphenoid, sphenotic and prootic, and the
posterior one by the sphenotic and pterotic. The
parietal, frontal, and supraoccipital are contributed in
the formation fontanel that is covered by a connective
tissue and extended longitudinally (Fig. 2a).
Jaws: The upper jaw comprises the maxilla and
praemaxilla (Fig. 4a). The premaxilla is a narrow L-
shaped bone and composed of two parts, i.e.
ascending and alveolar premaxilla processes. The
horizontal part is arc-shaped and vertical part is
narrower and longer (Fig. 4a). The maxilla is a large
laminar bone and slightly twisted along its
longitudinal axis. The lower jaw is composed of the
dental, retroarticular, articular and coronomeckelian
(Fig. 4b). The dental is the largest of this set and
includes two parts i.e. the narrow ramus dentalis and
wider coronoid process. This bone is connected to the
articular postero-dorsally and to the retroarticular
dorsally. The articular is connected to the dental
anteriorly, retroarticular ventrally and quadrate
posteriorly. The coronomeckelian is a small and
triangular bone which is positioned in the dorso-
medial part of the dental (Fig. 4b).
Suspensorium: The suspensorium comprises the
autopalatine, endopterygoid, ectopterygoid,
metapterygoid, hyomandibular, quadrate and
symplectic bones (Fig. 5). The autopalatine possesses
a blade-like process in the middle and it is connected
159
to the prevomer laterally. Also, this bone is
connected to the prepalatine, preethmoid-II and
endopterygoid anteriorly and posteriorly,
respectively. The endopterygoid is elongated and it is
connected to the metapterygoid and ectopterygoid
ventrally. This bone is connected to the autopalatine
by a condyle anteriorly. The metapterygoid is almost
rectangular in shape and positioned between the
hyomandibular and quadrate. The hyomandibular is
extended longitudinally and its dorsal part is wider.
This bone has two developed processes in its anterior
and posterior margins and connected to the interhyal
and symplectic ventrally and to the metapterygoid
anteriorly. There are two hyomandibular condyles in
the dorsal margin of the hyomandibular for
articulation to the neurocranium.
The quadrate has a pointed and stretched ventral
process inclined posteriorly (Fig. 5). Also, this bone
is connected to the endopterygoid dorsally and
metapterygoid posteriorly. The symplectic is almost
triangular in shape and situated under the
endopterygoid and posterior to the quadrate. The
ectopterygoid bears a pointed process anteriorly and
a downward process ventrally.
Opercular series: The opercular series consist of the
opercle, preopercle, subopercle, and interopercle
(Fig. 5). The opercle is the largest element in this
series and has a rod-shaped process antero-dorsally
for connecting to the levator operculi muscle. The
ventral margin of the opercle is connected to the
subopercle. The subopercle is a stretched bone and
connected to the interopercle anteriorly. The
preopercle is narrow and arc-shaped, and situated on
the interopercle. The interopercle is elongated and
wider at its middle part.
Branchial arches: The branchial arch consists of the
unpaired basibranchial and paired hypobranchial,
ceratobranchial, epibranchial, and pharyngobranch-
ials (Fig. 6). There are four basibranchials in this
series that the fourth one is very small. Five
ceratobranchials are the largest elements of the
branchial arch that fifth one is modified into the
pharyngeal teeth. The numbers of the epibranchial
 Iranian Journal of Ichthyology (September 2015), 2(3): 155-164
Fig.6. Branchial apparatus of Paraschistura nielseni.
Bbr: basibranchial; Cbr: ceratobranchial; Ebr:
epibranchial; Hbr: hypobranchial; Pbr:
pharyngobranchials.
are four; also there are two pharyngobranchials and
three hypobranchials.
Hyoid arch: The hyoid arch is composed of the
basihyal, hypohyals, ceratohyal, epihyal, interhyal,
urohyal and branchiostegal rays (Fig. 7). The
unpaired basihyal is T-shaped and its anterior part is
wider. The hypohyals are consisted of the dorsal and
ventral parts. The ceratohyal is the largest bone in the
hyoid arch which is situated between the hypohyal
and epihyal. The epihyal is almost triangular in shape
with a pointed process posteriorly. The interhyal is a
cylindrical bone connecting to the epihyal ventrally,
and to the hyomandibular and symplectic dorsally.
The unpaired urohyal has two ventral and dorsal
parts. The dorsal part is blade-like and perpendicular
to the ventral part. The branchiostegal rays are
narrow and extended to the dorsal margin of the
subopercle. The first and second branchiostegal rays
are connected to the middle of the ceratohyal and at
the junction of the ceratohyal and epihyal,
respectively. Also, the third one is situated in the
160
Fig.7. Hyoid arch of Paraschistura nielseni. Bhy:
basihyal; Br: branchiostegale; Chy: ceratohyal; Dhy
and Vhy: dorsal and ventral hypohyal; Ehy: epihyal;
Ihy: interhyal; Uhy: urohyal.
middle of the epihyal.
Pectoral girdle: The pectoral girdle includes the
cleithrum, supracleithrum, coracoid, mesocoracoid,
scapula, posttemporal, supratemporal and radials
(Fig. 8). The small supratemporal is positioned
antero-lateral to the elongated posttemporal. The
supracleithrum is wide and it is connected to the
cleithrum dorso-ventrally. The posttemporal is a long
bone situating between the supracleithrum and
supratemporal. Also, this bone is connected to the
epiotic posteriorly, through which the pectoral girdle,
is connected to the neurocranium. The cleithrum is
the largest element of the pectoral girdle and it is
connected to the supracleithrum dorsally and to the
coracoid through mesocoracoid latero-medially. The
anterior part of the coracoid is narrow, while its
posterior part is wide. The scapula has a large
opening (scapula foramen) and it is positioned
between the cleithrum and coracoid. The radials are
four in number among which the third one is
significantly wider than the others.
Dorsal fin skeleton: The dorsal fin skeleton bears 10
pterygiophores and one stay. The first pterygiophore
is positioned in the front of the seventh centrum and
 Azimi et al.-Osteological characteristics of Paraschistura nielseni

of Cobitis avicennae

Fig.8. Pectoral girdle of Paraschistura nielseni. Cl: cleithrum; Cor: coracoid; Mcor: mesocoracoid; pect-R: ray of the
pectoral fin; Rad: ossified pectoral radial; Sc: scapula.

Fig.9. Dorsal fin (a), anal fin (b) and pelvic girdle (c) of Paraschistura nielseni. Adp: anal distal pterygiophore; Dfr:
dorsal fin rays; Dfs: dorsal fin spin; Dr: distal radial; Mp: mesial pterygiophore; Mr: medial radial; Pp: pterygiophore;
Sty: stay.

a stay is located at the end of the tenth pterygiophore
(Fig. 9a). The numbers of unbranched and branched
rays are four and 8½, respectively.
Anal fin skeleton: The anal fin skeleton includes
seven pterygiophores and one stay. The first
161
pterygiophore is positioned in the front of the
nineteenth centrum. In addition, this fin bears three
unbranched and 5½ branched rays (Fig. 9b).
Pelvic girdle: The pelvic girdle consists of the paired
pelvic bones and radials (Fig. 9c). These bones are
 Iranian Journal of Ichthyology (September 2015), 2(3): 155-164
Fig.10. Caudal skeleton of Paraschistura nielseni.
Epu: epural; Hp: hypural; Npu2: neural spine of the
second preural centrum; Hpu2: hemal processes of the
second preural centrum; Ph: parhypural; Pst:
pleurostyle.
enclosed by muscles. The pelvic bones have three
processes consisting the anterior (pubic) process,
postero-lateral (iliacus) process that rays connected
on it and posterior (ischiadicus) process. The anterior
part of the pelvic bones (pubic process) is narrower
(Fig. 9c). On each side of the paired pelvic bones,
there are three small and rounded radials positioning
between the pelvic bones and rays. Furthermore, a
developed styloid bone is found lateral to the
unbranched rays.
Caudal skeleton: The hypural-1 of the caudal
skeleton is wide and connected to the centrum by a
rod-shaped process and parhypural ventrally (Fig.
10). The parahypural is relatively flat and it is
connected to the centrum posteriorly. The hypurals-
3, 4, and 5 are positioned between the pleurostyle and
hypural-2. The epural is connected to the pleurostyle,
and it is situated between the pleurostyle and
rudimentary neural arch.
Weberian apparatus and swim bladder capsule: The
Weberian apparatus comprises of the claustrum,
scaphium, intercalarium, and tripus (Figs. 11a-b).
162
Fig.11. The swim bladder capsule of Paraschistura
nielseni. Cla: claustrum; Dpr-2 -4: descending
processes of the second and fourth centra; Hpr-2-4:
horizontal processes of the second and fourth centra;
Na4: neural arch 4; Sca: scaphium; Sn2: supraneural
2; Sn3: supraneural 3.
The claustrum is a round and small bone situating on
the scaphium ventrally and it is connected to the
supraneural-2 dorsally. The intercalarium is a small
bone locating between the scaphium and tripus. The
first centrum does not participate in the formation of
the bony capsule. The second and third centra are
fused and the second, third, and fourth centra
participate in the formation of the bony capsule. The
third centrum is located between the supraneural-2
and fourth centrum. The parapophysis of the forth
centrum is modified and participated in the posterior
part of the bony capsule. In lateral side of bony
capsule, two openings are present and the posterior
one is larger. The bony capsule is rectangular in
 Azimi et al.-Osteological characteristics of Paraschistura nielseni
of Cobitis avicennae
shape and its ventral surface is not alveolar. Also, the
right and left lobes of the bony capsule are
symmetrical and divided by the manubrium (Fig.
11b).
Discussion
In the present study, the skeletal system of P. nielseni
was described in detail. Based on the results, the
length of the occipital region in P. nielseni is less than
one-third of the neurocranium length similar to the
majority of the loach species (Prokofiev 2010). In
P. nielseni, the supraethmoid-ethmoid and prevomer
are fused similar to other loaches, with the exception
of Lefua spp., Oreonectes platycephalus, Yunnanilus
pleurotaenia, Triplophysa microphthalma, T. tenuis
(Prokofiev 2010) and Schistura fasciolata (Sawada
1982). The connection of the lateral ethmoid to the
neurocranium in P. nielseni is at the level of the
anterior margin of the orbitosphenoid, similar to
Oxynoemacheilus kiabii (Mafakheri et al. 2014) and
O. bergianus (Jalili & Eagderi 2015). However,
based on Prokofiev (2010), this bone in loaches is
connected to the supraethmoid-ethmoid.
There is no preethmoids-I in P. nielseni, whereas
in other genera such as Lefua, Oreonectes,
Yunnanilus, Eonemachilus, Micronoemacheilus,
Hedinichthys, Orthrias and Triplophysa, paired
preethmoids-I present connecting to the lateral edge
of the anterolateral processes of the prevomer
(Prokofiev 2010). In P. nielseni, the sesamoids are
not completely ossified, but in O. kiabii, the
sesamoids are clearly ossified (Mafakheri et al.
2014). Whereas, the presence of the sesamoid
ossifications in loaches was rejected (Sawada 1982);
the sesamoid ossifications were reported in
Paracobitis malapterura, P. longicauda, Dzihunia
amudarjensis and Oxynoemacheilus angorae
(Prokofiev 2004, 2009). The prevomer is square-
shaped in P. nielseni similar to O. kiabii (Mafakheri
et al. 2014) whereas, this bone is elongated and
rectangular in O. bergianus (Jalili & Eagderi 2015).
The sphenotic and epiotic are contacted in P. nielseni,
similar to Paraschistura cristata and the members of
163
the genus Oxynoemacheilus and Paracobitis
(Prokofiev 2010).
The anterior facet of the hyomandibular in
P. nielseni, is formed by the pterosphenoid and
sphenotic, and the posterior one by the pterotic and
sphenotic while in O. kiabii (Mafakheri et al. 2014)
and O. bergianus (Jalili & Eagderi 2015), the anterior
facet is formed by the prootic, pterotic and sphenotic.
Whereas, Prokofiev (2010) noted that the anterior
facet is formed by the sphenotic and prootic, and the
posterior one by the sphenotic, prootic and pterotic.
The coronomeckelian of P. nielseni is situated on
the dorso-medial part of the articular similar to that
of O. kiabii (Mafakheri et al. 2014) and O. bergianus
(Jalili & Eagderi 2015), but Prokofiev (2010) noted
that the coronomeckelian is connected to the base and
dorsal edge of the coronoid process in loaches.
There are four basibranchials in P. nielseni
similar to O. kiabii (Mafakheri et al. 2014) and
O. bergianus (Jalili & Eagderi 2015), but Prokofiev
(2010) reported three basibranchials in the genus
Paraschistura. The two extra urohyals were not find
in P. nielseni; however, in O. kiabii (Mafakheri et al.
2014) and O. bergianus (Jalili & Eagderi 2015), they
exist. Also, these bones are not reported by Prokofiev
(2010) for thee members of this family. Paraschistura
nielseni has five hypurals in its caudal skeleton, while
O. bergianus (Jalili & Eagderi 2015) has six hypurals.
In P. nielseni, the base of the right pelvic bones is
separated from the main body which is not observed
in O. kiabii (Mafakheri et al. 2014) and O. bergianus
(Jalili & Eagderi 2015). In P. nielseni, the bony swim
bladder capsule is rectangular-shaped and its ventral
face is not alveolar similar to O. bergianus (Jalili &
Eagderi 2015), while the ventral face in O. kiabii
(Mafakheri et al. 2014) is alveolar.
In conclusion, the findings of this research
showed that P. nielseni has a number of distinct and
important osteological characteristics including a
square-shaped prevomer, four basibranchials, a semi-
ossified sesamoid, a non-alveolar bony swim bladder
capsule and separated pelvic bone bases that can be
used to distinguish this species from other loach
 Iranian Journal of Ichthyology (September 2015), 2(3): 155-164
species. It may also be possible to use these
osteological characteristics as a reference in the
identification, taxonomy and phylogenetic
relationship of other loach species in this genus in the
future.
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