Int. J. Aquat. Biol. (2016) 4(2): 117-124; DOI:  

ISSN: 2322-5270; P-ISSN: 2383-0956

Journal homepage: www.ij-aquaticbiology.com 
© 2016 Iranian Society of Ichthyology 

Original Article 
The complete description of the skeletal structure of Hafez loach,  

Turcinoemacheilus hafezi (Cypriniformes, Nemacheilidae) 
  

Nasrin Nikmehr, Soheil Eagderi*,1Hadi Poorbagher, Hamid Farahmand 

 
Department of Fisheries, Faculty of Natural Resources, University of Tehran, Karaj, Iran. 

 

 

 

 

 

 

Article history: 
Received 21 January 2016 

Accepted 17 April 2016 

Available online 2 5 April 2016 

Keywords:  
Nemacheilidae 

Osteology 

Loach 

Turcinoemacheilus 
Iran 

Abstract: The osteological characteristics is an important tool for clarification of the phylogenetic 
statue of the family Nemechelidae. Since any information is available about osteological features of 

the genus Turcinoemacheilus, hence the present study provides a detailed osteological characteristics 

of the Turcinoemacheilus hafezi as representative of this genus. Ten specimens of T. hafezi were 

collected from the Shalamzar Stream, Tigris basin, Iran and cleared and stained for osteological 

examination. According to the results, T. hafezi is osteologicaly characterized by absence of the 

preethmoideum-I and postcleithrum, having four basibranchials, presence of the sesamoid 

ossification, free and short epural with the reduced neural process and pleurostyle, connection of the 

hypural-1 to the parahypural, no connection of the hypurals 3, 4, and 5 to the pleurostyle, and no 

bony bridge between the parietal. 
 

Introduction 

Nemacheilids are small and benthic species that 

found in flowing waters of Europe, Asia, and 

northeast of Africa (Coad, 2016). Loaches of the 

genus Turcinoemacheilus, a member of the family 
Nemacheilidae, is easily distinguished from other 

members of this family by a more anterior position 

of the anus (Bânârescu and Nalbant, 1964; Freyhof 

et al., 2011). Turcinoemacheilus was a monotypic 
genus for 47 years (Conway et al., 2011; 

Golzarianpour et al., 2013). Turcinoemacheilus 
kosswigi Bănărescu and Nalbant, 1964 was recorded 
for the first time in Iran from the Karoun River 

drainage (Golzarianpour et al., 2009). Turcinoema-
cheilus hafezi, a new species from Karoun and Dez 
rivers drainages of Iran (Golzarianpour et al., 2013), 

was second species of this genus from Iran. Also, 

Esmaeili et al. (2014) described two new species of 

the genus Turcinoemacheilus from Iran, including 
T. bahaii from the Zayandeh River and T. saadii from 
the Karoun River drainage in Iran. 

Turcinoemacheilus hafezi Golzarianpour et al. 

                                                           
* Corresponding author: Soheil Eagderi 

E-mail address: soheil.eagderi@ut.ac.ir 

2013 is distinguished from the other species of this 

genus by the more posterior position of the anus, 

which is closer to the anal-fin origin than to the 

pelvic-fin origin (distance from anus to anal-fin 

origin 0.2-0.4 times in distance from pelvic-fin 

origin to anal-fin origin), a mottled colour pattern or 

a prominent irregular dark brown or black mid-

lateral stripe disconnect from blotches and saddles 

on back, a slightly emarginated caudal fin and a 

completely scaleless body (Golzarianpour et al., 

2013).  

Due to small size and low economic value, the 

members of the family Nemacheilidae are less 

known (Kottelat, 1990) and their classification is still 

complicated; therefore, ichthyologists are trying to 

reveal their phylogenic status (Prokofiev, 2010). 

Hence, the osteological characteristics can play an 

important role in this regard (Sawada, 1982; 

Mafakheri et al., 2015). The only comprehensive 

osteology and phylogenetic studies on nemacheilids 

performed by Prokofiev (2009, 2010), but in these 

studies, the Turcinoemacheilus did not include. 



118 
 

Nikmehr et al./ Osteology of Turcinoemacheilus hafezi 

Since, there is no information is available about the 

osteological features of the genus Turcinoema-
cheilus, therefore, this study aimed to provide a 
detailed osteological description of Hafez loach, 
T. hafezi from the Karoun river drainage.  
 

Materials and Methods 

Ten specimens of T. hafezi with average standard 
length of 38.22(±0.64) (Mean±SD) were collected 

by electrofishing device from the Shalamzar Stream 

(Charmahal-e Bakhtiari Province, Iran) (Fig. 1). 

Then, the specimens were anesthetized in 1% clove 

solution and fixed in 10% buffered formaldehyde. 

For osteological examination, the specimens were 

cleared and stained using alcian blue and alizarn red 

based on Taylor and Van Dyke (1985). Images of the 

stained skeletal structure obtained by a scanner 

(Epson V600) equipped to a glycerin bath. Skeletal 

structures were observed and studied by a 

stereomicroscope (Leica MS5). Drawings of the 

skeletal elements were performed using CorelDraw 

X6 software. Nomenclature and abbreviation of the 

skeletal elements were based on Prokofiev (2009). 

 

Results 

The neurocranium is wider posteriorly, forming the 

maximum width of the skull at the level of the 

pterotic. The ethmoid region is comprised of a pair 

of the lateral ethmoid, and unpaired prevomer and 

supraethmoid-ethmoid bones (Fig. 2A). The paired 

L-shaped lateral ethmoids posteriorly form the 

anterior part of the orbit. The prevomer is relatively 

flat having two processes with rounded edges antero-

laterally. This bone is connected to the 

orbitosphenoid and parasphenoid posteriorly (Fig. 

2B). The supraethmoid-ethmoid is narrower in the 

middle part. The kinethmoid is a small and free bone 

situated between two maxillae. This bone has two 

pointed lateral processes in the middle part, forming 

its maximum width. 

The orbital region includes the frontal, 

orbitosphenoid, ptersphenoid, parasphenoid, 

lachrymal, and sclerotic bones. The paired large 

frontals are asymmetric and have two processes in 

latero-middle part; its anterior part is narrow. These 

bones include about half of the length of the 

neurocranium separating by the fontanel posteriorly. 

The frontal is connected to the orbitosphenoid 

antero-laterally. The posterior margins of the frontals 

contribute in formation of the fontanel. The 

ptersphenoid is connected to the frontal dorsally and 

to the sphenoid posteriorly, forming the posterior 

part of the orbit. The parasphenoid is bifurcated at 

two ends and its middle part is connected to the 

pterosphenoids. It has two small lateral and one large 

middle pores. The orbitosphenoid is connected to the 

lateral ethmoid ventrally, to the parasphenoid antero-

laterally, and to the pterosphenoid posteriorly (Fig. 

2). 

The otic region includes five bones viz. the 
parietal, epiotic, sphenotic, pterotic, and prootic. The 

parietal is posteriorly connected to the supraoccipital 

and epiotic and ventrally to the prootic. In addition, 

no bony bridge present between the parietal and 

pterotic. The parietal is roughly rectangular. The 

pterotic is almost triangular in shape. There is no 

connection between the parietals due to a long 

fontanel. The sphenotic bears a lateral process and a 

Figure 1. Turcinoemacheilus hafezi from the Shalamzari Stream River. 



119 
 

Int. J. Aquat. Biol. (2016) 4(2): 117-124 

 

(a) 

(c) 

(b) 

latero-ventral facet.  

The occipital region is composed of the 

supraoccipital, exoccipitals, and basioccipital. The 

supraoccipital has a pentagon shape; it is connected 

to the exoccipitals ventrally. The exoccipital 

encloses the foramen magnum and bears a small 

foramen laterally. The basioccipital bears a posterior 

process with a large cavity centrally. There is an 

occipital condyle at the posterior part of the 

basioccipital for connecting to the vertebrae. 

In the branchiocranium, the upper jaw consists of 

the maxilla and premaxila (Fig. 3a). The maxilla is a 

Figure 2. The neurocranium of Turcinoemacheilus hafezi. (a) Dorsal, (b) lateral, and (c) ventral views. Apl: autopalatine, fon: fontanel, fr-Exo: 
foramen exoccipital, pr-Bo: basioccipital process, Epo: epiotic, Exo: exoccipital, Fr: frontal, Let: lateral ethmoid, Orb: orbitosphenoid, Pa: parietal, 

Pe: prevomer, Pro: prootic, Ps: parasphenoid, Pto: pterotic, Pts: pterosphenoid, Se: supraethmoid, Soc: supraoccipital, and S po: sphenotic. 



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Nikmehr et al./ Osteology of Turcinoemacheilus hafezi 

large laminar bone and its middle part is the wider; 

its anterior part is narrow with an antero-ventral 

process tilting ventrally. The premaxila is L-shaped 

comprising two horizontal and vertical parts. The 

horizontal part is arc-shaped and slightly wider in its 

middle part and the vertical part is narrower and 

longer. There is two small triangular process at the 

middle portion of the vertical part.  

The lower jaw is composed of four bones, 

including the dental, angular, retroarticular and 

coronomeckelian (Fig. 3b). The dental is the largest 

element of the mandible, including a narrow antero-

ventral branch and a deep section (process 

coronoideus) postero-dorsally; it is connected to the 

articular postero-dorsally and to the retroarticular 

dorsally. The coronomeckelian is triangular in shape 

and situated in the medial side of the dental beneath 

the process coronoideus. 

The suspensorium is formed by the 

hyomandibular, ectopterygoid, endopterygoid, 

metapterygoid, symplectic, quadrate, and 

autopalatine (Fig. 4). The hyomandibular is a 

relatively large bone extending longitudinally that its 

dorsal part is wider. In addition, there are two 

anterior and posterior hyomandibular condyles on its 

dorsal margin. The quadrate bears a condyle antero-

ventrally where it is articulated to the dental. The 

ectopterygoid, endopterygoid, metapterygoid, 

symplectic, and quadrate form a complex connecting 

to the anterior part of the neurocranium via 

autopalatine. The autopalatine is wider and 

posteriorly connected to the endopterygoid and 

anteriorly to the prevomer. The endopterygoid is 

pointed posteriorly connecting to the ectopterygoid 

Figure 3. The upper (a) and lower (b) jaws of Turcinoemacheilus hafezi. Mx: maxilla, Pmx: premaxilla, Art: articular, Cm: coronomeckelian, 
Den: dental, and Ret: retroarticular. 

Figure 4. The suspensorium, palatine, and opercular series of Turcinoemacheilus hafezi. Apl: atutopalatine; Ect: ectopterygoid, End: 
endopterygoid, Hm: hyomandibular, Io: interopercle, Mtp: metapterygoid, Op: opercle, Q: quadrate, So: subopercle, and Sym: symplectic. 



121 
 

Int. J. Aquat. Biol. (2016) 4(2): 117-124 

 

antero-ventrally and metapterygoid posteriorly. The 

metapterygoid bears several pores on its lateral face. 

The ectopterygoid is situated anterior to the 

quadrate.  

The opercular series consists of the opercle, 

peropercle, subopercle, and interopercle (Fig. 4). 

The opercle is the largest element of this series and 

possesses a condyle anteriorly. The paddle-shaped 

subopercular is located ventral part to the opercle. 

The narrow preopercular is pointed anteriorly and 

bears a notch at its posterior rim. The interopercular 

is a curved bone with pointed anterior part. 

The branchial apparatus includes five pairs of the 

ceratobranchial, four pairs of the epibranchiasls, 

three pairs of the hypobranchials and 

pharyngobranchials, and four unpaired basi-

branchials (Fig. 5a). The anterior part of the first and 

second basibranchials are the wider. Two first 

hypobranchials are similar in shape and the last one 

is the smallest. Four pairs of the ceratobranchials are 

situated between the hypobranchials and 

epibranchials and the last ceratobranchials are 

modified as the tooth plats. The last epibranchial 

curves dorsally and its anterior part is narrower. 

The hyoid arch consists of the paired interhyal, 

epihyals, ceratohyals, dorsal and ventral hypohyals, 

and the unpaired urohyal and basihyal, and three 

pairs of the branchiostegals (Fig. 5b). The urohyal is 

T-shaped and anteriorly bifurcated. The basihyal is 

also T-shaped with a deep notch on its anterior end. 

The dorsal and ventral hypohyals are connected 

firmly. The ceratohyals are the largest elements of 

the hyoid arch. There are three curved long 

branchiostegals; the first one is attached to the 

ceratohyal, the second one connected between the 

ceratohyal and epihyal, and the third one is attached 

to the epihyal. The interhyal is small with pointed 

posterior end (Fig. 5b). 

In the postcranial skeleton, the pectoral girdle 

consists of the cleithrum, supracleithrum, coracoid, 

mesocoracoid, scapula, posttemporal, supratemporal 

and radials (Fig. 6a). The largest element of this 

complex is the cleithrum that bears a longer vertical 

and a horizontal sections. The dorsal part of the 

supracleithrum is wider. The coracoid is connected 

to the cleihtrum posteriorly and middle-ventrally. 

The mesocoracoid is narrow and its ventral part ia 

wider and connected to the coracoid. The 

mesocoracoid is firmly connected to the cleithrum. 

The scapula is trapezoid in shape and connected to 

the coracoid ventrally. The pectoral girdle bears four 

radials that overlap together and first one is the 

widest. The pectoral fin has one unbranched and 9 

branched rays.  

Figure 5. Dorsal view of the branchial apparatus (a) and hyoid arch (b) of Turcinoemacheilus hafezi. Bbr: basibranchial, Cbr: ceratobranchial, 
Ebr: epibranchial, Hbr: hypobranchial, Pbr: pharyngobranchial, Bhy: basihyal, Chy: ceratohyal, Dhy and Vhy: dorsal and ventral hypohyal, Epi: 

epihyal, Ihy: interhyal, and Uhy: urohyal. 



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Nikmehr et al./ Osteology of Turcinoemacheilus hafezi 

The pelvic girdle includes the paired pelvic bones, 

pelvic splint and radials (Fig. 6b). The pelvic bone is 

horizontally positioned in the middle part of the 

belly. It has two long processes anteriorly and its 

posterior part is wider. The pelvic bone has a round 

process latero-posteriorly. The pelvic girdle 

possesses three radials that the medial one is the 

smallest one. The styloid is situated lateral to the 

radials. The pelvic fin bears one unbranched and 7 

branched rays.  

The dorsal fin bears 4 unbranched and 6 branched 

rays, 8 pterygiophores and one stay bone. The first 

pterygiophor is next to the 17th or 18th vertebra. The 

largest bone of the dorsal fin is the first 

pterygiophores that supports 4 unbranched rays. The 

first and last petrygiophores are bifurcated ventrally. 

A triangular stay supports the last branched ray (Fig. 

7a). The anal fine originates at 26th centrum; it has 4 

Figure 6. Medial view of pectoral girdle (a) and pelvic girdle (b) of Turcinoemacheilus hafezi. Cl: cleitherum, Cor: coracoid, Mcor: mesocoracoid, 
Pb: pelvic bone, Ps: pelvic splint, R: radials, and Sc: scapula. 

Figure 7. Dorsal (a), anal (b) and caudal (d) fin skeleton of Turcinoemacheilus hafezi. Adp: anal distal pterygiophore; C 18-26: 18th and 26th 
centrum; Dfs: dorsal fin spine; Mp: mesial pterygiophore; Ph; parhyporal; Pp: pterygiophore; Sty: stay; Epu: epural; Hp1–5: hypural plates; Npu 

2: neural processes of the second preural centrum; Hpu 2: hemal processes of the second preural centrum; Ust: pleurostyle. 



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Int. J. Aquat. Biol. (2016) 4(2): 117-124 

 
unbranched and 5 branched rays and is supported by 

6 pterygiophores and one small stays bone (Fig. 7b).  

The caudal skeleton is made up of four centra 

along with the epural, parhypural, pleurostyle, 

uroneural and 5 hypurals (Fig. 7c). The anterior part 

of the hypural-1 is wider and its anterior end reaches 

to the last centrum. The hypural-1 is also connected 

to the parahypural ventrally. The parahypural is 

relatively flat, and its posterior end is connected to 

the last centrum. The hypurals 3, 4, and 5 are 

positioned between the pleurostyle and hypural-2. 

The neural process of the second pleural is long and 

narrow. The caudal fin bears 18 branched rays. The 

number of the dorsal procurrent is 5 and thar of the 

ventral procurrent is 6. 

 

Discussion 

The present study provided a detailed skeletal 

description of the T. hafezi. Based on the results, 
T. hafezi showed various differences with other 
members of the family Nemacheilidae. In the 

neurocranium, the lateral ethmoid, supraethmoid-

ethmoid and prevomer are fused, as other loaches, 

with the exception of Oreonectes platycephalus, 
Lefua spp., Yunnanilus pleurotaenia, Triplophysa 
microphthalma, T. tenuis and Schistura fasciolata 
(Sawada, 1982). In contrast to T. kosswigi, the L-
shaped lateral ethmoid of T. hafezi has a process 
posteriorly. Turcinoemacheilus hafezi bears a 
sesamoid ossification similar to T. kosswigi (Azimi, 
2014), Paracobitis malapterurus (Azimi et al., 
2015a), P. longicauda, Dzihunia amudarjensis and 
Oxynoemacheilus angorae (Prokofiev, 2004, 2009). 
However, the presence of the sesamoid ossifications 

in loaches had been rejected previously (Sawada, 

1982).  In T. hafezi similar to T. kosswigi, (Azimi, 
2014), the supraethmoid-ethmoid is tightly attached 

to the frontals, as other loaches with the exception of 

Indoreonectes evezardi (Sawada, 1982). There are 
two opening on the ventral part of the exoccipital in 

T. hafezi similar to T. kosswigi (Azimi, 2014),  
O. platycephalus, L. costata, Orthrias barbatulus, 
T. stoliczkae, T. alticeps, T. strauchii, D. amudarj-
ensis, O. angorae, Micronoemacheilus pulcher, 

Iskandaria kuschakewitschi and Afronemacheilus 
abyssinicus, with the exception of P. cristata 
(Prokofiev, 2010; Azimi et al., 2015b). 

In branchial apparatus of T. hafezi, there are four 
basibranchials similar to T. kosswigi (Azimi, 2014), 
O. angorae and O. brandti (Mafakheri et al., 2015) 
that fourth one is very small; Prokofiev (2010) are 

reported three basibranchials in Lefua spp., 
O. platycephalus, Y. pleurotaenia, Hedinicthys, in 
the majority of species of the genus Orthrias (with 
the exception of O. dgebuadzei), in I. 
kuschakewitschi, Paracobitis longicauda, P. malapt-
erurus, Schistura spp. Paraschistura, Nemacheilus 
masyae, Nun, Seminemacheilus, Acanthocobitis 
botia, and I. evezardi (Prokofiev, 2010; Azimi et al., 
2015a, b). In addition, the basihyal of T. hafezi has a 
deep notch compared to that of T. kosswigi (Azimi, 
2014; Azimi et al., 2015a, b).  

In the caudal skeleton, the hypoural-1 of 

T. kosswigi attaches to the terminal centrum similar 
to O. brandti (Prokofiev, 2010; Mafakheri et al., 
2015). The hypural-1 is connected to the parahypural 

ventrally in T. hafezi similar to T. kosswigi (Azimi, 
2014), and unlike majority of loaches with the 

exception of O. angorae, Triplophysa orientalis, and 
T. scleroptera (Prokofiev, 2010). The epural of 
T. hafezi is long and not connected to the reduced 
neural process of the terminal centrum similar to 

P. cristata (Prokofiev, 2010; Azimi et al., 2015b), 
whereas in T. kosswigi, it is fused to the urostyle 
(Azimi, 2014).  

According to Prokofiev (2010), the most stable 

and important diagnostic and phylogenic feature of 

the pectoral girdle in loaches is the number and shape 

of bony radials in the pectoral fin; unlike 

O. platycephalus and Lefua spp., which have two 
radials (Sawada, 1982), there are three pelvic radials 

in T. hafezi and T. kosswigi (Azimi, 2014) like other 
loaches. In loaches, the caudal skeleton has 5-6 

hypurals (Prokofiev, 2010), which five hypurals 

were observed in T. hafezi similar to T. kosswigi 
(Azimi, 2014). Finally, since the identification of the 

species of the genus Turcinoemacheilus is difficult 
based on morphometric characteristics and color 



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Nikmehr et al./ Osteology of Turcinoemacheilus hafezi 

pattern, the present study provides a useful 

distinguishing osteological features of this genus 

from the other members of the nemacheilid family. 

 

Acknowledgments 

We would like to express our sincere thanks to 

A. Jouladeh-Roudbar for his help during sampling, 

H. Mousavi-Sabet for providing some materials and 

P. Jalili for her help during clearing and staining of 

specimens. This study was financially supported by 

the University of Tehran. 

 

References 
Azimi H., Mousavi-Sabet H., Eagderi S. (2015a). 

Osteology of Hyrcanian Crested Loach, Paracobitis 
hircanica (Pisces, Nemacheilidae). Our Nature, 13(1): 
8-18.  

Azimi H., Mousavi-Sabet H., Eagderi S., Vatandoust S. 

(2015b). Osteological characteristics of Turkmenian 

stone loach, Paraschistura cristata (Cypriniformes: 
Nemacheilidae). International Journal of Aquatic 

Biology, 3(5): 290-300. 

Azimi H. (2014). Phylogeny of five Iranian genus of 

Balitoridae based on morphological and osteological 

characteristics. MSc Thesis, University of Guilan, 

Iran. 

Breil M., Bohlen J. (2001). First record of the loach fish 

Turcinoemacheilus kosswigi in the Basin of Euphrates 
River, with first observations on habitat and behavior. 

Zoology in Middle East, 23: 71-76.    

Bănărescu P., Nalbant T.T. (1964). Süßwasserfische 

derTürkei. 2. Teil Cobitidae. Mitteilungen aus dem 

Hamburgischen Zool. Museum und Institut, 61: 159-

201. 

Coad B.W. (2016). Freshwater Fishes of Iran. (Available 

at http://www.briancoad.com) (Accessed on 10 

February 2016). 

Conway K.W., Edds D.R., Shrestha J., Mayden R.L. 

(2011). A new species of gravel-dwelling loach 

(Ostariophysi: Nemacheilidae) from the Nepalese 

Himalayan foothills. Journal of Fish Biology, 79: 

1746-1759. 

Esmaeili H.R., Sayyadzadeh G., Özulug M., Geiger M., 

Freyhof J. (2014). Three new species of 

Turcinoemacheilus from Iran and Turkey (Teleostei: 
Nemacheilidae). Ichthyological Exploration of 

Freshwaters, 24(3): 257-273. 

Freyhof J., Erkakan F., Ozeren C., Perdices A.J. (2011). 

An overview of the western Palaearctic loach genus 

Oxynoemacheilus (Teleostei: Nemacheilidae). 
Ichthyological Exploration of Freshwaters, 22: 301-

312.  

Golzarianpour K., Abdoli A.,  Kiabi B.H., Freyhof J. 

(2009). First record of the freshwater loach, 

Turcinoemacheilus kosswigi (Bănărescu and Nalbant, 
1964) from Iran (Karoun drainage),” Zoolgy in 

Middle East, 47: 57-62. 

Golzarianpour K., Abdoli A., Patimar R., Freyhof J. 

(2013). Turcinoemacheilus hafezi, a new loach from 
Karoun River drainage, Iran (Teleostei: 

Nemacheilidae). Ichthyological Exploration of 

Freshwaters, 24: 41-48. 

Kottelat M. (1990). Indochinese Nemachilines: a revision 

of Nemachiline loaches (Pisces: Cypriniformes) of 

Thailand, Burma, Laos, Cambodia. Southern Viet 

Nam (Pfeil, München). 

Mafakheri P., Eagderi S., Farahmand H., Mosavii-Sabet 

H. (2015). Descriptive Osteology of 

Oxynoemacheilus kermanshahensis (Bănărescu and 
Nalbant, 1966) (Cypriniformes, Nemacheilidae). 

Croatian Journal of Fisheries, 73: 115-123 

Nalbant T.T., Bianco P.G. (1998). The Loaches of Iran 

and adjacent regions with description of six new 

species (Cobitoidea). Italian Journal of Zoology, 65: 

109-123. 

Prokofiev A.M. (2004). Osteology and relationships 

between loaches of the genus Dzihunia (Osteichthyes, 
Balitoridae). Zoologicheskii Zhurnal, 83: 826-838. 

Prokofiev A.M. (2009). Problems of the classification 

and phylogeny of nemacheiline loaches of the group 

lacking the preethmoid I (Cypriniformes: Balitoridae: 

Nemacheilinae). Journal of Ichthyology, 49(10): 874-

898. 

Prokofiev A.M. (2010). Morphological classification of 

loaches (Nemacheilinae). Journal of Ichthyology, 

50(10): 827-913. 

Sawada Y. (1982). Phylogeny and zoogeography of the 

superfamily Cobitoidea (Cyprinoidei, Cyprini-

formes). Memoirs of Faculty of Fisheries Hokkaido 

University, 28(2): 65-223. 

Taylor W.R., Van Dyke G.C. (1985). Revised procedures 

for staining and clearing small fishes and other 

vertebrates for bone and cartilage study. Cybium, 9: 

107-119.