euperipatoides kanangrensis is a species of velvet worm of the peripatopsidae family , described in 1996 from specimens collected in kanangra-boyd national park , new south wales .

it is endemic to australia .

euperipatoides kanangrensis

collection and culturing of female euperipatoides kanangrensis reid , 1996 was done as described previously ( 48 ) . in situ hybridization was carried out as described by eriksson et al . ( 49 ) .
neurogenesis in the onychophoran e . kanangrensis does not reflect the ancestral pattern of euarthropod neurogenesis .
here we show that the onychophoran nervous system is generated by the massive irregular segregation of single neural precursor cells . furthermore , we have identified homologs of the euarthropod achaete - scute , notch , and delta genes in the onychophoran euperipatoides kanangrensis and have analyzed their expression patterns in the embryonic ventral neuroectoderm . the data show that these genes are not regulated in a precise spatiotemporal pattern as in euarthropods , and thus the selection of neural precursors is distinct in the onychophoran .
in contrast , in e . kanangrensis , a large number of neural precursors segregate from the ventral neuroectoderm . the number and arrangement of neural precursors is different in each hemineuromere within the same embryo and also between embryos of the same stage . these data suggests that the neural precursors in onychophorans segregate in a random , irregular pattern ( fig . s5 a ) . if neural precursors were generated in stereotyped positions , one would expect to see a repetitive pattern in contiguous developmental stages , given the long phase of neurogenesis in e . kanangrensis .
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so far , 211 species of velvet worms have been described ( 85 of peripatidae and 126 of peripatopsidae ) , of which 191 are valid species ( 76 of peripatidae and 115 of peripatopsidae ) and 20 represent nomina dubia [ 1–9 ] . in addition to extant representatives , the record of unambiguous fossils includes four species with uncertain relationships to the extant taxa [ 10–13 ] . in the following , we provide an updated classification and a complete list of species of onychophora , which is based on oliveira et al . [ 1 ] .
velvet worms are comparable to
worms with legs
they have an obscurely segmented body with relatively small eyes . they have sensitive antennae used to find food and multiple pairs of legs with slime glands . they catch other smaller insects by entraping them with a mucus lasso . they are unique in that they give birth to live young that go through direct development to become an adult .
tropical regions in the southern hemisphere . they can most commonly be found hiding in rotting logs during the daytime .
the velvet worms are of interest to scientists who study their unique social behavior and reproductive habits .
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► cleavages are superficial and appear identical to that described in the south - african yolk - less ovoviviparous species . ► the initial positioning of the anterior to posterior axis is not fixed in the egg . ► we propose the terms anterior and posterior part of the blastopore to replace older terms . ► the expression of the gene wingless shows the position of the proctodeum . ► this study has clarified the processes that occur in stage 1 of embryonic development in onychophorans .
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m . r . s . conceived the project ; dissected , described and interpreted specimens ; and ran the phylogenetic analysis . j . o . - h . led the integration of developmental data into phylogenetic analysis and the interpretation of results . both authors contributed equally to data analysis , discussion of results and manuscript preparation .
density of scaly ornament in a hallucigeniid spine with three constituent elements ( geological survey of canada 136958 ) .
this file contains details of character coding for phylogenetic analysis , analytical methodology , and transformations implied by our most parsimonious tree .
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edited by thomas c . kaufman , indiana university , bloomington , in , and approved november 10 , 2010 ( received for review june 28 , 2010 )
the formation of neural precursors is first evident in the ventral neuroectoderm at late stage ii [ stages after walker and tait ( 33 ) ] , when the head and the first five trunk segments have formed . single cells delaminate from the neuroectoderm and form a loose basal layer between the outer ectoderm and the inner mesoderm ( fig . 1 a ) . due to the segregation of additional neural precursors , the basal layer increases to about two cells in width but does not expand further during neurogenesis ( fig . 1 b and c ) . in the basal position , the neural precursors divide to generate smaller intermediate neural precursors ( fig . 1 b ) . the latter form a second basal layer of approximately three to four cells in width , and the innermost intermediate neural precursors differentiate into neural cells . neurons extend their processes into the most basal lateral area of the developing hemi - neuromeres to form the neuropile ( fig . 1 d ) . during neurogenesis , the basal area of differentiating neurons expands , whereas the size of the neural precursor and intermediate neural precursor layers remains the same ( fig . 1 e ) . we detected segregating neural precursors in all stages of neurogenesis , indicating that the layer of segregated neural precursors is continuously replenished .
we observed scattered mitotic divisions in all layers of the developing neuromeres except for the basal layers of differentiating neural cells ( fig . 2 ) . large mitotic cells are visible in the ventral neuroectoderm that are partially aligned in longitudinal and diagonal rows ( fig . 2 a and c ) . the segregated neural precursors seem to divide equally to generate two smaller intermediate precursor cells ( fig . 2 g , inset ) . the intermediate neural precursors also proliferate , but are considerably smaller than the dividing neural precursors and neuroectodermal cells ( fig . 2 b ) .
we have analyzed the distribution of neural precursors that can clearly be identified by their position basal to the neuroectoderm , their rounded shape , and the accumulation of f - actin in the cell cortex ( fig . 2 d–g and fig . s1 ) . at early stages , the neural precursors segregate from all areas of the ventral neuroectoderm ( fig . 2 h ) . however , the formation of neural precursors becomes restricted to the center of the developing hemineuromeres during further development ( fig . 2 g and i–k ) , resulting in the generation of ganglion anlagen , which can clearly be distinguished as metameric units ( fig . 2 b and c ) . the number ( ranging from ∼60 to > 100 ) and arrangement of neural precursors is different in each developing hemineuromere within the same embryo but also between embryos of the same stage ( fig . 2 h–k ) .
( a–f ) expression pattern of ekash . light micrographs of flat preparations ( a–d ) and transverse sections ( e and f ) of embryos stained with a dig - labeled ekash probe ; anterior is toward the left in a–d , dorsal is toward the top in e and f . ( a and b ) ventral views show that ekash expression is homogeneous in the neuroectoderm . transcripts are up - regulated in segregated neural precursors ( arrows ) . circles indicate the position of the limb buds . arrowheads in b indicate the area between the neuromeres which shows low / absent levels of ekash . ( c and d ) sagittal views show that ekash is expressed at low , homogenous levels in the neuroectoderm but is strongly expressed in the segregated neural precursors ( arrows ) . arrowhead indicates the area between the developing neuromeres . ( e and f ) ekash is strongly expressed in the segregated neural precursors ( arrow in e ) and the intermediate precursors ( arrowhead in f ) . l1 to l4 , developing neuromeres of walking leg segments 1–4 ; ml , ventral midline ; ms , mesoderm ; vne , ventral neuroectoderm . ( scale bars : 100 μm in a , 100 μm in b , 50 μm in c , and 50 μm in e and f . )
( a–e ) expression pattern of ekdelta . light micrographs of flat preparations ( a–e ) and transverse sections ( e ) of embryos stained with a dig - labeled ekdelta probe ; anterior is toward the left in a–d , dorsal is toward the top in e . ( a–c ) ekdelta is strongly expressed in the ventral neuroectoderm ( arrows ) . expression is lower between the developing neuromeres ( arrowheads ) . ekdelta is expressed in the sensory organ precursors . ( d ) sagittal view of a stage ii embryo shows that ekdelta is expressed in the ventral neuroectoderm , segregated neural precursors , and mesoderm . arrowheads indicate area between segments . ( e ) at stage iv , ekdelta is expressed in the ventral neuroectoderm and segregated ( arrows ) and intermediate neural precursors ( arrowheads ) l1 to l6 , developing neuromeres of walking leg segments 1–6 ; ml , ventral midline ; ms , mesoderm ; sop , sensory organ precursors ; vne , ventral neuroectoderm . ( scale bars : a . 100 μm ; b , 50 μm ; c , 100 μm ; d , 50 μm ; and e , 50 μm . )
( a–c ) expression pattern of eknotch . light micrographs of flat preparations ( a and b ) and transverse section ( c ) of embryos stained with dig - labeled eknotch probe ; anterior is toward the left in a and c ; dorsal is toward the top in b . ( a and c ) eknotch is expressed homogenously in the ventral neuroectoderm ( arrows ) . ( b ) in addition , eknotch is expressed in the segregated ( arrow ) and intermediate neural precursors ( arrowhead ) , the neuropile and the most basal differentiating neurons ( small arrowheads ) , and the mesoderm ( small arrows ) . l1 , l9 to l11 , developing neuromeres of walking leg segments 9–11 ; ml , ventral midline ; ms , mesoderm ; vne , ventral neuroectoderm . ( scale bars : a and c , 100 μm ; b , 50 μm . )
our data demonstrate that neurogenesis in onychophorans does largely not reflect the ancestral pattern of euarthropod neurogenesis . in all four euarthropod groups , neural precursors are generated at stereotyped positions despite the differences in neural precursor formation ( npgs versus neuroblasts ) . in each hemineuromere , a fixed set of ∼30 npgs / neuroblasts segregates in a pattern that is similar in all euarthropod representatives that have been analyzed ( 35 , 36 ) ( fig . s5 b–d ) . the conservation strongly suggests that this pattern has been present in the last common ancestor of euarthropods .
we conclude that the selection of neural precursors , as well as their arrangement and number , are conserved in all four euarthropod groups , suggesting that this pattern has been present in the last common ancestor of euarthropods , although onychophorans do not share these neural characters .
second , the formation of neural precursor groups in many areas of drosophila neurogenesis strongly suggests that npgs are part of the ancestral pattern of euarthropod neurogenesis and therefore cannot be used to resolve euarthropod relationships . the pars intercerebralis and pars lateralis , for example , are part of the central neuroendocrine system in the drosophila brain and derive from npgs that invaginate and become attached to the brain primordium ( 37 ) . additional examples are the stomatogastric nervous system , which is generated by three large npgs , the bilateral invaginations of the optic primordium and the generation of large precursor groups in the developing polyclonal sensory organs of drosophila ( 38 – 40 ) .
finally , based on the following data , we suggest that the exclusive generation of neural cells from the central area of the ventral neuroectoderm of chelicerates and myriapods represents an ancestral character of arthropod neurogenesis . first , in onychophorans , this mode of neurogenesis can be observed in the ventral protocerebral primordium . in this area , the whole neuroectoderm becomes internalized and forms vesicles that have been termed hypocerebral organs . neural precursors continue to segregate from the hypocerebral organs after their invagination ( 24 ) . second , in drosophila , the embryonic procephalic neuroectoderm that generates the larval brain neuromeres generates almost exclusively neural cells . the epidermis extends over the brain primordium during head involution ( 41 ) . furthermore , fundamental differences in the formation of epidermal cells in the ventral neuroectoderm of insects and malacostracans suggest that the capacity to generate epidermal and neural cells has evolved independently in insects and crustaceans . in insects , the cells that remain in the neuroectodermal layer after segregation of the neuroblasts develop into epidermis . in contrast , in malacostracans , neuroblasts remain in the neuroectoderm and can generate both neural and epidermal cells ( 13 , 28 ) .
based on the data presented here , we suggest the following evolutionary scenario of arthropod neurogenesis . in the last common ancestor of arthropods , the nervous system was generated by a massive irregular segregation of individual neural precursor cells ( fig . s6 ) . proneural genes might have been expressed homogeneously in the whole neurogenic area to confer neural potential to all cells . we speculate that the whole neurogenic area exclusively generated neural precursors , similar to the case in chelicerates , myriapods , and the brain primordia of drosophila and onychophorans .
cell proliferation might have occurred mainly in the neuroectoderm . individual neural precursors became postmitotic , segregated from the ventral neurectoderm and directly differentiated into neural cells . this hypothesis is supported by the nature of the neural precursors in chelicerates and myriapods . in both euarthropod groups , the npgs are postmitotic and directly differentiate into neural cells after their segregation . this ancestral state has been retained in the stomatogastric nervous system of drosophila melanogaster , which is generated by neural precursors that invaginate as large groups and directly differentiate into neurons ( 38 ) . the hypothetical ancestral state is further supported by recent data on neurogenesis in an annelid ( 42 ) . annelids belong to the lophotrochozans , which are a sister group to the ecdysozoa [ e . g . , arthropods , nematodes ( 43 ) ] . in the polychaete capitella sp . i , one to several adjacent neural precursors delaminate from the procephalic neuroectoderm . the neural precursors do not divide after their delamination ; rather , they directly differentiate into neural cells . thus , subsequent stages of neurogenesis must have diverged in the lineage leading to the onychophorans , such as mitotic divisions of the segregated neural precursor cells , the evolution of intermediate neural precursors , and the generation of epidermal and neural cells from the ventral neuroectoderm .
if we assume that the presented evolutionary sequence is correct , we would have to conclude that neuroblasts have gradually evolved from npgs . there is indeed evidence from intermediate states that supports this theory . in the centipede strigamia maritima , all cells of a npg are attached to a single cell of the group rather than to the apical surface as in chelicerates and millipedes ( 14 ) . this cell expresses higher levels of delta transcripts as compared with the remaining cells of the group . thus , delta / notch signaling seems to generate single cells with distinct properties within the npgs in strigamia , although the whole group eventually invaginates and gives rise to neural cells . furthermore , in the basal insect schistocerca americana , groups of four to eight cells are selected in the ventral neuroectoderm ( 12 ) . one cell of the group , the neuroblast , delaminates , whereas the remaining cells of the group differentiate into a cap cell and sheath cells that surround the neuroblast and its lineage .
our data on onychophoran neural precursor formation contradict neither the mandibulata nor the myriochelata hypothesis . however , we suggest that the invagination of npgs and the exclusive generation of neural cells in the ventral neuroectoderm of chelicerates and myriapods represent plesiomorphic characters that cannot be used to support the myriochelata hypothesis . furthermore , the deployment of onychophoran characters for outgroup comparison has to be considered carefully . crown - group euarthropods and probable stem - group onychophorans already existed in the early cambrian period ( 46 – 48 ) . hence , the onychophoran lineage must have had a long evolutionary history diverged from the euarthropod stem lineage . therefore , extant onychophorans might not show the ancestral state of neurogenesis without modifications . indeed , our data show that the development of the nervous system in onychophorans largely does not reflect the ancestral pattern of euarthropod neurogenesis ; rather , it seems to show a mixture of derived characters and ancestral characters that have been present in the last common ancestor of arthropods but have been modified in the euarthropod lineage .
for the detection of f - actin , we used alexa phalloidin 488 ( invitrogen ) ; 2 u dissolved in 10 μl methanol was evaporated to remove the methanol and then resuspended in 200 μl pbs and added to the embryos . after 1 h , the embryos were washed in pbs and used for imaging . to detect mitotic cells , we used the polyclonal rabbit anti - alpha phospo histone 3 antibody ( sigma ) diluted 1 : 200 times in pbs with 0 . 5 % bsa and 0 . 1 % triton ×100 . antibody incubation was done as described in eriksson and budd ( 50 ) .
we thank the members of the a . s . laboratory for continuous stimulating discussions and petra ungerer for critical comments on the manuscript . we are grateful to edina balczo for excellent technical support . we thank noel tait for help with collecting animals . this work was supported by the biotechnology and biological sciences research council .
author contributions : a . s . designed research ; b . j . e . performed research ; a . s . contributed new reagents / analytic tools ; b . j . e . and a . s . analyzed data ; and a . s . wrote the paper .
data deposition : the sequences reported in this paper have been deposited in the genbank database ( accession nos . ekash , gu954550 ; ekdelta , gu954551 ; and eknotch , gu954552 ) .
early events in insect neurogenesis . i . development and segmental differences in the pattern of neuronal precursor cells
development of paraperipatus amboinensis n . sp . ( entwicklung von paraperipatus amboinensis n . sp . )
formation and differentiation of the postnaupliar germ band of diastylis rathkei ( crustacea , cumacea ) ii . differentiation and pattern formation of the ectoderm ( translated from german )
studies on the onychophora . vii . the early embryonic stages of peripatopsis , and some general considerations concerning the morphology and phylogeny of the arthropoda
neurogenesis in an annelid : characterization of brain neural precursors in the polychaete capitella sp . i
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rupela sejuncta is a moth in the crambidae family .

it was described by heinrich in 1937 .

it is found in north america , where it has been recorded from alabama , florida , oklahoma and texas .

the wingspan is 28 – 33 mm for males and 25 – 30 mm for females .

the wings are shining white .

adults have been recorded on wing from march to july and from september to november .

rupela sejuncta

rupela sejuncta heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 373 , pl . 26 , f . 14 - 14c , pl . 32 , f . 39 ; tl : harris co . , texas
rupela nivea walker , 1863 ; list spec . lepid . insects colln br . mus . 28 : 524
rupela ( schoenobiinae ) ; heinrich , 1937 , proc . u . s . nat . mus . 84 ( 3019 ) : 356 ; [ nacl ] , 75 ; [ globiz ]
rupela adunca heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 374 , pl . 27 , f . 16 - 16c ; tl : bolivia
rupela bendis heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 378 , pl . 29 , f . 21 ; tl : aroa , venezuela
rupela faustina heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 380 , pl . 29 , f . 22 ; tl : cabima , panama
rupela jana heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 381 , pl . 31 , f . 38 ; tl : chaco , argentina
rupela lara heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 382 , pl . 32 , f . 40 ; tl : cabima , panama
rupela herie heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 381 , pl . 30 , f . 27 ; tl : georgetown , british guiana
rupela orbona heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 384 , pl . 31 , f . 35 ; tl : mackenzie , british guiana
rupela liberta heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 364 , pl . 23 , f . 4 - 4c ; tl : durango , mexico
rupela vexativa heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 371 , pl . 25 , f . 11 - 11c ; tl : quirigua , guatemala
rupela scitula heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 374 , pl . 26 , f . 15 - 15c ; tl : tucuman , argentina
rupela antonia heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 378 , pl . 30 , f . 31 ; tl : sixola river , costa rica
rupela drusilla heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 379 , pl . 29 , f . 24 ; tl : castro , paraná , brazil
rupela edusa heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 379 , pl . 30 , f . 29 ; tl : castro , paraná , brazil
rupela gaia heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 380 , pl . 30 , f . 28 ; tl : castro , paraná , brazil
rupela procula heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 384 , pl . 32 , f . 41 ; tl : sta . catherina , brazil
rupela lumaria heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 375 , pl . 27 , f . 17 - 17c ; tl : carillo , costa rica
rupela canens heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 379 , pl . 29 , f . 23 ; tl : sao paulo de olivenca , brazil
rupela labeosa heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 363 , pl . 22 , f . 3 - 3d ; tl : castro , paraná , brazil
rupela pallidula heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 365 , pl . 23 , f . 5 - 5d ; tl : castro , paraná , brazil
rupela saetigera heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 367 , pl . 24 , f . 8 - 8c ; tl : castro , paraná , brazil
rupela cornigera heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 371 , pl . 25 , f . 12 - 12b ; tl : castro , paraná , brazil
rupela imitativa heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 372 , pl . 26 , f . 13 - 13c ; tl : castro , paraná , brazil
rupela spinifera heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 377 , pl . 28 , f . 19 - 19c ; tl : castro , paraná , brazil
rupela monstrata heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 377 , pl . 28 , f . 20 - 20c ; tl : castro , paraná , brazil
rupela candace heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 382 , pl . 29 , f . 25 - 25a ; tl : castro , paraná , brazil
rupela nereis heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 383 , pl . 31 , f . 33 - 34 ; tl : castro , paraná , brazil
rupela albinella ; heinrich , 1937 , proc . u . s . nat . mus . 84 ( 3019 ) : 362 , pl . 22 , f . 2 - 2d , pl . 29 , f . 26 - 26b
rupela gibbera heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 367 , pl . 24 , f . 7 - 7d ; tl : moengo , boven , cortica river , surinam
rupela maenas heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 383 , pl . 31 , f . 36 - 37 ; tl : ponte nova ( rio xingu ) , brazil
rupela segrega heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 366 , pl . 23 , f . 6 - 6d , pl . 30 , f . 32 ; tl : south bay , lake okeechobee , florida
rupela horridula heinrich , 1937 ; proc . u . s . nat . mus . 84 ( 3019 ) : 376 , pl . 27 , f . 18 - 18f , pl . 33 , f . 47 ; tl : campo bello , rio de janeiro , brazil
rupela leucatea ; heinrich , 1937 , proc . u . s . nat . mus . 84 ( 3019 ) : 360 , pl . 22 , f . 1 - 1d , pl . 30 , f . 30 , pl . 33 , f . 44 - 45 , 48
rupela tinctella ; heinrich , 1937 , proc . u . s . nat . mus . 84 ( 3019 ) : 368 , pl . 24 , f . 9 - 9c , pl . 32 , f . 42 - 43 , pl . 33 , f . 46 ; [ nacl ] , # 5311
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a taxon identifier is composed of name , author , year and attribute , all separated by a blank . these are all extracted from the original publication .
the name is reproduced exactly as proposed in the original publication . the name of a genus is made up of one word and species made up of two words ( genus and species ) separated by a blank .
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= scirpophaga ; hampson , 1896 , proc . zool . soc . lond . 1895 : 912
puerto rico , jamaica , cuba , hispaniola , martinique , antiqua , mexico , guatemala , nicaragua , panama , honduras , trinidad , venezuela , guianas , brazil , argentina , paraguay , peru . see [ maps ]
larva on echinochloa polystachya heinrich , 1937 , proc . u . s . nat . mus . 84 ( 3019 ) : 361
mexico , british honduras , guatemala , honduras , costa rica , guianas , brazil , colombia , ecuador , peru . see [ maps ]
albinella ( stoll , [ 1781 ] ) ; in cramer , uitl . kapellen 4 ( 29 - 31 ) : pl . 372 , f . d ; tl : surinam
florida , mexico , cuba , guianas , trinidad , brazil ( paraná ) , paraguay , n . argentina . see [ maps ]
panama , brazil ( paraná , pará ) , argentina ( gran chaco , goya , villa ana ) . see [ maps ]
ruplea nivea ; heinrich , 1937 , proc . u . s . nat . mus . 84 ( 3019 ) : 370 , pl . 25 , f . 10 - 10c
brazil ( rio de janeiro ) , guianas , trinidad . see [ maps ]
panama , guianas , brazil , peru , paraguay , argentina ( chaco , goya , villa ana ) . see [ maps ]
this information is not automatically synchronized with globiz and can sometimes be lagging behind .
[ maps ] warning ! the maps are automatically generated from the textual information , and the process does not always produce acceptable result ; see about maps for more info .
zeller , 1863 chilonidarum et crambidarum genera et species chil . cramb . gen . spec . : 56pp
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the stone loach ( barbatula barbatula ) is a species of fresh water ray-finned fish in the balitoridae family .

it is one of seventeen species in the genus barbatula .

it is found in baltic states , eastern europe , austria , belgium , bulgaria , czech republic , denmark , finland , france , germany , hungary , ireland , italy , liechtenstein , luxembourg , moldova , the netherlands , poland , romania , serbia and montenegro , slovakia , slovenia , spain , sweden , switzerland , and the united kingdom .

stone loaches live amongst the gravel and stones of fast flowing water where they can search for food .

the most distinctive feature of this 14 cm fish is the presence of barbels around the bottom jaw , which they use to detect their invertebrate prey .

the body is a mixture of brown , green and yellow .

stone loach

the stone loach is classified as least concern ( lc ) on the iucn red list ( 1 ) .
information on the stone loach ( barbatula barbatula ) is currently being researched and written and will appear here shortly .
descriptive osteology of a newly described stone loach , oxynoemacheilus chomanicus ( kamangar et al . , 2014 ) ( cypriniformes , nemacheilidae )
along a stream , we investigated whether the abundance of stone loach ( barbatula barbatula , l . ) was related to the presence of brown trout ( salmo trutta , l . ) and instream habitat variables . first , a field survey was carried out where different habitat variables and the densities of both species were quantified and subjected to principal components analysis . then the abundance of stone loach was related to the scores of the retained axes ( eigenvalues > 1 ) . the abundance of stone loach was positively correlated to substrate particle size , amount of shade , temperature , discharge and current velocity , but negatively correlated to brown trout abundance . secondly , a month - long field enclosure experiment in a stream was performed to test for any negative effects of brown trout on stone loach growth . four treatments were used : intraspecific competition ( stone loach at double density ) , interspecific competition ( stone loach + small trout ) , predation ( stone loach + large trout ) and a control ( stone loach alone ) . the results showed that large trout tended to have negative effects on final stone loach biomass . the absence of a negative effect of large trout on resource density suggests that nonlethal effects rather than resource competition caused this trend .
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the effect of interaction between the stone loach noemacheilus barbatulus ( l . ) and the bullhead cottus gobio ( l . ) on prey and habitat selection
the effect of interaction between the stone loach noemacheilus barbatulus ( l . ) and the bullhead cottus gobio ( l . ) on prey and habitat selection | springerlink
do instream habitat variables and the abundance of brown trout salmo trutta ( l . ) affect the distribution and growth of stone loach , barbatula barbatula ( l . ) ?
@ article { c7137605 - fdeb - 4889 - bf4e - 7097b2f95108 , abstract = { along a stream , we investigated whether the abundance of stone loach ( barbatula barbatula , l . ) was related to the presence of brown trout ( salmo trutta , l . ) and instream habitat variables . first , a field survey was carried out where different habitat variables and the densities of both species were quantified and subjected to principal components analysis . then the abundance of stone loach was related to the scores of the retained axes ( eigenvalues & gt ; 1 ) . the abundance of stone loach was positively correlated to substrate particle size , amount of shade , temperature , discharge and current velocity , but negatively correlated to brown trout abundance . secondly , a month - long field enclosure experiment in a stream was performed to test for any negative effects of brown trout on stone loach growth . four treatments were used : intraspecific competition ( stone loach at double density ) , interspecific competition ( stone loach + small trout ) , predation ( stone loach + large trout ) and a control ( stone loach alone ) . the results showed that large trout tended to have negative effects on final stone loach biomass . the absence of a negative effect of large trout on resource density suggests that nonlethal effects rather than resource competition caused this trend . } , author = { nilsson , erika and persson , anders } , issn = { 0906 - 6691 } , language = { eng } , number = { 1 } , pages = { 40 - - 49 } , publisher = { wiley - blackwell } , series = { ecology of freshwater fish } , title = { do instream habitat variables and the abundance of brown trout salmo trutta ( l . ) affect the distribution and growth of stone loach , barbatula barbatula ( l . ) ? } , url = urltoken volume = { 14 } , year = { 2005 } , }
along a stream , we investigated whether the abundance of stone loach ( barbatula barbatula , l . ) was related to the presence of brown trout ( salmo trutta , l . ) and instream habitat variables . first , a field survey was carried out where different habitat variables and the densities of both species were quantified and subjected to principal components analysis . then the abundance of stone loach was related to the scores of the retained axes ( eigenvalues > 1 ) . the abundance of stone loach was positively correlated to substrate particle size , amount of shade , temperature , discharge and current velocity , but negatively correlated to brown trout abundance . secondly , a month - long field enclosure experiment in a stream was performed to test for any . . .
p . mafakheri , s . eagderi , h . farahmand , and h . mousavi - sabet , “osteological structure of kiabi loach (
p . mafakheri , s . eagderi , h . farahmand , and h . mousavi - sabet , “descriptive osteology of persian loach (
j . freyhof , f . erk ' akan , c . özeren , and a . perdices , “an overview of the western palaearctic loach genus
stone loach and bullhead were given a choice of chironomus and asellus prey in experiments using solitary fish and fish in company . solitary fish ate more than fish in company . the effect of light and substratum type on feeding was investigated . both species ate more prey items on gravel than silt when a significant difference was observed . bullhead ate more than loach in the light on both substrata . the only experiment in which loach ate more than bullhead was on a silt substratum in the dark . it is concluded that these two species utilise different components of the available food resources in chalk streams by adopting different habitats .
b . b . kamangar , a . m . prokofiev , e . ghaderi , and t . t . nalbant , “stone loaches of choman river system , kurdistan , iran ( teleostei :
appearance : a small , elongate , bottom - dwelling fish with three pairs of mouth barbels 3 - 5 mm in length . small eyes located almost on top of the head . body shape more cylindrical , forebody somewhat fatter than the spined loach , in which the body is laterally more flattened . tips of dorsal and caudal fins rounded , posterior margins may be slightly notched . fins of spined loach are more rounded .
topmouth gudgeon ( pseudorasbora parva ) is one of the most invasive aquatic fish species in europe and causes adverse effects to ecosystem structure and functioning . knowledge and understanding of the species’ interactions with the environment and with native fish are important to stop and prevent the further spread of the species . creating species distribution models is a useful technique to determine which factors influence the occurrence and abundance of a species . we applied three different modelling techniques : general additive models , random forests and fuzzy habitat suitability modelling ( fhsm ) to assess the habitat suitability of topmouth gudgeon . the former two techniques indicated that the abundance of native fish ( i . e . biotic variables ) was more important than environmental variables when determining the abundance of topmouth gudgeon in flanders ( belgium ) . bitterling ( rhodeus amarus ) , stone loach ( barbatula barbatula ) , three - spined stickleback ( gasterosteus aculeatus ) and predator abundance were selected as the most important biotic variables and implemented in the fhsm to investigate species interactions . depending on the preferred food source and spawning behaviour , either coexistence or interspecific competition can occur with bitterling , stone loach and three - spined stickleback . in contrast , the presence of predators clearly had a top down effect on topmouth gudgeon abundance . these findings could be applied as a biological control measure and implemented in conservation strategies in order to reduce the abundance of earlier established populations of topmouth gudgeon .
usually found in flowing stretches of streams and medium - sized rivers with gravel to stone bottom , but also in a variety of other habitats , including sandy canals and lake shores . larvae are benthic . larvae and small juveniles prefer sand bottom and slow current , shifting to gravel bottom and fast current when growing . adults prey on relatively large benthic invertebrates such as gammarids , chironomids , insect larvae . they breed on gravel , sand or among aquatic vegetation . tolerate moderate organic pollution and stream canalization and very sensitive to pollution by heavy metals ( ref . 59043 ) . sensitive to pollution and low oxygen levels , therefore , its presence in a river can be taken as an indication of good water quality ( ref . 6111 ) .
stone loaches , the family nemacheilidae , are small benthic fishes [ 1 ] found in fresh waters of asia and its islands , europe , and northeast africa [ 2 ] . this family have a great diversity in iranian inland waters [ 1 , 3 , 4 ] , with more than 40 reported species , including 24 endemic species [ 3 – 9 ] . they are less known due to small size and low economic value [ 10 , 11 ] and their classification is still complicated ; therefore , ichthyologists are trying to reveal their phylogenetic status [ 12 ] . given that , the osteological characteristics can play an important role in this regard [ 13 ] , since osteology is a useful tool to study the taxonomy and phylogenetic relationships among fishes [ 14 , 15 ] .
colouring : generally yellowish brown with darker bands and mottling . indistinct dark line joining eye to snout .
distribution and habitat : a freshwater species found throughout finland as far north as the lower reaches of the river kemijoki . avoids brackish water because of its salinity and occurs in the sea only in freshwater estuaries and in the easternmost part of the gulf of finland close to finland’s eastern border . inhabits stony bottoms in well - oxygenated water . uses its barbels to hunt for food , foraging under stones . unable to tolerate oxygen deficiency in eutrophic lakes but copes with eutrophication better in rivers because of the higher oxygen levels .
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often erroneously called nemacheilus barbatulus , cobitis barbatula or orthrias barbatula . several species are apparently confused under the name b . barbatula . recent molecular studies suggest that several species are lumped under this name and a taxonomic revision is needed to settle this problem .
justification : a widespread species with no known major widespread threats . european union 27 = lc . rationale same as above .
europe north of caucasus , pyrénées and alps , from loire and rhône drainages eastward ; british isles ( except northern scotland ) , southern sweden and finland ( northward to about 66°n ) ; danube and vardar drainages . several similar species in asia , as far as japan ( including
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department of fisheries , faculty of natural resources , university of tehran , p . o . box 4314 , karaj , iran
copyright © 2016 parvin mafakheri et al . this is an open access article distributed under the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original work is properly cited .
oxynoemacheilus chomanicus is a newly described species of the family nemacheilidae from the choman river drainage of the tigris basin . this study was conducted to provide the detailed osteological characteristics of this species and comparing them with those of other endemic species of the genus oxynoemacheilus from inland water basins of iran , namely , kiabii , o . persa , o . brandtii , o . kermanshahensis , and o . bergianus . for this purpose , nine specimens of o . chomanicus were collected , cleaned , and stained for osteological examination . then , a detailed description of their skeletal structure was provided . the results showed that o . chomanicus can be distinguished from other studied species of the genus oxynoemacheilus due to possessing an orbital shelf , number of the vertebrae , number of the hypurals , number of the unbranched rays in anal fin , features of the hemal and neural spines , and connection pattern of the parietal and frontal .
the genus oxynoemacheilus is a species rich genus of the family nemacheilidae known from albania eastwards to central iran [ 16 ] and has 11 reported species from iran of which three include o . chomanicus , o . kurdestanicus , and o . zagrodensesis recently described from the choman river drainage of the tigris basin [ 4 ] . the newly described species of o . chomanicus is found in the choman river drainage ( baneh , kurdistan province ) . identification of this species is based on the morphological features that show many similarities to other members of this genus [ 4 ] .
regarding high diversity and morphological similarity of the members of the family nemacheilidae , using combined data , including osteological and molecular data , can help to better understand their taxonomical relationship . therefore , this study was conducted to provide detailed osteological characteristics of o . chomanicus and comparing them with those of other species of the genus oxynoemacheilus from the inland water basins of iran that their osteological data are available . the finding of this study can provide the osteological features of this species that can help to discrete this species from others and be used as a basis for further phylogenetic study of the members of this genus based on the osteological data .
nine specimens of o . chomanicus with standard length of ( mean ± sd ) were collected using electrofishing device from the choman river ( kurdistan province , iran ) . then , specimens were anesthetized with 1 % clove solution and fixed in 4 % formaldehyde . for osteological examination , the specimens were cleared and stained using alcian blue and alizarin red based on taylor and van dyke [ 17 ] . pictures of the stained skeletal structures were obtained using an epson v600 scanner equipped with a glycerol bath . the skeletal structures of each sample were observed and studied by an ms5 leica stereomicroscope . the scanned images were illustrated by coreldraw x6 software . nomenclature and abbreviation of the skeletal elements follow prokofiev [ 12 , 18 ] . detailed descriptions of the osteological features of o . kiabii , o . bergianus , o . persa , o . brandtii , and o . kermanshahensis were provided by mafakheri et al . [ 19 ] , jalili and eagderi [ 20 ] , mafakheri et al . [ 21 ] , and mafakheri [ 22 ] , respectively .
the anterior part of the neurocranium is narrow with wider posterior part and its maximum width is about 60 % of its length . the ethmoid region consists of the paired lateral ethmoids and unpaired supraethmoid - ethmoid and prevomer . the elongated supraethmoid - ethmoid bone is vertically fused to the prevomer and posteriorly sutured with the frontal ( figures 1 ( a ) and 1 ( b ) ) . the anterior part of the prevomer is heart - shaped ( figure 1 ( b ) ) and is posteriorly connected to the orbitosphenoid ( figure 1 ( c ) ) . the lateral ethmoid is medially attached to the orbitosphenoid ( figure 1 ) . the anterior part of the lateral ethmoid is projected and its posterior part is convex in shape ( figure 1 ) .
figure 1 : ( a ) dorsal , ( b ) lateral , and ( c ) ventral views of the neurocranium of oxynoemacheilus chomanicus . 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 .
the orbital region includes the paired frontal , pterosphenoid , and sclerotic , and the unpaired orbitosphenoid and parasphenoid . the frontal is the largest bony element of the neurocranium’s roof . this bone is laterally connected to the orbitosphenoid , pterosphenoid , and sphenotic and posteriorly to the parietal ( figures 1 ( a ) and 1 ( b ) ) . the frontal is connected to the lateral ethmoid via the orbital shelf ( figure 1 ( a ) ) . the two frontals are connected medially but they are separated posteriorly and form the anterior part of the fontanelle ( figure 1 ( a ) ) . the orbitosphenoid is posteriorly connected to the heavy bone of the pterosphenoid . in the ventral plan , the parasphenoid is connected to the prevomer , orbitosphenoid , pterosphenoid , prootic , and basioccipital ( figure 1 ( c ) ) . the parasphenoid , orbitosphenoid , and pterosphenoid enclose the orbital foramen ( figure 1 ( b ) ) . the parasphenoid meets the prootic and pterosphenoid via its lateral wings and form a large foramen in ventral plan of the neurocranium ( figure 1 ( c ) ) . two small bones of the sclerotic enclose the orbits .
the otic region consists of the parietals , epiotics , pterotics , prootics , and sphenotics ( figure 1 ) . the parietal is located between the frontal and supraoccipital ( figure 1 ( a ) ) . this bone is anterolaterally connected to the sphenotic and posterolaterally to the epiotic ( figures 1 ( a ) and 1 ( b ) ) . the pterotic is almost triangle in shape that it locates between the sphenotic and epiotic dorsally , and the prootic and exoccipital ventrally ( figures 1 ( a ) and 1 ( c ) ) . connection of the epiotic and sphenotic avoids the relationship between the pterotic and parietal ( figure 1 ( b ) ) . the pterosphenoid , sphenotic , and prootic form the anterior articulatory facet for articulating the anterior condyle of the hyomandibular and the sphenotic and pterotic form the posterior articulatory facet for articulating the posterior condyle of the hyomandibular ( figures 2 ( b ) and 2 ( c ) ) .
figure 2 : ( a ) anterior part of the ethmoid region and ( b ) lateral plan of kinethmoid and autopalatine of oxynoemacheilus chomanicus . apl : autopalatine ; ke : kinethmoid ; mx : maxilla ; pe : prevomer ; peth ii : preethmoid ii ; pmx : premaxilla ; ppl : prepalatine ; ses : sesamoid .
the occipital region consists of both unpaired supraoccipital and basioccipital and the paired exoccipital ( figure 1 ) . the supraoccipital forms the posterior part of the fontanelle . this bone is connected to the sphenotic by its anterolateral wings and posteriorly to the exoccipitals ( figure 1 ( a ) ) . the exoccipital bears a large foramen and it is connected to the prootic anteroventrally ( figure 1 ( c ) ) . the basioccipital is posteriorly narrower having a ring - shaped process ( figure 1 ( c ) ) . this bone is situated between the exoccipitals and connected to the prootics anteriorly ( figure 1 ( c ) ) . the basioccipital is posteriorly articulated to the first centrum . in a ligamentous space anterior to the ethmoid region , the paired preethmoid ii , prepalatine , and sesamoid and unpaired kinethmoid are present ( figure 2 ) . the prepalatine is located between the autopalatine and maxilla . the prepalatine is medially connected to the preethmoid ii ( figure 2 ( a ) ) . the rod - shaped preethmoid ii is anteriorly connected to the maxilla and posteriorly to the prevomer . the cylindrically shaped kinethmoid is vertically located between jaws ( figure 2 ( a ) ) .
the upper jaw consists of the premaxilla and maxilla ( figure 3 ) . the premaxilla has two processes , that is , pr . ascenden and pr . alveolar . the premaxilla is ligamentously connected to the kinethmoid . the maxilla is a large laminar bone which has a small anteroventral process connecting to the preethmoid ii and prepalatine and an anteroventral process that is tilted downwardly reaching its counterpart ( figure 2 ( a ) ) . the lower jaw is composed of the dental , articular , retroarticular , and coronomeckelian . the large dental has two processes , including narrow ramus dentalis and wide coronoid processes . the articular is overlapped with the dental and posteriorly articulated to the quadrate . the small bone of the coronomeckelian is attached to the medial face of the articular . the small bone of the retroarticular is articulated to the posteroventral part of the articular .
figure 3 : ( a ) upper jaw and ( b ) medial view of lower jaw in oxynoemacheilus chomanicus art : articulare ; cm : coronomeckelian ; den : dental ; mx : maxilla ; pmx : premaxillary ; rar : retroarticular .
the branchial arch consists of the basibranchials , hypobranchials , ceratobranchials , epibranchials , and infrapharyngobranchials ( figure 4 ) . there are four basibranchials , which locate between the hypobranchials . the two first basibranchials are similar in shape . the hypobranchials are three pairs and almost square - shaped . in each side , there are five ceratobranchials of which the last one has different shape possessing pharyngeal teeth ( figure 4 ) . the four paired epibranchials situate between the rectangularly shaped ceratobranchials and the small infrapharyngobranchials . the infrapharingobranchials are horizontally arranged on the epibranchials .
figure 4 : branchial apparatus of oxynoemacheilus chomanicus . bbr : basibranchial ; cbr : ceratobranchial ( cbr5 : pharyngeal bones ) ; ebr : epibranchial ; hbr : hypobranchial ; pbr : infrapharyngobranchials .
the hyoid arch is composed of the basihyal , dorsal hypohyal , ventral hypohyal , ceratohyal , epihyal , interhyal , urohyal , extra urohyal , and branchiostegals ( figure 5 ) . the large cylindrical ceratohyal intervenes between the hypohyals and the epihyal ( figure 5 ) . the epihyal is triangular in shape . a small rod - shaped interhyal fits in a small groove on the posterodorsal part of the epihyal ventrally . the dorsal hypohyal and ventral hypohyal are connected firmly . two small rod - shaped extra urohyals are located between the hypohyals and ligamentously connected to the urohyal and basihyal . the laminar bone of the urohyal bears a depression anteriorly and wings anterolaterally ( figure 5 ) . the unpaired basihyal is t - shaped . there are three curved branchiostegals of which the first one is attached to the ceratohyal , the connection point of the second one is located between ceratohyal and epihyal , and the third one is connected to the epihyal .
figure 5 : hyoid arch of oxynoemacheilus chomanicus . bhy : basihyal ; br : branchiostegal ; chy : ceratohyal ; dhy and vhy : dorsal and ventral hypohyal ; ehy : epihyal ; ihy : interhyal ; uhy : urohyal ; uhye : extra urohyal .
the suspensorium consists of the autopalatine , endopterygoid , ectopterygoid , metapterygoid , hyomandibular , quadrate , and symplectic ( figure 6 ) . the autopalatine is a heavy bone articulating to the endopterygoid posteriorly ( figure 2 ( b ) ) . there is a strong ligamentous attachment between the endopterygoid , ectopterygoid , metapterygoid , and quadrate . the triangle - shaped bone of the symplectic is inserted into the depression of the quadrate . a condyle for articulation to the articular presents in the anteroventral part of the quadrat . the hyomandibular bears two articular heads dorsally for the craniohyomandibular articulations .
figure 6 : medial view of the suspensorium of oxynoemacheilus chomanicus . apl : autopalatine ; ect : ectopterygoid ; end : endopterygoid ; hm : hyomandibular ; io : interopercle ; mtp : metapterygoid ; op : opercle ; po : preopercle ; q : quadrate ; so : subopercle ; sym : symplectic .
this series consists of the opercle , subopercle , interopercle , and preopercle ( figure 6 ) . the opercle is the largest element of the opercular series and its dorsal margin is concaved . this bone has a large condyle , which is articulated to the hyomandibular . the opercle covers the paddle - shaped subopercle ventrally ( figure 7 ) . the narrow curved preopercle almost locates at the rear of the interopercle ( figure 6 ) . the interopercle is not completely ossified and connected to the subopercle posteriorly ( figure 6 ) .
figure 7 : medial views of the pectoral girdle of oxynoemacheilus chomanicus . cl : cleithrum ; cor : coracoid ; mcor : mesocoracoid ; ptt : posttemporal ; rad : ossified pectoral radial ; sc : scapula ; scl : supracleithrum ; stt : supratemporal .
the pectoral girdle includes the cleithrum , coracoid , scapula , mesocoracoid , supratemporal , posttemporal , supracleithrum , and radials ( figure 7 ) . the cleithrum is the largest bony elements of the pectoral girdle possessing the vertical and horizontal sections . its vertical section is connected to the supracleithrum , whereas the horizontal one is connected to the scapula and coracoid . the coracoid is narrow anteriorly . the scapula is located between the cleithrum and coracoid . there is a large foramen on its middle face . the mesocoracoid is a small rod - shaped bone , which is medially located between the cleithrum and coracoid . the mesocoracoid indirectly connects these two bones ( figure 7 ) . the supracleithrum bears a ridge on its lateral face . the elongated posttemporal is situated in the anterolateral depression of the supracleithrum . this bone is posterolaterally fused to the supratemporal . the supratemporal is a small tubular - shaped bone . there are four radials of which first two ones are narrow and elongated and the most medial one is the smallest and widest one .
the pelvic girdle consists of the paired pelvic bones and radials . the pelvic bones are horizontally located in belly area . the pelvic bones are firmly connected by a strong ligament medially . they have posterolateral processes , that is , pr . iliacus that along with the posterior process , that is , pr . ischiadicus , forms a depression for positioning the rays and radials . the pelvic bone is anteriorly narrow ( pubic process ) . there are three small radials . there are one unbranched ray , seven branched rays , and one small curved pelvic splint in the pelvic fin .
there are 37 centra of which first four centra form the weberian apparatus and bony capsule . the hemal and neural spines are narrow . the dorsal fin has 9 pterygiophores and one stay and the first pterygiophore is the largest reaching the 11th or 12th vertebra . there are five unbranched and 8 1 / 2 branched rays in dorsal fin ( figure 8 ( a ) ) . the number of the pterygiophores in anal fin is six with one stay in which first pterygiophore reaches the 24th centrum . the number of unbranched and branched rays in anal fin is 4 and 5 1 / 2 , respectively ( figure 8 ( b ) ) .
figure 8 : ( a ) dorsal fin and ( b ) anal fin of oxynoemacheilus chomanicus . dr : distal radial ; mr : medial radial ; pr : proximal radial ; sty : stay .
the caudal complex consists of the second preural centrum and its neural and hemal processes , first preural centrum , ural - 1 + ural - 2 , pleurostyle , hypurals , parhypural , epural , and principal caudal rays , and procurrent rays ( figure 9 ) . the hemal process of the second preural is articulated to its centrum and its neural process is fused to its centrum . the last centrum is formed by the fused ural - 1 , ural - 2 , and preural - 1 . the wide rectangularly shaped parhypural along with the hypural - 1 is articulated to the last centrum . the parhypural is dorsally connected to the hypural - 1 . there are six hypurals of which first two ones are large . the hypural - 2 is fused to the centrum and others are inserted into the groove of the pleurostyle . the last hypural is small . the small epural is located between the pleurostyle and neural rudimentary . the pleurostyle is elongated and fused to the centrum . the principal caudal rays are supported by the hypurals .
figure 9 : caudal skeleton of oxynoemacheilus chomanicus . epu : epural ; hp : hypural ; hpu2 : hemal process of the second preural centrum ; npu - 2 : neural process of the second preural centrum ; ph : parhypural ; ust : pleurostyle .
the members of the genus oxynoemacheilus , with 11 reported species , are distributed in the most of the inland waters basins of iran and their identification is difficult based on their external morphology . hence , the osteological features are suitable characteristics for their distinction [ 12 , 18 ] . in addition , the osteological data can be well interpreted in taxonomy and phylogenetic relationships of fish species even in higher level [ 23 ] . there are various works on the phylogenetic relationships of the superfamily cobitoidea using osteological characteristics containing important information [ 12 , 13 , 18 , 24 – 27 ] . therefore , the present study provided detailed osteological characteristics of o . chomanicus for further taxonomical studies of these taxa .
prokofiev [ 12 ] has pointed out that the coronomeckelian is located on the base and dorsal edge of the coronoid processes in the genus oxynoemacheilus , whereas it is attached to the articular in o . chomanicus as mentioned by sawada [ 13 ] . in addition , prokofiev [ 12 ] mentioned that the lateral ethmoid is stationarily jointed with the supraethmoid - ethmoid , while such a feature was not observed in o . chomanicus . in o . chomanicus , similar to other compared species of this genus , the prootic , pterosphenoid , and sphenotic form the anterior articulatory facet , and the pterotic and sphenotic form the posterior articulatory facet , whereas prokofiev [ 12 ] mentioned that , in the members of the genus oxynoemacheilus , the anterior articulatory facet is restricted by the sphenotic and prootic , and the posterior articulatory facet is restricted by the sphenotic , prootic , and pterotic .
based on the results , this species shows some osteological features that can be considered as its osteological identification key in comparison to o . kiabii [ 19 ] , o . persa [ 21 ] , o . brandtii , o . kermanshahensis [ 22 ] , and o . bergianus [ 20 ] . there is an orbital shelf of the frontal in o . chomanicus , whereas such shelf is not present in o . persa [ 21 ] . the posterior process of the orbitosphenoid in o . persa is well developed [ 21 ] unlike that of o . chomanicus .
in o . kiabii , the supraethmoid - ethmoid is shorter and wider [ 19 ] than that of o . chomanicus . the connection pattern of the frontal and parietal in o . kiabii had a zigzag pattern [ 19 ] versus straight one of o . chomanicus . furthermore , there is a small bone between the frontal and parietal in o . persa [ 21 ] . the shape of the metapterygoid in o . chomanicus is rectangular , while it is square - shaped in o . kiabii [ 19 ] . furthermore , the hypohyals’ processes of o . kiabii are well developed [ 19 ] unlike that of o . chomanicus . in o . brandtii and o . bergianus , the hemal and neural spines are wider and well developed [ 20 , 22 ] , while those of o . chomanicus are thin and narrow similar to o . kiabii , o . persa , and o . bergianus .
oxynoemacheilus chomanicus has six hypurals , whereas o . kiabii , o . brandtii , o . persa , and o . kermanshahensis have five hypurals . the number of the vertebrae in o . chomanicus is 37 , whereas o . persa and o . kermanshahensis have 34 - 35 and 39 - 40 vertebrae , respectively [ 21 , 22 ] . the dorsal margin of the maxilla in o . chomanicus is smooth , whereas it is convex - shaped in o . kermanshahensis [ 22 ] .
in o . chomanicus , the number of unbranched rays in dorsal and anal fins is five and four , respectively , which is different from the original description of this species ( three and three ) by kamangar et al . [ 4 ] . these extra unbranched rays were visible only in cleared and stained specimens . the number of branched rays in dorsal and anal fins of o . chomanicus is 8 1 / 2 and 5 1 / 2 , respectively , corresponding to description of this species by kamangar et al . [ 4 ] . the branched rays of the dorsal fin in o . chomanicus are more than those of o . kermanshahensis [ 19 ] .
some species of the genus oxynoemacheilus were previously placed in other genera , including barbatula and triplophysa , but osteological investigations revealed their differences and ascribed them to related genera [ 12 , 13 ] . of the most important differences of the genus oxynoemacheilus with the members of the genera barbatula and triplophysa are lack of the preethmoid - i and the number and shape of the radials of the pectoral fin . in the examined species , including o . persa , o . bergianus , o . brandtii , o . kermanshahensis , o . kiabii , and o . chomanicus , the preethmoid - i is absent . the numbers of the radials in the pectoral fins of triplophysa is 3 and barbatula is 4 with different shape compared to the members of the genus oxynoemacheilus . the members of the genus oxynoemacheilus bear two small sesamoid bones , whereas this bone is absent in triplophysa and barbatula [ 12 , 13 ] . in oxynoemacheilus , the number of the branched rays of the anal fin is 5 , whereas this number can be higher in barbatula and triplophysa .
based on the results , the presence of the orbital shelf , number of the vertebrae , number of the hypurals , number of the unbranched rays in anal fin , features of the hemal and neural spines , and connection pattern of the parietal and frontal can be offered as identified keys for o . chomanicus from other examined species of the genus oxynoemacheilus from inland water of iran .
the authors declare that there is no conflict of interests regarding the publication of this paper .
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j . freyhof , g . sayyadzadeh , h . r . esmaeili , and m . geiger , “review of the genus
h . mousavi - sabet , g . sayyadzadeh , h . r . esmaeili , s . eagderi , r . patimar , and j . freyhof , “
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p . jalili , s . eagderi , n . nikmehr , and y . keivany , “descriptive osteology of
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phylogenetic of iranian genus oxynoemacheilus ( nemacheilidae ) using morphological characters [ m . s . thesis ]
l . s . ramaswami , “skeleton of cyprinoid fishes in relation to phylogenetic studies . v . the skull and the gasbladder capsule of the cobitidae , ”
n . c . bird and l . p . hernandez , “morphological variation in the weberian apparatus of cypriniformes , ”
freshwater ; demersal ; ph range : 7 . 0 - 7 . 7 ; dh range : 10 - 15 ; potamodromous ( ref . 51243 ) . temperate ; ? - 18°c ( ref . 13371 ) ; 67°n - 39°n , 6°w - 59°e
eurasia : europe north of caucasus , pyrénées and alps , from loire and rhone drainages eastward ; british isles ( except northern scotland ) , southern sweden and finland ( northward to about 66°n ) ; northeastern italy ; danube and vardar drainages ( ref . 59043 ) ; asia to china ( ref . 6111 ) .
maturity : l m ? range ? - ? cm max length : 21 . 0 cm tl male / unsexed ; ( ref . 1441 ) ; common length : 12 . 0 cm sl male / unsexed ; ( ref . 5504 ) ; max . published weight : 200 . 00 g ( ref . 5504 ) ; max . reported age : 7 years ( ref . 6111 )
dorsal spines ( total ) : 3 ; dorsal soft rays ( total ) : 6 - 8 ; anal spines : 3 ; anal soft rays : 5 - 6 . distinguished from its congeners in europe by the following combination of characters : caudal fin usually slightly emarginate ( truncate in a few populations ) ; pelvic origin beneath dorsal origin or under branched dorsal rays 1 - 2 ; caudal peduncle depth 1 . 4 - 2 . 2 ( usually 1 . 6 - 2 . 0 ) times in its length , 1 . 2 - 1 . 8 times in body depth ; often lacking dark blotches along back between nape and dorsal ( ref . 59043 ) . body elongated , anteriorly somewhat depressed , posteriorly laterally compressed . three pairs of mouth barbels . no erectile spine below eye . posterior margin of caudal fin slightly notched . caudal fin with 15 - 17 rays ( ref . 2196 ) .
spawns once a year for several years in low productivity streams , but exhibits multiple spawning within a season in high productivity environments ( ref . 40290 , 40756 ) . releases eggs in open open water , often close to surface . eggs drift and adhere to different substrates and are often covered by sand or detritus . individual females may spawn daily for a short period ( ref . 59043 ) .
kottelat , m . and j . freyhof , 2007 . handbook of european freshwater fishes . publications kottelat , cornol and freyhof , berlin . 646 pp . ( ref . 59043 )
phylogenetic diversity index ( ref . 82805 ) : pd 50 = 0 . 5000 [ uniqueness , from 0 . 5 = low to 2 . 0 = high ] .
bayesian length - weight : a = 0 . 01000 ( 0 . 00386 - 0 . 02592 ) , b = 3 . 04 ( 2 . 82 - 3 . 26 ) , in cm total length , based on lwr estimates for this ( sub ) family - body shape ( ref . 93245 ) .
trophic level ( ref . 69278 ) : 3 . 3 ±0 . 0 se ; based on diet studies .
resilience ( ref . 69278 ) : medium , minimum population doubling time 1 . 4 - 4 . 4 years ( tm = 2 - 3 ) .
prior r = 0 . 69 , 2 sd range = 0 . 35 - 1 . 38 , log ( r ) = - 0 . 37 , sd log ( r ) = 0 . 34 , based on : 2 k , 8 tgen , 1 tmax , 1 fec records
vulnerability ( ref . 59153 ) : moderate vulnerability ( 37 of 100 ) .
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the general appearance is of a slim , long bodied pale brown fish dappled with darker patches of brown . it has six barbels on the mouth .
very clean , unpolluted stony streams . it will not tolerate even mildly polluted waters .
spawns once a year in low productivity streams , but exhibits multiple spawning within a season in high productivity environments . eggs are a dull white and laid among stones and water plants .
widespread and fairly frequent in all of britain except in the north of scotland .
fairly common in leicestershire and rutland . widely distributed in smaller rivers and streams . including the swift , rothley brook , gaddesby brook , wreake , mease / gilwiskaw , eastern and western sence , soar , chater and charnwood streams .
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robins , richard c . , reeve m . bailey , carl e . bond , james r . brooker , ernest a . lachner , et al .
disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .
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azimi , hoda ; mousavi - sabet , hamed ; eagderi , soheil ; vatandoust , saber .
international journal of aquatic biology ; karaj vol . 3 , iss . 5 , ( oct 2015 ) : 290 - 300 .
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pieterjan verhelst is a recipient of a ph . d . grant financed by the agency for innovation by science and technology in flanders ( iwt ) and is affiliated with ghent university and the research institute for nature and forest ( inbo ) . pieter boets was supported by a postdoctoral fellowship from the research foundation flanders ( fwo - vlaanderen ) . this work was supported by the interreg iia two seas project rinse . the author would like to thank the fishing team of inbo groenendaal and tom de boeck for data extraction .
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