lepidocottus is a genus of goby that lived in what is now southern france in the uppermost oligocene , 24 to 23 million years ago .

lepidocottus

† lepidocottus gracilis laube from freshwater sediments of the lower oligocene of eastern germany [ 78 ] , [ 79 ] .
† lepidocottus aries is additionally known from oligocene strata near martigues , bouches - du - rhône , france [ 57 ] .
1875 lepidocottus aries ( agassiz ) . – sauvage , notes sur les poissons fossiles , p . 635–637 , pl . 23 : fig . 1 .
a single marine † lepidocottus species has been described from the coastal lower miocene ( aquitanian ) site la paillade in southern france ( see [ 31 ] , as “genus eleotridarum” sectus ] . this species probably represents † l . martinii , rather than † l . sectus . this record represents the only known case in which † lepidocottus co - occurs with numerous gobiidae [ 31 ] . evidence of † lepidocottus younger than early miocene ( aquitanian ) remains unknown to date .
the here presented description of the cranial and postcranial skeleton , otolith , and scales of † lepidocottus aries , type species of † lepidocottus , demonstrates that several of the characters are present that define the gobioidei ( [ 6 ] , [ 11 ] , [ 50 ] , [ 51 ] , see also [ 56 ] ) . they include :
† lepidocottus sectus ( stinton & kissling ) from upper oligocene and lower miocene brackish and freshwater deposits in the western paratethys ( see [ 82 ] , as “genus eleotridarum” ) .
the extant butidae consist of 11 genera living in freshwater and estuarine habitats of the tropical indo - and w pacific , africa , asia and oceania [ 7 ] , [ 55 ] ( table 2 ) . all these genera display at least one important morphological feature that discriminates them from † lepidocottus ( table 2 ) . the extant genera of the eleotridae of which otoliths are known possess otoliths with a centered sulcus and thus cannot be identical with † lepidocottus . as a result , † lepidocottus is not a synonym of any extant butid or eleotrid and represents a valid genus name .
we hypothesize a phylogenetic setting that uses the presence of 15 branched caudal fin rays , quadratic otoliths and a centered sulcus as plesiomorphic characters ( fig . 8 ) . the differentiation of the otolith shape is considered a synapomorphy for the butidae . the trapezoid otolith shape is defining the butis lineage , while rectangular otoliths are interpreted as a synapomorphy for a clade that comprises † lepidocottus and the extant butidae , with the exception of butis . the differentiation of the sulcus , i . e . appearance of an anteriorly extended sulcus , is a synapomorphy for † lepidocottus and the extant butidae , except butis and oxyeleotris . secondary cycloid predorsal scales characterize a clade consisting of † lepidocottus , kribia and bostrychus , with ophiocara as sister . in addition , † lepidocottus is interpreted as sister to kribia based on the reduced number of caudal fin rays ( 13 vs . 15 in other butids ) .
thus , † lepidocottus clearly belongs to the gobioidei , and we can use our new data for the rare opportunity to incorporate a variety of characters from an exceptionally preserved fossil to scrutinize its phylogenetic position among an extant group of fishes .
overview of the extant genera of the butidae with number of species , zoogeographic distribution and habitats ( after [ 7 ] , [ 49 ] , [ 52 ] , [ 55 ] ) and selected morphological differences that separate them from †lepidocottus .
overview of the extant genera of the butidae with number of species , zoogeographic distribution and habitats ( after [ 7 ] , [ 49 ] , [ 52 ] , [ 55 ] ) and selected morphological differences that separate them from † lepidocottus .
gaudant j . 2015 . — présence du genre lepidocottus sauvage , 1875 ( teleostei , gobioidei ) dans l’oligocène inférieur des environs de céreste ( alpes - de - haute - provence , france ) . geodiversitas 37 ( 2 ) : 229 - 235 . urltoken
in addition , three otolith - based species , in previous studies described as “ gobius ” and / or “genus eleotridarum” ( in the old definition , i . e . including the present - day eleotridae and butidae ) can now be identified as belonging to † lepidocottus :
overview of the extant genera of the butidae with number of species , zoogeographic distribution and habitats ( after [ 7 ] , [ 49 ] , [ 52 ] , [ 55 ] ) and selected morphological differences that separate them from † < i > lepidocottus < / i > .
résumé des squelettes en connexion de poissons gobioidei provenant de la formation de campagne - calavon , d ' âge rupélien ( = stampien ) sont décrits dans deux gisements des environs de céreste ( alpes de haute - provence ) . ils partagent avec les eleotridae et les butidae actuels plusieurs caractères anatomiques . ces poissons , qui sont étroitement apparentés à l ' espèce lepidocottus aries ( agassiz , 1839 ) , de l ' oligocène terminal saumâtre d ' aix - en - provence , en diffèrent cependant légèrement par la composition de leurs nageoires dorsale postérieure et anale . c ' est pourquoi ils sont désignés comme lepidocottus cf . aries ( agassiz , 1839 ) . leur présence dans ces sédiments lacustres s ' accorde parfaitement avec le mode de vie des eleotridae et des butidae actuels .
the studied specimens are determined as † lepidocottus aries ( agassiz ) , because they largely correspond to the original description of this species by agassiz [ 34 ] ( p . 12 , 186–187 ) and were found at the type locality ( aix - en - provence ) . in addition , our specimens correspond well to specimens determined as gobius aries from previous collections at aix - en - provence ( nmw 1910 - v12 , hlmd smff - 356 ) .
according to our results , † lepidocottus cannot be assigned to the rhyacichthyidae because the type species of this family , rhyacichthys aspro ( valenciennes ) , is clearly different from † l . aries . according to hoese [ 6 ] and hoese and gill [ 7 ] , r . aspro possesses a lateral line on the body ( absent in † l . aries ) , three epurals ( two in † l . aries ) , and scales with several rows of transforming ( = distally truncated ) ctenii ( one row of ctenii and no transforming ctenii in † l . aries ) . the otolith morphology of r . aspro is unknown .
the caudal endoskeleton , usually a significant source of diagnostic characters in fish taxonomy , may display apomorphies that help in correctly identifying butid or eleotrid genera [ 7 ] , [ 51 ] , [ 65 ] , but it does not provide any diagnostic feature that can be interpreted as a synapomorphy at the family level [ 7 ] . the presence of two epurals and one parhypural , as in † lepidocottus aries , occurs in several butid and eleotrid taxa [ 7 ] . the number of branched and segmented caudal rays is 15 in butidae , with the exception of kribia , which has 11–13 branched and segmented caudal rays [ 7 ] . eleotridae usually possess 13 branched and segmented caudal rays ( precisely as in kribia and † l . aries ) .
we have compared the otoliths of † lepidocottus aries with the otoliths of those extant butids and eleotrids of which data on otolith morphology are available ( figs . 6 , 7 ) . clearly , the rectangular otolith shape and the anteriorly extended sulcus ( with the ostium tip very close to the anterior otolith margin ) of † l . aries are also present in otoliths of several butidae ( figs . 6b , 7 ; see also [ 30 ] : pl . 4 , [ 65 ] : figs . 11 , 12 ) . in contrast , otoliths of eleotridae do not show these characters ; they are usually quadratic or higher than long ( exceptions occur in eleotris ) and their sulcus consistently is centered ( fig . 6c , see also [ 65 ] ) .
agassiz has indicated that his new species cottus aries is figured on plate 18 ; however , this plate was never printed and thus no figure of the holotype exists ( see also [ 35 ] : 635 ) . our efforts to find the holotype in the natural museum of history , paris yielded no success , and thus the holotype has to be considered as lost . sauvage [ 35 ] considered , like agassiz , cottus aries as a member of the cottidae , but recognized that it is different from the extant cottus linnaeus and therefore introduced the new genus name † lepidocottus for it . sauvage [ 35 ] provided a description and also a figure of † l . aries ( agassiz ) based on two newly collected specimens from aix that he had received from a private collector , but also these specimens are apparently lost .
characters of the cranium of † lepidocottus aries are the presence of an infraorbital bone ( fig . 4b ) and irregular ridges along the orbital margins of the frontal bones ( fig . 4c 1 ) . considering the butidae and eleotridae , the only taxa with which † l . aries shares an infraorbital bone are bostrychus and oxyeleotris ( both belong to the butidae ) , and the only taxon with which † l . aries shares the irregular ridges on the frontals is butis ( data from [ 52 ] ) . it is possible that these two characters represent a synapomorphy of the family butidae ( vs . eleotridae ) because both characters are absent in the eleotridae ( at least what can be said based on the available data ) . if this is correct , then the presence of these characters in † l . aries is indicative of affinities with the butidae , and the absence of these features in certain butids represents a secondary loss . however , a more complete data set on these characters in extant eleotridae is necessary to rule out the possibility that the infraorbital bone and irregular ridges along the orbital margins of the frontals represent homoplasies .
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the classification of gobioid fishes is still under discussion . several lineages , including the eleotridae and butidae , remain difficult to characterize because synapomorphies are rare ( eleotridae ) or have not yet been determined ( butidae ) . moreover , the fossil record of these groups is scarce .
the butidae and eleotridae are currently distributed in w - africa , madagascar , asia and australia , but they do not appear in europe and also not in the mediterranean sea . the new results indicate that several species of the butidae thrived in europe during the oligocene and early miocene . similar to the recent butidae and eleotridae , these fishes were adapted to a wide range of salinities and thrived in freshwater , brackish and marginal marine habitats . the fossil butidae disappeared from europe and the mediterranean and paratethys areas during the early miocene , due probably to their lack of competitiveness compared to other gobioidei that radiated during this period of time . in addition , this study documents the great value of otoliths for gobioid systematics .
citation : gierl c , reichenbacher b , gaudant j , erpenbeck d , pharisat a ( 2013 ) an extraordinary gobioid fish fossil from southern france . plos one 8 ( 5 ) : e64117 . urltoken
copyright : © 2013 gierl et al . this is an open - access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .
funding : the work was supported by the department for earth and environmetal sciences at lmu university . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
a . location of aix - en - provence in southern france . b . position of the fish fossil - bearing outcrop ( indicated by star ) in aix - en - provence . c . lithofacies of the sediments ( hammer for scale ) at the studied outcrop .
specimens were collected in 1974 by one of us ( ap ) in the city of aix - en - provence , at the corner of avenue philippe solari and chemin du pin , during the construction of a house [ 45 ] ( figs . 1b–c ) . the lithofacies of the laminated marly sediments indicates that they belong to the aix - en - provence formation and most probably to its second member , i . e . the calcaires et marnes des stations d’essence . a more precise correlation is not possible because , contrary to the feebly inclined strata of the aix - en - provence formation along the road to avignon , the strata of chemin du pin show a rather steep eastward dip ( about 40° ) ( fig . 1c ) .
the material is comprised of nine articulated skeletons today deposited in the palaeontological collection of the museum g . cuvier ; montbéliard ( france ) , under accession numbers mc - p - 2011 - 01 - tf1 to - tf9 . four of the skeletons possess saccular otoliths in situ . in addition , three specimens were available for comparison from the collections of the hessian state museum in darmstadt ( hlmd smff - 356 , determined as gobius aries ( agassiz ) ; hlmd 1910 - v - 2543 , determined as ophidion barbatum linnaeus ) and the museum for natural history in vienna ( nmw 1910 - v - 12 , determined as gobius aries ) .
osteological , meristic and morphometric characters of the skeletons and otolith characters were studied under a stereomicroscope equipped with a digital camera . measurements were taken with a calliper to the nearest 0 . 1 mm . the d1 pterygiophore formula ( e . g . 4 ( 22110 ) ) follows birdsong et al . [ 5 ] ; the first number indicates the position of the interneural space with the first pterygiophore ( e . g . behind vertebra 4 ) , each figure within the brackets represents an interneural space , starting with the one into which the first pterygiophore is inserted ( e . g . behind vertebra 4 ) , and the number indicates the number of pterygiophores inserting at that position ( e . g . , two pterygiophores behind vertebrae 4 and 5 , respectively ; one pterygiophore behind vertebrae 6 and 7 , respectively ; no pterygiophore behind vertebra 8 ) .
branched rays . the counts of the predorsal scales and the number of scales in the longitudinal and transverse rows follow masuda et al .
a . counting of the numbers of scales in the longitudinal row ( lr ) and transverse row ( tr ) . b . counting of the predorsal scales ( prsc ) in the dorsal midline in front of the dorsal fin .
information on the skeletons of extant gobioids for comparison with our fossils was gathered from the primary literature , mainly from the studies by regan [ 47 ] , hoese [ 6 ] , akihito et al . [ 48 ] , birdsong et al . [ 5 ] , harrison and miller [ 49 ] , hoese and gill [ 7 ] , johnson and brothers [ 50 ] , pezold [ 10 ] , winterbottom [ 51 ] , miller [ 9 ] , akihito et al . [ 12 ] , larson and murdy [ 52 ] , harrison et al . [ 53 ] , kindermann et al . [ 54 ] , froese and pauly [ 55 ] , and gill and mooi [ 19 ] . the information used in the comparison of the otoliths was obtained from specimens of extant eleotrids and butids kept in the collections of the institut royal des sciences naturelles de belgique and of dr . w . schwarzhans ( hamburg ) .
institutional abbreviations used : hlmd , hessian state museum in darmstadt , germany ; irsnb , institut royal des sciences naturelles de belgique , brussels , belgium ; mc , museum g . cuvier , montbéliard , france ; nmw , museum of natural history of vienna , austria ; usnm , national museum of natural history , smithsonian institution , washington , d . c . , u . s . a . ; wam , western australian museum , perth , western australia ; zmh , zoological museum hamburg , germany ; zmuc , zoological museum , copenhagen , denmark .
no permits were required for the described study , which complied with all relevant regulations .
the classification follows nelson [ 1 ] . for a newly proposed classification , see also wiley and johnson [ 56 ] .
a1 . specimen mc - p - 2011 - 01 - tf8 , right pectoral fin ( pect ) and anterior portion of the vertebral column . the first pterygiophore of the first dorsal fin ( pt1 d1 ) inserts behind the neural spine ( ns ) of vertebra 4 ( v4 ) , and the first pterygiophore of the second dorsal fin ( pt1 d2 ) inserts behind the neural spine of vertebra 9 ( v9 ) . the interneural gap ( intn gap ) lies between vertebrae 8 and 9 . a2 . close - up of the putative separated pelvic fins ( pelv ) . a3 . close - up of the dorsal part of the caudal endoskeleton , showing the neural spines of the pen - and antepenultimate vertebrae ( pu2 , pu3 ; note the short ns of pu2 ) , the two epurals ( ep1 , ep2 ) and the dorsal hypural plates ( hy3 + 4 , hy5 ) . b . specimen mc - p - 2011 - 01 - tf7 , anterior part of the vertebral column , showing the spines of the first dorsal fin , the prominent pterygiophore supporting the last spine ( pt6 d1 ) , the interneural gap ( intn gap ) between the neural spines of vertebrae 8 and 9 ( ns v8 , ns v9 ) , and the first pterygiophore of the second dorsal fin ( pt1 d2 ) . c . close - up of caudal fin of specimen mc - p - 2011 - 01 - tf3 ( see fig . 4d ) , showing the expanded neural and haemal spines ( hs ) of the antepenultimate vertebra ( pu3 ) , two epurals ( ep1 , ep2 ) , two large hypural plates ( hy1 + 2 , hy3 + 4 ) and a small one ( hy5 ) , the parhypural ( ph ) and the branched rays ( 7 / 6 ) . note that numbering of epurals indicates position and does not imply homology .
( agassiz ) ( a ) ; extant butidae ( b ) , eleotridae ( c ) , gobiidae ( d ) and odontobutidae ( e–f ) .
1833–43 cottus aries sp . nov . – agassiz , recherches sur les poissons fossiles , vol . iv , p . 186–187 .
1975 gobius aries ( agassiz ) . – gaudant , intérêt paléoécologique de la découverte de gobius aries , p . 112 , pl . 1 .
1978 gobius aries ( agassiz ) . – gaudant , sur les conditions de gisement de l’ichthyofaune oligocène d’aix - en - provence , table 1 .
1981 gobius aries ( agassiz ) . – gaudant , mise au point sur l’ichthyofaune oligocène des anciennes plâtrières d’aix - en - provence ( bouches - du - rhône ) , p . 1111 .
nine articulated skeletons , mc - p - 2011 - 01 - tf1 to - tf9 .
corner of avenue philippe solari and chemin du pin , city centre aix - en - provence ( bouches - du - rhône , france ) .
aix - en - provence formation , member calcaires et marnes des stations d’essence .
the size ranges between 70 and 96 mm total length and 51 to 81 mm standard length ( sl ) . the body is subcylindric , with a relatively long caudal peduncle ( 22–26 % of sl , table 1 ) ; body depth is about one - quarter ( 23–28 % ) of sl ( table 1 , figs . 3 , 4 ) . the body is covered with ctenoid scales ( fig . 3b 4 ) , but cycloid scales occur in the predorsal region ( fig . 3b 3 ) . the lateral line is absent .
the head is large and robust ; its length is included about three times in the standard length ( table 1 ) . the orbit is of medium size , subdorsal , and slightly elliptic ; the eye diameter is approximately one sixth of the head length ( fig . 3b 1 ) . the mouth is terminal , the gape oblique , with the posterior end slightly in front of the orbit ( figs . 3b 1–b2 ) .
the ethmoid region is short and bears a short mesethmoid ; other ethmoid bones and the nasal bone are not recognizable . the vomer is short and rounded and apparently bears no teeth . the frontal bones display an elongate supraorbital and a widened postorbital section and show prominent crests of the supra - and postorbital sensory canals ( figs . 4b–c 1 ) . a small infraorbital ( io2 or io3 ) is present ( fig . 4b ) . in the otic region appear round epiotic bones , in four specimens with otoliths in situ ( figs . 3a 1 , 4c–d ) . parietal bones are absent . the parasphenoid is visible in the lower third of the orbit ( fig . 3b 2 ) .
the shape of the saccular otolith ( termed otolith in the following ) is rectangular , the lateral ( outer ) side slightly convex and the medial ( inner ) side almost flat . the dorsal otolith margin displays a fine crenulation and a prominent posterodorsal projection . the posterior otolith margin is first concave ( below the posterodorsal projection ) , then it runs slightly oblique in posterior - ventral direction and meets the ventral margin with an angle of about 70–80° . the ventral otolith margin is crenulated and faintly bent ; it bears a small praeventral projection . the anterior otolith margin is slightly concave ; its junction with the dorsal margin is angular or faintly rounded .
the medial side of the otolith presents a sulcus that shows the “shoe - sole - like” shape that is present in most gobioid otoliths ( fig . 3a 4 ) . the sulcus covers approximately 85 % of the median otolith length and is anteriorly extended , i . e . positioned closer to the anterior than to the posterior otolith margin . a distinct , but thin line borders the tip of the ostium ; a small prae - ostial area is visible between the ostium tip and the anterior otolith margin . a slightly thickened crista superior is present above the middle and posterior part of the sulcus , while a thin crista inferior appears along the entire sulcus . above the sulcus appears the elongate , moderately incised dorsal area . the ventral line is running along the ventral rim of the otolith and ascending posteriorly .
note that the otolith preserved in situ in specimen mc - p - 2011 - 01 - tf1 ( fig . 3a 4 ) displays an ostium that seems to be widely opened to the anterior margin , but this is an artifact produced by the poor preservation of this otolith ; the thin suture bordering the ostium tip is destroyed due to corrosion .
the general structure of the upper jaw is well exposed in several specimens . the premaxilla is bent in the anterior section and bears a prominent processus articularis ( figs . 3a 1–2 , 4a ) and an almost rectangular - shaped , large posterior processus ( fig . 3b 2 ) ; the processus ascendens is longer than the processus articularis ( visible in specimen nmw 1910 - v - 12 ) . the oral edge of the premaxilla has three rows of irregularly arranged alveoles and conical teeth of different sizes , i . e . small ( 0 . 08–0 . 17 mm ) , medium - sized ( 0 . 25 mm ) or rather large ( 0 . 4 mm ) ( fig . 3b 5 ) ; the largest teeth insert mostly along the outer margin of the premaxilla . the maxilla is bent anteriorly and slightly expanded posteriorly , with a prominent articular head ( fig . 4a ) ; it is toothless .
the lower jaw is robust , but not preserved completely . the dentary is associated with a wedge - shaped angulo - articular ( fig . 3a 3 ) , the oral edge of the dentary displays two or three rows of irregularly arranged alveoles and conical teeth of different sizes ; the largest teeth reach 0 . 55 mm ( the total length of this specimen is 90 mm ) .
the palatine is elongate , robust , and l - shaped , i . e . it has a prominent anterior process ( fig . 4a ) . the quadrate is three - dimensionally preserved in several of the specimens . it is triangular , with a well - developed posterior process and a large articular head . posteriorly , the quadrate is articulated with the symplectic that is rod - shaped anteriorly and v - shaped posteriorly ( fig . 3a 1 ) , anteriorly it is associated with the elongate ectopterygoid and entopterygoid ( fig . 3b 2 ) . a gap between the symplectic and the preopercle is visible in the comparative material ( nmw 1910 - v - 12 , hlmd 1910 - v - 2543 ) .
the hyomandibula is large , but usually damaged . the further hyoid region includes a large ceratohyal , which can be subdivided into an anterior ceratohyal and a triangular - shaped posterior ceratohyal ( = epihyal ) . two narrow branchiostegal rays ( rays 1–2 ) and four robust branchiostegal rays ( rays 3–6 ) articulate with the anterior ceratohyal , between rays 1–2 and rays 3–6 is a distinct gap ( fig . 3b 6 ) .
the opercular bones are large , but mostly not well preserved . the subopercle is semi - circular ( fig . 4d ) . the crescent - shaped preopercle ( best visible in specimen nmw 1910 - v - 12 ) has a lower and upper arm of almost equal length , and the opercle is rounded - triangular with the tip pointing to the ventral margin , a thickening is present at its anterior and posterior margins ( visible in specimen hlmd 1910 - v - 2543 ) .
many conical pharyngeal teeth ranging in size between 0 . 05 and 0 . 35 mm are present in the branchial region of most specimens . slender ceratobranchials and numerous , very small , spiny gill rakers are also visible ( fig . 4d ) .
it comprises 25–26 ( 10 + 15–16 ) vertebrae ( v ) , which have a delicate net - like structure ( fig . 4c 2 ) . the anteriormost vertebral centra ( v1 to v4 / v5 ) are less elongate and less constricted in the middle than the subsequent ones .
the neural spines of v1 to v3 are expanded and triangular . the other neural spines are of almost equal length and elongate ( figs . 3a 1 , b1 ) , with the exception of the short neural spine of the second preural centrum ( pu2 ; fig . 5a 3 ) . in addition , the neural spines of pu3–5 are more oblique than the preceding ones ; they appear at the posterior end of the respective centrum .
the haemal spines are usually as long as the neural spines , with the exception of the haemal spine of the first caudal vertebra ( v11 ) , which is slightly shortened ( fig . 5a 1 ) , and the haemal spine of pu2 , which is enlarged ( fig . 5c ) .
all abdominal vertebrae have long and prominent parapophyses , in some specimens preserved in connection with the ribs . seven to eight rib pairs are recognizable , the last two rib pairs are slightly shorter than the preceding ones ( fig . 5a 1 ) . the ribs are long , have a strong and thickened proximal portion , and a pointed distal tip ; long epipleurals are also present ( fig . 4d ) . there are no supraneurals .
the cleithrum is long , slender and reveals a broad to triangular widening in its ventral portion ( visible in specimen nmw 1910 - v - 12 ) . the supracleithrum is rather robust ( fig . 3a 1 ) . the endoskeleton of the pectoral fin consists of four well - developed hour - glass radials ( r ) with ovate - shaped gapes in between ( fig . 3b 7 ) . a scapula and coracoid are not recognizable .
the pectoral fins , which are inserted in the lower third of the flank , comprise 14–16 rays . the pelvic fins are inserted just under or slightly behind the pectoral fins and are probably separated ( fig . 5a 2 ) . each pelvic fin includes one spine ( visible in specimen nmw 1910 - v - 12 ) and five rays . the endoskeleton of the pelvic fin consists of an elongate , slightly triangular basipterygium ( fig . 5a 2 ) .
the first dorsal fin ( d1 ) has six unbranched and unsegmented , medially paired rays , increasing in length from ray 1 to 3 and then slightly descending ; the last ray follows with a short gap the preceding ones . every ray is supported by an elongate pterygiophore , distally ending in a slightly concave depression ( for the articulation of the ray ) ( fig . 5b ) . the first pterygiophore inserts behind the neural spine of vertebra 4 and the last pterygiophore is situated behind the neural spine of vertebra 7 ; the d1 pterygiophore formula is 4 ( 22110 ) ( figs . 5a 1 , b ) .
the second dorsal fin ( d2 ) is inserted slightly in front of the insertion of the anal fin . the first ray may perhaps be a spine , and then follow 10 segmented and branched rays ( fig . 3b 1 ) . the first ray ( or spine ) is supported by two pterygiophores , which both insert in the interneural space behind the vertebra 9 ( figs . 5a 1 , b ) . every pterygiophore supporting the rays is associated to a neural spine .
the anal fin consists of a very small spine and 8 to 9 rays . the spine and the first ray are supported by a single pterygiophore . the first three or four pterygiophores insert in front of the haemal spine of the second caudal vertebra ( v12 ) , they are slightly longer than the following ones ( fig . 3b 1 ) .
the caudal endoskeleton bears two large , triangular hypural plates ( hy1 + 2 , hy3 + 4 ) and an additional small hypural plate in the dorsal part ( hy5 ) , which is separated by a thin suture from hy3 + 4 ( fig . 5c ) . a short and slender parhypural is present and is closely associated with hy1 + 2 ; its proximal region is reduced and does not reach the terminal centrum ( fig . 5c ) .
two epurals ( ep ) are present ( figs . 5a 3 , c ) . the anterior one is characterized by a longitudinal median rib and pointed proximally ; its proximal end is close to the terminal centrum . the posterior epural is slightly shorter than the anterior epural and also shows a longitudinal rib .
the number of principal caudal fin rays that are segmented and branched is 13 . the caudal fin formula is 7 / 6 ( fig . 5c ) . in addition to the segmented and branched caudal fin rays , three long and ten short unbranched rays are present dorsally , and six rather long and an undetermined number of short rays appear ventrally ( fig . 5c ) .
the uppermost segmented and branched caudal fin ray is supported by hy5 , the next six rays are supported by hy3 + 4 , the next five rays are supported by hy1 + 2 , and the parhypural supports the lowermost segmented and branched ray ( fig . 5c ) . in addition , the epurals and the widened neural spine of pu3 contribute to the caudal endoskeleton by supporting the dorsal unbranched rays ( fig . 5c ) . the expanded haemal spine of pu2 supports the first ( longest ) ventral unbranched ray ( fig . 5c ) .
ctenoid scales are present all over the body except in front of the first dorsal fin . they do not differ much in size and shape in the dorsal and ventral body parts ( figs . 3a 1 , b1 ) and display regularly arranged radii and tiny ctenii ( fig . 3b 4 ) . specimen mc - p - 2011 - 01 - tf2 shows well preserved rows of scales with an average width of 0 . 9 mm for the not - imbricated part of the scale ( fig . 3b 4 ) ; thus , the number of scales along the lateral series ( = longitudinal scale row , see fig . 2 ) can be estimated as being about 50 .
cycloid scales are present in the predorsal region ( figs . 3a 1 , b1 ) . they show distinct radii , numerous fine circuli , and are thinner and relatively higher than the ctenoid scales . in specimen mc - p - 2011 - 01 - tf2 , the width of the not - imbricated part of a predorsal scale is about 0 . 5 mm and the predorsal segment that is covered with this type of scales is about 12 . 8 mm . therefore the number of predorsal scales can be calculated to have been about 25 .
specimen mc - p - 2011 - 01 - tf2 displays well - preserved content of the stomach and gut ( fig . 3b 1 ) , which consists of densely packed gastropod shells .
it should be mentioned that there are some differences between our results and the descriptions given by agassiz [ 34 ] and sauvage [ 35 ] . both observed a lower number of rays in the second dorsal fin and pelvic fin ( nine and four , vs . eleven and six in our specimens ) . moreover , sauvage [ 35 ] reported spines both on the preopercle and opercle ( not visible in our specimens ) . it is possible that these differences result from different preservation states of the individual specimens , or that sauvage [ 35 ] believed that he had seen spines on the preopercle and opercle because it was an argument in favour of an attribution to the cottids .
otoliths with a shoe - sole - like sulcus ( see [ 29 ] ) .
there is no consensus of the phylogenetic relations and number of families within the gobioidei ( see [ 19 ] , [ 58 ] , [ 59 ] ) . several family - based clades have been introduced , i . e . the rhyacichthyidae ( see [ 60 ] ) , odontobutidae ( see [ 61 ] ) , butidae and eleotridae ( see [ 13 ] , [ 21 ] ) , gobionellidae ( gobionellinae sensu pezold ( 1993 ) , see [ 23 ] ) , gobiidae ( see [ 62 ] ) , milyeringidae ( see [ 18 ] ) , and thalasseleotrididae ( see [ 19 ] ) . the butidae and eleotridae sensu thacker [ 13 ] have been previously treated as subfamilies ( butinae , eleotrinae ) of the eleotridae ( e . g . [ 1 ] , [ 5 ] , [ 7 ] , [ 63 ] ) . thacker’s families consisted of the same genera as the subfamilies . the milyeringidae represent small blind cave fishes that were previously assigned to the butidae and odontobutidae , respectively [ 13 ] , [ 14 ] . the thalasseleotrididae comprise two marine genera from australia and new zealand that were previously considered as members of the eleotridae [ 19 ] .
† l . aries may thus belong either to the butidae or eleotridae , with which it shares the presence of an entopterygoid and an l - shaped palatine . the presence of an interneural gap between the two dorsal fins does not conflict this assignment . even though the interneural gap is a typical character of the gobiidae , it also occurs in a few butidae and eleotridae [ 5 ] . in addition , the insertion of the first two pterygiophores of the second dorsal fin in the same interneural space , as observed in † l . aries , is consistent with those few butidae and eleotridae that display an interneural gap , but differs from almost all gobiidae with separate dorsal fins ( see [ 5 ] ) .
the separation of the butidae and eleotridae is supported by molecular data [ 13 ] , but is less clear if anatomical data is taken into consideration [ 5 ] , [ 7 ] , [ 12 ] . specializations both of the jaw musculature and caudal cartilage are the known synapomorphies used to characterize the eleotridae [ 7 ] . however , muscles and cartilages are not preserved in fossil gobioids . moreover , no synapomorphy is presently available to classify the butidae ; rather , this family is currently identified as lacking the apomorphies that define the eleotridae [ 7 ] .
the composition of the unpaired and paired fins , the number of longitudinal and transverse scales , as well as the number of vertebrae are not appropriate to separate between butidae and eleotridae . examples are the counts of i8 for the second dorsal fin , i9 for the anal fin , 9 for the number of transverse scales , and 10 + 15–16 for the precaudal / caudal vertebrae that all are present both in butid and eleotrid species ( see table 3 ) . moreover , it appears that the only meristic characters that are diagnostic for an individual genus include the number ( or ranges ) of predorsal and longitudinal scales ( table 3 ) . adding support to this suggestion comes from the “keys” for the genera of butidae and eleotridae [ 52 ] , in which the number ( or ranges ) of these scales is used as a key character in the identification of genera . as a result , the number of predorsal or longitudinal scales represents an autapomorphy for a given genus of the butidae and eleotridae , but does not discriminate between the families .
the scales of extant butids and eleotrids usually are ctenoid , but several exceptions exist . for example , eleotris possesses small cycloid scales in the predorsal region , at the pectoral - fin base , at the abdomen , and in one or two rows along the borders of the median fins , whereas large ctenoid scales cover the sides of its trunk [ 66 ] . the eleotrid philypnodon grandiceps ( krefft ) also shows a combination of both scale types along the body , whereas erotelis has exclusively small cycloid body scales [ 9 ] , [ 67 ] . among the butidae , pogoneleotris displays ctenoid plus auxiliary cycloid body scales , kribia produces cycloid predorsal scales , but large and ctenoid body scales ( with the exception of a few cycloid scales that may appear on the border of the second dorsal fin ) , while bostrychus is the only butid genus that is covered exclusively by cycloid scales [ 52 ] , [ 53 ] . as a result , the presence or absence of cycloid or ctenoid scales cannot be used for a general discrimination between butidae and eleotridae .
however , pezold and cage [ 66 ] have shown that the occurrence of many small cycloid body scales ( termed as secondary cycloid scales in the following ) can be interpreted as a derived character in erotelis , while the appearance of large ctenoid scales is regarded as conservative . accordingly we hypothesize that , during evolution , secondary cycloid scales successively replaced ctenoid scales in both butidae and eleotridae . if this hypothesis is accurate , then the many small cycloid scales of bostrychus would be more derived than the scale pattern of any other butid genus . this is consistent with the position of bostrychus on the molecular tree presented by thacker [ 13 ] . as a result , the presence of secondary cycloid scales may represent a tool for determining relationships within the butidae and eleotridae .
the morphological characters of the saccular otolith ( termed otolith in the following ) hitherto have received little attention in phylogenetic studies . otolith formation involves hormonally - regulated calcium carbonate deposition within an organic framework , but otoliths are not part of the skeleton and have evolved independently [ 68 ] . their significance in phylogenetic analyses remains to be fully explored ; however , it is long since known that otolith morphology can be characteristic not only at the species level , but also at genus and family level ( [ 29 ] , [ 69 ] and many others ) .
a . butis butis ( hamilton ) , sumatra , coll . schwarzhans , leg . zmh . b . ophiocara porocephala ( valenciennes ) , manus island , bismarck archipel , zmuc p . 781771–78 . c . oxyeleotris lineolata ( steindachner ) , northern australia , coll . schwarzhans , leg . wam . d . bostrychus sinensis lacepède , china , coll . schwarzhans , leg . zmh .
among the butidae , the otolith of † l . aries shares the rectangular shape and the anteriorly extended sulcus with bostrychus africanus ( steindachner ) , b . strigogenys nichols , b . sinensis lacepède ( fig . 7d ) , kribia kribensis ( boulenger ) ( fig . 6b ) , ophiocara porocephala ( valenciennes ) ( fig . 7b ) , and oxyeleotris lineolata ( steindachner ) ( fig . 7c ) . the otolith of butis butis ( hamilton ) is different from the former taxa in that it is rectangular to trapezoid in shape and possesses an almost centered sulcus ( fig . 7a ) . otoliths of oxyeleotris species other than o . lineolata are rectangular in shape , but their sulcus is centered ( figured in [ 65 ] : figs . 12l–n ) , which might suggest that their assignment to the butidae may deserve further investigation .
for genus apomorphies we used the number of scales in the longitudinal row ( lr ) according to larson and murdy [ 52 ] .
our phylogenetic hypothesis suggests an early divergence of butis from the other butids , which is consistent with the molecular phylogeny of thacker [ 13 ] . in addition , oxyeleotris appears to diverge earlier than ophiocara , which also does not conflict the molecular phylogeny of thacker [ 13 ] . a difference between our scenario and thacker’s phylogeny concerns the position of bostrychus as sister to kribia ( bostrychus is sister to ophiocara and kribia is positioned within a polyphyletic oxyeleotris clade in thacker ) .
we thank t . malvesy ( mc montbéliard ) , n . micklich ( hlm darmstadt ) and u . göhlich ( nm wien ) for the loan of specimens . n . micklich and u . göhlich are additionally acknowledged for helpful comments on the original provenience of the museum specimens . we thank d . nolf ( irsnb ) and w . schwarzhans ( hamburg ) for the loan of otoliths and constructive discussion . we are especially grateful to d . nolf for providing his unpublished drawings of odontobutidae . we thank j . stolarski ( editor of acta palaeontologica polonica ) for the permission for re - figuration of some previously published otoliths . we are grateful to michael krings ( munich ) for constructive comments and improving our english , and we thank the reviewers g . arratia ( lawrence ) and g . carnevale ( pisa ) for their constructive remarks and suggestions .
conceived and designed the experiments : cg br . performed the experiments : cg br jg ap . analyzed the data : cg br jg de . contributed reagents / materials / analysis tools : cg br ap . wrote the paper : cg br jg .
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pezold f ( 1993 ) evidence for a monophyletic gobiinae . copeia 1993 : 634–643 .
, new genus and species of western pacific fish ( gobiidae , xenisthminae ) , with discussions of gobioid osteology and classification . washington , d . c . : smithsonian institution press . 40 p .
akihito , iwata a , kobayashi t , ikeo k , imanishi t , et al . ( 2000 ) evolutionary aspects of gobioid fishes based upon a phylogenetic analysis of mitochondrial cytochrome b genes . gene 259 : 5–15 .
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wang h - y , tsai m - p , dean j , lee s - c ( 2001 ) molecular phylogeny of gobioid fishes ( perciformes : gobioidei ) based on mitochondrial 12s rrna sequences . mol phylogenet evol 20 : 390–408 .
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rüber l , agorreta a ( 2011 ) molecular systematics of gobioid fishes . in : patzner ra , van tassell jl , kovacic m , kapoor bg , editors . the biology of gobies . enfield , nh : science publishers inc . pp . 23–50 .
gill ac , mooi rd ( 2012 ) thalasseleotrididae , new family of marine gobioid fishes from new zealand and temperate australia , with a revised definition of its sister taxon , the gobiidae ( teleostei : acanthomorpha ) . zootaxa 3266 : 41–52 .
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gaudant j ( 1996 ) find of the oldest known gobioids ( teleostean fishes ) in the marine lower lutetian of catalonia ( spain ) . c r acad sci ii 322 : 71–76 .
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19 specimens of peronochaeta are known from the greater phyllopod bed , where they comprise < 0.1 % of the community .

peronochaeta

peronochaeta – from the greek perone , “needle , ” and khait , “long hair , ” in reference to its bristles .
peronochaeta is a genus of annelid known from the middle cambrian burgess shale . { gpb abundance | 19 | < 0 . 1 } . . .
peronochaeta dubia ( rom 61133 ) . complete specimen associated with an indeterminate fossil ( top right ) . specimen length = 8 mm . specimen dry – polarized light . walcott quarry .
peronochaeta bears some resemblance to modern polychaetes but it cannot be placed in any extant group ( conway morris , 1979 ) suggesting a position as a stem - group polychaete ( eibye - jacobsen , 2004 ) .
peronochaeta was considered one of the rarest annelids from the burgess shale but additional material has now been collected from the walcott quarry representing 0 . 03 % of the specimens counted in the community ( caron and jackson , 2008 ) .
this annelid worm was originally described as a species of canadia by charles walcott ( 1911 ) . when simon conway morris ( 1979 ) re - examined these fossils , he concluded that the differences between this species and canadia were too great to be contained within a single genus , and erected the new genus peronochaeta .
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current links for doi : 10 . 2110 / palo . 2003 . p05 - 070r
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the society for sedimentary geology is an international not - for - profit society based in tulsa , oklahoma . through its network of international members , the society is dedicated to the dissemination of scientific information on sedimentology , stratigraphy , paleontology , environmental sciences , marine geology , hydrogeology , and many additional related specialties .
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dubia – from the latin dubius , “uncertain , ” presumably reflecting walcott’s uncertainty regarding his original classification of this worm as canadia .
lectotype – unsm 83936a ; paralectotype – unsm 83936d , in the national museum of natural history , smithsonian institution , washington , dc , usa .
middle cambrian , bathyuriscus - elrathina zone ( approximately 505 million years ago ) .
this worm reached one to two centimetres in length , but its body is only 1 mm wide , or 2 mm wide , if its spines ( setae ) are included . the worm has approximately 25 segments , each bearing a pair of short lateral projections called parapodia . these are simple ( uniramous ) and the setae are short . a straight gut runs the length of its body . a pair of tentacles appears to be preserved on the sides of the head , although due to the small size and poor preservation , it is difficult to assert this with confidence .
on account of the scarcity of material , the ecology of this animal is difficult to ascertain . it may have been a scavenger , and its setae probably assisted in locomotion and perhaps even in burrowing .
caron , j . - b . and d . a . jackson . 2008 . paleoecology of the greater phyllopod bed community , burgess shale . palaeogeography , palaeoclimatology , palaeoecology , 258 : 222 - 256 .
conway morris , s . 1979 . middle cambrian polychaetes from the burgess shale of british columbia . philosophical transactions of the royal society of london . series b , biological sciences , 285 ( 1007 ) : 227 - 274 .
eibye - jacobsen , d . 2004 . a reevaluation of wiwaxia and the polychaetes of the burgess shale . lethaia , 37 ( 3 ) : 317 - 335 .
walcott , c . d . 1911 . middle cambrian annelids . smithsonian miscellaneous collections , 57 ( 2 ) : 109 - 144 .
morris , s . c . ( 1979 ) . middle cambrian polychaetes from the burgess shale of british columbia . philosophical transactions of the royal society b : biological sciences . 285 ( 1007 ) : 227 - 274 . , available online at urltoken [ details ]
read , g . ; fauchald , k . ( ed . ) ( 2018 ) . world polychaeta database .
( of canadia dubia walcott , 1911 † ) walcott , charles d . 1911 . cambrian geology and paleontology . middle cambrian annelids . smithsonian miscellaneous collections , 57 ( 5 ) : 109 - 144 . , available online at urltoken page ( s ) : 119 [ details ]
( of canadia dubia walcott , 1911 † ) walcott , charles d . 1931 . addenda to descriptions of burgess shale fossils . smithsonian miscellaneous collections 85 ( 3 ) : 1 - 46 , available online at urltoken [ details ]
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parent taxon : annelida according to j . - b . caron and d . a . jackson 2008
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the yucatan flycatcher ( myiarchus yucatanensis ) is a species of bird in the family tyrannidae .

it is endemic to the yucatan peninsula of mexico and the northern bordering parts of belize and guatemala .

it is a typical myiarchus flycatcher ; partially crested with a brown to gray back and head , a rufous tail and yellow to pale underparts .

it is closely related to and similar in appearance to the dusky-capped flycatcher ( m. tuberculifer ) , brown-crested flycatcher ( m. tyrannulus ) and the great crested flycatcher ( m. crinitus ) .

these species are best distinguished by voice .

its natural habitats are subtropical or tropical dry forests and heavily degraded former forest .

yucatan flycatcher

specific limits of the yucatan flycatcher , myiarchus yucatanensis . american museum novitates ; no . 2229
details - specific limits of the yucatan flycatcher , myiarchus yucatanensis . american museum novitates ; no . 2229 - biodiversity heritage library
ty - book ti - specific limits of the yucatan flycatcher , myiarchus yucatanensis . american museum novitates ; no . 2229 ur - urltoken py - 1965 au - lanyon , wesley e . er -
@ book { bhl169241 , title = { specific limits of the yucatan flycatcher , myiarchus yucatanensis . american museum novitates ; no . 2229 } , url = urltoken note = urltoken publisher = { } , author = { lanyon , wesley e . } , year = { } , pages = { 0 } , }
joseph , l . ( 2018 ) . yucatan flycatcher ( myiarchus yucatanensis ) . in : del hoyo , j . , elliott , a . , sargatal , j . , christie , d . a . & de juana , e . ( eds . ) . handbook of the birds of the world alive . lynx edicions , barcelona . ( retrieved from urltoken on 9 july 2018 ) .
as its name suggests , this flycatcher is endemic to the yucatán peninsula , where it ranges as far south as northern guatemala and northern belize , as well as being found on cozumel island . like many myiarchus flycatchers , identification is most reliably based on voice , though visual identification is not impossible . three subspecies are generally recognized , of which the most widespread geographically , the nominate , is strikingly rufous - crowned , making it comparatively easily identified . there is also substantial rufous in the primaries . the yucatan flycatcher is restricted to the lowlands , where it is found in both semi - arid and humid forests , including their edges .
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del hoyo , j . , collar , n . j . , christie , d . a . , elliott , a . , fishpool , l . d . c . , boesman , p . and kirwan , g . m . 2016 . hbw and birdlife international illustrated checklist of the birds of the world . volume 2 : passerines . lynx edicions and birdlife international , barcelona , spain and cambridge , uk .
justification : this species has a very large range , and hence does not approach the thresholds for vulnerable under the range size criterion ( extent of occurrence < 20 , 000 km2 combined with a declining or fluctuating range size , habitat extent / quality , or population size and a small number of locations or severe fragmentation ) . despite the fact that the population trend appears to be decreasing , the decline is not believed to be sufficiently rapid to approach the thresholds for vulnerable under the population trend criterion ( > 30 % decline over ten years or three generations ) . the population size may be moderately small to large , but it is not believed to approach the thresholds for vulnerable under the population size criterion ( < 10 , 000 mature individuals with a continuing decline estimated to be > 10 % in ten years or three generations , or with a specified population structure ) . for these reasons the species is evaluated as least concern .
partners in flight estimated the population to number fewer than 50 , 000 individuals ( a . panjabi in litt . 2008 ) , thus it is placed in the band 20 , 000 - 49 , 999 individuals here . trend justification : the population on cozumel has proven difficult to locate , suggesting a recent decline ( del hoyo et al . 2004 ) .
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) , in neotropical birds online ( t . s . schulenberg , editor ) . cornell lab of ornithology , ithaca , ny , usa . retrieved from neotropical birds online :
id certainty 90 % . ( archiv . tape 71 side a track 43 seq . a )
id certainty 90 % . ( archiv . tape 71 side a track 40 seq . a )
from a bird in low vine and scrub next to the old airstrip . after playback of xc126557 , this call after the bird popped out and saw four people pointing bins at it .
from a bird in low vine and scrub next to the old airstrip . after playback of xc126557
sonogram images © xeno - canto foundation . sonogram images share the same license terms as the recording they depict .
hbw alive contains information on descriptive notes , voice , habitat , food and feeding , breeding , movements , status and conservation plus a list of bibliographical references for this species account .
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see m . tyrannulus ( below ) . geographical variation , including populations on cozumel i ( lanyoni ) , seems ecophenotypic ( correlated with rainfall and vegetation ) ; nominate race and navai apparently intergrade in c yucatán peninsula ; sampling needed to test validity of current taxonomy . three subspecies recognized .
lawrence , 1871 – e mexico in extreme e tabasco ( near balancán ) and n & c yucatán peninsula .
parkes & a . r . phillips , 1967 – cozumel i , off ne quintana roo ( mexico ) .
( most populations ) with sometimes distinct facial pattern . nominate race has broad . . .
dawn song a fairly loud , clear , monotonously repeated “hoor - eep” or “hoow’eep” , second part rising . . .
humid to semi - arid forest , scrubby woodland and edge ; in deciduous forest found primarily in . . .
not globally threatened . fairly common to common . occurs in lamanai archaeological reserve and río bravo conservation and management area , in belize , and laguna del tigre . . .
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highly speciose family , herein divided into nine subfamilies ( with ten tribes ) ; recent studies # r # r suggest that the first four listed ( platyrinchinae , pipritinae , tachurisinae , pipromorphinae ) are sister to rest of family , and may merit treatment as 1–4 separate families .
phylogenetic study based on mtdna # r found that , with exception of m . semirufus , this genus comprises two clades , one of predominantly caribbean and central and north american taxa , the other of south american taxa . see also rhytipterna ( above ) .
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myavibase allows you to create and manage your own lifelists , and produce useful reports to help you plan your next birding excursion .
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myiarchus yucatanensis yucatanensis : e mexico ( extreme e tabasco and n and cent . yucatán pen . )
myiarchus yucatanensis lanyoni : cozumel i . ( off yucatán peninsula of e mexico )
howard and moore 4th edition ( incl . corrigenda vol . 1 - 2 ) :
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new acquaintances
) , published continuously and numbered consecutively since 1921 , are short papers that contain descriptions of new forms and reports in zoology , paleontology , and geology . new numbers are published at irregular intervals .
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biodivlibrary @ jzed _ trees @ kew _ laa awesome ! we ' re so honored to have this treasure in # bhlib . thanks @ kew _ laa ! now we can enjoy p… urltoken


collix blosyra is a moth in the family geometridae .

it is found on borneo .

the habitat consists of upper montane forests .

adults are somewhat dark , uniform dull brown with weakly developed submarginal white spots .

collix blosyra

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walker , 1862 , list specimens lepid . insects colln br . mus . , 24 : 1249 .
only one bornean specimen has been located , from near pontianak in kalimantan . no altitude data are given .
bell ( ms ) described the biology in s . india . the eggs are laid singly on the leaves , twigs and axils of the host - plant , and the young larvae prefer to feed on young leaves . the larva is cylindrical , narrowing slightly at each end . the head is very slightly and broadly bilobed , wider than t1 , and yellow - green in colour . the body is a rosy pinkish green , variable in shade , with a white , longitudinal dorsolateral line and a pale band in the spiracular region . young larvae sit on the edges of leaves in a variety of postures , older ones being more on the underside and eating holes in the lamina . the pupa is formed in a slight silken cell in a rolled leaf edge . the host - plants recorded were ardisia and embelia ( myrsinaceae ) .
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curators : konstantin efetov , vasiliy feoktistov , svyatoslav knyazev , evgeny komarov , stan korb , alexander zhakov .
species catalog enables to sort by characteristics such as expansion , flight time , etc . .
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to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
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this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .


the lord howe stag beetle ( lamprima insularis ) is a species of beetles in the family lucanidae , that is endemic to lord howe island .

lamprima insularis

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the most colourful of the lord howe island beetles , occasionally seen during summer months .
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urn : lsid : biodiversity . org . au : afd . taxon : d9d5460f - 9b21 - 4484 - 88ae - 401893e5dab9
urn : lsid : biodiversity . org . au : afd . taxon : f47ac029 - 83b5 - 4725 - b299 - 6cc0263ad57d
urn : lsid : biodiversity . org . au : afd . taxon : 4ac6d4ff - a6e2 - 46e7 - ae1e - df40a4af4ba2
urn : lsid : biodiversity . org . au : afd . name : 345149
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bibio xanthopus is a species of march fly ( bibionidae ) first identified by wiedemann in 1828 .

it is one of the most common species of bibio in north america .

the body is black with pale coloured pile and legs that are predominantly rufous .

the size of the species varies considerably , with wingspans ranging from 5.5 to 9.5 mm .

bibio xanthopus

detail of male bibio xanthopus . note relatively rounded head ( in dorsal view ) and holoptic eyes . see first image of series for more info .
southern canada to central mexico . one of the most widespread and common species of bibio .
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upcoming events 2018 bugguide gathering in virginia july 27 - 29 : registration and discussion photos of insects and people from the 2015 gathering in wisconsin , july 10 - 12 photos of insects and people from the 2014 gathering in virginia , june 4 - 7 . photos of insects and people from the 2013 gathering in arizona , july 25 - 28 photos of insects and people from the 2012 gathering in alabama photos of insects and people from the 2011 gathering in iowa photos from the 2010 workshop in grinnell , iowa photos from the 2009 gathering in washington
contributed by john f . carr on 22 may , 2009 - 6 : 19pm last updated 14 april , 2016 - 8 : 04am
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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 .