leucoblepsis neoma is a moth in the drepanidae family .

it was described by swinhoe in 1905 .

it is found in singapore and on peninsular malaysia , sumatra and borneo .

adults are white , suffused with pale chestnut-brown irrorations in parts .

there are two nearly straight brown lines from the abdominal margin of the hindwings .

the first from one-third , the other from the middle , running parallel across both wings and suddenly converge on the middle of vein 5 of the forewings , and run from there in a single line to the costa near the apex .

between these lines on the forewings is a large hyaline spot and the rest of the wing is lightly suffused .

there is a fairly broad pale pinkish band on the outer margin , composed of large pale pinkish spots joined together .

the marginal line is brown .

the hindwings have a nearly white inner area , the outer area from the outer line to the margin with dark suffusion and a dentated white line through the middle of the disc .

the marginal line is white .

leucoblepsis neoma

leucoblepsis neoma ; [ mob8 ] : 33 , pl . 6 , f . 38
problepsidis neoma swinhoe , 1905 ; ann . mag . nat . hist . ( 7 ) 15 ( 86 ) : 150
leucoblepsis excisa ; [ mob8 ] : 33 , pl . 6 , f . 39
leucoblepsis renifera ; [ mob8 ] : 32 , pl . 6 , f . 35 , 36
leucoblepsis warren , 1922 ; gross - schmett . erde 6 : 462 ; ts : problepsidis carneotincta warren
leucoblepsis ostia swinhoe , 1903 ; in annandale & robinson , fasc . malay zool . 1 : 59
leucoblepsis fenestraria ; holloway , 1976 , moths of borneo with special reference to mt . kinabalu : 94 ; [ mob8 ] : 33 , pl . 6 , f . 37
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nb : the taxon name search is for single names only . for example , to locate dysodia vitrina flammata warren , 1904 you should enter flammata only .
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the species is rare in lowland forest , though one specimen has been taken at 1618m on bukit retak , brunei .
problepsidis tristis swinhoe , 1905 ; ann . mag . nat . hist . ( 7 ) 15 ( 86 ) : 150
ne . himalaya , taiwan , peninsular malaysia , sumatra , borneo . see [ maps ]
problepsidis carneotincta warren , 1901 ; novit . zool . 8 : 191 ; tl : sarawak , bukan
ne . hiamalaya , taiwan , peninsular malaysia , borneo . see [ maps ]
[ 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 .
warren , 1901 drepanulidae , uraniidae , and geometridae from the palaearctic and indo - australian regions novit . zool . 8 : 190 - 201
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this article has been cited by the following publications . this list is generated based on data provided by crossref .
brehm , gunnar 2017 . a new led lamp for the collection of nocturnal lepidoptera and a spectral comparison of light - trapping lamps . nota lepidopterologica , vol . 40 , issue . 1 , p . 87 .
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wolfling , mirko becker , mira c . uhl , britta traub , anja and fiedler , konrad 2016 . how differences in the settling behaviour of moths ( lepidoptera ) may contribute to sampling bias when using automated light traps . european journal of entomology , vol . 113 , issue . , p . 502 .
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zou , yi sang , weiguo hausmann , axel and axmacher , jan christoph 2016 . high phylogenetic diversity is preserved in species - poor high - elevation temperate moth assemblages . scientific reports , vol . 6 , issue . 1 ,
lamarre , g p a mendoza , i rougerie , r decaëns , t hérault , b and bénéluz , f 2015 . stay out ( almost ) all night : contrasting responses in flight activity among tropical moth assemblages . neotropical entomology , vol . 44 , issue . 2 , p . 109 .
akite , perpetra telford , richard j . waring , paul akol , anne m . and vandvik , vigdis 2015 . temporal patterns in saturnidae ( silk moth ) and sphingidae ( hawk moth ) assemblages in protected forests of central uganda . ecology and evolution , vol . 5 , issue . 8 , p . 1746 .
jonason , dennis franzén , markus ranius , thomas and brigham , r . mark 2014 . surveying moths using light traps : effects of weather and time of year . plos one , vol . 9 , issue . 3 , p . e92453 .
thompson , jessica l . zack , richard s . crabo , lars and landolt , peter j . 2014 . survey of macromoths ( insecta : lepidoptera ) of a palouse prairie remnant site in eastern washington state . pan - pacific entomologist , vol . 90 , issue . 4 , p . 191 .
teston , josé augusto novaes , jessé bucioli and almeida júnior , josé otávio barros 2012 . abundância , composição e diversidade de arctiinae ( lepidoptera , arctiidae ) em um fragmento de floresta na amazônia oriental em altamira , pa , brasil . acta amazonica , vol . 42 , issue . 1 , p . 105 .
beck , jan schwanghart , wolfgang khen , chey vun and holloway , jeremy d . 2011 . predicting geometrid moth diversity in the heart of borneo . insect conservation and diversity , vol . 4 , issue . 3 , p . 173 .
ozaki , kenichi sayama , katsuhiko ueda , akira ito , masato tabuchi , ken and hironaga , teruhiko 2011 . short - term , efficient sampling strategies for estimating forest moth diversity using light traps . annals of the entomological society of america , vol . 104 , issue . 4 , p . 739 .
garcía - lópez , alejandra micó , estefanía zumbado , manuel a . and galante , eduardo 2011 . sampling scarab beetles in tropical forests : the effect of light source and night sampling periods . journal of insect science , vol . 11 , issue . 95 , p . 1 .
teston , josé augusto and delfina , márcia cristina 2010 . diversidade de arctiinae ( lepidoptera , arctiidae ) em área alterada em altamira , amazônia oriental , pará , brasil . acta amazonica , vol . 40 , issue . 2 , p . 387 .
scalercio , stefano infusino , marco and woiwod , ian p . 2009 . optimising the sampling window for moth indicator communities . journal of insect conservation , vol . 13 , issue . 6 , p . 583 .
hawes , joseph da silva motta , catarina overal , william l . barlow , jos gardner , toby a . and peres , carlos a . 2009 . diversity and composition of amazonian moths in primary , secondary and plantation forests . journal of tropical ecology , vol . 25 , issue . 03 , p . 281 .
schooler , s . s . mcevoy , p . b . hammond , p . and coombs , e . m . 2009 . negative per capita effects of two invasive plants , lythrum salicaria and phalaris arundinacea , on the moth diversity of wetland communities . bulletin of entomological research , vol . 99 , issue . 03 , p . 229 .
scanlon , annette t . and petit , sophie 2008 . biomass and biodiversity of nocturnal aerial insects in an adelaide city park and implications for bats ( microchiroptera ) . urban ecosystems , vol . 11 , issue . 1 , p . 91 .
experimental comparisons were made between samples of moths obtained by rothamsted tungsten - filament and robinson mercury - vapour light - traps operating in a lowland dipterocarp forest in peninsular malaysia . the rothamsted trap gave more uniform and consistent samples and performed better in the tropical conditions than the robinson trap . there were no significant differences between the overall measurement of diversity for the group geometroidea between the two trap designs although the robinson trap was much more erratic from night to night . the total catch of non - geometroidea moths was remarkably similar in the two trap types . the choice of appropriate light - trap designs for biodiversity studies in tropical rainforest is discussed .
) seasonality and diversity of macrolepidoptera in two lowland sites in the dumoga - bone national park , sulawesi utara . pp .
) insects as indicators of land - use change : a european perspective , focusing on moths and ground beetles . pp .
) describing and comparing high invertebrate diversity in tropical forest – a case study of small moths in borneo . pp .
jelebu district , negeri sembilan , forest resource reconnaissance survey of malay , report no . 21
) flying in the face of change : the rothamsted insect survey . pp .
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the tetrapodomorpha ( also known as choanata ) are a clade of vertebrates consisting of tetrapods ( four-limbed vertebrates ) and their closest sarcopterygian relatives that are more closely related to living tetrapods than to living lungfish .

advanced forms transitional between fish and the early labyrinthodonts , such as tiktaalik , have been referred to as " fishapods " by their discoverers , being half-fish , half-tetrapods , in appearance and limb morphology .

the tetrapodomorpha contains the crown group tetrapods ( the last common ancestor of living tetrapods and all of its descendants ) and several groups of early stem tetrapods , which includes several groups of related lobe-finned fishes , collectively known as the osteolepiforms .

the tetrapodamorpha minus the crown group tetrapoda are the stem tetrapoda , a paraphyletic unit encompassing the fish to tetrapod transition .

among the characters defining tetrapodomorphs are modifications to the fins , notably a humerus with convex head articulating with the glenoid fossa ( the socket of the shoulder joint ) .

another key trait is the internal nostril or choana .

most fish have two pairs of nostrils , one on either side of the head for incoming water ( incurrent nostrils ) and another pair for outgoing water ( excurrent nostrils ) .

early tetrapodomorphs such as kenichthys had excurrent nostrils that had migrated to the edge of the mouth .

in later tetrapodomorphs , including tetrapods , the excurrent nostril is positioned inside the mouth , where it is known as the choana .

tetrapodomorph fossils are known from the early devonian onwards , and include osteolepis , panderichthys , kenichthys and tungsenia .

tetrapodomorpha

jarvik , 1985 ( sarcopterygii , tetrapodomorpha ) from east greenland . journal of vertebrate paleontology 28 : 637–655 .
clément , g . 2002 . large tristichopteridae ( sarcopterygii , tetrapodomorpha ) from the late famennian evieux formation of belgium . palaeontology , 45 , 577–593 .
johanson z , ahlberg pe ( 2001 ) devonian rhizodontids and tristichopterids ( sarcopterygii ; tetrapodomorpha ) from east gondwana . transactions of the royal society of edinburgh : earth sciences 92 : 43–74 .
tetrapodomorpha was named by ahlberg ( 1991 ) . it is extant . it was reranked as the subclass tetrapodomorpha by betancur - r et al . ( 2013 ) . it was assigned to rhipidistia by ahlberg ( 1991 ) ; to sarcopterygii by daeschler et al . ( 2006 ) ; to dipnotetrapodomorpha by long ( 2011 ) ; and to dipnotetrapodomorpha by betancur - r et al . ( 2013 ) .
snitting , d . 2008 . a redescription of the anatomy of the late devonian spodichthys buetleri jarvik , 1985 ( sarcopterygii , tetrapodomorpha ) from east greenland . journal of vertebrate paleontology , 28 , 637–655 .
johanson , z . and ahlberg , p . e . 2001 . devonian rhizodontids and tristichopterids ( sarcopterygii ; tetrapodomorpha ) from east gondwana . transactions of the royal society of edinburgh , earth sciences , 92 , 43–74 .
clement , g . , snitting , d . , ahlberg , p . 2009 . a new tristichopterid ( sarcopterygii , tetrapodomorpha ) from the upper famennian evieux formation ( upper devonian ) of belgium . palaeontology , 52 , 4 , 823–836 .
sarcopterygii [ 21 ] ; rhipidistia [ 22 ] , [ 23 ] ; tetrapodomorpha [ 24 ] ; eotetrapodiformes [ 13 ] ; tinirau clackae gen . et sp . nov . urn : lsid : zoobank . org : pub : 5dee6139 - 42e1 - 4995 - bab2 - 5e0461aa57a0 .
martin d . brazeau ; a new genus of rhizodontid ( sarcopterygii , tetrapodomorpha ) from the lower carboniferous horton bluff formation of nova scotia , and the evolution of the lower jaws in this group . canadian journal of earth sciences ; 42 ( 8 ) : 1481–1499 . doi : urltoken
the tetrapodomorpha are defined as londoners > lungfish . basally , they differ little from the basic rhipidistian pattern . the synapomorphies of the group , as determined by cloutier & ahlberg ( 1996 ) , are relatively small matters of head and hand . in the head , the pineal foramen is open . the parasymphysial tooth whorl is lost , and the vomers meet on the midline of the palate . something is clearly going on with the nares , but there is much disagreement about exactly what .
tristichopterids ( sarcopterygii , tetrapodomorpha ) form a monophyletic group of exclusively devonian fishes . this thesis consists of descriptions of new material of tristichopterids and closely related taxa , as well as new interpretations and descriptions of previously figured material . redescribed specimens were originally figured as far back as 1861 , and publications as old as this are almost always of limited use as anatomical and systematical references , in addition to being difficult to acquire . the possibility of using new techniques and new theoretical frameworks also provides good justification for taking a second look at such specimens . in the case of this thesis , this includes the use of computed tomography scanning methods , and the cladistic approach to describing the interrelationships of taxa .
tetrapodomorpha here defines total - group tetrapods , and i restrict the use of the term tetrapod to the crown - group . i use the monophyletic definition of elpistostegalia [ 3 ] , [ 26 ] to refer to the clade consisting of panderichthys and crownward taxa . moreover , following from the phylogenetic result presented below , i use canowindridae as a stem - based name to refer to the clade constituting marsdenichthys , canowindra , koharalepis , and beelarongia , use the stem - based megalichthyiformes [ 13 ] to reference the formerly paraphyletic ( here recovered monophyletic , see supplementary information ) ‘osteolepidids’ , and apply the stem - based tristichopteridae to define any taxon more closely related to tristichopterus than to elpistostege . in turn , i use ‘osteolepiform’ to encapsulate the grade of tetrapodomorph that includes canowindrids + megalichthyiforms + tristichopterids , and eotetrapodiformes [ 13 ] as a node - based definition to refer to tristichopterids and elpistostegalians . because of the curious morphology and phylogenetic position of the newly described taxon , i avoid calling this animal an elpistostegalian , and let future studies confirm or refute the phylogenetic hypothesis presented here . in addition , following from the revised phylogenetic placement of platycephalichthys bischoffi [ 13 ] , i refer to this taxon by its name only , as opposed to calling it a tristichopterid or an elpistostegalian .
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lu , jing ; zhu , min ; long , john a . ; zhao , wenjin ; senden , tim j . ; jia , liantao ; qiao , tuo
( c ) 2012 : nature publishing group , a division of macmillan publishers limited . all rights reserved .
recent discoveries of advanced fish - like stem - tetrapods ( for example , panderichthys and tiktaalik ) have greatly improved our knowledge of the fin - to - limb transition . however , a paucity of fossil data from primitive finned tetrapods prevents profound understanding of the acquisition sequence of tetrapod characters . here we report a new stem - tetrapod ( tungsenia paradoxa gen . et sp . nov . ) from the lower devonian ( pragian , ~ 409 million years ago ) of china , which extends the earliest record of tetrapods by some 10 million years . sharing many primitive features with stem - lungfishes , the new taxon further fills in the morphological gap between tetrapods and lungfishes . the x - ray tomography study of the skull depicts the plesiomorphic condition of the brain in the tetrapods . the enlargement of the cerebral hemispheres and the possible presence of the pars tuberalis in this stem - tetrapod indicate that some important brain modifications related to terrestrial life had occurred at the beginning of the tetrapod evolution , much earlier than previously thought .
discover a faster , simpler path to publishing in a high - quality journal . plos one promises fair , rigorous peer review , broad scope , and wide readership – a perfect fit for your research every time .
the origin of terrestrial vertebrates represents one of the major evolutionary and ecological transformations in the history of life , and the established timing and environment of this transition has recently come under scrutiny . the discovery and description of a well - preserved fossil sarcopterygian ( fleshy - limbed vertebrate ) from the middle devonian of nevada helps to refine and question aspects of the temporal and anatomical framework that underpins the tetrapod condition . this new taxon , tinirau clackae , demonstrates that substantial parallelism pervaded the early history of stem - tetrapods , raises additional questions about when digited sarcopterygians first evolved , and further documents that incipient stages of the terrestrial appendicular condition began when sarcopterygians still retained their median fins and occupied aquatic habitats .
citation : swartz b ( 2012 ) a marine stem - tetrapod from the devonian of western north america . plos one 7 ( 3 ) : e33683 . urltoken
editor : andrew a . farke , raymond m . alf museum of paleontology , united states of america
copyright : © 2012 brian swartz . 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 gregory family and several generous anonymous donors . no additional external funding received for this study . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
the origin and early evolution of tetrapodomorphs ( total - group tetrapods ) has been firmly established by numerous studies over the last two decades [ 1 ] – [ 7 ] . however , knowledge of the interrelationships among fish - like ‘osteolepiform’ - grade taxa and the earliest elpistostegalians has remained elusive [ 8 ] – [ 11 ] . phylogenetic analyses have reinforced hypotheses of ‘osteolepiform’ paraphyly and parallelism among devonian stem - tetrapods , but lack of robust statistical support for particular topologies has limited our knowledge of branching and divergence in these early lineages [ 8 ] , [ 12 ] . few studies recover support for larger clades within the ‘osteolepidids’ [ 13 ] , and several establish the close relationship of tristichopterids and elpistostegalians with a robust sister relationship between panderichthys and early digited forms [ 3 ] , [ 12 ] , [ 14 ] . however , no new taxa so far known document the assembly of traits leading from tristichopterids to elpistostegalians .
the discovery of a new stem - tetrapod from the middle devonian of western north america helps to fill this gap and provides a stronger phylogenetic backbone upon which future studies can build . the new material includes several specimens from marine sediments and represents an animal with numerous elpistostegalian apomorphies , yet also many symplesiomorphies , suggesting that early tetrapodomorph features have a more crownward distribution than previously considered . this mélange of characters extends ancestral tetrapodomorph traits across the early history of the first digited forms , and as part of a phylogenetic hypothesis speaks to the length of current ghost ranges implied by the early middle devonian zachełmie ( polish ) trackways [ 15 ] . however , considering the late middle devonian age of this taxon , its congruence with the stratophylogenetic records of other stem - tetrapods , and the phylogenetic distribution of locomotor gaits among crown - group sarcopterygians , questions about when the first digited sarcopterygians first evolved should be considered a more open question than what a strict reading of the trace fossil record might imply .
the material was discovered and excavated in the mid - late 1970s by university of california , berkeley paleontologist joseph t . gregory and his graduate students at a field site in northeastern nevada known as red hill i . the red hill i beds are a series of silty limy mudstones alternating with thick - bedded limestones , bounded below and above by the denay and devils gate formations , respectively [ 16 ] . this university of california museum of paleontology field site ( ucmp v74084 ) is located in the northern simpson park mountains in eureka county , nevada . conodont biostratigraphy places red hill i in the lower klapperina disparilis zone [ 16 ] , [ 17 ] , the late givetian stage of the middle devonian . the described sarcopterygian material was recovered from levels 8–12 of the roughly 1 . 5 m thick sequence of vertebrate - bearing beds immediately above the denay limestone ( figure 1 ) .
geographic location and stratigraphic position of the red hill i field site ( ucmp v74084 ) in eureka co . , nevada , usa .
black patterning within eureka county represents exposed devonian outcrops . stars represent where the fossil material was collected . red hill i section courtesy of h . - p . schultze .
the fauna and geology indicate that the sedimentary rocks comprising red hill i were deposited in a marine environment . cnidarians such as conulariids , a clade known elsewhere only from marine strata [ 18 ] , are preserved in levels 21 - 5 ( figure 1 ) . moreover , the widespread deposition of limestone and shale along the western margin of laurentia suggests that the regional geology of the northern simpson park range represents an open marine paleoenvironment [ 19 ] , and in particular the outer continental shelf [ 16 ] , [ 20 ] . trace fossils preserved between levels one and two suggest a short - term nearshore paleoenvironment [ 20 ] .
tinirau ( tea - knee - / r / áu ) is a character of legend in polynesian culture and traces to islands located at approximately the same latitude as nevada during the middle devonian . according to the rarotonga and mangaia islanders , tinirau was a half - man , half - fish lord of the ocean creatures [ 25 ] . the specific name clackae honors the cambridge palaeontologist and former advisor jenny clack , for her contributions to our understanding of the earliest digited sarcopterygians .
( a ) ucmp 118605 , holotype , in dorsal , lateral and ventral view . see main text for details . right is anterior . scale bar equals 10 cm ; ( b ) complete restoration ; preserved elements outlined in black , inferred margins outlined in dashed black , hypothesized elements outlined in gray . see methods section for anatomical abbreviations . note the reduced postaxial fibular processes on the fibulae ( fib . p ) .
this description is based on six specimens ( ucmp 117884 , 118283 , 118605 , 123135 , 190998 , 190999 ) from a single locality . all specimens preserve complete or partial skull remains . two specimens ( ucmp 118605 , 190999 ) preserve postcrania and appendicular elements in some degree of articulation . specimens ucmp 118283 and 123135 were preserved in association with one another , adjacent on the same small block but not articulated . not all specimens of tinirau preserve every available character state , but consistent features among all specimens indicate that they represent a single taxon . these features include : elongate glenoid fossae ( ucmp 118065 , 190999 ) , reduced posterior processes on the maxillae ( ucmp 118065 , 190999 ) , fused parietals ( ucmp 117884 , 118238 , 118065 , 190999 ) , fused anterior tectals and lateral rostrals ( ucmp 11784 , 118283 ) , a row of non - fang teeth on the elongate posterior coronoids ( ucmp 118605 , 123135 ) , and similar proportions and dentitions of the dermopalatines and entopterygoids ( ucmp 190998 , 190999 ) .
usa , eureka co . , nevada , simpson park mountains north of the denay valley , ucmp locality v74084 .
lower disparilis conodont zone of the red hill i beds , immediately above the denay formation .
an eotetrapodiform sarcopterygian distinguished from known tristichopterids by ( i ) an elongate posterior jugal process ( figures 2 , s1 ) , ( ii ) a dermal cheek plate with fused squamosal , preopercular , and quadratojugal elements ( figures 2 , s1 ) , ( iii ) deep tongue - and - groove embayments along the posteromedial margins of the intertemporals ( figures 3a , s2 ) , ( i v ) fused anterior tectals with lateral rostrals ( figures 3a , s3 ) , ( v ) medially straight anterior parietal margins in the unfused skull - table ( figure s3 ) , ( vi ) a fused ethmoid skull - table in larger specimens—i . e . , later ontogenetic stages ( figures 2 , 3a , s1 , s4 ) , ( vii ) ectopterygoids that contribute to the subtemporal fossae ( figure 3b ) , ( viii ) splenials that remain unsutured to the prearticular ( figure 3c ) , and ( ix ) reduced postaxial fibular processes ( figures 2 , s1 , s6 ) . moreover , it is differentiated from elpistostegalians by ( i ) facially positioned anterior nostrils ( figure 3a ) , ( ii ) a ( inferred ) lateral component to the ventral orbital margins ( figure 2 , s1 ) , ( iii ) the presence of a median postrostral ( figure s3 ) , ( iv ) the absence of frontal bones ( figures 2 – 3 , s1 , s3 , s4 ) , ( v ) the presence of a ( anteriorly positioned ) postspiracular ( figure s4 ) , ( vi ) long posterior vomerine processes ( figure s2 ) , ( vii ) an absence of jugal - quadratojugal contact ( figures 2 , s1 , 2 ) , ( viii ) a small scapulocoracoid ( figures 3c , s7 ) , and ( ix ) round body scales ( figure 3c ) .
( a - i ) ucmp 117884 , ethmoid skull . anterior is toward the top of the page . scale bar equals 2 cm ; ( a - ii ) dorsal skull reconstruction with infilled gray ethmoid region following from ( a - i ) ; ( b ) left palatal fragment of ucmp 190998 . right is anterior . scale bar equals 5 cm ; ( c ) skull , partial shoulder , and interpretive drawing of ucmp 190999 . uniform stipple covering distal jaw elements indicate unexposed portions of the specimen still covered by bioplastic ; similarly , the dotted line posterior to the parasphenoid ( psph ) notes the division between ethmoid and oticoccipital regions recovered from x - ray imaging . anterior is toward the top of the page . scale bar equals 5 cm . see methods section for anatomical abbreviations . note the elongate glenoid fossa ( gle ) on the left scapulocoracoid ( sco ) .
the snout of tinirau has one pair of facially positioned external nostrils as in all tetrapodomorphs except kenichthys and elpistostegalians . however , in tinirau , the nares penetrate a single , fused element consisting of the anterior tectal and lateral rostral ( figure 3a ) . similar to ‘osteolepiforms’ , platycephalichthys , and elpistostegalians less crownward than ventastega , the premaxilla forms a broad part of the choanal margin ( figure s2 ) . moreover , and differing from ventastega and acanthostega , a single median postrostral and several nasal bones create a solid snout lacking a dorsal fontanelle ( figures 2a , 3a , s1 , s3 ) .
the anterior skull roof of tinirau is plesiomorphic among tetrapodomorphs : about 25 % of the skull extends anterior to the mid - orbital margins ( figures 2a , 3a , s1 ) . such proportions are more similar to those of rhizodonts and canowindrids than to those of other eotetrapodiforms . the anterior - most paired roofing bones are the parietals , which are pierced by a pineal foramen that lies posterior to the orbits in larger specimens , or later ontogenetic stages ( figures 3a , s3 ) . this condition is similar to early diverging ‘osteolepiforms’ such as koharalepis , canowindra , and gyroptychius , and later - diverging tristichopterids more phylogenetically distal than eusthenopteron . a functional dermal intracranial joint is unknown considering the tongue - and - groove articulations of the intertemporal and supratemporal bones that span this region . however , because the skull tends to be preserved in two parts , with the symplesiomorphic condition at least across the parietal / postparietal region , such a ‘joint’ is scored as present in tinirau ( figures 2a , 3a , s1 , s4 ) . the condition in tinirau is thus either autapomorphic ( considering that dermal suturing in panderichthys involves only the parietals and postparietals ) or ‘intermediate’ because of the simultaneous suturing and simple abutment found across its dermal intracranial division . interestingly , platycephalichthys also has posteriorly recessed intertemporals suggesting a similar intracranial configuration [ 27 ] .
the postparietal shield is not extremely wide posteriorly , as in canowindrids , nor do the parietals narrow to a point caudally , as in rhizodonts . instead , the tabulars extend to the posterior margin of a postparietal shield that is approximately as wide as the ethmoid , a condition akin to that seen in tristichopterids , panderichthys , tiktaalik , and ventastega ( figure s4 ) . lateral to the tabular resides a postspiracular ( = extratemporal ) situated in the plesiomorphic anterior position , similar to the condition in devonian tetrapodomorphs except tristichopterids phylogenetically distal of spodichthys ( figure s4 ) . the postspiracular is lost in known elpistostegalians .
surrounding the orbit , the anterior and posterior supraorbitals ( = prefrontals and postfrontals ) are of similar size and contact one another anterior to the mid - orbital margin . the posterior supraortbitals do not extend anterior to the orbits , similar to the condition in other devonian tetrapodomorphs except a few late - diverging tristichopterids ( figures 2a , s1 , s3 ) . the lacrimal and jugal meet approximately at the mid - ventral orbital margin where , unlike in mandageria and eusthenodon , the postfrontal and lacrimal do not make contact ( figures 2a , s1 ) . moreover , unlike in elpistostegalians , the squamosal ( here , bound up in a fused cheek plate ) precludes abutting of the jugal and quadratojugal ( figures 2 , s1 , s4 ) . it is not known directly if the postorbital contributes to the orbit of tinirau , but based on the topology of this element and neighboring bones in ucmp 118605 , it is inferred to make a minor contribution ( figures 2a , s1 ) .
the jaws of tinirau are characteristically eotetrapodiform in form , although contain a unique combination of plesiomorphic and apomorphic traits . the premaxillary teeth are all of similar size as in early diverging tristichopterids and elpistostegalians ( figure s2 ) . however , the maxilla lacks a posterodorsal process , a state shared with platycephalichthys and elpistostegalians such as panderichthys on crownward , but also with derived tristichopterids such as cabonnichthys and mandageria ( figures 2a , s1 , s4 ) . dentary fangs are present , similar to platycephalichthys and elpistostegalians , though this character is also known in rhizodonts , megalichthyids , and derived tristichopterids ( figure 3c ) . the posterior coronoid is much longer than the anterior two coronoids , yet only carries one fang pair followed by a row or 5 + medium - sized teeth ( figure s5 ) . this state combination is not present in any tristichopterid , and only shared with platycephalichthys and early elpistostegalians such as panderichthys . in other words , tristichopterids with long posterior coronoids also bear two posterior fang pairs , and those tristichopterids with one fang pair do not have very long posterior coronoids . a distinct meckelian groove is visible in the lower jaw of ucmp 190999 , and similar to the condition in non - elpistostegalian tetrapodomorphs , it bears an ossified posterior meckelian region separating the prearticular / angular contact ( figure 3c ) .
the operculogular elements in ucmp 190999 are similar in shape and proportion to those of other devonian ‘osteolepiforms’ , and therefore are not diagnostic of a physical neck ( i . e . , a discrete , disconnecting region ) between the shoulders and head ( figures 2a , s1 , s4 ) . similar to kenichthys and platycephalichthys , a large preoperculum is sutured to the squamosal in a cheek plate and is also visible in visceral view in ucmp 118605 and 190999 ( figures 2a , s1 , s4 ) . the spiracular notch is not well - preserved , but judging from the narrow space between the squamosal and postparietal shield , it is inferred to be small and thus more like the condition in most ‘osteolepiforms’ rather than to that of gogonasus and elpistostegalians ( figures 2a , 3a - ii , s1 ) . the presence and size of a median gular remain unknown .
the neurocranium is plesiomorphic in many ways , although it shares some similarities with those of tristichopterids . a fully ossified ethmoid extends below a narrow tectum orbitale and articulates with its posterior otic - occipital counterpart via an endoskeletal intracranial joint . in turn , a basicranial fanestra spans this division ( figures 3c ) . these states are present in all devonian tetrapodomorphs except for kenichthys and taxa crownward of tiktaalik . by contrast , tinirau shares with tristichopterids a relatively anterior ventral hyomandibular facet ( figure 3c ) . in other words , this state is generally considered to diagnose tristichopterids , but is here reconstructed to be either convergent among these taxa , or to ancestrally diagnose eotetrapodiforms only primitively .
the cephalic branches of the sensory canal system are typical of most other devonian tetrapdomorphs , although tinirau retains a few traits—such as the postorbital junction of supra - and infraorbital canals , a line of continuous pores that comprise the mandibular canal , and a surangular pitline—that are otherwise lost in taxa crownward of tiktaalik and acanthostega ( figures 2a , s1 , s5 ) . as in glyptopomus , marsdenichthys , tristichopterids , platycephalichthys , panderichthys , and tiktaalik , the sensory canals course through a tuberculate dermal skeleton that lacks the starburst ornamentation characteristic of the first digit - bearing elpistostegalians ( figures 3a - i , 3c , s3 , s4 , s5 ) . such elements also lack the thick ‘shine’ characteristic of cosmine - covered sarcopterygians such as megalichthyiforms .
the shoulder is typically tetrapodomorph in form , but it bears a few differences from those of key taxa . the anterior median extrascapular margin is “long” and therefore unlike those of canowindrids and mandageria ( figures 2a , 3a - ii , s1 , s4 ) . a postbranchial lamina is present on the cleithrum ( figures 2a , s1 ) , although posttemporals , supracleithra , anocleithra , and an interclavicle are not preserved . unlike in elpistostegalians such as panderichthys and tiktaalik , a small scapulocorocoid is elevated from the ventral plane formed by the clavicles . however , the glenoid is relatively elongate and bears a medial ‘accessory’ region that is less reflexed than the condition seen in megalichthyiforms such as medoevia and tristichopterids such as eusthenopteron ( figures 2a , 3c , s1 ) . although the humerus is crushed , judging from the shape of the glenoids , it appears that the convex caput humeri retains less of the oblate shape than is typical of ‘osteolepiforms’ . such an elongate condition is more characteristic of elpistostegalians .
paired appendages are only preserved in ucmp 118605 ( figures 2a , s1 ) . the left humerus is crushed and situated below the cleithrum , but it articulates with the rest of a well - preserved pectoral limb . the right humerus is missing , but the elongate glenoid and distal pectoral elements remain . the pectoral limb is symplesiomorphic , and generally similar to the ‘osteolepiform’ condition . as in ‘osteolepiforms’ and elpistostegalians such as panderichthys and tiktaalik , the ulna is about half as long as the radius and articulates with an ulnare and intermedium . as in ‘osteolepiforms’ , the ulnare retains a postaxial process and only articulates with two additional distal radials . proximal lepidotrichia are about three times longer than more distal ones ( figures 2a , s1 ) .
caudally , the pelvis articulates with a femur that is preserved in association with the acetabulum , despite the disassociation of distal elements ( figures 2a , s1 , s6 ) . as in eusthenopteron , the right and left disarticulated fibulae bear preaxial radial facets positioned about one half - step proximal to their postaxial counterparts . however , and unlike in gooloogongia and eusthenopteron , the postaxial fibular process is highly reduced and not simply the ‘fibula - equivalent’ of the condition seen in the ulnare . interestingly , the pelvic limb of panderichthys also displays a similar ‘lip’ overhanging the postaxial edge of the fibulare ( [ 28 ] , figure 1 , pg . 1146 ) .
the vertebral elements are preserved in near complete articulation , and are known only from ucmp 118605 ( figures 2a , s1 ) . paired intercentra are visible entirely in part / counterpart , and stout non - imbricate ribs radiate laterally , immediately posterior to the cleithrum . the axial skeleton proceeds through a left twist at ∼90° around mid - body , and posterior to the pelvis folds over itself so that the distal tip of the heterocercal caudal fin skeleton comes to face the more anterior ( dorsal ) neural spines . paired pleurocentra are not preserved and are presumed to have been cartilaginous . there is no evidence for dorsal fin radials , although dorsal fins are hypothesized to have been present . by contrast , a dissociated anal fin basal and radial are preserved immediately dorsal to the caudal fin . the notochordal canal is visible and arches dorsally through the neural and haemal arches of the caudal fin skeleton ( figures 2a , s1 ) .
a phylogenetic analysis using paup [ 29 ] recovered a single most parsimonious tree . a bayesian analysis [ 30 ] , [ 31 ] of the same data provided an additional metric . there are no major polytomies among the ‘osteolepiform’ grade taxa . instead , the major clades , rhizodontidae , canowindridae , megalichthyiformes , and tristichopteridae form successive sister taxa to more crownward groups . tinirau emerges as the sister to platycephalichthys and elpistostegalians , one step crownward of tristichopterids ( figure 4 ) .
analysis includes 46 taxa and 204 characters . tree length = 454 , consistency index = 0 . 5572 , retention index = 0 . 8481 ; consistency index excluding the four autapomorphic ( uninformative ) characters = 0 . 5532 , retention index = 0 . 8481 . numbers corresponding to respective nodes represent : bremer decay value / bayesian posterior probability . ghost ranges are calibrated after the early middle devonian ( eifelian ) zachełmie tracks ( [ 15 ] and “scenario 1” from friedman and brazeau ( 2011 ) . tetrapodomorphs include all taxa that are not total - group lungfishes . rhizodonts are in green , canowindrids are in yellow , megalichthyiforms are in blue , tristichopterids are in purple , devonian elpistostegalians are in red , and carboniferous elpistostegalians are in orange . the character list and data matrix are available as supplementary information .
this phylogenetic hypothesis implies that , ( 1 ) tristichopterid synapomorphies ( see text s1 ) have evolved in parallel during the early history of eotetrapodiforms ; and ( 2 ) the 18 + elpistostegalian synapomorphies are cut in half ( see text s1 ) as taxa such as tinirau and platycephalichthys fill the graduated history of the tetrapod stem . this is predicted by current evidence , especially with the recent finding of marine , digit - bearing tracks that predate the earliest elpistostegalian body fossils by 10 ma [ 15 ] . the discovery of tinirau fills a phylogenetic gap missing from previous discoveries even though its stratigraphic range conforms with the timing of the body fossil record . yet because ‘genus’ - level preservation rates for devonian tetrapodomorphs are an order of magnitude lower than ‘species’ - level rates for groups considered to have dense records [ 32 ] , the stratigraphic range of tinirau is not surprising . thus , when combined with the age of the trackways data , the late middle devonian ( givetian ) age of tinirau , its phylogenetic position as stem to the first digited forms , and its many symplesiomorphies may suggest a rich , yet undiscovered early tetrapodomorph record .
however , the phylogenetic distribution of potential sarcopterygian trackmakers does bring into question whether digited tetrapodomorphs even produced the zachełmie trackways . digit - bearing molds are preserved alongside continuous trackways but the ‘digits’ themselves are known only from isolated prints . crown - group coelacanths , lungfishes , and tetrapods are known to engage in trotting gaits [ 33 ] – [ 35 ] , and thus suggest substrate - based locomotor abilities in stem - tetrapods as well . moreover , recent work [ 36 ] has shown that african lungfish using a bipedal pelvic - driven gait can produce the three trackways patterns known from the zachełmie quarry : ( 1 ) alternating doublets ; ( 2 ) alternating singlets ; and ( 3 ) opposite , ladder - like prints . considering this range of potential explanations , the non - congruence of continuous digited prints with trackways patterns , and the increasingly strong stratophylogenetic congruence in the stem - tetrapod body fossil record , it might be wise to approach questions about timing of origins with a pluralistic eye and a bit of additional skepticism .
questions about palaeoenvironment are more complicated , but tinirau ' s marine preservation is consistent with the marine influenced environments of the zachełmie tracks and other closely related taxa [ 7 ] , [ 37 ] , [ 38 ] , although likely not with others [ 3 ] , [ 39 ] .
overall , the skeleton of tinirau retains many ‘fish - like’ traits , but they are combined with a suite of elpistostegalian apomorphies . because the utility of many of these characters remains obscure , here i elaborate on two traits that emerge as relevant to current discussions in tetrapodomorph evolution : the origins of the shoulder and pelvic limbs in the first digit - bearing elpistostegalians .
as in tristichopterids such as eusthenopteron and elpistostegalians such as panderichthys , the shoulder of tinirau retains the full osteichthyan complement of dermal and endochondral components . however , despite these general similarities , its glenoid is anteroposteriorly elongate and in this respect more similar to the condition found in panderichthys , acanthostega , and juvenile specimens of tiktaalik ( figure 5a ) . interestingly , glenoids in the largest tiktaalik specimens are less elongate than those of smaller individuals [ 3 ] , [ 40 ] ( personal observations ) and may reflect ontogenetic changes . nonetheless , despite this possible autapomorphy , elongate glenoids in the first digited taxa correlate with parallel changes observed in the flattening of the caput humeri [ 2 ] , [ 41 ] – [ 45 ] . although the glenoids in medoevia , eusthenopteron , tinirau , and panderichthys have a strong posterior component , fossae in the former two taxa are more oblate than the condition present in the latter forms . this reinforces the hypothesis that mosaic changes in the pectoral limb began proximally before the distal portions acquired a more characteristic tetrapod - like morphology [ 46 ] .
glenoids are illustrated in posterior view and highlighted in blue , fibulae are highlighted in green . the glenoid of tiktaalik is depicted from two different perspectives , posterior view ( above ) and posteroventral view ( below ) . the largest tiktaalik specimens are more oblate , which may be autapomorphic relative to the condition in more crownward devonian and carboniferous taxa . the glenoid of panderichthys was based on the shape of its caput humerus . see text for additional details . the in - plane glenoid measurement ( height at maximum extent divided by maximum length ) diagnoses an elongate glenoid fossa : medoevia = 0 . 60 ; eusthenopteron = 0 . 60 ; tinirau = 0 . 42 ; panderichthys = 0 . 48 ; tiktaalik = 0 . 44 ; acanthostega = 0 . 45 .
the femur , tibia , and fibula represent the only pelvic elements preserved in tinirau , but they share an interesting similarity with panderichthys , the only non - digit bearing elpistostegalian from which reasonable pelvic material is known [ 28 ] . one major difference between the fibulae of a rhizodont ( e . g . , gooloogongia ) or a tristichopterid ( e . g . , eusthenopteron ) and an elpistostegalian ( e . g . , panderichthys ) is that the postaxial process in panderichthys is reduced to a mere lip or overhang bordering the posterior margin of the distal fibulare [ 4 ] , [ 28 ] , [ 47 ] ( figures 4b , s6 ) . in this respect , the lack of a prominent postaxial process in the fibula of tinirau is more similar to the condition observed in crownward taxa . this pattern underscores previous phylogenetic reconstructions of the appendicular skeleton in which conventional crown group limb characteristics first originate in the pelvic fins [ 48 ] .
the new phylogeny also helps to displace eusthenopteron as our iconic surrogate piscine ‘ancestor’ . eusthenopteron shares with other tristichopterids a sequence of traits that nest it well within tristichopterids and not immediately along the tetrapod stem ( figure 6 ) . instead , this result builds upon the work of coates and friedman ( 2010 ) , whereby tinirau and platycephalichthys fill this position and provide an anatomical record on the transition to land . these taxa spread primitive tetrapodomorph traits along the early history of elpistostegalians , raise additional questions about when digited sarcopterygians first evolved , and fill a gap between tristichopterids and the first digited sarcopterygians in interesting and unexpected ways .
character states supporting this topology : ( 1 ) a parasymphyseal plate not sutured to the anterior coronoid ; ( 2 ) posterior coronoids longer than more anterior coronoids ; ( 3 ) 33–40 % of the dermatocranium anterior to the orbits ; ( 4 ) a posteriorly displaced postspiracular ; ( 5 ) posterior coronoids one third longer than the anterior coronoids ; ( 6 ) two ectopterygoid fang pairs ; and ( 7 ) a diphycercal caudal fin .
204 morphological characters were used to assess the phylogenetic position of the new taxon described above ( tinirau clackae ) relative to other early tetrapodomorphs . primary character sources [ 3 ] , [ 8 ] , [ 11 ] , [ 13 ] , [ 49 ] – [ 51 ] are indicated parenthetically following each character description in the supplementary information . numbers following the citations refer to the character number in the original source . characters modified from their original source are noted where applicable . very few characters are shared between this analysis and coates and friedman ( 2010 ) ; this was intentional with the goal of assessing how largely independent data sets converge on a similar result .
characters were polarized by comparison to outgroup taxa such as porolepis , glyptolepis , powichthys , youngolepis , diabolepis , and dipterus . these taxa were selected because they represent a range of total - group lungfish that are known from reasonable material , are well studied , and generally accepted as sister to total - group tetrapods .
characters were coded based on a combination of published descriptions , specimen illustrations , and firsthand examination of fossil material . care was taken to avoid simply recycling codings in the published literature . specimens from the following museums were examined , and are noted following each taxon in the supplementary information : australian museum , sydney ( amf ) , australian national university ( anu ) , geologisk museum , copenhagen , denmark ( mguh ) , latvian museum of natural history ( ldm ) , muséum national d ' histoire naturelle , paris ( mnhn ) , museum victoria , melbourne , australia ( nmv ) , the natural history museum , london ( mnh ) , palaeontological institute of the russian academy of sciences , moscow ( pin ) , national museums of scotland ( nms ) , nunavut fossil vertebrate collection ( nufv ) , swedish museum of natural history , stockholm ( nr ) , university of california museum of paleontology ( ucmp ) , university museum of zoology cambridge ( umzc ) .
the data matrix was subjected to a maximum parsimony analysis in the software package paup 4 . 0b10 [ 29 ] and a bayesian analysis using the software package mr . bayes 3 . 2 [ 30 ] , [ 31 ] . all characters were assigned an equal weight , multistate characters were run unordered , and a heuristic search algorithm was used in paup to search for the shortest networks—rooted on porolepis , glyptolepis , powichthys , youngolepis , diabolepis , and dipterus . bremer decay indices were calculated using paup [ 29 ] and tnt [ 52 ] , [ 53 ] , and bayesian posterior probabilities were calculated with mr . bayes following an analysis that included 500 , 000 mcmc generations , sampling every 1 , 000 generations , and with 20 samples discarded as burnin . character evolution was examined in macclade [ 54 ] , which was also used to produce the character state distributions in the supplementary information .
the material was prepared by an acid immersion procedure including baths of 30 % formic acid or 10–20 % acetic acid for 10–48 hours , followed by washing in running water for one month , and air - drying for 12–24 hours . exposed elements were strengthened with glyptal or duco cement . the three ucmp specimens 117884 , 118283 , and 123125 were studied 30 years ago by former uc berkeley graduate student john reed , although never published [ 55 ] . because so much has changed in the record , systematics , and nomenclature of stem - tetrapods , it was necessary to redo the study completely .
the electronic version of this document does not represent a published work according to the international code of zoological nomenclature ( iczn ) , and hence the nomenclatural acts contained in the electronic version are not available under that code from the electronic edition . therefore , a separate edition of this document was produced by a method that assures numerous identical and durable copies , and those copies were simultaneously obtainable for the purpose of providing a public and permanent scientific record , in accordance with article 8 . 1 of the code . the separate print - only edition is available on request from plos by sending a request to public library of science , 1160 battery street , suite 100 , san francisco , ca 94111 , usa along with a check for $ 10 ( to cover printing and postage ) payable to ‘public library of science’ . in addition , this published work and the nomenclatural acts it contains have been registered in zoobank , the proposed online registration system for the iczn . the zoobank lsids ( life science identifiers ) can be resolved and the associated information viewed through any standard web browser by appending the lsid to the prefix ‘ urltoken ’ . the lsid for this publication is : urn : lsid : zoobank . org : pub : 5dee6139 - 42e1 - 4995 - bab2 - 5e0461aa57a0 tinirau clackae swartz gen . et sp . nov . for the genus , the lsid is : urn : lsid : zoobank . org : act : f459d126 - ad40 - 4f69 - a0c3 - 1b6885e891a7 ; and for the species , the lsid is : urn : lsid : zoobank . org : act : fbd69da1 - 884c - 4a2d - 87c4 - f7f8d85ab376 .
ba . a , anal basal ; urltoken , basal articulation of the basipterygoid process ; basb1 , basibranchial # 1 ; bas . f , basicranial fenestra ; ch , ceratohyal ; clth , cleithrum ; clv , clavicle ; co 3 , posterior coronoid ; de . f , dentary fang ; dpt , dermopalatine ; enpt , entopterygoid ; ept , ectopterygoid ; exsc . l , lateral extrascapular ; exsc . m , median extrascapular ; fe , femur ; fib , fibula ; fib . p , posterior process of the fibula ; gle , glenoid fossa ; gu , lateral gular ; hh , hypohyal ; hu , humerus ; hyo , hyomandibular ; urltoken , hyomandibular articulation ; ic , intercentrum ; int , intermedium ; it , intertemporal ; ju , jugal ; la , lacrimal ; mk , meckelian bone ; mk . grv , meckelian groove ; mx , maxilla ; na . a , anterior naris ; nc . c , notochordal canal ; ns , neural spine ; op , operculum ; urltoken , parietal pitline ; part , prearticular ; pin . f , pineal foramen ; plv , pelvis ; pmx , premaxilla ; po , postorbital ; pop , preoperculum ; pp , postparietal ; pq , palatoquadrate ; psph , parasphenoid ; qj , quadratojugal ; quad , quadrate ; r , radius ; ra , radial ; ra . a , anal radial ; ra . c , caudal radial ; ri , rib ; sang , surangular ; sbm , submandibular ; sca , scale ; sco , scapulocoracoid ; spl , splenial ; st , supratemporal ; st . f , subtemporal fossa ; so . p , posterior supraorbital ; sq , squamosal ; tab , tabular ; te . a / ro . l , anterior tectal + lateral rostral ; tib , tibia ; u , ulna ; ul , ulnare ; uh , urohyal ; vo , vomer . ( l ) or ( r ) refers to left or right when displaced from natural side .
supplementary text . part a : taxa and characters used in the phylogenetic analysis ; part b : taxon - by - character matrix and character optimizations .
close - up of ucmp 118605 and specimen drawing . ucmp 118605 , holotype , in dorsal , lateral and ventral view . see main text for details ; right is anterior . abbreviations : ba . a , anal basal ; clth , cleithrum ; clv , clavicle ; co 3 , posterior coronoid ; exsc . l , lateral extrascapular ; exsc . m , median extrascapular ; fe , femur ; fib , fibula ; fib . p , posterior process of the fibula ; gle , glenoid fossa ; hu , humerus ; ic , intercentrum ; int , intermedium ; it , intertemporal ; ju , jugal ; la , lacrimal ; mx , maxilla ; na . a , anterior naris ; nc . c , notochordal canal ; ns , neural spine ; part , prearticular ; plv , pelvis ; pmx , premaxilla ; po , postorbital ; pop , preopercular ; pp , postparietal ; qj , quadratojugal ; r , radius ; ra , radial ; ra . a , anal radial ; ra . c , caudal radial ; ri , rib ; sca , scale ; sco , scapulocoracoid ; st , supratemporal ; so . p , posterior supraorbital ; sq , squamosal ; tab , tabular ; tib , tibia ; u , ulna ; ul , ulnare ; scale bar equals 10 cm .
ethmoid palatal region and interpretive drawing of ucmp 117884 . anterior is toward the top of the page . abbreviations : urltoken , autopalatine articulation ; urltoken , basal articulation of basipterygoid process ; cho , choana ; ‘cn’ ii , optic nerve ; it , intertemporal , nc , neurocranium ; p . con , processes connectens ; pmx , premaxilla ; pro . f , profundus foramen ; psph , parasphenoid ; vo , vomer ; vo . f , vomerine fang . ‘cn’ is in scare quotes because the optic nerve is not a real cranial nerve but a special - sensory extension of the diencephalon . scale bar equals 5 cm .
ethmoid skull roof and interpretive drawing of juvenile specimen ucmp 118283 . aside from the fusion of the anterior tectal and lateral rostral ( similar to the adult specimen , ucmp 117884 ) , many of the remaining roofing bones are unfused . the snout of this specimen is also proportionally shorter than the adult ( when pineal foramina are aligned ) , suggesting substantial allometric change during ontogeny . in addition , it lacks the recessed tongue - and - groove articulations spanning the dermal intracranial joint , suggesting acquisition later in life . anterior is toward the top of the page . abbreviations : it , intertemporal ; na , nasal ; pa , parietal ; pin . f , pineal foramen ; pmx , premaxilla ; ro . p , median postrostral ; so . a , anterior supraorbital ; soc , supraorbital canal ; te . a / ro . l , ( fused ) anterior tectal / lateral rostral . scale bar equals 5 mm .
skull , partial shoulder , and interpretive drawing of ucmp 190999 . anterior is toward the top of the page . abbreviations : clth , cleithrum ; clv , clavicle ; de , dentary ; exsc . l , lateral extrascapular ; exsc . m , median extrascapular ; gu , lateral gular ; hyo , hyomandibular ; ju , jugal ; la , lacrimal ; mx , maxilla ; op , operculum ; pa , parietal ; part , prearticular ; pop , preoperculum ; pp , postparietal ; psp , postspiracular ; qj , quadratojugal ; ro . p , median postrostral ; sco , scapulocoracoid ; sop , suboperculum ; sq , squamosal ; st , supratemporal ; ta , tabular ; te . a / ro . l , ( fused ) anterior tectal / lateral rostral . ( l ) or ( r ) refers to left or right when displaced from natural side . scale bar equals 5 cm .
lower jaw of ucmp 123135 . ( a ) dorsal view ; ( b ) lateral view and interpretive drawing . left is anterior . abbreviations : add . f , adductor fossa ; ang , angular ; art , articular ; co 1 , anterior coronoid ; co 2 , middle coronoid ; co 3 , posterior coronoid ; co . f , coronoid fang ; de , dentary ; mc , mandibular canal ; pspl , postsplenial ; sang , surangular ; spl , splenial . scale bar equals 10 mm .
close - up of the pelvic region of ucmp 118605 highlighting the reduced postaxial fibular processes . abbreviations : fe , femur ; fib , fibula ; plv , pelvis ; ns , neural spine ; ri , rib . ( l ) or ( r ) refers to left or right . scale bar equals 10 cm .
close - up of the elongate glenoid fossa of ucmp 190999 . abbreviations : clth , cleithrum ; gle , glenoid fossa ; sbm , submandibular ; sco , scapulocoracoid . ( l ) refers to left . scale bar equals 5 cm .
this work would not have been possible without the assistance of several others . i express my gratitude for the help provided by kevin padian , marvalee wake , tony barnosky , ted daeschler , martin brazeau , hans - peter schultze , jenny clack , matt friedman , john long , gavin young , tim senden , ken campbell , brian choo , nick matzke , gaël clément , john reed , zerina johanson , thomas mörs , per ahlberg , catherine boisvert , and marcus wong . i additionally thank kevin padian , marvalee wake , mike coates , david wake , per ahlberg , ted daeschler , and jenny clack for their assistance and thoughts on earlier drafts of this manuscript .
conceived and designed the experiments : bs . performed the experiments : bs . analyzed the data : bs . contributed reagents / materials / analysis tools : bs . wrote the paper : bs .
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vorobyeva ei , schultze h - p ( 1991 ) description and systematics of panderichthyid fishes with comments on their relationship to tetrapods . in : schultze h - p , trueb l , editors . origins of the higher groups of tetrapods : controversy and consensus . ithaca : cornell university press . pp . 68–109 .
ahlberg pe , johanson z ( 1998 ) osteolepiforms and the ancestry of tetrapods . nature 395 : 792–793 .
chang m - m , yu x ( 1997 ) reexamination of the relationship of middle devonian osteolepids : fossil characters and their interpretations . american museum novitates 1–20 .
friedman m , coates mi , anderson psl ( 2007 ) first discovery of a primitive coelacanth fin fills a major gap in the evolution of lobed fins and limbs . evolution & development 9 : 329–337 .
long ja , young gc , holland t , senden tj , fitzgerald emg ( 2006 ) an exceptional devonian fish from australia sheds light on tetrapod origins . nature 444 : 199–202 .
and characteristics of stem tetrapod neurocrania . in : elliott dk , maisey jg , yu x , miao d , editors . morphology , phylogeny and paleobiogeography of fossil fishes . münchen : verlag dr . friedrich pfeil . pp . 389–416 .
niedźwiedzki g , szrek p , narkiewicz k , narkiewicz m , ahlberg pe ( 2010 ) tetrapod trackways from the early middle devonian period of poland . nature 463 : 43–48 .
johnson jg , sandberg ca , poole fg ( 1988 ) early and middle devonian paleogeography of united states and their biostratigraphic responses . in : mcmillan nj , embry af , glass dj , editors . devonian of the world volume i , regional synthesis . calgary : canadian society of petroleum geologists . pp . 161–182 .


istiblennius edentulus , the rippled rockskipper , is a species of combtooth blenny found in coral reefs in the pacific and indian oceans .

it is also commonly known as the rippled blenny , smooth-lipped blenny , toothless blenny , or coral blenny .

males of this species can reach a maximum of 16 cm ( 6.3 in ) tl , while females can reach a maximum of 13.2 cm ( 5.2 in ) sl .

istiblennius edentulus

taxon concept istiblennius _ edentulus last modified 2013 - 11 - 25 10 : 42 : 21 . 399
istiblennius edentulus is of no interest to the fisheries industry , but is commercially collected for use in the aquarium trade ( burgess et . al 1990 ) .
alticops edentulus , alticops oryx , blennius cinereus , blennius edentulus , blennius truncatus , istiblennius edentululus , istiblennius enosimae , istilbennius edentulus , salarias atratus , salarias atrimarginatus , salarias azureus , more . . . salarias diproktopterus , salarias fluctatus , salarias garmani , salarias gilberti , salarias insulae , salarias marcusi , salarias melanocephalus , salarias quadricornis , salarias rechingeri , salarias sindonis , salarias sumatranus , scartella enosimae , scartichthys basiliscus , scartichthys enosimae hide . . .
a rippled rockskipper , istiblennius edentulus , at wooli , new south wales , april 2015 . source : ian shaw / inaturalist . org . license : cc by attribution - noncommercial
there are no known species - specific conservation measures in place for istiblennius edentulus . however , its range does overlap several marine protected areas ( world database of protected areas 2010 ) .
springer , v . g . and j . t . williams , 1994 . the indo - west pacific blenniid fish genus istiblennius reappraised : a revision of istiblennius , blenniella , and paralticus , new genus . . smithson . contrib . zool . 565 : 193 p
springer , v . g . and j . t . williams , 1994 . the indo - west pacific blenniid fish genus istiblennius reappraised : a revision of istiblennius , blenniella , and paralticus , new genus . smithson . contrib . zool . 565 : 1 - 193 . ( ref . 9962 )
springer , v . g . & williams , j . t . 1994 ,
the indo - west pacific blenniid fish genus istiblennius reappraised : a revision of istiblennius , blenniella , and paralticus , new genus
, smithsonian contributions to zoology , vol . 565 , pp . 1 - 193 figs 1 - 73
istiblennius edentulus is found throughout the indo - west pacific , including south africa , east africa , madagascar , mascarenes , and seychelles , north to the red sea and persian gulf , east to western australia , new south wales , lord howe and rapa islands , new caledonia , wake atoll , the marquesan , line , and pitcairn islands , north to the tuamoto and ogasawara islands , and southern japan ( forster and schneider 1801 , myers 1991 ) .
greek , istios = sail + greek , blennios = mucus ( ref . 45335 )
marine ; brackish ; reef - associated ; depth range 0 - 5 m ( ref . 9710 ) , usually 0 - 1 m ( ref . 9710 ) . tropical ; 32°n - 32°s
indo - pacific : red sea and east africa to the line , marquesan , and tuamoto islands , north to southern japan , south to lord howe and rapa .
maturity : l m ? range ? - ? cm max length : 16 . 0 cm tl male / unsexed ; ( ref . 48636 ) ; 13 . 2 cm sl ( female )
dorsal spines ( total ) : 12 - 13 ; dorsal soft rays ( total ) : 19 - 21 ; anal spines : 2 ; anal soft rays : 21 - 23 . males darkly dusky with 5 - 6 pairs of bands on body and pale stripes on dorsal fin ; develop a crest . females paler in color , bands broken into spots posteriorly , dorsal fin spotted ( ref . 4404 ) .
intertidal ( ref . 31184 , 48636 ) . adults are common in areas with large rubble pieces which are often used to built breakwaters or to support jetty - pylons ( ref . 48636 ) . they hide in cracks or holes when not feeding . they jump out of the water in energetic skippings to another pool when pursued ( ref . 2158 , 48636 ) . may remain out of water under rocks or seaweeds ( ref . 31184 ) . they breathe air when out of water ( ref . 31184 ) . feeds on filamentous algae ( ref . 89972 ) . oviparous . eggs are demersal and adhesive ( ref . 205 ) , and are attached to the substrate via a filamentous , adhesive pad or pedestal ( ref . 94114 ) . larvae are planktonic , often found in shallow , coastal waters ( ref . 94114 ) .
) : 24 . 7 - 29 . 3 , mean 28 . 4 ( based on 2954 cells ) .
phylogenetic diversity index ( ref . 82805 ) : pd 50 = 0 . 5001 [ uniqueness , from 0 . 5 = low to 2 . 0 = high ] .
trophic level ( ref . 69278 ) : 2 . 0 ±0 . 00 se ; based on food items .
resilience ( ref . 69278 ) : high , minimum population doubling time less than 15 months ( preliminary k or fecundity . ) .
vulnerability ( ref . 59153 ) : low to moderate vulnerability ( 27 of 100 ) .
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thank you for taking the time to provide feedback on the iucn red list of threatened species website , we are grateful for your input .
eschmeyer , w . n . ( ed . ) . 2014 . catalog of fishes . updated 27 august 2014 . available at : urltoken . ( accessed : 27 august 2014 ) .
williams , j . t . & smith - vaniz , w . f .
this is a widespread species that is locally abundant with no known threats and occurs in marine protected areas in parts of its range . it is therefore listed as least concern .
australia ; bangladesh ; cambodia ; china ; djibouti ; egypt ; eritrea ; fiji ; french polynesia ( marquesas ) ; hong kong ; india ; indonesia ; iran , islamic republic of ; iraq ; japan ; kenya ; kuwait ; madagascar ; malaysia ; mozambique ; myanmar ; new caledonia ; new zealand ; oman ; pakistan ; papua new guinea ; philippines ; saudi arabia ; seychelles ; singapore ; somalia ; south africa ; sudan ; taiwan , province of china ; tanzania , united republic of ; thailand ; united arab emirates ; united states minor outlying islands ( wake is . ) ; viet nam ; yemen
this species is common and locally abundant throughout most of its range ( williams , j . t . pers . comm . 2009 ) .
williams , j . t . & smith - vaniz , w . f . 2014 .
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a pale greenish grey rockskipper with dark greyish divided bars on sides that extend onto the dorsal - fin base , vertical orange wavy lines in the spaces between the bars , and a pale - edged grey bar through the eye across the upper lip . females have orange - brown spots on rear of body and on dorsal and anal fins . video of rippled rockskippers in a tide pool in shirahama , japan .
inhabits the edge of rocky shorelines in relatively protected areas where wave action is not severe .
allen , g . r . , hoese , d . f . , paxton , j . r . , randall , j . e . , russell , b . c . , starck , w . a . , talbot , f . h . & whitley , g . p . 1976 . annotated checklist of the fishes of lord howe island .
allen , g . r . & smith - vaniz , w . f . 1994 . fishes of cocos ( keeling ) islands .
the marine fishes of north - western australia . a field guide for anglers and divers
. perth , wa : western australian museum vi 201 pp . , 70 pls .
castelnau , f . l . de 1875 . researches on the fishes of australia . intercolonial exhibition essays . 2 . pp . 1–52 in ,
forster , j . r . & schneider , j . g . in bloch , m . e . & schneider , j . g . 1801 .
francis , m . 1993 . checklist of the coastal fishes of lord howe , norfolk , and kermadec islands , southwest pacific ocean .
hobbs , j - p . a . , s . j . newman , g . e . a . mitsopoulos , m . j . travers , c . l . skepper , j . j . gilligan , g . r . allen , h . j . choat & a . m . ayling . 2014 . checklist and new records of christmas island fishes : the influence of isolation , biogeography and habitat availability on species abundance and community composition .
hobbs , j - p . a . , s . j . newman , g . e . a . mitsopoulos , m . j . travers , c . l . skepper , j . j . gilligan , g . r . allen , h . j . choat & a . m . ayling . 2014 . fishes of the cocos ( keeling ) islands : new records , community composition and biogeographic significance .
johnson , j . w . 2010 . fishes of the moreton bay marine park and adjacent continental shelf waters , queensland , australia . pp . 299 - 353 in davie , p . j . f . & phillips , j . a . proceedings of the thirteenth international marine biological workshop , the marine fauna and flora of moreton bay .
. sydney , nsw , australia : new holland publishers xvii , 434 pp .
ogilby , j . d . 1899 . additions to the fauna of lord howe island .
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hadorn , r . & fraussen , k 2003 ,
the deep - water indo - pacific radiation of fusinus ( chryseofusus ) subgen . nov . ( gastropoda : fasciolariidae )
, iberus , vol . 21 , no . 1
daniels , g . 1978 ,
a catalogue of the type specimens of diptera in the australian museum
, records of the australian museum , vol . 31 , no . 11 , pp . 411 - 471
mckeown , k . c . 1948 ,
a reference list of types of coleoptera in the australian museum .
, records of the australian museum , vol . 22 , no . 1 , pp . 95 - 139
urn : lsid : biodiversity . org . au : afd . taxon : d707bc53 - 9254 - 4f27 - a573 - 76d44a42e5eb
urn : lsid : biodiversity . org . au : afd . taxon : 94f6eb0c - fac9 - 433c - b609 - 77a9c8f340a1
urn : lsid : biodiversity . org . au : afd . name : 592863
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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 .
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neotype usnm 292529 , tahiti , society islands ( original type locality huanaine island ) .
castelnau , f . l . de 1875 . researches on the fishes of australia . intercolonial exhibition essays . 2 . pp . 1–52
holotype whereabouts unknown ( qm 1 . 862 labelled type , with 2 specimens , both too large to be holotype and holotype presumed lost ) , lord howe island .
. paris : levrault vol . 11 506 pp . pls 307 - 343 .
lectotype mnhp a . 2003 , mauritius . paralectotype ( s ) mnhp 7067 ; mnhp a , 2002 ; mnhp a , 2005 ; mnhp a , 2007 ; mnhp a . 2304 ; mnhp 846 .
dirk hartog island ( 26°08 ' s ) to cartier island ( 12°33 ' s ) , wa and lizard island , qld ( 14°39 ' s ) to sydney , nsw ( 33°48 ' s ) ; tropical , indo - west - central pacific .
johnson , j . w . 2010 . fishes of the moreton bay marine park and adjacent continental shelf waters , queensland , australia . pp . 299 - 353
davie , p . j . f . & phillips , j . a . proceedings of the thirteenth international marine biological workshop , the marine fauna and flora of moreton bay .
mcculloch , a . r . 1929 . a check - list of the fishes recorded from australia . part iii .
allen , g . r . 1985 . fishes of western australia . book 9 . pp . 2207 - 2534 526 pls
intertidal ( ref . 31184 , 48636 ) . adults are common in areas with large rubble pieces which are often used to built breakwaters or to support jetty - pylons ( ref . 48636 ) . they hide in cracks or holes when not feeding . they jump out of the water in energetic skippings to another pool when pursued ( ref . 2158 , 48636 ) . may remain out of water under rocks or seaweeds ( ref . 31184 ) . they breathe air when out of water ( ref . 31184 ) . feeds on filamentous algae ( ref . 89972 ) . oviparous . eggs are demersal and adhesive ( ref . 205 ) , and are attached to the substrate via a filamentous , adhesive pad or pedestal ( ref . 94114 ) . larvae are planktonic , often found in shallow , coastal waters ( ref . 94114 ) .
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oxyurichthys keiensis also known as the kei goby is a species of goby native to marine and brackish waters along the coasts of mozambique , south africa , madagascar and the seychelles .

this species can reach a length of 7 centimetres ( 2.8 in ) tl .

kei goby

a study by harrison ( 2005 ) found the kei goby to be one of the most frequently captured species in open subtropical estuaries in south africa .
threatened kei goby ( oligdepis keimsis ) . more familiar to many people are the two breeding species of turtles : hawksbill ( eretmochelys imbricata ) and green ( chelonia mydas ) . more
the kei goby ( oligolepis keiensis ) is a species of fish in the gobiidae family . it is found in mozambique and south africa . source - * skelton , p . 1996 . oligolepis keiensis . more
i absolutely agree with you kei - those colours are out of this world ! i love how the blues melt into each other thanks to the shallow depth of field and the small goby pops out of the image unexpectedly due to its beautiful colouring around the eyes .
the kei goby ( oligolepis keiensis ) occurs from mozambique to the sundays estuary in the eastern cape ( south africa ) , as well as around the seychelles and madagascar . this species has a severely fragmented distribution ( a . k . whitfield pers . comm . 2008 ) .
tridacnidae has much color pattern , but the blue dot pattern is beautiful in particular . i met the goby which coexisted with a tridacnidae with the beautiful blue dot pattern for the first time last year . i like the small goby which coexists with this tridacnidae . the in them loving design , color and the favorite location exist . i could release the shutter by his favorite point . this galaxy and goby look very mysterious .
there are no known major threats to the kei goby . due to its preference for perennial river flow systems , changes to the flow regime of freshwater systems by activities such as abstraction may pose a threat to this species . it is also likely to be impacted by threats such as water pollution from industrial and domestic activities , dredging , and shipping traffic . however , these are localised threats and not known across the entire distribution of this species .
justification : the kei goby ( oligolepis keiensis ) has been assessed as least concern . this species has a wide range in eastern africa . it is restricted to estuarine habitats and although it is not known to be under any specific threat , it may be affected by habitat degradation through activities in estuaries within its range . given its wide range , lack of specific threats , and no evidence of population decline , this species is assessed as least concern . monitoring of the population numbers and rate of decline is needed so that changes to the threat status of this species can be noted .
the kei goby is a benthic species , most commonly found in estuaries . in the warm - temperate and subtropical regions of south africa , this species appears to be exclusively found in estuarine environments , as there are no records from freshwater or marine systems . its main food items include crustaceans and polychaetes . it is most abundant on sandy mud substrata in the middle and upper reaches of permanently open estuaries ( salinity 5–15 psu ) , but may also be found in some temporarily closed estuaries ( a . k . whitfield pers . comm . 2008 ) . estuaries with a perennial river flow appear to be the favoured type ( a . k . whitfield pers . comm . 2008 ) .
we tested the reactions of free embryos of the amphidromous goby , rhinogobius brunneus , to light under both artificial and ambient conditions along streams in which their downstream migration occurs . the embryos showed a positive phototaxis to 500 1ux light but a negative response to light of more than 5000 lux . they were able to swim at 1 . 54 cm sec −1 t in still water and showed positive rheotaxis , but their swimming ability was not sufficient to allow active movement in rapids . ambient natural light conditions varied among locations in relation to local topographical features . the variation in the diel periodicity of their migration could be explained by the interaction between behavioral reactions of embryos and environmental factors along river courses .
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eschmeyer , w . n . , fricke , r . , and ven der laan , r . ( eds . ) . 2017 . catalog of fishes : genera , species , references . updated 31 july 2017 . available at : urltoken .
de silva , r . , milligan , h . , lutz , m . , batchelor , a . , jopling , b . , kemp , k . , lewis , s . , lintott , p . , sears , j . , wilson , p . , smith , j . & livingston , f .
there are no species - specific conservation measures in place for oligolepis keiensis , however its distribution may coincide with a number of marine protected areas . further research is needed to monitor population trends and the status of its estuarine habitat .
this errata assessment has been created because the map was accidentally left out of the version published previously .
( errata version published in 2017 ) . the iucn red list of threatened species 2010 : e . t15239a115125895 .
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greek , oligos = small + greek , lepis = scale ( ref . 45335 )
western indian ocean : inhaca , mozambique to the fish river mouth , south africa ; including seychelles and madagascar .
maturity : l m ? range ? - ? cm max length : 7 . 0 cm tl male / unsexed ; ( ref . 2798 )
dorsal spines ( total ) : 7 ; dorsal soft rays ( total ) : 112 ; anal spines : 1 ; anal soft rays : 12 . body grey or dark brown with brown oblique streak ; upper caudal base with a black spot ; adults with blackish pelvic fins ( ref . 2798 ) .
hoese , d . f . , 1986 . gobiidae . p . 774 - 807 . in m . m . smith and p . c . heemstra ( eds . ) smiths ' sea fishes . springer - verlag , berlin . ( ref . 2798 )
) : 24 . 3 - 27 . 6 , mean 26 . 9 ( based on 70 cells ) .
phylogenetic diversity index ( ref . 82805 ) : pd 50 = 0 . 5156 [ uniqueness , from 0 . 5 = low to 2 . 0 = high ] .
bayesian length - weight : a = 0 . 00851 ( 0 . 00448 - 0 . 01616 ) , b = 3 . 04 ( 2 . 88 - 3 . 20 ) , in cm total length , based on lwr estimates for this species & ( sub ) family - body ( ref . 93245 ) .
trophic level ( ref . 69278 ) : 3 . 7 ±0 . 5 se ; based on size and trophs of closest relatives
resilience ( ref . 69278 ) : high , minimum population doubling time less than 15 months ( preliminary k or fecundity . ) .
vulnerability ( ref . 59153 ) : low vulnerability ( 15 of 100 ) .
at ibm , when performance review time rolls around , staff get judged not on their past achievements but also on how they might perform . . .
samsung has taken the ‘make in india’ initiative to another level by launching ‘ . . .
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max length : 7 . 0 cm tl male / unsexed ; ( ref . 2798 )
pd 50 = 0 . 5156 many relatives ( e . g . carps ) 0 . 5 - 2 . 0 few relatives ( e . g . lungfishes )
high , minimum population doubling time less than 15 months ( preliminary k or fecundity . )
balon , e . k . & a . goto . 1989 . styles in reproduction and early ontogeny . pp . 1–47 .
a . goto & k . maekawa ( ed . ) reproductive behavior in fishes : styles and strategies , tokai university press , tokyo . ( in japanese . )
blaxter , j . h . s . 1973 . monitoring the vertical movements and light responses of herring and plaice larvae . j . mar . biol . ass . u . k . 53 : 635–647 .
blaxter , j . h . s . & k . f . ehrlich . 1974 . changes in behaviour during starvation of herring and plaice larvae . pp . 575–588 .
j . h . s . blaxter ( ed . ) the early life history of fishes , springer - verlag , hidelberg .
kani , t . 1944 . ecology of torrent - inhabiting insects . pp . 171–317 .
h . furukawa ( ed . ) insects 1 , kenkyu - sha press , tokyo . ( in japanese . )
. chuokoron - sha press , tokyo . 176 pp ( in japanese . )
mcdowall , r . m . 1988 . diadromy in fishes . migration between freshwater and marine environments . croom helm , london . 308 pp .
gill with a proposition on the relationship between land - locking and speciation of some freshwater gobies in japan . men . col . sc . univ . kyoto ser . b 27 : 97–115 .
mizuno , n . & h . kawanabe . 1981 . a topographical classification of streams , with an introduction of the system widely used in japan . 1 . reach type , stream zone and stream type . ver . internat . verein limnol . 21 : 913 .
mizuoka , s . 1967 . studies on fluvial variations in the gobioid fish , ‘yoshinobori’ . iv . distributions and variations in color pattern and the pectoral fin ray numbers . bull . fac . educ . hiroshima univ . 16 : 43–52 . ( in japanese . )
nagoshi , m . 1982 . diel vertical migration of zooplankters and fish larvae in lake biwa . bull . fac . fish . mie univ . 9 : 1–10 .
northcote , t . g . 1984 . mechanisms of fish migration in rivers . pp . 317–355 .
j . d . mccleave , g . p . arnold , j . j . dodson & w . h . neil ( ed . ) mechanisms of migration in fishes , plenum press , new york .
pavlov , d . s . , a . n . pakhorukov , g . n . kuragina , v . k . nezdoliy , n . p . nekrasoba , d . a . brodskiy & a . l . ersler . 1977 . some features of the downstream migrations of juvenile fishes in the volga and kuban rivers . journal of ichtyology 9 : 157–179 .
woodhead , p . m . j . & d . woodhead . 1955 . reactions of herring larvae to light : a mechanisms of vertical migration . nature 176 : 349–350 .
iguchil , k . & mizuno , n . environ biol fish ( 1991 ) 31 : 295 . urltoken
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cfm script by eagbayani , 30 . 11 . 04 , , php script by cmilitante , 05 / 11 / 2010 , last modified by cmilitante , 14 / 03 / 2013


manduca pellenia is a moth of the family sphingidae .

it is known from mexico , belize , guatemala , nicaragua , costa rica , panama , colombia and ecuador .

the wingspan is 107 – 126 mm .

the underside of the abdomen is shaded with brown scales , especially in the male .

there are heavy , discal , black patches found on the forewing upperside with , forming a band .

there is probably one generation per year in costa rica with adults on wing from september to november .

they feed on flower nectar .

the larvae feed on solanum hayesii and cestrum megalophyllum .

manduca pellenia

manduca pellenia ( herrich - schäffer , 1854 ) = pellenia ( herrich - schäffer , 1854 ) .
manduca pellenia , female , upperside . guatemala , izabal dpt . , morales sierra de caral , finca firmeza
manduca pellenia , female , underside . guatemala , izabal dpt . , morales sierra de caral , finca firmeza
transferred to manduca by d ' abrera , [ 1987 ] , sphingidae mundi : 26 .
family : sphingidae , latreille , 1802 subfamily : sphinginae , latreille , [ 1802 ] tribe : sphingini , latreille , 1802 genus : manduca hubner , 1807 . . . . . . . . . . . species : pellenia herrich - schaffer , 1854
i believe the specimen to the right from yasuni , ecuador , september 7 , 2002 - 2 : 06 am , courtesy of steve graser is m . pellenia , but it could also be m . scutata scutata .
antenna almost as stout as in manduca sexta . abdomen underside shaded with brown scales , especially in the male . foretarsus with 1st segment externally with 4 or 5 moderately long spines and numerous small ones above them . no pulvillus . forewing upperside with discal , black , pubescent patches heavy , forming a band that is strongly angled near vein m3 ; the oblique , black apical line and the posterior part of the black postdiscal line both very heavy ; submarginal zigzag line creamy buff rather than white ; the cells between veins m3 and cup more or less russet between the discal band and postdiscal line . cate
( wing span : ( 107 - 126 mm ) ) , flies in
tropical america
, given as the specimen type locality . the moth can be found in
in costa rica there is probably only one generation annually with moths on the wing in from september until november .
females call in the males with a pheromone released from a gland at the tip of the abdomen . adults nectar at flowers .
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please send sightings / images to bill . i will do my best to respond to requests for identification help .
show appreciation for this site by clicking on flashing butterfly to the left . the link will take you to a page with links to many insect sites .
june 1 , 2015 , 1675m , 109mm , courtesy of terry stoddard , id by bill oehlke .
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except where otherwise noted , content on this site is licensed under a creative commons attribution cc by licence .
confused by a class within a class or an order within an order ? please see our brief essay .
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 .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
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 .
this system is free software released under gnu / gpl 3 . 0 - portal version 1 . 0
this work is licensed under a creative commons attribution - noncommercial - sharealike 3 . 0 united states license .
( bopisduval 1875 ) wingspan 135 - 140mm . please choose from drop tab .
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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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