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 .
ahlberg pe , johanson z ( 1997 ) second tristichopterid ( sarcopterygii , osteolepiformes ) from the upper devonian of canowindra , new south wales , australia , and phylogeny of the tristichopteridae . journal of vertebrate paleontology 17 : 653–673 .
jarvik : postcranial anatomy , basal tetrapod interrelationships and patterns of skeletal evolution . transactions of the royal society of edinburgh : earth sciences 87 : 363–421 .
daeschler eb , shubin nh , jenkins fa jr ( 2006 ) a devonian tetrapod - like fish and the evolution of the tetrapod body plan . nature 440 : 757–763 .
lebedev oa ( 1995 ) morphology of a new osteolepidid fish from russia . bulletin du museum national d ' histoire naturelle section c sciences de la terre paleontologie geologie mineralogie 17 : 287–341 .
long , 1985 , from the upper devonian gogo formation , western australia . records of the australian museum supplements 53 : 1–89 .
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 .


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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( bopisduval 1875 ) wingspan 135 - 140mm . please choose from drop tab .
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tetramorium simillimum , is a species of ant in the subfamily myrmicinae .

it is a small pale colored widespread species that can be found in almost all the continents .

tetramorium simillimum

senior synonym of tetramorium pygmaeum : forel , 1916 pdf : 421 ; of tetramorium auropunctata brevispinosa : borgmeier , 1937b pdf : 241 ; of tetramorium simillimum insulare : yarrow , 1967 pdf : 28 ; of tetramorium simillimum denticulatum , tetramorium simillimum opacior , tetramorium parallela : bolton , 1977 pdf : 131 ; of tetramorium pusillum bantouana , tetramorium pusillum exoleta and material of the unavailable name tetramorium breve referred here : bolton , 1980 pdf : 320 .
the above specimen data are provided by antweb . please see tetramorium simillimum for further details
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denticulatum . tetramorium simillimum r . denticulatum forel , 1902c : 235 ( w . ) india . junior synonym of simillimum : bolton , 1977 : 131 .
opacior . tetramorium simillimum var . opacior forel , 1913k : 81 ( w . ) sri lanka . junior synonym of simillimum : bolton , 1977 : 131 .
senior synonym of tetramorium pygmaeum : forel , 1916 pdf : 421 ; of tetramorium brevispinosa : borgmeier , 1937b pdf : 241 ; of tetramorium insulare : yarrow , 1967 pdf : 28 ; of tetramorium denticulatum , tetramorium opacior , tetramorium parallela : bolton , 1977 pdf : 131 ; of tetramorium bantouana , tetramorium exoleta and material of the unavailable name tetramorium breve referred here : bolton , 1980 : 320 .
jennifer hammock split the classifications by plazi from tetramorium simillimum ( smith , 1851 ) to their own page .
exoleta . tetramorium pusillum var . exoleta santschi , 1914d : 366 ( w . ) nigeria . junior synonym of simillimum : bolton , 1980 : 320 .
the small species is t . simillimum , and the larger species is t . bicarinatum .
insulare . tetramorium simillimum var . insulare santschi , 1928c : 69 ( w . ) fiji is . [ unresolved junior secondary homonym of insularis menozzi , above . ] junior synonym of simillimum : yarrow , 1967 : 28 ; bolton , 1977 : 131 .
bantouana . tetramorium pusillum var . bantouana santschi , 1910c : 382 , fig . 10 ( w . q . m . ) congo . junior synonym of simillimum : bolton , 1980 : 320 .
the range for tetramorium simillimum given by brown and taylor ( 1985 ) is probably understated . unlike some other exotic myrmicines in australia , this species does not seem to adversely affect the native ant fauna .
very similar to tetramorium caldarium , as detailed in the identification section of this other species .
tetramorium simillimum is a small reddish species that sometimes has a darker brown to black gaster . it is slow moving , but can recruit strongly and defend food resources . in the field it is impossible to distinguish from tetramorium caldarium . the species tends to nest and forage on the ground in disturbed areas .
parallela . myrmica parallela smith , f . 1859a : 147 ( w . ) indonesia ( aru i . ) . combination in tetramorium : donisthorpe , 1932c : 455 . junior synonym of simillimum : bolton , 1977 : 131 .
pygmaeum . tetramorium pygmaeum emery , 1877b : 371 ( q . ) ethiopia . emery , 1901e : 62 ( m . ) ; emery , 1915g : 17 ( w . ) . junior synonym of simillimum : forel , 1916 : 421 .
weakly sculptured , pale red much smaller than tetramorium guineense . length : 1 . 6 - 2 mm ( collingwood 1979 ) .
mayr , g . 1861 . die europäischen formiciden . nach der analytischen methode bearbeitet . wien : c . gerolds sohn , 80 pp . ( page 61 , combination in tetramorium )
brevispinosa . wasmannia auropunctata subsp . brevispinosa borgmeier , 1928a : 36 , figs . 4 , 5 ( w . ) brazil . [ unresolved junior secondary homonym of brevispinosus stitz , above . ] junior synonym of simillimum : borgmeier , 1937b : 241 .
bolton , b . 1979 . the ant tribe tetramoriini ( hymenoptera : formicidae ) . the genus tetramorium mayr in the malagasy region and in the new world . bull . br . mus . ( nat . hist . ) entomol . 38 : 129 - 181 ( see also )
bolton , b . 1977 ,
the ant tribe tetramoriini ( hymenoptera : formicidae ) . the genus tetramorium mayr in the oriental and indo - australian regions , and in australia
, bulletin of the british museum ( natural history ) entomology , vol . 36 , pp . 67 - 151
bolton , b . , 1979 , the ant tribe tetramoriini ( hymenoptera : formicidae ) . the genus tetramorium mayr in the malagasy region and in the new world . , bulletin of the british museum ( natural history ) entomology 38 , pp . 129 - 181 : 170 - 171 , ( download )
bharti , h . & kumar , r . 2012 . taxonomic studies on genus tetramorium mayr ( hymenoptera , formicidae ) with report of two new species and three new records including a tramp species from india with a revised key . zookeys . 207 : 11 - 35 . doi : 10 . 3897 / zookeys . 207 . 3040
bolton , b . 1977 . the ant tribe tetramoriini ( hymenoptera : formicidae ) . the genus tetramorium mayr in the oriental and indo - australian regions , and in australia . bulletin of the british museum ( natural history ) . entomology . 36 : 67 - 151 . pdf ( page 131 , senior synonym of denticulatum , opacior and parallela )
agavekar , g . , hita garcia , f . , economo , e . p . 2017 . taxonomic overview of the hyperdiverse ant genus tetramorium mayr ( hymenoptera , formicidae ) in india with descriptions and x - ray microtomography of two new species from the andaman islands . peerj 5 : e3800 ( doi 10 . 7717 / peerj . 3800 ) .
bolton , b . 1980 . the ant tribe tetramoriini ( hymenoptera : formicidae ) . the genus tetramorium mayr in the ethiopian zoogeographical region . bull . br . mus . ( nat . hist . ) entomol . 40 : 193 - 384 ( page 320 , senior synonym of bantouana and exoleta , and material of the unavailable name breve referred here . )
a tramp species that has developed a pantropical distribution and is even found in temperature areas , albeit in protected areas such as heated greenhouses .
collingwood ( 1979 ) - this cosmopolitan species occasionally occurs in heated glasshouses in europe and has been recorded from denmark and also on several occasions in england .
wheeler ( 1908 ) - in culebra a few colonies were found nesting under stones and logs on the beach , in coamo springs several colonies were seen under stones in the creek bottom near the baths .
a common species found as far north as st . johns county , florida . nests are usually in soil in open areas , often around buildings or parking lots . pest status : none . first published florida record : wheeler 1932 . ( deyrup , davis & cover , 2000 . )
the following information is derived from barry bolton ' s new general catalogue , a catalogue of the world ' s ants .
smith , f . 1851 : 118 ( w . ) great britain . meinert , 1861 : 331 ( gynandromorph ) ; andré , 1883a : 289 ( q . m . ) ; imai , baroni urbani ,
referred here : bolton , 1980 : 320 . see also : emery , 1909d : 696 .
unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description .
( length3 / 4 - 1 line ) . head and thorax pale ferruginous , the legs and antennae more pallid , the coxae a little coloured , the eyes black ; the abdomen is rufo - fuscous , pale towards the apex ; the head is evenly longitudinally striate ; the thorax above is without the usual transverse suture , but is a little compressed at the sides about the middle , and gradually slightly narrowed from the prothorax towards the nodes of the peduncle ; the metathorax is truncate at the apex , and the spines are short , broad , and acute ; the abdomen is furnished with a few scattered erect hairs .
myrmica simillima : syntype , worker ( s ) ( apparently lost ) , dorset , england , united kingdom of great britain and northern ireland .
the following notes on f . smith type specimens have been provided by barry bolton ( details ) :
holotype worker in oxford university museum of natural history . specimen without locality label , but with a label “ simillima ” in smith’s writing . type - locality “aru” according to original description .
andré , e . 1883a . les fourmis . [ part ] . pp . 281 - 344 in : andré , edm . 1881 - 1886 . species des hyménoptères d ' europe et d ' algérie . tome deuxième . beaune : edmond andré , 919 + 48 pp . ( page 289 , queen , male described )
borgmeier , t . 1937b . formigas novas ou pouco conhecidas da américa do sul e central , principalmente do brasil ( hym . formicidae ) . arch . inst . biol . veg . ( rio j . ) 3 : 217 - 255 ( page 241 , senior synonym of brevispinosa )
collingwood , c . a . 1979 . the formicidae ( hymenoptera ) of fennoscandia and denmark . fauna entomol . scand . 8 : 1 - 174 .
collingwood , c . a . , pohl , h . , guesten , r . , wranik , w . and van harten a . 2004 . the ants ( insecta : hymenoptera : formicidae ) of the socotra archipelago . fauna of arabia 20 : 473 - 495 pdf
deyrup , m . , davis , l . & cover , s . 2000 . exotic ants in florida . transactions of the american entomological society 126 , 293 - 325 .
emery , c . 1909f . beiträge zur monographie der formiciden des paläarktischen faunengebietes . ( hym . ) teil ix . dtsch . entomol . z . 1909 : 695 - 712 ( page 696 , see also )
forel , a . 1916 . fourmis du congo et d ' autres provenances récoltées par mm . hermann kohl , luja , mayné , etc . rev . suisse zool . 24 : 397 - 460 ( page 421 , senior synonym of pygmaeum )
heterick , b . e . 2009 . a guide to the ants of south - western australia . records of the western australian museum , supplement 76 : 1 - 206 . pdf
imai , h . t . ; baroni urbani , c . ; kubota , m . ; sharma , g . p . ; narasimhanna , m . h . ; das , b . c . ; 1984 . karyological survey of indian ants . jpn . j . genet . 59 : 1 - 32 ( page 8 , karyotype described )
meinert , f . 1861 . bidrag til de danske myrers naturhistorie . k . dan . vidensk . selsk . skr . ( 5 ) 5 : 273 - 340 ( page 331 , gynandromorph )
sharaf , m . r . , fisher , b . l . , collingwood , c . a . , aldawood , a . s . 2017 . ant fauna ( hymenoptera : formicidae ) of the socotra archipelago ( yemen ) : zoogeography , distribution and description of a new species . journal of natural history 51 , 317–378 ( doi 10 . 1080 / 00222933 . 2016 . 1271157 ) .
smith , f . 1851 . list of the specimens of british animals in the collection of the british museum . part vi . hymenoptera , aculeata . london : british museum , 134 pp . ( page 118 , worker described )
wheeler , w . m . 1905c . the ants of the bahamas , with a list of the known west indian species . bull . am . mus . nat . hist . 21 : 79 - 135
wheeler , w . m . 1908a . the ants of porto rico and the virgin islands . bull . am . mus . nat . hist . 24 : 117 - 158 .
yarrow , i . h . h . 1967 . on the formicidae of the azores . bol . mus . munic . funchal 21 : 24 - 32 ( page 28 , senior synonym of insulare )
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meinert , 1861 : 331 ( gynandromorph ) ; andré , 1883a : 289 ( q . m . ) ; imai , baroni urbani , et al . 1984 : 8 ( k . ) .
is an uncommon tramp species in costa rica . i have only collected it once in costa rica , on a picnic table at the headquarters of santa rosa national park .
wild , a . l . , 2007 , a catalogue of the ants of paraguay ( hymenoptera : formicidae ) . , zootaxa 1622 , pp . 1 - 55 : 38 , ( download )
collingwood , c . a . , 1979 , the formicidae ( hymenoptera ) of fennoscandia and denmark . , fauna entomologica scandinavica 8 , pp . 1 - 174 : 85 , ( download )
wheeler , w . m . , 1922 , the ants collected by the american museum congo expedition . , bulletin of the american museum of natural history 45 , pp . 39 - 269 : 193 , ( download )
forel , a . , 1893 , formicides de l ' antille st . vincent . récoltées par mons . h . h . smith . , transactions of the entomological society of london 1893 , pp . 333 - 418 : 382 - 383 , ( download )
ward , p . s . , 2005 , a synoptic review of the ants of california ( hymenoptera : formicidae ) . , zootaxa 936 , pp . 1 - 68 : - 1 , ( download )
forel , a . , 1907 , the percy sladen trust expedition to the indian ocean in 1905 , under the leadership of mr . j . stanley gardiner . no . vi . - fourmis des seychelles , amirantes , farquhar et chagos . , transactions of the linnean society of london 12 , pp . 91 - 94 : - 1 , ( download )
santschi , f . , 1914 , formicidae . , voyage de ch . alluaud et r . jeannel en afrique orientale ( 1911 - 1912 ) . résultats scientifiques . hyménoptères 2 , pp . 41 - 148 : 106 , ( download )
47 times found in tropical dry forest , 37 times found in urban / garden , 23 times found in coastal scrub , 16 times found in rainforest , 6 times found in port of entry / city , 23 times found in forest , 18 times found in mixed forest , 15 times found in urban garden , 7 times found in tropical dry forest on tsingy , 10 times found in spiny forest / thicket , . . .
57 times sifted litter ( leaf mold , rotten wood ) , 52 times ground forager ( s ) , 44 times ex rotten log , 23 times under stone , 16 times on low vegetation , 13 times under rootmat , litter on rock , 11 times ex soil , 9 times sifted litter , 11 times under tree bark , live tree , 2 times foraging on ground , 6 times ex rotten stick on ground , . . .
30 times mw 50 sample transect , 5m , 9 times h , 15 times aspirating ; pb bait , 10 times mw 25 sample transect , 5m , 7 times 9 maxiwinks , mixed samples , 11 times winkler , 14 times hand collected , 11 times pitfall trap , 2 times 2 maxi winks , 7 times l , 2 times aspirating , . . .
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, and accurate separation of these two species may require a taxonomic specialist or a side by side comparison with previously determined specimens . the characters presented in the comparison chart are often difficult to observe . if the user cannot confidently differentiate between the two species , it is recommended to determine the specimen as
near
for an example of how similar they can appear in the field . although a careful examination under the microscope is required ,
is believed to be native to africa and is now widely distributed across the pacific and other tropical regions . the species can achieve dense populations in disturbed habitats and is likely to adversely affect native biodiversity .
at peanut butter bait ( suva , fiji ) . notice the small size , pale color , slow movement and relatively short appendages .
smith , f . 1851 : 118 ( w . ) great britain . meinert , 1861 : 331 ( gynandromorph ) ; andré , 1883a : 289 ( q . m . ) ; imai , baroni urbani , et al . 1984 : 8 ( k . ) . combination in
bolton , b . ( 1977 ) the ant tribe tetramoriini ( hymenoptera : formicidae ) . the genus
mayr in the oreintal and indo - australian regions , and in australia . bulletin of the british museum ( natatural history ) entomology , 36 , 67 - 151 .
bolton , b . ( 1995 ) a new general catalogue of the ants of the world . harvard university press , cambridge , massachusetts , 504 pp .
wilson , e . o . & taylor , r . w . ( 1967 ) the ants of polynesia ( hymenoptera : formicidae ) . pacific insects monograph , 14 , 1 - 109 .
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meinert , 1861 : 331 ( gynandromorph ) ; andré , 1883a : 289 ( q . m . ) ; imai , baroni urbani , et al . 1984 : 8 ( k . ) .
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