crypturellus reai is a species of miocene fossil bird in the tinamou family .

it was described in 2012 from material originally excavated by american palaeontologist barnum brown in 1899 in patagonia .

the specific epithet honours amadeo m. rea , an educator , mentor and friend of the describer .

crypturellus reai

–musculus fibularis longus , sesamoid : absent ( 0 ) ; present ( 1 ) . specimens of crypturellus soui , crypturellus cinnamoneus , and crypturellus boucardi show a sesamoid enclosed in the tendon of the m . fibularis longus ( = m . peronaeus longus of hudson et al . , 1972 ) .
reduced strict consensus of the phylogenetic analysis of fossils and living tinamous under implied weight . differences between topologies under equal versus implied weights analyses are marked with an ‘ ’ . alternative positions of the fossils excluded from the strict consensus a posteriori are indicated above branches ( a , macn - sc - h ; b , macn - sc - t ; c , nothura sp . ; d , crypturellus reai ) . support values of the phylogenetic analysis of extant species and analysis including fossils , respectively , are below branches .
–musculus pterygoideus , fasciculus caudalis : absent ( 0 ) ; present ( 1 ) . absent in nothura , tinamus , and crypturellus ( elzanowski , 1987 ) .
musculus supinator , distinctly longer : absent ( 0 ) ; present ( 1 ) . this muscle is much longer in tinamus and crypturellus ( hudson et al . , 1972 ) .
abstract a new species of tinamou from the early - middle miocene ( santacrusian ) , santa cruz formation of argentina is named . the new species is approximately 16 million year old and has an affinity with the modern genus crypturellus based on the unique characteristics of the humerus , hence , the designation aff . crypturellus . fossil species and the zooarchaeological record of modern tinamous are given .
–musculus orbicularis palpebrarum , origin from ossicula supraorbitales : absent ( 0 ) ; present ( 1 ) . crypturellus , tinamus , eudromia , nothocercus , and some species of nothoprocta share the presence of the ossiculum supraorbitalis on the dorsal surface of the area interorbitalis ; however , the origin of the m . orbicularis palpebrarum from these ossicles has been described only for crypturellus ( elzanowski , 1987 ) . this character has been coded as non - comparable for other tinamous ( without ossiculum supraorbitalis ) .
–musculus depressor palpebrae ventralis : well developed ( 0 ) ; vestigial ( 1 ) . in tinamus and crypturellus , the m . depressor palpebrae ventralis is vestigial and fused to the m . tensor periorbitae . by contrast , in other tinamous both muscles are well developed ( elzanowski , 1987 ) .
–musculus protractor pterygoidei et quadrati : undivided ( 0 ) ; bipartite ( 1 ) . in most tinamous ( except for tinamus and crypturellus ) , the m . protractor et quadrati consist of two parts : pars superficialis and pars profunda . no division of this muscle has been described for ratites ( elzanowski , 1987 ) .
–musculus adductor mandibulae externus , pars caudalis : absent ( 0 ) ; present ( 1 ) . the m . adductor mandibulae externus of most birds is divided into four parts : superficialis , medialis , profunda , and caudalis . in most tinamous ( except for tinamus and crypturellus ) , the pars caudalis is absent ( elzanowski , 1987 ) .
–musculus pseudotemporalis , attachment to os suprangulare : absent ( 0 ) ; present ( 1 ) . in tinamous , the insertion of this muscle occurs on the tuberculum pseudotemporale of the mandible , near the processus medialis mandibularis ; only in crypturellus is it also attached to the os supraangulare ( on the dorsal area of the tuberculum ) ( elzanowski , 1987 ) .
–ligamentum jugomandibulare externum , insertion : undivided ( 0 ) ; bipartite ( 1 ) . only in crypturellus does this ligament bifurcate at the end , and thus the insertion at the supraangular area of the mandible is clearly bipartite ( elzanowski , 1987 ) . the ligamentum jugomandibulare externum is present in all birds except for galliformes and anhimidae ( dzerzhinskii , 1983 ) , therefore these taxa were coded as non - comparable for this character .
–coracoid , proximal margin of cotyla scapularis with pneumatic openings ( fig . 9 ) : absent or few small foramina ( 0 ) ; perforated with large foramina ( 1 ) . after bertelli and chiappe ( 2005 , character 24 ) . pneumatization of this area is present in the living crypturellus , eudromia , tinamotis , and the extinct taxa eudromia olsoni , nothura sp . , macn - sc - 10 , and macn - sc - 13 .
based on the present analysis , tinamidae is subdivided into two groups : the forest - dwelling tinaminae ( crypturellus , tinamu s , nothocercus ) and open - area nothurinae ( taoniscus , nothura , nothoprocta , eudromia , tinamotis , and rhynchotus ; fig . 1 and 2 ) . with the exception of nothura , almost all currently recognized polytypic genus - level taxa of tinamous were recovered with high support ( gc > 91 ; fig . 1 ) .
–musculus quadratomandibularis , insertion : beyond the dorsal margin of the mandible ( 0 ) ; does not extend so far dorsally ( 1 ) . the m . quadratomandibularis attaches on the medial surface of the mandible of tinamous ( fossa caudalis mandibulae ) , and extends beyond the dorsal margin of the mandible in rhynchotus , eudromia , tinamotis , and nothoprocta ( state 0 ) ; the alternative condition ( state 1 ) is present in tinamus , nothura , and crypturellus ( elzanowski , 1987 ) .
+ musculus adductor mandibulae externus , pars profunda , and superficialis : fused ( 0 ) ; partially separated ( 1 ) ; separated ( 2 ) . in rhynchotus and nothoprocta , the m . adductor mandibulae externus is partially tripartite ( with pars superficialis and pars profunda partially fused state 1 ) ; alternatively , it is bipartite ( pars superficialis and profunda fused into pars rostralis , state 0 ) in eudromia , tinamotis , and nothura , and tripartite ( condition 2 ) in crypturellus and tinamus ( elzanowski , 1987 ) .
a–d , tarsometatarsus , plantar view : tinamotis pentlandii ( a ) ; apteryx australis ( b ) ; nothocercus bonapartei ( c ) ; and crypturellus erythopus ( d ) . e–h , tarsometatarsus , proximal view : megapodius freycinet ( e ) ; tinamotis pentlandii ( f ) ; crypturellus soui ( g ) ; and eudromia elegans ( h ) . i–l , tarsometatarsus , distal end ( dorsal view ) : tinamotis pentlandii ( i ) ; rhynchotus rufescens ( j ) ; eudromia elegans ( k ) ; and megapodius freycinet ( l ) . abbreviations : ci , crista intermedia hypotarsi ; cl , crista lateralis hypotarsi ; cm , crista medialis hypotarsi ; fp , foramen vasculare proximale ; fd , foramen vasculare distale ; fm , fossa metatarsi i ; fv , foramen vasculare distale ; il , incisura intertrochlearis lateralis ; ii , trochlea metatarsi ii ; iii , trochlea metatarsi iii ; iv , trochlea metatarsi iv ; la , cotyla lateralis ; me , cotyla medialis . the numbers denote characters and character states as listed in appendix 2 . figures are not drawn to scale .
a–h , sternum , ventral view , proximal end : eudromia elegans ( a ) ; nothocercus julius ( b ) ; nothura maculosa ( c ) ; nothoprocta cinerascens ( d ) ; crypturellus erythropus ( e ) ; taoniscus nanus ( f ) ; tinamotis pentlandii ( g ) ; and tinamus major ( h ) . i , j , clavicula of nothoprocta pentlandii ( caudal view ; i ) and crypturellus brevirostris ( cranial view ; j ) . k , l , scapula ( medial view ) of nothoprocta pentlandii ( k ) and tinamus major ( l ) . abbreviations : a , acromion ; c , carina sterni ; cl , crista lateralis carinae ; es , extremitas sternalis claviculae ; fa , facies articularis acrocoracoidea ; fh , facies articularis humeralis ; fn , foramen pneumaticum ; ic , incisura intercostalis ; il , incisura lateralis ; pa , processus acromialis ; pc , processus craniolateralis ; sa , sulcus articularis coracoideus ; sc , sulcus carinae ; si , spina interna ; tl , trabecula lateralis . the numbers denote characters and character states as listed in appendix 2 . figures are not drawn to scale .
–sternum , rostrum , relative width of spina interna rostri : slender , rostral end narrower than craniolateral process ( 0 ) ; wide , broader than craniolateral process ( 1 ) . the width of the internal spine was measured at the cranial portion of this process . within tinamous , the spina interna is narrow only in the genus crypturellus and a few other species ( e . g . nothoprocta cinerascens ) . with the exception of galliformes , this character was scored as not comparable for the out - group because these taxa lack an internal spine ( character 59 ) .
+ musculus ulnimetacarpalis dorsalis , origin : only dorsal head present ( 0 ) ; bipartite , small ventral head ( 1 ) ; bipartite , large ventral and dorsal head ( 2 ) . the origin of this muscle on the distal end of the ulna of tinamous is divided into a large ventral and dorsal head in tinamus and crypturellus , the ventral head is poorly developed in other tinamous ( and completly absent in tinamotis ) . the m . ulnimetacarpalis dorsalis is vestigial in apteryx ( mcgowan , 1982 ) , and was coded as non - comparable for this character .
–orbita , margo supraorbitalis , glandular depressions ( fossae glandularum nasales , fig . 3 ) : absent ( 0 ) ; present ( 1 ) . the portion of the frontal bordering the orbit forms a sharp supraorbital margin in most tinamous ; only in tinamus and crypturellus glandular depressions ( i . e . fossae glandularum nasales ) are developed on the dorsal supraorbital border ; these depressions are shallow and narrow , bounded laterally by the ossicula supraorbitales , and separated by a wide space ( they do not reach the midline as in other birds , such as charadriiformes , procellariiformes , and others ) .
–orbita , interorbital area ( os frontale ) , dorsal surface ( fig . 3 ) : broader than or of similar width as the internarial area ( 0 ) ; narrower ( 1 ) . this character describes the variation observed in the width of the interorbital area of tinamous . the interorbital surface ( excluding the fossae glandularum nasales ) is wider than or subequal to the internarial area ( at the caudal end of the external nares ) in some tinamous ( e . g . eudromia , taoniscus , and rhynchotus ) , whereas it is distinctly narrower in nothura , crypturellus , and nothoprocta , among others .
+ ligamentum quadratomandibulare rostrale : extensive , not divided ( 0 ) ; bipartite , with medial and lateral parts ( 1 ) ; only medial part present ( 2 ) . both medial and lateral parts of this ligament are developed in crypturellus . only the medial part of this ligament is present in tinamus , and the lateral part is present in nothura and eudromia ; an extensive ligament that occupies the position of both parts ( lateral and medial ) occurs in rhynchotus ( elzanowski , 1987 ) . because of the absence of this structure , the condition for nothoprocta and tinamotis was coded as non - comparable .
–musculus latissimus dorsi , pars metapatagialis , relative position to the m . serratus superficialis : over dorsal surface ( 0 ) ; adjacent ( 1 ) . the pars metapatagialis of m . latissimus is very rare among birds ( baumel , 1993 ) , and , if present , inserts into the humeral feather tract with pars metapatagialis of m . serratus superficialis . this uncommon muscle is present in tinamous , and lies adjacent to the m . serratus superficialis in crypturellus , tinamus , and eudromia , whereas it passes over the dorsal surface of this muscle in other tinamous ( hudson et al . , 1972 ) .
a–f , pelvis , lateral view : crypturellus cinnamomeus ( a ) ; megapodius freycinet ( b ) ; tinamus solitarius ( c ) ; nothocercus bonapartei ( d ) ; nothoprocta perdicaria ( e ) ; and nothoprocta taczanowskii ( f ) . g–i , pelvis , dorsal view : tinamus solitarius ( g ) ; tinamotis pentlandi ( h ) ; and nothura darwinii ( i ) . j–l , pelvis , ventral view : nothura darwinii ( j ) ; crypturellus cinnamomeus ( k ) ; and tinamus solitarius ( l ) . abbreviations : ac , foramen acetabuli ; at , antitrochanter ; c , crista for attachment of ilioischiatic membrane ; cd , crista dorsolateralis ilii ; cs , crista spinosa synsacri ; cv , corpus vertebrae ; f , fenestra ischiopubica ; fc , fovea costalis ; fe , foramen ilioischiadicum ; fi , foramina intertransversaria ; fn , fenestra intertransversaria ; is , ischium ; pc , processus costalis ; pr , ala praeacetabularis ilii ; po , ala postacetabularis ilii ; pt , processus transversus ; pu , pubis ; si , sutura iliosynsacralis ; sy , synsacrum ; tb , tuberculum praeacetabulare ; va , vertebra acetabularis . the numbers denote characters and character states as listed in appendix 2 . figures are not drawn to scale .
–mandibula , processus medialis mandibularis , facies articularis parasphenoidalis ( fig . 6 ) : vestigial or not developed ( 0 ) ; distinct ( 1 ) . a medial facet , near the tip of the processus medialis mandibularis , is well developed in most forest tinamous ( e . g . tinamus , nothocercus , and some crypturellus ) and tinamotis , usually connected to the cotylae caudalis and lateralis , forming a common articular surface . the shape and position of these articular surfaces correlates with a process present in the lateral area of the lamina parasphenoidalis that articulates with the facet of the mandible ( baumel and witmer , 1993 ) .
–musculi iliotrochanterici cranialis and medius : separated ( 0 ) ; fused ( 1 ) . the m . iliotrochanterici , related to protraction movements of the femur , originate on the praeacetabular ilium and insert on the trochanter femoris ( baumel , 1993 ) . the m . iliotrochantericus cranialis ( = iliotrochantericus iliacus or anterior of hudson et al . , 1972 ) and medius are present in megapodiidae and most other birds ( george and berger , 1966 ) . these muscles are also separated in most tinamous , except for tinamus and crypturellus , in which the m . iliotrochantericus cranialis is fused throughout the m . iliotrochantericus medialis ( hudson et al . , 1972 ) .
–musculus pterygoideus , pars medialis : separated into three unnipennate portions ( 0 ) ; complex multipennate system ( 1 ) . this muscle ( including the pseudotemporalis profundus and depressor mandibulae ) participates in the movement ( closing ) of the mandible ( baumel , 1993 ) . in tinamous , it is subdivided into pars medialis and lateralis ( elzanowski , 1987 : fig . 53 ) . the morphology of the pars medialis shows two different patterns in tinamous : tinamus , crypturellus , and nothura have three well - separated unnipenate portions , whereas in rhynchotus , nothoprocta , tinamotis , and eudromia these portions are interconnected , forming a multipennate pars medialis ( elzanowski , 1987 ) .
next , tinamus and crypturellus are grouped together with high support ( gc 8 ; fig . 1 ) , and share the presence of glandular depressions ( character 11 , fig . 3a ) and a paired , complete row of ossicula supraorbitales ( character 12 , fig . 3a ) on the skull , a sternum with poorly projected processus craniolateralis ( character 58 , fig . 8h ) , an expanded scapular blade ( character 68 , fig . 8l ) , a coracoid with an overhanging tuberculum brachiale ( character 70 , fig . 9q ) , and a groove for the origin of ligamentum acrocoracohumerale contacting the facies articularis clavicularis ( character 71 , fig . 9p ) .
–coracoid , shape of the facies articularis clavicularis in medial view ( fig . 9 ) : circular to ovate ( 0 ) ; crescent shaped , with overhanging tuberculum brachiale protruding ventromedially ( 1 ) . after bertelli and chiappe ( 2005 : character 19 ) . this character occurs in tinamus , crypturellus , and eudromia . ratites were scored as non - comparable ( and following characters 70–74 ) because both scapula and coracoid are fused , obscuring the morphology of the proximal end of the coracoid . eudromia olsoni and the early miocene coracoids show a crescent - shaped facies articularis ( 1 ) , whereas it is ovate in the extinct nothura parvula and nothura sp . ( 0 ) .
–os lacrimale , lacrimal duct ( fig . 4 ) : perforating the lacrimal as a foramen or almost complete foramen ( 0 ) ; forming a wide notch ( incisura ductus lacrimalis ) ( 1 ) ; not perforating the lacrimal ( 2 ) . because its delicate nature makes it prone to damage , this character could not be confirmed in some taxa ( e . g . crypturellus and nothocercus ) with partially broken lacrimals . the incisura ductus lacrimalis ( 1 ) was coded as present if the lacrimal notch was distinctly wide dorsoventrally , forming a slender pediculus lacrimalis ( elzanowski , 1987 ) . this character was scored as non - applicable in galliformes , in which the ventral projection of the lacrimal in front of the orbit is absent .
+ aponeurosis parabasalis , attachment : to postmeatic area ( 0 ) ; to both the postmeatic area and lamina basitemporalis ( 1 ) ; to lamina basitemporalis ( 2 ) . according to webb ( 1957 ) the aponeurosis parabasalis is present in rhea among the out - group taxa , with attachment on the postmeatic area of the ala tympanica caudalis , on the occipital surface of the skull . among tinamous , a postmeatic attachment was also described for crypturellus and tinamus ( elzanowski , 1987 ) . unlike these taxa , the attachment is exclusively ( or at least mainly ) to the basal plate ( lamina basitemporalis ) in eudromia , tinamotis , rhynchotus , and nothoprocta ; an intermediate state is present in nothura that combines both conditions equally ( elzanowski , 1987 ) .
–musculus pseudotemporalis , ventral temporal portion : absent ( 0 ) ; present ( 1 ) . the m . pseudotemporalis originates from the temporal fossa and inserts to the medial surface of the mandible ( george and berger , 1966 ) . the subdivision of this muscle into three portions ( two pars orbitales and one pars temporalis ) is a generalized character in tinamous ( and also in most ratites ) , in which the temporal fossa is almost entirely occupied by the m . pseudotemporalis ( elzanowski , 1987 ) . in tinamus , crypturellus , and nothura , the origin extends ventrally onto the pleuroesphenoid area ( ventral head ) ( elzanowski , 1987 ) . the presence of a ventral head of pars temporalis was also described for several neognathous birds ( elzanowski , 1987 ) .
–humerus , incisura capitis obstructed by a tubercle projecting from the border of humeral head ( ericson , 1997 : character 54 ; fig . 9 ) : absent ( 0 ) ; present ( 1 ) . livezey ( 1997 : character 203 ) described the presence of a similar tubercle as the insertion of m . scapulohumeralis cranialis in some gruiforms . the presence in the forest tinamou crypturellus of an incisura capitis enclosed by a distal projection of caput humeri was described by bertelli and chiappe ( 2005 : character 30 ) . this condition is also present in galliformes . because of its highly apomorphic morphology ( character 77 ) , the humerus of ratites cannot be coded for this character ( and the following characters 79–81 ) . eudromia olsoni and nothura parvula lack this tubercle on the caudal surface of the humerus .
–orbita , sulcus , and foramen n . olfactorii forming a wide transversal fenestra , which perforates the septum interorbitale ( fig . 4 ) : yes ( 0 ) no ( 1 ) . next to the dorsal margin of the septum interorbitale , the sulcus for the olfactory nerve and the ethmoid artery ( i . e . sulcus n . olfactorii ; baumel and witmer , 1993 ) is developed as a shallow and thin groove in the open - area tinamous ; the foramen n . olfactorii is relatively smaller in eudromia and tinamotis than in other open - area groups . by contrast , both foramen and sulcus are greatly excavated in the forest taxa nothocercus , tinamus , and crypturellus . we consider this character to be non - comparable in apteryx , where this region of the skull is obscured by the highly modified orbital area .
skulls in ventral view : tinamotis pentlandii ( a ) ; crypturellus cinnamomeus ( b ) ; nothura minor ( c ) ; eudromia formosa ( d ) ; nothoprocta perdicaria ( e ) ; and rhynchotus maculicollis ( f ) . abbreviations : cm , condylus medialis ; fch , fossa choanalis ; fnh , foramen n . hypoglossi ; fnv , foramen n . vagi ; foe , foramen v . occipitalis externa ; fof , foramen n . ophthalmici , jm , jugamentum maxillopalatinum ; lch , pars choanalis ( os palatinum ) ; lp , lamina parasphenoidale ; occ , ostium canalis carotici ; pb , processus basipterygoideus ; pc , processus caudomedialis ; pmp , processsus maxillaris ( os palatinum ) ; ppm , processus maxillopalatinus ( os maxillare ) ; pp , processus paroccipitalis ; rm , rostrum maxillare ; rp , rostrum parasphenoidalis ; ta , tuba auditiva . the numbers denote characters and character states as listed in appendix 2 . figures are not drawn to scale .
–musculus latissimus dorsi , pars caudalis , origin extensive : absent ( 0 ) ; present , attached to two ribs ( 1 ) ; very extensive , attached to four ribs ( 2 ) . the large size of the pars caudalis of m . latissimus dorsi ( = m . latissimus dorsi posterior of hudson et al . , 1972 ) , with an extensive origin , is typical of tinamous and a few other groups ( e . g . loons , auks , penguins ; hudson et al . , 1972 ) . tinamous show two different conditions : in tinamus and crypturellus , the origin is very extensive , between the musculus serratus superficialis , pars metapatagialis and pars caudalis , including the second to fifth ribs , and the cranial area of the ilium . the alternative condition and the most generalized is a less extensive surface of origin , including only the fourth and fifth ribs and the cranial area of the ilium . this character was considered ordered .
a–e , mandible , caudal end ( dorsal view ) : tinamus major ( a ) ; crypturellus cinnamomeus ( b ) ; eudromia formosa ( c ) ; rhynchotus maculicollis ( d ) ; and nothoprocta ornata ( e ) . f–k , quadrate , medial view : nothocercus nigrocapillus ( f ) ; tinamus major ( g ) ; nothura darwinii ( h ) ; eudromia elgans ( i ) ; nothoprocta taczanowskii ( j ) ; and nothoprocta perdicaria ( k ) . abbreviations : cc , condylus caudalis ; ccl , cotylae caudalis et lateralis ; cm , condylus medialis ; cp , condylus pterygoideus ; cr , crest from prominentia submeatica ( elzanowski , 1987 ) ; fap facies articularis parasphenoidalis ; fn , foramen pneumaticum ; ot , processus oticus ; pm , processus medialis mandibularis ; po , processus orbitalis ; pr , processus retroarticularis ; ps , prominentia submeatica ( elzanowski , 1987 ) . the numbers denote characters and character states as listed in appendix 2 . figures are not drawn to scale .
ter - qua 2011 proceedings : robert m . chandler . 2012 . a new species of tinamou ( aves : tinamiformes , tinamidae ) from the early - middle miocene of argentina . – palarch’s journal of vertebrate palaeontology 9 ( 2 ) ( 2012 ) , 1 - 8 . issn 1567 - 2158 . 8 pages + 2 figures , 1 table . | palarch
ter - qua 2011 proceedings : robert m . chandler . 2012 . a new species of tinamou ( aves : tinamiformes , tinamidae ) from the early - middle miocene of argentina . – palarch’s journal of vertebrate palaeontology 9 ( 2 ) ( 2012 ) , 1 - 8 . issn 1567 - 2158 . 8 pages + 2 figures , 1 table .
new material of borealosuchus from the bridger formation , with notes on the paleoecology of wyoming’s eocene crocodylians . –
9 ( 5 ) ( 2012 ) , 1 - 7 . issn 1567 - 2158 . 7 pages + 3 figures , 1 table .
ter - qua 2011 proceedings : steven e . fields , h . gregory mcdonald , james l . knight & albert e . sanders . 2012 .
9 ( 3 ) ( 2012 ) , 1 - 19 . issn 1567 - 2158 . 19 pages + 7 figures , 1 table .
to see submission guidelines for palarch ' s journal for vertebrate palaeontology , click here . for archaeology of egypt / egyptology click here . for archaeology of northwest europe , click here . the procedure for submission is outlined at the bottom of the page .
the palarch foundation offers possibilities for fast , scientific and accessible publication . the journals are so - called web based publications and publish scientific papers in various disciplines .
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f . barroso - barcenilla , m . berrocal - casero , h . a . blain , p . m . callapez , o . cambra - moo , f . escaso , c . martín - closas , f . ortega , a . pérez - garcía , i . prieto , j . rodríguez - lázaro , a . ruiz - galván , j . l . sanz , m . segura & p . sevilla ( 2017 )
geological and palaeontological context of three new barremian ( lower cretaceous ) vertebrate sites in the iberian peninsula ( cuenca province , central spain ) .
structure of the nasal region of non - mammalian cynodonts and mammaliaforms : speculations on the evolution of mammalian endothermy .
jorge i . noriega , emilio a . jordan , raúl i . vezzosi & juan i . areta ( 2017 )
a new species of opisthodactylus ameghino , 1891 ( aves , rheidae ) , from the late miocene of northwestern argentina , with implications for the paleobiogeography and phylogeny of rheas .
jorge i . noriega , emilio a . jordan , raúl i . vezzosi & juan i . areta
sociologist ( phd ) , lecturer and researcher ( sociology of science ) , paleontologist ( researcher in micropaleontology ) . majors in sociology and biology , minor in geology . main interests in paleontology : microfossils , molecular fossils , paleobiology and paleoecology . ( read more about me )
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sara bertelli , luis m . chiappe , gerald mayr ; phylogenetic interrelationships of living and extinct tinamidae , volant palaeognathous birds from the new world , zoological journal of the linnean society , volume 172 , issue 1 , 1 september 2014 , pages 145–184 , urltoken
tinamous ( tinamidae ) include 47 species of central and south american birds inhabiting forested as well as open environments . although volant , the flight capabilities of these neotropical birds are limited ( cabot , 1992 ) . numerous studies have recognized the monophyly of tinamous and their relationship to the flightless ratites ( ostriches , emus , and their relatives ) , placing both groups within palaeognaths , an early diverging group of modern birds ( cracraft , 1974 ; lee , feinstein & cracraft , 1997 ; livezey & zusi , 2007 ; hackett et al . , 2008 ; harshman et al . , 2008 ; bourdon , ricqlés & cubo , 2009 ; haddrath & baker , 2012 ; worthy & scofield , 2012 ; smith , braun & kimball , 2013 ) .
here we review the interrelationships of tinamidae based on osteological and myological characters , including known fossil representatives of this clade . the distribution of the fossil tinamous within the resultant cladograms is discussed in light of their significance for understanding the evolution of the two main ecological subdivisions of these birds : the forest - dwelling taxa , traditionally classified in the taxon tinaminae , and the open - area tinamous ( nothurinae ) .
specimens of the taxa listed have been observed directly ( except for nothura sp . , taken from the respective bibliographic references ) .
no available information of this specimen ( missing from the museo de la plata palaeontological collection ) .
osteological characters supporting tinamou relationships . a , pelvis , relative length of cranial and caudal portions of ilium ( character 92 ) , extension of caudal end ( character 97 ) , and development of the tuberculum praeacetabulare ( character 99 ) ; b , tarsometatarsus , opening of hypotarsal sulcus / canal for m . flexor digitorum longus ( character 111 ) ; c , coracoid , development of processus lateralis ( character 76 ) ; d , quadratum , projection of prominentia submeatica ( character 47 ) ; e , shape of cotylae mandibulares ( characters 42–44 ) . the numbers denote characters and character states as described in in appendix 2 . figures not drawn to scale .
skulls in lateral view : tinamotis pentlandii ( a ) ; cypturellus soui ( b ) ; nothoprocta ornata ( c ) ; tinamus major ( d ) ; nothura maculosa ( e ) ; and rhynchotus maculicollis ( f ) . abbreviations : e , os ectethmoidale ; en , external nares ; fi , fonticuli interorbitales ; fdl , foramen ductus lacrimalis ; fno , foramen n . olfactorii ; fo , fonticuli orbitocraniales ; fol , foramen orbitonasale laterale ; ft , fossa temporalis ; ila , incisura ductus lacrimalis ; j , os jugale ; la , os lacrimale ; me , os mesethmoidale ; po , processus postorbitalis ; pp , processus paroccipitalis ; pr , processus orbitalis ; ps , prominentia suprameatica ; pz , processus zygomaticus ; q , os quadratum ; qj , os quadratojugale ; rm , rostrum maxillare ; si , septum interorbitale ; so , sulcus n . olfactorii ; sf , sutura frontoparietalis . the numbers denote characters and character states as listed in appendix 2 . figures are not drawn to scale .
a , b , e , f , axis , left lateral side : rhynchotus maculicollis ( a ) ; eudromia elegans ( b ) ; nothoprocta pentlandii ( e ) ; and tinamotis pentlandii ( articulated with atlas , f ) . c , d , g , h , cervical vertebra 3 , left lateral view : rhynchotus maculicollis ( c ) ; tinamotis pentlandii ( d ) ; eudromia elegans ( g ) ; nothoprocta pentlandii ( h ) . i–k , notarium ( lateral view ) : nothocercus bonapartei ( i ) ; tinamus solitarius ( j ) ; and nothoprocta taczanowskii ( k ) . abbreviations : a , atlas ; ca , corpus axis ; cn , corpus notarii ; cr , crista spinosa notarii ; cv , corpus vertebrae ; d , dens ; ec , eminentia costolateralis ; fa , facies articularis costalis fc , facies articularis caudalis ; fe , fenestra intercristalis ; fi , fenestra intertransversaria ; fn , foramen pneumaticum ; ic , incisura caudalis ; lt , lamina transversa notarii ; p , processus costalis ; pc , processus articularis caudalis ; ps , processus spinosus ; pv , processus ventralis ; zca , zygapophysis caudalis ; zcr , zygapophysis cranialis . the numbers denote characters and character states as listed in appendix 2 . figures are not drawn to scale .
a , femur , craniomedial view , of nothoprocta taczanowskii . b–d , femur , proximal end ( cranial view ) : megapodius freycinet ( b ) ; nothoprocta ornata ( c ) ; and tinamus tao ( d ) . e , f , femur , distal end ( medial view ) of rhynchotus maculicollis ( e ) and tinamotis pentlandii ( f ) . g–l , tibiotarsus , distal end ( cranial view ) : nothoprocta perdicaria ( g ) ; rhynchotus maculicollis ( h ) ; megapodius freycinet ( i ) ; tinamotis penlandii ( j ) ; tinamus tao ( k ) ; and eudromia elegans ( l ) . m , n , tibiotarsus , proximal end ( cranial view ) of tinamus tao ( m ) and tinamotis pentlandii ( n ) . abbreviations : cf , caput femoris ; cl , condylus lateralis ; cm , condylus medialis ; cnc , crista cnemialis cranialis ; cnl , crista cnemialis lateralis ; ct , crista trochanteris ; ft , fossa trochanteris ; ps , pons supratendineus ; se , sulcus extensorius ; si , incisura intercondylaris ; su , sulcus intercondylaris ; tf , trochanter femoris . the numbers denote characters and character states as listed in appendix 2 . figures are not drawn to scale .
the tinaminae is diagnosed by seven osteological apomorphies : an incomplete row of supraorbital ossicles ( character 12 , fig . 3c ) , a wide sulcus and foramen n . olfactorii ( character 15 , fig . 4b , d ) , a mediolaterally wide processus maxillares and maxillopalatini ( character 32 , fig . 5b ) , a mandibula with converging grooves of the ventral surface ( character 36 ) , a distinct facies articularis parasphenoidalis ( character 43 , fig . 6b ) , a rounded processus supracondylaris dorsalis of the humerus ( character 85 , fig . 9g , h ) , a tibiotarsus with a long condylus lateralis ( character 104 , fig . 11h , i , k , l ) , and a nearly enclosed canal for m . flexor digitorum longus ( character 111 , fig . 12g ) . the osteological evidence supporting the clade also brings a considerable measure of conflict ( gc 24 ; fig . 1 ) . when tinaminae is not recovered as a monophyletic group , the resultant topologies are moderately suboptimal , implying four extra steps when it is depicted as non - monophyletic .
within the forest - dwelling tinamous , nothocercus ( gc 99 ; fig . 1 ) is the earliest diverging taxon of tinaminae , and the relationships within this clade are unresolved . the monophyly of this genus was supported by the following osteological synapomorphies : a mandible with a weakly developed processus parasphenoidalis medialis ( character 24 ) , and a medial crest marked on lamina parasphenoidalis ( character 25 ) , a poorly developed processus lateralis of coracoid ( character 76 , fig . 9s ) , a processus flaxorius of the humerus that projects beyond the ventral condyle ( character 87 , fig . 9g ) , the cranial end of the ilium being rounded and markedly expanded laterally ( character 94 , fig . 10i , j ) , and a deeply excavated fossa parahypotarsalis lateralis of tarsometatarsus ( character 113 , fig . 12c ) .
a monophyletic tinamus ( gc 96 ; fig . 1 ) is supported by the following ostcranial characters : synsacrum with a flat centrum and wide processus costales ( character 54 ) , spina interna rostri markedly elongate ( character 62 , fig . 8h ) , ulna distinctly longer than humerus ( character 88 ) , preacetabular and postacetabular portions of pelvis of subequal length ( character 92 ; fig . 10c , g ) , markedly projected crista trochanteris of femur ( character 100 , fig . 11d ) , tarsometatarsus about the same length as the femur ( character 101 ) , and condylus lateralis with angular proximal margin , widening distally ( 108 , fig . 11e ) . tinamus major + tinamus guttatus ( pelzeln , 1863 ) form a relatively well supported clade ( gc 59 ; fig 1 ) , characterized by a notarium with three fused vertebrae ( character 53 , fig . 7i ) and a deep fossa parahypotarsalis lateralis of the tarsometatarsus ( character 113 ; fig . 12c ) .
the clade consisting of taoniscus , nothura , nothoprocta , and rhynchotus is recovered with high support ( gc 95 ; fig . 1 ) . a vestigial temporal notch ( character 5 , reversed in rhynchotus ; fig . 4c , e ) , large fonticuli orbitocraniales ( character 14 , fig . 4c , e ) , a long and curved processus orbitalis of the quadratum ( character 48 , fig . 6h , j , k ) , and an elongated cotyla medialis of the mandibula ( character 44 , fig . 6d , e ) are characteristic features of the skull of the members of this clade . the postcranium also displays several diagnostic characters : the synsacrum has a distinctly depressed dorsal surface ( character 56 , fig . 10i ) and a flat centrum , with incompletely fused processus costales ( character 48 , fig . 10j ) , the sternum possesses greatly elongated processus craniolaterales ( character 58 , fig . 8c , f ) , the carpometacarpus has rounded caudal rims with deep infratrochlear pits ( character 91 , fig . 9j ) , the pelvis is characterized by a praeacetabular area that has twice or more the length of the postacetabular portion ( character 92 , fig . 10e , f ) , an illium with expanded cranial ( character 94 , fig . 10i , j ) and caudal ( character 97 , fig . 10i , j ) ends , and a well - projected tuberculum praeacetabulare ( character 99 ; fig . 10e , f ) .
the monophyly of rhynchotus is highly supported ( gc 99 , fig . 1 ) by the following synapomorphies : a marked crista nuchalis sagittalis ( character 2 , fig . 3h ) , prominent processus paraoccipitalis ( character 3 , fig . 3h ) , a large fossa temporalis ( character 5 , fig . 4f ) , notched ventral margin of the processus zygomaticus ( character 8 , fig . 4f ) , a broad interorbital area of os frontale ( character 13 , fig . 3c ) , a narrow lacrimal–ectethmoid complex ( character 19 , fig . 4f ) , a rostrocaudally wide pars choanalis of os palatinum ( character 29 , fig . 5f ) , absence of lateral grooves of maxilla and mandibula ( character 35 ) , long and strongly curved cotyla lateralis and caudalis of mandibula ( character 42 , fig . 6d ) , tibiotarsus , condylus lateralis distinctly longer than medialis ( character 104 ; fig . 11h , l ) .
with the exception of the early miocene specimens , most tertiary fossils fall within the open - area nothurinae in all the most parsimonious cladograms . the late miocene eudromia sp . is positioned outside the clade including eudromia and tinamotis ( fig . 1 ) . most of the coded character states of the coracoid are unknown in the coracoid of eudromia sp . and therefore are currently optimized as ambiguous apomorphies of this node ; however , the coracoid of the species of the ( eudromia + tinamotis ) clade lacks a cranially projected processus acrocoracoideus , an unambiguous apomorphy absent in eudromia sp . , in which the process is cranially projected .
the phylogenetic placement of the pliocene tinamou species eudromia olsoni and nothura parvula is consistent with their original assignment ( fig . 1 ) . the placement of eudromia olsoni within the extant eudromia clade is supported by the presence of a coracoid with a crescent - shaped facies articularis clavicularis and an overhanging tuberculum brachiale .
nothura parvula is nested within the extant nothura , depicted in the reduced consensus tree as a paraphyletic taxon with respect to the ( nothoprocta + rhynchotus ) clade ( fig . 1 ) . this position is supported by the presence of pneumatic openings on the impression for the m . sternocoracoidei of the coracoid and a rounded and compact processus supracondylaris dorsalis of the humerus . furthermore , the femur of the ( rhynchotus + nothoprocta ) clade exhibits a strongly curved shaft , an unambiguous apomorphy absent in nothura parvula .
the pleistocene nothura sp . ( excluded from the consensus a posteriori ) takes multiple positions within the open - area taxa taoniscus , nothura , and nothoprocta ( fig . 1 ) . this uncertainty is related to both a lack of morphological data ( missing entries ) for critical characters and conflict in the character - state distribution in the unstable position of the fossil ; however , its original identification as nothura sp . is not contradicted by the present analysis .
the current analysis of osteological and myological characters recovered a tree structure similar to the molecular study of porzecanski ( 2003 ) : the open - area and forest - dwelling tinamous form monophyletic groups in all most - parsimonious hypotheses ( see fig . 1 ) . therefore , the early classification of nothurinae ( open - area tinamous ) and tinaminae ( forest - dwelling tinamous ) proposed by miranda - ribeiro ( 1937 ) is confirmed in the present analysis . the monophyly of the currently recognized polytypic tinamou genera is also recovered with high support : only nothura is recovered as paraphyletic .
the monophyly of the open - area tinamous has been consistently supported by integumentary ( bertelli et al . , 2002 ; bertelli & giannini , 2013 ) , molecular ( porzecanski , 2003 ) , and less inclusive osteological data sets ( bertelli & chiappe , 2005 ) . in the present analysis we added twice as many osteological and myological characters , increasing support for the monophyly of this entire clade , most polytypic genera ( except for nothura ) , and nothurine suprageneric clades .
within nothurinae , the recovered sister - group relationship between the clades ( eudromia + tinamotis ) and ( taoniscus ( nothura ( nothoprocta + rhynchotus ) ) ) ( fig . 1 ) is also congruent with the molecular analysis of porzecanski ( 2003 ) , but such a relationship was previously not recovered in other morphological studies ( bertelli et al . , 2002 ; bertelli & chiappe , 2005 ; bertelli & giannini , 2013 ) . a sister - group relationship between tinamotis and eudromia , by contrast , is uncontroversial , and has been recognized for over a century ( salvadori , 1895 ) ; the basal position of these taxa within nothurinae has , however , not been suggested by previous morphological studies . the present analysis further recovered a clade formed by the remaining open - area tinamous ( taoniscus , nothura , nothoprocta , and rhynchotus ) . given the absence of information on taoniscus , myological apomorphies provide unambiguous support for this clade in only some of the optimal trees .
in summary , osteological data were shown to support most nodes within the phylogenetic tree of tinamous . myological characters also contribute to the support of several nodes , but a number of muscular traits included in the current analysis could not be optimized as synapomorphies because of a lack of information for some taxa ( e . g . tinaminae ) . overall , the new data bring considerable additional evidence in support of well - established groups ( fig . 1 ) .
tnt : tree analysis using new technology , vers . 1 . 1 ( willi hennig society edition )
higher - order phylogeny of modern birds ( theropoda , aves : neornithes ) based on comparative anatomy . ii . analysis and discussion
data set used in the present analysis . characters 12 , 17 , 19 , 24 , 34 , 39 , 42 , 44 , 47 , 54 , 57 , 62 , 69 , 80 , 92 , 95 , 99 , 100 , 101 , 111 , 115 , and 116 were considered additive .
list of morphological characters used in the cladistic analysis . characters for which states could be arranged in a linear transformation series were coded as additive ( non - additive and additive characters are indicated by ‘–’ and ‘ + ’ , respectively ) .
–os supraoccipitale , position of foramen v . occipitalis externa relative to the prominentia cerebellaris ( fig . 3 ) : ventral of the prominentia ( 0 ) ; dorsal , proximal of the crista nuchalis transversa ( 1 ) .
–os supraoccipitale , development of sulcus v . occipitalis externa ( fig . 3 ) : short grooves ( 0 ) ; ventrally extended and curved grooves ( 1 ) . ventrally extended and curved grooves were observed in rhynchotus and nothoprocta .
–os supraoccipitale , crista nuchalis sagittalis ( fig . 3 ) : absent ( 0 ) ; present ( 1 ) . the area of the prominentia cerebellaris of tinamous is typically smooth ( not marked by a crista nuchalis sagittalis ) , with the exception of rhynchotus , tinamotis pentlandii , and nothoprocta taczanowskii , in which its surface is scarred by a marked crest .
–os exoccipitale , processus paroccipitalis ( fig . 3 ) : not developed , flat or poorly developed as a wing - like projection , approaching but not extending below the ventral margin of the otic cavity ( 0 ) ; distinctly prominent , projecting ventrally ( 1 ) . a poorly developed processus paroccipitalis is found in some out - group taxa ( galliformes ) and in all tinamous ( except for rhynchotus and nothoprocta taczanowskii ) . only this tinamou species posses a markedly prominent processus paroccipitalis ; a similar condition is also present in other out - group taxa ( e . g . ratitae ) .
–os exoccipitale , foramen n . vagi , nearer to the foramen n . ophthalmici than to foramen n . hypoglossum ( fig . 3 and 5 ) : no ( 0 ) ; yes ( 1 ) .
+ fossa temporalis ( fig . 4 ) : small temporal notch , smaller than otic cavity , not extending caudally beyond the cotyla quadratica squamosi ( 0 ) ; larger or about the same size as the otic cavity , caudal extension about or beyond the position of the cotyla ( 1 ) . a well - developed fossa temporalis in tinamidae ( condition 1 ) and related palaeognathae was described by pycraft ( 1900 ) ; however , we observed variation in the development of this fossa among tinamous , for example , a distinct small temporal notch enclosed by a rostroventrally orientated processus zygomaticus together with a pointed processus postorbitalis characterizes the skull of the open area tinamous ( with the exception of rhynchotus ) and some out - group taxa . we consider the highly derived skull morphology of apteryx to be non - comparable for this character .
–os squamosum , prominentia suprameatica ( elzanowski , 1987 ) developed as a distinct crest ( fig . 4 ) : absent ( 0 ) ; present ( 1 ) . the prominentia suprameatica flares out from the base of the processus zygomaticus , dorsal of the otic cavity ( elzanowski , 1987 ) . although this structure is absent or minimally developed in most tinamous , it projects as a wing - like crest in some open area taxa such as nothura , nothoprocta , and rhynchotus , among others . in these taxa , the suprameatic prominence is clearly visible both in lateral and ventral view .
–os squamosum , processus zygomaticus ( fig . 4 ) : absent or vestigial ( 0 ) ; present , variably developed ( 1 ) ; present , laterally compressed ( 2 ) . the body of the processus zygomaticus in rhynchotus and nothoprocta ( also in eudromia , although less developed ) is short and laterally compressed , projecting as a crest , which has been tentatively indentified as the area of attachment of the aponeurosis zygomatica ( elzanowski , 1987 ) . this process is also distinctly compressed in the ratites ( e . g . rhea and apteryx ) . a lower ridge is also distinguishable on the processus zygomaticus of other open - area tinamous . see character 33 of mayr ( 2011 ) for conditions in out - group taxa .
–os squamosum , ventral margin of the processus zygomaticus ( fig . 4 ) : straight ( 0 ) ; notched ( 1 ) . the notched margin of the proc . zygomaticus forms a border with the ala parasphenoidalis , roofing the otic cavity in rheidae , rhynchotus , and some species of nothoprocta ( e . g . n . ornata ) . a non - comparable state was assigned to galloanseres ( the process is absent or vestigial in these taxa , see character 7 ) .
–sutura frontoparietalis ( fig . 3 and 4 ) : absent ( 0 ) ; present ( 1 ) ( mayr , 2011 : character 32 ) .
–os frontale , dorsal surface , distinct fossa at midline ( fig . 3 ) : absent ( 0 ) ; present ( 1 ) . the interorbital surface of the skull of most taxa is essentially flat ; only in rhea , eudromia , and tinamotis is it scarred by a rounded depression at midline . this condition differs from that of some anseriformes , where the depression is a longitudinal concavity on the frontal region of the skull ( depressio frontalis of baumel et al . , 1993 ) .
–orbita , fonticuli orbitocraniales ( fig . 4 ) : vestigial or absent ( 0 ) ; present , large foramina present ( 1 ) . only the open area taxa taoniscus and nothura present these rounded openings in the caudal wall of the orbita . because of the highly modified nature of the skull in apteryx , we consider the character non - comparable for this taxon .
–orbita , septum interorbitale extensively ossified ( fig . 4 ) : yes ( 0 ) ; no ( 1 ) . after bertelli and chiappe ( 2005 : character 1 ) . the fonticuli interorbitales are typically present in most tinamous ( only absent in tinamotis and eudromia ) .
+ os mesethmoidale ( fig . 4 ) : does not reach rostrally beyond nasal–frontal hinge ( 0 ) ; extends rostrally immediately beyond the nasal–frontal hinge ( 1 ) ; extends far rostrally ( 2 ) . pycraft ( 1900 : 198 ) discussed this feature and noted that the rostral extension in struthio and rhea exceeded that of other ratites . this feature was coded by mayr ( 2011 ) , albeit only with two states , recognizing the derived one in apteryx and rheidae . cracraft ( 1986 : character 50 ) reported the absence of a ‘largely ossified’ septum in the nasal region ( mesethmoid ) of tinamidae , whereas mayr ( 2011 : character 8 ) noted the absence of this feature in both tinamidae and rheidae . in ichthyornis this character has been inferred from the morphology of the praemaxilla ( clarke , 2004 ) .
–os ectethmoidale ( fig . 4 ) : separated from os lacrimale ( 0 ) ; contacting or fused to os lacrimale , forming a lacrimal–ectetmoid complex ( cracraft , 1968 ) ( 1 ) ; extensively fused with os lacrimale and nasal capsule ( 2 ) . after bertelli and chiappe ( 2005 , character 5 ) . ericson ( 1997 : character 6 ) coded the contact between these bones as present in rheidae ; we find this contact to be absent in all rhea specimens examined ( for a discussion of this character , see cracraft , 1968 : 330 ) . because these bones form part of the highly modified praeorbital region in apteryx , we assigned an additional state ( 2 ) for this taxon .
+ lacrimal–ectethmoid complex ( fig . 4 ) : narrow plate on ventromedial part of the antorbital wall , enclosing a wide and circular opening ( foramen orbitonasale laterale ) ( 0 ) ; wide plate that covers most of the antorbital wall ( foramen orbitonasale laterale narrow ( 2 ) ; plate and enclosed foramen with triangular outline ( 1 ) . bertelli and chiappe ( 2005 , character 6 ) observed variation in the degree of development of the lacrimal–ectethmoid complex within tinamidae . a narrow plate , enclosing a circular foramen orbitonasale laterale characterizes the antorbital area of the skull of the forest tinamous ) . among the open - area groups , this condition is only present in rhynchotus , taoniscus , and the species nothura minor . a non - comparable state was assigned to out - group taxa in which the lacrimal–ectethmoid complex is absent ( character 18 ) .
–os lacrimale , dorsal surface of the head ( fig . 3 ) : expanded ( 0 ) ; very narrow ( 1 ) . in tinamous , the lacrimal contacts the frontal and nasal ; this condition is similar in apteryx and galliformes ; by contrast , in rhea and anhimidae it articulates only with the nasal ( cracraft , 1968 ) . the development of the dorsal surface of the lacrimals varies across species in tinamous . only in the open - area tinamous rhynchotus , taoniscus , nothura darwinii , nothura maculosa , and most species of nothoprocta ( except nothoprocta cinerascens ) are the lacrimals distinctively narrower than in any other tinamous or out - group taxa ( state 1 ) . this character was scored as non - applicable for apteryx , the lacrimals of which are not exposed on the dorsal surface of the skull . in rhea this condition was scored as expanded ( 0 ) , because the main body of the lacrimal is dorsally wide , regardless of the presence of a supraorbital process projecting caudolaterally .
–os parasphenoidale , processus basipterygoideus , developed as a large and ovoid facet for articulation with the pterygoid ( mayr 2011 : character 24 ; fig . 5 ) : no ( 0 ) , yes ( 1 ) .
–os parasphenoidale , processus basipterygoideus ( fig . 5 ) : stout process ( 0 ) ; elongated ( 1 ) . the presence of a slender processus basipterygoideus is the most widely distributed condition within tinamous . because of morphological differences ( character 22 ) , we coded this character as non - comparable for galloanseres .
+ os parasphenoidale , processus parasphenoidalis medialis , aspect in caudal view ( fig . 3 ) : absent ( 0 ) ; weakly developed ventrally as a low bump ( 1 ) ; prominent , developed as a conspicuous bony knob ( 2 ) . mayr ( 2011 ; character 30 ) distinguished only two states for this character . within tinamous , these processes are prominent only in nothoprocta taczanowskii , and are poorly developed in nothocercus . processus mediales parasphenoidales of the lamina parasphenoidalis are equivalent to the ‘mamillary processes’ of pycraft ( 1900 ) , the ‘basal tubercles of the basitemporal plate’ of lee et al . ( 1997 ) , as well as to ‘medial processes of the parashenoid’ ( bock , 1963 ) .


microderes is a genus of beetles in the family carabidae , containing the following species : microderes akbensis jedlicka , 1958 microderes brachypus ( steven , 1809 ) microderes breviformis ( tschitscherine , 1898 ) microderes diversopunctatus ( solsky , 1874 ) microderes intermittens ( solsky , 1874 ) microderes namanganensis ( heyden , 1885 ) microderes nanulus ( tschitscherine , 1898 ) microderes subtilis ( tschitscherine , 1898 ) microderes taschketensis ( jedlicka , 1958 ) microderes undulatus ( gebler , 1841 )

microderes

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microderes ( microderes ) brachypus steven , 1809 e : ab ar kz st uk a : in iq ki kz mg td tr uz xin ( löbl , smetana , 2003 ) .
microderes is a genus of beetles in the family carabidae , containing the following species : . . .
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