oligobuninae is an extinct subfamily of the family mustelidae .

the subfamily was described by j. a. baskin in 1998 ; of the genera that he assigned to this clade , eight are recognized today - brachypsalis , megalictis , oligobunis , potamotherium , promartes , zodiolestes , floridictis and parabrachypsalis - representing fifteen separate species .

oligobuninae

oligobuninae is an extinct subfamily of the family mustelidae known from miocene deposits in north america .
citation : valenciano a , baskin ja , abella j , pérez - ramos a , álvarez - sierra má , morales j , et al . ( 2016 ) megalictis , the bone - crushing giant mustelid ( carnivora , mustelidae , oligobuninae ) from the early miocene of north america . plos one 11 ( 4 ) : e0152430 . urltoken
the new specimens of megalictis ferox described here ( f : am 54079 , f : am 25430 and amnh 54076 ) give us a broader understanding of the morphology of m . ferox and lead us to conclude that the holotypes of both m . ferox ( amnh 12880 ) and aelurocyon brevifacies ( cm 1590 ) are conspecific and thus the latter should be subsumed into m . ferox . we argue that there are 3 species ascribed to megalictis : m . ferox , m . frazieri and m . simplicidens . however , the fourth potential congener , “m” . petersoni , might be best ascribed to a different genus . our cladistic analysis suggests that m . ferox is the sister taxon of the clade composed by m . simplicidens — m . frazieri . our phylogenetic hypothesis supports the subfamily oligobuninae as being a stem mustelid .
megalictis ferox matthew , 1907 [ 1 ] is a giant mustelid of the subfamily oligobuninae and belongs to the paraphyletic group of “paleomustelids” [ 2 ] . it lived in the early miocene during the late arikareean ar4 north american land mammal age 22 . 7–18 . 5 mya [ 3 , 4 ] of the central great plains of united states in the states of nebraska , south dakota , and wyoming [ 1 , 5 – 7 ] . the ar4 lithostratigraphic units containing giant oligobunines have been revised . hunt [ 8 ] named the anderson ranch formation for the terminal formation of the arikaree group in nebraska and wyoming formerly referred to as the upper harrison beds of peterson [ 5 , 9 ] and the lower marsland formation of schultz [ 10 ] . the black bear formation replaces the upper rosebud formation of south dakota [ 11 ] .
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full reference : j . a . baskin . 1998 . mustelidae . evolution of tertiary mammals of north america 152 - 173
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copyright : © 2016 valenciano et al . this is an open access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .
data availability : all relevant data are within the paper and its supporting information files .
funding : a . v . has received funding from the european union’s seventh framework programme ( fp7 / 2007 - 2013 ) under grant agreement n° 226506 ( synthesys ; se - taf - 3637 ) , the usc school of medicine ( columbia , south carolina , usa ) , the amnh ( collection study grant program 2014 ) and the international travel grant 2015 from the vertebrate paleontology department of flmnh from uf . a . v . is researcher in formation in the csic program jae - pre _ cp2011 ( csic program
junta para la ampliación de estudios
) , co - funded by the european social fund . a . p . r . is a pre - doctoral fpi fellowship ( bes - 2013 - 065469 ) of the project cgl2012 - 37866 . this study was also supported by the spanish ministerio de economía y competitividad ( research project cgl2015 - 68333 , cgl2011 - 28877 and cgl2011 - 28681 ) , the research group bsch - ucm 910607 and university of south carolina school of medicine ( columbia , south carolina , usa ) .
abbreviations : acm , amherst college beneski museum of natural history , massachusetts , usa ; ahr , comparative anatomy research collection , university of south carolina school of medicine , columbia , usa ; amnh , american museum of natural history , division of paleontology and division of mammalogy , new york , usa ; cm , carnegie museum of natural history , pittsburgh , usa ; f : am , collection housed in the frick collection of the division of paleontology , amnh , new york , usa ; lgput , laboratory of geology and palaeontology , university of thessaloniki , greece ; mncn , museo nacional de ciencias naturales madrid , spain ; mncncomp , comparative anatomy research collection of paleobiology department of museo nacional de ciencias naturales madrid , spain ; nrm , naturhistoriska rikmuseet , stockholm , sweden ; pin , palaeontological institute , russian academy of sciences , moscow , russia ; sam - pql , iziko south african museum , cape town south african museum , south africa ; tmm , texas memorial museum at the vertebrate paleontology laboratory , the jackson school of geosciences , university of texas , austin , usa ; uf , vertebrate paleontology collection of the florida museum of natural history ( flmnh ) , university of florida , gainesville , usa ; unsm , university of nebraska state museum , lincoln , nebraska , usa ; usnm , united states national museum of natural history , smithsonian institution , washington , d . c . , usa
megalictis ferox [ 1 ] was described from the black bear formation , stanley county , south dakota , usa . a second giant oligobunine , aelurocyon brevifacies peterson , 1907 [ 5 ] , was described from the niobrara canyon local fauna , anderson ranch formation in sioux county , nebraska , usa . hunt and skolnick [ 7 ] established that the actual publication date for a . brevifacies was one week after matthew described m . ferox in 1907 , not in 1906 as indicated in the journal . after these initial descriptions , riggs [ 6 ] described new cranial and postcranial material of both taxa . hunt and skolnick [ 7 ] synonymized megalictis ferox , aelurocyon brevifacies , and the large oligobunine mustelid paroligobunis simplicidens ( peterson , 1907 ) [ 5 ] .
here , we describe an important unpublished sample of craniomandibular remains of megalictis ferox ( f : am 25430 , f : am 54079 , and amnh 54076 ) , housed at the american museum of natural history ( new york , usa ) . although f : am 25430 and f : am 54079 were found in the late 1930s and have been used to obtain metric , morpho - functional and phylogenetic data ( e . g . , [ 2 , 7 , 12 – 16 ] ) , they have never been fully described . therefore , the main objective of the present paper is to describe these unpublished skulls and mandibles , and provide new data on the taxonomy and systematics of the genus in order to shed new light on the paleobiology of megalictis .
dental nomenclature follows ginsburg [ 17 ] and smith and dodson [ 18 ] . anatomical descriptions are based primarily on scapino [ 19 ] , turnbull [ 20 ] , barone [ 21 , 22 ] , waibl et al . [ 23 ] , evans and de lahunta [ 24 , 25 ] , and hartstone - rose et al . [ 26 ] . the terminology conforms to the standard of the nomina anatomica veterinaria [ 23 ] with the exception of the masseter and temporalis muscle complexes for which we follow hartstone - rose et al . [ 26 ] . the megalictis material ( figs 1 – 4 ) has been compared to all the other material of megalictis and paroligobunis on the basis of published descriptions , figures , measurements and photographs . we have re - measured the dentition of amnh 12880 and 22632 ( cast of cm 1590 ) measured initially by matthew [ 1 ] and peterson [ 5 ] and completed the measures of paroligobunis petersoni loomis , 1932 [ 27 ] using a cast tmm 40966–1 . measurements were made using mitutoyo absolute digital calipers to the nearest 0 . 1 mm ( tables 1 and 2 ) .
a lateral view ; b rostral view ; c dorsal view ; d caudal view ; e ventral view . scale bar equals 5 cm .
a right mandible lateral view ; b occlusal view ; c left mandible lingual view of lower dentition . scale bar equals 5 cm .
a1–4 cranium f : am 54079 , lateral view ( a1 ) , dorsal view ( a2 ) , ventral view ( a3 ) , and caudal view ( a4 ) ; b1–3 right hemimandible f : am 54079 , lateral view ( b1 ) , medial view ( b2 ) , and occlusal view ( b3 ) ; c1–3 left hemimandible f : am 54079 lateral view ( c1 ) , medial view ( c2 ) , and oclussal view ( c3 ) ; d1–3 right hemimandible of amnh 54076 , lateral view ( d1 ) , medial view ( d2 ) , and occlusal view ( d3 ) . scale bar equals 5 cm .
a , and b megalictis ferox holotype amnh 12880 , lateral view ( a ) , ventral view ( b ) ; c , and d megalictis ferox cm 1590 ( genotype of aelurocyon brevifacies ) , lateral view ( c ) , ventral view ( d ) ; megalictis ferox f : am 25430 lateral view ( e ) , ventral view ( f ) ; g , and h megalictis ferox f : am 54079 lateral view ( g ) , ventral view ( h ) . scale bar equals 5 cm . c and d courtesy of the carnegie museum of natural history .
f : am 25430 ( figs 1 and 2 , s1 video ) : relatively complete skull with i1 - 3 , c , p1 - 4 and m1 - 2 , missing only its left zygomatic arch , a broken frontal area plus a portion of the right parietal region missing and filled with plaster , a hole in its right parietal bone , and a complete mandible with i1 - 3 , c , p1 - 4 and m1 - 2 ; f : am 54079 ( fig 3 , s2 video ) : right side of a partial skull without the premaxilla , with worn c , p2 - m1 and partial mandibles with a nearly complete right one with c - m2 and a broken mandibular symphysis and a left one just with the mandibular corpus preserved and a broken p2 , and a complete both p3 and m1 ; amnh 54076 ( fig 3 ) : partial mandibular corpus with m1 - 2 .
in order to better understand the phylogenetic relationships of the oligobunines megalictis ferox ( amnh 12880 , cm 1590 , f : am 25430 and f : am 54079 ) , m . simplicidens ( = paroligobunis simplicidens ) ( cm 1553 and cm 2389 ) , m . frazieri ( = paroligobunis frazieri ) ( uf 23928 ) , “ m . ” petersoni ( = paroligobunis petersoni ) ( acm 2011 ) , and oligobunis crassivultus ( amnh 6903 ) , we have performed a cladistic analysis ( fig 5 ) including 18 taxa ( m . ferox is represented in the analysis as 4 separate operational taxonomic units ( otu ) ) and 73 equally weighted and unordered craniomandibular characters ( s1 – s3 appendices ) . cladistic analysis was performed using in paup * 4 . 0b10 [ 38 ] . the analysis was rooted using c . lupus as the outgroup .
searches were performed using the branch and bound and a bootstrap analysis through 1000 replicates to test the clade support in the analysis . the outgroup was c . lupus . strict consensus tree of 6 trees ( length 194 steps , consistency index ( ci ) = 0 . 41 , retention index ( ri ) = 0 . 65 ) for knowing the relationships between the different specimens of megalictis ferox , megalictis simplicidens , megalictis frazieri , “ megalictis ” petersoni , oligobunis crassivultus , and a sample of extant musteloids and a canid . numbers below nodes are bremer indices , and numbers above nodes are bootstrap support percentages ( only shown when ≥ 50 ) . character / taxa matrix is detailed in the s1 – s3 appendices . silhouette of megalictis ferox based on hunt and skolnick [ 7 ] , silhouette of megalictis simplicidens , megalictis frazieri , “ megalictis ” petersoni and oligobunis crassivultus based on megalictis ferox but rescaled according the size of the dentition .
virtual models of the mandibles and skulls of megalictis ferox ( f : am 25430 , f : am 54079 , amnh 12880 and amnh 22632 ) as well as megalictis frazieri uf 23928 and megalictis simplicidens ( cast of cm 1553 ) were derived by means of a 3d nextengine hd laser surface scanner ( s1 – s6 videos ) .
diagnosis : large to giant size mustelid ; robust mandible with a high , wide and distally curved ascending ramus ; deep masseteric fossa with a stout crest that extends from the dorsal border of the coronoid process to below the m2 ; robust dentition ; p1 present ; p2–4 with distal cingula high - crowned ; p4 relatively enlarged with mesial and distal accessory cuspids ; m1 trigonid widened , with a strong lingual concavity between the paraconid and protoconid ; m1 metaconid reduced to absent , present in the older and smaller forms and absent in the giant forms , m1 talonid low and narrow with a short , trenchant and labially located hypoconid ; and m2 with reduced metaconid .
type species : megalictis ferox matthew , 1907 p1 . ii , fig . 1 [ 1 ]
included species : megalictis simplicidens ( = paroligobunis simplicidens ) ( peterson , 1907 ) [ 5 ] and megalictis frazieri ( = paroligobunis frazieri ) ( frailey , 1978 ) [ 28 ] .
synonyms : senior subjective synonym [ 7 ] of aelurocyon brevifacies peterson , 1907 , p . 68 [ 5 ] , “upper harrison formation” , sioux county , nebraska and paroligobunis peterson , 1910 [ 29 ] . hunt and skolnick [ 7 ] synonymized megalictis , aelurocyon , and paroligobunis simplicidens into a single , sexually - dimorphic chronospecies m . ferox . this hypothesis has been generally accepted ( e . g . , [ 3 , 13 , 42 ] ) .
megalictis ferox , hunt and skolnick , 1996 ( pars ) . [ 7 ]
holotype : amnh 12880 , a partial reconstructed skull ( fig 4 , s3 video ) with right p4 , m1 - 2 , a fragmented right mandible with almost complete coronoid process , m1 trigonid and m2 , and very few postcranial remains figured by matthew , 1907 , p . 196 , fig . 10–13 , 15 [ 1 ] .
type locality : rosebud 22 , porcupine butte , black bear formation , stanley county , south dakota .
other localities : rosebud 5 , porcupine butte , stanley county , south dakota , usa ( amnh 12881 ) ; niobrara canyon local fauna , sioux county , nebraska , usa ( cm 1590 ) , “high daemonelix beds” , niobrara canyon local fauna , sioux county , nebraska , usa ( f : am 25430 ) ; j - m district , south of lusk , goshen county , wyoming , usa [ 6 ] ; “high brown sand” , 16 mile district , goshen county , wyoming , usa ( f : am 54079 ) ; 8 north of lusk , goshen county , wyoming , usa ( f : am 54076 ) .
age : upper part of the anderson ranch formation and its equivalents , south dakota , nebraska , and wyoming , late late arikareean ( ar4 ) , 22 . 7–18 . 5 mya [ 4 ] early miocene .
diagnosis : baskin [ 2 ] diagnosed of aelurocyon brevifacies ( which he considered the senior subjective synonym of megalictis ferox because of the presumed earlier publication date at the time he submitted the chapter ) . new or revised characters follow . megalictis ferox is the largest of the oligobunines ; coronoid process high and caudally curved ; enlarged masseteric fossa with a robust crest extending from the dorsal border of the coronoid process to below the m2 ; laterocaudal area of the ventral edge of the mandibular corpus laterally projected ; p2 with distal accessory cusp ; robust p3 ; robust p4 with strong parastyle and protocone ; p4 carnassial notch present ; m1 with enlarged stylar area ; m2 with paracone and protocone ; p2–4 with high - crowned distal cingula ; p3 with mesial and distal accessory cuspid ; p4 relatively enlarged with presence of mesial accessory cuspid and stout distal accessory cuspid ; m1 trigonid widened ; m1 with strong lingual concavity between paraconid and protoconid ; m1 protoconid higher than paraconid ; m1 hypoconid short , trenchant and buccally located ; m1 with a lingual cingulum in the entoconid position ; m2 reduced with metaconid .
differential diagnosis : megalictis ferox differs from m . simplicidens , m . frazieri , “m . ” petersoni and oligobunis crassivultus in its larger size , m1 without metaconid and m1 talonid with a closed lingual morphology with a lingual cingulum between the metacristid and entocristid . additionally , it differs from m . simplicidens and m . frazieri in having a higher and more robust mandibular symphysis , a reduced p2 and a more robust p4 and m1 . it further differs from “m . ” petersoni in much larger size and p3–4 with mesial accessory cuspids . it further differs from oligobunis crassivultus in having a more rectangular p2 , smaller m1 than p4 , enlarged m1 stylar area , higher paracone than metacone on the m1 , reduced p2 , p2–3 high - crowned distal cingula , more developed p3 distal accessory cuspid , relatively enlarged p4 , and higher protoconid than paraconid on the m1 .
comments : specimens that can be referred to m . ferox s . s . are from the latest arikareean ( ar4 ) upper part of the anderson ranch formation and its equivalents .
a nearly complete skull with i1 - 3 , c , p1 - 4 and m1 - 2 ( fig 1 , s1 video ) and a complete mandible with i1 - 3 , c , p1 - 4 and m1 - 2 ( fig 2 , s1 video ) . the left zygomatic arch is missing . part of the frontal and a region of the right parietal bones are missing and filled with plaster . there is a subrectangular and anthropogenic hole in its right parietal bone located above the most caudal area of the zygomatic arch .
locality : “high daemonelix beds” , anderson ranch formation , niobrara canyon local fauna , sioux county , nebraska , usa .
skull and upper dentition : the very well preserved skull f : am 25430 ( fig 1 ) has a basicranial length of 189 . 5 mm . it is slightly domed dorsally at the frontal bone , the bullae are broken and the left zygomatic arch is missing . in general terms the skull is high , domed with a short rostrum and high snout ( fig 1a ) . the nasal aperture is large ( fig 1b ) , and the crushed nasal bones are robust . they are crushed in the mid - sagittal plane and anteriorly the left nasal bone is partially above the right one . the reconstructed frontal region is quite domed . the interorbital region is broad . the postorbital processes are absent . the moderately developed infraorbital foramen is rounded and located above the distal accessory cusp of the p3 . the rostral margin of the orbit ends at the level of the distal margin of the p4 paracone . the orbits are large and rounded . the lacrimal foramen is rounded and relatively large . the sagittal crest is moderately developed and extends caudally where it divides into the nuchal crests , forming a y - pattern ( fig 1a , 1c and 1d ) . in lateral view , the outline of the skull is convex in the temporal region and concave between the temporal bone and nuchal crests .
the zygomatic arches are robust , especially caudally near the glenoid cavity . both m . masseter pars superficialis and m . masseter pars profunda have their origin on the ventrolateral side of the zygomatic arch . the frontal processes of the zygomatic arches are triangular and dorsoventrally high .
mandible and lower dentition : the mandible of f : am 25430 is very robust ( fig 2a and 2b ) . it has a total length of 149 . 0 mm . the tooth row is slightly convex and is aligned with the articular process . the mandibular corpus is high and robust . the ventral margin is convex at the level of the m1 . there is single rounded mental foramen under p2 . the ascending ramus is tall and rostrocaudally broad ( fig 2a ) . its tip is distally oriented . the coronoid process is laterally rotated with an angle of ~ 75 degrees , compared to the articular process . there is a robust crest from the dorsal border of the coronoid to beneath the m2 where the tendon of the m . temporalis is attached . this area is especially enlarged and laterally projected around the area of the m2 ( fig 2a and 2b ) . the masseteric fossa is large and deep . its rostral margin lies at the level of talonid of m1 , and ventrolaterally is limited by a strong area where the m . masseter pars superficialis and m . masseter pars profunda insert . the articular process is large and robust . the angular process is robust and shows a medial crest for the muscular attachment of the m . pterygoideus medialis .
the lower dentition ( 3 / 1 / 4 / 2 ) is also preserved in its entirety ( fig 2 ) . the lower incisors are heavily worn . the canine is large , stout and markedly curved distally ( fig 2a and 2b ) . it has a swollen base and is oval in cross - section . the p1 is oval , single - cusped and distally wide ( fig 2b ) . the p2–4 are stout , subrectangular and wider distally . these premolars have strong cingula at their bases , and the distal cingula are high - crowned . the p2 has a single messially - located cuspid . the p3 has a low mesial accessory cuspid and a more developed distal one . the p4 is the largest lower premolar and has more strongly developed mesial and distal accessory cuspids . the m1 is a relatively short and stout tooth ( fig 2 ) . the very robust trigonid occupies almost three fourths of the total length of the tooth , with the greatest width at the base of the protoconid . the paraconid is lower than the protoconid and there is no metaconid . the m1 shows a markedly lingual concavity in the base of the crown between the trigonid cuspids ( fig 2b ) . the stout talonid lacks an entoconid . the hypoconid is low , trenchant and buccally located . there is a smooth cristid from the top of the protoconid to the hypoconid that encloses a deep lingual depression ( fig 2c ) . the m2 is rounded and low ( fig 2b ) . the paraconid is low , and located in the mesial corner . the protoconid is the highest cuspid , located buccally in the middle of the tooth . the metaconid is situated over the lingual corner . it is less developed than the protoconid . the hypoconid is low and located in the distal corner . there is a cingulum around the whole tooth .
partial skull with worn c , p2–m1 and partial mandible with worn p1–4 and m1–2 ( fig 3a1–4 , 3b1–3 and 3c1–3 , s2 video ) .
locality : “high brown sand” , 16 mile district , anderson ranch formation , goshen county , wyoming , usa .
skull and upper dentition : the skull f : am 54079 ( fig 3a1–4 ) only preserves its right side . it has a maximum length of 180 . 2 mm . the premaxilla is missing , so the basicranial length is unknown . in general terms , f : am 54079 resembles f : am 25430 ( fig 1 ) . the frontal bone and dorsal area of the parietal bone are absent ( fig 3a2 ) . the zygomatic arches are more robust than those of f : am 25430 , especially in the rostral and the central part of the arches , and the origin of m . masseter pars superficialis and m . masseter pars profunda are also more developed . however , the frontal processes of the zygomatic arches are lower than those of f : am 25430 . the glenoid fossa is stout with a very well developed postglenoid process ( fig 3a3 ) . the complete right auditory bulla is large and swollen . the external auditory meati are rounded . the postglenoid foramen , the rostral foramen lacerum and the foramen ovale are similar to those of f : am 25430 . the mastoid process is also robust and expanded . the right occipital condyle is preserved but the caudodorsal area of the skull is not . the paroccipital process is triangular , stout and caudally oriented ( fig 3a1 and 3a3–4 ) .
c , p2–4 and are preserved . the p1 is missing . they are more worn than are those of f : am 25430 . the c has a large lingual wear facet . the morphology of p2–4 ( fig 3a3 ) is almost identical to that of f : am 25430 . the p3 is more quadrangular than that of f : am 25430 , but the mesiolingual corner of the p3 is missing . the p4 paracone , protocone and metastyle are greatly - worn ( fig 3a3 ) . the m1 ( fig 3a3 ) has the same development of the cusps as that found in f : am 25430 , and shows a very similar morphology as that of amnh 12880 . the m2 and its alveoli are not preserved .
mandible and lower dentition : the right hemimandible ( fig 3b1–3 ) has a fragmented corpus that is missing its symphyseal end but includes a complete ascending ramus with p1–4 and m1–2 . its morphology is almost identical to that of f : am 25430 . the left hemimandible ( fig 3c1–3 ) is missing its ascending ramus but includes a complete mandibular corpus , a complete p1 , a fragmented p2 , a highly worn p3 , a complete m1 and a fragmented m2 . the p1–4 and m1 are almost identical to those of f : am 25430 though there is more substantial occlusal wear in p2–4 and m1 than in f : am 25430 . the m2 is oval and has a more developed metaconid than the m2 of f : am 25430 .
locality : 8 north of lusk , goshen county , anderson ranch formation , wyoming , usa .
mandible and lower dentition : amnh 54076 is a fragmented mandibular corpus missing its symphysis ( fig 3d1–3 ) . it has roots for p2–3 , and complete m1–2 . the mandibular corpus is high and robust . the m1 is identical to those of f : am 54079 and f : am 25430 . it has a stout trigonid , and a low talonid composed of a trenchant hypoconid , lingually located and a lingual depression . the m2 is rounded and low . it has a distinguishable protoconid and metaconid , and a continuous basal cingulum .
hunt and skolnick [ 7 ] did not consider the other two species referred to paroligobunis : the small p . petersoni loomis , 1932 [ 27 ] and p . frazieri frailey , 1978 [ 28 ] . as discussed below , we consider the material referred to both p . simplicidens and p . frazieri to be valid species : megalictis frazieri and m . simplicidens .
the results of the cladistic analysis indicate that the specimens we assign to m . ferox form a monophyletic group ( fig 5 ) . we agree with hunt and skolnick [ 7 ] in that m . ferox and a . brevifacies are the same taxon , and that m . ferox has priority . morphologically , the specimens f : am 54079 , f : am 25430 and amnh 54076 , as well as cm 1590 and amnh 12880 , are practically identical to each other ( figs 1 – 4 ) . f : am 54079 differs from f : am 25430 and cm1590 in having a more robust p3 and a relatively longer m2 . cm 1590 has a reduced lingual expansion of p3 and a stronger parastyle of p4 than f : am 54079 , f : am 25430 and amnh 12880 . the morphology of f : am 25430 is clearly different from the skull of amnh 12880 , and shows that the reconstructed parts of the latter were incorrect , in which the temporal , frontal and a part of the zygomatic arch bones are misinterpreted ( fig 4a and 4b ) . f : am 25430 allows us to complete the knowledge about the morphology of the skull of m . ferox and showing that the holotype of m . ferox ( amnh 12880 ) and the holotype of a . brevifacies ( cm 1590 ) belong to the same species . consequently , f : am 54079 , f : am 25430 and amnh 54076 should be assigned to m . ferox . we agree with hunt and skolnick [ 7 ] that the difference observed in the specimens of m . ferox can be explained by intraspecific variability ( sexual dimorphism and intrapopulational differences ) or small temporal differences .
megalictis ferox ( figs 1 – 4 ) is characterized by several traits : long external auditory meatus ; high and caudally curved coronoid process ; enlarged masseteric fossa with a robust crest from the dorsal border of the coronoid process to just beneath the m2 ; latero - caudal area of the ventral edge of the mandibular corpus is laterally projected , with the ventral edge of the angular process also laterally projected ; i3 is enlarged ; p2 with a distal accessory cusp ; robust p3 ; robust p4 with carnassial notch ; enlarged stylar area of m1 , and a m2 with paracone and protocone differentiated ; p2–4 distal cingula high - crowned ; distal accessory cuspid on p3 ; relatively enlarged p4 with a stout mesial accessory cuspid ; relatively stout m1 with a widened trigonid , a strong lingual concavity between the paraconid and protoconid , no metaconid , protoconid higher than paraconid , with a short , trenchant and buccally located hypoconid and a lingual rim in the entoconid position ; reduced m2 with a metaconid .
all the three species that have been referred to paroligobunis ( fig 6 ) are known from limited material . the genotype of paroligobunis , megalictis simplicidens ( cm 1590 , peterson , 1907 , 1910 ) [ 5 , 29 ] comes from the “agate stock farm” , sioux county nebraska . the exact locality is unknown and it is either from the harrison formation ( ar3 ) or the basal part of the anderson ranch formation [ 7 ] . additional material first referred to p . simplicidens [ 29 ] and later to megalictis ferox [ 7 ] is from quarry 3 , beardog hill , agate fossil beds national monument , from the basal part of the anderson ranch formation . the small “m” . petersoni ( loomis , 1932 ) [ 27 ] is from a locality near van tassel , wyoming , “upper harrison beds” ( = anderson ranch formation ) and p . frazieri frailey , 1978 [ 28 ] is from the sb - 1a local fauna , florida , latest oligocene , early late arikareean ( ar3 ) . hunt ( in tedford et . al , 2004 : p . 205 [ 3 ] ) recognized that “‘ paroligobunis’ frazieri is an earlier form preceding the late arikareean species of megalictis ” .
( a ) megalictis simplicidens , type specimen , cm1553 ( peterson , 1907 ) [ 5 ] , lateral view of the mandible , ( b ) megalictis simplicidens cm 1553 ( peterson , 1907 ) [ 5 ] , medial view , ( c ) megalictis simplicidens cm 2389 ( peterson , 1910 ) [ 29 ] , lateral view of the mandible , ( d ) megalictis simplicidens cm 2389 ( peterson , 1910 ) [ 29 ] , medial view , ( e ) megalictis frazieri ( frailey , 1978 ) [ 28 ] , holotype uf 23928 , lateral view of the mandible , ( f ) megalictis frazieri ( frailey , 1978 ) [ 28 ] , uf 23928 , medial view , ( g ) “ megalictis” petersoni ( loomis , 1932 ) [ 27 ] , holotype acm 2011 , lateral view of the mandible , ( h ) “ megalictis” petersoni ( loomis , 1932 ) [ 27 ] acm 2011 , medial view , ( i ) megalictis simplicidens cm1553 ( peterson , 1907 ) [ 5 ] , occlusal view , ( j ) megalictis simplicidens cm 2389 ( peterson , 1910 ) [ 29 ] , occlusal view , ( k ) megalictis frazieri ( frailey , 1978 ) [ 28 ] , uf 23928 , occlusal view , ( l ) “ megalictis” petersoni ( loomis , 1932 ) [ 27 ] acm 2011 , occlusal view . scale bar equals 5 cm . a - d , i and j courtesy of the carnegie museum of natural history . e - f and k courtesy of the florida museum of natural history . g - h and l , beneski museum of natural history at amherst college , courtesy of the trustees of amherst college .
there are no derived characters uniting the three named species of paroligobunis that are not shared with megalictis ( s2 appendix ) . our phylogenetic analysis ( fig 5 ) shows that these three species are paraphyletic with m . ferox . the larger p . frazieri and p . simplicidens are both referred to megalictis . the differences in morphology and size between the three species of megalictis with respect to “ m . ” petersoni ( fig 6 ) suggest that “ m . ” petersoni could be excluded from the genus megalictis .
megalictis simplicidens and m . frazieri ( fig 6 ) resemble m . ferox in several characters , such as a high , wide and distally curved ascending ramus , and a deep masseteric fossa with a robust crest that extends from the dorsal border of the coronoid process to below the m2 . both taxa have a p1 , the distal cingula of p2–4 are high - crowned , and the p4 is relatively enlarged with mesial and distal accessory cuspids . the m1 trigonid is widened , with a strong lingual concavity between the paraconid and protoconid , a low , and narrow talonid with a short , trenchant and labially located hypoconid , and a reduced m2 with presence of a metaconid . however they differ from m . ferox in having a non - reduced p2 , the presence of a stout m1 metaconid , relatively more slender p4 and m1 , m1 talonid with an open lingual morphology between the metacristid and entocristid , and a lower and more slender mandibular symphysis .
hunt and skolnick [ 7 ] partially described and measured some of the unsm and cm specimens of megalictis from the basal part of the anderson ranch formation at beardog hill that we refer to m . simplicidens . aside from their more primitive morphology ( e . g . , presence of a metaconid on m1 ) , they are smaller than m . ferox from the upper anderson ranch formation . the upper and lower dental measurements indicate a size similar to g . gulo .
megalictis frazieri ( fig 6e , 6f and 6k ) differs from m . simplicidens ( fig 6a–6d , 6i and 6j ) in having a less massive mandible and a more distinctive distal cingulum with a higher crown in p2–4 than m . simplicidens . the c and p2 of m . frazieri are also more robust . the m1 hypoconid is higher and the talonid is relatively larger , slightly basined with a very low internal rim .
“megalictis” petersoni ( fig 6g , 6h and 6l ) differs from m . simplicidens and m . frazieri in the absence of mesial accessory cuspids on p3–4 , a relatively stouter p4 with a shorter mesial part and a relatively more robust m1 with a taller and stouter metaconid .
metrically the new megalictis ferox sample described above ( f : am 54079 , f : am 25430 and amnh 54076 ) together with amnh 12880 and cm 1590 form a single picture of m . ferox with dental biometric variability similar to the largest extant terrestrial mustelids gulo and mellivora ( figs 7 and 8 ) . however , if m . simplicidens is considered as a synonym of m . ferox , this variability exceeds the extant one . such variability is much more pronounced when all the specimens of m . simplicidens , m . frazieri and the small “ m . ” petersoni ( figs 7 and 8 ) are included .
a life appearance ; b , reconstructed skull and mandible ; c , skull and mandible f : am 25430 . artwork by adam hartstone - rose .
the preservation of the of m . ferox specimen f : am 25430 represents by far the most complete and best preserved craniomandibular specimen of any giant mustelids . based on the size of the skull , m . ferox emerges as the largest terrestrial mustelid ever known–even larger than the extinct late miocene giant mustelid ekorus , eomellivora , and plesiogulo [ 13 , 32 , 33 , 35 , 37 , 70 ] . this new material sheds light on a new paleobiological interpretation of megalictis as a hyena - like , bone - crushing mustelid , instead of the cat - like ecomorphotype previously ascribed to the genus .
s1 table . list of the extant specimens of carnivorans used in this paper .
s1 video . video of the cranium and mandible of megalictis ferox f : am 25430 .
s2 video . video of the cranium and mandible of megalictis ferox f : am 54079 .
s3 video . video of the cranium and mandible of the holotype of megalictis ferox amnh - 12880 .
s4 video . video of the cranium and mandible megalictis ferox amnh - 22632 ( cast of cm 1590 ) .
s5 video . video of the mandible of megalictis simplicidens ( cm 1553 ) and megalictis frazieri uf 23928 .
s6 video . video of the reconstructed head of megalictis ferox f : am 25430 .
conceived and designed the experiments : av jm ahr . performed the experiments : av jab jm . analyzed the data : av jab ja maas jm ahr . contributed reagents / materials / analysis tools : av ja apr . wrote the paper : av jab ja maas jm ahr .
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the subfamily was described by j . a . baskin in 1998 ; of the genera that he assigned to this clade , six are recognized today -
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the subfamily was described by j . a . baskin in 1998 , who assigned seven genera to it -
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the masked mountain tanager ( buthraupis wetmorei ) is a vulnerable species of bird in the tanager family .

this large and colourful tanager is endemic to elfin forest , woodland and shrub in the andean highlands of southern colombia , ecuador and northern peru .

it is generally rare or uncommon , and is threatened by habitat loss .

masked mountain tanager

masked mountain - tanager , reserva ecológica cayambe - coca , ecuador . nov 9 , 2014 .
information on the masked mountain - tanager is currently being researched and written and will appear here shortly .
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black - backed bush - tanager and pale - naped brush - finch in the flock as well .
hilty , s . ( 2018 ) . masked mountain - tanager ( tephrophilus wetmorei ) . in : del hoyo , j . , elliott , a . , sargatal , j . , christie , d . a . & de juana , e . ( eds . ) . handbook of the birds of the world alive . lynx edicions , barcelona . ( retrieved from urltoken on 9 july 2018 ) .
20 cm ; two birds 62 g and 63 g . large , stubby - billed tanager with rather odd plumage pattern . male has crown and nape yellowish - olive , becoming plain olive - green on back and . . .
20 . 5 cm . lethargic , yellowish tanager . yellow - olive crown and nape , yellow forehead and outline to black facial area , olive upperparts with yellow rump , underparts yellow , lightly mottled blackish on the flanks , blue fringing to coverts giving blue shoulder and wing - bar .
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sacc . 2005 and updates . a classification of the bird species of south america . available at : urltoken .
. in colombia , there are records from puracé national park and its environs , cauca , and recently from nariño ( strewe and kreft 1999 ) . it is restricted to the east andes in ecuador ( carchi , napo , morona - santiago , azuay , loja and zamora - chinchipe ) ( krabbe
, and occurs on cerro chinguela , eastern piura , peru . it is generally considered rare or uncommon , but is fairly common at cajanuma , podocarpus national park , which may protect a significant population . nevertheless , it is unlikely that the total population exceeds 5 , 000 birds , and it is inferred to be declining .
the population is placed in the band 2 , 500 - 9 , 999 individuals , equating to 1 , 667 - 6 , 666 mature individuals , rounded here to 1 , 500 - 7 , 000 mature individuals . trend justification : a slow and on - going population decline is suspected , based on rates of habitat loss within its range .
it inhabits very humid elfin forest , scattered bushes , bamboo , giant grasses and dense brush , on the páramo - forest ecotone from 2 , 900 to 3 , 600 m , and possibly higher before human alteration of the treeline .
. in south - west colombia , the proportion of timberline habitat remaining is estimated at less than 10 % , and human pressure is increasing ( p . g . w . salaman
. temperate forest has been replaced with exotic pine plantations near the known site in nariño ( strewe and kreft 1999 ) , and other threats include firewood - gathering and potato cultivation ( p . g . w . salaman
. the area where the species occurs in peru is being heavily deforested and burned for agriculture and cattle ranching ; fragmentation is severe around cerro chinguela ( f . angulo
conduct surveys to clarify its distribution . in particular , survey suitable habitat in tabaconas namballe national sanctuary , since it is not present in any protected area in peru ( f . angulo
, and prohibit the burning of páramo in national parks . educate and encourage local people to take a leading role in land - use management and restoration schemes ( fjeldså and kessler 1996 )
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) , in neotropical birds online ( t . s . schulenberg , editor ) . cornell lab of ornithology , ithaca , ny , usa . retrieved from neotropical birds online :
adult near fledgling . shrubbery above treeline . filtered version on moore et al . ( 2013 ) urltoken
in bushes in humid primary treeline forest . reference : lxxa 287 - 297 , - 302 , - 304 , - 310 , - 319 ( butwet6 ) . krabbe & nilsson ( 2003 ) ( isbn 90 - 75838 - 06 - 9 ) . filtered version on moore et al . ( 2013 ) urltoken
low páramo scrub . reference : jn1 . krabbe & nilsson ( 2003 ) ( isbn 90 - 75838 - 06 - 9 ) .
3 birds sunnying themselves in bush tops in humid primary treeline forest . reference : ( lxxa 369 , 402 - 406 , 413 - 415 , 461 - 480 , 484 - 490 , 521 - 527 ) . krabbe & nilsson ( 2003 ) ( isbn 90 - 75838 - 06 - 9 ) .
in bushes in humid primary treeline forest . reference : lxxa 263 - 267 ( butwet5 ) . krabbe & nilsson ( 2003 ) ( isbn 90 - 75838 - 06 - 9 ) .
in bushes in humid primary treeline forest . reference : lxxa 217 - 220 ( butwet4 ) . krabbe & nilsson ( 2003 ) ( isbn 90 - 75838 - 06 - 9 ) .
female . collected ( mecn 6614 ) . upper edge of humid forest . reference : lxiia 253 - 258 ( butwet3 ) . krabbe & nilsson ( 2003 ) ( isbn 90 - 75838 - 06 - 9 ) . filtered version on moore et al . ( 2013 ) urltoken
sonogram images © xeno - canto foundation . sonogram images share the same license terms as the recording they depict .
classified as vulnerable ( vu ) on the iucn red list ( 1 ) .
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recommended citation birdlife international ( 2018 ) species factsheet : tephrophilus wetmorei . downloaded from urltoken on 09 / 07 / 2018 . recommended citation for factsheets for more than one species : birdlife international ( 2018 ) iucn red list for birds . downloaded from urltoken on 09 / 07 / 2018 .
c andes of sw colombia ( quindío , and s on both slopes in cauca and huila to nariño ) , s through ecuador ( mostly on e slope , from carchi , w napo and w morona - santiago s to e loja and s zamora - chinchipe ) and adjacent nw peru ( along piura–cajamarca border ) .
notably quiet . infrequently heard song a rather long series of high - pitched “tsee” notes , variable . . .
low mossy forest and very humid elfin woodland , bamboo , and giant grasses at or near tree - line , and . . .
fruit and apparently some insects . contents of one stomach a large number of seeds . found singly , in pairs and in little groups up to four . . .
vulnerable . restricted - range species : present in central andean páramo eba . generally rare to uncommon ; localized . estimated global population not exceeding 5000 . . .
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“core tanagers” , with over 100 species , including a clade most members of which ( lophospingus , diuca , gubernatrix , paroaria ) were previously treated in emberizidae and one species ( pseudosaltator rufiventris ) previously treated in cardinalidae # r .
resurrected for single species for long treated in buthraupis but found by molecular study to be notably isolated , with sporathraupis as its closest apparent relative # r # r .
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lars petersson , dusan m . brinkhuizen , agustin carrasco , dubi shapiro , scott olmstead , nick athanas , william price , brayan coral jaramillo .
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howard and moore 4th edition ( incl . corrigenda vol . 1 - 2 ) :
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11 - 13 november 2015 . the antisana ecological reserve can easily be reached via a day trip from quito . given the frequent afternoon weather , it is best to arrive early in the morning to improve your chances of having an unobstructed view of the massive peak of antisana . antisana rises to a height of 18 , 891 ft ( 5 , 758 m ) and is absolutely [ . . . ]
16 – 21 september 2015 . we really ran short of time in southern colombia , unfortunately . we had hoped to visit pnn cueva de los guacharos , the mocoa area , the sibundoy area , and the junin area ( rna pangan and rio ñambi ) but we only had a few days left on our colombian visas so we decided to check out the mocoa and sibundoy areas as there [ . . . ]
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