Machaeroprosopus

{{Short description|Extinct genus of reptiles}} {{Automatic taxobox | fossil_range = Late Triassic, {{fossil_range|228|209}} | image = Phytosaurs.jpg | image_caption = Skulls in New Mexico | taxon = Machaeroprosopus | authority = Mehl ''et al.'', 1916 <ref name=Mehletal1916/> | type_species = {{extinct}}'''''Machaeroprosopus buceros''''' | type_species_authority = Cope, 1881 | subdivision_ranks = Species | subdivision = *{{extinct}}'''''M. andersoni''''' Mehl, 1922 *{{extinct}}'''''M. buceros''''' (Cope, 1881) *{{extinct}}'''''M. jablonskiae''''' (Parker & Irmis, 2006) <ref name=ParkerIrmis06/> *{{extinct}}'''''M. lottorum''''' Hungerbühler ''et al.'', 2013 <ref name=M.lottorum/> *{{extinct}}'''''M. mccauleyi''''' (Ballew, 1989) <ref name=Ballew89/> *{{extinct}}'''''M. pristinus''''' (Mehl, 1928) *{{extinct}}'''''M. validus''''' Mehl in Mehl ''et al.'', 1916 <ref name=Mehletal1916/> | synonyms = * ''Arribasuchus'' Long & Murry, 1995 <ref name="LongMurry95"/> * ''Machaeroprosopus tenuis'' Camp, 1930 * ''Pseudopalatus'' Mehl, 1928 }}

'''''Machaeroprosopus''''' (from {{langx|el|μαχαίρα}} {{Transliteration|el|machaíra}}, 'large knife' and {{langx|el|πρόσωπος}} {{Transliteration|el|prósōpos}}, 'bordering on')<ref>{{cite book |last1=Colbert |first1=Edwin H. (Edwin Harris) |last2=Knight |first2=Charles Robert |title=The dinosaur book : the ruling reptiles and their relatives |date=1951 |publisher=McGraw-Hill |location=New York |page=150 |url=https://archive.org/details/bookruli00colb}}</ref> is an extinct genus of mystriosuchin leptosuchomorph phytosaur from the Late Triassic of the southwestern United States. ''M. validus'', once thought to be the type species of ''Machaeroprosopus'', was named in 1916 on the basis of three complete skulls from Chinle Formation, Arizona. The skulls have been lost since the 1950s, and a line drawing in the original 1916 description is the only visual record of the specimen.<ref name="chin">{{cite web |url=http://chinleana.fieldofscience.com/2009/02/tragic-tale-of-machaeroprosopus-and.html |last=Parker |first=Bill |title=The Tragic Tale of ''Machaeroprosopus'' and ''Acompsosaurus'' |date=7 February 2007 |work=Chinleana |publisher=Field of Science |access-date=11 May 2012 |archive-url=https://web.archive.org/web/20120415145412/http://chinleana.fieldofscience.com/2009/02/tragic-tale-of-machaeroprosopus-and.html |archive-date=15 April 2012 |url-status=dead }}</ref> Another species, ''M. andersoni'', was named in 1922 from New Mexico, and the species ''M. adamanensis'', ''M. gregorii'', ''M. lithodendrorum'', ''M. tenuis'', and ''M. zunii'' were named in 1930. Most species have been reassigned to the genera ''Smilosuchus'',<ref name=Stocker10>{{cite journal|first=Michelle R. |last=Stocker |date=2010 |title=A new taxon of phytosaur (Archosauria: Pseudosuchia) from the Late Triassic (Norian) Sonsela Member (Chinle Formation) in Arizona, and a critical reevaluation of ''Leptosuchus'' Case, 1922 |journal=Palaeontology |volume=53 |issue=5 |pages=997–1022 |doi=10.1111/j.1475-4983.2010.00983.x |s2cid=83536253 |doi-access=free }}</ref> ''Rutiodon'', or ''Phytosaurus''.<ref name=GJT62>{{cite journal |last=Gregory |first=J.T. |date=1962 |title=The relationships of the American phytosaur ''Rutiodon'' |journal=American Museum Novitates |issue=2095 |pages=1–22 |url=http://digitallibrary.amnh.org/dspace/bitstream/handle/2246/3397/v2/dspace/ingest/pdfSource/nov/N2095.pdf?sequence=1}}</ref> Until recently, ''M. validus'' was considered to be the only species that has not been reassigned. Thus, ''Machaeroprosopus'' was considered to be a ''nomen dubium'' or "doubtful name" because of the lack of diagnostic specimens that can support its distinction from other phytosaur genera. However, a taxonomic revision of ''Machaeroprosopus'', conducted by Parker ''et al.'' in 2013, revealed that UW 3807, the holotype of ''M. validus'', is not the holotype of ''Machaeroprosopus'', while the species ''Machaeroprosopus buceros'', ''Machaeroprosopus'' being a replacement name, with a fixed type species, for ''Metarhinus'', is the ''combinatio nova'' of the type species of the genu: ''Belodon buceros''. Therefore, the name ''Pseudopalatus'' must be considered a junior synonym of ''Machaeroprosopus'', and all species of the former must be reassigned to the latter.<ref name=Parkeretal13>{{Cite journal | last1 = Parker | first1 = W. G. | last2 = Hungerbühler | first2 = A. | last3 = Martz | first3 = J. W. | title = The taxonomic status of the phytosaurs (Archosauriformes) ''Machaeroprosopus'' and ''Pseudopalatus'' from the Late Triassic of the western United States | doi = 10.1017/S1755691013000339 | journal = Earth and Environmental Science Transactions of the Royal Society of Edinburgh | volume = 103 | issue = 3–4 | pages = 265–268 | year = 2013 | s2cid = 84739408 }}</ref> This revised taxonomy was already accepted in several studies, including Stocker and Butler (2013).<ref name="StockerButler13">{{Cite journal | last1 = Stocker | first1 = M. R. | last2 = Butler | first2 = R. J. | title = Phytosauria | doi = 10.1144/SP379.5 | journal = Geological Society, London, Special Publications | volume = 379 | pages = 91–117 | year = 2013 | issue = 1 | bibcode = 2013GSLSP.379...91S | s2cid = 219192243 }}</ref> Stocker and Butler (2013) also treated ''M. andersoni'' as a valid species,<ref name="StockerButler13" /> and not a junior synonym of ''Machaeroprosopus buceros'' as was previously suggested by Long and Murry (1995).<ref name="LongMurry95">{{cite journal | last1=Long | first1= R. A. |last2= Murry | first2= P. A. | date = 1995 | title = Late Triassic (Carnian and Norian) tetrapods from the southwestern United States | journal = New Mexico Museum of Natural History and Science Bulletin | volume = 4 | pages = 1–254 }}</ref>

==Species==

===''M. andersoni''=== thumb|left|Skull of ''M. andersoni'' ''M. andersoni'' was first described and named by Maurice G. Mehl in 1922, on the basis of the holotype FMNH UC 396, partial skull.<ref name=StockerButler13/> It was probably collected from the Bull Canyon Formation of the Chinle Group or Dockum Group, probably at the Bull Canyon, in the Guadalupe County of New Mexico.<ref name=LongMurry95/> This taxon was considered to be a junior synonym of ''M. buceros'' by Long and Murry (1995) and later authors,<ref name=LongMurry95/><ref name=Irmis05/><ref name=ParkerIrmis06/> although Stocker and Butler (2013) treated ''M. andersoni'' as a valid species.<ref name=StockerButler13/>

The holotype is the only known specimen of this species, although there are other specimens from the Bull Canyon Formation that were referred to ''Arribasuchus buceros'' by Long and Murry (1995),<ref name=LongMurry95/> but not by Stocker and Butler (2013).<ref name=StockerButler13/>

===''M. buceros''=== thumb|left|''M. buceros'' skull ''M. buceros'' was first described and named by Edward Drinker Cope in 1881 as ''Belodon buceros'', on the basis of the holotype AMNH 2318, partial skull. It was collected from the Norian-aged Petrified Forest Member of the Chinle Formation, at the Arroyo Seco drainage, Orphan Mesa in the Rio Arriba County of New Mexico.<ref name=Lucasetal02>{{cite journal |last1 = Lucas | first1 = S. G. | last2 = Heckert | first2 = A. B. | last3 = Zeigler | first3 = K. E. | last4 = Hunt | first4 = A. P. |year=2002 |title=The type locality of ''Belodon buceros'' Cope, 1881, a phytosaur (Archosauria: Parasuchidae) from the Upper Triassic of north-central New Mexico | editor1-first = A. B. | editor1-last = Heckert | editor2-first = S. G. | editor2-last = Lucas | journal = Upper Triassic Stratigraphy and Paleontology, New Mexico Museum of Natural History and Science Bulletin |volume=21 |pages=189–192}}</ref> Later, the skull was occasionally referred to ''Phytosaurus buceros''. Jaekel (1910) erected a new genus for the species, creating ''Metarhinus buceros'', however this generic name was preoccupied by ''Metarhinus'' Osborn, 1908, a brontotheriid mammal. Mehl (1915) referred this species to ''Lophoprosopus'', as ''Lophoprosopus buceros'', but as the type species of this genus is considered to be synonymous with ''Nicrosaurus kapffi'', Mehl (1916) erected a new genus for ''B. buceros'', creating ''Machaeroprosopus buceros'', but also explicitly indicated ''M. validus'' as the type. Subsequent studies accordingly considered ''M. validus'' to be the type species of ''Machaeroprosopus''. Later, the skull was referred to as ''Rutiodon buceros'', ''Machaeroprosopus buceros'' or ''Pseudopalatus buceros''. Long and Murry (1995) erected a new genus for the species, not knowing that ''M. validus'' was not in fact the type species of ''Machaeroprosopus'', creating ''Arribasuchus buceros''.<ref name=LongMurry95/> Subsequent studies, including Hungerbühler (2002), Lucas ''et al.'' (2002),<ref name=Lucasetal02/> Zeigler ''et al.'' (2002), Irmis (2005)<ref name=Irmis05>{{cite journal | first = R. B. | last = Irmis | date = 2005 | title = The vertebrate fauna of the Upper Triassic Chinle Formation in northern Arizona | editor1-first = S. J. | editor1-last = Nesbitt | editor2-first = W. G. | editor2-last = Parker | editor3-first = R. B. | editor3-last = Irmis | journal = Guidebook to the Triassic Formations of the Colorado Plateau in Northern Arizona: Geology, Paleontology, and History. Mesa Southwest Museum Bulletin | volume = 9 | pages = 63–88 }}</ref> and Parker and Irmis (2006),<ref name=ParkerIrmis06>{{cite journal | last1 = Parker | first1 = W. G. | first2 = R. B. | last2 = Irmis | date = 2006 | url = http://socrates.berkeley.edu/~irmisr/pseudjab.pdf | title = A new species of the Late Triassic phytosaur Pseudopalatus (Archosauria: Pseudosuchia) from Petrified Forest National Park, Arizona | journal = Museum of Northern Arizona Bulletin | volume = 62 | pages = 126–143 }}</ref> referred the species back to ''Pseudopalatus buceros'', and considered ''Arribasuchus'' to be a junior synonym of ''Pseudopalatus''. A taxonomic revision of ''Machaeroprosopus'', conducted by Parker ''et al.'' in 2013, revealed that ''Machaeroprosopus buceros'' is the ''combinatio nova'' of the type species of the genus because Article 67-8 ICZN rules that with replacement names the original type species (i.c. Cope's ''Beladon buceros'') is maintained. Therefore, the name ''Pseudopalatus'' was considered a junior synonym of ''Machaeroprosopus'', and all species of the former were reassigned to the later, including ''P. buceros''.<ref name=Parkeretal13/><ref name=StockerButler13/> Following the revision, ''M. andersoni'' was treated as a valid species by Stocker and Butler (2013),<ref name=StockerButler13/> and not a junior synonym of ''Machaeroprosopus buceros'' as was previously suggested by Long and Murry (1995). Other specimens were referred to ''P. buceros'' by Long and Murry (1995),<ref name=LongMurry95/> although all specimens from Arizona and Texas was later reassigned to other species (namely ''M. lottorum'', ''M. mccauleyi'' and ''M. validus''), and as ''M. andersoni'' from the Dockum Group was re-validated, ''M. buceros'' is currently known only from Chinle Formation of New Mexico.<ref name=StockerButler13/><ref name=M.lottorum/>

===''M. jablonskiae''=== ''M. jablonskiae'' was first described and named by William G. Parker and Randall B. Irmis in 2006, on the basis of the holotype PEFO 31207, posterior skull roof and braincase missing the rostrum and palate. This specimen was initially referred to ''Pseudopalatus cf. mccauleyi'' by Parker and Irmis (2004) based on the morphology of the squamosals and the opisthotic, then to ''Pseudopalatus sp.'' by Parker and Irmis (2005) and finally to ''Pseudopalatus jablonskiae'' by Parker and Irmis (2006). It was collected in September 2002 from the lower Jim Camp Wash beds, Sonsela Member of the Chinle Formation, at locality PFV 295, near Mountain Lion Mesa in Petrified Forest National Park, Arizona. The specific name honors Pat Jablonsky, who discovered the holotype and only known specimen.<ref name=ParkerIrmis06/> Following the taxonomic revision of ''Machaeroprosopus'' by Parker ''et al.'' (2013), all species of ''Pseudopalatus'', including ''P. jablonskiae'', were reassigned to ''Machaeroprosopus''.<ref name=Parkeretal13/> This was already accepted by Stocker and Butler (2013).<ref name=StockerButler13/>

Although ''M. jablonskiae'' is known from an incomplete specimen, it can be diagnosed by at least one autapomorphy and a unique suite of traits, and it includes a well-preserved braincase, which is rarely preserved or described in detail for most phytosaur specimens. A phylogenetic analysis of mystriosuchin phytosaurs performed by Parker and Irmis (2006) found the species to be the most basal species of ''"Pseudopalatus"''.<ref name=ParkerIrmis06/>

===''M. lottorum''=== thumb|''M. lottorum'' skull ''M. lottorum'' was first described and named by Axel Hungerbühler, Bill Mueller, Sankar Chatterjee and Douglas P. Cunningham in 2013. The specific name honors John Lott and Patricia Lott Kirkpatrick, for their support during the work at the TTU VPL 3870. It is known from two complete skulls, the holotype TTU-P10076 and the paratype TTU-P10077 housed at Texas Tech University. The skulls were collected at Patricia Site (TTU Vertebrate Paleontology Locality 3870), 13 km South of Post, Garza County of west Texas, from the upper unit of the Norian Cooper Canyon Formation, Dockum Group. Other vertebrates known from this site include TTU-P10074, a partial skull referred to ''Machaeroprosopus'' sp., a phytosaur postcranial skeleton, fish, a temnospondyl amphibian, ''Typothorax'', ''Postosuchus'', ''Shuvosaurus'' and a theropod dinosaur.<ref name=M.lottorum>{{Cite journal | last1 = Hungerbühler | first1 = A. | last2 = Mueller | first2 = B. | last3 = Chatterjee | first3 = S. | last4 = Cunningham | first4 = D. P. | title = Cranial anatomy of the Late Triassic phytosaur Machaeroprosopus, with the description of a new species from West Texas | doi = 10.1017/S1755691013000364 | journal = Earth and Environmental Science Transactions of the Royal Society of Edinburgh | volume = 103 | issue = 3–4 | pages = 269 | year = 2013 | s2cid = 129442164 }}</ref>

A phylogenetic analysis of mystriosuchin phytosaurs performed by Hungerbühler ''et al.'' (2013) found the species to be a derived ''Machaeroprosopus'' species, most closely related to the type species of ''Redondasaurus'', ''"R." gregorii''. This clade in addition to ''Machaeroprosopus'' sp. (TTU-P10074) was recovered as the sister taxon of the clade formed by ''M. pristinus'' and ''M. buceros''. ''M. jablonskiae'', ''M. mccauleyi'' and ''"Redondasaurus" bermani'' were found to be basal species of ''Machaeroprosopus''.<ref name=M.lottorum/>

===''M. mccauleyi''=== thumb|''M. mccauleyi'' skull ''M. mccauleyi'' was first described and named by Karen A. Ballew in 1989 as a species of ''Pseudopalatus'', on the basis of the holotype UCMP 126999, an incomplete skull, lacking the anterior half of the rostrum, and probably associated lower jaws. The specific name honors John D. McCauley and Mrs. Molly McCauley McLean from Winslow, Arizona, the owners of the land at "Billings Gap" from which the holotype was found.<ref name=Ballew89>{{cite book |last=Ballew |first=K. L. |date=1989 |chapter=A phylogenetic analysis of Phytosauria from the Late Triassic of the western United States |editor1-last = Lucas | editor1-first = S. G. | editor2-last = Hunt | editor2-first = A. P. | title = Dawn of the age of dinosaurs in the American Southwest | publisher = New Mexico Museum of Natural History | location = Albuquerque |pages=309–339 |chapter-url=http://nmnaturalhistory.org/assets/files/Bulletins/DawnAgeDinos/dawn_18_ballew.pdf}}</ref> The specimen was originally informally designated as ''Pseudopalatus'' "bilingsensis" by Ballew (1986).<ref name=Stocker10/> It was collected at Dry Creek Tank SE (also known as UCMP V82040, UCMP 7043 and PFV 55), Apache County of Arizona, from the Norian-aged Upper Petrified Forest Member / Formation according to most authors,<ref name=Stocker10/><ref name=StockerButler13/><ref name=LongMurry95/> or possibly Sonsela Member of the Chinle Formation according to Parker & Irmis (2005). Ballew (1989) also referred to this species USNM 15839, another incomplete skull lacking the anterior half of the rostrum from Arizona.<ref name=Ballew89/> Long and Murry (1995) restricted this species to its holotype, although recent studies suggest that USNM 15839 is referable to it.<ref name="HCO’k13">{{Cite journal | last1 = Holloway | first1 = W. L. | last2 = Claeson | first2 = K. M. | last3 = O’Keefe | first3 = F. R. | doi = 10.1080/02724634.2013.747532 | title = A virtual phytosaur endocast and its implications for sensory system evolution in archosaurs | journal = Journal of Vertebrate Paleontology | volume = 33 | issue = 4 | pages = 848 | year = 2013 | s2cid = 30650543 }}</ref> Long and Murry (1995) also considered ''M. mccauleyi'' to be a species of their ''Arribasuchus'', probably synonymous with ''A. buceros'',<ref name=LongMurry95/> although most subsequent studies, like Hungerbühler (2002), Parker and Irmis (2006),<ref name=ParkerIrmis06/> Stocker (2010)<ref name=Stocker10/> and Stocker and Butler (2013)<ref name=StockerButler13/> treated ''M. mccauleyi'' as a valid species. Other specimens that are currently referred to ''M. mccauleyi'' include PEFO 31219,<ref name=Stocker10/> complete skull, lower jaws and articulated postcranial skeleton, from Petroglyph phytosaur site (also known as PFV 42) collected by UCMP in 1985, and possibly UCMP 27149, a large skull, from Cowboy (UCMP A257), both from the Petrified Forest Member, Arizona. Both specimen were originally referred to ''A. buceros'' by Long and Murry (1995).<ref name=LongMurry95/>

===''M. pristinus''=== thumb|''M. pristinus'' skull ''M. pristinus'' was first described and named by Maurice G. Mehl in 1928 as the type species of ''Pseudopalatus'', ''Pseudopalatus pristinus'', on the basis of the holotype MU 525, nearly complete skull. It was collected, from the Norian-aged Upper Petrified Forest Member of the Chinle Formation, near Adamana, Arizona. Additional skulls and postcranial material, some articulated skeletons, from the same member were referred to this species by Colbert (1946) and Long and Murry (1995) from Arizona, and by Lawler (1979), Ballew (1986) and Long and Murry (1995) from New Mexico.<ref name="LongMurry95" /> Charles Camp (1930) described and named ''Machaeroprosopus tenuis'' on the basis of UCMP 27018, a nearly complete skull, lower jaws, and some complete postcranial material, from the Billings Gap locality (UCMP 7043, Upper Petrified Forest Member), Apache County of Arizona. This specimen was occasionally referred to as ''Rutiodon tenuis'', although since Long and Murry (1995) it is considered to be a junior synonym of ''M. pristinus''.<ref name="LongMurry95" /> Long and Murry (1995) also suggested that ''Redondasaurus gregorii'' from the Redonda Formation of New Mexico is a junior synonym of ''M. pristinus'',<ref name="LongMurry95" /> although this was not accepted by subsequent authors.<ref name="ParkerIrmis06" /> Other, then unnumbered, specimens from the upper Church Rock Member (Chinle Formation, Utah), Bull Canyon Formation and Travesser Formation (New Mexico) and Cooper Canyon Formation (Texas), were referred to ''M. pristinus'' by Long and Murry (1995),<ref name="LongMurry95" /> although recent studies suggest that ''M. pristinus'' is currently known only from the Upper Petrified Forest Member of Arizona and New Mexico.<ref name="StockerButler13" />

===''M. validus''=== thumb|left|''M. validus'' restoration ''M. validus'' was first described and named by Maurice G. Mehl in Mehl ''et al.'', in 1916 on the basis of the holotype UW 3807, a complete skull, and the two paratypes UW 3808 and UW 3809, partial skulls,<ref name="chin" /> from Norian-aged Upper Petrified Forest Member of the Chinle Formation, Coconino County of Arizona.<ref name="LongMurry95" /> The skulls have been lost since the 1950s, and a line drawing in the original 1916 description is the only visual record of the specimen. ''M. validus'', once thought to be the type species of ''Machaeroprosopus'', was suggested by some authors to represent the same species as ''Pseudopalatus pristinus''. If this is the case, the name ''Machaeroprosopus'' would have precedence over ''Pseudopalatus'' because ''Pseudopalatus'' was named in 1928, twelve years after ''Machaeroprosopus'' was named.<ref name="chin"/> Long and Murry (1995) erected a new genus for ''Belodon buceros'', not knowing that it was in fact the type species of ''Machaeroprosopus'' (and not ''M. validus''), creating ''Arribasuchus buceros''. They also suggested that ''M. validus'' might be a junior synonym of ''A. buceros''.<ref name="LongMurry95" /> thumb|Line drawing of the skull of ''M. validus'' in Mehl ''et al.'' (1916)<ref name=Mehletal1916>{{cite journal | last1 = Mehl | first1 = Maurice | first2 = W. C. | last2 = Toepelmann | first3 = G. M. | last3 = Schwartz | date = 1916 | title = New or little known reptiles from the Trias of Arizona and New Mexico with notes from the fossil bearing horizons near Wingate, New Mexico | journal = University of Oklahoma Bulletin | volume = 103 | pages =1–44 }}</ref> Until recently, ''M. validus'' was considered to be the only species of ''Machaeroprosopus'' that has not been reassigned. Thus, ''Machaeroprosopus'' was considered to be a ''nomen dubium'' or "doubtful name" because of the lack of diagnostic specimens that can support its distinction from other phytosaur genera. However, a taxonomic revision of ''Machaeroprosopus'', conducted by Parker ''et al.'' (in press), revealed that UW 3807 is not the holotype of ''Machaeroprosopus'', while the species ''Machaeroprosopus buceros'' is in fact the correct new combination for the type species of the genus.

==References== {{Reflist}}

Category:Phytosauria Category:Prehistoric reptile genera Category:Norian genera Category:Late Triassic reptiles of North America Category:Triassic Arizona Category:Paleontology in Arizona Category:Triassic geology of New Mexico Category:Paleontology in New Mexico Category:Triassic geology of Texas Category:Paleontology in Texas Category:Fossil taxa described in 1916 Category:Taxa named by Maurice Mehl