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''Cutleria'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of basal
sphenacodontid Sphenacodontidae (Greek: "wedge point tooth family") is an extinct family (biology), family of small to large, advanced, carnivore, carnivorous, Late Pennsylvanian to Guadalupian, middle Permian pelycosaurs. The most recent one, ''Dimetrodon ang ...
s or derived stem- sphenacodontoid known from the Early Permian
period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
(
Sakmarian In the geologic timescale, the Sakmarian is an age or stage of the Permian. It is a subdivision of the Cisuralian Epoch or Series. The Sakmarian lasted between 293.52 and million years ago (Ma). It was preceded by the Asselian and followed by t ...
stage) of the
Colorado Colorado (, other variants) is a state in the Mountain states, Mountain West subregion of the Western United States. It encompasses most of the Southern Rocky Mountains, as well as the northeastern portion of the Colorado Plateau and the wes ...
, United States. It contains a single species, ''Cutleria wilmarthi''.


Discovery

''Cutleria'' is known only from the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...
specimen USNM 22099, a fractured but three-dimensionally preserved, a nearly complete
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
and articulated partial
postcrania Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...
l skeleton (including
vertebral column The vertebral column, also known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic of a vertebrate in which the notochord (a flexible rod of uniform composition) found in all chordate ...
, ribs and several girdle and limb elements). It was collected at locality 3, near Placerville of
San Miguel County, Colorado San Miguel County is a county located in the U.S. state of Colorado. As of the 2020 census, the population was 8,072. The county seat is Telluride. The county is named for the San Miguel River. History San Miguel County was given the Spanish ...
, from the
Cutler Formation The Cutler Formation or Cutler Group is a rock unit that is exposed across the U.S. states of Arizona, northwest New Mexico, southeast Utah and southwest Colorado. It was laid down in the Early Permian during the Wolfcampian epoch. Descri ...
, dating to the
Sakmarian In the geologic timescale, the Sakmarian is an age or stage of the Permian. It is a subdivision of the Cisuralian Epoch or Series. The Sakmarian lasted between 293.52 and million years ago (Ma). It was preceded by the Asselian and followed by t ...
stage of the Cisuralian
series Series may refer to: People with the name * Caroline Series (born 1951), English mathematician, daughter of George Series * George Series (1920–1995), English physicist Arts, entertainment, and media Music * Series, the ordered sets used in ...
. MCZ 2987, a tip of the
rostrum Rostrum may refer to: * Any kind of a platform for a speaker: **dais **pulpit * Rostrum (anatomy), a beak, or anatomical structure resembling a beak, as in the mouthparts of many sucking insects * Rostrum (ship), a form of bow on naval ships * Ros ...
and some teeth collected 2.5 km from the type locality (from localities 11-13), was originally referred to ''C. wilmarthi'' by Lewis and Vaughn (1965). A redescription of sphenacodonts by
Michel Laurin Michel Laurin is a Canadian-born French vertebrate paleontologist whose specialities include the emergence of a land-based lifestyle among vertebrates, the evolution of body size and the origin and phylogeny of lissamphibians. He has also made impo ...
(1993 and 1994), revealed that it can't be assigned to any named sphenacodont genus. Although its teeth also bear cutting edges without serrations, they are more
bulbous In botany, a bulb is structurally a short stem with fleshy leaves or leaf basesBell, A.D. 1997. ''Plant form: an illustrated guide to flowering plant morphology''. Oxford University Press, Oxford, U.K. that function as food storage organs durin ...
, not as strongly compressed
lateral Lateral is a geometric term of location which may refer to: Healthcare *Lateral (anatomy), an anatomical direction * Lateral cricoarytenoid muscle * Lateral release (surgery), a surgical procedure on the side of a kneecap Phonetics *Lateral co ...
ly and not curved distally. A new genus wasn't erected for MCZ 2987, as it is very fragmentary and lacks sufficient diagnostic features.


Description

''Cutleria'' was assigned to the
Haptodontinae ''Haptodus'' is an extinct genus of basal sphenacodont, member of the clade that includes therapsids and hence, mammals. It was at least in length. It lived in present-day France during the Early Permian. It was a medium-sized predator, feedi ...
by Lewis and Vaughn (1965) on the basis of similarities to ''
Haptodus baylei ''Haptodus'' is an extinct genus of basal sphenacodont, member of the clade that includes therapsids and hence, mammals. It was at least in length. It lived in present-day France during the Early Permian. It was a medium-sized predator, feedi ...
'' and '' H. longicaudata'' (now '' Palaeohatteria''). Currie (1977) described the second haptodontine from North America, ''
Haptodus garnettensis ''"Haptodus" garnettensis'' is an extinct species of basal sphenacodont from the Late Carboniferous ( Pennsylvanian) of Kansas, USA. Taxonomy The holotype of ''"Haptodus" garnettensis'' is RM 14156, a partially articulated skeleton ( ...
''. In 1979, he synonymized all European haptodontines, as well as ''C. wilmarthi'', with ''H. baylei''. Later, Laurin (1993 and 1994) considered Haptodontinae to represent a
polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of conver ...
grade of basal sphenacodonts, and revalidated ''Cutleria'' among other genera and species. His revision has been accepted since. Only three
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analyses included ''Cutleria''. Laurin (1994) and Fröbisch ''et al.'' (2011) recovered ''Cutleria'' as the sister taxon of Sphenacodontoidea, while the significantly larger analysis of Benson (in press) recovered it as the basalmost
sphenacodontid Sphenacodontidae (Greek: "wedge point tooth family") is an extinct family (biology), family of small to large, advanced, carnivore, carnivorous, Late Pennsylvanian to Guadalupian, middle Permian pelycosaurs. The most recent one, ''Dimetrodon ang ...
. Benson (in press) listed three synapomorphies between ''Cutleria'' and more derived sphenacodontids: their frontal is more than 2.5 times the length of the parietal (as it was noted by Laurin in 1994), a strongly posteroventrally inclined temporal roof (absent in ''
Secodontosaurus ''Secodontosaurus'' (meaning "cutting-tooth lizard") is an extinct genus of "pelycosaur" synapsids that lived from between about 285 to 272 million years ago during the Early Permian. Like the well known ''Dimetrodon'', ''Secodontosaurus'' is a c ...
''), and a lateral centrale that covers the proximal ends of the first-third distal carpals. However, only one additional step was required to place ''Cutleria'' outside Sphenacodontidae, as the sister taxon of Sphenacodontoidea (which is the node "Therapsida + Sphenacodontidae"). The
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
below shows ''Cutleria'' phylogenetic position among other synapsids following Benson (2012).


Etymology

''Cutleria'' was first described and named by George Edward Lewis and Peter Paul Vaughn in 1965 and the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specime ...
is ''Cutleria wilmarthi''. The generic name is derived from the name of the
Cutler Formation The Cutler Formation or Cutler Group is a rock unit that is exposed across the U.S. states of Arizona, northwest New Mexico, southeast Utah and southwest Colorado. It was laid down in the Early Permian during the Wolfcampian epoch. Descri ...
in which the holotype was found. The specific name honors Wilmarth for finding the holotype.


References

{{Taxonbar, from=Q5196889 Sphenacodontidae Fossil taxa described in 1965 Cisuralian synapsids of North America Prehistoric synapsid genera Cisuralian genus first appearances