Katepensaurus
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''Katepensaurus'' ("hole lizard") is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
rebbachisaurid Rebbachisauridae is a Family (biology), family of sauropod dinosaurs known from fragmentary fossil remains from the Cretaceous of South America, Africa, North America, Europe and possibly Central Asia. Taxonomy In 1990 sauropod specialist Jack M ...
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
known from the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...
of south-central
Chubut Province Chubut ( from Tehuelche language, Tehuelche 'transparent'; ) is a provinces of Argentina, province in southern Argentina, situated between the 42nd parallel south (the border with Río Negro Province), the 46th parallel south (bordering Santa ...
of central
Patagonia Patagonia () is a geographical region that includes parts of Argentina and Chile at the southern end of South America. The region includes the southern section of the Andes mountain chain with lakes, fjords, temperate rainforests, and glaciers ...
,
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
. It contains a single species, ''Katepensaurus goicoecheai''.


Fossil record

The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
and only known specimen of ''Katepensaurus goicoecheai'' is UNPSJB-PV 1007, a partial skeleton from the Lower Member of the
Bajo Barreal Formation The Bajo Barreal Formation is a geological Formation (geology), formation in the Golfo San Jorge Basin of Chubut Province, Chubut and Santa Cruz Province, Argentina, Santa Cruz, Argentina whose strata date back to the Cenomanian, Middle Cenomanian ...
of
Chubut Province Chubut ( from Tehuelche language, Tehuelche 'transparent'; ) is a provinces of Argentina, province in southern Argentina, situated between the 42nd parallel south (the border with Río Negro Province), the 46th parallel south (bordering Santa ...
,
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
. The specimen preserves one skull bone (a frontal), three
cervical vertebrae In tetrapods, cervical vertebrae (: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In saurop ...
, six
dorsal vertebrae In vertebrates, thoracic vertebrae compose the middle segment of the vertebral column, between the cervical vertebrae and the lumbar vertebrae. In humans, there are twelve thoracic vertebra (anatomy), vertebrae of intermediate size between the ce ...
, five
caudal vertebrae Caudal vertebrae are the vertebrae of the tail in many vertebrates. In birds, the last few caudal vertebrae fuse into the pygostyle, and in apes, including humans, the caudal vertebrae are fused into the coccyx. In many reptiles, some of the caud ...
, and several difficult-to-identify fragments, which probably include portions of additional vertebrae as well as possible fragments of some limb bones (a
metacarpal In human anatomy, the metacarpal bones or metacarpus, also known as the "palm bones", are the appendicular bones that form the intermediate part of the hand between the phalanges (fingers) and the carpal bones ( wrist bones), which articulate ...
or
metatarsal The metatarsal bones or metatarsus (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges ( toes). Lacking individual names, the metatarsal bones are ...
and an
astragalus Astragalus may refer to: * ''Astragalus'' (plant), a large genus of herbs and small shrubs *Astragalus (bone) The talus (; Latin for ankle or ankle bone; : tali), talus bone, astragalus (), or ankle bone is one of the group of foot bones known ...
). It represents an individual that was at least seven years old and had not finished growing when it died. The Lower Member of the Bajo Barreal Formation is considered to date to the
Cenomanian The Cenomanian is, in the International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...
or
Turonian The Turonian is, in the International Commission on Stratigraphy, ICS' geologic timescale, the second age (geology), age in the Late Cretaceous epoch (geology), Epoch, or a stage (stratigraphy), stage in the Upper Cretaceous series (stratigraphy), ...
age of the early
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...
. Additional rebbachisaurid specimens from the Bajo Barreal Formation have been described, including a cervical vertebra (UNPSJB-PV 1005), an isolated caudal vertebra (UNPSJB-PV 580), and a partial tail consisting of four incomplete caudal vertebrae and two
chevrons Chevron (often relating to V-shaped patterns) may refer to: Science and technology * Chevron (aerospace), sawtooth patterns on some jet engines * Chevron (anatomy), a bone * '' Eulithis testata'', a moth * Chevron (geology), a fold in rock la ...
(UNPSJB-PV 1004). It is possible that these specimens also belong to ''Katepensaurus'', although the available evidence is insufficient to determine their affinities more precisely than Rebbachisauridae. Remains of a rebbachisaurid taxon distinct from ''Katepensaurus'' have been found in the Bajo Barreal Formation, and it is possible that these specimens belong to this other unnamed species.


Discovery and naming

The only known specimen of ''Katepensaurus goicoecheai'' was collected from the "2005 Quarry" at Estancia Laguna Palacios, a ranch owner at which the National University of the Patagonia San Juan Bosco regularly conducted field work. In
2013 2013 was the first year since 1987 to contain four unique digits (a span of 26 years). 2013 was designated as: *International Year of Water Cooperation *International Year of Quinoa Events January * January 5 – 2013 Craig, Alask ...
, Lucio M. Ibiricu, Gabriel A. Casal, Rubén Dario Martínez, Matthew C. Lamanna, Marcelo Luna and Leonardo Salgado described this specimen as representing a new genus and species, which they named ''Katepensaurus goicoecheai''. The generic name is derived from Tehuelche ''katepenk'', "hole", referring to a distinctive opening in the transverse processes of the dorsal vertebrae. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
honours Alejandro Goicoechea, the owner of the Estancia Laguna Palacios. At the time the species was named, only three cervical vertebrae, three dorsal vertebrae, and two caudal vertebrae had been excavated and identified, but subsequent excavations conducted at the quarry uncovered additional bones from the same skeleton.


Description

''Katepensaurus'' is distinguished from other
rebbachisaurids Rebbachisauridae is a family of sauropod dinosaurs known from fragmentary fossil remains from the Cretaceous of South America, Africa, North America, Europe and possibly Central Asia. Taxonomy In 1990 sauropod specialist Jack McIntosh included t ...
primarily by various details of its dorsal vertebrae. As in other sauropods, the vertebrae of ''Katepensaurus'' were highly
pneumatized Skeletal pneumaticity is the presence of air spaces within bones. It is generally produced during development by excavation of bone by pneumatic diverticula (air sacs) from an air-filled space, such as the lungs or nasal cavity. Pneumatization is h ...
, with openings in the bone, such as the lateral pneumatic fossae, that would have been filled by air sacs in life. In ''Katepensaurus'', the lateral pneumatic fossae of the dorsal vertebrae were divided in two by a ridge of bone, which is an uncommon trait among sauropods not known in other rebbachisaurids but present in the more distantly related ''
Supersaurus ''Supersaurus'' (meaning "super lizard") is a genus of diplodocid sauropod dinosaur that lived in North America during the Late Jurassic period. The type species, ''S. vivianae'', was first discovered by Vivian Jones of Delta, Colorado, in the ...
''. ''Katepensaurus'' is unique among sauropods in exhibiting an additional set of pneumatic openings, termed laterodiapophyseal fenestrae, through which air sacs would have filled the transverse processes of the dorsal vertebrae. It is these openings that give ''Katepensaurus'' its name. The arrangement of the vertebral laminae, a complex arrangement of bony ridges that are characteristic of sauropod vertebrae, is generally similar between ''Katepensaurus'' and other rebbachisaurids. However, ''Katepensaurus'' shows a unique pair of laminae, termed the dorsal parapophyseal centrodiapophyseal fossa lamina and the ventral parapophyseal centrodiapophyseal fossa lamina, that have not been seen in any other rebbachisaurid. Other unique traits that have been documented in the dorsal vertebrae of ''Katepensaurus'' include vertical ridges on the junction between the neural arch and centrum, well-defined, rounded fossae on the lateral aspect of the postzygapophyses, and a teardrop-shaped profile of the posterior articular surface of the centrum.


Classification

''Katepensaurus'' exhibits a combination of traits characteristic of nigersaurines/ rebbachisaurines and traits characteristic of limaysaurines, and its phylogenetic affinities are not certain. An analysis by Ibiricu and colleagues in 2015 recovered ''Katepensaurus'' as a limaysaurine; in contrast, analyses by Fanti and colleagues in 2015 and Canudo and colleagues in 2018 recovered ''Katepensaurus'' as the basalmost rebbachisaurine. Analyses by Mannion and colleagues in 2019 could not resolve the affinities of ''Katepensaurus'' precisely, and found differing results depending on their analysis protocol. In one of their analyses, ''Katepensaurus'' was recovered as a limaysaurine, but in the other, it was recovered outside Khebbashia, as a basal rebbachisaurid.


Footnotes


References

{{Taxonbar, from=Q15179026 Rebbachisauridae Dinosaur genera Cenomanian dinosaurs Bajo Barreal Formation Fossil taxa described in 2013 Taxa named by Matt Lamanna Dinosaurs of Argentina