''Pulanesaura'' 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 basal

sauropodiform Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lon ...

known from the

Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic� ...

(late

Hettangian The Hettangian is the earliest age and lowest stage of the Jurassic Period of the geologic timescale. It spans the time between 201.3 ± 0.2 Ma and 199.3 ± 0.3 Ma (million years ago). The Hettangian follows the Rhaetian (part of the Triass ...

Sinemurian In the geologic timescale, the Sinemurian is an age (geology), age and stage (stratigraphy), stage in the Early Jurassic, Early or Lower Jurassic epoch (geology), Epoch or series (stratigraphy), Series. It spans the time between 199.5 ±0.3 annu ...

) Upper

Elliot Formation The Elliot Formation is a geological formation and forms part of the Stormberg Group, the uppermost geological group that comprises the greater Karoo Supergroup. Outcrops of the Elliot Formation have been found in the northern Eastern Cape, so ...

of the Free State,

South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...

. It contains a single species, ''Pulanesaura eocollum'', known from partial remains of at least two subadult to adult individuals.

Discovery and naming

The remains of ''Pulanesaura'' were discovered in a small quarry in the farm Spion Kop 932 in the

Senekal Senekal is a town situated on the banks of the Sand River (Free State), Sand River in the eastern part of the Free State (South African province), Free State province of South Africa. It was named after Commandant FP Senekal, who led the encro ...

District of the Free State in 2004 by paleontologist Matthew Bonnan. The bones were excavated between 2004 and 2006, and studied by Blair McPhee as part of his dissertation since 2011. ''Pulanesaura'' was then described and named officially by Blair W. McPhee, Matthew F. Bonnan, Adam M. Yates, Johann Neveling and Jonah N. Choiniere in

2015 2015 was designated by the United Nations as: * International Year of Light * International Year of Soil __TOC__ Events January * January 1 – Lithuania officially adopts the euro as its currency, replacing the litas, and becomes ...

with the

type species In International_Code_of_Zoological_Nomenclature, 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 spe ...

''Pulanesaura eocollum''. The generic name is derived from the Sesoth word for "rain-maker/bringer", ''Pulane'', in reference to the heavy rain conditions under which the remains were collected, and the feminine form of the common dinosaur name suffix, ''saura'', meaning "lizard" in

Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...

. 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 ...

is derived from

Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...

''eo'', meaning "dawn", and Latin ''collum'', meaning "neck", in reference to ''Pulanesaura'' being a very basal

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 ...

not yet showing the most archetypal trait of more advanced sauropods - their very long necks. ''Pulanesaura'' was one of eighteen dinosaur taxa from 2015 to be described in open access or free-to-read journals. ''Pulanesaura'' is known from partial remains of at least two subadult to adult individuals. 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 ...

, BP/1/6982, represents the front dorsal

vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...

missing the tip of the neural spine. In addition the referred material consists of two isolated teeth, a middle

cervical vertebra 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 sauropsid s ...

, five back vertebral arches, a single right dorsal rib, three tail vertebrae, a left

clavicle The clavicle, collarbone, or keybone is a slender, S-shaped long bone approximately long that serves as a strut between the scapula, shoulder blade and the sternum (breastbone). There are two clavicles, one on each side of the body. The clavic ...

, a distal right

humerus The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...

, a left

ulna The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to the wrist. It is on the same side of the forearm as the little finger, running parallel to the Radius (bone), radius, the forearm's other long ...

, possibly the fourth right middle hand bone, three

ischia Ischia ( , , ) is a volcanic island in the Tyrrhenian Sea. It lies at the northern end of the Gulf of Naples, about from the city of Naples. It is the largest of the Phlegrean Islands. Although inhabited since the Bronze Age, as a Ancient G ...

, a left and a right shinbones, and two hindlimb first

claws A claw is a curved, pointed appendage found at the end of a toe or finger in most amniotes (mammals, reptiles, birds). Some invertebrates such as beetles and spiders have somewhat similar fine, hooked structures at the end of the leg or tars ...

. The remains are considered to be conspecific with the holotype due to their close association (in an area of three to three and a half meters) in fine and stable sandstone, their consistent

morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...

, and the fact the same elements from different individuals show no conflict in traits. The remains were collected on the farm Spion Kop 932, in a quarry located just over a kilometer East-North East another dinosaur rich quarry in a higher stratigraphic position within the probably

part of the upper

, that yielded the less advanced

sauropodomorph Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lo ...

s ''

Aardonyx celestae ''Aardonyx'' (Afrikaans ''aard'', "earth" + Greek , "nail, claw") is a genus of basal sauropodomorph dinosaur. It is known from the type species ''Aardonyx celestae'' found from the Early Jurassic Elliot Formation of South Africa. ''A. celestae' ...

'' and the much smaller '' Arcusaurus pereirabdalorum''.

Phylogeny

''Pulanesaura'' is a medium-sized transitional sauropodiform. A

phylogenetic analysis In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...

resolved its position as either one of the least derived

s or as the

sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...

to Sauropoda, depending on the

definition A definition is a statement of the meaning of a term (a word, phrase, or other set of symbols). Definitions can be classified into two large categories: intensional definitions (which try to give the sense of a term), and extensional definitio ...

for Sauropoda used (node or stem based). The following

cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...

is simplified after this analysis (members of bold taxa are not shown). The following cladogram shows the position of ''Pulanesaura'' within Massopoda, according to Oliver W. M. Rauhut and colleagues, 2020:

Paleoecology

''Pulanesauras posture and skeletal build indicate that the animal was a low browser, unlike the prosauropods it shared its habitat with. Studies by Blair McPhee et al. indicate that ''Pulanesaura'' is thought to have coexisted with other sauropodomorphs found in the same formation due to

niche partitioning In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for e ...

. Its flexible neck would have further allowed it to feed without moving its body very often and expending valuable energy; a trait that later sauropods would take to extreme lengths. Studies of the Upper Elliot Formation suggest that the environment was a predominantly arid floodplain where vegetation was concentrated most heavily around the river channels that flowed through the area, further allowing the coexistence of ''Pulanesaura'' with other sauropodomorphs such as ''Aardonyx'' and ''Arcusaurus''.

Discovery and naming

Phylogeny

Paleoecology

References

Further reading