Geikia Temporal range: | |
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G. locusticeps skull, showing large orbits and shortened face | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Suborder: | † Anomodontia |
Clade: | † Dicynodontia |
Family: | † Geikiidae |
Genus: | † Geikia Newton, 1893 |
Type species | |
†G. elginensis Newton, 1893 | |
Species | |
Geikia is an extinct genus of dicynodont therapsids from the late Permian. The abundance and diversity of dicynodonts during this period, combined with incomplete or inadequately prepared specimens, have led to challenges in determining relationships within this taxon. [1] Only two species, Geikia locusticeps and Geikia elginensis have been assigned to this genus. [2] While this is the currently accepted classification, fossil record limitations have led to repeated debate on the genus assignments of these species. [2] [3]
This genus was established in 1893 following Edwin Tulley Newton’s discovery of a new dicynodont fossil, now known as G. elginensis, one of the Elgin Reptiles found near Elgin in Scotland. [4] Newton discovered this specimen in conjunction with other new reptiles, but believed that G. elginensis’ characteristics were sufficiently unique to justify a new genus. [4] The holotype is the only known occurrence, and is housed at the Institute of Geological Sciences in London. [5] [6]
There have been two occurrences of the second species, G. locusticeps, both from the Ruhuhu Basin of Tanzania. [5] Originally discovered by Friedrich von Huene in 1942, [6] they are housed at the Institut und Museum für Geologie und Paläontologie der Universität Tübingen. [2]
The name Geikia was proposed by E.T. Newton, who investigated the G. elginensis holotype in 1892. [4] This name was a dedication to Sir Archibald Geikie, the Director-General of the Geological Survey at the time. [4]
The only G. elginensis specimen is a natural mold of a nearly complete skull and mandible, associated left humerus, and an isolated metapodial or proximal phalanx. [6] The G. locusticeps holotype is a skull lacking the tip of premaxilla, right quadrate, left temporal arch, and mandible. [6] Two occurrences of this species exist in the fossil record. [7] Analyses of G. elginensis and G. locusticeps indicate affinities to each other, but each also shares many characteristics with other taxa, including Pelanomodon , Oudenodon , and Ptychognathus ( Lystrosaurus ). [2] [4] [6] [3] The absence of some expected characteristics in G. elginensis could be explained under the assumption that it is subadult and, therefore, not fully developed. [2] However, ontogenetic changes of both Geikia and Pelanomodon, which would enable better analyses, remain uncertain. [2]
A high level of skull specialization was significant in the classification of Geikia. [3] Maisch and Gebauer considered the squared off anterior snout tip and reduced exposure of squamosal in occiput to be characteristics exclusively expressed in Geikia. [2] Prior to their analysis, Rowe described the generic diagnosis of Geikia as “dicynodonts having no tusk or postcanine teeth; highly vaulted palate; anterior palatal ridges of premaxilla reduced or absent; large palatine having rugose palatal surface; palatine having extensive contact with maxilla and premaxilla; length of interpterygoidal vacuity not less than half the length of the interpterygoidal fossa; interpterygoidal vacuity lying entirely within roof of interpterygoidal fossa; well developed maxillary caniniform process having pronounced lateral ridge; sharp occlusal margin of beak; sharp ridge or “keel” developed on central edge of maxilla behind caniniform process; septomaxilla having exposure on lateral surface of snout behind external nares; anterior surface of premaxilla flat, oriented vertically, and meeting lateral surface of premaxilla in abrupt “corner”; single, prominent preorbital protuberance". [6]
G. locusticeps was recognized as Dicynodon locusticeps until Timothy Rowe referred it to Geikia in 1980. [2] [6] It has been suggested that G. locusticeps may be a juvenile Pelanomodon tuberosus, and the two have been used synonymously. [2] In 2005, it was proposed that P. tuberosus be referred to G. locusticeps as a junior subjunctive synonym. [2]
Both belonging to the family Geikiidae, the genera Pelanomodon and Geikia are closely related. [2] Morphological differences between G. elginensis and G. locusticeps, as well as individual similarities to other species (especially those within Pelanomodon) have been utilized in debates regarding their classifications. [2] This exemplifies unresolved aspects of dicynodont taxonomy; it has even been suggested that single or incomplete dicynodont specimens should be considered incertae sedis until conclusions can be better ascertained through additional specimens or better preparation. [1] More recent literature attributes the cross-genus similarities between Pelanomodon and Geikia to plesiomorphic geikiid traits. [2] With these considerations, it is currently maintained that the generic distinction of Geikia is warranted. [2]
Compared to other dicynodonts, the shortened skull could be indicative of specific herbivory habits, such as biting off small pieces of vegetation. [3] Pertaining to mastication, crushing action was likely more emphasized than slicing, due to structural limitations of lower jaw movement. Additionally, these limitations could have conferred a "selective browser" role upon Geikia. [3] [8] Aside from jaw specialization, the forward position and large size of the orbits could suggest a degree of stereoscopic vision. [3] Rotational ability of the eyes could have enabled Geikia to see in a variety of directions, such as through notches in the frontals. [3] Cruickshank indicated that these characteristics, along with the loss of tusks, could be suggestive of nocturnal behaviors. [3]
G. elginensis was discovered in the Cutties Hillock Sandstone Formation in Scotland. [9] The coarse, hard sand presented difficulty in the development of the specimen, [4] which was recovered from a pebbly layer near the base of the formation. [9] The pebbles are characteristic of water deposition, suggesting a fluvial environment. [7] Analyses have suggested that this formation is of Permian age, specifically Late Tatarian. [9] Assuming accuracy of age assessment, this could be representative of offshore Zechstein Sea deposits. [7]
Also estimated to be of Tatarian age, G. locusticeps was discovered in the Usili Formation (formerly Kawinga Formation) of Kingori in southwest Tanzania. [2] [10] The environment was identified as being terrestrial. [5] In a 2010 publication, Sidor et al. concluded that subsidence events during this time conferred a transition from alluvial fans to an axial braided channel, ultimately equilibrating as an alluvial plain with rivers and lakes. [10]
Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivores that typically bore a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, typically toothless beak, unique amongst all synapsids. Dicynodonts first appeared in Southern Pangaea during the mid-Permian, ca. 270–260 million years ago, and became globally distributed and the dominant herbivorous animals in the Late Permian, ca. 260–252 Mya. They were devastated by the end-Permian Extinction that wiped out most other therapsids ca. 252 Mya. They rebounded during the Triassic but died out towards the end of that period. They were the most successful and diverse of the non-mammalian therapsids, with over 70 genera known, varying from rat-sized burrowers to elephant-sized browsers.
Robertia is an extinct genus of small herbivorous dicynodonts from the Middle to Late Permian of South Africa, between 260 and 265 million years ago. It is a monospecific genus, consisting of the type-species R. broomiana, which was classified by Lieuwe Dirk Boonstra in 1948 and named in honor of Robert Broom for his study of South African mammal-like reptiles.
Dicynodon is a genus of dicynodont therapsid that flourished during the Upper Permian period. Like all dicynodonts, it was an herbivorous animal. This synapsid was toothless, except for prominent tusks, hence the name. It probably cropped vegetation with a horny beak, much like a tortoise, while the tusks may have been used for digging up roots and tubers.
Endothiodon is an extinct genus of medium to large dicynodont from the Late Permian. Like other dicynodonts, Endothiodon was an herbivore, but it typically lacked the two tusks that characterized most other dicynodonts and instead had long rows of teeth inset in the jaws that replaced in waves. The anterior portion of the upper and lower jaw are curved upward, creating a distinct beak that is thought to have allowed them to be specialized grazers.
Theriognathus is an extinct genus of therocephalian therapsid belonging to the family Whaitsiidae, known from fossils from South Africa, Zambia, and Tanzania. Theriognathus has been dated as existing during the Late Permian. Although Theriognathus means mammal jaw, the lower jaw is actually made up of several bones as seen in modern reptiles, in contrast to mammals. Theriognathus displayed many different reptilian and mammalian characteristics. For example, Theriognathus had canine teeth like mammals, and a secondary palate, multiple bones in the mandible, and a typical reptilian jaw joint, all characteristics of reptiles. It is speculated that Theriognathus was either carnivorous or omnivorous based on its teeth, and was suited to hunting small prey in undergrowth. This synapsid adopted a sleek profile of a mammalian predator, with a narrow snout and around 1 meter long. Theriognathus is represented by 56 specimens in the fossil record.
Dicynodontoides is a genus of small to medium-bodied, herbivorous, emydopoid dicynodonts from the Late Permian. The name Dicynodontoides references its “dicynodont-like” appearance due to the caniniform tusks featured by most members of this infraorder. Kingoria, a junior synonym, has been used more widely in the literature than the more obscure Dicynodontoides, which is similar-sounding to another distantly related genus of dicynodont, Dicynodon. Two species are recognized: D. recurvidens from South Africa, and D. nowacki from Tanzania.
Daptocephalus is an extinct genus of dicynodont synapsid, which was found in Late Permian strata, in a biozone known precisely for the presence of fossils of this dicynodont, the Daptocephalus Assemblage Zone, in the Karoo Basin in South Africa. An additional species, D. huenei, is known from the Usili Formation in Tanzania and was formerly assigned to the genus Dicynodon before a study in 2019 recognised that the type specimen belonged to Daptocephalus.
Eosimops is an extinct genus of pylaecephalid dicynodonts. They were small synapsids superficially resembling modern mammals. Eosimops is known from several skull specimens, as well as one complete skeleton. Eosimops lived during the Middle Permian of South Africa.
Kawingasaurus is an extinct genus of dicynodont therapsid from the Late Permian Usili Formation of Tanzania. It is a member of the family Cistecephalidae, and like other cistecephalids it is thought to have been fossorial. It is a member of the family Cistecephalidae. Cistephalidae includes genera Cisteceohalus, Cistecephaloides and Kawingasaurus. Greek for Saurus meaning “lizard” appears as a suffix denoting a reptilian origin. Living between 254.17 and 259.9 million years ago in the late Permian and believed to have the first and last recorded appearance in this time period. It lived in deep burrows as a suggested by most burrowing dicynodonts from evaluation of cranial sutures, vestibule inflation and enlarged stapes foot plates which are thought to be functionally correlated with bone-conduction hearing; all observed in fossorial vertebrates which use seismic signals as communication.
Odontocyclops is an extinct genus of Dicynodonts that lived in the Late Permian. Dicynodonts are believed to be the first major assemblage of terrestrial herbivores. Fossils of Odontocyclops have been found in the Karoo Basin of South Africa and the Luangwa Valley of Zambia. The phylogenetic classification of Odontocyclops has been long under debate, but most current research places them as their own genus of Dicynodonts and being very closely related to Rhachiocephalus and Oudenodon.
Pelanomodon is an extinct genus of dicynodont therapsids that lived in the Late Permian period. Fossil evidence of this genus is principally found in the Karoo Basin of South Africa, in the Dicynodon Assemblage Zone. Lack of fossil record after the Late Permian epoch suggests that Pelanomodon fell victim to the Permian-Triassic extinction event.
Sangusaurus is an extinct genus of large dicynodont synapsid with two recognized species: S. edentatus and S. parringtonii. Sangusaurus is named after the Sangu stream in eastern Zambia near to where it was first discovered + ‘saur’ which is the Greek root for lizard. Sangusaurus fossils have been recovered from the upper parts of the Ntawere Formation in Zambia and of the Lifua Member of the Manda Beds in Tanzania. The earliest study considered Sangusaurus a kannemeyeriid dicynodont, but more recent phylogenetic analyses place Sangusaurus within the stahleckeriid clade of Dicynodontia. Until recently, little work had been done to describe Sangusaurus, likely due to the fact that only four incomplete fossil specimens have been discovered.
Tetragonias is an extinct genus of dicynodont from the Anisian Manda Beds of Tanzania. With tetra meaning “four,” and goni meaning “angle,” the name references the square shape of the Tetragonias skull when viewed dorsally. Not to be confused with the plant Tetragonia,Tetragonias were dicynodont anomodonts discovered in the late 1960s by paleontologist A. R. I. Cruickshank in the Manda Formation. Only the type species, T. njalilus, has been recognized.
"Dixeya" nasuta is a extinct species of gorgonopsian that lived during the Late Permian of East Africa, known from fossils found in what is now Tanzania. The species has a complicated taxonomic history, it was originally named as a second species of the genus Dixeya which is now considered a junior synonym of Aelurognathus. "D." nasuta itself, however, was not moved to Aelurognathus, and although it was instead tentatively referred to Arctognathus at first it has since been recognised to not belong to this genus either. This situation leaves "Dixeya" nasuta without a formal genus name. It was proposed to belong to a new distinct genus, named "Njalila", that was informally proposed for the species in a PhD thesis, but this name has not yet been formally published and is currently a nomen nudum. "D." nasuta has been characterised from other gorgonopsians by a combination of its straight snout profile, upturned and "pinched" nose, and curved jaw margin. The fossil record of the Usili Formation shows that the taxon was contemporary with many other gorgonopsians, even alongside large representatives such as Inostrancevia and rubidgeines.
Syops is an extinct genus of dicynodont therapsid. The type species S. vanhoepeni was first named in 1938 as Dicynodon vanhoepeni. Fossils of the genus have been found in the Cistecephalus Assemblage Zone in the Usili Formation of the Ruhuhu Basin, Tanzania and the Upper Madumabisa Mudstone Formation of the Luangwa Basin, Zambia. Its phylogenetic placement is somewhat uncertain, with multiple different studies finding it as either a basal geikiid, rhachiocephalid a dicynodontoid more derived than the most basal genera but less derived than Lystrosauridae, or a lystrosaurid.
Gordonia is an extinct genus of dicynodont therapsid from the Late Permian of Scotland. Fossils have been found from the Elgin sandstone of Cutties Hillock Sandstone in Elgin, Moray. These are among the many amniote fossils referred to as the Elgin Reptiles. Gordonia was named in 1893 with four species: G. traquairi, G. duffiana, G. huxleyana, and G. juddiana. Currently, the only recognized species is the type G. traquairi. All other species are considered synonyms of the type.
The Usili Formation is a Late Permian geologic formation in Tanzania. It preserves fossils of many terrestrial vertebrates from the Permian, including temnospondyls, pareiasaurs, therapsids and the archosauromorph Aenigmastropheus.
Counillonia is an extinct genus of dicynodont therapsid from the area of Luang Prabang in Laos, Southeast Asia that lived at around the time of the Permian-Triassic boundary and possibly dates to the earliest Early Triassic. Its type and only known species is C. superoculis. Counillonia was related to the Triassic dicynodonts such as Lystrosaurus and the Kannemeyeriiformes that survived the Permian mass extinction, but it was more closely related to the Permian genus Dicynodon than to either of these lineages. Counillonia may then possibly represent another line of dicynodonts that survived the Permian mass extinction into the Triassic period, depending on its age. The discovery of Counillonia in Laos and its unexpected evolutionary relationships hint at the less well understood geographies of dicynodont diversity across the Permo-Triassic boundary outside of well explored regions like the Karoo Basin in South Africa.
Repelinosaurus is an extinct genus of dicynodont from the Purple Claystone Formation of Luang Prabang in Laos, Southeast Asia that lived at around the time of the Permian-Triassic boundary and possibly dates to the earliest Early Triassic. Its type and only known species is R. robustus. Repelinosaurus was originally described as the earliest known kannemeyeriiform dicynodont, supporting the idea of a more rapid radiation of the Triassic kannemeyeriiform dicynodonts during the Early Triassic following the Permian mass extinction. However, it may alternatively be more closely related to the Permian Dicynodon. The discovery of a potential early kannemeyeriiform in an understudied locality like Laos highlights the importance of such places in dicynodont research, which has been largely focused on historically important localities such as the Karoo Basin of South Africa.
Taoheodon is an extinct genus of dicynodont therapsid from the Sunjiagou Formation in the Shanxi province of China, dated to the Wuchiapingian age of the Late Permian. Its type and only known species is T. baizhijuni. Taoheodon was a close relative of the well known Dicynodon, and may represent a biogeographical link between the South African Dicynodon and similar dicynodonts found in Laos.