Geikia elginensis
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''Geikia'' 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
dicynodont Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivorous animals with a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, typic ...
therapsid Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more ...
s from the
late Permian Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effect, ...
. 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. Only two species, ''Geikia locusticeps'' and ''Geikia elginensis'' have been assigned to this genus. While this is the currently accepted classification, fossil record limitations have led to repeated debate on the genus assignments of these species.


Description

A high level of skull specialization was significant in the classification of ''Geikia''. Maisch and Gebauer considered the squared off anterior snout tip and reduced exposure of squamosal in occiput to be characteristics exclusively expressed in ''Geikia''. 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".


Discovery and naming

This genus was established in 1893 following
Edwin Tulley Newton Edwin Tulley Newton (4 May 1840 – 28 January 1930) was a British paleontologist. Newton originally worked at handicrafts, but was able to attend Thomas Henry Huxley's lectures and by 1865, was appointed as his assistant. In 1882, he becam ...
’s discovery of a new dicynodont fossil, now known as ''G. elginensis'', one of the
Elgin Reptiles Elgin Reptiles is the name given to the Permian and Triassic fossils found in the sandstone deposits in and around the town of Elgin, in Moray, Scotland. They are of historical and scientific importance, and many of the specimens are housed in th ...
found near Elgin in Scotland. Newton discovered this specimen in conjunction with other new reptiles, but believed that ''G. elginensis’'' characteristics were sufficiently unique to justify a new genus. The holotype is the only known occurrence, and is housed at the Institute of Geological Sciences in London. There have been two occurrences of the second species, ''G. locusticeps,'' both from the Ruhuhu Basin of Tanzania. Originally discovered by Friedrich von Huene in 1942, they are housed at the Institut und Museum für Geologie und Paläontologie der Universität Tübingen. The name ''Geikia'' was proposed by E.T. Newton, who investigated the ''G. elginensis''
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 ...
in 1892. This name was a dedication to Sir
Archibald Geikie Sir Archibald Geikie (28 December 183510 November 1924) was a Scottish geologist and writer. Early life Geikie was born in Edinburgh in 1835, the eldest son of Isabella Thom and her husband James Stuart Geikie, a musician and music critic. T ...
, the Director-General of the Geological Survey at the time.


Specimens

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. The ''G. locusticeps'' holotype is a skull lacking the tip of premaxilla, right quadrate, left temporal arch, and mandible. Two occurrences of this species exist in the fossil record. Analyses of ''G. elginensis'' and ''G. locusticeps'' indicate affinities to each other, but each also shares many characteristics with other taxa, including ''
Pelanomodon ''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.Kammerer, C. F., K. D ...
'', '' Oudenodon'', and ''Ptychognathus'' (''
Lystrosaurus ''Lystrosaurus'' (; 'shovel lizard'; proper Greek is λίστρον ''lístron'' ‘tool for leveling or smoothing, shovel, spade, hoe’) is an extinct genus of herbivorous dicynodont therapsids from the late Permian and Early Triassic epochs ( ...
''). The absence of some expected characteristics in ''G. elginensis'' could be explained under the assumption that it is subadult and, therefore, not fully developed. However,
ontogenetic Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the st ...
changes of both ''Geikia'' and ''Pelanomodon'', which would enable better analyses, remain uncertain.


Classification

''G. locusticeps'' was recognized as ''Dicynodon locusticeps'' until Timothy Rowe referred it to ''Geikia'' in 1980. It has been suggested that ''G. locusticeps'' may be a juvenile ''Pelanomodon tuberosus'', and the two have been used synonymously. In 2005, it was proposed that ''P. tuberosus'' be referred to ''G. locusticeps'' as a junior subjunctive synonym. Both belonging to the family
Geikiidae Geikiidae is a family of Late Permian dicynodonts. Fossils are known from Scotland, South Africa, and Tanzania. The family was first named by Franz Nopcsa in 1923, although Friedrich von Huene's 1948 description of the family brought it into com ...
, the genera ''Pelanomodon'' and ''Geikia'' are closely related. 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. This exemplifies unresolved aspects of dicynodont taxonomy; it has even been suggested that single or incomplete dicynodont specimens should be considered ''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertain ...
'' until conclusions can be better ascertained through additional specimens or better preparation. More recent literature attributes the cross-genus similarities between ''Pelanomodon'' and ''Geikia'' to plesiomorphic geikiid traits. With these considerations, it is currently maintained that the generic distinction of ''Geikia'' is warranted.


Paleobiology

Compared to other dicynodonts, the shortened skull could be indicative of specific herbivory habits, such as biting off small pieces of vegetation. 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''. Aside from jaw specialization, the forward position and large size of the orbits could suggest a degree of
stereoscopic vision Stereopsis () is the component of depth perception retrieved through binocular vision. Stereopsis is not the only contributor to depth perception, but it is a major one. Binocular vision happens because each eye receives a different image becaus ...
. Rotational ability of the eyes could have enabled ''Geikia'' to see in a variety of directions, such as through notches in the frontals. Cruickshank indicated that these characteristics, along with the loss of tusks, could be suggestive of nocturnal behaviors.


Paleoecology

''G. elginensis'' was discovered in the Cutties Hillock Sandstone Formation in Scotland. The coarse, hard sand presented difficulty in the development of the specimen, which was recovered from a pebbly layer near the base of the formation. The pebbles are characteristic of water deposition, suggesting a fluvial environment. Analyses have suggested that this formation is of Permian age, specifically Late Tatarian. Assuming accuracy of age assessment, this could be representative of offshore Zechstein Sea deposits. Also estimated to be of Tatarian age, ''G. locusticeps'' was discovered in the Usili Formation (formerly Kawinga Formation) of Kingori in southwest Tanzania. The environment was identified as being terrestrial. In a 2010 publication, Sidor et al. concluded that subsidence events during this time conferred a transition from
alluvial fan An alluvial fan is an accumulation of sediments that fans outwards from a concentrated source of sediments, such as a narrow canyon emerging from an escarpment. They are characteristic of mountainous terrain in arid to semiarid climates, but a ...
s to an axial braided channel, ultimately equilibrating as an alluvial plain with rivers and lakes.


References


The main groups of non-mammalian synapsids at Mikko's Phylogeny Archive
{{Taxonbar, from=Q5530121 Dicynodonts Permian synapsids Prehistoric synapsids of Europe Fossil taxa described in 1892 Fossil taxa described in 1942 Anomodont genera