Pelanomodon
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''Pelanomodon'' is an extinct genus 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 ...
therapsids that lived in the Late
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleo ...
period. Fossil evidence of this genus is principally found in the
Karoo Basin The Karoo Supergroup is the most widespread stratigraphic unit in Africa south of the Kalahari Desert. The supergroup consists of a sequence of units, mostly of nonmarine origin, deposited between the Late Carboniferous and Early Jurassic, a perio ...
of South Africa, in the ''Dicynodon'' Assemblage Zone.Kammerer, C. F., K. D. Angielczyk, and Jörg Fröbisch. 2015. Redescription of the geikiid Pelanomodon (Therapsida, Dicynodontia), with a reconsideration of ‘Propelanomodon’. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2015.1030408. http://www.tandfonline.com/doi/full/10.1080/02724634.2015.1030408 Lack of fossil record after the Late Permian epoch suggests that ''Pelanomodon'' fell victim to the Permian-Triassic extinction event. The name ''Pelanomodon'' can be broken up into three parts; “pelos” meaning mud, “anomo” meaning irregular and “odon” meaning tooth. Together, this suggests ''Pelanomodon'' to be a mud dwelling
anomodont Anomodontia is an extinct group of non-mammalian therapsids from the Permian and Triassic periods. By far the most speciose group are the dicynodonts, a clade of beaked, tusked herbivores.Chinsamy-Turan, A. (2011) ''Forerunners of Mammals: Ra ...
(a group of therapsids that are characterized by their lack of teeth). The Karoo Basin during this period was characterized by its extensive flood plains, so to hypothesize a mud based habitat for this genus is not far fetched. ''Pelanomodon'' is in the family Geikiidae along with ''
Aulacephalodon ''Aulacephalodon'' is an extinct genus of medium-sized dicynodonts, or non-mammalian synapsids, that lived during Permian period, about 299-252 million years ago. Individuals of ''Aulacephalodon'' are commonly found in the Lower Beaufort Group o ...
'' and ''
Geikia ''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 relat ...
. Aulacephalodon'' is believed to have lived alongside ''Pelanomodon'' in the Karoo Basin, where as records of ''Geikia'' have been discovered in Scotland and Tanzania. ''Pelanomodon'' is primarily characterized and distinguished from ''Aulacephalodon'' by its lack of tusks. Other crananial features have been used by paleontologists to establish two species of ''Pelanomodon;'' ''P. moschops'' and ''P. rubidgei.'' However, recent analysis points to the conclusion that these may in fact the same species.


Description


Skull

The fossil record of ''Pelanomodon'' is principally made up of both complete and partial skulls. It is for this reason that morphology of the skull is used to distinguish ''Pelanomodon'' from other genus in the Geikiines family. The absence of tusks is a significant feature that is used to differentiate ''Pelanomodon'' from ''Aulacephalodon'', in addition to bosses on the post orbital bar and twisting of the zygoma. ''Pelanomodon'' is distinguished from ''Geikia'' due to its longer temporal fenestra and snout, lesser developed oesophageal crest and flush pineal foramen. In addition to these distinctions, Pelanomodon's skull has many other characteristic features. One of the most apparent of these features being that its skull is wider than it is tall, resulting in a triangular shape, when viewed dorsally. In connection with this, the temporal fenestra angle out posterolaterally. Compared to other dicynodonts, the external naris is retracted and located relatively high on the snout. Likewise, the orbits are also placed higher than is typically seen in dicynodonts. In accordance with other Anomodonts, ''Pelanomodon'' have cheek teeth but lack their molar teeth. In addition, like other therapsids, the premaxilla extends on the ventral side in order to form a secondary palate.


Discovery and naming

''Pelanomodon'' was first discovered by
Robert Broom Robert Broom FRS FRSE (30 November 1866 6 April 1951) was a British- South African doctor and palaeontologist. He qualified as a medical practitioner in 1895 and received his DSc in 1905 from the University of Glasgow. From 1903 to 1910, he ...
, in the Karoo Basin of South Africa, in 1913. Its classification was based on a nearly complete skull, missing only the lower jaw. The species was originally named ''Dicynodon moschops'', however, in 1969 an analysis done by A. W. Keyser reclassified the specimen as ''Pelanomodon moschops.''Keyser, A.W (1969). Re-evaluation of the systematics and morphology of certain anomodont Therapsida. Ph.D. Thesis, University of the Witwatersrand, Johannesburg'' ''Another species, ''Pelanomodon rubidgei,'' was classified by Broom in 1938, and was established from a complete skull found in the same region, by S. H Rubidge. Between the years of 1913 and 1950, Broom described five different ''Pelanomodon'' species, based on differences observed between many partial, and complete skulls. However, since this time, several paleontologists have re-evaluated the morphology of the skulls and reclassified them all as either ''Pelanomodon rubidgei'' or ''Pelanomodon moschops.'''' ''Since fossil records can never provide a complete understanding of the past, these classifications are still debated among paleontologists. In 2016, a new analysis suggested that the morphological features used to distinguish between ''Pelanomodon moschops'' and ''Pelanomodon rubidgei'' are actually a result of
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...
within the same species. The same analysis claims that ''Propelanomodon'', another tuskless dicynodont genus, first described by Broom in 1913, is in fact a juvenile form of ''Pelanomodon''. The authors of this analysis would classify all these species as ''Pelanomodon moschops''. The basis for these conclusions are discussed below.


Paleobiology


Sexual dimorphism

In the past, skull features have also been used to distinguish possible ''Pelanomodon'' species from one another. However, more recent analysis suggests that these disparities between specimens may not suggest more than one species, but rather sexual dimorphism within the same species. Two species have been previously classified, ''Pelanomodon moschops'' (''P. moschops''), identified by having a skull longer than 18 cm in length and relatively small cranial bosses and ''Pelanomodon rubidge''i (''P. rubidgei''), identified by having a skull longer than 18 cm but with relatively large cranial bosses. ''P. moschops'' and ''P. rubidgei'' have been hypothesized to be of the same species by several paleontologists, due to the fact that few structural differences exist between the skulls of the two species. The most prominent distinction between these two skull types is the presence and size of the cranial bosses. ''P. moschops'' lacks the supraorbital ridge and postorbital bosses that are seen in ''P. rubidgei'', and although it maintains the nasal and prefrontal bosses, they are much smaller. Other than these seemingly superficial features, the species share all other cranial features. It is for this reason that it is hypothesized the differences are a result of sexual dimorphism within one species, ''P. moschops''.


Ontogeny

''Propelanomodon tylorhinus'' is another dicynodont species that has recently been hypothesized to be a juvenile form of ''P. moschops''. It has a shorter skull, typically shorter than long. Several key cranial features differ within this species when compared to ''P. moschops''’ skulls, which leaves there to be debate among paleontologists about its relationship to ''Pelanomodon''. For example, the temporal fenestra of ''Propelanomodon'' point anterposteriorly, rather than being angled slightly laterally. As a result of this, its skull does not have the distinct triangular shape of ''Pelanomodon''. In addition, ''Propelanomodon''’s snout is relatively shorter and its orbits relatively larger. The cranial bosses that are very distinct of ''Pelanomodon'' are also absent, except for the nasal bosses. Supporters of this hypothesis claim that it is not unusual to see large cranial changes in vertebrae as they mature.


Diet

Like other Dicynodonts, ''Pelanomodon'' is believed to have been an herbivore as its lack of canine teeth would inhibit it from having a carnivorous diet. In addition, ''Pelanomodon'' is described to have a similar overall skull to ''Aulacephalodon.'' Keyeser describes that the tip of the snout is supported by ridges along the palatal and snout's external surface which restricts the chewing motion to just the tip of the jaw. This combined with some of the other cranial features described above, such as the placement of the nares and orbit, suggests that the animal may have consumed plants from shallow waters.


Paleoecology

The fossil record of ''Pelanomodon'' has thus far been restricted to the Karoo Basin in Southern Africa. This region chronicles constant sediment accumulation from the Carboniferous period and into the Jurassic period. ''Pelanomodon'', along with other dicynodonts are predominantly found as a part of the
Beaufort Group The Beaufort Group is the third of the main subdivisions of the Karoo Supergroup in South Africa. It is composed of a lower Adelaide Subgroup and an upper Tarkastad Subgroup. It follows conformably after the Ecca Group and unconformably underli ...
. The Beaufort Group geological strata ranges from the
middle Permian The Guadalupian is the second and middle series/ epoch of the Permian. The Guadalupian was preceded by the Cisuralian and followed by the Lopingian. It is named after the Guadalupe Mountains of New Mexico and Texas, and dates between 272.95 ± ...
to middle Triassic period. Sedimentary analysis suggests that during this time the region was characterized by the extensive flood plains of a few large rivers. The climate during the late Permian period, in this region was semi-arid and rainfall was seasonal. This climate, combined with the important role that rivers played in establishing the environment, allowed for a variety of riparian vegetation to grow, which in turn lead to a high diversity of animals that were all able to fill a different niche. In fact, dicynodonts have been found to be the most abundant
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 ...
in this range.


References

{{Taxonbar, from=Q15712862 Dicynodonts Permian synapsids of Africa Fossil taxa described in 1938 Taxa named by Robert Broom Anomodont genera Permian genus extinctions