Aleodon Skull Lateral
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''Aleodon'' is an extinct genus of
cynodont Cynodontia () is a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 Megaannum, mya), and extensively diversified after the Permian–Triassic extinction event. Mammals are cynodonts, as are their extin ...
s that lived from the Middle to Late Triassic. Relatively few analyses have been conducted to identify the
phylogenetic 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 dat ...
placement of ''Aleodon'', although some have placed it as a sister taxon to '' Chiniquodon''. Two species of ''Aleodon'' are recognized: ''A. brachyrhamphus'' which was discovered in Tanzania, and ''A. cromptoni'' which was discovered most recently in Brazil. The name for the genus ''Aleodon'' was created when Alfred W. "Fuzz" Crompton initially discovered the type species, ''Aleodon brachyrhamphus''. The genus name, "''Aleodon''" referred to the grinding nature of the postcanine teeth, while "''brachyrhamphus''" referred to the relatively short snout of the specimen. The most recently discovered species, ''A. cromptoni'' was named after Crompton.


Discovery and classification

In 1955, ''Aleodon'' was initially classified as a gomphodont cynodont based on the partial skull and lower jaw fossils found in 1933 in Tanzania. The classification was based on the presence of three types of post-canines that were identified in the fossil that were similar to another well-known gomphodont, ''Diademodon tetragonus''. It was later argued in 2001 by Hopson and Kitching that ''Aleodon'' be classified under
Chiniquodontidae Chiniquodontidae is an extinct family of basal probainognathian cynodonts that lived in what is now Africa and South America during the Middle and Late Triassic. It is currently thought to include four valid genera: ''Aleodon'', '' Chiniquodon'' ...
based on less-worn dentition of unpublished specimens. This classification was accepted by many sources. However, a study done the following year by Abdala and Gianni found, based on fossils found in Namibia, that ''Aleodon'' (and '' Cromptodon'') both had well developed lingual cingular platforms in their post-canines, a characteristic that Chiniquodontidae did not possess. While not many analyses have been done to clarify the phylogenetic relationships of ''Aleodon'' thus far, those that have been done describe this genus as the sister taxon of ''Chiniquodon''.


Description


Dentition

The dentition of ''Aleodon'' is the most significant morphological feature to distinguish it from other genera. Crompton pointed out three distinct regions of the postcanine row: anterior circular, transversely expanded ovate, and sectorial posterior teeth (described as “shearing” by Crompton). These three regions of post canines were very similar to a well-known gomphodont, '' Diademodon tetragonus'' (which led Crompton to believe that ''Aleodon'' was part of Gomphodonta). Crompton also noted that ''Aleodon'' uniquely possessed two incisors on the lower jaw which differs from most cynodonts which primitively had four. Based on more fossils, ''Aleodon'' was also found to have a long secondary palate, a characteristic that was shared with members of the monophyletic group Chiniquodontidae and resulted in its reidentification to
Probainognathia Probainognathia is one of the two major subgroups of the clade Eucynodontia, the other being Cynognathia. The earliest forms were carnivorous and insectivorous, though some groups eventually also evolved herbivorous diets. The earliest and most b ...
from Gomphodonta. Shortly after, specimens of ''Aleodon'' showed the possession of well-developed lingual cingular platforms on the post canines. This finding resulted in some researchers removing ''Aleodon'' from
Chiniquodontidae Chiniquodontidae is an extinct family of basal probainognathian cynodonts that lived in what is now Africa and South America during the Middle and Late Triassic. It is currently thought to include four valid genera: ''Aleodon'', '' Chiniquodon'' ...
due to the group being not characterized by the possession of well-developed lingual cingular platforms. The lingual cingular platform that is expanded in both labiolingual and mesiodistal dimensions relative to the labial portion of the crown is also the most developed of all Triassic probainognathians. These observations led some researchers to believe that ''Aleodon'' (along with ''
Candelariodon ''Candelariodon'' is an extinct genus of carnivorous probainognathian cynodonts from the Middle to Late Triassic Santa Maria Formation of the Paraná Basin in Rio Grande do Sul state, Brazil. ''Candelariodon'' is known from a partial mandible h ...
'') may be specimens that exhibit the gradual development of the cingular platform.


Skull

Based on specimens found in Brazil, ''Aleodon'' was found to share the same general cranial morphology of Chiniquodontidae. Diagnostic features of Chiniquodontidae are zygomatic arches flaring laterally, angulation between the ventral edge of the maxillary zygomatic process and the anteroventral margin of the jugal, elongated pterygoid flanges that end in a thin projection, and a long secondary palate. When ''Aleodon'' was first described by Crompton, he noted the presence of a pineal foramen, though later specimens showed no evidence of a pineal foramen on this taxon's skull. ''Aleodon'' is also characterized by its relatively short snout.


Geological/Paleoenvironmental information


Manda Formation (Tanzania)

Specimens of ''Aleodon'' were first recovered from the Triassic Manda Beds of Tanzania by Parrington in the 1930s. The Manda Beds were first surveyed by Gordon M. Stockley from June to October 1930. There was further collection by Parrington and Nowack in the 1930s in which ''Aleodon brachyrhamphus'' was later described by Crompton. The collections showed that the Manda Beds contained fauna such as thecodontians, cynodonts, and dicynodonts, as well as a large amphibian and a rhynchosaur. The formation was described as being 2000 meters thick with purple to brown mudstones with grey sandstones in between. From 60 to 150 meters below the top of the formation, reptile bones were found while most (65%) of the specimens were found 1180 and 1310 meters below the surface.


Omingonde Formation (Namibia)

Fossil vertebrates in the
Omingonde Formation The Omingonde Formation is an Early Triassic, Early to Middle Triassic (Anisian to Ladinian) geologic Formation (geology), formation, part of the Karoo Supergroup, in the western Otjozondjupa Region and northeastern Erongo Region of north-central N ...
of Namibia were first found and published by Keyser in 1973. Among the fossils, he described amphibians, cynodonts and dicynodonts. The cynodonts were initially represented by ''
Cynognathus ''Cynognathus'' is an extinct genus of large-bodied cynodontian therapsids that lived in the Middle Triassic. It is known from a single species, ''Cynognathus crateronotus''. ''Cynognathus'' was a predator closely related to mammals and had a s ...
'' and herbivorous gomphodonts. After the discovery of ''Aleodon'' among other cynodont specimens such as ''Chiniquodon'', '' Luangwa'', and an unidentified traversodontid in 2009, the Omingonde Formation is considered to possess the most diverse fauna of Middle Triassic cynodonts in the world. The discovery also provided researchers a definite link between two faunas in South America and East Africa. From this, the Omingonde Formation was also considered the key in "reconstructing the biogeography of Southern Gondwana during the Middle Triassic".


Rio Grande do Sul (Brazil)

Most recently, a new species of ''Aleodon'' (''A. cromptoni'') was found in
Rio Grande do Sul Rio Grande do Sul (, ; ; "Great River of the South") is a Federative units of Brazil, state in the South Region, Brazil, southern region of Brazil. It is the Federative units of Brazil#List, fifth-most populous state and the List of Brazilian s ...
, southern Brazil. The specimens were first collected in 1988 by Daniel and Abraão Cargnin.


References

{{Taxonbar, from1=Q2081682, from2=Q20716967 Probainognathia Middle Triassic synapsids of Africa Middle Triassic synapsids of South America Late Triassic synapsids of South America Fossils of Brazil Triassic Tanzania Fossils of Tanzania Triassic Namibia Fossils of Namibia Omingonde Formation Rio Grande do Sul Fossil taxa described in 1955