''Tropidostoma'' is a medium-sized
herbivorous
A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
oudenodontid
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, typicall ...
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 ...
that lived during 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 Paleoz ...
(
Lopingian
The Lopingian is the uppermost series/last epoch of the Permian. It is the last epoch of the Paleozoic. The Lopingian was preceded by the Guadalupian and followed by the Early Triassic.
The Lopingian is often synonymous with the informal terms l ...
) period in
South Africa
South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring countri ...
. The first ''Tropidostoma'' fossil was described by
Harry Govier Seeley
Harry Govier Seeley (18 February 1839 – 8 January 1909) was a British paleontologist.
Early life
Seeley was born in London on 18 February 1839, the second son of Richard Hovill Seeley, a goldsmith, and his second wife Mary Govier. When his fat ...
in 1889. Later two
subspecies
In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
were identified. ''Tropidostoma''
fossil
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s are an
index fossil
Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock Stratum, strata by using the fossil assemblages contained within them.Hine, Robert. “Biostratigraphy.” ''Oxford Reference: Dictiona ...
in a
biozone
In biostratigraphy, biostratigraphic units or biozones are intervals of geological strata that are defined on the basis of their characteristic fossil taxa, as opposed to a lithostratigraphic unit which is defined by the lithological properties of ...
of the
Karoo Basin
The Karoo Supergroup is the most widespread stratigraphy, stratigraphic unit in Africa south of the Kalahari Desert. The group (stratigraphy), supergroup consists of a sequence of units, mostly of nonmarine origin, deposited between the Pennsylvan ...
known as the
''Tropidostoma'' Assemblage Zone. This
biozone
In biostratigraphy, biostratigraphic units or biozones are intervals of geological strata that are defined on the basis of their characteristic fossil taxa, as opposed to a lithostratigraphic unit which is defined by the lithological properties of ...
is characterized by the presence of this species in association with another
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, typicall ...
species, ''
Endothiodon uniseries''.
History of discovery
The first ''Tropidostoma''
fossil
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
material was found during a field expedition in the Upper
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 Paleoz ...
-aged Teekloof Formation 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 underlie ...
. This material was later described by
Seeley (1889) in a study in which he described two
fossil
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s which had been named ''Dicynodon microtrema'' and ''Tropidostoma dunni''. In 1915, several years after
Seeley's death, the paleontologist,
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 ...
, reexamined the same material and discovered that the fossil material in question was of the same
species
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
. This new single species was renamed ''Tropidostoma microtrema''. Some years later, the name ''T. microtrema'' was changed to ''Tropidostoma dubium'', and ''Tropidostoma dunni'' is now considered to be the type species.
Description
Two
subspecies
In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
exist among the specimens referred to as ''Tropidostoma dubium'' and ''Tropidostoma dunni'' . ''T. dubium'' is observed to have two
cranial
Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
morphs, one being robust form with a tall snout and large
tusk
Tusks are elongated, continuously growing front teeth that protrude well beyond the mouth of certain mammal species. They are most commonly canine teeth, as with pigs and walruses, or, in the case of elephants, elongated incisors. Tusks share c ...
s and the other more gracile with a low snout and small or no tusks. The robust and gracile forms are considered to either represent
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 ani ...
or individual variation.
Classification
''Tropidostoma'' is currently classified as an oudenodontid within the larger
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, typicall ...
clade
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
Bidentalia
Bidentalia is a group of dicynodont therapsids. Bidentalia was one of the first names used to describe dicynodonts; the group was established in 1876, while the name "bidentals" dates back as far as 1845. With the increasing prominence of phyloge ...
. This
clade
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
is characterized collectively by their reduced
dentition
Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiolo ...
with only their
maxilla
The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
ry
tusk
Tusks are elongated, continuously growing front teeth that protrude well beyond the mouth of certain mammal species. They are most commonly canine teeth, as with pigs and walruses, or, in the case of elephants, elongated incisors. Tusks share c ...
s being intact. However, many species in this
clade
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
sporadically lack tusks completely and their
fossil
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s only hold evidence that they retained their
keratin
Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up scales, hair, nails, feathers, ho ...
ous beaks. Many ''Tropidostoma'' fossils previously collected have been misidentified as other species, such as of ''
Oudenodon bainii
''Oudenodon'' is an extinct genus of dicynodont. It was common throughout southern Africa during the Late Permian. Several species of ''Oudenodon'' are known. Both ''O. bainii'', the type species, and ''O. grandis'' are known from South Africa ...
'' due to their remarkable similarity. In addition, the type fossil of the recently discovered ''
Bulbasaurus phylloxyron'' was misidentified as a ''Tropidostoma''
fossil
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
for several years.
See also
*
List of therapsids
This list of therapsids is an attempt to create a comprehensive listing of all genera that have ever been included in the Therapsida excluding mammals and purely vernacular terms. The list includes all commonly accepted genera, but also genera tha ...
References
{{Taxonbar, from=Q7846231
Dicynodonts
Lopingian synapsids of Africa
Fossil taxa described in 1876
Lopingian genus first appearances
Lopingian genus extinctions
Anomodont genera