Rhineceps
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''Rhineceps'' is an extinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order (biology), order of small to giant tetrapods—often considered Labyrinth ...
amphibian in the family
Rhinesuchidae Rhinesuchidae is a family (biology), family of Tetrapoda, tetrapods that lived primarily in the Permian period. They belonged to the broad group Temnospondyli, a successful and diverse collection of semiaquatic tetrapods which modern amphibians ...
. ''Rhineceps'' was found in Northern
Malawi Malawi, officially the Republic of Malawi, is a landlocked country in Southeastern Africa. It is bordered by Zambia to the west, Tanzania to the north and northeast, and Mozambique to the east, south, and southwest. Malawi spans over and ...
(formerly Nyasaland) in
Southern Africa Southern Africa is the southernmost region of Africa. No definition is agreed upon, but some groupings include the United Nations geoscheme for Africa, United Nations geoscheme, the intergovernmental Southern African Development Community, and ...
known only from its
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
''R. nyasaensis''. ''Rhineceps'' was a late
Permian The Permian ( ) is a geologic period and System (stratigraphy), 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 251.902 Mya. It is the s ...
(256-258 Mya) semi-aquatic carnivore that lived in streams, rivers, lakes or lagoons. ''Rhineceps'' is an early divergent Stereopondyl within the family
Rhinesuchidae Rhinesuchidae is a family (biology), family of Tetrapoda, tetrapods that lived primarily in the Permian period. They belonged to the broad group Temnospondyli, a successful and diverse collection of semiaquatic tetrapods which modern amphibians ...
, which only existed in the late
Permian The Permian ( ) is a geologic period and System (stratigraphy), 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 251.902 Mya. It is the s ...
(Lopingian) and failed to survive the Permian-Triassic extinction unlike other
stereospondyl The Stereospondyli are a group of extinct temnospondyl amphibians that existed primarily during the Mesozoic period. They are known from all seven continents and were common components of many Triassic ecosystems, likely filling a similar ecologi ...
families.


History

''Rhineceps'' was originally discovered in the “Upper Bone Beds” of the Chiweta Beds, North Waller Area in Northern
Malawi Malawi, officially the Republic of Malawi, is a landlocked country in Southeastern Africa. It is bordered by Zambia to the west, Tanzania to the north and northeast, and Mozambique to the east, south, and southwest. Malawi spans over and ...
. The fossil was first described and named by Haughton in 1926, whereby he described the fossil of a left mandibular ramus of ''R. nyasaensis'' as within the
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
''Rhinesuchus'' due to its similarity with the fossils of ''
Rhinesuchus ''Rhinesuchus'' (meaning "rasp crocodile" for the ridged surface texture on its skull bones) is a large temnospondyl. Remains of the genus are known from the Permian of the South African Karoo Basin's ''Tapinocephalus'' and ''Cistecephalus'' as ...
whaitsi.'' Watson later determined in a letter to Parrington that the original fossil was actually the anterior end of a right ramus and not a left ramus. Watson in 1962 described fossils newly found from the same locality collected by Parrington as fossils of ''R. nyasaensis'' and determined they were from the same individual.  Watson also placed ''R. nyasaensis'' as part of a new
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
''Rhineceps'' that is distinct from other ''Rhinesuchus'' species.


Classification

''Rhineceps'' belongs to the family
Rhinesuchidae Rhinesuchidae is a family (biology), family of Tetrapoda, tetrapods that lived primarily in the Permian period. They belonged to the broad group Temnospondyli, a successful and diverse collection of semiaquatic tetrapods which modern amphibians ...
, which is within the clade
Stereospondyli The Stereospondyli are a group of extinct temnospondyl amphibians that existed primarily during the Mesozoic period. They are known from all seven continents and were common components of many Triassic ecosystems, likely filling a similar ecologi ...
(a
Temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order (biology), order of small to giant tetrapods—often considered Labyrinth ...
clade). Until recently, both
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
, historical placement and
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 ...
analysis has placed ''Rhineceps'' as a
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to ''
Rhinesuchus ''Rhinesuchus'' (meaning "rasp crocodile" for the ridged surface texture on its skull bones) is a large temnospondyl. Remains of the genus are known from the Permian of the South African Karoo Basin's ''Tapinocephalus'' and ''Cistecephalus'' as ...
'' within Rhinesuchinae, an early diverging group of Rhinesuchidae. In 2017, Marsicano et al. created a new
phylogeny A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or Taxon, taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, M ...
that places ''Rhineceps'' as a
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to ''
Uranocentrodon ''Uranocentrodon'' is an extinct genus of temnospondyls in the family Rhinesuchidae. Known from a skull, ''Uranocentrodon'' was a large predator with a length up to . Originally named ''Myriodon'' by van Hoepen in 1911, it was transferred to a ...
'' rather than within Rhinesuchinae (which is no longer a
monophyletic group In biology, a clade (), also known as a monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach to taxonomy ...
). This discrepancy in clearly describing
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 ...
relationships is likely due to the
adaptive radiation In evolutionary biology, adaptive radiation is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic int ...
of
stereospondylomorphs Stereospondylomorpha is a clade of temnospondyls. It includes the superfamily Archegosauroidea and the more diverse group Stereospondyli. Stereospondylomorpha was first proposed by Yates and Warren (2000), who found Archegosauroidea and Stereosp ...
that took place during the late
Permian The Permian ( ) is a geologic period and System (stratigraphy), 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 251.902 Mya. It is the s ...
and
early Triassic The Early Triassic is the first of three epochs of the Triassic Period of the geologic timescale. It spans the time between 251.9 Ma and Ma (million years ago). Rocks from this epoch are collectively known as the Lower Triassic Series, which ...
. Terrestrial
temnospondyls Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order of small to giant tetrapods—often considered primitive amphibians—that flourished wo ...
were devastated during this period, caused by selective pressures for semi-aquatic or fully aquatic
temnospondyls Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order of small to giant tetrapods—often considered primitive amphibians—that flourished wo ...
to acquire many new niches in a relatively short time period.


Description and paleobiology

''Rhineceps'' was a medium-sized, semi-aquatic, carnivorous, basal
stereospondyl The Stereospondyli are a group of extinct temnospondyl amphibians that existed primarily during the Mesozoic period. They are known from all seven continents and were common components of many Triassic ecosystems, likely filling a similar ecologi ...
of the
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 te ...
of Malawi. They lived in fresh water bodies, including rivers lakes and lagoons, where they had diets mainly of fish.
Stereospondyls The Stereospondyli are a group of extinct temnospondyl amphibians that existed primarily during the Mesozoic period. They are known from all seven continents and were common components of many Triassic ecosystems, likely filling a similar ecologi ...
were historically compared with contemporary
crocodilia Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
ns and
giant salamander The Cryptobranchidae (commonly known as giant salamanders) are a family of large salamanders that are fully aquatic. The family includes some of the largest living amphibians. They are native to China, Japan, and the eastern United States. Giant ...
s due to their similar position as freshwater predators. However, they likely existed in a niche that is unique and independent from
crocodilia Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
ns and
giant salamander The Cryptobranchidae (commonly known as giant salamanders) are a family of large salamanders that are fully aquatic. The family includes some of the largest living amphibians. They are native to China, Japan, and the eastern United States. Giant ...
s. These
niche Niche may refer to: Science *Developmental niche, a concept for understanding the cultural context of child development and growth *Ecological niche, a term describing the relational position of an organism's species *Niche differentiation, in ec ...
differences are inferred from cranial stress and structural patterns along with
labyrinthodont "Labyrinthodontia" (Greek, 'maze-toothed') is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras (about 390 to 150 million years ago). Traditionally conside ...
dentition ( labyrinthodonty refers to infolding of enamel and
dentin Dentin ( ) (American English) or dentine ( or ) (British English) () is a calcified tissue (biology), tissue of the body and, along with tooth enamel, enamel, cementum, and pulp (tooth), pulp, is one of the four major components of teeth. It i ...
e within a tooth). The
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
of ''Rhineceps'' thus likely restricts its ability to hunt in specific ways that
crocodilia Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
ns and
giant salamander The Cryptobranchidae (commonly known as giant salamanders) are a family of large salamanders that are fully aquatic. The family includes some of the largest living amphibians. They are native to China, Japan, and the eastern United States. Giant ...
s such as sweeping sideways bites. ''Rhineceps'' also used its unique
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
in service of catching prey, including its strong powerful bite and its palatal tooth row to grasp and hold fish in its jaw. Many
temnospondyls Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order of small to giant tetrapods—often considered primitive amphibians—that flourished wo ...
including ''Rhineceps'' likely shared certain
physiological Physiology (; ) is the science, scientific study of function (biology), functions and mechanism (biology), mechanisms in a life, living system. As a branches of science, subdiscipline of biology, physiology focuses on how organisms, organ syst ...
characteristics with modern amphibians that cannot be directly observed in the fossil record. ''Rhineceps'' likely possessed heightened
phenotypic plasticity Phenotypic plasticity refers to some of the changes in an organism's behavior, morphology and physiology in response to a unique environment. Fundamental to the way in which organisms cope with environmental variation, phenotypic plasticity encompa ...
among juvenile individuals like seen in modern
amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s. This
plasticity Plasticity may refer to: Science * Plasticity (physics), in engineering and physics, the propensity of a solid material to undergo permanent deformation under load * Behavioral plasticity, change in an organism's behavior in response to exposur ...
allowed
physiological Physiology (; ) is the science, scientific study of function (biology), functions and mechanism (biology), mechanisms in a life, living system. As a branches of science, subdiscipline of biology, physiology focuses on how organisms, organ syst ...
changes during an organism's lifetime to alter its
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
during development to specific and different environmental factors experienced.
Plasticity Plasticity may refer to: Science * Plasticity (physics), in engineering and physics, the propensity of a solid material to undergo permanent deformation under load * Behavioral plasticity, change in an organism's behavior in response to exposur ...
was a means for
stereospondyls The Stereospondyli are a group of extinct temnospondyl amphibians that existed primarily during the Mesozoic period. They are known from all seven continents and were common components of many Triassic ecosystems, likely filling a similar ecologi ...
to adaptively radiate to many different
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
and niches. ''Rhineceps'' like other
temnospondyls Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order of small to giant tetrapods—often considered primitive amphibians—that flourished wo ...
were ecological
indicator species A bioindicator is any species (an indicator species) or group of species whose function, population, or status can reveal the qualitative status of the environment. The most common indicator species are animals. For example, copepods and other sma ...
like modern
amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s, and thus likely were sensitive to
chemical A chemical substance is a unique form of matter with constant chemical composition and characteristic properties. Chemical substances may take the form of a single element or chemical compounds. If two or more chemical substances can be combin ...
changes,
UV radiation Ultraviolet radiation, also known as simply UV, is electromagnetic radiation of wavelengths of 10–400 nanometers, shorter than that of visible light, but longer than X-rays. UV radiation is present in sunlight and constitutes about 10% of t ...
and other
mutagen In genetics, a mutagen is a physical or chemical agent that permanently changes genetic material, usually DNA, in an organism and thus increases the frequency of mutations above the natural background level. As many mutations can cause cancer in ...
s within its
ecosystem An ecosystem (or ecological system) is a system formed by Organism, organisms in interaction with their Biophysical environment, environment. The Biotic material, biotic and abiotic components are linked together through nutrient cycles and en ...
. ''Rhineceps'' like modern
amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s had obligatory aquatic young. ''Rhineceps'' like other
stereospondyls The Stereospondyli are a group of extinct temnospondyl amphibians that existed primarily during the Mesozoic period. They are known from all seven continents and were common components of many Triassic ecosystems, likely filling a similar ecologi ...
was the result of
evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
ary gigantism which resulted in stunted
metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth transformation or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and different ...
with few morphological differences between adults and larvae.
Metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth transformation or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and different ...
was likely lost in
stereospondyls The Stereospondyli are a group of extinct temnospondyl amphibians that existed primarily during the Mesozoic period. They are known from all seven continents and were common components of many Triassic ecosystems, likely filling a similar ecologi ...
because extreme
phenotypic plasticity Phenotypic plasticity refers to some of the changes in an organism's behavior, morphology and physiology in response to a unique environment. Fundamental to the way in which organisms cope with environmental variation, phenotypic plasticity encompa ...
allowed environmental acclimation through changes in
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
without needing a binary developmental life history with larval and adult stages. This eventually led to
neoteny Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the Physiology, physiological, or Somatic (biology), somatic, development of an organism, typically an animal. Neoteny i ...
whereby the larval stage is morphologically very similar to the adult stage.
Stereospondyls The Stereospondyli are a group of extinct temnospondyl amphibians that existed primarily during the Mesozoic period. They are known from all seven continents and were common components of many Triassic ecosystems, likely filling a similar ecologi ...
including ''Rhineceps'' only had gradual changes between juvenile and
adult An adult is an animal that has reached full growth. The biological definition of the word means an animal reaching sexual maturity and thus capable of reproduction. In the human context, the term ''adult'' has meanings associated with social an ...
stages, and their fossils do not suggest the presence of any type of
metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth transformation or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and different ...
, major or minor. Greater ambiguity is created because
stereospondyls The Stereospondyli are a group of extinct temnospondyl amphibians that existed primarily during the Mesozoic period. They are known from all seven continents and were common components of many Triassic ecosystems, likely filling a similar ecologi ...
including ''Rhineceps'' have traits traditionally associated with
metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth transformation or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and different ...
, including amphibious
dermal The dermis or corium is a layer of skin between the epidermis (with which it makes up the cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from stress and strain. It is divided i ...
pitting. The presence of the types of
gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow r ...
s in
temnospondyls Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order of small to giant tetrapods—often considered primitive amphibians—that flourished wo ...
has largely been difficult to determine because conflicting interpretation of
osteological Osteology () is the scientific study of bones, practiced by osteologists . A subdiscipline of anatomy, anthropology, archaeology and paleontology, osteology is the detailed study of the structure of bones, skeletal elements, teeth, microbone morp ...
evidence.
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 ...
information has also created uncertainty of where
external gills External gills are the gills of an animal, most typically an amphibian, that are exposed to the environment, rather than set inside the pharynx and covered by gill slits, as they are in most fishes. Instead, the respiratory organs are set on a fri ...
evolved and internal
gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow r ...
s were lost. The evolutionary explanation for this contradiction known as Bystrow's Paradox is that
temnospondyls Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order of small to giant tetrapods—often considered primitive amphibians—that flourished wo ...
had both internal
gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow r ...
s and
external gills External gills are the gills of an animal, most typically an amphibian, that are exposed to the environment, rather than set inside the pharynx and covered by gill slits, as they are in most fishes. Instead, the respiratory organs are set on a fri ...
, with external gills only present in larvae and internal gills that are homologous with
Osteichthyes Osteichthyes ( ; ), also known as osteichthyans or commonly referred to as the bony fish, is a Biodiversity, diverse clade of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondricht ...
(bony fish) gills that later become modern
amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
gills. Thus, ''Rhineceps'' larvae likely had external gills and adult ''Rhineceps'' had internal gills, although this is difficult to confirm given the lack of evidence for
metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth transformation or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and different ...
in
stereospondyls The Stereospondyli are a group of extinct temnospondyl amphibians that existed primarily during the Mesozoic period. They are known from all seven continents and were common components of many Triassic ecosystems, likely filling a similar ecologi ...
including ''Rhineceps''. ''Rhineceps'' as a
temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order (biology), order of small to giant tetrapods—often considered Labyrinth ...
, possesses
diagnostic Diagnosis (: diagnoses) is the identification of the nature and cause of a certain phenomenon. Diagnosis is used in a lot of different academic discipline, disciplines, with variations in the use of logic, analytics, and experience, to determine " ...
temnospondyl traits including wide vomers, large and round interpterygoid vacuities, otic notches, contact between post-pareital and exoccipital skull bones, and a stapes that articulates with the parasphenoid.Rainer R. Schoch. 2013. The evolution of major temnospondyl clades: an inclusive phylogenetic analysis, Journal of Systematic Palaeontology, 11:6, 673-705, DOI: 10.1080/14772019.2012.699006 ''Rhineceps'' fossils are differentiated from other rhinesuchids by the following traits “presence of a vomerine depression immediately anterior to cultriform process of the parasphenoid; ectopterygoids with enlarged tusks at their anterior end; transverse vomerine tooth row anteriorly convex; quadrate condyles projected behind the tip of the tabular horns; vomers with a continuous raised field of denticles; parasphenoid plate wider than long; well-developed transversely wide ‘pockets’; internarial vacuity between nasals and premaxillae; mandible with two anterior meckelian foraminae; chordatympanic foramen located on the suture between the articular and the prearticular.”.


Geological and paleoenvironmental information

''Rhineceps'' was found in the South African
Karoo Supergroup 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 per ...
, a
group A group is a number of persons or things that are located, gathered, or classed together. Groups of people * Cultural group, a group whose members share the same cultural identity * Ethnic group, a group whose members share the same ethnic iden ...
of
sedimentary rock Sedimentary rocks are types of rock (geology), rock formed by the cementation (geology), cementation of sediments—i.e. particles made of minerals (geological detritus) or organic matter (biological detritus)—that have been accumulated or de ...
s that span from the
Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
period to the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
period. These
sedimentary Sedimentary rocks are types of rock formed by the cementation of sediments—i.e. particles made of minerals (geological detritus) or organic matter (biological detritus)—that have been accumulated or deposited at Earth's surface. Sedime ...
layers were created from
sediment Sediment is a solid material that is transported to a new location where it is deposited. It occurs naturally and, through the processes of weathering and erosion, is broken down and subsequently sediment transport, transported by the action of ...
shed from lakes and rivers near the Gondwanide mountains on the
continent A continent is any of several large geographical regions. Continents are generally identified by convention (norm), convention rather than any strict criteria. A continent could be a single large landmass, a part of a very large landmass, as ...
of
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
. ''Rhineceps'' was found in the “Upper Bone Beds” of the Chiweta beds in the North Waller Area in Northern
Malawi Malawi, officially the Republic of Malawi, is a landlocked country in Southeastern Africa. It is bordered by Zambia to the west, Tanzania to the north and northeast, and Mozambique to the east, south, and southwest. Malawi spans over and ...
, which is in South
Central Africa Central Africa (French language, French: ''Afrique centrale''; Spanish language, Spanish: ''África central''; Portuguese language, Portuguese: ''África Central'') is a subregion of the African continent comprising various countries accordin ...
. The
sedimentary rock Sedimentary rocks are types of rock (geology), rock formed by the cementation (geology), cementation of sediments—i.e. particles made of minerals (geological detritus) or organic matter (biological detritus)—that have been accumulated or de ...
s from the Chiweta beds originated from mostly river sediments which often contain fossil bones surrounded by pedogenic carbonates. The lacustrine (lake derived) sediment does not contain bone, but instead contains pollen that were used to identify the age range of the Chiweta beds to be less than 263 Mya. The overlap of
stratigraphic Stratigraphy is a branch of geology concerned with the study of rock layers (strata) and layering (stratification). It is primarily used in the study of sedimentary and layered volcanic rocks. Stratigraphy has three related subfields: lithost ...
ranges of specific therapsid
taxa In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
place the
Cistecephalus Assemblage Zone The ''Cistecephalus'' Assemblage Zone is a tetrapod assemblage zone or biozone found in the Adelaide Subgroup of the Beaufort Group, a majorly fossiliferous and geologically important geological group of the Karoo Supergroup in South Africa. T ...
in the Chiweta beds (the zone where ''Rhineceps'' was found) within 256-258 Mya. ''Rhineceps'' lived during the late
Permian The Permian ( ) is a geologic period and System (stratigraphy), 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 251.902 Mya. It is the s ...
of northern
Malawi Malawi, officially the Republic of Malawi, is a landlocked country in Southeastern Africa. It is bordered by Zambia to the west, Tanzania to the north and northeast, and Mozambique to the east, south, and southwest. Malawi spans over and ...
256-258 Mya. The
ecosystem An ecosystem (or ecological system) is a system formed by Organism, organisms in interaction with their Biophysical environment, environment. The Biotic material, biotic and abiotic components are linked together through nutrient cycles and en ...
s of
Southern Africa Southern Africa is the southernmost region of Africa. No definition is agreed upon, but some groupings include the United Nations geoscheme for Africa, United Nations geoscheme, the intergovernmental Southern African Development Community, and ...
were mainly tropical and were dominated by many rivers and river systems. Terrestrial ecosystems are dominated by Therapsids and
river ecosystem River ecosystems are flowing waters that drain the landscape, and include the biotic (living) interactions amongst plants, animals and micro-organisms, as well as abiotic (nonliving) physical and chemical interactions of its many parts.Angelier, ...
s are dominated by fishes and aquatic carnivorous temnospondyls. The late
Permian The Permian ( ) is a geologic period and System (stratigraphy), 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 251.902 Mya. It is the s ...
is also characterized by reductions in
biodiversity Biodiversity is the variability of life, life on Earth. It can be measured on various levels. There is for example genetic variability, species diversity, ecosystem diversity and Phylogenetics, phylogenetic diversity. Diversity is not distribut ...
that limited the number of existing
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
and
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
. Most groups of
temnospondyls Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order of small to giant tetrapods—often considered primitive amphibians—that flourished wo ...
that existed terrestrially went
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
and consequently
stereospondyls The Stereospondyli are a group of extinct temnospondyl amphibians that existed primarily during the Mesozoic period. They are known from all seven continents and were common components of many Triassic ecosystems, likely filling a similar ecologi ...
were the only
temnospondyls Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order of small to giant tetrapods—often considered primitive amphibians—that flourished wo ...
to survive into the late
Permian The Permian ( ) is a geologic period and System (stratigraphy), 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 251.902 Mya. It is the s ...
and early
Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
by adapting to a mostly or fully aquatic lifestyle. The
ecosystem An ecosystem (or ecological system) is a system formed by Organism, organisms in interaction with their Biophysical environment, environment. The Biotic material, biotic and abiotic components are linked together through nutrient cycles and en ...
s that ''Rhineceps'' lived in experienced significant
ecological Ecology () is the natural science of the relationships among living organisms and their environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere levels. Ecology overlaps with the closely re ...
change causing dwindling
biodiversity Biodiversity is the variability of life, life on Earth. It can be measured on various levels. There is for example genetic variability, species diversity, ecosystem diversity and Phylogenetics, phylogenetic diversity. Diversity is not distribut ...
before the traditional date for the Permian-Triassic mass extinction event (252 Mya), likely caused by
global cooling Global cooling was a conjecture, especially during the 1970s, of imminent cooling of the Earth culminating in a period of extensive glaciation, due to the cooling effects of aerosols or orbital forcing. Some press reports in the 1970s specu ...
and growing
glacier A glacier (; or ) is a persistent body of dense ice, a form of rock, that is constantly moving downhill under its own weight. A glacier forms where the accumulation of snow exceeds its ablation over many years, often centuries. It acquires ...
s at the end of the
Guadalupian The Guadalupian is the second and middle Series (stratigraphy), series/Epoch (geology), 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 an ...
(end of middle Permian).


See also

*
List of prehistoric amphibians This list of prehistoric amphibians is an attempt to create a comprehensive listing of all genera from the fossil record that have ever been considered to be amphibians, excluding purely vernacular terms. The list includes all commonly accepted gen ...


References

{{Taxonbar, from=Q7320560 Stereospondyli Permian temnospondyls Lopingian genera Lopingian amphibians of Africa Permian Malawi Fossils of Malawi Fossil taxa described in 1962 Monotypic prehistoric amphibian genera