''Manubrantlia'' was a
genus
Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
lapillopsid from the
Early Triassic
The Early Triassic is the first of three epochs of the Triassic Period of the geologic timescale. It spans the time between Ma and Ma (million years ago). Rocks from this epoch are collectively known as the Lower Triassic Series, which is a un ...
Panchet Formation
Panchet is a census town in Nirsa CD block in Dhanbad Sadar subdivision of Dhanbad district in the Indian state of Jharkhand.
History
During construction of dam several ruin of Tilakampa kingdom were submerged in the dam. Telkupi was capital of ...
of
India
India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
. This genus is only known from a single
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 several ...
left jaw, given the designation
ISI
ISI or Isi may refer to:
Organizations
* Intercollegiate Studies Institute, a classical conservative organization focusing on college students
* Ice Skating Institute, a trade association for ice rinks
* Indian Standards Institute, former name of ...
A 57. Despite the paucity of remains, the jaw is still identifiable as belonging to a relative of ''
Lapillopsis
''Lapillopsis'' is an extinct genus of stereospondyl temnospondyl within the family Lapillopsidae. Fossils belonging to the genus have been found in the Arcadia Formation ( Rewan Group) of Queensland, Australia.
History of study
The type sp ...
''. For example, all three of its coronoid bones possessed teeth, the articular bone is partially visible in lateral (outer) view, and its postsplenial does not contact the posterior meckelian foramen. However, the jaw also possesses certain unique features which justify the erection of a new genus separate from ''Lapillopsis''. For example, the mandible is twice the size of any jaws referred to other lapillopsids. The most notable unique feature is an enlarged "pump-handle" shaped arcadian process at the back of the jaw. This structure is responsible for the
generic name of this genus, as "''Manubrantlia''" translates from
Latin
Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
to the
English
English usually refers to:
* English language
* English people
English may also refer to:
Peoples, culture, and language
* ''English'', an adjective for something of, from, or related to England
** English national ide ...
expression "pump-handle". The
type and only known
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 ...
of this genus is ''Manubrantlia khaki''. The
specific name Specific name may refer to:
* in Database management systems, a system-assigned name that is unique within a particular database
In taxonomy, either of these two meanings, each with its own set of rules:
* Specific name (botany), the two-part (bino ...
refers to the greenish-brown
mudstone
Mudstone, a type of mudrock, is a fine-grained sedimentary rock whose original constituents were clays or muds. Mudstone is distinguished from '' shale'' by its lack of fissility (parallel layering).Blatt, H., and R.J. Tracy, 1996, ''Petrology. ...
s of the Panchet Formation, with a color that had been described as "
khaki" by the first British geologists who studied the formation.
Lapillopsids such as ''Lapillopsis'' and ''Manubrantlia'' are part of
Temnospondyli, a diverse order of tetrapods which may include the ancestors of modern amphibians. Different authors disagree on the precise placement of Lapillopsids among temnospondyls. Some place them near or within
dissorophoids, a group of terrestrial insectivorous temnospondyls.
However, most other authors consider lapilopsids to be unusual
stereospondyls
The Stereospondyli are a group of extinct Temnospondyli, 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 ...
.
Stereospondyls are typically large and
crocodilia
Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest living ...
n-like
Mesozoic
The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...
temnospondyls with flattened skulls and semiaquatic habits, but lapillopsids differ from this body plan and more closely resemble the dissorophoids, at least superficially.
Description
The mandible is approximately 11.15 centimeters (4.4 inches) long, about twice the size of that of ''Lapillopsis''. This would have made ''Manubrantlia'' the largest lapillopsid by a wide margin. This has been construed as a primitive feature, as it falls more in line with the size of other temnospondyls rather than the derived minute size of more advanced lapillopsids.
Like other lapillopsids, ''Manubrantlia'' had two rows of teeth on each lower jaw. The teeth on the outer row, also known as "marginal teeth", are attached to the dentary bone and are sharp and curved. They are also slightly flattened in an anteroposterior (front-to-back) direction, although not to the extent seen in
capitosaurs
Capitosauria is an extinct group of large temnospondyl amphibians with simplified stereospondyl vertebrae. Mainly living as piscivores in lakes and rivers, the Capitosauria and its sister taxon Trematosauria were the only major labyrinthodonts t ...
such as ''
Mastodonsaurus
''Mastodonsaurus'' (meaning "teat tooth lizard") is an extinct genus of temnospondyl amphibian from the Middle Triassic of Europe. It belongs to a Triassic group of temnospondyls called Capitosauria, characterized by their large body size and pre ...
''. At the front of the jaw, the dentary tooth row also twists slightly outwards. The front of the jaw also possesses an enlarged tooth known as a parasymphyseal tusk, although this tooth is less than twice as wide as the rest of the marginal teeth. A deep longitudinal groove known as an oral sulcus is visible below the marginal tooth row.

The inner row of conical teeth form out of the three coronoid bones which lie inwards from the dentary. This feature is practically unique to lapillopsids among early Triassic temnospondyls, as most other groups have much more reduced coronoid teeth.
Lydekkerinids, for example, only have a patch of small denticles on the last coronoid. The only other stereospondyls with a continuous row of teeth on all three coronoid bones are the
plagiosaurids. The last coronoid also extends outwards, and is visible from the outer side of the mandible just above the rear tip of the dentary.
When seen in lateral or labial view (a.k.a. from the outside), a large portion of the mandible is formed by the
angular bone The angular is a large bone in the lower jaw (mandible) of amphibians and reptiles (birds included), which is connected to all other lower jaw bones: the dentary (which is the entire lower jaw in mammals), the splenial, the suprangular, and the art ...
(at the back), with a thinner portion under the dentary formed by the
splenial (at the front) and the postsplenial (at the middle). On the other hand, the splenial and postsplenial are much more exposed on the inner portion of the jaw (seen in lingual or medial view a.k.a. from the inside) while the angular is less so. Prominent ridges and grooves radiate from the middle of the angular bone (as seen from the outside of the jaw), similar to the condition in most temnospondyls but unlike other lapillopsids.
A large portion of the inner part of the jaw is also formed by the prearticular bone in the rear half of the mandible. A hole known as the posterior meckelian foramen is also visible from the inner part of the jaw. Unlike with most temnospondyls but in common with ''Lapillopsis'' (and a few other taxa, such as
archegosauroids and ''Eryops''), this hole has no contact with the postsplenial, instead being surrounded solely by the prearticular and angular.
The most prominent and unique features of ''Manubrantlia'' lie at the rear part of the jaw, in the general area of the jaw joint. The jaw joint is formed by the
articular bone, which is most visible in medial view but also wraps around to be visible in lateral view at the rear lower tip of the outer side of the bone. This lateral exposure of the articular is also known in ''Lapillopsis'', and is believed to be unique to lapillopsids. The upper rear portion of the lateral face of the mandible is formed by the
surangular bone. The rear tip of the surangular forms a bony projection known as an arcadian process. In most temnospondyls this projection is small and blunt, similar in size to the adjacent retroarticular process of the articular bone which is projects inwards and backwards. However, in ''Manubrantlia'' the arcadian process is enlarged greatly, turning into an elongated extension described as "resembling a pump-handle".
The arcadian process of ''Lapillopsis'' is also elongated, although it is much more slender as well.
The only other temnospondyls with arcadian processes near the size of ''Manubrantlia'' is the
rhytidosteid ''
Arcadia
Arcadia may refer to:
Places Australia
* Arcadia, New South Wales, a suburb of Sydney
* Arcadia, Queensland
* Arcadia, Victoria
Greece
* Arcadia (region), a region in the central Peloponnese
* Arcadia (regional unit), a modern administrative un ...
'' and lydekkerinids such as ''Lydekkerina''.
However, they also possess enlarged retroarticular processes, which contrasts with the condition in ''Manubrantlia''. Thus, the large size of the arcadian process is believed to be an example of convergent evolution between lapillopsids and lydekkerinids, as many other factors separate the two families.
Paleoecology
The fauna and flora of the Panchet Formation is practically identical to that of the
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 ...
n
''Lystrosaurus'' assemblage zone of the
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 perio ...
. This biozone is dated to the very beginning of the Triassic period about 250 million years ago, when life was still recovering from the severity of the
Permian-Triassic extinction event. It is assumed that the Panchet Formation, due to its similarities with the ''Lystrosaurus'' zone, had an equivalent age.
At this time the continents would have been formed into one supercontinent known as Pangaea. The
Indian tectonic plate was situated in the far southern hemisphere near South Africa at this point in time, as the plate tectonics which moved India towards its modern position in southern Asia had not yet activated. This proximity explains the similarity between the South African and Indian ecosystems.

The largest and most common animal within the Panchet Formation was the
herbivorous 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 ...
''
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 (a ...
.'' Several different species of ''Lystrosaurus'' have been reported from this formation in the past, but a 2005 analysis concluded that they were all referable to only a single species: ''Lystrosaurus murrayi.''
More recently, ''Lystrosaurus declivis'' and a species close to ''Lystrosaurus curvatus'' have also been found in lower portions of the Formation. A genus of
cynodont
The cynodonts () (clade Cynodontia) are a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 mya), and extensively diversified after the Permian–Triassic extinction event. Cynodonts had a wide variety ...
, ''
Panchetocynodon'', was also present in the formation. Fossils of ''
Thrinaxodon'', a cynodont common to the Karoo supergroup, have also been reported in the Panchet Formation, although these claims remain unsubstantiated. Other
amniote
Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are disti ...
remains are also known from the formation, including the dubious
proterosuchid ''
Ankistrodon
''Ankistrodon'' is an extinct genus of archosauriform known from the Early Triassic Panchet Formation of India. First thought to be a theropod dinosaur, it was later determined to be a proterosuchid. The type species is ''A. indicus'', described ...
'' (which may be synonymous with the South African ''
Proterosuchus'') and fragmentary
procolophonid fossils.
Apart from ''Lystrosaurus'', the most common animals in the Panchet Formation are a diverse assortment of stereospondyls, among them ''Manubrantlia''. These include ''
Indobenthosuchus
''Indobenthosuchus'' is an extinct genus of stereospondyl temnospondyl.
See also
* Prehistoric amphibian
* List of prehistoric amphibians
This list of prehistoric amphibians is an attempt to create a comprehensive listing of all genera from t ...
'' (a
lydekkerinid), ''
Indobrachyops
''Indobrachyops'' is an extinct genus of temnospondyl amphibian from the Early Triassic of India. It is known from a nearly complete fossil skull that was first described by paleontologists Friedrich von Huene and M. R. Sahni in 1958 from the Pan ...
'' (a
rhytidosteid), and various
trematosaurids
Trematosauridae are a family of large marine temnospondyl amphibians with many members. They first appeared during the Induan age of the Early Triassic, and existed until around the Carnian stage of the Late Triassic, although by then they were v ...
(''
Gonioglyptus
''Gonioglyptus'' is an extinct genus of trematosaurian temnospondyl within the family Trematosauridae. It is known from the Early Triassic Panchet Formation of India. It contains two species: ''G. longirostris'' (sometimes classified in the genu ...
'', ''
Panchetosaurus'', ''
Indolyrocephalus
''Indolyrocephalus'' is an extinct genus of prehistoric amphibian belonging to the family Trematosauridae. It contains a single species, ''I. huxleyi'', from the Early Triassic Panchet Formation of India. It was initially classified in ''Gonioglyp ...
,
Glyptognathus''). Many species remain poorly described and may qualify as ''
nomina dubia
In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application.
Zoology
In case of a ''nomen dubium'' it may be impossible to determine whether a s ...
'' ("dubious names").
Some stereospondyls from the formation, such as ''
Pachygonia'', are diagnostic yet too aberrant to be placed into a specific family. Fish scales are also known from the formation.The basal stereospondyl
Capulomala has also been found here.
Plant life includes remains of the
seed ferns
A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosperm pl ...
''
Glossopteris,
Lepidopteris
''Lepidopteris'' ("scaly fern") is a form genus for leaves of Late Permian to Late Triassic Period Pteridospermatophyta, or seed ferns, which lived from around 260 to 200 million years ago in what is now Australia, Antarctica, India, South Amer ...
,'' and ''
Dicroidium
''Dicroidium'' is an extinct genus of fork-leaved seed ferns that were widely distributed over Gondwana during the Triassic (). Their fossils are known from South Africa, the Arabian Peninsula, Australia, New Zealand, South America, Madagascar ...
''. However, no
coal
Coal is a combustible black or brownish-black sedimentary rock, formed as rock strata called coal seams. Coal is mostly carbon with variable amounts of other elements, chiefly hydrogen, sulfur, oxygen, and nitrogen.
Coal is formed when dea ...
seams are present in this formation unlike the case in many other Permian and Triassic formations of India. The abundance of
cryptogams (spore-bearing plants) suggests that India had a warm, humid climate during the early Triassic. However, many studies instead suggest that the climate was arid.
Geological analyses have indicated that the sandstones and mudstones of the Panchet Formation were mainly formed in a fluvial environment, meaning that the formation was dominated by large rivers during the time of ''Manubrantlia''. Some sand grains recovered from Panchet sandstones preserve the geological signatures of complex interactions between water and sand. These interactions include the formation of natural levees, tidal inlets, and beaches dominated by sand dunes. This indicates that large lakes were also present in the environment.
Earthquake
An earthquake (also known as a quake, tremor or temblor) is the shaking of the surface of the Earth resulting from a sudden release of energy in the Earth's lithosphere that creates seismic waves. Earthquakes can range in intensity, from ...
s also affected the area as shown by deformation in some sediments.
References
{{Taxonbar, from=Q55640996
Stereospondyls
Fossils of India
Triassic temnospondyls
Fossil taxa described in 2002