Pallisteria angustimentum
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''Mambawakale'' is a genus of large sized basal
paracrocodylomorph Paracrocodylomorpha is a clade of pseudosuchian archosaurs. The clade includes the diverse and unusual group Poposauroidea as well as the generally carnivorous and quadrupedal members of Loricata, including modern crocodylians. Paracrocodylomorp ...
from the
Manda Beds The Manda Formation (also known as the Manda Beds) is a Middle Triassic (Anisian?) or possibly Late Triassic (Carnian?) geologic formation in Tanzania. It preserves fossils of many terrestrial vertebrates from the Triassic, including some of the e ...
of
Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands ...
. It was informally named ''Pallisteria'' before being officially published under its current name almost 60 years after its discovery. It contains a single species, ''Mambawakale ruhuhu''.


History and naming

In 1963, following the independence of Tanzania, Alan Charig participated in a joint expedition of the
Natural History Museum, London The Natural History Museum in London is a museum that exhibits a vast range of specimens from various segments of natural history. It is one of three major museums on Exhibition Road in South Kensington, the others being the Science Museum an ...
and the
University of London The University of London (UoL; abbreviated as Lond or more rarely Londin in post-nominals) is a federal public research university located in London, England, United Kingdom. The university was established by royal charter in 1836 as a degree ...
, as well as researchers from
Uganda }), is a landlocked country in East Africa. The country is bordered to the east by Kenya, to the north by South Sudan, to the west by the Democratic Republic of the Congo, to the south-west by Rwanda, and to the south by Tanzania. The sou ...
,
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. It is bounded to the south by of coastline that stretch along the Atlantic Ocean, South Atlantic and Indian Oceans; to the ...
and
Edinburgh Edinburgh ( ; gd, Dùn Èideann ) is the capital city of Scotland and one of its 32 council areas. Historically part of the county of Midlothian (interchangeably Edinburghshire before 1921), it is located in Lothian on the southern shore of t ...
, to Tanzania and
Zambia Zambia (), officially the Republic of Zambia, is a landlocked country at the crossroads of Central, Southern and East Africa, although it is typically referred to as being in Southern Africa at its most central point. Its neighbours are t ...
. The expedition heavily relied on the support of locals who had discovered the localities and fossils within them, although they went unnamed in the field notes. The fossils discovered in 1963 were collected and stored in the Natural History Museum of London. Among these fossils was the incomplete skull of a large crocodylomorph, noted for its large size and informally referred to as ''Pallisteria angustimentum'' (after Charig's friend John Weaver Pallister and the Latin words "angustus" and "mentum", meaning "narrow chin). Little information was given on ''Pallisteria'', with neither details, figures or even a specimen number being noted down. The manuscript, although listed as "in press", was never published nor recovered from the archives, rendering ''Pallisteria'' (and the family Pallisteriidae) a '' nomen nudum''. Following the formal description of '' Teleocrater'', ''
Mandasuchus ''Mandasuchus'' is an extinct genus of loricatan pseudosuchian from the Manda Formation of Tanzania, which dates back to the Anisian stage of the Middle Triassic. Although this genus was first mentioned by Alan Charig in 1956,Charig, A. J. (1956 ...
'', ''
Hypselorhachis ''Hypselorhachis'' is a genus of extinct reptile, possibly a ctenosauriscid archosaur related to ''Ctenosauriscus''. It lived during the Triassic Period. It is currently known only from a single vertebra found from the Middle Triassic Manda Bed ...
'' and ''
Nyasasaurus ''Nyasasaurus'' (meaning " Lake Nyasa lizard") is an extinct genus of avemetatarsalian archosaur from the putatively Middle Triassic Manda Formation of Tanzania that may be the earliest known dinosaur. The type species ''Nyasasaurus parringtoni' ...
'', ''Pallisteria'' was the last of the significant fossil archosaurs reported by Charig to be formally published. A formal description of the holotype specimen (NHMUK R36620) was eventually published 59 years later in 2022 by Richard J. Butler and colleagues, who named it ''Mambawakale''. In addition to the skull, Butler's team also described associated postcranial material mentioned in the field notes, including elements of the cervical series and a left manus. The name ''Mambawakale'' is a composite of the
Kiswahili Swahili, also known by its local name , is the native language of the Swahili people, who are found primarily in Tanzania, Kenya and Mozambique (along the East African coast and adjacent litoral islands). It is a Bantu language, though Swahili ...
words "mamba" (crocodile) and "wakale" (ancient). The species name refers to the Ruhuhu Basin where the fossils have been found.


Description

Based on a skull length of 75 cm, ''Mambawakale'' would have been a large sized
Pseudosuchian Pseudosuchia is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely related to crocodilians than to birds. Pseudosuchians are also informally known as "crocodilian-line archosaurs". Prior to ...
of unknown age. Although only the lower edge of the external nares is preserved, it's extent over most of the
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...
and the front most part of the
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. T ...
suggests they were large in size, comparable to ''
Batrachotomus ''Batrachotomus'' is a genus of prehistoric archosaur. Fossils of this animal have been found in southern Germany and dated from the Ladinian stage of the Middle Triassic period, around 242 to 237 million years ago. ''Batrachotomus'' was descr ...
''. The contact between the premaxilla and maxilla is marked by a slight constriction visible when viewing the skull from below and is located within a short toothless section of the jaw (
diastema A diastema (plural diastemata, from Greek διάστημα, space) is a space or gap between two teeth. Many species of mammals have diastemata as a normal feature, most commonly between the incisors and molars. More colloquially, the condition ...
). The surface of the
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly separ ...
shows a series of pits that likely received the teeth of the mandible when the jaw was closed. Each maxilla contains four teeth, the last of which being notably larger than the first three and more similar to those of the maxilla. This notable heterodonty of the premaxillary teeth is described as an
autapomorphy In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to ...
of ''Mambawakale'' and differs greatly from most other archosauriforms. The surface of the maxilla, although badly preserved, shows no signs of rugosity and appears smooth. Generally, the maxilla share several characteristics with both
Saurosuchus ''Saurosuchus'' (meaning "lizard crocodile") is an extinct genus of large loricatan pseudosuchian archosaur that lived in South America during the Late Triassic period. It was a heavy, ground-dwelling, quadrupedal carnivore, likely being the ape ...
and
Prestosuchus ''Prestosuchus'' (meaning "Prestes crocodile") is an extinct genus of pseudosuchian in the group Loricata, which also includes '' Saurosuchus'' and '' Postosuchus''. It has historically been referred to as a " rauisuchian", and was the defining ...
. In overall shape the maxilla are straight, diverging from one another as they move away from the premaxilla. The maxilla appear to contain ten teeth each showing little variation in size unlike those of the premaxilla. There is however a slight reduction over the last 4 teeth. The hemimandibles of the lower jaw form an elongated symphisys, which gives the chin a narrow appearance as noted by Charig when naming ''P. angustimentum''. Although at one point considered a derived trait of ornithosuchids, this morphology can also be observed in other groups such as
crocodyliforms Crocodyliformes is a clade of crurotarsan archosaurs, the group often traditionally referred to as "crocodilians". They are the first members of Crocodylomorpha to possess many of the features that define later relatives. They are the only pseudo ...
,
poposauroids Poposauroidea is a clade of advanced pseudosuchians ( archosaurs closer to crocodilians than to dinosaurs). It includes poposaurids, shuvosaurids, ctenosauriscids, and other unusual pseudosuchians such as ''Qianosuchus'' and '' Lotosaurus''. Ho ...
and
phytosaurs Phytosaurs (Φυτόσαυροι in greek) are an extinct group of large, mostly semiaquatic Late Triassic archosauriform reptiles. Phytosaurs belong to the order Phytosauria. Phytosauria and Phytosauridae are often considered to be equivalen ...
. The shallow dentary contains fifteen to sixteen ziphodont teeth. The centrum of the
axis An axis (plural ''axes'') is an imaginary line around which an object rotates or is symmetrical. Axis may also refer to: Mathematics * Axis of rotation: see rotation around a fixed axis * Axis (mathematics), a designator for a Cartesian-coordinat ...
in ''Mambawakale'' is similar to that of ''
Stagonosuchus ''Stagonosuchus'' is an extinct genus of loricatan, or possibly a species of ''Prestosuchus''. Fossils have been found from the Late Triassic Manda Formation in Tanzania that are Anisian in age. Unlike other rauisuchians, which have a prominen ...
'', making it a possibility that the two taxa may represent the same species. However, the absence of well preserved postcranial material in ''Mambawakale'' and well preserved cranial material in ''Stagonosuchus'' makes it impossible to make a definitive statement on the matter. ''Mambawakale'' is easily distinguished from both '' Parringtonia'' and ''
Nundasuchus ''Nundasuchus'' is an extinct genus of crurotarsan, possibly a suchian archosaur related to Paracrocodylomorpha. Remains of this genus are known from the Middle Triassic Manda beds of southwestern Tanzania. It contains a single species, ''Nun ...
'' and likely does not represent ''
Hypselorhachis ''Hypselorhachis'' is a genus of extinct reptile, possibly a ctenosauriscid archosaur related to ''Ctenosauriscus''. It lived during the Triassic Period. It is currently known only from a single vertebra found from the Middle Triassic Manda Bed ...
'' or ''
Mandasuchus ''Mandasuchus'' is an extinct genus of loricatan pseudosuchian from the Manda Formation of Tanzania, which dates back to the Anisian stage of the Middle Triassic. Although this genus was first mentioned by Alan Charig in 1956,Charig, A. J. (1956 ...
'' either. Based on comparison with the South American ''Prestosuchus'', ''Mambawakale'' may have had a skull length of up to 75 cm and would have been one of the largest Pseudosuchians of the middle Triassic, comparable in size with '' Etjosuchus''.


Phylogeny

The strict consensus tree recovered ''Mambawakale'' as a basal paracrocodylomorph in a polytomy with two other Manda Bed Pseudosuchians,
Poposauroidea Poposauroidea is a clade of advanced pseudosuchians (archosaurs closer to crocodilians than to dinosaurs). It includes poposaurids, shuvosaurids, ctenosauriscids, and other unusual pseudosuchians such as ''Qianosuchus'' and ''Lotosaurus''. How ...
and
Loricata Loricata is a clade of archosaur reptiles that includes crocodilians and some of their Triassic relatives, such as '' Postosuchus'' and ''Prestosuchus''. More specifically, Loricata includes Crocodylomorpha (the persistent archosaur subset which ...
. The inclusion of ''
Nundasuchus ''Nundasuchus'' is an extinct genus of crurotarsan, possibly a suchian archosaur related to Paracrocodylomorpha. Remains of this genus are known from the Middle Triassic Manda beds of southwestern Tanzania. It contains a single species, ''Nun ...
'' did not affect this placement in the phylogenetic tree. However, overall its position within Archosauria is only weakly resolved and poorly supported. It can be clearly identified as an archosaur, however its placement in Pseudosuchia is less certain, as many traits that would nest it in the group are more broadly found across other archosaur clades.


References

{{Taxonbar, from=Q110895907 Paracrocodylomorphs Middle Triassic reptiles of Africa Anisian life Triassic Tanzania Fossils of Tanzania Fossil taxa described in 2022 Prehistoric pseudosuchian genera