Mandasuchus
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''Mandasuchus'' is an extinct genus of
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 ...
n
pseudosuchia Pseudosuchia, from Ancient Greek ψεύδος (''pseúdos)'', meaning "false", and σούχος (''soúkhos''), meaning "crocodile" is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely relat ...
n from the
Manda Formation The Manda Formation (also known as the Manda Beds) is a Middle Triassic (Anisian?) or possibly Late Triassic (Carnian?) Formation (stratigraphy), geologic formation in Tanzania. It preserves fossils of many terrestrial vertebrates from the Triassi ...
of
Tanzania Tanzania, officially the United Republic of Tanzania, is a country in East Africa within the African Great Lakes region. It is bordered by Uganda to the northwest; Kenya to the northeast; the Indian Ocean to the east; Mozambique and Malawi to t ...
, which dates back to the
Anisian In the geologic timescale, the Anisian is the lower stage (stratigraphy), stage or earliest geologic age, age of the Middle Triassic series (stratigraphy), series or geologic epoch, epoch and lasted from million years ago until million years ag ...
stage of the
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epoch (geology), epochs of the Triassic period (geology), period or the middle of three series (stratigraphy), series in which the Triassic system (stratigraphy), system is di ...
. Although this genus was first mentioned by Alan Charig in 1956,Charig, A. J. (1956). New Triassic archosaurs from Tanganyika, including ''Mandasuchus'' and ''Teleocrater'': Dissertation Abstracts. Cambridge University. a formal description was not published until 2018.


History

The name was first used in a 1956 doctoral dissertation by Alan J. Charig of the
University of Cambridge The University of Cambridge is a Public university, public collegiate university, collegiate research university in Cambridge, England. Founded in 1209, the University of Cambridge is the List of oldest universities in continuous operation, wo ...
, along with '' Teleocrater'', an archosaur formally named in 2017. Several well preserved specimens have been found, although there is little cranial material. The family
Prestosuchidae Prestosuchidae (in its widest usage) is a polyphyletic grouping of carnivorous archosaurs that lived during the Triassic. They were large active terrestrial apex predators, ranging from around in length. They succeeded the Erythrosuchidae as th ...
was erected in 1967 by
Alfred Romer Alfred Sherwood Romer (December 28, 1894 – November 5, 1973) was an American paleontologist and biologist and a specialist in vertebrate evolution. Biography Alfred Romer was born in White Plains, New York, the son of Harry Houston Romer an ...
to include ''Mandasuchus'' and three other formally named genera of "
rauisuchia "Rauisuchia" is a paraphyletic group of mostly large and carnivorous Triassic archosaurs. Rauisuchians are a category of archosaurs within a larger group called Pseudosuchia, which encompasses all archosaurs more closely related to crocodilians ...
ns". Charig and two coauthors suggested in a 1965 study dealing with
saurischia Saurischia ( , meaning "reptile-hipped" from the Greek ' () meaning 'lizard' and ' () meaning 'hip joint') is one of the two basic divisions of dinosaurs (the other being Ornithischia), classified by their hip structure. Saurischia and Ornithi ...
ns that ''Mandasuchus'' was a possible ancestor of the "prosauropods" ( basal
sauropodomorphs Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lo ...
), without explanation. In his 1993 study of the phylogeny of Crocodylotarsi (croc-line archosaurs, now known as
Pseudosuchia Pseudosuchia, from Ancient Greek ψεύδος (''pseúdos)'', meaning "false", and σούχος (''soúkhos''), meaning "crocodile" is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely relat ...
), J. Michael Parrish argued that ''Mandasuchus'' was not a distinct genus. He proposed that it was in the same genus as the European archosaur '' Ticinosuchus ferox'' due to the similarity of some homologous postcranial bones, but withheld final taxonomic determination of ''Mandasuchus'' until Charig provided a published description of the material. However, Charig died in 1997 without publishing on many of the archosaurs he discovered. ''Mandasuchus'' was finally formally described in 2018 by
Richard J. Butler Richard James Butler is a vertebrate palaeontologist at the University of Birmingham, where he holds the title of professor of palaeobiology. His research focuses on ornithischian dinosaur evolution, dinosaur origins, and fossil tetrapod macroe ...
and colleagues, as part of a multinational investigation into the fossils of Triassic
rift In geology, a rift is a linear zone where the lithosphere is being pulled apart and is an example of extensional tectonics. Typical rift features are a central linear downfaulted depression, called a graben, or more commonly a half-graben ...
systems in
Tanzania Tanzania, officially the United Republic of Tanzania, is a country in East Africa within the African Great Lakes region. It is bordered by Uganda to the northwest; Kenya to the northeast; the Indian Ocean to the east; Mozambique and Malawi to t ...
and
Zambia Zambia, officially the Republic of Zambia, is a landlocked country at the crossroads of Central Africa, Central, Southern Africa, Southern and East Africa. It is typically referred to being in South-Central Africa or Southern Africa. It is bor ...
. A
phylogenetic analysis 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 data ...
recovered it as a
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 ...
n more basal than ''
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 mem ...
'' and not synonymous with ''Ticinosuchus,'' albeit not too distantly related either.


Description


Skull

Skull material for ''Mandasuchus'' is limited to
maxilla In vertebrates, the maxilla (: maxillae ) 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 two maxil ...
e and part of a
dentary In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone ...
. The maxilla is low, with an elongated
antorbital fenestra An antorbital fenestra (plural: fenestrae) is an opening in the skull that is in front of the eye sockets. This skull character is largely associated with Archosauriformes, archosauriforms, first appearing during the Triassic Period. Among Extant ...
and at least 12 tooth sockets separated by discrete interdental plates. The antorbital fenestra is surrounded by an inset basin, the antorbital fossa, as with other archosaurs. However, ''Mandasuchus'' has a restricted and weakly differentiated antorbital fossa compared to other
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 ...
ns and ''
Ticinosuchus ''Ticinosuchus'' is an extinct genus of suchian archosaur from the Middle Triassic (Anisian– Ladinian) of Switzerland and Italy. Description One of only a handful of fossil reptiles that have been found in Switzerland, ''Ticinosuchus'' (mea ...
''. The ascending (or dorsal) process of the maxilla, which lies in front of the antorbital fenestra, is short and very thin. This process is also uniquely diagonally oriented when seen from above, with its rear edge set inwards from the front edge. The only preserved tooth is thin and serrated. The dentary fragment is incomplete, but seemingly slender and similar to the maxilla.


Postcrania

The centrum (main spool-shaped component) of ''Mandasuchus'''s longest cervical (neck) vertebrae are about 1.8 times longer than high. This is similar to ''Ticinosuchus'' and intermediate between other loricatans (which have shorter vertebrae) and
poposauroids Poposauroidea is a clade of advanced pseudosuchians. It includes poposaurids, shuvosaurids, ctenosauriscids, and other unusual pseudosuchians such as '' Qianosuchus'' and ''Lotosaurus''. It excludes most large predatory quadrupedal " rauisuchia ...
(which have longer vertebrae). Smaller specimens have slightly shorter cervical vertebrae. The lower edge and sides of the cervicals are concave while the rib facets (dia- and parapophyses) and
articular processes The articular process or zygapophysis ( + apophysis) of a vertebra is a projection of the vertebra that serves the purpose of fitting with an adjacent vertebra. The actual region of contact is called the ''articular facet''.Moore, Keith L. et al. ...
(pre- and post-zygapophyses) are stout. A long ridge runs along the base of the neural spine while smaller ridges connect the articular processes to the front and rear edges of the neural spines, defining deep pockets in the process. The neural spines themselves are short and expanded at their upper extent to form "spine tables", with those near the shoulder resembling rearward-pointed teardrops when seen from above. Cervical ribs are thick and slightly contact each other. Dorsal (back) vertebrae are taller, wider, and their parapophyses gradually shift upwards, but dorsals are otherwise similar to the cervicals. Dorsal neural spines overhang the rear of their respective vertebrae and possess elliptical spine tables. The single known sacral (hip) vertebra is small, simple, and poorly preserved, but it seems to retain a small rear pocket. The caudal (tail) vertebrae gradually lengthen and simplify down the tail, they gain
chevrons Chevron (often relating to V-shaped patterns) may refer to: Science and technology * Chevron (aerospace), sawtooth patterns on some jet engines * Chevron (anatomy), a bone * '' Eulithis testata'', a moth * Chevron (geology), a fold in rock la ...
starting at the fifth caudal, and their neural spines gradually shorten. The neural spines have small spine tables, less distinct than those of the cervicals and dorsals.
Osteoderm Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amph ...
s (bony plates) extended down the back in two rows which were "staggered". i.e. offset from each other so that the armor is asymmetrical. Individual osteoderms were short, thick, and overlapped the pointed front extent of succeeding osteoderms. The
scapula The scapula (: scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on either side ...
expands towards its upper extremity (more so than ''Ticinosuchus'') and has concave front and rear edges. A small muscle scar is present on the rear edge near the
glenoid The glenoid fossa of the scapula or the glenoid cavity is a bone part of the shoulder. The word ''glenoid'' is pronounced or (both are common) and is from , "socket", reflecting the shoulder joint's ball-and-socket form. It is a shallow, pyrif ...
(shoulder socket), similar to one observed in ''
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 descri ...
,'' albeit less distinct. The
humerus The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
is narrow and has prominent and well-developed muscle attachments, while the
ulna The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to the wrist. It is on the same side of the forearm as the little finger, running parallel to the Radius (bone), radius, the forearm's other long ...
is shorter and quite robust. The ilium is low and similar to that of ''Ticinosuchus'', with muscle scars for two sacral ribs on its inner surface and no vertical ridge on its outer surface. The pubis is somewhat elongated, with its base forming part of the
acetabulum The acetabulum (; : acetabula), also called the cotyloid cavity, is a wikt:concave, concave surface of the pelvis. The femur head, head of the femur meets with the pelvis at the acetabulum, forming the Hip#Articulation, hip joint. Structure The ...
and possessing a small perforation known as an obturator fenestra. The tip of the pubis slightly expands into a bulbous pubic boot resembling that of other basal loricatans. The
ischium The ischium (; : is ...
is only slightly shorter than the pubis and also possesses a three-dimensional expansion at its tip. Although the left and right ischia would have contact each other extensively like those of ''Ticinosuchus'' and
paracrocodylomorphs 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. Paracrocodylomor ...
, this contact was apparently weak enough that the bones were separated during fossilization. The
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
has many archosaurian hallmarks near the hip, such as enlarged proximal tuberosities, a mound-like
fourth trochanter The fourth trochanter is a shared characteristic common to archosaurs. It is a protrusion on the posterior-medial side of the middle of the femur shaft that serves as a muscle attachment, mainly for the '' musculus caudofemoralis longus'', the m ...
, and possibly a groove on the upper surface of the femoral head. It also possessed a scar for the ''iliofemoralis'' muscle and a distinct oval pit on its rear edge, which is not known in other pseudosuchians. The
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
is straight and possesses typical pseudosuchian features, while the incomplete
fibula The fibula (: fibulae or fibulas) or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. ...
has a knob-shaped ''iliofibularis'' scar in its upper half. Although no foot bones are preserved, ''Mandasuchus'' does have two well-preserved proximal tarsals (ankle bones): the
astragalus Astragalus may refer to: * ''Astragalus'' (plant), a large genus of herbs and small shrubs *Astragalus (bone) The talus (; Latin for ankle or ankle bone; : tali), talus bone, astragalus (), or ankle bone is one of the group of foot bones known ...
and
calcaneum In humans and many other primates, the calcaneus (; from the Latin ''calcaneus'' or ''calcaneum'', meaning heel; : calcanei or calcanea) or heel bone is a bone of the tarsus of the foot which constitutes the heel. In some other animals, it is t ...
. Mandasuchus has a "crocodile-normal" or "
crurotarsal A crurotarsal joint is one that's situated between the bones of '' crus'', i.e. shin (tibia and fibula) and the proximal tarsal bones, i.e. astragalus and calcaneum. The ankle joint of therian mammals ( marsupials and placentals) is a crurotarsa ...
" ankle, with a prominent peg on the astragalus fitting into a shallower pit on the calcaneum. The ankle has many
suchia Suchia (from Ancient Greek σούχος (''soúkhos''), meaning "crocodile") is a clade of archosaurs containing the majority of pseudosuchians (crocodilians and their extinct relatives). It was defined as the least inclusive clade containing ''A ...
n features, including a flexed (albeit indistinct) tibial facet on the astragalus, and a wide, expanded calcaneal tuber offset from the calcaneum's contact with the fourth distal tarsal.


References


External links


Press release of the 2018 description on Sci-News, with life reconstruction by Mark Witton


{{Taxonbar, from=Q6747875 Paracrocodylomorpha Middle Triassic reptiles of Africa Anisian life Triassic Tanzania Fossils of Tanzania Fossil taxa described in 2018 Prehistoric pseudosuchian genera