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''Zambiasaurus'' 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
dicynodont Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivores that typically bore a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, t ...
s that was discovered in 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 ...
(
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 ...
)
Ntawere Formation The Ntawere Formation is a Middle Triassic (Anisian) geological formation in Zambia, preserving fossils of synapsids, archosaurs, and temnospondyls. Geology Several different facies are present in the Ntawere Formation, reconstructing a floodpl ...
of
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 ...
, southern Africa. It was a large
dicynodont Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivores that typically bore a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, t ...
, reconstructed using several fossil fragments, in majority belonging to probably a juvenile ''Zambiasaurus submersus''. ''Zambiasaurus submersus'' is the
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 ...
of the genus ''Zambiasaurus''. It is a stahleckeriid, the first stahleckeriid known outside
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
, and it is also the oldest known stahleckeriid.


Description


Skull

The skull of ''Zambiasaurus submersus'' was reconstructed by using eighteen different juvenile
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
fragments and fragments of an adult Zambiasaurus. The immature skull had the dimensions of long and wide while the adults had a skull long and wide. There are no teeth in both upper and lower jaws making it a herbivorous species. Skull tapers
anteriorly Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek language, Greek roots, describe something in its standard anatomical position. Thi ...
, widest across
occiput The occipital bone () is a cranial dermal bone and the main bone of the occiput (back and lower part of the skull). It is trapezoidal in shape and curved on itself like a shallow dish. The occipital bone lies over the occipital lobes of the ...
. Wide inter-orbital region and narrow inter-temporal region. A blunt
snout A snout is the protruding portion of an animal's face, consisting of its nose, mouth, and jaw. In many animals, the structure is called a muzzle, Rostrum (anatomy), rostrum, beak or proboscis. The wet furless surface around the nostrils of the n ...
and a short median suture between nasals.
Pineal foramen A parietal eye (third eye, pineal eye) is a part of the epithalamus in some vertebrates. The eye is at the top of the head; is photoreceptive; and is associated with the pineal gland, which regulates circadian rhythmicity and hormone production ...
surrounded by parietals. Preparietal bone absent. Parietals slightly concave
antero-posteriorly Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position provi ...
and from bulk of intertemporal bar.
Interparietal bone An interparietal bone (os interparietale or Inca bone or ''os inca var.'') is a dermal bone situated between the parietal and supraoccipital. It is homologous to the postparietal bones of other animals. In humans, it corresponds to the upper ...
doesn't extend far forward. No sharp median intertemporal ridge. Sharp transition between dorsal and
occipital The occipital bone () is a cranial dermal bone and the main bone of the occiput (back and lower part of the skull). It is trapezoidal in shape and curved on itself like a shallow dish. The occipital bone lies over the occipital lobes of the cere ...
surface. Occipital wings of
squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone. In most tetrapods, the squamosal and quadratojugal bones form the cheek series of the skull. The bone forms an ancestra ...
extend laterally and somewhat posteriorly.
Palatal 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 sepa ...
surface of
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 mammals h ...
bears pair of anterior ridges.


Post-cranial elements

Fragmented
vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...
e showed the
prezygapophyses 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. ...
is concave and the postzygapophyses is convex. Fragments of ribs did not provide any useful information. Reconstructed from a lower and upper fragment, 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 ...
showed a long and narrow blade, low
scapular spine The spine of the scapula or scapular spine is a prominent plate of bone, which crosses obliquely the medial four-fifths of the scapula at its upper part, and separates the Supraspinatous fossa, supra- from the infraspinatous fossa. Structure It b ...
running along the antero-lateral edge.
Acromion process In human anatomy, the acromion (from Greek: ''akros'', "highest", ''ōmos'', "shoulder", : acromia) or summit of the shoulder is a bony process on the scapula (shoulder blade). Together with the coracoid process, it extends laterally over the shou ...
is not very large. Origin of scapular ligament of
triceps The triceps, or triceps brachii (Latin for "three-headed muscle of the arm"), is a large muscle on the ventral, back of the upper limb of many vertebrates. It consists of three parts: the medial, lateral, and long head. All three heads cross the ...
muscle on postero-dorsal edge above 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 ...
. There is no notch on the lower edge of 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 ...
so the
foramen In anatomy and osteology, a foramen (; : foramina, or foramens ; ) is an opening or enclosed gap within the dense connective tissue (bones and deep fasciae) of extant and extinct amniote animals, typically to allow passage of nerves, artery, ...
laid entirely inside the precoracoid. Fragments of
clavicle The clavicle, collarbone, or keybone is a slender, S-shaped long bone approximately long that serves as a strut between the scapula, shoulder blade and the sternum (breastbone). There are two clavicles, one on each side of the body. The clavic ...
provided no useful information. Sternum is hexagonal and ventral side is slightly concave. Dorsal side has postero-lateral bosses that have cartilage-ornaments on the ends. Reconstructed from two complementary fragments, 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 strongly twisted expressing very poorly muscular insertion and articulation areas. The
radius In classical geometry, a radius (: radii or radiuses) of a circle or sphere is any of the line segments from its Centre (geometry), center to its perimeter, and in more modern usage, it is also their length. The radius of a regular polygon is th ...
is also incomplete and reconstructed from two
distal Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position provi ...
fragments. It is slender and configured to be longer than 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 ...
. A complete
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 ...
shows a slender bone and convex distal ends.


Classification

''Zambiasaurus'' is characterized as a stahleckeriid, because of many shared characteristics such as; * Blunt
snout A snout is the protruding portion of an animal's face, consisting of its nose, mouth, and jaw. In many animals, the structure is called a muzzle, Rostrum (anatomy), rostrum, beak or proboscis. The wet furless surface around the nostrils of the n ...
* Wide but low
occiput The occipital bone () is a cranial dermal bone and the main bone of the occiput (back and lower part of the skull). It is trapezoidal in shape and curved on itself like a shallow dish. The occipital bone lies over the occipital lobes of the ...
* Short
temporal opening The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
* Lack of a high intertemporal crest * absence of preparietal * Nasals meet for a short distance in the midline or separated by a junction * Narrow
neural spine Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...
similar to anterior vertebrae of juvenile ''
Stahleckeria ''Stahleckeria'' is an extinct genus of Middle Triassic (Ladinian) dicynodonts.''Stahle ...
'' *
Coracoid A coracoid is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is present as part of the scapula, but this is n ...
,
Sternum The sternum (: sternums or sterna) or breastbone is a long flat bone located in the central part of the chest. It connects to the ribs via cartilage and forms the front of the rib cage, thus helping to protect the heart, lungs, and major bl ...
, and
Pelvis The pelvis (: pelves or pelvises) is the lower part of an Anatomy, anatomical Trunk (anatomy), trunk, between the human abdomen, abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also c ...
generally similar * Absence of
tusk Tusks are elongated, continuously growing front teeth that protrude well beyond the mouth of certain mammal species. They are most commonly canine tooth, canine teeth, as with Narwhal, narwhals, chevrotains, musk deer, water deer, muntjac, pigs, ...
s signify a close relation to ''
Stahleckeria ''Stahleckeria'' is an extinct genus of Middle Triassic (Ladinian) dicynodonts.''Stahle ...
'' rather than ''
Dinodontosaurus ''Dinodontosaurus'' (meaning "terrible-toothed lizard") is a genus of dicynodont therapsid. It was medium to large dicynodont of the Triassic (with skull up to long) and had a beak corneum. It lived in the Middle Triassic but disappeared in the ...
'' or '' Chanaria'', the other two Stakleckeriid
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 ...


History

''Zambiasaurus'' was named after the country of origin
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 ...
and ''Zambiasaurus submersus'' means the likelihood that the species drowned. Fossil reptiles were first found in upper
Luangwa River The Luangwa River is one of the major Tributary, tributaries of the Zambezi River, and one of the four biggest rivers of Zambia. The river generally floods in the rainy season (December to March) and then falls considerably in the dry season. I ...
valley by Dixey in 1928 and 1935. The area was later re-examined in 1960 and 1960 by Dr. A. R. Drysdall and James Kitching. They examined an area north-west of Sitwe for four months and collected almost 500
specimen Specimen may refer to: Science and technology * Sample (material), a limited quantity of something which is intended to be similar to and represent a larger amount * Biological specimen or biospecimen, an organic specimen held by a biorepository f ...
s. Later an expedition to Sitwe was done by a joint effort of the
British Museum The British Museum is a Museum, public museum dedicated to human history, art and culture located in the Bloomsbury area of London. Its permanent collection of eight million works is the largest in the world. It documents the story of human cu ...
(Natural History) and
University of London The University of London (UoL; abbreviated as Lond or more rarely Londin in Post-nominal letters, post-nominals) is a collegiate university, federal Public university, public research university located in London, England, United Kingdom. The ...
. They spent six weeks collecting 220 specimens. Even though there was a large collection of specimens the specimens themselves were imperfect or poorly documented.


Paleobiology

Thought to be
herbivorous A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat n ...
because of the lack of teeth and movement of lower jaws that appeared to be efficient in
mastication Chewing or mastication is the process by which food is comminution, crushed and ground by the teeth. It is the first step in the process of digestion, allowing a greater surface area for digestive enzymes to break down the foods. During the mast ...
. ''Zambiasaurus'' had a barrel shaped body that was strongly built but very slow. The body was held off the ground but it either had a sprawling stance or a more upright stance.


See also

*
List of therapsids This list of therapsids is an attempt to create a comprehensive listing of all Genus, genera that have ever been included in the Therapsida excluding mammals and purely vernacular terms. The list includes all commonly accepted genera, but also gene ...
*
Dicynodont Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivores that typically bore a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, t ...
* Stahleckeriids


References


External links


The main groups of non-mammalian synapsids at Mikko's Phylogeny Archive
{{Portal bar, Paleontology Kannemeyeriiformes Anisian life Middle Triassic synapsids of Africa Fossil taxa described in 1969 Anomodont genera