Rhynchosaurs are a group of extinct herbivorous
Triassic
The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
archosauromorph
Archosauromorpha ( Greek for "ruling lizard forms") is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs (such as crocodilians and dinosaurs, including birds) rather than lepidosaurs (such as tuataras, l ...
reptile
Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s, belonging to the order Rhynchosauria. Members of the group are distinguished by their triangular skulls and elongated, beak like
premaxillary bones. Rhynchosaurs first appeared in the Middle Triassic or possibly the Early Triassic, before becoming abundant and globally distributed during the
Carnian
The Carnian (less commonly, Karnian) is the lowermost stage of the Upper Triassic Series (or earliest age of the Late Triassic Epoch). It lasted from 237 to 227 million years ago (Ma). The Carnian is preceded by the Ladinian and is followed by t ...
stage of the Late Triassic.
Description
Rhynchosaurs were herbivores, and at times abundant (in some fossil localities accounting for 40 to 60% of specimens found), with stocky bodies and a powerful beak. Early primitive forms, like ''
Mesosuchus
''Mesosuchus'' ("middle crocodile") is an extinct genus of basal Rhynchosaur from early Middle Triassic (early Anisian stage) deposits of Eastern Cape, South Africa. It is known from the holotype SAM 5882, a partial skeleton, and from the ...
'' and ''
Howesia'', were generally small and more typically lizard-like in build, and had skulls rather similar to the early diapsid ''
Youngina
''Youngina'' is an extinct genus of diapsid reptile from the Late Permian Beaufort Group ('' Tropidostoma''-''Dicynodon'' zones) of the Karoo Red Beds of South Africa. This, and a few related forms, make up the family Younginidae, within the Ord ...
'', except for the beak and a few other features. Later and more advanced genera grew to medium to medium large size, up to two meters in length. The skull in these forms were short, broad, and triangular, becoming much wider than long in the most advanced forms like ''
Hyperodapedon
''Hyperodapedon'' is a genus of rhynchosaurs (beaked, archosaur-like reptiles) from the Late Triassic period (Carnian stage). Fossils of the genus have been found in Africa, Asia, Europe and North and South America. Its first discovery and namin ...
'' (= ''
Scaphonyx
''Hyperodapedon'' is a genus of rhynchosaurs (beaked, archosaur-like reptiles) from the Late Triassic period (Carnian stage). Fossils of the genus have been found in Africa, Asia, Europe and North and South America. Its first discovery and naming ...
''), with a deep cheek region, and 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 b ...
extending outwards and downwards to form the upper beak. The broad skull would have accommodated powerful jaw muscles. The lower jaw was also deep, and when the mouth was closed it clamped firmly into 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. The t ...
(upper jaw), like the blade of a
penknife
Penknife, or pen knife, is a British English term for a small folding knife. Today the word ''penknife'' is the common British English term for both a pocketknife, which can have single or multiple blades, and for multi-tools, with additional too ...
closing into its handle. This scissors-like action would have enabled rhynchosaurs to cut up tough plant material.
The teeth were unusual; those in the maxilla and palate were modified into broad tooth plates. The hind feet were equipped with massive claws, presumably for digging up roots and
tuber
Tubers are a type of enlarged structure used as storage organs for nutrients in some plants. They are used for the plant's perennation (survival of the winter or dry months), to provide energy and nutrients for regrowth during the next growin ...
s by backwards scratching of the hind limbs.
Like many animals of this time, they had a worldwide distribution, being found across
Pangea
Pangaea or Pangea () was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia during the Carboniferous approximately 335 million y ...
. These abundant animals might have died out suddenly at the end of the
Carnian
The Carnian (less commonly, Karnian) is the lowermost stage of the Upper Triassic Series (or earliest age of the Late Triassic Epoch). It lasted from 237 to 227 million years ago (Ma). The Carnian is preceded by the Ladinian and is followed by t ...
(Middle of the Late Triassic period), perhaps as a result of the extinction of the ''
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 ...
'' flora on which they may have fed. On the other hand, Spielmann, Lucas and Hunt (2013) described three distal ends of
humeri
The humerus (; ) 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 and ulna, and consists of three sections. The humeral upper extremity consists of a round ...
from early-mid
Norian
The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian.
Stratigraphic defi ...
Bull Canyon Formation in
New Mexico
)
, population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano)
, seat = Santa Fe
, LargestCity = Albuquerque
, LargestMetro = Tiguex
, OfficialLang = None
, Languages = English, Spanish ( New Mexican), Navajo, Ker ...
, which they interpreted as bones of rhynchosaurs belonging to the species ''
Otischalkia elderae''; thus, the fossils might indicate that rhynchosaurs survived until the Norian.
Classification
Taxonomy
Phylogeny
The Rhynchosauria included a single family, named Rhynchosauridae. All rhynchosaurs, apart from the four
Early and
Middle Triassic
In the geologic timescale, the Middle Triassic is the second of three epochs of the Triassic period or the middle of three series in which the Triassic system is divided in chronostratigraphy. The Middle Triassic spans the time between Ma and ...
monospecific genera, ''
Eohyosaurus
''Eohyosaurus'' is an extinct genus of basal rhynchosaur known from the early Middle Triassic (early Anisian stage) Burgersdorp Formation of Free State, South Africa. It contains a single species, ''Eohyosaurus wolvaardti''.
Discovery
''Eohyo ...
'', ''
Mesosuchus
''Mesosuchus'' ("middle crocodile") is an extinct genus of basal Rhynchosaur from early Middle Triassic (early Anisian stage) deposits of Eastern Cape, South Africa. It is known from the holotype SAM 5882, a partial skeleton, and from the ...
'', ''
Howesia'' and ''
Noteosuchus
''Noteosuchus'' is an extinct genus of basal rhynchosaur known from the earliest Triassic deposits of Eastern Cape Province, South Africa. It was first named by David Meredith Seares Watson in 1912 and the type species is ''Eosuchus colletti'' ...
'', are included in this family.
[ Hyperodapedontidae named by Lydekker (1885) was considered its ]junior synonym
The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently.
* In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linna ...
. However, Langer ''et al.'' (2000) noted that Hyperodapedontidae was erected by Lydekker to include ''Hyperodapedon
''Hyperodapedon'' is a genus of rhynchosaurs (beaked, archosaur-like reptiles) from the Late Triassic period (Carnian stage). Fossils of the genus have been found in Africa, Asia, Europe and North and South America. Its first discovery and namin ...
gordoni'' and ''H. huxleyi'', clearly excluding ''Rhynchosaurus articeps
''Rhynchosaurus'' (''beaked lizard'') is a genus of rhynchosaur that lived during the Middle Triassic Period (geology), period. It lived in Europe. It was related to the archosauria, archosaurs, but not within that group. The type species of ''R ...
'', which was the only other rhynchosaur known at that time. Thus, they defined it as the stem-based taxon
Phylogenetic nomenclature is a method of nomenclature for taxa in biology that uses phylogenetic definitions for taxon names as explained below. This contrasts with the traditional approach, in which taxon names are defined by a '' type'', which ...
that includes all rhynchosaurs more closely related to ''Hyperodapedon'' than to ''Rhynchosaurus''.
Within Hyperodapedontidae, which is now a subgroup of Rhynchosauridae, two subfamilies have been named. Stenaulorhynchinae named by Kuhn (1933) is defined sensu Langer and Schultz (2000) to include all species more closely related to ''Stenaulorhynchus
''Stenaulorhynchus'' (possibly meaning "narrow tube beak") is an extinct genus of hyperodapedontid rhynchosaur known from the Middle Triassic (late Anisian stage) deposits of Tanganyika Territory, Tanzania. It was found in the Lifua Member of th ...
'' than to ''Hyperodapedon''. Hyperodapedontinae named by Chatterjee (1969) was redefined by Langer ''et al.'' (2000) to include "all rhynchosaurs closer to ''Hyperodapedon'' than to ''"Rhynchosaurus" spenceri''" (now ''Fodonyx
''Fodonyx'' (meaning "digging claw") is an extinct genus of rhynchosaur from the middle Triassic epoch of Devon in England. Its fossils (25 specimens) were discovered in Otter Sandstone Formation (late Anisian age) and were first assigned to '' R ...
'').
The cladogram
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
below is based on Schultz ''et al.'' (2016) which is the most genera inclusive rhynchosaur phylogenetic analysis
In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
to date,[ with the position of ''Noteosuchus'' taken from other recent analyses (since it was removed in Schultz ''et al.'' (2016)), all in consensus with one another.][Ezcurra MD. (2016) The phylogenetic relationships of basal archosauromorphs, with an emphasis on the systematics of proterosuchian archosauriforms. ''PeerJ'', 4:e177]
/ref>
References
Bibliography
* Michael J. Benton, Benton, M. J. (2000), ''Vertebrate Paleontology'', 2nd ed. Blackwell Science Ltd.
* Carroll, R. L. (1988), ''Vertebrate Paleontology and Evolution'', W.H. Freeman & Co.
* Dilkes, D. W. 1998. The Early Triassic rhynchosaur ''Mesosuchus browni'' and the interrelationships of basal archosauromorph reptiles. '' Philosophical Transactions of the Royal Society of London: Biological Sciences'', 353:501-541.
External links
Dinosaurs of Rio Grande do Sul, Brasil.
{{Taxonbar, from=Q131768
Triassic reptiles
Triassic taxonomic orders
Anisian life
Carnian life
Norian life
Olenekian life
Early Triassic reptiles
Middle Triassic reptiles
Late Triassic reptiles
Olenekian first appearances
Norian extinctions