Rhynchosaurs are a group of extinct herbivorous Triassic archosauromorph reptiles, 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 Early Triassic, reaching their broadest abundance and a global distribution during the Carnian stage of the Late Triassic.

Description

Life restoration of Hyperodapedon sanjuanensis.
Skull of Bentonyx

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 and Howesia, were generally small, typically lizard-like in build, and had skulls rather similar to the early diapsid Youngina, except for the beak and a few other features. Later and more advanced genera grew to 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 with a deep cheek region, and the premaxilla 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 (upper jaw), like the blade of a penknife 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 tubers by backwards scratching of the hind limbs. They had a fixed number of teeth, and similar to elephants, those that were further back in the jaws replaced teeth that were worn out as the animal grew in size and the teeth were worn out because of a diet of very tough plants. Rhynchosaur teeth were ankylothecodont, similar to the acrodonty of modern tuataras and some lizards but differing in the presence of deep roots.

Evolution

Rhynchosaurs first appeared during the Induan, the first stage of the Triassic, appearing shortly after the end-Permian mass extinction, as part of a major diversification of reptiles during this period. Like many animals of this time, they had a worldwide distribution, being found across Pangea, being especially common in the southern Gondwanan part of Pangaea, though with records also in Europe and North America. Over their evolution, rhynchosaurs saw a progressive increase in their body size. It has been suggested that rhynchosaurs went extinct at the end of the Carnian stage during the early Late Triassic as part of an end-Carnian extinction event. Spielmann, Lucas and Hunt (2013) described three distal ends of humeri from early-mid Norian Bull Canyon Formation in New Mexico, which they interpreted as bones of rhynchosaurs belonging to the species Otischalkia elderae; thus, the fossils might indicate that rhynchosaurs survived until the Norian. These fossils were later reinterpreted as belonging to malerisaurine azendohsaurids.

Classification

Relationships with other reptiles

Historically, rhynchosaurs were considered to be closely related to Rhynchocephalia, the clade that includes the living tuatara and its extinct relatives. Today however the two groups are considered to be unrelated, and rhynchosaurs are considered to be archosauromorphs, more closely related to modern crocodilians and birds than to rhynchocephalians and lizards, as members of the archosauromorph subclade Crocopoda.

Cladogram of Archosauromorpha after Schoch et al. 2025:

ArchosauromorphaJesairosaurusProtorosaurusProlacertaTanysauria Crocopoda †AllokotosauriaRhynchosauriaTeyujagua Archosauriformes
Jesairosaurus
ProtorosaurusProlacertaTanysauria Crocopoda †AllokotosauriaRhynchosauriaTeyujagua Archosauriformes
Protorosaurus
ProlacertaTanysauria Crocopoda †AllokotosauriaRhynchosauriaTeyujagua Archosauriformes
Prolacerta
Tanysauria
CrocopodaAllokotosauriaRhynchosauriaTeyujagua Archosauriformes
Allokotosauria
RhynchosauriaTeyujagua Archosauriformes
Rhynchosauria
Teyujagua Archosauriformes
Teyujagua
Archosauriformes

Phylogeny

Hyperodapedon huxleyi (=Paradapedon)
Mesosuchus browni
Illustration of the ventral surface of a tooth plate of Hyperodapedon.

The Rhynchosauria included a single family, named Rhynchosauridae. All rhynchosaurs, apart from the four Early and Middle Triassic monospecific genera, Eohyosaurus, Mesosuchus, Howesia and Noteosuchus, are included in this family. Hyperodapedontidae named by Lydekker (1885) was considered its junior synonym. However, Langer et al. (2000) noted that Hyperodapedontidae was erected by Lydekker to include Hyperodapedon gordoni and H. huxleyi, clearly excluding Rhynchosaurus articeps, which was the only other rhynchosaur known at that time. Thus, they defined it as the stem-based taxon 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 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).

The cladogram below is based on Schultz et al. (2016) which is the most genera inclusive rhynchosaur phylogenetic analysis 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.

RhynchosauriaNoteosuchus colletti Mesosuchus browni Howesia browni Eohyosaurus wolvaardti Rhynchosauridae Rhynchosaurus articeps Hyperodapedontidae Stenaulorhynchinae Ammorhynchus navajoi Mesodapedon kuttyi Brasinorhynchus mariantensis Stenaulorhynchus stockleyi Bentonyx sidensis Langeronyx brodiei Fodonyx spenceri Hyperodapedontinae Isalorhynchus genovefae Teyumbaita sulcognathus Hyperodapedon spp.
Noteosuchus colletti
Mesosuchus browni Howesia browni Eohyosaurus wolvaardti Rhynchosauridae Rhynchosaurus articeps Hyperodapedontidae Stenaulorhynchinae Ammorhynchus navajoi Mesodapedon kuttyi Brasinorhynchus mariantensis Stenaulorhynchus stockleyi Bentonyx sidensis Langeronyx brodiei Fodonyx spenceri Hyperodapedontinae Isalorhynchus genovefae Teyumbaita sulcognathus Hyperodapedon spp.
Mesosuchus browni
Howesia browni Eohyosaurus wolvaardti Rhynchosauridae Rhynchosaurus articeps Hyperodapedontidae Stenaulorhynchinae Ammorhynchus navajoi Mesodapedon kuttyi Brasinorhynchus mariantensis Stenaulorhynchus stockleyi Bentonyx sidensis Langeronyx brodiei Fodonyx spenceri Hyperodapedontinae Isalorhynchus genovefae Teyumbaita sulcognathus Hyperodapedon spp.
Howesia browni
Eohyosaurus wolvaardti Rhynchosauridae Rhynchosaurus articeps Hyperodapedontidae Stenaulorhynchinae Ammorhynchus navajoi Mesodapedon kuttyi Brasinorhynchus mariantensis Stenaulorhynchus stockleyi Bentonyx sidensis Langeronyx brodiei Fodonyx spenceri Hyperodapedontinae Isalorhynchus genovefae Teyumbaita sulcognathus Hyperodapedon spp.
Eohyosaurus wolvaardti
RhynchosauridaeRhynchosaurus articeps Hyperodapedontidae Stenaulorhynchinae Ammorhynchus navajoi Mesodapedon kuttyi Brasinorhynchus mariantensis Stenaulorhynchus stockleyi Bentonyx sidensis Langeronyx brodiei Fodonyx spenceri Hyperodapedontinae Isalorhynchus genovefae Teyumbaita sulcognathus Hyperodapedon spp.
Rhynchosaurus articeps
HyperodapedontidaeStenaulorhynchinae Ammorhynchus navajoi Mesodapedon kuttyi Brasinorhynchus mariantensis Stenaulorhynchus stockleyi Bentonyx sidensis Langeronyx brodiei Fodonyx spenceri Hyperodapedontinae Isalorhynchus genovefae Teyumbaita sulcognathus Hyperodapedon spp.
StenaulorhynchinaeAmmorhynchus navajoi Mesodapedon kuttyi Brasinorhynchus mariantensis Stenaulorhynchus stockleyi
Ammorhynchus navajoi
Mesodapedon kuttyi
Brasinorhynchus mariantensis Stenaulorhynchus stockleyi
Brasinorhynchus mariantensis
Stenaulorhynchus stockleyi
Bentonyx sidensis Langeronyx brodiei Fodonyx spenceri Hyperodapedontinae Isalorhynchus genovefae Teyumbaita sulcognathus Hyperodapedon spp.
Bentonyx sidensis
Langeronyx brodiei
Fodonyx spenceri Hyperodapedontinae Isalorhynchus genovefae Teyumbaita sulcognathus Hyperodapedon spp.
Fodonyx spenceri
HyperodapedontinaeIsalorhynchus genovefae Teyumbaita sulcognathus Hyperodapedon spp.
Isalorhynchus genovefae
Teyumbaita sulcognathus Hyperodapedon spp.
Teyumbaita sulcognathus
Hyperodapedon spp.

List of genera

List of rhynchosaur genera
GenusSpeciesAgeLocationUnitNotes
AmmorhynchusA. navajoiAnisianUS (Arizona)Moenkopi FormationA stenaulorhynchine
BeesiiwoB. cooowuseCarnianUS (Wyoming)Popo Agie FormationA hyperodapedontine previously referred to Hyperodapedon
BrasinorhynchusB. mariantensisLadinianBrazilSanta Maria FormationA stenaulorhynchine, previously known as the "Mariante Rhynchosaur"
BentonyxB. sidensislate AnisianUK (England)Otter Sandstone FormationA basal hyperodapedontid
EifelosaurusE. triadicusearly AnisianGermanyRöt FormationA basal rhynchosaurid
ElorhynchusE. carrollilate Ladinian? - earliest Carnian?ArgentinaChañares Formation (Tarjadia Assemblage Zone)A stenaulorhynchine
EohyosaurusE. wolvaardtiearly AnisianSouth AfricaBurgersdorp Formation (Cynognathus Assemblage Zone)A basal (non-rhynchosaurid) rhynchosaur
FodonyxF. spencerilate AnisianUK (England)Otter Sandstone FormationA basal hyperodapedontid
HowesiaH. browniearly AnisianSouth AfricaBurgersdorp Formation (Cynognathus Assemblage Zone)A basal (non-rhynchosaurid) rhynchosaur
HyperodapedonH. gordoniCarnianUK (Scotland)Lossiemouth Sandstone FormationA hyperodapedontine, one of the most abundant and speciose rhynchosaur genera. Six valid species have been named, the most of any rhynchosaur.
H. hueneiCarnianBrazilSanta Maria Formation
H. huxleyiCarnianIndiaLower Maleri Formation
H. mariensisCarnianBrazil ArgentinaSanta Maria Formation Ischigualasto Formation
H. sanjuanensisCarnianArgentina BrazilIschigualasto Formation Santa Maria Formation
H. tikiensisCarnianIndiaTiki Formation
IsalorhynchusI. genovefaeCarnianMadagascarMakay Formation (Isalo II)A hyperodapedontine occasionally referred to Hyperodapedon
IsodapedonI. varzealisCarnianBrazilSanta Maria FormationA hyperodapedontine
LangeronyxL. brodieiAnisianUK (England)Bromsgrove Sandstone FormationA basal hyperodapedontid
MesodapedonM. kuttyiAnisianIndiaYerrapalli FormationA stenaulorhynchine
MesosuchusM. browniearly AnisianSouth AfricaBurgersdorp Formation (Cynognathus Assemblage Zone)A basal (non-rhynchosaurid) rhynchosaur
NoteosuchusN. collettiearly InduanSouth AfricaKatberg Formation (Lystrosaurus Assemblage Zone)A basal (non-rhynchosaurid) rhynchosaur; the earliest known species of rhynchosaur, and the only known Early Triassic representative
OryctorhynchusO. bairdilatest Carnian?-earliest Norian?Canada (Nova Scotia)Wolfville FormationA hyperodapedontine previously referred to Hyperodapedon
RhynchosaurusR. articepsAnisianUK (England)Tarporley Siltstone FormationA basal rhynchosaurid
StenaulorhynchusS. stockleyilate AnisianTanzaniaManda FormationA stenaulorhynchine
SupradapedonS. stockleyiMiddle - Late TriassicTanzaniaTunduru districtA hyperodapedontine previously referred to Hyperodapedon
TeyumbaitaT. sulcognathuslate Carnian - early NorianBrazil ArgentinaCaturrita Formation Ischigualasto FormationThe latest surviving species, and the only rhynchosaur known with confidence to have survived into the Norian stage

Bibliography

External links

Data related to Rhynchosauria at Wikispecies