# Protuberum

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Extinct genus of cynodonts

Protuberum Temporal range: Middle Triassic PreꞒ Ꞓ O S D C P T J K Pg N Scientific classification Kingdom: Animalia Phylum: Chordata Clade: Synapsida Clade: Therapsida Clade: Cynodontia Family: †Traversodontidae Subfamily: †Gomphodontosuchinae Genus: †Protuberum Reichel et al., 2009 Species P. cabralense[1] Reichel et al., 2009 (type)

***Protuberum*** was a [traversodontid](/source/Traversodontidae) [cynodontian](/source/Cynodontia) from the [Middle Triassic](/source/Middle_Triassic) of Brazil[2] known from a single species, ***P. cabralense.***[1] It is most notable for its thickened skull and ribs, vertebrae and illium covered in bony bumps from which it gets its name, and was among the largest triassic cynodonts.

## Discovery and naming

The first specimen of *Protuberum* was collected in 1977. It consists of several ribs and vertebrae. The second, collected in 1989, consists of a partial articulated skeleton and a skull with most of the dentition still present. Both were collected by [Father Daniel Cargnin](/source/Daniel_Cargnin_(paleontologist)). Additional specimens were discovered including [thoracic](/source/Thorax) (chest) ribs, a [proximal](/source/Anatomical_terms_of_location) (close to the midline) fragment of a right [cervical](/source/Cervical_vertebrae) (neck) [rib](/source/Rib), an [ulna](/source/Ulna) (one of the two bones of the lower arm), and two [humeri](/source/Humerus) (lower arm bones).[2][3] In 2009 they were described by a team consisting of Míriam Reichel and colleagues. The second specimen was made the [holotype](/source/Holotype).[2]

The genus name refers to the large amount of protuberances on the ribs and [ilia](/source/Ilium_(bone)) (part of the hip), while the species name is in honor of the Municipality of [Novo Cabrais](/source/Novo_Cabrais), where the type specimen was collected.[2]

## Description

*Protuberum* was a large traversodontid, with the holotype subadult specimen weighting about 60.35 kg (133.0 lb), and other, probably adult specimens being 40% larger.[4][5]

### Skull and teeth

The cranium was big, measuring about 20 cm (7.9 in) long, sporting a short and high [parietal](/source/Parietal_bone) crest, 15 millimetres (0.59 in) long or only 7.5% or the skull's length. The preorbital bones, [lambdoid](/source/Lambdoid_suture) crest and the skull roof are thick, making the skull overall very robust.[2] This robustness along with the large size of the skeletal elements indicate that the holotype individual was probably a subadult.[3]

On the upper jaw, *Protuberum* had four [incisors](/source/Incisor), one [canine](/source/Canine_tooth), and [postcanines](/source/Cheek_teeth) on each side. The incisors are large and point slightly forward, and have a thick coat of [enamel](/source/Tooth_enamel) on the [labial](/source/Glossary_of_dentistry#Labial) surface (= the side touching the lips), with the largest being the third incisor. The canines are not preserved, with only a single scrap of a tooth remaining, although the fourth incisor was originally misinterpreted as a canine. The paracanine fossa (a depression on the upper jaw to accommodate the canines of the [mandible](/source/Mandible) when the mouth is closed) is larger than that of other traversodontids, and was originally interpreted a being so extensive it would create a uniquely long [diastema](/source/Diastema) (gap) between the canine and postcanine teeth, although this was later rejected. The postcanine teeth possess two cusps connected by a sharp transverse process. Unlike *[Massetognathus](/source/Massetognathus),* *[Exaeretodon](/source/Exaeretodon)* and some other [gomphodonts](/source/Gomphodontia), they lack a shouldering pattern. They are worn sequentially from the front of the mouth back, indicating that the postcanines erupted from the back of the mouth over time. However they likely did not move as new teeth grew, as indicated by a straight root.[2][5]

### Postcranial skeleton

*Protuberum* is most striking for its neural spines, presacral ribs and [illium](/source/Ilium_(bone)) which are covered in bumps. Analogous rib ornaments are found in the [ankylosaur](/source/Ankylosauria) *[Spicomellus](/source/Spicomellus)* and the [pseudosuchian](/source/Pseudosuchia) *[Euscolosuchus](/source/Euscolosuchus)*, although in the former they are epidermal.[6][7] *Thrinaxodon* had a similar condition with a series of tubercules on the proximal ribs.[8] *[Kayentatherium](/source/Kayentatherium)* was another cynodont with bump-like vertebral processes.[2][9] On the illium they are present on the iliac blade, above the hips, and are aligned with the bumps of the ribs. These bumps are made of compact bone tissue, possibly [pachyostotic](/source/Pachyostosis), and were most likely not covered in [cartilaginous tissue](/source/Cartilage).[2]

Like with other cynodonts, the ribcage can be separated into two distinct regions, [thoracic](/source/Thorax) and [lumbar](/source/Lumbar), based on the presence of costal plates. The thoracic region bears overlapping costal plates, where the proximal half of each rib is widened. Each rib is lined with a single row of bumps, which decrease in size [distally](/source/Anatomical_terms_of_location) (away from the midline), so that the largest are the near the spine. The longest middle thoracic ribs bear up to eight of these processes, and their number generally decreases with the length of the rib.[2]

## Classification

In the first phylogeny ran in 2009, *Protuberum* was recovered deeply nested within [Gomphodontosuchinae](/source/Gomphodontosuchinae)[2]

Diademodon Trirachodon Andescynodon Pascualgnathus Massetognathus Exaeretodon Scalenodontoides Menadon Protuberum Gomphodontosuchus Dadadon Santacruzodon Traversodon Luangwa Scalenodon hirschsoni Scalenodon angustifrons

In 2024 *Protuberum* was recovered as outside Gomphodontosuchinae:[5]

Thrinaxodon liorhinus Cynognathia Cynognathus crateronotus Gomphodontia Diademodontidae Titanogomphodon crassus Diademodon tetragonus Neogomphodontia Langbergia modisei Trirachodon berryi Sinognathinae Cricodon metabolus Beishanodon youngi Sinognathus gracilis Traversodontidae BP_1_5538 CGP_100 Etjoia dentitransitus Scalenodon ribeiroae Scalenodon angustifrons Pascualgnathus polanskii Andescynodon mendozensis Rosieria delsatei Habayia halbadieri Microscalenodon nanus Mandagomphodon attridgei Mandagomphodon hirschoni Luangwa sudamericana Luangwa drysdalii "Traversodontinae" Paratraversodon fransiscaensis Traversodon staleckeri Protuberum carabalense Gomphodontosuchinae Gomphodontosuchus brasiliensis Menadon Brazil Menadon Madagascar Proexaeretodon vincei Siriusgnathus niemeyerorum Santagnathus mariensis Exaeretodon riograndensis Exaeretodon argentinus Scalenodontoides macrodontes Massetognathinae Massetognathus ochagaviae Massetognathus pascuali Santacruzodon hopsoni Dadadon isaloi Arctotraversodontinae Maubeugia lotharingica Arctotraversodon plemmyridon Plinthogomphodon herpetairus Boreogomphodon jeffersoni Nanogomphodon wildi

## Paleobiology

The [anterior](/source/Anatomical_terms_of_location) (front) portion of the [iliac blade](/source/Iliac_crest) is strongly curved, giving more surface for muscle attachment and indicating that strong hindlimb muscles would have been present in life. This is suggests the presence of powerful hindlimbs and may point toward burrowing habits. This is also supported by the high cortical thickness of the ribs, which is characteristic of aquatic or burrowing mammals.[2][10] As is typical for traversodontids, protuberum was a specialized herbivore with teeth adapted for grinding vegetation, and may have used its powerful limbs to dig out roots. While the use of the bumps is not clear, these alonside the thickened skull and costal plates have been suggested to be defensive or burrowing adaptations.[2]

Like other therapsids, *Protuberum* experienced a very rapid growth in its first years of life. One of the specimens was an adult or close to skeletal maturity based on the thick skull and wide patches of peripheral [lamellar bone](/source/Bone) in the ulna, and shows continuous growth until maturity like other non-mammalian cynodonts.[10]

## Paleoecology

Protuberum was found in the lower [*Dinodontosaurus* Assemblage Zone](/source/Santa_Maria_Formation) of the [Santa Maria Formation](/source/Santa_Maria_Formation) of Brazi, in the 'Sítio Cortado' an 'Rincão do Pinhal' outcrops.[2][11][5] It would have lived alongside the traversodontids [*Exaertodon*](/source/Exaeretodon),[12] [Luangwa](/source/Luangwa_(cynodont)),[13] *[Massetognathus](/source/Massetognathus),*[13] *[Scalenodon](/source/Scalenodon),*[14] and *[Traversodon](/source/Traversodon),*[15] the probainognathians *[Aleodon](/source/Aleodon),*[16] *[Bonacynodon](/source/Bonacynodon),*[17] [*Candeariodon*](/source/Candelariodon),[18] *[Chiniquodon](/source/Chiniquodon),*[19] and *[Protheriodon](/source/Protheriodon)*, the [kannemeyeriiform](/source/Kannemeyeriiformes) [dicynodonts](/source/Dicynodontia) *[Dinodontosaurus](/source/Dinodontosaurus)* and *[Stahleckeria](/source/Stahleckeria),* the [rhynchosaur](/source/Rhynchosauria) *[Brasinorhynchus](/source/Brasinorhynchus),*[20] the [proterchampsiids](/source/Proterochampsidae) *[Retymaijychampsa](/source/Retymaijychampsa)*[21] and *[Pinheirochampsa](/source/Pinheirochampsa),*[22] the [pseudosuchians](/source/Pseudosuchia) [*Archeopelta,*](/source/Archeopelta)[23] *[Decuriasuchus](/source/Decuriasuchus),*[24] *[Dynamosuchus](/source/Dynamosuchus),*[25] *[Pagosvenator](/source/Pagosvenator),*[26] *[Parvosuchus](/source/Parvosuchus),*[27] *[Prestosuchus](/source/Prestosuchus)*,[28] *[Procerosuchus](/source/Procerosuchus),*[29] and *[Schultzsuchus](/source/Schultzsuchus),*[30] the [silesaurids](/source/Silesauridae) *[Gamatavus](/source/Gamatavus)*[31] and *[Gondwanax](/source/Gondwanax),*[32] the [aphanosaur](/source/Aphanosauria) *[Spondylosoma](/source/Spondylosoma),*[33] and the [owenetiid](/source/Owenettidae) [procolophonoid](/source/Procolophonoidea) *[Candelaria](/source/Candelaria_(reptile))*[34]

In the Triassic, the Santa Maria Formation would have been a arid to semi-arid environment, with long dry seasons and shorter wet seasons, comprising [ephemeral lakes](/source/Ephemeral_lakes).[35] The *Dinodontosaurus* Assemblage Zone represented a [mudflat](/source/Mudflat), and would have been wetter than later environments.[36]

## References

1. ^ [***a***](#cite_ref-neuter_1-0) [***b***](#cite_ref-neuter_1-1) The specific name *cabralense* is "necessarily emended from the original *P. cabralensis* because *Protuberum* is a neuter generic name". Kammerer, Christian F. (2014). "Chapter 10. Theriodonta: Introduction". In Christian F. Kammerer; Kenneth D. Angielczyk; Jörg Fröbisch (eds.). *Early Evolutionary History of the Synapsida*. Springer Science+Business Media Dordrecht. p. 167. [doi](/source/Doi_(identifier)):[10.1007/978-94-007-6841-3_10](https://doi.org/10.1007%2F978-94-007-6841-3_10).

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Taxon identifiers Protuberum Wikidata: Q149194 GBIF: 8599834 IRMNG: 1477101 Open Tree of Life: 6158348 Paleobiology Database: 172338

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