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Meraxes; Temporal range: Late Cretaceous, (Cenomanian), ~95–93.9 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
	Reconstructed skull
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Clade:	Dinosauria
Clade:	Saurischia
Clade:	Theropoda
Family:	†Carcharodontosauridae
Tribe:	†Giganotosaurini
Genus:	†Meraxes; Canale et al., 2022
Species:	†M. gigas
Binomial name
	†Meraxes gigas; Canale et al., 2022

Meraxes is a genus of large carcharodontosaurid theropod dinosaur from the Late Cretaceous Huincul Formation of Patagonia, Argentina. The genus contains a single species, Meraxes gigas.^[1]

Discovery and naming

The holotype of Meraxes, MMCh-PV 65, was discovered in 2012. Known from a nearly complete skull, pectoral and pelvic elements, partial forelimbs, complete hindlimbs, fragmentary ribs and cervical and dorsal vertebrae, a sacrum, and several complete caudal vertebrae, it represents the most complete carcharodontosaurid skeleton known from the Southern Hemisphere.^[1] The specimen was referred to as the "Campanas carcharodontosaurid" before its description as a new species in 2022.^[2]

Meraxes gigas was described in 2022 by Canale et al. based on these remains. The generic name, "Meraxes", honors a female dragon ridden by Queen Rhaenys Targaryen in the George R. R. Martin novel series, A Song of Ice and Fire. The specific name, "gigas", means "giant" in Greek, in reference to its large size.^[1]

Description

Meraxes is one of the largest theropods, weighing approximately 4.26 metric tons (4.70 short tons).^[1] Henderson (2023) listed a body length estimate of 9–10 metres (30–33 ft), referencing Canale et al. (2022), but also estimated a body length of 10.2–11.6 metres (33–38 ft) using the pelvic area.^[3] Its skull is 1.27 m (4.2 ft) long, similar to that of Acrocanthosaurus, which has a skull length of 1.23–1.29 m (4.0–4.2 ft).^[1]^[4] The shapes and proportions of various bones, including the skull, scapula, metacarpals, ischial shaft, and foot, indicate that Meraxes and Acrocanthosaurus had similar proportions and body size. Meraxes possessed reduced forelimbs, an instance of convergent evolution that occurred independently in four different lineages: Carcharodontosauridae, Abelisauridae, Tyrannosauridae, and Alvarezsauridae. Additionally, the second toes possess an enlarged claw, almost twice as long as the claw on the fourth toe.^[1]

Osteohistological analysis of the holotype suggests the individual could have been between 39 and 53 years old when it died, having reached skeletal maturity approximately 4 years prior to its death (between 35 and 49 years old), making it the longest-lived non-avian theropod currently known. Meraxes was determined to have grown to large size by extending its growth period (hypermorphosis), rather than increasing its relative growth rate (acceleration) through development as in Tyrannosaurus, to which it was compared.^[1]^[2]

Classification

Canale et al. (2022) recovered Meraxes as the earliest diverging member of the tribe Giganotosaurini within the Carcharodontosauridae. The results of their phylogenetic analyses are displayed in the cladogram below:^[1]

Carcharodontosauridae

Neovenator

Concavenator

Eocarcharia

Lajasvenator

Lusovenator

Acrocanthosaurus

Shaochilong

Carcharodontosaurinae

Carcharodontosaurus spp.

Giganotosaurini

Meraxes

Tyrannotitan

	Giganotosaurus

	Mapusaurus

Cau (2024) also recovered similar relationships for Meraxes, but not in a clade with Tyrannotitan, Giganotosaurus, and Mapusaurus, as suggested by Canale et al. (2022).^[5]

Carcharodontosauridae

Neovenator

	Sauroniops

	Veterupristisaurus

	Lusovenator

	Eocarcharia (type skull roof)

	Concavenator

Carcharodontosaurus iguidensis (holotype maxilla)

Acrocanthosaurus

Eocarcharia (referred maxilla)

	Meraxes

	Carcharodontosaurus iguidensis (referred cranial material)

Lajasvenator

	Labocania

	Shaochilong

	Carcharodontosaurus saharicus (neotype)

	Carcharodontosaurus saharicus (described by Stromer in 1931)

Tyrannotitan

	Mapusaurus

	Giganotosaurus

Paleoenvironment

The fossil remains of Meraxes were recovered from the Huincul Formation. A substantial number of taxa are known to have inhabited this paleoenvironment. Theropods from the formation include the paravian Overoraptor, the elaphrosaurine Huinculsaurus, the abelisaurs Skorpiovenator, Tralkasaurus, and Ilokelesia, the megaraptoran Aoniraptor, and the fellow giant carcharodontosaurids Mapusaurus and Taurovenator.^[6]^[7] Meraxes was found in an older rock layer than either Mapusaurus and Taurovenator, so it is unlikely they coexisted.^[1]^[8] The herbivores of the area are represented by the rebbachisaurid sauropods Cathartesaura and Limaysaurus, the titanosaurs Argentinosaurus, Choconsaurus, and Chucarosaurus, and indeterminate iguanodonts.^[9]^[10]^[11]

References

^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ Canale, Juan I.; Apesteguía, Sebastián; Gallina, Pablo A.; Mitchell, Jonathan; Smith, Nathan D.; Cullen, Thomas M.; Shinya, Akiko; Haluza, Alejandro; Gianechini, Federico A.; Makovicky, Peter J. (July 2022). "New giant carnivorous dinosaur reveals convergent evolutionary trends in theropod arm reduction". Current Biology. 32 (14): 3195–3202.e5. Bibcode:2022CBio...32E3195C. doi:10.1016/j.cub.2022.05.057. PMID 35803271. S2CID 250343124.
^ ^a ^b Cullen, Thomas M.; Canale, Juan I.; Apesteguía, Sebastián; Smith, Nathan D.; Hu, Dongyu; Makovicky, Peter J. (25 November 2020). "Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution". Proceedings of the Royal Society B: Biological Sciences. 287 (1939): 20202258. doi:10.1098/rspb.2020.2258. PMC 7739506. PMID 33234083. S2CID 227154091.
^ Henderson, Donald M. (2023). "Growth constraints set an upper limit to theropod dinosaur body size". The Science of Nature. 110 (1). 4. Bibcode:2023SciNa.110....4H. doi:10.1007/s00114-023-01832-1.
^ Currie, Philip J.; Carpenter, Kenneth (2000). "A new specimen of Acrocanthosaurus atokensis (Theropoda, Dinosauria) from the Lower Cretaceous Antlers Formation (Lower Cretaceous, Aptian) of Oklahoma, USA". Geodiversitas. 22 (2): 207–246.
^ Cau, Andrea (2024). "A Unified Framework for Predatory Dinosaur Macroevolution" (PDF). Bollettino della Società Paleontologica Italiana. 63 (1): 1-19. doi:10.4435/BSPI.2024.08 (inactive 20 November 2024).{{cite journal}}: CS1 maint: DOI inactive as of November 2024 (link)
^ Matías J. Motta; Federico L. Agnolín; Federico Brissón Egli; Fernando E. Novas (2020). "New theropod dinosaur from the Upper Cretaceous of Patagonia sheds light on the paravian radiation in Gondwana". The Science of Nature. 107 (3): Article number 24. Bibcode:2020SciNa.107...24M. doi:10.1007/s00114-020-01682-1. hdl:11336/135530. PMID 32468191. S2CID 218913199.
^ Cerroni, M.A.; Motta, M.J.; Agnolín, F.L.; Aranciaga Rolando, A.M.; Brissón Egli, F.; Novas, F.E. (2020). "A new abelisaurid from the Huincul Formation (Cenomanian-Turonian; Upper Cretaceous) of Río Negro province, Argentina". Journal of South American Earth Sciences. 98: 102445. Bibcode:2020JSAES..9802445C. doi:10.1016/j.jsames.2019.102445. S2CID 213781725.
^ Rolando, Alexis M. Aranciaga; Motta, Matías J.; Agnolín, Federico L.; Tsuihiji, Takanobu; Miner, Santiago; Brissón-Egli, Federico; Novas, Fernando E. (9 October 2024). "A new carcharodontosaurid specimen sheds light on the anatomy of South American giant predatory dinosaurs". The Science of Nature. 111 (6): 56. doi:10.1007/s00114-024-01942-4. ISSN 1432-1904.
^ Calvo, Jorge O.; Salgado, Leonardo (1995). "Rebbachisaurus tessonei sp. nov. a new Sauropoda from the Albian-Cenomanian of Argentina; new evidence on the origin of the Diplodocidae" (PDF). Gaia. 11: 13–33. Archived from the original (PDF) on 23 September 2021.
^ Baiano, Mattia A.; Coria, Rodolfo A.; Cau, Andrea (June 2020). "A new abelisauroid (Dinosauria: Theropoda) from the Huincul Formation (lower Upper Cretaceous, Neuquén Basin) of Patagonia, Argentina". Cretaceous Research. 110: 104408. Bibcode:2020CrRes.11004408B. doi:10.1016/j.cretres.2020.104408. S2CID 214118853.
^ Agnolin, Federico L.; Gonzalez Riga, Bernardo J.; Aranciaga Rolando, Alexis M.; Rozadilla, Sebastián; Motta, Matías J.; Chimento, Nicolás R.; Novas, Fernando E. (2 February 2023). "A new giant titanosaur (Dinosauria, Sauropoda) from the Upper Cretaceous of Northwestern Patagonia, Argentina". Cretaceous Research. 146: 105487. Bibcode:2023CrRes.14605487A. doi:10.1016/j.cretres.2023.105487. ISSN 0195-6671.