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The Romualdo Formation (Araripe Basin) is worldwide known for the large number of well-preserved fossils but the dinosaur record is rather scarce
which is the first tetrapod recovered from the basal layers of this stratigraphic unit that consist of dark shales
is known by an incomplete but articulated right hind limb with the distal portion of the femur
The new species differs from other coelurosaurs by a medial fossa in the tibia and digits II
The phylogenetic analysis recovered Aratasaurus museunacionali closely related to Zuolong salleei
The paleohistology indicate that the specimen is a juvenile
with an estimated body length around 3.12 m
The new taxon represents the first occurrence of basal coelurosaurians in the Araripe Basin and suggests a widespread distribution of this group during the Lower Cretaceous
Location map of the Mina Pedra Branca, Ceará State. The crossed geologic hammers indicate where Aratasaurus museunacionali gen. et sp. nov. was found. Figure created by Renan Alfredo Machado Bantim on PS Adobe Photoshop CC, version 20.0.6.
Outcrop of Mina Pedra Branca where Aratasaurus museunacionali gen
was recovered with indication of the stratigraphy and where the dinosaur came from
Figure created by Renan Alfredo Machado Bantim on PS Adobe Photoshop CC
Composed stratigraphic section of Mina Pedra Branca quarry (Municipality of Santana do Cariri
showing the fossiliferous calcareous nodule level (A) and the dark shale horizon (dinosaur) where Aratasaurus museunacionali gen
were recovered from this layer such as fishes (e.g.
the sole tetrapod known form this deposit is the dinosaur described here
From the combination of “ara” and “atá” from the Tupi language meaning born and fire
Incomplete but articulated right hind limb with the distal portion of the femur
proximal half of the tibia and mid-distal regions of metatarsals I–IV
The specimen (MPSC R 2089) is housed at the Museu de Paleontologia of the Universidade Regional do Cariri
and a cast will be deposited at the Museu Nacional/UFRJ
in a dark shale located about 2.5 m above the contact with Ipubi Formation
Aratasaurus museunacionali differs from other basal coelurosaurs by the following combination of characters: tibia exhibiting a medial fossa; symmetric pes
with digits II and IV subequal in total length; distal condyles of metatarsi II
III and IV symmetric mediolaterally and with subequal width; width of metatarsi II and IV similar
presenting the dorsal surface of the distal articulation bulbous
The holotype (MPSC R 2089) of Aratasaurus museunacionali gen
showing the femur and tibia before preparation
Only a section of the distal portion of the femur is preserved (110 mm; Fig. 5). It is observable only from the medial view. The most distal region is in articulation with the proximal surface of the tibia, covering most of the posterior intercondylar fossa. A deep intercondylar fossa is observable. A marked groove separates the condyles.
It is bulbous and exhibits a lateral ridge
The fibular condyle forms a right angle with the anteroposterior axis of the articulation
The medial surface of the tibia is marked by a fossa
located close to the proximal articulation
the fibular condyle is continuous with the fibular crest
A deep fossa separates the lateral cnemial ridge from the fibular crest
Part of the holotype (MPSC R 2089) of Aratasaurus museunacionali gen
showing the (A) photo and (B) drawing of the right pes
first to second phalanx of pedal digit II; pph1d3
first phalanx of pedal digit III; pph1-3d4
first to third phalanx of pedal digit IV; u1
it is reduced and has the same length as the first phalange of pedal digit I
The proximal articulation is flattened and blade-like
This bone contacts the mid-distal region of the medial surface of metatarsal II
Metatarsals II and IV are morphologically and proportionally similar, being expanded mediolaterally. All exhibit collateral ligament pits. The longest is metatarsal III, which, based on the relation of the foot relative to the tibia, was about 243 mm long (Fig. 4)
The dorsal surface of the distal articulation of metatarsals II and IV are bulbous
The articulation of metatarsal III is markedly ginglymoid
with an extensor pit on the dorsal surface
The collateral ligament pits are present in all metatarsals
being deeper in metatarsal III and shallower in metatarsal IV
Digits II and IV are about the same length. The preserved pedal phalangeal formula is I-1, II-2, III-3 and IV-4 (Fig. 6)
Although most of the phalanges are compressed
some of the ones of digit III were preserved in their original shape and exhibit an ellipsoid cross-section
with digit IV possessing the shortest phalanges compared to the remaining digits
The collateral ligament pits of the phalanges of digits II and III are deep and symmetrical
Although these pits are also deep in digit IV
they exhibit a slight mediolateral asymmetry
being deeper in the lateral side in digit III
The dorsal surface of the proximal articulation of phalanges II-1 and III-1 is bulbous
The distal articulation of phalanges II-1 and III-2 are marked by an extensor pit marks on the dorsal surface
III-2 and III-3 and all of the digit IV show an asymmetric shaft
with the proximal half of the ventral surface showing flexor processes
The phalanges III-2 and III-3 also exhibit a concave ventral surface
Unguals I, II and III are preserved (Fig. 7). Most of the dorsal surface of the ungual I is covered by rock matrix. The ventral surface of all unguals show a faint flexor tubercle. The lateral and medial surfaces of the unguals II and III exhibit ridges, especially in the ungual II.
Pedal unguals (MPSC R 2089) of Aratasaurus museunacionali gen
B) Photo and schematic drawing of the second pedal digit and (C
D) photo and schematic drawing of the third pedal digit
The rounded cnemial crest is also present in Zuolong sallei
The lateral ridge on the cnemial crest is also observed in Zuolong sallei
A rounded fibular condyle and an elongated fibular crest is shared with Aratasaurus museunacionali and Zuolong sallei
This condyle in Australovenator wintonensis presents a ventral convexity
which is different from the flattened surface of Aratasaurus museunacionali
III and IV are about the same in Aratasaurus museunacionali
while Zuolong sallei shows a metatarsal III twice the width of the metatarsals II and IV
the distal articulation of metatarsals II and IV in Aratasaurus museunacionali are similar and differs from the condition of Aarun zhaoi
in which metatarsals II is the widest and tallest among the other metatarsi of the pes
The unguals of both Zuolong sallei and Aratasaurus museunacionali are also similar
presenting flexor tubercles and symmetrical grooves in lateral facets
the material known from the Aratasaurus museunacionali differs from derived coelurosaurian groups (e.g
Ornithomimosauria) and Megaraptora mainly regarding by the cnemial crest and the disposition and morphology of metatarsals
the new Brazilian theropod has a tibia similar to that of Zuolong salleei
and the pes more similar with that of Aarun zhaoi and Tanycolagreus topwilsoni
Osteohistological section of the second metatarsal of Aratasaurus museunacionali gen
showing the four growth cycles (numbers 1–4) marked by two lines of arrested growth and one annulus
po—primary osteons; LAG—lines of arrested growth
Stratigraphic chart modified from Cohen et al
based on the plesiomorphic characters and close relationships with Zuolong salleei
Aratasaurus museunacionali integrates the most basal lineage of Coelurosauria
Life reconstruction of Aratasaurus museunacionali gen
secondary osteons and a high number of primary osteons implies on a juvenile/young adult ontogenetic stage for this animal
The ontogenetic stage attributed to Aratasaurus museunacionali probably explains its reduced proportions
because its asymptotic size was not reached
indicating that this animal could have grown further
traditional search tree bisection and reconnection (TBR) branch swapping with zero random seed
3,000 replicates and 10 saved trees per replication
The obtained trees were reanalyzed in TBR with the parameter “stop when maxtrees hit”
with additional coding for Santanaraptor placidus and Timimus hermani as follows:
Two casts were also made to preserve external morphological data
The bone was sectioned in previous existing breaking area
A bone sample with approximately 1 cm of thickness was obtained
It was embedded in clear epoxy resin Resapol T-208 catalyzed with Butanox M50
cut with a micro rectify (Dremel 4000 with extender cable 225) mounted to a diamond disk
the mounting side was wet ground and polished using a metal polishing machine (AROPOL-E
Abrasive sandpaper of different grits were used in this step (grit size 60 / P60
the section was examined and photographed under a transmitted light microscope (Zeiss Inc
Spain) mounted to an AxioCam camera with Axio Imager
The M2 imaging software was used in the examination procedure
An amendment to this paper has been published and can be accessed via a link at the top of the paper
First Early Cretaceous theropod dinosaur from Brazil with comments on Spinosauridae
Brusatte, S. L., Candeiro, C. R. A. & Simbras, F. M. The last dinosaurs of Brazil: The Bauru Group and its implication for the end-Cretaceous mass extinction. An. Acad. Bras. Ciênc.89, 1465–1485. https://doi.org/10.1590/0001-3765201720160918 (2017)
Machado, E. B., Campos, D. A. & Kellner, A. W. A. On a theropod scapula (Upper Cretaceous) from the Marília Formation, Bauru Group, Brazil. PalZ82, 308–313. https://doi.org/10.1007/BF02988897 (2008)
A new abelisauroid from the upper cretaceous of Brazil
Morphology and internal structure of two new abelisaurid remains (Theropoda
Dinosauria) from the Adamantina Formation (Turonian - Maastrichtian)
Description of uncommon pneumatic structures of a noasaurid (Theropoda
Dinosauria) cervical vertebra from the Bauru Group (Upper Cretaceous)
Bandeira, K. L. N. et al. The Baurusuchidae vs Theropoda record in the Bauru Group (Upper Cretaceous, Brazil): a taphonomic perspective. J. Iber. Geol.44, 25–54. https://doi.org/10.1007/s41513-018-0048-4 (2018)
On the dentition of Baurusuchus pachecoi Price (Crocodyliformes
Metasuchia) from the Upper Cretaceous of Brazil
On a new maniraptoran dinosaur (Theropoda) from the Upper Cretaceous of Neuquén
Ocorrência de um novo crocodiliano no Cretáceo Inferior da Bacia do Araripe
Santana Fossils: An Illustrated Atlas (T.H.F
Guia Para Trabalhos de Campo em Paleontologia na Bacia do Araripe
Vila Nova, B. C., Saraiva, A. A. F., Moreira, J. K. R. & Sayão, J. M. Controlled excavations in the Romualdo Formation Lagerstätte (Araripe Basin, Brazil) and pterosaur diversity: remarks based on new findings. Palaios26, 173–179. https://doi.org/10.2110/palo.2010.p10-072r (2011)
Buchmann, R. & Rodrigues, T. The evolution of pneumatic foramina in pterosaur vertebrae. An. Acad. Bras. Ciênc.91, e20180782. https://doi.org/10.1590/0001-3765201920180782 (2019)
Simões, T., Caldwell, M. W. & Kellner, A. W. A. A new Early Cretaceous lizard species from Brazil, and the phylogenetic position of the oldest known South American squamates. J. Syst. Palaeontol.13, 601–614. https://doi.org/10.1080/14772019.2014.947342 (2015)
Sayão, J. M. et al. Paleohistology of Susisuchus anatoceps (Crocodylomorpha, Neosuchia): Comments on Growth Strategies and Lifestyle. PLoS ONE11, e0155297. https://doi.org/10.1371/journal.pone.0155297 (2016)
Oliveira, G. R. & Kellner, A. W. A. Rare hatchling specimens of Araripemys Price, 1973 (Testudines, Pelomedusoides, Araripemydidae) from the Crato Formation, Araripe Basin. J. S. Am. Earth Sci.79, 137–142. https://doi.org/10.1016/j.jsames.2017.07.014 (2017)
A new crested maniraptoran dinosaur from the Santana Formation (Lower Cretaceous) of Brazil
Coelurosauria) from the Santana Formation (Romualdo Member
Skeletal remains of a small theropod dinosaur with associated soft structures from the Lower Cretaceous Santana Formation of northeastern Brazil
Vertebrate paleontology in Brazil—a review
Kellner, A. W. A. A reconstrução do Museu Nacional: bom para o Rio, bom para o Brasil. Ciência e Cultura71, 4–5. https://doi.org/10.21800/2317-66602019000300001 (2019)
Partes vegetativas de carófitas fossilizadas no Membro Romualdo (Albiano
As condições ecológicas e faciológicas da Formação Santana na Chapada do Araripe (Nordeste do Brasil)
Proposta de Revisão da Coluna Litoestratigráfica da Bacia do Araripe in Anais do Congresso Brasileiro de Geologia
Una Nueva Propuesta Estratigráfica Para La Tecnosecuencia Post-rifte de La Cuenca de Araripe
Martill, D. M. The age of the Cretaceous Santana Formation fossil Konservat Lagerstätte of northeast Brazil: a historical review and an appraisal of the biochronostratigraphic utility of its paleobiota. Cretac Res.28, 895–920. https://doi.org/10.1016/j.cretres.2007.01.002 (2007)
Custódio, M. A. et al. The transgressive-regressive cycle of the Romualdo Formation (Araripe Basin): sedimentary archive of the Early Cretaceous marine ingression in the interior of Northeast Brazil. Sedim. Geol.359, 1–15. https://doi.org/10.1016/j.sedgeo.2017.07.010 (2017)
Sequências deposicionais do Andar Alagoas da Bacia do Araripe
Oliveira, G. R., Saraiva, A. A. F., Silva, H. P., Andrade, J. A. F. G. & Kellner, A. W. A. First turtle from the Ipubi Formation (Early Cretaceous), Santana Group, Araripe Basin, Brazil. Rev. Bras. Paleontol.14, 61–66. https://doi.org/10.4072/rbp.2011.1.06 (2011)
Fabin, C. E. et al. Stratigraphic relations of the Ipubi Formation: siliciclastic–evaporitic succession of the Araripe Basin. An. Acad. Bras. Ciênc.90(2 Suppl. 1), 2049–2071. https://doi.org/10.1590/0001-3765201820170526 (2018)
Lima, F. J. et al. Fire in the paradise: evidence of repeated palaeo-wildfires from the Araripe Fossil Lagerstätte (Araripe Basin, Aptian-Albian), Northeast Brazil. Palaeobio. Palaeoenv.99, 367–378. https://doi.org/10.1007/s12549-018-0359-7 (2019)
Quelques observations sur la palynologie de l'Aptien supérieur et de l'Albien du bassin d'Araripe (N.E
du Brésil) in Atas 1° Simpósio Sobre a Bacia do Araripe e Bacias Interiores do Nordeste (eds
Heimhofer, U. & Hochuli, P.-A. Early Cretaceous angiosperm pollen from a low-latitude succession (Araripe Basin, NE Brazil). Rev. Palaeobot. Palyno.161, 105–126. https://doi.org/10.1016/j.revpalbo.2010.03.010 (2010)
Choiniere, J. N. et al. A juvenile specimen of a new coelurosaur (Dinosauria: Theropoda) from the Middle-Late Jurassic Shishugou Formation of Xinjiang, People’s Republic of China. J. Syst. Palaeontol.12, 177–215. https://doi.org/10.1080/14772019.2013.781067 (2013)
New small theropod from the Upper Jurassic Morrison Formation of Wyoming
New mid-Cretaceous (latest Albian) dinosaurs from Winton
Tyrannosaurus and other Cretaceous carnivorous dinosaurs
Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull
et al.) 137–150 (University of California Press
et al.) 184–195 (University of California Press
Choiniere, J. N., Forster, C. A. & de Klerk, W. J. New information on Nqwebasaurus thwazi, a coelurosaurian theropod from the Early Cretaceous Kirkwood Formation in South Africa. J. Afr. Earth Sci.71–72, 1–17. https://doi.org/10.1016/j.jafrearsci.2012.05.005 (2012)
Delcourt, R. & Grillo, O. N. Reassessment of a fragmentary maxilla attributed to Carcharodontosauridae from Presidente Prudente Formation, Brazil. Cretac. Res.84, 515–524. https://doi.org/10.1016/j.cretres.2017.09.008 (2018)
Sereno, P. The evolution of dinosaurs. Science284, 2137–2147. https://doi.org/10.1126/science.284.5423.2137 (1999)
New Patagonian Cretaceous theropod sheds light about the early radiation of Coelurosauria
a new compsognathoid dinosaur from the Upper Jurassic
The earliest dromaeosaurid theropod from South America
A reappraisal of the Cretaceous non-avian dinosaur faunas from Australia and New Zealand: evidence for their Gondwanan affinities
Note on the paleobiogeography of Compsognathidae (Dinosauria: Theropoda) and its paleoecological implications
Müller, R. T. Are the dinosauromorph femora from the Upper Triassic of Hayden Quarry (New Mexico) three stages in a growth series of a single taxon?. An. Acad. Bras. Ciênc.89, 835–839. https://doi.org/10.1590/0001-3765201720160583 (2017)
Erickson, G. M. Assessing dinosaur growth patterns: a microscopic revolution. Trends. Ecol. Evol.20, 677–684. https://doi.org/10.1016/j.tree.2005.08.012 (2005)
Dinosaur growth patterns and rapid avian growth rates
Parson, W. L. & Parson, K. M. Morphological variations within the Ontogeny of Deinonychus antirrhopus (Theropoda, Dromaeosauridae). PLoS ONE10, e0121476. https://doi.org/10.1371/journal.pone.012147643 (2015)
Canale, J. I., Cerda, I., Novas, F. E. & Haluza, A. Small-sized abelisaurid (Theropoda: Ceratosauria) remains from the Upper Cretaceous of northwest Patagonia, Argentina. Cretac. Res.62, 18–28. https://doi.org/10.1016/j.cretres.2016.02.001 (2016)
Bone histology as a clue in the interpretation of functional adaptations in the Thalattosuchia (Reptilia
Osteohistological evidence for determinate growth in the American Alligator
Andrade, R. C. L. P. & Sayão, J. M. Paleohistology and Lifestyle Inferences of a Dyrosaurid (Archosauria: Crocodylomorpha) from Paraíba Basin (Northeastern Brazil). PLoS ONE9, e102189. https://doi.org/10.1371/journal.pone.0102189 (2014)
A new method to calculate allometric length mass relationships of dinosaurs
Basal Avialae in The Dinosauria (2nd edn) (eds
Padian, K., Horner, J. R. & De Ricqlès, A. Growth in small dinosaurs and pterosaurs: the evolution of archosaurian growth strategies. J. Vert. Paleontol.24, 555–571. https://doi.org/10.1671/0272-4634(2004)024[0555:GISDAP]2.0.CO;2 (2004)
The palaeohistology of pterosaur bone: An overview
Eleutério, L. H. S. et al. Biomechanical and physiological influences on the osteohistological deposition of Anhangueria (Pterosauria, Pterodactyloidea). Rev. Bras. Paleontolog.18, 403–412. https://doi.org/10.4072/rbp.2015.3.06 (2015)
Wang, X. et al. Eggshell and histology provide insight on the life history of a pterosaur with two functional ovaries. An. Acad. Bras. Cienc.87, 1678–2690. https://doi.org/10.1590/0001-3765201520150364 (2015)
Osteohistology of Confuciusornis sanctus (Theropoda: Aves)
Vermiform bones and the evolution of giantism in Megalania—how a reptilian fox became a lion
Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs
Andrade, R. C. L. P. et al. New data about the presence and absence of the External Fundamental System in Archosaurs. Cad. Cult. Cienc.14, 200–211. https://doi.org/10.14295/cad.cult.cienc.v14i1.932 (2015)
Variation in dinosaur skeletochronology indicators: implications for age assessment and physiology
and age estimates in sauropod humeri and femora
Neoceratopsians and ornithomimosaurs: dinosaurs of Gondwanan origin?
TNT version 1.5 including full implementation of phylogenetic morphometrics
In Preparation and Sectioning of Specimens (eds Padian
E.-T.) 55–160 (University of California Press
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We would like to thank Plácido Cidade Nuvens (deceased in 2016) for obtaining the specimen studied here and Bruno C
Vila Nova (UFPE) for preparing part of the material
Diogenes de Almeida Campos (CPRM) and Francisco de Freitas Leite (URCA) are thanked for discussions regarding the name of the new species
acknowledges Lucy Gomes de Souza (Museu Nacional/Universidade Federal do Rio de Janeiro) for helping with the TNT software
The paleoartist Maurilio Oliveira is acknowledged for the life reconstruction of this new dinosaur
This study was partially founded by the Fundação Carlos Chagas Filho de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ #E-26/202.905/2018 to A.W.A.K.)
the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq #420687/2016-5 and #313461/2018-0 to A.W.A.K.; #311715/2017-6 to J.M.S.; #305705/2019-9 to F.J.L.)
the Fundação Cearense de Apoio ao Desenvolvimento Científico e Tecnológico (FUNCAP #BMD-0124-00302.01.01/19 to R.A.M.B.
and the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001 (CAPES #88887.162865/2018-00 to R.A.M.B.)
acknowledges the doctoral fellowship from Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES - PROANTAR: 88887.336584/2019-00 and Finance code 001 CAPES #88887.162865/2019-00 to RAMB)
Laboratório de Paleobiologia e Microestruturas
Juliana Manso Sayão & Rafael Cesar Lima Pedroso de Andrade
Laboratory of Systematics and Taphonomy of Fossil Vertebrates
Museu Nacional/Universidade Federal do Rio de Janeiro
organized the curation and preparation of the specimen
performed the anatomical descriptive research
preformed the phylogenetic analysis; J.M.S
All authors contributed and reviewed the manuscript
The authors declare no competing interests
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