The Crato Formation is a geologic formation of Early Cretaceous (Aptian) age in northeastern Brazil's Araripe Basin. It is an important Lagerstätte (undisturbed fossil accumulation) for palaeontologists. The strata were laid down mostly during the Aptian age, about 113 million years ago. It thought to have been deposited in a semi-arid lacustrine wetland environment.

The Crato Formation earns the designation of Lagerstätte due to an exceedingly well preserved and diverse fossil faunal assemblage. Some 25 species of fossil fishes are often found with stomach contents preserved, enabling paleontologists to study predator-prey relationships in this ecosystem. There are also fine examples of pterosaurs, reptiles and amphibians, invertebrates (particularly insects), and plants. Even dinosaurs are represented: a new maniraptor was described in 1996. The unusual taphonomy of the site resulted in limestone accretions that formed nodules around dead organisms, preserving even soft parts of their anatomy.

History

Schematic development of the depositional environments of the Santana Group

Fish fossils in the area were noted in 1823. When they were first methodically published, in 1993, the Crato Formation limestones provided a new site for pterosaurs, one that also preserved insects that fell into a brackish lagoon and semionotid fish preserved in phosphatized nodules. The fossils are usually compacted and preserved in layers of limestone. Fossil Odonata (dragonflies) and damselflies are especially rich in the Crato lagerstätte: currently 384 specimens have been recovered, 264 adults and 120 larvae. Hemiptera (true bugs) and Orthoptera (grasshoppers and crickets) are also abundant in number of species and in number of specimens. There are also plant remains.

Local mining activities for cement and construction damage the sites. Trade in illegally collected fossils has sprung up in the last decade, driven by the remarkable state of preservation and beauty of these fossils and amounting to a considerable local industry. An urgent preservation program is being called for by paleontologists.

In addition, the weathering of Crato and Santana Formation rocks has contributed soil conditions unlike elsewhere in the region. The Araripe manakin (Antilophia bokermanni) is a very rare bird that was discovered only in the late 20th century; it is not known from anywhere outside the characteristic forest that grows on the Chapada do Araripe soils formed ultimately from Crato and Santana Formation rocks.

Definition

Lake level cyclicity in the Santana Group

The Crato Formation has often historically been considered the lowest member of the Santana Formation (or, alternatively, the Araripina Formation) of the Araripe Group, later redefined as the Romualdo Formation of the Santana Group. The Crato Member is the product of a single phase, where complicated sequence of sediment strata reflect changeable conditions in the opening sea. The age of this strata has been controversial, though most workers have agreed that it lies on or near the Aptian-Albian boundary, about 112 million years ago.

The extent of the Crato unit and its relationship to the Romualdo Formation had long been ill-defined. It was not until a 2007 volume on the unit by Martill, Bechly and Loveridge that the Crato Formation was given a formal type locality, and was formally made a distinct formation separate from the Santana, which is about 10 Ma younger. The Crato Formation is considered time equivalent with the Paracuru Formation.

Fossil content

Insects

(Note: Many more insects have been described than are present in the table below)

Insects of the Crato Formation
GenusSpeciesPresenceDescriptionImages
AraripenymphesA. seldoniNova Olinda MemberA Cratosmylidae lacewing
AestuacridaA. stereofemorisNova Olinda MemberA Locustopsid Grasshopper
A. mikronaulionNova Olinda MemberA Locustopsid Grasshopper
CratomastaxC. mariellaaeNova Olinda MemberA Eumastacoid Grasshopper
GracilepteryxG. pulchraAn Eolepidopterigidae moth
MakarkiniaM. adamsi M. kerneriA Kalligrammatid lacewing
MickoleitiaM. longimanusA Coxoplectopteran insectMickoleitia longimanus
NetoxenaN. nanaAn Eolepidopterigidae moth
PrincipialaP. incertaAn Ithonidae lacewing, type species of Principiala
PsamateiaP. calipsaAn Eolepidopterigidae moth
RafaelianaR. maximaNeuropterida incertae sedis
UndopterixU. cariensisAn Eolepidopterigidae moth
AstraeopteraA. vitreaAn Astraeopteridae mayfly.
A. cretacica
A. oligovenata
EosophobiaE. acutaAn Astraeopteridae mayfly.
VulcanidrisV. cratensisA hell ant

Arachnids

Arachnids of the Crato Formation
GenusSpeciesPresenceDescriptionImages
MesoproctusM. rayoliWhip Scorpion
ProtoischnurusP. axelrodorumScorpion
CratosolpugaC. wunderlichiSolifuge

Fish

Fish of the Crato Formation
GenusSpeciesPresenceDescriptionImagesNotes
AraripelepidotesAraripelepidotes temnurus
BelonostomusBelonostomus sp.
CalamopleurusCalamopleurus cylindricus
CladocyclusCladocyclus gardneriAn Ichthyodectidae fish
CratoamiaCratoamia gondwanica
DastilbeDastilbe crandalli
LepidotesLepidotes wenzae
PlacidichthysPlacidichthys bidorsalis
SantanichthysSantanichthys diasii

Amphibians

Amphibians of the Crato Formation
GenusSpeciesPresenceDescriptionImages
ArariphrynusArariphrynus placidoi
CratiaCratia gracilis
CratopipaCratopipa novaolindensis
EurycephalellaEurycephalella alcinae
KururubatrachusKururubatrachus gondwanicus
PipoideaPossible indeterminate pipoid remains.

Squamata

Squamates of the Crato Formation
GenusSpeciesPresenceDescriptionImages
CalangubanC. alamoiA non-iguana lizard
TetrapodophisT. amplectusA stem group snake with limbs

Dinosaurs

Dinosaurs of the Crato Formation
GenusSpeciesPresenceNotesImages
?Avialae?Avialan speciesNumerous isolated feathers
CratoavisC. cearensisAn enantiornithine
KaririavisK. materAn ornithuromorph
?Spinosauroidea?Spinosaur speciesIsolated tooth listed in a book appendix. It has later been suggested that the lithostratigraphic provenance of this specimen might have been incorrect.
"Ubirajara""U. jubatus"An informally-named compsognathid known from a partial skeleton preserving integument
SauropodaSauropoda footprints.

Crocodylomorphs

Crocodylomorphs of the Crato Formation
GenusSpeciesPresenceDescriptionImages
SusisuchusSusisuchus anatoceps
cf. Susisuchus sp.Undescribed species

Pterosaurs

Pterosaurs of the Crato Formation
GenusSpeciesPresenceDescriptionImages
ArthurdactylusA. conandoylei
AymberedactylusA. cearensisNova Olinda MemberA basal member of the Tapejarinae.Holotype
BrasileodactylusB. sp.Likely represents a specimen of Ludodactylus
LacusovagusL. magnificensNova Olinda Member
LudodactylusL. sibbickiAn anhangueridLudodactylus sp.
TupandactylusT. imperator T. navigansTupandactylus navigans
?Tupuxuara?T. sp.Tupuxuara sp.
Chaoyangopteridae indet.IndeterminateNova Olinda MemberUFC 721, a series of four articulated cervical vertebrae; may be referrable to Lacusovagus
Tapejaridae indet.Indeterminate
Azhdarchidae indet.IndeterminateNova Olinda MemberSMNK 2342 PAL, a partially articulated, distal portion of a left wing; may be referrable to Lacusovagus

Flora

Flora of the Crato Formation
SpeciesNotes
Araucaria cartellei, Brachyphyllum obesum, B. castilhoi, B. insigne, Iara iguassu, Caytoniales sp., Ephedra sp., Araripia florifera, Araucarites vulcanoi, Cariria orbiculiconiformis, Cearania heterophylla, Cratonia cotyledon, Endressinia brasiliana,Klitzchophyllites flabellatus, Novaolindia dubia, Pluricarpellatia peltata, Podozamites lanceolatus, Protananas lucenae, Ruffordia goeppertii, Tomaxellia biforme, Welwitschiaprisca austroamericana, Welwitschiophyllum brasiliense, Welwitschiostrobus murili, Araucariostrobus sp., Frenelopsis sp., Isoetites sp., Lindleycladus sp., Schizoneura sp.

Fungi

Fungi of the Crato Formation
GenusSpeciesPresenceDescriptionImages
GondwanagaricitesGondwanagaricites magnificus
EdaphagaricitesEdaphagaricites conicus

Other fossils

  • Araripenaeus timidus
  • CratoalloneuraCratoalloneura acuminataCratoalloneura verdandia
  • CratoatractocerusCratoatractocerus grimaldii
  • CratoborelliaCratoborellia gorbi
  • CratochrysaCratochrysa martinsnetoiCratochrysa sublapsaCratochrysa willmanni
  • CratocoraCratocora crassa
  • CratocorduliaCratocordulia borschukewitzi
  • CratocorisCratocoris shevchenkoae
  • CratocorydalopsisCratocorydalopsis brasiliensis
  • CratocossusCratocossus magnus
  • CratodactylusCratodactylus ferreiraiCratodactylus kellneri
  • CratoelcanaCratoelcana damianiiCratoelcana zessini
  • CratoenigmaCratoenigma articulata
  • CratogomphusCratogomphus erraticus
  • CratogryllusCratogryllus cigueliCratogryllus guimaraesaeCratogryllus pentagonalis
  • CratohageniusCratohagenius erichweberi
  • CratohaglopsisCratohaglopsis santanaensis
  • CratohexagenitesCratohexagenites longicercusCratohexagenites minor
  • CratokalotermesCratokalotermes santanensis
  • CratolindeniaCratolindenia knuepfae
  • CratolocustopsisCratolocustopsis araripensisCratolocustopsis contumax' – type locality for species †Cratolocustopsis cretacea
  • CratomacerCratomacer ephippigerCratomacer immersus
  • CratomastotermesCratomastotermes wolfschwenningeri
  • CratomyiaCratomyia cretacicaCratomyia macrorrhyncha
  • CratonemonyxCratonemonyx martinsnetoi
  • CratonemopteryxCratonemopteryx audaxCratonemopteryx robustaCratonemopteryx speciosa
  • CratonepaCratonepa enigmatica
  • CratonerthraCratonerthra corinthianaCratonerthra estevezae
  • CratoneuraCratoneura dividensCratoneura longissimaCratoneura pulchella
  • CratonymphaCratonympha microcelata
  • CratopelocorisCratopelocoris carpinteroi
  • CratopetaliaCratopetalia whiteheadi
  • CratopetaluraCratopetalura petruleviciusiCratopetalura petruleviciusi
  • CratopsychopsisCratopsychopsis maiseyi
  • CratopteryxCratopteryx nemopteroidesCratopteryx robertosantosi
  • CratoraricrusCratoraricrus oberlii
  • CratoscalaphaCratoscalapha electroneura
  • CratosirexCratosirex sennlaubi
  • CratosisyropsCratosisyrops gonzagai
  • CratosmylusCratosmylus magnificus
  • CratostenophlebiaCratostenophlebia schwickerti
  • CratotabanusCratotabanus stonemyomorphus
  • CratotetraspinusCratotetraspinus fossorius
  • CratotipulaCratotipula latialata
  • CratovitismaCratovitisma oldreadi
  • CratovoluptiaCratovoluptia criptoneura
  • CratozeunerellaCratozeunerella amedegnatoiCratozeunerella godoiiCratozeunerella neotropicaCratozeunerella nervosaCratozeunerella soaresiCratozeunerella titanella

See also

Bibliography

  • Aparecida dos Reis Polck, Márcia; Sardenberg Salgado de Carvalho, Marise; Miguel, Raphael; Gallo, Valéria (2015). Guia de identificação de peixes fósseis das Formações Crato e Santana da Bacia do Araripe. Serviço Geológico do Brasil (CPRM). pp. 1–74.
  • Assine, Mario L (1992). (PDF). Revista Brasileira de Geociências. 22 (3): 289–300. doi:.
  • Báez, A.M.; Moura, G.J.B.; Gómez, R.O. (2009). "Anurans from the Lower Cretaceous Crato Formation of northeastern Brazil: implications for the early divergence of neobatrachians". Cretaceous Research. 30 (4): 829–846. Bibcode:. doi:.
  • Bechly, Günter; Makarkin, Vladimir N. (March 2016). "A new gigantic lacewing species (Insecta: Neuroptera) from the Lower Cretaceous of Brazil confirms the occurrence of Kalligrammatidae in the Americas". Cretaceous Research. 58: 135–140. Bibcode:. doi:.
  • de Souza Carvalho, I.; Agnolin, F. L.; Rozadilla, S.; Novas, F. E.; Ferreira Gomes Andrade, J. A.; Xavier-Neto, J. (2021). "A new ornithuromorph bird from the Lower Cretaceous of South America". Journal of Vertebrate Paleontology. 41 (4) e1988623. Bibcode:. doi:. S2CID . S2CID
  • Fabin, Carlos E.; Correia Filho, Osvaldo J.; Alencar, Márcio L.; Barbosa, José A.; de Miranda, Tiago S.; Neumann, Virgínio H.; Gomes, Igor F.; de Santana, Felipe R. (2018). (PDF). Anais da Academia Brasileira de Ciências. 90 (2 suppl 1): 2049–2071. doi:. PMID .
  • Figueiredo, R.G.; Kellner, A.W.A. (2009). . Paläontologische Zeitschrift. 83 (2): 323–331. Bibcode:. doi:. S2CID .
  • Jorge de Lima, Flaviana; Feitosa Saraiva, Antonio Álamo; Manso Sayão, Juliana (2012). . Estudos Geológicos. 22 (1): 99–115. Bibcode:. doi:.
  • Leite da Silva, Agnelo; Neumann, Virginio Enrique (2003). (PDF). 2o Congresso Brasileiro de P&D em Petróleo & Gás. pp. 1–6.
  • Makarkin, Vladimir N.; Menon, Federica (October 2007). "First record of fossil 'rapismatid-like' Ithonidae (Insecta, Neuroptera) from the Lower Cretaceous Crato Formation of Brazil". Cretaceous Research. 28 (5): 743–753. Bibcode:. doi:.
  • Martill, David M.; Bechly, Günter; Loveridge, Robert F. (2007). . Cambridge University Press. p. 236. ISBN 978-1-139-46776-6.
  • Pinheiro, Allysson P.; Saraiva, Antônio Á.F.; Santana, William (2014). (PDF). Anais da Academia Brasileira de Ciências. 86 (2): 1–8. doi:. PMID . Archived from (PDF) on 2017-08-24.
  • Ribeiro, Alexandre Cunha; Ribeiro, Guilherme Cunha; Varejão, Filipe Giovanini; Battirola, Leandro Dênis; Pessoa, Edlley Max; Simões, Marcello Guimarães; Warren, Lucas Veríssimo; Riccomini, Claudio; Poyato-Ariza, Francisco José (May 2021). "Towards an actualistic view of the Crato Konservat-Lagerstätte paleoenvironment: A new hypothesis as an Early Cretaceous (Aptian) equatorial and semi-arid wetland". Earth-Science Reviews. 216 103573. Bibcode:. doi:. hdl:. S2CID .
  • Salisbury, S.W.; Frey, E.; Martill, D.M.; Buchy, M.C. (2003). . Palaeontographica, Abteilung A. 270 (1–3): 3–47. Bibcode:. doi:. S2CID .
  • Scherer, C.M.d.S.; Jardim de Sá, E.F.; Córdoba, V.C.; Sousa, D.d.C.; Aquino, M.M.; Cardoso, F.M.C. (2013). (PDF). Journal of South American Geosciences. 49: 1–43. Bibcode:. doi:. hdl:.
  • Weishampel, David B.; et al. (2004). Dinosaur distribution (Early Cretaceous, South America)in:Weishampel, David B.; Dodson, Peter; and Osmólska, Halszka (eds.): The Dinosauria. Berkeley: University of California Press. pp. 563–570. ISBN 0-520-24209-2.{{cite book}}: CS1 maint: publisher location (link)

Further reading

  • Bétard, François; Peulvast, Jean-Pierre; de Oliveira Magalhães, Alexsandra; Carvalho Neta, Maria de Lourdes; de Freitas, Francisco Idalecio (2017). (PDF). Geoheritage. 10 (4): 1–18. Bibcode:. doi:. Archived from (PDF) on 2018-08-21.
  • Neumann, V.H.; Borrego, A.G.; Cabrera, L.; Dino, R. (April 2003). "Organic matter composition and distribution through the Aptian–Albian lacustrine sequences of the Araripe Basin, northeastern Brazil". International Journal of Coal Geology. 54 (1–2): 21–40. Bibcode:. doi:.
  • Jouault, Corentin; Pouillon, Jean-Marc; Nel, André (31 August 2020). "The first fossil horntail wasp (Hymenoptera: Siricidae) from Lower Cretaceous Crato Formation in Brazil". Palaeoentomology. 3 (4): 382–389. Bibcode:. doi:.
  • Nel, André; Pouillon, Jean-Marc (30 June 2020). "The second genus of the 'libelluloid' family Araripephlebiidae (Odonata, Clavilabiata)". Palaeoentomology. 3 (3): 240–244. Bibcode:. doi:.