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Crato Formation
Crato Formation
Stratigraphic range: Latest Aptian; ~115–113 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
Type	Geological formation
Unit of	Santana Group
Sub-units	Nova Olinda Member
Underlies	Romualdo & Ipubi Formations
Overlies	Barbalha Formation
Lithology
Primary	Mudstone, limestone
Other	Siltstone
Location
Coordinates	7°06′S 39°42′W / 7.1°S 39.7°W
Approximate paleocoordinates	8°36′S 8°00′W / 8.6°S 8.0°W
Region	Ceará, Pernambuco
Country	Brazil
Extent	Araripe Basin
Type section
Named for	Crato, Ceará
	; Extent of the Santana Group, to which the Crato Formation belongs, in blue

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.^[1]

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

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

The Crato Formation has often historically been considered the lowest member of the Santana Formation (or, alternatively, the Araripina Formation) of the Araripe Group,^[2] later redefined as the Romualdo Formation of the Santana Group.^[3]^[4] 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.^[5]

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.^[5] The Crato Formation is considered time equivalent with the Paracuru Formation.^[6]

Fossil content

Insects

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

Inects of the Crato Formation
Genus	Species	Presence	Description	Images
Araripenymphes^[7]	A. seldoni	Nova Olinda Member	A Cratosmylidae lacewing^[8]
Gracilepteryx	G. pulchra		An Eolepidopterigidae moth
Makarkinia^[9]	M. adamsi M. kerneri		A Kalligrammatid lacewing
Mickoleitia	M. longimanus		A Coxoplectopteran insect	Mickoleitia longimanus
Netoxena	N. nana		An Eolepidopterigidae moth
Principiala^[10]	P. incerta		An Ithonidae lacewing, type species of Principiala
Psamateia	P. calipsa		An Eolepidopterigidae moth
Rafaeliana	R. maxima		Neuropterida incertae sedis
Undopterix	U. cariensis		An Eolepidopterigidae moth
Astraeoptera^[11]	A. vitrea		An Astraeopteridae mayfly.
	A. cretacica
	A. oligovenata
Eosophobia^[11]	E. acuta		An Astraeopteridae mayfly.

Arachnids

Arachnids of the Crato Formation
Genus	Species	Presence	Description	Images
Mesoproctus^[12]	M. rayoli		Whip Scorpion
Protoischnurus	P. axelrodorum		Scorpion
Cratosolpuga	C. wunderlichi		Solifuge

Fish

Fish of the Crato Formation
Genus	Species	Presence	Description	Images	Notes
Araripelepidotes	Araripelepidotes temnurus				^[13]
Belonostomus	Belonostomus sp.				^[14]
Calamopleurus	Calamopleurus cylindricus				^[15]
Cladocyclus	Cladocyclus gardneri		An Ichthyodectidae fish		^[16]
Cratoamia	Cratoamia gondwanica				^[17]
Dastilbe	Dastilbe crandalli				^[18]
Lepidotes	Lepidotes wenzae				^[19]
Placidichthys	Placidichthys bidorsalis				^[20]
Santanichthys	Santanichthys diasii				^[21]

Amphibians

Amphibians of the Crato Formation
Genus	Species	Presence	Description	Images
Arariphrynus^[22]	Arariphrynus placidoi^[22]
Cratia^[22]	Cratia gracilis^[22]
Cratopipa^[23]	Cratopipa novaolindensis
Eurycephalella^[22]	Eurycephalella alcinae^[22]
Kururubatrachus^[24]	Kururubatrachus gondwanicus
Pipoidea^[22]	Possible indeterminate pipoid remains.^[22]

Squamata

Squamates of the Crato Formation
Genus	Species	Presence	Description	Images
Calanguban	C. alamoi		A non-iguana lizard
Tetrapodophis	T. amplectus		A stem group snake with limbs

Dinosaurs

Dinosaurs of the Crato Formation
Genus	Species	Presence	Notes	Images
?Avialae	?Avialan species		Numerous isolated feathers
Cratoavis	C. cearensis		An enantiornithine
Kaririavis^[25]	K. mater		An ornithuromorph
?Spinosauroidea	?Spinosaur species		Isolated tooth listed in a book appendix.^[5] It has later been suggested that the lithostratigraphic provenance of this specimen might have been incorrect.^[26]
"Ubirajara"	"U. jubatus"		An informally-named compsognathid known from a partial skeleton preserving integument
Sauropoda			Sauropoda footprints.^[27]

Crocodylomorphs

Crocodylomorphs of the Crato Formation
Genus	Species	Presence	Description	Images
Susisuchus	Susisuchus anatoceps^[28]
Susisuchus	cf. Susisuchus sp.^[29]		Undescribed species

Pterosaurs

Pterosaurs of the Crato Formation
Genus	Species	Presence	Description	Images
Arthurdactylus	A. conandoylei
Aymberedactylus	A. cearensis		A basal member of the Tapejarinae.	Holotype
Brasileodactylus	B. sp.
Lacusovagus	L. magnificens	Nova Olinda Member
Ludodactylus	L. sibbicki		An anhanguerid	Ludodactylus sp.
Tupandactylus	T. imperator T. navigans			Tupandactylus navigans
?Tupuxuara	?T. sp.			Tupuxuara sp.

Flora

Flora of the Crato Formation
Species	Notes
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.	^[30]

Other fossils

Araripenaeus timidus^[31]
†Cratoalloneura
- †Cratoalloneura acuminata
- †Cratoalloneura verdandia
†Cratoatractocerus
- †Cratoatractocerus grimaldii
†Cratoborellia
- †Cratoborellia gorbi
†Cratochrysa
- †Cratochrysa martinsnetoi
- †Cratochrysa sublapsa
- †Cratochrysa willmanni
†Cratocora
- †Cratocora crassa
†Cratocordulia
- †Cratocordulia borschukewitzi
†Cratocoris
- †Cratocoris shevchenkoae
†Cratocorydalopsis
- †Cratocorydalopsis brasiliensis
†Cratocossus
- †Cratocossus magnus
†Cratodactylus
- †Cratodactylus ferreirai
- †Cratodactylus kellneri
†Cratoelcana
- †Cratoelcana damianii
- †Cratoelcana zessini
†Cratoenigma
- †Cratoenigma articulata
†Cratogomphus
- †Cratogomphus erraticus
†Cratogryllus
- †Cratogryllus cigueli
- †Cratogryllus guimaraesae
- †Cratogryllus pentagonalis
†Cratohagenius
- †Cratohagenius erichweberi
†Cratohaglopsis
- †Cratohaglopsis santanaensis
†Cratohexagenites
- †Cratohexagenites longicercus
- †Cratohexagenites minor
†Cratokalotermes
- †Cratokalotermes santanensis
†Cratolindenia
- †Cratolindenia knuepfae
†Cratolocustopsis
- †Cratolocustopsis araripensis
- †Cratolocustopsis contumax' – type locality for species
- †Cratolocustopsis cretacea
†Cratomacer
- †Cratomacer ephippiger
- †Cratomacer immersus
†Cratomastotermes
- †Cratomastotermes wolfschwenningeri
†Cratomyia
- †Cratomyia cretacica
- †Cratomyia macrorrhyncha
†Cratonemonyx
- †Cratonemonyx martinsnetoi
†Cratonemopteryx
- †Cratonemopteryx audax
- †Cratonemopteryx robusta
- †Cratonemopteryx speciosa
†Cratonepa
- †Cratonepa enigmatica
†Cratonerthra
- †Cratonerthra corinthiana
- †Cratonerthra estevezae
†Cratoneura
- †Cratoneura dividens
- †Cratoneura longissima
- †Cratoneura pulchella
†Cratonympha
- †Cratonympha microcelata
†Cratopelocoris
- †Cratopelocoris carpinteroi
†Cratopetalia
- †Cratopetalia whiteheadi
†Cratopetalura
- †Cratopetalura petruleviciusi
- †Cratopetalura petruleviciusi
†Cratopsychopsis
- †Cratopsychopsis maiseyi
†Cratopteryx
- †Cratopteryx nemopteroides
- †Cratopteryx robertosantosi
†Cratoraricrus
- †Cratoraricrus oberlii
†Cratoscalapha
- †Cratoscalapha electroneura
†Cratosirex
- †Cratosirex sennlaubi
†Cratosisyrops
- †Cratosisyrops gonzagai
†Cratosmylus
- †Cratosmylus magnificus
†Cratostenophlebia
- †Cratostenophlebia schwickerti
†Cratotabanus
- †Cratotabanus stonemyomorphus
†Cratotetraspinus
- †Cratotetraspinus fossorius
†Cratotipula
- †Cratotipula latialata
†Cratovitisma
- †Cratovitisma oldreadi
†Cratovoluptia
- †Cratovoluptia criptoneura
†Cratozeunerella
- †Cratozeunerella amedegnatoi
- †Cratozeunerella godoii
- †Cratozeunerella neotropica
- †Cratozeunerella nervosa
- †Cratozeunerella soaresi
- †Cratozeunerella titanella

References

^ Ribeiro et al., 2021
^ Scherer et al., 2013, p. 28. Retrieved 2018-10-05.
^ Assine, 1992, p. 291. Retrieved 2018-10-05.
^ Fabin et al., 2018, p. 2050. Retrieved 2018-10-05.
^ ^a ^b ^c Martill et al., 2007. Retrieved 2018-10-06.
^ Leite da Silva, 2003. Retrieved 2018-10-06.
^ Myskowiak, 2016
^ Makarkin, Vladimir N.; Heads, Sam W.; Wedmann, Sonja (October 2017). "Taxonomic study of the Cretaceous lacewing family Babinskaiidae (Neuroptera: Myrmeleontoidea: Nymphidoidae), with description of new taxa". Cretaceous Research. 78: 149–160. doi:10.1016/j.cretres.2017.06.007. ISSN 0195-6671.
^ Bechly et al., 2016. Retrieved 2018-10-06.
^ Makarkin & Menon, 2007. Retrieved 2018-10-06.
^ ^a ^b Storari, Arianny P.; Staniczek, Arnold H.; Godunko, Roman J. (2023-07-20). "A new Gondwanan mayfly family from the Lower Cretaceous Crato Formation, Brazil (Ephemeroptera: Siphlonuroidea: Astraeopteridae fam. nov.)". Scientific Reports. 13 (1): 11735. Bibcode:2023NatSR..1311735S. doi:10.1038/s41598-023-36778-x. ISSN 2045-2322. PMC 10359310. PMID 37474555.
^ Santana, W.; Pinheiro, A. P.; Silva, T. A.; Lima, D. (2024). "Description of a new fossil Thelyphonida (Arachnida, Uropygi) and further record of Cratosolpuga wunderlichi Selden, in Selden and Shear, 1996 (Arachnida, Solifugae) from Crato Formation (Aptian/Albian), Araripe Basin, Brazil". PeerJ. 12. e16670. doi:10.7717/peerj.16670. PMC 10771091. PMID 38188171.
^ Aparecida et al., 2015, p. 25
^ Aparecida et al., 2015, p. 35
^ Aparecida et al., 2015, p. 29
^ Aparecida et al., 2015, p. 36
^ Aparecida et al., 2015, p. 31
^ Aparecida et al., 2015, p. 51
^ Aparecida et al., 2015, p. 26
^ Aparecida et al., 2015, p. 33
^ Aparecida et al., 2015, p. 47
^ ^a ^b ^c ^d ^e ^f ^g ^h Báez et al., 2009. Retrieved 2018-10-06.
^ Ismar de Souza Carvalho; Federico Agnolin; Mauro A. Aranciaga Rolando; Fernando E. Novas; José Xavier-Neto; Francisco Idalécio de Freitas; José Artur Ferreira Gomes de Andrade (2019). "A new genus of pipimorph frog (Anura) from the Early Cretaceous Crato formation (Aptian) and the evolution of South American tongueless frogs". Journal of South American Earth Sciences. 92: 222–233. Bibcode:2019JSAES..92..222C. doi:10.1016/j.jsames.2019.03.005. hdl:11336/123956. S2CID 134070810.
^ Federico Agnolin; Ismar Souza Carvalho; Alexis M. Aranciaga Rolando; Fernando E. Novas; José Xavier-Neto; José Artur Ferreira Gomes Andrade; Francisco Idalécio Freitas (2020). "Early Cretaceous neobatrachian frog (Anura) from Brazil sheds light on the origin of modern anurans". Journal of South American Earth Sciences. 101: Article 102633. Bibcode:2020JSAES.10102633A. doi:10.1016/j.jsames.2020.102633. S2CID 218957427.
^ de Souza Carvalho et al., 2021
^ Caetano, João Marcus Vale; Delcourt, Rafael & Ponciano, Luiza Corral Martins de Oliveira (March 2023). "A taxon with no name: Ubirajara jubatus (Saurischia: Compsognathidae) is an unavailable name and has no nomenclatural relevance". Zootaxa. 5254 (3): 443–446. doi:10.11646/zootaxa.5254.3.10. PMID 37044710.
^ Ismar de Souza, Carvalho Vale; Leonardi, Giuseppe; Rios-Netto, Aristoteles de Moraes; Borghi, Leonardo; Freitas, Alexandre de Paula; Andrade, José Artur & Freitas, Francisco Idalécio (2021). "Dinosaur trampling from the Aptian of Araripe Basin, NE Brazil, as tools for paleoenvironmental interpretation". Cretaceous Research. 117: 104626. Bibcode:2021CrRes.11704626C. doi:10.1016/j.cretres.2020.104626.
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^ Jorge de Lima et al., 2015, p.102
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