Tlayúa Formation
Geological formation
PeriodCretaceous
PrilithologyLimestone
OtherlithologyMudstone
NamedforTlayúa Quarry
RegionPuebla
CountryMexico
Coordinates19.1°N 98.2°W
Paleocoordinates17.1°N 61.6°W
SubunitsLower, Middle and Upper Members
UnderliesUnconformity with Eocene-Oligocene Pie de Vaca Formation
OverliesOrizaba Formation
ThicknessLower: 50 m Middle: 35 m Upper: 40 m

The Tlayúa Formation is an Cretaceous Konservat-Lagerstätte near Tepexi de Rodríguez, Puebla, Central Mexico.[1] It consists of three members (Lower, Middle and Upper), spanning the lower Albian of the Early Cretaceous to the lower Cenomanian of the Late Cretaceous. It consists of a series of limestone quarries that preserve lagoonal palaeoenvironments, such as a shelf lagoon, a shallow lagoon surrounded by a peneplain, and a tidal flat. It is notable for preserving high quantities of vertebrate and invertebrate fossils, and is thus considered a Lagerstätte.

Description

The Tlayúa Formation was first described by Jerjes Pantoja-Alor in 1992.[2] It is located in the Tlayúa ravine, which itself lies in the southern portion of Puebla, Central Mexico, near Tepexi de Rodríguez,[1] and consists of a series of limestone quarries that has seen commercial use for decades.[3]

Stratigraphy

The Tlayúa Formation consists of two stratigraphic units (Tlayúa and Barranca Abuelo)[4] and three members. The Lower Member is lower Albian in age. It consists of bioturbated limestones, and is characterised by the presence of the non-rudist bivalve Chondrodonta and the rudist bivalve Toucasia polygyra. The Middle Member, corresponding to the Tlayúa Quarry, is the most fossiliferous.[3] Its age has been difficult to ascertain, as the foraminiferan Spiroloculina cretacea is known exclusively from lower Cenomanian strata,[5] though it has since been determined to be Upper Albian.[3] The Middle Member consists of fine-grained lithographic limestones, interspersed with hematitic layers that preserve its vertebrate fauna. The Upper Member is Cenomanian in age, and consists of a sequence of dolomites. It is characterised by the presence of the miliolid foraminiferan Dicyclina schlumbergi.[3]

Depositional environments

The depositional environment of the Tlayúa Formation was likely a shallow, coastal lagoon,[6] with some freshwater influence, as indicated by the presence of crocodilian and turtle fossils.[6][7] The Lower Member of the Tlayúa Formation represents a carbonate environment which apparently had stressed biodiversity.[4] The depositional environment of the Middle Member was likely arid for much of the year, with the exception of seasonal rains and storms.[7] The Upper Member likely represents a tidal flat.[4] The Tlayúa Formation may have been part of an island, though a connection to the North American mainland cannot be discounted.[7]

Paleobiota

The formation contains a diverse array of vertebrate and invertebrate fossils. For this reason, it is considered a Konservat-Lagerstätte.[4][7] About 70% of the macrofossils are osteichthyan fish.[8] Other vertebrates include chelonians, pterosaurs, lepidosaurs, and crocodiles.[9] Cyanobacteria, foraminifera, algae, gymnosperms, sponges, cnidarians, annelids, gastropods, ammonites, bivalves, arachnids, insects, isopods, anomurans, brachyurans, crinoids, echinoids, holothuroids, stelleroids, and ophiuroids, have also been recovered from the Tlayúa Formation.[9][10][11]

Archelosaurs

The remains of several indeterminate archelosaur taxa have been recovered from the Tlayúa Formation. Indeterminate crocodilians and turtles have been identified.[6][7] Partial, articulated wings of an unidentified pterosaur have been discovered from the formation. Though tentatively assigned to Pteranodon sp. and Nyctosaurus sp. in 2008,[12] they likely represent either an early azhdarchoid or an indeterminate ornithocheiromorph.[13]

Lepidosaurs

GenusSpeciesMaterialNotes
Huehuecuetzpalli[14]H. mixtecusNear-complete specimens of an adult and a juvenileA primitive lizard
Pamizinsaurus[15]P. tlayuaensisA single, crushed skull of a juvenileAn osteoderm-covered sphenodontian
Ankylosphenodon[16]A. pachyostosusFront half of a partial skeletonAn aquatic sphenodontian
Tepexisaurus[17]T. tepexiiNear-complete skeleton, minus the tailA basal scincomorph

Fish

Mainly after González-Rodríguez (2016)[18] and Applegate et al. (2006)[3]

Fish
GenusSpeciesHigher taxonNotesImages
Amblysemius?cf. A. spCaturidaeOnly non-amiid amiiform from Tlayúa[18]
AraripichthysA. weberiTeleosteiPossibly related to Late Cretaceous genera from North America?[19]
ArmigatusA. felixi, A. carrhenoaeEllimmichthyiformesFirst records of Armigatus from America, alongside the oldest species in the genus,[20] the two species from Tlayúa also seem to clade closest to each other[21]
Aspidorhynchidae spp.InapplicableTeleosteiUndescribed species of Vinctifer and Belonostomus have been recorded[18]
AxelrodichthysA. cf. A. araripensisMawsoniidaeOnly known specimen was lost before description[18]
Bananogmius?cf. B. spTselfatiiformesOne of the few Mexican members of the clade other than Tselfatia[18]
CipactlichthysC. scutatusHalecomorphiBears large plate-like scutes along its midline[22]
EllimmichthysE. sp[23]EllimmichthyiformesType genus of its order
Elopomorpha spp.UnapplicableTeleosteiPossible species of Brannerion, Paraelops and Megalops are known[18]
EpaelopsE. martineziElopiformesRelated to the Jurassic Anaethalion[24]
Gonorynchiformes indet.UnapplicableOstariophysiOne specimen reported in the 1990s, but has not been studied since[23]
Lycoptera?cf. L. spLycopteridaeOnly osteoglossiform fossil from Cretaceous Mexico[18]
MacrosemiocotzusM. americanusMacrosemiidaeSimilar to Macrosemius, but differs in a divided dorsal fin[25]
MichinM. csernaiPachyrhizodontidaeLikely the most basal pachyrhizodontid[26]
NeoproscinetesN. sp[18]PycnodontidaeRelated to Proscinetes
NotagogusN. novomundiMacrosemiidaeClosest to Notagogus helenae[27]
NusaviichthysN. nerivelaiPachyrhizodontoideiFormerly misidentified as Notelops fossils[28]
PachyamiaP. mexicanaVidalamiinaeAlso known from the Middle East[29]
ParaclupeaP. seilacheriEllimmichthyiformesFirst fossil of the genus outside of China[30]
RanulfoichthysR. dorsonudusClupeomorphaSomewhat resembles Ellimmichthyiformes, but lacks their dorsal scutes, therefore it is likely a basal clupeomorph[31]
QuetzalichthysQ. perilliataeIonoscopiformes (Halecomorphi)Likely closest to Oshunia[32]
TahnaichthysT. magnuserrataPycnodontidaeBears two humps in front of its dorsal fin, a novel trait in pycnodonts[33]
TeoichthysT. kallistos, T. brevipina[34]Ophiopsidae (Halecomorphi)Name of the type species (Teoichthys kallistos) translates to "God's most beautiful fish", in reference to the exquisite preservation[35]
TepexichthysT. aranguthyorumPycnodontidaeLikely fed on coral, as shown by gut contents and the large amount of wear on its teeth[36]
TlalocbatosT. applegateiRhinopristiformesOnly chondrichthyan known from Tlayúa[37]
TlayuamichinT. iztliSemionotiformesSpecific name translates to "obsidian", after the colour of the scales[38]
UnamichthysU. espinosaiIchthyodectiformesDiscovered alongside two other currently undescribed ichthyodectiforms[39]
Yabrudichthys?cf. Y. sp[18]AulopiformesLikely an enchodontid, mainly known from the Ein Yabrud site (hence its generic name)

Arthropods

Arthropods
GenusSpeciesHigher taxonNotesImages
ArchaeoniscusA. aranguthyorumSphaeromatidea?Bears an unusual axial pouch unlike all extant isopods, may have been a fish ectoparasite or scavenger like modern cymothoids[40]
AtocatleA. ranulfoiMesothelae?Indeterminate family due to poor preservation, likely within Mesothelae due to its distinct abdominal segments,[40] but more recently its affinity as a spider is questioned[41]
IxtahuaI. benjaminiAnisoptera?Only known from a nymph, matches dragonfly nymphs in shape[40]
ProtaeglaP. minisculaAeglidaeEarliest aeglid fossil known[40]
TepexicarcinusT. tlayuaensisTepexicarcinidae (basal Eubrachyura)Formerly assigned questionably to Homolidae due to an absence of clear carapace regions,[40] but then separated into its own family as a basal eubrachyuran[42]
Tipulidae indet.UnapplicableTipuloideaOnly known from a wing pair[40]

Echinoderms

Arthropods
GenusSpeciesHigher taxonNotesImages
AstropectenA. spAstropectinidaeKnown from juvenile fossils preserving both dorsal and ventral sides[43]
EchinasterE. spEchinasteridaeOnly known from a fossil in dorsal view[43]
OphiactisO. applegateiOphiactidaeOne of the few articulated ophiuroids from the Early Cretaceous[44]
OphidiasterO. spOphidiasteridaeOne of the few starfish from this formation which is not known from juvenile specimens[43]
PaleopentactaP. alencasteraeCucumariidaeLikely an infaunal suspsension feeder based on extant relatives[45]
ParapsolusP. tlayuensisPsolidaeLikely a suspension-feeder attached to hard substrates based on extant psolids[45]
PlutonasterP. spAstropectinidaeKnown from a juvenile specimen[43]
TamariaT. spOphidiasteridaeKnown from a juvenile[43]

Mollusks

Other invertebrates

Plants

References

  1. ^ Applegate, S. (1988). "A new genus and species of a holostean belonging to the family Ophiopsidae, Teoichthys kallistos, from the Cretaceous, near Tepexi de Rodriguez, Puebla". Revista Mexicana de Ciencias Geológicas. 7 (2): 200–205.
  2. ^ Pantoja-Alor, Jerjes (1992). "Geología y paleoambiente de la Cantera Tlayúa, Tepexi de Rodríguez Estado de Puebla". Rev. Instituto Geol. UNAM.. 9 (2): 156–169.
  3. ^ Applegate, Shelton P.; Espinosa-Arrubarrena, Luis; Alvarado-Ortega, Jesús; Benammi, Mouloud (2006), "Revision of Recent Investigations in the Tlayúa Quarry", Studies on Mexican Paleontology, Vega, Francisco J. (ed.), Dordrecht: Springer Netherlands, pp. 275–304, doi:10.1007/1-4020-3985-9_13. ISBN 978-1-4020-3985-0
  4. ^ Juárez-Arriaga, Edgar; Barragán, Ricardo; Núñez-Useche, Fernando; Moreno-Bedmar, Josep A. (2023-11-01). "Sedimentary environments in the prelude to the lagerstätten conditions of the Tlayúa Formation (Albian) in central Mexico: A microfacies approach". Journal of South American Earth Sciences. 131. Bibcode:2023JSAES.13104650J. doi:10.1016/j.jsames.2023.104650. ISSN 0895-9811
  5. ^ Chiocchini, Maurizio; Mancinelli, Anna; Romano, Antonio (1984). "Stratigraphic distribution of benthic Foraminifera in the Aptian, Albian and Cenomanian carbonate sequences of the Aurunci and Ausoni Mountains (Southern Lazio, Italy)". Benthos Research. 1: 167–181.
  6. ^ Juárez-Aguilar, E. Aldrin; Sánchez-Beristain, Francisco; Bernal, Juan Pablo; Morton-Bermea, Ofelia; García-Barrera, Pedro (2025-03-01). "Palaeoenvironmental inferences of a Mexican Konservat-Lagerstätte (Tlayúa Quarry; Lower Cretaceous) based on the geochemistry of rare earth elements". Cretaceous Research. 167. Bibcode:2025CrRes.16706045J. doi:10.1016/j.cretres.2024.106045. ISSN 0195-6671
  7. ^ Espinosa-Arrubarrena, Luis & Applegate, Shelton Pleasants (1996). "A paleoecological model of the vertebrate bearing beds in the Tlayúa Quarries, near Tepexi de Rodriguez, Puebla, Mexico". Mesozoic Fishes – Systematics and Paleoecology. Arriata, Gloria & Viohl, Günter (eds.). Proceedings of the International Meeting Eichstätt. pp. 539–550. ISBN 978-3-923871-90-2.
  8. ^ Alvarado-Ortega J, González-Rodríguez KA, Blaco-Piñón A, Espinosa-Arrubarrena L, Ovalles-Damián E (2006). "Mesozoic Osteichthyans of Mexico". Studies on Mexican Paleontology, Topics on Geobiology 24.. Vega, FJ; TG Nyborg; MC Perrilliat; M. Montellano-Ballesteros; SRS Cevallos-Ferriz. (eds.). Dordrecht, Netherlands: Springer Verlag. pp. 169–207. ISBN 1-4020-3985-9.
  9. ^ Vega FJ, Bruce NL, Serrano L, Bishop GA, Perrilliat MD (2005). "A review of the Lower Cretaceous (Tlayúa Formation: Albian) Crustacea from Tepexi de Rodríguez, Puebla, Central Mexico". Bulletin of the Mizunami Fossil Museum. ' (32): 25–30. Archived 2011-06-12 at the Wayback Machine.
  10. ^ Buitrón-Sánchez, Blanca Estela; Durán-Gónzalez, Alicia; Martín-Cao-Romero, Carolina; Solís-Marín, Francisco Alonso; Laguarda-Figueras, Alfredo. "Lower Cretaceous (Albian) Asteroidea (Echinodermata) from Tepexi de Rodriguez, Puebla, Mexico". Revista de Biología Tropical. 63 (2): 7–15.
  11. ^ Solé, Jesús; Pi-Puig, Teresa; Bermúdez-Chávez, Cynthia; Garduño-Martínez, Diana; Alvarado-Ortega, Jesús (2022-12-05). "Mineralogy, geochemistry, and K-Ar dating of feldspars and clays from an exceptional Cretaceous fossil locality (Tlayúa, Puebla, Mexico): Insights into the depositional and diagenetic ages and processes". Chemical Geology. 612. Bibcode:2022ChGeo.61221134S. doi:10.1016/j.chemgeo.2022.121134. ISSN 0009-2541
  12. ^ Barrett, Paul M.; Butler, Richard J.; Edwards, Nicholas P.; Milner, Andrew R. (2008). "Pterosaur distribution in time and space: an atlas". Zitteliana Reihe B. 28: 61–107.
  13. ^ Rivera-Sylva, Héctor E.; Carpenter, Kenneth; Frey, Eberhard (2014). Dinosaurs and other reptiles from the Mesozoic of Mexico. Life of the past. Bloomington: Indiana university press. ISBN 978-0-253-01183-1.
  14. ^ Reynoso, V.-H. (29 March 1998). "Huehuecuetzpalli mixtecus gen. et sp. nov: a basal squamate (Reptilia) from the Early Cretaceous of Tepexi de Rodríguez, Central México". Philosophical Transactions of the Royal Society B: Biological Sciences. 353 (1367): 477–500. doi:10.1098/rstb.1998.0224. JSTOR 56466. PMC 1692218
  15. ^ Reynoso, Victor-Hugo (1997-04-16). "A "beaded" sphenodontian (Diapsida: Lepidosauria) from the Early Cretaceous of central Mexico". Journal of Vertebrate Paleontology. 17 (1): 52–59. Bibcode:1997JVPal..17...52R. doi:10.1080/02724634.1997.10010953. ISSN 0272-4634
  16. ^ Reynoso, V. H. (2000). "An Unusual Aquatic Sphenodontian (Reptilia: Diapsida) from the Tlayua Formation (Albian), Central Mexico". Journal of Paleontology. 74 (1): 133–148. Bibcode:2000JPal...74..133R. doi:10.1017/S0022336000031310. JSTOR 1306891
  17. ^ Vega, Francisco J.; Nyborg, Torrey G.; María del Carmen Perrilliat; Marisol Montellano-Ballesteros; Sergio R.S Cevallos-Ferriz; Sara A. Quiroz-Barroso (2006). "Research on Fossil Amphibians and Reptiles". Studies on Mexican Paleontology. Vol. 24. Dordrecht, Netherlands: Springer. p. 214. ISBN 1-4020-3882-8. Retrieved 31 May 2011.
  18. ^ Rodríguez, Katia González & Fielitz, Christopher. "CRETACEOUS OSTEICHTHYAN FISH ASSEMBLAGES FROM MEXICO". Cretaceous Period: Biotic Diversity and Biogeography. New Mexico Museum of Natural History and Science Bulletin. 71
  19. ^ Alvarado-Ortega, Jesús & Brito, Paulo M. (2011). "A New Species of Araripichthys (teleostei, Elopocephala) from the Tlayúa Formation (cretaceous, Albian), Mexico". Journal of Vertebrate Paleontology. 31 (6): 1376–1381. Bibcode:2011JVPal..31.1376A. doi:10.1080/02724634.2011.602779. ISSN 0272-4634. JSTOR 41407692
  20. ^ Alvarado-Ortega, Jesús; Than-Marchese, Bruno Andrés; Melgarejo-Damián, María del Pilar (2020). "On the Albian occurrence of Armigatus (Teleostei, Clupeomorpha) in America, a new species from the Tlayúa Lagerstätte, Mexico". Palaeontologia Electronica. doi:10.26879/1107
  21. ^ Than-Marchese, Bruno Andrés & Alvarado-Ortega, Jesús (October 2022). "Armigatus felixi sp. nov. An Albian double armored herring (Clupeomorpha, Ellimmichthyiformes) from the Tlayúa lagerstätte, Mexico". Journal of South American Earth Sciences. 118. Bibcode:2022JSAES.11803905T. doi:10.1016/j.jsames.2022.103905
  22. ^ Brito, Paulo M. & Alvarado-Ortega, Jesus (4 September 2013). "Cipactlichthys scutatus, gen. nov., sp. nov. a New Halecomorph (Neopterygii, Holostei) from the Lower Cretaceous Tlayua Formation of Mexico". PLOS ONE. 8 (9). Bibcode:2013PLoSO...873551B. doi:10.1371/journal.pone.0073551. PMID 24023885
  23. ^ reassess
  24. ^ Alves, Yuri Modesto; Alvarado-Ortega, Jesús; Brito, Paulo M. (June 2020). "†Epaelops martinezi gen. and sp. nov. from the Albian limestone deposits of the Tlayúa quarry, Mexico – A new late Mesozoic record of Elopiformes of the western Tethys". Cretaceous Research. 110. Bibcode:2020CrRes.11004260A. doi:10.1016/j.cretres.2019.104260
  25. ^ González-Rodríguez, Katia; Applegate, Shelton P.; Espinosa-Arrubarrena, Luis (2004). "A New World Macrosemiid (Pisces: Neopterygii-Halecostomi) from the Albian of México". Journal of Vertebrate Paleontology. 24 (2): 281–289. Bibcode:2004JVPal..24..281G. doi:10.1671/1862. ISSN 0272-4634. JSTOR 4524716
  26. ^ Alvarado-Ortega, Jesús; de Mayrinck, Diogo; Brito, Paulo M. (July 2008). "A basal Pachyrhizodontid fish (Actinopterygii, Teleostei) from the Lower Cretaceous of the Tlayúa Quarry, Central Mexico". Comptes Rendus Palevol. 7 (5): 269–275. Bibcode:2008CRPal...7..269A. doi:10.1016/j.crpv.2008.03.006
  27. ^ Arratia Fuentes, Gloria & Tintori, Andrea (2004). Mesozoic fishes: proceedings of the International meeting, Serpiano, 2001. München: Verl. Dr. Friedrich Pfeil. ISBN 3-89937-053-8.
  28. ^ Alvarado-Ortega, Jesús & Alves, Yuri Modesto (July 2022). "Nusaviichthys nerivelai gen. et sp. nov., an Albian crossognathiform fish from the Tlayúa lagerstätte, Mexico". Cretaceous Research. 135. doi:10.1016/j.cretres.2022.105189
  29. ^ Grande, Lance & Bemis, William E. (10 April 1998). "A Comprehensive Phylogenetic Study of Amiid Fishes (Amiidae) Based on Comparative Skeletal Anatomy. an Empirical Search for Interconnected Patterns of Natural History". Journal of Vertebrate Paleontology. 18 (sup1): 1–696. Bibcode:1998JVPal..18S...1G. doi:10.1080/02724634.1998.10011114
  30. ^ Alvarado-Ortega, Jesús & Melgarejo-Damián, María Del Pilar (29 November 2017). "Paraclupea seilacheri sp. nov., un arenque de doble armadura (Clupeomorpha, Ellimmichthyiformes) de las calizas albianas de la cantera Tlayúa, Puebla, México". Revista Mexicana de Ciencias Geológicas. 34 (3): 234. Bibcode:2017RMxCG..34..234A. doi:10.22201/cgeo.20072902e.2017.3.528
  31. ^ Alvarado-Ortega, Jesús (July 2014). "Ancient herring from the Tlayúa Quarry (Cretaceous, Albian) near Tepexi de Rodíguez, Puebla State, central Mexico, closing the gap in the early diversification of Clupeomorpha". Cretaceous Research. 50: 171–186. Bibcode:2014CrRes..50..171A. doi:10.1016/j.cretres.2014.02.022
  32. ^ Alvarado-Ortega, Jesús & Espinosa-Arrubarrena, Luis (January 2008). "A new genus of ionoscopiform fish (Halecomorphi) from the Lower Cretaceous (Albian) lithographic limestones of the Tlayúa Quarry, Puebla, Mexico". Journal of Paleontology. 82 (1): 163–175. Bibcode:2008JPal...82..163A. doi:10.1666/04-152.1
  33. ^ Pacheco-Ordaz, Stephanie; Mejía, Omar; Alvarado-Ortega, Jesús (February 2025). "Tahnaichthys magnuserrata gen. and sp. nov., a double-hump pycnodontid (Actinopterygii, Pycnodontiformes) fish from the Albian limestones of the Tlayúa Quarry, Puebla, Mexico". Journal of South American Earth Sciences. 152. Bibcode:2025JSAES.15205277P. doi:10.1016/j.jsames.2024.105277
  34. ^ Machado, Giselle P.; Alvarado-Ortega, Jesús; Machado, Lúcio Paulo; Brito, Paulo M. (March 2013). "Teoichthys brevipina , sp. nov., a new ophiopsid fish (Halecomorphi, Ionoscopiformes) from the Lower Cretaceous Tlayúa Formation, Central Mexico". Journal of Vertebrate Paleontology. 33 (2): 482–487. Bibcode:2013JVPal..33..482M. doi:10.1080/02724634.2013.729962
  35. ^ Applegate, Shelton P. (1988). "A new genus and species of a holostean belonging to the family Ophiopsidae, Teoichthys kallistos, from the Cretaceous, near Tepexi de Rodríguez, Puebla". Revista Mexicana de Ciencias Geológicas. 7 (2): 200–205.
  36. ^ Applegate, Shelton P. (1992). "A new genus and species of pycnodont from the Cretaceous (Albian) of Central Mexico, Tepexi de Rodriguéz, Puebla". Revista Mexicana de Ciencias Geológicas. 10 (2): 164–178.
  37. ^ Brito, Paulo M.; Villalobos-Segura, Eduardo; Alvarado-Ortega, Jésus (March 2019). "A new early cretaceous guitarfish (chondrichthyes, batoidea) from the Tlayúa Formation, Puebla, Mexico". Journal of South American Earth Sciences. 90: 155–161. Bibcode:2019JSAES..90..155B. doi:10.1016/j.jsames.2018.12.005
  38. ^ López-Arbarello, Adriana & Alvarado-Ortega, Jesús (26 January 2011). "New semionotiform (Neopterygii) from the Tlayúa Quarry (Early Cretaceous, Albian), Mexico". Zootaxa. 2749 (1): 1–24. doi:10.11646/zootaxa.2749.1.1
  39. ^ Alvarado-Ortega, Jesús (2004). "Description and Relationships of a New Ichthyodectiform Fish from the Tlayúa Formation (Early Cretaceous: Albian), Puebla, Mexico". Journal of Vertebrate Paleontology. 24 (4): 802–813. doi:10.1671/0272-4634(2004)024[0802:DAROAN2.0.CO;2]. ISSN 0272-4634. JSTOR 4524776
  40. ^ Feldmann, Rodney M.; Vega, Francisco J.; Applegate, Shelton P.; Bishop, Gale A. (1998). "Early Cretaceous Arthropods from the Tlayúa Formation at Tepexi de Rodríguez, Puebla, México". Journal of Paleontology. 72 (1): 79–90.
  41. ^ Dunlop, J. A., Penney, D. & Jekel, D. 2023. A summary list of fossil spiders and their relatives. In World Spider Catalog. Natural History Museum Bern, online at http://wsc.nmbe.ch , version 23.5
  42. ^ Luque, Javier (March 2015). "The oldest higher true crabs (Crustacea: Decapoda: Brachyura): insights from the Early Cretaceous of the Americas". Palaeontology. 58 (2): 251–263. doi:10.1111/pala.12135
  43. ^ Buitrón-Sánchez, Blanca Estela; Durán-Gónzalez, Alicia; Martín-Cao-Romero, Carolina; Solís-Marín, Francisco Alonzo; Laguarda-Figueras, Alfredo (June 2015). "Lower Cretaceous (Albian) Asteroidea (Echinodermata) from Tepexi de Rodriguez, Puebla, Mexico". Revista de Biología Tropical. 63 (2). doi:10.15517/rbt.v63i2.23124
  44. ^ Martin‐Medrano, Leonora; Thuy, Ben; García-Barrera, Pedro (January 2009). "NEW ALBIAN (EARLY CRETACEOUS) OPHIUROIDS FROM THE TLAYÚA QUARRY, PUEBLA, MEXICO". Palaeontology. 52 (1): 83–94. doi:10.1111/j.1475-4983.2008.00836.x
  45. ^ Applegate, Shelton P.; Buitrón-Sánchez, Blanca E.; Solís-Marín, Francisco A.; Laguarda-Figueras, Alfredo; Ibanez, Jason C. (January 2009). "Two Lower Cretaceous (Albian) Fossil Holothurians (Echinodermata) from Tepexi De Rodriguez, Puebla, Mexico". Proceedings of the Biological Society of Washington. 122 (1): 91–102. doi:10.2988/07-41.1

Further reading

  • L. Martin-Medrano and P. Garcia-Barrera. 2006. Fossil Ophiuroids of Mexico. In F. J. Vega, T. G. Nybor, M. D. C. Perrillat, M. Montellano-Ballesteros, S. R. S. Cevallos-Ferriz, S. A. Quiroz-Barroso (eds.), Topics in Geobiology 24:115-131