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Last edited December 21, 2025


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The Cambay Shale Formation is an Early Eocene-aged geologic formation in the Cambay Basin, India. It varies in thickness from a few meters on the margins of the basin to more than 2,500m in the depressions. It directly overlies the Olpad Formation and is, in turn, overlain by the Anklesvar Formation in the southern part of the basin and by Kalol Formation in the northern part of the basin. Further north, the Cambay Shale, in its lower part, is gradually replaced by tongues of paralic-deltaic Kadi Formation and finally by Tharad Formation.[1]

Paleobiota[edit | edit source]

Vertebrates[edit | edit source]

Mammals[edit | edit source]

Source:[2]

Anthracobunians from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Cambaytherium C. thewissi A cambaytheriid.
C. gracilis
C. bidens
Kalitherium K. marinus Another cambaytheriid.
Indobune I. vastanensis A possible anthracobunid.
Primates[3] from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Asiadapis A. cambayensis An Asiadapine.
A. tapiensis
Marcgodinotius M. indicus Another Asiadapine.
Vastanomys V. gracilis A omomyid.
V. major
Hyaenodonts from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Indohyaenodon I.raoi A hyaenodont.
Dichobunids from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Gujaratia G. indica A Diacodexeid.
G. pakistanensis
Tapiromorphs from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Cambaylophus[4] C. vastanensis A tapiromorph.
Vastanolophus[5] V. holbrooki A tapiromorph.
Cimolestans from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Suratilestes S. gingerichi A cimolestan.
Anthraconyx A. hypsomylus A esthonychid
Bats from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Archaeonycteris A. storchi A bat.
Protonycteris P. gunnelli A bat.
Cambaya C. complexus A bat.
Hassianycteris H. kumari A bat.
Icaronycteris I. sigei A bat.
Jaegeria J. cambayensis A bat.
Microchiropteryx M. folieae A bat.
Other mammals from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Frugivastodon F. cristatus A apatemyid.
Indolestes I. kalamensis An adapisoriculid.
Indodelphis I. luoi A opossum.
Meldimys[6] M. musak A rodent.
Indonyctia[7] I. cambayensis A nyctitheriid.
Pahelia P. mysteriosa A herbivorous mammal of uncertain affinities.[8]

Birds[edit | edit source]

Birds from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Vastanavis V.eocaena A parrot.

Reptiles[edit | edit source]

Lizards from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Indiagama I. gujarata A iguanian.
Suratagama S. neeraae A iguanian.
Tinosaurus T. indicus A iguanian.
Vastanagama V. susanae A iguanian.
Heterodontagama H. borsukae A priscagamid iguanian.
Snakes from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Platyspondylophis P. tadkeshwarensis A Madtsoiid.
Russellophis R. crassus A russelophiid.
Procerophis P. sahnii A snake.
Thaumastophis T. missiaeni A snake.
Palaeophis P. vastaniensis A paleophiid snake.

Amphibians[edit | edit source]

Amphibians from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Eobarbourula E. delfinoi A toad.
Indorana I.prasadi A frog.

Ray-finned fish[edit | edit source]

Ray-finned fish from the Cambay Shale formation
Genus Species Stratigraphic position Material Notes Images
Avitoplectus A. molaris Lower jaw A bizarre tetraodontiform.[9]
Diodon D. sp. Teeth A porcupinefish.[10]
Egertonia E. sp. Teeth A phyllodontid elopomorph.[10]
?Enchodus ?E. sp. Teeth An enchodontid aulopiform. Among the latest known remains of this otherwise Cretaceous genus,[10] but may potentially represent reworked material from lower layers.[11]
Eotrigonodon E. indicus Teeth A pycnodont.[10]
Eutrichiurides E. sp. Teeth A cutlassfish.[10]
Osteoglossidae indet. Teeth, scales A bonytongue.[10]
Sphyraena S. sp. Teeth A barracuda.[10]
Stephanodus S. lybicus Teeth A pycnodont.[10]

Arthropods[edit | edit source]

Crabs[edit | edit source]

Crabs from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Philyra P. karkata A leucosiid crab.

Insects[edit | edit source]

Beetles from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Protoclaviger[12] P.trichodens A Beetle.
Cambaltica C. paleoindica A Flea Beetle.
Paleosorius P. cambayensis A Rove Beetle.
Hemipterans[13] from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Heteromargarodes H. hukamsinghi A sand pearl.
H. americanus
Normarkicoccus N. cambayae A diaspidid scale insect.
Neuropterans from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Spiloconis[14] S. sexguttata A Aleuropterygine Coniopterygid.
S. glaesaria
S. oediloma
S. eominuta
Bees from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Melikertes[15] M. (Paramelikertes) gujaratensis A bee.
M. (Melikertes) kamboja
Wasps from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Trichelyon T. tadkeshwarense A braconid wasp.
Termites[16] from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Nanotermes N. isaacae A termitid termite.
Prostylotermes P. kamboja A Stylotermitid termite.
Parastylotermes P. krishnai A Stylotermitid termite.
Zophotermes Z. ashoki A Rhinotermitid termite
Flies from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Sycorax[17] S. longistyla A moth fly.
Phlebotoiella P. eoindianensis A moth fly of unceritian affinities.
Dicranomyia D. (Dicranomyia) indica The oldest known Crane fly and tipulomorph.[18]
Stempellina S.pollex A non-biting midge
S.stebneri
Tanytarsus T.forfex A non-biting midge
T.ramus
Camptopterohelea C. odora A non-biting midge
Eohelea E. indica A biting midge
Gedanohelea G. gerdesorum A biting midge
Indorrhina I.sahnii A biting midge
Lygistorrhina L. indica A biting midge
Meunierohelea M. borkenti A biting midge
Meunierohelea M. cambayana A biting midge
Meunierohelea M. orientalis A biting midge
Palaeognoriste P. orientale A biting midge
Webspinners from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Kumarembia K. hurleyi One of the few fossil Webspinners known.

Arachnids[edit | edit source]

Whip spiders from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Paracharonopsis[19] P. cambayensis A Paracharontid.

Plants[edit | edit source]

Plants from the Cambay Shale formation
Genus Species Stratigraphic position Notes Images
Pterospermoxylon P. suratensis An angiosperm.
Anthocephalophyllum A. vastanicum An angiosperm.
Ebenoxylon E. cambayense An angiosperm.
Gardeniophyllum G. cambayum An angiosperm.
Calophyllaceophyllum C. eocenicum An angiosperm.
Carallioipollenites C. integerrimoides An angiosperm known from fossilised pollen.
Gynocardia G. eocenica An angiosperm.


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References[edit | edit source]

  1. Chowdhary, L. R. (2004). Petroleum geology of the Cambay Basin, Gujarat., India. Dehradun: Indian Petroleum Publishers. ISBN 978-81-900361-3-9.
  2. "Vastan Lignite Mine, Gujarat, India".
  3. Rose, Kenneth D.; Rana, Rajendra S.; Sahni, Ashok; Kumar, Kishor; Missiaen, Pieter; Singh, Lachham; Smith, Thierry (April 2009). "Early Eocene Primates from Gujarat, India". Journal of Human Evolution. 56 (4): 366–404. doi:10.1016/j.jhevol.2009.01.008. PMID 19303624.
  4. Kapur, Vivesh V.; Bajpai, Sunil (31 December 2015). "Oldest South Asian tapiromorph (Perissodactyla, Mammalia) from the Cambay Shale Formation, western India, with comments on its phylogenetic position and biogeographic implications". Journal of Palaeosciences. 64 (1–2): 95–103. doi:10.54991/jop.2015.104.
  5. Smith, Thierry; Solé, Floréal; Missiaen, Pieter; Rana, Rajendra; Kumar, Kishor; Sahni, Ashok; Rose, Kenneth (8 September 2015). "First early Eocene tapiroid from India and its implication for the paleobiogeographic origin of perissodactyls". Palaeovertebrata. doi:10.18563/pv.39.2.e5.
  6. Rana, Rajendra S.; Kumar, Kishor; Escarguel, Gilles; Sahni, Ashok; Rose, Kenneth D.; Smith, Thierry; Singh, Hukam; Singh, Lachham (January 2008). "An Ailuravine Rodent from the Lower Eocene Cambay Formation at Vastan, Western India, and Its Palaeobiogeographic Implications". Acta Palaeontologica Polonica. 53 (1): 1–14. doi:10.4202/app.2008.0101.
  7. Das, Debasis P.; Carolin, Nora; Bajpai, Sunil (3 July 2022). "A nyctitheriid insectivore (Eulipotyphla, Mammalia) of Asian affinity from the early Eocene of India". Historical Biology. 34 (7): 1157–1165. doi:10.1080/08912963.2021.1966002.
  8. Zack, Shawn P.; Rose, Kenneth D.; Holbrook, Luke T.; Kumar, Kishor; Rana, Rajendra S.; Smith, Thierry (2021). "An enigmatic new ungulate-like mammal from the early Eocene of India". Papers in Palaeontology. 7 (1): 497–520. doi:10.1002/spp2.1288. ISSN 2056-2802.
  9. Bemis, Katherine E.; Tyler, James C.; Bemis, William E.; Kumar, Kishor; Rana, Rajendra Singh; Smith, Thierry (2017-11-02). "A gymnodont fish jaw with remarkable molariform teeth from the early Eocene of Gujarat, India (Teleostei, Tetraodontiformes)". Journal of Vertebrate Paleontology. 37 (6) e1369422. doi:10.1080/02724634.2017.1369422. ISSN 0272-4634.
  10. 10.0 10.1 10.2 10.3 10.4 10.5 10.6 10.7 Rana, R. S.; Kumar, K.; Singh, H. (2004). "Vertebrate fauna from the subsurface Cambay Shale (Lower Eocene), Vastan Lignite Mine, Gujarat, India". Current Science. 87 (12): 1726–1733. ISSN 0011-3891.
  11. Davis, Matthew P.; Fielitz, Christopher (December 2010). "Estimating divergence times of lizardfishes and their allies (Euteleostei: Aulopiformes) and the timing of deep-sea adaptations". Molecular Phylogenetics and Evolution. 57 (3): 1194–1208. Bibcode:2010MolPE..57.1194D. doi:10.1016/j.ympev.2010.09.003. PMID 20854916.
  12. "Specialized Myrmecophily at the Ecological Dawn of Modern Ants: Current Biology".
  13. Vea, Isabelle M.; Grimaldi, David A. (2015). "Diverse New Scale Insects (Hemiptera: Coccoidea) in Amber from the Cretaceous and Eocene with a Phylogenetic Framework for Fossil Coccoidea". American Museum Novitates (3823): 1–15. doi:10.1206/3823.1. S2CID 73702369.
  14. Grimaldi, David; Engel, Michael S.; Nascimbene, Paul c.; Singh, Hukam (2013). "Coniopterygidae (Neuroptera: Aleuropteryginae) in Amber from the Eocene of India and the Miocene of Hispaniola". American Museum Novitates (3770): 20–39. doi:10.1206/3770.2. S2CID 56567508.
  15. Engel, Michael S.; Ortega-Blanco, Jaime; Nascimbene, Paul C.; Singh, Hukam (17 December 2013). "The bees of Early Eocene Cambay amber (Hymenoptera: Apidae)". Journal of Melittology (25): 1–12. doi:10.17161/jom.v0i25.4659. hdl:1808/14444.
  16. Engel, Michael; Grimaldi, David; Nascimbene, Paul; Singh, Hukam (21 November 2011). "The termites of Early Eocene Cambay amber, with the earliest record of the Termitidae (Isoptera)". ZooKeys (148): 105–123. doi:10.3897/zookeys.148.1797. PMC 3264413. PMID 22287892.
  17. Wagner, RüDiger; Agnihotri, Priya; Singh, Hukam (25 August 2022). "A new species of Sycorax (Sycoracinae: Psychodidae) from the Lower Eocene amber of Tadkeshwar, Gujarat, India". Palaeoentomology. 5 (4). doi:10.11646/palaeoentomology.5.4.4.
  18. Kania, Iwona; Krzemiński, Wiesław; Stebner, Frauke; Singh, Hukam (June 2016). "The first representative of Tipulomorpha (Diptera) from Early Eocene Cambay amber (India)". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 107 (2–3): 263–269. doi:10.1017/S1755691017000433.
  19. Engel, Michael S.; Grimaldi, David A (6 August 2014). "Whipspiders (Arachnida: Amblypygi) in amber from the Early Eocene and mid-Cretaceous, including maternal care". Novitates Paleoentomologicae (9): 1. doi:10.17161/np.v0i9.4765. hdl:1808/15287.