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Northumberland Formation
Stratigraphic range: Late Cretaceous, late Campanian
TypeGeological formation
Unit ofNanaimo Group
UnderliesGeoffery Formation
OverliesDe Courcy Formation
Location
Country Canada

The Northumberland Formation (previously also known as the Lambert Formation) is a Late Cretaceous geological formation exposed in the Hornby, Denman, Mayne, and Gabriola Islands, all located near Vancouver Island in British Columbia, Canada. Belonging to the larger Nanaimo Group, the Northumberland Formation has been dated to the upper Campanian, and numerous vertebrate, invertebrate, and plant fossils have been recovered from the formation, including birds, pterosaurs, sharks, coleoids, ammonites, angiosperms, and gymnosperms.[1][2][3][4][5]

Geology

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The Northumberland Formation is exposed in small strips of the Gabriola and Mayne Islands,[6] east half of Denman Island,[7] and along the western to southeastern shores of Hornby Island.[5][7] All of these islands are situated east of Vancouver Island in the Strait of Georgia. Certain areas of Hornby Island that expose the Northumberland Formation were formerly mapped as the defunct Lambert Formation and the Spray Formation, being reassigned with evolving interpretations and nomenclature.[5] Part of the Nanaimo Group, the Northumberland Formation overlies the De Courcy Formation and underlies the nonfossiliferous Geoffrey Formation, both dominated by thick-bedded sandstone and conglomerate.[7]


Paleoenvironment

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Various interpretations of the Northumberland Formation paleoenvironment have been proposed. A deep-water submarine fan complex has been put forth by several authors based on lithology.[7] Contrasting interpretations argue that the Northumberland Formation was deposited in a shallower environment. Jenkins et al. (2017) suggest an middle to outer shelf 100-300 m in depth based on macrofossil and lithological evidence, supported by foraminiferal data.

Paleobiota

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Color key
Taxon Reclassified taxon Taxon falsely reported as present Dubious taxon or junior synonym Ichnotaxon Ootaxon Morphotaxon
Notes
Uncertain or tentative taxa are in small text; crossed out taxa are discredited.

Bivalves

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Bivalves
Genus Species Location Stratigraphic position Material Notes Images
Acila A. grahami Collishaw Point Numerous shells
Acila castrensis, an extant relative
Amphidonte A. parasitica[8] Collishaw Point
Anomia A. vancouverensis[1]
Anomia ephippium, an extant relative
Caryocorbula C. traskii[9][10] Northwest side of Hornby Island Three partial right valves
Caryocorbula contracta, an extant relative
Glycymerita G. veatchii[1] Hornby Island
Inoceramus I. ex. gr. balticus Collishaw Point
I. (Platyceramus) cf. cycloides
I. ex gr. regularis Collishaw Point
I. vancouverensis[1] Collishaw Point
Nemodon N. vancouverensis[1] Collishaw Point
Paraesa? P.? lens[11] Gabriola Island
Solemya S. sp.[1] Collishaw Point
Atlantic awning clam (Solemya velum), an extant relative
Teredina T. suciensis[1] Collishaw Point

Scaphopods

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Scaphopods
Genus Species Location Stratigraphic position Material Notes Images
Antalis A. cooperi[1] Collishaw Point
Common tusk shell (Antalis vulgaris), an extant relative

Gastropods

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Gastropods
Genus Species Location Stratigraphic position Material Notes Images
Nonacteonina N. sp.[1] Collishaw Point
Tessarolax T. loullae[12] Collishaw Point, Phipps Point

Ammonites

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Ammonites
Genus Species Location Stratigraphic position Material Notes Images
Anagaudryceras A. politissimum[1] Collishaw Point
Baculites B. occidentalis[5] Southeastern shore of Hornby Island to Collishaw Point Spans nearly the entirety of the Northumberland Formation Includes material previously referred to B. chicoensis
Life reconstruction of Baculites vertebralis
Diplomoceras D. (Diplomoceras) cylindraceum[1][5] Western coast of Hornby Island, from Shingle Spit to Collishaw Point; one isolated occurrence from southeastern shore[5] Includes material previously referred to Hamites obstrictus
Life reconstruction of Diplomoceras
D. (Diplomoceras) cf. cylindraceum[5] Mid-western coast of Hornby Island Cannot be confidently assigned to D. (D.) cylindraceum, although fits within the parameters of the species
Exiteloceras E. (Exiteloceras) denicostatum[5] Collishaw Point Upper part of formation
Exiteloceras jenneyi
E. (Neancyloceras) aff. bipunctatum[5] Southeastern to mid-western shore of Hornby Island Lower part of formation
Fresvillia F. constricta[5] Western shore of Hornby Island, from Shingle Spit to Collishaw Point
Gaudryceras G. denmanense'[1] Denman and Hornby Islands
Gaudryceras denseplicatum
G. aff. venustum[13] Between Shingle Spit and Phipps Point G. venustum now a junior synonym of G. izumiense
G. sp.[14] Between Shingle Spit and Savoie's wharf Body chamber fragment of large individual Ornamentation similar to G. mitis
Hypophylloceras H. (Neophylloceras) ramosum[1] Collishaw Point
H. (Neophylloceras) hetonaiense[14] North side of Hornby Island Senior synonym of H. (N.) lambertense[15]
H. (Neophylloceras) surya[1] Collishaw Point
Nostoceras N. (Didymoceras?) adrotans[5] Southeastern coast of Hornby Island Lower part of formation Includes nomen dubium Anisoceras cooperi
Nostoceras fossil from the Late Cretaceous of Hornby Island
N. (Nostoceras) hornbyense[1][5] Western shores of Hornby Island, from Shingle Spit to Collishaw Point Upper part of formation
N. (Nostoceras) aff. pauper[5] Western shores of Hornby Island, from Shingle Spit to Collishaw Point Upper part of formation
Pachydiscus P. hornbyense[1] Collishaw Point
Pachydiscus japonicus
P. cf. ootacodensis
P. suciaensis[1] Collishaw Point
Phyllopachyceras P. forbesianum
Phylloptychoceras P. horitai[5] Southeastern shore of Hornby Island to Collishaw Point Spans nearly the entirety of the Northumberland Formation
Pseudophyllites P. indra[1] Collishaw Point
Solenoceras S. exornatus[5] Southeastern shore of Hornby Island Lower part of formation
S. cf. reesidei[5] Collishaw Point Upper part of formation

Coleoids

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Coleoids
Genus Species Location Stratigraphic position Material Notes Images
Actinosepia A. canadensis[16][17] Collishaw Point[1]
Cyrtobelus C. hornbyense Collishaw Point[1]
Enchoteuthis E. sp.[18]
Reconstruction of Enchoteuthis melanae

Crustaceans

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Crustaceans
Genus Species Location Stratigraphic position Material Notes Images
Archaeopus A. hoploserratus[19] "southeastern coast of Hornby Island"[19]
A. vancouverensis[1][19][20] Collishaw Point; "western coast of Hornby Island"[19]
Bournelyreidus B. grahamae[21] near Collishaw Point Carapace
Branchiocarcinus B. sp.[22] Collishaw Point Carapace and incomplete legs
Homolopsis H. sp.[1] Collishaw Point Carapace
Linuparus L. vancouverensis[1] Hornby Island
Japanese spear lobster (Linuparus tirgonus), an extant relative
Linuparus fossil from the Late Cretaceous of Hornby Island
Sabellidromites S. laneae[23] "near Manning Point"[23] Carapace and partial chelipeds Initially incorrectly spelled as S. lanae
Ostracoda indet.[24]

Insects

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Insects
Genus Species Location Stratigraphic position Material Notes Images
Curculionidae indet.[1] Collishaw Point Wing covers
Isoptera indet.[1] Collishaw Point Galleries within fossil driftwood, containing fecal pellets

Echinoderms

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Echinoderms
Genus Species Location Stratigraphic position Material Notes Images
Echinoidea indet.[24] Spines and plates
Goniasteridae indet.[1] Single incomplete arm
Holothuria indet.[24]

Sharks

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All following information follows Cappetta, Morrison, & Adnet (2021),[25] unless stated otherwise.

Sharks
Genus Species Location Stratigraphic position Material Notes Images
Carcharias C. dominguei Collishaw Point, Manning Point Over 1000 teeth from Collishaw Point, 130 teeth from Manning Point
Sand tiger shark (Carcharias taurus), an extant relative
Centrosqualus C. mustardi Collishaw Point, Manning Point 57 teeth from Collishaw Point, 34 teeth from Manning Point
Centroscymnus C. sp. Collishaw Point, Manning Point 4 teeth from Collishaw Point, 2 teeth from Manning Point
Portuguese dogfish (Centroscymnus coelolepis), an extant relative
Chlamydoselachus C. balli Collishaw Point, Manning Point 1 tooth from Collishaw Point, 2 teeth from Manning Point
Frilled shark (Chlamydoselachus anguineus), an extant relative
Dykeius D. garethi Collishaw Point, Manning Point 7 teeth from Collishaw Point, 2 teeth from Manning Point Estimated to be 7 m in length by Cappetta et al.
Echinorhinus E. sp.[26] Collishaw Point, Manning Point 220 teeth from Collishaw Point, 43 teeth from Manning Point Previously referred to E. lapaoi[25]
Prickly shark (Echinorhinus cookei), an extant relative
cf. Eoetmopterus cf. E. supracretaceus Collishaw Point, Manning Point 220 teeth from Collishaw Point, 43 teeth from Manning Point Estimated to be around 50 cm in length by Cappetta et al.
Velvet belly lanternshark (Etmopterus spinax), an extant relative
Florenceodon F. johnyi Collishaw Point, Manning Point 136 teeth from Collishaw Point, 29 teeth from Manning Point
Hemiscyllium H. hermani Collishaw Point 1 tooth
Epaulette shark (Hemiscyllium ocellatus), an extant relative
Hessinodon H. wardi Collishaw Point, Manning Point 2 teeth from Collishaw Point, 2 teeth from Manning Point
Cookiecutter shark (Isistius brasiliensis), a possible extant relative with similar lower teeth
Hexanchus H. microdon Collishaw Point, Manning Point 395 lower teeth from Collishaw Point, 151 lower teeth from Manning Point; numerous upper teeth from both sites
Bigeyed sixgill shark (Hexanchus nakamurai), an extant relative
Komoksodon K. kwutchakuth Collishaw Point, Manning Point 60 teeth from Collishaw Point, 26 teeth from Manning Point
Meristodonoides M. sp. Collishaw Point 1 tooth
Meristodonoides teeth from Santonian-early? Campanian Russia
Notidanodon N. pectinatus Collishaw Point, Manning Point 33 teeth from Collishaw Point, 14 teeth from Manning Point
Paraorthacodus P. rossi Collishaw Point, Manning Point 28 teeth from Collishaw Point, 7 teeth from Manning Point
Articulated keleton of Paraorthacodus jurensis
Plicatoscyllium P. cf. globidens Collishaw Point 1 tooth
Nurse shark (Ginglymostoma cirratum), an extant relative
Pristiophorus P. pricei Collishaw Point 30 teeth
Japanese sawshark (Pristiophorus japonicus), an extant relative
Skeleton of Pristophorus/Propristiophorus tumidens, a Late Cretaceous sawshark from Lebanon
P. smithi Collishaw Point, Manning Point 63 teeth from Collishaw Point, 1 tooth from Manning Point
Protocentrophorus P. steviae Collishaw Point, Manning Point 82 teeth from Collishaw Point, 39 teeth from Manning Point
Gulper shark (Centrophorus granulosus), an extant relative
Protoheptranchias P. lowei Collishaw Point, Manning Point 197 teeth from Collishaw Point, 58 teeth from Manning Point
Sharpnose sevengill shark (Heptranchias perlo), an extant relative
Rhinoscymnus R. clarki Manning Point 1 tooth
Little sleeper shark (Rhinoscymnus rostratus), an extant relative previously classified as Somniosus
Rolfodon R. ludvigseni Collishaw Point, Manning Point 213 teeth from Collishaw Point, 26 teeth from Manning Point
R. cf. thomsoni Manning Point 1 tooth
Squaliodalatias S. savoiei Collishaw Point, Manning Point 334 teeth from Collishaw Point, 74 teeth from Manning Point
Smalleye pygmy shark (Squaliolus aliae), an extant relative
Squalus S. nicholsae Collishaw Point, Manning Point 107 teeth from Collishaw Point, 27 teeth from Manning Point Notably large size compared to other Cretaceous Squalus species
Spiny dogfish (Squalus acanthias), an extant relative
S. vondermarcki Collishaw Point, Manning Point 258 teeth from Collishaw Point, 85 teeth from Manning Point Differs slightly from holotypic teeth from Campanian Germany, warrants taxonomic reanalysis[27]
S. sp. Collishaw Point 1 tooth
Synechodus S. dereki Collishaw Point, Manning Point 48 teeth from Collishaw Point, 4 teeth from Manning Point
Articulated skeleton of Synechodus ungeri
Xampylodon X. dentatus Collishaw Point, Manning Point 22 teeth from Collishaw Point, 7 teeth from Manning Point Estimated to be 3-4 m in length by Cappetta et al.
Scyliorhinidae indet. Collishaw Point 1 tooth
Chain catshark (Scyliorhinus retifer), an extant relative
Scyliorhinus elongatus, a Late Cretaceous catshark from Lebanon

Osteichthyans

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Osteichthyans
Genus Species Location Stratigraphic position Material Notes Images
Enchodontidae indet.[1] Collishaw Point Lower jaw
Life reconstruction of Enchodus petrosus
Saurodontidae indet.[28] Collishaw Point Left? maxilla Holotype of Gwawinapterus beardi, previously thought to be an istiodactylid pterosaur[29]
Life reconstruction of Saurodon leanus

Archosaurs

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Archosaurs
Genus Species Location Stratigraphic position Material Notes Images
Maaqwi M. cascadensis Near Phipps Point Associated right coracoid, humerus, ulna, and radius Coracoid previously described as Ornithurae indet.[3]
Holotype of Maaqwi cascadensis
Azhdarchidae? indet.[4] Collishaw Point Associated humerus, dorsal vertebrae, and indeterminate bone fragments; likely one individual
Crocodylia indet.[3] "a series of sites at Collishaw and Manning points"[3]
Euenantiornithes indet.[2][3] Collishaw Point Right radius, left coracoid,[2] and left femur[3] from different individuals Marked size difference between radius and coracoid
Ornithurae indet.[2][3] Collishaw Point Two left tarsometatarsi from different taxa; two distal ulnae of distinct sizes; left tibiotarsus;[2] left carpometacarpus[3]

Non-archosaur reptiles

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Non-archosaur reptiles
Genus Species Location Stratigraphic position Material Notes Images
Tylosaurus T. sp.[1] Collishaw Point Partial skull
Life reconstruction of Tylosaurus pembinensis
Mosasauridae indet.[1] Collishaw Point Isolated teeth
Testudines indet.[3] "a series of sites at Collishaw and Manning points"[3]

Mammals

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Mammals
Genus Species Location Stratigraphic position Material Notes Images
Multituberculata indet.[25] Undescribed tooth

Plants

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Ferns
Genus Species Location Stratigraphic position Material Notes Images
Appendicisporites A. sp.
Aquilapollenites A. quadrilobus
Atli A. morinii Collishaw Point Stem segment
Baculatisporites B. sp. Reported as "Baculitesporites" (lapsus calami)
Cibotiumspora C. sp.
Cicatricososporites C. norrisii
Cingutriletes C. sp.
Clavifera C. sp.
Cunninghamia C. hornbyensis Collishaw Point Twigs and leaves
Cunninghamia lanceolata
Cyathidites C. minor
C. sp.
Cycadeoidea C. maccafferyi Collishaw Point Ovulate cones
Dictyophyllidites D. sp. Reported as "Dictyophyllites" (lapsus calami)
Foraminisporis F. asymmetricus
Gleicheniidites G. spp.
Hamulatisporis H. sp.
Mancicorpus M. tripodiformis
Marcellopites M. tolmanensis
Microcachryidites M. sp.
Murospora? M.? sp.
Parviprojectus P. trialatus
Pilosisporites P. sp.
Picea P. spp.
Reticorpus R. delicatus
R. senonicus
Retitriletes R. sp.
Sabalites S. sp.[1] Collishaw Point Leaf-stalks
Sabalites from the Eocene Green River Formation in Wyoming
Siberiapollis S. spp.
Tricolpites T. sp.
Trilobosporites T. sp.
Triprojectus T. attenuatus
T. minshuensis
Trudopollis T. meekeri
T. sp.
Tschudypollis T. retusus
T. thalmannii
Varirugosisporites T. meekeri

See also

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Footnotes

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References

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  1. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad Ludvigsen, Rolf; Beard, Graham (1997). West Coast fossils: a guide to the ancient life of Vancouver Island (2nd ed.). Harbour Publishing. p. 216. ISBN 1-55017-179-8.
  2. ^ a b c d e Morrison, Kurt; Dyke, Gareth J.; Chiappe, Luis M. (2005). "Cretaceous fossil birds from Hornby Island (British Columbia)". Canadian Journal of Earth Sciences. 42 (12): 2097–2101. doi:10.1139/e05-081.
  3. ^ a b c d e f g h i j Dyke, Gareth; Wang, Xia; Kaiser, Gary (2011). "Large fossil birds from a Late Cretaceous marine turbidite sequence on Hornby Island (British Columbia)". Canadian Journal of Earth Sciences. 48 (11): 1489–1496. doi:10.1139/e11-050.
  4. ^ a b Martin-Silverstone, Elizabeth; Witton, Mark P.; Arbour, Victoria M.; Currie, Phillip J. (2016). "A small azhdarchoid pterosaur from the latest Cretaceous, the age of flying giants". Royal Society Open Science. 3 (8). doi:10.1098/rsos.160333.
  5. ^ a b c d e f g h i j k l m n o p McLachlan, Sandy M. S.; Haggart, James W. (2018). "Reassessment of the late Campanian (Late Cretaceous) heteromorph ammonite fauna from Hornby Island, British Columbia, with implications for the taxonomy of the Diplomoceratidae and Nostoceratidae". Journal of Systematic Paleontology. 16 (15): 1247–1299. doi:10.1080/14772019.2017.1381651.
  6. ^ Englert, Rebecca G.; Hubbard, Stephen M.; Coutts, Daniel S.; Matthews, William A. (2018). "Tectonically controlled initiation of contemporaneous deep-water channel systems along a Late Cretaceous continental margin, western British Columbia, Canada". Sedimentology. 65 (7): 2404–2438. doi:10.1111/sed.12472.
  7. ^ a b c d Katnick, Deanne C.; Mustard, Peter S. (2003). "Geology of Denman and Hornby islands, British Columbia: implications for Nanaimo Basin evolution and formal definition of the Geoffrey and Spray formations, Upper Cretaceous Nanaimo Group". Canadian Journal of Earth Sciences. 40 (3): 375–393. doi:10.1139/e03-005.
  8. ^ Squires, Richard L. (2017). "Late Cretaceous oysters from the Pacific slope of North America: Revision of named species and discovery of new species". Contributions in Science. 525: 25 53. doi:10.5962/p.318346.
  9. ^ Whiteaves, Joseph F. (1879). "On the fossils of the Cretaceous rocks of Vancouver and adjacent islands in the Strait of Georgia". Geological Survey of Canada. 1 (2): 93–190. doi:10.4095/216116.
  10. ^ Squires, Richard L.; Saul, Louella R. (2004). "Cretaceous corbulid bivalves of the Pacific slope of North America". The Veliger. 47 (2): 103–129.
  11. ^ Saul, L. R. (1993). "Pacific Slope Cretaceous bivalves: eight venerid species". Journal of Paleontology. 67 (6): 965–979. doi:10.1017/S0022336000025282.
  12. ^ Saul, Louella R.; Squires, Richard L. (2015). "Pacific slope of North America record of the Cretaceous Aporrhaid Gastropod Tessarolax: Evolutionary trends, mode of life, and paleobiogeography of the genus". Contributions in Science. 523: 37–65. doi:10.5962/p.318346.
  13. ^ Haggart, James W. (1989). "New and revised ammonites from the Upper Cretaceous Nanaimo group of British Columbia and Washington state". Geological Survey of Canada. 396: 181–221. doi:10.4095/127723.
  14. ^ a b Usher, J. L. (1952). "Ammonite faunas of the Upper Cretaceous rocks of Vancouver Island, British Columbia". Geological Survey of Canada. 21: 1–182. doi:10.4095/101522.
  15. ^ Jones, David L. (1963). "Upper Cretaceous (Campanian and Maestrichtian) ammonites from southern Alaska". U.S. Geological Survey Professional Paper. 432: 1–53. doi:10.3133/pp432.
  16. ^ Tanabe, Kazushige; Trask, Pat; Ross, Rick; Hikida, Yoshinori (2008). "Late Cretaceous octobrachiate coleoid lower jaws from the north Pacific regions". Journal of Paleontology. 82 (2): 398–408. doi:10.1666/07-029.1.
  17. ^ Larson, Neal L. (2010). "Fossil Coleoids from the Late Cretaceous (Campanian & Maastrichtian) of the Western Interior". Ferrantia. 59: 78–113.
  18. ^ Fuchs, Dirk; Iba, Yasuhiro; Heyng, Alexander; Iijima, Masaya; Klug, Christian; Larson, Neal L.; Schweigert, Günter (2020). "The Muensterelloidea: phylogeny and character evolution of Mesozoic stem octopods". Papers in Palaeontology. 6 (1): 31–92. doi:10.1002/spp2.1254.
  19. ^ a b c d Nyborg, Torrey; McLachlan, Sandy M. S.; Garassino, Alessandro; Vega, Francisco J.; Phillippe, Steven C.; Champagne, Donald E. (2019). "New and revised species of Archaeopus Rathbun, 1908 (Decapoda: Brachyura: Retroplumidae) from the eastern Pacific". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 292 (1): 25–56. doi:10.1127/njgpa/2019/0807.
  20. ^ Schweitzer, Carrie E.; Feldmann, Rodney M.; Fam, John; Hessin, William A.; Hetrick, Steven W.; Nyborg, Torrey G.; Ross, Richard L.M. (2003). Cretaceous and Eocene Decapod Crustaceans from Southern Vancouver Island, British Columbia, Canada. Ottawa, Ontario, Canada: NRC Research Press. p. 1–66. doi:10.1139/9780660190921. ISBN 0-660-19092-3.
  21. ^ Nyborg, Torrey; Garassino, Alessandro; Vega, Francisco J.; Ross, Richard L. M. (2024). "A new fossil frog crab (Brachyura, Raninoidea) from the late Campanian of Hornby Island (British Columbia, Canada)". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 310 (3): 209–217. doi:https://doi.org/10.1127/njgpa/2023/1177. {{cite journal}}: Check |doi= value (help); External link in |doi= (help)
  22. ^ Nyborg, Torrey; Ossó, Àlex; Vega, Francisco J. (2014). "A new species of icriocarcinid crab (Crustacea, Portunoidea) from the uppermost Cretaceous of California, USA: palaeobiogeographic implications". Scripta Geologica. 147: 83–93.
  23. ^ a b Nyborg, Torrey; Garassino, Alessandro; Ross, Richard L. M. (2020). "A new dromioid crab from the late Campanian of Hornby Island, Canada". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 295 (2): 141–147. doi:10.1127/njgpa/2020/0878.
  24. ^ a b c McGugan, Alan (1979). "Biostratigraphy and paleoecology of Upper Cretaceous (Campanian and Maestrichtian) foraminifera from the Upper Lambert, Northumberland, and Spray Formations, Gulf Islands, British Columbia, Canada". Canadian Journal of Earth Sciences. 16 (12): 2263–2274. doi:10.1139/e79-211.
  25. ^ a b c Cappetta, Henri; Morrison, Kurt; Adnet, Sylvain (2021). "A shark fauna from the Campanian of Hornby Island, British Columbia, Canada: an insight into the diversity of Cretaceous deep-water assemblages". Historical Biology. 33 (8): 1121–1182. doi:10.1080/08912963.2019.1681421.
  26. ^ Bogan, Sergio; Agnolín, Federico L. (December 2022). "The fossil record of the Bramble-shark Echinorhinus (Echinorhiniformes, Echinorhinidae) in South America". Journal of South American Earth Sciences. 120: 104083. doi:10.1016/j.jsames.2022.104083.
  27. ^ Pollerspöck, Jürgen; Beaury, Bernhard; Straube, Nicolas; Feichtinger, Iris (2021). "Oldest evidence of the genus Squalus in the north alpine realm with remarks on its evolution and distribution through time" (PDF). Paleoichthys. 2: 1–9.
  28. ^ Vullo, Romain; Buffetaut, Eric; Everhart, Michael J. (September 2012). "Reappraisal of Gwawinapterus beardi from the Late Cretaceous of Canada: a saurodontid fish, not a pterosaur". Journal of Vertebrate Paleontology. 32 (5): 1198–1201. doi:10.1080/02724634.2012.681078.
  29. ^ Arbour, Victoria M.; Currie, Philip J. (January 2011). "An istiodactylid pterosaur from the Upper Cretaceous Nanaimo Group, Hornby Island, British Columbia, Canada" (PDF). Canadian Journal of Earth Sciences. 48 (1): 63–69. doi:10.1139/E10-083.
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