Pygocephalid pancrustaceans from the late Carboniferous of Nova Scotia, Canada, with comments on the cuticular microstructures of Pygocephalamorpha
DOI:
https://doi.org/10.4138/atlgeo.2026.001Keywords:
Eumalacostraca, Joggins Fossil Cliffs, paleoecology, Systematics, PennsylvanianAbstract
Pygocephalomorphs are fossil eumalacostracans with a shrimp to lobster-like morphology, known from the Late Devonian (Famennian) to the early Permian (Cisuralian; Artinskian). In the late Paleozoic, pygocephalamorphs formed an important component of marginal marine, brackish and freshwater communities. Several species, including species of Tealliocaris and Pygocephalus, have been described from the Carboniferous of Canada, in particular from the Pennsylvanian. A revision of Pygocephalus from the Carboniferous of Nova Scotia, Canada, is proposed based on original material described by Copeland and on undescribed material from the Joggins Fossil Cliffs UNESCO World Heritage Site. The presence of three taxa is confirmed: Pygocephalus cooperi, Pygocephalus cf. cooperi and Pygocephalus dubius. The pygocephalomorphs of Joggins Fossil Cliffs, and more generally Nova Scotia, were likely benthic walkers based on their leg morphology and the paleoichnological evidence. A comparison of the studied assemblages with previously known assemblages from the Mississippian of Canada shows a faunal turnover among these assemblages, which could reflect paleoenvironmental changes in Atlantic Canada during the Carboniferous and thus various ecological affinities among taxa. We also observed three types of microstructures on the shield of Pygocephalus dubius: canal-like pores, which are most probably organule canals, setal attachment site, and polygonal reticular ornamentation.
References
Adami-Rodrigues, K., Pazinato, P.G., and Pinto, I.D. 2017. New species of Pygocephalomorpha (Crustacea) from the Irati Formation, Paraná Basin, RS, southern Brazil. Revista Brasileira de Paleontologia, 19, pp. 389–400. https://doi.org/10.4072/rbp.2016.3.04
Allen, J.P., Fielding, C.R., Gibling, M.R., and Rygel, M.C. 2011. Fluvial response to paleo-equatorial climate fluctuations during the late Paleozoic ice age. Geological Society of America Bulletin, 123, pp. 1524–1538. https://doi.org/10.1130/B30314.1
Anderson, L.I., Dunlop, J.A., Horrocks, C.A., Winkelmann, H.M., and Eagar, R.M.C. 1997. Exceptionally preserved fossils from Bickershaw, Lancashire UK (Upper Carboniferous, Westphalian A (Langsettian)). Geological Journal, 32, pp. 197–210. https://doi.org/10.1002/(SICI)1099-1034(199709)32:3<197::AID-GJ739>3.0.CO;2-6
Bell, W.A. 1922. A new genus of Characeae and new Merostomata from the Coal Measures of Nova Scotia. Transactions of the Royal Society of Canada, 16, pp. 159–167.
Beurlen, K. 1930. Vergleichende Stammesgeschichte: Grundlagen, Methoden, Probleme unter besonderer Berücksichtigung der höheren Krebse. Fortschritte der Geologie und Palaontologie, 8, pp. 317–586.
Birshtein, Y.A. 1966. Notes on Paleozoic higher crustaceans (Malacostraca) SSSR. Pygocephalomorpha. Paleontologicheskii Zhurnal, 1966, pp. 49–56 (in Russian).
Briggs, D.E.G. and Clarkson, E.N.K. 1985. The Lower Carboniferous shrimp Tealliocaris from Gullane, East Lothian, Scotland. Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 76, pp. 173–201. https://doi.org/10.1017/S0263593300010439
Broda, K. and Zatoń, M. 2017. A set of possible sensory system preserved in cuticle of Late Devonian thylacocephalan arthropods from Poland. Historical Biology, 29, pp. 1045–1055. https://doi.org/10.1080/08912963.2017.1284834
Brooks, H.K. 1962. The Paleozoic Eumalacostraca of North America. Bulletins of American Paleontology, 44, pp. 159–338.
Brooks, H.K. and Moore, R.C. 1969. Eocarida. In Treatise on Invertebrate Paleontology: Part R, Arthropoda 4. Edited by H.K. Brooks, V.A. Zullo, M.F. Glaessner, W.A. Newman, R.C. Moore, P. Tasch, L. McCormick, S.M. Manton, R.R. Hessler, R.B. Manning, W.D.I. Rolfe, F.M. Carpenter, L.B. Holthuis, R.L. Hoffman, G. Hahn, T.H. Withers, and A.R. Palmer. University of Kansas and The Geological Society of America, Lawrence, Kansas. pp. R332–R345.
Broom, R. 1931. On the Pygocephalus-like crustacean of the South African Dwyka. Proceedings of the Zoological Society of London, 101, pp. 571–574. https://doi.org/10.1111/j.1096-3642.1931.tb01032.x
Browne, G.H. and Plint, A.G. 1994. Alternating braidplain and lacustrine deposition in a strike-slip setting; the Pennsylvanian Boss Point Formation of the Cumberland Basin, Maritime Canada. Journal of Sedimentary Research, 64, pp. 40–59. https://doi.org/10.1306/D4267F41-2B26-11D7-8648000102C1865D
Budil, P. and Hoerbinger, F. 2007. Exoskeletal structures and ultrastructures in Lower Devonian dalmanitid trilobites of the Prague Basin (Czech Republic). Bulletin of Geosciences, 82, pp. 27–36.
Calder, J.H. 1998. The Carboniferous evolution of Nova Scotia. Geological Society, London, Special Publications, 143, pp. 261–302. https://doi.org/10.1144/GSL.SP.1998.143.01.19
Chaudoir, H. 1950. Étude du gisement houiller de la Campine — contribution à l’étude stratigraphique et paléontologique du Westphalien B supérieur: la zone d’Eikenberg. Association pour l’Étude de la Paléontologie et de la Stratigraphie houillères, 5, pp. 3–87.
Clark, N.D.L. 2013. Tealliocaris: a decapod crustacean from the Carboniferous of Scotland. Palaeodiversity, 6, pp. 107–133.
Clark, N.D.L. 2024. New malacostracan crustaceans from the Lower Coal Measures (Langsettian, Carboniferous) north Lanarkshire, Scotland. Scottish Journal of Geology, 60, sjg2024-004, 8 p. https://doi.org/10.1144/sjg2024-004
Clark, N.D.L. and Ross, A.J. 2024. Caridoid crustaceans from the Ballagan Formation (Tournaisian, Lower Carboniferous) of Willie’s Hole, Chirnside, Scottish Borders, UK. Earth and Environmental Science Transactions of The Royal Society of Edinburgh, 115, pp. 1–12. https://doi.org/10.1017/S1755691024000045
Clark, N.D.L., Miller, R.F., and Ross, A.J. 2016. The distribution of Schramocaris (Eumalacostraca, Crustacea) along the northwestern coast of the Rheic Ocean during the Lower Carboniferous. Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 107, pp. 91–98. https://doi.org/10.1017/S1755691017000159
Copeland, M.J. 1957. The arthropod fauna of the upper Carboniferous rocks of the Maritime Provinces. E. Cloutier, Queen’s Printer, Ottawa, 110 p. https://doi.org/10.4095/101505
Crawford, T. 1995. Carbonates and associated sedimentary rocks of the upper Visean to Namurian Mabou Group, Cape Breton Island, Nova Scotia: evidence for lacustrine deposition. Atlantic Geology, 31, pp. 167–182. https://doi.org/10.4138/2110
Dalingwater, J.E., Siveter, D.J., and Mutvei, H. 1999. Cuticular microstructure of some Silurian homalonotid trilobites from Sweden. Journal of Paleontology, 73, pp. 256–262. https://doi.org/10.1017/S002233600002775X
Dalinval, A. 1947. Découverte d’Anthrapalaemon et d’un second horizon à Leaia dans le Terrain houiller du Nord. Annales de la Société Géologique du Nord, 67, pp. 27–33.
Davies, S.J. and Gibling, M.R. 2003. Architecture of coastal and alluvial deposits in an extensional basin: the Carboniferous Joggins Formation of eastern Canada. Sedimentology, 50, pp. 415–439. https://doi.org/10.1046/j.1365-3091.2003.00553.x
Davies, S.J., Gibling, M.R., Rygel, M.C., Calder, J.H., and Skilliter, D.M. 2005. The Pennsylvanian Joggins Formation of Nova Scotia: sedimentological log and stratigraphic framework of the historic fossil cliffs. Atlantic Geology, 41, pp. 87–102. https://doi.org/10.4138/182
Davies, S.J., Gibling, M.R., Rygel, M.C., Calder, J.H., and Skilliter, D.M. 2006. The Pennsylvanian Joggins Formation of Nova Scotia: sedimentological log and stratigraphic framework of the historic fossil cliffs. Atlantic Geology, 41, pp. 115–142. https://doi.org/10.4138/182
Dawson, J.W. 1864. On the fossils of the genus Rusophycus. Canadian Naturalist and Geologist, 1, pp. 363–367.
Dawson, J.W. 1872. V.—Note on footprints from the Carboniferous of Nova Scotia, in the collection of the Geological Survey of Canada. Geological Magazine, 9, pp. 251–253. https://doi.org/10.1017/S0016756800465052
Dawson, J.W. 1877. Note on two Palaeozoic crustaceans from Nova Scotia. The Geological Magazine, 4, pp. 56–58. https://doi.org/10.1017/S0016756800148587
Dawson, J.W. 1878. Supplement to the second edition of Acadian geology: containing additional facts as to the geological structure, fossil remains, and mineral resources of Nova Scotia, New Brunswick, and Prince Edward Island. Macmillan and Company, 144 p. https://doi.org/10.5962/bhl.title.18064
Dewey, C.P. and Fåhræus, L.E. 1982. Peracarids (Crustacea) from Mississippian strata of western Newfoundland. Canadian Journal of Earth Sciences, 19, pp. 666–670. https://doi.org/10.1139/e82-055
Dix, E. 1930. An arthropod (Anthrapalaemon cf. russelianus (Salter)), from the North Staffordshire coalfield. Naturalist, March, pp. 103–104.
Edwards, W. and Stubblefield, C.J. 1947. Marine bands and other faunal marker-horizons in relation to the sedimentary cycles of the Middle Coal Measures of Nottinghamshire and Derbyshire. Quarterly Journal of the Geological Society of London, 103, pp. 209–260. https://doi.org/10.1144/GSL.JGS.1947.103.01-04.16
Etheridge, R. 1877. On the occurrence of a macrurous decapod (Anthrapalæmon? Woodwardi, sp. nov.) in the Red Sandstone, or lowest group, of the Carboniferous Formation in the south-east of Scotland. Quarterly Journal of the Geological Society of London, 33, pp. 863–878. https://doi.org/10.1144/GSL.JGS.1877.033.01-04.49
Garm, A. and Watling, L. 2015. The crustacean integument: setae, setules, and other ornamentation. In Functional morphology and diversity. Edited by L. Watling, L. and M. Thiel. Oxford University Press, New-York. pp. 167–198. https://doi.org/10.1093/acprof:osobl/9780195398038.003.0006
Grobben, K. 1892. Zur Kenntniss des Stammbaumes und des Systems der Crustaeeen. Sitzungsberichte. Abt. 1, Mineralogie, Krystallographie, Botanik, Physiologie der Pflanzen, Zoologie, Paläontologie, Geologie, physische Geographie und Reisen, 101, pp. 237–274.
Gueriau, P., Charbonnier, S., and Clément, G. 2014. First decapod crustaceans in a Late Devonian continental ecosystem. Palaeontology, 57, pp. 1203–1213. https://doi.org/10.1111/pala.12111
Hessler, R.R. 1983. A defense of the caridoid facies; wherein the early evolution of the Eumalacostraca is discussed. Crustacean Issues, 1, pp. 145–164.
Huxley, F.R.S. 1857. Description of a new crustacean (Pygocephalus Cooperi, Huxley) from the Coal-Measures. Quarterly Journal of the Geological Society of London, 13, pp. 363–369. https://doi.org/10.1144/GSL.JGS.1857.013.01-02.45
Jones, T.R. and Woodward, H. 1899. II.—Contributions to fossil Crustacea. Geological Magazine, 6, pp. 388–395. https://doi.org/10.1017/S0016756800142487
Jones, W.T., Feldmann, R.M., Schram, F.R., Schweitzer, C.E., and Maguire, E.P. 2016. The proof is in the pouch: Tealliocaris is a peracarid. Palaeodiversity, 9, pp. 75–88. https://doi.org/10.18476/pale.v9.a5
Jongmans, W.J. 1928. Stratigraphie van het Karboon in het algemeen en van Limburg in het bijonder. Mededeeling van het Geologisch Bureau voor het Nederlandsche Mijngebied te Heerlen, 6, pp. 1–50.
Keppie, J.D. 2000: Geological map of the Province of Nova Scotia. Nova Scotia Department of Natural Resources, Minerals and Energy Branch, map ME 2000-1, scale 1:500 000.
Kunkel, J. 2013. Modeling the calcium and phosphate mineralization of American lobster cuticle. Canadian Journal of Fisheries and Aquatic Sciences, 70, pp. 1601–1611. https://doi.org/10.1139/cjfas-2013-0034
Kunkel, J., Nagel, W., and Jercinovic, M. 2012. Mineral fine structure of the American lobster cuticle. Journal of Shellfish Research, 31, pp. 515–526. https://doi.org/10.2983/035.031.0211
Latreille, P.A. 1802. Histoire naturelle, générale et particulière, des crustacés et des insectes. Imprimerie de F. Dufart, Paris, 476 p. https://doi.org/10.5962/bhl.title.15764
Laville, T., Haug, C., Haug, J.T., Forel, M.-B., and Charbonnier, S. 2021. Morphology and anatomy of the Late Jurassic Mayrocaris bucculata (Eucrustacea?, Thylacocephala) with comments on the tagmosis of Thylacocephala. Journal of Systematic Palaeontology, 19, pp. 289–320. https://doi.org/10.1080/14772019.2021.1910584
Lerosey-Aubril, R. and Meyer, R. 2013. The sensory dorsal organs of crustaceans. Biological Reviews, 88, pp. 406–426. https://doi.org/10.1111/brv.12011
Lima, J.H.D., Minter, N.J., and Netto, R.G. 2017. Insights from functional morphology and neoichnology for determining tracemakers: a case study of the reconstruction of an ancient glacial arthropod-dominated fauna. Lethaia, 50, pp. 576–590. https://doi.org/10.1111/let.12214
Malay, B.C., Stimson, M.R., King, O.A., Faulkner, S., Heberts, B.L., Calder, J.H., MacRae, R.A., and Lucas, S.G. 2024. A new cubichnium ichnogenus and ichnospecies, Pygocephalichnium reidi, from the Pennsylvanian UNESCO World Heritage Site at Joggins Fossil Cliffs, Canada, and associated ichnotaxa. Atlantic Geoscience, 60, pp. 185–203. https://doi.org/10.4138/atlgeo.2024.009
Melchor, R.N. and Cardonatto, M.C. 2014. Insights on the behavior of late Paleozoic aquatic crustaceans (Pygocephalomorpha?): compound trace fossils from western Argentina. Ichnos, 21, pp. 76–99. https://doi.org/10.1080/10420940.2013.879868
Miller, R.F. and Purdy, S.C. 1998. Lower Carboniferous Tealliocaris woodwardi (Crustacea, Malacostraca) from the Macumber Formation, New Brunswick (NTS 21 H/5). New Brunswick Department of Natural Resources and Energy, Minerals and Energy Division, Mineral Resources Report, 98, pp. 17–24.
Mossman, D.J. and Grantham, R.G. 2000. Vertebrate trackways in the Parrsboro Formation (Upper Carboniferous) at Rams Head, Cumberland County, Nova Scotia. Atlantic Geology, 35, pp. 186–196. https://doi.org/10.4138/2032
Naimark, E.B. and Chaika, S.Y. 2025. The cuticle of Agnostina (basal Euarthropoda) was three-layered. Arthropod Structure and Development, 86, 101447, 18 p. https://doi.org/10.1016/j.asd.2025.101447
Naylor, R.D. and C.M. Kennedy. 1997. Stratigraphy and sedimentology of the late Carboniferous Parrsboro Formation. In Report of Activities, Nova Scotia Department of Natural Resources 1997, pp. 115–126.
Paproth, E. and Schmidt, W. 1955. Ein Anthrapalaemon-Fund aus sehr hohem Karbon. Paläontologische Zeitschrift, 29, pp. 83–87. https://doi.org/10.1007/BF03041758
Pastiels, A. 1951. Étude du gisement houiller de la Campine — contribution à l’étude stratigraphique et paléontologique du Westphalien B: quelques éléments de la faune non marine. Association pour l’Étude de la Paléontologie et de la Stratigraphie houillères, 7, pp. 3–18.
Pazinato, P., Soares, M.B., and Adami‐Rodrigues, K. 2016. Systematic and palaeoecological significance of the first record of Pygocephalomorpha females bearing oöstegites (Malacostraca, Peracarida) from the lower Permian of southern Brazil. Palaeontology, 59, pp. 817–826. https://doi.org/10.1111/pala.12260
Pazinato, P., Haug, C., Leipner, A., and Haug, J. 2022. Pygocephalomorphan crustaceans further emphasise the similarities between the Carboniferous Piesberg quarry in Germany and the Mazon Creek Lagerstätte in North America. Palaeontologia Electronica, 25, 25.1.a2, 21 p. https://doi.org/10.26879/1051
Peach, B.N. 1908. Monograph on the higher Crustacea of the Carboniferous rocks of Scotland. Printed for H.M. Stationery Office by J. Hedderwick and Sons, Ltd, Glasgow, 82 p. https://doi.org/10.5962/bhl.title.10510
Peach, B.N. 1914. On some Carboniferous arthropods, with description of a new genus of myriapod. Proceedings of the Royal Physical Society of Edinburgh, 19, pp. 142–148.
Piñeiro, G., Morosi, E., Ramos, A., and Scarabino, F. 2012. Pygocephalomorph crustaceans from the early Permian of Uruguay: constraints on taxonomy. Revista Brasileira de Paleontologia, 15, pp. 33–48. https://doi.org/10.4072/rbp.2012.1.03
Pinto, I.D. and Adami-Rodrigues, K. 1996. Pygocephalomorph Crustacea. New data and interpretations, with emphasis on Brazilian and South African forms. Pesquisas em Geociências, 23, pp. 41–50. https://doi.org/10.22456/1807-9806.21226
Pinto, I.D. and Würdig, N.L. 2014. Contribution to the knowledge of the genus Pygocephalus Huxley, 1857: morphology and taxonomy. Revista Brasileira de Paleontologia, 17, pp. 275–288. https://doi.org/10.4072/rbp.2014.3.01
Piper, D.J.W. and Pe-Piper, G. 2021. Evolution of late Paleozoic shearing in the Cobequid Highlands: constraints on the fragmentation of the Appalachian Orogen in Nova Scotia along intra-continental shear zones. In Pannotia to Pangaea: Neoproterozoic and Paleozoic orogenic cycles in the Circum-Atlantic Region. Edited by J.B. Murphy, R.A. Strachan, and C. Quesada. Geological Society, Special Publication, 503, pp. 1–20. https://doi.org/10.1144/SP503-2019-239
Prestwich, J. 1840. XXXIII.—On the geology of Coalbrook Dale. Transactions of the Geological Society of London, 5, pp. 413–495. https://doi.org/10.1144/transgslb.5.3.413
Pruvost, P. 1930. La faune continentale du terrain houiller de la Belgique. Mémoires du Musée royale d’Histoire naturelle de Belgique, 44, pp. 103–280.
Renier, A., Stockmans, F., Demanet, F., and Van Straelen, V. 1938. Flore et faune houillères de la Belgique: introduction à l’é̕tude paléontologique du terrain houiller. Patrimoine du Musée royal d’histoire naturelle de Belgique, Bruxelles, 366 p.
Rhodes, F.H.T. and Wilson, A.A. 1957. The arthropod species Anthrapalaemon dubius and related forms. Journal of Paleontology, 31, pp. 1159–1166.
Roer, R. and Dillaman, R. 1984. The structure and calcification of the crustacean cuticle. American Zoologist, 24, pp. 893–909. https://doi.org/10.1093/icb/24.4.893
Rolfe, W.D.I. 1962. The cuticle of some middle Silurian ceratiocaridid Crustacea from Scotland. Palaeontology, 5, pp. 30–51.
Ryan, R.J., Boehner, R.C., and Calder, J.H. 1991. Lithostratigraphic revisions of the upper Carboniferous to lower Permian strata in the Cumberland Basin, Nova Scotia and the regional implications for the Maritimes Basin in Atlantic Canada. Bulletin of Canadian Petroleum Geology, 39, pp. 289–314.
Rygel, M.C., Sheldon, E.P., Stimson, M.R., Calder, J.H., Ashley, K.T., and Salg, J.L. 2014. The Pennsylvanian Springhill Mines Formation: sedimentological framework for a portion of the Joggins Fossil Cliffs UNESCO World Heritage Site. Atlantic Geology, 50, pp. 249–289. https://doi.org/10.4138/atlgeol.2014.013
Salter, J.W. 1861. On some of the higher Crustacea from the British Coal-Measures. Quarterly Journal of the Geological Society of London, 17, pp. 528–533. https://doi.org/10.1144/GSL.JGS.1861.017.01-02.44
Salter, J.W. 1863a. On some fossil Crustacea from the Coal-Measures and Devonian rocks of British North America. Quarterly Journal of the Geological Society of London, 19, pp. 75–80. https://doi.org/10.1144/GSL.JGS.1863.019.01-02.14
Salter, J.W. 1863b. On a new crustacean from the Glasgow Coal-field. Quarterly Journal of the Geological Society of London, 19, pp. 519–521. https://doi.org/10.1144/GSL.JGS.1863.019.01-02.51
Savage, N.M. 1971. A varvite ichnocoenosis from the Dwyka series of Natal. Lethaia, 4, pp. 217–233. https://doi.org/10.1111/j.1502-3931.1971.tb01290.x
Schenk, P.E., Matsumoto, R., and von Bitter, P.H. 1994. Loch Macumber (early Carboniferous) of Atlantic Canada. Journal of Paleolimnology, 11, pp. 151–172. https://doi.org/10.1007/BF00686863
Schindelin, J., Arganda-Carreras, I., Frise, E., Kaynig, V., Longair, M., Pietzsch, T., Preibisch, S., Rueden, C., Saalfeld, S., Schmid, B., Tinevez, J.-Y., White, D.J., Hartenstein, V., Eliceiri, K., Tomancak, P., and Cardona, A. 2012. Fiji: an open-source platform for biological-image analysis. Nature Methods, 9, pp. 676–682. https://doi.org/10.1038/nmeth.2019
Schram, F.R. 1969. The stratigraphic distribution of the Paleozoic Eumalacostraca. Fieldiana Geology, 12, pp. 213–234. https://doi.org/10.5962/bhl.title.5338
Schram, F.R. 1974. The Mazon Creek caridoid Crustacea. Fieldiana Geology, 30, pp. 9–65. https://doi.org/10.5962/bhl.title.3154
Schram, F.R. 1979. British Carboniferous Malacostraca. Fieldiana Geology, 40, pp. 1–129.
Schram, F.R. 1980. Pygocephalus from the Upper Carboniferous of the Soviet Union. Journal of Paleontology, 54, pp. 50–56.
Schram, F.R. 1981. Late Paleozoic crustacean communities. Journal of Paleontology, 55, pp. 126–137.
Schram, F.R. and Koenemann, S. 2022. Evolution and phylogeny of Pancrustacea: a story of scientific method. Oxford University Press, New York, 872 p. https://doi.org/10.1093/oso/9780195365764.001.0001
Sjoberg, D.D., Whiting, K., Curry, M., Lavery, J., and Larmarange, J. 2021. Reproducible summary tables with the gtsummary package. The R Journal, 13, pp. 570. https://doi.org/10.32614/RJ-2021-053
Taylor, R.S., Taylor, R.S., Shen, Y.-B., and Schram, F.R. 1998. New pygocephalomorph crustaceans from the Permian of China and their phylogenetic relationships. Palaeontology, 41, pp. 815–834.
Teigler, D.J. and Towe, K.M. 1975. Microstructure and composition of the trilobite exoskeleton. Fossils and Strata, 4, pp. 137–149. https://doi.org/10.18261/8200049639-1975-07
Van der Heide, S. 1951. Les arthropodes du terrain houiller du Limbourg méridional. Mededeelingen van de Geologische Stichting Serie C, IV. Paleontologie, 5, pp. 1–84.
Waldron, J.W., Rygel, M.C., Gibling, M.R., and Calder, J.H. 2013. Evaporite tectonics and the late Paleozoic stratigraphic development of the Cumberland Basin, Appalachians of Atlantic Canada. Geological Society of America Bulletin, 125, pp. 945–960. https://doi.org/10.1130/B30718.1
Waugh, D.A., Feldmann, R.M., and Schweitzer, C.E. 2009. Systematic evaluation of raninid cuticle microstructure. Bulletin of the Mizunami Fossil Museum, 35, pp. 15–41.
Wheatly, M. 1993. Physiological adaptations in decapodan crustaceans for life in fresh water. In Advances in comparative and environmental physiology. Edited by I. Novak, M. Hidaka, M. Wheatly, J. Balthazart, and G.F. Ball. Springer, Berlin, Heidelberg. pp. 77–132. https://doi.org/10.1007/978-3-642-77528-4_3
Woodward, H. 1867. LXIII. Notes on some fossil Crustacea and a chilognathous myriapod, from the Coal Measures of the west of Scotland. Transactions of the Geological Society of Glasgow, 2, pp. 234–248. https://doi.org/10.1144/transglas.2.3.234
Woodward, H. 1879. On Necroscilla wilsoni, a supposed stomatopod crustacean from the Middle Coal-measures, Cossall near Ilkeston. Quarterly Journal of the Geological Society of London, 35, pp. 551–552.
Woodward, H. 1905. II.—Notes on some crustaceans and two myriapods from the Lower Coal-Measures near Colne, Lancashire. The Geological Magazine, 2, pp. 437–444. https://doi.org/10.1017/S0016756800128365
Woodward, H. 1907. IV. On the genus Pygocephalus (Huxley), a primitive schizopod crustacean, from the Coal-Measures. Geological Magazine, 4, pp. 400–407. https://doi.org/10.1017/S0016756800133746
Woodward, H. 1911. VI.—On a large form of Anthrapalæmon from the clay-ironstone nodules of the Middle Coal-Measures, Sparth Bottoms, Rochdale. Geological Magazine, 8, pp. 361–366. https://doi.org/10.1017/S0016756800139251
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