Patterns of evolution and extinction in the last harpetid trilobites during the Late Devonian (Frasnian)

52 1 January 11 33 10.1111/j.1475-4983.2008.00832.x

MCNAMARA, K. J., FEIST, R., EBACH, M. C. 2009. Patterns of evolution and extinction in the last harpetid trilobites during the Late Devonian (Frasnian). Palaeontology52, 1, 11–33.

Kenneth J. McNamara, Raimund Feist and Malte C. Ebach Late Devonian (Frasnian) harpetid trilobites have hitherto only been described from the western side of the Protethys Ocean, in what is now Europe and North Africa, as well as from Gondwana-derived northwestern Kazakhstan (Mugodjar). However, late Frasnian strata in the Canning Basin, Western Australia, that were deposited on the eastern side of this ocean, contain a rich harpetid fauna. Described herein are two new harpetids: Eskoharpes gen. nov. and Globoharpes gen. nov., within which are placed six species: E. palanasus sp. nov., E. wandjina sp. nov., E. boltoni sp. nov., E. guthae sp. nov., G. teicherti sp. nov. and G. friendi sp. nov. The ontogenetic development of E. palanasus, E. wandjina and G. teicherti are described, including the first unequivocal harpetid protaspis. Globoharpes exhibits evidence of sexual dimorphism in the development of a pronounced preglabellar boss in some specimens. This structure is thought to have functioned as a brood pouch. Such structures have previously only been described in Cambrian and Ordovician trilobites, and never before in harpetids. It is suggested that the characteristic harpetid fringe functioned as a secondary respiratory structure. The Eskoharpes lineage shows evolutionary trends that mirror changes seen in ontogenetic development of the youngest species, suggesting the operation of peramorphic processes. This is the first record of heterochrony in harpetids and the first documented example of peramorphosis in Devonian trilobites. These harpetids demonstrate a stepped pattern of extinction during the late Frasnian, probably related to the effects of the two Kellwasser biocrises that have been well documented in European Frasnian sections. Highly vaulted species of Eskoharpes and the strongly vaulted Globoharpes became extinct at the Lower Kellwasser Event. The flatter species of Eskoharpes became extinct at the base of the Upper Kellwasser Event shortly prior to the Frasnian/Famennian boundary. The extinction of these harpetids, along with contemporaneous forms from Europe, which are also discussed herein, marks the end of the trilobite order Harpetida worldwide.
  • BARRIENTOS, Y. and LAVERACK, M. S. 1986. The larval crustacean dorsal organ and its relationship to the trilobite median tubercle. Lethaia, 19, 309–313.
  • BECKER, R. T., FEIST, R., FLAJS, G., HOUSE, M. R. and KLAPPER, G. 1989. Frasnian–Famennian extinction events in the Devonian at Coumiac, southern France. Comptes Rendus Académie des Sciences de Paris, 309, 259–266.
  • BECKER, R. T., HOUSE, M. R., KIRCHGASSER, W. T. and PLAYFORD, P. E. 1991. Sedimentary and faunal changes across the Frasnian/Famennian boundary in the Canning Basin of Western Australia. Historical Biology, 5, 183–196.
  • CHATTERTON, B. D. E. 1980. Ontogenetic studies of Middle Ordovician trilobites from the Esbataottine Formation; Mackenzie Mountains, Canada. Palaeontographica (Abt A), 171, 1–74.
  • CHATTERTON, B. D. E. and SPEYER, S. E. 1997. Ontogeny. 173–247. In WHITTINGTON, H. B., CHATTERTON, B. D. E., SPEYER, S. E., FORTEY, R. A., OWENS, R. M., CHANG, W. T., DEAN, W. T., JELL, P. A., LAURIE, J. R., PALMER, A. R., REPINA, L. M., RUSHTON, A. W. A., SHERGOLD, J. H., CLARKSON, E. N. K., WILMOT, N. V. and KELLY, S. R. A. Treatise on invertebrate paleontology. Part O, Revised, Trilobita, Introduction, Order Agnostina, Order Redlichiida. Geological Society of America, Boulder and University of Kansas, Lawrence, 530 pp.
  • CRÔNIER, C. and FEIST, R. 1997. Morphologie et évolution ontogénétique de Trimerocephalus lelievrei nov. sp., premier trilobite phacopidé aveugle du Famennien nord-African. Geobios, Memoire Spécial, 20, 161–170.
  • CRÔNIER, C., RENAUD, S., FEIST, R. and AUFFRAY, J.-C. 1998. Ontogeny of Trimerocephalus lelievrei (Trilobita, Phacopida), a representative of the Late Devonian phacopine paedomorphocline: a morphometric approach. Paleobiology, 24, 359–370.
  • DRUCE, E. C. 1976. Conodont biostratigraphy of the Upper Devonian reef complexes of the Canning Basin, Western Australia. Australian Bureau Mineral Resources, Bulletin, 158, 1–303.
  • EBACH, M. and McNAMARA, K. J. 2002. A systematic revision of the family Harpetida (Trilobita). Records of the Western Australian Museum, 21, 235–267.
  • FEIST, R. 1991. The Late Devonian trilobite crises. Historical Biology, 5, 197–214.
  • FEIST, R. 1995. Effect of paedomorphosis in eye reduction on patterns of evolution and extinction in trilobites. 225–244. In McNAMARA, K. J. (ed.). Evolutionary change and heterochrony. Wiley, Chichester, 286 pp.
  • FEIST, R. 2002. Trilobites from the latest Frasnian Kellwasser Crisis in North Africa (Mrirt, central Moroccan Meseta). Acta Palaeontologica Polonica, 47, 203–223.
  • FEIST, R. and McNAMARA, K. J. 2007. Biodiversity, distribution and patterns of extinction of the last odontopleuroid trilobites during the Devonian (Givetian, Frasnian). Geological Magazine, 144, 777–796.
  • FEIST, R. and SCHINDLER, E. 1994. Trilobites during the Frasnian Kellwasser Crisis in European Late Devonian cephalopod limestones. Courier Forschungs-Institut Senckenberg, 169, 195–223.
  • FEIST, R., McNAMARA, K., CRÔNIER, C. and LEROSEY-AUBRIL, R. 2009. Patterns of extinction and recovery of phacopid trilobites during the Frasnian–Famennian (Late Devonian) mass extinction event, Canning Basin, Western Australia. Geological Magazine, 146, 12–33.
  • FORTEY, R. A. 2000. Olenid trilobites: the oldest known chemoautotrophic symbionts? Proceedings of the National Academy of Science, 97, 6574–6578.
  • FORTEY, R. A. and CLARKSON, E. N. K. 1976. The function of the glabellar ‘tubercle’ in Nileus and other trilobites. Lethaia, 9, 101–106.
  • FORTEY, R. A. and HUGHES, N. C. 1998. Brood pouches in trilobites. Journal of Paleontology, 72, 638–649.
  • FORTEY, R. A. and OWENS, R. M. 1999. Feeding habits in trilobites. Palaeontology, 42, 429–465.
  • GEORGE, A. D. and CHOW, N. 2002. The depositional record of the Frasnian/Famennian boundary interval in a fore-reef succession, Canning Basin, Western Australia. Palaeogeography, Palaeoclimatology, Palaeoecology, 181, 347–374.
  • GIRARD, C., KLAPPER, G. and FEIST, R. 2005. Subdivision of the terminal Frasnian linguiformis conodont Zone, revision of the correlative interval of Montagne Noire Zone 13, and discussion of stratigraphically significant associated trilobites. 181–198. In OVER, J. R., MORROW, J. R. and WIGNALL, P. B. (eds). Understanding Late Devonian and Permian–Triassic biotic and climatic events: towards an integrated approach. Elsevier, Amsterdam.
  • GLAUERT, L. 1910. New fossils from the Napier Range, Kimberley. Geological Survey of Western Australia, Bulletin, 36, 111–114.
  • GLENISTER, B. F. and KLAPPER, G. 1966. Upper Devonian conodonts from the Canning Basin, Western Australia. Journal of Paleontology, 40, 777–842.
  • GOLDFUSS, A. 1839. Beitrage zur Petrefactenkunde. Verhandlung K. Leopold-Caroline Akademie fur Naturforschung, 19, 327–364.
  • HAHN, G. and HAHN, R. H. 1975. Die Trilobiten des Ober-Devon, Karbon und Perm. Gebrüder Borntraeger, Berlin.
  • HAWLE, I. and CORDA, A. J. C. 1847. Prodrom einer Monographie der bohmischen Trilobiten. Abhandlungen der Koniglichen Bohemischen Gesellschaft der Wissenschaften, 5, 1–176. J. C. Calve, Prague.
  • JELL, P. A. 1978. Trilobite respiration and genal caeca. Alcheringa, 2, 251–260.
  • KLAPPER, G. 1988. The Montagne Noire Frasnian (Upper Devonian) conodont succession. Canadian Society of Petroleum Geologists, Memoir, 14, 449–468.
  • KLAPPER, G. 2007. Frasnian (Upper Devonian) conodont succession at Horse Spring and correlative sections, Canning Basin, Western Australia. Journal of Paleontology, 81, 513–537.
  • LEROSEY-AUBRIL, R. and McNAMARA, K. J. 2008. The cephalic median organ of trilobites. 229–235. In RABANO, I., GOZALO, R. and GARCíA-BELLIDO, D. (eds). Advances in trilobite research. Cuadernos del Museo Geominero 9, Instituto Geológia de España, Madrid.
  • MARTIN, J. W. and LAVERACK, M. S. 1992. On the distribution of the crustacean dorsal organ. Acta Zoologica, 73, 357–368.
  • MAXIMOVA, Z. A. 1955. Trilobity sretsnego i verkhnego Devona urala i severnykh Mougodzhar. [Trilobites of the Middle and Upper Devonian of the Urals and northern Mugodzhar Hills]. Trudy Vsesoyuznogo nauchno-issledovatelskogo Geologischeskogo Instituta, 3, 1–263.
  • McNAMARA, K. J. 1986. The role of heterochrony in the evolution of Cambrian trilobites. Biological Reviews, 61, 121–156.
  • McNAMARA, K. J. and FEIST, R. 2006. New styginids from the Late Devonian of Western Australia – the last corynexochid trilobites. Journal of Paleontology, 80, 981–992.
  • McNAMARA, K. J., YU, F. and ZHOU, Z. 2003. Ontogeny and heterochrony in the oryctocephalid trilobite Arthricocephalus from the Early Cambrian of China. Special Papers in Palaeontology, 70, 103–126.
  • McNAMARA, K. J., YU, F. and ZHOU, Z. 2006. Ontogeny and heterochrony in the oryctocephalid trilobites Changaspis, Duyunaspis and Balangia from the Lower Cambrian of China. Palaeontology, 48, 1–19.
  • NICOLL, R. S. 1984. Conodont distribution in the marginal-slope facies of the Upper Devonian reef complex, Canning Basin, Western Australia. Geological Society of America, Special Papers, 19, 127–141.
  • PLAYFORD, P. E. 1980. Devonian ‘Great Barrier Reef’ of the Canning Basin, Western Australia. American Association of Petroleum Geologists, 64, 814–840.
  • PLAYFORD, P. E. 1981. Devonian reef complexes of the Canning Basin, Western Australia. Fifth Australian Geological Convention Field Excursion Guidebook, Geological Society of Australia, Sydney1–64.
  • PLAYFORD, P. E.. 1984. Platform-margin and marginal-slope relationships in Devonian reef complexes of the Canning Basin. 189–213. In PURCELL, P. G. (ed.). The Canning Basin. Proceedings GSA/PESA Canning Basin Symposium, Perth.
  • PLAYFORD, P. E. and LOWRY, D. C. 1966. Devonian reef complexes of the Canning Basin, Western Australia. Geological Survey of Western Australia, Bulletin, 118, 1–150.
  • PRANTL, F. and PŘIBYL, A. 1954. On the Bohemian representatives of the family Harpedidae (Hawle and Corda). Rozpravy ústředního ústavu geologického, 18, 121–180.
  • PŘIBYL, A. and VANĚK, J. 1986. A study of the morphology and phylogeny of the family Harpetidae Hawle and Corda, 1847 (Trilobita). Sborník Národního Muzea v Praze, 42B, 1–72.
  • RICHTER, R. 1920. Von Bau und Leben der Trilobiten II. Der Aufenthalt auf dem Boden. Der Schutz. Die Ernhrung. Senckenbergiana, 2, 23–43.
  • RICHTER, R. 1921. Beitrge zur Kenntnis devonischer Trilobiten. III. Über die Organisation von Harpes, einen Sonderfall unter Crustaceen. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 37, 179–218.
  • RICHTER, R. and RICHTER, E. 1924. Unterlagen zum Fossilium Catalogus, Trilobita I. Senckenbergiana, 6, 229–233.
  • RICHTER, R. and RICHTER, E. 1926. Die Trilobiten der Oberdevons. Beitrge zur Kenntnis devonischer Trilobiten. IV. Abhandlungen der Preussischen Geologischen Landesanstalt, 99, 1–314.
  • SCHINDLER, E. 1990. The late Frasnian (Upper Devonian) Kellwasser Crisis. 151–159. In KAUFFMAN, E. G. and WALLISER, O. H. (eds). Extinction events in Earth history. Lecture Notes in Earth History, 30. Springer-Verlag, Heidelberg.
  • SEDDON, G. 1970. Frasnian conodonts from the Sadler Ridge-Bugle Gap area, Canning Basin, Western Australia. Journal of the Geological Society of Australia, 16, 723–753.
  • TEICHERT, C. 1943. The Devonian of Western Australia, a preliminary review. American Journal of Science, 241, 69–184.
  • TEICHERT, C. 1949. Stratigraphy and palaeontology of Devonian portion of the Kimberley Division, Western Australia. Australian Bureau of Mineral Resources, 2, 1–55.
  • VANĚK, J. 1963. Nové rody čeledi Harpidae Hawle et Corda, 1847 (Trilobita) z Barrandienu. Časopis. Národního Muzea. Oddíl Přírodovědný, 132, 226–229.
  • WALOSSEK, D. 1993. The Upper Cambrian Rehbachiella and the phylogeny of Branchiopoda and Crustacea. Fossils and Strata, 32, 1–202.
  • WEBER, V. N. 1932. Trilobity Turkestana. Trudy geologicheskogo komiteta, Moscow, Leningrad, 178, 1–157 [in Russian with English summary].
  • WEBSTER, M., SHEETS, H. D. and HUGHES, N. C. 2001. Allometric patterning in trilobite ontogeny: testing for heterochrony in Nephrolenellus. 105–144. In ZELDITCH, M. L. (ed.). Beyond heterochrony. Wiley-Liss, New York.
  • WEYER, D., FEIST, R. and GIRARD, C. 2003. Conodonta, Trilobita, and Anthozoa near the Late Frasnian Upper Kellwasser Event of the Geipel Quarry section in Schleiz, Thuringian Mountains (Germany). Mitteilungen des Museums für Naturkunde, Geowissenschaftliche Reihe, 6, 71–78.
  • WHITTINGTON, H. B. 1950. British trilobites of the family Harpidae. Palaeontographical Society Monograph. 55 pp.
  • WHITTINGTON, H. B. 1956. Silicified Middle Ordovician trilobites: the Odontopleuridae. Bulletin of the Museum of Comparative Zoology, Harvard, 14, 155–288.
  • ZIEGLER, W. and SANDBERG, C. A. 1990. The Late Devonian standard conodont zonation. Courier Forschungsinstitut Senckenbergia, 121, 1–115.
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