Hostname: page-component-586b7cd67f-l7hp2 Total loading time: 0 Render date: 2024-11-22T05:56:42.834Z Has data issue: false hasContentIssue false

Cladistic analysis of the Early Cambrian olenelloid trilobites

Published online by Cambridge University Press:  20 May 2016

Bruce S. Lieberman*
Affiliation:
Department of Organismic and Evolutionary Biology, 26 Oxford Street, Harvard University, Cambridge, MA 02138

Abstract

A phylogenetic analysis was used to determine evolutionary relationships within the Early Cambrian superfamily Olenelloidea Walcott, 1890. Phylogenetic patterns within the suborder Olenellina Walcott, 1890, which contains the Olenelloidea and the Fallotaspidoidea Hupé, 1953, are also discussed. The Olenelloidea are monophyletic, and synapomorphies uniting them include the condition of the ocular lobes where they intersect the frontal lobe of the glabella, and the condition of the lateral margins of the glabellar lobes. In contrast, taxa formerly assigned to the Fallotaspidoidea are shown to represent a paraphyletic grade of several genera, some more closely related to the Olenelloidea, and some more closely related to the Redlichiina Richter, 1933. Seventy-nine exoskeletal characters were coded for 26 taxa within the Olenellina. These included 22 ingroup Olenelloidea and four outgroup taxa that have traditionally been assigned to the Fallotaspidoidea. When subjected to parsimony analysis these character data yielded a single most parsimonious cladogram that provides an hypothesis of relationship for the generic clades within the superfamily. Two new genera are recognized herein, Fritzolenellus and Lochmanolenellus. It has been argued that genetic flexibility was so great and trilobite morphology was so plastic in the Early Cambrian that suprageneric classification of Early Cambrian trilobites is precluded. Although levels of intraspecific variability may have been slightly higher in the Early Cambrian relative to the mid Paleozoic, based on the extent of polymorphic character codings, it was not so high as to obviate attempts at recovering phylogenetic structure in a major clade of Early Cambrian taxa. In addition, the consistency index recovered by this analysis is not unduly low for a phylogenetic database of this size. The phylogenetic analysis also has bearing on patterns of allometric heterochrony, which have often been held to be significant in Early Cambrian trilobites. The paedomorphic retention of advanced genal spines into the adult probably evolved at least four times. Three of the episodes can be best described as neoteny, the fourth, as progenesis. Finally, based on the phylogeny, it is likely that rates of speciation in trilobites may have been two to three times higher in the Early Cambrian than in the mid Paleozoic.

Type
Research Article
Copyright
Copyright © The Paleontological Society 

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Ahlberg, P. 1991. Trilobites in the Lower Cambrian of Scandinavia. Geologiska Föreningens i Stockholm Forhandlingar, 113:7475.Google Scholar
Ahlberg, P., Bergström, J., and Johansson, J. 1986. Lower Cambrian olenellid trilobites from the Baltic faunal province. Geologiska Föreningens i Stockholm Forhandlingar, 108:3956.Google Scholar
Archie, J. W. 1989. Homoplasy excess ratios: new indices for measuring levels of homoplasy in phylogenetic systematics and a critique of the consistency index. Systematic Zoology, 38:253269.CrossRefGoogle Scholar
Barnaby, R. J., and Read, J. F. 1990. Carbonate ramp to rimmed shelf evolution: Lower to Middle Cambrian continental margin, Virginia Appalachians. Geological Society of America Bulletin, 102:391404.Google Scholar
Bergström, J. 1973. Classification of olenellid trilobites and some Balto-Scandian species. Norsk Geologisk Tidsskrift, 53:283314.Google Scholar
Bowring, S. A., Grotzinger, J. P., Isachsen, C. E., Knoll, A. H., Pelechaty, S. M., and Kolosov, P. 1993. Calibrating rates of early Cambrian evolution. Science, 261:12931298.Google Scholar
Bremer, K. 1994. Branch support and tree stability. Cladistics 10, 295304.CrossRefGoogle Scholar
Briggs, D. E. G., Fortey, R. A., and Wills, M. A. 1992. Morphological disparity in the Cambrian. Science, 256:16701673.Google Scholar
Budd, G. E. 1995. Kleptothule rasmusseni gen. et sp. nov.: an ?ole-nellinid-like trilobite from the Sirius Passet fauna (Buen Formation, Lower Cambrian, north Greenland). Transactions of the Royal Society of Edinburgh, Earth Sciences, 86:112.Google Scholar
Campbell, L., and Kauffman, M. E. 1969. Olenellus fauna of the Kinzers Formation, southeastern Pennsylvania. Proceedings of the Pennsylvania Academy of Science, 43 Cowie, J. W. 1968. Lower Cambrian faunas from Ellesmere Island, district of Franklin. Geological Survey of Canada Bulletin, 163:1127.Google Scholar
Edgecombe, G. D. 1992. Trilobite phylogeny and the Cambrian/Ordovician “event”: cladistic reappraisal, pp. 144177. In Novacek, M. J. and Wheeler, Q. D. (eds.), Extinction and Phylogeny. Columbia University Press, New York.Google Scholar
Eldredge, N., and Cracraft, J. 1980. Phylogenetic Patterns and the Evolutionary Process: Method and Theory in Comparative Biology. Columbia University Press, New York, 349 p.Google Scholar
Emmons, E. 1844. The Taconic System; based on observations in New York, Massachusetts, Maine, Vermont and Rhode Island. Albany, New York, 653 p.Google Scholar
Emmons, E. 1860. Manual of Geology. New York.Google Scholar
Fortey, R. A., and Whittington, H. B. 1989. The Trilobita as a natural group. Historical Biology, 2:125138.Google Scholar
Fortey, R. A., and Owens, R. M. 1990. Evolutionary radiations in the Trilobita, pp. 139164. In Taylor, P. D. and Larwood, G. P. (eds.), Major Evolutionary Radiations. Systematics Association Special Volume No. 42. Clarendon Press, Oxford.Google Scholar
Fritz, W. H. 1972. Lower Cambrian trilobites from the Sekwi Formation type section, Mackenzie Mountains, northwestern Canada. Geological Survey of Canada Bulletin, 212:190.Google Scholar
Fritz, W. H. 1973. Medial Lower Cambrian trilobites from the Mackenzie Mountains, northwestern Canada. Geological Survey of Canada, Paper 73-24, 43 p.Google Scholar
Fritz, W. H. 1991. Lower Cambrian trilobites from the Illtyd Formation, Wernecke Mountains, Yukon Territory. Geological Survey of Canada Bulletin, 409:177.Google Scholar
Fritz, W. H. 1992. Walcott's Lower Cambrian olenellid trilobite collection 61K, Mount Robson area, Canadian Rocky Mountains. Geological Survey of Canada Bulletin, 432:165.Google Scholar
Fritz, W. H. 1993. New Lower Cambrian olenelloid trilobite genera Cirquella and Geraldinella from southwestern Canada. Journal of Paleontology, 67:856868.Google Scholar
Fritz, W. H. 1995. Esmeraldina rowei and associated Lower Cambrian trilobites (1f fauna) at the base of Walcott's Waucoban series, southern Great Basin, U.S.A. Journal of Paleontology, 69:708723.CrossRefGoogle Scholar
Geyer, G. 1993. The giant Cambrian trilobites of Morocco. Beringeria, 8:71107.Google Scholar
Geyer, G., and Palmer, A. R. 1995. Neltneriidae and Holmiidae (Trilobita) from Morocco and the problem of Early Cambrian intercontinental correlation. Journal of Paleontology, 69:459474.Google Scholar
Gould, S. J. 1989. Wonderful Life. W. W. Norton, New York, 347 p.Google Scholar
Gould, S. J. 1991. The disparity of the Burgess Shale arthropod fauna and the limits of cladistic analysis: why we must strive to quantify morphospace. Paleobiology, 17:411423.Google Scholar
Hall, J. 1859. Trilobites of the shales of the Hudson River Group, p. 5962. 12th Annual Report of the New York Cabinet for Natural History, Albany, New York.Google Scholar
Hall, J. 1862. Supplementary note to the thirteenth report of the Regents of the State Cabinet, p. 113119. 15th Annual Report of the New York Cabinet for Natural History, Albany, New York.Google Scholar
Harrington, H. J. 1956. Olenellidae with advanced cephalic spines. Journal of Paleontology, 30:5661.Google Scholar
Harrington, H. J., Henningsmoen, G., Howell, B. F., Jaanusson, V., Lochman-Balk, C., Moore, R. C., Poulsen, C., Rasetti, F., Richter, E., Richter, R., Schmidt, H., Sdzuy, K., Struve, W., Stormer, L., Stubblefield, C. J., Tripp, R., Weller, J. M., and Whittington, H. B. 1959. In Moore, R. C. (ed.), Treatise on Invertebrate Paleontology, Part O, Arthropoda 1. Geological Society of America and University of Kansas Press, Lawrence.Google Scholar
Hu, C.-H. 1971. Ontogeny and sexual dimorphism of lower Paleozoic Trilobita. Palaeontographica Americana, 7:31155.Google Scholar
Hughes, N. C. 1991. Morphological plasticity and genetic flexibility in a Cambrian trilobite. Geology, 19:913916.Google Scholar
Hupé, P. 1952. Sur les zones de trilobites du Cambrien inferieur marocain. Comptes Rendus de l'Academie des Sciences, Paris, 235:480481.Google Scholar
Hupé, P. 1953. Contribution a l'etude du Cambrien inferieur et du Precambrien III de l'Anti-Atlas marocain. Notes et Mémoires du Service Géologique (Morocco), no. 103, 402 p.Google Scholar
Hupé, P., and Abadie, J. 1950. Sur l'existence de Trilobites du Cambrien Inferieur marocain. Comptes Rendus de l'Academie de Sciences, 230:21122113.Google Scholar
Khomentovskii, V. V., and Repina, L. N. 1965. (The Lower Cambrian stratotype section of Siberia). Sibirskoe Otdelenie, Institut Geologii i Geofiziki, Akademiia Nauk SSSR.Google Scholar
Kiaer, J. 1917. The Lower Cambrian Holmia fauna at Tomten in Norway. Vedenskapsselskapets Skrifter, I. Matematisk naturviden skapelig Klasse, 10, 140 p.Google Scholar
Kir'ianov, V. V., and Chernysheva, N. E. 1967. (About Lower Cambrian deposits of northwestern Volini and the discovery of an ancient trilobite). Izvestiia Akademii Nauk SSSR, Seriia Geologicheskaia, 1967(7):119125.Google Scholar
Klassen, G. J., Mooi, R. D., and Locke, A. 1991. Consistency indices and random data. Systematic Zoology, 40:446457.Google Scholar
Knoll, A. H. 1996. Daughter of time. Paleobiology, 22:17.Google Scholar
Landing, E. 1994. Precambrian Cambrian boundary global stratotype ratified and a new perspective of Cambrian time. Geology, 22:179182.2.3.CO;2>CrossRefGoogle Scholar
Lauterbach, K.-E. 1983. Synapomorphen zwischen Trilobiten- und Cheliceratenzweig der Arachnata. Zoologischer Anzeiger, 210:213238.Google Scholar
Lazarenko, N. P. 1960. (New species of ancient plants and invertebrates of the USSR, Pt. 2). In Markoviskiy, B. P. (ed.), VSEGEI, Moscow, 552 p.Google Scholar
Lazarenko, N. P. 1962. (New Lower Cambrian trilobites from the Soviet Arctic). Sbornik Statey po Paleontologii i Biostratigrafii, 29:2977.Google Scholar
Lermontova, E. V. 1951. (Lower Cambrian trilobites and brachiopods from eastern Siberia). Gosgeolizdat. Moscow, 218 p.Google Scholar
Lieberman, B. S. 1994. Evolution of the trilobite subfamily Proetinae Salter, 1864 and the origin, diversification, evolutionary affinity and extinction of the Middle Devonian proetid fauna of Eastern North America. Bulletin of the American Museum of Natural History, 223:1176.Google Scholar
Linnarsson, J. G. O. 1871. Om några forsteningar från Sveriges och Norges “Primordialzon.” Ofversigt af Kongliga Svenska Vetenskaps Akademiens Forhandlingar, 6:789796.Google Scholar
Lochman, C. 1952. Trilobites. In Cooper, G. A., Arellano, A. R. V., Johnson, J. H., Okulitch, V., Stoyanow, A., and Lochman, C. (eds.), Cambrian stratigraphy and paleontology near Caborca, northwestern Sonora, Mexico. Smithsonian Miscellaneous Collections, 119:60184.Google Scholar
Ludvigsen, R., and Westrop, S. R. 1985. Three new Upper Cambrian stages for North America. Geology, 13:139143.2.0.CO;2>CrossRefGoogle Scholar
Maddison, W. P., and Maddison, D. R. 1992. MacClade: Analysis of phylogeny and character evolution. Version 3.04. Sinauer Associates, Sunderland, MA.Google Scholar
Matthew, G. F. 1890. On Cambrian organisms in Acadia. Royal Society of Canada Proceedings and Transactions, 7:135162.Google Scholar
Matthew, G. F. 1897. What is the Olenellus fauna? American Geologist, 19:396407.Google Scholar
Mcnamara, K. J. 1978. Paedomorphosis in Scottish olenellid trilobites (Early Cambrian). Palaeontology, 21:635655.Google Scholar
Mcnamara, K. J. 1986. The role of heterochrony in the evolution of Cambrian trilobites. Biological Reviews, 61:121156.Google Scholar
Moberg, J. C., and Segerberg, C. O. 1906. Bidrag till Kannedomen om Ceratopyge regionen med sarskild hansyn till des utveckling i Fogelsangtraken. Meddelande fran Lunds Geologiska Faltkubb (series B), 2:1113.Google Scholar
Morris, P. J., Ivany, L. C., Schopf, K. M., and Brett, C. E. 1995. The challenge of paleoecological stasis: reassessing sources of evolutionary stability. Proceedings of the National Academy of Sciences, USA, 92:1126911273.Google Scholar
Nelson, C. A. 1976. Late Precambrian Early Cambrian stratigraphic and faunal succession of eastern California and the Precambrian Cambrian boundary, p. 3141. In Moore, J. N. and Fritsche, A. E. (eds.), Depositional environments of Lower Paleozoic rocks in the White Inyo Mountains, Inyo County, California. Society of Economic Paleontologists and Mineralogists, Pacific Section, Los Angeles.Google Scholar
Neltner, L., and Poctey, N. 1949. Quelques faunes georgiennes du Maroc. Notes du Service Géologique Maroc, 2, no. 74:5383.Google Scholar
Pack, P. D., and Gayle, H. D. 1971. A new olenellid trilobite, Biceratops nevadensis, from the Lower Cambrian near Las Vegas, Nevada. Journal of Paleontology, 45:893898.Google Scholar
Palmer, A. R., and Halley, R. B. 1979. Physical stratigraphy and trilobite biostratigraphy of the Carrara Formation (Lower and Middle Cambrian) in the southern Great Basin. U. S. Geological Survey, Professional Paper, 1047:1131.Google Scholar
Palmer, A. R., and Halley, R. B., and Peel, J. S. 1979. New Cambrian faunas from Peary land, eastern North Greenland. Gronlands Geologiske Unders⊘gelse Rapport, 91:2936.Google Scholar
Palmer, A. R., and Halley, R. B., and Repina, L. N. 1993. Through a glass darkly: taxonomy, phylogeny and biostratigraphy of the Olenellina. University of Kansas Paleontological Contributions, New Series, 3, 35 p.Google Scholar
Peach, B. N. 1894. Additions to the fauna of the Olenellus zone of the north-west highlands. Quarterly Journal of the Geological Society of London, 50:661675.Google Scholar
Peach, B. N., and Horne, J. 1892. The Olenellus zone in the northwest highlands of Scotland. Quarterly Journal of the Geological Society of London, 48:227242.Google Scholar
Poulsen, V. 1974. Olenellacean trilobites from eastern North Greenland. Bulletin of the Geological Society of Denmark, 23:79101.Google Scholar
Ramsköld, L.,and Edgecombe, G. D. 1991. Trilobite monophyly revisited. Historical Biology, 4:267283.Google Scholar
Raw, F. 1936. Mesonacidae of Comley in Shropshire, with a discussion of classification within the family. Quarterly Journal of the Geological Society of London, 92:236293.Google Scholar
Repina, L. N. 1961. (On the discovery of olenellids in the Lena Stage of Batenevskiy Ridge). Doklady Akademii Nauk SSSR, 136:4045.Google Scholar
Repina, L. N. 1979. (Dependence of morphologic features on habitat conditions in trilobites and evaluation of their significance for the systematics of the superfamily Olenelloidea). Akademiia Nauk SSSR, Sibirskoe Otdelenie, Trudy Instituta Geologii i Geofizikii, 431:1130.Google Scholar
Repina, L. N. 1990. (Evolution of trilobites in the beginning stages of their historical development, p. 3444). In Betekina, O. A. and Zhuravleva, I. T. (eds.), (Environment and life of the geologic past). Akademiia Nauk SSSR, Sibirskoe Otdelenie, Trudy Instituta Geologii i Geofizikii 764.Google Scholar
Resser, C. E. 1928. Cambrian fossils from the Mohave Desert. Smithsonian Miscellaneous Collections, 81, no. 2, 14 p.Google Scholar
Resser, C. E. 1938. Cambrian system (restricted) of the southern Appalachians. Geological Society of America Special Papers, No. 15, 140 p.Google Scholar
Resser, C. E., and Howell, B. F. 1938. Lower Cambrian Olenellus zone of the Appalachians. Geological Society of America Bulletin, 49:195248.Google Scholar
Riccio, J. F. 1952. The Lower Cambrian Olenellidae of the southern Marble Mountains, California. Bulletin of the Southern California Academy of Sciences, 51:2549.Google Scholar
Richter, R. 1933. Crustacea (Palaontologie): Handworterbuch der Naturwissenschaften, 2:840864.Google Scholar
Sanderson, M. J., and Donoghue, M. J. 1989. Patterns of variation in levels of homoplasy. Evolution, 43:17811795.Google Scholar
Savitskii, V. E., Evtushenko, V. M., Egorova, L. I., Kontorovich, A. E., and Shabanov, I. I. 1972. Cambrian of the Siberian platform (Iudomo Oleneksk type section. Kuonamsk complex of sediments). Trudy sibirskoe nauchno issled Institut Geologii i Geofizik mineralogii Syr', 130:1200.Google Scholar
Sdzuy, K. 1961. Das Kambrium Spaniens. Teil II: Trilobiten. 1. Abschnitt. Akademie der Wissenschaften und der Literatur Mainz, Abhandlungen der mathematisch naturwissenschaftlichen Klasse, 1961:217312.Google Scholar
Sdzuy, K. 1978. The Precambrian-Cambrian boundary beds in Morocco (preliminary report). Geological Magazine, 115:8394.Google Scholar
Shaw, A. B. 1955. Paleontology of northwestern Vermont. V. The Lower Cambrian fauna. Journal of Paleontology, 29:775805.Google Scholar
Stitt, J. H.,and Clark, R. L. 1984. A complete specimen of Peachella brevispina Palmer an unusual olenellid trilobite (Arthropoda: Olenellida) from the lower Cambrian of California. Transactions of the San Diego Society of Natural History, 20:145150.Google Scholar
Suvorova, N. P. 1960. Trilobity kembriia vostoka Sibirskoi Platformy. Vypusk 2, Olenellidy Granuliariidy. Trudy Paleontologicheskogo Akademii Nauk SSSR, 84:1230.Google Scholar
Swofford, D. L. 1993. PAUP (Phylogenetic analysis using parsimony) version 3.1.1.Google Scholar
Waggoner, B. M. 1996. Phylogenetic hypotheses of the relationships of arthropods to Precambrian and Cambrian problematic fossil taxa. Systematic Biology, 45:190222.Google Scholar
Walcott, C. D. 1884. Paleontology of the Eureka district (Nevada). U. S. Geological Survey Monograph, 8:1298.Google Scholar
Walcott, C. D. 1885. Paleozoic notes: new genus of Cambrian trilobites, Mesonacis. America. Journal of Science (series 3), 29:328330.Google Scholar
Walcott, C. D. 1890. The fauna of the Lower Cambrian or Olenellus zone. U. S. Geological Survey, 10th Annual Report:509763.Google Scholar
Walcott, C. D. 1910. Olenellus and other genera of the Mesonacidae. Smithsonian Miscellaneous Collections, 53(6):231422.Google Scholar
Walcott, C. D. 1913. Cambrian geology and paleontology, No. 11. New Lower Cambrian subfauna. Smithsonian Miscellaneous Collections, 57(11):309326.Google Scholar
Wanner, A. 1901. A new species of Olenellus from the Lower Cambrian of York County, Pennsylvania. Proceedings of the Washington Academy of Sciences, 3:267272.Google Scholar
Westrop, S. R.,and Ludvigsen, R. 1987. Biogeographic control of trilobite mass extinction at an Upper Cambrian “biomere” boundary. Paleobiology, 13:8499.Google Scholar
Whitfield, R. P. 1884. Notice of some new species of primordial fossils in the collections of the Museum, and corrections of previously described species. Bulletin of the American Museum of Natural History, 1:139154.Google Scholar
Whittington, H. B. 1989. Olenelloid trilobites: type species, functional morphology and higher classification. Philosophical Transactions of the Royal Society of London B, 324:111147.Google Scholar
Wiley, E. O. 1979. An annotated Linnaean hierarchy, with comments on natural taxa and competing systems. Systematic Zoology, 28:308337.Google Scholar