Hostname: page-component-78c5997874-ndw9j Total loading time: 0 Render date: 2024-11-19T08:42:23.375Z Has data issue: false hasContentIssue false

Subspeciation within a punctuated equilibrium evolutionary event: Phylogenetic history of the Lower-Middle Ordovician Paroistodus originalis—P. horridus complex (Conodonta)

Published online by Cambridge University Press:  14 July 2015

Guillermo L. Albanesi
Affiliation:
CONICET, CRILAR, Mendoza y Entre Ríos, 5301 Anillaco, La Rioja, Argentina,
Christopher R. Barnes
Affiliation:
School of Earth and Ocean Sciences, University of Victoria, P.O. Box 3055, Victoria, British Columbia, V8W 3P6, Canada,

Abstract

A microevolutionary event involving the conodont Paroistodus lineage is documented in the Gualcamayo Formation (Middle Ordovician), Argentine Precordillera. A detailed sampling of limestones throughout the upper part of the San Juan Formation and the lower member of the Gualcamayo Formation yielded over 14,000 well-preserved conodont elements. Paroistodus originalis (Sergeeva, 1963) was recorded through the upper 230 m of the San Juan Formation and the lower member (10 m thick) of the Gualcamayo Formation. The derived species Paroistodus horridus (Barnes and Poplawski, 1973) was recorded throughout the middle member of the Gualcamayo Formation (65 m thick). The intermediate linking forms between both species are identified as two new taxa: Paroistodus horridus primus Albanesi, 1998b, and P. h. secundus Albanesi, 1998b. They were recorded in the uppermost 70 cm of the lower member. Apparently, the speciation event occurred under stressed environmental conditions with the drowning of the carbonate platform, i.e., the San Juan Formation, and the beginning of a deeper and restricted environment represented by the Gualcamayo black shales. The demise of the carbonate production was caused by a sea level rise and a significant influx of volcanic ashes. The punctuated speciation event occurred within an allopatric setting while the Precordillera occupied an isolated (Iapetus) oceanic position in its overall drift from Laurentia to Gondwana.

Type
Research Article
Copyright
Copyright © The Paleontological Society 2000

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

Aceñolaza, F. G., and Toselli, A. J. 1973. Consideraciones estratigráficas y tectónicas sobre el Paleozoico Inferior del noroeste argentino. II Congreso Latinoamericano de Geología, Caracas, Memorias, 2:755763.Google Scholar
Albanesi, G. L. 1998a. Biofacies de conodontes de las secuencias ordovícicas del cerro Potrerillo, Precordillera Central de San Juan, R. Argentina. Actas XII Academia Nacional de Ciencias., Córdoba, 7598.Google Scholar
Albanesi, G. L. 1998b. Taxonomía de conodontes de las secuencias ordovícicas del cerro Potrerillo, Precordillera Central de San Juan, R. Argentina. Actas XII Academia Nacional de Ciencias., Córdoba, 101253.Google Scholar
Albanesi, G. L., and Astini, R. A. 1994. Conodontofauna de los niveles cuspidales de la Formación San Juan (Llanvirniano) en el perfil de Las Chacritas, Provincia de San Juan. VI Congreso Argentino de Paleontología y Bioestratigrafía, Trelew, Chubut, Resúmenes, 48.Google Scholar
Albanesi, G. L., and Barnes, C. R. 1996. The origin of the Middle Ordovician conodont Paroistodus horridus in the Argentine Precordillera. Sixth European Conodont Symposium, ECOS VI, Warszawa, Abstracts, 1.Google Scholar
Albanesi, G. L., Hünicken, M. A., and Barnes, C. R. 1998a. Bioestratigrafía de conodontes de las secuencias ordovícicas del cerro Potrerillo, Precordillera Central de San Juan, R. Argentina. Actas XII Academia Nacional de Ciencias., Córdoba, 772.Google Scholar
Albanesi, G. L., Ortega, G., and Hünicken, M. A. 1995a. Conodontes y graptolitos de la Formación Yerba Loca (Arenigiano-Llandeiliano) en las quebradas de Ancaucha y El Divisadero, Precordillera de San Juan, Argentina. Boletín de la Academia Nacional de Ciencias, Córdoba, Argentina, 60(3-4):365400.Google Scholar
Albanesi, G. L., Ortega, G., and Hünicken, M. A. 1998b. High resolution conodont-graptolite biostratigraphy controlling the diachronous boundary between the San Juan and Gualcamayo formations (Arenig-Llanvirn) in the Argentine Precordillera. ECOS VII (Seventh International Conodont Symposium Held in Europe), Bologna-Modena, Italia. Abstracts:24.Google Scholar
An, T.-X, and Zheng, Z. 1990. The conodonts of the marginal areas around the Ordos Basin, north China. Science Press, 1101.Google Scholar
An, T.-X, Du, G., and Gao, Q. 1985. Ordovician conodonts from Hubei, China. Geological Publications House, Beijing, 164.Google Scholar
Astini, R. A. 1994a. Geología e interpretación de la Formación Gualcamayo en su localidad clásica (suroeste de Guandacol y cordón de Perico-Potrerillo), Precordillera septentrional. Revista de la Asociación Geológica Argentina, 49:5570.Google Scholar
Astini, R. A. 1994b. Análisis secuencial y paleoambientes de las pelitas negras (aloformación Gualcamayo) que suprayacen a las sucesiones carbonáticas eo-ordovícicas en la Precordillera argentina. Revista de la Asociación Geológica Argentina, 49:7184.Google Scholar
Astini, R. A. 1995. Paleoclimates and paleogeographic paths of the Argentine Precordillera during the Ordovician: evidence from climatically sensitive lithofacies, p. 177180. In Cooper, J. D., Droser, M. L., and Finney, S. C. (eds.), Ordovician Odyssey. Short Papers for the Seventh International Symposium on the Ordovician System, Las Vegas, SEPM, Fullerton, CA.Google Scholar
Astini, R. A., Benedetto, J. L., and Vaccari, N. E. 1995. The early Paleozoic evolution of the Argentine Precordillera as a Laurentian rifted, drifted, and collided terrane: A Geodynamic model. Geological Society of America Bulletin, 107:253273.2.3.CO;2>CrossRefGoogle Scholar
Astini, R. A., Ramos, V. A., Benedetto, J. L., Vaccari, N. E., and Cañas, F. L. 1996. La Precordillera: un terreno exótico a Gondwana. XIII Congreso Geológico Argentino y III Congreso Exploración de Hidrocarburos, Buenos Aires, Actas, V:293324.Google Scholar
Baldis, B. A. J. 1979. Ensayo de análisis paleoecológicos con trilobites ordovícicos argentinos. Ameghiniana, 15:314.Google Scholar
Baldis, B. A. J., and Beresi, M. 1981. Biofacies de culminación del ciclo deposicional calcáreo del Arenigiano del oeste de Argentina. Anais II Congresso Latinoamericano de Paleontologia, Porto Alegre, 1:1119.Google Scholar
Baldis, B. A. J., and González, S. B. 1987. Kainisiliellinae, nueva subfamilia de Asaphidae (Trilobita) del Ordovícico de la Precordillera (sierra de Villicum, provincia de San Juan, Argentina). IV Congreso Latinoamericano de Paleontología, Sta. Cruz de la Sierra, Memorias, 1:5973.Google Scholar
Baldis, B. A. J., Beresi, M., Bordonaro, O., and Vaca, A. 1982. Síntesis evolutiva de la Precordillera Argentina. V Congreso Latinoamericano de Geología, Buenos Aires, Actas, IV:399445.Google Scholar
Baldis, B. A. J., Beresi, M., Armella, C., Beresi, M., Cabaleri, N., Peralta, S., and Bastías, H. 1989. La cuenca paleozoica inferior de la Precordillera Argentina, p. 101121. In Chebli, G. A. and Spalletti, L. A. (eds.), Cuencas sedimentarias argentinas. Serie Correlación Geológica No. 6.Google Scholar
Barnes, C. R. 1984. Early Ordovician eustatic events in Canada, p. 5163. In Bruton, D. L. (ed.), Aspects of the Ordovician System. Palaeontological Contributions from the University of Oslo, 295, Universitetsforlaget, Oslo.Google Scholar
Barnes, C. R. 1988. The proposed Cambrian-Ordovician global Boundary stratotype and point (GSSP) in Western Newfoundland, Canada. Geological Magazine, 125:381414.CrossRefGoogle Scholar
Barnes, C. R., and Fåhraeus, L. E. 1975. Provinces, communities, and the proposed nektobenthic habit of Ordovician conodontophorids. Lethaia, 8:133149.CrossRefGoogle Scholar
Barnes, C. R., and Poplawski, M. L. S. 1973. Lower and Middle Ordovician conodonts from the Mystic Formation, Quebec, Canada. Journal of Paleontology, 47:760790.Google Scholar
Benedetto, J. L., Cañas, F., and Astini, R. 1986. Braquiópodos y trilobites de la zona de transición entre las Formaciones San Juan y Gualcamayo en el área de Guandacol (La Rioja, Argentina). IV Congreso Argentino de Paleontología y Bioestratigrafía, Mendoza, Actas, 1:103111.Google Scholar
Benedetto, J. L., Herrera, Z. A., and Ortega, G. 1991. Bioestratigrafía del Ordovícico en el flanco occidental del Cerro Potrerillo, Provincia de San Juan. Revista de la Asociación Geológica Argentina, 46:6068.Google Scholar
Benedetto, J. L., Herrera, Z. A., Carrera, M., and Sánchez, T. M. 1995. The evolution of faunal provincialism in the Argentine Precordillera during the Ordovician: new evidence and paleogeographic implications, p. 181184. In Cooper, J. D., Droser, M. L. and Finney, S. C. (eds.), Ordovician Odyssey. Short Papers for the Seventh International Symposium on the Ordovician System, Las Vegas, SEPM, Fullerton, California.Google Scholar
Bergström, S. M. 1979. Whiterockian (Ordovician) conodonts from the H⊘londa Limestone of the Trondheim Region, Norwegian Caledonides. Norsk Geologisk Tidsskrift, 59:295307.Google Scholar
Bergström, S. M., Huff, W. D., Kolata, D. R., Krekeler, M. P. S., Cingolani, C., and Astini, R. A. 1996. Lower and Middle Ordovician K-bentonites in the Precordillera of Argentina: a progress report. XIII Congreso Geológico Argentino y III Congreso de Exploración de Hidrocarburos, Buenos Aires, Actas V:481490.Google Scholar
Boucot, A. J. 1992. Evolution of communities, p. 391394. In Briggs, D. E. G. and Crowther, P. R. (eds.), Paleobiology. A Synthesis. Blackwell Scientific Publications, Oxford.Google Scholar
Bradshaw, L. E. 1969. Conodonts from the Fort Peña Formation (Middle Ordovician), Marathon Basin, Texas. Journal of Paleontology, 43:11371168.Google Scholar
Broadhead, T. W., Driese, S. G., and Harvey, J. L. 1990. Gravitational settling of conodont elements: Implications for paleoecologic interpretations of conodont assemblages. Geology, 18:850853.2.3.CO;2>CrossRefGoogle Scholar
Cañas, F. L. 1995a. Estratigrafía y evolución paleoambiental de las sucesiones carbonáticas del Cámbrico tardío y Ordovícico temprano de la Precordillera Septentrional, República Argentina. Tesis Doctoral en Ciencias Geológicas, Facultad de Ciencias Exactas, Físicas y Naturales, Universidad Nacional de Córdoba, 1216.Google Scholar
Cañas, F. L. 1995b. Early Ordovician carbonate platform facies of the Argentine Precordillera: restricted shelf to open platform evolution, p. 221224. In Cooper, J. D., Droser, M. L. and Finney, S. C. (eds.), Ordovician Odyssey. Short Papers for the Seventh International Symposium on the Ordovician System, Las Vegas, SEPM, Fullerton, California.Google Scholar
Carrera, M. G. 1997. Significado paleoambiental de los poríferos y briozoos de la Formación San Juan (Ordovícico), Precordillera Argentina. Ameghiniana, 34:179199.Google Scholar
Carrera, M. G., and Astini, R. A. 1998. Valoración de las restricciones ambientales durante la transición Arenigiano-Llanvirniano, Ordovícico de la Precordillera argentina. Ameghiniana, 53(1):4156.Google Scholar
Cas, R. A. F., and Vandemberg, A. H. M. 1988. Ordovician. In Geology of Victoria, Revised Edition, Douglas, J. G. and Ferguson, J. A. (eds.); Victorian Division of the Geological Society of Australia, 63102.Google Scholar
Charlesworth, B. 1992. Speciation, p. 100106. In Briggs, B. E. G. and Crowther, P. R. (eds.), Paleobiology: A Synthesis. Blackwell Scientific Publications, Oxford.Google Scholar
Chauffe, K. M., and Nichols, P. A. 1995. Differentiating evolution from environmentally induced modifications in Mid-Carboniferous conodonts. Palaeontology, 38:875895.Google Scholar
Cheetham, A. H. 1987. Tempo of evolution in a Neogene bryozoan: are trends in single morphologic characters misleading? Paleobiology, 13:286296.CrossRefGoogle Scholar
Cocks, L. R. M., Mckerrow, W. S., and van Staal, C. R. 1997. The margins of Avalonia. Geological Magazine, 134:627636.CrossRefGoogle Scholar
Dalziel, I.W.D., Dalla Salda, L. H., Cingolani, C., and Palmer, P. 1996. The Argentine Precordillera: A Laurentian Terrane? Penrose Conference Report, GSA Today, p. 1618.Google Scholar
Darwin, C. 1878. The origin of species by means of natural selection, or the preservation of favoured races in the struggle for life. J. Murray ed. (sixth edition), London, 458 p.Google Scholar
Dumoulin, J. A., and Harris, A. G. 1987. Lower Paleozoic carbonate rocks of the Baird Mountains quadrangle, western Brooks Range, Alaska, p. 311336. In Tailleur, I. L. and Weimer, P. (eds.), Alaskan North Slope geology. Society of Economic Paleontologists and Mineralogists, Pacific Section, 50, Bakersfield, California.Google Scholar
Dzik, J. 1983. Relationships between Ordovician Baltic and North American Midcontinent conodont faunas. Fossils and Strata, 15:5986.Google Scholar
Dzik, J. 1989. Conodont evolution in high latitudes of the Ordovician. Courier Forschungs-institut Senckenberg, 117:128.Google Scholar
Dzik, J. 1991. Evolution of oral apparatuses in the conodont chordates. Acta Palaeontologica Polonica, 36:265323.Google Scholar
Dzik, J. 1994. Conodonts of the Mójcza Limestone, p. 43128. In Dzik, J., Olempska, E., and Pisera, A. (eds.), Ordovician carbonate platform ecosystem of the Holy Cross Mountains. Palaeontologia Polonica, 53.Google Scholar
Dzik, J. 1999. Relationship between rates of speciation and phyletic evolution: stratophenetic data on pelagic conodont chordates and benthic ostracods. GEOBIOS, 32, 2:205221.CrossRefGoogle Scholar
Eldredge, N., and Gould, S. J. 1972. Punctuated equilibria; an alternative to phyletic gradualism, p. 82115. In Schopf, T. J. M. (ed.), Models in paleobiology. Freeman, San Francisco.Google Scholar
Epstein, A. G., Epstein, J. B., and Harris, L. D. 1977. Conodont color alteration—An index to organic metamorphism. U. S. Geological Survey Professional Paper, 995:127.Google Scholar
Erdtmann, B. D. 1986. Early Ordovician eustatic cycles and their bearing on punctuations in early nematophorid (planktic) graptolite evolution, p. 139152. In Walliser, O. H. (ed.), Global Bio-Events. Springer-Verlag, Berlin and Heidelberg.CrossRefGoogle Scholar
Fortey, R. A. 1984. Global earlier Ordovician transgressions and regressions and their biological implications, p. 3750. In Bruton, D. L. (ed.), Aspects of the Ordovician System. Palaeontological Contributions from the University of Oslo 295, Universitetsforlaget, Oslo.Google Scholar
Furque, G., and Cuerda, A. 1979. Precordillera de La Rioja, San Juan y Mendoza. Segundo Simposio de Geología Regional Argentina, Córdoba, Academia Nacional de Ciencias, 1:455522.Google Scholar
Gingerich, P. D. 1985. Species in the fossil record: concepts, trends, and transitions. Paleobiology, 11:2741.CrossRefGoogle Scholar
Gingerich, P. D. 1992. Stratophenetics, p. 437442. In Briggs, D. E. G. and Crowther, P. R. (eds.), Paleobiology: A Synthesis. Blackwell Scientific Publications, Oxford.Google Scholar
Gould, S. J. 1982. The meaning of punctuated equilibrium and its role in validating a hierarchical approach to macroevolution, p. 83104. In Milkman, R. (ed.), Perspectives on Evolution. Sinauer, Sunderland, Massachusetts.Google Scholar
Gould, S. J., and Eldredge, N. 1977. Punctuated equilibria: the tempo and mode of evolution reconsidered. Paleobiology, 3:115151.CrossRefGoogle Scholar
Hallam, A. 1965. Environmental causes of stunting in living and fossil marine benthonic invertebrates. Palaeontology, 8:132155.Google Scholar
Harris, A. G., Bergström, S. M., Ethington, R. L., and Ross, R. J. Jr. 1979. Aspects of Middle and Upper Ordovician conodont biostratigraphy of carbonate facies in Nevada and southeast California and comparison with some Appalachian successions, p. 743. In Sandberg, C. A. and Clark, D. L. (eds.), Conodont Biostratigraphy of the Great Basin and Rocky Mountains. Brigham Young University Geology Studies, 26.Google Scholar
Holland, S. M. 1995. The stratigraphic distribution of fossils. Paleobiology, 21:92109.CrossRefGoogle Scholar
Huff, W. D., Bergström, S. M., Kolata, D. R., Cingolani, C., and Davis, D. W. 1995. Middle Ordovician K-bentonites discovered in the Precordillera of Argentina: geochemical and paleogeographical implications, p. 343349. In Cooper, J. D., Droser, M. L. and Finney, S. C. (eds.), Ordovician Odyssey. Short Papers for the Seventh International Symposium on the Ordovician System, Las Vegas, SEPM, Fullerton, California.Google Scholar
Hünicken, M. A. 1985. Lower Ordovician conodont biostratigraphy in Argentina. Boletín de la Academia Nacional de Ciencias, Córdoba, Argentina, 56:309321.Google Scholar
Ji, Z., Barnes, And C. R. 1994. Lower Ordovician conodonts of the St. George Group, Port au Port Peninsula, western Newfoundland, Canada. Palaeontographica Canadiana, Geological Association of Canada and Canadian Society of Petroleum Geologists, Calgary, 11:1149.Google Scholar
Keller, M. 1995. Continental slope deposits in the Argentine Precordillera: sediments and geotectonic significance, p. 215215. In Cooper, J. D., Droser, M. L., and Finney, S. C. (eds.), Ordovician Odyssey. Short Papers for the Seventh International Symposium on the Ordovician System, Las Vegas, SEPM, Fullerton, California.Google Scholar
Kellogg, D. E. 1983. Phenology of morphologic change in radiolarian lineages from deep-sea cores: implications for macroevolution. Paleobiology, 9:355362.CrossRefGoogle Scholar
Krekeler, M. P. S., Huff, W. D., Kolata, D. R., Bergström, S. M., and Cingolani, C. 1995. Mineralogy and grain characteristics of Middle Ordovician K-bentonites from the Precordillera of Argentina, p. 355356. In Cooper, J. D., Droser, M. L. and Finney, S. C. (eds.), Ordovician Odyssey. Short Papers for the Seventh International Symposium on the Ordovician System, Las Vegas, SEPM, Fullerton, California.Google Scholar
Lande, R. 1986. The dynamics of peak shifts and the pattern of morphological evolution. Paleobiology, 12:343354.CrossRefGoogle Scholar
Landing, E. 1976. Early Ordovician (Arenigian) conodont and graptolite biostratigraphy of the Taconic allochton, eastern New York. Journal of Paleontology, 50:614646.Google Scholar
Lazarus, D. B., and Prothero, D. R. 1984. The role of stratigraphic and morphologic data in phylogeny. Journal of Paleontology, 58:163172.Google Scholar
Lehnert, O. 1993. Bioestratigrafía de los conodontes arenigianos de la Formación San Juan en la localidad de Niquivil (Precordillera sanjuanina, Argentina) y su correlación intercontinental. Revista Española de Paleontología, 8:153164.Google Scholar
Lehnert, O. 1995. Ordovizische Conodonten aus der Präkordillere Westargentiniens: Ihre Bedeutung für Stratigraphie und Paläogeographie. Erlanger Geologische Abhandlungen, 125, 193 p.Google Scholar
Lieberman, B. S., Brett, C. E., and Eldredge, N. 1995. A study of stasis and change in two species lineages from the Middle Devonian of New York state. Paleobiology, 21:1527.CrossRefGoogle Scholar
Lindström, M. 1964. Conodonts. Elsevier, Amsterdam, 196 p.Google Scholar
Lindström, M. 1971. Lower Ordovician conodonts of Europe, p. 2161. In Sweet, W. C. and Bergström, S. M. (eds.), Symposium on Conodont Biostratigraphy. Geological Society of America Memoir, 127.Google Scholar
Lindström, M. 1984. Baltoscandic conodont life environments in the Ordovician: Sedimentologic and paleogeographic evidence, p. 3342. In Clark, D. L. (ed.), Conodont Biofacies and Provincialism. Geological Society of America Special Paper, 196.Google Scholar
Löfgren, A. 1978. Arenigian and Llanvirnian conodonts from Jämtland, northern Sweden. Fossils and Strata, 13:1129.Google Scholar
Löfgren, A. 1994. Arenig (Lower Ordovician) conodonts and biozonation in the Eastern Siljan District, Central Sweden. Journal of Paleontology, 68:13501368.CrossRefGoogle Scholar
Löfgren, A. 1995. The probable origin of the Ordovician conodont “Cordylodushorridus . Geobios, 28:371377.CrossRefGoogle Scholar
Löfgren, A. 1996. Lower Ordovician conodonts, reworking, and biostratigraphy of the Orreholmen quarry, Västergötland, south-central Sweden. Geologiska Föreningens i Stockholm Förhandlingar (GFF), 118:169183.Google Scholar
Löfgren, A. 1997. Reinterpretation of the Lower Ordovician conodont apparatus Paroistodus . Paleontology, 40:913929.Google Scholar
Margalef, R. 1977. Ecología. Ediciones Omega, Barcelona, 951 p.Google Scholar
McCracken, A. D. 1991. Middle Ordovician conodonts from the Cordilleran Road River Group, northern Yukon Territory, Canada, p. 4163. In Orchard, M. J. and McCracken, A. D. (eds.), Ordovician to Triassic Conodont Paleontology of the Canadian Cordillera. Geological Survey of Canada Bulletin, 417: 41-63.Google Scholar
McGhee, G. R. Jr., Bayer, U., and Seilacher, A. 1991. Biological and evolutionary responses: to transgressive-regressive cycles, p. 696708. In Einsele, G., Ricken, W. and Seilacher, A. (eds.), Cycles and events in Stratigraphy. Springer-Verlag, Berlin and Heidelberg.Google Scholar
McGoff, H. J. 1991. The hydrodynamics of conodont elements. Lethaia, 24:235247.CrossRefGoogle Scholar
Miller, J. F. 1980. Taxonomic revisions of some Upper Cambrian and Lower Ordovician conodonts, with comments on their evolution. University of Kansas Paleontological Contributions Paper, 99:144.Google Scholar
Miller, W. III. 1986. Paleoecology of benthic community replacement. Lethaia, 19:225231.CrossRefGoogle Scholar
Nehm, R. H., and Geary, D. H. 1994. A gradual morphologic transition during a rapid speciation event in marginellid gastropods (Neogene: Dominican Republic). Journal of Paleontology, 68:787795.CrossRefGoogle Scholar
Nicoll, R. S. 1987. Form and function of the Pa element in the conodont animal, p. 7790. In Aldridge, R. J. (ed.), Paleobiology of Conodonts. Ellis Horwood Limited, Chichester.Google Scholar
Nicoll, R. S. 1991. Differentiation of Late Cambrian-Early Ordovician species of Cordylodus (Conodonta) with biapical basal cavities. Bureau of Mineral Resources Journal of Australian Geology and Geophysics, 12:223244.Google Scholar
Nicoll, R. S. 1992. Evolution of the conodont genus Cordylodus and the Cambrian-Ordovician boundary, p. 105113. In Webby, B. D. and Laurie, J. R. (eds.), Global Perspectives on Ordovician Geology. Balkema, Rotterdam.Google Scholar
Nowlan, G. S., and Thurlow, J. G. 1984. Middle Ordovician conodonts from the Buchans Group, central Newfoundland, and their significance for regional stratigraphy of the central volcanic belt. Canadian Journal of the Earth Sciences, 21:284296.CrossRefGoogle Scholar
Ortega, G., and Albanesi, G. L. In press. Graptolitos de la Formación Gualcamayo (Arenigiano-Llanvirniano) en el cerro Potrerillo, Precordillera Central de San Juan, Argentina. Boletín de la Academia Nacional de Ciencias, Córdoba, Argentina, 64.Google Scholar
Ortega, G., Albanesi, G. L., and Hünicken, M. A. 1995. Bioestratigrafía en base a conodontes y graptolitos de las Formaciones San Juan (techo) y Gualcamayo (Arenigiano-Llanvirniano) en el Cerro Potrerillo, Precordillera de San Juan, Argentina. Boletín de la Academia Nacional de Ciencias, Córdoba, Argentina, 60:317364.Google Scholar
Ortega, G., Toro, B., and Brussa, E. 1993. Las zonas de graptolitos de la Formación Gualcamayo (Arenigiano tardío-Llanvirniano temprano) en el norte de la Precordillera (provincias de La Rioja y San Juan), Argentina. Revista Española de Paleontología, 8:207219.Google Scholar
Peralta, S. H. 1995. La Formación Gualcamayo en la Sierra de Villicum: sus Graptolitos y faunas asociadas. Boletín de la Academia Nacional de Ciencias, Córdoba, Argentina, 60:401408.Google Scholar
Pohler, S. M. L. 1994. Conodont biofacies of Lower to lower Middle Ordovician Mega conglomerates, Cow Head Group, Western Newfoundland. Geological Survey of Canada Bulletin, 459:171.Google Scholar
Ridley, M. 1993. Evolution. Blackwell Scientific Publications, Oxford, 670 p.Google Scholar
Sadler, P. M. 1981. Sediment accumulation rates and the completeness of stratigraphic sections. Journal of Geology, 89:569584.CrossRefGoogle Scholar
Sánchez, T. M., Carrera, M. G., and Benedetto, J. L. 1996. Variaciones faunísticas en el techo de la Formación San Juan (Ordovícico temprano, Precordillera Argentina): significado paleoambiental. Ameghiniana, 33:185200.Google Scholar
Sarmiento, G. N. 1991. Conodontos de la Zona suecicus (Llanvirniano inferior) en la sierra de Villicum, Precordillera de San Juan, Argentina. Revista Española de Micropaleontología, 23:113132.Google Scholar
Sarmiento, G. N., and Rábano, I. 1992. Nuevas precisiones bioestratigráficas sobre la Formación Gualcamayo (Ordovícico inferior) en la sierra de Villicum, San Juan, Argentina. Zentralblatt für Geologie und Paläontologie, H., 6:17851797.Google Scholar
Sarmiento, G. N., Vaccari, N. E., and Peralta, S. H. 1988. Conodontes ordovícicos de La Rinconada, Precordillera de San Juan, Argentina. IV Congreso Argentino de Paleontología y Bioestratigrafia, Mendoza, Actas, 3:219224.Google Scholar
Schindel, D. E. 1980. Microstratigraphic sampling and the limits of paleontologic resolution. Paleobiology, 6:408426.CrossRefGoogle Scholar
Schoch, R. M., and Meredith, A. 1984. Punctuated patterns in the fossil record: the devolutionary mechanism. III Congreso Latinoamericano de Paleontología, México. Memoria, 624630.Google Scholar
Schopf, J. W. 1994. Disparate rates, differing fates: Tempo and mode of evolution changed from the Precambrian to the Phanerozoic. National Academy of Sciences USA Proceedings, Colloquium Paper 91:67356742.CrossRefGoogle ScholarPubMed
Sheldon, P. R. 1992. Microevolution and the Fossil Record, p. 106110. In Briggs, D. E. G. and Crowther, P.R. (eds.), Paleobiology: A Synthesis. Blackwell Scientific Publications, Oxford.Google Scholar
Skelton, P. W. 1992. Adaptation, p. 139146. In Briggs, D. E. G. and Crowther, P. R. (eds.), Paleobiology: A Synthesis. Blackwell Scientific Publications, Oxford.Google Scholar
Stone, J. 1987. Review of investigative techniques used in the study of conodonts, p. 1734. In Austin, R. L. (ed.), Conodonts: Investigative Techniques and Applications, Ellis Horwood Limited, Chichester.Google Scholar
Stouge, S. 1984. Conodonts of the Middle Ordovician Table Head Formation, western Newfoundland. Fossils and Strata, 16:1145.Google Scholar
Stouge, S., and Bagnoli, G. 1988. Early Ordovician conodonts from the Cow Head Peninsula, western Newfoundland. Palaeontographia Italica, 75:89179.Google Scholar
Sweet, W. C. 1988. The Conodonta: Morphology, taxonomy, paleoecology, and evolutionary history of a long-extinct animal phylum. Oxford Monographs on Geology and Geophysics, 10 (Oxford University Press, New York), 212 p.Google Scholar
Thomas, W. A., and Astini, R. A. 1996. The Argentine Precordillera: A traveler from the Ouachita Embayment of North American Laurentia. Science, 273:752757.CrossRefGoogle ScholarPubMed
Zhang, J. 1998. Conodonts from the Guniutan Formation (Llanvirnian) in Hubei and Hunan Provinces, south-central China. Acta Universitatis Stockholmensis, Stockholm Contributions in Geology, 46, 161 p.Google Scholar