Relationships Between Ordovician Baltic and North American Midcontinent Conodont Faunas
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Relationships between Ordovician Baltic and North American Midcontinent conodont faunas JERZY DZIK FOSSILS AND STRATA Dzik, Jerzy 1983 12 15: Relationships between Ordovician Baltic and North American Midcontinent conodont faunas. Fossils and Strata. No. 15, pp. 59-85. Oslo. ISSN 0300-9491. ISBN 82-0006737-8. During most of the Early and Middle Ordovician, epicontinental seas of Laurentia, Baltica, and islands in the Iapetus Ocean between them were centers of diversification of different groups of conodonts. The most important were Phragmodontidae in Laurentia, Periodontidae in the Iapetus area, and Balognathidae in Baltica. Spectra of fossil assemblages (analyzed in terms of conodont lineages) were very stable through time in each of these biogeographic provinces. Relatively few lineages passed the biogeographic boundaries and successful immigrations were rare. An especially stable composition characterizes the Baltic faunas. A profound change in the composition of faunas in Baltica took place in the Oanduan (late Caradocian) through influx of several lineages previously confined to islands in the Iapetus. Conodont faunas of Laurentia, at times enriched by Iapetus-born lineages, evolved rather gradually until the end of the Ordovician. Diversity of Ordovician conodont assemblages seems to depend more on local ecologic factors and bathymetry than on climate. OConodonta, biostratigraphy, biogeography, evolution, Ordovician. ECOS III Jerzy Dzik, Department of Geology and Mineralogy, The Ohio State University, Columbus, Ohio 43210, A contribution to the Third European U.S.A. (Present address: Zaktad Paleobiologii PAN, Aleja Żwirki i Wigury 93, PL-02-089 Warszawa, Conodont Symposium, Lund, 1982 Poland); 11 th August, 1982. Ordovician bioprovincialism has received much attention systems of terminology for elements in the conodont apparatus from conodont workers and several important contributions (for review, see Sweet 1981 b; also Barnes et al. 1979 and Dzik & to its recognition have been published (for review, see Trammer 1980) two seem to be applicable for most apparat- Jaanusson 1979; Sweet & Bergstrom 1974; Jaanusson & uses. Those are Jeppsson's (1971) and Sweet's (1981b) Bergstrom 1980; Lindstrom 1976 b). The profound difference systems. When applied to sextimembrate apparatuses they between conodont faunas of the North American Midconti- are easily transferable to each other. nent and those of the southeastern Appalachians and Europe For the purpose of the present paper Jeppsson's notation has been recognized since the Sweet el al. (1959) paper. A has been chosen. It was originally introduced as a tool for boundary separating these two distinct biogeographic units expressing homology of elements. Symbols of all types of has been traced along the Helena-Saltville fault in the elements are derived by abbreviations of former form- southern Appalachians and corresponding structural features taxonomic generic names, which seems to be in good agree- to the north (Bergstrom 1971; Jaanusson & Bergstrom 1980). ment with tradition in biological terminology. In the same way Although changes in the distribution of particular Late terms for, say, larval stages (echinospira, pilidium, calyptopis, Ordovician conodont taxa have been discussed in detail by zoea, etc.), or organs (stigmaria, helens, aptychi, etc.) have Sweet & Bergstrom (1974) and faunal changes across the been introduced, all being based on names of former taxa. Appalachians in the Middle Ordovician have been presented This does not leave a place for uncertainty regarding reference by Bergstrom & Carnes (1976), much remains to be done for homologization. Particular types of elements are identified regarding the pattern of distribution and shifts of Early and in any apparatus by homologization with particular elements Middle Ordovician conodont lineages in Europe as well as of the Ozarkodina apparatus. There is no assumed a priori order precisely tracing relationships of those lineages that are in arrangement of the types of elements in the apparatus, thought to be typical of provinces. It must be stated, however, therefore no trouble appears when insertion of a new type of that data are still far from complete, and numerous key faunas element into the transition series appears necessary. (for example those from the Middle Ordovician of Great The apparatus of Amorphognathus is proposed here as a Britain and the Sudeten Mountains) have not yet been homologization standard for septimembrate apparatuses. described in terms of multielement taxonomy. Accordingly, The complete set of elements in such apparatuses contains, this paper is just a preliminary report on a study that remains according to this notation (in supposed order), sp, oz, tr, pi, to be completed. ke, hi, and ne types. Sweet (1981b) and Bergstrom (1981) proposed to consider the keislognathiform element of Amor- phognathus as pi (Sb) and consequently created 'location' Sd for Terminology tetraprioniodontiform elements. There are, however, analog- Choice of the best element-notation system begins to be a ous tetraramous elements in the pi (Sb) location in apparat- difficult task for conodontologists. Among several proposed uses other than those of the Balognathidae (for example 60 Jerzy Dzik FOSSILS AND STRATA 15 (1983) Microzarkodina and Paraprioniodus), while it has been asserted change in the distribution of some organisms. The concepts of that the keislognathiform element is a homologue of one of the island biogeography (MacArthur & Wilson 1967) are applic- Sc elements in those Balognathidae that do not have it (Dzik able to such biogeographic units. It is not difficult to find 1976). In the typical apparatus of the Balognathidae, Sa, boundaries in the terrestrial environment with the features tetraprioniodontiform (Sb), keislognathiform, and Sc ele- required by this definition. The marine environment is much ments form a single symmetry transition series, and a keislo- more continuous, but discontinuities in the distribution of gnathiform element should be inserted between Sb and Sc particular marine environments such as margins of the elements. This could be done either by introducing a third continental shelf, areas of convergence of warm and cold letter to the notation (like Sbb) or by replacing Sc elements water, thermoclines, a range of uplifts of oceanic currents sensu Sweet (1981b) to Sd location and reserving the Sc carrying biogenes, etc., can be used to delineate boundaries location only for homologues of the keislognathiform elements between marine ecosystems. All these factors may separate of Amorphognathus (which I would prefer). Kuwano (1982) oceanic water into discrete cells that are different in the proposed symbols Sa-b and Sb for homologues of pi and ke composition of their faunal and floral assemblages. Features of elements in the Ozarkodina excavata (Branson & Mehl) these cells may change with the evolution of climate and even apparatus, respectively. more profoundly with the tectonic evolution of continents Meanings of other terms used in the text are explained in the (Williams 1976). In the present paper the Ordovician epicon- Treatise on Invertebrate Paleontolog)! (Sweet 1981 b). There also all tinental seas of Laurentia (the Midcontinent province), taxonomic, auctor references that are not listed in this paper Baltica (the Baltic province), and islands in the Iapetus Ocean can be found. between them (the North Atlantic province) are considered to be paleobiogeographic units of this kind (Fig. 1). Comments on methodology Four main kinds of faunal processes may be connected with the evolution of biogeographic units defined above: (1) One basic trouble in establishing a good scientific framework Phyletic evolution of particular species. (2) Change in the for a discussion of conodont provincialism is the lack of distribution of particular species within boundaries of the unequivocal meaning for units in paleobiogeography. Con- unit. (3) Immigrations of species from other ecosystems, cepts of high-rank biogeographic units are usually based on which displace local species from their niches, but do not more or less objectively determined differences in the composi- change the state of faunal equilibrium. (4) Immigrations of tion of faunistic assemblages from different areas. This does species that occupy previously uninhabited niches, or destruc- not create special difficulties unless problems of the boundary tion of the previous distribution of niches, in both cases between particular units and their change in geologic time are changing faunal equilibrium. involved. Then any biogeographic unit appears to have a How can one recognize these basically different processes particular meaning that depends on methods used in recogniz- from the limited fossil record? It would be especially important ing it and the particular group of fossils used, rather than on to have a tool that permits distinction between rebuilding of objective factors (Jaanusson 1979). For these reasons Lind- the fossil assemblage caused by a local environmental change strom (1976 a) has proposed to separate the territorial and and immigration of extrinsic faunal elements. Solution of this faunistic aspects of provincialism, and to abandon the first one problem seems to be the first step to the serious considerations and shift the discussion toward aspects of the evolution of of bioprovincialism (see discussion in Sweet & Bergstróm particular faunas. These may completely change their dis- 1974; Barnes et al. 1973; Bergstrom