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(Upper Devonian) Conodont Zonations

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(Upper Devonian) Conodont Zonations Bollettino della Società Paleontologica Italiana Modena, Novembre 1999 Comparison ofFrasnian (Upper Devonian) Conodont Zonations Gilbert KLAPPER R. Thomas BECKER Deparrmem of Geology lnstitut fiir Palaomologie University oflowa Humboldt Universitat, Berlin KEYWORDS- Frasnian conodont zonations, Martenberg and Montagne Noire sequences, Palmatolepis taxonomy. ABSTRACT- Previously it has no t been possible to correlate the thirteen-fold Frasnian conodont zonation first developed in the Montagne No ire, southern France, with the nine-fold standard zonation ofthe Frasnian. The impasse has been due to the Jact that the taxonomies underlying the two zonations are so disparate that comparisons offounallists and range charts lead to substantial misalignments. We propose here an alignment of the two zonations, based on our sampling of the Martenberg reference section of the standard zonation in the Rfienish Siate Mountains of Germany, combined with application oj the taxonomic concepts underlying the Montagne Noire zonation to the conodonts recovered. RIASSUNTO- [Confronto tra le biozonacure a conodomi del Frasniano (Devoniano sup.)] -Per molto tempo non è stato possibile correlare le tredici biozone della biozonatura a conodonti del Frasniano, basata sulle sequenze della Montagna Nera, con le nove della Biozonatura Standard del Frasniano. Questo era dovuto principalmente al fotto che i criteri tassonomici alla base delle due biozonature erano talmente diversi che il confronto di elenchi di faune e tabelle di distribuzione provocava notevoli errori nei tentativi di correlazione. Una correlazione tra le due biozonature viene qui proposta, sulla base di una nuova campionatura della sezione di Martenberg {sezione di riferimento della biozonatura standard, nelle Montagne del Reno, in Germania), alla quale sono stati applicati i criteri tassonomici che stanno alla base della biozonatura della Montagna Nera. INTRODUCTION Because the taxonomies underlying the two zonations are so disparate, correlation between the two A thirteen-fold conodont zonation of the Frasnian zonations can no t be resolved by comparison of faunal was proposed for the Montagne Noire sequence in lists and range charts. This impasse has been southern France (Klapper, 1989), originally as a local emphasized severa! times (Sandberg et al., 1992; zonation describing that sequence. The Montagne Klapper & Foster, 1993, p. 3; Ziegler & Sandberg, Noire sequence, however, has since been replicated at 1994, p. 119). Previous attempts to align the two other Devonian tropical sites, for example, western New zonations (e.g., Sandberg et al., 1992, p. 30, fig. 12; York (Kirchgasser, 1994; Kralick, 1994), the Alberta Becker & House, 1997, fig. 9) are inaccurate due t o Rockies and centrai Alberta subsurface in Canada the disparate use of species concepts of Ancyrognathus (McLean & Klapper, 1998), the Canning Basin of (especially A. triangularis Youngquist, 1945) and Western Australia, and the Timan-Pechora region of principally of Palmatolepis, which accounts for the the Russian Platform (Klapper et al., 1996). The radically different ranges of such species as P. hassi underlying taxonomy is based o n shape analysis of Pa Mi.iller & Mi.iller, 1957 (Ziegler & Sandberg, 1990, elements of Palmatolepis, multielement taxonomy, and fig. 2; Klapper & Foster, 1993, fig. 2). visual discrimination of species of Palmatolepis and We have attempted to resolve the problem of other genera in which mudtielement taxonomy and alignment of the two zonations by sampling the shape analysis have not yet been determined. Species reference sections in the Rhenish Siate Mountains of ofAncyrodella, Ancyrognathus, an d Ozarkodina that are Germany that were the basis for the zonation ofZiegler used in the Montagne Noire zonation show first & Sandberg ( 1990), an d then applying the taxonomic occurrences as consistent as those of Palmatolepis, as concepts underlying the zonation first proposed for the determined through a Frasnian Comrosite Standard Montagne Noire Frasnian sequence to the conodonts developed from graphic correlation (Klapper, 1997; recovered. herein). A nine-fold zonation, based on largely different taxonomic concepts, has been proposed for the Frasnian MULTIELEMENT TAXONOMY OF PALMATOLEPIS (Ziegler & Sandberg, 1990) on the basis of German and Great Basin (Utah and Nevada) sequences. This lt would seem unnecessary to justif}r the use of is said to be a standard zonation, the zones are conceived multielement taxonomy to discriminate species of of as stricdy time, rather than biostratigraphic, units, Palmatolepis a t this late date, in view of numerous papers and the taxonomy of the species defining zona! that present evidence on this subject (e. g., Lange, 1968; boundaries is almost exclusively based on visual Boogaard & Kuhry, 1979, Puchkov et al., 1982; discrimination of Pa elements of Palmatolepis. Klapper & Foster, 1986, 1993; Schi.ilke, 1997). 340 G. KLAPPER, R. T. BECKER Nevertheless, in a formai 27-page systematic section on Palmatolepis and Mesotaxis, no other elements but MD N the Pa elementare considered by Ziegler & Sandberg Zone l Ancyrodella rotundiloba early form .52 2 (1990, pp. 43-69), although they state in an earlier section (p. 8) that they have "provisionally assigned Zone 2 Ancyrodella rotundiloba late form .41 3 most Pb and other elements of Palmatolepis." Zone 3 Ancyrodella rugosa .39 3 Furthermore, Ziegler & Sandberg (1990, p. 10, first Zone 6 Ancyrognathus primus .92 lO paragraph) disparaged the multielement reconstructions of Palmatolepis winchelli (Stauffer, 1938 Zone 7 Ozarkodina nonaginta .89 7 = P subrecta Miller & Youngquist, 1947) and P Zone 4 Palmatolepis transitans 1.74 15 bogartensis (Stauffer, 1938 = P rotunda Ziegler & Zone 5 Palmatolepis punctata 1.70 14 Sandberg, 1990) published by Klapper & Foster (1986) and Klapper (1989, p. 458). The first opportunity to Zone 8 Palmatolepis aff. P proversa .96 5 present the evidence in detail for these two Zone 9 Palmatolepis proversa 1.26 13 reconstructions was given by Klapper & Foster (1993, Zone 10 Palmatolepis plana .99 13 pp. 18, 26, 31, figs. 19, 20). The reconstruction of Palmatolepis bogartensis was independently replicated Zone 11 Palmatolepis sp. B .60 8 by Schi.ilke (1997, p. 44, pl. 3, figs. 1-20), on the Zone 12 Palmatolepis winchelli 4.60 lO evidence ofMontagne Noire collections from La Serre Trench C. Klapper & Foster (1986, 1993) had based Zone 13 Palmatolepis bogartensis 1.98 4 their reconstruction of the same species on the fused Upper rhenana Palmatolepis rhenana 3.46 7 cluster of Lange (1968) and isolated specimens from Zone the Lower Coumiac Quarry and Causses-et-Veyran South in the Montagne Noire. Zone 5 Polygnathus timanicus 1.29 7 Zone 12 Polygnathus samueli 1.35 3 Zone 12 Ancyrodella ioides .17 5 TESTING THE FRASNIAN ZONATIONS BY GRAPHIC CORRELATION Zone 11 Ancyrognathus seddoni .67 2 Zonations, especially zonations that are claimed to be geographically widespread or even global (Ziegler T ab. l - The fìrst column of values (MD) is the mean distance & Sandberg, 1990, p. 12) rest on the assumption that berween the lowest occurrence of the zonally defìning the first occurrences of species used to define zonal species and the line of correlation (LOC) in ali the graphs boundaries are isochronous (or "virtually synchro- of the Frasnian Composite Standard, given in composite nous"). Before relying o n this as a working assumption, standard units. The second column is the number (N) of occurrences used to calculate the mean. For further the isochronous character of zonally defining species explanation see the discussion of graphic correlation in must be demonstrated (Klapper, 1997, p. 113). Testing the text. of this proposition can be carried out by means of graphic correlation, which, among other attributes, permits the assessment of the degree of departure of zonally defining species from isochronous first occurrences at ali sections graphed. Tests of diachronism sequence, plus one of the zonally defining species, in the light of graphic correlation and indications of Pa. rhenana rhenana Bischoff (1956), of the zonation the way in which this can be portrayed are given in the of Ziegler & Sandberg (1990), a species that has bee n papers of Belka et al. ( 1997, fig. 8) for the Middle used consistently in both zonations. Four other species Devonian ofMorocco and ofK.lapper (1997, pp. 116- are shown at the botto m of the table. The first column 120, fig. 4) for Frasnian sections in the Montagne Noire of values (MD) is the mean distance between the lowest and western Canada. occurrence and the line of correlation (LOC) in ali the The Frasnian Composite Standard based on the graphs, given in composite standard units, and the method of graphic correlation has been developed over second column is the number of occurrences used to a l 0-year peri od and now consists of 38 graphed calculate the mean. Occurrences that are no t considered sections (two of which are regional composites of an in this calculation are those that are used to locate the additional number of sections). Various results derived LOC in a given graph and those that have first from the Frasnian Composite Standard have been given occurrences on a sampling terrace. in several papers (Klapper & Foster, 1993, fig. 2; Study of one of the key graphs in the Frasnian Klapper et al., 1995; 1996; and Klapper, 1997). Composite Standard (CS), that of the La Serre Trench Table l lists ali the zonally defining species for the D section plotted against the CS (Klapper, 1997, fig. 13 zones first proposed for the Montagne Noire l), indicates that many of the first occurrences of species COMPARISON OF FRASNIAN CONODONT ZONATIONS 341 of the conodont genera Ancyrodella, Ancyrognathus, rhenana Zone, and Zone 13 with a position within the and Ozarkodina fall as dose to the LOC as those of Upper rhenana Zone. Previously we equated Zone 11 Palmatolepis. In other words, they are as consistent in entirely with the jamieae Zone an d the base ofZone 12 their first occurrences as Palmatolepis. The results with the base of the Lower rhenana Zone (Klapper & presented in Table l expand the implications from the Becker, 1998, p.
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