Bollettino della Società Paleontologica Italiana Modena, Novembre 1999

Comparison ofFrasnian (Upper ) 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, 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 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 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 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 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. 54). That interpretation was based on La Serre Trench D graph into the generalization that the columnar diagram for the Martenberg sequence Ancyrodella, Ancyrognathus, an d Ozarkodina have some published in Ziegler & Sandberg (1990, fig. 3) , which species that are no greater in their degree of diachronism shows the southwest section equivalent ofBed Q of the than those of Palmatolepis, and in fact some are north section entirely within the jamieae Zone (as is demonstrably closer to being isochronous. repeated here in Text-fig. 2). However, in unpublished Palmatolepis winchelli of the Montagne Noire collections from the north section made in 1984 and zonation and P. rhenana rhenana of the standard 1987, Ziegler & Sandberg (1999, written communi- zonation are highly diachronous in the sections graphed for the Frasnian Composite Standard and are thus suspect as zonal indices. The four species at the bottom ofTable l have not "M o n tagne N o i re " S ta nd ard Zones a t Z been used previously as zonally defining species, but, o n es Martenberg as demonstrated by graphic correlation, are highly useful as zonally characterizing species (in the termi- l ingu iformis nology of Murphy, 1977). 1 3 In conclusion to this section (and as long ago Upp e r rhenana demonstrated by Shaw, 1964), grafhic correlation offers an inherently higher degree o resolution than any kind of zonation. 12

L ower rhe n ana CORRELATION OF THE TWO ZONATIONS 11 iamieae Correlation of Montagne Noire (the numbered zones refer to the Frasnian Montagne Noire sequence 1 0 in all instances) Zones 5 through 12 with the standard U pp e r h ass i zonation (Ziegler & Sandberg, 1990) has been 9 accomplished through our detailed sampling and analysis of the Martenberg sequence of the Rhenish 8 Siate Mountains, the reference section for the lower Lo w er h assi boundaries of the punctata through Lower rhenana Zones in the standard zonation. This is supplemented 7 by the correlation ofZone 13 with the Upper rhenana and linguiformis Zones, through analysis of additional 6 samples at Martenberg and selected samples at punctata Steinbruch Schmidt, the reference section for the latter 5 two zones. We have taken the boundaries of the standard Frasnian zonation, as shown in Text-fig. l , from Ziegler & Sandberg's (1990, fig. 3) columnar 4 tra n sitans diagram representing the southwest corner profìle at Martenberg. lt must be noted, however, that the lower Text-fìg. l - Correlation ofMomagne No ire zones with the standard boundaries of some of the standard zones (jamieae zones of the Frasnian. The reference section for most through Upper rhenana Zones) were misaligned with of the standard zones is Martenberg near Adorf in the the southwest column through drafting errors in the Rhenish Siate Moumains (Ziegler & Sandberg, 1990). cited 1990 figure (W. Ziegler & C.A. Sandberg, 1999, The alignment proposed here is based o n our sampling oral communication). of the Martenberg section and the applica rio n of the taxonomic concepts used for the Montagne Noire Salient levels of correlation are as follows, in all zonation to the conodoms recovered. The question instances referring to the lower boundaries of zones mark at the position of the Zone 8-9 boundary is (Text-fig. 1): Zone 5 with the punctata Zone, Zone 7 because our samples from Bed R could represem either with the Lower hassi Zone, Zone 9 approximatelywith uppermost Zone 8 or Zone 9 (Table 2). It is unwarramed to conclude from this diagram that the rh e U p per hassi Zone, Zone 11 either with the jamieae two zonations can be equated in exact!y this Zone or perhaps near the base of the Lower rhenana confìguration at sections other than Martenberg. The Zone, Zone 12 with a position within the Lower equation at other sections is a matter for future testing. 342 G. KLAPPER, R. T BECKER

MN Zones North Side Southwest Corner

k 13 llngulform;s 13 13 upper gigas

13 upper rhenana

2

n

12 o lower glgas lower rhenana

12 p 12

11 6 7 11 jamieae q Ancyrognatflus 8 triangulari s l l 9

10 r--q 10

upmos8·9 11 upper hassi 12

8 i.b.r 13 uppe r asymmetr i ca 14 lower hass ;

s 15

6 m1ddle asymmetrica 16 puncla/a 5 u

Texr-fìg. 2- Columnar diagrams of part of the sequence at Martenberg near Adorf in the Rhenish Siate Mountains. Left column after that of Ziegler (1971, fìR. 6), which depicts the section on the north side. Apparenrly rhe same column, re-drafted, was used to represent the Ma VI section at the southwest corner in Ziegler & Sandberg (1990, fig. 3), here repeated as the right column. Montagne No ire (MN) Frasnian zones shown along left margin, earlier standard zones (Ziegler, 1971) are to the right of rh e north column, and revised standard zones (Ziegler & Sandberg, 1990) are along right margin of diagram. Bed letters from Ziegler (1971) are immediarely to the left of the north section; sample numbers for the Ma VI' section (southwest corner section) are immediately to right of the right column. Note that the jamieae!Lower rhenana zona! boundary is near the base of Bed Q of the north section according to unpublished information kindly provided by W. Ziegler & C.A. Sandberg (1999, written communication; see discussion in text under Correlation of the Two Zonations). Furthermore, the exact alignment of some of the revised standard zones (jamieae through Upper rhenana Zones) with the Ma VI' column is currenrly under revision by Ziegler & Sandberg (1999, ora! communication; see also discussion in cited section of the text).

cation) have identifìed the base of the Lower rhenana time since there is no controversy over the taxonomic Zone (or Early rhenana Zone in their terms) at a Iower identity of the nominai species, Palmatolepis position, near the base ofBed Q. This means that Zone linguiformis Miiller (1956), in contrast with most of 11 mainiy correlates with a part of the Lower rhenana the other defìning species of the two zonations. Listing Zone, an d perhaps oniy the Iowermost p art ofZone 11 of ali our collections from Martenberg is given in Tabies correlates with the jamieae Zone (Text-fìg. 1), if at ali. 2 and 3, providing evidence for the foregoing zona! The exact relationship must remain uncertain pending identifìcations. the current revision of their Martenberg sampies by W The emphasis in this study has been on the interval Ziegier & C. A. Sandberg. from Zone 5 to Zone 13, as that has been the principal The uppermost part ofZone 13 correlates with the probiem of alignment of the two zonations. In the linguiformis Zone, as has been recognized for some interest of compieteness, it can be stated that the base COMPARISON OF FRASNIAN CONODONT ZONATIONS 343 of Zone 4 correlates with the base of the transitans the specimens on Plate 6 with a well differentiated Zone (Sandberg et al., 1992, p. 30, fig. 12; Becker & lobe are figs. 1-3 (the holorype), 6, and 8, whereas House, 1997, fig. 9), since the lower boundary ofboth those in figs. 4, 5, 7, 9, 10 belong to the second is defined by the lowest occurrence of Palmatolepis morphotype. It is unclear, however (compare Bultynck transitans Mi.iller (1956) about which there has been et al., 1998, p. 58). some degree of consistency in taxonomic concept. The species termed in open nomenclature as Correlation of Zones 1-3 is with most of the folsiovalis Palmatolepis sp. B (Klapper & Foster, 1986) has been Zone (Becker & House, 1997, fig. 9). synonymized under P. jamieae both by Ziegler & Sandberg (1990, p. 50) and Klapper (1997, p. 125; also the intention, though not discussed, in the range CORRELATIONS AT OTHER FRASNIAN SECTIONS chart of Klapper & Foster, 199 3, fig. 2). In terms of platform o udine an d the lack of sinuses strongly demar- Resolution of the correlation of a substantial part cating the outer lobe in the Pa element, Palmatolepis of the Montagne Noire zonation with the standard sp. B can be distinguished from the holorype of P. zonation as it is represented at Martenberg and jamieae Ziegler & Sandberg (1990, Pl. 6, figs. 1-3) and Steinbruch Schmidt does not mean that the two also specimens on their Plate 6, figs. 6 and 8. An almost zonations can be routinely equated with confidence at straight anterior and posterior carina is commonly sections elsewhere. The identification of the Lower characteristic of P. sp. B, including the specimens in rhenana Zone (or Early rhenana Zone) at the Lion Ziegler & Sandberg's Plate 6, figs. 4, 7. Palmatolepis Quarry access road, Frasnes, Belgium, as given by sp. B, which should be the basis of a new species, is the Sandberg et al. (1992, table 2) from Bed 113 up defining species for the base of MN Zone 11, whereas through Bed 164 ofMouravieff(l970), may no longer the holorype of P. jamieae is from the Lower rhenana be an apparent discrepancy, however, in contrast with Zone at Steinbruch Schmidt. The latter has a much our earlier interpretation (Klapper & Becker, 1998, p. more sigmoidal carina than in representative specimens 54). The associateci faunas in these beds include species of P. sp. B. that range no higher than the lower part of Zone 11 (Ancyrognathus seddoniKlapper, 1990, in Bed 155 and P. 'aomanicensis"ofKlapper, 1989, as high as Bed 157b PALMATOLEPIS aff. P. WINCHELLI Klapper & Lane,l989 in the collections of Mouravieff, 1970). Additionally, the specimen in Bed 125 at Lion Quarry identified by 1989 Palmatolepis aff. P winchelli (Stauffer) - .I

element of the new species can be distinguished from (1993, p. 5) and involve more than the discriminant the related P. sp. B, in that the latter characteristically analysis of the platform oudine (contra Ziegler & has a much straighter carina and a proportionately Sandberg, 1994, p. 113). wider platform. Thus, we consider that the specimens of Ziegler & Sandberg (1990, pl. 6, figs. 4, 7) are better identified as P. sp. B, as opposed to the synonymy PALMATOLEPIS HASS I Mi.iller & Mi.iller, 1957 s.s. given in Bultynck et al. (1998, p. 60). 1957 Palmatolepis (Manticolepis) hassi MùLLER & MùLLER, pp. 1102-1103, pl. 139, fig. 2; pl. 140, fi gs. 2-4 (fig. 4 = PALMATOLEPIS DOMANICENSIS Ovnatanova, 1976 holotype). 1993 Palmatolepis hassi Mtiller & Mtiller -KlAPPER & FosTER, p. 22, figs . 15.1-15.9 (fig. 15.4 = reillustration of holorype); 197 6 Palmatolepis domanicensis OVNATANOVA, p. l 09, p l. 9, figs. fig , 18.12; figs . 19.16-19.18 [see funher synonymy]. l, 2 (fig. 2 = holorype). 1998 Palmatolepis hassi Mtiller & Mtiller s.s. -BULTYNCK, HELSEN, 1996 Palmatolepis domanicensis Ovnatanova- KlAPPER, Kuz'MIN, & 0 VNATANOVA, p. 145, fig. 9.12 (only; reillustration of & HAYD UCKI EWIC H, p. 58, pl. 2, fig. 6; pl. 3, figs. 2-7. holorype). Remarks- Klapper & Foster (1993) and Bultynck Remarks- Restudy of the holotype of Palmatolepis et al. (1998) have given descriptions of the Pa element domanicensis in ]une 1998 clearly indicates that i t falls (and the Pb element in the former paper) and have outside the range of variation of specimens included advocated a restricted concept of this species centered within the species by Klapper (1989, and Klapper & around the type material. As such, the species ranges Foster, 1993). This was already suggested by Klapper from the upper part of MN Zone 12 into the lower et al. ( 1996), in that the holotype differs from the latter half ofZone 13 in terms of equivalencies in the Frasnian specimens by having a distinct centrai node and an Composite Standard. In our sampling at Martenberg extremely weak posterior carina. Visually, its platform (Appendix), the species does not occur below the level oudine differs as well. of MN Zone 13. The lower range at Martenberg In a discriminant analysis running the holotype as reported by Ziegler & Sandberg (1990, table l) as low an unknown against sets of Palmatolepis foliacea, P. as their sample 15 in the Lower hassi Zone (Bed S in luscarensis, and specimens attributed toP. domanicensis Text-fig. 2) must involve other species, if one accepts by Klapper & Foster (1993, fig. 3.3), the holotype plots the restricted definition of P. hassi. Except for their in the field between the three separate clusters but reillustration of the holotype (Ziegler & Sandberg, closest to that of P. foliacea. Thus, statistica! analysis of 1990, pl. 2, fig. 2) , the rest of the figured specimens the platform oudine of the holotype confirms the identified as P. hassi on their plates 2 and 12 represent results of visual discrimination cited above. Although a variety of other species that are incongruent with the there is some resemblance to the Pa element of P. holotype. They are distinguishable from P. hassi s.s. o n foliacea, it is interpreted here as an instance of the basis of a number of morphologic features, convergence because the holotype of P. domanicensis is especially the details of the platform o udine. from the upper part of the Middle Member of the Domanik Formation (which has conodont faunas correlative with MN Zone 6), whereas P. foliacea ranges p ALMATOLEPIS PLANA Ziegler & Sandberg, 1990 from MN Zone 12 to the lowest part ofZone 13. Thus, the specimens incorrecdy assigned to 1989 Palmatolepis domanicensis Ovnatanova- KlAPPER, 1989, p. Palmatolepis domanicensis by Klapper (1989) and 458, p l. l , figs. l , 2. Klapper & Foster (1993, p. 5) should now have the 1990 Palmatolepis plana ZIEGLER & SANDBERG, p. 46, pl. 3, figs. 1-10. name P. plana Ziegler & Sandberg (1990, p. 46, pl. 3, 1992 Palmatolepis plana Ziegler & Sandberg - SANDBERG et al. , figs. 1-1 O). This does no t mean, however, that ali pp. 70, 72, pl. 2, fig . lO; pl. 3, fig. 10. specimens identified as P. plana are necessarily 1993 Palmatolepis domanicensis Ovnatanova - KlAPPER & FosTER, congruent with P. "domanicensis" of Klapper (1989), pp. 5, 8, figs . 4.1-4.8, 6.10-6.13, 7. 15, 7.16, 10.9-10.12 because Ziegler & Sandberg (1990, table l) show P. lsee further synonymy]. 1996 Palmatolepis domanicensis Ovnatanova - KlAPPER, Kuz'MIN, plana ranging as low as the Lower hassi Zone at & 0 VNATANOVA, p. 145, fig . 9.1 1 (onJy; fig. 9.12 = Martenberg. In our sampling (Appendix) , P. plana in reillustration of holorype). a restricted sense (= P. "domanicensis') does no t range below a position high in the Upper hassi Zone, Remarks - Palmatolepis plana (= P. "domanicensis" correlative with MN Zone 10 (Text-fig. 1). Lower of Klapper, 1989) can be distinguished from the occurrences of P. plana at Martenberg probably refer holotype of P. domanicensis and other closely similar instead to P. bohemica Klapper & Foster (1993), a specimens from the Middle Member of the Domanik species which Ziegler & Sandberg (1994, p. 113) do Formation (MN Zone 6) in the southern Timan region not recognize. Differences between P. bohemica and P. of Russia, as discussed in the Remarks on the latter "domanicensis"have been specified by Klapper & Foster specres. COMPARISON OF FRASNIAN CONODONT ZONATIONS 345

APPENO IX OF CONODONTS IDENTIFIED FROM occurrences of Palmatolepis winchelli and Ancyrodella MARTENBERG ioides Ziegler (1958) are diagnostic for the identification of the base ofZone 12. Ancyrodella ioides In Tables 2 and 3, bed designations are those from is used in a restricted definition, with no platform on the col umnar diagram in Ziegler (1971, fig. 6; al so either side of the secondary carinae of the Pa element. shown in House & Ziegler, 1977, fig. 2), which Bed P, like Bed Q, is rich in goniatites (manticoceratids represents the section at Martenberg on the north side. and tornoceratids) and bivalves. All samples collected by RTB and GK from the north Bed M, lower 5-6 cm of 17 cm thick bed. section, except for that from Bed U, which is from the Polygnathus n. sp. BT is an unusual species of northwest section shown in Ziegler (1958, fig. 2, Polygnathus that occurs just below and in the Lower sample 3a). Authors of all species names are cited in Kellwasser Limestone at Causses-et-Veyran South and the text on first mention. Upper Coumiac Quarry in the Montagne Noire, and just below (5 cm) the Lower Kellwasser Limestone at Bed U, 6 cm thick. One specimen of Polygnathus Steinbruch Schmidt. The upper few cm ofBed M has timanicus Ovnatanova (1969) has been illustrateci by a rich goniatite fauna, almost a goniatite coquina with GK (in Ziegler, 1991, p. 153-154, Polygnathus- Plate orthocones and Buchiola. We interpret the upper part 14, fig. 3). Polygnathus linguiformis linguiformis gamma of Bed M as the equivalent of the Lower Kellwasser morphotype of Bultynck (1970) is interpreted as Limestone. reworked at this level. Bed K, lower 6 cm an d 6-1 O cm above base of 23 Bed T, 12 cm thick. Lowest occurrences of cm thick bed. These lower to middle parts of Bed K Ancyrognathus primus Ji (1986) and Ancyrodella gigas represent the Upper Kellwasser Limestone horizon. The form 2 of Klapper (1989) are indicators of Zone 6. samples in the lower 6 cm an d a t 6-1 O cm above base Bed S, upper 4 cm. Mesotaxis asymmetrica (Bischoff have Palmatolepis linguiformis (Tab. 3) and the higher & Ziegler, 1957) is probably reworked and Polygnathus of the two samples has Ancyrognathus ubiquitus linguiformis linguiformis gamma morphotype is Sandberg et al. (1988), which are diagnostic of the definitely reworked at this level. The lowest occurrence linguiformis Zone. The upper l O cm ofBed K contains of Ozarkodina nonaginta Klapper et al. (1996) is a mixed fauna of Frasnian Zone 13 and species that diagnostic for Zone 7. have first occurrences in the lower Lower Sample from bed immediately below Bed R (i.b.R). triangularis Zone (P. triangularis Sannemann, 1955a) The lowest occurrences ofAncyrognathus coeni Klapper and Upper triangularis Zone (P. minuta minuta Branson (1990) andAncyrodella gigas form 3 ofKlapper (1989) & Mehl, 1934, and P. tenuipunctata Sannemann, and the highest occurrences of Ozarkodina nonaginta 1955b). and O. trepta (Ziegler, 1958) are in Zone 8. Bed R, samples from upper 7 cm an d upper l Ocm. Possible zonal range is from uppermost part ofZone 8 ACKNOWLEDGMENTS to Zone 9, because diagnosti c markers to distinguish We thank William T. Kirchgasser for a criticai reading of che the two zones are lacking. Palmatolepis mucronata manuscript. Nicholas Mouravieff kindly allowed study of his Klapper et al. ( 1996) has its lowest occurrence in the collections from che Lion Quarry access road in 1987 and 1990. Rhawn Denniston signifìcandy helped with the preparation of upper part of Zone 8 a t Horse Spring in the Canning Text-fìg. 2. Thanks are also extended to Pierre Bultynck, Ruth Basin and P. ljaschenkoae Ovnatanova (1976) has its Mawson, and John Talent for reviews of an earlier version of che lowest occurrence in the Frasnian Composite Standard manuscript. Las t but no t least, we thank Willi Ziegler and Charles in the equivalent of the uppermost part of Zone 8. A. Sandberg for keeping us fully informed of current revisions of Bed between R and Q, 13 cm thick. Palmatolepis che Martenberg conodont sequence. plana is at least partly a synonym of P. "domanicensis" of Klapper (1989, and Klapper & Foster, 1993; see Taxonomic Notes). REFERENCES Bed Q, 20-25 cm above base and 3-13 cm below BECKER, R.T. & HousE, M.R., 1997, Sea-level changes in che Upper top ofbed, which is 37 cm thick. The lowest occurrence Devonian of che Canning Basin, Western Australia. In House, of Palmatolepis sp. B defines the lower boundary of M.R. & Ziegler, W. (eds.), On sea-level fluctuations in che Zone 11 (Kfapper, 1989). Note that this is the lowest Devonian: Courier Forschungsinstitur Senckenberg, 199: 129- occurrence of Ancyrognathus triangularis in our 146. sampling. The basal5 cm ofBed Q is a layered, rather BELKA, Z., KAuFMANN , B. & BuLTYN CK, P., 1997, Conodont-based quantitative biostratigraphy forche Eifelian of che eastern Anti- dark bioclastic (crinoidal) limestone with spariti c sheet Adas,Morocco: Geologica! Society of America Bulletin, 109: cracks, without macrofauna and conodonts. The main 643-651. part of the bed is a gray and partly red (haematitic) BISCHOFF, G., 1956, Oberdevonische Conodonten (co Iù) aus dem very fossiliferous micritic limestone, with goniatites, Rheinischen Schiefergebirge: Notizblatt cles Hessischen Landesamtes ftir Bodenforschung, 84: 115-137,3 pls. orthocones, bivalves, and some gastropods. BISCHOFF, G. & ZIEGLER, W., 1957, Die Conodontenchronologie Bed P, lower 8 cm of 24 cm thick bed. The lowest cles Mitteldevons und cles tiefsten Oberdevons: Abhandlungen 346 G. KLAPPER, R. T. BECKER

undiag- upmos. ZONE 5 6 7 8 lO 11 Il n ostie 8to9 BED u T s s i. b. R R R-Q Q Q up. 7, 20-25 3-13 INTERVAL 6 12 low. 14 up. 4 lO 13 up. 10 a. b. b.t. M. ovalis x M. asymmetrica x R? M. sp. x A g. ancyrognathoideus x

Pa. gutta x

An. africana x P. timanicus x I . sp. . x P. sp. x x

P a. transitans x x P a. l. linguiformis gamma R R

An. gigas form l x x x

Pa. punctata x x x x x x x P. webbi x x P. decorosus cf. cf. cf. cf. x x P a. spinata x An. gigas form 2 x Oz. bidentatiformis x P. dubius x x x P. aff. P. dengleri x x x Ag. primus x x P a. bohemica x x Oz. trepta x x x l. symmetricus x x x x x x An. eu rva t a early form x x x x Oz. nonaginta x x A g. amplicavus x x P a. aff. Pa. proversa ? ? x A g. coeni x x x An. gigas form 3 x x x A g. barba x P a. mucronata x x P a. ljaschenkoae x x A g. tsiensi x x Pa. plana x Pa. proversa x P a. aff. Pa. winchelli x P a. sp. B x x Pa. jamieae cf.

An. nodosa of authors x x A g. triangularis x An. curvata late form x P. politus x I. alternatus x

T ab. 2 - For explication see next page. COMPARISON OF FRASNIAN CONODONT ZONATIONS 347

ZONE 12 12 12 12 12 13 13 13 13 13-FA.

BED p p o N N M M L L K K K

INTERVAL 0-8 8-18 up.IO low.ll up.5 low.5-6 up.IO low.IO up.8 low.G 6-10 up.IO

Pa. sp. B x An. ioides x x x Pa. winchelli x x x x x An. nodosa of authors x x x x x x x x l. alternatus x x x x x x x P. webbi x x x x x x x Pa. sp. x x A g. triangularis x x x x x x x Pa. aff. Pa. winchelli x x x x x P. sp. inder. x x x x An. nodosa -An. ioides x x x x An. curvata early form x x P. n. sp. BT x x l. symmetricus x x x Ag. amana x x An. curvata late form x x x x x Pa. bogartensis x x x x x Pa. bassi s.s. x P. decorosus x ? Pa. rhenana x x x A g. asymmetricus x x x x Pa. boogaardi x x Pa. linguiformis x x x A g. ubiquitus x Mehlina sp. x Be/ode/la sp. x Pa. triangularis x Pa. minuta minuta x Pa. tenuipunctata x l. iowaensis x

Tabs. 2 and 3- Distribution of conodonts at Martenberg near Adorf (Ziegler, 1958; 1971; Ziegler & Sandberg, 1990). The pro file we have collected is that of the north side. Zone numbers are those of the Frasnian wnation fìrst developed in the Monragne No ire. Bed letters are shown in Text-fìgure 2 for the north side column; intervals are in cm. Abbreviations: low. = lower, up. = upper, upmos. = uppermost, a.b. = above base, b.t. = below top, i.b. = immediately below, FA. = Famennian, R = reworked. Pa. = Palmatolepis, P. = Polygnathus, Ag. =Ancyrognathus, An. =Ancyrodella, Oz. = Ozarkodina, I. = Icriodus, M. = Mesota.xis. Remarks o n some wnal assignments are given in the Appendix.

des Hessischen Landesamtes fiir Bodenforschung, 22: 136 pp., succession and biofacies in upper Frasnian formations 21 pls. (Devonian) from the southern and centrai parts of the Dinant BooGAARD, M. VAN DEN & KuHRY, B., 1979, Statistica! Synclinorium (Belgium) - (Timing of facies shifting and reconstruction of the Palmatolepis apparatus (Late Devonian correlation with late Frasnian events): Bulletin de l'Institut conodonrophorids) at the generic, subgeneric, and specifìc Royal des Sciences Naturelles de Belgique, Sciences de la leve!: Scripta Geologica, 49: 57 pp. Terre, 68: 25-75, 8 pls. BRANSON, E. B. & MEHL, M. G., 1934, Conodonts from the Grassy HousE, M.R. & ZIEGLER, W, 1977, The goniatite and conodont Creek Shale of Missouri: University of Missouri Srudies, 8: sequences in the early Upper Devonian at Adorf, Germany: 171-259, 9 pls. (imprint 1933). Geologica et Palaeontologica, 11: 69-108, 6 pls. BuLTYNCK, P., 1970, Révision stratigraphique et paléontologique }1, Q., 1986, Conodonts. In Hou, H ., Ji, Q., Xian, S. & Wang, J. (Brachiopodes et Conodontes) de la coupe type du Couvinien: (eds.), Middle-Upper Devonian boundary in Maanshan of Mémoires de l'Institut Géologique de l'Université de Louvain, Xiangzhou, Guangxi: Beijing, Geological Publishing House,: 26: 152 pp., 39 pls. 18-50, 113-115. BuLTYNCK, P., HELSEN, S. & HAYDUCKIEWICH, J., 1998, Conodont KlRCHGASSER, WT., 1994, Early morphorypes of Ancyrodella 348 G. KLAPPER, R. T BECKER

rotundiloba a t the Middle-Upper Devonian boundary, Genesee OVNATANOVA, N.S., 1969, Novye verkhnedevonskie konodonry Formation, west-central New York.In Landing, E. (ed.), Studies tsentral'nykh rayonov Russkoy Platformy i Timana: in stratigraphy and paleontology in honor ofDonald W Fisher: Vsesoyuznyy Nauchno-Issledovatel'skiy Geologorazvedochnyy New York State Museum Bulletin, 481: 117-134, 3 pls. Neftyanoy Institut, Trudy, Vypusk 93: 139-141, l p!. KiAPPER, G., 1989, The Montagne Noire Frasnian (Upper Devonian) OVNATANOVA, N.S., 1976, Novye pozdnedevonskie konodonry conodont succession. In McMillan, N .J ., Embry, A. F. & Glass, Russkoy Platformy: Paleontologischeskiy Zhurnal, 1976 (2): D.J. (eds.), Devonian of the World: Canadian Society of 106-115, l pl. Perroleum Geologists, Memoir 14, vol. III: 449-468, 4 pls. PuCHKOV, V.N., KlAPPER, G. & MAsHKOVA, T.V., 1982, Natura! (imprint 1988). assemblages of Palmatolepis from the Upper Devonian of the KiAPPER, G., 1990, Frasnian species of the Late Devonian conodont northern Urals: Acta Pafaeontologica Polonica, 26: 281-298 genus Ancyrognathus: Journal of Paleontology, 64: 998-1025. (imprint 1981). KLAPPER, G., 1997, Graphic correlation of Frasnian (Upper SANDBERG, C.A., ZIEGLER, W, DREESEN, R. & BUTLER, J .L., 1988, Devonian) sequences in Montagne Noire, France, and western Late Frasnian mass extinction: Conodont evenr stratigraphy, Canada. In Klapper, G., Murphy, M.A. & Talent, J.A. (eds.), global changes, and possible causes: Courier Forschungsinstitut Paleozoic sequence stratigraphy, biostratigraphy, and Senckenberg, 102: 263-307, 2 pls. biogeography: Studies in honor ofJ. Granville ("Jess") Johnson: SANDBERG, C.A., ZIEGLER, W, DREESEN, R. & BUTLER, J .L., 1992, Geological Sociery of America Special Paper, 321: 113-129. Conodont biochronology, biofacies, taxonomy, and event KlAPPER, G. & BECKER, R.T., 1998, Comparison of Frasnian stratigraphy around Middle Frasnian Lion mudmound (F2h), (Upper Devonian) conodont zonations. In Bagnoli, G. (ed.), Frasnes, Belgium: Courier Forschungsinstitut Senckenberg, ECOS VII, Seventh international conodont symposium held 150, 87 pp., 10 pls. in Europe, Abstracts: 53-54. SANNEMANN, D., 1955a, Beitrag zur Untergliederung des KlAPPER, G. & FosTER, C.T., Jr., 1986, Quantification of outlines Oberdevons nach Conodonren: Neues Jahrbuch fiir Geologie in Frasnian (Upper Devonian) platform conodonts: Canadian und Palaonrologie, Abhandlungen, l 00: 324-331, l p!. Journal ofEarth Sciences, 23: 1214-1222, 2 pls. SANNEMANN, D., 1955b, Oberdevonische Conodonren (ro Ila): KlAPPER, G. & FosTER, C.T., Jr., 1993, Shape analysis ofFrasnian Senckenbergiana lethaea, 36: 123-156, 6 pls. species of the Late Devonian conodont genus Palmatolepis: ScHùLKE, I., 1997, Conodonr clusters an d multielement Paleontological Sociery Memoir 32 Qournal of Paleontology, reconstructions from the Upper Kellwasser horizon a t La Serre 67 (4), supplement): 35 pp. (Late Frasnian, Montagne Noire, sourhern France): Geologica KlAPPER, G. & LANE, H.R., 1989, Frasnian (Upper Devonian) et Palaeontologica, 31, 37-66, 5 pls. conodont sequence a t Luscar Mountain and Mount Haultain, SHAW, A.B., 1964, Time in stratigraphy: 365 pp., McGraw-Hill, Alberta Rocky Mountains.In McMillan, N.J., Embry, A.F. & In c. Glass, D.J. (eds.), Devonian of the World: Canadian Sociery STAUFFER, C. R., 1938, Conodonts of the Olentangy Shale: Journal ofPerroleum Geologists, Memoir 14, vol. III: 469-478, 2 pls. ofPaleontology, 12:411-433, 6 pls. (imprint 1988). YouNGQUIST, WL., 1945, Upper Devonian conodonts from the KlAPPER, G., KiRCHGASSER, WT. & BAESEMANN,J.F., 1995, Graphic Independence Shale (?) oflowa: Journal of Paleontology, 19: correlation of a Frasnian (Upper Devonian) composite 355-367, 3 pls. standard. In Mann, K.O, & Lane, H.R. (eds.), Graphic ZIEGLER, W., 1958, Conodonrenfeinstratigraphische Unrer- Correlation: Sociery for Sedimentary Geology (SEPM), Special suchungen an der Grenze Mirreldevon/Oberdevon und in der Publication, 53: 177-184. Adorfstufe: Notizblatt des Hessischen Landesamtes fiir KlAPPER, G., Kuz'MIN, A.V. & 0VNATANOVA, N.S., 1996, Upper Bodenforschung, 87: 7-77, 12 pls. Devonian conodonts from the Timan-Pechora regio n, Russia, ZIEGLER, W., 1971, A field trip guidebook; post-symposium and correlation with a Frasnian composite standard: Journal excursion, Sept. 15-18, 1971 ro Rhenish Siate Mountains and ofPaleonrology, 70: 131-152. Harrz Mountains: Philipps-University of Marburg, KRALICK, J .A., 1994, The conodont gen us Ancyrodella in the mi dd! e Symposium on conodonr taxonomy: 47 pp. Genesee Formation (Lower Upper Devonian, Frasnian), ZIEGLER, W. (ed.), 1991, Catalogue of Conodonrs, V. E. western New York: Journal of Paleontology, 68: 1384-1395. Schweizerbart'sche Verlagsbuchhandlung, Stuttgart: 212 pp., LANGE, F.-G., 1968, Conodonten-Gruppenfunde aus Kalken des 13 pls. tieferen Oberdevon: Geologica et Palaeontologica, 2: 37-57, ZIEGLER, W & SANDBERG, C.A., 1990, The Late Devonian standard 6 pls. conodont zonation: Courier Forschungsinstitut Senckenberg, McLEAN, R.A. & KLArrER, G., 1998, Biosrratigraphy of Frasnian 121: 115 pp., 17 pls. (Upper Devonian) strata in western Canada, based on ZIEGLER, W. & SANDBERG, C.A., 1994, Conodont phylogenetic- conodonts and rugose corals: Bulletin of Canadian Petroleum zone concept: Newsletters on Stratigraphy, 30: 105-123. Geology, 46: 515-563, 2 pls. MILLER, A.K. &YouNGQUIST, W, 1947, Conodonts from the rype section of the Sweetland Creek Shale in Iowa: Journal of (manuscript received December 29, 1998 Paleontology, 21: 501-517, 4 pls. acceptedjune 15, 1999) MouRAVIEFF, A. N., 1970, Conodontes du Frasnien de la Belgique: Biostratigraphie et aspects écologiques: Unpublished Ph.D. disserration, Université Catholique de Louvain: 141 pp., 8 Gilberr KlAPPER pls. Deparrmenr of Geology, Universiry of Iowa MùLLER, K.J., 1956, Zur Kennrnis der Conodonten-Fauna des Iowa Ciry, Iowa 52242, U.S.A europaischen Devons, l. Die Garrung Palmatolepis: Abhand- e-mail: [email protected] lungen der Senckenbergischen Naturforschenden Gesellschaft, 494: 70 pp., 11 pls. MùLLER, K.J. & MùLLER, E.M., 1957, Early Upper Devonian R. Thomas BECKER (Independence) conodonts from Iowa, part I: Journal of Institut fiir Palaonrologie, Museum fiir Naturkunde Paleontology, 31: l 069-1108, 8 pls. Humboldt Universitat, MuRPHY, M.A., 1977, On time-stratigraphic units: Journal of Invalidenstrasse 43, D-10115 Berlin, Germany Paleontology, 51:213-219. e-mail: [email protected]