First record of the Early Devonian ammonoid

E

N. Jb. Geol. Paläont. Abh. 282/2 (2016), 201–208Article Stuttgart, November 2016

First record of the Early Devonian ammonoid Teicherticeras from the Eifel (Germany): biogeographic and biostratigraphic importance Christof Übelacker, Ulrich Jansen, and Kenneth De Baets With 4 figures

Abstract: We herein report the discovery of Teicherticeras cf. teicherti in the lower Emsian of the Eifel region (Germany), which is important for the correlation of the neritic Rhenish facies and the hemipelagic Hercynian facies as well as the alignment of regional brachiopod biozonation with global ammonoid biostratigraphy. The specimens are similar in morphology and ornamentation to early Teicherticeras species like T. senior and particularly T. teicherti. The temporal overlap of Anetoceras and Teicherticeras indicates a correlation of the Neichnerberg Formation with the upper part of the Praecursor Zone and lower part of the Barrandei Zone. Interestingly, Teicherticeras is also more widely distributed than Anetoceras, which might be related with its better dispersal capabilities and/ or sampling biases against Anetoceratinae. Key words: Ammonoidea, Teicherticeratidae, Devonian, Emsian, Rhenish Mountains, biostratigraphy, biogeography.

Emsian (Mittmeyer 1982; De Baets et al. 2009, 2013a). We herein report the first find of Teicherticeras. Such finds of taxa typical for hemipelagic Hercynian facies in sections dominated by neritic Rhenish facies are crucial for the correlation of both facies and the alignment of regional zonation schemes with global ones (Jansen et al. 2007; Carls et al. 2008; De Baets et al. 2013a; Jansen 2016)

1. Introduction Early ammonoids are characteristic for the hemipelagic Hercynian facies (Erben 1964b), but they can occasionally also occur in sections containing faunal elements of both neritic Rhenish and hemipelagic Hercynian facies known for instance from the Rhenish Mountains in Belgium and Germany (e.g., De Baets et al. 2009, 2013a, 2013b) or the Dra Valley in Morocco (Jansen et al. 2007; De Baets et al. 2010). Unfortunately, the historical collection of ammonoids without clear reference to stratigraphy and tectonic situation in the Rhenish Mountains (particularly in the Hunsrück Slate) often hampers the usefulness for correlation of biostratigraphic schemes in both facies. An exception are rare ammonoid finds from the Eifel region, which are found in the Rhenish facies (Fuchs 1982). So far these finds have been restricted to the Anetoceratinae (Anetoceras arduennense, rarely Ivoites schindewolfi) in the lower

2. Material and methods A nicely preserved specimen of Teicherticeras described herein derives from brownish-gray siltstones assigned to the Neichnerberg Formation in the Finsterbach valley near Hasborn (N 50° 2.511520’, E 06° 50.846470’; Gauss-Krüger coordinates r 256075 h 554541; topographical mapsheet 1:25,000; no. 5907 Hasborn, locality FU 698) in the Eifel region, part of the Rhenish Mountains (Fig. 1). The same locality has yielded a number of brachiopods, corals and

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DOI: 10.1127/njgpa/2016/0613

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202 C. Übelacker et al.

Fig. 1. Geological map of the Eifel (modified from De Baets et al. 2009) showing lower Emsian ammonoid localities: 1) Steffeshausen, 2) Neuerburg, 3) Zendscheid, 4) Sankt Johann, 5) Auf dem Lauzert, 6) Gemünden (near Daun), 7) Hasborn.

by brachiopod biostratigraphy (Mittmeyer 1982, 2008). The paleogeographic reconstruction was made with the freeware Gplates (version 1.5.0; http://www.gplates.org/) using the standard plate model. The Phanerozoic Plate Motion model (Wright et al. 2013) was used as a rotation model.

few trilobite remains (see below). The fossils are deposited in the Senckenberg Museum in Frankfurt [SMF] and were collected by Günther Fuchs in 1971. The Neichnerberg Formation is traditionally assigned to the Singhofen Substage (Fuchs 1982). The stratigraphic assignment is also supported

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First record of the Early Devonian ammonoid Teicherticeras from the Eifel (Germany) 203

We compiled data on the ammonoid faunas, their respective stratigraphic ages (in ma) and coordinates, which were converted to shape files with QGIS (version 2.8.2). We compiled stratigraphic age on the dacryoconaride zone resolution following the correlations suggested in Becker et al. (2012). The key to descriptions, abbreviations and measurements in the Systematic paleontology section follows De Baets et al. (2013b). Institutional abbreviation: SMF: Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt am Main. The abbreviations for open nomenclature and synonymy list follow the recommendations made by Matthews (1973) and Bengtson (1988).

3. Systematic paleontology Class Cephalopoda Cuvier, 1797 Subclass Ammonoidea Zittel, 1884 Order Agoniatitida Ruzhencev, 1957 Suborder Agoniatitina Ruzhencev, 1957 Superfamily Mimosphinctaceae Erben, 1953 Family Teicherticeratidae Bogoslovsky, 1969 Subfamily Teicherticeratinae Bogoslovsky, 1969

Fig. 2. Teicherticeras cf. teicherti (SMF 60513) from the Neichnerberg Formation near Hasborn. Scale bar: 5cm.

discus: Mimagoniatites discus Erben, 1953: 202, Hasselfelde Limestone, Harz Mountains (Germany). erbeni: Teicherticeras (Convoluticeras) erbeni Bogoslovsky, 1963: 35, Vagran Formation, North Urals (Russia). ?ilanense: Teicherticeras ilanense Shen, 1975: 97, Tangding Formation, Guangxi (China). ?lakhranense: Teicherticeras lakhranense Nikolaeva, 2007, Lower Emsian, Northern Caucasus (Russia). ?lineare: Teicherticeras lineare Shen, 1975: 95, Tangding Formation, Guangxi (China). ?nantanense: Teicherticeras nantanense Shen, 1975: 97, Tangding Formation, Guangxi (China). planum: Teicherticeras planum Bogoslovsky, 1980: 59, Dzhaus Beds, Zeravshan Range (Uzbekistan). senior: Teicherticeras senior Chlupáč & Turek, 1983: 29, Zlíchov Limestone, Bohemia (Czech Republic). teicherti: Teicherticeras teicherti Chlupáč & Turek, 1983: 26 (= replacement name for Gyroceratites desideratus Teichert, 1948; but see also Becker et al. 2010 for an alternative view), Taravale Formation, Victoria (Australia).

Genus Teicherticeras Erben, 1960 Type species: Gyroceratites desideratus Teichert, 1948 (= Teicherticeras teicherti), Taravale Formation, lower Emsian, Victoria (Australia), by subsequent designation by Erben (1960). Not to be confused with homonym Teicherticeras desideratum (Walcott, 1884). Diagnosis: See De Baets et al. (2013b) for an up-to-date diagnosis of the genus. Remarks: We herein follow the definition of the genus Teicherticeras as introduced by Klug (2001) and Korn & Klug (2002) with the exception of “Teicherticeras” primigenitum, which has in the meantime been assigned to Mimosphinctes (De Baets et al. 2013b), and inclusion of the newly erected species Teicherticeras lakhranense (Nikolaeva 2007). The genus of Teicherticeras is in need of further revision (Nikolaeva 2007; Becker et al. 2010; De Baets et al. 2013b), but this falls outside the scope of our study. Our form is similar to the type species (T. teicherti), so its inclusion in Teicherticeras remains warranted, even after revision of the questionably assigned species (marked here with question marks).

Teicherticeras cf. teicherti Chlupáč & Turek, 1983 Fig. 2 non 1884 Goniatites desideratum. – Walcott, p. 203, pl. 17, fig. 10. cf. p. 1948 Gyroceratites desideratus. – Teichert, p. 65, pl. 16, figs. 1, 4-6 (non pl. 16, figs. 2, 3 = Teicherticeras sp. D in Erben 1965, which is currently synonymized with Talenticeras talenti; see Klug 2001). cf. p. 1958 Gyroceratites desideratus. – Teichert & Talent, pp. 12, 13, 17.

Species included (modified from De Baets et al. 2013b): buluti: Convoluticeras buluti Erben,1965: 297, Dede Formation, Bithynia (Turkey). coskuni: Convoluticeras coskuni Erben, 1965: 295, Dede Formation, Bithynia (Turkey). ?desideratum: Goniatites desideratum Walcott, 1884: 203, Lower Nevada Limestone, Nevada (USA).

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Fig. 3. Proposed correlations (modified from De Baets et al. 2013a); time-scale (modified from Kaufmann 2006), stratigraphic correlations (Weddige 1996; Weddige et al. 2005a, 2005b), ranges of some important taxa used for biozonation or discussed in the text are listed.

cf. 1960 Teicherticeras desideratum. – Erben, pp. 63, 65. cf. 1964a Teicherticeras desideratum. – Erben, fig. 9. cf. 1965 Teicherticeras desideratum. – Erben, pp. 289291. cf. 1983 Teicherticeras teicherti. – Chlupáč & Turek, p. 26.

and the simple, rursiradiate ribbing speak for its inclusion in Teicherticeras. Species of Teicherticeras can be roughly grouped based on their morphology and stratigraphic ranges. Our specimen is most similar in coiling and ribbing to older Teicherticeras like T. teicherti and T. senior. Based on coiling and ornamentation, our specimen is most similar to Teicherticeras teicherti, which has been found in the Nothoperbonus conodont Zone (Mawson 1987) – correlative with Praecursor and Barrandei dacryoconaride zones (Becker et al. 2012). Teicherticeras senior has been reported from the upper part of the Praecursor dacryoconarid Zone. The holotype of T. senior was also compressed by compaction, but it has a lower whorl expansion rate and less densely spaced ribbing at similar diameters. Additionally, specimens provisionally assigned to T. senior (Becker & House 1994; De Baets et al. 2010) have a thicker, more rounded whorl profile than T. teicherti and our specimen. Teicherticeras teicherti is still poorly known – only one specimen shows faint remains of the original sculpture, and two specimens (= Teicherticeras sp. D of Erben 1965) originally included in it have subsequently been assigned to Talenticeras by Klug (2001). Furthermore, T. teicherti has so far only been reported from Buchan (Victoria, Australia), which was paleogeographically far away from the Eifel (Fig. 4). As we only have one specimen, which is not that well-preserved, we describe it in open nomenclature. Teicherticeras buluti and T. coskuni have a stronger coiling and distinctly different ornamentation (more widely spaced plications). Furthermore, both are stratigraphically younger than T. teicherti and T. senior, as they are associated with the bactritoid Pseudobactrites bicarinatus and dacryo-

Material: One internal (compound) mould (SMF 60513). Description: Specimen SMF 60513 has an estimated maximum diameter of 70 mm and completes at least 3 whorls. The inner whorls are poorly preserved, but suggest that there is an umbilical window. The specimen is thinly discoidal (but no precise estimate of original whorl width can be made). The specimen has a moderately high whorl expansion rate (WER ~ 2.14) and a moderately wide umbilicus (uw/dm = 0.45) at 70 mm diameter. The specimen is advolute, and no evidence for a whorl overlap could be found. The specimen shows fine, densely spaced, rursiradiate plications, which are strongly bent backwards towards the ventral side and show a rounded hyponomic sinus. The spacing between ribs increases from about 0.8 mm at wh = 16 mm to 1.6 at wh = 24 mm. This corresponds with a rise of the relative rib index RRI of about 7 at wh = 19.5 mm to 11 at wh = 22.5 mm. Comparisons: The specimen is strongly compressed by diagenetic compaction (making it impossible to estimate precisely the original whorl width), but a subtrapezoidal cross section can still be recognized. The moderately high whorl expansion rate, advolute coiling, its cross section shape,

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• Australia (T. teicherti, T. sp. E; cf. Teichert 1948;

conarides Nowakia cf. barrandei and Viriatellina pseudogeinitziana (Haas 1968). Teicherticeras? planum is only weakly ornamented in later ontogeny and also is younger (Bogoslovsky 1980) than T. senior and T. teicherti.

Erben 1965; Chlupáč & Turek 1983; Klug 2001);

• China (T. ilanense, T. lineare, T. nantanense; see

Shen 1975, Ruan 1981, 1996; Yu & Ruan 1988);

• the Czech Republic (Bohemia: T. senior, Chlupáč

& Turek 1983);

4. Discussion

• Germany (Harz and Rhenish mountains: T. discus,

4.1. Implications for ammonoid biostratigraphy and biogeography

Teicherticeras sp.; see Erben 1953, 1960; De Baets et al. 2013b);

So far, Anetoceratinae, predominantly Anetoceras (A. arduennense) as revised by De Baets et al. (2009), are the only early Emsian ammonoids that have been reported from the Eifel region (Mittmeyer 1982, 2008). Although no Anetoceras has so far been reported from the Neichnerberg Formation (upper Singhofen), the taxon has been reported from the underlying Gefell Formation (lower Singhofen and the overlying Stadtfeld Formation (Mittmeyer 1982; De Baets et al. 2009, 2013a). This speaks for at least a temporal overlap of Teicherticeras and Anetoceras during part of the deposition of the Neichnerberg Formation (Fig. 3). The overlapping ranges of both genera in the lower Emsian are also known from the lower Emsian of South China (Yu & Ruan 1988; Ruan 1996), the Anti-Atlas of Morocco (De Baets et al. 2010) and the Barrandian (Chlupáč et al. 1979; Chlupáč & Turek 1983; Chlupáč & Lukeš 1999). Both Teicherticeras and Anetoceras are also known from the Bundenbach/Gemünden area (Erben 1965; De Baets et al. 2009, 2013b), but it is unclear if they derive from the same layers due to historical collection without clear reference to stratigraphy. Anetoceras appears earlier in the fossil record (at least in South China and potentially also in Morocco). Cooccurrence of Anetoceras and Teicherticeras seems to be typical of the upper part of the Praecursor Zone and lowermost part of the Barrandei Zone in places where they are well biostratigraphically constrained in the Czech Republic (Chlupáč et al. 1979; Chlupáč & Turek 1983), Morocco (De Baets et al. 2010; Aboussalam et al. 2015), and South China (Shen 1975; Ruan 1981; Yu & Ruan 1988; Ruan 1996). This also further corroborates that the ammonoid content of the Hunsrück Slate ranges from the Praecursor to Elegans zones (De Baets et al. 2013b). Interestingly, Teicherticeras has a wider paleogeographic distribution during the Praecursor and Barrandei zones than Anetoceras (De Baets et al. 2013c, 2013d); see Fig. 4. The genus Teicherticeras is confidently known from:

• Morocco (Anti-Atlas: T. aff. senior, T. cf. senior,

Becker & House 1994; De Baets et al. 2010);

• Russia (Northern Caucasus: T. lakhranense,

Nikolaeva 2007; North Urals: T. erbeni, Bogoslovsky 1963);

• the United States of America (Nevada: T. desidera-

tum, Walcott 1884; see also House 1965);

• Uzbekistan (Zeravshan-Gissar Range: T. planum,

Bogoslovsky 1980).

Only from Novaya Zemlya, Anetoceras (A. oriens) has been reported to be present in absence of Teicherticeras (Yatskov 1994). However, early ammonoid studies focused on Anetoceratinae (Yatskov 1990), while other taxa are just listed (Yatskov 1994) and not described or figured and have been reported in open nomenclature, making it hard to verify this report. In summary, Anetoceras and Teicherticeras are known to co-occur in sections ranging from the equatorial (South China; point 6 in Fig. 4) to higher latitudes (e.g., Bohemia in the Czech Republic: point 10 in Fig. 4; Anti-Atlas in Morocco: point 9 in Fig. 4) in a wide range of facies (limestone and marl to claystone and siltstone) including neritic sediments (Rhenish facies of the Eifel; point 8 in Fig. 4), so that differences in facies distribution cannot necessarily explain this discrepancy. Teicherticeras has a more strongly coiled conch morphology and a smaller embryonic shell compared with more basal ammonoids (House 1996; Klug & Korn 2004; De Baets et al. 2012, 2013b, Klug et al. 2015), which relate with better swimming abilities and/or more beneficial life history characteristics (higher fecundity) in Teicherticeras than in Anetoceras. However, a collection bias cannot be excluded as Anetoceratinae are more easily fragmented and are often overlooked (Becker et al. 2010; De Baets et al. 2013a, 2013b).

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Fig. 4. Paleogeographic distribution of ammonoid localities in the lower Emsian. 1 = Canada; 2 = Novaya Zemlya, Russia; 3 = Northern Urals, Russia; 4 = Northern Caucasus, Russia; 5 = Zeravshan Range, Uzbekistan; 6 = South China; 7 = Nevada, USA; 8 = Eifel-Hunsrück, Germany; 9 = Tafilalt, Morocco; 10 = Bohemia, Czech Republic; 11 = Pendik, Turkey; 12 = Victoria, Australia.

4.2. Biostratigraphic constraints of the associated fauna

the mid-early Emsian Neichnerberg Formation (upper Singhofen Group) mainly by lithostratigraphic correlation. This further corroborates that at least part of the Hunsrück Slate yielding Mimagoniatites fecundus must be younger than Ulmen-Singhofen age as was already suggested by De Baets et al. (2013a, 2013b).

The locality has also yielded the following taxa: Pleurodictyum problematicum Goldfuss, 1829 Rugosa indet. Arduspirifer latestriatus prolatestriatus Mittmeyer, 1973 Euryspirifer assimilis ssp. (fragments) Tropidoleptus rhenanus Frech, 1897 Plebejochonetes semiradiatus (Sowerby, 1842) Burmeisterella sp. (fragment of cranidium) Diamenocrinus sp. crinoid columnals fish remains (?)

5. Conclusions Based on its morphology, the studied ammonoid specimen from the Eifel is safely assigned to Teicherticeras. An assignment to the species T. teicherti seems most likely. Overlapping ranges of Anetoceras and Teicherticeras in the Eifel can be assumed based on stratigraphic correlation and can be considered typical for the upper Praecursor and lower Barrandei zones. Teicherticeras has a wider paleogeographic distribution than Anetoceras during this interval. This discrepancy cannot easily be attributed to facies controls (Erben 1964b) as both taxa are known to co-occur in a wide range of facies. It could potentially be related with better swimming capabilities or life history strategies reflected in differences in conch morphology and embryonic shell size, or a collection bias putting the more

Most significant with respect to biostratigraphy are the finds of sufficiently preserved Ard. latestriatus prolatestriatus, showing an earliest to mid-early Emsian (Ulmen–Singhofen) age in terms of the Rhenish subdivision (Jansen 2016). This age is supported by Eur. assimilis, which is restricted to nearly the same temporal range, Trop. rhenanus (mid-Siegenian to early Emsian) and the Emsian trilobite Burmeisterella. G. Fuchs (unpublished database) assigned the outcrop to

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Acknowledgements We would like to thank Mihaela-Cristina Krause for introducing us to GPlates. This research also benefitted from a visit to the National Museum (Prague) for comparative purposes supported by the SYNTHESYS Project http://www. synthesys.info/ which is financed by European Community Research Infrastructure Action under the FP7 Integrating Activities Programme (CZ-TAF-4374). We would also like to thank the reviewers Christian Klug (Zürich) and Dieter Korn (Berlin) for their constructive comments on a previous version of this manuscript.

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K enneth De Baets (corresponding author), Christof Übelacker, Fachgruppe PaläoUmwelt, Geozentrum Nordbayern, Friedrich-Alexander Universität Erlangen-Nürnberg, Loewenichstraße 28, 91054 Erlangen, Germany; e-mails: [email protected]; christof.uebelacker@ gmail.com Ulrich Jansen, Senckenberg Forschungsinstitut und Naturmuseum, Senckenberganlage 25, 60325 Frankfurt am Main, Germany; e-mail: [email protected]

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