New Data on the Middle Devonian Bryozoans of ...

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In Systême Silurien du Centre de la Bohême par Joachim Bar- rande. Recherches Paleontologiques, 8(1), 230 pp., Prague. Schlüter, C., 1889, Anthozoen des ...
New Data on the Middle Devonian Bryozoans of Germany Andrej Ernst1 Abstract This study presents new data on taxonomic composition, distribution and paleobiogeographic relations of the Middle Devonian bryozoans in Germany. The compilation is performed on the material from ten representative localities of the Eifelian and Givetian age in the Rhenish Slate Massif. The bryozoan fauna contains many genera previously not known from the Devonian in Germany as well as in Europe. They show connections to the Middle Devonian of North America Eastern Americas Realm (EAR) and Paleotethys. An analysis of the fauna reveals that it takes an intermediate position between EAR and Paleotethyan provinces.

Introduction The insufficient taxonomic data presently available for bryozoan faunas of the Eifel erroneously support the impression that bryozoans are only rarely represented in benthic faunas of this region. This is definitively wrong as bryozoans are common in the Devonian deposits of the Eifel, but they have been neglected for many decades, even though the first taxonomic studies started with occasional descriptions during the early 19th century. The first descriptions of fossils from the Eifel included bryozoan specimens, which were also recognized as an integral part of the fauna (Steininger 1853; Goldfuss 1826-33; Schlüter 1889). However, following these early investigations, little data was added during the 20th century, a consequence of the immense diversity of other, stratigraphically more relevant and accordingly more “important” fossil groups. The development of the detailed shelf stratigraphy of the Eifel is generally based on brachiopods (mainly atrypids) and to some extent on trilobites (Struve 1996a-b), which are therefore rather well known in comparison to other common faunal elements. A few publications were made during the last century on Devonian Bryozoa from Germany (Solle 1937, 1952, 1968; Kräusel 1953, 1957, 1981; Toots 1951; Langer 1980), but continuous work on the group was not carried out. Moreover, Solle (1937, 1952, 1968) focused mostly on hederellids which are most certainly not bryozoans (Taylor and Wilson, this volume).

Institut für Geowissenschaften der Christian-Albrechts-Universität zu Kiel, Ludewig-Meyn-Str. 10, D-24118 Kiel, Germany. . 1

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Bryozoa are also common components (10-15%) in biodetrital limestones (arenitic limestones) of the Upper Emsian Heisdorf Formation, upper Nohn Formation (Lower Eifelian) and Cürten Formation (Givetian). They are, however, fragmented due to high energy conditions, and in some cases are even impregnated with hematite or limonite. But even within such deposits, a remarkably high diversity is recognizable and common faunal elements include fenestellids, encrusting fistuliporids and cryptostomes, as their colonies were to some extent able to resist the high mechanical stress of reworking in turbulent waters (Utescher 1992). The present paper delivers an overview of the taxonomic composition, distribution and paleobiogeography of bryozoan faunas from the Middle Devonian of Germany, using material from the most representative localities in the Rhenish Slate Massif.

Distribution and taxonomic composition of bryozoan faunas in the Middle Devonian of Germany Material for this investigation comes mainly from 10 locations in the Rhenish Slate Massif, SW-Germany (Fig. 1). The stratigraphic nomenclature of the Middle Devonian of the Eifel carbonate synclines follows not Steininger and Piller (1999) but Struve (1961, 1992). The terms “Formation,” “Subformation” and “Member” are considered as independent and hierarchic (Bohatý 2005, 2006). Approximately 250 oriented thin sections were made, which contain the following bryozoans:

and Schroeder, 2007). Biodetrital rudstones containing a rich fauna of brachiopods, corals, echinoderms, and the following 19 species of bryozoans: Eridopora orbiculata (Kiepura 1973), Corynotrypa basiplata Kiepura, 1973, Leioclema passitabulatum Duncan, 1939, Leptotrypa prima (Duncan 1939), Eostenopora primiformis Duncan, 1939, Nematopora tenuis sp. n., Acanthoclema distilum Bigey, 1988, Intrapora germanica Ernst and Schroeder, 2007, Prolixicella bifurcata Ernst and Schroeder (in press), Bashkirella cf. devonica (Dessilly 1967), Fenestella spp. (3 species), Spinofenestella antiqua (Goldfuss, 1826), Alternifenestella spp. (2 species), Hemitrypa mimicra McKinney and Kříž, 1986, Reteporidra inflata Bigey, 1988, and Reteporidra sp. Loc. 5 –Western Gerolstein, Auberg, Gerolstein syncline, Flesten Member of the Ahrdorf Formation (Middle Eifelian). Muddy limestone containing corals, echinoderms, brachiopods, and the following bryozoans: Eostenopora globosa (Goldfuss 1826) (Figures 3.3-4), Eridopora orbiculata (Kiepura 1973), Fistulipora sp. and indeterminable fenestrates. Loc. 6–Southwestern development area of the village Gondelsheim, Prüm syncline. Nims Member of the Upper Junckenberg Formation (Eifelian). Muddy limestone containing corals, brachiopods, echinoderms and the following three species of bryozoans: Cyclotrypa cyclostoma (Schlütter, 1889), Fistuliphragma sp. (Figures 2.3-4), and Eostenopora clivosa (Schlütter, 1889). Loc. 7–“Weinberg” quarry near Kerpen, Bohnert Member of the Freilingen Formation (Upper Eifelian). Packstones containing brachiopods, echinoderms and the following four species of bryozoans: Fistuliphragma sp., Streblotrypella sp., Penniretepora sp., and an indeterminable bifoliate cryptostome species. Loc.8–Üxheim-Ahütte, Müllertchen quarry, Hillersheim syncline, Olifant Member of the Ahbach Formation (Upper Eifelian) and Müllert Subformation of the Ahbach-Formation (Lowermost Givetian). Muddy limestone containing sponges, corals, crinoids, brachiopods, and the following 8 species of bryozoans: Cliotrypa sp. (Figures 2.5-7), Leioclema sp., Intrapora sp., Acanthoclema sp., Hemitrypa sp., Reteporidra sp., Fenestrapora transcaucasica Morozova and Lavrentjeva, 1998, and indeterminable rhabdomesine cryptostomes. Loc. 9 –Weinsheim industrial area near “Niesenberg,” Prüm syncline, Upper Loogh Formation (Lower Givetian) containing the following >4 species of bryozoans: Sulcoretepora cf. obliqua McNair, 1937 (Figures 2.8-9), Acanthoclema sp., Hemitrypa sp., and trepostome bryozoans. Loc. 10 – Brühlborn near Rommersheim, Prüm syncline, Upper Nims Member of the Junckenberg Formation (Eifelian). Rudstones and bindstones containing brachiopods, corals, echinoderms and the following 11 species of bryozoans: Fistulipora sp., Eridopora orbiculata (Kiepura, 1973), Cyclotrypa cyclostoma (Schlütter, 1889), Cliotrypa sp., Eostenopora clivosa (Schlütter, 1889), Leioclema sp., Microcampylus sp., Cuneatopora sp., Acanthoclema sp., Bashkirella devonica (Dessilly, 1967), Fenestrapora transcaucasica Morozova and Lavrentjeva, 1998 (Figures 3.8-9), Hemitrypa sp., and Spinofenestella sp.

Figure 1. Middle Devonian bryozoan localities in the Rhenish Mountains (Germany). (������������������������������������������������������������� 1) Blankenheim, (2) Drolshagen, (3) Lüdenscheid, (4) Freilingen, (5) Gerolstein, (6) Gondelsheim, (7) Kerpen, (8) Üxheim-Ahütte, (9) Weinsheim, (10) Brühlborn (map sketch created at http://www.aquarius. geomar.de/omc/make_map.html).

Loc. 1–Railway cut west of Blankenheim, Blankenheim syncline, Junkerberg Formation, Grauberg Subformation, Nims Member (Eifelian). Silty limestone with a rich fauna of brachiopods and corals, and the following 14 species of bryozoans: Fistulipora pachycysta (Bigey 1988), Fistulipora spp. (2 species; Figures 2.1-2), Canutrypa francqana Bassler, 1952, Eridotrypellidae spp. (2 species), Leioclema sp., Anomalotoechus spp. (2 species), Intrapora sp. (Figures 3.5-7), Acanthoclema distilum Bigey, 1988, Bashkirella devonica (Dessilly 1967), ? Rectifenestella sp., Reteporidra sp. Loc. 2–Drolshagen, Middle Devonian (Eifelian). Silty shales, strongly weathered, containing brachiopods and the following four species of bryozoans: Filites bohemicus Barrande in Počta, 1894, Reteporidra sp., Intrapora sp., and indeterminable fenestrates. Loc. 3–Lüdenscheid, Honsel and Werdohl Groups (Givetian). Silty limestones with a rich fauna of brachiopods, corals, echinoderms and the following 19 species of bryozoans: Fistulipora sp., Eridopora sp., Canutrypa francqana Bassler, 1952, ? Leptotrypa sp., ? Leptotrypella sp., Eostenopora primiformis Duncan, 1939, Eridotrypellidae spp. (2 species), Acanthoclema distilum Bigey, 1988, Intrapora sp., Bashkirella devonica (Dessilly 1967), Prolixicella bifurcata Ernst and Schroeder, 2007, Hemitrypa sp., Hemitrypa cf. mimicra McKinney and Kříž, 1986, Reteporidra sp., Reteporina sp., Spinofenestella antiqua (Goldfuss 1826), ? Spinofenestella sp., and ? Speotrypa sp. Loc. 4–Freilingen, Cürten Formation (Givetian) (Ernst

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Figure 2. Cystoporate bryozoans from the Middle Devonian of Germany. (1, 2) Fistulipora sp.��������������������������������������������������������� , Nims Member of the Grauberg Subformation of the Junkerberg Formation (Eifelian), loc. 1. (1) longitudinal section; (2) tangential section, scale bars = 0.5 mm. (3, 4) Fistuliphragma sp., Nims Member of the upper Junckenberg Formation (Eifelian), loc. 6. (3) tangential section; (4) longitudinal section showing hemiphragms, scale bars = 0.5 mm. (5, 6, 7) Cliotrypa sp. Olifant Member of the Ahbach Formation (Upper Eifelian), loc. 8. (5) longitudinal section, scale bar = 1 mm; (6) tangential section, scale bar = 0.5 mm; (7) transverse section, scale bar = 1 mm. (8, 9) Sulcoretepora cf. obliqua McNair, 1937, Upper Loogh Formation (Lower Givetian), loc. 9. Colony surface views, scale bar for 8 = 1 mm and for 9 = 0.5 mm.

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Figure 3. Trepostome, cryptostome and fenestrate bryozoans from the Middle Devonian of Germany. (1, 2) Undescribed trepostome genus, Honsel Group (Givetian), loc. 3. (1) longitudinal section; (2) tangential section, scale bars = 0.2 mm. (3, 4) Eostenopora globosa (Goldfuss 1826). Flesten Member of the Ahrdorf Formation (Middle Eifelian), loc. 5. (3) tangential section, scale bar = 0.5 mm; (4) longitudinal section, scale bar = 1 mm. (5, 6, 7) Intrapora sp. Nims Member of the Grauberg-Subformation of the Junkerberg Formation (Eifelian), loc. 1. (5) transverse section; (6) longitudinal section; (7) tangential section, scale bars = 0.5 mm. (8, 9) Fenestrapora transcaucasica Morozova and Lavrentjeva, 1998. Upper Nims Member of the Junkenberg Formation (Eifelian), loc. 10. (8) tangential section showing autozooecia and “aviculomorphs”, scale bar = 1 mm; (9) “aviculomorphs” on the reverse side of the colony, scale bar = 0.5 mm.

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The encrusting cystoporate Fistuliphragma sp. (Figures 2.3-4) from locality 6 builds tubular colonies and possesses thick hemiphragms. Similar hemiphragms are also known in the ramose genus Cliotrypa Ulrich and Bassler in Bassler, 1929 (Figures 2.5-7). A similar species also occurs in the Givetian of Morocco (unpublished data). Another unusual cystoporate is Canutrypa francqana Bassler, 1952. This species was detected at localities 1 (Eifelian) and 3 (Givetian). Canutrypa francqana produces usually ramose colonies. However, it occurs as encrusting sheets at investigated localities. Such a colony form of Canutrypa francqana was mentioned also by Bigey (1991: 29, plate 1, figs. 9-11) from the Ferques Formation (Frasnian) of Boulonnais, northern France. This bryozoan contains hemispherical cyst-like structures in its autozooecia. The bifoliate cystoporate Sulcoretepora, previously unknown from Europe, was detected at locality 9 (Lower Givetian; Figures 2.8-9). Trepostome bryozoans also possess previously unknown combinations of morphologic characters. An encrusting eridotrypellid bryozoan (Figs. 3.1-2) possesses hemiphragms and cystiphragms. Another species is ramose with thick hemiphragms and basal diaphragms (localities 1 and 3). The genus Eostenopora Duncan, 1939 is reported for the first time from the Devonian of Germany with three species occurring in the Eifelian and Givetian (localities 4-6 and 10). Cryptostome bryozoans are represented by branched rhabdomesine and bifoliate species, belonging mostly to the genus Intrapora Hall, 1883. The rhabdomesine Acanthoclema Hall, 1886 (apparently several species) is the most ubiquitous genus occurring at all localities except 2 and 5-7. Bifoliate Intrapora is represented by four species and occurs abundantly both in the Eifelian and Givetian (localities 1, 4, 8 and 10; Figures 3.5-7). Finally, fenestrate bryozoans occur at the largest number of localities. The most ubiquitous genera are Reteporidra and Hemitrypa. Fenestrapora was found at localities 8 and 10. Other common genera are Spinofenestella and Alternifenestella.

Paleobiogeographic relations of Devonian bryozoans in Germany

Upper Devonian (Frasnian and Famennian) of China and Kazakhstan (Nekhoroshev 1960, Troitzkaya 1968, 1979, Xia 1997), and two species of Sulcoretepora are known from the Devonian of China (Liu 1980, Xia 1997). The connection to the Traverse Group of EAR is also confirmed by three trepostome species which were identified from the Cürten Formation (Givetian): Leioclema passitabulatum Duncan, 1939, Leptotrypa prima (Duncan 1939), and Eostenopora primiformis Duncan, 1939 (Ernst and Schroeder 2007). The genus Fenestrapora Hall, 1885 is known with several species from the Middle Devonian of North America. However one species, Fenestrapora transcaucasica Morozova and Lavrentjeva, 1998, was also discovered in the Eifelian of the Caucasus. This species was identified from locality 10 (Figures 3.8-9). The genus Fenestrapora is distinguished by the presence of special structures called “aviculomorphs” (McKinney 1998). This distribution suggests a transitional character for Middle Devonian bryozoan faunas in Germany (as well as in Europe). Whereas North American species tend to be older in age (mostly Eifelian), Asiatic species of the same genera are younger (Frasnian to Famennian). European faunas seem to assume an intermediate position in age (Eifelian to Givetian).

Summary Middle Devonian bryozoans from Germany presented here indicate higher diversity than previously known. Bryozoans are mainly associated with biodetrital and silty limestones exposed in the Rhenish Slate Massif. Bryozoan diversity varies from 3 to 20 species per locality. Bryozoan faunas display paleobiogeographic affinities with the EAR, on one hand, and with China and Kazakhstan, on the other. However, these connections are not completely contemporaneous, so that bryozoan faunas of the Middle Devonian of Germany are intermediate in space and time between North American and Paleotethyan provinces.

Acknowledgments

The bryozoan fauna from the Middle Devonian of Germany shows various paleobiogeographic relations. Most of genera and several species are present in contemporaneous strata elsewhere in Europe. However, some others show more distant connections. The Devonian species of Intrapora and Sulcoretepora are known mostly from the Middle Devonian of EAR and China. McNair (1937) described several species of both genera from the Traverse Group, roughly corresponding to the Eifelian. Several Intrapora species are also known from the

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I would like to thank to J. Bohatý, M. Grigo and S. Schröder (all University of Cologne) and R. Leunissen (Wollersheim) for providing of material and assistance in the field. J. Bohatý made helpful comments to the present manuscript. The present study was accomplished during the project ER 278/4-1 supported by the Deutsche Forschungsgemeinschaft (DFG). This paper is a contribution to IGCP 499 “Devonian land-sea interaction: evolution of ecosystems and climate”. M. Key (Carlisle), F. Bigey (Paris) and J. Pachut (Indianapolis) are thanked for their helpful and constructive reviews.

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