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Tarphycerida, Nautiloidea ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Denisia Jahr/Year: 2014 Band/Volume: 0032 Autor(en)/Author(s): King Andy H. Artikel/Article: Taxonomic review of early Darriwilian estonioceratids (Tarphycerida, Nautiloidea) from Sweden, Estonia, and the ‘Diluvium-GeschiebeÂ’ of northern Germany and Poland 47-57 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Taxonomic review of early Darriwilian estonioceratids (Tarphycerida, Nautiloidea) from Sweden, Estonia, and the ‘Diluvium-Geschiebe’ of northern Germany and Poland A.H. KING Abstract: Estonioceratid nautiloids are a common and attractive component of mid-Ordovician (early Darriwilian) cephalopod faunas in Baltoscandia. Their distinctive conchs are typically loosely coiled, and vary from forms which possess an apically coiled portion with divergent final whorl, to open gyrocones and even torticones. This paper briefly reviews the taxonomy of several mainly Swedish and Estonian estonioceratid genera which were formerly poorly known at the time of the publication of the Tarphycerida section of the ‘Nautiloid Treatise Part K’ (FURNISH & GLENISTER 1964) or have been described since. Keywords: Systematics, Estonioceratidae, Kundan, Aserian Introduction REMELÉ, 1886, Tragoceras REMELÉ, 1890, Eichwaldoceras BALASHOV, 1955, Bentoceras STUMBUR, 1962 and Ase- Fossil tarphycerid nautiloids are widespread and rioceras STUMBUR, 1962. The torticonic genus Pakrioce- commonly encountered within the Mid Ordovician ras was extensively described by STUMBUR & MUTVEI, ‘Orthoceratite Limestone’ facies of Baltoscandia, espe- 1983, and is only briefly summarised here for reasons of cially Sweden and Estonia, and coeval glacial erratic completeness. boulders (‘Diluvium-Geschiebe’) of northern Germany and Poland. The majority of forms that occur within Revision of a number of these genera is long over- the Kundan and Aserian Stages (early Darriwilian) be- due. Within the Tarphycerida section of the ‘Nautiloid long to the family Estonioceratidae. Their loosely coi- Treatise Part K’ (FURNISH & GLENISTER, 1964, pp. led gyroconic shells are very distinctive and in Sweden K357-K359) Falcilituites was regarded as synonymous these nautiloids are referred to as ‘böjda Cephalopoda’ with Estonioceras; Eichwaldoceras was “Poorly known. meaning literally ‘bent Cephalopoda’. Apparently like Tragoceras” and Tragoceras was conside- red “similar to Aphetoceras and possibly synonymous”. In common with other groups of cephalopods, the Although Bentoceras and Aserioceras were erected befo- preservation of tarphycerids within the ‘Orthoceratite re publication of the Treatise Part K, they were omitted Limestone’ facies is generally very good. Although ma- from the volume. The re-examination of type material ny of the sequences are highly condensed, the fine-grai- along with study of additional specimens from Sweden ned limestones often enable detail of external shell or- and Estonia now enables the status of all these genera nament to be discerned and many conchs are relatively to be re-assessed, and provisional lists of their constitu- uncrushed facilitating examination of whorl cross-secti- ent species to be presented here. A more thorough re- ons and early apical stages. Occasionally limestone vision of the Baltoscandian estonioceratids, including beds are phosphatised, which permits extremely fine assessment of the range of intraspecific variation and detail of the siphuncle wall and structure of connecting the description of new taxa, is currently in preparation rings to be observed (MUTVEI & DUNCA, 2011). by the author. This paper briefly reviews the status and taxonomy of estonioceratid nautiloids from the early Darriwilian Denisia 32, Stratigraphy zugleich Kataloge des Stage of Baltoscandia. The taxa covered include Esto- oberösterreichischen nioceras HYATT in ZITTEL, 1900 along with five previo- Within Baltoscandia, the estonioceratid nautiloids Landesmuseums Neue Serie 157 (2014): usly little known or obscure genera, namely Falcilituites are mainly found within limestones assigned to the ear- 47-57 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Table 1: Biostratigraphical correlation of the Baltoscandian !! "!! ! Stages/Substages with trilobite, graptolite !!! "!! ! and conodont zonation schemes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able 2: Character matrix table and stratigraphic occurrence of Baltoscandian estonioceratid genera. ly Darriwilian Kundan or Aserian stages. The correlati- Taxonomy on of these stages (and their substages) with standard In this section and figure captions, the following ab- international graptolite and conodont zonation sche- breviations are used when referring to material or muse- mes is provided in Table 1. The Kundan and Aserian um collections: stages (represented in Sweden by the Holen Limestone and Segerstad Limestone formations respectively) equa- GIT – Institute of Geology, University of Tallinn, Esto- te approximately to the Didymograptus ‘bifidus’ (Undulo- nia graptus austrodentatus to Nicholsongraptus fasiculata) and RM – Museum of Natural History, (Naturhistoriska lower Didymograptus murchisoni (Pterograptus elegans) Riksmuseet), Stockholm, Sweden graptolite zones; and the upper part of the Lenodus varia- MNH – Museum of Natural History, London, UK bilis to near top of the Eoplacognathus suecicus conodont MfN – Museum für Naturkunde, Berlin, Germany zones. Megistaspid, asaphid and illaenid trilobites are al- so useful locally within the condensed Swedish and Es- tonian ‘Orthoceratite Limestone’ sequences to facilitate correlation. 48 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Order Tarphycerida Genus Estonioceras NOETLING, 1883 (Fig. 1A–D) FLOWER in FLOWER & KUMMEL, 1950 Remeleceras HYATT, 1894 Family Estonioceratidae HYATT in ZITTEL, 1900 Type species: Lituites lamellosus HISINGER, 1837 [by original de- Diagnosis: Smooth or ribbed gyroconic to loosely signation; NOETLING 1883: 275] coiled, rarely torticonic tarphyceracones, typically with Diagnosis: Moderately expanding, relatively stout divergent last whorl. Whorl section depressed to com- conch, early whorls in contact, with depressed or fusi- pressed, dorsal impression absent or only weakly develo- form section and typically with umbilical perforation; ped. Siphuncle ventral to subventral. Septal necks or- last whorl divergent, slightly depressed to circular in thochoanitic, connecting rings thick. Growth lines and section. Conch narrowly to widely camerate, sutures raised lirae form conspicuous broad, shallow to deep hy- straight, transverse. Siphuncle subventral, septal necks ponomic sinus (adapted from FURNISH & GLENISTER short, orthochoanitic; connecting rings very thick and 1964: K357). layered. External ornament indistinct, consisting of fine Discussion: Following FURNISH & GLENISTER growth lines which form an inconspicuous, very broad (1964), the Estonioceratidae is retained here as a sepa- ventral sinus. rate family and distinguished from the similar Tarphyce- Discussion: Estonioceras is a common tarphycerid in ratidae (HYATT, 1894) on the general basis of conch the Kundan Stage of Baltoscandia; its conch form and form (more tarphyceraconic in the Tarphyceratidae) expansion rate readily distinguish it from other estonio- and for tarphyceratids to possess an impressed area in ceratid genera. The differences with Falcilituites are no- the dorsal region of the whorl, related to tightness of ted under that genus below. coiling. The Trocholitidae SCHRÖDER, 1891 differs from both the Tarphyceratidae and Estonioceratidae in pos- HYATT (1894: 525) erected Remeleceras for forms sessing a dorsal siphuncle. closely allied to Estonioceras but differing in possessing a depressed, elliptical whorl section with deeper dorsal Estonioceratid faunas in Sweden, Estonia and Rus- impressions and an “extraordinary form of annular mus- sia, and from coeval glacial erratic boulders in northern cle”. Examination by the present author of material be- Germany and Poland have much in common. All taxa longing to the type species of Remeleceras, Estonioceras possess similar siphuncular features (orthochoanitic sep- impressum (HYATT, 1894, also figured by MUTVEI 1957: tal necks and thickened connecting rings) and distincti- pl. 2) failed to provide sufficient evidence for distinguis- on at generic and specific levels tends to reply upon hing the two genera. Some species of Estonioceras, nota- overall conch form and expansion rate, whorl section, bly E. perforatum SCHRÖDER, 1891, also exhibit a very position of the siphuncle, shape of the suture and type shallow, weak but conspicuous impressed area. The ot- of external
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