Taxonomy and Paleobiogeography of the Ordovician ...

ACTA UNIVERSITATIS CAROLINAE - GEOLOGICA 1999. 43 (1/2)

Taxonomy and Paleobiogeography of the Ordovician Clisospiridae and Onychochilidae (MoUusca) J. FRYDA", D. M . ROHR^' 1) Czech Geological Survey, Kldrov 3/131, 118 21 Praha 1, Czech Republic; e-mail: [email protected] 2) Department of Geology, Sul Ross State University, Alpine, Texas 79832; e-mail: [email protected]

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INTRODUCTION

uppermost part of the Klabava Formation of the Prague Basin, as the type species of Mimospira Koken in Pemer, Opinions on the higher taxonomic position of the families 1925. The latter genus is knovra from Ordovician Clisospiridae Miller and Onychochilidae Koken as well as gastropod faunas in many parts of world. Two species of on their relationships have often been changed. Wenz Mimospira were found in the Ordovician rocks of the (1938) placed the Clisospiridae as a family of the Prague Basin: Mimospira helmhackeri in the Lower and Trochonematacea. On the other hand, Onychochilus was lower Middle Ordovician strata (Fryda and Rohr, in prep.) placed as the sole genus in its own subfamily within the and the younger species Mimospira barrandei Fryda, 1989 family Subulitidae. Both groups were thus regarded as in the uppermost part of the Kraluv Dvur Formation unrelated taxa within the Archaeogastropoda. This (Kralodvor, "Ashgillian"). Species closely related to the opinion was later followed by Knight et al. (1960), but in genotype of Mimospira were recorded from Arenig age contrast to Wenz (1938), they considered onychochylids to strata of Wales (being assigned to Matherella (?) be dextral hyperstrophic gastropods like the macluritids. acuticostata Bates, 1963; Fryda and Rohr, in prep.) and On the other hand, clisospirids were interpreted as from the Late Ordovician ("Caradocian"?) Lower sinistral orthostrophic gastropods with an uncertain higher Edinburg Formation of Virginia (Wangberg-Eriksson taxonomic position. Discoveries of many new clisospirid 1979). and onychochilid taxa have revealed that both families The majority of species of Mimospira occur in the (Clisospiridae and Onychochilidae) are closely related Balto-Scandian region where we know hitherto at least (Horny 1964; Golikov and Starobogatov 1975; Peel 1986; seven species of this genus. The oldest species {Mimospira Wangberg-Eriksson 1979). atava (Moberg and Segerberg, 1906)) is poorly known, Dzik (1983) on the basis of his study of early shell and comes from the upper Tremadocian Ceratopyge Beds ontogeny of the genus Mimospira proposed to separate (Oelandian). Mimospira similis Wangberg-Eriksson, both the Clisospiridae and Onychochilidae from the 1979, (Kalla Limestone and lowermost Dalby Limestone, suborder Macluritina and established a new suborder Viruan) and Mimospira tenuistriata Wangberg-Eriksson, Mimospirina for them. On the other hand, Linsley and 1979, (Kalla, Furudal and lowermost Dalby limestones, Kier (1984) proposed to unite the Onychochiloidea Viruan) are characteristic species for the Middle and early (including Clisospiridae and Onychochilidae), Late Ordovician of Scandian region. The Late Ordovician Macluritoidea, and possibly the Euomphaloidea in the new Mimospira kallholniensis Wangberg-Eriksson, 1979, is order Hyperstrophina of the new class Paragastropoda. knovra hitherto only from its type area (Boda Limestone of The latter class united untorted molluscs. However, quite Dalarna; Harjuan). Two additional species of Mimospira different early shell ontogeny of members of the are known from the Late Ordovician of Estonia; Onychochiloidea and Euomphaloidea (Dzik 1983; Fryda Mimospira puhmuense Isakar and Peel, 1997, (Rakvere 1989, 1995; Bandel and Fryda 1998; Fryda and Rohr, in Stage of Viruan) and unnamed Mimospira species prep.) suggests that the class Paragastropoda is an (Kukruse Stage of Viruan) described under the name artificial group with no zoological validity. Clisospira aff rugosa by Opik (1930). Dzik (1983) In this short paper the stratigraphic ranges of the recorded the occurrence of additional unnamed species of Ordovician Clisospiridae and Onychochilidae are Mimospira from late Middle and early Late Ordovician of summarized as well as some aspects of their Poland. The imperfectly known Middle Ordovician paleogeographic distribution are discussed. Clisospira ingrica Koken, 1897, from the Kunda Stage of the St. Peterburg area may also belong to the genus Mimospira as suggested by Wangberg-Eriksson (1979). FAMILY CLISOSPIRIDAE The Ordovician Clisospira balclatchiensis Longstaff, 1924, from the Balclatchie Group of Girvan (Scotland) is Mimospira transferred to the genus Mimospira (Fryda and Rohr, in Knight (1937) chose the early Middle Ordovician prep.) Onychochilus helmhackeri Pemer, 1900, from the

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Clisospira

The genus Clisospira Billings, 1865, is based on the Ordovician ("Arenigian"?) Clisospira curiosa Billings, 1865, from the "Beekmantovm", latest Early Ordovician age (Whittington 1953), of Quebec (Canada). The Late Ordovician ("Ashgillian") Clisospira reticulata Longstaff, 1924, from the Drummock Group of Girvan (Scotland) resembles the type species in its reticulate ornamentation. The only knovm shell of Clisospira bassleri Raymond, 1908, from the Chazy Formation (Middle Ordovician) of Vermont is imperfectly preserved and so its genus level position is uncertain. Also, Clisospira occidentalis Whitfield, 1877, from the Ordovician (Mohawkian) of Wisconsin is based on internal molds. An undescribed species of Clisospira with distinct collabral shell ornamentation was recently found in the Whiterockian age Shallow Bay Formation of the Cow Head Group, Newfoundland (Rohr, unpubl. data).

Scandian region, the genus Laeogyra is also represented by Laeogyra arvetensis Wangberg-Eriksson, 1979 from the Upper Ordovician Boda Limestone (Dalarna) and by three species, Laeogyra volhynica, Laeogyra alta, and Laeogyra sp., described by Hynda (1986) from the Middle Ordovician of the south-western part of European Platform. As suggested by Wangberg-Eriksson (1979) Clisospira reinwaldi Opik, 1930, from the Kukruse Stage (Viruan) of Estonia may also belong to the genus Laeogyra. Rohr (1996) described an occurrence of lowspired, sinistrally coiled shell in the Middle Ordovician (Whiterockian) of Nevada which resembles the genus Laeogyra. Antispira

The genus Antispira Perner, 1903, was based on Antispira praecox Barrande in Perner, 1903, from the Late Ordovician (middle Beroun, "Caradocian") of the Prague As early mentioned by Wangberg-Eriksson (1979) and Basin. The genus Antispira has been recently emended Peel (1986) the shell of the Upper Ordovician (Harjuan) (Fryda and Rohr, in prep.) as a taxon resembling tlie Clisospira rugosa Koken, 1897 from the Red Jonstorp genus Laeogyra but having a cancellate shell Formation of Ostergotland (Sweden) is ornamented only ornamentation. Besides the type species, it includes the by distinct collabral ribs in contrast to the type species of Balto-Scandian Antispira gracilis (Koken, 1897) from the Clisospira. Thus, the genus-level position of Clisospira Upper Ordovician Boda Limestone (Dalarna); Antispira rugosa is uncertain. Also, the position of two additional ferrigena (Perner, 1911) from the early Late Ordovician Baltic species from the Late Ordovician (Ashgilian) (Beroun, "Caradocian") of the Prague Basin and Antispira ascribed to Clisospira, Clisospira christianiae Koken in n. sp. from the late Late Ordovician (Kosov, "Ashgillian") Pemer, 1925 (Norway) and Clisospira borkholrniensis of the Prague Basin. Invertospira Horny, 1964, is a poorly Koken in Perner, 1925 (Porkuni Stage of Estonia), is still knovra genus similar to Antispira. The type species of this uncertain because of insufficient knowledge of their shell genus, Invertospira lamellifera Horny, 1964 from tlie Middle Ordovician (Sarka Formation, Darriwilian) of the characters. Prague Basin, is based on a single specimen. We tentatively place Invertospira, as well as Pervertina Ferrogyra The genus Ferrogyra Horny, 1964, is based on Clisospira Horny, 1964, in synonymy with Antispira. antiqua Perner, 1903, from the Upper Ordovician (Beroun, "Caradocian") of the Prague Basin. The shape of the shell resembles that of Laeogyra and Versispira, but the last whorl bears basal lamella. These lamellae have the same shape as those in the type species of Mimospira. Up to now Ferrogyra is knovra only from the Prague Basin (Vinice, Zahofany and Bohdalec fms).

Versispira

Perner (1911) placed four Late Ordovician species in his new subgenus Scaevogyra (Versispira): S. (V.) bohemica Barrande in Perner, 1907, S. (V.) ferrigena Perner, 1911, S. (V.) rugosa Perner, 1911, and S. (V.) contraria Barrande in Perner, 1903, from the Prague Basin. The last species was later selected as the type species of Versispira. Unfortunately, this species was based on a single steinkern Undospira from the early Late Ordovician (Letna Formation, Beroun, Wangberg-Eriksson (1979) established the genus "Caradocian") of the Prague Basin and so the knowledge Undospira for her new species Undospira striata from the of its shell character is very limited. In addition, its lowMiddle Ordovician limestones (Kalla and Furudal spired shell as well as that of S. (V) bohemica show no limestones) of Sweden. Besides this species, she important feature distinguishing them from the Late tentatively placed two additional species into Undospira: Cambrian genus Scaevogyra. The remaining two species, tlie Late Ordovician Undospira turrita (Koken in Perner, S. (V.) ferrigena and S. (V.) rugosa Perner, 1911, were 1925) and fragmentary specimens from the Gigas recently moved to Antispira and Laeogyra respectively Limestone. (Fryda and Rohr, in prep.). FAMILY ONYCHOCHILIDAE Helicotis Laeogyra

The type species of Laeogyra Perner, 1903, is L. bohemica Barrande in Perner, 1903, from the Middle Ordovician (Sarka Formation, Darriwilian) of the Prague Basin. Beside tliis species, Laeogyra rugosa (Perner, 1911) from the early Late Ordovician (Vinice Formation, Beroun, "Caradocian") of the same area is knovra. In the Balto-

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Helicotis Koken in Perner, 1925, is an imperfectly known genus with only one knovra species, Helicotis rufiger (Koken, 1897). This species was based on a single incomplete specimen which was lost (Wangberg-Eriksson 1979, p. 4). It occurs in the Lower Ordovician Lower Grey Orthoceras Limestone of the Sm0land area (Sweden). Judging from the photo of Knight (1941, pi. 90, fig. 7),

this species may represent a species of Mimospira or Laeogyra.

Bodospira, Angulospira and Tapinogyra, is typical for Late Ordovician time.

Avalonia The Avalonia microplate include areas of Europe (Wales, England, southeastern Ireland, the Ardennes and the Rhenohercynian zone) as well as of North America (coastal New England, Nova Scotia and the Avalon Peninsula of Newfoundland). Paleogeographic reconsfructions for the Early Ordovician time (Torsvik and Trench 1991; Tait et al. 1997) have suggested that Angulospira Avalonia microplate was at a high paleolatitude (about Angulospira Wangberg-Eriksson, 1979, is closely related 60° south) close to Gondwana. During Arenig/Llanvim to the genus Bodospira. The type species, Angulospira times the Avalonia microplate started its northward drift striata Wangberg-Eriksson, 1979, occurs together with from Gondwana. In Late Ordovician times the Avalonia species of Bodospira in the Upper Ordovician Boda microplate was located close to Baltica in temperate Limestone (Dalarna). A single specimen of an unnamed paleolatitudes (Tait et al. 1997). species of Angulospira (Mimospira of Rohr, 1996) is The occurrence of species belonging to the family known from the Middle Ordovician (Whiterockian) of Onychochilidae is recorded in the "Arenig" of Wales Nevada. (Bates, 1963; Fryda and Rohr, in prep.), and in the Upper Ordovician Lower Edinburg Formation of Virginia Matherellina (Wangberg-Eriksson 1979). Both of these species of The genus Matherellina Kobayashi, 1937 is based on Mimospira seem to be closely related to Mimospira Matherella walcotti Kobayashi, 1933 and is known only helmhackeri (Pemer, 1900) from the early Middle from the Tremadocian Wanjuanshu Formation of the Ordovician of the Prague Basin (Perunica). Sichuan Province (China). Armorica terrane asseblage Tapinogyra During the last 20 years the term Armorica has been used The higher taxonomic position of the poorly known genus in slightly different ways (e.g. Havlicek et al. 1994; Tait et Tapinogyra is still uncertain (Fryda and Rohr, in prep.). al. 1997). Generally, Armorica unites the various terranes The only know species, Tapinogyra glaphyra Wangberg- with Cadomian basement situated within the Variscan and Eriksson, 1979, comes from the Upper Ordovician Boda Alpine fold belts of Europe. Young (1990) suggested that Limestone (Dalarna) and is based on only one specimen. the Armorica microplate was elongated north/south with latitudinal range of about 20° between Iberia and the more PALEOBIOGEOGRAPHY northerly Bohemia. In contrast to this, Havlicek et al. (1994) on the basis of detailed comparison of brachiopod Laurentia and ttilobite assemblages of Baltica, Gondwana, Avalonia The North American paleocontinent (including also and Armorica s.l. demonsfrated that the Ordovician Northern Ireland and Scotland) was situated near the faunas of Bohemia had an independent evolution within paleoequator during Ordovician tune. The clisospirid Armorica s. 1. For this reason, Havlicek et al. (1994) fauna is characterized by the occurrence of Clisospira established a term Perunica (=Tepla-Barrandian terrane) species with a reticulate shell ornament (Clisospira for the microplate occurring in proximity of Baltica during curiosa from the "Beekmantown" of Quebec (Canada) and Cambrian/Ordovician time. This position of Perunica also Clisospira reticulata from the Drummock Group of seems to be consistent with paleomagnetic data (EJ:S et al. Girvan (Scotland)). Unusual occurrences of the Baltic 1986, 1987; but see also Tait et al. 1997). During Middle genera Angulospira and Laeogyral in the Middle Ordovician time, Perunica was separated maximally from Ordovician (Whiterockian) of Nevada need a Baltica by the Tomquist's Sea. According to Havlicek et paleogeographic evaluation. al. (1994), gradual closing of Tomquist's Sea during the Late Ordovician again enabled faunal exchange between Baltica Perunica and Baltica. Different drift histories of IberoDuring Ordovician time the Baltica paleocontinent had Armorica and Perunica were recently suggested by Cocks drifted northwards from high southern latitudes rotating (1998) and Franke (1998). anti-clockwise (Torsvik et al. 1992). The high biodiversity Up to now we have no evidence for existence of any of Ordovician clisospirids and onychochilids is a typical feature of Baltic fauna. The Early Ordovician may be Ordovician clisospirids and onychochilids in the Iberocharacterized by the genera Mimospira, Undospira and Armorica area. In conttast to this, these faunal elements imperfectly knovra Helicotis. During the Middle are relatively conunon in Ordovician sttata of the Prague Ordovician we know the occurrence of several species of Basin (Perunica). In Lower Ordovician sttata, one species Mimospira, Undospira and Laeogyra. The increase of of the Baltic genus Mimospira was found. Mimospira clisospirid and onychochilid species, belonging to genera survived until the early Middle Ordovician (the uppermost Mimospira, Clisospira, Undospira, Laeogyra, Antispira, part of Klabava Formation). The late Middle Ordovician (Darriwilian) is characterized by species of Laeogyra and Bodospira

Bodospira Wangberg-Eriksson, 1979, is known only from the Balto-Scandian region and is represented by two species; Bodospira undulata Wangberg-Eriksson, 1979, and Bodospira kallhooniensis Wangberg-Eriksson, 1979. Both species are from the Upper Ordovician Boda Limestone (Dalarna).

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Invertospira {=Antispira). Absence of both clisospirids and onychochilids is typical for Middle/Upper Ordovician strata (Dobrotiva and Liben fms.) of the Prague Basin. Relatively high numbers of the clisospirid and onychochilid species belonging to genera Ferrogyra, Laeogyra, Antispira, and Mimospira are known from Upper Ordovician strata. The last three genera are also known to occur in Upper Ordovician strata of the Baltica paleocontiment. CONCLUSIONS The stratigraphic ranges of the Ordovician members of the Clisospiridae and Onychochilidae as well as of their paleogeographic distribution suggest: 1. The presence of common genera (Mimospira, Laeogyra and Antispira) in the Ordovician strata of Baltica and Perunica indicate the possibility of faunal exchange between these two areas. On the other hand, the absence of any clisospirids and onychochilids in IberoArmorica and Gondwana indicate the unusual position of the Perunica microplate within the Mediterranean Province. These facts seem to be in contrast with model of Tait et al. (1997) placing the Tepla-Barrandian (= Perunica) block closer to Gondwana than the rest of Armorica s.l. 2. The stratigraphic ranges of clisospirids and onychochilids in the Prague Basin (repeated occurrence of Mimospira species in Early-early Middle and Late Ordovician times; absence of clisospirids and onychochilids in Middle/Upper Ordovician strata) as well as an increase in the species number during Late Ordovician time fits well with the model of Havlicek et al. (1994). REFERENCES Bandel K . , Fryda J., 1998: Position of Euomphalidae in the system of the Gastropoda. Senck. Lethaea 78(1/2): 103-131. Bates D.E.B., 1963: A Lower Ordovician gastropod from Anglesey. Geol. Mag. 100: 258-259. Billings E. 1865: Paleozoic fossils. Can. Geol. Surv. 186-187:420. Cocks L . R . M . , 1998: Faunal integrity of the terranes surrounding Gondwana. Schr. Staatl. Mus. Min. Geol. Dresden 9: 115. Dzik J. 1982: Larval development and relationships of Mimospira; a presumably hyperstrophic Ordovician gastropod. Geol. Foer. Stockh. Foerh. 104(3): 231-239. Franke W., 1998: Tectonic and plate tectonic units at the North Gondwana Margin: evidence from the Central European Variscides. Schr. Staatl. Mus. Min. Geol. Dresden 9: 132-134. Fryda J. 1989: A new species of Mimospira (Clisospiridae, Gastropoda) from the Late Ordovician of Bohemia. Vestnik ryUG6A(A):Til-2A\. Fryda J. 1995: Shell ontogeny of some Lower Paleozoic gastropods and its significance for higher taxonomy. Twelfth International Malacological Congress, Vigo, pp. 305-306. Golikov A . N . , Starobogatov Y.I. 1975: Systematics of prosobranch gastropods. Ma/aco/. 15(1): 185-232. Havlicek V . , Vanek J., Fatka O., 1994: Perunica microcontinent in the Ordovician (its posij^ion within the Mediterranean Province, series division, benthic and pelagic associations). Sbor. geol. ved 46: 23-56. Horny, R. 1964: New Lower Paleozoic gastropod genera of Bohemia (Mollusca). Cas. Ndr. Muzea, R. pf. 133 (4): 211216.

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