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Revue de micropalontologie 51 (2008) 3966
Original article
Upper Ordovician microphytoplankton of the Bills Creek Shale and Stonington Formation, Upper Peninsula of Michigan, U.S.A.: Biostratigraphy and paleogeographic signicance Microphytoplancton du Shale de Bills Creek et de la Formation de Stonington (Ordovicien Sup erieur), P eninsule sup erieure du Michigan, Etats-Unis : biostratigraphie et signication pal eog eographique
Reed Wicander a, , Geoffrey Playford b
b a Department of Geology, Central Michigan University, Mount Pleasant, Michigan 48859, USA Department of Earth Sciences, The University of Queensland, Brisbane, Queensland 4072, Australia
Abstract An abundant, diverse, and well-preserved organic-walled microphytoplankton assemblage is described from the Upper Ordovician Bills Creek Shale and the lower Stonington Formation (Bay de Noc Member) in the Upper Peninsula of Michigan, U.S.A. Based on graptolite and conodont evidence, the Bills Creek Shale and Stonington Formation are Richmondian (=Ashgill) in age. The assemblage is dominated by acritarchs, which comprise 29 species (including the enigmatic palynomorph Gloeocapsomorpha prisca) assigned to 20 genera. The prasinophyte phycomata are represented by undifferentiated species of Leiosphaeridia and Tasmanites. In addition, chitinozoans are abundant, and scolecodonts and graptolite fragments are common. Paleontologic-palynologic and sedimentologic evidence indicates that the Bills Creek Shale was deposited in a low-energy, shallow, nearshore marine environment. The overlying Bay de Noc Member of the Stonington Formation also accumulated in a lowenergy, normal marine environment, but in a more offshore, somewhat deeper water setting. Both formations experienced minor transgressive and regressive episodes as indicated by uctuations in the composition of the palynoora. The combined Bills Creek/Stonington acritarch assemblage closely resembles those described from the Richmondian-aged Maquoketa Shale (Missouri and Kansas), Sylvan Shale (Oklahoma), and Vaur eal Formation (Anticosti Island, Qu ebec, Canada). The overall composition of the acritarch assemblage from these four formations reects a distinctive, recognizably Laurentian character. Nonetheless, many of the Bills Creek/Stonington acritarchs have been reported from Upper Ordovician localities elsewhere, providing additional evidence for Late Ordovician cosmopolitanism of the marine microphytoplankton community. Additionally, the restricted stratigraphic range of many of the taxa further enhances their biostratigraphic application, both regionally and globally, and reafrms the Richmondian (=Ashgill) age of the Bills Creek Shale and Stonington Formation. 2007 Elsevier Masson SAS. All rights reserved. R esum e ` paroi organique abondant, diversi Un assemblage de microphytoplancton a e et bien pr eserv e est d ecrit dans lOrdovicien Sup erieur du Shale de Bills Creek et de la Formation Stonington inf erieure (Membre Baie de Noc) dans la P eninsule sup erieure du Michigan, Etats-Unis. Dapr` es le contenu en graptolites et conodontes, le Shale de Bills Creek et la Formation Stonington sont d age Richmondien (=Ashgill). Lassemblage nigmatique Gloeocapsomorpha prisca) r est domin e par des acritarches, comprenant 29 esp` eces (dont le palynomorphe e eparties dans 20 genres. Les prasinophytes phycomata sont repr esent es par des esp` eces indiff erenci ees de Leiosphaeridia et Tasmanites. De plus, les chitinozoaires sont abondants et les fragments de graptolites et scol ecodontes sont communs. Les r esultats s edimentologiques et pal eonto-palynologiques indiquent nergie. Le Membre Baie de Noc de la que le Shale de Bills Creek sest d epos e dans un environnement marin proximal, peu profond et de faible e galement accumul nergie mais dans des eaux plus distales Formation Stonington susjacent sest e e dans un environnement marin normal de faible e pisodes transgressifs et r et plus profondes. Les deux formations ont connu des e egressifs mineurs, indiqu es par les uctuations dans la composition ` acritarches de Bills Creek et Stonington ressemble a ` ceux d de la palynoore. Lassemblage a ecrits dans le Shale de Maquoketa (Missouri et
Corresponding author. E-mail address: reed.wicander@cmich.edu (R. Wicander).
0035-1598/$ see front matter 2007 Elsevier Masson SAS. All rights reserved. doi:10.1016/j.revmic.2007.01.001
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R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966
Kansas), le Shale de Sylvan (Oklahoma) et la Formation Vaur eal ( Ile dAnticosti, Qu ebec, Canada) d age Richmondien. La composition g en erale ` acritarches de ces quatre formations re` de lassemblage a ete un caract` ere Laurentien distinctif et reconnaissable. Cependant, la plupart de ces t acritarches de Bills Creek et Stonington ont e e signal es dans dautres localit es de lOrdovicien Sup erieur, ce qui constitue une preuve suppl ementaire du cosmopolitisme de la communaut e microphytoplanctonique marine de lOrdovicien Sup erieur. De plus, la r epartition stratigraphique restreinte ` la fois au niveau r de la plupart des taxons renforce leur application biostratigraphique a egional et global et r eafrme l age Richmondien (=Ashgill) du Shale de Bills Creek et de la Formation Stonington. 2007 Elsevier Masson SAS. All rights reserved.
Keywords: Acritarchs; Bills Creek Shale; Biostratigraphy; Paleoenvironment; Paleogeography; Stonington Formation; Upper Ordovician Mots cl es : Acritarches ; Shale de Bills Creek ; Biostratigraphie ; Pal eoenvironnement ; Pal eog eographie ; Formation de Stonington ; Ordovicien Sup erieur
1. Introduction Organic-walled microphytoplankton have been widely used for stratigraphic correlation both regionally and globally for several decades, and their utility in dating Proterozoic and Paleozoic marine sedimentary rocks is widely recognized (Playford, 2003). Additionally, these palynomorphs are being applied increasingly in paleoenvironmental studies (Vecoli, 2000; Li et al., 2004), and more recently, as they relate to global biodiversication, and to paleoclimatic and paleogeographic changes (e.g., Servais et al., 2004; Vecoli and Le H eriss e, 2004). Acritarchs and prasinophyte phycomata are the major components of Proterozoic and Paleozoic microphytoplankton communities. Acritarchs (from the Greek akritos, uncertain, confused; and arche, origin) are an informal and undoubtedly polyphyletic group of organic-walled microfossils of unresolved biologic afnities. Most palynologists accept that acritarchs are probably the cysts of various marine microphytoplanktonic groups, and that many of them are pre-dinoagellates (Playford, 2003). The Prasinophyceae are an extant class of marine green algae, whose fossil record consists of the cyst-producing (or phycomata) stage in their life cycle. Acritarchs and prasinophytes were the primary producers of the Proterozoic and Paleozoic oceans. Thus, being at the base of the marine food web, uctuations in their diversity and abundance would, presumably, have inuenced to some extent the evolution of the marine ecosystem. The Ordovician Period was a time when acritarchs experienced their greatest radiation (more than 1500 species are known from the Ordovician) and also mirrored the provincialism of marine invertebrates during the Early and Middle Ordovician, as well as the cosmopolitanism of the Late Ordovician (Vecoli and Le H eriss e, 2004). Although the Lower Ordovician has been more intensively studied palynologically than the Upper Ordovician, a fairly extensive literature exists regarding the taxonomy, biostratigraphy, and paleogeographic distribution of Upper Ordovician organic-walled microphytoplankton taxa. A profuse and varied assemblage of acritarchs and prasinophytes from two Upper Ordovician (Richmondian = Ashgill) localities in Michigans Upper Peninsula is described here, following preliminary observations by Wicander and Playford (1999). The assemblage is compared to three other coeval Laurentian acritarch suites, and the biostratigraphic, paleoenvironmental, and paleogeographic signicance of the assemblage is discussed in both regional and global contexts.
2. Stratigraphy and age 2.1. Stratigraphy The Bills Creek Shale, Stonington Formation, and Big Hill Formation are the three uppermost Ordovician marine lithostratigraphic units exposed successively in the Upper Peninsula of Michigan, northern U.S.A. (Catacosinos et al., 2001). Hussey (1926: p. 121) named the Bills Creek beds (=Bills Creek Shale) for thinly bedded shales and argillaceous limestones exposed along the banks of Bills Creek in Michigans Upper Peninsula and placed them in the Richmond portion of the Cincinnatian series. Hussey (1952: p. 41) later described an outcrop of Bills Creek beds at Haymeadow Creek, also in the Upper Peninsula. However, he noted that the lower Bills Creek beds, exposed at the falls on Haymeadow Creek, may not belong to the Richmond. It now seems best to correlate these lower beds, at least, with the Collingwood [Middle Ordovician]. . . (p. 43). Hussey (1952: p. 45) remarked that these lower beds differed lithologically from the typical Bills Creek beds at Bills Creek in being darker and not weathering to a light gray color. He termed them the Haymeadow Creek Member, which he stated was formerly the basal part of the Bills Creek beds of Richmond age (Hussey, 1952: p. 13). The Haymeadow Creek Member is not formally recognized, the beds exposed at Haymeadow Creek being considered the lowermost portion of the Bills Creek Shale (Catacosinos et al., 2001). The Bills Creek Shale consists of up to 26.8 m of mostly thin-bedded gray shale and calcareous shale, with numerous alternations of shale and argillaceous limestone near its top (Hussey, 1926, 1952). It rests disconformably on limestone of the Gross Quarry Member of the Middle Ordovician Trenton Formation (Catacosinos et al., 2001). A disconformity also separates the Bills Creek Shale from the overlying Stonington Formation, which is divided into two members (Hussey, 1926: p. 132; Catacosinos et al., 2001). The lower Bay de Noc Member comprises ca. 3.1 m of massive grayish-brown argillaceous limestone and gray calcareous shale, succeeded by ca. 4.6 m of alternating, thinly bedded gray argillaceous limestone and gray calcareous shale. The 0.9 to 6.1 m-thick Ogontz Member conformably overlies the Bay de Noc, and consists largely of gray to dark brown, massive and irregularly bedded cherty limestone with some lenses of argillaceous limestone.
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The Big Hill Formation is usually separated from the underlying Stonington Formation by a covered interval. Where exposed, it consists predominantly of gray, argillaceous limestone (Hussey, 1926). Both the Bills Creek Shale and Stonington Formation contain abundant and diverse marine faunas. In general, fossils are plentiful and well-preserved in the limestones, and locally abundant and well-preserved in the shales of the Bills Creek Shale (Hussey, 1926, 1952). The fauna comprises graptolites, crinoids, locally abundant bryozoans, brachiopods, bivalves, and ostracodes, as well as trilobites and Cornulites. The paleontologic and sedimentologic characteristics of the Bills Creek Shale indicate deposition in a low-energy, shallow, normal marine environment. Desiccation cracks, although rare, are preserved in some of the calcareous shale layers, indicating at least occasional aerial exposure. The argillaceous limestone layers may represent more offshore conditions, farther from the terrestrial sources for the shales. An erosional break between the Bills Creek Shale and Stonington Formation is indicated not only by their disconformable relationship, but also by distinct differences in their faunal content. Only a few taxa (e.g., the brachiopod Ranesquina alternata, bivalve Pterinae demissa, and ostracode Tetradella regularis) occur in both formations (Hussey, 1926: p. 130). The Bay de Noc Member of the Stonington Formation contains abundant marine fossils, mainly in the argillaceous limestones. In the Ogontz Member, fossils are most numerous in the cherty limestones and include gastropod and trilobite taxa not found in the Bay de Noc Member. Typical Stonington fossils include crinoids, brachiopods, bivalves, gastropods, trilobites, and Cornulites. Deposition of the Bay de Noc Member was probably in a low-energy, offshore, normal marine environment, with periodic terrigenous input as indicated by the shales. The Ogontz Member was very likely deposited under low-energy, offshore, normal marine conditions. 2.2. Age The latest global Ordovician chronostratigraphy and regional series and stage correlations (Webby et al., 2004) indicate that the base of the Amorphognathus ordovicicus conodont Biozone occurs within the upper part of the Amplexograptus manitoulinensis graptolite Biozone, and thus slightly predates the base of the Dicellograptus complanatus graptolite Biozone (Fig. 1). Furthermore, the base of the A. ordovicicus condodont Biozone is slightly higher than the base of the North American Richmondian stage and the base of the British Ashgill series. Accordingly, the respective bases of the Richmondian and Ashgill are nearly coincident (Webby et al., 2004: Figs. 2.1, 2.2). Goldman and Bergstr om (1997: p. 980981) suggested a possible two-part subdivision of the North American A. manitoulinensis graptolite Biozone, its upper part assigned to the lower portion of the Richmondian stage (Fig. 1). Although not formally proposing this subdivision, they thought it likely to have biostratigraphic utility in the North American Midcontinent successions. Additionally, Goldman and Bergstr om (1997:
p. 981) noted that several Midcontinent successions contain co-occurring conodonts and graptolites, thus facilitating precise correlation between the biozonal schemes based on each of these groups. Particularly important for regional correlation is the fact that Goldman and Bergstr oms (1997) data support the interpretation that the base of the A. ordovicicus conodont Biozone is equivalent to a level within the upper A. manitoulinensis graptolite Biozone. The Stonington Formation was assigned by Goldman and Bergstr om (1997: p. 980) to the D. complanatus graptolite Biozone (Richmondian = Ashgill) and the A. ordovicicus conodont Biozone (Fig. 1). Moreover, Bergstr om (personal communication, 2001) has conrmed that his conodont collections from shore exposures of the Stonington Formation signify a postArnheim Richmondian age. The Arnheim Formation in the type Cincinnati region is placed in the lower Richmondian stage, which equates to the C3 Cincinnati sequence (Goldman and Bergstr om, 1997: p. 983, Text-g. 10). Bergstr om and Mitchell (1986: p. 256) also noted that based on then available graptoloid evidence, particularly the occurrence of Glyptograptus anacanthus, Mitchell and Bergstr om (1977: 261) concluded that the basal Richmondian Arnheim Formation is coeval with the upper A. manitoulinensis Zone in our 1977 study. Thus, the Stonington Formation is attributable to the upperlower to middle Richmondian (=D. complanatus graptolite Biozone). Catacosinos et al. (2001: p. 37) placed the Stonington Formation in the Upper Ordovician, Cincinnatian series, Richmond Group. The underlying Bills Creek Shale is assigned to the upper A. manitoulinensis graptolite Biozone, which is equivalent to the lower Richmondian (Goldman and Bergstr om, 1997: p. 980, Text-g. 8). No conodonts have been described from either the Bills Creek Shales type section or the Haymeadow Creek section. However, conodonts recovered by Votaw (1980a: p. 19) from the upper part of the Bills Creek Shale in the Escanaba region of Michigans Upper Peninsula indicate attribution to the Amorphognathus ordovicicus or possibly the uppermost A. superbus conodont Biozone (lowest Richmondian; Goldman and Bergstr om, 1997: p. 971): Fig. 1. Goldman and Bergstr om (1997: p. 971) re-examined the graptolite fauna of the Haymeadow Creek Shale (=Haymeadow Creek Member of Hussey, 1952; lower Bills Creek Shale of Catacosinos et al., 2001) exposed at Haymeadow Creek and assigned it to the A. manitoulinensis graptolite Biozone. Furthermore, they stated that the Haymeadow Creek section does not contain the typical lower A. manitoulinensis graptolite Biozone fauna [=upper Maysvillian stage] such as is present in the Blue Mountain Formation of Manitoulin Island and in the Lorranine Group siltstones and shales of New York (p. 971). Thus, it appears that the Bills Creek Shale can, with reasonable condence, be placed in the lower Richmondian stage (upper A. manitoulinensis graptolite Biozone). It should be noted that Goldman and Bergstr om (1997: p. 980, Text-g. 8) showed only the upper A. manitoulinensis and D. complanatus graptolite Biozones of the Richmondian in the North American Midcontinent region. According to Bergstr om and Mitchell (1986: p. 259), there are no graptolite data useful for classication of the middle and upper Richmondian in
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Fig. 1. Chronostratigraphic correlation of Richmondian palyniferous formations within the Cincinnatian series of North America. Correlation among the global series, British stages, North American series and stages, graptolite and conodont biozones, and time slices is based on Webby et al. (2004). Radiometric dates based on Sadler and Cooper (2004). Biostratigraphic placement of the palyniferous Richmondian formations within the Ordovician time scale of Webby et al. (2004) based on Jacobson and Achab (1985), Bergstr om and Mitchell (1986), and Goldman and Bergstr om (1997). Fig. 1. Corr elations chronostratigraphiques des formations fossilif` eres d age richmondienne dans les s eries du Cincinnatien en Am erique du Nord. Corr elations entre les tages Britanniques, les s tages dAm ` conodontes : dapr` s eries globales, les e eries et les e erique du Nord, les biozones a es Webby et al. (2004). Datations radiom etriques ` l dapr` es Sadler et Cooper (2004). Position biostratigraphique des formations fossilif` eres d age richmondienne en rapport a echelle chronostratigraphique de Webby et al. (2004) : dapr` es Jacobson et Achab (1985), Bergstr om et Mitchell (1986), et Goldman et Bergstr om (1997).
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terms of graptolite zones but indirect evidence suggests that this interval is coeval with (part of?) the Dicellograptus complanatus Zone, and perhaps the lowermost Climacograptus inuiti Zone. Jacobson and Achab (1985: p. 168, Text-g. 3) showed the Dicellograptus anceps graptolite Biozone of Britain and Scandinavia correlating to a position between the D. complanatus and Amplexograptus inuiti graptolite Biozones of North America. This interval corresponds to the complexus and pacicus graptolite Biozones of Britain and North America (Webby et al., 2004). Therefore, the A. inuiti graptolite Biozone of North America correlates to the Climacograptus extraordinarius graptolite Biozone of Britain and Scandinavia (Jacobson and Achab, 1985), which is approximately equivalent to the Hirnantian stage of Britain and the Gamachian stage of North America (Webby et al., 2004). Fig. 1 thus shows the Sylvan Shale of Oklahoma, Maquoketa Formation of Missouri, and the Stonington Formation and Bills Creek Shale of Michigans Upper Peninsula as occurring in the graptolite and conodont biozones indicated by Goldman and Bergstr om (1997). A dashed line indicates that the upper biostratigraphic extent of those formations within the Richmondian is not known in accordance with the Ordovician timescale of Webby et al. (2004). Placement of the Vaur eal Formation of Anticosti Island, Qu ebec, Canada, is based on Jacobson and Achab (1985). 3. Materials and methods Representative lithologies of the Bills Creek Shale and Stonington Formation were sampled at two locations in Michigans Upper Peninsula (Fig. 2). Eight samples of the Bills Creek Shale (Locality 1) were collected from a beach exposure on the east side of Little Bay de Noc; specically, along the section line between sec. 14 and 23, T. 39 N, R. 22 W, Delta County (Votaw, 1980b: p. 24; Hussey, 1926: p. 126; Hussey, 1952: p. 41). Here, the 2.6 m interval of the Bills Creek Shale (Fig. 3) consists of gray-brown, blocky shale weathering to a bluish-gray aggy shale (samples BC1, 3, 5, 6, 8) alternating with browngray, argillaceous, ne-grained crystalline limestone weathering bluish-gray (samples BC2, 4, 7). Nine samples from a 3.95 m section of the Bay de Noc Member of the Stonington Formation (Locality 2) were collected from a bluff along the east shore of Little Bay de Noc, south of Stonington Community Hall (SE 1/4, sec. 26, T. 39 N, R. 22 W, Delta County; Stop 5 of Votaw, 1980c: p. 56; Hussey, 1926: p. 132). The sampled interval (Fig. 3) comprises the following: 0.65 m of gray, argillaceous, ne- to medium-grained crystalline limestone, with occasional stringers of gray shale (samples S0, 1); 1.8 m of gray to gray-brown, calcareous, blocky shale (samples S2-5); 0.5 m of gray, argillaceous, ne- to medium-grained crystalline limestone, with a few stringers of gray shale (sample S6); 0.5 m of gray, calcareous, blocky shale, weathering graybrown (sample S7); and 0.5 m of gray, argillaceous, ne- to medium-grained crystalline limestone with occasional stringers of bluish-gray, calcareous, blocky shale (sample S8). These are all composite samples, with thicknesses ranging from 1060 cm, designed to provide an overview of acritarch
Fig. 2. Map showing the two sampling localities in Delta County of the Upper Peninsula of Michigan from which eight studied samples of the Bills Creek Shale and nine of the overlying Stonington Formation were collected for this study (Fig. 3). Fig. 2. Localisation des deux sites d echantillonnage dans le Comt e de Delta de la P eninsule sup erieure du Michigan.
diversity and abundance for each lithology. Thus, major changes in acritarch composition between successive lithologies can be ascertained, as well as variations among similar rock types at differing stratigraphic levels (e.g., the thicker portions of the Stonington Formation and Bills Creek Shale). Laboratory preparation of the samples followed the standard palynologic technique of dissolving 2530 g of rock successively in cold HCl, HF, and HNO3 for respective removal of carbonates, silicates, and suldes. Samples were neutralized with distilled H2 O between acid treatments. The resultant residues were ltered through 52 m and 20 m nylon screens, thus yielding >52 m, 2052 m, and <20 m fractions from each sample. Five slides each of the >52 m and 2052 m fractions, and two slides each of the <20 m fraction were prepared using Elvacite as the mounting medium. Three slides each of the >52 m and 2052 m fraction and one slide of the
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Sample S4 (Stonington Formation) was prepared for the picking of selected single specimens and their examination under the scanning electron microscope (SEM). Residues, in absolute ethanol, from the >20 m fraction were allowed to dry on microscope slides where they were then picked using a micromanipulator (Lefngwell and Hodgkin, 1971) and transferred to a circular no. 1 coverslip (12 mm diameter) that was pre-coated with a very thin adhesive. The coverslip was mounted on an aluminum stub and gold-platinum-coated for SEM examination and imaging. Following SEM examination, the coverslip was permanently mounted using Eukitt mounting medium on a microscope slide. Light photomicrographs were taken under Nomarski differential interference contrast illumination using an Olympus BH2 microscope equipped with an automatic photomicrographic system. Specimens were photographed using Kodak black and white 35 mm ISO 400 lm. A JEOL JSM-840A SEM instrument was used for detailed examination and electronic image capture. All illustrated microphytoplankton specimens (Plates 14) are deposited in the Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, U.S.A., and are assigned catalog numbers CMNH 19086 through CMNH 19147 (see Appendix A).
4. Systematic paleontology The vast majority of the palynomorphs encountered in the Bills Creek Shale and Stonington Formation are acritarchs, with minor amounts of leiospheres and tasmanitids, which are attributed to the prasinophycean green algae. The acritarchs recorded hereunder are arranged alphabetically by genera under the informal incertae sedis group name Acritarcha, and are treated as form genera and species in accordance with the provisions of the International Code of Botanical Nomenclature (I.C.B.N.; Greuter et al., 2000). A complete synonymy is not provided for all species. The original binomial name and any subsequent generic transfers are listed; and, where appropriate, a reference to a complete synonymy is cited. Morphologic terminology follows Williams et al. (2000). The dimensions for all species are given, and where there are three numbers, the rst number is the minimum value, the second number in parenthesis is the arithmetic mean, and the third number is the maximum value. The dimensions of each species are followed by the number of specimens measured.
Fig. 3. Stratigraphic sections of the Richmondian (Upper Ordovician) Bills Creek Shale and overlying Stonington Formation, Delta County, Upper Peninsula of Michigan, showing lithofacies and composite intervals sampled for this study. Samples from Locality 1 are prexed BC (=Bills Creek Shale) in text and Fig. 4; samples from Locality 2 are prexed S (=Stonington Formation) in text and Fig. 5. Fig. 3. Sections stratigraphiques des formations de Bills Creek Shale et de Stonington, d age richmondienne (Ordovicien Sup erieur), Comt e de Delta, P eninsule chantillonn sup erieure du Michigan, montrant les lithofacies et les niveaux e es chantillons provenant de la localit au cours de l etude. Les e e 1 sont indiqu es avec le pr exe BC (=Bells Creek Shale) dans le texte et dans la Fig. 4 ; chantillons provenant de la localit les e e 2 sont indiqu es avec le pr exe S (=Stonington Formation) dans le texte et dans la Fig. 5.
<20 m fraction were examined for identication and counting of the individual specimens so as to determine relative abundances of taxa in each sample and for quantitative and qualitative comparisons between samples.
Plate 1. Fig. 1. Tasmanites sp. 400. Figs. 2, 6. Leiosphaeridia sp. 2. 400. 6. 490. Figs. 3, 7. Baltisphaeridium adiastaltum Wicander, Playford and Robertson, 1999. 400. Figs. 4, 8. Baltisphaeridium perclarum Loeblich and Tappan, 1978. 400. Fig. 5. Aremoricanium squarrosum Loeblich and MacAdam, 1971. 400. Figs. 9, 10. Dactylofusa ctenista (Loeblich and Tappan, 1978) Fensome, Williams, Barss, Freeman and Hill, 1990. 400. Figs. 11, 12, 16, 17. Dactylofusa playnetrella (Loeblich and Tappan, 1978) Fensome, Williams, Barss, Freeman, and Hill, 1990. 11, 12. 400. 16. 600. 17. Showing discontinuous nature of ridges, 1200. Figs. 13, 14, 15. Dorsennidium hamii (Loeblich, 1970) Sarjeant and Stancliffe, 1994. 13. 400. 14. 600. 15. 400. Figs. 18, 19. Dorsennidium undosum Wicander, Playford and Robertson, 1999. 400. Figs. 20, 21. Estiastra sp. A. 400. Planche 1. Fig. 1. Tasmanites sp. 400. Figs. 2, 6. Leiosphaeridia sp. 2. 400. 6. 490. Figs. 3, 7. Baltisphaeridium adiastaltum Wicander, Playford et Robertson, 1999. 400. Figs. 4, 8. Baltisphaeridium perclarum Loeblich et Tappan, 1978. 400. Fig. 5. Aremoricanium squarrosum Loeblich et MacAdam, 1971. 400. Figs. 9, 10. Dactylofusa ctenista (Loeblich et Tappan, 1978) Fensome, Williams, Barss, Freeman et Hill, 1990. 400. Figs. 11, 12, 16, 17. Dactylofusa playnetrella (Loeblich et Tappan, 1978Loeblich et Tappan, 1978) Fensome, Williams, Barss, Freeman et Hill, 1990. 11, 12. 400. 16. 600. 17. Montrant la nature discontinue des rides, 1200. Figs. 13, 14, 15. Dorsennidium hamii (Loeblich, 1970) Sarjeant et Stancliffe, 1994. 13. 400. 14. 600. 15. 400. Figs. 18, 19. Dorsennidium undosum Wicander, Playford et Robertson, 1999. 400. Figs. 20, 21. Estiastra sp. A. 400.
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Plate 2. Figs. 1, 2. Excultibrachium concinnum Loeblich and Tappan, 1978. 800. Figs. 3, 4, 7. Hoegklintia sp. cf. H. radicosa (Loeblich, 1970) Jacobson and Achab, 1985. 3. 800. 4. 2000. 7. 2000. Figs. 5, 6. Elektoriskos sp. A. 800. Planche 2. Fig. 1, 2. Excultibrachium concinnum Loeblich et Tappan, 1978. 800. Fig. 3, 4, 7. Hoegklintia sp. cf. H. radicosa (Loeblich, 1970) Jacobson et Achab, 1985. 3. 800. 4. 2000. 7. 2000. Fig. 5, 6. Elektoriskos sp. A. 800.
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Plate 3. Figs. 1, 2. Leiofusa litotes Loeblich and Tappan, 1978. 800. Figs. 3, 4. Lophosphaeridium varum Wicander, Playford and Robertson, 1999. 800. Fig. 5. Lophosphaeridium edenense Loeblich and Tappan, 1978. 800. Fig. 6. Multiplicisphaeridium irregulare Staplin, Jansonius and Pocock, 1965. 800. Figs. 7, 10. Micrhystridium prolixum Wicander, Playford and Robertson, 1999. 800. Figs. 8, 11. Micrhystridium hirticulum Wicander, Playford and Robertson, 1999. 800. Fig. 9. Veryhachium europaeum Stockmans and Willi` ere, 1960. 800. Planche 3. Fig. 1, 2. Leiofusa litotes Loeblich et Tappan, 1978. 800. Fig. 3, 4. Lophosphaeridium varum Wicander, Playford et Robertson, 1999. 800. Fig. 5. Lophosphaeridium edenense Loeblich et Tappan, 1978. 800. Fig. 6. Multiplicisphaeridium irregulare Staplin, Jansonius et Pocock, 1965. 800. Fig. 7, 10. Micrhystridium prolixum Wicander, Playford et Robertson, 1999. 800. Fig. 8, 11. Micrhystridium hirticulum Wicander, Playford et Robertson, 1999. 800. Fig. 9. Veryhachium europaeum Stockmans et Willi` ere, 1960. 800.
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4.1. Prasinophyte phycomata Division CHLOROPHYTA Pascher, 1914 Class PRASINOPHYCEAE Christensen, 1962 Order PYRAMIMONADALES Chadefaud, 1950 Family LEIOSPHAERIDIACEAE Timofeev, 1956 nom. corr. M adler, 1963 Genus Leiosphaeridia Eisenack, 1958a Type species: Leiosphaeridia baltica Eisenack, 1958a; by original designation. Leiosphaeridia spp. Plate 1, Figs. 2 and 6 Measurements: Vesicle diameter 54 (63) 70 m. (5 specimens). Discussion: There are a number of specimens that have a simple, psilate, spherical vesicle and we attribute them, nonspeciated, to Leiosphaeridia. Occurrence: Bills Creek Shale and Stonington Formation (present study). Leiosphaeridia Eisenack, 1958a is a widely occurring and stratigraphically long-ranging genus (Proterozoic-Recent). Order PTEROSPERMATALES Schiller, 1925 Family TASMANITACEAE Sommer, 1956 Genus Tasmanites Newton, 1875 Type species: Tasmanites punctatus Newton, 1875; by original designation. Tasmanites spp. Plate 1, Fig. 1 Measurements: Vesicle diameter 44 m, 132 m, 145 m. (3 specimens). Discussion: The specimens have a simple, psilate, thickwalled, spherical vesicle, and are attributed here, without any attempt at speciation, to Tasmanites. They differ from Leiosphaeridia in being generally larger with a thicker eilyma. Occurrence: Bills Creek Shale and Stonington Formation (present study). Tasmanites Newton, 1875 is a widely occurring and stratigraphically long-ranging genus (ProterozoicRecent).
4.2. Acritarchs Group ACRITARCHA Evitt, 1963 Genus Aremoricanium Deunff, 1955 Type species: Aremoricanium rigaudiae Deunff, 1955; by original designation. Aremoricanium squarrosum Loeblich and MacAdam, 1971 Plate 1, Fig. 5 1971. Aremoricanium squarrosum Loeblich and MacAdam, p. 44, Pl. 18, Figs. 18. 1971. Aremoricanium syringosagis Loeblich and MacAdam, p. 44, Pl. 18, Fig. 9. Measurements: Vesicle diameter 76 m; 12 processes; process length 38+ m; process width at base 78 m; neck length 36 m; neck width 20 m. (1 specimen). Discussion: We follow Playford and Wicander (2006) and others (e.g., Jacobson and Achab, 1985) in accepting that Loeblich and MacAdams (1971) species Aremoricanium squarrosum and A. syringosagis nom. corr. syringosage are conspecic. Occurrence: Stonington Formation (present study). Previously reported from the Edenian (upper Caradoc) of Indiana and Ohio (Loeblich and MacAdam, 1971); Richmondian (Ashgill) of Oklahoma (Loeblich and MacAdam, 1971; Playford and Wicander, 2006) and Anticosti Island, Qu ebec, Canada (Jacobson and Achab, 1985);?CaradocAshgill of Gasp e, Qu ebec, Canada (Martin, 1980); and from coeval strata in Europe, the Middle East, and North Africa (e.g., Molyneux et al., 1996: Pl. 2, Fig. 7; Vavrdov a, 1997: Fig. 3). Genus Baltisphaeridium Eisenack, 1958b ex Eisenack, 1959 emend. Eisenack, 1969 Type species: Baltisphaeridium longispinosum (Eisenack, 1931 ex O. Wetzel, 1933) Eisenack, 1959; by original designation. Discussion: Despite being the repository of more species than any other acritarch genus, Baltisphaeridium Eisenack, 1958b ex Eisenack, 1959 emend. Eisenack, 1969, remains one of the most unsatisfactorily dened acritarch genera (Wicander et al., 1999: p. 5). Because of the continued nomenclatural
Plate 4. Figs. 1, 2, 10. Leiofusa fusiformis (Eisenack, 1934) Eisenack, 1938. 1. Showing ne striations, 400. 2. 400. 10. 325. Fig. 3. Lophosphaeridium acinatum Wicander, Playford and Robertson, 1999. 400. Figs. 4, 11. Gloeocapsomorpha prisca Zalessky, 1917 emend. Foster et al., 1989. 4. 400. 11. 800. Figs. 5, 6. Peteinosphaeridium septuosum Wicander, Playford and Robertson, 1999. 400. Figs. 7, 8. Navifusa sp. A. 400. Fig. 9. Orthosphaeridium rectangulare (Eisenack, 1963) Eisenack, 1968. 400. Figs. 12, 13. Peteinosphaeridium septuosum Wicander, Playford and Robertson, 1999. 12. 600. 13. Showing trilaminate nature of processes, 800. Figs. 1416. Veryhachium oklahomense Loeblich, 1970. 14. 400. 15, 16. Showing psilate nature of eilyma and processes, 1000. Figs. 17, 18. Polygonium gracile Vavrdov a, 1966 emend. Sarjeant and Stancliffe, 1996. 17. 400. 18. 600. Figs. 19, 23. Veryhachium trispinosum (Eisenack, 1938) Stockmans and Willi` ere, 1962 complex. 19. 400. 23. Showing psilate-microgranulate surface of eilyma, 600. Figs. 20, 24. Villosacapsula setosapellicula (Loeblich, 1970) Loeblich and Tappan, 1976. 20. 400. 24. Showing excystment by epityche, 600. Figs. 21, 22. Sylvanidium paucibrachium Loeblich, 1970. 400. Planche 4. Fig. 1, 2, 10. Leiofusa fusiformis (Eisenack, 1934) Eisenack, 1938. 1. Montrant les tr` es nes rides, 400. 2. 400. 10. 325. Fig. 3. Lophosphaeridium acinatum Wicander, Playford et Robertson, 1999. 400. Fig. 4, 11. Gloeocapsomorpha prisca Zalessky, 1917 emend. Foster, Reed et Wicander, 1989. 4. 400. 11. 800. Fig. 5, 6. Peteinosphaeridium septuosum Wicander, Playford et Robertson, 1999. 400. Fig. 7, 8. Navifusa sp. A. 400. Fig. 9. Orthosphaeridium rectangulare (Eisenack, 1963) Eisenack, 1968. 400. Fig. 12, 13. Peteinosphaeridium septuosum Wicander, Playford et Robertson, 1999. 12. 600. 13. Montrant la structure trilaminate des processus, 800. Fig. 1416. Veryhachium oklahomense Loeblich, 1970. 14. 400. 15, 16. Montrant la nature psilate de leilyma et des processus, 1000. Fig. 17, 18. Polygonium gracile Vavrdov a, 1966 emend. Sarjeant et Stancliffe, 1996. 17. 400. 18. 600. Fig. 19, 23. Veryhachium trispinosum (Eisenack, 1938) Stockmans et Willi` ere, 1962 complex . 19. 400. 23. Montrant la surface psilate-microgranulate de leilyma, 600. Fig. 20, 24. Villosacapsula setosapellicula (Loeblich, 1970) Loeblich et Tappan, 1976. 20. 400. 24. Montrant lexcystement par epityche, 600. Fig. 21, 22. Sylvanidium paucibrachium Loeblich, 1970. 400.
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confusion concerning this genus, we follow Wicander et al. (1999) in regarding assignment of the baltisphaerid species herein as only provisional. Baltisphaeridium adiastaltum Wicander, Playford and Robertson, 1999 Plate 1, Figs. 3 and 7 1999. Baltisphaeridium adiastaltum Wicander, Playford and Robertson, p. 5, 7, Fig. 4.64.9. For additional synonymy, see Wicander et al. (1999: p. 5). Measurements: Vesicle diameter 40 (49) 60 m; 1521 processes; process length 21 (34) 50 m; process width at base 2 (3.3) 4 m. (13 specimens). Discussion: For a complete discussion of this species and comparison to similar species, see Wicander et al. (1999: p. 7). Although Baltisphaeridium adiastaltum Wicander, Playford and Robertson, 1999 and B. oligopsakium Loeblich and Tappan, 1978 are morphologically similar, and tend to co-occur, all of our specimens can denitely be attributed to B. adiastaltum. Occurrence: Bills Creek Shale and Stonington Formation (present study). Previously reported from the CaradocAshgill of Kansas (Wright and Myers, 1981); Richmondian (Ashgill) of Missouri (Wicander et al., 1999), Oklahoma (Playford and Wicander, 2006), and Anticosti Island, Qu ebec, Canada (Jacobson and Achab, 1985). Baltisphaeridium perclarum Loeblich and Tappan, 1978 Plate 1, Figs. 4 and 8 1978. Baltisphaeridium perclarum Loeblich and Tappan, p. 1253, Pl. 6, Figs. 57. For additional synonymy, see Wicander et al. (1999: p. 9). Measurements: Vesicle diameter 50 (56) 70 m; 69 processes; process length 60 (63) 70 m; process width at base 6 (8.8) 12 m; plug height 4 m. (7 specimens). Occurrence: Bills Creek Shale and Stonington Formation (present study). Previously reported from the CaradocAshgill of Kansas (Wright and Myers, 1981); Richmondian (Ashgill) of Missouri (Robertson, 1997; Wicander et al., 1999) and Oklahoma (Loeblich and Tappan, 1978; Playford and Wicander, 2006); CaradocAshgill of Gasp e, Qu ebec, Canada (Martin, 1980); Ashgill of northeast Libya (Molyneux, 1988; Hill and Molyneux, 1988); ?LlanvirnAshgill of Iran (Ghavidel-syooki, 2001); and possibly from the CaradocAshgill of Gotland, Sweden (Eiserhardt, 1989) and Estonia (Uutela and Tynni, 1991).
species assigned to Eupoikilofusa by Loeblich and Tappan (1978). Dactylofusa ctenista (Loeblich and Tappan, 1978) Fensome, Williams, Barss, Freeman and Hill, 1990 Plate 1, Figs. 9 and 10 1978. Eupoikilofusa ctenista Loeblich and Tappan, p. 1263, Pl. 8, Figs. 8, 9. 1990. Dactylofusa ctenista (Loeblich and Tappan, 1978) Fensome, Williams, Barss, Freeman and Hill, p. 180. Measurements: Vesicle length 150 (160) 176 m; maximum vesicle width 27 (33) 39 m. (5 specimens). Discussion: Loeblich and Tappan (1978) named four new species of Eupoikilofusa recovered from the Upper Ordovician Sylvan Shale, Oklahoma. All were based on very few specimens (apparently only one specimen in three of the species, and ve specimens in the other). Loeblich and Tappan (1978: p. 1263) differentiated E. ctenista from E. parvuligranosa on the basis of the latter possessing small grana that may be aligned in rows (see also Wright and Myers, 1981: p. 24, Pl. 3, Fig. H). Such a distinction based on so few specimens and the fact that all other morphologic features, including size are the same, appears tenuous. However, all of our specimens are nongranulate and clearly t the circumscription for Dactylofusa ctenista. Occurrence: Bills Creek Shale and Stonington Formation (present study). Previously reported from the Richmondian (Ashgill) of Oklahoma (Loeblich and Tappan, 1978; Playford and Wicander, 2006); Ashgill of northeast Libya (Molyneux, 1988); Ordovician/Silurian boundary of northwest Argentina (Rubinstein and Vaccari, 2004). Dactylofusa platynetrella (Loeblich and Tappan, 1978) Fensome, Williams, Barss, Freeman and Hill, 1990 Plate 1, Figs. 11, 12, 16 and 17 1978. Eupoikilofusa platynetrella Loeblich and Tappan, p. 12631264, Pl. 8, Fig. 10. 1990. Dactylofusa platynetrella (Loeblich and Tappan, 1978) Fensome, Williams, Barss, Freeman and Hill, p. 215. Measurements: Vesicle length 99 (116) 135 m; maximum vesicle width 33 (41) 44 m. (4 specimens). Discussion: Dactylofusa platynetrella (Loeblich and Tappan, 1978) is characterized as having an excentric, thin-walled relatively broad fusiform vesicle ornamented with discontinuous ridges (Loeblich and Tappan, 1978: p. 1263). It is probably conspecic with D. striata (Staplin, Jansonius and Pocock, 1965) Fensome, Williams, Barss, Freeman and Hill, 1990, the only difference seemingly being that the vesicle of D. platynetrella is broader and more asymmetric than D. striata. These features were noted by Loeblich and Tappan (1978: p. 1264) and possibly result from postdepositional compression (Jacobson and Achab, 1985: p. 182). All of our specimens tend to be of the broad and compressed variety as illustrated by the holotype (Loeblich and Tappan, 1978: Pl. 8, Fig. 10) and gured as Eupoikilofusa striata in Jacobson and Achab (1985: Pl. 2, Fig. 2).
Genus Dactylofusa Brito and Santos, 1965 Type species: Dactylofusa maranhensis Brito and Santos, 1965; by original designation. Discussion: The generic categorization of sculptured fusiform acritarchs such as Dactylofusa Brito and Santos, 1965, Poikilofusa Staplin, Jansonius and Pocock, 1965, and Eupoikilofusa Cramer, 1971 is confused and controversial (Fensome et al., 1990; Playford and Wicander, 2006). We follow Fensome et al. (1990) in attributing to Dactylofusa the
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Occurrence: Bills Creek Shale and Stonington Formation (present study). Previously reported from Richmondian (Ashgill) strata of Oklahoma (Loeblich and Tappan, 1978); Anticosti Island, Qu ebec, Canada (Jacobson and Achab, 1985); Ashgill of Algerian Sahara and southern Tunisia (Vecoli, 1999). Genus Dorsennidium Wicander, 1974 emend. Sarjeant and Stancliffe, 1994 Type species: Dorsennidium patulum Wicander, 1974; by original designation. Dorsennidium hamii (Loeblich, 1970) Sarjeant and Stancliffe, 1994 Plate 1, Figs. 13, 14 and 15 1970. Veryhachium hamii Loeblich, p. 741, Fig. 35AF. 1994. Dorsennidium? hamii (Loeblich, 1970) Sarjeant and Stancliffe, p. 40 (assignment provisional). Measurements: Vesicle sides 42 (53) 60 m long side and 20 (31) 40 m short side; 3 major processes in same plane 18 (30) 40 m long, 4 (5.5) 7 m wide at base; 13 supplementary processes 20 (30) 36 m long, 4 (5) 6 m wide at base (13 specimens). Discussion: Although Sarjeant and Stancliffes (1994: p. 40) assignment of Veryhachium hamii Loeblich, 1970 to Dorsennidium Wicander, 1974 was only made provisionally, we regard such assignment as appropriate for the reasons stated by Wicander et al. (1999: p. 11). Occurrence: Bills Creek Shale and Stonington Formation (present study). Previously reported from the CaradocAshgill of Kansas (Wright and Myers, 1981); Richmondian (Ashgill) of Missouri (Miller, 1991; Wicander et al., 1999), Oklahoma (Loeblich, 1970; Playford and Wicander, 2006), and Anticosti Island, Qu ebec, Canada (Jacobson and Achab, 1985); Mohawkian (Caradoc) and Edenian (upper Caradoc) of St. Lawrence Lowland, Qu ebec and southeast Ontario, Canada (Martin, 1983); CaradocAshgill of Czech Republic (Vavrdov a, 1988), Morocco (Elaouad-Debbaj, 1988), and Gotland, Sweden (Eiserhardt, 1992). Dorsennidium undosum Wicander, Playford and Robertson, 1999 Plate 1, Figs. 18 and 19 1981. Veryhachium hamii auct. non Loeblich, 1970. Wright and Meyers, p. 2930, Pl. 3, Figs. J, N, Pl. 8, Fig. G only. 1999. Dorsennidium undosum Wicander, Playford and Robertson, p. 11, 13, Fig. 7.17.4. Measurements: Vesicle diameter 30 (32) 38 m; 610 processes; process length 22 (27) 34 m; process width at base 2 (3.8) 4 m. (5 specimens). Occurrence: Bills Creek Shale and Stonington Formation (present study). Previously reported from the CaradocAshgill of Kansas (Wright and Myers, 1981); Richmondian (Ashgill) of Missouri (Wicander et al., 1999) and Oklahoma (Playford and Wicander, 2006). Li et al. (2006) recorded Dorsennidium cf. D. undosum from the Caradoc of Xinjiang, northwestern China.
Genus Elektoriskos Loeblich, 1970 Type species: Elektoriskos aurora Loeblich, 1970; by original designation. Elektoriskos sp. A Plate 2, Figs. 5 and 6 Description: Vesicle originally spherical, outline subcircular. Eilyma psilate, ca. 0.5 m thick. Numerous (30+), discrete, generally evenly distributed, elongate, spine-like, homomorphic, straight to curving, solid, psilate processes; proximal contacts orthogonal to slightly curved at base; processes tapering gently to acuminate tips. No excystment structure observed. Measurements: Vesicle diameter 30 38 m, 24 32 m; process length 1218 m, 1416 m; process width at base ca. 1 m. (2 specimens). Discussion: This species supercially resembles Elektoriskos aktinotos Wicander, Playford and Robertson, 1999 but has a larger vesicle and longer processes. Because the ratio of process length to vesicle diameter is nearly the same for both species, it is possible that these specimens are just larger. However, because only two specimens were found, we prefer to leave them in open nomenclature. Occurrence: Stonington Formation (present study).
Genus Estiastra Eisenack, 1959 emend. Sarjeant and Stancliffe, 1994 Type species: Estiastra magna Eisenack, 1959; by original designation. Estiastra sp. A Plate 1, Figs. 20 and 21 Description: Vesicle outline stellate. Vesicle formed by the conuence of 57 broad-based, hollow processes, not in the same plane. Processes open into and freely communicate with vesicle interior. Processes taper to acuminate tip. Eilyma and process walls thin, <1 m, psilate. No excystment structure observed. Measurements: Vesicle diameter 40 (49) 66 m; process length 31 (48) 61 m; process width at base 7 (12) 18 m; overall diameter including processes 115 (138) 165 m. (4 specimens). Discussion: We concur with the emendation of Estiastra by Sarjeant and Stancliffe (1994) to include only those stellate forms composed of broad-based processes arising in more than one plane and whose conuence forms the central vesicle and whose distal tips are acuminate (not blunt or rounded). The present specimens do not readily t any previously erected species. They are much smaller overall than Estiastra magna Eisenack, 1959, which ranges up to 515 m, larger in overall size than E. oklahomensis Loeblich and Tappan, 1978 (115165 m vs. 60 m and 63 m), and have 57 processes versus only four processes for the two described species. Because only four well-preserved specimens were recovered, with the rest being deformed, we are leaving this species in open nomenclature. Occurrence: Bills Creek Shale and Stonington Formation (present study).
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Genus Excultibrachium Loeblich and Tappan, 1978 emend. Turner, 1984 Type species: Excultibrachium concinnum Loeblich and Tappan, 1978; by original designation. Excultibrachium concinnum Loeblich and Tappan, 1978 Plate 2, Figs. 1 and 2 1978. Excultibrachium concinnum Loeblich and Tappan, p. 12671268, Pl. 9, Figs. 36. 1979. Ordovicidium gracile Colbath, 1979, p. 23, Pl. 8, Figs. 47. 1984. Excultibrachium oligocladatum Turner, p. 110, Pl. 9, Figs. 7, 9. Measurements: Vesicle diameter 44 (55) 66 m; 1524 processes; process length 14 (19) 26 m; process width at base 2 (3.6) 4 m; furcation length 4 (6) 8 m. (13 specimens). Discussion: As noted by Wicander et al. (1999: p. 13), only minor morphologic differences exist between Excultibrachium concinnum Loeblich and Tappan, 1978, E. oligocladatum Turner, 1984, and Ordovicidium gracile Colbath, 1979, such that they should be considered conspecic. Our specimens, while falling within the range of published measurements have, on average, fewer and shorter processes, but otherwise t the circumscription of E. concinnum. Occurrence: Bills Creek Shale and Stonington Formation (present study). Previously reported from the Edenian (upper Caradoc) of Indiana (Loeblich and Tappan, 1978; Colbath, 1979); Richmondian (Ashgill) of Missouri (Wicander et al., 1999) and Anticosti Island, Qu ebec, Canada (Jacobson and Achab, 1985); CaradocAshgill of Labrador Sea, Canada (Legault, 1982); Caradoc (Harnagian-Actonian) of Shropshire, England (Turner, 1984); LlanvirnCaradoc (Viruan) of Gotland, Sweden (G orka, 1987); Caradoc and Ashgill of Gotland, Sweden (Eiserhardt, 1992); possibly from LlanvirnAshgill of Estonia (Uutela and Tynni, 1991); Caradoc of Xinjiang, northwestern China (Li et al., 2006). Genus Gloeocapsomorpha Zalessky, 1917 emend. Foster, Reed and Wicander, 1989 Type species: Gloeocapsomorpha prisca Zalessky, 1917; by monotypy. Gloeocapsomorpha prisca Zalessky, 1917 emend. Foster, Reed and Wicander, 1989 Plate 4, Figs. 4 and 11 1917. Gloeocapsomorpha prisca Zalessky, p. 814, Figs. 13, 69. 1989. Gloeocapsomorpha prisca Zalessky, 1917 emend. Foster, Reed and Wicander, p. 743745, Pl. 1, Figs. 112; Pl. 2, Figs. 116; Text-gs. 2, 57.1, 7.3, 7.5, 7.6. Measurements: The size of the cells, colonies, and supercolonies varies, depending on the number of cell divisions and whether they have been broken during palynologic processing. Discussion: Gloeocapsomorpha prisca Zalessky, 1917 emend. Foster et al., 1989 is an enigmatic palynomorph whose biological afnities are still unclear. Until its systematic position is resolved, we provisionally place it with the acritarchs. Occurrence: Bills Creek Shale and Stonington Formation (present study). Recorded worldwide from Ordovician
sequences, but most commonly in Middle Ordovician strata (Wicander et al., 1996). Genus Hoegklintia Dorning, 1981 1981. Hogklintia Dorning, p. 192. 1988. Hoegklintia Dorning nom. corr. Eley and Legault, 1988, p. 58, 63. Type species: Hoegklintia visbyensis (Eisenack, 1959) Dorning, 1981; by original designation. Hoegklintia sp. cf. H. radicosa (Loeblich, 1970) Jacobson and Achab, 1985 Plate 2, Figs. 3, 4 and 7 1970. cf. Multiplicisphaeridium radicosum Loeblich, p. 730, Fig. 23AE. 1985. cf. Hoegklintia radicosa (Loeblich, 1970) Jacobson and Achab, p. 183, Pl. 4, Fig. 2. Description: Vesicle subcircular in outline. Eilyma thin (<1 m thick), psilate. Seventeen to 23, broad-based, stout, hollow processes, drawn out from and communicating freely with vesicle interior. Process trunks taper gradually towards blunted distal ends that typically furcate to second order, although some furcate to third order, and a few remain acuminate. Processes surface psilate. No excystment structure observed. Measurements: Vesicle diameter 25 (28) 33 m; process length 917 m; process width at base 3.46.6 m; overall diameter including processes 37 (50) 61 m. (4 specimens). Discussion: Only four specimens of this distinctive form were recovered. Whereas they are well-preserved and t the description of Hoegklintia radicosa (Loeblich, 1970) Jacobson and Achab, 1985, they all have a smaller vesicle and shorter processes than specimens described from elsewhere in North America; thus a cf. designation is preferred. However, the present specimens are of comparable size to the smaller specimens of Multiplicisphaeridium radicosum Loeblich, 1970 as described and illustrated by Uutela and Tynni (1991: p. 9596, Pl. XXII, Fig. 224). Playford and Wicander (2006) did not consider the specimens of M. radicosum recorded by Uutela (1989) and Uutela and Tynni (1991) from the Finnish and Estonian Ordovician as authentic representatives of H. radicosa. Pending availability of additional specimens revealing a size-continuum linking the smaller specimens with those considered typical of H. radicosa, a cf. designation for the smaller forms seems prudent at this time. Occurrence: Stonington Formation (present study). Hoegklintia radicosa has previously been reported from the Richmondian (Ashgill) of Missouri (Miller, 1991; Wicander et al., 1999), Oklahoma (Loeblich, 1970; Playford and Wicander, 2006), and Qu ebec, Canada (Martin, 1980; Jacobson and Achab, 1985); Mohawkian (Caradoc) and Edenian (upper Caradoc) of Qu ebec and southeast Ontario, Canada (Martin, 1983). Genus Leiofusa Eisenack, 1938 Type species: Leiofusa fusiformis (Eisenack, 1934) Eisenack, 1938; by original designation. Leiofusa fusiformis (Eisenack, 1934) Eisenack, 1938 Plate 4, Figs. 1, 2 and 10 1934. Ovum hispidum fusiformis Eisenack, p. 6566, Pl. 4, Fig. 19.
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1938. Leiofusa fusiformis (Eisenack, 1934) Eisenack, p. 28. Measurements: Vesicle length 140 (176) 236 m; maximum vesicle width 24 (31) 54 m. (29 specimens). Discussion: Our specimens include those whose psilate vesicle is both symmetrical and slightly asymmetrical around the long axis and bears equal processes, together with those that have essentially the same morphologic range but feature just-visible discontinuous longitudinal striae that are probably induced by compression along the long axis. This range of morphology encompasses a variety of species assigned to Leiofusa; viz., L. fusiformis as illustrated by Jacobson and Achab (1985), Eupoikilofusa striatifera as illustrated by Cramer (1971) and Jacobson and Achab (1985), Dactylofusa striatifera as illustrated by Molyneux et al. (1996), L. elenae as illustrated by Cramer (1971), and L. asymmetrica as illustrated by Colbath (1979). It is beyond the scope of this study to determine the amount of morphologic variation acceptable in circumscribing these species. There is a morphologic continuum (i.e., between symmetrical and slightly asymmetrical vesicle shape, and psilate to faintly striate eilyma) that is probably reective of variation found in natural populations. Accordingly, we assign our specimens to L. fusiformis (Eisenack, 1934) Eisenack, 1938, which ts the morphology most commonly encountered in our samples. Occurrence: Bills Creek Shale and Stonington Formation (present study). Depending on which species are considered synonymous, the range of L. fusiformis is Upper Ordovician through Silurian. Leiofusa litotes Loeblich and Tappan, 1978 Plate 3, Figs. 1 and 2 1978. Leiofusa litotes Loeblich and Tappan, p. 1271, Pl. 12, Figs. 1, 2. Measurements: Vesicle length 176 (196) 220 m; maximum vesicle width 14 (20) 26 m. (7 specimens). Occurrence: Bills Creek Shale and Stonington Formation (present study). Previously reported from the CaradocAshgill of Kansas (Wright and Myers, 1981); Richmondian (Ashgill) of Missouri (Miller, 1991) and Oklahoma (Loeblich and Tappan, 1978). Provisionally reported (as Leiofusa aff. L. litotes) by Hill and Molyneux (1988) and Molyneux (1988) from Well E1-81, Libya, in rocks dated as Ashgill. Elaouad-Debbaj (1988: p. 237) cited a questionable and also reputedly Ashgill occurrence (Leiofusa litotes?) from the Anti-Atlas of Morocco, but did not illustrate any specimens. Rubinstein and Vaccari (2004) recorded Leiofusa cf. L. litotes from the Ordovician/Silurian boundary in northwest Argentina. Genus Lophosphaeridium Timofeev, 1959 ex Downie, 1963 Type species: Lophosphaeridium rarum Timofeev, 1959; by subsequent designation of Downie (1963). Discussion: We follow Colbaths (1990: p. 116) suggestion that Lophosphaeridium Timofeev, 1959 ex Downie, 1963 be restricted to species with simple processes that are less than twice as long as wide at base, thus facilitating distinction of
that genus from Gorgonisphaeridium Staplin, Jansonius and Pocock, 1965. Lophosphaeridium acinatum Wicander, Playford and Robertson, 1999 Plate 4, Fig. 3 1999. Lophosphaeridium acinatum Wicander, Playford and Robertson, p. 15, Fig. 8.68.11. Measurements: Vesicle diameter 33 m, 50 m, 54 m; grana height ca. 1.1 m; grana width 11.5 m. (3 specimens). Occurrence: Stonington Formation (present study). Previously reported from the Richmondian (Ashgill) of Missouri (Wicander et al., 1999). Lophosphaeridium edenense Loeblich and Tappan, 1978 Plate 3, Fig. 5 1978. Lophosphaeridium edenense Loeblich and Tappan, p. 12721273, Pl. 14, Figs. 4, 5. Measurements: Vesicle diameter 34 m, 42 m; grana height ca. 0.8 m; grana width ca. 0.6 m. (2 specimens). Occurrence: Bills Creek Shale and Stonington Formation (present study). Previously reported from the Caradoc and Ashgill of Kansas (Wright and Myers, 1981); Richmondian (Ashgill) of Missouri (Wicander et al., 1999) and Anticosti Island, Qu ebec, Canada (Jacobson and Achab, 1985); Edenian (upper Caradoc) of Kentucky and Indiana (Loeblich and Tappan, 1978); Caradoc of Tarim Basin, China (Li and Wang, 1997) and Xinjiang, northwestern China (Li et al., 2006). Lophosphaeridium varum Wicander, Playford and Robertson, 1999 Plate 3, Figs. 3 and 4 1999. Lophosphaeridium varum Wicander, Playford and Robertson, p. 1516, Fig. 9.19.5. Measurements: Vesicle diameter 32 (37) 44 m; grana height ca. 11.5 m; grana width ca. 0.81 m. (4 specimens). Occurrence: Bills Creek Shale and Stonington Formation (present study). Previously reported from the Richmondian (Ashgill) of Missouri (Wicander et al., 1999). Genus Micrhystridium Deandre, 1937 Type species: Micrhystridium inconspicuum Deandre, 1937; by original designation. Micrhystridium hirticulum Wicander, Playford and Robertson, 1999 Plate 3, Figs. 8 and 11 1999. Micrhystridium hirticulum Wicander, Playford and Robertson, p. 17, Fig. 9.69.8. Measurements: Vesicle diameter 22 (25) 30 m; >30 processes; process length 10 (11) 14 m; process width at base ca. 1.5 m. (9 specimens). Occurrence: Bills Creek Shale and Stonington Formation (present study). Previously reported from the Richmondian (Ashgill) of Missouri (Wicander et al., 1999) and Oklahoma (Playford and Wicander, 2006).
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Micrhystridium prolixum Wicander, Playford and Robertson, 1999 Plate 3, Figs. 7 and 10 1999. Micrhystridium prolixum Wicander, Playford and Robertson, p. 17, Figs. 9.13, 10.7. Measurements: Vesicle diameter 14 (19) 21 m; 1116 processes; process length 16 (19) 21 m; process width at base 23 m. (5 specimens). Occurrence: Bills Creek Shale and Stonington Formation (present study). Previously reported from the Richmondian (Ashgill) of Missouri (Wicander et al., 1999) and Oklahoma (Playford and Wicander, 2006); Caradoc of Xinjiang, northwestern China (Li et al., 2006). Genus Multiplicisphaeridium Staplin, 1961 emend. Sarjeant and Vavrdov a, 1997 Type species: Multiplicisphaeridium ramispinosum Staplin, 1961 emend. Sarjeant and Vavrdov a, 1997; by original designation. Multiplicisphaeridium irregulare Staplin, Jansonius and Pocock, 1965 Plate 3, Fig. 6 1965. Multiplicisphaeridium irregulare Staplin, Jansonius and Pocock, p. 183, Pl. 18, Fig. 18 (non Fig. 17). Measurements: Vesicle diameter 20 (23) 28 m; 1218 processes; process length 12 (15) 19 m; process width at base 23 m. (7 specimens). Discussion: Playford and Wicander (2006) discussed the differences between Multiplicisphaeridium bifurcatum Staplin, Jansonius and Pocock, 1965 and M. irregulare Staplin, Jansonius and Pocock, 1965, which are morphologically similar and typically occur together. The main difference is that M. irregulare exhibits a greater degree of process heteromorphy than M. bifurcatum. Despite the fact these two species tend to co-occur, all of our specimens from the Stonington Formation exhibit a high degree of process heteromorphy (not the near-homomorphic processes with regular rst-order distal bifurcation characterizing M. bifurcatum). Occurrence: Stonington Formation (present study). See Wicander et al. (1999) for a listing of most previous occurrences. Molyneux et al. (1996) considered this species to be cosmopolitan in basal Caradoc through upper Ashgill strata. Genus Navifusa Combaz, Lange and Pansart, 1967 ex Eisenack, 1976 Type species: Navifusa navis (Eisenack, 1938) Eisenack, 1976; by original designation. Navifusa sp. A Plate 4, Figs. 7 and 8 Description: Vesicle navicular, sides straight and parallel, ends rounded to broadly rounded. Eilyma thin, psilate. No excystment structure observed. Measurements: Vesicle length 171 (193) 218 m; maximum vesicle width 59 (65) 71 m. (4 specimens). Occurrence: Bills Creek Shale (present study).
Discussion: Navifusa sp. A supercially resembles N. indianensis Loeblich and Tappan, 1978 and Leiovalia teretis (Loeblich, 1970) Loeblich and Tappan, 1978. N. indianensis was described as ornamented with a dense cover of ne, variably spaced punctae (Loeblich and Tappan, 1978: p. 1277), whereas the present specimens are psilate. However, the punctae are not obvious on the high-magnication gure of the eilyma of N. indianensis (Loeblich and Tappan, 1978: Pl. 13, Fig. 4). Furthermore, only one measurement was given (holotype), and our specimens are larger than the holotype. Navifusa sp.A is comparable in size to L. teretis as well as having a psilate eilyma. Loeblich and Tappan (1978: p. 1272) transferred teretis from Navifusa to Leiovalia because of the elongate-ovate vesicle and absence of parallel sides in the vesicle of the 19 specimens observed. Our specimens have straight, parallel sides and resemble the specimens illustrated by Loeblich (1970) and Loeblich and Tappan (1978). However, because of the scarcity and somewhat poor preservation of our specimens, we are leaving them in open nomenclature, rather than making a denite attribution. Genus Orthosphaeridium Eisenack, 1968 Type species: Orthosphaeridium rectangulare (Eisenack, 1963) Eisenack, 1968; by original designation. Orthosphaeridium rectangulare (Eisenack, 1963) Eisenack, 1968 Plate 4, Fig. 9 1963. Baltisphaeridium rectangulare Eisenack, p. 211, Pl. 20, Figs. 13, 10. 1968. Orthosphaeridium rectangulare (Eisenack, 1963) Eisenack, p. 92, Pl. 25, Fig. 1. 1970. Orthosphaeridium inatum Loeblich, p. 733734, Fig. 29ae. For additional synonymy, see Wicander et al. (1999: p. 1921). Measurements: Vesicle diameter 52 (59) 66 m; 4 processes; process length 62 (75) 106 m; process width 46 m; process basal plug 56 m thick (9 specimens). Discussion: All specimens encountered of Orthosphaeridium rectangulare (Eisenack, 1963) Eisenack, 1968 are either fragments or half-vesicles, having broken apart along the median excystment split. Occurrence: Bills Creek Shale and Stonington Formation (present study). Previously reported from the Richmondian (Ashgill) of Missouri (Miller, 1991; Wicander et al., 1999) and Oklahoma (Loeblich, 1970; Playford and Wicander, 2006); RichmondianGamachian (Ashgill) of Anticosti Island, Qu ebec, Canada (Jacobson and Achab, 1985; Martin, 1988); LlanvirnCaradoc (Viruan) of Gotland, Sweden (Kjellstr om, 1971);?LlanvirnAshgill of Iran (Ghavidel-syooki, 2001, 2003); Caradoc and Ashgill of Gotland, Sweden (Eisenack, 1968; Eiserhardt, 1985); and Ashgill of Estonia (Uutela and Tynni, 1991), Morocco (Elaouad-Debbaj, 1988), and Jordan (Keegan et al., 1990). Molyneux et al. (1996: Text-g. 8) described this species as cosmopolitan with a stratigraphic range of basal Caradoc through upper Ashgill.
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Genus Peteinosphaeridium Staplin, Jansonius and Pocock, 1965 emend. Playford, Ribecai and Tongiorgi, 1995 Type species: Peteinosphaeridium bergstroemii Staplin, Jansonius and Pocock, 1965 emend. Playford, Ribecai and Tongiorgi, 1995; by original designation. Peteinosphaeridium septuosum Wicander, Playford and Robertson, 1999 Plate 4, Figs. 5, 6, 12, and 13 1981. Baltisphaeridium sp. b of Wright and Myers, p. 22, Pl. 5, Figs. AC. 1999. Peteinosphaeridium septuosum Wicander, Playford and Robertson, p. 21, 23, Figs. 11.511.9, 12.2, 12.3. Measurements: Vesicle diameter 48 (63) 75 m; 14 (15) 18 processes; process length 6 (8) 11 m; process width at base 2 m. (23 specimens). Pylome diameter 14 (15) 18 m. (9 specimens). Occurrence: Bills Creek Shale and Stonington Formation (present study). Previously reported from the Caradoc and Ashgill of Kansas (Wright and Myers, 1981); Richmondian (Ashgill) of Missouri (Wicander et al., 1999) and Oklahoma (Playford and Wicander, 2006). Genus Polygonium Vavrdov a, 1966 emend. Sarjeant and Stancliffe, 1994 Type species: Polygonium gracile Vavrdov a, 1966; by original designation. Polygonium gracile Vavrdov a, 1966 emend. Sarjeant and Stancliffe, 1996 Plate 4, Figs. 17 and 18 1966. Polygonium gracile Vavrdov a, p. 413, Pl. 1, Fig. 3, Pl. 3, Fig. 1. 1996. Polygonium gracile Vavrdov a, 1966 emend. Sarjeant and Stancliffe, p. 359360. For complete synonymy, see Sarjeant and Stancliffe (1996: p. 359360). Measurements: Vesicle diameter 22 (29) 41 m; 916 processes; process length 16 (24) 32 m; process width at base 2 (2.5) 4 m. (23 specimens). Discussion: The Bills Creek Shale and Stonington Formation specimens t the species emendation of Sarjeant and Stancliffe (1996). The specimens are, however, somewhat variable in vesicle diameter, number of processes, and process length. As discussed by Wicander et al. (1999: p. 23), this represents a morphologic continuum that evidently reects morphologic variation within a population. If only the end members of the continuum were observed, they might be considered as specically distinct, but viewed as a whole, the entire population must be considered a single species. Occurrence: Bills Creek Shale and Stonington Formation (present study). Previously reported from the Richmondian (Ashgill) of Missouri (Wicander et al., 1999), Oklahoma (Playford and Wicander, 2006), and Anticosti Island, Qu ebec, Canada (Jacobson and Achab, 1985). Upper Cambrian through Devonian strata globally elsewhere (Playford and Wicander, 2006).
Genus Sylvanidium Loeblich, 1970 Type species: Sylvanidium paucibrachium Loeblich, 1970; by original designation, monotypy. Sylvanidium paucibrachium Loeblich, 1970 Plate 4, Figs. 21 and 22 1965. Acritarchous hystrichosphere of Staplin, Jansonius and Pocock, p. 185, Pl. 19, Fig. 15. 1970. Sylvanidium paucibrachium Loeblich, p. 737, Fig. 32AF. Measurements: Vesicle length 54 (69) 80 m; vesicle width 44 (59) 74 m; 46 processes; process length 14 (22) 34 m; process width at base 2 (4.1) 6 m. (11 specimens). Discussion: Jacobson and Achab, (1985: p. 193) commented on the supercial resemblance between Sylvanidium paucibrachium Loeblich, 1970 and Dorsennidium hamii, and on their tendency to co-occur. Playford and Wicander (2006) also noted the similarity in morphology and conjectured that the two species may intergrade morphologically. We have observed the same similarity in morphology between the two species, and follow Playford and Wicander (2006) in separating them primarily on vesicle shape; that is, bell-shaped for D. hamii, and bean-shaped for S. paucibrachium. Occurrence: Stonington Formation (present study). Previously reported from the Richmondian (Ashgill) of Oklahoma (Loeblich, 1970; Playford and Wicander, 2006) and Qu ebec, Canada (Staplin et al., 1965; Martin, 1980; Jacobson and Achab, 1985). According to Molyneux et al. (1996: Text-g. 8), S. paucibrachium is restricted to the middle Ashgill of Laurentia. Genus Veryhachium Deunff, 1954 ex Downie, 1959 Type species: Veryhachium trisulcum (Deunff, 1951) ex Deunff, 1959; by subsequent designation of Downie (1959). Veryhachium europaeum Stockmans and Willi` ere, 1960 Plate 3, Fig. 9 1960. Veryhachium europaeum Stockmans and Willi` ere, p. 3, Pl. 2, Fig. 25. Measurements: Vesicle diameter 48 m, 50 m; 3 processes drawn out from plane of vesicle, 23 supplementary processes arising from plane of vesicle; process length 14 m, 22 m. (2 specimens). Occurrence: Bills Creek Shale and Stonington Formation (present study). Veryhachium europaeum Stockmans and Willi` ere, 1960 is a cosmopolitan species with a previously recorded stratigraphic range of Silurian through Devonian (Wicander and Wood, 1981). Veryhachium oklahomense Loeblich, 1970 Plate 4, Figs. 1416 1970. Veryhachium oklahomense Loeblich, p. 742743, Fig. 36F, G. 1985. Veryhachium lairdii (auct. non Deandre) Deunff ex Downie, 1958. Jacobson and Achab, p. 195, Pl. 9, Fig. 2. Measurements: Vesicle length 14 (28) 42 m; vesicle width 14 (34) 48 m; process length 8 (22) 30 m; process width at base 4 (4.5) 6 m. (6 specimens).
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Occurrence: Bills Creek Shale and Stonington Formation (present study). See Wicander et al. (1999) for a complete listing of previous occurrences, all but one of which are Upper Ordovician, and that one (Uutela and Tynni, 1991 ArenigLlanvirn) we consider to be questionable. Ghavidel-syooki (2001, 2003) has recorded the species from Iranian strata dated as LlanvirnAshgill, and Rubinstein and Vaccari (2004) noted its occurrence from the Ordovician/Silurian boundary of northwest Argentina. Veryhachium trispinosum (Eisenack, 1938) Stockmans and Willi` ere, 1962 complex Plate 4, Figs. 19 and 23 1938. Hystrichosphaeridium trispinosum Eisenack, p. 14, 16, Text-gs. 2, 3. 1954. Veryhachium trispinosum (Eisenack, 1938) Deunff, p. 306. [combination invalid: ICBN, Article 33.2]. 1962. Veryhachium trispinosum (Eisenack, 1938) Stockmans and Willi` ere, p. 4647, Pl. II, Figs. 25, 26, Text-g. 1. 1981. Veryhachium trispinosum (Eisenack, 1938) Deunff, 1954 complex. Wicander and Wood, p. 6771. For additional synonymy, see Wicander and Wood (1981: p. 6770). Measurements: Vesicle diameter 28 (37) 40 m; process length 14 (22) 26 m. (14 specimens). Discussion: The Bills Creek Shale and Stonington Formation specimens form a morphologic continuum in which the vesicle sides are straight to weakly convex, the spine-like processes are drawn out from the vesicle corners, and the eilyma and process wall are thin and psilate or scabrate under light microscopy, and psilate to microgranulate under scanning electron microscopy. When viewed as a whole population, it is reasonable to regard these morphologically simple specimens as part of a complex rather than speciating them on the basis of minor morphologic variations (Wicander and Wood, 1981). Occurrence: Bills Creek Shale and Stonington Formation (present study). This species complex has a cosmopolitan distribution and ranges from Ordovician through Permian (Wicander and Wood, 1981: p. 71). Genus Villosacapsula Loeblich and Tappan, 1976 Type species: Villosacapsula setosapellicula (Loeblich, 1970) Loeblich and Tappan, 1976; by original designation. Villosacapsula setosapellicula (Loeblich, 1970) Loeblich and Tappan, 1976 Plate 4, Figs. 20 and 24 1970. Veryhachium setosapellicula Loeblich, p. 743, Figs. 36a,b, 37a,b. 1971. Veryhachium calandrae Cramer, p. 106, Pl. 6, Fig. 99, Text-g. 29a. 1976. Villosacapsula setosapellicula (Loeblich, 1970) Loeblich and Tappan, p. 307. Measurements: Vesicle diameter 28 (32) 40 m; process length 12 (20) 30 m; process width at base 2 (2.5) 3 m. (9 specimens).
Discussion: As noted by Wicander et al. (1999: p. 27), Cramers (1971) description and line-drawing of a specimen from the Maysville Formation (Upper Ordovician: Caradoc) of Ohio indicate that it probably belongs to Villosacapsula setosapellicula (Loeblich, 1970) Loeblich and Tappan, 1976. Cramer (1971) assigned the specimen to Veryhachium calandrae Cramer, 1971 but omitted to designate a holotype for that species, which accordingly is invalid. Occurrence: Bills Creek Shale and Stonington Formation (present study). Previously reported from the Edenian (upper Caradoc) of Indiana (Colbath, 1979); CaradocAshgill of Kansas (Wright and Myers, 1981); Richmondian (Ashgill) of Missouri (Miller, 1991; Robertson, 1997; Wicander et al., 1999) and Oklahoma (Loeblich, 1970; Playford and Wicander, 2006); Mohawkian (Caradoc) and Edenian (upper Caradoc) of Qu ebec and Ontario, Canada (Martin, 1983); upper Arenig-Llanvirn and Caradoc-?Ashgill of Algerian Sahara (Jardin e et al., 1974); Ashgill of Algerian Sahara and southern Tunisia (Vecoli, 1999); Caradoc of Shropshire, England (Turner, 1984) and northwest Libya (Deunff and Massa, 1975); CaradocAshgill of Czech Republic (Vavrdov a, 1988) and Jordan (Keegan et al., 1990); Ashgill of northeast Libya (Molyneux and Paris, 1985; Hill and Molyneux, 1988); Ordovician/Silurian boundary of northwest Argentina (Rubinstein and Vaccari, 2004). Probably also from the upper Caradoc of Ohio (Cramer, 1971; see above synonymy and discussion). The reputedly pre-Caradoc occurrence by Jardin e et al. (1974) lacks descriptive and illustrative documentation, in addition to a questionable palynological age attribution, and is accordingly considered unsubstantiated (Vecoli, 1999). Ghavidel-syooki (2001, 2003) recorded this species from Iranian strata he dated as LlanvirnAshgill. 5. Composition of the palynoora All 17 sampled intervals from the Bills Creek Shale (eight samples) and Bay de Noc Member of the Stonington Formation (nine samples) yielded a generally diverse and advantageously preserved palynomorph assemblage (Figs. 4 and 5). The majority of samples yielded abundant specimens based on total prasinophyte phycomata and acritarch counts from the three >52 m slides and three 2052 m slides counted per sample. No counts were made from the <20 m-fraction slides, but they were examined to ensure that no small specimens had been excluded from the recovered assemblages. Palynomorphs per sample ranged from a low of 94+ specimens (sample BC8) to a high of 533+ specimens (sample S4). Diversity of the palynoora ranged from a low of 10 species (sample BC8) to a high of 25 species (samples S3, S4, S5). Typically, the Stonington Formations palynoora is more abundant and diverse than that of the Bills Creek Shale. In addition to the prasinophyte phycomata and acritarchs, scolecodonts and graptolite fragments were frequently encountered in the strew slides, as well as abundant and diverse chitinozoans in many of the slides. The palynoora recovered from the Bills Creek Shale and Bay de Noc Member of the Stonington Formation is overwhelmingly represented by acritarchs. The acritarchs (including the enigmatic palynomorph Gloeocapsomorpha prisca) comprise
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Fig. 4. Occurrence of prasinophyte and acritarch species in the studied samples of the Bills Creek Shale. All specimens present on the rst three slides of the >52 m and 2052 m fractions were counted to determine relative abundances of species per sample. The term very abundant (=va) applies to those species represented by >100 specimens; abundant (=a) for 51100 specimens; common (=) for 1050 specimens; and rare (=r) for <10 specimens. Because of the large numbers of Gloeocapsomorpha prisca masses, its presence is indicated by a double asterisk (**) for high abundance, or a single asterisk (*) for lesser abundance. The total number of specimens counted per sample is indicated at the base of the table. When 100 specimens of a particular species were reached in the count, that species was no longer counted. Thus, most total counts are followed by a plus (+) sign to signify >100 specimens were encountered. Fig. 4. Pr esence des esp` eces dacritarches et de prasinophytes dans les chantillons pr e elev es dans la Formation de Bill Creek Shale. Le calcul de t labondance relative des esp` eces a e e effectu e sur la base de tous les sp ecimens pr esents dans les trois premi` eres lames palynologiques des fractions >52 m et 2025 m. Cl e des abr eviations : va = >100 sp ecimens ; a = 51100 sp ecimens ; c = 1050 sp ecimens ; r = <10 sp ecimens. Pour lesp` ece tr` es commune Gloeocapsomorpha prisca le double ast erisque (**) indique une grande abondance et last erisque simple (*) indique une abondance mineure.
Fig. 5. Occurrence of prasinophyte and acritarch species in the studied samples of the Stonington Formation. For explanation of quantitative methods used to determine relative abundance, see Fig. 4. Fig. 5. Pr esence des esp` eces dacritarches et de prasinophytes dans les chantillons pr e elev es dans la Formation de Stonington. Pour les explications des symboles et des m ethodes utilis es pour les comptages voir Fig. 4.
29 species assigned to 20 genera. Of the acritarch species, 24 are previously named species, one is designated cf., one is categorized as a complex, and three are left in open nomenclature as sp. A. The prasinophyte phycomata are represented by undifferentiated species of Leiosphaeridia and Tasmanites. The abundance and diversity of the acritarch assemblage obtained from the Bills Creek Shale is more uniform than that from the Stonington Formation (Figs. 4 and 5). Except for sample BC8, which contains nine species, the other seven samples contain between 12 and 17 species (Fig. 4). The two most abundant acritarchs in the Bills Creek assemblage are Leiofusa fusiformis and members of the Veryhachium trispinosum complex. Other commonly occurring acritarch species from the Bills Creek Shale include Peteinosphaeridium septuosum and Villosacapsula setosapellicula. The rest of the species are generally rare to common and their abundance is fairly consistent throughout the eight samples. Two prasinophyte genera occur in the Bills Creek Shale (Fig. 4). Leiosphaeridia spp. is present
in all sampled intervals and, except for samples BC2 and BC7 where it is common, it is rare throughout the sampled section. Tasmanites spp. occurs in ve of the eight sampled intervals and is rare in each of those samples. The acritarch assemblage of the Bay de Noc Member of the Stonington Formation is generally more abundant and diverse than that of the Bills Creek Shale (Figs. 4 and 5). As in the Bills Creek assemblage, the two most abundant forms are L. fusiformis and members of the V. trispinosum complex. Other common to abundant species include Dorsennidium hamii, Leiofusa litotes, Micrhystridium hirticulum, Peteinosphaeridium septuosum, Polygonium gracile, and Villosacapsula setosapellicula. Diversity ranges from 11 to 25 species per sample with D. hamii, L. fusiformis, P. gracile, and the V. trispinosum complex typically being the most abundant taxa in each sample (Fig. 5). As with the Bills Creek Shale assemblage, the same two prasinophyte genera are present. Leiosphaeridia spp. was recovered in all nine sampled intervals and was rare in three of the samples and common in six samples. Tasmanites spp. only occurred in four samples (S2S5), being common in samples S2S4 and rare in sample S5.
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6. Paleoenvironmental interpretation As stated earlier, sedimentologic and paleontologic evidence indicates that the Bills Creek Shale was deposited in a low-energy, shallow, normal marine environment, with at least occasional aerial exposure as indicated by rare desiccation cracks within some of the calcareous shale layers. The interbedded argillaceous limestones may represent more offshore conditions resulting from intermittent transgressions. The Bay de Noc Member of the Stonington Formation accumulated in a low-energy, offshore, normal marine environment, with periodic regressions resulting in terrigenous input as evidenced by the shales. The palynologic assemblages recovered from these two formations generally supports the paleoenvironmental interpretation suggested by the sedimentologic and paleontologic evidence. Staplin (1961) rst demonstrated a relationship between differing acritarch morphotype assemblages and distance from Upper Devonian reefs in Alberta, Canada. Later studies have shown that qualitative changes in microphytoplankton morphotypes and diversity and abundance uctuations can assist in determining nearshore to offshore trends (e.g., Jacobson, 1979; Dorning, 1981; Al-Ameri, 1983; Vecoli, 2000; Li et al., 2004; Stricanne et al., 2004; Vecoli and Le H eriss e, 2004). However, just as interrelated physical and ecological factors are responsible for changes in the composition and distribution patterns of modern microphytoplankton assemblages, such as dinoagellates (Dale, 1996; Marret and Zonneveld, 2003; Rochon and Marret, 2004), these same factors are also most likely responsible for the distribution of Paleozoic microphytoplankton (Colbath, 1980; Strother, 1996). These various complex interrelating factors must therefore be taken into consideration before any rm relationships can be established between the preserved palynomorph assemblage and the paleoenvironment. Nonetheless, compositional changes in the palynomorph assemblages preserved throughout a stratigraphic section can contribute to paleoenvironmental syntheses. The paleoenvironmental interpretation based on the palynologic composition of the Bills Creek Shale complements that deduced from sedimentologic and paleontologic studies. The Bills Creek Shale palynomorph assemblage represents a nearshore to inner offshore environment based on the assumptions and criteria of Dorning (1981), Vecoli (2000), and Li et al. (2004). However, the Bills Creek palynomorphs do not readily t into a specic environmental category based on Jacobsons (1979) form-class assignments. This is partly because leiofusid acritarchs were not included in his classes, and this category comprises a large proportion of the Bills Creek palynologic assemblage. According to Dorning (1981), the Eupoikilofusa striatifera and Leiofusa estrecha complex (i.e., leiofusid acritarchs) appear to prefer an inshore environment, although they do range from nearshore to deep water. Inshore and nearshore species of Veryhachium commonly have three or four processes on a planar vesicle, whereas offshore forms commonly have four to six processes on an inated vesicle.
Li et al. (2004) stated that the nearshore marine environment of the Yangtze Platform, South China, was dominated by fusiform, leiosphaerid, and polygonomorph acritarchs. Surprisingly, they did not mention any veryhachid components. They observed that both generic and specic diversity increased from nearshore to offshore, a trend also noted by Dorning (1981), Vecoli (2000), and Stricanne et al. (2004), and in studies focusing on the distribution of extant microphytoplankton. The two most abundant palynomorphs of the Bills Creek assemblage are Leiofusa fusiformis and members of the Veryhachium trispinosum complex, followed by Villosacapsula setosapellicula. In addition, the leiofusid acritarchs Leiofusa litotes, Dactylofusa ctenista, and D. playnetrella are rare to common, as are such taxa as Peteinosphaeridium septuosum, Polygonium gracile, and the veryhachids Veryhachium europaeum and V. oklahomense. Except for D. ctenista and D. playnetrella, all of the aforementioned species occur in all or seven of the eight Bills Creek Shale samples. Additionally, Leiosphaeridia spp. and Lophosphaeridium varum are present in all samples, and Lophosphaeridium edenense occurs in six samples. Taking into account the composition of the palynomorph assemblage, as well as species diversity and abundance for each sample (Fig. 4), the palynologic evidence also points to a nearshore marine environment for the Bills Creek Shale, with transgressive episodes resulting in an innermost offshore environment. The palynomorph assemblage of the Bay de Noc Member of the Stonington Formation is more diverse and abundant than that of the Bills Creek Shale, and is not dominated by a few species. Based on those palynologic observations alone, the Stonington Formation represents a more offshore marine environment than the Bills Creek Shale (cf. Dorning, 1981; Vecoli, 2000; Li et al., 2004; Stricanne et al., 2004). The Stonington Formation assemblage conforms to Jacobsons (1979) offshore, open-marine suite. This suite comprises the baltisphaerid-veryhachid-Polygonium-micrhystridid formclasses, all of which commonly occur throughout the sampled Stonington. As noted earlier, whereas Leiosphaeridia and related psilate sphaeromorphs are indicative of Jacobsons nearshore environment, they are also commonly associated with diverse acritarch assemblages in offshore situations. The Silurian offshore shelf assemblage of Dorning (1981) has high diversity and moderate abundance per sample, with no single taxon dominating. Although the Eupoikilofusa striatifera and Leiofusa estrecha complex is more abundant nearshore, it is still very common in the farther offshore region of the shelf. The veryhachids also are common from the nearshore to offshore environment, with forms having four to six processes on an inated vesicle (like Dorsennidium hamii) more common offshore. As stated by Vecoli (2000), Li et al. (2004), and others, diversity increases along a nearshore to offshore transect. In addition, Li et al. (2004) indicated that the acanthomorph genera Baltisphaeridium and Peteinosphaeridium, as well as polygonomorph genera such as Polygonium, were the dominant elements of the EarlyMiddle Ordovician Yangtze Platform offshore acritarch assemblage. Whereas Baltisphaeridium adi-
59
astaltum and B. perclarum are rare to abundant, they do occur consistently throughout the sampled Stonington section. Furthermore, Peteinosphaeridium septuosum and Polygonium gracile are likewise abundant and persistent elements. Thus, the Stonington Formations paleoenvironment based on palynologic evidence mirrors that of the sedimentologic and paleontologic data, by indicating a marine, offshore shelf environment. Furthermore, the Stonington Formation (Bay de Noc Member) is considered to have accumulated farther offshore than the underlying Bills Creek Shale. 7. Comparison with other acritarch assemblages The numerous publications dealing with Caradoc and Ashgill acritarch/prasinophyte palynooras are mainly from Laurentia, Baltica, Avalonia, the northern Gondwana margin, and South China (e.g., Molyneux et al., 1996; Servais et al., 2004; Vecoli and Le H eriss e, 2004). Unfortunately, despite a fairly large database of taxa for the Upper Ordovician, many of the publications lack denitive stratigraphic information or independent age control, particularly in relation to graptolite, conodont, or chitinozoan biozones. Because the Bills Creek Shale and Stonington Formation are assigned to the upper Amplexigraptus manitoulinensis (Bills Creek Shale) and Dicellograptus complanatus (Stonington Formation) graptolite Biozones and the upper Amorphognathus superbus and lower A. ordovicicus conodont Biozones, we focus our comparisons to other acritarch assemblages on those dated reliably as Richmondian (North American terminology) or the equivalent, globally recognized Ashgill (Fig. 1). 7.1. Laurentia As indicated by Wicander et al. (1999), Wicander (2004), and Playford and Wicander (2006), knowledge of Cincinnatian (including Richmondian) palynooras from Laurentia is by no means comprehensive. However, the majority of them are welldocumented in terms of stratigraphic and paleontologic control. When considering palynomorph assemblages of exclusively Richmondian (=Ashgill) age, denitive comparisons can be made to the Maquoketa Shale of Missouri and Kansas, the Sylvan Shale of Oklahoma, and the Vaur eal Formation of Anticosti Island, Qu ebec, Canada (Figs. 1 and 6). Martin (1980) described an acritarch palynoora from part of the Whitehead Formation (CaradocAshgill) of the Perc e region, Gasp e Peninsula, Qu ebec, Canada, but only four of the 15 acritarch species she listed also occur in the Bills Creek/Stonington assemblage (viz., Aremoricanium squarrosum, Baltisphaeridium perclarum, Multiplicisphaeridium irregulare, and Sylvanidium paucibrachium). A total of 16 genera and 25 named or cf. species comprise the present organic-walled microphytoplankton assemblage recovered from the Bills Creek Shale and Bay de Noc Member of the Stonington Formation (excluding Leiosphaeridia, Tasmanites, and Gloeocapsomorpha prisca): Fig. 6. Of the 25 species, 76% also occur in the Maquoketa Shale of Kansas and Missouri, 72% in the Sylvan Shale of Oklahoma, and 60% in the Vaur eal Formation of Anticosti Island, Qu ebec, Canada. In
Fig. 6. Comparison of the Bills Creek Shale and Stonington Formation acritarch assemblage with published age-equivalent (Richmondian) palynooras from the Maquoketa Shale of Missouri and Kansas (Wright and Myers, 1981; Miller, 1991; Wicander et al., 1999), the Sylvan Shale of Oklahoma (Loeblich, 1970; Loeblich and MacAdam, 1971; Loeblich and Tappan, 1978; Playford and Wicander, 2006), and the Vaur eal Formation of Anticosti Island, Qu ebec, Canada (Staplin et al., 1965; Jacobson and Achab, 1985). ` acritarches des Formations de Bills Fig. 6. Comparaison des assemblages a Creek Shale et de Stonington avec des associations palynologiques publi ees d age richmondienne : Maquoketa Shale du Missouri et Kansas (Wright et Myers, 1981 ; Miller, 1991; Wicander et al., 1999), Sylvan Shale de lOklahoma (Loeblich, 1970; Loeblich et MacAdam, 1971 ; Loeblich et Tappan, 1978 ; Playford et Wicander, 2006), et la Formation de Vaur eal de l le dAnticosti, Qu ebec, Canada (Staplin et al., 1965; Jacobson et Achab, 1985).
addition, 32% of the Bills Creek/Stonington assemblage are common to all three of the other Laurentian Richmondian-age formations (Maquoketa Shale, Sylvan Shale, and Vaur eal Formation), and 48% of the Bills Creek/Stonington assemblage occur in at least two of the three aforementioned formations (Fig. 6). Among the noteworthy and consistently occurring Bills Creek/Stonington species that also are found in the Maquoketa, Sylvan, and/or Vaur eal assemblages are: Baltisphaeridium adiastaltum, Dorsennidium hamii, D. undosum, Leiofusa fusiformis, L. litotes, Lophosphaeridium edenense, L. varum, Micrhystridium hirticulum, M. prolixum, Multiplicisphaeridium irregulare, Peteinosphaeridium septuosum, Polygonium gracile, Veryhachium oklahomense, and Villosacapsula setosapellicula. Other Bills Creek/Stonington species of stratigraphic importance, but occurring less commonly, are: Aremoricanium squarrosum, Baltisphaeridium perclarum, Dactylofusa ctenista, Excultibrachium concinnum, Orthosphaeridium rectangulare, and Sylvanidium paucibrachium.
60
Clearly, a high degree of similarity exists among the acritarch assemblages of the coeval Bills Creek Shale/Stonington Formation, Maquoketa Shale, Sylvan Shale, and Vaur eal Formation (Fig. 6). As intimated by Playford and Wicander (2006), and herein conrmed, the occurrence of a large percentage of the acritarchs from these formations collectively constitutes a distinct Laurentian palynostratigraphic entity that facilitates precise regional correlation. In addition, many of these taxa have a global distribution, further corroborating and enhancing intercontinental correlations. 7.2. Extra-Laurentia Numerous publications on palynomorph assemblages from outside of Laurentia include Ashgill acritarchs, but many are based on imprecisely dated and unsatisfactorily documented strata or from erratics, thus hindering reliable correlation and meaningful comparison. Whereas numerous species identied from the Bills Creek Shale and Stonington Formation appear restricted to Laurentia, a signicant number have been reported from elsewhere. Wicander et al. (1999: p. 30, Fig. 16) depicted the global distribution of published Ordovician acritarch occurrences. Most of these are dated as Caradoc, and few, if any, of the component species occur in our Ashgill assemblage. Among the extra-Laurentian Ashgill assemblages, those from northeast Libya (Molyneux, 1988; Hill and Molyneux, 1988), the Algerian Sahara and southern Tunisia (Vecoli, 1999), Morocco (Elaouad-Debbaj, 1988), and Estonia (Uutela and Tynni, 1991) have the most acritarch species in common with the Bills Creek Shale/Stonington Formation palynoora (see Systematic paleontology for sources of data). Those species are discussed in the next section. 8. Palynologic age and biostratigraphy According to Molyneux et al. (1996) and Vecoli and Le H eriss e (2004), Ashgill assemblages are characterized by common to abundant baltisphaerids and netromorphs, together with species of Ordovicidium, Orthosphaeridium, Peteinosphaeridium, and Polygonium. This is exemplied by the Laurentian Ashgill assemblages discussed in this paper. Not only are many of the species found in the Bills Creek Shale, Stonington Formation, Maquoketa Shale, Sylvan Shale, and Vaur eal Formation stratigraphically useful from a regional (Laurentian) perspective, but many also have global biostratigraphic signicance (Fig. 7). Among the species from the Bills Creek/Stonington acritarch assemblage that have been recorded only from Laurentia, and are stratigraphically restricted therein, are the following: Hoegklintia radicosa (middle CaradocAshgill); Baltisphaeridium adiastaltum, Dorsennidium undosum, and Peteinosphaeridium septuosum (upper CaradocAshgill); and Lophosphaeridium acinatum, L. varum, Micrhystridium hirticulum, and Sylvanidium paucibrachium (Ashgill). Essentially cosmopolitan species with restricted stratigraphic ranges include Excultibrachium concinnum and Orthosphaeridium rectangulare, both evidently conned to the CaradocAshgill [i.e., aside from supposed occurrences of
Fig. 7. Chronostratigraphic ranges of previously reported acritarch species occurring in the Bills Creek Shale and Stonington Formation. Ranges are based on what are considered authentic identications. See Systematic paleontology section for sources of data. Arrows indicate range extensions into older (Llanvirn) and younger (Silurian) strata except for Polygonium gracile, which ranges from the Upper Cambrian through Devonian. Fig. 7. Distributions chronostratigraphiques, bas ee sur les donn ees publi ees, des esp` eces dacritarches dans les formations de Stonington et Bills Creek Shale. Voire le texte (Systematic paleontology) pour plus dexplications et les sources ` des donn ees. Les ` eches indiquent lextension vers le Llanvirn ou le Silurien, a lexception de lesp` ece Polygonium gracile, qui a une r epartition stratigraphique allant du Cambrien Sup erieur jusquau D evonien.
E. concinnum in the Llanvirn of Sweden (G orka, 1987) and Estonia (Uutela and Tynni, 1991); and of O. rectangulare in the Llanvirn of Sweden (Kjellstr om, 1971) and Iran (Ghavidel-syooki, 2001, 2003)]. Multiplicisphaeridium irregulare, Veryhachium oklahomense, and Villosacapsula setosapellicula all have a wide geographic distribution and are restricted to the CaradocAshgill; and Aremoricanium squarrosum ranges through the upper CaradocAshgill. Leiofusa litotes is a typical upper CaradocAshgill Laurentian species; however, Vecoli and Le H eriss e (2004) recorded L. cf. L. litotes as being restricted to the Ashgill along the northern Gondwana margin, and Rubinstein and Vaccari (2004) recorded it from Ordovi-
61
cian/Silurian boundary strata in northwest Argentina. Similarly, Lophosphaeridium edenense and Micrhystridium prolixum characterize the upper CaradocAshgill of Laurentia, although both have also been reported from the Chinese Caradoc (Li and Wang, 1997; Li et al., 2006). Baltisphaeridium perclarum is likely conned to the Ashgill, its earlier (supposedly Caradoc) occurrences requiring conrmation. Dactylofusa ctenista is restricted to the Ashgill, not only in Laurentia, but also in northeast Libya (Molyneux, 1988) and northwest Argentina (Rubinstein and Vaccari, 2004). Lastly, Dactylofusa platynetrella is limited to the Ashgill, not only in Laurentia but also in the Algerian Sahara and southern Tunisia (Vecoli, 1999). Based on the taxonomic association and the known stratigraphic ranges (Fig. 7) of the various species recorded from independently dated sections, the Bills Creek Shale and Stonington Formation palynoora is clearly of Ashgill age. 9. Paleogeographic implications During the Late Ordovician, Laurentia was located between ca. 20 north and 20 south of the equator (Fig. 8). The microphytoplankton assemblages of the Bills Creek Shale, Stonington Formation, Maquoketa Shale, Sylvan Shale, and Vaur eal Formation all occur just south of the equator, in roughly the same latitudinal belt (Fig. 8, in which the Bills Creek/Stonington locality is shown with a star symbol). Microphytoplankton provinciality during the Ordovician, particularly the Early and Middle Ordovician, is welldocumented (e.g., Vavrdov a, 1974, 1997; Playford et al., 1995; Tongiorgi et al., 1998; Vecoli, 1999; Li and Servais, 2002;
Servais et al., 2003, 2004; Vecoli and Le H eriss e, 2004; Wicander, 2004). Beginning in at least the late Tremadoc and continuing until the early Caradoc, there was a clear paleobiogeographic differentiation between the so-called Baltic and peri-Gondwana assemblages (Vecoli and Le H eriss e, 2004). A breakdown in microphytoplankton provincialism began during the Caradoc and continued throughout the Ashgill, resulting in greater cosmopolitanism and in assemblages of lesser diversity than pre-Caradoc assemblages (Vecoli and Le H eriss e, 2004). This increase in microphytoplankton cosmopolitanism could have been caused by changes in oceanic currents, particularly as they relate to the conguration of Laurentia, Baltica, Avalonia, and Gondwana during Ashgill time (Wilde, 1991). Fig. 8 illustrates the various geographic occurrences of selected Ashgill organic-walled microphytoplankton assemblages. Whereas the published assemblages from Laurentia constitute a reasonably cohesive entity, many of the constituent taxa are geographically widespread. These include: Aremoricanium squarrosum, Baltisphaeridium perclarum, Dactylofusa ctenista, D. platynetrella, Dorsennidium hamii, Excultibrachium concinnum, Leiofusa fusiformis, L. litotes, Lophosphaeridium edenense, Micrhystridium prolixum, Multiplicisphaeridium irregulare, Orthosphaeridium rectangulare, Polygonium gracile, Veryhachium europaeum, V. oklahomense, V. trispinosum complex, and Villosacapsula setosapellicula. Of the 25 named or cf. species from the Bills Creek Shale and Stonington Formation acritarch assemblage, 17 (68%) also occur elsewhere. It should be noted that many of the Upper Ordovician assemblages from Laurentia and elsewhere have not been described comprehensively, and that preservational
Fig. 8. Geographic occurrences (solid circles) of selected Ashgill organic-walled microphytoplankton assemblages. The star shows the location in Laurentia of the studied sections of the Bills Creek Shale and Stonington Formation. For published records of Upper Ordovician microphytoplankton assemblages, see Wicander et al. (1999: p. 30). Additional localities since Wicander et al. (1999) include Vecoli (1999), Ghavidel-syooki (2001, 2003), Rubinstein and Vaccari (2004), and Li et al. (2006). Paleogeographic reconstruction (450 Ma) based on Li and Powell (2001). NC = North China Block; SC = South China Block. toiles indiquent la position pal Fig. 8. Localisations g eographiques dassemblages palynologiques s electionn ees d age ashgillienne. Les e eog eographique des formations de Stonington et Bills Creek Shale. Pour les sources bibliographiques des assemblages palynologiques de lOrdovicien Sup erieur, voir Wicander et al. (1999 : p. 30), Vecoli (1999), Ghavidel-syooki (2001, 2003), Rubinstein et Vaccari (Rubinstein and Vaccari, 2004), Li et al. (2006). Reconstruction pal eog eographique (450 Ma) dapr` es Li et Powell (2001). NC = North China Block ; SC = South China Block.
62
factors also affect whether some species are considered synonymous. Hence, the reality or degree of cosmopolitanism among CaradocAshgill organic-walled microphytoplankton assemblages cannot yet be regarded as fully comprehended. 10. Summary
France) for critically reviewing the manuscript and making useful suggestions for improvement. This is a contribution to IGCP Project No. 503 Ordovician Palaeogeography and Palaeoclimate, and the phytoPal project, funded by the Leverhulme Trust Research Interchange Grant reference F/00212/F. Appendix A
An abundant, diversied, and well-preserved microphytoplankton assemblage is hosted by the Upper Ordovician Bills Creek Shale and Bay de Noc Member of the Stonington Formation exposed in Michigans Upper Peninsula. The assemblage comprises 29 acritarch species, including the enigmatic palynomorph Gloeocapsomorpha prisca, together with undifferentiated species of the prasinophyte phycomata Leiosphaeridia and Tasmanites. In addition, chitinozoans, scolecodonts, and graptolite fragments are frequent components. Based on published paleontologic information, in conjunction with the stratigraphic ranges of many of the acritarchs, the Bills Creek Shale and Stonington Formation are condently assigned to the North American Richmondian stage of the Cincinnatian series and are correlative with the lower part of the international, British-typied Ashgill interval. The two most abundant acritarchs of the Bills Creek/Stonington assemblage are Leiofusa fusiformis and members of the Veryhachium trispinosum complex. These are commonly associated with Baltisphaeridium adiastaltum, Dorsennidium hamii, Leiofusa litotes, Micrhystridium hirticulum, Peteinosphaeridium septuosum, Polygonium gracile, and Villosacapsula setosapellicula. The combined Bills Creek Shale and Stonington Formation acritarch assemblage shares 76% of its named and cf. species with the Maquoketa Shale of Kansas and Missouri, 72% with the Sylvan Shale of Oklahoma, and 60% with the Vaur eal Formation of Anticosti Island, Qu ebec, Canada. Although exhibiting a distinctive Richmondian Laurentian character, this acritarch assemblage includes many taxa known from Upper Ordovician localities in Baltica, Avalonia, South China, and Gondwana, thus evincing Late Ordovician cosmopolitanism among marine microphytoplankton communities. Paleontologic-palynologic and sedimentologic data indicate that the Bills Creek Shale was deposited in a low-energy, shallow, nearshore, marine environment. The overlying Bay de Noc Member (=lower Stonington Formation) accumulated in a low-energy, offshore, normal marine environment, albeit in somewhat deeper water and farther from the paleoshoreline. Both formations experienced minor transgressions and regressions as indicated by uctuations in the composition of the palynoora. Acknowledgments We thank Geoffrey Williams (Central Michigan University Biology Department Microscopy Facility) for his skilled assistance with scanning electron microscopy, and Josh Perlinski (Central Michigan University) for drafting of text-gures. We also thank Drs. Marco Vecoli and Thomas Servais (Universit e Lille 1, Sciences de la Terre, Laboratoire de Pal eontologie LP3,
A.1. Register of illustrated specimens Catalog numbers are those assigned by the Carnegie Museum of Natural History (Pittsburgh, Pennsylvania, U.S.A.) where the specimens are deposited. Field Finder coordinates refer to the Teledyne Gurley Field Finder, deposited with the gured specimens. For locality and stratigraphic details pertaining to the samples, refer to Figs. 2 and 3.
Species/type Pl./Fig. Sample S6 S4 BC1 Slide >52 m3 SEM S1 >52 m1 >52 m1 >52 m4 >52 m1 >52 m1 >52 m3 2052 m2 >52 m3 2052 m1 2052 m2 SEM S1 >52 m1 >52 m2 SEM S1 >52 m1 >52 m1 2052 m2 2052 m2 >52 m1 >52 m1 >52 m1 >52 m1 2052 m1 SEM S1 Field Finder Q16/3,3 P12/0,3 D15/1,4 M18/1,3 H9/4,1 G10/3,1 W17/2,1 L14/2,3 D17/1,1 P12/3,1 T12/3,2 G4/3,3 P13/4,1 V13/4,1 P4/0,4 Q13/1,5 V7/2,1 M2/1,4 W10/5,0 Z4/2,4 E11/4,0 L15/3,4 H11/1,4 S14/2,2 F11/1,1 P13/5,0 Catalog no. 19086 19087 19088 19089 19090 19091 19092 19093 19094 19095 19096 19097 19098 19099 19100 19101 19102 19103 19104 19105 19106 19107 19108 19109 19110 19111
Leiosphaeridia sp. Hypotype 1/2 Hypotype 1/6 Tasmanites sp. Hypotype 1/1
Aremoricanium squarrosum Hypotype 1/5 S3 Baltisphaeridium adiastaltum Hypotype 1/3 S0 Hypotype 1/7 S4 Baltisphaeridium perclarum Hypotype 1/4 S4 Hypotype 1/8 S3 Dactylofusa ctenista Hypotype 1/9 Hypotype 1/10 S0 BC3
Dactylofusa platynetrella Hypotype 1/11 S5 Hypotype 1/12 S2 Hypotype 1/16, 17 S4 Dorsennidium hamii Hypotype 1/15 Hypotype 1/13 Hypotype 1/14 Dorsennidium undosum Hypotype 1/18 Hypotype 1/19 Elektoriskos sp. A Hypotype 2/5 Hypotype 2/6 Estiastra sp. A Hypotype 1/20 Hypotype 1/21 S4 S4 S4 S2 S2 S0 S0 S4 S6
Excultibrachium concinnum Hypotype 2/1 S4 Hypotype 2/2 S4 Gloeocapsomorpha prisca Hypotype 4/4 S4 Hypotype 4/11 S4
63
References
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Pal aontologische Zeitschrift 13, 74118. Hoegklintia sp. cf. H. radicosa Hypotype 2/3 S5 Hypotype 2/4 S4 Hypotype 2/7 S4 Leiofusa fusiformis Hypotype 4/1 Hypotype 4/2 Hypotype 4/10 Leiofusa litotes Hypotype 3/1 Hypotype 3/2 S5 S0 S4 BC3 S2
Lophosphaeridium acinatum Hypotype 4/3 S0 Lophosphaeridium edenense Hypotype 3/5 S5 Lophosphaeridium varum Hypotype 3/3 S1 Hypotype 3/4 S1 Micrhystridium hirticulum Hypotype 3/8 S2 Hypotype 3/11 S2 Micrhystridium prolixum Hypotype 3/7 S5 Hypotype 3/10 S4 Multiplicisphaeridium irregulare Hypotype 3/6 S4 Navifusa sp. A Hypotype 4/7 Hypotype 4/8 BC4 BC4
Orthosphaeridium rectangulare Hypotype 4/9 S4 Peteinosphaeridium septuosum Hypotype 4/5 S6 Hypotype 4/6 BC1 Hypotype 4/12 S4 Hypotype 4/13 S4 Polygonium gracile Hypotype 4/17 Hypotype 4/18 S5 S4
Sylvanidium paucibrachium Hypotype 4/21 S3 Hypotype 4/22 S4 Veryhachium europaeum Hypotype 3/9 BC1 Veryhachium oklahomense Hypotype 4/14 S0 Hypotype 4/15 S4 Hypotype 4/16 S4 Veryhachium trispinosum Hypotype 4/19 S2 Hypotype 4/23 S4 Villosacapsula setosapellicula Hypotype 4/20 S4 Hypotype 4/24 S4
64
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Revue de micropalontologie 51 (2008) 3966

Corresponding author. E-mail address: reed.wicander@cmich.edu (R. Wicander).

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

R. Wicander, G. Playford / Revue de micropalontologie 51 (2008) 3966

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