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Palaeozoogeography of Western European Lower Jurassic (Pliensbachian and Toarcian) Ostracoda
Geobios 38 (2005) 697–724 http://france.elsevier.com/direct/GEOBIO/
Palaeozoogeography of Western European Lower Jurassic (Pliensbachian and Toarcian) Ostracoda Paléozoogéographie des ostracodes du Jurassique inférieur (Pliensbachien et Toarcien) de l’Ouest de l’Europe Carmen Arias a,*, Robin Whatley b a
Departamento de Paleontología, Facultad de CC Geológicas, Universidad Complutense de Madrid, 28040 Madrid, Spain b Department of Geology, University of Wales, Aberystwyth, Dyfed SY23 3BD, United Kingdom Received 3 January 2002; accepted 28 October 2002 Available online 05 October 2005
Abstract The palaeozoogeography of Lower Jurassic (Pliensbachian-Toarcian) Ostracoda is studied in terms of faunal similarity between 13 geographical areas. A list of 270 marine ostracod species from Europe, North Africa and North America has been compiled and subjected to quantitative analysis (using multivariate methods) in order to ascertain the degree of similarity between Spanish assemblages and those described from these areas. The cluster analysis demonstrates that degree of similarity generally reflects proximity in palaeogeographical position. The results indicate extensive intercommunication of taxa between the basins, and also the absence of clear differentiation between Tethyan and Boreal faunas. © 2005 Elsevier SAS. All rights reserved. Résumé La paléozoogéographie des ostracodes du Jurassique inférieur (Pliensbachien-Toarcien) est étudiée en termes de similitudes fauniques entre 13 zones géographiques. Une liste de 270 espèces d’ostracodes marins de l’Europe du Nord de l’Afrique et d’Amérique du Nord a été compilée et soumise à l’analyse quantitative (méthodes d’analyse multivariée) de façon à déterminer le degré de similitude entre les assemblages d’Espagne et ceux décrits à partir de ces régions. Les classifications hiérarchiques démontrent que le degré de similitude reflète généralement la proximité paléogéographique. Les résultats indiquent une communication très importante des taxons entre les basins et aussi une différentiation claire entre les faunes téthysiennes et boréales. © 2005 Elsevier SAS. All rights reserved. Resumen Este estudio analiza la paleozoografía de los ostrácodos del Jurásico inferior (Pliensbachiense-Toarciense), teniendo en consideración el grado de semejanza faunística entre 13 áreas geográficas. Se ha compilado una lista de 270 especies de ostrácodos marinos descritos en Europa, Africa y Norte América. El resultado ha sido sometido a un ánalisis cuantitativo (usando métodos de análisis multivariante) con el fín de conocer el grado de semejanza entre las asociaciones de ostrácodos españolas y aquellas descritas en dichas áreas. El análisis de tipo cluster demuestra que el grado de semejanza es mayor entre áreas paleogeograficamente próximas. Los resultados muestran una intercomunicación muy amplia de taxones entre las cuencas estudiadas. Además, los resultados de este estudio indican la inesistencia de una clara diferenciación entre fanuas típicas del Tethys y faunas boreales. © 2005 Elsevier SAS. All rights reserved.
* Corresponding author. E-mail address: [email protected] (C. Arias). 0016-6995/$ - see front matter © 2005 Elsevier SAS. All rights reserved. doi:10.1016/j.geobios.2002.10.001
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Keywords: Ostracoda; Palaeobiogeography; Lower Jurassic (Pliensbachian-Toarcian); Cluster-Analysis Mots clés : Ostracodes ; Paléobiogéographie ; Jurassique inférieur (Pliensbachien-Toarcien) ; Classifications hiérarchiques Palabras claves: Ostrácodos; Palaeobiogeografía; Jurásico Inferior (Pliensbaquiense-Toarciense); Análisis de tipo Cluster
1. Introduction Palaeozoogeography is the study of the distribution of animal taxa in space and time, which in most cases is largely a consequence of their ecological requirements. Thus, zoogeography transcends the classical study of the geographical distribution of organisms, involving as it does a range of other factors, including ecological processes, which limit the distribution of species. In an ecological context, zoogeographers have used geographical patterns of distribution to infer the role of such physico-chemical factors as temperature, water depth, O2 concentration, pH, light, etc. Biotic factors, although more difficult to quantify are, notwithstanding, of equal importance. However, in the study of fossils, it is also necessary to embrace an evolutionary dynamic with respect to the phylogenetic history of the taxa concerned, together with evolving palaeogeographies and their consequences in terms of palaeoenvironmental successions; the whole, of course, being complicated by hiatuses and taphonomic processes. The intermittent nature of preservation of strata and outcrop, the wide distribution of Lower Jurassic sediments over the European continent, together with the occurrence of some lithologies highly prejudicial to the preservation or the recovery of Ostracoda, will naturally bias the results of this study to an unknown degree. The number of papers concerning the general distribution of Jurassic marine Ostracoda (Bate, 1977; Lord, 1978, 1982, 1988; Herrig, 1988) is less than for other groups of fossils invertebrates, such as ammonites, brachiopods, bivalves or foraminifers, etc. The present paper has been compiled from a comprehensive literature of ostracod studies for each of the 13 selected geographical areas across Europe. It considers the main composition of their Pliensbachian and Toarcian ostracod assemblages and also their faunal interrelationships in that both similarities and differences are analysed. 2. Provinciality and faunal provinces during the Lower Jurassic In spite of the abundant literature on Jurassic Ostracoda, rather little attention has been paid to their spatial occurrence. This has been not the case with other invertebrate groups. The distribution patterns of such groups as ammonites, brachiopods and bivalves have been used to propose the existence of two major provinces the Northern Hemisphere Lower Jurassic. These are the Boreal and Tethyan provinces (Neumayr, 1882, 1883; Uhlig, 1911; Arkell, 1956; Hallam, 1975, 1983; Enay, 1980; Taylor et al., 1984; Cariou et al, 1985; Damborenea, 1993, etc.).
However, in those studies based on microfossils such provinces have not been so readily recognised. This is the case, for example, with foraminiferal assemblages. Gordon (1970) distinguished five types of foraminiferal assemblages (Gordon, 1970; Exton and Gradstein, 1984). Three are characteristic of shelf seas (composed mainly of Nodosaridae), while the other two are characteristic of the Tethys and its margins (which included planktonic and arenaceous foraminifera). In addition to differences in their geographical distribution (in relation to Boreal and Tethyan ammonite provinces these types of foraminiferal assemblages alternate with each other the same localities and are obviously local ecofacies. The study of the distribution of Northern Hemisphere Mesozoic Ostracoda has allowed the identification of five provinces: European, Tethyan, North African, American and East African (Bate, 1977). Bate indicates that only the first two provinces can be clearly recognised in the Lower Jurassic. The European Province extends from Newfoundland in the west, to the Ukraine in the east and from Greenland in the north to the northern margin of the Tethys in the south. Although, its total inclusion is open to doubt, the Iberian Peninsula is also usually included in this province (Bate, 1977). The Tethyan Province (Bate, 1977; Lord, 1988) extends from the eastern coast of Canada to Northwest Africa (Tunisia, Morocco, Algeria, etc.). For this study the palaeogeographical framework of Ziegler (1988, 1991, 1992) and Bassoullet et al. (1992) has been adopted (Fig. 1). The Lower Jurassic was marked by the development of an extensive epicontinental sea that covered the entire present western European continent. Ziegler (1988) recognized three North European provinces based on facies: Baltic and Polish, North Sea-North Germany and southwestern Europe. The first one is a continental and estuarine, clastic facies province, situated between the Fenno-Scandian High and the Bohemian Massif. The second was an open, colder water marine shale dominated area, which extended from the Norwegian-Greenland Sea into the Netherlands and from the Scottish Highlands into northern Germany. The Southwestern Europe province was a warmer water, carbonate-shale depositional area that extended from the southwest of the Bohemian-Rhenish-London-Brabant massifs to the Irish Massif axis. In the Tethys area, Bassoullet et al. (1992) distinguished carbonate platforms of North Africa (Moroccan High Atlas, Algerian Sahara and Atlas), basinal environment argillaceous sedimentation (Subbetic and Tellian basins) and the pelagic or hemipelagic deposits of the Ammonitico Rosso limestones in the western Tethys.
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Fig. 1. Location of the studied basins. (Palaeogeographical reconstruction after Ziegler, 1988 and Bassoullet et al., 1993). Abbreviations, CI, Cordillera Iberica, DK, Denmark, FB, Fasnet Basin, GB, Grand Banks, DO, Dorset, I, Italy, IL, Ilminster, MA, Morocco, NOD, Northeast Germany, NC, North Celtic Sea, NS, North Sea, NED, Northwest Germany, PB, Paris Basin, S, Sweden, SD, Southwest Germany, Sz, Switzerland, W, Wales, YO, Yorkshire, P, Zambujal. Fig. 1. Localisation des basins étudiés (reconstitution paléogéographique d’après Ziegler, 1988 et Bassoullet et al., 1993).
3. Material and methods The aim of this study is to assess the similarity between the Spanish and other European ostracod assemblages by means of analytical methods based on measures of similarity or dissimilarity. A taxonomic evaluation of published literature was compiled and a faunal list of 270 marine ostracod species was elaborated by combining this data with that added by unpublished information obtained from various authors (Appendix A and B). Since the number of individual localities that have been considered in this paper is large, these sites have been grouped at the level of locally defined geographical areas. These areas are: C: Grand Banks, Canada; CI: the Cordillera Iberica, Spain; DK: Danish Embayment, Denmark and Sweden; FB: the Fasnet Basin and North Celtic basins (Mochras borehole included); GB: British basins (Mochras borehole was not included); It: Italy; M: Morocco; NEG: Northeast Germany; NWG: Northwest Germany; PB: Paris
Basin, France; Q: Quercy, France; SWG: Southwest Germany; Sz: Switzerland; P: the Lusitanian Basin, Portugal (Fig. 1). Due to clearly noticeable differences described between Pliensbachian and Toarcian ostracod assemblages, two data matrices were constructed, one for Pliensbachian and the other for Toarcian assemblages. Having compiled the raw data, the initial data matrix was edited with respect to the occurrence (presence/absence) of taxa with respect to a set of studied localities of the same age. Each similarity matrix was symmetrical and was made in the form of Q-mode (Reyment, 1971, 1979, 1987; Keesey and Whittaker, 1976; Digby and Kempton, 1987). A problem in relation to the use of statistical methods is the wide choice of clustering methods and indices (BaroniUrbani and Buser, 1976). Many authors have suggested differents ways in which these analyses should be carried out (Morisita, 1959; Cheetham and Hazel, 1969; Hohn, 1976; Huhta, 1979; Kenkel and Booth, 1979; Pielou, 1979; Archer
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Table 1 Differences of selected characteristics in the Similarity Indices used in the study Différences des caractères sélectionnés pour les indices de similarités utilisés dans cette étude. Indices (Authors)
Formula
Range
a=0
b=c=0
Independent of d
Affected by sample size
Emphasis
Ignore
Recommended
References
Simpson
a ------a+b
0.1
0
1
yes
little
Useful to measure dissimilarity between samples of different sizes
Absence from both samples
Hazel 1969; Fallaw 1979; Shi 1993
2a ------2a + b + c
0.1
0
1
yes
moderate
Low emphasis in mutual presence
Absence from both samples
Hazel, 1969; Pierce, 1984; Shi, 1993
a ------a+b+c
0
1
yes
much
Presence in both samples
Absence from both samples
Shi, 1993
0.1
Cheetham and Hazel, 1969; Hazel, 1970; Henderson and Heron, 1976; Fallaw, 1979; Archer and Mapples, 1978; Shi, 1993b Cheetham and Hazel, 1969; Hazel, 1970; Archer and Mapples, 1987 Cheetham and Hazel, 1969; Hazel, 1970; Henderson and Heron, 1976; Cambell and Valentine, 1977; Flessa and Miyazaki, 1978; Archer and Mapples, 1987; Shi, 1993
(a + b) < (a + c)
Dice
Jaccard
a = Number of taxa common to both samples. b = Number of taxa restricted to sample 1. c = Number of taxa restricted to sample 2. d = Number of taxa absent from both samples.
and Mapples, 1987, 1989; Mapples and Archer, 1988; Shi, 1993). For the present study, the selection of indices was made by considering those that would satisfy the following conditions: (a) range from 0 (totally different) to 1 (identical) (Goodall, 1973; Lamont and Grant, 1979; Shi, 1993); (b) independent of double absence (absence in both areas) (Goodall, 1973; Henderson and Heron, 1976; Janson and Vegelius, 1981; Hubalek, 1982; Shi, 1993); (c) indices that emphasise presence should be more reliable, because the presence of a taxon in a locality is much less ambiguous than absence (poor sampling, loss through taphonomical process, difficulty with taxonomic identification of taxa, etc.); (d) not susceptible to differences in sample size (Wolda, 1981; Koch, 1987). On the basis of these criteria, three binary similarity coefficients have been chosen which are empirical (non-probabilistic): Simpson, Dice and Jaccard (Table 1). The clustering methodology (Anderberg, 1973; Everitt, 1980; Gordon, 1981; Romesburg, 1984; Podam, 1989) chosen was the agglomerative and hierarchical cluster analysis: Single-Linkage, Complete-Linkage and Average-Linkage (Hazel, 1970; Sneath and Sokal, 1973; Shi and Waterhouse, 1990; Shi, 1993). For this study, the programmes of Microsoft Excel 3.0 for the Macintosh and Ntsys-pc. (Version 1.5) have been used.
4. Results from the cluster analysis 4.1. Pliensbachian Table 2 and Figs. 2–4 summarise the results of the analysis carried out on Pliensbachian taxa. The dendrograms based on the Dice and Jaccard similarity indices have grouped the geographical regions in two major clusters: (1) the Cordillera Iberica-Portugal (CI-P) and (2) Northeast-Southwest Germany (NEG-SWG). The first cluster contains all western areas, with two subclusters: (1a) the Paris Basin-Quercy-the Cordillera Iberica (PB-Q-CI) and (1b) Portugal-the Fasnet Basin (P-FB). The second cluster contains two subclusters that grouped all German and Danish assemblages: (2a) Northwest Germany-Southwest Germany (NWG-SWG) and (2b) Northeast Germany-Denmark (NEG-Dk). Less similar, Swiss, Italian and Canadian assemblages have been joined to the remaining areas as independent geographical areas. Swiss and Italian assemblages show the highest values of similarity with southwestern German and the Paris Basin assemblages. Canadian assemblages show minimum similarity values with all other studied areas, what may be a consequence of their very low diversity. Surprisingly, British assemblages do not cluster with the Fasnet Basin, but form
Table 2 Values of the similarity indices between the Pliensbachian ostracod assemblages, which have been described in the different sample localities former Valeurs des indices de similarités entre les assemblages d’ostracodes du Pliensbachien qui ont été décrites dans les différentes localités échantillonnées auparavant. C. NE. NW Germany Iberica Germany * 0,421052632 0,526315789 * 0,411764706 *
Canada
Denmark
Paris Basin
Quercy
0 0,019607843 0,021276596 0,032258065 *
0,473684211 0,470588235 0,425531915 0,338709677 0,5 *
0,736842105 0,196078431 0,29787234 0,274193548 1 0,266666667 *
0,947368421 0,156862745 0,212765957 0,129032258 0 0,177777778 0,285714286 *
0,197530864 0,407079646 0,550458716 *
0 0,037735849 0,040816327 0,0625 *
0,28125 0,5 0,434782609 0,392523364 0,042553191 *
0,595744681 0,253164557 0,373333333 0,377777778 0,133333333 0,328767123 *
0,109589041 0,255555556 0,379746835 *
0 0,019230769 0,020833333 0,032258065 *
0,163636364 0,333333333 0,277777778 0,244186047 0,02173913 *
0,424242424 0,144927536 0,229508197 0,232876712 0,071428571 0,196721311 *
SIMPSON Pliensbachian
*
0,228571429 0,303030303 * 0,428571429 *
DICE Pliensbachian
*
0,129032258 0,178571429 * 0,272727273 *
JACCARD Pliensbachian
Great Britain 0,421052632 0,196078431 0,276595745 0,209677419 0 0,133333333 0,178571429 0,333333333 *
Italy
Portugal
Switzerland
0,473684211 0,215686275 0,319148936 0,338709677 1 0,377777778 0,607142857 0,259259259 0,366666667 0,1875 0,842105263 *
0,210526316 0,137254902 0,191489362 0,258064516 0,5 0,155555556 0,178571429 0,074074074 0,1 0,1875 0,105263158 0,096774194 *
C. Iberica NE. Germany NW Germany SW Germany Canada Denmark Paris Basin Quercy Great Britain Italy Fastnet Basin Portugal Switzerland
0,782608696 0,205128205 0,27027027 0,179775281 0 0,222222222 0,290909091 *
0,326530612 0,12345679 0,155844156 0,195652174 0 0,186666667 0,24137931 0,137931034 *
0,114285714 0,149253731 0,19047619 0,230769231 0 0,229508197 0,318181818 0,186046512 0,173913043 *
0,285714286 0,27027027 0,371428571 0,305882353 0,08 0,352941176 0,235294118 0,44 0,264150943 0,153846154 *
0,36 0,268292683 0,384615385 0,451612903 0,121212121 0,447368421 0,576271186 0,24137931 0,360655738 0,127659574 0,592592593 *
0,228571429 0,208955224 0,285714286 0,41025641 0,111111111 0,229508197 0,227272727 0,093023256 0,130434783 0,1875 0,102564103 0,127659574 *
C. Iberica NE. Germany NW Germany SW Germany Canada Denmark Paris Basin Quercy Great Britain Italy Fastnet Basin Portugal Switzerland
0,642857143 0,114285714 0,15625 0,098765432 0 0,125 0,170212766 *
0,195121951 0,14084507 0,203125 0,164556962 0 0,086956522 0,094339623 0,1875 *
0,060606061 0,080645161 0,105263158 0,130434783 0 0,12962963 0,189189189 0,102564103 0,095238095 *
0,166666667 0,15625 0,228070175 0,180555556 0,041666667 0,214285714 0,133333333 0,282051282 0,152173913 0,063829787 *
0,219512195 0,154929577 0,238095238 0,291666667 0,064516129 0,288135593 0,404761905 0,137254902 0,22 0,068181818 0,421052632 *
0,129032258 0,116666667 0,166666667 0,258064516 0,058823529 0,12962963 0,128205128 0,048780488 0,069767442 0,103448276 0,054054054 0,068181818 *
C. Iberica NE. Germany NW Germany SW Germany Canada Denmark Paris Basin Quercy Great Britain Italy Fastnet Basin Portugal Switzerland
0,105263158 0,098039216 0,127659574 0,14516129 0 0,155555556 0,25 0,148148148 0,133333333 *
701
Fastnet Basin 0,315789474 0,196078431 0,276595745 0,209677419 0,5 0,266666667 0,214285714 0,407407407 0,233333333 0,1875 *
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SW Germany 0,421052632 0,450980392 0,638297872 *
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Fig. 2. Dendogram from cluster analysis of the Pliensbachian ostracod assemblages of the studied areas. The similarity Index used is Simpson and the clustering techniques are Single-Linkage, Complete-Linkage and AverageLinkage (UPGMA and WPGMA). Abbreviations, C, Grand Banks (Canada), CI, Cordillera Iberica, DK, Denmark, FB, Fasnet Basin, GB, Great Britain, It, Italy, NEG, Northeast Germany, NWG, Northwest Germany, Q, Quercy, PB, Paris Basin, SWG, Southwest Germany, Sz, Switzerland, P, Portugal. Fig. 2. Dendogramme des assemblages d’ostracodes du Pliensbachien des zones étudiées. Indice de similarité utilisé = Indice Simpson–Techniques utilisées = Single-Linkage, average-linkage (UPGMA et WPGMA).
an independent cluster related to other European assemblages. They show especially high similarity values with the Cordillera Iberica. The Simpson index results (Table 2 and Fig. 2) coincide with those obtained from Dice and Jaccard indices, but British assemblages are related to the Cordillera Iberica-Quercy (CI-Q-GB) cluster and Canadian to the Paris Basin-the Fasnet Basin-Portugal cluster (PB-P-FB). The results shows two extra cluster: (1) Cordillera Iberica-Quercy-Great Britain and (2) Canada-the Paris Basin-the Fasnet Basin-Portugal. 4.1.1. Discussion The results show that Spanish assemblages have the highest similarity values with French assemblages; they are more similar to Quercy than to the Paris Basin assemblages. Spanish Pliensbachian assemblages from alternating series of limestones, marly limestones and bioclastic limestones (Calizas y Dolomias Tableadas de Cuevas Labradas, Calizas y Margas de Almonacid de la Cuba, Margas grises del Cerro del Pez and Calizas Bioclásticas de Barahona formations) are dominated by healdiids (Ogmoconchella adenticulata, O. aequalis, O. contractula, O. danica, O. gruendeli, O. propinqua), cytheraceans (Kinkelinella tenuicostata Ektyphocythere aff. E. vitosa, Gramannella apostolescui) and two very abundant species, Liasina lanceolata and Cytherella
Fig. 3. Dendogram from cluster analysis of the Pliensbachian ostracod assemblages of the studied areas. The similarity Index used is Dice and the clustering techniques are Single-Linkage, Complete-Linkage and AverageLinkage (UPGMA and WPGMA). Abbreviations, C, Grand Banks (Canada), CI, Cordillera Iberica, DK, Denmark, FB, Fasnet Basin, GB, Great Britain, It, Italy, NEG, Northeast Germany, NWG, Northwest Germany, Q, Quercy, PB, Paris Basin, SWG, Southwest Germany, Sz, Switzerland, P, Portugal. Fig. 3. Dendogramme des assemblages d’ostracodes du Pliensbachien des zones étudiées. Indice de similarité utilisé = Indice Dice–Techniques utilisées = Single-Linkage, average-linkage (UPGMA et WPGMA).
toarcensis (Arias, 1989, 1991, 1995; Arias and Comas Rengifo, 1992; Arias et al., 1992; Arias and Lord, 1999a, 1999b). Species of the genus Hermiella have not yet been recorded (Fig. 5 and Table 3). Spanish ostracod assemblages share more than 90% of the species with those described in the marly and marly calcareous sediments of the Brian-de-Vere, Valeyres and Barre à Pecten formations of Quercy, southwestern Aquitanian Basin, France (Cubaynes and Ruget, 1985; Bodergat et al., 1991, 1998; Andreu et al., 1998). Their assemblages are dominated by healdiids (Ogmoconchella gruendeli, O. transversa, O. pseudospina and Ogmoconcha gr. amalthei), together with some large cytheraceans (Gramannella apostolescui) and cypridaceans. No species of Hermiella have yet been described either (Fig. 5 and Table 3). The Paris Basin assemblages show a higher similarity to the Iberian Peninsula, with 80% of species in common (Fig. 5 and Table 3). Pliensbachian carbonate and argillaceous sediments from the Paris Basin yielded ostracod assemblages, which are characterised by a very important representation of cytherellids (Cytherelloidea), healdiids (Ogmoconcha, Ogmoconchella, Hermiella and Pseudohealdia), bairdiids and cytheraceans (Ektyphocythere, Kinkelinella and Gramannella). However, the Paris Basin assemblages differ from the
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Fig. 4. Dendogram from cluster analysis of the Pliensbachian ostracod assemblages of the studied areas. The similarity Index used is Jaccard and the clustering techniques are Single-Linkage, Complete-Linkage and AverageLinkage (UPGMA and WPGMA). Abbreviations, C, Grand Banks (Canada), CI, Cordillera Iberica, DK, Denmark, FB, Fasnet Basin, GB, Great Britain, It, Italy, NEG, Northeast Germany, NWG, Northwest Germany, Q, Quercy, PB, Paris Basin, SWG, Southwest Germany, Sz, Switzerland, P, Portugal. Fig. 4. Dendogramme des assemblages d’ostracodes du Pliensbachien des zones étudiées. Indice de similarité utilisé = Indice Jaccard–Techniques utilisées = Single-Linkage, average-linkage (UPGMA et WPGMA).
703
Cordillera Iberica assemblages in the occurrence of Hermiella (Colloque sur le Lias français in 1961, Oertli, 1985; Donze, 1967; Magné and Obert, 1966; Bodergat et al., 1985; Depêche, 1985; Bodergat and Donze, 1988). In the cluster analysis for the Pliensbachian (Figs. 2–4), Portugal was clustered with the Fasnet Basin and both were linked to the Spanish and French assemblages. Ostracod assemblages from the shales, interbedded limestones and bluegrey marls of the Brenha Formation from the Lusitanian Basin (Central Portugal) and the Whale Unit from the Grand Banks of the Newfoundland Basin (Canada), are dominated by cladocopids (Fig. 6), bairdiaceans (Bairdiacypris), healdiids, cytheraceans, cytherellids and Liasina lanceolata (Exton, 1979; Lord, 1982; Exton and Gradstein, 1984; Boomer et al., 1998). The assemblages have over 15% of species common with Spanish and French assemblages (Table 3). Canadian assemblages are characterised by the intermittent preservation of their elements that result in assemblages impoverished in both diversity and abundance. Western and southwestern Ireland basins (Porcupine, Slyne, Erris and Donegal, Fasnet, Celtic Seas and Cardigan Bay) assemblages are clearly related to both former clusters (CI-PB-Q). Pliensbachian sediments are represented by an interbedded shale and siltstone, fine-grained sandstone sequence of the marginal to shallow marine Scalpa Sandstone Formation. To the south, in the Fasnet Basin, this becomes a nearshore marine argillaceous sequence, Croyde and Kikhampton formations (Ainsworth, 1986a, 1986b, 1987; Ainsworth et al., 1987). These sediments yield very rich and diverse ostracod assemblages (Fig. 6) with Bairdiidae and Healdiidae as the dominant components (> 60%), together with many new species of Liasina, Polycope and Procytherura. These assemblages show high commonalties (more
Fig. 5. Number of ostracod species recorded per Lower Jurassic ammonite zone in the Paris Basin and Quercy (France) and Cordillera Iberica (Spain). (Legend as for Fig. 9). Fig. 5. Nombre d’espèces d’ostracodes enregistrées par zone d’ammonite du Jurassique inférieur dans le Bassin de Paris et du Quercy (France) ainsi que dans la Cordillière Ibérique (légende comme pour la Fig. 9).
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Table 3 Distribution of selected ostracod species in the Pliensbachian. Each number represents the sites where these species have been recorded Distribution des espèces d’ostracodes sélectionnées dans le Pliensbachien. Chaque numéro représente les sites où cette espèce a été enregistrée. ostracod species Acrocythere gassumensis Michelsen, 1975 Acrocythere oeresundensis Michelsen, 1975 Acrocythere troesteri Riegraf, 1984 Bairdia clio Bizon, 1960 Bairdia crassa Drexler, 1958 (= B. fortis) Bairdia donzei Herrig, 1979 Bairdia guttulae Herrig, 1979 Bairdia hahni Lord and Moorley, 1974 Bairdia kempfi Ainsworth, 1989 Bairdia michelseni Herrig, 1979 Bairdia molesta Apostolescu, 1959 Bairdia praehilda Herrig, 1979 Bairdia rostrata Issler, 1908 Bairdia thuringica Herrig, 1979 Bairdia sp. A Ohm, 1986 Bairdiacypris anisica brevis Herrig, 1979 Bairdiacypris triasica postera Herrig, 1979 Bairdiacypris tumida Ainsworth, 1987 Bythocypris fabaeformis Drexler, 1958 Cardobairdia liassica (Drexler, 1958) Cardobairdia Nr. 103 Klingler, 1962 Cardobairdia sp. K (Apostolescu, 1959) Cristacythere betzi (Klingler and Neuweiler, 1959) Cythere? terquemiana Jones, 1872 Cytherella lindseyensis Lord, 1974 Cytherella praecadomensis (Knitter and Riegraf, 1984) Cytherella toarcensis Bizon, 1960 Cytherelloidea anningi Lord, 1974 Cytherelloidea drexlerae (Field, 1976) Cytheropteron cavatum Michelsen, 1975 Cytheropteron diversum Herrig, 1969 Cytheropteron foveolatum Michelsen, 1975 Cytheropteron sp. A Riegraf, 1985 Ektyphocythere champeauae (Bizon, 1960) Ektyphocythere luxuriosa (Apostolescu, 1959) Ekt. multicostata (Klingler and Neuweiler, 1959) Ektyphocythere quadrata Boomer and Lord, 1988 Ektyphocythere aff. E. vitiosa (Apostolescu, 1959) Ektyphocythere vulgaris (Klingler and Neuweiler, 1959) Eucytherura liassica (Bate and Coleman, 1975) Eucytherura tricostata (Michelsen, 1975) Fabalicypris symmetrica Herrig, 1979 Gammacythere ubiquita Malz and Lord, 1976 Gramannella apostolescui (Gramann, 1962) Gramannella laevigata Michelsen, 1975 Gramannella tatei (Gramann, 1962) Gramannicythere aubachensis Riegraf, 1984 Gramannicythere bachi Herrig, 1982a Gramannicythere coniuncta Herrig, 1982a Gramannicythere sp. Malz and Nagy, 1984 Hutsonia decorata (AP. Magné and Malmoustier, 1961) Isobythocypris cylindrica (Herrig, 1979) Isobythocypris dorsoconversa Ainsworth, 1986b Isobythocypris fabaeformis Drexler, 1958 Isobythocypris pliensbachiensis Ainsworth, 1986 Isobythocypris unispinata Apostolescu, 1959
1
2
C
3 C C
4
5
C
C
C
C
C
6
7
8
C C C C
C C
C
C
C
9
10
11
12
13
14
15
16
17
C
C
C C
C
C
C
C C
C
C C
C C C C
C C C C
C C C C C
C
C C
C
C C
C
C
C
C C
C
C
C C
C
C
C
C C
C
C
C
C C
C C
C
C
C C
C
C
C
C C
C
C
C
C C C C
C
C
C C
C
C
C
C
C
C
C
C
C C
C
C C C
C C
C
C C C
C C
C C
C C C C C
C
C
C C C C
C C C
C C C
C C C C
C
C
C
C
C
C
C
C C
C C
C C
C
C
C
C C
C
C C
C
C
C C C C C
C C
C
C
C
C
C (continued on next page)
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Table 3 (continued) ostracod species Isobythocypis tatei (Coryell, 1963) Kinkelinella tenuicostata Martin, 1960 Klinglerella elongata Michelsen, 1975 Klinglerella foveolata Michelsen, 1975 Klinglerella herrigi Ainsworth, 1989 Klinglerella intermedia (Klingler and Neuweiler, 1959) Klinglerella lacunosa Ainsworth 1989 Klinglerella katsloesensis Sivhed 1980 Klinglerella moorei (Brady, 1872) Klinglerella variabilis (Klingler and Neuweiler, 1959) Ledahia conveniens Herrig, 1980 Ledahia septenaria (Gründel, 1964) Liasina lanceolata (Apostolescu, 1959) Liasina vestibulifera Gramann, 1962
1
2
3 C
4 C
5 C
6 C
7
8
9
10
11
12
13
C
C C
C C
C
14
15
16
17 C
C
C C C C
C C
C
C
C
C C C C
C C
C C C C
C C C
C C C
C C
C C C
C
C
Key to localities: 1. North Sea, 2. Shane, Sweden, 3. Danish Embayment, 4. NE Germany, 5. NW Germany, 6. SW Germany, 7. Switzerland, 8. Paris Basin, 9. Quercy, 10. Great Britain, 11. Fastnet Basin, 12. Cordillera Iberica, 13. Portugal, 14. Canada, 15. Italy, 16. Australia, 17. Argentina.
than 70%) with Portuguese assemblages and share 31% of common species with the Cordillera Iberica (Fig. 6 and Table 3). Relatively few publications have dealt with British Pliensbachian ostracods. The cluster analysis shows that British assemblages are most similar to those of the Paris Basin and the Cordillera Iberica. The broad belt of Lower Pliensbachian clay-shale and sands deposits from Belemnite Marls, Green Ammonite Beds, Three Tiers, Eype Clay, Down Cliff Sands formations (Howarth, 1978) and the Upper Pliensbachian yellow weathering sands of the Thorncombe Sands and the oolitic limestones of the Marlstone Rock Bed (Lord, 1974; Boomer, 1992) yield assemblages dominated by healdiids (Pseudohealdia bispinosa, Ogmoconchella propinqua, Ogmo-
concha contractula, Ledahia septenaria). Other elements are Liasina lanceolata and some cytheraceans (Gramannella apostolescui, Kinkelinella tenuicostata, Ektyphocythere quadrata and E. champeau) (Fig. 6 and Table 3). The Lower Jurassic sequence of grey mudstones, siltstones and occasional limestones of the Mochras Borehole, on the North Wales coast of Cardigan Bay, yielded an interesting assemblage that differs from the other British assemblages (Boomer, 1991; Boomer and Ballent, 1996) in its large number of cytheraceans (Eucytherura liassica, Cytheropteron cavatum, Gammacythere ubiquita, Gramannella apostolescui and Lophodentina striata). Healdiids (Ogmoconchella adenticulata, Ogmoconcha convexa and O. contractula) are also abundant. Since British assemblages share less than one third of
Fig. 6. Number of ostracod species recorded per Lower Jurassic ammonite zone in Great Britain, Fasnet Basin and Zambujal (Portugal). (Legend as for Fig. 9). Fig. 6. Nombre d’espèces d’ostracodes enregistrées par zone d’ammonite du Jurassique inférieur en Grande-Bretagne, dans le Bassin de Fasnet et à Zambujal (Portugal) (légende comme dans la Fig. 9).
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Fig. 7. Dendogram from cluster analysis of the Toarcian ostracod assemblages of studied areas. The similarity Index used is Dice and the clustering techniques are Single-Linkage, Complete-Linkage and Average-Linkage (UPGMA and WPGMA). Abbreviations, C, Grand Banks (Canada), CI, Cordillera Iberica, FB, Fasnet Basin, GB, Great Britain, Q, Quercy, PB, Paris Basin, SWG, Southwest Germany, Sz, Switzerland, P, Portugal. Fig. 7. Dendogramme des assemblages d’ostracodes du Toarcien des zones étudiées. Indice de similarité utilisé = Indice Dice–Techniques utilisées = Single-Linkage, average-linkage (UPGMA et WPGMA).
Fig. 8. Dendogram from cluster analysis of the Toarcian ostracod assemblages of studied areas. The similarity Index used is Jaccard and the clustering techniques are Single-Linkage, Complete-Linkage and Average-Linkage (UPGMA and WPGMA). Abbreviations, C, Grand Banks (Canada), CI, Cordillera Iberica, FB, Fasnet Basin, GB, Great Britain, Q, Quercy, PB, Paris Basin, SWG, Southwest Germany, Sz, Switzerland, P, Portugal. Fig. 8. Dendogramme des assemblages d’ostracodes du Toarcien des zones étudiées. Indice de similarité utilisé = Indice Jaccard–Techniques utilisées = Single-Linkage, average-linkage (UPGMA et WPGMA).
their species with other areas (Figs. 6–8), they form an independent group under the Dice and Jaccard indices. Using the Simpson Index, British assemblages have been included in the same cluster as the Cordillera Iberica, because Spanish assemblages have almost 40% cent of species common with British assemblages. Ostracod assemblages described from Northeast and Central Europe show a higher similarity, and this is reflected in a major cluster. Danish Embayment and southern Sweden basin assemblages are sparse and poorly preserved. The Pliensbachian Fjerritslev Formation in the Norwegian-Danish basins consists of a relatively uniform series of marine claystone or shale, with a varying silt content, sandstone intercalations and calcareous beds (Michelsen, 1975, 1978). The assemblages share almost 50% species with the Cordillera Iberica (Table 3) and are dominated by healdiids and cytheraceans (cytherurids, bythocytherids), and significant numbers of bairdiaceans and polycopids (Fig. 9). Danish assemblages are also broadly comparable in composition with those described from Germany and Switzerland. The deposition of black shale (Posidonienschiefer facies) during the Sinemurian-Middle Aalenian characterised German and Swiss sedimentation and their ostracod assemblages are well known (Fischer, 1961b; Riegraf et al., 1984). Lower Pliensbachian offshore clay shales of the Schwarzjuratone Beta and Schwarzjuratone Lower Delta formations, together with the Upper Pliensbachian marly limestones with intercalated micritic limestones of the Schwarzjuratone Upper
Delta Formation yielded ostracod assemblages more diverse and abundant than those described in north-western Europe (Fischer, 1961a; Harloff, 1993; Riegraf et al., 1984; Riegraf, 1985; Fischer et al., 1986). They are generally characterised by the occurrence of healdiaceans (including Hermiella), large cytheraceans, cytherurids, bythocytherids, healdiids and bairdiids (Fig. 9) and share more than 40% of species with the Cordillera Iberica (Table 3). 4.2. Toarcian Toarcian ostracod assemblages are represented in the dendrograms (Figs. 7,8,10 and Table 4). They are grouped in two major clusters. They are, from the top to the bottom: (1) Cordillera Iberica to Great Britain (CI-GB) and (2) CanadaPortugal-Fasnet Basin (C-P-FB). In the first cluster, where the majority of the studied assemblages have been included, the highest similarity is observed between southwestern German and Swiss assemblages, with Spanish, French and British assemblages included in the same cluster, although they show less similarity among them in relation to the Pliensbachian. Results obtained with the Jaccard index (Fig. 8) show the highest value of similarity to be between the German and Swiss basins assemblages, which form a cluster that has its maximum similarity with the Paris Basin and British basins assemblages. Canada, Portugal and the Fasnet Basin assemblages form a different cluster that has a rather low similarity with the cluster (CI-SWG-Sz-PB-GB-Q).
C. Arias, R. Whatley / Geobios 38 (2005) 697–724
707
Fig. 9. Number of ostracod species recorded per Lower Jurassic ammonite zone in the Danish Embayment, Northeast Germany and Northwest Germany. Ammonites zones, p, Planorbis, l, Liasicus, a, Angulata, b, Bucklandi, s, Semicostatum, t, Turneri, o, Obtusum, o, Oxynotum, r, Raricostatum, j, Jamesoni, i, Ibex, d, Davoei, m, Margaritatus, s, Spinatum, t, Tenuicostatum, f, Falciferum, b, Bifrons, v, Variabilis, t, Thouarsense, l, Levesquei. Fig. 9. Nombre d’espèces d’ostracodes enregistrées par zone d’ammonite du Jurassique inférieur au nord est et au nord ouest de l’Allemagne.
Dendrograms (Table 4 and Fig. 10) based on the Simpson index contain two major clusters: (1) SW Germany-CanadaSwitzerland (SWG-C-Sz) and (2) the rest of assemblages have been incorporated into the Cordillera Iberica-Fasnet Basin (CI-FB) cluster, which contains two subclusters: (2a) the Cordillera Iberica-Quercy-the Paris Basin (CI-Q-PB) and (2b) Great Britain-Portugal (GB-P). These clusters are different from those obtained when the Dice and Jaccard indices were considered. The different results can be explained in the light of the differences in the size of the compiled samples, particularly, in the case of Canadian and Portuguese assemblages.
Fig. 10. Dendogram from cluster analysis of the Toarcian ostracod assemblages of studied areas. The similarity Index used is Simpson and the clustering techniques are Single-Linkage, Complete-Linkage and AverageLinkage (UPGMA and WPGMA). Abbreviations, C, Grand Banks (Canada), CI, Cordillera Iberica, FB, Fasnet Basin, GB, Great Britain, Q, Quercy, PB, Paris Basin, SWG, Southwest Germany, Sz, Switzerland, P, Portugal. Fig. 10. Dendogramme des assemblages d’ostracodes du Toarcien des zones étudiées. Indice de similarité utilisé = Indice Simpson–Techniques utilisées = Single-Linkage, average-linkage (UPGMA et WPGMA).
4.2.1. Discussion The Lower Toarcian carbonate platform facies of the Cordillera Iberica, developed as an alternating series of marly limestones and marls, formally named the Alternancia de Margas y Calizas de Turmiel Formation (Goy, 1974; Goy et al., 1976; Arche et al., 1977; Comas-Rengifo, 1985), yielded a very rich and diverse ostracod fauna (Fig. 5) dominated by Cytheracea (Arias and Comas-Rengifo, 1992; Arias et al., 1992; Arias and Lord, 1999a, 1999b). The Tenuicostatum Zone and the lower part of the Serpentinus Zone (Falciferum Zone), ostracod assemblages are characterised by the occurrence of Cytherella toarcensis, Liasina lanceolata, Gramannella apostolescui, Ektyphocythere dharennsourensis, E. aff. E. vitiosa, and Ogmoconchella aff. O. aspinata, O. adenticulata and O. aequalis. In the base of the Serpentinus Zone, healdiids disappear, cypridaceans and platycopids continue
708
C. Arias, R. Whatley / Geobios 38 (2005) 697–724
Table 4 Values of the similarity indices between the Toarcian ostracod assemblages, which have been described in the different sample localities former Valeurs des indices de similarités entre les assemblages d’ostracodes du Toarcien qui ont été décrites dans les différentes localités échantillonnées auparavant. C. Iberica
SW Germany
Canada
Paris Basin
Quercy
Great Britain
Fastnet Basin
Portugal
Switzerland
*
0,425925926 *
0,6 1 *
0,675675676 0,648648649 0,135135135 *
0,923076923 0,461538462 0 0,615384615 *
0,575 0,575 0,125 0,525 0,25 *
0,368421053 0,421052632 0,210526316 0,473684211 0,157894737 0,421052632 *
0,631578947 0,578947368 0,263157895 0,578947368 0,263157895 0,631578947 0,421052632 *
0,46728972 0,527472527 0,238095238 *
0,289156627 0,179104478 0 0,32 *
0,418181818 0,489361702 0,222222222 0,545454545 0,377358491 *
0,157303371 0,219178082 0,333333333 0,321428571 0,1875 0,271186441 *
0,269662921 0,301369863 0,416666667 0,392857143 0,3125 0,406779661 0,421052632 *
0,189393939 0,208695652 0,106382979 *
0,126315789 0,082191781 0 0,137931034 *
0,172932331 0,196581197 0,1 0,214285714 0,158730159 *
0,072916667 0,098765432 0,142857143 0,138461538 0,085714286 0,119402985 *
0,118811881 0,130952381 0,172413793 0,164179104 0,135135135 0,169014085 0,173913043 *
0,466666667 0,933333333 0,088888889 0,577777778 0,133333333 0,4 0,155555556 0,222222222 * 0,365217391 0,848484848 0,16 0,634146341 0,206896552 0,423529412 0,28 0,3125 * 0,154411765 0,29787234 0,074074074 0,240740741 0,09375 0,174757282 0,122807018 0,135135135 *
SIMPSON Toarcian *
0,270588235 *
0,074074074 0,169491525 *
DICE Toarcian *
0,156462585 *
0,038461538 0,078125 *
JACCARD Toarcian
decreasing, while the Bairdiacea increase (Bairdiacypris dorisae). In the remainder of the Lower Toarcian, ostracod assemblages are characterised by Cytheracea (Kinkelinella sermoisensis, Ektyphocythere bucki, E. anterocosta, and Praeschuleridea pseudokinkelinella). The Upper Toarcian and Aalenian sediments of the Turmiel and Chelva formations consist of rhythmic alternations of marls and limestones, mudstone and bioclastic wackestone and yield ostracod assemblages comprising many of the Lower Toarcian species plus Cytheropteron alafastigatum, Cytheropterina criba, Kinkelinella fischeri, Ektyphocythere furcata, Praeschuleridea gallemannica, P. ventriosa, Otocythere callosa and Cytherella cadomencis (Goy et al., 1994, 1996). Almost all species cited are common with the other areas of this first (SWG-Sz-PB-GB) cluster (Fig. 5 and Table 5). The Cordillera Iberica assemblages share more species with the Paris Basin (60% of their species) than with those described from Germany, Switzerland and Great Britain. All these areas (CI-PB-SWC-Sz) form a first independent cluster. Spanish assemblages, however, are less similar (share less than 30%) to Canadian and Portuguese assemblages, which form an independent cluster together with the Fasnet Basin assemblages (C-P-FB). Both clusters result when the Dice and Jaccard indices have been used. In contrast with the results described for the Pliensbachian, Toarcian ostracod assemblages recorded from the Quercy area do not show the same
C. Iberica SW Germany Canada Paris Basin Quercy Great Britain Fastnet Basin Portugal Switzerland C. Iberica SW Germany Canada Paris Basin Quercy Great Britain Fastnet Basin Portugal Switzerland C. Iberica SW Germany Canada Paris Basin Quercy Great Britain Fastnet Basin Portugal Switzerland
close similarity to the Cordillera Iberica assemblages and in this way, they are linked to the previous cluster (CI-PB-SWCSz). The reasons may be in the reduced number of species that have been described so far from this area. Results obtained from the cluster analysis show the highest values of similarity between southwestern German and Swiss assemblages. The Lower Toarcian black organic rich clay and interbedded black-shale of the Posidonienschiefer Formation yielded ostracod assemblages dominated by cytheraceans of the genera Kinkelinella, Ektyphocythere, Praeschuleridea, Monoceratina and Trachycythere. The Upper Toarcian ostracod assemblages recorded from the marls, marly limestones and interbedded calcareous shales of the Upper Schwarzjuramergel Formation and for the dark marine shales of the Lower Opalinuston Formation (Knitter and Ohmert, 1983, 1986; Knitter and Riegraf, 1984; Riegraf, 1985; Richter, 1987) are characterised by cytheraceans of the genera Praeschuleridea (P. angulata, P. bernierensis, P. ventriosa) and Kinkelinella (K. sermoisensis, K. costata); Cytherelloidea cadomensis is also well represented (Table 5 and Fig. 11). These assemblages show a high similarity to the Cordillera Iberica, with almost with 40% of common species. The other difference in clustering results between the Dice or Jacard indices and the Simpson index concerns the cluster for southwestern German and Swiss assemblages (SWGSz). These assemblages are linked with Canada when the Sim-
C. Arias, R. Whatley / Geobios 38 (2005) 697–724
709
Table 5 Distribution of selected ostracod species in the Toarcian. Each number represents the sites where these species has been recorded Distribution des espèces d’ostracodes sélectionnées dans le Toarcien. Chaque numéro représente les sites où cette espèce a été enregistrée. SPECIES Bairdia cassiana Bairdia clio Bairdia fortis Bairdia italica Bairdia michelseni Bairdia molesta Bairdia rostrata Bairdiacypris anisica brevis Bairdiacypris dorisae Bairdiacypris trias. postera Cardobairdia liassica Cardobaridia sp. K Cytherella praecadomensis Cytherella toarcensis Cytherelloidea drexlerae Cytheropteron alafastigatum Cytheropteron diversum Ektyphocythere bucki Ektyph. dharennsourensis Ektyphocythere vitiosa Eucytherua liassica Eucytherura spp. Eucytherura tricostata Gen. et sp. D Bate et al. Gramannacyhere ubiquita Hermiella ambo Hermiella comes Hermiella hyblea Isobythocypris cylindrica Isobythocypris tatei Kinkelinella costata Kinkelinella sermoisensis Kinkelinella tenuicostata Liasina lanceolata Monoceratia scrobiculata Monoceratina striata Monoceratina vulsa Monoceratina ungulina Ogmoconcha amalthei Ogmoconcha inflata Ogmoconchella bispinosa Ogmoconchella contractula Paracypris liassica Paracypris redcarensis Polycope cerasia Polycope cincinnata Polycope pelta Pontocyprella cavata Prae. pseudokinkelinella Praeschuleridea ventriosa Pseudohealdia etaulensis Pseudom. subtriangularis Ptychobairdia aselfingensis Ptychob. schaubergensis
LUSITANIA BASIN
GRAND BANKS
SICILY
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MOROCCO
ALGERIA
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Fig. 11. Number of ostracod species recorded per Lower Jurassic ammonite zone in Northwest Germany, Southwest Germany and Switzerland. (Legend as for Fig. 9). Fig. 11. Nombre d’espèces d’ostracodes enregistrées par zone d’ammonite du Jurassique inférieur au nord ouest et au sud ouest de l’Allemagne et en Suisse (légende comme pour la Fig. 9).
pson index is used because Canada shares its entire low diversity fauna with these areas. Thus, the percentage of common species of both central European areas with Canada is higher than with the remaining areas, although the number of common species is very low. However, when the comparison is made with more diverse assemblages, such as, the Paris Basin, Great Britain or the Cordillera Iberica, German ostracod assemblages share a large number of species, although the percentage of common species (total) is smaller than with the Canadian assemblages. This explains why because the southwestern German and Swiss assemblages (SWG-Sz) form a cluster with the Paris Basin, Great Britain and the Cordillera Iberica assemblages with Jaccard or Dice indices, but with Canada with the Simpson index. French assemblages (Fig. 5) from the Paris Basin are closely related to the Cordillera Iberica and Great Britain (CIGB) cluster, and shares more than 30% species. However there is a lack of data for the beginning of the Toarcian. The Spinatum Zone regressive phase is reflected in non-deposition and erosion during the Tenuicostatum Zone, resulting in a barren level. The rest of the Toarcian is characterised in many areas of the Paris Basin (during the Falciferum Zone) by the deposition of marls and marly limestones, together with black shales (″Schistes carton″). Toarcian ostracod assemblages (Cubaynes and Ruget, 1985; Bodergat and Donze, 1988; Bodergat et al., 1985) are less diverse and become dominated by cytheraceans (Ektyphocythere neumannae, E. quadrata and Pleurifera harpa), small cytherurids (species of Cytheropteron, Infracytheropteron, Rutlandella, Eucytherura), cytherellids and bythoceratids after the disappearance of the healdiids at the base of the Toarcian (Table 5 and Fig. 5). They share more than 50% species with the Cordillera Iberica.
Quercy assemblages exhibit a lower level of similarity with the Cordillera Iberica than in the Pliensbachian. Lower Toarcian deposits in the Quercy area (Cubaynes and Faure, 1989; Cubaynes, 1986) comprise marly and marly-calcareous sediments of the Penne and Lexos formations. At the base, green marls and bioclastic limestones of the Barre à Pecten (Upper Pliensbachian-Lower Toarcian) contain only a few species, such as, Ogmoconcha aequalis, Cytherella praecadomensis and Kinkelinella sermoisensis. The Penne Formation includes black-shales of the Schistes carton (Tenuicostatum Zone) and marls and marly limestones (Serpentinus to Insigne ammonite zones) and yield a rich and diverse ostracod fauna (Cubaynes, 1986; Andreu et al., 1995) dominated by large cytheraceans (Kinkelinella, Ektyphocythere, Praeschuleridea, Camptocythere), meanwhile cytherellids and bairdiids (Cytherella toarcensis, C. praecadomensis and Bairdiacypris dorisae) are very common. These assemblages contain a large number of species in common, almost a 75%, with the Cordillera Iberica (Table 5 and Fig. 5). British assemblages show many similarities with the above assemblages, especially with those from the Paris Basin, and have more than 50% species in common with the Cordillera Iberica (Table 5). In the south of England the succession of thin, hard, micritic limestones, separated by non sequences and marked erosion planes in the Lower Toarcian Junction Bed (Hesselbo and Jenkyns, 1995) yielded only a few species of Ogmoconcha, Ogmoconchella and Hermiella, which disappeared at the base of the Falciferum Zone (Boomer, 1992), followed by an increase in the Cytheracea. A better knowledge of the Lower Toarcian ostracod assemblages was obtained from the study of seven boreholes in the East Midlands (Central England) and a section at Ilminster (South-
C. Arias, R. Whatley / Geobios 38 (2005) 697–724
west England). Their assemblages comprised a rich and diverse ostracod fauna (Bate and Coleman, 1975; Boomer, 1992) characterised by the occurrence of the cytheraceans (Kinkelinella sermoisensis, K. costata, Ektyphocythere anterocosta, E. intrepida, E. debilis, Praeschuleridea pseudokinkelinella), bairdiids and cytherellids, showing a large similarity with German and French assemblages and with more than 50% of species common with the Cordillera Iberica (Table 5 and Figs. 7,8,10). Ostracod assemblages from the Grand Banks (Canada), Zambujal (Portugal) and the Fasnet Basin were grouped into one cluster (Figs. 7,8 and 10). Toarcian sediments of the Brenha Formation and the Whale Unit in Zambujal and the Grand Banks are represented by a sequence of shales, marls and limestones, with no black-shale deposition (Mouterde et al., 1971; Exton, 1979; Lord, 1982; Exton and Gradstein, 1984; Boomer et al., 1998) which have yielded ostracod assemblages dominated by species of Bairdiacypris, large cytheraceans and many species of Paracypris. Both assemblages share more than 50% of species with the Cordillera Iberica (Tables 5 and 6). Inner shelf Toarcian-Aalenian dark calcareous bituminous shales of the Stratton Formation in the Fasnet, Celtic and Cardigan Bay basins, yielded less abundant assemblages, where healdiids are absent and large Cytheracea dominate, with many new species of Bairdia, Cardobairdia, Isobythocypris, Bairdiacypris, Liasina and Procytherura (Ainsworth, 1986b; Ainsworth et al., 1987) (Tables 5,6 and Fig. 6). These assemblages have only minor similarity with those described from the Cordillera Iberica, with less than 20% of their ostracod species in common.
5. The distinction of the Tethyan ostracod province Although our knowledge of Tethyan ostracod assemblages has been improved during recent years they are considerably less well known than western European assemblages. In many cases this is due to the small number of species recorded and/or to their poor preservation. Tethyan assemblages have been described from several localities in Italy and North Africa (Barbieri, 1964; Maupin, 1977; Lord (in Farinacci et al., 1978); Lord, 1982, 1988; Baloge, 1981; Boutakiout et al., 1982; Arias, 1993). Italian assemblages are characterised by the occurrence of members of the superfamilies Bairdiacea and Healdiacea, with a lesser representation of Cytheracea (Bate, 1977; Lord, 1988; Arias, 1993). Only a few species recorded in the Ragusa1 borehole (Barbieri, 1964) have been also described in the Cordillera Iberica. In the Umbria-Marche area, northern Apennines, Pliensbachian pelagic sedimentation represented by micritic limestones with chert nodules of the Corniola Formation and clayey and marly sediments of the Marne del Monte Serrone Formation (Farinacci et al., 1978; Colacicchi et al., 1988) yielded assemblages dominated by healdiids, bairdiaceans (Cardobairdia, Bairdiacypris and Bairdia) and heal-
711
diids (Ogmoconchella, Ogmoconcha and Hermiella) and Liasina lanceolata (Lord, 1978 (in Farinacci et al., 1978); Arias, 1993) (Tables 3 and 6). Italian assemblages are more similar to German assemblages than to those situated in the area of the Grand Banks-Portugal and they share more than 50% of their species with Spanish assemblages. Ostracod assemblages from the shallow carbonate platforms of North Africa (Morocco and Algeria) show close affinities with those recorded in southern European basins (Maupin, 1977; Baloge, 1981; Boutakiout et al., 1982; Lord, 1982; Bassoullet et al., 1991). Boutakiout et al. (1982) and Bassoullet et al. (1991) described assemblages from three sections situated at rides sud-rifaines (Atlas Meseta and eastern Moroccan Meseta). Upper Pliensbachian bioclastic limestones and Toarcian grey marls and marly limestones yielded ostracod assemblages characterised by the important occurrence of cytheraceans, platycopids and healdiids, together with some species of the genera Liasina, Bairdia and Bairdiacypris (Table 6). Ostracod assemblages from the western part of the Oman Meseta, have been studied by Maupin (1977). Upper Pliensbachian assemblages are characterised by the occurrence of healdiids (Hermiella ambo, Ogmoconcha amalthei) and Toarcian by species of Bairdia, Cytherella, Isobythocypris, Paracypris, Liasina and some large cytheraceans. These assemblages are very similar to those recorded by Baloge (1981) from the Rosso-Ammonitico of Western Algeria. Upper Pliensbachian assemblages from this area comprise species of Hermiella, Ogmoconchella, Bairdia and Bairdiacypris and Toarcian assemblages, which are of low diversity, are characterized by the lack of healdiids and the occurrence of Bairdiacypris, Cytherella, Polycope and Kinkelinella (Table 6). However, a different record has been obtained from the study of a borehole (DSDP Leg 79, Site 547) located off North Africa (Bate et al., 1984). Eighteen Jurassic samples (from Hettangian to Pliensbachian) yielded a very peculiar fauna, with many new ostracod taxa and with many left in open nomenclature and where smooth taxa dominate (healdiids and bairdiids). As a whole, these assemblages show only a few species in common with previously recorded assemblages from NW Europe. A complete analysis of the Tethyan Lower Jurassic ostracod assemblages (Table 6) reflects a great similarity in composition with those from Central Europe, although with differences in the representation of their taxa. The main differences are that the presence of large Cytheracea are less common than in European assemblages and that ornamented forms of bairdiids and healdiids (Ptychobairdia, Hermiella) are present even during the Toarcian. The present knowledge about Tethyan ostracod assemblages considered here does support the recognition of a separate Tethyan province. 6. Conclusions This paper considers the degree of similarity among European (and surrounding areas) Lower Jurassic Ostracoda
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Table 6 Distribution of the most representative Tethyan ostracod species considered in this study Distribution des espèces téthysiennes d’ostracodes les plus représentatives considérées dans cette étude. ostracod species Monoceratina amlingstadtensis Triebel and Bartenstein, 1938 Monoceratina mesoliassica Triebel and Bartenstein, 1938 Monoceratina michelseni Riegraf, 1984 Monoceratina seebergensis Triebel and Bartenstein, 1938 Monoceratina stimulea (Schwager, 1866) Monoceratina striata Triebel and Bartenstein, 1938 Monoceratina ungulina Triebel and Bartenstein, 1938 Nanacythere (D.) firma Herrig, 1969b Nanacythere (D.) fissicosta Herrig, 1969b Nanacythere (G.) minor Michelsen, 1975 Nanacythere (N.) simplex Herrig, 1969b Ogmoconcha (Hermiella) ambo Lord and Moorley, 1974 Ogmoconcha (Hermiella) cincta Malz, 1975 Ogmoconcha (H.) circumvallata Dreyer, 1967 Ogmoconcha (Hermiella) cista Malz, 1975 Ogmoconcha (Hermiella) comes Malz, 1975 Ogmoconcha (Hermiella) hyblea (Barbieri, 1964) Ogmoconcha (Hermiella) inflata (Ainsworth, 1987) Ogmoconcha (H.) intercedens Dreyer, 1967 Ogmoconcha (Hermiella) klingleri Malz, 1975 Ogmoconcha amalthei (Quenstedt, 1967) Ogmoconcha amalthei rotunda Dreyer, 1967 Ogmoconcha contractula Triebel, 1941 Ogmoconcha convexa Boomer, 1991 Ogmoconcha dentata (Issler, 1908) Ogmoconcha eocontractula Park 1984 Ogmoconchella aenticulata (Pietrzenuk, 1961) Ogmoconchella aequalis (Herrig, 1969a) Ogmoconchella aspinata (Drexler, 1958) Ogmoconchella danica Michelsen, 1975 Ogmoconchella impressa Malz, 1975 Ogmoconchella michelseni (Michelsen, 1975) Omoconchella mouhersensis (Apostolescu, 1959) Ogmoconchella propinqua Malz, 1971 Ogmoconchella secunda Herrig, 1981a Ogmoconchella sp. A Lord, 1974 Ogmoconchella sp. B (Apostolescu, 1959) Oligocythereis? Mochrarensis Boomer, 1991 Paracypris redcarensis (Blake, 1876) Paracypris sp. 1 Exton, 1979 Paracypris sp. A Riegraf, 1985 Pleurifera harpa harpa (Klinger and Neuweiler, 1959) Pleurifera harpa harpoidea (Gramann, 1962) Pleurifera vermiculata (Apostolescu, 1959) Polycope cerasia Blake, 1876 Polycope cincinnata Apostolescu, 1959 Polycope decorata Apostolescu, 1959 Polycope discus Fischer, 1961a Polycope minor Michelsen, 1975 Polycope pelta Fischer, 1961a Polycope plumhoffı Bate and Coleman, 1975 Polycope tenuireticulata (Herrig, 1981c) Progonoidea reticulata (Klingler and Neuweiler, 1959) Pseudohealdia etaulensis (Apostolescu, 1959) Pseudohealdia gruendeli Malz, 1971 Pseudohealdia pseudospina (Herrig, 1969a)
1
2
3 C
4
C
C
C C
C C C C
C C
5 C C C C C C C C C
C
C
C
C
C C
C C C
C C C
C
C
C
C
C
C C C
7
8 C
9
10
11
12
C
13 C
14
15
16
17
C
C
C
C
C
C
C
C
C
C C C
6
C C C C C C
C C C C
C C C C C
C C C C
C C C C C C C C C C C
C C
C C C
C C C
C C
C C
C C C C
C C
C C
C C
C C
C C
C C C
C
C C
C
C C C C C C C C C C
C C C
C
C
C C C
C C C C C C
C C
C C
C C C C C C C
C
C
C C C C
C C
C
C C
C C
C
C C
C
C
C C C
C
C C
C C
C
C
C C
C C
C
C
C
C
C
C
C C
C C
C
C
C C
C C (continued on next page)
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Table 6 (continued) ostracod species Pseudohealdia pseudohealdidae (Gründel, 1964) Pseudohealdia transversa (Gründel, 1964) Pseudohealdia truncata Malz, 1971 Pseudomacr. subtriangularis Michelsen, 1975 Trachycythere angusta Triebel and Klingler, 1959 Trachycythere horrida Triebel and Klingler, 1959 Trachycythere tub. seratina Triebel and Klingler, 1959 Trachycythere tub. tubulosa Triebel and Klingler, 1959 Trachycythere verrucosa Triebel and Klingler, 1959 Trachycythere multistriata Michelsen, 1975 Triassocythere? sp. 4135 (Michelsen, 1975)
1
2
3
4 C
C
7
8 C C
9
10 C C
11
12
C
C C C
6
C
C C C C
5
C C C C C C
C C C C
13
14
15
16 C
17
C C
C C
C C C
C
C
C
Key to localities: 1. North Sea. 2. Shane, Sweden. 3. Danish Embayment. 4. NE Germany. 5. NW Germany. 6. SW Germany. 7. Switzerland. 8. Paris Basin. 9. Quercy. 10. Great Britain. 11. Fastnet Basin. 12. Cordillera Iberica. 13. Portugal. 14. Canada. 15. Italy. 16. Australia. 17. Argentina.
assemblages, with particular attention to their respective relationships with the assemblages of the Cordillera Iberica (Spain). The results indicate that during the Pliensbachian, the Cordillera Iberica assemblages show a higher similarity with French basins (Quercy and the Paris Basin) than those described in the remaining areas of northwestern Europe. Spanish assemblages are situated in the same major cluster that grouped all western European assemblages (French, Portuguese and British). It is remarkable that the grade of similarity among western European assemblages is slightly lower than that which they show individually with Central and northeastern Europe assemblages (Danish and German basins). Canadian, Italian and Swiss assemblages show a low grade of similarity to remaining European assemblages, probably as a consequence of our relatively poor knowledge of the faunas and their poor representation. In the Pliensbachian, the Paris Basin assemblages form a group with the rest of western European assemblages (PB-QCI-GB-P-FB), meanwhile southwestern German basins are within the cluster (NEG-NWG-SWG-Dk) with the nortwestern and Central European assemblages, indicating a first level disjunction. It is noteworthy that northwestern African assemblages are very similar to those described in Central Europe. Showing the occurrence of a open connection between Tethys and the epicontinental sea to the north during the Early Jurassic. For the Toarcian, the cluster analyses show the Cordillera Iberica assemblages have a composition very similar to those of Eastern Europe (German, French and British basins) and their assemblages are included in the same cluster. Meanwhile, Canadian, Fasnet Basin and Portuguese assemblages are grouped together at a high level of similarity, forming another major cluster. Thus, Toarcian Cordillera Iberica assemblages are now more related to the Paris Basin assemblages than they were during the Pliensbachian. Similarly, the Paris Basin assemblages show higher similarity to south-
western German assemblages in the Toarcian than they showed in the Pliensbachian. The results indicate that during the Pliensbachian there was a better communication among western European areas and less between western and Central Europe than during the Toarcian. This latter communication was increased during the Toarcian, meanwhile the most western European areas (the Fasnet Basin, Portugal, Mochras borehole, Canada) show a clear differentiation in relation to the rest of western areas (Iberian Peninsula, French and British Basins). In the Lower Jurassic, especially in the Toarcian, the European ostracod fauna show no clear differentiation into two major biogeographical provinces as do other groups of fossil invertebrates (ammonites, brachiopods). It must be said, however, that Tethyan ostracod assemblages, especially in the post-Pliensbachian, are relatively poorly known.
Acknowledgements The content and presentation of this paper was enhanced by the thoughtful and highly constructive reviews of Prof. A. Lord and Prof. E. Herrig to whom I owe our greatest debts of gratitude. I owe a special debt to gratitude to Dr. Jean Vannier for the critical comments of the early version of our paper and for his patience. I also want to thank, the members of the editorial board of Geobios for their valuable assistance during all stages of this manuscript. C.A. owe special thanks to Fabienne Ongaro for her excellent handling of the numerous misprints and typographical errors. The ostracoda database is based on CA’s ostracod compilation over the past seven years. We acknowledge the TMR Mobility Program (European Union), the Comunidad de Madrid postdoctoral Program and the DGCYT Projects PB93-0459, PB97-0274 and BTE 2001.1924 from the Ministerio de Educación (MEC) y Ciencia and Ministerio de Ciencia y Tecnología of Spain (MCYT), for financial support.
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Annexe A. List of studied localities and references
Pliensbachian Northeastern Germany (NEG)
Northwestern Germany (NWG)
Southwestern Germany (SWG)
Canada (C) Celtic Sea-Fastnet Basin-Porcupine (FB)
Danish Embayment, Denmark (Dk) and Skane, Sweden(S) Paris Basin, France (PB)
Quercy, France (Q)
Great Britain (GB)
Italy (It) Portugal (P)
Northwestern Switzerland (Sz)
Pietrzenuk, 1961 Dreyer, 1967 Herrig, 1969a,b Herrig, 1979 Herrig, 1980 Herrig, 1981a,b,c Herrig, 1982a,b Triebel and Bartenstein, 1938 Triebel and Klingler, 1959 Gramann, 1963 Malz, 1971 Ohm (Fischer et al., 1986) Fischer, 1961b,c Klingler, 1962 Lord and Morley, 1974 Malz, 1975 Urlichs, 1977 Riegraf, 1984, 1985 Harloff, 1993 Harloff and Jagger, 1994 Exton and Gradstein, 1984 Ainsworth and Horton, 1986 Ainsworth, 1987 Ainsworth et al., 1987 Ainsworth, 1990 Michelsen, 1975 Sivhed, 1980 Apostolescu, 1959, 1961 Bizon, 1960, 1961 Apostolescu et al., 1961 Bizon and Oertli, 1961 Cousin and Apostolescu, 1961 Champeau, 1961 Oertli and Grosdidier, 1961 Magné and Malmoustier, 1961 Magné et al., 1961 Maupin, 1978 Donze, 1985 Bodergat et al., 1985 Riegraf, 1985 Bodergat and Donze, 1988 Cubaynes and Ruget, 1985 Andreu et al., 1998 Bodergat et al., 1998 Lord, 1972, 1974 Lord and Bown, 1987 Lord, 1978, 1988 Boomer, 1991 Boomer, 1992 Lord, 1978, 1988 Arias, 1993 Exton, 1979 Exton and Gradstein, 1984 Boomer et al., 1998 Richter, 1987
Dobbertin and Eisenach Brandenburg Greifswald Thüringen Thüringen Thüringen Thüringen Baden (Niedersachten) Hannover Weseker Sattel Niedersachten Empelde Baden-Württemberg Baden-Württemberg Baden-Württemberg Baden-Württemberg Baden-Württemberg Baden-Württemberg Baden-Württemberg Kalk-Alpen Bayern and North Tyrol Grand Banks, Newfoundland Fasnet Basin Fasnet Basin Fasnet Basin Porcupine, Slyne, Evris and Donegal Danish Embayment Southern Sweden Paris Basin Basse Normandie Paris Basin Lorraine Ardennes Southeastern Paris Basin Paris Basin Thouars Thouars Vendée France Paris Basin Truc de Balduc Paris Basin Quercy Quercy Quercy England Dorset and Isle of Wight Great Britain Mochras, Wales Ilminster, Southwestern England Strettura Umbria Zambujal Zambujal Zambujal Basel
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Toarcian Southwestern Germany (SWG)
Canada (C) Celtic Sea-Fastnet Basin-Porcupine (FB)
Paris Basin, France (PB)
Quercy, France (Q)
Great Britain (GB)
Portugal (P)
Northwestern Switzerland (Sz)
Fischer, 1961a,b Knitter, 1983, 1984 Knitter and Riegraf, 1984 Riegraf, 1984, 1985 Harloff, 1993 Exton and Gradstein, 1984 Ainsworth and Horton, 1986 Ainsworth et al., 1987 Ainsworth, 1990 Apostolescu, 1959 Apostolescu, 1961 Champeau, 1961 Magné et al., 1961 Oertli and Grosdidier, 1961 Depêche, 1985 Bodergat et al., 1985 Riegraf, 1985 Bodergat and Donze, 1988 Bodergat et al., 1991 Cubaynes and Ruget, 1985 Cubaynes, 1986 Bodergat et al., 1991 Lord, 1974 Bate and Coleman, 1975 Lord, 1978 Boomer, 1991 Boomer, 1992 Exton, 1979 Exton and Gradstein, 1984 Boomer et al., 1998 Richter, 1987
Annexe B. Alphabetical list of species considered in this study A.1. List of the taxa Acrocythere gassumensis Michelsen, 1975 Acrocythere mimica Bate and Coleman, 1975 Acrocythere oeresundensis Michelsen, 1975 Acrocythere rectangula Michelsen, 1975 Acrocythere reticulata Knitter, 1983 Acrocythere troesteri Riegraf, 1984 Ambigocythere concentricostata Herrig, 1985 Aphelocythere kuhni Triebel and Klingler 1959 Bairdia cariata Drexler, 1958 Bairdia clio Bizon, 1960 Bairdia crassa Drexler, 1958 Bairdia donzei Herrig, 1979 Bairdia fortis Drexler, 1958 Bairdia guttulae Herrig, 1979 Bairdia hahni Lord and Morley, 1974 Bairdia kempfi Ainsworth 1989 Bairdia michelseni Herrig, 1979 Bairdia molesta Apostolescu, 1959 Bairdia ohmerti Knitter, 1983 Bairdia praehilda Herrig, 1979 Bairdia rostrata Issler, 1908 Bairdia sp. A Ainsworth, 1986b
715
Baden-Württemberg Baden-Württemberg Baden-Württemberg Baden-Württemberg Baden-Württemberg Grand Banks, Newfoundland Fasnet Basin Fasnet Basin Porcupine, Slyne, Evris and Donegal Paris Basin South of Paris Basin Southeastern Paris Basin Thouars Paris Basin France Paris Basin Truc de Balduc Paris Basin Paris Basin Quercy Quercy Quercy England Empingham and Upwood Great Britain Mochras, Wales Ilminster Zambujal Zambujal Zambujal Basel
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Bairdia sp. A Ainsworth, 1990 Bairdia sp. A Ohm, 1986 Bairdia thuringica Herrig, 1979 Bairdia undulata Herrig, 1979 Bairdia verrucosa Herrig, 1979 Bairdia? eirensis Ainsworth, 1986a Bairdiacypris dorisae (Knitter, 1984) Bairdiacypris rectangularis Ainsworth, 1986b Bairdiacypris sartriensis Donze, 1966 Bairdiacypris triangularis Ainsworth, 1986b Bairdiacypris triasica postera Herrig, 1979 Bairdiacypris tumida Ainsworth, 1987 Bythocypris fabaeformis Drexler, 1958 Cardobairdia fastnetensis Ainsworth, 1987 Cardobairdia liassica (Drexler, 1958) Cardobairdia Nr. 103 Klingler, 1962 Cardobairdia posteroprolata Ainsworth, 1987 Cardobairdia sp. A Ainsworth, 1987 Cardobairdia sp. K (Apostolescu, 1959) Cardobairdia toarcensis Ainsworth 1986b Cristacythere betzi (Klingler and Neuweiler, 1959) Cristacythere costata Michelsen, 1975 Cristacythere crassireticulata Michelsen, 1975 Cytherella demiexensis Ainsworth, 1989 Cytherella drexlerae Field, 1966 Cytherella lindseyensis Lord, 1974 Cytherella praecadomensis (Knitter and Riegraf, 1984) Cytherella toarcensis Bizon, 1960 Cytherelloidea anningi Lord, 1974 Cytherelloidea cadomensis (Bizon, 1960) Cytherelloidea circumscripta (Blake, 1876) Cytherelloidea drexlerae (Field, 1966) Cytherelloidea lacertosa Apostolescu, 1959 Cytherelloidea modesta Apostolescu, 1959 Cytheropteron alafastigatum Fischer, 1962 Cytheropteron byfieldensis Boomer and Bodergat, 1992 Cytheropteron cavatum Michelsen, 1975 Cytheropteron diversum Herrig, 1969b Cytheropteron foveolatum Michelsen, 1975 Cytheropteron sp. B Riegraf, 1985 Cytheropteron? sp. A Riegraf 1985 Cytheropteron? sp. Boomer, 1991 Ektyphocythere acuminata (Riegraf, 1984) Ektyphocythere ambo Boomer, 1988 Ektyphocythere anterocosta Boomer, 1988 Ektyphocythere bucki (Bizon, 1960) Ektyphocythere champeauae (Bizon, 1960) Ektyphocythere debilis Bate and Coleman, 1975 Ektyphocythere dharennsourensis Boutakiout et al., 1982 Ektyphocythere intrepida Bate and Coleman, 1975 Ektyphocythere knitteri Riegraf, 1984 Ektyphocythere lanceolata Boomer, 1988 Ektyphocythere laqueata (Klingler and Neuweiler, 1959) Ektyphocythere luxuriosa (Apostolescu, 1959) Ektyphocythere multicostata (Klingler, and Neuweiler, 1959) Ektyphocythere neumannae (Maupin, 1978)
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Ektyphocythere quadrata Boomer and Lord, 1988 Ektyphocythere triebeli (Klingler and Neuweiler, 1959) Ektyphocythere vitilis furcata (Wienholz, 1958) Ektyphocythere vitilis vitilis (Apostolescu et al., 1961) Ektyphocythere vitiosa (Apostolescu, 1959) Ektyphocythere vulgaris (Klingler and Neuweiler, 1959) Eocytheropterina jutlandica (Michelsen, 1975) Eucytherura sp. Riegraf, 1985 Fabalicypris symmetrica Herrig, 1979 Gammacythere ubiquita Malz and Lord, 1976 Gen ind. sp. A Boutakiout et al., 1982 Gramannella apostolescui (Gramann, 1962) Gramannella laevigata Michelsen, 1975 Gramannella tatei (Gramann, 1962) Gramannicythere aubachensis Riegraf, 1984 Gramannicythere bachi bachi Herrig, 1982a Gramannicythere bachi ornata Herrig, 1982a Gramannicythere coniuncta Herrig, 1982a Gramannicythere sp. Malz and Nagy, 1989 Hemiparacytheridea (Tuberocytheridea) angulocostata Knitter, 1983 Infracytheropteron gwashense Bate and Coleman, 1975 Infracytheropteron pulchellum Bate and Coleman, 1975 Infracytheropteron rarum Knitter, 1983 Infracytheropteron supraliasicum (Herrig, 1981a) Isobythocypis tatei (Coryell, 1963) Isobythocypris aff. elongata (Blake, 1876) Sivhed, 1980 Isobythocypris cf. elongata (Blake, 1876) Sivhed, 1980 Isobythocypris cf. elongata (Blake, 1876) Drexler, 1958 Isobythocypris cf. elongata (Blake, 1876) Michelsen, 1975 Isobythocypris cylindrica (Herrig, 1979) Isobythocypris dorsoconversa Ainsworth, 1986b Isobythocypris elongata (Blake, 1876) Isobythocypris ovalis Bate and Coleman, 1975 Isobythocypris pliensbachiensis Ainsworth, 1986b Isobythocypris unispinata Apostolescu, 1959 Isobythocypris? sp. Dreyer, 1967 Kinkelinella costata Knitter, 1984 Kinkelinella kadeshensis Rosenfeld et al., 1987 Kinkelinella persica Bate and Coleman, 1975 Kinkelinella procera Herrig, 1985 Kinkelinella sermoisensis (Apostolescu, 1959) Kinkelinella tenuicostata Martin, 1960 Kinkelinella? idrissensis Boutakiout et al., 1982 Klinglerella auleata (Klingler and Neuweiler, 1959) Klinglerella bipartita (Klingler and Neuweiler, 1959) Klinglerella elongata (Michelsen, 1975) Klinglerella foveolata Michelsen, 1975 Klinglerella glabellata (Klingler and Neuweiler, 1959) Klinglerella herrigi Ainsworth, 1989 Klinglerella intermedia (Klingler and Neuweiler, 1959) Klinglerella katsloesensis Sivhed, 1980 Klinglerella lacunosa Ainsworth, 1989 Klinglerella medioreticulata (Michelsen, 1973) Klinglerella moorei (Brady, 1872) Klinglerella sinemuriana Ainsworth, 1989 Klinglerella sulcata (Klingler and Neuweiler, 1959)
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Klinglerella translucens Ainsworth, 1989 Klinglerella variabilis (Klingler and Neuweiler, 1959) Ledahia conviniens Herrig, 1980 Ledahia septenaria (Gründel, 1964) Ledahia telata (Drexler, 1958) Liasina cf. vestibulifera Sivhed 1980 Liasina cylindrica Ainsworth, 1986b Liasina lanceolata (Apostolescu 1959) Liasina vestibulifera Gramann 1963 Monoceratina amlingstadtensis Triebel and Bartenstein, 1938 Monoceratina frentzeni Triebel and Bartenstein, 1938 Monoceratina mesoliassica Triebel and Bartenstein, 1938 Monoceratina michelseni Riegraf, 1984 Monoceratina seebergensis Triebel and Bartenstein, 1938 Monoceratina stimulea (Schwager, 1866) Monoceratina striata Triebel and Bartenstein, 1938 Monoceratina ungulina Triebel and Bartenstein, 1938 Nanacythere (D.) firma Herrig, 1969b Nanacythere (D.) fissicosta Herrig, 1969b Nanacythere (G.) circumcostata Michelsen, 1975 Nanacythere (G.) elegans (Drexler, 1958) Nanacythere (G.) elongata Michelsen, 1975 Nanacythere (G.) minor Michelsen, 1975 Nanacythere (G.) paracostata Michelsen, 1975 Nanacythere (N.) simplex Herrig, 1969b Nanacythere persicaeformis Riegraf, 1984 Ogmoconcha (Hermiella) ambo Lord and Moorley 1974 Ogmoconcha (Hermiella) cincta Malz, 1975 Ogmoconcha (Hermiella) circumvallata Dreyer, 1967 Ogmoconcha (Hermiella) cista Malz, 1975 Ogmoconcha (Hermiella) comes Malz, 1975 Ogmoconcha (Hermiella) hyblea (Barbieri 1964) Ogmoconcha (Hermiella) intercedens Dreyer, 1967 Ogmoconcha (Hermiella) klingleri Malz, 1975 Ogmoconcha amalthei amalthei (Quenstedt, 1967) Ogmoconcha amalthei rotunda Dreyer, 1967 Ogmoconcha contractula Triebel, 1941 Ogmoconcha convexa Boomer, 1991 Ogmoconcha dentata (Issler, 1908) Ogmoconcha eocontractula Park, 1984 Ogmoconcha hagenowi Drexler, 1958 Ogmoconchella adenticulata (Pietrzenuk, 1961) Ogmoconchella aequalis (Herrig, 1969a) Ogmoconchella aspinata (Drexler, 1958) Ogmoconchella celticensis Ainsworth, 1989 Ogmoconchella danica Michelsen, 1975 Ogmoconchella gruendeli Malz, 1971 Ogmoconchella impressa Malz, 1971 Ogmoconchella michelseni Michelsen, 1975 Ogmoconchella mouhersensis (Apostolescu, 1959) Ogmoconchella propinqua Malz, 1971 Ogmoconchella scanica Sivhed, 1977 Ogmoconchella serratostriata Ainsworth, 1989 Ogmoconchella sp. A Lord, 1974 Ogmoconchella sp. B (Apostolescu, 1959) Ogmoconchella sp. B Cubaynes, 1986
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Ogmoconchella transversa (Gründel, 1970) Paracypris liassica (Bate and Coleman, 1975) Paracypris redcarensis (Blake, 1876) Paracypris sp. 1 Exton, 1979 Paracypris sp. 2 Exton, 1979 Paracypris sp. A Riegraf, 1985 Paracypris sp. C Ainsworth, 1986b Paracypris sp. Cubaynes, 1986 Paracypris? longiformis Sivhed, 1980 Paradoxostoma? fusiforme (Drexler, 1958) Paradoxostoma? pulchellum Michelsen, 1975 Paranotacythere (Unicosta) decorata (Apostolescu et al., 1961) Paranotacythere (Unicosta) mochrarensis (Boomer, 1991) Paranotacythere (Unicosta) terquemiana (Jones, 1872) Pleurifera harpa harpa (Klingler and Neuweiler, 1959) Pleurifera harpa harpoidea (Gramann, 1962) Pleurifera perplexa (Klingler and Neuweiler, 1959) Pleurifera plicata (Apostolescu, 1959) Pleurifera vermiculata (Apostolescu, 1959) Polycope cerasia Blake 1876 Polycope cincinnata Apostolescu 1959 Polycope decorata Apostolescu 1959 Polycope discus Fischer 1961a Polycope minor Michelsen 1975 Polycope pelta Fischer 1961a Polycope plumhoffı Bate and Coleman 1975 Polycope tenuireticulata (Herrig, 1981c) Pontocyprella fabaeformis (Drexler, 1958) Praeschuleridea arguta Ainsworth 1986a Praeschuleridea aspera Knitter 1983 Praeschuleridea bernierensis (Apostolescu, 1959) Praeschuleridea costata Ainsworth 1986a Praeschuleridea ellipsoidea Ainsworth 1986b Praeschuleridea gallemannica Malz 1966 Praeschuleridea levita Ainsworth 1986b Praeschuleridea pseudokinkelinella Bate and Coleman 1975 Praeschuleridea punctulata Plumhoff 1963 Praeschuleridea reticulata Ainsworth 1986a Praeschuleridea sp. A Ainsworth 1986b Praeschuleridea ventriosa Plumhoff 1963 Praeschuleridea whatleyi Ainsworth 1986b Praescleridea magnycourtensis (Apostolescu, 1959) Procytheridea laneuvevillae Donze 1967 Procytheridea lotharingiae Apostolescu 1959 Procytheridea multiforata (Klingler and Neuweiler, 1959) Procytheridea perplexa (Klingler and Neuweiler, 1959) Procytheridea spinaecostata (Klingler and Neuweiler, 1959) Procytheridea undulata Apostolescu 1959 Procytheridea? jardensis Maupin 1978 Procytherura celtica Ainsworth 1986b Procytherura euglyphea Ainsworth, 1986b Procytherura hastata Bate and Coleman, 1975 Procytherura mediocostata Bate and Coleman, 1975 Procytherura multicostata Ainsworth, 1986b Procytherura suebica (Herrig and Richter, 1990) Procytherura werneri (Riegraf, 1984)
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Procytherura? exquisita Ainsworth, 1986b Procytherura? liassica Ainsworth, 1986b Procytherura? sp. Boomer 1992 Progonoidea acuticostata (Klingler and Neuweiler, 1959) Progonoidea auleata (Gramann, 1962) Progonoidea polygonata Sivhed 1980 Progonoidea reticulata (Klingler and Neuweiler, 1959) Pseudohealdia etaulensis (Apostolescu, 1959) Pseudohealdia grosdidieri (Viaud, 1963) Pseudohealdia nasuta (Drexler, 1958) Pseudohealdia pseudohealdiae (Gründel, 1964) Pseudohealdia pseudospina (Herrig, 1969a) Pseudohealdia sp. A Ainsworth, 1987 Pseudohealdia transversa (Gründel, 1970) Pseudohealdia truncata Malz 1971 Pseudomacrocypris cf. subtriangularis Michelsen 1975 Pseudomacrocypris subaequalis Michelsen 1975 Pseudomacrocypris? sp. A Ainsworth 1986b Renicytherura liassica (Bate and Coleman, 1975) Renicytherura tricostata (Michelsen, 1975) Rutlandella tatei Ainsworth 1986b Trachycythere tubulosa seratina Triebel and Klingler 1959 Trachycythere tubulosa tubulosa Triebel and Klingler 1959 Trachycythere verrucosa Triebel and Klingler 1959 Triassocythere multiestriata Michelsen, 1975 Triassocythere? sp. 4135 (Michelsen, 1975) Triceratina sp. Boomer 1992 Wicherella semiora kirtonensis Lord 1972 Wicherella semiora semiora Lord 1972 References Ainsworth, N.R., 1986a. Rhaetian, Hettangian and Sinemurian Ostracoda from the Fasnet Basin, offshore Southwest Ireland. Bulletin of the Geological Survey of Ireland 4, 107–150. Ainsworth, N.R., 1986b. Toarcian and Aalenian Ostracoda from the Fasnet Basin, offshore Southwest Ireland. Bulletin of the Geological Survey of Ireland 3, 277–336. Ainsworth, N.R., 1987. Pliensbachian Ostracoda from the Fasnet Basin, offshore Southwest Ireland. Bulletin of the Geological Survey of Ireland 4, 41–62. Ainsworth, N.R., 1989. Remarks on the nomenclature of two ostracod species from the Pliensbachian and upper Toarcian-Aalenian of the Fasnet Basin, offshore Southwest Ireland. Bulletin of the Geological Survey of Ireland 4, 165–166. Ainsworth, N.R., 1990. Uppermost Rhaetian to lower Bajocian Ostracoda from the Porcupine Slyne, Erris and Donegal Basins, offshore west Ireland. Bulletin of the Geological Survey of Ireland 4, 169–200. Ainsworth, N.R., Horton, N.F., 1986. Mesozoic micropaleontology of exploration well Elf 55/30-1 from the Fasnet Basin, offshore southwest Ireland. Journal of Micropaleontology 5, 19–29. Ainsworth, N.R., O’Neill, M., Rutherford, M.M., Clayton, G., Horton, N.F., Penney, R.A., 1987. Biostratigraphy of the Lower Cretaceous, Jurassic and uppermost Triassic of the North Celtic Sea and Fasnet Basin. In: Brooks, J., Glennie, K. (Eds.), Petroleum Geology of North West Europa. Graham and Trotman, pp. 611–622. Anderberg, M.R., 1973. Cluster Analysis for Applications. Academic Press, New York. Andreu, B., Bodergat, A.M., Brunel, F., Colin, J.P., Cubaynes, R., 1998. Ostracodes du Carixien supérieur-Domérien (Jurassique inférieur) du Quercy, Bassin d’Aquitaine, France. Palaeontographica A 250, 68–122.
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Palaeozoogeography of Western European Lower Jurassic (Pliensbachian and Toarcian) Ostracoda Paléozoogéographie des ostracodes du Jurassique inférieur (Pliensbachien et Toarcien) de l’Ouest de l’Europe Carmen Arias a,*, Robin Whatley b a
Departamento de Paleontología, Facultad de CC Geológicas, Universidad Complutense de Madrid, 28040 Madrid, Spain b Department of Geology, University of Wales, Aberystwyth, Dyfed SY23 3BD, United Kingdom Received 3 January 2002; accepted 28 October 2002 Available online 05 October 2005
Abstract The palaeozoogeography of Lower Jurassic (Pliensbachian-Toarcian) Ostracoda is studied in terms of faunal similarity between 13 geographical areas. A list of 270 marine ostracod species from Europe, North Africa and North America has been compiled and subjected to quantitative analysis (using multivariate methods) in order to ascertain the degree of similarity between Spanish assemblages and those described from these areas. The cluster analysis demonstrates that degree of similarity generally reflects proximity in palaeogeographical position. The results indicate extensive intercommunication of taxa between the basins, and also the absence of clear differentiation between Tethyan and Boreal faunas. © 2005 Elsevier SAS. All rights reserved. Résumé La paléozoogéographie des ostracodes du Jurassique inférieur (Pliensbachien-Toarcien) est étudiée en termes de similitudes fauniques entre 13 zones géographiques. Une liste de 270 espèces d’ostracodes marins de l’Europe du Nord de l’Afrique et d’Amérique du Nord a été compilée et soumise à l’analyse quantitative (méthodes d’analyse multivariée) de façon à déterminer le degré de similitude entre les assemblages d’Espagne et ceux décrits à partir de ces régions. Les classifications hiérarchiques démontrent que le degré de similitude reflète généralement la proximité paléogéographique. Les résultats indiquent une communication très importante des taxons entre les basins et aussi une différentiation claire entre les faunes téthysiennes et boréales. © 2005 Elsevier SAS. All rights reserved. Resumen Este estudio analiza la paleozoografía de los ostrácodos del Jurásico inferior (Pliensbachiense-Toarciense), teniendo en consideración el grado de semejanza faunística entre 13 áreas geográficas. Se ha compilado una lista de 270 especies de ostrácodos marinos descritos en Europa, Africa y Norte América. El resultado ha sido sometido a un ánalisis cuantitativo (usando métodos de análisis multivariante) con el fín de conocer el grado de semejanza entre las asociaciones de ostrácodos españolas y aquellas descritas en dichas áreas. El análisis de tipo cluster demuestra que el grado de semejanza es mayor entre áreas paleogeograficamente próximas. Los resultados muestran una intercomunicación muy amplia de taxones entre las cuencas estudiadas. Además, los resultados de este estudio indican la inesistencia de una clara diferenciación entre fanuas típicas del Tethys y faunas boreales. © 2005 Elsevier SAS. All rights reserved.
* Corresponding author. E-mail address: [email protected] (C. Arias). 0016-6995/$ - see front matter © 2005 Elsevier SAS. All rights reserved. doi:10.1016/j.geobios.2002.10.001
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Keywords: Ostracoda; Palaeobiogeography; Lower Jurassic (Pliensbachian-Toarcian); Cluster-Analysis Mots clés : Ostracodes ; Paléobiogéographie ; Jurassique inférieur (Pliensbachien-Toarcien) ; Classifications hiérarchiques Palabras claves: Ostrácodos; Palaeobiogeografía; Jurásico Inferior (Pliensbaquiense-Toarciense); Análisis de tipo Cluster
1. Introduction Palaeozoogeography is the study of the distribution of animal taxa in space and time, which in most cases is largely a consequence of their ecological requirements. Thus, zoogeography transcends the classical study of the geographical distribution of organisms, involving as it does a range of other factors, including ecological processes, which limit the distribution of species. In an ecological context, zoogeographers have used geographical patterns of distribution to infer the role of such physico-chemical factors as temperature, water depth, O2 concentration, pH, light, etc. Biotic factors, although more difficult to quantify are, notwithstanding, of equal importance. However, in the study of fossils, it is also necessary to embrace an evolutionary dynamic with respect to the phylogenetic history of the taxa concerned, together with evolving palaeogeographies and their consequences in terms of palaeoenvironmental successions; the whole, of course, being complicated by hiatuses and taphonomic processes. The intermittent nature of preservation of strata and outcrop, the wide distribution of Lower Jurassic sediments over the European continent, together with the occurrence of some lithologies highly prejudicial to the preservation or the recovery of Ostracoda, will naturally bias the results of this study to an unknown degree. The number of papers concerning the general distribution of Jurassic marine Ostracoda (Bate, 1977; Lord, 1978, 1982, 1988; Herrig, 1988) is less than for other groups of fossils invertebrates, such as ammonites, brachiopods, bivalves or foraminifers, etc. The present paper has been compiled from a comprehensive literature of ostracod studies for each of the 13 selected geographical areas across Europe. It considers the main composition of their Pliensbachian and Toarcian ostracod assemblages and also their faunal interrelationships in that both similarities and differences are analysed. 2. Provinciality and faunal provinces during the Lower Jurassic In spite of the abundant literature on Jurassic Ostracoda, rather little attention has been paid to their spatial occurrence. This has been not the case with other invertebrate groups. The distribution patterns of such groups as ammonites, brachiopods and bivalves have been used to propose the existence of two major provinces the Northern Hemisphere Lower Jurassic. These are the Boreal and Tethyan provinces (Neumayr, 1882, 1883; Uhlig, 1911; Arkell, 1956; Hallam, 1975, 1983; Enay, 1980; Taylor et al., 1984; Cariou et al, 1985; Damborenea, 1993, etc.).
However, in those studies based on microfossils such provinces have not been so readily recognised. This is the case, for example, with foraminiferal assemblages. Gordon (1970) distinguished five types of foraminiferal assemblages (Gordon, 1970; Exton and Gradstein, 1984). Three are characteristic of shelf seas (composed mainly of Nodosaridae), while the other two are characteristic of the Tethys and its margins (which included planktonic and arenaceous foraminifera). In addition to differences in their geographical distribution (in relation to Boreal and Tethyan ammonite provinces these types of foraminiferal assemblages alternate with each other the same localities and are obviously local ecofacies. The study of the distribution of Northern Hemisphere Mesozoic Ostracoda has allowed the identification of five provinces: European, Tethyan, North African, American and East African (Bate, 1977). Bate indicates that only the first two provinces can be clearly recognised in the Lower Jurassic. The European Province extends from Newfoundland in the west, to the Ukraine in the east and from Greenland in the north to the northern margin of the Tethys in the south. Although, its total inclusion is open to doubt, the Iberian Peninsula is also usually included in this province (Bate, 1977). The Tethyan Province (Bate, 1977; Lord, 1988) extends from the eastern coast of Canada to Northwest Africa (Tunisia, Morocco, Algeria, etc.). For this study the palaeogeographical framework of Ziegler (1988, 1991, 1992) and Bassoullet et al. (1992) has been adopted (Fig. 1). The Lower Jurassic was marked by the development of an extensive epicontinental sea that covered the entire present western European continent. Ziegler (1988) recognized three North European provinces based on facies: Baltic and Polish, North Sea-North Germany and southwestern Europe. The first one is a continental and estuarine, clastic facies province, situated between the Fenno-Scandian High and the Bohemian Massif. The second was an open, colder water marine shale dominated area, which extended from the Norwegian-Greenland Sea into the Netherlands and from the Scottish Highlands into northern Germany. The Southwestern Europe province was a warmer water, carbonate-shale depositional area that extended from the southwest of the Bohemian-Rhenish-London-Brabant massifs to the Irish Massif axis. In the Tethys area, Bassoullet et al. (1992) distinguished carbonate platforms of North Africa (Moroccan High Atlas, Algerian Sahara and Atlas), basinal environment argillaceous sedimentation (Subbetic and Tellian basins) and the pelagic or hemipelagic deposits of the Ammonitico Rosso limestones in the western Tethys.
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Fig. 1. Location of the studied basins. (Palaeogeographical reconstruction after Ziegler, 1988 and Bassoullet et al., 1993). Abbreviations, CI, Cordillera Iberica, DK, Denmark, FB, Fasnet Basin, GB, Grand Banks, DO, Dorset, I, Italy, IL, Ilminster, MA, Morocco, NOD, Northeast Germany, NC, North Celtic Sea, NS, North Sea, NED, Northwest Germany, PB, Paris Basin, S, Sweden, SD, Southwest Germany, Sz, Switzerland, W, Wales, YO, Yorkshire, P, Zambujal. Fig. 1. Localisation des basins étudiés (reconstitution paléogéographique d’après Ziegler, 1988 et Bassoullet et al., 1993).
3. Material and methods The aim of this study is to assess the similarity between the Spanish and other European ostracod assemblages by means of analytical methods based on measures of similarity or dissimilarity. A taxonomic evaluation of published literature was compiled and a faunal list of 270 marine ostracod species was elaborated by combining this data with that added by unpublished information obtained from various authors (Appendix A and B). Since the number of individual localities that have been considered in this paper is large, these sites have been grouped at the level of locally defined geographical areas. These areas are: C: Grand Banks, Canada; CI: the Cordillera Iberica, Spain; DK: Danish Embayment, Denmark and Sweden; FB: the Fasnet Basin and North Celtic basins (Mochras borehole included); GB: British basins (Mochras borehole was not included); It: Italy; M: Morocco; NEG: Northeast Germany; NWG: Northwest Germany; PB: Paris
Basin, France; Q: Quercy, France; SWG: Southwest Germany; Sz: Switzerland; P: the Lusitanian Basin, Portugal (Fig. 1). Due to clearly noticeable differences described between Pliensbachian and Toarcian ostracod assemblages, two data matrices were constructed, one for Pliensbachian and the other for Toarcian assemblages. Having compiled the raw data, the initial data matrix was edited with respect to the occurrence (presence/absence) of taxa with respect to a set of studied localities of the same age. Each similarity matrix was symmetrical and was made in the form of Q-mode (Reyment, 1971, 1979, 1987; Keesey and Whittaker, 1976; Digby and Kempton, 1987). A problem in relation to the use of statistical methods is the wide choice of clustering methods and indices (BaroniUrbani and Buser, 1976). Many authors have suggested differents ways in which these analyses should be carried out (Morisita, 1959; Cheetham and Hazel, 1969; Hohn, 1976; Huhta, 1979; Kenkel and Booth, 1979; Pielou, 1979; Archer
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Table 1 Differences of selected characteristics in the Similarity Indices used in the study Différences des caractères sélectionnés pour les indices de similarités utilisés dans cette étude. Indices (Authors)
Formula
Range
a=0
b=c=0
Independent of d
Affected by sample size
Emphasis
Ignore
Recommended
References
Simpson
a ------a+b
0.1
0
1
yes
little
Useful to measure dissimilarity between samples of different sizes
Absence from both samples
Hazel 1969; Fallaw 1979; Shi 1993
2a ------2a + b + c
0.1
0
1
yes
moderate
Low emphasis in mutual presence
Absence from both samples
Hazel, 1969; Pierce, 1984; Shi, 1993
a ------a+b+c
0
1
yes
much
Presence in both samples
Absence from both samples
Shi, 1993
0.1
Cheetham and Hazel, 1969; Hazel, 1970; Henderson and Heron, 1976; Fallaw, 1979; Archer and Mapples, 1978; Shi, 1993b Cheetham and Hazel, 1969; Hazel, 1970; Archer and Mapples, 1987 Cheetham and Hazel, 1969; Hazel, 1970; Henderson and Heron, 1976; Cambell and Valentine, 1977; Flessa and Miyazaki, 1978; Archer and Mapples, 1987; Shi, 1993
(a + b) < (a + c)
Dice
Jaccard
a = Number of taxa common to both samples. b = Number of taxa restricted to sample 1. c = Number of taxa restricted to sample 2. d = Number of taxa absent from both samples.
and Mapples, 1987, 1989; Mapples and Archer, 1988; Shi, 1993). For the present study, the selection of indices was made by considering those that would satisfy the following conditions: (a) range from 0 (totally different) to 1 (identical) (Goodall, 1973; Lamont and Grant, 1979; Shi, 1993); (b) independent of double absence (absence in both areas) (Goodall, 1973; Henderson and Heron, 1976; Janson and Vegelius, 1981; Hubalek, 1982; Shi, 1993); (c) indices that emphasise presence should be more reliable, because the presence of a taxon in a locality is much less ambiguous than absence (poor sampling, loss through taphonomical process, difficulty with taxonomic identification of taxa, etc.); (d) not susceptible to differences in sample size (Wolda, 1981; Koch, 1987). On the basis of these criteria, three binary similarity coefficients have been chosen which are empirical (non-probabilistic): Simpson, Dice and Jaccard (Table 1). The clustering methodology (Anderberg, 1973; Everitt, 1980; Gordon, 1981; Romesburg, 1984; Podam, 1989) chosen was the agglomerative and hierarchical cluster analysis: Single-Linkage, Complete-Linkage and Average-Linkage (Hazel, 1970; Sneath and Sokal, 1973; Shi and Waterhouse, 1990; Shi, 1993). For this study, the programmes of Microsoft Excel 3.0 for the Macintosh and Ntsys-pc. (Version 1.5) have been used.
4. Results from the cluster analysis 4.1. Pliensbachian Table 2 and Figs. 2–4 summarise the results of the analysis carried out on Pliensbachian taxa. The dendrograms based on the Dice and Jaccard similarity indices have grouped the geographical regions in two major clusters: (1) the Cordillera Iberica-Portugal (CI-P) and (2) Northeast-Southwest Germany (NEG-SWG). The first cluster contains all western areas, with two subclusters: (1a) the Paris Basin-Quercy-the Cordillera Iberica (PB-Q-CI) and (1b) Portugal-the Fasnet Basin (P-FB). The second cluster contains two subclusters that grouped all German and Danish assemblages: (2a) Northwest Germany-Southwest Germany (NWG-SWG) and (2b) Northeast Germany-Denmark (NEG-Dk). Less similar, Swiss, Italian and Canadian assemblages have been joined to the remaining areas as independent geographical areas. Swiss and Italian assemblages show the highest values of similarity with southwestern German and the Paris Basin assemblages. Canadian assemblages show minimum similarity values with all other studied areas, what may be a consequence of their very low diversity. Surprisingly, British assemblages do not cluster with the Fasnet Basin, but form
Table 2 Values of the similarity indices between the Pliensbachian ostracod assemblages, which have been described in the different sample localities former Valeurs des indices de similarités entre les assemblages d’ostracodes du Pliensbachien qui ont été décrites dans les différentes localités échantillonnées auparavant. C. NE. NW Germany Iberica Germany * 0,421052632 0,526315789 * 0,411764706 *
Canada
Denmark
Paris Basin
Quercy
0 0,019607843 0,021276596 0,032258065 *
0,473684211 0,470588235 0,425531915 0,338709677 0,5 *
0,736842105 0,196078431 0,29787234 0,274193548 1 0,266666667 *
0,947368421 0,156862745 0,212765957 0,129032258 0 0,177777778 0,285714286 *
0,197530864 0,407079646 0,550458716 *
0 0,037735849 0,040816327 0,0625 *
0,28125 0,5 0,434782609 0,392523364 0,042553191 *
0,595744681 0,253164557 0,373333333 0,377777778 0,133333333 0,328767123 *
0,109589041 0,255555556 0,379746835 *
0 0,019230769 0,020833333 0,032258065 *
0,163636364 0,333333333 0,277777778 0,244186047 0,02173913 *
0,424242424 0,144927536 0,229508197 0,232876712 0,071428571 0,196721311 *
SIMPSON Pliensbachian
*
0,228571429 0,303030303 * 0,428571429 *
DICE Pliensbachian
*
0,129032258 0,178571429 * 0,272727273 *
JACCARD Pliensbachian
Great Britain 0,421052632 0,196078431 0,276595745 0,209677419 0 0,133333333 0,178571429 0,333333333 *
Italy
Portugal
Switzerland
0,473684211 0,215686275 0,319148936 0,338709677 1 0,377777778 0,607142857 0,259259259 0,366666667 0,1875 0,842105263 *
0,210526316 0,137254902 0,191489362 0,258064516 0,5 0,155555556 0,178571429 0,074074074 0,1 0,1875 0,105263158 0,096774194 *
C. Iberica NE. Germany NW Germany SW Germany Canada Denmark Paris Basin Quercy Great Britain Italy Fastnet Basin Portugal Switzerland
0,782608696 0,205128205 0,27027027 0,179775281 0 0,222222222 0,290909091 *
0,326530612 0,12345679 0,155844156 0,195652174 0 0,186666667 0,24137931 0,137931034 *
0,114285714 0,149253731 0,19047619 0,230769231 0 0,229508197 0,318181818 0,186046512 0,173913043 *
0,285714286 0,27027027 0,371428571 0,305882353 0,08 0,352941176 0,235294118 0,44 0,264150943 0,153846154 *
0,36 0,268292683 0,384615385 0,451612903 0,121212121 0,447368421 0,576271186 0,24137931 0,360655738 0,127659574 0,592592593 *
0,228571429 0,208955224 0,285714286 0,41025641 0,111111111 0,229508197 0,227272727 0,093023256 0,130434783 0,1875 0,102564103 0,127659574 *
C. Iberica NE. Germany NW Germany SW Germany Canada Denmark Paris Basin Quercy Great Britain Italy Fastnet Basin Portugal Switzerland
0,642857143 0,114285714 0,15625 0,098765432 0 0,125 0,170212766 *
0,195121951 0,14084507 0,203125 0,164556962 0 0,086956522 0,094339623 0,1875 *
0,060606061 0,080645161 0,105263158 0,130434783 0 0,12962963 0,189189189 0,102564103 0,095238095 *
0,166666667 0,15625 0,228070175 0,180555556 0,041666667 0,214285714 0,133333333 0,282051282 0,152173913 0,063829787 *
0,219512195 0,154929577 0,238095238 0,291666667 0,064516129 0,288135593 0,404761905 0,137254902 0,22 0,068181818 0,421052632 *
0,129032258 0,116666667 0,166666667 0,258064516 0,058823529 0,12962963 0,128205128 0,048780488 0,069767442 0,103448276 0,054054054 0,068181818 *
C. Iberica NE. Germany NW Germany SW Germany Canada Denmark Paris Basin Quercy Great Britain Italy Fastnet Basin Portugal Switzerland
0,105263158 0,098039216 0,127659574 0,14516129 0 0,155555556 0,25 0,148148148 0,133333333 *
701
Fastnet Basin 0,315789474 0,196078431 0,276595745 0,209677419 0,5 0,266666667 0,214285714 0,407407407 0,233333333 0,1875 *
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SW Germany 0,421052632 0,450980392 0,638297872 *
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Fig. 2. Dendogram from cluster analysis of the Pliensbachian ostracod assemblages of the studied areas. The similarity Index used is Simpson and the clustering techniques are Single-Linkage, Complete-Linkage and AverageLinkage (UPGMA and WPGMA). Abbreviations, C, Grand Banks (Canada), CI, Cordillera Iberica, DK, Denmark, FB, Fasnet Basin, GB, Great Britain, It, Italy, NEG, Northeast Germany, NWG, Northwest Germany, Q, Quercy, PB, Paris Basin, SWG, Southwest Germany, Sz, Switzerland, P, Portugal. Fig. 2. Dendogramme des assemblages d’ostracodes du Pliensbachien des zones étudiées. Indice de similarité utilisé = Indice Simpson–Techniques utilisées = Single-Linkage, average-linkage (UPGMA et WPGMA).
an independent cluster related to other European assemblages. They show especially high similarity values with the Cordillera Iberica. The Simpson index results (Table 2 and Fig. 2) coincide with those obtained from Dice and Jaccard indices, but British assemblages are related to the Cordillera Iberica-Quercy (CI-Q-GB) cluster and Canadian to the Paris Basin-the Fasnet Basin-Portugal cluster (PB-P-FB). The results shows two extra cluster: (1) Cordillera Iberica-Quercy-Great Britain and (2) Canada-the Paris Basin-the Fasnet Basin-Portugal. 4.1.1. Discussion The results show that Spanish assemblages have the highest similarity values with French assemblages; they are more similar to Quercy than to the Paris Basin assemblages. Spanish Pliensbachian assemblages from alternating series of limestones, marly limestones and bioclastic limestones (Calizas y Dolomias Tableadas de Cuevas Labradas, Calizas y Margas de Almonacid de la Cuba, Margas grises del Cerro del Pez and Calizas Bioclásticas de Barahona formations) are dominated by healdiids (Ogmoconchella adenticulata, O. aequalis, O. contractula, O. danica, O. gruendeli, O. propinqua), cytheraceans (Kinkelinella tenuicostata Ektyphocythere aff. E. vitosa, Gramannella apostolescui) and two very abundant species, Liasina lanceolata and Cytherella
Fig. 3. Dendogram from cluster analysis of the Pliensbachian ostracod assemblages of the studied areas. The similarity Index used is Dice and the clustering techniques are Single-Linkage, Complete-Linkage and AverageLinkage (UPGMA and WPGMA). Abbreviations, C, Grand Banks (Canada), CI, Cordillera Iberica, DK, Denmark, FB, Fasnet Basin, GB, Great Britain, It, Italy, NEG, Northeast Germany, NWG, Northwest Germany, Q, Quercy, PB, Paris Basin, SWG, Southwest Germany, Sz, Switzerland, P, Portugal. Fig. 3. Dendogramme des assemblages d’ostracodes du Pliensbachien des zones étudiées. Indice de similarité utilisé = Indice Dice–Techniques utilisées = Single-Linkage, average-linkage (UPGMA et WPGMA).
toarcensis (Arias, 1989, 1991, 1995; Arias and Comas Rengifo, 1992; Arias et al., 1992; Arias and Lord, 1999a, 1999b). Species of the genus Hermiella have not yet been recorded (Fig. 5 and Table 3). Spanish ostracod assemblages share more than 90% of the species with those described in the marly and marly calcareous sediments of the Brian-de-Vere, Valeyres and Barre à Pecten formations of Quercy, southwestern Aquitanian Basin, France (Cubaynes and Ruget, 1985; Bodergat et al., 1991, 1998; Andreu et al., 1998). Their assemblages are dominated by healdiids (Ogmoconchella gruendeli, O. transversa, O. pseudospina and Ogmoconcha gr. amalthei), together with some large cytheraceans (Gramannella apostolescui) and cypridaceans. No species of Hermiella have yet been described either (Fig. 5 and Table 3). The Paris Basin assemblages show a higher similarity to the Iberian Peninsula, with 80% of species in common (Fig. 5 and Table 3). Pliensbachian carbonate and argillaceous sediments from the Paris Basin yielded ostracod assemblages, which are characterised by a very important representation of cytherellids (Cytherelloidea), healdiids (Ogmoconcha, Ogmoconchella, Hermiella and Pseudohealdia), bairdiids and cytheraceans (Ektyphocythere, Kinkelinella and Gramannella). However, the Paris Basin assemblages differ from the
C. Arias, R. Whatley / Geobios 38 (2005) 697–724
Fig. 4. Dendogram from cluster analysis of the Pliensbachian ostracod assemblages of the studied areas. The similarity Index used is Jaccard and the clustering techniques are Single-Linkage, Complete-Linkage and AverageLinkage (UPGMA and WPGMA). Abbreviations, C, Grand Banks (Canada), CI, Cordillera Iberica, DK, Denmark, FB, Fasnet Basin, GB, Great Britain, It, Italy, NEG, Northeast Germany, NWG, Northwest Germany, Q, Quercy, PB, Paris Basin, SWG, Southwest Germany, Sz, Switzerland, P, Portugal. Fig. 4. Dendogramme des assemblages d’ostracodes du Pliensbachien des zones étudiées. Indice de similarité utilisé = Indice Jaccard–Techniques utilisées = Single-Linkage, average-linkage (UPGMA et WPGMA).
703
Cordillera Iberica assemblages in the occurrence of Hermiella (Colloque sur le Lias français in 1961, Oertli, 1985; Donze, 1967; Magné and Obert, 1966; Bodergat et al., 1985; Depêche, 1985; Bodergat and Donze, 1988). In the cluster analysis for the Pliensbachian (Figs. 2–4), Portugal was clustered with the Fasnet Basin and both were linked to the Spanish and French assemblages. Ostracod assemblages from the shales, interbedded limestones and bluegrey marls of the Brenha Formation from the Lusitanian Basin (Central Portugal) and the Whale Unit from the Grand Banks of the Newfoundland Basin (Canada), are dominated by cladocopids (Fig. 6), bairdiaceans (Bairdiacypris), healdiids, cytheraceans, cytherellids and Liasina lanceolata (Exton, 1979; Lord, 1982; Exton and Gradstein, 1984; Boomer et al., 1998). The assemblages have over 15% of species common with Spanish and French assemblages (Table 3). Canadian assemblages are characterised by the intermittent preservation of their elements that result in assemblages impoverished in both diversity and abundance. Western and southwestern Ireland basins (Porcupine, Slyne, Erris and Donegal, Fasnet, Celtic Seas and Cardigan Bay) assemblages are clearly related to both former clusters (CI-PB-Q). Pliensbachian sediments are represented by an interbedded shale and siltstone, fine-grained sandstone sequence of the marginal to shallow marine Scalpa Sandstone Formation. To the south, in the Fasnet Basin, this becomes a nearshore marine argillaceous sequence, Croyde and Kikhampton formations (Ainsworth, 1986a, 1986b, 1987; Ainsworth et al., 1987). These sediments yield very rich and diverse ostracod assemblages (Fig. 6) with Bairdiidae and Healdiidae as the dominant components (> 60%), together with many new species of Liasina, Polycope and Procytherura. These assemblages show high commonalties (more
Fig. 5. Number of ostracod species recorded per Lower Jurassic ammonite zone in the Paris Basin and Quercy (France) and Cordillera Iberica (Spain). (Legend as for Fig. 9). Fig. 5. Nombre d’espèces d’ostracodes enregistrées par zone d’ammonite du Jurassique inférieur dans le Bassin de Paris et du Quercy (France) ainsi que dans la Cordillière Ibérique (légende comme pour la Fig. 9).
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Table 3 Distribution of selected ostracod species in the Pliensbachian. Each number represents the sites where these species have been recorded Distribution des espèces d’ostracodes sélectionnées dans le Pliensbachien. Chaque numéro représente les sites où cette espèce a été enregistrée. ostracod species Acrocythere gassumensis Michelsen, 1975 Acrocythere oeresundensis Michelsen, 1975 Acrocythere troesteri Riegraf, 1984 Bairdia clio Bizon, 1960 Bairdia crassa Drexler, 1958 (= B. fortis) Bairdia donzei Herrig, 1979 Bairdia guttulae Herrig, 1979 Bairdia hahni Lord and Moorley, 1974 Bairdia kempfi Ainsworth, 1989 Bairdia michelseni Herrig, 1979 Bairdia molesta Apostolescu, 1959 Bairdia praehilda Herrig, 1979 Bairdia rostrata Issler, 1908 Bairdia thuringica Herrig, 1979 Bairdia sp. A Ohm, 1986 Bairdiacypris anisica brevis Herrig, 1979 Bairdiacypris triasica postera Herrig, 1979 Bairdiacypris tumida Ainsworth, 1987 Bythocypris fabaeformis Drexler, 1958 Cardobairdia liassica (Drexler, 1958) Cardobairdia Nr. 103 Klingler, 1962 Cardobairdia sp. K (Apostolescu, 1959) Cristacythere betzi (Klingler and Neuweiler, 1959) Cythere? terquemiana Jones, 1872 Cytherella lindseyensis Lord, 1974 Cytherella praecadomensis (Knitter and Riegraf, 1984) Cytherella toarcensis Bizon, 1960 Cytherelloidea anningi Lord, 1974 Cytherelloidea drexlerae (Field, 1976) Cytheropteron cavatum Michelsen, 1975 Cytheropteron diversum Herrig, 1969 Cytheropteron foveolatum Michelsen, 1975 Cytheropteron sp. A Riegraf, 1985 Ektyphocythere champeauae (Bizon, 1960) Ektyphocythere luxuriosa (Apostolescu, 1959) Ekt. multicostata (Klingler and Neuweiler, 1959) Ektyphocythere quadrata Boomer and Lord, 1988 Ektyphocythere aff. E. vitiosa (Apostolescu, 1959) Ektyphocythere vulgaris (Klingler and Neuweiler, 1959) Eucytherura liassica (Bate and Coleman, 1975) Eucytherura tricostata (Michelsen, 1975) Fabalicypris symmetrica Herrig, 1979 Gammacythere ubiquita Malz and Lord, 1976 Gramannella apostolescui (Gramann, 1962) Gramannella laevigata Michelsen, 1975 Gramannella tatei (Gramann, 1962) Gramannicythere aubachensis Riegraf, 1984 Gramannicythere bachi Herrig, 1982a Gramannicythere coniuncta Herrig, 1982a Gramannicythere sp. Malz and Nagy, 1984 Hutsonia decorata (AP. Magné and Malmoustier, 1961) Isobythocypris cylindrica (Herrig, 1979) Isobythocypris dorsoconversa Ainsworth, 1986b Isobythocypris fabaeformis Drexler, 1958 Isobythocypris pliensbachiensis Ainsworth, 1986 Isobythocypris unispinata Apostolescu, 1959
1
2
C
3 C C
4
5
C
C
C
C
C
6
7
8
C C C C
C C
C
C
C
9
10
11
12
13
14
15
16
17
C
C
C C
C
C
C
C C
C
C C
C C C C
C C C C
C C C C C
C
C C
C
C C
C
C
C
C C
C
C
C C
C
C
C
C C
C
C
C
C C
C C
C
C
C C
C
C
C
C C
C
C
C
C C C C
C
C
C C
C
C
C
C
C
C
C
C
C C
C
C C C
C C
C
C C C
C C
C C
C C C C C
C
C
C C C C
C C C
C C C
C C C C
C
C
C
C
C
C
C
C C
C C
C C
C
C
C
C C
C
C C
C
C
C C C C C
C C
C
C
C
C
C (continued on next page)
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Table 3 (continued) ostracod species Isobythocypis tatei (Coryell, 1963) Kinkelinella tenuicostata Martin, 1960 Klinglerella elongata Michelsen, 1975 Klinglerella foveolata Michelsen, 1975 Klinglerella herrigi Ainsworth, 1989 Klinglerella intermedia (Klingler and Neuweiler, 1959) Klinglerella lacunosa Ainsworth 1989 Klinglerella katsloesensis Sivhed 1980 Klinglerella moorei (Brady, 1872) Klinglerella variabilis (Klingler and Neuweiler, 1959) Ledahia conveniens Herrig, 1980 Ledahia septenaria (Gründel, 1964) Liasina lanceolata (Apostolescu, 1959) Liasina vestibulifera Gramann, 1962
1
2
3 C
4 C
5 C
6 C
7
8
9
10
11
12
13
C
C C
C C
C
14
15
16
17 C
C
C C C C
C C
C
C
C
C C C C
C C
C C C C
C C C
C C C
C C
C C C
C
C
Key to localities: 1. North Sea, 2. Shane, Sweden, 3. Danish Embayment, 4. NE Germany, 5. NW Germany, 6. SW Germany, 7. Switzerland, 8. Paris Basin, 9. Quercy, 10. Great Britain, 11. Fastnet Basin, 12. Cordillera Iberica, 13. Portugal, 14. Canada, 15. Italy, 16. Australia, 17. Argentina.
than 70%) with Portuguese assemblages and share 31% of common species with the Cordillera Iberica (Fig. 6 and Table 3). Relatively few publications have dealt with British Pliensbachian ostracods. The cluster analysis shows that British assemblages are most similar to those of the Paris Basin and the Cordillera Iberica. The broad belt of Lower Pliensbachian clay-shale and sands deposits from Belemnite Marls, Green Ammonite Beds, Three Tiers, Eype Clay, Down Cliff Sands formations (Howarth, 1978) and the Upper Pliensbachian yellow weathering sands of the Thorncombe Sands and the oolitic limestones of the Marlstone Rock Bed (Lord, 1974; Boomer, 1992) yield assemblages dominated by healdiids (Pseudohealdia bispinosa, Ogmoconchella propinqua, Ogmo-
concha contractula, Ledahia septenaria). Other elements are Liasina lanceolata and some cytheraceans (Gramannella apostolescui, Kinkelinella tenuicostata, Ektyphocythere quadrata and E. champeau) (Fig. 6 and Table 3). The Lower Jurassic sequence of grey mudstones, siltstones and occasional limestones of the Mochras Borehole, on the North Wales coast of Cardigan Bay, yielded an interesting assemblage that differs from the other British assemblages (Boomer, 1991; Boomer and Ballent, 1996) in its large number of cytheraceans (Eucytherura liassica, Cytheropteron cavatum, Gammacythere ubiquita, Gramannella apostolescui and Lophodentina striata). Healdiids (Ogmoconchella adenticulata, Ogmoconcha convexa and O. contractula) are also abundant. Since British assemblages share less than one third of
Fig. 6. Number of ostracod species recorded per Lower Jurassic ammonite zone in Great Britain, Fasnet Basin and Zambujal (Portugal). (Legend as for Fig. 9). Fig. 6. Nombre d’espèces d’ostracodes enregistrées par zone d’ammonite du Jurassique inférieur en Grande-Bretagne, dans le Bassin de Fasnet et à Zambujal (Portugal) (légende comme dans la Fig. 9).
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Fig. 7. Dendogram from cluster analysis of the Toarcian ostracod assemblages of studied areas. The similarity Index used is Dice and the clustering techniques are Single-Linkage, Complete-Linkage and Average-Linkage (UPGMA and WPGMA). Abbreviations, C, Grand Banks (Canada), CI, Cordillera Iberica, FB, Fasnet Basin, GB, Great Britain, Q, Quercy, PB, Paris Basin, SWG, Southwest Germany, Sz, Switzerland, P, Portugal. Fig. 7. Dendogramme des assemblages d’ostracodes du Toarcien des zones étudiées. Indice de similarité utilisé = Indice Dice–Techniques utilisées = Single-Linkage, average-linkage (UPGMA et WPGMA).
Fig. 8. Dendogram from cluster analysis of the Toarcian ostracod assemblages of studied areas. The similarity Index used is Jaccard and the clustering techniques are Single-Linkage, Complete-Linkage and Average-Linkage (UPGMA and WPGMA). Abbreviations, C, Grand Banks (Canada), CI, Cordillera Iberica, FB, Fasnet Basin, GB, Great Britain, Q, Quercy, PB, Paris Basin, SWG, Southwest Germany, Sz, Switzerland, P, Portugal. Fig. 8. Dendogramme des assemblages d’ostracodes du Toarcien des zones étudiées. Indice de similarité utilisé = Indice Jaccard–Techniques utilisées = Single-Linkage, average-linkage (UPGMA et WPGMA).
their species with other areas (Figs. 6–8), they form an independent group under the Dice and Jaccard indices. Using the Simpson Index, British assemblages have been included in the same cluster as the Cordillera Iberica, because Spanish assemblages have almost 40% cent of species common with British assemblages. Ostracod assemblages described from Northeast and Central Europe show a higher similarity, and this is reflected in a major cluster. Danish Embayment and southern Sweden basin assemblages are sparse and poorly preserved. The Pliensbachian Fjerritslev Formation in the Norwegian-Danish basins consists of a relatively uniform series of marine claystone or shale, with a varying silt content, sandstone intercalations and calcareous beds (Michelsen, 1975, 1978). The assemblages share almost 50% species with the Cordillera Iberica (Table 3) and are dominated by healdiids and cytheraceans (cytherurids, bythocytherids), and significant numbers of bairdiaceans and polycopids (Fig. 9). Danish assemblages are also broadly comparable in composition with those described from Germany and Switzerland. The deposition of black shale (Posidonienschiefer facies) during the Sinemurian-Middle Aalenian characterised German and Swiss sedimentation and their ostracod assemblages are well known (Fischer, 1961b; Riegraf et al., 1984). Lower Pliensbachian offshore clay shales of the Schwarzjuratone Beta and Schwarzjuratone Lower Delta formations, together with the Upper Pliensbachian marly limestones with intercalated micritic limestones of the Schwarzjuratone Upper
Delta Formation yielded ostracod assemblages more diverse and abundant than those described in north-western Europe (Fischer, 1961a; Harloff, 1993; Riegraf et al., 1984; Riegraf, 1985; Fischer et al., 1986). They are generally characterised by the occurrence of healdiaceans (including Hermiella), large cytheraceans, cytherurids, bythocytherids, healdiids and bairdiids (Fig. 9) and share more than 40% of species with the Cordillera Iberica (Table 3). 4.2. Toarcian Toarcian ostracod assemblages are represented in the dendrograms (Figs. 7,8,10 and Table 4). They are grouped in two major clusters. They are, from the top to the bottom: (1) Cordillera Iberica to Great Britain (CI-GB) and (2) CanadaPortugal-Fasnet Basin (C-P-FB). In the first cluster, where the majority of the studied assemblages have been included, the highest similarity is observed between southwestern German and Swiss assemblages, with Spanish, French and British assemblages included in the same cluster, although they show less similarity among them in relation to the Pliensbachian. Results obtained with the Jaccard index (Fig. 8) show the highest value of similarity to be between the German and Swiss basins assemblages, which form a cluster that has its maximum similarity with the Paris Basin and British basins assemblages. Canada, Portugal and the Fasnet Basin assemblages form a different cluster that has a rather low similarity with the cluster (CI-SWG-Sz-PB-GB-Q).
C. Arias, R. Whatley / Geobios 38 (2005) 697–724
707
Fig. 9. Number of ostracod species recorded per Lower Jurassic ammonite zone in the Danish Embayment, Northeast Germany and Northwest Germany. Ammonites zones, p, Planorbis, l, Liasicus, a, Angulata, b, Bucklandi, s, Semicostatum, t, Turneri, o, Obtusum, o, Oxynotum, r, Raricostatum, j, Jamesoni, i, Ibex, d, Davoei, m, Margaritatus, s, Spinatum, t, Tenuicostatum, f, Falciferum, b, Bifrons, v, Variabilis, t, Thouarsense, l, Levesquei. Fig. 9. Nombre d’espèces d’ostracodes enregistrées par zone d’ammonite du Jurassique inférieur au nord est et au nord ouest de l’Allemagne.
Dendrograms (Table 4 and Fig. 10) based on the Simpson index contain two major clusters: (1) SW Germany-CanadaSwitzerland (SWG-C-Sz) and (2) the rest of assemblages have been incorporated into the Cordillera Iberica-Fasnet Basin (CI-FB) cluster, which contains two subclusters: (2a) the Cordillera Iberica-Quercy-the Paris Basin (CI-Q-PB) and (2b) Great Britain-Portugal (GB-P). These clusters are different from those obtained when the Dice and Jaccard indices were considered. The different results can be explained in the light of the differences in the size of the compiled samples, particularly, in the case of Canadian and Portuguese assemblages.
Fig. 10. Dendogram from cluster analysis of the Toarcian ostracod assemblages of studied areas. The similarity Index used is Simpson and the clustering techniques are Single-Linkage, Complete-Linkage and AverageLinkage (UPGMA and WPGMA). Abbreviations, C, Grand Banks (Canada), CI, Cordillera Iberica, FB, Fasnet Basin, GB, Great Britain, Q, Quercy, PB, Paris Basin, SWG, Southwest Germany, Sz, Switzerland, P, Portugal. Fig. 10. Dendogramme des assemblages d’ostracodes du Toarcien des zones étudiées. Indice de similarité utilisé = Indice Simpson–Techniques utilisées = Single-Linkage, average-linkage (UPGMA et WPGMA).
4.2.1. Discussion The Lower Toarcian carbonate platform facies of the Cordillera Iberica, developed as an alternating series of marly limestones and marls, formally named the Alternancia de Margas y Calizas de Turmiel Formation (Goy, 1974; Goy et al., 1976; Arche et al., 1977; Comas-Rengifo, 1985), yielded a very rich and diverse ostracod fauna (Fig. 5) dominated by Cytheracea (Arias and Comas-Rengifo, 1992; Arias et al., 1992; Arias and Lord, 1999a, 1999b). The Tenuicostatum Zone and the lower part of the Serpentinus Zone (Falciferum Zone), ostracod assemblages are characterised by the occurrence of Cytherella toarcensis, Liasina lanceolata, Gramannella apostolescui, Ektyphocythere dharennsourensis, E. aff. E. vitiosa, and Ogmoconchella aff. O. aspinata, O. adenticulata and O. aequalis. In the base of the Serpentinus Zone, healdiids disappear, cypridaceans and platycopids continue
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Table 4 Values of the similarity indices between the Toarcian ostracod assemblages, which have been described in the different sample localities former Valeurs des indices de similarités entre les assemblages d’ostracodes du Toarcien qui ont été décrites dans les différentes localités échantillonnées auparavant. C. Iberica
SW Germany
Canada
Paris Basin
Quercy
Great Britain
Fastnet Basin
Portugal
Switzerland
*
0,425925926 *
0,6 1 *
0,675675676 0,648648649 0,135135135 *
0,923076923 0,461538462 0 0,615384615 *
0,575 0,575 0,125 0,525 0,25 *
0,368421053 0,421052632 0,210526316 0,473684211 0,157894737 0,421052632 *
0,631578947 0,578947368 0,263157895 0,578947368 0,263157895 0,631578947 0,421052632 *
0,46728972 0,527472527 0,238095238 *
0,289156627 0,179104478 0 0,32 *
0,418181818 0,489361702 0,222222222 0,545454545 0,377358491 *
0,157303371 0,219178082 0,333333333 0,321428571 0,1875 0,271186441 *
0,269662921 0,301369863 0,416666667 0,392857143 0,3125 0,406779661 0,421052632 *
0,189393939 0,208695652 0,106382979 *
0,126315789 0,082191781 0 0,137931034 *
0,172932331 0,196581197 0,1 0,214285714 0,158730159 *
0,072916667 0,098765432 0,142857143 0,138461538 0,085714286 0,119402985 *
0,118811881 0,130952381 0,172413793 0,164179104 0,135135135 0,169014085 0,173913043 *
0,466666667 0,933333333 0,088888889 0,577777778 0,133333333 0,4 0,155555556 0,222222222 * 0,365217391 0,848484848 0,16 0,634146341 0,206896552 0,423529412 0,28 0,3125 * 0,154411765 0,29787234 0,074074074 0,240740741 0,09375 0,174757282 0,122807018 0,135135135 *
SIMPSON Toarcian *
0,270588235 *
0,074074074 0,169491525 *
DICE Toarcian *
0,156462585 *
0,038461538 0,078125 *
JACCARD Toarcian
decreasing, while the Bairdiacea increase (Bairdiacypris dorisae). In the remainder of the Lower Toarcian, ostracod assemblages are characterised by Cytheracea (Kinkelinella sermoisensis, Ektyphocythere bucki, E. anterocosta, and Praeschuleridea pseudokinkelinella). The Upper Toarcian and Aalenian sediments of the Turmiel and Chelva formations consist of rhythmic alternations of marls and limestones, mudstone and bioclastic wackestone and yield ostracod assemblages comprising many of the Lower Toarcian species plus Cytheropteron alafastigatum, Cytheropterina criba, Kinkelinella fischeri, Ektyphocythere furcata, Praeschuleridea gallemannica, P. ventriosa, Otocythere callosa and Cytherella cadomencis (Goy et al., 1994, 1996). Almost all species cited are common with the other areas of this first (SWG-Sz-PB-GB) cluster (Fig. 5 and Table 5). The Cordillera Iberica assemblages share more species with the Paris Basin (60% of their species) than with those described from Germany, Switzerland and Great Britain. All these areas (CI-PB-SWC-Sz) form a first independent cluster. Spanish assemblages, however, are less similar (share less than 30%) to Canadian and Portuguese assemblages, which form an independent cluster together with the Fasnet Basin assemblages (C-P-FB). Both clusters result when the Dice and Jaccard indices have been used. In contrast with the results described for the Pliensbachian, Toarcian ostracod assemblages recorded from the Quercy area do not show the same
C. Iberica SW Germany Canada Paris Basin Quercy Great Britain Fastnet Basin Portugal Switzerland C. Iberica SW Germany Canada Paris Basin Quercy Great Britain Fastnet Basin Portugal Switzerland C. Iberica SW Germany Canada Paris Basin Quercy Great Britain Fastnet Basin Portugal Switzerland
close similarity to the Cordillera Iberica assemblages and in this way, they are linked to the previous cluster (CI-PB-SWCSz). The reasons may be in the reduced number of species that have been described so far from this area. Results obtained from the cluster analysis show the highest values of similarity between southwestern German and Swiss assemblages. The Lower Toarcian black organic rich clay and interbedded black-shale of the Posidonienschiefer Formation yielded ostracod assemblages dominated by cytheraceans of the genera Kinkelinella, Ektyphocythere, Praeschuleridea, Monoceratina and Trachycythere. The Upper Toarcian ostracod assemblages recorded from the marls, marly limestones and interbedded calcareous shales of the Upper Schwarzjuramergel Formation and for the dark marine shales of the Lower Opalinuston Formation (Knitter and Ohmert, 1983, 1986; Knitter and Riegraf, 1984; Riegraf, 1985; Richter, 1987) are characterised by cytheraceans of the genera Praeschuleridea (P. angulata, P. bernierensis, P. ventriosa) and Kinkelinella (K. sermoisensis, K. costata); Cytherelloidea cadomensis is also well represented (Table 5 and Fig. 11). These assemblages show a high similarity to the Cordillera Iberica, with almost with 40% of common species. The other difference in clustering results between the Dice or Jacard indices and the Simpson index concerns the cluster for southwestern German and Swiss assemblages (SWGSz). These assemblages are linked with Canada when the Sim-
C. Arias, R. Whatley / Geobios 38 (2005) 697–724
709
Table 5 Distribution of selected ostracod species in the Toarcian. Each number represents the sites where these species has been recorded Distribution des espèces d’ostracodes sélectionnées dans le Toarcien. Chaque numéro représente les sites où cette espèce a été enregistrée. SPECIES Bairdia cassiana Bairdia clio Bairdia fortis Bairdia italica Bairdia michelseni Bairdia molesta Bairdia rostrata Bairdiacypris anisica brevis Bairdiacypris dorisae Bairdiacypris trias. postera Cardobairdia liassica Cardobaridia sp. K Cytherella praecadomensis Cytherella toarcensis Cytherelloidea drexlerae Cytheropteron alafastigatum Cytheropteron diversum Ektyphocythere bucki Ektyph. dharennsourensis Ektyphocythere vitiosa Eucytherua liassica Eucytherura spp. Eucytherura tricostata Gen. et sp. D Bate et al. Gramannacyhere ubiquita Hermiella ambo Hermiella comes Hermiella hyblea Isobythocypris cylindrica Isobythocypris tatei Kinkelinella costata Kinkelinella sermoisensis Kinkelinella tenuicostata Liasina lanceolata Monoceratia scrobiculata Monoceratina striata Monoceratina vulsa Monoceratina ungulina Ogmoconcha amalthei Ogmoconcha inflata Ogmoconchella bispinosa Ogmoconchella contractula Paracypris liassica Paracypris redcarensis Polycope cerasia Polycope cincinnata Polycope pelta Pontocyprella cavata Prae. pseudokinkelinella Praeschuleridea ventriosa Pseudohealdia etaulensis Pseudom. subtriangularis Ptychobairdia aselfingensis Ptychob. schaubergensis
LUSITANIA BASIN
GRAND BANKS
SICILY
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STRETTURA
MOROCCO
ALGERIA
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Fig. 11. Number of ostracod species recorded per Lower Jurassic ammonite zone in Northwest Germany, Southwest Germany and Switzerland. (Legend as for Fig. 9). Fig. 11. Nombre d’espèces d’ostracodes enregistrées par zone d’ammonite du Jurassique inférieur au nord ouest et au sud ouest de l’Allemagne et en Suisse (légende comme pour la Fig. 9).
pson index is used because Canada shares its entire low diversity fauna with these areas. Thus, the percentage of common species of both central European areas with Canada is higher than with the remaining areas, although the number of common species is very low. However, when the comparison is made with more diverse assemblages, such as, the Paris Basin, Great Britain or the Cordillera Iberica, German ostracod assemblages share a large number of species, although the percentage of common species (total) is smaller than with the Canadian assemblages. This explains why because the southwestern German and Swiss assemblages (SWG-Sz) form a cluster with the Paris Basin, Great Britain and the Cordillera Iberica assemblages with Jaccard or Dice indices, but with Canada with the Simpson index. French assemblages (Fig. 5) from the Paris Basin are closely related to the Cordillera Iberica and Great Britain (CIGB) cluster, and shares more than 30% species. However there is a lack of data for the beginning of the Toarcian. The Spinatum Zone regressive phase is reflected in non-deposition and erosion during the Tenuicostatum Zone, resulting in a barren level. The rest of the Toarcian is characterised in many areas of the Paris Basin (during the Falciferum Zone) by the deposition of marls and marly limestones, together with black shales (″Schistes carton″). Toarcian ostracod assemblages (Cubaynes and Ruget, 1985; Bodergat and Donze, 1988; Bodergat et al., 1985) are less diverse and become dominated by cytheraceans (Ektyphocythere neumannae, E. quadrata and Pleurifera harpa), small cytherurids (species of Cytheropteron, Infracytheropteron, Rutlandella, Eucytherura), cytherellids and bythoceratids after the disappearance of the healdiids at the base of the Toarcian (Table 5 and Fig. 5). They share more than 50% species with the Cordillera Iberica.
Quercy assemblages exhibit a lower level of similarity with the Cordillera Iberica than in the Pliensbachian. Lower Toarcian deposits in the Quercy area (Cubaynes and Faure, 1989; Cubaynes, 1986) comprise marly and marly-calcareous sediments of the Penne and Lexos formations. At the base, green marls and bioclastic limestones of the Barre à Pecten (Upper Pliensbachian-Lower Toarcian) contain only a few species, such as, Ogmoconcha aequalis, Cytherella praecadomensis and Kinkelinella sermoisensis. The Penne Formation includes black-shales of the Schistes carton (Tenuicostatum Zone) and marls and marly limestones (Serpentinus to Insigne ammonite zones) and yield a rich and diverse ostracod fauna (Cubaynes, 1986; Andreu et al., 1995) dominated by large cytheraceans (Kinkelinella, Ektyphocythere, Praeschuleridea, Camptocythere), meanwhile cytherellids and bairdiids (Cytherella toarcensis, C. praecadomensis and Bairdiacypris dorisae) are very common. These assemblages contain a large number of species in common, almost a 75%, with the Cordillera Iberica (Table 5 and Fig. 5). British assemblages show many similarities with the above assemblages, especially with those from the Paris Basin, and have more than 50% species in common with the Cordillera Iberica (Table 5). In the south of England the succession of thin, hard, micritic limestones, separated by non sequences and marked erosion planes in the Lower Toarcian Junction Bed (Hesselbo and Jenkyns, 1995) yielded only a few species of Ogmoconcha, Ogmoconchella and Hermiella, which disappeared at the base of the Falciferum Zone (Boomer, 1992), followed by an increase in the Cytheracea. A better knowledge of the Lower Toarcian ostracod assemblages was obtained from the study of seven boreholes in the East Midlands (Central England) and a section at Ilminster (South-
C. Arias, R. Whatley / Geobios 38 (2005) 697–724
west England). Their assemblages comprised a rich and diverse ostracod fauna (Bate and Coleman, 1975; Boomer, 1992) characterised by the occurrence of the cytheraceans (Kinkelinella sermoisensis, K. costata, Ektyphocythere anterocosta, E. intrepida, E. debilis, Praeschuleridea pseudokinkelinella), bairdiids and cytherellids, showing a large similarity with German and French assemblages and with more than 50% of species common with the Cordillera Iberica (Table 5 and Figs. 7,8,10). Ostracod assemblages from the Grand Banks (Canada), Zambujal (Portugal) and the Fasnet Basin were grouped into one cluster (Figs. 7,8 and 10). Toarcian sediments of the Brenha Formation and the Whale Unit in Zambujal and the Grand Banks are represented by a sequence of shales, marls and limestones, with no black-shale deposition (Mouterde et al., 1971; Exton, 1979; Lord, 1982; Exton and Gradstein, 1984; Boomer et al., 1998) which have yielded ostracod assemblages dominated by species of Bairdiacypris, large cytheraceans and many species of Paracypris. Both assemblages share more than 50% of species with the Cordillera Iberica (Tables 5 and 6). Inner shelf Toarcian-Aalenian dark calcareous bituminous shales of the Stratton Formation in the Fasnet, Celtic and Cardigan Bay basins, yielded less abundant assemblages, where healdiids are absent and large Cytheracea dominate, with many new species of Bairdia, Cardobairdia, Isobythocypris, Bairdiacypris, Liasina and Procytherura (Ainsworth, 1986b; Ainsworth et al., 1987) (Tables 5,6 and Fig. 6). These assemblages have only minor similarity with those described from the Cordillera Iberica, with less than 20% of their ostracod species in common.
5. The distinction of the Tethyan ostracod province Although our knowledge of Tethyan ostracod assemblages has been improved during recent years they are considerably less well known than western European assemblages. In many cases this is due to the small number of species recorded and/or to their poor preservation. Tethyan assemblages have been described from several localities in Italy and North Africa (Barbieri, 1964; Maupin, 1977; Lord (in Farinacci et al., 1978); Lord, 1982, 1988; Baloge, 1981; Boutakiout et al., 1982; Arias, 1993). Italian assemblages are characterised by the occurrence of members of the superfamilies Bairdiacea and Healdiacea, with a lesser representation of Cytheracea (Bate, 1977; Lord, 1988; Arias, 1993). Only a few species recorded in the Ragusa1 borehole (Barbieri, 1964) have been also described in the Cordillera Iberica. In the Umbria-Marche area, northern Apennines, Pliensbachian pelagic sedimentation represented by micritic limestones with chert nodules of the Corniola Formation and clayey and marly sediments of the Marne del Monte Serrone Formation (Farinacci et al., 1978; Colacicchi et al., 1988) yielded assemblages dominated by healdiids, bairdiaceans (Cardobairdia, Bairdiacypris and Bairdia) and heal-
711
diids (Ogmoconchella, Ogmoconcha and Hermiella) and Liasina lanceolata (Lord, 1978 (in Farinacci et al., 1978); Arias, 1993) (Tables 3 and 6). Italian assemblages are more similar to German assemblages than to those situated in the area of the Grand Banks-Portugal and they share more than 50% of their species with Spanish assemblages. Ostracod assemblages from the shallow carbonate platforms of North Africa (Morocco and Algeria) show close affinities with those recorded in southern European basins (Maupin, 1977; Baloge, 1981; Boutakiout et al., 1982; Lord, 1982; Bassoullet et al., 1991). Boutakiout et al. (1982) and Bassoullet et al. (1991) described assemblages from three sections situated at rides sud-rifaines (Atlas Meseta and eastern Moroccan Meseta). Upper Pliensbachian bioclastic limestones and Toarcian grey marls and marly limestones yielded ostracod assemblages characterised by the important occurrence of cytheraceans, platycopids and healdiids, together with some species of the genera Liasina, Bairdia and Bairdiacypris (Table 6). Ostracod assemblages from the western part of the Oman Meseta, have been studied by Maupin (1977). Upper Pliensbachian assemblages are characterised by the occurrence of healdiids (Hermiella ambo, Ogmoconcha amalthei) and Toarcian by species of Bairdia, Cytherella, Isobythocypris, Paracypris, Liasina and some large cytheraceans. These assemblages are very similar to those recorded by Baloge (1981) from the Rosso-Ammonitico of Western Algeria. Upper Pliensbachian assemblages from this area comprise species of Hermiella, Ogmoconchella, Bairdia and Bairdiacypris and Toarcian assemblages, which are of low diversity, are characterized by the lack of healdiids and the occurrence of Bairdiacypris, Cytherella, Polycope and Kinkelinella (Table 6). However, a different record has been obtained from the study of a borehole (DSDP Leg 79, Site 547) located off North Africa (Bate et al., 1984). Eighteen Jurassic samples (from Hettangian to Pliensbachian) yielded a very peculiar fauna, with many new ostracod taxa and with many left in open nomenclature and where smooth taxa dominate (healdiids and bairdiids). As a whole, these assemblages show only a few species in common with previously recorded assemblages from NW Europe. A complete analysis of the Tethyan Lower Jurassic ostracod assemblages (Table 6) reflects a great similarity in composition with those from Central Europe, although with differences in the representation of their taxa. The main differences are that the presence of large Cytheracea are less common than in European assemblages and that ornamented forms of bairdiids and healdiids (Ptychobairdia, Hermiella) are present even during the Toarcian. The present knowledge about Tethyan ostracod assemblages considered here does support the recognition of a separate Tethyan province. 6. Conclusions This paper considers the degree of similarity among European (and surrounding areas) Lower Jurassic Ostracoda
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Table 6 Distribution of the most representative Tethyan ostracod species considered in this study Distribution des espèces téthysiennes d’ostracodes les plus représentatives considérées dans cette étude. ostracod species Monoceratina amlingstadtensis Triebel and Bartenstein, 1938 Monoceratina mesoliassica Triebel and Bartenstein, 1938 Monoceratina michelseni Riegraf, 1984 Monoceratina seebergensis Triebel and Bartenstein, 1938 Monoceratina stimulea (Schwager, 1866) Monoceratina striata Triebel and Bartenstein, 1938 Monoceratina ungulina Triebel and Bartenstein, 1938 Nanacythere (D.) firma Herrig, 1969b Nanacythere (D.) fissicosta Herrig, 1969b Nanacythere (G.) minor Michelsen, 1975 Nanacythere (N.) simplex Herrig, 1969b Ogmoconcha (Hermiella) ambo Lord and Moorley, 1974 Ogmoconcha (Hermiella) cincta Malz, 1975 Ogmoconcha (H.) circumvallata Dreyer, 1967 Ogmoconcha (Hermiella) cista Malz, 1975 Ogmoconcha (Hermiella) comes Malz, 1975 Ogmoconcha (Hermiella) hyblea (Barbieri, 1964) Ogmoconcha (Hermiella) inflata (Ainsworth, 1987) Ogmoconcha (H.) intercedens Dreyer, 1967 Ogmoconcha (Hermiella) klingleri Malz, 1975 Ogmoconcha amalthei (Quenstedt, 1967) Ogmoconcha amalthei rotunda Dreyer, 1967 Ogmoconcha contractula Triebel, 1941 Ogmoconcha convexa Boomer, 1991 Ogmoconcha dentata (Issler, 1908) Ogmoconcha eocontractula Park 1984 Ogmoconchella aenticulata (Pietrzenuk, 1961) Ogmoconchella aequalis (Herrig, 1969a) Ogmoconchella aspinata (Drexler, 1958) Ogmoconchella danica Michelsen, 1975 Ogmoconchella impressa Malz, 1975 Ogmoconchella michelseni (Michelsen, 1975) Omoconchella mouhersensis (Apostolescu, 1959) Ogmoconchella propinqua Malz, 1971 Ogmoconchella secunda Herrig, 1981a Ogmoconchella sp. A Lord, 1974 Ogmoconchella sp. B (Apostolescu, 1959) Oligocythereis? Mochrarensis Boomer, 1991 Paracypris redcarensis (Blake, 1876) Paracypris sp. 1 Exton, 1979 Paracypris sp. A Riegraf, 1985 Pleurifera harpa harpa (Klinger and Neuweiler, 1959) Pleurifera harpa harpoidea (Gramann, 1962) Pleurifera vermiculata (Apostolescu, 1959) Polycope cerasia Blake, 1876 Polycope cincinnata Apostolescu, 1959 Polycope decorata Apostolescu, 1959 Polycope discus Fischer, 1961a Polycope minor Michelsen, 1975 Polycope pelta Fischer, 1961a Polycope plumhoffı Bate and Coleman, 1975 Polycope tenuireticulata (Herrig, 1981c) Progonoidea reticulata (Klingler and Neuweiler, 1959) Pseudohealdia etaulensis (Apostolescu, 1959) Pseudohealdia gruendeli Malz, 1971 Pseudohealdia pseudospina (Herrig, 1969a)
1
2
3 C
4
C
C
C C
C C C C
C C
5 C C C C C C C C C
C
C
C
C
C C
C C C
C C C
C
C
C
C
C
C C C
7
8 C
9
10
11
12
C
13 C
14
15
16
17
C
C
C
C
C
C
C
C
C
C C C
6
C C C C C C
C C C C
C C C C C
C C C C
C C C C C C C C C C C
C C
C C C
C C C
C C
C C
C C C C
C C
C C
C C
C C
C C
C C C
C
C C
C
C C C C C C C C C C
C C C
C
C
C C C
C C C C C C
C C
C C
C C C C C C C
C
C
C C C C
C C
C
C C
C C
C
C C
C
C
C C C
C
C C
C C
C
C
C C
C C
C
C
C
C
C
C
C C
C C
C
C
C C
C C (continued on next page)
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Table 6 (continued) ostracod species Pseudohealdia pseudohealdidae (Gründel, 1964) Pseudohealdia transversa (Gründel, 1964) Pseudohealdia truncata Malz, 1971 Pseudomacr. subtriangularis Michelsen, 1975 Trachycythere angusta Triebel and Klingler, 1959 Trachycythere horrida Triebel and Klingler, 1959 Trachycythere tub. seratina Triebel and Klingler, 1959 Trachycythere tub. tubulosa Triebel and Klingler, 1959 Trachycythere verrucosa Triebel and Klingler, 1959 Trachycythere multistriata Michelsen, 1975 Triassocythere? sp. 4135 (Michelsen, 1975)
1
2
3
4 C
C
7
8 C C
9
10 C C
11
12
C
C C C
6
C
C C C C
5
C C C C C C
C C C C
13
14
15
16 C
17
C C
C C
C C C
C
C
C
Key to localities: 1. North Sea. 2. Shane, Sweden. 3. Danish Embayment. 4. NE Germany. 5. NW Germany. 6. SW Germany. 7. Switzerland. 8. Paris Basin. 9. Quercy. 10. Great Britain. 11. Fastnet Basin. 12. Cordillera Iberica. 13. Portugal. 14. Canada. 15. Italy. 16. Australia. 17. Argentina.
assemblages, with particular attention to their respective relationships with the assemblages of the Cordillera Iberica (Spain). The results indicate that during the Pliensbachian, the Cordillera Iberica assemblages show a higher similarity with French basins (Quercy and the Paris Basin) than those described in the remaining areas of northwestern Europe. Spanish assemblages are situated in the same major cluster that grouped all western European assemblages (French, Portuguese and British). It is remarkable that the grade of similarity among western European assemblages is slightly lower than that which they show individually with Central and northeastern Europe assemblages (Danish and German basins). Canadian, Italian and Swiss assemblages show a low grade of similarity to remaining European assemblages, probably as a consequence of our relatively poor knowledge of the faunas and their poor representation. In the Pliensbachian, the Paris Basin assemblages form a group with the rest of western European assemblages (PB-QCI-GB-P-FB), meanwhile southwestern German basins are within the cluster (NEG-NWG-SWG-Dk) with the nortwestern and Central European assemblages, indicating a first level disjunction. It is noteworthy that northwestern African assemblages are very similar to those described in Central Europe. Showing the occurrence of a open connection between Tethys and the epicontinental sea to the north during the Early Jurassic. For the Toarcian, the cluster analyses show the Cordillera Iberica assemblages have a composition very similar to those of Eastern Europe (German, French and British basins) and their assemblages are included in the same cluster. Meanwhile, Canadian, Fasnet Basin and Portuguese assemblages are grouped together at a high level of similarity, forming another major cluster. Thus, Toarcian Cordillera Iberica assemblages are now more related to the Paris Basin assemblages than they were during the Pliensbachian. Similarly, the Paris Basin assemblages show higher similarity to south-
western German assemblages in the Toarcian than they showed in the Pliensbachian. The results indicate that during the Pliensbachian there was a better communication among western European areas and less between western and Central Europe than during the Toarcian. This latter communication was increased during the Toarcian, meanwhile the most western European areas (the Fasnet Basin, Portugal, Mochras borehole, Canada) show a clear differentiation in relation to the rest of western areas (Iberian Peninsula, French and British Basins). In the Lower Jurassic, especially in the Toarcian, the European ostracod fauna show no clear differentiation into two major biogeographical provinces as do other groups of fossil invertebrates (ammonites, brachiopods). It must be said, however, that Tethyan ostracod assemblages, especially in the post-Pliensbachian, are relatively poorly known.
Acknowledgements The content and presentation of this paper was enhanced by the thoughtful and highly constructive reviews of Prof. A. Lord and Prof. E. Herrig to whom I owe our greatest debts of gratitude. I owe a special debt to gratitude to Dr. Jean Vannier for the critical comments of the early version of our paper and for his patience. I also want to thank, the members of the editorial board of Geobios for their valuable assistance during all stages of this manuscript. C.A. owe special thanks to Fabienne Ongaro for her excellent handling of the numerous misprints and typographical errors. The ostracoda database is based on CA’s ostracod compilation over the past seven years. We acknowledge the TMR Mobility Program (European Union), the Comunidad de Madrid postdoctoral Program and the DGCYT Projects PB93-0459, PB97-0274 and BTE 2001.1924 from the Ministerio de Educación (MEC) y Ciencia and Ministerio de Ciencia y Tecnología of Spain (MCYT), for financial support.
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Annexe A. List of studied localities and references
Pliensbachian Northeastern Germany (NEG)
Northwestern Germany (NWG)
Southwestern Germany (SWG)
Canada (C) Celtic Sea-Fastnet Basin-Porcupine (FB)
Danish Embayment, Denmark (Dk) and Skane, Sweden(S) Paris Basin, France (PB)
Quercy, France (Q)
Great Britain (GB)
Italy (It) Portugal (P)
Northwestern Switzerland (Sz)
Pietrzenuk, 1961 Dreyer, 1967 Herrig, 1969a,b Herrig, 1979 Herrig, 1980 Herrig, 1981a,b,c Herrig, 1982a,b Triebel and Bartenstein, 1938 Triebel and Klingler, 1959 Gramann, 1963 Malz, 1971 Ohm (Fischer et al., 1986) Fischer, 1961b,c Klingler, 1962 Lord and Morley, 1974 Malz, 1975 Urlichs, 1977 Riegraf, 1984, 1985 Harloff, 1993 Harloff and Jagger, 1994 Exton and Gradstein, 1984 Ainsworth and Horton, 1986 Ainsworth, 1987 Ainsworth et al., 1987 Ainsworth, 1990 Michelsen, 1975 Sivhed, 1980 Apostolescu, 1959, 1961 Bizon, 1960, 1961 Apostolescu et al., 1961 Bizon and Oertli, 1961 Cousin and Apostolescu, 1961 Champeau, 1961 Oertli and Grosdidier, 1961 Magné and Malmoustier, 1961 Magné et al., 1961 Maupin, 1978 Donze, 1985 Bodergat et al., 1985 Riegraf, 1985 Bodergat and Donze, 1988 Cubaynes and Ruget, 1985 Andreu et al., 1998 Bodergat et al., 1998 Lord, 1972, 1974 Lord and Bown, 1987 Lord, 1978, 1988 Boomer, 1991 Boomer, 1992 Lord, 1978, 1988 Arias, 1993 Exton, 1979 Exton and Gradstein, 1984 Boomer et al., 1998 Richter, 1987
Dobbertin and Eisenach Brandenburg Greifswald Thüringen Thüringen Thüringen Thüringen Baden (Niedersachten) Hannover Weseker Sattel Niedersachten Empelde Baden-Württemberg Baden-Württemberg Baden-Württemberg Baden-Württemberg Baden-Württemberg Baden-Württemberg Baden-Württemberg Kalk-Alpen Bayern and North Tyrol Grand Banks, Newfoundland Fasnet Basin Fasnet Basin Fasnet Basin Porcupine, Slyne, Evris and Donegal Danish Embayment Southern Sweden Paris Basin Basse Normandie Paris Basin Lorraine Ardennes Southeastern Paris Basin Paris Basin Thouars Thouars Vendée France Paris Basin Truc de Balduc Paris Basin Quercy Quercy Quercy England Dorset and Isle of Wight Great Britain Mochras, Wales Ilminster, Southwestern England Strettura Umbria Zambujal Zambujal Zambujal Basel
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Toarcian Southwestern Germany (SWG)
Canada (C) Celtic Sea-Fastnet Basin-Porcupine (FB)
Paris Basin, France (PB)
Quercy, France (Q)
Great Britain (GB)
Portugal (P)
Northwestern Switzerland (Sz)
Fischer, 1961a,b Knitter, 1983, 1984 Knitter and Riegraf, 1984 Riegraf, 1984, 1985 Harloff, 1993 Exton and Gradstein, 1984 Ainsworth and Horton, 1986 Ainsworth et al., 1987 Ainsworth, 1990 Apostolescu, 1959 Apostolescu, 1961 Champeau, 1961 Magné et al., 1961 Oertli and Grosdidier, 1961 Depêche, 1985 Bodergat et al., 1985 Riegraf, 1985 Bodergat and Donze, 1988 Bodergat et al., 1991 Cubaynes and Ruget, 1985 Cubaynes, 1986 Bodergat et al., 1991 Lord, 1974 Bate and Coleman, 1975 Lord, 1978 Boomer, 1991 Boomer, 1992 Exton, 1979 Exton and Gradstein, 1984 Boomer et al., 1998 Richter, 1987
Annexe B. Alphabetical list of species considered in this study A.1. List of the taxa Acrocythere gassumensis Michelsen, 1975 Acrocythere mimica Bate and Coleman, 1975 Acrocythere oeresundensis Michelsen, 1975 Acrocythere rectangula Michelsen, 1975 Acrocythere reticulata Knitter, 1983 Acrocythere troesteri Riegraf, 1984 Ambigocythere concentricostata Herrig, 1985 Aphelocythere kuhni Triebel and Klingler 1959 Bairdia cariata Drexler, 1958 Bairdia clio Bizon, 1960 Bairdia crassa Drexler, 1958 Bairdia donzei Herrig, 1979 Bairdia fortis Drexler, 1958 Bairdia guttulae Herrig, 1979 Bairdia hahni Lord and Morley, 1974 Bairdia kempfi Ainsworth 1989 Bairdia michelseni Herrig, 1979 Bairdia molesta Apostolescu, 1959 Bairdia ohmerti Knitter, 1983 Bairdia praehilda Herrig, 1979 Bairdia rostrata Issler, 1908 Bairdia sp. A Ainsworth, 1986b
715
Baden-Württemberg Baden-Württemberg Baden-Württemberg Baden-Württemberg Baden-Württemberg Grand Banks, Newfoundland Fasnet Basin Fasnet Basin Porcupine, Slyne, Evris and Donegal Paris Basin South of Paris Basin Southeastern Paris Basin Thouars Paris Basin France Paris Basin Truc de Balduc Paris Basin Paris Basin Quercy Quercy Quercy England Empingham and Upwood Great Britain Mochras, Wales Ilminster Zambujal Zambujal Zambujal Basel
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Bairdia sp. A Ainsworth, 1990 Bairdia sp. A Ohm, 1986 Bairdia thuringica Herrig, 1979 Bairdia undulata Herrig, 1979 Bairdia verrucosa Herrig, 1979 Bairdia? eirensis Ainsworth, 1986a Bairdiacypris dorisae (Knitter, 1984) Bairdiacypris rectangularis Ainsworth, 1986b Bairdiacypris sartriensis Donze, 1966 Bairdiacypris triangularis Ainsworth, 1986b Bairdiacypris triasica postera Herrig, 1979 Bairdiacypris tumida Ainsworth, 1987 Bythocypris fabaeformis Drexler, 1958 Cardobairdia fastnetensis Ainsworth, 1987 Cardobairdia liassica (Drexler, 1958) Cardobairdia Nr. 103 Klingler, 1962 Cardobairdia posteroprolata Ainsworth, 1987 Cardobairdia sp. A Ainsworth, 1987 Cardobairdia sp. K (Apostolescu, 1959) Cardobairdia toarcensis Ainsworth 1986b Cristacythere betzi (Klingler and Neuweiler, 1959) Cristacythere costata Michelsen, 1975 Cristacythere crassireticulata Michelsen, 1975 Cytherella demiexensis Ainsworth, 1989 Cytherella drexlerae Field, 1966 Cytherella lindseyensis Lord, 1974 Cytherella praecadomensis (Knitter and Riegraf, 1984) Cytherella toarcensis Bizon, 1960 Cytherelloidea anningi Lord, 1974 Cytherelloidea cadomensis (Bizon, 1960) Cytherelloidea circumscripta (Blake, 1876) Cytherelloidea drexlerae (Field, 1966) Cytherelloidea lacertosa Apostolescu, 1959 Cytherelloidea modesta Apostolescu, 1959 Cytheropteron alafastigatum Fischer, 1962 Cytheropteron byfieldensis Boomer and Bodergat, 1992 Cytheropteron cavatum Michelsen, 1975 Cytheropteron diversum Herrig, 1969b Cytheropteron foveolatum Michelsen, 1975 Cytheropteron sp. B Riegraf, 1985 Cytheropteron? sp. A Riegraf 1985 Cytheropteron? sp. Boomer, 1991 Ektyphocythere acuminata (Riegraf, 1984) Ektyphocythere ambo Boomer, 1988 Ektyphocythere anterocosta Boomer, 1988 Ektyphocythere bucki (Bizon, 1960) Ektyphocythere champeauae (Bizon, 1960) Ektyphocythere debilis Bate and Coleman, 1975 Ektyphocythere dharennsourensis Boutakiout et al., 1982 Ektyphocythere intrepida Bate and Coleman, 1975 Ektyphocythere knitteri Riegraf, 1984 Ektyphocythere lanceolata Boomer, 1988 Ektyphocythere laqueata (Klingler and Neuweiler, 1959) Ektyphocythere luxuriosa (Apostolescu, 1959) Ektyphocythere multicostata (Klingler, and Neuweiler, 1959) Ektyphocythere neumannae (Maupin, 1978)
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Ektyphocythere quadrata Boomer and Lord, 1988 Ektyphocythere triebeli (Klingler and Neuweiler, 1959) Ektyphocythere vitilis furcata (Wienholz, 1958) Ektyphocythere vitilis vitilis (Apostolescu et al., 1961) Ektyphocythere vitiosa (Apostolescu, 1959) Ektyphocythere vulgaris (Klingler and Neuweiler, 1959) Eocytheropterina jutlandica (Michelsen, 1975) Eucytherura sp. Riegraf, 1985 Fabalicypris symmetrica Herrig, 1979 Gammacythere ubiquita Malz and Lord, 1976 Gen ind. sp. A Boutakiout et al., 1982 Gramannella apostolescui (Gramann, 1962) Gramannella laevigata Michelsen, 1975 Gramannella tatei (Gramann, 1962) Gramannicythere aubachensis Riegraf, 1984 Gramannicythere bachi bachi Herrig, 1982a Gramannicythere bachi ornata Herrig, 1982a Gramannicythere coniuncta Herrig, 1982a Gramannicythere sp. Malz and Nagy, 1989 Hemiparacytheridea (Tuberocytheridea) angulocostata Knitter, 1983 Infracytheropteron gwashense Bate and Coleman, 1975 Infracytheropteron pulchellum Bate and Coleman, 1975 Infracytheropteron rarum Knitter, 1983 Infracytheropteron supraliasicum (Herrig, 1981a) Isobythocypis tatei (Coryell, 1963) Isobythocypris aff. elongata (Blake, 1876) Sivhed, 1980 Isobythocypris cf. elongata (Blake, 1876) Sivhed, 1980 Isobythocypris cf. elongata (Blake, 1876) Drexler, 1958 Isobythocypris cf. elongata (Blake, 1876) Michelsen, 1975 Isobythocypris cylindrica (Herrig, 1979) Isobythocypris dorsoconversa Ainsworth, 1986b Isobythocypris elongata (Blake, 1876) Isobythocypris ovalis Bate and Coleman, 1975 Isobythocypris pliensbachiensis Ainsworth, 1986b Isobythocypris unispinata Apostolescu, 1959 Isobythocypris? sp. Dreyer, 1967 Kinkelinella costata Knitter, 1984 Kinkelinella kadeshensis Rosenfeld et al., 1987 Kinkelinella persica Bate and Coleman, 1975 Kinkelinella procera Herrig, 1985 Kinkelinella sermoisensis (Apostolescu, 1959) Kinkelinella tenuicostata Martin, 1960 Kinkelinella? idrissensis Boutakiout et al., 1982 Klinglerella auleata (Klingler and Neuweiler, 1959) Klinglerella bipartita (Klingler and Neuweiler, 1959) Klinglerella elongata (Michelsen, 1975) Klinglerella foveolata Michelsen, 1975 Klinglerella glabellata (Klingler and Neuweiler, 1959) Klinglerella herrigi Ainsworth, 1989 Klinglerella intermedia (Klingler and Neuweiler, 1959) Klinglerella katsloesensis Sivhed, 1980 Klinglerella lacunosa Ainsworth, 1989 Klinglerella medioreticulata (Michelsen, 1973) Klinglerella moorei (Brady, 1872) Klinglerella sinemuriana Ainsworth, 1989 Klinglerella sulcata (Klingler and Neuweiler, 1959)
717
718
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Klinglerella translucens Ainsworth, 1989 Klinglerella variabilis (Klingler and Neuweiler, 1959) Ledahia conviniens Herrig, 1980 Ledahia septenaria (Gründel, 1964) Ledahia telata (Drexler, 1958) Liasina cf. vestibulifera Sivhed 1980 Liasina cylindrica Ainsworth, 1986b Liasina lanceolata (Apostolescu 1959) Liasina vestibulifera Gramann 1963 Monoceratina amlingstadtensis Triebel and Bartenstein, 1938 Monoceratina frentzeni Triebel and Bartenstein, 1938 Monoceratina mesoliassica Triebel and Bartenstein, 1938 Monoceratina michelseni Riegraf, 1984 Monoceratina seebergensis Triebel and Bartenstein, 1938 Monoceratina stimulea (Schwager, 1866) Monoceratina striata Triebel and Bartenstein, 1938 Monoceratina ungulina Triebel and Bartenstein, 1938 Nanacythere (D.) firma Herrig, 1969b Nanacythere (D.) fissicosta Herrig, 1969b Nanacythere (G.) circumcostata Michelsen, 1975 Nanacythere (G.) elegans (Drexler, 1958) Nanacythere (G.) elongata Michelsen, 1975 Nanacythere (G.) minor Michelsen, 1975 Nanacythere (G.) paracostata Michelsen, 1975 Nanacythere (N.) simplex Herrig, 1969b Nanacythere persicaeformis Riegraf, 1984 Ogmoconcha (Hermiella) ambo Lord and Moorley 1974 Ogmoconcha (Hermiella) cincta Malz, 1975 Ogmoconcha (Hermiella) circumvallata Dreyer, 1967 Ogmoconcha (Hermiella) cista Malz, 1975 Ogmoconcha (Hermiella) comes Malz, 1975 Ogmoconcha (Hermiella) hyblea (Barbieri 1964) Ogmoconcha (Hermiella) intercedens Dreyer, 1967 Ogmoconcha (Hermiella) klingleri Malz, 1975 Ogmoconcha amalthei amalthei (Quenstedt, 1967) Ogmoconcha amalthei rotunda Dreyer, 1967 Ogmoconcha contractula Triebel, 1941 Ogmoconcha convexa Boomer, 1991 Ogmoconcha dentata (Issler, 1908) Ogmoconcha eocontractula Park, 1984 Ogmoconcha hagenowi Drexler, 1958 Ogmoconchella adenticulata (Pietrzenuk, 1961) Ogmoconchella aequalis (Herrig, 1969a) Ogmoconchella aspinata (Drexler, 1958) Ogmoconchella celticensis Ainsworth, 1989 Ogmoconchella danica Michelsen, 1975 Ogmoconchella gruendeli Malz, 1971 Ogmoconchella impressa Malz, 1971 Ogmoconchella michelseni Michelsen, 1975 Ogmoconchella mouhersensis (Apostolescu, 1959) Ogmoconchella propinqua Malz, 1971 Ogmoconchella scanica Sivhed, 1977 Ogmoconchella serratostriata Ainsworth, 1989 Ogmoconchella sp. A Lord, 1974 Ogmoconchella sp. B (Apostolescu, 1959) Ogmoconchella sp. B Cubaynes, 1986
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Ogmoconchella transversa (Gründel, 1970) Paracypris liassica (Bate and Coleman, 1975) Paracypris redcarensis (Blake, 1876) Paracypris sp. 1 Exton, 1979 Paracypris sp. 2 Exton, 1979 Paracypris sp. A Riegraf, 1985 Paracypris sp. C Ainsworth, 1986b Paracypris sp. Cubaynes, 1986 Paracypris? longiformis Sivhed, 1980 Paradoxostoma? fusiforme (Drexler, 1958) Paradoxostoma? pulchellum Michelsen, 1975 Paranotacythere (Unicosta) decorata (Apostolescu et al., 1961) Paranotacythere (Unicosta) mochrarensis (Boomer, 1991) Paranotacythere (Unicosta) terquemiana (Jones, 1872) Pleurifera harpa harpa (Klingler and Neuweiler, 1959) Pleurifera harpa harpoidea (Gramann, 1962) Pleurifera perplexa (Klingler and Neuweiler, 1959) Pleurifera plicata (Apostolescu, 1959) Pleurifera vermiculata (Apostolescu, 1959) Polycope cerasia Blake 1876 Polycope cincinnata Apostolescu 1959 Polycope decorata Apostolescu 1959 Polycope discus Fischer 1961a Polycope minor Michelsen 1975 Polycope pelta Fischer 1961a Polycope plumhoffı Bate and Coleman 1975 Polycope tenuireticulata (Herrig, 1981c) Pontocyprella fabaeformis (Drexler, 1958) Praeschuleridea arguta Ainsworth 1986a Praeschuleridea aspera Knitter 1983 Praeschuleridea bernierensis (Apostolescu, 1959) Praeschuleridea costata Ainsworth 1986a Praeschuleridea ellipsoidea Ainsworth 1986b Praeschuleridea gallemannica Malz 1966 Praeschuleridea levita Ainsworth 1986b Praeschuleridea pseudokinkelinella Bate and Coleman 1975 Praeschuleridea punctulata Plumhoff 1963 Praeschuleridea reticulata Ainsworth 1986a Praeschuleridea sp. A Ainsworth 1986b Praeschuleridea ventriosa Plumhoff 1963 Praeschuleridea whatleyi Ainsworth 1986b Praescleridea magnycourtensis (Apostolescu, 1959) Procytheridea laneuvevillae Donze 1967 Procytheridea lotharingiae Apostolescu 1959 Procytheridea multiforata (Klingler and Neuweiler, 1959) Procytheridea perplexa (Klingler and Neuweiler, 1959) Procytheridea spinaecostata (Klingler and Neuweiler, 1959) Procytheridea undulata Apostolescu 1959 Procytheridea? jardensis Maupin 1978 Procytherura celtica Ainsworth 1986b Procytherura euglyphea Ainsworth, 1986b Procytherura hastata Bate and Coleman, 1975 Procytherura mediocostata Bate and Coleman, 1975 Procytherura multicostata Ainsworth, 1986b Procytherura suebica (Herrig and Richter, 1990) Procytherura werneri (Riegraf, 1984)
719
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