Palaeozoogeography of Western European Lower Jurassic (Pliensbachian and Toarcian) Ostracoda

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, Spainb 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 geo-graphical areas. A list of 270 marine ostracod species from Europe, North Africa and North America has been compiled and subjected toquantitative analysis (using multivariate methods) in order to ascertain the degree of similarity between Spanish assemblages and thosedescribed from these areas. The cluster analysis demonstrates that degree of similarity generally reflects proximity in palaeogeographicalposition. The results indicate extensive intercommunication of taxa between the basins, and also the absence of clear differentiation betweenTethyan 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 fauniquesentre 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 étécompilée et soumise à l’analyse quantitative (méthodes d’analyse multivariée) de façon à déterminer le degré de similitude entre les assem-blages d’Espagne et ceux décrits à partir de ces régions. Les classifications hiérarchiques démontrent que le degré de similitude reflètegénéralement la proximité paléogéographique. Les résultats indiquent une communication très importante des taxons entre les basins et aussiune 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 elgrado de semejanza faunística entre 13 áreas geográficas. Se ha compilado una lista de 270 especies de ostrácodos marinos descritos enEuropa, Africa y Norte América. El resultado ha sido sometido a un ánalisis cuantitativo (usando métodos de análisis multivariante) con el fínde conocer el grado de semejanza entre las asociaciones de ostrácodos españolas y aquellas descritas en dichas áreas. El análisis de tipo clusterdemuestra que el grado de semejanza es mayor entre áreas paleogeograficamente próximas. Los resultados muestran una intercomunicaciónmuy amplia de taxones entre las cuencas estudiadas. Además, los resultados de este estudio indican la inesistencia de una clara diferenciaciónentre fanuas típicas del Tethys y faunas boreales.© 2005 Elsevier SAS. All rights reserved.

* Corresponding author.E-mail address: [email protected] (C. Arias).

Geobios 38 (2005) 697–724

http://france.elsevier.com/direct/GEOBIO/

0016-6995/$ - see front matter © 2005 Elsevier SAS. All rights reserved.doi:10.1016/j.geobios.2002.10.001

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 ani-mal taxa in space and time, which in most cases is largely aconsequence of their ecological requirements. Thus, zooge-ography transcends the classical study of the geographicaldistribution of organisms, involving as it does a range of otherfactors, including ecological processes, which limit the dis-tribution of species. In an ecological context, zoogeogra-phers have used geographical patterns of distribution to inferthe role of such physico-chemical factors as temperature,water depth, O2 concentration, pH, light, etc. Biotic factors,although more difficult to quantify are, notwithstanding, ofequal importance. However, in the study of fossils, it is alsonecessary to embrace an evolutionary dynamic with respectto the phylogenetic history of the taxa concerned, togetherwith evolving palaeogeographies and their consequences interms of palaeoenvironmental successions; the whole, ofcourse, being complicated by hiatuses and taphonomic pro-cesses. The intermittent nature of preservation of strata andoutcrop, the wide distribution of Lower Jurassic sedimentsover the European continent, together with the occurrence ofsome lithologies highly prejudicial to the preservation or therecovery of Ostracoda, will naturally bias the results of thisstudy to an unknown degree.

The number of papers concerning the general distributionof Jurassic marine Ostracoda (Bate, 1977; Lord, 1978, 1982,1988; Herrig, 1988) is less than for other groups of fossilsinvertebrates, such as ammonites, brachiopods, bivalves orforaminifers, etc. The present paper has been compiled froma comprehensive literature of ostracod studies for each of the13 selected geographical areas across Europe. It considersthe main composition of their Pliensbachian and Toarcianostracod assemblages and also their faunal interrelationshipsin that both similarities and differences are analysed.

2. Provinciality and faunal provinces during the LowerJurassic

In spite of the abundant literature on Jurassic Ostracoda,rather little attention has been paid to their spatial occur-rence. This has been not the case with other invertebrategroups. The distribution patterns of such groups as ammo-nites, brachiopods and bivalves have been used to proposethe existence of two major provinces the Northern Hemi-sphere Lower Jurassic. These are the Boreal and Tethyan prov-inces (Neumayr, 1882, 1883; Uhlig, 1911; Arkell, 1956; Hal-lam, 1975, 1983; Enay, 1980; Taylor et al., 1984; Cariou etal, 1985; Damborenea, 1993, etc.).

However, in those studies based on microfossils such prov-inces have not been so readily recognised. This is the case,for example, with foraminiferal assemblages. Gordon (1970)distinguished five types of foraminiferal assemblages (Gor-don, 1970; Exton and Gradstein, 1984). Three are character-istic of shelf seas (composed mainly of Nodosaridae), whilethe other two are characteristic of the Tethys and its margins(which included planktonic and arenaceous foraminifera). Inaddition to differences in their geographical distribution (inrelation to Boreal and Tethyan ammonite provinces these typesof foraminiferal assemblages alternate with each other thesame localities and are obviously local ecofacies.

The study of the distribution of Northern HemisphereMesozoic Ostracoda has allowed the identification of fiveprovinces: European, Tethyan, North African, American andEast African (Bate, 1977). Bate indicates that only the firsttwo provinces can be clearly recognised in the Lower Juras-sic. The European Province extends from Newfoundland inthe west, to the Ukraine in the east and from Greenland in thenorth to the northern margin of the Tethys in the south.Although, its total inclusion is open to doubt, the Iberian Pen-insula is also usually included in this province (Bate, 1977).The Tethyan Province (Bate, 1977; Lord, 1988) extends fromthe eastern coast of Canada to Northwest Africa (Tunisia,Morocco, Algeria, etc.).

For this study the palaeogeographical framework of Zie-gler (1988, 1991, 1992) and Bassoullet et al. (1992) has beenadopted (Fig. 1). The Lower Jurassic was marked by the devel-opment of an extensive epicontinental sea that covered theentire present western European continent. Ziegler (1988) rec-ognized three North European provinces based on facies: Bal-tic and Polish, North Sea-North Germany and southwesternEurope. The first one is a continental and estuarine, clasticfacies province, situated between the Fenno-Scandian Highand the Bohemian Massif. The second was an open, colderwater marine shale dominated area, which extended from theNorwegian-Greenland Sea into the Netherlands and from theScottish Highlands into northern Germany. The Southwest-ern Europe province was a warmer water, carbonate-shaledepositional area that extended from the southwest of theBohemian-Rhenish-London-Brabant massifs to the Irish Mas-sif axis.

In the Tethys area, Bassoullet et al. (1992) distinguishedcarbonate platforms of North Africa (Moroccan High Atlas,Algerian Sahara and Atlas), basinal environment argilla-ceous sedimentation (Subbetic and Tellian basins) and thepelagic or hemipelagic deposits of the Ammonitico Rossolimestones in the western Tethys.

698 C. Arias, R. Whatley / Geobios 38 (2005) 697–724

3. Material and methods

The aim of this study is to assess the similarity betweenthe Spanish and other European ostracod assemblages bymeans of analytical methods based on measures of similarityor dissimilarity. A taxonomic evaluation of published litera-ture was compiled and a faunal list of 270 marine ostracodspecies was elaborated by combining this data with that addedby unpublished information obtained from various authors(Appendix A and B). Since the number of individual locali-ties that have been considered in this paper is large, thesesites have been grouped at the level of locally defined geo-graphical areas.

These areas are: C: Grand Banks, Canada; CI: the Cordil-lera Iberica, Spain; DK: Danish Embayment, Denmark andSweden; FB: the Fasnet Basin and North Celtic basins(Mochras borehole included); GB: British basins (Mochrasborehole was not included); It: Italy; M: Morocco; NEG:Northeast Germany; NWG: Northwest Germany; PB: Paris

Basin, France; Q: Quercy, France; SWG: Southwest Ger-many; Sz: Switzerland; P: the Lusitanian Basin, Portugal(Fig. 1).

Due to clearly noticeable differences described betweenPliensbachian and Toarcian ostracod assemblages, two datamatrices were constructed, one for Pliensbachian and the otherfor Toarcian assemblages. Having compiled the raw data, theinitial data matrix was edited with respect to the occurrence(presence/absence) of taxa with respect to a set of studiedlocalities of the same age. Each similarity matrix was sym-metrical and was made in the form of Q-mode (Reyment,1971, 1979, 1987; Keesey and Whittaker, 1976; Digby andKempton, 1987).

A problem in relation to the use of statistical methods isthe wide choice of clustering methods and indices (Baroni-Urbani and Buser, 1976). Many authors have suggested dif-ferents 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

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).

699C. Arias, R. Whatley / Geobios 38 (2005) 697–724

and Mapples, 1987, 1989; Mapples and Archer, 1988; Shi,1993). For the present study, the selection of indices was madeby considering those that would satisfy the following condi-tions: (a) range from 0 (totally different) to 1 (identical)(Goodall, 1973; Lamont and Grant, 1979; Shi, 1993); (b) inde-pendent of double absence (absence in both areas) (Goodall,1973; Henderson and Heron, 1976; Janson and Vegelius,1981; Hubalek, 1982; Shi, 1993); (c) indices that emphasisepresence should be more reliable, because the presence of ataxon in a locality is much less ambiguous than absence (poorsampling, loss through taphonomical process, difficulty withtaxonomic identification of taxa, etc.); (d) not susceptible todifferences in sample size (Wolda, 1981; Koch, 1987). Onthe basis of these criteria, three binary similarity coefficientshave 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) cho-sen 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 MicrosoftExcel 3.0 for the Macintosh and Ntsys-pc. (Version 1.5) havebeen used.

4. Results from the cluster analysis

4.1. Pliensbachian

Table 2 and Figs. 2–4 summarise the results of the analy-sis carried out on Pliensbachian taxa. The dendrograms basedon the Dice and Jaccard similarity indices have grouped thegeographical regions in two major clusters: (1) the CordilleraIberica-Portugal (CI-P) and (2) Northeast-Southwest Ger-many (NEG-SWG). The first cluster contains all westernareas, with two subclusters: (1a) the Paris Basin-Quercy-theCordillera Iberica (PB-Q-CI) and (1b) Portugal-the FasnetBasin (P-FB). The second cluster contains two subclustersthat grouped all German and Danish assemblages: (2a) North-west Germany-Southwest Germany (NWG-SWG) and (2b)Northeast Germany-Denmark (NEG-Dk).

Less similar, Swiss, Italian and Canadian assemblages havebeen joined to the remaining areas as independent geographi-cal areas. Swiss and Italian assemblages show the highest val-ues of similarity with southwestern German and the ParisBasin assemblages. Canadian assemblages show minimumsimilarity values with all other studied areas, what may be aconsequence of their very low diversity. Surprisingly, Britishassemblages do not cluster with the Fasnet Basin, but form

Table 1Differences of selected characteristics in the Similarity Indices used in the studyDifférences des caractères sélectionnés pour les indices de similarités utilisés dans cette étude.

Indices(Authors)

Formula Range a = 0 b = c = 0 Independentof d

Affectedby samplesize

Emphasis Ignore Recommended References

Simpson a-------a + b

(a + b) <(a + c)

0.1 0 1 yes little Useful tomeasuredissimilaritybetweensamples ofdifferent sizes

Absencefrombothsamples

Hazel 1969;Fallaw 1979;Shi 1993

Cheetham andHazel, 1969;Hazel, 1970;Henderson andHeron, 1976;Fallaw, 1979;Archer andMapples, 1978;Shi, 1993b

Dice 2a-------2a + b + c

0.1 0 1 yes moderate

Lowemphasisin mutualpresence

Absencefrom bothsamples

Hazel, 1969;Pierce, 1984;Shi, 1993

Cheetham andHazel, 1969;Hazel, 1970;Archer andMapples, 1987

Jaccard a-------a + b + c

0.1 0 1 yes much

Presencein bothsamples

Absencefrom bothsamples

Shi, 1993 Cheetham andHazel, 1969;Hazel, 1970;Henderson andHeron, 1976;Cambell andValentine, 1977;Flessa andMiyazaki, 1978;Archer andMapples, 1987;Shi, 1993

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 absentfrom both samples.


Table 2Values of the similarity indices between the Pliensbachian ostracod assemblages, which have been described in the different sample localities formerValeurs des indices de similarités entre les assemblages d’ostracodes du Pliensbachien qui ont été décrites dans les différentes localités échantillonnées auparavant.

C.Iberica

NE.Germany

NW Germany SWGermany

Canada Denmark Paris Basin Quercy GreatBritain

Italy FastnetBasin

Portugal Switzerland

* 0,421052632 0,526315789 0,421052632 0 0,473684211 0,736842105 0,947368421 0,421052632 0,105263158 0,315789474 0,473684211 0,210526316 C. Iberica

* 0,411764706 0,450980392 0,019607843 0,470588235 0,196078431 0,156862745 0,196078431 0,098039216 0,196078431 0,215686275 0,137254902 NE. Germany

* 0,638297872 0,021276596 0,425531915 0,29787234 0,212765957 0,276595745 0,127659574 0,276595745 0,319148936 0,191489362 NW Germany

* 0,032258065 0,338709677 0,274193548 0,129032258 0,209677419 0,14516129 0,209677419 0,338709677 0,258064516 SW Germany

* 0,5 1 0 0 0 0,5 1 0,5 Canada

SIMPSON * 0,266666667 0,177777778 0,133333333 0,155555556 0,266666667 0,377777778 0,155555556 Denmark

* 0,285714286 0,178571429 0,25 0,214285714 0,607142857 0,178571429 Paris Basin

Pliensbachian * 0,333333333 0,148148148 0,407407407 0,259259259 0,074074074 Quercy

* 0,133333333 0,233333333 0,366666667 0,1 Great Britain

* 0,1875 0,1875 0,1875 Italy

* 0,842105263 0,105263158 Fastnet Basin

* 0,096774194 Portugal

* Switzerland

* 0,228571429 0,303030303 0,197530864 0 0,28125 0,595744681 0,782608696 0,326530612 0,114285714 0,285714286 0,36 0,228571429 C. Iberica

* 0,428571429 0,407079646 0,037735849 0,5 0,253164557 0,205128205 0,12345679 0,149253731 0,27027027 0,268292683 0,208955224 NE. Germany

* 0,550458716 0,040816327 0,434782609 0,373333333 0,27027027 0,155844156 0,19047619 0,371428571 0,384615385 0,285714286 NW Germany

* 0,0625 0,392523364 0,377777778 0,179775281 0,195652174 0,230769231 0,305882353 0,451612903 0,41025641 SW Germany

* 0,042553191 0,133333333 0 0 0 0,08 0,121212121 0,111111111 Canada

DICE * 0,328767123 0,222222222 0,186666667 0,229508197 0,352941176 0,447368421 0,229508197 Denmark

* 0,290909091 0,24137931 0,318181818 0,235294118 0,576271186 0,227272727 Paris Basin


* 0,173913043 0,264150943 0,360655738 0,130434783 Great Britain

* 0,153846154 0,127659574 0,1875 Italy

* 0,592592593 0,102564103 Fastnet Basin

* 0,127659574 Portugal

* Switzerland

* 0,129032258 0,178571429 0,109589041 0 0,163636364 0,424242424 0,642857143 0,195121951 0,060606061 0,166666667 0,219512195 0,129032258 C. Iberica

* 0,272727273 0,255555556 0,019230769 0,333333333 0,144927536 0,114285714 0,14084507 0,080645161 0,15625 0,154929577 0,116666667 NE. Germany

* 0,379746835 0,020833333 0,277777778 0,229508197 0,15625 0,203125 0,105263158 0,228070175 0,238095238 0,166666667 NW Germany

* 0,032258065 0,244186047 0,232876712 0,098765432 0,164556962 0,130434783 0,180555556 0,291666667 0,258064516 SW Germany

* 0,02173913 0,071428571 0 0 0 0,041666667 0,064516129 0,058823529 Canada

JACCARD * 0,196721311 0,125 0,086956522 0,12962963 0,214285714 0,288135593 0,12962963 Denmark

* 0,170212766 0,094339623 0,189189189 0,133333333 0,404761905 0,128205128 Paris Basin


* 0,095238095 0,152173913 0,22 0,069767442 Great Britain

* 0,063829787 0,068181818 0,103448276 Italy

* 0,421052632 0,054054054 Fastnet Basin

* 0,068181818 Portugal

* Switzerland 701C

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an independent cluster related to other European assem-blages. They show especially high similarity values with theCordillera Iberica.

The Simpson index results (Table 2 and Fig. 2) coincidewith those obtained from Dice and Jaccard indices, but Brit-ish assemblages are related to the Cordillera Iberica-Quercy(CI-Q-GB) cluster and Canadian to the Paris Basin-the Fas-net Basin-Portugal cluster (PB-P-FB). The results shows twoextra cluster: (1) Cordillera Iberica-Quercy-Great Britain and(2) Canada-the Paris Basin-the Fasnet Basin-Portugal.

4.1.1. DiscussionThe results show that Spanish assemblages have the high-

est similarity values with French assemblages; they are moresimilar to Quercy than to the Paris Basin assemblages. Span-ish Pliensbachian assemblages from alternating series of lime-stones, marly limestones and bioclastic limestones (Calizasy Dolomias Tableadas de Cuevas Labradas, Calizas y Mar-gas de Almonacid de la Cuba, Margas grises del Cerro delPez and Calizas Bioclásticas de Barahona formations) aredominated by healdiids (Ogmoconchella adenticulata, O.aequalis, O. contractula, O. danica, O. gruendeli, O. pro-pinqua), cytheraceans (Kinkelinella tenuicostata Ektypho-cythere aff. E. vitosa, Gramannella apostolescui) and twovery abundant species, Liasina lanceolata and Cytherella

toarcensis (Arias, 1989, 1991, 1995; Arias and ComasRengifo, 1992; Arias et al., 1992; Arias and Lord, 1999a,1999b). Species of the genus Hermiella have not yet beenrecorded (Fig. 5 and Table 3).

Spanish ostracod assemblages share more than 90% of thespecies with those described in the marly and marly calcare-ous sediments of the Brian-de-Vere, Valeyres and Barre àPecten formations of Quercy, southwesternAquitanian Basin,France (Cubaynes and Ruget, 1985; Bodergat et al., 1991,1998; Andreu et al., 1998). Their assemblages are dominatedby healdiids (Ogmoconchella gruendeli, O. transversa, O.pseudospina and Ogmoconcha gr. amalthei), together withsome large cytheraceans (Gramannella apostolescui) and cyp-ridaceans. No species of Hermiella have yet been describedeither (Fig. 5 and Table 3).

The Paris Basin assemblages show a higher similarity tothe Iberian Peninsula, with 80% of species in common (Fig. 5and Table 3). Pliensbachian carbonate and argillaceous sedi-ments from the Paris Basin yielded ostracod assemblages,which are characterised by a very important representationof cytherellids (Cytherelloidea), healdiids (Ogmoconcha,Ogmoconchella, Hermiella and Pseudohealdia), bairdiids andcytheraceans (Ektyphocythere, Kinkelinella and Graman-nella). However, the Paris Basin assemblages differ from the

Fig. 2. Dendogram from cluster analysis of the Pliensbachian ostracod assem-blages of the studied areas. The similarity Index used is Simpson and theclustering techniques are Single-Linkage, Complete-Linkage and Average-Linkage (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 deszones étudiées. Indice de similarité utilisé = Indice Simpson–Techniquesutilisées = Single-Linkage, average-linkage (UPGMA et WPGMA).

Fig. 3. Dendogram from cluster analysis of the Pliensbachian ostracod assem-blages of the studied areas. The similarity Index used is Dice and the clus-tering techniques are Single-Linkage, Complete-Linkage and Average-Linkage (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 deszones étudiées. Indice de similarité utilisé = Indice Dice–Techniquesutilisées = Single-Linkage, average-linkage (UPGMA et WPGMA).


Cordillera Iberica assemblages in the occurrence of Herm-iella (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 werelinked to the Spanish and French assemblages. Ostracodassemblages from the shales, interbedded limestones and blue-grey marls of the Brenha Formation from the Lusitanian Basin(Central Portugal) and the Whale Unit from the Grand Banksof the Newfoundland Basin (Canada), are dominated by cla-docopids (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 commonwith Spanish and French assemblages (Table 3). Canadianassemblages are characterised by the intermittent preserva-tion of their elements that result in assemblages impover-ished in both diversity and abundance.

Western and southwestern Ireland basins (Porcupine,Slyne, Erris and Donegal, Fasnet, Celtic Seas and CardiganBay) assemblages are clearly related to both former clusters(CI-PB-Q). Pliensbachian sediments are represented by aninterbedded shale and siltstone, fine-grained sandstonesequence of the marginal to shallow marine Scalpa Sand-stone Formation. To the south, in the Fasnet Basin, thisbecomes a nearshore marine argillaceous sequence, Croydeand Kikhampton formations (Ainsworth, 1986a, 1986b, 1987;Ainsworth et al., 1987). These sediments yield very rich anddiverse ostracod assemblages (Fig. 6) with Bairdiidae andHealdiidae as the dominant components (> 60%), togetherwith many new species of Liasina, Polycope and Pro-cytherura. These assemblages show high commonalties (more

Fig. 4. Dendogram from cluster analysis of the Pliensbachian ostracod assem-blages of the studied areas. The similarity Index used is Jaccard and theclustering techniques are Single-Linkage, Complete-Linkage and Average-Linkage (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 deszones étudiées. Indice de similarité utilisé = Indice Jaccard–Techniquesutilisées = Single-Linkage, average-linkage (UPGMA et WPGMA).

Fig. 5. Number of ostracod species recorded per Lower Jurassic ammonite zone in the Paris Basin and Quercy (France) and Cordillera Iberica (Spain). (Legendas 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 dansla Cordillière Ibérique (légende comme pour la Fig. 9).


Table 3Distribution of selected ostracod species in the Pliensbachian. Each number represents the sites where these species have been recordedDistribution des espèces d’ostracodes sélectionnées dans le Pliensbachien. Chaque numéro représente les sites où cette espèce a été enregistrée.

ostracod species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17Acrocythere gassumensis Michelsen, 1975 C

Acrocythere oeresundensis Michelsen, 1975 C

Acrocythere troesteri Riegraf, 1984 C C

Bairdia clio Bizon, 1960 C C C C C C

Bairdia crassa Drexler, 1958 (= B. fortis) C C

Bairdia donzei Herrig, 1979 C C C C

Bairdia guttulae Herrig, 1979 C C C C C

Bairdia hahni Lord and Moorley, 1974 C C C C

Bairdia kempfi Ainsworth, 1989 C

Bairdia michelseni Herrig, 1979 C C C C C

Bairdia molesta Apostolescu, 1959 C C C C C C C C C C C C C

Bairdia praehilda Herrig, 1979 C C C

Bairdia rostrata Issler, 1908 C C C C C

Bairdia thuringica Herrig, 1979 C C

Bairdia sp. A Ohm, 1986 C

Bairdiacypris anisica brevis Herrig, 1979 C C C C

Bairdiacypris triasica postera Herrig, 1979 C C C C C C

Bairdiacypris tumida Ainsworth, 1987 C C C

Bythocypris fabaeformis Drexler, 1958 C C

Cardobairdia liassica (Drexler, 1958) C C C

Cardobairdia Nr. 103 Klingler, 1962 C

Cardobairdia sp. K (Apostolescu, 1959) C C C C C C C

Cristacythere betzi (Klingler and Neuweiler, 1959) C

Cythere? terquemiana Jones, 1872 C

Cytherella lindseyensis Lord, 1974 C

Cytherella praecadomensis (Knitter and Riegraf, 1984) C C C

Cytherella toarcensis Bizon, 1960 C

Cytherelloidea anningi Lord, 1974 C C C C C C

Cytherelloidea drexlerae (Field, 1976) C C

Cytheropteron cavatum Michelsen, 1975 C C

Cytheropteron diversum Herrig, 1969 C

Cytheropteron foveolatum Michelsen, 1975 C C

Cytheropteron sp. A Riegraf, 1985 C

Ektyphocythere champeauae (Bizon, 1960) C C

Ektyphocythere luxuriosa (Apostolescu, 1959) C

Ekt. multicostata (Klingler and Neuweiler, 1959) C C

Ektyphocythere quadrata Boomer and Lord, 1988 C C C

Ektyphocythere aff. E. vitiosa (Apostolescu, 1959) C C C

Ektyphocythere vulgaris (Klingler and Neuweiler, 1959) C

Eucytherura liassica (Bate and Coleman, 1975) C C C C

Eucytherura tricostata (Michelsen, 1975) C C C C C C C

Fabalicypris symmetrica Herrig, 1979 C C C

Gammacythere ubiquita Malz and Lord, 1976 C C C C C C C

Gramannella apostolescui (Gramann, 1962) C C C C C C C C

Gramannella laevigata Michelsen, 1975 C C

Gramannella tatei (Gramann, 1962) C

Gramannicythere aubachensis Riegraf, 1984 C C C C

Gramannicythere bachi Herrig, 1982a C C C C C C

Gramannicythere coniuncta Herrig, 1982a C

Gramannicythere sp. Malz and Nagy, 1984 C

Hutsonia decorata (AP. Magné and Malmoustier, 1961) C

Isobythocypris cylindrica (Herrig, 1979) C

Isobythocypris dorsoconversa Ainsworth, 1986b C

Isobythocypris fabaeformis Drexler, 1958 C C

Isobythocypris pliensbachiensis Ainsworth, 1986 C

Isobythocypris unispinata Apostolescu, 1959 C C C C C

(continued on next page)


than 70%) with Portuguese assemblages and share 31% ofcommon species with the Cordillera Iberica (Fig. 6 andTable 3).

Relatively few publications have dealt with British Pliens-bachian ostracods. The cluster analysis shows that Britishassemblages are most similar to those of the Paris Basin andthe Cordillera Iberica. The broad belt of Lower Pliensba-chian clay-shale and sands deposits from Belemnite Marls,Green Ammonite Beds, Three Tiers, Eype Clay, Down CliffSands formations (Howarth, 1978) and the Upper Pliensba-chian yellow weathering sands of the Thorncombe Sands andthe 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 areLiasina lanceolata and some cytheraceans (Gramannellaapostolescui, Kinkelinella tenuicostata, Ektyphocytherequadrata and E. champeau) (Fig. 6 and Table 3). The LowerJurassic sequence of grey mudstones, siltstones and occa-sional limestones of the Mochras Borehole, on the NorthWales coast of Cardigan Bay, yielded an interesting assem-blage that differs from the other British assemblages (Boomer,1991; Boomer and Ballent, 1996) in its large number ofcytheraceans (Eucytherura liassica, Cytheropteron cava-tum, Gammacythere ubiquita, Gramannella apostolescui andLophodentina striata). Healdiids (Ogmoconchella adenticu-lata, Ogmoconcha convexa and O. contractula) are also abun-dant. Since British assemblages share less than one third of

Table 3(continued)

ostracod species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17Isobythocypis tatei (Coryell, 1963) C C C C C

Kinkelinella tenuicostata Martin, 1960 C C C

Klinglerella elongata Michelsen, 1975 C C

Klinglerella foveolata Michelsen, 1975 C C C

Klinglerella herrigi Ainsworth, 1989 C

Klinglerella intermedia (Klingler and Neuweiler, 1959) C

Klinglerella lacunosa Ainsworth 1989 C

Klinglerella katsloesensis Sivhed 1980 C

Klinglerella moorei (Brady, 1872) C C

Klinglerella variabilis (Klingler and Neuweiler, 1959) C

Ledahia conveniens Herrig, 1980 C C C

Ledahia septenaria (Gründel, 1964) C C C C C C

Liasina lanceolata (Apostolescu, 1959) C C C C C C C C

Liasina vestibulifera Gramann, 1962 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.

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).


their species with other areas (Figs. 6–8), they form an inde-pendent group under the Dice and Jaccard indices. Using theSimpson Index, British assemblages have been included inthe same cluster as the Cordillera Iberica, because Spanishassemblages have almost 40% cent of species common withBritish assemblages.

Ostracod assemblages described from Northeast and Cen-tral Europe show a higher similarity, and this is reflected in amajor cluster. Danish Embayment and southern Sweden basinassemblages are sparse and poorly preserved. The Pliensba-chian Fjerritslev Formation in the Norwegian-Danish basinsconsists of a relatively uniform series of marine claystone orshale, with a varying silt content, sandstone intercalations andcalcareous beds (Michelsen, 1975, 1978). The assemblagesshare almost 50% species with the Cordillera Iberica (Table 3)and are dominated by healdiids and cytheraceans (cytheru-rids, bythocytherids), and significant numbers of bairdi-aceans and polycopids (Fig. 9). Danish assemblages are alsobroadly comparable in composition with those described fromGermany and Switzerland.

The deposition of black shale (Posidonienschiefer facies)during the Sinemurian-Middle Aalenian characterised Ger-man and Swiss sedimentation and their ostracod assem-blages are well known (Fischer, 1961b; Riegraf et al., 1984).Lower Pliensbachian offshore clay shales of the Schwarzjura-tone Beta and Schwarzjuratone Lower Delta formations,together with the Upper Pliensbachian marly limestones withintercalated micritic limestones of the Schwarzjuratone Upper

Delta Formation yielded ostracod assemblages more diverseand 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 characterisedby the occurrence of healdiaceans (including Hermiella), largecytheraceans, cytherurids, bythocytherids, healdiids and bair-diids (Fig. 9) and share more than 40% of species with theCordillera Iberica (Table 3).

4.2. Toarcian

Toarcian ostracod assemblages are represented in the den-drograms (Figs. 7,8,10 and Table 4). They are grouped in twomajor clusters. They are, from the top to the bottom: (1) Cor-dillera Iberica to Great Britain (CI-GB) and (2) Canada-Portugal-Fasnet Basin (C-P-FB). In the first cluster, wherethe majority of the studied assemblages have been included,the highest similarity is observed between southwestern Ger-man and Swiss assemblages, with Spanish, French and Brit-ish assemblages included in the same cluster, although theyshow less similarity among them in relation to the Pliensba-chian.

Results obtained with the Jaccard index (Fig. 8) show thehighest value of similarity to be between the German andSwiss basins assemblages, which form a cluster that has itsmaximum similarity with the Paris Basin and British basinsassemblages. Canada, Portugal and the Fasnet Basin assem-blages form a different cluster that has a rather low similaritywith the cluster (CI-SWG-Sz-PB-GB-Q).

Fig. 7. Dendogram from cluster analysis of the Toarcian ostracod assembla-ges of studied areas. The similarity Index used is Dice and the clusteringtechniques are Single-Linkage, Complete-Linkage and Average-Linkage(UPGMA and WPGMA). Abbreviations, C, Grand Banks (Canada), CI, Cor-dillera Iberica, FB, Fasnet Basin, GB, Great Britain, Q, Quercy, PB, ParisBasin, SWG, Southwest Germany, Sz, Switzerland, P, Portugal.Fig. 7. Dendogramme des assemblages d’ostracodes du Toarcien des zonesétudiées. 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 assembla-ges of studied areas. The similarity Index used is Jaccard and the clusteringtechniques are Single-Linkage, Complete-Linkage and Average-Linkage(UPGMA and WPGMA). Abbreviations, C, Grand Banks (Canada), CI, Cor-dillera Iberica, FB, Fasnet Basin, GB, Great Britain, Q, Quercy, PB, ParisBasin, SWG, Southwest Germany, Sz, Switzerland, P, Portugal.Fig. 8. Dendogramme des assemblages d’ostracodes du Toarcien des zonesétudiées. Indice de similarité utilisé = Indice Jaccard–Techniques utili-sées = Single-Linkage, average-linkage (UPGMA et WPGMA).


Dendrograms (Table 4 and Fig. 10) based on the Simpsonindex contain two major clusters: (1) SW Germany-Canada-Switzerland (SWG-C-Sz) and (2) the rest of assemblages havebeen incorporated into the Cordillera Iberica-Fasnet Basin(CI-FB) cluster, which contains two subclusters: (2a) the Cor-dillera Iberica-Quercy-the Paris Basin (CI-Q-PB) and (2b)Great Britain-Portugal (GB-P). These clusters are differentfrom those obtained when the Dice and Jaccard indices wereconsidered. The different results can be explained in the lightof the differences in the size of the compiled samples, par-ticularly, in the case of Canadian and Portuguese assem-blages.

4.2.1. DiscussionThe Lower Toarcian carbonate platform facies of the Cor-

dillera Iberica, developed as an alternating series of marlylimestones and marls, formally named the Alternancia deMargas y Calizas de Turmiel Formation (Goy, 1974; Goy etal., 1976; Arche et al., 1977; Comas-Rengifo, 1985), yieldeda very rich and diverse ostracod fauna (Fig. 5) dominated byCytheracea (Arias and Comas-Rengifo, 1992; Arias et al.,1992; Arias and Lord, 1999a, 1999b). The TenuicostatumZone and the lower part of the Serpentinus Zone (FalciferumZone), ostracod assemblages are characterised by the occur-rence of Cytherella toarcensis, Liasina lanceolata, Graman-nella apostolescui, Ektyphocythere dharennsourensis, E. aff.E. vitiosa, and Ogmoconchella aff. O. aspinata, O. adenticu-lata and O. aequalis. In the base of the Serpentinus Zone,healdiids disappear, cypridaceans and platycopids continue

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.

Fig. 10. Dendogram from cluster analysis of the Toarcian ostracod assem-blages of studied areas. The similarity Index used is Simpson and the clus-tering 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. 10. Dendogramme des assemblages d’ostracodes du Toarcien des zonesétudiées. Indice de similarité utilisé = Indice Simpson–Techniquesutilisées = Single-Linkage, average-linkage (UPGMA et WPGMA).


decreasing, while the Bairdiacea increase (Bairdiacypris dori-sae). In the remainder of the Lower Toarcian, ostracod assem-blages are characterised by Cytheracea (Kinkelinella sermoi-sensis, Ektyphocythere bucki, E. anterocosta, andPraeschuleridea pseudokinkelinella).

The Upper Toarcian and Aalenian sediments of the Turm-iel and Chelva formations consist of rhythmic alternations ofmarls and limestones, mudstone and bioclastic wackestoneand yield ostracod assemblages comprising many of theLower Toarcian species plus Cytheropteron alafastigatum,Cytheropterina criba, Kinkelinella fischeri, Ektyphocytherefurcata, Praeschuleridea gallemannica, P. ventriosa, Oto-cythere callosa and Cytherella cadomencis (Goy et al., 1994,1996). Almost all species cited are common with the otherareas of this first (SWG-Sz-PB-GB) cluster (Fig. 5 andTable 5).

The Cordillera Iberica assemblages share more specieswith the Paris Basin (60% of their species) than with thosedescribed from Germany, Switzerland and Great Britain. Allthese areas (CI-PB-SWC-Sz) form a first independent clus-ter. Spanish assemblages, however, are less similar (share lessthan 30%) to Canadian and Portuguese assemblages, whichform an independent cluster together with the Fasnet Basinassemblages (C-P-FB). Both clusters result when the Diceand Jaccard indices have been used. In contrast with the resultsdescribed for the Pliensbachian, Toarcian ostracod assem-blages recorded from the Quercy area do not show the same

close similarity to the Cordillera Iberica assemblages and inthis way, they are linked to the previous cluster (CI-PB-SWC-Sz). The reasons may be in the reduced number of speciesthat have been described so far from this area.

Results obtained from the cluster analysis show the high-est values of similarity between southwestern German andSwiss assemblages. The Lower Toarcian black organic richclay and interbedded black-shale of the PosidonienschieferFormation yielded ostracod assemblages dominated by cyther-aceans of the genera Kinkelinella, Ektyphocythere, Praes-chuleridea, Monoceratina and Trachycythere. The UpperToarcian ostracod assemblages recorded from the marls, marlylimestones and interbedded calcareous shales of the UpperSchwarzjuramergel Formation and for the dark marine shalesof the Lower Opalinuston Formation (Knitter and Ohmert,1983, 1986; Knitter and Riegraf, 1984; Riegraf, 1985; Rich-ter, 1987) are characterised by cytheraceans of the generaPraeschuleridea (P. angulata, P. bernierensis, P. ventriosa)and Kinkelinella (K. sermoisensis, K. costata); Cytherel-loidea cadomensis is also well represented (Table 5 andFig. 11). These assemblages show a high similarity to theCordillera Iberica, with almost with 40% of common spe-cies.

The other difference in clustering results between the Diceor Jacard indices and the Simpson index concerns the clusterfor southwestern German and Swiss assemblages (SWG-Sz). These assemblages are linked with Canada when the Sim-

Table 4Values of the similarity indices between the Toarcian ostracod assemblages, which have been described in the different sample localities formerValeurs 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 0,675675676 0,923076923 0,575 0,368421053 0,631578947 0,466666667 C. Iberica

* 1 0,648648649 0,461538462 0,575 0,421052632 0,578947368 0,933333333 SW Germany

* 0,135135135 0 0,125 0,210526316 0,263157895 0,088888889 Canada

* 0,615384615 0,525 0,473684211 0,578947368 0,577777778 Paris Basin

* 0,25 0,157894737 0,263157895 0,133333333 Quercy

SIMPSON * 0,421052632 0,631578947 0,4 Great Britain

* 0,421052632 0,155555556 Fastnet Basin

Toarcian * 0,222222222 Portugal

* Switzerland

* 0,270588235 0,074074074 0,46728972 0,289156627 0,418181818 0,157303371 0,269662921 0,365217391 C. Iberica

* 0,169491525 0,527472527 0,179104478 0,489361702 0,219178082 0,301369863 0,848484848 SW Germany

* 0,238095238 0 0,222222222 0,333333333 0,416666667 0,16 Canada

* 0,32 0,545454545 0,321428571 0,392857143 0,634146341 Paris Basin

* 0,377358491 0,1875 0,3125 0,206896552 Quercy

DICE * 0,271186441 0,406779661 0,423529412 Great Britain

* 0,421052632 0,28 Fastnet Basin


* Switzerland

* 0,156462585 0,038461538 0,189393939 0,126315789 0,172932331 0,072916667 0,118811881 0,154411765 C. Iberica

* 0,078125 0,208695652 0,082191781 0,196581197 0,098765432 0,130952381 0,29787234 SW Germany

* 0,106382979 0 0,1 0,142857143 0,172413793 0,074074074 Canada

* 0,137931034 0,214285714 0,138461538 0,164179104 0,240740741 Paris Basin

* 0,158730159 0,085714286 0,135135135 0,09375 Quercy

JACCARD * 0,119402985 0,169014085 0,174757282 Great Britain

* 0,173913043 0,122807018 Fastnet Basin


* Switzerland


Table 5Distribution of selected ostracod species in the Toarcian. Each number represents the sites where these species has been recordedDistribution des espèces d’ostracodes sélectionnées dans le Toarcien. Chaque numéro représente les sites où cette espèce a été enregistrée.

SPECIES LUSITANIA GRAND SICILY STRETTURA MOROCCO ALGERIABASIN BANKS

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. pseudokinkelinellaPraeschuleridea ventriosa -------------Pseudohealdia etaulensis -------------Pseudom. subtriangularis -------------Ptychobairdia aselfingensis -------------Ptychob. schaubergensis -------------


pson index is used because Canada shares its entire low diver-sity fauna with these areas. Thus, the percentage of commonspecies of both central European areas with Canada is higherthan with the remaining areas, although the number of com-mon species is very low. However, when the comparison ismade with more diverse assemblages, such as, the Paris Basin,Great Britain or the Cordillera Iberica, German ostracodassemblages share a large number of species, although thepercentage of common species (total) is smaller than withthe Canadian assemblages. This explains why because thesouthwestern German and Swiss assemblages (SWG-Sz) forma cluster with the Paris Basin, Great Britain and the Cordil-lera Iberica assemblages with Jaccard or Dice indices, butwith Canada with the Simpson index.

French assemblages (Fig. 5) from the Paris Basin areclosely related to the Cordillera Iberica and Great Britain (CI-GB) cluster, and shares more than 30% species. However thereis a lack of data for the beginning of the Toarcian. The Spina-tum Zone regressive phase is reflected in non-deposition anderosion during the Tenuicostatum Zone, resulting in a barrenlevel. The rest of the Toarcian is characterised in many areasof the Paris Basin (during the Falciferum Zone) by the depo-sition of marls and marly limestones, together with blackshales (″Schistes carton″). Toarcian ostracod assemblages(Cubaynes and Ruget, 1985; Bodergat and Donze, 1988;Bodergat et al., 1985) are less diverse and become dominatedby cytheraceans (Ektyphocythere neumannae, E. quadrataand Pleurifera harpa), small cytherurids (species of Cythe-ropteron, Infracytheropteron, Rutlandella, Eucytherura),cytherellids and bythoceratids after the disappearance of thehealdiids 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 withthe Cordillera Iberica than in the Pliensbachian. Lower Toar-cian deposits in the Quercy area (Cubaynes and Faure, 1989;Cubaynes, 1986) comprise marly and marly-calcareous sedi-ments of the Penne and Lexos formations. At the base, greenmarls and bioclastic limestones of the Barre à Pecten (UpperPliensbachian-Lower Toarcian) contain only a few species,such as, Ogmoconcha aequalis, Cytherella praecadomensisand Kinkelinella sermoisensis. The Penne Formation includesblack-shales of the Schistes carton (Tenuicostatum Zone) andmarls and marly limestones (Serpentinus to Insigne ammo-nite zones) and yield a rich and diverse ostracod fauna(Cubaynes, 1986; Andreu et al., 1995) dominated by largecytheraceans (Kinkelinella, Ektyphocythere, Praeschuleridea,Camptocythere), meanwhile cytherellids and bairdiids(Cytherella toarcensis, C. praecadomensis and Bairdiacyp-ris dorisae) are very common. These assemblages contain alarge number of species in common, almost a 75%, with theCordillera Iberica (Table 5 and Fig. 5).

British assemblages show many similarities with the aboveassemblages, especially with those from the Paris Basin, andhave more than 50% species in common with the CordilleraIberica (Table 5). In the south of England the succession ofthin, hard, micritic limestones, separated by non sequencesand marked erosion planes in the Lower Toarcian JunctionBed (Hesselbo and Jenkyns, 1995) yielded only a few speciesof Ogmoconcha, Ogmoconchella and Hermiella, which dis-appeared at the base of the Falciferum Zone (Boomer, 1992),followed by an increase in the Cytheracea. A better knowl-edge of the Lower Toarcian ostracod assemblages wasobtained from the study of seven boreholes in the East Mid-lands (Central England) and a section at Ilminster (South-

Fig. 11. Number of ostracod species recorded per Lower Jurassic ammonite zone in Northwest Germany, Southwest Germany and Switzerland. (Legend as forFig. 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).


west England). Their assemblages comprised a rich anddiverse ostracod fauna (Bate and Coleman, 1975; Boomer,1992) characterised by the occurrence of the cytheraceans(Kinkelinella sermoisensis, K. costata, Ektyphocythereanterocosta, E. intrepida, E. debilis, Praeschulerideapseudokinkelinella), bairdiids and cytherellids, showing alarge similarity with German and French assemblages andwith more than 50% of species common with the CordilleraIberica (Table 5 and Figs. 7,8,10).

Ostracod assemblages from the Grand Banks (Canada),Zambujal (Portugal) and the Fasnet Basin were grouped intoone cluster (Figs. 7,8 and 10). Toarcian sediments of theBrenha Formation and the Whale Unit in Zambujal and theGrand Banks are represented by a sequence of shales, marlsand limestones, with no black-shale deposition (Mouterde etal., 1971; Exton, 1979; Lord, 1982; Exton and Gradstein,1984; Boomer et al., 1998) which have yielded ostracodassemblages dominated by species of Bairdiacypris, largecytheraceans and many species of Paracypris. Both assem-blages share more than 50% of species with the CordilleraIberica (Tables 5 and 6).

Inner shelf Toarcian-Aalenian dark calcareous bitumi-nous shales of the Stratton Formation in the Fasnet, Celticand Cardigan Bay basins, yielded less abundant assem-blages, where healdiids are absent and large Cytheracea domi-nate, with many new species of Bairdia, Cardobairdia, Iso-bythocypris, Bairdiacypris, Liasina and Procytherura(Ainsworth, 1986b; Ainsworth et al., 1987) (Tables 5,6 andFig. 6). These assemblages have only minor similarity withthose described from the Cordillera Iberica, with less than20% of their ostracod species in common.

5. The distinction of the Tethyan ostracod province

Although our knowledge of Tethyan ostracod assem-blages has been improved during recent years they are con-siderably less well known than western European assem-blages. In many cases this is due to the small number ofspecies recorded and/or to their poor preservation. Tethyanassemblages have been described from several localities inItaly 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 ofmembers of the superfamilies Bairdiacea and Healdiacea, witha lesser representation of Cytheracea (Bate, 1977; Lord, 1988;Arias, 1993). Only a few species recorded in the Ragusa-1 borehole (Barbieri, 1964) have been also described in theCordillera Iberica. In the Umbria-Marche area, northernApen-nines, Pliensbachian pelagic sedimentation represented bymicritic limestones with chert nodules of the Corniola For-mation and clayey and marly sediments of the Marne delMonte Serrone Formation (Farinacci et al., 1978; Colacicchiet al., 1988) yielded assemblages dominated by healdiids, bair-diaceans (Cardobairdia, Bairdiacypris and Bairdia) and heal-

diids (Ogmoconchella, Ogmoconcha and Hermiella) and Lia-sina lanceolata (Lord, 1978 (in Farinacci et al., 1978); Arias,1993) (Tables 3 and 6). Italian assemblages are more similarto German assemblages than to those situated in the area ofthe Grand Banks-Portugal and they share more than 50% oftheir species with Spanish assemblages.

Ostracod assemblages from the shallow carbonate plat-forms of North Africa (Morocco and Algeria) show closeaffinities 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) andBassoullet et al. (1991) described assemblages from three sec-tions situated at rides sud-rifaines (Atlas Meseta and easternMoroccan Meseta). Upper Pliensbachian bioclastic lime-stones and Toarcian grey marls and marly limestones yieldedostracod assemblages characterised by the important occur-rence of cytheraceans, platycopids and healdiids, together withsome species of the genera Liasina, Bairdia and Bairdiacyp-ris (Table 6).

Ostracod assemblages from the western part of the OmanMeseta, have been studied by Maupin (1977). Upper Pliens-bachian assemblages are characterised by the occurrence ofhealdiids (Hermiella ambo, Ogmoconcha amalthei) and Toar-cian by species of Bairdia, Cytherella, Isobythocypris, Para-cypris, Liasina and some large cytheraceans. These assem-blages are very similar to those recorded by Baloge (1981)from the Rosso-Ammonitico of Western Algeria. UpperPliensbachian assemblages from this area comprise speciesof Hermiella, Ogmoconchella, Bairdia and Bairdiacypris andToarcian assemblages, which are of low diversity, are char-acterized by the lack of healdiids and the occurrence of Bair-diacypris, Cytherella, Polycope and Kinkelinella (Table 6).

However, a different record has been obtained from thestudy of a borehole (DSDP Leg 79, Site 547) located off NorthAfrica (Bate et al., 1984). Eighteen Jurassic samples (fromHettangian to Pliensbachian) yielded a very peculiar fauna,with many new ostracod taxa and with many left in opennomenclature and where smooth taxa dominate (healdiids andbairdiids). As a whole, these assemblages show only a fewspecies in common with previously recorded assemblagesfrom NW Europe.

A complete analysis of the Tethyan Lower Jurassic ostra-cod assemblages (Table 6) reflects a great similarity in com-position with those from Central Europe, although with dif-ferences in the representation of their taxa. The maindifferences are that the presence of large Cytheracea are lesscommon than in European assemblages and that ornamentedforms of bairdiids and healdiids (Ptychobairdia, Hermiella)are present even during the Toarcian. The present knowledgeabout Tethyan ostracod assemblages considered here doessupport the recognition of a separate Tethyan province.

6. Conclusions

This paper considers the degree of similarity among Euro-pean (and surrounding areas) Lower Jurassic Ostracoda


Table 6Distribution of the most representative Tethyan ostracod species considered in this studyDistribution des espèces téthysiennes d’ostracodes les plus représentatives considérées dans cette étude.

ostracod species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17Monoceratina amlingstadtensis Triebel and Bartenstein, 1938 C C C C

Monoceratina mesoliassica Triebel and Bartenstein, 1938 C

Monoceratina michelseni Riegraf, 1984 C C C C

Monoceratina seebergensis Triebel and Bartenstein, 1938 C

Monoceratina stimulea (Schwager, 1866) C

Monoceratina striata Triebel and Bartenstein, 1938 C C C C C

Monoceratina ungulina Triebel and Bartenstein, 1938 C

Nanacythere (D.) firma Herrig, 1969b C C C

Nanacythere (D.) fissicosta Herrig, 1969b C C C C C

Nanacythere (G.) minor Michelsen, 1975 C C

Nanacythere (N.) simplex Herrig, 1969b C C

Ogmoconcha (Hermiella) ambo Lord and Moorley, 1974 C C

Ogmoconcha (Hermiella) cincta Malz, 1975 C

Ogmoconcha (H.) circumvallata Dreyer, 1967 C C C C C

Ogmoconcha (Hermiella) cista Malz, 1975 C C

Ogmoconcha (Hermiella) comes Malz, 1975 C C C C C C C

Ogmoconcha (Hermiella) hyblea (Barbieri, 1964) C

Ogmoconcha (Hermiella) inflata (Ainsworth, 1987) C C C

Ogmoconcha (H.) intercedens Dreyer, 1967 C C C C C

Ogmoconcha (Hermiella) klingleri Malz, 1975 C C C C

Ogmoconcha amalthei (Quenstedt, 1967) C C C C C C C

Ogmoconcha amalthei rotunda Dreyer, 1967 C C C

Ogmoconcha contractula Triebel, 1941 C C C C C C C C C C

Ogmoconcha convexa Boomer, 1991 C C C

Ogmoconcha dentata (Issler, 1908) C C

Ogmoconcha eocontractula Park 1984 C C C

Ogmoconchella aenticulata (Pietrzenuk, 1961) C C C C C C C C C

Ogmoconchella aequalis (Herrig, 1969a) C C C C C C C C

Ogmoconchella aspinata (Drexler, 1958) C C C C C

Ogmoconchella danica Michelsen, 1975 C C

Ogmoconchella impressa Malz, 1975 C

Ogmoconchella michelseni (Michelsen, 1975) C

Omoconchella mouhersensis (Apostolescu, 1959) C C C C C C

Ogmoconchella propinqua Malz, 1971 C C C

Ogmoconchella secunda Herrig, 1981a C C

Ogmoconchella sp. A Lord, 1974 C C

Ogmoconchella sp. B (Apostolescu, 1959) C C

Oligocythereis? Mochrarensis Boomer, 1991 C C

Paracypris redcarensis (Blake, 1876) C C C C

Paracypris sp. 1 Exton, 1979 C C

Paracypris sp. A Riegraf, 1985 C C

Pleurifera harpa harpa (Klinger and Neuweiler, 1959) C C C C C C C C

Pleurifera harpa harpoidea (Gramann, 1962) C C

Pleurifera vermiculata (Apostolescu, 1959) C C

Polycope cerasia Blake, 1876 C C C C C

Polycope cincinnata Apostolescu, 1959 C C C C C C C C

Polycope decorata Apostolescu, 1959 C C C C C C

Polycope discus Fischer, 1961a C

Polycope minor Michelsen, 1975 C

Polycope pelta Fischer, 1961a C C C C C C C C C C

Polycope plumhoffı Bate and Coleman, 1975 C C C C

Polycope tenuireticulata (Herrig, 1981c) C C

Progonoidea reticulata (Klingler and Neuweiler, 1959) C C

Pseudohealdia etaulensis (Apostolescu, 1959) C C C C

Pseudohealdia gruendeli Malz, 1971 C C C C

Pseudohealdia pseudospina (Herrig, 1969a) C C C C

(continued on next page)


assemblages, with particular attention to their respectiverelationships with the assemblages of the Cordillera Iberica(Spain). The results indicate that during the Pliensbachian,the Cordillera Iberica assemblages show a higher simi-larity with French basins (Quercy and the Paris Basin) thanthose described in the remaining areas of northwesternEurope.

Spanish assemblages are situated in the same major clus-ter that grouped all western European assemblages (French,Portuguese and British). It is remarkable that the grade ofsimilarity among western European assemblages is slightlylower than that which they show individually with Centraland northeastern Europe assemblages (Danish and Germanbasins). Canadian, Italian and Swiss assemblages show a lowgrade of similarity to remaining European assemblages, prob-ably as a consequence of our relatively poor knowledge ofthe faunas and their poor representation.

In the Pliensbachian, the Paris Basin assemblages form agroup with the rest of western European assemblages (PB-Q-CI-GB-P-FB), meanwhile southwestern German basins arewithin the cluster (NEG-NWG-SWG-Dk) with the nortwest-ern and Central European assemblages, indicating a first leveldisjunction. It is noteworthy that northwesternAfrican assem-blages are very similar to those described in Central Europe.Showing the occurrence of a open connection between Tethysand the epicontinental sea to the north during the Early Juras-sic.

For the Toarcian, the cluster analyses show the CordilleraIberica assemblages have a composition very similar to thoseof Eastern Europe (German, French and British basins) andtheir assemblages are included in the same cluster. Mean-while, Canadian, Fasnet Basin and Portuguese assemblagesare grouped together at a high level of similarity, forminganother major cluster. Thus, Toarcian Cordillera Ibericaassemblages are now more related to the Paris Basin assem-blages than they were during the Pliensbachian. Similarly,the Paris Basin assemblages show higher similarity to south-

western German assemblages in the Toarcian than theyshowed in the Pliensbachian. The results indicate that duringthe Pliensbachian there was a better communication amongwestern European areas and less between western and Cen-tral Europe than during the Toarcian. This latter communica-tion was increased during the Toarcian, meanwhile the mostwestern European areas (the Fasnet Basin, Portugal, Mochrasborehole, Canada) show a clear differentiation in relation tothe rest of western areas (Iberian Peninsula, French and Brit-ish Basins).

In the Lower Jurassic, especially in the Toarcian, the Euro-pean ostracod fauna show no clear differentiation into twomajor biogeographical provinces as do other groups of fossilinvertebrates (ammonites, brachiopods). It must be said, how-ever, that Tethyan ostracod assemblages, especially in thepost-Pliensbachian, are relatively poorly known.

Acknowledgements

The content and presentation of this paper was enhancedby the thoughtful and highly constructive reviews of Prof. A.Lord and Prof. E. Herrig to whom I owe our greatest debts ofgratitude. I owe a special debt to gratitude to Dr. Jean Vannierfor the critical comments of the early version of our paperand for his patience. I also want to thank, the members of theeditorial board of Geobios for their valuable assistance dur-ing all stages of this manuscript. C.A. owe special thanks toFabienne Ongaro for her excellent handling of the numerousmisprints and typographical errors. The ostracoda databaseis based on CA’s ostracod compilation over the past sevenyears. We acknowledge the TMR Mobility Program (Euro-pean Union), the Comunidad de Madrid postdoctoral Pro-gram and the DGCYT Projects PB93-0459, PB97-0274 andBTE 2001.1924 from the Ministerio de Educación (MEC) yCiencia and Ministerio de Ciencia y Tecnología of Spain(MCYT), for financial support.

Table 6(continued)

ostracod species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17Pseudohealdia pseudohealdidae (Gründel, 1964) C C C C

Pseudohealdia transversa (Gründel, 1964) C C C C

Pseudohealdia truncata Malz, 1971Pseudomacr. subtriangularis Michelsen, 1975 C C C

Trachycythere angusta Triebel and Klingler, 1959 C C C

Trachycythere horrida Triebel and Klingler, 1959 C C

Trachycythere tub. seratina Triebel and Klingler, 1959 C C C C

Trachycythere tub. tubulosa Triebel and Klingler, 1959 C C C C C C

Trachycythere verrucosa Triebel and Klingler, 1959 C C C C C

Trachycythere multistriata Michelsen, 1975 C C C C

Triassocythere? sp. 4135 (Michelsen, 1975) 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.


Annexe A. List of studied localities and references

PliensbachianNortheastern Germany (NEG) Pietrzenuk, 1961 Dobbertin and Eisenach

Dreyer, 1967 BrandenburgHerrig, 1969a,b GreifswaldHerrig, 1979 ThüringenHerrig, 1980 ThüringenHerrig, 1981a,b,c ThüringenHerrig, 1982a,b Thüringen

Northwestern Germany (NWG) Triebel and Bartenstein, 1938 Baden (Niedersachten)Triebel and Klingler, 1959 HannoverGramann, 1963 Weseker SattelMalz, 1971 NiedersachtenOhm (Fischer et al., 1986) Empelde

Southwestern Germany (SWG) Fischer, 1961b,c Baden-WürttembergKlingler, 1962 Baden-WürttembergLord and Morley, 1974 Baden-WürttembergMalz, 1975 Baden-WürttembergUrlichs, 1977 Baden-WürttembergRiegraf, 1984, 1985 Baden-WürttembergHarloff, 1993 Baden-WürttembergHarloff and Jagger, 1994 Kalk-Alpen Bayern and North Tyrol

Canada (C) Exton and Gradstein, 1984 Grand Banks, NewfoundlandCeltic Sea-Fastnet Basin-Porcupine (FB) Ainsworth and Horton, 1986 Fasnet Basin

Ainsworth, 1987 Fasnet BasinAinsworth et al., 1987 Fasnet BasinAinsworth, 1990 Porcupine, Slyne, Evris and Donegal

Danish Embayment, Denmark (Dk) Michelsen, 1975 Danish Embaymentand Skane, Sweden(S) Sivhed, 1980 Southern SwedenParis Basin, France (PB) Apostolescu, 1959, 1961 Paris Basin

Bizon, 1960, 1961 Basse NormandieApostolescu et al., 1961 Paris BasinBizon and Oertli, 1961 LorraineCousin and Apostolescu, 1961 ArdennesChampeau, 1961 Southeastern Paris BasinOertli and Grosdidier, 1961 Paris BasinMagné and Malmoustier, 1961 ThouarsMagné et al., 1961 ThouarsMaupin, 1978 VendéeDonze, 1985 FranceBodergat et al., 1985 Paris BasinRiegraf, 1985 Truc de BalducBodergat and Donze, 1988 Paris Basin

Quercy, France (Q) Cubaynes and Ruget, 1985 QuercyAndreu et al., 1998 QuercyBodergat et al., 1998 Quercy

Great Britain (GB) Lord, 1972, 1974 EnglandLord and Bown, 1987 Dorset and Isle of WightLord, 1978, 1988 Great BritainBoomer, 1991 Mochras, WalesBoomer, 1992 Ilminster, Southwestern England

Italy (It) Lord, 1978, 1988 StretturaArias, 1993 Umbria

Portugal (P) Exton, 1979 ZambujalExton and Gradstein, 1984 ZambujalBoomer et al., 1998 Zambujal

Northwestern Switzerland (Sz) Richter, 1987 Basel


Annexe B. Alphabetical list of species considered in this study

A.1. List of the taxa

Acrocythere gassumensis Michelsen, 1975Acrocythere mimica Bate and Coleman, 1975Acrocythere oeresundensis Michelsen, 1975Acrocythere rectangula Michelsen, 1975Acrocythere reticulata Knitter, 1983Acrocythere troesteri Riegraf, 1984Ambigocythere concentricostata Herrig, 1985Aphelocythere kuhni Triebel and Klingler 1959Bairdia cariata Drexler, 1958Bairdia clio Bizon, 1960Bairdia crassa Drexler, 1958Bairdia donzei Herrig, 1979Bairdia fortis Drexler, 1958Bairdia guttulae Herrig, 1979Bairdia hahni Lord and Morley, 1974Bairdia kempfi Ainsworth 1989Bairdia michelseni Herrig, 1979Bairdia molesta Apostolescu, 1959Bairdia ohmerti Knitter, 1983Bairdia praehilda Herrig, 1979Bairdia rostrata Issler, 1908Bairdia sp. A Ainsworth, 1986b

ToarcianSouthwestern Germany (SWG) Fischer, 1961a,b Baden-Württemberg

Knitter, 1983, 1984 Baden-WürttembergKnitter and Riegraf, 1984 Baden-WürttembergRiegraf, 1984, 1985 Baden-WürttembergHarloff, 1993 Baden-Württemberg

Canada (C) Exton and Gradstein, 1984 Grand Banks, NewfoundlandCeltic Sea-Fastnet Basin-Porcupine (FB) Ainsworth and Horton, 1986 Fasnet Basin

Ainsworth et al., 1987 Fasnet BasinAinsworth, 1990 Porcupine, Slyne, Evris and Donegal

Paris Basin, France (PB) Apostolescu, 1959 Paris BasinApostolescu, 1961 South of Paris BasinChampeau, 1961 Southeastern Paris BasinMagné et al., 1961 ThouarsOertli and Grosdidier, 1961 Paris BasinDepêche, 1985 FranceBodergat et al., 1985 Paris BasinRiegraf, 1985 Truc de BalducBodergat and Donze, 1988 Paris BasinBodergat et al., 1991 Paris Basin

Quercy, France (Q) Cubaynes and Ruget, 1985 QuercyCubaynes, 1986 QuercyBodergat et al., 1991 Quercy

Great Britain (GB) Lord, 1974 EnglandBate and Coleman, 1975 Empingham and UpwoodLord, 1978 Great BritainBoomer, 1991 Mochras, WalesBoomer, 1992 Ilminster

Portugal (P) Exton, 1979 ZambujalExton and Gradstein, 1984 ZambujalBoomer et al., 1998 Zambujal

Northwestern Switzerland (Sz) Richter, 1987 Basel


Bairdia sp. A Ainsworth, 1990Bairdia sp. A Ohm, 1986Bairdia thuringica Herrig, 1979Bairdia undulata Herrig, 1979Bairdia verrucosa Herrig, 1979Bairdia? eirensis Ainsworth, 1986aBairdiacypris dorisae (Knitter, 1984)Bairdiacypris rectangularis Ainsworth, 1986bBairdiacypris sartriensis Donze, 1966Bairdiacypris triangularis Ainsworth, 1986bBairdiacypris triasica postera Herrig, 1979Bairdiacypris tumida Ainsworth, 1987Bythocypris fabaeformis Drexler, 1958Cardobairdia fastnetensis Ainsworth, 1987Cardobairdia liassica (Drexler, 1958)Cardobairdia Nr. 103 Klingler, 1962Cardobairdia posteroprolata Ainsworth, 1987Cardobairdia sp. A Ainsworth, 1987Cardobairdia sp. K (Apostolescu, 1959)Cardobairdia toarcensis Ainsworth 1986bCristacythere betzi (Klingler and Neuweiler, 1959)Cristacythere costata Michelsen, 1975Cristacythere crassireticulata Michelsen, 1975Cytherella demiexensis Ainsworth, 1989Cytherella drexlerae Field, 1966Cytherella lindseyensis Lord, 1974Cytherella praecadomensis (Knitter and Riegraf, 1984)Cytherella toarcensis Bizon, 1960Cytherelloidea anningi Lord, 1974Cytherelloidea cadomensis (Bizon, 1960)Cytherelloidea circumscripta (Blake, 1876)Cytherelloidea drexlerae (Field, 1966)Cytherelloidea lacertosa Apostolescu, 1959Cytherelloidea modesta Apostolescu, 1959Cytheropteron alafastigatum Fischer, 1962Cytheropteron byfieldensis Boomer and Bodergat, 1992Cytheropteron cavatum Michelsen, 1975Cytheropteron diversum Herrig, 1969bCytheropteron foveolatum Michelsen, 1975Cytheropteron sp. B Riegraf, 1985Cytheropteron? sp. A Riegraf 1985Cytheropteron? sp. Boomer, 1991Ektyphocythere acuminata (Riegraf, 1984)Ektyphocythere ambo Boomer, 1988Ektyphocythere anterocosta Boomer, 1988Ektyphocythere bucki (Bizon, 1960)Ektyphocythere champeauae (Bizon, 1960)Ektyphocythere debilis Bate and Coleman, 1975Ektyphocythere dharennsourensis Boutakiout et al., 1982Ektyphocythere intrepida Bate and Coleman, 1975Ektyphocythere knitteri Riegraf, 1984Ektyphocythere lanceolata Boomer, 1988Ektyphocythere laqueata (Klingler and Neuweiler, 1959)Ektyphocythere luxuriosa (Apostolescu, 1959)Ektyphocythere multicostata (Klingler, and Neuweiler, 1959)Ektyphocythere neumannae (Maupin, 1978)


Ektyphocythere quadrata Boomer and Lord, 1988Ektyphocythere 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, 1985Fabalicypris symmetrica Herrig, 1979Gammacythere ubiquita Malz and Lord, 1976Gen ind. sp. A Boutakiout et al., 1982Gramannella apostolescui (Gramann, 1962)Gramannella laevigata Michelsen, 1975Gramannella tatei (Gramann, 1962)Gramannicythere aubachensis Riegraf, 1984Gramannicythere bachi bachi Herrig, 1982aGramannicythere bachi ornata Herrig, 1982aGramannicythere coniuncta Herrig, 1982aGramannicythere sp. Malz and Nagy, 1989Hemiparacytheridea (Tuberocytheridea) angulocostata Knitter, 1983Infracytheropteron gwashense Bate and Coleman, 1975Infracytheropteron pulchellum Bate and Coleman, 1975Infracytheropteron rarum Knitter, 1983Infracytheropteron supraliasicum (Herrig, 1981a)Isobythocypis tatei (Coryell, 1963)Isobythocypris aff. elongata (Blake, 1876) Sivhed, 1980Isobythocypris cf. elongata (Blake, 1876) Sivhed, 1980Isobythocypris cf. elongata (Blake, 1876) Drexler, 1958Isobythocypris cf. elongata (Blake, 1876) Michelsen, 1975Isobythocypris cylindrica (Herrig, 1979)Isobythocypris dorsoconversa Ainsworth, 1986bIsobythocypris elongata (Blake, 1876)Isobythocypris ovalis Bate and Coleman, 1975Isobythocypris pliensbachiensis Ainsworth, 1986bIsobythocypris unispinata Apostolescu, 1959Isobythocypris? sp. Dreyer, 1967Kinkelinella costata Knitter, 1984Kinkelinella kadeshensis Rosenfeld et al., 1987Kinkelinella persica Bate and Coleman, 1975Kinkelinella procera Herrig, 1985Kinkelinella sermoisensis (Apostolescu, 1959)Kinkelinella tenuicostata Martin, 1960Kinkelinella? idrissensis Boutakiout et al., 1982Klinglerella auleata (Klingler and Neuweiler, 1959)Klinglerella bipartita (Klingler and Neuweiler, 1959)Klinglerella elongata (Michelsen, 1975)Klinglerella foveolata Michelsen, 1975Klinglerella glabellata (Klingler and Neuweiler, 1959)Klinglerella herrigi Ainsworth, 1989Klinglerella intermedia (Klingler and Neuweiler, 1959)Klinglerella katsloesensis Sivhed, 1980Klinglerella lacunosa Ainsworth, 1989Klinglerella medioreticulata (Michelsen, 1973)Klinglerella moorei (Brady, 1872)Klinglerella sinemuriana Ainsworth, 1989Klinglerella sulcata (Klingler and Neuweiler, 1959)


Klinglerella translucens Ainsworth, 1989Klinglerella variabilis (Klingler and Neuweiler, 1959)Ledahia conviniens Herrig, 1980Ledahia septenaria (Gründel, 1964)Ledahia telata (Drexler, 1958)Liasina cf. vestibulifera Sivhed 1980Liasina cylindrica Ainsworth, 1986bLiasina lanceolata (Apostolescu 1959)Liasina vestibulifera Gramann 1963Monoceratina amlingstadtensis Triebel and Bartenstein, 1938Monoceratina frentzeni Triebel and Bartenstein, 1938Monoceratina mesoliassica Triebel and Bartenstein, 1938Monoceratina michelseni Riegraf, 1984Monoceratina seebergensis Triebel and Bartenstein, 1938Monoceratina stimulea (Schwager, 1866)Monoceratina striata Triebel and Bartenstein, 1938Monoceratina ungulina Triebel and Bartenstein, 1938Nanacythere (D.) firma Herrig, 1969bNanacythere (D.) fissicosta Herrig, 1969bNanacythere (G.) circumcostata Michelsen, 1975Nanacythere (G.) elegans (Drexler, 1958)Nanacythere (G.) elongata Michelsen, 1975Nanacythere (G.) minor Michelsen, 1975Nanacythere (G.) paracostata Michelsen, 1975Nanacythere (N.) simplex Herrig, 1969bNanacythere persicaeformis Riegraf, 1984Ogmoconcha (Hermiella) ambo Lord and Moorley 1974Ogmoconcha (Hermiella) cincta Malz, 1975Ogmoconcha (Hermiella) circumvallata Dreyer, 1967Ogmoconcha (Hermiella) cista Malz, 1975Ogmoconcha (Hermiella) comes Malz, 1975Ogmoconcha (Hermiella) hyblea (Barbieri 1964)Ogmoconcha (Hermiella) intercedens Dreyer, 1967Ogmoconcha (Hermiella) klingleri Malz, 1975Ogmoconcha amalthei amalthei (Quenstedt, 1967)Ogmoconcha amalthei rotunda Dreyer, 1967Ogmoconcha contractula Triebel, 1941Ogmoconcha convexa Boomer, 1991Ogmoconcha dentata (Issler, 1908)Ogmoconcha eocontractula Park, 1984Ogmoconcha hagenowi Drexler, 1958Ogmoconchella adenticulata (Pietrzenuk, 1961)Ogmoconchella aequalis (Herrig, 1969a)Ogmoconchella aspinata (Drexler, 1958)Ogmoconchella celticensis Ainsworth, 1989Ogmoconchella danica Michelsen, 1975Ogmoconchella gruendeli Malz, 1971Ogmoconchella impressa Malz, 1971Ogmoconchella michelseni Michelsen, 1975Ogmoconchella mouhersensis (Apostolescu, 1959)Ogmoconchella propinqua Malz, 1971Ogmoconchella scanica Sivhed, 1977Ogmoconchella serratostriata Ainsworth, 1989Ogmoconchella sp. A Lord, 1974Ogmoconchella sp. B (Apostolescu, 1959)Ogmoconchella sp. B Cubaynes, 1986


Ogmoconchella transversa (Gründel, 1970)Paracypris liassica (Bate and Coleman, 1975)Paracypris redcarensis (Blake, 1876)Paracypris sp. 1 Exton, 1979Paracypris sp. 2 Exton, 1979Paracypris sp. A Riegraf, 1985Paracypris sp. C Ainsworth, 1986bParacypris sp. Cubaynes, 1986Paracypris? longiformis Sivhed, 1980Paradoxostoma? fusiforme (Drexler, 1958)Paradoxostoma? pulchellum Michelsen, 1975Paranotacythere (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 1876Polycope cincinnata Apostolescu 1959Polycope decorata Apostolescu 1959Polycope discus Fischer 1961aPolycope minor Michelsen 1975Polycope pelta Fischer 1961aPolycope plumhoffı Bate and Coleman 1975Polycope tenuireticulata (Herrig, 1981c)Pontocyprella fabaeformis (Drexler, 1958)Praeschuleridea arguta Ainsworth 1986aPraeschuleridea aspera Knitter 1983Praeschuleridea bernierensis (Apostolescu, 1959)Praeschuleridea costata Ainsworth 1986aPraeschuleridea ellipsoidea Ainsworth 1986bPraeschuleridea gallemannica Malz 1966Praeschuleridea levita Ainsworth 1986bPraeschuleridea pseudokinkelinella Bate and Coleman 1975Praeschuleridea punctulata Plumhoff 1963Praeschuleridea reticulata Ainsworth 1986aPraeschuleridea sp. A Ainsworth 1986bPraeschuleridea ventriosa Plumhoff 1963Praeschuleridea whatleyi Ainsworth 1986bPraescleridea magnycourtensis (Apostolescu, 1959)Procytheridea laneuvevillae Donze 1967Procytheridea lotharingiae Apostolescu 1959Procytheridea multiforata (Klingler and Neuweiler, 1959)Procytheridea perplexa (Klingler and Neuweiler, 1959)Procytheridea spinaecostata (Klingler and Neuweiler, 1959)Procytheridea undulata Apostolescu 1959Procytheridea? jardensis Maupin 1978Procytherura celtica Ainsworth 1986bProcytherura euglyphea Ainsworth, 1986bProcytherura hastata Bate and Coleman, 1975Procytherura mediocostata Bate and Coleman, 1975Procytherura multicostata Ainsworth, 1986bProcytherura suebica (Herrig and Richter, 1990)Procytherura werneri (Riegraf, 1984)


Procytherura? exquisita Ainsworth, 1986bProcytherura? liassica Ainsworth, 1986bProcytherura? sp. Boomer 1992Progonoidea acuticostata (Klingler and Neuweiler, 1959)Progonoidea auleata (Gramann, 1962)Progonoidea polygonata Sivhed 1980Progonoidea 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, 1987Pseudohealdia transversa (Gründel, 1970)Pseudohealdia truncata Malz 1971Pseudomacrocypris cf. subtriangularis Michelsen 1975Pseudomacrocypris subaequalis Michelsen 1975Pseudomacrocypris? sp. A Ainsworth 1986bRenicytherura liassica (Bate and Coleman, 1975)Renicytherura tricostata (Michelsen, 1975)Rutlandella tatei Ainsworth 1986bTrachycythere tubulosa seratina Triebel and Klingler 1959Trachycythere tubulosa tubulosa Triebel and Klingler 1959Trachycythere verrucosa Triebel and Klingler 1959Triassocythere multiestriata Michelsen, 1975Triassocythere? sp. 4135 (Michelsen, 1975)Triceratina sp. Boomer 1992Wicherella semiora kirtonensis Lord 1972Wicherella semiora semiora Lord 1972

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Palaeozoogeography of Western European Lower Jurassic (Pliensbachian and Toarcian) Ostracoda