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European Jurassic Floras: Statistics and Palaeoenvironmental Proxies

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European Jurassic Floras: Statistics and Palaeoenvironmental Proxies Acta Palaeobotanica 54(2): 173–195, 2014 DOI: 10.2478/acpa-2014-0011 European Jurassic floras: statistics and palaeoenvironmental proxies MARIA BARBACKA1,4,*, EMESE BODOR 2,3,*, AGATA JARZYNKA4, EVELYN KUSTATSCHER 5,6, GRZEGORZ PACYNA7, MIHAI E. POPA8, GIOVANNI G. SCANU 9, FRÉDÉRIC THÉVENARD 10, and JADWIGA ZIAJA4 1 Hungarian Natural History Museum, Botanical Department, H-1476 Budapest, P.O. Box 222, Hungary; e-mail: [email protected] 2 Hungarian Geological and Geophysical Institute, Geological and Geophysical Collections, 1143 Budapest, Stefánia út 14, Hungary 3 Eötvös Loránd University, Department of Palaeontology, 1117 Budapest, Pázmány Péter sétány 1/C, Hungary; e-mail: [email protected] 4 W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Kraków, Poland; e-mail: [email protected] 5 Naturmuseum Südtirol, Bindergasse 1, 39100 Bolzano/Bozen, Italy 6 Department für Geo- und Umweltwissenschaften, Paläontologie und Geobiologie, Ludwig-Maximilians- Universität und Bayerische Staatssammlung für Paläontologie und Geobiologie, Richard-Wagner-Straße 10, 80333 München, Germany; e-mail: [email protected] 7 Department of Palaeobotany and Palaeoherbarium, Institute of Botany, Jagiellonian University, ul. Lubicz 46, 31-512 Kraków, Poland; e-mail: [email protected] 8 University of Bucharest, Faculty of Geology and Geophysics, Laboratory of Palaeontology, 1, N. Balcescu Ave., 010041, Bucharest, Romania; e-mail: [email protected] 9 Department of Chemical and Geological Sciences, University of Cagliari, via Trentino 51, I-09127 Cagliari, Italy; e-mail: [email protected] 10 UMR5276 CNRS, Laboratoire de Géologie de Lyon, Terre, Plančtes et Environnement, Université Claude Bernard Lyon 1, 69622 Villeurbanne cedex, France; e-mail: [email protected] Received 15 September 2014; accepted for publication 5 November 2014 ABSTRACT. The Jurassic floras of Europe show considerable diversity. To examine the extent of this diver- sity and its possible causes we used multivariate statistical methods (cluster analysis, PCA, NMDS) to com- pare all significant Jurassic floras in Europe. Data were based on 770 taxa from 46 fossiliferous occurrences (25 units) from France, Germany, Greenland, Hungary, Italy, Norway, Poland, Romania, Scotland, Serbia, Sweden, Switzerland, and the United Kingdom. Statistical analyses were applied at species level and genus level, and also performed for the major plant groups. The genus cladograms show affinities between differ- ent localities based on environmental factors, while the cladograms based on species affinities indicate only taxonomical correlations. The study shows that locality age does not seem to be of paramount importance for floral composition. KEYWORDS: flora composition, autochthonous and allochthonous floras, cluster analysis, PCA, NMDS, Jurassic, Europe INTRODUCTION In the last decades, as taxonomic studies of different floras/localities has become a focus of numerous Mesozoic localities have been com- interest. Different methods have been applied pleted, palaeoenvironmental reconstruction of to study plants’ adaptation to environmental conditions and the mechanisms of plant distri- * These authors contributed equally to this article and share bution and migration. Methods include analy- first co-authorship ses of macromorphology and cuticular structure 174 (Barbacka et al. 2006, Pott & McLaughlin 2009, compositional differences between the various Steinthorsdottir et al. 2011), traditional quanti- floras and/or time slices in relation to their tative methods (Knobloch & Mai 1986, Kostina environmental or latitudinal settings. Since & Herman 2013), comparisons between macro- the number of species and genera described and microfloras (Kustatscher et al. 2010), anal- from the different floras over the years is very yses of the affinities between palynomorphs high, only multivariate statistical methods can and their “mother” plants or sporomorph eco- be used. This is the first study comparing the groups (Abbink et al. 2004), and considerations various macrofloras and relating them to their of plants’ environmental requirements based on palaeoecological settings. statistical analyses (Spicer & Hill 1979, Rees et al. 2000, McElwain et al. 2007, Jasper et al. 2010, Barbacka 2011). Statistical analyses have MATERIAL AND METHODS also been used to compare floras on the global scale (Spicer & Hill 1979, Herman & Spicer The database comprises 770 taxa from 46 Early, 1997). These methods provide extensive infor- Middle and Late Jurassic localities in Europe mation helpful in understanding the relation- (Table 1). For statistical reasons, floras with less than ships between localities and taxa, and they can five well-defined species were not considered. The data are based mostly on classic and modern peer- indicate possible migration pathways and dis- reviewed works published up to 2013. For floras that tribution mechanisms, as is done for phytoprov- were revised the latest list of taxa was used. Records inces (Vakhrameev 1991). Significant palaeo- from classic works which were not re-examined or only geographic and palaeoclimatic hypotheses have partly re-examined were used in their original form, been proposed. supplemented by the new data. Because a large num- ber of references are cited, they are given in tables The aim of the present study was to com- (taxonomic references Table 2, environmental refer- pare fossil floras on a regional scale in Europe ences Table 3) and are not repeatedly mentioned in (except Russia) and to determine the most the text. Records from France, Italy, Poland, Romania, important factors influencing the similarities and Greenland contain, besides published data, also and dissimilarities between particular regions unpublished data on taxa from revisions, most of them done by us. Taxa with uncertain determinations (“cf.”) (i.e. palaeogeographical, temporal, environ- were recorded in the database as definite. Indetermi- mental, and climatic factors), and in particu- nable taxa (sp.) were included only when they were the lar the degree to which the palaeoenvironment only representatives of a given genus. In other cases influenced the plant composition at the sites. undetermined species (e.g. “sp.”, “sp. A”) were omitted. The study is based on literature data and on Outcrops of the same age and depositional set- ting in a given area were treated as single units and our unpublished data. Fortunately, in recent labelled as such; examples are Yorkshire (United King- times some classic localities (France, Germany, dom), Franken (Germany), Scania (Sweden), Hungary, Italy, Sweden, United Kingdom, some localities some Late Jurassic localities in France, the Holy Cross in Poland) have been taxonomically revised and Mts. (Poland), the Veneto area (Italy), and Serbia (see Table 3). re-examined according to the methodological The database was prepared using MS Excel soft- standards of modern palaeobotany and supple- ware in a 1/0 (presence-absence) matrix for species and mented by local environmental data. A number genera. The full database from the studied localities of new localities have been described (Serbia, is attached as a supplement (http://botany.pl/ibwyd/ some small Polish sites, Switzerland, Norway), acta_paleo/base1/act-p54_2_Barbacka.html). The statistical and mathematical analyses em- and work on some localities is in progress (Hun- ployed the R program (R Development Core Team gary, Romania). For all localities, more or less 2011). Sampling was affected by different factors at comprehensive information on their palaeoenvi- each locality (e.g. sample accessibility, size of outcrop, ronmental settings is available. subjective selection of material, preservation); this To date there has been no detailed study should be borne in mind when interpreting the re- sults. Rarefaction was not possible in this case as only correlating different European floras of the binary data were available, without abundance data. Jurassic period except for a recent comparison Statistical analyses: of Middle Jurassic floras of Europe and north- 1) Diversity indices. Alpha diversity (Bush & Bam- ern palaeo-Africa that demonstrated dissimi- bach 2004) was calculated for all taxonomical groups larity in the floras of this period (Scanu et al. in each locality. Both the total number of taxa and the alpha diversity of the localities and epochs were calcu- 2014). Such work is needed if we are to under- lated. Alpha diversity provides important information stand how uniform/diverse the floras of Europe about the locality but is strongly affected by differences were during the Jurassic, or to understand the in the manner of collecting fossils (open mines, dumps, Table 1. Localities and locality units, with their depositional environments during the Jurassic of Europe Locality/environment Lower Jurassic (L) Middle Jurassic (M) Upper Jurassic (U) Country depositional depositional depositional unit name localities unit name localities unit name localities setting setting setting Studzianna Huta Poland Holy Cross-L Odrowąż Gromadzice fluvial Grojec-M Grojec fluvial Wólka-U Wólka lagoon Chmielów Podszkodzie Causses-M Causses lagoon Armaille Causses basin allochthonous Creys France Vendée-L Deux-Sèvres Vendée lagoon – marine Jura-U lagoon – delta Orbagnoux Hettange Mamers-M Mamers lagoon – paralic basin Jura Anina Sirinia Romania Reşiţa-L fluvial Cristian Holbav Hungary Mecsek-L Mecsek Mts. delta Solnhofen Bayreuth Germany Franken-L fluvial Solnhofen-U
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