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Relationships Between Ecosystems and Plant Assemblages As Responses to Environmental Conditions in the Lower Jurassic of Hungary and Romania

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Relationships Between Ecosystems and Plant Assemblages As Responses to Environmental Conditions in the Lower Jurassic of Hungary and Romania Acta Palaeobotanica 55(1): 3–17, 2015 DOI: 10.1515/acpa-2015-0005 Relationships between ecosystems and plant assemblages as responses to environmental conditions in the Lower Jurassic of Hungary and Romania MARIA BARBACKA1,2, MIHAI E. POPA3, JÓZEF MITKA4, EMESE BODOR 5,6 and GRZEGORZ PACYNA7 1 Hungarian Natural History Museum, Botanical Department, H-1476 Budapest, P.O. Box 222, Hungary; e-mail: [email protected] 2 W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Kraków, Poland 3 University of Bucharest, Faculty of Geology and Geophysics, Laboratory of Palaeontology, N. Bălcescu Ave. 1, 010041 Bucharest, Romania; e-mail: [email protected] 4 Jagiellonian University, Botanical Garden, Kopernika 27, 31-501 Kraków, Poland; e-mail: [email protected] 5 Hungarian Geological and Geophysical Institute, Geological and Geophysical Collections, 1143 Budapest, Stefánia út 14, Hungary 6 Eötvös Loránd University, Department of Palaeontology, 1117 Budapest, Pázmány Péter sétány 1/C, Hungary; e-mail: [email protected] 7 Department of Palaeobotany and Palaeoherbarium, Institute of Botany, Jagiellonian University, Lubicz 46, 31-512 Kraków, Poland; e-mail: [email protected] Received 24 April 2015; accepted for publication 26 May 2015 ABSTRACT. Two Early Jurassic localities, the Mecsek Mts in Hungary and Anina in Romania, are similarly significant and both floras are of autochthonous/paraautochthonous origin. In the Early Jurassic the Hungar- ian locality was a delta plain; the Romanian locality was an intramontane depression filled predominantly by a braided river system. The floristic composition of the two localities (52 genera, 120 species), although superfi- cially similar (25 common genera), differs at species level (only 9 common species) as well as in the proportions of taxa in major plant groups. These differences can be explained by differences in environmental conditions resulting from palaeogeographic and topographic factors. Based on previous and recent studies, alpha diversity as well as statistically (DCA, PCA) differentiated ecogroups are compared and discussed. For common species, the GLM method was used to classify them to particular environmental response types. Their environmental requirements in both ecosystems are evaluated. Some of the shared species showed different preferences at the localities, explainable by their broad ecological tolerance. KEYWORDS: macroflora, palaeoecology, Early Jurassic, Hungary, Romania INTRODUCTION A comparison of Hungarian and Roma- preservation (Popa 1998, Barbacka 2011). Both nian Early Jurassic localities is of interest of them have been systematically sampled for to us, since these localities are close to each more than 20 years. There are exhaustive lists other, the straight-line distance being ca of taxa recognised so far, and the geological 370 km. Both floras are of autochthonous/par- settings are well known (Barbacka 1991, 1992, aautochthonous origin, associated with coal- 1994a, b, 1997, 2000, 2001, 2002, 2009, Bar- bearing deposits, and with similar modes of backa & Bodor 2008, Bodor & Barbacka 2008, 4 Givulescu & Popa 1994, 1998, Kędzior & Popa turbed swamp (Komlopteris nordenskioeldii, 2013, Nagy & Nagy 1969, Popa 1992, 1997a, Elatocladus sp., Baiera furcata, Sphenobaiera b, 1998, 2000a, b, 2001a, b, 2005, 2014, Popa longifolia, Pagiophyllum sp., Brachyphyllum & Kędzior 2008, Popa & Van Konijnenburg- sp., Sphenobaiera leptophylla, Equisetites van Cittert 1999, 2006, Popa & Meller 2009, muensteri). Thévenard & Barbacka 2000). Statistical stud- Anina (formerly known as Steierdorf) is ies of both localities have distinguished plant a historical coal mining centre in the middle ecogroups (Barbacka 2011, Barbacka et al., area of the Reşiţa Basin, where the plant-bear- MS.). In the present work we applied statisti- ing Steierdorf Formation reaches 250 m thick- cal methods to both localities in order to com- ness (Bucur 1991, 1997, Popa & Kędzior 2008, pare their ecological backgrounds and plant Popa 2009). The Steierdorf Formation is coal- environmental responses. Although they are bearing, yielding eight bituminous coal seams, palaeogeographically close, they differ in their Hettangian-Sinemurian in age. The Steierdorf taxonomical composition and type of geological Formation is formed mainly by a braided river setting. system occurring in a depression during the The Hungarian locality in the Mecsek Mts Hettangian-Sinemurian, where mires, lakes, is a typical delta plain (facies: delta – limnic – flood plains next to levees, and river channels lacustrine – delta – lagoonal – plain marine – occurred (Popa 2009, Kędzior & Popa 2013). lagoonal – plain marine), which is Hettangian For plant ecology, PCA was performed (based and earliest Sinemurian in age. The recon- alike in Mecsek on co-occurrence of taxa in the structed palaeoenvironment was characterised same hand specimen), revealing four ecogroups as a system of river channels with river levees whose taxonomical composition depended and marine barriers, crevasse splays, swampy mainly on the moisture/disturbance gradient areas, lakes, and channels (Nagy & Nagy 1969, and the temperature gradient (Barbacka et al. Paál-Solt 1969, Barbacka 2011). Detrended in prep). correspondence analysis (DCA) of floral com- 1. Podozamites group, in moderately position (based on co-occurrence of taxa on the wet and disturbed habitat, not influenced by same slabs) gave five ecogroups interpreted as temperature (Podozamites paucinervis, Sphe­ depending on the two most important factors – nobaiera sp. and Pinites sp.). moisture and degree of disturbance (Barbacka 2. Schizoneura group. Lower tempera- 2011): ture and higher moisture/higher disturbance, 1. Sagenopteris group. Moderately dis- a typical flood plain association (Neocalamites turbed, relatively dry (non-flooded) inland (Schizoneura) carcinoides, Dictyophyllum nils­ areas (Nilssonia revoluta, Anomozamites mar­ sonii, Cladophlebis nebbensis, C. haiburnen­ ginatus, Cladophlebis denticulata, Maratti­ sis, Dictyophyllum nervulosum, Coniopteris opsis hoerensis, Sagenopteris sp., Nilssonia murrayana, Matonia braunii, Thaumatopteris obtusa). brauniana). 2. Thaumatopteris group. Highly dis- 3. Zamites group. Higher temperature turbed, short-lived, moderately wet areas and higher moisture, probably swampy in its formed by alluvial deposits (islands, peninsu- last, closing moments, when it was filled up las, forelands), fully damaged by river flooding, with sediment (Zamites schmiedelii, Baiera occupied by pioneer plants (Thaumatopteris sp., Ptilophyllum sp., Cladophlebis denticu­ brauniana, Phlebopteris angustiloba, Equiseti­ lata, Geinitzia sp., Ginkgoites sp., Komlopteris tes sp., Dictyophyllum rugosum, Cladophlebis nordenskioeldii, Ptilozamites cycadea). haiburnensis) 4. Nilssonia group. Relatively dry, moder- 3. Ptilozamites group. Weakly disturbed, ately warm and undisturbed conditions corre- moderately wet canopy (Nilssonia polymorpha, sponding with levees, which were the highest- Equisetites columnaris, Pterophyllum subae­ elevation relief forms in the basin (Nilssonia quale, Ptilozamites cycadea, Dictyophyllum sp. 1). nilssonii, Bjuvia simplex, Desmiophyllum sp., Since the climate of the Jurassic is known to Phlebopteris sp.) have been relatively stable (Vakhrameev 1991), 4. Ginkgoites group. Weakly disturbed microclimatic, palaeotopographic, or palaeo- wetland (Ginkgoites marginatus). geographic factors influenced the floristic com- 5. Komlopteris group. Moderately dis- position of particular localities. A genus-level 5 cluster analysis of European Jurassic localities same slab. The taxa were coded as binary (0-1) (Barbacka et al. 2014) placed the Reşiţa Basin variables (for details see Barbacka 2011, Bar- (including Anina) and the Mecsek Mts in the backa et al. 2014, Barbacka et al. in prep). In same branch: Reşiţa was paired with York- order to estimate the responses of particular shire (UK) and Mecsek with Scoresby Sound taxa along PCoA axis 1 and axis 2 we applied (Greenland), all being of the river-delta type a logistic regression model, the General Lin- of environment. On species level, Reşiţa and ear Model (GLM) using the logit link. A bino- Mecsek were in different clades, confirming mial distribution of the response variable a significant difference in their plant composi- was assumed (Agresti 2007). In that way the tion. Similarity of genus composition accompa- response variables (species) were related to nied by dissimilarity of species content is not a predictor – sample loadings along PCoA axis an unusual combination, as observed in a sta- 1 and axis 2 (Barbacka et al. in prep, Fig. 2). tistical approach to Mecsek flora (Barbacka Forward (stepwise) selection starting from the 2011). The same genera can occur in different null model was used to find the fitted model habitats, but species of the same genus almost for the particular species, based on the F-test always occupy different ecological niches (Bar- criterion and corresponding I-type error based backa 2011). on 499 runs. The calculations were performed In this paper we discuss the presumed with CANOCO 5 (ter Braak & Šmilauer 2012). taxonomical similarity between the two flo- Finally, seven species common to the two sites ras and the mechanisms governing local flo- were considered (see Table 1). GLMs for the ristical changes, in the light of environmental two localities revealed the response of the spe- variation. cies along the two PCoA axes and thus enabled us to classify them to particular
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