Vegetation Dynamics in Europe During the Neogene
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Johanna Kovar-Eder Staatliches Museum für Naturkunde, Stuttgart Vegetation dynamics in Europe during the Neogene Kovar-Eder, J., 2003 - Vegetation dynamics in Europe during the Neogene - in: Reumer, J.W.F. & Wessels, W. (eds.) - DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA. A VOLUME IN HONOUR OF HANS DE BRUIJN - DEINSEA 10: 373-392 [ISSN 0923-9308] Published 1 December 2003 Based on the leaf, seed-fruit, and pollen record, the main trends of vegetational evolution in South and Central Europe are outlined. Paleoclimatic parameters are inferred partly from the comparis- on of the fossil assemblages with modern vegetation units and their climatic requirements and part- ly by summarizing the available data from the literature. Correspondence: J. Kovar-Eder, Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, D- 70191 Stuttgart, Germany; email: [email protected] Keywords: Europe, Neogene, vegetation history INTRODUCTION ly restricted to only one of the plant organs In Europe, the Neogene is characterized by (leaf or seed-fruit or pollen assemblages, paleogeographic reorganization due to the respectively). As different plant organ assem- collision of the African with the Eurasian blages are bound to specific taphonomic con- plate. Orogenic belts evolved in central and ditions, the derived interpretations may diver- southern regions and large foredeep basins ge considerably. Although the different plant and wetland areas extended along the arising organs often co-occur, they are only rarely orogenes. Altitudinal vegetation zonations evaluated jointly. In such cases, they offer a successively developed in the new orogenes, detailed and complementary rather than con- and increasing altitudinal differences ultima- tradictory picture of the vegetation (e.g. tely constituted floristic barriers. In addition Buzek et al. 1985, Kovar-Eder et al. 1998). to the global climatic changes, the tectonic Differences in preservation, investigation development and sea level changes severely methods, reliability of age determination, and influenced the climate in Eurasia; the annual different "schools" of naming and treatment course of precipitation, its quantity, and its further complicate our profound large-scale regional distribution changed, thus also alte- understanding of the floristic and vegetational ring the temperature realm. development. The Neogene European floral records are very rich and diverse (leaves, seeds-fruits, THE MEDITERRANEAN pollen, wood). However, they are not equally distributed over the continent but rather clus- Early/Middle Miocene (Figs. 1-3) tered in regions with appropriate sedimenta- Macrofloras of this interval are relatively tion; some areas largely lack a fossil record. scarce in the Mediterranean region. On the In certain regions, the latest investigations Iberian Peninsula, the leaf-floras from date to the 19th and early 20th century. The Martorell (Vallès-Penedès Basin, surroun- fossil record and its interpretation are typical- dings of Barcelona) have been assigned to the 373 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003 Early and Middle Miocene (Sanz de Siria corresponding to the type Quercus sect. Ilex Catalan 1981a, 1982). The prevalence of and/or Suber is a remarkable feature. The flo- nanophyllous and microphyllous leaf size ras from Greece and former Yugoslavia also classes (see Glossary) there is interpreted to include relict taxa that were widespread in document warm and possibly seasonally dry earlier periods in more northerly European climatic conditions. regions, e.g. Platanus neptuni from Radoboj, The record from former Yugoslavia is com- Pancevo Bridge (Kovar 1982), or Calocedrus parably richer. Dating of Early Miocene in Kimi (Kvacek 1999). records has largely been based on their floris- Floristic and leaf-physiognomical differen- tic composition due to the lack of objective ces are clearly present between the correlation possibilities. Marine and brackish Mediterranean and Central Europe. The main sedimentation offers better age control of the differences are in the abundance of taxa that Middle Miocene floras, e.g. Pancevo Bridge fall into smaller leaf size classes (nano-, near Beograd (Pantic & Mihajlovic 1977), or microphyllous) and in the abundance of enti- Radoboj, Croatia (Pantic 1992). The floras re-margined leaves (usually evergreen) versus from the Balkan Peninsula and from Evia non-entire margined (often deciduous) in the Island (Aliveri, Velitzelos et al. 1992; Kimi, Mediterranean floras. However, floristic simi- Unger 1867) correspond physiognomically. larities may mask these differences at first Among these floras, the common presence of glance, e.g. due to the presence of azonal (see probably sclerophyllous (see Glossary) leaves Glossary) taxa such as Liquidambar, Figure 1 The flora of Martorell,Vallès-Penedès Basin, Spain, Figure 2 The flora from the Martorell,Vallès-Penedès Basin, Early Miocene (Sanz de Siria Catalan 1981), 0.8 X of the ori- Spain, Middle Miocene (Sanz de Siria Catalan 1982), 1 X; ginal plate size; composed of entire-margined small/narrow composed of entire-margined small/narrow leaves, leaf-size leaves, leaf-size class nano-/microphyllous. class nano-/microphyllous. 374 KOVAR-EDER: Neogene vegetation dynamics Populus, and Zelkova in the Mediterranean to trace floristic and vegetational differences and in Central Europe. Floristic and leaf-phy- not only in the Mediterranean but also siognomical differences are also present between the Mediterranean and Central between the West and East Mediterranean. In Europe (Gregor 1990: 335). this context, it is noteworthy that in the Early Aquitanian pollen profiles from southern Oligocene, the physiognomical similarities France (Provence, Languedoc) are interpreted between the Tard Clay flora (Hungary) and to reflect a littoral Avicennia mangrove; the Spanish record are closer than between megathermic taxa are reported to infer sea- the former and the Central European record. sonal tropical/subtropical conditions. The This indicates subxeric conditions for the maximum development of the Avicennia Western Mediter-ranean and the Tard Clay mangrove along the coast of Languedoc and flora (see chapter on Mediterranean vegeta- the best representation of megathermic taxa tion), although the floristic correspondence of are recognized during the Langhian. In the the Tard Clay flora is closer to Central Serravalian, the megathermic taxa and the Europe (Hably & Fernandez Marron 1998). Avicennia mangrove disappear, indicating cli- Long-distance correlation based on floristic matic deterioration (Bessedik et al. 1984). similarities as applied by Gregor (1990) is an The decrease in temperature and precipitation unsuitable approach for age determination regime is assigned to the Antarctic glaciation covering time-equivalent regional floristic and to changes in sea currents due to paleo- differences. Consequently, Gregor is unable geographic evolution (the isolation of the Figure 3 The flora of Kaludra, Serbia,Yugoslavia, Early Miocene (Mihajlovic 1988: pls. 1,2), composed of - broad-leafed (microphyl- lous), entire-margined, evergreen taxa: 1, 2 Lauraceae (2 Daphnogene), - broad-leafed (microphyllous/mesophyllous), non-entire- margined, deciduous taxa: 3 Populus, 4 Zelkova, 5 Liquidambar and - probably sclerophyllous taxa: 6 Quercus sect. Ilex and/or Suber, 7 entire-margined, small (nanophyllous) taxa of uncertain taxonomic affinity. 375 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003 Mediterranean and Indian Ocean) (Suc 1986). early Late Miocene (Sarmatian/Pannonian) The pollen record from Middle/Late correspond floristically as well. However, the Miocene (Aragonian/Vallesian) deposits in characteristic presence of sclerophyllous the southeastern part of the Duero Basin in Quercus sect. Ilex and/or Suber, for example, Spain is interpreted to indicate open wood- and of relicts such as Engelhardia and land and steppe (Rivas-Carballo 1991). Daphnogene along with laurophyllous species in the Italian and Greek record indicate a clo- Late Miocene (Figs. 4-6) ser relationship among the Mediterranean flo- The floral record of this time interval is far ras and climatic differences between the richer throughout the Mediterranean than the Mediterranean and more northerly European previous one. Based on the floras of different regions. Mai (1995) summarized the Italian Miocene age in the Vallès Penedès Basin, and Greek floras as the Likudi-Vegora floral Sanz de Siria Catalan (1993) traced clear complex. changes towards temperate climatic condi- tions in the Late Miocene. This is inferred Was there an influence of the "Messinian from the appearance of broad-leafed deci- salinity crisis" on the vegetation? Partly due duous taxa (leaf size class mesophyllous/ to the relatively short time interval (less than microphyllous) such as Acer tricuspidatum 2 MA), the Messinian floral record is not and Platanus leucophylla in the Late very rich (macro: Murviel-lès Béziers, Miocene (MN9) (Sanz de Siria Catalan 1981 b, 1993). The flora of Terrasse (MN10, Sanz de Siria Catalan 1997) yields also broad- leafed evergreen taxa (Lauraceae). In the lacustrine basin of La Cerdaña (Eastern Pyrenees), the Vallesian leaf and pollen flora of Vilella (Barrón 1999) supports this floristic and climatic interpretation. In the flora of Azambuja near Alenquer Tejo Basin (Lisboa region, Portugal, MN9), modern taxa (Castanea sativa, Ulmus campestris, Quercus ilex, Quercus suber) have been