Field guide to the Neogene of the Island of Evia-Early Miocene flora of Kymi

Dimitrios Velitzelos; Chryssanthi Ioakim; Thomas Denk

The fossil plant assemblage of Steinheim am Albuch, reference locality of mammal Neogene zone MN 7, is revised. The plant-bearing calcareous marls (sulcatus-beds) were deposited in an impact crater lake underlying the mammalbearing strata (trochiformis- and oxystoma-beds). Currently, MN7-8 are correlated chronostratigraphically to the Serravallian and to the Sarmatian (Central Paratethys stage). The plant fossils are preserved as mere impressions, very poor in contrast. The assemblage includes aquatic plants from the lake and its margin as well as mainly leaves of woody taxa from the terrestrial surroundings. Aquatic plants include Characeae, Cladiocarya trebovensis, Phyllites sp. – ?floating leaf, Equisetum, ?Lythraceae, ?Potamogeton geniculatus, ?Trapa, and reed and sedge-like foliage (“Typha” latissima, Monocotyledoneae). In the terrestrial flora oaks are most diverse (Quercus drymeja, Q. ?gigas, Q. kubinyii, Q. mediterranea, Q. pseudocastanea). Foliage of ?Quercus sp. is more numerous than that of all other oak species but its systematic position remains unclear. Celtis japeti, C. lacunosa, Podocarpium podocarpum, Populus balsamoides, Zelkova zelkovifolia, and Sapindales leaflets of compound leaves (Sapindales fam., gen. et sp. 1 and 2) are well documented. Acer pseudomonspessulanum vel A. integrilobum, vel Acer angustilobum, ?Carpinus sp., Buxus pliocenica, Chaneya oeningensis, cf. Daphnogene polymorpha, Leguminosites cf. hesperidum, and “Parrotia” pristina are documented based on single specimens only. Not confirmed are, among others, the following taxa described by Schweigert (1993) which would be ecologically and/or climatically relevant: Isoëtes braunii recognised here as indeterminable remain, Isoëtes sp. as leaf rhachis and petiole of compound leaves, Smilax which is Celtis japeti, Pistacia miochinensis determined here as Sapindales fam., gen. et sp. 2, Zanthoxylum as ?Sapindales fam., gen. et sp. 1, “Dikotylenrest (Phyllocladium)” as fragment of a damaged leaf, and Eichhornia as indeterminable remain. The plant assemblage at Steinheim is peculiar in several respects: it lacks conifers as well as winged fruits except for Chaneya oeningensis and a poor fragment (?Ailanthus sp.). Lauraceae are practically absent except for a single specimen (cf. Daphnogene polymorpha). Betulaceae are documented by a single leaf (?Carpinus sp.) only, while taxa otherwise characteristic of this time interval like beech, elms and willows are completely absent. Assuming that the terrestrial plant record represents the close lake surroundings, the composition of taxa, leaf physiognomy and assumed autecology indicate subhumid sclerophyllous forest environment and seasonal changes in precipitation under warm-temperate climatic conditions. The Steinheim flora is compared to the regional floristic record and to selected European plant assemblages. • Key words: leaf flora, impact crater lake, vegetation reconstruction, middle Miocene.

The remarkable collections of fossil leaves of Messinian age (ca. 5,7 Ma) from two sites of the Vena del Gesso Fm. (Monte Tondo and Tossignano, Bologna and Ravenna provinces, Italy) are studied morphologically to assess the taxonomic composition of the assemblages and carry out a palaeoenvironmental analysis. The flora of Tossignano includes so far 42 taxa of vascular plants, with 10 conifers and 32 angiosperms (29 dicots and 3 monocots). In the plant material from Monte Tondo 72 taxa of vascular plants were recognized, with 9 conifers and 63 angiosperms (59 dicots, 3 monocots, 1 plant incertae sedis). Several fossil-species, well known in the Miocene of central Europe and Italy, have been detected. The most frequent ones are, at Monte Tondo, Pinus cf. rigios, Daphnogene polymorpha, Fagus gussonii, Platanus leucophylla and Quercus pseudocastanea, at Tossignano: Sequoia sp., Quercus roburoides, Ailanthus pythii. Other species are frequent in both sites: Taxodium dubium, Laurophyllum sp. A and B, Leguminosae gen. et sp. 1. The general palaeovegetation structure of the Monte Tondo and Tossignano plant assemblages corresponds to the vegetation transect of the Evaporitic Messinian with swamp, riparian vegetation, and zonal “subtropical humid forest”. For the last one, the results of IPR vegetation analysis indicate a transitional (ecotone) vegetation type between “Broad-leaved Evergreen Forest” and “Mixed Mesophytic Forest". The Coexistence Approach (CA) results indicate that both floras existed under comparable climatic conditions , with a MAT 14-16.2 °C for Tossignano and 16-16.5 °C for Monte Tondo, hence warmer than at present (MAT around 13.5 °C).

The early Burdigalian (MN3) plant assemblage of the Güvem area (northwestern Central Anatolia) is preserved in lacustrine sediments of the Dereköy pyroclastics. Its age is well constrained by radiometric dates of basaltic rocks bracketing the pyroclastics, making the Güvem flora one of the extremely few precisely dated early Miocene floras in the Mediterranean region. The rich assemblage of impression fossils comprises ferns and fern allies (2 species), gymnosperms (12 spp.) and angiosperms (129 spp.). Ilex miodipyrena sp. nov. is described as a new fossil-species. The most diverse families in the assemblage are the Fagaceae with 12 taxa and the Fabaceae with 12 leaf morphotypes and one fruit taxon. Aquatic plants are represented by seven taxa, riparian (including palms) and swamp forest elements by >35 taxa, and lianas by three taxa (Smilax spp., Chaneya). The relatively large number of aquatic and riparian/swamp elements is congruent with the rich fish, amphibian and reptile record of the Güvem area. Another characteristic feature of the plant assemblage is the presence of various lobed leaves which show similarities with modern species of different families (e.g. Alangium, various Malvales). Trees and shrubs growing on well-drained soils and forming closed-canopy and open-canopy forests are the most diversified group (>70 taxa). In terms of number of specimens in the collection and based on field observations, by far the most abundant leaf fossils belong to evergreen oaks of Quercus drymeja and Q. mediter-ranea and to various types of foliage that cannot be assigned to a particular extant or extinct genus of Fagaceae. These sclerophyllous trees must have covered vast areas surrounding the wetlands that developed during the early Miocene in the Güvem Basin. Based on a recent reassessment of the ecology and taxonomic affinity of these trees, they are considered to reflect humid temperate climatic conditions but with a brief drier season during the winter months. These forests are more similar to the laurel forests of the southeastern United States and those stretching in a narrow belt south of the Himalayas to eastern central China. The large number of Fabaceae may indicate the presence of warm subtropical environments but this is difficult to assess, as they are known for having wide ecological ranges today and in the past. All in all, a larger part of the plant taxa point to forested vegetation. This is in agreement with previous palynological studies which detected only small amounts of her-baceous and grass pollen. Open patches of vegetation may have been restricted to river banks and to rocky areas in a volcanic landscape. The biogeographic patterns detected for the early Miocene of the Güvem assemblage are manifold; most taxa are widespread Northern Hemispheric elements. A substantial part of the species migrated from Asia into Europe during the (late) Paleogene and reached Anatolia during the early Miocene (Fagus, Paliu-rus, Chaneya, Ailanthus, Quercus kubinyii, Davallia haidingeri, Acer angustilobum, A. palaeosaccharinum). Fewer taxa may have been in Anatolia before they migrated to Europe (e.g. Nerium, Smilax miohavanensis, Quercus sosnowskyi). Finally, very few taxa are Anatolian endemics (e.g. Ilex miodipyrena).

t AUGUST 29 - SEPTEMBER 2, 2002 ATHENS, GREECE Field guide to the Neogene of the Island of Evia Early Miocene flora of Kymi Geology and Stratigraphy: Palaeontology and Palaeobotany: Editing: Thomas Kottis Chryssanthi Ioakim Evangelos Velitzelos Zlatko Kvacek Thomas Denk Dimitrios Velitzelos Dimitrios Velitzelos KIRA • PANAG/:D 0 'i) Fig.l. The geological map of Euboea island Greece. Geological map of Greece. 1:50.000 Bomovas, 1., Rondoyanni - Tsiambaou Th. , IGME, 1983. Recent Formations Debris, Slide rocks, eluvial deposits and conglomerates Ultrabasic Conglomerates Mixed conglomerates (with basic and ultrabasic elements), sands, clays. (120 m) Second Lignite Formation Formation with sands, clays and marls with intercalations of mud and conglomerates on the top. Lignite seams. (120m) I Mud Formation Mud, partly with horizons of Ostracodes, Planorbis, flora remains. Intercalations of clays and sands. (90 m) Calcareous Mud Formation Calcareous mud with fossils and flora remains. First Lignite Formation Sands, clays, silts with fossils and flora remains. Lignite seams. (55 m) t Basement Formation Limestones, dolomites. Fig. 2. Synthetic stratigraphical column of the Neogene - Quaternmy deposits. (Yiakkoupis et. al. 1998). INTRODUCTION The island of Evia (Euboea) is the second largest Greek island facing the southeastern part of the Hellenic peninsula. It is situated between the peninsular massifs Othrys and Pelion in the northwest and the island complex in the southeast (Fig. 1). The Neogene of the island of Evia (Euboea) has attracted the attention of palaeontologists and geologists since the very beginning of the scientific exploration of this area in the 19th century. During the last few decades the intensity of the palaeontological research has increased and many new data have accumulated. Several new sites have been discovered and are currently under investigation. Neogene continental deposits containing fossils are exposed in various parts of the island and adjacent territories. One of the islands of particular interest is Illiodroma (i.e. Alonissos in the present geography of Greece), which is the type locality of the common Tertiary conifer Glyptostrobus europaeus. The excursion will concentrate on the central part of Evia, where the famous site of the Early Miocene flora and fauna of Kymi (or Kumi) is situated (Fig. 1). The exposures are easily accessible and promising for collecting fossils. A monograph of the flora is in preparation, and the participants of the excursion are kindly asked to let the collected specimens be identified by the excursion leaders. The partial lignite basin of Aliveri on the southwestern coast of Evia is filled with lignite-bearing deposits corresponding partly to the outcrops of Kymi. Fossil remains were explored there in the 1980-90s (e.g., Katsikatos et al. 1981, Benda and de Bruijn 1982, Gregor 1983, Velitzelos et al. 1992), when the lignite was exploited in the open pit Plakes at the Prinias village. Unfortunatelly, the fossiliferous marl is no longer accessible there. Therefore, the excursion will not include this area. GEOLOGY AND STRATIGRAPHY GEOTECTONICS From the viewpoint of geotectonics, southern Evia consisting of metamorphic formations is part of the crystalline massif Attica-Cyclades (Bornovas et al. 1983). It is regarded as a multiphase double window (Katsikatsos et al. 1986). The other part - the carbonate nonmetamorphosed series of the central and northern zones belong to the sedimentary series of the Inner Hellenides (Guernet 1971, Dercourt et al. 1977, Katsikatsos 1970) formely known as Subpelagonian zone (Aubouin 1957, 1963, Aubouin et al. 1963) or as East-Greek series (Renz, 1940). THE NEOGENE DEPOSITS According to Katsikatsos et al. (1981) all Neogene deposits of the island of Evia, except for minor brackish intercalations in a section near Vlachia, are of continental origin. The successions found in the three major sedimentary basins (the Aliveri-Kymi Basin, the Palioura-Gides Basin, and the Limni-Istiea Basin, (Fig.1) are lithologically roughly similar. In each of these basins the Neogene can be divided into a "lower" unit of predominantly finegrained lacustrine sediments, which locally contain lignite deposits locally, and an "upper" unit consisting of mainly coarse-grained fluviatile sediments. The lithological similarity of the various basins has generally been attributed to their similar overall geological history, and this interpretation has subsequently led to the time correlation of similar lithologies. The "lower" unit has been considered by most workers to be of Early Miocene age, the "upper" unit of Late Miocene age (Fig.2). These age determinations are based largely on the flora (Gaudry, 1860, De Saporta, 1868, Deprat, 1904, Guernet & Sauvage, 1969), but local occurrences of a mollusc fauna (GorceI 1878; Guernet 1971) and a mammal fauna (Mitzopoulos 1947, Deprat, 1904, CordelIa 1878, Woodward 1901) have also been considered. The occurrences of Neogene sediments near Styra near Almyropotamos along the southwestern coast and near Vlachica is of minor geographical as well as stratigraphical importance. They consist almost exclusively of coarsegrained red-bed deposits, except for the deposit near Almyropotamos. THE ALIVERI-KYMI BASIN GENERAL The Neogene sedimentary basin of Aliveri-Kymi (Fig.l) extends roughly in a northerly direction from north of Aliveri to the fault-controlled north-east coast. The on-land part of the basin covers an area of around 200 km 2 between the mountains of Mavrovouni in the northwest and those of Octonia in the southeast. It is not known how far this basin extends northwards into the Aegean Sea. The Neogene sediments can be divided into two units, which in the southern part of the basin, near Aliveri, are separated by a distinct unconformity. In the northern part of the basin, near Kymi, where the section is much thicker, these units pass gradually merge into each other. The lower unit. The total thickness of the predominated lacustrine sediments of the lower unit varies considerably from some 40m in the open cast mine of Aliveri to several hundreds of meters in the area of Kymi, but the general succession is similar throughout the area. From bottom to top can be distinguished a unit of alternating plastic clays, sandstone and conglomerates, which in the southwestern part of the basin and the nortrhwestern part of the basin is overlain by lignites. The geographically quite restricted lignite occurrences (Fig.2) are overlain by well stratified marls and marly limestones which at the lower parts are slightly folded. The sediments of the lower unit are pierced by small volcanoes in the area south of Kymi. The age of the volcanic rocks has been determined as around 13 Ma by Fytikas et a1. (1976). The upper unit. The contact between the predominantly fluviatile sediments of the upper unit and the lacustrine deposits of the lower unit is clearly unconformable in the area of the Aliveri lignite mine where the red-coloured sandy clays, sandstones, and conglomerates of the upper unit cover an erosional surface. The components of the conglomerates suggest that the source area for these deposits has to be searched for in the metamorphic rocks of southern Evia. The conglomerates, which contain boulders as large as 5 m 3 show a general trend to became coarser from bottom to top and from west to east. The thickness of the upper unit ranges from about 50 meters in the open-cast mine of Aliveri to over 200 meters in the area of Avlonori-Octonia. The age of the Neogene deposits of Aliveri-Kymi. The lower unit has been studied intensively because of the economic importance of the lignite. Palaeontological age determinations suggested for these deposits by previous workers are based primarily on the fossil flora. The lignite deposits near Aliveri and Kymi, which mainly consist of typical swamp vegetation, have been considered to be of Aquitanian age by De Saporta (1868), Guernet & Sauvage (1969), De Bruijn et al. (1980), Katsikatsos et al. (1981) Velitzelos et aI1982,1983,1985: Ioakim,1984,1988, Doukas, 1986). These authors have considered the whole sequence to be of Early Miocene age. Deprat (1904), however, placed the lacustrine marls, which conformably overlie the lignites into the Sarmatian. Fytikas et al. (1976) dated the volcanic rocks, which pierce the series south of Kymi to around 13 Ma, which determines a maximum age for the top of these deposits. PALAEONTOLOGY AND PALAEOBOTANY The French geologist Souvage, who noticed occurrences of fossil plants and fish remains in the marls overlying the lignite, discovered the palaeontological locality of Kymi on the Evia Island as early as 1846. Nearly 20 years later the fossil flora of Kymi was explored in more detail. Brongniart (in Gaudry 1860) was the first who reported on the plants of Kymi. Later studies were undertaken by the Austrian botanist and palaeobotanist Franz Unger (1861, 1862) and Gaston de Saporta (in Gaudry 1862). These two authorities worked independently and created many synonyms for plant taxa occurring in Kymi. Some of the type specimens for these pioneer studies are kept at present in the Natural History Museum in Vienna and Paris respectively and have been under the revision (Kvacek & Velitzelos 2000, Kvacek & Erdei 2001). A comprehensive monograph of the Kymi flora was presented by Unger (1867) and commented and complemented by Saporta (1868, 1973). Since that time extensive fieldwork and collections have been made and many specimens spread into various institutions and museums in Europe. Engelhardt (1910) and Frittel (1921, 1922) published additions to the Kymi flora. The material of Kymi is well recognizable by the characteristic firm yellowish marlstone, usually cut into rectangular slabs. Therefore, unlabeled specimens can be ascribed to Kymi in many cases (e.g. Erdei & Kvacek 1997). During the last decades the sites at Kymi have been under investigation by Prof. E. Velitzelos and his team and a couple of thousands of new samples including those suitable for cuticular studies have been gathered at the University of Athens (Velitzelos & Kvacek 1996). A comprehensive monograph of this material including the revisions of more important collections made earlier is planned for the near future. Several exposures in the surroundings of Kymi are accessible and local fossil assemblages slightly differ in composition and quality of preservation. About three levels/facies can be distinguished so far: 1) the lowermost layers of the marl along the coast (the Fagus level), 2) the lignite and the adjacent dark marl with plant compressions, and 3) the classical yellow, finely bedded marl, exposed at several places in the upper part of the section. The lowermost assemblage is dominated by Fagus leaves identical to Fagus pristina Saporta from Aquitanian deposits of southern France. Betulaceae leaf impressions are the second dominant plant group there along with leaves of Daphnogene. This assemblage has not been fully excavated and studied. The lignite has yielded a poor carpological assemblage containing a single element of common swampy coal-forming vegetation - Stratiotes sp. (Velitzelos & Gregor 1990). The adjacent marls further away from the lignite mine have yielded an assemblage of leaf compressions dominated by Myricaea, Lauraceae, and the Mediterranean-type of oaks. Leaves assigned to Myrica vindobonensis exhibit bi-seriate glandular trichomes. A lauraceous leaf type has been identified as Laurophyllum acutimontanum, typical of the European late Palaeogene, another broader form has yielded the cuticle structure of Laurophyllum pseudoprinceps. Additional elements are Glyptostrobus europaeus, Cupressus sp., Pinus engelhardtii,Acer tricuspidatum, Populus populina (Vienna collections). The vegetation of this level is reconstructed as a mixed evergreen and deciduous riparian forest on moist soils. The yellowish marl exposed at several places in the upper part of the slope has yielded the majority of fossils known from Kymi including fish skeletons, insects (Bachmayer et al. 1981) and plant remains. It contains a much more diversified flora than the other two parts. Unger (1861) was believing that he had found plant elements native today in the Southern Hemisphere, such as Callitris, Eucalyptus, Proteaceae, Nothofagaceae, etc. Gymnosperms: A peculiar member of the flora is cycadalean foliage described by Saporta (1874) as Encephalartos, confined to Africa today. A revision (Kvacek & Velitzelos 2000) has shown that it has much more in common with the genus Dioon growing in Central America and Mexico. Among conifers, Glyptostrobus , Tetraclinis , and species of Pinus prevail. Glyptostrobus in Evia develops relatively smaller cones and heteromorphic foliage , which is in contrast to Early Miocene specimens from Central Europe. Tetraclinis is represented mostly by the xeromorphic T. brachyodon. Only a single fragment of T. salicomioides has been recovered so far. Seed cones of Cupressus (or Chamaecyparis ?) were misidentified by Unger (1867) as Sequoia. In addition, a single twig of Calocedrus suleticensis was also found in the old collections in Vienna. Pine species occur rarely in this assemblage. They are representatives of either Pinus subgen. Pinus (i.e. Diploxylon) or subgen. Strobus (i.e. Haploxylon). Besides seed cones and seeds, also male cones and needles in fascicles of two, three, and five are found. Angiosperms: Among angiosperms, Myricaceae, Fagaceae, and Lauraceae predominate. Most taxa are based on leaf impressions and difficult to assign to modern genera. Several new elements have been recovered, and taxonomic affinities of many taxa have been re-evaluated (Velitzelos & Gregor 1990, Velitzelos & Kvacek 1996). A large-leaved Myrica solonis was identified originally as a Proteaceae by Unger, but soon transferred to Myricaceae by Saporta. Besides the foliage of Myrica lignitum is quite abundant, and a single fertile twig of the latter bearing fruits of Myrica ceriferiformis has been recovered. Two more morpho-species, Myrica vindobonensis and Comptonia difformis cooccur. Two leaf types belong to Alnus. The xeromorphic small-leavedA. cycladum has not been found elsewhere in Europe. The rarer large-leaved species A. gaudinii links the Kymi flora with other sites of the European mid-Tertiary. Two kinds of fructifications co-occur, larger and smaller ones. The larger form - A. kefersteinii most probably belongs to A. gaudinii. A species of Betula, Betula oreadum , which was wrongly identified by Unger as Carpinus, is rather common in Kymi. Among the Fagaceae, two morpho-species - Quercus mediterranea and Q. drymeja predominate. Both are interconnected by transitional leaf forms. Cupules of Quercus were wrongly identified by Unger as Nephelium and by Velitzelos, Gregor and Jahnichen (1983) as Aesculus. Another member of the Fagaceae has often been assigned to Laurus primigenia. Givulescu was able to demonstrate its affinities to Fagaceae. A smaller form of this kind of leaves resembles Trigonobalanopsis. The true leaves of Lauraceae mostly belong to Daphnogene polymorpha , a common species of the European Miocene. It occurs in various leaf forms, both broader and narrower ones. Another common European species, Laurophyllum pseudoprinceps, is also present in the assemblage. There may be more lauroids among impression material, which are not identifiable without cuticular anatomy. Leguminosae (Fabaceae) foliage and fruits are present, but by no means do dominate the assemblage. To evaluate their precise taxonomic affinities a special study would be required. A common recently re-evaluated element of the Kymi flora is the narrow-leaved species Berberis kymeana, originally assigned by Unger to the Proteaceae. This element is also represented in the Aquitanian of France (Lomatites sensu Saporta). Another broader-leaved Berberis has been recovered recently but is much rarer. Elements previously assigned to Proteaceae -seeds of the Embothrium salicinum-type are now re-interpreted as Saportaspermum. Common capsular fruits identified by Unger as Royena (Diospyros sensu Saporta) are similar to the theaceous genus Gordonia. Among accessory elements, which are represented by few specimens only, the following can be safely recognized so far. Very few leaves of Acer tricuspidatum have been recovered. Some of them have crenulate margins and resemble A. tricuspidatum forma crenatifolium. Leaves of Populus populina, Ziziphus paradisiaca , and Zelkova zelkovaefolia also occur, but rarely. Very rare fruits and leaflets of Engelhardia have been recovered. Finally, en do carps of Mastixicarpum (Velitzelos & Gregor 1982), a single infructescence of Tilia, fruits of Cedrelospermum, few samples of disseminules of Ceratostratiotes (identified by Gregor), asymmetrical leaflets assigned by Palamarev and Petkova to Cedrela, and a narrow-leaved Smilax can be added to the assemblage. There are several leaf forms of doubtful affinities. Unger assigned some of them to Rhus, Pittosporum, and even Nothofagus. Velitzelos, Gegor and lahnichen (1983) suggested one larger palmately compound leaf to be a leaf of Aesculus damaged by insects. Saporta compared the same type of leaves from France to Oreopanax of the Araliaceae. All these and many other leaf forms require more detailed studies. The so far revised elements of the Kymi flora do not show any affinities with the Southern Hemisphere taxa. Instead, the flora illustrates well the mid-Tertiary assemblages of the Mediterranean Province, and correlates particularly well with southern France (Aix-enProvence) and Bosna-Hertzegovina (Miljevina). Mediterraean, East Asiatic and North American geo-elements prevail there. The flora of Kymi can serve as a reference floral assemblage for the eastern Mediterranean Early Miocene (Kymi-Arjuzanx sensu Mai 1995). The vegetation corresponds to the broad-leaved nothophyllous to microphyllous evergreen forest. A subtropical sub-humid climatic regime without a pronounced arid period can be suggested as the most probable model for this part of Europe in Early Miocene times. The exact age of the Kymi flora is still not certain. Small mammals occurring at the site Kazarma near Kymi (Katsikatsos et al. 1981) suggested to belong to the MN4 zone, i.e. Ottnangian (Benda et al. 1982, de Bruin, personal communication 2002). The discovery of a mastixioid element characteristic of the Oligocene in the classical collections from Kymi made by Velitzelos and Gregor. (1982) allows to assume the time span Oligocene to Early Miocene for some of the lignite occurrences at Kymi. At present we are inclined to believe that the section at Kymi encompasses levels at the Oligocene/Miocene boundary (correlated with the Linz Floral Assemblage sensu Kvacek & Walther 2001), namely the lowermost strata, which yielded leaf fossils of Fagus. Sedimentation including the lignite and the overlying marlstone continued to the middle Early Miocene, i.e. Ottnangian. It is important to stress that so far not a single specimen of Eotngonobalanus furcinervis has been recovered at Kymi (Unger 1867: pl. 4, fig. 18 is a misidentification). This species is most characteristic of the Oligocene in Evros, northeastern Greece (Velitzelos et al. 1999) and elsewhere in Europe. It is also noteworthy that many taxa characteristic of Kymi are also known from Aixen-Provence in southern France, which is independently dated as latest Oligocene. Revised list of the flora of Kymi: "Encephalartos" gorceixianus Pinus holothana Pinus hampeana Pinus cf. engelhardtii Pinus spp. (needles, pollen cones, seeds) Cupressus sp. Tetraclinis salicomioides Tetraclinis brachyodon Calocedrus suleticensis Glyptostrobus europaeus ? Taxodium dubium vel Sequoia abietina? Platanus neptuni Alnus kefersteinii Alnus cycladum Alnus cf gracilis Alnus gaudinii (= Carpinus betuloides) Alnus kefersteinii Carpinus grandis Betula oreadum Myrica vindobonensis Myrica solonis Myrica lignitum/M. ceriferiformis (- Olea noti, Asclepias podatyrii, Neritinium longifolium etc.) Comptonia difformis f. dryandroides Engelhardia macroptera Engelhardia orsbergensis Fagus sp. Quercus drymeja Quercus mediterranea Trigonobalanopsis rhamnoides Fagaceae gen. Daphnogene polymorpha Laurophyllum pseudoprinceps Laurophyllum acutimontanum Laurophyllum sp. Berberis kymeana (= Grevillea kymeana) Berberis sp. Populus populina Cycadaceae Pinaceae Cupressaceae Taxodiaceae Platanaceae Betulaceae Myricaceae Juglandaceae Fagaceae Lauraceae Berberidaceae Salicaceae Tilia sp. Zelkova zelkovaeJolia Cedrelospermum aquense Saportaspermum sp. Cedrela attica ? Rosa sp. Leguminosae gen. div. Acer tricuspidatum Ziziphus paradisiaca Mastixicarpum cacaoides ? Aesculus sp. vel Araliaceae ? Gordonia sp. (Royena graeca) "flex" cyclophylla Smilax sp. Ceratostratiotes sinijanus Dicotylophyllum sp. div. Tiliaceae Ulmaceae Rosaceae Fabaceae Aceraceae Rhamnaceae Cornaceae (Mastixiaceae) Hippocastanaceae vel Araliaceae Theaceae Aquifoliaceae Smilacaceae Ceratophyllaceae A tentative revision of the plant fossils figured in Unger (1867) PI. 2 1, la ? Cupressus sp. (as Callitris brongniartii) 2 indet. (as Callitris brongniartii) 3 - 11 Glyptostrobus europaeus PI. 2 1,9 Pinus holothana 2- 8, 10-11 Pinus sp. (as Pinus holothana) 12 extant Pinus for comparison 13 - 14 Pinus hampeana 15 Pinus sp. (as Pinus hampeana) 16 ? Pinus sp. (as Pinus furcata) 17-21 Cupressus sp. (as Sequoia langsdorfii) 22-23 ? Sequoia abietina vel Taxodium dubium (as Sequoia langsdoifii) 24-25 Dicotylophyllum sp. (as Podocarpus taxites) PI. 3 1-4 Alnus kefersteinii (as Alnus sporadum) 5 reconstruction 6-7 Pinus sp. male cones (as Alnus sporadum) 8 Alnus gaudinii (as Alnus sporadum) 9-10 ? Alnus gracilis (as Alnus cycladum) 11- 22 Alnus cycladum 23 Carpinus grandis (as Carpinus betuloides) 24-26 ? Alnus vel Carpinus (as Carpinus betuloides) 27 (17) ? Quercus drymeja (as Carpinus betuloides) 28 ? Zelko va zelkovaefolia (as Carpinus betuloides) 29 ? Fagus sp. (as Carpinus betuloides) 30-34 Alnus gaudinii (as Cmpinus betuloides) 35-37 catkins of ? Carpinus or Ostrya PI. 4 1-9 Alnus gaudinii (as Carpinus betuloides) 10-16 Zelko va zelkovaefolia (as Planera ungeri) 17 Dicotylophyllum sp. (as Quercus cyclophylla) 18 ? Quercus drymeja (as Quercus furcinervis) 19 ? Quercus drymeja (as Fagus pygmea) 20-22,28-30 Myrica vindobonensis 23-27 Myrica lignitum (as Myrica vindobonensis) 31 -36 ? Laurophyllum pseudoprinceps (as Ficus aglajae) PI. 5 1-17 Quercus drymeja (as Quercus lonchitis) 18-20 Alnus gaudinii (as Quercus kamischinensis) 21 -22 Quercus sp., cupules (as Quercus lonchitis) PI. 6 1-22 Quercus mediterranea 23 -27 Quercus dlymeja (as Quercus zoroastri) 29 Populus populina (as Populus attenuata) 30 ? Populus stipule (as Populus attenuata) 31 Berberis kymeana (as Grevillea kymeana) PI. 7 1-10 Daphnogene polymO/pha (as Cinnamomum lanceolatum) 11-24 Daphnogene polymorpha (as Cinnamomum scheuchzeri) 25-30 Daphnogene polymO/pha (as Cinnamomum subrotundum) 31 ? Cinnamomum fruits (as Cinnamomum rossmaessleri) 32 Ziziphus ziziphoides f. paradisiaca (as Cinnamomum rossmaessleri) 33-38 ? Lauraceae vel Fagaceae (as Law'us lalages) 39 ? Laurophyllum pseudoprinceps (as Cinnamomum buchii) PI. 8 1-7 ? Lauraceae vel Fagaceae (as Launls primigenia) 8-10 ? Laurophyllum pseudoprinceps (as Laurus princeps) 11 Myrica lignitum (as Laurinastrum dubium) 12 ? Laurophyllum sp. (as Protaea graeca) 13 ? Dicotylophyllum sp. (as Persoonia euboea) 14 Comptonia difformis f. dlyandroides (as Dryandra thesei) 15 ? Laurophyllum sp. (as Grevillea kymeana) 16-31 Berberis kymeana (as Grevillea kymeana) 32-33 ? Pinaceae seeds (as Hakea attica) 34-35 extant Proteaceae for comparison • PI. 9 1-3 Myrica solonis (as Banksia solonis) 4-13, 15 Myrica lignitum (as Dryandroides hakeaefolia) 14 ? Quercus dlymeja (as Dryandroides hakeaefolia) 16-18 Comptonia difformis f. dryandroides (as Dlyandra ungeri) 19-22 Saportaspermum sp. (as Embothrium salicinum) 23 ? Cedrelospelmum aquense (as Embothrium boreale) PI. 10 1-12 Myrica lignitum (as Olea noti) 13-24 Myrica lignitum (as Asclepias podalyrii) 25 Myrica lignitum (as Nelitinium longifolium) 26 ? Trigonobalanopsis rhamnoides (as Apocynophyllum carissa) PI. 11 1-3 Laurophyllum sp. (as Sideroxylon putterliki) 4 ?? Mastixicarpum sp. (as Sideroxylon putterliki) 5-6 extant fruits of Sideroxylon for comparison 7-9 ? Lauraceae vel Fagaceae (as Sideroxylon hepios) 10 Cwpolithes sp. (as Sideroxylon hepios) 11 extant fruits of Sideroxylon for comparison 12-13 Dicotylophyllum sp. (as Chlysophyllum atticum) 14-15 Carpolithes sp. (as Chrysophyllum atticum) 16-28 Dicotylophyllum sp. - partly Leguminosae ? (as Chrysophyllum olympicum) 29 Leguminosae (as Bumelia kymeana) 30 ? Leguminosae (as Bumelia oreadum) 31 -34 Dicotylophyllum sp. (as Bumelia minor) 35-36 Laurophyllum sp. (as Myrsine selenes) 37 ? Laurophyllum sp. (as Myrsine grandis) 38 (as 39) Dicotylophyllum sp. (as Myrsine graeca) 39 Myrica lignitum (as Euclea relicta) 40-50? Gordonia sp. vel "Diospyros" rugosa (as Royena graeca) 51 Dicotylophyllum sp. (as Royena graeca) 52 extant Royena for comparison PI. 12 1-23 Dicotylophyllum sp. (as Sapindus graecus) 24 Quercus sp., cupule s.n. extant Nephelium for comparison 25-27 ? Trigonobalanopsis rhamnoides (as Sapindus graecus) 28-29 Acer tricuspidatum (as Acer trilobatum) 30 Acer sp. (as Acer trilobatum) PI. 13 1-6 ? Trigonobalanopsis rhamnoides (as Rhamnus brevi/olius) 7-9 Quercus mediterranea (as Celastrus persei) 10-11 ? Berberis (as Celastrus oxyphyllos) 12-13 Dicotylophyllum (as Celastrus graecus) 14 "!lex" cyclophylla 15-18 "!lex" cyclophylla (as !lex neogena) 19-25 ? Rosa sp. (as !lex ambigua) 26 Alnus gaudinii (as Prinos eubaeos) 27 Dicotylophyllum sp. (as Pittosporum ligustrinum) 28-30 ? Laurophyllum sp. (as Rhus zanthoxyloides) PI. 14 1 Dicotylophyllum sp. (as Royena amalthese) 2-4 Dicotylophyllum sp. (as Royena euboea) 5-6 Dicotylophyllum sp. (as Royena myosotis) 7-8 Antholites sp. (as Royena myosotis) 9 Dicotylophyllum sp. (as Royena penteliei) 10 Fagaceae - Lauraceae (as Andromeda protogaea) 11-12 Cedrela attica (as luglans attica) 13 Dicotylophyllum sp. (as Carya bilinica) 14-15 ? Trigonobalanopsis rhamnoides (as Rhus heladotherii) 16 ? Leguminosae (as Rhus antilopum) 17-19 ? Rosa sp. (as Amyris berenices) 20 ? Leguminosae fruit (as Amyris berenices) • 21 ? Populus populina (as Amyris canopi) 22-25 Leguminosae leaflets (as Omphalobium relictum) 26 extant Omphalobium for comparison 27-29 ? Platanus neptuni (as Prunus aegaea) 30-33 Carpolithes sp. (P,unus aegaea) PI. 15 1 Dicotylophyllm sp. (as Eucalyptus agaea) 2-4 Dicotylophyllm sp. (as Callistemon eocenicum) 5 Dicotylophyllm sp. (as Myrtus paradisiaca) 6 Dicotylophyllm sp. (as Rhynchosia populina) 7 extant Rhynchosia for comparison 8 Dicotylophyllm sp. (as Rhynchosia ammonia) 9 extant Rhynchosia for comparison 10 ? Leguminosae leaflet (as Rhynchosia osiridis) 11 extant Rhynchosia for comparison 12-15 ? Leguminosae leaflets (as Rhynchosia isidis) 16 extant Rhynchosia for comparison 17-20 Leguminosae leaflets (as Glycine glycyside) 21 ? Leguminosae leaflets (as Caesalpinia antiqua) 22 extant Caesalpinia for comparison 23-25 ? Leguminosae leaflets (as Caesalpinia europaea) 26-27 extant Caesalpinia for comparison 28-30? Leguminosae leaflets (as Cassia aegaea) 31 extant Cassia for comparison 32-33 ? Trigonobalanopsis rhamnoides (as Cassia memnonia) 34 Leguminosae leaflet (as Cassia vetula) 35 extant Cassia for comparison 36 Daphnogene polymorpha (as Bauhinia olympica) 37 ? Leguminocarpon sp. (as Copaijera kymmeana) 38 ? Trigonobalanopsis rhamnoides (as Copaijera kymmeana) 39-41 Leguminosae leaflets (as Copaijera kymmeana) PI. 16 1-3 Leguminosae leaflets (as Prosopsis kymmeana) 4 extant Prosopsis for comparison 5? Leguminosae leaflet (as Acacia prisca ) 6-7 Leguminosae leaflets (as Mimosa mediaeae) 8-9 Leguminocarpon sp. (as Mimosa mediaeae) 10 Lauraceae - Fagaceae (as Inga icari) 11 extant Inga for comparison 12 reconstruction of a seed cone of Pinus megalopsis 13 Pinus sp. needles (as Pinus megalopsis) 14 Dicotylophyllum sp. (as Grevillea pandorae) 15 cortex 16 ramus 17-20 insects PI. 17 1 Araliaceae (as Cussonia polydrys) 2 extant Cussonia for comparison References: Aronis, G. 1952: Aliveri. Geological and mining survey of the lignite basin. Inst. Geol. Subs. Res. Ath. 2: 97-140. Aronis, G. , Anastopoulos, J. 1950: Interim report on the lignite deposits of Kymi. IGSR, Athens. Bachmayer, F., Symeonidis, N., Theodorou, G. 1981: Neue Insektenreste aus den jungtertiaren Susswasser-Ablagerungen von Kumi (Insel Euboa, Griechenland). Ann. Geol. Pays Hellen. 30: 763-766. Benda, L., de Bruijn, H. , Gregor, H.-J. 1982: Biostratigraphic correlations in the Eastern Mediterranean Neogene. Newsl. Stratigr. 11: 128-135. Bornovas, J., Rondoyanni-Tsiambaou Th., 1983: Geological map of Greece, 1:500.000, Ed. IGME. Bruijn, H. de, Meulen, van der J., 1979: A review of the Neogene Rodent succession in Greece. Ann. Geol. Pays Hell. Hors ser., Fasc. I, p207-217, VIIth Int. Congo On Mediterranean Neogene, Athens. Bruijn, H. de, Meulen, van der, J., Katsikatos, G. 1980: The mammals from the Lower Miocene of Aliveri (Island of Evia, Greece). Part 1. The Scuridae. Proc. K. Nederl. Akad. Wet. B, 83: 231-261. Deprat, J. 1904: Etude geologique et petrographique de I' ile de Eubee. Becanson, Paris. De Saporta, G. 1868: Sur la flore fossile de Coumi (Eubee). Bull. Soc. Geol. Fr., 2 (25) : 315-328. Doukas, C.S. 1986: The mammals from the Lower Miocene of Aliveri (Island of Evia, Greece). Part 5. The Insectivores. Proc. K. Nederl. Akad. Wet. , B 89 (1): 15-36. Engelhardt, H . 1910: Prilog poznavanju fosilne flore iz Kumi. Glansik zem. Muz. Bosni Herceg. 1910: 671-683. Erdei, B., Kvacek, Z. 1997: A newly recovered collection of the Early Miocene flora of Kymi (Greece) previously misinterpreted as the Upper Miocene flora ofTxllya (NE Hungary). Ann. Hist.-nat. Mus. Nat. Hung. 89: 5-10. Fytikas M., Giulliani 0., Innocenti F, Manetti P., Mazzuolli R , Peccerillo A, Villari L., 1976: Geochronological data on recent magmatism of the Aegean Sea. Tectonophysics 31: 129-134. Fytikas M., Giulliani 0., Innocenti F, Manetti P., Mazzuolli R , Peccerillo A, Villari L., 1979: Neogene volcanism of the Northern and central Aegean Region . Ann. Geol. Pays Hell., 30: 106-129. Fritel, P.H. 1921: Revision de la flore aquitanienne de Coumi (Grece). Bull. Mus. Hist. Nat. 27 : 576-580. Fritel, P.H. 1922: Revision de la flore aquitanienne de Coumi (Grece). Bull. Mus. Hist. Nat. 28 : 123-128. Gaudry, A 1860: Plantes fossils de l'ile de l'Eubee. C. R Acad. Sci. Paris 50 : 1093-1095. Gaudry, A 1862: Animaux fossiles et geologie de l'Attique. - F Savy, Paris. Gorceix, H. 1874: Notice sur Ie basin miocenique d'eau douce de Koumi (Eubea). Ann. Ec. Norm. Sup. 2: 317-32l. Gregor, H.-l. 1983: A Miocene fruit- and seed flora from the browncoal of Aliveri (Island of Evia, Greece. Documenta Naturae 6: 1-20. Guernet, C. 1971: Contribution a I'etude geologique de l'Eubee et des regions voisines. Theses, Univ. Paris VI, Paris. Guerner, c., Sauvage, l . 1969 : Sur la microflore des lignites et calcaires marneux des bassins neogenes de Kymi et de Gides (Eubee, Grce). c.R. Acad. Sci. Paris 269: 1611-1613. loakim Chr., 1984: Palynological - Stratigraphical of D4 and D5 Boreholes Vitala-Kymi. Euboea Int. Reports. IGME. loakim Chr., 1988: Palynology and Stratigraphy of E5 borehole Ents-kymi, Euboea. Int. Report. IGME. Katsikatsos, G., de Brujin, H., van der Meullen, A. l. 1981: The Neogene of the Island of Euboea (Evia), a review. Geo1. Mijnb. 60: 509-516. Kvacek, Z., Erdei, B. 2001: Putative proteaceous elements of the Lomatites-type reinterpreted as new Berberis of the European Tertiary. Plant Syst. Evol. 226: 1-12. Kvacek, Z., Velitzelos, E. 2000: The cycadalean foliage "Encephalartos" gorceixianus Saporta (Zamiaceae) from the Lower Miocene of Greece (Kymi, Island of Evia) revisited. Feddes Repert. 111: 151-163. Kvacek, Z., Walther, H. 2001: The Oligocene of Central Europe and the development of forest vegettaion in space and time based on megafossils. Palaeontographica B, 259: 125-148. Mai, D.H. 1996: TerWire Vegetationsgeschichte Europas. - G. Fischer VerI. , lena. Mitzopoulos, M., 1947: Die Verbreitung der Pikermistufe auf der insel Euboea. Ann. Geo1. Pays Hellen. 1: 209-216. Papanikolaou, C. 2001: Atlas of Greek coal Petrographic characteristics.Biomarkers.2nd E.U.Framework Program.IGME.Athens,426p. Saporta, G. 1868: Note sur la flore de Coumi (Eubee). Bull. Soc. Geo1. France, ser. 2, 25 : 315-328. Saporta, G. 1873: Examan critique d'une collection de plants fossiles de Koumi (Eubee). Ann. Scient. Ecole norm. super. 2e ser. , 2 : 323-352. Saporta, G. 1874: Sur la presence d'une Cycadee dans Ie depot miocene de Koumi (Eubee). c.R. Acad. Sci. Paris 78: 1318-132l. Symeonidis, N. K. 1975 : Ein bemerkenswerter Wirbeltierfund aus dem Lignit von Atalanti (Phyniotis, Griechenland). Ann. Geo1. Pays Hellen 26: 306-313. Unger, F. 1861: Neu-Holland in Europa. - W. Braunmiiller, Wien. Unger, F. 1862: Wissenshcaftliche Ergebnisse einer Reise in Griechenland und in den jonischen Inseln. - W. Braunmiiller, Wien. Unger, F. 1867: Die fossile Flora von Kumi auf der Insel Eubpoea. Denkschr. K. Akad. Wiss. , math.-naturwiss. Cl. 27: 27-87 . • Velitzelos, E., Buzek, a., Kvacek, Z . 1992: Contributions to the Lower Miocene flora of Aliveri (Island of Evia, Greece). Documenta Naturae, 74: 10-25. Velitzelos, E., Gregor, H.-J. 1982: Der erste Nachweis von Mastixiaceen im Tertiar von Euboea (Griechenland). Ann. Geol. Pays hellen. 31: 107-112. Velitzelos, E., Gregor, H.-J. 1990: Some aspects of the Neogene floral history in Greece. Rev. Palaeobot. Palyn. 62: 291-307. Velitzelos, E., Gregor, H.-J., Jahnichen, H. 1983: Fossile Vertreter der Rosskastanien (Aesculus LINNE) in der untermiozanen Flora von Kumi (Eubba, Griechenland). Documenta Naturae, 14: 1-23. Velitzelos, E., Kvacek, Z. 1996: New data of the Lower Miocene flora of Kymi (Island of Evia), Greece. Abstr. 5th lOP Conference, Santa Barbara, p. 105. Velitzelos, E., Kvacek, Z., Walther, H. 1999: Erster Nachweis von Eotrigonobalanus jurcinervis (Rossm.) Walther & Kvacek (Fagaceae) in Griechenland. Feddes Repert. 110: 349-358. Yiakkoupis P., VakalopoulosS P., Xenakis M. 1998: The Kymi lignite deposit ents lignite field western margins. IGME, 172p. Woodward, A. 1901: On the bonebeds of Pikermi, Attica and on similar deposits in Northern Euboea. Geol. Mag. 4: 481486 . • PLATES Pl. 1 1, la ? Cupressus sp. (as Callitris brongniartii) 2 indet. (as Callitris brongniartii) 3 - 11 Glyptostrobus europaeus • 1.2. Callitris Brongniarti End!. sp. 3-11. Glyptostrobus europaeus Brong. sp. Pl. 2 1, 9 Pinus holothana 2- 8, 10-11 Pinus sp. (as Pinus holothana) 12 extant Pinus for comparison 13 - 14 Pinus hampeana 15 Pinus sp. (as Pinus hampeana) 16 ? Pinus sp. (as Pinus furcata) 17-21 Cupressus sp. (as Sequoia langsdorfii) 22-23 ? Sequoia abietina vel Taxodium dubium (as Sequoia langsdorfii) 24-25 Dicotylophyllum sp. (as Podocarpus taxites) 1 / / . // / " ..- 25. 1-11. Pinus holothana Ung. 13-15. Pinus Hampeana Ung. 16. Pinus furcata Ung. 24-25. Podocarpus Taxites Ung. 17-23. Sequoia Langsdolji Er. sp. Pl. 3 1-4 Alnus keJersteinii (as Alnus sporadum) 5 reconstruction 6-7 Pinus sp. male cones (as Alnus sporadum) 8 Alnus gaudinii (as Alnus sporadum) 9-10 ? Alnus gracilis (as Alnus cycladum) 11- 22 Alnus cycladum 23 Carpinus grandis (as CaJpinus betuloides) 24-26 ? Alnus vel Carpinus (as Carpinus betuloides) 27 (17) ? Quercus drymeja (as Carpinus betuloides) 28 ? Zelko va zelkovaeJolia (as Carpinus betuloides) 29 ? Fagus sp. (as Carpinus betuloides) 30-34 Alnus gaudinii (as Carpinus betuloides) 35-37 catkins of ? Carpinus or Ostrya , .\ 33. 36. 1-8. Alnus Sparadum Ung. 9-22. Alnus Cycladum Ung. \ 35. 23-37. Carpinus belulaides Ung. Pl. 4 1-9 Alnus gaudinii (as Cmpinus betuloides) 10-16 Zelkova zelkovaefolia (as Planera ungeri) 17 Dicotylophyllum sp. (as Quercus cyclophylla) 18 ? Quercus dlymeja (as Quercus furcinervis) 19 ? Quercus drymeja (as Fagus pygmea) 20-22,28-30 Myrica vindobonensis 23-27 Myrica lignitum (as Myrica vindobonensis) 31-36 ? Laurophyllum pseudoprinceps (as Ficus aglajae) • 2. i / 31. i 135. 28. ! 1-9. Carpinus betuloides Ung. 18. Quersus furcinervis Roism. sp. 10-16. Planera Ungeri Ett. 17. Quersus cyclophyUa Ung. 19. Fagus pygmaea Ung. 20-30. Myrica vindobonensis Ett. sp 31-36. Ficus AgZajae Ung. Pl. 5 1-17 Quercus drymeja (as Quercus lonchitis) 18-20 Alnus gaudinii (as Quercus kamischinensis) 21-22 Quercus sp., cupules (as Quercus lonchitis) 1-17, 21, 22. Quersus Lonchiris Ung. 18-20. Quersus Kamischinensis Gopp sp. Pl. 6 1-22 Quercus mediterranea 23-27 Quercus drymeja (as Quercus zoroastri) 29 Populus populina (as Populus attenuata) 30 ? Populus stipule (as Populus attenuata) 31 Berberis kymeana (as Grevillea kymeana) 1-22. Quel"Sus mediterranea Ung. 23-28. Quersus Zoroastri Ung. 31. Grevillea Kymeana Ung. 29, 30. Populus attenuata A.Br. Pl. 7 1-10 Daphnogene polymorpha (as Cinnamomum lanceolatum) 11-24 Daphnogene polymorpha (as Cinnamomum scheuchzeri) 25-30 Daphnogene polymOlpha (as Cinnamomum subrotundum) 31 ? Cinnamomum fruits (as Cinnamomum rossmaessleri) 32 Ziziphus ziziphoides f. paradisiaca (as Cinnamomum rossmaessleri) 33-38 ? Lauraceae vel Fagaceae (as Laurus lalages) 39 ? Laurophyllum pseudoprinceps (as Cinnamomum buchii) • 1-10. Cinnamomum lanceolatum Ung. 5p. 11-24. Cinnamomum Scheuchxeri Heer. 25-30. Cinnamomum subrotundum Heer. 31-32. Cinnamomum Rofsmafsleri Heer. 33-38. Laurus Lalages Ung. 39. C. Buchi Heer. Pl. 8 1-7 ? Lauraceae vel Fagaceae (as Lawus primigenia) 8-10 ? Laurophyllum pseudoprinceps (as Lawus princeps ) 11 Myrica lignitum (as Laurinastrum dubium) 12 ? Laurophyllum sp. (as Protaea graeca) 13 ? Dicotylophyllum sp. (as Persoonia euboea) 14 Comptonia difformis f. dlyandroides (as Dlyandra thesei) 15 ? Laurophyllum sp. (as Grevillea kymeana) 16-31 Berberis kymeana (as Grevillea kymeana ) 32-33 ? Pinaceae seeds (as Hakea attica) 34-35 extant Proteaceae for comparison • 13.* I L_32. _______.-J 1-7. Laurus primigenia Ung. 8-10. Laurus princeps Heel'. 11. Laurinastrum dubium Ung. 12. Protaea graeca Ung. 13. Persoonia Eubraea Ung. 14. Dlyandra Thesei Ung. 15-31. Grevillea Kymeana Ung. 32.33. Hakea attica Ung. Pl. 9 1-3 Myrica solonis (as Banksia solonis) 4-13 , 15 Myrica lignitum (as Dryandroides hakeaefolia) 14 ? Quercus drymeja (as Dryandroides hakeaefolia) 16-18 Comptonia difformis f. dryandroides (as Dryandra ungeli) 19-22 Saportaspermum sp. (as Embothlium salicinum) 23 ? Cedrelospermum aquense (as Embothrium boreale) • 1-3. Banksia Solonis Ung. 4-15. Dlyandroides hakeaef olia Ung. 16-18. Dryandra Ungeri Ett. 19-22. E mbothrium Salicinum Heer. 23. Embothrium boreale Ung. Pl. 10 1-12 Myrica lignitum (as Olea noti) 13 -24 Myrica lignitum (as Asclepias podalyrii) 25 Myrica lignitum (as Neritinium longifolium) 26 ? Trigonobalanopsis rhamnoides (as Apocynophyllum carissa) • 1-12. Olea Noti Ung. 13-24. Asclepias Podalyrii Ung. 25. Neritinium longifolium Ung. 26. Apocynophyllum Carissa Ung. Pl. 11 1-3 Laurophyllum sp. (as Sideroxylon putterliki) 4 ?? Mastixicarpum sp. (as Sideroxylon putterliki) 5-6 extant fruits of Sideroxylon for comparison 7-9 ? Lauraceae vel Fagaceae (as Sideroxylon hepios) 10 Carpolithes sp. (as Sideroxylon hepios) 11 extant fruits of Sideroxylon for comparison 12-13 Dicotylophyllum sp. (as Chrysophyllum atticum) 14-15 Carpolithes sp. (as Chrysophyllum atticum) 16-28 Dicotylophyllum sp. - partly Leguminosae ? (as Chrysophyllum olympicum) 29 Leguminosae (as Bumelia kymeana) 30 ? Leguminosae (as Bumelia oreadum) 31-34 Dicotylophyllum sp. (as Bumelia minor) 35-36 Laurophyllum sp. (as Myrsine selenes) 37 ? Laurophyllum sp. (as Myrsine grandis) 38 (as 39) Dicotylophyllum sp. (as }\{yrsine graeca) 39 Myrica lignitum (as Euclea relicta) 40-50 ? Gordonia sp. vel "Diospyros" rugosa (as Royena graeca) 51 Dicotylophyllum sp. (as Royena graeca) 52 extant Royena for comparison • 1-4. Sideroxylon Putterliki Ung. 7-10. Sideroxylon hepios Ung. 12-15. Chlysophyllum atticum Ung. 16-28. Chrysophyllum olympicum Ung. 29. Bumelia Kymeana Ung. 30. Bumelia Oreadum Ung. 31-34. Bumelia minor Ung. 35-36. Myrsine Selenes Ung. 37. Myrsine grandis Ung. 38. Myrsine gr(Eca Ung. 39. Euclea relieta Ung. 40-51. Royena gr(Eca Ung. Pl. 12 1-23 Dicotylophyllum sp. (as Sapindus graecus) 24 Quercus sp., cupule s.n. extant Nephelium for comparison 25-27 ? Trigonobalanopsis rhamnoides (as Sapindus graecus) 28-29 Acer tricuspidatum (as Acer trilobatum) 30 Acer sp. (as Acer trilobatum) ./ - It --'---\/ ._-"- ｜ｾＭ ' '\' -- -I. .....- .. __ ｾ＠ 1-23. Sapindus graxus Ung. J 24-27. Nephelium Jovis Ung. 28-30_ Acer trilobatum A.Br. Pl. 13 1-6 ? Trigonobalanopsis rhamnoides (as Rhamnus brevifolius) 7-9 Quercus mediterranea (as Celastrus persei) 10-11 ? Berberis (as Celastrus oxyphyllos) 12-13 Dicotylophyllum (as Celastrus graecus) 14 "flex" cyclophylla 15-18 "flex" cyclophylla (as flex neogena) 19-25 ? Rosa sp. (as flex ambigua) 26 Alnus gaudinii (as Prinos eubaeos) 27 Dicotylophyllum sp. (as Pittosporum ligustrinum) 28-30 ? Laurophyllum sp. (as Rhus zanthoxyloides) • "-----, 6. ---'", : . ｾ｟ＭＮｊ / 'r ' " ./ • ｾ｜＠ /,\ / \ ｾｩ＠ -', 6. * / " '-...,.r, , ＬＭｾ＠ ''\ \1/ / i I \' 25. \1 t, o 1/ l , \ 25.*" .. , l' \ ' , ': ( l \'·t Ｇｾ＠ ) j r f I S 27. I ｾ＠ " , , \ 1 \0/JI ｾ＠ . ' , ,' 24. *\ \, • ｾ＠ 23 . 1-6. Rhamnus brevifolius Reer. 7-9. Celastrus Persei Ung. 10-11. Celastrus oxyphyllus Ung. 12-13. Celastrus graxus Ung. 14. flex cyclophylla Ung. 15-18. flex neogena Ung. 19-25. flex ambiqua Ung. 26. Prinos Eubceos Ung. 27. Pittosporum ligustrinum Ung. 28-30. Rhus zanthoxyloides Ung. Pl. 14 1 Dicotylophyllum sp. (as Royena amalthese) 2-4 Dicotylophyllum sp. (as Royena euboea) 5-6 Dicotylophyllum sp. (as Royena myosotis) 7 -8 Antholites sp. (as Royena myosotis) 9 Dicotylophyllum sp. (as Royena penteliei) 10 Fagaceae - Lauraceae (as Andromeda protogaea) 11-12 Cedrela attica (as Juglans attica) 13 Dicotylophyllum sp. (as Carya bilinica) 14-15 ? Trigonobalanopsis rhamnoides (as Rhus heladotherii) 16 ? Leguminosae (as Rhus antilopum) 17-19 ? Rosa sp. (as Amyris berenices) 20 ? Leguminosae fruit (as Amyris berenices) 21 ? Populus populina (as Amyris canopi) 22-25 Leguminosae leaflets (as Omphalobium relictum) 26 extant Omphalobium for comparison 27 -29 ? Platanus neptuni (as Prunus aega ea ) 30-33 Carpolithes sp. (Prunus aegaea) , - -3. \ I 1 ! ( I \12.* :I " 5. J ｾＮ｜＠ + I! l 6. 7 / / /' j 1. Royena Almathece Ung. 2-4. Royena EubO?ea Ung. 5-8. Royena myosotis Ung. 9. Royena Pentelici Ung. 10. Andromeda protogcea Ung. 11-12. Juglans attica Ung. 13. Carya biliniea Ung. 14-15. Rhus Helladotherii Ung. 16. Rhus Antilopum Ung. 17-20. Amyris Berenices Ung. 21. Amyris Canopi Ung. 22-25. Omphalobium relictum Ung. 27-33. Prunus O?gO?a Ung. Pl. 15 1 Dicotylophyllm sp. (as Eucalyptus agaea) 2-4 Dicotylophyllm sp. (as Callistemon eocenicum) 5 Dicotylophyllm sp. (as Myrtus paradisiaca) 6 Dicotylophyllm sp. (as Rhynchosia populina) 7 extant Rhynchosia for comparison 8 DicotylophyUm sp. (as Rhynchosia ammonia) 9 extant Rhynchosia for comparison 10 ? Leguminosae leaflet (as Rhynchosia osiridis) 11 extant Rhynchosia for comparison 12-15 ? Leguminosae leaflets (as Rhynchosia isidis) 16 extant Rhynchosia for comparison 17-20 Leguminosae leaflets (as Glycine glycyside) 21 ? Leguminosae leaflets (as Caesalpinia antiqua) 22 extant Caesalpinia for comparison 23-25 ? Leguminosae leaflets (as Caesalpinia europaea) 26-27 extant Caesalpinia for comparison 28-30 ? Leguminosae leaflets (as Cassia aegaea) 31 extant Cassia for comparison 32-33 ? Trigonobalanopsis rhamnoides (as Cassia memnonia) 34 Leguminosae leaflet (as Cassia vetula) 35 extant Cassia for comparison 36 Daphnogene polymorpha (as Bauhinia olympica) 37 ? Leguminocarpon sp. (as Copaijera kymmeana) 38 ? Trigonobalanopsis rhamnoides (as Copaijera kymmeana) 39-41 Leguminosae leaflets (as Copaijera kymmeana) • 7. -'\ .. \ 28. '. I I 'i 29. 30. 35. / 1. Eucalyptus agcea Ung. 2-4. Callistemon eocenicum Ett. 5. Myrtus paradisiana Ung. 6. Rhynchosia populina Ung. 8. Rhynchosia Ammonia Ung. 10. Rhynchosia Osiridis Ung. 12-15. Rhynchosia lsidis Ung. 17-20. Glycine glycyside Ung. 21. Ccesalpinia antiqua Ung. 23-25. Ccesalpinia eyropcea U. 28-30. Cafsia cegcea Ung. 32-33. Cafsia Memnonia Ung. 34. Cafsia vetula Ung. 36. Bauhinia olympica Ung. 37-41. Copaifera Kymeana Ung. Pl. 16 1-3 Leguminosae leaflets (as Prosopsis kymmeana) 4 extant Prosopsis for comparison 5? Leguminosae leaflet (as Acacia prisca) 6-7 Leguminosae leaflets (as Mimosa mediaeae) 8-9 Leguminocarpon sp. (as Mimosa mediaeae) 10 Lauraceae - Fagaceae (as Inga icari) 11 extant Inga for comparison 12 reconstruction of a seed cone of Pinus megalopsis 13 Pinus sp. needles (as Pinus megalopsis) 14 Dicotylophyllum sp. (as Grevillea pandorae) 15 cortex 16 ramus 17 -20 insects 2. 3. /\ I ,0 19. 13. , 19.* 18. 1-3. Prosopis Kymeana Ung. 5. Acacia prisca Ung. 6-8. Mimosa Medrrrr Ung. 10. Inga Jcari Ung. 12-13. Pinus megalopsis Ung. 14. Grevillea Pandorrr Ung. 15. Cortex 16. Ramus 17. Hydrophylus vexatorius Heer. 18. Helops Atticus L. R. 19. Calosoma excrobiculatum Heer. 20. Bombus pristinus Roghj Pl. 17 1 Araliaceae (as Cussonia polydrys) 2 extant Cussonia for comparison • ｾ f / --' -..., '-. J Ｍ \ \ ,-,t) \ ' · . [. / ll " "1.- ! ' :, ., \ Cussonia polydlys Ung.

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Field guide to the Neogene of the Island of Evia-Early Miocene flora of Kymi

Field guide to the Neogene of the Island of Evia-Early Miocene flora of Kymi

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