Miocene Cupressinoxylon from Gökçeada (Imbros), Turkey with Protophytobia cambium mining and the study of ecological signals of wood anatomy

Premise The recognition of the Miocene Climate Optimum (MCO) in terrestrial palaeoenvironments of the Eastern Mediterranean is restricted to Lesbos and Lemnos Islands, Greece. This area is significant for its wood microfossils. A recently-discovered fossil wood assemblage from Gökçeada (Imbros) Island, Turkey, including tree species similar to the Greek findings, is thought to have an early Miocene age. Here, we revise the age of the latter plant fossiliferous locality, re-evaluate the area for the study of MCO for the terrestrial palaeoecosystems of the Eastern Mediterranean and the nomenclature errors referring to the occurrence of fossil wood. We present the plant–insect–environment interactions using detailed anatomical descriptions, of an extinct conifer and its extinct cambium miner feeding traces observed in its secondary xylem. Methods Three thin sections were prepared with standard palaeoxylotomical techniques from a small section of the silicified wood; the sections were observed under a light microscope. The anatomy of the conifer and its damage patterns were compared with those of extant and fossil Cupressaceae and Agromyzidae, respectively. Pivotal results The common anatomical features of the studied wood specimen and Hesperocyparis macrocarpa (Hartw.) Bartel and a shared characteristic (the number of the cross-field pits – a feature we consider of diagnostic value) with Xanthocyparis vietnamensis Farjon & T.H. Nguyên led to its assignment to the Hesperocyparis–Xanthocyparis–Callitropsis clade. The detailed study of the wound scars and anatomical abnormalities, the anatomical–environmental associations, and structural–functional reactions follow the identification of the wood’s anatomy sensu Carlquist providing decisive results. Conclusions Based on the distinctive characteristics presented, we identify our macrofossil as Cupressinoxylon matromnense Grambast, a stem or an extinct lineage of the Hesperocyparis–Xanthocyparis vietnamensis–Callitropsis nootkatensis clade with feeding traces of the fossil cambium miner of the genus Protophytobia Süss (Diptera: Agromyzidae), and anatomical damage and reaction tissue on adventitious shoots. The use of Protopinaceae and Pinoxylon F. H. Knowlton from the eastern Mediterranean are re–evaluated and corrections are provided. The age of the studied plant fossiliferous locality in Gökçeada is revised as middle Miocene, allowing the proposal of an eastern Mediterranean MCO hotspot, including Lesbos, Lemnos, and Gökçeada (Imbros) Islands

Έρευνα της παλαιοχλωρίδας του καινοφυτικού στην Ανατολική Μεσόγειο (νήσοι: Λέσβος, Λήμνος, Αλόννησος, Γαύδος)

The presence of plant (macro) fossils in Greece is known since ancient times and is of high significance by the Greek and International scientific community because of: their age, the species and plant organs diversity and the information about the palaeoenvironment and paleoclimate which can be excluded. The present work is dealing with: the research for new plant fossiliferous localities of Cenophytic age at the Eastern Mediterranean (Lesbos, Lemnos, Alonissos, Gavdos Islands), the study of the new findings under the perspective of palaeobotanical analysis (petrified wood anatomy with the usage of microscope, study of leaves imprints) and use of older (method of the Nearest Living Relative) and new methodologies (e.g., CLAMP, IPR) in order to export significant conclusions about the geological history of the Eastern Mediterranean and of Greece.Η ύπαρξη φυτικών (μακρο)απολιθωμάτων στον ελλαδικό χώρο είναι γνωστή από την αρχαιότητα και χαίρει ιδιαίτερης σπουδαιότητας από την ελληνική και παγκόσμια επιστημονική κοινότητα λόγω της ηλικίας, της ποικιλότητας ειδών και οργάνων και των πληροφοριών για το παλαιοπεριβάλλον και το παλαιοκλίμα. Μέσα από την παρούσα εργασία πραγματοποιείται: έρευνα νέων θέσεων εύρεσης φυτικών απολιθωμάτων Καινοφυτικής ηλικίας στην Ανατολική Μεσόγειο (νήσος Λέσβος, Λήμνος, Αλόννησος, Γαύδος), προσπάθεια προσέγγισης των νέων ευρημάτων από τη σκοπιά της παλαιοβοτανικής ανάλυσης με ό,τι αυτή περιλαμβάνει (ανατομία απολιθωμένου ξύλου με χρήση μικροσκοπίου, μελέτη αποτυπωμάτων φύλλων) αλλά και χρήση παλαιότερων (μέθοδος του Πλησιέστερου Σημερινού Συγγενή) και νέων μεθοδολογιών (π.χ. CLAMP, IPR) με στόχο την εξαγωγή σημαντικών συμπερασμάτων για τη γεωλογική ιστορία της Ανατολικής Μεσογείου και γενικότερα του ελλαδικού χώρου

Miocene Cupressinoxylon from Gokceada (Imbros), Turkey with Protophytobia cambium mining and the study of ecological signals of wood anatomy

Premise: The recognition of the Miocene Climate Optimum (MCO) in terrestrial palaeoenvironments of the Eastern Mediterranean is restricted to Lesbos and Lemnos Islands, Greece. This area is significant for its wood microfossils. A recently -discovered fossil wood assemblage from Gokceada (Imbros) Island, Turkey, including tree species similar to the Greek findings, is thought to have an early Miocene age. Here, we revise the age of the latter plant fossiliferous locality, re-evaluate the area for the study of MCO for the terrestrial palaeoecosystems of the Eastern Mediterranean and the nomenclature errors referring to the occurrence of fossil wood. We present the plant-insect-environment interactions using detailed anatomical descriptions, of an extinct conifer and its extinct cambium miner feeding traces observed in its secondary xylem.Methods: Three thin sections were prepared with standard palaeoxylotomical techniques from a small section of the silicified wood; the sections were observed under a light microscope. The anatomy of the conifer and its damage patterns were compared with those of extant and fossil Cupressaceae and Agromyzidae, respectively.Pivotal results: The common anatomical features of the studied wood specimen and Hesperocyparis macrocarpa (Hartw.) Bartel and a shared characteristic (the number of the cross -field pits - a feature we consider of diagnostic value) with Xanthocyparis vietnamensis Farjon & T.H. Nguyen led to its assignment to the Hesperocyparis-Xanthocyparis-Callitropsis clade. The detailed study of the wound scars and anatomical abnormalities, the anatomical-environmental associations, and structural-functional reactions follow the identification of the wood's anatomy sensu Carlquist providing decisive results.Conclusions: Based on the distinctive characteristics presented, we identify our macrofossil as Cupressinoxylon matromnense Grambast, a stem or an extinct lineage of the Hesperocyparis-Xanthocyparis vietnamensis-Callitropsis nootkatensis clade with feeding traces of the fossil cambium miner of the genus Protophytobia Suss (Diptera: Agromyzidae), and anatomical damage and reaction tissue on adventitious shoots. The use of Protopinaceae and Pinoxylon F. H. Knowlton from the eastern Mediterranean are re-evaluated and corrections are provided. The age of the studied plant fossiliferous locality in Gokceada is revised as middle Miocene, allowing the proposal of an eastern Mediterranean MCO hotspot, including Lesbos, Lemnos, and Gokceada (Imbros) Islands

The first plant megafossil in the Early Jurassic of Greece: Brachyphyllum (Coniferales) from the Lower Posidonia Beds (Toarcian) in the Ionian zone (NW Greece) and its palaeogeographic implications

General information on the geology of the Lower Jurassic in the Hellenides is presented, focused on the Chionistra section, where the first land plant macrofossil was recovered. This specimen is interpreted as a new record of Brachyphyllum elegans (SAPORTA) BARALE and compared with the type material from the Upper Jurassic of France and some other Mesozoic fossils of this kind from Eurasia. Its stomatal complexes with papillate subsidiary cells and the co-occurrence with abundant Classopolis pollen indicate an affinity to the Cheirolepidiaceae. The reconstructed environment usually assigned to the Brachyphyllum nepos type may suggest mangrove-like vegetation that thrived under tropical-subtropical climatic conditions. Moreover, the finding of Brachyphyllum elegans (SAPORTA) BARALE from the Lower Jurassic (Toarsian) of western Greece provides confirmation of the palaeogeographic evolution model of the Ionian zone during this period

The first report of an ‘evergreen Castanopsis type’ wood (Fagaceae) for the Late Miocene–Early Pliocene of Europe (Bulgaria, Blagoevgrad Graben)

In the present article, the establishment of the species Castanopsis: C. bulgarica Mantzouka, Ivanov, and Bozukov is proposed after the study of a new fagaceous fossil wood stem discovered in 2016 from a new Late Miocene to Early Pliocene plant fossiliferous locality east of Boboshevo town, south-west Bulgaria. A detailed palaeoxylotomical study of the fossil wood revealed characteristics of the ‘evergreen Quercus type’ (e.g. the gradual porosity of the vessels, the existence of two types of rays: uniseriate and multiseriate aggregate, the oval shape of the solitary vessels outline, the occurrence of tracheids) as well as heterocellular compound-aggregate rays, typical for Castanopsis. Similarities and differences of the anatomical characteristics/features of the studied specimen with the descriptions of the fossil representatives of Fagaceae (Quercoxylon, Lithocarpoxylon, and Castanopsis) along with its botanical affinities are discussed. A taxonomic list of the Bulgarian fossil fagaceous record of the same age and their nearest living relatives is provided. Emended xylotomical keys with the addition of Castanopsis have been created. Moreover, the presence of evergreen Castanopsis species with heterocellular rays of two distinct sizes in the fossil record of the ‘evergreen oak woods’ is supported

The first report of Lesbosoxylon Suss & Velitzelos from the early-middle Miocene of eastern Anatolia

A new fossil pine species from eastern Turkey is described and its botanical affinities are discussed. The sample was collected from the city of Kemaliye, Erzincan province, Turkey, and derives from the early-middle Miocene Divrigi formation. Transverse, tangential and radial sections were taken from the petrified wood, and its palaeoxylotomical features were investigated. Based on its anatomical features including idioblastic cells in rays a new fossil-species of the genus Lesbosoxylon Suss & Velitzelos was identified as Lesbosoxylon kemaliyensis Akkemik & Mantzouka, sp. nov. Diagnostic features of the new species are: Transition from earlywood to latewood mostly gradual; axial and radial resin canals with thin-walled epithelial cells present; latewood tracheids thin to thick walled; bordered pits on radial walls of tracheids 1-2(-3) seriate; crassulae common; rays heterocellular, uniseriate, partly biseriate; uniseriate rays up to 27 cells high; fusiform rays up to 30 cells high; axial parenchyma occasionally present; ray tracheids 2-3 rows; cell walls of ray tracheids smooth; cross-field pitting pinoid, 1-2(-6) pits per cross-field. Detailed investigation of the botanical affinities of the new fossil wood suggested that the most closely related modern species is Pinus canariensis C. Sm in Buch, a relict species from the Canary Islands

The first paleoxylotomical evidence from the Mid-Eocene Climate Optimum from Turkey

Petrified wood is important evidence of forest type and past climate. Studies from mainly the Neogene of Turkey described many fossil woods, and revealed valuable information about the paleoenvironment structure and climate. This is the first occurrence of plant macrofossils belonging to the Middle Miocene Climate Optimum, a crucial period in Earth's history, found in Turkey. The purpose of the present study is to investigate, for the first time, the Middle Eocene Climate Optimum of Turkey through the first identification of two paleoxylotomical findings of Lutetian age from Central-North Turkey. After working on the thin wood sections, a new fossil genus of Lauraceae, Actinodaphnoxylon gen. nov. was described with a type species of Actinodaphnoxylon zileensis sp. nov. Another coniferous specimen was described as Pinuxylon cf. P. tarnocziense. These species indicate the presence of warm, humid rainforest and lower mountain forest 40-41 million years ago in the middle Black Sea region (Tokat-Zile) of North-Central Turkey. (C) 2020 Elsevier B.V. All rights reserved

Glyptostrobus europaeus Unger. Cette

<p> <i>Glyptostrobus europaeus</i> (Brongniart) Unger (Fig. 4 A-F)</p> <p> <i>Taxodium europaeum</i> Brongniart, 1833: 168 (Iliodroma = Alonissos).</p> <p> <i>Taxodites europaeus</i> – Endlicher 1847: 278 (Iliodroma = Alonissos).</p> <p> <i>Glyptostrobus europaeus</i> – Unger 1850: 434 (Iliodroma and others). — Schneider & Velitzelos 1973: 246, pl. 35, fig. 1 (Vegora). — Velitzelos & Schneider 1977: 173, figs 1-3 (Vegora). — Mai & Velitzelos 1997: 510, pl. 4, figs 1-4 (Vegora).</p> <p> ORIGINAL DIAGNOSIS. — “ <i>Ramis fastigiatis</i>, <i>elongatis</i>, <i>gracilibus: Foliis subtristichis</i>, <i>alternis</i>, <i>brevissimis</i>, <i>basi decurrentibus</i>, <i>apice acutiusculis: Strabilis subglobosis vel ovoideis: Squamis sub octofariis</i>, <i>disco terminali</i>, <i>margine superne armato</i>, <i>crenato</i>, <i>medio crista transversali prominente partito</i>, <i>saliis radiantibus in parte superiore notato</i> ” (Brongniart 1833: 168-176).</p> <p>DESCRIPTION</p> <p> According to the new study of the holotype the following characteristics have been observed: Branched twigs with cupressoid foliage (Fig. 4A). No taxodioid foliage was observed. Ripe cone (male) 1.6 cm long and it consists of about 16 elongated imbricated scales (Fig. 4B). Branches/ shoots with leaves scale-like, cupressoid type (or even linearsubulate as discussed in Ma <i>et al.</i> 2013) or if one follows the terminology by Vickulin <i>et al.</i> (2003) the leaves are squamate with cupressoid appearance, helically/spirally arranged or even slightly decussate. The leaves are short (leaf length: 1-2 cm and width: 0.5-1 cm) and their apex is not mucronate as in the modern <i>G. pensilis</i> but globose to broadly ovoid/concave (Fig. 4D, F). Cones (Fig. 4C, D) and perennial branchlets/shoots with cupressoid foliage (Fig. 4E) are observed.</p> <p>DISCUSSION</p> <p> According to Kvaček <i>et al.</i> (2002), <i>Glyptostrobus</i> in Euboea develops relatively smaller cones and heteromorphic foli- age, which is in contrast to Early Miocene specimens from Central Europe.</p> <p> <i>Glyptostrobus europaeus</i> represents a species found in several localities in Greece during the Cainozoic. Apart from the findings of this species from Kymi (Euboea Island) and Alonissos Island of Burdigalian age (Early Miocene) and from Vegora (Macedonia) of Messinian age (Upper Miocene) given in LePage (2007: 393), the geographical distribution of this species in Greece includes additional localities (Velitzelos <i>et al.</i> 2014), such as Aliveri and Nea Styra (Euboea Island) and Moudros (Lemnos Island) of Burdigalian age, Komnina (Macedonia), Platana (Western Peloponnese), Prosilio and Lava (Kozani, eastern West Macedonia), Likoudi and Drymos (Elassona, Northern Thessaly) and Iliokomi-Kormitsa (Strymon Basin- Thrace) of Upper Miocene (Messinian) age, Crete (Makrilia) of Middle Tortonian age, Pikermi-Chomateri (Attica) of Tortonian-Messinian age, Ptolemaida (West Macedonia) of Lower Pliocene age, Skoura (Sparti, Laconia) of Pliocene age, Kolymbia (Rhodes) of Pleistocene age.</p> <p> According to Manchester (1999), <i>Glyptostrobus</i> has a continuous presentation at the fossil floras of Europe from the Palaeocene until the Pliocene/Pleistocene, of North America during Palaeocene, Eocene and Miocene and a constant one from Palaeocene to present in Asia.</p> <p> <i>Glyptostrobus europaeus</i> taxonomic and biogeographic history started in Canada (Aptian) and is given in detail in LePage (2007). Unfortunately, the assignment of the fossil conifer described in Li <i>et al.</i> (2018) to <i>Glyptostrobus europaeus</i> is considered ambiguous because the sample has seed cones falling into scales, a characteristic not in accordance with any known fossil representative of <i>Glyptostrobus europaeus</i> until now. Recent studies (Dolezych & van der Burgh 2004) had correlated for the first time <i>Glyptostrobus europaeus</i> with <i>Glyptostroboxylon rudolphii</i> Dolezych & van der Burgh. Numerous information for the “whole-plant” reconstruction of the species providing by the connection between fossil stumps, trunks and twigs of the taxon are also given in Vassio <i>et al.</i> (2008) or by an indirect association of different organs within the same site (Teodoridis & Sakala 2008).</p> <p> <i>Glyptostrobus pensilis</i> Koch is the only extant species in its genus inhabited today in SE China, Viet Nam and Lao PDR (Henry & McIntyre 1926; Farjon & Filer 2013).</p> <p>Among the newly collected specimens we were able to identify a xylitic remnant with anatomical characteristics resembling a conifer. Its microscopical study is also presented below.</p>Published as part of <i>Mantzouka, Dimitra, Sakala, Jakub, Kvaček, Zlatko, Koskeridou, Efterpi & Ioakim, Chryssanthi, 2019, Two fossil conifer species from the Neogene of Alonissos Island (Iliodroma, Greece), pp. 125-142 in Geodiversitas 41 (3)</i> on pages 130-132, DOI: 10.5252/geodiversitas2019v41a3, <a href="http://zenodo.org/record/3692007">http://zenodo.org/record/3692007</a&gt

Two fossil conifer species from the Neogene of Alonissos Island (Iliodroma, Greece)

Mantzouka, Dimitra, Sakala, Jakub, Kvaček, Zlatko, Koskeridou, Efterpi, Ioakim, Chryssanthi (2019): Two fossil conifer species from the Neogene of Alonissos Island (Iliodroma, Greece). Geodiversitas 41 (3): 125-142, DOI: 10.5252/geodiversitas2019v41a