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Cedroxylon Lesbium (Unger) Kraus from the Petrified Forest of Lesbos, Lower Miocene of Greece and Its Possible Relationship to Cedrus

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Cedroxylon Lesbium (Unger) Kraus from the Petrified Forest of Lesbos, Lower Miocene of Greece and Its Possible Relationship to Cedrus N. Jb. Geol. Paläont. Abh. 284/1 (2017), 75–87 Article E Stuttgart, April 2017 Cedroxylon lesbium (UNGER) KRAUS from the Petrified Forest of Lesbos, lower Miocene of Greece and its possible relationship to Cedrus Dimitra Mantzouka, Vasileios Karakitsios, and Jakub Sakala With 6 figures Abstract: The Petrified Forest of Lesbos has been the subject of the palaeobotanical research since the 19th century, but a number of inconsistencies still remain. One of them concerns the fossils described over 100 years ago that are characterized by lack of the accompanied illustrations, missing or even lost type material, rather general and uninformative descriptions and finally weak evidence about stratigra- phy and exact location of fossiliferous sites. We present here an accurate interpretation of Cedroxylon lesbium (UNGER) KRAUS from Sigri (Petrified Forest area, western peninsula of Lesbos Island), which is hosted at the collections of the Natural History Museum of Vienna, Austria. The specimen, which is designated here as a lectotype, is compared with living Cedrus wood, its attribution to Cedroxylon is discussed and finally, a new combination for its denomination is proposed: Taxodioxylon lesbium (UNGER) MANTZOukA & SAKALA, comb. nov. Key words: fossil conifer wood, modern wood of Cedrus, Petrified Forest of Lesbos, early Miocene, Mediterranean, Greece. 1. Introduction presents those in the annual fossils’ exhibition at the Landesmuseum Joanneum (Graz, Austria) in 1842 Lesbos Island is highly appreciated by the scientific (GROSS 1999). The Austrian Professor of Botany and community because of the occurrence of the famous Director of this museum’s Botanical garden FRANZ Miocene Petrified Forest at the western peninsula UNGER described this material (UNGER 1844, 1845, 1847, of the Island (Fig. 1), although there are numerous 1850). One of the samples was identified by UNGER as important plant fossiliferous findings also from other Peuce lesbia UNGER. Unfortunately no stratigraphic parts of the island (e.g., MANTZOukA et al. 2016). The information, full anatomical description or illustrations first references about the extraordinary fossil flora of accompanied the identification. The present paper gives the island belong to the famous Greek philosopher for the first time a detailed description of the lectotype and father of botany THEOPHRASTUS (3rd century B.C.) of Peuce lesbia, known today as Cedroxylon lesbium (MANTZOukA et al. 2013, in press). In the first half of (UNGER) KRAUS, and also brings a comparison with the the 19th century AD the Archduke Johann collects modern Cedrus species for investigating its botanical fossil samples from the Petrified Forest of Lesbos and affinities. ©2017 E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, Germany www.schweizerbart.de DOI: 10.1127/njgpa/2017/0653 0077-7749/2017/0653 $ 3.25 eschweizerbart_xxx 76 Dimitra Mantzouka et al. Fig. 1. Lesbos Island palaeogeographical map during early Miocene. 2. Geology and stratigraphy of the study The central and western part of the Island is covered area by Neogene volcanic rocks (of calcium–alkaline and shoshonitic composition) originating from the volcanic Lesbos Island is located at the NE part of Aegean activity which took place at the Central-Northern and it belongs to the Pelagonian geotectonic zone of Aegean area (volcanic arc) and ended at the Western Greece which represents a fragment of the Cimmerian Anatolia during the upper Oligocene – middle Miocene Continent (MOUNTRAKIS 1986) with the bending of the (FYTIKAS et al. 1984). geotectonic zones to the E / NE to the Sakaraya Zone Concerning Cedroxylon findings from Lesbos of Asia Minor (THOMAIDOU 2009). Island, FLICHE (1898) described without giving further The geology of Lesbos has been described by details 4 types of Cedroxyla among his findings from several scientists (FYTIKAS et al. 1984; HECHT 1971- Ordymnos area (Cedroxylon type 1: samples No. 4, 31, 1974; KATSIKATSOS et al. 1982, 1986; MOUNTRAKIS 1986; Cedroxylon type 2: samples No 13-16, 23, Cedroxylon PE-PIPER 1980; PE-PIPER & PIPER 1993; THOMAIDOU type 3: samples No 3, 18, 24-30, Cedroxylon type 4: 2009), and it consists of: a) an autochthonous unit of sample No 8). Only one is permineralised (Cedroxylon Permo–Triassic age (basically extended in the south- type 4) among these fossils, the others are lignitic. eastern part of the Island) and b) two allochthonous Taking into account that the lignitic horizon is located units representing the volcanosedimentary nappe and in Lapsarna (Fig. 1) we come to the hypothesis that the ophiolite nappe. eschweizerbart_xxx Cedroxylon lesbium (UNGER) KRAUS from the Petrified Forest of Lesbos 77 FLICHE performed two different samplings. One with 4. Systematic part all lignitic samples was at the Lapsarna area, which some millions years ago (most probably in Ottnangian Family Cupressaceae GRAY sensu FARJON 2005 stage, MANTZOukA 2009; VASILEIADOU & ZOUROS 2012) Genus Taxodioxylon HARTIG hosted a palaeolake with a general low palaeoaltitude. The other one, hosting the permineralised tree trunks Taxodioxylon lesbium (UNGER) MANTZOukA & studied by FLICHE, was at the higher altitudes of SAKALA, comb. nov. Ordymnos Mountain. There is the Petrified Forest Park Fig. 2A-H (Bali Alonia). Although there is no stratigraphic evidence about Homotypic (nomenclatural) synonyms: the exact fossiliferous locality, the fossils UNGER (1844) 1844 Peuce lesbia UNGER, p. 37. had in his disposal come very possibly from Sigri area 1848 Pinites lesbius (UNGER). – GOEPPERT, p. 976. (or from Sigri until Ordymnos area, western peninsula 1870 Cedroxylon lesbium (UNGER). – KRAUS, p. 372. of Lesbos Island) which was easily accessible by boat. Material: One piece of wood (Fig. 2F) and four slides Moreover, this hypothesis is also supported by the among them three old and one new, sample No. D.38, reddish colour and the type of silicification (the fossil UNGER’s collection in the Natural History Museum of Vienna in hand is permineralised) of the original material that (lectotype, designated here). we re-studied. The reddish colour is connected with the Emended diagnosis: Homoxylous wood. Distinct growth reddish early Miocene stratigraphic horizon (enriched rings with abrupt transition from earlywood to latewood. in iron oxides) which hosts mostly permineralised Tracheid pitting opposite, uniseriate to predominantly stumps. According to PE-PIPER & PIPER (1993, 2002), biseriate. Crassulae present, helical and other wall Sigri pyroclastic deposits belong to the Skoutaros thickenings absent. Axial parenchyma diffuse to tangentially Formation and have been dated to 18.4 ± 0.5 Ma with zonate with smooth transverse end walls. Rays uniseriate or partly biseriate and very height (more than 30 cells). the use of K/Ar method. Ray tracheids absent. Ray walls, both horizontal and end (tangential), thin and smooth. Cross-field pits taxodioid in earlywood and cupressoid in latewood, mostly 1-3 per field 3. Material and methods arranged in one horizontal row (in marginal cells up to 4 per field arranged in two horizontal rows). Axial and radial The studied sections belong to the sample D.38 of UNGER’s intercellular (resin) canals absent. collection housed in the Natural History Museum of Vienna. Apart from the already existing sections, the Natural History Description: The slides come from a reddish permineralised Museum of Vienna made a new transverse one from the sample, about 10 centimeters in size. same type material UNGER had studied. The anatomical Transverse section: Nine distinct growth ring boundaries description is in accordance with IAWA Softwood List with a width from 0.4 to 1.2 mm (mean: 0.8 mm), transition (IAWA COmmITTEE 2004). For their microscopic observation from earlywood to latewood abrupt. The latewood is the facilities of the Faculty of Botany, Department of relatively wide and composed of 11-19 cells (mean: 14, Biology, National and Kapodistrian University of Athens SD: 2.4). False rings are absent. Latewood tracheids thin- (Olympus CX41 optical microscope, Nikon D5000, 12.3 walled, torus well-defined, disc-shaped (Fig. 2A, B), axial megapixel camera), Laboratory of Electron Microscopy of parenchyma mainly diffuse, sometimes tangentially zonate the Agricultural University of Athens (Scanning Electron (Fig. 2A, B), axial intercellular (resin) canals absent. Microscope Jeol 6360, Olympus BX40 Light microscope Tangential longitudinal section: axial parenchyma is equipped with accessories for fluorescence microscopy, present, transverse end walls smooth (Fig. 2H), the rays are dark field and phase contrast, DP71 Olympus 12.5 megapixel very high, with more than 30 cells (Fig. 2G, H) and up to digital camera and “Soft Imaging System-CELL” Olympus biseriate in width (Fig. 2G, H), radial intercellular (resin) image analysis software), and Institute of Geology and canals absent. Paleontology, Faculty of Science, Charles University in Radial longitudinal section: bordered pits in tracheids Prague (Olympus BX51 microscope, Olympus DP73 camera (Fig. 2C, E). The pits in radial walls of tracheids can be and QuickPHOTO MICRO 3.0 image analysis software) uniseriate but mainly tracheids have 2 rows of bordered pits were used. The modern wood specimens have been taken (tracheid pitting predominantly biseriate), arrangement of from “Julia & Alexander N. Diomedes” Botanic Garden of tracheid pitting in radial walls opposite (occasionally seems the University of Athens. The preparations of the modern alternate but it is due to the crowded pits),
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