PHYTOLOGIA BALCANICA 11 (2): 119–131, Sofia, 2005 119
Sequoioideae (Cupressaceae) woods from the upper Oligocene of European Turkey (Thrace)
Ünal Akkemik1, Nesibe Köse1, Imogen Poole2
1 Department of Forest Botany, Faculty of Forestry, Istanbul University, 34473 Bahceköy- Istanbul, Turkey, e-mail: [email protected] , [email protected] 2 Wood Anatomy Section, National Herbarium of the Netherlands, University of Utrecht Branch, P.O. Box 80102, 3585 CS Utrecht, The Netherlands, e-mail: [email protected] Received: May 06, 2005 ▷ Accepted: May 28, 2005
Abstract. Six Upper Oligocene mummified fossil wood samples originating from the Agacli Lignite Quarry near Istanbul (Turkey) were anatomically studied and identified as Sequoioxylon. This record increases the understanding of the palaeoenvironments of Turkey during the Oligocene and thus represents an important contribution to the poorly understood vegetation existing between Europe and Asia during the Tertiary.
Key words: fossil wood, Oligocene, palaeobotany, Sequoioideae, Sequoioxylon, Turkey
Introduction fossil material, including wood of Pinus sp., Taxus sp., Juniperus sp., Juglans sp., Quercus sp., and Salix sp. Identification of tree species based on wood anatomy (Aytug & Sanli 1974), alongside Sequoia type pollen is of interest not only to taxonomists studying extant (Bati 1996). The only record of taxodiaceous wood vegetation but also for studies focusing on past envi- was the material of Sequoioxylon (S. egemeni) de- ronments and ecosystems especially when other vege- scribed from Tertiary sediments within the Northern tative or reproductive parts are not available (Visscher Thrace Basin (Özgüven 1971) and the wood from a & Jagels 2003). Few palaeobotanical studies have been second lignite quarry near Çiftalan village (Eyüp dis- undertaken on the fossiliferous deposits of Turkey trict, Istanbul; Fig. 1) with the greatest similarity to and thus little is known about the Tertiary vegetation Sequoiadendron giganteum (Kayacik & al. 1995). and environments of this region. Those palaeobotani- Since our understanding of the vegetation that cal investigations that have been undertaken, on both grew in Turkey during the Tertiary is relatively poor, microfossils (Sanli 1982) and macrofossils (Özgüven the aim of this study was to identify the fossil wood 1971; Eroskay & Aytug 1982; Sanli 1982; Kayacik & al. samples in order to further the knowledge pertaining 1995; Aras & al. 2003), have concentrated on Tertiary to both the woody component of the vegetation and sediments around Anatolia (the Asian part of Turkey) thereby to the palaeoenvironment that existed across and Thrace (European Turkey) (Fig. 1). Within the Turkey during Oligocene. Northern Thrace Basin (Fig. 1) evidence from petri- fied woods for Carya sp. and Juglans sp. (Eroskay & Material and methods Aytug 1982) have been found alongside the pollen of such more exotic taxa as Ginkgo sp., Podocarpus sp., Pseudotsuga sp., Cedrus sp. and Pinus sp. (Sanli 1982). This work focuses on six pieces of Tertiary mummi- The Agacli Lignite Quarry has yielded ample Tertiary fied fossil wood (Plate I, Figs 1–6) originating from 120 Akkemik, Ü. & al. • Sequoioideae woods from the upper Oligocene
verse, tangential- and radial longitu- dinal). Anatomical analyses were per- formed with a light microscope and described using the terminology of the IAWA Committee (2004) for soft- wood identification wherever possi- ble. Mean and maximum numbers of cells were determined from 50 rep- licates for each character measure- ment. Due to compression, measuring the tracheid lengths was problemati- cal and only 20 (rather than 25) meas- urements per character were carried out for each specimen. Since accurate identification requires Fig. 1. Locality map indicating the location of Agacli Lignite Quarry and other places mentioned in the text. detailed descriptions and comparisons with well-described extant wood, pref- the Agacli Lignite Quarry (41° 17' 20'' N, 28° 44' 30'' E) erably from vouchered specimens (Visscher & Jagels situated near the village of Çiftalan in the Eyüp district 2003), the fossil woods were compared with pub- of Istanbul, Turkey (Fig. 1). The specimens were col- lished literature (e.g. Jacquiot 1955; Hejnowicz 1973; lected by Dr. Burhan Aytug in 1995 and are deposited Gromyko 1982; Kayacik & al. 1995; Visscher & Jagels in the Akkemik Collection housed in the University 2003) and sections of modern material housed in of Istanbul, Faculty of Forestry and Department of the National Herbarium of the Netherlands (Utrecht Forest Botany. Branch) and the Royal Botanic Garden, Kew (UK). On the basis of extensive palynological studies it is Finally, the specimens were compared with simi- widely accepted that the age of the coal-bearing stra- lar fossils formerly assigned to the Taxodiaceae (e.g. ta within the sequence from which the wood materi- Greguss 1955; Özgüven 1971; Basinger 1981; Aras & al was found (depicted with the bold horizontal lines al. 2003). in Fig. 2) represents the Upper Oligocene Danismen The largest piece (Specimen 6) of the six wood to Çöpköy Formations (Nakoman 1968; Ediger & samples collected could be used for tree-ring meas- Bati 1988; Ediger & al. 1990; Elsik & al. 1990; Akyol urements. A tree-ring chronology was construct- & Akgün 1995). Of the six samples retrieved from the quarry, one sample (Specimen 6) had previously been studied and assigned to S. giganteum on the basis of tracheid and ray characters (Kayacik & al. 1995), but has been re- $PBM examined and included here for completeness. Aras & CFBSJOH al. (2003) later identified a second large stem (approx- TUSBUB imately 115 cm) from the same lignite quarry as S. gi- ganteum. Although Taxodiaceae have recently been syno- nymised into the Cupressaceae (Gadek & al. 2000; Farjon 2001), in this work the fossil material previous- ly assigned to the Taxodiaceae will be referred to as ‘taxodiaceous’ rather than ‘cupressaceous’ for clarity. Thin sections (20–40 µm) of the wood materi- Fig. 2. Simplifi ed al were cut with a Richert microtome and prepared stratigraphy of the Agacli Lignite by standard methods, which include the staining, Quarry aft er Edi- in Safranin T, of the three planes of section (trans- ger & Bati (1988). Phytol. Balcan. 11(2) • Sofia • 2005 121 ed with the help of CRONOL computer program (in 88–98 %)] uniseriate, occasionally [mean 5 % (range DPL) (Grissino-Mayer & al. 1996) based on the 416 2–12 %)] biseriate within the body of the rays. Ray tree rings of the fossil wood No 6. This chronology height ranges from 1 to 28 cells in specimens 1, 2, 3 was compared with the latest 1000-year period of the and 6 (Plate II, Fig. 4), and up to 41 and 61 in spec- 3220-year long Giant Sequoia Master Chronology imen 5 and specimen 4 (Plate IV, Fig. 3) respective- (Hughes & al. 1994). The frequency of extreme neg- ly. Ray cell walls in the tangential section measure ative years was investigated in both the current and 2.5–6.25 µm. Axial parenchyma cells generally have the fossil tree ring series. To do this the mean of the smooth end walls (Plate II, Figs 4–5), but sometimes below-average values of the chronology was calculat- and more rarely the horizontal end walls are slight- ed and the “lower mean” obtained. Those values fall- ly dentate. ing below the lower mean were considered to repre- Radial section: Tracheids range from 4.3 to 9 mm sent extremely negative years. in length. Tertiary or spiral thickenings are absent. Tracheid walls have abundant circular bordered pits in uniseriate rows and oppositely arranged pits in bi- Results seriate rows; some triseriate rows were observed in Specimens 4 and 5 (Plate IV, Fig. 2) located predom- The description of the wood is based on the anatomi- inantly in the earlywood tracheids. Pits are close- cal characteristics of all six pieces of large organ/trunk ly spaced, sometimes crowded (Plate II, Fig. 6 and wood, measuring 1.2–24.5 cm in diameter, and 11– Plate III, Fig. 1) and measure 16–23 µm in diameter. 97 cm in length (Table 1; Plate I, Figs 1–6). Degradation around the edges of some bordered pits Transverse section: Wood material is mummified, can be clearly seen in Specimen 5 (Plate IV, Fig. 5). with some areas having undergone significant decay. Crassulae are frequently observed between pits form- Growth rings in Specimen 6 number over 416 tree- ing bi- to triseriate rows but are less frequent between rings. Growth rings in the other specimens range uniseriate pits (Plate II, Fig. 6 and Plate III, Fig. 1). from 13–51 in number (Table 1). Tree rings measure The numbers of successive biseriate-bordered pits between 0.5 and 2 mm in width. The transition be- are in the range of 19–20 in Specimens 4 and 5, and tween earlywood and latewood is generally distinct – 17–42 in the remainder. Most rays appear to be ho- abrupt or semi-abrupt (Plate II, Fig. 1, Plate IV, Fig. 1). mogeneous although ray tracheids are present in Tracheids of both earlywood and latewood are more all specimens. Ray tracheids are most frequent in or less angular in cross section. Radial diameters of Specimens 4 and 5, where they are located in the up- the earlywood tracheids measure 55–90 µm (mean = per or lower borders of some rays (Plate III, Fig. 2). 71 µm), whereas tangential diameters measure 40– Ray cell walls of all specimens are generally smooth 67 µm (mean = 52 µm). In well-preserved regions, tra- and seldom dentate. Horizontal ray cell walls meas- cheid cell wall thickness ranges from 2.5–3.8 µm in ure 2.5–5 µm in thickness. Half-bordered pit pairs in the earlywood to 6.3–12 µm in the latewood. Resin ca- the cross-fields number 2–4(-7) (-8 in Specimen 2) nals are absent. Wood parenchyma is present and dif- (Plate III, Fig. 3). Cross-field pits measure 8–14 µm fuse, especially in the transition zone from earlywood in diameter and are arranged in one or two horizon- to latewood and within the latewood. Parenchyma tal rows in the cross-fields of the wider earlywood cells are filled with resin-like globules (Plate II, Fig. 3) tracheids (Plate IV, Fig. 4), and in one or two vertical and have smaller or equivalent radial and tangential rows in the narrower latewood tracheids. Pit borders dimensions when compared with the surrounding are broad, elliptical to round; apertures are general- tracheids. ly oval and commonly horizontally orientated. Pits Tangential longitudinal section: Tracheidal pitting are more abundant in the uppermost and lowermost is abundant, with circular bordered pits located on marginal ray cells, arranged in one or two horizon- tangential walls of some earlywood tracheids (Plate II, tal rows. Axial parenchyma is composed of 6–11 cells Fig. 2), the circular pits ranging from 14–23 µm in di- in Specimens 4 and 5, and up to 14 cells in the re- ameter. Occasionally, more or less solitary pits were maining specimens with the marginal cells tall (1.4– observed on some tangential walls of the latewood 1.94 mm relative to the cells (0.13–0.23 mm) located tracheids. Rays are predominantly [mean 95 % (range in the middle of the axial chain. 122 Akkemik, Ü. & al. • Sequoioideae woods from the upper Oligocene
All samples can be assigned to the Sequoioideae er hand, ray tracheids present in the fossil material of the Cupressaceae based on the following features: have only been observed in Sequoia, Sequoiadendron, (1) generally abrupt transition from earlywood to and Metasequoia, in particular M. glyptostroboides latewood; (2) absence of true resin canals; (3) pre- Cheng & Hu (Greguss 1955). The presence of these dominantly two rows of opposite pits on the radial features increases the similarity of these three genera walls of tracheids; (4) crassulae conspicuous; (5) rays with the fossil material. homogeneous, with some rays having ray tracheids The presence of occasional nodules (indentures) present; (6) cross-field pitting predominantly taxo- in the ray cell walls of the fossils (Table 1) suggests dioid; (7) wood parenchyma present, diffuse, resin- a slight but possible affinity to Glyptostrobus and ous; and (8) horizontal end walls of axial parenchy- Taxodium (Taxodioideae) and Metasequoia. Wood ma are either smooth or very seldom nodular. The anatomical features of Metasequoia and Glyptostrobus six fossil woods under consideration could be sep- are very similar and thus these genera have been arated further on the basis of (1) ray height, where difficult to separate. However, a recent study by Specimens 1,2,3 and 6 attain maximum heights of Visscher & Jagels (2003) used quantitative characters < 30 cells (mean 4–6 cells) when compared with the to successfully separate these two genera (Table 2). rays of Specimens 4 and 5, where rays can reach a According to Visscher & Jagels (2003), Metasequoia maximum of c. 60 cells (mean 9–10 cells); (2) ray is characterized by the occasional separation of ray number per mm, where Specimens 1,2,3, and 6 have cells – a feature also observed in the fossil materi- ray abundances of > 40, whereas those of Specimens al. The abrupt transition from earlywood to late- 4 and 5 have abundances of < 40; and (3) occasional wood and the generally smooth horizontal end walls triseriate bordered pits can be observed in the trac- of the wood parenchyma cells are additional charac- heids of Specimens 4 and 5. However, the differenc- ters shared by the fossils and Metasequoia. However, es exhibited with regard to ray height and abundance anatomical differences between Metasequoia and the could be due to different relative ontogenetical ag- fossils include the arrangement of the cross-field pits. es of the specimens under study, whereas increase Those of Metasequoia are arranged in single hori- in the number of rows of tracheid pits in two of the zontal rows within the body of the ray cells but are specimens could be related to the organ of origin (cf. random in marginal ray cells, whereas in the fossils Falcon-Lang 2005). Therefore, although slight differ- cross-field pits are arranged in one or two horizon- ences amongst the specimens have been noted, they tal rows. Although the quantitative differences ex- do not warrant further subdivision. hibited by the cross-field pits could be a distinguish- ing taxonomic character, it cannot be ruled out that there might have been an evolutionary trend towards Discussion a reduction in the number of cross-field pits. Although ray height varies greatly both within a Comparisons with extant and fossil taxa species and throughout a single tree, by itself this fea- Identification of taxodiaceous wood to generic level is ture is considered an unreliable character for separat- often very difficult. The main wood features of modern ing species (Basinger 1981; Visscher & Jagels 2003; taxodiaceous genera are summarised in Table 2. These Falcon-Lang 2005). However, maximum ray height genera may be divided into two groups on the basis may be useful for generic determination if extremes of nodular (Glyptostrobus and Taxodium) or smooth are reached (Basinger 1981). The fossils exhibited a (Sequoia, Sequoiadendron, Metasequoia, Cryptomeria, range in ray heights with those of Specimens 1,2,3, Athrotaxis, Taiwania, and Cunninghamia) horizontal and 6 of greatest similarity to the heights attained walls of the wood parenchyma cells (Greguss 1955; by Sequoiadendron, whereas Specimens 4 and 5 have Basinger 1981). However, the findings of Jacquiot heights more similar to those exhibited by Metasequoia (1955) and the studies of modern taxodiaceous woods and Sequoia respectively (Tables 1, 2). However, we undertaken during the course of this study show that consider separation at generic level difficult using this the horizontal end walls of the parenchyma can be ei- feature alone. ther smooth or slightly nodular in modern Sequoia, Tracheid length is thought to distinguish Sequoia Sequoiadendron and Metasequoia wood. On the oth- sempervirens, Sequoiadendron giganteum and Meta- Phytol. Balcan. 11(2) • Sofia • 2005 123 sequoia glyptostroboides, in particular from other con- scribed. One sample formerly assigned to S. gigan- ifers (Basinger 1981 citing Panshin & DeZeeuw 1964). teum, namely Specimen 6, originally described by When compared with the fossil material, no clear dis- Kaya cik & al. (1995) was re-studied herein and de- tinction could be made between tracheid length of scribed as Sequoioxylon. One another big trunk iden- the fossils and the wood from modern Sequoioideae tified by Aras & al. (2003) as Sequoiadendron gigan- (Table 1). Sometimes striations similar to spiral thick- teum, which exhibits very similar wood anatomy to enings could be seen on the radial walls of tracheids the material identified as taxodiaceous herein, should in the fossils but these could represent spiral cavities also be considered as Sequoioxylon (Table 1). In the caused by soft rot following the microfibrillar struc- light of the above discussions and age of the mate- ture of the cell wall (Blanchette & Simpson 1992), rial, we consider that our specimens and the former rather than true spiral thickening and are, therefore, specimens from Ağaçli Lignite Quarry should also be not considered further. placed in Sequoioxylon sp. Although slight differences On the basis of the above findings, along with the amongst the specimens have been noted, they do not presence of ray tracheids and generally smooth ray cell warrant further subdivision. walls, we have considered that greatest affinity lies with the genera Sequoia, Sequoiadendron, and Metasequoia Palaeoenvironment of the subfamily Sequoioideae (Leurss.) Quinn (Gadek & al. 2000; Farjon 2001). However, anatomical differ- The Thrace region of Turkey lay at a palaeolatitude of ences between these fossil woods and those of mod- c. 37° N (Meulenkamp & Sissingh 2003) during the ern Sequoioideae do exist. Such differences include mid-Tertiary, with members of the Sequoioideae form- (i) a greater number of successive biseriate-bordered ing an important component of the vegetation. pits, often up to 42, in the fossil woods; (ii) the great- Taxodiaceous fossils from Late Cretaceous to er diameters of the bordered tracheid pits in the fos- Middle Tertiary sediments often dominate the floris- sil specimens relative to modern Sequoioideae; (iii) the tic components of lowland swamp and braided riv- number of cross-field pits in the fossil material being er delta forests of North America, Europe and Asia generally 2–4, seldom up to seven, (although eight oc- (Momohara 1994; Kumagai & al. 1995; Stockey & cur in Specimen 2), whereas in modern material there al. 2001). The Sequoioideae are well represented in are usually no more than six; and (iv) cross-fields of the fossil record from the Cretaceous onwards, with earlywood tracheids in the fossil material are often a common and widespread occurrence across two higher in number relative to modern Sequoioideae or more continents. Today their distribution has wood. to be interpreted as relict (Farjon 2001). Sequoia, Although we have suggested that greatest affini- once more widely distributed across the Northern ty of the fossils lies with the extant members of the Hemisphere until the Pleistocene, is now restrict- Sequoioideae, based on the combination of wood an- ed to California and Oregon in the United States of atomical features outlined above and summarised America (Farjon & Page 1999; Farjon 2001) where it in Tables 1 and 2, certain identification of the fossils is confined to elevations generally below 300 m (oc- described herein remains problematical. Therefore, casionally up to 1000 m) and within 60 km of the we have assigned the material to the morphogenus coast: a region characterised by fog (Watson 1993). Sequoioxylon erected for fossil woods with similari- Sequoiadendron occurs in mixed montane coniferous ties to the former Taxodiaceae or Sequoioideae of the forests between 900–2700 m in California and in iso- Cupressaceae. lated groves along the western foothills of the Sierra Sequoioideae wood of the Neogen age from Thrace Nevada (Watson 1993; Farjon 2001). Metasequoia has also been identified. Özgüven (1971) described ma- was widely distributed in the Tertiary, reaching 80°N terial which he assumed as belonging to Sequoioxylon during the Eocene, but became almost extinct in the egemeni on the basis of tracheid diameters and the Pleistocene (Farjon 2001). Today it is restricted to a presence of taxodioid type cross-field pits, homoge- small area in Central China near the Sichuan-Hubei neous ray cells occasionally with ray tracheids, wood border (ca. 30° 10' N, 108° 45' E) in E Sichuan, SW parenchyma cells and crassulae. Moreover, wood con- Hubei and NW Hunan, at altitudes between 750– sidered to belong to Sequoiadendron has also been de- 1500 m (Silba 1986). 124 Akkemik, Ü. & al. • Sequoioideae woods from the upper Oligocene
The occurrence of Sequoioideae woods (Özgüven widespread in Southern Thrace to the north, with the 1971; Aras & al. 2003) and Sequoia pollen (Ediger & al. exception of several small stands of P. nig ra represent- 1990) from the same horizon of the Upper Oligocene ing the relict Miocene flora, conifers characteristic of deposits, which represent former swamp-communities, the Tertiary no longer dominate the vegetation of that could be used to support the hypothesis of warm, humid area (Yaltirik 1966). conditions existing in Turkey at that time (Nakoman From growth ring analysis the numbers of ex- 1968). The occurrence of Taxodioxylon wood of the Late treme negative years, below the lower mean (Fig. 3 and Oligocene – Miocene age on the Lesvos Island (Greece) Table 3) in each 100-year period ranged between 13– (Velitzelos & Zouros 1997) also supports this hypothe- 22 in the present chronology and 17–20 in the fossil sis. This is also in agreement with other studies which wood. On this basis we can conclude that climate in the indicate that the mean annual temperature in Turkey in Thrace region during the upper Oligocene, when the the Early Oligocene was c. 15 °C and had subsequent- woods were living, is similar to the prevailing climate ly dropped to c. 10 °C by the beginning of the Miocene where the Sequoiadeae are distributed today. (Wolfe 1978; Aleksandrova & al. 1987). This cooling trend from the Early Oligocene through the Miocene In conclusion this paper outlines new records resulted in a change from subtropical to temperate-sub- of fossil woods with anatomy most similar to the tropical humid conditions (Ediger & al. 1990), which Sequoioideae (Cupressaceae) from Oligocene coal de- also reflected on the vegetation. posits of the Thrace Basin to the northwest of Istanbul. From the Upper Oligocene into the Miocene there These woods represent an important contribution was a change in the microflora represented in the to the poorly understood vegetation that existed be- Agacli Lignite Quarry. That change is characterized tween Europe and Asia during the Middle Tertiary. by taxodiaceous taxa being replaced by members of These fossils help confirm that taxodiaceous conifers the Pinaceae (Nakoman 1968) so that the new vege- formed an important part of the swamp community tation was distinguished by species of Pinus (such as under a prevailing warm temperate-subtropical cli- P. nig ra , P. pine a and P. brutia) that came to dom- mate regime, prior to the onset of the post Oligocene inate the forests across that region (Aytug & Sanli climatic deterioration. It is through the documenta- 1974). Today, vegetation of the Northern Thrace can tion of records such as this that the vegetation dynam- be separated into different zones: the humid broad- ics leading to the evolution of the present-day ecosys- leaved forest with Quercus petraea, Q. robur, Q. frai- tem in Europe can be more fully understood. netto, Fagus orientlis, Castanea sativa, Carpinus bet- ulus, Tilia argentea, Ulmus minor; the anthropogenic steppe; the macchia-pseudo-macchia; and the coastal Acknowledgements. We are indebted to Prof. Dr. Burhan Aytug zone (Dönmez 1968). Even though Pinus brutia is still for providing the material for study, Prof. Dr. Elizabeth Wheeler for her interest and for supplying the relevant literature, and Prof. Dr. Nüzhet Dalfes for his comments on an earlier draft of the manu- script. Furthermore, we thank Dr P. Rudall for allowing access to the wood collections housed in the Royal Botanic Gardens, Kew.
Fig. 3. Tree-ring chronology of specimen 6. Phytol. Balcan. 11(2) • Sofia • 2005 125 D S PXT S MP JBUF S
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Summary of the anatomical features of the six genera of the six genera of of features Summary the anatomical of UB 8U SN CV J U B IFJET QSFTFOU CSEFSFQ POUB POUB M J personal observation from cultivated trees in Istanbul; Ø – diameter; no. – number. Ø – diameter; no. in Istanbul; trees cultivated observation personal from FO I S 8 B TGJFMQ GP H QF SJ[ J O %JTU JOEFOUVSF SBZUS D $SP T SBEJ IPSJ[ IF OP &8- UB H P IP EW-LW – earlywood-latewood; max. – maxsimum; data from: from: data – earlywood-latewood; max. – maxsimum; EW-LW Table 2. Table 1952, Phytol. Balcan. 11(2) • Sofia • 2005 127
Plate I
DN DN
DN DN
DN
DN
Figs 1-6. Mummifi ed fossil wood specimens with the fi gure numbers coinciding with the specimen number. 128 Akkemik, Ü. & al. • Sequoioideae woods from the upper Oligocene
Plate II
Figs 1-6. Photomicrographs of fossil Sequoioideae wood specimens assigned to Group 1: 1, specimen 6 showing abrupt transition from earlywood to latewood; 2, specimen 6 showing bordered pits in earlywood tracheids and rays of two cells in tangential section; 3, specimen 2 showing wood parenchyma cells (arrowed) in transverse section with fi lled a prob- able resinous material; 4, specimen 6 showing smooth end wall of axial parenchyma cells (arrowed) and uniseriate rays; 5, specimen 2 showing axial parenchyma cells with fi lled resinous material and smooth end walls; 6, specimen 6 showing crassulae between successive uniseriate bordered pits. Phytol. Balcan. 11(2) • Sofia • 2005 129
Plate III
Figs 1-3. Photomicrografs of radial longitudinal sections of fossil Sequoioideae woods: 1, crassulae between bordered pit paris of specimen 6; 2, ray tracheids (arrowed) in specimen 3; 3, ray cells in specimen 2 with up to eight taxodioid pits per cross fi eld.
Table 3. The frequency of extremely negative years in 100-year period both the fossil and the current master chronologies.
1FSJPET $ISPOPMPHZPG 1FSJPET $VSSFOUNBTUFS UIFGPTTJMXPPE DISPOPMPHZ
TU ZFBS TU ZFBS OE ZFBS OE ZFBS SE ZFBS SE ZFBS UI ZFBS UI ZFBS UI ZFBS UI ZFBS UI ZFBS UI ZFBS UI ZFBS UI ZFBS 130 Akkemik, Ü. & al. • Sequoioideae woods from the upper Oligocene
Plate IV
Figs 1-5. Photomicrographs of fossil Sequoioideae woods: 1, tangential section of specimen 4 showing abrupt transition from earlywood to latewood; 2, triseriatae bordered pits in radial longi- tudinal section (RLS) of specimen 4; 3, specimen 4 tangential longitudinal section showing uniseriate rays including part of the tallest ray (total 61 cells) seen in specimen; 4, RLS showing taxodioid cros-fi eld pits in the earlywood to the specimen 5 aligned in two rows in upper and lower (marginal) cells and in one row in the cells of the body of the ray; 5, RLS showing tracheidal pitting in specimen 5 with outer borders showing degradation. Phytol. Balcan. 11(2) • Sofia • 2005 131
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