- Email: [email protected]
Leaves and pollen of bamboos from the Polish Neogene
Review of Palaeobotany and Palynology 133 (2005) 39 – 50 www.elsevier.com/locate/revpalbo
Leaves and pollen of bamboos from the Polish Neogene Elybieta Worobiec, Grzegorz Worobiec* WlCadyslCaw Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, PL-31-512 Krako´w, Poland Received 9 January 2004; accepted 26 August 2004
Abstract Recently discovered fossil bamboo leaves and pollen are described from the Neogene of Poland. Morphological analysis of fossil leaves from the Late Miocene deposits of the Beychato´w Lignite Mine, Central Poland have allowed them to be determined as bBambusaQ lugdunensis Saporta. It is the first record of this species in the Cenozoic of Poland. The fossils have been compared with other records of bamboos from the Cenozoic of Europe. Pollen grains from the Middle Miocene (Badenian) deposits from the Legnica brown coal deposit (Lower Silesia) provide additional evidence of bamboos. These are assigned to Graminidites bambusoides Stuchlik, and are similar to those of recent Arundinaria. We infer that plant (plants?) that produced the pollen of G. bambusoides could have grown in swamp forests and/or reed marshes. This report provides a general survey of the occurrence of bamboos in the European Neogene, from which the leaves and pollen documented here present important data on the distribution of this group of monocots in Poland and eastern Europe. D 2004 Elsevier B.V. All rights reserved. Keywords: bamboos; fossil leaves; pollen grains; Neogene; Poland; Beychato´w Lignite Mine; Legnica brown coal deposit; wetland vegetation
1. Introduction The bamboos include at present about 45 genera and belong to a distinctive group of the family Poaceae (grasses) that are assigned either to subfamily (Bambusoideae; Dahlgreen and Clifford, 1982) or tribe (Bambuseae; Hutchinson, 1973). This group is distinguished from other grasses in having usually woody culms (most are shrubs or trees), fruits including nuts, berries or caryopsis, and having three, six or more stamens per flower (see Hutchinson, * Corresponding author. Fax: +48 12 4219790. E-mail address: [email protected] (G. Worobiec). 0034-6667/$ - see front matter D 2004 Elsevier B.V. All rights reserved. doi:10.1016/j.revpalbo.2004.08.004
1973). Leaves of bamboos are characterised by the presence of a petiole between the leaf blade and the sheath (McClure, 1967; Hutchinson, 1973). This petiole is termed the bBambusaQ type (Dahlgreen and Clifford, 1982) and is found in family Poaceae mainly in bamboos, and rarely in other grasses. This feature is of crucial importance in systematics of fossil macroremains of leaves of Poaceae and is a key trait for including fossil leaves within bamboos. Fossil leaves of monocotyledons morphologically corresponding to bamboos were found together with numerous other plant remains from the Late Miocene leaf flora collected from the Beychato´w Lignite Mine, Central Poland. Deposits of similar age from Legnica
40
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
brown coal deposit in western Poland yield pollen grains morphologically typical of bamboos. The state of preservation of the fossils is generally very good and the material has proven to be suitable for detailed examination aimed to determine their taxonomical position. Accompanying plant remains, mostly very well preserved, were also investigated. Results of these investigations have offered the basis of deductions on the palaeoecology and the palaeogeography of bamboos in the Neogene of Poland and the Neogene floras of Central Europe.
2. Material and methods Macroremains of bamboo leaves were collected from the Beychato´w Lignite Mine in Central Poland, in the region between the rivers Warta and Pilica, ca. 15 km south of the Beychato´w city (Fig. 1). In this area, Tertiary deposits with lignites occur within a series of tectonic depressions called the Kleszczo´w Graben. Czarnecki et al. (1992), Stuchlik and Szynkiewicz (1998), and Matl (2000) distinguished four main lithological units from the Tertiary deposits from Beychato´w: a subcoal unit, a coal unit including the main economic seam, and two overlying sedimentary units; a clayey–coaly unit and the youngest clayey– sandy unit. Quaternary sediments are deposited on the Neogene deposits. Comprehensive geological, palae-
Fig. 1. Map of Poland with the location of the Beychato´w Lignite Mine and the Legnica brown coal deposit.
obotanical and palaeozoological study of the Tertiary deposits of the outcrop at the Beychato´w Lignite Mine has allowed the age of the deposits filling the Beychato´w Graben to be determined as Neogene, those of the main coal seam as Early Miocene, and those of the sedimentary series of the overburden as Middle/Late Miocene–Pliocene (Stuchlik et al., 1990; Stuchlik and Szynkiewicz, 1998; Worobiec and Lesiak, 1998; Wo´jcicki and Zastawniak, 1998; Szynkiewicz, 1999, 2000; Worobiec, 2003). The fossil flora containing remains of bamboo leaves was found in 1995. Deposits with the fossil flora were found within the base of the clayey–sandy unit. These sediments were deposited in an eroded depression (a fossil channel) at the boundary between the clayey–coaly unit and clayey–sandy unit. A layer of grey clays about 12 cm thick, with abundant leaf remains, graded upwards into a thick layer (ca. 25 cm) of compressed leaf litter. According to Wilczyn´ski (1992), deposits belonging to the floor part of the clayey–sandy unit were formed in fluvial environment that was transitional from a braided to meandering river system, and they originated as the result of filling of stagnant water bodies that may represent ox-bow lakes. Therefore, the sediments containing leaf remains of bamboo may represent an ox-bow lake deposits. The age of this flora is considered to be Late Miocene (Pannonian or Pontian) based on analysis of stratigraphical range of leaf taxa (Worobiec, 2003). Collected in the mine outcrop, bamboo leaves were slowly dried to avoid the specimens breaking up. The material was only investigated for its morphology as attempts to prepare leaf epidermal tissues failed. Morphological descriptions of fossil leaves were based on the terminology of Hickey and Peterson (1978) and Ash et al. (1999). Macrophotographs of the fossil leaves were made using NIKON Optiphot microscope and NIKON SMZ 800 stereomicroscope fitted with NIKON Coolpix 995 digital camera. For comparison with extant bamboos, the authors made use of the herbaria housed at the Wyadysyaw Szafer Institute of Botany, Polish Academy of Sciences, Krako´w. All the examined fossil specimens from the Beychato´w Lignite Mine are kept in the Department of Palaeobotany, Wyadysyaw Szafer Institute of Botany PAS, Krako´w (KRAM-P 217). The pollen grains of bamboos were found in the Legnica brown coal deposit complex that is situated in
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
the southwestern Poland on the Fore–Sudetic Block. This is a platform type deposit that extends over a large area in the Legnica Depression. Neogene deposits of the Legnica region are mainly of continental origin and are composed of clays, sands, gravels and lignites. In this area the following litostratigraphic units have been distinguished: the Z˙ary series with the third (S´cinawa) brown coal seam, the Silesian–Lusatian series with the second (Lusatian) seam, the Muyako´w series ended by the first (Henryk) seam, the Poznan´ series, and the Gozdnica series (Dyjor, 1978, 1986). These lithostratigraphic units are still used for the Legnica region even though, according to the lithostratigraphical scheme of the Polish Lowland, new stratigraphic units (formations and members) have been distinguished (Piwocki and Ziembin´ska-Tworzydyo, 1997). Exploratory drillings in the Legnica deposit were carried out at the end of the 1970s. The studied profile Legnica 33/56, situated in the eastern part of the Legnica brown coal deposit, was initially analysed by Sadowska et al. (1981). On the basis of sedimentological and mineralogical data, the Tertiary/Quaternary boundary was determined and, on the basis of palynological research, the Tertiary sedimentary series and brown coal seams were distinguished. In our study material from a depth of 74.0–77.0 and 90.0– 112.0 m has been used for palynological studies. The samples were taken at different intervals, mainly every 0.5 m or 1 m. A total of 46 samples were collected. Material for pollen analysis was prepared by a modified Erdtman’s acetolysis method (Moore et al., 1991), using HF. Only three studied samples were barren, and the data from the remaining 43 samples have been used to construct pollen diagrams. The sporomorphs were identified on the basis of available publications and the reference palynological collection of the Department of Palaeobotany, Wyadysyaw Szafer Institute of Botany, Polish Academy of Sciences in Krako´w. Microphotographs were taken using Leitz Laaborlux S microscope with Wild MPS 52 microcamera. Results of the pollen analysis show that the sedimentation of the examined deposits was limited to Badenian (?Late Karpatian–Late Badenian). Pollen grains of bamboos were found in samples taken from the Muyako´w series, a unit composed mainly of grey clays (Worobiec, 2000).
41
3. Macroremains of bamboo leaves from Belchato´w Lignite Mine Poaceae Bambusoideae bBambusaQ lugdunensis Saporta (Plate I, 1a–4; Plate II, 1a–2c) 1869 Bambusa lugdunensis Saporta; Saporta, p. 760. 1876 Bambusa lugdunensis Saporta; Saporta and Marion, p. 94, pl. 23, figs. 9–16. Material studied: KRAM-P 217: 48/II, 53/III, 58, 66/III, 76, 79, 83, 84, 123/IV, 127/II, 134/II, 145/II, 198, 225/II, 239. Description: Fragmentary, elongate leaves up to 6.0 cm long and 0.8–2.0 cm wide. Leaf base widely cuneate-obtuse, decurrent on petiole (Plate I, 1b, 3; Plate II, 1a). Leaf margin finely serrate, teeth very small, acute, strongly upmedially curved and orientated almost parallel to the margin (Plate II, 1b, 2b). Venation parallelodromous. Mid-vein distinct. On both sides of the mid-vein vein lie 4 to 6 (most commonly 4) less distinct lateral veins spaced 1.0–1.5 mm apart, between them lie 5 to 9 (most commonly 6 or 7) delicate, third order parallel veins, spaced 0.15– 0.20 mm apart (Plate II, 2a). Cross (transverse) veins connect adjacent parallel veins. The cross veins are oriented mostly perpendicular to them, spaced 0.15– 0.35 mm apart (Plate II, 2c). Remarks: Investigations on the morphology of monocotyledonous leaves found in the Late Miocene fossil flora from the Beychato´w Lignite Mine support their placement within the Poaceae. Because the bases of these leaves pass into petioles, they obviously represent leaves of a bamboo. The petiole is of the so-called bBambusaQ type typical of the subfamily Bambusoidae (Plate II, 3), and only rarely found in other grasses (Dahlgreen and Clifford, 1982). In the genus Phragmites Adanson, whose leaves are morphologically rather similar (Plate II, 4), the leaves lack a petiole and pass directly into a sheath. The remaining features, such as the parallel venation pattern and the leaf margin with minute teeth, are generally typical of grasses. The leaves from Beychato´w are most similar to bBambusaQ lugdunensis Saporta described from the Pliocene of Me´ximieux, France, by Saporta (1869), and illustrated by Saporta and Marion (1876). The
42
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
leaves under discussion clearly differ from another species of fossil bamboo, Bambusa ilinskiae Shvareva described from the Miocene of the Precarpatians (Shvareva, 1970), which has more lateral parallel veins (8–10 pairs at each side of the mid-vein vein, between which are 6–10 third order veins) and broader leaves, up to 5.5 cm. Sasa kodorica Kolakovsky, fossil bamboo leaves described from the Neogene of Kodor in Abkhazia (Kolakovsky, 1964), differ markedly from bBambusaQ lugdunensis. They are over twice as wide (up to 3.5 cm). This feature was considered by Kolakovsky (op. cit.) as characteristic of Sasa Mak. and Shibata and Pseudosasa Mak. For this reason, a new combination Sasa lugdunensis (Saporta) Givulescu created by Givulescu (1984) for relatively narrow leaves of bBambusaQ lugdunensis seems incorrect. The leaves from Beychato´w are similar in size and venation to those of Bambusium sp. B. from the Late Miocene flora of Tatsumigote (Honsiu, Japan, Ozaki, 1980). Poacites mengeanus Heer, which somewhat resembles bBambusaQ lugdunensis morphologically was reported by Heer (1869) from the Lower Miocene of Rozewie, and compared with the extant genera Uniola L., Panicum L., Bambusa Schreber and Arundinaria Michx. These remains described as Poacites mengeanus Heer may in fact belong to bBambusaQ lugdunensis. The genus name bBambusaQ is used with quotation marks as it is not possible to undoubtedly decide if the fossil bamboo leaves under discussion really belong to the recent genus Bambusa Schreber or not. This is due to the fact that, although leaf morphology and anatomy exhibit characteristics that could be used as diagnostic on the species level, they are not useful features to assign bamboo leaves to genera (Metcalfe, 1956; Bedell, 1997). Bedell (op. cit.) in her monograph on bamboo cites Gamble (1896) and Brandis
Plate I. 1a–4. 1a. 1b. 2. 3. 4.
(1907) namely that counting of the density of parallel veins on a quarter inch in bamboo leaves facilitate the identification of species. However, she wrote that in most bamboo genera the number of veins varies between 20 and 50 in a quarter inch and there is no information about the influence of the environment on this feature in her book. For example, leaves of genus Arundinaria have 21 to 36 nerves in quarter inch and leaves of Bambusa 20 to 40, the difference is indistinct and then this feature in taxonomy of bamboos genera seems not to be useful, at least in case of not all bamboos genera. Considering the above data on present knowledge of the morphology and anatomy of extant bamboo leaves, the genus name of fossil bamboo leaves from Beychato´w was remained as bBambusaQ.
4. Fossil pollen grains of bamboos from the Legnica brown coal deposit Poaceae Bambusoideae Graminidites Cookson 1947 ex Potonie´ 1960 Graminidites bambusoides Stuchlik 1994 (Plate III, 1–3) 1964 Gramineae bBambusaQ type; Stuchlik, p. 76, pl. 24, figs. 1–2. 1970 Graminidites sp. A (Bambusa—typus); Krutzsch, p. 51, pl. 1, fig. 1. 1994 Graminidites bambusoides Stuchlik sp. nov.; Ziembin´ska-Tworzydyo et al., p. 14, pl. 8, figs. 4–5. Description: Pollen grains monoporate, oval to spheroidal, 50–72 Am in diameter. Pore 4–6 Am in diameter, with annulus 4.0–4.5 Am broad. Exine composed of two layers, very thin (1.0–1.5 Am thick), with secondary folds, surface very finely granulate (Plate III, 1–3). Pollen grains often deformed.
bBambusaQ lugdunensis Saporta. Leaf, the Beychato´w Lignite Mine, No. KRAM-P 217/53. Enlarged leaf base with petiole, showing details of network of venation, No. KRAM-P 217/53. Leaf fragment, the Beychato´w Lignite Mine, No. KRAM-P 217/84. Fragmentary basal part of leaf with petiole, the Beychato´w Lignite Mine, No. KRAM-P 217/239. Fragments of leaves, the Beychato´w Lignite Mine, No. KRAM-P 217/58.
Scale bar=1 cm. 1a, 1b—phot. S. Florjan.
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
43
44
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
Remarks: These pollen grains closely correspond to the fossil taxon Graminidites bambusoides Stuchlik (Ziembin´ska-Tworzydyo et al., 1994). Pollen grains of this taxon have been relatively often encountered in various parts of the Rypin profile, from which Stuchlik (1964) compared them with those of the recent genera Bambusa Schreber, Oryza L. and Dendrocalamus C.G. Nees. In addition, Graminidites bambusoides from Legnica shows similarities to the recent pollen grains of Arundinaria Michx. (see Plate III, 4). Pollen grains of G. bambusoides were encountered in 12 samples from the Muyako´w series (Middle Miocene, Badenian) in the Legnica 33/56 profile, accounting for up to 2–6% (Worobiec, 2000). These rather large pollen grains of G. bambusoides that occur regularly in the Legnica profile were compared with the pollen of recent cereals to rule out the possibility that the pollen grains under study could be an artefact of contamination by recent pollen of cereals. They differ from pollen of the recent genus Triticum in having a thinner exine (Triticum—about 2 Am) and a more delicate annulus (see Plate III, 6). From the recent genus Secale they differ in shape and thinner exine, as well as fine grain surface (see Plate III, 5). The pollen grains of this fossil taxon, named by Stuchlik (1964) Gramineae bBambusaQ type, were subsequently named Graminidites sp. A (Bambusa typus) Krutzsch by Krutzsch (1970). Stuchlik (in Ziembin´ska-Tworzydyo et al., 1994) later named this species Graminidites bambusoides Stuchlik. The pollen grains of Graminidites cf. sp. A bBambusaQ type (Stuchlik) Planderova´ described from the Lower Miocene of southern Slovakia by Planderova´ (1990) essentially differ from Gramineae bBambusaQ type
Plate II. 1a–2c. 1a. 1b. 2a. 2b. 2c. 3. 4.
(=G. bambusoides) in a thicker exine and more distinct sculpture of the surface as well as in dimensions (pollen grain illustrated in Planderova´ (op. cit.) on pl. 54, fig. 1 is about 40 Am in diameter).
5. Discussion 5.1. Palaeoecology of fossil bamboos bBambusaQ lugdunensis, is in opinion of Laurent (1904–1905) closely related to extant bamboos of the genera Phyllostachys Sieb. and Zucc. and Arundinaria Michx., and especially to Arundinaria japonica Sieb. Grangeon (1958) regarded leaves of the genera Bambusa and Arundinaria as most similar to bBambusaQ lugdunensis. The present authors are of the opinion that bBambusaQ lugdunensis is similar to recent North American species of the genus Arundinaria: A. gigantea (Walter) Muhlenberg (=A. macrosperma Michx.) and A. tecta (Walter) Muhlenberg (Plate II, 3). This opinion is confirmed by comparison of the ecology of these two Arundinaria species and fossil bBambusaQ lugdunensis. Palaeoecological analysis of the fossil flora KRAM-P 217 in which leaves of bBambusaQ lugdunensis were found, points to the presence of wetland, swamp and riparian vegetation. Fossil taxa including Alnus gaudinii (Heer) Knobloch et Z. Kvacˇek, Alnus menzelii Raniecka-Bobrowska, Byttneriophyllum tiliaefolium (Al. Braun) Knobloch et Z. Kvacˇek, Sequoia abietina (Brongniart) Knobloch, Pinus sp., Smilax cf. weberi Wessel in Wessel et Weber and Smilax cf. protolancaefolia Kolakovsky were elements of the Neogene swamp forest, and Pterocarya paradisiaca (Unger) Iljinskaya and Dio-
bBambusaQ lugdunensis Saporta. Leaf base with short fragment of petiole, the Beychato´w Lignite Mine, No. KRAM-P 217/225. Enlargement of leaf margin, showing fine teeth, the Beychato´w Lignite Mine, No. KRAM-P 217/225. Details of leaf venation, the Beychato´w Lignite Mine, No. KRAM-P 217/58. Fine teeth on leaf margin, the Beychato´w Lignite Mine, No. KRAM-P 217/58. Details of leaf venation, showing cross veins, the Beychato´w Lignite Mine, No. KRAM-P 217/58. Arundinaria tecta (Walter) Muhlenberg, leaf base with petiole, recent. Herbarium of the Institute of Botany, Polish Academy of Sciences in Krako´w (KRAM), No. 048071. Phragmites australis (Cav.) Trin. ex Steud., leaf base, recent. Herbarium of Department of Palaeobotany, Institute of Botany, Polish Academy of Sciences in Krako´w.
1b, 2b—scale bar=0.5 mm; 1a, 2a, 2c—scale bar=1 mm; 3, 4—scale bar=1 cm. 3, 4—phot. S. Florjan.
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
45
46
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
spyros anceps Heer of the riparian forest vegetation (Worobiec, 2003). Recent Arundinaria gigantea and A. tecta occur in a similar type of wetland communities. Arundinaria gigantea is a component of swamp communities of southeastern North America, where it grows in swamp forests, for example in bDismal SwampsQ (Kearney, 1901). Besides, A. gigantea grows along rivers and stream banks as well as at open places and reaches up to 5 m height. Sometimes it forms an association (bcanbrakeQ) in which it dominates. The other species, A. tecta, at the same area is a component of bottomland forests and often grows on the riverbanks (Kearney, 1901). Similar deductions concerning the palaeoecology of fossil bamboos could be inferred on the basis of the analysis the palynological spectra from Legnica (Worobiec, 2000). In all samples Graminidites bambusoides Stuchlik is accompanied by pollen grains of representatives of the Taxodiaceae and Cupressaceae, genera Alnus, rarely Betula and Salix, and often by smaller pollen grains of other grasses (e.g. Graminidites pseudogramineus Krutzsch, G. subtiliglobosus (Trevisan) Krutzsch, G. neogenicus Krutzsch, and G. laevigatus Krutzsch). Single pollen grains of the family Cyperaceae (mainly of the Mariscus Vahl. type) and fern spores (Osmunda and Polypodiaceae) also occur. Taxodium and Alnus grow in wet habitats today, and accordingly they most probably were components of swamp forests that overgrew areas permanently inundated by water. Grasses and sedges could form swamp–aquatic reed associations, which are considered to be an early stage of the succession of the plant communities within Miocene mires. Deposits of these mires were the source of lignite seams (Teichmu¨ller, 1958; Sadowska, 1977). Traces of the reed associations are very rarely reported from the Tertiary pollen diagrams. As the extant counterparts to the Tertiary reed communities recent extensive reed marshes, so-called bEvergladesQ that occur in the
Plate III. 1–3. 4. 5. 6.
southern part of Florida could be considered (Gleason and Cronquist, 1968). To conclude it would seem that in the Neogene bamboos could be both components of reed associations and occur as an admixture in swamp and riparian forests like the recent Arundinaria gigantea and A. tecta in southeastern North America (Kearney, 1901; Johnson and Hoagland, 1999; Wunderlin and Hansen, 2003). The rare occurrence of bamboo pollen in fossil floras may be caused by the irregular and unpredictable in most species flowering cycle of these plants (McClure, 1967; Bedell, 1997). 5.2. Distribution of fossil remains of bamboos in the Neogene deposits of Europe Fossil macroremains of bamboos are very rare in the Cenozoic deposits of Europe. Most frequently are found leaves of bBambusaQ lugdunensis Saporta predominantly in Late Neogene fossil floras. bBambusaQ lugdunensis is known mainly from the Pliocene of southern France (Saporta, 1869; Saporta and Marion, 1876; Boulay, 1892; Laurent, 1904– 1905; Grangeon, 1958; Roiron, 1991) and is also known from Paleogene of Momcˇilovcˇy of Bulgaria (Kitanov and Palamarev, 1962), Sarmatian of Vienna (Berger and Zabusch, 1953), Pliocene of Transcarpathians (Iljinskaya, 1968), Pliocene of Asti, Italy (Martinetto, 2003) and Pliocene of Duab in Abkhazia (Kolakovsky, 1956). Shvareva (1970) described leaves of another fossil bamboo Bambusa ilinskiae Shvareva from Miocene deposits of Precarpathians. Leaves of Sasa kodorica Kolakovsky, described for the first time from Neogene flora of Kodor in Abkhazia (Kolakovsky, 1964) were found also in Sarmatian deposits of Ruzˇinci in Bulgaria (Palamarev and Petkova, 1987). Bamboo leaves were also reported from several fossil assemblages of Italy from Oligocene (Pavone d’Alessandria) through Miocene
Graminidites bambusoides Stuchlik, pollen, borehole Legnica 33/56, 99.0 m, sample No. 21. 1000. Arundinaria sp., pollen, recent, reference slide collection of Department of Palaeobotany, Institute of Botany, Polish Academy of Sciences in Krako´w. 1000. Secale cereale L., pollen, recent, reference slide collection of Department of Palaeobotany, Institute of Botany, Polish Academy of Sciences in Krako´w. 1000. Triticum aestivum L., pollen, recent, reference slide collection of Department of Palaeobotany, Institute of Botany, Polish Academy of Sciences in Krako´w. 1000.
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
47
48
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
(Nizza Monferrato) and Pliocene (Arboschio-L4) to Middle Pleistocene (Oriolo near Faenza) (Peola, 1900; Pavia, 1970; Martinetto and Sami, 2001; Brambilla and Gallo, 2002), from Pliocene of southern France (Roiron, 1979) and Neogene deposits of Chiuzbaia bHQ in Romania (Givulescu, 1984). In the Cenozoic fossil floras of Poland macroremains of bamboo leaves have not been reported so far, in contrast to the known occurrence of pollen grains of morphology corresponding to members of the bamboos—Gramineae bBambusaQ type. Stuchlik (1964) described such examples from the Miocene deposits of Rypin and until now it was the only locality of pollen grains of the bBambusaQ type (=Graminidites bambusoides) in the Cenozoic of Poland. In the Neogene of Europe they are extremely rare as well.
Acknowledgements Dr. Syawomir Florjan (Jagiellonian University in Krako´w) is acknowledged for help in preparing some macrophotographs. Dr. Jason Hilton is gratefully acknowledged for comments on the English text. The authors also thank the reviewers for their constructive comments that greatly improved the manuscript. The investigations on macroremains of bamboo leaves were financially supported by the grant No. 6 P04D 055 18 (bCharacteristics of the natural environment of the Neogene in central Poland on the basis of the leaf flora from the Lignite Mine in Beychato´wQ) of the State Committee for Scientific Research, Poland.
References Ash, A., Ellis, B., Hickey, L.J., Johnson, K., Wilf, P., Wing, S., 1999. Manual of Leaf Architecture—morphological description and categorization of dicotyledonous and net-veined monocotyledonous angiosperms by Leaf Architecture Working Group. Washington (Smithsonian Inst.). Bedell, P.E., 1997. Taxonomy of Bamboos. APC Publications PVT, New Delhi. Berger, W., Zabusch, F., 1953. Die obermioz7ne (sarmatische) Flora der Tqrkenschanze in Wien. N. Jb. Geol. Pal7ontol. 98 (2), 226 – 276. Boulay, N., 1892. Flore Plioce`ne du Mont-Dore (Puy-de-Doˆme). Savy, Paris.
Brambilla, G., Gallo, M., 2002. Analisi stratigrafica e paleobotanica della successione messiniana di Bric Santa Margherita (Nizza Monferrato, Asti, Italia NW). Boll. Mus. Reg. Sci. Nat. Torino 19 (1), 191 – 283. Brandis, D., 1907. Remarks on the structure of bamboo leaves. Trans. Linn. Soc. London., Second Ser., Bot. 7, 69 – 92. Czarnecki, L., Frankowski, R., Kuszneruk, J., 1992. Syntetyczny profil litostratygraficzny utworo´w trzeciorze˛du zyoya bBeychato´wQ. In: Lipiarski, I. (Ed.), Proc. XV Symp. bGeol. of Coal-bear. Strata of PolandQ. University of Mining and Metallurgy, Cracow, pp. 19 – 23. Dahlgreen, R.M.T., Clifford, H.T., 1982. The monocotyledones. A comparative study. In: Heywood, V.H. (Ed.), Botanical Systematics. An Occasional Series of Monographs. Academic Press, London. Dyjor, S., 1978. Wyksztaycenie i stratygrafia utworo´w trzeciorze˛dowych na obszarze Legnicko-Gyogowskiego Okre˛gu Miedziowego. In: Jerzman´ski, J. (Ed.), Przewodnik 50 Zjazdu PTG, pp. 210 – 214. Dyjor, S., 1986. Evolution of Sedimentation and Palaeogeography of Near-frontier Areas of the Silesian Part of the Parathetys and the Tertiary Polish–German Basin. Zesz. Nauk. Akad. Go´r.Hut., Geol. 12 (3), 7 – 23. Gamble, J.S., 1896. The Bambuseae of British India. Ann. R. Bot. Gard., 1 – 133. Givulescu, R., 1984. Die fossile Flora des Fundortes Chiuzbaia bHQ (Kreis Maramures, Rum7nien). D.S. Inst. Geol. Geofiz., Paleontol. 69 (1982), 69 – 93. Gleason, H.A., Cronquist, A., 1968. The Natural Geography of Plants. Columbia University Press, New York. Grangeon, P., 1958. Contribution a` l’e´tude de la pale´ontologie ve´ge´tale du Massif du Coiron (Arde`che). Me´m. Soc. Hist. Nat. 6, 1 – 299. Heer, O., 1869. Miocene baltische Flora. Beitr. Naturkd. Preuss. 2, 1 – 104. Hickey, L.J., Peterson, R.K., 1978. Zingiberopsis, a fossil genus of the ginger family from Late Cretaceous to early Eocene sediments of Western Interior North America. Can. J. Bot. 56, 1136 – 1152. Hutchinson, J., 1973. The Families of Flowering Plants. Clarendon Press, Oxford. Iljinskaya, I.A., 1968. Neogenovye flory Zakarpatskoi oblasti USSR (summary: Neogene floras of the Transcarpathian Region of the U.S.S.R.). Izdatel’stvo Nauka. Leningrad. Johnson, F.L., Hoagland, B.W., 1999. Catalog of the Woody Plants of Oklahoma: Descriptions and Range Maps (http://www. biosurvey.ou.edu/shrub/argit8.htm). Oklahoma Biological Survey and the University of Oklahoma. Kearney, T.H., 1901. Report on botanical survey of the Dismal swamp region. Contr. Nat. Herb., Washington 5 (6), 1 – 585. Kitanov, B., Palamarev, E., 1962. Beitrag zum Studium der Terti7rflora Bulgarensis (in Bulgarian, German summary). Ann. Univ. de Sophia 54–55 (1), 1 – 16. Kolakovsky, A.A., 1956. Pliotsenovaya flora Duava (summary: Pliocene flora of Duab). Tr. Sukhum. Bot. Sada 9, 213 – 309. Kolakovsky, A.A., 1964. Pliotsenovaya flora Kodora (summary: A Pliocene flora of the Kodor River). Sukhum. Bot. Sad. Monogr. 1, 1 – 209.
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50 Krutzsch, W., 1970. Atlas der mittel- und jungterti7ren dispersen Sporen- und Pollen- sowie der Mikroplanktonformen des nfrdlichen Mitteleuropas VII. VEB Gustav Fischer Verlag, Jena. Laurent, L., 1904–1905. Flore Plioce`ne des cine´rites du Pas-de-laMougudo et de Saint-Vincent-La Sabie (Cantal). Ann. Mus. D’Histoire Nat. de Marseille, Geol. 9, 1 – 313. Martinetto, E., 2003. Leaves of terrestrial plants from the Pliocene shallow marine and transitional deposits of Asti (Piedmont NW Italy). Boll. Soc. Paleontol. Ital. 42 (1–2), 75 – 111. Martinetto, E., Sami, M., 2001. Paleoflora delle bsabbie gialleQ mediopleistoceniche di Oriolo, presso Faenza (Ravenna). Quad. Studi. Nat. Romagna 14, 1 – 28. Matl, K., 2000. Wybrane elementy budowy geologicznej zyoya Beychato´w (summary: selected elements of geological structure of the Beychato´w deposit). In: Syomka, T., Wagner, M. (Eds.), Charakter petrograficzny i warunki sedymentacji wybranych komplekso´w litologicznych z profilu miocenu w zyoyu we˛gla brunatnego Beychato´w (Petrological studies and sedimentological conditions of select lithologic series in Miocene from Beychato´w Lignite deposit, Poland). Komisja Nauk Geologicznych PAN, oddziay w Krakowie, Prace Geologiczne 147, 11–19. McClure, F.A., 1967. The Bamboos. A Fresh Perspective. Harvard University Press, Cambridge. Metcalfe, C.R., 1956. Some thoughts on the structure of bamboo leaves. Bot. Mag. Tokyo 69, 391 – 400. Moore, P.D., Webb, J.A., Collinson, M.E., 1991. Pollen Analysis. Blackbell Scient. Publ., Oxford. Ozaki, K., 1980. Late Miocene Tatsumitoge Flora of Tottori Prefecture, Southwest Honshu, Japan (III). Sci. Rep. Yokohama Natl. Univ., Sect. 1, Math., Phys., Chem. 2 (27), 19 – 45. Palamarev, E.H., Petkova, A.S., 1987. Les fossiles de Bulgarie, 8/1: La macroflore du Sarmatien. L’Academie Bulgare des Sciences, Sophia. Pavia, G., 1970. Resti di Anancus arvensis e flora ad affinita´ plioceniche nel Villafranchiano inferiore della cava di Arboschio (Villafranca d’Asti). Mem. Soc. Geol. Ital. 9, 157 – 176. Peola, P., 1900. Flora tongriana di Pavone d’Alessandria. Boll. Soc. Geol. Ital. 19, 36 – 62. Piwocki, M., Ziembin´ska-Tworzydyo, M., 1997. Neogene of the Polish Lowlands—litostratigraphy and pollen-spore zones. Kwart. Geol. 41 (1), 21 – 40. Planderova´, E., 1990. Miocene microflora of Slovak Central Paratethys and its biostratigraphical significance. Diony´z Sˇtu´r Instytute of Geology, Bratislava. Roiron, P., 1979. Recherches sur les Flores Plio-Quaternaires Mediterraneennes: La Macroflore Pliocene de Pichegu Pres de Saint-Gilles (Gard). These, Acad. de Montpellier. Univ. des Sci. et Tech. du Languedoc, 1–221. Roiron, P., 1991. Les macroflores d’age Miocene Superieur des diatomites de Murat (Cantal France). Implication paleoclimatiques. Palaeontographica B 223, 169 – 203. Sadowska, A., 1977. Ros´linnos´c´ i stratygrafia go´rnomiocen´skich pokyado´w we˛gla Polski poyudniowo-zachodniej (summary: vegetation and stratigraphy of Upper Miocene coal seam of the South-Western Poland). Acta Palaeobot. 18 (1), 87 – 122.
49
Sadowska, A., Grodzicki, A., Kuszell, T., 1981. Badania stratygraficzne serii skalnych rejonu zyoya Legnica (Pole bwschodnieQ). Archive of the Inst. of Geol. Sciences. Wrocyaw University, Wrocyaw. Saporta, G., 1869. Sur l’existence de plusieurs espe`ces actuelles observe´es dans la flore Plioce`ne de Me´ximieux. Bull. Soc. Ge´ol. Fr. 2 (26), 752 – 773. Saporta, G., Marion, A.F., 1876. Recherches sur les ve´ge´taux fossiles Me´ximieux. Arch. Mus. d’Hist. Nat. Lyon. 1, 131 – 335. Shvareva, N.Y., 1970. Miotsenovaya flora Pistynki v Predkarpat’e [The Miocene flora of Pistynka (Cis-Carpathian Area)]. Bot. Z. 55 (3), 443 – 456. Stuchlik, L., 1964. Pollen analysis of the Miocene deposits at Rypin. Acta Palaeobot. 5 (2), 1 – 111. Stuchlik, L., Szynkiewicz, A., 1998. General geological situation and palynological investigations of the brown coal deposits. In: Sadowska, A., Szynkiewicz, A. (Eds.), Guide to Excursion No 2: Tertiary–Quaternary (Pleistocene) floras of Beychato´w (Middle Poland) and several localities in south-western Poland. The Fifth European Palaeobotanical–Palynological Conference in Cracow, 26-30.06.1998. IB PAN, Krako´w, pp. 6–11. Stuchlik, L., Szynkiewicz, A., xan´cucka-S´rodoniowa, M., Zastawniak, E., 1990. Wyniki dotychczasowych badan´ paleobotanicznych trzeciorze˛dowych we˛gli brunatnych zyoya bBeychato´wQ (summary: Results of the hitherto palaeobotanical investigations of the Tertiary brown coal bed bBeychato´wQ (Central Poland). Acta Palaeobot. 30 (1–2), 259 – 305. Szynkiewicz, A., 1999. Wiek utworo´w trzeciorze˛dowych odkrywki bBeychato´wQ. XX Konferencja Terenowa Sekcji Tektonicznej PTG bMyodoalpejski Ro´w Kleszczowa: rozwo´j i uwarunkowania w tektonice regionu. Syok k. Beychatowa, 15–16 paz´dziernika 1999, 143–147. Szynkiewicz, A., 2000. Wiek we˛gla brunatnego na tle pozycji geologicznej badanych pro´bek (KWB bBeychato´wQ) (Age of the brown coal deposits from Beychato´w lignite mine (Central Poland). Prz. Geol. 48, 1038 – 1044. Teichmqller, M., 1958. Rekonstruktionen verschiedner Moortypen des Hauptflfzes der niederrheinischen Braunkohle. Fortschr. Geol. Rheinl. Westfal. 2, 599 – 612. Wilczyn´ski, R., 1992. Dotychczasowe wyniki badan´ podstawowych serii poznan´skiej w s´wietle geologiczno-inyynierskich problemo´ w prowadzenia robo´ t go´ rniczych w KWB bBeychato´wQ (summary: the hitherto existing results of the Poznan´ suite in the light of geological-engineering problems of carrying mining works in the bBeychato´wQ brown coal open mine). Pr. Geol.-Mineral. (Acta Univ. Wratislav.) 1354 24, 91 – 108. Worobiec, E., 2000. Palinoflora neogenu wschodniej cze˛s´ci legnickiego kompleksu zyo´ y we˛gla brunatnego (Neogene palynoflora of the eastern part of the Legnica brown coal deposit complex). PhD Thesis. Archive of the W. Szafer Institute of Botany, Polish Academy of Sciences, Krako´w. Worobiec, G., 2003. New fossil floras from Neogene deposits in the Beychato´w Lignite Mine. Acta Palaeobot., Suppl. 3, 1 – 133. Worobiec, G., Lesiak, M., 1998. Plant megafossils from the Neogene deposits of Stawek-1A (Beychato´w, Middle Poland). Rev. Palaeobot. Palynol. 101, 179 – 208.
50
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
Wo´jcicki, J.J., Zastawniak, E., 1998. Trapa srodoniana, a new fossil species from the Pliocene of Beychato´w (Middle Poland). Acta Palaeobot. 38 (1), 167 – 174. Wunderlin, R.P., Hansen, B.F., 2003. Atlas of Florida Vascular Plants (http://www.plantatlas.usf.edu/). [Landry, S.M., Campbell, K.N. (application development), Florida Center for Community Design and Research.] Institute for Systematic Botany, University of South Florida, Tampa.
Ziembin´ska-Tworzydyo, M., Grabowska, I., Kohlman-Adamska, A., Skawin´ska, K., Syodkowska, B., Stuchlik, L., Sadowska, A., Wayyn´ska, H., 1994. Taxonomical revision of selected pollen and spores taxa from Neogene deposits. In: Stuchlik L. (Ed.), Neogene pollen flora of Central Europe. Part 1. Acta Palaeobot., Suppl. 1, pp. 5–30.
Leaves and pollen of bamboos from the Polish Neogene Elybieta Worobiec, Grzegorz Worobiec* WlCadyslCaw Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, PL-31-512 Krako´w, Poland Received 9 January 2004; accepted 26 August 2004
Abstract Recently discovered fossil bamboo leaves and pollen are described from the Neogene of Poland. Morphological analysis of fossil leaves from the Late Miocene deposits of the Beychato´w Lignite Mine, Central Poland have allowed them to be determined as bBambusaQ lugdunensis Saporta. It is the first record of this species in the Cenozoic of Poland. The fossils have been compared with other records of bamboos from the Cenozoic of Europe. Pollen grains from the Middle Miocene (Badenian) deposits from the Legnica brown coal deposit (Lower Silesia) provide additional evidence of bamboos. These are assigned to Graminidites bambusoides Stuchlik, and are similar to those of recent Arundinaria. We infer that plant (plants?) that produced the pollen of G. bambusoides could have grown in swamp forests and/or reed marshes. This report provides a general survey of the occurrence of bamboos in the European Neogene, from which the leaves and pollen documented here present important data on the distribution of this group of monocots in Poland and eastern Europe. D 2004 Elsevier B.V. All rights reserved. Keywords: bamboos; fossil leaves; pollen grains; Neogene; Poland; Beychato´w Lignite Mine; Legnica brown coal deposit; wetland vegetation
1. Introduction The bamboos include at present about 45 genera and belong to a distinctive group of the family Poaceae (grasses) that are assigned either to subfamily (Bambusoideae; Dahlgreen and Clifford, 1982) or tribe (Bambuseae; Hutchinson, 1973). This group is distinguished from other grasses in having usually woody culms (most are shrubs or trees), fruits including nuts, berries or caryopsis, and having three, six or more stamens per flower (see Hutchinson, * Corresponding author. Fax: +48 12 4219790. E-mail address: [email protected] (G. Worobiec). 0034-6667/$ - see front matter D 2004 Elsevier B.V. All rights reserved. doi:10.1016/j.revpalbo.2004.08.004
1973). Leaves of bamboos are characterised by the presence of a petiole between the leaf blade and the sheath (McClure, 1967; Hutchinson, 1973). This petiole is termed the bBambusaQ type (Dahlgreen and Clifford, 1982) and is found in family Poaceae mainly in bamboos, and rarely in other grasses. This feature is of crucial importance in systematics of fossil macroremains of leaves of Poaceae and is a key trait for including fossil leaves within bamboos. Fossil leaves of monocotyledons morphologically corresponding to bamboos were found together with numerous other plant remains from the Late Miocene leaf flora collected from the Beychato´w Lignite Mine, Central Poland. Deposits of similar age from Legnica
40
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
brown coal deposit in western Poland yield pollen grains morphologically typical of bamboos. The state of preservation of the fossils is generally very good and the material has proven to be suitable for detailed examination aimed to determine their taxonomical position. Accompanying plant remains, mostly very well preserved, were also investigated. Results of these investigations have offered the basis of deductions on the palaeoecology and the palaeogeography of bamboos in the Neogene of Poland and the Neogene floras of Central Europe.
2. Material and methods Macroremains of bamboo leaves were collected from the Beychato´w Lignite Mine in Central Poland, in the region between the rivers Warta and Pilica, ca. 15 km south of the Beychato´w city (Fig. 1). In this area, Tertiary deposits with lignites occur within a series of tectonic depressions called the Kleszczo´w Graben. Czarnecki et al. (1992), Stuchlik and Szynkiewicz (1998), and Matl (2000) distinguished four main lithological units from the Tertiary deposits from Beychato´w: a subcoal unit, a coal unit including the main economic seam, and two overlying sedimentary units; a clayey–coaly unit and the youngest clayey– sandy unit. Quaternary sediments are deposited on the Neogene deposits. Comprehensive geological, palae-
Fig. 1. Map of Poland with the location of the Beychato´w Lignite Mine and the Legnica brown coal deposit.
obotanical and palaeozoological study of the Tertiary deposits of the outcrop at the Beychato´w Lignite Mine has allowed the age of the deposits filling the Beychato´w Graben to be determined as Neogene, those of the main coal seam as Early Miocene, and those of the sedimentary series of the overburden as Middle/Late Miocene–Pliocene (Stuchlik et al., 1990; Stuchlik and Szynkiewicz, 1998; Worobiec and Lesiak, 1998; Wo´jcicki and Zastawniak, 1998; Szynkiewicz, 1999, 2000; Worobiec, 2003). The fossil flora containing remains of bamboo leaves was found in 1995. Deposits with the fossil flora were found within the base of the clayey–sandy unit. These sediments were deposited in an eroded depression (a fossil channel) at the boundary between the clayey–coaly unit and clayey–sandy unit. A layer of grey clays about 12 cm thick, with abundant leaf remains, graded upwards into a thick layer (ca. 25 cm) of compressed leaf litter. According to Wilczyn´ski (1992), deposits belonging to the floor part of the clayey–sandy unit were formed in fluvial environment that was transitional from a braided to meandering river system, and they originated as the result of filling of stagnant water bodies that may represent ox-bow lakes. Therefore, the sediments containing leaf remains of bamboo may represent an ox-bow lake deposits. The age of this flora is considered to be Late Miocene (Pannonian or Pontian) based on analysis of stratigraphical range of leaf taxa (Worobiec, 2003). Collected in the mine outcrop, bamboo leaves were slowly dried to avoid the specimens breaking up. The material was only investigated for its morphology as attempts to prepare leaf epidermal tissues failed. Morphological descriptions of fossil leaves were based on the terminology of Hickey and Peterson (1978) and Ash et al. (1999). Macrophotographs of the fossil leaves were made using NIKON Optiphot microscope and NIKON SMZ 800 stereomicroscope fitted with NIKON Coolpix 995 digital camera. For comparison with extant bamboos, the authors made use of the herbaria housed at the Wyadysyaw Szafer Institute of Botany, Polish Academy of Sciences, Krako´w. All the examined fossil specimens from the Beychato´w Lignite Mine are kept in the Department of Palaeobotany, Wyadysyaw Szafer Institute of Botany PAS, Krako´w (KRAM-P 217). The pollen grains of bamboos were found in the Legnica brown coal deposit complex that is situated in
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
the southwestern Poland on the Fore–Sudetic Block. This is a platform type deposit that extends over a large area in the Legnica Depression. Neogene deposits of the Legnica region are mainly of continental origin and are composed of clays, sands, gravels and lignites. In this area the following litostratigraphic units have been distinguished: the Z˙ary series with the third (S´cinawa) brown coal seam, the Silesian–Lusatian series with the second (Lusatian) seam, the Muyako´w series ended by the first (Henryk) seam, the Poznan´ series, and the Gozdnica series (Dyjor, 1978, 1986). These lithostratigraphic units are still used for the Legnica region even though, according to the lithostratigraphical scheme of the Polish Lowland, new stratigraphic units (formations and members) have been distinguished (Piwocki and Ziembin´ska-Tworzydyo, 1997). Exploratory drillings in the Legnica deposit were carried out at the end of the 1970s. The studied profile Legnica 33/56, situated in the eastern part of the Legnica brown coal deposit, was initially analysed by Sadowska et al. (1981). On the basis of sedimentological and mineralogical data, the Tertiary/Quaternary boundary was determined and, on the basis of palynological research, the Tertiary sedimentary series and brown coal seams were distinguished. In our study material from a depth of 74.0–77.0 and 90.0– 112.0 m has been used for palynological studies. The samples were taken at different intervals, mainly every 0.5 m or 1 m. A total of 46 samples were collected. Material for pollen analysis was prepared by a modified Erdtman’s acetolysis method (Moore et al., 1991), using HF. Only three studied samples were barren, and the data from the remaining 43 samples have been used to construct pollen diagrams. The sporomorphs were identified on the basis of available publications and the reference palynological collection of the Department of Palaeobotany, Wyadysyaw Szafer Institute of Botany, Polish Academy of Sciences in Krako´w. Microphotographs were taken using Leitz Laaborlux S microscope with Wild MPS 52 microcamera. Results of the pollen analysis show that the sedimentation of the examined deposits was limited to Badenian (?Late Karpatian–Late Badenian). Pollen grains of bamboos were found in samples taken from the Muyako´w series, a unit composed mainly of grey clays (Worobiec, 2000).
41
3. Macroremains of bamboo leaves from Belchato´w Lignite Mine Poaceae Bambusoideae bBambusaQ lugdunensis Saporta (Plate I, 1a–4; Plate II, 1a–2c) 1869 Bambusa lugdunensis Saporta; Saporta, p. 760. 1876 Bambusa lugdunensis Saporta; Saporta and Marion, p. 94, pl. 23, figs. 9–16. Material studied: KRAM-P 217: 48/II, 53/III, 58, 66/III, 76, 79, 83, 84, 123/IV, 127/II, 134/II, 145/II, 198, 225/II, 239. Description: Fragmentary, elongate leaves up to 6.0 cm long and 0.8–2.0 cm wide. Leaf base widely cuneate-obtuse, decurrent on petiole (Plate I, 1b, 3; Plate II, 1a). Leaf margin finely serrate, teeth very small, acute, strongly upmedially curved and orientated almost parallel to the margin (Plate II, 1b, 2b). Venation parallelodromous. Mid-vein distinct. On both sides of the mid-vein vein lie 4 to 6 (most commonly 4) less distinct lateral veins spaced 1.0–1.5 mm apart, between them lie 5 to 9 (most commonly 6 or 7) delicate, third order parallel veins, spaced 0.15– 0.20 mm apart (Plate II, 2a). Cross (transverse) veins connect adjacent parallel veins. The cross veins are oriented mostly perpendicular to them, spaced 0.15– 0.35 mm apart (Plate II, 2c). Remarks: Investigations on the morphology of monocotyledonous leaves found in the Late Miocene fossil flora from the Beychato´w Lignite Mine support their placement within the Poaceae. Because the bases of these leaves pass into petioles, they obviously represent leaves of a bamboo. The petiole is of the so-called bBambusaQ type typical of the subfamily Bambusoidae (Plate II, 3), and only rarely found in other grasses (Dahlgreen and Clifford, 1982). In the genus Phragmites Adanson, whose leaves are morphologically rather similar (Plate II, 4), the leaves lack a petiole and pass directly into a sheath. The remaining features, such as the parallel venation pattern and the leaf margin with minute teeth, are generally typical of grasses. The leaves from Beychato´w are most similar to bBambusaQ lugdunensis Saporta described from the Pliocene of Me´ximieux, France, by Saporta (1869), and illustrated by Saporta and Marion (1876). The
42
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
leaves under discussion clearly differ from another species of fossil bamboo, Bambusa ilinskiae Shvareva described from the Miocene of the Precarpatians (Shvareva, 1970), which has more lateral parallel veins (8–10 pairs at each side of the mid-vein vein, between which are 6–10 third order veins) and broader leaves, up to 5.5 cm. Sasa kodorica Kolakovsky, fossil bamboo leaves described from the Neogene of Kodor in Abkhazia (Kolakovsky, 1964), differ markedly from bBambusaQ lugdunensis. They are over twice as wide (up to 3.5 cm). This feature was considered by Kolakovsky (op. cit.) as characteristic of Sasa Mak. and Shibata and Pseudosasa Mak. For this reason, a new combination Sasa lugdunensis (Saporta) Givulescu created by Givulescu (1984) for relatively narrow leaves of bBambusaQ lugdunensis seems incorrect. The leaves from Beychato´w are similar in size and venation to those of Bambusium sp. B. from the Late Miocene flora of Tatsumigote (Honsiu, Japan, Ozaki, 1980). Poacites mengeanus Heer, which somewhat resembles bBambusaQ lugdunensis morphologically was reported by Heer (1869) from the Lower Miocene of Rozewie, and compared with the extant genera Uniola L., Panicum L., Bambusa Schreber and Arundinaria Michx. These remains described as Poacites mengeanus Heer may in fact belong to bBambusaQ lugdunensis. The genus name bBambusaQ is used with quotation marks as it is not possible to undoubtedly decide if the fossil bamboo leaves under discussion really belong to the recent genus Bambusa Schreber or not. This is due to the fact that, although leaf morphology and anatomy exhibit characteristics that could be used as diagnostic on the species level, they are not useful features to assign bamboo leaves to genera (Metcalfe, 1956; Bedell, 1997). Bedell (op. cit.) in her monograph on bamboo cites Gamble (1896) and Brandis
Plate I. 1a–4. 1a. 1b. 2. 3. 4.
(1907) namely that counting of the density of parallel veins on a quarter inch in bamboo leaves facilitate the identification of species. However, she wrote that in most bamboo genera the number of veins varies between 20 and 50 in a quarter inch and there is no information about the influence of the environment on this feature in her book. For example, leaves of genus Arundinaria have 21 to 36 nerves in quarter inch and leaves of Bambusa 20 to 40, the difference is indistinct and then this feature in taxonomy of bamboos genera seems not to be useful, at least in case of not all bamboos genera. Considering the above data on present knowledge of the morphology and anatomy of extant bamboo leaves, the genus name of fossil bamboo leaves from Beychato´w was remained as bBambusaQ.
4. Fossil pollen grains of bamboos from the Legnica brown coal deposit Poaceae Bambusoideae Graminidites Cookson 1947 ex Potonie´ 1960 Graminidites bambusoides Stuchlik 1994 (Plate III, 1–3) 1964 Gramineae bBambusaQ type; Stuchlik, p. 76, pl. 24, figs. 1–2. 1970 Graminidites sp. A (Bambusa—typus); Krutzsch, p. 51, pl. 1, fig. 1. 1994 Graminidites bambusoides Stuchlik sp. nov.; Ziembin´ska-Tworzydyo et al., p. 14, pl. 8, figs. 4–5. Description: Pollen grains monoporate, oval to spheroidal, 50–72 Am in diameter. Pore 4–6 Am in diameter, with annulus 4.0–4.5 Am broad. Exine composed of two layers, very thin (1.0–1.5 Am thick), with secondary folds, surface very finely granulate (Plate III, 1–3). Pollen grains often deformed.
bBambusaQ lugdunensis Saporta. Leaf, the Beychato´w Lignite Mine, No. KRAM-P 217/53. Enlarged leaf base with petiole, showing details of network of venation, No. KRAM-P 217/53. Leaf fragment, the Beychato´w Lignite Mine, No. KRAM-P 217/84. Fragmentary basal part of leaf with petiole, the Beychato´w Lignite Mine, No. KRAM-P 217/239. Fragments of leaves, the Beychato´w Lignite Mine, No. KRAM-P 217/58.
Scale bar=1 cm. 1a, 1b—phot. S. Florjan.
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
43
44
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
Remarks: These pollen grains closely correspond to the fossil taxon Graminidites bambusoides Stuchlik (Ziembin´ska-Tworzydyo et al., 1994). Pollen grains of this taxon have been relatively often encountered in various parts of the Rypin profile, from which Stuchlik (1964) compared them with those of the recent genera Bambusa Schreber, Oryza L. and Dendrocalamus C.G. Nees. In addition, Graminidites bambusoides from Legnica shows similarities to the recent pollen grains of Arundinaria Michx. (see Plate III, 4). Pollen grains of G. bambusoides were encountered in 12 samples from the Muyako´w series (Middle Miocene, Badenian) in the Legnica 33/56 profile, accounting for up to 2–6% (Worobiec, 2000). These rather large pollen grains of G. bambusoides that occur regularly in the Legnica profile were compared with the pollen of recent cereals to rule out the possibility that the pollen grains under study could be an artefact of contamination by recent pollen of cereals. They differ from pollen of the recent genus Triticum in having a thinner exine (Triticum—about 2 Am) and a more delicate annulus (see Plate III, 6). From the recent genus Secale they differ in shape and thinner exine, as well as fine grain surface (see Plate III, 5). The pollen grains of this fossil taxon, named by Stuchlik (1964) Gramineae bBambusaQ type, were subsequently named Graminidites sp. A (Bambusa typus) Krutzsch by Krutzsch (1970). Stuchlik (in Ziembin´ska-Tworzydyo et al., 1994) later named this species Graminidites bambusoides Stuchlik. The pollen grains of Graminidites cf. sp. A bBambusaQ type (Stuchlik) Planderova´ described from the Lower Miocene of southern Slovakia by Planderova´ (1990) essentially differ from Gramineae bBambusaQ type
Plate II. 1a–2c. 1a. 1b. 2a. 2b. 2c. 3. 4.
(=G. bambusoides) in a thicker exine and more distinct sculpture of the surface as well as in dimensions (pollen grain illustrated in Planderova´ (op. cit.) on pl. 54, fig. 1 is about 40 Am in diameter).
5. Discussion 5.1. Palaeoecology of fossil bamboos bBambusaQ lugdunensis, is in opinion of Laurent (1904–1905) closely related to extant bamboos of the genera Phyllostachys Sieb. and Zucc. and Arundinaria Michx., and especially to Arundinaria japonica Sieb. Grangeon (1958) regarded leaves of the genera Bambusa and Arundinaria as most similar to bBambusaQ lugdunensis. The present authors are of the opinion that bBambusaQ lugdunensis is similar to recent North American species of the genus Arundinaria: A. gigantea (Walter) Muhlenberg (=A. macrosperma Michx.) and A. tecta (Walter) Muhlenberg (Plate II, 3). This opinion is confirmed by comparison of the ecology of these two Arundinaria species and fossil bBambusaQ lugdunensis. Palaeoecological analysis of the fossil flora KRAM-P 217 in which leaves of bBambusaQ lugdunensis were found, points to the presence of wetland, swamp and riparian vegetation. Fossil taxa including Alnus gaudinii (Heer) Knobloch et Z. Kvacˇek, Alnus menzelii Raniecka-Bobrowska, Byttneriophyllum tiliaefolium (Al. Braun) Knobloch et Z. Kvacˇek, Sequoia abietina (Brongniart) Knobloch, Pinus sp., Smilax cf. weberi Wessel in Wessel et Weber and Smilax cf. protolancaefolia Kolakovsky were elements of the Neogene swamp forest, and Pterocarya paradisiaca (Unger) Iljinskaya and Dio-
bBambusaQ lugdunensis Saporta. Leaf base with short fragment of petiole, the Beychato´w Lignite Mine, No. KRAM-P 217/225. Enlargement of leaf margin, showing fine teeth, the Beychato´w Lignite Mine, No. KRAM-P 217/225. Details of leaf venation, the Beychato´w Lignite Mine, No. KRAM-P 217/58. Fine teeth on leaf margin, the Beychato´w Lignite Mine, No. KRAM-P 217/58. Details of leaf venation, showing cross veins, the Beychato´w Lignite Mine, No. KRAM-P 217/58. Arundinaria tecta (Walter) Muhlenberg, leaf base with petiole, recent. Herbarium of the Institute of Botany, Polish Academy of Sciences in Krako´w (KRAM), No. 048071. Phragmites australis (Cav.) Trin. ex Steud., leaf base, recent. Herbarium of Department of Palaeobotany, Institute of Botany, Polish Academy of Sciences in Krako´w.
1b, 2b—scale bar=0.5 mm; 1a, 2a, 2c—scale bar=1 mm; 3, 4—scale bar=1 cm. 3, 4—phot. S. Florjan.
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
45
46
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
spyros anceps Heer of the riparian forest vegetation (Worobiec, 2003). Recent Arundinaria gigantea and A. tecta occur in a similar type of wetland communities. Arundinaria gigantea is a component of swamp communities of southeastern North America, where it grows in swamp forests, for example in bDismal SwampsQ (Kearney, 1901). Besides, A. gigantea grows along rivers and stream banks as well as at open places and reaches up to 5 m height. Sometimes it forms an association (bcanbrakeQ) in which it dominates. The other species, A. tecta, at the same area is a component of bottomland forests and often grows on the riverbanks (Kearney, 1901). Similar deductions concerning the palaeoecology of fossil bamboos could be inferred on the basis of the analysis the palynological spectra from Legnica (Worobiec, 2000). In all samples Graminidites bambusoides Stuchlik is accompanied by pollen grains of representatives of the Taxodiaceae and Cupressaceae, genera Alnus, rarely Betula and Salix, and often by smaller pollen grains of other grasses (e.g. Graminidites pseudogramineus Krutzsch, G. subtiliglobosus (Trevisan) Krutzsch, G. neogenicus Krutzsch, and G. laevigatus Krutzsch). Single pollen grains of the family Cyperaceae (mainly of the Mariscus Vahl. type) and fern spores (Osmunda and Polypodiaceae) also occur. Taxodium and Alnus grow in wet habitats today, and accordingly they most probably were components of swamp forests that overgrew areas permanently inundated by water. Grasses and sedges could form swamp–aquatic reed associations, which are considered to be an early stage of the succession of the plant communities within Miocene mires. Deposits of these mires were the source of lignite seams (Teichmu¨ller, 1958; Sadowska, 1977). Traces of the reed associations are very rarely reported from the Tertiary pollen diagrams. As the extant counterparts to the Tertiary reed communities recent extensive reed marshes, so-called bEvergladesQ that occur in the
Plate III. 1–3. 4. 5. 6.
southern part of Florida could be considered (Gleason and Cronquist, 1968). To conclude it would seem that in the Neogene bamboos could be both components of reed associations and occur as an admixture in swamp and riparian forests like the recent Arundinaria gigantea and A. tecta in southeastern North America (Kearney, 1901; Johnson and Hoagland, 1999; Wunderlin and Hansen, 2003). The rare occurrence of bamboo pollen in fossil floras may be caused by the irregular and unpredictable in most species flowering cycle of these plants (McClure, 1967; Bedell, 1997). 5.2. Distribution of fossil remains of bamboos in the Neogene deposits of Europe Fossil macroremains of bamboos are very rare in the Cenozoic deposits of Europe. Most frequently are found leaves of bBambusaQ lugdunensis Saporta predominantly in Late Neogene fossil floras. bBambusaQ lugdunensis is known mainly from the Pliocene of southern France (Saporta, 1869; Saporta and Marion, 1876; Boulay, 1892; Laurent, 1904– 1905; Grangeon, 1958; Roiron, 1991) and is also known from Paleogene of Momcˇilovcˇy of Bulgaria (Kitanov and Palamarev, 1962), Sarmatian of Vienna (Berger and Zabusch, 1953), Pliocene of Transcarpathians (Iljinskaya, 1968), Pliocene of Asti, Italy (Martinetto, 2003) and Pliocene of Duab in Abkhazia (Kolakovsky, 1956). Shvareva (1970) described leaves of another fossil bamboo Bambusa ilinskiae Shvareva from Miocene deposits of Precarpathians. Leaves of Sasa kodorica Kolakovsky, described for the first time from Neogene flora of Kodor in Abkhazia (Kolakovsky, 1964) were found also in Sarmatian deposits of Ruzˇinci in Bulgaria (Palamarev and Petkova, 1987). Bamboo leaves were also reported from several fossil assemblages of Italy from Oligocene (Pavone d’Alessandria) through Miocene
Graminidites bambusoides Stuchlik, pollen, borehole Legnica 33/56, 99.0 m, sample No. 21. 1000. Arundinaria sp., pollen, recent, reference slide collection of Department of Palaeobotany, Institute of Botany, Polish Academy of Sciences in Krako´w. 1000. Secale cereale L., pollen, recent, reference slide collection of Department of Palaeobotany, Institute of Botany, Polish Academy of Sciences in Krako´w. 1000. Triticum aestivum L., pollen, recent, reference slide collection of Department of Palaeobotany, Institute of Botany, Polish Academy of Sciences in Krako´w. 1000.
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
47
48
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
(Nizza Monferrato) and Pliocene (Arboschio-L4) to Middle Pleistocene (Oriolo near Faenza) (Peola, 1900; Pavia, 1970; Martinetto and Sami, 2001; Brambilla and Gallo, 2002), from Pliocene of southern France (Roiron, 1979) and Neogene deposits of Chiuzbaia bHQ in Romania (Givulescu, 1984). In the Cenozoic fossil floras of Poland macroremains of bamboo leaves have not been reported so far, in contrast to the known occurrence of pollen grains of morphology corresponding to members of the bamboos—Gramineae bBambusaQ type. Stuchlik (1964) described such examples from the Miocene deposits of Rypin and until now it was the only locality of pollen grains of the bBambusaQ type (=Graminidites bambusoides) in the Cenozoic of Poland. In the Neogene of Europe they are extremely rare as well.
Acknowledgements Dr. Syawomir Florjan (Jagiellonian University in Krako´w) is acknowledged for help in preparing some macrophotographs. Dr. Jason Hilton is gratefully acknowledged for comments on the English text. The authors also thank the reviewers for their constructive comments that greatly improved the manuscript. The investigations on macroremains of bamboo leaves were financially supported by the grant No. 6 P04D 055 18 (bCharacteristics of the natural environment of the Neogene in central Poland on the basis of the leaf flora from the Lignite Mine in Beychato´wQ) of the State Committee for Scientific Research, Poland.
References Ash, A., Ellis, B., Hickey, L.J., Johnson, K., Wilf, P., Wing, S., 1999. Manual of Leaf Architecture—morphological description and categorization of dicotyledonous and net-veined monocotyledonous angiosperms by Leaf Architecture Working Group. Washington (Smithsonian Inst.). Bedell, P.E., 1997. Taxonomy of Bamboos. APC Publications PVT, New Delhi. Berger, W., Zabusch, F., 1953. Die obermioz7ne (sarmatische) Flora der Tqrkenschanze in Wien. N. Jb. Geol. Pal7ontol. 98 (2), 226 – 276. Boulay, N., 1892. Flore Plioce`ne du Mont-Dore (Puy-de-Doˆme). Savy, Paris.
Brambilla, G., Gallo, M., 2002. Analisi stratigrafica e paleobotanica della successione messiniana di Bric Santa Margherita (Nizza Monferrato, Asti, Italia NW). Boll. Mus. Reg. Sci. Nat. Torino 19 (1), 191 – 283. Brandis, D., 1907. Remarks on the structure of bamboo leaves. Trans. Linn. Soc. London., Second Ser., Bot. 7, 69 – 92. Czarnecki, L., Frankowski, R., Kuszneruk, J., 1992. Syntetyczny profil litostratygraficzny utworo´w trzeciorze˛du zyoya bBeychato´wQ. In: Lipiarski, I. (Ed.), Proc. XV Symp. bGeol. of Coal-bear. Strata of PolandQ. University of Mining and Metallurgy, Cracow, pp. 19 – 23. Dahlgreen, R.M.T., Clifford, H.T., 1982. The monocotyledones. A comparative study. In: Heywood, V.H. (Ed.), Botanical Systematics. An Occasional Series of Monographs. Academic Press, London. Dyjor, S., 1978. Wyksztaycenie i stratygrafia utworo´w trzeciorze˛dowych na obszarze Legnicko-Gyogowskiego Okre˛gu Miedziowego. In: Jerzman´ski, J. (Ed.), Przewodnik 50 Zjazdu PTG, pp. 210 – 214. Dyjor, S., 1986. Evolution of Sedimentation and Palaeogeography of Near-frontier Areas of the Silesian Part of the Parathetys and the Tertiary Polish–German Basin. Zesz. Nauk. Akad. Go´r.Hut., Geol. 12 (3), 7 – 23. Gamble, J.S., 1896. The Bambuseae of British India. Ann. R. Bot. Gard., 1 – 133. Givulescu, R., 1984. Die fossile Flora des Fundortes Chiuzbaia bHQ (Kreis Maramures, Rum7nien). D.S. Inst. Geol. Geofiz., Paleontol. 69 (1982), 69 – 93. Gleason, H.A., Cronquist, A., 1968. The Natural Geography of Plants. Columbia University Press, New York. Grangeon, P., 1958. Contribution a` l’e´tude de la pale´ontologie ve´ge´tale du Massif du Coiron (Arde`che). Me´m. Soc. Hist. Nat. 6, 1 – 299. Heer, O., 1869. Miocene baltische Flora. Beitr. Naturkd. Preuss. 2, 1 – 104. Hickey, L.J., Peterson, R.K., 1978. Zingiberopsis, a fossil genus of the ginger family from Late Cretaceous to early Eocene sediments of Western Interior North America. Can. J. Bot. 56, 1136 – 1152. Hutchinson, J., 1973. The Families of Flowering Plants. Clarendon Press, Oxford. Iljinskaya, I.A., 1968. Neogenovye flory Zakarpatskoi oblasti USSR (summary: Neogene floras of the Transcarpathian Region of the U.S.S.R.). Izdatel’stvo Nauka. Leningrad. Johnson, F.L., Hoagland, B.W., 1999. Catalog of the Woody Plants of Oklahoma: Descriptions and Range Maps (http://www. biosurvey.ou.edu/shrub/argit8.htm). Oklahoma Biological Survey and the University of Oklahoma. Kearney, T.H., 1901. Report on botanical survey of the Dismal swamp region. Contr. Nat. Herb., Washington 5 (6), 1 – 585. Kitanov, B., Palamarev, E., 1962. Beitrag zum Studium der Terti7rflora Bulgarensis (in Bulgarian, German summary). Ann. Univ. de Sophia 54–55 (1), 1 – 16. Kolakovsky, A.A., 1956. Pliotsenovaya flora Duava (summary: Pliocene flora of Duab). Tr. Sukhum. Bot. Sada 9, 213 – 309. Kolakovsky, A.A., 1964. Pliotsenovaya flora Kodora (summary: A Pliocene flora of the Kodor River). Sukhum. Bot. Sad. Monogr. 1, 1 – 209.
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50 Krutzsch, W., 1970. Atlas der mittel- und jungterti7ren dispersen Sporen- und Pollen- sowie der Mikroplanktonformen des nfrdlichen Mitteleuropas VII. VEB Gustav Fischer Verlag, Jena. Laurent, L., 1904–1905. Flore Plioce`ne des cine´rites du Pas-de-laMougudo et de Saint-Vincent-La Sabie (Cantal). Ann. Mus. D’Histoire Nat. de Marseille, Geol. 9, 1 – 313. Martinetto, E., 2003. Leaves of terrestrial plants from the Pliocene shallow marine and transitional deposits of Asti (Piedmont NW Italy). Boll. Soc. Paleontol. Ital. 42 (1–2), 75 – 111. Martinetto, E., Sami, M., 2001. Paleoflora delle bsabbie gialleQ mediopleistoceniche di Oriolo, presso Faenza (Ravenna). Quad. Studi. Nat. Romagna 14, 1 – 28. Matl, K., 2000. Wybrane elementy budowy geologicznej zyoya Beychato´w (summary: selected elements of geological structure of the Beychato´w deposit). In: Syomka, T., Wagner, M. (Eds.), Charakter petrograficzny i warunki sedymentacji wybranych komplekso´w litologicznych z profilu miocenu w zyoyu we˛gla brunatnego Beychato´w (Petrological studies and sedimentological conditions of select lithologic series in Miocene from Beychato´w Lignite deposit, Poland). Komisja Nauk Geologicznych PAN, oddziay w Krakowie, Prace Geologiczne 147, 11–19. McClure, F.A., 1967. The Bamboos. A Fresh Perspective. Harvard University Press, Cambridge. Metcalfe, C.R., 1956. Some thoughts on the structure of bamboo leaves. Bot. Mag. Tokyo 69, 391 – 400. Moore, P.D., Webb, J.A., Collinson, M.E., 1991. Pollen Analysis. Blackbell Scient. Publ., Oxford. Ozaki, K., 1980. Late Miocene Tatsumitoge Flora of Tottori Prefecture, Southwest Honshu, Japan (III). Sci. Rep. Yokohama Natl. Univ., Sect. 1, Math., Phys., Chem. 2 (27), 19 – 45. Palamarev, E.H., Petkova, A.S., 1987. Les fossiles de Bulgarie, 8/1: La macroflore du Sarmatien. L’Academie Bulgare des Sciences, Sophia. Pavia, G., 1970. Resti di Anancus arvensis e flora ad affinita´ plioceniche nel Villafranchiano inferiore della cava di Arboschio (Villafranca d’Asti). Mem. Soc. Geol. Ital. 9, 157 – 176. Peola, P., 1900. Flora tongriana di Pavone d’Alessandria. Boll. Soc. Geol. Ital. 19, 36 – 62. Piwocki, M., Ziembin´ska-Tworzydyo, M., 1997. Neogene of the Polish Lowlands—litostratigraphy and pollen-spore zones. Kwart. Geol. 41 (1), 21 – 40. Planderova´, E., 1990. Miocene microflora of Slovak Central Paratethys and its biostratigraphical significance. Diony´z Sˇtu´r Instytute of Geology, Bratislava. Roiron, P., 1979. Recherches sur les Flores Plio-Quaternaires Mediterraneennes: La Macroflore Pliocene de Pichegu Pres de Saint-Gilles (Gard). These, Acad. de Montpellier. Univ. des Sci. et Tech. du Languedoc, 1–221. Roiron, P., 1991. Les macroflores d’age Miocene Superieur des diatomites de Murat (Cantal France). Implication paleoclimatiques. Palaeontographica B 223, 169 – 203. Sadowska, A., 1977. Ros´linnos´c´ i stratygrafia go´rnomiocen´skich pokyado´w we˛gla Polski poyudniowo-zachodniej (summary: vegetation and stratigraphy of Upper Miocene coal seam of the South-Western Poland). Acta Palaeobot. 18 (1), 87 – 122.
49
Sadowska, A., Grodzicki, A., Kuszell, T., 1981. Badania stratygraficzne serii skalnych rejonu zyoya Legnica (Pole bwschodnieQ). Archive of the Inst. of Geol. Sciences. Wrocyaw University, Wrocyaw. Saporta, G., 1869. Sur l’existence de plusieurs espe`ces actuelles observe´es dans la flore Plioce`ne de Me´ximieux. Bull. Soc. Ge´ol. Fr. 2 (26), 752 – 773. Saporta, G., Marion, A.F., 1876. Recherches sur les ve´ge´taux fossiles Me´ximieux. Arch. Mus. d’Hist. Nat. Lyon. 1, 131 – 335. Shvareva, N.Y., 1970. Miotsenovaya flora Pistynki v Predkarpat’e [The Miocene flora of Pistynka (Cis-Carpathian Area)]. Bot. Z. 55 (3), 443 – 456. Stuchlik, L., 1964. Pollen analysis of the Miocene deposits at Rypin. Acta Palaeobot. 5 (2), 1 – 111. Stuchlik, L., Szynkiewicz, A., 1998. General geological situation and palynological investigations of the brown coal deposits. In: Sadowska, A., Szynkiewicz, A. (Eds.), Guide to Excursion No 2: Tertiary–Quaternary (Pleistocene) floras of Beychato´w (Middle Poland) and several localities in south-western Poland. The Fifth European Palaeobotanical–Palynological Conference in Cracow, 26-30.06.1998. IB PAN, Krako´w, pp. 6–11. Stuchlik, L., Szynkiewicz, A., xan´cucka-S´rodoniowa, M., Zastawniak, E., 1990. Wyniki dotychczasowych badan´ paleobotanicznych trzeciorze˛dowych we˛gli brunatnych zyoya bBeychato´wQ (summary: Results of the hitherto palaeobotanical investigations of the Tertiary brown coal bed bBeychato´wQ (Central Poland). Acta Palaeobot. 30 (1–2), 259 – 305. Szynkiewicz, A., 1999. Wiek utworo´w trzeciorze˛dowych odkrywki bBeychato´wQ. XX Konferencja Terenowa Sekcji Tektonicznej PTG bMyodoalpejski Ro´w Kleszczowa: rozwo´j i uwarunkowania w tektonice regionu. Syok k. Beychatowa, 15–16 paz´dziernika 1999, 143–147. Szynkiewicz, A., 2000. Wiek we˛gla brunatnego na tle pozycji geologicznej badanych pro´bek (KWB bBeychato´wQ) (Age of the brown coal deposits from Beychato´w lignite mine (Central Poland). Prz. Geol. 48, 1038 – 1044. Teichmqller, M., 1958. Rekonstruktionen verschiedner Moortypen des Hauptflfzes der niederrheinischen Braunkohle. Fortschr. Geol. Rheinl. Westfal. 2, 599 – 612. Wilczyn´ski, R., 1992. Dotychczasowe wyniki badan´ podstawowych serii poznan´skiej w s´wietle geologiczno-inyynierskich problemo´ w prowadzenia robo´ t go´ rniczych w KWB bBeychato´wQ (summary: the hitherto existing results of the Poznan´ suite in the light of geological-engineering problems of carrying mining works in the bBeychato´wQ brown coal open mine). Pr. Geol.-Mineral. (Acta Univ. Wratislav.) 1354 24, 91 – 108. Worobiec, E., 2000. Palinoflora neogenu wschodniej cze˛s´ci legnickiego kompleksu zyo´ y we˛gla brunatnego (Neogene palynoflora of the eastern part of the Legnica brown coal deposit complex). PhD Thesis. Archive of the W. Szafer Institute of Botany, Polish Academy of Sciences, Krako´w. Worobiec, G., 2003. New fossil floras from Neogene deposits in the Beychato´w Lignite Mine. Acta Palaeobot., Suppl. 3, 1 – 133. Worobiec, G., Lesiak, M., 1998. Plant megafossils from the Neogene deposits of Stawek-1A (Beychato´w, Middle Poland). Rev. Palaeobot. Palynol. 101, 179 – 208.
50
E. Worobiec, G. Worobiec / Review of Palaeobotany and Palynology 133 (2005) 39–50
Wo´jcicki, J.J., Zastawniak, E., 1998. Trapa srodoniana, a new fossil species from the Pliocene of Beychato´w (Middle Poland). Acta Palaeobot. 38 (1), 167 – 174. Wunderlin, R.P., Hansen, B.F., 2003. Atlas of Florida Vascular Plants (http://www.plantatlas.usf.edu/). [Landry, S.M., Campbell, K.N. (application development), Florida Center for Community Design and Research.] Institute for Systematic Botany, University of South Florida, Tampa.
Ziembin´ska-Tworzydyo, M., Grabowska, I., Kohlman-Adamska, A., Skawin´ska, K., Syodkowska, B., Stuchlik, L., Sadowska, A., Wayyn´ska, H., 1994. Taxonomical revision of selected pollen and spores taxa from Neogene deposits. In: Stuchlik L. (Ed.), Neogene pollen flora of Central Europe. Part 1. Acta Palaeobot., Suppl. 1, pp. 5–30.