A reconsideration of the diversity of Symplocos in the ...

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A taxonomic reassessment of Symplocos from the Neogene of Europe is proposed ... we dramatically reduced the number of species in the European Neogene.

Review of Palaeobotany and Palynology 140 (2006) 1 – 26 www.elsevier.com/locate/revpalbo

A reconsideration of the diversity of Symplocos in the European Neogene on the basis of fruit morphology D.H. Mai a,b , E. Martinetto a,⁎ b

a Università degli Studi di Torino, Italy Museum für Naturkunde Humboldt Universität zu Berlin, Germany

Received 4 November 2004; received in revised form 3 January 2006; accepted 4 February 2006 Available online 24 March 2006

Abstract A taxonomic reassessment of Symplocos from the Neogene of Europe is proposed based on morphological and anatomical features of fruit stones at the macro- and mesoscopic scale. In light of the large extent of polymorphism observed in fossil and recent stones, we dramatically reduced the number of species in the European Neogene. Only 11 out of 26 fossil species cited in the literature of the last two decades were still considered valid, and, due to the species concept we applied, we suggest that this number roughly corresponds to the number of ancient botanical species which actually grew in the lowlands of Europe from 30 to 2 My ago. The characters of the valid species are extensively illustrated to provide a guide for species identification. An updated overview of the stratigraphic and geographic distribution of the species of the European Neogene is also provided. A comparative analysis of fossil fruits and analogous modern material (poorly available) has been carried out with the aim to derive climatic information from modern plants. © 2006 Elsevier B.V. All rights reserved. Keywords: Symplocos; fruits; diversity; Europe; Neogene

1. Introduction The genus Symplocos, which presently is not native to Europe, has a rich fossil record in this area (Fig. 1), both of pollen (Krutzsch, 1989) and macrofossils (mainly fruits). Symplocos fruits are also known in North America from a few Eocene localities (Manchester, 1994; Tiffney, 1999) and the early Miocene Brandon Lignite (Tiffney, 1994, 1999). All the modern species of Symplocos produce drupes containing a decay-resistant lignified endocarp (stone), which often shows species-diagnostic characters (Fig. 2). Thus, the

⁎ Corresponding author. E-mail address: [email protected] (E. Martinetto). 0034-6667/$ - see front matter © 2006 Elsevier B.V. All rights reserved. doi:10.1016/j.revpalbo.2006.02.001

analysis of the morphological diversity of fossil endocarps can be considered a useful tool to reconstruct the past specific diversity in Symplocos, especially in central Europe, where such remains have been reported from several sites, sometimes in huge quantities (Kirchheimer, 1950, 1957; Mai, 1970; Basilici et al., 1997). In Palaeogene localities of Europe, 14 fruit species assigned to Symplocos have been reported (Ettingshausen, 1877; Chandler, 1926; Reid and Chandler, 1933; Kirchheimer, 1939a; Chandler, 1960b, 1961, 1962, 1963; Mai, 1970), but in our opinion only 7 species (Symplocos casparyi Ludwig, Symplocos headonensis Chandler, Symplocos kirstei Kirchheimer, Symplocos lakensis Chandler, Symplocos minutula (Sternberg) Kirchheimer, Symplocos trilocularis Reid and Chandler,

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Fig. 1. Map of Europe showing localities cited in the text. 1: Lower Rhine Embayment (Adendorf, Düren, Fortuna-Garsdorf I, Hambach, Konzendorf, Niederpleis, Reuver); 2: Sessenheim; 3: Vogelsberg (Salzhausen) and Wetterau (Dorheim, Weckesheim); 4: Oberpfalz; 5: Lusatia (Dauban, Hartau, Kausche, Merka, Puschwitz, Vetschau, Wiesa) and Saxony (Brandis, Regis, Weisselster Basin); 6: Turow and Lower Silesia; 7: Stare Gliwice and Wieliczka, 8: Rypin; 9: Oberdorf and Weingraben; 10: Pont-de-Gail; 11: Boca, Buronzo, Caʻ Viettone, Candelo, Castellengo, Castelletto Cervo II, Sento II, Pocapaglia; 12: central Italy (Tuscany, Umbria).

Kirchheimer) show sufficient diagnostic characters for a reliable assignment to this genus. Three of these species (S. headonensis, S. lakensis, and S. trilocularis) are restricted to the Paleogene, and we are not able to confirm, at present, if they actually represent separate taxonomic entities. On the other hand we studied in detail those species which also occur in Neogene deposits (S. casparyi, S. kirstei, S. minutula, and S. pseudogregaria), and we concluded that only three of

Fig. 2. Morphological characters of Symplocos stones.

them represent separate taxonomic entities (S. kirstei is a synonym of S. pseudogregaria, see below). In Neogene localities of Europe, more than 30 fruit species assigned to Symplocos have been reported, but only 26 species are cited in the literature of the last two decades. The recent findings of very rich populations of fossil endocarps from the Neogene of the Lower Rhine Embayment, Lusatia and Italy have been of fundamental importance for understanding the full extent of variation within several taxonomic entities, and suggested to us that a reconsideration of the specific diversity was needed. Therefore, we provide here a complete taxonomic revision of Neogene fruits of Symplocos from Europe, including an extensive iconography to facilitate species identification. We also provide a contribution to palaeoclimatic analyses. In fact, the occurrence of mega-mesotherm plants, such as the Symplocaceae, has been traditionally used to show that the early–middle Cenozoic climate in Europe was warmer than today (Mai, 1995). This concept has been recently incorporated into the “coexistence approach” (Mosbrugger and Utescher, 1997), which aims to assess a more precise quantification of the palaeoclimatic conditions under which fossil taxa occurred based on the climatic preferences of their nearest living relatives. The correct definition of the living relatives within Symplocos requires a careful comparison of fruit characters in modern and fossil species of the genus. Unfortunately, this target cannot

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be fully achieved in the present work because the fruit characters and their variation in many modern species of Southeast Asia (Nooteboom, 1975) have not been described in sufficient detail. Furthermore, the fruits of such species are not adequately represented in the reference collections of modern material available to us. Thus, our conclusions are largely based on the analysis of Caribbean material (Mai, 1986), whereas the features of East Asian material have been inferred by the literature (Nooteboom, 1975; Nagamasu, 1993), supplemented by direct observations on several species obtained from carpological collections (MNB, Szafer Botanical Institute of Crakow). 2. Material and methods The present study is based on re-examination of the morphological and anatomical features of Symplocos stones from the European Neogene at the macro- and mesoscopic scale. Microscopic characters, such as presence/absence of crystal idioblasts (which had been considered taxonomically relevant by Kirchheimer, 1936, 1950), have been excluded from consideration in this treatment, because we observed that they can be variable within a single species (e.g. Symplocos casparyi Ludwig). A thorough exam of the taxonomic value of this character requires assessment through a large number of thin sections, which is at present not available. In particular, we studied all the Neogene fruits assigned to Symplocos which are stored in the collections of the Museum für Naturkunde of Berlin (specimens cited with the prefix, MNB), Forschungsinstitut Senckenberg of Frankfurt am Main (FSF) and Museo di Geologia e Paleontologia of the Turin University (PU). We also studied selected material, including all-type specimens, stored at British Museum of Natural History in London (BMNH), Museum für Mineralogie und Geologie in Dresden (MMGD), Staatliches Museum für Naturkunde (Stuttgart), Geol.Palaeont. Institut (Tübingen), Laboratory of Palaeobotany and Palynology (Utrecht), Geological Survey of The Netherlands (Utrecht), and the Szafer Botanical Institute (Crakow). Whenever possible, the type material of each species (or at least the topotypic material) has been comparatively re-analysed, with particular attention to diagnostic features, and character variation within each population. As in Mai (1986), we deem it useful to distinguish several fruit morphological types in both fossil and

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modern material based on shape (terminology follows Anderberg, 1994), number of seed locules and apical pores, features of the apical pit, permanent or ephemeral connection of the endocarp to the mesocarp, and type of ornamentation: Ampulliformis-type (Mai, 1986). Named after the shape: Stones ampulliform to urceolate, sometimes nearly oval, with an apical neck bearing a marginal bulge, but without an apical pit. Base rounded to conical. Axis straight, surface rugose to longitudinally striate. Locules 3 in the early stage of development, but later always aborted to 1, slightly curved locule. Central canal moved to one side. Endocarp wall thin and elastic. No fossil species in Europe. Modern species in America: Symplocos ovalis Wright ex Griseb., Symplocos salicifolia Griseb., Symplocos guadeloupensis Krug et Urb., and Symplocos domingensis Urb. Ciponimastrum-type (Mai, 1986). Named after subsection Ciponimastrum Brand, 1901. Endocarp obovoid or ellipsoid, apex broad and truncate, with a deep pit, usually lacking a marginal bulge (apart the fossil Symplocos pseudogregaria), base rounded, longitudinal axis occasionally oblique. Stone surface longitudinally striate or rugose but not ribbed. Mesocarp sometimes persistent, providing the external aspect of the fossil fruits. Locules 3 to 5, often unevenly developed, rarely reduced to unrecognisability (sterile). Central canal never laterally shifted. Endocarp thick, stony. Fossil species: Symplocos arecaeformis, Symplocos pseudogregaria, Symplocos gothanii. Modern species in America: Symplocos limonicillo Humb. et Bonpl., Symplocos berteroi (A. DC.) Miers, Symplocos lindeniana Krug et Urb., Symplocos martinicensis Jacq., Symplocos latifolia Krug et Urb., Symplocos tubulifera Krug et Urb. In Asia: Symplocos lucida (Thunb.) Sieb. and Zucc., Symplocos henschelii (Mor.) Benth ex Clarke, Symplocos tingifera Chen. Germanica-type (Mai, 1986). Named after the fossil Symplocos germanica Mai. Endocarps narrowly ovoid to obovoid, rarely ellipsoid, apically truncate, with deep pit and thin margin, basally attenuate or rounded, longitudinal axis more or less straight to oblique. Surface with distinct transverse wrinkles interrupted by a fine longitudinal striation. Locules 3 in the early stage of development, but not more than 2 locules become fertile. Central canal laterally shifted. Endocarp thin or of medium thickness. Fossil species: only Symplocos

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germanica. Modern species in America: Symplocos lanata Krug et Urb., Symplocos leonis Britt. and Wils., Symplocos moaensis Borhidi. In Southeast Asia: Symplocos filipes Nooteboom. Minutula-type (Mai, 1986). Named after the fossil Symplocos minutula (Sternberg) Kirchheimer from the Miocene of central Europe. Stones ovoid. Apically attenuate without a marginal bulge around the apical pit. Base rounded, longitudinal stone-axis curved. Surface nearly smooth to finely striated. Locules 3, unequally developed, mostly aborted to 1, sometimes oblique. Central canal laterally shifted. Endocarp of medium thickness. Fossil species: only Symplocos minutula. Modern species in America: Symplocos tinctoria L'Herit. In Southeast Asia: Symplocos ramosissima Wallich ex G. Don. Palura-type. Named after section Palura G. Don. Similar to the ampulliformis-type, but stone dorsiventrally flattened, apical pit present without a marginal bulge; locules 2 in the early stage of development, only one (rarely 2) of which becomes large and fertile; central canal lacking; small basal pit laterally shifted and crossed by a horseshoeshaped peripheral furrow. Fossil species: Symplocos paucicostata. Modern species: Symplocos paniculata (Thunb. ex Murray) Miq. s. l. Sulcata-type. Named after the modern Symplocos sulcata Kurz. Stones narrowly obovoid to ellipsoid to globose, apex abruptly truncate, with a deep pit, small to large (Symplocos schereri, fossil) without marginal bulge, basally rounded or conical. Surface with strong and often nearly straight longitudinal ridges, but also with faint ridges to nearly smooth in a single fossil species (Symplocos casparyi). Locules 3, rarely 4, mostly equally developed. Central canal never laterally shifted. Endocarp of medium thickness, stony. Fossil species: Symplocos casparyi, Symplocos schereri, Symplocos pseudoschereri, Symplocos tetraporina. Modern species: Symplocos sulcata Kurz, Symplocos tanakae Matsamura, etc. (see under Symplocos casparyi and Symplocos schereri). Previously Mai (1986) included this in the Ciponimasrum-type, but here it is distinguished for its importance and easy identification in the Neogene of Europe. The foregoing fruit-types, apart from the Paluratype, do not seem to have phylogenetic relevance, since they are widely distributed within the “traditional” subgenera and sections (Nooteboom, 2004), which, however, are regarded to be of limited taxonomic value to judge from ongoing DNA analyses

(Wang et al., 2004), suggesting that the infrageneric classification of Symplocos is in strong need of reassessment. 3. Valid species Symplocos arecaeformis (Schlotheim 1820) Kirchheimer 1950 Plate I, 1–6 Original: Carpolithus arecaeformis Schlotheim (1820: 420, without fig., nomen illegit.), from Liblar, Lower Rhine Embayment (Germany), middle Miocene, not found in the Schlotheim collection at MNB (lost). Synonyms: Symplocos sapperi Kirchheimer (1934: 789, Fig. 18) from Plessa near Senftenberg, Lusatia (Germany), coll. Braunkohlenmuseum Senftenberg, destroyed in World War II. Discussion: The name Carpolithus arecaeformis Schlotheim, 1820 is nomenclaturally non binding because it was introduced before the date of 31 December 1820, fixed as the beginning for valid publication of fossil plant names (ICBN, Art. 13.1.f). After Kilpper (1969, p. 123) a possible valid new name is Cocos faujasii (Brongniart, 1828). Geographic and stratigraphic distribution: Very rare, to our knowledge a single fragmentary specimen (Plate I, 6) is available in existing collections (MNB). The few lost specimens were obtained from lower and middle Miocene brown coal beds of Germany. Description: Fruit with a solid thick mesocarp, persistent around the endocarp, ovoid, of the ciponimastrum-type, 5-locular, mostly one locule abortive, thin-ribbed, unusually large: 25–45 mm long. Apical pit 1/3 of total fruit diameter, shallow, round-edged. Similar modern species: A modern fruit from Guatemala (Jalapa, Tierra Fria, mountain rain forest), labelled “Symplocos purpurea” in the MNB collection (the name does not exist within the genus), has comparably large, 5-loculed coarsely rugose stones, whose external appearance is presumably given by the mesocarp. Symplocos limonicillo Humb. et Bonpl. is similar but smaller and narrower; the specimens observed by us were again collected in Guatemala (Jalapa). According to Nooteboom's description (1975), Symplocos henschelii (Mor.) Benth ex Clarke also seems to produce similar fruits: about 30 mm long, obovoid, often oblique, 3–4 loculed, mesocarp rather hard, fleshy or even woody. It grows in the rainforests of Burma, Indochina, Malaya, Sumatra, West Java and Borneo, at 45–1700 m a.s.l. Climatic indications: The modern species grow under subtropical moist conditions, MAT > 20 °C,

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MAP > 2000 mm. Additionally, as a brown-coal plant, Symplocos arecaeformis should have grown under moist, warm temperate to subtropical conditions. Symplocos braunii Kirchheimer 1935c Plate I, 7; Text—Fig. 3 Original: Symplocos braunii (Kirchheimer, 1935c:718 and 737, Fig. 20) from “CarpolithenKohle” of Salzhausen, Vogelsberg (Germany), middle Miocene. Represented only by three specimens, formerly stored at Wett. Ges. f. d. Naturkunde Hanau am Main (Kirchheimer, 1937), but lost during World War II (V. Wilde, personal communication). Geographic and stratigraphic distribution: Endemic to the Vogelsberg volcanic area. Description: An unclear species, also because no more specimens are available for accurate comparative studies. Stone with mesocarp sticking on the lobed endocarp of the sulcata-type, always 3-locular, 9– 12 mm long, obovoid. Apical pit 1/3 of total fruit diameter, deep, thick-edged. Wall with cavities. Similar modern species: Unknown. Symplocos casparyi Ludwig 1857 Plate I, 8–15; Plate II, 1–8, and Plate III, 1–8 Original: Symplocos casparyi Ludwig (1857: 99, Pl. 20, Figs. 6a–f), from Dorheim in the Wetterau near Friedberg (Germany), lower Pliocene. Type-material: 1 specimen detected (Nov. 2003) in Ludwig's collection from Dorheim (MNB 2004/825), in between the stones labelled as Symplocos ludwigii, is designated here as the neotype. Synonyms: Myrica granulosa Ludwig (1857: 91, Pl. 20, Figs. 29a– b) = Symplocos granulosa (Ludwig) Mai (1973: 101, Pl. 5, Figs. 1–3), from Dorheim near Friedberg, Weterau, Germany, preserved in MNB (original – no. 964). Symplocos globosa Ludwig (1857: 98–99, Pl. 20, Figs. a–e), from Dorheim, type material lost. Carpinus salzhausenensis Ludwig (1860: Pl. 33, Fig. 8) = Symplocos salzhausenensis (Ludwig, 1860) Kirchheimer (1935c, Fig. 19), from Salzhausen (Germany), middle Miocene; holotype stored in MNB (2004/821: Plate I, 13). Carpolithus lignitarum Quenstedt (1867: 914, Pl. 86, Fig. 35) = Symplocos lignitarum (Quenstedt, 1867) Kirchheimer (1935c: 718, Figs. 17–18), from Salzhausen (Germany), middle Miocene, according to Kirchheimer (1957) stored in Geol.-Palaeont. Institut Tübingen, but not found in 2003, most probably lost. Abundant material from Salzhausen stored at MNB (2004/821: Plate II, 1) and FSF (Plate III, 3).

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Symplocos jugata E. M. Reid (1920: 73, Pl. 4, Fig. 7), from Pont de Gail, France, upper Miocene, holotype (Plate III, 5) stored at BMNH, V25697. Symplocos microcarpa E. M. Reid (1920: 75–6, Pl. 4, Fig. 9), from Pont de Gail, France, upper Miocene, holotype stored at BMNH. Symplocos lusatica Mai (1964: 116, Pl. 16, Figs. 3–5; text—Fig. 9c) from Hartau near Zittau (Germany), lower Miocene. Holotype stored at MNB, 1993/1008a, see Plate III, 8. Symplocos durensis Mai (1970: 484, Pl. 67, Figs. 23–26) from Düren, NW Germany, upper Miocene, stored at MNB (No. 1993/698). Only one of the two syntypes is 4-locular (Plate I, 15), the other is 3locular (Plate I, 14) and falls within the variation range of the granulosa-morphotype. Thus, the 4locular condition is here reconsidered as an intraspecific variation, as observed in many modern species. Symplocos pliocaenica Van der Burgh (1978: 194–6, Pl. 5, Figs. 1–5) from Fortuna-Garsdorf I, lower Pliocene. Type material stored at the Laboratory of Palaeobotany and Palynology, Utrecht. Possibly, also Symplocos elongata Ludwig (1857: 99, Pl. 20, Figs. 7a–c), whose Pliocene type material from Weckesheim in Wettarau (Germany) is lost. In addition, we assign to Symplocos casparyi the specimens from Hartau (Miocene of Saxony, Germany) previously assigned to Symplocos anglica Chandler by Mai (1964: Pl. 16, Figs. 1–2), the specimen from the Oberpfalz assigned to Symplocos sp. 1 by Gregor (1980: Pl. 13, Fig. 23), the stone from Wieliczka (Poland) assigned to Symplocos minutula by Łańcucka-Środoniowa and Zastawniak (1997: Pl. 2, Figs. 9, 10), and the two specimens from Pocapaglia (Pliocene of northern Italy) assigned to S. minutula by Cavallo and Martinetto (1996: Pl. 5, Figs. 10, 11). We also revise as Symplocos casparyi the following records of Symplocos gothanii: – Düren in NW Germany, Miocene (stored at the Geol. Landesamt Nordrh. Westfalen, Krefeld: Thomson, 1958: 551, Pl. 2, Figs. 14, 15); – Kausche, Lusatia, NE Germany, Miocene (Kirchheimer, 1935a: Pl. 10, Fig. 29f); – Wiesa, Lusatia, NE Germany, Miocene (Kirchheimer, 1937: Fig. 25 and Kirchheimer, 1957: 318, Pl. 50, Figs. 189a–c); – Turow in Poland, Miocene (Czeczott and Skirgiełło, 1967: 159–160, Pl. 7, Figs. 11–13; Pl. 10, Fig. 5);

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– Stare Gliwice in W-Poland, Miocene (Szafer, 1961: 79, Pl. 21, Figs. 10–11); – Rypin near Dobrzyn (Wisła) in Poland, Miocene (Łańcucka-Środoniowa, 1957: 33, Pl. 5, Figs. 14–20>);

– Caʻ Viettone (Bertoldi and Martinetto, 1995: Fig. 4, n.1), Candelo and Castellengo (Martinetto, 1995, p. 80), Sento II (Martinetto, 1995: Pl. 2, Fig. 2; Basilici et al., 1997: Pl. 2, Fig. 7), NW Italy, Pliocene.

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Geographic and stratigraphic distribution: Symplocos casparyi, in the new, broad sense introduced in this work, occurs since the lower Oligocene and is widely distributed in the Miocene of Europe, from western and central France to Denmark and Poland (Stachurska et al., 1971; Günther and Gregor, 1990; Mai, 2000, 2001). Very similar distribution in the lower–middle Pliocene, but also southwards to central Italy (Martinetto, 2001), single place where the taxon persists in the upper Pliocene (Gelasian). Possibly also present along the East coast of the Black Sea (Chochieva, 1975). Description: The neotype (Plate III, 2) is an ovoid– ellipsoid stone of the sulcata-type, 5.6 mm long and 3.4 mm broad, with 3 equally developed pores around the central canal; surface with blunt, nearly straight, longitudinal ridges; abundant remains of a thin mesocarp; apically truncate, without bulge, margins thinedged; basally rounded with a small cavity; maximum breadth in the middle. Despite the fact that only two specimens are presently available (incl. the type of “Symplocos granulosa”) from the type-locality Dorheim, some neighbouring Pliocene (less probably upper Miocene, see Martinetto et al., 1997) sites, such as Sessenheim

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Fig. 3. Schematic cross-section of a stone of Symplocos braunii, after Kirchheimer (1936).

(Geissert et al., 1990), provided rich populations which include stones identical to the neotype and to the type of S. granulosa (Ludwig) Mai. Since the variability of such polymorphous populations agrees very well with those of Salzhausen, Wiesa, Hambach and Sento II, all of which include more than one hundred specimens, we decide to merge into a single, much variable species (for priority Symplocos casparyi Ludwig) all the globose–ovoidal to narrowly ovoid to ellipsoid, 3loculed forms with endocarps of the sulcata-type. Within the large populations of this species, one may

Plate I. 1.–5. 1. 2. 3. 4. 5. 6. 6a. 6b. 6c. 7. 7a. 8–15. 8. 8a. 8b. 9–13. 9. 10. 11. 12. 13. 14–15. 14. 15. 15a. 15b. 15c.

Symplocos arecaeformis (Schlotheim) Kirchheimer, stones from Plessa (1.–4.) and Gohra (5.), lower Miocene of Germany; after Kirchheimer (1935a: Pl. 10, Figs. 30a–e). ×1.5. Lateral view, holotype of “Symplocos sapperi” Kirchheimer 1934 (text—Fig. 18). Basal view. Apical view. Cross-section of the fruit from Plessa. Cross-section of the fruit from Gohra. Symplocos arecaeformis (Schlotheim) Kirchheimer, fragmentary stone from Annahütte, middle Miocene of Germany (original by Mai, 2001: Pl. 18, Fig. 8). MNB 1999/2117. ×1.7. Apical view. Internal view. Lateral view. Symplocos braunii Kirchheimer (1936: Pl. 10, Figs. 3a–b; type). Stone from Salzhausen, Miocene of Germany. ×4. Apical view.7b.. Lateral view. Symplocos caparyi Ludwig, stones. ×5. Holotype of “Myrica granulosa Ludwig”; Dorheim, Pliocene of Germany. After Mai (1973: Pl. 5, Figs. 1–3); MNB Berlin 964. Lateral view. Basal view. From Salzhausen, Miocene of Germany. MNB 2004/820. Cross-section. Apical view. Lateral view. Basal view. Holotype of “Carpinus salzhausenensis Ludwig” in two views. After Mai (1970: Pl. 67, Fig. 19); MNB 2004/819. From Düren, Miocene of Germany, “Symplocos durensis Mai”. After Mai (1970: Pl. 67, Fig. 23–24); MNB 1993/698. Apical view of 3-pored specimen, broken to the right of the apical pit to show seed locules. Holotype of Symplocos durensis Mai, with 4 apical pores. MNB 1993/698. Apical view, pit with 4 pores. Lateral view. Basal view.

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distinguish four main morphotypes (“pliocaenica”, “granulosa”, “lignitarum” and “jugata”, linked by intermediate forms), for which we provide the following description, with reference to the site of origin of the material described:

“pliocaenica”-morphotype (Fortuna-Garsdorf I, lower Pliocene, type locality of “Symplocos pliocaenica Van der Burgh”; 334 stones): endocarp ovoid or pear-shaped, mostly somewhat asymmetrical, crooked, variable in size, 3 to 7 (− 10) mm long, with fine,

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attenuated longitudinal ridges, sometimes curved and frequently forming a regular spiral when seen from the basal pole. Complete transition to smooth specimens. This morphotype is also very frequent in the assemblages of Adendorf (MNB), Boca (PU), Hambach (MNB), Sento (PU) and Wiesa (MNB), where smooth specimens were assigned by Mai (1964, Pl. 5, Figs. 11–13) to “Symplocos anglica Chandler”. “granulosa”-morphotype (a single stone is available from the type locality of “Symplocos granulosa (Ludwig) Mai”, Dorheim, thus variation is described based on abundant material from the Miocene of Wiesa in Germany (MNB), the ?Pliocene of Sessenheim in Alsace (Staatliches Museum für Naturkunde, Stuttgart) and the Pliocene of Sento II in Italy (PU): endocarp of the sulcata-type, globose to ovoidal, variable in size, but mostly 3.4–7.8 mm long. Frequent coarse or fine longitudinal ridges, mainly straight, but sometimes curved. Complete transition to almost smooth and totally smooth specimens. Apical pit of variable size, rounded or quadrangular, not more than 1/2 the diameter of the fruit. This morphotype is frequent in the upper Oligocene to upper Miocene of Germany, Denmark, Austria, Poland. Also in the Early (to Middle?) Pliocene of the Rhine valley and Italy. The apical pores are usually 3, but in a few localities rare 4pored stones occur (“Symplocos durensis Mai”). “lignitarum”-morphotype (Salzhausen, middle Miocene, type locality of “Symplocos lignitarum (Quenstedt) Kirchheimer; several hundred specimens): endocarp narrowly ovoid to ellipsoid, rather symmetrical, variable in size, 3 to 9.5 mm long, bearing strong and nearly straight, sharp longitudinal ridges (Plate III, 4), continuous from the apex to the base. The poor drawing in Quenstedt (1867) does not illustrate the diagnostic features of Symplocos endocarps, although it shows a narrowly ovoid specimen, bearing strong and nearly straight longitudinal ridges. The foregoing description is based on several analo-

gous specimens from Salzhausen, stored at MNB (Plate III, 4) and FSF (Plate III, 3). This morphotype occurs widely although it is less common than the pliocaenica- and granulosa-ones in the assemblages of Adendorf, Hambach and Wiesa (MNB). “jugata”-morphotype (Pont de Gail, France, upper Miocene, type locality of “Symplocos jugata” E. M. Reid; one specimen (Plate III, 5). Variability from Cava Toppetti I (Plate II, 8), middle or late Pliocene of central Italy, see Martinetto, 2001): endocarp ellipsoid to banana-shaped, frequently asymmetrical and curved, variable in size, 5 to 9 mm long (Plate II, 8), bearing straight longitudinal ridges which are both strong and continuous from the apex to the base, and fine, covering only 1/3–1/2 of the length. This morphotype is always very rare in the populations from central Europe, but it occurs widely in Italy, and it represents the dominant one in the middle and late Pliocene Symplocos stone assemblages of the Umbria region in central Italy, where a clear transition to the “lignitarum”-morphotype is observed (Plate III, 7), but no “pliocaenica”- or “granulosa”-morphotypes are present. For this reason such population (Plate II, 8) would be better regarded as belonging to a distinct variety of Symplocos casparyi. Similar modern species: Reid (1920) and Czeczott and Skirgiełło (1967) showed that, respectively, Symplocos glandulifera Brand and “Symplocos yunnanensis Brand” (= Symplocos sulcata Kurz, after Wu and Nooteboom, 1996) have shape and ornamentation very similar to the “jugata”-morphotype. Symplocos sulcata Kurz was formerly considered a variety of Symplocos macrophylla Wall. ex G. Don, which, after Nooteboom (1975) is a much polymorphous species as for fruitshape: “ovoid to ellipsoidal to cylindrical, 6–20 mm by 3–8 mm”. Distribution: India, Ceylon, Burma, Thailand, Indo-China and S China. Even more similar to the “jugata”-morphotype fossils are the fruits of Symplocos ophirensis Clarke:

Plate II. 1–8. 1. 2. 3. 4. 5. 6. 7. 8. 9.

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Symplocos casparyi Ludwig, populations of stones showing variation. “Symplocos lignitarum (Quenstedt) Kirchheimer” from Salzhausen, Miocene of Germany. MNB 2004/821. ×1.8. “Symplocos lusatica Mai” from Hartau, Miocene of Germany. MNB 1993/1008. ×1.2. “Symplocos salzhausenensis (Ludwig) Kirchheimer” from Salzhausen, Miocene of Germany. FSF, B12689. ×2. Other specimens assigned to “Symplocos salzhausenensis (Ludwig) Kirchheimer” from Salzhausen, MNB 2004/820. ×1.8. Globose to ellipsoid stones from Sento II, Pliocene of Italy. PU105115 ×1.7. From Hambach F7, Miocene of Germany. MNB1993/490. ×1.3. Globose to ellipsoid stones from Wiesa, Miocene of Germany. FSF, B12675 (coll. Kirchheimer). ×1.7. Stones of the “jugata”-morphotype from Cava Toppetti I, Pliocene of Italy. PU105116. ×1.5. Symplocos schereri Kirchheimer from Wiesa (Symplocos wiesaensis Kirchheimer), Miocene of Germany. MNB 1993/427a. ×1.5.

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“ampulliform, rarely ellipsoidal or cylindrical stones with coarse surface, few lengthwise ridges, ridges interrupted, locules 3”. Distributed in Sumatra, Malaya, Borneo, Celebes and Philippines.

Less similar is “Symplocos touranensis Guillaumin” (= Symplocos adenophylla Wall. ex G. Don var. touranensis (Guillaumin) Nooteboom, after Nooteboom, 1975).

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Symplocos foliosa Wight (material from mountain evergreen forests of Mt. Nilagiri, India) is very similar to the pliocaenica-morphotype, possessing an endocarp with rounded and slightly curved longitudinal ridges, 3 locules and thick walls (1 specimen observed). This differs, however, because the stone surface shows a diffuse fine granulation, similar to that observed in another modern species, Symplocos lucida (Thunb.) Sieb. and Zucc. Future research could indicate which character (longitudinal ridges, diffuse fine granulation) has better systematic value. Stones collected in China by Dr. C. Gee from a plant identified as “Symplocos mollifolia Dunn” (= Symplocos lancifolia Sieb. and Zucc., after Wu and Nooteboom, 1996) show an external morphology which is nearly identical to the granulosa-morphotype, however they are unilocular, with a single fertile seed, so they probably have a far phylogenetic relation to Symplocos casparyi. Similarly, Symplocos microphylla Wight and Symplocos paniculata (Thunb. ex Murray) Miq., indicated as modern relatives of Symplocos durensis Mai and Symplocos granulosa (Ludwig) Mai (Mai, 1970, 1973), have unilocular ampulliform stones, with a single curved fertile seed, so they differ consistently from the granulosa-morphotype. Trilocular stones are present in the Caribbean species of subgen. Symplocos (Mai, 1986) and in the Japanese sections Palaeosymplocos Brand and Okinawenses Nagamasu of subgen. Hopea (L.) Clarke (Nagamasu,

1993). However, we are not sure if they are truly of the “sulcata” type. Climatic indications: The similar modern species are distributed in the Asiatic Mixed Mesophytic and Broadleved Evergreen Forest regions or in the wet mountain forests (“cloud forests”) of the tropics of SE Asia and Middle America, additional research is needed to pinpoint the species which are closest to the fossil one. Symplocos germanica Mai 1970 Plate IV, 1–3 and Plate V, 1–11) Original: Symplocos germanica Mai (1970: 485–486, Pl. 68, Figs. 18–31, text—Fig. 15), from Brandis, NW Saxony, Germany (Floral zone IV) stored in MNB (No. 1993/3141). Synonyms: We revise as Symplocos germanica the specimen from Vetschau near Cottbus (Germany, lower Miocene, Floral Zone VII) assigned by Mai (1967: 214, Pl. II, Fig. 20) to Symplocos lakensis Chandler. Geographic and stratigraphic distribution: Lower Miocene in E Germany (Mai and Walther, 1991; Mai, 2000); upper Miocene in E and NW Germany, very common (Van der Burgh, 1987; Mai, 2001; Pingen, 2001). Description: Endocarps sometimes with a thin sticking mesocarp, ellipsoid to narrowly ovoid to obovoid, transversely wrinkled, of the germanica-type, no more than 2-locular, mostly only one locule developed (Plate V, 6,11). Apical pit 2/3 of the diameter of the fruit, deep and thin edged, always bearing two regular pores

Plate III. 1–8. 1. 1a. 1b. 1c. 2. 2a. 2b. 2c. 3. 3a. 3b. 3c. 4. 4a. 4b. 4c. 5. 6. 7. 7a. 7b. 7c. 8.

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Symplocos casparyi Ludwig, stones. ×7. From Hambach F7, MNB 1998/491b. Apical view. Lateral view. Basal view. From Dorheim, Pliocene of Germany. Neotype of Symplocos casparyi Ludwig, MNB 2004/825. Apical view. Lateral view. Basal view. “Symplocos lignitarum (Quenstedt) Kirchheimer” from Salzhausen, Miocene of Germany. FSF B12691/a. Apical view. Lateral view. Basal view. “Symplocos lignitarum (Quenstedt) Kirchheimer” from Salzhausen, Miocene of Germany. MNB 2004/821a. Apical view. Lateral view. Basal view. Holotype of “Symplocos jugata Reid” from Pont-de-Gail, Miocene of France. BMNH London, V25697. Stone of the “jugata”-morphotype from Cava Toppetti I, Pliocene of Italy. PU105117. Stone of the “lignitarum”-morphotype from Cava Toppetti I. PU105118. Lateral view. Apical view. Basal view. Holotype of “Symplocos lusatica Mai”, from Hartau, Miocene of Germany. MNB 1993/1008a.

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(Plate V, 3−5). Length 4.5–10.0 (6.6) mm; breadth 1.5–3.5 mm. Somewhat similar to Symplocos lakensis Chandler (1960a) from England, but more transversally wrinkled and in any case 1- or 2-locular.

Similar modern species: The most similar stones are produced by Symplocos filipes Nooteboom from Mindoro/Mt. Halcon, Philippines: stone spindle-shaped, with shallow longitudinal grooves, 1-loculed, 10 by 4 mm, with thin, fleshy mesocarp (Nooteboom, 1975, No. 42).

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Also some Caribbean species produce similar stones (Mai, 1986, pp. 3–4), such as Symplocos lanata Krug and Urb., Symplocos leonis Britt. and Wils. (sect. Urbaniocharis Brand) and Symplocos moaensis Borhidi (sect. Pseudosymplocos Brand). Climatic indications: Subtropical mountain climate, humid to semihumid. Symplocos gothanii Kirchheimer 1935a Plate IV, 6 and Plate VI, 1–7 Original: Symplocos gothanii Kirchheimer (1935a: 75– 76, Pl. 10, Fig. 20a–e, g), upper Miocene of Konzendorf near Düren, Lower Rhine Embayment, Germany. Preserved in MNB 2004/178(3245). Geographic and stratigraphic distribution: Upper Miocene in the Lower Rhine Embayment (Inden beds and allies), possibly endemic (Mai, 1970; Pingen, 2001. See under Symplocos granulosa a series of erroneous records. Description: Endocarps of the ciponimastrum-type, more or less longitudinally wrinkled, occasionally with single furrows, broadly obovoid to ellipsoid, 4– 5 mm long and 3.5–4 mm broad, with 3 or 4 unequally developed locules. Apical pit broader than half of the stone diameter, shallow and thick-edged. Similar in the ellipsoid form and wrinkled instead of ribbed surface to Symplocos pseudogregaria Kirchheimer (1938). Similar modern species: “Symplocos theaefolia D. Don” and “Symplocos tetragona Chen” (both = Symplocos lucida (Thunberg) Sieb. and Zucc., after Wu and Nooteboom, 1996). Climatic indications: Mixed mesophytic forests of China; MAT 11.1–16 °C, MAP 1000–1500 mm/a; frost free season 230 to 280 days, monthly mean temperature never below 0 °C, extreme minimum −14 °C. Symplocos minutula (Sternberg, 1825) Kirchheimer 1957 (Plate IV, 4,5 and Plate V, 12–22) Original: Carpolithus minutulus Sternberg (1825: 44, Pl. 53, Fig. 8) = Symplocos minutula (Sternberg, 1825)

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Kirchheimer (1957: 320, Pl. 50, Figs. 187a–b), from Salzhausen, Vogelsberg (Germany), middle Miocene. Kirchheimer (1957) reported that the type was not found in the Mus. of Nat. History of Prague, however topotypic material is stored in other collections, e.g. MNB (Plate IV, 5), Geol.-Palaeont. Institut Tübingen, FSF. Synonyms: Carpolithus gregarius Bronn (in Leonhard, 1840: 50 and 54, nomen nudum) = Symplocos gregaria (Bronn) A. Braun in Unger (1866: 31–32, Pl. 11, Figs. 1a–h, 2h and 2i), also from Salzhausen, topotypic material stored at MNB. Symplocos ludwigii Kirchheimer (1950: 16, Pl. 1, Fig. 8), from Dorheim near Friedberg, Wetterau (Germany), Pliocene. Stored at MNB (original no. 972). Pinus tumida Ludwig (1857: 88, Pl. 19, Figs. 3g–n) = Symplocos tumida (Ludwig) Kirchheimer (1936: 119, Pl. 13, Figs. 1a–i), also from Dorheim, Pliocene. Stored at MNB. If Symplocos elongata Ludwig (1857: 99, Pl. 20, Figs. 7a–c) does not represent Symplocos casparyi (see there), it would be referred to this species, as suggested by Kirchheimer (1957, p. 317). Geographic and stratigraphic distribution: Late Oligocene to middle Miocene of Germany, Poland, Switzerland, Austria (details in Mai, 1970, 2001). Upper Miocene in the Lower Rhine Embayment (Schürmann, 1961; Schultz, 1962; Van der Burgh, 1987). Pliocene of Wetterau (Germany), early Pliocene of northern Italy (Basilici et al., 1997, Pl. 2, Fig. 10). Description: Endocarps of the minutula-type, sometimes bearing remains of the thin mesocarp, with 1 to 2 fertile locules out of 3, finely striped to smooth, ellipsoid to ovoid, 6 to 11 mm long; breadth 3.5 to 5.5 mm. Apical pit 1/8–1/2 of the diameter of the stone, rounded to triangular, thin edged. Wall with cavities and crystal idioblasts. Similar modern species: Symplocos tinctoria L'Hérit., southeastern USA to Texas, Fagus–Magnolia forests (Knapp, 1965, p. 59), has an ovoid drupe, apically narrowed, 3-loculed, sometimes 1–2 locules aborted,

Plate IV. 1–3. 1. 2. 3. 4–5. 4. 5. 6.

Symplocos germanica Mai, populations of stones showing variation. ×1.8. From Hambach F7, Miocene of Germany. MNB 1993/494. From Hambach F7, Miocene of Germany. MNB 2001/157. From Brandis, Miocene of Germany, after Mai (1970: Pl. 68, Figs. 19–23), types. MNB 1993/3141. Symplocos minutula (Sternberg) Kirchheimer. ×1.8. Four stones from Dorheim, Pliocene of Germany. “Symplocos ludwigii” Kirchheimer (1936: Pl. 13, Figs. 1a–c). MNB 2004/826 (972), types. Population of stones from Salzhausen, Miocene of Germany, showing variation. MNB 2004/818. Symplocos gothanii Kirchheimer. Population of stones showing variation; from Hambach, Miocene of Germany, coll. F. v. d. Hocht, MNB 1998/491. ×1.8.

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surface nearly smooth to faintly striate, central canal pushed aside. Symplocos ramosissima Wallich ex G. Don (material from central Nepal) has endocarps with nearly identical

shape and ornamentation, but with thinner walls and with scarcely apparent sterile locules. The same is true for Symplocos glauca (Migo) from Honshu, with quite similar endocarps, mostly large as the largest Symplocos

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minutula, but again with poorly visible sterile locules and thinner walls. Climatic indications: Thompson et al. (2000) provide accurate data for the climatic preferences of Symplocos tinctoria L'Hérit.: 80% of its growing places have a mean annual temperature (MAT) comprised between 15.5 and 19.1 °C, the minimal MAT is 9.5 °C and the maximal is 22.1 °C. Mean January (coldest month) temperature: (0.2–) 4.7–11.3 (–15.9) °C; mean July temperature: (18.2–) 25.6–27.7 (–28.8) °C. Mean annual precipitation 1095–1630 mm; mean July precipitation: 80–214 mm. The data available for the similar Asian species are much less informative. Symplocos paucicostata (C. and E. M. Reid 1915) nov. comb. Plate VI, 16–19 and Plate VII, 1 Original: Karwinskia paucicostata C. and E. M. Reid (1915: 114, Pl. 12, Fig. 5). From Reuver, The Netherlands, middle Pliocene. Stored at the Geological Survey of The Netherlands (Utrecht). Synonyms: Symplocos aff. paniculata (Thunb. ex Murray) Miq. – Cavallo and Martinetto (2001: 317, Pl. 3, Figs. 4, 5).

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Geographic and stratigraphic distribution: Middle Pliocene of The Netherlands and ?late Pliocene of northern Italy (Cavallo and Martinetto, 2001). Description (based on the holotype and new specimens from Buronzo and Castelletto Cervo II, northern Italy): Endocarps of the Palura-type, dorsiventrally flattened, with an ovate outline, 4 to 9 mm long and 4 to 7 mm broad (L/W ratio from 1.6 to 1.0), mostly 1-locular, rarely 2-locular. Wall very thin (Plate VII, 1b), thus the fossils are usually strongly compressed during diagenesis. Apical pit 1/4 to 1/2 of the diameter of the stone, thin-edged. Small basal pit laterally shifted and crossed by a horseshoe-shaped peripheral furrow. Surface with fine anastomosing longitudinal striations. Similar modern species: The stones of living representatives of section Palura are very similar in all characters, even when different species are considered (Nagamasu, 1993), thus Symplocos paucicostata can definitely be assigned to this section. Climatic indications: Section Palura has the northern limit in northern Hokkaido (Japan), where the MAT is around 8 °C (Numata, 1974). Southwards it reaches central Vietnam (Nooteboom, 1998), probably in elevated areas, where the MAT can be estimated around

Plate V. 1–11. 1.–9. 1–3. 4. 5. 6. 7–9. 10–11. 10. 11. 12–22. 12–16. 12. 13a. 13b. 13c. 14. 15. 16. 17. 18. 19–22. 19. 20. 21a. 21b. 21c. 22.

Symplocos germanica Mai, stones, ×5. From Hambach F7, Miocene of Germany, Apical view of 3 specimens. MNB 1993/494a. Cross-section. Basal view. Cross-section. Three specimens in lateral view. From Brandis, Miocene of Germany, Holotype in lateral view. MNB 1993/3141. Other specimen in lateral view. Symplocos minutula (Sternberg) Kirchheimer, stones, ×5. From Salzhausen, Miocene of Germany, paratypes. MNB 2004/818. Apical view. Apical view. Lateral view. Basal view. Lateral view. Basal view. Cross-section. Longitudinally broken stone showing two locules. Cross-section. From Dorheim, Pliocene of Germany. Paratypes of “Symplocos tumida” Kirchheimer (1936: Pl. 13, Figs. 1a–g). MNB 2004/826 (972). Cross-section. Longitudinally broken stone showing a wide fertile locule and a thin sterile locule to the right. Apical view. Lateral view. Basal view. Cross-section.

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16–22 °C. In all its range the MAP is higher than 1000 mm. Symplocos pseudogregaria Kirchheimer 1938

Plate VI, 8–15 and Plate VII, 2–16 Original: Symplocos pseudogregaria Kirchheimer (1938: 354–355, Pl. 7, Figs. 1–2) from Niederpleis near Siegburg, lower Miocene of the Lower Rhine

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Embayment, stored in collection Schulte, Siegburg, not seen by the authors. Material (15 stones) from the type locality studied in 2003 at FSF. Synonyms: Symplocos kirstei Kirchheimer (1939b: 285–287, Pl. 3, Fig. 3) from Regis near Borna, Saxony, German lower Oligocene (“Haselbach beds”), stored in MMGD. Symplocos poppeana Kirchheimer (1941: 217–220, Figs. 12, 13) from Wiesa, stored at the MMGD. Formerly mentioned as Symplocos poppeana Kirchheimer (1940b: 32, without Fig., nomen nudum) from Wiesa near Kamenz, Saxony (Germany), lower Miocene, floral zone VI. Symplocos anglica Chandler (1957: 116, Pl. 17, Figs. 181–186) from the Oligocene of Bovey Tracey (England). Remarks: The Kirchheimer collection includes 15 stones from Niederpleis, the type locality of Symplocos pseudogregaria, which were labelled by Kirchheimer as “Symplocos sp.”. A few specimens, indeed, show the characteristic combination of diagnostic features indicated by Kirchheimer (1938) for this species. Nevertheless, a broad extent of variation is apparent, and the whole population does not differ significantly from the one of Symplocos kirstei from Borna (again Kirchheimer collec-

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tion, FSF). The remarkable apical bulge, which was considered a diagnostic character of Symplocos pseudogregaria by Kirchheimer (1957), is well developed only in 7 out of 15 specimens from Niederpleis and occurs in 4 out of 15 specimens from Borna. Also the absence of crystal idioblasts in the wall (Kirchheimer, 1939b) is a problematic and possibly an inconstant character. Additionally, in both localities, an apparent transition from urceolate to ellipsoid–ovoid to ellipsoid stones is observed. The latter ones are identical to the types of Symplocos poppeana from Wiesa (Kirchheimer, 1941), thus suggesting that this last species was defined simply by selecting ellipsoid specimens from a polymorphic population, also including urceolate to ellipsoid–ovoid stones assigned to Symplocos anglica by Mai (1970) and Mai and Walther (1991). Symplocos anglica was defined by Chandler (1957) based on a group of 34 stones from the Oligocene of Bovey Tracey (England). Even if such specimens are heavily compressed, they do not seem to differ significantly from the Niederpleis population cited above, suggesting that also the name S. anglica must be considered a younger synonym for Symplocos pseudogregaria. In conclusion, Symplocos pseudogregaria is reinterpreted here as a polymorphic species, showing a

Plate VI. 1–7. 1. 2. 3. 3a. 3b. 4. 5. 6. 7. 8–15. 8. 9. 10. 11. 12. 13. 14. 15. 16–19. 16. 17–19. 17. 18. 19a. 19b.

Symplocos gothanii Kirchheimer. Original material by Kirchheimer (1935a: Pl. 10, Figs. 29a–e), Konzendorf, Miocene of Germany. MNB 2004/178(3245). ×7. Apical view. Lateral view. Holotype by Kirchheimer (1935a: Pl. 10, Figs. 29a, 29c). Apical view. Lateral view. Apical view. Lateral view. Basal view. Cross-section. Symplocos pseudogregaria Kirchheimer. Stones from Phoenix–Nord, upper Eocene: “Symplocos kirstei Kirchheimer” by Mai and Walther (1985, p. 90). MNB 1993/10505a. ×7. Apical view. Basal view Lateral view. Cross-section. Apical view. Apical view. Lateral view. Lateral view. Symplocos paucicostata (C. Reid and E.M. Reid) n. comb. Stones. Holotype of “Karwinskia paucicostata” C. Reid and E.M. Reid (1915: Pl. 12, Fig. 5), fragmentary stone in lateral view from Reuver, Pliocene of the Netherlands. ×5. From Buronzo, ?late Pliocene of Italy. ×5. Apical view. sample A7Q, PU105119. Ventral lateral view. Sample A3M, PU105120. Ventral lateral view of 2-pored specimen. Sample A3M, PU105121. Apical view.

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range of stone shapes similar to that discussed for Symplocos casparyi. In this case, however, we provide a unified overview of the stone polymorphism, rather than a separate description of definite morphotypes.

Description: Endocarps at times bearing remains of a 1 mm-thick mesocarp, forming a stone of the ciponimastrum-type, globose, urceolate, ovoid, ellipsoid–ovoid to ellipsoid; nearly 5 to 11 mm long, 3 to 6 mm broad,

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mostly 3-locular, but 4- or 5-locular as well. Locules usually regularly developed. Surface mostly with fine anastomosing longitudinal wrinkles, otherwise longitudinally striate or bumpy, occasionally nearly smooth. Apex abruptly cut. Apical pit rounded, 9/10 to 1/3 of the diameter of the stone; thin to thick edged, at times surrounded by a remarkable bulge. Geographic and stratigraphic distribution: Upper Eocene to lower Oligocene in the Weisselster Basin, Germany (Mai and Walther, 1978, 1985). Lower Miocene (Floral-zone IV) of Leipnitz near Grimma, northwest Saxony (Mai and Walther, 1991: 89, Pl. 10, Fig. 19: stored at Museum für Mineralogie und Geologie Dresden, No. 1993/2422b) and Lusatia (Mai, 2000: 51, Pl. 14, Figs. 6–7). Lower to middle Miocene in other parts of Germany (Gregor, 1978, 1980; Mai, 2000, 2001; Pingen, 2001), Czech Republic (Holý, 1975) and Western Poland (Szafer, 1961). Lower Miocene of Oberdorf (identified as Symplocos cf. pseudogregaria and Symplocos poppeana by Meller, 1998, Pl. 16, Figs. 10–13) and middle Miocene of Weingraben in Austria (Jechorek and Kovar-Eder, 2004).

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Similar modern species: Symplocos anomala Brand: drupe oblong–globose–ellipsoid, 3-locular, sometimes 1–2 locules aborted, 7–10 by 4–6 mm, woody, faintly ribbed, mesocarp fleshy, thin. Widely distributed from Indonesia to Japan, in Yunnan 400 to 3000 m a.s.l., mixed forests (Nooteboom, 1975). Symplocos tingifera Chen (Hangchow, China) and Symplocos kuroki Nagamasu (=“Symplocos lucida Sieb and Zucc.”) from the Evergreen broad-leaved forests of southern Japan, which, after Nagamasu (1993), has 2-locular (rarely 3-locular) endocarps, while the modern material we observed (MNB) had 3-locular endocarps only. Symplocos foliosa Wight is less similar (see under Symplocos casparyi). Climatic indications: After Mai (2000, p. 167) climatic type Cfa/I, MAT 18–21 °C, CM 4–10 °C, WM 25– 28 °C, MAP 800–2000 mm. Symplocos pseudoschereri Mai 1970 Plate VIII, 1 Original: Symplocos pseudoschereri Mai (1970: 480– 481, Pl. 67, Figs. 1–2, text–Fig. 15c) from the borehole Dauban NSL 13/63 near Bautzen in Lusatia (Germany),

Plate VII. 1. 1a. 1b. 2.–16. 2. 3. 4–7. 4–5. 4a. 4b. 5a. 5b. 6–7. 6a. 6b. 7. 8. 9. 10. 10a. 10b. 10c. 11–14. 11. 12. 12a. 12b. 13. 14. 15. 16.

Symplocos paucicostata (C. Reid and E.M. Reid) n. comb. Stone from Buronzo, ?Late Pliocene, sample A3M, PU105122. Ventral lateral view. Lateral view, hole showing thin wall. Symplocos pseudogregaria Kirchheimer. Stones from different German sites. From Spremberg 37/60: Mai, 2000: Pl. 14, Fig. 6. MNB 1993/2047. ×5. From Leipnitz, lower Miocene: “Symplocos kirstei Kirchheimer” by Mai and Walther, 1991: Pl. 10, Fig. 19. MNB 1993/2422. ×5. “Symplocos poppeana Kirchheimer”. From Adendorf 2. MNB 2001/1029a. ×5. Apical view. Lateral view. Apical view. Lateral view. FromWiesa, lower Miocene. MNB 1993/415. Apical view. ×5. Lateral view. ×5. Variation of three stones. ×2. From Hartau, lower Miocene: Mai, 2000: Pl. 14, Fig. 20. MNB 1993/1017. ×5. From Spremberg 43/59: Mai, 2000: Pl. 14, Fig. 7. MNB 1993/2184. ×5. From Regis near Borna, lower Oligocene, type locality of “Symplocos kirstei” Kirchheimer (1939, p. 427). FSF B12740. ×5. Apical view. Lateral view. Basal view. “Symplocos sp.” in the Kirchheimer collection, FSF B12705. From Niederpleis near Siegburg, Miocene. Type locality of Symplocos pseudogregaria Kirchheimer (1938, p. 354). FSF B12705a. ×5. FSF B12705c. ×5. Lateral view. Basal view. FSF B12705b. ×5. FSF B12705. ×1.7. From Phoenix–Nord, upper Eocene: “Symplocos kirstei Kirchheimer” by Mai and Walther, 1985, p. 90. MNB 1993/10505a. ×1.7. From Nerchau, Miocene: “Symplocos anglica Chandler” by Mai, 1970, Pl. 68, figs. 9–12. MNB 1998/4963. ×3.

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lower Miocene (Floral Zone VI), stored at MNB (No. 1993/2311). Geographic and stratigraphic distribution: Only a few specimens at the type locality, endemic?

Description: Endocarp of the sulcata-type, finely ridged, straight–obovoid, 5 mm long and 4 mm broad, with 3 equally developed locules. Longitudinal ribs straight, crest-shaped, irregular and sometimes broken.

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Apical pit 2/3 of the diameter of the stone, deep and thin-edged. Similar to Symplocos schereri Kirchheimer, but remarkably smaller and with different ridges. Similar modern species: Unknown. Symplocos schereri Kirchheimer 1935b Plate II, 9 and Plate VIII, 2–7 Original: Symplocos schereri Kirchheimer (1935b: 291, Fig. 8), from Konzendorf near Düren Germany), Miocene. Stored at MNB (No. 2004/179a). Synonyms: Symplocos wiesaensis Kirchheimer (1940a: 288–290, Fig. 5), from Wiesa near Kamenz, Saxony (Germany), lower Miocene, floral zone VI, stored at MMGD. Geographic and stratigraphic distribution: Symplocos schereri (incl. Symplocos wiesaensis) is common in the lower–middle Miocene (most of the German basins: Kirchheimer, 1940a; Mai, 1964, 1970; Gregor, 1978, 1980; Mai, 2000, 2001. Bohemia: Kvaček and Holý, 1977. Poland: Czeczott and Skirgiełło, 1967. Austria: “Symplocos cf. schereri Kirchheimer” by Meller, 1998: Pl. 16, Figs. 8, 9). It also occurs in the upper Miocene of the Lower Rhine Embayment (Schultz, 1962; Mai, 2000) and Lower Silesia (Łańcucka-Środoniowa and Zastawniak, 1993), in the lower Pliocene (NW Germany, N Italy: Cavallo and

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Martinetto, 1996; Basilici et al., 1997), and even the middle Pliocene in central Italy (Martinetto, 2001). Description (Wiesa): Endocarps of the sulcata-type, straight–obovoid, 7 to 15 mm long and 9 mm broad, with thin, wing-shaped, irregularly developed longitudinal ridges, commonly 3-locular, but mostly one locule aborted, often tripointed at the base. Wall thin, with a few cavities only. Apical pit 2/3 diameter of the stone, deep, thinly edged. Similar modern species: Symplocos tanakae Matsamura (based on endocarp description by Nagamasu, 1993, Fig. 4b). Southern Japan, warm-temperate montane evergreen forests. Possibly similar (after Nooteboom, 1975): Symplocos costata (Blume) Choisy: endocarp ribbed, with 8 high ridges. West Java, 900–2000 m. Symplocos crassilimba Merr. (Nooteboom, 1975, Pl. 8, Fig. d): endocarp ribbed, with 7 to 8 ridges. Philippines. Similar in cross-section, but with definitely larger fruits, are the following species: “Symplocos barringtoniifolia Brand” (= Symplocos rigida Clarke, after Nooteboom, 1975, Pl. 4, Fig. e): stone woody, with 8 very high ridges, ovoid to ellipsoid, 25–40 by 12– 20 mm, mesocarp chartaceous. Indochina, Malay Peninsula, Borneo. Symplocos cerasifolia Wallich (Nooteboom, 1975, Pl. 7, Fig. f): endocarp woody,

Plate VIII. 1. 1a. 1b. 1c. 2–5. 2. 3a. 3b. 4. 5. 6–7. 6. 7. 7a. 7b. 7c. 7d. 8–16. 8. 9–16. 9. 10. 11–14. 15. 16.

Symplocos pseudoschereri Mai, holotype. Stone from Dauban, Miocene of Germany, after Mai (1970, Pl. 67, fig.1). MNB 1993/2311. ×6. Lateral view. Apical view. Basal view. Symplocos schereri Kirchheimer. Stones from Wiesa (“Symplocos wiesaensis” Kirchheimer), Miocene of Germany. MNB 1993/427. ×5. Apical view. Lateral view. Basal view. Cross-section Lateral view. Symplocos schereri Kirchheimer stones from Konzendorf, Miocene of Germany. ×5. Apical view of the paratype. MNB 2004/179. Holotype, MNB 2004/179a. Cross-section. Lateral view. Apical view. Basal view. Symplocos tetraporina Mai, stones. From Klettwitz, Miocene of Germany, lateral view. MNB 1993/930. ×6.5. From Puschwitz, Miocene of Germany. MNB 1993/250. Apical view. ×6.5. Holotype after Mai (1970: Pl. 67, Fig. 3), lateral view. MNB 1993/250a. ×6.5. Specimens in lateral view. ×4. Cross-section. ×8. Basal view. ×6.5.

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with 8 high ridges, ellipsoid, 3-loculed with central canal, often only 1 locule developed, 22–40 by 8–18 mm. Thailand, Malaya, Sumatra, Borneo, New Guinea. Climatic indications: The similar modern species prefers warm temperate to subtropical humid conditions (Cfa or Cwa-type). Symplocos tetraporina Mai 1970 Plate VIII, 8–16 Original: Symplocos tetraporina Mai (1964: 65, nomen nudum), from Merka near Bautzen, lower Miocene (Floral Zone VI); Symplocos tetraporina Mai (1970: 482–483, Pl. 67, Figs. 3–4, text–Fig. 15e) from Puschwitz near Bautzen, lower Miocene (Floral Zone VI), stored at MNB (no. 1993/250). Geographic and stratigraphic distribution: Lower and middle Miocene in the Lusatia region of Germany (Mai 1964, 1970, 2000, 2001). Description: Endocarp of the sulcata-type, narrowly ellipsoid, 5 to 8 (most commonly 6) mm long and 2.4–3 (most commonly 2.8) mm broad, with 11 to 20 unevenly high and broad, often anastomosed, longitudinal ridges. Four locules, always regularly developed. Apical pit nearly as broad as the diameter, very shallow, rounded to quadrangular. Although resembling the lignitarum-morphotype of Symplocos casparyi, Symplocos tetraporina is always tetralocular and slender, with thinner and sharper ridges. Similar modern species: Very similar to Symplocos sulcata Kurz, which thrives in mixed forests and evergreen broad-leaved forests in southern China, Burma and, above 1500 m, in Thailand and Indo-China. Climatic indications: The similar modern species prefers warm temperate to subtropical humid conditions (Cfa or Cwa-type). 4. Doubtful species Symplocos chvarbetica Chochieva 1975 Original: Symplocos chvarbetica Chochieva (1975: 62, Pl. 15, Figs. 7a–b). From Khvarbeti/Guria, Plio/ Pleistocene. Discussion: Original material not seen, probably a Symplocos casparyi, lignitarum-morphotype. Symplocos urceolata E. M. Reid 1920 Original: Symplocos urceolata E. M. Reid (1920: 74– 75, Pl. 4, Figs. 8a, b – same specimen) from Pont-deGail, central France, late Miocene (formerly early Pliocene), stored at BMNH (no. V25 592). Discussion: The single endocarp from Pont-de-Gail, recently observed by the authors, does not permit us

to confirm that Symplocos urceolata is a distinct species. Most probably, it is an anomalous specimen of Symplocos casparyi, found in the same assemblage (sub Symplocos jugata and Symplocos microcarpa). 5. Conclusions The re-examination of abundant fossil material allowed us to reduce the number of Symplocos species represented in the European Neogene to 11 (Table 1). Despite the difficult delimitation of Symplocos species, even among living populations (Nooteboom, 1998), we suggest that the species concept we applied is such that the forgoing number of fossil species roughly corresponds to the number of ancient botanical species which grew in the lowlands of Europe from 30 to 2 My ago. This is, of course, a minimal number, because additional species could have grown in elevated areas, away from any sedimentary basin, thus leaving no fossil record. The taxonomic revision and the rough separation of European Neogene species into several morphological type that are also recognized within living species, permit us to limit the search for the nearest relatives to restricted groups of species. At present, it is possible to provide a first approximation of the modern distribution of such fruit-types in America and Asia. Future investigations will probably enhance the precision of fossil–modern relationships, with a consequent refinement of the climatic indications which they can provide for the European Neogene. Our reassessment confirms that the diversity of Symplocos in Europe was maximal in the early and middle Miocene, characterized by the occurrence of 7 common species (Symplocos arecaeformis, Symplocos casparyi, Symplocos germanica, Symplocos minutula, Symplocos pseudogregaria, Symplocos schereri, Symplocos tetraporina) and 2 rare ones (Symplocos braunii, Symplocos pseudoschereri). The diversity started to decrease in the late Miocene (possible extinction of S. arecaeformis, S. pseudogregaria and S. tetraporina), despite the local appearance of Symplocos gothanii (Lower Rhine Embayment), and fell down sharply in the early Pliocene. Only two species (Symplocos casparyi and Symplocos paucicostata) remained in central Europe after that time. Until recently, the genus Symplocos was thought to disappear from Europe in the final part of the middle Pliocene (“Reuverian”), before the onset of glacial– interglacial cycles. However, recent findings in central Italy prove that the genus survived there with one species (Symplocos casparyi) until the late Pliocene (Martinetto, 2001). At the same time in northern Italy a

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Table 1 Main fruit characters of the Symplocos species from the European Neogene Species

Mesocarp sticking

Locules Locule Endocarp Shape development length (mm)

Endocarp ornament

S. arecaeformis

Yes, thick

5

1 usually abortive

Non-visible

S. braunii

Yes

3

Equal

9–12

Obovoid

S. pseudoschereri No

3

Equal

5

Obovoid

S. schereri

No

3

1 usually abortive

7–15

Obovoid

S. casparyi s. l.

No

3 (4)

Equal

3–10

S. gothanii

No

3, 4

Unequal

4–5

Globose, ovoid, narrowly ovoid, ellipsoid, pearand bananashaped Broadly obovoid to ellipsoid Urceolate, ovoid to ellipsoid

25–45

Broadly obovoid

1/3 diameter, shallow, round-edged Non-visible 1/3 diameter, deep, thick-edged Fine crest2/3 diameter, shaped deep, thinridges edged Winged ridges 2/3 diameter, deep, thick-edged

Coarse or fine Rounded or ridges, rarely quadrangular, smooth variable size, not more than 1/2 diameter Longitudinally Broader than 1/2 wrinkled diameter Longitudinally Rounded, 9/10 to 1/ wrinkled, 3 diameter, thin to striate or thick edged, at bumpy, rarely times remarkable smooth bulge Rounded to 11 to 20 quadrangular, as unevenly broad as diameter, high and very shallow broad, often anastomosed, longitudinal ridges Transversely 2/3 diameter wrinkled

S. pseudogregaria Rarely, 1 3 (4–5) Equal mm thick

5–11

S. tetraporina

No

5–8

S. germanica

Sometimes, 2 thin

Usually only 4.5–10 1 developed

Ellipsoid, narrowly obovoid or ovoid

S. minutula

Sometimes, 3 thin

1 to 2 fertile

6–11

Ellipsoid to ovoid

Finely striped to smooth

S. paucicostata

No

Unknown

4–9

Bilaterally flattened, ovate

Fine anastomosing longitudinal striations

4

1 (2)

Equal

species (Symplocos paucicostata) of Symplocos section Palura, which today includes one or a few deciduous, rather cool-tolerant species, survived until the end of the Pliocene (Cavallo and Martinetto, 2001). It was one of several plant genera (Carya, Magnolia, Phellodendron, etc.) which were eliminated by the following climatic deteriorations that brought cool and dry conditions to southernmost Europe, and thus hampered the survival of plants which require a humid temperate climate.

Ellipsoid

Apical pit

Rounded to triangular, 1/8–1/2 diameter, thin-edged 1/4–1/2 diameter, thin-edged

Stratigraphic distribution Lower and middle Miocene ?Middle Miocene Lower Miocene Lower Miocene to middle Pliocene Upper Oligocene to upper Pliocene Upper Miocene Upper Eocene to middle Miocene Lower and middle Miocene

Lower and upper Miocene Upper Oligocene to Pliocene Pliocene

6. Key (A) Fruit with solid mesocarp sticking at the stone (Fig. 3) (1) Fruit stone 5-locular, unusually large, up to 45 mm long. Symplocos arecaeformis (2) Stones 3-locular, only up to 12 mm long. Symplocos braunii (B) Fruit with a leathery or fleshy mesocarp, loosely attached to the stone and usually not preserved

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(a) Stone with more or less winged ridges (1) thinly crest-shaped longitudinal ridges, 5 mm long. Symplocos pseudoschereri (2) Stone with thicker, irregularly developed, ridges, stone up to 15 mm long and 9 mm broad. Symplocos schereri (b) Stone with low longitudinal ridges (1) Stone narrowly ellipsoid or ovoid. Locules 3 (rarely 4), mostly all fertile, apical pit medium deep and 1/3 to 1/1 of the diameter. Symplocos casparyi (2) Stone narrowly ellipsoid. Locules 4 equally developed. Apical pit shallow, as broad as the stone. Symplocos tetraporina (3) Stone globose, with frequent coarse or fine longitudinal ridges, mainly straight, but sometimes curved. Complete transition to almost smooth and totally smooth specimens. Symplocos casparyi (granulosa-morphotype) (c) Stone more or less longitudinally rugose, sometimes with isolated furrows. (1) Stone broadly ovoid to ellipsoid, 4 to 5 mm long, 3.5 to 4 mm broad, with 3 or 4 inequally developped locules; apical pit broader than 1/2 of the stone diameter. Symplocos gothanii (2) Stone urceolate, ovoid to ellipsoid, 5 to 11 mm long and 3 to 6 mm broad, mostly 3-locular but as well 4 to 5-locular. Locules usually regularly developed, apical pit broad, often surrounded by a bulge. Symplocos pseudogregaria (3) Stone ellipsoid, narrowly ovoid to obovoid, transversely wrinkled, 2- or 1locular, apical pit deep with thin edges. Symplocos germanica (d) Stone nearly smooth or longitudinally striate, with 1 or 2 fertile locules. (1) stone relatively thick walled, often bearing remains of the thin mesocarp, ellipsoid, length 6 to 11 mm; breadth 3.5 to 5.5 mm. Fertile locules one or two. Wall with cavities. Apical pit rounded to triangular, thin edged. Symplocos minutula (2) stone thin walled, bilaterally flattened; basal pit laterally shifted and crossed by a horseshoe-shaped peripheral furrow; fertile locule one, rarely two. Symplocos paucicostata.

Acknowledgements We wish to thank Dr. Peter W. Fritsch for the careful revision and improvement of the preliminary text and useful information on the phylogeny of Symplocos. We are grateful to Dr. Hans P. Nooteboom for resignments on the taxonomy of modern Symplocos in Asia. We also thank Dr. Volker Wilde for help in the Forschunginstitut Senckenberg collection, and for providing up-to-date information on fossils from Wetterau. Dr. Hidetoshi Nagamasu sent several fruit samples of sect. Palura from Japan and Dr. Carole Gee gave us samples of Symplocos fruits recently collected in China. The reviewers, Dr. Steven R. Manchester and Dr. Bruce Tiffney, suggested important improvements and linguistic corrections. References Anderberg, A.-L., 1994. Atlas of Seeds: Part 4. Resedaceae– Umbelliferae. Swedish Museum Nat. Hist., Stockholm. 281 pp. Basilici, G., Martinetto, E., Pavia, G., Violanti, D., 1997. Paleoenvironmental evolution in the Pliocene marine–coastal succession of Val Chiusella (Ivrea, NW Italy). Boll. Soc. Paleont. It. 36 (1–2), 23–52. Bertoldi, R., Martinetto, E., 1995. Ricerche paleobotaniche (palinologiche e paleocarpologiche) sulla successione “villafranchiana” del Rio Ca' Viettone. Il Quaternario 8 (2), 403–422. Brand, A., 1901. In: Engler, A. (Ed.), Symplocaceae. Planzenreich IV, vol. 242, pp. 1–100. Brongniart, A., 1828. Prodrome D'une Histoire Des Végétaux Fossiles. Levrault, Paris. 223 pp. Cavallo, O., Martinetto, E., 1996. Flore plioceniche del bacino del Tanaro. Alba Pompeia 17 (1), 5–31. Cavallo, P., Martinetto, E., 2001. Flore carpologiche del Pliocene di Castelletto Cervo (Biella). Boll. Mus. Reg. Sci. Nat. Torino 18 (2), 277–343. Chandler, M.E.J., 1926. The upper Eocene flora of Hordle, Hants II. Monogr. Palaeont. Soc. London 78, 33–52. Chandler, M.E.J., 1957. The Oligocene flora of the Bovey Tracey Lake Basin, Devonshire. Bull. Br. Mus., Nat. Hist., Hist. Ser. 3 (3), 71–123. Chandler, M.E.J., 1960a. The Lower Tertiary Floras of Southern England: I. Palaeocene Floras – London Clay Flora (Supplement). Brit. Mus. Natur. Hist., London. 357 pp. Chandler, M.E.J., 1960b. Plant remains of the Hengistbury and Barton Beds. Bull. Brit. Mus. Nat. Hist., Geol. 4 (6), 193–238. Chandler, M.E.J., 1961. Flora of the Lower Headon Beds of Hampshire and the Isle of Wight. Bull. Brit. Mus. Nat. Hist., Geol. 5 (5), 93–157. Chandler, M.E.J., 1962. The Lower Tertiary Floras of Southern England: II. Flora of the Pipe-Clay Series of Dorset (Lower Bagshot). Brit. Mus. Natur. Hist., London. 176 pp. Chandler, M.E.J., 1963. The Lower Tertiary Floras of Southern England: III. Flora of the Bournemouth Beds; the Boscombe, and the Highcliff Sands. Brit. Mus. Natur. Hist., London. 169 pp. Chochieva, K.I., 1975. Chvarbetskij iskopaemyi chvoinyj lec. Mecniereba, Tbilisi. 184 pp. Czeczott, H., Skirgiełło, A., 1967. The fossil flora of Turow near Bogatynia III. Prace Muz. Ziemi 10, 97–166.

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