Acta Palaeobotanica 47(2): 419–424, 2007 Trapa kvacekii (Trapaceae), a remarkable new fossil species from the late Miocene of Greece JAN J. WÓJCICKI 1 and DIMITRIOS VELITZELOS 2 1 W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Kraków, Poland; e-mail: jan.wojcicki@ib-pan.krakow.pl 2 National and Kapodistrian University of Athens, Faculty of Geology and Geoenvironment, Panepistimioupolis Zografou, Athens, Greece; e-mail: veljim@otenet.gr Received 16 August 2006; accepted for publication 16 July 2007 ABSTRACT. Trapa kvacekii Wójcicki & D. Velitzelos (Trapaceae), a new fossil species from the late Miocene of Likudi near Elassone (Thessalia, Greece) is described, illustrated and briefly discussed. It differs markedly from its congeners by fruit morphology, primarily by the characteristic fruit body oblong-triangular in outline, well-developed high ring at the base, and stout lower horns inserted 3/5 to 3/4 the distance from the base of the fruit. KEY WORDS: Trapa, Trapaceae, new species, fruit morphology, late Miocene, Greece INTRODUCTION The late Miocene diatomites in Likudi near Elassona (Thessalia, Greece) are known from rich and well-preserved leaf flora (Velitzelos & Gregor 1985, 1986, 1990, Knobloch & Velitzelos 1986, 1987, Mai 1995, Velitzelos et al. 2000). Their unique character is manifested by four new species of trees – Quercus dubia Knobloch & Velitzelos, Q. likudensis Knobloch & Velitzelos, Ostrya likudensis Knobloch & Velitzelos, and Salix massalongii Knobloch & Velitzelos, restricted to this locality (Knobloch & Velitzelos 1987) – and one legume Gymnocladocarpum velitzelosii Gregor of the subfamily Caesalpinioideae, probably ancestral to SE North American Gymnocladus dioicus (L.) K. Koch (Gregor 1986). Recent collections of the second author yielded new material from Likudi (Fig. 1), including several relatively well-preserved impressions of Trapa fruits of interesting morphology, which represent a new fossil species described here. Fig. 1. Geographic location of the site of Trapa kvacekii Wójcicki & D. Velitzelos sp. nov. at Likudi near Elassona, Greece 420 MATERIAL AND METHODS Nine Trapa specimens exist as impressions in soft whitish or yellowish finely bedded diatomite, sometimes with remnants of oxidised and/or carbonised endocarp. They were collected from an erosion-exposed slope in a valley between Likudi and Kleisura, ca. 10 km NW of Elassona (Fig. 1) marked on a schematic map provided by Velitzelos and Gregor (1986, Fig. 3) as “Likudi 5”. Trapa fossils are reported from this place as Trapa sp. by Velitzelos and Gregor (1990), but they have not been described so far. The age of the locality is not precise, but according to Knobloch and Velitzelos (1987) it is late Miocene, younger than late Badenian (see also Kovar-Eder et al. 2006). Most fruit impressions from Likudi are relatively well preserved (Figs 2, 3) and they bear characteristics that permit more precise interpretation of their morphology. Six of them are impressions of almost complete fruit compressions in the plane of the upper horns (Figs 2A–C, 3A–E, G). Two other fruits were positioned differently in the sediment, one in the plane of the lower horns (Fig. 3F) and the other obliquely in the plane of the upper horns (Figs 2D, 3H), enabling confirmation of the presence of well-developed lower horns, and their characterisation. To stabilise the morphology of the Trapa fruit impressions and to prevent degradation and removal of organic matter, the samples were coated with arti- ficial resin (Poraloid dissolved in acetone) after preparation of the specimens. Photographic documentation was made with a standard Nikon Coolpix 995 digital camera. The contrast was enhanced by low-angle light. The material described here is housed in the palaeobotanical collection of the Department of Historical Geology and Palaeontology, Athens University, Athens (DHGPA; eight specimens), and one specimen, kindly donated by the second author, is stored in the Palaeobotanical Museum of the W. Szafer Institute of Botany, Polish Academy of Sciences, Kraków (KRAM-P). Comparative fossil plant material was studied by the first author in the palaeobotanical collections of Barcelona (Departament d’Estratigrafia, Paleontologia i Geociències Marines, Universitat de Barcelona), Beijing (Institute of Botany, Chinese Academy of Sciences), Berlin (Museum für Naturkunde), Brno (The Moravian Museum), Bucharest (Department of Geology and Palaeontology, University of Bucharest), Budapest (Hungarian Natural History Museum and Geological Institute of Hungary), Cottbus (Museum der Natur und Umwelt), Dresden (Staatliche Sammlungen, Museum für Mineralogie und Geologie), Frankfurt (Palaeobotanical Section of the Forschungsinstitut Senckenberg), Kraków (W. Szafer Institute of Botany, Polish Academy of Sciences), London (Palaeontological Collection of the British Museum – Natural History), Munich (Bayer- Fig. 2. Trapa kvacekii Wójcicki & D. Velitzelos sp. nov. from Likudi near Elassona. A – No. DHGPA L/Tr/1 (= Fig. 3A), B – DHGPA L/Tr/5 (= Fig. 3D), C – DHGPA L/Tr/2 (= Fig. 3B), D – DHGPA L/Tr/9 (= Fig. 3H); A – holotype, B, C – paratype. a – apical aperture, bh – base of lower horn, h – head, lh – lower horn, n – neck, r – ring, ri – ribs, s – scar, t – trace of tubercle, uh – upper horn; scale bar 1 cm 421 ische Staatssammlung für Paläontologie und Geologie in Munich), Nanjing (Institute of Geology and Palaeontology, Chinese Academy of Sciences), Stockholm (Swedish Museum of Natural History), St. Petersburg (Komarov Botanical Institute, Russian Academy of Sciences), Turin (Museo di Geologia e Paleontologia of the Turin University) and Vienna (Geological-Palaeontological Department of the Natural History Museum). Extant plant material was studied in or obtained on loan from the following herbaria: B, BM, BP, CAL, FI, KRAM, KW, LE, LY, MW, MHA, P, PE, PR, PRC, S, W, WU, Z (acronyms follow Holmgren et al. 1990). Fig. 3. Trapa kvacekii Wójcicki & D. Velitzelos sp. nov. from Likudi near Elassona. A – No. DHGPA L/Tr/1 (= Fig. 2A), B – DHGPA L/Tr/2 (= Fig. 2B), C – KRAM-P 249, D – DHGPA L/Tr/5 (= Fig. 2B), E – DHGPA L/Tr/6, F – DHGPA L/Tr/7, G – DHGPA L/Tr/8, H – DHGPA L/Tr/9 (= Fig. 2D), I – neck of H with visible upward-pointing hairs closing apical aperture; A – holotype, B, C, D – paratypes. a – apical aperture, bh – base of lower horn, f – frame, h – head, lh – lower horn, n – neck, r – ring, ri – ribs, s – scar, t – trace of tubercle, uh – upper horn; scale bars 1 cm 422 SYSTEMATICS Trapaceae Doum. nom. cons. Trapa kvacekii Wójcicki & D. Velitzelos sp. nov. Figs 2A–D, 3A–I H o l o t y p e (designated here). Coll. file No. DHGPA L/Tr/1 (Figs 2A & 3A). P a r a t y p e s. Coll. file Nos DHGPA L/Tr/2 (Figs 2C & 3B), L/Tr /5 (Figs 2B & 3D); KRAMP 249 (Fig. 3C). F u r t h e r m a t e r i a l. Coll. file Nos DHGPA L/Tr/3, L/Tr/6 (Fig. 3E), L/Tr/7 (Fig. 3F), L/Tr/8 (Fig. 3G), L/Tr/9 (Figs 2D & 3H). T y p e l o c a l i t y. Erosion exposed slope in a valley between Likudi and Kleisura near Elassona, Thessalia, Greece (ALikudi 5@ according to Velitzelos & Gregor 1986). T y p e h o r i z o n. Yellowish diatomite. A g e. Late Miocene. D e r i v a t i o n o f t h e n a m e. Named after the eminent Czech palaeobotanist Professor Zlatko Kvaček in recognition of his contribution to Tertiary palaeobotany. D i a g n o s i s a n d d e s c r i p t i o n. Large fruits, oblong-triangular in outline, with two pairs of solid horns; fruit 31–41 mm high (including neck), width of fruit at upper horns 40–55 mm; fruit about 1.3 times as wide as high; fruit head pronounced, 8–10 mm long, its upper end situated below the line joining the bases of the upper horns, bearing a welldeveloped neck usually somewhat narrowing towards the apex; neck 3–4 mm long and 3.5–5.0 mm broad, slightly protruding beyond the line joining the bases of the upper horns, corona lacking; apical aperture with a ring of upward-pointing hairs; surface of fruit head and neck finely ribbed; upper horns triangular in outline, (14) 16–22 mm long, at least slightly raised at the base, gradually attenuating into straight, elongate, thin, spine-like tips, ascending (50°–65°), with a smooth surface except for poorly marked, at least 6 mm long, retrorsely barbed spines (harpoons); presence of mat areas excluded; lower horns narrowly triangular in outline, up to 14 mm long, and about 8 mm wide at the base, at least slightly retrorse, straight or with gently upward-pointing apical part, inserted usually 3/5 to 3/4 the distance from the base of the fruit; the frame of the fruit well developed; on the fruit frame between the bases of the upper and lower horns, solid, probably conical (truncate?) tubercles developed, traces of their bases up to 4 mm in diameter; lower part of the fruit body obtriangular in outline, truncate at the base, its surface (on one side only) covered with five protruding longitudinal ribs; fruit base with a smooth ring, up to 3 mm high; basal scar probably up to 3 mm in diameter. DISCUSSION Trapa kvacekii is a new well-defined fossil species of the late Miocene of Europe with a unique combination of characters never reported previously (see Wójcicki 2002, Wójcicki & Zastawniak 2003 and Kovar-Eder et al. 2005 for references). It is characterized by having fruits with the fruit body oblong-triangular in outline, solid, relatively long lower horns inserted 3/5 to 3/4 the distance from its base and a well developed high ring at the base (Figs 2, 3). In addition, Trapa kvacekii possesses the largest fruit body of all Tertiary fossil Trapa described so far. The fruits from Likudi seem not to vary much, and the variation observed is probably due to fossilisation and the position of the fruit in the sediment (Fig. 3). From the Balkan Peninsula there are several late Neogene localities with fossil Trapa remains reported. One is the Kreka Basin near Tuzla in Bosnia of the late Miocene. On the basis of single specimens, Janković and Pantić (1953; see also Janković 1958) described four separate fossil species from this locality: Trapa bosniaca Janković & Pantić, T. tuzlensis Janković & Pantić, T. pontica Janković & Pantić, and T. praemuzzanensis Janković & Pantić. From the protologues and figures it is evident that T. bosniaca and T. tuzlensis differ in fruit shape and size from the species newly described from Likudi. Unlike T. kvacekii, their fruits are broadly rounded at the base, with a pair of relatively short upward-pointing upper horns and reduced lower horns inserted probably 1/3 the distance from the base of the fruit. It is possible that T. bosniaca and T. tuzlensis represent the same fossil species. The original descriptions of 423 the other two species, T. pontica and T. praemuzzanensis, are based on poorly preserved, incomplete specimens showing some similarity to T. silesiaca Goeppert emend. Wójcicki & Zastawniak, widely distributed in Europe in the late Miocene (Wójcicki & Zastawniak 2002, Kovar-Eder et al. 2005). The identity of the fruits from Kreka remains unsolved, requiring detailed reinvestigation of the original material. Other localities with Trapa remains from the Balkan region are of Pliocene age, reported from south-western Romania (Dacic Basin) by Givulescu and Ţicleanu (1986) and Ţicleanu (1995, 2003). The characteristic morphology of Trapa fruits from these localities (e.g., stalked and/or abruptly elongated fruit base) suggests that they are evidently of separate evolutionary lineage (Wójcicki & Ţicleanu, in preparation). The fruits of T. kvacekii show some similarity to those from the Pliocene of Arboschio (Villafranca d’Asti, Italy), originally classified by Pavia (1970: tab. 4, fig. 1) as T. natans L., but they differ in some aspects. Like T. kvacekii, the specimens from Arboschio are obtriangular and somewhat elongate in outline, with solid lower horns, relatively big tubercles on the fruit frame between the bases of the upper and lower horns, and a well-developed smooth basal ring up to 2 mm high. The difference of the specimens from Italy is best marked by their smaller size (about 25 mm high and 32 mm wide at upper horns), the position of the lower horns (inserted approximately in the centre of the fruit body), and the presence of a corona on the neck, a character unknown in representatives of the Trapa genus described from the Miocene. There are at least a few unnamed modern morphotypes scattered in the southern part of Europe, placed in the still poorly recognized T. natans complex, with a visibly elongated fruit body obtriangular in outline, relatively stout lower horns located above the centre of the fruit, bearing at least small tubercles on the fruit frame between the lower and upper horns, and a smooth basal ring (Wójcicki, personal observation). Despite the obvious morphological differences, such a combination of characters suggests their similarity to T. kvacekii, and leads us to suspect that the newly described species may have given rise to some extant morphotypes, but convergence cannot be ruled out. Similarly to some other fossil Trapa species of the Miocene of Europe (Kovar-Eder et al. 2005), T. kvacekii was probably endemic to the late Miocene of Thessalia; it highlights the unique character of its flora. ACKNOWLEDGEMENTS Special thanks are due to J. Kovar-Eder (Stuttgart), E. Zastawniak (Kraków) and Z. Kvaček (Prague) for valuable scientific discussions and to the two anonymous reviewers for helpful comments on the manuscript. The first author is additionally grateful to the curators and keepers of the palaeobotanical collections of Barcelona (C. Martin-Closas), Berlin (D.H. Mai), Beijing (C.S. Li), Brno (R. Gregorova), Bucharest (N. Ţicleanu), Budapest (L. Hably, L. Kordos), Cottbus (U. & R. Striegler), Dresden (L. Kunzmann), Frankfurt (V. Wilde), Kraków (E. Zastawniak), London (P. Kenrick), Munich (W. Jung, H. Mayr), Nanjing (S.X. Guo), Stockholm (M.E. Friis), St. Petersburg (S.G. Zhilin), Turin (E. Martinetto) and Vienna (J. Eder), as well as to the curators of the herbaria B, BM, BP, CAL, FI, KRAM, KW, LE, LY, MW, MHA, P, PE, PR, PRC, S, W, WU, Z for kindly allowing him to study comparative fossil and extant material, to J.W. Wieser (Kraków) for drawings of Trapa specimens, and to M. Jacobs (Miechów) for verification of the English text. 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