Palaeobiodiversity and Palaeoenvironments https://doi.org/10.1007/s12549-020-00461-0 ORIGINAL PAPER Fossil Gordonia (s.l.)–like (Theaceae) winged seeds from the early Miocene of the Mecsek Mts, W Hungary Boglárka Erdei1 & Lilla Hably1 Received: 3 March 2020 /Revised: 5 May 2020 /Accepted: 21 September 2020 # The Author(s) 2021 Abstract Winged seeds were recovered from two sites of the late early Miocene (Karpatian) flora of Magyaregregy, Mecsek Mts, W Hungary. The seeds are assigned to the fossil-genus and species, Mecsekispermum gordonioides Hably and Erdei gen. nov. et sp. nov., and are tentatively related to the family Theaceae. Based on the overall character of the winged seeds and the isodiametric surface pattern of the seed coat, the seeds are most comparable with species of Gordonia J. Ellis (s.l.,) in Theeae (Laplacea Kunth or Polyspora Sweet). A comparison with winged seeds of other fossil genera, e.g. Saportaspermum Meyer and Manchester, and winged seeds of modern genera in various families is also given. The fossil flora is preserved in the fish scale-bearing clay marl belonging to the Feked Formation and Komló Claymarl Member and dated as Karpatian (late Burdigalian, standard chronostratigraphy). Keywords Fossil flora . Magyaregregy . Saportaspermum . Mecsekispermum . Fish scale-bearing clay marl . Karpatian Introduction Kentucky and Tennessee (Grote and Dilcher 1992; Martínez-Millán 2010). Winged fruits or seeds have frequently been reported in the The classification of the family Theaceae varied in many European fossil record. Disseminules having an apical or lat- authors, according to the characters emphasised (Prince and eral wing have been described variously as Cedrelospermum Parks 2001). In the Cronquist system, the family involves ~ 40 Saporta, Gordonia J. Ellis, Saportaspermum Meyer and genera and 600 species in four subfamilies (Cronquist 1981). Manchester, or Embothrites Unger and have been assigned Molecular data showed the family as non-monophyletic (e.g. to various plant groups, e.g. Ulmaceae, Theaceae, and Morton et al. 1996;Soltisetal.2000), and APG (1998)sug- Malvales (Unger 1850; Manchester 1987; Meyer and gested a narrow definition of the family with Theaceae s.str. Manchester 1997; Kvaček and Walther 1998; Kvaček and and a separate Ternstroemiaceae (Ternstroemieae in Wilde 2010). Fruits or seeds assigned to Theaceae are rare Pentaphylacaceae). Based on molecular data, three main line- not only in the European fossil record but also worldwide. ages of Theaceae s.str. were identified, i.e. Theeae, Grote and Dilcher (1989, 1992) gave a summary of fossil Stewartieae, and Gordonieae (Prince and Parks 2001). specimens reported as remains of Theaceae. The earliest fossil According to the same study, the monophyly of the genus fruits assigned to Gordonia (Gordonia lamkinensis Grote and Gordonia was not supported, and sequence data placed its Dilcher and Gordonia warmanensis Grote and Dilcher) were species in Gordonieae (Gordonia s. str.) and Theeae described from the middle Eocene Claiborne flora in (Laplacea Kunth and Polyspora Sweet). Although a common character of the seeds of Gordonia s.l. species is a prominent flattened apical wing, micromorphological study of the seed coat was used to corroborate the above placement of This article is a contribution to the special issue “Palaeobotanical contributions in honour of Volker Mosbrugger” Gordonia s.l. species (Gunathilake et al. 2015). Leaves of Theaceae have been reported from some locali- * Boglárka Erdei ties in the Northern Hemisphere fossil record during the [email protected] Cenozoic. Today the area of the family occupies subtropical- tropical Asia and America (Prince and Parks 2001); however, 1 Botanical Department, Hungarian Natural History Museum, palaeofloristic data indicate the presence of the family also in Könyves Kálmán krt. 40., Budapest 1087, Hungary Europe. Leaf records of Theaceae (s.l.) have been documented Palaeobio Palaeoenv in the European floras from the middle Eocene up to the late by the Budafa Formation, traditionally regarded as of Miocene. From the middle Eocene flora of Messel (Germany; Karpatian (late Burdigalian) age (Gyalog 1996;Gyalogand Wilde 1989) and from the early Oligocene flora of Flörsheim Budai 2004; Budai et al. 2015).TheBudafaFormation (Mainz Basin, Germany; Kvaček 2004), leaves of two genera, consisted of three members, the Komló Claymarl Member, Gordonia and Ternstroemites Berry, were mentioned. Leaves Pécsvárad Limestone Member, and the overlying Budafa preserved with cuticle were reported as Gordonia hradekensis Sandstone Members. The succession is overlain by (Kvaček and Bůžek) Bozukov and Palamarev, Gordonia Badenian (Langhian) Leitha limestones in the littoral zone pseudoknauensis Kvaček, and Ternstroemites floersheimensis and offshore sands and silts in the basins, locally with coal- Kvaček and Walther from the rich, thermophilous flora. From bearing swamp deposits along the shores. Based on the ab- the early Miocene flora of Hrádek (North Bohemia, sence of marine fossils in the Pécsvárad Limestone and in Czech Republic; Holy et al. 2012) and from the flora of the most part of the Komló Claymarl, Sebe et al. (2019)conclud- Cypris Shale (Western Bohemia, Czech Republic; Bůžek et al. ed that these rocks are lacustrine deposits. As suggested by the 1996), leaf remains of Gordonia hradekensis were mentioned. lake sediments and its fauna, instead of a system of smaller From the early Miocene (Ottnangian) flora of Köflach lakes, a contiguous water body, called “Lake Mecsek”,existed (Austria) ?Gordonia oberdorfensis Kovar-Eder and ?Cleyera in the area from the Karpatian up to the early Badenian ac- schilcheriana Kovar-Eder (Kovar-Eder and Meller 2001) cording to K/Ar age and biostratigraphic dating of the overly- were described based on leaf remains and preserved cuticle. ing marine deposits (Sebe et al. 2019). The Pécsvárad and From the late early/early middle Miocene flora of Parschlug Komló Members are both interpreted as lake deposits and (Austria; Kovar-Eder et al. 2004), there are also some leaves differ very much from the Budafa Sandstone Member; assigned to Theaceae, cf. ?Gordonia oberdorfensis and therefore, Sebe et al. (2019) united the two members as a Ternstroemites pereger (Unger)Kovar-EderandKvaček. separate formation, the Feked Formation. From the middle Miocene (Badenian) of the Satovcha Consequently, the “fish scale-bearing clay marl” Graben in the Western Rhodopes (Bulgaria), Bozukov and sediments preserving the plant fossils belong to the Feked Palamarev (1995) described a flora as a rich refuge of the Formation and Komló Claymarl Member, dated as Karpatian representatives of Theaceae (s.l.), e.g. Eurya aff. (late Burdigalian) (Sebe et al. 2019). It is composed of dark acuminatissima Merril and Chun, Gordonia hradekensis, grey, greenish-grey, massive, or laminated silty clay marl, Gordonia stefanovii Palamarev and Bozukov, Hartia calcareous silt, and fine sands, with numerous fish scales and palaeorhodopensis Bozukov and Palamarev, Stewartia bones, and sometimes with tuff interbeds. K/Ar dating mea- stefanovii Palamarev and Bozukov, and Stewartia surements from tuff interbeds within the Komló Claymarl submonadelpha Tanai and Onoe. In Europe, Theaceae pre- showed an age of 16.82 ± 0.65 Ma, i.e. Karpatian (late sumably survived up to the late Miocene or Pliocene based Burdigalian) (Sebe et al. 2019). on reports from the Sessenheim flora in France (Geissert et al. Palaeontological research of the Mecsek Mts dates back to 1990), Fritzlar (Hessen) in Germany (Hottenrott et al. 1996), decades, attested by numerous studies, e.g. nannoplankton and refuge areas of the Pannonian Basin. A rich late Miocene (Báldi-Beke 1963), foraminifers (Korecz-Laky 1968), and thermophilous flora was described from Mataschen (Austria, palynology (Nagy 1969). After floristic works published by Kovar-Eder and Hably 2006) with four species assigned to Staub (1882), Andreánszky (1955), and Pálfalvy (1953, 1961, Theaceae. Gordonia emanuelii Kovar-Eder, Gordonia 1964, 1967), Hably L. and her colleagues have contributed pannonica Kovar-Eder, Gordonia styriaca Kovar-Eder, and with studies to the knowledge on the Miocene flora of the Schima mataschensis Kovar-Eder were described based on Mecsek Mts (Hably 1992, 2001, 2002; Hably and Thiébaut leaf remains with cuticle. The flora of Magyaregregy also 2002). A monographic study of the plant fossils of yielded leaf remains described as Ternstroemites pereger, Magyaregregy is going to be published by Hably (2020). which may be related to the Theaceae (s.l.) family (Hably The Magyaregregy flora is extremely diverse with more than 2020). 100 taxa hitherto described, including Glyptostrobus europaeus (Brongniart) Unger, Zizyphus paradisiaca (Unger) Heer, Daphnogene cinnamomifolia (Brongniart) Geology Unger,Laurophyllumsp., Myrica lignitum (Unger) Saporta, Engelhardia orsbergensis (P.Wessel and C. O. Weber) The area of the Mecsek Mts is situated in the SW Pannonian Jähnichen, Mai and Walther, Cedrelospermum flichei Basin, on the Tisza-Dacia Megaunit, a tectonic block south of (Saporta) Hably and Thiébaut, C. aquense Saporta, Zelkova the mid-Hungarian Shear Zone. The Neogene stratigraphy of zelkovifolia (Unger) Bůžek and Kotlaba, legumes the Mecsek area was summarised by Hámor (1970), Chikán (Podocarpium podocarpum (A. Braun) Herendeen, (1991), and Barabás (2010, 2011). Widespread lower Leguminosae gen. et sp.), Ailanthus confucii Unger, and Miocene fluvial clastics (Szászvár