Acta Palaeobotanica 47(1): 135–143, 2007

The fl oral change in the Tertiary of the Rhön mountains (Germany)

DIETER HANS MAI

Museum für Naturkunde, Humboldt Universität zu Berlin, Invalidenstraße 43, 10115 Berlin, Germany

Received 9 May 2006, accepted for publication 10 December 2006

ABSTRACT. A total of 438 species from 8 stratigraphically well-dated macro- and microfloras from the Rhön mountains (Central Germany) are biostratigraphically and palaeoecologically analysed. The formations are stratigraphically discontinuous but span the age range mid/late Eocene to late Pliocene (ca. 39 Ma to 2.15 Ma). The primary compositional elements and ecological character of each formation’s fl ora is as follows: – Kaolinitic Formation: Eocene with Beckettia, Steinhauera, Sterculia, Wardensheppeya; Pollenfl ora of SPP- zone 16/17 (Bartonian). – Sieblos Formation: early Oligocene. Palynological spectrum including Boehlensipollis hohlii; invasion of Arctotertiary elements. Many Nymphaea remains. – Kühnstein Formation: “Marxheim”– pollen-spectrum of early late Oligocene. No macrofl ora. – Oberleichtersbach Formation: dated to the late Oligocene by small mammals and gastropods. 100 plant species of fruits and seeds. Mixed mesophytic forest of the Carpino-Ostryon-type; many semihumid elements and herbs. – Escheri Formation: beds south of Theobaltshof; small fl ora with Alnus julianiformis, Fagus deucalionis, Quercus drymeja and Q. kubinyi. Palynological ratio between Triatriopollenites coryphaeus and T. rurensis indi- cates early Miocene. – Kaltennordheim Formation: fossil fl oras of Kaltennordheim, Hochrain, Bauersberg and Lettengraben with more than 65 plant species, characterized by typical elements of the so called Younger Mastixioid fl oras (Eomas- tixia hildegardis); optimal warm, humid climatic phase. Latest early Miocene to early middle Miocene. – Wollbach Formation: dominated by Quercus pseudocastanea, together with Acer pseudomiyabei, Alnus ducalis, Buxus pliocenica, and Eucommia europaea. Warm temperate, deciduous broad-leaved forest of late Miocene affi nity. – Borsoni Formation: 130 plant species from six localities belonging to 59 families. Deciduous broad-leaved warm-temperate forests and different aquatic- and swamp-plant taphocoenoses with many exotic species. Marked east-European infl uence. Two different fl oral complexes document a late Pliocene warming (Gerstungen, Oberzella) and an intra-Pliocene cooling (Kaltensundheim).

KEY WORDS: macrofloras, microfloras, floral change, Tertiary, Rhön mountains, Germany

INTRODUCTION

The Rhön mountains, a volcanic low moun- been investigated. More than 450 fossil plant tain range in the centre of Germany (Fig. 1), species have been discovered and described. lies between Bavaria, Hesse and Thuringia These plant fossils are well placed stratigraphi- (Schmeer 1964) and yield numerous Tertiary cally by association with fossil gastropods and fl oras of different ages. Knowledge of these plant mammalian remains, together with several fossils goes back to the 19th century (Zenker radiometric dates of the volcanites (Lippoldt 1833, Hassencamp 1858, 1860, Heer 1859) and 1978). The sequence of fl oras (Fig. 2) started increased after investigations in recent years with many gaps in the Eocene and terminated (Geyer 2002, Gümbel & Mai 2002, 2004, 2006, in the Upper Pliocene (Martini et al. 1994). Mai in press). Fossil fruits and seeds, leaves, The fl oral change over this long time (39 Ma pollen and spores, woods and charophytes have – 2.47 Ma) was very signifi cant. 136

Fig. 1. The distribution of volcanic rocks with the main fossiliferous places in the Rhön mountains. After Gümbel & Mai (2006) – localities, and Schmeer (1964) – volcanic rocks 137

Fig. 2. The stratigraphical assignment of the sedimentary and fossiliferous beds of the Rhön mountains (no true to time scale and to massivness of the sediments). After Martini et al. (1994), changed

EOCENE Intratriporopollenites ceciliensis Krutzsch, I. minimus Mai, Labrapollis labraferus KAOLINITIC FORMATION (R. Pot.) Krutzsch, and Plicatopollis plicatus (Kr.) Krutzsch. Palaeocarpological remains were described Age. The pollen assemblage places the only from the research drilling Sieblos 1994/1 Paleogene basal layers into the Eocene. This near the Wasserkuppe in a depth of 36.65 to might be correlated with the SPP – zone 16/17 38.10 m. The macrofl ora of 16 species from after Krutzsch et al. (1992); Bartonian, ca. clayey coal seam sediment (Kaolinitic Forma- 39 Ma. tion) lying immediately above Lower Muschel- References. Krutzsch et al. (1992), Hot- kalk indicates middle to late Eocene age. tenrott (1998), Mai (1998). Flora. Beckettia mastixioides Reid & Chan- dler, Steinhauera subglobosa Presl, Sterculia subovoidea (Reid & Chandl.) Mai, and War- OLIGOCENE densheppeya marginata (Chandl.) Eyde are most important species. The palynological SIEBLOS FORMATION investigation of the borehole Sieblos 1994/1 yielded microfl oras only from the underlying The abandoned coal mine of Sieblos at the substratum. Signifi cant pollen are Composit- Wasserkuppe was famous for its fossil contents oipollenites rhizophorus (R. Pot.) R. Potonié, already during the 19th century and has been 138 accepted as a classical locality of the German Hottenrott (1988, 1998), Martini & Rothe palaeobotany. Since most of the fossils so far (1998, 2005). collected were from mining dumps, they could not be positioned into a relative stratigraphi- KÜHNSTEIN FORMATION cal succession. The Tertiary period was re-exa- At the bend of the street no. 284 south-east mind by scientifi c drillings in the years 1994, of the Wasserkuppe near Gersfeld in a land- 1998, and 1999. The results of the investiga- slide a small deposit of brown-coal clay and tions of the whole Sieblos Formation were pub- dysodile was investigated in 1975. This was lished in monographical studies. Many details named after the neighbouring hill as Kühn- about the sedimentology, palaeoecology, vol- stein beds. canology, and geochemistry of this remarkable Flora. A microfl ora (only 56 species) freshwater deposit in the Oligocene of Central from these beds shows very characteristical Europe are well known. The structure Sieblos amounts of Sparganiaceaepollenites, Cyper- is suggested as a suberosional depression. aceaepollis, Alnus and Platanus pollen grains. Flora. A dysodile bed of Sieblos has Very typical elements are also Polypodiaceois- included many plants of intrazonal vegetation, porites marxheimensis (Mürr. & Pfl ug) Krut- predominantly Nymphaea nymphaeoides (Ett.) zsch, Diplopollis kockelii Pfl anzl (= Calamus), Weyland (= N. rhoenensis Kurtz 1894), and Momipites punctatus (R. Pot.) Nagy (= Engel- Spirematospermum wetzleri (Heer) Chandl. hardia), and Tricolporopollenites cingulum The association of Glyptostrobus europaeus (R. Pot.) R. Pot. & Pfl ug. One uncertain speci- (Brongn.) Heer – Nyssa disseminata (Ludw.) men of Boehlensipollenites hohlii Krutzsch Kirchh. verifi ed the swampy forest, possibly possibly indicates early late Oligocene. together with Myrica banksiaefolia Unger, Age. The differences between this “Marx- and the fern Lygodium cf. kaulfussii Heer. heim pollen spectrum” of the Kühnstein For- The zonal vegetation was characterized by the mation and the younger microfl oras of Hesse members of a mixed mesophytic forest. Impor- and the Rhön mountains are evident. No mac- tant elements of this forest were Tetraclinis rofl ora has been discovered. salicornioides (Ung.) Kvaček, Carya rostrata The lithological composition, thickness, and (Schloth.) Schimper, Engelhardia orsbergensis regional extent of the Kühnstein Formation is (Wessel & Web.) Jähnichen et al., Quercus unknown. Therefore nothing can be said about lonchitis Ung., Tilia irtyshensis (Shaparenko) the overlapping with volcanites (tuffs) and their Grubov, Zanthoxylum europaeum Ung., and radiometric age. The eruptions in the region of many Lauraceae. This is very similar to the the Wasserkuppe started approximately before conditions known from the volcanodetritic 25.0 Ma., that is in accordance with the new series in the Bohemian mountain range. The time scale late Oligocene (Chattian). macrofl ora of only 32 species contrasts with References. Hottenrott (1992), Krutzsch a richer microfl ora of 48 species with typical (1993), Hottenrott et al. (1998). elements as Cicatricosisporites paradorogensis Krutzsch, Momipites quietus (R. Pot.) Nichols., OBERLEICHTERSBACH FORMATION and Boehlensipollis hohlii Krutzsch, verifying unambiguously older Oligocene. Worth men- The fossil remains at the locality Oberleich- tioning is the strong invasion of Arctotertiary tersbach near Bad Brückenau (Bavarian Rhön) elements as in the “fl oral complex Valeč” at the derive from a blue-black clayey sediment of same time. the fi lling of a doline in the Muschelkalk. This Age. The palynological picture, mainly sediment is very rich in fossil fruits and seeds, Boehlensipollis, demonstrates typical Oli- ostracods, snails, shells, and mammal bones in gocene. Based on recently found teeth of excellent preservation. smaller mammals, especially by the presence Age. Based on gastropods, e.g. the strati- of Melissiodon (family Melissiodontideae) the graphically important Ferussina (Stropho- age of the Sieblos Formation can be deter- nema) tricarinata M. Braun, and a great mined more precisely as early Oligocene. diversity especially of small mammals the References. Hassencamp (1858, 1860), sediments can be placed in the latest late Heer (1859), Schenk (1891), Kurtz (1894), Oligocene (Neochattian, Paleogene mammal Müller-Stoll (1936), Jähnichen & Rüffl e (1988), zone – MP 30, Coderet). The assignment of 139 a very rich macrofl ora to a generally new fl o- Next to the deposit of Theobaltshof escheri ral assemblage “Oberleichtersbach” (24.5 Ma beds are known from Kaltennordheim, Eis- to 23.0 Ma) is quite within bounds of prob- graben near Hausen, Erdpfahl near Roth, and ability. Bauersberg near Bischofsheim, mostly with- Flora. On the basis of fossil fruits and out fossil plants. Plant macrofossils are well- seeds, 100 plant species belonging to 63 fami- known only from the “underlying brown clays” lies have been described from the locality till (tuffi tes or ash-tuffi tes) and lacustric marls now. Many other species await a description. in Kaltennordheim. The poor fl ora (16 spe- This well-preserved fl ora is one of the richest cies) yields Alnus julianiformis (Stbg.) Kvaček and most interesting Tertiary fl oras of Cen- & Holý, Engelhardia macroptera (Brongn.) tral Europe. The composition refl ects a mixed Ung., Fagus deucalionis Ung., Quercus drymeja mesophytic forest of the Carpino – Ostryon Ung., Q. kubinyi (Ett.) Czeczott, Trigonobalan- -type with many semihumid – humid alter- opsis rhamnoides (Rossm.) Kvaček & Walth., nating elements. Dominant species are and Zelkova zelkovifolia (Ung.) Bůžek & Kotl., Carpinus cordataeformis Mai and Ostrya which are of stratigraphical interest. scholzii Gregor together with Carya rostrata The dysodiles of the Dietrichsberg Member (Schloth.) Schimper, Celtis lacunosa (Reuss) near Vacha with a fi sh fauna of Palaeoleu- Kirchh., Cynoxylon carolii Mai, Ehretia euro- ciscus dietrichsbergensis M. Böhme and an paea E. M. Reid, Gironniera carinata Mai, undescribed macrofl ora with many winged Schisandra moravica (Mai) Gregor, Toddalia fruits and seeds Buzekia tertiaria (Weyland) latisiliquata (Ludw.) Gregor, Wikstroemia Manchester may be contemporaneous with thomasii Gregor & Geissert, and Zizyphus the Escheri Formation. striatus (Ludw.) Mai & Gregor. Palaeobiologi- Age. Important to the biostratigraphy how- cal signifi cance had also some annual plants of ever are the palynological investigations. The open environments, e.g. Amethystea heckiana diagram of the relations between coryphaeus/ Mai, Chenopodium wetzleri Gregor, Polanisia rurensis (= Triatriopollenites coryphaeus punc- sibirica Dorof., Silene microsperma Mai etc. tatus (R. Pot.) Th. & Pf. /Triatriopollenites A new aquatic plant of paratethyan affi nity is rurensis Th. & Pf.) shows the assignment of Ceratostratiotes antiquus Mai. So far, this fos- the Escheri Formation to the Lower Miocene sil fl ora is unique. (M 1) in Hesse. References. Martini (2000), Mai (in press). The molar of Amphitragalus cf. boulangeri Pomel, a precursor of the deer family, is also a sure reference of early Miocene (Neogene MIOCENE mammal zone MN3). References. Knobloch (1971), Moayed- ESCHERI FORMATION pour (1977), Gregor (1982), Hottenrott (1988), The stratigraphically important series of Böhme (1996), Geyer (2002), Gümbel & Mai Miocene sediments in the Rhön mountains is (2002). developed south of Theobaltshof near Tann. KALTENNORDHEIM FORMATION Tuff and tuffi tes verify an intensive volcanism at the time of sedimentation. Above a basal The Kaltennordheim Formation covers tuff-limestone horizon follows the original a changing succession of clayey and calcare- escheri – horizon, gray-green, coarse-grained ous marly sediments together with plentiful tuffi tes with the snail Brotia escheri aqui- brown-coal seams, which were the objects of tanica, in the upper part mixed with clay, many mines in the former time. Plant fossils sand or tuffi tes. Country-loving snails and were found at different localities. A Gyraulus mammals, and also the plant remains were horizon, named after the lacustric gastropod introduced from the surrounding woodlands. Gyraulus trochiformis dealbatus Thomae, Flora. The aquatic vegetation was obvi- a leading horizon of tuffi tic and lacustric sedi- ously dominated by charophytes. Important ments meshed with basalt, may be present macrofossils are Fagus attenuata Goepp. resp. in all the local basins of the Kaltennordheim F. saxonica Kvaček & Walth., Betulaceae aff. Formation (Fig. 2). Alnus, Carya sp. aff. ventricosa (Stbg.) Ung., Flora. The main localities of fossil fl oras Celtis lacunosa (Reuss) Kirchh., and Pinus sp. are Kaltennordheim, Hochrain near Gersten- 140 grund, Bauersberg north of Bischofsheim, and discovered, but the composition of the fauna is the mine “Barbara” at the Lettengraben (Her- of a Lower to Middle Miocene character. renwasser) near Wüstensachsen – Ehrenberg. Age: The fossil fl ora is characterized by All these localities contain fossils of evergreen, typical elements of the so-called Mastixioid fl o- broad-leaved and mixed mesophytic forests ras, which generally attest to an optimal warm with many characteristic geofl oral elements. humid climatic phase of the Miocene. The age A very rich macrofl ora comes from the mine of this phase is Lower to Middle Miocene “Barbara” where 63 plant species belonging to based on the very likely assignment of the 38 families were described. Important are the Lettengraben fl ora to the fl oral assemblage newly discovered evergreen trees and shrubs “Eichelskopf – Wiesa” (zone VI – VIII) or the of the families Taxaceae (Amentotaxus), Lau- Kaltennordheim fl ora to the fl oral assemblage raceae, Magnoliaceae, Rutaceae (Todda lia, “Kleinleipisch – Františkove Lázně”(zone X Zanthoxylum), Sabiaceae (Meliosma), Sym- – XII). The age 16.8 Ma to 17.5 Ma is conform plocaceae, Ericaceae (Vaccinioides), Fagaceae to the radiometric dates of the neighbouring (Trigonobalanopsis), Hamamelidaceae (Disty- basalts (18 Ma). The palynostratigraphic cor- lium), Menispermaceae (Parabaena), Mora- relation of the Bauersberg lignite-seam with ceae, Myricaceae, Staphyleaceae (Turpinia), the sediments of the Prososthenia – beds near and Styracaceae. This fl ora represents a typical Frankfurt am Main support the latest Early warm-temperate rain forest that is assigned to Miocene age. the fl oral assemblage “Eichelskopf – Wiesa” by References. Zenker (1833), Heer (1859), 74.6 % of all species. Hassencamp (1860), Müller-Stoll (1936), From the brown-coal bed and the clayish Kelber & Gregor (1987), Gregor (1990), Hot- intermediate sediments at the mine “Einig- tenrott (1992), Geyer (2002), Gümbel & Mai keit” of the Bauersberg comes also a macro- (2002, 2006). fl ora of 65 species partly known since the WOLLBACH FORMATION 19th century. Remarkable is the enriching of aquatic plants as Potamogeton schenkii Cross-bedded gravels in small areas are Kirchh., Salvinia cerebrata Nikitin, or Stra- preserved in the south-eastern foothills of the tiotes kaltennordheimensis (Zenker) Keilhack Rhön mountains. The conservation of such in the brown-coal. Palynological investiga- relics of a Tertiary fl uvial system are related tions of the Bauersberg lignite-seam reveal to the subrosion of Triassic salt layers in the a microfl ora of more than 45 species and allow underground. In the gravel pit Wollbach near the correlation of this seam with lignite-bear- Bad Neustadt a lense of plastic clay contains ing sediments from the area of Frankfurt am a very poor leaf fl ora of not more than 34 spe- Main (Prososthenia – Beds). cies from 16 families. From the brown-coal layers of Kaltenord- Flora. The fl oral assemblage, mainly domi- heim and the neighbouring Hochrain in the nated by Quercus pseudocastanea Goepp., and same basin 36 plant species belonging to 27 also with the presence of Acer pseudomiya- families have been described on the base of fos- bei Baikovskaja, Aesculus sp., Alnus ducalis sil fruits and seeds. The most important plant Gaudin emend. Knobloch (younger synonym fossils are Eomastixia hildegardis (Ung.) Holý, A. hoernesii Stur), Buxus pliocenica Saporta and Actinidia germanica Mai, two species of & Marion, shows unmistakeably a fl oristical middle Miocene range. relationship to fl oral assemblages of the Pan- Fauna. Small mammals were found in the nonian or Pontian in the Paratethyan region of marly limestone beds of the Kaltenordheim Europe. The fl ora refl ects a warm-temperate, Formation (see Geyer 2002): Glirundinus sp., deciduous broad-leaved forest. Heteromyoxus wetzleri Schlosser, Melissiodon Age. The fl ora is of Upper Miocene affi nity. schlosseri Schaub, Miopetaurista sp., Peridyro- In contradiction, mammals from a similar fl u- mys sp., Pseudotheridomys parvulus Schlosser. viatile series of Ostheim vor der Rhön indicate Remains of the big mammal Aceratherium sp. late Pliocene age. New fossil material and the derive from the coal clay of Kaltennordheim. future investigation of the whole fl ora may con- Common at most localities was the fi sh Leu- tribute to solve the problem of age correlation. ciscus (Palaeoleuciscus) dietrichsbergensis References. Kelber (1980, 1988), Knobloch M. Böhme. No index mammalian species were (1986). 141

PLIOCENE Dorof., Betula cholmechensis Dorof., Eleo- charis praemaximowiczii Dorof., Ficus tam- BORSONI FORMATION bovica Dorof., Myrica goretzkyi Dorof., Pota- The borsoni beds are of fl uvial origin and mogeton aculeatus Dorof., P. borysthenicus represent the youngest Neogene sediments of Dorof., P. felixii Dorof., Ranunculus tanaiticus the Rhön. At some places the sediments are Dorof., Teucrium pripiatense (Dorof.) Wieliczk. sunken into subrosion depressions and are & Zastawniak, and Typha pseudovata Dorof. clearly limited. At other places they are only as pieces of evidence of east-European infl u- relics of thick gravel bodies marking former ence in the west-European uppermost Neo- river courses. On account of the time of their gene. Pollenanalytical investigations from genesis and the remains of mammalian teeth Oberzella, Kaltensundheim and Mellrichstadt at many localities the term “arvernensis-time” shows typical spectra of the Upper Pliocene. is applied. In the northern foreland of the Fauna. Signifi cant in some localities of mountains near the river Werra the fi llings the Borsoni Formation of the Rhön are fossil of local dolines were found at Gerstungen, teeth of mastodons (Fulda, Sülzfeld, Jüchsen, Oberzella near Vacha, Barchfeld near Bad Ostheim vor der Rhön) or skeletons (Kalten- Salzungen, and Kaltensundheim at the Felda sundheim): Mammuthus (Zygolophodon) bor- river, and near the river Haune the dolines of soni Hays, Anancus arvernensis Croiz. & Job., Buchenau and Hünfeld with minimal brown- and Tapirus arvernensis Croiz & Job., and coal seams. They represent fossiliferous clay some deer-like mammals as Hypolagus and lenses with fl oral remains, mostly fruits, Metacervoceros. All these mammalian fossils seeds, and pollen grains . facilitate to date these deposits into the Late Flora. In summary from the six localities Pliocene. 127 plant species belonging to 59 families were Age. The middle to late Pliocene age of the described and evaluated palaeoecologically and Borsoni Formation in the Rhön mountains is biostratigraphically. Deciduous warm-temper- based on mammals as well as on palaeomag- ate forests and different aquatic- and swamp- netic dates of the sediments (Wiegank 1981, plant taphocoenoses are recognized. Important 1982). The time interval lies between 2.8 exotic trees and shrubs are Acanthopanax Ma to 2.1 Ma. The palaeofl oristically well uralensis Dorof., Alnus lusatica Mai, Carpinus investigated fl oral assemblages “Ceyssac miocenica Negru, Cunninghamia minisperma – Kaltensundheim” (2.8 Ma to 2.56 Ma) and Mai & Walth., Glyptostrobus brevisiliquatus “Berga – Reuver” (2.65 Ma to 2.47 Ma) most (Ludw.) Mai, Ilex thuringiaca Mai, Magnolia probably correspond to the late Pliocene. The cor Ludw., Meliosma miessleri Mai, Nyssa fi rst assemblage documents a cooling, the sec- disseminata (Ludw.) Kirchh., Phellodendron ond a warming, . The assemblage of Hünfeld, elegans C. & E.M. Reid, Picea omoricoides which may be “Perrier – Rippersroda” (2.3 Ma C.A. Web., P. rotunde-squamosa (Ludw.) to 2.15 Ma) is uppermost Pliocene represent- Mai & Walth., Pseudolarix schmidtgenii Kr., ing a new forest phase (Krutzsch 1988). Pterocarya limburgensis C. & E.M. Reid, References. Blankenhorn (1901), Hofmann Sequoia abietina (Brongn.) Knobl., Stachyurus (1941), Leschik (1952, 1954), Schaarschmidt merkaensis Mai, Stewartia beckerana (Ludw.) (1958) Duphorn (1961), G. Böhme (1963, Kirchh., Symplocos casparyi Ludw., Taxo- 1992), Krutzsch & Majewski (1965), Wiegank dium rossicum Dorof., Trema lusatica Mai, (1981, 1982), Kahlke & Ukraintseva (1986), Trichosanthes fragilis C. & E.M. Reid, Tsuga Grüger in Büttner (1988), Krutzsch (1988), moenana Kirchh., etc. Remarkable species of Mai & Walther (1988), Böhme (1996), Gümbel the aquatic vegetation are Lobelia pliocenica & Mai (2004). (Dorof.) Mai, Microdiptera menzelii (E.M. Reid) Mai, Nuphar canaliculata C. & E.M. Reid, ACKNOWLEDGEMENTS Ranunculus gailensis E.M. Reid, Salvinia I am deeply grateful to Steven R. Manchester miocenica Dorof., and Trapa heerii Fritsch. (Gainesville, Florida) and David Lazarus (Berlin) for Biostratigraphically important and phytogeo- reading and correcting the English version of the manuscript, and Ms Elke Siebert (Berlin) for graphi- graphically very interesting are species such cal assistance. I thank also the fossil-hunters in the as Acorus palaeocalamus Dorof., Acorellus Rhön Tertiary which offered me the free disposal of distachyoformis Łań.-Środon., Alnus tanaitica their plant fossils. 142

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