Acta Palaeobotanica 51(2): 127–179, 2011

Biodiversity and the reconstruction of Early Jurassic fl ora from the Mecsek Mountains (southern Hungary)

MARIA BARBACKA

Hungarian Natural History Museum, Department of Botany, H-1476, P.O. Box 222, W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Kraków, Poland; e-mail: [email protected]

Received 15 June 2011; accepted for publication 27 October 2011

ABSTRACT. Rich material from Hungary’s Early Jurassic (the Mecsek Mts.) was investigated in a palaeoen- vironmental context. The locality (or, more precisely, area with a number of fossiliferous sites) is known as a delta plain, showing diverse facies, which suggest different landscapes with corresponding plant assemblages. Taphonomical observations proved that autochthonous and parautochthonous plant associations were present. The reconstruction of the biomes is based on the co-occurrence of taxa and their connection with the rock matrix and sites in the locality, as well as the environmental adaptation of the plants expressed in their morphology and cuticular structure. The climatic parameters were confi rmed as typical for the Early Jurassic by resolution of a palaeoatmospheric CO2 level based on the stomatal index of one of the common species, Ginkgoites mar- ginatus (Nathorst) Florin. Plant communities were differentiated with the help of Detrended Correspondence Analysis (DCA); the rela- tionship between taxa and sites and lithofacies and sites, were analysed by Ward’s minimal variance and cluste- red with the help of the UPGMA algorithm. These methods aided the recognition of fi ve phytocoenoses in the Mecsek delta: swamp, dominated by the pteridosperm Komlopteris associated with cheirolepidiaceous and other conifers and ginkgophytes; swamp areas along river banks with conifers; seasonally drier fl oodplain colonised by cycads, bennettites and sphenophytes; wetter fl oodplain with dominant Ginkgoites marginatus; highly disturbed, well-drained islands occupied mainly by ferns; and relatively dry areas dominated by Sagenopteris and associated with cycads and ferns.

KEYWORDS: plant assemblages, palaeoecology, statistic methods, Early Jurassic, Hungary

INTRODUCTION

Fossil plants and their responses to pal- through investigations of plant adaptation as aeoenvironments, their assemblages and expressed in their morphology and cuticular mutual interactions have met with increas- structure, up to modern mathematic modelling ing interest from the second half of the 20th and statistical analyses are used in palaeo- century onwards. Based on taxonomic studies, botany. which are the starting point for palaeoecologi- The Early Jurassic fl ora from the Mecsek cal patterns, different methods were provided Mts. (South Hungary) has been collected and for trying to understand the functional mecha- studied since 1989 (Barbacka 1992, 1994a,b, nisms active in palaeo-communities. 2001, 2002, 2009). The fl ora now consists of 55 Individual localities with their specifi c con- species. Taxonomical studies were often sup- ditions of plant preservation offer varied oppor- plemented with taphonomical remarks and fol- tunities for palaeoecological reconstructions. lowed by environmental conclusions. The envi- To reach this aim, different methods, from ronmental hypotheses were mainly based on very simple listing of plants that give informa- supposed plant adaptations to differentiated tion about the content of plant assemblages, conditions. Working on the fossil fl ora from 128

Hungary, certain tendencies were observed in the regular repetition of species co-existing on the same rock slabs, and possible relations between taxa and their location within the area, as well as between taxa and the type of rock. The aim of the present paper, based on all available data, is to introduce the Hungarian fossil plant sites in an ecosystem model show- ing differentiated plant co-existence, interrela- tions of plant species and rock type, and the real distribution of plant assemblages. The objectivity of the results is provided by statistic methods. Such analyses are rather rarely used in palaeobotany, because they need a large record appropriate for statistics and proof of the autochthonous or parautochthonous origin of the material. The obtained results can be applied to a revi- sion of the plant assemblage-environment rela- tionship, and may form a basis for updating palaeoecological models based on palynological analyses. It can also contribute to the general knowledge of Jurassic plant assemblages and supply data for comparative studies.

PART 1 Geological and Paleoenviromental Background

GEOLOGY AND PALAEOGEOGRAPHY

The Early Jurassic plant macrofossil local- ity in Hungary is confi ned to the Mecsek Coal Formation (Hetényi in Császár 1997) in the Mecsek Mountains (Pécs and Komló regions, southern Hungary) and closely connected with coal mining. The coal-bearing area extends in a N – S direction, for a distance of about 20–30 km, from Nagymányok to Pécs (Fig. 1). The mining region occupies about Fig. 1. Map of sites in the Mecsek Mts. locality (A. Sojka 2 drawing): 1 – settlements, 2 – dumps, 3 – open mines, 4 – 350–400 km . The beginning of coal mining shafts in Hungary dates back more than 200 years. After the most intense exploitation period 1992) and palynological (Góczan 1956) inves- between 1960 and 1990, the underground tigations. mines were closed, and in 2004, open mining The Mecsek Coal Formation, which is the also came to an end. most interesting from a palaeobotanical view- The age of coal measures were determined point, comprises the Upper Triassic and the as Early Liassic (Hantken 1878) and narrowed Lower Jurassic sequences consisting of sand- down to the Hettangian by subsequent geologi- stone, claystone, mudstone and coal. The coal cal (Paál-Solt 1969, Nagy & Nagy 1969), pal- seams occur mainly in the middle and upper aeontological (Földi 1967, Nagy 1970, Szente parts of the sequence, which are dated as 129

Early Jurassic (Landis et al. 2003), but thin Beginning from the basal (Late Rhaetian) coal seams appear already in the fl uvial succes- part of the Mecsek Coal Formation the facies sion of the latest Rhaetian. At the beginning of show a well expressed trend of environment the Liassic, fl uvial–lacustrine–palustrine sedi- changes: delta – limnic – lacustrine – delta mentation continued, with paralic coal-swamp – lagoonal – plain marine – lagoonal – plain deposits becoming predominant in the later marine (Nagy 1969). During the Hettangian phase of the sedimentary record. sedimentation, it was mainly fl uvial with The formation is built of a cyclic alternation channel, fl ood plain and swamp facies. Fur- of arkosic sandstone, siltstone, claystone and ther up the sequence, coquinas of brackish- coal seams. The thickness and carbonization water molluscs appear in increasing fre- degree of the coal deposits are diversifi ed due quency. The Lower Sinemurian member of to asymmetry in the depository basin caused the formation may have been deposited in by unequal subsidence (Fig. 2): according to a tidal-fl at marsh environment. In the mid- Nagy (1969) the thickness is at its lowest near dle member of the Mecsek Coal Formation, Nagymányok (120 m), gradually increasing thin (0.5–1.5 m) rhyolitic tuffi te interlayers southward to Pécs (1000–1200 m thick). This occur (Haas 2001). asymmetry, observed also in the Upper Trias- Plant beds sometimes alternate with mol- sic Karolina völgy Sandstone Formation, is lusc-bearing marine sublittoral layers in the a result of non-uniform extension of the half- same horizons of the succession. In the upper graben (Nagy 1969). In the Alpine-Carpathian member of the Mecsek Coal Formation, crinoids region, Lower Liassic, coal bearing siliciclastic appear, indicating a temporary establishment sequences, showing features similar to those of “normal salinity” marine conditions. of the Mecsek Zone, are classifi ed as “Gresten The coal formation is overlain by fi ne- Facies” – a characteristic facies of the south- grained sandstone and dark-grey shale with ern margin of the European shelf of the Tethys thinning upward sandstone interlayers (Vasas (Haas 2001). Marl). In the lower part of the succession,

Fig. 2. Map of the Mecsek basin (modifi ed, after Nagy 1969): 1 – granitoid denudated area (sedimentary basin margine), 2 – carbonate denudated area (sedimentary basin margine), 3 – isolines of hypothetical basin thickness, 4 – direction of the sea current, 5 – direction of granitoid clast transport 130

Liogryphaea beds occur which may have formed 3. The uppermost section (Lower Sinemu- in a very shallow marine environment. Locally, rian) is of paralic type, characterized by thin other bivalves and crinoids, as well as sporadic coal seams that were not suitable for coal pro- poorly preserved plant remains, also occur. duction. In the southern part of the basin, the The upper part of the formation is rich facies is heterotypic, coastal, marine, with grey in molluscs, echinoderms, brachiopods and calcareous siltstones, fi ne- and coarse grained foraminifera, indicating a deeper marine ramp sandstones and marls. The uppermost section environment of normal salinity. The entire is subdivided into three parts: sequence suggests transgression and a grad- 3.1. The lower part, without coal seams, rep- ual deepening of the sea. The rise in sea-level resents marine facies with syngenetic deforma- and the resultant retreat of the shoreline may tions created by sandstones with crinoids and have caused a decrease in terrigenous input, “wormtracks”; because the delta system diverged from the 3.2. The middle part is a typical paralic suc- study area. cession of shales, aleurolites and sandstones According to Némedi Varga (1998) the coal- with thin coal seams; bearing succession can be subdivided into three 3.3. The upper part is of transitional type, major units and several subunits: with siltstones and sandstones. 1. The lowermost unit (Rhaetian) is divided Nagy & Nagy (1969) and Némedi Varga into two parts: (1995, 1998) interpreted this section as paralic, 1.1. The lower part showing laminated, open-lagoon, coastal and marine environments paludal (river channel – fl oodplain) facies: grey in a transgressive context. Nagy & Nagy (1969) to dark-grey siltstones, black shales and thin accepted the deltaic character of the deposits, coal seams. The overlying bed contains layers and suggested a river delta-plain leading to of a phyllopod Isaura hungarica Vadász. lagoon systems. 1.2. In the upper part of this section, a lim- The changes of facies in the sedimentary nic facies dominates, represented by nonlami- series vary at particular sites within the same nated green-grey siltstones with occasional locality (shafts and open mines, Fig. 3). The rootlet tracks, shale and subordinate sand- stratigraphic sequences may strongly differ stones. from each other at short distances as a result 2. The middle section (Hettangian) contains of the tectonic activity associated with strong the thickest coal seams; the coal production is subsidence, which produced fl exures and folds. the most intensive therein and almost entirely Consequently, data from different shafts are limited to them. They are strongly variable in often of local signifi cance only (e.g. see Nagy thickness and very diffi cult to follow because of & Nagy 1969). the presence of foldings, fl exures and reverse faults. The middle section is divided into fi ve FACIES IN WHICH FOSSIL PLANT levels: REMAINS WERE COMMON 2.1. Limnic level a, with thin coal seams and predominating siltstones and shales, sub- In the Mecsek Mts. the facies sequence ordinately sandstone intercalations; includes small cycles beginning from Early 2.2. Limnic level b, with dominant sand- Hettangian limnic, up to Early Sinemurian, stones, subordinately siltstones, without coal marine (Fig. 4). Plant remains are mostly asso- seams; ciated with a Hettangian paludal (river chan- 2.3. Limnic, with thick coal seams, domi- nel – fl ood plain) facies. nating siltstones, shales, sandstones (in some At Pécsbánya and Vasas this part of the areas), and a rhyolitic tuffi te layer at Pécs; sequence is the thickest (almost equally thick) 2.4. Limnic – paralic transitional level. and starts in the Hettangian, while at Komló Dominant siltstones – sandstones, without coal (Kossuth shaft) it begins in the early Sinemu- seams, containing a tuffi te layer with phyl- rian and narrows towards the NW of the sedi- lopods (Howellites princetonensis var. minor mentary basin. Nagy) indicating freshwater environments; Changes are not only observed in the thick- 2.5. Paralic transitional level with coal ness of the sedimentary cycle. Near Pécs- seams of medium thickness; dominant silt- bánya and Vasas deltaic and fl oodplain facies stones and shales. dominate, with regularly repeated swamp 131

Fig. 3. The Mecsek Mts., sequences of different sites (modifi ed, after Nagy 1969): 1 – overlying complex, 2 – coal seams with their serial numbers, 3 – tuffi te, 4 – underlying complex facies. In the Late Hettangian they are inter- cyclic swamp facies, and in the Early Sinemu- rupted by a thin marine facies followed then rian it turns into fl oodplain alternating with by fl oodplain. Near the Komló-Béta shaft, the swamp and lagoon. The Komló-Kossuth shaft deltaic facies are only in the lower half of the shows an increase in swamp facies alternating sequence (Hettangian), which also contains with fl oodplain. 132

Fig. 4. Facies in the Mecsek Mts. locality (modifi ed, after Nagy 1969)

PALAEOENVIRONMENT thus differences in plant content between par- ticular localities only partly depended on cli- The Mecsek delta plain (today’s coal fi eld matic zones and fl oristical provinces. The main area) was situated in the Jurassic at the lati- groups of plants, even at a generic level, were tude of today’s Spain (Csontos & Vörös 2004, present all over the world. It seems, however, fi g.23), the climate was generally stable, warm that plant specifi c diversifi cation observed at and wet during the whole period of the Jurassic the various fossil plant sites might be partly (Vakhrameev 1991). According to the global caused by local conditions like palaeogeographic palaeogeography, the main continents still position, altitude above sea level, intensity of formed one supercontinent (Golonka 2000), solar radiation, distance from fresh-water and 133 sea coasts, air humidity, moisture, type and Bóna (1963) proposed the fi rst, and still quality of soil. The taxonomic content of the used, palaeoenvironmental model with three plant cover depended on adaptation to these vegetation zones: conditions, and on the strength of competition (1) the swamp zone (dominant allochthonous between species. saccate conifer pollen grains The Mecsek Mts. fl ora grew in specifi c con- (2) the fl at marsh zone (allochthonous sac- ditions characterized by short-term cycles as cate conifer pollen grains and autochthonous a consequence of the dynamic character of the Calamospora (Sphenophyta) delta plain. Oscillation in the water level, the (3) the marshland forest zone (fern spores, fl ooded area, new areas formed by deposited and pollen grains from seed ferns, Podozami- sand, cyclic devastation and creation have taceae and Palissyaceae. made the territory interesting for the palaeo- A detailed palynofacies analysis focusing botanist (see also the fi gure in Thomas & Cleal on palaeoenvironment and palaeoclimate was 1993, p. 29). addressed for the fi rst time by Ruckwied et al. (2008). They proved that the palynomorph assemblages represent a typical Rhaetian/ PALAEONTOLOGY AND PALYNOLOGY Hettangian microfl ora. The Upper Rhaetian The coals generally contain monospecifi c part of the sedimentary series is dominated by bivalve assemblages of Unio vizeri Nagy bisaccate pollen grains, trilete spores and pol- accompanied by phyllopods, both indicat- len of the Circumpolles group, and by a small ing predominantly freshwater deltaic envi- amount of Corollina spp. Up-section, a strik- ronments. They alternate with subordinate ing increase in trilete spores, mainly species strata containing diverse gastropods and of the spore genera Concavisporites and Del- bivalves, indicating marine paralic environ- toidospora, was recognized. ments (e.g. Coelesina and Gryphea). The They also noted the sudden increase in bivalves from Pécsbánya are usually char- Dictyophyllum harrisii within the Triassic/ acteristic of lagoonal associations, except for Jurassic boundary interval, followed by a peak the two genera, Plagiostoma and Ceratomya, abundance of Inaperturopollenites spp. This which are characteristic of near-shore marine signal is repeated up-section in a small-scale environments (Szente 1995). In a vertical cyclic pattern characteristic of the entire Het- direction the coal deposits show a tendency tangian sedimentary series following swamp – towards transition from freshwater limnic or fl uvial changes. deltaic facies to lagoonal or marine paralic Within the small-scale cycles, river ele- facies, which are also heteropic facies at the ments of the palynomorph assemblage (Con- same time. baculatisporites mesozoicus, Concavisporites Among terrestrial vertebrates, the Gralla- spp., Deltoidospora spp., Stereisporites spp., tor-type footprint, Comlosaurus carbonis Kor- Todisporites spp., Vitreisporites pallidus) dom- dos, were found in both open mines, Vasas and inate in fl uvial sandstones, whereas swamp Pécsbánya (Kordos 1983, Ősi et al. 2011). elements (predominantly Dictyophyllum spp., The palynological investigations in the Inaperturopollenites spp.) are characteristic Mecsek Mts. locality that started in the 1960s, of the coal beds; deltaic sand- and siltstones were of interest to the mining industry, con- show a nearly equal amount of both groups, stituted a basis for the designation of the Tri- i.e. river and swamp elements, pointing to assic/Jurassic boundary and helped to corre- both the standing and agitated water bodies late the distinct coal seams (Bóna 1963, 1969, of the delta plain. 1979, 1983, 1995, Bóna et al. 1995). Based on Götz et al. (2011) made a palaeoecological rich palynological assemblages, Bóna (1983) reconstruction of the Mecsek locality accord- established that the coal deposits were Upper ing to the model proposed by Abbink (1998). Hettangian – Lower Sinemurian in age, and This model was discussed in relation with the intercalated between the Uppermost Triassic macro fl ora and its adaptation to different types (Rhaetian) strata and the coal measures of the of environment (see also page 162). Bucklandian Zone (Lower Sinemurian). 134

PART 2 the only NLE for Ginkgoales, has been exten- sively studied for its SI from different locali- Reconstruction of the Environment ties. Both dried material from herbaria, and Based on Plant Macroremains freshly collected material were studied (Royer et al. 2001), as well as samples growing under controlled experimental conditions in ambient PALAEOATMOSPHERIC CO2 and elevated CO2 (e.g. Beerling et al. 1998). COCENTRATION The species Ginkgoites marginatus from the Hettangian Thaumatopteris zone (Nagy 1961) Palaeobotanical climate proxy methods of the Mecsek Mts. was selected for stomatal complementing geological and geochemical measurements. According to Beerling and approaches in palaeoclimatic studies are now Royer (2002a), the SI is species-dependent, well-established (Beerling 1999, Poole & Kür- so data from the same species should be com- schner 1999, Mosbrugger 1999). The response pared. of plants to external conditions and their inher- Samples for investigation were restricted ent adaptation mechanisms make it possible to mainly to Pécsbánya, where 22nd horizon was monitor changes in atmospheric pCO2 on the most productive for macrofl oral remains. basis of leaf structures. In the last decades, The best preserved 18 samples of Ginkgoites a method that uses cuticle characteristics has marginatus from the Mecsek Mts. were stud- been developed. Stomatal parameters have ied. Ginkgoales are represented in the Hungar- proved particularly useful, because stomata ian locality by three genera and fi ve species; density is closely related to atmospheric com- their richness is mainly manifested mainly position and displays an inverse correlation to in the high numbers of specimens. The most pCO2 (Woodward 1987). common is Ginkgoites marginatus. Usually it An estimation of palaeoatmospheric CO2 has a cuticle well-preserved for determination, concentration is possible using parameters but for our study, the quantity of the material such as the stomatal density (SD), stomatal was limited to the best samples only. A lot of index (SI), and its relation to the nearest liv- leaves have cuticles partly damaged by pyrite, ing equivalent species (NLEs), defi ned as the which made the cell outlines unrecognizable; stomatal ratio (SR). A detailed discussion of other samples have cuticles too delicate for this method is provided by Beerling and Royer preparing satisfactory large pieces. The cuti- (2002a, b). cle pieces were collected in each case from the Numerous previous studies have investi- middle part of the leaf and prepared in Schulze gated the SD and SI for different plant groups solution followed by 3% KOH. In some cases, from different time intervals of Earth’s his- unprepared cuticle pieces were examined tory, from the oldest land plants (McElwain under a fl uorescent microscope. Only the lower & Chaloner 1995, McElwain 1998) up to mod- cuticle was used for examination. Epidermal ern plants. For the Mesozoic, mostly gymno- cells, including the subsidiary cells, were sperms have been used, including Cycadales counted according to the method described by (e.g. Ctenis and Nilssonia: McElwain Chaloner Jones & Rowe (1999) and Kürschner (1996). 1995, McElwain et al. 1999), Bennettitales The counting was done on microphotographs (Anomozamites and Pterophyllum: Steinthors- – for each sample as many counts as possible dottir et al. 2011), Pteridospermales (Steno- depending on the size and quality of cuticle: pteris: McElwain et al. 1999, McElwain 1998, this amounted to 116 measurements of 18 Lepidopteris, Tatarina, Rhachiphyllum: Retal- specimens. The investigated cuticle area was lack 2001, Bonis et al. 2010), Coniferales 300 × 300 μm = 0.09 mm2 (the minimum area (Brachyphyllum and Pagiophyllum: McElwain for SI calculation is 0.03 mm2 – Jones & Rowe 1998) and Ginkgoales (Baiera, Ginkgoites, 1999). This area was chosen because it was Ginkgo: Xie et al. 2006, Beerling et al. 1998). possibly the largest fragment of cuticle within Ginkgoales represent an especially suitable a stomatal band. SD was calculated from the group, because of its wide occurrence, com- same parts of cuticle. monly well-preserved cuticle and continuous SI was calculated according to Salisbury’s presence in global fl ora from the Palaeozoic equation (Jones & Rowe 1999) SI = [SD/ up to the present. The recent Ginkgo biloba, (ED+SD)] ×100, where SI – stomatal index, 135

SD – stomatal density (number of stomata/ may cause fl uctuation in estimations of palaeo area), ED – number of epidermal cells/area. CO2 level. To avoid inaccuracy during com- The atmospheric CO2 level was estimated as parison of different data, in the present paper 600 ppm × SR when SR is the Stomatal Ratio, (Table 3) the SI of G. biloba = 9.1 at 350 ppm i.e. SI of the nearest living equivalent divided was used after Beerling and Royer (2002a), by SI of fossil plant: SR = SI of NLEs/SI fossil. which is the nearest to the mean value of Carboniferous standardization was used after accessible records at 350 ppm.

Berner and Kothavala (2001). The gained SI and pCO2 level data were SI for the extant Ginkgo biloba was used compared with those from other localities according to Beerling and Royer (2002a), as 9.1 and time slices, measured on different species at 350 ppm. belonging to the Ginkgoales. The list of spe- The statistical test employed was the InStat cies and their SI is given according to Beer- 997: GraphPad InStat Demo, Version 3.00 ling and Royer (2002a), supplemented with for Win95/NT. GraphPad Software Incl., San data published later. For instance, measure- Diego. ments were completed on Ginkgoites mar- ginatus from Sweden and its SI estimated by RESULTS AND INTERPRETATION Beerling et al. (1998). The value of SI given by the authors for the Early Jurassic is 5.7, The mean value of stomatal density for which is a little higher than the value for the Hungarian specimens is 4.23, and the mean Hungarian specimens, but the difference is not SI = 4.96. signifi cant. There is also a record for the Late Based on the obtained SI values, the pal- Triassic, which shows a higher value (= 6.5), aeoatmospheric CO2 concentration was calcu- which denotes a slight increase in pCO2 at the lated and presented on Table 1. Triassic-Jurassic transition (see also Beerling et al. 1998, McElwain et al. 1999, Bonis et al. Table 1. Ginkgoites marginatus – SI, SD and SR values for Hungary; SD and SI counted for 0.09 mm2 cuticle areas 2010). Comparing all the data from Table 3, it can be observed that SI values of different Ginkgoites marginatus, Hungary species from the same localities correspond

number of specimens/ 18/116 well with each other. The level of pCO2 dur- measurements ing the Mesozoic fl uctuates generally between SI average (mean) 4.96 666 and 1162, except for two peaks at 2100 Std deviation 1.056 (China Middle Jurassic – G. yimanensis) and Std error 0.2488 Minimum 3.220 1606 (China E. Cretaceous – G. coriacea (Chen Maximum 6.328 et al. 2001). The pCO2 value for the Hungar- Median 5.053 ian Early Jurassic is placed among the higher Lower 95% CI 4.437 values above 1000 ppm in the curve given by Upper 95% CI 5.487 Beerling and Royer (2002a, p. 546). These data SD 2.30–6.30 also fi t into the curve of CO and temperature SD average 4.23 2 SR 1.83 trends over the past 300 million years given by Kürschner (2001). As shown on Table 2, the SI values obtained The Hungarian data confi rm the Hettang- from recent Ginkgo biloba by different authors ian climate of this locality as typical for this vary even at the same CO2 ppm level, which period, without signifi cant local oscillation.

Table 2. Ginkgo biloba, SI values according to different authors and possibly fl uctuation in pCO2 in Hungary estimated on the bases of these values

Beerling Beerling Royer et McElwain Chen et Retallack Retallack Beerling Beerling et al. et al. al. et al. al. (2001) (2001) (2001) & Royer & Royer (1998) (1998) (2001) (1999) 350 ppmv ppmv (2002) (2002) 350 ppm 560 ppm 560 ppmv 300 ppmv 350 ppmv SI of Ginkgo biloba 8.7 8.0 12.1 11.33 9.3 9.7 8.0 12.1 9.1 SR 1.75 1.61 2.44 2.28 1.88 1.95 1.61 2.44 1.83 palaeoatmospheric 1050 966 1464 1368 1128 1170 966 1464 1100 CO2 level 136

Table 3. SI values for the Mesozoic ginkgoalean species (on the basis of Beerling and Royer 2002), pCO2 level was estimated using Si of NLE Ginkgo biloba = 9.1 at 350 ppm (after Beerling & Royer 2002)

Species of the genus Study Locality Period SI CO level Ginkgoites 2

Beerling et al. (1998) G. troedssonii Sweden Late Triassic (Rhaetian) 5.9 925 G. marginatus Sweden Late Triassic (Rhaetian) 6.5 840 G. marginatus Sweden Early Jurassic 5.7 958 G. huttoni Yorkshire Middle Jurassic 5.6 975 McElwain et al. (1999) G. obovata Sweden Late Triassic (Rhaetian) 4.7 1162 G. obovata Sweden Late Triassic (Rhaetian) 6.8 803 G. acosmia Greenland Late Triassic (Rhaetian) 8.5 642 Chen et al. (2001) G. obrutschewii China Early Jurassic 6.7 815 G. yimanensis China Middle Jurassic 2.6 2100 G. huttoni Yorkshire Middle Jurassic 5.5 993 G. coriacea China Early Cretaceous 3.4 1606 Retallack (2001) G. matatiensis S. Africa Late Triassic (Carnian) 8.2 666 G. telemachus S. Africa Layte Triassic (Carnian) 7.6 718 G. lunzenzis Austria Late Triassic (Carnian) 6.7 815 G. troedssonii Sweden Late. Triassic (Rhaetian) 6.0 910 G. manchurica China Late Jurassic (K-Tit.) 7.4 738 G. manchurica China Early Cretaceous (Berr.) 5.0 1092 G. coriacea China Early Cretaceous (Val-Haut) 6.2 881 G. polaris Antarctica Early Cretaceous (Barr.) 6.7 815 Beerling & Royer (2002) G. dahlii Norway Middle Jurassic 4.5 1213 Xie et al. (2006) G. chilinense China Middle Jurassic. (Bajocian) 5.3 1030 G. obrutschewi China Middle Jurassic. E. Bajocian 5.9 925 G. huttoni China Early Jurassic. Aalenian 6.1 895 G. aganzhenense China Early Jurassic. (Pliensbachian) 5.8 941 present paper G. marginatus Hungary Early Jurassic (Hettangian) 4.96 1100

STATISTIC ANALYSES The present study is based on all leaf spe- cies from the Mecsek Mts. stored in the palaeo- botanical collection in the Hungarian Natural DATA COLLECTION AND THEIR DESCRIPTION History Museum, Department of Botany. The collection at the moment contains more than The fossil plant macroremains from the 5000 hand specimens. Hettangian–Sinemurian Karolinavöldi Coal The remains are represented mostly by com- Formation of the Mecsek Mountains were col- plete or incomplete leaves, but also by leafy lected from 1950 in the shafts of Béta, Kossuth shoots (conifers), fragments of axes, simple or and Zóbak, in the open mines of Pécsbánya, branched, seeds, roots, and occasional fructifi - Rücker and Vasas and the dumps of Budafa cations. The remains are distributed on rock and Dáviföld the Komló and Pécs regions, slabs in various directions, from one to several Southern Hungary (Fig. 1). They are found in on each slab; often slabs of the fi ne laminated layers between exploited coal lenses, whose siltstone are entirely covered by plant com- thickness varies from a few centimetres to pressions; the fossil leaves are also present in several dozen meters; they are usually pre- between rock lamellas. The number of differ- served in siltstone and its varied forms like ent taxa represented on particular slabs usu- grey siltstone, micaceous grey siltstone, brown ally varies from 1 to 3, but often rises to 5. siltstone with a high level of organic matter The aim of the study was to detect relation- and the form containing mica and foliated silt- ships between plant species in the community stone and the form with mica, foliated siltstone and their coexistence in ecosystems, clarify pos- with a high crystal sulphur content, very fi ne sible plant segregation into groups dependent grained siltstone and fi ne laminated siltstone on local landscapes, on the kind of rocks and and the form with crystal sulphur, and sand- on the site in the depository basin, with the aid stone. of numerical methods. The acquired data are 137 used for a model of the delta ecosystem. Per- MACROFOSSILS AND PLANT ASSEMBLAGE sonal observations during whole period of col- The basic character of the fossil fl ora of lection (1989–2003) confi rmed that the same the Mecsek Mts. was determined by environ- species in the same combinations repeated mental changes typical for delta plains: river- independently on level on particular sites. This side–swamp–lake–lagoon. Such alternations fact allows assumption that collected samples induced changes in the plant cover of the area. cover all depositional subtypes and assem- The ecosystem was infl uenced by both damag- blages of the Mecsek Coal Formation. ing and creating processes caused by fl uctua- Specimens used for the study were larger tions in the water level or changes in the riv- than 10 cm, and possibly represented more erbed. than one plant remain on a rock surface. Data Plant fossils have been sporadically col- were collected from both sides of slabs and lected in the Mecsek Mts. since the begin- they were treated separately. ning of the 19th century, when the coal min- The following features were recorded into ing started. Later, lists of plant fossils were the database: recorded in a number of publications (among (1) the site of collection (name of shaft, open others Stúr & Böck 1874, Hantken 1878, Klei- mine, dump); dorfer 1898, Hofmann 1907, Némedi Varga (2) the names of all species on the slab. In 1998, Wein 1952), as supplementary data for some cases, data only regard the generic level: the geological documentation needed for com- Sagenopteris is partly undeterminable due to its mercial mining. The fi rst short description of cuticle being poorly preserved, and the genus is the macroremains was given by Nagy (1961). generally quite uniform in environmental toler- The taxonomic revision of the described taxa ance, and Brachyphyllum was found in small and systematic work on the continuously col- amounts; both species, B. mamillare and B. cru- lected material has been carried out by Bar- cis, show no differences either in rock matrix or backa (1992, 1994a,b, 2001, 2002, 2009). in co-occurrence, so they are handled together. The remains mostly occur in the Hettang- Also, Pagiophyllum was used for analysis as ian deposits which are richest in coal seams, in a genus, since P. peregrinum is very common, between the coal seams, and are predominantly and occurs together with the rare P. ordina- preserved in siltstone, fi ne laminated siltstone tum. Coniopteris hymenophylloides, Sphenop- and, rarely, in sandstone. From a palaeoeco- teris sp., Pachypteris banatica, Pseudoctenis sp., logical aspect, the coal seams can be regarded Ctenis sp., and Geinitzia sp. were not included as one unit. Therefore, this is treated as one into data base for the analyses, because they locality with several sites (shafts, open mines occur only as single specimens not associated and dumps). with other taxa. Generally, the plant remains from the Mec- The state of preservation and fragmentation sek Mts. are characterized by quite a high of some ginkgoalean and equisetalean remains generic variability, while the species diversity made them undeterminable at a specifi c level, in each genus is low. Almost all the main plant but they were found in such large amounts, that groups occur in the fl ora; each group is repre- they cannot be omitted. They are indicated in sented by a number of genera, usually up to the material as indeterminable ginkgophytes 5, with the exception of the fern genera, rep- and Equisetites sp., independent of the deter- resented by up to 9. The number of species in mined species from those groups; each genus varies from usually 1 up to 3 in (3) length of remains. When the remain was only four genera: Cladophlebis, Sagenopteris, fragmented by rock breaking, but it was obvi- Nilssonia, and Brachyphyllum. Table 4 shows ously fossilized as a larger piece, the length is the complete list of taxa (leaves only). given as “more than” × cm. In the case of very dense and numerous fragments, the measure- SHORT CHARACTERIZATION OF THE FLORA ments were done on the fragments that were ELEMENTS IN THE MECSEK LOCALITY easily distinguishable and a length was given from × to y. The term “large” used in the text Horsetails are represented by two genera, always means a size over 6 cm in length. Equisetites (E. columnaris – Pl. 1, fi g. 1, and (4) type of remain (leaf, stalk, cone, etc.); E. muensteri) and Neocalamites (N. carci- (5) type of rock. noides, see Barbacka 2009). Both genera are 138

Table 4. List of leaf taxa described from the Mecsek Mts. so far

SPHENOPHYTA Equisetites columnaris (Brongniart) Phillips Equisetites muensteri Sternberg Equisetites sp. Neocalamites carcinoides Harris

PTERIDOPHYTA Marattiaceae Marattiopsis hoerensis Schimper Osmundaceae Cladophlebis denticulata (Brongniart) Fontaine Cladophlebis haiburnensis (Lindley et Hutton) Brongniart Cladophlebis roesserti (Schenk) Saporta Todites goeppertianus (Münster) Krasser Todites princeps (Presl) Gothan Matoniaceae Phlebopteris angustiloba (Presl) Hirmer et Hörhammer Phlebopteris sp. Dipteridaceae Clathropteris meniscoides Brongniart Dictyophyllum nilssonii (Brongniart) Göppert Dictyophyllum rugosum Lindley et Hutton Thaumatopteris brauniana Popp Dicksoniaceae Coniopteris hymenophylloides (Brongniart) Seward Sphenopteris sp.

PTERIDOSPERMATOPHYTA Caytoniales Sagenopteris hallei Harris Sagenopteris nilssoniana Brongniart Sagenopteris pilosa Barbacka incerte sedis Komlopteris nordenskioeldii (Nathorst) Barbacka Pachypteris banatica (Humml) Doludenko Ptilozamites cycadea (Ctenozamites cycadea) (Berger) Schenk

CYCADOPHYTA Cycadales Bjuvia simplex Florin Ctenis sp. Nilssonia obtusa (Nathorst) Harris Nilssonia polymorpha Schenk Nilssonia revoluta Harris Pseudoctenis sp. Bennettitales Anomozamites marginatus (Unger) Nathorst Pterophyllum subaequale Hartz

GINKGOPHYTA Ginkgoales Baiera furcata (Lindley & Hutton) Braun Ginkgoites marginatus (Nathorst) Florin Ginkgoites minuta (Nathorst) Harris Sphenobaiera leptophylla (Harris) Florin Sphenobaiera longifolia (Pomel) Florin

CONIFEROPHYTA Coniferales Elatocladus sp. Geinitzia sp. Cheirolepidiaceae or Brachyphyllum crucis Kendall Araucariaceae Brachyphyllum mamillare Lindley & Hutton Brachyphyllum papareli Saporta emend. Thévenard Cheirolepidiaceae Hirmeriella airelensis Muir et van Konijnenburg-van Cittert Pagiophyllum ordinatum Kendall emend. Harris Pagiophyllum peregrinum (Lindley et Hutton) Schenk emend. Kendall Podocarpaceae Podozamites lanceolatus Lindley et Hutton Podozamites sp. Incertae sedis Desmiophyllum sp. common and numerous specimens have sur- Neocalamites is rarer than Equisetites. vived as large fragments. In the case of Equise- It is supposed that it occurred in a different tites, besides stems, the underground rhizomes habitat, possibly dryer than Equisetites (Bar- together with roots in their growing position backa 2009). have been preserved, which suggests undis- Ferns are the richest group, being repre- turbed conditions in fi ne, muddy soil. sented by 14 species from fi ve families (see 139

Table 4). Except for the Dicksoniaceae, which fronds. Segments of C. meniscoides are very have a very fi ne structure and are small, ferns long, toothed, while T. brauniana has long, from the Mecsek Mts. have large, palmately narrow pinnules (up to 1.5 cm wide). dissected fronds (often exceeding 1 m) with The rarest are representatives of Dickso- long pinnate segments or toothed or incised niaceae, Coniopteris hymenophylloides (1 spec- margins; the species of Marattiaceae, Osmun- imen) and Sphenopteris sp. (17 specimens), daceae and Dicksoniaceae are pinnate. Usually and two species from Osmundaceae, C. roes- large fragments, or even whole fronds were pre- sertii (16 specimens), and T. princeps (14 spe- served, but they had always been fragmented cimens). The remaining two species, Todites during mining processes, or erosion. Thus, the goeppertianus and Clathropteris meniscoides specimens contain fragments of rachises with are very common, but the number of co-occur- pinnules, fragments of segments and often ring specimens is relatively low. complete leaf bases. Extended fronds with Pteridosperms are very widespread in the a large surface area correspond with humid Mecsek locality, although the number of their and warm conditions (van Konijnenburg-van taxa is low. The commonest is Komlopteris nor- Cittert 2002). denskioeldii (Pl. 2, fi gs. 5, 7), which is one of the Four genera are represented by more than dominant elements in the Mecsek community. one species and usually these species differ This seed fern has pinnate leaves about 20 cm signifi cantly from each other. In the genus in length with varied pinnules attached to the Cladophlebis, C. haiburnensis has the widest rachis, most often with a narrowed base. This pinnules, with entire margins and rounded type of adherence is quite fragile, but despite apices. Pinnules of C. denticulata (Pl. 2, fi g. 3) that, leaves are very often found whole, or as are elongated, relatively narrow and dentate rachis fragments with pinnules. Leaves of this in different degrees, apex acute or subacute. species often remained preserved as debris, C. roessertii (Pl. 2, fi g. 4) is the smallest, and in fi nely laminated aleurolite, also crowded its pinnules are narrow, elongated, shorter between lamellas. The structure of the leaves than 1 cm, margins entire and apex acute shows an adaptive specialization (shade (Barbacka & Bodor 2008). leaves with thin cuticle and sun leaves with Todites goeppertianus (Pl. 1, fi g. 2) and thick ones, Barbacka & van Konijnenburg-van T. princeps (Pl. 2, fi g. 1) differ in their whole Cittert 1998, Guignard et al. 2001), but the structure. The former is large and its fronds and general structure of its stomata suggests an pinnules resemble C. haiburnensis in size and adaptation to wet habitats (Barbacka 1994c). shape. T. princeps has small fi ne fronds with The cuticles vary in the presence and number minute pinnules (ca. 0.5 cm long). Reduced size of multicellular hair bases, whose structure can sometimes be a sign of arid circumstances, is very similar to the secreting glands (hyda- althrough C. hymenophylloides and Spheno- todes) of recent Gonocaryum (Icacinaceae, see pteris sp. certainly are not of a xeromorphic type. Hejnowicz 1980). Their function is connected The fronds of Phlebopteris angustiloba (Pl. 1, with the secretion of superfl uous water (Bar- fi g. 3) are not large, and the pinnules are very backa 1994b, Barbacka et al. 2010). Possibly, narrow (0.5 cm or less in width), and touching the plant was a tree (Barbacka & van Koni- each other. Phlebopteris sp. (Pl. 2, fi g. 2) has jnenburg-van Cittert 1998). larger fronds and more delicate, long, sparse The genus Sagenopteris (Pl. 3, fi g. 1) is rep- pinnules. resented by three species (Barbacka 1992), rec- The two species of Dictyophyllum differ in ognizable only by cuticle details. Its seemingly the same manner from each other. D. nilssonii compound leaves with a rather short petiole (Pl. 1, fi g. 5) has long segments with incisions, and usually four (rarely fi ve) nearly oval pin- while those of D. rugosum (Pl. 1, fi g. 6) are nules are rather small – the size of developed long and relatively narrow (ca 2 cm), elongated, leaf varies from 5 to 15 cm. The structure of more sparsely arranged and often incised. the leaf is always very delicate, the pinnules’ Their apex is acute. The lamina substance is place of attachment is very thin and easily thicker than in D. nilssonii. Two other genera breaks, but still complete or almost complete from the same family occur as a single species: leaves are often found in addition to numerous Thaumatopteris brauniana and Clathro pteris separate pinnules. The cuticle is always thin meniscoides (Pl. 1, fi g. 4). Both have large and the stomata have large and exposed guard 140 cells. Species from the Mecsek Mts. often have which has the largest leaves of the three spe- numerous hair bases. The cuticle structure cies. The length of its leaves is estimated at up suggests adaptation to high air humidity. to 30 cm (no whole leaf preserved), and width Ptilozamites (Ctenozamites) cycadea (Pl. 2, at up to 5.6 cm. The cuticle is very fi ne, with fi g. 6) is common. Its large, bipinnate leaves thin stomata with hollow papillae on subsidi- are quite strong and coriaceous. Small pin- ary cells and occasional hair bases. This spe- nules (up to 3 cm) join to the rachis with their cies shows features mostly related to wet con- whole base. The cuticle is thick, sometimes ditions. a little thinner (sun and shade leaves effect? Bjuvia simplex (Pl. 3, fi g. 3) is very common. Barbacka & van Konijnenburg-van Cittert Its very large leaves resemble those of recent 1998) and the stomata are strongly protected bananas. Because of their size, the leaves are by cuticle thickenings which restrict the sto- always found in fragments, sometimes form- matal pit to a very small opening. Guard cells ing debris. The cuticle is moderately thick, and are large and well cutinised. These features stomata are small and surrounded by a thick- are of a xeromorphic type. ened ring (sometimes papillae). The large Pachypteris banatica, excluded from the surface of the leaf blade, which is exposed to analyses, is very rare, almost unique (3 speci- allow intensive evaporation, signals rather wet mens found). The pinnate leaves have small conditions and is justifi ed by the need to pro- pinnules (up to 1.5 cm) with a thick cuticle; the tect the stomata, since the plant is large with stomata are varied, protected by papillae or by leaves placed on the top of a high stem (see a slightly thickened ring, with sunken guard reconstruction, Florin 1933, text fi g. 32). cells. The morphology and cuticle (presence of Pseudoctenis sp. and Ctenis sp. are very glands) suggest xeromorphic or halophytic tol- rare, only a few fragments being collected, so erance (Barbacka 1994 a,c). they were not used in the analyses. They have The most widespread cycad genera in the pinnate leaves with a rather thick cuticle and Mecsek locality are Nilssonia and Bjuvia (Bar- thickenings around the stomatal pit. backa 2001). All species of Nilssonia, N. poly- Bennettites are only represented by two morpha (Pl. 3, fi g. 2), N. obtusa and N. revoluta species belonging to two genera: Anomozamites have elongated leaves with entire or almost marginatus and Pterophyllum subaequale. entire margins (N. polymorpha is more or less They are not numerous in the locality and irregularly pinnate). This type of leaf grows in their leaves are mainly found as large frag- clusters from short branches (Nilssoniocladus ments. Their leaves are pinnate with narrow Kimura & Sekido). In most cases they are found segments and both species are rather small in as complete leaves; only the largest species has their categories. Their cuticles are moderately fragmented remains. The leaves often form thick and do not show any special adaptive fea- debris in fi ne laminated aleurolite, frequently tures. Usually bennettites are connected with crowded between laminas. These plants were rather dry conditions (Abbink 1998, Abbink probably of the shrub type (see reconstruction et al. 2004). of similar species, Watson & Cusack 2005, text Ginkgophytes are very numerous (Barbacka fi g. 33). Species from the Mecsek Mts. differ 2002); especially abundant are Ginkgoites mar- from each other in their size and with regard ginatus (Pl. 3, fi g. 4) and Baiera furcata (Pl. 3, to their cuticles. The smallest is N. revoluta, fi g. 6). Sphenobaiera longifolia (Pl. 3, fi g. 5) whose leaves are narrow (less than 1 cm) and and S. leptophylla are less common. In most do not exceed 9 cm in length. Its cuticle is cases, the leaves were preserved complete; even thick and stomata small, surrounded by a cuti- leaves of such fi ne plants as Sphenobaiera lep- nised ring. These features may be regarded as tophylla or S. longifolia. Ginkgophytes in the xeromorphic. N. obtusa is of medium size, its Mecsek often form debris, but even in these leaves are up to 17 cm long, and the cuticle is cases, fragmentation is not strong. thinner than in N. revoluta. The stomata have All taxa from this locality are character- thicker subsidiary cells than ordinary cells ized by relatively narrow segments (2–5 mm (sometimes papillae) and numerous one-cell in different taxa). The broadest segments hair bases occur on the cuticle. This species belong to Ginkgoites marginatus, up to 17 mm shows transitional features between xeromor- in the form with two lobes. They have thin to phic N. revoluta and the fi nest N. polymorpha, moderately thick cuticles with papillae on the 141 subsidiary cells or at least thickenings around categories of size/type of preservation for the the stomatal pit in all species, sometimes also fourteen most abundant species (over 30 speci- on the ordinary cells. mens), from particular groups. The following Ginkgoites minuta, not used in the analy- categories were considered: large fragments ses, is very rare, being found only once, and all (above 6 cm), debris (numerous fragments the leaves came from the same slab. Its mor- between 2–6 cm), litter (a crowd of large frag- phology differs from other ginkgophytes from ments or complete/almost complete leaves, in this locality in its size (it is much smaller), between rock lamellae as well), and small frag- thicker cuticle and in the presence of resin ments (separate, below 6 cm). Leaves, which bodies. In all the other species, resin bodies may have been preserved complete, i.e. Kom- were not observed, even in Sphenobaiera lepto- lopteris or Sagenopteris, have additional cate- phylla, which was described by Harris (1935). gories like “complete” or “pinnae”, and ginkgo- It seems that their presence may depend on phytes have a category for “segments”. the environment (Barbacka 2002). The state of preservation of macroremains The conifers are strongly dominated by proves that most of them are autochthonous Elatocladus sp. (Pl. 3, fi g. 9). The genera or parautochthonous in origin and there were Brachyphyllum, represented by three species good conditions for fossilization (the presence (B. mamillare, B. papareli and B. crucis), as of cuticles and their good state). As it issues well as Pagiophyllum (Pl. 3, fi g. 8) with its two from Fig. 5A-N the autochtonic character of species (P. peregrinum and P. ordinatum) are the fl ora may be proved by: not very common, but often co-occurred with a. domination of large and undamaged frag- other taxa. They are commonly preserved in ments large shoot fragments, often branched, but in b. fragile attachments of pinnules in compo- debris the fragments can be rather small (1–2 und fronds or leaves are in most cases unda- cm or even separated leaves). Their morphol- maged ogy is different: Elatocladus sp. has needle- c. conifer shoots (mainly Elatocladus) are like leaves of very fi ne structure; its cuticle is very often preserved in branching state and thin and unsuitable for preparation. It is usu- often with male or female cones attached ally interpreted as living in wet conditions. d. occurrence of underground rhizomes of Brachyphyllum and Pagiophyllum have thick Equisetites with thin roots in growing position and small leaves adjacent to their shoots, their e. predominance of monospecifi c debris in cuticle is thick and sunken guard cells are pro- which leaves are often complete (Ginkgoales) tected by a thickened cuticular ring suggest- or fragments are relatively large (e.g. sepa- ing the xeromorphic character of these plants rate, but entire pinnules). (Thévenard et al. 2005). Some other coniferalean taxa are present in MULTIVARIATE ANALYSIS the locality, like Geinitzia sp. and Podozamites schenkii (Pl. 3, fi g. 7). Both genera are fi ne in The data encompass the species composition structure, having a thin cuticle unsuitable for of 1122 palaeontological samples, which were preparation. Geinitzia (not used in the analy- classifi ed into types of rock substratum accord- sis) is found as branched shoots and, similarly ing to the geological criteria. Generally, 37 spe- to Elatocladus, Podozamites is usually preser- cies of plant and 10 types of rock substratum ved as rather large fragments, in contrast to were recognized. The occurrence of species in its fi ne shoots and leaves with narrow bases. samples was coded in a nominal, binary 0–1 They are usually associated with wet conditi- scale, i.e. as absence/presence data. ons (Harris 1979). Additionally, each sample was character- ized regarding the type of rock substratum. DEGREE OF FRAGMENTATION, Nominal variables (types of rock substratum) AUTOCHTONOUS ORIGIN OF THE REMAINS were represented by a series of dummy vari- ables each representing a category yik = 1 or 0, High numbers of samples allow the analysis depending on whether sampling unit i belongs of different degrees of remains fragmentation, or not to category k. The data set prepared in which is inevitable even in autochthonous or the form of data matrix was then used for the parautochthonous fl ora. Figure 5A-N shows clustering and ordination. 142

Fig. 5A-N. Degree of leaf fragmentation in selected species

Clustering nominal, binary 0–1 scale, forming a synthetic table. Then, the sites were classifi ed accord- The palaeontological samples originated ing to the species occurrence. Additionally, the from 7 sites (Fig. 1). In each site the occurrence relationships between the type of substratum of species and the type of rock substratum were and species were coded in the nominal, binary evaluated, and their presence was coded in the 0–1 scale and displayed in the form of a data 143

Fig. 5. Continued matrix. The matrix was then used for the Ordination classifying of species and rock substratum in respect of their occurrence in particular sites. To display the relationships between the A UPGMA algorithm (Sokal & Michener 1958) samples and species, and to show joint occur- was used, implemented in the NTSYSpc pro- rence of species in the samples, a Detrended gram (Rohlf 2000–2005). Correspondence Analysis was used (DCA – Hill 144

Gauch 1980). This is a heuristic modifi cation as well as groups formed by both the fi rst and of CA design (Hill 1973) to remedy both the second axes. edge effect and the arch effect, which distort In the fi rst step, out of the 37 examined spe- the original distances among the samples. cies, six species were removed by the used pro- The edge effect is removed in DCA by non- gramm as outliers: Clathropteris meniscoides, linear rescaling of the axis. The arch effect, Todites goeppertianus, Todites princeps, Podo- a more serious problem in CA, is removed in zamites schenkii, Podozamites sp. and Neoca- DCA by the heuristic method of “detrending- lamites carcinoides, thus the fi nal analysis was by-segments”, which was used in the present carried out on 31 species. analysis. DCA is popular, especially in fi eld The lowest scores on Axis 1 are possessed ecology; however it may be used in all types by Nilssonia revoluta, Cladophlebis denticu- of data, given that they are reasonably repre- lata and Anomozamites marginatus, forming sentative of sections of the major underlying a distinct cluster, together with Sagenopteris environmental gradients. The main premise is sp., Marattiopsis hoerensis and Nilssonia that each species displays a unimodal (Gaus- obtusa (Fig. 6). The position at the opposite sian) distribution in two or more dimensions, extreme along Axis 1 is occupied by Equisetites defi ned by the specifi c species’ responses to muensteri. The gradient along Axis 1 divides the hypothetical environmental gradients the plant species into three clusters (see also (Ter Braak & Prentince 1988). In particular, Table 5). in the context of the present study, it means On Axis 2, the lowest DCA score is possessed that groups of species had similar patterns by Thaumatopteris brauniana, and Nilssonia of ecological response to the underlying envi- polymorpha shows the highest value. The gra- ronmental gradients, and that the patterns dient along the axis also groups species into have been preserved, or at least not seriously three clusters, which occupy equal parts of the disturbed, in the palaeobotanical samples. axis (Fig. 6). The groups of species with the uniform type Axes 1 and 2 enable a distinction to be of response (deduced from their joint occur- made between fi ve groups of plants, includ- rence) are considered as having similar eco- ing the monospecifi c Ginkgoites marginatus logical requirements, thus forming ecological group (Fig. 6, Tab. 5). The percentage of plant groups. groups in the particular samples is shown in The length of gradient for DCA Axis 1 Fig. 7A–D. amounted to 10.19 standard deviation, and the percentage of variance accounted for 4.0% INTERPRETATION OF RESULTS and 3.7% of the variance of DCA Axis 1 and AND DISCUSSION Axis 2, respectively. The inspection of the DCA results showed that fi ve species were outliers DCA Ecogroups (see Results). The length of gradient of the reduced data set for DCA Axis 1 amounted to The DCA ordination provides an oppor- 8.06 standard deviation, and the percentage tunity for the study of species co-occurrence of variance accounted for 4.6% of the Axis 1, within the same samples, and, assuming the and 4.3% of the Axis 2. The samples that dif- autochthonous or parautochthonous character fer by 4 standard deviations are to be expected of the fl ora, it may be interpreted as their joint to have no species in common. Total inertia occurrence in similar ecological conditions. equaled to 19.494. The distribution of species Taxa showing the closest association along the along the DCA Axis 1 and Axis 2 is displayed Axes 1 and 2 are grouped into fi ve clusters, in Figure 6. i.e. fi ve putative ecological groups. The groups are named from the dominant taxon and show their centre of occurrence along the DCA Axis 1 RESULTS and 2 (Fig. 6A). The DCA analysis was applied for species The many factors which may determine co-occurrence and separation of plant groups the origin of plant assemblages include: pal- presumably growing together in the same eco- aeogeographic position, altitude above sea logical niches. Fig. 6 shows the obtained dis- level, marine transgressions and regressions, tribution of species along Axis 1 and Axis 2, distance from the sea coast (infl uence of salty 145

Fig. 6. A. DCA: ecogroups, B – ecogroups with extended niches water), distance from a fresh-water reservoir, and only local environmental changes are con- air humidity, type of soil, its moisture and sidered as a possibility. quality, succession and competition. Considering the delta environment and Since the taxa are in this case from the changes typical of such a landscape, the most same area, the fi rst two factors may be omitted probable factors strongly infl uencing changes 146

Table 5. Groups of taxa obtained in DCA analysis according to Axis1, Axis 2 and Axes 1/2 (following the gradient)

Axis 1 Axis 2 Axes 1/2

1 Nilssonia revoluta Thaumatopteris brauniana Nilssonia revoluta Cladophlebis denticulata Phlebopteris angustiloba Anomozamites marginatus Marattiopsis hoerensis Dictyophyllum rugosum Cladophlebis denticulata Sagenopteris sp. Equisetites sp. Marattiopsis hoerensis Nilssonia obtusa Sagenopteris sp. Anomozamites marginatus Nilssonia obtusa

2 Thaumatopteris brauniana Nilssonia revoluta Thaumatopteris brauniana Phlebopteris angustiloba Cladophlebis denticulata Phlebopteris angustiloba Dictyophyllum rugosum Sphenobaiera leptophylla Equisetites sp. Equisetites sp. Elatocladus sp. Dictyophyllum rugosum Cladophlebis haiburnensis Cladophlebis haiburnensis Cladophlebis haiburnensis Phlebopteris sp. ginkgophytes Desmiophyllum sp. Baiera furcata Bjuvia simplex Marattiopsis hoerensis Dictyophyllum nilssonii Sagenopteris sp. Ptilozamites cycadea Equisetites muensteri Pterophyllum subaequale Komlopteris nordenskioeldii Equisetites columnaris Pagiophyllum sp. Nilssonia polymorpha Brachyphyllum sp. Sphenobaiera longifolia 3 unidentifi ed ginkgophytes Nilssonia obtusa Nilssonia polymorpha Komlopteris nordenskioeldii Phlebopteris sp. Equisetites columnaris Ginkgoites marginatus Desmiophyllum sp. Pterophyllum subaequale Elatocladus sp. Anomozamites marginatus Ptilozamites cycadea Baiera furcata Bjuvia simplex Dictyophyllum nilssonii Sphenobaiera longifolia Ginkgoites marginatus Bjuvia simplex Pagiophyllum sp. Dictyophyllum nilssonii Desmiophyllum sp. Sphenobaiera leptophylla Ptilozamites cycadea Phlebopteris sp. Brachyphyllum sp. Pterophyllum subaequale Equisetites muensteri Equisetites columnaris Nilssonia polymorpha 4 Ginkgoites marginatus 5 Komlopteris nordenskioeldii Elatocladus sp. Baiera furcata Sphenobaiera longifolia Pagiophyllum sp. Brachyphyllum sp. Sphenobaiera leptophylla Equisetites muensteri in fl oristic composition are the moisture gra- both axes fi ts in to a curve of normal distri- dient of (Axis 1) and degree of environmental bution with lower extreme values and higher stability (Axis 2). The degree of disturbance moderate values (Fig. 8A wetness gradient may infl uence fl oristic changes detectable in – blue and stability gradient – red). It corre- the fossil state in two ways: by selection of sponds well with ecological rules, when the taxa that adapt to given conditions and by most diverse populations are connected with plant succession stopped at different levels moderate conditions, while extreme conditions by fl ood or other catastrophic events, which is need higher specialization, which restricts manifested in the contents of particular groups the number of taxa. It implies from Fig. 8A and adaptation of their components. that in both cases (Axes 1 and 2) the dry and Presuming that in a delta plain a large ter- disturbed conditions characterised by low val- ritory is well watered, the gradient of Axis 1 ues restricted the number of taxa to a higher possibly follows humidity in the present case degree than in case of the opposite values, i.e. from the relatively less wet regions to wet, wet and more stable. swampy zones. The morphology and cuticular The quantitative contribution of particular structure of plants change along this axis in groups in the plant cover displays the reverse correspondence with this supposition. trend (Fig 8B). The Komlopteris group and the The appearance of the groups in relation to Sagenopteris group (they show the opposite 147 tendencies on Axis 1) are most rich in speci- mens. The groups from the medium values of Axis 1 are not so numerous. If the biomass is proportional to the extension of environments, it would mean that in the Mecsek delta ter- ritory, fl oodplains and swamps dominated, as well as more drained territories, while highly

Fig. 8. A. Distribution of taxa amount along Axis 1 (wetness gradient – blue) and 2 (stability gradient – red) according to DCA; B – Distribution of specimens amount along Axis 1

disturbed or the most stabile, moderated wet areas were smaller, although more diverse.

DCA – Extension of Niches The segregation of taxa according to the extent of their ecological profi les is in comp- liance with the given environmental circums- tances, so they form distinct ecogroups. Axes 1 and 2 form a space in which particular taxa are distributed according to their putative ecologi- cal adaptations (Fig. 6B). It is worth noticing that, although all the ecogroups tend to extend and overlap with each other, none of the taxa extended along the entire gradient either that of Axis 1, or that of Axis 2. Additional DCA analyses for particular species give more detailed information on their predilections and show the scope of their environmental tolerance, i.e. may defi ne their niches (Figs 9−13). It is natural that overlap- ping niches do not fully follow the groups sepa- rated on the grounds of maximum occurrence of taxa and that taxa with wide niches can occur in more than one ecogroup. The widest niches were occupied by 5 species: Komlop- teris nordenskioeldii, Elatocladus sp., Sagen- Fig. 7. A-D. – Percentage of taxa in particular ecogroups opteris sp., Phlebopteris sp. and Dictyophyllum 148 nilssonii, while Brachyphyllum sp. and Equi- ginkgophytes and cycads do not combine with setites columnare show the narrowest niches. each other. The niches of Sphenobaiera leptophylla and Most combinations show both species of Nilssonia revoluta are not very restricted, but Dictyophyllum, Phlebopteris sp., Komlopteris quite distinctive and only overlap each with nordenskioeldii and Elatocladus sp., and also other to a minimal degree. Desmiophyllum sp. (see also Fig. 14). Taxa reaching the opposite sides of the Axis 1, presumably representing the moisture gradient, belong to completely different types DESCRIPTION OF RECONSTRUCTED of environment, and Nilssonia revoluta and LANDSCAPES Brachyphyllum sp., in particular, are mutu- ally exclusive. Nilssonia revoluta occupies an Based on the DCA ecogroups, the following exceptional niche, since it extends to presum- environmental types were distinguished within ably drier and more disturbed surroundings the delta landscape of the Mecsek locality: (Fig. 9), while the narrow occurrence of Brach- yphyllum sp. (Fig. 12) can be explained by 1. Moderately disturbed, relatively dry its high specialisation. Equisetites muensteri, (non-fl ooded) inland areas, which correspond which expands to maximally wet conditions, to the occurrence of the Sagenopteris group. has a wider adaptation and co-occurs with more taxa (Fig. 12). Dominant element. Sagenopteris sp. Some species with xeromorphic features, This habitat mostly seems to correspond like Brachyphyllum sp., Pagiophyllum sp., to the “upland” widely applied in palaeoeco- Ptilozamites cycadea or Bjuvia simplex, show logical reconstructions, here used according their maxima in presumably wet environment, to Batten's defi nition (Baten 1974, p. 446), in but their niches extend towards dry habitats that case probably only minimally raised, yet (Figs 12 and 11 respectively). On the other high enough not to be fl ooded. In this habi- hand, xeromorphic features occur not only tat, taxa are highly spread: from dry and dis- in ancient plants living in strict arid condi- turbed conditions favouring Nilssonia revoluta tions, but also in semi-arid sites, which are and Cladophlebis denticulata, passing towards seasonally or periodically dry (Batten 1974). areas of higher moisture, where N. obtusa and In the recent biomes, several plants living in Marattiopsis hoerensis began to appear, while wet conditions, e.g. in high peat bogs, also Sagenopteris sp. and Anomozamites margina- displayed xeromorphic features due to physi- tus were attracted to conditions of heightened ological dryness. stability. The majority of species listed in this group Taxa co-occurrence based justifi es their position on the Axis 1 as rep- on extended niches resenting the driest conditions in the Mecsek Based on the range of niches, all possible ecosystem. Nilssonia revoluta has the narrow- co-occurrences of taxa within the Mecsek delta est leaves among the Nilssonia species with were determined (Table 6). entire margins, and possess bulges on the sub- The amount of possible combinations of sidiary cells; features which were interpreted taxa is high, but certain tendencies are notice- by Harris (1964) as xeromorphic. Its niche able. Most strikingly, none of the ferns, cycads is rather narrow and within the drier inter- and bennettites occurs together with Brachy- val extends towards the more disturbed con- phyllum. Cladophlebis denticulata does not dition (Fig. 9). Cladophlebis denticulata has co-occur with any ginkgophytes, similarly to relatively narrow and elongated pinnules with Phlebopteris, although the latter may occur dentate margins. Its niche continues towards with Ginkgoites marginatus. Thaumatopteris more dry and disturbed circumstances (Fig. 9). brauniana does not co-occur with bennet- Anomozamites marginatus, a bennettite with tites, and the majority of ginkgophytes. Also, thick, small and short pinnules and Nilssonia Nilssonia revoluta avoids ginkgophytes; the obtusa with its moderately thick cuticle and other cycads co-occur only with G. marginatus. narrow entire leaves also corresponds with the Generally, besides Brachyphyllum, which has suggested type of environment. Its occurrence exceptionally few combinations, some ferns, expands towards a less disturbed section. 149

Fig. 9. DCA for species from Sagenopteris group

Fig. 10. DCA for species from Thaumatopteris group 150

Fig. 11. DCA for species from Ptilozamites group

What is surprising is the presence of Marat- places with high air humidity, and not neces- tiopsis hoerensis and Sagenopteris sp., which sarily a high concentration of moisture in the are characterized by rather delicate leaves. soil. This way they may be imagined as under- Sagenopteris has a thin cuticle and exposed storey, or a climbing form, in the case of Sage- guard cells; Marattiopsis is usually connected nopteris. The suggested shrub or tree appear- with warm, moist circumstances (van Koni- ance of the latter (Harris 1964, Gordenko jnenburg-van Cittert 2002, Wang 2002). Its 2008) would have been more justifi ed if sun recent relative Marattia grows in the shaded leaves had been found. The described trunk of fl oors of wet tropical forests (Thomas 1985), Sagenopteris does not exclude it from a climb- and also often along river banks (van Konij- ing life style (e.g. extant genus Hedera). This nenburg-van Cittert pers. com.) which is here fact would explain such a wide co-occurrence confi rmed by its niche stretching towards wet- of Sagenopteris, always living in the shadow ter and more disturbed environments. The of other plants. range of Sagenopteris is much expanded, and The other taxa, which are possible in this also occupies territories, which are variably type of environment because of the range of disturbed, reaching moderately wet areas. The their niches, are: Ptilozamites cycadea, Bjuvia structure of both the genera Marattiopsis and simplex, Cladophlebis haiburnensis, and Dic- Sagenopteris suggests a preference for shaded tyophyllum rugosum. 151

Fig. 12. DCA for species from Komlopteris group

Fig. 13. DCA for Ginkgoites marginatus 152 brown – gikgophytes, grey conifers. Table. 6 . Taxa co-occurrence based on extended niches. Explanation of colours: dark blue – sphenophytes, light brown ferns, green seed ferrns, violet – cycads, blue bennettites, Pagiophyllum Brachyphyllum Elatocladus Desmiophyllum Baiera furcata Sphenobaiera leptophylla Sphenobaiera longifolia Ginkgoites marginatus Pterophyllum subaequale Anomozamites marginatus Bjuvia simplex Nilssonia revoluta Nilssonia obtusa Nilssonia polymorpha Ptilozamites cycadea Komlopteris nordenskioeldii Sagenopteris Thaumatopteris brauniana Dictyophyllum rugosum Dictyophyllum nilssonii Phlebopteris Phlrbopteris angustiloba Cladophlebis denticulata Cladophlebis haiburnensis Marattiopsis hoerensis Equisetites muensteri Equisetites columnaris sp. sp. sp. sp. sp. sp. +++++++++++++++–+++++++++++ +++++++++++++++++++++++++–+ ++–+––+–+––+––––+–+++++++++ ++++–++++–+++++–+++++++++++ +–+++++++–+++++–+++++–+++–+ +–+++++++–++++++++++–––++–– +++++–+++–+++++++++++++++–+ +++++++++++++–++++++–––++–+ +–+–––+++–++++–+++++–––++–+ +–+–+–+++–+++++–++++–––++–– +++++++++++++++–+++++++++++ ++++++++++++++++++++––+++–+ +++++++++++++++–++++–++++–+ +++++++++++++++–+++++++++–+ +–++++++++++++++++++––+++ –+ +–+–+–++–_+++++–+++++––++–– –+–––––––––+–––––––++–+–+++ –+++––++++++––––+–+++++++++ –+–+––+++––+––––+––++++++–+ ––+++–––+++––+++++–––––+––– ––++++++++++––++––––––+++–– –++++++++++++++++++++++++–+ –+++––+++–++–++–+–+++++++++ ––++++++++++––+––+++–––++–– –++++++++++++–+++++––––++–– –+++++++++++––+++++++++++–+ –+–++–+++–++––+–+––++++++++ Equisetites columnaris Equisetites muensteri Marattiopsis hoerensis Cladophlebis haiburnensis Cladophlebis denticulata Phlebopteris angustiloba

Phlebopteris sp.

Dictyophyllum nilssonii Dictyophyllum rugosum Thaumatopteris brauniana

Sagenopteris sp.

Komlopteris nordenskioeldii Ptilozamites cycadea Nilssonia polymorpha

Nilssonia obtusa

Nilssonia revoluta

Bjuvia simplex

Anomozamites marginatus Pterophyllum subaequale Ginkgoites marginatus Sphenobaiera longifolia Sphenobaiera leptophylla

Baiera furcata

Desmiophyllum sp.

Elatocladus sp.

Brachyphyllum sp.

Pagiophyllum sp. 153

Taxa that were not used in the analyses but as indicators of seasonally dry vegetation could be attributed to this habitat are: Todites (Falcon-Lang et al. 2009). Cheirolepidiaceae goeppertii, Todites princeps, and Clathropteris are often assigned to drier habitat (including meniscoides. Only 3 specimens of Todites goep- the seasonally dry) as drought resistant, ther- pertianus (total number 69) were associated mophilous plants (e.g. Batten 1974, Nguyen with Nilssonia obtusa. Therefore, since the et al. 1999, Abbink 1998, Abbink et al. 2004). niche of N. obtusa extends from medium wet Other groups often connected with dry envi- to drier habitats, it is probable that T. goepper- ronments are cycads and bennettites (Batten tianus also belong to this group. This is justi- 1974, Abbink et al. 2004). fi ed by the presence of Cladophlebis, which is Caytoniales, which in the Mecsek Mts. are closely relative to Todites (Harris 1961). It is at their maximum extent in rather dry condi- also probable that its niche extended towards tions, appeared to have a very wide tolerance. more disturbed areas, where they formed mon- In a deltaic environment, they are usually pla- ospecifi c thickets. ced into transitions between the fl oodplain and No specimens of Todites princeps were asso- backswamp peats, or moist, lush, generally ciated with other plants. The small amount of warm and wet habitats (Abbink 1998, Abbink these in the locality suggests a narrow niche. et al. 2004, Hesselbo et al. 2003). However, The morphology, although fi ne, suggests adap- Thorn (2001) interpreted many Caytoniales as tation to rather dry conditions, according to its occurring in dry habitat with conifers, arbores- relatively thick rachis and rigid appearance cent ferns, lycopods and bryophytes. (see also Barbacka & Bodor 2008). Schweitzer Ferns and sphenophytes are not considered (1978) discussed the lifestyle of this spe- as typical of dry conditions, but are some- cies as possibly being epiphytic (after Harris times mentioned (Batten 1974). For the ferns, 1961) or living on periodically dried clay (after this mostly regards Matoniaceae, which have Weber 1968). Its strongly distinct occurrence leaves protected by a resistant cuticle (van in the Mecsek locality rather suggests the last Konijnenburg-van Cittert & van der Burgh solution. 1996, Abbink et al. 2004, van Konijnenburg- The majority of Clathropteris meniscoides van Cittert 2002). They mentioned species of did not associate with other taxa, but some Phlebopteris as being tolerant to semiarid con- samples showed co-occurrence (18% of studied ditions, similarly to Hesselbo et al. (2003), who specimens). Single fragments occurred with stressed that the same was true for Osmun- Anomozamites marginatus, Sagenopteris sp., daceae. Phlebopteris sp., P. angustiloba, Elatocladus sp., Thaumatopteris brauniana, Dictyophyllum 2. Highly disturbed short-lived, modera- rugosum, and Komlopteris nordenskioeldii, but tely wet territories formed by alluvial deposits it was mostly associated with Nilssonia obtusa (islands, peninsulas, forelands), fully damaged (10 specimens). Considering the types of niche by river fl ood, occupied by pioneer plants from characteristic of these species, the niche of Cla- the group Thaumatopteris. thropteris meniscoides could extend to medium wet and medium disturbed habitats. The dominant species. Thaumatopteris Four selected taxa were analysed from brauniana. a taphonomical point of view: Sagenopteris The Thaumatopteris group exclusively con- sp. (Fig. 5B), Clathropteris meniscoides (Fig. sists of Sphenophyta and Pteridophyta. Taxa 5L), Todites goeppertianus (Fig. 5H), Nilsso- of this group show very close moisture values, nia obtusa (Fig. 5G). All of them show signifi - but are spread along the stability gradient of cant dominance of large fragments; in the case the environment. Thaumatopteris brauniana, of Sagenopteris separated pinnae are most Equisetites sp., Dictyophyllum rugosum, and abundant. The amount of debris specimens Cladophlebis haiburnensis show their maxi- is not very high, and production of litter was mal appearance in this habitat. Other possi- rather low. ble taxa whose niches reach these territories R e m a r k s. Generally conifers are often con- are also mainly pteridophytes: Phlebopteris nected with drier conditions (Batten 1974, angustiloba, Cladophlebis denticulata, Dic- Kustatscher et al. 2010, Abbink 1998, Abbink tyophyllum nilssonii, Phlebopteris sp., Maratti- et al. 2004, Wang et al. 2005) and also treated opsis hoerensis, and Clathropteris meniscoides. 154

Fig. 14. Degree of co-occurence of particular taxa: A – according to taxonomy, B – in order of range of charge

The composition of this assemblage is typical preserved in large fragments with a minimal of disturbed areas, since pteridophytes and tendency for production of debris or litter (Fig. sphenophytes, which are here the main ele- 5 I). Other species of ferns (compare Fig. 5 H, ments, tend to occupy new territories as pio- J, L) also show the same type of fragmentation. neer plants, and often occur in a stage of early In this type of habitat, when areas are quickly succession (Thomas 1985). From the other covered by water and sand, fragmentation of plant groups, Komlopteris nordenskioeldii, such a type is not remarkable. Observations Sagenopteris sp., Baiera furcata, and Elatocla- of the Hungarian records correspond with the dus sp. tend to appear in this early colonized Iranian ones that were regarded by Schweitzer stage as well (or as the next succession stage (1978) as forming extended thickets, or in asso- in these areas). ciation with Dictyophylum nilssonii or species Coniopteris hymenophylloides and Sphenop- from the genus Sagenopteris. teris sp., which were not used in the analyses, always occur alone. Their morphology, slender R e m a r k s. By analogy with their contem- rachises and fi ne pinnules suggest medium porary species, ferns and sphenophytes may wet to wet habitats. often be regarded as pioneer plants coloni- Thaumatopteris brauniana was mainly zing disturbed habitats. Bomfl eur and Kerp 155

(2010) discussed this predilection with regard conditions. These are: Ginkgoites margina- to Dipteridaceae. They concluded that „Clath- tus, Pterophyllum subaequale, Ptilozamites ropteris/Polyphacelus locally became dominant cycadea, Dictyophyllum nilssonii, D. rugo- elements during the early phase of colonization sum, Cladophlebis haiburnensis, Phlebopteris of disturbed sites after catastrophic volcanic sp., Sagenopteris sp., and Desmiophyllum sp. events” (Bomfl eur & Kerp 2010, p. 151). Tho- Finally, taxa that probably appeared there mas (1985) considered ferns as a usual element as well are the earlier discussed Coniopteris of conifer dominated open forests, but rarely hymenophylloides, Sphenopteris sp., and Cla- as a strictly dominant factor. He pointed out thropteris meniscoides, as well as some taxa Dicksoniaceae and Cyatheaceae together with which were not used in the analysis: Ginkgoi- horsetails as forming monospecifi c thickets as tes minuta, Geinitzia sp., and Podozamites sp. a stage of succession (see also Retallack 1977). Ginkgoites minuta, similar to all other gink- Apart from ferns, Francis et al. (2008) also gophytes, is placed in the Komlopteris group included liverworts, mosses, cycadophytes and and so is Geinitzia sp., which is comparable angiosperms in the disturbed fl ora, and regar- to Elatocladus sp. Both species of Podozamites ded Taeniopteris as forming thickets in distur- may probably also be included in this group, bed clearings. since they are usually connected with wet con- ditions. 3. Moderately disturbed swamp occupied by Members of this assemblage show differ- the Komlopteris ecogroup. ences in remains fragmentation (Fig. 5). Kom- lopteris nordenskioeldii and Baiera furcata Dominant element. Komlopteris nor- possess the high amount of large fragments denskioeldii. or complete leaves usual for the locality, while This ecogroup occupies a habitat continu- conifers like Elatocladus sp., Pagiophyllum ously passing from moderately wet to swampy, sp., and Podozamites sp. are much more frag- occasionally submerged areas. Here gym- mented: the highest in number are specimens nosperms show their maximum occurrence with small fragments, or debris with a rela- accompanied by Equisetites muensteri, which tively low production of litter. This situation reaches the highest value on Axis 1. The major- would be justifi ed when those conifers also ity of them show leaf structures characteristic grew along the swampy river banks and a por- of wet conditions. To this group belongs Kom- tion of their fragments would be destroyed by lopteris nordenskioeldii with exposed stomata river stream. The other taxa possibly preferred (Barbacka 1994b) and a type of hydatodes, as the swampy surroundings of stagnant water well as Elatocladus sp. connected usually with (lakes, canals, river branches with a slow a wet, even swampy habitat, and all species stream). of ginkgophytes from this locality. Their cuti- cles are either moderately thick (Ginkgoites R e m a r k s. Usually equisetes and ferns were and Baiera) or rather thin (Sphenobaiera). All attributed to swamp (fl oodplain) or marsh forms found in the Mecsek Mts. have papillate (van Konijnenburg-van Cittert 2002, 2008, van cuticles, at least around the stomatal pit (Bar- Konijnenburg-van Cittert & van der Burgh backa 2002). According to Zhou (2009, p. 16), 1989, 1996, Kustatscher et al. 2010, Krassilov the Mesozoic ginkgophytes are often associated 1973, Batten 1974, Abbink et al. 2004). Kras- with plants adapted to “different climates from silov (1973) revealed Equisetites as forming hot and dry to wet and temperate, from coastal monospecifi c thickets in marshes, and he also plain and lowland to inland riparian/swamp ascribed Dictyophyllum, Todites, and Elatides environments”. Sun et al. (2008, p. 220) meas- to this habitat. Elatides was also attributed ured the stomatal index and carbon isotope for to moist lowland by Abbink et al (2004), and the Middle Jurassic Ginkgo leaves, and conclu- Geinitzia and Brachyphyllum were attributed ded that “the fossil Ginkgo in a ‘green house’ to the same by Nguyen et al. (1999). Marat- world is more effi cient in water use than the tiales, Osmundaceae, Dipteridaceae, Dicksoni- living Ginkgo”. aceae and others were interpreted as unders- Other plants, whose niches extend to this torey, either in marshes or along riverbanks habitat, do not reach the maximum mois- (van Konijnenburg-van Cittert 2002, Abbink ture, but rather extend only to slightly wetter et al. 2004, and Hesselbo et al. 2003 without 156

Osmundaceae). In the Mecsek Mts., only recent opinions, Cheirolepidiaceae are found two species of Dictyophyllum, one species of in very variable environments, from dry to Cladophlebis and Phlebopteris sp., which show swampy and coastal (Vakhrameev 1970, 1991, fi ne frond structure, partly occupy the wetter Alvin 1982, Hesselbo et al. 2003, Greb et al. territories, but none of them are abundant in 2006, Wang et al. 2005, Popa & van Konijnen- the areas with maximum moisture. burg-van Cittert 2006); and details of their Chun (1996) discussed a swamp-lake envi- cones confi rm adaptation to wet conditions ronment of the Lower Mesozoic in Korea with (Barbacka et al. 2007). Since the genus Brach- the dominancy of Dictyophyllum and Clathrop- yphyllum is not homogeneous and certain teris species together with Cladophlebis, Ptero- species can be of Cheirolepidiaceae or Arau- phyllum, Anomozamites, and Nilssonia as well cariaceae affi liation (van Konijnenburg-van as ginkgophytes. Out of the Hungarian species Cittert & Morgans 1999), it is only possible of bennettites and cycads, only one, Pterophyl- to indicate that the Brachyphyllum from the lum subaequale expanded to a wetter canopy. Mecsek locality shows a very low tolerancy and Considering that the composition of this can- is strongly restricted to a narrow niche wit- opy complemented by other taxa extended to hin the swamp, reaching maximum wetness this habitat, it corresponds well with the lush, on Axis 1, and probably occasionally in sub- moist vegetation from Sutherland (van Koni- merged areas. jnenburg-van Cittert & van der Burgh 1996). Popa and van Konijnenburg-van Cittert 4. Weakly disturbed, moderately wet canopy (2006, p. 207) interpreted seed ferns from the – Ptilozamites group. genera of Sagenopteris, Pachypteris, Cteno- Dominant species. Ptilozamites cycadea. zamites (Ptilozamites), and Komlopteris as being „swamp margin plants” together with The Ptilozamites group is the most diverse conifers, when cycads together with ferns were compared with representatives of different considered as growing „in the fi nal phase of plant groups, showing here their maximum peat swamp development”. It agrees with ear- frequency: horsetails – Equisetites columnaris; lier Popa’ s suggestions (Popa 1997) that seed ferns – Dictyophyllum nilssonii and Phlebo- ferns of „Thinnfeldia” group were associated pteris sp.; seed ferns – Ptilozamites cycadea; with swamp areas. cycads – Bjuvia simplex and Nilssonia polymor- All species of Hungarian Sphenobaiera were pha; bennettites – Pterophyllum subaequale; found in swampy habitat, although Retallack as well as Desmiophyllum sp.(incertae sedis, (1977) in his reconstruction regarded the genus probably Gnetales). Other taxa whose niches as xerophytic. In the Kimmeridgian fl ora from extend to this habitat are: Nilssonia obtusa, Sutherland, the ginkgophytes, Brachyphyllum, Komlopteris nordenskioeldii, Anomozamites Elatocladus together with Nilssonia, Ptero- marginatus, Sagenopteris sp., and Ginkgoites phyllum, and Sphenopteris are attributed to marginatus. upland forest (van Konijnenburg-van Cittert The composition of this assemblage unites & van der Burgh 1996). Similarly to the Hun- taxa usually connected with both wetter and garian locality, Batten (1974) interpreted coni- drier conditions. Nilssonia polymorpha, dis- fers as growing along river banks. playing a relatively narrow niche limited to this According to Abbink et al. (2004) as well environment (Fig. 10), was generally regarded as van Konijnenburg-van Cittert (2008), river as being connected with wet conditions, simi- SEG was also favourable for ferns, especially larly to Desmiophyllum, which has a delicate from the families Osmundaceae, Dicksoniaceae, cuticle. Contrary to this, Bjuvia simplex with Schizaceae, Dipteridaceae, Cyatheaceae, and its rather thick cuticle and stomata protected Pteridaceae, a fi nding which is not excluded in by a thickened ring shows adaptation to dry the case of the Mecsek fl ora, however, it is not surroundings. It may be a sign of periodical confi rmed either. dryness, but its tall stature and large leaves A little unexpectedly, the genera Pagiophyl- at the top also suggest that these leaves might lum and Brachyphyllum in the Mecsek Mts. have been exposed to sun, therefore, their pro- are limited to highly watered areas contrary to tection is justifi ed even in water-rich areas. their xerophytic micro- and macromorphology The other species with typical xeromorphic (Thévenard et al. 2005). However, according to features is Ptilozamites cycadea, which has 157 coriaceous and rather small pinnules, and sto- Ptilozamites cycadea, which here is the mata protected by a small, very thick cuticu- dominant element, was rather interpreted as lar ring on the upper surface that narrows growing in dry conditions, which is clear from the opening. The guard cells are also thick, its xeromorphic structure. Also, carbon-cycle although large. Nevertheless, xeromorphic interpretations of Yorkshire fl ora placed the features occur not only in plants living in arid genus Ctenozamites (Ptilozamites) in rather circumstances, but also in seasonally or peri- dry conditions (Hesselbo et al. 2003, Kustat- odically dry ones (Rees et al. 2000). scher et al. 2010) even in the coastal zone. Equisetites columnaris from the Hungarian This canopy, similarly to the drier one, groups locality is, compared with E. muensteri, char- cycads and bennettites as displaying here their acterized by smaller, narrower stems, which maximum frequency. Cycads are often con- possibly may be the effect of adaptation to nected with rather wet circumstances (Abbink survival in unfavourable circumstances. Taxa et al. 2004), especially Nilssonia with entire not used in the analysis but attributed to this margins (Vakhrameev 1991). However, this environment are: Neocalamites hoerensis, study demonstrates that, even within this type Cladophlebis roessertii, Ctenis sp., and Pseu- of leaves, the preferences of particular species doctenis sp.. might be different. In Batten (1974), cycads co- Neocalamites carcinoides shows a low co- occurred with ginkgophytes and conifers, they occurrence. On single specimens, it is found were located also together with bennettites in together with Equisetites sp., Bjuvia simplex, open forest, while Kustatscher et al. (2010) Nilssonia obtusa, Sagenopteris sp., Ginkgoites located cycads (including Bjuvia and Nilsso- sp., Ptilozamites cycadea, and on 8 specimens nia) in lowland together with pteridosperms with Komlopteris nordenskioeldii. Cladophle- (including Sagenopteris). All of these composi- bis roessertii occurs together with Nilssonia tions are similar to those in the Mecsek local- obtusa and Phlebopteris sp. on two specimens; ity. However, Thorn (2001) mentioned that in possibly, it can be placed in the same environ- New Zealand (Middle Jurassic) cycads occur ment. Pseudoctenis sp. and Ctenis sp. are very far from the other gymnosperms. In this can- rare and are not associated with any other opy, both species of Mecsek bennettites meet, taxa. Their morphology, which is similar to since their niches overlap at the border of hab- that of Bjuwia simplex, may allow their place- itats. Their preference corresponds with those ment in the same group. described by Gordenko (2008), who interpreted The degree of fragmentation of selected spe- her bennettitalean records as mesophytic. cies from this ecogroup (Nilssonia polymorpha, According to Bomfl eur and Kerp (2010), Phlebopteris sp., Ptilozamites cycadea, (Fig. 5 Clathropteris may occur in a bennettitalean J, K, N) shows a predominance of large frag- dominated assemblage forming herbaceous ments in comparison with small fragments or understorey in open vegetation dominated debris. Litter production is minimal, probably by Bennettitales. In the Hungarian assem- because of the nature of the plants. blages, Clathropteris co-occurred with Nilsso- nia obtusa and Dictyophyllum rugosum, which R e m a r k s. This habitat is the most mixed means that its niche extends rather in the with regard to the structural adaptation of direction of more disturbed territories, while the composing taxa. According to Wnuk and the bennettitalean/cycad assemblage tends Pfefferkorn (1984, p. 348), “coexistence of toward stability. mesomorphic and xeromorphic species in the Thorn (2001) placed arborescent and herba- same community suggests that distributions ceous ferns as well as cycads into a damp area, are being controlled by microenvironmental pointing that cycads grew far from the other factors”, which may mean that small, better gymnosperms (Coniferales and Caytoniales). drained places exist in the canopy occupied by The Neocalamites assigned here forms such types of plant. These authors also indi- mainly monospecifi c thickets, which is uder- cated that xeromorphy is not connected exc- standable because of its size and biology. lusively with dryness, but can also be viewed Probably its growth was similar to Calamites, as a response to light intensity, which in con- whose expansive and thick rhizomes excluded sequence induces succession of plants with all other plants from the habitat (Thomas such tolerancy. 1985). 158

5. Weakly disturbed wetland – Ginkgoites THE COASTAL ZONE group. The possible species appearing in this envi- This type of environment is predominated ronment are Pachypteris banatica and possibly by Ginkgoites marginatus, which is at its max- Brachyphyllum sp., while Pachypteris banatica, imum expanse here. It is in the same mois- occurred alone (three specimens only). Its semi ture range as the highly wet, swampy habitat xeromorphic features (Barbacka 1994c) and of the Komlopteris group, but with a tendency the fact, that another species from this genus toward more stable circumstances. Other spe- (P. papillosa) was regarded as a halophytic cies occurring in these surroundings are: Kom- plant, imply that this species might be placed lopteris nordenskioeldii, Sphenobaiera longi- in the coastal zone. The genus Brachyphyllum, folia, Elatocladus sp., Ptilozamites cycadea, particularly B. crucis of which a few specimens Pterophyllum subaequale, Desmiophyllum sp., are known from the Mecsek locality,was also Phlebopteris sp., and Equisetites columnaris. presumed to be tolerant of salty habitats (Har- The fragmentation of Ginkgoites marginatus ris 1979, Abbink 1998, Abbink et al. 2004, van (Fig. 5M) differs from that of Baiera furcata Konijnenburg-van Cittert & Morgans 1999). (Fig. 5A) in having more fragmented leaves R e m a r k s. Sometimes equisetes are attrib- and the highest production of litter out of all uted to a coastal/tidally infl uenced habitat species from the locality. In this aspect, Sphe- (Batten 1974, Gee et al. 2003). Usually Chei- nobaiera longifolia shows a transition between rolepidiaceae are known as being tolerant to both species. This very high litter production salty soil and interpreted as a type of man- can be explained by relatively stable (the niche grove (Batten 1974, Nguyen et al. 1999). Kus- of Ginkgoites marginatus extends towards even tatscher et al. (2010) interpreted Ptilozamites more stable, Fig. 13), and highly wet circum- sandbergeri as a potential halophytic plant, stances; thus, fallen leaves (ginkgophytes were placing it into the coastal zone together with deciduous) had good conditions for fossilization lycopods. Pachypteris is generally known as in this state. a genus with some halophytic species (Har- R e m a r k s. Compositionally, this environment ris 1964, van Konijnenburg-van Cittert & van shows a transition between moderately dis- der Burgh 1996, van Konijnenburg-van Cittert turbed swamp and less disturbed, moderately 2008, Abbink et al. 2004). wet habitats, where apart from the dominant Ginkgoites marginatus some elements of both canopies are present. The monospecifi c domi- GENERAL REMARKS nancy of Ginkgoites may be explained by anal- ogy with Ginkgo biloba, which shows moderate The Mecsek locality represents typical low- allelopathy. Nam (Nam et al. 1997) revealed land vegetation with a diverse, but not spe- that its leaves produce substances that, after cies-rich, fl ora which fi ts in with the general falling, dissolve in soil and inhibit the develop- rules of wetland ecology presented by Wing ment of seeds of certain other trees. and DiMichele (1995). The proposed habitats The canopy occupied by Ginkgoites mar- and taxa connected with them were dynamic ginatus well corresponds with the statement to a certain degree. Fluctuation in water lev- that during the Mesozoic Ginkgoales preferred els was the factor, which mostly infl uenced “stable and ecologically saturated” places Zhou changes and the succession of vegetation. (2009, p. 15), while Abbink et al. (2004) also Since plant associations within groups were ascribed ginkgophytes to wetland. Francis et based exclusively on taxa co-occurrence which al. (2008) interpreted stable wetland as being was rather stable, these changes were of minor covered by araucarian conifers, ferns and signifi cance. cycads, while Hesselbo et al. (2003) stated that Generally, in the Mecsek delta the circum- in the Yorkshire Jurassic more ginkgophytes stances of medium to less wetness appeared to were associated with wet than dry conditions, be optimal for ferns, for which all species are although they occurred in both types of habitat. found in these categories. The environmental Popa & van Konijnenburg-van Cittert (2006) preferences of ferns are known to be shady, supposed Ginkgoales (including Sphenobaiera warm and humid (van Konijnenburg-van longifolia) as upland representatives. Cittert 2002, Wang 2002), but unexpectedly, 159 here they avoid very wet or swampy surround- Danube delta these levees were covered by oak ings. forests. The ecological conditions of the terri- All cycads prefer less disturbed places, and tory are diverse and strongly depend on fl uc- all ginkgophytes together with conifers are tuation in water levels. grouped in richly irrigated places. Seed ferns The Ptilozamites group would possibly cor- show the widest tolerance. respond with the ecosystem of calcareous cliff Plant distribution in the presented model in the Danube delta. Specifi c morphohydro- reveals differences in environmental prefer- graphic conditions provided an environment ences between species within genera. In most for diverse fl ora adapted to semi-arid condi- cases, different species of the same genus show tions and high insolation. maximum growth in different niches exclud- The most extended, swampy area with the ing species of Sphenobaiera, which are at Komlopteris group is comparable with wetland, their maximum extent in the same niche. The probably encompassing lakeside terrain, or on niches of some species intersect, so they may a riparian formation on the river levees along occur together, like Phlebopteris, Dictyophyl- the channels or in depressions. These are ter- lum, Cladophlebis, and two of the three spe- ritories that, dependent on water level, may cies of Nilssonia, (N. obtusa–N. revoluta, and be periodically submerged. The group could N. polymorpha- N. revoluta), as well as some occasionally reach the slopes of the marine species whose niches are completely separate, levees, too. like those belonging to Equisetites, as well as The Thaumatopteris group could be placed Nilssonia polymorpha with N. obtusa. on territories, which are mostly disturbed, at the riverbanks, on easily damageable alluvial DELTA ENVIRONMENT deposits or newly formed islands, possibly also forming fl oating islands (Dr. M. Dorothei pers. For better understanding of ecosystems, com.), extending also to river levees or swamp. a study of the extant Danube delta can reveal The percentages of particular groups in the the location of particular plant groups in a hypo- plant cover (Fig. 15) and thus putative propor- thetical biome. A complex study of this delta tions among environments within the hypo- has been carried out during the last decade thetic delta suggest that swamps occupied the (Gâştescu et al. 1998, Szilassi 2004, Hanganu largest territories, followed by marine levees et al. 2002). Within the Danube delta, the fol- and drier territories, then cliffs and semi-arid lowing ecosystems were recognised: areas, the smallest territory being occupied by – coastal lagoons highly disturbed zones and fl oating islands. – standing brackish and salt waters Habitat containing Ginkgoites marginatus can – channels and canals with active or free be treated as transition between swamp and water circulation semi-arid areas. – standing fresh water: lakes with active or Deltas have a mosaic character and envi- reduced change of water table ronments can change even within small areas, – wetlands: fl ooded reed beds, fl oating reed and also within short time intervals (seasonal beds, riparian formations, frequently fl ooded changes). Such changes were also indicated by river levees – forests (dunes, fl oodplains) – shrubs and herbaceous vegetation (cal- careous cliffs, predeltaic fi elds, low marine levees) – open places with little or no vegetation (dunes, beaches) If an attempt is made to situate the Jurassic ecogroups on a delta model using existing eco- systems similar to those of Danube delta, the habitat of the Sagenopteris group is most simi- lar to those of the marine levees – these are slightly raised territories perpendicular to the Fig. 15. The percentages of particular groups in the plant main branches of the river. Recently, in the cover 160 pollen analysis (see Götz et al. 2011). Altera- paradox can be explained by local hydrological tion from fern-dominated to conifer-dominated conditions, i.e. permanently high humidity and assemblages was also observed by Batten a high water table only in restricted depres- (1974) and Wang et al. (2005). Fluctuations in sions with dense vegetation and peat accumu- water level (at the rivers, channels and lakes) lation, while drier, seasonal climate dominated and changes in salinity (infl uence of sea move- in the surrounding land. Therefore, it can be ments) may cause plant taxa to alternate in suggested that the occurrence of coal depos- short cycles (too short to be visible in the fos- its in this setting is not connected with an sil state). In this case, the general content of overall humid climate, but either with humid eco-groups would be plus minus constant, but periods or differentiated topography character- within them there would certainly not be any ized by locally wetter depressions (Pieńkowski, cases in which all taxa occurred together in the pers. com.). Mineralogical studies generally same time. confi rm associations between sandstones and channels (the presence of Ostracoda), between siltstones and periodically dried fl oodplain, RELATIONS BETWEEN TAXA, TYPE and between laminated siltstones and swamps OF ROCK OR SITE IN THE LOCALITY (Bodor pers. com.). Associations between rocks and sites group All leaf taxa and distinctive rock types con- two open mines Vasas and Pécsebánya and taining plant fragments were used in the anal- two shafts – Zobák and Béta and separate ysis of the associations between taxa and rock them from the dumps (Fig. 16). More inter- types (in previous analyses not all taxa were esting are associations between taxa and sub- considered). It seems that the obtained relation- strate. It is noticeable from Table 7 that plant ships do not follow the order of the ecogroups, groups do not really follow a certain rock type. which means that connections between type There are two taxa with completely opposite of rock and taxa are insignifi cant. Although, trends: Brachyphyllum sp. which has a lim- macroscopically, ten types of rock were distin- ited niche in one group appeared in all types guished, microscopic examination proved that of rocks, while Cladophlebis haiburnensis a lot of them have similar structures which occurs only in one type of rock, but belongs enable them to unite within the same facies, to three assemblages. It may be interpreted thus rock types were simplifi ed to 4 categories that for Brachyphyllum sp., habitat wetness (Table 7). Similarly to the previous analysis, and given ecogroup were the main factors that taxa do not seem to be in close connection with infl uenced its distribution, and not the type of a given type of facies. environment: fl oodplain, swamp, or channel. Three lithofacies can be distinguished in the Contrary to this, Cladophlebis haiburnensis rock unit in which most plant remains were seems to favour certain types of environment found. Sandstones could represent environ- (here lakes or swamps), but might belong to ments containing higher energy depositional more ecogroups tolerating this type of habitat. conditions, like channels; siltstones could rep- Data from Tables 7 and 8 suggest that local resent environments of low, episodically higher conditions varied within small areas and were depositional energy, like alluvial plain or delta mixed to a high level, while most of the plants plain; and fi ne lamellated siltstones could origi- show a wide tolerance. nate in more distal environments, like lakes and swamps. The presence of calcite cementation perpendicular to lamination provides evidence for carbonated pedogenic levels whose presence might indicate a semi-dry or dry climate, at least in the lower part of the sequence. The occurrence of calcite molusc shells indicates a lack of acidifi cation (at least tem- poral) of the water in the basin, which should be expected in a humid climate. On the other hand, the presence of coal seams and rich fl o- Fig. 16. Relationships between rocks and sites, UPGMA ral remains points to a humid climate. This analysis 161

Table 7. Association between substrate and taxa in relation to ecogroups and facies. 1 – Sagenopteris group, 2 – Thaumatop- teris group, 3 – Ptilozamites group, 4 – Ginkgoides group, 5 – Komlopteris group

Fine lam. Grey silt Fol silt Sandst. Species silt. Groups river or delta fl oodplain lakes or swamps channels Anomozamites marginatus XX 1, 3 Baiera furcata XXX 2, 5 Bjuvia simplex XXX 1, 3 Brachyphyllum sp. X X X X 5 Cladophlebis denticulata XXX 1, 2 Cladophlebis haiburnensis X1, 2, 5 Clathropteris meniscoides XX X2, 5 Dictyophyllum nilssonii X X 2, 3, 5 Dictyophyllum rugosum XX 1, 2, 5 Elatocladus sp. X X X 2, 4, 5 Equisetites columnaris X3, 4 Equisetites muensteri XX X5 Ginkgoides minuta X5 Ginkgoites marginatus XXX 3, 4 Komlopteris nordenskioeldii X X X 2, 3, 4, 5 Marattiopsis hoerensis XXX 1, 2 Neocalamites carcinoides XX 3 Nilssonia obtusa XXX 1, 3 Nilssonia polymorpha XXX 3 Nilssonia revoluta XXX 1 Pagiophyllum sp. X X X X 5 Phlebopteris angustiloba XX X2 Phlebopteris sp. X X X 2, 3, 4, 5 Podozamites sp. X X 5 Pterophyllum subaequale XXX 3, 4, 5 Ptilozamites cycadea X X X 1, 3, 4, 5 Sagenopteris sp. X X X 1, 2, 3, 5 Sphenobaiera leptophylla XX 5 Sphenobaiera longifolia XX 4, 5 Thaumatopteris brauniana XX X2 Todites goeppertianus XXXX1 Todites princeps XX 1

Figure 17 illustrates the association generally correspond to the geographical dis- between taxa and sites. On the basis of this tinction between the region of Komló and analysis, the most similar sites appeared to be region of Pécs. Turning back to Figure 4, it Vasas (an open mine) and Budafa (a dump), which proves that unproductive rocks from Vasas were mostly transported to Budafa (so in further analyses they are united into one site). A similar situation pertains with the Zobák shaft, Kossuth shaft and Dávidföld (a dump – but this dump is connected with more shafts, some of which are unknown, so is omitted from further discussion). Based on the cladogram, similarities divide the sites into two groups: Pécsbánya–Béta shaft–Vasas+Budafa Fig. 17. Relationships between taxa and sites, UPGMA anal- and Kossuth shaft–Zobák shaft. These groups ysis 162 can be observed that passing from Pécsbánya facies changes. Pécsbánya (Fig. 18C) and Béta towards Komló, the delta facies comes to an (Fig. 18D) are considerably poorer and there, end at the Béta shaft, so in this region fl ood- the infl uence of the deltaic facies is higher. plain dominated, with less river facies than in the region of Pécs. The macrofl oral composition of both zones noticeably follows this distinc- COMPARISON OF MECSEK PLANT tion in proportion to particular components: ASSEMBLAGE OBTAINED USING TWO the Komló region is generally richer in plants, METHODS: THE SEGS MODEL and its main disproportions regard increased AND DCA CO-OCCURRENCE MODEL amounts of Komlopteris nordenskioeldii, Ptilo- zamites cycadea, Bjuvia simplex, ginkgophytes, The reconstruction of the Mecsek palaeoen- and Elatocladus sp., thus dominant elements vironment started with palynological investi- of the Komlopteris and Ptilozamites ecogroups, gations, fi rst of all made by Bóna (1963, 1969). but slightly fewer ferns, Sagenopteris, and the He distinguished three zones according to pal- genus Nilssonia (Sagenopteris and Thaumat- ynomorph occurrence: a deep water zone which opteris ecogroups). This implies that the region was characterised by allochthonous pollen and of Komló could have been wetter and less dis- spores transported by wind from land; shallow turbed, while the region of Pécs was could have swamp and swamp forest with allochthonous been more disturbed and drier (Fig. 18 F, G). gymnosperm pollen and autochthonous horse- The distribution of the main fl oristic com- tail spores; and inshore areas with mainly ponents in particular sites is shown in Figure conifer pollen (Lachkar et al. 1984). 18A–F. Two neighbouring sites, nearest to New investigations based on microfl oral the opposite sides of the border between the data were provided by Ruckwied et al. (2008). regions represent quite different compositions. The authors dealt mainly with the Triassic– In the Béta shaft, ferns predominate (Fig. Jurassic boundary and from this point of view 18D), while in Kossuth, it is the Komlopteris distinguished environmental types character- ecogroup (Fig. 18A), which is dominant. The istic of the Upper Triassic as upland and river Zobák shaft lies between the Kossuth shaft and which turn into warmer lowland and drier low- Szászvár in a NW direction. Here again, river land at the lowermost Hettangian. and delta facies appear, but evidently fl ood- The authors also pointed out diffi culties in plain dominates. As far as plant composition is reconstruction based on microremains alone, concerned, this resulted in a relatively low per- which often appear in problems of association centage of all elements with a predominance between palynomorphs and mother plants, as of ferns, Ptilozamites cycadea and species from well as in the unknown tolerance of plants to the genus Nilssonia. Also Ginkgoites margina- stress or disturbance. The next attempt to recon- tus occurs here in higher numbers (ecogroup struct the palaeoecology in the Mecsek locality G. marginatus, low disturbance and moderate was based on micro and macroremains (Götz wetness). The proportion of other taxa ranges et al. 2011). The authors discussed changes in between 2–7% (Fig. 18B). the palynomorph assemblages refl ecting a long Pécsbánya is most exposed to the South, term trend from river dominated to swamp where the delta facies dominates alternately dominated elements, as well as small-scale with the fl oodplain. Here the Cheirolepidiaceae cyclic changes of deltaic–swamp–river domi- and Desmiophyllum are absent and ginkgo- nated elements. This way, temporal changes phytes are rare. Komlopteris is at less than were detected, in which habitat types proposed 5 %, but the ferns, Nilssonia and Elatocladus, by Abbink (1998), Abbink et al. (2001, 2004) dominate. It seems that the Thaumatopteris were represented by different frequency of ecogroup is dominant (disturbed and moder- palynomorphs through the Rhaetian and Het- ately wet habitat), and elements occur from wet tangian (Götz et al. 2011, fi g. 8). and drier habitats that tolerate disturbance According to Abbink’s environmental types (Fig. 18C). Cheirolepidiaceae (Pagiophyllum) (Abbink 1998, Abbink et al. 2001, 2004) and the are most abundant at Vasas and the Zobák corresponding SEGs (Sporomorph Ecogroups) shaft, where alluvial facies are most developed he proposed, a model of the Mecsek vegetation (Fig. 18E,B respectively). It can be observed was discussed. According to fi gure 8 from Götz that richness in fl oral elements depends on et al. (2011), the participation of sporomorphs 163

Fig 18. A-F. Percentage of taxa in particular sites 164 characteristic of upland is very low during the & Beall 1990, DiMichele et al. 2008), sedi- Hettangian. Drier lowland shows medium and mentology and macrofl ora (Falcon-Lang et al. fl uctuated contribution values, the warmer 2009), coal-ball investigations (Galtier 1997), lowland and wetter lowland show maximum stomatal index calculation (for references see values, and the elements of river deposits page 134, about SI, this paper) and carbon show the most variable values. The microfl oral isotope and stomatal CO2 densities (Hesselbo assemblages and their environmental connec- et al. 2003). tions were discussed in relation to sporomorph Investigations consider biotic changes over – mother plant combinations,with a macrofl ora long periods (DiMichele et al. 2008, Greb et al. adaptability approach. 2006), or focus on a given period or locality. Table 8 contains a comparison of both Many reconstructions have been made for the models. The DCA co-occurrence model inclu- Palaeozoic, mainly Carboniferous or Permian des all taxa according to their niches in their wetlands (Di Michele & Phillips 1994, DiM- widest meaning, complemented by taxa which ichele et al. 2006, DiMichele & Gastaldo 2008, were not considered in the analysis. The names Libertin et al. 2009, Bercovici et al. 2009), or of habitats used by Götz et al. (2011), after coal swamps (Phillips & Peppers 1984, Phillips Abbink (1998), Abbink et al. (2001, 2004), with et al. 1985, Thomas & Cleal 1993), providing small modifi cations, are changed according to good records for such investigations. local conditions which better correspond with Well-documented and detailed reviews of the suggested reconstruction and in the table wetlands and their role as ecosystem over are supplemented by two additional habitats. a time period were given by Greb et al. (2006) The environmental types do not correspond and of their taphonomy by Gastaldo et al. with each other in each category, but were col- (1995). lated based on their defi nitions. Both models in Special methods were applied for sporo- general outline correspond with each other. The morph analyses based on familiar ordinary most similar is the composition of the swamp plant adaptations and comparisons with (“wetter lowland”), although the present model extant relatives (Abbink 1998, Abbink et al. does not contain the cycads and Pachypteris (2001, 2004). (which in this case were placed in the coas- The Jurassic (or closely related period from tal habitat) suggested by the SEG model. The Late Triassic to pre-angiosperm Cretaceous) preferences of cycads and ginkgophytes are ecosystems have also been worked up using different. In seasonally dried fl oodplain (“drier different methods (Batten 1974, van Koni- lowland”) in the present model the Cheirolepi- jnenburg-van Cittert & van der Burgh 1996, diaceae are completely absent and the partici- Abbink 1998, Nguyen et al. 1999, Thorn 2001, pation of Dicksoniaceae is doubtful. The most Shao et al. 2003, Hesselbo et al. 2003, Abbink different is the “river”, which in both models et al. 2004, McElwain et al. 2007, Sun et al. does not contain any common taxa. The “high- 2008, Jansson et al. 2008, Francis et al. 2008, land” (drier inland or levees) is also different Gordenko 2008, Kustatscher et al. 2010). in having only two common elements. The general rule in those studies was to mainly use a generic or higher systematic level for plant taxonomy and in this way certain GENERAL DISCUSSION interpretations were generalized. Taxonomy of fossils is often problematic and analyses on In the last decades, palaeoecological recon- a specifi c level may be regarded as running structions have become very popular. They a risk of producing false results, particularly encompass very different types of investiga- in the mathematic models (Rees et al. 2000, tion, based on plants and animal assemblages 2002). Palynological studies cannot also be used in a locality (Lerner et al. 2009), evidence of for specifi c analyses, because sporomorphs are fossil micro- and megafl ora (Thorn 2001, Ber- not specialized to such a degree. covici et al. 2009), analyses of palaeosols (Shel- It ensues that fl oristical compositions of dif- don & Tabor 2009), analyses of peats (DiMi- ferent landscapes are more or less universal. chele & Phillips 1994), sedimentology (Shao et The present analysis, which in most cases al. 2003), complex sedimentology, palaeozoo- was based on species, showed that this method logical and palaeobotanical data (DiMichele can provide interesting results, showing that 165

Table 8. Comparison of assemblages obtained by using two methods of paleoecological reconstruction (SEGs and DCA co- occurrence model). Taxa in systematic order. Names written in bold – taxa showing maximum frequency in the present model. Underlined – taxa not attributed to given habitat by SEGs. Interrupted underline – taxa which niches extend to the given habitat, or not used in analysis but attributed as possible elements

Analysis of sporomorph/macrofl ora Abbink’s model Present model – taxa coocurrence Present paper (Götz et al. 2011) wetter lowland Sphenophyta Komlopteris nordenskioeldii swamp (swampy Osmundaceae (Todites, Cladophlebis) Baiera furcata fl oodplain) Pachypteris (Corystospermales) Sphenobaiera longifolia (Komlopteris, Sphenobaiera leptophylla Ptilozamites) Equisetites muensteri Cycadales Elatocladus sp. Ginkgoales Pagiophyllum sp. Dicksoniaceae (Sphenopteris, Brachyphyllum sp. Coniopteris) Ginkgoites marginatus Bennettitales Pterophyllum subaequale Dipteridaceae (Clathropteris, Ptilozamites cycadea Thaumatopteris, Dictyophyllum) Dictyophyllum nilssonii Coniferales (Elatocladus, Geinitzia) Dictyophyllum rugosum Cladophlebis haiburnensis, Phlebopteris sp. Sagenopteris sp. Desmiophyllum sp. Coniopteris hymenophylloides Sphenopteris sp. Ginkgoites minuta Genitzia sp. Clathropteris meniscoides Podozamites sp. river pteridosperms (Komlopteris, Elatocladus sp. swamps along the Ptilozamites) Pagiophyllum sp. river banks Sphenopteris Podozamites sp. Dipteridaceae (Clathropteris, Thaumatopteris) Caytoniales (Sagenopteris) warmer lowland Osmundaceae (Todites, Cladophlebis) Thaumatopteris brauniana disturbed areas Pachypteris (Corystospermales) Equisetites sp. (small islands, Komlopteris, Dictyophyllum rugosum fl oating islands, Ptilozamites) Cladophlebis haiburnensis river banks) Sphenophyta Phlebopteris angustiloba Coniferales (Elatocladus) Baiera furcata Dicksoniaceae (Sphenopteris, Elatocladus Coniopteris) Cladophlebis denticulata Cycadales Dictyophyllum nilssonii Ginkgoales Komlopteris nordenskioeldii Marattiopsis Dipteridaceae (Clathropteris, hoerensis Thaumatopteris, Dictyophyllum) Sagenopteris sp. Bennettitales Phlebopteris sp. Clathropteris meniscoides Coniopteris hymenophylloides? Sphenopteris sp.? drier lowland Cycadales Nilssonia polymorpha weakly disturbed Cheirolepidiaceae Pterophyllum subaequale moderately wet bennettitales Ptilozamites cycadea (cliffs, seasonally pteridosperms (Komlopteris, Bjuvia simplex semi-arid habitats) Ptilozamites) Dictyophyllum nilssonii Dicksoniaceae (Sphenopteris, Equisetites columnaris Coniopteris) Desmiophyllum sp. Phlebopteris sp. Nilssonia obtusa Komlopteris nordenskioeldii Anomozamites marginatus Sagenopteris sp. Ginkgoites marginatus Neocalamites hoerensis Cladophlebis roessertii Ctenis sp. Pseudoctenis sp. 166

Table 8. Continued

Analysis of sporomorph/macrofl ora Abbink’s model Present model – taxa coocurrence Present paper (Götz et al. 2011) highland Caytoniales(Sagenopteris) Sagenopteris sp. drier areas (marine Ginkgoales Nilssonia revoluta levees) Cheirolepidiaceae Anomozamites marginatus Coniferales (Geinitzia) Cladophlebis denticulata pteridosperms (Komlopteris, Marattiopsis hoerensis Ptilozamites) Nilssonia obtusa Ptilozamites cycadea Bjuvia simplex Cladohlebis haiburnensis Dictyophyllum rugosum Todites goeppertian Todites princeps Clathropteris meniscoides Ginkgoites marginatus weakly disturbed Komlopteris nordenskioeldii, Sphenobaiera wetland longifolia, Elatocladus sp., Ptilozamites cycadea, Pterophyllum subaequale, Desmi- ophyllum sp., Phlebopteris sp., Equisetites columnaris Pachypteris banatica? coastal Brachyphyllum sp.? different species of the same genus often tend al. 2002). On the other hand, the preferences to choose different life conditions. of genera are easier to interprete and general- Numerous environmental reconstructions ize. were based on the sporomorph ecotype in The standard reconstructions of plant Abbink’s model, or in the case of macrofl ora, assemblages rarely consider the wide toler- on structural adaptations of plants and com- ance of plants, instead placing certain taxa parison with their extant relatives. Since in quite narrow environmental categories. most remains were preserved in wetland The multivariate analyses, though used in circumstances (Greb et al. 2006), and these palaeobotany for more than 30 years (Spicer remains provided most information thanks to & Hill 1979), have so far been employed for their state of preservation (grossmorphology more general databases. Jasper et al. (2010) and cuticle), the majority of reconstructions applied a DCA method which appeared to be regard this ecosystem and consequently, types suitable for palaeoecological reconstructions. of environments in different localities are The authors differentiated plant associations comparable. The classic model of wetland, or (miospores) depending on the environment of delta plain, contains territories as proposed by the Duckmanian wetland. Abbink (1998): upland, lowland, river, pioneer, The CA multivariate analysis of the Per- coastal and tidally-infl uenced areas, which are mian fl oras/climates over the world was usually slightly modifi ed, depending on local ap plied by Rees et al. (2000, 2002) as a method surroundings. of providing the best available proxy for the Abbink (1998, Abbink et al. 2001, 2004) original vegetation. The authors documented in creating his model, studied macrofl oral global geographic patterns of Permian climate assemblages and their possible attribution to parameters using fossil fl oras and climate- a given type of canopy. Since then, Abbink’s sensitive sediments based on co-occurrences model has become popular not only in palyno- and distribution patterns of fossil leaves at logy, but also in macrofl ora research. a generic level. This way, an arrangement of The common feature for these reconstruc- fossil genera from 644 localities in the Nort- tions was the study of a macrofl ora at a generic hern Hemisphere was obtained. The authors level, due to lack of suffi cient information on used a mathematical method for grouping the ecological requirements of different species plants of the same parameters and tracing within one genus, as well as to taxonomical their association with sedimentological and uncertainties. This was especially important palaeoclimatological patterns over the world. in the use of multivariate analyses (Rees et The present investigation tried to group plants 167 in a similar way, based on their co-occurrence, natural powerful events like storms, hurrica- but in a much narrower context, within one nes, and so on. Contrary to this, it is worth locality area. noticing that detritus or litter in most cases is When making a mathematic model for pal- monospecifi c, which suggests that its deposi- aeoecological reconstructions, some factors tion in each case is rather restricted to small that infl uence the state of a fl ora, but cannot basins. Generally, large swampy territories be considered in the analysis, should be taken and numerous lakes provided good conditions into account. for the fossilization of plants growing together T i m e. The palaeoenvironment developed or close to each other. and changed according to the cycles menti- The analyses employed in the presented oned above over a long time. Development of investigations provided very interesting results. a single stratum in the Mecsek Coal Forma- They were mainly used to demonstrate how it tion was estimated as taking ca 7000 years. is possible to apply statistic methods to the In the region of Pécs, the development time interpretation of relationships between fossil for the entire coal measure was estimated taxa and environmental changes from a given as 4.5 million years (Nagy 1969). In spite of locality. The other question was how the results such a long period and accidental gathering, could help in reconstructing the palaeoecology the fl ora seems to be quite compact (species and what type of information can be detected assemblages may persist for a million years – for understanding mechanisms functioning in DiMichele 1994), which has also been proved plant communities. Based on plant co-occur- by palynological records. Its low variability rence, we obtained signs of regularity in the suggests that during this time its elements grouping of taxa into assemblages, which may have not changed to a high degree. It can be be interpreted as a response to local conditions. assumed that coexistence of given species was Group of plants segregated by this method are nearly constant since it was caused by environ- clearly interpretable and fi t in with our knowl- mental conditions and not by temporal factors. edge about the ecology of the delta. However, time as a factor cannot be entirely overlooked. CONCLUSIONS The casual character of records. In the open mines, the remains were collected at 1. A DCA method appeared appropriate different points of the entire sequence, but not for the reconstruction of a palaeoenvironment continuously from consecutive strata. In the based on the co-occurrence of taxa. Mecsek Coal Formation, the beds with plant 2. Such investigations may be used not only remains were distributed sparsely as irregular at a generic (or higher) level, but also at a spe- lenses and not as continuous sequences that cifi c level, which can help to detect the varied were caused partly by a process of biostrati- tolerance of different species belonging to the nomy, and partly by tectonic activity. A portion same genus. of the material was collected from the dump 3. Using a DCA method, it was possible to heaps. Although the collecting was accidental separate fi ve plant associations whose composi- in character, co-occurrence of given taxa shows tion depended on environmental circumstances the same tendencies and does not vary much. like soil moisture and stability/disturbance of Circumstances of fossilization. Most their living areas. The same type of analysis taxa are of autochthonous or parautochtho- made for particular species shows their wider nous character, according to Gastaldo’s defi ni- extension within the whole community, which tion (Gastaldo et al. 1995). This is revealed by made it possible to defi ne the range of their the good state of the remains, unbroken large ecological tolerance as well as determine all fragments, often almost complete compound possible combinations of taxa within a given leaves, branching shoots and delicate pinnules type of environment. etc. However, there are also slabs with broken 4. Association between rock matrix and and crowded fragments, which is unavoida- taxa, as well as between taxa and site within ble during a long time period, and may have the locality proved that in the Mecsek local- been caused by the character of the delta and ity, the biodiversity of the areas displaying 168 a dominance of delta facies was poorer than BARBACKA M. 1994b. Komlopteris Barbacka nov. those in which alluvial facies predominated. gen., a segregate from Pachypteris Brongniart. 5. Comparison with the environmental recon- Rev. Palaeobot. Palynol., 83: 339–349. struction of the same locality obtained on the BARBACKA M. 1994c. Reconstructions of stomata of basis of sporomorphs and macrofl ora remains Komlopteris and Pachypteris and their adaptation to climate. Annls. Hst.-Nat. Mus. Natural. Hung., shows that most similar results regarded the 86: 5–11. plant compositions of swamps and warmer BARBACKA M. 2001. The cycads of Hungarian Lias- lowland (here interpreted as moderately wet, sic. Rev. Paléobiol., 20(2): 525–541. highly disturbed areas). The contents of other BARBACKA M. 2002. The Liassic Ginkgoales from environmental categories signifi cantly diverge the Mecsek Mountains, Hungary. Rev. Paléobiol., in both methods. 21(2): 697–715. BARBACKA M. 2009. Sphenophyta from the Early ACKNOWLEDGEMENTS Jurassic of the Mecsek Mts., Hungary. Acta Palaeo- bot., 49(2): 221–231. I would like to express my thanks to Ass. Prof. BARBACKA M. & BODOR E. 2008. Systematic and Józef Mitka for his assistance and help in statistic palaeoenvironmental investigations of fossil ferns analyses of my material. I am very grateful Bodor Cladophlebis and Todites from the Liassic of Hun- Emese, who helped to collect the database and made gary. Acta Palaeobot., 48(2): 133–149. thin sections for mineralogical analysis of rocks and Dr. György Szakmány for help in this analysis. I also BARBACKA M. & van KONIJNENBURG-van CITERT thank very much Prof. Johanna van Konijnenburg- J.H.A. 1998. Sun and shade leaves in two Jurassic van Cittert, Prof. Mihai Popa and Prof. Leon Stuch- species of Pteridosperms. Rev. Palaeobot. Palynol., lik, for critically reading the manuscript and their 103: 209–221. suggestions, Prof. Géza Császár and Prof. Krzysztof BARBACKA M., ZIAJA J. & WCISŁO-LURANIEC E. Birkenmajer for remarks on the geological part, and 2010. Taxonomy and palaeoecology of the Early prof. Grzegorz Pieńkowski for his help in facies inter- Jurassic macrofl ora from Odrowąż, central Poland. pretation. I am very grateful prof. Botond Kiss and Acta Geol. Pol., 60(3): 373–392. Dr. Mihai Dorotfei for materials about Danube delta and suggestions concerning plant ecology of delta, as BARBACKA M., ZIAJA J., WCISŁO-LURANIEC E. well as M.S. Agnieszka Sojka for help in drawing and & REYMANÓWNA M. 2007. 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PLATES

Plate 1

1. Equisetites columnaris, No BP 96.144.1. 2. Todies goeppertianus, No BP 89.224.1. 3. Phlebopteris angustiloba, leaf base, No BP 89.440.1. 4. Clathropteris meniscoides, fragment of leaf base, No BP 89.315.1. 5. Dictyophyllum nilssonii, near the leaf base, No BP 96.80.2. 6. Dictyophyllum rugosum, No BP 94.273.1. Plate 1 175

M. Barbacka Acta Palaeobot. 51(2) 176

Plate 2

1. Todites princeps, No BP 94.692.1. 2. Phlebopteris sp. No BP 96.239.1. 3. Cladophlebis denticulata, No BP 2004.1040.1. 4. Cladophlebis roessetii, No BP 2004.1002.1. 5. Komlopteris nordenskioeldii, shade leaf, No BP 89.368.1. 6. Ptilozamites cycadea, No BP 94..1. 7. Komlopteris nordenskioeldii, sun leaf, No BP 89.275.1. Plate 2 177

M. Barbacka Acta Palaeobot. 51(2) 178

Plate 3

1. Sagenopteris nilssoniana, No BP 89.385.1. 2. Nilssonia polymorpha, No BP 94.879.1. 3. Bjuvia simplex, No BP 98.1054.1. 4. Ginkgoites marginatus, No BP 2011.230.1. 5. Sphenobaiera longifolia, BP 94.989.1. 6. Baiera furcata, No BP 98.596.1. 7. Podozamites schenkii, No BP 89.35.1. 8. Pagiophyllum sp. No BP 98.1216.1. 9. Elatocladus sp. No BP 2000.1202.1. Plate 3 179

M. Barbacka Acta Palaeobot. 51(2)