Relationships Between Ecosystems and Plant Assemblages As Responses to Environmental Conditions in the Lower Jurassic of Hungary and Romania

Acta Palaeobotanica 55(1): 3–17, 2015 DOI: 10.1515/acpa-2015-0005

Relationships between ecosystems and plant assemblages as responses to environmental conditions in the Lower Jurassic of Hungary and Romania

MARIA BARBACKA1,2, MIHAI E. POPA3, JÓZEF MITKA4, EMESE BODOR 5,6 and GRZEGORZ PACYNA7

1 Hungarian Natural History Museum, Botanical Department, H-1476 Budapest, P.O. Box 222, Hungary; e-mail: [email protected] 2 W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Kraków, Poland 3 University of Bucharest, Faculty of Geology and Geophysics, Laboratory of Palaeontology, N. Bălcescu Ave. 1, 010041 Bucharest, Romania; e-mail: [email protected] 4 Jagiellonian University, Botanical Garden, Kopernika 27, 31-501 Kraków, Poland; e-mail: [email protected] 5 Hungarian Geological and Geophysical Institute, Geological and Geophysical Collections, 1143 Budapest, Stefánia út 14, Hungary 6 Eötvös Loránd University, Department of Palaeontology, 1117 Budapest, Pázmány Péter sétány 1/C, Hungary; e-mail: [email protected] 7 Department of Palaeobotany and Palaeoherbarium, Institute of Botany, Jagiellonian University, Lubicz 46, 31-512 Kraków, Poland; e-mail: [email protected]

Received 24 April 2015; accepted for publication 26 May 2015

ABSTRACT. Two Early Jurassic localities, the Mecsek Mts in Hungary and Anina in Romania, are similarly significant and both floras are of autochthonous/paraautochthonous origin. In the Early Jurassic the Hungar- ian locality was a delta plain; the Romanian locality was an intramontane depression filled predominantly by a braided river system. The floristic composition of the two localities (52 genera, 120 species), although superfi- cially similar (25 common genera), differs at species level (only 9 common species) as well as in the proportions of taxa in major plant groups. These differences can be explained by differences in environmental conditions resulting from palaeogeographic and topographic factors. Based on previous and recent studies, alpha diversity as well as statistically (DCA, PCA) differentiated ecogroups are compared and discussed. For common species, the GLM method was used to classify them to particular environmental response types. Their environmental requirements in both ecosystems are evaluated. Some of the shared species showed different preferences at the localities, explainable by their broad ecological tolerance.

KEYWORDS: macroflora, palaeoecology, Early Jurassic, Hungary, Romania

INTRODUCTION

A comparison of Hungarian and Roma- preservation (Popa 1998, Barbacka 2011). Both nian Early Jurassic localities is of interest of them have been systematically sampled for to us, since these localities are close to each more than 20 years. There are exhaustive lists other, the straight-line distance being ca of taxa recognised so far, and the geological 370 km. Both floras are of autochthonous/par- settings are well known (Barbacka 1991, 1992, aautochthonous origin, associated with coal- 1994a, b, 1997, 2000, 2001, 2002, 2009, Bar- bearing deposits, and with similar modes of backa & Bodor 2008, Bodor & Barbacka 2008, 4

Givulescu & Popa 1994, 1998, Kędzior & Popa turbed swamp (Komlopteris nordenskioeldii, 2013, Nagy & Nagy 1969, Popa 1992, 1997a, Elatocladus sp., Baiera furcata, Sphenobaiera b, 1998, 2000a, b, 2001a, b, 2005, 2014, Popa longifolia, Pagiophyllum sp., Brachyphyllum & Kędzior 2008, Popa & Van Konijnenburg- sp., Sphenobaiera leptophylla, Equisetites van Cittert 1999, 2006, Popa & Meller 2009, muensteri). Thévenard & Barbacka 2000). Statistical stud- Anina (formerly known as Steierdorf) is ies of both localities have distinguished plant a historical coal mining centre in the middle ecogroups (Barbacka 2011, Barbacka et al., area of the Reşiţa Basin, where the plant-bear- MS.). In the present work we applied statisti- ing Steierdorf Formation reaches 250 m thick- cal methods to both localities in order to com- ness (Bucur 1991, 1997, Popa & Kędzior 2008, pare their ecological backgrounds and plant Popa 2009). The Steierdorf Formation is coal- environmental responses. Although they are bearing, yielding eight bituminous coal seams, palaeogeographically close, they differ in their Hettangian-Sinemurian in age. The Steierdorf taxonomical composition and type of geological Formation is formed mainly by a braided river setting. system occurring in a depression during the The Hungarian locality in the Mecsek Mts Hettangian-Sinemurian, where mires, lakes, is a typical delta plain (facies: delta – limnic – flood plains next to levees, and river channels lacustrine – delta – lagoonal – plain marine – occurred (Popa 2009, Kędzior & Popa 2013). lagoonal – plain marine), which is Hettangian For plant ecology, PCA was performed (based and earliest Sinemurian in age. The recon- alike in Mecsek on co-occurrence of taxa in the structed palaeoenvironment was characterised same hand specimen), revealing four ecogroups as a system of river channels with river levees whose taxonomical composition depended and marine barriers, crevasse splays, swampy mainly on the moisture/disturbance gradient areas, lakes, and channels (Nagy & Nagy 1969, and the temperature gradient (Barbacka et al. Paál-Solt 1969, Barbacka 2011). Detrended in prep). correspondence analysis (DCA) of floral com- 1. Podozamites group, in moderately position (based on co-occurrence of taxa on the wet and disturbed habitat, not influenced by same slabs) gave five ecogroups interpreted as temperature (Podozamites paucinervis, Sphe depending on the two most important factors – nobaiera sp. and Pinites sp.). moisture and degree of disturbance (Barbacka 2. Schizoneura group. Lower tempera- 2011): ture and higher moisture/higher disturbance, 1. Sagenopteris group. Moderately dis- a typical flood plain association (Neocalamites turbed, relatively dry (non-flooded) inland (Schizoneura) carcinoides, Dictyophyllum nils areas (Nilssonia revoluta, Anomozamites mar sonii, Cladophlebis nebbensis, C. haiburnen ginatus, Cladophlebis denticulata, Maratti sis, Dictyophyllum nervulosum, Coniopteris opsis hoerensis, Sagenopteris sp., Nilssonia murrayana, Matonia braunii, Thaumatopteris obtusa). brauniana). 2. Thaumatopteris group. Highly dis- 3. Zamites group. Higher temperature turbed, short-lived, moderately wet areas and higher moisture, probably swampy in its formed by alluvial deposits (islands, peninsu- last, closing moments, when it was filled up las, forelands), fully damaged by river flooding, with sediment (Zamites schmiedelii, Baiera occupied by pioneer plants (Thaumatopteris sp., Ptilophyllum sp., Cladophlebis denticu brauniana, Phlebopteris angustiloba, Equiseti lata, Geinitzia sp., Ginkgoites sp., Komlopteris tes sp., Dictyophyllum rugosum, Cladophlebis nordenskioeldii, Ptilozamites cycadea). haiburnensis) 4. Nilssonia group. Relatively dry, moder- 3. Ptilozamites group. Weakly disturbed, ately warm and undisturbed conditions corre- moderately wet canopy (Nilssonia polymorpha, sponding with levees, which were the highest- Equisetites columnaris, Pterophyllum subae elevation relief forms in the basin (Nilssonia quale, Ptilozamites cycadea, Dictyophyllum sp. 1). nilssonii, Bjuvia simplex, Desmiophyllum sp., Since the climate of the Jurassic is known to Phlebopteris sp.) have been relatively stable (Vakhrameev 1991), 4. Ginkgoites group. Weakly disturbed microclimatic, palaeotopographic, or palaeo- wetland (Ginkgoites marginatus). geographic factors influenced the floristic com- 5. Komlopteris group. Moderately dis- position of particular localities. A genus-level 5 cluster analysis of European Jurassic localities same slab. The taxa were coded as binary (0-1) (Barbacka et al. 2014) placed the Reşiţa Basin variables (for details see Barbacka 2011, Bar- (including Anina) and the Mecsek Mts in the backa et al. 2014, Barbacka et al. in prep). In same branch: Reşiţa was paired with York- order to estimate the responses of particular shire (UK) and Mecsek with Scoresby Sound taxa along PCoA axis 1 and axis 2 we applied (Greenland), all being of the river-delta type a logistic regression model, the General Lin- of environment. On species level, Reşiţa and ear Model (GLM) using the logit link. A bino- Mecsek were in different clades, confirming mial distribution of the response variable a significant difference in their plant composi- was assumed (Agresti 2007). In that way the tion. Similarity of genus composition accompa- response variables (species) were related to nied by dissimilarity of species content is not a predictor – sample loadings along PCoA axis an unusual combination, as observed in a sta- 1 and axis 2 (Barbacka et al. in prep, Fig. 2). tistical approach to Mecsek flora (Barbacka Forward (stepwise) selection starting from the 2011). The same genera can occur in different null model was used to find the fitted model habitats, but species of the same genus almost for the particular species, based on the F-test always occupy different ecological niches (Bar- criterion and corresponding I-type error based backa 2011). on 499 runs. The calculations were performed In this paper we discuss the presumed with CANOCO 5 (ter Braak & Šmilauer 2012). taxonomical similarity between the two flo- Finally, seven species common to the two sites ras and the mechanisms governing local flo- were considered (see Table 1). GLMs for the ristical changes, in the light of environmental two localities revealed the response of the spe- variation. cies along the two PCoA axes and thus enabled us to classify them to particular response types. The group responses were interpreted MATERIAL AND METHODS as common occurrence in similar ecological conditions, that is, forming the putative eco- The comparison of the two floras from the Mecsek logical groups. Mts and Anina localities was based on samples stored at the Hungarian Natural History Museum (3256 samples belonging to 42 taxa, collected by Barbacka since 1989) and samples stored at the University RESULTS of Bucharest and the National Geological Museum (1384 samples belonging to 89 taxa, collected by Popa since 1990). ALPHA DIVERSITY Alpha diversity was estimated based on complete lists of taxa based on vegetative plant remains with The flora from Anina appears to be more quantitative values. Since the studied material from diverse than the flora from the Mecsek Mts. It Mecsek contains twice as many specimens as that contains 48 genera and 88 species belonging from Anina, the values are given as percentages, mak- to 9 plant groups (Popa 1992, 1998, 2000a, b, ing the data comparable. Taxa determined as ‘sp.’ are assigned letters for 2009, Popa & Van Konijnenburg-van Cittert Anina (sp. A, sp. B) and numbers for Mecsek (sp. 1, 2006, Popa & Meller 2009), as compared with sp. 2). 29 genera and 42 species from 8 plant groups (including one incertae sedis) in Mecsek (Bar- backa 1991, 1992, 1994a, b, 1997, 2000, 2001, STATISTICAL ANALYSES 2009, 2011, Barbacka & Bodor 2008, Thévé- nard & Barbacka 2000; Table 1, Fig. 1A, B). We used palaeobotanical databases of the The plant groups represented in both locali- 3256 samples from Mecsek and 1384 samples ties correspond to each other; lycopods and from Anina for the calculations. First the data Czekanowskiales additionally occur in the were ordinated separately for the two locali- material from Anina, while Desmiophyllum sp. ties by principal coordinate analysis (PCoA). (incertae sedis) is present in the material from Their distribution along axis 1 and axis 2 was Mecsek. The two localities have 25 genera in influenced by the similarities and differences common but only 9 species in common. in taxonomical composition. The data are for In terms of plant taxonomical groups, sphe- taxa based on vegetative plant remains that nophytes are represented in Anina by 2 genera co-occurred with at least one other taxon in the and 3 species, representing 13% of the total 6

Table 1. List of species in Mecsek and Anina localities. Common species in bold letters

Taxon Anina Mecsek Taxon Anina Mecsek Aninopteris formosa Popa 1 Komlopteris sp. 1 Anomozamites marginatus (Unger) Kylikipteris arguta Lindley et Hutton 20 31 Nathorst Kylikipteris sp. 1 Baiera furcata (Lindley et Hutton) Marattia (Marattiopsis) intermedia 57 4 Braun (Münster) Kilpper Baiera sp. 2 Marattia (Marattiopsis) sp. 3 Bjuvia simplex Florin 28 Marattiopsis hoerensis (Schimper) 45 Bjuvia sp. 2 Thomas Brachyphyllum sp. 1 18 Matonia braunii (Göppert) Harris 70 Brachyphyllum sp. 2 1 Neocalamites (Schizoneura) carcino- 142 46 Cladophlebis cf. aktashensis Turutanova- ides Harris 2 Ketova Neocalamites sp. A 8 Cladophlebis denticulata (Bron- Nilssonia obtusa (Nathorst) Harris 181 65 58 gniart) Fontaine Nilssonia polymorpha Schenk 30 Cladophlebis haiburnensis Lindley 30 36 Nilssonia revoluta Harris 18 et Hutton Nilssonia sp. A 78 Cladophlebis nebbensis (Brongniart) 58 Nilssonia sp. B 9 Nathorst Nilssonia sp. C 26 Cladophlebis roessertii (Schenk) Saporta 17 Nilssoniopteris sp. 4 Cladophlebis sp. 119 Osmundopsis cf. sturi (Raciborski) Harris 2 Cladophlebis sp. X 6 Otozamites sp. A 1 Clathropteris meniscoides Bron- 1 194 Otozamites gniart sp. B 1 Coniopteris hymenophylloides (Bron- Pachypteris banatica (Humml) Doludenko 1 2 Pachypteris rhomboidalis (Ettingshau- gniart) Seward 1 Coniopteris murrayana Brongniart 8 sen) Doludenko Pachypteris speciosa (Ettingshausen) Coniopteris sp. A 1 64 Andrae Coniopteris sp. B 52 Pachypteris sp. 2 Ctenis cf. grandifolia Fontaine 12 Pagiophyllum sp. 1 127 Ctenis sp. 1 Pagiophyllum sp. A 6 Cupressinocladus sp. 2 Pagiophyllum sp. B 7 Czekanowskia rigida Heer 1 Phlebopteris angustiloba (Presl) 1 75 Desmiophyllum sp. 76 Hirmer et Hörhammer Dicksonia sp. 6 Phlebopteris formosa Givulescu et Popa 9 Dictyophyllum nervulosum Kilpper 5 Phlebopteris polypodioides Brongniart 1 Dictyophyllum nilssonii (Brongniart) 60 56 Phlebopteris sp. 1 112 Göppert Phlebopteris sp. A 4 Dictyophyllum rugosum Lindley et Hutton 48 Phlebopteris woodwardii Leckenby 18 Dictyophyllum sp. 2 Phoenicopsis angustifolia Heer 7 Eboracia lobifolia (Philips) Thomas 1 Phoenicopsis sp. 10 Elatides sp. 6 Pinites sp. 37 Elatocladus sp. 1 366 Podozamites distans (Braun) Presl 7 Elatocladus sp. A 13 Podozamites lanceolatus (Lindley et Hut- 1 Equisetites columnaris Brongniart 53 ton) Braun Equisetites muensteri (Sternberg) Harris 56 Podozamites paucinervis Boersma et Van 43 Equisetites sp. 1 24 Konijnenburg-van Cittert Equisetites sp. A 12 Podozamites sp. 1 29 Geinitzia sp. 17 Podozamites sp. A 11 Ginkgoites marginatus (Nathorst) Florin 121 Pseudoctenis sp. 1 1 Ginkgoites minuta (Nathorst) Harris 1 Pseudoctenis sp. A 1 Ginkgoites sp. 1 93 Pseudocycas sp. 2 Ginkgoites sp. A 41 Pterophyllum brevipenne Kurr 6 Hausmannia buchii (Andrae) Seward 5 Pterophyllum longifolium Brongniart 7 Hausmannia cf. dentata Oishi 2 Pterophyllum subaequale (Hartz) Harris 9 Hausmannia sp. 1 Pterophyllum sp. 24 Isoetites sp. 1 Ptilophyllum sp. 22 Komlopteris nordenskioeldii Ptilozamites cycadea (Berger) Schenk 2 140 2 552 (Nathorst) Barbacka Ptilozamites leckenbyi (Leckenby) Nathorst 1 7

Table 1. Continued common species (of the total 9) were ferns, but their shares differed between the two assem- Taxon Anina Mecsek blages: Cladophlebis denticulata (Anina 10.8% Raphaelia sp. 2 of ferns, Mecsek 6.9%), Clathropteris menis Sagenopteris sp. 1 312 coides (Anina 1 specimen, Mecsek 23.3% of all Sagenopteris sp. A 13 ferns), Dictyophyllum nilssonii (Anina 10%, Solenites sp. 5 Phlebopteris angustiloba Sphenobaiera dragastanii Givulescu 1 Mecsek 6.7%), and Sphenobaiera grandis Kilpper 3 (Anina 1 specimen, Mecsek 9% of ferns). Sphenobaiera leptophylla (Harris) Florin 21 Seed ferns (Fig. 5) are not very numerous Sphenobaiera longifolia (Pomel) Florin 31 in Anina, represented by 4 genera and 8 spe- Sphenobaiera spectabilis (Nathorst) 2 cies (Popa 1997b, Popa 2000a), but they have Florin a 6.7% share of the total number of speci- Sphenobaiera sp. 54 mens (Fig. 2A, B). In Mecsek the same 4 gen- Sphenopteris sp. 2 Storgaardia johannae nomen nudum 1 era include 4 species (Barbacka 1991, 1992, Storgaardia spectabilis Harris 3 1994a, b, 1997) but their share is 30.8% of the Storgaardia sp. 8 total number of specimens, indicating their Thaumatopteris brauniana Popp 18 94 dominance in the flora. In Anina the individual Thaumatopteris sp. 21 species are not frequent and the most numer- Todites goepertianus (Münster) Krasser 78 ous one is Pachypteris speciosa (74.4% of all Todites princeps Presl 15 seed ferns), while Komlopteris nordenskioeldii Todites sp. 1 Zamites aninaensis (Semaka) Givulescu 3 is represented by a single specimen. In Mecsek, Zamites schmiedelii Presl 34 Zamites sp. 14 A Taxa 89 42 summa of samples 1384 3256 40 genus 35 species number of specimens. In Mecsek, sphenophytes 30 account for 2 genera, 4 species and a 5% share 25 (Figs 2A, B, 3). The genera are the same but 20 the species are different. For Mecsek, 3 spe- 15 cies of Equisetites were noted (E. columnaris, 10 29.6% of sphenophytes; E. muensteri, 31.3%; 5 Equisetites sp. 1, 13.4%) and one species of Neo 0 calamites (Schizoneura) carcinoides, 25.7%.

For Anina there was one species, Equisetites Ferns

Cycads

Conifers

ycophytes

sp. A (7.4% of sphenophytes) and 2 species of Seed ferns

Ginkgophytes Neocalamites, but only one of them, N. carci Sphenophytes

Bennettitaleans noides (named here Schizoneura carcinoides), B Czekanowskiales occurred in a significant amount (87.7%). 16 Lycophytes were recorded only from Anina, genus 14 as one specimen of Isoetites sp. species Ferns (Fig. 4) are the most diverse in both 12 localities. In Anina they are represented by 10 16 genera and 35 species (Popa 1997a, 2001a, 8 Popa 2005, Givulescu & Popa 1994, 1998, 6 Popa & Van Konijnenburg-van Cittert 1999), 4 and the total number of specimens constitutes 2 43.4% of all the material (Fig. 2A, B); ferns 0 dominated the whole assemblage. In Mecsek,

9 genera and 14 species of fern were recorded Ferns

Cycads

Conifers

(Barbacka 2011, Barbacka & Bodor 2008, Seed ferns

Ginkgophytes

incertae sedis Bodor & Barbacka 2008), and the specimens Sphenophytes

Bennettitaleans formed 25.5% of all collected samples, the sec- Fig. 1. Numbers of genera and species from the major taxo- ond biggest group besides seed ferns. Four nomical plant groups (A) in Anina, (B) in Mecsek 8

A B 76; 2.3% 1; 0.08% 18; 1.4% 179; 5.5% 162; 12,6% 162; 12,6% Sph 541; 16.6% 832; 25.5% Sph F F 106; 8.2% SF SF Cy 324; 10.7% Cy 65; 5% Be Be G 40; 1.2% G 127; 9.8% Con 259; 7.9% Con Lyc incertae 86; 6.7% 560; 43.4% Czek sedis 1005; 30.8%

Fig. 2. Number of specimens and percentage of the major taxonomical plant groups (A) in Anina, (B) in Mecsek. Sph – sphenophytes, F – ferns, SF – seed ferns, Cy – cycads, Be – bennettitaleans, G – ginkgophytes, Con – conifers, Lyc – lycophytes, Czek – Czekanowskiales

Pachypteris is very rare (single specimen of Nilssonia sp. A (60% of cycads) is most numer- P. banatica) and Komlopteris occurs in large ous in Anina; Nilssonia obtusa represents numbers (54.9% of seed ferns). Sagenopteris 69.9% of the cycad material in Mecsek. Bjuvia sp. is also rare in Anina but in Mecsek it has simplex is more frequent in Mecsek (10.8%); a 31% share of seed ferns. Ptilozamites cycadea Bjuvia sp. accounts for only 2 specimens in is much more frequent in Mecsek (13.9%, ver- Anina. sus 2 specimens in Anina). This group shows Bennettitaleans (Fig. 7) are not very fre- the largest quantitative disproportions in the quent in either locality. Although in Anina presence of a given taxon. (Popa 2001b, 2014) they are quite diverse Cycads (Fig. 6) are present in almost the (6 genera, 12 species), their number is not same proportions in the two localities: in high (8.4% of the whole flora). In Mecsek there Anina, 6 genera and 10 species (Popa & Van are only 2 genera (one of them, Pterophyllum, Konijnenburg-van Cittert 2006), constituting shared with Anina) and 2 species (both differ- 9.8% of the total number of specimens (Fig. 2 ent from Anina), together forming 1.2% of the A, B); in Mecsek, 4 genera and 6 species (Bar- entire material (Fig. 2A, B). backa 2001), forming 7.9% of the total material. Ginkgophytes (Fig. 8) are less diverse: 4 genera and 7 species were recorded in Anina 160 Anina (Popa & Van Konijnenburg-van Cittert 2006), 140 7.8% of the whole flora, while in Mecsek (Bar- Mecsek 120 backa 2002) there were 3 genera and 6 species noted (10%, Fig. 2A, B). All genera from Mec- 100 sek are also present in Anina, but there are no 80 common species. In Anina the commonest is 60 Sphenobaiera sp. A (50% of ginkgophytes); in 40 Mecsek the most numerous is Ginkgoites mar 20 ginatus (37.3%). Conifers (Fig. 9) in Anina were repre- 0

) sented by 9 genera and 15 species, 11.7% of

A A the whole flora (Fig. 2A, B). In Mecsek, 4 gen-

sp.

sp. sp. era and 5 species were recognised (Barbacka 2011, Thévénard & Barbacka 2000), 16.6% of

Schizoneura ( the total number of specimens. In Anina the

Equisetites Equisetites most common conifer was Podozamites pauci

Neocalamites

carcinoides

Equisetites muensteri

Equisetites columnaris nervis (26.5%), and in Mecsek Elatocladus sp. (67.7%). Neocalamites One specimen of Czekanowskia rigida indi- Fig. 3. Sphenophytes represented in Anina and Mecsek cates the presence of the order Czekanowskiales 9

Todites sp. Todites princeps

Anina Todites goeppertianus

Mecsek Thaumatopteris sp. Thaumatopteris brauniana Sphenopteris sp.

Raphaelia sp. Phlebopteris sp. 1

Phlebopteris sp. B Phlebopteris sp. A Phlebopteris woodwardii

Phlebopteris polypodioides Phlebopteris formosa

Phlebopteris angustiloba Osmundopsis cf. sturii

Matonia braunii Marattiopsis (Marattia) sp. B

Marattiopsis (Marattia) sp. A Marattiopsis (Marattia) intermedia Marattiopsis hoerensis

Kylikipteris sp. Kylikipteris arguta

Hausmannia sp.

Hausmannia cf. dentata Hausmannia buchii Eboracia losifolia Dictyophyllum sp. Dictyophyllum rugosum Dictyophyllum nilssonii

Dictyophyllum nervulosum Dicksonia sp. Coniopteris sp. B

Coniopteris sp. A

Coniopteris murrayana Coniopteris hymenophylloides

Clathropteris meniscoides Cladophlebis sp. X Cladophlebis sp. A

Cladophlebis roessertii

Cladophlebis nebbensis Cladophlebis haiburnensis

Cladophlebis denticulata Cladophlebis cf. aktashensis 0 0 90 80 70 60 50 40 30 20 10

200 190 180 170 160 150 140 130 120 Fig.11 100 4. Ferns represented in Anina and Mecsek 10

552 181 312 150 150 140 140 Anina Anina 130 130 Mecsek Mecsek 120 120 110 110 100 100 90 90 80 80 70 70 60 60 50 50 40 40 30 30 20 20 10 10 0

0 1

sp.

Bjuvia

Ctenis

grandifolia

cf.

Bjuvia simplex

Nilssonia

Nilssonia obtusa

Pseudocycas

Pachypteris

Komlopteris

Nilssonia revoluta

Pseudoctenis

Sagenopteris

Ctenis

Sagenopteris

Nilssonia polymorpha

Pachypteris banatica

Ptilozamites cycadea

Pachypteris speciosa

Ptilozamites leckenbyi

Pachypteris rhomboidalis

Komlopteris nordenskioeldii

Fig. 5. Seed ferns represented in Anina and Mecsek Fig. 6. Cycads represented in Anina and Mecsek

121 93

40 60 Anina 35 Anina 50 Mecsek Mecsek 30 40 25

20 30

15 20 10 10 5

0 0

sp.

Zamites

Baiera

Solenites

Nilssoniopteris

Baiera furcata

Ptilophyllum

Pterophyllum

Ginkgoites

Otozamites

Zamites aninensis

Ginkgoites minuta

Zamites schmiedelii

Sphenobaiera

Sphenobaiera grandis

Ginkgoites marginatus

Sphenobaiera longifolia

Pterophyllum longifolium

Pterophyllum brevipenne

Sphenobaiera spectabilis

Sphenobaiera leptophylla

Pterophyllunm subaequale

Anomozamites marginatus

Sphenobaiera dragastanii

Fig. 7. Bennettitaleans represented in Anina and Mecsek Fig. 8. Ginkgophytes represented in Anina and Mecsek 11

366 127 AninaAnina

MecsekMecsek 50

sp.

Pinites

Elatides

Geinitzia

Storgaardia

Elatocladus

Podozamites

Pagiophyllum

Podozamites

Pagiophyllum

Brachyphyllum

Podozamites distans

Cupressinocladus

Storgaardia johannae

Storgaardia soectabilis

Podozamites paucinervis

Podozamites lanceolatus Fig. 9. Conifers represented in Anina and Mecsek in Anina (Popa 1992), which in Mecsek is In Anina the 7 shared species were distrib- absent. Desmiophyllum sp. was not observed uted in two groups: Podozamites and Schizo in Anina but was noted in Mecsek as Gymno- neura (Table 3). In the Podozamites group, spermae incertae sedis, forming 2.3% of the Cladophlebis haiburnensis accounted for 17.0– entire flora. 17.1% of the variance along both PCO axes; along PCO axis 2, Neocalamites (Schizoneura) GLM OF COMMON SPECIES carcinoides accounted for 27% and Dictyophyl lum nilssonii for 83.7% of the total variance. Seven of the 9 taxa common to the two These species correspond to the Ptilozamites localities were used in the analyses; 2 species and Thaumatopteris ecogroups from Mecsek. (Phlebopteris angustiloba and Clathropteris In the Schizoneura group only Cladophlebis meniscoides) were represented by only one denticulata had a statistically significant GLM specimen in Anina. response, accounting for 13.5% and 3.6% of the In Mecsek these 7 species were distrib- total variance along PCO axes 1 and 2 respec- uted in four groups (according to ecogroup tively. This species corresponds to the Sagen as defined by Barbacka 2011): Komlopte opteris group from Mecsek. ris, Sagenopteris, Ptilozamites, and Thau The remaining species (5 from Mecsek, matopteris. Among them, only Ptilozamites 3 from Anina), though concordant in their cycadea and Komlopteris nordenskioeldii, response profiles, did not have statistically sig- found in the common Komlopteris group, nificant responses (p < 0.05, Figs 10, 11). showed statistically significant (p ≤ 0.05) GLM responses (Table 2). Komlopteris nor denskioeldii accounted for 67.0–74.06% of the DISCUSSION total variance and P. cycadea only 1.3% along PCO axis 1. The species within the remain- Cluster analyses of European Jurassic flo- ing groups had statistically non-significant ras (Barbacka et al. 2014) produced groupings (p > 0.05) GLM responses which accounted for of localities with similar taxonomical composi- < 0.1–2% of the total variance. tions. The analyses were done on both genus 12

Table 2. Site A. Stepwise selection of response variables (species in groups discriminated in PCA analysis) along PCO axes 1 and 2, , according to logistic regression model with logit link function and binomial response assumed. R2 – coefficient of determination, F – a partial F test, p – I-type error

Response R2[%] F p R2[%] F p Ordination PCO 1 PCO 2 Komlopteris Group Ptilozamites cycadea 1.3 5.2 0.0223 < 0.1 < 1 0.63854 Komlopteris nordenskioeldii 74.6 791.9 < 0.00001 67.0 710.4 < 0.00001 Sagenopteris Group Cladophlebis denticulata 1.2 < 1 0.26045 < 0.1 < 1 0.74282 Ptilozamites Group Cladophlebis haiburnensis 2.0 < 1 0.30447 < 0.1 < 1 0.88202 Dictyophyllum nilssonii 0.1 < 1 0.60082 < 0.1 < 1 0.78458 Thaumatopteris Group Thaumatopteris brauniana 0.2 < 1 0.57259 1.0 4.0 0.04661 Neocalamites (Schizoneura) carcinoides 0.1 < 1 0.72214 0.7 1.4 0.23091

Table 3. Site B. Stepwise selection of response variables (species in groups discriminated in PCA analysis) along PCO axes 1 and 2, according to logistic regression model with logit link function and binomial response assumed. R2 – coefficient of determination, F – a partial F test, p – I-type error

Response R2[%] F p R2[%] F p Axis PCO 1 PCO 2 Group 1 Cladophlebis haiburnensis 17.1 12.2 0.00056 17.0 12.2 0.00057 Dictyophyllum nilssonii 73.9 125.5 < 0.00001 83.7 142.1 < 0.00001 Neocalamites (Schizoneura) carcinoides 0.1 < 1 0.64899 27.0 42.1 < 0.00001 Thaumatopteris brauniana 0.5 < 1 0.51394 1.1 1.0 0.31199 Group 2 Cladophlebis denticulata 13.5 24.7 < 0.00001 3.6 6.60 0.01054 Komlopteris nordenskioeldii 1.0 < 1 0.71227 1.3 < 1 0.68558 Ptilozamites cycadea 1.4 < 1 0.73992 0.8 < 1 0.66795 and species levels. The genus cluster tended Bearing in mind that 25 genera but only to separate delta and fluvial ecosystems from 9 species are shared between the two locali- coastal/lagoonal ones. This suggests that the ties, we stress that for the present case the delta and fluvial environments maintained much more accurate palaeoenvironment indi- similar conditions for the corresponding genera. cator is species-level taxonomy rather than Within the genus cluster, Mecsek is on the genus-level taxonomy. same branch with Scoresby (Lower Jurassic, A statistical study of the flora from Mecsek Greenland), while Reşiţa (Anina, with some (Barbacka 2011) showed that species of the smaller localities from the same unit), together same genera had their maximum occurrence with Yorkshire (the Middle Jurassic, UK), in different ecogroups. This suggests that fine occupies the neighbouring final branches. In differences in conditions led to differentiation the species cluster, however, the two localities of species composition, while the same genera are far from each other, on different primary appeared in similar types of ecosystems. Our branches. comparison of the ecogroups distinguished in When genera are considered without quanti- Anina and Mecsek, and of their taxonomical tative data, the two localities seem similar, but structure, supports this supposition. In both a quantitative comparison at species level shows localities, ecogroups of pioneer type are pre- 8 species in common; apart from them, only the sent, consisting exclusively of ferns (in Anina ferns Cladophlebis denticulata and Dictyophyl the Schizoneura group, in Mecsek the Thau lum nilssonii occur in relatively equal numbers, matopteris group), but in Anina there are , spe- and the amounts of the remaining species are cies that do not occur in Mecsek: Cladophlebis very disproportionate. Both of these fern spe- nebbensis, Dictyophyllum nervulosum, Coniop cies are widespread, especially Cladophlebis teris murrayana, and Matonia braunii. Their denticulata (Barbacka et al. 2014). environmental preferences tend towards open 13

Komlopteris nordenskioeldii Komlopteris nordenskioeldii

1.0 a 1.0 b

Ptilozamites cycadea

Response Group Komlopteris

0.0

0.0 -2 PCO 1 3 -3 PCO 2 3

1.0 c 0.8 d

Cladophlebis haiburnensis Dictyophyllum nilssonii

Ptilozamites Group Cladophlebis haiburnensis Dictyophyllum nilssonii

0.0

0.0 -2 3 -3 3

1.0 Neocalamites 1.0 (Schizoneura) carcinoides e f Neocalamites (Schizoneura) carcinoides

Thaumatopteris

Thaumatopteris brauniana

Response Group

Thaumatopteris brauniana

0.0

0.0 -2 3 -3 3

0.06 g 0.030 h

Cladophlebis denticulata

Cl.dent.

Sagenopteris Group

Cladophlebis denticulata

0.01

0.018 -2 PCO 1 3 -3 PCO 2 3

Fig. 10. Logistic regressions of GLM for response variables (species) in four palaeoecological species groups along PCO 1 and PCO 2. a – Komlopteris, Ptilozamites – PCO 1; b – Komlopteris, Ptilozamites – PCO 2; c – Dictyophyllum, Cladophlebis – PCO 1; d – Dictyophyllum, Cladophlebis – PCO 2; e – Neocalamites (Schizoneura), Thaumatopteris – PCO 1; f – Neocalamites, Thaumatopteris – PCO 2; g – Cladophlebis denticulata – PCO 1; h – Cladophlebis denticulata – PCO 2. The same pattern of response (decreasing or increasing) in a group of species means their ecological profile are similar. Differences in shapes of response curves depend on the distribution and abundance of species along a putative environmental gradient 14

1.0 Dictyophyllum nilssonii a Dictyophyllum nilssonii b

Neocalamites (Schizoneura) 1 carcinoides

Neocalamites (Schizoneura) carcinoides Response Group Thaumatopteris brauniana Thaumatopteris brauniana Cladophlebis Cladophlebis haiburnensis haiburnensis

0.0 -2 PCO 1 3 -3 PCO 2 3

0.6 Ptilozamites cycadea c d Komlopteris sp.

Ptilozamites

Response Group cycadea

Cladophlebis Cladophlebis denticulata Komlopteris sp. denticulata

0.0 -2 3 -3 3

Fig. 11. Logistic regressions of GLM for response variables (species) in four palaeoecological species groups along PCO 1 and PCO 2. a – Species Group 1 – PCO 1; b – Species Group 1 – PCO 2; c – Species Group 3 – PCO 1; d – Species Group 3 – PCO 2. The same pattern of response (decreasing or increasing) in a group of species means their ecological profile are similar. Differences in shapes of response curves depend on the distribution and abundance of species along a putative environmental gradient plains or clearings, more typical of periodi- moisture provided favourable conditions for cally inundated flood plains, while the pioneer swampy habitats to develop. This increase of assemblage from Mecsek was interpreted temperature is comparable to the Hettangian as growing mainly on easily eroded channel interval in Mecsek, when mires also occurred banks or floating islands (Barbacka 2011). but with slightly different phytocoenoses (Popa The moderately wet and weakly disturbed 2000b, 2009, Popa & Van Konijnenburg-van habitat represented in Mecsek by the Ptilo Cittert 2006). The plant assemblages from zamites group corresponds to the moderately this stage are interpreted as characteristic for wet and disturbed habitat in Anina occupied the closing drier phase of mire development. by the Podozamites group, but the taxonomi- Probably this phase does not correspond fully cal composition of these habitats differs com- with the Hettangian swamp phase from Mec- pletely between the two localities. The differ- sek. This phase was manifested in taxa assem- ences probably are due to differences in the blages which in Mecsek are connected to both climatic conditions of these localities, related habitats: swampy (Komlopteris nordenskioel mainly to mean annual temperature. This dii or possibly Baiera sp.; in Anina they occur ecogroup in Mecsek was connected with cal- in trace amounts) and slightly drier (Clad careous river cliffs with at least periodic semi- ophlebis denticulata or Ginkgoites sp., Ptilo arid conditions and high insolation (Barbacka zamites cycadea). The remaining taxa from 2011). In Anina, the Reşiţa Basin is bordered this ecogroup (e.g. Zamites schmiedelii, Gein by crystalline heights of the Sebes-Lotru meta- itzia sp., Ptilophyllum sp.) occur in Anina and morphic series and partly by Variscan (Upper correspond well with the described conditions. Carboniferous – Lower Permian) promontories, Relatively dry, moderately warm and undis- contributing to different edaphic conditions. turbed levees in Anina were occupied mainly In Anina during the Sinemurian, the by Nilssonia sp. This habitat in Mecsek would increase of temperature and relatively high correspond to the Sagenopteris ecogroup with 15

Nilssonia obtusa, N. revoluta, Cladophlebis linked with axis 2: temperature, which did not denticulata, Marattiopsis hoerensis, Sagenop play a role in Mecsek (in the literature, no teris sp., and Anomozamites marginatus. In mention of temperature change in Mecsek dur- Mecsek this type of environment was inhab- ing the Hettangian and Sinemurian). Hence, ited by a very developed plant succession. The the types of GLM responses in particular spe- monospecific character of the Anina site might cies groups are not represented by the same be due to edaphic factors. ecological groups. For example, Neocalamites In Mecsek the monospecific group of Gink (Schizoneura) carcinoides and Thaumatopt goites marginatus is explained possibly by eris brauniana reacted differently in Mecsek expansive growth, and by analogy with Ginkgo and in the opposite direction to Dictyophyllum biloba with its moderate allelopathy (Nam et nilssonii and Cladophlebis haiburnensis, while al. 1997, Barbacka 2011). in Anina they show the same preferences. Sim- The occurrence of species shared by the ilarly, Cladophlebis denticulata, which in Mec- localities makes it possible to compare their sek was opposite to Komlopteris nordenskioel joint responses. While in Mecsek they represent dii and Ptilozamites cycadea, had the same four ecogroups of putatively different ecological tendencies as the latter two in Anina. Only profiles, in Anina they are distributed in two Komlopteris nordenskioeldii and Ptilozamites groups (Schizoneura and Zamites groups). cycadea showed similar trends for both locali- The Schizoneura group in Anina encom- ties and were similarly opposite to Dictyophyl passes the Ptilozamites and Thaumatopteris lum nilssonii and Cladophlebis haiburnensis. ecogroups from Mecsek, both well supported For Mecsek, Thaumatopteris brauniana statistically. This could be an effect of having and Neocalamites (Schizoneura) carcinoides similar ecological niches in Anina. The Zamites were interpreted as connected to the pioneer group also encompasses two ecogroups previ- assemblage of a highly disturbed and mod- ously recognised in Mecsek, namely Sagen erately wet habitat. According to a previous opteris and Komlopteris (Barbacka 2011). Of analysis, Dictyophyllum nilssonii and Clad these, only Sagenopteris was supported statis- ophlebis haiburnensis were in one putative tically. This could mean that the Komlopteris ecogroup with Thaumatopteris brauniana group was in fact ecologically distinct from (Barbacka 2011). In another analysis employ- Sagenopteris and that its joint presence here ing PCA ordination, pairs of species differed was a chance event. This fully corresponds in their preferences in the Mecsek locality. In with our knowledge of the plant assemblages Anina they formed one ecogroup (Schizoneura) from Mecsek. Komlopteris nordenskioeldii was in PCA (Barbacka et al. in prep). It was asso- in fact a species of very wide tolerance, and ciated with high moisture/high disturbance its co-occurrence with taxa from different eco- and relatively low temperature, interpreted as groups is highly probable (Barbacka 2011). a flood plain association. For the statistical analyses we assumed Ptilozamites cycadea and Komlopteris nor that common GLM responses along the ordi- denskioeldii are numerous in Mecsek; the lat nation axes denote species with the same eco- ter is the most numerous fossil in this local- logical requirements. Figures 10 and 11 show ity, but in Anina they are very rare (Zamites the curves for shared species with the same group). Only Cladophlebis denticulata from requirements. For example, the Komlopteris this ecogroup occurs in almost equal amounts Group (Fig. 10a, b) differs from the Thau at both sites. In Mecsek, Ptilozamites cycadea matopteris Group (Fig. 10e, f) by the difference indicates moderately wet and undisturbed in response along axis 2. An exact comparison habitat, while Komlopteris nordenskioeldii is hampered by probable differences in the prefers wetter, swampy places. However, the main environmental factors influencing the latter’s wide tolerance enables it to appear in plant composition of the two localities. Accord- different ecogroups. In Mecsek, Cladophlebis ing to a previous interpretation (Barbacka denticulata was associated with rather drier 2011), two factors were decisive in Mecsek: the and moderately disturbed conditions; in Anina humidity gradient according to axis 1 of the it belongs to an association of swampy habi- plot, and the disturbance gradient according tat but in the last, not so wet, closing stage to axis 2. In Anina, humidity and disturbance of mire development (Barbacka 2011, Popa were together on axis 1, and a third factor was 1998, 2014). Since both K. nordenskioeldii and 16

Ptilozamites cycadea are very rare in Anina, BARBACKA M. 1991. New data about Liassic flora their real preferences in this area cannot be in the Mecsek Mountains. Annls. Hist.-Nat. Mus. unambiguously interpreted. Natural. Hung., 83: 17–23. Species from the Zamites group in Anina BARBACKA M. 1992. The Liassic seed ferns of the first appeared in the Sinemurian after an Mecsek Mountains (S. Hungary): 257–263. In: Kovar-Eder J. (ed.), Palaeovegetational development increase of temperature, when the climate in Europe and regions relevant to its palaeoflo- became warmer and slightly more humid than ristic evolution. Proc. Pan. Eur. Palaeobot. Conf., during the Hettangian (Popa 1998, 2000b, Vienna, 19–23 Sept. 1991. 2009). In general, the species common to both BARBACKA M. 1994a. Komlopteris Barbacka nov. localities show similar trends, occupying cor- gen., a segregate from Pachypteris Brongniart. responding habitats within two similar though Rev. Palaeobot. Palynol., 83: 339–349. not identical environments. BARBACKA M. 1994b. Pachypteris banatica from the Mecsek Mountains Liassic. Acta Palaeobot., 34(1): 5–19. CONCLUSIONS BARBACKA M. 1997. Ctenozamites cycadea (Berger) Schenk from the Mecsek Mountains Liassic – 1. The environments of Mecsek (delta – S. Hungary. Mededel. Nederl. Inst. Toegep. Geovet. TNO, 58: 81–85. limnic – lagoonal) and Anina (fluvial) gener- ally differ but have similar topographic ele- BARBACKA M. 2000. Bennettitales from the Mec- sek Mountains Liassic, Hungary. Acta Palaeobot., ments such as river/channel banks, flooded or 40(2): 111–127. swampy areas, lakes, river levees, or marine barriers (Mecsek). BARBACKA M. 2001. The cycads of Hungarian Lias- sic. Rev. Paléobiol., 20(2): 525–541. 2. Although the generic composition of the floras seems similar, fine differences in local BARBACKA M. 2002. The Liassic Ginkgoales from the Mecsek Mountains, Hungary. Rev. Paléobiol., conditions led to significant quantitative and 21(2): 697–715. qualitative dissimilarity of species composi- BARBACKA M. 2009. Sphenophyta from the Early tion. Jurassic of the Mecsek Mts., Hungary. Acta Palae 3. The eight common species do not occur obot., 49(2): 221–231. in strictly corresponding ecogroups but their BARBACKA M., 2011. Biodiversity and the recon- environmental preferences are similar. All of struction of Early Jurassic flora from the Mecsek them show quite wide tolerance. Mountains (southern Hungary). Acta Palaeobot., 4. Differences in climate (mean annual 51: 127–179. temperature, humidity) and edaphic condi- BARBACKA M. & BODOR E. 2008. Systematic and tions (different provenance areas differing in palaeoenvironmental investigations of fossil ferns petrographic composition) explain the differ- Cladophlebis and Todites from the Liassic of Hun- ences between floras at species level under gary. Acta Palaeobot., 48(2): 133–149. similar genus-level composition. BARBACKA M., BODOR E., JARZYNKA A., KUS- TATSCHER E., PACYNA G., POPA M.E. SCANU G.G., THÉVENARD F. & ZIAJA J. 2014. European ACKNOWLEDGEMENTS Jurassic floras: statistics and palaeoenvironmental proxies. Acta Palaeobot., 54(2): 173–195. This study was supported by the Hungarian National Science Foundation (grant OTKA 100658) BODOR E. & BARBACKA M. 2008. Notes on some and by the W. Szafer Institute of Botany, Polish Acad- doubtful taxonomical aspects in the genera Clado emy of Sciences though its statutory funds. Part of this phlebis Brongniart and Todites Seward. Palaeo- research (by Grzegorz Pacyna) received support from world, 17(3–4): 201–214. the SYNTHESYS Project http://www.synthesys.info/, BUCUR I.I. 1991. Proposition pour une nomenclature which is financed by European Community − Research formelle des depots paleozoiques et mesozoiques de Infrastructure Action under the FP7 “Capacities” Spe- la zone de Reşiţa-Moldova Nouă (Carpathes Merid- cific Programme. ionales, Roumanie). Studia Universitatis Babeş- Bolyai, Geologie, 36: 3–14. BUCUR I.I. 1997. Formaţiunile mesozoice din REFERENCES zona Reşiţa-Moldova Nouă. Presa Universitară Clujeană, Cluj-Napoca. AGRESTI A. 2007. An introduction to categorical data GIVULESCU R. & POPA M.E. 1994. Eine neue Dic analysis. 2nd Ed. John Wiley & Sons, Inc., Hoboken, tyophyllum – Art aus dem unteren Lias von Anina New Jersey. (Rumänien). Documenta Naturae, 84: 42–46. 17

GIVULESCU R. & POPA M.E. 1998. Aninopteris for POPA M.E. 2001b. Aspects of Romanian Early Juras- mosa Givulescu et Popa, gen. et sp. nov., a new sic palaeobotany and palynology. Part IV. A new Liassic matoniaceous genus and species from species of Weltrichia from Anina. Studia Universi- Anina, Banat, Romania. Rev. Palaeobot. Palynol., tatis Babe-Bolyai, Geologie, 46: 69–76. 104: 51–66. POPA M.E. 2005. Aspects of Romanian Early Jurassic KĘDZIOR A. & POPA M.E. 2013. Sedimentology of Palaeobotany and Palynology. Part VI. Anina, an the Early Jurassic terrestrial Steierdorf Formation exceptional locality. Acta Palaeont. Romaniae, 5: in Anina, Colonia Cehă Quarry, South Carpath- 375–378. ians, Romania. Acta Geol. Pol., 63(2): 175–199. POPA M.E. 2009. Late Palaeozoic and Early Mesozoic NAGY E. & NAGY J. 1969. Stratigraphie. In: Nagy E. continental formations of the Reşiţa Basin. Editura (ed.), Unterlias-Kohlenserie des Mecsek-Gebirges, Universităţii din Bucureşti, Bucharest. Geologie. Ann. Inst. Geol. Publ. Hung. 51(2): POPA M.E. 2014. Early Jurassic bennettitalean repro- 280–287. ductive structures of Romania. Palaeobiodiversity NAM S.J., KIM K.U., SHIN D.H. & HWANG S.J. 1997. and Palaeoenvironments, 94: 327–362. Identification of biologically active substances from POPA M.E. & KĘDZIOR A. 2008. High resolution Ginkgo biloba L. Kor. J. Weed Sci., 17(4): 421–430. paleobotany and sedimentology of the Steier- PAÁL-SOLT M. 1969. Kohlenpetrographie. In: Nagy E. dorf Formation, Reşiţa Basin: 57–59. In: Bucur (ed.), Unterlias-Kohlenserie des Mecsek-Gebirges, I.I. & Filipescu S. (eds), Annual scientific session Geologie. Ann. Inst. Geol. Publ. Hung., 51(2): “Ion Popescu Voiteşti”. Cluj University Press, Cluj- 473–575. Napoca. POPA M.E. 1992. The Early Liassic of Anina: new POPA M.E. & Van KONIJNENBURG-van CITTERT paleobotanical aspects. Documenta Naturae, 1–3: J.H.A. 1999. Aspects of Romanian Early Jurassic 1–9. palaeobotany and palynology. Part I. In situ spores POPA M.E. 1997a. Liassic ferns from the Steierdorf from the Getic Nappe, Banat, Romania. Acta Pal- Formation, Anina. In: Herngreen G.F.W. (ed.), Pro- aeobot., Suppl. 2: 181–195. ceedings 4th European Palaeobotanical and Paly- POPA M.E. & Van KONIJNENBURG-van CITTERT nological Conference. Meded. Nederl. Inst. Toegep. J.H.A. 2006. Aspects of Romanian Early – Middle Geowetens. TNO, 58: 139–148. Jurassic palaeobotany and palynology. Part VII. POPA M.E. 1997b. Corystospermal pteridosperms in Successions and floras. Progr. Natur. Sci., 16: the Liassic continental deposits of Romania. Acta 203–212. Palaeont. Romaniae, 1: 81–87. POPA M.E. & MELLER B. 2009. Review of Juras- POPA M.E. 1998. The Liassic continental flora of sic plants from the Anina (Steierdorf) coal mining Romania: Systematics, Stratigraphy and Paleo- area, South Carpathians, in the collections of the ecology. Acta Botanica Horti Bucurestensis, 1997– Geological Survey of Austria. Jb. Geol. Bundesan- 1998: 177–184. stalt, 149: 487–498. POPA M.E. 2000a. Early Jurassic land flora of the TER BRAAK C.J.F. & ŠMILAUER P. 2012. Canoco Getic Nappe. Faculty of Geology and Geophysics, reference manual and user’s guide: software for University of Bucharest, Bucharest. ordination (version 5.0). Microcomputer Power, Ithaca, NY, USA. POPA M.E. 2000b. Aspects of Romanian Early Juras- sic palaeobotany and palynology. Part III. Phyto- THEVENARD F. & BARBACKA M. 2000. Two leaf stratigraphy of the Getic Nappe. Acta Palaeontol. morphotypes of the Pagiophyllum peregrinum Romaniae, 2: 377–386. (Lindley et Hutton) Schenk emend Kendall from the Mecsek Mountains, Hungary. Acta Palaeobot., POPA M.E. 2001a. Ponor SSSI (Site of Special Scien- Suppl., 2: 219–231. tific Interest). Lower Jurassic Paleoflora: 167–171. In: Bucur I.I., Filipescu S. & Săsăran E. (eds), VAKHRAMEEV V.A. 1991. Jurassic and Cretaceous Algae and carbonate platforms in western part of floras and climates of the Earth. Cambridge Uni- Romania. Field trip guidebook. Babes-Bolyai Uni- versity Press, Cambridge. versity, Cluj-Napoca.