40 5.2.1 Terrestrial Palynomorphs Laevigate Trilete Spores

5.2.1 Terrestrial Palynomorphs

Laevigate trilete spores

Genus: Aulisporites LESCHIK 1954

Aulisporites astigmosus (LESCHIK 1956a) KLAUS 1960

1956 Calamospora astigmosa sp. nov . – LESCHIK, p. 22, Plate 2 Fig. 17 1960 Aulisporites astigmosus (Leschik) nov. Comb . – K LAUS , p. 119 - 120, Plate 28, fig. 2.

Genus: Calamospora SCHOPF , WILSON & B ENTALL 1944

Calamospora tener (LESCHIK 1955) DE JERSEY 1962

1955 Laevigatisporites tener sp. nov. – LESCHIK , p. 13, Plate 1., Fig. 20. 1955 Punctatisporites flavus – LESCHIK , p. 31, Plate 4, Fig. 2. 1958 Calamospora mesozoicus – COUPER , p. 132, Plate 15/3+4. 1960 Calamospora nathorstii – KLAUS , p. 116, Plate 28, Fig. 1. 1962 Calamaospora tener (L ESCHIK ) n. comb. - de Jersey, p. 3-4, Plate 1, fig 9 –10. 1964 Calamospora tener (LESCHIK 1955) n. comb. – MÄDLER (a), p. 92, Plate 8, Fig. 2.

Genus: Cingulizonates DYBOVA & J ACHOWICS 1957

Cingulizonates rhaeticus (R HEINHARDT ) S CHULZ 1967

1962 Cingulatizonates rhaeticus sp. nov. – RHEINHARDT , P. 702, P LATE 2 F IG . 3 1964 Anulatisporites drawehni MAEDLER , P. 177, P LATE 2, F IGS . 1 – 2 1966 Cingulatizonates delicatus ORLOWSKA – Z WOLINSKAP I 014, PLATE 7 FIGS 36 - 38 1967 Cingulatizonates rhaeticus – SCHULZ P. 584, PLATE .13, F IG . 6- 7

Genus: Concavisporites THOMSON & P FLUG 1953 p. 49, Plate 1, Fig. 19

1953 Concavisporites gen. nov. – THOMSON & P FLUG , p. 49. 1959 Toroisporites gen. nov. – KRUTZSCH ,, p. 90.

Concavisporites crassexinius NILSSON 1958 p.35, Plate 1, Fig. d

1958 Concavisporites crassexinius sp. nov. – NILSSON , p . 35, Plate 1, Fig. 11.

Concavisporites mesozoicus sensu BÓNA

Comment: Sporomorphs described by Bóna as C. mesozoicus are comparable to Concavisporites variverrucatus described by COUPER 1958.

Concavisporites parvulus sensu BONA The spores, described by Bóna as C. parvulus with a diameter of 20-24 µm are smaller as the other species of this genus.

Concavisporites rhaetoliassicus ACHILLES 1981 1981 Concavisporites rhaetoliassicus sp. nov. – ACHILLES , p. 13 –14 , Plate 1, Fig. -13 - 15.

Genus: Cornutisporites SCHULZ 1962 p. 310, plate 1, Fig. 7 - 8

Cornutisporites seebergensis SCHULZ 1962

1967 Cornutisporites seebergensis SCHULZ , p. 310, Plate 1, Fig. 7 - 8.

Genus: Cyathidites COUPER 1953 p. 27, Plate 2, Fig. 11, 12.

Cyathidites australis COUPER 1953

1953 Lygodiosporites adriennis POTONIE & G ELLETICH f. mesozoicus Thiergart; COOKSON , p. 470, Plate 2, Fig. 29. 1953 Cyathidites australis COUPER , p. 27, Plate 2, Fig. 11, 12. 1961 Lygodiumsporites adriennis POTONIE & G ELLETICH ; B OLKOVITINA , p. 104-105, Plate 31, Fig. 3e (illustration of specimen figured by COOKSON 1953). 1961 Cyathidites australis COUPER ; Dev, p. 43, Plate 1, Fig. 1. 1963 Leiotriletes sp. Type A.; SAAD , p. 120, Plate 33, Fig. 1-5. 1963 Cyathidites australis COUPER ; DETTMANN , p. 22, Plate 1, Fig. 1-3. 1965 Cyathidites cf. australis COUPER ; DORING , p. 19, Plate 3, Fig. 1-3. 1966 Cyathidites australis COUPER ; BURGER , p. 237, Plate 5, Fig. 2. 1968 Deltoidospora australis ; CHALONER & MUIR , Plate 16, Fig. a. 1969 Cyathidites australis COUPER ; NORRIS , p. 582, Plate 102, Fig. 1. 2003 Cyathidites australis COUPER ; CHEN & Z HANG , p. 305, Plate 1, Fig. 6-8. 2003 Deltoidospora australis (COUPER ) POCOCK ; QUATTROCCHIO , et al. p. 66, Plate 1, Fig. 1. 2004 Cyathidites australis COUPER ; CHEN & Z HANG , p. 205, Plate 1, Fig. 7, 8. 2006 Deltoidospora australis (COUPER ) POCOCK ; QUATTROCCHIO , et al. p. 594, Fig. 6E. 2006 Cyathidites australis COUPER ; MACPHAIL & CANTRILL , p. 619, Plate 1, Fig. 11. Natural affinity: Filicopsida; various genera and families; Cyatheaceae ( Cyathea ), Dicksoniaceae, Schizaeaceae (Lygodium ).

Cycathidites minor COUPER , 1953

1953 Cyathidites australis COUPER , p. 28, Plate 2, Fig. 13

Genus: Deltoidospora MINER 1935 p. 613 plate 24 fig. 7

Deltoidospora sp. sensu BONA

Description: Trilete spores, concavely triangular to subcircular; Y mark distinct, rays at least 2/3 radius; exine two-layered, smooth or infrapunctate, with or without exinal folds (kyrtome or less) along the Y mark; 25-80 µm.

Genus : Densosporites BERRY 1937 p. 157

Densosporites fissus ( REINHARDT 1964) S CHULZ 1967

1964 Densoisporites fissus sp.nov. REINHARDT p. 54 plate 2 fig 1 – 3 1964 Cingulatizonites rhaeticus MAEDLER 1964 p. 184 plate 2 fig. 18 - 19 1967 Densosporites fissus (R HEINHARDT ) comb. nov. SCHULZ 582, PLATE 12, FIG . 5 - 6

Densoisporites sp. Plate 1, Fig. c

Genus : Dictyophyllidites COUPER , 1958 p. 140 plate 21 fig. 5 - 6

Dictyophyllidites harrisii COUPER , 1958

1958 Dictyophyllidites harrisii COUPER , p. 140 plate 21 fig. 5 - 6

Original diagnosis: Trilete, laesurae long, distinct, commissures clearly raised, bordered by a distinct margo; equatorial contour triangular, distal surface markedly convex, proximal less so; exine smooth and comparatively thin. The genus is intended for dispersed spores of the type met within the Jurassic fern Dictyophyllum .

Genus: Leiotriletes NAUMOVA 1939 p. 355 emend. POTONIÉ & K REMP 1955

Leiotriletes sp.

Genus: Stereisporites THOMSON & P FLUG 1953 p. 53

Stereisporites spp. DE JERSEY & R AINE 1990

1990 Stereisporites sp.; DE JERSEY & RAINE , p. 22, Plate 1, Fig. L. Natural affinity: Bryophyta.

Genus Todisporites COUPER 1958 P. 134

Todisporites major COUPER 1958

1958 Todisporites major COUPER , p. 134, Plate 16, Fig. 6-8. 1964 Punctatisporites major (COUPER ) KEDVES & S IMONCSICS , p. 13, Plate 3, Fig. 1, 2. 1966 Todisporites major COUPER ; Helal, p. 86, Plate 31, Fig. 5. 1977 Punctatisporites major (COUPER ) DORHOFER , pp. 20-21, Plate 2, Fig. 10-12 (in part). (includes T. minor COUPER in synonymy) 1978 Todisporites major COUPER ; GUY -OHLSON , pp. 40-41, Plate 6, Fig. 58. 1988 Punctatisporites major (COUPER ) DORHOFER ; PONS , p. 82, Plate 17, Fig.7. 1989 Todisporites cf. major COUPER ; PLANDEROVA , p. 40, Plate 11, Fig. 6. 1992 Puncatisporites sp. cf. P. major (COUPER ) DORHOFER ; BRENNER & Bickoff, p. 164, Plate 5, Fig. 11. 1993 Todisporites major COUPER ; MANDAOKAR , p. 134-135, Plate 2, Fig. 11, 14, 18. 1995 Punctatosporites COUPER ; RAVN , p. 64, Plate 8, Fig. 21. 42

1996 Todisporites major COUPER ; PRAMPARO et al., p. 398. Natural affinity: Filicopsida; Osmundaceae?.

Todisporites minor COUPER 1958 p. 135 Plate 16, Fig. 9 -10

See 1955 Laevigatosporites globosus LESCHIK , p. 11, Plate 1, Fig. 5. 1958 Todisporites minor COUPER , p. 135, Plate 16, Fig. 9, 10. 1968 Todisporites sp. A.; JAIN , p. 9, Plate 1, Fig. 5. 1969 Todisporites minor COUPER ; VAGVOLGYI & H ILLS , p. 160, Plate 1, Fig. 13. 1973 Todisporites cf. T. minor COUPER ; STONE , p. 64, Plate 10, Fig. 51. 1974 Todisporites minor COUPER ; RAMANUJAM & S RISAILAM , p. 73, 75, Plate 2, Fig. 11. 1975 Todisporites cf. minor COUPER ; VOLKHEIMER & QUATTROCCHIO , p. 204-205, Plate 1, Fig. 2. 1976 Todisporites minor COUPER ; ROGALSKA , Plate 10, Fig. 153-156. 1977 Punctatisporites globosus (LESCHIK ) LUND , p. 52, Plate 1, Fig. 11. 1977 Punctatisporites major (COUPER ) DORHOFER ,p. 20-21 (in part). 1977 Todisporites minor COUPER ; STAPLETON & B EER , pp. 6-7, Plate 2, Fig. 9. 1985 Todisporites rotundiformis (MALJAVKINA ) POCOCK ; Yu et al., p. 65-66, Plate 12, Fig. 22. 1986 Todisporites minor COUPER ; ASHRAF & E RBEN , p. 128, Plate 2, Fig. 10. 1986 Punctatisporites minor (COUPER ) BRENNER , p. 143, Fig. 6.10. 1086 Todisporites minor COUPER ; Nanjing Institute etc., p. 200, Plate 1, Fig. 18, 39, 40, 46. 1991 Punctatisporites globosus (LESCHIK ) LUND ; DYBKJAER , p. 19, Plate 1, Fig. 8. 1991 Punctatisporites minor COUPER ; ZHANG & ZHAN , p. 76, Plate 3, Fig. 18. 1992 Todisporites cf. T. minor COUPER ; KUMAR , p. 84, Plate 1, Fig. 8. 1992 Todisporites minor COUPER ; ELA & M AHROUS , p. 607, Fig. 5.1. 1998 Todisporites cf. minor COUPER ; SONG , p. 344, Plate 1, Fig. 23, 24. 1998 Todisporites minor COUPER ; SHANG , p. 444, Plate 3, Fig. 2. Natural affinity: Filicopsida; ?Osmundaceae.

Genus: Uvaesporites DOERING 1965 P. 39

Uvaesporites argenteaeformis (BOLKOVITINA 1953) SCHULZ 1967

1953 Stenozonatriletes argenteaeformis BOLKOVITINA , p. 51, Plate 7, Fig. 9. 1962 Triletes reissingeri RHEINHARDT , p. 707, plate 2 fig. 1-2 1967 Uvaesporites argenteaeformis (BOLKOVITINA ) SCHULZ , p. 560, Plate 2, Fig. 10, 11. 1997 Uvaesporites argenteaeformis (BOLKOVITINA ) SCHULZ ; ZHANG & GRANT -MACKIE , p. 18, Plate 2, Fig. 7; Plate 4, Fig. 11, 12. 2000 Uvaesporites argenteaeformis (BOLKOVITINA ) SCHULZ ; GAO et al., p. 222, Plate 8, Fig. 16. 2001 Uvaesporites argenteaeformis (BOLKOVITINA ) SCHULZ ; VAJDA , pp. 417, 421, Fig. 10I, 15H. Natural affinity: Lycopsida.

Sculptured trilete Spores

Genus: Acanthotriletes (Naumova 1939) p. 355 POTONIÉ & K REMP 1954

Acanthotriletes varius NILSSON 1958 page 42 Plate 2, Fig. 10

1958 Acanthotriletes varius sp. nov. – NILSSON , p. 42, Plate 2, Fig. 10. 43

1964 Anemiidites spinosus sp. nov. – MÄDLER (b), p. 180, Plate 2, Fig. 11.

Genus: Baculatisporites PFLUG & T HOMSON 1953 p. 56

Baculatisporites sp. 1934 Baculatisporltes primarius WOLFF 1953 PFLUG & THOMSON in THOMSON & PFLUG , 1953, p. 56.

Comment: Krutzsch 1967 after having studied spores of all recent species of Osmunda , proposes to maintain Verrucosisporites for Paleozoic spores of this affinity and morphology, and to broaden the diagnosis of Baculatisporites so it will encompass all Mesozoic spores with osmundoid character of ornamentation, which includes ruguiate and baculate sculpture. Similar sculpture can be found, amongst others, in the genera Todea and Lepidopteris of the Osmundaceae . Krutzsch states that at present only few fossil spores can be identified positively with extant Osmunda species.

Genus: Camerosporites LESCHIK 1956 p. 40

Camerosporites secatus (LESCHIK 1956)

1956 Camerosporites secatus sp. nov. – LESCHIK p. 40 plate 5 fig 11 – 13 1970 Camerosporites pseudoverrucatus - SCHEURING P . 87 – 88, PLATE 29 fig 253 – 267, pl. 30, fig 268 – 281, plate 31 fig. 282 – 292 Comment: According to SCHEURING 1980 C. pseudoverrucatus is a junior synonym of C. secatus .

Genus: Carnisporites MÄDLER 1964 p. 74

Carnisporites ornatus (MÄDLER 1964)

1964 Carnisporites ornatus sp. nov. – MÄDLER . p. 96, plate 8, fig. 10

Carnisporites spininger (LESCHIK 1955) MORBEY 1975

1955 Apiculatisporites spiniger sp. nov. – LESCHIK , p. 18, Plate 2, Fig. 6, 7. 1958 Sporites telephorus Pautsch, p. 323, plate 1, fig.12 1960 Anapiculatisporites telephorus (Pautsch) Klaus, p 124, plate 29, fig. 17 1962 Anapiculatisporites spiniger (Leschik) Reinhardt, p. 707, plate 1, fig. 1 1964 Carnisporites telephorus (Pautsch) Maedler, p. 95, plate 8 fig. 9 1975 Carnisporites spiniger (LESCHIK 1955) comb. nov. – MORBEY , p. 12, Plate 1, Fig. 10-12.

Genus: Conbaculatisporites KLAUS 1960 p. 125

Conbaculatisporites mesozoicus KLAUS 1960

1960 Conbaculatisporites mesozoicus , KLAUS , p. 126, plate 29, fig. 15

Genus: Converrucosisporites POTONIE & K REMP 1954

Converrucosisporites luebbenensis SCHULZ 1967

1967 Converrucosisporites luebbenensis sp. nov. – SCHULZ 1967, p. 561, Plate 2, Fig. 15-17, Plate 25, Fig. 1.

Genus: Kraeuselisporites LESCHIK 1956 p. 36

Kraeuselisporites sp. Plate 2, Fig. b

Genus: Lycopodiacidites COUPER 1953 p. 26 emend. POTONIE 1956

Lycopodiacidites rugulatus (COUPER 1958) SCHULZ 1967

1958 Perotrilites rugulatus sp. nov. – COUPER , p. 147, Plate 25, Fig. 7, 8. 1967 Lycopodiacidites rugulatus comb. nov. – (COUPER ) SCHULZ , p. 573 - 574, Plate 7, Fig. 15, 16.

Genus: Nevesisporites DE JERSEY & P ATEN 1964

Nevesisporites lubricus ORLOWSKA -ZWOLINSKA 1972

1972 Nevesisporites lubricus sp. nov. – OLOWSKA -ZWOLINSKA , p. 309, Plate 6, Fig. 41, 42.

Genus: Polypodiisporites POTONIE 1931

Polypodiisporites polymicrofeoratus (ORLOWSKA -ZWOLINSKA 1966) LUND 1977

1966 Foveosporites polymicroforatus ORLOOWSKA -ZWOLINSKA p. 1011 plate 2 fig 15 - 16 1977 Polypodiisporites polymicroferratus (ORLOWSKA - Z WOLINSKA ,) L UND , P. 58, PLATE 3 FIG 6

Genus: Porcellispora SCHEURING 1970 p. 103 emend. MORBEY 1975

Porcellispora longdonensis (Clarke 1965) SCHEURING 1970 EMEND . M ORBEY 1975

1965 Conbaculatisporites longdonensis sp. nov. – CLARKE , p. 299, Plate 36, Fig. 1-5. 1970 Porcellispora longdonensis (CLARKE 1965) comb. nov. – SCHEURING , p. 103 - 104, Plate 37, Plate 38 Plate 39 Fig. 408 - 409. 1975 Porcellispora longdonensis (SCHEURING 1970) emend. – MORBEY , p. 23 - 24, Plate 6-8, Fig. 1-7.

Genus: Punctatisporites IBRAHIM 1933 P. 21

Punctatisporites sp. Plate 2, Fig. d 45

Genus: Semiretisporis REINHARDT 196 P. 709

Semiretisporis gothae REINHARDT 1962

1962 Semiretisporis gothae sp. nov. – REINHARDT , p. 709, Plate 2, Fig. 5 - 6.

Genus: Trachysporites NILSSON 1958 P.38

Trachysporites fuscus NILSSON 1958 Plate 1, Fig. a

1958 Trachysporites fuscus sp. nov. – NILSSON , p. 38, Plate 2, Fig. 1. 1975 Microreticulatisporites fuscus (NILSSON 1958) n. comb. – MORBEY , p. 17, Plate 4, Fig. 1-4.

Genus: Taurocusporites STOVER 1962 p. 55 emend. PLAYFORD & D ETTMANN 1965 p. 146

Taurucosporites sp. A MORBEY 1975

1975 Taurucosporites sp. A sp. nov. – MORBEY , p. 19, Plate 4, Fig. 5-8.

Genus: Tigrisporites KLAUS 1960 p. 140

Tigrisporites sp.

Genus: Verrucosisporites IBRAHIM 1933 P.24

Verrucosisporites sp. sensu ZHANG & G RANT -MACKIE 1997 Plate 2, Fig. e, f

1997 Verrucosisporites sp.; ZHANG & GRANT -MACKIE , p. 15. 2001 Verrucosisporites spp.; ZHANG & GRANT -MACKIE , p. 602, Fig. 11H, 11I. Natural affinity: Filicopsida.

Genus: Zebrasporites Klaus 1960 p. 37

Zebrasporites fimbriatus KLAUS 1960

1960 Zebrasporites fibriatus .; KLAUS P 39 PLATE 30 FIG 29.

Monolete Spores

Genus: Aratrisporites (LESCHIK 1956) p. 38 emend. MÄDLER 1964

1955 Aratrisporites gen. nov. – LESCHIK , p. 38. 1960 Saturnisporites gen. nov. – KLAUS , p. 142. 1964 Aratrisporites (LESCHIK 1955) emend. – MÄDLER (a), p. 79

Aratrisporites sp. sensu JEANS et al. 2003

2003 Aratrisporites sp. nov. – JEANS et al., p. 72, 77, Fig. 14L. Natural affinity: Lycopsida.

Genus: Gibeosporites (LESCHIK 1959) p. 59

Gibeosporites gemini sensu BONA

Comment: The described monolete spores are spores of the genus Gibeosporites , but the official name of these spores is Gibeosporites lativerrucosus (LESCHIK 1959).

Genus: Simplicesporites LESCHIK 1956 p. 34

Simplicesporites sp. sensu BONA

Comment: The described spores are very poorly preserved.

Pollen Circumpolles

Genus: Corollina MALJAVKINA 1949 p.124 EMEND . C ORNET & T RAVERSE 1975

1949 Corollina MALJAVKINA . 1953 Classopollis gen. nov. – PFLUG , p. 91. 1958 Classopollis (PFLUG ) emend. – COUPER 1958, p. 156. 1960 Circulina MALJAVKINA ex. KLAUS , p. 165. 1961 Classopollis (PFLUG 1953) emend. – POCOCK & J ANSONIUS , p. 443. 1964 Corollina (MALJAVKINA 1949) emend. – VENKATACHALA & G OZCAN , p. 215. 1966 Gliscopollis gen. nov. – VENKATACHALA , p. 99. 1975 Corollina (MALJAVKINA 1949) emend. – CORNET & T RAVERSE 1975, p. 16.

Corollina meyeriana (KLAUS 1960)VENKATACHALA & G ÓZCÁN 1964 Plate 3, Fig. f; Plate 4, Fig. a

1960 Circulina meyeriana sp. nov. – KLAUS , p. 165, Plate 36, Fig. 57-60. 1964 Corollina meyeriana (KLAUS 1960) comb. nov. – VENKATACHALA & G ÓZCÁN , p. 219, Plate 3, Fig.1, 5 ,21 1966 Gliscopollis meyeriana (KLAUS 1960) emend. – VENKATACHALA & G ÓZCÁN , p. 99. 1968 Classopollis reclusus (THIERGART 1949) comb. nov. – MÄDLER , p. 306, Plate 30, Fig. 5-9. 1973 Classopollis meyeriana (KLAUS ) comb. nov. –DE JERSEY , p. 130 – 131 plate 3 fig 5 – 10, plate 4 fig 4 -6 47

Corollina torosa (REISSINGER 1950) CORNET & TRAVERSE 1975

1950 Pollenites torosus sp. nov. – REISSINGER , p. 115, Plate 14, Fig. 20. 1953 Classopollis classoides sp. nov. – PFLUG , p . 91, Fig. 4 j-m, Plate 16, Fig. 20-25, 29-37. 1958 Classopollis torosus (REISSINGER 1950) comb. nov. – COUPER , p. 156, Plate 28, Fig. 2-7. 1961 Classopollis classoides (PFLUG 1953) emend. – POCOCK & J ANSONIUS , p. 443, Plate 1, Fig. 1-9. 1975 Corollina torosa (MALJAVKINA 1949) KLAUS 1960 emend. – CORNET & T RAVERSE , p. 17, Plate 5,

Genus: Geopollis BRENNER 1986

Geopollis zwolinskae (LUND 1977) BRENNER 1986

1972 Circulina sp. nov. – FISHER 1972, p. 105, Plate 8, Fig. 22 1975 Gliscopollis meyeriana (KLAUS 1960) VENKATACHALA 1966 in MORBEY 1975: only Plate 12, Fig. 10. 1975 Granuloperculatipollis cf. rudis (VENKATACHALA & G ÓZCÁN 1964) emend. – MORBEY , p. 38, Plate 12, F Fig. 14-17. 1977 Corollina sp. – SCHUURMAN p.215, plate 22, fig.8 1977 Corollina zwolinskae n. sp. – LUND , p. 70, Plate 7, Fig. 5 a, b. 1987 Geopollis zwolinskae (LUND 1977) comb. nov. – BRENNER 1986, p. 158, Plate 8, Fig. 12.

Genus: Granuloperculatipollis VENKATACHALA & G ÓZCÁN 1964 p. 219

Granuloperculatipollis rudis VENKATACHALA & G ÓZCÁN 1964 emend. Morbey 1975 p. 35

1964 Granuloperculatipollis rudis sp. nov. – VENKATACHALA & G ÓZCÁN 1964, p . 219 - 220, Plate 3, Fig. 22- 29.

Genus: Paracirculina KLAUS 1960 P.162 Paracirculina quadruplicis SCHEURING 1970

1970 Paracirculina quadruplicis SCHEURINGP . 94 – 95 PLATE 35 FIG 358 - 363.

Various pollen

Genus : Callialasporites DEV 1961 P. 48 Callialasporites dampieri DEV , 1961 Plate 1, Fig. e

1937 Nelumbium type SIMPSON , p. 673, fig 2a 1957 Zonalapollenites dampieri BALME p. 32, pl. 8, Figs. 88, 90. 1958 Zonalapollenites dampieri BALME ; L ANTZ p. 925, pl. 3, Figs. 34, 35; pl. 4, fig. 36. 1958 Zonalapollenites cf. Z. dampieri BALME ; H UGHES & C OUPER p. 1482, Figs. 1c, 1d. 1959 Zonalapollenites dampieri BALME ; DE JERSEY p. 362, pl. 3, Fig. 8. 1961 Callialasporites dampieri (BALME ) D EV p. 48, pl. 4, Figs 26, 27. 1962 Pflugipollenites dampieri (BALME ) – P OCOCK p 72 plate 12 fig 183 – 184 1962 Appalanopsis dampieri (BALME ) D OERING p.113, plate 16, fig 11 – 15 1962 Appalanopsis lenticularis (B ALME ) D OERING p.113, plate 16, fig 9 -10 1963 Tsugaepollenites dampieri (B ALME ) L EVET -CARETTE p. 107 plate 6 figs 13-17

Genus : Eucommiidites ERDTMAN , 1948 emmend. P OTONIE , 1958

Eucommiidites troedssonii (E RDTMANN 1948) P OTONIE , 1958

1948 Tricolpites (Eucommiidites) troedssonii ERDTMANN P . 267, figs 5 –10, 13 – 15 1953 Protoquercus agdijakendensis BOLKHOVITINA , p. 93, plate 15 figs 29 - 32 1958 Eucommiidites troedssonii POTONIE p. 87, plate 10, fig 117 1960 Eucomiidites minor GROOT – P ENNY , p.234, plate 2, fig 14 1961 Psilatricolpites psilatus PIERCE , p. 49, plate 3, figs 98, 99.

Comment: Potonié, 1958 first used the name Eucommiidites as a single, definite form genus, interpreting the designation ' Tricolpites ', placed before it by Erdtman, as a suprageneric morphographic grouping. Potonié compares the form with Schopfipollenites and Bennettiteaepollenites , and groups the genus with the Praecolpates rather than the tricolpate angiospermous grains. He changes the generic diagnosis thus: Shape ovaloid, exine smooth or infrapunctate; the form is probably praecolpate rather than "tricolpate" as described by Erdtman, as the three colpate features are rather different in length than equal.

Genus: Inaperturopollenites PFLUG & T HOMSON IN THOMSON & P LUG 1953 P. 64

Inaperturopollenites reissingerii sensu BONA

Comment: Bóna’s Inaperturopollenites reissingerii is very similar to specimens of Inaperturopollenites dubius (POTONIE & V ENITZ 1934) T HOMSON & P FLUG 1953.

Genus: Monosulcites COOKSON 1947 EX . C OUPER 1953

Monosulcites minimus COOKSON 1947 p. 135 plate 15 fig 47 - 50 1947 Monosulcites minimus COOKSON p 135 plate 15 fig 47 - 50

Genus: Rhaetipollis SCHULZ 1967 emend. SCHUURMAN 1977

1967 Rhaetipollis gen. nov. – SCHULZ , p. 605. 1977 Rhaetipollis (SCHULZ 1967) emend. – SCHUURMAN , p. 217.

Rhaetipollis germanicus SCHULZ 1967 emend. SCHUURMAN 1977

1967 Rhaetipollis germanicus sp. nov. – SCHULZ ,p. 605 – 606 Plate 22, Fig. 10-15., plate 26 fig 4 1977 Rhaetipollis germanicus SCHULZ 1967 emend. – SCHUURMAN 1977, p. 217.

Genus: Ricciisporites LUNDBLAD 1954 p. 400

Ricciisporites tuberculatus LUNDBLAD 1954 Plate 3, Fig. b

1954 Ricciisporites tuberculatus . – LUNDBLAD P . 401, PLATE 4 FIG . 8.

Genus: Tsugaepollenites POTONIE & V ENIZ 1934 EX . P OTONIE 1958

Tsugaepollenites macroverrucosus sensu BONA

Comment: The described trisaccate pollen grains are probably similar to Tsugaepollenites pseudomassulae (MÄDLER 1964) Morbey 1975.

Bisaccate Pollen

Genus: Alisporites DAUGHTERY 1941 p. 98

Alisporites sp. sensu JANSONIUS 1971

1971 Alisporites sp. – JANSONIUS , Pollen et Spores, Vol. 3, p. 355.

Comment: After a discussion of earlier emendations and a re-examination of the holotype, Jansonius proposed the following new diagnosis: Bisaccate, sulcate pollen grains; equatorial outline oval to broadly oval; cappa not strongly thickened, but clearly differentiated from the sacci; sacci proximally attached in equatorial region, but distally more or less strongly overlapping the central part; distal bases of sacci not clearly marked, enclosing a cappula that reaches towards the equator; on distal side a more or less narrow nexinal sulcus is reflected by nexinal folds parallel to and approximately underlying, or even involving, the distal bases of the sacci; a more or less distinct nexinal body is present; sacci usually not strongly distally pendent, with fine reticulation.

Alisporites minutisaccus CLARKE 1965

1965 Alisporites minutisaccus Clarke P. 310 311 plate 35 fig. 12

Alisporites robustus NILSSON 1958 P. 82

1958 Alisporites robustus NILSSON P 82 PLATE 8 FIG 2-3

Disturbopollenites sensu BONA

Comment: Disturbopollenites is not an official pollen genus. The described material is bisaccate and looks similar to pollen of the genus Alisporites .

Genus : Chasmatosporites NILSSON 1958 P. 51 - 53

Chasmatosporites apertus NILSSON 1958

1954 Pollenites apertus – ROGALSKA , p45 plate 12 fig 13 – 1958 Chasmatosporites apertus (ROGALSKA ) N ILSSON 1958, p.56 Plate 2 fig. 5 – 6 1962 Verrucipollenites apertus (ROGALSKA ) B ONA p. 23 plate 2 fig 2 1958 Chasmatosporites crassus NILSSON p. 57 plate5 fig 3

Chasmatosporites minimus sensu BONA

Comment: Chasmatosporites minimus is not officially introduced into the scientific literature. The described pollen grains are relatively small monosulcate gymnosperm pollen. The morphology is relatively similar to Chasmatosporites canadensis (POCOCK 1970). 50

Genus: Cycadopites (WODEHOUSE 1933) ex. WILSON & WEBSTER 1946

Cycadopites spp.

1933 Cycadopites – WODEHOUSE p. 482.

Comment: Wodehouse did not assign any species to the original describtion 1933.

Cycadopites nitidus (B ALME 1957) DE JERSEY & H AMILTON 1967

1957 Entylissa nitidus BALME , p. 30, Plate 6, Fig. 78-80. 1962 Ginkgocycadophytus nitidus (BALME ) DE JERSEY , p. 12, Plate 5, Fig. 1-3. 1964 Cycadopites nitidus De Jersey p 10 . 1969 Cycadopites nitidus (B ALME ) N ORRIS p599 – 6—plate 111 figs 11 – 12

Genus: Ginkgocycadophytus SAMOILOVICH 1953 P.30

Ginkgocycadophytus sp. sensu BONA

Comment: Based on the image material of the internal report by Bóna, Ginkgocycadophytus spp. is probably the same as Cycadopites nitidus (BALME ) DE JERSEY 1964

Genus: Lunatisporites LESCHIK 1956 p. 56 emend. SCHEURING 1970

Lunatisporites sp. sensu JEANS et al. 2003

2003 Lunatisporites sp; JEANS et al., pp. 73, 77, Fig. 14N. Natural affinity: Gymnospermopsida.

Lunatisporites rhaeticus ( SCHULZ 1967 WARRINGTON 1974 Plate 3, Fig. c

1967 Lunatisporites rhaeticus SCHULZ , p. 30. 1974 Lunatisporites rhaeticus WARRINGTON .

Genus: Ovalipollis KRUTZSCH 1955 emend. SCHEURING 1970

Ovalipollis minimus ( SCHEURING 1970 P. 34)

1970 Ovalipollis minimus sp. nov. – SCHEURING plate 2 Fig. 6 – 7, Plate 4 Fig. 21, 22.

Ovalipollis ovalis (KRUTZSCH 1955) SCHEURING 1970

1955 Ovalipollis longiformis sp. nov. – KRUTZSCH , p. 70, Plate 1, Fig. 1-5. 1954 Ovalipollis breviformis sp. nov. – KRUTZSCH , p. 70, Plate 6. 51

1956 Unatexisporites mohri LESCHIK p. 60 plate 8 fig 9 1960 Ovalipollis lunzensis sp. nov. – KLAUS , p. 152, Plate 34, Fig. 46-49. 1960 Ovalipollis rarus sp. nov. – KLAUS , p. 154, Plate 35, Fig. 50. 1970 Ovalipollis ovalis (KRUTZSCH 1955) emend. – SCHEURING , p . 31, Plate 1, Fig. 1; Plate 2, Fig. 3; Plate 3, Fig. 16.

Genus: Perinopollenites COUPER 1958

Perinopollenites elatoides COUPER 1958 p.152 Plate 3, Fig. a

1958 Perinopollenites elatoides COUPER , p. 152, Plate 27, Fig. 9-11. Natural affinity: Gymnospermopsida.

Ballosporites hians sensu BONA

1964 Ballosporites hians MÄDLER , p. 179, Plate 2, Fig. 6-10 1965 Perinopollenites elatoides COUPER ; NORRIS , p. 259, Fig. 4c, 63.

Comment: The material classified as Ballosporites hians does not look exactely like the material from Mädler (1964) and therefore it is not similar to Perinopollenites elatoides . However, the material is too poor preserved to be described as new species and therefore, the name Ballosporites hians is used in this study.

Genus: Pinuspollenites RAATZ 1938 EX POTONIE 1958

Pinuspollenites minimus (COUPER 1958) KEMP 1970

1958 Abietinaeepollenites minimus sp. nov. COUPER P .153, PL .28 FIG 14 - 15. 1966 Pinus minima (C OUPER ) A NDERSON p. 17 plate 10, fig 5. 1970 Pinuspollenites minimus (COUPER ) K EMP p. 116, plate 24, figs 1 – 6.

Genus: Platysaccus NAUMOVA 1939 ex Ishchenko 1952

Platysaccus sp.

Genus: Quadraeculina (M ALIAVKINA 1949)

Quadraeculina bitorosus sensu BONA

Comment: The described material is probably identical with Quadraeculina anaellaeformis MALIAVKINA (1949).

Genus: Schizosaccus MÄDLER 1964 P. 119

Schizosaccus keuperi MÄDLER 1964

1964 Schizosaccus keuperi sp. nov. – MAEDLER P . 119 PLATE 11, FIG 9-12. Natural affinity: Gymnospermopsida. 52

Genus: Triadispora KLAUS 1964 p. 120

Triadispora sp. sensu JEANS et al. 2003

2003 Triadispora sp. . – JEANS et al., p. 74. Natural affinity: Gymnospermopsida.

Genus: Vitreisporites LESCHIK 1956 p. 1953 emend. JANSONIUS 1962

Vitreisporites pallidus (REISSINGER 1938) NILSSON 1958

1938 Pityosporites pallidus sp. nov. – REISSINGER 1938, p. 14 1950 Pityopollenites pallidus comb. nov. – REISSINGER , p. 109, Plate 15, Fig. 1-5. 1958 Caytonipollenites pallidus (REISSINGER 1950) comb. nov. – COUPER , p. 150, Plate 26, Fig. 7,8. 1958 Vitreisporites pallidus (R EISSINGER ) N ILSSON p 77 - 78

5.2.2 Aquatic Palynomorphs

Dinoflagellates

Subclass: Peridiniphycidae FENSOME et al. 1993 Order: Gonyaulacales TAYLER 1980 Suborder: Rhaetogonyaulacineae NORRIS 1978 Family: Shublikodiniaceae WIGGINS 1973

Genus: Dapcodinium EVITT 1961, emend. BELOW 1987

Dapcodinium priscum EVITT 1961, emend. BELOW 1987 Plate 2, Fig. c

1961 Dapcodinium priscum n. gen. n. sp. – EVITT 1961, p. 996, Plate 119, Fig. 1-14. 1987 Dapcodinium priscum EVITT 1961 emend. – BELOW 1987, Plate 23, Fig. 6-11, 19.

Genus: Rhaetogonyaulax SARJEANT 1966 p. 152 153, emend. BELOW 1987

Rhaetogonyaulax rhaetica (SARJEANT ) LOEBLICH & L OEBLICH (1968) – emend. BELOW 1987 Plate 3, Fig. d

1963 Gonyaulax rhaetica – SARJEANT , p. 353, Fig. 1, 2. 1963 Gonyaulax chaloneri SARJEANT p. 354, Fig 2. 1966 Rhaetogonyaulax chaloneri SARJEANT nom. nud. SEARGEANT p. 153 1968 Rhaetogonyaulax rhaetica (SARJEANT 1963) comb. nov. – LOEBLICH & L OEBLICH 1968, p. 213.

Acritarchs

Group: Acritarcha (EVITT ) DIVER & P EAT 1979 Subgroup: Acanthomorphitae DOWNIE , E VITT & S ARJEANT 1963

Genus: Micrhystridium DEFLANDRE 1937, Micrhystridium spp.

Subgroup: Herkomorphitae DOWNIE , E VITT & S ARJEANT 1963

Genus: Cymatiosphaera O. WETZEL 1933, ex DEFLANDRE 1954

Cymatiosphaera sp.

Subgroup: Pteromorphitae DOWNIE , E VITT & S ARJEANT 1963

Genus: Pterosper mopsis W. WETZEL 1952 p. 411

Pterospermopsis sp. Plate 4, Fig. e

Tasmanites

Division: Chlorophyta PASCHER 1914 Class: Chlorophycea KÜTZING 1843 Family: Tasmannaceae SOMMER 1956

Genus: Tasmanites NEWTON 1875 p. 341

Tasmanites spp. Plate 4, Fig. b

Division: Chlorophyta PASCHER 1914 Class: Chlorophycea KÜTZING 1843 Schizosporis COOKSON & D ETTMANN 1959

Schizosporis sp.

6 Biostratigraphy 6.1 Biostratigraphy of the Triassic/Jurassic boundary interval

In Central Europe numerous tectonostratigraphic units represent a broad spectrum of facies types across the Triassic/Jurassic boundary. Here, the main focus is given on the biostratigraphy of the most complete sections. The bivalve Rhaetavicula contorta is the key biostratigraphic marker of the topmost Rhaetian, developed in marly facies (“Grès

infraliasique”) that overlies the marginal marine Keuper. In the eastern part of the Paris Basin and around the Ardennes Massif the basal Jurassic is represented by a Gryphaea -bearing limestone of early Hettangian age as indicated by ammonoids of the genus Psiloceras . It is of historical interest that the Hettangian Stage was first described on the basis of a sequence at Hettange in Lorraine (Pálfy 2008). In the Western Carpathian Tatra Mountains, biostratigraphically important Late Triassic microfossilsare rare (Ga ździcki 1974, 1978; Ga ździcki et al. 1979; Ga ździcki & Michalík 1980; Błaszyk & Ga ździcki 1982; Ga ździcki 1983; Michalík & Ga ździcki 1983; Fijałkowska & Uchman 1993; Ga ździcki et al. 2000). Rapid evolutionary changes of Involutinidae, Ammodiscidae and Ophthalmidiinae permit the establishment of a detailed foraminiferal zonation. The sequence of the Rhaetian Glomospirella friedli -Triasina hantkeni assemblage zone and the Hettangian -Sinemurian Ophthalmidium leischneri -Ophthalmidium walfordi assemblage zone was detected by Ga ździcki (1978). The Glomospirella friedli -Triasina hantkeni zone was correlated with both the Choristoceras haueri and Ch. marshi ammonoid zones (Rhaetian), and its extent also corresponds to that of the Misikella posthernsteini conodont zone. The extent of the Early Jurassic Ophthalmidium leischneri -Ophthalmidium walfordi zone may correspond to the Planorbis to Angulata, and possibly also the Bucklandi standard ammonite zones of the Hettangian -Sinemurian, defining the age of the basal Jurassic strata in the Tatra Mountains (Michalík et al. 2007). In the Northern Hungarian Cs ővár section the biostratigraphic definition of the Triassic/Jurassic boundary is based on ammonoids, radiolarians, foraminifera and conodonts (Pálfy et al. 2007). Ammonoids are scarce and the poor preservation hinders firm identification. Choristoceras findings in the lowermost part of the section indicate the late Rhaetian marshii zone. The Triassic/Jurassic boundary interval yields findings of ex situ phylloceratids (Pálfy et al. 2007). A pronounced radiolarian turnover from mainly multicyrtid nasselarians to saturnalid spumellarians is observed (Pálfy et al. 2007). Radiolarian assemblages of the Globolaxtorum tozeri zone indicate an Upper Rhaetian age. Lower Hettangium limestone beds comprise radiolarians of the Canoptum merum zone. Foraminifers have been investigated by Pálfy et al. (2007). In the lower part of the section platform derived forms are rare and the assemblage is dominated by agglutinated forms. The Rhaetian age of this interval is suggested by the

occurrence of Variostoma cochlea , V. coniforme , V. crassum , Ophthalmidium leischneri , Miliopora cuvillieri , Galeanella panticae , Ammobaculites rhaeticus and A. eomorphus (Pálfy et al. 2007). Upsection, much impoverished assemblages were encountered. Few persisting Triassic species include Aulotortus tenuis and A. tumidus. An oligospecific association dominated by encrusting P. carinata occurs, except for facies-controlled assemblages of abundant smooth, elongated forms ( Eoguttulina, Ramulina and Dentalina ) in laminated beds. The latter assemblage, which is interpreted to indicate relatively deeper (neritic to bathyal) environments, re-occurs in lesser abundance in the upper part of the section. Involutina liassica , a diagnostic Jurassic species, first appears near the top of the section. The conodont association of the lowermost part of the Cs ővár section corresponds to the lower part of the Misikella posthernsteini Assemblage Zone of Kozur & Mock (1991), including specimens of Norigondolella steinbergensis and representatives of two Misikella species, M. hernsteini and M. posthernsteini as well as elements of Norigondolella (Orchard

2005), Chirodella dinodoides and Neohindeodella rhaetica. Stratigraphically higher collections are assigned to the Misikella ultima zone of Kozur & Mock (1991) based on the appearance of the zonal nominal species, although it is subordinate to M. posthernsteini throughout this interval. Single specimens of M. ultima and Neohindeodella sp. were recovered in the younger, here assumed as Hettangian part of the section. The latter record may support the suggestion of Kozur (1993) that at least one conodont lineage survived into the earliest Jurassic (Pálfy et al. 2007).

Biostratigraphic data of the S Hungarian Mecsek Mountains are nearly absent. In the

Hettangian/Sinemurian deposits of the Komló area, footprint findings of the dinosaur

Komlosaurus carbonis are more or less the only published palaeontological data of this region

(Kordos 1983). In the upper part of the Pécs coal pit (Jurassic sediments) marine bivalves are present, but up to now these fossils are not described systematically. Palaeobotanical findings of the Jurassic pteridosperm Komlopteris nordenskioeldii were described by Barbacka (1994).

6.2 Current palynostratigraphic zonations

Latest Triassic and Early Jurassic palynological assemblages are well documented by a number of studies in the German and Danish Triassic basins (e.g., Schulz 1967, Herngreen &

De Boer 1974, Lund 1977, Guy-Ohlson 1981, Brenner 1986, Lund 2003) and the British

Rhaetian-Hettangian (e.g., Orbell 1973, Warrington 1974, Hounslow et al. 2004). A compilation is given in Appendix 1.

Lund (1977) divided the Rhaetian of the North Sea into a Rhaetipollis Limbosporites zone and a Ricciisporites Polypodiisporites zone; the Hettangian is build up by the Pinuspollenites

Trachysporites zone. Orbell (1973) has distinguished a Late Triassic Rhaetipollis zone and an

Early Hettangian Heliosporites zone in the Austrian Kössen Beds. The latter is characterized by an acme of Naiaditaspora spp. ( Naiaditaspora harrisii is considered by Morbey (1975) as a junior synonym of Porcellispora longdonensis ) following the rapid decline of palynomorphs characterizing the Rhaetipollis zone ( Rhaetipollis germanicus and Ovalipollis pseudoalatus ) and a marked increase in the abundance of Heliosporites .

Kürschner et al. (2007) separates a Rhaetipollis-Porcellispora zone and a Trachysporites

Porcellispora zone within the Rhaetian of the Northern Calcareous Alps. The Hettangian is defined by palynomorphs of the Trachysporites-Heliosporites Zone. Kürschner et al. (2007) suggested Cerebropollenites thiergartii as a marker species of the Hettangian. Weiss (1989) divided the Rhaetoliassic of S Germany into a Rhaetian Concavisporites-Duplexisporites problematicus-Ricciisporites tuberculatus zone and a Hettangian Concavisporites-

Duplexisporites problematicus zone. Brenner (1986) described sporomorph assemblages of the SW German realm, but forbeard from defining zones. Sediments of the Polish Basin were investigated by Orlowska-Zwolinska in 1983. She distinguished a Rhaetian Assemblage

(Assemblage V) and a Hettangian Assemblage (Assemblage VI). Ashraf et al. (1999) studied the Rhaetian Haojiagou Formation and the Liassic Badaowan Formation of the Chinese 57

Junggar Basin. Palynofloras of Eastern North America are the only investigated assemblages in the world that are affected by a mass extinction. The sudden decrease in sporomorph diversity at the Triassic/Jurassic boundary is described by Fowell & Olsen (1993) from the

Newark Basin.

6.3 Microfloral zonation of the studied Rhaetoliassic material

Due to the poor preservation of the palynomorph assemblages in most of the sections studied, a definition of zones was not possible. In the Furkaska section of the Slovakian Tatra

Mountains palynomorphs are well preserved and show characteristic changes within the boundary interval. The sudden increase in the abundance of trilete spores, the last appearance of Corollina spp., and the first appearance of Concavisporites spp. and Pinuspollenites minimus are striking features for a subdivision of two palynomorph assemblages. The differences of both assemblages can be detected by means of multivariate statistical analysis which allows the quantification of similarity/dissimilarity of a variety of samples that were examined according to different attributes.

A Principal Components Analysis (PCA) was performed for the dataset consisting of 11 samples and 70 sporomorph species (variables). The principal plane for the first two principal components is shown in Figure 28. The distances among the projected points on this plane are a measure of similarity (cf. Marinoni 2006). The smaller this distance, the higher is the similarity. The PCA results show that two data clusters can be identified: The Jurassic samples that were taken above sample horizon 408 and the Triassic samples that were sampled below horizon 408 (Fig. 28). In contrast to the Triassic samples, the Jurassic samples are characterised by a very high abundance and diversity of trilete spores.

Fig 28: PCA plot for the Furkaska section. Principle plane for components 1 and 2.

Sample 400/401 can clearly be identified as an outlier, indicating a different pattern regarding the variables analysed. Sample 400/401 is the lowest and oldest sample of the Furkaska section. Previous studies detected a change in the microfloral assemblage from Mid Rhaetian to Upper Rhaetian (Kürschner et al. 2007). The presented PCA results can therefore be regarded as multivariate evidence for this microfloral change between the Middle and Upper

Rhaetian.

Fig. 29: Hierarchical tree plot of the Furkaska data set.

A very similar clustering pattern can be identified in the Cluster analysis, which was carried out with the same data set as used with the PCA. Figure 29 shows the subdivision of the samples into two clusters. The Triassic samples of the lower part of the section form one cluster whereas the Jurassic samples are grouped into another cluster. Sample 400/401 which was identified an outlier in the PCA is part of the Triassic cluster, however its degree of similarity is fairly low (just below .4 see left axis in Fig. 29) and it is therefore separated very early from the remaining Triassic samples.

It is necessary to point out, that the number of samples must be considered very low for a multivariate analysis. Regarding the minimum size of the sample population there are no general rules except that he sample population should be “large enough and representative”

(Bortz 1999). Tabachnick & Fidell (1996) recommended that the number of samples being examined by means of PCA should be at least the same as the number of variables. Though 60

the results of both, PCA and Cluster analysis are based on a small sample population their results are interpretable and confirm each other. Due to the fact that the change within the assemblage is not isochronic with the lithofacies change from limestone to clay but some centimetres lower in the section, the microfloral change is supposed to be independent from this lithologic change.

Fig. 30: Spore spike within Rhaetoliassic sediments of the Mecsek Mountains as stratigraphic correlation tool.

The palynomorph assemblages of the Mecsek Mountains can be subdivided into two different assemblage zones, displaying a characteristic cyclic vegetation pattern of a fluvial system turning periodically to a swamp area. A biostratigraphic definition of zones to subdivide

Rhaetian and Hettangian deposits was not possible. Anyway, all investigated sections show the same characteristic increase in the abundance of trilete spores within the boundary interval

(Fig. 30). This spore spike is a striking feature for correlation of sediments deposited in different palaeoenvironments of the NW Tethyan Realm.

6.4 Correlation with established zonations

Appendix 1 shows the stratigraphical occurrence of the most important palynomorphs in the different study areas in comparison with previous works. The sporomoph assemblage of the

Tatra Mountains is very similar to the assemblages of the Polish Basin (Orlowska-Zwolinska

1983) and to the assemblages of the Austrian Kössen Beds (e.g., Kürschner et al. 2007). The close palaeogeographic relation of these areas during Rhaetian and Hettangian times is the cause for this resemblance. The North Hungarian Cs ővár section comprises sporomorphs, which show affinity with those from the Germanic realm. Due to the small number of samples with well preserved palynomorphs, a detailed comparison was not possible. Palynomorph assemblages from the South Hungarian Mecsek Mountains differ from other assemblages.

Marker species of the Germanic and Alpine realm such as Ovalipollis, Corollina and

Rhaetipollis are lacking. The kind of vegetation seems to be close related to those of the other study areas, but the plant species are often different, which points to a great distance to the other areas during the Triassic/Jurassic boundary interval. Palynomorph assemblages of North

America are very different from assemblages of the NW Tethyan realm. The high increase of

Corollina spp. at the Triassic/Jurassic boundary in North American sections is in contrast to the last appearance of this genus in the Tatra Mountains during the same time interval. 62

7 Palaeoclimate reconstruction

7.1 Palynomorphs and their botanical affinity

A realistic interpretation of Mesozoic sporomorph signals is strongly dependent on the ability to distinguish palaeocommunities by using information on the ecology of mother plants

(Abbink et al. 2001, 2004a). The mother plants of the most sporomorphs of the Mesozoic are unknown. Some pollen and spores have been found in situ together with macroplant fossils.

These findings are an important tool for reconstructing the palaeoenvironment during

Rhaetian/Hettangian times. The majority of the data are taken from Traverse (1988) and

Abbink et al. (2001, 2004a). Comparison with recent vegetation pattern with respect to climatic values like temperature and humidity are another crucial part of this paleoclimatic interpretation. The sporomorph ecogroup model of Abbink et al. (2001), generated for the

Upper Jurassic and the Lower Cretaceous of the North Sea, was applied for the Furkaska section and the Komló cores 176 and 137.

Monolete Spores

Aratrisporites

According to Helby & Martin (1965) Aratrisporites was produced as a microspore by the lycopsids Cyclostrobus and Lycostrobus . Ash (1979) found Aratrisporites in the heterosporous lycopsid cone Skilliostrobus . Spores that are similar to Aratrisporites minimus

SCHULZ are found by Grauvogel-Stamm & Duringer (1983) in the lycopsid frutification

Annalepis zeilleri FLICHE . Aratrisporites microspores are related to Banksisporites and

Nathorstisporites megaspores (Scott & Playford 1985). 63

Trilete Spores

Calamospora

Calamospora spores were identified in Triassic fossil Equisetites by Couper (1958).

Cyathidites

Couper (1958) identified Cyatidites microspores in Jurassic ferns referable to Dicksoniacea .

In the Lower Cretaceous Cyathidites was found in Coniopteris (Douglas 1973) and

Onychiopsis ferns (Sukh-Dev 1980).

Deltoidospora

Deltoidospora -like microspores were obtained from Lower Cretaceous Onchiopsis ferns by

Sukh-Dev in 1980. According to Van Kronijnenburg-Van Cittert (1989, 1993) they are related to Dicksoniacea , Cyatheacea and Dipteridaceae .

Granulatisporites

Granulatisporites -type spores were identified by Litwin (1985) in Triassic Clathropteris material.

Punctatisporites

Punctatisporites microspores were described by Grauvogel-Stamm & Grauvogel (1980) from the Triassic fern Anomopteris.

Todisporites

Todisporites microspores were obtained by by Couper (1958) from Jurassic osmundaceous ferns, and from the Triassic fern WINGATEA by Litwin (1985). In situ spores from Todites spp. are referable to Todisporites (Van Kronijnenburg-Van Cittert 1978).

Rol FernPhoto courtesy Wisconsin State Herbarand Emmet J. Judziewicz

Bisaccate Pollen

Alisporites

Several species of the genus Alisporites were found by Grauvogel-Stamm (1978) from

Willsiostrobus cones, a conifer from the early Triassic. Alisporites pollen was also found in

Triassic organs of the pteridosperm Pteruchus and the conifer Masculostrobus (Townrow

1962). Alisporites has been found on Dicroidium , a probable member of the

Corystospermaceae and from the conifer frutification Lelestrobus by Srivastava (1984).

Lunatisporites

Lunatisporites was found in a conifer cone from the late Permian by Clement-Westerhof

(1974) and in Triassic Pteruchus and Masculosrobus cones (see Alisporites ) by Townrow

(1962).

Platysaccus

Platysaccus has been found in cones associated with Dicroidium (Anderson & Anderson

1983).

Vitreisporites

Vitreisporites pallidus is known to be the dispersed pollen of Jurassic Caytonanthus

(Chaloner 1968). The bisaccate nature of Vitreisporites has always been a stumbling block in efforts to connect the Caytoniales with angiosperm anciestry (Traverse 1988).

Inaperturate Pollen

Inaperturopollenites

Inaperturopollenites limbatus BALME pollen was shown to be produced by Lower Cretaceous

Brachyphyllum cones by Gamerro (1968). Pollen attributed to the genus Inaperturopollenites was also found by Grauvogel-Stamm (1978) as immature grains in Lower Triassic Darneya conifer cones, of which the mature pollen was Triadispora. Inaperturopollenites dubius

(Potonie & Venitz 1934) THOMSON & PFLUG 1953 is related to taxodiaceous conifers (Raine et al. 2006).

Perinopollenites

Perinopollenites pollen was found by Couper (1958) in cone preparations of the Jurassic taxodiaceous conifer, Elatoides . Harris (1973) confirmed this, noting that the pollen is very variable from one cone to another.

Circumpolloid Pollen

Classopollis

Classopollis (= Corollina ) pollen was produced by the Lower Cretaceous conifer cone

Tomaxiella (Gamerro 1968). Couper (1958) identified it in male cones of Pagiophyllum and

Hirmiella . However, circumpolloiud pollen of various sorts has been obtained from male 66

cones of a variety of Mesozoic conifers, including Brachyphyllum, Hirmiella, Pagiophyllum and Masculostrobus . The primary association seems to be Hirmiella (Cheirolepidaceae) cones and Classopollis pollen (Francis 1983).

Monosulcate Pollen

Cycadopites

Cycadopites -like pollen was obtained from Jurassic Sahnia (Pentoxylaceae ), according to

Sukh-Dev (1980) and from Lepidopteris ( Peltaspermales ) by Anderson & Anderson (1983).

7.2 Taxonomy and ecology of mother plants

Division Pteridophyta

Ferns

Ferns have a popular image of growing in moist, shady woodland nooks, but the reality is far more complex. Recent ferns grow in a wide variety of habitats, ranging from remote mountain elevations to dry desert rock faces to bodies of water to open fields. Ferns in general may be thought of as largely being specialists in marginal habitats, often succeeding in places where various environmental delimiters limit the success of flowering plants. On the other hand, some ferns are among the world's most serious weed species, such as the bracken growing in the British highlands, or the mosquito fern ( Azolla ) growing in tropical lakes. There are four particular types of habitats that are often key places to find ferns: the afore-mentioned moist, shady forest cove; the sheltered rock face, especially when sheltered from the full sun; acid bogs and swamps; and tropical trees, where many species are epiphytes. Many ferns depend on associations with mycorrhizal fungi. Many ferns only grow within specific pH ranges; e.g., the climbing fern ( Lygodium ) of eastern North America will only

grow in moist, intensely acid soils, while the bulblet bladder fern ( Cystopteris bulbifera ) with overlapping range is only ever found on limestone rock.

Class Polypodiopsida (= Filicopsida)

(Polypodiacaea) Polypodiaceae is a family of polypod ferns, which includes today approximately 50 genera divided into several tribes containing around 1000 species. Nearly all are epiphytes, but some are terrestrial. Their stems range from erect to long-creeping. The fronds are entire, pinnatifid, or variously forked or pinnate. The petioles lack stipules. The scaly rhizomes are generally creeping in nature. Polypodiaceae species indicate wet climates, most commonly they exist in rain forests.

Horsetails (Equisetum) Equisetum is a genus of vascular plants that reproduce spores rather than seeds. The genus includes 15 species, commonly known as horsetails and scouring rushes. Equisetum is the only one in the family Equisetaceae, which in turn is the only family in the order Equisetales and the class Equisetopsida. This class is often placed as the sole member of the Division Equisetophyta (also called Arthrophyta in older works), though some recent molecular analyses place the genus within the ferns (Pteridophyta), related to Marattiales. Other classes and orders of Equisetophyta are known from the fossil record, where they were important members of the world flora during the Carboniferous period. The name horsetail, often used for the entire group, arose because the branched species somewhat resemble a horse's tail, the name Equisetum being from the Latin equus, "horse", and seta, "bristle". The genus is near-cosmopolitan, being absent only from Australasia and Antarctica. They are perennial plants, either herbaceous, dying back in winter (most temperate species) or evergreen (tropical species). In these plants the leaves are greatly reduced, in whorls of small, segments fused into nodal sheaths. The stems are green and photosynthetic, also distinctive in being hollow, jointed, and ridged. The spores are borne in cone-like structures (strobilus, Plate strobili) at the tips of some of the stems. In many species the cone-bearing stems are unbranched, and in some they are non-photosynthetic, produced early in spring separately from photosynthetic sterile stems. In some other species (e.g., E. palustre ) they are very similar to sterile stems, photosynthetic and with whorls of branches. Many plants in this genus prefer wet sandy soils, though some are aquatic and others adapted to wet clay soils. 68

Gymnosperms Gymnosperms (Gymnospermae) are a group of seed-bearing plants with ovules on the edge or blade of an open sporophyll, the sporophylls usually arranged in cone-like structures. The term gymnosperm derives from the Greek word gumnospermos, meaning "naked seeds" and referring to the unenclosed condition of the seeds, as when they are produced they are found naked on the scales of a cone or similar structure. Gymnosperms are heterosporous, producing microspores that develop into pollen grains and megaspores that are retained in an ovule. After fertilization (joining of the micro- and megaspore), the resulting embryo, along with other cells comprising the ovule, develops into a seed. The seed is a sporophyte resting stage. In early classification schemes, the gymnosperms "naked seed" plants were regarded as a "natural" group. However, certain fossil discoveries suggest that the angiosperms evolved from a gymnosperm ancestor, which would make the gymnosperms a paraphyletic group if all extinct taxa are included. Modern cladistics only accepts taxa that are monophyletic, traceable to a common ancestor and inclusive of all descendants of that common ancestor. So, while the term gymnosperm is still widely used for non-angiosperm seed-bearing plants, the plant species once treated as gymnosperms are usually distributed among four groups, which can be given equal rank as divisions within the Kingdom Plantae: Pinophyta, Ginkgophyta, Cycadophyta and Gnetophyta .

Pinophyta The conifers, division Pinophyta , also known as division Coniferae , are one of 13 or 14 division level taxa within the Kingdom Plantae. They are cone-bearing seed plants with vascular tissue; all extant conifers are woody plants, the great majority being trees with just a few being shrubs. Typical examples of conifers include cedars, cypresses, firs, junipers, pines, and redwoods. Species of conifers can be found growing naturally in almost all parts of the world, and are frequently dominant plants in their habitats, as in the taiga, for example. The division contains approximately 700 living species.

Ginkgophyta The Ginkgo ( Ginkgo biloba ) is an unique tree with no close living relatives. It is classified in its own division, the Ginkgophyta , comprising the single class Ginkgoopsida , order Ginkgoales , family Ginkgoaceae , genus Ginkgo and is the only extant species within this

group. It is one of the best known examples of a living fossil. In the past it has also been placed in the divisions Spermatophyta or Pinophyta . Its seeds are not protected by an ovary wall. The apricot-like structures produced by female Ginkgo trees are technically not fruits, but are the seeds having a shell that consists of a soft and fleshy section (the sarcotesta ), and a hard section (the sclerotesta ). Ginkgos are known since the Permian. They diversified and spread throughout Laurasia during the Middle Jurassic and Cretaceous, but became much rarer thereafter. It is in fact doubtful whether the Northern Hemisphere fossil species of Ginkgo can be reliably distinguished; given the slow pace of evolution in the genus, there may have been only two in total; what is today called G. biloba (including G. adiantoides) , and G. gardneri from the Paleocene of Scotland. At least morphologically, G. gardneri and the Southern Hemisphere species are the only known post-Jurassic taxa that can be unequivocally recognised, the remainder may just as well have simply been ecotypes or subspecies. The implications would be that G. biloba had occurred over an extremely wide range, had remarkable genetic flexibility and though evolving genetically never showed much speciation. The occurrence of G. gardneri , it seems a Caledonian mountain endemic, and the somewhat greater diversity on the Southern Hemisphere, suggests that old mountain ranges on the Northern Hemisphere could hold other, presently undiscovered, fossil Ginkgo species. Since the distribution of Ginkgo was already relictual in late prehistoric times, the chances that ancient DNA from subfossils can shed any light on this problem seem remote. While it may seem improbable that a species may exist as a contiguous entity for many millions of years, many of the Ginkgo's life-history parameters fit. These are extreme longevity, slow reproduction rate, (in Cenozoic and later times) a wide, apparently contiguous, but steadily contracting distribution coupled with, as far as can be demonstrated from the fossil record, extreme ecological conservatism (being restricted to light soils around rivers), and a low population density (Holt & Rothwell 1997) . Extreme examples of the Ginkgo's tenacity may be seen in Hiroshima, Japan, where four trees, growing 1-2 km from the 1945 atom bomb explosion, were among the few living things in the area to survive the blast. While almost all other plants (and animals) in the area were destroyed, the Ginkgos, though charred, survived and were healthy. The trees are alive to this day (Lewington & Parker 1999). With special respect to a probable mass extinction event at the T/J boundary, Ginkgos can be interpreted as “event resistent”.

Cycadophyta Cycads are an ancient group of seed plants characterized by a large crown of compound leaves and a stout trunk. They are evergreen, dioecious plants having large pinnately compound leaves. They are frequently confused with and mistaken for palms or ferns, but are unrelated to either, belonging to the division Cycadophyta . Cycads are found across much of the subtropical and tropical parts of the world. They are found in South and Central America (where the greatest diversity occurs), Australia, the Pacific Islands, Japan, China, India, Madagascar, and southern and tropical Africa, where at least 65 species occur. Some are renowned for survival in semi-desert climates, and can grow in sand or even on rock. They are able to grow in full sun or shade, and some are salt tolerant. Though they are a minor component of the plant kingdom today, during the Jurassic period they were extremely common. They have very specialized pollinators and have been reported to fix nitrogen in association with a cyanobacterium living in the roots. These blue-green algae produce a neurotoxin called BMAA that is found in the seeds of cycads. The cycad fossil record dates to the Early Permian. There is controversy over older cycad fossils that date to the late Carboniferous period. One of the first colonizers of terrestrial habitats, this clade probably diversified extensively within its first few million years, although the extent to which it radiated is unknown as relatively few fossil specimens have been found. The regions to which cycads are restricted probably indicate their former distribution on the supercontinents Laurasia and Gondwana. The family Stangeriaceae , consisting of only three extant species, is thought to be of Gondwanian origin as fossils have been found in Lower Cretaceous deposits in Argentina. Zamiaceae is more diverse, with a fossil record extending from the Middle Triassic to the Eocene in North and South America, Europe, Australia, and Antarctica, implying that the family was present before the break-up of Pangaea. Cycads are the only genus in the family and contain 99 species, the most of any cycad genus. Molecular data has recently shown that Cycad species in Australasia and the east coast of Africa are recent arrivals, suggesting that adaptive radiation may have occurred. The current distribution of cycads may be due to radiations from a few ancestral types sequestered on Laurasia and Gondwana, or could be explained by genetic drift following the separation of already evolved genera. Both explanations account for the strict endemism across present continental lines. The probable former range of cycads can be inferred from their current global distribution. For example, the family Stangeriaceae only contains three extant species in Africa. Diverse

fossils of this family have been dated to 135 ma, indicating that diversity may have been much greater before the Jurassic and late Triassic mass extinction events. However, the cycad fossil record is generally poor and little can be deduced about the effects of each mass extinction event on their diversity (Gonzàlez-Astorga et al. 2003). Instead, correlations can be made between the number of extant gymnosperms and angiosperms. It is likely that cycad diversity was affected more by the great angiosperm radiation in the mid-Cretaceous than by extinctions. Very slow cambial growth was first used to define cycads, and because of this characteristic the group could not compete with the rapidly growing, relatively short-lived angiosperms, which now number over 95000 species, compared to the 947 remaining gymnosperms. It is surprising that the cycads are still extant, having been faced with extreme competition and five major extinctions in Earth’s history. The ability of cycads to survive in relatively dry environments where plant diversity is generally lower and their great longevity may explain their long persistence (Norstog & Nicholls 1997).

7.3 Sporomorph ecogroups

The Sporomorph Ecogroup Model (SEG Model) of Abbink et al. (2001) was applied to the palynological data set of the Furkaska section (App. 1) and the Komló cores 176 and 137 (App. 2, 3). In order to define SEGs for the Mesozoic and to model the response of SEGs to palaeoenvironmental change, the establishment of an ecological framework for the source communities is necessary. According to Grime (1979), two factors determine the type of plants defining a specific vegetation: stress and disturbance. The variation of the level of stress and disturbance results in three different plant strategies: a competitive strategy low stress, low disturbance, a stress-tolerating strategy high stress, low disturbance, and a ruderal strategy low stress, high disturbance, whereas highly disturbed habitats with severe and continuous stress are not viable as plant habitats (Grime 1979). Following this concept, Abbink et al. (2001) established six SEGs for the Late Jurassic-Early Cretaceous of NW Europe (see chapter 2.5). The Lowland SEG reflects the vegetation present in the main lowland area. The lowland represents an area with an optimum of nutrients and fresh water, low stress and low disturbance. The lowland plants are following a competitive strategy, and consequently, the ecological boundaries of the plants reflected by this SEG are determined by less than optimum conditions surrounding this lowland area. The plants within the communities reflected by other SEGs will follow a stress-tolerating or ruderal strategy.

Seaward, ecological stress is introduced by the influence of salt water in a tidally influenced area and salt spray in a coastal area. Landward, the possible deficiency of nutrients and/or the decreased availability of fresh water in the upland area may introduce ecological stress. Along rivers, periodical submersion and erosion of riverbanks will cause plants to follow a more ruderal strategy. A ruderal strategy will also be employed by first colonizers or pioneer plants. As for plants in the lowland area, the plants within the other ecologically defined areas will also show a competitive strategy, as they are optimized for the ecology of that particular habitat (Abbink et al. 2001). However, the grouping of fossil pollen grains and spores into these different ecogroups remains difficult. The application of the SEG Model to the Furkaska data set shows the limitations. The paleoenvironmental reconstruction based on sedimentological data points to a very shallow carbonate platform setting during late Rhaetian times. Sediments of the Triassic- Jurassic boundary interval document the change of a marine setting into a (probably brackish) prograding deltaic system. The sudden supply of freshwater and clastic material is caused by an increase of humidity (rainfall, weathering). Numerous interdisciplinary studies support this hypothesis for the NW Tethyan realm (e.g., Michalík 2003, Kuerschner et al. 2007). The SEGs of Abbink et al. (2001) show a different trend. During Rhaetian times the palynomorph assemblage is characterized by a high amount of pollen grains and spores of the “coastal” and “upland” SEG. But in the upper part of the section, the “warmer lowland” and “drier lowland” SEGs dominate the assemblage. Surprisingly, the “river” SEG is only represented by two spore genera. The number of spores of this SEG is decreasing upsection. Several factors may be responsible for this discrepancy between the paleoenvironmental and palaeoclimatological interpretation based on SEGs and other disciplines. First, the SEG model was created for the Upper Jurassic/Lower Cretaceous of the North Sea and it has to be taken into account that the climatological conditions during this period differed from the conditions at the Triassic/Jurassic boundary. A “drier” Upper Cretaceous climate was probably still wetter than a wet climate during the Hettangian. Therefore, the warmer/cooler and wetter/drier trends are not applicable for Triassic/Jurassic boundary palynomorph assemblages. Second, the parent plants for many Mesozoic sporomorphs are still unknown. Therefore, the SEG model is incomplete and many classifications are only based on morphological features. Third, the tolerance of many plant genera with respect to stress and disturbance are not well investigated. Therefore, many plants could be indicators of different environments, e.g. river and lowland (Abbink et al. 2004b). Thus, some of the sporomorph taxa of this study have been re-grouped with respect to the different conditions at the

Triassic/Jurassic boundary: Spore genera such as Concavisporites and Deltoidospora have been removed from the “drier” lowland SEG and placed within the “wetter” lowland and “river” SEG. In the Komló core sections of the S Hungarian Mecsek Mountains a similar trend like in the Furkaska section was recognized: The Rhaetian palynomorph assemblages are characterized by pollen grains and spores of the “upland” and “river” SEGs. Due to the terrestrial setting, the “coastal” SEG is hardly represented. In the upper part of the section, the “warmer lowland” and “drier lowland” SEGs dominate the assemblage. After the above described re- grouping, “wetter lowland” sporomorphs are dominating, accompanied by “river” SEG indicators.

7.4 Changes within the palynomorph assemblages and possible causes

The Triassic/Jurassic boundary interval is marked by one of the five biggest biotic extinctions during the Phanerozoic (Sepkoski 1996). Therefore, the lack of mass extinction within the microfloral assemblages of the NW Tethyan realm is surprising. However, the ultimate cause of the global biotic and environmental changes remains enigmatic. There are three main contenders that are considered as the possible drivers of these changes. The first is the emplacement of a Large Igneous Province (the Central Atlantic Magmatic Province, CAMP) that was associated with the initial break up of Pangaea (Wilson 1997), while the second is the possible impact of a large meteorite (Olsen et al. 2002) similar in size (10 km in diameter) to the one that is inferred to have impacted Earth 65 Ma ago at the K-Pg boundary (formerly referred to as the K/T boundary). The third involves the sudden dissociation of large amounts of methane hydrate (Beerling & Berner 2002). Although the influence of any one of these mechanisms would not necessarily rule out the operation of the others, the recognition and precise quantification of the environmental effects that took place across the Triassic/Jurassic boundary can help to identify and understand the most likely cause of the global changes at that time (Cohen & Coe 2007). While the evidence for large-scale volcanism near the Triassic–Jurassic boundary is incontrovertible (Marzoli et al. 1999), support for an extraterrestrial impact at that time is much more contentious. Shocked quartz has been reported from a locality in Italy (Bice et al. 1992), although the significance of this evidence has been challenged (Hallam & Wignall 1997), while very minor enrichments of Ir have been detected in organic-rich layers close to the Triassic-Jurassic boundary at some localities in the

Newark Basin of North America (Olsen et al. 2002). These mildly elevated Ir abundances were interpreted by the authors as evidence that a meteorite impact had occurred at the Triassic/Jurassic boundary. The authors further inferred that the effects of the proposed impact had been responsible for the environmental changes at that time, which, amongst other things, had led to the rise of the dinosaurs. However, their evidence for a substantial impact at the Triassic-Jurassic boundary is considered to be controversial because of (1) the small magnitude of the enrichment, with maximum Ir abundances of only up to 285 ppt (compared, for example, with the much higher levels of Ir, sometimes up to hundreds of ppb, in some K- Pg boundary strata; cf. Koeberl et al. 1994, Koeberl & MacLeod 2002); (2) the association of the purported Ir anomaly with coaly layers and organic-rich strata, which have the ability to sequester Ir and associated elements during deposition and subsequent diagenesis; and (3) the absence of any other firm evidence for a large impact at that time in either these or in any other sedimentary deposits of a similar age (Cohen & Coe 2007).

Fig. 31: The “Neves effect”. Transport of different palynomorph morphotypes in different depositional environments (after Chaloner & Muir 1968).

The most obvious change in the microfloral assemblages that could be detected in all sections is the sudden increase in the abundance of trilete spores. Spores are produced by plants, which need a high humid environment like ferns and horsetails. The cause of the spore spike is 76

supposed to be an increase in humidity during the boundary interval. Due to the fact that the study areas belong to different palaeoenvironments, the sorting of the palynomorphs during transportation should be considered. A striking demonstration of the fact that spores and pollen are selectively sorted by sedimentation factors is given in Figure 27. Saccate pollen are carried by both, wind and water into marine environments, whereas Lepidendron microspores are produced in swamps and are not much transported out of the swamps. Some other spores are produced in other palaeoenvironments and float reasonably well towards the sea (this phenomenon is known as the “Neves effect”). Palynofloras from coals are practically 100 % authochthonous (Traverse 1988). The fact that the spore spike in the marine sections shows the upper extremity underlines the dimension of this floral change. Van der Schootbrugge et al (2007) observed a similar increase in the abundance of trilete spores in sections in Germany and the North Sea region. He interpreted the vegetation change as a “waldsterben” event related to the high amount of CO2, which was released to the atmosphaere by the volcanic activity in the CAMP region. Conifers are plants that are sensitive to acidification and would suffer due to acid rain. Many spore producing plants prefer a more “sour” soil. Van der Schootbrugge et al also described a simultaneous increase in the amount of spores and prasinophytes that are interpreted as disaster species. The increase of Kaolinite in the Furkaska section is also interpreted as a result of soil weathering due to acidic rain.

8 Palaeoenvironmental interpretation of the studied areas

8.1 Tatra Mountains During late Triassic and early Jurassic times, the sections studied were part of the Tatro- Verporic Unit, situated at the NW Tethyan shelf area (Fig. 7). The sediment series of the Kardolína and Furkaska sections, belonging to the Zliechov subunit, were deposited in a shallow marine environment. In the lower part of the Kardolína section, findings of brachiopods, crinoids, bivalves and gastropods point to normal marine conditions. The carbonate platform was populated by small coral biostromes. Upsection, the input of siliciclastics is increasing and hinders the carbonate production. The sporomorph assemblage reflects the palaeoenvironmental conditions of the hinterland, which was part of the Bohemian massive. The relatively low content of Corollina spp. within the Rhaetian part of the section points to warm subtropical climate (cf. Vakhrameev 1981). These results disagree with studies from the Northern Calcareous Alps, where a high amount of Corollina points to semi- arid conditions (Holstein 2004).

In the boundary interval of the Furkaska section, a lithofacies change from carbonates to clays indicates a characteristic palaeoenvironmental turn over. The sudden increase of clastic sediment input and the simultaneous increase of trilete spores within the palynomorph assemblages point to an increase of humidity. The distribution of clay minerals supports this hypothesis. Kaolinite, an important climate-sensitive clay mineral indicator, appears only in the boundary claystones, while underlying strata are free of kaolinite. Moreover, the Furkaska section is also characterized by a relatively continuous increase of mixed-layer illite-smectite content at the expense of discrete illite (Fig. 32).

Fig. 32: Distribution of clay minerals, T/J boundary interval of the Furkaska section (from Ruckwied et al. 2006).

These changes are interpreted here as response of increasing intensity of chemical weathering in the hinterland due to increasing humidity (Michalik et al. submitted). Heavy rainfalls caused the development of a receiving stream that transported the sediment in a deltaic system towards the coast. The clays of the lower Hettangian Kopienic Formation are interpreted as deposits of the distal part of the deltaic system (Fig. 33). The presence of marine plankton indicates marginal marine conditions. 78

Fig. 33: Palaeoenvironmental reconstruction of the Tatra Mountains during Rhaetian-Hettangian times (from Michalík et al. 2007).

8.2 S Hungary During late Triassic and early Jurassic times, the Mecsek Mountains were part of the Tizsa Unit, which belongs to the NW Tethyan realm (Fig. 7). The terrestrial environment was dominated by a fluvial system and turned periodically to a swamp. During upper Hettangian/Sinemurian times, a transgression caused the flooding of this area. The palynomorph assemblage of the Upper Rhaetian part of the section comprises a small amount of Corollina spp., which points to relatively temperate climatic conditions (cf. Vakhrameev 1981). The increase of trilete spores within the Triassic/Jurassic boundary interval (Fig. 230) indicates an increase of humidity. This global palaeoclimatic signal is superimposed by a local palaeoenvironmental change from fluvial to swamp conditions. These changes are documented by two distinguished microfloral assemblages that are related to different palaeoenvironments. Coal layers contain assemblage A which is dominated by Inaperturopollenites sp.; sand- and siltstone layers comprise assemblage B, mainly composed of trilete spores and bisaccate pollen grains (Fig. 34). 79

Fig. 34: Palaeoenvironment reconstruction of the two vegetation phases detected in the Mecsek Mountains, S Hungary.

8.3 N Hungary The Cs ővár section, situated on the Dachstein carbonate platform, belongs to the Tethyan Transdanubian Range (Fig. 7). Facies analysis of the Rhaetian-Hettangian deposits reveals a long-term change in sea level, superimposed by short-term fluctuations. After a period of highstand platform progradation in the Late Norian, a significant sea-level fall occurred in the

Early Rhaetian, exposing large parts of the platform. A renewed transgression led to the formation of smaller build-ups fringing the higher parts of the previous foreslope that served as habitat of crinoids, representing the main source of carbonate turbidites. The higher part of the Rhaetian is characterised by proximal turbidites with intercalated lithoclastic debris flows. Distal turbidites and radiolarian basin facies become prevalent upsection, dominating in the earliest Hettangian. The next significant facies change in the Early Hettangian is marked by the appearance of redeposited oncoid-grapestone beds, indicating the end of the Rhaetian to earliest Hettangian sequence (Haas & Tardy-Filácz 2004).

Palynofacies of the sedimentary series exposed in the Pokol-völgy quarry is dominated by terrestrial components, reflecting a high supply from the hinterland. Numerous needle-shaped opaque particles, as well as a high amount of large translucent plant fragments within the phytoclast group, may point to the transport mechanism of sedimentary organic matter, strongly related to the occurrence and frequency of turbidites along the slope. The presence of prasinophytes is characteristic of a permanently stratified deeper basin. The most striking feature within the boundary interval is the synchronous peaks of prasinophytes and trilete spores. The co-occurrence of spikes in both the marine and terrestrial signals is described for the first time from a marine boundary section. The prasinophyte and spore peaks also correspond to the previously documented prominent negative carbon isotope excursion and are proposed as a potentially powerful correlation tool. The inferred marine algal bloom and the temporary dominance of ferns in the terrestrial vegetation may signal the biotic response to the same environmental stress, which also affected the carbon cycle. The relative high amount of Corollina spp. points to semi-arid conditions of the hinterland. Palynofacies of the carbonates exposed in the upper part of the Vár-hegy section, dated as Lower Hettangian, is dominated by degraded organic matter, small equidimensional phytoclasts and foraminiferal test linings, pointing to a distal basinal setting. The dominance of turbidites, together with the preservation and composition of sedimentary organic matter, supports the complex basin topography proposed by Galácz (1988; Fig. 35).

Fig. 35: Palaeoenvironmental reconstruction of the Transdanubian Range (N Hungary) during late Triassic – early Jurassic times (modified after Galácz 1988).

9 Summary

Late Rhaetian/Hettangian microfloras of three depositional environments of the NW Tethyan realm have been studied: platform to basinal limestones of the Csövar section (N Hungary), shallow marine limestones and marls of the Tatra Mountains (N Slovakia) and terrestrial coal deposits of the Mecsek Mountains (S Hungary). The comparison of the palynological assemblages builds the base for interpretation and correlation of the different depositional environments. Additionally, the sporomorph distribution and diversity were investigated with respect to the Triassic/Jurassic mass extinction event and the processes that may have caused this crisis.

The palynomorph assemblages of the settings studied display typical Rhaetian/Hettangian microfloras, dominated by bisaccate pollen grains, trilete spores and pollen of the Circumpolles group. A floral mass extinction was not recognised in the NW Tethyan realm. Palynomorph assemblages of the Cs ővár section are similar to the assemblages of the Germanic realm comprising marker species such as Rhaetipollis germanicus and a high amount of Classopollis spp. in the Upper Rhaetian part of the section. Due to the close paleogeographic position of the Tatra Mountains and the Northern Calcaerous Alps, the palynomorph assemblages of these areas are very similar. The sporomorphs of the S Hungarian Mecsek Mountains can be divided into two different assemblages, displaying a

characteristic cyclic vegetation pattern of a fluvial system turning periodically to a swamp area. Coal layers contain assemblage A, dominated by pollen grains of the genus Inaperturopollenites . Sand- and siltstone layers comprise assemblage B, mainly composed of trilete spores and bisaccate pollen grains.

Due to the lack of Lower Jurassic marker species such as Cerebropollenites thiergartii , a correlation of the study areas using FADs and LADs is not possible. All investigated assemblages comprehend a significant shift in the abundance of trilete spores within the Triassic/Jurassic boundary interval. The abundance of trilete spores is at least 15% higher in the Hettangium than in the Rhaetian. This striking quantitative change is a very valuable tool for correlation and points to an increase of humidity during this period, representing an overregional signal. In N Hungary a prasinophyte bloom was detected together with a spore spike during T/J boundary times. The occurrence of prasinophyte blooms in sections from different regions may support this hypothesis. Thus the prasinophyte spike in the Cs ővár section is likely to record a geographically widespread event rather than a local palaeoecological phenomenon. Similar to our observations at Cs ővár, a spore spike at the Triassic/Jurassic boundary was reported from the Newark Basin in the eastern US (Olsen et al., 1990; Fowell et al., 1994). A sudden increase in the relative abundance of spores was also recognized in several other sections of the NW Tethyan realm (Ruckwied et al 2009) a well as in the Danish-Germanic Basin (Heunisch et al. 2008) and in Sweden (Schootbrugge et al 2007). Therefore this change in the sporomorph assemblage is seen as a reflection of a supra-regional change in the hinterland vegetation. Due to the fact that this signal is observed in sediments deposited in different paleoenvironmental settings, it is unlikely to be caused by sorting or preservation. Schootbrugge et al. (2007b) investigated an abrupt change in the microfloral assemblages of two cores from Germany and Sweden. They reported that conifers, seed ferns and cycads- ginkgophytes have been replaced by herbaceous ferns and fern allies during the T/J boundary interval and proposed that this floral turn-over could be triggered by the CAMP volcanism. The release of sulfur, causing short-term cooling and a regional acidification of terrestrial ecosystems through the formation of sulfuric acid (H 2SO 4) rain. Heunisch et al. (2008) observed a contemporaneous turnover of marine phytoplankton communities and interpreted this changes to be not only driven by changes in humidity/aridity and/or sea-level changes but as a response of severe environmental changes that were most likely triggered by the CAMP volcanism. A more pronounced fern spike is well known from the Cretaceous/Paleogene

boundary (e.g. Tschudy et al., 1984). The spore spike, recording a sudden change of the terrestrial vegetation, and the prasinophyte bloom, reflecting a similarly abrupt event in the marine realm, indicate a significant perturbation of the biosphere at the Triassic/Jurassic boundary. A correlative signal is also documented in the δ13 C isotope record. Hesselbo et al. (2002) discussed a causal relation of the negative δ13 C excursion and the initial volcanic activity of the Pangaean Atlantic rifting. Recent 40 Ar/ 39 Ar dating of flood basalts in Morocco and Portugal (Nomade et al., 2007; Verati et al., 2007) confirm the isochroneity of the Central Atlantic Magmatic Province (CAMP) volcanism and major changes in marine and terrestrial ecosystems at the Triassic/Jurassic boundary. Alternatively, Olsen et al. (2002) postulate a bolide impact as the main trigger for the drastic changes at the end of the Triassic. In the lack of substantial evidence for an end-Triassic impact, our data from Csővár are fully consistent with the model where the initial volcanic activity of the CAMP is related to climatic change, the negative δ13 C excursion, and also leads to the perturbation in marine and terrestrial ecosystems at the Triassic/Jurassic boundary.

Fig. 36: Distrubition of Early Jurassic plateau basalts of the Central Atlantic Magmatic Provence (CAMP), from McHone (2000).

10 Outlook

Triassic/Jurassic boundary sections of the Tatra Mountains and the Transdanubian range are well studied with respect to palaeoenvironmental changes during this period. Palynological investigations built one important tool for multi-disciplinary interpretation of climatic change and possible causes. 85

Sedimentary deposits of the S Hungarian Mecsek Mountains yield a high potential for high- resolution stratigraphy due to their cyclic sedimentation pattern. Therefore, these successions are seen as key sections for ongoing studies focussing the short-term climatic change during the Triassic/Jurassic boundary interval. The palynological results of this study, combined with existing palaeobotanic data of the Mecsek Coal (e.g., Barbacka 2000, 2001, 2002), may build the base for a reconstruction of vegetation pattern based on the SEG model of Abbink et al. (2001). Other areas of interest are the Newark Basin (USA), the Junggar Basin (China) and the Madygen area in Kyrgyzstan (Central Asia). These areas are characterized by thick continous Rhaetian/Hettangian sediment series and palaeoclimatic investigations of these areas are crucial for a global palaeoclimatic reconstruction of the Triassic/Jurassic boundary interval. Finally, the integration of palynological data into a multi-disciplinary study, using sedimentology, geochemistry, clay mineralogy and micro- as well as macropalaeontology will clarify our picture of the Earth during Rhaetian-Hettangian times. The IGCP 458 project represented the first integrated approach of such a global multi-proxy study.

11 References

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