Lunz flora in the Austrian Alps — a standard for Carnian floras

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Palaeogeography, Palaeoclimatology, Palaeoecology 143 (1998) 307–345

Lunz flora in the Austrian Alps — a standard for Carnian floras Inna A. Dobruskina * The Hebrew University of Jerusalem, The Institute of Earth Sciences, Jerusalem, 97904, Israel Received 1 September 1997; accepted 1 March 1998

Abstract The Lunz flora of Niedero¨sterreichische Alpen is the most famous and for a long time was the best known flora of the Carnian Stage (Upper Triassic). It occurs in marine deposits with marine invertebrates, so that its geological age is exactly known, so it provides a standard by which to correlate terrestrial floras of Carnian age to the marine Triassic timescale. Fossil plants from the Lunz beds were collected from more than ten localities in the Northern Limestone Alps not far from Lunz, where they are confined to the so-called ‘Lunzer Sandstein,’ which is part of the ‘Lunzer Schichten.’ The flora, reviewed and analysed here in detail, consists of numerous sterile leaves of Cycadophyta, of abundant ferns and Sphenopsida and of rare Ginkgophyta and Coniferophyta.  1998 Elsevier Science B.V. All rights reserved. Keywords: Upper Triassic; correlation; floras; Austrian Alps

1. Significance of the Lunz fossil plant determinations and interregional correlation The Lunz flora of Niedero¨sterreichische Alpen (Fig. 1) is the most famous and for a long time was the best known flora of the Carnian Stage (Upper Triassic). It was discovered about 150 years ago, and many species of fossil plants were established for the first time in this flora. Later, such species were determined in other floras by comparison with Lunz plants. The Lunz flora occurs in marine deposits with marine invertebrates, so that its geological age is exactly known. This flora was the basis for the correlation of German and Alpine facies of the European Triassic. The question for a long time was which flora from the German basin — Lettenkohle or Schilfsandstein — was to be correlated with the Carnian Lunz flora. Ł Fax:

C972 (2) 622581. E-mail: [email protected]

Thus, the exact correlation of the Triassic of the German basin and the Alps depends very much on the exact determination of the Lunz plants. The Lunz flora was at the same time the basis for the dating of many intercontinental floras of Eurasia and America which had no association with marine

Fig. 1. Geographical position of localities Lunz (1), Raibl (2), Basel (3), Salzburg (4), Bleiberg, Viktorhu¨tte (5), Liptovsky Gradok (6), San Cassian (7).

c 1998 Elsevier Science B.V. All rights reserved. 0031-0182/98/$ – see front matter PII: S 0 0 3 1 - 0 1 8 2 ( 9 8 ) 0 0 1 1 6 - 3

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fossils: the age of the Upper Triassic floras of the Donez basin (Stanislavskiy, 1976), Priuralye (Brick, 1941; Neuburg, 1959), the Urals, Middle Asia, Mongolia (Dobruskina, 1980), and of China (Sze, 1956) was established through comparison with it (and with coeval floras of the German basin). The Lunz flora was basic to inferring the Late Triassic age of the North American flora of Virginia, USA, at the end of the last century (Stur, 1888). Thus, it is a standard flora. The term ‘standard flora’ is used here to denote floras whose geological age can be determined directly with marine fossils (Dobruskina, 1982), as the geological timescale for the Triassic (as well as for most of the Phanerozoic) was established by a succession of marine fossils. Apart from the Lunz flora, we have some other standard floras for the Carnian: the floras from Svalbard, Soviet Primorye and Japan. However, the stratotype of the Carnian Stage occurs in the Alps, and the Carnian in other regions was established after a correlation with the stratotype by using marine fossils. The direct correlation of plant-bearing beds with the Alpine stratotype is certainly more precise. The Lunz flora is very rich and diverse, fertile ferns and fertile parts of other plants being most important; good preservation of fossils permits the study of the anatomy of leaves and spores in situ. Thus, the importance of the Lunz flora is due to more precise correlation with the stratotype of the Carnian (Zalzkammergut), a longer history of study, and the original description of many species and the possibility of cuticular and palynological studies. A large collection of the Lunz flora in the Museum of the Geologische Bundesanstalt in Vienna is the basis of this study. The collection was originally studied by D. Stur; there are still some labels written by him. It contains more than 16,000 specimens with more than three thousand imprints of fossil plants. Smaller collections are found in the Naturhistorisches Museum of Vienna, Palaeontological, Botanical and Ge¨. ological Institutes of Vienna University, and N.O Landesmuseum in Vienna. Other important collections are stored at numerous museums in Europe, in Austria and elsewhere. In Austria they are in Graz (Loanneum), Lunz (Amonhaus), Waidhofen, Mo¨dling, Innsbruck (Botanical Institute of University), and Leoben (the last collection is not accessible

now). R. Kra¨usel mentioned Lunz material from the Geological Survey of Berlin, Naturhistorisches Museum in Berlin, Staatliche Naturaliensammlung in Stuttgart, Geological Institute of Tu¨bingen University, Senckenberg Museum in Frankfurt, Naturhistorical Museum in Stockholm and the Geological Museum in Strasbourg. I have seen a collection in Lyon University and photos of specimens in Meiningen, Germany. All these collections are very important as it is impossible now to obtain new material, because the main source for Lunz collections were coal mines that are closed. In the middle of the last century some of those were operating exclusively to dig out fossil plants; that is the reason for the abundance of Lunz fossils in various collections in Europe. The importance of the careful study of the Lunz flora is evident, and it is strange that we still do not have a complete description of it. There were many good palaeobotanists who dealt with this flora, but none managed to complete investigations. There are more than a dozen of papers on the Lunz flora, but only the first of them (Stur, 1885) gives a complete (though preliminary) list of all plants from the Lunz beds, comparison of the flora as a whole with other fossil floras (especially with Basel and Raibl), and determination of plant groups. Some specimens (the most interesting from the botanical point of view) were described several times, while most were not described at all. This gave a distorted picture of the relationship of the main plant groups in this flora. Many specific and even generic names are used without any detailed study after the preliminary determinations of Stur, often following names inscribed on the labels; for many species, it is a tradition. But is clear that such determinations cannot be valid. The complete picture of the flora under consideration may be compiled from studies of all the collections and their systematic monographic investigation. The present paper has resulted from the first stage of investigations and may be considered as an inventory of the greatest part of the Lunz collection. A review of this flora from different points of view is given here: analysis of its geological and stratigraphical position, localities, composition of different collections, palaeobotanical publications, and the history of study. Such analysis was made on the basis of earlier ideas on the position of the Lunz flora among the Triassic floras of Eurasia (Dobruskina, 1980, 1982).

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Fig. 2. Localities of the Lunz plants in the Northern Alps.

2. Position, geological situation, stratigraphy of plant-bearing beds Fossil plants from the Lunz beds were collected from more than ten localities in the Northern Limestone Alps not far from Lunz; they extend in a WSW– ENE oriented narrow belt within the Lunzer Decke nearly from Lunz to Vienna (Fig. 2). They are confined to the so-called ‘Lunzer Sandstein,’ which is part of the ‘Lunzer Schichten.’ ‘Lunzer Schichten’ is a terrigenous coal-bearing formation which underlies and overlies limestones, as in Fig. 3 (after Verloop, 1908). In other parts of the Northern Limestone Alps this terrigenous formation is replaced by limestones. Terrigenous deposits of the Lunz beds, but without fossil plants, extend northeast into Slovakia; palynological study demonstrates that they are coeval with the plant-bearing beds (Planderova, 1972).

Palynological investigations of the Lunz beds were made also in the Lunz environs. Five stratigraphically important species were described (Klaus, 1960) from the Lunz beds; a palynological assemblage (37 miospore species) was described from coal interlayers of the Lunz beds (Bhardwaj and Singh, 1956). Klaus (1960) and Kavary (1966) also investigated palynological assemblages from the Lunz beds correlatives: Cardita beds of the Gailtal Alps (Bleiberg and Viktorhu¨tte) and the underlying Halobia beds in the vicinity of Salzburg. Pelecypods are known throughout the Lunz section: in the upper part of the Reiflinger Kalke, Reingrabener Schiefer, Lunzer Schichten and the lower part of the Opponitzer Schichten. Tetrapods are known from the lower part of the Reingrabener Schiefer and from the Lower Opponitzer Schichten: cf. Francosuchus trauthi Huene (Huene, 1939) from

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Fig. 3. Position of plant-bearing beds in Lunz section (after Verloop, 1908): 1 D dolomites; 2 D limestones; 3 D marls; 4 D sandstones; 5 D argillites; 6 D coals; 7 D conglomerates; 8 D fossil plants.

the Lunzer Bergstollen near Lunz together with Anoplopora (Anodontophora) lettica Qu and other pelecypods. Fishes are known from the Reingrabener Schiefer. Following the investigations of Krystyn (1978), the Lunz beds are referred to the uppermost part of the lower Carnian (Julian), substage to the upper part of the Austriacum Zone. The thickness of the Lunzer Schichten is about 140 m (Verloop, 1908).

3. Lunz flora collection in the Geological Survey of Austria Collections of the Geologische Bundesanstalt represent the material studied by D. Stur. Specimens carry labels written by him or rewritten later after earlier ones were destroyed. The majority of the specimens are deposited in the order of their description (without figures) in two of Krasser’s papers of 1909; described and figured specimens are contained in special separate collections in agreement with the

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respective papers. During my first short visit to the GBA, I looked through all the material and numbered all specimens in agreement with this arrangement. Unfortunately, I could not understand how it was possible to distinguish the originals from Krasser (1909a) and Krasser (1909b), because figures were absent, as well as references to the concrete specimens or to their numbers (some of the specimens had original numbers but without reference to the papers or any sort of inventory). There are seven collections of Lunz plants in GBA: (1) 1909=2 corresponds to the paper by Krasser, 1909a (573 spec.); (2) 1909=3 corresponds to the paper by Krasser, 1909b (982 spec.); (3) 1917=1 corresponds to the paper by Krasser, 1917 (56 spec.); (4) 1919=2 corresponds to the paper by Krasser, 1919 (9 spec.); (5) 1942=1 corresponds to the paper by Kra¨usel, 1942 (1943) (34 spec.); (6) 1947=1 corresponds to the paper by Langer, 1945 (1 spec.); and (7) 1966 corresponds to the paper by Kra¨usel and Schaarschmidt, 1966 (3 spec.). There are 1658 specimens, together with more than 3414 imprints of fossil plants. Actually, there are many more imprints, because many specimens are bedded rocks in which many layers with imprints are distinguishable, but it is impossible to make a preparation without destroying the upper layer of imprints. Concrete localities of the Lunz plants in the GBA are shown in Fig. 2. Most specimens in the GBA show the exact location, but the locations of some of them are reduced to one word, ‘Lunz’. Whether it means that the specimen occurs in Lunz itself, or the Lunz area, is not clear. Unfortunately, publications by Kra¨usel often show the same location — Lunz without specification. The same occurs in the collections of Lunz plants in Lyon and Meiningen. Table 1 shows the distribution of different taxa of the Lunz flora in specific localities in the Lunz area. Most specimens were collected in Pramelreuth near Lunz (Fig. 4). Evidently, this is due to the special fossil-yielding mines and not to the original fossil richness of the area. The second place belongs to Ahornberg, also near Lunz, the third, to Seestollen at Lunz. From these three localities, more than one thousand specimens (2633 imprints) were collected, among them more than 800 occurred in Pramelreuth (2239 imprints). There are about 3000 specimens in

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the Lunz area sensu stricto (Lunz environs: 5622 imprints) deposited in GBA collections that are not illustrated in publications. There are 53 specimens with 103 imprints from Kirchberg at the Pielach River in the GBA collections. About 100 specimens with 157 imprints come from the vicinity of Lilienfeld. From the first discovered locality — Tonibaueralpe — there are 37 specimens with 54 imprints in the GBA. From 1 to 15 specimens are from other areas. Tables 1 and 2 show that various groups of plants are distributed unevenly, especially sphenopsids. Whereas they constitute less than 4% in the richest localities of Pramelreuth and Seestollen, they reach about 50% in Ahornberg. It seems that this picture reflects the original situation. In the Kirchberg area they constitute 25%, and in the Lilienfeld area, 36% of all plants. Various ferns are known only from Lunz localities (Pramelreuth, Ahornberg and Seestollen); from others, ferns are absent or represented only by Asterotheca. Sterile leaves of Pterophyllum are abundant in all the localities, whereas Taeniopteris (Macrotaeniopteris) is confined to Lunz localities and to Lilienfeld. In the rest of the localities, they are absent or few of the imprints were found. Fertile parts of Cycadophyta are known from all the localities of the Lunz area and some others. Their quantity seems to correspond to more intense collecting and not to the original richness of this plant group here. The Glossophyllacea are abundant only in Pramelreuth (203 of 2239 imprints), where they compose about 9.6% of all the plants. Their relative role is the same in Seestollen (8.7%) as well as in Lunz (6.6%). It is much less in Ahornberg (0.2%) of the same area. As a whole, they are more abundant in the west of the studied area (Hollenstein 9.5%, Tonibaueralpe 7.4%, Lunz environs 7.5%), less in the centre (Kirchberg 0.3%) and absent in the east (Lilienfeld, Kleinzell, Ramsau). The majority of Glossophyllaceae is Glossophyllum; Ginkgoites is represented by sixteen imprints from the Lunz localities and one imprint from Tonibaueralpe. In the rest of the localities, they are not known. Imprints of Stachyotaxus were collected in relatively large quantities (166 imprints) from about half of the localities, and the majority are confined to the Lunz area: 34 imprints were collected in Pramel-

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Fig. 4. Geological section in Pramelreuth (after Verloop, 1908).

reuth, 4 in Ahornberg, 5 in Seestollen and 114 imprints in the area indicated as ‘Lunz’. From other areas, only single specimens are known. Since this genus was originally described as Pterophyllum?, one cannot exclude that there are more representatives of this conifer in the Lunz flora than we know now. Described regularities in the distribution of fossil taxa in a west–east direction to some degree depends on the intensity of collection by I. Haberfelner in the Lunz environment. Based on the results of Dr. R. Sachsenhofer (oral communication) on regularities

of coalification there were original peculiarities in the distribution of plant assemblages. It is possible, that richer assemblages originally existed in the west of the area under consideration. Table 1 also shows that the main role in the Lunz flora belongs to the Cycadophyta (4079 imprints of 6630), and especially to sterile leaves (3855 imprints). Among them, the main role belongs to numerous Pterophyllum (2846 imprints). Cycadophyta constitute more than half of all imprints. Fertile parts of the Cycadophyta are represented by 241 imprints only. More than half occur in Pramelreuth.

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Table 1 Composition of different collections of the Lunz flora (quantity of imprints) Lunz flora (1986–1987)

GBA

Sphenopsids

396

139

64

14

Ferns Dipteridaceae Asterotheca Bernoullia Danaeopsis Pecopteris others

758 163 319 26 99 20 131

292 57 104 5 50 17 59

79 3 67 1 2 – 6

36 1 24 1 7 1 2

Pteridopsperms?

8

–

–

–

Cycadophyta sterile (incl. Pteroph.) fertile Glossophyllum

Naturhist. Museum

1923 1198 1744 1157 (1215) (944) 179 41 244

55

Ioanneum

708 695 (505) 13

Ammonhaus

107 102 (74) 5

¨ NO Landesmuseum

Innsbruck

Waidhofen

Lyon

Meiningen

6

3

6

2

3

15 2 12 – – – 1

13 2 6 – 4 – 1

2 – 1 – 1 – –

24 4 12 2 3 2 1

6 – 5 – 1 – –

–

–

–

1

–

65 63 (37) 2

12 12 (8) –

2 – (1) 1

59 59 (40) –

23 23 (22) –

127

9

11

5

–

11

–

Ginkgoites

17

1

–

–

–

–

–

–

–

Stachyotaxus

68

61

50

3

2

–

–

1

1

(Total 6499)

3414

1746

1028

169

99

33

10

98

33

Sterile leaves were collected in all localities in equal amounts. The second place belongs to the ferns. They are more scarce, though the absolute quantity is also impressive: 1225 imprints. Cycadophyta and ferns compose the majority of the Lunz flora. The role of Ginkgophyta and Sphenophyta is considerably less: their amount is nearly the same. There are very few conifers; pteridosperms and lycopsids are either entirely absent or represented by doubtful remains (for discussion of the systematic position of the Glossophylaceae, see below). Only 86 specimens from GBA collections are figured and discussed in detail in publications, they belong to fertile Cycadophyta, Ginkgophyta and Coniferophyta. The ferns and sphenopsids from GBA collections have not figured in publications.

4. Lunz flora collection in the Naturhistorisches Museum, Vienna The second great collection of Lunz plants belongs to the Vienna Naturhistorisches Museum. Most of the specimens were received from J. Haberfel-

ner in 1881–1888 (especially in 1885). More than 30 specimens were received from C. Ettingshausen in 1878 and 1879. Two specimens have labels – ‘Pramelreuth. A. Handlirsch, 1905’, three – ‘Lunz A. Mariani, 1931’, nine – ‘Neuer Stollen Ybbsthal. F. Mu¨ller, 1934’. All other labels are written by Haberfelner. Only these specimens have exact localities, all others show ‘Lunz’ labels. Because the majority of them were sent by Haberfelner, it means that they occur in the nearest vicinity of Lunz, possibly all of them from Pramelreuth. Very large specimens which are in the permanent exhibition of the Museum are of extraordinary interest because of their great dimensions. Such very large specimens are rare in collections; in these specimens we can see parts of Lunz plants which are unusual. And it is very important for this flora, because the majority of plants had giant fronds, and we have the possibility in this Museum to see the whole fronds of ferns and cycadophytes, not only different parts of them. During the war Prof. R. Kra¨usel studied Lunz plants from this collection as well as from GBA. In the boxes one can see sheets of paper with his

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Table 2 Distribution of fossil plant taxa in Lunz localities (quantity of imprints)

notes that he selected for studying Glossophyllum, Stachyotaxus, and fertile parts of cycadophytes. But after his publications were ready he did not mark figured specimens and now it is difficult to identify this material. This work has to be done by palaeobotanists who are interested in the Lunz flora. Now, Dr. P. Crane from Chicago has begun to deal with Kra¨usel’s originals. Most of the collection consists of sterile leaves of Cycadophyta; as usual nearly 70% are imprints, Pterophyllum being the most common. Fertile parts are not more than 5% of all remains of Cycadophyta. The second place belongs to ferns, the third to the sphenopsids, the fourth to Glossophyllaceae and the last to the conifer Stachyotaxus. It corresponds very well to the percentage of these groups in the whole Lunz flora.

5. Lunz flora collection in Joanneum, Graz This collection was received from Prof. F. Krasser in 1902, 1904 and 1909. In addition, there are 39 specimens received as gifts from Graz University and 126 specimens collected by Marktanner. Some of the specimens have old labels, but nobody hasdealt with them for nearly a century. This collection does not contain extraordinary material, as far as I can judge after quick acquaintance with it, but confirms the relationships of main groups in the flora and provides good material for study. It consists of 234 specimens with 1026 imprints, the majority of which are also sterile cycadophytes: 497 imprints among 709 cycadophytes are Pterophyllum, and only 14 imprints belong to fertile parts of Cycadophyta. It is natural that F. Krasser did not send the most

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interesting specimens because at that time he had already begun to study this part of the Lunz flora. A relatively large quantity of Glossophyllum is present in this collection. Ferns, sphenopsids and conifers (Stachyotaxus) are present in equal quantities.

6. Lunz flora collection in Niedero¨sterreichisches Landesmuseum in Vienna There is a small, but very interesting collection in ¨ Landesmuseum, because the specimens have the NO indications of the exact localities from which they were obtained. The knowledge of the exact distribution of fossil plant taxa may be interesting from different points of view. The first point is ecology of the Triassic in the area of the Lunz beds, the second is regularities in sedimentation and relationships with coalification, and the third is associations in plant assemblages. From the last point of view, the locality of Schrambach is of special interest. Antholites wettsteini — a male fructification — was described from this locality by R. Kra¨usel. He connected this fructification with Ginkgoites lunzensis, and some palaeobotanists later connected it with Glossophyllum florinii. But among specimens which have labels ‘Schrambach’ there are no Glossophyllum or Ginkgoites. Seventeen ‘new’ specimens from ¨ Landesmuseum also do not the collection of NO contain these genera. This collection contains 42 specimens with 99 imprints. Specimens from Schrambach were collected by Prof. R.J. Mayrhofer in 1929, except one specimen marked ‘1882’. Five specimens were received from J. Haberfelner in 1895, one from C. Ettingshausen in 1879, eight in 1932 from Lunz (name of collector is absent) and four in 1914 from the environ of Gross Hollenstein a.d. Ybbs (name of collector is absent, too). As usual, 65% of the imprints are sterile cycadophytes, nearly all being Pterophyllum, Glossophyllum (from Lunz and Gr. Hollenstein), and ferns are nearly equal (11% and 15%); sphenopsids are 6%, and Stachyotaxus is 2%.

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7. Lunz flora collection in Amonhaus in Lunz-am-See All specimens in this collection have no other location than ‘Lunz’ and no indication as to collector or year of collecting. I have seen 96 specimens with 169 imprints. These are in the Biological Station in Lunz-am-See and in private collections (for example, several good Danaeopsis with Dr. A. Ruttner). The collection of Amonhaus also, as usual, contains 63% sterile leaves of cycadophytes, mostly Pterophyllum longifolium; there are many ferns (22%); sphenopsids and Glossophyllaceae are nearly equal (8% and 9%); and the conifer Stachyotaxus is least common (2%).

8. Lunz plants in Waidhofen The Heimatmuseum in Waidhofen is now under reconstruction and I could see only four specimens from the Lunz beds: Equisetites and Pterophyllum from the private collection of F. Almer.

9. Lunz flora collection in the Botanical Institute of the University of Innsbruck There are ten specimens, eight of which have labels ‘Holzapfel H. GAMS’ with different years of collecting: 1920, 1929, 1933. They contain 33 imprints. Twelve of them belong to cycadophytes, thirteen of ferns, five of Glossophyllum, and three of Equisetites.

10. Lunz flora collection in Lyon The Geological Department at Lyon University possesses a smaller collection of Lunz plants, which had come from the Ecole de Mines in Paris. All specimens have a short address ‘Lunz’ and collector ‘De Ribera’. No other data are indicated. While in Lyon I looked through the whole collection and was allowed to take several specimens with me for a year to study them in Moscow. The collection consists of 100 imprints, the majority of which, as in the Vienna collection, belong to sterile leaves of Cycadophyta (Fig. 5). Ferns are

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Fig. 5. Relationships of main groups of fossil plants in the Lunz flora.

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second, as in the Vienna collection. Among the ferns, the most interesting is Bernoullia lunzensis. Its nervation is shown in Fig. 6. The Glossophyllaceae comes third. Sphenopsids and conifers are rare, and no Ginkgoites has been discovered. There is one fertile plant comparable to Pramelreuthia or Peltaspermum. It is not clear yet if it is a male or female strobil: it is necessary to try to extract spores from it. As in the Austrian collections, lycopsids, czekanowskiales and cycadocarpidiales are absent, as well as pteridosperms, unless the plant is not a pteridosperm, and if the Glossophyllaceae belongs to Gingkophyta. The short description of the Lyon collection follows. 10.1. Sphenopsids Sphenopsids are represented only by badly preserved remains of trunks. 10.2. Ferns Ferns consist of Clathropteris meniscioides Brongn. (Fig. 6A,B), Clathropteris lunzensis Stur (Fig. 6C), Danaeopsis lunzensis Stur (Fig. 6E), D. marantacea (Presl) Heer (Fig. 7A–K), Asterotheca marianii (Brongn.) Stur (Fig. 6D), and Bernoullia lunzensis Stur (Fig. 8A–C). The most interesting fern in the Lyon collection is that with the label ‘Bernoullia lunzensis Stur’ and which shows the venation perfectly. The genus Bernoullia was established by O. Heer on material from the Schilfsandstein of Basel. The brief description includes a very schematic picture. The pictures of venation in the present paper (Figs. 4 and 10A–C) give a more complete representation of this genus, but it is not enough for revision. It is very important to study it carefully because the genus Bernoullia was mentioned many times from Eurasia (Priuralye, China, Viet Nam) and comparison was made on the base of the poor illustration of the Basel specimen. It would be interesting to see if we have the same genus in western Europe and in other parts of Eurasia. 10.3. Cycadophytes Pterophyllum (Fig. 9A–J) consists of large leaves dissected in narrow segments. I think that in Stur’s

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list the quantity of species was exaggerated and all of them belong to one, two or maybe three species. The majority of them belong to P. filicoides (Schloth.) Thomas. Cuticles are preserved on many of them; epidermal study will help us to compare Lunz Pterophyllum with the same genus from Basel studied in detail by Kra¨usel and Schaarschmidt (1966). There is Ctenis lunzensis Stur (Fig. 8D,E) in the Lyon collection besides Pterophyllum and Taeniopteris. 10.4. Ginkgophytes? The remains of Glossophyllum (nearly 15%) are of great importance also. Unfortunately, whole leaves are practically absent and nervation hardly can be seen, especially near the base of the leaf. Cuticle preserved on leaves makes it possible not only to study the anatomy of leaves but also to clear up the systematic position of these plants and to compare them with similar plants from other regions. Again, the question is raised if Ginkgophyta were present or not in the Lunz flora. F. Krasser considered them not to be present; he thought that Clathrophyllum lunzense belongs to Cordaitales. R. Kra¨usel redetermined this latter species as Ginkgoites lunzensis and found also the leaves (previously determined by Krasser as Noeggerathiopsis sp. and by himself as Baiera lunzensis), which he considered to be a new genus, Glossophyllum. Kra¨usel attributed both of them to Ginkgophyta. Kra¨usel considered Glossophyllum to be Ginkgophyta because of two vascular bundles in the base of the leaf. Ginkgoites lunzensis and Sphenobaiera furcata seemed to him typical Ginkgophyta by their morphology. Tralau (1968) and Samylina (1970) established a special family, Glossophyllaceae. The present author follows this point of view (Dobruskina, 1980). Meyen (1984) included Glossophyllum with related plants in the order Peltaspermales. Unfortunately, I have not seen in the Lyon collection Sphenobaiera-like or Ginkgoites-like leaves. And I have no complete specimens of Glossophyllum here (Fig. 9K,L, Fig. 10); I have brought with me only pieces of cuticle of this genus. That is why in the present paper I give only preliminary data on the epidermal structure of Glossophyllum with some illustrations. As may be seen in Figs. 11–17, leaves have stomata on both sides of the cuticle. There are 30–50 stomata per mm2 on the lower side and 13–20

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Fig. 6. (A, B) Clathropteris meniscioides Brongn., Ft. 27, Ft. 23. (C) Clathropteris lunzensis Stur, Ft. 38. (D) Asterotheca merianii (Brongn.) Stur, Ft. 27-3, ð0.82, 65. (E) Danaeopsis lunzensis Stur, specimen Ft. 13-6. (F–H) Strobil of peltaspermous pteridosperm (?), Ft. 23-6: ð0.82, ð1.64, ð0.82, ð1.64, ð2.46, ð2.46, ð2.46.

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Fig. 7. (A–E) Danaeopsis marantacea (Presl) Heer, Ft. 13 (scale see on plate). (F–K) Danaeopsis marantacea (Presl) Heer, Ft. 14, ð0.82.

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Fig. 8. (A–C) Bernoullia lunzensis Stur, Ft. 29, ð0.83, ð2.49, ð2.49. (D, E) Ctenis lunzensis Stur, Ft. 23, ð0.74 (scale see on plate).

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Fig. 9. (A–J) Pterophyllum filicoides (Schloth.) Thomas, Ft. 27-4, Ft. 13-5, Ft. 38-1, Ft. 14, Ft. 13-3, Ft. 13-2, Ft. 38-4, Ft. 38-2, Ft. 38-3. (K, L) Glossophyllum florinii Kra¨usel, Ft. 38-5, Ft. 27-6, 7.

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Fig. 10. (A–I) Glossophyllum florinii Kra¨usel, resin-bodies, Ft. 2, ð82.

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Fig. 11. (A–D) Glossophyllum florinii Kra¨usel, lower cuticle, Ft. 10, ð82. (E) Glossophyllum florinii Kra¨usel, lower cuticle, Ft. 2, ð82.

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Fig. 12. (A–D) Glossophyllum florinii Kra¨usel, upper cuticle, Ft. 10, ð85. (E) Glossophyllum florinii Kra¨usel, upper cuticle, Ft. 2, ð85.

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Fig. 13. (A–D) Glossophyllum florinii Kra¨usel, upper cuticle, Ft. 48, ð100.

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Fig. 14. (A–D) Glossophyllum florinii Kra¨usel, lower cuticle, Ft. 48, ð82. (E) Glossophyllum florinii Kra¨usel, lower cuticle. Ft. 2, ð82.

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Fig. 15. (A–D) Glossophyllum florinii Kra¨usel, upper cuticle, Ft. 13-9, ð100.

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Fig. 16. (A–D) Glossophyllum florinii Kra¨usel, lower cuticle, Ft. 13-9, ð100.

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Fig. 17. (A–D) Glossophyllum florinii Kra¨usel: (A, B) upper cuticle; (C, D) lower cuticle, Ft. 38-7, ð100.

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to 40 (rarely to 60 stomata) per mm2 on the upper side. In contrast to this observation, R. Kra¨usel noted that the side with the best orientation of epidermal cells (lower side?) had less stomata. The greatest density of stomata (after Kra¨usel) was in the middle part of leaf; their quantity becomes less upward and downward. The lower cuticle consists of elongated epidermal cells, often arranged in longitudinal rows parallel to veins; typically they are polygonal, but sometimes roughly square. The latter ones may occur from near the petiole. Average dimensions of epidermal cells: 30 ð 50 µm, maximal D 60 ð 50 µm, minimal D 10 ð 10 µm. Stomata are typically arranged in rows and files; between them there are files without stomata. Such alteration of stomatal zones and zones without stomata is not always present and the orientation is strongly marked only in the middle part of the leaf. Walls of epidermal cells are straight; there are often papillae in the centre of the cell (after Kra¨usel, the greatest density of central papillae is in the middle of the leaf and they are absent in the upper part); the whole upper surface of the cell is often cutinized. The upper cuticle consists typically of polygonal isodiametrical epidermal cells without orientation with irregularly distributed stomata. After Kra¨usel, stomata are more numerous near the margin of the leaf. Sometimes epidermal cells are square and a bit elongated. Walls are straight; papillae in the middle of the cells are round, and they are less common there than on the lower cuticle and smaller. Secretory canals are rare on both sides, but during maceration of specimen FT-2 resin bodies are isolated (Figs. 11 and 12D,E). Stomata have one ring of 5–7 subsidiary cells smaller (typically 20 ð 30 µm, max. 20 ð 30 µm, min. 10 ð 15 µm) than ordinary epidermal cells and more cutinized; sometimes the upper surface is cutinized completely, sometimes it has papillae in the centre of the cell or near its margin. Stomata are of round or oval-round form, but sometimes their outline is uneven, while the whole form is round. After Kra¨usel’s description, there were 2–4 polar subsidiary cells with other subsidiary cells situated at sides of the stomatal pit. But in our material, distinguishing polar and side subsidiary cells may be done very rarely. As a whole, stomata on the upper cuticle are larger.

Guard cells 20–40 µm, more rarely to 60 µm, tend to be parallel to veins in nearly half of the material, the majority of them not being sunken. Differences in individual specimens consist of the degree of orientation of epidermal cells, degree of cutinization of subsidiary cells, and quantity of sunken guard cells. In the Triassic deposits of Eurasia, many Glossophyllum-like leaves (Cordaites-like leaves) were described (Dobruskina, 1980, p. 93). Among them there are few in which the cuticle was studied. Principles of determinations of such leaves were discussed earlier (Dobruskina, 1982). Differences in the epidermal structure of Glossophyllum florinii and Desmiophyllum imhoffii were already discussed by Kra¨usel (1943); he indicated the difference in structure and distribution of stomata and in the absence of papillae; Yuccites hadroclada differs as well. Only in this species of Cordaites-like leaves from the European Triassic was the cuticle described. Many leaves very similar to Glossophyllum are preserved in the Triassic of Priural’ye (Dobruskina, 1982, figs. 10–12). Previously they were assigned to Glossophyllum (Neuburg, 1959) or Yuccites (Brick, 1952). However, the present author considered differences in cuticle to be of generic level and proposed to distinguish an independent genus, Maria (Dobruskina, 1980). The genus Maria has the same morphology as Glossophyllum but differs in the absence of orientation of epidermal cells along the elongated leaf both in the lower and upper cuticle, and in the absence of stomatal rows and fills by the greater role of papillae in the centre of epidermal cells, by greater similarity of the subsidiary and ordinary epidermal cells (although the former are smaller, as in Glossophyllum), by proximal cutinization of subsidiary cells forming ‘stars’ or ‘rings’ and by sunken guard cells. At the same time, we have in the Triassic of Eurasia leaves with other morphologies (Sphenobaiera-type and Ginkgoites-type), with cuticles more similar to Glossophyllum. In fossil plants from the Pechora basin, I have found great similarity in cuticle structure in the leaves from the Pechora (genus Kalantarium Dobruskina), Lunz (both Glossophyllum florinii and Ginkgoites lunzensis), Basel (Sphenobaiera furcata), Svalbard (Glossophyllum) and the Tunguska basin (Kirjamkenia lobata, Sphenobaiera porrecta, S. vittaefolia, Glossophyllum claviforma).

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Prynada (1970) was the first to note the peculiarity of such cuticle. He had described Sphenobaiera-like leaves from the Tunguska basin under the generic name Kirjamkenia and considered them to be pteridosperms rather than ginkgophytes. His paper (published after his death) was prepared in the forties and he did not know the paper of Kra¨usel with the description of Glossophyllum. Similarity in the cuticle structure of Kirjamkenia and Kalantarium on the one hand and Glossophyllum on the other consists of: (1) orientation of cells on lower cuticle; (2) the presence of stomatal zones alternating with zones without stomata on the lower cuticle; on the upper cuticle this feature is absent in Kalantarium and Kirjamkenia and is less marked in Glossophyllum; (3) the presence of central papillae in Glossophyllum in one species of Kalantarium and sometimes in Kirjamkenia; (4) the difference in size and cutinization of subsidiary cells and epidermal cells; the former are smaller and constitute a ring around the stomatal pit; (5) unsunken guard cells which tend to be parallel to veins; and (6) approximately the same density of stomata (30–50 and 13– 20 to 40 per mm2 in Glossophyllum, 36–50 and 8–10 per mm2 in Kalantarium). The genus Kirjamkenia according to the descriptions of Prynada (1970) and Sadovnikov (1983) differs from Glossophyllum in the polygonal form of the epidermal cells on the upper cuticle, by the absence of regularity in the distribution of stomata (stomatal files can be seen sometimes on the lower side of the leaf, but stomata themselves have no orientation), and by the presence of proximal papillae on subsidiary cells (papillae at the end of a cell which typically overlaps the stomatal pit). The genus Kalantarium (Dobruskina, 1980) differs in the absence of stomatal files on the upper cuticle, in the absence of central papillae at epidermal cells (except one species), in the greater difference between ordinary epidermal cells and subsidiary cells (the latter are small, have a bracket shape and constitute a narrow ring around the stomatal pit), by proximal papillae on subsidiary cells, and by rare sunken guard cells. I have paid attention (Dobruskina, 1980) to the fact that in three different regions (Central Europe, Petchora, Tunguska) we have the same association of morphological types of leaves (Glossophyllum-like

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and Sphenobaiera-like) which have the same and peculiar type of cuticle. In the genus Kalantarium, I have united both types of leaves; in Lunz and in the Tunguska basin they were described under different generic names. I think that such associations indicate that in these regions we have related plants or maybe a single plant had leaves of two types. For practical purposes I would not like to change the generic names and would prefer to keep the original names for each region. But the name Sphenobaiera is used for very different plants that have similar morphology and very different cuticle and associated fructifications. I think that for the Palaeozoic and early Mesozoic this genus may be considered to be formal and must be revised. As concerns this group of plants, I think that the plants with Sphenobaiera-type morphology and Glossophyllum-type of cuticle would be better referred to the genus Kirjamkenia. Earlier I proposed to include these plants in the family Glossophyllaceae Tralaee (Dobruskina, 1980, 1982). Its volume and systematic position is discussed below. Here I would like to mention that in Lyon a collection of female fructifications is present (Figs. 5 and 8F–H) which will be described later. This seems to be a new genus which also may belong to the family Glossophyllaceae. 10.5. Conifers I have seen one small twig which may be considered a conifer (Stachyotaxus?).

11. Lunz flora collection in Meiningen, Germany I have been able to study this collection thanks to Dr. H. Kozur, who sent me photos of Lunz plants stored in the museum of Meiningen. I have 33 photos, among which are Pterophyllum (23 spec.), Asterotheca (6 spec.), Neocalamites (3 spec.), and Stachyotaxus (1 spec.) (Fig. 5).

12. History of Lunz flora studies W. Haidinger was the first to collect fossil plants from Lunz sandstones in 1842. The first localities

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were Tonibaueralpen near Wienerbruckl and Gaming near Lunz (see Fig. 2). Ettingshausen (1851), in his ‘Flora der Vorwelt,’ gave a first picture of Equisetites gamingianus from Gaming. W. Haidinger was right to compare them with the Keuper plants of Wu¨rfejberg (German basin). Unfortunately, later palaeobotanists, H.R. Goppert and F. Unger, referred them to the Lias due to the disorder in fossil plant collections: in one collection there were ones not only from the above localities but also from Grossau, Hinterberg and Pechgraben. It was D. Stur who explained the difference between the two groups of plants, having first studied the rocks in the museum. Later he confirmed his results by fieldwork in 1863– 1864. Present investigations are in agreement with Stur’s conclusions. In a separate paper, Stur (1885) gave a full list of fossil plants determined by him from the Lunz sandstone (Table 3). The same paper contained also a list of fossil plants from the Aon Schiefer of Raibl also determined by him. Stur compared these two floras and explained the difference in their composition not only by age (upper-middle Carnian in the first case, and lower Carnian in the second), but by the difference in ecology. That difference was explained by the different relationship to the Tethys basin. He thought that the Raibl flora was the flora of limestone reefs southward of the Central Range, which could have been subjected to storms of the open sea; the Lunz flora was the flora of a peat swamp at the southern margin of the Bohemian Highland north of the Central Range. Coeval vegetation of north Switzerland (Basel environs) was in the same position as the Lunz one. The second of Stur’s papers on the Lunz flora (Stur, 1888) was provoked by a publication by Fontaine (1883) on the Triassic plants of Virginia, USA. Stur found great similarity in the Lunz and Virginia floras (maybe somewhat exaggerated). He received some American specimens from Fontaine and had the possibility to compare imprints — not only figures and descriptions. He recognized the priority of Fontaine’s names for similar plants because the American plants were described and figured according to the rules, while he gave only a preliminary list of determinations for the Lunz flora. Stur indicated synonyms of the Alpine plants in the Virginia flora, and later F. Krasser gave new names to the

Alpine plants in agreement with Stur’s comments (Table 3). Unfortunately, this was all that Stur did with the Lunz flora. There are two collections of the Lunz flora in the Museum of the Geologische Bundesanstalt in Vienna with labels written by Stur; most of these labels have already been destroyed. These collections in the GBA are arranged in correspondence with two papers by Krasser (1909a,b); in the first one, Krasser diagnosed new taxa of marratiacean ferns introduced by Stur; in the second, he gave short descriptions of ferns and some other plants for which Stur’s names had remained ‘nomina nuda’; therefore, he dealt with these two papers before finishing his own investigations on Lunz Cycadophyta. Unfortunately, none of these papers have figures of or references to concrete specimens. It is not clear what specific specimens Krasser bore in mind while describing plants: the arrangement of collections was made later and not by Krasser. Later, Krasser published two papers (Krasser, 1917, 1919) with the descriptions and good figures of fertile regions of Lunz Cycadophyta (Table 4); besides, we can find one figure of Lunz Williamsonia in the paper ‘Williamsonia in Sardinien’ (Krasser, 1912). G. Nathorst, who received specimens of Lunz ferns from Haberfelner for the Stockholm Museum, extracted spores from sporangia of Asterotheca mariani and Danaeopsis lunzensis and published their photos (Nathorst, 1908). In another paper (Nathorst, 1907), he mentioned the identity of Camptopteris lunzensis and C. spiralis (not giving either a description or figure of the Lunz fern). Kra¨usel began to study Lunz plants in the twenties. In 1922, he described Baiera lunzensis (Stur) Kra¨usel, Macrotaeniopteris simplex Stur and Pterophyllum longifolium Heer, using collections from Berlin (Kra¨usel, 1922). In 1923, he published photos of cuticle in Baiera lunzensis from the Senckenbergmuseum in Frankfurt. In the forties, after 20 years of interruption, he revised part of these determinations (Tables 4 and 5). In the forties, Langer (1943, 1945) put the collections of the Lunz flora in the Geologische Bundesanstalt in order. Judging by his short papers he did enormous work. But he published the results in brief and without figures: only the most indispensable data on Stur’s ‘originals’. First of all, he showed that the concept ‘original’ can hardly be applied to

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Table 3 Lunz plants after preliminary determinations of Stur (1885), diagnoses and short descriptions of Krasser (1909a,b) and their synonyms in Virginia flora Stur (1885) 31 a 32 33 34 35 36 37 38 39 40

Calamites merianii Brongn. Equietum arenaceum Jaeger i E. lunzense Stur E. gamingianum Ett. sp. E. aratum Stur E. majus Stur E. haidingeri Stur E. neuberi Stur E. aequale Stur E. constrictum Stur

1 2 3 4 5 6

Coniopteris lunzensis Stur Speirocarpus pusillus Stur S. haberfelneri Stur S. neuberi Stur S. aurieulactus Stur S. lunzensis Stur

7

S. dentiger Stur S. microphyllus b Stur Oligocarpia lunzensis Stur O. robustior Stur

8 9 12 10 11 13 14

Asterotheca meriani Bgt. sp. A. lacera Stur c A. intermedia Stur d Bernoullia lunzensis Stur Danaeopsis lunzensis Stur e

15 22 23 24

D. marantacea Presl sp. Laccopteris lunzensis Stur Clathropteris lunzensis Stur C. reticulata Kurr m.

25 26 30 16 19 18 20 21 47 28 29 42 43 44 45 46

C. repanda Stur Thaumatopteris lunzensis Stur Camptopteris lunzensis Stur Taeniopteris simplex Stur T. lattior Stur T. angustior Stur T. haidingeri Geopp MS nec. Ett. T. lunzensis Stur T. parvula Stur Ctenis lunzensis Stur C. angustior Stur Pterophyllum pichleri Stur P. lunzense Stur P. gumbeli Stur P. haueri Stur P. neuberi Stur

Stur (1885): synonyms in Virginia Font. i

Schizoneura virginensis Equisetum rogersi Schimper

Krasser (1909a,b) (b) Neocalamites merianii (Brongn.) Halle (b) Equisetites arenaceus (Jaeg.) (b) E. gamingianus (Ett.) (b) E. laevis=E. platyodon

Lonchopteris virginensis Font. i

Aerostichides rhombifolius Font. f,i (may be part of it is a linneaefolius Bunb. sp.) A. microphyllus Font. Mertensides distans Font. i Mertensides bullatus Bunb. sp. i Asterocarpus virginensis Font. g,i A. platyrrachis Font. i A. penticarpus Font. i Pseudodanaeopsis reticulata Font. h,i

Clathropteris platyphylla var. expansa Font. i

Macrotaeniopteris magnifolia Roger sp. Macrotaeniopteris magnifolia Roger sp.

Ctenophyllum grandifolium Font. i

(b) E. (Equisetostchys) suecicus (Nath.) Halle (a,b) Coniopteris lunzensis Stur (a,b) Oligocarpia distans (Font.) Stur (a,b) Speirocarpus virginensis (Font.) Stur (a,b) S. neuberi Stur n. sp. (a,b) S. auriculatus Stur n. sp. (a,b) Stenuifolius (Emmons) Krasser

(a,b) Coniopteris lunzensis Stur n. sp. (a,b) Oligocarpia distans (Font.) Stur (a,b) O. bullata (Bunb.) Stur (a,b) O. coriacea Stur n. sp.-MS (a,b) Asterotheca meriani (Brongn.) Stur (a,b) Bernoullia lunzensis Stur n. sp. (a,b) Pseudodanaeopsis plana (Emmons) Font. (a,b) P. morantacea (Presl) Krasser (b) Laccopteris lunzensis Stur (b) Clathropteris lunzensis Stur (b) C. reticulata Kurr (b) C. repanda Stur (b) Thaumatopteris lunzensis Stur (b) Camptopteris lunzensis Stur (a) Macrotaeriopteris simplex Krasser n. sp. (a) M. lattior Krasser n. sp. (a) M. angustior Krasser n. sp. (a) M. haidingeri Krasser n. sp. (a) M. lunzensis Stur (a) M. simplex and M. angustior (b) Ctenis lunzensis Stur (b) did not find an original o (b) P. pichleri Stur A (b) P. lunzense Stur ) (b) P. gumbeli Stur (b) P. haueri Stur C (b) P. grandifolium (Font.) Krasser

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Table 3 (continued) Stur (1885)

Stur (1885): synonyms in Virginia

47 48 49 50 51 52 53 54 55 56

P. rectum Stur P. cf. pulchellum Heer P. cteniforme Stur P. haberfelneri Stur P. brevipenne Kurr P. longifolium Jaeg. P. macrophyllum Kurr P. approximatum Stur P. pectiniforme Stur P. riegeri Stur

57 41 27

P. irregulare Stur Dioonites cf. pennaeformis Schenk Clathrophyllum lunzense Stur

ex gr. P. longifolium ?

58

Pterophyllum ? lipoldi Stur

Palissya braunii Emmons

Krasser (1909a,b) (b) P. longifolium Jaeg.

Ctenophyllum taxinum Font. small leaves of P. longifolium P. inaequale Font.; P. decussatuni Emm. ex gr. P. longifolium ? ex gr. P. longifolium ? Ctenophyllum brauniannin Font. i

(b) after Fontaine D P. meriani Heer (b) P. longifolium Jaeg. (b) P. brevipenne Kurr (b) P. longifolium Jaeg. (b) P. longifolium Jaeg. (b) P. longifolium Jaeg. (b) P. riegeri Stur (b) P. jaegeri Brongn. (b) P. taxinum (L. et H.) Goepp (b) Nilssonia sturi Krasser n. sp. (b) ‘Dioonites cf. pennaeformis Schenk’ (b) Clathrophyllum lunzense Stur (b) Noeggerathiopsis sp. (b) Palissya lipoldi

o 9 > > > > > = > > > > > ;

B

B

o oD B

a Ordinal

number in Stur’s list. This species was mentioned only in Stur (1888). c Stur (1888) considered the upper part of Asterotheca meriani. d Stur (1888) considered as the middle part of A. meriani. e Stur (1888) called this species Heeria lunzensis Stur. f Aerostichides rhombifolius Font., 1883 coincides with Odontopteris tenuifolius Emmons 1857, i.e. is to be called Aerostichides tenuifolius (Emmons) Font. [Krasser, 1909a, p. 7]. g Asterocarpus virginensis Font., 1883 coincides with Peeopteris falcatus Emmons 1836, i.e. is to be called Asterocarpus falcatus (Emmons) Font. [Krasser, 1909a, p. 7]. h Pseudodanaeopsis reticulata Font., 1883 coincides with Stangerites plana Emmons 1857, i.e. is to be called Pseudodanaeopsis plana (Emmons) Font. [Krasser, 1909a, p. 7]. i D. Stur had received Virginian specimens from W. Fontaine (others he had seen only in Fontaine’s paper). b

Stur’s species due to the absence of descriptions. Even Krasser’s descriptions cannot help because he did not indicate concrete specimens. Now, 40 years later, after numerous movements of collections, it is impossible to outline the correspondence of descriptions to the concrete specimens. He underlined that the originals of Krasser’s Cycadophyta were real originals. J. Langer mentioned Kra¨usel’s work of the twenties and works by K. Frentzen and R. von Lilienstern on the German Triassic floras; in these works, the Lunz plants were mentioned for comparison. Langer found nearly all Krasser’s originals (Krasser, 1912, 1917, 1919) and paid special attention to those originals which he thought to be lost or nearly destroyed. He tried to find other specimens of the same species in GBA or other collections. He discussed several species. (1) In connection with Clathrophyllum lunzensis Stur he mentioned Kra¨usel’s papers in which this

species was referred to Ginkgoites lunzensis, while describing similar specimens from Berlin and Frankfurt. To be more correct, in the twenties Kra¨usel used the name Baiera lunzensis for such plants; in the forties, after Florin’s epidermal investigations, he referred them to Ginkgoites lunzensis (Table 5). (2) In 1943, J. Langer suggested that the original of Pseudoptilophyllum titzei was lost, but in 1945 he found that ‘Dioonites cf. pennaeformis (Schenk) Stur’ (with the label of R. Kra¨usel, not D. Stur) is the original of Krasser’s Pseudoptilophyllum titzei (microsporophylle), as well as the specimen with Stur’s label ‘Cycadeen Frucht-Hu¨llblatter’. J. Langer followed R. von Lilienstern, who considered these plants Dioonitocarpidium pennaeforme (Schenk) Ruhle von Lilienstern. Later, Kra¨usel referred it to a new species of the same genus: Dioonitocarpidium keuperianum (Krasser) Kra¨usel. (3) J. Langer considered Weltrichia keuperiana

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Table 4 Described and figured Lunz Cycadophyta Krasser, 1912, 1917, 1919; Kra¨usel, 1922 (1920); Florin, 1933; Langer, 1943, 1945

Langer, 1943, 1945; Kra¨usel, 1943, 1948 Kra¨usel, 1949, 1953; Kra¨usel and Schaarschmidt, 1966; Townrow, 1962; Potonie, 1962; Klaus, 1966, 1986; Meyen, 1983, 1984, 1987; Gomankov and Meyen, 1986 Taeniopteris haidingeri Goepp. K1949: XVI-5, Senckenberg

Taeniopteris sp. (cuticle only) K1922 (1920): XI-4,5, Berlin

Taeniopteris sp. (cuticle only) KS1966: XI-6,7 Stockholm

Macrotaeniopteris simplex Krasser K1922 (1920): X-1,2,3, textf. 5 (cuticle) K1922 (1920): IX-3–6; X-4,5; XI-6 (with cuticle and spores), Berlin

Leguminanthus siliguosus (Leuth.) Kra¨usel et Schaar KS1966: p. 41

Pterophyllum longifolium Brongn. K1922 (1920): IX-6b, XI-1,2,3 (with cuticle), Berlin Pterophyllum longifolium Heer P1921: Fig. 220

Pterophyllum longifolium K1986: Fig. 153 (after Kollmann)

Pterophyllum haberfelneri Stur P1897: p. 112 Pterophyllum sp. K1943: V-9, MW Lunzia austriaca Krasser K1917: I–VI, GBA 1917=1=1,2,3,4,11 with cuticle and spores

Lunzia austriaca Krasser P1962: XVI-440a,b (after Krasser)

Haitingeria krasseri (Schuster) K1919: I, GBA=Cycadospadis sp. In Krasser, 1910, C. krasseri Schuster in Schuster 1911=with cuticle

Haitingeria krasseri (Schuster) Krasser K1949: IX-8,9; X-8–11; XI-1–4; Texttab. 5–8, GBA (D Krasser), MW, Graz, Stockholm, Prague with cuticle and spores KS1966: IX-6 Stockholm P1962: XVII-466 (after Krasser) K1966: Fig. 160 (after Krasser) Westersheimia pramelreuthensis Krasser L1943: without figures: distorted specimen of Krasser C 2 others C 1 in Cambridge K1949: XVI; XVII; XVIII; Texttab. 14–16 GBA (D Krasser), Senckenberg with cuticle

Westersheimia pramelreuthensis Krasser K1917: Textf. 1 GBA

Williamsonia juvenilis Krasser K1917: without figures GBA

Bennetticarpus sp. K1949: XV-1,2 GBA

Williamsonia wettsteinii Krasser K1912: II-9 GBA=Williamsonia sp. in Krasser 1910=

Bennetticarpus wettsteinii (Krasser) Kra¨usel K1949: XI–XIII; Textf. 9–11 GBA (D Krasser), MW, Senckenberg with cuticle K1966: Abb. 155 (after Krasser)

F1933: Textab. 2 (cuticle) ? Williamsonia wettsteinii Krasser K1917: without figures GBA Weltrichia ? keuperiana Krasser K1917: IV-5 GBA 1917=1=13 L1943: considered as lost L1945: has found the original and two other specimens D Dioonitocarpidium pennoeforme (Schenk) R. v. Lilienstern

Cycadolepis wettsteinii Kra¨usel K1949: XIV–XVI; Textab. 12–13, GBA, MW, Stuttgart with cuticle

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Table 4 (continued) Krasser, 1912, 1917, 1919; Kra¨usel, 1922 (1920); Florin, 1933; Langer, 1943, 1945

Langer, 1943, 1945; Kra¨usel, 1943, 1948 Kra¨usel, 1949, 1953; Kra¨usel and Schaarschmidt, 1966; Townrow, 1962; Potonie, 1962; Klaus, 1966, 1986; Meyen, 1983, 1984, 1987; Gomankov and Meyen, 1986

Pseudoptiloptilophyllum titzei Krasser K1917: IV-6 GBA 1917=1=12 L1943: considered as lost but showed the similarity to ‘Dioonites cf. pennaeformis’ (Schenk) Stur L1945: considered the specimen with a label ‘Dioonites cf. pennaeformis’ as Stur’s and Krasser’s original. The label was written by Kra¨usel, not Stur

Dioonitocarpidium keuperianum (Krasser) Kra¨usel K1949: IX-5–7; X-1–6; Textf. 4 GBA 1917=1=13,12, MW, BW K1953: I-1,2 Stockholm

Dioonites cf. pennaeformis (Schenk) Stur L1943: Dioonitocarpidium pennaeforme (Schenk) R.v. Lilienstern L1945: considered this specimen as original of Pseudoptilophyllum titzei Krasser (the label was written by Kra¨usel) – Dioonitocarpidium pennaeforme (Schenk) R.v. Lilienstern ‘Cycadeen-Frucht-Hullblatter’ (a label of Stur) L1945: Dioonitocarpidium pennaeforme (Schenk) R.v. Lilienstern

Pramelreuthia haberfelneri Krasser K1917: I-5,6 GBA

Pramelreuthia II L1943: without figures, GBA

Dioonitocarpidium liliensterni Kra¨usel K1953: I-34 Stockholm K1949: XVII-5,7; XVIII-4–10; Textab. 17, 18, GBA (D Krasser) with spores T1962: 7 a–d (spores?) P1962: XV-441, 442 (after Kra¨usel and Townrow ?) K1966: without figures (spores) K1986: Abb. 160 (after Kra¨usel) Sturiella langeri Kra¨usel K1948: Fig. 1–7, GBA Sturianthus langeri K1986: Fig. 154 (after Kra¨usel, 1948)

Discostrobus treitlii Krasser K1917: I–II Antholithus wettsteinii Krasser K1917: I-7; IV-1 BW (Schrambach)

Antholithus wettsteinii Krasser K1943: VI-11; XI-6, XIII-6,7: Textab. 8 (right) BW (D Krasser), MW with cuticle and spores L1943: type of Stachyotaxus, GBA: Kra¨usel considers as Antholithus wettsteinii of bed preservation K1949: without figures P1962: XVII-477 (spores after Kra¨usel?) M1983: after Kra¨usel M1984: Fig. 16-14 after Kra¨usel, considered to be Antevsia GM1986: Fig. 18-14 after Kra¨usel, new combination Antevsia wettsteinii (Krasser) S. Meyen M1987: Fig. 49-14 after Kra¨usel, considered to be Antevsia Leguminethus siliquosus (Leuth.) Kra¨usel et Schaarschmidt KS1966: XI-3,5; XII, XIII-1,2,5,10,11; XV, GBA, MW, PW, Senckenberg, Basel, with cuticle and spores

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Table 5 Described and figured Lunz Ginkgophyta and Desmiophyllum imhoffi Clathrophyllum lunzense Stur Krasser (1909b): without figure Langer (1943): considered as lost

Baiera lunzensis (Stur) Kra¨usel Kra¨usel (1922) (1920): IX-1,2,7; X-6-8; text 1-3, Berlin (RB) with cuticle

Ginkgoites lunzensis (Stur) Florin Kra¨usel (1943): I-1-8; II-1-8; IV-1-5 GBA; 1942=1-6; Berlin (RB) with cuticle Klaus (1986): Abb. 151 (after Kra¨usel) Ginkgoites sp. Kra¨usel (1943): I-9,10; GBA: 1942=1=7 Kra¨usel (1949): VI-2-3; Textf. 1

Noeggerathiopsis sp. Krasser (1909b): without figure

Baiera lunzensis (Stur) Kra¨usel Kra¨usel (1923): III-1; IV-2 Senckenberg cuticle only

Glossophyllum florini Kra¨usel Kra¨usel (1943): II-9-11; III-6-10; IV-1-9; V; VI-1-10; VII; VIII; IX; X; text: 1-5; GBA: 1942=1=13-25; MW: Senckenberg; Berlin (RB), with cuticle Klaus (1986): Abb. 151 (after Kra¨usel) Desmiophyllum im hoffi Florin Kra¨usel (1943): IV-10,11

to be a protective leaf of the fertile part of the same plant, though not the microsporophyle itself. In 1943, he considered this original to be lost, though in 1945 he reported finding not only the original but two more specimens. (4) Besides the original of Westersheimia pramelreuthensis, Langer also saw two other specimens of this species and recalled that another one had been lent to Dr. H. Thomas in Cambridge. (5) Clathropteris lunzensis was represented in the GBA by a single specimen. But Langer reported that some specimens were present in Berlin collections (after Gothan). Langer considered it impossible to distinguish three species of Clathropteris, as in Stur’s list. In addition, J. Langer described, without figures, two fertile specimens that F. Krasser had not seen. The first one he named ‘Pramelreuthis II’. It was later named by R. Kra¨usel Sturiella langeri. The second one remained the Langer type of Stachyotaxus, but he mentioned that R. Kra¨usel had considered it Antholithus wettsteinii badly preserved. R. Florin’s works on the epidermis of fossil Ginkgophyta, Cycadophyta and Coniferophyta had a great influence on Kra¨usel’s investigations at the second stage of his work on the Lunz flora (in the forties). Florin (1933), himself, described and figured one picture of cuticle of Lunz Williamsonia wettsteinii (later redescribed by Kra¨usel as Bennet-

ticarpus wettsteinii). In the forties, R. Kra¨usel investigated the Lunz plants in Vienna, especially in GBA; the important part of his work was the study of Krasser’s originals. In GBA as well as in the Naturhistorical Museum in Vienna he has found a great number of long leaves which were only briefly mentioned by Krasser (1909b) as Noeggerrathiopsis sp. R. Kra¨usel investigated their cuticle and referred them to Ginkgophyta, distinguishing a new genus Glossophyllum (Kra¨usel, 1943). In the same work he described uncommon Ginkgoites lunzensis and included in this species (partly) Baiera lunzensis from his earlier papers (defined in the lists of Stur and Krasser as Clathrophyllum lunzensis) (Table 4). R. Kra¨usel in the same publication inferred Antholithus wettsteinii redescribed by him to be the male fructification of Ginkgoites lunzensis because the latter’s pollen grains discovered on the cuticle were identical to those extracted from Antholithus wettsteinii. Other investigations of Kra¨usel on the Lunz plants were performed on the Cycadophyta and Coniferophyta. In 1948, he described the new genus Sturiella (earlier Pramelreuthia II of Langer in GBA), which he referred to Bennettitales. He described Dioonitocarpidium after collections in GBA and the Naturhistorical Museum in Vienna (1949) and Stockholm (1953). He assumed the Lunz species to be different from the German one. He described Lunz cycadales as D. keuperianum (Krasser) Kra¨usel and

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Table 6 Described and figured Lunz Coniferophyta

Table 7 Described and figured Lunz ferns

Stachyotaxus lipoldi (Stur) Kra¨usel Kra¨usel (1949): VI, VII, VIII, IX-1, Textab. 2, 3 (Pterophyllum ? lipoldi in Stur (1885), Palissya lipoldi in Stur (1888) and Krasser (1909b)) GBA, MW, Senckenberg, Stockholm with cuticle and spores Potonie (1962): IX-523a,b: spores (after Kra¨usel) Klaus (1966): spores without figures Klaus (1986): Abb. 159 (after Kra¨usel)

Asterotheca merianii (Brongn.) Stur Nathorst (1908): I-10-12 (spores only); Stockholm Bhardwaj and Singh (1956): (leaves and spores), coll. of author Potonie (1962): IX-213, 214a,b (spores after Nathorst)

Stachyotaxus sp. Kra¨usel (1949): IX-2,3,4; GBA, MW

D. liliensternii Kra¨usel. He included a synonymy of the former Weltrichia keuperiana Krasser and Pseudoptilophyllum titzei Krasser. R. Kra¨usel (1949) redescribed Krasser’s Williamsonia as Bennetticarpus and Cycadolepis (Table 4) Together with F. Schaarschmidt, he defined in 1966 a new genus, Leguminanthus with a species L. siliqueosus after collections in Vienna (RW, MW, PW), Frankfurt and Basel. One of the specimens from the paper of 1922 (1920) earlier designated as Macrotaeniopteris simplex was included in this species. From collections in Vienna (RW, MW), Strasbourg and Frankfurt, Kra¨usel (1949) separated leaves which were earlier unconfidently defined as Pterophyllum and Palissya and ascribed them to the conifer Stachyotaxus lipoldi (Table 6). In the fifties, Bhardwaj and Singh (1956) described a fern, Asterotheca merianii, with its spores in situ based on their own material (Table 7). Roselt (1954), studying German sphenopsids, described Lunz Neocalamites merianii, with one illustration (Table 8). In the sixties, ‘in situ’ spores investigation was performed by Townrow (1962): Pramelreuthia haberfelneri (pictures published); and by Klaus (1966): Pramelreuthia haberfelneri and Stachyotaxus lipoldi (without figures). Klaus compared the spores of the first species with the dispersed spore species Lueckisporites junior Klaus and the second with Decussatisporites martini (Leschik). R. Potonie’s synopsis ‘Sporae in situ’ (1962) included such species as Asterotheca meranii, Danaeopsis lunzensis (after Nathorst), Pramelreuthia haberfelneri (after Kra¨usel and Townrow), Antholithus

Asterotheca-Synangien Klaus (1986): Abb. 145 Danaeopsis lunzensis Stur Nathorst (1908): I-8 (spores only); Stockholm Potonie (1962): IX-224 (spores after Nathorst) Klaus (1986): Abb. 146 (leaf) Clathropteris lunzensis Stur Langer (1943): without figures (fertile leaf) Klaus (1986): Abb. 1476 (after Stur) Clathropteris repanda Potonie (1897–1898): p. 112 Clathropteris sp. Potonie (1921): fig. 27 Camptopteris lunzensis Stur Nathorst (1907): mentioned that it coincides completely with C. spiralis Nath. Thaumatopteris lunzensis Stur Nathorst (1907): mentioned while describing T. schenkii

wettsteinii (after Kra¨usel), and Leguminanthus siliquosus (after Kra¨usel and Schaarschmidt). Figures of the Lunz plants are also presented in reviews and textbooks. H. Potonie’s textbook of palaeobotany contains pictures of Clathropteris repanda (1897–1898), Clathropteris sp. and Pterophyllum longifolium (1921). Klaus’ textbooks (Klaus, 1986, 1987) describes the Lunz flora as the stage in the development of fossil floras and presents illustrations of Asterotheca synangien, Danaeopsis lunzensis, Clathropteris lunzensis (after Stur), Ginkgoites lunzensis (reconstruction after Kra¨usel), Glossophyllum florini (after Kra¨usel), Table 8 Described and figured Lunz sphenopsids Equisetites gamingianus Ettingsh. Ettingshausen (1851): 90, VIII-2; Graz Neocalamites merianii Roselt (1954): V-1

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Pterophyllum longifolium (after H. Kollmann), Sturianthus langeri, Bennetticarpus wettsteinii, Pramelreuthia haberfelneri, and Stachyotaxus lipoldi (after Kra¨usel). Meyen (1983) produced a picture of Antholithus wettsteini Krasser (after Kra¨usel), erroneously connecting it (following Krassilov, 1972) with Glossophyllum florini (Kra¨usel connected it with Ginkgoites lunzensis). The microsporophyle of Antholithus wettsteini and microsporophyle of Sphenobaiera furcata are referred to the genus Antevsia in Meyen’s review (Meyen, 1984, fig. 16), in ‘Tatarina flora’ (Gomankov and Meyen, 1986, fig. 18) and in ‘Fundamentals of palaeobotany’ (Meyen, 1987, fig. 44). The combination Antevsia wettsteinii (Krasser) S. Meyen was proposed by him in ‘Tatarina flora’ (Gomankov and Meyen, 1986, p. 50). He later inferred the association of A. wettsteinii, Glossophyllum florini and Sphenobaiera lunzensis from Lunz with S. furcata and Leuthardtia ovata Kra¨usel et Schaar. from Basel. This connection is evidenced by the similarity of epidermal structure of Glossophyllum florini, Sphenobaiera furcata and S. lunzensis (after Dobruskina, 1980, 1982) and the similarity of microsporophyles (after Meyen). Considering these microsporophyles as Antevsia, Meyen ascribed the family Glossophyllaceae to Peltaspermales.

Table 9 Papers with descriptions and figures (C), only descriptions ( ), repeated figures after original works (?) and discussion or mention (without mark) of the Lunz plants C Bhardwaj and Singh (1956) Depape and Doubinger (1963) Dobruskina (1980) Dobruskina (1982) C Ettingshausen (1851) C Florin (1933) Florin (1936) Frentzen (1922) ? Gomankov and Meyen (1986) Halle (1908) Klaus (1966) ? Klaus (1986) ? Klaus (1987)

C C C C C C C C C

13. Present state of studies of the Lunz flora There are more than 30 papers which deal with the Lunz plants one way or another (Table 9). Nine papers with illustrations (Krasser, 1917, 1919; Kra¨usel, 1922 (1920),Kra¨usel, 1923, 1943, 1948, 1949, 1953; Bhardwaj and Singh, 1956) and five without illustrations (Krasser, 1909a,b; Langer, 1943, 1945; Klaus, 1966) are devoted just to the Lunz plants. Six other papers present original drawings or photos of a single Lunz plant, its cuticle or its spores along with other fossil plants (Ettingshausen, 1851; Nathorst, 1908; Schuster, 1910–1911; Krasser, 1912; Florin, 1933; Townrow, 1962). In ten papers the Lunz plants are mentioned to compare or to identify them with early known species or to discuss their systematic position (Nathorst, 1907; Halle, 1908; Frentzen, 1922; von Lilienstern, 1928; Schuster, 1932; Florin, 1936; Meyen, 1983, 1984, 1987; Gomankov and Meyen,

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? ? ? C ? ? ? C

C

Krasser (1909a) Krasser (1909b) Krasser (1912) Krasser (1917) Krasser (1919) Kra¨usel (1922) (1920) Kra¨usel (1923) Kra¨usel (1943) Kra¨usel (1948) Kra¨usel (1949) Kra¨usel (1953) Langer (1943) Langer (1945) von Lilienstern (1928) Meyen (1983) Meyen (1984) Meyen (1987) Nathorst (1907) Nathorst (1908) Potonie (1897–1898) Potonie (1921) Potonie (1962) Roselt (1954) Schuster (1910–1911) Schuster (1932) Townrow (1962)

Asterotheca merianii Noeggerathiopsis sp. Glossophyllaceae All list of species Equisetites gamingianus Williamsonia wettsteinii (cuticle) Ginkgoites lunzensis Dioonites pennaeformis Anthiolithus wettsteinii Equisetites majus two species of spores and pollen 12 figures in textbook Spores in situ, two species (fig. to 1966) diagnoses of ferns, 20 species short descriptions of 47 species Williamsonia wettsteinii 9 species of Cycadophyta 1 species of Cycadophyta Ginkgophyta and Cycadophyta, 4 species Ginkgophyta, cuticle of 1 species Ginkgophyta, 4 species Sturiella langeri Cycadophyta, 8 species, 1 conifer Dioonitocarpidium, 2 species 6 original of Krasser, 2 new 2 originals of Krasser Dioonites pennaeformis Antholithus wettsteinii Antholithus wettsteinii Antholithus wettsteinii Camptopteris, Thaumatopteris spores of ferns, 2 species Clathropteris repanda Clathropteris sp., Pterophyllum 6 species of spores in situ Neocalamites meriani Cycadopspadix krasseri 4 species of Cycadophyta spores of Pramelreuthia

1986). Finally, several reviews and textbooks on palaeobotany contain figures of the Lunz plants or their spores reproduced from the original works (Potonie, 1897–1898; Potonie, 1921; Potonie, 1962; Klaus, 1986).

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Table 10 Described and figured Lunz plants Equisetites gamingianus Neocalamites merianii Asterotheca merianii Danaeopsis lunzensis Clathropteris lunzensis C. repanda Clathropteris sp. Taeniopteris haidingeri Taeniopteris sp. Pterophyllum longifolium P. haberfelneri Pterophyllum sp. Macrotaeniopteris simplex Lunzia austriaca Haitingeria krasseri Westersheimia pramelreuthensis Bennetticarpus wettsteinii Bennetticarpus sp. Cycadolepis wettsteinii Dioonitocarpidium keuperianum D. liliensternii Pramelreuthia haberfelneri Sturiella langeri Antholithus wettsteinii Discostrobus treitlii Leguminanthus siliquosus Ginkgoites lunzensis Ginkgoites sp. Glossophyllum florinii Desmiophyllum imhoffii Stachyotaxus lipoldi Stachyotaxus sp.

E 1851 R 1954 N 1908, BS 1956, P 1962, K 1986 N 1908, P 1962, K 1986 L 1943, K 1986 P 1897 P 1921 K 1949 K 1922 (1920), KS 1966 K 1922 (1920), P 1921, K 1986 P 1897 K 1943 K 1922 (1920) K 1917, P 1962 K 1917, K 1919, K 1949, P 1962, KS 1966, K 1986, S 1911 (Cycadospadix krasseri) K 1917, L 1943, K 1949 K 1949, K 1912 (Williamsonia wettsteinii), F 1933 (Williamsonia wettsteinii) K 1949, K 1917 (?Williamsonia juvenieis) K 1949, K 1917 (?Williamsonia wettsteinii) K 1949, K 1953, K 1917 (Weltrichia keuperiana and Pseudoptilophyllum titzei), L 1943 (Dioonitocarpidium pennaeforme), L 1945 (D. pennaeformae) K 1953 K 1917, K 1949, T 1962, P 1962, K 1966, K 1986 K 1948, L 1943 (‘Pramelreuthia II’) K 1917, K 1943, L 1943, K 1949, P 1962, M 1983, M 1984 (Antevsia wettsteinii), GM 1986 (A. wettsteinii), M 1987 (A. wettsteinii) K 1917 KS 1966, K 1922 (1920): Macrotaeniopteris simplex K 1943, K 1949, K 1986, K 1922 (1920): Baiera lunzensis, L 1943 (Clathrophyllum lunzense) K 1943, K 1949 K 1943, K 1949, K 1986, K 1923 (Baiera lunzensis), K 1909b (Noeggerathiopsis sp.) K 1943 K 1949, K 1966, K 1986, K 1909b (Palissya lipoldi) K 1949

The resulting list of published (described and figured) Lunz species is shown in Table 10. It is interesting to compare it with Table 3, which gives the list of all defined plants (except the new species of Krasser and Kra¨usel distinguished later). Attention in publications was mostly focused on the Cycadophyta, especially on their fertile parts. The same specimens were studied by F. Krasser, R. Kra¨usel, J. Langer, J. Townrow, and W. Klaus. For example, there are more than four publications on Pramelreuthia, and more than three on Antholithus wettsteinii, etc. It seems that all fertile Cycadophyta were studied and reported in papers, while numerous sterile leaves of the Cycadophyta were mentioned and illustrated in no more than five figures. They are

evidently monotonous and less interesting from the botanical point of view, but their diversity is clearly illustrated by Stur’s list (Table 3). GBA collections present more than 1395 samples of such leaves. Only one Taeniopteris leaf and four Pterophyllum leaves were figured in different papers, including textbooks (Table 4). The second place in publications belongs to the Ginkgophyta. In this case it cannot be said that all specimens were studied and illustrated; maybe all Ginkgoites but not all Glossophyllum: they are more numerous, and a large part of them is figured (Table 5). Among numerous ferns (see also Table 3) only one species is described in detail and figured on the

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basis of the Bhardwaj and Singh collection. At the same time, the bulk of material in the GBA was not studied after Stur and Krasser, who did not illustrate any ferns. The same with the sphenopsids: two species were published based on Graz and German material. A huge body of GBA data was only mentioned in short notes of Krasser. Occasional conifers (one species, Stachyotaxus lipoldi) are represented in publications quite satisfactorily. Table 3 presents the taxonomic diversity of fossil plants in Lunz but does not show quantities of each group. Quantitative relations of the main groups of plant may be seen in Fig. 5. The most numerous specimens are the sterile leaves of Cycadophyta, the second place belongs to ferns, and the third to Sphenopsida. Ginkgophyta and Coniferophyta are few. All these papers contain illustrations of about 190 Lunz specimens; some 90 of them belong to the GBA collections. The first two papers of F. Krasser as well as two short papers of J. Langer give no pictures at all. Taking into account that the Geologische Bundesanstalt collection alone comprises more than 2600 specimens, it is clear that only a small part of the Lunz collection is published. It is true that the most interesting species are published in the above-mentioned papers, as well as most of the reproductive organs of Cycadophyta. The unpublished species are mostly sterile parts of similar leaves. This means that the ratio of published and unpublished taxa is much less than is inferred from formal estimates. Some groups, however, are almost absent from illustrated publications (for instance, ferns), others need revision, and the interrelation of sterile and fertile parts is often unclear. This means that the above publications fail to produce the general picture of the flora, to say nothing of the relations of different plant groups in the Lunz flora.

14. Problems of systematics and taxonomy Because the Lunz flora studies are very disproportional, it is worth giving a short review of its systematic composition with an account of controversies and unsolved problems. Lycopsida are absent from the Lunz flora. Sphenopsida are represented by two genera (Neocalamites and Equisetites) that are widespread in

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the Triassic of Eurasia. They are very monotonous throughout all Eurasia. At first glance the Lunz genera do not differ from Middle Asian ones. The number of recognized species is controversial. It is necessary to study the Lunz sphenopsids at the species level. Two species were published (Table 8): the picture of E. gamingianus Ett. was the first illustration of a Lunz fossil plant in 1851 and the second, Neocalamites merianii, was reported exactly a century later, in 1954. Ferns are abundant, all featuring great fronds, often fertile, very diverse. Most of them belong to Marrattiaceae and Dipteridaceae. Because the majority of the Lunz ferns are not published, it is difficult to determine their role in the Lunz flora from publications. For example, papers treat only two parts of Asterotheca fronds, whereas the collections represent very abundant and large fronds. Dipteridaceae are also very abundant and have large fronds. These were only occasionally pictured in the textbooks on palaeobotany. They were not given due place in the Carnian floras; it seemed that the abundance of Dipteridaceae evidently occurred later, in the Norian, while in dispersed spectra they were abundant already in the Carnian. Thus, the Lunz ferns, especially Dipteridaceae need a most comprehensive study. The Lunz ferns were discussed more than once because D. Stur had read the paper by Fontaine (1883) on the Triassic Virginia flora after his publication in 1885. The Virginia flora is mostly represented by ferns. Stur inferred them to be very similar or even identical with Lunz ferns. His investigation of Virginia rock material received from W. Fontaine permitted him to indicate a striking resemblance of rocks and fossils, and he gave a list of the Lunz plants synonymous in the Virginia flora. After Stur’s death, F. Krasser brought to order Stur’s notes and published the diagnoses and short descriptions of the Lunz ferns (Krasser, 1909a,b), in accordance with Stur’s point of view and rules of priority (see Table 3 and notices of Krasser in a footnote). However, these papers lack figures and references to particular specimens either. The Lunz and Virginia plants similarity does not seem to me so evident, identity at the species level seems to be absent and the generic identity has yet to be proved. It may be achieved only through monographic study.

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Only Asterotheca merianii was studied monographically (Table 7) by Bhardwaj and Singh (1956), with illustrations of leaves and spores ‘in situ’. The picture of Asterotheca synangia from Lunz is given by Klaus (1986) in his textbook. Spores from Asterotheca merianii and Danaeopsis lunzensis sporangia were extracted by A.G. Nathorst and sketched in 1908. The sketch was included in R. Potonie’s ‘Synopsis of sporae in situ’. Danaeopsis lunzensis and Clathropteris lunzensis fronds were pictured in the ‘Einfu¨hrung in die Pala¨obotanik’ by Klaus (1986) and those of Clathropteris sp. in H. Potonie’s (Potonie, 1921) ‘Lehrerbuch der Pala¨obotanik’. While studying sphenopsids we are faced with the problem of revision of their specific composition; addressing ferns calls for revision of their generic composition and belonging to one or another family, the studies of pteridosperms bring forth the problem of whether this group exists at all in the Lunz flora. The publications on the Lunz flora do not mention this group, but Lunz Glossophyllaceae may belong to pteridosperms. The present paper refers this family to Ginkgophyta with some degree of uncertainty. Some reproductive organs earlier referred to Cycadophyta or unstudied remains may also belong to pteridosperms. Even in this case, only several specimens are to be included in this group. Rather infrequent ‘Fertile regions of Cycadophyta’ are the most interesting part of the collection from a botanical point of view. It was only natural that nearly all palaeobotanists who performed research on the Lunz flora tried to study them. Curiously, there are more than eleven genera of fertile parts that correspond to two genera for sterile leaves: Pterophyllum and Taeniopteris (Macrotaeniopteris) (Table 4). It is worth remembering that in the time of Krasser and Kra¨usel, Taeniopteris or Macrotaeniopteris was ascribed to ferns. These two genera are represented in GBA by more than 500 sterile leaves. Fertile parts are represented by 179 specimens, the majority of which is described. Only Westersheimia showed the direct attachment of sporophylles to the stem: the connection with sterile leaves can only be supposed. It was attributed to the Cycadophyta (the most part to Bennettitales) based on botanical considerations. The Lunz Bennettitales are the earliest representatives of this group. The systematic position of these organs remains controversial up to the

present. Seven special monographs were published on the Lunz Cycadophyta, and the problem is usually treated by all palaeobotanical reviews. The systematic position of Lunz ‘Fertile regions of Cycadophyta’ is highly debatable. For example, F. Krasser considered Haitingeria to be a female organ, but later R. Kra¨usel obtained some pollen grains from it. Antholithus wettsteinii was assigned by F. Krasser to Cycadophyta, by R. Kra¨usel to Ginkgophyta, and by S.V. Meyen to pteridosperms. There are different views on the position of Pramelreuthia and Bennetticarpus, etc. As far as I know, no species or genus of fertile Lunz Cycadophyta were discovered in any other flora except Antholithus and Leguminanthus (in Basel). The abundance of fertile parts itself is an unusual feature of the Lunz flora. Ginkgophyta are represented in the Lunz flora by the family Glossophyllaceae (Glossophyllum florinii and Ginkgoites lunzensis). It is, however, problematic whether this family belongs to Ginkgoales or to Peltaspermales (pteridosperms). Tralau (1968) established for Glossophyllum a special family of Ginkgoales because it differed from other representatives of Ginkgoales (Ginkgoaceae and Sphenobaieraceae). Due to the similarity of the epidermis of Glossophyllum florinii, Ginkgoites lunzensis and S. furcata as well as some Pechora and Siberia species (see also description of Glossophyllum in Section 4), I proposed to widen the notion of the family Glosophyllaceae by including in it Glossophyllum florinii from Lunz, G. spetsbergense Vasilevskaja, Glossophyllum sp. from Svalbard, G. claviforme Mogutcheva from the Tunguska basin, Kalantarium krauselii Dobruskina, K. prosundum Dobruskina from Petchora, and Kirjamkenia lobata from Prynada, including in it Sphenobaiera vittaefolia Mogutcheva and S. porrecta Mogutcheva from Tunguska, Kirjamkenia? lunzensis (Stur) Dobruskina from Lunz, and K.? furcata (Kra¨usel) Dobruskina from Basel (Dobruskina, 1980, p. 112). This means that I referred to the genus Kirjamkenia two ‘Sphenobaiera’ species from the Tunguska basin and (with question mark) also Ginkgoites lunzensis Kra¨usel from Lunz and Sphenobaiera furcata Kra¨usel from Basel. Later, while discussing the volume of the family Glossophyllaceae I preferred to leave original generic names: Glossophyllum florinii Kra¨usel, Sphenobaiera lunzensis (Stur) Dobrusk., Sphenobaiera furcata (Heer)

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Florin, Antholithus wettsteinii Krasser, Kirjamkenia lobata Pryn., Sphenobaiera porrecta Mogutch., S. vittaefolia Mogutch., Kalantarium kraeuselii Dobrusk., and K. prosundum Dobrusk. (Dobruskina, 1982, p. 66). But, here I consider Ginkgoites lunzensis Stur as Sphenobaiera. Originally it was determined by Stur as Clathrophyllum lunzensis Stur, then redetermined by Kra¨usel (1922) as Baiera lunzensis (Stur) Kra¨usel and in 1943 as Ginkgoites lunzensis (Stur) Florin (Table 5). The final redeterminations may be made with assurance only after detailed revision of Lunz and Basel plants of this type as well as of other representatives of this family. In any case, independent of the correct generic determinations, all these plants are very similar and belong to the family Glossophyllaceae. In epidermal structure, all these plants share features with the pteridosperms. V.D. Prynada was the first to speak about it while studying Kirjamkenia from Siberia in the forties. But, they differ from the studied Ginkgoaceae and Spenobaieraceae as well as from the studied pteridosperms (Dobruskina, 1980). As for fertile parts that may be connected with Glossophyllaceae there are Antholithes wettsteinii Krasser from Lunz and the structure on the twig of Sphenobaiera furcata from Basel. These two, Kra¨usel considered as male fructifications of Ginkgophyta. Only Sphenobaiera furcata has a microsporophyle (of Anthevsia-type) in direct attachment to a leaf. Female fructifications of Leuthardtia crassa Mogutcheva (1973) were found in association with Kirjamkenia in the Tunguska basin. Kra¨usel (1943) related Antholithus wettsteinii to Ginkgoites lunzensis (not with Glossophyllum florinii as was mentioned later) because the pollen grains extracted from Antholithus wettsteinii are of the same type he had found with the Ginkgoites lunzensis cuticle. One (of two) imprint of Antholithus lies near the imprint of Glossophyllum; but the second one was found in Schrambach, a locality in which Glossophyllum is not known, as well as in all nearly localities (Lilienfeld group of localities and Kirchberg group). No pollen grains were found on Glossophyllum cuticle or on any other plant except Ginkgoites lunzensis. Meyen (1984) referred Antholithus wettsteinii Krasser and the microsporophylle of Sphenobaiera furcata to Antevsia. This gave him grounds to as-

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sign all Glossophyllaceae to Peltaspermales. But it seems now that stomata of the Lunz Glossophyllum when scanned are similar to Ginkgoales stomata (oral communication of V.A. Samylina). To my mind the question is debatable and I would not currently change the earlier accepted systematic position of this group. All the plants referred to the Glossophyllaceae have been collected in the Triassic: the Lower Triassic in the Tunguska basin, the Ladinian–Carnian of Priural’ye, and the Carnian of Svalbard, Lunz and Basel. I think that some plants of similar morphology but without cuticles may also belong to Glossophyllaceae. I mean the Ladinian–Carnian floras of the Donez basin, Middle Asia, Mongolia, China, and Soviet Primor’ye. From the geological distribution of Glossophyllaceae this family does not exist (or nearly does not exist) in floras younger than the Carnian; we must look for related forms in more ancient floras. I have written earlier that Glossophyllaceae are the palaeophytic element in the Triassic floras (Dobruskina, 1980). The real Ginkgophyta (fam. Sphenobaieraceae) appear in the Norian–Rhaetian. The conifers show only one species: Stachyotaxus lipoldii. About 15 leaves (Table 6) were illustrated; in collections their number now is 186. There were no controversies up to now concerning this species. Klaus (1966) extracted pollen grains from it. It is strange that the usual Triassic conifers, Podozamites, Cycadocarpidium, and Voltzia were not found in the Lunz flora, while they are abundant in nearly coeval floras. Thus, in different groups we are faced with quite different problems.

Acknowledgements This paper was originally completed and submitted to the Geologische Bundesanstatt, Vienna, in 1988.

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