+ MODEL

ARTICLE IN PRESS
PALBO-02802; No of Pages 21

Review of Palaeobotany and Palynology xx (2006) xxx – xxx

www.elsevier.com/locate/revpalbo

Morphology and epidermal anatomy of Nilssonia (cycadalean

foliage) from the Upper Triassic of Lunz (Lower Austria)
Christian Pott a,⁎, Hans Kerp a , Michael Krings b
a
Forschungsstelle für Paläobotanik am Geologisch-Paläontologischen Institut, Westfälische Wilhelms-Universität Münster, Hindenburgplatz 57,
D-48143 Münster, Germany
LMU
b
Bayerische Staatssammlung für Paläontologie und Geologie und GeoBio-Center , Richard-Wagner-Straβe 10, D-80333 Munich, Germany
Received 11 April 2006; received in revised form 12 July 2006; accepted 14 July 2006

Abstract

The Carnian flora from Lunz (Lower Austria) ranks among the richest and most diverse fossil floras from the Upper Triassic.
It is one of the first modern Triassic floras with bennettitaleans. Although this flora is often referred to in the literature, modern
taxonomic studies are mostly absent; only some of the reproductive structures have been studied in detail. Many of the plant
remains yield excellently preserved cuticles. During a systematic study of the Pterophyllum leaves from Lunz, it appeared that
several species previously accommodated in that taxon have to be transferred to other genera. This paper deals with four species
that are transferred to Nilssonia (cycadalean foliage); the macromorphology and epidermal anatomy are described and discussed.
The following new combination is introduced: Nilssonia riegeri nov. comb. Two new species are described (i.e. Nilssonia
lunzensis and Nilssonia neuberi) based on material originally mentioned in a species list as Ctenis lunzensis Stur nom. nud. and
Pterophyllum neuberi Stur nom. nud. The diagnosis for a fourth species, Nilssonia sturii Krasser, is emended. The Nilssonia
species from Lunz range among the earliest representatives of the genus Nilssonia.
© 2006 Elsevier B.V. All rights reserved.

Keywords: Nilssonia; cuticular analysis; Carnian; fossil cycads; Austria; palaeoecology

1. Introduction cycadalean from bennettitalean foliage is by cuticular

analysis (Florin, 1933; Watson and Sincock, 1992;
The very rich and diverse Carnian flora from Lunz Watson and Cusack, 2005). Thus far, no unequivocal
(Austria) is dominated by pinnate leaves, most of representatives of cycadalean foliage, recognized based
which are attributed to the Bennettitales. Only a few on cuticles, have been reported from Lunz, with the
fossils have been assigned to Ctenis Lindley et Hutton, exception of several cycadalean megasporophylls
a genus of cycadalean foliage (Stur, 1885); however, assigned to Dioonitocarpidium Rühle von Lilienstern
the cycadalean nature was presumed only based on by Kräusel (1953).
macromorphology. The best method to discriminate Stur (1871) was the first who mentioned pinnate
cycad-like foliage from Lunz, and in 1885, he published
⁎ Corresponding author. Tel.: +49 251 8323934; fax: +49 251 a long list of taxa. However, none of these two publi-
8321739. cations includes descriptions and illustrations. Stur's list
E-mail address: christian.pott@uni-muenster.de (C. Pott). (1885) includes 17 Pterophyllum species, 13 of which
0034-6667/$ - see front matter © 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.revpalbo.2006.07.007

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
2 C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx

are new names. Moreover, the list also contains two new The Lunz Formation (Lunzer Schichten) is subdi-
names for Ctenis species. All the new names introduced vided into three units (Verloop, 1908). The middle unit
by Stur (1885) are nomina nuda. Krasser (1909) pub- is named Lunzer Sandstein and consists of sandstones at
lished a classification of the taxa based on Stur's hand- the base overlain by marine marls gradually grading
written notes, giving short descriptions of the taxa upwards into terrestrial sands, shales and coal. The coal-
accompanied by a brief discussion. Because Krasser bearing part of the sequence is overlain by marls and
(1909) could not find the specimens originally assigned with a sandstone layer at the top of the subunit. The
to Ctenis by Stur, he did not deal with the material plant fossils occur in the shales associated with the coal
allocated to Ctenis. Krasser (1909) classified the Ptero- beds.
phyllum species into six groups, but his classification is The lower unit of the Lunzer Schichten (= Reingrab-
difficult to follow. Pterophyllum is now generally ac- ener Schiefer) is dated as Julian based on the occurrence
cepted as a type of bennettitalean foliage (e.g., Boyd, of ammonoids (Krystyn, 1978). Palynological studies
2000). Krasser (1909) was the first who reported Nils- indicate a Carnian (Bhardwaj and Singh, 1964) and a
sonia from the Lunz flora, by renaming Pterophyllum Julian age (Dunay and Fisher, 1978). The Opponitzer
irregulare Stur Nilssonia sturii Krasser. However, P. ir- Limestone, the upper subunit of the Lunzer Schichten
regulare (non P. irregulare Nathorst) is a nomen nudum. was dated as Tuvalian (Dunay and Fisher, 1978).
Krasser (1909) only gave a brief description of the Outcrops in the area around Lunz are today poor
species without illustrations and did not study cuticles. since coal mining finally ceased after World War II.
Although Pterophyllum leaves from Lunz are shown Spoil tips are not well accessible and most of the
in many museum exhibitions and are illustrated in many material is strongly weathered. The old specimens are
textbooks and popular articles (e.g., Potonié, 1899; stored in museum collections throughout Europe (see
Schmidt, 1928; Bachmayer and Kollmann, 1969; Turek Appendix).
et al., 1990; GBA, 2002), a modern revision of the genus Almost all remains are preserved as impressions or
that includes cuticular analyses is still absent. compressions, the latter often with excellently preserved
A new revision of the Lunz flora, focusing on foliage cuticles. Cuticles were prepared according to procedures
taxa, reveals that several species previously assigned to summarized in Kerp (1990), and Kerp and Krings
Pterophyllum actually represent cycadalean leaves. The (1999). Samples with plant remains were dissolved in
cycadalean nature of these fossils is evidenced by epi- hydrofluoric acid in order to obtain cuticles, or cuticles
dermal anatomy, notably the morphology of the stomatal were picked directly from the rock surface. Cuticles
complexes. Based on leaf morphology and epidermal were macerated according to the standard procedure
anatomy, these forms are accommodated in Nilssonia using Schulze's reagent (35% HNO3 with a few crystals
Brongniart. Four species are described here that repre- of KClO3) and 5–10% potassium hydroxide. Macerated
sent the first unequivocal cycadalean leaves from cuticles were washed in distilled water, gently dehy-
Lunz. drated in pure glycerine, and afterwards mounted in
permanent glycerine-jelly on microscope slides. Slides
2. Material and methods are stored in the collection of the Forschungsstelle für
Paläobotanik am Geologisch-Paläontologischen Institut,
The material studied comes from the Upper Triassic Universität Münster, Germany; accession numbers are
of Lunz in the Northern Calcareous Alps of Lower given in the figure captions.
Austria, approximately 100 km west of Vienna (Fig. 1). Hand specimens were photographs with a Nikon D
The material was collected during the late 19th and 100 digital camera; in order to increase contrast cross-
early 20th centuries in coalmines at several localities in polarization (i.e. polarized light sources with polarizing
the area around Lunz-am-See. With more than 30,000 filter over the camera lens) was used. Cuticles were
specimens, the Lunz flora is one of the richest and most analysed with a Leitz Diaplan light microscope equipped
diverse Late Triassic floras of the Northern Hemi- with a Nikon DS-5M digital camera.
sphere. The flora consists of sphenophytes, ferns, and
numerous gymnosperms (i.e., ?ginkgophytes, cycada- 3. Taxonomy
leans, bennettitaleans and coniferophytes); to date more
than 70 taxa have been reported. The most up-to-date Subdivision: Cycadophytina
overviews were published by Dobruskina (1989, 1998), Class: Cycadopsida
however, these papers are essentially based on older Order: Nilssoniales
publications. Genus: Nilssonia Brongniart (1825)

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx 3

Fig. 1. Position of Lunz-am-See in the Northern Calcareous Alps, Lower Austria.

Remark on the orthography of the generic name Nils- Repository: Palaeobotanical Collection of the Geolo-
sonia: Brongniart (1825) introduced the genus Nilsonia gische Bundesanstalt, Vienna.
for leaves from the Lower Jurassic of Scania (Sweden).
Brongniart (1825, p. 204), mentioned that the genus was Emended diagnosis
named after M. Nilson. Nathorst (1909) commented that Leaves petiolate, pinnate, lanceolate in outline, distally
the name was misspelled because the family name is tapering; lamina subdivided into parallel-sided to
Nilsson, and he corrected the generic name. The spelling tapering leaf segments, variable in width, with rounded
Nilsonia is a typographical error (Schweitzer et al., apices; oppositely to sub-oppositely positioned, inserted
2000). The spelling Nilssonia is today widely accepted to the adaxial (upper) side of the rachis, each with
in the literature (e.g., Harris, 1932, 1964; Schweitzer several unforked, delicate parallel veins. Leaves hypos-
et al., 2000; Watson and Cusack, 2005). tomatic with delicate cuticles, epidermal cells narrow,
polygonal to rectangular, anticlinal cell walls straight.
Nilssonia sturii Krasser (1909) emend. Stomatal apparati monocyclic and actinocytic, in
Plates I, II1–3 and III intercostal fields; guard cells with prominent dorsal
thickenings. Abaxial epidermis with short hollow
Selected references papillae.
1885 Pterophyllum irregulare Stur, Flora Lunzer-
Schichten, p. 99 (nom. nud.) Description
non 1879 Pterophyllum irregulare Nathorst, Om Floran Leaves are petiolate, pinnate (segmented), of almost
i Skånes kolförande Bildningar, p. 68, Plate XVII, Fig. 1 regular, oblong, more or less lanceolate shape, apex
1909 Pterophyllum irregulare Krasser, Kenntnis Flora unknown. Lamina subdivided into numerous, irregularly
Lunzer Schichten, pp. 116–120, no ill. oppositely positioned segments (Plate I1–8). Length of
1909 Nilssonia sturii Krasser, Kenntnis Flora Lunzer segments continuously decreases towards the apex of the
Schichten, pp. 120–121, no ill. leaf. Segments are clearly crescent-to sword-shaped, all
1917 Nilssonia sturii Seward, Fossil Plants, p. 576, no ill. of same shape, distally tapering, slightly widened at the
base. The width of the individual segments may vary
Lectotype: GBA 1909/003/0388 (Plate I4). considerably, some segments being twice as wide as
others (Plates I1–6 and II2–3). The distance between the
Comment: The name Nilssonia sturii was introduced by individual segments remains the same within a single
Krasser (1909) who gave a very brief diagnosis but did leaf, unless the segments are widened basally. Segments
not illustrate any specimens of the species. Because no are attached to the upper side of the rachis (Plates I5,9
holotype was designated, the specimen GBA 1909/003/ and II1,3).
0388 is here selected as the lectotype. This specimen Numerous parallel, non-bifurcated veins enter the seg-
belongs to the original set of fossils studied by Stur ments and run straight to the apex (Plate I5–9). In adaxial
(1885) and Krasser (1909); originally labelled Ptero- surface view, the prominent rachis is nearly completely
phyllum irregulare by Stur (1885). covered by the leaf segment bases (Plate I4–6, 8–9). The

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
4 C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx

leaf apex is not preserved. The proximal part of the leaf Material studied: Approximately 60 specimens from the
consists of a naked petiole, which is, however, rarely following collections: NHM, GBA, NRM, RUU, JOA,
preserved. The petiole may reach a length of up to SPO.
17.1 cm.
Incomplete leaves are up to 54.5 cm long and 26.2 cm Remarks
wide. The length of the leaf segments varies according The leaf remains described above are assigned to the
to their position within the leaf, the longest being up to genus Nilssonia because macromorphological features
13.2 cm. Within a single leaf, the width of segments may (e.g. narrow, nearly band-like or oblong-oval, irregularly
vary considerably, ranging from 6.7 mm up to 18.8 mm segmented lamina and segments, attached to the upper
at their base. side of the rachis and covering it) correspond well with
Leaves are hypostomatic and have delicate cuticles. those seen in typical representatives of this genus (Van
Costal and intercostals fields are distinguishable on the Konijnenburg-van Cittert et al., 2001; Watson and
abaxial (Plate III1) but not on the adaxial side of the leaf Cusack, 2005). Additional characters include a short
(Plate III2). distance between the segments, and a fine, parallel ve-
Adaxial cuticle — The cuticle of the adaxial side is very nation with unforked veins (Schimper and Schenk, 1890;
delicate and difficult to macerate. Epidermal cells are Nathorst, 1909; Harris, 1964). Leaf variability as shown
polygonal or rectangular in outline, elongate with acute by the Lunz specimens (Plates I1–2 and II1–2) is typical
or pointed ends, 35–80 μm long and 20–37.5 μm wide. for Nilssonia (Nathorst, 1909; Harris, 1932, 1964;
Anticlinal cell walls are smooth. Stomata are absent on Schweitzer et al., 2000).
the adaxial side, which does not show any other special The leaf morphology of the Lunz material is superficially
features, neither cuticular thickenings nor trichome similar to that of the Nilssonia tenuicaulis-complex, as
bases or papillae (Plate III2). has already been stated by Krasser (1909) and Seward
Abaxial cuticle — The abaxial cuticle is thin, but (1917). These authors, however, did not figure Lunz
slightly thicker than the adaxial cuticle. The epidermis material. The leaves from Lunz here described as Nils-
shows a clear differentiation into costal and intercostal sonia sturii are much larger than those of N. tenuicaulis
fields. Costal fields are composed of about 3 to 4 lines Seward, as was already mentioned by Krasser (1909).
of cells (Plate III1) without stomata and trichome According to Harris (1943), N. tenuicaulis leaves reach
bases; cells are narrow, rectangular, or polygonal to a maximum length of 20.0 cm, although he later states
isodiametric in outline, slightly elongate, acute, 25– that the leaves must have been larger (Harris, 1964).
62.5 μm long and 15–25 μm wide. Anticlinal cell walls Nevertheless, N. tenuicaulis is still smaller than N. sturii.
are straight and periclinal walls are smooth. Every Moreover, both species also differ considerably with
second or third cell bears a short, thick-walled, hollow regard to epidermal anatomy (cf. Dower et al., 2004).
papilla (20–25 μm in diameter) positioned at the end of The density of papillae in Nilssonia tenuicaulis is lower
the cell (Plate III1, 4, 8, 10). Intercostal fields are 100– than in Nilssonia sturii (Plate III1, 4); in N. tenuicaulis
150 μm wide; cells are polygonal and isodiametric to papillae occur only occasionally (Harris, 1943). Harris
broadly rectangular or slightly elongate (Plate III1, 4), (1943) also describes small resin bodies for N. tenui-
ranging from 15–40 μm long to 12.5–30 μm wide. caulis that do not occur in N. sturii.
Anticlinal cell walls are straight. Stomata are regularly Pterophyllum irregulare described and illustrated by
distributed but irregularly oriented within the intercos- Nathorst (1879) from the Rhaeto–Liassic of Bjuv
tal fields. Actinocytic stomatal apparati are mono- to (Sweden) is very different from Nilssonia sturii, in
diacyclic, 32.5–45 μm long and 15–22.5 μm wide, having more slender leaf segments; the name P. irre-
with 6 to 8 trapezoid to rectangular subsidiary cells gulare Stur is a nomen nudum, used for the species later
(Plate III3, 5–7, 9). The stomatal pores vary in length named N. sturii (Krasser, 1909).
from 10–17.5 μm. The kidney-shaped guard cells are Several of the specimens originally assigned to Ctenis
not sunken. The central parts of the dorsal walls are (and Ctenophyllum Schimper) by Stur (1885) and
strongly cutinized, whereas the ventral part and the Krasser (1909), kept in the collections of NHM, GBA
polar ends of the guard cells are weakly cutinized and NRM, can be identified as Nilssonia sturii. None of
(Plate III3, 5–7, 9). these specimens shows anastomosing veins, which is a
typical feature for Ctenis.
Locality: Lunz-am-See, Lower Austria.
Source Strata: Lunzer Schichten, Julian, middle Car- Nilssonia riegeri (Stur ex Krasser, 1909) nov. comb.
nian, Triassic. Plate II4–6

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx 5

Plate I (caption on page 8).

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
6 C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx

Plate II (caption on page 8).

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx 7

Plate III (caption on page 8).

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
8 C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx

Basionym: Pterophyllum riegeri, (Stur, 1888 nomen Comment: The name Pterophyllum riegeri was introduced
nudum) Krasser (1909) — Krasser, F., 1909. Zur by Stur (nomen nudum) and validated by Krasser (1909),
Kenntnis der fossilen Flora der Lunzer Schichten. who gave a very brief diagnosis but did not illustrate the
Jahrbuch der kaiserlich-königlich geologischen Reich- specimens. Because no holotype was designated, the
sanstalt 59, 1–26. specimen GBA 1909/003/0589 is here selected as the
lectotype. This specimen belongs to the original set of
References fossils studied by Stur (1885) and Krasser (1909);
1871 Pterophyllum riegeri Stur, Geologie der Steier- originally labelled Pterophyllum riegeri by Stur (1885).
mark, p. 250, (nom. nud.)
1885 Pterophyllum riegeri Stur, Flora Lunzer-Schich- Repository: Palaeobotanical Collection of the Geolo-
ten, p. 99, (nom. nud.) gische Bundesanstalt, Vienna.
1888 Pterophyllum riegeri Stur, Lunzer Flora in
Virginia“, p. 8, (nom. nud.) Diagnosis
1909 Pterophyllum riegeri Krasser, Kenntnis Flora Leaves slender pinnate, overall outline lanceolate;
Lunzer Schichten, pp. 119–121, no ill. lamina subdivided into narrow segments of equal
width, tapering towards their apex, apices rounded;
Lectotype: GBA 1909/003/0589 (Plate II5) leaf segments insert to the upper side of the rachis; each

Plate I. Nilssonia sturii Krasser (1909) emend. (see on page 4)

1. well preserved leaf with irregularly faced leaf segments (NHM, 1886/0001/0014), scale bar = 1 cm;
2. well preserved leaf displaying irregularly faced leaf segments (GBA, 1909/003/0400), scale bar = 2 cm;
3. well preserved leaf with well recognisable character of the leaf segments (NHM, 1884/D/1202), scale bar = 1 cm;
4. well preserved leaf showing different sizes of the leaf segments, lectotype, originally labelled Pterophyllum irregulare by Stur (1885)
(GBA, 1909/003/0388), scale bar = 1 cm;
5. detail of central part of a leaf with leaf segments inserted to the upper side of the rachis (NHM, 1886/0001/0014), scale bar = 1 cm;
6. apical part of a well preserved leaf with the leaf segments decreasing in length towards the leaf apex (NRM, S148681), scale bar = 1 cm;
7. central part of a large and well preserved leaf (GBA, 1909/003/0392), scale bar = 2 cm;
8. large and excellently preserved leaf portion (NHM, 2006B0008/0033), scale bar = 1 cm;
9. detail of a leaf with covered rachis and simple veins (NHM, 2006B0008/0012), scale bar = 2 mm.

Plate II. Nilssonia sturii Krasser (1909) emend. and Nilssonia riegeri (Stur ex Krasser, 1909) nov. comb. (see on page 5)

1. N. sturii, central part of a leaf with leaf segments inserted to the upper side of the rachis (NHM, 2006B0008/0033), scale bar = 5 mm;
2. N. sturii, apical part of an excellently preserved leaf with the leaf segments decreasing in length towards the leaf apex (NHM, 1885/D/
4027), scale bar = 1 cm;
3. N. sturii, central part of a leaf with leaf segments inserted to the upper side of the rachis (NRM, S148246), scale bar = 1 cm;
4. N. riegeri, central part of a leaf with several lanceolate leaf segments (GBA, 1909/003/0585), scale bar = 1 cm;
5. N. riegeri, excellently preserved leaf displaying the delicate architecture of this species, lectotype (GBA, 1909/003/0589), scale
bar = 1 cm;
6. N. riegeri, central part of a leaf with leaf segments inserted to the upper side of the rachis, lectotype (GBA, 1909/003/0589), scale
bar = 5 mm.

Plate III. Nilssonia sturii Krasser (1909) emend. (see on page 6)

1. abaxial cuticle, overview displaying cell pattern with short papillae and stomata (GBA, 1909/002/0518/0007), scale bar = 50 μm;
2. adaxial cuticle, overview of cell pattern (GBA, 1909/002/0518/0004), scale bar = 50 μm;
3. abaxial cuticle, stoma with strongly cutinized guard cells (GBA, 1909/002/0518/0007), scale bar = 50 μm;
4. abaxial cuticle, overview showing the arrangement of cells, stomata, and papillae (GBA, 1909/002/0518/0006), scale bar = 50 μm;
5. abaxial cuticle, haplocheilic stoma with strongly cutinized parts of the guard cells (GBA, 1909/002/0518/0006), scale bar = 20 μm;
6. abaxial cuticle, haplocheilic stoma with surrounding subsidiary cells forming an actinocytic stomatal apparatus (NHM, 2006B0008/0035/
0002), scale bar = 20 μm;
7. abaxial cuticle, haplocheilic stoma with surrounding subsidiary cells forming an actinocytic stomatal apparatus (NHM, 2006B0008/0035/
0001), scale bar = 10 μm;
8. abaxial cuticle, short, hollow, and strongly cutinized papilla (GBA, 1909/002/0518/0006), scale bar = 10 μm;
9. abaxial cuticle, haplocheilic stoma with radial striae on the walls of the guard cells (NHM, 2006B0008/0035/0001), scale bar = 10 μm;
10. abaxial cuticle, short, hollow, and strongly cutinized papilla (GBA, 1909/002/0518/0006), scale bar = 10 μm.

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx 9

segment with several simple, delicate parallel veins. of the dorsal walls are strongly cutinized, whereas their
Leaves amphistomatic with delicate cuticles; costal and ventral parts and the polar ends are only weakly cutinized.
intercostals fields distinguishable on the abaxial, but not
on the adaxial side of the leaf. Epidermal cells narrow, Locality: Lunz-am-See, Lower Austria.
polygonal to rectangular, cell walls straight; stomata Source Strata: Lunzer Schichten, Julian, middle Car-
haplocheilic, arranged in intercostal fields; guard cells nian, Triassic.
with prominent dorsal thickenings; abaxial epidermis Material studied: Five specimens from the following
with short hollow papillae. collections: GBA, NRM

Description Remarks
Leaves are oblong to lanceolate in outline, pinnate, leaf Krasser (1909) originally described the specimens
segments densely spaced. Lamina subdivided into studied here and gave a short diagnosis. Although he
numerous, irregularly faced, narrow and lanceolate noticed some characteristic features (insertion of the leaf
segments; (Plate II4–5). Segments attached to the upper segments, venation) that are typical for Nilssonia, he
side of the rachis. They cover most of the adaxial surface assigned the material to Pterophyllum riegeri, thereby
of the rachis (Plate II6). Individual segments gradually following Stur (1885). Cuticular analysis clearly demon-
decrease in length towards the apex of the leaf; individual strates that the specimens cannot be retained in Ptero-
segments taper towards their apex, resulting in an irregular phyllum, a genus for bennettitalean foliage, but rather
outline of the leaf. Leaf segments narrow, basally slightly belong to the Cycadales. Based on the pinnate nature of
expanded, apex rounded. Segments almost equal in width, the leaf, the insertion of the individual leaf segments, the
bent slightly towards the leaf apex, more than five times as venation pattern and the epidermal anatomy, this species
long as broad. Five to eight parallel, non-bifurcated veins is assigned to Nilssonia.
enter the segments at a somewhat acute angle (Plate II4– Cuticles of Nilssonia riegeri are very similar to those of
6). Petiole not preserved. Nilssonia sturii and it is difficult to distinguish both
Incomplete leaves are up to 16.0 cm long and 8.9 cm species based on the cuticles. However, N. riegeri
wide. The length of the leaf segments varies depending leaves are amphistomatic, whereas those of N. sturii are
on their position within the leaf; they are up to 5.2 cm hypostomatic. Moreover, the overall morphology of the
long; their width ranges from 1.7 mm to 3.3 mm. leaves is quite different in that N. riegeri leaves are
Leaves are amphistomatic with delicate cuticles. Costal much smaller and possess narrower leaf segments.
and intercostals fields are distinguishable on the abaxial, Nilssonia riegeri appears to be a relatively rare species.
but not on the adaxial side of the leaf. Only five specimens have been recorded to date.
Adaxial cuticle — Epidermal cells are polygonal or Unfortunately, not all of these specimens yield well
rectangular, elongate with acute or pointed ends. The preserved cuticles, because some are covered with var-
adaxial cuticle of Nilssonia riegeri is largely similar to nish and cuticles are strongly oxidized.
that of Nilssonia sturii, morphologically as well as with Nilssonia riegeri is quite similar to Nilssonia feriziensis
regard to cell sizes, but N. riegeri differs from N. sturii Fakhr from the Rhaeto–Jurassic flora of the Kerman
in having a few stomata on the adaxial side. Hollow Basin, Iran (Schweitzer et al., 2000), but much smaller;
papillae occur occasionally but in some instances may cuticles of the latter species are not known.
be reduced to thickenings of the periclinal walls.
Abaxial cuticle — Cells over veins are narrow, Nilssonia lunzensis Stur ex Pott, Kerp and Krings nov.
rectangular or polygonal to isodiametric in shape, slightly spec.
elongate, acute. Costal fields are composed of about 2 to 3 Plates IV and V1–2
lines of cells. The abaxial cuticle of Nilssonia riegeri is
largely similar to that of Nilssonia sturii, morphologically Selected references
as well as with regard to cell sizes. Periclinal walls are 1885 Ctenis lunzensis Stur, Flora Lunzer-Schichten, p.
smooth; short, thick-walled, hollow papillae-bearing 98 (nom. nud.)
cells are scattered regularly between the normal epider- 1885 Ctenis angustior Stur, Flora Lunzer-Schichten, p.
mal cells. The monocyclic stomata are regularly 98 (nom. nud.)
distributed within the intercostal fields and oriented 1909 Ctenis lunzensis Krasser, Kenntnis Flora Lunzer
irregularly. Stomata are surrounded by an actinocytic ring Schichten, p. 113 (nom. nud.)
of 6 to 8 trapezoid to rectangular subsidiary cells. The 1909 Ctenophyllum lunzense Krasser, Kenntnis Flora
kidney-shaped guard cells are surficial. The central parts Lunzer Schichten, p. 113 (nom. nud.)

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
10 C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx

Holotype: GBA 1909/003/0196 (Plate IV6) cutinized (Plate V3, 7, 11–12). The periclinal walls
Epitype: NRM, S148602 (Plate IV5) sometimes show faint longitudinal striae. Stomata are
confined to the intercostal fields, irregularly oriented, 55–
Repository: Palaeobotanical collection of the Geolo- 80 μm long and 22.5–32.5 μm wide; they are monocyclic
gische Bundesanstalt, Vienna with a ring of 6 to 7 polygonal subsidiary cells. The
stomata are slightly sunken, with pores of 15–22.5 μm in
Diagnosis length. Papillae and trichomes are absent (Plate V1, 4, 5).
Leaves imparipinnate, broad-lanceolate to acute-oval in Abaxial cuticle — Costal fields are composed of about 3 to
outline; leaf subdivided into tongue-shaped segments of 4 lines of cells. Epidermal cells are similar in outline and
unequal width. Leaf segments strongly decurrent at the size to those of the adaxial side; however, cutinisation of the
base, tapering towards the apices, apices rounded; leaf anticlinal walls is weaker. Cells are narrow, elongate,
segments inserted to the upper side of the rachis, several rectangular in outline, with straight anticlinal walls (Plate
delicate veins enter each segment and run straight towards V2, 4), 50–115 μm long and 20–40 μm wide. Periclinal
the apex. The leaf apex consists of a large terminal walls are normally smooth, but longitudinal striae may
segment, rhomboidal in outline. Leaves amphistomatic, occasionally be present. Intercostal fields are 200–225 μm
cuticles delicate. Epidermal cells elongate, rectangular to wide, cells rectangular to acutely ending, polygonal or
isodiametric in outline; anticlinal cell walls straight; isodiametric, irregularly oriented (Plate V2), 35–80 μm
stomata sunken; stomatal apparati monocyclic, arranged long and 17.5–45 μm wide. Anticlinal cell walls are
in intercostal fields. straight. Stomatal apparati irregularly oriented, monocyclic,
45–65 μm long and 20–35 μm wide, with 6 to 7 polygonal
Description subsidiary cells encircling the stomatal pit; stomata sunken
Leaves imparipinnate, individual segments attached to (Plate V6, 8–9, 10), stomatal pores 12.5–17.5 μm long.
the upper side of the rachis, strongly decurrent basisco- Papillae may occur in the proximal portion of the segments;
pically and tapering towards the apices, resulting in a trichome bases are absent.
rather open appearance of the leaf (Plate IV1–2, 4–9).
The unequal widths of the individual segments, some Locality: Lunz-am-See, Lower Austria.
twice as wide as the adjacent ones, create an irregular Source Strata: Lunzer Schichten, Julian, middle Car-
appearance (Plate IV1–2, 5–7, 9). The overall outline of nian, Triassic.
the leaf is oblong to pointed-oval. Segment length grad- Material studied: Approximately 30 specimens from the
ually decreases towards the leaf apex. The leaf apex following collections: NHM, GBA, NRM, RUU, JOA
consists of a large terminal segment that is rhomboidal in
outline (Plate IV5, arrow). Leaf segments are more than Remarks
twice as long as wide and bent towards the leaf apex. The The leaf morphology and epidermal anatomy, especially
apices of the individual segments are obtuse-rounded. the shape of the stomatal apparati, justify the accommo-
Numerous parallel veins enter the segments at an angle of dation of this species in the genus Nilssonia. Many of the
ca. 80°, and run straight towards the apex without bi- specimens included here in Nilssonia lunzensis were
furcating (Plate IV3, 8); each vein consists of two thin originally labelled as Ctenis lunzensis by Stur (1871,
vascular strands. The petiole is unknown. 1885) and Krasser (1909). However, this latter name is a
Incomplete leaves are up to 24.6 cm long and 13.9 cm nomen nudum because the species was never validly
wide. Individual leaf segments are up to 85 mm long and described. The assignment to Ctenis was based on the
16.8 to 42.1 mm wide; their length varies depending to presumed anastomosing venation (Krasser, 1909). How-
their position within the leaf. ever, careful re-examination of Stur's and Krasser's
Leaves are amphistomatic; cuticles are very delicate and original material revealed that the specimens do not show
difficult to macerate. A differentiation into costal and anastomosing veins. Some of the leaf segments were
intercostal fields occurs in the adaxial and abaxial epidermis. folded during embedding in the sediment and, as a result,
Adaxial cuticle — Cells over veins are elongate, leaf parts overlap each other (Plate IV3). The veins of the
rectangular to isodiametric in outline, occasionally ending underlying leaf portion are often still visible in the
acutely, 55–105 μm long and 25–37.5 μm wide. overlying leaf segment, which suggests the presence of
Anticlinal cell walls are smooth. Cells of intercostal fields anastomosing veins. Therefore, these anastomoses are
are isodiametric, polygonal to broadly rectangular in regarded as a taphonomic feature, which does not allow
outline, ending acutely, 45–95 μm long and 30–60 μm assignment of the material to the genus Ctenis. Several
wide. Anticlinal cell walls are smooth and heavily specimens from the collection of the Geologische

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx 11

Table 1
Macromorphological and epidermal features of the Nilssonia species from Lunz

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
12 C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx

Bundesanstalt in Vienna were labelled by Stur as Cteno- 1990 non Pterophyllum grandifolium Cornet and Olson,
phyllum lunzense. However, the genus Ctenophyllum is no Flora Richmond and Taylorsville Basins, p. 52, pl. 18.
longer used and, according to Seward (1917), is a synonym
of Pseudoctenis Seward. According to Krasser (1909), Holotype: GBA 1909/003/0576 (Plate VI1)
these are the specimens referred to as C. lunzensis by Stur Epitype: GBA 2006/004/0014 (Plate VI5)
in his 1885 paper. Ctenis differs from Nilssonia in having
laterally inserted leaf segments. Repository: Palaeobotanical collection of the Geolo-
Nilssonia lunzensis differs from the other Nilssonia gische Bundesanstalt, Vienna
species from Lunz in having more loosely arranged leaf
segments; a comparison of the Nilssonia species from Diagnosis
Lunz is presented in Table 1. Leaves large, robust, subdivided into long, parallel-sided,
A species closely resembling Nilssonia lunzensis is Nils- irregularly alternating segments with rounded apices.
sonia kendallii Harris from the Middle Jurassic of Leaf segments insert to the upper side of the rachis. Each
Yorkshire, UK (Harris, 1964). Although N. kendallii segment with several, non-bifurcated, parallel, delicate
shows a similar gross morphology and the cuticles also are veins. Leaves hypostomatic, cuticles thin. Epidermal
superficially similar, this species has much smaller leaves cells on both leaf sides elongate to rectangular with
and the resin bodies and papillae mentioned by Harris straight anticlinal cell walls. Epidermis of the abaxial side
(1964) for N. kendalii are missing in N. lunzensis. differentiated into costal and intercostals fields; stomata
Nilssonia acuminata Göppert from the Rhaeto–Liassic restricted to the intercostal fields. Stomatal apparati
of Franconia (Germany) is another species that shows haplocheilic; guard cells with prominent dorsal thicken-
some similarity to Nilssonia lunzensis (Schenk, 1867). ings and radial striae. Abaxial epidermis with a few short
However, leaves of this species apparently also have hollow papillae.
resin bodies, which were interpreted as sporangia by
Schenk (1867). Description
Leaves are large and robust, regularly pinnate; leaf
Nilssonia neuberi Stur ex Pott, Krings and Kerp nov. segments widely spaced (Plate VI1–3, 5); individual
spec. leaf segments slightly decurrent, insert to the upper side
Plate VI of the rachis, long and narrow, hardly tapering towards
their tips. The striate rachis is remarkably thin. Leaf
Nomenclatural remarks: The name Pterophyllum neu- petiole and apex unknown. Venation dense, consisting
beri, from which the specific epithet is derived, was of a large number of parallel, non-bifurcated veins that
introduced by Stur (1885) as a nomen nudum. Krasser enter the leaf segments at 90° angles (Plate VI5).
(1909) gave a very brief description and placed this Incomplete leaves are up to 52.5 cm long and 39.3 cm
species in the synonymy of Ctenophyllum grandifolium wide. The length of the leaf segments varies depending
Fontaine (1883), thereby transferring the latter species to on their position within the leaf; leaf segments may be up
Pterophyllum. Cornet and Olson (1990), and probably to 23.3 cm long and 12.3–26.4 mm wide.
not aware of Krasser's (1909) paper, again transferred C. The leaves are hypostomatic and have very delicate
grandifolium to Pterophyllum. Cornet and Olson (1990) cuticles. Costal and intercostals fields are distinguish-
refigured one of the very schematic drawings of Fontaine able only on the abaxial side.
(1883). Because Fontaine's illustrations are schematic Adaxial cuticle — Unfortunately, cuticles are rather poorly
and do not show essential details such as the venation, we preserved. Cells are rectangular, elongate, 67.5–142.5 μm
feel that a comparison of the Lunz material to Fontaine's long and 27.5–32.5 μm wide. Anticlinal cell walls are
specimens, which have never been illustrated photo- smooth as far as visible. Periclinal walls are smooth.
graphically, is not possible. Therefore, a new species Abaxial cuticle — Cells over the veins are narrow,
based on the material studied by Stur (1885) and Krasser rectangular or elongate to isodiametric in outline;
(1909) is formally proposed here. dimensions are difficult to ascertain, cells are about
20–30 μm wide (Plate VI6, 9). Costal fields are
Selected references composed of 9 to 10 lines of cells (Plate VI9). Anticlinal
1885 Pterophyllum neuberi Stur, Flora Lunzer-Schich- cell walls are straight, periclinal walls smooth, some
ten, p. 99, (nom. nud.) bearing a thick-walled hollow papilla. Intercostal fields
1909 Pterophyllum grandifolium Krasser, Kenntnis are 350–375 μm wide, individual cells not recognizable.
Flora Lunzer Schichten, p. 115–120, no ill. Stomata are regularly distributed in the intercostal fields,

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx 13

Plate IV (caption on page 16).

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
14 C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx

Plate V (caption on page 16).

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx 15

Plate VI (caption on page 16).

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
16 C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx

randomly oriented, monocyclic, 40–50 μm long and easily be distinguished from other Nilssonia species by its
17–25 μm wide (Plate VI4, 6–9). Stomata are slightly large dimensions (Plate VI1, 5).
sunken, guard cells with radial striae on the dorsal walls The material from Lunz originally labelled as Ptero-
(Plate VI4, 8). Dorsal walls are strongly cutinized in the phyllum neuberi cannot be assigned to Pterophyllum
central part, but the small polar ends are only weakly because this latter genus is a type of bennettitalean
cutinized. foliage. The material here assigned to Nilssonia neuberi
has haplocheilic stomata that are typical for cycadalean
Locality: Lunz-am-See, Lower Austria. leaves (Florin, 1933; Watson and Cusack, 2005).
Source Strata: Lunzer Schichten, Julian, middle Car- Although only cuticles can give definite proof for the
nian, Triassic. generic assignment of this type of pinnate foliage, it
Material studied: About 30 specimens from the cannot be denied that some Pterophyllum species are
following collections: GBA, NRM superficially similar to N. neuberi. If no cuticles are
available, generic assignments remain speculative. This
Remarks latter category includes several species from the Upper
The leaf architecture and epidermal anatomy, particularly Triassic and Jurassic, which either have been assigned to
the insertion of the leaf segments and the morphology of Pterophyllum, Pseudoctenis or Nilssonia.
the stomatal apparati justify the accommodation of this Pterophyllum braunsii Schenk looks somewhat similar but
species in the genus Nilssonia. Nilssonia neuberi can the insertion of the leaf segments is not clear and cuticles are

Plate IV. Nilssonia lunzensis nov. spec. (see on page 7)

1. detail of central part of a leaf, epitype (NRM, S148602), scale bar = 1 cm;
2. large and well preserved leaf showing characters of the leaf segments (NHM, 2006B0008/0031), scale bar = 1 cm;
3. detail of a leaf, showing vein courses of overlapping leaf portions, cf. text (GBA, 1909/003/0196), scale bar = 5 mm;
4. central part of a well preserved leaf (GBA, 1909/002/0208), scale bar = 1 cm;
5. excellently preserved leaf with recognisable apical region, epitype (NRM, S148602), scale bar = 1 cm;
6. central part of a well preserved leaf, holotype (GBA, 1909/003/0196), scale bar = 2 cm;
7. a part of an excellently preserved leaf (NHM, 1883/C/5900), scale bar = 1 cm;
8. detail of a leaf, with leaf segments inserted at the upper side of the rachis and well recognisable vein courses (NHM, 1885/D/4020), scale
bar = 5 mm;
9. upper part of a leaf showing vein courses (GBA, 1909/003/0195), scale bar = 5 mm.

Plate V. Nilssonia lunzensis nov. spec. (see on page 8)

1. adaxial cuticle, overview of cell pattern (NRM, S148241/0006), scale bar = 100 μm;
2. abaxial cuticle, overview of cell pattern with stomata (NRM, S148241/0006), scale bar = 100 μm;
3. adaxial cuticle, overview with densely arranged epidermal cells (NRM, S148241/0001), scale bar = 100 μm;
4. adaxial cuticle, with haplocheilic stoma (NRM, S148241/0006), scale bar = 50 μm;
5. adaxial cuticle, showing cuticular bars (NRM, S148241/0002), scale bar = 10 μm;
6. abaxial cuticle, sunken stoma (NRM, S148241/0006), scale bar = 10 μm;
7. adaxial cuticle, epidermal cells with solid cell walls (NRM, S148241/0001), scale bar = 50 μm;
8. adaxial cuticle, sunken stoma (NRM, S148241/0002), scale bar = 10 μm;
9. adaxial cuticle, sunken stoma (NRM, S148241/0002), scale bar = 10 μm; 1
10. adaxial cuticle, stoma with monocytic arranged subsidiary cells (NRM, S148241/0002), scale bar = 20 μm;

Plate VI. Nilssonia neuberi nov. spec. (see on page 9)

1. central part of a well preserved leaf, holotype (GBA, 1909/003/0576), scale bar = 1 cm;
2. part of an excellently preserved leaf (NRM, S148579), scale bar = 1 cm;
3. detail of a leaf, with leaf segments inserted at the upper side of the rachis, epitype (GBA, 2006/004/0014), scale bar = 1 cm;
4. abaxial cuticle, stoma with strongly cutinized dorsal walls of the guard cells (NRM, S148579/0007), scale bar = 10 μm;
5. very large and excellently preserved leaf, epitype (GBA, 2006/004/0014), scale bar = 3 cm;
6. abaxial cuticle, overview showing arrangement pattern of stomata (NRM, S148579/0004), scale bar = 100 μm;
7. abaxial cuticle, stoma with strongly cutinized dorsal walls of the guard cells (NRM, S148579/0004), scale bar = 10 μm;
8. abaxial cuticle, stoma with strongly cutinized dorsal walls of the guard cells (NRM, S148579/0004), scale bar = 10 μm;
9. abaxial cuticle, overview showing arrangement pattern of stomata and vein courses (NRM, S148579/0007), scale bar = 100 μm.

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx 17

unknown (Schenk, 1867). Leaf segments attached to the leaf remains with well preserved cuticles, and thus en-
upper side of the rachis and non-bifurcated veins that are ables the characterisation of the four Nilssonia species
typical features for Nilssonia characterize Pterophyllum described above.
robustum Compter from the Keuper (Upper Triassic) of The Lunz flora is dominated by bennettitaleans, e.g.,
Franconia, southern Germany (Compter, 1894; Rühle von Pterophyllum. In addition, cycadaleans such as the here
Lilienstern, 1933; Kelber and Hansch, 1995). The size of described species of Nilssonia are common. Other typical
the leaves (Compter, 1874, 1894; Kelber and Hansch, 1995, elements include marattialean and osmundalean ferns
Fig. 155) is similar to that of Nilssonia neuberi. Compter (e.g., Asterotheca Presl ex Corda, Speirocarpus Stur), and
(1894) discussed the assignment of P. robustum to Ptero- sphenopsids (Equisetites Sternberg, Neocalamites Halle).
phyllum, whereas Kelber (1998) suggested that this species Conifers have been recorded, but are rare. More than 70
might be assignable to Pseudoctenis. taxa have to date been reported from the Lunz flora
Nilssonia princeps Oldham et Morris from the Rajmahal (Dobruskina, 1998). Although several other rich floras
Group (Jurassic) of India is another species that is very have been reported from the Carnian [e.g. Neuewelt near
similar to Nilssonia neuberi (Seward, 1917; Sahni and Basel, Switzerland (Heer, 1865; Leuthardt, 1901, 1903;
Rao, 1931, 1934). This species was originally described Kräusel and Leschik, 1955, 1959; Kräusel and Schaarsch-
as Pterophyllum princeps (Oldham and Morris, 1863; midt, 1966), Raibl, NE-Italy (Bronn, 1858; Schenk, 1866;
Feistmantel, 1877) and assigned to the cycads. Menen- Stur, 1885), Franconia, S-Germany (Kelber and Hansch,
dez (1952) gave a detailed description of N. princeps 1995)], the Lunz flora still remains the richest and most
from the Rhaetian Llantenes flora of Argentina. Nils- diverse Late Triassic flora of the Northern Hemisphere
sonia princeps is characterized by very large leaves with (Dobruskina, 1989, 1994, 1998).
segments that insert to the upper side of the rachis and The here reported species of Nilssonia are the
non-bifurcated, densely spaced veins. The epidermal oldest well-documented representatives of the genus.
anatomy of N. princeps is unknown. However, the leaf Nilssonia is known from Lunz, but has not been
segments of N. princeps are always shorter than those of recorded for more or less coeval floras such as
N. neuberi. Neuewelt and Raibl. However, cuticular studies with
regard to coeval floras primarily focussed on fertile
4. Discussion organs and not on sterile foliage. Without cuticles, it is
difficult, if not impossible, to distinguish between
As has already been mentioned before, only cuticles bennettitalean and cycadalean leaves. Broglia Loriga
can give definite proof for the assignment of isolated et al. (2002) briefly mentioned the occurrence of
pinnate foliage to the Cycadales or Bennettitales. If cu- Nilssonia in the Anisian flora from the Kühwiesen-
ticles are unavailable, assignments remain speculative and kopf in the Dolomites (N-Italy). Schenk (1867)
result in an artificial system of morphotaxa that do not reported two Nilssonia species from the Upper Keuper
necessarily reflect biological relationships. The assign- (Rhaetian) of Franconia, and Ash (2001) mentioned a
ment of several foliage morphotypes previously accom- Nilssonia species from the Upper Triassic Chinle
modated in the bennettialean foliage morphogenus Formation of North America. Nilssonia is very
Pterophyllum to the cycadalean foliage morphogenus common in the Jurassic, and known from various
Nilssonia based on a complement of macromorphological parts of the world, e.g., Europe (Nathorst, 1879, 1909;
and epidermal characters demonstrates the value of cutic- Lundblad, 1950; Harris, 1964; Kvacek, 1995; Kelber,
ular analyses in more accurately depicting the systematic 1998; Watson and Cusack, 2005), Iran and Afghani-
position of compression foliage fossils based on biolog- stan (Sadovnikov, 1989; Schweitzer et al., 2000), India
ical criteria. (Sahni and Rao, 1931, 1934), Japan (Yabe, 1925),
However, even if the generic assignment to Nilssonia Greenland (Harris, 1932, 1946; Boyd, 2000).
is clear, individual species within this genus are often Within the Triassic of the paratropics several phases
difficult to discriminate because they are highly variable in the development of the flora can be recognized.
(Nathorst, 1909; Harris, 1932; Boyd, 2000; Schweitzer Earliest Triassic floras were dominated by lycopsids
et al., 2000). Moreover, morphologically very similar (Pleuromeia Corda ex Giebel). In the early Anisian a
species may show distinct differences in the epidermal conifer-dominated flora with Voltzia Brongniart, Pe-
anatomy (Schweitzer et al., 2000). According to these lourdea Seward and Aethophyllum Brongniart appeared.
latter authors, species can only be adequately character- Ladinian floras include a number of elements also
ized by studying several complete leaves. The Carnian known from the Anisian, e.g., the lycopsid Annalepis
flora of Lunz includes a considerable number of large zeilleri Fliche, the fern Anomopteris mougeotii

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
18 C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx

Brongniart, together with Pelourdea vogesiaca Schim- During the Carnian, Lunz was positioned at a
per et Mougeot and several species of Voltzia. New palaeolatitude of c. 30° N (Scotese, 2003; Blakey,
elements include the conifer Elatocladus Halle, the fern 2005). According to reconstructions of Scotese (2003),
Cladophlebis Brongniart and several types of cycada- the climate in this region was cool temperate to locally
lean foliage, e.g., Bjuvia Florin and Sphenozamites arid. Palaeogeographical reconstructions show that the
(Brongniart) Miquel (Kerp, 2000). Although the earliest Lunz flora grew in a lowland setting. The coal-bearing
unequivocal bennettitalean fossils (Pterophyllum jae- beds are intercalated between fully marine beds. The
geri [Schlotheim] Brongniart) have been reported from presence of coal seams indicates that the environment
the Ladinian, it is a rare element (Kelber and Hansch, was stable for longer periods of time, enabling the
1995). Bennettitaleans did not become common until accumulation of large amounts of plant material (Kerp,
the Carnian, when they were dominant, at least locally 2000). The preservation of the large amounts of peat,
like in Lunz (Kelber, 1998; Pott et al., subm.). which were later altered into coal, requires special
The major floristic change took place after the Carnian conditions, including minimal oxidation and bacterial
and before the Rhaetian. Rhaetian floras show a much decay. These latter conditions are usually reached in
more modern aspect than Carnian floras, and are char- peat bogs with a high groundwater table, reduced
acterized by the first major occurrence of a number of oxygen supply and low pH values. The abundance of
leptosporangiate ferns, e.g., Dipteridaceae and Matonia- ferns and sphenophytes, the latter often occurring in
ceae, and conifers, e.g., Cheirolepidiaceae (Hirmeriella- monotypical associations, indeed indicate humid to very
ceae) (Kelber and Hansch, 1995; Kerp, 2000). Putative humid environments. The abundance of very large
Cheirolepidiaceae have been reported from the Chinle randomly oriented and well preserved leaves suggests
Formation (Carnian) of North America, e.g., Pagiophyl- minimal transport of the plant material.
lum (Ash, 1970), but Cheirolepidiaceae are never The Lunz flora contains a number of elements that are
common before the Rhaetian (Cleal, 1993). This is also regarded as typical for younger Mesozoic floras, i.e.
evidenced by the palynological record. Although cheir- Rhaetian (Nathorst, 1876, 1878–1886; Johansson, 1922;
olepidiaceous pollen is known from older Triassic strata, Harris, 1946; Lundblad, 1950; Kelber, 1998) and
the first real abundance of Corollina (Classopollis) has younger. Lunz is one of the earliest occurrences of the
been recorded for the Rhaetian (Visscher and Brugman, bennettitaleans, notably the morphogenus Pterophyllum
1981; Traverse, 1988). (Cleal, 1993; Kelber, 1998, 2005). Moreover, the Lunz
The earliest Dipteridaceae (e.g., Clathropteris Brong- flora contains very early representatives of the bennetti-
niart) have been described from the middle Carnian of talean genus Nilssoniopteris Nathorst (Pott et al.,
Franconia (Kelber and Hansch, 1995), and this genus is subm.). The description of the cycadalean foliage
also known from Lunz (Stur, 1885). The earliest morphogenus Nilssonia further substantiates the signif-
Matoniaceae (Phlebopteris Brongniart) are known icance of the Lunz flora with regard to a more complete
from the Carnian Chinle Formation of North America understanding of the vegetational changes and evolu-
(Ash et al., 1982; Ash, 2005). However, altogether tionary innovations that occurred during the Middle to
Rhaetian floras are much more similar to Jurassic floras Late Triassic, especially the first appearances of genera
than to older Triassic floras (Kelber and Hansch, 1995, that are very common and widespread in the Rhaetian–
Kelber, 1998). Unfortunately, very little is known about Jurassic.
the Norian vegetation because diverse Norian macro- As a result, the Lunz flora obviously represents a
floras are very rare, and most Norian floras are dom- mixture, in which typical elements of older Mesozoic,
inated by conifers (Kelber and Hansch, 1995; Ash, i.e. pre-Rhaetian, floras co-occur with forms that are
1999). characteristic constituents of younger and more modern,
Although the fossils Nilssonia from Lunz do not post-Carnian floras. A number of recent studies indicate
represent the earliest record of the genus it is noteworthy that several plant groups and genera evolved much
that Nilssonia, a genus traditionally regarded as a earlier in the Triassic than previously thought (Wachtler
typically Jurassic group, is represented by four different and Van Konijnenburg-van Cittert, 2000; Broglia Loriga
species. In addition, Nilssonia is relatively common in et al., 2002; Passoni and Van Konijnenburg-van Cittert,
the Lunz flora. The earliest Cycadales have been 2003; Kustatscher et al., 2004; Kustatscher and Van
described from the Lower Permian of China (Zhu and Konijnenburg-van Cittert, 2005). Therefore, it may be
Du, 1981; Gao and Thomas, 1989a,b). Nevertheless, concluded that, within the Triassic, a major floral
cycadalean foliage is relatively rare in pre-Carnian turnover did not occur, but rather the floral composition
strata. changed gradually.

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx 19

Acknowledgments Bhardwaj, D.C., Singh, H.P., 1964. An Upper Triassic miospore

assemblage from the coals of Lunz, Austria. Palaeobotanist 12, 28–44.
Blakey, R., 2005. Global Earth History. Published online at http://jan.
Financial support was provided by the Deutsche ucc.nau.edu/~rcb7/RCB.html. Last hit: 21.03.2006.
Forschungsgemeinschaft (DFG grant KR 2125/3-1 to Boyd, A., 2000. Bennettitales from Early Cretaceous floras of West
MK and HK). We are indebted to E. M. Friis (Stock- Greenland: Pterophyllum and Nilssoniopteris. Palaeontographica
holm, Sweden), M. Gross (Graz, Austria), M. Harzhau- Abteilung B Paläophytologie 255, 47–77.
Broglia Loriga, C., Fugagnoli, A., Van Konijnenburg-van Cittert, J.H.A.,
ser (Vienna, Austria), H. Steininger (St. Pölten, Austria),
Kustatscher, E., Posenato, R., Wachtler, M., 2002. The Anisian
J. H. A. van Konijnenburg-van Cittert (Leiden and macroflora from the Northern Dolomites (Monte prà della Vacca/
Utrecht, The Netherlands) and I. Zorn (Vienna, Austria) Kühwiesenkopf, Braies): a first report. Rivista Italiana di Paleonto-
for making available the Lunz specimens for cuticular logia e Stratigrafia 108, 381–390.
analysis, and to K. P. Kelber (Würzburg) for helpful Brongniart, A., 1825. Observations sur les Végétaux fossiles
comments on the manuscript. renfermés dans les Grès de Hoer en Scanie. Annales des Sciences
Naturelles 4, 200–224.
Bronn, H.G., 1858. Beiträge zur Triasischen Fauna und Flora der
Appendix A bituminösen Schiefer von Raibl, nebst Anhang über die Kurr'sche
Sippe Chiropteris aus dem Lettenkohlen-Sandsteine. Schweizer-
Abbreviations used in the text are as follows: bart, Stuttgart.
GBA — Geologische Bundesanstalt, Wien (Geological Cleal, C.J., 1993. In: Benton, M.J. (Ed.), Gymnospermophyta. The
Fossil Record, vol. 2. Chapman and Hall, London.
Survey of Austria, Vienna, Austria), Compter, G., 1874. Ein Beitrag zur fossilen Keuperflora. Nova Acta
NHM — Naturhistorisches Museum, Wien (Museum of der kasierlichen Leopoldinisch-Carolinischen Deutschen Akade-
Natural History, Vienna, Austria), mie der Naturforscher 37, 10–13.
NRM — Naturhistoriska Riksmuseet, Stockholm Compter, G., 1894. Die fossile Flora des unteren Keupers von
(Museum of Natural History, Stockholm, Sweden), Ostthüringen. Zeitschrift für Naturwissenschaften 37, 205–230.
Cornet, B., Olson, P.E., 1990. Early to Middle Carnian (Triassic) Flora
MNB — Museum für Naturkunde, Berlin (Museum of and Fauna of the Richmond and Taylorsville Basins, Virginia and
Natural History, Berlin, Germany), Maryland, U.S.A. Virginia Museum of Natural History, Martinsville.
RUU — Laboratory of Palaeobotany and Palynology, Dobruskina, I.A., 1989. The alpine Lunz-flora — a standard flora for
University Utrecht, The Netherlands, the carnian stage of the Triassic. International Geological Review
NAT — Nationaal Naturhistorisch Museum Naturalis, 31, 1209–1215.
Dobruskina, I.A., 1994. Triassic Floras of Eurasia. Springer, Wien.
Leiden (Museum of Natural History, Leiden, The Dobruskina, I.A., 1998. Lunz flora in the Austrian Alps — a standard
Netherlands), for Carnian floras. Palaeogeography, Palaeoclimatology, Palaeoe-
JOA — Landesmuseum Joanneum, Graz (State Museum cology 143, 307–345.
Joanneum, Graz, Austria), Dower, B.L., Bateman, R.M., Stevenson, D.W., 2004. Systematics,
ontogeny, and phylogenetic implications of exceptional anatomi-
SPO — Niederösterreichisches Landesmuseum, St. Pölten
cally preserved cycadophyte leaves from the Middle Jurassic of
(State Museum of Lower Austria, St. Pölten, Austria), Bearreraig Bay, Skye, Northwest Scotland. Botanical Review 70,
PBO — Forschungsstelle für Paläobotanik, Münster 105–120.
(Palaeobotany Research Group, Münster, Germany). Dunay, R.F., Fisher, M.J., 1978. The Carnian palynofloral succession
in the Northern Calcarous Alp, Lunz-am-See, Austria. Pollen et
References Spores 20, 177–187.
Feistmantel, O., 1877. Jurassic (Liassic) flora of the Rajmahal Group
Ash, S.R., 1970. Pagiophyllum simpsonii, a new conifer from the in the Rajmahal Hills. Memoirs of the Geological Society of India,
Chinle Formation (Upper Triassic) of Arizona. Journal of Palaeontologia Indica Serie II 53–162.
Paleontology 44, 945–952. Florin, R., 1933. Studien über die Cycadales des Mesozoikums.
Ash, S., 1999. An Upper Triassic upland flora from North Central New Kungliga Svenska Vetenskapsakademiens Handlingar, Tredje
Mexico, U.S.A. Review of Palaeobotany and Palynology 105, Serien 12, 1–134.
183–199. Fontaine, W.M., 1883. Contributions to the Knowledge of the Older
Ash, S., 2001. New cycadophytes from the Upper Triassic Chinle Mesozoic Flora of Virginia. Government Printing, Washington.
Formation of the southwestern United States. Paleobios 21, 15–28. Gao, Z., Thomas, B.A., 1989a. Occurrence of earliest cycads in the
Ash, S., 2005. A new Upper Triassic flora and associated invertebrate Permian of China and its bearing on their evolution. Chinese
fossils from the basal beds of the Chinle Formation, near Cameron, Science Bulletin 34, 766–769 [in Chinese].
Arizona. Paleobios 25, 17–34. Gao, Z., Thomas, B.A., 1989b. A review of fossil cycad megaspor-
Ash, S., Litwin, R.J., Traverse, A., 1982. The Upper Triassic fern ophylls, with new evidence of Crossozamia Pomel and its
Phlebobteris smithii (Daugherty) Arnold ant its spores. Palynology associated leaves from the Lower Permian of Taiyuan, China.
6, 203–219. Review of Palaeobotany and Palynology 60, 205–223.
Bachmayer, F., Kollmann, H.A., 1969. Schätze im Boden—Bilder aus Geologische Bundesanstalt Wien, GBA, 2002. Rocky Austria-Eine
Österreichs Vergangenheit, 2nd edn. Naturhistorisches Museum bunte Erdgeschichte von Österreich. Geologische Bundesanstalt
Wien, Wien. Wien, Wien.

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
20 C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx

Harris, T.M., 1932. The fossil flora of Scoresby Sound East Greenland — Lundblad, A.B., 1950. Studies in the Rhaeto–Liassic floras of Sweden.
Part 3: Caytoniales and Bennettitales. Meddelelser om Grønland 85, I. Pteridophyta, Pteridospermae and Cycadophyta from the mining
1–133. district of NW Scania. Kungliga Svenska Vetenskapsakademiens
Harris, T.M., 1943. Notes on the Jurassic Flora of Yorkshire. Annals Handlingar, Fjärde Serien 1, 1–82.
and Magazine of Natural History, Series II 10, 838–854, 7–9. Menendez, C.A., 1952. La Flora Mesozoica de la Formacion
Harris, T.M., 1946. Liassic and Rhaetic plants collected in 1936–38 Llantenes. Revista del Instituto Nacional de Investigacion de las
from East Greenland. Meddelelser om Grønland 114, 1–40. Ciencas Naturales 11, 147–261.
Harris, T.M., 1964. The Yorkshire Jurassic Flora II. Trustees of the Nathorst, A.G., 1876. Anmärkningar om den fossilen floran vid Bjuf i
British Museum (Natural History), London. Skåne. Öfversigt af Konglika Vetenskaps-Akademiens Förhan-
Heer, O., 1865. Die Urwelt der Schweiz. Schulthess, Zürich. dlingar 1, 29–41.
Johansson, N., 1922. Die rhätische Flora der Kohlengruben bei Nathorst, A. G., 1878–1886. Om floran i Skånes kolförande
Stabbarp und Skromberga in Schonen. Kungliga Svenska bildningar-I. Floran vid Bjuf. Sveriges Geologiska Undersökning
Vetenskapsakademiens Handlingar 63, 1–78. Serie C 27, 33, 85, 1–126.
Kelber, K.-P., 1998. Phytostratigraphische Aspekte der Makrofloren Nathorst, A.G., 1879. Om floran i Skånes kolförande bildningar-I.
des süddeutschen Keupers. Documenta Naturae 117, 89–115. Floran vid Bjuf (andra häftet). Sveriges Geologiska Undersökning.
Kelber, K.-P., 2005. Beyond the Permian–Triassic extinction events: the Ser. Ca, Avhandlingar Och Uppsatser 335, 55–82.
highly diverse lower Keuper flora (Ladinian, Triassic) of southern Nathorst, A.G., 1909. Über die Gattung Nilssonia Brongn. Kungliga
Germany. Workshop on Permian–Triassic Palaeobotany and Paly- Svenska Vetenskapsakademiens Handlingar 43, 3–37.
nology Bolzano/Bozen (Italy), June 16–18, 2005, Abstract Book. Oldham, T., Morris, J., 1863. Fossil Flora of the Rajmahal Series in the
Kelber, K.-P., Hansch, W., 1995. Keuperpflanzen. Die Enträtselung Rajmahal Hills. Memoirs of the Geological Society of India,
einer über 200 Millionen Jahre alten Flora. Museo 11, 1–157. Palaeontologia Indica Series II 1–52.
Kerp, H., 1990. The study of fossil gymnosperms by means of Passoni, L., Van Konijnenburg-van Cittert, J.H.A., 2003. New taxa of
cuticular analysis. Palaios 5, 548–569. fossil Carnian plants from Mount Pora (Bergamasc Alps,
Kerp, H., 2000. The modernization of landscapes during the Late Northern Italy). Review of Palaeobotany and Palynology 123,
Paleozoic–Early Mesozoic. In: Gastaldo, R.A., DiMichele, W.A. 321–346.
(Eds.), Phanerozoic Terrestrial Ecosystems. Paleontological Soci- Potonié, H., 1899. Lehrbuch der Pflanzenpalaeontologie mit beson-
ety Papers, pp. 79–113. derer Rücksicht auf die Bedürfnisse des Geologen. Dümmler,
Kerp, H., Krings, M., 1999. Light microscopy of cuticules. In: Jones, Berlin.
T. (Ed.), Fossil Plants and Spores: Modern Techniques. Geological Rühle von Lilienstern, H., 1933. Die Pflanzenwelt des unteren
Society, London, pp. 52–56. Keupers. Mitteilungen der Gemeinde der Steinsburgfreunde 2,
Krasser, F., 1909. Zur Kenntnis der fossilen Flora der Lunzer 12–22.
Schichten. Jahrbuch der Kaiserlich-Königlichen Geologischen Sadovnikov, G.N., 1989. Taeniopteris, Nilssoniopteris and Nilssonia
Reichsanstalt 59, 1–26. in the Late Triassic flora of Iran. Paleontological Journal 23,
Kräusel, R., 1953. Ein neues Dioonitocarpidium aus der Trias von 95–100.
Lunz. Senckenbergiana 34, 105–108. Sahni, B., Rao, A.R., 1931. On some Jurassic plants from the
Kräusel, R., Leschik, G., 1955. Die Keuperflora von Neuewelt bei Rajmahal Hills. Journal and Proceedings of the Asiatic Society of
Basel-I. Koniferen und andere Gymnospermen. Schweizer Bengal (New Series) XXVII 183–208.
Paläontologische Abhandlungen 71, 1–27. Sahni, B., Rao, A.R., 1934. Rajmahalia paradoxa gen. et sp. nov. and
Kräusel, R., Leschik, G., 1959. Die Keuperflora von Neuewelt bei other Jurassic plants from the Rajmahal Hills. Proceedings of the
Basel-III. Equisetaceen. Schweizer Paläontologische Abhandlun- Indian Academy of Sciences, I 258–269.
gen 77, 5–19. Schenk, A., 1866. Über die Flora der schwarzen Schiefer von Raibl.
Kräusel, R., Schaarschmidt, F., 1966. Die Keuperflora von Neuewelt Würzburger Naturwissenschaftliche Zeitschrift VI, 10–20.
bei Basel-IV. Pterophyllen und Taeniopteriden. Schweizer Paläon- Schenk, A., 1867. Die fossile Flora der Grenzschichten des Keupers
tologische Abhandlungen 84, 5–79. und Lias Frankens. Kreidel, Wiesbaden.
Krystyn, L., 1978. Eine neue Zonengliederung im alpin-mediterranen Schimper, W., Schenk, A., 1890. Palaeophytologie. In: Zittel, K.A. (Ed.),
Unterkarn. In: Zapfe, H. (Ed.), Beiträge zur Biostratigraphie der Handbuch der Palaeontologie. Oldenbourg, München, pp. 211–232.
Tethys-Trias. Springer, Wien. Schmidt, M., 1928. Die Lebewelt unserer Trias. Hohenlohe'sche
Kustatscher, E., Van Konijnenburg-van Cittert, J.H.A., 2005. The Ladinian Buchhandlung, Öhringen.
flora (Middle Triassic) of the Dolomites: palaeoenvironmental Schweitzer, H.-J., Kirchner, M., Van Konijnenburg-van Cittert, J.,
reconstructions and palaeoclimatic conclusions. GeoAlp 2, 31–52. 2000. The Rhaeto–Jurassic flora of Iran and Afghanistan. 12.
Kustatscher, E., Wachtler, M., Van Konijnenburg-van Cittert, J.H.A., Cycadophytina II. Nilssoniales. PAlaeontographica Abteilung B
2004. A number of additional and revised taxa from the Ladinian Paläophytologie 254–263.
flora of the Dolomites, Northern Italy. GeoAlp 1, 57–69. Scotese, C.R., 2003. Paleomap Project. Published online at www.
Kvacek, J., 1995. Cycadales and Bennettitales leaf compressions of the scotese.com. Last hit: 21.03.2006.
Bohemian Cenomanian, central Europe. Review of Palaeobotany Seward, A.C., 1917. Fossil Plants. Hafner, New York. reprint 1969.
and Palynology 84, 389–412. Stur, D., 1871. Lunzer Sandstein. In: Direction des geognostisch-
Leuthardt, F., 1901. Beiträge zur Kenntnis der Flora und Fauna der montanen Vereines für Steiermark (Ed.), Geologie der Steiermark.
Lettenkohle von Neuewelt bei Basel. Eclogae Geologicae Helve- Verlag des geognostisch-montanen Vereines für Steiermark, Graz,
tiae 125–128. pp. 242–316.
Leuthardt, F., 1903. Die Keuperflora von Neuewelt bei Basel-I. Teil Stur, D., 1885. Die obertriadische Flora der Lunzer-Schichten und des
Phanerogamen. Abhandlungen der Schweizer Paläontologischen bituminösen Schiefers von Raibl. Denkschriften der kaiserlichen
Gesellschaft 30, 1–23. Akademie der Wissenschaften Wien 3, 93–103.

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.
 ARTICLE IN PRESS
C. Pott et al. / Review of Palaeobotany and Palynology xx (2006) xxx–xxx 21

Stur, D., 1888. Die Lunzer (Lettenkohle-)Flora in den “older mesozoic beds Watson, J., Cusack, H.A., 2005. Cycadales of the English Wealden.
of the Coal-Field of Eastern Virginia”. Verhandlungen der Kaiserlich- Monograph of the Palaeontographical Society 1–189.
Königlichen Geologischen Reichsanstalt Wien 10, 203–217. Watson, J., Sincock, C.A., 1992. Bennettitales of the English Wealden.
Traverse, A., 1988. Paleopalynology. Unwin Hyman, Boston. Monograph of the Palaeontographical Society 1–228.
Turek, V., Marek, J., Benes, J., 1990. Fossilien. Aventinum, Praha. Yabe, H., 1925. Notes on some Mesozoic plants from Japan, Korea
Van Konijnenburg-van Cittert, J.H.A., Mikuz, V., Pavsic, J., 2001. and China, in the collection of the Institute of Geology and
Pterophyllum (Cycadopsida) from Carnian beds in Poljane valley Paleontology of Tôhoku Imperial University. The Scientific
(Slovenia). Geologija 44, 317–323. Reports of the Tôhoku Imperial University, Sendai, Japan —
Verloop, J.H., 1908. Profil der Lunzer Schichten in der Umgebung von 2nd Series (Geology) 7, 1–28.
Lunz. Zeitschrift der Deutschen Geologischen Gesellschaft 60, 81–88. Zhu, J., Du, X., 1981. A new cycad — Primocycas chinensis gen.
Visscher, H., Brugman, W.A., 1981. Ranges of selected palynomorphs et sp. nov. discovers from the Lower Permian in Shanxi, China and
in the Alpine Triassic of Europe. Review of Palaeobotany and its significance. Acta Botanica Sinica 23, 401–404 [in Chinese].
Palynology 34, 115–128.
Wachtler, M., Van Konijnenburg-van Cittert, J.H.A., 2000. The fossil
flora of the Wengen Formation (Ladinian) in the Dolomites (Italy).
Beiträge Zur Paläontologie 25, 105–141.

Please cite this article as: Christian Pott, et al., Morphology and epidermal anatomy of Nilssonia (cycadalean foliage) from the Upper Triassic of
Lunz (Lower Austria), Review of Palaeobotany and Palynology (2006), doi:10.1016/j.revpalbo.2006.07.007.