Kustatscher et al. 2006 - science . naturalis

[Palaeontology, Vol. 50, Part 5, 2007, pp. 1277–1298] 

HORSETAILS AND SEED FERNS FROM THE MIDDLE 

TRIASSIC (ANISIAN) LOCALITY KÜHWIESENKOPF 

(MONTE PRÀ DELLA VACCA), DOLOMITES, 

NORTHERN ITALY 

by EVELYN KUSTATSCHER*, MICHAEL WACHTLER and 

JOHANNA H. A. VAN KONIJNENBURG-VAN CITTERTà 

*Naturmuseum Südtirol, Bindergasse 1, 39100 Bolzano, Italy; e-mail: Evelyn.Kustatscher@naturmuseum.it 

P.-P. Rainerstrasse 11, 39038 Innichen, Italy; e-mail: michael@wachtler.com 

àLaboratory of Palaeobotany and Palynology, Budapestlaan 4, 3584 CD Utrecht, and National Natural History Museum ‘Naturalis’, PO Box 9517, 2300 RA Leiden, 

the Netherlands; e-mail: J.H.A.vanKonijnenburg@bio.uu.nl 

Typescript received 8 March 2006; accepted in revised form 30 October 2006 

Abstract: Well-preserved floras from the Alpine Early– 

Middle Triassic are rare, and thus our understanding of 

the vegetation in this area during this period of time continues 

to be incomplete. As a result, every new find represents 

a significant piece of information that deserves 

thoughtful consideration. Anisian (Middle Triassic) sphenophytes 

and pteridosperms have recently been recovered 

from the Kühwiesenkopf locality (Monte Prà della Vacca) 

in northern Italy. The sphenophytes are represented by 

stem fragments, strobili and isolated sporangiophore heads 

of Equisetites, as well as by a few specimens of Neocalamites 

sp. and Echinostachys sp. The pteridosperms include 

abundant remains of the peltasperm foliage type Scytophyl- 

Early to early Middle Triassic floras from the German 

Basin are known from a number of localities in France 

(e.g. Schimper and Mougeot 1844; Fliche 1910; Grauvogel-Stamm 

1978) and Germany (e.g. Blanckenhorn 1886; 

Schimper 1869; Frentzen 1915; Mägdefrau 1931; Gothan 

1937; Fuchs et al. 1991). By contrast, contemporaneous 

floras from the Alpine Triassic are very rare; the most 

famous of these comes from the Recoaro area in Italy 

(e.g. de Zigno 1862; Schenk 1868). This flora, along 

with a few other, less diverse records from localities elsewhere, 

suggests that the vegetation that occurred in the 

Alpine realm during the Early–early Middle Triassic was 

dominated by conifers (de Zigno 1862; Schenk 1868; 

Gümbel 1879). Recently, an additional, much richer, 

early Middle Triassic impression ⁄ compression flora has 

been discovered in the marine Anisian succession at 

Kühwiesenkopf (¼ Monte Prà della Vacca) in the Pragser 

(¼ Braies) Dolomites of northern Italy (Broglio Lori- 

lum bergeri. A second Scytophyllum species in this flora, 

S. apoldense, is regarded as conspecific with S. bergeri based 

on epidermal anatomy; the two morphotypes are interpreted 

as sun and shade leaves of a single biological 

species. The seed-bearing disc Peltaspermum bornemannii 

sp. nov. probably represents the female reproductive 

structure of S. bergeri. Additional pteridosperm remains 

include foliage assignable to Sagenopteris sp. and Ptilozamites 

sp., in both cases perhaps the earliest records of 

these genera. 

Key words: fossil Equisetales, Pteridospermae, Dolomites, 

Italy, Middle Triassic, Anisian. 

ga et al. 2002; Kustatscher 2004; van Konijnenburg-van 

Cittert et al. 2006). 

This flora is significant because it represents one of 

only a few floras with well-preserved plant fossils (some 

also yield excellently preserved cuticles) from the Alpine 

early Middle Triassic, and thus adds substantially to a 

more complete understanding of the vegetation during 

this period of time. 

Broglio Loriga et al. (2002) presented a preliminary 

report on the macrofloral remains from Kühwiesenkopf. 

van Konijnenburg-van Cittert et al. (2006) provided a 

detailed analysis and taxonomic revision of the ferns. 

Here we present the second part of a detailed analysis of 

the Anisian Kühwiesenkopf flora, which comprises 

descriptions and illustrations of the sphenophytes and 

pteridosperms. Taxa are revised where necessary. Comparisons 

with other coeval floras for the groups concerned 

are provided. 

ª The Palaeontological Association doi: 10.1111/j.1475-4983.2007.00707.x 1277

1278 PALAEONTOLOGY, VOLUME 50 

MATERIAL AND METHODS 

The section containing the plant horizon discussed crops 

out for several hundred metres along the western slope of 

Kühwiesenkopf on the north-eastern margin of the Dolomites 

(Text-fig. 1, GPS data for the locality, 12°3¢E, 

46°43¢N). It is well known through the detailed study of 

Bechstädt and Brandner (1970) and Senowbari-Daryan 

et al. (1993), and has been referred to the Dont Formation 

(for details, see Broglio Loriga et al. 2002; Kustatscher 

et al. 2006), a carbonate-terrigenous sequence more 

than 200 m thick in this section. The plant-bearing horizon 

is about 75 m above a massive carbonate platform 

attributed to the Gracilis Formation (De Zanche et al. 

1992; Broglio Loriga et al. 2002; van Konijnenburg-van 

Cittert et al. 2006). 

The plant-bearing horizon is 1 m thick. The plant 

remains are concentrated in some cm-thick lens-shaped 

layers of siltstone, which change laterally in number and 

thickness and alternate with silty and marly limestone 

layers with only sparse plant remains. A few marine fossils 

(bivalves, brachiopods, ammonoids and fishes) occur in 

association. For more details, see Broglio Loriga et al. 

(2002). 

TEXT-FIG. 1. Geographical map indicating the locality of 

Kühwiesenkopf ⁄ Monte Prà della Vacca. 

The Dont Formation is considered to be Pelsonian– 

Illyrian in age (Delfrati et al. 2000 and references therein), 

although the base of the formation may be diachronous 

at different locations in the basin (see also Bechstädt and 

Brandner 1970; Kustatscher et al. 2006). Studies of brachiopod 

(Bechstädt and Brandner 1970) and foraminiferal 

(Fugagnoli and Posenato 2004) assemblages suggested a 

Pelsonian age for this section, but recent integrated studies 

of palynomorphs and ammonoids in the section have 

narrowed the time interval for the deposition of the plant 

horizon down to the boundary between the middle and 

late Pelsonian (Kustatscher and Roghi 2006; Kustatscher 

et al. 2006). 

The specimens recovered have been studied with the 

aid of a dissecting microscope and, when possible, cuticle 

and in situ spore preparations were made. For this purpose, 

small leaf pieces of organic material were macerated 

in Schulze’s reagent (KClO 3 and 30 per cent HNO 3) and 

neutralized with 5 per cent NH 4OH. The cuticles were 

then separated with the aid of needles into upper and 

lower components, and sporangia into single or groups of 

spores (depending on their maturity), which were then 

mounted in glycerine jelly and sealed with paraplast. 

Most of the macrofossil plant collection, including all 

figured specimens, is stored at the Naturmuseum Südtirol 

in Bozen ⁄ Bolzano (Italy) along with the cuticle and spore 

slides. Their numbers are prefixed by either ‘Küh’ or 

‘Pal’. The remainder of the collection is in Wachtler’s 

Museum Dolomythos at Innichen (San Candido, Italy). 

SYSTEMATIC PALAEONTOLOGY 

Division SPHENOPHYTA 

Order EQUISETALES Dumortier, 1829 

Family EQUISETACEAE Michaux, ex DC 1804 

Genus EQUISETITES Sternberg, 1833 

Equisetites mougeotii (Brongniart, 1828a) Wills, 1910 

Plate 1; Plate 2, figure 1 

Selected synonymy 

1827 Calamites arenaceus minor Jaeger, p. 37, pl. 3, 

figs 1–7; pl. 5, figs 1–3; pl. 6, fig. 1. 

1828a Calamites mougeotii Brongniart, p. 137, pl. 25, 

figs 4–5. 

1844 Calamites mougeotii Brongniart; Schimper and 

Mougeot, p. 58, pl. 29, figs 1–3. 

1844 Equisetum brongniartii Schimper and Mougeot, 

p. 53, pl. 27. 

1869 Equisetitum mougeotii Brongniart; Schimper, 

p. 278, pls 12, 13, figs 1–4. 

1886 Equisetitum mougeotii Brongniart; Blanckenhorn, 

p. 141, pl. 20, figs 13–16a.

KUSTATSCHER ET AL.: TRIASSIC HORSETAILS AND SEED FERNS FROM THE DOLOMITES 1279 

1894 Equisetites singularis Compter, p. 215, pl. 3, 

figs 3–7. 

1910 Equisetites mougeotii Brongniart; Wills, p. 282, 

text-fig. 20, pl. 15, fig. 3. 

1910 Equisetum mougeoti Brongniart; Fliche, p. 117, 

pl. 9, fig. 2; pl. 12, fig. 1; pl. 15, fig. 1. 

1915 Equisetites mougeotii Brongniart; Frentzen, pp. 

14–21, pls 10–11; pl. 12, figs 1–5. 

1922 Equisetites singularis Compter; Frentzen, pp. 3, 10. 

1928 Equisetites mougeotii Brongniart; Schmidt, p. 74, 

fig. 90. 

1937 Equisetites mougeotii Brongniart; Gothan, p. 254, 

pl. 31, figs 1–2. 

1978 Equisetites mougeotii Brongniart; Grauvogel-Stamm, 

p. 23, pl. 1, fig. 3. 

? 1994 Equisetitum mougeotii Brongniart; Sander and Gee, 

p. 120, fig. 12.5. 

Description. This taxon is rare in the Kühwiesenkopf flora; some 

stem fragments (Küh 349, 585–588, 674, 762, 777, 1030–1031, 

1193, Pal 562, 808–809) and a few fructifications (Küh 676, 714, 

797) have been found so far. The preservation of the stem fragments 

varies considerably; they occur as imprints of the vascular 

bundles (Küh 586–588, 674, 777, 1030, 1193, Pal 562, 808–809) 

or as thick, almost three-dimensionally preserved carbonized 

woody stems (Küh 349, 585, 762). The imprints are up to 24 cm 

long and 3Æ5 cm wide. One specimen (Küh 1030) is characterized 

by a 22Æ1-cm-long and 1Æ5-cm-wide (basal) stem fragment 

with four nodes and, from the top to the bottom, respectively, 

88-, 42-, 23- and 13-mm-long, internodes. The base of the stem 

is slightly curved to the right. The preservation allows observation 

of the upper part the outer side of the stem, while in the 

lower part the vascular bundles can be seen from the inner side 

(Pl. 1, fig. 1). A more apical fragment (Pl. 1, fig. 2; Küh 674; 

57 mm long, 36 mm wide) shows six nodes with 8–10-mm-long 

internodes. Another specimen (Küh 587) represents a stem apex, 

with two narrow nodes c. 3 mm apart. 

When preserved, the vascular bundles continue over the nodes 

(Pl. 1, fig. 1). Generally, the distance between two bundles is less 

than 1 mm; in one specimen only (Küh 588) the distance reaches 

1–1Æ5 mm. 

The almost three-dimensionally preserved stem fragments are 

up to 14 cm long and 4Æ3 cm wide. One of the specimens (Küh 

762; Pl. 1, fig. 3) is a compressed cast 33 · 12 mm in diameter 

with faint imprints of vascular bundles, enclosed by a thin 

woody layer (less than 1 mm thick). The other two specimens 

(Küh 349, 585) are characterized by thicker woody remains (12 

and 33 mm thick, respectively). In Küh 585 the vascular bundles 

seem to be preserved, with a distance between single bundles of 

c. 1 mm. 

One fertile specimen (Pl. 1, fig. 4; Küh 676) shows two strobili 

attached to the same branch. From this branch (80 mm 

long, 6Æ5 mm wide, without any articulation) two smaller branches 

(21–24 mm long, 3–3Æ5 mm wide) arise apically, each terminating 

in a strobilus. The obovate strobili are up to 40 mm 

long and 25–28 mm wide and consist of up to 5–6 whorls of 

sporangiophores. The penta- to hexagonal sporangiophore heads 

are c. 7Æ5–8Æ5 mm in diameter (Pl. 1, fig. 4). 

An almost three-dimensionally preserved strobilus (Küh 714) 

shows the sporangiophore heads very well (Pl. 1, fig. 5), while 

one isolated sporangiophore head (Küh 797) measures 9 mm in 

diameter. Both sporangiophore heads (Küh 676, 714) proved to 

be immature; some slightly immature trilete spores, 30–45 lm 

in diameter, were recovered (Pl. 1, figs 6, 7). 

One specimen (Küh 1307; Pl. 2, fig. 1) shows a disintegrated 

strobilus with rhomboedric strobili 6 mm diameter and a probable 

diaphragm of 15 · 18 mm with an imprint of the vascular 

bundles at the outer margin (distance between adjacent vascular 

bundles, 0Æ5 mm). 

Discussion. Schimper (1869, p. 279) considered Calamites 

mougeotii Brongniart, Equisetum brongniartii Schimper 

and Mougeot, 1844 and Calamites arenaceus Jaeger, 1827 

to be conspecific. Most authors have agreed with him 

concerning the first two species (e.g. see Grauvogel- 

Stamm 1978) occurring in the ‘Buntsandstein’ floras of 

France and Germany, but Equisetites arenaceus (Jaeger) 

Schenk, 1864, a characteristic species of the German 

Keuper, differs in sterile and fertile morphology (for a 

detailed discussion, see Frentzen 1915; usually stems of 

E. mougeotii are considerably narrower than those of 

E. arenaceus, and the vascular bundles are more widely 

spaced in the latter). Information on the vegetative parts 

of our material is limited to the structure of the stem and 

its vascular bundles; it is missing for the leaf sheaths, 

which are almost always preserved in E. arenaceus and 

very characteristic. Moreover, our strobili differ markedly 

from those in the emended diagnosis for this species as 

proposed by Kelber and van Konijnenburg-van Cittert 

(1998). Our material is distinctly larger (strobili 40 · 25– 

28 mm vs. 35 · 22 mm) and is characterized by 5–6 

whorls of pentagonal to hexagonal sporangiophores with 

pointed heads c. 7Æ5–9 mm in diameter, whereas E. arenaceus 

is characterized by nine whorls of 10–12 pentagonal 

to rounded hexagonal sporangiophores 1Æ5–5 mm in 

diameter. 

Another species that is probably conspecific with 

E. mougeotii is E. singularis Compter, 1894 from the 

lower Keuper flora of Apolda, Thuringia, Germany. The 

main difference, according to Compter (1911), is the 

slightly finer stem striations, and the nodes are sometimes 

swollen, but the latter also occurs in E. mougeotii 

(a preservational feature?), and the two species are 

exactly the same size. 

Genus NEOCALAMITES Halle, 1908 

Remarks. This genus was introduced by Halle (1908) for 

horsetail macro-remains with general Equisetites-like features 

but completely free leaves attached at the nodes. 

Material nowadays attributed to it has commonly been


assigned previously to Equisetites (e.g. Brongniart 1828a; 

Unger 1850), Calamites (e.g. Schenk 1864; Schönlein- 

Schenk 1865) or Schizoneura (e.g. Schimper 1869; Compter 

1874, 1894, 1911). 

Neocalamites sp. 

Plate 2, figures 2–4 

Description. Three specimens (Pl. 2, figs 2–4; Küh 037, 039, 611) 

have been attributed to Neocalamites, all preserved as stems with 

attached leaves at nodal level. From a 3Æ5–4Æ7-mm-thick stem 

arise 12–15 fragmentary leaves, although in one specimen (Küh 

037; Pl. 2, fig. 2) the diameter is wider, at 10Æ6 mm, and the 

number of counted leaves is 17 (only half of the number of 

leaves are exposed); they are 1Æ4–2Æ3 mm wide, and the maximum 

length (never complete) is 15Æ6 mm. In some leaves a central 

vein seems to be preserved. 

Discussion. The specimens have been assigned to Neocalamites 

because of their free leaves, attached to the stem at 

nodal level. However, the lack of any additional information 

prevents attribution to a species. Neocalamites is 

quite common in Middle and Late Triassic floras (e.g. see 

Grauvogel-Stamm 1978; Kelber and Hansch 1995), not 

only in France and Germany but also elsewhere (e.g. 

China; Wang 1996). 

Family ECHINOSTACHYACEAE Grauvogel-Stamm, 1978 

Genus ECHINOSTACHYS Brongniart, 1828b 

Remarks. Echinostachys was erected by Brongniart (1828b, 

p. 457) with the following diagnosis (translated): ‘fructification 

with elongated spike, flower or fructification, sessile, 

contiguous, subconical and echinate’, and the type 

species Echinostachys oblonga. Subsequently, Schimper and 

Mougeot (1844, p. 45) described Echinostachys cylindrica, 

and De Zigno (1862) described Echinostachys massalongii. 

After a detailed macro- and micromorphological study on 

both species, Grauvogel-Stamm (1978) considered E. oblonga 

to be the male (with in situ microspores) and 

E. cylindrica the respective female strobilus (with in situ 

megaspores) of the new combination Schizoneura-Echinostachys 

paradoxa (Schimper and Mougeot) Grauvogel- 

Stamm, 1978 (p. 70). 

Echinostachys sp. 


Description. A few specimens of Echinostachys-type strobili have 

been found in the Kühwiesenkopf flora (Küh 240, 1252, Pal 

552). The strobili are up to 4 cm long and 7Æ7 mm wide. The 

axis is surrounded by spirally arranged, slightly imbricate sporophylls. 

The head of each sporophyll is more or less rhomboidal, 

1–2 · 1–1Æ5 mm in size. No spores could be recovered from any 

of the strobili. 

Discussion. Although our specimens show the macromorphology 

typical of Echinostachys, the absence of in situ 

spores prevents their attribution to any of the French species 

noted above. Our material differs in any case from 

Echinostachys massalongii De Zigno because strobili of the 

latter are round to ovate, apically truncated or expanded. 

The ovate to lanceolate sporophylls with their broad bases 

and acuminate apices are slightly imbricate and twice as 

long as wide. 

Order PELTASPERMALES Taylor, 1981 

Family PELTASPERMACEAE Pilger and Melchior, in Melchior 

and Werdermann 1954 

Genus SCYTOPHYLLUM Bornemann, 1856 

Remarks. Bornemann (1856, p. 75) created the genus Scytophyllum, 

with the type species S. bergeri Bornemann, 1856 

(from the Middle Triassic of Thuringia), for pinnately dissected 

leaves of leathery consistency with a strong midrib 

and indistinct (or even invisible) secondary veins. The leaf 

is amphistomatic and the epidermis of the upper leaf surface 

contains fewer stomata and larger polygonal epidermal 

cells than that of the lower leaf surface. The stomata are 

deeply sunken and protected by c. 6 subsidiary cells. 

According to Bornemann (1856), his S. bergeri is 

conspecific with the original material of Odontopteris cycadea 

Berger, 1832, renamed subsequently as Odontopteris 

bergeri Goeppert, 1836 and Zamites bergeri Presl, in Sternberg 

1838. However, this is not the case, owing to a different 

frond morphology and cuticular anatomy (e.g. Harris 

1964). Odontopteris cycadea has been reassigned to Ctenozamites 

and is now widely known from mainly Jurassic 

sediments as Ctenozamites cycadea (Berger) Schenk, 1887 

(for synonymy, see Harris 1964, p. 95). 

EXPLANATION OF PLATE 1 

Figs 1–7. Equisetites mougeotii (Brongniart, 1828) Wills, 1910. 1, 3, stem fragments Küh 1030 and 762, respectively; · 0Æ5 and · 1. 

2, broad stem fragment, Küh 674; · 1Æ5. 4, two fertile strobili connected by two thin stems, Küh 676; · 1; 5, detail of fertile strobilus 

preserved in three dimensions, Küh 714; · 2. 6, group of immature spores, Küh 676; · 350. 7, immature spores, Küh 714; · 550.

6 

1 

4 

7 

KUSTATSCHER et al., Equisetites 

PLATE 1 

2 3 

5


Bornemann (1856, p. 76, pl. 7, figs 7–8) also described 

Scytophyllum dentatum, which differs from the type species 

in the more denticulate shape of the lobes. However, 

the cuticle seems to correspond to that of the type species, 

with only slightly larger epidermal cells with thinner 

walls; hence, we believe this material to be conspecific. 

Linnell (1933, p. 311) stated that the material figured by 

Bornemann (1856) corresponds only to a fragment of a 

single pinna and not to a leaf-fragment as he had supposed. 

Therefore, he emended (p. 310) the original diagnosis 

(translated here) to include in the genus ‘pinnate 

and bipinnate leaves with an apical pinna. The lanceolate 

or elongate pinnae are attached opposite each other or 

alternately. They arise from the primary rachis with a 

short petiole or are broadly to obliquely attached to the 

primary rachis. The venation is characteristic, with a distinct 

midrib and dichotomous secondary and tertiary 

veins that never anastomose. The epidermis of the 

amphistomatic leaves is thick and composed of polygonal 

epidermal cells with straight walls. The haplocheilic and 

irregularly scattered stomata consist of two sunken guard 

cells and a circle of subsidiary cells.’ 

Linnell (1933) considered S. dentatum to be conspecific 

with S. bergeri. He distinguished another species, Scytophyllum 

apoldense (Compter, 1874) Linnell, 1933, in the 

Keuper flora of Thale (Harz, Germany); this was originally 

described as Cycadites apoldensis Compter (1874) from the 

Keuper flora of Apolda. It consists of pinnate or bipinnate 

leaves with a thick rachis and one, or occasionally two, 

apical pinnae. The pinnae (largest fragment 17 cm long, 

3 cm wide) are attached alternately or opposite each other 

on the rachis, forming an angle of c. 45 degrees with it. 

They are elongate to lanceolate with a rounded apex and 

broad, often decurrent, base. In specimens with a broad 

lamina the margin becomes undulating. The venation is 

composed of a distinct midrib with secondary veins arising 

at an acute angle, forking at least once. The whole 

texture of the leaf is thinner than in S. bergeri, but the epidermis 

of the two species is identical. 

The high variability of leaf-shape within the Scytophyllum 

material was discussed by Linnell (1933, p. 326), who 

considered not only Cycadites apoldensis Compter (1874, 

p. 8, pl. 2, fig. 6) to be identical with his material from 

Thale, but also provisionally attributed Cycadites rumpfii 


Schenk (1864, p. 61, pl. 6, fig. 1; Compter 1894, p. 8, 

pl. 2, fig. 5) to the same species. The latter differs in leaf 

and pinna width, but according to Linnell (1933, p. 326) 

is identical in cuticle structure. However, Linnell did not 

consider Scytophyllum apoldense to be conspecific with 

S. bergeri because he noted some morphological differences 

and had no intermediate forms in his collection. 

We are of the opinion that the two species (occurring 

together not only at Thale but also in our flora) are conspecific 

and probably represent sun and shade leaves 

of one natural species (see discussion below under 

S. bergeri). 

Doweld (2001) considered Scytophyllum Bornemann, 

1856 to be a junior homonym of the extant angiosperm 

genus Scytophyllum Ecklon and Zeyher, 1835 and consequently 

proposed the new generic name Dellephyllum. 

However, according to the Index Nominum Genericorum 

Plantarum (Farr et al. 1979; website ING 2005) Scytophyllum 

Ecklon and Zeyher is a nomen rejectum; as a result, 

Scytophyllum Bornemann, 1856 does not have to be 

replaced by a new name (Dr Gea Zijlstra, IAPT, pers. 

comm. 2005). 

Scytophyllum bergeri Bornemann, 1856 

Plate 2, figures 7–9; Plates 3–4; Plate 5, figures 1–2; 

Text-figures 2–3 

Selected synonymy 

1856 Scytophyllum bergeri Bornemann, p. 75, pl. 7, 

figs 1–6. 

1856 Scytophyllum dentatum Bornemann, p. 76, pl. 7, 

figs 7–8. 

? 1864 Cycadites rumpfii Schenk, pp. 111–112, pl. 6, fig. 1. 

? 1874 Cycadites rumpfii Schenk; Compter, p. 8, pl. 2, 

figs 5, 7–8. 

1874 Cycadites apoldensis Compter, p. 8, pl. 2, fig. 6. 

? 1894 Cycadites rumpfii Schenk; Compter, p. 218. 

1894 Cycadites pinnatilobatus Compter, p. 219, pl. 4, 

fig. 1. 

1894 Cycadites apoldensis Compter, p. 219. 

1911 Thinnfeldia apoldensis Compter, p. 108, figs 41–42. 

1922 Thinnfeldia apoldensis Compter, p. 38, pl. 2, 

fig. 31. 

1922 Scytophyllum dubium Compter, p. 39, pl. 3, fig. 32. 

Fig. 1. Equisetites mougeotii (Brongniart, 1828) Wills, 1910; a possible diaphragm on the lower left side, on the upper right side a 

disaggregated cone, Küh 1307; · 1. 

Figs 2–4. Neocalamites sp., leaves attached at nodal level, Küh 037, 039 and 611, respectively; all · 1. 

Figs 5–6. Echinostachys sp., cone fragments Küh 240 and 1252, respectively; both · 2. 

Figs 7–9. Scytophyllum bergeri Bornemann, 1856. 7, leaf fragment showing a decurrent lower lamina, leaf type 1, Küh 1302; · 1. 8–9, 

apical leaf fragments, leaf type 1, Küh 963 and 977, respectively; · 1 and · 2.

KUSTATSCHER et al., Anisian plants 

PLATE 2 

1 2 3 4 

5 

6 7 

8 

9


1928 Scytophyllum dubium Compter; Schmidt, p. 88, 

fig. 132. 

1928 Scytophyllum dentatum Bornemann; Schmidt, 

p. 89, fig. 133. 

1928 Scytophyllum bergeri Bornemann; Schmidt, p. 89, 

fig. 134. 

? 1928 Danaeopsis rumphii Schenk; Schmidt, p. 67, fig. 69. 

1933 Scytophyllum bergeri Bornemann; Linnell, pp. 311– 

321 pl. 2, figs 1–5, text-figs 1–3. 

1933 Scytophyllum apoldense (Compter) Linnell, pp. 

321–328, pl. 2, figs 6–9, text-figs 4–7. 

1990 Scytophyllum cf. bergeri; Wang and Wang, p. 129, 

pl. 24, figs 1–3. 

? 1995 Cycadites rumpfii; Kelber and Hansch, p. 70, 

fig. 144. 

non 1995 Scytophyllum bergeri; Kelber and Hansch, p. 62, 

fig. 132. 

? 1996 Scytophyllum cf. bergeri Bornemann; Wang, pl. 4, 

fig. 1. 

2002 Scytophyllum; Broglio Loriga et al., pp. 384–385. 

2004 Scytophyllum sp.; Kustatscher, pp. 144–145, pl. 6, 

figs 2–3. 

Emended diagnosis. Paripinnate leaves. Axis stout, covered 

by small scales. Pinnae lanceolate to broadly lanceolate 

with rounded apex; pinnae inserted at an acute angle (c. 

45 degrees) opposite, subopposite or alternately to the 

rachis. Lamina not constricted at base, or showing a decurrent 

proximal and restricted distal margin. In sun 

leaves, pinna margin entire, lamina narrow. Shade leaves 

with crenate-lobate to undulating margin, giving rise to 

lobes of very variable dimensions, but never reaching 

midrib. Midrib distinct, secondary and ⁄ or tertiary veins 

rarely visible, arising at an angle of c. 45 degrees, and 

usually forking at least once. 

Epidermal cells isodiametric, slightly elongated above 

veins. Stomata sunken, surrounded by 6–7 subsidiary 

cells. Epidermal cells of sun leaves smaller and more often 

covered and protected by papillae than shade leaves. Stomata 

of sun leaves disposed along bands in intravenal 

areas of lower epidermis and rarely on upper epidermis. 

Stomata of shade leaves in bands between veins on lower 

surface, and irregularly scattered on upper surface. Epidermal 

cells in sun leaves smaller and more protected by 

papillae than in shade leaves. 

Description. Scytophyllum bergeri is a common fossil in the flora 

of Kühwiesenkopf (over 100 specimens), occurring usually as 

fragmentary or entire pinnae (e.g. Küh 410, 463, 541, 548, 624, 

626, 781, 804, 937–938, 958), although several specimens show 

leaf fragments (up to 30 cm long) with several pinnae attached 

(e.g. Küh 479, 541–542, 721, 1195–1196). 

Leaf-type 1. Pinnate leaf fragments up to 30 cm long, with pinnae 

variable in length and width. The most entire apical leaf fragment 

is paripinnate (Küh 479; Text-figs 2, 3A), 15 cm long and 

15 cm wide. The 1Æ8-mm-wide axis is covered by scales c. 3Æ5 mm 

in diameter. Four pairs of lanceolate pinnae attached opposite 

each other arise from the axis at a distance of c. 20 mm and an 

angle of 45 degrees. Apically the axis bifurcates and gives rise to 

two distinct pinnae, respectively 50 and 60 mm long and 15 mm 

wide, with their lamina attached broadly to the rachis. The basal 

pinnae with rounded apices are c. 10 cm long and 15 mm broad. 

The margin of the pinnae is always entire, the venation never distinct. 

Other apical leaf-fragments are small (e.g. Küh 963, 977, 

1003). Here too (e.g. Küh 963; Pl. 2, fig. 8) the rachis is covered 

by scales (2 · 1Æ2 mm). The elongate pinnae with an entire margin 

(32 · 8Æ5 mm) are attached opposite (Pl. 2, fig. 8) or subopposite 

each other on the rachis (Pl. 2, figs 7, 9). 

In the apical fragment of a young leaf (Küh 1046; 

10Æ2 · 6Æ6 cm), pinnae (40 · 11 mm basally – 24 · 7Æ5 mm 

apically) arise from a 3Æ5-mm-wide axis. The midrib is stout; the 

pinnae are attached opposite to it and almost touch each other 

(Pl. 3, fig. 1). 

Not all the narrow pinnate leaf-fragments, however, are apical 

fragments. Basal leaf-fragments occur as well (e.g. Küh 666, 

1003, 1194). In this case, the oppositely attached pinnae with 

entire margins show a decurrent lower lamina (e.g. Küh 119, 

1195, 1302; Pl. 2, fig. 7). 

Leaf-type 2. Other specimens (e.g. Küh 313, 463, 542, 563, 655, 

709AB, 804, 938, 960, 978, 996) are characterized by a rachis up 

to 7–18 mm broad and (sub)alternately attached pinnae up to 

60 mm apart. The latter (maximum length 100–140 mm, up to 

25 mm wide) have a crenate-lobate to undulating margin and 

arise without constriction from the rachis or are slightly constricted 

on the upper side (Küh 655; Pl. 3, fig. 2; Text-fig. 3B). 

In some specimens the incisions on the pinnae margin [e.g. Küh 

623, 804 (Pl. 3, fig. 3), 938] are pronounced, giving rise to lobes 

of very variable dimensions (4–17 · 8–16 mm). However, since 

the incisions never reach the midrib (2Æ5–4 mm wide) and show 

no constant pattern, these lobes cannot be considered as pinnules. 

In one case (Küh 1196; Pl. 3, fig. 4) the incisions give rise 

to ‘pinnules’ with a dentate shape (9Æ2 · 6Æ2 mm), resembling 

Scytophyllum dentatum as described by Bornemann (1856). 

All specimens studied are characterized by having a distinct 

midrib (1–3Æ5 mm wide), while the secondary and ⁄ or tertiary 

veins are rarely visible. When they are seen, the secondary veins 


Figs 1–7. Scytophyllum bergeri Bornemann, 1856. 1, apical leaf fragment, leaf type 1, Küh 1046; · 1. 2, fragment of leaf type 2, Küh 

655; · 1. 3–4, leaf type 2, pinnae with incisions on the pinnae margin, Küh 804 and 1196, respectively; both · 1. 5, leaf fragment 

with leaflets and interstitial pinna, Küh 731; · 1. 6–7, venation pattern, Küh 563 and Pal 464, respectively; · 1 and · 1Æ5.

1 

KUSTATSCHER et al., Scytophyllum 

PLATE 3 

3 4 5 6 

2 

7


TEXT-FIG. 2. Scytophyllum bergeri Bornemann, 1856, fragment of leaf type 1 (Küh 479), and detail; · 1 and · 1Æ5, respectively. 

A B C 

TEXT-FIG. 3. Line drawings of sun and shade leaves of Scytophyllum bergeri. A, schematic drawing of a sun leaf fragment, Küh 

479; · 0Æ5. B, schematic drawing of a shade leaf fragment, Küh 655; · 0Æ5. C, venation pattern, Küh 563; · 1Æ5.


arise at an angle of 45 degrees, and fork usually at least once 

(e.g. Küh 563, 969, Pal 464; Pl. 3, figs 6–7; Text-fig. 3C). 

Küh 731 (¼ Pal 519) shows a leaf fragment with a rachis and 

one almost complete pinna attached, basal fragments of two 

other pinnae, and the remains of an interstitial pinna (Pl. 3, 

fig. 5). Of particular interest is the presence of a pinna (124 mm 

long and 23 mm wide) with an undulating-incised margin and a 

distinct midrib (Küh 887; Pl. 4, fig. 1). Several small pinnae 

(14Æ5–15 · 2Æ5–3 mm) arise from the apex of this pinna. A similar 

structure was figured by Linnell (1933, p. 324, text-fig. 7b) as 

a fragment probably corresponding to a basal part of a small 

leaf, but we consider this specimen to correspond to a possible 

seedling, proof of agamous reproductive capacity of the species. 

Cuticle. That the two leaf-types discussed above belong to the 

same natural species is supported by our cuticular analysis. The 

epidermal cells of both are isodiametric, slightly elongated above 

the veins. The stomata are sunken and surrounded by 6–7 subsidiary 

cells, slightly thicker around the stomatal pit (Pl. 4, figs 6– 

8; Pl. 5, fig. 2). Although the general pattern is the same in both 

leaf-types, there are, however, also some small differences. The 

epidermal cells of the narrow-pinnate leaf-type 1 are smaller and 

more often covered and protected by papillae (Pl. 4, figs 4–5) 

than the leaf-fragments with broad pinnae of type 2 (Pl. 5, 

fig. 1). In leaf-type 1 the stomata are also disposed along bands 

between the veins on the cuticle of the lower surface (Pl. 4, 

fig. 4), whereas stomata are very rare on the cuticle of the upper 

surface (Pl. 4, fig. 5). Leaf-type 2, on the other hand, is characterized 

by epidermal cells of greater dimensions, only occasionally 

protected by papillae (Pl. 5, fig. 1). The stomata are 

disposed along bands between the veins on the lower surface, 

whereas on the upper surface they are scattered irregularly. The 

stomata of the upper leaf-surface (Pl. 5, fig. 2) are even less frequent 

than on the lower surface, but more abundant than on 

the upper surface of leaf-type 1. 

Particularly interesting also are some pinnae that reveal bite 

traces, probably caused by insects (e.g. Küh 119, 398, 551, 604, 

761, 887, 962, 977, 997, 1061; Pl. 4, figs 2–3). These traces are 

the only possible evidence of insects in the Middle Triassic of 

the Dolomites; hitherto, no insect remains have been recorded 

from this area. It is clear that the chewing took place during the 

life of the plant because the chewed leaf parts of several specimens 

have thicker margins. 

Discussion. According to our observations on the macromorphology 

of the Scytophyllum material from Kühwiesenkopf 

and on the cuticle analyses, we believe that all 

different leaf-shapes belong to the same natural variability 

of one species. This natural species includes specimens 

that were previously attributed to Scytophyllum bergeri, 

S. apoldense and S. dentatum as described from the 

Lettenkohle of Germany by Bornemann (1856) and Linnell 

(1933), of which S. bergeri Bornemann has priority of 

publication. 

The narrow-pinnate leaf-fragments of type 1 resemble 

the material described by Linnell, and by various other 

authors as Cycadites apoldensis (Compter 1874, 1894), 

Thinnfeldia apoldensis Compter (Compter 1911, 1922), or 

Scytophyllum apoldense (Linnell 1933; Mägdefrau 1953). 

Part of the specimens described as Cycadites rumpfii 

(Schenk 1864; Compter 1874, 1894) may also belong to 

this form of S. bergeri (in Kelber and Hansch 1995 as 

S. apoldense), but this is not clear as some authors have 

described the presence of sporangia on the margins of the 

pinnae (Compter 1874). 

The broad pinnate specimens of leaf-type 2 strongly 

resemble the material described as S. bergeri (Bornemann 

1856; Schmidt 1928, 1938; Linnell 1933), although ‘bipinnate’ 

forms are very rare. Compter (1894, p. 219, pl. 4, 

fig. 1) described Cycadites pinnatilobus, also from Apolda, 

as a new species for such a form, while in 1922 (p. 38, pl. 

3, fig. 32) he apparently described the same specimen as 

Scytophyllum dubium. However, there is no doubt that 

this type of leaf belongs to S. bergeri. 

The two different shape-types of the pinnae studied, as 

well as their respective cuticle patterns, may suggest an 

adaptation of the pinnae to, for example, different exposures 

to the sun. In this case, S. apoldense leaf-type 1 with 

narrow lamina and well-protected stomata would correspond 

to the sun-leaves of the species, whereas S. bergeri 

leaf-type 2 with broader pinnae and less protected stomata 

would correspond to the shade-leaves of the same 

species. 

The specimen described and figured as Scytophyllum 

bergeri in Kelber and Hansch (1995, p. 62, fig. 132) is not 

this species but a fragment of a Cladophlebis pinna. 

The number of species included in Scytophyllum is relatively 

low. Apart from the S. bergeri complex, which has 

been recorded extensively from Germany and herein from 

northern Italy, various other species have been described 

by Dobruskina (1969, 1980, 1995) from a number of 

Middle and Late Triassic floras in the former USSR, and 

by Mogucheva (1973) from the Early Triassic flora of the 

Tunguska Basin. Scytophyllum persicum (Schenk) Kilpper 

was recorded from a number of Norian and Rhaetian 

localities in Iran (Schweitzer and Kirchner 1998). All of 

these species are similar in gross morphology but differ in 

venation patterns, and, where known, in cuticular structure. 

Genus PELTASPERMUM Harris, 1937 

Remarks. The morphogenus Peltaspermum was introduced 

by Harris (1937, p. 34), although material had 

been already described and discussed under the name 

‘cupulate disc’ (Harris 1932, p. 65, pl. 6, figs 3–9; pl. 7, 

figs 3–9; pl. 8, figs 1–3, 5–6, 9–10; text-fig. 28). Peltaspermum 

is a genus for seed-bearing fructifications composed 

of an axis with lateral branches terminating in peltate


discs. Each peltate disc bears a ring of seeds under the 

surface. These seeds have a prominent micropyle, a free 

and cutinized nucellus, and a cutinized megaspore membrane. 

The stomata are haplocheilic; the guard cells are 

sunken and protected by a ring of subsidiary cells and 

papillae. 

Harris (1937) distinguished two species, the type species 

Peltaspermum rotula Harris, 1937 (for discs associated 

with leaves of Lepidopteris ottonis (Goeppert) Schimper, 

1869 and the male reproductive organ Antholithus zeilleri 

Nathorst, 1908 [ ¼ Antevsia zeilleri (Nathorst) Harris, 

1937] and Peltaspermum thomasii Harris, 1937 (for those 

associated with leaves of L. nataliensis and the male reproductive 

organ Antevsia extans), distinguished by their 

dimensions, the type of margin, the number of seeds, and 

the presence or absence of the typical blister-like swellings 

on the disc and the branches (see also Thomas 1933). The 

continuous association of these two sets of three morphospecies 

as well as their structural affinities, such as the 

presence or absence of blister-like swellings on the leafrachis 

and the axis of the reproductive organs, and the 

same cuticular morphology, indicate the same natural species 

(Harris 1932; Lundblad 1950; Townrow 1960). 

Townrow (1960) emended slightly the original diagnosis 

including ovuliferous organs with 5–15 (probably vascular) 

marginal lobes where the seeds may be disposed in 

pairs on each side of the branch (P. thomasii) or in numbers 

of 10–12 in a ring around the axis (P. rotula), and 

with Lepidopteris-like stomata. 

Poort and Kerp (1990) stated, however, that the name 

of the type species, Peltaspermum rotula Harris, was illegitimate, 

and made the new combination Peltaspermum 

ottonis (Harris) Poort and Kerp, 1990, because Harris 

(1932) had first described the seed-bearing discs as female 

fructifications associated with Lepidopteris ottonis. At the 

same time these authors published an emended diagnosis 

of Peltaspermum as a natural genus, including leaves and, 

for some species, also male inflorescences. However, 

according to Article 11.7 in the ICBN (International Code 

of Botanical Nomenclature; Greuter et al. 2000), this cannot 

be done; a morphogenus cannot become a natural 

genus (e.g. see example 25 in the ICBN, directly after Art. 

11.7). Moreover, Poort and Kerp (1990) assigned dispersed 

ovuliferous organs of supposed peltaspermaceous 

affinity but unknown linkage to sterile foliage and previously 

attributed to the genus Peltaspermum, to the morphogenus 

Lopadangium Zhao et al., 1980 as emended by 

Gomankov and Meyen (1986) if radially symmetrical, or 

to Autuniopsis Poort and Kerp, 1990 if bilaterally symmetrical. 

Schweitzer and Kirchner (1998) discussed the proposal 

of Poort and Kerp (1990) in their study on Rhaeto-Jurassic 

plant remains from Iran. They agreed that the leaf 

morphogenus Lepidopteris has female reproductive organs 

belonging to Peltaspermum but would not substitute Lepidopteris 

by Peltaspermum to create a natural genus. This 

decision was motivated also by the fact that Peltaspermum 

seed-bearing discs have also been attributed to Scytophyllum 

leaves by both Dobruskina (1969, p. 44; 1980, p. 99) 

and themselves, and that no specimens of Peltaspermum 

have so far been found in anatomical connection with 

either Lepidopteris or Scytophyllum. Therefore, Schweitzer 

and Kirchner (1998) maintained the morphogenus Peltaspermum. 

Holmes and Anderson (2005) noted that as 

Peltaspermum and Lepidopteris are both morphotaxa 

according to the ICBN, a new name would be required 

for a ‘natural genus’. As indicated above, we agree and 

therefore refer our material to Peltaspermum. 

Peltaspermum bornemannii sp. nov. 


2002 Peltaspermum; Broglio Loriga et al., pp. 384–385. 

2004 Peltaspermum sp.; Kustatscher, p. 144, pl. 6, fig. 3. 

Derivation of name. After Dr J. G. Bornemann, who first 

described Scytophyllum bergeri, the leaves to which this female 

fructification belongs. 

Types. Holotype: Küh 1301 ⁄ PAL464 (Pl. 5, figs 6–7). Paratypes: 

Küh 1079 (Pl. 5, fig. 3), Küh 2112B ⁄ Pal 525 (Pl. 5, fig. 4). 

Other material. Küh 848, 962A ⁄ B, 1300, 2112A. 

Diagnosis. Isolated ovuliferous organ of Peltaspermumtype 

having a more or less flattened, umbrella-shaped 

disc, c. 15–25 mm in diameter, with a central depression 

corresponding to the attachment area of the stalk and at 

least 15 marginal lobes. No attached seeds found so far. 

Cuticle of disc surface with isodiametrical epidermal cells 

and stomata arranged in irregular rows, consisting of two 

sunken guard cells and 6–8, slightly thickened subsidiary 

cells without papillae. 


Figs 1–7. Scytophyllum bergeri Bornemann, 1856. 1, presumed seedling, Küh 887; · 1. 2–3, bite traces, probably caused by insects, Küh 

1195 and 761, respectively; · 1Æ5 and · 1. 4–5, cuticle, leaf type 1, lower and upper sides, respectively, Küh 701; · 200. 6–7, cuticle, 

leaf type 1, stoma of the upper and lower sides, respectively, Küh 701; · 600.

1 

KUSTATSCHER et al., Scytophyllum 

5 

PLATE 4 

2 3 

6 7 

4


Description. This taxon is rare in the Kühwiesenkopf section; 

only a few specimens have been found so far. The isolated ovuliferous 

organs consist of more or less flattened, umbrella-shaped 

discs. Unfortunately, owing to the way they are preserved, only 

the upper surface is visible in all specimens, while the seeds are 

attached to the lower surface of the disc. 

The holotype (Pl. 5, fig. 6) is composed of an almost completely 

preserved umbrella-shaped disc, probably seen from the 

lower side. The maximum diameter of this specimen is 

21 mm, with an attachment area of the axis that is 3Æ7– 

5Æ5 mm wide. The disc seems to be covered by at least 16 

(immature?) seeds. Around the margin several marginal lobes 

with acute apices are visible. The maceration of some of the 

organic matter permitted the extraction of fragments of ovules 

(Pl. 5, fig. 7). 

The most complete specimen (Küh 1079; Pl. 5, fig. 3) is 

about 21 mm in diameter, 16 mm corresponding to the inner 

sterile part, with the area of the attachment of the stalk characterized 

by a depression of c. 2 mm. Around the margin at least 

15 lobes can be counted. Another specimen (Küh 962; Pl. 5, 

fig. 5), 15 mm in diameter, shows 16 marginal lobes. The upper 

surface of this specimen seems to be characterized by radially 

disposed ribs. The margin of a third specimen (diameter 

14 mm) has at least 15 lobes. Küh 2112A ⁄ B, the specimen yielding 

the largest cuticle fragments, is 18 mm in diameter (but is 

incomplete) and has a central depression of 3 mm (Pl. 5, fig. 4). 

The number of marginal lobes is not clear because part of the 

margin has not been preserved, but is probably around 16 (Pl. 

5, fig. 11). 

If the number of lobes on the margin and the number of ribs of 

the upper surface correspond to an identical number of seeds, as 

proposed by various authors (Harris 1932, pl. 8, figs 1–2, 9; Lundblad 

1950; Poort and Kerp 1990) for peltasperm ovuliferous 

organs, there should be at least 15–16 seeds in our ovuliferous 

structures. 

Küh 2112 (Pl. 5, figs 8–10) yielded cuticle fragments that consist 

of more or less isodiametrical epidermal cells, with stomata 

arranged in irregular rows, always with a number of normal epidermal 

cells between them. The stomata are sunken, and have 

two guard cells and 6–8 slightly thickened subsidiary cells without 

papillae. 

Discussion. Several of our specimens (Küh 962, 1301, 

2112) are closely associated, although not in organic connection, 

with a pinna attributed to Scytophyllum bergeri 


Bornemann (Pl. 5, fig. 5). This suggests that the leaves of 

S. bergeri and these ovuliferous organs belong to the same 

natural species, especially since Peltaspermum and Scytophyllum 

are so far the only representatives of the Peltaspermaceae 

found in the Kühwiesenkopf flora. 

Ovuliferous remains of Peltaspermum have also been 

found in association with Scytophyllum leaves in other 

Anisian localities of northern Italy. Remains of both have 

been collected from Furkelpass, a locality situated more 

to the west than Kühwiesenkopf that is possibly of the 

same age or slightly younger. An impression of Peltaspermum 

has also been recorded from another nearby locality, 

Hochalpenkopf. In the Carnian Alps (Val Pesarina) a 

three-dimensionally preserved umbrella-shaped disc has 

been recovered from the ‘Marne a Daonella’ Formation 

[ammonoids dated by Prof. Paolo Mietto (University of 

Padova) to the Avisianum subzone, Parakellnerites Zone], 

and is therefore Illyrian in age. 

As noted above in the discussion of Peltaspermum, 

most Triassic species have been associated with foliage 

belonging to Lepidopteris, but Dobruskina (1969, 1980, 

1995) also recorded Peltaspermum material from Russia 

in association with Scytophyllum foliage (e.g. with 

S. neuburgianum Dobruskina, 1980), and Schweitzer and 

Kirchner (1998) recorded Peltaspermum decipiens from 

the Late Triassic floras of Iran associated with Scytophyllum 

persicum. None of these has, however, been recorded 

together with S. bergeri foliage. Wang and Wang (1989, 

1990) described Peltaspermum lobalatum Wang and 

Wang, 1989 and P. calycinum Wang and Wang, 1990 

from two Early Triassic floras in North China. In at least 

one of these floras (the upper one) S. cf. bergeri foliage 

has also been found. P. calycinum is c. 2Æ5–3Æ5 cm in 

diameter and has a deeply divided margin with 8–10 

lobes with acute apices; this differs from our Peltaspermum 

material. P. lobalatum is 2–3 cm in diameter, and 

has 12–14 rounded marginal lobes that are similar in 

shape to the lobes in our peltate discs. However, as the 

number of lobes is slightly lower in our material, and the 

diameter of the discs can be less than 2 cm, we are not 

sure that it is conspecific with P. lobalatum. However, we 

Figs 1–2. Scytophyllum bergeri Bornemann, 1856. 1, cuticle, leaf type 2, upper side, Küh 566; · 200. 2, cuticle, leaf type 2, stoma of the 

upper side, Küh 542; · 600. 

Figs 3–11. Peltaspermum bornemannii sp. nov. 3, paratype, complete ovuliferous peltate disc, Küh 1079; · 2. 4, paratype, ovuliferous 

peltate disc, Küh 2112B ⁄ Pal 525; · 1. 5, ovuliferous peltate disc showing characteristic radially disposed midribs, and associated 

with pinnae of Scytophyllum, Küh 962; · 1. 6, holotype, partially preserved ovuliferous peltate disc, Küh 1301 ⁄ Pal 464; · 2. 7, 

fragmentary remains of ovules, Küh 1301 ⁄ Pal 464; · 150. 8, cuticle, Küh 2112B ⁄ Pal 525; · 150. 9, stoma; the guard cells have 

been lost during fossilization, Küh 2112B ⁄ Pal 525; · 100. 10, epidermal cells, Küh 2112B ⁄ Pal 525; · 100. 11, ovuliferous peltate 

disc viewed from the upper side showing bulges indicating the position of the ovules, Küh 2136 ⁄ Pal 529; · 1Æ5.

3 

6 

7 

KUSTATSCHER et al., Peltaspermum, Scytophyllum 

1 

8 

PLATE 5 

4 5 

10 11 

2 

9


are convinced that it is the female fructification of the 

type species, S. bergeri. 

Order CAYTONIALES Thomas, 1925 

Family CAYTONIACEAE Thomas, 1925 

Genus SAGENOPTERIS Presl, in Sternberg 1838 

Remarks. The leaf genus Sagenopteris was described for 

the first time by Presl in Sternberg (1838, p. 164) for pinnate 

fronds composed of four (or fewer often two) pinnae 

(¼ leaflets) with a strong midrib extending almost to the 

margin, and forking and anastomosing lateral veins forming 

irregular and narrow hexagonal meshes. Harris (1964, 

p. 3) emended the diagnosis to ‘petiolate leaves with each 

petiole bearing normally two pairs of leaflets at its apex, 

and petiole and leaflets being shed by absciss layers. The 

leaflets are lanceolate with a main vein at a greater or less 

distance distal to the mid-line. The lateral veins arise at a 

small angle from the midrib, then curve outwards, forking 

and anatomising to form obliquely elongated meshes. The 

ultimate veins end freely at the margin. Stomata are confined 

to the lower side of the cutinized leaves, oval and 

with sunken guard cells. The stomata are surrounded by a 

perigenous (unspecialized) ring of subsidiary cells.’ He 

also indicated as type species Sagenopteris nilssoniana 

(basionym Filicites nilssoniana Brongniart, 1825), 

although the first species noted by Presl in Presl in Sternberg 

(1838) was Sagenopteris rhoifolia, later considered to 

A B C 

TEXT-FIG. 4. A–B, Sagenopteris sp., leaf fragment, Küh 231, and leaflet fragment, Küh 1156, respectively; both · 2. C–D, 

Ptilozamites sp., pinna fragments, Küh 906 and 1122, respectively; · 1Æ5 and · 0Æ5. 

D 

be a synonym of S. nilssoniana (Brongniart, 1825) Ward, 

1900. 

Sagenopteris sp. 

Text-figure 4A–B 

Selected synonymy (only the last two are from the Dolomites) 

? 1990 Glossopteris shanxiensis Wang and Wang, p. 127, 

pl. 19, figs 5–8; pl. 20, fig. 3. 

? 1995 Gen. et sp. indet., glossopteridische Fieder; Kelber 

and Hansch, fig. 131. 

non 1995 Sagenopteris sp.; Kelber and Hansch, p. 66, 

figs 140–142. 

? 1996 Neoglossopteris shanxiensis Wang Zi-qiang, p. 129, 

pl. 2, figs 1–5. 

2002 ?Sagenopteris; Broglio Loriga et al., pp. 384–385. 

2004 ?Sagenopteris sp.; Kustatscher, p. 143, pl. 5, fig. 6. 


only a few specimens (Küh 231–234, 913, 1118, 1149, 1156) have 

been found so far. These are mostly fragments of leaf(lets) with 

a maximum length of 55 mm and a maximum width of 21 mm. 

The most complete specimen (Küh 231; Text-fig. 4A) is a nearly 

entire leaf(let) and a partially preserved second leaf(let) arising 

at an almost perpendicular angle. The more complete leaf(let) 

fragment is 52 mm long and 30Æ4 mm wide, but shows only 

about one-half of the leaf(let) (maximum width of the half-leaf 

is 23 mm); its base is restricted, widening upwards very quickly. 

The second leaf(let), although only partially preserved, is 42 mm 

long and 29 mm wide. The midrib is 1 mm wide at its base and 

decreases apically (width 0Æ5 mm). Fine secondary veins arise


from it; these dichotomise a few times and anastomose forming 

wide meshes (Text-fig. 4A–B). The single veins are up to 1– 

1Æ5 mm apart. Only the base of the second leaf(let) is visible 

with the midrib and a few secondary veins. In some specimens 

the large, lingulate shape of the leaf is visible (Küh 233, 1156; 

Text-fig. 4B). In these leaves, the dichotomising and anastomosing 

veins are more distant at the base (up to 1 mm) than at the 

apex (0Æ5 mm). 

Discussion. Our material has been attributed to Sagenopteris 

because of the ovoid to lanceolate shape of the leaflets, 

the strong midrib, and the dichotomising and 

anastomosing lateral veins forming elongate meshes, with 

one specimen (Küh 231) showing two partially preserved 

leaflets with their bases attached. Palynological samples 

yielded Vitreisporites pallidus, a bisaccate pollen grain that 

is typical of the Caytoniales to which Sagenopteris also 

belongs. Unfortunately the preservation of the specimens 

(no complete leaf or leaflet) is insufficient for an attribution 

to species level. 

The forking and anastomosing venation with elongate 

meshes opening at the margin might also suggest an attribution 

to the morphogenus Chiropteris, but this is not, 

however, possible owing to the presence of the well-developed 

midrib and the two attached but distinct leaflets in 

our material; in Chiropteris the funnel-shaped and incised 

leaves show only a vague midrib, and the leaflets are not 

really separated. The shape of the leaf(lets) and the bifurcating 

and slightly anastomosing veins might also suggest 

an attribution to the genus Linguifolium, but leaves of the 

latter are never composed of 2–4 leaflets attached at their 

bases but are placed opposite each other on the rachis. 

The pinnae of Linguifolium are characterized by a distinct 

straight midrib, which, however, stops well below the 

apex, whereas in Sagenopteris the midrib almost reaches 

the apex. 

Our material resembles specimens described first as 

Glossopteris shanxiensis by Wang and Wang (1990, p. 127, 

pl. 19, figs 5–8; pl. 20, fig. 3) and figured later as Neoglossopteris 

shanxiensis by Wang (1996, pl. 2, figs 1–5). This 

species is characterized by large fronds with simple leaves 

attached ‘in groups’ on a longitudinally ridged axis 1 cm 

wide. According to these authors, the lingulate leaves with 

an obtuse apex may reach a maximum length of 10 cm 

and a maximum width of 5 cm. The thick midrib 

becomes narrower apically, dissolving near the apex, while 

the lateral veins arise almost parallel from the mid-vein, 

curving outwards and finally forming an angle of 50–60 

degrees with the mid-vein. The venation is repeatedly 

dichotomising and anastomosing forming narrow, elongate 

meshes that become smaller near the margin. 

According to Wang (1996, p. 129), Neoglossopteris represents 

an endemic Early Triassic plant in China and 

includes pteridospermous fronds characterized by anasto- 

mosing venation. He considered it to be morphologically 

related to Gondwanan Glossopteris but differing from it 

because of its bifurcating apex, although this feature is 

not clearly figured. Although our specimens are represented 

just by small fragments, they differ in being half the 

dimensions of the Chinese specimens described and 

figured by Wang and Wang (1990) and Wang (1996) 

(maximum dimensions of 55 · 21 mm vs. 100 · 55 mm) 

and in the absence of a bifurcate apex, and are therefore 

unlikely to belong to the same species. 

Only a very few pre-Rhaetian (latest Triassic) Sagenopteris 

species are known so far from Europe: the type species 

Sagenopteris nilssoniana (Brongniart, 1825) Ward, 

1900, with its junior synonyms S. diphylla Presl, in Sternberg 

1838, S. rhoifolia Presl, in Sternberg (all three originally 

from the Rhaetian–Liassic area around Bamberg and 

Bayreuth) and S. semicordata Presl, in Sternberg 1838 

(from the Carnian flora of Sinsheim), has been recorded 

from Sweden, Germany, Hungary and Romania (e.g. see 

Kelber and Hansch 1995); Stur (1885) mentioned a Sagenopteris 

sp. in a list of taxa from Raibl but did not describe 

or figure the material; Kerner (1907, 1908) recorded 

Sagenopteris from Dalmatia, which he did not figure, but 

in 1907 he described in detail the five taxa of the florule, 

including material that he believed to belong to Sagenopteris 

based on the macromorphology of leaflet fragments 

and the typical net-venation, and in 1908 he gave the age 

of the florule as Ladinian, and assigned the Sagenopteris 

to S. cf. rhoifolia Presl. 

Unfortunately our material is not well enough preserved 

to attribute it to any of these European Triassic 

species, but it is the oldest known Sagenopteris material 

so far. Sagenopteris-type leaves also occur in the nearby 

Anisian locality of Furkelpass (see above under Peltaspermum 

bornemannii). 

Order indeterminate 

Genus PTILOZAMITES Nathorst, 1878 

Remarks. This genus was created by Nathorst (1878, pp. 

21–23) for linear pinnate and petiolate leaves. The pinnae 

are attached with their entire base to the rachis, the distal 

margin is perpendicular or slightly concave, and the proximal 

margin is rounded. The veins are usually dichotomising, 

disposed radially, and directed towards the basal 

margin of the pinnae. According to Nathorst, the genus differs 

from Anomozamites owing to its radiating veins, from 

Ptilophyllum and Otozamites because of the broadly 

attached pinnae, the rounded proximal margin of the pinnae 

and the thickness of the cuticle, and from Ctenozamites 

by being bi- or tripinnate and not just once pinnate.


Antevs (1914, pp. 3–8) emended the diagnosis and described 

the cuticle for the first time as ‘characterised by irregular 

to isodiametric epidermis cells, which become 

elongated and organised in regular rows above the veins 

and the rachis. The often papillate epidermis cells have 

straight or slightly undulated walls. The cuticle is thick; the 

leaves are amphistomatic or hypostomatic, with stomata 

concentrated in the intravenal areas of the lower cuticle. 

The guard cells are sunken and surrounded by irregularly 

shaped subsidiary cells, which form a ring-like thickening.’ 

It should be noted that Anomozamites, Ptilophyllum 

and Otozamites belong to the Bennettitales whereas Ctenozamites 

and Ptilozamites are seed ferns (e.g. see Harris 

1932). 

Ptilozamites sp. cf. P. sandbergeri (Schenk) Kustatscher and 

van Konijnenburg-van Cittert, 2007 

Text-figure 4C–D 

2004 ?Ptilozamites sp.; Kustatscher, p. 145, pl. 6, fig. 1. 

2007 Ptilozamites sp. cf. Pt. sandbergeri (Schenk) 

Kustatscher and van Konijnenburg-van Cittert, 

p. 84, fig. 4I, K. 


only a few specimens (Küh 070, 681, 903, 906, 1003, 1122, 1140) 

have been provisionally attributed to Ptilozamites. The leaf fragments 

are characterized by rectangular pinnae attached with 

their whole margin to the rachis of the leaves. They have a 

maximum length of 72 mm and a maximum width of 13 mm. 

One characteristic specimen (Küh 1122; Text-fig. 4D) is 28 mm 

long and 7 mm broad. Several almost square pinnae are attached 

on the 1Æ5-mm-wide rachis with their entire broad bases, five on 

the left side and seven on the right; they are 3 · 2Æ5 mm in size. 

A few small cuticle fragments have been extracted from one 

specimen (Küh 906; Text-fig. 4C). Although not very well preserved, 

they are thick and display polygonal epidermal cells with 

straight cell walls. Around some poorly preserved stomatal pits, 

surrounded by 4–6 subsidiary cells, a more or less well-preserved 

ring-like structure can be observed, as is typical for stomata in 

this genus. 

Discussion. Unfortunately, the specimens lack details of 

venation pattern and stomatal distribution; in gross morphology 

they resemble the macro-remains described as Ptilozamites 

heeri from Ladinian deposits in the Dolomites 

(Wachtler and van Konijnenburg-van Cittert 2000; Kustatscher 

2004; Kustatscher et al. 2004; Kustatscher and van 

Konijnenburg-van Cittert 2005). A recent study of both the 

original collections of the genus in Stockholm and the collections 

from the Alpine area resulted in a new attribution 

of the Ladinian specimens (Wachtler and van Konijnenburg-van 

Cittert 2000) to Ptilozamites sandbergeri (Schenk) 

Kustatscher and van Konijnenburg-van Cittert, 2007. As 

there are no real macromorphological and cuticular differences 

from the Ladinian–Carnian P. sandbergeri, we provisionally 

place our Anisian specimens from Kühwiesenkopf 

in this taxon. 

COMPARISON WITH OTHER COEVAL 

EUROPEAN FLORAS 

Anisian localities in Europe are well known in Germany 

and France. However, only few have been described from 

England (Grauvogel-Stamm 1972; Dobruskina 1994), 

Spain (Virgili 1958; Dobruskina 1994; Grauvogel-Stamm 

and Alvarèz Ramis 1996; Diez et al. 1996) and Poland 

(Schmidt 1928; Dobruskina 1994), and only one has been 

known so far from Italy (Recoaro: De Zigno 1862; Schenk 

1868; Gümbel 1879); all of these have yielded only a small 

number of horsetail, lycopsid and fern taxa, but many 

different conifers have been found (Kustatscher 2004). 

German Anisian localities are abundant and have been 

well studied over a long period of time (e.g. Blanckenhorn 

1886; Frentzen 1915; Schmidt 1928; Mägdefrau 

1931; Gothan 1937; Fuchs et al. 1991; Sander and Gee 

1994). The floras recovered, as well as those from coeval 

deposits in France (e.g. Brongniart 1828a, b; Schimper 

and Mougeot 1844; Fliche 1910; Depape and Doubinger 

1963; Grauvogel-Stamm 1969, 1978, 1991, 1993; Grauvogel-Stamm 

and Duringer 1983), mostly contain an abundance 

of horsetails, ferns and conifers. Our finding of 

horsetails at Kühwiesenkopf was therefore to be expected, 

though a few taxa typical of the German Basin, such as 

representatives of Schizoneura, are missing. The situation 

with respect to seed ferns is, however, entirely different. 

So far their remains have not been found at any other 

Anisian localities in Europe. Scytophyllum bergeri was previously 

known mainly from the Keuper (supposedly Ladinian) 

flora of Thuringia (Bornemann 1856; Compter 

1874; Linnell 1933). Peltaspermum has been described 

previously from, for example, the Permian of Germany 

and Italy (Poort and Kerp 1990), the Upper Triassic of 

Afghanistan (Schweitzer and Kirchner 1998), Russia 

(Dobruskina 1969) and South Africa (Holmes and Anderson 

2005), and the Rhaetian of Greenland (Harris 1932, 

1937) but never associated with S. bergeri. Only Wang 

and Wang (1990, 1990) have described a species of Peltaspermum 

that is also associated with Scytophyllum cf. bergeri 

foliage; these occur in Early Triassic floras of North 

China. 

Sagenopteris, by contrast, is well known from Rhaetian– 

Jurassic sediments (Harris 1964), but very few pre-Rhaetian 

species of this genus are known so far from Europe 

(e.g. Sagenopteris cf. rhoifolia from the Ladinian of 

Dalmatia, and Sagenopteris sp. and S. semicordata,

espectively, from the Carnian of Raibl and Sinsheim; for 

details, see above under Sagenopteris sp.). Therefore, the 

specimens from Kühwiesenkopf are the earliest record of 

this genus. 

The same is true for Ptilozamites, well known for the 

Swedish Rhaeto-Liassic species P. nilssonii, P. heeri and 

P. blasii. As noted above in the discussion of Ptilozamites 

sp. cf. P. sandbergeri, one species has been described from 

Carnian and Ladinian sediments of the Alpine area. The 

specimens from Kühwiesenkopf, although not well 

enough preserved to be attributed unequivocally to 

P. sandbergeri, represent the earliest record of this genus 

so far. 

Acknowledgements. We thank Dr Gea Zijlstra for her advice 

on nomenclature. Prof. Zhou Zhiyan provided us with Chinese 

literature and translations of some descriptions, for which 

we are very grateful. We especially thank Mr Paolo Fedele, 

Cortina, for his part in collecting the material. Sebastian and 

Fabian Pfeifhofer and Giorgio Zardini also provided us with 

specimens from the Dolomites; Luca Simonetto (Museo Friulano 

di Storia Naturale, Udine, Italy) provided the Peltaspermum 

specimen from the Carnian Alps. The manuscript 

benefited greatly from the comments of Dr M. Krings 

(Munich) and Mr K.-P. Kelber (Würzburg), and the editorial 

work of Prof. D. J. Batten (Manchester). 

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