Catalog of Middle Triassic plants

“My” plants of the Middle Triassic of the Dolomites and Germanic Basin (but not only)
You have found some plant fossils of Middle Triassic age but are not able give them a name? You have been searching for a general text book or article, which helps you to understand them – but are not able to find any?

What can you do? Of course you can always contact me! Please send me a picture using the form on the right, and I will try to help you! Or, you can have first a look at the general list of species that have been found in the Middle Triassic of the Dolomites and the Germanic Basin. I’ll give you a short, general description and some pictures of the material I have studied so far. At the end you will find also a list of books and articles that can help you to understand more about the fossils you have found.

N.B: of course these descriptions can include only material and species I have studied and seen so far. Names that are considered synonyms are not discussed here. Thus, this list must be considered “in evolution” and will updated once I have published new data. Inedited data will be avoided as much as possible in order to permit you to have only “correct” data.


Division Lycophyta

Selaginellites leonardii Kustatscher et al., 2010

Description: Heterosporous strobilus (at least 17 x 3 mm) with helically to sometimes almost decussately arranged sporophylls which are aligned in four irregular vertical files. The sporophylls are closely arranged, slightly imbricate. The sporophylls (1.5-2 x 1-1.2 mm) are ovate with a long, acuminate apex (about 2 mm long) and an entire margin. Sporophylls arranged in rows of microsporophylls and megasporophylls. Maceration of sporophyll fragments gave small cuticle pieces with isodiametric cells and sporangia containing masses of immature, in situ microspores and megaspores. The epidermis is characterized by isodiametric cells. The microspores are organised in compact tetrads of c. 56 (45-62.5) µm in diameter. Well-developed, separated trilete microspores are 46 (42.5-50) µm in diameter and characterized proximally by a punctate (granulate?) or smooth (psilate) and distally by a rugulate spore wall. Megaspores are oval to circular and around 350 µm in diameter. The trilete aperture is delicate or not yet well-developed, and only rarely indicated by plicae, the spore wall is about 10 µm thick.
Distribution
: Anisian of the Dolomites (Italy).


Isoetites brandneri Kustatscher et al., 2010

Description: Herbaceous lycophytes of up to 150 mm height and 120 mm width. From the unlobed corm (about 20 mm wide) arise simple rootlets (total structure about 37 mm long). Root traces of 1.0-1.5 mm diameter cover the surface. On the short stem are helically inserted elongate, 80-150 mm long and 3-5 mm wide sporophylls with a pointed apex. The leaf bases enlarge up to 3.5-5.0 mm and leaving rhomboidal leaf scars with delicate strap-like structures. Apically the elongate to lanceolate leaves are narrower, usually with a fold in the middle part. From the base of the leaves have been extracted immature groups of in situ microspores and mature megaspores.
The epidermis is thick on both sides (2.5-3.0 µm) and very variable. Epidermal cells are isodiametric in the central part (around 30-40 x 25-35 µm wide), and become more elongate near the margins. Abaxial epidermal cells are protected by thick papillae (15-25 µm in diameter). Stomata on the abaxial side are abundant and arranged in short, irregular rows. They are sunken, often heavily protected by papillae of the 5-7 subsidiary cells (sometimes covering the stomatal pit). On the margin of the leaves, more elongated “hair-like” appendices (up to 20 µm long and 10 µm wide) can be present. Adaxial epidermis has only few or no stomata, epidermal cells bear almost no papillae, only rarely slight thickenings. Variations of the “normal” epidermis sometimes occur with rows of more more elongated cells (3-5 cells wide) give origin to “vein-like” epidermal structures (?air channels) (cells 50 x 10-15 µm) or stomata common on both sides or sculptured subsidiary cells (instead of papillae).The immature microspores are reniform, up to 35-40 µm in longest diameter and are probably monolete. They could belong to the dispersed genus Aratrisporites. The oval to circular megaspores, 270-300 µm in diameter are convolute to verrucate with a thick spore wall. Proximally the ornamentation reduces in height. The trilete aperture is delicate, only a few times indicated by plicae.
Distribution
: Anisian of the Dolomites (Italy).

Lepacyclotes zeilleri (Fliche) Retallack, 1997

Description: The more or less wedge-shaped sporophylls are usually 25-45 mm long and 10-20 mm wide but in the Dolomites smaller specimens have been recorded (10-12 x 7 mm). The distal part is triangular, the proximal part is trapezoidale with a central, tongue-shaped sporangium (2 mm width). The sporangia contain either small monolete microspores, or large trilete megaspores.
Distribution: Anisian of the Vosges (France), upper Ladinian of the Dolomites (Italy) and of the German Basin (Thale, Schweinfurt) and the Middle Triassic of China.


Lepacyclotes bechstaedtii Kustatscher et al., 2010

Description: Herbaceous lycophytes. From the stem (11-15 mm in diameter and at least 20 mm height) arise several closely spaced sporophylls, which are arranged in whorls or helically. The corm is quadrilobate, 14-17 mm in diameter, with 23 mm long and 2 mm wide root fragments. The sporophylls are spateolate, c. 16-17 mm long and 7-10 mm wide; the central fertile area is oval, 9 mm long and 5 mm wide. In some cases, the apical part does not seem just simply triangular in shape, but almost trilobite. The sterile leaves are at the innermost circles, 5-6 mm wide and up to 20 mm long, with a central costae.
The normal epidermal cells are elongate (40-50 x 10 µm), on one side with and on the other without papillae. A few stomata are irregularly distributed on both epidermal sides. The stomata are simple, and the guard cells large and elongate.
Distribution
: upper Anisian of the Dolomites (Italy).


Lycopia dezanchei Kustatscher et al., 2010

Description: Subarborescent lycophyte. The stem fragments are up to 500 mm long and up to 40-60 mm wide. The biggest dichotomising rhizome fragment is 480 mm long and 135 mm wide with stem fragments arising alternately, first parallel to the rhizome. Afterwards they curve upwards and proceed perpendicularly to the rhizome. The rhizome is covered by small rootlets of 1.0-0.5 mm diameter and 2-5 mm length. Proximally, stems with rhomboidal leaf scars; apically, stems with leaves inserted in dense helices. A side branch is commonly going off apically. The leaves are long, lanceolate to elongate with a central midrib/costae, at least 125-150 mm long and 4.5 mm wide. The leaf scars are rhomboidal, 7-10 mm high and 9-11.5 mm wide. In the older and bigger stem fragments these leaf scars can even get as high as wide, seeming almost circular.
The cuticle is thick, amphistomatic. The epidermal cells are isodiametric to rectangular (25-30 x 40-50 µm). The stomata are on one side (upper?) few and scattered, common on the other (lower?) side and there arranged in rows. The stomata are sunken and protected by papillae on the 4-6 subsidiary cells.
Distribution
: upper Anisian of the Dolomites (Italy)


Division Sphenophyta

Equisetites arenaceus (Jaeger, 1827) Schenk, 1864

Description: The stem fragments are characterized by nodes and internodes that reduce apically and basally. A few specimens show stem fragments arising perpendicularly from a horizontal rhizome fragment (56 x 17 mm). In some cases stem fragments show ochreoles, interpreted as basal leaf sheaths of the lateral branches. Ochreoles are attached below the stem node; each ochreole is circa 3 mm in diameter and characterised by a whorl of 1-2 mm wide leaves. Leaves are fused basally, giving origin to leaf sheath fragments up to 115 mm long and 75 mm wide. The most complete leaf sheath displays 32 leaf teeth. Leaves are up to 8 mm long and 2.5 mm wide; the pointed apex is often missing, showing the sub-trapezoidal attachment area of 2.5-3 x 2-2.5 mm. Complete leaves show 5-15 mm long acuminate teeth.
The strobili are roundish, 15-21 mm in diameter and composed of whorls of penta- to hexagonal sporangiophore heads 3.5-4 mm in diameter.
Distribution
: Anisian of the Vosges, upper Ladinian of Germany, France and of the Dolomites (Italy), Carnian of the German Basin, Neue Welt (CH) and Lunz (A), “Lettenkohle” of Kazakstan and China, Keuper of Spain, Middle-Upper Triassic of Southern Priuralye and Southern Fergana, Triassic of Libya, Central Pamir, Thailand and Mongolia.
Remarks
: The fragments from the Ladinian of the Dolomites are smaller (stem fragments of 260 x 27-43 mm)

Equisetites mougeotii (Brongniart) Wills, 1910

Description: Stem fragmens preserved as imprints of the vascular bundles or as thick, almost three-dimensional preserved carbonised woody stems. The fragments are up to 24 cm long and 3·5 cm wide. The fructifications assigned to this species are two strobili attached to the same branch. From the branch (80 mm long, 6·5 mm wide, without any articulation) two smaller branches (21-24 mm long, 3-3·5 mm wide) arise apically, each terminating in a strobilus. The obovate strobili (40×25-28 mm) consist of up to 5-6 whorls of sporangiophores. The penta- to hexagonal sporangiophore heads are c. 7.5-9 mm in diameter.  The immature strobili yielded immature smooth, trilete spores, 30-45µm in diameter.
Distribution: Anisian of the Dolomites (Italy), Vosges (France) and Germany.


Neocalamites merianii (Brongniart) Halle 1908

Description: The stem fragments are up to 150 mm long and 40-60 mm wide, often threedimensionally preserved, and with 7-17 mm wide “costae” (ridges), which correspond to the outer layer (cortex). In specimens that lack the outer layer the vascular bundles beneath are exposed. Their number is much higher than the number of costae (6-10 times). At the nodes the stems are swollen, and in one case several leaves are attached.
Distribution: Ladinian and Carnian of the German Basin (Baden-Württemberg, Franken, Thüringen), Carnian of Lunz (A), Middle and Upper Triassic of China, Norian of Ukraina, and the Jurassic of Germany, Kazakhstan and Korea.


Neocalamites asperrimus (Franke) Shen 1990

Description: The cortex fragments are up to 9 cm long and 7 cm wide, and characterised by ornamentation. In most specimens the ornamentation is discontinuously undulating, the undulations can be reduced, giving origin to short undulations intercalated with a coarse granulate structure. In some specimens vascular bundles typical for sphenophytes are visible below the “cortical” structure. When the nodal point is preserved leaf scars are hinted but cannot be clearly distinguished, a slight nodal thickening can be observed.
Distribution: Ladinian of the German Basin (Thale am Harz, Molsdorf, Grossrettbach, Erfurt, Neudietendorf, Gotha), Late Triassic of China Tibet (Shensi, Yenchang District, Ichün District).


Neocalamites sp.

Description: Stem fragments with whorls of 12-17 fragmentary preserved leaves. The stem fragments are 3.5-10 mm thick. The leaves are 1.4 – 2.3 mm wide, the maximal length (never complete) is 15·6 mm. In some leaves a central vein seems to be preserved.
Distribution: Anisian of the Dolomites (Italy).


Echinostachys sp.

Description: These strobili are up to 4 cm long and 7.7 mm wide. The axis is surrounded by spirally arranged, slightly imbricate sporophylls. The head of each sporophyll is more or less rhomboidal, 1-2 x 1-1.5 mm in dimension. No spores could be recovered from any of the strobili.
Distribution: Anisian of the Dolomites (Italy).


Radicites sp.

Description: This is a morphotaxon used for compound equisetalean roots. The fragments (up to 100 x 50 mm) are characterised by a 2-3 mm thick main root and several secondary roots (30 x 1 mm). These secondary roots may be arranged into “whorls”. Sometimes also threedimensional, layer-crossing holes are visible in the rocks (that belong to secondary roots systems.
Distribution: Ladinian of the German Basin (e.g. Thale, Schweinfurt).


Division Pteridophyta


Anomopteris mougeotii Brongniart, 1828

Description: The frond fragments (up to 34 cm long) have a thick rachis (c. 5–10 mm) with long, almost perpendicular pinnae. The pinnae fragments are up to 11.5 cm long and 5 mm wide und show at their basis a roundish aphlebia. They are closely spaced but never in contact; their rachis is c. 1 mm wide. The pinnules arise perpendicularly and vary in size between 1. 2 mm and 2. 3 mm. Sterile pinnules often have a basal lobe. The midrib is delicate and forks in the apical part of the pinnule. Secondary veins arise at 60–90° and usually fork only once, the basal ones may fork twice. The basal part of the pinnae is sterile, the lower surface of the distal pinnules can be covered with sporangia.

Distribution: Anisian of the Vosges and Germany, upper Ladinian of the Dolomites (Italy)

Remarks: Grauvogel-Stamm and Grauvogel (1980) describe: circular, trilete in situ spores with 25–40 µm diameter and a punctate exospore.

Cladophlebis leuthardtii Leonardi, 1953

Description: The pinna fragments (20-25 mm long) are characterized by slightly falcate pinnules. These are attached with their whole base and are usually ca. 2.2 mm long and 2.5 mm wide. The venation is indistinct, except for the midrib, which ends long before the pinnule apex.

Distribution: Ladinian of the Dolomites (Italy)

Cladophlebis ruetimeyeri (Heer) Leonardi, 1953

Description: The bipinnate fronds are characterised by a narrow rachis and falcate pinnules (up to 83 x 8 mm) which reduce only slightly toward the tip (15 mm long). Midrib and secondary veins are delicate, the lower secondary veins fork twice, upper veins only once.

Distribution: Ladinian of the Dolomites (Italy)

Remarks: could be a junior synonym of Cladophlebis remota (see below).

Cladophlebis remota (Presl) Konijnenburg-van Cittert et al., 2006

Description: At least bipinnate fern. The frond fragments found so far are up to 20 cm long with a narrow rachis (2–3 mm wide). The slightly falcate pinnae arise alternately at an angle of 45° or even less near the apex. The pinnae are c. 10 cm long and basally about 2 cm wide with a 1.5 mm wide rachis. The pinnules (3–16×2–8 mm) are attached alternately with their whole base but are not adnate. There is a clear midrib and secondary veins that fork twice in the basal part of the pinnules, and only once more apically.

Distribution: Anisian of the Dolomites (Italy)

Cladophlebis sp.1

Description: At least bipinnate frond fragments with a relatively thick rachis (2-4 mm wide) which pinnae arising perpendicularly at 6-8 mm distance. The pinnae fragments are up to 7 cm long and 5 mm wide with a 1-1.5 mm wide rachis. The small, triangular-like pinnules (2-4 x 1.5-2.5 mm) arise perpendicularly from it. The venation consists of a midrib only, no secondary veins have been observed so far. Also apparently fertile fragments have been found but because of the poor preservation these fragments do not show any details of the sporangia

Distribution: Anisian of the Dolomites (Italy)

Remarks: This material has been assigned so far to no distinct because the specimens are to few and badly preserved.

Neuropteridium elegans (Brongniart) Schimper, 1869

Description: This small fern is characterised by small leaves (fragments up to 100 mm) and (round to ovoid) rhizomes of about 5-6 cm diameter From the rhizomes arise only few 2-3 cm long petioles. The leaves are characterised by small basal pinnules (4 x 2-3 mm) but increase to 15-20 x 6-7 mm. The venation is typically neuropterid with a marked midrib that extends 1/2-2/3 of the length. The secondary veins diverge and fork up to three times.

Distribution: Anisian and Ladinian of the Dolomites (Italy)

Neuropteridium voltzii (Brongniart) Schimper, 1879

Description: This species is characterised by bigger leaves than N. elegans. The rachis reaches about 3–6 mm width. Pinnules are up to 50 mm long and 4–9 mm wide, with a slightly pointed apex. The base is slightly contracted, the veination is neuropterid, with a clear midrib extending about 2/3 of the pinnule. Secondary veins are numerous and fork 2–3 times.

Distribution: Anisian of the Dolomites (Italy)

Gordonopteris lorigae Van Konijnenburg – van Cittert et al., 2006

Description: Tripinnate fern frond with broad, smooth rachis. The first-order pinnae arise alternately to suboppositely at angles of 45–60°, in close contact, but never overlapping. From the pinnae rachis (c. 2 mm wide) arise at 60–90 degrees relatively short (longest 4.5 cm) second-order pinnae, in close contact, but never overlapping. The roundish pinnules are relatively small (c. 2–3×2–3 mm), attached with the whole base. They containe a short midrib and forking secondary veins (neuropterid venation). Fertile pinnules have a reduced, rounded lamina with sporangia covering the lower side. The spores are globose, trilete,  45–60 µm in diameter and the exospore is finely punctate.

Distribution: upper Anisian and upper Ladinian of the Dolomites (Italy)

Marattiopsis sp.

Description: There are a few specimens that may represent pinna fragments of Marattiopsis. The pinnae fragments are up to 6 cm long and 1 cm wide basally, tapering towards the apex. The midrib is distinct, c. 1 mm wide at the base. Secondary veins arise at an angle of c. 60°, then curve outwards and fork once in the middle part of the lamina. The secondary veins reach the margin almost perpendicularly except in the apical area where the angle is smaller.

Distribution: Anisian of the Dolomites (Italy)

Danaeopsis sp.

Description: So far only pinnules fragments (up to 11.5 mm long and 2.5 cm wide)have been found, none of which is complete. The secondary veins arise from a broad midrib (2.5–4 mm wide) at an angle of c. 70°, then bend downwards and fork, usually once, near the midrib. Sometimes the veins fork more in the middle part of the lamina. They reach the margin more or less perpendicularly at a concentration of 8-12 each cm.

Distribution: Anisian of the Dolomites (Italy)

Scolopendrites scolopendrioides (Brongniart) Van Konijnenburg-van Cittert et al., 2006

Description: The biggest leaf fragments are about 10 cm long with a broad rachis (4-6 mm wide) and hanging pinnules (8-11 x 2-3 mm). Oval sporangia (600 x 200 µm) cover the lower surface of the pinnules. The spores are trilete, circular in equatorial outline (35–45 µm) with a scabrate to granulate exospore.

Distribution: upper Anisian of the Dolomites (Italy)

Scolopendrites grauvogelii Van Konijnenburg-van Cittert et al., 2006

Description: These fertile leaves are simply pinnate. From the relatively narrow rachis (0.6 – 3.0 mm) arise hanging pinnules (4-8 x 2-3 mm). The complete lower side of pinnules is covered with globular sporangia of 300-400 µm diameter. The spores are trilete, circular in equatorial outline (c. 35–45 µm diameter) with a scabrate to granulate exospore.

Distribution: upper Anisian of the Dolomites (Italy)

Scolopendrites sp.

Description: The so far only fragment is 46 mm long and 9 mm wide. The rachis is 2 mm wide, and pinnules arise oppositely and at almost rectangular angles. The pinnules are 3-4 mm long and 1.5-2 mm wide, and their whole lower surface is covered with sporangia.

Distribution: upper Ladinian of the Dolomites (Italy)

Sphenopteris schoenleiniana (Brongniart) Presl, 1838

Description: Tripinnate fern fronds with thin rachis (2–3 mm wide). The pinnae fragments are up to 15 cm long with several pinnae attached to the 1–2 mm wide rachis. Pinnules (10×3 mm) are usually attached with only part of their base at an angle of 30–60°, and are elongated or slightly falcate, giving an undulating margin. The venation consists of a midrib that does not reach the apex, and once forking secondary veins arising at c. 60 degrees. The fertile pinnules have sporangial attachment areas at the end of the secondary veins. Details of the sporangia are still unclear.

Distribution: upper Anisian of the Dolomites (Italy)

Gen. et sp. indet.

Description: Frond fragments (up to 8.5×3.3 cm) of a fern-like plant. The rachis is c. 3 mm wide and bears rhomboid pinnules varying in size between 17-20 x 14-16 mm. The venation arises from the lower basal angle of the pinnules as an undevelopped midvein that forks several times, and some veins below this ‘midvein’ arise directly from the rachis. The secondary veins end near the margin. The apex of the pinnules is relatively acute.

Distribution: upper Anisian of the Dolomites (Italy)

Remarks: Since the affinity of this species is still unclear (fern or seedfern) and only 2 frond fragments were found so far, this material has not yet been assigned to any genus and species.

Lugardonia paradoxa Kustatscher et al., 2009

Description: The strobili consist of a long axis (more than 200 mm long and 3.5-4.0 mm wide). Groups of 3-4 microsporangia (or pollen sacs) (c. 2.5-3.0 x 1.5-1.7 mm) are clustered together on small, less than 1 mm long stalks, which are helically arranged on the axis. The elongated microsporangia (c. 2.5-4.0 x 0.5-1.0 mm) yielded large spores (prepollen).
The microsporangia (c. 96-116 x 76-100 µm) are almost circular in equatorial outline, trilete, with a smooth inner layer (exospore) and a granulate-verrucate outer layer (perispore). The trilete mark is distinct, 3/4 – 7/8 of the radius, but not accompanied by a margo. The sculpture gets less pronounced under prolonged maceration and finally the perispore disappears.
TEM analysis of the spores (pollen) reveals that their aperture consists of a strongly protruding fold and that the wall in this area is made up of four or five layers (5 µm distally, 6 µm proximally, 8 µm at the laesural ridge). The outermost layer is of variable thickness (usually 0.5 to 2.0 µm) and corresponds to the perispore described above. This  outer perispore (OP) is electron lucent and often shows an apparently cellular structure between adjacent spores within the sporangium, individual cells having a slightly more electron-dense centre. The inner perispore (IP) is electron dense and of variable thickness (1 to 4 µm); it is thinnest over the apertural fold. The IP is permeated by numerous cavities of roughly circular outline and of less than 0.05 µm diameter, which seem to be distinctly radial aligned in the apertural area and distal wall. The exospore consists in the proximal part of the wall and particularly on both sides of the apertural fold of the outer and middle exospore (OE and ME respectively, about 1.5 to 7.0 µm in thickness). The OE is homogeneous in composition. The ME (where distinguishable) is slightly more electron dense and interrupted by rather angular cavities, laying lie parallel to the wall and may be indicative of otherwise obscured lamination within this part of the exospore. Tiny (0.02 µm) dark particles occur in the cavities indicating perhaps that these were filled with a material that has subsequently become coalified. The inner exospore (IE) is extremely thin, perhaps only 0.1 µm, and extends into (and defines) the commissure where it thins to nothing before reaching the outer edge of the OE.

Distribution: upper Anisian of the Dolomites (Italy)

Remarks: The affinity of this species is still unclear (fern or seedfern), although a fern affinity is considered more likely.

Division Pteridospermatophyta

Scytophyllum bergeri Bornemann, 1856

Description: Paripinnate leaves with stout axis, covered by small scales; the biggest fragments are 30 cm long. The pinnae (40-140 x 8-26 mm) are lanceolate to broadly lanceolate with a rounded apex inserted loosely, on a 4-6 mm wide rachis; apically they reduce in dimension (24-40 x 7.5-26 mm). Pinnae are attached oppositely in pairs (occasionally suboppositely or alternately) and arise at an angle of circa 45°. Pinnae are inserted at an acute angle (c. 45°) oppositely, sub-oppositely or alternately to the rachis (5-18 mm wide). The pinna base is constricted on the upper (acroscopic) side and decurrent on the lower (basiscopic) side. In sun-leaves the pinna margin is entire with a narrow lamina; shade leaves have a crenate-lobate to undulate margin, giving rise to lobes with very variable dimensions, but never reaching the midrib. The midrib is distinct (1-3.5 mm wide), secondary and/or tertiary veins rarely visible, arising at an angle of c. 45°, and forking usually at least once.
Epidermal cells are isodiametric, slightly elongated above the veins. Stomata sunken, surrounded by 6-7 subsidiary cells. Epidermal cells of sun leaves are smaller and more often covered and protected by papillae than in shade leaves. Stomata of sun leaves are disposed along bands in intravenal areas of the lower epidermis and rarely on upper epidermis. Stomata of shade leaves are arranged in bands between the veins on the lower surface, and irregularly scattered on the upper surface. Epidermal cells in sun leaves are smaller and more protected by papillae than in shade leaves.
These pinna fragments (max. 94 mm long and up to 26 mm wide) have a margin and arise from the rachis either without any constriction or are slightly constricted at the acroscopic side

Distribution: upper Anisian of the Dolomites (Italy) and China, upper Ladinian of the German Basin (Thuringia, Würzburg, Thale am Harz), Lower-Middle Keuper of Madygen (southern Fergana).

Remarks: To this species belongs as “sun”-leaves also Scytophyllum apoldense (Compter) Linnell 1933.

Peltaspermum bornemannii Kustatscher et al., 2007

Description: Isolated ovuliferous organ of the Peltaspermum type, with a more or less flattened, umbrella-shaped disc, c. 15 – 25 mm in diameter, with a central depression (3.7-5.5 mm diameter) corresponding to the attachment area of the stalk and at least 15 marginal lobes with a marked rib and an acute apex. No attached seeds has been found so far.
The cuticle of the disc is characterized by isodiametric epidermal cells and stomata arranged in irregular rows, consisting of 2 sunken guard cells and 6-8, slightly thickened subsidiary cells without papillae.

Distribution: upper Anisian of the Dolomites (Italy) and Carnian Alps.

Remarks: This organ belongs probably to Scytophyllum bergeri, although no organic connection has been found so far.

?Peltaspermum sp.

Description: The ovuliferous discs arise in pairs (2-3) from the axis (136 x 3.3 mm). These ovuliferous discs are only partially and not very well preserved, but show lobed, umbrella-shaped structures. The best preserved megasporophyll (circa 27-35 mm) shows 5 or 6 different lobes  suggesting that the complete disc consisted of approximately 10-15 lobes.

Distribution: upper Anisian of the Dolomites (Italy).

Ptilozamites sandbergeri (Schenk) Kustatscher et Van Konijnenburg – van Cittert 2007

Description: The leaf fragments (up to 115×15 mm), are characterised by oppositely to suboppositely arranged pinnae. The pinnae are subrectangular to subquadrate with rounded apex and laterally attached to the striate rachis and decrease in size towards the apex (8 x 6-8 mm). The veins arise perpendicularly from the rachis, with a concentration of 7-9 each cm and run parallelly and almost completely without forking up to the margin. The margin is characterized by a narrow vein-free band. The cuticle is thick (upper side c. 4-6µm, lower side c. 2-4µm) and composed of irregular polygonal epidermal cells. Stomata are scattered on the upper side and more frequent on the lower side of the leaf. The epidermal cells of the lower cuticle are more elongate over the veins. Stomata are irregularly distributed and never share subsidiary cells; they are concentrated in bands between the veins, on the upper side of the rachis and almost completely absent on the lower side.

Distribution: Ladinian of Monte San Giorgio (CH), upper Ladinian of the Dolomites, Carnian of the Raibl and Dogna (Julian Alps, Italy).

Remarks: Some material from the Anisian of the Dolomites (Kühwiesenkopf/Monte Prá della Vacca) can be also attributed to this specie; because of the bad preservation and missing cuticle these specimens have been assigned so far only on cf. level.

Sagenopteris sp.

Description: The leaf(let) fragments reach so far a maximum length of 70 mm and a maximal width of 30 mm. The basis of the large, lingulate leaf(lets) is restricted, expanding upwards very quickly. The midrib is 0.5-1.0 mm wide at its base and decreases apically (width 0.5 mm). Fine secondary veins arise from it, which dichotomise a few times and anastomose forming wide meshes; the dichotomising and anastomosing veins are more distant at the base (up to 1 mm) than at the apex (0.5 mm).

Distribution: Anisian of the Dolomites (Italy), Ladinian of the German Basin (Thale am Harz).

Division Cycadophyta

Order Cycadales

Bjuvia dolomitica Wachtler et Van Konijnenburg – van Cittert, 2000

Description: The large, entire-margined leaves have an estimated length on about 60 cm and about 15 cm width with an up to 12 cm long and 5-6 mm wide petiole. The leaf lamina increases gradually from the base, probably the apex is smoothed. Secondary veins arise almost perpendicularly from the rachis, and reach the margin without bifurcating (14-18/cm). The leaves are amphistomatic with rectangular epidermal cells on the upper leaf side, with straight to slightly wavy anticlinal walls. The veins are evidenced by slightly more elongated cells, sttomata occurred only in the intervenal areas, where they are distrbuted in irregular rows. On the lower cuticle the venal and intervenal areas are more clearly defined with stomata arranged in 2-4 rows. The stomata are haplocheilic with slightly sunken guard cells and 4-6 subsidiary cells forming a thickened ring around the stomatal pit.

Distribution: upper Ladinian of the Dolomites (Italy).

Bjuvia thalensis Kustatscher et Van Konijnenburg-van Cittert, 2010

Description: The large lanceolate leaves (up to 200 mm long fragments) are entire margined and reach a maximal width of 150 mm in the middle portion. The lamina is attached to the upper side of the stout, striate rachis (10-12 mm) and, thus, the rachis as seen from above is less than 5 mm wide. Undulations on the lamina might be the result of dessication or may be due to depositional circumstances. The leaf width decreases gradually towards the apex, which is rounded. Fine, but distinct secondary veins arise perpendicularly from the rachis, and reach the margin unforked at a concentration of 25-30/cm.
The leaf is amphistomatic, with one side of the cuticle thicker (3-4 µm) than the other (1-2 µm). Normal epidermal cells are isodiametric (25-40 µm in diameter, varying in shape from square to hexagonal), bearing a central (slightly hollow) papilla; anticlinal walls are straight and distinct. Stomata are more abundant on the thicker (lower?) cuticle. Stomata are arranged in irregular bands and separated by stomata-free zones (circa 150 µm wide), which probably indicate the veins. Stomata are usually oriented almost perpendicular to the supposed direction of the veins, and consist of two guard cells and commonly 4-6 subsidary cells, each of which bear a papilla overhanging the stomatal pit. Stomata sometimes have only two subsidiary cells. Stomatal complexes (including subsidiary cells) are around 50-65 µm in diameter.

Distribution: upper Ladinian of the German Basin (Thale am Harz)

Apoldia tenera (Compter) Zijlstra et al. 2009

Description: Pinnate leaves with one terminal pinna; fragments are up to 410 mm long and 220 mm wide. Pinnae are inserted oppositely to (sub)oppositely and widely spaced on the rachis, arising at 40-80° and never covering the stout rachis (4-10 mm). The pinnae are attached laterally to the rachis and never overlap each other. Each pinna is asymmetrically spatiolate to ovoid in shape with a wedge-shaped base (up 10 mm wide) and rounded truncate apex. The entire margin has sometimes irregular incisions, probably a result of preservation. The largest pinnae occur in the central portion of the leaf (up to 130 mm long and 70 mm wide); they decrease in width and length towards the apex and the base. Numerous veins arise from the constricted base, dichotomizing several times and extending almost completely to the margin; near the base the veins have a concentration of circa 27/cm, more apically 35/cm.
The epidermal cells are isodiametric (square to polygonal), irregularly distributed and covered by papillae. Stomata are arranged in bands between non-stomatiferous areas (probably indicating the veins). Stomata are more common on the lower than on the upper side of the leaf. Near the leaf margin the stomata become less common. The guard cells are sunken, partly covered by papillae from the 4-8 subsidiary cells. The cuticle of the subsidiary cells is generally thicker than that of the normal epidermal cells. Some stomata have only two subsidiary cells and are reminiscent of syndetocheilic stomatal complexes.

Distribution: upper Ladinian of the German Basin (Thale am Harz, Apolda, Nauendorf, Thuringia)

Sphenozamites wengensis Wachtler et Van Konijnenburg – van Cittert, 2000

Description: Simple pinnate leaves with pinnae attached (sub)oppositely to the upper side of the broad rachis. The pinnae are relatively small (up to 50 x 15 mm) with a contracted basis and a rounded apex. The veins fork in the lower and middle part of the pinnae and reache the margin at a concentration of about 16/cm.

Distribution: upper Ladinian of the Dolomites (Italy)

Sphenozamites sp. cf. S. bronnii (Schenk) Passoni et Van Konijnenburg – van Cittert, 2003

Description: The impari-pinnate leaves are characterised by lanceolate-rhomboidal pinnae attached (sub)oppositely to the stout rachis (5-20 mm). The pinnae arise slightly apically inclined (~70-85°) and are about 25-55 mm wide and about 100-250 mm long with a restricted basis (~ 5 mm). The apex is truncate, asymmetric and with smoothed margins. The marked veins are straight, bifurcating in the lower part of the pinnae (vein density 25-30/cm).
The leaves are hypostomatic. In the upper cuticle the epidermal cells are irregular, elongated, more or less arranged in rows. There is no indication of veins, and papillae are absent. In the lower cuticle the veins stomata-free zones are characterised by elongated epidermal cells. Stomata are arranged in irregular rows between the veins. The stomata consist of sunken guard cells surround by 5-7 papillate subsidiary cells. The papillae usually covering the stomatal pit.

Distribution: upper Ladinian of the Dolomites (Italy)

Nilssonia cf. neuberi Stur ex Pott et al. 2007
Description: The leaf fragments are up to 160 mm long and 80 mm wide; however, neither the length nor width of the lamina is complete. From the upper side of the striate rachis (6-11 mm wide) arise segment fragments (up to 70 x 11-16 mm). The segments expand proximally; at the point of attachment to the rachis they measure 17-22 mm. The cuticle fragments show a slightly thicker (upper?) and slightly thinner (lower?) cuticle. Epidermal cells are isodiametric to rectangular. The stomata are haplocheilic.

Distribution: upper Ladinian of the German Basin (Thale am Harz, Apolda, Nauendorf, Piffelbach, Baden)

Dioonitocarpidium moroderi (Leonardi) Kustatscher et al., 2004

Description: The species is composed by two macrosporophyll apical fragments (part and counterpart) of 90 mm length and 25 mm width with a 5 mm wide rachis with longitudinal striae. Only the sterile part of the sporophyll has been preserved, the basal fertile part is missing. The lamina segments (10-20 x 2 mm) arise at an angle of c. 70°.

Distribution: upper Ladinian of the Dolomites (Italy)

Dioonitocarpidium pennaeformis (Schenk 1864) Rühle von Lilienstern 1928

Description: The (macro)sporophylls are preserved only as fragments (65 mm long and 20 mm wide). The axis (3.5 mm at the base, 1-2 mm at the apex) is characterised by one (or two) central furrow(s). The apical part of the (macro)sporophylls is pinnate, the lateral appendices are lanceolate, inserted (sub)alternately and inclined apically (slightly falcate). They are largest in the middle portion (8.5-11.0 x 1.0-1.5 mm), and decrease gradually towards the base and noticeably towards the apex (4.0-6.0 x 0.5-1.0 mm), and are characterised by a central thickening.

Distribution: upper Ladinian and Carnian of the German Basin (Thale am Harz, Frankonia, Thuringia, near Stuttgart and Darmstadt).

?Order Bennettitales

“Pterophyllum jaegeri” Brongniart, 1828

Description: The leaf fragments are up to 33 cm long and 6 cm wide with a ca. 4 cm long petiole. The pinnae are linear with a truncate tip (up to 40-50 mm long and ca. 4 mm wide); they are inserted perpendicularly on the 4 mm wide axis. The venation is parallel.

Distribution: upper Ladinian of the Dolomites (Italy)

Order indet.

Taeniopteris kelberi Kustatscher et Van Konijnenburg-van Cittert, 2010

Description: Simple lanceolate leaves of almost 500 mm lenght and 20-28 mm wide in their middle portion and basally 20 mm. The margin is entire and may be somewhat undulating. At the base the leaf restricts to a narrow lamina (petiole, 11 mm), the attachment is slightly crescent-shaped. The apex is acute. The lamina is inserted to the upper part of the striate, stout rachis (8 mm at the base reducing at the apex to 2 mm).Veins arise almost perpendicularly (80°) from the rachis at the central part of the leaf, near the apex under a smaller angle (50-60°), bifurcate rarely at the base of the lamina and reach the margin with a density of 20-25/cm.

Distribution: upper Ladinian of the German Basin (Thale am Harz, Schleerieth, Apolda, Eubingheim), Carnian of Neue Welt (CH), possibly Anisian of the Dolomites (I).

Taeniopteris sp.

Description: The leaf fragments are characterised by a entire, parallel margin, the strong midrib, the distinct, parallel secondary veins which arise perpendicularly from the midrib and do almost never fork.

Distribution: upper Ladinian of the Dolomites (Italy)

Division Coniferophyta

Elatocladus cassinae Stockar et Kustatscher, 2010

Description: Unbranched shoots, up to 165 mm long and 40 mm wide. The axis is proximally up to 3.5 mm broad, but decreases to 1 mm wide at the tip. The leaves are arranged in a loose helix. The leaves are dorsi-ventrally flattened, coriaceous, linear, with a rounded apex, and a constricted base. They are up to 30 mm long and 2.5-3 mm wide but decrease in length both toward the tip and base (length around 8-12mm). The leaves are coriaceous and probably were vascularized by 6-8 veins. Branching has not been observed among the shoot fragments. In the young shoots the axis is still very short, with leaves inserted in a dense helix. The leaves in these shoots are ~12 mm long and 2 mm wide (more apically about 3.5 x 1 mm) and have a smoothed tip.

Distribution: Ladinian of Monte San Giorgio (CH).

Elatocladus sp.

Description: The only shoot fragments is 17 cm long and 7 cm wide. The leaves are spirally arranged, flat, 20 – 35 mm long and 15 – 20 mm wide:. The apex is rounded and the basis contracted. No midrib can be distinct.

Distribution: upper Ladinian of the Dolomites (Italy)

Pelourdea vogesiaca (Schimper et Mougeot) Seward 1917

Description: Herbaceous conifer with elongated to lanceolate leaves with smoothed apex of up to 50 cm lenght and 15-22 mm width. The leaves restrict slightly basally but attach to the axis with an enlarged basis. The veins are distinct, numerous, parallel and bifurcate rarely.

Distribution: Anisian of the Vosges (France), upper Ladinian of the Dolomites (Italy)

Voltzia dolomitica Wachtler et Van Konijnenburg – van Cittert, 2000

Description: This coniferal shoots have the ultimate shoots arranged in one plane, arising with an angle of ~ 45˚. The leaves are falcate to triangular in shape with a more or less acute apex; they are spirally arranged. The overlap partially each other. The leaves are hypostomatic. The adaxial cuticle has large, irregular, isodiametric epidermal cells, in the abaxial cuticle the epidermal cells are smaller. Stomata are arranged in short, irregular rows, never share subsidiary cells. The guard cells re sunken, surrounded by a ring of 4-7 subsidiary cells. The male cones consists of an axis with spirally arranged sporophylls. No pollen sacs are known. The female cones consist of spirally arranged bract/ovuliferous scale complexes, the seeds are 3 mm long and round.

Distribution: upper Ladinian of the Dolomites (Italy)

Voltzia ladinica Wachtler et Van Konijnenburg – van Cittert, 2000

Description: This species is characterised often only by small shoot fragments. The ultimate shoots arise at angles of 60˚. The falcate leaves are spirally arranged, 1-2 mm wide and 4-10 mm long but with up to 25 mm long heterophyllous leaves. The leaves are amphistomatic with less stomata on the adaxial than on the abaxial side. The epidermal cells are more or less rectangular between stomatal rows and more isodiametric within stomatal rows. The stomata are quite densely placed within the rows but never share subsidiary cells. The guard cells are sunken, surrounded by 6-10 subsidiary, which can sometimes be protected by papillae. The male cones (50 x 1 mm) show spirally attached microsporophylls, the female cones consist of 6-8 mm long bracts and 8-10 mm long and 8-10 mm wide ovuliferous scales with five lobes with 2 seeds per scale.

Distribution: upper Ladinian of the Dolomites (Italy)

Voltzia pragsensis Wachtler et Van Konijnenburg – van Cittert, 2000

Description: This conifer has penultimate and ultimate shoots in one plane with the ultimate shoots arising at angles of about 30-45°. The leaves are triangulate with acute apex, spirally arranged, closely attached to the stem The cuticle of this species is still unknown and well as the male cones. The female cones consist spirally arranged bract/scale complexes.

Distribution: upper Ladinian of the Dolomites (Italy)

Voltzia zoldana Leonardi, 1953

Description: The so far only sample is 58 mm long and 13 m wide with a curved axis and 7-8 x 1.5-2 mm slightly falcate leaves.

Distribution: upper Ladinian of the Dolomites (Italy)

Indet.

Phylladelphia strigata Bronn, 1858

Description: Spatulate to tongue-shaped leaf-like structures (up to 96 x 44 mm long), with a broad basal attachment area (c. 18-30mm). The leaf lamina width expands slowly from the basal part of the leaf to the middle part, from where it restricts again, ending in an acute apex. In the central part, a “midrib” is visible, crossing the whole leaf and becoming more distinct in the basal half of the leaf, working almost like a symmetry axis. Laterally to it several (4-5) ribs (costae) run parallel to it at a distance of 1-2 mm, up to the apical margin. In the distal part, where the leaf is broader, 2-3 additional ribs cross the entire lamina from one margin to the other. Basally the leaf appears to be thicker, and the attachment area is almost straight. In the basal central part and near the margin of the apex the ribs are sometimes connected by very small wrinkles (1-5 mm long); in basal fragments the ribs are much more prominent. When the leaves of leaf fragments cover each other partly they seem slightly more rhomboidal than in the isolated specimens.

Distribution: Carnian of Raibl/Cave del Predil (Julian Alps, Italy)

Evelyn Kustatscher

Here you can find more about the above mentioned taxa

Kustatscher E. & Van Konijnenburg-van Cittert J.H.A., 2008. Considerations on Phylladelphia strigata Bronn from the historical Raibl flora (Carnian, lower Upper Triassic, Italy). – Geo.Alp, 5: 69-81.

Kustatscher E. & Van Konijnenburg-van Cittert J.H.A., 2010. Seed ferns and Cycadophytes from the Triassic Flora of Thale (Germany). Neues Jahrbuch für Geologie und Paläontologie, 258(2): 195-217.

Kustatscher E. & Van Konijnenburg-van Cittert J.H.A., 2005. The Ladinian Flora (Middle Triassic) of the Dolomites: palaeoenvironmental reconstructions and palaeoclimatic considerations. – Geo.Alp, 2: 31-51.

Kustatscher E. & Van Konijnenburg-van Cittert J.H.A., 2008. Lycophytes and horsetails from the Triassic Flora of Thale (Germany). – Neues Jahrbuch für Geologie und Paläontologie, 250(1): 65-77.

Kustatscher E., Hemlsey A.R. & Van Konijnenburg-van Cittert J.H.A., 2009. Lugardonia paradoxa, a new fertile strobilus with unknown affinities from the Anisian flora of Kühwiesenkopf, the Dolomites, Italy. – Review of Palaeobotany and Palynology, 156: 90–97.

Kustatscher E., Meller B. & Van Konijnenburg-van Cittert J.H.A., 2006. Old treasures newly discovered: Scytophyllum bergeri from the Ladinian of the Dolomites in the historical collections of the Geologische Bundesanstalt Wien. – Geo.Alp 3: 47-54.

Kustatscher E., Wachtler M. & Van Konijnenburg-van Cittert J.H.A., 2004. Some additional and revised taxa from the Ladinian Flora of the Dolomites, Northern Italy. – Geo.Alp, 1: 57-70.

Kustatscher E., Wachtler M., & Van Konijnenburg-van Cittert J.H.A., 2010. Lycophytes from the Middle Triassic (Anisian) locality Kühwiesenkopf (Monte Prà della Vacca) in the Dolomites (Northern Italy). – Palaeontology, 53(3): 595-626.

Kustatscher E. & Van Konijnenburg-van Cittert J.H.A., 2007. Taxonomical and palaeogeographic considerations on the seedfern genus Ptilozamites. – Neues Jahrbuch der Geologie und Paläontologie Abhandlungen, 243(1): 71-100.

Kustatscher E., Wachtler M. & Van Konijnenburg-van Cittert J.H.A., 2007. Horsetails and seedferns from the Middle Triassic (Anisian) locality Kühwiesenkopf (Monte Prà della Vacca) in the Dolomites (Northern Italy). – Palaeontology 50/5: 1277–1298.

Stockar R. & Kustatscher E., 2010. The Ladinian Macroflora from the Cassina beds (Monte San Giorgio, Switzerland): a first report. – Rivista Italiana di Paleontologia e Stratigrafia, 116(2): 161-176.

Van Konijnenburg-van Cittert J.H.A., Kustatscher E. & Wachtler M., 2006. Pteridophytes from the Anisian locality Kühwiesenkopf (Dolomites, Northern Italy). – Palaeontology, 49(5): 943-968.

Wachtler M. & Van Konjinenburg-van Cittert J.H.A., 2000a. The fossil flora of the Wengen Formation (Ladinian) in the Dolomites (Italy). – Beiträge zur Paläontologie, 25: 105-141.