LYCOPHYTA from the EARLY-MIDDLE TRIASSIC (ANISIAN) PIZ DA PERES (DOLOMITES - NORTHERN ITALY) by MICHAEL WACHTLER

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LYCOPHYTA FROM THE EARLY-MIDDLE TRIASSIC (ANISIAN) PIZ DA PERES (DOLOMITES - NORTHERN ITALY) by MICHAEL WACHTLER

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 1 DOLOMYTHOS

Published by the Dolomythos Museum, Innichen, South Tyrol, Italy January 2012

Dolomythos includes results of original research on systematic, evolutionary, morphological and ecological biology, including paleontology. Syntheses and other theoretical papers, based on research, are also welcome. Dolomythos is intended primarily for papers by the staff of the Dolomythos Museum or on research using material in this Museum.

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Print: Technolab Communication srl, TLAB Editrice Viale Pecori Giraldi, 20/B 36061 Bassano del Grappa (VI) - IT www.technolab.it Wachtler M.: The Genesis of plants Preliminary researches about the Early-Middle Triassic Fossil Floras from the Dolomites - A Compendium; ISBN 978-88-904127

Please cite this articles as:

Wachtler, M., (12/2011): Lycophyta from the Early-Middle Triassic (Anisian) Piz da Peres (Dolomites - Northern Italy), in “The Genesis of plants Preliminary researches about the Early- Middle Triassic Fossil Floras from the Dolomites - A Compendium” Dolomythos, Innichen. p. 165 - 211

Dolomythos, 2011 2 LYCOPHYTA FROM THE EARLY-MIDDLE TRIASSIC (ANISIAN) PIZ DA PERES (DOLOMITES - NORTHERN ITALY) by Michael Wachtler P. P. Rainerstrasse 11, 39038 Innichen, Italy, e-mail [email protected]

Abstract Although the most dominant group of plants in the Carboniferous, Lycophyta currently inhabit the shadows of more successful plants. Therefore, discoveries of new missing links not only throw light on the evolution of this important family, but also help us understand climatic and paleoecological cataclysms in the past. Four different genera containing five new species from the Early-Middle (Anisian) Triassic age in the Eastern Alps are described here. They were well preserved and enabled the determination of their preferred living habitat and symbiosis to other plants and animals due to their abundance and detailed stratigraphical classification. Surprisingly, the most abundant form was the sub-arborescent club moss Lycopia dezanchei gen. et sp. nov., exhibiting dichotomous branching and creeping root horizons. They could act as an intermediate form between Carboniferous Lepidodendrales and extant Lycopodiales. The fertile morphogenus was the homosporous Lycopodostrobus gaiae gen. et sp. nov. Extant Sellaginella-like forms were represented by Selaginellites leonardii sp. nov., with its heterosporous strobili and Selaginellites venierii sp. nov., with characteristic anisophyllous shoots. Another group of extant Lycophyta, the Isoetales, were represented by Isoetites brandneri sp. nov., resembling mostly extant Isoëtes and Lepacyclotes bechstaedtii sp. nov., with its bulbous corpus. Never recorded in these Anisian strata was the most enigmatic Early Triassic lycophyte, Pleuromeia sternbergii. Surprisingly, in the near but slightly older Werfen strata of the Carinthia, it still constitutes the dominant plant assemblage. This work compares the living conditions in the Early-Middle Triassic age and develop information and solutions about the rapid extinction of the historic and often discussed Pleuromeiales.

Online: December 2011. Key words: fossil lycophyta, Dolomites, Italy, Early-Middle Triassic, Anisian.

Remarks and annotations were based on my older This study summarises fifteen years of my archives and a large part of these studies lonely and dangerous work in the mountains lacked further cuticle analyses and pollen of the Eastern Alps in which I made a lot preparation. I apologise for this and hope to of interesting discoveries. In long-lasting make a revised version for when my country studies, cataloguing and descriptions, I learns to appreciate science. tried to shed more light on a lost world. I Although agreeing with some parts of the thought that all this could be useful for content of ”Lycophytes from the Middle humanity to understand more about our Triassic (Anisian) locality Kühwiesenkopf past. Unfortunately, one day, the authorities (Monte Prà della Vacca) in the Dolomites confiscated and removed all my specimens (northern Italy)“ (2010), this publication from my house, damaging a large part of was, however, never fully authorised by Mi- my research. My computers, photos and chael Wachtler and edited under unaccepta- notations were also sequestered and taken ble pressure, with several formerly-agreed away, the cataloguing labels changed conditions of a reciprocal contract not be- to cover the traces of my findings and ing maintained. Therefore, in my opinion, studies, and my name cancelled from my the right authorship for this work has to be collection and anonymised. Many photos read as MICHAEL WACHTLER, EVELYN KUS-

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 165 Fig. 1: Fossil-plant bearing localities in the Eastern Southern Alps. Orange: carboniferous fossil points; yellow: Permian fossil points; red: Early Triassic - Olenekian fossil points; and green: Early-Middle Triassic (Anisian) fossil points.

TATSCHER, JOHANNA H. A. VAN KONIJNEN- Paleobiology and paleoecology BURG-VAN CITTERT or it has to be seen as an illegal publication or one without agree- During the Early Triassic age, the Eastern- ment from all authors. Michael Wachtler re- Southern Alps were located near a beach grets the pronounced interdiction of research close to a shallow water basin. The high by the authorities, as well as police force, at- sedimentation rate suggests frequent storm tempting to exercise his intellectual research events (hurricanes). After intense volcanic rights. activities in the Lower Permian, their first, not yet, marine deposited cycles contributed to the Gröden-Formation during the Upper Localities Permian. During the following Early Triassic The fossil plant-rich Piz da Peres area covers age, the now oceanic-influenced sediments about 8 km of the Northern part of the Do- contributed to the Werfen-Formation. They lomite Mountains, starting from the western consist of a strongly varying sequence side of Piz da Peres over the Ladinian village of mixed terrigenous siliciclastic and of St. Vigil and reaching the Kühwiesenkopf carbonatic lithofacies (BRANDNER ET over the hamlet of Prags. It traverses sever- AL., 2009). Since the 19th century, the al mountains, which the native people refer Werfen-Formation has been divided into to as a big mountain sundial. Starting from two sections: one beginning from the the Kühwiesenkopf, we have the ”Zehner” Permian-Triassic boundary subdivided into (Tenner) “Elfer” (Elevener)and “Zwölfer” Tesero-, Mazzin-, Andraz-, Seis-Member and (Twelver), also called Hochalpenkopf, the Gastropodenoolith, which are characterised “Einser” (Oner), also called Maurerkopf, and by an abundance of the scallop-like bivalve the “Zweier”(Twoer). Other mountains with mollusc Claraia especially found in the fossil plant lenses are the Flatschkofel and Seis-Member, and the other composed of the Drei-Fingerspitze. The main fossil plant Campill-, Val Badia-, Cencenighe- and San lenses lie in the sometimes-exposed open Lucano-Member, showing an increase in rock slopes that are occasionally interrupt- terrigenous input. Hemipelagic terrigenous ed by shrubs and woods in which the lay- and subordinate carbonate sediments ers cannot be traced. Other places impor- were deposited continuously, while storm tant for the comprehension of Early Triassic events also sometimes delivered a lot of lycophyte evolution lie in the neighbouring plant debris. These basin sediments, with a Carinthian Mountains, where the Induan/ conspicuous clastic input, continue until the Olenekian Werfen layers hold a rich Pleu- Anisian Dont Formation (=Pragser Schichten romeia-dominated flora. Usually, the recov- sensu PIA, 1937). The fine silty marine ered specimens are well preserved and give Werfen layers reach 500 metres in thickness a deep insight into the ecosystems after the in the Northern Alps, and 200 metres in the Permian-Triassic boundary. Southern Alps (BRANDNER ET AL., 2009).

Dolomythos, 2011 166 Kühwiesenkopf Hochalpenkopf Maurerkopf Zehner Elfer Zwölfer Einser Zweier

Flatschkofel Drei-Fingerspitze

Piz da Peres

Fig. 2: The Piz da Peres fossil area. It covers about 8 km from Piz da Peres to Kühwiesenkopf, around a mythical sundial of the native people. Zehner = Tenner, Elfer = Elevener, Zwölfer = Twelver, Einser = Oner, Zweier = Twoer. The main plant fossil lenses lie on open rock.

The Campill-Val Badia Member in the Eastern and Selaginellites venierii) are common. In Southern Alps could be correlated with the spite of intensive research, a total absence Middle Buntsandstein, or more detailed of Pleuromeia has to be accepted. The gi- Volpriehausen-Hardegsen-Formation of ant horsetail Equisetites mougeotii has been Germany (about 249.7 to 245.9 ± 0.7 Mya recovered in the Werfen-Formation (Campill- old) stretching from Induan to Olenekian. Val Badia Member), as well as in the follow- While the South Tyrolean Werfen-Formation ing successions from Anisian to Ladinian and has not revealed to date any determinable Carnian, showing only minimal changes in plant species, the nearby Gailtaler Alps its aspect and reproduction cycles. and other mountain chains in Carinthia Most Carinthian Triassic localities are display well fossilised floras (Ulrichberg- situated inside crowded vegetation below Klagenfurt, Wunderstätten-St. Paul, Nötsch- the forest line (in the Dolomites, mainly Villach, Golsernock-Stockenboi and Laas- all fossil plant-bearing horizons lie over Kötschach-Mauthen). They are characterised the forest limit); therefore, only newly by a dominance of the lycophyte built forest trails enable the exploration of Pleuromeia sternbergii, accompanied by underlying rocks. Sufficient Early Triassic the sphenophyte Equisetites and some plant layers have been opened over the isolated Voltzia remains (THIEDIG & KABON, years and the excavated fossil slabs 2011) and can therefore be defined as a vegetation-rich, but species-poor flora. The locality Bleiriese-Golsernock, discovered in 2011 by Georg Kandutsch, Helmut Prasnik and Michael Wachtler, exhibits abundant and well-preserved Pleuromeia sternbergii material, accompanied by Equisetites mougeotii. It is remarkable that other plant groups like ferns, seed ferns, conifers or cycads are till now missing. The sub-arborescent lycophyte Pleuromeia, dominant worldwide during the Early Triassic period, seems to have been effectively extinct during the Olenekian- Anisian boundary (RETALLACK G., 1975). Ecological support is given by the slightly younger Anisian Formations of the Dolomites where rich lycophyte assemblages (Lyco- pia dezanchei, Isoetites brandneri, Lepacy- clotes bechstaedtii, Selaginellites leonardii Exploring the newly discovered Olenekian fossilised- plant point Bleiriese-Golsernock in the Gailtaler Alps.

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 167 Open lying isolated tree-trunks, root-horizons and ripple marks from Kühwiesenkopf - main plant layer.

give a good insight into ancient habitat the almost exploding diversity of the Anisian systems. All the findings mainly show landscapes that followed a little later. The predominance of the lycophyte Pleuromeia theory of the commonly thought vegetation- sternbergii, identical to the Central German poor and arid Permian landscapes Buntsandstein one. It could be assumed that has to be revised. New discoveries in vast areas of the Northern hemisphere in the Dolomites (Bletterbach, Tregiovo) the Early Triassic were colonised by only one suggest an extraordinary abundance of plant species. The Pleuromeia monocultures humidity-loving horsetails (Equisetites), are effectively strange when compared to ferns and seed ferns (Callipteris),

Dolomythos, 2011 168 Kühwiesenkopf - Main plant lenses conifers, cycadophyta (Taeniopteris) and total of 214 metres, of which the middle 52 gingkophyta (Sphenobaiera). Therefore, metres are covered by a dense forest where comparisons with today’s Okavango-delta no rocks protrude. On the right side, there with its seasonal floodings, salt islands and are plant layers 42.5 metres long and one- associated salt-resistant plants, and semi- metre thick. There is also the presence of arid zones facilitate the understanding of the only terrestrial reptile, Megachirella this ancient landscape. wachtleri (RENESTO, 2003). The lenses hold a variegated flora containing equal propor- The Kühwiesenkopf lycophyte paradise tion of lycophyta, conifers, cycadophyta, ferns and seed ferns. The Anisian Dont Formation, consisting of a After the overgrowing patch of forest, the carbonate-terrigenous sequence more than layers crop out for 119.5 metres, interrupt- 200 metres thick, reaches the most distin- ed once by a geological fault that elevates guished development in the Braies Dolo- the lenses 25 metres high. These beddings mites. The main plant-bearing horizon on are characterised by a rich fish assemblage Kühwiesenkopf lays circa 75 metres above belonging to five different families (Dipter- a massive carbonate platform attributed to onotus, Saurichthys, Bobasatrania, Gyro- the Bythinian-Anisian Gracilis Formation (DE lepis and coelacanths), of which the first ZANCHE ET AL. 1992; BROGLIO LORIGA ET three genera have never or only rarely been AL. 2002; VAN KONIJNENBURG–VAN CIT- recorded in the Early Triassic (TINTORI A. ET TERT ET AL. 2006). AL. 2001). Most of the fish skeletons have One club moss-rich layer on Kühwiesen- been found in a narrow, only three-metre kopf has enabled, like in a few other places long section in the middle. The preservation worldwide, an inimitable insight into a con- of invertebrates in this area can be relat- tiguous beach more than 240 million years ed to the very rapid burial events made by old. This main lens can be prosecuted for a

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 169 heavy storms and flood deposits originating by Isoetites brandneri, Lepacyclotes from terrestrial domains. bechstaedtii and Selaginellites leonardii. All the fishes seem to be well adapted to live Apart from Isoetites brandneri and near the coastline. Dipteronotus sp., known Lepacyclotes bechstaedtii, which have only from a dozen specimens in Europe, is never been recorded in the other layers one of the best indicators of strong fresh wa- of Kühwiesenkopf, Hochalpenkopf or Piz ter influx in the Middle Triassic. Saurichthys sp., widespread in the Thetydalean ocean in the Triassic period, was an able predator. It is generally encountered in sediments of ma- ______rine environments, as well as sporadically in rocks of brackish or continental origin. Boba- 12,5 m satrania sp., found nearly worldwide from ______Madagascar to Cina, Greenland, Svalbard and Canada in Early Triassic beds, was another 15 m predator, but in comparison to Saurichtys sp., Lycophyta point it was a steady swimmer, scrunching with great accuracy its preys with a battery of Megachirella point minute teeth. Gyrolepis sp. has been found in sediments of marine coastal environ- ______ments, especially in the Ladinian Fossillag- erstätte of Monte San Giorgio. The skeletons of the coelacanths recovered allow, with cau- 15 m tion, the insertion of the species Heptanema ______(known from the Western Alps) in the Middle Triassic. The ichthyofauna attests a marine depositional environment with a very strong Wood influx from nearby land, testified by a huge amount of plant remains. In the whole section, apart from the 52 m ubiquitous ferns, seed ferns, conifers Wood and abundant cycadophyta species, there is the most complete club moss ______association. The most abundant is the sub- arborescent Lycopia dezanchei, followed 37,5 m

Cycad-point

______Tectonic fault ______3 m 6 m______

Lycophyta point

Ripple 17 m Fish - bassin 32 m 24 m ______Fish - bassin ____

Kühwiesenkopf: Lower lens layer demonstrating the Megachirella finding point, the plant lenses (especially lycophyta) and the fish-basin. Total length: 214 metres.

Dolomythos, 2011 170 da Peres, Lycopia dezanchei is found Materials and methods everywhere and in large quantities, whereas Selaginellites leonardii is much To date (2011), the Lycophyta fossil record more abundant in the Wachtler gorge in is characterised by: the west side of Piz da Peres, with another Pleuromeia sternbergii (about 200 species) species, Selaginellites venierii, holding only known from the Olenekian strata in anisophyllous foliage. Carinthia All in all, we have to do it with an ancient Lycopia dezanchei (about 200 pieces) marine depression probably near a highly Lycopodostrobus gaiae (about 10 pieces) saline coastline or also influenced by geo- Isoetites brandneri (about 50 pieces) thermal events such as hot springs or heat- Lepacyclotes bechstaedtii (about 10 pieces) ed soils offering the best requirements for Selaginellites leonardii (about 20 pieces) lycopods to predominate over other plant Selaginellites venieri (2 pieces) families. Whether several storm events burst the plants in this depression or over- Repository flooding at the coastline will be determined Most of the macrofossil plant collection is by some fossilised root horizon. stored at the Natural History Museum Südti- Just below this main Kühwiesenkopf horizon, rol, Bozen. The remainder of the collection a bank of nodular and bioclastic limestone is in the Museum Dolomythos, Innichen, or contains a rich marine epifaunal assem- in the collection of Georg Kandutsch, Arriach blage, dominated by brachiopods (Puncto- (Carinthia). spiriferella fragilis, Tetractinella trigonella, Specimens were photographed under natu- Mentzelia mentzelii, Spiriferina-paläotypus ral light using Nikon D200, Lenses AF MICRO and Angustothyris angustaeformis) and bi- NIKKOR 60 mm 1:2.8 D and AF-S NIKKOR valves (Mysidioptera cainalloi, Neomorpho- 17-35 mm 1:2.8 D for larger pieces. The tis compta, Entolium discites, Newaagia digital images were processed using Adobe noetlingi, Pleuromya cf. musculoides, P. Photoshop CS version 12.1. brevis and Neoschizodus? sp. Others, such as Plagiostoma striatum, Pseudoplacunopsis fissistriata and Prospondylus comptus have Explanation of the table on the next pages: been found only in the plant bearing depos- Like an expulsion from the Garden Eden, we have the it, or in fossiliferous horizons located in the biggest modifications in plant kingdom at the Carbonif- upper part of the formation (Parallelodon erous-Permian boundary. Surprisingly, mainly all plant esinensis, Pteria? sp., Entolium kellneri and families developed during this time. Chlamys (Praechlamys) cf. schroeteri) (PO- In the Permian, we encounter an impoverished and decreasing number of plant families. SENATO, 2008). From the Early Triassic onwards, we have a general re- After this fish- and club moss-rich section, covery and mutation of all plant associations. this lens becomes abruptly increasingly From a cycadophyte-related group originates the proto- thinner to the end in well-preserved ripple- angiosperms as the true cycads. In Early Triassic, we marks that wedge into the mainly sterile have the division of “feather-sporophyll”-bearing cycads and plant-free horizons. Although other like extant Cycas and strobili-holding cycads like Bowe- layers in Kühwiesenkopf and Piz da Peres nia, Ceratozamia, Dioon, Encephalartos, Stangeria and Zamia. are rich in club mosses, they never reach The conifers split into the araucarian line (Voltziales) the abundance of this lens. Furthermore, and pinaceae-cupressaceae line (Alpiales). extant club moss communities are known Seed ferns were dominated for a long time by Peltas- from stress-conditioned biocenosis, like the permum-Scytophyllum-Thinnfeldia and a Sagenopteris Tepuis of Venezuela or some high-plateau line. Both seemed to disappear at the Jurassic-Creta- in the Andes. Such marine-near and also ceous boundary without leaving any extant descendants. different genera-rich lycophyte zones are Equisetaceae diminished from the Triassic period to unknown nowadays. The richness of all present. plant families in this Anisian stratum, from Lycophyta remained all the time mainly unmodified in conifers to ferns and cycadophyta, does not the Isoetaceae and Selaginellaceae, whereas Lycopia allow extraordinary conclusions to be made met the same fate as Equisetites. It was reduced to about global cataclysms or totally different surviving as Lycopodiales. The Pleuromeiales became extinct in a very short time after the Early Triassic. living conditions during that period.

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 171 Dolomythos, 2011 172 Araucariacea Gingko Lycopodiacea Isoetacea Ferns Feather Strobilus angiosperms Equisetacea Pinaceae cycads cycads Cupressaceae Selaginellacea

? ? Bjuvia simplex Sagenopteris rhoifolia Equisetites Voltzia Alpia Thinnfeldia arenacea dolomitica ladinica rhomboidales

Nilssonia lunzensis

Osmundaceae Bjuvia Marattiales dolomitica Lindsaeacea

Lycopia Selaginellites Isoetites Alpia Sagenopteris Bjuvia Voltzia Schizolepis Pizperesia venieri brandneri keilmannniiolangensis rhaetia Selaginellites Isoetes Pleuromeia agordica Nilssonia Equisetites Scytophyllumbraiensis leonardii beestoni sternbergii mougeotii bergeri ? Pseudo- Spheno- ? Taeniopteris voltzia Peltaspemum eckardtii baiera martinsii Ortiseia Ullmannia

? Proto angiospermales

Peltaspermum Walchia conferta Isophyllous Taeniopteris Selaginel- laceae multinervis Anisophyllous Selaginella- ceae Taeniopteris Callipteris jejunata Selaginellites Neuropteris Isoetites Lepidodendron Sigillaria Calamites Cordaites Pecopteris Alethopteris Odontopteris

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 173 SYSTEMATIC PALAEONTOLOGY cally four-lobed and the stem in the lower Division LYCOPHYTA part is covered with traces of scars from Order PLEUROMEIALES fallen leaves. Foliage in the upper part is Family PLEUROMEIACEAE lanceolatic, displaying a prominent mid-vein and attached on their enlarged basis to the GENUS Pleuromeia CORDA in GERMAR stem. Cones are well defined, sitting termi- nally and solitary on the axis. Sporophylls 1852 have a circular to oval lamina and a large The lycophyte Pleuromeia constitutes one globose sporangium covering nearly all the of the best-described and most interpreted adaxial surface. They contain either micro- plants in paleobotany. It was found in the spores or megaspores. first decade of the nineteenth century in the German Buntsandstein and described by the Discussion Czech botanist Karl Joseph Corda in 1852. This description tried to work out only new After that, a lot of similar species or subspe- stratigraphical and paleoecological phenom- cies were recovered around the world from ena from the not so well-known fossil de- France, Spain and Austria to Russia, China, posits in the Eastern Alps and left behind Japan, Argentina and Australia. detailed specifications indicating only some references (TAYLOR ET AL. 2009; GRAUVO- Pleuromeia sternbergii (MÜNSTER GEL-STAMM, 1993; GRAUVOGEL-STAMM & 1842), CORDA in GERMAR 1852 LUGARDON, 2001; RETALLACK, 1975). 1839 Sigillaria sternbergii MÜNSTER, P. 47, PL 3, FIG. 10 The consistent Pleuromeia findings from Carinthia comes from the Induan-Olenekian 1852 Pleuromeia sternbergii (MÜNSTER) CORDA in GERMAR, p. 184 Werfen-Formation (Campill-Val Badia-Mem- 1904 Pleuromeia sternbergii (MÜNSTER) CORDA in PO- ber), corresponding to the German Mittlerer TONIE, n. 38 Buntsandstein (THIEDIG & KABON, 2011; 1904 Pleuromeia sternbergii (MÜNSTER) CORDA in KANDUTSCH & WACHTLER, 2011). In con- MÄGDEFRAU, p. 119, pl. III-VII trast to the continental German deposits, 2011 Pleuromeia sternbergii (MÜNSTER) CORDA in they belong to a marine environment with THIEDIG & KABON, p. 203 – 224 accompanying casts and impressions of sea- 2011 Pleuromeia sternbergii (MÜNSTER) CORDA in shells like Neoschizodus and Bakevellia sp. KANDUTSCH & WACHTLER The horizon newly discovered by Georg Kandutsch, Helmuth Prasnik and Michael Type localities Wachtler in 2011 on the forest trail of Carinthia: Bleiriese-Golsernock, Laaser Wald Bleiriese-Golsernock near the Upper – Kötschach-Mauthen, Ulrichberg - Kla- Carinthian locality of Stockenboi revealed genfurt, St. Paul Mountains, Rottensteiner an especially rich and well-preserved Kreuz, Wunderstätten, Dobratsch, Waidisch- Pleuromeia sternbergii vegetation, bach accompanied by the giant horsetail Equisetites mougeotii. It was interesting Type horizon and age that other plant groups like ferns, seed ferns, cycads and conifers were absent. Lower Triassic, Induan-Olenekian, Werfen About 200 specimens were recovered from Formation the locality Bleiriese-Golsernock and some from the Laaser Wald near Kötschach- Repository Mauthen, including typical four-lobed Georg Kandutsch Collection, Arriach - Carin- rhizomes, stems bearing leaf scars, leaves thia, Museum Dolomythos - Innichen (South and cones with mega- and microsporophylls. Tyrol) They confirmed that we have to do it with a mainly in pure stands growing plant. Diagnosis The location near the ancient Tethys Ocean confirmed once more that the Pleuromeiales Sub-arborescent lycophyte with unbranched, occupied a wide-ranging habitat, growing erect trunk. Rhyzomorphic roots are typi- as a coastal halophyte as well as an inland

Dolomythos, 2011 174 3 1 cm

1 3 cm

4 1 cm

2 1 cm 5 1 cm

1) GOL 30. Pleuromeia sternbergii. Several stems from juvenile plants. 2) GOL 22. Pleuromeia sternbergii. Part of a lower stem fragment with cleavage impressions. 3) GOL 05. Pleuromeia sternbergii. Typical four-lobed rhizomorphous root. 4) GOL 14. Pleuromeia sternbergii. Internal part of a stem with outstanding leaf fragments from the upper side. 5) GOL 02. Pleuromeia sternbergii. Single leaf with mid-vein.

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 175 xerophyte in the Early Triassic German the Early Triassic, and the creeping and Basin, perhaps also in a desert or stormy dichotomising Lycopiales of Anisian strata environment. Surprisingly, this lycophyte, have a completely different structural which dominated the floras of the lower composition, we could not hypothesise an Triassic, did not survive the Olenekian- apparent successor of Pleuromeia. We can Anisian boundary because a club moss- only vaguely argue that maybe Carboniferous rich vegetation was encountered (Lycopia Sigillaria have some similarities. It seems dezanchei, Isoetites brandneri, Lepacyclotes that the Pleuromeiales came from nothing bechstaedtii, Selaginellites leonardii and and disappeared without leaving any trace. S. venierii) in the slightly younger Anisian This appears strange because in the Eastern- strata around the neighbouring Piz da Peres, Southern Alps, we have a mainly continuous but no Pleuromeia was found. fossil plant record on a small area from the Of interest are therefore the reasons why Carboniferous over Permian and Triassic the outstanding giant club moss Pleuromeia periods. It is dissatisfying to establish became extinct in a geologically short time, Pleuromeia as a pioneer plant after the whereas the giant horsetail Equisetites, which Permian-Triassic cataclysm. A lot of other occupied the same ecological niches, passed plants like the Peltaspermales, Coniferales, unchanged on to the following periods. Since Cycadophyta and Equisetales survived, also quillworts like Isoetites and Selaginella- mainly unmodified by these supposedly all- like club mosses have been recorded from destroying catastrophes.

f a c e

Pleuromeia sternbergii. (a) Entire plant (b) leaf-arrangement on the upper side, (c) leaf-arrangement on the lower side, (d) internal part of the stem, (e) root system, (f) entire strobilus and (g) megasporophyll.

Dolomythos, 2011 176 7 5 mm

5 mm 5 mm

8 9

1 cm 6 10 1 cm 11 5 mm

6) GOL 01. Pleuromeia sternbergii. Strobilus with attached part of the stem. 7) GOL 01 Pleuromeia sternbergii. Detail of the cone with macrospore. 8) GOL 22 Pleuromeia sternbergii. Single sporangium. 9) GOL 22 Pleuromeia sternbergii. Single sporangium. 10) GOL 04 Pleuromeia sternbergii. Cone with microspores. 11) GOL 21 Pleuromeia sternbergii. Cone with microspores.

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 177 SYSTEMATIC PALAEONTOLOGY Type localities Division LYCOPHYTA Order ISOETALES Prantl, 1874 Kühwiesenkopf Family ISOETACEAE Reichenbach 1828 Type horizon and age Genus Isoetites MÜNSTER, 1842 Lower to Middle Triassic, Anisian, Pelson-Il- lyrian The Isoetaceae are a group of widespread lycophytes currently occurring in temperate Diagnosis regions, as well as at high altitudes in tropical regions. Known as quillworts, these Low-growing herbaceous lycophytes, which perennial plants, superficially resembling resemble extant quillworts. Plant displays grasses, grow mostly submerged in shallow a small corm-like stem, leaves and rhizo- water or in wet soils or flood plains. A morphous roots are attached close together. habitat of permanent or temporary lakes, Foliage is spreading and protruding, single riverbanks and saliniferous lagoons could leaves are narrowly erect and ending as lan- also be true for their ancestors. ceolate. Heterosporous sporangia are em- The generic name Isoetites, for fossilised bedded on the spoon-shaped basis of the quillworts, was first introduced by Georg leaves. Megasporangia occur on the outer Graf zu MÜNSTER (1842, p. 107-8, pl. IV, leaf bases and microspores on the interior fig. 4) in the description of the specimen part. Isoetites crociformis from the Lower Jurassic of Daiting near Monheim. He chose this Description name to express the resemblance of the Roots: Holotype KÜH 035, a mostly entire tuberous stem and elongated leaves to those plant (at all 15 cm high), shows in the low- of the extant Isoëtes. This concept to use er part the organisation of the 3.7 cm long the name Isoetites for quillwort-resembling roots. They do not have traditional roots, fossil forms has gradually become accepted; but some of its leaves are modified to act the same happened for the generic term like rhizomorphs. The roots are not creeping Equisetites for horsetails being similar to or protruding, but try to claw compactly in extant Equisetes. All in all, this herbaceous the soil substrate. lycophyte has evolved very little from Early Stems: Bulbous to reduced, or completely Triassic to the present. missing. An indeterminate number of fertile to vegetative leaves form on the lower part Isoetites brandneri WACHTLER et. al., a rounded short corm (KÜH 035). 2010 Leaves: Tufted, erect to somewhat lax. Leaves narrowly cylindrical at the tips and 2010 Isoetites brandneri; Wachtler et al. Pl 2 - 4 gradually widen towards the base (until 0.6 cm), where they abruptly expand and flat- Holotype ten into a spoon-like form. Usually a fold KÜH035 in the middle part of the leaves can be noted. They can reach a length of 15 cm, Paratypes leaving rhomboidal leaf scars on the stem when released. On KÜH 035, several leaves KÜH634 corm, KÜH011 microspores, arise spirally, up to 8 cm long and 0.5 cm KÜH751 megaspore wide, from the base of the 4 cm long corm. Leaves narrow constantly until they reach a Material width of 0.3 cm in the middle and end api- KÜH 030, KÜH 041, KÜH 062, KÜH 459, cally lanceolate. KÜH 1246 shows the or- KÜH 246 ganisation of Isoetites brandneri from the upper side. Two plants growing close to- Etymology gether display stems that measure 1.35 cm and 1.15 cm, the closely arranged rolling After Rainer Brandner, who studied the geol- and papery leaves varying from 0.3 to 0.35 ogy and stratigraphy of the Braies Dolomites. cm in width. Leaf cuticles are 2.5 to 3.0-µm

Dolomythos, 2011 178 1 1 cm 2 1 cm

3 1 cm 1) KÜH 035. Isoetites brandneri sp. nov. Holotype. Complete plant with roots. 2) KÜH 002. Isoetites brandneri sp. nov. Old stem fragment covered by leaf scars. 3) KÜH 011. Isoetites brandneri sp. nov. Paratype. The specimen yield immature microspores.

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 179 thick, both on the upper and lower side. sian layers in the European Alps and Isoet- Most of the leaves are fertile with sporan- ites sagittatus from the late early Triassic gia of two kinds embedded in the leaf bas- of North China (WANG, 1991), this plant es. varies only in minimum details. Epidermis is thick on the upper and lower Closely related species to Isoetites are prob- side. Epidermal cells are isodiametric in the ably Lepacyclotes bechstaedtii found in central part, uncommonly more elongated the same Anisian sediments and Lepacy- near the margins. Abaxial epidermal cells clotes (Annalepis) zeilleri recorded from the are protected by papillae. Stomata are in ir- younger Ladinian deposits in Europe. The regular rows, sunken and covered partially Pleuromeiaceae and Carboniferous Sigillaria by papillae of the surrounding 5 to 7 subsid- could eventually be added to this group of iary cells. Adaxial epidermis occur with few single-standing lycopods. They stand any- or no stomata and epidermal cells without way in discrepancy to the Lyopodiales bear- papillae. Variations of the “normal” epider- ing creeping and protruding roots, such as mis occur sometimes (KÜH 751, KÜH 004, Lycopiaceae, Selaginellaceae and Carbonif- KÜH 035). erous Lepidodrendales. Sporangia: Plants hold two kinds of sporophylls. Each sporangium is solitary and embedded in the hollowed leaf bases. Meg- SYSTEMATIC PALAEONTOLOGY asporangia occur in the outer leaf bases, Division Lycophyta while the following circle of leaves bear Order Isoeteales PRANTL, 1874 microsporangia and the innermost leaves Family Isoetaceae REICHENBACH, 1828 are sterile. Megasporangia are large. Some immature groups of spores have been ex- Genus Lepacyclotes EMMONS, 1856 tracted from KÜH 011 and mature megaspores from KÜH 751. In KÜH 011, the mi- Lepacyclotes has to be regarded as a world- crospores are reniform and up to 35 to 40 wide ranging low-growing bulbed lycophyte. µm in diameter. They could belong to the It has been recorded from the Early-Middle dispersed genus Aratrisporites. Several ov- Triassic (Anisian) beds in China as Lepa- al to circular megaspores, 270 to 300 µm cyclotes (Isoetes) ermayinensis (WANG, in diameter, have been found in the cuticle 1991), from the coeval sediments of Küh- slides (KÜH 751). Megaspores are convo- wiesenkopf in the Dolomites as Lepacyclotes lute to verrucate with a thick spore. The or- bechstaedtii, from the Middle to Later Tri- namentation reduces proximally in height. assic as Lepacyclotes (Tomiostrobus) con- The trilete aperture is delicate, only a few vexus (BRIK, 1952) in Kazakhstan, and as times indicated by plicae (WACHTLER ET Lepacyclotes zeilleri from the Middle Triassic AL. 2010). (Ladinian) sites in the Lettenkohle in France, Germany and the Wengen-Formation in the Remarks Dolomites. Although many specimens are difficult to interpret because their leaves could eas- Lepacyclotes bechstaedtii WACHTLER ily be confused with those of a lot of oth- ET AL., 2010 er plants, especially when lacking fertile 2010 Lepacyclotes bechstaedtii; Wachtler et al. Pl 5 - 6 parts, compressions of Triassic Isoetaceae are known around the world. It is there- Holotype fore possible to suggest that the origins of this characteristic heterosporous short- KÜH 1285 stemmed lycopod date back to the Carbon- iferous if not to the Devonian time. Begin- Paratypes ning from the Early Triassic (Olenekian) KÜH 007, KÜH 547 with Isoetes beestonii, which is known from Australia (Blackwater – Queensland, Material Coal Cliff Sandstone) (RETALLACK, 1997), as well as Isoetites brandneri from the Ani- KÜH 2142

Dolomythos, 2011 180 4 1 cm

5 1 cm 6 1 cm

4) KÜH 1246. Isoetites brandneri sp. nov. Two plants growing very closely together. 5) KÜH 030. Isoetites brandneri sp. nov. Detail of the enlarged leaf bases containing sporangia. 6) KÜH 025. Isoetites brandneri sp. nov. View from the upper side.

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 181 Etymology elongated guard cells (WACHTLER ET AL., 2010). After Thilo Bechstädt, who studied with Sporophylls: The plant holds sterile and Rainer Brandner the geology and stratigra- fertile leaves, which are difficult to separate phy of the Braies Dolomites. because they are always densely packed. KÜH 547 displays an isolated sporophyll Type localities 1.5 cm long and 0.8 cm wide, with a cen- Kühwiesenkopf tral elongated fertile part that is 0.9 cm long and 0.5 cm wide. It has a putative quadrilo- Type horizon and age bate corm. Lower to Middle Triassic, Anisian, Pelson-Il- Remarks lyrian In 1856, the American geologist Ebenezer Diagnosis EMMONS described leaf circlets from the Late Triassic (Carnian) Pekin-Formation Low-growing lycophyta with closely-spaced in North Carolina as Lepacyclotes circula- foliage forming a rosette. Leaves are ob- ris and several detached scales as Lepacy- long and thickened, ending acuminate to clotes ellipticus. For a long time, the speci- triangular with sterile leaves holding central men passed unobserved in the U.S. National costae. Sporophylls with elongated sporan- Museum. In 1910, the French palaeontolo- gium occur in the basal part. Corm is quad- gist Paul FLICHE named some isolated and rilobate. disarticulated sporophylls Annalepis zeilleri. These fertile parts have also been misun- Description derstood and regarded as araucarian seed Roots: KÜH 2142 shows part of the 3 cm scales (GRAUVOGEL-STAMM & LUGARDON, long rhyzomorphs. They resemble those 2001). The confusion increased when simi- of Isoetites and are quite different from lar rosette-like structures or elongated tri- the protruding and creeping roots of Ly- angular sporophylls were described from copia. other parts of the world as Tomiostrobus, Stem: Lepacyclotes bechstaedtii spread Skilliostrobus or Cylostrobus (RETALLACK, into large clusters by forming 4 to 10 cm 1997). Finally, Lea Grauvogel-Stamm and high and wide rosettes. Holotype KÜH 1285 Philippe Duringer in 1983 cognised that the shows a mainly complete preserved side- scales of Annalepis zeilleri were sporophylls faced plant. From a 2 cm high stem, with a belonging to the lycopsids with megaspores diameter of 1.1 to 1.5 cm, arise spirally the of Tennellisporites and microspores assign- closely-spaced fertile and sterile leaves. The able to Aratrisporites. Although the first de- whole plant was 4.5 cm. KÜH 007, as well scribed Lepacyclotes found in considerable as KÜH 547 evidence compressed rosettes numbers lack clearly identified spores, but from the upper side, both have a total diam- are abundantly associated with them, the eter of 7 cm. resemblance of the fossilised specimen and Leaves: The oblong leaves open flattish especially the sporophylls suggest that we and are 1.6 to 1.7-cm long and 0.7 to 1 have to do it with a conspecific group of her- cm wide. The adaxial surface is flat to con- baceous lycophyta, with Lepacyclotes (EM- cave, broader in the middle and ends in a MONS, 1856) having priority over the other mucronate three-cornered structure. The one. basal leaves from KÜH 007 are up to 2 cm long, 0.6 cm wide and with a central costa. Discussion Only small cuticle fragments (KÜH 2122 Several low growing Triassic lycophyta are and KÜH 007) are obtained. The normal known from a lot of locations worldwide. epidermal cells are elongated (40 to 50 x Difficulties occur in differentiating if we have 10 µm), with papillae on one side and with- to do it with detached fertile parts from out on the other. A few stomata are irregu- sub-arborescent club mosses or small-sized larly distributed on both epidermal sides. plants to be inserted into a wider group of They show simple stomata with large and Isoetaceae-like forms.

Dolomythos, 2011 182 2 1 cm

1 1 cm

3 1 cm 4 1 cm

1) KÜH 1285. Lepacyclotes bechstaedtii sp. nov. Holotype. Mainly complete plant seen from the lateral side. 2) KÜH 2142. Lepacyclotes bechstaedtii sp. nov. Plant with crowded foliage. 3) KÜH 547. Lepacyclotes bechstaedtii sp. nov. Paratype. Plant seen from the upper side. 4) KÜH 007. Lepacyclotes bechstaedtii sp. nov. Paratype. Specimen with an inner circle of elongated leaf fragments.

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 183 Different fructifications from the Lower reduced leaves but bear more resemblances Triassic of Australia, belonging probably with Isoetaceae. In all of them, the leaves to sub-arborescent lycophytes, have have a relatively short upper leaf zone and been described as Tomiostrobus, a sheathing base comprising two-thirds Cylostrobus, Austrostrobus, Lycostrobus or more of the total leaf length. In Isoëtes and Skilliostrobus. Most are regarded as novo-granadensis, the up to 5-cm long tri- a synonym of Tomiostrobus (RETALLACK angular foliage is arranged in a half-buried 1997, TAYLOR ET AL. 2009). This is valid for rosette. The stems are generally fleshly and Takhtajanodoxa mirabilis from the Siberian buried completely in marshes or swamps. deposits. Although all these sporophylls Lepacyclotes bechstaedtii and Isoetites have some affinities with Lepacyclotes brandneri have only been found in the Küh- – producing even the same microspores wiesenkopf “Dead Fish-Basin”, together with of dispersed genus Aratrisporites - they the other lycophytes Lycopia dezanchei and differ in having a distal elongated limb and Selaginellites leonardii. The latter two are bluntly pointed to mucronate ending tips common in other parts of the Piz da Peres (GRAUVOGEL-STAMM & LUGARDON, 2001). area. The basin harbours abundant skel- Furthermore, Tomiostrobus is characterised etons of fish like coelacanths, Gyrolepis, by a short stem (RETALLACK, 1997). Bobasatrania, Dipteronotus and Saurichtys, The genus Lepacyclotes was a worldwide- as well as being unusually rich in land ranging Mesozoic lycophyta, found from the plants. Besides the lycophytes and some Alps to China and Australia and appearing isolated Equisetites mougeotii, it is rich in in the Early-Middle Triassic until the Triassic, ferns like Neuropteridium or Gordonopt- when it began to disappear on the border eris, seed ferns such as Scytophyllum and with the Jurassic. The Isoetales, in contrast, Sagenopteris, conifers like Voltzia and Alpia, was just present in the earliest Triassic in as well as several cycadophyta species like Australia as Isoetes beestonii and slightly Bjuvia and Nilssonia and some enigmatic later as Isoetites brandneri, remaining only proto-angiospermales. Therefore, it is diffi- marginally modified until the present. cult to deduce globally active theories about While the Pleuromeiales with their un- extraordinary climatic changes from the branched erect stem have only vague affini- plant assemblage. More reasonable would ties, some fossilised remains especially from be the notion of a salt-saturated soil with Australia, India and Siberia could be more frequently ephemeral ponds as well as over- closely related to Lepacyclotes. However, flooded marshlands, where Lepacyclotes and this is only valid if all this morphogenus for Isoetites occupied the most exposed places. fructifications is not regarded as terminal This is the preferred living space for extant cones of other shrubby or bushy plants. Isoëtes species worldwide, which further Anisian Lepacyclotes bechstaedtii was much supports the idea that this plant never aban- smaller than Ladinian Lepacyclotes (An- doned its ecological niche and ancestral liv- nalepis) zeilleri, bearing sporophylls up to ing habitat in 300 million years. 5 cm long and 2 cm wide and reaching a plant size of 15 cm. Lepacyclotes circularis described by Emmons has nearly the same size as that of Lepacyclotes zeilleri. Some affinities persist with coeval Lepacyclotes er- mayensis holding elongated sporophylls with triangular mucronate tips. No extant plants have enough affinities with Lepacyclotes. Stylites andicola, found in extremely crowded colonies on a moist and limy substrate around the margins of glaciers at an elevation of 4,750 metres in Peru, also share resemblances with Lepa- cyclotes, but much more with Isoëtes. Oth- ers, like South American Isoëtes novo-granadensis or Tasmanian Isoëtes gunnii, have

Dolomythos, 2011 184 d

a a b

Isoetites brandneri: (a) Entire plant (KÜH 035), (b) Lepacyclotes bechstaedtii: (a) Entire plant (KÜH 1285), (b) megasporophyll and microsporophyll with leaves and (c) interior view of the plant (KÜH 1285), (c) upper view ((KÜH lateral side showing the arrangement of fertile parts. 007) and (d) single sporophyll.

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 185 SYSTEMATIC PALAEONTOLOGY Selaginellites leonardii sp. nov. WACHT- Division LYCOPHYTA Order SELAGINELLALES Prantl, 1874 LER ET AL. 2010 Family SELAGINELLACEAE Willkomm, 1854 Holotype Genus Selaginellites ZEILLER, 1906 KÜH 1140 Extant Selaginellaceae belong to a widely distributed family of herbaceous lycopods, well- Paratype adapted to various climate conditions and soil KÜH 956 types. Some species can resist extreme weath- er, such as those prevalent in alpine or artic cir- Material cles. They can also colonise barren and dry de- serts. One of the most well-known, Selaginella PIZ 622, PIZ 623, KÜH523, PIZ 563, PIZ lepidophylla, is called the resurrection plant be- 165 cause it can survive years without rainfall. They reach their highest diversity in tropi- Etymology cal areas under the forest canopy, protected Remembering Piero Leonardi, a pioneer in from direct sunshine or around riverbanks, the research on the geology of the Dolomites marshes or waterfalls. All extant Selaginel- laceae are included in one large genus of Type localities about 700 species. The presence or absence of leaves of two distinct sizes has been used Piz da Peres, Kühwiesenkopf as a criterion for subdividing extant Sell- aginellaceae (KORALL & KENRICK, 2002). Type horizon and age Anisophylly is characteristic of the subgen- Early-Middle Triassic, Anisian, Pelson era Heterostachys and Stachygynandrum, whereas Tetragonostachys, Selaginella and Diagnosis Ericetorum are isophyllous. Selaginellaceae are delimited from herbaceous Lycopodiace- Stems branch dichotomously and are reg- ae by their heterosporous fructifications in ularly or irregularly forked or branched. contrast to the isosporous Lycopods. Leaves tightly appressed to ascending on the The earliest fossil evidence of Selaginellaceae stems. Foliage structure linear-lanceolate to comes from the Carboniferous (Visean slightly falcate at the marginal ranks. Strobili 345.3 to 328.3 Ma) and from the late with clusters of overlapping sporophylls are Carboniferous (Selaginellites gutbieri), when arranged spirally to decussately. Sporophylls branching stems that bore minute leaves ovate, differentiated from vegetative sterile were widespread in coal measure floras. leaves. Heterosporous sporangia in the axils Surprisingly, in that time, isophyllous and of sporophylls are divided into compact tet- anisophyllous species coexisted (THOMAS rads of trilete microspores, while megaspores 1992, 1997). Although fossil Selaginellas are oval to circular with thick spore walls. are known from such a long time ago, their remains are very rare, often despite their Description inconspicuousness and fragility. Renè Zeiller Vegetative branches: Stems are densely instituted the genus Selaginellites in 1906 cespitose, ascending, slightly to greatly for fossil heterosporous lycopods resembling branched, up to 5 cm high and with recent Selaginellas. Even if Paleozoic-Mesozoic dichotomising axes without an appartenent Selaginellites are presumably congeneric with decrease in the thickness of lateral Selaginella, this classification concept has branches. PIZ 622 is a good example of become largely accepted. A delimitation to a mainly complete specimen. It shows other Lycopodaceae is that Selaginellites is an erect, many-branched stem holding restricted to heterosporous species, whereas juvenile and adult shoots. Some of the Lycopodites includes isosporous taxa and ultimate branchlets are merely short spurs, other species that are not known to be slightly enlarged at the tip and only a few heterosporous (ZEILLER, 1906). millimetres in length. The thickness of the

Dolomythos, 2011 186 3 1 cm

1 1 cm

4 5 mm

2 1 cm 5 1 cm 1) PIZ 622. Selaginellites leonardii. Mainly complete branch. 2) PIZ 623. Selaginellites leonardii. Branches show dichotomous forking. 3) KÜH 956. Selaginellites leonardii. Paratype. Axes evidence slightly enlarged short spurs. 4) KÜH 1140. Selaginellites leonardii. Holotype. Minute axes. 5) PIZ 165. Selaginellites leonardii. Sterile branch.

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 187 branches varies from 1.2 to 1.8 mm in PIZ an entire immature mass of microspores, 623. suggesting that the sporophylls were prob- Leaves: On juvenile shoots of PIZ 563, ably distributed in rows of microsporophylls leaves are 1 to 3 mm long and protrud- and megasporophylls (WACHTLER ET AL. ing, a little awl-shaped, displaying a subtle 2010). needle-like characteristic. Adult branchlets are covered with several rows of minute Selaginellites venieri sp. nov. WACHT- appressed, narrowly lanceolate and highly overlapping leaves. In that manner, they LER, 2011 are mostly invisible, suggesting naked shoots (PIZ 622, PIZ 623, KÜH 1140, KÜH Holotype 956 and PIZ 165). PIZ 548 Strobili: Holotype KÜH 1140 shows two compact heterosporous strobilus fragments, preserved on slightly different Material horizons in the block, but not in organic PIZ 548B counterplate connection with the vegetative parts. The complete larger strobilus (1.7 x 0.3 cm) Etymology bears helically to decussately arranged For Umberto Venier, which dedicate his life micro- and macrosporophylls, which are for searching fossil plants in the Eastern aligned in four irregular vertical files (1.5 Alps. to 2 x 1 to 1.2 mm). They are ovate with a long, acuminate apex (about 2 mm long) and an entire margin and are therefore Type localities completely different from the sterile leaves, Piz da Peres which are minute and acicular. PIZ 622 display in the lower part a semi-destroyed Type horizon and age strobilus attached to a branchlet. Strobili, in Early-Middle Triassic, Anisian, Pelson comparison to the shoots and the dimension of the whole plant, are extraordinarily long. Maceration of sporophyll fragments gave Diagnosis small cuticle pieces with isodiametric cells Cespitose low-growing lycopod with a and sporangia containing masses of imma- dichotomising axis. Leaves anisophyllous, ture, in situ microspores and megaspores. characterised by their opposite pairs of The microspores are mostly organised in im- dorsal and ventral leaves that differ in size mature, compact tetrads containing spores and form. The larger leaves are attached approximately 46 (42.5 to 50) µm in diame- laterally and the smaller ones medially on ter. Well-developed, separated trilete micro- the upper surface. Leaves mostly entire to spores are 56 (45 to 62.5) µm in diameter, minutely denticulate. proximally with a punctate (possibly granu- late) or smooth (psilate) spore wall and dis- Description tally rugulate. Several oval to circular mega- Vegetative branches: Erect plant, dichot- spores of 312 x 356 (270 to 340 x 300 to omously branched, with leaves crowded 410) µm in diameter have been found dis- throughout the stem. Selaginellites venieri persed in the sediment; clusters of mega- is characterised by their anisophyllous mor- spores have been extracted from the sporo- phology, with its leaves in two ranks and the phylls. Megaspores are psilate to punctate, ventral or lateral set smaller than the dorsal with a 10-µm thick spore wall. The trilete set. aperture is delicate or not well-developed, Vegetative leaves: Both median and lat- and only rarely indicated by plicae. Mega- eral leaves are 2-pairs ranked, closely imbri- spores, and to a lesser degree microspores, cate, narrowly rounded at the tip, strongly are found both in the lower and upper part oblique to the axis, and ascending at an ap- of cone fragments. During maceration, an proximately 45o angle. The lateral side is 1.8 entire sporophyll with adhering sediment to 2.2 mm long and 0.5 to 0.7 mm wide, yielded a small cluster of megaspores and

Dolomythos, 2011 188 6 3 mm 8 1 cm 9 1 cm

7 1 mm 10 5 mm 6) KÜH 1140. Selaginellites leonardii. Holotype. Heterosporous strobilus. 7) KÜH 1140. Selaginellites leonardii. Holotype. Heterosporous strobilus. Detailed. 8) PIZ 622. Selaginellites leonardii. Badly conserved, but attached strobilus. 9) PIZ 563. Selaginellites leonardii. Shrub with well-preserved or juvenile leaves. 10) PIZ 563. Selaginellites leonardii. Detail of subtle leaves. while the central part is 0.6 to 0.8 mm long ecosystem in the Dolomites. The isophyl- and 0.5 to 0.7 mm wide. The surface area lous Selaginellites leonardii, characterised of the external standing leaves is therefore by their same-sized leaves, and the aniso- about three times greater than that of the phyllous Selaginellites venieri, with its pos- smaller internal leaves (PIZ 548). Anisophyl- session of two sets of different foliage on ly is typical for many species of extant Se- the same rank. Unfortunately, only a few, laginella species. sometimes, poorly preserved Triassic heterosporous Selaginellites species are currently Discussion known, most of them found in Upper Trias- sic (Rhaetian) rocks in Arizona (Selaginella At least two quite different Selaginella spe- anasazia), Greenland (Selaginellites pola- cies formed part of the Early-Middle Triassic

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 189 a b

Selaginella venieri: a) Part of a branch, b) evidencing the arrangement of the anisophyllous leaves

a b c d

Selaginella leonardii: (a) Entire twig, (b) part of an ultimate branch, (c) sporophyll and (d) internal part of a sporophyll displaying macro- and microsporangia.

ris), Sweden (Selaginellites hallei and Lyco- lites venieri, but not enough to classify them podites scanicus) and China (Selaginellites as conspecific. The most closely resembling yunnanensis). They have some affinities species to Selaginellites leonardii is Rhae- with Selaginellites leonardii and Selaginel-

Dolomythos, 2011 190 1 1 cm 2 3 mm

1) PIZ 548. Selaginellites venieri. Holotype. Branch with anisophyllous leaves. 2) PIZ 548. Selaginellites venieri. Holotype. Detail of the anisophyllous leaves. tian Selaginellites hallei, but it bears much occur in almost the same manner amidst smaller strobili. a variegated flora consisting of conifers, It is not possible - with the dried charac- ferns, seed ferns, cycads and horsetails. teristic, its close-fitted needles and also, To understand the desiccated disposition of in parts, the naked stems of Selaginellites Selaginellites leonardii, it would be much leonardii - to trace parallels within all the more reasonable to search them in local extant Selaginella species. The well-known ecological phenomena and not to use them Selaginella lepidophylla has some similari- to diagnose global or even catastrophic ties, but exhibits a much more equipped events. This spike-moss found its preferred leaf arrangement and is also anisophylous habitat probably in some rocky, sun- in contrast to the isophyllous Selaginellites exposed, up-rushed coastal zone. leonardii. Other is the situation for Selaginellites It would therefore be conducive to compare venieri. A habitat in the shadow of the the living parts and the visual nature of other plants like today’s anisophyllous Early Triassic and extant Selaginellaceae. Selaginellaceae could be likely accepted. Selaginellites leonardii is found mainly The ecological advantages of double-sized at two points in Piz da Peres: in the leaves are still unknown. Nonetheless, lower strata of Kühwiesenkopf and in the heterophyllous foliage characterises most slightly younger layers of the Piz da Peres Early Triassic plants like conifers (Voltzia Wachtler gorge. The species at these heterophylla and Schizolepis ungeri), seed two sites are remarkably common, in ferns (Scytophyllum bergeri) and cycado- association with Selaginellites venieri, and phyta.

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 191 SYSTEMATIC PALAEONTOLOGY Type horizon and age Division LYCOPHYTA Subdivision LYCOPHYTINA Kenrick & Crane, Dont Formation, Richthofen Conglomerate, 1997 Morbiac Limestone. Order LYCOPIALES Wachtler, 2011 Lower to Middle Triassic, Anisian, Pelson-Il- Family LYCOPIACEAE Wachtler, 2011 lyrian Diagnosis Genus Lycopia WACHTLER ET AL., 2010 Sub-arborescent lycophyte with wide creeping rhizomes, rooting at irregular Diagnosis intervals and covered by subtle hairy Sub-arborescent to arborescent lycophyte, appendices. Main stems erect, isophyllous, stems are wide creeping and rooting sparsely leafy, branching dichotomously and at irregular intervals. Main stems are anisotomously, bearing spirally-arranged dichotomously or anisotomously branched. foliage. Leaves are lanceolate to elongated, Leaves herbaceous, acicular, entire and maintaining an uniform width. Cuticle spreading on the apical part. Strobili is thick and epidermal cells isodiametric terminal and solitary on a short stalk, to rectangular. Stomata are sunken and bearing spirally-arranged sporophylls. protected by the papillae. Cones are of Sporangia are adaxially near the sporophyll Lycopodostrobus gaiae-type, with long base. helically-arranged decreasing sporophylls on a central axis. Sporangia are borne adaxially Etymology at the sporophyll base. After their insertion as lycophyte and com- Description memorating Julius Pia, and Austrian geologist and director of the Natural History Mu- Roots: Rhizomes grow mainly prostrate seum of Vienna, who during the First World along the ground, from which branches the War conducted intensive research in the main and erect stems. Two root fragments Braies Dolomites. (KÜH 1162 is 48 cm long and 13.5 cm wide, while KÜH 2438 is 40 cm long and 14 cm wide) display alternate arising stems. The Lycopia dezanchei Wachtler et. al., 2010 rhizomes are densely covered with 2 to 5 2004 Lycophyta gen indet. sp. indet.; Kustatscher mm long hairy rootlets and subtle 1 mm- ET AL., p. 131, pl. 1, fig. 4. wide pit marks. On KÜH 1162, the main 2010 Lycopia dezanchei; Wachtler ET AL. Pl 7, Fig 1, pl. stems arising reach a diameter of 4 cm and 8, Fig. 1 - 4 arise in a dorsolateral position. Stems: Stems are of considerable size, the Holotype fragment of KÜH 1423 is up to 60 cm long KÜH 1425 and 7 cm wide. Sometimes on the lower part, they are covered with rhomboidal Material cleavage compressions, a result of abscised leaves (KÜH 1229). Branches are sparsely PIZ 97, KÜH 1423, KÜH 1424, KÜH 655, leafy, isophyllous and covered with spirally- KÜH192, KÜH529, KÜH0641, KÜH1115, arranged, long, simple leaves especially KÜH1322 in the upper part. A dichotomy resulting in daughter axes of approximately equal Etymology diameter (KÜH 1264) as well as anisotomous After Vittorio De Zanche, who made exten- systems with one structurally dominant sive and detailed studies about the stratig- branch have been noted (KÜH 1424). Leaves raphy in the Dolomites. display a tendency to be bushy on the apical part and are downwardly curved. Type localities Leaves: On the base, the leaves are densely appressed and sometimes also scale-like. Kühwiesenkopf, Piz da Peres, Val Duron On older or larger stems, rhomboidal leaf

Dolomythos, 2011 192 1 2 cm 2 2 cm

1) KÜH 1425 (PAL 1264). Lycopia dezanchei (Holotype) Isotomous forking apical stem fragment Kühwiesenkopf. 2) PIZ 97. Lycopia dezanchei. Stem fragment with long leaves. Piz da Peres.

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 193 scars from released foliage are typical. In Etymology the higher parts, the foliage is spreading and protruding. Leaves are herbaceous, After Gaia, the primordial Earth goddess in lanceolate to elongated, with an entire ancient Greek mythology margin showing one central unbranched vein or costa. It extends through the entire Type localities length of the leaf. They taper only slight- Piz da Peres, Kühwiesenkopf ly from the base to the apex. The length of the leaves varies from 15 to 30 cm on Type horizon and age apical mature tree crowns, and their width ranges from 0.4 to 0.7 cm (KÜH 1423). Dont Formation, Richthofen Conglomerate, Thick leaf cuticle is amphistomatic, while Morbiac Limestone. normal epidermal cells are isodiametric to Lower to Middle Triassic, Anisian, Pelson-Il- rectangular (25 to 30 x 40 to 50 µm). Sto- lyrian mata are sunken and protected by papillae. Diagnosis Strobili: Cones are terminal and solitary on Elongated strobili with long gradually taper- a short stalk, cylindrical, up to 10 cm long ing sporophylls. Spores are homosporous and 3.5 to 4 cm wide. Sporophylls are scale- and clustered on the upper side near the like and overlapping, broadly ovate to del- apex. Sporophylls encase the cone at a ju- toid, narrowing to an extralong pointed tip. venile stage to become increasingly protrud- Sporangia are borne adaxially near the spo- ing when mature. rophyll base and are homosporous. Description Genus Lycopodostrobus nov. gen. Several lycophyta cones at all growth stag- WACHTLER, 2011 es were recovered from Piz da Peres. They help elucidate the evolution of club moss Diagnosis cones from the Palaeozoic to the present. While the mostly mature cones from Küh- Strobili of Triassic Lycopodiales with helically wiesenkopf yielded no in situ spores, four arranged homosporous sporophylls. Sporo- juvenile to semi-fully grown strobili from phyll scales end in a long pointed append- Piz da Peres exhibited the position of the age. Sporangia occur adaxially on the lower spores inside the sporophylls. Holotype PIZ leaf axils. 98, with 7-cm long preserved parts, shows the organisation of these strobili well. The Etymology estimated length of the cone body of Ly- After their affinity to the strobili of the Lyco- copodostrobus gaiae probably reached 10 podiales. to 20 cm in length (KÜH 1276: 15.5 cm long) and 3 to 4 cm in width. The cones can continue for a further 5 cm, showing a Lycopodostrobus gaiae nov. sp. characteristic tapering appendix. PIZ 213 WACHTLER, 2011 represents a juvenile specimen with a cone length of 5 cm, held by a 1-cm long stalk. Holotype Sporophylls are attached at an angle of about 80 degrees to the cone apex. The PIZ 98 single sporophylls are complete, with a triangular to slightly lanceolate ending and Paratype 2 to 5-cm long pointed prolongations. In KÜH 1262 mature cones (KÜH 1262 and KÜH 1426), the characteristic appendices become in- Material creasingly protruding, whereas in juvenile forms, they tend to encase the cones. The PIZ 99, PIZ 213, KÜH 1426, PIZ 317, KÜH spore-bearing zone, probably reniform, 778, KÜH 1276 lies on the adaxial side of the sporophyll

Dolomythos, 2011 194 1 cm 4

3 4 cm 5 3 cm

3) KÜH 1423. Lycopia dezanchei. Extraordinarily long stem fragment. 4) KÜH 655. Lycopia dezanchei. Root segmented with subtle hairs. 5) KÜH 1424. Lycopia dezanchei. Bifurcating anisotomous apical stem fragment with bushy leaves.

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 195 near the axis and is 10 to 15 mm long and 5 characterised by their non-forking stems. mm wide. Differences in the root system (four-lobed tap root-like in Pleuromeia versus creeping Remarks and protruding) and different heterosporous sporangia from the homosporous structure Michael Wachtler first discovered Lycopia in of Lycopia dezanchei are other distinguishing 1999 on the slopes of Kühwiesenkopf in the features. Braies Dolomites. In the following years, he It seems that Lycopia stands were isolated collected other materials from Piz da Peres among the Mesozoic lycophyta, occupying and finally some badly preserved specimens mostly the ecological niche of Pleuromeia. from Val Duron belonging to the younger which was never recorded in the Early-Mid- Morbiac Limestone of the Illyrian age, which dle Triassic strata of Piz da Peres, but was enhanced knowledge on the appearance and very abundant in the Induan-Olenekian Wer- living habitat of this interesting lycophyte. fen-Formation of the neighbouring Carinthia Lycopia constitutes one of the most common and in the global distribution during the Ear- flora elements in the Anisian strata of the ly Triassic. Dolomites and was in sub-arborescent com- The generic name Lycopodites was intro- petition with conifers (Voltzia, Alpia and Al- duced to describe the isophyllous to aniso- bertia) and giant horsetails (Equisetites). phyllous leaves of the suggested Lycophyta, It should not be completely discarded that ranging from Devonian till Pleistocene (TAY- Triassic Lycopia has some affinity with ex- LOR ET AL. 2009). Especially for Palaeozoic tant Lycopodiales as well as Carboniferous fossils, it is nearly impossible to distinguish Lepidodendrales. If the descendants of Tri- minute branches from the distal twigs of assic Equisetites are observed in the Car- Lepidodendrales, particularly while the at- boniferous Calamites, this is not so mislead- tached reproduction organs are missing. ing. Most Lepidodendrales are thought to Lycopodites amazonica described from the have become extinct in Europe and North Middle Devonian of Brazil and Oxrodia gra- America at the end of the Carboniferous, cilis from the Mississippian of Scotland are whereas in China, they persisted into the thought to represent herbaceous lycopsids. Middle Permian. Others, like Synlycostrobus tyrmensis and A large number, probably more than 350 Onychiopsis psilotoides, belong to younger Lepidodendron species, has been described Cretaceous deposits and can therefore be since the genus was first recognized by regarded as the descendants of Lycopia. Kaspar Graf von Sternberg in 1820. Most Many more affinities have been discovered species are based either on fragments of with the extant group of Lycopodiales. Ex- large stems lacking leaves or on generally tant Lycopodiales are represented by four smaller branches with foliage still attached. genera (about 480 species in Lycopodium, Only rarely does good evidence of a 430 species in Huperzia, 41 species in Ly- connection between leafy shoots, main copodiella and ones species in Phylloglos- axis fragments and still fertile organs sum. All are homosporous, generally di- exist (CHALONER & MEYER-BERTHAUD, chotomously branched and represent small 1983). A feature of Lepidodendron is their herbaceous plants. Their stems arise ad- expanded pyramidal and also diamond- ventitiously from a horizontal rhizome. The shaped decurrent leaf base. This is most Lycopodiales have a global distribution, but evident when the leaf has been abscised and their favoured habitat spans mainly from a characteristic leaf cushion remains, called temperate to subarctic regions. ”Blattpolster“ by older authors in German. Both extant Lycopodiales and Lycopia show Comparing Lycopia to other Triassic a comparable root system with a reptant lycopsida, such as Isoetites or Lepacyclotes, stem and rootlets. Lycopia also bears either they have to be immediately discharged aboveground or underground running rhi- because of their herbaceous characteristic, zomes or lateral branches. The abscised leaf their non-protruding rhizome structure cushions are pyramidal in Lepidodendron and non-dichotomising branching system. and more rhomboidal in Lycopia, but simi- This is also the main difference between lar in every case. The robot-like structure of arborescent Pleuromeiales, which are Lepidodendron, commonly called Stigmaria,

Dolomythos, 2011 196 2 5 mm

1 1 cm

1 cm 3 1 cm 4 5 1 cm 1) PIZ 98. Lycopodostrobus gaiae (Holotype). Suggested strobilus of Lycopia dezanchei. 2) PIZ 98. Lycopodostrobus gaiae (Holotype). Detail of sporophylls . 3) PIZ 99. Lycopodostrobus gaiae. Strobilus with sporophylls. 4) PIZ 213. Lycopodostrobus gaiae. Strobilus with attached sterile leaves of Lycopia dezanchei on the lower part. 5) KÜH 1262. Lycopodostrobus gaiae. Mature strobilus bearing spatiolate sporophylls.

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 197 resembles the rhizomes of Lycopia in many heterosporous reproduction. The discov- parts, with its shallow propagation structure ery of abundant material of Isoetaceae and and helically-arranged lateral appendages. Selaginellaceae in the Anisian layers con- In Stigmaria and Lycopia, the appendages solidates the argument that other relatives are abscised during plant growth, leaving have to be searched for. small subtle circular scars in Lycopia and Altogether, Lycopia constitutes a highly in- bigger ones in Stigmaria. teresting plant and, if not a direct descend- Lepidodendron and Lycopia hold apically di- ant of giant Lepidodendron, it could at least chotomising axes and do not have ligulate be integrated into the same family. It seems leaves. The organography of branching was that the arborescent lycophyte crossed the fundamentally dichotomous in extinct Lepi- Permian by reducing their size enormously dodendron, varying from equal to unequal. and survived the Mesozoic by further de- When unequal, overtopping would have oc- creasing its size to occupy ecological niches curred, leading to a determinate branch. as dormant dwarfs, maybe increasing in size The same could be stated for Triassic Lyco- when circumstances changed. pia as well as the recent Lycopodiales. Both also bear homosporous strobili. Lycopia could be considered a “Giant Lycopod” in the General discussion same manner as the “Giant horsetails Equi- setites” from the same Anisian strata. Why I conducted this research Lepidostrobus is a loosely defined morpho- Since 1990, I spent most of my time in genus, named and briefly described by Alex- the mountains around Piz da Peres. First, andre BRONGNIART after a cone determined I had to learn how to walk and then hear for the first time by Parkinson (1804) and the sound of the past, at least to see inside currently stored in the London Natural His- the rocks. This took years and years of hard tory Museum. For the next 150 years, all work. Only after that, I felt mature enough cones that even vaguely fitted BRONGNI- to understand the rocks. At the end, the ART’s criteria were included there (BRACK- rocks began to tell me their living history. HANES, S.D. & THOMAS, B.A., 1983). Since Thus, I discovered a lot of new and, at that the holotype yielded only microsporangia, time, unknown fossils. I often read in books but bisporangiate cones were found in Lepi- the manner in which I can understand the dodendron, a distinction was made that the past, but the best way is by exploring and morphogenus Flemingites has to be used for searching until you have the missing links that kind of strobilus, whereas Lepidostro- in your hand. These are then the incontro- bus should only include microsporangiate vertible arguments for understanding how strobili (TAYLOR ET AL. 2009). life evolved or how they could have evolved. Although never found in complete organic Unfortunately, we now live in a time when connection, Lycopodostrobus gaiae many scientists perform their studies in the strobili constitute the fertile organ of privacy of their own home and not in nature. Lycopia dezanchei. The cone morphology Although cladistic analyses on a writing is comparable with that of Carboniferous desk can be useful, they are nothing against Lepidostrobus cones belonging to many years of studying in the wilderness. Lepidodendron. They are characterised by All that time, I lied in wet soil or scrambled their homosporous strobili borne on distal onto exposed rocks. Avalanches buried me, branches in the crown of the tree. Such I was struck by lightning, and I suffered strobili found in the Stephanian sediments from cold and heat. However, I felt like a on Stangnock in Carinthia also bear typical free man, something that I could not affirm protruding leafy appendices. Only the size of from the people in the valleys, who had to Triassic Lycopodostrobus is smaller, reaching carry out often nonsensical work. Some- half of the Carboniferous one in the best times, my absence from normal life was cases. noticed and the police put pressure on me Compared to the extant Lycopodiopsida, with their guns. I saw villages expanding the Isoetales and Selaginellas have to be and an increasing number of ski slopes that eliminated because of their totally differ- restricted the areas I was investigating. ent composition, root system and mainly

Dolomythos, 2011 198 g h i j

f e

d c

a b

Lycopia dezanchei: (a) Entire plant, (b) root horizon, (c) stem, (d) Lycopodostrobus gaiae: Cone with open sporophylls, (e) sporophyll displaying microsporangia, (f) closed sporophyll, (g) single leaf, (h) single shoot, (i) isotomous growth pattern and (j) anisotomous forking stem with one part more dominant than the other.

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 199 Shortly, the authorities imposed a research numerous spirally-arranged microphylls, and ban on me and punished me. However, I reproduction by spores produced in large never suffered from boredom or despera- sporangia borne adaxially to the leaves, re- tion. I said to myself that if I helped to de- main remarkably unchanged from the old- code only one drop of water in the big ocean est member of the group (HUEBER, 1983). of science, my life was just worth living. Selaginellaceae has also been recorded in The significance of lycopods in the past the Carboniferous, whereas the oldest mem- Although extant lycophytes constitute a bers of Isoetaceae have only been recorded modest group of herbaceous plants, they as far back as the Early Triassic in Australia have played an enormous role in the history (Isoetes beestonii; RETALLACK, 1975); the of land vegetation and have the longest fos- Alps (Isoetites brandneri; WACHTLER ET AL. sil record of any vascular plant, extending 2010). Certainly they have to date further over 400 million years from the Early De- back because just in that time of global dis- vonian, maybe from the late Silurian until tribution. the present (CLEAL & THOMAS, 2009). They dominated vegetation around the world and The modification of landscape from the were the largest known living organisms in Carboniferous to the Triassic the Carboniferous. The oldest representa- tives of the group are found in the Early An impoverished flora after the Carbon- Devonian with Baragwanathia and Drepano- iferous. In a geologically short time, the phycus (SCHWEITZER, 1990). The presence dominant giant lycophyta disappeared com- of lycopods - from herbaceous to arbores- pletely. In the lower Permian, only isolated cent - can be traced continuously from then remains were found (Rotliegend: Subsigil- until the present. All the characteristics that laria brardii and Lepidostrobus) (BARTHEL, typify Lycopodium today, such as creeping, 2009). This is also valid for the Alps, where dichotomous and anisotomous branching, only some doubtful Stigmaria ficoides roots

An entirely recovered Sigillaria stem from the Carinthian Stangnock area with a diameter of up to one metre (Collec- tion Georg Kandutsch).

Dolomythos, 2011 200 from the Early Permian Laas-Formation and global plant species in that period, rang- Wunderstätten have been published (FRITZ ing from the Carinthian mountains (Gail- ET AL. 1990; WANK & PRINZ, 1994). taler Alps, Karawanken), Germany (Pleu- Huge delta swamps in the Early Trias- romeia sternbergii), over the Caspian Sea to sic. In the Early Triassic, big parts of Europe Wladiwostok, China and Japan (Pleuromeia and other landscapes consisted of enormous rossica, P. obrutschewii, P. epicharis, P. ji- flat basins and delta swamps. The palaeoe- aochengensis and P. hattai), even occur- cological milieu was more humid with an in- ring in Australian locations from the former tertonguing of river channels and floodplains and lakes (MADER, 1990). This would have been consistent with the presence of Pleu- romeia and the abundant richness of humid- loving Equisetacea (Equisetites mougeotii), whereas other plants (the ferns Anomopteris mougeotii and Neuropteridium, the conifer Voltzia heterophylla, and the seed fern Scy- tophyllum bergeri) filled the niches of drier landscapes or higher-lying areas. Thus, the dominance of Pleuromeia and Equisetites is explainable without catastrophism theories. The short heyday of Pleuromeiales. The lycophyte Pleuromeia sternbergii is recorded extensively in the Induan-Olenekian and constitutes one of the most well-known

1 cm

Lepidostrobus sp. cone from Carboniferous (Upper 1 cm Stephanian) Carinthian Stangnock, belonging to Lepido- dendron (Collection Georg Kandutsch). Upper Carboniferous Selaginellites gutbieri with two at- Compared to the Carboniferous Lepidodendron cones, tached bisporangiate cones and well developed planar Lycopia strobili were nearly as large, but from half to branching ovate to lanceolote leaves with acute apices. one-third smaller in length. The composition of long Westfal D. Zwickau. Collection Jürgen Meyer, Zwickau and protruding leafy appendices on sporophylls is surprisingly similar.

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 201 Gondwana continent (P. longicaulis). As longing to the Marattiacea, Gleichenaceae, stated by other authors (RETALLACK, 1975; Osmundacea or Lindsaeaceae were highly LOOY ET AL. 1999; TAYLOR ET AL. 2009), diffused. This is also valid for the conifers the Pleuromeiales apparently came from dividing them in Voltziaceae and Alpiaceae, nothing, dominated the planet for a short which evolved into mainly all conifers pre- geological time and became completely ex- sent nowadays like Araucarians, Pinaceae tinct on the Olenekian-Anisian boundary. It and Cupressaceae. There was an enormous is also possible that they are the descend- diversification of the cycadophyta (Bjuvia ants of Carboniferous Sigillariaceae. Trac- and Nilssonia), with a splitting of mainly all ing an evolutionary line from Pleuromeiales living cycad genera and a rise in an unusual to any extant lycopods like Isoëtes is highly group of proto-angiospermales (Ladinia and improbable because Isoetales co-inhabited Pizperesia). Moreover, the lycophyta (Lyco- with Pleuromeiales in the Early Triassic. pia, Isoetites and Selaginellites) and sphe- The irresistible ascension of land plants nophyta (Equisetites) maintained their eco- in the Early-Middle Triassic. After nearly logical niche, even though very reduced in 50 million years of a difficult time for plants size and diffusion. Only the Peltaspermales beginning from the Early Permian to the (Scytophyllum, Sagenopteris), although Early Triassic, we relive the rise and spread highly present in the Early-Middle Triassic, of mainly all extant plants in the Anisian. A seemed to not have survived the Jurassic- lot of new-age ferns (Gordonopteris, Maran- Creataceous boundary. toidea, Ladinopteris, Sphenopteris, Anom- The importance of the new lycopod spe- opteris, Wachtleria and Neuropteridium) be- cies from Piz da Peres. The discovery of

1 2

The Induan-Olenekian (249.7 to 245 million years) Carinthian Pleuromeia area: (1) Pleuromeia sternbergii and (2) Equisetites mougeotii. In the background: The conifers Voltzia heterophylla, Alpia, the fern Neuropteridium, the seed fern Scytophyllum and Anomopteris.

Dolomythos, 2011 202 several new species and one new genus of assemblages and combinations of floras, like Lycophytes in the lower Middle Triassic of the club moss vegetation capable of surviv- the Southern Alps shed some interesting ing high-stress conditions of soil minerali- light on the evolution of this group. The Tri- sation and temperature. However, marked assic was a time of important radiation and differences in vegetation composition and change in several key morphological charac- physiognomy were not so distinctive to de- teristics of this group. After the predominant duce further speculations regarding global position of giant Lycopods in the Carbonifer- catastrophic events. ous, there was a decline in the Permian and The invariability of Isoetaceae. The quill- a much reduced level of radiation of dwarf worts maintained their aspect mainly un- lycopods in the Early Triassic, with nearly all varied at least from the Early Triassic until groups present then still extant now. today. Their evolutionary line traces more Possible geothermal events as a cause to the group of Carboniferous Sigillariaceae of this unusual lycophyta assemblage. than the Lepidodendrales, even though this The early Middle Triassic Kühwiesenkopf hypothesis is not completely acceptable. Dead-Fish basin is distinguished by their Moreover, this evolutionary line began in the variegated club moss association. Special Carboniferous or even the Devonian. It is, reasons for this local environment could lie however, suggested that there is a close re- in local geothermal features such as hot lationship between Triassic Isoetites (brand- springs, lakes, fumaroles, geysers, heated neri) and Lepacyclotes (bechstaedtii). soils or hot streams. These distinctive envi- The early polypetalous diversification ronmental conditions allowed unusual plant of Selaginellales. In the Early Triassic, we

1 2 3 4 5 6

The Anisian (245 to 237 million years) Kühwiesenkopf Dead-Fish-Rift with the Lycopod coast: (1) Lycopia dezanchei with (2) Lyopodostrobus gaiae, (3) Selaginella leonardii, (4) Selaginella venieri, (5) Lepacyclotes bechstaedtii, (6) Isoetites brandneri and (7) Equisetites mougeotii. In the fish basin: Saurichtys, Bobasatrania, Dipteronotus and coelacanths. In the background: Neuropteridium elegans, Neuropteridium voltzii, Voltzia unescoensis, Gordonop- teris lorigae, Alpia anisica, Bjuvia olangensis, Scytophyllum bergeri, Sagenopteris keilmannii and Aethophyllum stipulare.

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 203 Early-Middle Triassic lycophyta from the Alps

Pleuromeiales Isoetales

Isoetites brandneri Anisian - Early Middle Triassic

Adaxial view

cone

Megasporophyll Whole plant

Pleuromeia sternbergii Lepacyclotes bechstaedtii Olenekian - Early Triassic Anisian - Early Middle Triassic

Dolomythos, 2011 204 Early-Middle Triassic lycophyta from the Alps Selaginellales Lycopodiales

Selaginellites leonardii Anisian - Early Middle Triassic

Cone

Anisophyllous leave

Selaginellites venieri Lycopia dezanchei Anisian - Early Middle Triassic Anisian - Early Middle Triassic

Michael Wachtler: Lycophyta from the Early-Middle Triassic Dolomites 205 encounter Selaginella species with isoto- Equisetes as their direct descendants and phyllous (Selaginellites leonardii) and also probably originated in the colossal Lepido- anisophyllous leaves (Selaginellites venieri). dendron or Calamites trees of the Carbon- The advantage of one or the other could not iferous. In contrast to Isoetites or Selagin- be explained satisfactorily. Furthermore, ellites, which were always small-sized, they there is currently a nearly equal distribution became smaller over the million years until of anysophyllos genera like Heterostachys today as dwarf scrubbed plants in the copse and Stachygynandrum, or isophyllous spe- of other vegetation. Astonishingly, Lycopia cies such as Tetragonostachys, Selaginella resembling lycopods were, until now, never or Ericetorum. Their heterosporophylly is an recorded from other parts of the world or in old attribute, still remaining unvaried. The the Early Triassic and following periods. Oth- same is true for their often branching and er midget characteristics have been noted in low-growing characteristics. the Early Triassic, with the shrubby Voltzia The interesting evolution of Lyco- conifer Aethophyllum stipulare in contrast pia. One of the most surprising discover- to the arborescent Voltziales, as well as the ies involved the sub-arborescent Lycopia current enormous difference in the growing dezanchei. It seems to have the same re- habitus of giant Sequoia and shrubby dwarf duction history as the Equisetites. Both were pines. widespread in the Early-Middle Triassic floras The showdown of the dinosaurs. The of Tethydealan Dolomites and can be defined Anisian layers of Piz da Peres also hold a as “Giant horsetails” or “Giant Lycopods”. In rich ichnofauna dominated by Archosauri- addition, both have extant Lycopodium and formes (Chirotherium, Brachychirotherium

Lycopods are distributed all over the world. Club mosses like Lycopodium are homosporous, but spike mosses like Selaginellaceae and quillworts are heterosporous, with female spores larger than the male ones. Left: Huperzia den- tata, right: Lyopodium clavatum.

Dolomythos, 2011 206 and Isochirotherium) and some dinosauro- G., (1976). Bericht über erste Pflanzenfunde aus der morpha (Sphingopus ladinicus n. sp.; AVAN- Trias-Basis der westlichen Gailtaler Alpen bei Kötschach (Kärnten, Österreich), Carinthia II, S. 103-112 Klagen- ZINI & WACHTLER, 2011). The emergence furt. of the dinosaurs was therefore linked to the ANDREWS, H. N. (1961). Studies in Palaeobotany. Wi- transformation of vegetation. The increas- ley and Sons Ed., New York and London, 487 pp. ingly widespread bushlands benefitted tetra- ASH, S. R. (1972). Late Triassic plants from the Chinle pods, which were able to elevate themselves Formation in North-Eastern Arizona. Palaeontology, and walk two-legged. Sphingopus, with its 15(4), 598-618. reduced forelegs and three-clawed hind leg, ANDREWS, H. N. (1979). Skilliostrobus gen. nov., a new lycopsid cone from the early Triassic of Australia. achieved all the properties of true dinosaurs Alcheringa, 3, 73-89. that followed a little later. In this Early Trias- AVANZINI M., WACHTLER, M. (2011). Sphingopus la- sic, there was the dawn of a long-lasting al- dinicus isp. nov, from the Anisian of Pragser Dolomiten liance and partnership between animals and (Southern Alps, Italy), in press. plants. BALME, B. E. 1963. Plant microfossils from the Lower Triassic of Western Australia. Paleontology, 6 (1), 12- 40. References BANERJI, J. (1989). Some Mesozoic plant remains from AMERON, H. W. J., van BOERSMA M. (1974). Vorläufige Bhuj Formation with remarks on the depositional envi- Untersuchungsergebnisse an älteren und neu ronment of beds. The Palaeobotanist, 37, 159-168. aufgesammelten jungpaläozoischen Pflanzenfunden BARTHEL, M. (2009). Die Rotliegendflora des Thüringer der Ostalpen (Österreich). – Carinthia II, 84: 9–15, Waldes. – Veröffentlichungen des Naturhistorischen Mu- Klagenfurt. seums Schleusingen AMERON, H. W. J. van, BOERSMA, M. NlEDERMAYR, BECHSTÄDT, T. H. AND BRANDNER, R. (1970). Das Anis

Left: The recent club moss Lycopodiella cernua is found throughout the tropics and subtropics. It occurs along forest fringes, in young secondary forest, often in swamp margins, in grassland, on moist cliff-faces, hillsides and mountain slopes. Right: Anisophyllous Selaginella kraussiana.

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