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

Home , Cycad, Isoetales, Megachirella, Pleuromeia, Pleuromeiaceae, Sigillaria

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

Wachtler, M.: Lycophyta 1 Dolomythos Published online by the Dolomythos Museum, Innichen, South Tyrol, Italy.

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 Dolomy- thos Museum or on research using material in this Museum.

Editors: Edith Campei, Michael Wachtler

Dolomythos is published at frequent but irregular intervals. Manuscripts, orders for publications, and all correspondence concerning publications should be sent to:

Museum Dolomythos Rainerstraße 11 39038 Innichen Italy mail: [email protected]

Please cite this articles as: Wachtler, M., (12/2011): Lycophyta from the Early-Middle Triassic (Anisian) Piz da Peres (Dolo- mites - Northern Italy), Dolomythos, 3-49, Innichen.

1 Michael Wachtler, P. P. Rainerstrasse 11, 39038 Innichen, Italy, e-mail michael@wachtler. com.

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 Lycopodia- les. 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. Therefore, this work compared the living conditions in the Early-Middle Triassic age and developed information and solutions about the rapid extinction of the historic and often discussed Pleuromeiales.

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

Remarks for when my country learns to appreciate science. Michael Wachtler wanted to publish This study summarises fifteen years of my his discovery of a new Lycophyta species in lonely and dangerous work in the mountains Piz da Peres and Kühwiesenkopf. Although of the Eastern Alps in which I made a lot of agreeing with some parts of the content of interesting discoveries. In long-lasting stud- ”Lycophytes from the Middle Triassic (Ani- ies, cataloguing and descriptions, I tried to sian) locality Kühwiesenkopf (Monte Prà shed more light on a lost world. I thought della Vacca) in the Dolomites (northern It- that all this could be useful for humanity aly)“ (2010), this publication was, however, to understand more about our past. Unfor- never fully authorised by Michael Wachtler tunately, one day, the authorities confis- and edited under unacceptable pressure, cated and removed all my specimens from with several formerly-agreed conditions of my house, damaging a large part of my re- a reciprocal contract not being maintained. search. My computers, photos and notations Therefore, in my opinion, the right author- were also sequestered and taken away, the ship has to be read as MICHAEL WACHTLER, cataloguing labels changed to cover the EVELYN KUSTATSCHER, JOHANNA H. A. VAN traces of my findings and studies, and my KONIJNENBURG-VAN CITTERT or it has to be name cancelled from my collection and seen as an illegal publication or one with- anonymised. Many photos and annotations out agreement from all authors. He regrets were based on my older archives and a large the pronounced interdiction of research by part of these studies lacked further cuticle the authorities, as well as police force, at- analyses and pollen preparation. I apologise tempting to exercise his intellectual research for this and hope to make a revised version rights.

Wachtler, M.: Lycophyta 3 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.

Localities not yet, marine deposited cycles contributed to the Gröden-Formation during the Upper The fossil plant-rich Piz da Peres area covers Permian. During the following Early Triassic about 8 km of the Northern part of the Do- age, the now oceanic-influenced sediments lomite Mountains, starting from the western contributed to the Werfen-Formation. They side of Piz da Peres over the Ladinian village consist of a strongly varying sequence of of St. Vigil and reaching the Kühwiesenkopf mixed terrigenous siliciclastic and carbonatic over the hamlet of Prags. It traverses sever- lithofacies (BRANDNER ET AL., 2009). Since al mountains, which the native people refer the 19th century, the Werfen-Formation has to as a big mountain sundial. Starting from been divided into two sections: one begin- the Kühwiesenkopf, we have the ”Zehner” ning from the Permian-Triassic boundary (Tenner) “Elfer” (Elevener)and “Zwölfer” subdivided into Tesero-, Mazzin-, Andraz-, (Twelver), also called Hochalpenkopf, the Seis-Member and Gastropodenoolith, which “Einser” (Oner), also called Maurerkopf, and are characterised by an abundance of the the “Zweier”(Twoer). Other mountains with scallop-like bivalve mollusc Claraia especial- fossil plant lenses are the Flatschkofel and ly found in the Seis-Member, and the other the Drei-Fingerspitze. The main fossil plant composed of Campill-, Val Badia-, Cence- lenses lie in the sometimes-exposed open nighe- and San Lucano-Member, showing an rock slopes that are occasionally interrupt- increase in terrigenous input. Hemipelagic ed by shrubs and woods in which the lay- terrigenous and subordinate carbonate sedi- ers cannot be traced. Other places impor- ments were deposited continuously, while tant for the comprehension of Early Triassic storm events also sometimes delivered a lycophyte evolution lie in the neighbouring lot of plant debris. These basin sediments, Carinthian Mountains, where the Induan/ with a conspicuous clastic input, continue Olenekian Werfen layers hold a rich Pleu- until the Anisian Dont Formation (=Pragser romeia-dominated flora. Usually, the recov- Schichten sensu PIA, 1937). The fine silty ered specimens are well preserved and give marine Werfen layers reach 500 metres in a deep insight into the ecosystems after the thickness in the Northern Alps, and 200 me- Permian-Triassic boundary. tres in the Southern Alps (BRANDNER ET AL., 2009). The Campill-Val Badia Member in the Eastern Southern Alps could be corre- Paleobiology and paleoecology lated with the Middle Buntsandstein, a more During the Early Triassic age, the Eastern- detailed Volpriehausen-Hardegsen-Forma- Southern Alps were located near a beach tion of Germany (about 249.7 to 245.9 ± close to a shallow water basin. The high 0.7 Mya old) stretching from Induan to Ole- sedimentation rate suggests frequent storm nekian. events (hurricanes). After intense volcanic While the South Tyrolean Werfen-Formation activities in the Lower Permian, their first, has not revealed to date any determinable

Dolomythos, 2011 4 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. plant species, the nearby Gailtaler Alps and and Carnian, showing only minimal changes other mountain chains in Carinthia display in its aspect and reproduction cycles. well fossilised floras (Ulrichberg-Klagenfurt, Most Carinthian Triassic localities are situ- Wunderstätten-St. Paul, Nötsch-Villach, ated inside crowded vegetation below the Golsernock-Stockenboi and Laas-Kötschach- forest line (in the Dolomites, mainly all fos- Mauthen). They are characterised by a dom- sil plant-bearing horizons lie over the for- inance of the lycophyte Pleuromeia stern- est limit); therefore, only newly built forest bergii, accompanied by the sphenophyte trails enable the exploration of underlying Equisetites and some isolated Voltzia re- rocks. Sufficient Early Triassic plant layers mains (THIEDIG & KABON, 2011) and can have been opened over the years and the therefore be defined as a vegetation-rich, excavated fossil slabs give a good insight but species-poor flora. into ancient habitat systems. All the find- The locality Bleiriese-Golsernock, discov- ings mainly show predominance of the ly- ered in 2011 by Georg Kandutsch, Hel- cophyte Pleuromeia sternbergii, identical mut Prasnik and Michael Wachtler, exhibits to the Central German Buntsandstein one. abundant and well-preserved Pleuromeia It could be assumed that vast areas of the sternbergii material, accompanied by Eq- Northern hemisphere in the Early Triassic uisetites mougeotii. It is remarkable that were colonised by only one plant species. other plant groups like ferns, seed ferns, conifers or cycads are missing. The sub-arborescent lycophyte Pleuromeia, dominant worldwide during the Early Triassic period, seems to have been effectively extinct during the Olenekian-Anisian boundary (RE- TALLACK G., 1975). Ecological support is given by the slightly younger Anisian For- mations of the Dolomites where rich lycophyte assemblages (Lycopia dezanchei, Isoetites brandneri, Lepacyclotes bechstaedtii, Selaginellites leonardii and Se- laginellites venierii) are common. In spite of intensive research, a total absence of Pleuromeia has to be accepted. The giant horsetail Equisetites mougeotii has been recovered in the Werfen-Formation (Campill- Val Badia Member), as well as in the following successions from Anisian to Ladinian Exploring the newly discovered Olenekian fossilised- plant point Bleiriese-Golsernock in the Gailtaler Alps.

Wachtler, M.: Lycophyta 5 Open lying isolated tree-trunks, root-horizons and ripple marks from Kühwiesenkopf - main plant layer.

The Pleuromeia monocultures are effectively ity-loving horsetails (Equisetites), ferns and strange when compared to the almost ex- seed ferns (Callipteris), conifers (Ortiseia, ploding diversity of the Anisian landscapes Ullmannia), cycadophyta (Taeniopteris) and that followed a little later. The theory of the gingkophyta (Sphenobaiera). Therefore, commonly thought vegetation-poor and arid comparisons with today’s Okavango-delta Permian landscapes has to be revised. New with its seasonal floodings, salt islands and discoveries in the Dolomites in the Upper associated salt-resistant plants, and semi- Permian Grödner-Sandstein-Formation sug- arid zones facilitate the understanding of gest an extraordinary abundance of humid- this ancient landscape.

Dolomythos, 2011 6 Kühwiesenkopf - Main plant lenses

The Kühwiesenkopf lycophyte paradise containing equal proportion of Lycophy- ta, conifers, cycadophyta, ferns and seed The Anisian Dont Formation, consisting of a ferns. carbonate-terrigenous sequence more than After the overgrowing patch of forest, the 200 metres thick, reaches the most distin- layers crop out for 119.5 metres, interrupt- guished development in the Braies Dolo- ed once by a geological fault that elevates mites. The main plant-bearing horizon on the lenses 25 metres high. These beddings Kühwiesenkopf lays circa 75 metres above are characterised by a rich fish assemblage a massive carbonate platform attributed to belonging to five different families (Dipter- the Bythinian-Anisian Gracilis Formation (DE onotus, Saurichthys, Bobasatrania, Gyro- ZANCHE ET AL. 1992; BROGLIO LORIGA ET lepis and coelacanths), of which the first AL. 2002; VAN KONIJNENBURG–VAN CIT- three genera have never or only rarely been TERT ET AL. 2006). recorded in the Early Triassic (TINTORI A. ET One club moss-rich layer on Kühwiesen- AL. 2001). Most of the fish skeletons have kopf has enabled, like in a few other plac- been found in a narrow, only three-metre es worldwide, an inimitable insight into a long section in the middle. The preservation contiguous beach more than 240 million of invertebrates in this area can be relat- years old. This main lens can be prose- ed to the very rapid burial events made by cuted for a total of 214 metres, of which heavy storms and flood deposits originating the middle 52 metres are covered by a from terrestrial domains. dense forest where no rocks protrude. On All the fishes seem to be well adapted to the right side, there are plant layers 42.5 live near the coastline. Dipteronotus sp., metres long and one-metre thick. There known only from a dozen specimens in Eu- is also the presence of the only terrestrial rope, is one of the best indicators of strong reptile, Megachirella wachtleri (RENESTO, fresh water influx in the Middle Triassic. 2003). The lenses hold a variegated flora Saurichthys sp., widespread in the Thety-

Wachtler, M.: Lycophyta 7 dalean ocean in the Triassic period, was an lites leonardii is much more abundant in the able predator. It is generally encountered in Wachtler gorge in the west side of Piz da Pe- sediments of marine environments, as well res, with another species, Selaginellites ven- as sporadically in rocks of brackish or conti- ierii, holding anisophyllous foliage. nental origin. Bobasatrania sp., found nearly All in all, we have to do it with an ancient worldwide from Madagascar to Cina, Green- marine depression probably near a highly land, Svalbard and Canada in Early Trias- saline coastline or also influenced by geo- sic beds, was another predator, but in comparison to Saurichtys sp., it was a steady swimmer, scrunching with great accuracy ______its preys with a battery of minute teeth. Gy- rolepis sp. has been found in sediments of 12,5 m marine coastal environments, especially in ______the Ladinian Fossillagerstätte of Monte San Giorgio. The skeletons of the coelacanths 15 m recovered allow, with caution, the insertion Lycophyta point of the species Heptanema (known from the Western Alps) in the Middle Triassic. The Megachirella point ichthyofauna attests a marine depositional environment with a very strong influx from ______nearby land, testified by a huge amount of plant remains. In the whole section, apart from the ubiq- 15 m uitous ferns, seed ferns, conifers and abun- ______dant cycadophyta species, there is the most complete club moss association. The most abundant is the sub-arborescent Lycopia Wood dezanchei, followed by Isoetites brandneri, Lepacyclotes bechstaedtii and Selaginellites leonardii. Apart from Isoetites brandneri 52 m and Lepacyclotes bechstaedtii, which have Wood never been recorded in the other layers of Kühwiesenkopf, Hochalpenkopf or Piz da Pe- ______res, Lycopia dezanchei is found everywhere and in large quantities, whereas Selaginel- 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 8 thermal events such as hot springs or heat- Isoetites brandneri (about 50 pieces) ed soils offering the best requirements for Lepacyclotes bechstaedtii (about 10 pieces) lycopods to predominate over other plant Selaginellites leonardii (about 20 pieces) families. Whether several storm events Selaginellites venieri (2 pieces) burst the plants in this depression or over- flooding at the coastline will be determined by some fossilised root horizon. Repository Just below this main Kühwiesenkopf horizon, Most of the macrofossil plant collection is a bank of nodular and bioclastic limestone stored at the Natural History Museum Südti- contains a rich marine epifaunal assem- rol, Bozen. The remainder of the collection blage, dominated by brachiopods (Puncto- is in the Museum Dolomythos, Innichen, or spiriferella fragilis, Tetractinella trigonella, in the collection of Georg Kandutsch, Arriach Mentzelia mentzelii, Spiriferina_paläotypus (Carinthia). and Angustothyris angustaeformis) and bi- Specimens were photographed under natu- valves (Mysidioptera cainalloi, Neomorpho- ral light using Nikon D200, Lenses AF MICRO tis compta, Entolium discites, Newaagia NIKKOR 60 mm 1:2.8 D and AF-S NIKKOR noetlingi, Pleuromya cf. musculoides, P. 17-35 mm 1:2.8 D for larger pieces. The brevis and Neoschizodus? sp. Others, such digital images were processed using Adobe as Plagiostoma striatum, Pseudoplacunopsis Photoshop CS version 12.1. fissistriata and Prospondylus comptus have been found only in the plant bearing depos- it, or in fossiliferous horizons located in the upper part of the formation (Parallelodon esinensis, Pteria? sp., Entolium kellneri and Chlamys (Praechlamys) cf. schroeteri) (PO- SENATO, 2008). After this fish- and club moss-rich section, this lens becomes abruptly increasingly thinner to the end in well-preserved ripple- marks that wedge into the mainly sterile Explanation of the table on the next pages: and plant-free horizons. Although other lay- Like an expulsion from the Garden Eden, we have the ers in Kühwiesenkopf and Piz da Peres are biggest modifications in plant kingdom at the Carbonif- rich in club mosses, they never reach the erous-Permian boundary. Surprisingly, mainly all plant abundance of this lens. Furthermore, ex- families developed during this time. tant club moss communities are known from In the Permian, we encounter an impoverished and de- creasing number of plant families. stress-conditioned biocenosis, like the Tepu- From the Early Triassic onwards, we have a general re- is of Venezuela or some high-plateau in the covery and mutation of all plant associations. Andes. Such marine-near and also different From a cycadophyte-related group originates the proto- genera-rich lycophyte zones are unknown angiosperms as the true cycads. In Early Triassic, we nowadays. The richness of all plant fami- have the division of “feather-sporophyll”-bearing cycads lies in this Anisian stratum, from conifers to like extant Cycas and strobili-holding cycads like Bowe- ferns and cycadophyta, does not allow ex- nia, Ceratozamia, Dioon, Encephalartos, Stangeria and Zamia. traordinary conclusions to be made about The conifers split into the araucarian line (Voltziales) global cataclysms or totally different living and pinaceae-cupressaceae line (Alpiales). conditions during that period. Seed ferns were dominated for a long time by Peltas- permum-Scytophyllum-Thinnfeldia and a Sagenopteris line. Both seemed to disappear at the Jurassic-Creta- Materials and methods ceous boundary without leaving any extant descendants. To date (2011), the Lycophyta fossil record Equisetaceae diminished from the Triassic period to is characterised by: present. Pleuromeia sternbergii (about 200 species) Lycophyta remained all the time mainly unmodified in only known from the Olenekian strata in the Isoetaceae and Selaginellaceae, whereas Lycopia Carinthia met the same fate as Equisetites. It was reduced to surviving as Lycopodiales. The Pleuromeiales became Lycopia dezanchei (about 200 pieces) extinct in a very short time after the Early Triassic. Lycopodostrobus gaiae (about 10 pieces)

Wachtler, M.: Lycophyta 9 Dolomythos, 2011 10 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

Wachtler, M.: Lycophyta 11 SYSTEMATIC PALAEONTOLOGY adaxial surface. They contain either micro- Division LYCOPHYTA spores or megaspores. Order PLEUROMEIALES Family PLEUROMEIACEAE Genus PLEUROMEIA Corda in Germar, 1852 Discussion The lycophyte Pleuromeia sternbergii constitutes one of the best-described and most in- Pleuromeia sternbergii (Münster) terpreted plants in paleobotany. It was found Corda in Germar 1852 in the first decade of the nineteenth century 1839 Sigillaria sternbergii MÜNSTER, P. 47, PL 3, FIG. 10 in the German Buntsandstein and described 1852 Pleuromeia sternbergii (MÜNSTER) CORDA in by the Czech botanist Karl Joseph Corda in GERMAR, p. 184 1852. After that, a lot of similar species or 1904 Pleuromeia sternbergii (MÜNSTER) CORDA in PO- subspecies were recovered around the world TONIE, n. 38 from France, Spain and Austria to Russia, 1904 Pleuromeia sternbergii (MÜNSTER) CORDA in China, Japan, Argentina and Australia. MÄGDEFRAU, p. 119, pl. III-VII This description therefore tried to work out 2011 Pleuromeia sternbergii (MÜNSTER) CORDA in only new stratigraphical and paleoecologi- THIEDIG & KABON, p. 203 – 224 cal phenomena from the not so well-known 2011 Pleuromeia sternbergii (MÜNSTER) CORDA in KANDUTSCH & WACHTLER fossil deposits in the Eastern Alps and left behind detailed specifications indicating only some references (TAYLOR ET AL. 2009; Type localities GRAUVOGEL-STAMM, 1993; GRAUVOGEL- STAMM & LUGARDON, 2001; RETALLACK, Carinthia: Bleiriese-Golsernock, Laaser Wald 1975). – Kötschach-Mauthen, Ulrichberg - Kla- The consistent Pleuromeia findings from genfurt, St. Paul Mountains, Rottensteiner Carinthia comes from the Induan-Olenekian Kreuz, Wunderstätten, Dobratsch, Waidisch- Werfen-Formation (Campill-Val Badia-Mem- bach ber), corresponding to the German Mittlerer Buntsandstein (THIEDIG & KABON, 2011; Type horizon and age KANDUTSCH & WACHTLER, 2011). In contrast to the continental German deposits, Lower Triassic, Induan-Olenekian, Werfen they belong to a marine environment with Formation accompanying casts and impressions of sea- shells like Neoschizodus and Bakevellia sp. The horizon newly discovered by Georg Repository Kandutsch, Helmuth Prasnik and Michael Georg Kandutsch Collection, Arriach - Carin- Wachtler in 2011 on the forest trail of Bleir- thia, Museum Dolomythos - Innichen (South iese-Golsernock near the Upper Carinthian Tyrol) locality of Stockenboi revealed an especially rich and well-preserved Pleuromeia sternbergii vegetation, accompanied by the giant Diagnosis horsetail Equisetites mougeotii. It was in- Sub-arborescent lycophyte with unbranched, teresting that other plant groups like ferns, erect trunk. Rhyzomorphic roots are typi- seed ferns, cycads and conifers were absent. cally four-lobed and the stem in the lower About 200 specimens were recovered from part is covered with traces of scars from the locality Bleiriese-Golsernock and some fallen leaves. Foliage in the upper part is from the Laaser Wald near Kötschach-Mau- lanceolatic, displaying a prominent mid-vein then, including typical four-lobed rhizomes, and attached on their enlarged basis to the stems bearing leaf scars, leaves and cones stem. Cones are well defined, sitting termi- with mega- and microsporophylls. They con- nally and solitary on the axis. Sporophylls firmed that we have to do it with a mainly in have a circular to oval lamina and a large pure stands growing plant. globose sporangium covering nearly all the The location near the ancient Tethys Ocean confirmed once more that the Pleuromeiales

Dolomythos, 2011 12 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.

Wachtler, M.: Lycophyta 13 occupied a wide-ranging habitat, growing as Selaginella-like club mosses have been re- a coastal halophyte as well as an inland xe- corded from the Early Triassic, and the rophyte in the Early Triassic German Basin, creeping and dichotomising Lycopiales of perhaps also in a desert or stormy environ- Anisian strata have a completely different ment. Surprisingly, this lycophyte, which structural composition, we could not hypoth- dominated the floras of the lower Trias- esise an apparent successor of Pleuromeia. sic, did not survive the Olenekian-Anisian We can only vaguely argue that maybe Car- boundary because a club moss-rich vegeta- boniferous Sigillarias have some similarities. tion was encountered (Lycopia dezanchei, It seems that the Pleuromeiales came from Isoetites brandneri, Lepacyclotes bechstae- nothing and disappeared without leaving any dtii, Selaginellites leonardii and S. venierii) trace. This appears strange because in the in the slightly younger Anisian strata around Eastern-Southern Alps, we have a mainly the neighbouring Piz da Peres, but no Pleu- continuous fossil plant record on a small ar- romeia was found. ea from the Carboniferous over Permian and Of interest are therefore the reasons why Triassic periods. It is dissatisfying to estab- the outstanding giant club moss Pleuromeia lish Pleuromeia as a pioneer plant after the became extinct in a geologically short time, Permian-Triassic cataclysm. A lot of other whereas the giant horsetail Equisetites, plants like the Peltaspermales, Coniferales, which occupied the same ecological niches, Cycadophyta and Equisetales survived, also passed unchanged on to the following pe- mainly unmodified by these supposedly all- riods. Since quillworts like Isoetites and destroying catastrophes.

d b

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 14 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.

Wachtler, M.: Lycophyta 15 SYSTEMATIC PALAEONTOLOGY Description Division LYCOPHYTA Order ISOETALES Prantl, 1874 Roots: The holotype KÜH 035, mostly the Family ISOETACEAE Reichenbach 1828 entire plant (at all 15 cm high), shows in Genus ISOETITES Muenster 1842 the lower part the organisation of the 3.7- cm long roots. They do not have traditional roots, but some of its leaves are modified Isoetites brandneri WACHTLER et. to act like rhizomorphs. The roots are not al., 2010 creeping or protruding, but try to claw com- pactly in the soil substrate. 2010 Isoetites brandneri; Wachtler et al. Pl 2 - 4 Stems: Bulbous to reduced, or completely missing. Mostly an indeterminate number of Holotype fertile to vegetative leaves form on the lower part of a rounded short corm (KÜH 035). KÜH035 Leaves: Tufted, erect to somewhat lax. Leaves narrowly cylindrical at the tips and gradually widen towards the base (until 0.6 Paratypes cm), where they abruptly expand and flatten KÜH634 corm, KÜH011 microspores, into a spoon-like form. Usually a fold in the KÜH751 megaspore middle part of the leaves can be noted. They can reach a length of 15 cm, leaving rhomboidal leaf scars on the stem when released. Material On KÜH 035, several leaves arise spirally, KÜH 030, KÜH 041, KÜH 062, KÜH 459, up to 8 cm long and 0.5 cm wide, from the KÜH 246 base of the 4cm-long corm. Leaves narrow constantly until they reach a width of 0.3 cm in the middle and end apically lanceolate. Etymology KÜH 1246 shows the same organisation as that of Isoetites brandneri from the up- After Rainer Brandner, who studied the ge- per side. Two plants growing close together ology and stratigraphy of the Braies Dolo- that have been preserved in this slab dis- mites. play stems that measure 1.35 cm and 1.15 cm, the closely arranged rolling and papery Type localities leaves varying from 0.3 to 0.35 cm in width. Leaf cuticles are 2.5 to 3.0-µm thick, both Kühwiesenkopf on the upper and lower side. Most of the leaves are fertile with sporangia of two kinds Type horizon and age embedded in the leaf bases. Epidermis is thick on the upper and lower Lower to Middle Triassic, Anisian, Pelson-Il- side. Epidermal cells are isodiametric in the lyrian central part, uncommonly more elongated near the margins. Abaxial epidermal cells are protected by papillae. Stomata are in ir- Diagnosis regular rows, sunken and covered partially Low-growing herbaceous lycophytes, which by papillae of the surrounding 5 to 7 sub- resemble extant quillworts. Plant displays sidiary cells. Adaxial epidermis occur with a small corm-like stem, leaves and rhizo- few or no stomata and epidermal cells with- morphous roots attached close together. out papillae. Variations of the “normal” epi- Foliage is spreading and protruding, sin- dermis occur sometimes (KÜH751, KÜH004, gle leaves are narrowly erect and ending KÜH035). as lanceolate. Heterosporous sporangia Sporangia: Plants hold two kinds of sporo- are embedded on the spoon-shaped basis phylls. Each sporangium is solitary and em- of the leaves. Megasporangia occur on the bedded in the hollowed leaf bases. Megaspo- outer leaf bases and microspores on the in- rangia occur in the outer leaf bases, while terior part. the following circle of leaves bear microspo-

Dolomythos, 2011 16 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.

Wachtler, M.: Lycophyta 17 rangia and the innermost leaves are sterile. Triassic (Olenekian) with Isoetes beestonii, Megasporangia are large. Some immature which is known from Australia (Blackwater groups of spores have been extracted from – Queensland, Coal Cliff Sandstone) (RETAL- KÜH 011 and mature megaspores from KÜH LACK, 1997), as well as Isoetites brandneri 751. In KÜH 011, the microspores are re- from the Anisian layers in the European Alps niform and up to 35 to 40 µm in diameter. and Isoetites sagittatus from the late early They could belong to the dispersed genus Triassic of North China (WANG, 1991), this Aratrisporites. Several oval to circular meg- plant varies only in minimum details. aspores, 270 to 300 µm in diameter, have Closely related species to Isoetites are been found in the cuticle slides (KÜH 751). probably Lepacyclotes bechstaedtii found Megaspores are convolute to verrucate with in the same Anisian sediment and Lepacy- a thick spore. The ornamentation reduces clotes (Annalepis) zeilleri recorded from the proximally in height. The trilete aperture is younger Ladinian deposits in Europe. For delicate, only a few times indicated by plicae Gondwanian Tomiostrobus, Lycostrobus and (WACHTLER ET AL. 2010) Cylostrobus, it is important to determine if Lepacyclotes is conspecific with Isoetites. Pleuromeiaceae and Carboniferous Sigil- Remarks laria could probably be added to this group Isoetaceae is a group of widespread lyco- of single-standing lycopods. They stand any- phytes currently occurring in temperate re- way in discrepancy to the Lyopodiales bear- gions, as well as at high altitudes in tropical ing creeping and protruding roots, such as regions. Known as quillworts, these peren- Lycopiaceae, Selaginellaceae and Carbonif- nial plants, superficially resembling grasses, erous Lepidodrendales. grow mostly submerged in shallow water or in wet soils or flood plains. A habitat of per- manent or temporary lakes, riverbanks and SYSTEMATIC PALAEONTOLOGY saliniferous lagoons could also be true for Division LYCOPHYTA their ancestors. Order ISOETALES Prantl, 1874 The generic name Isoetites, for fossilised Family ISOETACEAE Reichenbach 1828 quillworts, was first introduced by Georg Genus LEPACYCLOTES Emmons 1856 Graf zu Münster (1842, p. 107-8, pl. IV, fig. 4) in the description of the specimen Isoet- ites crociformis from the Lower Jurassic of Lepacyclotes bechstaedtii WACHT- Daiting near Monheim. He chose this name LER et al., 2010 to express the resemblance of the tuber- 2010 Lepacyclotes bechstaedtii; Wachtler et al. Pl 5 - 6 ous stem and elongated leaves to those of the extant Isoëtes. This concept to use the name Isoetites for quillwort-resembling fos- Holotype sil forms has gradually become accepted; the same happened for the generic term KÜH 1285 Equisetites for horsetails being similar to extant Equisetes. All in all, this herbaceous Paratypes lycophyte has evolved very little from Early Triassic to the present. KÜH 007, KÜH 547 Although many specimens are difficult to in- terpret because their leaves could easily be Material confused with those of a lot of other plants, especially when lacking fertile parts, com- KÜH 2142 pressions of Triassic Isoetaceae are known around the world. It is therefore possible to suggest that the origins of this charac- Etymology teristic heterosporous short-stemmed lyco- After Thilo Bechstädt, who studied with pod date back to the Carboniferous if not to Rainer Brandner the geology and stratigra- the Devonian time. Beginning from the Early phy of the Braies Dolomites.

Dolomythos, 2011 18 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.

Wachtler, M.: Lycophyta 19 Type localities 1.5 cm long and 0.8 cm wide, with a central elongated fertile part that is 0.9 cm long Kühwiesenkopf and 0.5 cm wide. It has a putative quadrilobate corm. Type horizon and age Lower to Middle Triassic, Anisian, Pelson-Il- Remarks lyrian In 1856, the American geologist Eben- ezer Emmons described leaf circlets from the Late Triassic (Carnian) Pekin-Formation Diagnosis in North Carolina as Lepacyclotes circula- Low-growing lycophyta with closely-spaced ris and several detached scales as Lepacy- foliage forming a rosette. Leaves are oblong clotes ellipticus. For a long time, the speci- and thickened, ending acuminate to triangu- men passed unobserved in the U.S. National lar with sterile leaves holding central costae. Museum. In 1910, the French palaeontolo- Sporophylls with elongated sporangium oc- gist Paul Fliche named some isolated and cur in the basal part. Corm is quadrilobate. disarticulated sporophylls Annalepis zeilleri. These fertile parts have also been misun- derstood and regarded as araucarian seed Description scales (GRAUVOGEL-STAMM & LUGARDON, Roots: KÜH 2142 shows part of the 3-cm 2001). The confusion increased when simi- long rhyzomorphs. They resemble those of lar rosette-like structures or elongated tri- Isoetites and are quite different from the angular sporophylls were described from protruding and creeping roots of Lycopia. other parts of the world as Tomiostrobus, Stem: Lepacyclotes bechstaedtii spread into Skilliostrobus or Cylostrobus (RETALLACK, large clusters by forming 4 to 10-cm high and 1997). Finally, Lea Grauvogel-Stamm and wide rosettes. Holotype KÜH 1285 shows a Philippe Duringer in 1983 cognised that the mainly complete preserved side-faced plant. scales of Annalepis zeilleri were sporophylls From a 2-cm high stem and with a diameter belonging to the lycopsids with megaspores of 1.1 to 1.5 cm, arise spirally the closely- of Tennellisporites and microspores assign- spaced fertile and sterile leaves. In this case, able to Aratrisporites. Although the first de- we classified it with a 4.5-cm high plant. KÜH scribed Lepacyclotes found in considerable 007, as well as KÜH 547 showing compres- numbers lack clearly identified spores, but sion rosettes from the upper side, both have are abundantly associated with them, the a total diameter 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-cm have to do it with a conspecific group of her- wide. The adaxial surface is flat to concave, baceous lycophyta, with Lepacyclotes (EM- broader in the middle and ends in a mucro- MONS, 1856) having priority over the other nate three-cornered structure. The basal one. leaves from KÜH 007 are up to 2 cm long, Lepacyclotes has to be regarded as a world- 0.6 cm wide and with a central costa. On- wide ranging low-growing bulbed lycophyte. ly small cuticle fragments (KÜH2122 and It has been recorded from the Early-Middle KÜH007) are obtained. The normal epider- Triassic (Anisian) beds in China as Lepa- mal cells are elongated (40 to 50 x 10 µm), cyclotes (Isoetes) ermayinensis (WANG, with papillae on one side and without on the 1991), from the coeval sediments of Küh- other. A few stomata are irregularly distrib- wiesenkopf in the Dolomites as Lepacyclotes uted on both epidermal sides. They show bechstaedtii, from the Middle to Later Tri- simple stomata with large and elongated assic as Lepacyclotes (Tomiostrobus) con- guard cells (WACHTLER ET AL. 2010). vexus (BRIK, 1952) in Kazakhstan, and as Sporophylls: The plant holds sterile and Lepacyclotes zeilleri from the Middle Triassic fertile leaves, which are difficult to separate (Ladinian) sites in the Lettenkohle in France, because they are always densely packed. Germany and the Wengen-Formation in the KÜH 547 displays an isolated sporophyll Dolomites.

Dolomythos, 2011 20 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.

Wachtler, M.: Lycophyta 21 Discussion mayensis holding elongated sporophylls with triangular mucronate tips. Several low growing Triassic lycophyta are No extant plants have enough affinities known from a lot of locations worldwide. with Lepacyclotes. Stylites andicola, found Difficulties occur in differentiating if we have in extremely crowded colonies on a moist to do it with detached fertile parts from and limy substrate around the margins of sub-arborescent club mosses or small-sized glaciers at an elevation of 4,750 metres in plants to be inserted into a wider group of Peru, also share resemblances with Lepa- Isoetaceae-like forms. cyclotes, but much more with Isoëtes. Different fructifications from the Lower Tri- Others, like South American Isoëtes novo- assic of Australia, belonging probably to granadensis or Tasmanian Isoëtes gunnii, sub-arborescent lycophytes, have been de- have reduced leaves but bear more resem- scribed as Tomiostrobus, Cylostrobus, Aus- blances with Isoetaceae. In all of them, the trostrobus, Lycostrobus and Skilliostrobus. leaves have a relatively short upper leaf Most are regarded as a synonym of Tomi- zone and a sheathing base comprising two- ostrobus (RETALLACK 1997, TAYLOR ET AL. thirds or more of the total leaf length. In 2009). This is valid for Takhtajanodoxa mi- Isoëtes novo-granadensis, the up to 5-cm rabilis from the Siberian deposits. Although long triangular foliage is arranged in a half- all these sporophylls have some affinities buried rosette. The stems are generally with Lepacyclotes – producing even the fleshly and buried completely in marshes same microspores of dispersed genus Ara- or swamps. trisporites - they differ in having a distal Lepacyclotes bechstaedtii and Isoetites elongated limb and bluntly pointed to mu- brandneri have only been found in the Küh- cronate ending tips (GRAUVOGEL-STAMM & wiesenkopf “Dead Fish-Basin”, together with LUGARDON, 2001). Furthermore, Tomios- the other lycophytes Lycopia dezanchei and trobus is characterised by a short stem (RE- Selaginellites leonardii. The latter two are TALLACK, 1997). common in other parts of the Piz da Peres The genus Lepacyclotes was a worldwide- area. The basin harbours abundant skel- ranging Mesozoic lycophyta, found from the etons of fish like coelacanths, Gyrolepis, Alps to China and Australia and appearing Bobasatrania, Dipteronotus and Saurichtys, in the Early-Middle Triassic until the Triassic, as well as being unusually richn in land when it began to disappear on the border plants. Besides the lycophytes and some with the Jurassic. The Isoetales, in contrast, isolated Equisetites mougeotii, it is rich in was just present in the earliest Triassic in ferns like Neuropteridium or Gordonopteris, Australia as Isoetes beestonii and slightly (as well as the now missing Anomopteris, a later as Isoetites brandneri, remaining only fern characteristic of the Early-Middle Trias- marginally modified until the present. sic), seed ferns such as Scytophyllum and While the Pleuromeiales with their un- Sagenopteris, conifers like Voltzia and Alpia, branched erect stem have only vague affini- as well as several cycadophyta species like ties, some fossilised remains especially from Bjuvia and Nilssonia and some enigmatic Australia, India and Siberia could be more proto-angiospermales. Therefore, it is diffi- closely related to Lepacyclotes. However, cult to deduce globally active theories about this is only valid if all this morphogenus for extraordinary climatic changes from the fructifications is not regarded as terminal plant assemblage. More reasonable would cones of other shrubby or bushy plants. be the notion of a salt-saturated soil with Anisian Lepacyclotes bechstaedtii was much frequently ephemeral ponds as well as over- smaller than Ladinian Lepacyclotes (An- flooded marshlands, where Lepacyclotes and nalepis) zeilleri, bearing sporophylls up to Isoetites occupied the most exposed places. 5 cm long and 2 cm wide and reaching a This is the preferred living space for extant plant size of 15 cm. Lepacyclotes circularis Isoëtes species worldwide, which further described by Emmons has nearly the same supports the idea that this plant never aban- size as that of Lepacyclotes zeilleri. Some doned its ecological niche and ancestral liv- affinities persist with coeval Lepacyclotes er- ing habitat in 300 million years.

Dolomythos, 2011 22 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.

Wachtler, M.: Lycophyta 23 SYSTEMATIC PALAEONTOLOGY Type horizon and age Division LYCOPHYTA Order SELAGINELLALES Prantl, 1874 Early-Middle Triassic, Anisian, Pelson Family SELAGINELLACEAE Willkomm, 1854 Genus SELAGINELLITES Zeiller, 1906 Diagnosis Selaginellites leonardii sp. nov. Stems branch dichotomously and are reg- WACHTLER et. al 2010 ularly or irregularly forked or branched. Leaves are tightly appressed to ascending on the stems. Foliage structure is linear- Holotype lanceolate to slightly falcate at the marginal ranks. Strobili with clusters of overlapping KÜH 1140 sporophylls are arranged spirally to decussately. Sporophylls are ovate, differentiated from vegetative sterile leaves. Heterospor- Paratype ous sporangia in the axils of sporophylls are KÜH 956 divided into compact tetrads of trilete microspores, while megaspores are oval to circular with thick spore walls. Material PIZ 622, PIZ 623, KÜH523, PIZ 563, PIZ Description 165 Vegetative branches: Stems are densely cespitose, ascending, slightly to greatly Etymology branched, up to 5 cm high and with dichot- Remembering Piero Leonardi, a pioneer in omising axes without an appartenent de- the research on the geology of the Dolomites crease in the thickness of lateral branches. PIZ 622 is a good example of a mainly complete specimen. It shows an erect, many- Type localities branched stem holding juvenile and adult shoots. Some of the ultimate branchlets are Piz da Peres, Kühwiesenkopf

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.

Dolomythos, 2011 24 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 show slightly enlarged short spurs. 4) KÜH 1140. Selaginellites leonardii. Holotype. Minute axes. 5) PIZ 165. Selaginellites leonardii. Sterile branch.

Wachtler, M.: Lycophyta 25 merely short spurs, slightly enlarged at the yielded a small cluster of megaspores and tip and only a few millimetres in length. The an entire immature mass of microspores, thickness of the branches varies from 1.2 to suggesting that the sporophylls were prob- 1.8 mm in PIZ 623. ably distributed in rows of microsporophylls Leaves: On juvenile shoots of PIZ 563, and megasporophylls (WACHTLER ET AL. leaves are 1 to 3 mm long and protruding, a 2010). little awl-shaped, displaying a subtle needle- like characteristic. Adult branchlets are cov- Selaginellites venieri sp. nov. ered with several rows of minute appressed, WACHTLER, 2011 narrowly lanceolate and highly overlapping leaves. In that manner, they are mostly in- visible, suggesting naked shoots (PIZ 622, Holotype PIZ 623, KÜH 1140, KÜH 956 and PIZ 165). Strobili: The holotype KÜH 1140 shows two PIZ 548 compact heterosporous strobilus fragments, preserved on slightly different horizons in the block, but not in organic connection with Material the vegetative parts. The complete larger PIZ 548B counterplate strobilus (1.7 x 0.3 cm) bears helically to decussately arranged micro- and macrospor- ophylls, which are aligned in four irregu- Etymology lar vertical files (1.5 to 2 x 1 to 1.2 mm). For Umberto Venier, which dedicate his life They are ovate with a long, acuminate apex for searching fossil plants in the Eastern (about 2 mm long) and an entire margin. Alps. They are therefore completely different from sterile leaves, which are minute and acicular. PIZ 622 display in the lower part a semi- Type localities destroyed strobilus attached to a branchlet. Piz da Peres Strobili, in comparison to the shoots and the dimension of the whole plant, are extraordinarily long. Type horizon and age Maceration of sporophyll fragments gave small cuticle pieces with isodiametric cells Early-Middle Triassic, Anisian, Pelson and sporangia containing masses of immature, in situ microspores and megaspores. Diagnosis The microspores are mostly organised in immature, compact tetrads containing spores Cespitose low-growing lycopod with a di- approximately 46 (42.5 to 50) µm in diame- chotomising axis. Leaves are anisophyllous, ter. Well-developed, separated trilete micro- characterised by their opposite pairs of dor- spores are 56 (45 to 62.5) µm in diameter, sal and ventral leaves that differ in size and proximally with a punctate (possibly granu- form. The larger leaves are attached later- late) or smooth (psilate) spore wall and dis- ally and the smaller ones medially on the tally rugulate. Several oval to circular mega- upper surface. Leaves are mostly entire to spores of 312 x 356 (270 to 340 x 300 to minutely denticulate. 410) µm in diameter have been found dispersed in the sediment; clusters of megaspores have been extracted from the sporo- Description phylls. Megaspores are psilate to punctate, Vegetative branches: Erect plant, di- with a 10-µm thick spore wall. The trilete chotomously branched, with leaves crowded aperture is delicate or not well-developed, throughout the stem. Selaginellites venieri is and only rarely indicated by plicae. Mega- characterised by their anisophyllous morphol- spores, and to a lesser degree microspores, ogy, with its leaves in two ranks and the ven- are found both in the lower and upper part tral or lateral set smaller than the dorsal set. of cone fragments. During maceration, an Vegetative leaves: Both median and lat- entire sporophyll with adhering sediment eral leaves are 2-pairs ranked, closely

Dolomythos, 2011 26 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. imbricate, narrowly rounded at the tip, Discussion strongly oblique to the axis, and ascending at an approximately 45o angle. The later- Extant Selaginellaceae belong to a widely al side is 1.8 to 2.2-mm long and 0.5 to distributed family of herbaceous lycopods, 0.7-mm wide, while the central part is 0.6 well-adapted to various climate conditions to 0.8 mm long and 0.5 to 0.7 mm wide. and soil types. Some species can resist ex- The surface area of the external stand- treme weather, such as those prevalent in ing leaves is therefore about three times alpine or artic circles. They can also colonise greater than that of the smaller internal barren and dry deserts. One of the most leaves (PIZ 548). Anisophylly is typical for well-known, Selaginellites lepidophylla, is many species of extant Selaginella spe- called the resurrection plant because it can cies. survive years without rainfall.

Wachtler, M.: Lycophyta 27 a b

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

They reach their highest diversity in tropi- remains are very rare, often despite their in- cal areas under the forest canopy, protected conspicuousness and fragility. Renè Zeiller from direct sunshine or around riverbanks, instituted the genus Selaginellites in 1906 marshes or waterfalls. All extant Selaginel- for fossil heterosporous lycopods resembling laceae are included in one large genus of recent Selaginellas. Even if Paleozoic-Meso- about 700 species. The presence or absence zoic Selaginellites are presumably congener- of leaves of two distinct sizes has been used ic with Selaginella, this classification concept as a criterion for subdividing extant Sell- has become largely accepted. A delimitation aginellaceae (KORALL & KENRICK, 2002). to other Lycopodaceae is that Selaginel- Anisophylly is characteristic of the subgen- lites is restricted to heterosporous species, era Heterostachys and Stachygynandrum, whereas Lycopodites includes isosporous whereas Tetragonostachys, Selaginella and taxa and other species that are not known Ericetorum are isophyllous. Selaginellaceae to be heterosporous (ZEILLER, 1906). are delimited from herbaceous Lycopodiace- At least two quite different Selaginella spe- ae by their heterosporous fructifications in cies formed part of the Early-Middle Triassic contrast to the isosporous Lycopods. ecosystem in the Dolomites. The isophyl- The earliest fossil evidence of Selaginel- lous Selaginellites leonardii, characterised laceae comes from the Carboniferous (Vi- by their same-sized leaves, and the aniso- sean 345.3 to 328.3 Ma) and from the late phyllous Selaginellites venieri, with its pos- Carboniferous (Selaginellites gutbieri), when session of two sets of different foliage on branching stems that bore minute leaves the same rank. Unfortunately, only a few, were widespread in coal measure floras. sometimes, poorly preserved Triassic heter- Surprisingly, in that time, isophyllous and osporous Selaginellites species are currently anisophyllous species coexisted (THOMAS known, most of them found in Upper Trias- 1992, 1997). Although fossil Selaginellas sic (Rhaetian) rocks in Arizona (Selaginella are known from such a long time ago, their anasazia), Greenland (Selaginellites pola-

Dolomythos, 2011 28 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. ris), Sweden (Selaginellites hallei and Lyco- younger layers of the Piz da Peres Wachtler podites scanicus) and China (Selaginellites gorge. The species at these two sites are yunnanensis). They have some affinities remarkably common, in association with with Selaginellites leonardii and Selaginel- Selaginellites venieri, and occur in almost lites venieri, but not enough to classify them the same manner amidst a variegated flora as conspecific. The most closely resembling consisting of conifers, ferns, seed ferns, cy- species to Selaginellites leonardii is Rhae- cads and horsetails. To understand the des- tian Selaginellites hallei, but it bears much iccated disposition of Selaginellites leonar- smaller strobili. dii, it would be much more reasonable to It is not possible - with the dried charac- search them in local ecological phenomena teristic, its close-fitted needles and also, and not to use them to diagnose global or in parts, the naked stems of Selaginellites even catastrophic events. This spike-moss leonardii - to trace parallels within all the ex- found its preferred habitat probably in tant Selaginella species. The well-known ex- some rocky, sun-exposed, up-rushed coast- tant Selaginella lepidophylla has some simi- al zone. larities, but exhibits a much more equipped Other is the situation for Selaginellites ven- leaf arrangement and is also anisophylous ieri. A habitat in the shadow of the other in contrast to the isophyllous Selaginellites plants like today’s anisophyllous Selaginel- leonardii. laceae could be likely accepted. The eco- It would therefore be conducive to com- logical advantages of double-sized leaves pare the living parts and the visual nature are still unknown. Nonetheless, heterophyl- of Early Triassic and extant Selaginellace- lous foliage characterises most Early Triassic ae. Selaginellites leonardii is found mainly plants like conifers (Voltzia heterophylla and at two points in Piz da Peres: in the lower Schizolepis ungeri), seed ferns (Scytophyl- strata of Kühwiesenkopf and in the slightly lum bergeri) and cycadophyta.

Wachtler, M.: Lycophyta 29 SYSTEMATIC PALAEONTOLOGY Type localities Division LYCOPHYTA Subdivision LYCOPHYTINA Kenrick & Crane, Kühwiesenkopf, Piz da Peres, Val Duron 1997 Order LYCOPIALES Wachtler, 2011 Family LYCOPIACEAE Wachtler, 2011 Type horizon and age Dont Formation, Richthofen Conglomerate, Genus Lycopia Wachtler et al., Morbiac Limestone. Lower to Middle Triassic, Anisian, Pelson-Il- 2010 lyrian

Diagnosis Diagnosis Sub-arborescent to arborescent lycophyte, Sub-arborescent lycophyte with wide creep- stems are wide creeping and rooting at ir- ing rhizomes, rooting at irregular intervals regular intervals. Main stems are dichoto- and covered by subtle hairy appendices. mously or anisotomously branched. Leaves Main stems are erect, isophyllous, sparsely are herbaceous, acicular, entire and spread- leafy, branching dichotomously and anisot- ing on the apical part. Strobili are terminal omously bearing spirally-arranged foliage. and solitary on a short stalk, bearing spirally- Leaves are lanceolate to elongated, main- arranged sporophylls. Sporangia are adaxi- taining a uniform width. Cuticle is thick and ally near the sporophyll base. epidermal cells isodiametric to rectangular. Stomata are sunken and protected by the papillae. Cones are of Lycopodostrobus Etymology gaiae-type, with long helically-arranged de- After their insertion as lycophyte and com- creasing sporophylls on a central axis. Spo- memorating Julius Pia, and Austrian geolo- rangia are borne adaxially at the sporophyll gist and director of the Natural History Mu- base. seum of Vienna, who during the First World War conducted intensive research in the Braies Dolomites. Description Roots: Rhizomes grow mainly prostrate Lycopia dezanchei WACHTLER et. along the ground, from which branches the al., 2010 main and erect stems. Two root fragments (KÜH 1162 is 48 cm long and 13.5 cm wide, 2004 Lycophyta gen indet. sp. indet.; Kustatscher ET AL., p. 131, pl. 1, fig. 4. while KÜH 2438 is 40 cm long and 14 cm 2010 Lycopia dezanchei; Wachtler ET AL. Pl 7, Fig 1, pl. wide) display alternate arising stems. The 8, Fig. 1 - 4 rhizomes are densely covered with 2 to 5-mm long hairy rootlets and subtle 1-mm- wide pit marks. On KÜH 1162, the main Holotype stems arising reach a diameter of 4 cm and arise in a dorsolateral position. KÜH 1425 Stems: Stems are of considerable size, the fragment of KÜH 1423 is up to 60 cm long Material and 7 cm wide. Sometimes on the lower part, they are covered with rhomboidal PIZ 97, KÜH 1423, KÜH 1424, KÜH 655, cleavage compressions, a result of abscised KÜH192, KÜH529, KÜH0641, KÜH1115, leaves (KÜH 1229). Branches are sparsely KÜH1322 leafy, isophyllous and covered with spirally-arranged, long, simple leaves especially Etymology in the upper part. A dichotomy resulting in daughter axes of approximately equal di- After Vittorio De Zanche, who made exten- ameter (KÜH 1264) as well as anisotomous sive and detailed studies about the stratig- systems with one structurally dominant raphy in the Dolomites.

Dolomythos, 2011 30 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.

Wachtler, M.: Lycophyta 31 branch have been noted (KÜH 1424). Leaves Paratype display a tendency to be bushy on the apical part and are downwardly curved. KÜH 1262 Leaves: On the base, the leaves are densely appressed and sometimes also scale-like. On older or larger stems, rhom- Material boidal leaf scars from released foliage PIZ 99, PIZ 213, KÜH 1426, PIZ 317, KÜH are typical. In the higher parts, the foli- 778, KÜH 1276 age is spreading and protruding. Leaves are herbaceous, lanceolate to elongated, with an entire margin showing one cen- Etymology tral unbranched vein or costa. It extends After Gaia, the primordial Earth goddess in through the entire length of the leaf. They ancient Greek mythology taper only slightly from the base to the apex. The length of the leaves varies from 15 to 30 cm on apical mature tree crowns, Type localities and their width ranges from 0.4 to 0.7 cm Piz da Peres, Kühwiesenkopf (KÜH 1423). Thick leaf cuticle is amphisto- matic, while normal epidermal cells are isodiametric to rectangular (25 to 30 x 40 Type horizon and age to 50 µm). Stomata are sunken and pro- Dont Formation, Richthofen Conglomerate, tected by papillae. Morbiac Limestone. Strobili: Cones are terminal and solitary on Lower to Middle Triassic, Anisian, Pelson-Il- a short stalk, cylindrical, up to 10 cm long lyrian and 3.5 to 4 cm wide. Sporophylls are scale- like and overlapping, broadly ovate to del- toid, narrowing to an extralong pointed tip. Diagnosis Sporangia are borne adaxially near the sporophyll base and are homosporous. Elongated strobili with long gradually tapering sporophylls. Spores are homosporous Genus Lycopodostrobus Wachtler and clustered on the upper side near the apex. Sporophylls encase the cone at a ju- 2011 venile stage to become increasingly protruding when mature. Diagnosis Description Strobili of Triassic Lycopodiales with helically arranged homosporous sporophylls. Sporo- Several lycophyta cones at all growth stages phyll scales end in a long pointed append- were recovered from Piz da Peres. They help age. Sporangia occur adaxially on the lower elucidate the evolution of club moss cones leaf axils. from the Palaeozoic to the present. While the mostly mature cones from Kühwiesen- kopf yielded no in situ spores, four juvenile Etymology to semi-fully grown strobili from Piz da Pe- After their affinity to the strobili of the Lyco- res exhibited the position of the spores in- podiales. side the sporophylls. Holotype PIZ 98, with 7-cm long preserved parts, shows the or- Lycopodostrobus gaiae WACHT- ganisation of these strobili well. The esti- mated length of the cone body of Lycopo- LER 2011 dostrobus gaiae probably reached 10 to 20 cm in length (KÜH 1276: 15.5 cm long) and 3 to 4 cm in width. The cones can continue Holotype for a further 5 cm, showing a characteristic PIZ 98 tapering appendix. PIZ 213 represents a juvenile specimen with a cone length of 5 cm,

Dolomythos, 2011 32 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.

Wachtler, M.: Lycophyta 33 held by a 1-cm long stalk. Sporophylls are ised by their non-forking stems. Differences attached at an angle of about 80 degrees to in the root system (four-lobed tap root-like the cone apex. The single sporophylls are in Pleuromeia versus creeping and protrud- complete, with a triangular to slightly lan- ing) and different heterosporous sporangia ceolate ending and 2 to 5-cm long pointed from the homosporous structure of Lycopia prolongations. In mature cones (KÜH 1262 dezanchei are other distinguishing features. and KÜH 1426), the characteristic appendi- It seems that Lycopia stands were isolated ces become increasingly protruding, where- among the Mesozoic lycophyta, occupying as in juvenile forms, they tend to encase the mostly the ecological niche of Pleuromeia. cones. The spore-bearing zone, probably re- It was never recorded in the Early-Middle niform, lies on the adaxial side of the sporo- Triassic strata of Piz da Peres, but was very phyll near the axis and is 10 to 15 mm long abundant in the Induan-Olenekian Werfen- and 5 mm wide. Formation of the neighbouring Carinthia and in the global distribution during the Early Triassic. Remarks It should not be completely discarded that Michael Wachtler first discovered Lycopia in Triassic Lycopia has some striking affinity 1999 on the slopes of Kühwiesenkopf in the with extant Lycopodiales and Carbonifer- Braies Dolomites. In the following years, he ous Lepidodendrales. If the descendants collected other materials from Piz da Peres of Triassic Equisetites are observed in the and finally some badly preserved specimens Carboniferous Calamites, this is not so mis- from Val Duron belonging to the younger leading. Most Lepidodendrales are thought Morbiac Limestone of the Illyrian age, which to have become extinct in Europe and North enhanced knowledge on the appearance and America at the end of the Carboniferous, living habitat of this interesting lycophyte. whereas in China, they persisted into the Lycopia constitutes one of the most common Middle Permian. flora elements in the Anisian strata of the A large number, probably more than 350 Dolomites and was in sub-arborescent com- Lepidodendron species, has been described petition with conifers (Voltzia, Alpia and Al- since the genus was first recognized by Kas- bertia) and giant horsetails (Equisetites). par Graf von Sternberg in 1820. Most spe- The generic name Lycopodites was intro- cies are based either on fragments of large duced to describe the isophyllous to aniso- stems lacking leaves or on generally smaller phyllous leaves of the suggested Lycophyta, branches with foliage still attached. Only ranging from Devonian till Pleistocene (TAY- rarely does good evidence of a connection LOR ET AL. 2009). Especially for Palaeozoic between leafy shoots, main axis fragments fossils, it is nearly impossible to distinguish and still fertile organs exist (CHALONER & minute branches from the distal twigs of MEYER-BERTHAUD, 1983). A feature of Lepi- Lepidodendrales, particularly while the at- dodendron is their expanded pyramidal and tached reproduction organs are missing. also diamond-shaped decurrent leaf base. Lycopodites amazonica described from the This is most evident when the leaf has been Middle Devonian of Brazil and Oxrodia gra- abscised and a characteristic leaf cushion cilis from the Mississippian of Scotland are remains, called ”Blattpolster“ by older au- thought to represent herbaceous lycopsids. thors in German. Others, like Synlycostrobus tyrmensis and Lepidostrobus is a loosely defined morpho- Onychiopsis psilotoides, belong to younger genus, named and briefly described by Alex- Cretaceous deposits and can therefore be andre Brongniart after a cone determined for regarded as the descendants of Lycopia. the first time by Parkinson (1804) and cur- Comparing Lycopia to other Triassic lycopsi- rently stored in the London Natural History da, such as Isoetites or Lepacyclotes, shows Museum. For the next 150 years, all cones that they have to be immediately discharged that even vaguely fitted Brongniart’s crite- because of their herbaceous characteristic, ria were included there (BRACK-HANES, S.D. their non-protruding rhizome structure and & THOMAS, B.A., 1983). Since the holotype non-dichotomising branching system. This yielded only microsporangia, but bisporangi- is also the main difference between arbo- ate cones were found in Lepidodendron, a rescent Pleuromeiales, which are character- distinction was made that the morphogenus

Dolomythos, 2011 34 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.

Wachtler, M.: Lycophyta 35 Flemingites has to be used for that kind of are abscised during plant growth, leaving strobilus, whereas Lepidostrobus should on- small subtle circular scars in Lycopia and ly include microsporangiate strobili (TAYLOR bigger ones in Stigmaria. ET AL. 2009). Lepidodendron and Lycopia hold apically di- Although never found in complete organic chotomising axes and do not have ligulate connection, the Lycopodostrobus gaiae stro- leaves. The organography of branching was bili constitute the fertile organ of Lycopia fundamentally dichotomous in extinct Lepi- dezanchei. The cone morphology is com- dodendron, varying from equal to unequal. parable with that of Carboniferous Lepidos- When unequal, overtopping would have oc- trobus cones belonging to Lepidodendron. curred, leading to a determinate branch. They are characterised by their homospor- The same could be stated for Triassic Lyco- ous strobili borne on distal branches in the pia as well as the recent Lycopodiales. Both crown of the tree. Such strobili found in also bear homosporous strobili. Lycopia the Stephanian sediments on Stangnock in could be considered a “Giant Lycopod” in the Carinthia also bear typical protruding leafy same manner as the “Giant horsetails Equi- appendices. Only the size of Triassic Lyco- setites” from the same Anisian strata. podostrobus is smaller, reaching half of the Altogether, Lycopia constitutes a highly in- Carboniferous one in the best cases. teresting plant and, if not a direct descend- Of interest is the almost same manner in ant of giant Lepidodendron, it could at least which lycophyte could be regarded as a de- be integrated into the same family. It seems scendant of Lycopia. Compared to the extant that the arborescent lycophyte crossed the Lycopodiopsida, the genera Isoetales and Permian by reducing their size enormously Selaginellas have to be eliminated because and survived the Mesozoic by further de- of their totally different composition, root creasing its size to occupy ecological niches system and mainly heterosporous reproduc- as dormant dwarfs, maybe increasing in size tion. The discovery of abundant material of when circumstances changed. Isoetaceae and Selaginellaceae in the Ani- sian layers consolidates the argument that General discussion other relatives have to be searched for. Many more affinities have been discovered Why I conducted this research with the extant group of Lycopodiales. Ex- Since 1990, I spent most of my time in tant Lycopodiales are represented by four the mountains around Piz da Peres. First, genera (about 480 species in Lycopodium, I had to learn how to walk and then hear 430 species in Huperzia, 41 species in Ly- the sound of the past, at least to see inside copodiella and ones species in Phylloglos- the rocks. This took years and years of hard sum. All are homosporous, generally di- work. Only after that, I felt mature enough chotomously branched and represent small to understand the rocks. At the end, the herbaceous plants. Their stems arise ad- rocks began to tell me their living history. ventitiously from a horizontal rhizome. The Thus, I discovered a lot of new and, at that Lycopodiales have a global distribution, but time, unknown fossils. I often read in books their favoured habitat spans mainly from the manner in which I can understand the temperate to subarctic regions. past, but the best way is by exploring and Both extant Lycopodiales and Lycopia show searching until you have the missing links a comparable root system with a reptant in your hand. These are then the incontro- stem and rootlets. Lycopia also bears either vertible arguments for understanding how aboveground or underground running rhi- life evolved or how they could have evolved. zomes or lateral branches. The abscised leaf Unfortunately, we now live in a time when cushions are pyramidal in Lepidodendron many scientists perform their studies in the and more rhomboidal in Lycopia, but simi- privacy of their own home and not in nature. lar in every case. The robot-like structure of Although cladistic analyses on a writing Lepidodendron, commonly called Stigmaria, desk can be useful, they are nothing against resembles the rhizomes of Lycopia in many many years of studying in the wilderness. parts, with its shallow propagation structure All that time, I lied in wet soil or scram- and helically-arranged lateral appendages. bled onto exposed rocks. Avalanches buried In Stigmaria and Lycopia, the appendages me, I was struck by lightning, and I suf-

Dolomythos, 2011 36 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.

Wachtler, M.: Lycophyta 37 fered from cold and heat. However, I felt dominated vegetation around the world and like a free man, something that I could not were the largest known living organisms in affirm from the people in the valleys, who the Carboniferous. The oldest representa- had to carry out often nonsensical work. tives of the group are found in the Early Sometimes, my absence from normal life Devonian with Baragwanathia and Drepano- was noticed and the police put pressure on phycus (SCHWEITZER, 1990). The presence me with their guns. I saw villages expand- of lycopods - from herbaceous to arbores- ing and an increasing number of ski slopes cent - can be traced continuously from then that restricted the areas I was investigat- until the present. All the characteristics that ing. Shortly, the authorities imposed a re- typify Lycopodium today, such as creeping, search ban on me and punished me. How- dichotomous and anisotomous branching, ever, I never suffered from boredom or numerous spirally-arranged microphylls, and desperation. I said to myself that if I helped reproduction by spores produced in large to decode only one drop of water in the big sporangia borne adaxially to the leaves, re- ocean of science, my life was just worth liv- main remarkably unchanged from the old- ing. est member of the group (HUEBER, 1983). The significance of lycopods in the past Selaginellaceae has also been recorded in Although extant lycophytes constitute a the Carboniferous, whereas the oldest mem- modest group of herbaceous plants, they bers of Isoetaceae have only been recorded have played an enormous role in the history as far back as the Early Triassic in Australia of land vegetation and have the longest fos- (Isoetes beestonii; RETALLACK, 1975); the sil record of any vascular plant, extending Alps (Isoetites brandneri; WACHTLER ET AL. over 400 million years from the Early De- 2010) certainly have to date further back vonian, maybe from the late Silurian until because just in that time of global distribu- the present (CLEAL & THOMAS, 2009). They tion.

An entirely recovered Lepidodendron stem from the Carinthian Stangnock area with a diameter of up to one metre (Collection Georg Kandutsch).

Dolomythos, 2011 38 The modification of landscape from ta (Equisetites). The flora had no dominant species, as in the following Early Triassic the Carboniferous to the Triassic with Pleuromeia, but was not so desert- An impoverished flora after the Carbon- like as is often thought. Nevertheless, the iferous. In a geologically short time, the plants were very reduced in size, as well as dominant giant lycophyta disappeared com- scrubby and prickly, the result of an enor- pletely. In the lower Permian, only isolated mous flat and semi-arid grassland savannah remains were found (Rotliegend: Subsigil- crossed by many meandering rivers from a laria brardii and Lepidostrobus) (BARTHEL, 2009). This is also valid for the Alps, where only some doubtful Stigmaria ficoides roots from the Early Permian Laas-Formation and Wunderstätten have been published (FRITZ ET AL. 1990; WANK & PRINZ, 1994). A hunger flora in the Late Permian. In all Europe in the Late Permian, there was a species-poor flora with reduced foliage structures. As in the German Zechstein and Dolomitian Late Permian (Monte Ozol, Blet- terbach and Seceda), we encounter only some spinous conifers (Ullmannia, Pseudo- voltzia and Ortiseia), microleaved peltaspermales (Peltaspermum martinsii), gingkophyta (Sphenobaiera), early low-growing cycadophyta (Taeniopteris) and Sphenophy-

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 Compared to the Carboniferous Lepidodendron cones, attached bisporangiate cones and well developed pla- Lycopia strobili were nearly as large, but from half to nar branching ovate to lanceoolote leaves with acute one-third smaller in length. The composition of long apices. Westfal D. Zwickau. Collection Jürgen Meyer, and protruding leafy appendices on sporophylls is sur- Zwickau prisingly similar.

Wachtler, M.: Lycophyta 39 humid hinterland, such as the current Oka- global plant species in that period, rang- vango delta in Botswana. ing from the Carinthian mountains (Gail- Huge delta swamps in the Early Trias- taler Alps, Karawanken), Germany (Pleu- sic. In the Early Triassic, big parts of Europe romeia sternbergii), over the Caspian Sea to and other landscapes consisted of enormous Wladiwostok, China and Japan (Pleuromeia flat basins and delta swamps. The palaeoe- rossica, P. obrutschewii, P. epicharis, P. ji- cological milieu was more humid with an in- aochengensis and P. hattai), even occur- tertonguing of river channels and floodplains ring in Australian locations from the for- and lakes (MADER, 1990). This would have mer Gondwana continent (P. longicaulis). It been consistent with the presence of Pleu- seems plausible, as stated by other authors romeia and the abundant richness of humid- (RETALLACK, 1975; LOOY ET AL. 1999; TAY- loving Equisetacea (Equisetites mougeotii), LOR ET AL. 2009), that the Pleuromeiales whereas other plants (the ferns Anomopteris apparently came from nothing, dominated mougeotii and Neuropteridium, the conifer the planet for a short geological time and Voltzia heterophylla, and the seed fern Scy- then became completely extinct on the Ole- tophyllum bergeri) filled the niches of drier nekian-Anisian boundary. It is possible that landscapes or higher-lying areas. Thus, the they are the descendants of Carboniferous dominance of Pleuromeia and Equisetites is Sigillariaceae. Tracing an evolutionary line explainable without catastrophism theories. from Pleuromeiales to any extant lycopods The short heyday of Pleuromeiales. like Isoëtes is highly improbable because The lycophyte Pleuromeia sternbergii is re- Isoetales co-inhabited with Pleuromeiales in corded extensively in the Induan-Olenekian the Early Triassic. and constitutes one of the most well-known

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 40 The irresistible ascension of land plants their ecological niche, even though very in the Early-Middle Triassic. After nearly reduced in size and diffusion. Only the Pel- 50 million years of a difficult time for plants taspermales (Scytophyllum, Sagenopteris), beginning from the Early Permian to the although highly present in the Early-Middle Early Triassic, we relive the rise and spread Triassic, seemed to not have survived the of mainly all extant plants in the Anisian. A Jurassic-Creataceous boundary. lot of new-age ferns (Gordonopteris, Maran- The importance of the new lycopod spe- toidea, Ladinopteris, Sphenopteris, Anom- cies from Piz da Peres. The discovery of opteris, Neuropteridium, Wachtleria and several new species and one new genus of Marantoidea) belonging to the Marattiacea, Lycophytes in the lower Middle Triassic of Gleichenaceae, Osmundacea or Lindsae- the Southern Alps shed some interesting aceae were highly diffused. This is also valid light on the evolution of this group. The Tri- for the conifers dividing them in Voltziaceae assic was a time of important radiation and and Alpiaceae, which evolved into mainly all change in several key morphological charac- conifers present nowadays like Araucarians, teristics of this group. After the predominant Pinaceae and Cupressaceae. There was an position of giant Lycopods in the Carbonifer- enormous diversification of the cycadophy- ous, there was a decline in the Permian and ta (Bjuvia and Nilssonia), with a splitting of a much reduced level of radiation of dwarf mainly all living cycad genera and a rise in lycopods in the Early Triassic, with nearly all an unusual group of proto-angiospermales groups present then still extant now. (Ladinia and Pizperesia). Moreover, the lyco- Possible geothermal events as a cause phyta (Lycopia, Isoetites and Selaginellites) of this unusual lycophyta assemblage. and sphenophyta (Equisetites) maintained The early Middle Triassic Kühwiesenkopf

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.

Wachtler, M.: Lycophyta 41 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 42 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

Wachtler, M.: Lycophyta 43 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 of ronmental conditions allowed unusual plant Selaginellales. In the Early Triassic, we en- assemblages and combinations of floras, like counter Selaginella species with isotophyl- the club moss vegetation capable of surviv- lous (Selaginellites leonardii) and also an- ing high-stress conditions of soil minerali- isophyllous leaves (Selaginellites venieri). sation and temperature. However, marked The advantage of one or the other could differences in vegetation composition and not be explained satisfactorily. Furthermore, physiognomy were not so distinctive to de- there is currently a nearly equal distribution duce further speculations regarding global of anysophyllos genera like Heterostachys catastrophic events. and Stachygynandrum, or isophyllous spe- The invariability of Isoetaceae. The quill- cies such as Tetragonostachys, Selaginella worts maintained their aspect mainly un- or Ericetorum. Their heterosporophylly is an varied at least from the Early Triassic until old attribute, still remaining unvaried. The today. Their evolutionary line traces more same is true for their often branching and to the group of Carboniferous Sigillariaceae low-growing characteristics. than the Lepidodendrales, even though this The interesting evolution of Lyco- hypothesis is not completely acceptable. pia. One of the most surprising discover-

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 44 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 flo- The showdown of the dinosaurs. The Ani- ras of Tethydealan Dolomites and can be de- sian layers of Piz da Peres also hold a fined as Giant horsetails or Giant Lycopods. rich ichnofauna dominated by Archosau- In addition, both have extant Lycopodium riformes (Chirotherium, Brachychiroth- and Equisetes as their direct descendants erium and Isochirotherium) and some Di- and probably originated in the colossal Lepi- nosauromorpha (Parasphingopus ladinicus dodendron or Calamites trees of the Carbon- nov. gen. n. sp.; AVANZINI & WACHTLER, iferous. In contrast to Isoetites or Selagin- 2011). The emergence of the dinosaurs ellites, which were always small-sized, they was therefore linked to the transforma- became smaller over the million years until tion of vegetation. The increasingly wide- today as dwarf scrubbed plants in the copse spread bushlands benefitted tetrapods, of other vegetation. Astonishingly, Lycopia which were able to elevate themselves resembling lycopods were, until now, never and walk two-legged. Parasphingopus, recorded from other parts of the world or in with its reduced forelegs and three-clawed the Early Triassic and following periods. Oth- hind leg, achieved all the properties of er midget characteristics have been noted in true dinosaurs that followed a little later. the Early Triassic, with the shrubby Voltzia In this Early Triassic, there was the dawn conifer Aethophyllum stipulare in contrast of a long-lasting alliance and partnership to the arborescent Voltziales, as well as the between animals and plants.

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.

Wachtler, M.: Lycophyta 45 of the fossil quillworts called Isoetites. Journal of the References Washington Academy of Sciences, 48 (11), 358-361. AMERON, H. W. J., van BOERSMA M. 1974. Vorläufige CANTRILL, D. J., and WEBB, J. A. 1998. Permineralized Untersuchungsergebnisse an älteren und neu aufgesa- pleuromeid lycopsid remains from the Early Triassic mmelten jungpaläozoischen Pflanzenfunden der Ostal- Arcadia Formation, Queensland, Australia. Review of pen (Österreich). – Carinthia II, 84: 9–15, Klagenfurt. Palaeobotany and Palynology, 102, 189-211. AMERON, H. W. J. van, BOERSMA, M. NlEDERMAYR, CASSINIS, G., PEROTTI C. R. 2007. A stratigraphic and G., 1976. Bericht über erste Pflanzenfunde aus der tectonic review of the Italian Southern Alpine Permian, Trias-Basis der westlichen Gailtaler Alpen bei Kötschach Palaeoworld 16 – 140- 172 (Kärnten, Österreich), Carinthia II, S. 103-112 Klagen- CHALONER, W. 1967. Lycophyta. In: BOUREAU, E. furt. (Ed), Traité de Paleobotanique. Tome II. Bryophyta, Psi- ANDREWS, H. N. 1961. Studies in Palaeobotany. Wiley lophyta et Lycophyta. Masson et Cie Editeurs. Paris, p. and Sons Ed., New York and London, 487 pp. 437–802, 845 pp. ASH, S. R. 1972. Late Triassic plants from the Chinle CHALONER. W. G., & MEYER-BERTHAUD. B. 1983. Leaf Formation in North-Eastern Arizona. Palaeontology, and stem growth in the Lepidodendrales. Botanical 15(4), 598-618. Journal of the Linnean Society. 86, 135-48. ANDREWS, H. N. 1979. Skilliostrobus gen. nov., a new CLEAL, C. J. & THOMAS, B. A. 2009. Introduction to lycopsid cone from the early Triassic of Australia. Alche- plant fossils. Cambridge University Press, Cambridge, ringa, 3, 73-89. 237 pp. AVANZINI M., WACHTLER, M. 2011. Parasphingopus la- COLLINSON, M. E. 1991. Diversification of modern dinicus igen. nov., isp. nov, from the Anisian of Pragser heterosporous pteridophytes. In BLACKMORE, S. and Dolomiten (Southern Alps, Italy), in press. BARNER, S. H. (eds.). Pollen and Spores. Systematic BALME, B. E. 1963. Plant microfossils from the Lower Association Special Volume, 44, 119-150 (Clarence Triassic of Western Australia. Paleontology, 6 (1), 12-40. Press, Oxford). BANERJI, J. 1989. Some Mesozoic plant remains from COUPER, R. A. 1958. British Mesozoic microspores and Bhuj Formation with remarks on the depositional envi- pollen grains a systematic and stratigraphic study. Pal- ronment of beds. The Palaeobotanist, 37, 159-168. aeontographica, 103B, 75-179. BARTHEL, M. 2009. Die Rotliegendflora des Thüringer DAUGHERTY, L. H. 1941. The Upper Triassic Flora of Waldes. – Veröffentlichungen des Naturhistorischen Mu- Arizona. Carnegie Institute, Washington Publications, seums Schleusingen 526, 1-108. BECHSTÄDT, T., BRANDNER, R. 1970. Das Anis zwisch- DE JERSEY, N. J. 1979. Palynology of the Permian-Tri- en St. Vigil und dem Höhlensteintal (Pragser-und Ol- assic Transition in the Western Bowen basin. Geological anger Dolomiten, Südtirol). 9-103. In MOSTLER, H. Survey of Queensland, 374 (46), 1-39. (ed). Beiträge zur Mikrofazies und Stratigraphie von Ti- DE ZANCHE, V., FRANZIN, A., GIANOLLA, P., MIETTO, rol und Vorarlberg (Festband des Geol. Inst., 300-Jahr- P., SIORPAES, C. 1992. The Piz da Peres Section (Vald- Feier Univ. Innsbruck. Wagner aora-Olang, Pusteria Valley, Italy): a reappraisal of the BLANCKENHORN, M. 1886. Die fossile Flora des Bunt- Anisian stratigraphy in the Dolomites. Eclogae Geologi- sandsteins und des Muschelkalks der Umgegend von cae Helveticae, 85, 127–143. Commern. Palaeontographica, 32B, 117-153. DE ZANCHE, V., GIANOLLA, P., MIETTO, P., SIORPAES, BRANDNER, R., HORACEK, M., KEIM, L., SCHOLGER, R. C., VAIL, P. R. 1993. Triassic sequence stratigraphy in 2009. The Pufels/Bulla road section: deciphering envi- the Dolomites (Italy). - Mem. Sci. Geol., 45: 1 – 27. ronmental changes across the permian-Triassic bound- DETTMANN, M. E. 1961. Lower Mesozoic megaspores ary to the Olenekian by integrated litho-, magneto- and from Tasmania and South Australia. Micropaleontology, isotope stratigraphy. A field trip guide. In: GEO.ALP 6, 7 (1), 71-86. S. 116 - 132. DOBRUSKINA, I. A. 1974. Triassic Lepidophytes. Pale- BOCK, W. 1962. A study on fossil Isoetes. Journal of ontogical Zhurnal, 3, 111-124. Paleontology, 36, 53-59. DOBRUSKINA, I. A. 1985. Some problems of the sys- BOSE, M.N. and ROY, S.K. 1964. Studies on the upper tematics of the Triassic lepidophytes. Paleontogical Gondwana of Kutch - 2. Isoetaceae. Paleobotanist, 12, Zhurnal, 19(3), 74-88. 226-227 EMMONS, E., 1856. Geological report of the Midland BRACK-HANES, S.D., THOMAS, B.A., 1983: A re-exami- Counties of North Carolina. George P. Putnam and Co., nation of Lepidostrobus Brongniart, Botanical journal of New York, 352 pp. the Linnean Society. EMMONS, E., 1857. American Geology, part 6. Sprague, BROGLIO LORIGA, C., FUGAGNOLI, A., VAN KONIJNEN- Albany, New York, 152 pp. BURG-VAN CITTERT, J. H. A., KUSTATSCHER, E., PO- ERKAN, E. 1977. Uran- und gipsführendes Permoskyth SENATO, R. and WACHTLER, M. 2002. The Anisian mac- der östlichen Ostalpen, 1 Jahrb. Geol. B.-A. Band 120, roflora from the Northern Dolomites (Monte Prà della Heft 2 S. 343—400 Wien. Vacca/Kühwiesenkopf, Braies): a first report. Rivista Italiana di Paleontologia e Stratigrafia, 108, 381-390. FLICHE, P. 1910. Flore fossile du Trias en Lorraine et Franche-Comté. Ed. Berger-Levrault, Paris, Nancy, 297 BROWN, R.W., 1939. Some American fossil plants be- pp. longing to the Isoetales. Journal of the Washington Academy of Sciences, 29 (16), 261-269. FONTAINE, W.M. 1883. Contributions to the Knowledge of the Older Mesozoic Flora of Virginia. United States BROWN, R.W. 1958. Paleobotany. – New occurrences Geological Survey Monograph, 6, 1-144.

Dolomythos, 2011 46 FRENTZEN, K. 1914. Die Flora des Bundsandsteins KANDUTSCH G., WACHTLER M. 2011. Fossile Pflanzen Badens. Mitteilungen des Großherzolichen Badischen der Frühen Trias aus den Gailtaler Alpen, in press Geologischen Landesanstalt, 8, 63-158. KELBER, K.-P. and HANSCH, W. 1995. Keuperpflanzen. FRITZ, A. BOERSMA, M., KRAINER, K. 1990. Die Enträtselung einer über 200 Millionen Jahre alten Steinkohlenzeitliche Pflanzenfossilien aus Kärnten.- Flora. Museo, 11, 1-157. Carinthia II, Sonderheft KELBER, K.-P. .1999. Neue Befunde über die Schachtel- FUGAGNOLI, A. and POSENATO, R. 2004. Middle Trias- halme des Keupers.- In: Hauschke, N. & Wilde, V. sic (Anisian) benthic Foraminifera from the Monte Prà (eds.): Trias - Eine ganz andere Welt, III.14: 355-370; della Vacca/Kühwiesenkopf section (Dont Fm, Braies (Verl. Dr. F. Pfeil), München. Dolomites, Northern Italy). Bollettino della Società Pa- KENRICK, P. and CRANE, P.R. 1997. The Origin and leontologica Italiana, 43 (3), 347-360. Early Diversification of Land Plants: A Cladistic Study. GEYER, G. 1901. Geologische Aufnahmen im Weissen- Smithsonian Institution Press, Washington DC, 441 pp. bachthale , Kreuzengraben und in der Spitzegelkette KRASSILOV, V. A. and ZAKHAROV, Y. D. 1975. Pleu- (Oberkärnten). Verhandlungen der k. k. geologischen romeia from the Lower Triassic of the Far East of the Reichsanstalt, Wien. U.S.S.R. Review of Palaeobotany and Palynology, 19, GÖTZINGER, M. 1985. Mineralisationen in den Guten- 221-232. steiner Schichten (Anis) in Ostösterreich, Arch. f. KELBER, K.-P., VAN KONIJNENBURG-VAN CITTERT, J. Lagerst.forsch. Geol. B.-A. ISSN 0253-097X Band 6 H. A. 1998. Equisetites arenaceus from the Upper Trias- S.183-192 Wien, Juni 1985 sic of Germany with evidence for reproductive strate- GRAUVOGEL-STAMM, L. 1978. La flore du Grès à Volt- gies. Review of Palaeobotany and Palynology, 100, 1–6. zia (Buntsandstein supérieur) des Vosges du Nord KUSTATSCHER, E., WACHTLER, M. VAN KONIJNEN- (France). Sciences Géologiques, Mémoirs, 50, 1-225. BURG-VAN CITTERT, J. H. A. 2007. Horsetails and seed- GRAUVOGEL-STAMM, L. 1993. Pleuromeia sternbergii ferns from the Middle Triassic (Anisian) locality Küh- (Münster) Corda from the Lower Triassic Germany – wiesenkopf (Monte Prà della Vacca) in the Dolomites further observations and comparative morphology of its (Northern Italy). Palaeontology, 50 (5), 1277–1298. rooting organ. Review of Palaeobotany and Palynology, KRASSILOV, V. A. 1978. Mesozoic lycopods and ferns 77, 185-212. from the Bureja basin. – Palaeontographica, Abt. B., GRAUVOGEL-STAMM, L., DURINGER, P. 1983. Annalepis 166 (1-3): 16-29. zeilleri Fliche 1910 emend., un organe reproducteur de LEE, H. H. 1951. On some Selaginellites remains from Lycophyte de la Lettenkohle de l‘Est de la France. Mor- the Tatung Coal series. Science Record, 4 (2), 193-196. phologie, spores in situ et paléoécologie. Geologische Rundschau, 72, 23-51. LESCHIK, G. 1955. Die Keuperflora von Neuwelt bei Basel. II. Die Iso- und Microsporen. Schweizer Paläon- GRAUVOGEL-STAMM, L. and LUGARDON, B. 2001. The tologische Abhandlungen, 72, 1-70. Triassic Lycopsids Pleuromeia and Annalepis: Relation- ships, Evolution, and Origin. American Fern Journal, 91 LINDLEY, J., and HUTTON, W. 1836. The fossil flora of (3), 115-149. Great Britain. London, 208 pp. GREUTER W., MCNEILL J., BARRIE F.R., BURDET H. LOOY, C. V., COLLINSON, M. E., VAN KONIJNENBURG- M., DEMOULIN V., FILGUEIRAS T. S., NICOLSON D. H., VAN CITTERT, J. H. A., VISSCHER, H., BRAIN, A. P. SILVA P. C., SKOG J. E., TREHANE P., TURLAND N. J. R. (2005): The ultrastructure and botanical affinity of and HAWKSWORTH D. L. 2000. International Code of end-Permian spore tetrads. - Int. J.Plant Sci., 166(5): Botanical Nomenclature (Saint Louis Code). Koeltz Sci- 875-887. entific Books, Königstein, Germany, 474 pp. LUGARDON, B., GRAUVOGEL-STAMM, L., DOBRUSKINA, HALLE, T. G. 1907. Einige krautartige Lycopodiaceen I. 1999. The microspores of Pleuromeia rossica Neu- paläozoischen und mesozoischen Alters. Kungliga Sven- burg (Lycopsida, Triassic): comparative ultrastructure ska Vetenskapsakademiens Handlingar, 7 (5), 1-17. and phylogenetic implications. - Compt. Rend. Acad. Scienc., ser. II, fasc. A, Scienc. Terr. Plan., 329 (6): HARRIS, T. M. 1935. The fossil Flora of Scoresby Sound, 435–442. East Greenland. Meddelelser om Grønland, 112, 1-176. LUGARDON, B., GRAUVOGEL-STAMM, L., DOBRUSKINA, HELBY, R. and MARTIN A. R. H., 1965. Cyclostrobus I., 2000. Comparative ultrastructure of the megaspores gen. nov., cones of lycopsidean plants from the Nar- of the Triassic lycopsid Pleuromeia rossica Neuburg. rabeen group (Triassic) of New South Wales. Australian – Compt. Rend. Acad. Scienc., ser. II, fasc. A, Scienc. Journal of Botany, 13, 389-404. Terr. Plan., 330 (7), 501–508. HOOKER, W. J. 1861. The British ferns. L. Reeve, Lon- LUNDBLAD, B. 1948. Selaginelloid Strobilus from East don. 305 pp. Greenland (Triassic). Meddlilser fra Dansk Geologish Hueber, FM. 1983. A new species of Baragwanathia Forening Kobenhavn, 11 (3), 351-363. from the Sextant Formation (Emsian) Northern Ontario, LUNDBLAD, B. 1950a. Studies in the Rhaeto-Liassic Canada. Journal of the Linnean Society (Botany) 86, Floras of Sweden: I: 1 Pteridophyta, Pteridospermae, 57-79. and Cycadophyta from the mining district of NW Sca- JUNG, W. W., 1958. Zur Biologie und morphologie eini- nia. Kungliga Svenska Vetenskapsakademiens Handlin- ger disperser Megasporen, vergleichbar mit solchen von gar, ser. 4, 1 (8), 5-82. Lycostrobus scottii, aus dem Rhät-Lias Frankens. Geolo- LUNDBLAD, B. 1950b. On a fossil Selaginella from the gisches Blatt Nordost-Bayern, 8 (3), 114-130. Rhaetic Hyllinge, Scania. Svenska Botaniisk Tidskrift, KARRFALT, E., 1986. New observations on the structure 44 (3), 475-487. and development of Nathorstianella. Review of Palaeo- MADER, D. 1990. Palaeoecology of the flora in Bunt- botany and Palynology, 47, 1-8.

Wachtler, M.: Lycophyta 47 sandstein and Keuper in the Triassic of Middle Europe. RENESTO S., POSENATO R. 2003. A new Lepidosauro- [2 vols.]. Fischer, Stuttgart. 1582 pp. 71 fig. 103 plts. morph reptile from the Middle Triassic of the Dolomites MÄGDEFRAU, K. 1931. Zur Morphologie und phylogene- (Northern Italy). Rivista Italiana di Paleontologia e tischen Bedeutung der fossilen Pflanzengattung Pleuro- Stratigrafia, 109(3), p. 463-474. meia. Beiheften zum Botanischen Centralblatt, 48 (2), RETALLACK, G. J., 1975. The life and times of a Triassic 119-140. lycopod. Alcheringa 1, 3-29. ISSN 0311-5518. MÄGDEFRAU, K. 1956. Paläobiologie der Pflanzen, RETALLACK, G. J. 1997. Earliest Triassic origin of Stuttgart 1968 Isoetes and Quillwort evolutionary radiation. Journal of MENG FANSONG, 1996. Middle Triassic Lycopsids flora Palaeontology, 71 (3), 500-521. of South China and its palaeoecological significance. Pa- RETALLACK, G. J., METZGER, C. A., GRAVER, T., laeobotanist India, 45, 334-343. JAHREN, A. H., SMITH, R. M. H., SHELDON, N. D. MENG FANSONG 1998. Studies on Annalepis from (2006): Middle-Late Permian mass extinction on land. – Middle Triassic along the Yangtze Valley and its bearing Geol. Soc. Amer. Bull., 118: 1398-1411. on the Origin of Isoetes. Acta Botanica Sinica, 40 (8), RICHTER, P. B. 1909. Beiträge zur Flora der unteren 768-774. Kreide Quedlinburgs, II. Die Gattung Nathorstiana und MENG FANSONG, ZHANG ZHENLAI, NIU ZHIJUN and Cylindrites spongioides. Leipzig, 12 pp. CHEN DAYOU 2000. Primitive lycopsid flora in the Yang- ROTHWELL, G.W. and ERWIN, D.M. 1985. The rhizo- tze Valles of China and systematics and evolution of morph apex of Paurodendron; implications for homolo- Isoetales. 70-85. In: Changsha = Science and Technol- gies among the rooting organs of Lycopsida. American ogy Press, Hunan Province, 87 pp. Journal of Botany, 72, 86–98. M. MENNING & A. HENDRICH , 2002. Stratigraphische SADOVNIKOV, G.N. 1982. The morphology, systematics Tabelle von Deutschland. Deutsche Stratigraphische and distribution of the genus Tomiostrobus. Paleonto- Kommission logical Zhournal, 1982 (2), 100-109. MOUGEOT, A. 1849. Observations sur le Caulopteris Le- SCHIMPER, W. P. 1869. Traité de Paléontologie végétale sangeana (Schimper et Mougeot) et nécessité de créer ou la flore du monde primitif dans ses rapports avec les pour ce végétal fossile un genre nouveau dédié à Lesa- formations géologiques et la flore du monde actuel. I. ing. Annales de la Societé d’Emulation des Vosges, 7, J. B. Baillère et Fils éd., Paris, 738 pp. 7-16. SCHIMPER, W. P. and MOUGEOT, A. 1844. Monographie MOSTLER, H.: Die permoskythische Transgressions- des Plantes fossiles du Grès Bigarré de la Chaine des Serie der Gailtaler Alpen. — Mitt. Ges. Geol. Berg- Vosges. Ed. G. Engelmann, Leipzig, 83 pp. baustud., 20, S. 143—149, Wien 1972. SCHWEITZER H.-J, 1990. Pflanzen erobern das Land. MÜNSTER G., 1842. Beiträge zur Petrefacten-Kunde, Kleine Senckenberg-Reihe Nr. 18, Frankfurt am Main. Band 5, In Commission der Buchner’schen Buchhand- SCHWEITZER, H.-J., VAN KONIJNENBURG-VAN CITTERT, lung. J. H. A. AND VAN DER BURGH, J., 1997. The Rhaeto- NATHORST, A. G. 1908 Paläobotanische Mitteilungen, 3. Jurassic flora of Iran and Afghanistan. 10. Bryophyta, Lycostrobus scotti, eine grosse Sporophyllähre aus den Lycophyta, Sphenophyta, Pterophyta-Eusporangiatae rätischen Ablagerungen Schonens. Kungliga Svenska and -Protoleptosporangiatae.. Palaeontographica, 243B, Vetenskapsakademiens Handlingar, 43,1–9. 103-192. PASSONI, L., VAN KONIJNENBURG-VAN CITTERT, J. H. SENOWBARI-DARYAN, B., ZÜHLKE, R., BECHSTÄDT, T. A., 2003. New taxa of fossil Carnian plants from Mount and FLÜGEL, E. 1993. Anisian (Middle Triassic) Buildups Pora (Bergamasc Alps, Northern Italy). Review of Pa- of the Northern Dolomites (Italy): The Recovery of Reef leobotany and Palynology, 123, 321-346. Communities after the Perm/Triassic Crisis. Facies, 28, PIA, J. 1937. Stratigraphie und Tektonik der Pragser 181-256. Dolomiten in Südtirol, Wien. Selbstverl.. 1937. XV, 248 SEWARD, A. C. 1910. Fossil Plants II – A Textbook for S. : Ill., graph. Darst., Kt.. students of Botany and Geology. Cambridge University PIGG, K. B. 1992. Evolution of Isoetalean Lycopsids. Press, Cambridge, 624 pp. Annals of the Missouri Botanic Garden, 79, 589-612. SEWARD, A. C. 1913. A British Fossil Selaginella. New PIGG, K. B. 2001. Isoetalean Lycopsid Evolution: from Phytologist, 12 (3), 85-88. the Devonian to the Present. American Fern Journal, 91 SEWARD, A. C. 1917. Fossil plants, III Pteridospermae, (3), 99-114. Cycadofilices, Cordaitales, Cycadophyta. University POSENATO, R. 2008. Anisian (Middle Triassic) bivalves Press, Cambridge, 559 pp. from the Dolomites. N. Jb. Geol. Paläont. Abh.2008, SEWARD, A. C. 1931. Plant life through the ages – A vol. 247/1, p. 93–115, Stuttgart, January, published geological and botanical retrospect. University Press, online Cambridge, 603 pp. POTONIÉ, H. 1902. Pleuromeiacae. In: Enger A (ed.), SKOG, J. E., and HILL, C. R. 1992. The Mesozoic herba- Die natürlichen Pflanznefossilien von Engler, Teil 4, 754- ceous lycopsids. Annals of the Missouri Botanical Gar- 756, Leipzig. den, 79 (3), 648-675. POTONIÉ, H. 1904. Abbildungen und Beschreibungen SKOG, J. E., DILCHER, D. L. and POTTER, F. W. 1992. fossiler Pflanzen-Reste der paleozoischen und mesozois- A new species of Isoetites from the mid- Cretaceous chen Formationen, Königliche Preussische Geologische Dakota Group of Kansas and Nebraska. American Fern Landes-Anstalt und Bergakademie, II, 38-39, 1-15. Journal, 82, 151-161. PRANTL, K. 1874. Lehrbuch der Botanik. Engelmann, SNIGIREVSKAYA, N. S. 1980. Takhtajanodoxa Snig., a Leipzig. 240 pp. new link in evolution of lycopsids. In ZHILIN S.G. (ed.)

Dolomythos, 2011 48 Systematika i evolyutsiya vysshikh rastenii, Nauka, Middle Triassic (Anisian) locality Kühwiesenkopf (Monte Leningrad, 45-53, 122 pp. [In Russian] Prà della Vacca) in the Dolomites (Northern Italy). – SRIVASTAVA G. K., D. D. PANT, and P. K. SHUKULA. Palaeontology, 53(3) 1993. The Genus Isoetes L. in India. American Fern WACHTLER, M., 2011. Ferns and seed ferns from the Journal, 83 (4), 105-119. Early-Middle Triassic (Anisian) Piz da Peres (Dolomites - SPIEKER, T. 1854. Pleuromeia, eine neue fossile Pflan- Northern Italy), Dolomythos, 57-79, Innichen zengattung und ihre Arten, gebildet aus Sigillaria WACHTLER, M., 2011. Seed ferns from the Early-Middle Sternbergi Münst. des bunten Sandsteins zu Bernburg. Triassic (Anisian) Piz da Peres (Dolomites - Northern Zeitschrift für die Gesammten Naturwissenschaften, Italy), Dolomythos, 88-104, Innichen. 1852 (3), 177-191. WACHTLER, M., 2011. Evolutionary lines of conifers THIEDIG, F., KABON, H. 2011. Neue fossile Pflanzen- from the Early-Middle Triassic (Anisian) Piz da Peres funde von Pleuromeia sternbergii CORDA aus den (Dolomites - Northern Italy), Dolomythos, 3-72 In- Werfener Schichten (Olenekium-Stufe, Unter-Trias) im nichen. Oberostalpin Kärntens, Carinthia, Klagenfurt. 203-224. WALKOM, A.B, 1941. Fossil plants from Gingin W.A. THOMAS, B. A. 1992. Paleozoic herbaceous lycopsids Journal of the Royal Society of Western Australia, 28, and the beginnings of extant Lycopodium sens. lat. and 201-208. Selaginella sens. lat. – Ann. Miss. Bot. Garden, 79: WANG Qi 2008. Proposal to conserve the name Pleu- 624-631. romeia with that spelling (fossil lycopsida). TAXON, 57 TOLLMANN, A. 1968. Beitrag zur Frage der Skyth-Anis- (2), 659-660. Grenze in der zentralalpinen Fazies der Ostalpen. Verh. WANG ZIQIANG 1991. Advances on the Permo-Triassic Geol. B.-A., Jg. 1968, S. 28—45, Wien. lycopods in North. China. I. An Isoetes from the mid- TOLLMANN, A. 1968. Tektonische Karte der Nördlichen Triassic in Northern Shaanxi. province. Palaeontographi- Kalkalpen, 2. Teil. — Mitt. Geol. Ges. Wien, 61 124— ca, Abt. B, 222 (1-3), 1-30. 181, Taf. 1, Wien 1969. WANG ZIQIANG 1996. Recovery of vegetation from the TOLLMANN, A. 1972. Die Neuergebnisse über die Trias- terminal Permian mass extinction in North China. Re- Stratigraphie der Ostalpen. — Mitt. Ges. Geol. Berg- view of Palaeobotany and Palynology, 91, 121-142. baustud., 21, 65—113, Innsbruck. WANG ZIQIANG and WANG LIXIN 1990. New plant as- TOWNROW, J.A. 1968. A fossil Selaginella from the semblages from the bottom of the Mid-Triassic Ermay- Permian of New South Wales. - J. Linn. Soc. London ing Formation. Shanxi Geology, 5 (4), 303-318. Bot., 61: 13-23. WANK, M., PRINZ, G., 1994 Ein Schuppenbaumgewächs TAYLOR, W. A. 1989. Megaspore wall organization in bei Wunderstätten (St. Pauler Berge, Kärnten), Carin- Selaginella. - Pollen et Spores, 31: 251-288. thia II, S. 467-468. TAYLOR, T.N. and TAYLOR, E. J. 1993. The Biology and WARCH, A. 1979. Perm und Trias der nördlichen Gailt- Evolution of plants. Prentice-Hall Inc., Englewood Cliffs, aler Alpen. - Carinthia II, Sh. 35, 111 S., Klagenfurt New Jersey (USA), 982 pp. WARD, L.F. 1900. Status of the Mesozoic Floras of the TAYLOR W.C. and HICKEY R.J. 1992. Habitat, evolution, United States. First paper: The older Mesozoic. 211- and speciation in Isoetites. Annals of the Missouri Bo- 748. In 20th Annual Report 1898-1899, U. S. Geologi- tanical Garden, 79, 613-622. cal Survey, II General geology and paleontology, Wash- TAYLOR, T.N., TAYLOR, E.L., KRINGS M. 2009. Paleobot- ington. any. The Biology and Evolution of Fossil Plants. Burling- WILLKOMM, H.M. 1854 Anleitung zum Studien der Wis- ton MA, London, San Diego CA, New York NY, Elsevier/ senschaftlichen Botanik, 2. Specielle Botanik. Fleischer, Academic Press Inc., xxi + 1230 pp. Leipzig, 163 pp. TINTORI A., POSENATO R., KUSTATSCHER E., Michael WHITE, M. E. 1981. Cyclomeia undulata (Burges) gen WACHTLER M. 2001. New Triassic fish faunas from et comb. nov., a Lycopod of the early Triassic strata of paralic environments in the Alps. 3. International Meet- New South Wales. Records of the Australian Museum, ing on Mesozoic fishes, Serpiano (CH) 33 (16), 723-734. THOMAS, B. A. 1992. Paleozoic herbaceous lycopsids WOOD, G. R. and BEESTON, J. W. 1986. A late Permian and the beginnings of extant Lycopodium sens. lat. and lycopod cone Skilliostrobus sp. cf. S. australis Ash 1979 Selaginella sens. lat. Annals of the Missouri Botanical from Queensland. Geological Survey of Queensland, Garden, 79, 624-631. Publications, 387, 41-49. TRYON, R. M. and TRYON, A. F. 1982. Ferns and allied ZEILLER, R. 1906. Bassin houiller et Permien de Blanzy plants. Springer-Verlag, New York, USA, 875 pp. et du Creusot. II. Flore fossile. Études Gîtes Minéraux VAN KONIJNENBURG-VAN CITTERT, J. H. A., KUS- de la France. - 265 pp., Ministère des Travaux 1979 TATSCHER, E. and WACHTLER, M. 2006. Middle Triassic from Queensland. Geological Survey of Queensland, (Anisian) ferns from the locality Kühwiesenkopf (Monte Publications, 387, 41-49. Prà della Vacca) in the Dolomites (Northern Italy). Pal- aeontology, 49 (5), 943-968. WACHTLER, M., 2010. About the origin of Cycads and some enigmatic Angiosperm-like fructifications from the Early-Middle Triassic (Anisian) Braies Dolomites (North- ern Italy), Dolomythos, 1: 3-55, Innichen. WACHTLER, M., KUSTATSCHER E., VAN KONIJNEN- BURG-VAN CITTERT J.H.A. 2010. Lycophytes from the

Wachtler, M.: Lycophyta 49 Dolomythos, 2011 50