PERMIAN - Birth of a New World

PERMIAN Birth of a New World

Thomas Perner - Michael Wachtler PERMIAN_content_pages.indd 1

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Permian Birth of a New World

First Edition January 2015 All rights reserved Thomas Perner - Michael Wachtler Published by DoloMythos (Italy) and the Oregon Institut of Geological Research (USA) Printed in Italy by Athesia, Bozen ISBN 978-88-908815-3-4 F端r den Inhalt E-Mail: michael@wachtler.com E-Mail: fossilperner@arcor.de Copyrights In illustrations: Michael Wachtler unless otherwise credited In photographs: Michael Wachtler unless otherwise credited In text: Michael Wachtler unless otherwise credited

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Permian: Birth of a New World

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CONTENTS:

The End of an Old World..............................................................................Page 8 In the transition between the Carboniferous and Permian periods three hundred million years ago, the world fell apart at the seams as it had never done before. Powerful volcanic eruptions shook almost the entire northern hemisphere. The landscape changed completely.

The Complete Transformation of the Plant Kingdom.............Page 25 In the Permian period, there were profound changes in the flora. The large scale trees and giant horsetail died out, but this opened space for the gymnosperm conifers and cycads.

They characterised the flora on every continent after that.

The Life of Small Animals and Insects.......................................... Page 129 At the beginning of the Permian, many insects and small animals were already completely evolved. Some probably took up the function of pollen carrier for plants; a completely new challenge in the mutual interest of both parties.

Fish, a Model for Success..................................................................... Page 131 Strange fish evolved in rapid succession. There were lobe-finned fish living in fresh water and Xenacanthida that were the terror of the animal world in the Permian swamps. One group began a series of triumphs that continues today: the bony fish.

The Triumph of the Terrestrial Vertebrates............................... Page 161 More and more animals ventured onto land. At first only primitive amphibians sporadically left the water, but reptile-like creatures would soon conquer the continents and populate the land forever.

Saurians Conquer the Continents.................................................. Page 179 All animals became more and more complex and specialised.

Many completely different creatures soon roamed the Earth.

Two groups would become part of our collective memory: the mammals and the dinosaurs.

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The following collectors and collections kindly provided material: Private collections Barthel, Manfred, Berlin: 47 (2) Brandt, Silvio, Halle: 79 (2), 103 (4), 116, 127 (2), 128, 148, 149, 152, 153, 155, 156, 158, 159, 195 Dammann, Martin, Berlin: 30, 83, 173 (2), 178 Hesse, Norbert, Eisleben: 65, 79, 125, 148 (2), 149, 152, 153, 154, 155, 156, 158 (3), 159 (collection Thomas Schneider) Kahl, Matthias, Gera: 103 Kindlimann, René, Aathal, Swiss: 137 (3), 138 (2) Klopschar, Michael,Braunschweig: 148, 149, 160 Loth, Günther, Bovenden: 54, 132, 141, 146, 152, 159, 170 Noll, Robert, Tiefenthal: 29 (3), 32, 33 (2), 36, 45, 59 (4), 115 (6) Meyer, Jürgen, Zwickau: 8, 36 (2), 48 Perner, Thomas, Bad Homburg: 6, 27, 28, 34 (3), 45, 47 (2), 50 (2), 56, (3), 59 (3), 61 (4), 63 (2), 92 (4), 93 (4), 98, 100 (6), 104, 110 (6), 124, 133 (2), 164 (2), 168, 169, 173, 178 Schneider, Thomas, Helbra: 193 Valentini, Ferruccio, Tuenno, Italy: 29, 37 (3), 51 (2), 64 (4), 68, 69, 70, 71 (2), 76 (2), 77 (5), 101 (3), 105 (5), 120 (3), 121 (4), 124 (3), 190 Wachtler, Michael, Innichen, Italy: 38 (3), 40 (3), 51, 52, 65 (2), 73 (4), 85 (5),81 (4), 83 (5), 101 (4), 106 (3), 107 (5), 111 (6), 112 (3), 113 (8), 118, 119, (4), 120 (3), 189 Public Museums Bergakademie Freiberg: 174 Bundesanstalt für Geowissenschaften und Rohstoffe, Berlin, 151 Geologisch-Paläontologisches Institut Halle/Saale (GP Halle/Saale): 79 (2), 82, 102, 127, 150,(2), 193, 196 (2) Geopark Thüringen, Tambach-Dietharz: 180, 182, 183, 184 Kew Garden, London: 9 Museum für Naturkunde, Berlin (MUSNAT, Berlin): 87 (3), 88 (2), 89, 98, 125 (2), 127, 128, 144

Museum für Naturkunde Chemnitz (NM Chemnitz): 29, 31, 32 (2), 44 (2), 45, 69, 125, 185 Museum für Naturkunde, Gera: 52 (2), 65 (2), 102,(2), 103 (3), 128 Museum für Naturkunde, Gotha: 156, 180, 182, 183, 184 Museum für Naturkunde, Mainz: 134, 139, 145, 187 Museum für Naturkunde, Stuttgart: 171 Museum für Naturkunde, Tübingen: 176 Museum/Naturalienkabinett Waldenburg: 153 Naturhistorisches Museum Schloss Bertholdsburg Schleusingen (NM Schleusingen): 10, 45, 47 (2), 53 (2), 69 (2), 83, 90 (5), 91 (6), 128, 136, 137, 138, 154, 160, 166 (2), 168 (2), 177 (3), 180, 182, 183, 184, 189 (2), 191 Paläontologisches Museum Nierstein (PM Nierstein): 28, 29, 32, 33, 35 (2), 43 (2), 45, 49 (4), 54 (3), 55 (2), 58, 62 (2), 63 (2), 96 (3), 97 (4), 104 (4), 114, 124, 129 (4), 130 (2), 133, 135, 137, 142 (2), 143, 165, 169, 170 (2), 172, 175, 188, 190, 191, 197 Pfalzmuseum für Naturkunde – POLLICHIA-Museum, Bad Dürkheim (PMN Bad Dürkheim): 57, 137 (2), 139, 144, 162, 166, 167, 170, 172, 175, (2) Staatliches Museum für Naturkunde Karlsruhe (Wolfgang Munk): 156, 193 Westfälisches Museum für Naturkunde, Münster: 78, 195 Naturhistorisches Museum Wien: 9 (2), 155, 177, 181 (2), 186, 195 Národní Museum, Naturhistorisches Museum, Prag: 9, 33, 58 (2), 175 181 (2) Università e Museo Padova: 187 Università La Sapienza, Rome: 198, 199 (2) Museo Civico di Storia Naturale Brescia: 190 Museo di Scienze Naturali, Udine: 11 (4) Museo di Storia Naturale di Verona: 143 Museum Radein, Südtirol: 199 (2) Sauriermuseum, Aathal, Switzerland: 172 Museum für Naturkunde, Warschau, 196 Museum für Naturkunde Kotelnitsch, Russia, 196

Explanations: A page and the number of images in parentheses. The name or the institution in which it is kept were placed in parentheses like a seal in the book in some cases (e.g. (MoP Nierstein) = Museum of Paleontology Nierstein, or (NM Schleusingen) = Natural History Museum in Bertholdsburg Castle, Schleusingen).

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Permian: Birth of a New World

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Chronological divisions in the Permian and important geological formations System Age Name Germany Alps Eatern Europe

242 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Anisian

Untere

247.2 - - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Olenekian 251.2 Induan - - - - - Buntsandstein - - - -- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 252.17 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Changhsingian 254.14 - - - - - - - - - -Zechstein - - - - - - - - - - - - - - - - - - - - - - - Seceda, - - - - - -Bletterbach - - - - - - - -Bohuslavice - - - - - - - - (CZ) --Eisleben Recoaro Kotelnitsch (RU) Wuchiapiangian

259.8 +-0.4 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -- - Capitanian

Lopingian

Perm ian

Muschelkalk

Guadalupian

265.1 +-0.4 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -- -

Wordian 268.8 +-0.5 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Roadian

272.3 +-0.5 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -- - -

Kungurian

Tambach Cisuralian

Oberes Rotliegend

283.5 +-0.6 - - - - - - - -- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -

Artinskian

290.1 +-0.3 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -

Sakmarian

Pennsylvanian

Carboniferous

Tregiovo-FM

Collio-FM Oberhof-FM Goldlauter Unteres Rotliegend Manebach - - - - - - - - - - - - - - Ilmenau --------

Lodeve

Boscovice Broumov FM

295.0 +-0.2 - - - - - - ---------------------------Asselian 298.9 +-0.2- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Meisenheim-FM

Gzhelian 303.7 +-0.2 - - - - - - - - Stephanian - - - - - - - -BC - - - - - Niederhausen -----------------------------------Kasimovian

307.0 +-0.2 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -

Moscovian

315.2 +-0.2 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -

Bashkirian

323.2 +-0.4 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -

Serpukhovian

330.9 +-0.3 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -

237 - - - - - - - - - - - - -Keuper ---------------------------------------------Ladinian

Mittlere

Triassic

235___ _

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240___ _ _

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_ 245___ _ _ _

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_

250 _

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_

_

_ -255___ _ _

_ _ 260___ _ _

_ _ _ 265___ _ _

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_ 270___ _ _ _

_ _ 275___ _ _

_ _ _ 280___ _ _ _

_ _ 285___ _

_ _ _ _ 290__ _ _ _

_ _ 295__ _ _

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_ 300___ _ _

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_ 305___ _

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_ _ _ 310___ _ _ _ _ _ 315__ _ _ _

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_ _ _ _ 330___ _

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_ 340___ _ _ _ _ _ 345___ _

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Westphalian

Namurian

MIssissippian

Visèan

346.7 +-0.4 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -- - -

Tournaisian

Visèan

Tournaisian

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An amphibian sample from the large order Temnospondyli with particularly well preserved skin, where it is even possible to see the fine scales. Long, oval eye sockets and the sclerotic rings (the ring-shaped, bony reinforcement around the eyes) are visible.

This uniquely mutated form can belong to a new species.

We can expect even bigger surprises in the evolution of life, especially from the Permian period. Niederhausen, Perner collection.

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Permian: Birth of a New World

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Why a book about the Permian The period 300 to 250 million years ago is considered one of the most misunderstood periods of our planet, as far as geology and the development of life are concerned, and also one of the most mysterious. There is not yet enough evidence, but it has become more and more clear that the key to understanding all aspects of our current existence can be found in this period.

Its importance lies in the fact that the blueprint for conquering the continents, and the permanent colonisation of land was developed during the Permian. Amphibians and other predecessors of various animals diversified into reptiles, dinosaurs, and into the ancestors of all mammals alive today.

The evolution of flora at this time was equally dramatic. The era of the giant clubmosses and giant horsetail was over forever, and a new group of plants, which still dominates many parts of the world today, took front stage, the gymnosperms, and with them, the conifers and cycads. The most dominant plant group today, the flowering plants, must also have split off from these early plants.

As authors, this project became a search for clues, at the end of which we found ourselves in a period from the distant past. Our journey through time took us all over Europe - and the whole world. We dug through the evidence that scientists had collected over the course of centuries, and which is now exhibited in museums as the masterpieces of creation. We accompanied researchers on their searches for related artefacts. Some of these people were simple shepherds and herbalists, while others were academically trained in other fields but dedicated themselves to researching our existence in their spare time. Others were experts who had focused on their niche areas of the Earth‘s history since childhood.

We crossed established political borders, and the researchers sent by the governments opened up to us, as did the numerous unknown and otherwise forgotten private scholars working completely on their own. To our amazement, we found that in this world of explorers and discoverers, no one was superior or inferior to any other, and that na-

tional borders had been arbitrarily designated by humans and had never existed in nature. Something beyond human influence reigns over everything: „time“.

In the Permian, our planet experienced cycles of extreme climatic change, and these periods provided the most fertile breeding grounds for diversification. Dry lagoons forced the fauna that lived in water to emerge onto land, at least to find a new source of water. This may then have forced them to adapt to a life on dry land, in what was probably the birthplace of all current land animals, ultimately including humans.

Desert-like or icy cold continents offered „dormitories“ lending a marginal existence and thus new opportunities for development over millions of years. This is the only explanation for the sudden rise of conifers. Plants and animals mutated in the Permian, within a short period of time, to become new families, which have surprisingly survived to today, almost completely unchanged. The cycads are one example, another is the amphibians. Many theories expounding slow evolutionary processes falter in the face of such an explosion of life.

At the end of the project, we authors found ourselves in a completely new world. Large parts of the world are still „Permian“.

Using our understanding of today‘s biosphere, we are able to shed light on the biosphere that existed several hundred million years ago. We can then examine the relics of that time to understand the diverse biosphere of today.

Michael Wachtler and Thomas Perner January, 2015

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The End of an Old World Between the Silurian and Carboniferous periods, all living creatures experienced profound changes.

First, higher forms of plants conquered the land. In just over one hundred million years, formerly small flora that had only lived near the shore spread all over the world and reached heights and sizes that were never seen before, or seen again thereafter. In the same way, animals living in water conquered land and began a series of triumphs on this planet that was thought to be impossible.

The distribution of oceans and continents in what we today call Europe changed between the Devonian and Carboniferous periods. The north was surrounded by the Devonian Old Red Continent, and part of the Caledonian mountains. Central Europe was dominated for the most part by tropical seas, with elongated islands wedged between land masses, and the Alps were surrounded all the way to Spain by a markedly changing Tethys Ocean.

The landscape began to change to an alarming degree with the beginning of the Lower Carboniferous period. Due to their movement and the increasing proximity of all continental blocks, entire chains of mountains soon began to form.

The climate experienced profound change and volcanic activity started becoming more and more intense. Two mighty mountain ranges were born in Europe, the Armorican Massif, which extended from central Spain to southwest England and to the Mediterranean coast in France, and the approximately 500 kilometre long Variscan Massif, which ran straight through central Europe. Most

Europe of today in the Upper Carboniferous

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of today‘s Europe was located near the equator.

The warm and humid environmental conditions favoured the development of extensive primeval forests, comprising mainly giant clubmoss and giant horsetails, while new forms of life such as the largest insects in the history of the Earth took over the landscape. Over almost all Europe, in areas subject to frequent flooding by seawater and in the interior, massive layers of coal were formed, composed of the residue resulting from the enormous amounts of dead land plants.

Leaf cushions of Sigillaria. Together with the genus Lepidodendron, they combined to form enormous clubmoss forests in the Upper Carboniferous period. JĂźrgen Meyer collection, Zwickau

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Top: In the year 1820, Kaspar Graf Sternberg, the founder of modern palaeobotany, described the first Lepidodendron trunk (NĂĄrodnĂ­ Museum, Prague, Czech Republic). Right: Detail of the surface of Lepidodendron: spirally arranged rhombic leaf cushions cover the surface of the trunk like scales. Lower right: Part of a clubmoss trunk with a diameter of one meter: both are from Rosnitz, Czech Republic (Natural History Museum Vienna). Bottom: Reconstruction of a forest from the Upper Carboniferous period. Kew Garden, London.

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The Reign of the Scale Trees and Giant Horsetail Lycopods such as the Lepidodendron in particular, with its crown of bifurcating branches, reached enormous heights of up to 40 meters. The typical pattern on the surface of the bark, caused by dropped leaves, partially characterised the appearance of the forests of the Carboniferous period, as did the Sigillaria trees, which only grew half as high. The landscape also included

arborescent Calamites, which were only held in place by a hollow trunk and could reach heights of up to 20 meters and diameters of up to half a meter. Two groups of ferns dominated the continents, ancient low-growing ferns or tree ferns (Filicales) which are similar to today‘s, and, curiously enough, seed ferns (Pteridospermae), which were characterised by seeds and pollen organs. The abundance of biomass led to the creation of huge layers and seams of material that did not rot and, due to geological folding and continental drift, are useful for humans today in the form of hard coal, which we eagerly mine.

Ancient Dragonflies and Giant Millipedes Large winged insects, similar to the dragonflies of today, with proboscises and stiff wings, took over another niche, the air. These insects included Meganeura, the largest flying insect of all time with a wingspan of almost one meter.

Arthropods up to two meters long, but also smaller spiders, scorpions, and millipedes, colonised the underbrush of the primeval forests.

Palaeodictyoptera, with their stiff, non-folding wings, looking like modern dragonflies but much larger, began to spread. Amazingly, there were even flying insects with three pairs of wings. In contrast, muddy swamp bottoms offered an ideal habitat for a myriad of grasshoppers, roaches, and beetles, which significantly outnumbered all other animals living on land.

Armoured Fish and Labyrinthodonts

In many animals, the initial body plan was so good that they have remained almost unchanged into modern time. Top: Euproops bifidus, a primitive original horseshoe crab from the Upper Carboniferous from Piesberg near OsnabrĂźck (G. Sommer, Natural History Museum in Bertholdsburg Castle, Schleusingen). Bottom: Its descendant Limulus polyphemus, found especially in the flat sandy coasts of tropical seas.

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The open waters were colonised by ganoid fish such as the Palaeonisciformes, spiny sharks, and fish from the group including acanthodes, megalodons, and coelacanths. These are all fish families that would become important later in the Permian period. Amphibians such as the temnospondyls (Stegocephalia) began their first golden era in the Upper Carboniferous period. Four to five meter long crocodile-like creatures such as Anthracosaurus or Pteroplax, which was better adapted to life in water, reigned over prehistoric Europe and would become subsequently more diversified.

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Annularia stellata,, parts of a giant horsetail plant from the Carboniferous. Ferns such as Pecopteris or Alethopteris spread out over a large area. Many were even ferns that bore seeds. All specimens are from the top Carboniferous plant fossil site of the Alps, Krone/Pramollo, collection of the Museo di Scienze Naturali, Udine.

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The Lower Permian - a Changing Landscape During the Carboniferous, and even more so at the beginning of the Permian period, almost every piece of continental land came together to form a contiguous land mass spanning prehistoric Europe to North America, with serious consequences. Land bridges to Australia, Africa, and South America allowed the exchange of plants and animals almost everywhere in the world. The birthplace of our habitat today can be found in this unique cluster of continents.

Many regions in prehistoric Europe rose so high that they could hardly be covered by new sediments. Other lower regions, however, gained alluvial material due to rivers and floods, and were thus raised. Triggered by dramatic changes in the Earth‘s crust, intensive volcanic activity lasting millions of years began throughout Europe, from Scandinavia to the Mediterranean region. Accompanied by uplifts and subsidence, breaks, the formation of volcanoes and the associated lava flows, the result was continuous change and a radical transformation of the former landscapes. The legacy was the striking terrain still present today, including features such as the porphyry plate of Bozen in the Southern Alps, the Donnersberg (the highest mountain in the Palatinate, or the petrified forests in Chemnitz, which were preserved due to volcanic eruptions.

Partly due to drifting into the slightly more distant equatorial regions and due to partial isolation from clouds that provide rain, annual

cycles began to set in, in contrast to the wet and rainy Carboniferous period. The weather alternated between heavy monsoon rains in connection with the depositing of large quantities of red muddy sediments followed by longer dry periods. This produced the sequence of rock strata in central Europe referred to as the “Rotliegend”, which is divided into larger and smaller formations and into the older “Lower Rotliegend Group” and the “Upper Rotliegend”, the latter of which was in formation almost until the Middle Permian period. The name, however, is derived from the term used by miners to refer to “dead” layers of rock strata containing no ore.

In other parts of Europe such as the Southern Alps, the erosion of the thick lava flows from the Lower Permian also created red layers of rock which are referred to in geology as “Grödner Sandstone”. This sedimentation was especially significant in the Upper Permian, although the first deposits were produced in the Lower Permian.

The landscape today in the Rhineland-Palatinate: due to the erosion of flattened areas surrounding Bad Dürkheim, there are hints of previous life forms hidden there.

The quarries on the Bromacker on the northern edge of the small city of Tambach-Dietharz in Thuringia, the most important site for the fossils of terrestrial vertebrates (Tetrapoda) from the Lower Permian in Europe.

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Important sites for geological research of the Lower Permian

Sheets of lava in the Donnersberg Formation near Niederhausen, and in the foreground, light sandstone formations formed due to sediments from the sea. They are rich in primitive amphibians, fish, and very well preserved plants.

The area around the Collio in the Brescia Alps is rich in plant fossils and animal tracks. The Lower Rotliegend layers awakened the interest of scientists as early as the 19th century.

A Rich Marine Environment and Walking on Land The fact that the climate of the Lower Permian was not as dry as generally assumed is demonstrated by numerous small lake areas and large wetlands such as the Rßmmelbach-Humberg Sea between Lebach in Saarland, and Bad Kreuznach in Rhineland, which at 3,500 square kilometres could still be one of the largest lakes in today’s Europe.

The Petrified Forest of Chemnitz. An open air museum was built around the early Permian tree trunks found there. Although numerous petrified plants were found, hardly any animal skeletons were discovered.

Plants and animal tracks from the Tregiovo in the Trentino Dolomites were analysed for the first time in the 19th century. Many scientific papers on this area followed.

Fossils found there prove that there was an exceedingly abundant population of freshwater fish consisting of ganoid fish such as Paramblypterus, spinyfinned fish such as Acanthodes, large sharks such as the Xenacanthida, and even lungfish (Conchopoma) or freshwater coelacanths, which were able to spread out in selected habitats. Large temnospondyl amphibians (Sclerocephalus, Cheliderpeton), but also archegosaurians and small branchiosaurians, formed an additional, rich population, which triggered additional evolutionary steps towards colonising land.

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The Landscape in the Upper Permian At the beginning of the Upper Permian, even more dramatic changes took place. The territories had all finally merged to form a single supercontinent - Pangea - and a single ocean - Panthalassa - united all bodies of water.

North America and Europe of today were located almost on the equator. The southern hemisphere, together with large parts of Africa, India, South America, and Australia, was covered by enormous sheets of ice that regularly pushed forward, only to partially melt again. In the area between the Southern Alps and the Black Sea of today - the Tethys Ocean was able to break through the continent in a wide and narrow inlet, thus forming the basis for the break-up of the continents which began in the Triassic period.

A Decline in Volcanic Activity The heavy volcanic activity that characterised the Lower Permian subsequently subsided, while the erosion of the Variscan mountains was in full swing. Sands and boulders, which were red for the most part, completely filled the smaller basins. Towards the beginning of the Upper Permian, large parts of Europe sunk as a result of local movements of the Earth‘s crust, and formed expansive plains such as the Germanic Basin, while the area around the Alps began to push the coastlines due to the similar subsidence forces in parts of the prehistoric Tethys Ocean.

The European landscape during the time of the Zechstein (Wuchiapingian - Changhsingian 259.9 - 252.2 million years ago)

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About 258 million years ago (Lopingian) a rift was formed due to tectonic activity in the north between Greenland and Norway, which were once located directly next to each other, through which the ocean was able to penetrate and subsequently flood large parts of Central Europe for a short time. The birth of the Zechstein Sea had begun.

Due to the inflow of floodwater and the enormous amount of water, the sea level rose rapidly and covered areas that were previously land. At first only a few clay sediments were deposited there: these layers of sediment, which are referred to as copper slate due to their high ore content, triggered the formation of this former European inland sea, the beginning of which remained preserved until today. This sea would soon extend to cover almost one million square kilometres, which corresponds to about twice the surface area of the Black Sea. Its watershed extended from modern Greenland to England, then to the Baltic region, parts of Poland, and South-Western Germany.

Once the rate at which the sea was rising began to subside, the deposition of sediments raised. This led to the accumulation of digested sludge on the

The remains of animals can be preserved as evidence of life in earlier times for millions of years. However, good conditions for preservation are necessary, such as those of the bird preserved in the salts of the Ethiopian Danakil Depression.

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Current landscapes that match the conditions during the creation of the Zechstein Sea in Upper Permian After the Panthalassa Ocean penetrated into Central Europe through a rift and caused flooding, other areas would repeatedly dry out, as illustrated above by the rift in the Red Sea in the direction of the Ethiopian Danakil Depression (above) or the Great Salt Lake in North America (middle right). The sea level then increased dramatically, providing normal conditions, so that vegetation consisting of Araucaria-like conifers was able to colonise land right up to the shores (New Caledony).

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floor of the sea, and the salt content of the sea increased so much that it was similar to levels in the Great Salt Lake in America or the Dead Sea of today. The reason for this is found in the high rate of evaporation, which in turn was due to the warm climate and the small inlet to the sea. At times, the Zechstein Sea was completely isolated from the rest of the ocean, which led to the higher concentrations of salt and the formation of thick layers of salt.

group of Peltaspermales that adapted to the dry conditions, and early ancestors of the cycads.

In higher layers we do not find the same conditions for the preservation of fossils, and the fossil density decreases in these layers. Very few invertebrates, plants, and fish remains have been found in the black shale created from digested sludge. In the subsequent chalky and dolomitic layers, only the remains of invertebrates can be found.

The Birth of the Inland Zechstein Sea

The Reign of the Tethys Sea

During the recurring periods of increased rainfall, the overall salt content dropped so much that it was even possible for reefs to form that allowed the creation of environmental conditions which must have been similar to the conditions present today in tropical seas. The reduction in salt content may have been caused by a rise in overall sea levels due to the decline of the ice ages in the southern hemisphere. The repeated alternation between increasing and decreasing rates of salt and gypsum deposits, and adaptation to current conditions in the ocean is characterised today by the sequence of the layers deposited. This created the thickest salt deposits in Europe, and the large number of fish communities, terrestrial vertebrates and plants indicate a rich living environment. Although the outcrops of rock in the various areas of the former Zechstein Sea exhibit regional variations in terms of the distribution of fossils, it is still possible to speak of a typical Zechstein living environment towards the end of the Permian period, especially because of the primitive ray-finned fish Palaeoniscum which was found everywhere and which dominated the seas. Other fauna also belonging to this living environment, and of particular interest for the evolution of life on land includes the Pareiasaurus or the primitive Diapsid Protorosaurus and the earliest flying lizard Coelurosauravus. In contrast, the flora was dominated by several conifers, such as Pseudovoltzia or Ullmannia. There were also Sphenopteris ferns, seed ferns from the

Even in the modern Southern Alps region in northern Italy, or in France, where there were still extensive land masses during the Lower Permian, the sea began to take over more and more land towards the beginning of the Upper Permian, due to subsidence of the continent. In these zones, a plethora of animal tracks were preserved. The Bletterbach Gorge in the Dolomites is especially well known for such tracks. The large number of plants preserved provides an excellent glimpse of what the living environment was like 260 million years ago. Pachypes-traces assumed to be caused by pareiasaurians were primitive plant eaters about the size of a cow, with thick legs and a short tail. There were also early Lepidosauria and Therapsids, which are believed to be the ancestors of mammals.

There are indications of a flora rich in tropical/ subtropical plants such as the Cycadophyta, horsetails, ferns and seed ferns, and conifers, but the diversity of the species was low. Probably only a few species were able to colonise large parts of Europe, similar to the situation today on the extensive plains in the northern hemisphere. In the Southern Alps, the characteristic conifers Ortiseia and Majonica, a conifer bearing winged seeds, dominated the landscape.

Due to the large number of hygrophilous plants, however, one cannot consider the climate to have been desert-like or arid. It can be assumed that the Europe of that time consisted of extensive plains and coastal rainforests that were only rarely inter-

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A Complete Change in the Plant Kingdom During the transition from the Carboniferous to the Permian, there were dramatic changes to the flora in the northern part of the Pangea supercontinent. The changes were not abrupt, so that in large areas - such as North America - they took place over a long period of time and allowed the large tropical coal swamps to continue to survive. In spite of this, in a short geological period of a few million years the vegetation changed more drastically than ever before or ever after in the history of the Earth.

In Central Europe, the extensive clubmoss and horsetail forests were pushed back, starting in Westphal B 307 million years ago, disappearing almost completely in five to seven million years.

In their place, a new plant family appeared that would come to dominate large parts of the global flora up to modern times: the gymnosperms or “naked seeds�, and in particular the conifers from this group. The first remains of conifers, in the form of small cuticles, were found at Westphal B in Yorkshire, England, but in a short time, they would attain a series of triumphs unparalleled in history.

They not only pushed back the previously completely dominant clubmosses and overshadowed them until today, but also were

able in just a few million years to reach a stage of evolutionary development so successful that they have only changed marginally to this day.

The male and female cones, the growth habit, and the foliage of Early Permian conifers barely differs from those of today: quickly conifers bearing winged and nut-like seeds developed into the form commonly encountered today in most of the pines and cypresses found in the northern hemisphere. Even the seeds embedded deep in the scale trees, like those in the Araucaria in the southern hemisphere, were created at this time. Then, as today, there were monoecious and dioecious conifers. Even the ginkgos achieved their characteristic appearance with collar-like petiole bases from which the segmented leaf

Reconstruction of the living environment in Niederhausen in the Palatinate during the transition from the Carboniferous to the Permian period. Various primitives (Perneria) and conifers that were already highly evolved (Seymourina, Otovicia, Majonica) dominated the landscape. Seed ferns (Peltaspermales), common ferns, and horsetails (Calamites and Equisetites) were also present.

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The Rise and Fall of the Ferns Although the ferns, together with giant clubmoss plants and horsetail trees, dominated the forest landscapes during the Carboniferous period, this world of ferns suddenly declined during the transition from the Carboniferous to the Permian. While some families of ferns still survived into the beginning of the Lower Permian, almost the only species found at the beginning of the Middle Permian was the small-leaved and skeleton-like Sphenopteris. The Carboniferous period was characterised by an abundance of seed-bearing ferns such as Lyginopteris or the Medullosaceae, while only one significant family of seed ferns survived the rest of the Permian in the northern hemisphere: Peltaspermales.

With about 9,000 different species, ferns play a more important role in today‘s vegetation than horsetails or clubmosses. A wide variety of different ferns exist today, from 20 meter high tree ferns to creeping, moss-like ferns.

They colonise the rainforests, can be found in mangrove swamps, expand into arid regions, and even dare to go into mountainous areas up to the snow line and deep into the Arctic. The ferns of today are difficult to characterise, but the ferns of the past are even more so. The main factor contributing to this difficulty is the fragmentary

fossil finds, since many species only developed reproductive sporangia on certain areas or limited their production to certain special fronds.

The decline of the ferns During the Carboniferous, numerous types of ferns reached their peak and exhibited a great wealth of forms. The ferns that still exist today were in a more primitive form in the Carboniferous, such as the Marattiales order which can today be found in the tropics. A fern is considered primitive if the sporangia in a

This is what tree fern forests must have looked like during the transition from the Carboniferous to the Permian. Over the course of the Early Permian, most ferns were crowded out, and their niches were occupied by the cycads and conifers.

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The Rise and Fall of the Peltaspermales It is difficult to find any other family of plants that suddenly appeared mainly out of nowhere between the Carboniferous and Permian, conquered the world, and then inexplicably declined until they disappeared astonishingly in the Late Triassic, than one of the most mysterious seed fern genera: the Peltaspermales.

It has only been possible to infer a classification or relationships to other plants based on their foliage for a few of these species, which were particularly widespread in the northern hemisphere, the shape of their leaves is just too varied. One feature that makes them unmistakable, however, is their umbrella-shaped female fructification.

Like the Glossopteris plants, which appeared at the same time and were one of the seed ferns in the flora of the continent Gondwana, they form the most typical group of plants existing during the Permian period, with Autunia conferta as a higher level symbolic plant. This very interesting group of plants reveals a special problem in 1

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palaeobotany: the term Peltaspermum pertain to the female seeds, while the foliage is often referred to using other names such as Autunia, Rachyphyllum, or Lepidopteris. Since their umbrellashaped fructifications make them so unmistakable, tey became the distinct feature used for their classification. In the form genus Peltaspermum used for female fern fructifications - which usually have round to slightly elliptical, segmented (8 to 20 times), umbrella-shaped, and 5 to 20 millimetre long pods - the seeds were embedded around the stalk, which was connected to the main stem.

The wedge-shaped incisions on the opposite side are also very easy to recognise. Since such Peltaspermum heads have been found dating from the Carboniferous-Permian transition to the Triassic period, and several species of seed ferns with completely different fronds could usually have borne these organs, it is often impossible to classify them with certainty. The only aid in this regard is their connection to the foliage, which is very rarely found.

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Wachtlerophyllum schaalii. One of the most primitive Peltaspermales. The fronds are deeply lobed. The circular ovule bearing fruits sprout out from a short peduncle and are protected by leaflets from the main rachis. 1. Mainly complete frond, 40 cm long. 2. Detail of the bifurcating leaves. 3. two umbrella like fruits holding the seeds, reconstructions, all Niederhausen (Coll. Th. Perner).

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An exceptionally pretty specimen of Autunia conferta, Rockenhausen, Palatinate (NHMoP Bad D端rkheim).

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Female Cycas revoluta plants. All cycads are characterised by the fact that male and female cones grow on separate plants.

The Birth of the Cycads The origins and evolution of the cycads or palm ferns have fascinated palaeobotany researchers for decades. On the one hand, this is probably due to their early origins - with ancestors extending back more than 300 million years into the Carboniferous - and on the other hand because the plants that developed from them or parallel to them could enable researchers to gain knowledge of the evolution of the flowering plants.

The cycads still living today consist of 11 genera and 305 species, which make them the second largest group of gymnosperms after the conifers.

They spread over the tropics and subtropics north and south of the equator, and Central America is considered the region with the greatest diversity of cycads. The most extensive range belonged to the genus Cycas with representatives that colonised the northern hemisphere from Japan to China and Southeast Asia, and in the southern hemisphere throughout the Pacific, Australia, and the eastern coast of Africa, including Madagascar.

The palm ferns are generally divided into three large families: the Cycadaceae, the Stangeriaceae,

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and the Zamiaceae. Today, the Cycadaceae family only contains one large type genus: Cycas. The Stangeriaceae are divided into two genera, the Stangeria and the Bowenia, each of which only contains one species, while the Zamiaceae split into two subfamilies: the Encephalartoidea and the Zamiodeae.

Features common to all cycads All cycads exhibit a plethora of common features that point towards a single, common ancestor.

They all have separate sexes, with male and female fructifications located on separate plants.

With the exception of the genus Cycas, all species

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developed striking male and female cones that are easy to differentiate from those of other plants and were located individually or in groups at the tip of the stem between the fronds. They could reach a significant size and consisted of a receptacle with sporophylls arranged in a spiral around it. Only in the genus Cycas did all macrosporophylls protrude at one level. For the most part, the sporophylls of the cycads bore a total of two seeds that were protected by a pair of mirror-symmetrical covers. Only in the genus Cycas can they contain up to 16 seeds.

The male pollen organs of all cycads appear to be relatively uniform. They were wide, shield-shaped segments that bore microspores on their undersides. They ripened while covered by the tightfitting microsporophylls, and they only separated to open when releasing pollen due to stretching of the internodes of the receptacle.

All cycads living today bear finely branched or unbranched hairs on the tips of their cones, referred to as trichomes. With the exception of Bowenia, which is the only bipinnate species, all cycads exhibit pinnate leaflets without an end segment. Every two to three years, a rosette of

new fronds forms, which then unfolds like that of a fern. Most species also form a more or less long, spherical to beet-shaped tuber. Some species can also appear column-like or tree-like, but are always without branches and seldom exceed a height of two meters. All modern palm ferns produce the glycoside cycasin, which is a property unknown to other gymnosperms.

All cycads are equipped with two types of root organs: a primary, thick, fleshy, and elongated taproot from which the coralloid, lateral, highly branched roots extend. They live in symbiosis with algae that are able to filter nitrogen from the atmosphere. This unusual feature is unique in the plant kingdom.

The first leaf fragments that can be considered ancestors of the cycads appeared in the Carboniferous between the Mississippian and Pennsylvanian periods. Early descriptions from the 19th century placed the line of development of the cycads in the since similarities with the inflorescences of the conifers were observed.

Another theory focuses that the cycads originated from a group of seed ferns from the Carboniferous, the Medullosaceae.

As on ferns, the young shoots of cycads unroll slowly at first (left: Cycas revoluta). Different species of cycads have different types of fronds: in the middle is Lepidozamia peroffskyana, and on the right is Stangeria eriopus.

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Bjuvia tridentina. Strangely enough, the modern Cycas frond evolved from a leaf with an entire margin. Reconstruction of the Lower Permian cycad Bjuvia tridentina. 1.

Individual frond, and 2. complete plant. 3. Young frond, Tregiovo, Northern Italy (Coll. F. Valentini).

The relatives of the cycads Although fossil finds predominately come from America, Europe and China, in the northern hemisphere, they must have spread all over the world very early. Some interesting conclusions can be drawn: the process of formation of the modern cycads - in terms of their structure, the tubers, and the male and female seed parts - was already more or less complete in the Permian period. At the end, however, only marginal fundamental evolutionary processes can be detected over the following millions of years up until today. Even the frequently cited Permian-Triassic extinction event apparently swept over the plant kingdom without leaving any traces. All Cycadales of the Permian survived in a barely altered form into the Triassic. For a long time, doctrine has held that the genus Cycas can be considered the most primitive based on the unusual structure of their macrosporophyll, and that all others were derived from it through reduction. Fossil finds, however,

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indicate that the large Cycas and Zamia groups developed simultaneously, ever since the Palaeozoic era. For this reason, they must already have been separated between the Carboniferous and Permian, which makes it impossible to consider them more primitive and less developed cycads.

The development of the cycads between the Carboniferous and Permian In many parts of the world first taeniopterid leaves, thought to belong to primitive cycads, appear in the Carboniferous. They were classified as Taeniopteris polymorpha, T. multinervis, or T. coriacea. Even though an unambiguous classification as Cycadales has never been demonstrated by analyses of the cuticles or on the structure of the fertile organs, they can be considered as possible ancestors. This includes Phasmatocycas of the American Midwest from

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Lower Permian cycads 1 3

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1. Bjuvia tridentina, Lower Permian/Artinskian, Tregiovo, (Coll. F. Valentini, MUSE Trento. 8cm), 2. (Nilssonia) abnormis, Middle Rotliegend, Chemnitz Hilbersdorf, (Otto Weber estate, Museum of Natural History Chemnitz), 15cm. 3. Taeniopteris (Bjuvia) multinervis frond. 4. Detailed view of the venation. “Oberhofer Schichten�, Upper Rotliegend, Oberhof (NM Schleusingen).

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the Upper Carboniferous (Pennsylvanian) and Lower Permian layers (Artinskian-Kungurian).

The plants hold relatively long and slender fertile leaves joined on a central axis that surrounded a large number of seeds in two rows along a middle vein and which released them upon maturation.

This scenario is slightly different to that of Primocycas chinensis from the middle of the Lower Permian. Compared to Phasmatocycas, its sporophylls were smaller and only bore between one and four seeds of varying size on each side. The apical pinnule was well developed. Relatively well preserved Cycadophyte fronds came from the Early Permian series in South Korea. Pseudoctenis samchokense bore surprisingly well pinnated fronds, in contrast to the many taeniopterid leaves known from this period. It can therefore be assumed that some leaf shapes reached the typical pinnate structure of the cycads of today between the Carboniferous and Permian periods.

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Differences in the shapes of the cones and fronds Strangely enough, some ancestors of the cycads hold more or less large entire fronds that may have been torn several times due to the rigors of nature, like todays banana leaves. They were classified as Bjuvia, Ladinia, Phasmatocycas, or in the form genus Taeniopteris when there was no additional evidence for classification available, and they lived from the Carboniferous into the Cretaceous period. These species bore fertile leaves similar to those of the modern cycads where from just a few to numerous seeds grew closely together in two parallel rows. A feather-like and sterile end gave these seed-leaves an unmistakable appearance. These appendices, which have often been isolated finds, were therefore classified with their own name, Dioonitocarpidium, but also as Cycadospadix, Pseudoptilophyllum, and Palaeocycas, and were recognised from the 19th century as possible ancestors of the pistils of the modern genus Cycas.

Another group appearing in the Early Permian were the cycads with segmented fronds, whose leaves were characterised by fronds with a more or less geometric growth habit. These cycads were classified as Nilssonia, Pseudoctenis, or Apoldia.

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1. Thetydostrobus sp. Pollen cone of a Lower Permian cycad (Bjuvia, Nilssonia, 10cm), Tregiovo (Valentini collection). 2: Pollen cone of Stangeria eriopus, and a cross-section of the top and bottom of the cone.

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Lower Permian cycads: As early as the Early Permian, clearly structured and segmented fronds appear that were classified as Nilssonia or Pseudoctenis. They formed both male and female cones very early, which indicate the modern Zamiaceae.

A frond of Nilssonia perneri connected to the attached tuber so typical for cycads, and a reconstruction of the plant, Tregiovo, Northern Italy (Valentini/Wachtler collection, MUSE, Trento, 22cm).

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Fossilised male cycad cones were defined as Androstrobus, female cones pointing in the direction of the modern Zamiaceae as Tethydostrobus.

Much is still unclear about the development of the cycads between the Permian and Triassic, however, especially in terms of the decisive feature, the fructification.

The unique features of the cycads in the Lower Permian Due to the high number of fossil sites and the long period of time, which makes classification difficult, and which extends from the Upper Carboniferous period (Stephan C) to the Lower Middle Permian, precise chronological determination of when the first possible ancestors of the cycads appeared is not possible.

Female cone (top) and male cone (bottom) of Encephalartos ferox. They belong to the group of Zamiaceae, which bear relatively similar male and female cones. The female cones are generally somewhat more bulbous, and the male cones are thinner and longer. They are probably successors of the Permian Nilssonia cycads.

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Reconstruction of the cycad Nilssonia brandtii. A single frond, and the entire plant with two female cones.

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Upper Permian cycads Nilssonia brandtii

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Nilssonia brandtii. 1. Female cycad cone, 12cm. 2. Detailed view of the sporophylls. 3. Part of a frond. 4. + 5. Male cone. Like the cones of modern cycads, the male cone was more slender than the female cone. Ariche, Valli del Pasubio, Northern Italy (Coll. M. Wachtler)

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A comparison of the different female organs: megasporophylls of Cycas revoluta with seeds and of Zamia floridana.

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1 Pollen organs of Cycas revoluta and of Stangeria eriopus. The male cones of the cycads barely differ between the various genera.

In many cases, their leaves have an entire margin that was classified as Taeniopteris abnormis or Taeniopteris multinervis. For certain larger leafs, however, some of which were over 20cm long and 10cm wide and exhibited dense, parallel lateral veins, classification in the genus Bjuvia would be more justifiable.

In addition to central European flora, an abundance of cycadophytic plant material has also been found in the Southern Alps. With its irregular but still recognisably segmented fronds, Nilssonia perneri (Artinskian-Kungurian) can be

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Reconstruction of the Upper Permian cycad Bjuvia wachtleri. 1.

Complete plant. 2. Leaf. 3. Male cone.

considered a true ancestor of the cycads. The fact that the typical features of cycads developed very early can be seen in the tubers found, together with the rest of the plant. From the Permian and through the Triassic, tuberous roots with coralloid secondary roots are considered typical of cycads. The elongated roots of certain modern cycads evolved from them.

Other cycads with fronds bearing entire margins were classified as Bjuvia tridentina. In contrast to descending species appearing between Upper Permian and Triassic in the same area, how-

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Upper Permian cycads Bjuvia wachtleri 2

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Bjuvia wachtleri. 1. Part of a large frond. 2. Tip of a leaf. 3. Detailed view of a leaf.

4. Reconstruction. The leaf veins fork at the base. 5. After forking at the base, they continue in parallel and do not fork any more. 6. Part of a disintegrated male cone. Ariche, Valli del Pasubio (Coll. M. Wachtler).

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ever, they were smaller, had a maximum height of 30cm, and a width of 15-20cm, while Bjuvia fronds especially in the Triassic could reach lengths of up to one meter in the Triassic period.

They are the ancestors of the modern Cycas species.

The cycads in the Upper Permian There were no major changes in the development of the cycads up to the Upper Permian. After their sudden appearance between the Carboniferous and Permian, they became more numerous, especially in landscapes located near the equator or in the Southern Alps of today, but the shape and structure of their fronds barely changed at all. The pinnate veins on the blades of the upper part of the fronds of Bjuvia wachtleri, are reminiscent of Bjuvia multinervis from the Early Permian in eastern Germany.

Nilssonia brandtii, which bore segmented fronds, differs only slightly from modern Zamia cycads with its more robust female cones and more slender male cones. All things considered, they verify the developments starting in the Per-

mian that led to their division into the Cycas and Zamia plants.

Cycad-like plants of uncertain classification Although classification as a cycad based on fossils found in context, or on development trends, is likely for many genera like Bjuvia or Nilssonia, there are still other plants from the Permian whose association with a known group cannot be properly understood.

The origins of the largest family living today the flowering plants - cannot be clearly traced, but it may be possible to find their ancestors among these plantgroups. This is especially true because the structure of the Cycadophyta offers the best opportunities to answer this question, which Charles Darwin referred to as an “abominable mystery�. How and why the ovules became covered by a covering leave, and the origin of two pairs of pollen sacs located on the sides of the stamen needs to be explained first, based on complete fossil finds. The origin of the flowering plants (or angiosperms), however, must at least lie somewhere near the Carboniferous-Permian boundary.

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Wachtleropteris valentinii. An enigmatic plant from the Southern Alps during the Lower Permian. They combine features of the cycads and conifers, but also of certain seed ferns, 15cm. 1. Sterile plant with typical forking and root system. 2. Detailed view of the pinnate leaves with stem. 3. Reconstruction.

Tregiovo (Coll. F. Valentini, MUSE Museo Scienze Naturali, Trento).

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Wachtleropteris valentinii

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Wachtleropteris valentinii. Wachtleropteris valentinii. 4. Male fructification sprouting out from a leaf, 12cm. 5. Reconstruction, 6. Adult pollen cone. 7.

Isolated microsporophyll, 8. Female cone, 9. Reconstruction, 10. Detail of a single seed scale with two seeds and reconstruction Tregiovo, (F. Valentini collection, MUSE Museo Scienze Naturali, Trento).

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Enigmatic, potential ancestors of the flowering plants The classification of Wachtleropteris valentinii, a relatively well-known low-growing plant often found in the Lower Permian layers of the Trentino Dolomites, is unclear. The leaves extended upwards on a stem and branched twice, which is not typical for cycads, and each of the leaves split into two independent branches. In this primitive form of dichotomous leaf growth, this species bridges the gap to the most primitive Devonian plants. The leaves were typically taeniopteroid/tongue-shaped and had a single wide blade from which densely packed lateral veins extended at a ninety degree angle. The tubers typical of cycads did not yet exist, and they anchored themselves loosely in the ground instead. The leaves could still be designated as “cycad-like”, however, and the arrangement of the cones on the end of a pinnate leaf identifies them as gymnosperms, but without being able to draw clear parallels to conifers, ginkgos, or cycads.

Their classification is therefore questionable, even though they formed a bridge as the “last representatives of a primitive species” in the Carboniferous.

The botanical classification of Taeniopteris jejunata, which usually originates from the Stephan and the bottom of the Lower Permian in Central Europe, is also uncertain. Their structure, which appears more like the fronds of ferns, speaks against classification as an ancestor of the cycads.

Sobernheimia jonkeri fossils have been described from layers of almost the same age (Artinskian-Kungurian) in Germany. The fructifications bore marginal seeds on a fertile receptacle. The fact that these ovules, which supposedly had Taeniopteris-like leaves, were widespread in the Lower Permian is demonstrated by similar fossil finds in North America that have been classified as Phasmatocycas. Somewhat similar but more comprehensive material, including some complete finds, is available from the Virgilian stage of the Upper Carboniferous (Gzehlian 303.7-298.9 million years) as Phasmatocycas bridwellii, and from the Lower Permian Wellington Formation (Artinskian 283–290 million years) in the form of Phasmatocycas kansana. Relatively certain relationships can

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be found between the microtaeniopterid to macrotaeniopterid leaves of Taeniopteris (coriacea) and the fertile parts of Phasmatocycas. The fertile organs of Phasmatocycas are characterised by elongated, seed-leaves that protruded in a spiral from a central receptacle. There were two rows of 40 to 50 ovules on them, which were more or less

Sobernheimia jonkeri. a female cone from the Lower Permian whose classification is still unclear. Sobernheim, (University of Münster).

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Taeniopteris eckardtii

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Taeniopteris eckardtii.1. Original specimen from G. A. Kurtze from 1838. 2. The original label. 3. A fossil containing multiple leaves (both from the Geological-Palaeontological Collection of the University of Halle). 4. Individual leaf, Mansfeld (Coll. N. Hesse). 5. Large leaf that tapers to a point, Siersleben. 6. Extremely small leaf, Mansfeld (Coll. S. Brandt). Although a relationship to the cycads has not been proven, it appears to be plausible.

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surrounded by the leave depending on the level of maturity. The egg-shaped and somewhat pointy seeds usually had a length of 5 millimetres and a width of 2.5 millimetres, were slightly flattened, equipped with an apical cut, and connected at the lower part to the main vein of the fertile leave.

They probably only opened completely once they had reached maturity to enable the completely developed seeds to be released.

The abominable mystery of the Upper Permian Similarities with and major differences to Sobernheimia are exhibited by the Upper Permian species Pernerina pasubi from the Italian Alps. In this case, we are also dealing with a low growing plant with irregular to rounded leaf shapes and a pinnate leaf vein pattern.

Interestingly, they bore ovules and pollen organs on the same plant, as can be seen on fossil finds of complete specimens. Just as on Sobernheimia, the seeds were arranged in two rows of about twenty seeds each on a fertile leave, and the eggshaped 5 millimetre wide seeds, which became pointed at the tip, exhibit a recess in the middle that is typical for this type of plant. The shape of its leaf and its monoecious structure - with male and female sexual organs on the same plant and the lack of tuberous roots - differs from those of cycads. Isolated finds of leaves in the Dolomites prove that this plant survived the Permian and lived into the Triassic period. In any case, Pernerina pasubi is a highly intriguing plant whose classification is still extremely puzzling.

A plethora of tongue-shaped leaves from the Upper Permian of Central Europe is just as difficult to classify. The main problem here is their lack of fertile organs. Such leaves include those frequently found in 1839 by G. A. Kurtze in the Upper Permian layers of the German Zechstein (Wuchiapingian (269.9–254.2 million years ago)) and designated as Taeniopteris eckardtii. Some compound finds exhibit a growth habit similar to that of the cycads. Since similar small-leaved and low-growing leaf shapes from the Early Triassic (Ladinia) have been found with Dioonitocarpidium ovules sticking to them, they could certainly be

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placed among the ancestors of the cycads. The leaves were small, simple, tongue-shaped leaves with entire margins, and with veins emanating from the rachis at a right angle that either did not fork at all or only forked sporadically. In the Upper Permian in particular, these leaves could be round or could taper to a point at the tip. The first representatives appeared as early as the Lower Carboniferous. They became more numerous in the Permian and lived through the Triassic and into the Cretaceous period. It is certain, however, that not all of the species in the genus described in 1828

by Adolphe Brongniart belong to the ancestors of the cycads. Other possibilities include coniferlike plants such as the Pelourdea known from the Lower Triassic, or the Bennettitales.

Pernerina pasubi Reconstruction of the entire plant with female ovules, pollen organs, and individual leaves.

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Pernerina pasubi 1. Part of the plant with ovules attached and a pollen strobilus (holotype, 8cm). 2. Detailed view of an individual leaf. 3. Female organ. . 4. Reconstruction of an ovuliferous organ. 5. Detailed view of the seeds. Since this plant is a monoecious plant with male and female sex organs on the same plant, it cannot be considered a member of the cycads. They are therefore difficult to classify. Upper Permian. Valli del Pasubio, Northern Italy (Coll. M. Wachtler).

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The Enigmatic Cordaitales Tree-like plants with long, scoop-shaped leaves characterised the landscapes from the Lower Carboniferous to the Early Permian. Their cone-shaped reproductive organs give the impression that they are somehow related to the conifers, but they do not fit our image of conifers today.

Cordaites leaves, which were typically up to one meter long with delicate, parallel veins, have been found at many fossil sites in the northern hemisphere. Although they were still widespread during the Carboniferous period, their numbers dwindled rapidly over the course of the Lower Permian and they soon became extinct. Cordaites principalis in particular is often encountered up to the Lower Permian. One of the interesting features of these large-growing plants is their cone-like structure, which could reach sizes of up to twenty five centimetres and which are referred to as Cordaianthus (also Cordaitanthus), regardless of whether the cone is male or female.

The pollen organs were composed of an encasing of cover scales that were held in place by bracts.

Elongated pollen tubes were located inside them.

This makes them fundamentally different from the male cones of conifers.

In contrast, the female cones formed similar loose, cone-like structures, and a high number of sterile protecting leaves grew out of the axils of each bract. They encased the heart-shaped seeds. This type of structure makes it extremely difficult to classify a plant, probably due in part to their evolution early in the Carboniferous period. Certain features link them to the conifers, especially to the Walchiaceae and the Ortiseiacae, while other features point in a different direction. Since the entire genesis of the conifers must have had its roots in the Carboniferous period, it is perfectly possible that they are a side branch of the conifers.

Cordaites principalis. Gesamter Blattbüschel. A complete bunch of leaves. A famous fossil find that was depicted by Karl Mägdefrau in his book “Paläobiologie der Pflanzen” (The Palaeobiology of Plants, 1956). Carboniferous-Permian transition, Wettin, (Institute of Geosciences and Geography, Halle).

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Cordaites principalis. 1. 55 centimetre long, mainly complete single leaf. Niederhausen (Coll. M. Dammann). 2. Leaf with traces of feeding by insects (NM Schleusingen). 3. Reconstruction of a leaf with the typical parallel veins. 4. Female fructification and reconstruction of the ovuliferous organ.

5. Reconstruction. 6. Detail of the seed scales and seeds. 7. Male cone. 8. Reconstruction of the pollen cone and the microsporophylls, all from Lochbrunnen (Coll. M. Wachtler).

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Conifers still characterise the landscapes of the Earth today. The Araucarias (in the image: Araucaria araucana) dominated the southern hemisphere.

The Inexorable Rise of the Conifers One of the most impressive events in the evolution of life on Earth is the apparently sudden appearance and fast spread of the gymnosperms, and in particular of the conifers. While the conifers played no role at all at the height of the Carboniferous period, they had already appeared in a variety of forms during the transition from the Kasimovian to the Gzhelian age - 307 to 300 million years ago - and then spread all over the world.

Hardly anything - except for the forefather of all flowering plants - has fascinated the world of palaeobotany as much as the line of development of the gymnosperms, whether it be conifers, ginkgos, cycads, or one of the gymnosperms living a poor and marginal existence today such as the Gnetales or the enigmatic Weltwitschia.

The successful blueprint for the coniferous trees But what is it exactly that characterised all conifers (Coniferales, in many cases the Pinales or pinelike trees as well)? They are all gymnosperms, meaning plants whose ovules are not protected

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by closed carpels as in the flowering plants, the angiosperms. The sporophylls are usually arranged in a cone, which is why they are called conifers, which means “cone-bearing� in Latin.

The over 600 species living today are divided into eight families, but belong for the most part to the families Pinaceae, Podocarpaceae, and Cupressaceae. Strangely enough, the conifers spread primarily in the northern hemisphere, while the southern continents are dominated by other families such as the Araucariaceae and Podocarpaceae. This is very interesting in terms of understanding the course of evolution. In

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contrast, cypresses can be found on the southern and on the northern hemisphere.

Conifers are missing completely, in the tropical landscapes of Central America and the Amazon, but they do appear in the high altitude rainforests of Southeast Asia.

In the northern part of the world, conifers are the most characteristic group of plants, especially in the taiga of Siberia, but also in Alaska, Canada, the west coast of the USA, Mexico, China, and in the Alps. They come in a wide variety of shapes and sizes, from the smallest creeping species of pines such as the mountain pine (Pinus mugo) to the highest growing coastal redwood tree (Sequoia sempervirens), which can reach a maximum height of 112.8 meters. The conifer with the largest volume is a giant redwood tree (Sequoiadendron giganteum) which is almost 1,500 cubic meters in volume, and the thickest conifer is the Montezuma Cypress (Taxodium mucronatum) with a trunk diameter of over 11 meters. The long-living bristlecone pines of America (Pinus longaeva) reach ages of over 5,000 years.

The majority of conifers are monoecious, meaning a single tree will bear both female and male cones, but some such as the Araucaria, the yew, or the juniper are dioecious. The most common form of leaf is the needle, although some genera such as the cypresses (Cupressaceae) and the evergreens (Podocarpaceae) form flat leaves. Agathis conifers, with their tongue-shaped leaves, are a striking contradiction, while the genus Phyllocladus has short, flat, leaf-like shoots. There are needles that reach lengths of two to four hundred millimetres, such as those of the Apache pine (Pinus engelmannii).

The study of the unique features and differences between the modern conifers in particular can provide us with deep insights or parallels to the conifers of the past. In this regard, it is more than just their trunk structure and foliage, the differences and similarities of their cones, which can help us the most to understand their evolution.

Almost all conifer seeds develop in a protective cone whose size can range from a meagre two millimetres to an enormous six hundred millime-

Pines (in the image: Pinus cembra) and spruces (Picea abies) are considered typical trees in the northern hemisphere. The distinct features of almost every conifer alive today was formed in the Permian period.

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The Search for the Most Primitive Conifer Which ancestors of the gymnosperms from the Carboniferous-Permian transition are the most likely candidates for being the first ancestor of all conifers? Which fossil finds exhibit the most primitive features? Which species has the best prerequisites suggesting that all other conifers are derived from it? These questions are some of the most difficult to answer in the history of the evolution of plants.

The oldest conifer remains ever found, described as Swillingtonia denticulata were dated in the Moscovian (315 to 307 million years ago) and came from Yorkshire in England. Only microscopic remains were found, but these exhibited Y-shaped, forked tips. Amazingly, this pseudomonopodial growing pattern, in which

branching only took place on one level and one leaf dominated, extending slightly above the other, had existed since the Devonian in plants such as Sawdonia or other zosterophylls, from which all other plants subsequently derived.

While the clubmosses, horsetails and the ferns diversified in a short period of time until they

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Perneria thomsonii. 1. Detailed view of a twig with apical, tripinnate sterile leaves and sharp needles in the lower part. 2. Upper part of a twig. 3. A juvenile and an adult leaf, all from Lower Saxony (Coll. Th. Perner).

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Perneria thomsonii. 4. Female seed cone. 5. Detailed view the ovuliferous scales with bracts and seeds. 6. Male cone with pollen sacs attached to the underside. 7. Detailed view of the pollen sacs, all from Lower Saxony (Th. Perner).

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The Astonishing Development of Conifers with Winged Seeds Conifers with cones that were characterised by winged seeds appeared for the first time in the earliest part of the Lower Permian. They did not form a homogeneous group in the Permian, but it is certainly possible to draw parallels to the modern spruce or fir trees. The formation of winged seeds led to their inexorable rise, as their seedlings could travel long distances.

Spruce (Picea) and fir trees (Abies), but also larches (Larix) and Douglas firs (Pseudotsuga), are related to them, and other conifers with this blueprint still dominate many landscapes of the north today. Their woody seed cones are composed of numerous, spirally arranged ovuliferous scales which bear winged seeds on their upper surface that are only attached at the base. Certain species such as the Douglas fir (Pseudotsuga menziesii) or the bristlecone fir (Abies bracteata) are characterised by cover bracts that extend far beyond the ovuliferous scales, a feature of many Palaeozoic conifers.

The variety in appearance of these winged seeds today demonstrates the different ways in which

they have developed since the earliest stages of the Permian. It is certain that winged seeds had formed by the beginning of the evolution of conifers, and that the ability to spread seeds carried

A fir cone (Abies alba) disintegrating upon maturity. Right: A young cone with cover scales (bracts) extending past the seed scales. Top: Detailed image of the winged seed scales.

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by the wind over long distances became a model for success. This model was no less advanced than the nuts offered by other conifers to animals as food.

Wachtlerinaceae towards the modern Abietoideae, based on well-documented specimens covering the entire Permian period. We can only speculate about the origins of these bearers of winged seeds, but a series of distinct characteristics such as the structure of the needles and twigs and a certain number of sterile bracts point to a line of evolution diverging from that of the Araucarialike Walchiaceae. The split must have occurred relatively quickly at the Carboniferous-Permian border.

The Lower Permian Wachtlerina bracteata from Central Germany is characterised by relatively long bracts that extend well beyond the seed scales. In contrast to other conifers of that time, their bracts did not fork near the tip. The strongly divided ovuliferous scale with two formed winged

Wachtlerina, the bracted conifer At the Carboniferous-Permian transition (Kasimovian-Gzehlian) slightly more than 300 million years ago, conifers with the characteristic feature of winged seeds flourished, together with the most primitive Perneria and Araucarialike conifers. Although this type of seed has only been preserved in rare cases, and even then only in the finest sediments, it has still been possible to collect enough evidence for this important step in evolution. It is possible to trace the continuous development and perfection of these

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1. Pollen cones of a Norway spruce (Picea abies). 2. The upper side of a single seed scale with bract, and 3. A single seed from a fir (Abies alba). 4. Seed scale of a common spruce (Picea).

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Wachtlerina bracteata. 11. Twig. 2. Detailed image of the sickle-shaped, curved needles with median vascular bundle. 3. Reconstruction of the twig. 4. Female cone with non-laciniate bracts and reconstruction. 5. Disintegrated cone with just a few bracts and seed scales still attached to it. 6. A single bract. 7. Seed scale with attached bract. 8. A single winged seed. 9.

Reconstruction of a single bract and seed scale, Niederhausen, (Coll. Th. Perner).

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The advancement of a successful concept

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Wachtlerina suessi. 1. Twig. 2. Detailed image of the sickle-shaped, curved needles. 3. Male cone. They were usually small and round. 4. Reconstruction of a twig with male cone. 5. Immature female cone.

6. Seed scale with seeds. 7. Bract. 8. Female cone. 9. Ovuliferous scale with seeds from Collio in the Brescia Alps, (Coll. M. Wachtler).

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1 Wachtlerina suessi. A deposit of needles, probably due to the annual dropping of needles.

seeds embedded in it was located on the upper side of the bract - but connected directly to the rachis of the cone. The cone itself disintegrated exactly like the cones of modern fir trees upon maturity, leaving an empty rachis behind. Due to this separate arrangement, it is rare to find connected bracts and seed scales. The extending twigs bore sickle-shaped, curved needles, a feature common to many early Permian conifers and therefore unsuitable as an identifying characteristic for the genus.

Wachtlerina suessi from the Southern Alps represents a further development over time, until the Middle Lower Permian. The twigs and needles were similar to those of Wachtlerina bracteata, as well as the relatively long cone-bracts. The clearly divided seed scale was already more developed and more modern, and the concept of seeds surrounded by a wing is still clearly recognisable.

Layer upon layer of isolated and dropped needles were amazingly found at the main fossil site in Monte Dasdana in the Brescian Alps, and this is the first location at which it is possible to detect the highly interesting development of trees that dropped their needles annually. Whether or

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Majonica alpina. 1. Part of a twig. 2. A single leaf. 3. Reconstruction of a twig and a single leaf.

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Majonica alpina. 4. Juvenile seed cone. 5. Fully grown seed cone before it disintegrates. 6. Reconstruction of a twig with female cones. 7. Isolated bract. 8. Typical holding appendix of the ovule. 9. Ovule with two winged seeds. 10. Variability of the seeds. 11. Reconstruction of a seed scale. 12. male cone and reconstruction. Valli del Pasubio, Vicentinian Dolomites (Coll. M. Wachtler).

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not this was due to the generally unfavourable climatic conditions of this landscape during that time, and the raw climate associated with high altitudes, can only be discovered through further research. The other genera of conifers were also characterised by small leaves like those of Seymourina, or by the first appearance of massive numbers of low growing pine trees (Valentinia).

In any case, these conifers with winged seeds remained a dominant element in the landscape of conifers throughout the entire Permian period.

Majonica, the winged seed conifers Upper Permian genus Majonica was even more similar to modern spruce and fir trees. These conifers, which were classified for the first time by the Dutch palaeobotanist Johanna ClementWesterhof in the year 1987 based on fossil finds from the Dolomites, are a milestone in the history of research of the plant kingdom. Majonica alpina was characterised by its slender cones, which could be up to ten centimetres long, and also its relatively long bracts. The ovuliferous scales were similar to those of their Lower Permian ancestors, and also dropped off upon reaching maturity.

A unique feature of these conifers with winged seeds was a sterile, more lobed appendix, which has often been found isolated and which probably fulfilled the function of a holding scale.

The twigs spread out widely and irregularly, and the individual needles were long, slightly grooved, and tapered to a dull or sharp point at the tip.

These so characteristic conifers for the Southern Alps during the Upper Permian can be considered as ancestors of the modern Abietoideae, based on many features. Even more of the fir-like trees due to the seed scales and winged seeds that fell off as a whole. Similar winged seeds are also known, however, from the coeval Zechstein in Germany.

Bracts extending well past the seed scales, as here on Pseudotsuga bracteata, are still especially fascinating today, even though the evolutionary advantage of having a cover scale is not entirely clear. It probably is a functional relict from the earliest period of the evolution of the gymnosperms.

Dicranophyllum, an herbaceous conifer Another Lower Permian tree bearing winged seeds that is extremely difficult to fit into a generic concept is Dicranophyllum hallei. Although first found in Thuringia, it was the excellently preserved remains found in the Donnersberg Formation near the town of Kahlheckerhof in Rhineland-Palatinate in particular that made an

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Some of the foliage from the German copper shale likely represents conifers with winged seeds from the genus Majonica, Kamsdorf, Thuringia, (Coll. S. Brandt, 1.5cm).

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The Origins of the Pines Just like almost every other family of conifers, the modern Pinoideae, which are generally referred to as pines or Pinus, had already experienced an explosive development by the time of the transition from the Carboniferous to the Permian that brought them close to the modern pines in a very short time. Strangely enough, the multi-lobed and laciniate individual leaves of its Devonian ancestors played a role again in this case: they became longer, forming the long and slender needles we are familiar with today. The cones, in contrast, became extremely woody and had virtually reached their modern appearance during the Lower Permian.

The subfamily of the Pinoideae today consists of about 110 species that almost exclusively inhabit the northern hemisphere. They form monoecious trees or even bushes, and colonise the most inhospitable regions, including the highest mountains, due to their tenaciousness. They like to grow in the company of other pines and can cover extensive areas. Their needles rarely grow alone (except for Pinus monophylla), but usually in bundles of two to five, sometimes even in groups of eight, whereas the base of the needles is surrounded by a fascicle sheath. The seeds form in pairs on an ovuliferous scale and can either be winged, as on the Scots pine (Pinus sylvestris), or nut-like as on the Swiss pine (Pinus cembra). The cones themselves can reach lengths from just a few centimetres, as on the mountain pine (Pinus

mugo) or the Colorado pinyon (Pinus edulis), to 60 centimetres on the sugar pine (Pinus lambertiana).

The male pollen cones are usually small and grow in bunches arranged in a helix near the base of long, young shoots.

Valentinia, the most primordial pine The incredibly rapid rate at which development must have occurred at the beginning of the Permian, a rate which was never achieved again, can also be traced based on the development and divergence of the pine-like trees. Valentinia wachtleri, fossils of which have been found in the Collio Formation in the Southern Alps, in sediments deposited between the Sakmarian and Artinskian stages 283 to 280 million years ago, was a conifer with features that could easily be

Pine trees, with their easily recognisable bundled needles, dominate the scene in many landscapes of the northern hemisphere. They are characterised by their tenacious will to live. Left, Aleppo pines (Pinus halepensis) clawing onto the cliff on Mallorca, and on the right are Pinus edulis in the arid area of Utah.

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Valentinia wachtleri (holotype) from the Lower Permian: 1. Part of a twig with a cone hanging on it, and 2. Reconstruction. Collio, (Coll. M. Wachtler, 13cm).

confused with those of the twenty million year older, more primitive conifer Perneria, although its foliage is certainly recognisable as consisting of long needles. The female cones were significantly different from those of Perneria, and exhibited striking similarities to the woody cones of the modern genus Pinus. The seed cones grew on a minute stem like the cones of pines today, were no more than two to three centimetres high, and composed of a few megasporophylls, arranged in a helix, each one bore two winged seeds on their surface. The entire cones dropped off the twig upon maturity, but in contrast to the cones of fir trees at that time, the woody ovuliferous scales

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remained, inseparably connected to the cone, as on modern pine cones.

The astonishing changes that must have taken place in the short period of about five million years can be seen on pine trees found just a few kilometres away in Tregiovo, which came from an later stage, the Artinskian-Kungurian, 277 to 274 million years ago. In the meantime, the pines had clearly diverged along several lines. Valentinia angelellii still exhibited relatively short-needled, laciniate leaves, while Valentinia cassinissi, in contrast, was characterised by long-needled, bushy bundles similar to the modern, five-needle Swiss pine (Pinus cembra). The seed cones, howev-

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Valentinia wachtleri. Different individual needles with their typical fork-like appearance, 3.5cm. 3. Side view of cone and reconstruction. Note the relatively low number of seed scales, 3.0cm. 4. Top view of a female cone with seeds and a reconstruction, 2.5cm. 5. Bottom view with attached stem 2.5cm. Collio, Brescia Alps (Coll. M. Wachtler).

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The changes in Valentinia over the course of five million years heading towards the Artinskian. The relatively irregular dichotomous needles become more regular. Valentinia angelellii is characterised by relatively short needle leaves. 1. Individual leaves and reconstruction. 2 + 3. Parts of twigs. 4.

Female cone. 5. Winged seeds. 6. Reconstruction of a seed scale, Tregiovo, Trentino (Coll. F. Valentini).

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Valentinia cassinisii. These conifers are characterised by relatively long, bushy needles. 1. Part of a twig and reconstruction. 2. Individual needles. 3.

Female cone and reconstruction from different positions. 4. Pollen cone and reconstruction. Both male and female cones stood upright on modified leaves. Tregiovo, Trentino (Coll. F. Valentini).

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1. Arrangement of the needles on modern pine trees: Swiss pine (Pinus cembra) with five needles. 2. A bundled arrangement of two needles and a cone from the mountain pine (Pinus mugo). 3. Top, outer, and inner view of a mountain pine (Pinus mugo) cone with a detailed image of the seeds.

As early as the Lower Permian, there was a marked tendency towards collecting two (e.g. Valentinia angelellii or the modern black pine), or several needles in a bundle, as on Valentinia cassinisi or the modern Swiss pine.

er, only changed marginally, and have stayed that way until today. They remained woody and thick with deep grooves and furrows on the underside and bore an umbo at the end. The two seeds per scale were only a few millimetres in size and were slightly to heavily winged.

The relationship between ginkgos and pines An interesting feature of the needles of these early pines is their striking similarity to the ginkgo leaves of that period. It is especially difficult to differentiate them from Permian ginkgophyte

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(Baiera digitata), or a genus classified under the taxon Esterella (gracilis), especially when cones or ginkgo seeds are missing. It is therefore possible to derive a relationship between the ginkgos and the pines.

The structure of the cones and seeds are different, however, and if they are related, then they must have sprung from a common ancestor several million years earlier near the end of the boundary between the Carboniferous and Permian, which would certainly appear to be plausible.

In any case, the discovery of these very early bearers of winged seeds, which are otherwise extremely different in terms of type (whether it be Valentinia or Wachtlerina), and the long period in which they have retained their form, is one of the most astonishing occurrences in the development of the plant kingdom. Equally surprising is their function as a missing link between the earliest form of the Devonian plants and the vegetation still dominating the landscape today.

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The Strange Evolution of the Ginkgos Ginkgo biloba tree stands like a recluse among the flora of today and is still surrounded by mystery.

Its evolutionary history is also astonishing since it has been a witness to the history of the Earth from the Early Permian to today. They also demonstrate how many plants can develop over time.

From the most primitive gymnosperms, with leaves exhibiting dichotomous venation and veins that fork repeatedly, all other gymnosperms developed in quick succession. Plants with long leaves with dichotomous venation appeared for the first time in the Permian. Amazingly, these trees had already been identified as ginkgo trees in 1843 by the Bavarian apothecary Karl Friedrich Wilhelm Braun under the name Baiera (digitata). Clearly fan-shaped leaves with a repeatedly forking vein structure - typical of the modern Ginkgo biloba - are only known to have existed since the Tertiary, which makes Braun’s evaluation even more remarkable.

Baiera perneri from the Carboniferous-Permian transition can be classified as close to the original line of evolution of ginkgo trees due to its foliage, but even more distinctive are the characteristics of Baiera pohli from the Alps of the Lower Permian. Except for its lack of fan-shaped leaves, many of its features are similar to those of modern ginkgo trees. These characteristics in-

clude the collar-like ring from which the leaves emanate, just as on modern ginkgos, and the two upright ovules that form together. The ovules did not hang from the end of a distinctive stalk like those on a modern ginkgo, however, but were held by a modified leaf segment. On the one hand, their lobe-like needles are reminiscent of the much more primordial Perneria, but they are also easy to confuse with the needles of one of the most primitive Pinoideae, the Valentinia conifers. A fossil can only be identified as being from a ginkgo tree with certainty when the ovules are found, which is rare, so that in many cases the classification of sterile leaf material identified as Baiera, Sphenobaiera, or Esterella is questionable or, should more often be identified as one of the pine-like Valentiniaceae. The tattered foliage of the ancient ginkgos was retained for many millions of years. Baiera digitata from the Upper Permian in Europe differed only minimally from the Lower Permian version, and the same applies to Baiera taeniata from the Lias in Germany.

The colourful variety in appearances of the leaves can still be seen today on the Ginkgo biloba. Middle: In contrast, the Ginkgo adiantoides from the Palaeocene in North Dakota already exhibited characteristics of the modern ginkgo leaf.

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Baiera perneri, one of the earliest ginkgo trees from the Carboniferous-Permian transition. Left: Niederhausen, (Coll. Th. Perner, 3cm). Right: (PM Nierstein, 3cm). Reconstruction.

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The initial evolution of the ginkgos 4

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Psygmophyllum sp. Lower Permian, Nowo-Wladimir-Andrejewsk (Prof. Dr. Lahusen, Petersberg, now the Museum of Natural History Chemnitz, 8cm).

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1. This Baiera digitata ginkgo leaf specimen from the Upper Permian has been published numerous times 8cm. 2. Possible ginkgo seeds from the same fossil site. Both from Volkstedt, (Museum of Natural History at the Humboldt University of Berlin). 3. Another Baiera digitata leaf with four lobes, Mansfeld, (Coll. N. Hesse, 8cm). 4. Male pollen cone of Ginkgo biloba. 5. Ovule. 6. Leaves sprouting out of the typical Ginkgo shoots.

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Fish, a Model for Success More than half of all species of vertebrates existing today, which is about 33,000 species, are fish. All vertebrates living in water that propel themselves forward with fins, breathe with gills, and are not terrestrial vertebrates are called fish.

The oldest known (but jawless) fish worthy of fish is by examining the design of their the name originated in the Early Ordovician fins. These designs were important for the about 450–470 million years ago, although evolution of all living creatures, especially there were also fish-like chordates such as the land animals: the fins of the ray-finned Haikouella, found in rock strata in China that fish are formed by a row of narrow rays made is over 540 million years old. The cartilagiof a bony substance or cartilage, while on nous fish appeared in the Silurian/Devonian the lobe-finned fish, on the other hand, the transition around 420 million years ago. Bony fins are supported by individual basal bones fish have existed since the Devonian, although moved by muscles. The arms and appendages they may have begun their evolution as early as of all terrestrial vertebrates living today the Silurian, and then evolved to form numerprobably originated from the pectoral and ous families after that. The primitive lampreys pelvic fins extending from two sides of the (Petromyzontiformes) form an order of very body. primitive, gilled, fish-like vertebrates that have Other groups of fish important for an barely changed over the last 500 million years. understanding of life in the Permian period, They have eel-like, elongated bodies with dorbut extinct by the end of the Palaeozoic era sal fins and a tail fin. should also be mentioned: these groups To understand the fish of the Permian, it is a include the Placodermi, the armoured fish, good idea to obtain a superficial overview of and the spiny shark (Acanthodii). the large families of fish existing today. These include the cartilaginous fish (Chondrichthyes), in large groups such as the sharks and rays, and the bony fish (Osteichthyes), which are divided into the subclasses lobe-finned fish and rayfinned fish. The bony fish differ from the cartilaginous fish in that their skeleton is completely or at least partially ossified.

Almost all fish groups in the world, including all European freshwater fish, are classified as ray-finned fish. In contrast, the coelacanths and lungfish, which were so important for the evolution of all vertebrates, are Stingrays (Himantura Fai) and blacktip reef sharks (Carcharhinus melanopterus) prefer the classified as lobe-finned fish. The reefs. This probably applied to the rays and shark-like fish of the Permian. The elongation of best way to differentiate between the skull before the pectoral fins is the most obvious difference from the rays alive today. Fish ray-finned fish and lobe-finned populated the seas well before there was life on land. Sharks are one of the oldest fish.

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Lower Permian: The End of the Spiny Shark Over a period of almost 170 million years from the Lower Silurian to the Lower Permian, a single group of fish developed highly successful “jaws� and dominated the seas: the acanthodians, which are also referred to as spiny sharks. During their evolution, their mouth rims changed and evolved to form segments of cartilage or bone that were connected by a hinge mechanism. With this organisation they could grab and chew food. In contrast to other families that evolved in the course of the Devonian and Carboniferous periods, the spiny sharks did not experience much pressure to adapt or evolve any further. Their blueprints barely changed, and they eventually became extinct, which is why no fish like them exist today.

One special feature of the acanthodians was their scalation, which consisted of a pattern of cubical to rhomboid platelets that, in contrast to most fish today, did not overlap and were arranged in a mosaic pattern instead. All species of acanthodians had jaws, but often no teeth, and they also had only one dorsal fin. Their main skeleton, however, consisted of ring-shaped collars of bone that varied in size depending on growth stage, or even on environmental influences such as the supply of food, water, or variations in the climate. The result is that their preserved remains can differ greatly depending on the fossil site and on the rock layer they are found in, making proper classification difficult. They colonised the open sea in particular, and probably fed primarily on plankton and other marine creatures. Their skin-covered fins had bony spines along the front edge, which is why they are called spiny sharks, even though they are not related in any way to the sharks that dominate the oceans of today all over the world.

The spiny shark Acanthodes bronni, Odernheim (Coll. G. Loth, 14cm)

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Acanthodes bronni. 1. Niederkirchen (MoP Nierstein, 34cm). 2. Complete specimen, and 3. Detailed view of the tail fin with its typical scale pattern, Niederkirchen (Coll. Th. Perner, 10cm). 4. Reconstruction (From Heidtke, 1990). 5. Acanthodes sp. “Palatinate”, Niederkirchen (Th. Perner, 8cm). 6.

Acanthodes sp. “Palatinate” reconstruction (From Heidtke, 1990).

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The spiny shark Acanthodes Acanthodes bronni became known as a characteristic species of the Lower Permian in Europe, because numerous fossils have been found in Europe. They were spiny sharks that grew to relatively large sizes with a length of up to 80 centimetres. Like all fish in this family, Acanthodes had bony spines located on various parts of the body. Cubical scales covered its body like tiles. The skeleton was made of cartilage.

Most acanthodians, especially the young ones up to 30 centimetres long, were plankton eaters that ate by filtering the water with the help of their mouths and gill rakers. This applies to all small Acanthodes. In larger animals, the rakers on the lower jaw changed into pin-like objects similar to teeth and allowed them to hunt for food as predators. The remains of small amphibians and even small acanthodians have been found in the stomachs of large acanthodians. They were therefore definitely cannibalistic.

to be called Acanthodes “Palatinate� sp. They were probably also plankton eaters. A particularity about them was that only the tail and the part of the body from the middle to the rear had an arrow-like scale arrangement pointing towards the head.

The Acanthodian Westrichus The rare Westrichus kraetschmeri differs from the other acanthodians in that it has an elongated pelvic fin. Fully grown specimens therefore look bulky. The head, however, was relatively small.

It fed mainly on small crustaceans, but did not hesitate to eat small fish, amphibians, and other members of its own species.

The Acanthodes sp. Palatinate These unusual acanthodians are found especially in the German Palatinate, which is why they used Westrichus kraetschmeri (after Heidtke 1990)

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The spiny shark Westrichus kraetschmeri (holotype) from the Lower Permian, Niederkirchen (Mainz Museum of Natural History).

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Xenacanthida, the Terror of Permian Swamps The xenacanth sharks include a series of fossilized fish which were found primarily in freshwater sediments and dominated the lake areas of what is now called Europe from the Middle Carboniferous until the Upper Triassic period. Their skeletons could reach lengths from thirty centimetres to three meters. In addition to the difference in size, they also differed in terms of the shape and manner in which the spine on the neck was attached to the body, their pairs of opposing fins, and their teeth. The teeth consist of two larger tips on the sides and one normally smaller tip in the middle, arranged in several rows to create a set of “revolving teeth� in the jaw.

Only the front rows were actually used for feeding purposes. If a tooth fell out or broke off, then a new tooth from the row behind it immediately pushed forward into the front row. Xenacanthida fossils from the Lower Permian have been found, most commonly of the up to three meter long Orthacanthus, the much smaller Xenacanthus, and smaller genera with body lengths up to one meter long like Triodus and Plicatodus.

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The largest single-spine shark Orthacanthus The freshwater shark Orthacanthus senckenbergianus was the largest species of single-spine sharks known to date. The females, which were larger than males, could reach lengths of over three meters. They were the most feared predators in large and deep bodies of water in the Lower Permian. Their long, narrow bodies and the considerable size of their fins made them ideal hunters that stalked their prey near the bottom, or from a hiding place. Similar behaviour is seen today in fully grown catfish. Due to their large size, they were able to prey on practically every other creature in the water using fast surprise attacks. A special feature of these sharks was their single spine, which was located directly behind the head and fastened to the pectoral girdle on Orthacanthus. One distinctive feature was their teeth, which had three points that, due to their different shapes, are an important classification

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Orthacanthus senckenbergianus. 1. This is a two meter long male specimen with penis-like clasping organs on the pelvic fins, Niederkirchen (MoP Nierstein). 2. Reconstruction (from Heidtke, 2003).

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Abundance of Fish in the Zechstein Sea The changing climate in the late Lower Permian placed high demands on living creatures to adapt accordingly. Temperatures rose, and rainfall levels dropped, which caused the seas of the Rotliegend to dry out. Mountains eroded, and the eroded sediments were deposited in the valleys. Only a few creatures were able to adapt to the shifting climate and evolve further.

It was only after greater parts of Europe flooded from the northeast, and the formation of a large sea, that numerous marine creatures were able to inhabit modern Europe. This decisive turning point in nature was the reason the Zechstein Sea formed in Europe. Metals carried out to the sea from land and hydrothermal solutions rising up from the deeper, oxygen-poor parts of the Zechstein Sea, where digested sludge was deposited, led to the polymetallic assemblage of copper, and the formation of the German Copper Slate. During periods in which salinisation levels were low, the waters were suffused with light and dominated by an abundance of marine fauna, primarily by the bony fish, but also by the primitive cartilaginous fish. One fish in particular, which makes up 90 per cent of all specimens

found today in the seas of the Copper Slate, would become the characteristic fossil of these seas: Palaeoniscum freieslebeni, a slender Palaeoniscidae that evolved from a group of ganoid fish from the Lower Permian and then evolved even further. It inhabited the seas of the Upper Permian in the northern hemisphere, and large amounts of fossil remains have been found there. Other common bony fish were Platysomus and Pygopterus. In contrast to the Rotliegend period, lobe-finned fish adapted to a life in the sea swam in these seas, in addition to rare specimens of real bony fish from the family of Semionotidae such as Acentrophorus.

The cartilaginous ganoids were made up of new sharks (Euselachii) Wodnika and Hopleacanthus, petalodont fish from the genus Janassa, and chimaeras (Holocephali) such as Menaspis.

Three Palaeoniscum freieslebeni simultaneously embedded in the mud of the Copper Slate, Bad Sachsa (Coll. G. Loth).

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The Cartilaginous Fish of the Upper Permian Sharks with more than 500 species, rays with over 600 species, and the less well known chimaeras with 34 species all belong to the group of cartilaginous fish (Chondrichthyes). This means only about four per cent of the species of fish living today are cartilaginous fish. The whale shark, with a length of 14 meters and a maximum weight of 12 tons, is the largest cartilaginous fish living today.

They reach lengths that are much greater than the lengths of bony fish.

The petalodont fish Janassa Janassa bituminosa, from the family of Petalodontiformes (“petal or f lattened teeth�) was a medium-sized ray-like fish that reached a length of one meter and a width of 25 centimetres. It was the most common cartilaginous fish in the Zechstein Sea. The head tapered to a point while its mouth was located on its underside. With its revolving teeth in overlapping rows, it fed primarily on hard-shelled animals such as shellfish, mussels, brachiopods, or sea lilies. In addition to two large pectoral fins, it also had two pelvic fins that extended from both sides of its body, as on a ray and a dorsal fin at the beginning of the tail that ended in a round tail fin. The flat Janassa korni must have looked similar to Janassa bituminosa, although it had a different number of points on its teeth and a smaller set of teeth.

The ray-like fish Rays were traditionally viewed as a taxon of cartilaginous fish of the same rank as the sharks.

The primitive guitarfish, the sawfishes and electric rays propel themselves, like most sharks, by moving their tail fin from side to side. Real rays, however, swing their large pectoral fins like waves, and eagle rays beat them like wings.

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Representations of the creatures living in the Zechstein. According to NM Schleusingen: 1. Palaeoniscum 2. Pygopterus 3. Wodnika 4. Platysomus 5. Eurysomus 6. Acentrophorus 7. Dorypterus 8. Janassa 9. Coelacanthus

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1. Complete specimen of Janassa bituminosa, length 48cm, Bad Sachsa (photo + M. Klopschar collection). 2. Teeth of Janassa bituminosa, Eisleben (University of Halle, photo by S. Brandt). 3. Janassa korni, Eisleben (Coll. S. Brandt, 15cm), and 4. Teeth of Janassa korni (photo and Coll. N. Hesse). 5.

Reconstruction of Janassa bituminosa (from Haubold & Schaumberg, 1985).

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The shark-like fish

The rare Hopleacanthus

The Zechstein shark Wodnika Wodnika striatula, a medium-sized fish that could reach a length of one meter, was characterised by its slender head and relatively large eyes. The teeth were bean-shaped and slightly curved. It must have fed primarily on hard-shelled animals like mussels, brachiopods, and shellfish. Wodnika was equipped with two spikes in front of the two dorsal fins, where the front spike extended about one third of its length past the rear one.

Hopleacanthus richelsdorfensis was a mediumsized shark-like fish up to one meter long with a flattened head. The jaws were provided with teeth bearing a single point that appeared very much like those of modern species. This indicates a predatory lifestyle similar to that of modern sharks. It was equipped with two smooth backpointing spikes in front of each dorsal fin, but in contrast to Wodnika, the front spike was smaller than the rear spike. The dorsal fins were also shifted far to the rear of its body, and the front dorsal fin was located at about the same level as the rear pectoral fin. The pectoral fins were relatively large, while the anal fin flowed almost seamlessly into the tail fin.

A modern lemon shark (Negaprion brevirostris). The type and arrangement of its razor-sharp teeth has hardly changed at all up to today.

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Hopleacanthus richelsdorfensis.

1. A young specimen, 30cm, Bad Sachsa (photo + M. Klopschar collection). 2. Remains of a lower jaw, Eisleben (Coll. S.Brandt).

3. Wodnika striatula, Mansfeld (photo + N. Hesse collection). 4.

Reconstruction (From Schaumberg).

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1. Coelacanthus granulatus. Bad Sachsa, 30cm (photo + M. Klopschar collection). 2. A coelacanth eating a fish (Palaeoniscum freieslebeni). Only its tail is still sticking out. Bad Sachsa (NM Schleusingen). 3. A historical piece, Richelsdorf (MoNH Vienna). 4. Reconstruction (from Moy-Thomas, 1935, and Haubold & Schaumberg).

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Comparison with one of the lagoons more or less filled with water during the labyrinthodont period. A myriad of amphibians in the various stages of growth lived in such lagoons.

The Triumph of the Terrestrial Vertebrates To understand the development of the animal kingdom on land, it is necessary to trace the transition from life in water to life on land, which is one of the most fascinating events in the history of evolution. Amphibians, reptiles, birds, and mammals are classified today under the taxon Tetrapoda. They include all four-limbed vertebrates except for the fish and the jawless Agnatha.

Further subdivisions have been useful for all animals that can reproduce independently from water (amniotes) and for the non-amniotic, more primitive amphibians, which still need to lay their eggs today in water or wetlands to keep them from drying out, since their eggs lack a protective eggshell. They usually spend most of the first stage of their lives in the water as larva. Later, they go through a metamorphosis to become animals that can live in water or on land.

Fossil finds from the Devonian show that the ancestors of all modern land animals, from

amphibians to reptiles and even mammals, were once fish-like vertebrates. The first animals that risked going on land – even if only for a short time – were accordingly elongated fish equipped with a tail fin and two further pairs of fins, from which the limbs of all terrestrial vertebrates would then develop.

It is generally assumed that the following creatures existing in the Devonian could potentially be the ancestors of all terrestrial vertebrates living today: the lungfish (Dipnoi) and the lobe-finned fish (Crossopterygii).

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The strange way of life of the cut vertebrae The most widespread group worldwide between the Carboniferous and Permian was the Temnospondyli (“cut vertebrae”), a group of primitive amphibians including such important families as the Archegosaurus (Sclerocephalus, Cheliderpeton, Archegosaurus), the Dissorophidae (Micromelerpeton, Eimerisaurus), and the Branchiosaurians (Apateon, Melanerpeton). They ranged from animals that lived only in fresh water or near the sea, to creatures partially or completely adapted to land.

Due to the adaptation in varying degrees to life on land, the cut vertebrates formed lungs for breathing, and a spinal column from ossified bone with the associated shoulder, rib, and hip bones.

They also ranged considerably in terms of their size, with some species reaching a length of one to two meters, such as Archegosaurus – which looked like a modern crocodile from the outside – to the small to medium-sized Dissorophidae, which were

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not unlike modern salamanders. The ancestors of all modern amphibians must have sprung from this line. The most commonly found fossils from the Lower Permian are the Micromelerpeton – probably due to the fact that they lived primarily in ponds and small lakes and therefore lived in outstanding conditions for preservation – and the tiny branchiosaurians, which is the most commonly found animal in the group of amphibians.

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The strange world of the branchiosaurs in the Lower Permian: 1. Apateon caducus, Heimkirchen (MoP Bad Dürkheim). 2. Apateon dracyiensis, lower Goldlauter Formation, Cabarz near Tabarz. 3. Apateon flagrifera, including some parts with skin and gill filaments, Upper Goldlauter Formation, Friedrichroda (NM Schleusingen).

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Branchiosaurs, the gilled amphibians The now extinct family of branchiosaurs included several species, of which the well-known Apateon pedestris was probably the most common form in terms of numbers in the Lower Permian, especially in the Central Europe of today. Other species, such as Apateon caducus or the genus Melanerpeton were much rarer, but all of these species differed only slightly from each other.

The small branchiosaurs, which was only between five and ten centimetre long, was an important component of a wet belt that extended from

southern France to Rhineland-Palatinate, Hessen, Thuringia, and Saxony to the Czech Republic. In the Lower Permian, extensive flatlands formed in these areas and contained lakes, small bodies of water, rivers, and creeks that were fed by seasonal, monsoon-like rainfalls. However, these areas must also have experienced frequent and long

Reconstruction of branchiosaur Apateon pedestris.

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Apateon pedestris. 1. Preserved body, Odernheim (Natural History Museum of the Palatinate in Bad DĂźrkheim). 2. A large number of newts, just as today. From left to right: Alpine newt (Ichthyosaura alpestris), Bosca‘s newt (Lissotriton boscai), Carpathian newt (Lissotriton montandoni), great crested newt, twice (Triturus cristatus), fire salamander (Salamandra salamandra) (Natural History Museum Vienna).

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dry periods that could bring life to the brink of disaster. All in all, these were good prerequisites for forcing a large number of amphibians to adapt continuously to changes in environmental conditions and to develop new species.

The habitat and the appearance of branchiosaurs was probably similar to those of modern salamanders or newts. Their level of development forced them to search for water, even if only to lay their eggs. The structure of the limbs of fully grown animals suggests that they were not able to walk on land for very long. According to the current state of research, all species of branchiosaurs were equipped with external gills, regardless of their age, which means that they must have breathed through their gills their entire life and did not experience a true metamorphosis – like the salamanders – and therefore even reproduced in a larval stage.

They probably fed on everything they could overpower, which, depending on their size, could have been insects, worms, or even smaller members of their own species, and also fed on floating organic matter that they filtered from the water using their frilly gills. They were probably often preyed upon by larger amphibians, fish, or freshwater sharks, however. This made them an important part of the food chain, as demonstrated by fossilized faeces (coprolites).

Apateon caducus has only been found in a few geological layers and was significantly larger, with a skull length of thirty millimetres, and also much rarer – even however representatives in the form of Apateon pedestris lived together in the same waters, and their skulls only differed in terms of minor details.

Apateon dracyiensis, found particularly in Drachy/St Loup in France, in Saxony and Thuringia in Germany, spread out over a very extensive area. It probably lived in an environment similar to that of other Apateon species, but they exhibit different skull features and were also significantly smaller.

Melanerpeton humbergense differed from the Apateon species in that the front of its skull, the upper jaw, and the body skeleton were more slender.

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Various branchiosaurs from the group of Melanerpeton. 1. Melanerpeton eisfeldi, upper Goldlauter Formation, Friedrichroda, below: Melanerpeton arnhardtii, lower Goldlauter Formation, Lochbrunnen (NM Schleusingen) 3. Apateon dracyiensis, Thuringia (Coll. Th. Perner).

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Micromelerpeton, the Permian Salamander Several species belong to the family of Micromelerpetontidae, of which Micromelerpeton credneri is the most commonly found and best documented species. Overall, these species were all rarer than the branchiosaurs.

Lower Permian fauna: A juvenile Sclerocephalus haeuseri, together with the branchiosaurus Micromelerpeton, the palaeoniscoid fish Paramblypterus, and the large-eyed ganoid Aedulla indicate the diversity of life at that time, Odernheim, Palatinate (Coll. Th. Perner).

The skull of Micromelerpeton credneri was long and arrow-shaped, while the skull of the branchiosaurs was more semicircular. The arms and legs of Micromelerpeton were short in proportion to their bodies, and their eyes were smaller. Since their external gills were significantly smaller, it could be presumed that they left water, at least for certain period of time. They must therefore have been adapted to life on land, like modern salamanders, so that they only needed to return to water to lay their eggs. Specimens of Micromelerpeton credneri without gills, for which the process of metamorphosis was complete, have also been found. With a body length over twenty centimetres, Micromelerpeton was significantly larger than the branchiosaurs, which could only grow to a few centimetres in length. They did share the water with them, however, and were very likely preyed upon by Micromelerpeton. In addition, they fed on small fish and amphibians, as well as their larva, and did not refrain from eating worms and water insects.

Micromelerpeton credneri (after Boy, 2007).

Micromelerpeton credneri, Odernheim (PoM Nierstein)

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3 Micromelerpeton credneri, all from Odernheim. Various forms of preservation and mutations, with four to five toes. 1. D端rkheim Museum. 2. Coll. G. Loth. 3. MoP Museum.

4. + 5. The specimen below even shows imprints of the skin (MoP Nierstein).

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The Dominating Archegosaurus The largest amphibians in the Lower Rotliegend were the temnospondyl amphibians (Stegocephalia) with body lengths of up to two meters. In the larval stage, they still had outer gills and preferred to live in the water, but as they matured, their gills would retract and they would breath through their lungs, or possibly even through their skin. It can be assumed that these animals preferred to inhabit waters close to the shore. They could only move clumsily on land due to their primitively formed limbs, which is why they must have only left the water for short periods of time.

These animals, which are not unlike modern crocodiles in terms of their appearance, had relatively large, remarkably massive, ossified skulls consisting of typically round or nearly round polygonal indentations. The snout was elongated, the eye sockets were narrow and looked upwards since the roof of the head arched only slightly. The nostrils were relatively small and were located wide apart from each other on the tip of the snout. The stomach area was covered by a network of spindle-shaped, bony scales. The strong dentition and their powerful jaws point to a predatory lifestyle.

The temnospondyl amphibians also include the frequently found Sclerocephalus with its wide skull and a blunt, round snout, the rarer Cheliderpeton with its snout that tapered to a point, and Ar-

chegosaurus with an extremely elongated, pointy head. These large amphibians, which belong to the Eryiopidae, then died out towards the end of the Permian.

The hard-headed Sclerocephalus The best known specimens, particularly the outstandingly preserved specimens from the famous fossil site in Odernheim in Rhineland-Palatinate, were of Sclerocephalus haeuseri. This animal was the largest of its kind during the Lower Permian, and they could reach lengths of up to two meters. Since specimens in every stage of growth, from larva to the fully grown animal, have been found, its life cycle is relatively well known.

A particularly large specimen of Sclerocephalus haeuseri from the Lower Permian about 295 million years ago, Jeckenbach, Rhineland-Palatinate (Museum of Natural History, Stuttgart, 1950cm).

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Sclerocephalus haeuseri. 1. Skull, and 2. rows of teeth, Odernheim (MoP Nierstein). 3. Shape of the skull of a young animal, Odernheim (Siber collection, Aathal Dinosaur Museum). 4. An adult specimen, Rümmelbach Lebach (MoP Bad Dürkheim). 5. Sclerocephalus must have looked like the modern hellbender (Cryptobranchus alleganiensis) (from: “Fische, Reptilien und Lurche” (Fish, Reptiles, and Amphibians), Schleyer 1900).

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Im Stein erhaltene Lebensgemeinschaften 1

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Unknown amphibian together with a Seymourina conifer branch, Niederhausen. 2. Unknown amphibian in the company of the palaeoniscoid fish Paramblypterus. It must have been its preferred prey (both from Coll. M. Damann, Berlin). 3. Another excellently preserved conifer branch together with Sclerocephalus haeuseri, Niederhausen (Coll. Th. Perner).

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The Amphibian-like Lepospondyls This group contains terrestrial vertebrates that usually only reached a size of up to twenty centimetres and were similar modern caudates or caecilians. Its members were bizarre forms of amphibians that still generate differing opinions as to their origins. Some of these animals had extremely elongated, snake-like bodies with up to 230 vertebrae and only rudimentarily developed limbs. Others completely lacked extremities. It would appear that the lepospondyls were a highly specialised branch of the amphibians which led to a dead end.

There are theories suggesting that lepospondyls survived into modern times in the form of legless caecilians (Gymnophiona). Some primarily colonised land, while others remained inhabitants of water. Their skeleton was only weakly ossified, the enamel of their powerful teeth was not folded like a labyrinth, the snout was short, and the skull stout. The back edge of the skull was straight, and they were missing one ear slit. Their ribs were long and curved. The most important feature, from which their name is also derived, was their spool-shaped vertebrae. They included the group of Microsaurs, especially the Batropetes, the Nectridea, the Aistopoda, the Lysorophidae, the Adelogyrinidae, and the Acherontiscidae.

The multi-vertebrate Microsauria They were small animals that clearly did not belong to the amphibians or to the reptiles. Charac-

teristic for Microbrachis pelicani from the Czech Republic was their large number of trunk vertebrae (over forty) and their relatively short extremities. The lungs were poorly developed. That gave them an appearance analogue to that of the modern glass lizards, and they probably ate similar food and led a same life. Their body length was only between eight and fifteen centimetres. The group contains salamander-like as well as snakelike animals.

The small reptilian Batropetes Batropetes, which also belongs to the order Microsauria, was a small amphibian that reached a length of approximately seven centimetres. At first glance, poorly preserved specimens look similar to Apateon pedestris, with which they shared the same habitat, due to their size. The head was short and therefore appeared round.

The caudate Microbrachis pelicani, Nydany, Czech Republic (Natural History Museum Vienna).

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Additional features were its massively ossified vertebrae, the long ribs, and a long, ossified tail.

This makes it probable that we are dealing with a purely terrestrial animal with a lifestyle that had adapted, such as the lifestyles of small reptiles still living today. Its prey must have consisted mainly of insects, just like extant lizards.

The dolomitic reptile Tridentinosaurus While the amphibians - and the protoreptiles - from the Lower Permian in Germany can be considered satisfactorily understood, the same does not apply to the rest of Europe. A small, twenty-five centimetre long vertebrate from the Dolomites (Stromaiolo), is known by the name Tridentinosaurus antiquus, which, strangely enough, was “grilled� in the true sense of the word during a volcanic eruption. For this reason, remains of skin impressions were preserved, but the skeletons remained hidden under the skin.

Parts of the head are also missing. This lizard-like animal was characterised by a slender body and delicate front and hind legs. It can be assumed that the Dromopus tridactylus tracks found in large numbers from the Lower Permian were left behind by these animals.

A preserved skeleton of Batroptetes, Niederhausen (Naturmuseum Mainz).

Tridentinosaurus antiquus from the Dolomites of the Lower Permian (University and Museum of Padova).

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The Saurians in the Upper Permian Although one could say that the animals of the Lower Permian were numerous, preserved as fossils, partly due to the ideal conditions for preservation of amphibians (since they lived both in water and on land), this does not apply to the time after the Lower Permian. Evidence of the transformation, especially the diversification of the terrestrial vertebrates into the reptiles, precursors of the dinosaurs, or lizards, is scarce to say the least. Researchers have jumped at every partial skeleton in order to put the pieces of the puzzle into place.

As early as the Upper Permian, fossils of amphibians become rarer and rarer in favour of the real reptiles. The step from water to a permanent existence on land had been achieved.

From that point on, the diapsids, the most diverse group of reptiles ever in the history of the Earth would take front stage.

The first reptiles capable of flight, such as Coelurosauravus, rose into the air and conquered new niches. Others like Protorosaurus, however, continued to diverge unabated in the direction of the dinosaurs, or even the mammals. Plant eaters such as the cow-like pareiasaurs, which belong to the other large group of anapsids, began to reign over expansive plains while small lizards started spreading out in all directions. The primary reason for the lack of fossils was certainly not the disappearance of animals in the Upper Permian, but probably the lack of potential sites for preservation due to the more desert-like climate.

The Permian flying lizard Coelurosauravus The history of this oldest known flying reptile, which was capable of gliding, is long, especially because it has been found in Upper Permian strata in areas ranging from Madagascar to England and Germany. A skeleton discovered in the Copper Slate near Eisleben was purchased in 1913 by Otto Jaekel for the collection at the University of Greifswald and described by the German palaeontologist Johannes Weigelt in 1930 as Palaeochamaeleo jaekeli.

Although it was offered at that time as an example of a “flying dinosaur�, Weigelt considered the bone fragments of the gliding wings to be the fin rays of a lobe-finned fish that were incidentally embedded together with the reptile skeleton.

Since the name Palaeochamaeleo had already been

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The common flying dragon (Draco volans) is a small agamid from the tropical rainforests of Southeast Asia and looked similar to the Upper Permian Coelurosauravus (from Pantologia 1813).

assigned to the fossils of a species of agamids, it was later renamed Weigeltisaurus jaekeli.

Weigelt named the fragment of another fossil find that included one extremity Gracilisaurus ottoi. In 1926, however, the French researcher Jean Piveteau described a fossil from the Upper Permian Sakamena Formation as Coelurosauravus elivensis. In 1987, researchers realised that these three species needed to be combined under one name, and these primitive European diapsids were finally assigned the name Coelurosauravus

Coelurosauravus (Weigeltisaurus) jaekeli. Reconstruction (after Schaumberg, Unwin & Brandt 2007).

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1

2

3

Coelurosauravus (Weigeltisaurus) jaekeli. 1.

Specimen from the State Museum of Natural History Karlsruhe. 2. Holotype of the Gracilisaurus ottoi specimen described by Weigelt in 1930 based only on its extremities (Geological-Palaeontological Institute Halle). 3. Specimen found

by Thomas Schneider in Mansfeld (photograph

by N. Hesse).

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jaekeli. Similar to the modern flying dragon (Draco volans) from the reptilian family of agamids, Coelurosauravus had membranes that allowed them to glide through the air. In contrast to other reptiles capable of flight, the gliding membranes of Coelurosauravus were not spanned by bones that were already present on other nonflying ancestors of that time, but were made up of 28 pairs of curved, bony rods that were located on the sides of the rib cage and were newly created ossifications of the skin. These bony rods were similar to those still existing today on numerous reptiles or crocodiles, which are usually part of the animal‘s armour. With a wingspan of about 30 centimetres, a length of 50 centimetres, and an estimated weight of 200 grammes, it must have been an excellent glider. It was probably able to fly through the air from tree to tree over longer distances, possibly even nearby islands, and thus was able to expand into new habitats through its ability to conquer the air. This is why these Permian dragons were classified in the group Avicephala together with the Longisquama and drepanosaurs that evolved later, which therefore puts them in a lineage that is not related to the birds.

Protorosaurus, the lizard-like reptile The Protorosaurus sticks out among the few finds of reptile fossils from the Upper Permian in Europe. Today, this reptile is placed in the lineage of the archosauromorphic archosaurs, as quasi ancestors of the precursors of the dinosaurs, snakes, and lizards. Remains have been found in the Copper Slate in Saxony-Anhalt, Thuringia, North Rhine-Westphalia, Lower Saxony, and Hessen, but also as far away as England. The main species Protorosaurus speneri was named in honour of the first person to describe it, Christian Maximilian Spener, who amazingly had already described the first specimen in 1710, although he then considered it to be a crocodile that had died during the Great Flood. Over the course of time, a large number of Protorosaurus fossils were found, among which were complete skeletons, so that it is now one of the most well studied reptiles of the Upper Permian of Central Europe, which otherwise has produced so few

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fossils. Protorosaurus was a relatively long-necked lizard that, in spite of its conical, pointed teeth, ate plants, since seeds of the conifer Ullmannia were found together with gastroliths in the digestive tract of the animals, which is certainly

Protorosaurus speneri. First image of the specimen found in 1706 in Feldschacht zu Kupfersuhl near Eisenach. It was initially described and classified in 1710 by Christian Maximilian Spener as a crocodile.

an astonishing discovery. The animal itself was characterised by gastralia (abdominal ribs), which are skeletal structures that only exist today in a few reptiles such as the tuatara endemic to New Zealand. The more than two meter long reptile, which was similar to modern monitor lizards, must have been able to move on land and in the water, as suggested by the considerable number of fossil finds – totalling about 120 with about six specimens of complete skeletons. The Protorosaurus was also characterised by a long tail region and short front limbs.

Protorosaurus speneri, Reconstruction after www.reptileevolution. com.

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1

2 3

4 5

Protorosaurus speneri. 1. A complete specimen from Ibbenb端ren (LWL-Museum of Natural History in M端nster). 2. Another almost completely preserved skeleton from Gl端cksbrunn (Natural History Museum Vienna). 3. The best preserved complete skull, Richelsdorf (Museum of Natural History in Kassel). 4. Part of an isolated upper jaw (Geological-Palaeontological Institute Halle). 5. Ribs and parts the front extremities, Eisleben (Coll. S. Brandt).

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The Preparation of Fossils from Permian The process of exposing petrified fossils is one of the most complicated, but also most important tasks next to actually finding fossils. In general, fossils - in addition to the skeleton, this also includes any imprints of soft tissue present such as skin, organs, or stomach contents - are completely or partially embedded in rock and must first be exposed using a series of processing steps that serve to remove the surrounding rock.

For optimal and proper preparation, special tools such as air scribes, microsand blasters, and optical instruments like dental loupes and microscopes,

are needed. Extensive knowledge of their function and handling is absolutely necessary. In many cases, paper-thin layers of epoxy resins and fiberglass matting need to be applied to stabilise the rock.

The long-term preservation of plants has proven to be even more difficult since the fossil substance is extremely fine, generally in the range of one tenth or even one hundredth of a millimetre. Even experienced preparers require a good eye and a calm hand in particular, as well as a great deal of time.

Such fossils are usually only prepared using fine scribes and needles.

A fossil prepared with such skill and expertise then fulfils the best prerequisites for further scientific study, so that evidence of past life can then be displayed as a unique exhibition piece worth admiring, and can be exhibited anywhere in the world in museums or private collections.

By Stefan Perner

Top: Stefan Perner making the final preparations for the freshwater shark Orthacanthus. Only after finishing the long and arduous preparation process is it possible to see the beauty of this unique exhibition piece. Right: Before this, however, there are many months of hard work with an air scribe and dental loupe. Middle right: Stefan Perner hardening a paper shale plate using epoxy resin and fiberglass matting. Bottom right: The difficult to prepare amphibian Sclerocephalus in paper shale. All photos from the Oregon Institute of Geological Research, Portland.

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Thanks and reflections A book can only provide a snapshot of the current state of knowledge and may be out of date the next day. Although we were able to base this work on numerous new fossil finds from a number of collections, new discoveries can quickly put everything into question again.

Periods of time so far in the past are always tainted by the unknown, and very often by the unapproachable, however, that is no reason not to take a chance with this most interesting period of evolution, the Permian. It is not protagonism or a craving for sensation that sometimes forces us to push away time-honoured theories and replace them with new ones, but the large number of new facts and discoveries that provided us with a clear view through doors that were previously closed

but are now open. It is here that we must begin giving thanks and paying tribute.

Nothing seems more logical than to start at the bottom with the numerous people who have been disenfranchised, gone unrecognised, and very often led a marginal existence. A typical example is Ferruccio Valentini from the Italian town of Tuenno in the Dolomites. An at old age, the herbalist and woodsman set out to find new Permian fossil sites full of magical beauty accessible to people using his extremely deep knowledge of nature. Day after day, even in the cold and snow, he split stone slabs to bring new species and genera into the broad daylight of knowledge. Society and politicians should have thanked him, but instead they hit this person, who always gave and never thought about his own personal gain, with stiff penalties as if business and destruction were more important than meditation on nature and the fascinating powers of observation.

It‘s in our lemming-like nature, however, to only recognise genius when it is much too late.

Every year, thousands, maybe even millions, of scientifically valuable fossils are lost through the indiscriminate destruction of nature, and anyone who helps to save even just a few fossils from being ruined should be supported and not condemned.

How many private researchers have entered the field of palaeontology and produced spectacular results? I would like to thank and commemorate Harald Stapf and his father Arnulf at this point on behalf of all the others. The Palaeontological Museum in Nierstein, which they happily The two authors Michael Wachtler and Thomas Perner while studying the variety of literafounded together with friends, ture, the objects recently brought to light from numerous private collections, and the present is one of the most informative flora and fauna for the sake of comparison.

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From the basic idea of freedom of research to understanding

The German Arnulf Stapf collecting fossils near the locality of Niederwรถrresbach in the Palatinate. There will be no absolute Credo how or in which manner fossils can be dig out.

The woodsman and collector of wild herbs Ferruccio Valentini from the Non Valley in South Tyrol has made a large number of Permian fossil sites accessible to the public. Silvio Brandt from Halle combines the searcher and scientist into one person in a way like no other.

Fossils collected by Alfred Arnhardt in the Thuringian Forest and studied by the Swedish researcher Rudolf Florin. The complete collection is now in the Museum of Natural History in Schleusingen. On the right is a collection from the Zechstein belonging to the Senckenberg collections of the Museum of Natural History in Dresden.

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From understanding to ideas for all human beings

Knowledge of the world of the past has change considerably over the course of the last decades. On the left is the Museum of Natural History in Sobernheim, and on the right is a reconstruction of a Lower Permian landscape in the Museum of Natural History in Schleusingen

A Permian conifer is depicted on a postage stamp. In the storeroom of the Museum of Natural History in Berlin, the results of several centuries of collecting and research are available for further study.

Knowledge should be passed down from generation to generation. Professor Rudolf Daber, formerly the Director of the Museum of Natural History in Berlin, and author Michael Wachtler. The author Thomas Perner introduces young people to the fascinating world of the origins of life.

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museums in existence when it comes to this often misunderstood period, the Permian. As private researchers, they are not alone. With a willingness to help and background knowledge, the Upper Permian specialist Silvio Brandt stood by our side with advice and assistance. Michael Klopschar, Günther Loth, Norbert Hesse, René Kindlimann and Martin Dammann readily opened the archive they have filled over the course of their lifetimes, and we were amazed at how many “hidden” treasures could be found in private collections and outside of large museums.

Just as we bow to these scholars in reverence, we bow simultaneously and full of reverence to those who decided at a young age to dedicate their careers to studying, and have tried throughout their lives at museums to make this far away Permian period accessible to a larger community, and those who received the finds at their universities and research institutes and published papers about them.

In addition to many others, we would like to point out Barbara Mohr and Stephan Schultka from the Museum of Natural History in Berlin and Ralf Werneburg, the Director of the Natural History Museum in the Bertholdsburg Castle in Schleusingen, which is one of the best museums about the Permian in the world. We would also like to thank Norbert Hauschke from the Martin Luther University in Halle, Ronny Rößler, the director of the Museum of Natural History in Chemnitz, Wolfgang Munk, Natural History Museum Karlsruhe, Köbi Siber, Saurier-Museum Aathal, Paolo Schirolli from the Museum of Natural Sciences, Brescia and the Natural History Museum of the Palatinate in Bad Dürkheim under the direction of Reinhold Flösser.

Two experts should be noted individually, however: Professor Rudolf Daber, who has headed the Museum of Natural History in Berlin for many years, and Professor Giuseppe Cassinis, the Permian pioneer from the University of Pavia. They have not only gained international recognition as scientists, but we must also hold their philosophical studies, their capacity for empathy, and especially their intellectual noblesse in high regard.

Throughout their entire careers, they felt themselves subject to a single noble idea: research for

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humanity. Where others battled with petty animosity, they used all their energy to expand our knowledge of this world. They remained unrelenting to pedantic regimes and were willing to accept penalties and sanctions without ever allowing themselves to be broken. We bow down before such luminous figures and present them as examples to be followed.

We would also like to thank Gotlind Blechschmidt.

As a tireless fighter for all issues relating to nature and environmental protection, we would not have been able to find a more accomplished person.

We are concerned about the attempts made today to pack research and the acquisition of knowledge in a box dominated by small-minded, national, private, and territorial interests instead of viewing it as a public good. During our research, we also unfortunately encountered closed doors, although we actually wanted nothing more than to make the beauty of these former eras accessible to all of humanity. Due to our research, we have been sentenced to prison as if the courts and state institutions had the power to rule over the right of humans to acquire knowledge. While people have fought for racial and gender equality over the past several decades, there is still an urgent need for improvement in this respect. It would appear that these institutions are more rigid than a century ago. It‘s not the more powerful who should have the right to conduct research, but anyone who is willing to do so in a peaceful, competitive manner. We have an obligation to keep the doors to the numerous mysteries and unsolved puzzles regarding our evolution open, from its very beginning up to the present, for upcoming generations and especially for the youth of today. The study of nature lies in our future and not in its destruction through the pursuit of superficial human pleasures.

Contributing authors: Michael Wachtler wrote the majority of the book, including the entire section on the evolution of the plant world. Unless otherwise specified, all drawings and photos are his. Thomas Perner wrote some parts of the chapter on the evolution of the fish and terrestrial vertebrates. He also conducted research.

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Index Abies 108 Abies alba 108 109 Abies bracteata 108 Acanthodes 13 134 Acanthodes bronni 132, 133, 134 Acanthodes sp. ÂŤ Pfalz Âť 134 Acanthostega 162 163 Acentrophorus 146, 151 Acentrophorus glaphyrus 159 Acentrophorus varians 159 Acrolepis 157 Acrolepis segwickii 157, 158 Aeduella 141 169 Aeduella blainviellei 143 Alethopteris 11, 55 Amblypterus 141 Ambystoma tigrinum 164 Amphisauropus 72 180 Amphisauropus latus 188, 189, 190 Angiopteris 42, 48 Angiopteris angustfolia 46 Annularia carinata 35 Annularia spiceta 35 Annularia spinulosa 33, 35 Annularia stellata 11 Anthracosaurus 10 Apateon 61, 140, 162, 168 Apateon caducus 166, 167, 168 Apateon dracyiensis 166, 168 Apateon flagrifera 168 Apateon pedestris 167, 168 Araucaria araucana 84, 99, 106 Araucaria bidwillii 95 Araucaria heterophylla 96, 99 Araucaria rulei 95 Archegosaurus 176 Archegosaurus decheni 176 Archegosaurus latirostris 176 Archiulus 129 Arnhardtia scheibei 62 Arnulfia stapfii 129 Arthopleura armata 130 Arthropitys bistriata 31 Asterophyllites dumasii 32 35 Asterophyllites longifolius 35 Asteroteca sternbergi 48 Asteroxylon elberfeldense 94 Autunia conferta 56, 57, 58, 59 , 60, 62 Autunia dammannii 62 Autunia naumannii 62 Baiera 14, 123 Baiera digitata 122, 123, 125 Baiera perneri 123, 124 Baiera pohli 26, 123, 124 Barthelopteris 55 Batrachichnus salamandroides 188 Batroptetes 186, 187 Bjuvia 14, 24, 70, 74, 76 Bjuvia multinervis 76 Bjuvia tridentina 26, 68, 69, 74 Bjuvia wachtleri 26, 74 ,75, 76 Bowenia 66, 67

Bohemiacanthus 138 Branchiosaurus 174 Calamites 10, 25, 31, 32, 35, 37, 38 Calamites gigas 35 Calamites multiramis 33, 35 Calamites wachtleri 34, 35 Calamostachys dumasii 32, 35 Calamostachys tuberculata 33, 35 Callipteridium gigas 55 Callipteridium pteridium 55 Carchahinus melanopterus 131 Cassinisia ambrosii 26 Chelichnus tatzelwurmi 199 Cheliderpeton 13, 171, 176 Cheliderpeton latirostre 176 Cheliderpeton vranyi 175, 176 Claudiosaurus germaini 197 Cochleosaurus bohemicus 177 Coelacanthus 147, 160 Coelacanthus granulatus 160 Coelurosauravus 24, 192, 194 Coelurosauravus jaekeli 192, 193 Conchopoma 13, 144 Conchopoma gadiforme 144 Cordaianthus 82 Cordaites principalis 82, 83 Cryptobranchus alleganiensis 172 Cyathea arborescens 43 Cycadopites 62 Cycadospadix 70 Cycas 27, 66, 67, 68, 70, 76 Cycas revoluta 66, 67, 74 Deltavjatia 197 Diadectes 179, 182, 189 Diadectes absitus 182, 183 Dicksoniites pluckenetii 53, 54, 55 Dicksonites 53 Dicranophyllum hallei 114, 115 Dimetrodon 179, 185 Dimetrodon teutonis 184 Dimetropus 185, 191 Dimetropus leisnarianus 191 Dioonitocarpidium 70, 80 Discosauriscus austriacus 181 Discosauriscus pulcherrimus 181 Dorypterus 147 Dorypterus hoffmanni 154, 157 Draco volans 192 Dromopus didactylus 190, 191 Dromopus lacertoides 189, 190, 191 Dromopus tridactylus 187 Dvinosaurus 196 Dycinodotipus isp. 198 Ectosteorhachis sp. 140, 145 Eliginerpeton 163 Elonichthys eupterygius 141 Encephalartos ferox 72 Equisetites 14, 27, 31, 37, 39 Equisetites arenauceus 24, 31, 40 Equisetites mougeotii 24, 31, 40 Equisetites siberi 26, 31, 40 Equisetum 37

Equisetum arvense 39 Equisetum hyemale 31 Equisetum schaffneri 31 Equisetum telmateia 30 Ernestiodendron (filiciformis) 86 Ernestiodendron piniformis 89 Eryops 185 Esterella (gracilis) 122 Esterella 123 Eudibamus cursoris 184 Eupropoos bifidus 10 Eurysomus 147 Eurysomus macrurus 154, 157 Eusthenopteron 162 Filicites pluckenetii 53 Ginkgo adiantiodes 123 Ginkgo biloba 123, 125 Gomphostrobus bifidus 90, 104 Gracilisarus ottoi 192, 193 Haikouella 131 Haptodus 185 Himantura Fai 131 Hopleacanthus 146 Hopleacanthus richelsdorfensis 149 Hurumia lingulata 60, 61 Hynerpeton 163 Ichniotherium 183, 198 Ichniotherium accordii 198 Ichniotherium cottae 182, 183, 188, 189 Ichniotherium sphaerodactylum 183 Ichthyosaura alpestris 167 Ichthyostega 162, 163 Inostrancevia 196 Isoetes 27 Janassa bituminosa 147, 148 Janassa korni 147, 148 Janusichnus bifrons 199 Juniperus communis 126 Kotlassia 196 Ladinia 70, 80 Larix 108 Latimeria 144, 160, 162 Latimeria chalumnael 145 Lebachia (Walchia) hypnoides 90, 91 Lepidodendron 8, 10, 28, 29 Lepidopteris 56 Lepidopteris martinsii 26, 65 Lepidopteris meyeri 26, 64 Lepidosiren 162 Lepidozamia peroffskyana 67 Letoverpeton 182 Limulus polyphemus 10 Lissotriton montandoni 167 Longisquama 194 Lycopia dezanchei 24, 28 Lycopodiolithes 86, 87 Meganeura 10 Majonica 24, 26, 94, 114, 116 Majonica alpina 26, 112, 113, 114 Marattia 42 Melanerpeton 140, 167, 188 Melanerpeton arnhardtii 168

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Melanerpeton eisfeldi 168 Melanerpeton humbergense 168 Menaspis 146 Menaspis armata 150 Metaxygnathus 163 Microbrachis pelicani 186 Micromelerpeton 162, 169, 174, 181 Micromelerpeton credneri 165, 169 170 Nemejcopteris feminaeformis 48 Neocalamites 14, 31, 37, 39, 40 Neocalamites behnkeae 26, 38, 40 Neocalamites tregiovensis 26, 37, 39, 40 Neoceratodus fosteri 144 Neuropteris 55 Nilssonia 14 24 71 72 76 Nilssonia abnormis 69 Nilssonia brandtii 26, 73, 76 Nilssonia perneri 26, 71, 74 Obruchevichthys 163 Odontopteris latifrons 55 Odontopteris lingulata 52, 55 Odontopteris schlotheimii 55 Oestocephalus 164 Onchiodon 140, 176, 177 Onchiodon labyrinthicus 176 Onchiodon langenhani 177 Orabates pabsti 183 Orthacanthus 135, 138 Orthacanthus pollichiae 137 Orthacanthus senckenbergianus 135, 136, 137 Ortiseia 24, 26, 98, 106, 114 Ortiseia daberi 105, 106 Ortiseia leonardii 106, 107 Ortiseia zanettii 107 Osmunda 42, 43 Osteolepis 160 Otovicia hypnoides 25, 97, 104,105 Pachypes (dolomiticus) 24, 197, 198, 199 Palaeoniscum freieslebeni 24, 146, 147, 151, 152, 153, 160 Panderichthys 162 Pantelosaurus (haptodus) saxonicus 185 Pantelosaurus longicaulis 185 Paramblypterus 13, 140, 141, 143, 169, 173, 174 Paramblypterus dovernoyi 142 Paramblypterus gelberti 142 Parasaurus geinitzi 196, 197 Pareiasaurus 196, 197, 199 Pecopteris 11 Pelourdea 80 Peltaspermum 56, 59, 61, 62, 65 Peltaspermum martinsii 65 Perneria 25, 94, 109, 118, 123 Perneria thomsonii 92, 93, 94 Phasmatocycas 70, 78 Phasmatocycas bridwellii 78 Phyllobates 130 Picea 108, 109 Picea abies 85, 109 Pinus cembra 85, 117, 122 Pinus edulis 117 Pinus engelmannii 85 Pinus halepensis 117 Pinus lambertiana 117

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Pinus longaeva 85 Pinus monophylla 117 Pinus mugo 85, 117, 122 Pinus sylvestris 117 Pitisana (marattia) purpurescens 42 Platysomus 146, 147, 157 Platysomus gibbosus 155 Platysomus striatus 155, 157 Pleurojulus steuri 130 Pleuromeia 24, 28 Plicatodus 135 Primocycas chinensis 70 Protorosaurus speneri 24, 192, 194, 195 Psaronius 43, 44 Pseudoctenis 71, 72 Pseudoctenis samchokense 70 Pseudomariopteris 53 Pseudomariopteris busquetii 55 Pseudoptilophyllum 70 Pseudotsuga bracteata 108, 116 Pseudotsuga menziesii 108 Pseudovoltzia liebeana 24, 98, 102 103, 105 Psygmophyllum 125 Pterispermostrobus (kontheri) 62 Pteroplax 10 Pygopterus 146, 147 Pygopterus humdoldti 153 Remia pinnatifida 48, 49 Reticulolepis exsculpta 156, 157 Rhabdolepis macropterus 143 Rhachiphyllum 56, 62 Rhachiphyllum hauptmannii 60, 63 Rhachiphyllum schenkii 60, 63 Rhynchosauroides pallinii 198 Rhynchosauroides palmatus 198, 199 Salamandra atra 165 Salamandra salamandra 167 Samaropsis ulmiformis 115, 116 Stangeria 66 Stangeria eriopus 74 Sawdonia 92 Sawdonia spinossima 94 Sclerocephalus haueseri 169, 171, 172, 174 Sclerocephalus jogischneideri 174 Sclerocephalus, 13, 140, 173, 174, 176, 200 Scolecopteris 43, 46 Scolecopteris arborescens 45 Scolecopteris candolleana 47, 48 Scolecopteris cyathea 45 Scolecopteris densifolia 45 Scolecopteris elegans 46 Scolecopteris longifolium 38 Scolecopteris lothii 45, 47, 48 Scolecopteris polymorpha 45, 48 Scolecopteris sternbergii 47 Scolecopteris thonii 38, 39 Scutosaurus karpinski 196, 197 Scythophyllum 65 Selaginella 27, 28 Selaginellites zollwegii 28 Sequoia sempervirens 85 Sequoiadendron giganteum 85 Seymouria 25, 180, 182, 188 Seymouria sanjuanensis 180

Seymourina 98, 105, 114, 173, 179, 180 Seymourina geinitzi 101, 102 Seymourina niederhauseni 102 Seymourina viallii 26, 101, 102 Sigillaria 8, 10, 14 Sigillaria brardii 28, 29 Sinerpeton 163 Sobernheimia jonkeri 78, 80 Sphenophyllostachys 38 Sphenophyllum 37, 38 Sphenophyllum angustifolium 36, 38 Sphenophyllum oblongifolium 36, 38 Sphenophyllum thonii 36 Sphenopteris 14, 24, 52 Sphenopteris dichotoma 26, 52 Sphenopteris suessi 26, 51, 52 Stangeria eriopus 67, 70 Swillingtonia denticulata 92 Taeniopteris, 14 Taeniopteris coriacea 68 Taeniopteris eckardtii 79, 80 Taeniopteris jejunata 78 Taeniopteris multinervis 68, 69 74 Taeniopteris polymorpha 68 Tambachia 179 Taxodium mucronatum 85 Tethydostrobus 70, 72 Pernerina pasubi 26, 80, 81 Thuringiostrobus meyenii 98 Thuringiostrobus florinii 98 Todea 50 Todites muelleri 50 Tridentinosaurus antiquus 187, 190, 191 Triodus 135 Triodus sessilis 139 Triturus stiratus 167 Tubicaulis berthieri 44 Tulerpeton 163 Ullmannia 24, 126, 194 Ullmannia frumentaria 126, 127, 128, Uronectes fimbratus 129 Valentinia 14, 26, 94, 114, 122, 123 Valentinia angelellii 118, 120, 122 Valentinia cassinisi 118, 121, 122 Valentinia wachtleri 117, 118, 119 Ventastega 163 Voltzia 24, 98 Voltzia agordica 102 Voltzia dolomitica 102 Voltzia heaxagona 105 Wachtlerina 14, 25, 94, 122 Wachtlerina bracteata 109, 110, 112 Wachtlerina suessi 112, 111 Wachtleropteris valentinii 14, 26 76 77 Walchia 14, 86, 89, 90, 91, 98, 102, 106 Walchia (Lebachia) piniformis 98 Walchianthus 90 Walchiostrobus gothani 90 Weigeltisaurus jaekeli 192 Westrichus kraetschmeri 134 Wodnika 146, 147, 149 Wodnika striatula 149 Xenacanthus 135, 138, 139 Xenacanthus meisenheimenis 138 Zamia 27, 68, 76 Zamia floridana 74

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Veröff. Naturhist. Mus. Schleus., Schleusingen.

Val Trompia. Neue Jahrbuch für Mineralogie Geologie Paläontologie: 456-461

BARTHEL, M. & WEISS. H.J. (1997): Xeromophe Baumfarne im Rotliegend Sachsens, Veröffentl. Mus. für Naturk. Chemnitz 20 S.45-56 ,12 Abb. Chemnitz

GEINITZ, H. Br. & GUTBIER A.von (1843): Die Versteinerungen von Obersachsen und der Lausitz aus der Grauwacke, Steinkohlenformation und Rotliegend Gäa von Sachsen S.63-92

BARTHEL, M.; BETTAG,E. ;NOLL, R. (1998): Dicranophyllum hallei REMY & REMY im oberen Rotliegend Veröffentl. Mus. Für Naturk. Chemnitz 21 S.5-20 , 20 Abb. Chemnitz BARTHEL, M. (2000): Annularia stellata oder Annularia spinulosa. , Veröff. Naturkundemuseum Erfurt, Band 19: S. 37-42, Erfurt.

HAUBOLD, H., SCHAUMBERG, G. (1985). Die Fossilien des Kupferschiefers. Pflanzen- und Tierwelt zu Beginn des Zechsteins – eine Erzlagerstätte und ihre Paläontologie. – Neue Brehm-Bücherei, Bd. 333, 223 S., 139 Abb., 13 Tab., Wittenberg (A. Ziemsen-Verlag). ISSN 0138-1423

BRONGNIART, A. (1828): Histoire des végétaux fossiles -1, 488 S. 166 Taf. Paris.

Haubold, H. (2000): Tetrapodenfährten aus dem Perm – Kenntnisstand und Progress 2000. - Hallesches Jb. Geowiss., B 22: 1-16, 3 Tab., Halle.

CASSINIS, G., NICOSIA, U., PITTAU, P. & RONCHI, A., (2002). Paleontological and radiometric data from the Permian deposits of the central Southern Alps (Italy), and their stratigraphic implications: Association Géologique du Permien, Paris, Mémoire n. 2.

KERP, H.; NOLL, R. & UHL, D. (2007): Vegetationsbilder aus dem saarpfälzischen Permokarbon, Pollicha Sonderv. Nr.10 S.76 - 109 30 Abb.

CASSINIS, G. & RONCHI, A., (2001). Permian chronostratigraphy of the Southern Alps (Italy) – an update. – Contribution to Geology and Palaeontology of Gondwana in honour of Helmut Wopfner, Cologne p. 73-87.

CLEMENT-WESTERHOF, J., (1984). Aspects of Permian Palaeobotany and Palynology. IV. The conifer Ortiseia from the Val Gardena Formation of the Dolomites and the Vicentinian Alps (Italy) with special reference to a revised concept of the Walchiaceae (GOEPPERT) SCHIMPER. - Rev. Palaeobot. Palynol., n. 41, pp. 51-166 CLEMENT-WESTERHOF, J., (1987). Aspects of Permian Palaeobotany and Palynology; VII, The Majonicaceae, a new family of Late Permian conifers. - Rev. Palaeobot. Palynol. 52 (4), pp. 375-402.

CLEMENT-WESTERHOF, J. A., (1988) Morphology and Phylogeny of Palaeozoic conifers, in BECK, C. B. (Hrsg.): Origin and evolution of gymnosperms. Columbia University Press, New York, pp. 298-337.

DABER, R. (1978): Das Große Fossilienbuch 264 S. 98 Fotos Urania Verlag Leipzig-Jena-Berlin DABER, R.& HELMS, J. (1981): Fossile Schätze 231 S. Edition Leipzig ECKENWALDER, J.E. (2009). Conifers of the World: the complete reference. Portland (OR): Timber Press. 720 pp.

FLORIN, R. 1933. Studien über die Cycadales des Mesozoikums (Bennettitales, pp. 12-30). – Kungliga Svenska Vetenskapsakademiens Handlingar 12: 4-134 FLORIN, R. (1938 - 1945): Die Koniferen des Oberkarbons und des unteren Perms. I. - Palaeontographica, FLORIN, R., (1936): Die fossilen Ginkgophyten von Franz-Joseph-Land nebst Erörterungen über vermeintliche Cordaitales mesozoischen Alters. II. Allgemeiner Teil. Palaeontographica 1936, 82(B):1-72.

FLORIN, R., (1964). Über Ortiseia leonardii n. gen. et sp., eine Konifere aus den Grödener Schichten im Alto Adige (Südtirol). - Mem. Geopaleont. Univ. Ferrara, 1(1), pp. 3-11, n. 41, pp. 51-166.

GEINITZ, H. B., (1869). Über Fossile Pflanzenreste aus der Dyas von

KERP, J. H. F., (1988). Aspects of Permian palaeobotany and palynology. X. The West- and Central European species of the genus Autunia KRASSER emend. KERP (Peltaspermaceae) and the form-genus Rhachiphyllum KERP (callipterid foliage). Review of Palaeobotany and Palynology, n. 54, pp. 249-360.

KERP, J. H. F. & FICHTER, J. (1985): Die Makrofloren des saarpfälzischen Rotliegenden(?Oberkarbon - Unter Perm, SW-Deutschland) Mainzer geowiss. Mitt. 14 S.159-286 3 Abb. 42 Taf. Mainz Lohmann, U. & Sachs, S. 2001. Observations on the postcranial morphology, ontogeny and palaeobiology of Sclerocephalus haeuseri (Amphibia: Actinodontidae) from the Lower Permian of southwest Germany. Memoirs of the Queensland Museum 46, 771-781 MÄGDEFRAU, K.(1952): Vegetationsbilder der Vorzeit 24 S. 18 Taf. Gustav Fischer Verlag Jena.

MÄGDEFRAU, K.(1968): Paläobiologie der Pflanzen 549 S. 394 Abb Gustav Fischer Verlag Jena.

NOLL, R. & WILDE, V. (2002) : Koniferen aus den Uplands Permische Kieselhölzer aus der Mitte Deutschlands, Geheimnisse Versteinerte Pflanzen S. 88 - 103 35 Abb. RÖßLER, R. (2004): Fundmitteilung Das größte anatomisch erhaltene Schachtelhalmgewächs der Erdgeschichte - ein Calamit aus dem Perm von Chemnitz, Veröffentl. Mus. für Naturk. Chemnitz 27 S. 127129, 6 Abb. Chemnitz ROČEK, Zbyněk, (1998): Stammesgeschichte und Evolution. In Hofrichter, Robert. Amphibien. Evolution, Anatomie, Physiologie, Ökologie und Verbreitung, Verhalten, Bedrohung und Schutz. Augsburg: Natur Buch Verlag. 1998. S. 20-31.

RÖSSLER, R. (2001): Der Versteinerte Wald von Chemnitz 248 S. 53, 6 Abb. Museum für Naturkunde Chemnitz RÖSSLER, R. (2001): Studien zur Lebensweise und Fossilisation des Baumfarnes Psaronius im Versteinerten Wald von Chemnitz( Unterperm, Deutschland) Hallesches Jahrb. Geowiss. B.23 S.9-33 5 Taf. Halle SCHINDLER Th., ULRICH H., HEIDTKE, J., (Hrsg.) (2007): Kohlesümpfe,

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Seen und Halbwüsten. Pollichia Sonderveröffentlichung, 10: 4-37, Neustadt an der Weinstraße.

SCHWEITZER, H.J., (1994): Die ältesten Pflanzengesellschaften Deutschlands in Koenigswald & MEYER W. Erdgeschichte

Publications about Permian from the same authors

STEINER W. (Hrsg (1993: Europa in der Urzeit: Die erdgeschichtliche Entwicklung unseres Kontinents von der Urzeit bis heute. MosaikVerlag, München Schlotheim, E.F. von, (1820): Die Petrefactenkunde auf ihrem jetzigen Standpunkte durch die Beschreibung seiner Sammlung versteinerter und fossiler Überreste des Thier- und Pflanzenreichs der Vorwelt erläutert 1-457 [P. Wagner/P. Wagner/P. Wagner].

PERNER T. & WACHTLER, M., (2014): Permian Fossil Plants from Europe and their Evolution. The Niederhausen- and the Tregiovo-Flora, Dolomythos Museum, Innichen, South Tyrol, Italy, Oregon Institute of Geological Research, Portland, OR, (USA), ISBN 978-88-908815-1-0 POORT, R. J. & KERP, J. H. F., (1990): Aspects of Permian palaeobotany and palynology. XI. On the recognition of true peltasperms in the Upper Permian of West and Central Europe and a reclassification of the species formerly assigned to Peltaspermum HARRIS. Review of Palaeobotany and Palynology, n. 63, pp. 197-225.

POTT, C., McLOUGHLIN, S. & LINDSTROEM, A., (2010). Late Palaeozoic foliage from China displays affinities to Cycadales rather than to Bennettitales necessitating a re-evaluation of the Palaeozoic Pterophyllum species. Acta Palaeontologica Polonica, n. 55, pp.157-168.

REMY, W., REMY, R., 1975. Beiträge zur Kenntnis des Morpho-Genus Taeniopteris Brongniart. Argumenta Palaeobotanica vol. 4, pp. 31–37.

REMY, W. & REMY, R., (1978). Die Flora des Perms im Trompia-Tal und die Grenze Saxon/Thuring in den Alpen. Argumenta Palaeobotanica, Münster, vol. 5, pp. 57-90.

Volume 1

SCHUSTER, J. 1932. Cycadaceae. In: H. G. A. Engler, ed. 1900-1953. Das Pflanzenreich. Berlin. Vol. 99[IV,1], pp. l-168.

SCHWEITZER, H.-J., (1960). Die Sphenopteriden des Zechsteins.

Senckenbergiana Lethaea, 41, pp. 37-57, Frankfurt.

SCHWEITZER, H.-J., (1984). The land flora of the German and England Zechstein sequences. [In HARWOOD, G.M. & SMITH, D.B., (eds.). The English Zechstein and related topics.]. Geological Society Special Publication 22, pp. 31-54.

SUESS, E., (1869). Über das Rotliegende im Val Trompia. Sitzg. d. Akad.

Wiss. Wien. Math.-Naturwiss. Klasse, 1. Abt., vol. 59.

TAYLOR, T.N., TAYLOR, E.L., KRINGS M. (2009). Paleobotany. The Biology and Evolution of Fossil Plants. Burlington MA, London, San Diego CA, New York NY, Elsevier/Academic Press Inc., xxi + 1230 pp.

PERMIAN FOSSIL PLANTS FROM EUROPE AND THEIR EVOLUTION Rotliegend and Zechstein Floras from Germany and the Dolomites by

MICHAEL WACHTLER and THOMAS PERNER

WACHTLER, M. 2012. The Genesis of Plants. Preliminary researches about the Early-Middle Triassic Fossil Floras from the Dolomites. A Compendium. DoloMythos – Innichen. ISBN 978-88-904127 WACHTLER, M. 2014. The latest Artinskian/Kungurian (Early Permian) Flora from Tregiovo-Le Fraine in the Val di Non (Trentino, Northern Italy) - Additional and revised edition WEISS, H.J. (2002): Beobachtungen zur Variabilität der Synangien des Madenfarns, Veröffentl. Mus. für Naturk. Chemnitz 25, S.57 - 62, 8 Abb. Chemnitz

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DOLOMYTHOS Oregon Institute of Geological Research Volume 2

Volume 2

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