Tambach Formation

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Tambach Formation
Stratigraphic range: Early Permian, Sakmarian
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Bromacker 02.jpg
Traco quarry at Bromacker
Type Geological formation
Underlies Eisenach Formation
Overlies Rotterode Formation
Lithology
Primary Conglomerate, sandstone, mudstone
Location
Coordinates 50.8097° N, 10.6189° E
RegionFlag of Thuringia.svg  Thuringia
CountryFlag of Germany.svg  Germany
Type section
Named for Tambach-Dietharz village

The Tambach Formation is an Early Permian-age geologic formation in central Germany. It consists of red to brown-colored sedimentary rocks (red beds) such as conglomerate, sandstone, and mudstone, and is the oldest portion of the Upper Rotliegend within the Thuringian Forest Basin. [1] [2]

Contents

The overall geology records a history with three distinct stages of sedimentation within a mountainous environment. First, tectonic activity forms a basin (the Tambach Basin) dominated by high-energy debris flows, sheetfloods, and braided rivers. These incise underlying rhyolitic rock, depositing a coarse conglomerate known as the Bielstein Conglomerate. Second, calmer conditions allow the basin to widen, and the conglomerate is marginalized by finer sediments which were previously only common at the center of the basin, such as the characteristic Tambach Sandstone. These finer sediments were deposited through repeated sequences of flooding, followed by calm water, followed by exposure to air. The overall climate would have been similar to modern tropical savannas, with hot and dry periods broken up by heavy rainfall, likely multiple times in a year. The third stage involves a return of tectonic conditions, this time inducing wide deposits of polymictic (heterogenous) conglomerate known as the Finsterbergen conglomerate. [1] [3] [2]

The Tambach Formation also includes one of the most important Permian fossil localities in Europe: the Bromacker locality. This former sandstone quarry and surrounding sites preserves several different facies types, with different fossil components. Thick sandstone channel fills in the lower section at Bromacker are overlain with mudstone from ephemeral lakes. A diverse assemblage of trace fossils such as footprints are imprinted onto the mudstone drapings. The upper section of Bromacker contains siltstone deposited through sheetfloods, in which well-preserved articulated skeletons of terrestrial tetrapods have been discovered. These include early amphibians like Rotaryus and Tambaroter, and early reptiles like Eudibamus and Thuringothyris . Unlike most fossil-preserving Permian red beds, aquatic vertebrates are absent at Bromacker while carnivorous synapsids (like Dimetrodon ) are rare and herbivorous diadectids are abundant. Plant and arthropod fossils have also been found in shales at Bromacker. [1] [4]

History

From 1887 to 2010, the Ducal (Herzogliches) Museum in Gotha housed most of the Tambach Formation's most famous fossils. Thuringia Gotha asv2020-07 img28 Herzogliches Museum.jpg
From 1887 to 2010, the Ducal (Herzogliches) Museum in Gotha housed most of the Tambach Formation's most famous fossils.

Starting in the 1840s, five-toed footprints had been described from various sandstone quarries and roadcuts in the Gotha district of Germany. The stratigraphy of rocks near Tambach-Dietharz was mapped out in 1876, [5] and named as the "Tambacher Schichten" (Tambach strata) in 1895, although at the time it was believed to include several additional rock layers (now termed the Elgersberg and Eisenach Formations) which have since then been separated from it. [2] in 1887 the first fossilized footprints from Bromacker were discovered by a local fossil collector named Heinrich Friedrich Schäfer. The find was donated to the Ducal Museum in the city of Gotha, after which it was independently reported on by various German paleontologists who had received photographic evidence. [6] Wilhelm Pabst, the curator of the Ducal Museum's natural history department, collected and described 140 sandstone slabs from the Tambach Formation from 1890 until his death in 1908. The collection was rediscovered in the 1950s, and subsequently restudied by GDR paleontologists such as Hermann Schmidt, Arno Hermann Müller, [7] and Hartmut Haubold. [8]

Fossilized tetrapod bones were discovered in the upper beds of Bromacker by Thomas Martens in 1974, prompting further attention from Gotha paleontologists. These include Harald Lutzner, who formally delineated the Tambach Formation as a sequence including two conglomerate layers separated by a sandstone layer. [2] In the coming years, research contacts were made with Western paleontologists such as Jürgen Boy (University of Mainz) and David Berman (Carnegie Museum of Natural History). This allowed excavation to ramp up and the Tambach Formation to achieve worldwide fame. A 1993 German-American joint expedition recovered articulated tetrapod fossils, and trace fossil collecting was resumed after more than 80 years thanks to the excavation of a new sandstone quarry at Bromacker in 1995. The first Tambach body fossils outside of Bromacker were discovered in 2008, in a construction site in downtown Tambach-Dietharz. In 2010, the exhibits comprising Gotha's Museum of Nature began the process of being moved from the Ducal Museum (which was being converted to an art museum) to the Friedenstein Castle. Due to funding issues, collecting from Bromacker has been limited and the Tambach collection is being archived at the historic Perthesforum complex prior to the construction of a new Permian exhibit. [8]

Geology

Map of the modern Thuringian Forest Basin (in German). TB = "Tambacher Becken" (Tambach Basin) Thuringian Forest Geology.png
Map of the modern Thuringian Forest Basin (in German). TB = "Tambacher Becken" (Tambach Basin)

The Tambach Formation mainly lies within a basin currently occupied by the Thuringian Forest, and it is stratigraphically younger than the Rotterode Formation and older than the Eisenach Formation. It is the oldest part of the Thuringian Forest Basin's Upper Rotliegend succession, a name referring to a sequence of purely sedimentary rocks in the Lower Permian of Germany. The sediments of the Tambach Formation were deposited in a small Permian graben (termed the Tambach Basin), which was oriented in a northeast to southwest direction and incised into the igneous and sedimentary rocks of the Rotterode Formation. The Tambach Basin would have about 250 square km during the Permian, though modern outcrops only occupy about 50 square km, not counting the northeastern portion of the basin which has had its deposits erased by later geological processes. [1] [2]

Traditionally, the Tambach Formation is considered to be divided into three discrete layers: a lower and an upper conglomerate layer (Bielstein and Finsterbergen Konglomerate, respectively), separated by a somewhat narrower layer of finer sediments such as sandstone, the Tambach-Sandstein (Tambach sandstone) member. [9] [3] [2] However, the borders between these layers are often imprecise, and some geologists have offered a more complex depositional picture with three stratigraphic stages defined by changes in basin-wide sedimentology, rather than specific rock types. [1]

The lowest stage (stage I) experienced a period of high tectonic activity (part of the Variscan orogeny) to the southeast, the Oberhof uplift. The formation of the basin among this tectonic backdrop initially led to powerful debris flows and sheet floods, and then active braided rivers flowing along its edge with lower-energy rivers, floodplains, and lakes at its center. The coarse (cobble/boulder-scale) and rhyolitic Bielstein conglomerate was deposited among the high-energy marginal environments while early portions of the Tambach sandstone were deposited further away from the basin's edge. The most prominent flow direction in the well-preserved eastern part of the basin is northwest, towards the center of the basin. [1] [2] [10]

Decreasing tectonic activity in the middle stage (stage II) leads to increased erosion, lowering the relief along the edge of the basin. As a result, the braided rivers at the edge slowed down, lowering the size of the clasts to cobble/pebble conglomerate. The center of the basin graded into small streams and marshes, depositing sandstone, siltstone, shale and mudstone in fossiliferous red beds. Although the Tambach Basin may have been hydrologically isolated during this period, with its waterways draining internally, [1] some paleontologists instead consider its waters to flow into another basin in the northeast, which was not preserved. [2] The last stage (stage III) experienced a return of tectonic activity (the Ruhla crystalline uplift) to the northwest, although the relief was still fair flat. Ruhla-sourced alluvial fans and braidplains became more common, gradually allowing mineral-rich and polymictic pebble-sized conglomerate to build up and expand into the center of the basin, forming the Finsterbergen conglomerate. [1] [2] [10]

Bromacker

A sandstone slab from the lower beds of Bromacker, showing cracked mudstone drapings. Bromacker 01.JPG
A sandstone slab from the lower beds of Bromacker, showing cracked mudstone drapings.

The most famous and fossiliferous locality within the Tambach Formation is the Bromacker locality, a cluster of small abandoned quarries near the town of Tambach-Dietharz. Strata exposed to the surface at Bromacker corresponds to the center of the Tambach Basin, during the time of the upper portion of stage I and the lower portion of stage II. Stage I sediments at Bromacker are termed the "lower beds" [1] or "Bromacker sandstone" [3] and stage II sediments are the "upper beds" [1] or "Bromacker horizon". [3]

The lower beds are dominated by thick sheets of fine-grained sandstone, often with cross-bedding indicating that the paleocurrent was oriented towards the northeast. These sandstone layers are typically blanketed by homogeneous mudstone, which sometimes preserved mudcracks, plant fossils, invertebrate burrows, and tetrapod footprints. The thick sandstone layers (and their mudstone drapings) are not continuous, interrupted by a succession of finer and darker micaceous sediments such as shale, siltstone, and (rarely) very fine sandstone. This reconstructs a sequence of repeated flooding events, involving strong, straight rivers eroding channels through the silty floodplain at the center of the basin, leaving behind sandy channel fills (sandstone) and fine-grained overbank deposits (shale and other sediments). The mudstone drapings can be explained as precipitate from extensive ephemeral lakes that evaporated in the weeks following the floods, after which they became mudflats. Many of the mudcrack fragments were ripped up by the next flood, being incorporated into the subsequent sandstone sheets as intraclasts. [1]

A complete and fully articulated skeleton of Orobates pabsti from the upper beds of Bromacker. Orobates.png
A complete and fully articulated skeleton of Orobates pabsti from the upper beds of Bromacker.

The narrower upper beds also represent alternating flooding and still water conditions, albeit with finer clasts and lower topography. The sandstone channel fills are mostly replaced by homogeneous layers of red siltstone, which were prone to breaking along sharp edges. This siltstone contained mudcrack fragments, calcite-encased roots, and well-preserved partial or articulated skeletons of terrestrial tetrapods. These sediments indicate that reduced relief in stage II of the Tambach Formation had made sheetfloods the dominant erosional force, rather than individual channels. The lowest sheetflood deposit was particularly rich in diadectid fossils. The fine sediment successions were also altered, becoming dominated by finely laminated (and only occasionally micaceous) shale in which conchostracans and arthropod remains were fossilized. These indicate a transition to more permanent lakes and broad floodplain conditions at the center of the Tambach Basin, rather than the ephemeral fluvial environment of the lower beds. [1]

An extensive older sequence, the "Tambach-Wechsellagerung" (Tambach interbedding) was discovered in 2004 through borehole data. This sequence somewhat resembled the lower beds, with alternating fine micaceous deposits and thick sandstone sheets filled with a breccia of mudstone intraclasts. However, the sandstone layers had no evidence of cross-bedding, and the mud drapings responsible for most of the Tambach trace fossils were absent as well. Rare fragments of vertebrate fossils were present, along with calcite structures. [3]

Age

Uranium-Lead dating is not possible for the Tambach Formation, which lacks fresh volcanic rocks. The similar [2] Elgersburg Formation to the southeast contains rhyolite dated to 274 ± 4.9 million years ago. [11] However, it is unclear whether the strata at Elgersburg are younger, older, or equivalent in age to the Tambach Formation. [3] Biostratigraphy is more informative but still imprecise. Insect and conchostracan biostratigraphy places it into the Sakmarian-Artinskian Moravamylacris kukalovae [12] and late Artinskian Lioestheria monticula/andreevi [13] biozones, respectively. The only species of tetrapod known to exist in both the Tambach Formation and North American faunas is Seymouria sanjuanensis , which persisted for approximately 15 million years between the Asselian and the early Kungurian. Since the species of Dimetrodon present at Tambach is smaller than those present in the red beds of Texas, the Tambach Formation was likely older than those formations. [14] The Tambach Formation was placed within the Seymouran LVF (Land Vertebrate Faunachron) of Lucas (2006), a biozone which was estimated to include the Artinskian-Kungurian boundary. [15] Combining both invertebrate and tetrapod biostratigraphy, the age of the Tambach Formation was considered to be probably Artinskian in age. [12] In a study published in 2022, Menning and colleagues consider the age of the Tambach formation to be probably between 294 and 292 Ma, corresponding to the Sakmarian. [16] This estimate is based primarily on the radiometric age of 295.8 ± 0.4 Ma (late Asselian) of the Rotterode Formation which unconformably underlies the Tambach Formation, and on the estimate that the interval of geologic time not represented between the two formations is less than 2 million years. [16] [17] In addition, comparison of the footprint assemblage of the Tambach Formation with radiometrically dated Permian footprint assemblages from France and Italy also suggests a Sakmarian age. [16] [18]

Climate

Los Llanos in present-day South America, which may have had a similar climate to the Tambach Formation. NP Llanos27 lo (5853938480).jpg
Los Llanos in present-day South America, which may have had a similar climate to the Tambach Formation.

The sand and silt-rich portions of the Tambach Formation were likely deposited in a warm climate with both hot, dry parts of the year and periodic heavy rainfall events. The dry times were severe enough to evaporate the Tambach basin's flood-induced ephemeral lakes within a matter of days, restricting the ability of a permanent aquatic fauna to colonize the basin. However, most plant root fossils are horizontally (rather than vertically) oriented, indicating that the climate was generally humid enough that native plants would not need to evolve deep roots or other xerophytic adaptations. Because of this, the Tambach Formation would probably fall under the modern tropical savanna climate, despite its lack of grass. Modern climatic equivalents include the northern African savanna and the Llanos of Venezuela and Colombia. [1] However, from the sediment geochemistry of the Tambach Sandstone Member, Scholze and Pint proposed a mean annual temperature of only 10.9 to 15°C (12.7°C on average). [19] There is evidence that sub-zero temperatures may have occurred on some nights during the dry season, likely as a result of the basin's high elevation. [20] [2] [19] The climate may have been drier during the conglomerate-rich periods of the Tambach Formation. [1] [2]

Paleobiota

Artist's depiction of the Tambach paleoenvironment Tambach Formation Scene.jpg
Artist's depiction of the Tambach paleoenvironment

The ecosystem of the Tambach Formation is unusual for its lack of aquatic animals such as xenacanthid sharks, Eryops , or Diplocaulus , which are otherwise common in Early Permian red beds. This is best explained by its mountainous environment, isolated from the monsoonal lowland floodplains which deposited most of the red beds. In addition, the ephemeral nature of the Tambach Basin's lakes and rivers means that only aquatic animals adapted to such conditions, such as conchostracans, were able to flourish. The Tambach Basin did support a diverse amphibian fauna, but only terrestrially-adapted types including dissorophoids and seymouriamorphs. Large herbivorous tetrapods such as caseids and especially diadectids are the most common body fossils recovered from the formation, while carnivorous synapsids are relatively rare. This is in contrast to North American environments, where fossils of carnivores such as Dimetrodon outnumber herbivore fossils. The environmental conditions of Tambach likely created a food web which was very different from that of the lowlands. The most common plants were tough, drought-adapted types such as conifers, while seed ferns and other lowland plants were much rarer. Fibrous terrestrial plants encouraged colonization of the basin by herbivorous land animals, but the dry climate prevents the development of an aquatic food chain, inhibiting animals such as large species of Dimetrodon, which get a large portion of their food from waterways. [1] [4]

Color key
Taxon Reclassified taxonTaxon falsely reported as presentDubious taxon or junior synonym Ichnotaxon Ootaxon Morphotaxon
Notes
Uncertain or tentative taxa are in small text; crossed out taxa are discredited.

Flora

TaxonMemberMaterialNotesImages
Calamites gigas Tambach-Sandstein MemberSome specimens. Horsetails. [21]
Callipteris sp.Tambach-Sandstein MemberVery rare.A fern. [3]
Ernestiodendron filiciforme Tambach-Sandstein MemberSome specimens.A conifer. [3]
Metacalamostachys dumasii Tambach-Sandstein MemberSome specimens. Horsetails. [21]
Walchia piniformis Tambach-Sandstein MemberSome specimens.A conifer. [3]

Invertebrates

TaxonMemberMaterialNotesImages
cf. Anthracoblattina Tambach-Sandstein MemberSome specimens.A phylloblattid insect. [4]
Lioestheria andreevi Tambach-Sandstein MemberSome specimens.A conchostracan. Originally Lioestheria monticula, which was later considered a junior synonym of L. andreevi. [13]
Medusina limnica Tambach-Sandstein MemberMany specimens.A freshwater jellyfish. [22]
Moravamylacris kukalovae Tambach-Sandstein MemberSome specimens.A mylacrid insect. [12]
Opsiomylacris sp.Tambach-Sandstein MemberSome specimens.A mylacrid insect. [4]
Phylloblatta sp.Tambach-Sandstein MemberSome specimens.A phylloblattid insect. [4]

Scoyenia gracilis

Tambach-Sandstein Member

Numerous specimens.

Tiny burrow casts, possibly created by burrowing worms. [23]

Striatichnium bromackerense

Tambach-Sandstein Member

Numerous specimens.

Possibly feeding traces from underwater worms or arthropods raking along a substrate. [22]

Tambia spiralis

Tambach-Sandstein Member

Numerous specimens.

Small burrow casts with scratch marks, possibly created by large beetles [24] or small burrowing reptiles ( Thuringothyris ). [23]

TambiaClose.jpg

Basal Tetrapods

TaxonMemberMaterialNotesImages

Amphisauropus kablikae ?

Tambach-Sandstein Member

Rare, heavily eroded specimens.

Footprints likely created by seymouriamorphs such as Seymouria sanjuanensis . Tambach specimens may be misinterpreted examples of other ichnotaxa. [8]

Bromerpeton [25] MNG 16545 (holotype), a partial skeleton with skull and forelimb material.A brachystelechid microsaur.

Georgenthalia [26]

Tambach-Sandstein Member

MNG  11135 (holotype), a small but complete skull.

An amphibamiform dissorophoid temnospondyl.

Ichniotherium cottae

Tambach-Sandstein Member

Numerous specimens.

Footprints created by Diadectes absitus . [27]

Ichniotherium sphaerodactylum

Tambach-Sandstein Member

Numerous specimens.

Footprints created by Orobates pabsti . [27]

Orobates [28]

Tambach-Sandstein Member

Four individuals are known from the specimens MNG 10181 (holotype), MNG 8760, 8980, 11133, 11134 (paratypes).

A diadectid tetrapod.

Orobates1DB.jpg

Rotaryus [29]

Tambach-Sandstein Member

MNG 10182 (holotype), articulated partial well-preserved skull and both mandibles and a closely associated partial postcranial skeleton.

A trematopid dissorophoid temnospondyl.

Rotaryus gothae.jpg

Seymouria sanjuanensis [30]

Tambach-Sandstein Member

MNG 7727, 8759, 10553, 10554 (referred)

A seymouriid reptiliomorph.

Bromacker2 copy.jpg

Diadectes absitus [31]

Tambach-Sandstein Member

MNG 8853 (holotype), MNG 7721, 8978 (paratypes).

A diadectid tetrapod. May be its own genus, Silvadectes . [32]

Diadectes absitus (fossil amphibian) (Tambach Formation, Lower Permian; Bromacker Quarry, Thuringia, Germany) 1.jpg

Diadectes dreigleichenensis [33] Tambach-Sandstein MemberMNG 8747 (holotype), a skull with jaws. [33] A diadectid tetrapod.

Tambachia [34]

Tambach-Sandstein Member

MNG 7722 (holotype), a crushed skull and much of the postcranial skeleton.

A trematopid dissorophoid temnospondyl.

Tambachia.jpg

Tambaroter [10]

Finsterbergen conglomerate Member

MNG 14708 (holotype), an almost complete skull.

An ostodolepid microsaur.

Reptiles

TaxonMemberMaterialNotesImages

Eudibamus [35]

Tambach-Sandstein Member

MNG 8852 (holotype), an almost complete cranial and postcranial skeleton.

A bolosaurid parareptile.

Eudibamus BW.jpg

Thuringothyris [36]

Tambach-Sandstein Member

At least eight individuals are known from the specimens MNG 7729 (holotype), MNG 10183, 10647, 10652, 11191 (referred).

The basalmost known captorhinid eureptile. [37]

Synapsids

TaxonMemberMaterialNotesImages

Dimetrodon teutonis [38]

Tambach-Sandstein Member

MNG 10598 (holotype), partial vertebral column. [38] The referred specimens MNG 10654, 10655, 10693 represent much of the postcranial skeleton. The referred specimen MNG 13433 represents a right maxilla. [39]

A sphenacodontid.

Dimetropus leisnerianus

Tambach-Sandstein Member

Some specimens.

Footprints created by non-therapsid synapsids ("pelycosaurs") such as sphenacodontids. [40]

Martensius bromackerensis

Tambach-Sandstein Member

Four articulated specimens.

A caseid

Tambacarnifex unguifalcatus [41]

Tambach-Sandstein Member

MNG 10596 (holotype), partial skeleton. MNG 15037, partial left dentary.

A varanopid.

Tetrapoda indet.

TaxonMemberMaterialNotesImages

Megatambichnus sp.

Tambach-Sandstein Member

Some specimens.

Large burrows and scratch marks, likely created by diadectids. [23]

Tambachichnium schmidti

Tambach-Sandstein Member

Some specimens.

Footprints which may have been created by procolophonomorphs, [7] araeoscelidians, or varanopids. [42]

Varanopus microdactylus

Tambach-Sandstein Member

Some specimens.

Originally Ichnium microdactylum. Footprints which may have been created by captorhinomorphs ( Thuringothyris ), [42] varanopids, or Seymouria sanjuanensis . [8]

Related Research Articles

<i>Dimetrodon</i> Genus of carnivorous synapsids from the Permian

Dimetrodon is an extinct genus of non-mammalian synapsid belonging to the family Sphenacodontidae that lived during the Cisuralian age of the Early Permian period, around 295–272 million years ago. With most species measuring 1.7–4.6 m (5.6–15.1 ft) long and weighing 28–250 kg (62–551 lb), the most prominent feature of Dimetrodon is the large neural spine sail on its back formed by elongated spines extending from the vertebrae. It was an obligate quadruped and had a tall, curved skull with large teeth of different sizes set along the jaws. Most fossils have been found in the Southwestern United States, the majority of these coming from a geological deposit called the Red Beds of Texas and Oklahoma. More recently, its fossils have also been found in Germany and over a dozen species have been named since the genus was first erected in 1878.

<span class="mw-page-title-main">Diadectidae</span> Extinct family of tetrapods

Diadectidae is an extinct family of early tetrapods that lived in what is now North America and Europe during the Late Carboniferous and Early Permian, and in Asia during the Late Permian. They were the first herbivorous tetrapods, and also the first fully terrestrial animals to attain large sizes. Footprints indicate that diadectids walked with an erect posture. They were the first to exploit plant material in terrestrial food chains, making their appearance an important stage in both vertebrate evolution and the development of terrestrial ecosystems.

<i>Seymouria</i> Extinct genus of tetrapodomorphs

Seymouria is an extinct genus of seymouriamorph from the Early Permian of North America and Europe. Although they were amphibians, Seymouria were well-adapted to life on land, with many reptilian features—so many, in fact, that Seymouria was first thought to be a primitive reptile. It is primarily known from two species, Seymouria baylorensis and Seymouria sanjuanensis. The type species, S. baylorensis, is more robust and specialized, though its fossils have only been found in Texas. On the other hand, Seymouria sanjuanensis is more abundant and widespread. This smaller species is known from multiple well-preserved fossils, including a block of six skeletons found in the Cutler Formation of New Mexico, and a pair of fully grown skeletons from the Tambach Formation of Germany, which were fossilized lying next to each other.

<i>Tapinocephalus</i> Assemblage Zone

The Tapinocephalus Assemblage Zone is a tetrapod assemblage zone or biozone which correlates to the middle Abrahamskraal Formation, Adelaide Subgroup of the Beaufort Group, a fossiliferous and geologically important geological Group of the Karoo Supergroup in South Africa. The thickest outcrops, reaching approximately 2,000 metres (6,600 ft), occur from Merweville and Leeu-Gamka in its southernmost exposures, from Sutherland through to Beaufort West where outcrops start to only be found in the south-east, north of Oudshoorn and Willowmore, reaching up to areas south of Graaff-Reinet. Its northernmost exposures occur around the towns Fraserburg and Victoria West. The Tapinocephalus Assemblage Zone is the second biozone of the Beaufort Group.

Angelosaurus is an extinct genus of herbivorous caseid synapsids that lived during the late Lower Permian (Kungurian) and early Middle Permian (Roadian) in what is now Texas and Oklahoma in the United States. Like other herbivorous caseids, it had a small head, large barrel-shaped body, long tail, and massive limbs. Angelosaurus differs from other caseids by the extreme massiveness of its bones, particularly those of the limbs, which show a strong development of ridges, processes, and rugosities for the attachment of muscles and tendons. Relative to its body size, the limbs of Angelosaurus were shorter and wider than those of other caseids. The ungual phalanges looked more like hooves than claws. The few known cranial elements show that the skull was short and more robust than that of the other representatives of the group. Angelosaurus is also distinguished by its bulbous teeth with shorter and wider crowns than those of other caseids. Their morphology and the high rate of wear they exhibit suggests a diet quite different from that of other large herbivorous caseids, and must have been based on particularly tough plants. A study published in 2022 suggests that the genus may be paraphyletic, with Angelosaurus possibly only represented by its type species A. dolani.

<i>Eudibamus</i> Extinct genus of reptiles

Eudibamus is an extinct genus of biped bolosaurid reptile known from the Free State of Thuringia of central Germany. It had a very small size reaching only 25-26 cm in length.

<i>Orobates</i> Extinct genus of reptiliomorphs

Orobates is an extinct genus of diadectid reptiliomorphs that lived during the Early Permian. Its fossilised remains were found in Germany. A combination of primitive and derived traits distinguish it from all other well-known members of Diadectidae, a family of herbivorous reptiliomorphs. It weighed about 4 kg and appears to have been part of an upland fauna, browsing on high fibre plants.

<span class="mw-page-title-main">Brachystelechidae</span> Extinct family of tetrapods

Brachystelechidae is an extinct family of Early Permian microsaurs. The family was first named by Robert L. Carroll and Pamela Gaskill in 1978, with the only member being Brachystelechus fritschi. Brachystelechus fritschi has since been reassigned to the genus Batropetes. Genera assigned to the family include: Batropetes, from Germany; Carrolla, from Texas; Quasicaecilia, also from Texas; Diabloroter, from the Mazon Creek lagerstätte of Illinois; and Bromerpeton from the Tambach Formation of Germany.

<i>Tambachia</i> Extinct genus of amphibians

Tambachia is an extinct genus of dissorophoid temnospondyl within the family Trematopidae. It is known from the Early Permian Tambach Formation near the town of Tambach-Dietharz in Thuringia, Germany. Tambachia is the first trematopid to have been discovered outside the United States.

<span class="mw-page-title-main">Ostodolepidae</span> Extinct family of tetrapods

Ostodolepidae, also spelled Ostodolepididae, is an extinct family of Early Permian tetrapods. Initially they were considered microsaurs, but later were assigned to the group Recumbirostra. Ostodolepids were relatively large, reaching lengths of up to 2 feet (61 cm), terrestrial, and presumably fossorial. Ostodolepid remains have been found from Early Permian beds in Texas, Oklahoma, and Germany.

<i>Georgenthalia</i> Extinct genus of amphibians

Georgenthalia is an extinct genus of dissorophoid temnospondyl from the Lower Permian. It is an amphibamid which lived in what is now the Thuringian Forest of central Germany. It is known from the holotype MNG 11135, a small, complete skull. It was found in the Bromacker locality of the Tambach Formation. It was first named by Jason S. Anderson, Amy C. Henrici, Stuart S. Sumida, Thomas Martens and David S. Berman in 2008 and the type species is Georgenthalia clavinasica.

Tambaroter is an extinct genus of ostodolepid microsaur from the Early Permian of Germany. The type species T. carrolli was named in 2011. Tambaroter is known from a single skull found in the Tambach Formation, which is the lowermost unit of the Upper Rotliegend. It is the only vertebrate that has been found outside the Bromacker Quarry, the most productive locality of the formation. It is also the first ostodolepid known from outside North America.

Thuringothyris is an extinct genus of Early Permian eureptiles known from the Thuringian Forest in central Germany.

<i>Rotaryus</i> Extinct genus of amphibians

Rotaryus is an extinct genus of Early Permian trematopid dissorophoid temnospondyl known from the Free State of Thuringia of central Germany.

Tambacarnifex is an extinct genus of varanodontine synapsids known from the Early Permian Tambach Formation of Free State of Thuringia, central Germany. It was first named by David S. Berman, Amy C. Henrici, Stuart S. Sumida, Thomas Martens and Valerie Pelletier in 2013 and the type species is Tambacarnifex unguifalcatus.

The Arroyo Formation, sometimes termed the Lower Clear Fork Formation, is a geologic formation in Texas. It preserves fossils dating back to the Kungurian stage of the Permian period. It is the lower-most portion of the Clear Fork Group, part of a series of fossiliferous Permian strata in the south-central United States known as the red beds.

<span class="mw-page-title-main">Abo Formation</span> Geological formation in New Mexico

The Abo Formation is a geologic formation in New Mexico. It contains fossils characteristic of the Cisuralian epoch of the Permian period.

<span class="mw-page-title-main">Organ Rock Formation</span>

The Organ Rock Formation or Organ Rock Shale is a formation within the late Pennsylvanian to early Permian Cutler Group and is deposited across southeastern Utah, northwestern New Mexico, and northeastern Arizona. This formation notably outcrops around Canyonlands National Park, Natural Bridges National Monument, and Monument Valley of northeast Arizona, southern Utah. The age of the Organ Rock is constrained to the latter half of the Cisuralian epoch by age dates from overlying and underlying formations. Important early terrestrial vertebrate fossils have been recovered from this formation in northern Arizona, southern Utah, and northern New Mexico. These include the iconic Permian terrestrial fauna: Seymouria, Diadectes, Ophiacodon, and Dimetrodon. The fossil assemblage present suggests arid environmental conditions. This is corroborated with paleoclimate data indicative of global drying throughout the early Permian.

<span class="mw-page-title-main">Teekloof Formation</span> Late Permian geological formation that forms part of the Beaufort Group of South Africa

The Teekloof Formation is a geological formation that forms part of the Beaufort Group, one of the five geological groups that comprises the Karoo Supergroup in South Africa. The Teekloof Formation is the uppermost formation of Adelaide Subgroup deposits West of 24ºE and contains Middle to Late Permian-aged deposits and four biozones of the Beaufort Group. It overlies the Abrahamskraal Formation. The Teekloof Formation does not underlie other units other than the younger Karoo dolerites and sills that relate to the emplacement of the Early Jurassic Drakensberg Group to the east. Outcrops and exposures of the Teekloof Formation range from Sutherland through the mountain escarpments between Fraserburg and Beaufort West. The northernmost localities of the Teekloof Formation are found by Loxton, Victoria West and Richmond.

The Timezgadiouine Formation, sometimes spelled as the Timesgadiouine Formation, is a Triassic geological formation in the Argana Basin of Morocco. It is a succession of red bed sediments spanning from the Olenekian to at least the Carnian, encompassing members T3 to T5 of the Argana Group. It is preceded by the Permian Ikakern Formation and succeeded by the Late Triassic Bigoudine Formation.

References

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