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The Tambach Formation is an
Early Permian 01 or '01 may refer to: * The year 2001, or any year ending with 01 * The month of January * 1 (number) Music * '01 (Richard Müller album), 01'' (Richard Müller album), 2001 * 01 (Son of Dave album), ''01'' (Son of Dave album), 2000 * 01 (Urban ...
-age
geologic formation A geological formation, or simply formation, is a body of rock having a consistent set of physical characteristics ( lithology) that distinguishes it from adjacent bodies of rock, and which occupies a particular position in the layers of rock exp ...
in central
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
. It consists of red to brown-colored
sedimentary rock Sedimentary rocks are types of rock that are formed by the accumulation or deposition of mineral or organic particles at Earth's surface, followed by cementation. Sedimentation is the collective name for processes that cause these particles ...
s (
red beds Red beds (or redbeds) are sedimentary rocks, typically consisting of sandstone, siltstone, and shale, that are predominantly red in color due to the presence of ferric oxides. Frequently, these red-colored sedimentary strata locally contain ...
) such as
conglomerate Conglomerate or conglomeration may refer to: * Conglomerate (company) * Conglomerate (geology) * Conglomerate (mathematics) In popular culture: * The Conglomerate (American group), a production crew and musical group founded by Busta Rhymes ** Co ...
,
sandstone Sandstone is a clastic sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks. Most sandstone is composed of quartz or feldspar (both silicates) ...
, and
mudstone Mudstone, a type of mudrock, is a fine-grained sedimentary rock whose original constituents were clays or muds. Mudstone is distinguished from '' shale'' by its lack of fissility (parallel layering).Blatt, H., and R.J. Tracy, 1996, ''Petrology. ...
, and is the oldest portion of the Upper Rotliegend within the Thuringian Forest Basin. 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 river A braided river, or braided channel, consists of a network of river channels separated by small, often temporary, islands called braid bars or, in English usage, ''aits'' or ''eyots''. Braided streams tend to occur in rivers with high sediment l ...
s. These incise underlying
rhyolitic Rhyolite ( ) is the most silica-rich of volcanic rocks. It is generally glassy or fine-grained (aphanitic) in texture, but may be porphyritic, containing larger mineral crystals (phenocrysts) in an otherwise fine-grained groundmass. The mineral ...
rock, depositing a coarse
conglomerate Conglomerate or conglomeration may refer to: * Conglomerate (company) * Conglomerate (geology) * Conglomerate (mathematics) In popular culture: * The Conglomerate (American group), a production crew and musical group founded by Busta Rhymes ** Co ...
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. 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 In geology, a facies ( , ; same pronunciation and spelling in the plural) is a body of rock with specified characteristics, which can be any observable attribute of rocks (such as their overall appearance, composition, or condition of formatio ...
types, with different fossil components. Thick sandstone channel fills in the lower section at Bromacker are overlain with mudstone from
ephemeral Ephemerality (from the Greek word , meaning 'lasting only one day') is the concept of things being transitory, existing only briefly. Academically, the term ephemeral constitutionally describes a diverse assortment of things and experiences, fr ...
lakes. A diverse assemblage of trace fossils such as footprints are imprinted onto the mudstone drapings. The upper section of Bromacker contains
siltstone Siltstone, also known as aleurolite, is a clastic sedimentary rock that is composed mostly of silt. It is a form of mudrock with a low clay mineral content, which can be distinguished from shale by its lack of fissility.Blatt ''et al.'' 1980, p ...
deposited through sheetfloods, in which well-preserved articulated skeletons of terrestrial
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
s have been discovered. These include early amphibians like ''
Rotaryus ''Rotaryus'' is an extinct genus of Early Permian trematopid dissorophoid temnospondyl known from the Free State of Thuringia of central Germany. Discovery ''Rotaryus'' is known only from the holotype MNG 10182, articulated partial well-pr ...
'' and ''
Tambaroter ''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 The Tambach Formation ...
,'' and early reptiles like ''
Eudibamus ''Eudibamus'' is an extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, alth ...
'' and '' Thuringothyris''. Unlike most fossil-preserving Permian red beds, aquatic
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s are absent at Bromacker while carnivorous synapsids (like ''
Dimetrodon ''Dimetrodon'' ( or ,) meaning "two measures of teeth,” is an extinct genus of non-mammalian synapsid that lived during the Cisuralian (Early Permian), around 295–272 million years ago (Mya). It is a member of the family Sphenacodontid ...
'') are rare and herbivorous diadectids are abundant. Plant and arthropod fossils have also been found in shales at Bromacker.


History

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, 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. 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. 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 East Germany, officially the German Democratic Republic (GDR; german: Deutsche Demokratische Republik, , DDR, ), was a country that existed from its creation on 7 October 1949 until its dissolution on 3 October 1990. In these years the state ...
paleontologists such as
Hermann Schmidt Hermann or Herrmann may refer to: * Hermann (name), list of people with this name * Arminius, chieftain of the Germanic Cherusci tribe in the 1st century, known as Hermann in the German language * Éditions Hermann, French publisher * Hermann, Miss ...
, Arno Hermann Müller, and Hartmut Haubold. 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. 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 Friedenstein Palace (german: Schloss Friedenstein) is an early Baroque palace built in the mid-17th century by Ernest I, Duke of Saxe-Gotha at Gotha, Thuringia, Germany. In Germany, ''Friedenstein'' was one of the largest palaces of its time and o ...
. 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.


Geology

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 In geology, a graben () is a depressed block of the crust of a planet or moon, bordered by parallel normal faults. Etymology ''Graben'' is a loan word from German, meaning 'ditch' or 'trench'. The word was first used in the geologic contex ...
(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. Traditionally, the Tambach Formation is considered to be divided into three discrete layers: a lower and an upper
conglomerate Conglomerate or conglomeration may refer to: * Conglomerate (company) * Conglomerate (geology) * Conglomerate (mathematics) In popular culture: * The Conglomerate (American group), a production crew and musical group founded by Busta Rhymes ** Co ...
layer (Bielstein and Finsterbergen Konglomerate, respectively), separated by a somewhat narrower layer of finer sediments such as
sandstone Sandstone is a clastic sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks. Most sandstone is composed of quartz or feldspar (both silicates) ...
, the Tambach-Sandstein (Tambach sandstone) member. 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. 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 river A braided river, or braided channel, consists of a network of river channels separated by small, often temporary, islands called braid bars or, in English usage, ''aits'' or ''eyots''. Braided streams tend to occur in rivers with high sediment l ...
s flowing along its edge with lower-energy rivers,
floodplain A floodplain or flood plain or bottomlands is an area of land adjacent to a river which stretches from the banks of its channel to the base of the enclosing valley walls, and which experiences flooding during periods of high discharge.Goudi ...
s, and lakes at its center. The coarse (
cobble Cobble may refer to: * Cobble (geology), a designation of particle size for sediment or clastic rock * Cobblestone, partially rounded rocks used for road paving * Hammerstone, a prehistoric stone tool * Tyringham Cobble, a nature reserve in Tyr ...
/
boulder In geology, a boulder (or rarely bowlder) is a rock fragment with size greater than in diameter. Smaller pieces are called cobbles and pebbles. While a boulder may be small enough to move or roll manually, others are extremely massive. In c ...
-scale) and
rhyolitic Rhyolite ( ) is the most silica-rich of volcanic rocks. It is generally glassy or fine-grained (aphanitic) in texture, but may be porphyritic, containing larger mineral crystals (phenocrysts) in an otherwise fine-grained groundmass. The mineral ...
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. Decreasing tectonic activity in the middle stage (stage II) leads to increased
erosion Erosion is the action of surface processes (such as water flow or wind) that removes soil, rock, or dissolved material from one location on the Earth's crust, and then transports it to another location where it is deposited. Erosion is distin ...
, 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 A pebble is a clast of rock with a particle size of based on the Udden-Wentworth scale of sedimentology. Pebbles are generally considered larger than granules ( in diameter) and smaller than cobbles ( in diameter). A rock made predominant ...
conglomerate. The center of the basin graded into small streams and marshes, depositing sandstone,
siltstone Siltstone, also known as aleurolite, is a clastic sedimentary rock that is composed mostly of silt. It is a form of mudrock with a low clay mineral content, which can be distinguished from shale by its lack of fissility.Blatt ''et al.'' 1980, p ...
,
shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4) and tiny fragments (silt-sized particles) of other minerals, especial ...
and mudstone in fossiliferous
red beds Red beds (or redbeds) are sedimentary rocks, typically consisting of sandstone, siltstone, and shale, that are predominantly red in color due to the presence of ferric oxides. Frequently, these red-colored sedimentary strata locally contain ...
. Although the Tambach Basin may have been hydrologically isolated during this period, with its waterways draining internally, some paleontologists instead consider its waters to flow into another basin in the northeast, which was not preserved. 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 Conglomerate or conglomeration may refer to: * Conglomerate (company) * Conglomerate (geology) * Conglomerate (mathematics) In popular culture: * The Conglomerate (American group), a production crew and musical group founded by Busta Rhymes ** Co ...
to build up and expand into the center of the basin, forming the Finsterbergen conglomerate.


Bromacker

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" or "Bromacker sandstone" and stage II sediments are the "upper beds" or "Bromacker horizon". 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 Mudstone, a type of mudrock, is a fine-grained sedimentary rock whose original constituents were clays or muds. Mudstone is distinguished from '' shale'' by its lack of fissility (parallel layering).Blatt, H., and R.J. Tracy, 1996, ''Petrology. ...
, which sometimes preserved
mudcrack Mudcracks (also known as mud cracks, desiccation cracks or cracked mud) are sedimentary structures formed as muddy sediment Desiccation, dries and contracts.Jackson, J.A., 1997, ''Glossary of Geology'' (4th ed.), American Geological Institute, Ale ...
s, plant fossils,
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
burrows, and
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
footprints. The thick sandstone layers (and their mudstone drapings) are not continuous, interrupted by a succession of finer and darker
mica Micas ( ) are a group of silicate minerals whose outstanding physical characteristic is that individual mica crystals can easily be split into extremely thin elastic plates. This characteristic is described as perfect basal cleavage. Mica is ...
ceous sediments such as
shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4) and tiny fragments (silt-sized particles) of other minerals, especial ...
,
siltstone Siltstone, also known as aleurolite, is a clastic sedimentary rock that is composed mostly of silt. It is a form of mudrock with a low clay mineral content, which can be distinguished from shale by its lack of fissility.Blatt ''et al.'' 1980, p ...
, and (rarely) very fine sandstone. This reconstructs a sequence of repeated flooding events, involving strong, straight rivers eroding channels through the silty
floodplain A floodplain or flood plain or bottomlands is an area of land adjacent to a river which stretches from the banks of its channel to the base of the enclosing valley walls, and which experiences flooding during periods of high discharge.Goudi ...
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 Ephemerality (from the Greek word , meaning 'lasting only one day') is the concept of things being transitory, existing only briefly. Academically, the term ephemeral constitutionally describes a diverse assortment of things and experiences, fr ...
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. 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 Calcite is a Carbonate minerals, carbonate mineral and the most stable Polymorphism (materials science), polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on ...
-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 Clam shrimp are a group of bivalved branchiopod crustaceans that resemble the unrelated Bivalvia, bivalved molluscs. They are extant and also known from the fossil record, from at least the Devonian period and perhaps before. They were originally ...
and
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
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. 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 Breccia () is a rock composed of large angular broken fragments of minerals or rocks cemented together by a fine-grained matrix. The word has its origins in the Italian language, in which it means "rubble". A breccia may have a variety of di ...
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.


Age

Uranium-Lead dating is not possible for the Tambach Formation, which lacks fresh volcanic rocks. The similar Elgersburg Formation to the southeast contains
rhyolite Rhyolite ( ) is the most silica-rich of volcanic rocks. It is generally glassy or fine-grained (aphanitic) in texture, but may be porphyritic, containing larger mineral crystals (phenocrysts) in an otherwise fine-grained groundmass. The mineral ...
dated to 274 ± 4.9 million years ago. However, it is unclear whether the strata at
Elgersburg Elgersburg is a municipality situated in the district of Ilm-Kreis, Thuringia, Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Rus ...
are younger, older, or equivalent in age to the Tambach Formation.
Biostratigraphy Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock Stratum, strata by using the fossil assemblages contained within them.Hine, Robert. “Biostratigraphy.” ''Oxford Reference: Dictiona ...
is more informative but still imprecise. Insect and conchostrachan biostratigraphy places it into the Sakmarian- Artinskian '' Moravamylacris kukalovae'' and late Artinskian '' Lioestheria monticula/andreevi'' 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 ''Dimetrodon'' ( or ,) meaning "two measures of teeth,” is an extinct genus of non-mammalian synapsid that lived during the Cisuralian (Early Permian), around 295–272 million years ago (Mya). It is a member of the family Sphenacodontid ...
'' present at Tambach is smaller than those present in the red beds of Texas, the Tambach Formation was likely older than those formations. 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. Combining both invertebrate and tetrapod biostratigraphy, the age of the Tambach Formation was considered to be probably Artinskian in age. 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. This estimate is based primarily on the radiometric age of 295.8 ± 0.4 Ma (late Asselian) of the Rotterode Formation which
unconformably An unconformity is a buried erosional or non-depositional surface separating two rock masses or strata of different ages, indicating that sediment deposition was not continuous. In general, the older layer was exposed to erosion for an interval o ...
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. In addition, comparison of the footprint assemblage of the Tambach Formation with radiometrically dated Permian footprint assemblages from
France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
and
Italy Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the middle of the Mediterranean Sea, and its territory largely coincides with the homonymous geographical re ...
also suggests a Sakmarian age.


Climate

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 Tropical savanna climate or tropical wet and dry climate is a tropical climate sub-type that corresponds to the Köppen climate classification categories ''Aw'' (for a dry winter) and ''As'' (for a dry summer). The driest month has less than of p ...
, despite its lack of grass. Modern climatic equivalents include the northern African savanna and the
Llanos The Llanos (Spanish ''Los Llanos'', "The Plains"; ) is a vast tropical grassland plain situated to the east of the Andes in Colombia and Venezuela, in northwestern South America. It is an ecoregion of the tropical and subtropical grasslands, sav ...
of
Venezuela Venezuela (; ), officially the Bolivarian Republic of Venezuela ( es, link=no, República Bolivariana de Venezuela), is a country on the northern coast of South America, consisting of a continental landmass and many islands and islets in th ...
and
Colombia Colombia (, ; ), officially the Republic of Colombia, is a country in South America with insular regions in North America—near Nicaragua's Caribbean coast—as well as in the Pacific Ocean. The Colombian mainland is bordered by the Car ...
. However, from the sediment
geochemistry Geochemistry is the science that uses the tools and principles of chemistry to explain the mechanisms behind major geological systems such as the Earth's crust and its oceans. The realm of geochemistry extends beyond the Earth, encompassing the e ...
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). 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. The climate may have been drier during the conglomerate-rich periods of the Tambach Formation.


Paleobiota

The ecosystem of the Tambach Formation is unusual for its lack of aquatic animals such as xenacanthid sharks, '' Eryops'', or ''
Diplocaulus ''Diplocaulus'' (meaning "double caul") is an extinct genus of lepospondyl amphibians which lived from the Late Carboniferous to the Late Permian of North America and Africa. ''Diplocaulus'' is by far the largest and best-known of the lepospondy ...
'', 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 Clam shrimp are a group of bivalved branchiopod crustaceans that resemble the unrelated Bivalvia, bivalved molluscs. They are extant and also known from the fossil record, from at least the Devonian period and perhaps before. They were originally ...
, were able to flourish. The Tambach Basin did support a diverse
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
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 ''Dimetrodon'' ( or ,) meaning "two measures of teeth,” is an extinct genus of non-mammalian synapsid that lived during the Cisuralian (Early Permian), around 295–272 million years ago (Mya). It is a member of the family Sphenacodontid ...
'' 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 Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All extan ...
, while
seed ferns A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosperm pl ...
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.


Flora


Invertebrates


Basal Tetrapods


Reptiles


Synapsids


Tetrapoda ''indet.''


References

{{reflist Geologic formations of Germany Permian System of Europe Sakmarian Paleontology in Germany