Chirotherium

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Part of the original layer surface (layer underside) from Heßberg with Chirotherium barthii (bottom left, center and top right) in the Teylers Museum, Haarlem, Netherlands.

Chirotherium is a trace fossil genus fünffingeriger or -zehiger (pentadactyler) footprints and corresponding tracks of terrestrial vertebrates (Tetrapoda). In the ethological trace classification according to Seilacher it is therefore a repichnium (movementtrace) or a cursichnium (walking trace) in the subdivision of movement traces according to Müller . Chirotherium is typical for terrestrial sandstones of the Lower and Middle Triassic (approx. 250 to 230 mya ) and was probablycausedby the purely land-based ancestors of today's crocodiles , the "Rauisuchians".

Chirotherium or the type species Ch. Barthii is the first trace fossil ever named after Linnaeus' nomenclature .

In addition to the numerous described Chirotherium species, morphologically similar trace genera, such as Brachychirotherium , Isochirotherium , Protochirotherium or Synaptichnium , are informally referred to as “ Chirotherium ” .

etymology

The name is made up of the ancient Greek words χειρός, cheiros , "hand" and θηρίον, therion , "beast", "wild animal", which in the relevant German-language literature is usually translated as "hand animal" (spelling in 19th century literature). Century: "Handthier"). The name was coined in 1835 by the Darmstadt zoology professor Johann Jakob Kaup and refers to a certain similarity of the prints with human hands as well as the fact that Kaup, in his interpretation of the trace, gave preference to the possibility that the traces could have come from a mammal .

history

Part of the original layer surface from Hildburghausen in the Goldfuß Museum Bonn. The step seals are in the middle in the lower half and on the right in the upper half. Manus and Pes are each very close together.

First find and response

In the spring of 1833, the Hildburghausen high school director Friedrich Sickler discovered imprints on a sandstone slab that came from a quarry near Heßberg and was to be built into the foundation wall of his garden shed, which reminded him of the footsteps of animals. Sickler then offered the quarry workers money to watch out for further traces and to secure them. In the summer of 1834, a larger layer area with tracks was exposed, which Sickler described in the form of an open letter to the Göttingen anatomist and zoologist Johann Friedrich Blumenbach . In the following months, various naturalists published comments on Sickler's description or the trace finds, including a. the Heidelberg geology professor Heinrich Georg Bronn , his Bonn colleague Johann Jacob Nöggerath , the Darmstadt zoology professor Johann Jakob Kaup and also the famous Alexander von Humboldt .

In contrast to Bronn and Humboldt, who considered it probable that the tracks were from mammals, and to Sickler, who even went so far, because of the similarity of the prints with human hands, the origin of prehistoric monkeys ("Quadrumans") not To be able to exclude, Nöggerath pointed out that mammal fossils in sediments, which were the same age as the tracing sandstones of Hildburghausen ("either colorful sandstone or Keuper formation"), had never been found and a reptile as Cause of the "Tazzen reliefs" would be much more likely. At Kaup, who had meanwhile acquired a sandstone slab from Heßberg and was therefore able to inspect the track personally, the name Chirotherium finally appears for the first time . Since Kaup, on the one hand, did not intend to name the trace itself, but rather its producer, and on the other hand, it also tended to refer to a mammal, he chose this designation ("hand mammal"), but kept one later renaming to "Chirosaurus" ("Hand-Lizard"), if his assessment should prove to be wrong.

While research on trace fossils was of relatively minor importance in the 19th century, modern palichnology developed in the first half of the 20th century. The binary (Linnaeus') nomenclature was retained and now consistently applied exclusively to the traces, although the names were originally intended either to designate the trace producer or the traces themselves were considered fossilized organisms and were named as such. With this restriction, Chirotherium and its type species Chirotherium barthii (the species epithet pays tribute to the graphic artist Carl Barth, who was instrumental in the recovery of the type material in Sickler's absence ) are the first fossil land vertebrate traces in the history of paleontology to be named in this way. To commemorate this “premiere”, the Chirotherium Monument was inaugurated on September 12, 2004 on the market square in Hildburghausen.

Sickler's open letter about the tracks in the Hildburghausen red sandstone is not the first scientific mention of fossil tetrapod traces. Already in 1828 the pastor and naturalist J. Grierson reported about Footsteps before the Flood ("Footprints from before the Flood") in the "New Red Sandstone" (red sandstones of the Permian and Triassic) of Dumfriesshire (South Scotland), which in turn was already there were known since at least 1814. In addition, given the abundance of chirotherium in Triassic sandstones, it is very likely that humans were aware of these trace fossils well before the 19th century, but without realizing what they were.

Limitation of the producer

Owen's reconstruction of a "Labyrinthodontier" as a trace producer of Chirotherium from the year 1855. Note that Labyrinthodon (today mostly Mastodonsaurus ) was considered a "Batrachier", a relatively close relative of today's frogs, which is why the "thumb" is interpreted as an imprint of the first toe has been. Accordingly, the animal would have had to cross its legs when walking in order to create the track. It should also be noted that the distal extremity bones of Labyrinthodon were not even known at the time.

While, as stated above, German paleontologists wavered between reptiles and mammals as producers in the 1830s, in 1842 the famous British paleontologist Richard Owen was apparently one of the first to imagine that the cause of chirotherium was a relatively close relative of the crocodiles could be, and that the supposed “thumb” should actually be the fifth toe. However, he himself was convinced that the track came from labyrinthodontists , non-amniotic, distantly crocodile-like tetrapods. Owen regarded the “labyrinthodontians” as relatively close relatives of today's frogs (which is true to a certain extent - at least they are no more closely related to crocodiles than to frogs) and therefore preferred the interpretation of the “thumb” as the first toe.

In 1914, 80 years after the tracks were first mentioned scientifically, it was the Briton DMS Watson who first favored the ancestors of dinosaurs (at that time placed in a group called " Thecodontia ") and thus for the first time favored early archosaurs as the cause of the tracks. The original assumption that they were mammals was based in large part on the fact that the width of the track, i.e. That is, the distance between the step seal and the body axis of symmetry of the trace generator was very small and in the 1830s the view was still prevalent that without exception all prehistoric reptiles, just like today's lizards and turtles, have limbs that are strongly angled from the body (splaying gait) and not, like most mammals, with stretched limbs standing under the trunk (stalk). It was not until Watson's time that knowledge of the anatomy of prehistoric reptiles had advanced to such an extent that the splayed gait thesis was no longer tenable for archosaurs. In addition, it was also known until then that dinosaurs already lived in the late Triassic, which made a dinosaur ancestor more likely as a producer of Triassic traces than in Owen's time, when dinosaurs were only known from the Jurassic .

The work of the German Wolfgang Soergel from 1925, who also benefited from the advances made in paleontology up to that point in his search for the chirotherium producer, is considered particularly trend-setting . He compared the shape (morphology) of the footprints with the skeletons of the feet of the Triassic reptiles known in the 1920s and found the greatest agreement in a group of "Thecodonti", which at that time were summarized under the term "Pseudosuchia". The greatest similarity existed with the extremities of the Euparkeria capensis described by the famous Robert Broom from the Triassic of the Karoo basin of South Africa . Since Euparkeria was only known from South Africa (and is still today) and was also much too small to have produced traces of the size of most known Chirotherium species, it was not directly possible as the cause. Soergel therefore constructed a phantom image of a large "pseudo-searcher" similar to Euparkeria .

In 1965, Bernard Krebs published a work on the Central Triassic "Rauisuchier" (a subgroup of the "Pseudosuchier") Ticinosuchus from Monte San Giorgio . Ticinosuchus was significantly larger than Euparkeria and by reconstructing the locomotor system of this reptile, Krebs proved that it was actually able to produce “chirotheroid” tracks. However, these tracks could not be brought into agreement with any of the Chirotherium species (see also →  producers ).

morphology

Graphic representation of a Chirotherium track with three manus pes sentences (left) and the associated diagnostic trace parameters (right).

A set of Chirotherium step seals includes an imprint of the rear extremity (Pes) and a significantly smaller (half as large or smaller) imprint of the front extremity (Manus). Both imprints are fifty-toed or -fingered (pentadactyl), with the tip of toes I to IV ending in claws and toes II to IV being significantly stronger and longer than toes I and V. The third toe is the longest. On the manus, claws are less clearly developed and fingers I-IV are all about the same length and strength. A special feature is the clawless fifth toe or finger, which protrudes strongly to the side and is sometimes slightly bent backwards, which was interpreted as a “thumb” (i.e. first finger) by the first processors (see above ). The resemblance of the Pes imprint with a human hand, which is particularly evoked by this “thumb”, was ultimately the reason for the name “hand animal”.

The distance between two manus pes sentences within a track is a little more than the length of a manus pes sentence. The width of the track, i.e. the distance between the tread seals and the imaginary center line of the track, is very small, which is at a step angle (angle between the imaginary lines connecting a Manus or Pes triplet) of 160 to 170 ° expresses. At that time Bronn even used the huntsman's term lacing , as the tread seals are almost on one line. The longitudinal axis of a manus-pes sentence is usually not parallel to the center line of the track, but forms a small angle with it.

conservation

Chirotherium barthii in typical preservation as a convex hyporelief in sandstone (right) and a plaster cast made of it (left, trace prints colored), which corresponds to the original terrain surface before it was covered by the sand.

Chirotherium is typically preserved on the underside of the layers of sandstone banks (so-called convex hyporelief ) and is often associated with dry crack fillings that occasionally cross the tread seals. In this special case, the deposits are characteristic of a semi-arid alluvial plain : at the end of the rainy season, the rivers flood and overflow their banks. In the flooded areas, sediments are deposited that were carried along by the water masses. First sand, then silt and on top fine clay particles are deposited as mud. After the onset of the dry season, the floods recede and the mud, which covers large parts of the plain, begins to dry up. The first dry cracks are forming, but the mud is still moist enough that animals running over it can sink into it a little and leave step seals. However, the drying process progresses quickly and further cracks form, some of which also run through the tread seals. In the finally hardened sludge layer, dry cracks and step seals are now preserved at least until the end of the drying time. During the next flood, sand is deposited on top of the dried mud and fills the dry cracks and the tread seals. These fillings correspond to the reliefs of tracks and net-like structures that can be found on the undersides of many “track sandstones ”. The mudstone layers, which correspond to the mud layer that contains the original footsteps and dry cracks, are also geologically transmitted, but the mudstone is often very brittle and disintegrates when the sandstone banks are broken down.

In relatively rare cases, the footsteps contain impressions of the horn scales on the sole of the producer's foot.

Occurrence

Germany

In Germany, chirotherium is mainly known from the Upper Middle and Upper Buntsandstein in southern Lower Saxony, Hesse, Thuringia and northern Bavaria, where it is much more common than body fossils from terrestrial vertebrates.

The type occurrence of Chirotherium , or the type species Ch. Barthii and another species described by Kaup, Ch. Sickleri , is located near Hildburghausen in Thuringia in the highest part of the Solling Formation (uppermost Middle Buntsandstein). The corresponding stratigraphic horizon is now known far beyond the Hildburghausen region as Thuringian Chirotheriensandstein . Fragments of the approx. 120 m² track area uncovered in 1834 with the type material by Ch. Barthii are today in 30 different museums and other paleontological collections in various cities in Germany and Europe. a. in Berlin, Frankfurt, Prague, Vienna, Paris and London.

From the upper part of the Solling Formation on the southern edge of the Solling near Bad Karlshafen on the Lower Saxony-Hessian border, which is occupied by the so-called Karlshafen sandstone , the find of a trace association with u. a. Chirotherium sickleri .

Again in the Thuringian chirotherium sandstone, in a former quarry near Eiterfeld on the western edge of the Soisberger Kuppenrhön , is one of the largest known tracking plates with chirotherium in Germany (approx. 300 m²). The site was discovered during mapping work in 1963 and scientifically investigated in 1964. After the quarry was closed, the surrounding villages used the quarry as a building rubble dump and in 1977, when efforts to preserve the quarry as a natural monument promised to be crowned with success, the pit was so filled that plans were dropped. Another relatively important site in the Thuringian chirotheria sandstone, discovered by the local pastor Josef Vorbeck in 1841, is located near Aura near Bad Kissingen in the southern foreland of the Rhön (also known as the southern Rhön ). Some of the step seals have been ascribed to their own Chirotherium species, Ch. Vorbachi , and some originally even to a new genus of their own with the species " Saurichnites auraensis " (now Ch. Barthii ).

Sandstone slab with three longer tracks from Chirotherium sp. from the red quartzite from Külsheim near Würzburg , Baden-Württemberg. Clearly recognizable are both the differently sized footsteps of Manus (small) and Pes (large) as well as the "lacing" of the tracks.

In southern Germany, especially in Main Franconia and neighboring areas, chirotherium tracks can also be found in the Upper Buntsandstein ( Röt ). These deposits gave the corresponding horizons the name Franconian Chirotheriensschichten . A distinction is made between the predominantly clayey chirotheria slate at the red base from the border quartzite and the red quartzite of the higher red. The terms “Mittlerer Chirotheriensandstein” and “Oberer Chirotheriensandstein” are also used for border quartzite and red quartzite (the “Untere Chirotheriensandstein” is the Thuringian Chirotheriensandstein). The outcrop of the Upper Buntsandstein including the Franconian Chirotherienschichten stretches from the Südrhön to the southwest over the sandstone Spessart into the sandstone Odenwald and accordingly there are also tracking sites in this strip, u. a. in Gössenheim , Gambach , Külsheim , Hardheim and Ünglert near Mudau . The Gambach site is either in the border quartzite or in the red quartzite and is a type locality of the species " Saurichnites gambachensis " (today Chirotherium barthii ). The tracks recovered near Külsheim in the northern sandstone Odenwald in the northeast of Baden-Württemberg were given to the Natural History Museum in Stuttgart between 1989 and 1992 . A single, larger slab is on display north of Külsheim on a parking lot on the L 509 as part of a geological nature trail. In Hardheim, less than 10 kilometers southwest of Külsheim, Ch. Barthii and Ch. Sickleri were excavated in autumn 1992 as part of one of the largest recent track finds in Germany's red sandstone. It is not known whether the “chirotheria” found in Ünglert, also located in the northern sandstone Odenwald, also belong to the genus Chirotherium .

Apparently, chirotherium is rare in the red sandstone of the Palatinate . In this regard, an older individual find by Ch. Barthii in the Upper Buntsandstein near Pirmasens, which was documented in detail in 2012, deserves special mention.

In the German limestone are Chirotherium -Fährten handed. The corresponding layers do not represent a river landscape, but a mudflat area . The traces are not raised here on the underside of the layer, but as hollow shapes on the upper side of the layer ( concave epirelief ). An example of such a site is the Karlstadt Formation (Middle Muschelkalk, Anisium) from Bernburg , Saxony-Anhalt.

One of the stratigraphically youngest representatives of the genus in Germany is Chirotherium wondrai from the Ansbacher sandstone ( Stuttgart formation , "Schilfsandstein", Middle Keuper ) from Altselingsbach in Middle Franconia .

Rest of Europe

Movement from left manus and pes by Ch. Storetonense from Helsby sandstone, British red sandstone equivalent , Bebington near Liverpool, exhibited in the Oxford University Museum.

The first scientifically described finds of Chirotherium outside Germany were made in England in the Central Triassic ( Anisium ) Helsby sandstone formation (Sherwood sandstone group) in the Storeton quarries near Bebington , immediately southwest of Liverpool , in 1838. The quarries of Lymm near Warrington , about 30 kilometers east of Liverpool, which are not in the Helsby Formation, but in the somewhat younger Tarporley Siltstein Formation (Mercia-Claystone Group), produced traces as early as 1842. Storeton and Lymm formed the most important and productive chirotherium deposits in the British Isles, but are no longer accessible today. Today, three species are distinguished on the British Isles: Ch. Barthii , Ch. Sickleri and Ch. Storetonense , the latter only being known from Storeton. Fossil footsteps in the late Lower to early Middle Triassic Auchenew strata of the Isle of Arran off the west coast of Scotland , which were originally identified as Ch. Barthii in 2002 , are considered to represent the Ichnospecies Isochirotherium herculis after investigation of recent finds . I. herculis itself was also described as a Chirotherium species as early as 1838 . The type occurrence is probably located near Tarporley in the formation of the same name. Another species, which was identified in 1954 by Donald Baird on a track record from the Helsby sandstone by Storeton and named Ch. Lomasi , is today a representative of the trace genus Isochirotherium .

In France there are a number of trace sites with chirotherium , the geologically older ( Olenekian to Middle Anisium) on the western edge of the Vosges , the younger (Middle Anisium to Ladinium ) mainly along the east and southeast edge of the Massif Central . Ch. Barthii and, only in the more southern sites, Ch. Mediterraneum occur as identifiable species . One of the best-known sites is in the southern Massif Central near Lodève in the Hérault department .

Chirotherium occurs in Italy especially in the Middle Triassic of the Dolomites, u. a. in the "Giovi formation" (anisium) near Bad Gfrill above the Etsch valley in South Tyrol and in the Richthofen conglomerate (anisium) of the Val Gerlano ( Trentino ). From the Upper Triassic (Carnian) of Monte Pisano in Tuscany , Friedrich von Huene described , among other traces, the species Chirotherium angustum , although it is questionable whether this classification is still justified according to the current definition of the genus, and if so, whether Ch. Angustum actually represents a species of its own.

Movement from the left manus and pes of “ Chirosaurus ibericus ” from the anisium of the Moncayo massif (Iberian Mountains, Spain).

In Spain, Chirotherium is mainly known from the Triassic of the Iberian Mountains . Among the traces that can be identified at the species level, there is apparently only Ch. Barthii , including the species " Chirosaurus " ibericus described in 1906 by the Jesuit naturalist Longino Navás from the anisium of the Moncayo massif (near Tarazona , province of Saragossa ) . The first find in Spain , originally referred to as Chirotherium , in 1897, made near Rillo de Gallo in the east of the province of Guadalajara , is now regarded as an indeterminable "Chirotheriide" trace. The species " Ch. Catalaunicum " described in 1979 from the early Anisium of the Catalan coastal cordillera near Barcelona , which was based on tetradactyls, i.e. four-toed tread seals and therefore did not correspond to the definition of the genus Chirotherium , has meanwhile been declared a noun dubium .

Traces from the so-called Labyrinthodontidae layers ( red sandstone equivalents, Olenekian) of the Świętokrzyskie Mountains in Poland, described in 1990 as Ch. Hauboldi , were subsequently assigned to the genus Brachychirotherium and finally to the genus Protochirotherium .

The trace Chirotherium peabodyi , described in 1958 from the lower Muschelkalk of the well-known fossil site Winterswijk in the east of the Netherlands , is not Chirotherium , but Rhynchosauroides .

Outside of Europe

North America

Most of the Chirotherium discovery sites outside Europe are in the USA. Here Chirotherium occurs in numerous localities in the Moenkopi Formation (Olenekium Anisium) of the Colorado Plateau in the states of New Mexico , Arizona and Utah . After the first reports about Chirotherium in the Moenkopi Formation in 1935, the well-known palaeontologist Frank Elmer Peabody differentiated a total of 8 different Chirotherium species in a comprehensive monograph in 1948 , 6 of which he himself described again. Half of these species come from the Wupatki subformation. In the meantime, four of these species have been placed in other genera, so that Ch. Barthii , Ch. Sickleri , Ch. Rex and Ch. Moquinense are still considered representatives of the genus Chirotherium in the Moenkopi trace fauna that can be identified at the species level .

Another significant occurrence of Chirotherium in North America is in the Paleorifts in the east of the USA in the Upper Triassic layers of the Newark supergroup , especially in the Passaic Formation of the Newark Basin, which is the stratigraphically youngest occurrence of the trace genus. Of the total of five under the generic name Chirotherium described or mentioned in older literature species now only are Ch. Huberi and Ch. Lulli left. The quarries near Milford in Hunterdon County , New Jersey, are among the most important sites there .

From the Upper Carboniferous of Pennsylvania Art "described Chirotherium " reiteri should not be a member of the genus because of their high geological age already.

South America

In South America, Chirotherium is so far only known from Argentina . There it occurs in the presumably Middle Triassic Cerro de las Cabras Formation (in older literature: Higueras Formation) of the Cuyo Basin ( Mendoza Province ). The traces were originally described as Ch. Higuerensis , but are now considered to belong to the type species Ch. Barthii .

Asia

In the People's Republic of China there are two trace sites with chirotherium in the Guanling Formation (Middle Triassic) in the southwest of Guizhou Province . In one of the two localities (Niuchang), discovered by locals in 1960, the traces were not examined by paleontologists until the late 1980s and identified as a chirotherium . Subsequent processing resulted in a classification in the species Ch. Barthii . In 2003 the second locality (Longchang) was finally discovered. In 2014, finds of chirotheriids from the Upper Triassic Baoding Formation in the Sichuan province were reported. Although the first toe is missing from the pes imprints (concave epireliefs), these traces are classified as cf.  Chirotherium and the absence of the first toe is interpreted as being due to the substrate and / or as an expression of a peculiarity in the walker.

Africa

In Africa, Chirotherium is known from the Aglegal subformation of the Timezgadiwine Formation (Lower Triassic) of the Argana Basin in the High Atlas of Morocco . In 1981 the Ichnospecies Chirotherium atlensis was described from Upper Triassic layers of the Argana Basin . Unfortunately, this is not mentioned in current specialist articles, so that it is unclear whether it is a Chirotherium at all , and if so, actually a separate Chirotherium species.

Classification and systematics

There are several ways to classify trace fossils. One is the so-called ethological classification, i. H. the trace is designated according to the activity of the producing organism. Tetrapod tracks are therefore part of the movement tracks or repichnia . For traces of animals that move with the help of limbs on a substrate (sediment), the more concrete term Schreitspuren or Cursichnia is used. Chirotherium is therefore a cursichnium.

Cursichnia and especially the footsteps and tracks of terrestrial vertebrates (Tetrapoda) have a very specific morphology which, by comparing them with body fossils of roughly the same age, enables a relatively precise identification of their producers. Therefore, classification schemes exist in tetrapod palichnology, which are based on the agreement of the trace morphology with the morphology of the extremities of certain tetrapod groups.

According to the six-part classification scheme for fossil reptile tracks by Franz Nopcsa, Chirotherium is classified among the crocodiloid tracks or tracks. In 1935, Othenio Abel established the Ichno family Chirotheriidae for Chirotherium and morphologically similar traces, which in the systematic part of palichnological treatises is still partly classified into higher-ranking reptile taxa like a real reptile family . The trace taxonomy , however, is a parallel taxonomy that is detached from the vertebrate system and a classification of the Chirotheriiden in z. B. the taxon Archosauria is inadmissible, although the trace producers may actually have been early Archosaurs (see → History and → Producer ). The German Palichnologists around Hartmut Haubold summarize Chirotherium together with morphologically similar trace taxa under the informal group term "Chirotherien" (English: chirotheres ).

Synonyms

On the basis of some trace specimens that are classified by today's palichnologists in the genus Chirotherium , separate trace genera (Ichnogenera) have been established in the past. In some cases, however, the synonymy only includes different spellings of the generic name originally used, since the international rules for zoological nomenclature that are binding today did not yet exist at the time. Therefore, in the scientific literature u. a. following synonyms for chirotherium :

  • Chirosaurus K AUP , 1835 (reserved "replacement name" for Chirotherium , actually used by Lydekker (1890) , among others )
  • Palaeopithecus V OIGT 1835
  • Cheirotherium S ICKLER 1836 (sic)
  • Cheirotherion N OPCSA 1923 (sic)
  • Krokodilipus N OPCSA 1923
  • Saurichnites K IRCHNER 1927

species

In the 19th and even at the beginning of the 20th century, the trace genus Chirotherium was not yet particularly precisely defined, with the result that numerous species from the various regions of Germany and the rest of the world were described, the type species Ch. Barthii morphologically sometimes very much and sometimes less similar. Some even looked very vaguely like her. As a result, the long list of Ichnospecies originally described as Chirotherium has shrunk considerably with more recent investigations , since many of these species have now either been synonymous with the species described first or have been assigned to other genera or the fossil specimens on which some of these species were based , showed such a bad state of conservation that those species are now considered to be completely invalid (noun dubium). Some names are also based simply on mistakes or misunderstandings (e.g. the name " Ch. Kaupii " probably denotes nothing more than "the chirotherium named by Kaup ", ie Ch. Barthii ) or inadmissible subsequent renaming (" Ch. Majus " for Ch. Barthii and " Ch. Minus " for Ch. Sickleri ). Since numerous Chirotherium species listed in the past have not yet been subjected to a thorough re-examination, the following list is likely to be too long. Problematic taxa are marked with an asterisk or provided with a note (see also section Occurrences ).

  • Chirotherium angustum VON H UENE 1941 *
  • Chirotherium atlensis B IRON & D UTUIT 1981 *
  • Chirotherium barthii K AUP 1835 = Chirosaurus ibericus N AVÁS 1906, Chirotherium bairdi R EIG 1961, Ch. Barthi (K AUP 1835) (sic), Ch. Gallicum W ILLRUTH 1917, Ch. Higuerensis R USCONI 1951, Ch. Ibericum N AVÁS 1906 , Ch. Ibericus (N AVÁS 1906) (sic), Ch. Kaupii O WEN 1842 (obj. Syn.), Ch. Majus S ICKLER 1836 (obj. Syn.), Saurichnites auraensis K IRCHNER 1927, S. gambachensis S CHUSTER 1936
  • Chirotherium culmbachense S OERGEL 1925 *
  • Chirotherium gigas P OHLIG 1893 *
  • Chirotherium hessei S OERGEL 1925 *
  • Chirotherium huberi B OCK 1952 *
  • Chirotherium lulli B OCK 1952
  • Chirotherium mediterraneum D EMATHIEU & D URAND 1991
  • Chirotherium moquinense P EABODY 1948 = Ch. Moquinensis P EABODY 1956 (sic), = Ch. Moquiensis P EABODY 1956 (sic)
  • Chirotherium reiteri M OORE 1873
  • Chirotherium rex P EABODY 1948
  • Chirotherium sickleri K AUP 1835 = Ch. Beasleyi N OPCSA 1932, Ch. Bipes B ERTHOLD 1835, Ch. Bornemanni W ILLRUTH 1917, Ch. Minus S ICKLER 1836 (obj. Syn.), Ch. Pfeifferi S OERGEL 1925
  • Chirotherium storetonense M ORTON 1862 = Ch. Beasleyi N OPCSA 1932 (pars), Chirosaurus stortonensis L YDEKKER 1890 (sic), Dinosaurichnium postchirotheroides R EHNELT 1949
  • Chirotherium vorbachi K IRCHNER 1927
  • Chirotherium wondrai H ELLER 1952


Species described as Chirotherium , which are now considered to be representatives of other Ichnos (including synonyms):

  • other "chirotheria" genera
    • Brachychirotherium B EURLEN 1950
      • B. eyermani (B AIRD 1957)
      • B. harrasense (H AUBOLD 1967)
      • B. lorteti (H AUBOLD 1970)
      • B. praeparvum (H AUBOLD 1967)
      • B. thuringiacum ( RÜHLE VON L ILIENSTERN 1939)
    • Isochirotherium H AUBOLD 1971
      • I. coltoni (P EABODY 1948)
      • I. coureli (D EMATHIEU 1970)
      • I. herculis (E Gerton 1838) = Chirotherium Barthi var. Herculis (E Gerton , 1838)
      • I. hessbergense (H AUBOLD 1971)
      • I. Jenense (H AUBOLD 1971)
      • I. lomasi (B AIRD 1954)
      • I. marshalli (P EABODY 1948)
      • I. Soergeli (H AUBOLD 1967)
    • Protochirotherium F ICHTER & K UNZ 2004
      • P. hauboldi (P TASZYŃSKI in F UGLEWICZ & al. 1990) = Brachychirotherium hauboldi (P TASZYŃSKI in F UGLEWICZ & al. 1990), Brachychirotherium wiorense P TASZYŃSKI 2000 Isochirotherium gierlinskii P TASZYŃSKI 2000 Isochirotherium sanctacrucense P TASZYŃSKI in F UGLEWICZ al & . 1990, Synaptichnium chirotherioides P TASZYŃSKI in F UGLEWICZ & al. 1990
    • Synaptichnium N OPCSA 1923
      • S. diabolense (P EABODY 1948) = Chirotherium diabolensis P EABODY 1956 (sic)
      • S. cameronense (P EABODY 1948) = Chirotherium cameronensis P EABODY 1956 (sic)
      • S. Hildburghausense ( RÜHLE VON L ILIENSTERN 1939)
  • non- “chirotheroid” trace genera
    • Dicynodontipus R ÜHLE OF L ILIENSTERN 1944
      • D. geinitzi (H ORNSTEIN 1876)
    • Herpetichnus J ARDIN 1850
      • H. pabsti (N OPCSA 1923)
      • H. rubrum (N OPCSA 1923)
    • Rhynchosauroides M AIDWELL 1911
      • Rhynchosauroides peabodyi (F BUT 1958)


Species classified as Chirotherium in older literature but not originally described as Chirotherium , which today are considered representatives of other Ichno genera (including synonyms):

  • Brachychirotherium parvum (H ITCHCOCK 1889), originally combined: Otozoum parvum (= Chirotherium copei B OCK 1952)
  • Limnopus heterodactylus (K ING 1845), originally Comb .: Thenaropus heterodactylus
  • Paratetrasauropus swinnertoni (S ARJEANT 1970), originally combined: Otozoum swinnertoni

Noun dubia :

  • Chirotherium bipedale A BEL 1935
  • Chirotherium catalaunicum C ASANOVAS -C LADELLAS 1979 et al.
  • Chirotherium fucinii ( VON H UENE 1941), originally combined: Thecodontichnus fucinii

Producer

Batrachotomus kupferzellensis from the Keuper of Baden-Württemberg, possibly a close relative of the Chirotherium producers. The protruding 5th toe is relatively easy to see.
Skeleton of the right foot of Euparkeria capensis  (A) with corresponding drawing (B). This representative also has a protruding 5th toe.

Today it is generally accepted that chirotherium was caused by reptiles belonging to the archosaur line that leads to today's crocodiles ( Crurotarsi ), with representatives from the group of "Rauisuchia" being considered the safest candidates. The problem here, however, is the fact that there are usually few or no body fossils in trace layers. So far, no direct association between Chirotherium and Rauisuchier body fossils has been found anywhere in the world . Which known "Rauisuchier" of the Triassic could be responsible for certain occurrences of chirotheria in certain regions of the world has therefore been derived exclusively from indirect evidence:

  • The first “Rauisuchier” that was specifically associated with Chirotherium is Ticinosuchus . So far, however, its remains are only known from marine sediments ; That is, there is no direct connection to the mainland deposits in which Chirotherium occurs. However, due to the anatomy of its limbs , Ticinosuchus was able to generate “chirotheriide” tracks and the deposits in which it was found are about as old as many chirotherium- bearing layers.
  • For a special, large variant of the Ichnospecies Ch. Sickleri , as z. B. was found in Hardheim, is believed to be the producer of the "Rauisuchier" Ctenosauriscus or a closely related species. Corresponding bone remnants have been excavated in the Röt von Waldshut along with a few Chirotherium footsteps (although the Ch. Sickleri is not mentioned ).
  • Batrachotomus or a closely related genus is traded as another polluter candidate for chirotherium in Central England and Germany . Bone remnants suggesting such a relationship come from the Bromsgrove sandstone, a stratigraphic equivalent of the Helsby sandstone, near Warwick in Central England.
  • Also Arizonasaurus applies since its detection in the trace range Moenkopi Formation as a likely Chirotherium Generating Sets.
  • As a producer of the southern French track Ch. Mediterraneum is Euparkeria (no "Rauisuchier" but a core group representatives of Avesuchia ) accepted. The same problems exist here as with Soergel and Ch. Barthii (see →  history ).
Chirotherium Monument in Hildburghausen with a reconstruction of the original trace area in the background and a bronze sculpture of a reconstruction of the trace generator. The position of the left extremities corresponds to the typical Manus-Pes sentence of Chirotherium .

A purely hypothetical living reconstruction of the trace producer by Ch. Barthii can be found as part of the Chirotherium monument in Hildburghausen. It is a " chimera " of Saurosuchus and Euparkeria .

With regard to the chirotherium associations from the Storeton and Heßberg quarries, which both contain the small form Ch. Sickleri and a large form, Ch. Storetonense and Ch. Barthii , respectively , it has been speculated that the small and large forms are respectively the represent the same producer species but have been caused by individuals in different growth stages. The relatively small footsteps of Ch. Sickleri were therefore produced by young animals, the large footsteps of Ch. Storetonense or Ch. Barthii by adult individuals. A certain morphological variation within the tracks of Ch. Storetonenese and Ch. Barthii is also seen as an expression of a sexual dimorphism , whereby the somewhat smaller, more delicate step seals are said to come from females, the larger, more strongly developed step seals from males.

Biostratigraphic meaning

Chirotherium is the index taxon of a late Olenek-Early Anise biochron, which is also characterized by the trace taxa Rotodactylus , Isochirotherium and Synaptichnium (including a variant previously classified as " Brachychirotherium "). The use of tetrapod trace biostratigraphy is particularly recommended in mainland sediments with little or no body fossils, although the temporal resolution achieved is less than that of biostratigraphy using body fossils.

Artistic reception

On the occasion of the first chirotherium finds on the Heßberg near Hildburghausen, the poet Eduard Mörike , who had a penchant for “petrefactual”, wrote the following poem at the request of the paleontologist Albert Oppel :

“Poetic etiquette for Chirotherium Kaupii
(from the Heßberg sand quarries ) Whether a giant frog or a marsupial was unfortunately not yet to be fathomed; We would have the clear trail, But only an Oppel will find the beast himself. If it finally came to a batrachum , it might not be called bad iambicus , because if the frog doesn't jump, rather just walks, he puts the small forefoot in front of him first , he puts the hind foot on the same side after, the one that is much larger, just like figura shows, and thus has the regular iambic gait. An older sample of this verse can hardly be found than in Hessberg's Steinkodizibus. Today's frogs are known to move, our poetic ones , out of innate instinct in these dimensions with particular ease. For my part, Doctor, I would gladly give away without delay for a single Antediluvian Batrachier of the current species, preserved only in an emergency, the whole host of the newest Diluvii, which covers Parnassus from foot to summit, and all of my iambs cheaply. "























- Eduard Mörike : Complete Works. Second volume

Remarks

  1. ↑ In the zoological nomenclature, the word parts Thero- , -therium or -theria traditionally refer to mammals in some way. If a genus or species is originally considered a mammal and is therefore named accordingly, and if it later turns out that this classification was incorrect, the International Rules for Zoological Nomenclature require that the original name be retained.
  2. ↑ The term "Scheinkrokodil", which is often mentioned in popular or non-scientific writings on the subject of chirotherium, is the German translation of the word Pseudosuchia . In the meantime, the taxon Pseudosuchia has been redefined as a clade and its content roughly corresponds to the Crurotarsi (the latter name is less misleading, as this taxon also includes the actual crocodiles , Crocodylia). Therefore, one can still say that the chirotherium producer is a pseudosearcher, but should then put the word “basal” in front of the taxon name. For more information on the current systematic position of the Chirotherium producers, see the Producer section .
  3. ↑ In older literature these layers are often referred to as “Keuper” (or Cuyper), which goes back to the nomenclature of Edward Hull (1869) for the Triassic Central England. As far as we know today, the Helsby and Tarporley Formations are equivalent to the higher red sandstone and not to the Keuper of Central Europe.
  4. ↑ Haubold carried out the initial “ splitting ” of the genre in 1971.
  5. ↑ French and Spanish authors do not regard Ch. Gallicum as a synonym of Ch. Barthii , but as a representative of the trace genus Brachychirotherium .
  6. ↑ King et al. (2005) do not consider Ch. Gambachensis to be a synonym of Ch. Barthii .
  7. ↑ Comes from the Upper Carboniferous and is therefore very likely not a Chirotherium species.
  8. ↑ King et al. (2005) do not consider Ch. Pfeifferi to be a synonym of Ch. Sickleri , but of Ch. Barthii .
  9. ↑ King et al. (2005) consider the type material of I. herculis as a representative of a non-identifiable Chirotherium species, and thus I. herculis de facto as a noun dubium. However, Clark and Corrence (2009) consider the species to be valid as defined by Haubold (1971)
  10. ↑ King et al. (2005) consider the type material by O. swinnertoni as a representative of Chirotherium , but not as identifiable at the species level, as they do not list it under the name “ Chirotherium swinnertoni ” among the Chirotherium species recognized by them, and thus de facto as a noun dubium.

Individual evidence

  1. ↑ a b c d Johann J. Kaup: Mittheilungen, addressed to Professor Bronn. Thier tracks from Hildburghausen: Chirotherium or Chirosaurus. New yearbook for mineralogy, geognosy, geology and petrefacts. Year 1835, pp. 327-328, online
  2. ↑ a b c d e Fabien Knoll: Alexander von Humboldt and the hand-beast: A contribution to palaeontology from the last universal scholar. Comptes Rendus Palevol. Vol. 8, No. 4, 2009, pp. 427-436, doi : 10.1016 / j.crpv.2008.12.001
  3. ^ Friedrich KL Sickler: Letter to Sr. Hochwohlgeboren [...] Dr. JF Blumenbach [...] on the most remarkable reliefs, only discovered a few months ago, showing the tracks of primeval, large and unknown animals in the Hessberg sandstone quarries near the city of Hildburghausen. Kesselringsche Hofbuchhandlung, Hildburghausen 1834, 16 p., Urn : nbn: de: bvb: 12-bsb10231937-5
  4. ^ A b c Heinrich G. Bronn: Latest literature - excerpts: III. Petrefacts. New yearbook for mineralogy, geognosy, geology and petrefacts. Year 1835, pp. 230-234, online
  5. ↑ a b J. Jacob Nöggerath: Alleged tracks of primeval monkeys (Quadrumans) in sandstones. Non-profit and entertaining Rheinische Provinzial-Blätter, new series. Second year, first volume, second issue, 1835, pp. 143–148, limited preview in the Google book search
  6. ^ Ernst Probst: Johann Jakob Kaup: the great natural scientist from Darmstadt. GRIN-Verlag, 2011, p. 103, ISBN 978-3-640-84916-1
  7. ^ J. Grierson: On Footsteps before the Flood, in a specimen of Red Sandstone. The Edinburgh Journal of Science. Vol. 8, pp. 130-133, online
  8. ↑ H. Girard: About the tracks of pre-worldly animals in the sandstone, especially by Chirotherium. New yearbook for mineralogy, geognosy, geology and petrefacts. Jhrg. 1846, pp. 1-22, online
  9. ↑ Richard Owen: Description of parts of the Skeleton and Teeth of five species of the Genus Labyrinthodon (Lab.leptognathus, Lab.pachygnathus, and Lab.ventricosus , from the Coton-end and Cubbington Quarries of the Lower Warwick Sandstone; Lab. Jaegeri , from Guy's Cliff, Warwick; and Lab. scutulatus , from Leamington); with remarks on the probable identity of the Cheirotherium with this genus of extinct Batrachians. Pp. 515-543 (536 ff.), Online
  10. ↑ DMS Watson: The Cheirotherium. The Geological Magazine, New Series. Dekade 6, Vol. 1, No. 9, 1914, pp. 395-398, online
  11. ^ A b A. J. Bowden, GR Tresise, W. Simkiss: Chirotherium , the Liverpool footprint hunters and their interpretation of the Middle Trias environment. In: RTJ Moody, E. Buffetaut, D. Naish, DM Martill (Eds.): Dinosaurs and Other Extinct Saurians: A Historical Perspective. Geological Society, London, Special Publications. Vol. 343, 2010, pp. 209-228, doi : 10.1144 / SP343.12
  12. ^ Adolf Seilacher: Trace Fossil Analysis. Springer-Verlag, 2007, pp. 6/7, ISBN 978-3-540-47225-4
  13. ↑ a b Klaus Ebel, Franz Falkenstein, Frank-Otto Haderer, Rupert Wild: Ctenosauriscus koeneni (by Huene) and the Rauisuchier von Waldshut - Biomechanical interpretation of the spine and relationships to Chirotherium sickleri Kaup. Stuttgart Contributions to Natural History, Series B., No. 261, 1998, online
  14. ↑ a b c d e f Hendrik Klein, Hartmut Haubold: Differentiation of selected chirotheries of the Triassic by means of landmark analysis. Hallesches Jahrbuch für Geoswissenschaften. Vol. B 25, 2003, pp. 21-36
  15. ^ Georges Demathieu, Pierre Demathieu: Chirotheria and Other Ichnotaxa of the European Triassic. Ichnos. Vol. 11, No. 1-2, 2004, pp. 79-88, doi : 10.1080 / 10420940490444898
  16. ↑ a b Hartmut Haubold: Die Saurierfährten Chirotherium barthii Kaup, 1835 - the type material from the red sandstone near Hildburghausen / Thuringia and the "Chirotherium Monument". Publications of the Natural History Museum Schleusingen. Vol. 21, 2006, pp. 3-31
  17. ^ A b Georges Demathieu, Jürgen Fichter: The Karlshafener Fährten in the natural history museum of the city of Kassel. Philippia. Treatises and reports from the Natural History Museum in the Ottoneum in Kassel. Vol. 6, No. 2, 1989, pp. 111–154, online (PDF; 7.9 MB)
  18. ↑ Daniel Krause, Hartmut Haubold: The tracking area near Eiterfeld (East Hesse, Fulda district), in the chirotherium layers of the Solling series of the red sandstone. Geological Yearbook Hessen. Vol. 135, 2007, pp. 61-100
  19. ↑ Ludwig Rumpf: track prints from the colorful sandstone to Aura on the Saale; living frog in the shell limestone near Höchberg; Fragment of an antler from the Steinberge near Würzburg; Trigonotreta fragilis and Placodus gigas Agass. from the local shell limestone; Dolomite as the uppermost link of the Muschelkalk formation belongs to the Keuper structure. New Yearbook for Mineralogy, Geology and Paleontology, 1842, pp. 450–451 ( online ), panel VIII A ( online ; note: the traces shown therein are atypical and offer a fitting example of so-called extramorphological (ie substrate and / or conservation-related) variation )
  20. ↑ Ludwig Rumpf: About animal tracks in the colorful sandstone near Aura; Horseshoe-like impressions in the colored sandstone of Elfershausen. New Yearbook for Mineralogy, Geology and Paleontology, 1843, pp. 705–707 Archives
  21. ↑ W. Freudenberger, G. Geyer, B. Schröder: The red sandstone in Bavaria (north-western Franconia, quarry land and edge facies in the subsoil). In: German Stratigraphic Commission (Ed .; coordination and editing: J. Lepper & H.-G. Röhling for the sub-commission Perm-Trias): Stratigraphie von Deutschland XI. Red sandstone. Series of publications of the German Society for Geosciences. Vol. 69, 2013, pp. 547-582
  22. ↑ Heinrich Kirchner: About the animal tracks in the upper Buntsandstein Franconia. Paleontological Journal. Vol. 9, No. 1-3, 1927, pp. 112-122
  23. ↑ a b Hartmut Haubold: Expert opinion on the track find near Gössenheim (Main-Spessart), with preliminary determination. Halle / S., 2011, online (PDF; 1.6 MB)
  24. ↑ a b c d e Frank-Otto Haderer, Georges Demathieu, Ronald Böttcher: Vertebrate tracks from the red quartzite (Upper Buntsandstein, Middle Triassic) from Hardheim near Wertheim / Main (southern Germany). Stuttgart Contributions to Natural History, Series B. No. 230, 1995, online
  25. ↑ Annual reports of the State Museum for Natural History Stuttgart for the years 1989 , 1991 and 1992 , published in the annual journals of the Society for Natural History in Württemberg, volumes 145, 147 u. 148 (1990, 1992, 1993)
  26. ↑ 240 million year old Chirotherien track discovered in the Odenwald. dinosaurier-interesse.de, April 14, 2007 (last accessed June 29, 2013)
  27. ↑ a b Frank-Otto Haderer, Sven Sachs: A tracking plate with Chirotherium barthii Kaup and cf. Rotodactylus from the Upper Buntsandstein (Lower Triassic) of Pirmasens. Communications from POLLICHIA. Vol. 96, 2012, pp. 5-10
  28. ^ Cajus G. Dietrich: Middle Triassic chirotherid trackways on earthquake influenced intertidal limulid reproduction flats of the European Germanic Basin coasts. Central European Journal of Geosciences. Vol. 4, No. 3, 2012, pp. 495-529, doi : 10.2478 / s13533-011-0080-9
  29. ↑ Hendrik Klein, Hartmut Haubold: Tradition-related variation in chirotheria and notes on Ichnotaxonomy based on examples from the Middle to Upper Triassic (Anisium-Carnian) of Northern Bavaria. Hallesches Jahrbuch für Geoswissenschaften. Bd. B 26, 2004, pp. 1-16
  30. ↑ a b c d e f g h i j k l m Michael J. King, William AS Sarjeant, David B. Thompson, Geoffrey Tresise: A Revised Systematic Ichnotaxonomy and Review of the Vertebrate Footprint Ichnofamily Chirotheriidae from the British Triassic. Ichnos. Vol. 12, No. 4, 2005, pp. 241-299, doi : 10.1080 / 10420940591009312
  31. ^ Mike Batty: Chirotherium and Its Domain: A Redescription of Rediscovered Specimens from Northwest England. The Geological Curator. Vol. 8, No. 9, 2008, 437–454, online (complete issue; PDF; 5.9 MB)
  32. ^ NDL Clark, P. Aspen, H. Corrance: Chirotherium barthii Kaup 1835 from the Triassic of the Isle of Arran, Scotland. Scottish Journal of Geology. Vol. 38, No. 2, 2002, pp. 83-92, online (digital reprint; PDF; 18.2 MB)
  33. ^ A b N. DL Clark, H. Corrance: New discoveries of Isochirotherium herculis (Egerton 1838) and a reassessment of chirotheriid footprints from the Triassic of the Isle of Arran, Scotland. Scottish Journal of Geology. Vol. 45, No. 1, 2009, pp. 69–82, online (digital reprint; PDF; 812 kB)
  34. ↑ Geoffrey Tresise: 'Chirotherium herculis' - problem Posed by the First Find. Annals of Science. Vol. 48, No. 6, 1991, pp. 565-576, doi : 10.1080 / 00033799100200451
  35. ^ George Demathieu, Georges Gand: Les sites à traces de pas de vertébrés du Trias à l'Hettangien. Contenu et interprétation. Le Naturaliste Vendéen. Vol. 3, 2003, pp. 47–53 online (PDF; 263 kB)
  36. ↑ Diana Valdiserri, Marco Avanzini: A tetrapod Ichnoassociation from the Middle Triassic (Anisian, Pelsonian) of Northern Italy. Ichnos. Vol. 14, No. 1-2, 2007, pp. 105-116, doi : 10.1080 / 10420940601010703
  37. ^ Rossana Todesco, Massimo Bernardi: Una nuova icnoassociazione a vertebrati nel Triassico medio (Anisico) del Trentino meridionale (Val Gerlano, Vallarsa). Studi Trentini di Scienze Naturali. Vol. 88, pp. 203-218, online (PDF; 321 kB)
  38. ^ Friedrich von Huene: The tetrapod tracks in the Tuscan Verrucano and their meaning. New Yearbook for Mineralogy, Geology and Paleontology, Section B, supplement volume. Jhrg. 1941, pp. 1-34
  39. ^ RP Longino Navás: El Chirosaurus ibericus sp. nov. Boletín de la Sociedad Aragonesa de Ciencias Naturales. Vol. 5, 1906, pp. 208-213, online
  40. ↑ Ignacio Díaz-Martínez, Diego Castanera, José Manuel Gasca, José Ignacio Canudo: A reappraisal of the Middle Triassic chirotheriid Chirotherium ibericus Navás, 1906 (Iberian Range NE Spain), with comments on the Triassic tetrapod track biochronology of the Iberian Peninsula. PeerJ. Vol. 3, 2015, e1044, doi : 10.7717 / peerj.1044 ( Open Access )
  41. ↑ Ignacio Díaz-Martínez, Adán Pérez-García: Historical and Comparative Study of the First Spanish Vertebrate Paleoichnological Record and Bibliographic Review of the Spanish Chirotheriid Footprints. Ichnos. Vol. 19. No. 3, 2011, pp. 141-149, doi : 10.1080 / 10420940.2012.685565
  42. ↑ a b J. Fortuny, A. Bolet, AG Sellés, J. Cartanyà, À. Galobart: New insights on the Permian and Triassic vertebrates from the Iberian Peninsula with emphasis on the Pyrenean and Catalonian basins. Journal of Iberian Geology. Vol. 37, No. 1, 2011, pp. 65–86, doi : 10.5209 / rev_JIGE.2011.v37.n1.5
  43. ↑ Ryszard Fuglewicz, Tadeusz PTASZYNSKI, and Kazimierz Rdzanek: Lower Triassic footprints from the Świętokrzyskie (Holy Cross) Mountains, Poland. Acta Palaeontologica Polonica. Vol. 35, No. 3-4, 1990, pp. 109-164, online
  44. ^ Tadeusz Ptaszyński: Lower Triassic vertebrate footprints from Wióry, Holy Cross Mountains, Poland. Acta Palaeontologica Polonica. Vol. 45, No. 2, 2000, pp. 151-194, online
  45. ↑ a b Hendrik Klein, Grzegorz Niedźwiedzki: Revision of the Lower Triassic Tetrapod Ichnofauna from Wióry, Holy Cross Mountains, Poland. New Mexico Museum of Natural History and Science Bulletin. Vol. 56, 2012, online
  46. ↑ a b H. W. Oosterink: Fossiele voetstappen in de eerste Muschelkalk-groeve van Winterswijk. Grondboor & Hamer. Vol. 30, No. 5, 1976, pp. 130-144, online
  47. ↑ a b c d e f Michael Morales: Terrestrial Fauna and Flora from the Triassic Moenkopi Formation of the Southwestern United States. In: M. Morales, DK Elliot (Ed.): Triassic continental deposits of the American Southwest. Proceedings of a Special Symposium held at the Museum of Northern Arizona, Fragstaff, Arizona, USA, September 21-22, 1985. Journal of the Arizona-Nevada Academy of Science. Vol. 22, No. 1, 1987, p. 11
  48. ^ Paul E. Olsen, Hans-Dieter Sues: Correlation of continental Late Triassic and Early Jurassic sediments, and patterns of the Triassic-Jurassic tetrapod transition. In: K. Padian (Ed.): The Beginning of the Age of Dinosaurs. Faunal Change Across the Triassic-Jurassic Boundary. Cambridge University Press, New York 1986, pp. 321-351, ISBN 0-521-30328-1
  49. ^ A b c Earle E. Spamer, Edward Daeschler, L. Gay Vostreys-Shapiro: A Study of Fossil Vertebrate Types in the Academy of Natural Sciences of Philadelphia. The Academy of Natural Sciences of Philadelphia Special Publication. Vol. 16, 1995, pp. 287-293
  50. ^ Ricardo N. Melchor, Silvina de Valais: A Review of Triassic Tetrapod Track Assemblages from Argentina. Paleontology. Vol. 46, No. 2, pp. 355-372, doi : 10.1111 / j.1475-4983.2006.00538.x
  51. ↑ Lida Xing, Hendrik Klein, Martin G. Lockley, Jianjun Li, Jianping Zhang, Masaki Matsukawa, Jiafei Xiao: Chirotherium Trackways from the Middle Triassic of Guizhou, China. Ichnos. Vol. 20, No. 2, 2013, pp. 99-107, doi : 10.1080 / 10420940.2013.788505
  52. ^ Lida Xing, Hendrik Klein, Martin G. Lockley, Zezhong Kan, Jianping Zhang, Guangzhao Peng, Yong Ye: First chirothere and possible grallatorid footprint assemblage from the Upper Triassic Baoding Formation of Sichuan Province, southwestern China. Palaeogeography, Palaeoclimatology, Palaeoecology. Vol. 412, 2014, pp. 169–176, doi : 10.1016 / j.palaeo.2014.07.032
  53. ↑ Hendrik Klein, Sebastian Voigt, Hafid Saber, Jörg W. Schneider, Abdelkbir Hminna, Jan Fischer, Abdelouahed Lagnaoui, Andreas Brosig: First occurrence of a Middle Triassic tetrapod ichnofauna from the Argana Basin (Western High Atlas, Morocco). Palaeogeography, Palaeoclimatology, Palaeoecology. Vol. 307, No. 1–4, 2011, pp. 218-231, doi : 10.1016 / j.palaeo.2011.05.021
  54. ↑ a b c Hendrik Klein, Hartmut Haubold: Archosaur Footprints - Potential for Biochronology of Triassic Continental Sequences. In: Spencer G. Lucas, Justin A. Spielmann (Eds.): The Global Triassic. New Mexico Museum of Natural History and Science Bulletin. Vol. 41, pp. 120-130, online
  55. ^ Richard Lydekker: Catalog of the Fossil Reptilia and Amphibia in the British Museum (Natural History). Part IV. London 1890, pp. 215 ff., Online
  56. ^ WE Swinton: The history of Chirotherium. Geological Journal. Vol. 2, No. 3, 1961, pp. 443-473, doi : 10.1002 / gj.3350020309
  57. ^ Adrian P. Hunt, Spencer G. Lucas: Late Triassic Tetrapod Tracks of Western North America. In: Spencer G. Lucas, Justin A. Spielmann (Eds.): Triassic of the American West. New Mexico Museum of Natural History and Science Bulletin. Vol. 40, 2007, pp. 215-230, online
  58. ↑ Georges Gand, Raúl De La Horra, Belén Galán-Abellán, José López-Gómez, José F. Barrenechea, Alfredo Arche, M. Isabel Benito: New ichnites from the Middle Triassic of the Iberian Ranges (Spain): paleoenvironmental and paleogeographical implications . Historical Biology. Vol. 22, No. 1-3, 2010, pp. 40-56, doi : 10.1080 / 08912961003644096
  59. ↑ a b Hartmut Haubold: Tetrapod tracks from the Permian - state of knowledge and progress 2000. Hallesches Jahrbuch für Geoswissenschaften. Vol. B 22, 2000, pp. 1-16
  60. ↑ Oliver Perry Hay: Bibliography and Catalog of the Fossil Vertebrata of North America. Washington 1902, p. 512, online
  61. ↑ Sebastian Voigt: The Tetrapodenichnofauna of the continental Upper Carboniferous and Permian in the Thuringian Forest - Ichnotaxonomy, Paleoecology and Biostratigraphy. Göttingen 2005, pp. 77 f., ISBN 3-86537-432-8 .
  62. ↑ Chirotherium . Datasheet in the Paleobiology Database
  63. ↑ Chirotheriidae. Datasheet in the Paleobiology Database
  64. ↑ Peter M. Galton: Notes on bones and teeth of Bromsgroveia and other archosaurian reptiles from the lower Middle Triassic (Anisian) of the English Midlands. Revue de Paléobiologie. Vol. 31, No. 1, 2012, pp. 171–204 online (PDF; 3.1 MB)
  65. ^ Louis Courel, Georges Demathieu: Tentative Correlation Using Ichnological data from Continental Sandstone Series and Marine Faunas in the Middle Triassic of Europe. Albertiana. Vol. 15, 1995, pp. 83-91
  66. ↑ Geoffrey Tresise: Sex in the footprint bed. Geology Today. Vol. 12, No. 1, 1996, pp. 22-26, doi : 10.1046 / j.1365-2451.1996.00005.x
  67. ↑ Hendrik Klein, Spencer G. Lucas: Tetrapod footprints - their use in biostratigraphy and biochronology of the Triassic. In: Spencer G. Lucas (Ed.): The Triassic Timescale. Geological Society, London, Special Publications. Vol. 334, 2010, pp. 419-446, doi : 10.1144 / SP334.14
  68. ^ Eduard Mörike: Complete Works. Second volume: Poetry gleanings, idyll from Lake Constance, Wispeliaden. Published by the art warden by Karl Fischer. Callwey-Verlag, Munich around 1906, pp. 125/126 online

Web links

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