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''Chirotherium'', also known as ''Cheirotherium'' (‘hand-beast’), is a
Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
trace fossil A trace fossil, also called an ichnofossil (; ), is a fossil record of biological activity by lifeforms, but not the preserved remains of the organism itself. Trace fossils contrast with body fossils, which are the fossilized remains of part ...
consisting of five-fingered (pentadactyle) footprints and whole tracks. These look, by coincidence, remarkably like the hands of apes and bears, with the outermost toe having evolved to extend out to the side like a thumb, although probably only functioning to provide a firmer grip in mud. ''Chirotherium'' tracks were first found in 1834 in Lower Triassic
sandstone Sandstone is a Clastic rock#Sedimentary clastic rocks, clastic sedimentary rock composed mainly of grain size, sand-sized (0.0625 to 2 mm) silicate mineral, silicate grains, Cementation (geology), cemented together by another mineral. Sand ...
(
Buntsandstein The Buntsandstein (German for ''coloured'' or ''colourful sandstone'') or Bunter sandstone is a lithostratigraphy, lithostratigraphic and allostratigraphy, allostratigraphic unit (a sequence of rock strata) in the Subsurface (geology), subsurface ...
) in
Thuringia Thuringia (; officially the Free State of Thuringia, ) is one of Germany, Germany's 16 States of Germany, states. With 2.1 million people, it is 12th-largest by population, and with 16,171 square kilometers, it is 11th-largest in area. Er ...
,
Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
, dating from about 243
million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...
( mya). The creatures who made the footprints and tracks were probably
pseudosuchia Pseudosuchia, from Ancient Greek ψεύδος (''pseúdos)'', meaning "false", and σούχος (''soúkhos''), meaning "crocodile" is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely relat ...
n
archosaur Archosauria () or archosaurs () is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant taxon, extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistics ...
s related to the ancestors of the
crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large, semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term "crocodile" is sometimes used more loosely to include ...
s. They likely belonged to either prestosuchidae or rauisuchidae groups, which were both large carnivores with semi-erect gaits.


History

''Chirotherium'' tracks were first found in German Triassic sandstones in 1834, and later in
England England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...
in 1838. They were found before
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s were known and initial models of the trackmaker proposed that it was a bear or ape, which walked with its feet crossed. This proposal was necessary to explain the toe on the outside. The tracks were also proposed to be from a
marsupial Marsupials are a diverse group of mammals belonging to the infraclass Marsupialia. They are natively found in Australasia, Wallacea, and the Americas. One of marsupials' unique features is their reproductive strategy: the young are born in a r ...
. These fossil tracks have now been found on
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
,
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
,
North Africa North Africa (sometimes Northern Africa) is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region. However, it is sometimes defined as stretching from the Atlantic shores of t ...
,
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
, and
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
. British paleontologist
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomy, comparative anatomist and paleontology, palaeontologist. Owen is generally considered to have been an outstanding naturalist with a remarkabl ...
suggested in 1842 that the tracks were made by a
labyrinthodont "Labyrinthodontia" (Greek, 'maze-toothed') is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras (about 390 to 150 million years ago). Traditionally conside ...
amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
. Over the following years, new discoveries of archosaurian reptiles indicated that ''Chirotherium'' tracks were made by a pseudosuchian. The print's resemblance to mammals was only superficial; in reality, an external (lateral) ‘thumb’ was commonplace among Triassic archosaurs. In 1965, the skeleton of an animal probably closely related to the trackmaker of ''Chirotherium'' was found, called '' Ticinosuchus''. It had the external toe on its hind feet but not on its front feet and was possibly a more derived descendant, whose gait did not require a stabilizing front toe. Footprints of different size and proportions occurring together on one and the same bedding plane probably reflect a sex difference (
sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
) within the trackmaker species. Other Chirotheriidae include '' Isocheirotherium''.


Paleobiology

''Chirotherium'' trackways have been found in German sandstones that were likely deposited on flood plains. During the Middle Triassic, much of Central Europe was covered by a shallow epicontinental sea (the so-called Muschelkalk Sea). In one location, ''Chirotherium'' trackways were found alongside those of early
horseshoe crab Horseshoe crabs are arthropods of the family Limulidae and the only surviving xiphosurans. Despite their name, they are not true crabs or even crustaceans; they are chelicerates, more closely related to arachnids like spiders, ticks, and scor ...
s. The horseshoe crabs were likely breeding along the
intertidal zone The intertidal zone or foreshore is the area above water level at low tide and underwater at high tide; in other words, it is the part of the littoral zone within the tidal range. This area can include several types of habitats with various ...
while the ''Chirotherium'' trackmaker preyed on them during low tide. Smaller reptiles like '' Macrocnemus'', represented by the ichnogenus '' Rhynchosauroides'', likely fed on the horseshoe crabs’ eggs.


See also

* Ichnite


References

{{Taxonbar, from=Q311020 Vertebrate trace fossils Fossil trackways