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''Chirotherium'', also known as ''Cheirotherium'' (‘hand-beast’), is a
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest per ...
trace fossil A trace fossil, also known as an ichnofossil (; from el, ἴχνος ''ikhnos'' "trace, track"), is a fossil record of biological activity but not the preserved remains of the plant or animal itself. Trace fossils contrast with body fossils, ...
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 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 silicat ...
(
Buntsandstein The Buntsandstein (German for ''coloured'' or ''colourful sandstone'') or Bunter sandstone is a lithostratigraphic and allostratigraphic unit (a sequence of rock strata) in the subsurface of large parts of west and central Europe. The Buntsandst ...
) in
Thuringia Thuringia (; german: Thüringen ), officially the Free State of Thuringia ( ), is a state of central Germany, covering , the sixth smallest of the sixteen German states. It has a population of about 2.1 million. Erfurt is the capital and lar ...
,
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 betwee ...
, dating from about 243
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago ...
( mya). The creatures who made the footprints and tracks were probably
pseudosuchia Pseudosuchia is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely related to crocodilians than to birds. Pseudosuchians are also informally known as "crocodilian-line archosaurs". Prior to ...
n
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avia ...
s related to the ancestors of the
crocodile Crocodiles (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 even more loosely to include all extant me ...
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 country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea to the southwest. It is separated from continental Europe ...
in 1838. They were found before
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
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 any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in ...
. These
fossil track A fossil track or ichnite (Greek "''ιχνιον''" (''ichnion'') – a track, trace or footstep) is a fossilized footprint. This is a type of trace fossil. A fossil trackway is a sequence of fossil tracks left by a single organism. Over the yea ...
s have now been found on
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
,
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the List of South American countries by area, second-largest ...
,
North Africa North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region, and it is sometimes defined as stretching from the Atlantic shores of Mauritania in ...
,
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
, and
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
. British paleontologist
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. Ow ...
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 four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbo ...
. 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 gender difference (
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...
) 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 marine and brackish water arthropods of the family Limulidae and the only living members of the order Xiphosura. Despite their name, they are not true crabs or crustaceans: they are chelicerates, most closely related to ar ...
s. The horseshoe crabs were likely breeding along the
intertidal zone The intertidal zone, also known as the foreshore, is the area above water level at low tide and underwater at high tide (in other words, the area within the tidal range). This area can include several types of habitats with various species ...
while the ''Chirotherium'' trackmaker preyed on them during low tide. Smaller reptiles like ''
Macrocnemus ''Macrocnemus'' is an extinct genus of archosauromorph reptile known from the Middle Triassic (Late Anisian to Ladinian) of Europe and China. ''Macrocnemus'' is a member of the Tanystropheidae family and includes three species''. Macrocnemus ba ...
'', represented by the ichnogenus ''
Rhynchosauroides ''Rhynchosauroides'' is an ichnogenus, a form taxon based on footprints. The organism producing the footprints was likely a lepidosaur and may have been a sphenodont, an ancestor of the modern tuatara. The footprint consists of five digits, of ...
'', likely fed on the horseshoe crabs’ eggs.


See also

*
Ichnite A fossil track or ichnite ( Greek "''ιχνιον''" (''ichnion'') – a track, trace or footstep) is a fossilized footprint. This is a type of trace fossil. A fossil trackway is a sequence of fossil tracks left by a single organism. Over the y ...


References

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