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'''''Gwyneddichnium''''' is an [[Ichnotaxon|ichnogenus]] from the [[Late Triassic]] of North America and Europe. It represents a form of reptile footprints and trackways, likely produced by small [[Tanystropheidae|tanystropheids]] such as ''[[Tanytrachelos]]''. ''Gwyneddichnium'' includes a single species, '''''Gwyneddichnium major''''' (also spelled ''G. majore''). Two other proposed species, ''G. elongatum'' and ''G. minore'', are indistinguishable from ''G. major'' apart from their smaller size and minor [[Taphonomy|taphonomic]] discrepancies. As a result, they are considered junior synonyms of ''G. major''.<ref name=":0">Liquid error: wrong number of arguments (given 1, expected 2)</ref>
== Description ==
''Gwyneddichnium'' corresponds to footprints from an a quadrupedal animal with a small pentadactyl (five-fingered) manus (hand) and a notably larger five-toed pes (foot). The manus and pes are mesaxonic, meaning that the third digit is the longest digit, followed by the subequal second and fourth digits. The innermost digit (digit I) and the outermost digit (digit V) were short and located close to the rest of the foot. Sometimes the small fifth digit is poorly preserved, making the hand or foot appear to be tetradactyl (having only four digits). Overall, the digits are long and narrow, with minimal curvature. There is some variation with how the digits are positioned, with some specimens having digits which evenly radiate away from the sole,<ref name=":0" /> and others having digits which separate into two clumps comprised of digits I-III and IV-V.<ref name=":1" />
The skin made pad-like impressions with a characteristic nodular shape, and small, pointed claw impressions are also present. Some specimens preserved irregularly-shaped sole impressions and/or tail drag marks, but these are not always preserved. One purported ''Gwyneddichnium'' trackway (CU-MWC 159.10) has been interpreted as swimming traces due to the absence of manus prints. Skin webbing appears to be present between toes I-III,<ref name=":1" /> though the webbing has also been interpreted as sediment deformation. ''Gwyneddichnium'' trackways in general are widely spaced, with pes prints pointing forwards and manus prints rotated outwards. The positions of the pes and manus prints relative to each other are variable, corresponding to different speeds and gaits.<ref name=":0" />
The overall shape of the footprints are similar to ''[[Rhynchosauroides]]'', which sometimes occurs alongside ''Gwyneddichnium''. However, ''Rhynchosauroides'' has an exaxonic print (with the fourth digit longer than the third), more curvature in the digits, and a fifth digit more widely separated from the fourth.<ref name=":0" />
== Distribution ==
''Gwyneddichnium'' fossils were first collected by Wilhelm Bock at Gwynedd, a railroad outcrop in [[Montgomery County, Pennsylvania]].<ref name="Bock52b">Liquid error: wrong number of arguments (given 1, expected 2)</ref> Gwynedd preserves sediments from the late Triassic [[Lockatong Formation]], part of the [[Newark Supergroup]] exposed in the [[Newark Basin]]. Bock's specimens, now stored at the [[Academy of Natural Sciences of Drexel University|Academy of Natural Sciences]] in [[Philadelphia]], include ANSP 15212 (the [[holotype]] of ''G. major''), ANSP 15213 (the [[paratype]] of ''G. major''), ANSP 15214 (the holotype of "''G. elongatum''"), ANSP 15215 (the paratype of "''G. elongatum''", now considered lost), ANSP 15216 (the holotype of "''G. minore''"), and ANSP 15217 (the paratype of "''G. minore''"). Various outcrops throughout Pennsylvania and [[New Jersey]] which express the Lockatong Formation continue to produce ''Gwyneddichnium'' fossils.<ref name="olsen flynn 1989"></ref><ref name=":0" /><ref name="Fillmore17">Liquid error: wrong number of arguments (given 1, expected 2)</ref> ''Gwyneddichnium'' is also known to occur within the overlying [[Passaic Formation]].<ref name="Baird86">Liquid error: wrong number of arguments (given 1, expected 2)</ref><ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref> Outside of the Newark Basin, ''Gwyneddichnium'' tracks have also been found in the [[Bull Run Formation]] (also known as the "[[Balls Bluff Siltstone]]") at [[Manassas National Battlefield Park]] in the [[Culpeper Basin]] of [[Virginia]].<ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref><ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref><ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref> They are also known from the [[New Oxford Formation|New Oxford]] or [[Gettysburg Formation]] of the [[Gettysburg Basin]] in [[Maryland]].<ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref>
Reports of ''Gwyneddichnium'' trackways in the western United States began to surface in the 1990s. These included both typical trackways from walking animals, and unusual swimming trackways which were provisionally referred to the ichnogenus. The earliest western ''Gwyneddichnium'' tracks to be discovered hail from several sites in Northeast [[Utah]] and Northwest [[Colorado]]. Many of these sites lie within the boundaries of [[Dinosaur National Monument]]. The sites in the area preserve the Rock Point Member of the upper Triassic [[Chinle Formation]], though they were originally reported as representing the age-equivalent [[Popo Agie Formation]].<ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref><ref name=":1">Liquid error: wrong number of arguments (given 1, expected 2)</ref><ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref> Possible ''Gwyneddichnium'' prints are also known from the Chinle Formation's Owl Creek Member. These rare fossils were found at Brinkerhof Spring, a site in south-central Utah near [[Capitol Reef National Park]].<ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref><ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref><ref name=":0" /> Brinkerhof Spring is located in Circle Cliffs, a natural amphitheater formerly protected within the boundaries of [[Grand Staircase–Escalante National Monument|Grand Staircase-Escalante National Monument]], but excluded when the monument was shrunk in 2017.<ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref>
In 2007, a new ichnogenus and ichnospecies, ''Apachepus cottonorum'', was named from the [[Redonda Formation]] of [[New Mexico]].<ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref> However, ''Apachepus cottonorum'' was subsequently considered a species of ''Gwyneddichnium'' (''Gwyneddichnium cottonorum'') after pentadactyl manus prints were discovered. ''G. cottonorum'' was distinguished from ''G. major'' by its larger size and more evenly spaced digits.<ref name=":2">Liquid error: wrong number of arguments (given 1, expected 2)</ref> Further sampling has suggested that these differences are not as clear-cut as suggested, so ''Gwyneddichnium cottonorum'' is currently considered a junior synonym of ''Gwyneddichnium majore''.<ref name=":0" />
The only locale outside the United States known to produce ''Gwyneddichnium'' prints is the Berndorf site in [[Germany]]. Berndorf is located near [[Kemnath]] in [[Bavaria]] and preserves a [[Clastic rock|clastic]] portion of the lower [[Muschelkalk]] known as the [[Eschenbach Formation]]. The rocks of the Eschenbach Formation are from the lower [[middle Triassic]] ([[Anisian]] stage), making their prints the oldest known ''Gwyneddichnium'' fossils.<ref name=":0" /><ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref>
== Trackmaker ==
''Gwyneddichnium'' was originally tentatively suggested as being created by small [[Pseudosuchia|pseudosuchians]].<ref name="Bock52b" /> The discovery of the small [[Tanystropheidae|tanystropheid]] [[Archosauromorpha|archosauromorph]] ''[[Tanytrachelos]]'' provided a more specific and well-supported trackmaker for ''Gwyneddichnium''. ''Tanytrachelos'' and other tanystropheids have an unusual [[Metatarsal bones|metatarsal]]-like [[Phalanx bone|phalange]] in the fifth toe which points forwards (like that of ''Gwyneddichnium'') rather than curves back like other reptiles. ''Tanytrachelos'' in particular is the most likely trackmaker due to its wide gait, small size, co-occurrence with the tracks, and feet with a fourth digit shorter than the third.<ref name="Baird86" /><ref name="olsen flynn 1989" /><ref name=":0" /> Though ''Tanytrachelos'' is by far the most commonly considered trackmaker for ''Gwyneddichnium'', small [[Drepanosaur|drepanosaurs]] such as ''[[Hypuronector]]'' or ''[[Dolabrosaurus]]'' have also been proposed as trackmakers.<ref name=":1" /> This is considered unlikely due to their unusual morphology.<ref name=":0" /> At least in the Chinle and Redonda formations, small [[Lepidosauromorpha|lepidosauromorphs]] have been suggested as trackmakers for ''Gwyneddichnium''.<ref name=":2" />
== References ==
[[Category:Reptile trace fossils]]
== Description ==
''Gwyneddichnium'' corresponds to footprints from an a quadrupedal animal with a small pentadactyl (five-fingered) manus (hand) and a notably larger five-toed pes (foot). The manus and pes are mesaxonic, meaning that the third digit is the longest digit, followed by the subequal second and fourth digits. The innermost digit (digit I) and the outermost digit (digit V) were short and located close to the rest of the foot. Sometimes the small fifth digit is poorly preserved, making the hand or foot appear to be tetradactyl (having only four digits). Overall, the digits are long and narrow, with minimal curvature. There is some variation with how the digits are positioned, with some specimens having digits which evenly radiate away from the sole,<ref name=":0" /> and others having digits which separate into two clumps comprised of digits I-III and IV-V.<ref name=":1" />
The skin made pad-like impressions with a characteristic nodular shape, and small, pointed claw impressions are also present. Some specimens preserved irregularly-shaped sole impressions and/or tail drag marks, but these are not always preserved. One purported ''Gwyneddichnium'' trackway (CU-MWC 159.10) has been interpreted as swimming traces due to the absence of manus prints. Skin webbing appears to be present between toes I-III,<ref name=":1" /> though the webbing has also been interpreted as sediment deformation. ''Gwyneddichnium'' trackways in general are widely spaced, with pes prints pointing forwards and manus prints rotated outwards. The positions of the pes and manus prints relative to each other are variable, corresponding to different speeds and gaits.<ref name=":0" />
The overall shape of the footprints are similar to ''[[Rhynchosauroides]]'', which sometimes occurs alongside ''Gwyneddichnium''. However, ''Rhynchosauroides'' has an exaxonic print (with the fourth digit longer than the third), more curvature in the digits, and a fifth digit more widely separated from the fourth.<ref name=":0" />
== Distribution ==
''Gwyneddichnium'' fossils were first collected by Wilhelm Bock at Gwynedd, a railroad outcrop in [[Montgomery County, Pennsylvania]].<ref name="Bock52b">Liquid error: wrong number of arguments (given 1, expected 2)</ref> Gwynedd preserves sediments from the late Triassic [[Lockatong Formation]], part of the [[Newark Supergroup]] exposed in the [[Newark Basin]]. Bock's specimens, now stored at the [[Academy of Natural Sciences of Drexel University|Academy of Natural Sciences]] in [[Philadelphia]], include ANSP 15212 (the [[holotype]] of ''G. major''), ANSP 15213 (the [[paratype]] of ''G. major''), ANSP 15214 (the holotype of "''G. elongatum''"), ANSP 15215 (the paratype of "''G. elongatum''", now considered lost), ANSP 15216 (the holotype of "''G. minore''"), and ANSP 15217 (the paratype of "''G. minore''"). Various outcrops throughout Pennsylvania and [[New Jersey]] which express the Lockatong Formation continue to produce ''Gwyneddichnium'' fossils.<ref name="olsen flynn 1989"></ref><ref name=":0" /><ref name="Fillmore17">Liquid error: wrong number of arguments (given 1, expected 2)</ref> ''Gwyneddichnium'' is also known to occur within the overlying [[Passaic Formation]].<ref name="Baird86">Liquid error: wrong number of arguments (given 1, expected 2)</ref><ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref> Outside of the Newark Basin, ''Gwyneddichnium'' tracks have also been found in the [[Bull Run Formation]] (also known as the "[[Balls Bluff Siltstone]]") at [[Manassas National Battlefield Park]] in the [[Culpeper Basin]] of [[Virginia]].<ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref><ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref><ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref> They are also known from the [[New Oxford Formation|New Oxford]] or [[Gettysburg Formation]] of the [[Gettysburg Basin]] in [[Maryland]].<ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref>
Reports of ''Gwyneddichnium'' trackways in the western United States began to surface in the 1990s. These included both typical trackways from walking animals, and unusual swimming trackways which were provisionally referred to the ichnogenus. The earliest western ''Gwyneddichnium'' tracks to be discovered hail from several sites in Northeast [[Utah]] and Northwest [[Colorado]]. Many of these sites lie within the boundaries of [[Dinosaur National Monument]]. The sites in the area preserve the Rock Point Member of the upper Triassic [[Chinle Formation]], though they were originally reported as representing the age-equivalent [[Popo Agie Formation]].<ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref><ref name=":1">Liquid error: wrong number of arguments (given 1, expected 2)</ref><ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref> Possible ''Gwyneddichnium'' prints are also known from the Chinle Formation's Owl Creek Member. These rare fossils were found at Brinkerhof Spring, a site in south-central Utah near [[Capitol Reef National Park]].<ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref><ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref><ref name=":0" /> Brinkerhof Spring is located in Circle Cliffs, a natural amphitheater formerly protected within the boundaries of [[Grand Staircase–Escalante National Monument|Grand Staircase-Escalante National Monument]], but excluded when the monument was shrunk in 2017.<ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref>
In 2007, a new ichnogenus and ichnospecies, ''Apachepus cottonorum'', was named from the [[Redonda Formation]] of [[New Mexico]].<ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref> However, ''Apachepus cottonorum'' was subsequently considered a species of ''Gwyneddichnium'' (''Gwyneddichnium cottonorum'') after pentadactyl manus prints were discovered. ''G. cottonorum'' was distinguished from ''G. major'' by its larger size and more evenly spaced digits.<ref name=":2">Liquid error: wrong number of arguments (given 1, expected 2)</ref> Further sampling has suggested that these differences are not as clear-cut as suggested, so ''Gwyneddichnium cottonorum'' is currently considered a junior synonym of ''Gwyneddichnium majore''.<ref name=":0" />
The only locale outside the United States known to produce ''Gwyneddichnium'' prints is the Berndorf site in [[Germany]]. Berndorf is located near [[Kemnath]] in [[Bavaria]] and preserves a [[Clastic rock|clastic]] portion of the lower [[Muschelkalk]] known as the [[Eschenbach Formation]]. The rocks of the Eschenbach Formation are from the lower [[middle Triassic]] ([[Anisian]] stage), making their prints the oldest known ''Gwyneddichnium'' fossils.<ref name=":0" /><ref>Liquid error: wrong number of arguments (given 1, expected 2)</ref>
== Trackmaker ==
''Gwyneddichnium'' was originally tentatively suggested as being created by small [[Pseudosuchia|pseudosuchians]].<ref name="Bock52b" /> The discovery of the small [[Tanystropheidae|tanystropheid]] [[Archosauromorpha|archosauromorph]] ''[[Tanytrachelos]]'' provided a more specific and well-supported trackmaker for ''Gwyneddichnium''. ''Tanytrachelos'' and other tanystropheids have an unusual [[Metatarsal bones|metatarsal]]-like [[Phalanx bone|phalange]] in the fifth toe which points forwards (like that of ''Gwyneddichnium'') rather than curves back like other reptiles. ''Tanytrachelos'' in particular is the most likely trackmaker due to its wide gait, small size, co-occurrence with the tracks, and feet with a fourth digit shorter than the third.<ref name="Baird86" /><ref name="olsen flynn 1989" /><ref name=":0" /> Though ''Tanytrachelos'' is by far the most commonly considered trackmaker for ''Gwyneddichnium'', small [[Drepanosaur|drepanosaurs]] such as ''[[Hypuronector]]'' or ''[[Dolabrosaurus]]'' have also been proposed as trackmakers.<ref name=":1" /> This is considered unlikely due to their unusual morphology.<ref name=":0" /> At least in the Chinle and Redonda formations, small [[Lepidosauromorpha|lepidosauromorphs]] have been suggested as trackmakers for ''Gwyneddichnium''.<ref name=":2" />
== References ==
[[Category:Reptile trace fossils]]
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