Protichnites
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''Protichnites'' is an
ichnogenus An ichnotaxon (plural ichnotaxa) is "a taxon based on the fossilized work of an organism", i.e. the non-human equivalent of an artifact. ''Ichnotaxon'' comes from the Ancient Greek (''íchnos'') meaning "track" and English , itself derived from ...
of
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 the imprints made by the walking activity of certain
arthropods Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
. It consists of two rows of tracks and a medial furrow between the two rows. This furrow, which may be broken, set at an angle, and of varying width and depth, is thought to be the result of the tail region contacting the substrate.


The first footprints on land

Sir
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 ...
, the noted British paleontologist and anatomist who coined the term “ Dinosauria”, based ''Protichnites'' on trackways that were shipped to him from the
Cambrian The Cambrian ( ) is the first geological period of the Paleozoic Era, and the Phanerozoic Eon. The Cambrian lasted 51.95 million years from the end of the preceding Ediacaran period 538.8 Ma (million years ago) to the beginning of the Ordov ...
Potsdam Sandstone of
Quebec Quebec is Canada's List of Canadian provinces and territories by area, largest province by area. Located in Central Canada, the province shares borders with the provinces of Ontario to the west, Newfoundland and Labrador to the northeast, ...
.Owen, 1851 He was correct in recognizing that these were the oldest known footprints on land, having been produced in intertidal and supratidal environments during what would now be called Cambrian times.Collette and Hagadorn, 2010


The ''Protichnites'' makers

Owen first thought that these trackways were made by tortoises, but new material convinced him that “articulates” (a group that included the arthropods) were responsible.Owen, 1852 He further suggested a kinship with '' Limulus'', the modern horseshoe crab. Additional material was collected in Quebec,
Ontario Ontario is the southernmost Provinces and territories of Canada, province of Canada. Located in Central Canada, Ontario is the Population of Canada by province and territory, country's most populous province. As of the 2021 Canadian census, it ...
,
New York New York most commonly refers to: * New York (state), a state in the northeastern United States * New York City, the most populous city in the United States, located in the state of New York New York may also refer to: Places United Kingdom * ...
,
Wisconsin Wisconsin ( ) is a U.S. state, state in the Great Lakes region, Great Lakes region of the Upper Midwest of the United States. It borders Minnesota to the west, Iowa to the southwest, Illinois to the south, Lake Michigan to the east, Michig ...
, and
Missouri Missouri (''see #Etymology and pronunciation, pronunciation'') is a U.S. state, state in the Midwestern United States, Midwestern region of the United States. Ranking List of U.S. states and territories by area, 21st in land area, it border ...
for the next 150 years without a single fossil of the maker of these traces. Finally, body fossils of potential makers were found in two of the same quarries that preserved ''Protichnites'' from the Elk Mound Group of Blackberry Hill, Wisconsin, and the Potsdam Group of Melocheville, Quebec. The animals, '' Mosineia macnaughtoni'' and ''Mictomerus melochevillensis'', were euthycarcinoids, extinct arthropods that may have given rise to the
mandibulate The clade Mandibulata constitutes one of the major subdivisions of the phylum Arthropoda, alongside Chelicerata. Mandibulates include the crustaceans, myriapods (centipedes and millipedes, among others), and all true insects. The name "Mandibulat ...
s. Fossils that clearly tie euthycarcinoids to ''Protichnites'' were then found at Blackberry Hill.Collette et al., 2012 It is possible that other extinct arthropods, such as members of the
Aglaspidida Aglaspidida is an extinct order of marine arthropods known from fossils spanning the Middle Cambrian to the Upper Ordovician. Initially considered chelicerates, modern anatomical comparisons demonstrate that the aglaspidids cannot be accommodated ...
, may also have produced some of these trackways.
Trilobite Trilobites (; meaning "three-lobed entities") are extinction, extinct marine arthropods that form the class (biology), class Trilobita. One of the earliest groups of arthropods to appear in the fossil record, trilobites were among the most succ ...
s have been suggested as well; however, no aglaspidids or trilobites have been found thus far in the strata that contain this ichnogenus. Similar trackways are present in post-Cambrian strata; however, those are seldom referred to as ''Protichnites''.


Behavior

It has been suggested that one form of ''Protichnites'', ''P. eremita'' Form 1, may have been produced by an ambulating pair of individuals, as in
amplexus Amplexus (Latin "embrace") is a type of Mating, mating behavior exhibited by some External fertilization, externally fertilizing species (chiefly amphibians, Amphipoda, amphipods, and horseshoe crabs) in which a male grasps a female with his fro ...
. In this form, the medial furrow is discontinuous and each section is set at an angle, such that the series of medial imprints have a shingled appearance, as might result if the female were displacing her tail to minimize its interference with external fertilization. A second set of medial imprints observed in Form 1 is postulated to have been produced by the tail region of the male of the pair. ''Protichnites eremita'' was originally interpreted as early evidence of
hermit crab Hermit crabs are anomuran Decapoda, decapod crustaceans of the superfamily (taxonomy), superfamily Paguroidea that have adapted to occupy empty scavenged mollusc shells to protect their fragile exoskeletons. There are over 800 species of hermit c ...
-like behavior. It has also been suggested that some ''P. eremita'' may have been produced by distressed individuals attempting to return to a body of water after being stranded during high tide, which was more extensive than today due to the moon being closer to Earth in Cambrian times.Braddy and Gass, 2022


Relationship with ''Diplichnites''

Differing only in the presence or absence of a medial furrow, ''Protichnites'' and '' Diplichnites'' trackways could both be produced by the same individual animal. In cases where the feet penetrate the sediment more deeply than does the tail, ''Diplichnites'' could be created on the underlying layer while ''Protichnites'' is being produced on the surface.Hoxie, 2005


References


Further reading

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External links


Discusses a major source of ''Protichnites'' trackways.
{{Taxonbar, from=Q7251819 Cambrian arthropods Arthropod trace fossils Paleozoic life of Ontario