Protichnites

Sir Richard Owen, the noted British paleontologist and anatomist who coined the term “Dinosauria”, based Protichnites on trackways that were shipped to him from the Cambrian Potsdam Sandstone of Quebec.[2] 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.[3]

Owen first thought that these trackways were made by tortoises,[2] but new material convinced him that “articulates” (a group that included the arthropods) were responsible.[1] He further suggested a kinship with Limulus, the modern horseshoe crab. Additional material was collected in Quebec, Ontario, New York, Wisconsin, and Missouri 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.[3] The animals, Mosineia macnaughtoni and Mictomerus melochevillensis, were euthycarcinoids, extinct arthropods that may have given rise to the mandibulates.[4] Fossils that clearly tie euthycarcinoids to Protichnites were then found at Blackberry Hill.[5]

It is possible that other extinct arthropods, such as members of the Aglaspidida, may also have produced some of these trackways. Trilobites 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.

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. 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.[5] Protichnites eremita was originally interpreted as early evidence of hermit crab-like behavior.[6] 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.[7]

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.[8][9]

• Braddy, S. J. & Gass, K.C. (2022). "Fossils of Blackberry Hill, Wisconsin, USA: the first animals on land, 500 million years ago". Geology Today. 38 (1): 25–31. doi:10.1111/gto.12379. S2CID 246604495.{{cite journal}}: CS1 maint: multiple names: authors list (link)
• Collette, J. H., K. C. Gass & J. W. Hagadorn (2012). "Protichnites eremita unshelled? Experimental model-based neoichnology and new evidence for a euthycarcinoid affinity for this ichnospecies". Journal of Paleontology. 86 (3): 442–454. doi:10.1666/11-056.1. S2CID 129234373.{{cite journal}}: CS1 maint: multiple names: authors list (link)
• Collette, J. H.; J. W. Hagadorn (2010). "Three-dimensionally preserved arthropods from Cambrian Lagerstatten of Quebec and Wisconsin". Journal of Paleontology. 84 (4): 646–667. doi:10.1666/09-075.1. S2CID 130064618.
• Goldring, R.; A. Seilacher (1971). "Limulid undertracks and their sedimentological implications". Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 137: 422–442.
• Hagadorn, J. W.; A. Seilacher (2009). "Hermit arthropods 500 million years ago?". Geology. 37 (4): 295–298. Bibcode:2009Geo....37..295H. doi:10.1130/G25181A.1.
• Hesselbo, S. P. (1989). "Trace fossils of Cambrian aglaspidid arthropods". Lethaia. 21 (2): 129–146. doi:10.1111/j.1502-3931.1988.tb02064.x.
• Hoxie, C. T. (2005). "Late Cambrian arthropod trackways in subaerially exposed environments: Incentives to simplify a problematic ichnogenus". Unpublished B.A. Thesis: 1–89.
• C. T. Hoxie; J. W. Hagadorn (2005). "Late Cambrian arthropod trackways in subaerially exposed environments". Geological Society of America Abstracts with Programs, Northeastern Section. 37: 12.
• Ortega-Hernandez, J., Tremewan, J., & Braddy, S. J. (2010). "Euthycarcinoids". Geology Today. 26 (5): 195–198. doi:10.1111/j.1365-2451.2010.00770.x. S2CID 247664753.{{cite journal}}: CS1 maint: multiple names: authors list (link)
• Owen, R. (1851). "Description of the impressions on the Potsdam sandstone, discovered by Mr. Logan in Lower Canada". Geological Society of London Quarterly Journal. 7 (1–2): 250–252. doi:10.1144/GSL.JGS.1851.007.01-02.42. S2CID 140674536.
• Owen, R. (1852). "Description of the impressions and footprints of the Protichnites from the Potsdam sandstone of Canada". Geological Society of London Quarterly Journal. 8 (1–2): 214–225. doi:10.1144/GSL.JGS.1852.008.01-02.26. S2CID 130712914.
• N. E. Vaccari, G. D. Edgecombe & C. Escudero (2004). "Cambrian origins and affinities of an enigmatic fossil group of arthropods". Nature. 430 (6999): 554–557. Bibcode:2004Natur.430..554V. doi:10.1038/nature02705. PMID 15282604. S2CID 4419235.

• Discusses a major source of Protichnites trackways.