Palmichnium
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''Palmichnium'' ("palm trace") is an
ichnofossil 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 ...
genus, interpreted as a
eurypterid Eurypterids, often informally called sea scorpions, are a group of extinct marine arthropods that form the Order (biology), order Eurypterida. The earliest known eurypterids date to the Darriwilian stage of the Ordovician period, 467.3 Myr, mil ...
trackway. It has been found by many places around the world, such as Australia, Canada, United States and
Wales Wales ( ) is a Countries of the United Kingdom, country that is part of the United Kingdom. It is bordered by the Irish Sea to the north and west, England to the England–Wales border, east, the Bristol Channel to the south, and the Celtic ...
. Its trackways consist of three or four subcircular tracks that are symmetrical around a midline impression that is arranged en echelon with a high angle to the midline.


Ichnospecies

The
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 ...
contains eight ichnospecies. *''Palmichnium antarcticum'' Gevers ''et al.'', 1971 *''Palmichnium capensis'' Anderson, 1975 *''?Palmichnium culmicum'' Pfeiffer, 1968 *''Palmichnium kosinskiorum'' Briggs and Rolfe, 1983 *''Palmichnium macdonaldi'' Braddy, 1995 *''Palmichnium palmatum'' Richter, 1954 (
type Type may refer to: Science and technology Computing * Typing, producing text via a keyboard, typewriter, etc. * Data type, collection of values used for computations. * File type * TYPE (DOS command), a command to display contents of a file. * ...
) *''Palmichnium pottsae'' Braddy and Anderson, 1996 *''Palmichnium stoermeri'' Briggs and Rolfe, 1983


History of research

In 1975, Ann M. Anderson described a new ichnospecies of '' Petalichnus'' from the
Table Mountain Sandstone Table Mountain Sandstone (TMS), formally known by its geological name the Peninsula Formation Sandstone, is a group of rock formations within the Cape Supergroup sequence. While the term "Table Mountain Sandstone" remains widely used, it is n ...
of
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
, ''P. capensis'',
Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, and the second of twelve periods of the Phanerozoic Eon (geology), Eon. The Ordovician spans 41.6 million years f ...
in time. Its ichnospecific name ''capensis'' refers to the type locality in the
Cape Province The Province of the Cape of Good Hope (), commonly referred to as the Cape Province () and colloquially as The Cape (), was a province in the Union of South Africa and subsequently the Republic of South Africa. It encompassed the old Cape Co ...
of South Africa. Anderson noted that ''P. capensis'' was larger than the rest of the ''Petalichnus'' ichnospecies, as well as the presence of an unusual median drag line for the ichnogenus (although this was not present in all the specimens). However, in 1999, Simon J. Braddy and John E. Almond formally reclassified some specimens of ''Petalichnus capensis'' to ''Palmichnium'' due to the possession of four tracks and a medium line, the rest of the specimens were referred to ''Petalichnus brandenburgensis''. The trackway of ''P. capensis'' is defined as medium-sized (largest track 13.6 cm, 5.4 in wide) and consists of several symmetrical series of four tracks, each formed by a simple oval or tear-shaped mark with small impressions on the sides, sometimes bilobed (divided into two lobes) and intermittent formed by closely spaced series. These tracks were located at a low angle with respect to the sometimes present median line, which is believed to be due to the
telson The telson () is the hindmost division of the body of an arthropod. Depending on the definition, the telson is either considered to be the final segment (biology), segment of the arthropod body, or an additional division that is not a true segm ...
("tail") touching the substrate. The bilobed marks is thought to be "eight"-shaped due to the long spines of the sixth appendage of an onychopterellid eurypterid, presumably '' Onychopterella augusti'', using a swimming stroke-like motion. In 1983, Derek E. Gilmor Briggs and William D. Ian Rolfe described two new ichnospecies from
Pennsylvania Pennsylvania, officially the Commonwealth of Pennsylvania, is a U.S. state, state spanning the Mid-Atlantic (United States), Mid-Atlantic, Northeastern United States, Northeastern, Appalachian, and Great Lakes region, Great Lakes regions o ...
, ''P. kosinskiorum'' and ''P. stoermeri''. The trace fossil of both ichnospecies were collected at a party from the
Carnegie Museums of Pittsburgh Carnegie Museums of Pittsburgh is a nonprofit organization that operates four museums in Pittsburgh, Pennsylvania, United States. The organization is headquartered in the Carnegie Institute and Library complex in the Oakland neighborhood of Pitts ...
in 1948. Its large size attracted the attention of the press, which attributed it to three-clawed
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 ...
hopping bipedally. The median line, the nature of the footprints and the lack of a suitable hopping vertebrate at the time rule out any hypothesis of an amphibian or vertebrate as the author of the footprints. In the case of invertebrates, it was concluded that only a large eurypterid that walked on six legs (hexapodous) could have produced the tracks. Braddy erected in 1995 ''P. macdonaldi'' for Lower
Permian The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years, from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.902 Mya. It is the s ...
trackways found in the
Hueco Formation The Hueco Formation is a geologic formation in west Texas and southern New Mexico.Richardson 1904Richardson 1908Richardson 1914 It preserves fossils dating back to the early Permian period.Nelson 1940Bachman and Hayes 1958 Description The format ...
of
New Mexico New Mexico is a state in the Southwestern United States, Southwestern region of the United States. It is one of the Mountain States of the southern Rocky Mountains, sharing the Four Corners region with Utah, Colorado, and Arizona. It also ...
, becoming the youngest species of the ichnogenus. The specimens, including holotype P23404, are housed in the
New Mexico Museum of Natural History and Science The New Mexico Museum of Natural History and Science is a natural history and science museum in Albuquerque, New Mexico near Old Town Albuquerque. The Museum was founded in 1986. It operates as a public revenue facility of the New Mexico Departme ...
. The ichnospecies is named in honor of Jerry MacDonald, the discoverer and local collector of the ichnofauna of the
Robledo Mountains The Robledo Mountains are a mountain range in Doña Ana County, New Mexico, just northwest of Las Cruces. They are an uplifted block of Paleozoic rocks on the west side of the Rio Grande Rift. The range was named for Pedro Robledo, who died o ...
. The prints had a width of 25 mm (1 in) and consisted of symmetrical series of three to four pairs of impressions, large paddle-shaped outer tracks of 12 mm (0.5 in) in length and two or three internal linear impressions. In front of some outer tracks there is a slight curvilinear impression slightly convex outwards, possibly representing the recovery of a stroke of an appendage. There are also present inner tracks, small and linear scritches. An intermittent median groove preserved in the middle of the trackway is situated between the outer tracks. The producer of the footprints was a small arthropod with long and paddle-shaped posterior appendages and with three pairs of appendages in front, using an in-phase gait. Due to these characteristics it has been suggested that the producer was '' Adelophthalmus luceroensis'', which has been found in nearby deposits (
Madera Formation The Madera Group is a group of geologic formations in northern New Mexico. Its fossil assemblage dates the formation to the middle to late Pennsylvanian period. Description The group consists primarily of marine limestones, and it is exposed in ...
, central New Mexico). In 1843, the English geologist and paleontologist
William Buckland William Buckland Doctor of Divinity, DD, Royal Society, FRS (12 March 1784 – 14 August 1856) was an English theologian, geologist and paleontology, palaeontologist. His work in the early 1820s proved that Kirkdale Cave in North Yorkshire h ...
described the now called BGS GSM 26037 (housed at the
British Geological Survey The British Geological Survey (BGS) is a partly publicly funded body which aims to advance Earth science, geoscientific knowledge of the United Kingdom landmass and its continental shelf by means of systematic surveying, monitoring and research. ...
in
Nottingham Nottingham ( , East Midlands English, locally ) is a City status in the United Kingdom, city and Unitary authorities of England, unitary authority area in Nottinghamshire, East Midlands, England. It is located south-east of Sheffield and nor ...
,
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 ...
) specimen as the result of the repeated impressions of three
bony Bony may refer to: * Adjective relating to bone People * Bony Dashaco, Cameroonian businessman * Bony King, Belgian singer-songwriter * Bony Pierre (born 1991), Haitian footballer * Bony Ramirez (born 1996), Dominican-born American painter and ...
processes of the
pectoral fin Fins are moving appendages protruding from the body of fish that interact with water to generate thrust and help the fish aquatic locomotion, swim. Apart from the tail or caudal fin, fish fins have no direct connection with the vertebral column ...
of an ambulatory (related or adapted to walk)
fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
, ''Icthyopatolites''. The specimen was collected in
Mostyn Mostyn is a village and Community (Wales), community in Flintshire, Wales, and Wards and electoral divisions of the United Kingdom, electoral ward lying on the estuary of the River Dee, Wales, River Dee, located near the town of Holywell, Flin ...
,
Wales Wales ( ) is a Countries of the United Kingdom, country that is part of the United Kingdom. It is bordered by the Irish Sea to the north and west, England to the England–Wales border, east, the Bristol Channel to the south, and the Celtic ...
, and is Westphalian (Late Carboniferous) in age. However, in 1996, Braddy and Lyall I. Anderson recognized the track as clearly that of a moderately large (around 20 cm, 7.9 in long) hexapodous arthropod similar to the eurypterid traits that were described before in other papers, and named it ''Palmichnium pottsae''. Although the leg morphology of the Carboniferous eurypterids is poorly known or completely unknown, the trackway fits with the maximum leg span of ''Adelophthalmus'', approximately 10 cm (3.9 in). ''P. pottsae'' produced large trackways with asymmetric rows of three paired impressions along several irregular prints. It lacked a median line, but it has been suggested that this is due to the buoyancy of the water surrounding the abdomen of the animal or the active held up of the telson on the substrate without touching it, allowing better locomotion. The total length of the trackway was 37 cm (14.6 in). The outer tracks of the right and left series of the trackway were curvilinear or linear grooves, as well as the intermediate tracks, which were smaller than the outer ones. The internal tracks of both series were the smallest and with a bifid (cleft-like) form. There were also several irregular tracks that did not belong to any of the series. Although the assignment of ''P. pottsae'' to ''Palmichnium'' should mean the synonymy of ''Icthyopatolites'' with the first ichnogenus, Braddy and Anderson avoided formally synonymizing them because Buckland informally described ''Icthyopatolites'' and without an established diagnosis. Early Devonian trackways have also been found in Alken an der Mosel, Germany.


References

Eurypterida Fossil trackways {{trace-fossil-stub