Peytoia Nathorsti
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''Peytoia'' is a genus of
hurdiid Hurdiidae is an extinct cosmopolitan family of radiodonts, a group of stem-group arthropods, which lived during the Paleozoic Era. It is the most long-lived radiodont clade, lasting from the Cambrian period to the Devonian period. Descripti ...
radiodont Radiodonta is an extinct Order (biology), order of stem-group arthropods that was successful worldwide during the Cambrian period. They may be referred to as radiodonts, radiodontans, radiodontids, anomalocarids, or anomalocaridids, although the ...
that lived in the
Cambrian The Cambrian Period ( ; sometimes symbolized C with bar, Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million ...
period, containing two species, ''Peytoia nathorsti'' from the
Miaolingian The Miaolingian is the third Series of the Cambrian Period, and was formally named in 2018. It lasted from about to million years ago and is divided in ascending order into 3 stages: the Wuliuan, Drumian, and Guzhangian. The Miaolingian is prec ...
of Canada and ''
Peytoia infercambriensis ''Peytoia infercambriensis'' is a species of hurdiid radiodont in the genus ''Peytoia''. ''P. infercambriensis'' is the geologically oldest known radiodont; its remains date to the third age of the Cambrian. The type and only known specimen ...
'' from Poland, dating to
Cambrian Stage 3 Cambrian Stage 3 is the still unnamed third stage of the Cambrian. It succeeds Cambrian Stage 2 and precedes Cambrian Stage 4, although neither its base nor top have been formally defined. The plan is for its lower boundary to correspond approxi ...
. Its two frontal appendages had long
bristle A bristle is a stiff hair or feather (natural or artificial), either on an animal, such as a pig, a plant, or on a tool such as a brush or broom. Synthetic types Synthetic materials such as nylon are also used to make bristles in items such as br ...
-like spines, it had no fan tail, and its short stalked eyes were behind its large head. 108 specimens of ''Peytoia'' are known from the Greater
Phyllopod bed The Phyllopod bed, designated by USNM locality number 35k, is the most famous fossil-bearing member of the Burgess Shale fossil ''Lagerstätte''. It was quarried by Charles Walcott from 1911–1917 (and later named Walcott Quarry), and was t ...
, where they comprise 0.21% of the community. ''Peytoia nathorsti'' and its junior synonym ''Laggania cambria'' played a major role in the discovery of the radiodont body plan. Initially interpreted as a
jellyfish Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the subphylum Medusozoa, a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animals with umbrella- ...
and a
sea cucumber Sea cucumbers are echinoderms from the class Holothuroidea (). They are marine animals with a leathery skin and an elongated body containing a single, branched gonad. Sea cucumbers are found on the sea floor worldwide. The number of holothuria ...
respectively, they were eventually shown to be the mouthparts and body of a single animal, which bore ''
Anomalocaris ''Anomalocaris'' ("unlike other shrimp", or "abnormal shrimp") is an extinct genus of Radiodonta, radiodont, an Order (biology), order of early-diverging stem-group arthropods. The first fossils of ''Anomalocaris'' were discovered in the ''Ogygop ...
''-like appendages. ''Peytoia infercambriensis'' is the geologically oldest known radiodont species.


Classification

''Peytoia'' belongs to the clade
Hurdiidae Hurdiidae is an extinct cosmopolitan distribution, cosmopolitan Family (biology), family of radiodonts, a group of Crown group#Stem groups, stem-group arthropods, which lived during the Paleozoic Era. It is the most long-lived radiodont clade, la ...
, and is closely related to the contemporary genus ''
Hurdia ''Hurdia'' is an extinct genus of hurdiid radiodont that lived 505 million years ago during the Cambrian Period. As a radiodont like ''Peytoia'' and ''Anomalocaris'', it is part of the ancestral lineage that led to euarthropods. Description ' ...
''. ''Peytoia'' contains two named species: ''
Peytoia nathorsti ''Peytoia'' is a genus of hurdiid radiodont that lived in the Cambrian period, containing two species, ''Peytoia nathorsti'' from the Miaolingian of Canada and ''Peytoia infercambriensis'' from Poland, dating to Cambrian Stage 3. Its two front ...
'', the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
, from the
Burgess Shale The Burgess Shale is a fossil-bearing deposit exposed in the Canadian Rockies of British Columbia, Canada. It is famous for the exceptional preservation of the soft parts of its fossils. At old (middle Cambrian), it is one of the earliest foss ...
of Canada and the
Wheeler Wheeler may refer to: Places United States * Wheeler, Alabama, an unincorporated community * Wheeler, Arkansas, an unincorporated community * Wheeler, California, an unincorporated community * Wheeler, Illinois, a village * Wheeler, Indiana, a ...
and
Marjum Formation The Marjum Formation is a Cambrian geological formation that overlies the Wheeler Shale in the House Range, Utah. It is known for its occasional preservation of soft-bodied tissue, and is slightly younger than the Burgess Shale The Burgess S ...
s of the United States, and ''
Peytoia infercambriensis ''Peytoia infercambriensis'' is a species of hurdiid radiodont in the genus ''Peytoia''. ''P. infercambriensis'' is the geologically oldest known radiodont; its remains date to the third age of the Cambrian. The type and only known specimen ...
'' from the Zawiszyn Formation of Poland. Another species of ''Peytoia'' may be present in the Burgess Shale, represented by a single frontal appendage from the Tulip Beds locality. A specimen regarded as ''Peytoia'' cf. ''nathorsti'' is known from the
Balang Formation The Balang formation, is a Cambrian Period geological formation, which outcrops in western Hunan and eastern Guizhou in southern China. It contains Burgess Shale-type soft-bodied fossils. It is intermediate in age between the Chengjiang and Ka ...
of China.


History

The history of ''Peytoia'' is entangled with that of "''
Laggania ''Peytoia'' is a genus of hurdiid radiodont that lived in the Cambrian period, containing two species, ''Peytoia nathorsti'' from the Miaolingian of Canada and '' Peytoia infercambriensis'' from Poland, dating to Cambrian Stage 3. Its two fro ...
''" and ''
Anomalocaris ''Anomalocaris'' ("unlike other shrimp", or "abnormal shrimp") is an extinct genus of Radiodonta, radiodont, an Order (biology), order of early-diverging stem-group arthropods. The first fossils of ''Anomalocaris'' were discovered in the ''Ogygop ...
'': all three were initially identified as isolated body parts and only later discovered to belong to one type of animal. This was due in part to their makeup of a mixture of mineralized and unmineralized body parts; the oral cone (mouth) and frontal appendage were considerably harder and more easily
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
ized than the delicate body. The first was a detached frontal appendage of ''Anomalocaris'', described by Joseph Frederick Whiteaves in 1892 as a
phyllocarid Phyllocarida is a subclass of crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapoda, decapods, ostracoda, seed shrimp, branchiopoda, branchiopods, argulidae, fish lice, krill, remipe ...
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
, because it resembled the abdomen of that taxon. The first fossilized oral cone was discovered by
Charles Doolittle Walcott Charles Doolittle Walcott (March 31, 1850February 9, 1927) was an American paleontologist, administrator of the Smithsonian Institution from 1907 to 1927, and director of the United States Geological Survey.Wonderful Life (book) by Stephen Jay G ...
, who mistook it for a
jellyfish Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the subphylum Medusozoa, a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animals with umbrella- ...
and placed it in the genus ''Peytoia''. In the same paper, Walcott described a poorly-preserved body specimen as ''Laggania''; he interpreted it as a holothurian (sea cucumber). In 1978,
Simon Conway Morris Simon Conway Morris (born 1951) is an English palaeontologist, evolutionary biologist, and astrobiologist known for his study of the fossils of the Burgess Shale and the Cambrian explosion. The results of these discoveries were celebrated in ...
noted that the mouthparts of ''Laggania'' were identical to ''Peytoia'', but interpreted this as indicating that ''Laggania'' was a composite fossil of ''Peytoia'' and the sponge '' Corralio undulata''. Later, while clearing what he thought was an unrelated specimen,
Harry B. Whittington Harry Blackmore Whittington FRS (24 March 1916 – 20 June 2010) was a British palaeontologist who made a major contribution to the study of fossils of the Burgess Shale and other Cambrian fauna. His works are largely responsible for the conce ...
removed a layer of covering stone to discover the unequivocally connected arm thought to be a phyllocarid abdomen and the oral cone thought to be a jellyfish. Whittington linked the two species, but it took several more years for researchers to realize that the continuously juxtaposed ''Peytoia'', ''Laggania'' and frontal appendage represented one enormous creature. ''Laggania'' and ''Peytoia'' were named in the same publication, but Conway Morris selected ''Peytoia'' as the valid name in 1978, which makes it the valid name according to
International Commission on Zoological Nomenclature The International Commission on Zoological Nomenclature (ICZN) is an organization dedicated to "achieving stability and sense in the scientific naming of animals". Founded in 1895, it currently comprises 26 commissioners from 20 countries. Orga ...
rules. The discovery that ''Anomalocaris'', ''Laggania'', and ''Peytoia'' represented parts of a single type of animal led to the synonymization of the three genera, with ''Peytoia nathorsti'' reclassified as ''Anomalocaris nathorsti''. ''Peytoia nathorsti'' was subsequently considered a junior synonym of ''Anomalocaris canadensis'', while ''Laggania cambria'' became recognized as a distinct genus and species again, but in 2012 it was determined that ''Anomalocaris canadensis'' had an oral cone with only three large plates, unlike that of ''Laggania cambria'' and ''Peytoia nathorsti'' with four, and ''Peytoia'' was once again recognized as valid, with ''Laggania'' its junior synonym. A second species, ''
Peytoia infercambriensis ''Peytoia infercambriensis'' is a species of hurdiid radiodont in the genus ''Peytoia''. ''P. infercambriensis'' is the geologically oldest known radiodont; its remains date to the third age of the Cambrian. The type and only known specimen ...
'', was named in 1975 as ''Pomerania infercambriensis''. Its discoverer, Kazimiera Lendzion, interpreted it as a member of
Leanchoiliidae Megacheira ("great hands") is an extinct class of predatory arthropods that possessed a pair of great appendages, hence the class name as well as the common name "great appendage arthropods". Their taxonomic position is controversial, with stud ...
, a family which now known as part of the unrelated
megacheira Megacheira ("great hands") is an extinct class of predatory arthropods that possessed a pair of great appendages, hence the class name as well as the common name "great appendage arthropods". Their taxonomic position is controversial, with stud ...
ns (great appendage arthropods). It was subsequently renamed ''Cassubia infercambriensis'' because the name ''Pomerania'' had already been used for an ammonoid. ''C. infercambriensis'' was later recognized as a
radiodont Radiodonta is an extinct Order (biology), order of stem-group arthropods that was successful worldwide during the Cambrian period. They may be referred to as radiodonts, radiodontans, radiodontids, anomalocarids, or anomalocaridids, although the ...
. It was later determined that the specimen was a composite of a radiodont frontal appendage and the body of an unknown arthropod. Due to the close similarity of the appendage to ''Peytoia nathorsti'', ''C. infercambriensis'' was reassigned to ''Peytoia''.


Description

''P. nathorsti'' had body length about . The oral cone of ''Peytoia nathorsti'' has four large plates, similar to ''Hurdia'', as compared to three in ''Anomalocaris.'' However, unlike ''Hurdia'', the oral cone of ''Peytoia'' lacks inner rows of spines. The frontal appendages have 13 podomeres in their distal part, as is typical and likely ancestral for radiodonts. Like other hurdiid radiodonts, the frontal appendages have five blade-like endites, which have short auxillary spines. An intercalary podomere is present, separating the proximal and distal ends of the appendage. The appendages also have large medial spines, sometimes referred to as "gnathites", which face towards the opposite appendage. The trunk consists of 13 segments, which are associated with wide swimming flaps. Compared to ''Hurdia'', ''Peytoia'' has less prominent setal blades.


Ecology

It has been proposed that the frontal appendages of ''Peytoia'' were used to sift sediment for prey, however, some authors have considered this unlikely due to the small size and irregular spacing of the auxillary spines. It has been alternatively proposed that ''Peytoia'' was a predator, using its appendages to capture slow-moving, relatively large
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning "t ...
prey.


References


External links

* {{Taxonbar, from=Q20672130 Burgess Shale fossils Cambrian arthropods Anomalocaridids