Tuzoia
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''Tuzoia'' (from
Mount Tuzo Mount Tuzo, previously "Peak Seven" or "Shagowa", was named in 1907 after its first ascendant Henrietta L. Tuzo. It is located within the Valley of the Ten Peaks in the Canadian Rockies, along the Continental Divide, which forms the boundary be ...
, a mountain in the
Canadian Rockies The Canadian Rockies (french: Rocheuses canadiennes) or Canadian Rocky Mountains, comprising both the Alberta Rockies and the British Columbian Rockies, is the Canadian segment of the North American Rocky Mountains. It is the easternmost part ...
) is an extinct genus of large bivalved
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
known from Early to Middle
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 ...
marine environments from what is now North America, Australia, China, Europe and Siberia. The large, domed carapace reached lengths of , making them amongst the largest known Cambrian arthropods.


Description

The largest carapaces of ''Tuzoia'' are known to reach in length, suggesting a total body length of approximately . Along the sides of the carapace a spiked ridge is present, and the edges and midline of the carapace are also spiked in many species. These spines likely functioned to aid stability while swimming. The carapaces are marked with a reticulate (net-like) pattern, which was likely to increase the structural integrity of the valves while remaining lightweight. The head had a anterior/ocular sclerite at the top, a single pair of large stalked eyes, and a pair of segmented antennae. The head has pairs of cephalic appendages, which have curved setae. The first two limb pairs, dubbed the "cephalothoracic legs" have 7 podomeres, with spinose endites, with the legs ending in a terminal claw. The basipods are greatly elongated, and bears 5 to 6 endites, which become spinose at their ends. It is unclear whether exopods are present on these limbs. There are likely 10 pairs of trunk legs excluding the cephalothoracic legs. These had seven podomeres, the first with a spinose endite, with the limb ending in a terminal claw, and a similarly elongated basipod, which had a spine proximal to that of the first podomere. These limbs likely had paddle-shaped exopods. The body terminates with a large tail fan.


Ecology

''Tuzoia'' was likely an actively swimming organism. It had historically been considered a
pelagic The pelagic zone consists of the water column of the open ocean, and can be further divided into regions by depth (as illustrated on the right). The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or wa ...
organism, due to their wide distribution. However, the soft tissues described in 2022 suggest an alternative interpretation as an animal that swam close to the seafloor (nektobenthic) as a predator or a scavenger. ''Tuzoia'' was likely capable of walking along the seafloor with the carpace flexing outwards to allow the legs to contact the substrate.


Taxonomy

They were formerly considered to be the only members of the family Tuzoiidae until 2022, when a new genus ''
Duplapex ''Duplapex'' is an extinct genus of bivalved arthropod known from the Cambrian Stage 3 aged Qingjiang biota of Hubei, China, with a single species. ''D. anima''. It is thought to be a close relative of ''Tuzoia ''Tuzoia'' (from Mount Tuzo, a m ...
'' was described from the Cambrian of China, which differs from ''Tuzoia'' by having a flat carapace. Their phylogenetic position was long uncertain due to the lack of soft tissue remains, a close relationship with ''
Isoxys ''Isoxys'' (meaning "equal surfaces") is a genus of extinct bivalved Cambrian arthropod; the various species of which are thought to have been freely swimming predators. It had a pair of large spherical eyes (which are the most commonly preserve ...
'' was proposed based on similarities of their carapaces. However soft tissues described in 2022 suggest instead that ''Tuzoia'' was a member of
Mandibulata Mandibulata, termed "mandibulates", is a clade of arthropods that comprises the extant subphyla Myriapoda (millipedes and others), Crustacea and Hexapoda (insects and others). Mandibulata is currently believed to be the sister group of the clade ...
, as the earliest diverging member of a paraphyletic
Hymenocarina Hymenocarina is an order of extinct arthropods known from the Cambrian. They possess bivalved carapaces, typically with exposed posteriors. Members of the group are morphologically diverse and had a variety of ecologies, including as filter feed ...
, and that similarities to isoxyids may be due to retained
plesiomorphies In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, and ...
.


Species

After Vannier et al. 2007. and subsequent literature. * ''Tuzoia'' ''retifera'' Walcott, 1912 ( type)
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 ...
, Canada, Utah, USA * ''Tuzoia'' ''burgessensis'' Resser, 1929, Burgess Shale, Canada * ''Tuzoia'' ''guntheri'' Robison & Richards, 1981
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 ...
Utah, USA * ''Tuzoia'' ''bispinosa'' Yuan and Zhao, 1999 Kaili Formation,
Guizhou Guizhou (; formerly Kweichow) is a landlocked province in the southwest region of the People's Republic of China. Its capital and largest city is Guiyang, in the center of the province. Guizhou borders the autonomous region of Guangxi to t ...
, China * ''Tuzoia polleni'' Resser, 1929
Eager Formation Eager may refer to: *Eager (band) * Eager (horse), (1788 – after 1795), a British Thoroughbred racehorse * ''Eager'' (novel), a children's science-fiction novel written by Helen Fox * Eager (surname) *, an Admirable-class minesweeper built for t ...
, Canada,
Parker Shale Parker may refer to: Persons * Parker (given name) * Parker (surname) Places Place names in the United States *Parker, Arizona *Parker, Colorado * Parker, Florida * Parker, Idaho *Parker, Kansas * Parker, Missouri * Parker, North Carolina *Parke ...
, Vermont,
Pioche Shale The Pioche Shale is an Early to Middle Cambrian Burgess shale-type Lagerstätte in Nevada. It spans the Early–Middle Cambrian boundary; fossils from the Early Cambrian are preserved in botryoidal hematite, whereas those from the Middle Cambria ...
, Nevada
Kinzers Formation The Kinzers Formation is a geologic formation in Pennsylvania. It preserves fossils dating back to the Cambrian Period. The base of the Kinzers Formation is primarily a dark-brown shale. The middle is a gray and white spotted limestone and, lo ...
, Pennsylvania, USA * ''Tuzoia australis'' Glaessner, 1979
Emu Bay Shale The Emu Bay Shale is a geological formation in Emu Bay, South Australia, containing a major Konservat-Lagerstätte (fossil beds with soft tissue preservation). It is one of two in the world containing Redlichiidan trilobites. The Emu Bay Shale ...
, Australia * ''Tuzoia manchuriensis'' Resser and Endo, 1937 Tangshih Formation Liaoning, China (poorly known) * ''Tuzoia sinensis'' P'an, 1957 Wulongqing Formation Yunnan, China (poorly known) * ''Tuzoia tylodesa'' Luo et Hu et al., 2006 Wulongqing Formation Yunnan, China * ''Tuzoia multispinosa'' Zhao, 2015 Wulongqing Formation Yunnan, China * ''Tuzoia lazizhaiensis'' Wen et al., 2019
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 ...
, Hunan, China. * ''Tuzoia jianheensis'' Chen and Zhao, 2017 Tsinghsutung Formation, Guzihou, China Indeterminate remains are also known from the Buchava Formation and Jince Formation of the Czech Republic. As well as the Sinsk Lagerstatte of Siberia.Ivantsov, A. I., Zhuravlev, A. I., Krasilov, V. A., Leguta, A. V., Melnikova, L. M., Urbanek, A., et al. (2005). ''Unique Sinsk Localities of Early Cambrian Organisms (Siberia Platform)''. Moscow: Nauka. Rossiyskaya Akademia Nauk, 143. n Russian


References


External links

* {{Taxonbar, from=Q148514 Burgess Shale fossils Wheeler Shale Cambrian genus extinctions Hymenocarina