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''Tuzoia'' (from
Mount Tuzo Mount Tuzo is a mountain located within the Valley of the Ten Peaks in the Canadian Rockies, along the Continental Divide, which forms the provincial boundary between British Columbia and Alberta in Western Canada. It also lies on the boundary sh ...
, a mountain in the
Canadian Rockies The Canadian Rockies () 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 of the Canadian Cordillera, w ...
) is an extinct genus of large bivalved
arthropod 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 ...
known from Early to Middle
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 ...
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 also bore 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. The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or water column between the sur ...
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'' 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 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 "Mandibul ...
, as the earliest diverging member of a paraphyletic
Hymenocarina Hymenocarina is an Order (biology), order of extinct marine 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, i ...
, 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, an ...
.


Species

After Vannier ''et al.'', 2007, and subsequent literature: * ''Tuzoia'' ''retifera'' Walcott, 1912 (
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. * ...
)
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 fos ...
, 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 named after its type locality, Marjum Pass, and was defined in 1908. The formation is known for its occasional preservation of ...
Utah, USA * ''Tuzoia'' ''bispinosa'' Yuan and Zhao, 1999
Kaili Formation The Kaili Formation () is a Formation (stratigraphy), stratigraphic formation which was deposited during the Lower and Middle Cambrian (~513 to 506 million years ago). The formation is approximately thick and was named after the city Kaili City, ...
,
Guizhou ) , image_skyline = , image_caption = , image_map = Guizhou in China (+all claims hatched).svg , mapsize = 275px , map_alt = Map showing the location of Guizhou Province , map_caption = Map s ...
, China * ''Tuzoia polleni'' Resser, 1929 Eager Formation, Canada,
Parker Shale Parker may refer to: People * Parker (given name), including a list of people and fictional characters with the given name * Parker (surname), including a list of people and fictional characters with the surname Arts and entertainment * ''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 Cambrian ...
, Nevada
Kinzers Formation The Kinzers Formation is a geologic formation in Pennsylvania. It preserves fossils dating back to the fourth stage of the Cambrian Period. The base of the Kinzers Formation is primarily a dark-brown shale. The middle is a gray and white spotte ...
, Pennsylvania, USA * ''Tuzoia australis'' Glaessner, 1979
Emu Bay Shale The Emu Bay Shale is a Formation (stratigraphy), 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 trilob ...
, 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, Cambrian Period geological formation, which outcrops in western Hunan and eastern Guizhou in southern China. It contains Burgess Shale-type preservation, Burgess Shale-type soft-bodied fossils. It is intermedi ...
, 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 Biota The Sinsk Formation is a geologic formation in Central Siberia, Russia, exposed along the Lena River. It preserves fossils dating back to the Botomian stage of the Cambrian (corresponding to Cambrian Series 2). It notably contained the Sinsk Alga ...
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 N, or n, is the fourteenth letter of the Latin alphabet, used in the modern English alphabet, the alphabets of other western European languages, and others worldwide. Its name in English is ''en'' (pronounced ), plural ''ens''. History ...
Phylogeny of Hymenocarina after Izquierdo-López and Caron (2024)


See also

*
Paleobiota of the Burgess Shale This is a list of the Biota (ecology), biota of the Burgess Shale, a Cambrian lagerstätte located in Yoho National Park in Canada. The Burgess Shale is a fossil-bearing deposit exposed in the Canadian Rockies of British Columbia, Canada. It is fa ...


References


External links

* {{Taxonbar, from=Q148514 Burgess Shale fossils Wheeler Shale Cambrian genus extinctions Hymenocarina Fossil taxa described in 1912 Taxa named by Charles Doolittle Walcott