Tuzoia Temporal range: | |
---|---|
Carapace of Tuzoia retifera | |
Life restoration | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Order: | † Hymenocarina |
Family: | † Tuzoiidae |
Genus: | † Tuzoia Walcott, 1912 |
Type species | |
Tuzoia retifera Walcott, 1912 | |
Species | |
Tuzoia (from Mount Tuzo, a mountain in the Canadian Rockies) is an extinct genus of large bivalved arthropod known from Early to Middle Cambrian marine environments from what is now North America, Australia, China, Europe and Siberia. The large, domed carapace reached lengths of 180 millimetres (7.1 in), making them amongst the largest known Cambrian arthropods.
The largest carapaces of Tuzoia are known to reach 18 cm (7.1 in) in length, suggesting a total body length of approximately 23 cm (9.1 in). [1] 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. [2] [3] 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. [1]
Tuzoia was likely an actively swimming organism. It had historically been considered a pelagic 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. [1]
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 was proposed based on similarities of their carapaces. [4] However soft tissues described in 2022 suggest instead that Tuzoia was a member of Mandibulata, as the earliest diverging member of a paraphyletic Hymenocarina, and that similarities to isoxyids may be due to retained plesiomorphies. [1]
After Vannier et al., 2007, and subsequent literature: [2] [5]
Indeterminate remains are also known from the Buchava Formation and Jince Formation of the Czech Republic. [2] [9] As well as the Sinsk Biota of Siberia. [10] [5]
The Maotianshan Shales (帽天山页岩) are a series of Early Cambrian sedimentary deposits in the Chiungchussu Formation, famous for their Konservat Lagerstätten, deposits known for the exceptional preservation of fossilized organisms or traces. The Maotianshan Shales form one of some forty Cambrian fossil locations worldwide exhibiting exquisite preservation of rarely preserved, non-mineralized soft tissue, comparable to the fossils of the Burgess Shale of British Columbia, Canada. They take their name from Maotianshan Hill in Chengjiang County, Yunnan Province, China.
Sidneyia is an extinct arthropod known from fossils found from the Early to the Mid Cambrian of China and the Mid Cambrian Burgess Shale of British Columbia, Canada.
Canadaspis is an extinct genus of bivalved Cambrian arthropod, known from North America and China. They are thought to have been benthic feeders that moved mainly by walking and possibly used its biramous appendages to stir mud in search of food. They have been placed within the Hymenocarina, which includes other bivalved Cambrian arthropods.
Waptia is an extinct genus of arthropod from the Middle Cambrian of North America. It grew to a length of 6.65 cm (3 in), and had a large bivalved carapace and a segmented body terminating into a pair of tail flaps. It was an active swimmer and likely a predator of soft-bodied prey. It is also one of the oldest animals with direct evidence of brood care. Waptia fieldensis is the only species classified under the genus Waptia, and is known from the Burgess Shale Lagerstätte of British Columbia, Canada. Specimens of Waptia are also known from the Spence Shale of Utah, United States.
Chuandianella ovata is an extinct bivalved arthropod that lived during Cambrian Stage 3 of the Early Cambrian. It is the only species classified under the genus Chuandianella. Its fossils were recovered from the Chengjiang Biota in Yunnan, China.
Clypecaris is genus of bivalved Cambrian arthropod known from the Chengjiang biota of Yunnan, China. The genus was initially described for the type species C. pteroidea by Hou, 1999. A second species C. serrata was described by Yang et al. in 2016. The species are primarily distinguished by the presence of a serrated edge on the front of the carapace of C. serrata. C. serrata is noted for the modification of an anterior pair of limbs into spined grasping appendages, indicating a predatory lifestyle. It is unknown whether a similar structure was present in C. pteroidea.Clypecaris is considered to likely be a member or a close relative of Hymenocarina, and is closely related to Perspicaris. As well as to Ercaicunia.
Isoxys 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, and two large frontal appendages used to grasp prey.
Pectocaris is an extinct genus of bivalved arthropods from the Cambrian Maotianshan Shales, Yunnan Province of China. There are currently four known species within the genus.
Primicaris is genus of Cambrian arthropod from the Chengjiang biota of China and the Burgess Shale of Canada. It contains a single described species, P. larvaformis.
Radiodonta is an extinct order of stem-group arthropods that was successful worldwide during the Cambrian period. Radiodonts are distinguished by their distinctive frontal appendages, which are morphologically diverse and used for a variety of functions. Radiodonts are among the earliest large predators, but they also included sediment sifters and filter feeders. Some of the most famous species of radiodonts are the Cambrian taxa Anomalocaris canadensis, Hurdia victoria, Peytoia nathorsti, Titanokorys gainesi, Cambroraster falcatus and Amplectobelua symbrachiata. The later surviving members include the subfamily Aegirocassisinae from the Early Ordovician of Morocco and the Early Devonian member Schinderhannes bartelsi from Germany.
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.
Tokummia is a genus of fossil hymenocarine arthropod, known only by one species, Tokummiakatalepsis, from the middle Cambrian Burgess Shale as found in a quarry in Marble Canyon in Canada.
The Artiopoda is a grouping of extinct arthropods that includes trilobites and their close relatives. It was erected by Hou and Bergström in 1997 to encompass a wide diversity of arthropods that would traditionally have been assigned to the Trilobitomorpha. Trilobites, in part due to abundance of findings owing to their mineralized exoskeletons, are by far the best recorded, diverse, and long lived members of the clade. Other members, which lack mineralised exoskeletons, are known mostly from Cambrian deposits.
Retifacies abnormalis is an extinct arthropod that lived in the lower Cambrian. Its fossil remains have been found in the Maotianshan Shales of Yunnan, China. It is a member of the Artiopoda, and closely related to Pygmaclypeatus.
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 feeders and as predators. Recent research has generally considered them to be stem or crown group members of Mandibulata, due to the presence of mandibles in some species.
Erratus is an extinct genus of marine arthropod from the Cambrian of China. Its type and only species is Erratus sperare. Erratus is likely one of the most basal known arthropods, and its discovery has helped scientists understand the early evolution of arthropod trunk appendages. Some of the stem-arthropods like radiodonts did not have legs, instead they had flap like appendages that helped them swim. Erratus on the other hand had not only flaps but also a set of primitive legs. It also supported the theory that the gills of aquatic arthropods probably evolved into the wings and lungs of terrestrial arthropods later in the Paleozoic.
Laminacaris is a genus of extinct stem-group arthropods (Radiodonta) that lived during the Cambrian period. It is monotypic with a single species Laminacaris chimera, the fossil of which was described from the Chengjiang biota of China in 2018. Around the same time, two specimens that were similar or of the same species were discovered at the Kinzers Formation in Pennsylvania, USA. The first specimens from China were three frontal appendages, without the other body parts.
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, but unlike it, the reticulated bivalved carapace was interpreted to have opened at a wide angle. The carapace had a pair of spines projecting posteriorly. In its original description, it was suggested to have eyes on annulated stalks, but later research suggested that these may have been preservational artifacts instead.
Jugatacaris is an extinct genus of bivalved arthropod known from a single species, Jugatacaris agilis found in the Cambrian Stage 3 aged Chengjiang biota of Yunnan, China. The carapace is around 28 to 37 millimetres in length, with a pronounced ridge at the top of the carapace separating the two valves, which formed a fin-like structure raised above the carapace. The head has a pair of stalked eyes, as well as a dumbbell shaped medial eye between them. The head also bore a pair of mandibles as well as at least one and possibly two pairs of antennules. The trunk had up to 65 segments, each with biramous appendages. The appendages had thin endopods with 30 podomeres, each bearing a spiny endite, with the endopods ending with a terminal claw. The appendages also had overlapping flap-like exopods, which are elongated, being at maximum eight times as long as they are wide, which on their posterior edge are covered with setae. The trunk ended with a forked tail. It was likely an actively swimming filter feeder, using its constantly beating appendages to sift food from the water column, which was then passed forward along the U-shaped food groove between the appendage pairs towards the mouth. While initially placed as a crustaceanomorph, later studies considered to be a member of Hymenocarina, which contains numerous other similar bivalved Cambrian arthropods.
Isoxyids are members of the order Isoxyida and the family Isoxyidae, a group of basal arthropods that existed during the Cambrian period. It contains two genera, Isoxys, with 20 species found worldwide, and Surusicaris known from a single species found in the Burgess Shale of Canada. They are distinguished by their bivalved carapaces and pair of upward curving grasping frontal appendages.