- Diagram of Surusicaris
- Diagram of Isoxys curvirostratus
- The holotype, and only known fossil, of Surusicaris
- Fossil of Isoxys acutangulus
Isoxyida Temporal range: | |
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Restoration of Isoxys curvirostratus | |
Restoration of Surusicaris elegans | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Order: | † Isoxyida Simonetta & Delle Cave, 1975 |
Family: | † Isoxyidae Vogdes, 1893 |
Genera | |
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Synonyms | |
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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.
Isoxyids have a combination of features seen in both stem-group arthropods, as well as more advanced taxa. Isoxyids have semicircular bivalved carapaces, large, spherical eyes, a pair of large upward curling frontal appenages covered in spines, and pairs of biramous limbs running along the body. The trunk region appears to be unsegmented and lacks sclerotisation, similar to radiodonts, with a segmented and sclerotised (arthropodized) trunk being characteristic of most arthropods. [1] [2] Currently only two genera have been accepted to be members of this family. These being Isoxys , which has an almost cosmopolitan distribution, and is known from twenty species, and Surusicaris , which is limited to the Burgess shale site in British Columbia, and only contains a single species. These arthropods are thought to have been predators, hunting soft-bodied prey in either the water column, or close to the seabed. [2] [3] [4]
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This cladogram shows the relationships of the isoxyids compared to other arthropod groups, according to the results of Zhang et al. 2023. [1]
The true classification of the isoxyids has been somewhat controversial. Although they are often regarded as basal arthropods, where they actually fit into the arthropod family tree has been contested. Originally, it was thought that Isoxys was related to another bivalved arthropod from the Cambrian, Tuzoia , due to the similar aspects of their carapaces. However, a study in 2022 found that Tuzoia was a member of Hymenocarina, and that it was not closely related to the isoxyids. Currently the group has been placed into the Deuteropoda, a proposed clade of arthropods whose members are distinguished by an anatomical reorganization of the head region, including the appearance of a differentiated first appendage pair (the 'deutocerebral' pair), a multisegmented head, and a hypostome/labrum complex, [5] [6] [7] being placed outside the crown group of Arthropoda, with both Mandibulata and Chelicerata being more closely related to each other than to isoxyids. [1]
Species of the Isoxydia are thought to have been actively swimming predators, using their frontal appendages to capture soft-bodied prey, with the frontalmost pairs of biramous limbs aiding in food processing. [1] The various species of the order are thought to have occupied a variety of niches, from swimming just above the seafloor (nektobenthic) to open ocean swimmers (pelagic). [4] Swimming was likely accomplished by rhythmic movement of the legs. [1] Isoxyid species with elongated carapace spines are likely to have engaged in vertical migration up and down the water column, like many modern marine invertebrates. [4]
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.
Dinocaridida is a proposed fossil taxon of basal arthropods that flourished in the Cambrian period with occasional Ordovician and Devonian records. Characterized by a pair of frontal appendages and series of body flaps, the name of Dinocaridids refers to the suggested role of some of these members as the largest marine predators of their time. Dinocaridids are occasionally referred to as the 'AOPK group' by some literatures, as the group compose of Radiodonta, Opabiniidae, and the "gilled lobopodians" Pambdelurion and Kerygmachelidae. It is most likely paraphyletic, with Kerygmachelidae and Pambdelurion more basal than the clade compose of Opabiniidae, Radiodonta and other arthropods.
Anomalocaris is an extinct genus of radiodont, an order of early-diverging stem-group arthropods.
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.
Megacheira is an extinct class of predatory arthropods defined by their possession of spined "great appendages". Their taxonomic position is controversial, with studies either considering them stem-group euarthropods, or stem-group chelicerates. The homology of the great appendages to the cephalic appendages of other arthropods is also controversial. Uncontested members of the group were present in marine environments worldwide from the lower to middle Cambrian.
Tuzoia 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.
Schinderhannes bartelsi is a species of hurdiid radiodont (anomalocaridid) known from one specimen from the lower Devonian Hunsrück Slates. Its discovery was astonishing because previously, radiodonts were known only from exceptionally well-preserved fossil beds (Lagerstätten) from the Cambrian, 100 million years earlier.
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.
Radiodonta is an extinct 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 last two originally refer to the family Anomalocarididae, which previously included all species of this order but is now restricted to only a few species. Radiodonts are distinguished by their distinctive frontal appendages, which are morphologically diverse and used for a variety of functions. Radiodonts included the earliest large predators known, 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 gainessii, Cambroraster falcatus and Amplectobelua symbrachiata, the Ordovician Aegirocassis benmoulai and the Devonian Schinderhannes bartelsi.
Hurdia is an extinct genus of hurdiid radiodont that lived 505 million years ago during the Cambrian Period. Fossils have been found in North America, China and the Czech Republic.
Marble Canyon is a canyon surrounding Tokumm Creek just above its confluence with the Vermilion River, at the north end of Kootenay National Park in the Canadian Rockies of British Columbia, Canada. South of Marble Canyon on Highway 93 is Numa Falls on the Vermillion River.
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.
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 the presence of mandibles in at least some species.
Titanokorys is a genus of extinct hurdiid radiodont that existed during the mid Cambrian. It is the largest member of its family from the Cambrian, with a body length of 50 cm (20 in) long, making it one of the largest animals of the time. It bears a resemblance to the related genus Cambroraster. Fossils of T. gainesi were first found within Marble Canyon in 2018. The fossils were not named until 2021 because they were assumed to be giant specimens of Cambroraster.
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.
Balhuticaris is a genus of extinct bivalved hymenocarine arthropod that lived in the Cambrian aged Burgess Shale in what is now British Columbia around 506 million years ago. This extremely multisegmented arthropod is the largest member of the group, and it was even one of the largest animals of the Cambrian, with individuals reaching lengths of 245 mm (9 in). Fossils of this animal suggests that gigantism occurred in more groups of Arthropoda than had been previously thought. It also presents the possibility that bivalved arthropods were very diverse, and filled in a lot of ecological niches.
Pahvantia is an extinct genus of hurdiid radiodont from the Cambrian. It is known by a single species, Pahvantia hastata, described from Wheeler Shale and Marjum Formation in Utah. Although it was once considered as filter feeder using large number of putative setae, this structures are later considered as misidentification of trunk materials.
Surusicaris is an extinct genus of bivalved arthropod, known from the Cambrian Burgess Shale of British Columbia, Canada. It is considered to be closely related to Isoxys, and like it has spined grasping frontal appendages.