Amplectobeluidae | |
---|---|
![]() | |
Amplectobelua (left) and Lyrarapax (right) | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | † Dinocaridida |
Order: | † Radiodonta |
Clade: | † Amplectobeluidae Pates et al., 2019 |
Genera | |
|
Amplectobeluidae is a clade of Cambrian radiodonts. It currently includes five definitive genera, Amplectobelua , Lyrarapax , Ramskoeldia , Guanshancaris and a currently unnamed genus from the lower Cambrian aged Sirius Passet site in Greenland. [1] There is also a potential fifth genus, Houcaris , but that genus has become problematic in terms of its taxonomic placement. [2] [3]
In 2014, Amplectobeluidae was defined as the most inclusive clade including Amplectobelua symbrachiata but not Anomalocaris canadensis , Tamisiocaris borealis , or Hurdia victoria . [4]
Amplectobeluids could be recognized by frontal appendages with well-developed first distal endite, which forming a pincer-like structure that presumably better suited for a grasping function. [4] [5] Complete body fossils of amplectobeluids are only known by Amplectobelua and Lyrarapax , both showing combination of characters resembling Anomalocaris (i.e. streamlined body; small head with ovoid sclerites; well-developed swimming flaps; a pair of caudal furcae). [6] [7] [8] [9] [10] Another distinctive features only known in amplectobeluid genera were pairs of gnathobase-like structures (known by Amplectobelua and Ramskoeldia ), [9] [11] or an oral cone with combination of tetraradial arrangement and scale-like nodes (known by Lyrarapax and Guanshancaris). [8] [12] [13]
Early in 2014, "Anomalocaris" kunmingensis was tentatively assigned to Amplectobelua by Vinther et al. [4] Later that year, however, the discoverers of Lyrarapax unguispinus ignored that assessment and created a genus within Amplectobelua sensu Vinther et al. [7] Indeterminate frontal appendages assignable to this group are known from the Parker Formation of Vermont. [14]
An a posteriori-weighted phylogenetic analysis in 2014 found the following relationships within the Amplectobeluidae: [7]
Dinocaridida is a proposed fossil taxon of basal arthropods, which flourished during the Cambrian period and survived up to the Early to Middle Ordovician. The putative Early Devonian member Schinderhannes bartelsi has been considered questionable by some researchers.
Anomalocaris is an extinct genus of radiodont, an order of early-diverging stem-group arthropods.
Peytoia is a genus of hurdiid radiodont, an early diverging order of stem-group arthropods, 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 frontal appendages had long bristle-like spines, it had no fan tail, and its short stalked eyes were behind its large head.
Amplectobelua is an extinct genus of late Early Cambrian amplectobeluid radiodont, a group of stem arthropods that mostly lived as free-swimming predators during the first half of the Paleozoic Era.
Anomalocarididae is an extinct family of Cambrian radiodonts, a group of stem-group arthropods.
Pambdelurion is an extinct genus of panarthropod from the Cambrian aged Sirius Passet site in northern Greenland. Like the morphologically similar Kerygmachela from the same locality, Pambdelurion is thought to be closely related to arthropods, combining characteristics of "lobopodians" with those of primitive arthropods.
Parapeytoia is a genus of Cambrian arthropod. The type and only described species is Parapeytoia yunnanensis, lived over 518 million years ago in the Maotianshan shales of Yunnan, China. Unidentified fossils from the same genus also had been discovered from the nearby Wulongqing Formation.
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 latest surviving members include the subfamily Aegirocassisinae from the Early Ordovician of Morocco. The putative Early Devonian member Schinderhannes bartelsi from Germany has been considered questionable by some researchers.
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.
Caryosyntrips ("nutcracker") is an extinct genus of stem-arthropod which known from Canada, United States and Spain during the middle Cambrian.
Tamisiocaris is a radiodont genus from the Cambrian period. The taxon was initially described in 2010 based on frontal appendages discovered from the Sirius Passet lagerstatte in northern Greenland.. A subsequent study by Vinther and colleagues in 2014 revealed that the frontal appendages were segmented and bore densely packed auxiliary spines, which were adapted to suspension feeding in a manner analogous to modern baleen whales. It is assigned to the family Tamisiocarididae, and is measured about 22.8–33.6 cm (0.75–1.10 ft) long.
Tamisiocarididae is a family of radiodonts, extinct marine animals related to arthropods, that bore finely-spined appendages that were presumably used in filter-feeding. When first discovered, the clade was named Cetiocaridae after a speculative evolution artwork, Bearded Ceticaris by John Meszaros, that depicted a hypothetical filter-feeding radiodont at a time before any were known to exist. However, the family name was not valid according to the International Code of Zoological Nomenclature, as no real genus named "Cetiocaris" exists, and in 2019 it was formally replaced by the name Tamisiocarididae, after the only valid genus of the clade at the time. The family is only known from Series 2 of the Cambrian, unlike other radiodont families, which persisted longer into the Cambrian. All known species would have lived in tropical or subtropical waters, suggesting a preference for warmer waters.
Lyrarapax is a radiodont genus of the family Amplectobeluidae that lived in the early Cambrian period 520 million years ago. Its neural tissue indicates that the radiodont frontal appendage is protocerebral, resolving parts of the arthropod head problem and showing that the frontal appendage is homologous to the antennae of Onychophorans and labrum of euarthropods. Its fossilized remains were found in Yunnan in southwestern China. A second species was described in 2016, differing principally in the morphology of its frontal appendages. It is a small animal, with L. unguispinus and L. trilobus measuring up to 8.3 cm (3.3 in) and 3.9 cm (1.5 in) in body length respectively.
Ramskoeldia is a genus of amplectobeluid radiodont described in 2018. It was the second genus of radiodont found to possess gnathobase-like structures and an atypical oral cone after Amplectobelua. The type species, Ramskoeldia platyacantha, was discovered in the Chengjiang biota of China, the home of numerous radiodontids such as Amplectobelua and Lyrarapax.
Cambroraster is an extinct monotypic genus of hurdiid radiodont, dating to the middle Cambrian, and represented by the single formally described species Cambroraster falcatus. Hundreds of specimens were found in the Burgess Shale, and described in 2019. A large animal at up to 30 centimetres (12 in), it is characterized by a significantly enlarged horseshoe-shaped dorsal carapace (H-element), and presumably fed by sifting through the sediment with its well-developed tooth plates and short frontal appendages with hooked spines. Nicknamed the "spaceship" fossil when first found, for the way its dorsal carapace resembles the fictional Millennium Falcon, the specific epithet falcatus in its scientific name is a nod to that resemblance.
Squamacula is an extinct artiopodan arthropod from the Cambrian Series 2. The type species S. clypeata was described in 1997 from the Chengjiang biota of Yunnan, China. At the time of description there were only two known specimens of S. clypeata, but now there are at least six known specimens. In 2012 a second species S. buckorum was described from the Emu Bay Shale of Australia.
Houcaris is a possibly paraphyletic radiodont genus, tentatively assigned to either Amplectobeluidae, Anomalocarididae or Tamisiocarididae, known from Cambrian Series 2 of China and the United States. The type species is Houcaris saron which was originally described as a species of the related genus Anomalocaris. Other possible species include H. magnabasis and H. consimilis. The genus Houcaris was established for the two species in 2021 and honors Hou Xianguang, who had discovered and named the type species Anomalocaris saron in 1995 along with his colleagues Jan Bergström and Per E. Ahlberg.
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.
Pseudoangustidontus is a genus of hurdiid (peytoiid) radiodont from the Lower Ordovician of Morocco. This genus is known from two described species, P. duplospineus and P. izidigua, with some specimens that are hard to determine which species to belong to. This animal is only known from the Fezouata Formation, a fossil site in Morocco that is of Lagerstätte status, meaning that the fossils from this site are exceptionally preserved. Because of its partial remains, its classification was debated, but with more complete fossils it is identified as radiodont frontal appendage.
Guanshancaris is an extinct genus of amplectobeluid radiodont known from the Cambrian Stage 4 Guanshan Biota of southern China. It is only known from a single species. G. kunmingensis which was described in 2013 as a species of Anomalocaris before being placed in a new genus in 2023. Like many other radiodonts, it is only known from fragmentary remains, including its frontal appendages and the oral cone. It has been suggested to have been durophagous. It is estimated to have reached up to 22.2–32.8 cm (8.7–12.9 in) long.