Dinocaridida Temporal range: | |
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Assembly of dinocaridids ( Anomalocaris , Opabinia , Pambdelurion and Kerygmachela ) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | † Dinocaridida Collins, 1996 |
Subgroups | |
Dinocaridida [derivation 1] is a proposed fossil taxon [1] of basal arthropods that flourished in the Cambrian period with occasional Ordovician [2] and Devonian records. [3] Characterized by a pair of frontal appendages and series of body flaps, the name of Dinocaridids (Greek for deinos "terrible" and Latin for caris "crab") refers to the suggested role of some of these members as the largest marine predators of their time. [1] Dinocaridids are occasionally referred to as the 'AOPK group' by some literatures, [4] [5] [6] as the group compose of Radiodonta ( Anomalocaris and relatives), Opabiniidae ( Opabinia and relatives), and the "gilled lobopodians" Pambdelurion and Kerygmachelidae. [7] It is most likely paraphyletic, with Kerygmachelidae and Pambdelurion more basal than the clade compose of Opabiniidae, Radiodonta and other arthropods. [4] [8] [9]
Dinocaridids were bilaterally symmetrical, with a mostly non-mineralized cuticle and a body divided into two major groupings of tagmata (body-sections): head and trunk. The head apparently unsegmented [8] [9] and had a pair of specialized frontal appendages just in front of the mouth and eyes. The frontal appendages are either lobopodous (soft as in gilled lobopodians) or arthropodized (hardened and segmented as in Radiodonta) and usually paired, but highly fused into a nozzle-like structure in Opabiniidae. [7] Based on their preocular position and putative protocerebral origin, the frontal appendages are generally thought to be homologous to the labrum of euarthropods and primary antennae of onychophoran, [10] [9] while subsequent evidence also suggest a deutocerebral origin (homologous to the jaws of onychophora and great appendages/antennae/chelicerae of euarthropods). [11] [12] The trunk possessed multiple segments, each with its own gill branch and swimming flaps (lobes). [13] It is thought that these flaps moved in an up-and-down motion, in order to propel the animal forward [14] in a fashion similar to the cuttlefish. In gilled lobopodian genera, the trunk may have borne a lobopodous limb (lobopod) underneath each of the flaps. [13] The midgut of dinocaridids had paired digestive glands similar to those of siberiid lobopodians and Cambrian euarthropods. [15] The dinocaridid brain is relatively simple than those of a euarthropod (3-segmented), it is thought to be comprised either 1 (only protocerebrum [10] [16] ) or 2 cerebral ganglions (protocerebrum and deutocerebrum). [12]
| Dinocaridida |
Although some authors may rather suggest different taxonomic affinities (e.g. as cycloneuralia n relatives [17] [7] ), most of the phylogenetic studies suggest that dinocaridids are stem group arthropods. [18] [19] [20] [21] [13] [8] [9] [22] [23] [24] Under this scenario, Dinocaridida is a paraphyletic grade in correspond to the arthropod crown group (Euarthropoda or Deuteropoda) and also suggest a lobopodian origin of the arthropod lineage. [25] [8] In general, the gilled lobopodian genera Pambdelurion and Kerygmachela which have lobopodian traits (e.g. lobopodous appendage, annulation) occupied the basal position; while Opabiniidae and Radiodonta are more derived and closely related to the arthropod crown group, with the latter even having significant arthropod affinities such as arthropodization and head sclerites. [24] [26] [8]
In the original description, Dinocaridida was composed of only Opabiniidae and Radiodonta. [1] With the exclusion of questionable taxa (e.g. the putative opabiniid Myoscolex [27] ), the former were known only by Opabinia, while all radiodont species were grouped under a single family: Anomalocarididae (hence the previous common name 'Anomalocaridids' [18] ). In later studies, the gilled lobopodians Pambdelurion and Kerygmachela were also regarded to be dinocaridids, [7] two new opabiniid genera, Utaurora and Mieridduryn were described, [28] [29] [30] other strange dinocaridids like Parvibellus (which might actually be a juvenile siberiid lobopodian), [31] [29] many radiodonts were reassigned to other new families (Amplectobeluidae, Tamisiocarididae and Hurdiidae), [21] and a new family, Kerygmachelidae, [29] was named.
The group was geographically widespread, and has been reported from Cambrian strata in Canada, United States, Greenland, China, Australia and Russia, [32] as well as the Ordovician of Morocco and Wales [2] [33] [30] and Devonian of Germany. [3]
Lobopodians are members of the informal group Lobopodia, or the formally erected phylum Lobopoda Cavalier-Smith (1998). They are panarthropods with stubby legs called lobopods, a term which may also be used as a common name of this group as well. While the definition of lobopodians may differ between literatures, it usually refers to a group of soft-bodied, marine worm-like fossil panarthropods such as Aysheaia and Hallucigenia.
Opabinia regalis is an extinct, stem group arthropod found in the Middle Cambrian Burgess Shale Lagerstätte of British Columbia. Opabinia was a soft-bodied animal, measuring up to 7 cm in body length, and its segmented trunk had flaps along the sides and a fan-shaped tail. The head shows unusual features: five eyes, a mouth under the head and facing backwards, and a clawed proboscis that probably passed food to the mouth. Opabinia probably lived on the seafloor, using the proboscis to seek out small, soft food. Fewer than twenty good specimens have been described; 3 specimens of Opabinia are known from the Greater Phyllopod bed, where they constitute less than 0.1% of the community.
Anomalocaris is an extinct genus of radiodont, an order of early-diverging stem-group arthropods.
Anomalocarididae is an extinct family of Cambrian radiodonts, a group of stem-group arthropods.
Kerygmachela kierkegaardi is a kerygmachelid gilled lobopodian from the Cambrian Stage 3 aged Sirius Passet Lagerstätte in northern Greenland. Its anatomy strongly suggests that it, along with its relative Pambdelurion whittingtoni, was a close relative of radiodont and euarthropods. The generic name "Kerygmachela" derives from the Greek words Kerygma (proclamation) and Chela (claw), in reference to the flamboyant frontal appendages. The specific name, "kierkegaardi" honors Danish philosopher Søren Kierkegaard.
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.
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.
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.
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.
Cucumericrus ("cucumber-leg") is an extinct genus of stem-arthropod. The type and only species is Cucumericrus decoratus, with fossils discovered from the Maotianshan Shales of Yunnan, China.
Caryosyntrips ("nutcracker") is an extinct genus of stem-arthropod which known from Canada, United States and Spain during the middle Cambrian.
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. There is also a potential fifth genus, Houcaris, but that genus has become problematic in terms of its taxonomic placement.
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, measuring up to 8 cm (3.1 in) in total body length.
Aegirocassis is an extinct genus of giant radiodont arthropod belonging to the family Hurdiidae that lived 480 million years ago during the early Ordovician in the Fezouata Formation of Morocco. It is known by a single species, Aegirocassis benmoulai. Van Roy initiated scientific study of the fossil, the earliest known of a "giant" filter-feeder discovered to date. Aegirocassis is considered to have evolved from early predatory radiodonts. This animal is characterized by its long, forward facing head sclerite, and the endites on its frontal appendages that bore copious amounts of baleen-like auxiliary spines. This animal evolving filter-feeding traits was most likely a result of the Great Ordovician Biodiversification Event, when environmental changes caused a diversification of plankton, which in turn allowed for the evolution of new suspension feeding lifeforms. Alongside the closely related Pseudoangustidontus, an unnamed hurdiid from Wales, the middle Ordovician dinocaridid Mieridduryn, and the Devonian hurdiid Schinderhannes this radiodont is one of the few dinocaridids known from post-Cambrian rocks.
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. It contains two species, Houcaris saron and Houcaris magnabasis, both of which were originally named as species of the related genus Anomalocaris. 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.
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.
Deuteropoda is a proposed clade of arthropods whose members are distinguished from more basal stem-group arthropods like radiodonts by an anatomical reorganization of the head region, namely the appearance of a differentiated first appendage pair, a multisegmented head, a hypostome/labrum complex, and by bearing pairs of segmented biramous limbs.
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.
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