Deuteropoda | |
---|---|
Life restoration of Kylinxia zhangi , a basal deuteropod | |
Life restoration of Surusicaris , a likely basal deuteropod | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
(unranked): | Deuteropoda Ortega-Hernández 2016 [1] |
Subphyla and classes | |
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 (the 'deutocerebral' pair), a multisegmented head, a hypostome/labrum complex, and by bearing pairs of segmented biramous limbs. [1]
The clade contains all living arthropods (i.e. chelicerates and mandibulates) as well as several fossil groups that share these characteristics (e.g. Fuxianhuiida Megacheira, Isoxyida, and Artiopoda), while excluding other fossil groups that are more 'basal' or 'primitive' (e.g. radiodonts, opabiniids and lobopodians). [1] [3] Members of Deuteropda that are basal to the last common ancestor of chelicerates and mandibulates (and thus lie outside the crown group of Arthropoda) are often referred to as "upper stem group arthropods". [1]
Members of Deuteropoda are characterized by the presence of a differentiated labrum and a differentiated first 'deutocerebral' pair of appendages. [1] In contrast, lobopodians (part of the "lower stem group") and onychophorans have a pair of pre-ocular or 'protocerebral' appendages, which presumably have evolved to be the labrum of modern living arthropods. [3]
The cladogram below is a possible panarthropod phylogeny, taking into account the differentiation between "lower stem group arthropods" and "upper stem group arthropods", based on cladograms found in Zeng et al. 2020 and O'Flynn at al. 2022. [4] [5] (living groups are marked in bold). Not all parts of this cladogram are widely accepted, as some studies have found Isoxyida, Megacheira, Fuxianhuiida, and/or Artiopoda to lie outside of crown group Arthropoda, [6] [7] while some other studies have found a closer relationship to Chelicerata than to Mandibulata for Artiopoda. [7]
Panarthropoda | "lower stem group" "upper stem group" + crown group | |
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.
Dinocaridida is a proposed fossil taxon of basal arthropods, which flourished during the Cambrian period and survived up to Early Devonian. 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.
Cheloniellida is a taxon of extinct Paleozoic arthropods. As of 2018, 7 monotypic genera of cheloniellids had been formally described, whose fossils are found in marine strata ranging from Ordovician to Devonian in age. Cheloniellida has a controversial phylogenetic position, with previous studies associated it as either a member or relative of various fossil and extant arthropod taxa. It was later accepted as a member of Vicissicaudata within Artiopoda.
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.
The (pan)arthropod head problem is a long-standing zoological dispute concerning the segmental composition of the heads of the various arthropod groups, and how they are evolutionarily related to each other. While the dispute has historically centered on the exact make-up of the insect head, it has been widened to include other living arthropods, such as chelicerates, myriapods, and crustaceans, as well as fossil forms, such as the many arthropods known from exceptionally preserved Cambrian faunas. While the topic has classically been based on insect embryology, in recent years a great deal of developmental molecular data has become available. Dozens of more or less distinct solutions to the problem, dating back to at least 1897, have been published, including several in the 2000s.
Arthropods are invertebrates in the phylum Arthropoda. They possess an exoskeleton with a cuticle made of chitin, often mineralised with calcium carbonate, a body with differentiated (metameric) segments, and paired jointed appendages. In order to keep growing, they must go through stages of moulting, a process by which they shed their exoskeleton to reveal a new one. They form an extremely diverse group of up to ten million species.
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 the latest definitive radiodonts were known only from the Early Ordovician, at least 66 million years earlier than this taxon.
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 later surviving members include the subfamily Aegirocassisinae from the Early Ordovician of Morocco and the Early Devonian member Schinderhannes bartelsi from Germany.
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.
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.
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.
Fuxianhuiida is an extinct clade of arthropods from the Cambrian of China. All currently known species are from Cambrian Series 2 aged deposits in Yunnan Province, including the Chengjiang biota. Although historically suggested to be members of the arthropod stem group recent research has suggested that they may be closely related to mandibulates. Many specimens are known with exceptional soft tissue preservation, including preserved guts and neural tissue, which given their basal phylogenetic position makes them important in understanding the evolution of Arthropoda as a whole. They reach a size of up to 15 cm, and are interpreted as benthic predators and scavengers. The Fuxianhuiid exoskeleton is unmineralised, and the number of tergites ranges from 15 to over 40. The cephalon is covered by a head shield and contains stalked eyes connected by the anterior sclerite, antennae, a butterfly shaped hyposome and a posterior facing mouth. Fuxianhuiids possess specialized post-antennal appendages with serrated edges used for food processing. The presence of gnathobases in members of Chengjiangocardidae suggests that they were capable of durophagy. In most Fuxianhuiids, the thorax tergites narrow posteriorly, terminating in either a swimming paddle or paired flukes with a tail spine. In members of Fuxianhuiidae the thorax is divided into two sections, the anterior wide opisthothorax and the posterior narrow limbless tail-like abdomen.
Cheloniellon is a monotypic genus of cheloniellid arthropod, known only by one species, Cheloniellon calmani, discovered from the Lower Devonian Hunsrück Slate of Germany.
Kylinxia is a genus of extinct arthropod described in 2020. It was described from six specimens discovered in Yu'anshan Formation in southern China. The specimens are assigned to one species Kylinxia zhangi. Dated to 518 million years, the fossils falls under the Cambrian period. Announcing the discovery on 4 November 2020 at a press conference, Zeng Han of the Nanjing Institute of Geology and Paleontology, said that the animal "bridges the evolutionary gap from Anomalocaris to true arthropods and forms a key ‘missing link’ in the origin of arthropods," which was "predicted by Darwin’s evolutionary theory." The same day the formal description was published in Nature.
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.
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.