Radiodonta

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''.

Etymology

The name Radiodonta (Latin for ''radius'' "spoke of a wheel" and Greek for ''odoús'' "tooth") refers to the radial arrangement of tooth plates (oral cone) surrounding the mouth, although these features are suggested to be absent in some radiodont species.

Definition

The original diagnosis of order Radiodonta in 1996 is as follows:

In 2014, the clade Radiodonta was defined phylogenetically as a clade including any taxa closer to ''Anomalocaris canadensis'' than ''Paralithodes camtschaticus''. In 2019, it was redefined morphologically as animal bearing head carapace complex with central (H-) and lateral (P-) elements; outgrowths (endites) from frontal appendages bearing auxiliary spines; and reduced anterior flaps or bands of lamellae (setal blades) and strong tapering of body from anterior to posterior.

Description

Most radiodonts were significantly larger than other Cambrian fauna, with typical body lengths varying from 30 to 50 centimeters. The largest described radiodont is Ordovician '' Aegirocassis benmoulai'', which may have grown up to two meters long. A nearly complete specimen of a juvenile ''Lyrarapax unguispinus'' measured only , making it among the smallest radiodont specimens known, though adults reached a length of An isolated frontal appendage of a hurdiid with a length less than half that of the juvenile ''Lyrarapax'' is known, but it is not known whether this specimen pertains to an adult. The largest known Cambrian radiodont was ''Laminacaris'', although known from only by frontal appendages, had an estimated body length of up to based on ''Anomalocaris''. ''Anomalocaris'' and ''Amplectobelua'' are also large ones, reached and (there was an estimation that ''Houcaris saron'' (previously ''Anomalocaris saron'') reached , but specimen used for estimating the body length no longer belongs to that species); the Cambrian hurdiid ''Titanokorys'' approached it in size, with an estimated body length of approximately .

The body of a radiodont could be divided into two regions: head and trunk. The head is composed of only one body segment known as the ocular somite, covered by sclerites (head carapace complex), bore arthropodized frontal appendages, ventral mouthparts (oral cone), and stalked compound eyes. The tapering trunk is composed of multiple body segments, each associated with pairs of flaps and gill-like structures (setal blades).

Frontal appendage

The anterior structures on the head are a pair of frontal appendages which have been referred to as 'claws', 'grasping appendages', 'feeding appendages', or 'great appendages' in previous studies (the last term is discouraged since the homology between frontal appendages and the original, megacheiran great appendages is questionable.). They are sclerotized (hardened) and arthropodized (segmented), bearing ventral endites (spines) on most of their podomeres (segmental units), and the endites may bear additional rows of auxiliary spines on their anterior and posterior margins. The frontal appendage consists of two regions: the shaft ('peduncle', 'base' or 'promixal region' in some studies) and the distal articulated region (also referred to as 'claw'). A triangular region covered by soft cuticle (arthrodial membrane) may occur on the ventral side between podomeres and provide flexibility. Their pre-ocular and protocerebral origin suggest they are homologous to the primary antennae of Onychophora and the labrum of Euarthropoda (all arose from ocular somite), and not homologous with the chelicerae of Chelicerata nor the antennae or 'great appendages' of other arthropods, which are deutocerebral (arose from post-ocular somite 1). Since the morphology of the frontal appendages, especially those of the spines, always differs between species, it is one of the most important means of species identification. In fact, many radiodonts are only known from a handful of fossilized frontal appendages.

File:20191221 Radiodonta frontal appendage Anomalocarididae Amplectobeluidae.png, Frontal appendages of Anomalocarididae, Amplectobeluidae, and possibly related species
File:20191228 Radiodonta frontal appendage Tamisiocarididae Cetiocaridae.png, Frontal appendages of Tamisiocarididae
File:20191229 Radiodonta frontal appendage Hurdiidae.png, Frontal appendages of Hurdiidae

Oral cone

The mouth is on the ventral side of the head, behind the attachment point of frontal appendages and is surrounded by a ring of tooth plates, forming the mouthpart known as oral cone ('jaws' in previous studies). 3 or 4 tooth plates might be enlarged, giving the oral cone a triradial (e.g. ''Anomalocaris'') or tetraradial (e.g. Hurdiidae, ''Lyrarapax'') appearance. The inner margin of tooth plates have spikes facing towards the mouth opening. Additional rows of internal tooth plates may occur in some hurdiid genera. Detail reconstruction of some amplectobeluid oral cones are speculative, but they possibly did not present a typical radial arrangement.

Head sclerites, eyes and trunk

Three head sclerite (carapace) complex formed by a central H-element (anterior sclerite or head shield) and a pair of P-elements (lateral sclerites) cover the dorsal and laterovental surface of the animal's head. The P-elements may connect to each other as well as the H-element by a narrow anterior extension (P-element neck or 'beak'). The head sclerites are small and ovoid in Anomalocarididae and Amplectobeluidae, but often enlarged in Hurdiidae, corresponded to their distinct body shapes (streamlined in Anomalocarididae/Amplectobeluidae but often compact in Hurdiidae). The head bore two stalked compound eyes, which may have had mobility, and are located between the gaps formed by the posterior regions of the H-element and P-elements.

Contrary to the original diagnosis, the division of body segments (segmental boundaries) can be visible externally and no known member of Radiodonta (except the putative radiodont ''Cucumericrus'') is known to have pediform trunk appendages (legs). The trunk has numerous body segments (somites), tapering from anterior to posterior, with the anterior 3 or 4 segments significantly constricted into a neck region.

File:20210914 Radiodonta body flaps lobes structures variations.png, Variations of radiodont body flaps
File:Radiodonta body flaps movement.gif, The movement of radiodont flapping appendages
File:20200103 Radiodonta gnathobase‐like structures.png, Ventral view of a generalized GLS-bearing radiodont, showing gnathobase‐like structures (GLSs) associated with reduced anterior flaps

The trunk appendages were fin-like body flaps ('lateral flaps' or 'lobes' in some studies), usually one pair of ventral flaps per body segment, each slightly overlapping the one more anterior to it, but additional, non-overlapping sets of small dorsal flaps may occur in some Hurdiid species. The flaps may have numerous vein-like structures (referred to as 'strengthening rays', 'flap rays', 'tranverse rods', 'transverse lines' or 'veins'). The flaps on the neck region (referred to as 'reduced flaps', 'neck flaps', 'head flaps', 'anterior flaps' or 'differentiated flaps') are significantly reduced. In some species, jaw-like feeding appendages called gnathobase-like structures (GLSs) arose from each of the bases of their reduced neck flaps. Numerous elongated blade-like extensions (referred to as lanceolate blades or lamellae) arranged in a row, forming bands of gill-like structures known as setal blades, covered the dorsal surface of each body segment. At least in '' Aegirocassis'', each of the lanceolate blades are covered in wrinkles. The ventral flaps may be homologous to the endopod of the biramous limbs of euarthropods and lobopodous limbs (lobopods) of gilled lobopodians, and the dorsal flaps and setal blades may be homologous to the exite and gill-bearing dorsal flaps of the former taxa. The trunk may end either with a tail fan compose of 1 to 3 pairs of blades, a pair of long furcae, an elongated terminal structure, or a featureless blunt tip.

File:20220724 Hurdiidae.png, '' Stanleycaris'', '' Hurdia'', '' Aegirocassis'', '' Peytoia'' and '' Cambroraster'' are all examples of hurdiid radiodonts.
File:20210626 Anomalocaris.png, ''Anomalocaris'' is a member of the anomalocarididae family, which at one point included all radiodonts, but now only includes a few genera.
File:20210912 Amplectobeluidae.png, ''Amplectobelua'' and ''Lyrarapax'' are representatives of the amplectobeluidae which is a very inclusive family of mainly Chinese radiodonts.
File:20191228 Radiodonta frontal appendage Tamisiocarididae Cetiocaridae.png, ''"Anomalocaris" briggsi'' and ''Tamisiocaris'' are examples of the family tamisiocarididae which were exclusively suspension feeding radiodonts from the Cambrian.

Internal structures

Traces of muscles, digestive system and nervous system were described from some radiodont fossils. Pairs of well-developed muscles were connected to the ventral flaps located at the lateral cavities of each body segment. Between the lateral muscles is a sophisticated digestive system, formed by a widening of the foregut and hindgut, both connected by a narrow midgut associated with six pairs of gut divercula (digestive glands). Compared to the three-segmented brains of euarthropods and two-segmented brains of onychophorans, the brain of radiodonts is composed of only one brain segment originating from the ocular somite, the protocerebrum. The nerves of the frontal appendages and compound eyes arose from the anterior and lateral regions of the brain. Posterior to the brain was a pair of apparently unfused ventral nerve cords which ran through the animal's neck region.

Paleoecology

Physiology

Radiodonts were interpreted as nektonic or nektobenthic animals, with their morphology suggesting an active swimming lifestyle. The muscular, overlapping ventral flaps may have propelled the animal through the water, possibly by moving in a wave-like formation resembling modern rays and cuttlefish. Pairs of dorsal flaps, which make up a tail fan in some species, may have helped steering and/or stabilizing the animal during locomotion. In ''Anomalocaris'', morphology of the tail fan even suggests it could rapidly change its swimming direction efficiently. On the other hand, some hurdiids have features significantly specialized for a nektobenthic lifestyle, such as '' Cambroraster'' with its dome-like H-element similar to the carapace of a horseshoe crab. Bands of setal blades with wrinkling lanceolate blades may have increased the surface area, suggesting they were gills, providing the animal's respiratory function. Abundance of the remains of scleritzed structures such as disarticulated frontal appendages and head sclerite complexes, suggest that mass moulting events may have occurred among radiodonts, a behavior which also has been reported in some other Cambrian arthropods such as trilobites.

Diet

Radiodonts had diverse feeding strategies, which could be categorized as raptorial predators, sediment sifters, or suspension, filter feeders. For example, raptorial predators like ''Anomalocaris'' and Amplectobeluids might have been able to catch agile prey by using their raptorial frontal appendages; the latter even bore a robust endite for holding prey like a pincer. With the smaller head carapace complex and large surface of arthrodial membranes, frontal appendages of these taxa had greater flexibility. Stout frontal appendages of sediment sifters like '' Hurdia'' and '' Peytoia'' have serrated endites with mesial curvature, which could form a basket-like trap for raking through sediment and passing food items towards the well-developed oral cone. Endites of frontal appendages from suspension/filter feeders like ''Tamisiocaris'' and '' Aegirocassis'' have flexible, densely-packed auxiliary spines, which could filter out organic components such as mesozooplankton and phytoplankton down to 0.5mm. Frontal appendages of ''Caryosyntrips'', which are unusual for radiodonts in having the direction of endite-bearing surfaces opposing one another and may have been able to manipulate and crush prey in a scissor-like slicing or grasping motion.

Oral cones of radiodonts may have been used for suction and/or biting. Together with the great variety of frontal appendages in different species of radiodonts, differentiation of oral cones between species suggests preferences of different diets as well. For example, the triradial oral cone of ''Anomalocaris'' with irregular, tuberculated toothplates and a small opening may have been adapted to small and nektonic prey, ; while the rigid tetraradial oral cones of '' Peytoia'', '' Titanokorys'', '' Hurdia'', and one isolated oral cone attributed to '' Cambroraster'' with a larger opening and sometimes additional tooth plates may have been capable to consume larger food items relative to their body size and probably benthic or endobenthic prey.

Classification

Taxonomic affinities

File:20220213 Opabiniidae Opabiniids.png, The opabiniids '' Opabinia'' (top) and '' Utaurora'' (bottom)
File:20210310 Kylinxia zhangi.png, The radiodont/ opabiniid- euarthropod intermediate ''Kylinxia''
File:20220218 Erratus sperare diagrammatic reconstruction.png, A diagrammatic reconstruction of the basal deuteropod ''Erratus''
File:20221119 Mieridduryn bonniae diagrammatic reconstruction.png, '' Mieridduryn'' is a panarthropod from the Middle Ordovician that shares traits with both opabiniids and radiodonts

Most phylogenetic analyses suggest that radiodonts, alongside opabiniids ('' Opabinia'' and '' Utaurora''), are stem-group arthropods just basal to deuteropoda, a clade including upper stem (e.g. fuxianhuiids and bivalved arthropods) and crown Euarthropoda (e.g. Artiopoda, Chelicerata and Mandibulata). This interpretation is supported by numerous arthropod groundplan found on radiodonts and opabiniids, such as stalked compound eyes, digestive glands, trunk appendages forming by dorsal and ventral elements (precursor of arthropod biramous appendages). Compared to opabiniids, which possess posterior mouth opening and fused frontalmost appendages (comparable to euarthropod posterior-facing labrum/hypostome complex), radiodonts on the other hand featured euarthropod-like dorsal sclerite (H-element) and arthropodization (frontal appendages) on their head regions, alongside cuticularized gut termini. The fact that both radiodonts and opabiniids lack exoskeleton on their trunk region suggests that the origin of compound eyes and arthropodization (segmented appendages) precede arthrodization (full set of trunk exoskeleton) in the arthropod stem lineage. The constricted neck region with feeding appendicular structures of some radiodont may also shed light on the origin of the sophisticated arthropod head, which was formed by the fusion of multiple anterior body segments. Basal deuteropods that possess a mixture of radiodont/opabiniid characters like ''Kylinxia'' and ''Erratus'', may represent intermediate forms between radiodonts, opabiniids and other euarthropods.

File:20191217 Siberiida Siberion Megadictyon Jianshanopodia.png, The siberiid lobopodians '' Siberion'' (upper left), '' Megadictyon'' (bottom center) and '' Jianshanopodia'' (upper right)
File:20210730 Gilled lobopodians Pambdelurion Kerygmachela.png, The ' gilled lobopodians' ''Pambdelurion'' (left) and ''Kerygmachela'' (right)
File:20191028 Megacheirans Leanchoilia Haikoucaris Yohoia Fortiforceps.png, Megacheira or 'great appendage arthropods', a class of possible stem-chelicerate previously thought to be radiodont's close relative

Taxa just basal to the radiodont, opabiniid and euarthropod branch are ' gilled lobopodians' like ''Pambdelurion'' and ''Kerygmachela'', which occasionally united under the class Dinocaridida with opabibiids and radiodonts. They have body flaps, digestive glands, large (presumely compound) eyes and specialized frontal appendages like the former taxa, but their frontal appendages are not arthropodized nor fused, eyes sessile, gill-like structures less prominent, and certainlly bore lobopod underneath each of their flaps. Taxa even basal to 'gilled lobopodians' are siberiids like '' Megadictyon'' and '' Jianshanopodia'', a group of lobopodians bore robust frontal appendages and digestive glands, but no body flaps. Such intermediate forms between lobopodian and radiodont/euarthropod suggest that the total-group Arthropoda arose from a paraphyletic lobopodian grade, alongside the other two extant panarthropod phyla Tardigrada and Onychophora.

Previous studies may suggest radiodonts as a group other than stem-arthropods, such as a hitherto unknown phylum; cycloneuralian worms undergone convergent with arthropods (based on the cycloneuralian-like radial mouthparts); stem chelicerate euarthropods alongside megacheirans also known as great appendage arthropods (based on the similarity between radiodont frontal appendages, megacheiran great appendages and chelicerae); or ''Schinderhannes bartelsi,'' which resolved as a hurdiid radiodont in recent analyses, as a species more closely related to euarthropods than other radiodonts (based on some putative euarthropod-like features found on ''Schinderhannes''). However, neither each of them were supported by later investigations. The radial mouthparts are not cycloneuralian-exclusive and more likely present result of convergent evolution or ecdysozoan plesimorphy, since they also have been found in panarthropods such as tardigrade and some lobopodians; radiodonts lacking definitive euarthropod features such as trunk tergites and multiple head appendages, and the megacheiran great appendages were considered to be deutocerebral, which could be non-homologous to the radiodont protocerebral frontal appendages; putative euarthropod characters found on the single ''Schinderhannes'' fossil is questionable and may present other radiodont-like structures.

Interrelationships

Traditionally, all radiodont species have been placed within one family, Anomalocarididae, hence the previous common name 'anomalocaridid' and it was still occasionally used to refer the whole order even after reclassification. Since the reassignment done by Vinther ''et al.'' 2014, most of the radiodont species were reclassified within three new families: Amplectobeluidae, Tamisiocarididae (formerly Cetiocaridae), and Hurdiidae. Including Anomalocarididae, the four recent radiodont families may form the clade Anomalocarida.

The original description of the order Radiodonta included ''Anomalocaris'', ''Laggania'' (later known as '' Peytoia''), '' Hurdia'', '' Proboscicaris'', ''Amplectobelua'', ''Cucumericrus'', and ''Parapeytoia''. However, '' Proboscicaris'' is now regarded as a junior synonym of '' Hurdia'', and ''Parapeytoia'' is considered to be a Megacheiran. Due to the limited discovery, The position of ''Cucumericrus'' within Radiodonta is unclear, as it was either unselected by phylogenetic analysis or resolved in a polytomy with Radiodonta and Euarthropoda.

File:20210207 Cucumericrus decoratus trunk appendage.png, One of the poorly-known body parts (trunk appendage) of ''Cucumericrus decoratus''; this species may not represent a true radiodont.
File:20191228 Radiodonta frontal appendage Anomalocaris briggsi.png, Frontal appendage of ''"Anomalocaris" briggsi''. Despite the suggested generic name, this radiodont is unlikely to be a species of genus ''Anomalocaris'' based on phylogenetic analysis.

The first in-depth phylogenetic analysis of Radiodonta was conducted by Vinther ''et al''. in 2014, followed by a handful of subsequest studies with more or less modified results. In most analysis, ''Caryosyntrips'' is the basal-most genus, but resolved in a polytomy with other radiodonts and Euarthropoda (alongside ''Cucumericrus'' if included). With the exclusion of ''Caryosyntrips'' and ''Cucumericrus'', the monophyly of Radiodonta is widely supported, with a few results suggest possible paraphyly (either the Anomalocarididae+Amplectobeluidae clade or Hurdiidae sister to Euarthropoda). Putative synapomorphies of monophyletic Radiodonta including tripartite head sclerite complex and differentiated neck region. The genus ''Anomalocaris'' in a broader sense always found to be polyphyletic, usually with ''"Anomalocaris" kunmingensis'' and ''"Anomalocaris" briggsi'' resolved as a member of Amplectobeluidae and Tamisiocarididae respectively. Interrelationship of Amplectobeluidae is uncertain, as the amplectobeluid affinities of ''Lyrarapax'' and ''Ramskoeldia'' were occasionally questioned. Monophyly of the speciose family Hurdiidae was recovered by most analysis and well-supported by several synapomorphies (e.g. distal articulated region of frontal appendage with proximal 5 podomeres bearing subequal endites), with Tamisiocarididae often suggested to be its sister group.

* Radiodonta
** ?''Cucumericrus'' (putative radiodont)
** ''Caryosyntrips''
** Anomalocarida
*** ''Paranomalocaris'' (placed within Anomalocarididae by some studies.)
*** '' Laminacaris'' (placed within Amplectobeluidae by some studies.)
*** '' Houcaris'' (either placed within Anomalocarididae, Amplectobeluidae or Tamisiocarididae.)
*** '' Innovatiocaris''
*** Anomalocarididae
**** ''Anomalocaris'' (in a broader sense, some species may placed within the other families.)
**** '' Lenisicaris''
*** Amplectobeluidae
**** ''Lyrarapax'' (position questioned by some studies.)
**** ''Amplectobelua''
**** ''Ramskoeldia'' (position questioned by some studies.)
**** ''"Anomalocaris" kunmingensis'' (non-''Anomalocaris'' species await to be renamed. Labeled as ''Amplectobelua kunmingensis'' by Vinther et al. 2014.)
*** Tamisiocarididae
**** ''Tamisiocaris''
**** ''"Anomalocaris" briggsi'' (non-''Anomalocaris'' species await to be renamed.)
*** Hurdiidae
****'' Peytoia''
**** ''Schinderhannes''
**** '' Hurdia''
**** '' Aegirocassis''
**** '' Stanleycaris''
**** ''Pahvantia''
**** '' Ursulinacaris''
**** '' Cambroraster''
**** ?'' Zhenghecaris'' (putative radiodont)
**** '' Cordaticaris''
**** '' Buccaspinea''
**** '' Titanokorys''

History

File:Anomalocaris canadensis grasping claw, Burgess Shale.jpg, Frontal appendage of ''Anomalocaris canadensis''
File:Peytoia nathorsti Laggania cambria oral cone, Burgess Shale.jpg, Oral cone of '' Peytoia nathorsti''
File:Hurdia victoria USNM PAL 57718.jpg, H-element of '' Hurdia victoria''
File:USNM PAL 57490.jpg, Paired frontal appendages from an unnamed hurdiid radiodont

The history of radiodonts is complex. Incomplete specimens pertaining to different body parts of the same species had historically been interpreted as belonging to different species and even different phyla. Prior to their recognition as a group, radiodont specimens had been assigned to five different phyla: Porifera, Cnidaria, Echinodermata, Annelida, and Arthropoda.

The first known radiodont specimens were collected from the trilobite beds of Mount Stephen by Richard G. McConnell of the Geological Survey of Canada in 1886 or 1888. These specimens were named ''Anomalocaris canadensis'' in 1892 by GSC paleontologist Joseph Whiteaves. Whiteaves interpreted the specimens, now known to be isolated frontal appendages, as the abdomen of a phyllocarid crustacean. Additional radiodont specimens were described in 1911 by Charles Walcott.Walcott, C. D. 1911a
Middle Cambrian holothurians and medusae
Cambrian geoogy and paleontology II. Smithsonian Miscellaneous Collections, 57: 41-68. He interpreted an isolated oral cone, which he named '' Peytoia nathorsti'', as a jellyfish, and a poorly-preserved but relatively complete specimen, which he named ''Laggania cambria'', as a holothurian. In 1912 Walcott named '' Hurdia victoria'' and ''H. triangulata'' based on isolated H-elements, which he interpreted as the carapaces of crustaceans.WALCOTT, C. D. 1912
Middle Cambrian Branchiopoda, Malacostraca, Trilobita and Merostomata
Smithsonian Miscellaneous Collections, 57: 145–228. Isolated frontal appendages of ''Peytoia'' and ''Hurdia'', collectively known as "Appendage F" in Briggs 1979, were all identified as those of '' Sidneyia'' at that time. A ''Hurdia'' P-element was named ''Proboscicaris'' in 1962, and interpreted as the carapace of a bivalved arthropod.

The Geological Survey of Canada initiated a revision of Burgess Shale fossils in 1966, overseen by Cambridge University paleontologist Harry B. Whittington. This revision would ultimately lead to the discovery of the complete radiodont body plan. In 1978, Simon Conway Morris recognized that the mouthparts of ''Laggania'' were ''Peytoia''-like, but he interpreted this as evidence that it was a composite fossil made up of a ''Peytoia'' jellyfish and a sponge. In 1979, Derek Briggs recognized that the fossils of ''Anomalocaris'' were appendages, not abdomens, but interpreted them as walking legs alongside "Appendage F". It was not until 1985 that the true nature of the fossils of ''Anomalocaris'', ''Laggania'', and ''Peytoia'' was recognized, and they were all assigned to a single genus, ''Anomalocaris''. Subsequently, it was recognized that ''Anomalocaris'' was a distinct form from the other two, resulting in a split into two genera, the latter of which was variously named ''Laggania'' and ''Peytoia'' until it was determined that both represent the same species and ''Peytoia'' had priority. It was later recognized that some of the fossils assigned to these taxa belonged to another form, which was recognized as bearing a carapace made up of ''Hurdia'' and ''Proboscicaris'' elements. Finally, in 2009, these specimens were redescribed as ''Hurdia''. Even after these recognitions, partial misidentifications (e.g. oral cone and frontal appendages of ''Peytoia'' had been assigned to ''Anomalocaris'' and ''Hurdia'', respectively) had been revealed by subsequent studies as well.

The taxon Radiodonta itself was coined in 1996 by Desmond Collins, after it was established that ''Anomalocaris'' and its kin represented a distinctive lineage with arthropod affinities rather than a hitherto unknown phylum. Collins also established the class Dinocarida to contain the order Radiodonta as well as the Opabiniidae, which he recognized as distinct due to its lacking the distinctive oral cone structure of radiodonts. Radiodonta was first given a phylogenetic definition in 2014. Radiodonta was originally viewed as containing a single family, Anomalocarididae, but it was divided into four families in 2014: Amplectobeluidae, Anomalocarididae, Cetiocaridae, and Hurdiidae. The name Cetiocaridae did not conform to the International Code of Zoological Nomenclature and so was renamed Tamisiocarididae in 2019.

Until the 2010s, radiodonts were typically considered to be uniformly large apex predators, but discoveries of new species over the course of that decade led to a considerable increase in the known ecological and morphological diversity of the group.

References

{{Taxonbar, from=Q3291852

*
Cambrian Series 2 first appearances
Dinocarida
Early Devonian extinctions
Prehistoric arthropod orders