| Radiodonta | |
|---|---|
 | |
| Left to right, top to bottom:Amplectobelua symbrachiata,Anomalocaris canadensis,Aegirocassis benmoulai,Peytoia nathorsti,Lyrarapax unguispinus,Cambroraster falcatus, andHurdia victoria | |
| Scientific classification | |
| Kingdom: | |
| Phylum: | |
| Class: | |
| Order: | †Radiodonta Collins, 1996 |
| Families | |
| Synonyms | |
Anomalocarida | |
Radiodonta is an extinctorder ofstem-grouparthropods that was successful worldwide during theCambrian period. Radiodonts are distinguished by their distinctive frontal appendages, which are morphologically diverse and were used for a variety of functions. Radiodonts were among the earliest large predators, but they also included sediment sifters and filter feeders.[1] Some of the most famous species of radiodonts are theCambrian taxaAnomalocaris canadensis,Hurdia victoria,Peytoia nathorsti,Titanokorys gainesi,Cambroraster falcatus andAmplectobelua symbrachiata. The later surviving members include the subfamilyAegirocassisinae from theEarly Ordovician ofMorocco and theEarly Devonian memberSchinderhannes bartelsi from Germany.
The name Radiodonta (Latin forradius "spoke of a wheel" and Greek forodoús "tooth") refers to the radial arrangement of tooth plates (oral cone) surrounding the mouth,[2] although these features are suggested to be absent in some radiodont species.[3][4]
The original diagnosis of order Radiodonta in 1996 is as follows:[2]
Radiodontids are bilaterally symmetrical, elongate arthropods with a nonmineralized cuticle typically most robust in the jaws and claws. The body is subdivided into twotagmata, much like theprosoma andopisthosoma ofchelicerate arthropods. Typically, the front part shows no external segmentation, bears one pair of preoral claws, a pair of prominent eyes, and ventral jaws with radiating teeth. Some forms have additional rows of teeth and three or four postoralgnathobasic limb pairs. The trunk ismetameric, typically with about 13 segments laterally developing imbricating lobes for swimming and gills for respiration, and may end in a prominent three-part tail. Some forms have gnathobasic trunk limbs.
In 2014, the clade Radiodonta was definedphylogenetically as a clade including any taxa closer toAnomalocaris canadensis thanParalithodes camtschaticus.[5] In 2019, it was redefinedmorphologically 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.[6]
Members of Radiodonta are known as radiodonts,[6][4][7] radiodontans,[3][8] radiodontids,[2] anomalocarids,[5] or anomalocaridids,[9][10][11] although the last two originally refer to the familyAnomalocarididae, which previously included all species of this order but is now restricted to only a few species.[5]
Most radiodonts were significantly larger than the otherCambrian fauna, with typical body lengths of large taxa varying from 30 to 50 cm (12 to 20 in).[12] The largest described radiodont is theEarly Ordovician speciesAegirocassis benmoulai, which may have grown up to 2 m (6.6 ft) long.[11][7] A nearly complete specimen of a juvenileLyrarapax unguispinus measured only 18 mm (0.71 in), making it among the smallest radiodont specimens known, though adults reached a length of 8.3 cm (3.3 in)[7][13] An isolated frontal appendage of a hurdiid with a length less than half that of the juvenileLyrarapax is known, but it is not known whether this specimen pertains to an adult.[14] The largest known Cambrian radiodont wasAmplectobelua, reaching lengths of up to 90 cm (35 in) based on an incomplete specimen.[15]Anomalocaris canadensis was also relatively large, estimated up to 34.2–37.8 cm (13.5–14.9 in) long,[7] and the Cambrian hurdiidTitanokorys approached around 50 cm (20 in) long.[16]
The body of a radiodont could be divided into two regions: head and trunk. The head is composed of only one body segment[17] known as the ocular somite, covered by sclerites (head carapace complex), bore arthropodized frontal appendages, ventral mouthparts (oral cone), andstalkedcompound eyes. The tapering trunk is composed of multiple body segments, each associated with pairs of flaps and gill-like structures (setal blades).[6]
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, morphologically distinctmegacheirangreat appendages is questionable.[17][18]). They aresclerotized (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.[19][6] The frontal appendage consists of two regions: the shaft ('peduncle',[7] 'base'[20] or 'promixal region'[7] in some studies) and the distal articulated region[19] (also referred to as 'claw'[20]). A triangular region covered by soft cuticle (arthrodial membrane) may occur on the ventral side between podomeres and provide flexibility.[21][1] Their purported pre-ocular and protocerebral origin suggest they are homologous to the primary antennae ofOnychophora and thelabrum ofEuarthropoda (all arose from ocularsomite),[17][10] while subsequent studies also suggest a deutocerebral origin and homologous with thechelicerae ofChelicerata and theantennae or 'great appendages' of other arthropods (all arose from post-ocular somite 1).[22] 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.[19] In fact, many radiodonts are only known from a handful of fossilized frontal appendages.[21][19]
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[2]). 3 or 4 tooth plates might be enlarged, giving the oral cone a triradial (e.g.Anomalocaris,Echidnacaris) or tetraradial (e.g.Hurdiidae,Lyrarapax) appearance.[23][13][24] The inner margin of tooth plates have spikes facing towards the mouth opening. Additional rows of internal tooth plates may occur in somehurdiid genera.[9][6] Detail reconstruction of someamplectobeluid oral cones are speculative, but they possibly did not present a typical radial arrangement.[3][4]
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.[6] The P-elements may connect to each other as well as the H-element by a narrow anterior extension (P-element neck or 'beak').[9][6] The head sclerites are small and ovoid inAnomalocarididae andAmplectobeluidae,[3][6] but often enlarged inHurdiidae, corresponded to their distinct body shapes (streamlined in Anomalocarididae/Amplectobeluidae but often compact in Hurdiidae).[6] The head bore twostalkedcompound eyes, which may have had mobility,[25] and are located between the gaps formed by the posterior regions of the H-element and P-elements.[9][6] The compound eyes ofEchidnacaris are exceptionally unstalked.[14] Some species possess an additional median eye behind the H-element.[22]
Contrary to the original diagnosis, the division of body segments (segmental boundaries) can be visible externally[11][8][6] and no known member of Radiodonta (except the putative radiodontCucumericrus[11][26]) is known to have pediform trunk appendages (legs).[27] The trunk has numerous body segments (somites), tapering from anterior to posterior, with the anterior three or four segments significantly constricted into a neck region.[6]
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 someHurdiid species.[11] The flaps may have numerous vein-like structures (referred to as 'strengthening rays',[8] 'flap rays',[6] 'tranverse rods',[11] 'transverse lines'[28] or 'veins'[29]). The flaps on the neck region (referred to as 'reduced flaps',[3] 'neck flaps',[8] 'head flaps',[27] 'anterior flaps'[30] or 'differentiated flaps'[18]) 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.[3][4] Numerous elongated blade-like extensions (referred to as lanceolate blades or lamellae[6]) arranged in a row, forming bands ofgill-like structures known as setal blades, covered the dorsal surface of each body segment.[11] At least inAegirocassis, each of the lanceolate blades are covered in wrinkles.[11] The ventral flaps may be homologous to the endopod of the biramous limbs of euarthropods and lobopodous limbs (lobopods) of gilledlobopodians, and the dorsal flaps and setal blades may be homologous to the exite and gill-bearing dorsal flaps of the former taxa.[31][11] The trunk may end either with a tail fan compose of 1 to 3 pairs of blades,[29][27][6] a pair of long furcae,[29][13][6] an elongated terminal structure,[27] or a featureless blunt tip.[11]
Traces ofmuscles,digestive system andnervous 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.[27][10] 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).[27][8][32]
The brain of radiodonts was simpler than the three-segmented (compose of pro-, deuto- and tritocerebrum) brains ofeuarthropods, but further interpretations differ between studies. Based on Cong et al. 2014, the brain 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.[10][17] Based on Moysiuk & Caron 2022, the frontal appendage nerves arose from the ventral deutocerebrum, the second brain segment. The previous "frontal appendage nerves" actually represent median eye nerve.[22] In both interpretations, posterior to the brain was a pair of apparently unfusedventral nerve cords which ran through the animal's neck region.[10][22]
Radiodonts were interpreted asnektonic ornektobenthic 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 modernrays andcuttlefish.[33][34] Pairs of dorsal flaps, which make up a tail fan in some species, may have helped steering and/or stabilizing the animal during locomotion.[11][35] InAnomalocaris, morphology of the tail fan even suggests it could rapidly change its swimming direction efficiently.[36] On the other hand, some hurdiids have features significantly specialized for a nektobenthic lifestyle, such asCambroraster with its dome-like H-element similar to thecarapace of ahorseshoe crab.[6] Bands of setal blades with wrinkling lanceolate blades may have increased the surface area, suggesting they weregills, providing the animal'srespiratory function.[27][11] Abundance of the remains of scleritzed structures such as disarticulated frontal appendages and head sclerite complexes, suggest that massmoulting events may have occurred among radiodonts,[11][6] a behavior which also has been reported in some otherCambrian arthropods such astrilobites.[37]
Radiodonts had diverse feeding strategies, which could be categorized asraptorialpredators, sediment sifters, or suspension,filter feeders.[7][38][1][39][40] For example, raptorial predators likeAnomalocaris andAmplectobeluids 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 apincer.[26][21][3][1] With the smaller head carapace complex and large surface of arthrodial membranes, frontal appendages of these taxa had greater flexibility.[13] Stout frontal appendages of sediment sifters likeHurdia andPeytoia 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.[6][1] Endites of frontal appendages from suspension/filter feeders likeTamisiocaris andAegirocassis have flexible, densely-packed auxiliary spines, which could filter out organic components such asmesozooplankton andphytoplankton down to 0.5mm.[5][11] Frontal appendages ofCaryosyntrips, 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.[21][41]
Oral cones of radiodonts may have been used for suction and/or biting.[23][38][6] 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.[38][1] For example, the triradial oral cone ofAnomalocaris with irregular, tuberculated toothplates and a small opening may have been adapted to small and nektonic prey,[23][1] while the rigid tetraradial oral cones ofPeytoia,Titanokorys,Hurdia, and one isolated oral cone attributed toCambroraster 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.[23][38][6]
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| Summarized phylogeny between Radiodonta and other Ecdysozoan taxa[42] |
![The presumed radiodont/opabiniid-euarthropod intermediate Kylinxia, shares many of the characteristics found in both dinocaridids and euarthropods — later studies have considered this relationship far less direct[43]](/mt/?noimg=&dark=on&url=http%3a%2f%2fen.wikipedia.org%2fwiki%2fFile%3a20210310_Kylinxia_zhangi.png)
Mostphylogenetic analyses suggest that radiodonts, alongsideopabiniids (Opabinia andUtaurora[44]), arestem-grouparthropods just basal todeuteropoda,[42] a clade including upper stem (e.g.fuxianhuiids and bivalved arthropods) and crownEuarthropoda (e.g.Artiopoda,Chelicerata andMandibulata).[9][45][46][47][48][49][5][10][11][7][6][30][18][39][40][50][51][44] This interpretation is supported by numerous arthropod groundplan found on radiodonts and opabiniids, such asstalkedcompound eyes,[25] digestive glands,[32] trunk appendages forming by dorsal and ventral elements (precursor of arthropod biramous appendages).[11][51] Compared to opabiniids, which possess posterior mouth opening and fused frontalmost appendages (comparable to euarthropod posterior-facinglabrum/hypostome complex),[17][44] radiodonts on the other hand featured euarthropod-like dorsal sclerite (H-element) and arthropodization (frontal appendages) on their head regions,[52][17][44] alongsidecuticularized gut termini.[27] 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.[42][53][54] 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.[3][17] Basal deuteropods that possess a mixture of radiodont/opabiniid characters likeKylinxia andErratus, may represent intermediate forms between radiodonts, opabiniids and other euarthropods.[18][51]
Taxa just basal to the radiodont, opabiniid and euarthropod branch are 'gilled lobopodians' likePambdelurion andKerygmachela, which occasionally united under the classDinocaridida with opabibiids and radiodonts.[55][47] 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 certainly bore lobopod underneath each of their flaps.[56][11][57][44] Taxa even basal to 'gilled lobopodians' are siberiids likeMegadictyon andJianshanopodia,[42] a group oflobopodians that 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 aparaphyletic lobopodian grade, alongside the other two extantpanarthropod phylaTardigrada andOnychophora.[58][42][17][59][53][54]
Previous studies may suggest radiodonts as a group other than stem-arthropods, such as a hitherto unknownphylum;[33]cycloneuralian worms undergoneconvergent with arthropods (based on the cycloneuralian-like radial mouthparts);[60][55] stemchelicerate euarthropods alongsidemegacheirans also known as great appendage arthropods (based on the similarity between radiodont frontal appendages, megacheiran great appendages andchelicerae);[61] orSchinderhannes bartelsi, which resolved as ahurdiid radiodont in recent analyses,[42][5][11][7][6][39][40] as a species more closely related to euarthropods than other radiodonts (based on some putative euarthropod-like features found onSchinderhannes).[35] 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 orecdysozoanplesimorphy, since they also have been found inpanarthropods such astardigrade and somelobopodians;[62] radiodonts lacking definitive euarthropod features such as trunktergites and multiple head appendages,[42] and the megacheiran great appendages were considered to be deutocerebral,[63][64] which could be non-homologous to the radiodont protocerebral frontal appendages;[10][17] putative euarthropod characters found on the singleSchinderhannes fossil is questionable and may present other radiodont-like structures.[42]
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| Phylogeny of Radiodonta after Moysiuk & Caron 2021[39] |
Traditionally, all radiodont species have been placed within one family,Anomalocarididae,[2] hence the previous common name 'anomalocaridid'[26][9] and it was still occasionally used to refer the whole order even after reclassification.[10][11] Since the reassignment done by Vintheret al. 2014, most of the radiodont species were reclassified within three new families:Amplectobeluidae,Tamisiocarididae[7][6] (formerlyCetiocaridae[5]), andHurdiidae.[5][11][7][6] Including Anomalocarididae, the four recent radiodont families may form the clade Anomalocarida.[5]
The original description of the order Radiodonta includedAnomalocaris,Laggania (later known asPeytoia),Hurdia,Proboscicaris,Amplectobelua,Cucumericrus, andParapeytoia.[2] However,Proboscicaris is now regarded as a junior synonym ofHurdia, andParapeytoia is considered to be aMegacheiran.[9][27][11] Due to the limited discovery, The position ofCucumericrus within Radiodonta is unclear, as it was either unselected byphylogenetic analysis[5][6][7][39][40] or resolved in a polytomy with Radiodonta andEuarthropoda.[11][13]
The first in-depth phylogenetic analysis of Radiodonta was conducted by Vintheret al. in 2014,[5] followed by a handful of subsequest studies with more or less modified results.[10][11][7][13][6][39][40][44] In most analysis,Caryosyntrips is the basal-most genus, but either resolved in a polytomy with other radiodonts and Euarthropoda (alongsideCucumericrus if included[11][13]) or outside of Radiodonta, casting doubt on its radiodont affinity.[65] With the exclusion of questionableCaryosyntrips andCucumericrus, themonophyly of Radiodonta is widely supported,[5][10][11][7][13][6][39][40] with a few results suggest possibleparaphyly (either the Anomalocarididae+Amplectobeluidae clade or Hurdiidaesister to Euarthropoda).[30][44] Putativesynapomorphies of monophyletic Radiodonta including tripartite head sclerite complex and differentiated neck region.[6] The genusAnomalocaris in a broader sense always found to bepolyphyletic, usually with"Anomalocaris" kunmingensis and"Anomalocaris" briggsi resolved as a member ofAmplectobeluidae andTamisiocarididae respectively.[5][10][11][7][6][39][40] Interrelationship of Amplectobeluidae is uncertain, as the amplectobeluid affinities ofLyrarapax andRamskoeldia were occasionally questioned.[4][6][40] Monophyly of the speciose familyHurdiidae 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[19][6]). Tamisiocarididae was often suggested to be sister group of Hurdiidae in the 2010s,[5][11][7][6] but this position became questionable in subsequent studies.[22][24]
| Species | Original description | Year named | Family | Age | Location | Frontal appendage | Head sclerite complex |
|---|---|---|---|---|---|---|---|
| Cucumericrus decoratus | Hou, Bergström, & Ahlberg | 1995[26] | (unassigned) | Cambrian Stage 3 |  China | Unknown | Unknown |
| Caryosyntrips serratus | Daley & Budd | 2010[21] | (unassigned) | Wuliuan–Drumian |  Canada United States |  | Unknown |
| Caryosyntrips camurus | Pates & Daley | 2017[41] | (unassigned) | Wuliuan | Canada United States |  | Incomplete[80] |
| Caryosyntrips durus | Pates & Daley | 2017[41] | (unassigned) | Drumian | United States |  | Unknown |
| Paranomalocaris multisegmentalis | Wang, Huang, & Hu | 2013[67] | Anomalocarididae? | Cambrian Stage 4 | China |  | Unknown |
| Paranomalocaris simplex | Jiao, Pates, Lerosey-Aubril, Ortega-Hernandez, Yang, Lan, Zhang | 2021[68] | Anomalocarididae? | Cambrian Stage 4 | China |  | Unknown |
| Laminacaris chimera | Guo, Pates, Cong, Daley, Edgecombe, Chen, & Hou | 2018[69] | (controversial) | Cambrian Stage 3 | China |  | Unknown |
| Innovatiocaris maotianshanensis | Zeng, Zhao, Zhu | 2022[71] | (unassigned) | Cambrian Stage 3 | China |  | P-element unknown[71] |
| Innovatiocaris?multispiniformis | Zeng, Zhao, Zhu | 2022[71] | (unassigned) | Cambrian Stage 3 | China |  | Unknown |
| Anomalocaris canadensis | Whiteaves | 1892[81] | Anomalocarididae | Wuliuan | United States |  |  |
| Lenisicaris pennsylvanica (formerlyAnomalocaris pennsylvanica)[20] | Resser | 1929 | Anomalocarididae | Cambrian Stage 3 | United States |  | Unknown |
| Lenisicaris lupata | Wu, Ma, Lin, Sun, Zhang, & Fu | 2021[20] | Anomalocarididae | Cambrian Stage 3 | China |  | Unknown |
| Anomalocaris daleyae | Paterson, García-Bellidob & Edgecombe | 2023 | Anomalocarididae | Cambrian Stage 4 |  Australia |  | Unknown |
| Houcaris magnabasis (formerlyAnomalocaris magnabasis)[70] | Pates, Daley, Edgecombe, Cong & Lieberman | 2019 | (controversial) | Cambrian Stage 4 | United States |  | Unknown |
| Houcaris saron (formerlyAnomalocaris saron)[70] | Hou, Bergström, & Ahlberg | 1995 | (controversial) | Cambrian Stage 3 | China |  | Unknown |
| Echidnacaris briggsi[24] | Nedin | 1995 | Tamisiocarididae | Cambrian Stage 4 | Australia | | Possible H-element and unique lateral sclerites associated with compound eyes[14][24] |
| Ramskoeldia platyacantha | Cong, Edgecombe, Daley, Guo, Pates, & Hou | 2018[4] | Amplectobeluidae | Cambrian Stage 3 | China |  | Incomplete[4] |
| Ramskoeldia consimilis | Cong, Edgecombe, Daley, Guo, Pates, & Hou | 2018[4] | Amplectobeluidae | Cambrian Stage 3 | China |  | Incomplete[4] |
| Lyrarapax unguispinus | Cong, Ma, Hou, Edgecombe, & Strausfield | 2014[10] | Amplectobeluidae | Cambrian Stage 3 | China |  | P-element neck unknown |
| Lyrarapax trilobus | Cong, Daley, Edgecombe, Hou, & Chen | 2016[8] | Amplectobeluidae | Cambrian Stage 3 | China |  | P-element unknown |
| Amplectobelua symbrachiata | Hou, Bergström, & Ahlberg | 1995[26] | Amplectobeluidae | Cambrian Stage 3 | China |  |  |
| Amplectobelua stephenensis | Daley & Budd | 2010[21] | Amplectobeluidae | Wuliuan | United States |  | Unknown |
| Guanshancaris kunmingensis | Zhanget al. | 2023[73] | Amplectobeluidae | Cambrian Stage 4 | China |  | Unknown |
| Tamisiocaris borealis | Daley & Peel | 2010 | Tamisiocarididae | Cambrian Stage 3 |  Greenland |  | Incomplete[5] |
| Ursulinacaris grallae | Pates, Daley & Butterfield | 2019 | Hurdiidae | Wuliuan | Canada |  | Unknown |
| Schinderhannes bartelsi | Kühl, Briggs, & Rust | 2009[35] | Hurdiidae | Emsian |  Germany | Incomplete[6] | Incomplete[6] |
| Stanleycaris hirpex | Pates, Daley, & Ortega-Hernández | 2018[78] | Hurdiidae | Wuliuan | Canada |  | P-element is unknown, possibly absent[22] |
| Peytoia nathorsti | Walcott | 1911[82] | Hurdiidae | Wuliuan–Drumian | Canada United States |  | Incomplete[6] |
| Peytoia infercambriensis (formerlyCassubia infercambriensis)[83] | Lendzion | 1975 | Hurdiidae | Cambrian Stage 3 |  Poland |  | Unknown |
| Aegirocassis benmoulai | Van Roy, Daley, & Briggs | 2015[11] | Hurdiidae (Aegirocassisinae) | Tremadocian |  Morocco |  |  |
| Hurdia victoria | Walcott | 1912[84] | Hurdiidae | Wuliuan–Drumian | Canada Czechia |  |  |
| Hurdia triangulata | Walcott | 1912[84] | Hurdiidae | Wuliuan | Canada | |  |
| Cambroraster falcatus | Moysiuk & Caron | 2019[6] | Hurdiidae | Wuliuan | Canada |  |  |
| Pahvantia hastata | Robison & Richards | 1981 | Hurdiidae | Drumian | United States |  |  |
| Cordaticaris striatus | Sun, Zeng, & Zhao | 2020[79] | Hurdiidae | Drumian | China | Incomplete[79] |  |
| Zhenghecaris shankouensis | Vanner, Chen, Huang, Charbonnier, & Wang | 2006 | Hurdiidae | Cambrian Stage 3 | China | Unknown[66] |  |
| Buccaspinea cooperi | Pates, Lerosey-Aubril, Daley, Kier, Bonino & Ortega-Hernández | 2021[80] | Hurdiidae | Drumian | United States |  | Unknown |
| Titanokorys gainesi | Caron & Moysiuk | 2021[40] | Hurdiidae | Wuliuan | Canada |  |  |
| Pseudoangustidontus duplospineus | Van Roy & Tetlie | 2006 | Hurdiidae (Aegirocassisinae) | Tremadocian | Morocco |  | Unknown |
| Pseudoangustidontus izdigua | Potin, Gueriau & Daley | 2023 | Hurdiidae (Aegirocassisinae) | Tremadocian | Morocco |  | Incomplete[74] |
| Shucaris ankylosskelos | Wuet al. | 2024 | (controversial) | Cambrian Stage 2–Cambrian Stage 3 | China |  | Incomplete[72] |
| Stanleycaris qingjiangensis | Wuet al. | 2024 | Hurdiidae | Cambrian stage 3 | China |  | Incomplete[85] |
![Paired frontal appendages from an unnamed hurdiid radiodont[39]](/mt/?noimg=&dark=on&url=http%3a%2f%2fen.wikipedia.org%2fwiki%2fFile%3aUSNM_PAL_57490.jpg)
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.[2][9] Prior to their recognition as a group, radiodont specimens had been assigned to five different phyla:Porifera,Cnidaria,Echinodermata,Annelida, andArthropoda.[2]
The first known radiodont specimens were collected from thetrilobite beds ofMount Stephen by Richard G. McConnell of theGeological Survey of Canada in 1886[2] or 1888.[81] These specimens were namedAnomalocaris canadensis in 1892 by GSC paleontologistJoseph Whiteaves.[81] Whiteaves interpreted the specimens, now known to be isolated frontal appendages, as the abdomen of aphyllocaridcrustacean.[81] Additional radiodont specimens were described in 1911 byCharles Walcott.[82] He interpreted an isolated oral cone, which he namedPeytoia nathorsti, as a jellyfish, and a poorly-preserved but relatively complete specimen, which he namedLaggania cambria, as a holothurian.[82] In 1912 Walcott namedHurdia victoria andH. triangulata based on isolated H-elements, which he interpreted as the carapaces of crustaceans.[84] Isolated frontal appendages ofPeytoia andHurdia, collectively known as "Appendage F" in Briggs 1979, were all identified as those ofSidneyia at that time.[82] AHurdia P-element was namedProboscicaris in 1962, and interpreted as the carapace of a bivalved arthropod.[86]
The Geological Survey of Canada initiated a revision of Burgess Shale fossils in 1966, overseen byCambridge University paleontologistHarry B. Whittington.[2] This revision would ultimately lead to the discovery of the complete radiodont body plan. In 1978,Simon Conway Morris recognized that the mouthparts ofLaggania werePeytoia-like, but he interpreted this as evidence that it was a composite fossil made up of aPeytoia jellyfish and a sponge.[87] In 1979,Derek Briggs recognized that the fossils ofAnomalocaris were appendages, not abdomens, but interpreted them as walking legs alongside "Appendage F".[88] It was not until 1985 that the true nature of the fossils ofAnomalocaris,Laggania, andPeytoia was recognized, and they were all assigned to a single genus,Anomalocaris.[33] Subsequently, it was recognized thatAnomalocaris was a distinct form from the other two, resulting in a split into two genera, the latter of which was variously namedLaggania andPeytoia until it was determined that both represent the same species andPeytoia had priority.[23] 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 ofHurdia andProboscicaris elements. Finally, in 2009, these specimens were redescribed asHurdia.[9] Even after these recognitions, partial misidentifications (e.g. oral cone and frontal appendages ofPeytoia had been assigned toAnomalocaris[2] andHurdia,[9] respectively) had been revealed by subsequent studies as well.[23][89]
The taxon Radiodonta itself was coined in 1996 by Desmond Collins, after it was established thatAnomalocaris and its kin represented a distinctive lineage with arthropod affinities rather than a hitherto unknown phylum.[2] Collins also established the classDinocarida to contain the order Radiodonta as well as theOpabiniidae, which he recognized as distinct due to its lacking the distinctive oral cone structure of radiodonts.[2] Radiodonta was first given a phylogenetic definition in 2014.[5] Radiodonta was originally viewed as containing a single family,Anomalocarididae, but it was divided into four families in 2014:Amplectobeluidae, Anomalocarididae,Cetiocaridae, andHurdiidae.[5] The name Cetiocaridae did not conform to the International Code of Zoological Nomenclature and so was renamedTamisiocarididae in 2019.[90]
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.[5][11][7][6][91][80][39][40]
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