Dinocaridida Temporal range:
Assembly of dinocaridids (Anomalocaris, Opabinia, Pambdelurion and Kerygmachela)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	†Dinocaridida Collins, 1996
Subgroups
Parvibellus? Pambdelurion Omnidens? Kerygmachelidae Opabiniidae Radiodonta

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]

Anatomy

Eyes (deep blue and black), brain (light blue) and digestive system (yellow) of Radiodonta (left), Kerygmachela (Center) and Opabinia (Right)

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 though 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 an euarthropod (3-segmented), it is thought to be comprised either 1 (only protocerebrum^[10][16]) or 2 cerebral ganglions (protocerebrum and deutocerebrum).^[12]

Classification

Although some authors may rather suggest different taxonomic affinities (e.g. as cycloneuralian 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.

Distribution

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]

Notes

References