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DinocarididaGreek, "Terrible crabs" – sometimes informally spelt Dinocarida, but the second 'id' is linguistically correct – see is a proposed
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
taxon of basal
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s that flourished in the
Cambrian The Cambrian Period ( ; sometimes symbolized C with bar, Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million ...
period with occasional
Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start ...
Van Roy, P.; Briggs, D. E. G. (2011). "A giant Ordovician anomalocaridid". Nature 473 (7348): 510–513. doi:10.1038/nature09920. edit and
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
records. Characterized by a pair of frontal appendages and series of body flaps, the name of Dinocaridids comes from
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
, "deinos" and "caris" ("terrible crab"), referring to the suggested role of some of these members as the largest marine
predators Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
of their time. Dinocaridids are occasionally referred to as the 'AOPK group' by some literatures, as the group compose of
Radiodonta Radiodonta is an extinct Order (biology), 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 ...
(''
Anomalocaris ''Anomalocaris'' ("unlike other shrimp", or "abnormal shrimp") is an extinct genus of Radiodonta, radiodont, an Order (biology), order of early-diverging stem-group arthropods. The first fossils of ''Anomalocaris'' were discovered in the ''Ogygop ...
'' and relatives),
Opabiniidae Opabiniidae is an extinct family of bizarre marine stem-arthropods. Its type and best-known genus is '' Opabinia''. It also contains '' Utaurora''. Opabiniids closely resemble radiodonts, but their frontal appendages were basally fused into a ...
(''
Opabinia ''Opabinia regalis'' is an extinct, stem group arthropod found in the Middle Cambrian Burgess Shale Lagerstätte (505 million years ago) of British Columbia. ''Opabinia'' was a soft-bodied animal, measuring up to 7 cm in body length, and it ...
'' and relatives), ''
Pambdelurion ''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 ar ...
'' and ''
Kerygmachela ''Kerygmachela kierkegaardi'' is a 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 rela ...
.'' It is most likely
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
, with ''Kerygmachela'' and ''Pambdelurion'' more basal than the clade compose of Opabiniidae, Radiodonta and other arthropods.


Anatomy

Dinocaridids were
bilaterally symmetrical Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pla ...
, with a mostly non-mineralized
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
and a body divided into two major groupings of tagmata (body-sections): head and trunk. The head apparently unsegmented 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 Radiodonta is an extinct Order (biology), 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 ...
) and usually paired, but highly fused into a nozzle-like structure in
Opabiniidae Opabiniidae is an extinct family of bizarre marine stem-arthropods. Its type and best-known genus is '' Opabinia''. It also contains '' Utaurora''. Opabiniids closely resemble radiodonts, but their frontal appendages were basally fused into a ...
. Based on their preocular position and putative protocerebral origin, the frontal appendages are generally though to be homologous to the labrum of
euarthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chitin, oft ...
s and primary antennae of
onychophoran Onychophora (from grc, ονυχής, , "claws"; and , , "to carry"), commonly known as velvet worms (due to their velvety texture and somewhat wormlike appearance) or more ambiguously as peripatus (after the first described genus, '' Peripatus ...
, while subsequent evidence also suggest a deutocerebral origin (homologous to the jaws of onychophora and
great appendage Great appendages are claw-like appendages which attach to the heads of the "great appendage arthropods", a name usually refers to Megacheira, a class of extinct arthropod characterized by a pair of "short-great appendages" bearing in front of the ...
s/ antennae/
chelicerae The chelicerae () are the mouthparts of the subphylum Chelicerata, an arthropod group that includes arachnids, horseshoe crabs, and sea spiders. Commonly referred to as "jaws", chelicerae may be shaped as either articulated fangs, or similarly ...
of euarthropods). The trunk possessed multiple segments, each with its own gill branch and swimming flaps (lobes). It is thought that these flaps moved in an up-and-down motion, in order to propel the animal forward in a fashion similar to the
cuttlefish Cuttlefish or cuttles are marine molluscs of the order Sepiida. They belong to the class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique internal shell, the cuttlebone, which is used for control of ...
. In gilled lobopodian genera, the trunk may have borne a lobopodous limb (lobopod) underneath each of the flaps. The midgut of dinocaridids had paired digestive glands similar to those of siberiid lobopodians and cambrian euarthropods. The dinocaridid
brain A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a v ...
is relatively simple than those of an euarthropod (3-segmented), it is thought to be comprised either 1 (only protocerebrum) or 2 cerebral
ganglion A ganglion is a group of neuron cell bodies in the peripheral nervous system. In the somatic nervous system this includes dorsal root ganglia and trigeminal ganglia among a few others. In the autonomic nervous system there are both sympatheti ...
s (protocerebrum and deutocerebrum).


Classification

Although some authors may rather suggest different taxonomic affinities (e.g. as
cycloneuralia Cycloneuralia is a clade of ecdysozoan animals including the Scalidophora (Kinorhynchans, Loriciferans, Priapulids) and the Nematoida ( nematodes, Nematomorphs). It may be paraphyletic, or may be a sister group to Panarthropoda. Or perhaps P ...
n relatives), most of the
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
studies suggest that dinocaridids are
stem group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s. Under this scenario, Dinocaridida is a
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
grade in correspond to the arthropod crown group (Euarthropoda or Deuteropoda) and also suggest a
lobopodian The lobopodians, members of the informal group Lobopodia (from the Greek language, Greek, meaning "blunt feet"), or the formally erected phylum Lobopoda Cavalier-Smith (1998), are panarthropods with stubby legs called lobopods, a term which may ...
origin of the arthropod lineage. In general, the gilled lobopodian genera ''
Pambdelurion ''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 ar ...
'' and ''
Kerygmachela ''Kerygmachela kierkegaardi'' is a 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 rela ...
'' which have lobopodian traits (e.g. lobopodous appendage, annulation) occupied the basal position; while
Opabiniidae Opabiniidae is an extinct family of bizarre marine stem-arthropods. Its type and best-known genus is '' Opabinia''. It also contains '' Utaurora''. Opabiniids closely resemble radiodonts, but their frontal appendages were basally fused into a ...
and
Radiodonta Radiodonta is an extinct Order (biology), 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 ...
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. In the original description, Dinocaridida was composed of only Opabiniidae and Radiodonta. With the exclusion of questionable taxa (e.g. the putative opabiniid ''
Myoscolex ''Myoscolex'' is an early animal species known from the Cambrian Emu Bay Shale in South Australia. It was interpreted as an annelid and some supports that theory, while it was also considered as an arthropod close to '' Opabinia'', through other ...
''), the former were known only by ''Opabinia'', while all radiodont species were grouped under a single family:
Anomalocarididae Anomalocarididae (occasionally mis-spelt Anomalocaridae) is an extinct family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpo ...
(hence the previous common name 'Anomalocaridids'). In later studies, the gilled lobopodians ''Pambdelurion'' and ''Kerygmachela'' were also regarded to be dinocaridids, a new opabiniid genus ''
Utaurora ''Utaurora'' is an extinct genus of opabiniid, which were bizarre stem-arthropods closely related to true arthropods and radiodonts; the type species is ''U. comosa''. The animal's fossils come from the Cambrian of Utah. This genus is so far ...
'' was described, and many radiodonts were reassigned to other new families (
Amplectobeluidae ''Amplectobeluidae'' is a clade of Cambrian Radiodonta, radiodonts. Definition In 2014, ''Amplectobeluidae'' was defined as the most inclusive clade including ''Amplectobelua, Amplectobelua symbrachiata'' but not ''Anomalocaris, Anomalocaris ca ...
,
Tamisiocarididae Tamisiocarididae is a family of radiodonts, extinct marine animals related to arthropods, that bore finely-spined appendages that were presumably used in filter-feeding. When first discovered, the clade was named Cetiocaridae after a speculative ...
and
Hurdiidae Hurdiidae is an extinct cosmopolitan distribution, cosmopolitan Family (biology), family of radiodonts, a group of Crown group#Stem groups, stem-group arthropods, which lived during the Paleozoic Era. It is the most long-lived radiodont clade, la ...
).


Distribution

The group was geographically widespread, and has been reported from
Cambrian The Cambrian Period ( ; sometimes symbolized C with bar, Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million ...
strata in
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...
,
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
,
Greenland Greenland ( kl, Kalaallit Nunaat, ; da, Grønland, ) is an island country in North America that is part of the Kingdom of Denmark. It is located between the Arctic and Atlantic oceans, east of the Canadian Arctic Archipelago. Greenland is t ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
,
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
and
Russia Russia (, , ), or the Russian Federation, is a List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, Northern Asia. It is the List of countries and dependencies by area, largest country in the ...
, as well as the
Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start ...
of
Morocco Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
and
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
of
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
.


Notes


References

Cambrian arthropods Devonian arthropods Ordovician arthropods Cambrian first appearances Early Devonian extinctions Burgess Shale fossils Maotianshan shales fossils Sirius Passet fossils Paleozoic animals of Asia Paleozoic animals of North America Prehistoric China Paleozoic Greenland Arthropod classes Prehistoric protostome classes Paraphyletic groups {{paleo-arthropod-stub