Nektaspida (also called Naraoiida, Nektaspia and Nectaspida) is an extinct

order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...

of non- mineralised

artiopodan Artiopoda is a clade 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 Trilobitomor ...

arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...

s. They are known from the lower-

Cambrian The Cambrian ( ) is the first geological period of the Paleozoic Era, and the Phanerozoic Eon. The Cambrian lasted 51.95 million years from the end of the preceding Ediacaran period 538.8 Ma (million years ago) to the beginning of the Ordov ...

to the upper

Silurian The Silurian ( ) is a geologic period and system spanning 23.5 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the third and shortest period of t ...

. Originally classified as

trilobite Trilobites (; meaning "three-lobed entities") are extinction, extinct marine arthropods that form the class (biology), class Trilobita. One of the earliest groups of arthropods to appear in the fossil record, trilobites were among the most succ ...

s, which they superficially resemble, they are now placed as close relatives as members of the

Trilobitomorpha Artiopoda is a clade 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 Trilobitomor ...

within Artiopoda. The order is divided into three major families;

Emucarididae Emucarididae is an extinct family of soft-shelled trilobite-like arthropods (nektaspids) from the Lower Cambrian of South Australia and South China. It contains only two genera – '' Emucaris'' and '' Kangacaris''. Two species were described in ...

Liwiidae Liwiidae is a family of arthropods in the order Nektaspida. Members are known from the Cambrian and Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, ...

, and

Naraoiidae Naraoiidae is a family, of extinct, soft-shelled trilobite-like arthropods, that belongs to the order Nektaspida. Species included in the Naraoiidae are known from the second half of the Lower Cambrian to the end of the Upper Silurian. The total ...

Naming history and taxonomic placement

The order was originally proposed by Raymond in 1920 as Nektaspia. Størmer corrected it to Nectaspida for the 1959 ''

Treatise on Invertebrate Paleontology The ''Treatise on Invertebrate Paleontology,'' published from 1953–2007 by the Geological Society of America and the University of Kansas, then 2009–present by the University of Kansas Paleontological Institute, is a definitive multi-authore ...

'' to conform with the names of the other trilobite orders. Whittington described it in 1985 with the spelling Nektaspida; the revised 1997 Treatise by Raymond and Fortey uses this spelling, as do other modern works. Whittington (1985) placed the order in the

Trilobita Trilobites (; meaning "three-lobed entities") are extinction, extinct marine arthropods that form the class (biology), class Trilobita. One of the earliest groups of arthropods to appear in the fossil record, trilobites were among the most succ ...

. Cotton & Braddy (2000) place it in a new "Trilobite clade" containing the Trilobita, recognizing the close affinities of the nektaspids to trilobites. However this necessitates the inclusion of genera that look very little like trilobites., it was formerly placed in the stem-group to the

chelicerata The subphylum Chelicerata (from Neo-Latin, , ) constitutes one of the major subdivisions of the phylum Arthropoda. Chelicerates include the sea spiders, horseshoe crabs, and arachnids (including harvestmen, scorpions, spiders, solifuges, tic ...

subdivision of the Arthropoda phylum. However, it currently considered part of

Artiopoda Artiopoda is a clade 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 Trilobitomor ...

, the clade that contains trilobites and their close relatives.

Anatomy

The group is united by several morphological characters, including reduced or absent lateral eyes, a

hypostome In zoology, the hypostome can refer to structures in distinct animal groups: *Hypostome (trilobite), the ventral mouthpart plate in trilobites *Hypostome (tick) The hypostome (also called the maxilla, radula, or Labium (insect), labium) is a Os ...

with a natant attachment, extensive articulation overlap between unfused

trunk Trunk may refer to: Biology * Trunk (anatomy), synonym for torso * Trunk (botany), a tree's central superstructure, and the stem of woody plants * Trunk of corpus callosum, in neuroanatomy * Elephant trunk, the proboscis of an elephant Comput ...

tergites A ''tergum'' (Latin for "the back"; : ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or 'margin'. ...

, and fused

pleurae The pleurae (: pleura) are the two flattened closed sacs filled with pleural fluid, each ensheathing each lung and lining their surrounding tissues, locally appearing as two opposing layers of serous membrane separating the lungs from the media ...

that do not form

lateral Lateral is a geometric term of location which may also refer to: Biology and healthcare * Lateral (anatomy), a term of location meaning "towards the side" * Lateral cricoarytenoid muscle, an intrinsic muscle of the larynx * Lateral release ( ...

spines. Preserved soft tissue of the group indicates that the cephalic (head) shield of nektaspids and liwiids contained branched digestive glands. The Naraoiidae have their exoskeletons mostly composed of only two major shields, which have a single articulation point.

Ecology

The naraoiids have been interpreted as

benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from the Ancient Greek word (), meaning "the depths". ...

organisms that were opportunistic scavengers and predators of soft-bodied prey, with the spine-like endites of the limbs possibly allowing soft-bodied prey to be shredded before ingestion. The differences in gut morphology between some species of naraoiids suggests that some species only intermittently fed, while others regularly fed.

Evolutionary history

The group first appeared and was most diverse in the

, especially during the

Cambrian Series 2 Cambrian Series 2 is the unnamed 2nd series of the Cambrian. It lies above the Terreneuvian series and below the Miaolingian. Series 2 has not been formally defined by the International Commission on Stratigraphy, lacking a precise lower boundary ...

are only known from the Cambrian Series 2 of East

Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...

(now Australia) and the then nearby

South China South China ( zh, s=, p=Huá'nán, j=jyut6 naam4) is a geographical and cultural region that covers the southernmost part of China. Its precise meaning varies with context. A notable feature of South China in comparison to the rest of China is ...

were diverse in low latitudes during Cambrian Series 2 and the following

Miaolingian The Miaolingian is the third Series of the Cambrian Period, and was formally named in 2018. It lasted from about to million years ago and is divided in ascending order into 3 stages: the Wuliuan, Drumian, and Guzhangian. The Miaolingian is prec ...

Laurentia Laurentia or the North American craton is a large continental craton that forms the Geology of North America, ancient geological core of North America. Many times in its past, Laurentia has been a separate continent, as it is now in the form of ...

and in South China.

first appeared in

Baltica Baltica is a paleocontinent that formed in the Paleoproterozoic and now constitutes northwestern Eurasia, or Europe north of the Trans-European Suture Zone and west of the Ural Mountains. The thick core of Baltica, the East European Craton, i ...

during the Cambrian, and are absent from major Cambrian deposits elsewhere during this period, but are widespread in the southern high-latitude margin of Gondwana during the

Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, and the second of twelve periods of the Phanerozoic Eon (geology), Eon. The Ordovician spans 41.6 million years f ...

. The youngest known member of the group is the Laurentian naraoiid ''Naraoia bertensis'' from the

Bertie Formation The Bertie Group or Bertie Limestone, also referred to as the Bertie Dolomite and the Bertie Formation, is an Silurian, upper Silurian (Pridoli epoch, Pridoli, or Cayugan) geologic Group (stratigraphy), group and Lagerstätte in southern Ontario, ...

in Ontario, Canada, dating to the

Upper Silurian The Silurian ( ) is a geologic period and system spanning 23.5 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the third and shortest period of t ...

Gallery

References

External links

Are Naraoids trilobites?
��photographs of specimens and discussion of classification controversy.

��more technical overview with references and descriptions of known genera. {{Taxonbar, from=Q3337678 Prehistoric arthropod orders Cambrian first appearances Silurian extinctions