{{Short description|Extinct order of arthropods}} {{Automatic taxobox | fossil_range = {{fossil range|Atdabanian|Silurian}} | image = Naraoia compacta fossil.jpg | image_caption = ''Naraoia compacta'' fossils with preserved appendages | taxon = Nektaspida | authority = Raymond, 1920 | subdivision_ranks = Families | subdivision_ref = <ref name=":0"/> | subdivision = * †Emucarididae * †Liwiidae * †Naraoiidae * †''Buenaspis'' * †''Campanamuta''? * †''Panlongia''<ref>{{cite journal |last1=Luo |first1=Shengxiang |last2=Liu |first2=Jianni |title=Exceptionally preserved Panlongia tetranodusa from the early Cambrian sheds light on the evolution of arthropods |journal=Science Bulletin |date=February 2024 |volume=69 |issue=3 |pages=308–313 |doi=10.1016/j.scib.2023.11.043}}</ref> }}
'''Nektaspida''' (also called '''Naraoiida''', '''Nektaspia''' and '''Nectaspida''') is an extinct order of non-mineralised artiopodan arthropods. They are known from the lower-Cambrian to the upper Silurian. Originally classified as trilobites, which they superficially resemble, they are now placed as close relatives as members of the Trilobitomorpha within Artiopoda. The order is divided into three major families; Emucarididae, Liwiidae, and Naraoiidae.
== Naming history and taxonomic placement ==
The order was originally proposed by Raymond in 1920 as Nektaspia.<ref>{{cite journal |last=Raymond |first=P.E. |year=1920 |title=The appendages, anatomy, and relationships of trilobites |url=https://biodiversitylibrary.org/page/27744126 |journal=Memoirs of the Connecticut Academy of Arts and Sciences |volume=7 |pages=1–169}}</ref> Størmer corrected it to Nectaspida for the 1959 ''Treatise on Invertebrate Paleontology'' 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.<ref name="Budd1996">{{Cite journal |last1=Budd |first1=G. E. |year=1999 |title=A nektaspid arthropod from the Early Cambrian Sirius Passet fauna, with a description of retrodeformation based on functional morphology |journal=Palaeontology |volume=42 |issue=1 |pages=99–122 |doi=10.1111/1475-4983.00064|bibcode=1999Palgy..42...99B |s2cid=128566745 |doi-access=free }}</ref> Whittington (1985) placed the order in the Trilobita. 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.,<ref name="Budd1996" /> it was formerly placed in the stem-group to the chelicerata subdivision of the Arthropoda phylum.<ref name="Budd2000">{{cite journal |last1=Budd |first1=G. E. |last2=Jensen |first2=S. |year=2000 |title=A critical reappraisal of the fossil record of the bilaterian phyla |journal=Biological Reviews of the Cambridge Philosophical Society |volume=75 |issue=2 |pages=253–95 |doi=10.1111/j.1469-185X.1999.tb00046.x |pmid=10881389 |s2cid=39772232}}</ref> However, it currently considered part of Artiopoda, the clade that contains trilobites and their close relatives.<ref>{{Cite journal |last1=Pérez-Peris |first1=Francesc |last2=Laibl |first2=Lukáš |last3=Lustri |first3=Lorenzo |last4=Gueriau |first4=Pierre |last5=Antcliffe |first5=Jonathan B. |last6=Enright |first6=Orla G. Bath |last7=Daley |first7=Allison C. |date=March 2021 |title=A new nektaspid euarthropod from the Lower Ordovician strata of Morocco |url=https://www.cambridge.org/core/journals/geological-magazine/article/abs/new-nektaspid-euarthropod-from-the-lower-ordovician-strata-of-morocco/50E1FE68F9E38151AA2EC93E92222BFE |journal=Geological Magazine |language=en |volume=158 |issue=3 |pages=509–517 |doi=10.1017/S001675682000062X |bibcode=2021GeoM..158..509P |s2cid=221859513 |issn=0016-7568}}</ref>
== Anatomy == left|thumb|Digestive systems of various naraoiid species The group is united by several morphological characters, including reduced or absent lateral eyes, a hypostome with a natant attachment, extensive articulation overlap between unfused trunk tergites, and fused pleurae that do not form lateral spines.<ref>{{Cite journal |last1=Mayers |first1=Benjamin |last2=Aria |first2=Cédric |last3=Caron |first3=Jean‐Bernard |date=January 2019 |editor-last=Zhang |editor-first=Xi‐Guang |title=Three new naraoiid species from the Burgess Shale, with a morphometric and phylogenetic reinvestigation of Naraoiidae |url=https://onlinelibrary.wiley.com/doi/10.1111/pala.12383 |journal=Palaeontology |language=en |volume=62 |issue=1 |pages=19–50 |doi=10.1111/pala.12383 |bibcode=2019Palgy..62...19M |s2cid=135331371 |issn=0031-0239|url-access=subscription }}</ref> Preserved soft tissue of the group indicates that the cephalic (head) shield of nektaspids and liwiids contained branched digestive glands.<ref name=":0" /> The Naraoiidae have their exoskeletons mostly composed of only two major shields, which have a single articulation point.<ref>{{Cite journal |last1=Bond |first1=Andrew D. |last2=Edgecombe |first2=Gregory D. |date=January 2021 |editor-last=Zhang |editor-first=Xi‐Guang |title=Phylogenetic response of naraoiid arthropods to early–middle Cambrian environmental change |journal=Palaeontology |language=en |volume=64 |issue=1 |pages=161–177 |doi=10.1111/pala.12516 |bibcode=2021Palgy..64..161B |s2cid=229454194 |issn=0031-0239|doi-access=free }}</ref>
== Ecology == The naraoiids have been interpreted as benthic 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 fed only intermittently, while others fed regularly.<ref>{{Cite journal |last1=VANNIER |first1=JEAN |last2=CHEN |first2=JUN-YUAN |date=2007-01-02 |title=Digestive system and feeding mode in Cambrian naraoiid arthropods |url=http://dx.doi.org/10.1111/j.1502-3931.2002.tb00072.x |journal=Lethaia |volume=35 |issue=2 |pages=107–120 |doi=10.1111/j.1502-3931.2002.tb00072.x |issn=0024-1164|url-access=subscription }}</ref>
== Evolutionary history == The group first appeared and was most diverse in the Cambrian, especially during the Cambrian Series 2. Emucarididae are only known from the Cambrian Series 2 of East Gondwana (now Australia) and the then nearby South China. Naraoiidae were diverse in low latitudes during Cambrian Series 2 and the following Miaolingian in Laurentia and in South China. Liwiidae first appeared in Baltica 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.<ref name=":0">{{Cite journal |last1=Pérez-Peris |first1=Francesc |last2=Laibl |first2=Lukáš |last3=Lustri |first3=Lorenzo |last4=Gueriau |first4=Pierre |last5=Antcliffe |first5=Jonathan B |last6=Bath Enright |first6=Orla G |last7=Daley |first7=Allison C |date=2020-07-10 |title=A new nektaspid euarthropod from the Lower Ordovician strata of Morocco |url=http://dx.doi.org/10.1017/s001675682000062x |journal=Geological Magazine |volume=158 |issue=3 |pages=509–517 |doi=10.1017/s001675682000062x |s2cid=221859513 |issn=0016-7568}}</ref> The youngest known member of the group is the Laurentian naraoiid ''Naraoia bertensis'' from the Bertie Formation in Ontario, Canada, dating to the Upper Silurian.<ref>{{Cite journal |last1=Caron |first1=Jean-Bernard |last2=Rudkin |first2=David M. |last3=Milliken |first3=Stuart |date=November 2004 |title=A new Late Silurian (Pridolian) naraoiid (Euarthropoda: Nektaspida) from the Bertie Formation of southern Ontario, Canada—delayed fallout from the Cambrian explosion |url=https://www.cambridge.org/core/journals/journal-of-paleontology/article/abs/new-late-silurian-pridolian-naraoiid-euarthropoda-nektaspida-from-the-bertie-formation-of-southern-ontario-canadadelayed-fallout-from-the-cambrian-explosion/5E91487CE888A3A31FA30B8D384D7E2B |journal=Journal of Paleontology |language=en |volume=78 |issue=6 |pages=1138–1145 |doi=10.1666/0022-3360(2004)078<1138:ANLSPN>2.0.CO;2 |s2cid=131595361 |issn=0022-3360|url-access=subscription }}</ref>
== Gallery == {{gallery|Misszhouia.png|Life restoration of the naraoiid ''Misszhouia''|File:Buenaspis drawing.jpg|Drawing of the carapace of the ''incertae sedis'' nektaspid ''Buenaspis''|File:Emucaridae.jpg|Restoration of the emucaridids ''Kangacaris'' (blue, left) and ''Emucaris'' (yellow, right)|Liwia_reconstruction.png|Fossil of the Cambrian liwiid ''Liwia'' from Poland|File:Pseudonaraoia hammoni reconstruction.JPG|Life restoration of the naraoiid ''Pseudonaraoia''|File:Naraoia longicaudata.jpg|Fossil of ''Misszhouia longicaudata''|width=200|height=180}}
== References == {{Reflist}}
== External links == * [http://www.trilobites.info/naraoia.htm Are Naraoids trilobites?]—photographs of specimens and discussion of classification controversy. * [https://web.archive.org/web/20040529221712/http://www.peripatus.gen.nz/Taxa/Arthropoda/Trilobita/Nectaspida.html Order Nectaspida]—more technical overview with references and descriptions of known genera.
{{Artiopoda}} {{Taxonbar|from=Q3337678}}
Category:Nektaspida Category:Prehistoric arthropod orders Category:Cambrian first appearances Category:Silurian extinctions