Archaeoniscus

{{Short description|Extinct genus of crustaceans}} {{Automatic taxobox | fossil_range = {{fossilrange|Bajocian|Albian|latest=Santonian}} | image = Archaeoniscus brodiei.jpg | image_caption = Fossil of ''A. brodiei'' | display_parents = 4 | parent_authority = Haack, 1918 | taxon = Archaeoniscus | authority = Milne-Edwards, 1843 | type_species = {{extinct}}'''''Archaeoniscus brodiei''''' | type_species_authority = Milne-Edwards, 1843 | subdivision_ranks = Species | subdivision_ref =<ref name=WoRMS>{{BioRef|WoRMS|title=''Archaeoniscus'' H. Milne Edwards, 1843 † |id=607630 |access-date=29 December 2022}}</ref> | subdivision = *{{extinct}}'''''A. brodiei''''' Milne-Edwards, 1843 *{{extinct}}'''''A. edwardsii''''' Westwood, 1854 *{{extinct}}'''''A.? texanus''''' Wieder and Feldmann, 1992 *{{extinct}}'''''A. aranguthyorum''''' Feldmann ''et al.'', 1998 *{{extinct}}'''''A. coreaensis''''' Park ''et al.'', 2012 *{{extinct}}'''''A. italiensis''''' Jones ''et al.'', 2014 }}

'''''Archaeoniscus''''' ({{IPAc-en|ˌ|ɑːr|k|iː|'|oʊ|n|ɪ|s|k|ə|s}}) is a genus of prehistoric isopods that first appeared during the Bajocian stage of the Middle Jurassic. It is a widespread genus with a paleogeographic distribution encompassing the continental margin environments of the central Atlantic Ocean and the western Tethys Ocean. Fossils of ''Archaeoniscus'' suggest that this genus lived in diverse aquatic habitats, including the marine, paralic, and freshwater environments. While earlier descriptions suggested that it may have had an ectoparasitic association with fishes, some researchers argue that at least two species, ''A. aranguthyorum'' and ''A. coreaensis'', lived a benthic free-living lifestyle based on morphological characteristics that are either unsuitable for or unrelated to parasitic behavior.

Six species have been named: two species are known from the Jurassic period rocks in Europe, and the other four are known from the Cretaceous period rocks in Europe, North America and East Asia. Of these species, the taxonomic position of the only Late Cretaceous species, ''A. texanus'', is uncertain, for some researchers claim that this species should be neglected because of the significant morphological differences in comparison to the other species within the genus. Some specimens of the most completely known species, ''A. aranguthyorum'', suggest that the female individuals of this species possibly had brood pouches, indicating that the genus may have had sexual dimorphism. An alternative interpretation suggests that the brood pouches may actually be the impressions of the hindgut and anus. Additionally, two unnamed indeterminate species are known from the Early Cretaceous rocks in Mexico and Egypt.

==Discovery== Reverend Peter Bellinger Brodie first discovered the crustacean fossils from the Purbeck Group in the Vale of Wardour, England. He asked Henri Milne-Edwards to examine some specimens and formally describe them. The specimens sent to Milne-Edwards measured {{cvt|9|-|12|cm|in}} in length, though other descriptions indicate their lengths were about {{cvt|2.4|cm|in}}.<ref name="Feldmannetal1998" /> Brodie claimed that he also found larger specimens that are about the size of small trilobites. Milne-Edwards identified that the specimens belong to an order Isopoda, and suggested that it may belong within the range of a family Cymothoidae, possibly an intermediate species between ''Serolis'' and ''Cymothoa''. In 1843, Milne-Edwards proposed a genus name ''Archaeoniscus'' and a species name ''A. brodiei'' in honor of the discoverer Rev. P. B. Brodie.<ref>{{cite journal|last1=Milne-Edwards|first1=H.|year=1843|title=Note sur deux Crustacés fossiles de l'ordre des Isopodes|journal=Annales des Sciences Naturelles, 2e série (Zoologie)|volume=20|pages=326–329}}</ref><ref>{{cite journal|last1=Milne-Edwards|first1=H.|year=1844|title=XVI.—On a Fossil Crustacean of the Order Isopoda, discovered by the Rev. P. B. Brodie in the Wealden formation of Britain|journal=The Annals and Magazine of Natural History|volume=13|issue=82|pages=110–111|doi=10.1080/03745484409442579|url=https://www.biodiversitylibrary.org/part/3665 }}</ref> From the same lithostratigraphic group, Charles Willcox collected various insect fossils and sent them to John O. Westwood. Upon his examination, Westwood found several isopod specimens which varied in size, being either equal to or larger than ''A. brodiei''. Westwood considered them to be a new species of ''Archaeoniscus'', so he designated these specimens under a new species name ''A. edwardsii'' in honor of Milne-Edwards.<ref>{{cite journal|year=1854|last1=Westwood|first1=J.O.|title=Contributions to Fossil Entomology|journal=Quarterly Journal of the Geological Society|volume=10|issue=1–2|pages=378–396|doi=10.1144/gsl.jgs.1854.010.01-02.43|s2cid=129712238 |url=https://zenodo.org/record/25050 |doi-access=free}}</ref> While ''A. brodiei'' is discovered from the Late Jurassic-Early Cretaceous (Oxfordian-Berriasian) strata in England, France and Germany, ''A. edwardsii'' is only discovered from the Early Cretaceous (Berriasian) strata in England, though both species may be synonymous.<ref name=Vegaetal2019>{{cite journal|last1=Vega|first1=Francisco J.|first2=Niel L. |last2=Bruce|first3=Oscar |last3=González-León|first4=Jason |last4=Jeremiah|first5=María de Lourdes |last5=Serrano-Sánchez |first6=Jesús |last6=Alvarado-Ortega|first7=Josep A. |last7=Moreno-Bedmar|year=2019|title=Lower Cretaceous marine isopods (Isopoda: Cirolanidae, Sphaeromatidae) from the San Juan Raya and Tlayúa formations, Puebla, Mexico|journal=Journal of Crustacean Biology|volume=39|issue=2|pages=121–135|doi=10.1093/jcbiol/ruy107|s2cid=92289017|doi-access=free}}</ref><ref>{{Cite journal|last1=Bicknell |first1=R. D. C. |last2=Klompmaker |first2=A. A. |last3=Smith |first3=P. M. |last4=Hegna |first4=T. A. |year=2025 |title=Exploring the morphology and taphonomy of ''Archaeoniscus brodiei''—a gregarious, Early Cretaceous isopod |journal=The Science of Nature |volume=112 |issue=2 |at=16 |doi=10.1007/s00114-025-01962-8 |pmid=39928166 |pmc=11811265 |doi-access=free |bibcode=2025SciNa.112...16B }}</ref>

In 1918, Wilhelm Haack erected a monotypic family Archaeoniscidae and claimed that it is distinct from the suborder Oniscidea based on the large pleotelson, the shapes of antennulae, antennae and uropods.<ref>{{cite journal|last1=Haack|first1=W.|year=1918|title=Ueber einen Isopoden aus dem Sepulit des weslichen, Osnegs (''Archaeoniscus brodei'', Milne-Edwards)|journal=Jahrbuch der Königlichen Preussischen Geologischen Landesanstalt|volume=39|pages=73–102}}</ref> In 1992, Robert W. Wieder and Rodney M. Feldmann examined various isopod fossils from Austin Chalk in Texas, United States. Of these fossils, Wieder and Feldmann considered some specimens that are only known from posterior exuviae to belong to ''Archaeoniscus'' based on the raised axial node on the pleotelson which they claim to be distinct from almost every other known isopod species. Although they tentatively referred these specimens under a new species name, ''A. texanus'', they acknowledged that the taxonomic placement of these specimens is difficult; still, they considered this placement prudent enough and better than erecting a new genus name and establishing the taxonomic affinities for the specimens.<ref>{{cite journal|last1=Wieder|first1=Robert W.|last2=Feldmann|first2=Rodney M.|year=1992|title=Mesozoic and Cenozoic fossil isopods of North America|journal=Journal of Paleontology|volume=66|issue=6|pages=958–972|doi=10.1017/s0022336000021041|s2cid=132171864 }}</ref> The stratigraphic age of Austin Chalk is during the Late Cretaceous (Coniacian-Santonian), so ''A. texanus'' is the youngest known species of this genus.<ref name=Vegaetal2019/>

In 1998, Feldmann and his colleagues described a new species, ''A. aranguthyorum'', which was discovered from the Tlayúa Formation in Mexico.<ref name=Feldmannetal1998>{{cite journal|year=1998|last1=Feldmann|first1=R.M.|last2=Vega|first2=F.J.|last3=Applegate|first3=S.P.|last4=Bishop|first4=G.A.|title=Early Cretaceous Arthropods from the Tlayúa Formation at Tepexi de Rodriguez, Puebla, Mexico|journal=Journal of Paleontology|volume=72|issue=1|pages=79–90|url=https://isopods.nhm.org/pdfs/3061/3061.pdf|doi=10.1017/S0022336000024033|s2cid=2775985|jstor=1306680}}</ref> In 2012, Park and his colleagues described a new freshwater-living species, ''A. coreaensis'', which was discovered from the Jinju Formation in South Korea. Both species, ''A. aranguthyorum'' and ''A. coreaensis'', lived during the Early Cretaceous (Albian).<ref name=Parketal2012>{{Cite journal |last1=Park |first1=Tae-Yoon |last2=Wilson |first2=George D. F. |last3=Lee |first3=Dong-Chan |last4=Choi |first4=Duck K. |date=2012 |title=Occurrence of the Isopod ''Archaeoniscus coreaensis'' New Species from the Lower Cretaceous Jinju Formation, Korea |url=https://www.researchgate.net/publication/259453671 |journal=Journal of Paleontology |volume=86 |issue=4 |pages=626–640 |doi=10.1666/11-131R.1 |jstor=23253719 |s2cid=131617665 }}</ref> In 2014, a new species, ''A. italiensis'', was discovered from the Monte Fallano Plattenkalk, Italy. Within the genus, ''A. italiensis'' is the oldest known species that lived during the Middle Jurassic (Bajocian-Bathonian).<ref>{{cite journal|last1=Jones|first1=W.T.|last2=Feldmann|first2=R.M.|last3=Garassino|first3=A.|year=2014|title=Three New Isopod Species and a New Occurrence of the Tanaidacean ''Niveotanais brunnensis'' Polz, 2005 from the Jurassic Plattenkalk Beds of Monte Fallano, Italy|journal=Journal of Crustacean Biology|volume=34|issue=6|pages=739–753|doi=10.1163/1937240X-00002282|doi-access=free}}</ref> Unnamed Early Cretaceous species have been discovered from the Valanginian-Hauterivian stage rocks in Mexico and from the Albian stage rocks in Egypt.<ref name=Vegaetal2019/>

==Description== [[File:Archaeoniscus coreaensis.png|thumb|Speculative life restoration of ''Archaeoniscus coreaensis'']] ''Archaeoniscus'' generally has an oval-shaped body which is dorsoventrally compressed. The lateral margins of the pereion (the thorax of crustaceans) consist of broad, overlapping coxal plates with distinguishable sutures. The head is embedded into the first pereonite (the first segment of the pereion); it also possesses mandibles positioned transversely with a broad incisor process and has globular eyes on its lateral margin. The four anterior pleonites (parts of the abdomen of crustaceans called pleon) are long and wide, while the fifth pleonite is relatively reduced and does not extend to the lateral margin. ''Archaeoniscus'' lacks marginal spines on its broad and semicircular pleotelson, the structure formed by the fusion of the last abdominal segment of an arthropod and its telson. The unmodified ambulatory pereiopods have distinct coxal dorsal articulations, which are distinguishable from modern cymothoids, suggesting that the genus was not suited for ectoparasitic behavior. The antennulae is large and multiarticulate, and the spine-like uropodal rami have a narrow, parallel-sided protopod.<ref name=Parketal2012/> ''A. texanus'' has significant morphological differences in comparison to other species within this genus, including the presence of a long fifth pleonite, triangular epimeres, shorter pleotelson and more robust uropod.<ref name=Vegaetal2019/>

==Classification== ''Archaeoniscus'', initially described as a possible cymothoid and now contained in its own monotypic family Archaeoniscidae, has an unclear phylogenetic position within the order Isopoda. This controversial classification may have occurred because the earlier studies have misinterpreted the anatomical characteristics of ''Archaeoniscus'', such as the reconstructions provided by Haack in 1918 and by Vega and his colleagues in 2005, and the morphological data of this genus is insufficient for a definitive suprafamilial classification.<ref name=Parketal2012/> In 1969, Robert R. Hessler argued that some features of this genus resemble those of the suborder Oniscidea, but acknowledged that the features noted by Haack in 1918 does distinguish the genus from that suborder.<ref>{{cite book|year=1969|last1=Hessler|first1=R.R.|editor1=Moore, R.C.|chapter=Peracarida|title=Treatise on Invertebrate Paleontology. Pt. R. Arthropoda 4, Crustacea (except ostracoda)|pages=360–393|publisher=Geological Society of America and University of Kansas Press|location=Lawrence, Kansas|isbn=0-8137-3018-X}}</ref> Although it was commonly placed within the suborder Flabellifera, this view has been disputed since 1989 by Johann Wolfgang Wägele.<ref>{{cite journal|year=1989|last1=Wägele|first1=J.W.|title=Evolution und phylogenetisches System der Isopoda|journal=Zoologica|volume=140|pages=1–242|s2cid=83440947}}</ref> In 1998, Feldmann and his colleagues claimed that possible brood pouches found in some specimens of ''A. aranguthyorum'' may indicate that ''Archaeoniscus'' might belong to the suborder Sphaeromatidea; they assumed that the presence of brood pouch is a characteristic only observed in female sphaeromatid isopods.<ref name=Feldmannetal1998/> Following this suggestion, Vega and his colleagues in 2005 placed this genus within the superfamily Sphaeromatoidea of the suborder Sphaeromatidea.<ref>{{cite journal |first1=Francisco J. |last1=Vega |first2=Niel L. |last2=Bruce |first3=Lourdes |last3=Serrano |first4=Gale A. |last4=Bishop |first5=María del Carmen |last5=Perrilliat |year=2005 |journal=Bulletin of the Mizunami Fossil Museum |volume=32 |pages=25–30 |title=A review of the Lower Cretaceous (Tlayúa Formation: Albian) Crustacea from Tepexi de Rodríguez, Puebla, Central Mexico |url=https://decapoda.nhm.org/pdfs/10374/10374.pdf |s2cid=55376301}}</ref> However, Brandt and his colleagues in 2003 did not consider the genus within the suborder Sphaeromatidea, but considered it to be an ancestral taxon of terrestrial isopods without specific reasoning behind their argument.<ref>{{cite journal|last1=Brandt|first1=A.|last2=Crame|first2=J.A.|last3=Polz|first3=H.|last4=Thomson|first4=M.R.A.|year=2003|title=Late Jurassic Tethyan ancestry of recent southern high-latitude marine isopods (Crustacea, Malacostraca)|journal=Palaeontology|volume=42|issue=4|pages=663–675|doi=10.1111/1475-4983.00090|s2cid=129327151 |doi-access=free}}</ref> In 2012, Park and his colleagues argued that Feldmann and his colleagues have mistaken the impressions of the hindgut and anus of ''A. aranguthyorum'' as possible brood pouches and that all female isopods have brood pouches, and thus the taxonomic position of this genus is still unclear. They also argued that ''A. texanus'' should be removed from the genus, as it significantly differs from other species, possessing flat limbs and five subequal pleonites with the fifth being the longest; in contrast, other species of ''Archaeoniscus'' possess narrow, rod-like limbs and only four free subequal pleonites with the fifth being reduced.<ref name=Parketal2012/> Vega and his colleagues in 2019 also advocated the view that ''A. texanus'' is morphologically different from other species and that a new genus name should be erected for this species. They also considered that the latest occurrence of ''Archaeoniscus'' may more likely be the Albian stage of the Early Cretaceous.<ref name=Vegaetal2019/>

==Paleobiology== As the earlier descriptions of this genus indicated a taxonomic association with cymothoids, Feldmann and his colleagues in 1998 considered that ''Archaeoniscus'' may have been ectoparasites on fishes just like modern cymothoids.<ref name=Feldmannetal1998/> Park and his colleagues in 2012 disagreed with this view, claiming that the morphological characteristics of ''A. aranguthyorum'' and ''A. coreaensis'' suggest otherwise. The researchers pointed out that the anterior limbs of ''Archaeoniscus'' are unmodified and ambulatory, contrary to those of modern cymothoids which resemble hooks. They also stated that the shape of the limbs and the flattened body resemble that of serolids or other sphaeromatids which live in benthic zones. Their interpretation suggests that at least two species of ''Archaeoniscus'', ''A. aranguthyorum'' and ''A. coreaensis'', likely lived a benthic lifestyle without any parasitic association with fishes.<ref name=Parketal2012/> Additionally, while Feldmann and his colleagues suggested that ''Archaeoniscus'' had sexual dimorphism based on the possible existence of brood pouches,<ref name=Feldmannetal1998/> Park and his colleagues also disagreed with this claim, arguing that Feldmann and his colleagues misidentified the impressions of the hindgut and anus.<ref name=Parketal2012/>

==References== {{Reflist}}

==External links== {{Portal|Crustaceans|Palaeontology}} *{{Wikispecies-inline|Archaeoniscus|''Archaeoniscus''}}

Category:Isopod genera Category:Late Jurassic crustaceans Category:Late Cretaceous crustaceans Category:Middle Jurassic animals of Europe Category:Late Jurassic animals of Europe Category:Cretaceous arthropods of North America Category:Early Cretaceous arthropods of Asia Category:Bajocian genus first appearances Category:Santonian genus extinctions Category:Fossil taxa described in 1843 Category:Cretaceous Mexico Category:Cretaceous Egypt Category:Jurassic England Category:Jurassic France Category:Cretaceous England Category:Cretaceous Texas Category:Cretaceous South Korea Category:Jurassic Italy Category:Bathonian genera Category:Callovian genera Category:Oxfordian genera Category:Kimmeridgian genera Category:Tithonian genera Category:Valanginian genera Category:Berriasian genera Category:Hauterivian genera Category:Barremian genera Category:Aptian genera Category:Albian genera Category:Cenomanian genera Category:Turonian genera Category:Coniacian genera Category:Fossils of Texas Category:Fossils of England Category:Fossils of France Category:Fossils of Mexico Category:Fossils of Italy Category:Fossils of South Korea Category:Early Cretaceous arthropods of Europe Category:Jurassic arthropods of Europe Category:Middle Jurassic crustaceans Category:Early Cretaceous crustaceans Category:Crustaceans described in 1843 Category:Prehistoric crustacean genera Category:Sphaeromatidea