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{{Short description|Order of molluscs}}
{{Taxobox
{{Automatic taxobox
| fossil_range = Recent
| image = Acochlidium fijiiensis.png
| image = Acochlidium fijiiensis.png
| image_caption = ''[[Acochlidium fijiiensis]]''
| image_caption = ''[[Acochlidium fijiiensis]]''
| display_parents = 2
| regnum = [[Animal]]ia
| phylum = [[Mollusca]]
| taxon = Acochlidiacea
⚫
| authority = [[Nils Hjalmar Odhner|Odhner]], 1937<ref name="Odhner 1937">{{in lang|de}} [[Nils Hjalmar Odhner|Odhner N. H.]] (1937). "''[[Hedylopsis suecica]]'' n.sp. und die Nacktschneckengruppe Acochlidiacea (Hedylacea)". ''[[Zoologischer Anzeiger]]'' 120(3–4): 51–64. 52, 62.</ref>
| classis = [[Gastropoda]]
| unranked_familia = clade [[Heterobranchia]]<br/>clade [[Euthyneura]]<br/>clade [[Panpulmonata]]<br/>clade '''Acochlidiacea'''
⚫
| unranked_familia_authority = [[Nils Hjalmar Odhner|Odhner]], 1937<ref name="Odhner 1937">{{de icon}} [[Nils Hjalmar Odhner|Odhner N. H.]] (1937). "''[[Hedylopsis suecica]]'' n.sp. und die Nacktschneckengruppe Acochlidiacea (Hedylacea)". ''[[Zoologischer Anzeiger]]'' 120(3–4): 51–64. 52, 62.</ref>
| subdivision_ranks =
| subdivision =
| synonyms_ref = <ref name="Bouchet 2005">{{Bouchet 2005}}</ref><ref name="Gofas 2010"/>
| synonyms_ref = <ref name="Bouchet 2005">{{Bouchet 2005}}</ref><ref name="Gofas 2010"/>
| synonyms =
| synonyms =
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Acochlidea<br/>
Acochlidea<br/>
Acochlidia
Acochlidia
⚫
| diversity = 46 species
| diversity_link = Diversity of gastropods
| diversity_ref = <!--per Overview of species --><!--<ref name="Jörger 2012"/>--><!--33 species per Jörger 2012 + 1 Bathyhedyle-->
⚫
| diversity =
33 species + 9 undescribed in 2012
| diversity_ref = <ref name="Jörger 2012"/>
| subdivision_ranks = Families
| subdivision_ranks = Families
| subdivision =
| subdivision =
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(unranked) [[Microhedylacea]]
(unranked) [[Microhedylacea]]
* [[Asperspinidae]]:
* [[Asperspinidae]]:
* [[Microhedylidae]]
* [[Microhedylidae]]
* [[Bathyhedylidae]]
}}
}}


'''Acochlidiacea''', common name '''acochlidians''', are a [[taxonomic]] clade of very unusual [[sea snail]]s and sea and freshwater [[slug]]s, [[aquatic animal|aquatic]] [[gastropod]] [[mollusk]]s within the large clade [[Heterobranchia]]. '''Acochlidia''' is a variant spelling.
'''Acochlidiacea''', common name '''acochlidians''', are a [[Taxonomy (biology)|taxonomic]] clade of very unusual [[sea snail]]s and sea and freshwater [[slug]]s, [[aquatic animal|aquatic]] [[gastropod]] [[mollusk]]s within the large clade [[Heterobranchia]]. '''Acochlidia''' is a variant spelling.


[[File:Pseudunela cornuta.png|left|thumb|Schematic drawing of dorsal view of ''[[Pseudunela cornuta]]'' shows some characteristics of acochlidians, but characteristics of other acochlidians differ greatly:<br/>
[[File:Pseudunela cornuta svg hariadhi.svg|left|thumb|Schematic drawing of dorsal view of ''[[Pseudunela cornuta]]'' shows some characteristics of acochlidians, but characteristics of other acochlidians differ greatly:<br/>
f – [[Mollusca#Foot|foot]]<br/>
f – [[Mollusca#Foot|foot]]<br/>
hb – heart bulb<br/>
hb – heart bulb<br/>
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== Description ==
== Description ==
These are mostly very small animals, without a [[Mollusc shell|shell]] or [[gill]]s, distinguished by the [[viscera]]l mass being sharply set off from the rest of the body.<ref name=IZ>{{cite book |author= Barnes, Robert D. |year=1982 |title= Invertebrate Zoology |publisher= Holt-Saunders International |location= Philadelphia, PA|page= 376|isbn= 0-03-056747-5}}</ref>
These are mostly very small animals, without a [[Mollusc shell|shell]] or [[gill]]s, distinguished by the [[viscera]]l mass being sharply set off from the rest of the body.<ref name=IZ>{{cite book |author = Barnes, Robert D. |year=1982 |title= Invertebrate Zoology |publisher= Holt-Saunders International |location= Philadelphia, PA|page= 376|isbn= 0-03-056747-5}}</ref>


Being a small group with only 30 species worldwide known in 2010,<ref>There are 29 valid species plus one undescribed species</ref> and 32 species described in 2011,<ref name="Neusser 2011"/> and 33 in 2012 (+9 undescribed ''[[Pontohedyle]]'' species),<ref name="Jörger 2012"/> these slugs are morphologically and biologically highly aberrant and diverse, comprising a series of unusual characters (e.g. secondary [[gonochorism]], lack of copulatory organs, asymmetric [[radula]]e).<ref name="Jörger 2010"/> Most acochlidians live interstitially in marine sands, while some have conquered limnic systems (uniquely within opisthobranch gastropods).<ref name="Jörger 2010"/>
Being a small group with only 47 species worldwide known in 2023,<ref name="WoRMS">MolluscaBase eds. (2021). [https://molluscabase.org/aphia.php?p=taxdetails&id=411937 MolluscaBase. Acochlidiimorpha.] Accessed on 2023-02-13.</ref> this group has been the subject of active research since 2010, by which date only 32 species were named.<ref name="Neusser 2011"/> These slugs are morphologically and biologically highly aberrant and diverse, comprising a series of unusual characters (e.g. secondary [[gonochorism]], lack of copulatory organs, asymmetric [[radula]]e).<ref name="Jörger 2010"/> Most acochlidians live interstitially in marine sands, while some have conquered limnic systems (uniquely within opisthobranch gastropods).<ref name="Jörger 2010"/>


==Taxonomy==
==Taxonomy==
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Wawra (1987)<ref>Wawra E. (1987). "Zur Anatomie einiger Acochlidia (Gastropoda, Opisthobranchia) mit einer vorläufigen Revision des Systems und einem Anhang über Platyhedylidae (Opisthobranchia, Ascoglossa)". PhD thesis. Universität Wien.</ref> and various authors (2007–2010)<ref name="Jörger 2010"/><ref>Neusser T. P., Jörger K. M. & Schrödl M. (2007). "Exploring cerebral features in Acochlidia (Gastropoda: Opisthobranchia)". ''Bonn. Zool. Beitr.'' '''55''': 301–310.</ref><ref>Jörger K. M., Heβ M., Neusser T. P. & Schrödl M. (2009). "Sex in the beach: spermatophores, dermal insemination and 3D sperm ultrastructure of the aphallic mesopsammic ''[[Pontohedyle milaschewitchii]]'' (Acochlidia, Opisthobranchia, Gastropoda)". ''[[Marine Biology]]'' '''156''': 1159–1170. {{doi|10.1007/s00227-009-1158-5}}.</ref><ref name="Schrödl 2010">Schrödl M. & Neusser T. P. (2010). "Towards a phylogeny and evolution of Acochlidia (Mollusca: Gastropoda: Opisthobranchia)". ''[[Zoological Journal of the Linnean Society]]'' '''158''': 124–154. {{doi|10.1111/j.1096-3642.2009.00544.x}}.</ref> spelled this taxon as Acochlidia.
Wawra (1987)<ref>Wawra E. (1987). "Zur Anatomie einiger Acochlidia (Gastropoda, Opisthobranchia) mit einer vorläufigen Revision des Systems und einem Anhang über Platyhedylidae (Opisthobranchia, Ascoglossa)". PhD thesis. Universität Wien.</ref> and various authors (2007–2010)<ref name="Jörger 2010"/><ref>Neusser T. P., Jörger K. M. & Schrödl M. (2007). "Exploring cerebral features in Acochlidia (Gastropoda: Opisthobranchia)". ''Bonn. Zool. Beitr.'' '''55''': 301–310.</ref><ref>Jörger K. M., Heβ M., Neusser T. P. & Schrödl M. (2009). "Sex in the beach: spermatophores, dermal insemination and 3D sperm ultrastructure of the aphallic mesopsammic ''[[Pontohedyle milaschewitchii]]'' (Acochlidia, Opisthobranchia, Gastropoda)". ''[[Marine Biology]]'' '''156''': 1159–1170. {{doi|10.1007/s00227-009-1158-5}}.</ref><ref name="Schrödl 2010">Schrödl M. & Neusser T. P. (2010). "Towards a phylogeny and evolution of Acochlidia (Mollusca: Gastropoda: Opisthobranchia)". ''[[Zoological Journal of the Linnean Society]]'' '''158''': 124–154. {{doi|10.1111/j.1096-3642.2009.00544.x}}.</ref> spelled this taxon as Acochlidia.


Three families ([[Hedylopsidae]], [[Microhedylidae]] and [[Acochlidiidae]]) are classically recognized.<ref name="Bouchet 2005"/><ref name="Gofas 2010"/> Two controversial classifications (Rankin 1979,<ref name="Rankin 1979"/> [[Yaroslav Starobogatov|Starobogatov]] 1983)<ref name="Starobogatov 1983">{{ru icon}} [[Yaroslav Starobogatov|Starobogatov Ya. I.]] (1983). "Sistema otriada Acochlidiiformes [System of the order Acochlidiiformes]." ''Vsesoiuznoe soveshchanie po izucheniiu molliuskov'', Leningrad, '''7''': 30–32.</ref> have been proposed recently, but they have not been evaluated since.<ref name="Bouchet 2005"/><ref name="Gofas 2010"/>
Three families ([[Hedylopsidae]], [[Microhedylidae]] and [[Acochlidiidae]]) are classically recognized.<ref name="Bouchet 2005"/><ref name="Gofas 2010"/> Two controversial classifications (Rankin 1979,<ref name="Rankin 1979"/> [[Yaroslav Starobogatov|Starobogatov]] 1983)<ref name="Starobogatov 1983">{{in lang|ru}} [[Yaroslav Starobogatov|Starobogatov Ya. I.]] (1983). "Sistema otriada Acochlidiiformes [System of the order Acochlidiiformes]." ''Vsesoiuznoe soveshchanie po izucheniiu molliuskov'', Leningrad, '''7''': 30–32.</ref> have been proposed recently, but they have not been evaluated since.<ref name="Bouchet 2005"/><ref name="Gofas 2010"/>


An alternative classification by Burn (in Beesley et al., 1998) for the Australian species recognizes 2 superfamilies and 5 families.<ref name="Bouchet 2005"/><ref name="Gofas 2010"/>
An alternative classification by Burn (in Beesley et al., 1998) for the Australian species recognizes 2 superfamilies and 5 families.<ref name="Bouchet 2005"/><ref name="Gofas 2010"/>
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Most recent morphological analyses suggested a common origin with either the equally enigmatic [[Rhodopemorpha]], the diaphanid cephalaspidean ''[[Toledonia]]'', or with runcinid or philinoid [[cephalaspidea]]ns.<ref name="Jörger 2010"/> Molecular markers independent from direct ecological pressures suggested an unresolved basal opisthobranch origin for Acochlidia (based on nuclear 18S rRNA and 28S rRNA) (Vonnemann et al. 2005).<ref name="Jörger 2010"/><ref>{{cite journal
Most recent morphological analyses suggested a common origin with either the equally enigmatic [[Rhodopemorpha]], the diaphanid cephalaspidean ''[[Toledonia]]'', or with runcinid or philinoid [[cephalaspidea]]ns.<ref name="Jörger 2010"/> Molecular markers independent from direct ecological pressures suggested an unresolved basal opisthobranch origin for Acochlidia (based on nuclear 18S rRNA and 28S rRNA) (Vonnemann et al. 2005).<ref name="Jörger 2010"/><ref>{{cite journal
| author = Vonnemann V., Schrödl M., Klussmann-Kolb A. & Wägele H.
|author1=Vonnemann V. |author2=Schrödl M. |author3=Klussmann-Kolb A. |author4=Wägele H. | year = 2005
| year = 2005
| date=
| title = Reconstruction of the phylogeny of the Opisthobranchia (Mollusca: Gastropoda) by means of 18s and 28s rRNA gene sequences
| title = Reconstruction of the phylogeny of the Opisthobranchia (Mollusca: Gastropoda) by means of 18s and 28s rRNA gene sequences
| journal = Journal of Molluscan Studies
| journal = Journal of Molluscan Studies
Line 79: Line 72:
| issn =
| issn =
| doi = 10.1093/mollus/eyi014
| doi = 10.1093/mollus/eyi014
}}</ref> A first combined multi-gene dataset led to the surprising result of Acochlidia clustering in a pulmonate relationship, united in a clade with [[Pyramidelloidea]], [[Amphiboloidea]] and [[Eupulmonata]].<ref name="Jörger 2010"/> However, only three derived acochlids were included into analysis prior to 2010, with partially missing data.<ref name="Jörger 2010"/>
| doi-access = free}}</ref> A first combined multi-gene dataset led to the surprising result of Acochlidia clustering in a pulmonate relationship, united in a clade with [[Pyramidelloidea]], [[Amphiboloidea]] and [[Eupulmonata]].<ref name="Jörger 2010"/> However, only three derived acochlids were included into analysis prior to 2010, with partially missing data.<ref name="Jörger 2010"/>


=== 2005 taxonomy ===
=== 2005 taxonomy ===
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=== 2010 taxonomy ===
=== 2010 taxonomy ===
A first comprehensive [[cladistic analysis]] of their [[phylogeny]] has been established by Schrödl & Neusser (2010),<ref name="Schrödl 2010"/> but the identity of their sister group remained uncertain. Morphology-based analyses by Schrödl & Neusser,<ref name="Schrödl 2010"/> demonstrated that Acochlidia usually group with other [[mesopsammic]] (they live in interstitial spaces of marine sands) taxa, if any were included (i.e. with the sacoglossan ''[[Platyhedyle]]'', the rhodopemorph ''[[Rhodope (gastropod)|Rhodope]]'' or the cephalaspideans ''[[Philinoglossa]]'' or ''[[Philine exigua]]'').<ref name="Jörger 2010"/> Thus, it is likely that [[convergent]] adaptations to the interstitial habitat mask the truly phylogenetic signals.<ref name="Jörger 2010"/>
A first comprehensive [[cladistic analysis]] of their [[phylogeny]] has been established by Schrödl & Neusser (2010),<ref name="Schrödl 2010"/> but the identity of their sister group remained uncertain. Morphology-based analyses by Schrödl & Neusser,<ref name="Schrödl 2010"/> demonstrated that Acochlidia usually group with other [[mesopsammic]] (they live in interstitial spaces of marine sands) taxa, if any were included (i.e. with the sacoglossan ''[[Platyhedyle]]'', the rhodopemorph ''[[Rhodope (gastropod)|Rhodope]]'' or the cephalaspideans ''[[Philinoglossa]]'' or ''[[Philine exigua]]'').<ref name="Jörger 2010"/> Thus, it is likely that [[Convergent evolution|convergent]] adaptations to the interstitial habitat mask the truly phylogenetic signals.<ref name="Jörger 2010"/>


Schrödl & Neusser (2010)<ref name="Schrödl 2010"/> split Acochlidiacea into two (unranked) taxa and into six families like this:
Schrödl & Neusser (2010)<ref name="Schrödl 2010"/> split Acochlidiacea into two (unranked) taxa and into six families like this:
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A multi-locus molecular study by Jörger et al. (2010),<ref name="Jörger 2010"/> included six out of seven acochlidian families.<ref name="Jörger 2010"/> It confirmed Acochlidiacea in a pulmonate relationship, as sister to Eupulmonata.<ref name="Jörger 2010"/> Euthyneura, Opisthobranchia and Pulmonata as traditionally defined were found non-monophyletic.<ref name="Jörger 2010"/> The enigmatic amphibious and insectivorous [[Aitengidae]] clusters within Acochlidiacea, as sister to [[meiofauna]]l and brackish Pseudunelidae and limnic Acochlidiidae.<ref name="Jörger 2010"/> Inclusion of this small acochlidian group resulted in redefinition of major groups within [[Heterobranchia]], that has led to creation of the new clades [[Euopisthobranchia]] and [[Panpulmonata]].<ref name="Jörger 2010"/>
A multi-locus molecular study by Jörger et al. (2010),<ref name="Jörger 2010"/> included six out of seven acochlidian families.<ref name="Jörger 2010"/> It confirmed Acochlidiacea in a pulmonate relationship, as sister to Eupulmonata.<ref name="Jörger 2010"/> Euthyneura, Opisthobranchia and Pulmonata as traditionally defined were found non-monophyletic.<ref name="Jörger 2010"/> The enigmatic amphibious and insectivorous [[Aitengidae]] clusters within Acochlidiacea, as sister to [[meiofauna]]l and brackish Pseudunelidae and limnic Acochlidiidae.<ref name="Jörger 2010"/> Inclusion of this small acochlidian group resulted in redefinition of major groups within [[Heterobranchia]], that has led to creation of the new clades [[Euopisthobranchia]] and [[Panpulmonata]].<ref name="Jörger 2010"/>

=== 2016 taxonomy ===
Two more families were established. Both include slugs that are unusual among acochlidians for their anatomy and habitat (amphibious, terrestrial, or benthic in deeper waters):
* [[Aitengidae]] Swennen & Buatip, 2009
* [[Bathyhedylidae]] Neusser, Jörger, Lodde-Bensch, Strong & Schrödl, 2016<ref name="Neusser 2016">Neusser T. P., Jörger K. M., Lodde-Bensch E., Strong E. E. & Schrödl M. (2016). "The unique deep sea - land connection: Interactive 3D visualization and molecular phylogeny of ''Bathyhedyle boucheti'' n. sp. (Bathyhedylidae n. fam.) - the first panpulmonate slug from bathyal zones". ''PeerJ'' '''4''': e2738. {{doi|10.7717/peerj.2738}}.</ref>


== Phylogeny ==
== Phylogeny ==
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=== Cladogram ===
=== Cladogram ===
A [[cladogram]] showing phylogenic relations of some genera and species within Acochlidiacea:<ref name="Jörger 2010"/>
A [[cladogram]] showing phylogenetic relations of some genera and species within Acochlidiacea:<ref name="Jörger 2010"/>


{{clade
{{clade
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== Overview of species ==
== Overview of species ==
[[File:Acochlidium bayerfehlmanni.png|thumb|''[[Acochlidium bayerfehlmanni]]'']]
⚫
# ''[[Hedylopsis spiculifera]]'' (Kowalevsky, 1901) (Hedylopsidae)
# ''[[Hedylopsis ballantinei]]'' Sommerfeldt & Schrödl, 2005 (Hedylopsidae)
# ''[[Hedylopsis spiculifera]]'' (Kowalevsky, 1901) (Hedylopsidae) – marine
⚫
# ''[[Hedylopsis ballantinei]]'' Sommerfeldt & Schrödl, 2005 (Hedylopsidae) – marine
# ''[[Pseudunela cornuta]]'' (Challis, 1970) (Pseudunelidae) – marine and temporary brackish<ref name="Neusser 2011"/>
# ''[[Pseudunela cornuta]]'' (Challis, 1970) (Pseudunelidae) – marine and temporary brackish<ref name="Neusser 2011"/>
# ''[[Pseudunela eirene]]'' Wawra, 1988 (Pseudunelidae) – marine
# ''[[Pseudunela eirene]]'' Wawra, 1988 (Pseudunelidae) – marine
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# ''[[Pseudunela viatoris]]'' Neusser, Jörger & Schrödl, 2011 (Pseudunelidae) – marine<ref name="Neusser 2011">Neusser T. P., Jörger K. M. & Schrödl M. (2011). "Cryptic Species in Tropic Sands - Interactive 3D Anatomy, Molecular Phylogeny and Evolution of Meiofaunal Pseudunelidae (Gastropoda, Acochlidia)". ''[[PLoS ONE]]'' '''6'''(8): e23313. {{doi|10.1371/journal.pone.0023313}}.</ref>
# ''[[Pseudunela viatoris]]'' Neusser, Jörger & Schrödl, 2011 (Pseudunelidae) – marine<ref name="Neusser 2011">Neusser T. P., Jörger K. M. & Schrödl M. (2011). "Cryptic Species in Tropic Sands - Interactive 3D Anatomy, Molecular Phylogeny and Evolution of Meiofaunal Pseudunelidae (Gastropoda, Acochlidia)". ''[[PLoS ONE]]'' '''6'''(8): e23313. {{doi|10.1371/journal.pone.0023313}}.</ref>
# ''[[Aiteng ater]]'' Swennen & Buatip, 2009 (Aitengidae) – marine (and brackish)
# ''[[Aiteng ater]]'' Swennen & Buatip, 2009 (Aitengidae) – marine (and brackish)
# ''[[Aiteng marefugitus]]'' Kano, Neusser, Fukumori, Jörger & Schrödl, 2015 (Aitengidae) – marine
# ''[[Aiteng mysticus]]'' Neusser, Fukuda, Jörger, Kano & Schrödl, 2011 (Aitengidae) – from Japan<ref name="Neusser 2011"/><ref>Neusser T. P., Fukuda H., Jörger K. M., Kano Y. & Schrödl M. (2011) "Sacoglossa or Acochlidia? 3D-reconstruction, molecular phylogeny and evolution of ''Aiteng ater'' and ''Aiteng mysticus'' n. sp. (Aitengidae, Gastropoda)". ''[[Journal of Molluscan Studies]]'' '''77'''(4): 332–350. {{doi|10.1093/mollus/eyr033}}.</ref>
# ''[[Aiteng mysticus]]'' Neusser, Fukuda, Jörger, Kano & Schrödl, 2011 (Aitengidae) – from Japan<ref name="Neusser 2011"/><ref>Neusser T. P., Fukuda H., Jörger K. M., Kano Y. & Schrödl M. (2011) "Sacoglossa or Acochlidia? 3D-reconstruction, molecular phylogeny and evolution of ''Aiteng ater'' and ''Aiteng mysticus'' n. sp. (Aitengidae, Gastropoda)". ''[[Journal of Molluscan Studies]]'' '''77'''(4): 332–350. {{doi|10.1093/mollus/eyr033}}.</ref>
# ''[[Bathyhedyle boucheti]]'' Neusser, Jörger, Lodde-Bensch, Strong & Schrödl, 2016 (Bathyhedylidae) - marine<ref name="Neusser 2016"/>
# ''[[Strubellia paradoxa]]'' (Strubell, 1892) (Acochlidiidae) – freshwater
# ''[[Strubellia paradoxa]]'' (Strubell, 1892) (Acochlidiidae) – freshwater
# ''[[Strubellia wawrai]]'' Brenzinger, Neusser, Jörger & Schrödl, 2011 (Acochlidiidae) – freshwater
# ''[[Acochlidium amboinense]]'' (Strubell, 1892) (Acochlidiidae) – freshwater
# ''[[Acochlidium amboinense]]'' (Strubell, 1892) (Acochlidiidae) – freshwater
# ''[[Acochlidium bayerfehlmanni]]'' Wawra, 1980 (Acochlidiidae) – freshwater
# ''[[Acochlidium bayerfehlmanni]]'' Wawra, 1980 (Acochlidiidae) – freshwater
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# ''[[Tantulum elegans]]'' Rankin, 1979 (Tantulidae) – freshwater
# ''[[Tantulum elegans]]'' Rankin, 1979 (Tantulidae) – freshwater
# ''[[Asperspina brambelli]]'' (Swedmark, 1968) (Asperspinidae)
# ''[[Asperspina brambelli]]'' (Swedmark, 1968) (Asperspinidae)
⚫
# ''[[Asperspina loricata]]'' (Swedmark, 1968) (Asperspinidae)
# ''[[Asperspina murmanica]]'' (Kudinskaya & Minichev, 1978) (Asperspinidae)
# ''[[Asperspina murmanica]]'' (Kudinskaya & Minichev, 1978) (Asperspinidae)
# ''[[Asperspina rhopalotecta]]'' Salvini-Plawen, 1973 (Asperspinidae)
# ''[[Asperspina rhopalotecta]]'' Salvini-Plawen, 1973 (Asperspinidae)
⚫
# ''[[Asperspina loricata]]'' (Swedmark, 1968) (Asperspinidae)
# ''[[Asperspina riseri]]'' (Morse, 1976) (Asperspinidae)
# ''[[Asperspina riseri]]'' (Morse, 1976) (Asperspinidae)
⚫
# ''[[Microhedyle gerlachi]]'' Marcus & Marcus, 1959 - synonym: ''[[Parhedyle gerlachi]]'' (Ev. Marcus & Er. Marcus, 1959) (Microhedylidae)
# ''[[Microhedyle glandulifera]]'' (Kowalevsky, 1901) (Microhedylidae)
# ''[[Microhedyle glandulifera]]'' (Kowalevsky, 1901) (Microhedylidae)
# ''[[Microhedyle nahantensis]]'' (Doe, 1974) (Microhedylidae)
# ''[[Microhedyle nahantensis]]'' (Doe, 1974) (Microhedylidae)
⚫
# ''[[Microhedyle odhneri]]'' (Ev. Marcus & Er. Marcus, 1955) (Microhedylidae)
# ''[[Microhedyle remanei]]'' (Er. Marcus, 1953) (Microhedylidae)
# ''[[Microhedyle remanei]]'' (Er. Marcus, 1953) (Microhedylidae)
# ''[[Ganitus evelinae]]'' Marcus, 1953 (Microhedylidae s.l. / Ganitidae)
# ''[[Ganitus evelinae]]'' Marcus, 1953 (Microhedylidae s.l. / Ganitidae)
# ''[[Paraganitus ellynnae]]'' Challis, 1968 (Microhedylidae s.l. / Ganitidae)
# ''[[Paraganitus ellynnae]]'' Challis, 1968 (Microhedylidae s.l. / Ganitidae)
# ''[[Parhedyle cryptophthalma]]'' (Westheide & Wawra, 1974) (Microhedylidae)
# ''[[Parhedyle cryptophthalma]]'' (Westheide & Wawra, 1974) (Microhedylidae)
⚫
# ''[[Parhedyle odhneri]]'' (Ev. Marcus & Er. Marcus, 1955) - synonym: ''[[Microhedyle odhneri]]'' (Ev. Marcus & Er. Marcus, 1955) (Microhedylidae)
# ''[[Parhedyle tyrtowii]]'' (Kowalevsky, 1900) (Microhedylidae)
# ''[[Parhedyle tyrtowii]]'' (Kowalevsky, 1900) (Microhedylidae)
⚫
# ''[[Parhedyle gerlachi]]'' (Ev. Marcus & Er. Marcus, 1959) (Microhedylidae)
# ''[[Pontohedyle brasilensis]]'' (Rankin, 1979)<ref name="Jörger 2012"/> (Microhedylidae)
# ''[[Pontohedyle brasilensis]]'' (Rankin, 1979)<ref name="Jörger 2012"/> (Microhedylidae)
# ''[[Pontohedyle joni]]'' Jörger & Schrödl, 2013
# ''[[Pontohedyle kepii]]'' Jörger & Schrödl, 2013
# ''[[Pontohedyle liliae]]'' Jörger & Schrödl, 2013
# ''[[Pontohedyle martynovi]]'' Jörger & Schrödl, 2013
# ''[[Pontohedyle milaschewitchii]]'' (Kowalevsky, 1901) (Microhedylidae)
# ''[[Pontohedyle milaschewitchii]]'' (Kowalevsky, 1901) (Microhedylidae)
# ''[[Pontohedyle neridae]]'' Jörger & Schrödl, 2013
# ''[[Pontohedyle peteryalli]]'' Jörger & Schrödl, 2013
# ''[[Pontohedyle verrucosa]]'' (Challis, 1970) (Microhedylidae)
# ''[[Pontohedyle verrucosa]]'' (Challis, 1970) (Microhedylidae)
# ''Pontohedyle'' sp. 1 - ''Pontohedyle'' sp. 9<ref name="Jörger 2012"/>
# ''[[Pontohedyle wenzli]]'' Jörger & Schrödl, 2013
# ''[[Pontohedyle wiggi]]'' Jörger & Schrödl, 2013
# ''[[Pontohedyle yurihookeri]]'' Jörger & Schrödl, 2013
# ''[[Helicohedyle dikiki]]'' Drainas, Carlson, Jörger, Schrödl & Neusser, 2017 (unassigned to family) – marine


==References==
==References==
This article incorporates CC-BY-2.0 text from references<ref name="Jörger 2010">Jörger K. M., Stöger I., Kano Y., Fukuda H., Knebelsberger T. & Schrödl M. (2010). "On the origin of Acochlidia and other enigmatic euthyneuran gastropods, with implications for the systematics of Heterobranchia". ''[[BMC Evolutionary Biology]]'' '''10''': 323. {{doi|10.1186/1471-2148-10-323}}.</ref><ref name="Jörger 2012">Jörger K. M., Norenburg J. L., Wilson N. G. & Schrödl M. (2012). "Barcoding against a paradox? Combined molecular species delineations reveal multiple cryptic lineages in elusive meiofaunal sea slugs". ''[[BMC Evolutionary Biology]]'' '''12''': 245. {{doi|10.1186/1471-2148-12-245}}.</ref> and CC-BY-SA-3.0 text from the reference.<ref name="Gofas 2010">Gofas, S. (2010). Acochlidiacea. In: Bouchet, P.; Gofas, S.; Rosenberg, G. (2010) World Marine Mollusca database. Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetails&id=411937 on 2010-11-30</ref>
This article incorporates CC-BY-2.0 text from references<ref name="Jörger 2010">Jörger K. M., Stöger I., Kano Y., Fukuda H., Knebelsberger T. & Schrödl M. (2010). "On the origin of Acochlidia and other enigmatic euthyneuran gastropods, with implications for the systematics of Heterobranchia". ''[[BMC Evolutionary Biology]]'' '''10''': 323. {{doi|10.1186/1471-2148-10-323}}.</ref><ref name="Jörger 2012">Jörger K. M., Norenburg J. L., Wilson N. G. & Schrödl M. (2012). "Barcoding against a paradox? Combined molecular species delineations reveal multiple cryptic lineages in elusive meiofaunal sea slugs". ''[[BMC Evolutionary Biology]]'' '''12''': 245. {{doi|10.1186/1471-2148-12-245}}.</ref> and CC-BY-SA-3.0 text from the reference.<ref name="Gofas 2010">Gofas, S. (2010). Acochlidiacea. In: Bouchet, P.; Gofas, S.; Rosenberg, G. (2010) World Marine Mollusca database. Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetails&id=411937 on 2010-11-30</ref>
{{reflist}}
{{Reflist}}


== External links ==
== External links ==
{{commons category|Acochlidiacea}}
{{Commons category|Acochlidiacea}}
{{wikispecies|Acochlidiacea}}
{{Wikispecies|Acochlidiacea}}
* [http://www.seaslugforum.net/factsheet.cfm?base=acochlidea Acochlidea on Sea Slug Forum]
* [https://web.archive.org/web/20100614230258/http://seaslugforum.net/factsheet.cfm?base=acochlidea Acochlidea on Sea Slug Forum]

{{Taxonbar|from=Q1054947}}


[[Category:Panpulmonata]]
[[Category:Panpulmonata]]

Latest revision as of 01:43, 5 September 2024

Acochlidiacea
Acochlidium fijiiensis
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
Subclass: Heterobranchia
Infraclass: Euthyneura
Superorder: Acochlidiacea
Odhner, 1937[1]
Families

(unranked) Hedylopsacea

(unranked) Microhedylacea

Diversity
46 species
Synonyms[2][3]

Acochlidiomorpha
Acochlidiida
Acochlidea
Acochlidia

Acochlidiacea, common name acochlidians, are a taxonomic clade of very unusual sea snails and sea and freshwater slugs, aquatic gastropod mollusks within the large clade Heterobranchia. Acochlidia is a variant spelling.

Schematic drawing of dorsal view of Pseudunela cornuta shows some characteristics of acochlidians, but characteristics of other acochlidians differ greatly:
f – foot
hb – heart bulb
lt – labial tentacle
rh – rhinophore
vh – visceral hump.

Description

[edit]

These are mostly very small animals, without a shell or gills, distinguished by the visceral mass being sharply set off from the rest of the body.[4]

Being a small group with only 47 species worldwide known in 2023,[5] this group has been the subject of active research since 2010, by which date only 32 species were named.[6] These slugs are morphologically and biologically highly aberrant and diverse, comprising a series of unusual characters (e.g. secondary gonochorism, lack of copulatory organs, asymmetric radulae).[7] Most acochlidians live interstitially in marine sands, while some have conquered limnic systems (uniquely within opisthobranch gastropods).[7]

Taxonomy

[edit]

Nils Hjalmar Odhner established this taxon as a family in 1937, when he created the families Microhedylidae and Acochlidiidae.[1][2] In 1939, he treated this taxon as an order.[2][8]

Rankin (1979)[9] treated this taxon as an order, the order Acochlidioidea.[2][3]

Salvini-Plawen (1983)[10] wrote this taxon as Acochlidiomorpha.[3]

Anderson (1992)[11] treated this taxon as the order Acochlidiida.[2][3]

Burn in Beesley et al. (1998), wrote this taxon as the order Acochlidea.[3]

Wawra (1987)[12] and various authors (2007–2010)[7][13][14][15] spelled this taxon as Acochlidia.

Three families (Hedylopsidae, Microhedylidae and Acochlidiidae) are classically recognized.[2][3] Two controversial classifications (Rankin 1979,[9] Starobogatov 1983)[16] have been proposed recently, but they have not been evaluated since.[2][3]

An alternative classification by Burn (in Beesley et al., 1998) for the Australian species recognizes 2 superfamilies and 5 families.[2][3]

3D reconstructions like this one (Pseudunela cornuta) highly improved knowledge about acochlidians.

The Acochlidia, a traditional "order" of the Opisthobranchia since their establishment by Odhner have formed one of the unsolved mysteries within Euthyneura.[7] Their monophyly is widely accepted especially since a proposed sister group relationship of the acochlidian family Ganitidae with Sacoglossa (based on the dagger-shaped radula teeth) could be rejected based on a comprehensive parsimony analysis of morphological characters.[7] During the last years a series of studies have redescribed key acochlidian taxa in great detail, including 3D reconstructions, and added considerably to the morphological and biological knowledge of this previously little understood group.[7]

Most recent morphological analyses suggested a common origin with either the equally enigmatic Rhodopemorpha, the diaphanid cephalaspidean Toledonia, or with runcinid or philinoid cephalaspideans.[7] Molecular markers independent from direct ecological pressures suggested an unresolved basal opisthobranch origin for Acochlidia (based on nuclear 18S rRNA and 28S rRNA) (Vonnemann et al. 2005).[7][17] A first combined multi-gene dataset led to the surprising result of Acochlidia clustering in a pulmonate relationship, united in a clade with Pyramidelloidea, Amphiboloidea and Eupulmonata.[7] However, only three derived acochlids were included into analysis prior to 2010, with partially missing data.[7]

2005 taxonomy

[edit]

The taxonomy of Bouchet & Rocroi (2005)[2] tentatively follows Starobogatov (1983),[16] but they have downgraded his taxonomic ranks (suborders to superfamilies, superfamilies to families).[2][3] The group Acochlidiacea is arranged as follows:

2010 taxonomy

[edit]

A first comprehensive cladistic analysis of their phylogeny has been established by Schrödl & Neusser (2010),[15] but the identity of their sister group remained uncertain. Morphology-based analyses by Schrödl & Neusser,[15] demonstrated that Acochlidia usually group with other mesopsammic (they live in interstitial spaces of marine sands) taxa, if any were included (i.e. with the sacoglossan Platyhedyle, the rhodopemorph Rhodope or the cephalaspideans Philinoglossa or Philine exigua).[7] Thus, it is likely that convergent adaptations to the interstitial habitat mask the truly phylogenetic signals.[7]

Schrödl & Neusser (2010)[15] split Acochlidiacea into two (unranked) taxa and into six families like this:

Hedylopsacea

[edit]

Hedylopsacean Acochlidiacea, whose evolution involves several habitat shifts from marine interstitial to amphibious or freshwater benthic habitats, possess complex excretory and reproductive systems.[18]

(unranked) Hedylopsacea has no superfamilies defined:[15]

Microhedylacea

[edit]

Microhedylacean Acochlidiacea are exclusively found in interstitial spaces in sediment, and show a tendency toward reduction of complexity in major organ systems.[18]

(unranked) Microhedylacea has no superfamilies defined:[15]

A multi-locus molecular study by Jörger et al. (2010),[7] included six out of seven acochlidian families.[7] It confirmed Acochlidiacea in a pulmonate relationship, as sister to Eupulmonata.[7] Euthyneura, Opisthobranchia and Pulmonata as traditionally defined were found non-monophyletic.[7] The enigmatic amphibious and insectivorous Aitengidae clusters within Acochlidiacea, as sister to meiofaunal and brackish Pseudunelidae and limnic Acochlidiidae.[7] Inclusion of this small acochlidian group resulted in redefinition of major groups within Heterobranchia, that has led to creation of the new clades Euopisthobranchia and Panpulmonata.[7]

2016 taxonomy

[edit]

Two more families were established. Both include slugs that are unusual among acochlidians for their anatomy and habitat (amphibious, terrestrial, or benthic in deeper waters):

Phylogeny

[edit]

There is no fossil record of Acochlidiacea. Application of a molecular clock allowed estimation of divergence times for these groups. The split between Eupulmonata and Acochlidiacea took place in the Mesozoic, between the Triassic and Jurassic periods. The diversification of Acochlidia is estimated to have happened in the Jurassic with the split between Hedylopsacea and Microhedylacea.[7]

Cladogram

[edit]

A cladogram showing phylogenetic relations of some genera and species within Acochlidiacea:[7]

Acochlidiacea

Ecology

[edit]

The life cycle of Acochlidiacea is poorly known. With a typically low reproductive output in Acochlidiacea (max. of 40 eggs in Pontohedyle milaschewitchii), free veliger larvae are assumed to stay in the interstices of the sand grains rather than entering the water column thereby avoiding long distance dispersal.[18] Fertilized eggs are attached to sand grains and might promote dispersal via current driven sediment transport along shorelines.[18]

Overview of species

[edit]
Acochlidium bayerfehlmanni
  1. Hedylopsis spiculifera (Kowalevsky, 1901) (Hedylopsidae) – marine
  2. Hedylopsis ballantinei Sommerfeldt & Schrödl, 2005 (Hedylopsidae) – marine
  3. Pseudunela cornuta (Challis, 1970) (Pseudunelidae) – marine and temporary brackish[6]
  4. Pseudunela eirene Wawra, 1988 (Pseudunelidae) – marine
  5. Pseudunela espiritusanta Neusser & Schrödl, 2009 (Pseudunelidae) – in brackish water
  6. Pseudunela marteli Neusser, Jörger & Schrödl, 2011 (Pseudunelidae) – marine[6]
  7. Pseudunela viatoris Neusser, Jörger & Schrödl, 2011 (Pseudunelidae) – marine[6]
  8. Aiteng ater Swennen & Buatip, 2009 (Aitengidae) – marine (and brackish)
  9. Aiteng marefugitus Kano, Neusser, Fukumori, Jörger & Schrödl, 2015 (Aitengidae) – marine
  10. Aiteng mysticus Neusser, Fukuda, Jörger, Kano & Schrödl, 2011 (Aitengidae) – from Japan[6][20]
  11. Bathyhedyle boucheti Neusser, Jörger, Lodde-Bensch, Strong & Schrödl, 2016 (Bathyhedylidae) - marine[19]
  12. Strubellia paradoxa (Strubell, 1892) (Acochlidiidae) – freshwater
  13. Strubellia wawrai Brenzinger, Neusser, Jörger & Schrödl, 2011 (Acochlidiidae) – freshwater
  14. Acochlidium amboinense (Strubell, 1892) (Acochlidiidae) – freshwater
  15. Acochlidium bayerfehlmanni Wawra, 1980 (Acochlidiidae) – freshwater
  16. Acochlidium fijiiensis Haynes & Kenchington, 1991[21] (Acochlidiidae) – freshwater
  17. Palliohedyle sutteri (Wawra, 1979) (Acochlidiidae) – freshwater
  18. Palliohedyle weberi (Bergh, 1895) (Acochlidiidae) – in brackish waters
  19. Tantulum elegans Rankin, 1979 (Tantulidae) – freshwater
  20. Asperspina brambelli (Swedmark, 1968) (Asperspinidae)
  21. Asperspina loricata (Swedmark, 1968) (Asperspinidae)
  22. Asperspina murmanica (Kudinskaya & Minichev, 1978) (Asperspinidae)
  23. Asperspina rhopalotecta Salvini-Plawen, 1973 (Asperspinidae)
  24. Asperspina riseri (Morse, 1976) (Asperspinidae)
  25. Microhedyle gerlachi Marcus & Marcus, 1959 - synonym: Parhedyle gerlachi (Ev. Marcus & Er. Marcus, 1959) (Microhedylidae)
  26. Microhedyle glandulifera (Kowalevsky, 1901) (Microhedylidae)
  27. Microhedyle nahantensis (Doe, 1974) (Microhedylidae)
  28. Microhedyle remanei (Er. Marcus, 1953) (Microhedylidae)
  29. Ganitus evelinae Marcus, 1953 (Microhedylidae s.l. / Ganitidae)
  30. Paraganitus ellynnae Challis, 1968 (Microhedylidae s.l. / Ganitidae)
  31. Parhedyle cryptophthalma (Westheide & Wawra, 1974) (Microhedylidae)
  32. Parhedyle odhneri (Ev. Marcus & Er. Marcus, 1955) - synonym: Microhedyle odhneri (Ev. Marcus & Er. Marcus, 1955) (Microhedylidae)
  33. Parhedyle tyrtowii (Kowalevsky, 1900) (Microhedylidae)
  34. Pontohedyle brasilensis (Rankin, 1979)[18] (Microhedylidae)
  35. Pontohedyle joni Jörger & Schrödl, 2013
  36. Pontohedyle kepii Jörger & Schrödl, 2013
  37. Pontohedyle liliae Jörger & Schrödl, 2013
  38. Pontohedyle martynovi Jörger & Schrödl, 2013
  39. Pontohedyle milaschewitchii (Kowalevsky, 1901) (Microhedylidae)
  40. Pontohedyle neridae Jörger & Schrödl, 2013
  41. Pontohedyle peteryalli Jörger & Schrödl, 2013
  42. Pontohedyle verrucosa (Challis, 1970) (Microhedylidae)
  43. Pontohedyle wenzli Jörger & Schrödl, 2013
  44. Pontohedyle wiggi Jörger & Schrödl, 2013
  45. Pontohedyle yurihookeri Jörger & Schrödl, 2013
  46. Helicohedyle dikiki Drainas, Carlson, Jörger, Schrödl & Neusser, 2017 (unassigned to family) – marine

References

[edit]

This article incorporates CC-BY-2.0 text from references[7][18] and CC-BY-SA-3.0 text from the reference.[3]

  1. ^ a b (in German) Odhner N. H. (1937). "Hedylopsis suecica n.sp. und die Nacktschneckengruppe Acochlidiacea (Hedylacea)". Zoologischer Anzeiger 120(3–4): 51–64. 52, 62.
  2. ^ a b c d e f g h i j Bouchet, Philippe; Rocroi, Jean-Pierre; Frýda, Jiri; Hausdorf, Bernard; Ponder, Winston; Valdés, Ángel & Warén, Anders (2005). "Classification and nomenclator of gastropod families". Malacologia. 47 (1–2). Hackenheim, Germany: ConchBooks: 1–397. ISBN 3-925919-72-4. ISSN 0076-2997.
  3. ^ a b c d e f g h i j Gofas, S. (2010). Acochlidiacea. In: Bouchet, P.; Gofas, S.; Rosenberg, G. (2010) World Marine Mollusca database. Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetails&id=411937 on 2010-11-30
  4. ^ Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. p. 376. ISBN 0-03-056747-5.
  5. ^ MolluscaBase eds. (2021). MolluscaBase. Acochlidiimorpha. Accessed on 2023-02-13.
  6. ^ a b c d e Neusser T. P., Jörger K. M. & Schrödl M. (2011). "Cryptic Species in Tropic Sands - Interactive 3D Anatomy, Molecular Phylogeny and Evolution of Meiofaunal Pseudunelidae (Gastropoda, Acochlidia)". PLoS ONE 6(8): e23313. doi:10.1371/journal.pone.0023313.
  7. ^ a b c d e f g h i j k l m n o p q r s t u Jörger K. M., Stöger I., Kano Y., Fukuda H., Knebelsberger T. & Schrödl M. (2010). "On the origin of Acochlidia and other enigmatic euthyneuran gastropods, with implications for the systematics of Heterobranchia". BMC Evolutionary Biology 10: 323. doi:10.1186/1471-2148-10-323.
  8. ^ Odhner N. H. (1939). "Opisthobranchiate Mollusca from the western and northern coasts of Norway". Det Kongelige Norske Videnskabers Selskabs Skrifter 1939(1): 1–92. page 5.
  9. ^ a b Rankin J. J. (1979). "A freshwater shell-less mollusc from the Caribbean: structure, biotics, and contribution to a new understanding of the Acochlidioidea". Royal Ontario Museum, Life Sciences Contributions 116: 123 pp., page 83.
  10. ^ Salvini-Plawen L. v. (1983). Mollusca. In: Fauna & Flora des Mittelmeeres. Riedl R. (ed.) P. Parey. Hamburg & Berlin, 248–390. page 309.
  11. ^ Anderson C. (1992). Classification of organisms living and fossil. Golden Crown Press, Lancaster, Ohio, 69 pp., page 37.
  12. ^ Wawra E. (1987). "Zur Anatomie einiger Acochlidia (Gastropoda, Opisthobranchia) mit einer vorläufigen Revision des Systems und einem Anhang über Platyhedylidae (Opisthobranchia, Ascoglossa)". PhD thesis. Universität Wien.
  13. ^ Neusser T. P., Jörger K. M. & Schrödl M. (2007). "Exploring cerebral features in Acochlidia (Gastropoda: Opisthobranchia)". Bonn. Zool. Beitr. 55: 301–310.
  14. ^ Jörger K. M., Heβ M., Neusser T. P. & Schrödl M. (2009). "Sex in the beach: spermatophores, dermal insemination and 3D sperm ultrastructure of the aphallic mesopsammic Pontohedyle milaschewitchii (Acochlidia, Opisthobranchia, Gastropoda)". Marine Biology 156: 1159–1170. doi:10.1007/s00227-009-1158-5.
  15. ^ a b c d e f g Schrödl M. & Neusser T. P. (2010). "Towards a phylogeny and evolution of Acochlidia (Mollusca: Gastropoda: Opisthobranchia)". Zoological Journal of the Linnean Society 158: 124–154. doi:10.1111/j.1096-3642.2009.00544.x.
  16. ^ a b (in Russian) Starobogatov Ya. I. (1983). "Sistema otriada Acochlidiiformes [System of the order Acochlidiiformes]." Vsesoiuznoe soveshchanie po izucheniiu molliuskov, Leningrad, 7: 30–32.
  17. ^ Vonnemann V.; Schrödl M.; Klussmann-Kolb A.; Wägele H. (2005). "Reconstruction of the phylogeny of the Opisthobranchia (Mollusca: Gastropoda) by means of 18s and 28s rRNA gene sequences". Journal of Molluscan Studies. 71 (2): 113–125. doi:10.1093/mollus/eyi014.
  18. ^ a b c d e f Jörger K. M., Norenburg J. L., Wilson N. G. & Schrödl M. (2012). "Barcoding against a paradox? Combined molecular species delineations reveal multiple cryptic lineages in elusive meiofaunal sea slugs". BMC Evolutionary Biology 12: 245. doi:10.1186/1471-2148-12-245.
  19. ^ a b Neusser T. P., Jörger K. M., Lodde-Bensch E., Strong E. E. & Schrödl M. (2016). "The unique deep sea - land connection: Interactive 3D visualization and molecular phylogeny of Bathyhedyle boucheti n. sp. (Bathyhedylidae n. fam.) - the first panpulmonate slug from bathyal zones". PeerJ 4: e2738. doi:10.7717/peerj.2738.
  20. ^ Neusser T. P., Fukuda H., Jörger K. M., Kano Y. & Schrödl M. (2011) "Sacoglossa or Acochlidia? 3D-reconstruction, molecular phylogeny and evolution of Aiteng ater and Aiteng mysticus n. sp. (Aitengidae, Gastropoda)". Journal of Molluscan Studies 77(4): 332–350. doi:10.1093/mollus/eyr033.
  21. ^ Haynes A. & Kenchington W. (1991). "Acochlidium fijiiensis sp. nov. (Gastropoda: Opisthobranchia: Acochlidiacea) from Fiji". The Veliger 34(2): 166–171.
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