{{Short description|Clade of gastropods}} {{Automatic taxobox | taxon = Pneumopulmonata | authority = Krug et al., 2022 }}
'''Pneumopulmonata''' is a superorder of heterobranch pulmonate gastropods belonging to the subterclass Tectipleura. It was defined based on results of phylogenomic studies<ref>{{Cite journal |last1=Krug |first1=Patrick J. |last2=Caplins |first2=Serena A. |last3=Algoso |first3=Krisha |last4=Thomas |first4=Kanique |last5=Valdés |first5=Ángel A. |last6=Wade |first6=Rachael |last7=Wong |first7=Nur Leena W. S. |last8=Eernisse |first8=Douglas J. |last9=Kocot |first9=Kevin M. |date=2022-04-13 |title=Phylogenomic resolution of the root of Panpulmonata, a hyperdiverse radiation of gastropods: new insight into the evolution of air breathing |journal=Proceedings of the Royal Society B: Biological Sciences |language=en |volume=289 |issue=1972 |article-number=20211855 |doi=10.1098/rspb.2021.1855 |issn=0962-8452 |pmc=8984808 |pmid=35382597}}</ref><ref>{{Cite book |last=Teasdale |first=Luisa Cinzia |title=Phylogenomics of the pulmonate land snails. [Dissertation submitted in total fulfilment of the requirements of the degree of doctor of Philosophy] |date=2017 |publisher=School of BioSciences, The University of Melbourne |location=Melbourne}}</ref> and named by Krug et al. (2022). The superorder unites all of Panpulmonata except for Sacoglossa.
==Description== Synapomorphy of the group is the presence of a pneumostome. It as established for the taxa comprising all descendants of the last common ancestor shared by Siphonarioidea, Amphiboloidea and Stylommatophora.<ref name="WoRMS"> MolluscaBase eds. (2024). MolluscaBase. Pneumopulmonata. Accessed through: World Register of Marine Species at: https://www.marinespecies.org/aphia.php?p=taxdetails&id=1586068 on 2024-10-26</ref> It mainly contains clades occurring at the interface between sea and land, in freshwater, and in terrestrial habitats.
The following groups are included:
* Siphonarioidea * Acochlidiacea * Pylopulmonata ** Amphiboloidea ** Pyramidelloidea ** Glacidorboidea * Hygrophila * Eupulmonata ** Ellobiida ** Systellommatophora ** Stylommatophora
The clade largely overlaps with the former Pulmonata, but includes additional lineages as Pyramidelloidea. It is a sister group of Sacoglossa within Panpulmonata. Repeated colonizations of the freshwater and terrestrial realms occurred in this clade, and the biology and anatomy of its members is important for understanding the evolution of the highly diverse stylommatophoran land snails.
The individual subgroups differ in the digree of their terrestrialization as well as in the anatomical similarity to stylommatohora, sometimes reversing evolutionary trends seen in related groups:
* Siphonarioidea include a single family of marine limpet-like intertidal (but see the subtidal ''Williamia'') gastropods. They have a large pallial cavity containing a secondary gill on a large part of its roof and opening by a non-contractible pneumostome on the right side of the body. During the high tide the cavity fills with water, at low tide it is used a lung. The pneumostome lacks a sphincter. Anus is located at the pneumostome. There is no operculum, tentacles are absent. Radula has many small and rather uniform teeth in each row. The genital system has a common spermoviduct ending in a copulatory organ and a single genital opening. Sperm is transferred in a spermatophore. Development is through a free-swimming veliger larva.<ref name=":3">{{Citation |last=Heller |first=Joseph |chapter=Marine Ancestors of most Land Snails: Pulmonates |date=2015 |title=Sea Snails |pages=257–264 |chapter-url=https://link.springer.com/10.1007/978-3-319-15452-7_10 |access-date=2024-10-24 |place=Cham |publisher=Springer International Publishing |language=en |doi=10.1007/978-3-319-15452-7_10 |isbn=978-3-319-15451-0|chapter-url-access=subscription }}</ref><ref>{{Cite journal |last1=Simone |first1=Luiz Ricardo L. |last2=Seabra |first2=Maria Inês G. L. |date=2017-09-01 |title=Shell and body structure of the plesiomorphic pulmonate marine limpet Siphonaria pectinata (Linnaeus, 1758) from Portugal (Gastropoda: Heterobranchia: Siphonariidae) |url=https://www.foliamalacologica.com/Shell-and-body-structure-of-the-plesiomorphic-pulmonate-marine-limpet-Siphonaria,118015,0,2.html |journal=Folia Malacologica |volume=25 |issue=3 |pages=147–164 |doi=10.12657/folmal.025.012|doi-access=free }}</ref><ref name=":1">{{Cite book |last=Hubendick |first=B. |title=Pulmonates. Volume 2A. Systematics, Evolution and Ecology |publisher=Academic Press |year=1978 |isbn=0-12-267502-9 |editor-last=Fretter1 Peake2 |editor-first=Vera1 J.2 |location=London, New York, San Francisco |pages=1–47 |chapter=Systematics and comparative morphology of the Basommatophora}}</ref> * Acochlidiacea is a group of shell-less gastropods of tiny size, mostly with distinctive dorsal visceral hump. They are mostly marine, inhabiting the interstitial of coastal sands, but there are also freshwater species. Pallial cavity is small or missing. Anus is located on the right side, may be moved to the visceral hump in some species. Operculum is missing. Two pairs of tentacles (rhinophores and labial tentacles) are present, but do not bear eyes. Radula is narrow, with as few as just one tooth per row, in some species asymmetric. Some are secondarily gonochoric, otherwise there is a common spemoviduct either ending with a single opening, connected to a copulatory organ/penial sheath by an external groove, or there is vas deferens and a separate male opening. At least some species use spermatophores for sperm transfer. Some species have veliger larvae.<ref>{{Cite journal |last1=Jörger |first1=Katharina M |last2=Stöger |first2=Isabella |last3=Kano |first3=Yasunori |last4=Fukuda |first4=Hiroshi |last5=Knebelsberger |first5=Thomas |last6=Schrödl |first6=Michael |date=2010 |title=On the origin of Acochlidia and other enigmatic euthyneuran gastropods, with implications for the systematics of Heterobranchia |journal=BMC Evolutionary Biology |language=en |volume=10 |issue=1 |pages=323 |doi=10.1186/1471-2148-10-323 |doi-access=free |issn=1471-2148 |pmc=3087543 |pmid=20973994|bibcode=2010BMCEE..10..323J }}</ref><ref>{{Cite journal |last1=Schrödl |first1=Michael |last2=Neusser |first2=Timea P. |date=2010 |title=Towards a phylogeny and evolution of Acochlidia (Mollusca: Gastropoda: Opisthobranchia) |url=https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2009.00544.x |journal=Zoological Journal of the Linnean Society |language=en |volume=158 |issue=1 |pages=124–154 |doi=10.1111/j.1096-3642.2009.00544.x}}</ref><ref>{{Cite journal |last1=Neusser |first1=Timea P |last2=Heß |first2=Martin |last3=Schrödl |first3=Michael |date=2009 |title=Tiny but complex - interactive 3D visualization of the interstitial acochlidian gastropod Pseudunela cornuta (Challis, 1970) |journal=Frontiers in Zoology |language=en |volume=6 |issue=1 |pages=20 |doi=10.1186/1742-9994-6-20 |doi-access=free |issn=1742-9994 |pmc=2761907 |pmid=19747373}}</ref><ref>{{Cite journal |last1=Brenzinger |first1=B. |last2=Neusser |first2=T.P. |last3=Glaubrecht |first3=M. |last4=Haszprunar |first4=G. |last5=Schrödl |first5=M. |date=2010-12-20 |title=Redescription and three-dimensional reconstruction of the limnic acochlidian gastropod Strubellia paradoxa (Strubell, 1892) (Gastropoda: Euthyneura) from Ambon, Indonesia |url=https://www.tandfonline.com/doi/full/10.1080/00222933.2010.521862 |journal=Journal of Natural History |language=en |volume=45 |issue=3–4 |pages=183–209 |doi=10.1080/00222933.2010.521862 |bibcode=2010JNatH..45..183B |issn=0022-2933|url-access=subscription }}</ref> * Amphiboloidea occur in on intertidal mud flats in marine or brackish water and are active during the low tide. The pallial cavity is used for breathing air when the snail is exposed, but also for breathing in water.<ref>{{Cite journal |last1=Pilkington |first1=J. B. |last2=Little |first2=C. |last3=Stirling |first3=P. E. |date=1984 |title=A respiratory current in the mantle cavity of Amphibola crenata (Mollusca, Pulmonata). |url=http://www.tandfonline.com/doi/abs/10.1080/03036758.1984.10421734 |journal=Journal of the Royal Society of New Zealand |language=en |volume=14 |issue=4 |pages=327–334 |doi=10.1080/03036758.1984.10421734 |issn=0303-6758|url-access=subscription }}</ref> There is no contractile pneumostome. Operculum is developed. The is one pair of tentacles, eyes are located near their base. Radula has many similar teeth per row. The structure of the genital system varies: there is a single genital opening, but there may be a common spermoviduct (''Salinator'') as well as separated oviduct and vas deferens (''Maningrida'').<ref>{{Cite journal |last1=Golding |first1=Rosemary E. |last2=Byrne |first2=Maria |last3=Ponder |first3=Winston F. |date=2008 |title=Novel copulatory structures and reproductive functions in Amphiboloidea (Gastropoda, Heterobranchia, Pulmonata) |url=https://onlinelibrary.wiley.com/doi/10.1111/j.1744-7410.2007.00120.x |journal=Invertebrate Biology |language=en |volume=127 |issue=2 |pages=168–180 |doi=10.1111/j.1744-7410.2007.00120.x |issn=1077-8306|url-access=subscription }}</ref> There is a free-swimming veliger larva.<ref name=":3" /><ref name=":1" /> * Pyramidelloidea is a group of marine snails, ectoparasitic on bivalves and annelids. They occur from the subtidal to depths of at least 300 m. They do not have a pneumostome and the pallial cavity contains a well developed gill. Operculum is present. Head bears a pair of tentacles, eyes are positioned medially on the head. Radula is lost and there is a long sucking proboscis. The genital system has a spermoviduct leading to a single common opening, but there is an external ciliated groove leading to vas deferes, penis, and a separate male genital opening. Development is through a free-swimming veliger larva.<ref>{{Cite journal |last=Ponder |first=W. F. |date=1987 |title=The anatomy and relationships of the pyramidellacean limpet Amathina tricarinata (Mollusca: Gastropoda) |journal=Asian Marine Biology |volume=4 |pages=1–34}}</ref><ref>{{Cite journal |last1=Fretter |first1=V. |last2=Graham |first2=A. |date=1949 |title=The structure and mode of life of the Pyramidellidae, parasitic opistobranchs |url=https://plymsea.ac.uk/id/eprint/1371/1/The_structure_and_mode_of_life_of_the_pyramidellidae,_parasitic_opisthobranchs.pdf |journal=Journal of the Marine Biological Association of the United Kingdom |volume=28 |issue=2 |pages=493–532 |doi=10.1017/S0025315400023377 }}</ref><ref>{{Cite journal |last=Robertson |first=Robert |date=2012 |title=Pyramidellid Protoconchs, Eggs, Embryos and Larval Ecology: An Introductory Survey |url=https://www.bioone.org/doi/abs/10.4003/006.030.0201 |journal=American Malacological Bulletin |language=en |volume=30 |issue=2 |pages=219–228 |doi=10.4003/006.030.0201 |issn=0740-2783|url-access=subscription }}</ref><ref>{{Cite journal |last=Høisæter |first=Tore |date=2014-12-19 |title=The Pyramidellidae (Gastropoda, Heterobranchia) of Norway and adjacent waters. A taxonomic review |url=https://www.ntnu.no/ojs/index.php/fauna_norvegica/article/view/1672 |journal=Fauna Norvegica |volume=34 |pages=7–78 |doi=10.5324/fn.v34i0.1672 |issn=1891-5396|hdl=1956/10213 |hdl-access=free }}</ref> * Glacidorboidea are tiny freshwater snails. The pallial cavity is widely open and filled with water; there is no gill. Anus and the genital opening are within the pallial cavity. Operculum is present. There is one pair of tentacles with eyes positioned medially on the head. Predatory, radula with large median teeth and much reduced lateral teeth. Direct development.<ref>{{Cite journal |last1=Ponder |first1=Winston F. |last2=Avern |first2=G. J. |date=2000 |title=The Glacidorbidae (Mollusca: Gastropoda: Heterobranchia) of Australia |url=https://media.australian.museum/media/Uploads/Journals/17898/1318_complete.pdf |journal=Records of the Australian Museum |volume=52 |issue=3 |pages=307–353 |doi=10.3853/j.0067-1975.52.2000.1318 }}</ref> * Hygrophila is a clade of exclusively freshwater snails and limpets. They mostly breath oxygen from the air and regularly come to the surface to refill their pallial cavity with fresh air (this may not hold during the winter), but some live permanently submerged; secondary breathing surfaces for absorbing oxygen from water may be present. The basal Chilinidae (some species) and Latiidae breathe with their pallial cavity oxygen dissolved in the water. Pneumostome is contractible (but not in Chilinidae). Operculum is absent. There is one pair of tentacles that does not bear eyes. Radula has many small, similar teeth per row. In the genital system, the male and female tracts separate after a short spermoviduct (Chilinidae, Latiidae) or there is no common spermoviduct at all. Development is direct.<ref>{{Cite journal |last=Brace |first=R. C. |date=1983 |title=Observations on the Morphology and Behaviour of Chilina fluctuosa Gray (Chilinidae), with a Discussion on the Early Evolution of Pulmonate Gastropods |url=https://www.jstor.org/stable/2396031 |journal=Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences |volume=300 |issue=1101 |pages=463–491 |doi=10.1098/rstb.1983.0017 |jstor=2396031 |issn=0080-4622}}</ref><ref>{{Cite journal |last=Simone |first=Luiz Ricardo L. |date=2021-03-25 |title=Freshwater micro-gastropods from the Iguaçu National Park, Brazil: two new truncatelloid caenogastropods and anatomy of Chilina megastoma Hylton Scott, 1958 |url=https://www.foliamalacologica.com/Freshwater-micro-gastropods-from-the-Iguacu-National-Park-Brazil-two-new-truncatelloid,133953,0,2.html |journal=Folia Malacologica |volume=29 |issue=1 |pages=13–32 |doi=10.12657/folmal.029.002|doi-access=free }}</ref><ref>{{Cite journal |last=Cheatum |first=Elmer Philip |date=1934 |title=Limnological Investigations on Respiration, Annual Migratory Cycle, and Other Related Phenomena in Fresh-Water Pulmonate Snails |url=https://www.jstor.org/stable/3222383 |journal=Transactions of the American Microscopical Society |volume=53 |issue=4 |pages=348–407 |doi=10.2307/3222383 |jstor=3222383 |issn=0003-0023|url-access=subscription }}</ref><ref>{{Citation |last1=Cuezzo |first1=María Gabriela |chapter=Phylum Mollusca |date=2020 |title=Thorp and Covich's Freshwater Invertebrates |pages=261–430 |url=https://linkinghub.elsevier.com/retrieve/pii/B9780128042250000113 |access-date=2024-10-27 |publisher=Elsevier |language=en |doi=10.1016/b978-0-12-804225-0.00011-3 |isbn=978-0-12-804225-0 |last2=Gutiérrez Gregoric |first2=Diego Eduardo |last3=Pointier |first3=Jean-Pierre |last4=Vázquez |first4=Antonio Alejandro |last5=Ituarte |first5=Cristián |last6=Dreher Mansur |first6=Maria Cristina |last7=Arruda |first7=Janine Oliveira |last8=Barker |first8=Gary M. |last9=dos Santos |first9=Sonia Barbosa|url-access=subscription }}</ref><ref>{{Cite journal |last1=Greistorfer |first1=S. |last2=Miller |first2=I. |last3=von Byern |first3=J. |last4=Meyer-Rochow |first4=V. B. |last5=Geyer |first5=S. H. |last6=Weninger |first6=W. J. |last7=Steiner |first7=G. |date=2024 |title=In search of the glow—Three-dimensional reconstruction of Latia neritoides with specific focus on the mantel cavity (Mollusca; Gastropoda; Hygrophila) |journal=Journal of Zoology |language=en |volume=323 |issue=2 |pages=97–113 |doi=10.1111/jzo.13161 |issn=0952-8369|doi-access=free }}</ref><ref>{{Cite journal |last=Harbeck |first=K. |date=1996 |title=Die Evolution der Archaeopulmonata |url=https://repository.naturalis.nl/pub/317550/ZV1996305001.pdf |journal=Zoologische Verhandelingen |location=Leiden |volume=305 |pages=1-133, pls 1-33}}</ref> * Ellobiida mostly comprise snails, but also a single slug genus (''Smeagol'') and a limpet genus (''Trimusculus''). They are distributed from the marine upper littoral to moist terrestrial habitats. The conctractile pneustome is located on the right side of the body (quite posteriorly in Otina) and the pallial cavity functions as a lung. Operculum is absent, retained only in juveniles of ''Blauneria heteroclita''. The is one pair of contractile or partially retractable tentacles (but see ''Trimusculus''). Eyes are typically positioned medially of the tentacles, but in some ''Melampus'' they are on the tentacles at about 1/3 of their length. Radula typically has many similar teeth per row. The structure of the genital tract varies widely, from ''Pythia'' with a common spermoviduct leading to a single genital opening from which an external groove leads to the separate vas deferens and penis, to ''Leucopythia'' with female and male tracts separate from the carrefour and with two openings. There are free-swimming veliger larvae in marine species (e.g. ''Melampus'').<ref name=":2" /><ref name=":1" /><ref>{{Cite journal |last1=Tseng |first1=Roger |last2=Dayrat |first2=Benoit |date=2014 |title=Anatomical redescription of the limpet-like marine pulmonate Trimusculus reticulatus (Sowerby, 1835) |journal=The Veliger |volume=51 |issue=4 |pages=194–207}}</ref><ref>{{Cite journal |last=Morton |first=J. E. |date=1955 |title=The functional morphology of Otina otis, a primitive marine pulmonate. |journal=Journal of the Marine Biological Association of the United Kingdom |volume=34 |pages=113–150 |doi=10.1017/S002531540000864X }}</ref> * Systellommatophora include the mostly intertidal (subtidal to terrestrial) Onchidiidae as well as fully terrestrial Rathousiidae and Veronicellidae; all are shell-less slugs. Pallial cavity is used as a lung but may be gretaly reduced (Veronicellidae). Anus and the pneumostome (the latter except for Rathouisiidae) are shifted posteriorly and are located at the end of the body. Operculum is missing. There is one pair of retractable (Onchidiidae) or two pairs of contractile tentacles, eyes are located on tentacle tips. Radula has many similar teeth per row (but this is modified in Rathousiidae). There is a common spermoviduct, but then the male and female tracts divide and the male and female parts have widely separated openings. A free-swimming veliger larva is found in marine species of Ochidiidae, otherwise the development is direct.<ref name=":0">{{Cite journal |last=Dayrat |first=Benoît |date=2010 |title=Anatomical Re-Description of the Terrestrial Onchidiid Slug Semperoncis montana (Plate, 1893) |url=https://www.bioone.org/doi/abs/10.4002/040.052.0101 |journal=Malacologia |language=en |volume=52 |issue=1 |pages=1–20 |doi=10.4002/040.052.0101 |issn=0076-2997}}</ref><ref name=":2">{{Cite book |last=Barker |first=G. M. |url=http://www.cabidigitallibrary.org/doi/book/10.1079/9780851993188.0000 |title=The biology of terrestrial molluscs |date=2001 |publisher=CABI Publishing |isbn=978-0-85199-318-8 |editor-last=Barker |editor-first=G. M. |edition=1 |location=UK |pages=1–146 |language=en |chapter=Gastropods on Land: Phylogeny, Diversity and Adaptive Morphology |doi=10.1079/9780851993188.0000}}</ref><ref>{{Cite journal |last=Laidlaw |first=F. F. |date=1940 |title=Notes on some specimens of the genus Atopos (Mollusca Pulmonata) with microphotographs illustrating points in the anatomy of the genus. |url=https://lkcnhm.nus.edu.sg/wp-content/uploads/sites/11/app/uploads/2017/06/16brm121-132.pdf |journal=Bulletin of the Raffles Museum |volume=16 |pages=121-132, pl. 30-33}}</ref> * Stylommatophora is the most diverse group of land snails and slugs. Pallial cavity forms a spacious, air-filled lung. Anus is located on the right side close to the pneumostome. Operculum is missing. Two pairs of tentacle are retractable and the posterior pair bears eyes on the tips. Radula typically has many small, quite uniform teeth per row. The spermoviduct separates distally into an oviduct and vas deferens, but then there is a single genital opening for both male and female parts. Sperm transfer is via a spermatophore. Development is direct. A whole genome duplication occurred in the branch leading to Stylommatophora.<ref>{{Cite journal |last1=Chen |first1=Zeyuan |last2=Doğan |first2=Özgül |last3=Guiglielmoni |first3=Nadège |last4=Guichard |first4=Anne |last5=Schrödl |first5=Michael |date=2022-08-20 |title=Pulmonate slug evolution is reflected in the de novo genome of Arion vulgaris Moquin-Tandon, 1855 |journal=Scientific Reports |language=en |volume=12 |issue=1 |pages=14226 |doi=10.1038/s41598-022-18099-7 |issn=2045-2322 |pmc=9392753 |pmid=35987814}}</ref>
== References == {{Reflist}}
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Category:Gastropod taxa Category:Euthyneura Category:Tectipleura