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Pneumopulmonata is a clade of heterobranch pulmonate gastropods, defined based on results of phylogenomic studies[1][2] and named by Krug et al. (2020). The clade unites all of Panpulmonata except for Sacoglossa. Synapomorphy of the group is the presence of a pneumostome. It mainly contains clades occuring at the interface between sea and land, in freshwater, and in terrestrial habitats.

The following groups are included:

Siphonarioidea

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 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. Bursa copulatrix (gametolytic gland) is present. Development is through a free-swimming veliger larva.[3][4][5]

Acochlidiacea

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. Bursa copulatrix is present in some species. Some species have veliger larvae.[6][7][8][9]

Amphiboloidea

Pyramidelloidea

Glacidorboidea

Hygrophila

Ellobiida

Ellobiida mostly comprises snails, but also a single slug genus (Smeagol) and a limpet genus (Trimusculus). They are distributed from the upper littoral to moist terrestrial habitats. The conctractile pneustome is located on the right side of the body and the pallial cavity functions as a lung. Operculum is absent, but retained in juveniles of Blauneria heteroclita. The structure of the ganital 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.

Systellommatophora

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 (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. [10] Bursa copulatrix is present. A free-swiming veliger larva is found in marine species of Ochidiidae, otherwise the development is direct.[10][11][12]

Stylommatophora

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. Bursa copulatrix is present. Development is direct.

References

  1. ^ Krug, Patrick J.; Caplins, Serena A.; Algoso, Krisha; Thomas, Kanique; Valdés, Ángel A.; Wade, Rachael; Wong, Nur Leena W. S.; Eernisse, Douglas J.; Kocot, Kevin M. (2022-04-13). "Phylogenomic resolution of the root of Panpulmonata, a hyperdiverse radiation of gastropods: new insight into the evolution of air breathing". Proceedings of the Royal Society B: Biological Sciences. 289 (1972). doi:10.1098/rspb.2021.1855. ISSN 0962-8452. PMC 8984808. PMID 35382597.
  2. ^ Teasdale, Luisa Cinzia (2017). Phylogenomics of the pulmonate land snails. [Dissertation submitted in total fulfilment of the requirements of the degree of doctor of Philosophy]. Melbourne: School of BioSciences, The University of Melbourne.
  3. ^ Heller, Joseph (2015), "Marine Ancestors of most Land Snails: Pulmonates", Sea Snails, Cham: Springer International Publishing, pp. 257–264, doi:10.1007/978-3-319-15452-7_10, ISBN 978-3-319-15451-0, retrieved 2024-10-24
  4. ^ Simone, Luiz Ricardo L.; Seabra, Maria Inês G. L. (2017-09-01). "Shell and body structure of the plesiomorphic pulmonate marine limpet Siphonaria pectinata (Linnaeus, 1758) from Portugal (Gastropoda: Heterobranchia: Siphonariidae)". Folia Malacologica. 25 (3): 147–164. doi:10.12657/folmal.025.012.
  5. ^ Hubendick, B. (1978). "Systematics and comparative morphology of the Basommatophora". In Fretter1 Peake2, Vera1 J.2 (ed.). Pulmonates. Volume 2A. Systematics, Evolution and Ecology. London, New York, San Francisco: Academic Press. pp. 1–47. ISBN 0-12-267502-9.{{cite book}}: CS1 maint: numeric names: editors list (link)
  6. ^ Jörger, Katharina M; Stöger, Isabella; Kano, Yasunori; Fukuda, Hiroshi; Knebelsberger, Thomas; Schrödl, Michael (2010). "On the origin of Acochlidia and other enigmatic euthyneuran gastropods, with implications for the systematics of Heterobranchia". BMC Evolutionary Biology. 10 (1): 323. Bibcode:2010BMCEE..10..323J. doi:10.1186/1471-2148-10-323. ISSN 1471-2148. PMC 3087543. PMID 20973994.
  7. ^ Schrödl, Michael; Neusser, Timea P. (2010). "Towards a phylogeny and evolution of Acochlidia (Mollusca: Gastropoda: Opisthobranchia)". Zoological Journal of the Linnean Society. 158 (1): 124–154. doi:10.1111/j.1096-3642.2009.00544.x.
  8. ^ Neusser, Timea P; Heß, Martin; Schrödl, Michael (2009). "Tiny but complex - interactive 3D visualization of the interstitial acochlidian gastropod Pseudunela cornuta (Challis, 1970)". Frontiers in Zoology. 6 (1): 20. doi:10.1186/1742-9994-6-20. ISSN 1742-9994. PMC 2761907. PMID 19747373.
  9. ^ Brenzinger, B.; Neusser, T.P.; Glaubrecht, M.; Haszprunar, G.; Schrödl, M. (2010-12-20). "Redescription and three-dimensional reconstruction of the limnic acochlidian gastropod Strubellia paradoxa (Strubell, 1892) (Gastropoda: Euthyneura) from Ambon, Indonesia". Journal of Natural History. 45 (3–4): 183–209. Bibcode:2010JNatH..45..183B. doi:10.1080/00222933.2010.521862. ISSN 0022-2933.
  10. ^ a b Dayrat, Benoît (2010). "Anatomical Re-Description of the Terrestrial Onchidiid Slug Semperoncis montana (Plate, 1893)". Malacologia. 52 (1): 1–20. doi:10.4002/040.052.0101. ISSN 0076-2997.
  11. ^ Barker, G. M. (2001). "Gastropods on Land: Phylogeny, Diversity and Adaptive Morphology". In Barker, G. M. (ed.). The biology of terrestrial molluscs (1 ed.). UK: CABI Publishing. pp. 1–146. doi:10.1079/9780851993188.0000. ISBN 978-0-85199-318-8.
  12. ^ Laidlaw, F. F. (1940). "Notes on some specimens of the genus Atopos (Mollusca Pulmonata) with microphotographs illustrating points in the anatomy of the genus" (PDF). Bulletin of the Raffles Museum. 16: 121–132, pl. 30-33.