扁蜷科

扁蜷科（學名：Planorbidae），亦作扁卷螺科，是腹足綱之下的一個單系群支序，本科物種皆為可呼吸空氣的淡水螺。原屬有肺目基眼亞目椎實螺總科，今屬異鰓類泛有肺類喜溼螺類支序。 本科多個物種的外殼都是平旋壳（planispiral），與一般腹足綱物種的外殼形狀不同。外殼看似是右旋，但其實是螺把殼上下顛倒來揹在身上，使左旋的外殼看起來像是右旋。 因其外形，本科物種又稱羊角螺（英语：Ramshorn snail）。 由於本科物種的頭和腹足都比較細小，使得牠們頭上絲般幼的觸角顯得很細長。 對於部份分類元來說，是否應該歸入扁蜷科仍有爭議。這對於淡水帽貝Ferrissia跟Ancylus這兩個屬來說由其合適。這些屬有時會被歸入椎实螺科（Lymnaeidae Rafinesque, 1815）；另外亦有文獻把上述兩個屬放進各自的一個科裡。然而，在布歇特和洛克羅伊的腹足類分類 (2005年)的分類，這些屬被歸為扁蜷亞科（Planorbinae）之下的Ancylini族。以下將詳述這個分類的具體內容。 根據布歇特和洛克羅伊的腹足類分類 (2005年)，本科包括下列各個亞科： 扁蜷亞科 Planorbinae Rafinesque, 1815 扁蜷族 Planorbini Rafinesque, 1815：synonyms: Choanomphalinae P. Fisher & Crosse, 1880; Orygoceratidae Brusina, 1882 tribe Ancylini Rafinesque, 1815 - synonym: Pseudancylinae Walker, 1923 (inv.) tribe Biomphalariini H. Watson, 1954 - synonyms: Acrorbini Starobogatov, 1958; Drepanothrematini Zilch, 1959; Taphiinae Harry & Hubendick, 1964 tribe Planorbulini Pilsbry, 1934 tribe Segmentinini F.C. Baker, 1945 subfamily Bulininae P. Fischer & Crosse, 1880 tribe Bulinini P. Fischer & Crosse, 1880 - synonyms: Laevapicinae Hannibal, 1912; Isidorinae Annandale, 1922; Gundlachiinae Starobogatov, 1967 tribe Coretini Gray, 1847 - synonyms: Pompholicinae Dall, 1866 (inv.); Camptoceratinae Dall, 1870; Megasystrophinae Tryon, 1871 (inv.); Pompholycodeinae Lindholm, 1927; Helisomatinae F. C. Baker, 1928; Bayardellini Starobogatov & Prozorova, 1990; Planorbariini Starobogatov, 1990 tribe Miratestini P. & F. Sarasin, 1897 - synonyms: Ferrissiinae Walker, 1917; Ancylastrinae Walker, 1923; Protancylinae Walker, 1923; Physastrinae Starobogatov, 1958; Ameriannini Zilch, 1959; Patelloplanorbidae Franc, 1968 tribe Plesiophysini Bequaert & Clench, 1939 subfamily Neoplanorbinae Hannibal, 1912 - synonym: Payettiinae Dall, 1924 subfamily Rhodacmeinae Walker, 1917 Albrecht等(2007)分析過本科部分物種，based on sequences of mitochondrial 18S ribosomal DNA and cytochrome-c oxidase I (COI) genes, and on the basis of the results, they rearranged the taxonomy like this: "A-clade" sensu Albrecht et al. (2007) Burnupia Walker, 1912 Tribus Bulinini Bulinus O. F. Müller, 1781 Indoplanorbis Annandale, 1921 Tribus Ancylini Rafinesque-Schmaltz, 1815 Ancylus O. F. Müller, 1774 Ferrissia Walker, 1903 Gundlachia Pfeiffer, 1849 Laevapex Walker, 1903 Hebetancylus Pilsbry, 1914 "B-clade" sensu Albrecht et al. (2007) Glyptophysa Crosse, 1872 Protancylus Sarasin, 1897 Kessneria Walker & Ponder 2001 Leichhardtia Walker, 1988 Tribus Camptoceratini Planorbarius Duméril 1806 扁蜷族 Planorbini 多旋螺屬 Anisus Studer, 1820 Bathyomphalus Charpentier, 1837 旋螺屬 Gyraulus Charpentier, 1837 Choanomphalus Gerstfeldt, 1859 扁蜷屬 Planorbis O. F. Müller, 1774 Tribus Segmentinini Segmentina Fleming, 1818 平扁蜷屬 Hippeutis Charpentier, 1837 类扁蜷属 Polypylis Pilsbry, 1906，多脉扁螺属 "C-Clade" sensu Albrecht et al. (2007) Biomphalaria Preston, 1910 Menetus H. & A. Adams, 1855 Planorbella Haldeman, 1843 Planorbula Haldeman, 1843 The following is a cladogram that shows the phylogenic relationships within the Planorbidae according to Albrecht 2007: 本科的模式屬是 Planorbis Müller（英语：Otto Friedrich Müller）。以下列出主要根據布歇特及其他人(2005)的分類，因為Albrecht (2007)的分類並未整全的包括本科所有的屬。至於每個屬的亞屬則依照 Glöer (2002) 的文獻。以下綜合三份文獻的結果： 扁蜷亞科 Planorbinae Rafinesque, 1815 ? 多旋螺屬 Anisus S. Studer, 1820 subgenus Disculifer C. Boettger, 1944 Bathyomphalus Charpentier, 1837 旋螺属 Gyraulus Charpentier, 1837 subgenus Torquis Dall, 1905 subgenus Lamorbis Starobogatov, 1967 subgenus Armiger Hartmann, 1843 平扁蜷屬 Hippeutis Charpentier, 1837：又名圓扁螺屬 tribe Ancylini Rafinesque, 1815 Ancylus O. F. Müller, 1773 - type genus of tribe Ancylini tribe Biomphalariini H. Watson, 1954 Biomphalaria Preston, 1910 - type genus of tribe Biomphalariini Drepanotrema Crosse & P. Fischer, 1880 扁卷螺族 Planorbini Rafinesque, 1815 扁卷螺屬 Planorbis Müller, 1773 tribe Planorbulini Pilsbry, 1934 Planorbula Haldeman, 1840 - type genus of tribe Planorbulini tribe Segmentinini F.C. Baker, 1945 Segmentina Fleming, 1818 - type genus of tribe Segmentinini 淤泥螺亞科 Bulininae P. Fischer & Crosse, 1880 印度扁卷螺屬 Indoplanorbis Annandale & Prashad, 1920：單型屬，只有Indoplanorbis exustus一個物種。 Planorbarius Duméril, 1806 Planorbella Haldeman, 1842 Menetus H. Adams & A. Adams, 1855 subgenus Dilatata Clessin, 1885 tribe Bulinini P. Fischer & Crosse, 1880 Bulinus O. F. Müller, 1781 - type genus of subfamily Bulininae Gundlachia Pfeiffer, 1849 tribe Coretini Gray, 1847 Coretus Gray, 1847 - type genus of tribe Coretini tribe Miratestini P. & F. Sarasin, 1897 Miratesta P. & F. Sarasin, 1897 - type genus of tribe Miratestini Amerianna Strand, 1928 Ferrissia Walker, 1903[來源請求] Pettancyclus Iredale, 1943 tribe Plesiophysini Bequaert & Clench, 1939 Plesiophysa P. Fischer, 1883 - type genus of tribe Plesiophysini subfamily Neoplanorbinae Hannibal, 1912 Neoplanorbis Pilsbry, 1906 - type genus of subfamily Neoplanorbinae subfamily Rhodacmeinae Walker, 1917 Rhodacmea Walker, 1917 - type genus of subfamily Rhodacmeinae 亚科地位未定(other genera that are not yet sorted are listed here) Acrorbis Odhner, 1937 Africanogyrus Özdikmen & Darilmaz, 2007 - synonym: Afrogyrus Brown & Mandahl-Barth, 1973 Afroplanorbis Thiele, 1931 Amphigyra Pilsbry, 1906 Anisopsis Sandberger, 1875 Antillorbis Harry & Hubendick, 1964 Armigerus Clessin, 1884 Australorbis Pilsbry, 1934 Bayardella Burch, 1977 Berellaia Laubrière & Carez, 1880 Camptoceras Benson, 1843 Camptoceratops Wenz, 1923 Carinifex W.G. Binney, 1865 Carinogyraulis Polinski, 1929 Ceratophallus Brown & Mandahl-Barth, 1973 Choanomphalus Gerstfeldt, 1859 Culmenella Clench, 1927 Fossulorbis Pilsbry, 1934 Glyptophysa Crosse, 1872 Helicorbis Benson, 1855 Helisoma Swainson, 1840 Intha Annandale, 1922 Isidorella Tate, 1896 Kessneria Walker & Ponder, 2001 Leichhardtia Walker, 1988 Lentorbis Mandahl-Barth, 1954 Macrophysa (Meek) Dall, 1870 Paraplanorbis Hanna, 1922 Patelloplanorbis Hubendick, 1957 Pecosorbis D.W. Taylor, 1985 Pentagoniostoma Branson, 1935 Perrinilla Hannibal, 1912 Physastra Tapparone-Canefri, 1883 Physopsis Krauss, 1848 Pingiella F.C. Baker, 1945 Pitharella F. Edwards, 1860 Planorbifex Pilsbry, 1935 Planorbina Haldeman, 1842 Platyphysa P. Fischer, 1883 Platytaphius Pilsbry, 1924 多脉扁螺属 Polypylis Pilsbry, 1906 Promenetus F.C. Baker, 1935 Protancylus P. & F. Sarasin, 1897 Pygmanisus Iredale, 1943 Segmentorbis Mandahl-Barth, 1954 Sineancylus Gutiérrez Gregoric, 2014 Syrioplanorbis F.C. Baker, 1945 Trochorbis Benson, 1855 Vorticifex Meek in Dall, 1870 The genus Camptoceratops Wenz, 1923 is no longer considered to be a planorbid. It was recognised by Curry (1965, p. 360) as a euthecosomatous pteropod (Heterobranchia) (note by Arie W. Janssen, 092507). 属名 Taphius H. Adams & A. Adams, 1855 为Biomphalaria的异名。 The shells of most species in this family are disk-like or button-like, being coiled in one plane, although several groups have shells that are more higher-spired, and some are limpet-like. All coiled shell Planorbidae are sinistral in their shell coiling, as is proved by their internal anatomy (the respiratory and the genital orifice are situated on the left side), however the animals carry their shells with what would normally be the ventral (i.e. umbilical) surface uppermost, and because of this, the shells appear to be dextral. Indeed, formerly planorbids were thought to have dextral shells, and so species of this family were figured as if they had dextral shells. Although it is now understood that these species are sinistral in shell coiling, disk-like Planorbid shells are often still shown in illustrations oriented as if they were dextral. Most species of coiled planorbids have a rather thin and moderately smooth shell, although more distinct sculpture such as a keel occurs in, and is diagnostic of, certain species. In the flat, keeled species, the whorls tend to overlap. The aperture has a sharp outer lip. A peristome can be present, but often the lip is not thickened nor reflected. Those planorbid species which have a high-spired shell may have a narrow umbilicus, but frequently this is covered by callus. In height most species vary between 6 mm and 6 cm, however, disk-like shells are usually less than about 2 cm in maximum dimension. Like all pulmonate aquatic snails, ramshorn shells do not have an operculum to close the shell aperture. Flat-coiled planorbid gastropod shells are hard to understand in terms of their coiling and orientation. Many of the shells of species in this family are almost planispiral in coiling such that one side of the shell often looks rather like the other side, but it is important to bear in mind that nonetheless there is an umbilical side and a spire side of the shell. In addition these are in fact sinistral shells, despite the fact that the snail carries its shell as if it were a normal dextral shell. To make sense of the shell coiling, the following facts are useful: In life, these pond snails hold their shells upside down compared to the normal gastropod shell orientation, with the umbilicus facing upwards The spire of the shell is quite sunken in many species, in addition it is carried facing downwards The umbilicus of the shell is very wide and shallow In some species the umbilicus is not as deeply "dished" as the sunken spire is, so superficially it can be hard to tell one from the other However, once it is understood that the planorbid shell is sinistral, if the shell is held with the aperture on the left and facing the observer, then the sunken spire side of the shell is uppermost. This is a convenience for understanding the shell, but is the opposite of the way the shell is actually carried in life. The side of the shell which is in fact the spire (a sunken spire) faces down in the living animal, contrary to what is the case in almost all other shelled gastropods. Because the shell is carried "upside down" like this, the aperture of the shell is angled to face downwards also, so the aperture faces a little towards the spire, not away from it, as is usually the case in other shelled gastropods. 相對於其他的軟體動物，扁卷螺科物種的血液都含有血紅素。因此，這些動物呼吸氧氣時都較其他軟體動物來得更有效；而血紅素亦令這些動物的身體帶紅色，這尤其在白化物種身上更明顯。 扁卷螺科的大多數物種都只能在淡水生活，只有少數物種能存活於鹹淡水交界。所以對於其他未能存活於鹹淡水的物種，亦令農夫可以用食鹽來清除牠們。 Ancestors of ramshorn snails are known with certainty since the Jurassic period. Modern taxa developed since the Cretaceous. Species in this family occur worldwide. In Northwest Europe about 20 species are known (including non-indigenous species). In this region, various extinct taxa are known to have occurred, starting in the Jurassic period. 膀胱螺科 Rafinesque, C. S. Analyse de la Nature ou tableau de l'univers et des corps organisés. Palermo. 1815: 223 pp., page 143 （法语）.  Strong, E. E.; Gargominy, O.; Ponder, W. F.; Bouchet, P. Global Diversity of Gastropods (Gastropoda; Mollusca) in Freshwater. Hydrobiologia. 2008, 595: 149-166. hdl:10088/7390 . doi:10.1007/s10750-007-9012-6 （英语）.  Bouchet, P.; Rocroi, J.-P.; Frýda, J.; Hausdorf, B.; Ponder, W.; Valdés, Á. & Warén, A. Classification and nomenclator of gastropod families. Malacologia (Hackenheim, Germany: ConchBooks). 2005, 47 (1–2): 1–397. ISBN 3-925919-72-4. OCLC 61351079. OL 42989175M. S2CID 83847814.  《香港淡水貝類 - 腹足綱》. 香港自然生態論壇. 2015-02-15 [2015-04-30]. （原始内容存档于2015-07-04） （中文（繁體））.  planospiral什么意思及同义词. 滬江詞庫. [2016-09-30]. （原始内容存档于2016-10-02）.  Albrecht, C.; Kuhn, K.; Streit, B. A molecular phylogeny of Planorboidea (Gastropoda, Pulmonata): insights from enhanced taxon sampling. Zoologica Scripta. 2007, 36: 27–39. doi:10.1111/j.1463-6409.2006.00258.x （英语）.  Glöer, P. Die Süßwassergastropoden Nord- und Mitteleuropas (北歐和中歐的淡水腹足類動物). Die Tierwelt Deutschlands [德國的動物] ?. Hackenheim, Germany: ConchBooks. 2002: 19–20. ISBN 3-925919-60-0 （德语）.  Liu L et al. (2010) "The phylogeography of Indoplanorbis exustus (Gastropoda: Planorbidae) in Asia". Parasites & Vectors 3: 57. doi:10.1186/1756-3305-3-57. Mollusc Specialist Group (1996). Culmenella rezvoji. In: IUCN 2010. IUCN Red List of Threatened Species. Version 2010.4. <www.iucnredlist.org>. Downloaded on 08 March 2011. 维基共享资源上的相关多媒体资源：扁蜷科 （英文） NCBI Taxonomy Browser: Planorbidae （页面存档备份，存于互联网档案馆） （英文） Systema naturae 2000 (classification) - Taxon: Planorbidae （页面存档备份，存于互联网档案馆） Planorbidae. ITIS.  （英文） http://mkohl1.net/Planorbidae.html （页面存档备份，存于互联网档案馆） Frank Collins Baker, The Molluscan Family Planorbidae, The University of Illinois press, Urbana, 1945