Clausiliidae

**煙管蝸牛科**（學名：Clausiliidae），亦作**煙管螺科**或**密封蝸科**，是煙管蝸牛總科之下的一種小型和非常長的科級分類。本科物種主要都是左旋物種的陸生蝸牛、會呼吸空氣的有肺類腹足綱直神經亞綱柄眼目軟體動物:105。

本科舊屬柄眼目中尿道亞目（Mesurethra）及泛有肺類真有肺類柄眼類非正式群組彎尿道類。

截至 2008 年 10 月 (2008-10)，本科連同現生種及化石種合共有 1300 個物種，是陸生蝸牛當中其生物多樣性最高的分類單元（參照 Orthalicidae，在整個腹足綱來說也只是僅次於海生的小塔螺科）。

本科物種在台灣的名稱叫作「煙管蝸牛科」，這是得名於其殻口的獨特「煙管」解剖結構，叫作「腔襞」:116，讓牠們可以將牠們的殼口變得可以像一扇趟門那樣把殻口關閉起來。

## 型態描述

幾乎所有本科的貝殻都是左旋的:105，這在腹足綱物種的螺殻來說並不常見。殻的表面有肋樣構造:105。殻長約 10 到 15 mm:105。

These snails have shells which are extremely high-spired, with numerous whorls.

The shells tend to be club-shaped, tapering at both ends to a rounded nub. The aperture usually has visible folds.

### 「煙管」

煙管蝸牛科物種在肺螺類腹足綱軟體動物來說相當特殊，因為牠們大多數的殻口都有稱為「腔襞」的類似「煙管」構造，就像一扇趟門一樣。這種「煙管」與口蓋並不一樣，因為在肺螺類物種都已有外套膜肺，並沒有「口蓋」這種構造。

腔襞是一種石灰質構造，舌形或調匙形，可將其殻口完全關閉，以防止肉食性的甲蟲幼蟲入侵其柔軟的身體。腔襞有不同的形態，如「月形」:116 或「弓形」腔襞、「倒 C 形腔襞」:124、「工形腔襞」:118、「平行腔襞」:128 等，還可以分為多個部分，如：上腔襞、下腔襞、月形襞及主襞等:116。腔襞有「閉板」及「腔襞板」（clausilium slides）將其封閉：「閉板」是附生於殻軸的板狀構造，而腔襞板則是殻內壁的板狀突起:116，由殻壁摺疊而形成。

## 生物學

本科物種的染色體的單倍體介乎 21 到 30 之間:139,142。

## 分類學

本科的模式屬是煙管蝸牛屬（_Clausilia_ Draparnaud, 1805）。

根據 2005 年《布歇特和洛克羅伊的腹足類分類》，本科是彎尿道類非正式群組的成員，而這個群組在 2010 年的修訂被歸類為泛有肺類真有肺類演化支的成員。在 2017 年《布歇特等人的腹足類分類》，曲尿道類支序和泛有肺類都不再使用。

### 2005 年分類

根據 2005 年《布歇特和洛克羅伊的腹足類分類》，本科包括下列各亞科：

* 煙管蝸牛亞科 Clausiliinae Gray, 1855
  * tribe Clausiliini Gray, 1855 - synonym: Fusulinae Lindholm, 1924
  * tribe Gracillariini H. Nordsieck, 1979
* subfamily Alopiinae A. J. Wagner, 1913
  * tribe Alopiini A. J. Wagner, 1913
  * tribe Cochlodinini Lindholm, 1925 (1923) - synonym: Marpessinae Wenz, 1923
  * tribe Delimini Brandt, 1956 - synonym: Papilliferini Brandt, 1961 (n.a.)
  * tribe Medorini H. Nordsieck, 1997
  * tribe Montenegrinini H. Nordsieck, 1972
* subfamily Baleinae A. J. Wagner, 1913 - synonyms: Laciniariini H. Nordsieck, 1963; Tristaniinae Schileyko, 1999
* subfamily † Constrictinae H. Nordsieck, 1981
* 斜管螺亞科 Garnieriinae C.R. Boettger, 1926
  * 斜管螺族 Garnieriini C.R. Boettger, 1926
  * 傘管螺族 Tropidaucheniini H. Nordsieck, 2002
* subfamily † Eualopiinae H. Nordsieck, 1978
  * tribe † Eualopiini H. Nordsieck, 1978
  * tribe † Rillyini H. Nordsieck, 1985
* subfamily Laminiferinae Wenz, 1923
* subfamily Mentissoideinae Lindholm, 1924
  * tribe Mentissoideini Lindholm, 1924 - synonym: Euxininae I. M. Likharev, 1962
  * tribe Acrotomini H. Nordsieck, 1979
  * tribe Boettgeriini H. Nordsieck, 1979
  * tribe Euxinellini Neubert, 2002
  * tribe Filosini H. Nordsieck, 1979
  * tribe Olympicolini Neubert, 2002
  * tribe Strigileuxinini H. Nordsieck, 1994
  * tribe Strumosini H. Nordsieck, 1994
* subfamily Neniinae Wenz, 1923 - Neniastrinae H. B. Baker, 1930
* subfamily Phaedusinae A. J. Wagner, 1922
  * tribe Phaedusini A. J. Wagner, 1922
  * tribe Megalophaedusini Zilch, 1954 - synonym: Zaptyxini Zilch, 1954
* subfamily Serrulininae Ehrmann, 1927

### 2017 年分類

根據 2017 年《布歇特等人的腹足類分類》，煙管蝸牛科是柄眼目旋蝸牛亞目煙管蝸牛下目之下的煙管蝸牛總科的成員。本科之下的亞科從過往的 11 個亞科縮減至 8 個亞科 21 個族，分別如下：

* Subfamily Alopiinae A.J. Wagner, 1913
  * Tribe Alopiini A.J. Wagner, 1913
  * Tribe Cochlodinini Lindholm, 1925 (1923)
  * Tribe Delimini R. Brandt, 1956
  * Tribe Medorini H. Nordsieck, 1997
* Subfamily Clausiliinae Gray, 1855
* 斜管螺亞科 Garnieriinae C.R. Boettger, 1926
* Subfamily Laminiferinae Wenz, 1923
* Subfamily Mentissoideinae Lindholm, 1924
* Subfamily Neniinae Wenz, 1923
* Subfamily Peruiniinae H. Nordsieck, 2005
* 管螺亞科 Phaedusinae A.J. Wagner, 1922
  * 管螺族 Phaedusini A.J. Wagner, 1922
  * Tribe Serrulinini Ehrmann, 1927
  * Tribe Synprosphymini H. Nordsieck, 2007

## 屬

以下詳列本科之下各個亞科所屬的分類單元：

### 煙管蝸牛亞科

煙管蝸牛亞科（Clausiliinae）包括以下兩族：

* 煙管蝸牛族（Clausiliini）
  * 煙管蝸牛屬 _Clausilia_ Draparnaud, 1805 - with two subgenera: _Clausilia_ and _Strobeliella_ H. Nordsieck, 1977
  * _Erjaveci a_ Brusina, 1870
  * _Fusulus_ Fitzinger, 1833 - with two subgenera: _Erjaveciella_ H. Nordsieck, 1877 and _Fusulus_
  * _Julica_ H. Nordsieck, 1963
  * _Macrogastra_ Hartmann, 1841 - with three subgenera: _Macrogastra_, _Pseudovestia_ H. Nordsieck, 1877 and _Pyrostoma_ Vest, 1867
  * _Micridyla_ H. Nordsieck, 1973
  * _Neostyriaca_ A. J. Wagner, 1920
  * _Pseudofusulus_ H. Nordsieck, 1977 - has the only one species: _Pseudofusulus varians_
  * _Ruthenica_ Lindholm, 1924
* tribe Gracillariini
  * _Graciliaria_ Bielz, 1867

### Alopiinae

Alopiinae

tribe Alopiini

* _Alopia_ H. & A.Adams, 1855 - with two subgenera _Alopia_ and _Kimakowiczia_ Szekeres, 1969
* _Herilla_ H. & A. Adams, 1855
* _Triloba_ Vest, 1867

tribe Cochlodinini

* _Cochlodina_ A. Férussac, 1821 - with four subgenera: _Cochlodina_; _Cochlodinastra_ H. Nordsieck, 1977; _Paracochlodina_ H. Nordsieck, 1969; _Procochlodina_ H. Nordsieck, 1969
* _Macedonica_ O. Boettger, 1877

tribe Delimini

* _Barcania_ Brandt, 1956
* _Charpentieria_ Stabille, 1864 - with four subgenera: _Charpentieria_; _Gibbularia_ Monterosato, 1908; _Mauritanica_ O. Boettger, 1877; _Siciliaria_ Vest, 1867 and _Stigmatica_ O. Boettger, 1877
* _Delima_ Hartmann, 1842 - with four subgenera: _Delima_; _Dugiana_ Stamol & Slapnik, 2002; _Piceata_ O. Boettger, 1877 and _Semirugata_ O. Boettger, 1877
* _Dilataria_ Vest, 1867
* _Papillifera_ Hartmann, 1842

tribe Medorini H. Nordsieck, 1997

* _Agathylla_ H. & A.Adams, 1855 - with two subgenera _Agathylla_ and Agathyllina H. Nordsieck, 1969
* _Albinaria_ Vest, 1867 - also include former genera _Carinigera_ Moellendorff, 1873; _Cristataria_ Vest, 1867; _Isabellaria_ Vest, 1867; _Sericata_ O. Boettger, 1878.
* _Inchoatia_ Gittenberger & Uit de Weerd, 2006
* _Lampedusa_ - with two subgenera _Imitatrix_ Westerlund, 1884 and _Lampedusa_
* _Leucostigma_ A. J. Wagner, 1919
* _Medora_ H. & A. Adams, 1855
* _Muticaria_ Lindholm, 1925
* _Strigilodelima_ A. J. Wagner, 1924

tribe Montenegrinini

* _Montenegrina_ O. Boettger, 1877
* _Protoherilla_ A. J. Wagner, 1921

### Baleinae

* _Alinda_ H. & A. Adams, 1855 - with two subgenera: _Alinda_ and _Pseudalinda_ O. Boettger, 1877
* _Balea_ J. E. Gray, 1824
* _Bulgarica_ O. Boettger, 1877 - with three subgenera _Bulgarica_, _Pavlovicia_ H. Nordsieck, 1973 and _Strigilecula_ Kennard & Woodward, 1923
* _Lacinaria_ Hartmann, 1842
* _Likharevia_ H. Nordsieck, 1975
* _Menissa_ H. & A.Adams, 1855
* _Mentissella_ H. Nordsieck, 1973
* _Micropontica_ O. Boettger, 1881 - with two subgenera: _Baleopsina_ Lindholm, 1924 and _Micropontica_
* _Mucronaria_ O. Boettger, 1877 - with two subgenera: _Index_ O. Boettger, 1877 and _Mucronaria_
* _Quadriplicata_ O. Boettger, 1878
* _Vestia_ Hesse, 1916 - with three subgenera: _Brabenecia_ H. Nordsieck, 1974; _Vestia_ and _Vestiella_ H. Nordsieck, 1877

### Constrictinae

Fossil subfamily Constrictinae contains genera:

* † _Constricta_ O. Boettger, 1877 - type genus of the subfamily

### 斜管螺亞科

斜管螺亞科（Garnieriinae）包括下列各屬：

* _Garnieria_ Bourguignat, 1877
* 斜管螺屬 _Grandinenia_ H. Minato & C. Chen, 1984
* _Indonenia_ Ehrmann, 1927
* _Megalauchenia_ H. Nordsieck, 2007
* _Neniauchenia_ H. Nordsieck, 2002
* 傘管螺屬 _Tropidauchenia_ Lindholm, 1924
  * subsp. _Euryauchenia_ H. Nordsieck, 2007
  * subsp. _Tropidauchenia_

### Eualopiinae

Fossil subfamily Eualopiinae contains genera:

tribe † Eualopiini

* † _Eualopia_ O. Boettger, 1877 - type genus of the subfamily

tribe † Rillyini

* † _Rillya_ Munier-Chalmas [in P. Fischer], 1883 - type genus of the tribe

### Laminiferinae

* _Laminifera_ O. Boettger, 1863 - the type genus of the family Laminiferinae. Its type species is fossil. (Actual status of the genus?)
* _Bofilliella_ Ehrmann, 1927
* _Neniatlanta_ Bouguignat, 1876

### Mentissoideinae

* _Acrotoma_ O. Boettger, 1881 - with four subgenera: _Acrotoma_, _Acrotomina_ H. Nordsieck, 1977; _Bzybia_ H. Nordsieck, 1977 and _Castelliana_ Suvorov, 2002
* _Akramowskia_ H. Nordsieck, 1975
* _Armenica_ O. Boettger, 1877 - with two subgenera: _Armenica_ and _Astrogena_ Szekeres, 1970
* _Bitorquata_ O. Boettger, 1877
* _Boettgeria_ O. Boettger, 1863 - with two subgenera: _Boettgeria_and _Loosjesiella_ Neubert & Groh, 1998
* _Elia_ H. & A.Adams, 1855 - with four subgenera: _Acroeuxina_ O. Boettger, 1877; _Caucasica_ O. Boettger, 1877; _Elia_ and _Megaleuxina_ O. Boettger, 1877
* _Euxina_ O. Boettger, 1877 - with two subgenera: _Euxina_ and _Illunellaria_ Lindholm, 1924
* _Euxinastra_ O. Boettger, 1888 - with two subgenera: _Euxinastra_ and _Odonteuxina_ H. Nordsieck, 1875
* _Euxinella_ H. Nordsieck, 1973
* _Filosa_ O. Boettger, 1877
* _Galeata_ O. Boettger, 1877
* _Idyla_ H. & A. Adams, 1855 - with two subgenera: _Idyla_ and _Strigilidyla_ H. Nordsieck, 1994
* _Kazancia_ Neubert, 1992 - with two subgenera: _Kazancia_ and _Lasica_ Neubert, 1995
* _Macroptychia_ O. Boettger, 1877
* _Mentissoidea_ O. Boettger, 1877
* _Olympicola_ Hesse, 1916
* _Phrygica_ H. Nordsieck, 1994
* _Roseniella_ Thiele, 1931 - with two subgenera: _Chavchetia_ Neubert, 1992 and _Roseniella_
* _Sabaeola_ Lindholm, 1925
* _Sprattia_ O. Boettger, 1883
* _Strigileuxina_ H. Nordsieck, 1975
* _Strumosa_ O. Boettger, 1877
* _Sumelia_ H. Nordsieck, 1994

### 大口管螺亞科

大口管螺亞科（Neniinae Ehrmann, 1927）主要分佈於南美洲，但亦有少數分佈於中國南部、南亞及東南亞，包括以下各屬：

* _Andinia_ Polinski, 1922
* _Andiniella_ Weyrauch, 1958
* _Audiniastra_ H. Nordsieck, 2005
* _Bequaertinenia_ Weyrauch, 1964 - with two subgenera: _Bequaertinenia_ and _Miranenia_ Grego & Szekeres, 2004
* _Brevinenia_ Neubert & H. Nordsieck, 2005
* _Columbinia_ Polinski, 1924 - with three subgenera: _Columbinia_; _Paranenia_ Rehder, 1939 and _Steatonenia_ Pilsbry, 1926
* _Cyclonenia_ H. Nordsieck, 1999
* _Cylindronenia_ Ehrmann in Pilsbry, 1949 - with two subgenera: _Cylindronenia_ and _Cylindroneniella_ H. Nordsieck, 2007
* _Ehrmanniella_ Zilch, 1949
* _Gonionenia_ Pilsbry, 1926
* _Gracilinenia_ Polinski, 1922
* _Hemicena_ Pilsbry, 1949
* _Incaglaia_ Pilsbry, 1949 - with two subgenera: _Gibbonenia_ Zilch, 1954 and _Incaglaia_
* _Incania_ Polinski, 1922
* _Leuconenia_ H. Nordsieck, 2005
* 大口管螺屬 _Nenia_ H. & A. Adams, 1855
* _Neniaptyx_ H. Nordsieck, 2007
* _Neniatracta_ Pilsbry, 1926
* _Neniella_ Grego & Sekeres, 2004
* _Neniops_ Pilsbry, 1926
* _Nenisca_ Rehder, 1939
* _Parabalea_ Ancey, 1882
* _Peruinia_ Polinski, 1922
* _Pfeifferiella_ Weyrauch, 1957
* _Pseudogracilinenia_ Loosjes & Loosjes-van Bemmel, 1984
* _Steeriana_ Jousseaume, 1900
* _Symptychiella_ H. Nordsieck, 1999 - with two subgenera: _Divnenia_ H. Nordsieck, 2005 and _Symptychiella_
* _Temesa_ H. & A. Adams, 1855
* _Zilchiella_ Weyrauch, 1957

### 管螺亞科

管螺亞科（Phaedusinae Thiele, 1931）包括以下各屬：

管螺族 Phaedusini

* 管螺屬 _Phaedusa_ H. & A.Adams, 1855 - with four subgenera: _Metaphaedusa_ H. Nordsieck, 2001; _Phaedusa_; _Pseudophaedusa_ Tomiyama, 1984 and _Stereophaedusa_ O. Boettger, 1877

tribe Megalophaedusini

* _Leptacme_ Ehrmann, 1927
* _Oospira_ Blanford, 1872 - with five subgenera: _Atractophaedusa_ Ehrmann, 1927; _Formosana_ O. Boettger, 1877; _Formosanella_ H. Nordsieck, 2003; _Oospira_; _Paraformosella_ H. Nordsieck, 2003 and _Siphonophaedusa_ Lindholm, 1924
* _Zaptyx_ Pilsbry, 1901 - with two subgenera: _Prozaptyx_ Loosjes, 1950 and _Zaptyx_

tribe ?

* _Acanthophaedusa_ H. Nordsieck, 2007
* _Bathyptychia_ Lindholm, 1925 - with three subgenera: _Bathyptychia_; _Brachyptychia_ H. Nordsieck, 2001 and _Strictiphaedusa_ H. Nordsieck, 2001
* _Celsiphaedusa_ H. Nordsieck, 2001
* _Cylindrophaedusa_ O. Boettger, 1877 - with three subgenera: _Cylindrophaedusa_ and _Montiphaedusa_ H. Nordsieck, 2002
* _Dautzenbergiella_ Lindholm, 1924 - with two subgenera: _Dautzenbergiella_ and _Mansuyiella_ H. Nordsieck, 2003
* _Diceratoptyx_ Pilsbry, 1905
* 真管螺屬 _Euphaedusa_ O. Boettger, 1877： - with five subgenera: _Dentiphaedusa_ H. Nordsieck, 2003; _Euphaedusa_; _Papilliphaedusa_ H. Nordsieck, 2003; _Tauphaedusa_ H. Nordsieck, 2003 and _Telophaedusa_ H. Nordsieck, 2003：又名小煙管蝸牛屬。
* _Fuchsiana_ Gredler, 1887
* 擬管螺屬 _Hemiphaedusa_ O. Boettger, 1877：with 10 subgenera: _Dendrophaedusa_ H. Nordsieck, 2002; _Hemiphaedusa_; _Hemiphaedusoides_ H. Nordsieck, 2001; _Hemizaptyx_ Pilsbry, 1905; _Labyrinthiphaedusa_ H. Nordsieck, 2001; _Margaritiphaedusa_ H. Nordsieck, 2001; _Notoptychia_ Ehrmann, 1927; _Pinguiphaedusa_ Azuma, 1982; _Placeophaedusa_ Minato, 1994 and _Selenophaedusa_ Lindholm, 1924
* _Heterozaptyx_ Pilsbry, 1906
* _Juttingia_ Loosjes, 1965 - with two subgenera: _Juttingia_ and _Pseudohemiphaedusa_ H. Nordsieck, 2002
* _Lindholmiella_ Ehrmann, 1927
* _Liparophaedusa_ Lindholm, 1924
* _Loosjesia_ H. Nordsieck, 2002
* _Luchuphaedusa_ Pilsbry, 1901 - with two subgenera: _Luchuphaedusa_ and _Nesiophaedusa_ Pilsbry, 1905
* _Macrophaedusa_ Moellendorff, 1883
* _Macrophaedusella_ H. Nordsieck, 2001
* _Megalophaedusa_ O. Boettger, 1877 - with four subgenera: _Megalophaedusa_; _Mesophaedusa_ Ehrmann, 1929; _Mesozaptyx_ Kuroda, 1963 and _Mundiphaedusa_ Minato, 1979. (_Neophaedusa_ is a synonym of _Megalophaedusa_).
* _Metazaptyx_ Pilsbry, 1905
* _Miraphaedusa_ H. Nordsieck, 2005
* _Oligozaptx_ Pilsbry, 1905
* _Paraphedusa_ O. Boettger, 1877
* _Parazaptyx_ Pilsbry, 1905
* _Pliciphaedusa_ H. Nordsieck, 1998
* _Pulchraptyx_ Minato, 1981
* _Reinia_ Kobelt, 1876 - with three subgenera: _Parareinia_ H. Nordsieck, 1998; _Pictophaedusa_ Azuma, 1982 and _Reinia_
* _Renschiphaedusa_ Loosjes & Loosjes-van Bemmel, 1973
* _Selenoptyx_ Pilsbry, 1908
* _Serriphaedusa_ H. Nordsieck, 2001
* _Sinigena_ Lindholm, 1925
* _Stereozaptyx_ Pilsbry, 1905
* _Streptodera_ Lindholm, 1925
* _Synprosphyma_ Germain, 1919 - with two subgenera; _Excussispira_ Lindholm, 1925 and _Synprosphyma_
* _Thaumatoptyx_ Pilsbry, 1908
* _Tyrannophaedusa_ Pilsbry, 1901 - with two subgenera: _Aulacophaedusa_ Azuma, 1982 and _Tyrannophaedusa_
* _Tyrannozaptyx_ Käufel, 1930
* _Zaptychopsis_ Ehrmann, 1927

### Serrulininae

* _Caspiophaedusa_ Lindholm, 1924
* _Dobatia_ H. Nordsieck, 1973
* _Graecophaedusa_ Rähle, 1982
* _Laeviphaedusa_ Likharev & Stelkov, 1965
* _Microphaedusa_ H. Nordsieck, 1978
* _Nothoserrulina_ Németh & Szekeres, 1995
* _Pamphylica_ Németh & Szekeres, 1995
* _Pravispira_ Lindholm, 1924
* _Pontophaedusa_ Lindholm, 1924
* _Pontophaedusella_ H. Nordsieck, 1994
* _Sciocochlea_ C. Boettger, 1935
* _Serrulina_ Mousson, 1873
* _Serrulinella_ H. Nordsieck, 1984
* _Truncatophaedusa_ Majoros, Németh & Szili-Kovács 1994
* _Tsoukatosia_ Gittenberger, 2000 - with two subgenera: _Agiosspeleikos_ A. & P. Reischütz, 2003 and _Tsoukatosia_

## 保育

儘管非海洋軟體動物當中本科物種看似易危，但在《國際自然保護聯盟瀕危物種紅色名錄》只有 9 個本科的物種被列入名錄內。

## 參考文獻

## 延伸閱讀

* Maltz, T. K.; Sulikowska-Drozd, A. Life Cycles of Clausiliids of Poland — Knowns and Unknowns. _Annales Zoologici_. 2008, **58** (4): 857–880. doi:10.3161/000345408X396783 （英語）.
* Nordsieck H. (2007). _Worldwide Door Snails_. ConchBooks, 213 pp. ISBN 978-3-939767-07-7.
* Uit de Weerd D. R. (2004). "Molecular phylogenetic history of eastern Mediterranean Alopiinae, a group of morphologically indeterminate land snails". Doctoral thesis, Department of Biology, Faculty of Science, Leiden University. HTM （頁面存檔備份，存於網際網路檔案館）, PDF.
* Uit de Weerd D. R., Piel W. H. & Gittenberger E. (2004). "Widespread polyphyly among Alopiinae snail genera: when phylogeny mirrors biogeography more closely than morphology". _Molecular Phylogenetics and Evolution_ **33**(3): 533-548. doi:10.1016/j.ympev.2004.07.010

## 外部連結

* Nordsieck H. (2005–2007). www.clausilia.de at Internet Archive.
* Nordsieck H.: http://www.hnords.de （頁面存檔備份，存於網際網路檔案館）: Internet journal for the malacology of landsnails (with special regard to Clausiliidae).
* BioLib info （頁面存檔備份，存於網際網路檔案館）