Phylum Echiura Stephen, 1965
(systematics, ecology and distribution)
by V.Murina
The Echiura popularly called "spoon-worms" are
inhabitants of the shore of polar, temperate and tropical seas.
They have also been collected from the floor of oceans at great
depths.
Morphology. Echiura comprise unsegmented,
coelomate, bilaterally symmetrical and soft-bodied invertebrates.
They size varies from a few millimetres to 25 centimeters in
length. They are sausage-shaped creature, with highly muscular
sacs filled with fluid in wich the long alimentary canal and
other organ freely move. The name "spoon-worms" is
derived from the shape of the contracted proboscis which is
muscular, sesory and lies as the anterior end of trunk but cannot
be retractor within body. The mouth is anteriorly place usually
at the base of the long proboscis used gathering food and the
anus at the posterior part of trunk. A pair of setae are usually
present on the ventral surface of the body just posterior of the
mouth and one or two rings of anal setae encircle the posterior
region of two genera. One to numerous nephridia, usually present
in pairs, are attached to the ventral surface of the body wall. A
pair of anal vesicles serves as excretory organ. Ther sexes are
separate, fertilisation is usually external and the larvae is
trochophore.
Ecology. Echiurans are marine except for a
few species that in brackish water of India. They are soft bodied
, almost defenceless creatures, that always live in protected
places. Most borrow in sand or mud, where they live in U-shaped
burrow. They inhabit also rock crevices, empty shells,
sand-dollar tests, pholad holes, roots of marine angiosperms.
Echiurans are detritus feeders. With the aid of their highly
muscular and sometimes spoon-or scoop-like probiscis they suck
sand, mud, coral fragments and finely divided particles from
their surroundings into their mouths and extract organic matter
from the ingested material. Some species, however, trap bacteria
and very fine particles of food in a slim net which is secreted
from the surface of the animal.
to begin
Family BONELLIDAE Lacaze-Duthers, 1858
Family Bonellidae has 28 genera and 70 species. The sexual
dimorphism of bonellids distinguishes them from all other
families of the phylum. The early larva may develop into a male
or female, depending on its proximity to an adult female. It is
finds as fastens on the proboscis of the female the developing
larva becomes a male. If it does not make contact with a female
it eventually becomes a female.
Males are usually small, about 1-6 mm long, planarian-like or
nematoform, without a proboscis. Parasitic males usually found in
the nephridium of the female.
Distribution. Family Bonellidae occurs widely
and thier bathymetric range is extensive. L.A.Zenkevitch (1966)
considered that the Bonellidae inhabit mainly the abyssal and
ultraabyssal zones of World Ocean. Really 44 species or 63%
bonellids are known to occur at depth of over 3000 m. Vitjazema
sp. have been draged from depth 10210 m (Philippine Trench)
(Beljaev, 1989). All these worms are obligate deposite feeders,
sorting detritus from the bottom surface. Most of the species
have the Nearcontinental and the lesser part have Oceanic types
of distribution (Sokolova, Murina 1997). The distribution of 18
shallow-water species are limited tropical-temperate waters,
especially Indo-Westpacific region. The 8 bathyal species are
known as having single find with the exception of Eubonellia
valida.
Checklist of BONELLIDAE Species
- Acanthobonellia Fisher, 1948
- Ac. miyajimai (Ikeda, 1904)
- Ac. pirotanensis Jose, 1964
- Ac. rollandoe Menon, Datta-Gupta et Johnson, 1964
- Acanthohamingia Ikeda, 1910
- Ac. ijimai (Ikeda, 1908)
- Ac. shiplei Ikeda, 1910
- Achaetobonellia Fisher, 1953
- Ach. maculata Fisher, 1953
- Alomasoma Zenkevitch, 1958
- +Al. belyaevi Zenkevitch, 1964
- +Al. chaetiferum Zenkevitch, 1964
- +Al. nordpacificum Zenkevitch, 1964
- Al. rhynchollulus Datta-Gupta, 1981
- Amalosoma Fisher, 1948
- Am. eddystonense Stephen, 1956
- Am. paradolum (Fisher, 1946)
- Archibonellia Fisher, 1919
- Arch. michaelseni Fischer, 1919
- Arch. mjoebergi Fischer, 1921
- Binophorus Salvini-Plaven, 1972 (may be new family?)
- B. geminolobotus Salvini-Plaven, 1972
- Bonellia Rolando, 1821
- +B. minor Marion & Rietsh, 1886
- +B. pacifica Zenkevitch, 1958
- B. plumosa Datta-Gupta, 1981
- B. pumicea Sluiter, 1891
- B. suhmii Selenka, 1885
- B. thomense Fischer, 1922
- B. viridis Rolando, 1821
- Bonelliopsis Fisher, 1946
- B. alaskana Fisher, 1946
- B. minutus Datta-Gupta, 1981
- Bruunelia Zenkevitch, 1966
- +B. bandae Zenkevitch, 1966
- Charcotus Datta Gupta, 1981
- Ch. charcotus Datta-Gupta, 1981
- Ch. clavatum Datta-Gupta, 1981
- Choanostomelia Zenkevitch, 1964
- +Cho. bruuni (Zenkevitch, 1964)
- +Cho. filatovae Zenkevitch, 1964
- +Cho. vinogradovae
Murina, 1978*
- Eubonellia Fisher, 1946
- E. longistomus Datta-Gupta, 1981
- +E. noratlanticum Datta-Gupta, 1981
- +E. valida Fisher, 1946
- Hamingia Danielssen & Koren, 1881
- +H. arctica Danielssen & Koren, 1881
- Ikedella Monro, 1927
- +I. artemiivanovi
Murina, 1976*
- I. qingdaoense Li Fenglu, Wang Wei & Zhou
Hong, 1994
- +I. bogorovi Zenkevitch, 1964
- I. misakiensis (Ikeda, 1904)
- Jacobia Zenkevitch, 1958
- +J.birsteini Zenkevitch, 1958
- J.similaris Datta-Gupta, 1981
- Kurchatovus Datta Gupta, 1977
- K. epeedentatus Datta-Gupta, 1986
- +K. tridentatus D.-G.,1977
- Maxmuelleria Bock, 1942
- M. aulacoferum (Herubel, 1924)
- +M. faex (Selenka, 1885)
- M. gigas (M. Muller, 1852)
- +M. lankesteri (Herdman, 1898)
- M. verrucosum (Studer, 1879)
- Metabonellia Stephen & Edmonds6 1972
- M. gigas (Nielsen, 1963)
- M. haswelli (Johnston & Tiegs, 1920
- Nellobia Fisher, 1946
- Prometor Fisher, 1948
- +P. benthophila Fisher,1948
- +P. gracilis Zenkevitch, 1957
- +P. grandis Zenkeviotch, 1957
- P. pocula Hartman & Bernard, 1960
- Protobonellia Ikeda, 1908
- Pseudobonellia Johnston & Tiegs, 1919
- P. biuterina Johnston et Teigs. 1919
- P. iraidii Murina,
1984
- Pseudoikedella Murina, 1978*
- +P. achaeta (Zenkevitch, 1958)
- Sluiterina Monro, 1927
- Torbenwolffia Zenkevitch, 1966
- +T. galathea Zenkevitch, 1966
- Vitjazema Zenkevitch, 1958
- +V. aleutica Zenkevitch, 1958
- +V. planirostris
Murina, 1978*
- +V. ultraabyssalis Zenktvotch, 1958
- Zenkevitchiola Murina, 1978
to begin
Family ECHIURIDAE Quatrefages, 1847
Family of Echiuridae has 7 genera and 82 species. The ansence
of sexual dimorphism distinguishes the echiurids from the
bonellids.
Morphology. Proboscis usually well-developed,
sometimes very long but never bifid; often easly detached from
the trunk. Trunk sac-like to cylindrical in shape and usually
covered with papillae; sometimes green or redish in colour.
Longitudinal musculature is some genera is thicked to form
longitudinal muscle bands.
Distribution. Echiurids tend to inhabit
shallow waters. Only 4 species (Thalassema elapsus, Th.ovatum,
Th.steinbecki, Arhynchite arhynchite) are known in abyssal zone
at depth more 4000 m (Datta-Gupta, 1981. 1983). The whole of
genera of Echiuridae (exception of genus Echiurus) are limited
the tropical-temperate waters. Two species of Echiurus (E. e.
echiurus and E. antarcticus) are found in the cold-water Arctic
and Antarctic.
Checklist of ECHIURIDAE Species
- Subfamily Echiurinae Quatrefages, 1847
- Echiurus Guerin-Meneville. 1831
- +E. antarcticus Spengel, 1912
- E. abyssalis Skorikow, 1906
- +E. e. echiurus (Pallas, 1767)
- E. e. alascanus Fisher, 1946
- E. sitchaensis (Brandt, 1845)
- E. torulobotus Salvini -Plawen, 1972
- Subfamily Thalassematinae Forbes and Goodsir, 1841
- Arhinchite Sato, 1937
- Ar. arhynchite (Ikeda, 1924)
- Ar. californicus Fisher, 1949
- Ar. hiscocki Edmonds, 1960
- Ar. inamoenus Fisher, 1946
- Ar. pugettensis Fisher, 1949
- Ar. rugosus Chen & yeh, 1958
- Lissomyema Fisher, 1946
- L. exilii (F. Muller, 1883)
- L. muellita (conn, 1886)
- Listriolobus Spengel, 1912
- L. bahamensis Fischer, 1926
- L. billitonensis Fischer, 1926
- L. bulbocaudatus Edmonds, 1963
- L. brevirostris Chen &Yeh, 1958
- L. capense (Jones & Stephen, 1966)
- L. hexamyotus Fisher, 1949
- L. peloides Fisher, 1946
- L. riukiuensis Sato, 1939
- L. sorbillans (Lampert, 1883)
- Ochetostoma Leuckart & Ruppell, 1828
- O. arkati (Prashad, 1935)
- O. australiense Edmonds, 1960
- O. baronii ( Greeff, 1879)
- O. bombayense (Prashad & Awati, 1929)
- O. caudex (Lampert, 1883)
- O. edax Fisher, 1946
- +O. eryhrogrammon Leuckart & Ruppell,
1828
- O. decameron (Lanchester, 1905)
- O. formosolum (Lampert, 1883)
- O. glaucum (Wesenberg-Lund, 1957)
- +O. griffini ( Wharton, 1913)
- O. hornelli (Prashad, 1921)
- O. hupferi (Fischer, 1895)
- O. indosinense Wesenberg-Lund, 1939
- O. keferstein (ten Broeke, 1925)
- O. kempi (Prashad, 1919)
- O. kokotoniense (Fischer, 1892)
- O. manjuyodense (Ikeda, 1905)
- O. mercator Wesenberg-Lund, 1954
- O. multilineatum (Fischer, 1914)
- O. myersae Edmonds, 1963
- O. octomyotum Fisher, 1946
- O. palense (Ikeda, 1924)
- O. pellucidum (Fischer, 1895)
- O. senegalense Stephen, 1960
- O. septemyotum Datta Gupta, Menon &
Johnsom, 1963
- O. stuhlmannu (Fischer, 1892)
- O. zanzibarense Stephen & Robertson,
1952
- Anelassorhynchus Annandale, 1922
- +An. abyssalis Fisher, 1949
- An. adelaidensis Edmonds, 1960
- An. branchiorhynchus (Annandale &
Kemp, 1915
- An. chaetiferus Datta Gupta , Menon &
Johnson, 1963
- An. dendrorhynchus (Annanandale &
Kemp, 1915)
- An. fisheri Datta-Gupta, 1974
- An. inanensis (Ikeda, 1904)
- +An. indivisus (Sluiter, 1900)
- An. loborhynchus Datta-Gupta & Menon,
1965
- An. microrhynchus (Prashad, 1919)
- An. moebii (Greeff, 1879)
- An. mucosus (Ikeda, 1904)
- An. porcellus Fisher, 1948
- An. sabinus (Lanchester, 1905)
- An. semoni (Fischer, 1896)
- An. vegrandis (Lampert, 1883)
- Thalassema Lamarck, 1801
- Th. antarcticum Stephen, 1941
- Th. arcassonense Cuenot, 1902
- Th. diaphanes Sluiter, 1888
- Th. elapsum Sluiter, 1912
- Th. fuscum Ikeda, 1904
- Th. harmani Fisher, 1947
- Th. marshalti Prashad, 1935
- Th. mortenseni Fischer, 1923
- Th. ovatum Sluiter, 1902
- Th. owstoni Ikeda, 1904
- Th. philostracum Fisher , 1947
- Th. steinbecki Fisher, 1946
- Th. sydninse Edmonds, 1960
- Th. thalassemum (Pallas, 1766)
- Th. viride Verrill, 1879
Family Urechidae Fisher and Macginitie,
1928
- Urechis Seitz, 1907
- Urechis caupo Fisher & MacGinitie, 1928
- Urechis chilensis (M. Muller, 1852)
- Urechis novaezealandiae (Dendi, 1897)
- Urechis unicinctus (von Drasche, 1881)
Family Ikedidae Bock, 1942
- Ikeda Wharton, 1913
- Ikeda taenioides (Ikeda) 1904
total 157 species
to begin
Remarks:
+ these species are preserved in the
depository of Institute of Oceanology Russian Academy of
Science Moskow.
* species were described by Murina V.V.
as n. sp.
Principal publications
- Beljaev G.M. Deepsea Oceanic Troughs and its Fauna.
Moscow: Nauka, 1989, 255p.(In Russian)
- Zenkevich L.A. New Genus and two new species of deepsea
Echiurids from the Pacific Ocean. Proceeding of
Oceanology Institute (USSR), v.23, 1957, pp.291-295 (In
Russian)
- Zenkevich L.A., Murina V. V. Deepsea Echiurids from the
Pacific Ocean. Proceeding of Oceanology Institute (USSR),
v.69, 1964, pp.102-114 (In Russian)
- Zenkevich L.A. Deepsea Echiurids from Nord-West of the
Pacific Ocean. Proceeding of Oceanology Institute (USSR),
v.27, 1958, pp.192-203 (In Russian)
- Murina V.V. New Abyssal Species of Echiurans from the
Pacific and Atlantic Oceans. Zoological Journal (USSR),
v.55, 1976, N6,pp.837-843 (In Russian)
- Murina V. V. New and Rare Echiurids of the Family
Bonelliidae. Proceeding of Oceanology Institute (USSR),
v.113, 1978, pp.107-119 (In Russian)
- Murina V. V. On the Sipunculans and Echiurans Fauna of
Mediterranean and Iberian Basin. Proceeding of Oceanology
Institute (USSR), v.117, 1982, pp.178-191 (In Russian)
- Murina V. V. A New Species of the Bonnelidae and a New
Finding of an Echiurid. Zoological Journal (USSR), v.63,
1984, N4,pp.617-620 (In Russian)
- Murina V.V. Echiurans Structure and Distribution of the
Mediterranean Sea. Proceeding of Oceanology Institute
(USSR), v.119, 1984, pp.82-98 (In Russian)
- Datta-Gupta A.K. A new genus and a new species of
Bonellidae Echiura from the Carrieben deep-see//Proc.
Zool. Soc. Calcutta. 1977, 30:17-23.
- Datta-Gupta A.K. Atlantic Echiurans. Part I. Report on
twenty-two species of deep-sea echiurans of the North and
South Atlantic Ocean//Bull. Mus. nath.Hist. nat. Paris, 4
ser,3 1981, section A, n 2: 353-378.
- Datta-Gupta A.K. Echiurans collected during the SAFARI I
cruise in the Indian ocean, south of Madagascar//Bull.
Mus. natn. Hist. nat. Paris, 4 ser.5, 1983, section a,
N4:1045- 1050.
- Murina V. V. Researches on the coast of Somalia. The
shore and the Dune of sar Uanle. A small collection of
Sipuncula and Echiura//Monitore zool. italiano:
Ital.Jorn.of Zool., 1980.-Suppl. 14,N 2:7-15. 71-72
- Nishikawa T. Nomenclatural remarks on the family-group
names of the Phylum Echiura. Proc. of the Biological Soc.
of Washington. 111(2):249-256. 1998.
- Sato H. Studies on the Echiuroidea, Sipunculoidea and
Priapuloidea of Japan// Science Reports Tohoku Imp.Univ.
1939, 4, 14, pp.339-460.
- Stephen A. C. & Edmonds S.J. The Phyla Sipuncula and
Echiura.-Brit. Museum Natur History, London,1972. 588 p.
- Zenkevitch L.A. The systematics and distribution of
abyssal and hadal (ultra-abyssal) Echiuroidea//Galathea
Rep., 1966, vol. 8, p.175-184.
to begin
Dr. Murina Galena-Vantsetti has the total information of the
bathymetrical and geographical distribution of each from 157
species of the Phylum Echiura. She would be glad to identify of
echiurans collection from any region of the World Ocean.
E-mail: murina@ibss.iuf.net
odmi@alpha.mhi.iuf.net
Copyright © 1998. Institute of Biology of the
Southern Seas, National Academy of Sciences of Ukraine
2, Nakhimov av., Sevastopol, 335011, Crimea, Ukraine. Tel:
380-692-591725 Fax: 380-692-592813
For more information contact zalex@ibss.iuf.net