All three tentaculate
ctenophores above have been called "Pleurobrachia bachei " at one time or another.
They are (left to right): Pleurobrachia bachei, Euplokamis
dunlapae, and Hormiphora sp., all from Friday Harbor,
Washington, USA. Photographic
image copyright 1998
Claudia E. Mills.
The suggested citation for this work is listed below:
Mills, C.E. Internet
1998-present. Phylum Ctenophora: list of all valid species names.
Electronic internet document available at http://faculty.washington.edu/cemills/Ctenolist.html.
Published by the author, web page established March 1998, last
updated (see date at end of page).
Because the scientific literature on the Ctenophora is widely dispersed and much of it is difficult to locate, I have compiled a list of all classes, orders, families, genera and species of ctenophores that seem to be in use at the present time. This list is the result of a rather extensive search of the literature, combined with a little independent thinking. The phylogenetic order used in this list is derived primarily from the morphological work of Richard Harbison (1984, 1996) and Harbison and Madin (1982). It will undoubtedly be heavily modified as genetic studies of relationships between these animals are published. There is little question that some of the species listed are the same as others, but in most cases, no author has dealt with the problem species in the peer-reviewed, published literature, so the redundant names are still "current" and valid.
I suspect that there are actually 100-150 good species of described ctenophores, of which 30-35 (the platyctenids) are benthic in habitat. (There are probably at least 25 more known, but undescribed, ctenophores, many of them in the deep-sea -- stay tuned for descriptions of most of these eventually.) In many cases, I have listed synonyms to good species names in parentheses on the line following - the list of synonyms is not complete - it's only a start. There must be many other synonyms in this list of names (the genus Beroe, for instance, has many names that might ultimately be synonomized in future studies, but that may be a very distant future). A number of other lists of ctenophores, many of which seem to derive from this one, can be found elsewhere on the web; I do not claim to be the only authority about which names are presently in use for the Ctenophora. One of the purposes of this page is to have accurate access to the spelling, as well as describing author and date, for each species name.
Note that genus and species names are always written in italics. If the author and date following a species name are within parentheses, that is the taxonomists' code for indicating that the species has been moved from the genus in which it was originally described; author and date not in parentheses indicates that the genus has not changed from the original description. (I am not making this format up - it is strictly formalized and governed by an international body of taxonomists.)
This list has taken me several years to put together and I would really appreciate it if you cite it as an electronic publication if you use it in your studies (suggested citation above). The list is an ongoing work and is subject to modification at any time. I consider it my intellectual property and do not want to find it dumped wholesale onto someone else's web page - links to it are fine. I welcome any additions or corrections that anyone may wish to make - please contact me at the above address. In order to be true to its in-progress nature, I have left my personal shorthand in the electronic manuscript, so any entry preceeded by . means that I have not yet been able to check the original citation for accuracy. Although I have chosen to put this list of species names and authors on the web, it will be noted that I have not included the bibliography from which it derives on the web at this time.
I have included links to ctenophore images located elsewhere on the web. In the event that there is more than one good image of the same species, I have linked the best one or two, whose identification I am sure about, for you. In a few cases, I have linked images to species that I do not know well, so the identifications could be wrong, but I wouldn't know - such links are clearly labeled. Any additions to these photograph links are welcome - please let me know. Be aware that many ctenophore photographs on the web are misidentified.
Phylum CTENOPHORA Eschscholtz,
1829
Class Tentaculata Eschscholtz, 1825
Class/Subclass Typhlocoela Ospovat, 1985
Order Cydippida Gegenbaur, 1856 (or Lesson, 1843?)
Family Haeckeliidae Krumbach, 1925
Aulacoctena Mortensen, 1932
acuminata Mortensen, 1932
Haeckelia Carus, 1863
beehleri (Mayer, 1912)
bimaculata C. Carré and D. Carré, 1989 - image contains B. beehleri, B. rubra, and B. bimaculata
filigera (Chun, 1880)
(= Euchlora filigera Chun, 1880)
rubra (Kölliker, 1853)
(= Euchlora rubra (Kölliker, 1853))
Family Ctenellidae C. Carré and D. Carré, 1993
Ctenella C. Carré and D. Carré, 1993
aurantia C. Carré and D. Carré, 1993
Family Bathyctenidae Mortensen, 1932
Bathyctena Mortensen, 1932
chuni (Moser, 1909)
Family Lampeidae Krumbach, 1925
Lampea Stechow, 1921
. elongata (Quoy and Gaimard, 1824)
fusiformis (A. Agassiz and Mayer, 1902)
lactea (Mayer, 1912)
pancerina (Chun, 1879) - another image showing this species eating a salp chain
(= Lampetia pancerina Chun, 1880)
(= Gastrodes parasiticum Korotneff, 1888)
Family Pleurobrachiidae Chun, 1880
Ceroctena C. Carré and D. Carré, 1991
bicornis C. Carré and D. Carré, 1991
Hormiphora L. Agassiz, 1860 (see also the illustration at the top of this web page)
californensis (Torrey, 1904)
cilensis (Ghigi, 1909) (may = californensis above (Mills 1987))
cucumis (Mertens, 1833)
(= Janira cucumis (by Agassiz 1860 and Agassiz 1865))
(= Euplokamis cucumis (by Torrey, 1904))
elliptica (Eschscholtz, 1829)
. hormiphora (Gegenbaur, 1856)
"coeca" (= Hormiphora coeca Stanford, 1931, a nomen nudem)
labialis Ghigi, 1909
luminosa Dawydoff, 1946
ochracea (A. Agassiz and Mayer, 1902)
octoptera (Mertens, 1833)
palmata Chun, 1898
(= Hormiphora amboinae Moser, 1907 by Bigelow, 1912)
(= Hormiphora fusiformis Agassiz and Mayer, 1902 by Bigelow 1912)
(= Hormiphora japonica Moser, 1907 by Bigelow, 1912)
piriformis Ghigi, 1909
plumosa L. Agassiz, 1860 (image is glass model) or is it this? - I'll have to check (they are not the same)
(= Cydippe hormiphora Gegenbaur, 1856)
(= Cydippe plumosa M. Sars, 1859)
(= Cydippe densa Spagnolini, 1870)
polytrocha Dawydoff, 1946
punctata Moser, 1909
. sibogae Moser, 1903
spatulata Chun, 1898 (= P. brunnea by Bigelow, 1912)
Minictena C. Carré and D. Carré, 1993
luteola C. Carré and D. Carré, 1993
Moseria Ghigi, 1909 (likely absorbed by Pleurobrachia and Hormiphora)
This genus as established included M. australis (Benham), M. pigmentata (Moser) and M. striata (Moser). These assignments do not seem to be in current use, but Ghigi's' 1909 paper is not widely known. Moseria is also in use as a siphonophore genus name.
Pleurobrachia Fleming, 1822
. arctica Wagner, 1885 (in Moser)
australis (Benham, 1907)
bachei A. Agassiz, 1860 (see another illustration at the top of this web page)
brunnea Mayer, 1912
. cyanea (Chun, 1889) = Tinerfe? - see below
. globosa Moser, 1903
var. ceylonensis Browne, 1905
var. bengalensis Annandale and Kemp, 1915
latipharyngea Dawydoff, 1946
pigmentata Moser, 1903
pileus (O. F. Müller, 1776)
rhododactyla L. Agassiz, 1860
rhodopis Chun, 1879
striata Moser, 1907
Sabaudia Ghigi, 1909
liguriae Ghigi, 1909
Tinerfe Chun, 1898
cyanea (Chun, 1889)
Family Euplokamididae Mills, 1987 ( = Euplokamidae, changed in agreement with Cairns et al., 2002)
Euplokamis Chun, 1879
crinita (Moser, 1909)
dunlapae Mills, 1987 (see another more detailed illustration at the top of this web page)
helicoides (Ralph and Kaberry, 1950)
octoptera (Mertens, 1833) - likely to be Mertensia ovum
stationis Chun, 1879
Family Cryptocodidae Leloup, 1938
Cryptocoda Leloup, 1938
gerlachi Leloup, 1938
Family Mertensiidae L. Agassiz, 1860
Callianira Péron and Lesueur, 1808
antarctica Chun, 1897 - I do not know this species and cannot verify that the photo ID is as stated on that website (C.E.M).
bialata Della Chiaje, 1841
(= Eschscholtzia cordata Kölliker, 1853)
cristata Moser, 1909
compressa (Mertens, 1833) - may be Mertensia ovum
. diploptera Lamarck, 1843 (in Moser)
ficalbi Curreri, 1900
. hexagona Slabber
. triploptera Lamarck, 1843 (in Moser)
Charistephane Chun, 1879
fugiens Chun, 1879
. Mertensia Lesson, 1830
ovum (Fabricius, 1780) another image of this species
"Undescribed genus" Harbison and Mills, in prep. (with masses of fine pink-red tentilla)
"Species 1" Harbison and Mills, in prep.
"Species 2" Harbison and Mills, in prep.
"Species 3" Harbison and Mills, in prep.
Family Dryodoridae Harbison, 1996 (earlier?)
. Dryodora L. Agassiz, 1860
glandiformis (Mertens, 1833) note similarity of sheaths and canals to Cambodgia
Family not determined
Big red undescribed cydippid "Agmayeria
tortugensis"- but never formally described (also known as "Tortugas Red")
Cydippida, but family affiliation not determined (mostly juveniles):
Cydippe Eschscholtz, 1829
. densa Forskal, 1775 (in Moser)
ovata Lesson, 1843
. quadricostata Sars, 1835 (in Moser)
Eschscholthia Lesson 1836
dimidiata (Eschscholtz, 1829)
Eschscholtzia Kölliker, 1853
. borealis Wagner, 1885 (in Moser)
Paracelsia Dawydoff, 1946
quadriloba Dawydoff, 1946
Thoe Chun, 1879 1878?
paradoxa Chun, 1879 or 1878?
. Tinerfe coerulea Chun
Tizardia Dawydoff, 1946
phosphorea Dawydoff, 1946
FOSSIL CTENOPHORA - Order Cydippida
Archeocydippida Stanley and Stürmer, 1987
hunsrueckiana Stanley and Stürmer, 1987. Devonian: Hunsrück Slate, Germany
Paleoctenophora Stanley and Stürmer, 1983
brasseli Stanley and Stürmer, 1983. Devonian: Hunsrück Slate, Germany
. Podolites Ljaschendo 1969? (Berger 1981 Zoo. Record). Silurian
FOSSIL CTENOPHORA - Order Unknown
Family Ctenorhabdotidae Conway Morris and Collins, 1996
Ctenorhabdotus Conway Morris and Collins, 1996
capulus Conway Morris and Collins, 1996. Middle Cambrian: British Columbia, Canada
Family Fasciculidae Conway Morris and Collins, 1996
Fasciculus Simonetta and Della Cave, 1978
vesanus Simonetta and Della Cave, 1978. Middle Cambrian: British Columbia, Canada
Family Xanioascidae Conway Morris and Collins, 1996
Xanioascus Conway Morris and Collins, 1996
canadensis Conway Morris and Collins, 1996. Middle Cambrian: British Columbia, Canada
For a good review of fossil ctenophores
with all citations, see: Conway Morris, S. and D.H. Collins, 1996.
Middle Cambrian ctenophores from the Stephen Formation, British
Columbia, Canada. Phil Trans. R. Soc. Lond. B, 351: 279-308.
. Order Platyctenida
Bourne 1900 = Coeloplanidea Stechow, 1922
Family Ctenoplanidae Willey, 1896
Ctenoplana Korotneff, 1886
bengalensis Gnanamuthu and Nair, 1948
duboscqui Dawydoff, 1929
korotneffi Willey, 1896
kowalevskii Korotneff, 1886
maculomarginata Yosii (Yoshi), 1933 (also as maculomarainata in
original description)muculosa Yosii (Yoshi), 1933 (also maculosa, in original description's figure)
perrieri (Dawydoff, 1930)
rosacea Willey, 1896
sub-genus Diploctena Fricke and Plante, 1971
neritica Fricke and Plante, 1971
sub-genus Planoctena Dawydoff, 1936
agnae (Dawydoff, 1929)
caulleryi Dawydoff, 1936
yurii (Dawydoff, 1929)
Family Tjalfiellidae Komai, 1922
Tjalfiella Mortensen, 1910
tristoma Mortensen, 1910
Family Lyroctenidae Komai, 1942
Lyrocteis Komai, 1941
flavopallidus Robilliard and Dayton, 1972
imperatoris Komai, 1941
Family Savangiidae Harbison and Madin, 1982 (?)
Savangia Dawydoff, 1950
atentaculata Dawydoff, 1950
Family Coeloplanidae Willey, 1896
. Coeloplana Kowalevsky, 1880 - I do not know this species and cannot verify that the photo ID is as stated on that website, but it certainly is beautiful (C.E.M).
agniae Dawydoff, 1930 - apparently not the same as Ctenoplana agnae.
(Tentacles of both spp. compared in same figure, Dawydoff (1933) p. 122)var. striata Dawydoff, 1938
astericola Mortensen, 1927 - I do not know this species and cannot verify that the photo ID is as stated on that website (C.E.M).
bannwarthii Krumbach, 1933 - Scroll down. I do not know this species and cannot verify that the photo ID is as stated on that website (C.E.M).
bocki Komai, 1920
duboscqui Dawydoff, 1930
echinicola Tanaka, 1932
gonoctena Krempf, 1920
var. natalensis Pople, 1960
var. rosea Dawydoff, 1938
indica Devanesen and Varadarajan, 1942
komaii Utinomi, 1963
krusadiensis Devanesen and Varadarajan, 1942
lineolata Fricke, 1970
meteoris Thiel, 1968 - I do not know this species and cannot verify that the photo ID is as stated on that website (C.E.M).
mesnili Dawydoff, 1938
metschnikowii Kowalevsky, 1880
mitsukurii Abbott, 1902
perrieri Dawydoff, 1930
punctata Fricke, 1970
scaberiae Matsumoto and Gowlett-Holmes, 1996
sophiae Dawydoff, 1938
tattersalli Devanesen and Varadarajan, 1942
thomsoni Matsumoto, 1999
weilli Dawydoff, 1938
willeyi Abbott, 1902
wuennenbergi Fricke, 1970
sub-genus Benthoplana Fricke and Plante, 1971
meteoris Thiel, 1968
Vallicula Rankin, 1956
multiformis Rankin, 1956
Class/Subclass Cyclocoela Ospovat, 1985 (includes through Beroida)
Order Cambojiida Ospovat, 1985
Family Cambojiidae Ospovat, 1985
Cambodgia Dawydoff, 1946 (also spelled in same paper Cambodjia Dawydoff, 1946)
elegantissima Dawydoff, 1946 Note similarity of sheaths and canals to Dryodora
. Order Ganeshida Moser, 1908
Family Ganeshidae Moser, 1907
Ganesha Moser, 1907
elegans (Moser, 1903)
(= Lampetia elegans Moser in Moser 1908)
annamita Dawydoff, 1946
Order Cryptolobiferida Ospovat, 1985
Family Cryptolobatidae Ospovat, 1985
Cryptolobata Moser, 1909 (Ospovat, 1985, says this may be larva of Lobocrypta)
primitiva Moser, 1909
Lobocrypta Dawydoff, 1946
annamita Dawydoff, 1946
Lobata with uncertain affinity:
Axiotima Eschscholtz, 1829
gaedii Eschscholtz, 1829
"Die 6 eckige Beroe" von Slabber (in Moser)
Calymma Eschscholtz, 1829
trevirani Eschscholtz, 1829
Order Thalassocalycida Madin and Harbison, 1978
Family Thalassocalycidae Madin and Harbison, 1978
Thalassocalyce Madin and Harbison, 1978
inconstans Madin and Harbison, 1978
Order Lobata Eschscholtz, 1825
Family Bathocyroidae Harbison and Madin, 1982
Bathocyroe Madin and Harbison, 1978
fosteri Madin and Harbison, 1978
paragaster (Ralph and Kaberry, 1950)
Family Bolinopsidae Bigelow, 1912
Bolinopsis L. Agassiz, 1860 (=Bolina Mertens, 1833)
. chuni (von Lendenfeld, 1884)
elegans (Mertens, 1833)
indosinensis Dawydoff, 1946
infundibulum (O.F. Müller, 1776)
(= B. septentrionalis Mertens, 1833 by Mayer 1912)
microptera (A. Agassiz, 1865)
mikado (Moser, 1907)
ovalis (Bigelow, 1904)
rubripunctata Tokioka 1964
vitrea (L. Agassiz, 1860)
(= Bolina hydatina Chun, 1879)
Lesueuria Milne Edwards, 1841
hyboptera A. Agassiz, 1865
pinnata Ralph and Kaberry, 1950
tiedemanni (Eschscholtz, 1829) moved by Moser, 1908
vitrea Milne Edwards, 1841
MnemiopsisL. Agassiz, 1860
gardeni L. Agassiz, 1860 (said to be junior synonym of M. leidyi by Seravin (1994),
which it cannot be, because it has seniority by date)leidyi A. Agassiz, 1865
mccradyi Mayer, 1900 = junior synonym of M. leidyi by Seravin, 1994
(but I do not accept this synonomy)
Family Leucotheidae Krumbach, 1925 ( = Leucothoeae Lesson 1843; = Eucharidae Chun, 1880)
Leucothea Mertens, 1833 (=Alcinoe Della Chiaje, 1841, =Chiaia Lesson, 1843)
grandiformis Agassiz and Mayer, 1899
. harmata
japonica Komai, 1918
multicornis (Quoy and Gaimard, 1824)
(= Eucharis multicornis Eschscholtz, 1829)
(= Chiaia neapolitana Lesson, 1843)
(= Chiaja palermitana Milne Edwards, 1857)
(= Cydippe brevicostata Will, 1844 - larva, by Chun, 1879)
(= Eschscholtzia pectinata Kölliker, 1853 - larva, by Chun, 1879)
ochracea Mayer, 1912
pulchra Matsumoto, 1988
. Family Ocyropsidae (= Ocyroidae Lesson, 1843)
Alcinoe Rang, 1828 (in Moser 1908)
. rosea Mertens, 1833
vermicularis Rang 1828 (in Moser 1908))
Ocyropsis Mayer, 1912 (= Ocyroe Rang, 1828)
crystallina (Rang, 1828)
subsp. crystallina (Rang, 1828)
subsp. guttata Harbison and Miller, 1986
fusca (Rang, 1828)
maculata (Rang, 1828)
subsp. immaculata Harbison and Miller, 1986
subsp. maculata (Rang, 1828)
. pteroessa Bigelow 1904 (in Moser)
Family Eurhamphaeidae L. Agassiz, 1860
Eurhamphaea Gegenbaur, 1856 = probably only one species, E. vexilligera
chamissonis (Eschscholz, 1829)
heteroptera (Chamisso and Eysenhardt, 1821)
kuhlii (Eschscholtz, 1829)
schweiggeri (Eschscholtz, 1829)
vexilligera Gegenbaur, 1856
(= Mnemia elegans M. Sars, 1856)
[Family Deiopeidae Chun, 1880 (Harbison (1996) and Harbison, Matsumoto and Robison (2001) combined this family into the Eurhamphaeidae)]
Deiopea Chun, 1879
kaloktenota Chun, 1879
Kiyohimea Komai and Tokioka, 1940
aurita Komai and Tokioka, 1940
usagi Matsumoto and Robison, 1992
Family Lampoctenidae Harbison, Matsumoto and Robison, 2001
Lampocteis Harbison, Matsumoto and Robison, 2001
cruentiventer Harbison, Matsumoto and Robison, 2001
Family Lobatolampeidae Hirota, 2000
Lobatolampea Horita, 2000
tetragona Horita, 2000
Order Cestida Gegenbaur, 1856
Family Cestidae Gegenbaur, 1856
Cestum Lesueur, 1813
veneris Lesueur, 1813 another image of this species
(= Cestus veneris Chun, 1879 and 1880)
Velamen Krumbach, 1925
parallelum (Fol, 1869) another image of this species
(= Vexillum parallelum Fol, 1869)
(= Folia parallela Mayer, 1912)
Class Nuda Chun 1879 (at least, perhaps earlier?)
Order Beroida Eschscholtz, 1829 or 1825?
Family Beroidae Eschscholtz, 1825
Beroe Browne, 1756
abyssicola Mortensen, 1927
australis Agassiz and Mayer 1899
baffini Kramp, 1942
. basteri Lesson, 1830
campana Komai, 1918
compacta Moser, 1909
constricta Chamisso and Eysenhardt, 1821
cucumis Fabricius, 1780
(= Beroe roseola Agassiz, 1860 by Mayer, 1912)
. culcullus Martens, also Eschscholtz, 1829
flemingii (Eschscholtz, 1829)
forskalii Milne Edwards, 1841 another image of this species
gilva Eschscholtz, 1829
hyalina Moser, 1907
gracilis Künne, 1939 another image of this species
. macrostoma Péron and Lesueur, 1808 or 1809 (in Moser)
. mitraeformis Lesson, 1830 (in Moser)
mitrata (Moser, 1907)
. ovale Bosc 1802 (in Moser)
. ovata Bruguière, 1789 or Eschscholtz, 1829 - for a full description of the taxonomic problem presented by Beroe ovata, see the paragraph written by Richard Harbison in Mills et al. 1996, Scientia Marina, vol. 60(1), p. 161.
(= Idyiopsis affinis L. Agassiz, 1860 by Mayer, 1912)
(= Beroe clarkii (Agassiz, 1860) by Mayer, 1912)
(= Idya mertensii Mertens, 1833 by Mayer, 1912)
(= Beroe capensis Chamisso and Eysenhardt, 1821 by Chun, 1880)
(= Beroe punctata Chamisso & Eysenhardt, 1821 by Mayer, 1912)
(= Beroe shakespeari Benham, 1907 by Bigelow, 1912)
. pandorina (Moser, 1903)
penicillata (Mertens, 1833)
ramosa Komai, 1921
roseus Quoy and Gaimard, 1824 (in Moser 1908)
rufescens (Eschscholtz, 1829)
. Neis Lesson, 1843 or 1829? (this picture of Neis does not show bioluminescence, despite its caption) another image
cordigera Lesson, 1829