After Nelson, 1994, and Kinziger, Wood & Neely, 2005 (fig. 1), with Recent species from Froese & Pauly, 2001 [“The Fishbase”]
<==o Cottidae [Icelidae] [incl. Cottocomephoridae, Abyssocottidae, and Comephoridae] (simput, jääsimput; sculpins) [N: 70/300] [FB: 62/210] |?- †Eocottus veronensis M. Eoc. Eu. |?- †Paraperca Olig. Eu. |?- †Cottopsis Olig. SWAs. |?- †Kerocottus Mioc.-Plioc. NA. |?- †Lepidocottus ariens L. Olig. Eu. |?- †Lirosceles Mioc. NA. |?- Alcichthys alcicornis (Herzenstein, 1890) |?-o Antipodocottus [4] | |-- A. elegans Fricke & Brunken, 1984 | |-- A. galatheae Bolin, 1952 | |-- A. megalops DeWitt, 1969 | `-- A. mesembrinus (Fricke & Brunken, 1983) |?- Archaulus biseriatus Gilbert & Burke, 1912 (Scaled sculpin) |?- Argyrocottus zanderi Herzenstein, 1892 |?- Artedielloides auriculatus Soldatov, 1922 |?-o Artediellus [9] | |-- A. atlanticus Jordan & Evermann, 1898 (Atlantic hookear sculpin) | |-- A. camchaticus Gilbert & Burke, 1912 | |--o A. dydymovi Soldatov, 1915 | | |-- A. d. dydymovi Soldatov, 1915 | | `-- A. d. schmidti Soldatov, 1915 | |-- A. gomojunovi Taranetz, 1933 | |-- A. miacanthus Gilbert & Burke, 1912 | |-- A. ochotensis Gilbert & Burke, 1912 | |-- A. pacificus Gilbert, 1896 (Hookhorn sculpin) | |-- A. scaber Knipowitsch, 1907 (Hamecon) | `-- A. uncinatus (Reinhardt, 1835) (Arctic hookear sculpin) |?-o Artedius [5] | |-- A. corallinus (Hubbs, 1926) (Coralline sculpin) | |-- A. fenestralis Jordan & Gilbert, 1883 (Padded sculpin) | |-- A. harringtoni (Starks, 1896) (Scalyhead sculpin) | |-- A. lateralis (Girard, 1854) (Smoothhead sculpin) | `-- A. notospilotus Girard, 1856 (Bonehead sculpin) |?- Ascelichthys rhodorus Jordan & Gilbert, 1880 (Rosylip sculpin) |?- Asemichthys taylori Gilbert, 1912 (Spinynose sculpin) |?-o Astrocottus [3] | |-- A. leprops Bolin, 1936 | |-- A. matsubarae Katayama, 1942 | `-- A. oyamai Watanabe, 1958 |?- Atopocottus tribranchius Bolin, 1936 |?-o Bero [2] | |-- B. elegans (Steindachner, 1881) | `-- B. zanclus Snyder, 1911 |?- Chitonotus pugetensis (Steindachner, 1876) (Roughback sculpin) |?-o Clinocottus [5] | |-- C. acuticeps (Gilbert, 1896) (Sharpnose sculpin) | |-- C. analis (Girard, 1858) (Woolly sculpin) | |-- C. embryum (Jordan & Starks, 1895) (Calico sculpin) | |-- C. globiceps (Girard, 1858) (Mosshead sculpin) | `-- C. recalvus (Greeley, 1899) (Bald sculpin) |?-o Cottiusculus | |-- C. gonez Jordan, 1904 | `-- C. schmidti Jordan & Starks, 1904 |?- Daruma sagamia Jordan & Starks, 1904 |?-o Enophrys | |-- E. bison (Girard, 1854) (Buffalo sculpin) | |-- E. diceraus (Pallas, 1788) (Antlered sculpin) | |-- E. lucasi (Jordan & Gilbert, 1898) (Leister sculpin) | `-- E. taurina Gilbert, 1914 (Bull sculpin) |?-o Furcina [2] | |-- F. ishikawae Jordan & Starks, 1904 | `-- F. osimae Jordan & Starks, 1904 |?-o Gymnocanthus [6] | |-- G. detrisus Gilbert & Burke, 1912 | |-- G. galeatus Bean, 1881 (Armorhead sculpin) | |-- G. herzensteini Jordan & Starks, 1904 | |-- G. intermedius (Temminck & Schlegel, 1843) | |-- G. pistilliger (Pallas, 1814) (Threaded sculpin) | `-- G. tricuspis (Reinhardt, 1830) (Arctic staghorn sculpin) |?-o Hemilepidotus [6] | |-- H. gilberti Jordan & Starks, 1904 (Gilbert's Irish Lord) | |-- H. hemilepidotus (Tilesius, 1811) (Red Irish lord) | |-- H. jordani Bean, 1881 (Yellow Irish lord) | |-- H. papilio (Bean, 1880) (Butterfly sculpin) | |-- H. spinosus Ayres, 1854 (Brown Irish lord) | `-- H. zapus Gilbert & Burke, 1912 (Longfin Irish lord) |?-o Icelinus [9] | |-- I. borealis Gilbert, 1896 (Northern sculpin) | |-- I. burchami Evermann & Goldsborough, 1907 (Dusky sculpin) | |-- I. cavifrons Gilbert, 1890 (Pit-head sculpin) | |-- I. filamentosus Gilbert, 1890 (Threadfin sculpin) | |-- I. fimbriatus Gilbert, 1890 (Fringed sculpin) | |-- I. japonicus Yabe, Tsumura & Katayama, 1980 | |-- I. oculatus Gilbert, 1890 (Frogmouth sculpin) | |-- I. quadriseriatus (Lockington, 1880) (Yellowchin sculpin) | `-- I. tenuis Gilbert, 1890 (Spotfin sculpin) |?-o Icelus [13] | |-- I. bicornis (Reinhardt, 1840) (Twohorn sculpin) | |-- I. canaliculatus Gilbert, 1896 | |-- I. cataphractus (Pavlenko, 1910) | |-- I. euryops Bean, 1890 | |-- I. gilberti Taranetz, 1936 | |-- I. mandibularis Yabe, 1983 | |-- I. ochotensis Schmidt, 1927 | |-- I. perminovi Taranetz, 1936 | |-- I. rastrinoides Taranetz, 1936 | |-- I. spatula Gilbert & Burke, 1912 (Spatulate sculpin) | |-- I. spiniger Gilbert, 1896 (Thorny sculpin) | |-- I. stenosomus Andriashev, 1937 | `-- I. uncinalis Gilbert & Burke, 1912 |?- Jordania zonope Starks, 1895 (Longfin sculpin) |?- Leiocottus hirundo Girard, 1856 (Lavender sculpin) |?-o Megalocottus platycephalus (Pallas, 1814) (Belligerent sculpin) | |-- M. p. platycephalus (Pallas, 1814) (Belligerent sculpin) | `-- M. p. taeniopterus (Kner, 1868) (Southern flathead sculpin) |?- Mesocottus haitej (Dybowski, 1869) |?- Micrenophrys lilljeborgii (Collett, 1875) (Norway bullhead) |?- Microcottus sellaris (Gilbert, 1896) (Brightbelly sculpin) |?-o Myoxocephalus Mioc.-Plioc. R. NA. [12] | |-- M. aenaeus (Mitchill, 1814) (Grubby) | |-- M. brandti (Steindachner, 1867) | |-- M. incitus Watanabe, 1958 | |-- M. jaok (Cuvier, 1829) (Plain sculpin) | |-- M. matsubarai Watanabe, 1958 | |-- M. niger (Bean, 1881) (Warthead sculpin) | |-- M. octodecemspinosus (Mitchill, 1814) (Longhorn sculpin) | |-- M. polyacanthocephalus (Pallas, 1814) (Great sculpin) | |-- M. scorpioides (Fabricius, 1780) (Arctic sculpin) | |-- M. scorpius (Linnaeus, 1758) (isosimppu; Shorthorn sculpin) | |-- M. stelleri Tilesius, 1811 (Steller's sculpin) | `-- M. verrucosus (Bean, 1881) |?-o Ocynectes [2] | |-- O. maschalis Jordan & Starks, 1904 | `-- O. modestus Snyder, 1911 |?-o Oligocottus [5] | |-- O. latifrons (Gilbert & Thompson, 1905) | |-- O. maculosus Girard, 1856 (Tidepool sculpin) | |-- O. rimensis (Greeley, 1899) (Saddleback sculpin) | |-- O. rubellio (Greeley, 1899) (Rosy sculpin) | `-- O. snyderi Greeley, 1898 (Fluffy sculpin) |?- Orthonopias triacis Starks & Mann, 1911 (Snubnose sculpin) |?- Paricelinus hopliticus Eigenmann & Eigenmann, 1889 (Thornback sculpin) |?- Phallocottus obtusus Schultz, 1938 (Spineless sculpin) |?- Phasmatocottus ctenopterygius Bolin, 1936 |?-o Porocottus [8] | |-- P. allisi (Jordan & Starks, 1904) | |-- P. bradfordi Rutter, 1898 (Pored sculpin) | |-- P. camtschaticus (Schmidt, 1916) | |-- P. coronatus Yabe, 1992 | |-- P. japonicus Schmidt, 1935 | |-- P. minutus (Pallas, 1814) | |-- P. quadrifilis Gill, 1859 | `-- P. tentaculatus (Kner, 1868) |?-o Pseudoblennius [4] | |-- P. cottoides (Richardson, 1848) | |-- P. marmoratus (Döderlein, 1884) | |-- P. percoides Günther, 1861 | `-- P. zonostigma Jordan & Starks, 1904 |?-o Radulinus [3] | |-- R. asprellus Gilbert, 1890 (Slim sculpin) | |-- R. boleoides Gilbert, 1898 (Darter sculpin) | `-- R. vinculus Bolin, 1950 (Smooth-gum sculpin) |?- Rastrinus scutiger (Bean, 1890) |?- Ricuzenius nudithorax Bolin, 1936 |?-o Ruscarius [2] | |-- R. creaseri (Hubbs, 1926) (Roughcheek sculpin) | `-- R. meanyi Jordan & Starks, 1895 (Puget Sound sculpin) |?- Scorpaenichthys marmoratus Girard, 1854 (Cabezon) |?-o Sigmistes [2] | |-- S. caulias Rutter, 1898 (Kelp sculpin) | `-- S. smithi Schultz, 1938 (Arched sculpin) |?-o Stelgistrum [2] | |-- S. beringianum Gilbert & Burke, 1912 | `-- S. stejnegeri Jordan & Gilbert, 1898 |?- Stlegicottus xenogrammus Bolin, 1936 |?-o Stlengis [3] | |-- S. distoechus Bolin, 1936 | |-- S. misakia (Jordan & Starks, 1904) | `-- S. osensis Jordan & Starks, 1904 |?- Synchirus gilli Bean, 1890 (Manacled sculpin) |?- Taurocottus bergi Soldatov & Pavlenko, 1915 |?- Taurulus bubalis (Euphrasen, 1786) [Ceratocottus bubalis?] (piikkisimppu; Longspined bullhead) |?- Thyriscus anoplus Gilbert & Burke, 1912 |?- Trachidermus fasciatus Heckel, 1837 [Trachidermis] |?- Trichocottus [not recognized by the Fishbase] |?-o Triglops [9] | |-- T. forficatus (Gilbert, 1896) (Scissortail sculpin) | |-- T. jordani (Schmidt, 1903) (Jordan's sculpin) | |-- T. macellus (Bean, 1884) (Roughspine sculpin) | |-- T. metopias Gilbert & Burke, 1912 (Alaskan sculpin) | |-- T. murrayi Günther, 1888 (Moustache sculpin) | |-- T. nybelini Jensen, 1944 (Bigeye sculpin) | |-- T. pingelii Reinhardt, 1837 (Ribbed sculpin) | |-- T. scepticus Gilbert, 1896 (Spectacled sculpin) | `-- T. xenostethus Gilbert, 1896 (Scaly breasted sculpin) |?-o Triglopsis [2] | |-- T. quadricornis (Linnaeus, 1758) [Myoxocephalus quadricornis?] (härkäsimppu; Fourhorn sculpin) | `-- T. thompsonii Girard, 1851 (Deepwater sculpin) |?-o Vellitor [2] | |-- V. centropomus (Richardson, 1848) | `-- V. minutus Iwata, 1983 |?-o Zesticelus | |-- Z. ochotensis Yabe, 1995 | `-- Z. profundorum (Gilbert, 1896) (Flabby sculpin) `--+--+-- Leptocottus armatus Girard, 1854 (Pacific staghorn sculpin) | `-- Cottus kazika Jordan & Starks, 1904 `?=o Cottus Linnaeus, 1758 sensu lato R. As., Mioc. NZ., L.Olig.-R. Eu., Mioc. Plioc.-R. NA. (kivisimput) [34] |--+--+-- Cottus pollux Günther, 1873 (Japanese fluvial sculpin) | | `-- Cottus reinii Hilgendorf, 1879 | `--+--+-- Cottus poecilopus volki (Taranetz, 1933) | | `-- Cottus hangiongensis Mori, 1930 | `--o Cephalocottus -clade | |-- Cottus nozawae Snyder, 1911 | `-- Cottus amblystomopsis Schmidt, 1904 (Sakhalin sculpin) `--+-- Cottus poecilopus poecilopus Heckel, 1837 (Alpine bullhead) `--+--o Cottopsis -clade | |-- Cottus aleuticus Gilbert, 1896 (Coastrange sculpin) | |--+-- Cottus pitensis Bailey & Bond, 1963 (Pit sculpin) | | |-- Cottus perplexus Gilbert & Evermann, 1894 (Reticulate sculpin) | | `--+-- Cottus asper Richardson, 1836 (Prickly sculpin) | | `--+-- Cottus gulosus (Girard, 1854) (Riffle sculpin) | | `-- Cottus marginatus (Bean, 1881) (Margined sculpin) | `--+-- Cottus princeps Gilbert, 1898 (Klamath Lake sculpin) | `--+-- Cottus asperrimus Rutter, 1908 (Rough sculpin) | `--o Cottus klamathensis Gilbert, 1898 (Marbled sculpin) | |-- Cottus klamathensis polyporus | `--+-- Cottus klamathensis macrops | `-- Cottus klamathensis klamathensis Gilbert, 1898 `--+--o Cottus Linnaeus, 1758 sensu stricto [Cottus -clade sensu Kinziger, Wood & Neely, 2005] | |-- Cottus ricei (Nelson, 1876) (Spoonhead sculpin) | `--+-- Cottus sibiricus Kessler, 1899 (Siberian sculpin) | `--+-- Cottus ferrugineus Heckel & Kner, 1858 | `--o Cottus gobio Linnaeus, 1758 (kivisimppu; Bullhead) | |-- Cottus gobio gobio Linnaeus, 1758 (kivisimppu; Bullhead) | `-- Cottus gobio jaxartensis Berg, 1916 `--+--o “Uranidea” clade | |-- Cottus confusus Bailey & Bond, 1963 (Shorthead sculpin) | |--+-- Cottus greenei (Gilbert & Culver, 1898) (Shoshone sculpin) | | `--+-- Cottus leiopomus Gilbert & Evermann, 1894 (Wood River sculpin) | | `-- Cottus beldingii Eigenmann & Eigenmann, 1891 (Paiute sculpin) | `--+--+-- Cottus baileyi Robins, 1961 (Black sculpin) | | |-- Cottus paulus | | |--+-- Cottus sp. “smoky” | | | `-- Coppus sp. “Clinch” | | |--+-- Cottus carolinae (Gill, 1861) [1] (Banded sculpin) | | | `-- Cottus sp. “Holston” | | `--+-- Cottus sp. “Bluestone” | | `-- Cottus sp. cf. Cottus carolinae (Gill, 1861) [2] (Banded sculpin) | `--+--+--+-- Cottus rhotheus (Smith, 1882) (Torrent sculpin) | | | `-- Cottus cognatus Richardson, 1836 (Slimy sculpin) | | `--+-- Cottus bairdii punctulatus | | |--+-- C. extensus Bailey & Bond, 1963 (Bear Lake sculpin) | | | `-- Cottus bairdii semiscaber | | `--+-- Cottus hubbsi | | `-- Cottus bendirei | `--+--+-- Cottus hypselurus Robins & Robison, 1985 (Ozark sculpin) | | `-- Cottus bairdii bairdii Girard, 1850 [3&4] (Mottled sculpin) | `--+-- Cottus cognatus Richardson, 1836 [3] (Slimy sculpin) | |--+-- Cottus bairdii bairdii Girard, 1850 [1] | | `--+-- Cottus sp. cf. Cottus cognatus Richardson, 1836 [1] “checkered” | | `-- Cottus girardi Robins, 1961 (Potomac sculpin) | `--+-- C. caeruleomentum Kinziger, Raesly & Neely, 2000 (Blue Ridge sculpin) | `--+-- Cottus bairdii bairdii Girard, 1850 [2] | `-- Cottus bairdii kumlieni [1&2] `--o “Baikalian clade” |--o Paracottus knerii (Dybowski, 1874) (Stone sculpin) [paraphyletic Cottocomephoridae] | |-- P. k. knerii (Dybowski, 1874) (Stone sculpin) | `-- P. k. putorania Koryakov and Sidelov, 1976 |--+-- Leocottus kessleri | `--o Cottocomephorus [paraphyletic Cottocomephoridae] | |-- C. grewingkii (Dybowski, 1874) (Baikal yellowfin) | `-- C. inermis (Yakovlev, 1890) (Longfin Baikal sculpin) |--o Procottus [Metacottus] [paraphyletic Abyssocottidae] | |?- P. gurwici (Taliev, 1946) | |-- P. jeittelesii (Dybowski, 1874) (Red sculpin) | `-- P. major Taliev, 1949 `--+--o Comephorus [Comephoridae] (rasvasimput) | |-- C. baikalensis (Pallas, 1776) (Big Baikal oilfish) | `-- C. dybowskii Korotneff, 1904 (Little Baikal oilfish) `--+-- Cyphocottus eurystomus `--+?- Abyssocottus elochini Taliev, 1955 [paraphyletic Abyssocottidae] |?- Abyssocottus werestschagini Taliev, 1935 [paraphyletic Abyssocottidae] |--+--o Asprocottus [paraphyletic Abyssocottidae] | | |?- A. abyssalis Taliev, 1955 | | |?- A. parmiferus Taliev, 1955 | | |--+-- A. minor | | | `-- A. pulcher Taliev, 1955 | | `--+-- A. herzensteini Berg, 1906 (Herzenstein's rough sculpin) | | `-- A. platycephalus Taliev, 1955 | `--+-- Cottinella boulengeri (Berg, 1906) (Short-headed sculpin) [paraphyletic Abyssocottidae] | `-- Abyssocottus korotneffi Berg, 1906 [paraphyletic Abyssocottidae] `--+-- Abyssocottus gibbosus Berg, 1906 [paraphyletic Abyssocottidae] `--+?-o Batrachocottus [paraphyletic Cottocomephoridae] | |-- B. baicalensis (Bybowski, 1874) (Bighead sculpin) | |-- B. multiradiatus Berg, 1907 | `-- B. nikolskii (Berg, 1900) (Fat sculpin) `--o Limnocottus [incl. Neocottus??] [paraphyletic Abyssocottidae] |?- L. eurystomus (Taliev, 1955) |?- L. kozovi Taliev, 1946 (Vitim sculpin) |?- L. megalops (Gratzianov, 1902) |--+-- L. godlewskii (Dybowski, 1874) | `-- L. griseus (Taliev, 1948) `--+-- L. bergianus Taliev, 1935 `-- L. pallidus Taliev, 1948
Reference(s):
- Carroll, R. L., 1988: Vertebrate paleontology and evolution.
–W. H. Freeman and company, New York, 1988, 698 - Carroll, R. L., 1988: Appendix. 594-648
in Carroll, R. L., 1988: Vertebrate paleontology and evolution.
–W. H. Freeman and company, New York, 1988, 698 - Frickhinger, K. A., 1995: Fossil Atlas – Fishes.
–Mergus – Publishers for Natural History and Pet Books, Hans A.Baensch, Malle, Germany, 1995, pp. 1-1088 - Froese, R. & Pauly, D. (eds), 2001: FishBase.
–iNet: www.fishbase.org - Kinziger, A. P., Raesly, R. L. & Neely, D. A., 2000: New species of Cottus (Teleostei: Cottidae) from the middle Atlantic eastern United States.
–Copeia: Vol. 100, #4, pp. 1007-1018 - Kinziger, A. P., Wood, R. M. & Neely, D. A., 2005: Molecular systematics of the genus Cottus (Scorpaeniformes: Cottidae).
–Copeia: Vol. 105, #2, pp. 303-311 - Lahti, S., Malmström, K. K., Koli, L., Leikola, A., Syrjämäki, J. & Lahti, J., 1980: Zoo, Suuri Eläinkirja 5: Kalat, sammakkoeläimet, matelijat.
–Werner Söderström Osakeyhtiö, Porvoo-Helsinki-Juva, 1980.
Original work [in French]: Beauté Du Monde Animal: IX Reptiles/Amphibiens & X Poissons.
–Rizzoli Editore, Milano, 1968 - Nelson, J. S., 1994: Fishes of the world.
–John Wiley & Sons Inc., New York, 1994, xx-600