Haplochromis
Haplochromis | |
---|---|
Scientific classification | |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Actinopterygii |
Order: | Cichliformes |
Family: | Cichlidae |
Tribe: | Haplochromini |
Genus: | Haplochromis Hilgendorf, 1888 |
Type species | |
Chromis obliquidens Hilgendorf, 1888
| |
Synonyms | |
Many, see text |
Haplochromis is a ray-finned fish genus in the family Cichlidae. It has been used as the default "wastebin taxon" for Pseudocrenilabrinae cichlids of the East African Rift, and as such became the "largest" fish "genus". Many of these cichlids are popular aquarium fishes; like similar Haplochromini they are known as "haplos", "happies" or "haps" among aquarium enthusiasts.[1]
The genus was established by F.M. Hilgendorf in 1888. It was originally conceived as a subgenus of A.C.L.G. Günther's "Chromis", at that time an even larger "wastebin genus" for Pseudocrenilabrinae cichlids. The type species of Hilgendorf was H. obliquidens. "Chromis" of Günther turned out to be a junior homonym of G. Cuvier's ocean fish genus Chromis, already established in 1814, and was abolished. As the years went by, other genera of (mostly) Haplochromini were lumped with and split again from Haplochromis, and the final delimitation of the clade around H. obliquidens is not yet done.
Extinction crisis in Lake Victoria[]
The introduction of Nile perch (Lates niloticus) to Lake Victoria after 1954 severely upset the lake's ecosystem. By the late 1970s, the perch's population was approaching carrying capacity, and the smaller cichlids were fair game for the huge carnivorous Lates, the Nile tilapia (Oreochromis niloticus), an adaptable generalist, was also introduced and competed with the often specialized endemic cichlids of the lake for food and other resources. When the Nile perch stocks finally declined again in the late 1980s, an estimated 200 Haplochromini species (mostly Haplochromis) may have become extinct – many of these had only been scientifically described a few years before their demise, and additional ones were only known or suspected to exist, but never properly studied or described.[2]
While the stocks of those species that survived are in many cases recovering, the lake ecosystem has changed irrevocably. The entire trophic web has been observed to be upset. But still, evolution runs its course: those Lake Victoria Haplochromis species that still exist are in many cases adapting to new food sources, and in time, speciation is likely to set in and produce a new adaptive radiation of these fishes. Until then, however, the ecological balance of the lake is still on the brink, and many of the cichlids that survived the peak population of Lates are still critically endangered and close to extinction.[3]
Systematics and taxonomy[]
Haplochromis is the type genus of the tribe Haplochromini. Most of the tribe's members were at one time or another included in the present genus, but in many cases this was only temporary. Around the year 1900, as well 100 years later, the trend was to split up the genus; especially in the mid-20th century, on the other hand, most authors lumped any and all Haplochromini that were not conspicuously distinct in the type genus.[4]
While a number of African Rift Valley cichlids are certainly very close relatives of H. obliquidens, the type species of the present genus, it is not very clear where to draw the boundary of Haplochromis with regard to its relatives. Still, several genera are now recognized as distinct by many authors and scientific databases, such as FishBase (see below); in particular the Haplochromini from Lake Tanganyika and Lake Malawi are usually removed from Haplochromis. The genus delimitation in the entire tribe remains badly resolved, however, and further changes in taxonomy are likely in the future. In particular, between Haplochromis, Astatotilapia and Thoracochromis, species have been moved to and from over the years. The habit of Pseudocrenilabrinae to hybridize is hampering molecular phylogenetic studies based on mtDNA alone, while trophic morphs of a single species may appear to be distinct "species" if they are not phylogenetically studied. Several proposed genera are again included in Haplochromis at present, but it cannot be ruled out that some of these will eventually be recognized as valid again.[5]
Species[]
There are currently 229 recognized species in this genus:
- Haplochromis acidens Greenwood, 1967
- Haplochromis adolphifrederici Boulenger, 1914
- Haplochromis aelocephalus Greenwood, 1959
- Haplochromis aeneocolor Greenwood, 1973
- , 2003
- Regan, 1929
- Regan, 1922
- , 2004
- Haplochromis angustifrons Boulenger, 1914
- Haplochromis annectidens Trewavas, 1933
- & , 2010 [6]
- Haplochromis apogonoides Greenwood, 1967
- Haplochromis arcanus Greenwood & , 1969
- Haplochromis argens , & , 2013
- Haplochromis argenteus Regan, 1922
- Haplochromis artaxerxes Greenwood, 1962
- Regan, 1921
- Regan, 1929
- Haplochromis azureus & , 1998
- Haplochromis barbarae Greenwood, 1967
- , 1991
- Haplochromis bartoni Greenwood, 1962
- Haplochromis bayoni Boulenger, 1909
- Haplochromis beadlei Trewavas, 1933
- Haplochromis bicolor Boulenger, 1906
- Haplochromis boops Greenwood, 1967
- Haplochromis brownae Greenwood, 1962
- Trewavas, 1938
- & , 2010 [6]
- Haplochromis cassius Greenwood & , 1978
- Haplochromis cavifrons Hilgendorf, 1888
- Haplochromis chilotes Boulenger, 1911
- Greenwood & , 1969
- Haplochromis chromogynos Greenwood, 1959
- , 1991
- Greenwood & , 1969
- Boulenger, 1906
- & , 1981
- , 2004 (Kachira blue)
- & , 2010 [6]
- Haplochromis crassilabris Boulenger, 1906
- Haplochromis crebridens , , & , 1990
- Greenwood & , 1978
- Greenwood, 1959
- Haplochromis cryptodon Greenwood, 1959
- Greenwood & , 1969
- Haplochromis cyaneus , & , 1998
- Haplochromis decticostoma Greenwood & , 1969
- Haplochromis degeni Boulenger, 1906
- Haplochromis dentex Regan, 1922
- Haplochromis dichrourus Regan, 1922
- Regan & Trewavas, 1928
- Greenwood & , 1969
- Trewavas, 1933
- Haplochromis eduardianus Boulenger, 1914
- Regan, 1921
- Haplochromis elegans Trewavas, 1933
- Greenwood, 1960
- Trewavas, 1933
- Greenwood & , 1969
- Haplochromis erythromaculatus , & , 1991
- Haplochromis estor Regan, 1929
- Regan & Trewavas, 1928
- , 2004
- Seegers, 2008
- Haplochromis flavipinnis Boulenger, 1906
- Haplochromis flavus , & , 1998
- Hilgendorf & , 1903 (Fuelleborn's mouthbrooder)
- Regan, 1925
- Greenwood & , 1969
- Haplochromis gigas & , 1998
- Pfeffer, 1896
- Haplochromis gilberti Greenwood & , 1969
- Haplochromis goldschmidti , & , 2013
- Haplochromis gowersii Trewavas, 1928
- Haplochromis gracilior Boulenger, 1914 (Torpedostripe haplochromis)
- Haplochromis granti Boulenger, 1906
- Haplochromis graueri Boulenger, 1914
- Haplochromis greenwoodi & , 1998
- Haplochromis guiarti Pellegrin, 1904
- , 1991
- Haplochromis heusinkveldi & , 1987
- & , 1981
- Haplochromis howesi , 1992
- Boulenger, 1911
- Steindachner, 1866
- Haplochromis igneopinnis & , 1998
- Haplochromis insidiae , 1994
- & , 1981
- Haplochromis ishmaeli Boulenger, 1906
- Haplochromis kamiranzovu , & , 1984
- Haplochromis katavi Seegers, 1996 (Katavi mouthbrooder)
- & Tichy, 2010
- & , 2010 [6]
- , 1991
- Haplochromis labiatus Trewavas, 1933
- Nichols & , 1938
- Boulenger, 1906
- Haplochromis laparogramma Greenwood & , 1969
- Haplochromis latifasciatus Regan, 1929
- Haplochromis limax Trewavas, 1933
- Haplochromis lividus Greenwood, 1956
- Greenwood, 1971
- Haplochromis longirostris Hilgendorf, 1888
- Haplochromis luteus & , 1998
- Haplochromis macconneli Greenwood, 1974
- Haplochromis macrocephalus & , 1998
- Haplochromis macrognathus Regan, 1922
- Boulenger, 1911
- Haplochromis macropsoides Greenwood, 1973
- Haplochromis maculipinna Pellegrin, 1913
- & Poll, 1937
- , 1991
- Poll & , 1939
- Haplochromis mandibularis Greenwood, 1962
- Haplochromis martini Boulenger, 1906
- Haplochromis maxillaris Trewavas, 1928
- Haplochromis mbipi & , 1998
- Haplochromis megalops Greenwood & , 1969
- Haplochromis melanopterus Trewavas, 1928
- Regan, 1922
- Greenwood & , 1969
- Regan, 1925
- Haplochromis mento Regan, 1922
- Haplochromis michaeli Trewavas, 1928
- Haplochromis microchrysomelas , 1994
- Haplochromis microdon Boulenger, 1906
- Boulenger, 1913
- Haplochromis mylergates Greenwood & , 1978
- Haplochromis mylodon Greenwood, 1973
- Haplochromis nanoserranus Greenwood & , 1978
- Haplochromis nigrescens Pellegrin, 1909
- Haplochromis nigricans Boulenger, 1906
- Haplochromis nigripinnis Regan, 1921
- Haplochromis nigroides Pellegrin, 1928
- Greenwood, 1960
- Haplochromis nubilus Boulenger, 1906 (Blue Victoria mouthbrooder)
- Haplochromis nuchisquamulatus Hilgendorf, 1888
- Haplochromis nyanzae Greenwood, 1962
- Haplochromis nyererei & , 1985
- Boulenger, 1906
- Haplochromis obliquidens Hilgendorf, 1888
- Haplochromis obtusidens Trewavas, 1928
- Haplochromis occultidens , 1988
- , 2003
- Haplochromis olivaceus , , & , 1990
- Haplochromis omnicaeruleus & , 1998
- Haplochromis oregosoma Greenwood, 1973
- Haplochromis orthostoma Regan, 1922
- Haplochromis pachycephalus Greenwood, 1967
- Boulenger, 1911
- & , 2010 [6]
- Haplochromis pappenheimi Boulenger, 1914
- Haplochromis paradoxus & Kaufman, 2003
- Haplochromis paraguiarti Greenwood, 1967
- Haplochromis paraplagiostoma Greenwood & , 1969
- Haplochromis paropius Greenwood & , 1969
- Haplochromis parorthostoma Greenwood, 1967
- Boulenger, 1911
- Haplochromis paucidens Regan, 1921
- Regan, 1922
- Haplochromis percoides Boulenger, 1906
- Haplochromis perrieri Pellegrin, 1909
- Haplochromis petronius Greenwood, 1973
- Poll & , 1939
- Haplochromis pharyngomylus Regan, 1929
- Haplochromis phytophagus Greenwood, 1966
- Haplochromis piceatus Greenwood & , 1969
- Fowler, 1936
- Poll & , 1939
- Haplochromis plagiodon Regan & Trewavas, 1928
- Haplochromis plagiostoma Regan, 1922
- Greenwood & , 1978
- Trewavas, 1935
- Haplochromis prognathus Pellegrin, 1904
- Haplochromis pseudopellegrini Greenwood, 1967
- Greenwood & , 1978
- Haplochromis pundamilia & , 1998
- Haplochromis pyrrhocephalus & , 1987
- , 1991
- Haplochromis retrodens Hilgendorf, 1888
- Haplochromis riponianus Boulenger, 1911
- Haplochromis rubescens , 1994
- Haplochromis rubripinnis , & , 1998
- Haplochromis rudolfianus Trewavas, 1933
- Haplochromis rufocaudalis & , 1998
- Haplochromis rufus & , 1998
- Haplochromis sauvagei Pfeffer, 1896
- Haplochromis saxicola Greenwood, 1960
- Haplochromis scheffersi , & , 1987
- Haplochromis schubotzi Boulenger, 1914
- Haplochromis schubotziellus Greenwood, 1973
- Haplochromis serranus Pfeffer, 1896
- Regan, 1925
- Haplochromis simotes Boulenger, 1911
- Haplochromis simpsoni Greenwood, 1965
- Castelnau, 1861
- & , 2010 [7]
- Haplochromis spekii Boulenger, 1906
- & , 2010 [6]
- Haplochromis squamipinnis Regan, 1921
- Regan, 1922
- Greenwood & , 1978
- & , 1996
- Haplochromis taurinus Trewavas, 1933
- Haplochromis teegelaari Greenwood & , 1978
- & , 1981
- Greenwood, 1960
- & , 1996
- Haplochromis thuragnathus Greenwood, 1967
- Regan & Trewavas, 1928
- Haplochromis turkanae Greenwood, 1974 (Turkana haplo)
- Greenwood & , 1969
- , 2004
- , & , 2014 [8]
- & , 2010 [6]
- Haplochromis velifer Trewavas, 1933
- Haplochromis venator Greenwood, 1965
- Trewavas, 1933
- Haplochromis victoriae Greenwood, 1956
- Haplochromis victorianus Pellegrin, 1904
- Haplochromis vittatus Boulenger, 1901
- Haplochromis vonlinnei & , 2008
- Haplochromis welcommei Greenwood, 1966
- Haplochromis worthingtoni Regan, 1929
- Haplochromis xanthopteryx & , 1998
- Haplochromis xenognathus Greenwood, 1957
- Haplochromis xenostoma Regan, 1922
Undescribed species[]
These populations are typically referred to by the names they have in the aquarium fish trade. A number of them are likely to represent undescribed distinct species; others might just be subspecies or color morphs. Whether they all belong in Haplochromis is, of course, doubtful. Some of these populations are:
- Haplochromis aff. bloyeti[verification needed]
- Haplochromis sp. 'Amboseli'
- Haplochromis sp. 'backflash cryptodon'[verification needed]
- Haplochromis sp. 'black cryptodon'[verification needed]
- Haplochromis sp. 'Chala'
- Haplochromis sp. 'frogmouth'[verification needed]
- Haplochromis sp. 'Kyoga flameback'[verification needed]
- Haplochromis sp. 'long snout'[verification needed]
- Haplochromis sp. 'Migori'
- Haplochromis sp. 'parvidens-like'[verification needed]
- Haplochromis sp. 'ruby'
- Haplochromis sp. 'Rusinga oral sheller'[verification needed]
- Haplochromis sp. 'rainbow sheller'[verification needed]
- Haplochromis sp. 'small obesoid'[verification needed]
Formerly in Haplochromis[]
Among other genera of Haplochromini that were formerly included here, many are small or monotypic. The distinctness of these is highly doubtful, as they may just be distinct lineages of Haplochromis or other haplochromines. That nonwithstanding, Haplochromini genera to which some former "Haplochromis" have been removed are in particular:[9]
- Astatoreochromis (3 species)
- Astatotilapia (8 described species)
- Aulonocara (22 species)
- Buccochromis (7 species)
- Caprichromis (2 species)
- Champsochromis (2 species)
- Cheilochromis (monotypic)
- Chetia (6 species)
- Copadichromis (25 described species)
- Ctenochromis (5 living and 1 extinct described species)
- Ctenopharynx (3 species)
- Cyrtocara (monotypic)
- Dimidiochromis (4 species)
- Eclectochromis (2 species)
- Fossorochromis (monotypic)
- Hemitaeniochromis (monotypic)
- Lethrinops (24 species)
- Maylandia/Metriaclima (25 species)
- Mchenga (6 species)
- Melanochromis (27 species)
- Mylochromis (21 species)
- Naevochromis (monotypic)
- Nimbochromis (5 species)
- Nyassachromis (8 species)
- Orthochromis (14 species)
- Otopharynx (13 species)
- Pharyngochromis (2 species)
- Placidochromis (43 species)
- Protomelas (15 species)
- Pundamilia (4 species)
- Pseudocrenilabrus (3 species)
- Sargochromis (8 species)
- Schwetzochromis (monotypic)
- Sciaenochromis (4 species)
- Stigmatochromis (4 species)
- Taeniochromis (monotypic)
- Taeniolethrinops (4 species)
- Thoracochromis (12 species)
- Tramitichromis (5 species)
- Trematocranus (3 species)
- Tyrannochromis (4 species)
Some other Pseudocrenilabrinae were also – mainly by early authors – included in Haplochromis, though they are not members of its tribe. These are:[4]
- Anomalochromis thomasi of the Hemichromini
- Altolamprologus compressiceps of the Lamprologini
- of the Tilapiini
Synonyms of Haplochromis[]
With all the taxonomic and systematic confusion affecting Haplochromis and its allies, it is hardly surprising that the genus has a large number of junior synonyms. Most referred to small or monotypic genera that were once considered distinct, but are now included in Haplochromis again, if only to wait for a major review of their status. Synonyms are:
- Allochromis Greenwood, 1980
- Cleptochromis Greenwood, 1980
- Enterochromis Greenwood, 1980
- Gaurochromis Greenwood, 1980
- Harpagochromis Greenwood, 1980
- Labrochromis Regan, 1920
- Lipochromis Regan, 1920
- Platytaeniodus Boulenger, 1906
- Prognathochromis Greenwood, 1980
- Ptyochromis Greenwood, 1980
- Psammochromis Greenwood, 1980
- Tridontochromis Greenwood, 1980
- Xystichromis Greenwood, 1980
- Yssichromis Greenwood, 1980
Sometimes other genera listed above are also synonymized.
Footnotes[]
- ^ Linke & Staeck (1994)
- ^ Linke & Staeck (1994), Kishe-Machumu et al. (2008), IUCN (2009)
- ^ Kishe-Machumu et al. (2008), IUCN (2009)
- ^ a b FishBase [2009b]
- ^ Nagl et al. (2001), FishBase [2009b]
- ^ a b c d e f g de Zeeuw, Mietes, Niemantsverdriet, ter Huurne & Witte (2010). "Seven new species of detritivorous and phytoplanktivorous haplochromines from Lake Victoria". Zoologische Mededelingen. 84: 201–250.
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: CS1 maint: uses authors parameter (link) - ^ Wamuini Lunkayilakio; Vreven (2010). "'Haplochromis' snoeksi, a new species from the Inkisi River basin, Lower Congo (Perciformes: Cichlidae)" (PDF). . 21 (3): 279–287.
- ^ Schedel, F. D. B., Friel, J. P. & Schliewen, U. K. (2014): Haplochromis vanheusdeni, a new haplochromine cichlid species from the Great Ruaha River drainage, Rufiji basin, Tanzania (Teleostei, Perciformes, Cichlidae). Spixiana, 37 (1): 135-149.
- ^ Linke & Staeck (1994), FishBase [2009b]
References[]
Wikimedia Commons has media related to Haplochromis. |
- FishBase [2009a]: Valid Haplochromis species. Retrieved 2009-OCT-03.
- FishBase [2009b]: Haplochromis taxa. Retrieved 2009-OCT-03.
- International Union for the Conservation of Nature and Natural Resources (IUCN) (2009): 2009 IUCN Red List of Threatened Species. Version 2009.1. Retrieved 2009-SEP-20.
- Kishe-Machumu, Mary; Witte,Frans & Wanink, Jan H. (2008): Dietary shift in benthivorous cichlids after the ecological changes in Lake Victoria. Anim. Biol. 58(4): 401-417. doi:10.1163/157075608X383700 (HTML abstract)
- Linke, H. & Staeck, L. (1994): African cichlids II: Cichlids from East Africa. A handbook for their identification, care and breeding. Tetra Press, Germany. ISBN 1-56465-167-3
- Nagl, Sandra; Tichy, Herbert; Mayer, Werner E.; Samonte, Irene E.; McAndrew, Brendan J. & Klein, Jan (2001): Classification and Phylogenetic Relationships of African Tilapiine Fishes Inferred from Mitochondrial DNA Sequences. Molecular Phylogenetics and Evolution 20(3): 361–374. doi:10.1006/mpev.2001.0979
- Haplochromis
- Cichlid fish of Africa
- Cichlid genera
- Taxa named by Franz Martin Hilgendorf