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.
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 â 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.
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.
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.
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 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.
Species
There are currently 227 recognized species in this genus:
- Haplochromis acidens <small>Greenwood, 1967</small>
- Haplochromis adolphifrederici <small>Boulenger, 1914</small>
- Haplochromis aelocephalus <small>Greenwood, 1959</small>
- Haplochromis aeneocolor <small>Greenwood, 1973</small>
- Haplochromis akika <small>Lippitsch, 2003</small>
- Haplochromis albertianus <small>Regan, 1929</small>
- Haplochromis altigenis <small>Regan, 1922</small>
- Haplochromis ampullarostratus <small>Schraml, 2004</small>
- Haplochromis angustifrons <small>Boulenger, 1914</small>
- Haplochromis annectidens <small>Trewavas, 1933</small>
- Haplochromis antleter <small>Mietes & F. Witte, 2010</small>
- Haplochromis apogonoides <small>Greenwood, 1967</small>
- Haplochromis arcanus <small>Greenwood & Gee, 1969</small>
- Haplochromis argens <small>de Zeeuw, Westbroek & F. Witte, 2013</small>
- Haplochromis argenteus <small>Regan, 1922</small>
- Haplochromis artaxerxes <small>Greenwood, 1962</small>
- Haplochromis astatodon <small>Regan, 1921</small>
- Haplochromis avium <small>Regan, 1929</small>
- Haplochromis barbarae <small>Greenwood, 1967</small>
- Haplochromis bareli <small>van Oijen, 1991</small>
- Haplochromis bartoni <small>Greenwood, 1962</small>
- Haplochromis bayoni <small>Boulenger, 1909</small>
- Haplochromis beadlei <small>Trewavas, 1933</small>
- Haplochromis bicolor <small>Boulenger, 1906</small>
- Haplochromis bloyeti <small>(Sauvage, 1883)</small>
- Haplochromis boops <small>Greenwood, 1967</small>
- Haplochromis brownae <small>Greenwood, 1962</small>
- Haplochromis bullatus <small>Trewavas, 1938</small>
- Haplochromis bwathondii <small>Niemantsverdriet & F. Witte, 2010</small>
- Haplochromis cassius <small>Greenwood & Barel, 1978</small>
- Haplochromis cavifrons <small>Hilgendorf, 1888</small>
- Haplochromis chilotes <small>Boulenger, 1911</small>
- Haplochromis chlorochrous <small>Greenwood & Gee, 1969</small>
- Haplochromis chromogynos <small>Greenwood, 1959</small>
- Haplochromis chrysogynaion <small>van Oijen, 1991</small>
- Haplochromis cinctus <small>Greenwood & Gee, 1969</small>
- Haplochromis cinereus <small>Boulenger, 1906</small>
- Haplochromis cnester <small>F. Witte & Witte-Maas, 1981</small>
- Haplochromis commutabilis <small>Schraml, 2004</small> (Kachira blue)
- Haplochromis coprologus <small>Niemantsverdriet & F. Witte, 2010</small>
- Haplochromis crassilabris <small>Boulenger, 1906</small>
- Haplochromis crebridens <small>Snoeks, De Vos, Coenen & Thys van den Audenaerde, 1990</small>
- Haplochromis crocopeplus <small>Greenwood & Barel, 1978</small>
- Haplochromis cronus <small>Greenwood, 1959</small>
- Haplochromis cryptodon <small>Greenwood, 1959</small>
- Haplochromis cryptogramma <small>Greenwood & Gee, 1969</small>
- Haplochromis cyaneus <small>Seehausen, Bouton & Zwennes, 1998</small>
- Haplochromis decticostoma <small>Greenwood & Gee, 1969</small>
- Haplochromis degeni <small>Boulenger, 1906</small>
- Haplochromis dentex <small>Regan, 1922</small>
- Haplochromis dichrourus <small>Regan, 1922</small>
- Haplochromis diplotaenia <small>Regan & Trewavas, 1928</small>
- Haplochromis dolichorhynchus <small>Greenwood & Gee, 1969</small>
- Haplochromis dolorosus <small>Trewavas, 1933</small>
- Haplochromis eduardianus <small>Boulenger, 1914</small>
- Haplochromis eduardii <small>Regan, 1921</small>
- Haplochromis elegans <small>Trewavas, 1933</small>
- Haplochromis empodisma <small>Greenwood, 1960</small>
- Haplochromis engystoma <small>Trewavas, 1933</small>
- Haplochromis erythrocephalus <small>Greenwood & Gee, 1969</small>
- Haplochromis erythromaculatus <small>De Vos, Snoeks & Thys van den Audenaerde, 1991</small>
- Haplochromis estor <small>Regan, 1929</small>
- Haplochromis eutaenia <small>Regan & Trewavas, 1928</small>
- Haplochromis exspectatus <small>Schraml, 2004</small>
- Haplochromis fischeri <small>Seegers, 2008</small>
- Haplochromis flavipinnis <small>Boulenger, 1906</small>
- Haplochromis flavus <small>Seehausen, Zwennes & Lippitsch, 1998</small>
- Haplochromis fuelleborni <small>Hilgendorf & Pappenheim, 1903</small> (Fuelleborn's mouthbrooder)
- Haplochromis fuscus <small>Regan, 1925</small>
- Haplochromis fusiformis <small>Greenwood & Gee, 1969</small>
- Haplochromis gigliolii <small>Pfeffer, 1896</small>
- Haplochromis gilberti <small>Greenwood & Gee, 1969</small>
- Haplochromis goldschmidti <small>F. Witte, Westbroek & de Zeeuw, 2013</small>
- Haplochromis gowersii <small>Trewavas, 1928</small>
- Haplochromis gracilior <small>Boulenger, 1914</small> (Torpedostripe haplochromis)
- Haplochromis granti <small>Boulenger, 1906</small>
- Haplochromis graueri <small>Boulenger, 1914</small>
- Haplochromis guiarti <small>Pellegrin, 1904</small>
- Haplochromis harpakteridion <small>van Oijen, 1991</small>
- Haplochromis heusinkveldi <small>F. Wittee & Witte-Maas, 1987</small>
- Haplochromis hiatus <small>Hoogerhoud & F. Witte, 1981</small>
- Haplochromis howesi <small>van Oijen, 1992</small>
- Haplochromis humilior <small>Boulenger, 1911</small>
- Haplochromis humilis <small>Steindachner, 1866</small>
- Haplochromis insidiae <small>Snoeks, 1994</small>
- Haplochromis iris <small>Hoogerhoud & F. Witte, 1981</small>
- Haplochromis ishmaeli <small>Boulenger, 1906</small>
- Haplochromis kamiranzovu <small>Snoeks, Coenen & Thys van den Audenaerde, 1984</small>
- Haplochromis katavi <small>Seegers, 1996</small> (Katavi mouthbrooder)
- Haplochromis katonga <small>Schraml & Tichy, 2010</small>
- Haplochromis katunzii <small>ter Huurne & F. Witte, 2010</small>
- Haplochromis kujunjui <small>van Oijen, 1991</small>
- Haplochromis labiatus <small>Trewavas, 1933</small>
- Haplochromis labriformis <small>Nichols & La Monte, 1938</small>
- Haplochromis lacrimosus <small>Boulenger, 1906</small>
- Haplochromis laparogramma <small>Greenwood & Gee, 1969</small>
- Haplochromis latifasciatus <small>Regan, 1929</small>
- Haplochromis limax <small>Trewavas, 1933</small>
- Haplochromis lividus <small>Greenwood, 1956</small>
- Haplochromis loati <small>Greenwood, 1971</small>
- Haplochromis longirostris <small>Hilgendorf, 1888</small>
- Haplochromis luluae <small>(Fowler, 1930</small>
- Haplochromis luteus <small>Seehausen & Bouton, 1998</small>
- Haplochromis macconneli <small>Greenwood, 1974</small>
- Haplochromis macrognathus <small>Regan, 1922</small>
- Haplochromis macrops <small>Boulenger, 1911</small>
- Haplochromis macropsoides <small>Greenwood, 1973</small>
- Haplochromis maculipinna <small>Pellegrin, 1913</small>
- Haplochromis mahagiensis <small>L. R. David & Poll, 1937</small>
- Haplochromis maisomei <small>van Oijen, 1991</small>
- Haplochromis malacophagus <small>Poll & Damas, 1939</small>
- Haplochromis mandibularis <small>Greenwood, 1962</small>
- Haplochromis martini <small>Boulenger, 1906</small>
- Haplochromis maxillaris <small>Trewavas, 1928</small>
- Haplochromis megalops <small>Greenwood & Gee, 1969</small>
- Haplochromis melanopterus <small>Trewavas, 1928</small>
- Haplochromis melanopus <small>Regan, 1922</small>
- Haplochromis melichrous <small>Greenwood & Gee, 1969</small>
- Haplochromis mentatus <small>Regan, 1925</small>
- Haplochromis mento <small>Regan, 1922</small>
- Haplochromis michaeli <small>Trewavas, 1928</small>
- Haplochromis microchrysomelas <small>Snoeks, 1994</small>
- Haplochromis microdon <small>Boulenger, 1906</small>
- Haplochromis multiocellatus <small>Boulenger, 1913</small>
- Haplochromis mylergates <small>Greenwood & Barel, 1978</small>
- Haplochromis mylodon <small>Greenwood, 1973</small>
- Haplochromis nanoserranus <small>Greenwood & Barel, 1978</small>
- Haplochromis nigrescens <small>Pellegrin, 1909</small>
- Haplochromis nigripinnis <small>Regan, 1921</small>
- Haplochromis nigroides <small>Pellegrin, 1928</small>
- Haplochromis niloticus <small>Greenwood, 1960</small>
- Haplochromis nubilus <small>Boulenger, 1906</small> (Blue Victoria mouthbrooder)
- Haplochromis nuchisquamulatus <small>Hilgendorf, 1888</small>
- Haplochromis nyanzae <small>Greenwood, 1962</small>
- Haplochromis obesus <small>Boulenger, 1906</small>
- Haplochromis obliquidens <small>Hilgendorf, 1888</small>
- Haplochromis obtusidens <small>Trewavas, 1928</small>
- Haplochromis occultidens <small>Snoeks, 1988</small>
- Haplochromis oligolepis <small>Lippitsch, 2003</small>
- Haplochromis olivaceus <small>Snoeks, De Vos, Coenen & Thys van den Audenaerde, 1990</small>
- Haplochromis oregosoma <small>Greenwood, 1973</small>
- Haplochromis orthostoma <small>Regan, 1922</small>
- Haplochromis pachycephalus <small>Greenwood, 1967</small>
- Haplochromis pallidus <small>Boulenger, 1911</small>
- Haplochromis paludinosus <small>(Greenwood, 1980)f</small>
- Haplochromis pancitrinus <small>Mietes & F. Witte, 2010</small>
- Haplochromis pappenheimi <small>Boulenger, 1914</small>
- Haplochromis paradoxus <small>Lippitsch & Kaufman, 2003</small>
- Haplochromis paraguiarti <small>Greenwood, 1967</small>
- Haplochromis paraplagiostoma <small>Greenwood & Gee, 1969</small>
- Haplochromis paropius <small>Greenwood & Gee, 1969</small>
- Haplochromis parorthostoma <small>Greenwood, 1967</small>
- Haplochromis parvidens <small>Boulenger, 1911</small>
- Haplochromis paucidens <small>Regan, 1921</small>
- Haplochromis pellegrini <small>Regan, 1922</small>
- Haplochromis percoides <small>Boulenger, 1906</small>
- Haplochromis perrieri <small>Pellegrin, 1909</small>
- Haplochromis petronius <small>Greenwood, 1973</small>
- Haplochromis pharyngalis <small>Poll & Damas, 1939</small>
- Haplochromis pharyngomylus <small>Regan, 1929</small>
- Haplochromis phytophagus <small>Greenwood, 1966</small>
- Haplochromis piceatus <small>Greenwood & Gee, 1969</small>
- Haplochromis pitmani <small>Fowler, 1936</small>
- Haplochromis placodus <small>Poll & Damas, 1939</small>
- Haplochromis plagiodon <small>Regan & Trewavas, 1928</small>
- Haplochromis plagiostoma <small>Regan, 1922</small>
- Haplochromis plutonius <small>Greenwood & Barel, 1978</small>
- Haplochromis prodromus <small>Trewavas, 1935</small>
- Haplochromis prognathus <small>Pellegrin, 1904</small>
- Haplochromis pseudopellegrini <small>Greenwood, 1967</small>
- Haplochromis ptistes <small>Greenwood & Barel, 1978</small>
- Haplochromis pyrrhocephalus <small>F. Witte & Witte-Maas, 1987</small>
- Haplochromis pyrrhopteryx <small>van Oijen, 1991</small>
- Haplochromis retrodens <small>Hilgendorf, 1888</small>
- Haplochromis riponianus <small>Boulenger, 1911</small>
- Haplochromis rubescens <small>Snoeks, 1994</small>
- Haplochromis rubripinnis <small>Seehausen, Lippitsch & Bouton, 1998</small>
- Haplochromis rudolfianus <small>Trewavas, 1933</small>
- Haplochromis rufus <small>Seehausen & Lippitsch, 1998</small>
- Haplochromis sauvagei <small>Pfeffer, 1896</small>
- Haplochromis saxicola <small>Greenwood, 1960</small>
- Haplochromis scheffersi <small>Snoeks, De Vos & Thys van den Audenaerde, 1987</small>
- Haplochromis schubotzi <small>Boulenger, 1914</small>
- Haplochromis schubotziellus <small>Greenwood, 1973</small>
- Haplochromis serranus <small>Pfeffer, 1896</small>
- Haplochromis serridens <small>Regan, 1925</small>
- Haplochromis simpsoni <small>Greenwood, 1965</small>
- Haplochromis smithii <small>Castelnau, 1861</small>
- Haplochromis snoeksi <small>Wamuini Lunkayilakio & Vreven, 2010</small>
- Haplochromis spekii <small>Boulenger, 1906</small>
- Haplochromis sphex <small>ter Huurne & F. Witte, 2010</small>
- Haplochromis squamipinnis <small>Regan, 1921</small>
- Haplochromis squamulatus <small>Regan, 1922</small>
- Haplochromis sulphureus <small>Greenwood & Barel, 1978</small>
- Haplochromis tanaos <small>van Oijen & F. Witte, 1996</small>
- Haplochromis taurinus <small>Trewavas, 1933</small>
- Haplochromis teegelaari <small>Greenwood & Barel, 1978</small>
- Haplochromis teunisrasi <small>F. Witte & Witte-Maas, 1981</small>
- Haplochromis theliodon <small>Greenwood, 1960</small>
- Haplochromis thereuterion <small>van Oijen & F. Witte, 1996</small>
- Haplochromis thuragnathus <small>Greenwood, 1967</small>
- Haplochromis tridens <small>Regan & Trewavas, 1928</small>
- Haplochromis turkanae <small>Greenwood, 1974</small> (Turkana haplo)
- Haplochromis tyrianthinus <small>Greenwood & Gee, 1969</small>
- Haplochromis ushindi <small>van Oijen, 2004</small>
- Haplochromis vanheusdeni <small>F. D. B. Schedel, Friel & U. K. Schliewen, 2014</small>
- Haplochromis vanoijeni <small>de Zeeuw & F. Witte, 2010</small>
- Haplochromis velifer <small>Trewavas, 1933</small>
- Haplochromis venator <small>Greenwood, 1965</small>
- Haplochromis vicarius <small>Trewavas, 1933</small>
- Haplochromis victoriae <small>Greenwood, 1956</small>
- Haplochromis victorianus <small>Pellegrin, 1904</small>
- Haplochromis vittatus <small>Boulenger, 1901</small>
- Haplochromis vonlinnei <small>van Oijen & de Zeeuw, 2008</small>
- Haplochromis welcommei <small>Greenwood, 1966</small>
- Haplochromis worthingtoni <small>Regan, 1929</small>
- Haplochromis xanthopteryx <small>Seehausen & Bouton, 1998</small>
- Haplochromis xenognathus <small>Greenwood, 1957</small>
- Haplochromis xenostoma <small>Regan, 1922</small>
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:
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:
Some other Pseudocrenilabrinae were also â mainly by early authors â included in Haplochromis, though they are not members of its tribe. These are:
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 <small>Greenwood, 1980</small>
- Cleptochromis <small>Greenwood, 1980</small>
- Enterochromis <small>Greenwood, 1980</small>
- Gaurochromis <small>Greenwood, 1980</small>
- Harpagochromis <small>Greenwood, 1980</small>
- Labrochromis <small>Regan, 1920</small>
- Lipochromis <small>Regan, 1920</small>
- Platytaeniodus <small>Boulenger, 1906</small>
- Prognathochromis <small>Greenwood, 1980</small>
- Ptyochromis <small>Greenwood, 1980</small>
- Psammochromis <small>Greenwood, 1980</small>
- Tridontochromis <small>Greenwood, 1980</small>
- Xystichromis <small>Greenwood, 1980</small>
- Yssichromis <small>Greenwood, 1980</small>
Sometimes other genera listed above are also synonymized.
Footnotes
References
- [2009a]: Valid Haplochromis species. Retrieved 2009-OCT-03.
- [2009b]: Haplochromis taxa. Retrieved 2009-OCT-03.
- (2009): 2009 IUCN Red List of Threatened Species. Version 2009.1. Retrieved 2009-SEP-20.
- (2008): Dietary shift in benthivorous cichlids after the ecological changes in Lake Victoria. Anim. Biol. 58(4): 401-417. <small></small> (HTML abstract)
- (1994): African cichlids II: Cichlids from East Africa. A handbook for their identification, care and breeding. Tetra Press, Germany. <small></small>
- (2001): Classification and Phylogenetic Relationships of African Tilapiine Fishes Inferred from Mitochondrial DNA Sequences. Molecular Phylogenetics and Evolution 20(3): 361âÂÂ374. <small></small>