Haplochromis

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 – 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 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.
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
Haplochromis akika Lippitsch, 2003
Haplochromis albertianus Regan, 1929
Haplochromis altigenis Regan, 1922
Haplochromis ampullarostratus Schraml, 2004
Haplochromis angustifrons Boulenger, 1914
Haplochromis annectidens Trewavas, 1933
Haplochromis antleter Mietes & F. Witte, 2010
Haplochromis apogonoides Greenwood, 1967
Haplochromis arcanus Greenwood & Gee, 1969
Haplochromis argens de Zeeuw, Westbroek & F. Witte, 2013
Haplochromis argenteus Regan, 1922
Haplochromis artaxerxes Greenwood, 1962
Haplochromis astatodon Regan, 1921
Haplochromis avium Regan, 1929
Haplochromis azureus Seehausen & Lippitsch, 1998
Haplochromis barbarae Greenwood, 1967
Haplochromis bareli van Oijen, 1991
Haplochromis bartoni Greenwood, 1962
Haplochromis bayoni Boulenger, 1909
Haplochromis beadlei Trewavas, 1933
Haplochromis bicolor Boulenger, 1906
Haplochromis bloyeti (Sauvage, 1883)
Haplochromis boops Greenwood, 1967
Haplochromis brownae Greenwood, 1962
Haplochromis bullatus Trewavas, 1938
Haplochromis bwathondii Niemantsverdriet & F. Witte, 2010
Haplochromis cassius Greenwood & Barel, 1978
Haplochromis cavifrons Hilgendorf, 1888
Haplochromis chilotes Boulenger, 1911
Haplochromis chlorochrous Greenwood & Gee, 1969
Haplochromis chromogynos Greenwood, 1959
Haplochromis chrysogynaion van Oijen, 1991
Haplochromis cinctus Greenwood & Gee, 1969
Haplochromis cinereus Boulenger, 1906
Haplochromis cnester F. Witte & Witte-Maas, 1981
Haplochromis commutabilis Schraml, 2004 (Kachira blue)
Haplochromis coprologus Niemantsverdriet & F. Witte, 2010
Haplochromis crassilabris Boulenger, 1906
Haplochromis crebridens Snoeks, De Vos, Coenen & Thys van den Audenaerde, 1990
Haplochromis crocopeplus Greenwood & Barel, 1978
Haplochromis cronus Greenwood, 1959
Haplochromis cryptodon Greenwood, 1959
Haplochromis cryptogramma Greenwood & Gee, 1969
Haplochromis cyaneus Seehausen, Bouton & Zwennes, 1998
Haplochromis decticostoma Greenwood & Gee, 1969
Haplochromis degeni Boulenger, 1906
Haplochromis dentex Regan, 1922
Haplochromis dichrourus Regan, 1922
Haplochromis diplotaenia Regan & Trewavas, 1928
Haplochromis dolichorhynchus Greenwood & Gee, 1969
Haplochromis dolorosus Trewavas, 1933
Haplochromis eduardianus Boulenger, 1914
Haplochromis eduardii Regan, 1921
Haplochromis elegans Trewavas, 1933
Haplochromis empodisma Greenwood, 1960
Haplochromis engystoma Trewavas, 1933
Haplochromis erythrocephalus Greenwood & Gee, 1969
Haplochromis erythromaculatus De Vos, Snoeks & Thys van den Audenaerde, 1991
Haplochromis estor Regan, 1929
Haplochromis eutaenia Regan & Trewavas, 1928
Haplochromis exspectatus Schraml, 2004
Haplochromis fischeri Seegers, 2008
Haplochromis flavipinnis Boulenger, 1906
Haplochromis flavus Seehausen, Zwennes & Lippitsch, 1998
Haplochromis fuelleborni Hilgendorf & Pappenheim, 1903 (Fuelleborn's mouthbrooder)
Haplochromis fuscus Regan, 1925
Haplochromis fusiformis Greenwood & Gee, 1969
Haplochromis gigas Seehausen & Lippitsch, 1998
Haplochromis gigliolii Pfeffer, 1896
Haplochromis gilberti Greenwood & Gee, 1969
Haplochromis goldschmidti F. Witte, Westbroek & de Zeeuw, 2013
Haplochromis gowersii Trewavas, 1928
Haplochromis gracilior Boulenger, 1914 (Torpedostripe haplochromis)
Haplochromis granti Boulenger, 1906
Haplochromis graueri Boulenger, 1914
Haplochromis greenwoodi Seehausen & Bouton, 1998
Haplochromis guiarti Pellegrin, 1904
Haplochromis harpakteridion van Oijen, 1991
Haplochromis heusinkveldi F. Wittee & Witte-Maas, 1987
Haplochromis hiatus Hoogerhoud & F. Witte, 1981
Haplochromis howesi van Oijen, 1992
Haplochromis humilior Boulenger, 1911
Haplochromis humilis Steindachner, 1866
Haplochromis igneopinnis Seehausen & Lippitsch, 1998
Haplochromis insidiae Snoeks, 1994
Haplochromis iris Hoogerhoud & F. Witte, 1981
Haplochromis ishmaeli Boulenger, 1906
Haplochromis kamiranzovu Snoeks, Coenen & Thys van den Audenaerde, 1984
Haplochromis katavi Seegers, 1996 (Katavi mouthbrooder)
Haplochromis katonga Schraml & Tichy, 2010
Haplochromis katunzii ter Huurne & F. Witte, 2010
Haplochromis kujunjui van Oijen, 1991
Haplochromis labiatus Trewavas, 1933
Haplochromis labriformis Nichols & La Monte, 1938
Haplochromis lacrimosus Boulenger, 1906
Haplochromis laparogramma Greenwood & Gee, 1969
Haplochromis latifasciatus Regan, 1929
Haplochromis limax Trewavas, 1933
Haplochromis lividus Greenwood, 1956
Haplochromis loati Greenwood, 1971
Haplochromis longirostris Hilgendorf, 1888
Haplochromis luteus Seehausen & Bouton, 1998
Haplochromis macconneli Greenwood, 1974
Haplochromis macrocephalus Seehausen & Bouton, 1998
Haplochromis macrognathus Regan, 1922
Haplochromis macrops Boulenger, 1911
Haplochromis macropsoides Greenwood, 1973
Haplochromis maculipinna Pellegrin, 1913
Haplochromis mahagiensis L. R. David & Poll, 1937
Haplochromis maisomei van Oijen, 1991
Haplochromis malacophagus Poll & Damas, 1939
Haplochromis mandibularis Greenwood, 1962
Haplochromis martini Boulenger, 1906
Haplochromis maxillaris Trewavas, 1928
Haplochromis mbipi Lippitsch & Bouton, 1998
Haplochromis megalops Greenwood & Gee, 1969
Haplochromis melanopterus Trewavas, 1928
Haplochromis melanopus Regan, 1922
Haplochromis melichrous Greenwood & Gee, 1969
Haplochromis mentatus Regan, 1925
Haplochromis mento Regan, 1922
Haplochromis michaeli Trewavas, 1928
Haplochromis microchrysomelas Snoeks, 1994
Haplochromis microdon Boulenger, 1906
Haplochromis multiocellatus Boulenger, 1913
Haplochromis mylergates Greenwood & Barel, 1978
Haplochromis mylodon Greenwood, 1973
Haplochromis nanoserranus Greenwood & Barel, 1978
Haplochromis nigrescens Pellegrin, 1909
Haplochromis nigricans Boulenger, 1906
Haplochromis nigripinnis Regan, 1921
Haplochromis nigroides Pellegrin, 1928
Haplochromis niloticus Greenwood, 1960
Haplochromis nubilus Boulenger, 1906 (Blue Victoria mouthbrooder)
Haplochromis nuchisquamulatus Hilgendorf, 1888
Haplochromis nyanzae Greenwood, 1962
Haplochromis nyererei Witte-Maas & F. Witte, 1985
Haplochromis obesus Boulenger, 1906
Haplochromis obliquidens Hilgendorf, 1888
Haplochromis obtusidens Trewavas, 1928
Haplochromis occultidens Snoeks, 1988
Haplochromis oligolepis Lippitsch, 2003
Haplochromis olivaceus Snoeks, De Vos, Coenen & Thys van den Audenaerde, 1990
Haplochromis omnicaeruleus Seehausen & Bouton, 1998
Haplochromis oregosoma Greenwood, 1973
Haplochromis orthostoma Regan, 1922
Haplochromis pachycephalus Greenwood, 1967
Haplochromis pallidus Boulenger, 1911
Haplochromis paludinosus (Greenwood, 1980)f
Haplochromis pancitrinus Mietes & F. Witte, 2010
Haplochromis pappenheimi Boulenger, 1914
Haplochromis paradoxus Lippitsch & Kaufman, 2003
Haplochromis paraguiarti Greenwood, 1967
Haplochromis paraplagiostoma Greenwood & Gee, 1969
Haplochromis paropius Greenwood & Gee, 1969
Haplochromis parorthostoma Greenwood, 1967
Haplochromis parvidens Boulenger, 1911
Haplochromis paucidens Regan, 1921
Haplochromis pellegrini Regan, 1922
Haplochromis percoides Boulenger, 1906
Haplochromis perrieri Pellegrin, 1909
Haplochromis petronius Greenwood, 1973
Haplochromis pharyngalis Poll & Damas, 1939
Haplochromis pharyngomylus Regan, 1929
Haplochromis phytophagus Greenwood, 1966
Haplochromis piceatus Greenwood & Gee, 1969
Haplochromis pitmani Fowler, 1936
Haplochromis placodus Poll & Damas, 1939
Haplochromis plagiodon Regan & Trewavas, 1928
Haplochromis plagiostoma Regan, 1922
Haplochromis plutonius Greenwood & Barel, 1978
Haplochromis prodromus Trewavas, 1935
Haplochromis prognathus Pellegrin, 1904
Haplochromis pseudopellegrini Greenwood, 1967
Haplochromis ptistes Greenwood & Barel, 1978
Haplochromis pundamilia Seehausen & Bouton, 1998
Haplochromis pyrrhocephalus F. Witte & Witte-Maas, 1987
Haplochromis pyrrhopteryx van Oijen, 1991
Haplochromis retrodens Hilgendorf, 1888
Haplochromis riponianus Boulenger, 1911
Haplochromis rubescens Snoeks, 1994
Haplochromis rubripinnis Seehausen, Lippitsch & Bouton, 1998
Haplochromis rudolfianus Trewavas, 1933
Haplochromis rufocaudalis Seehausen & Bouton, 1998
Haplochromis rufus Seehausen & Lippitsch, 1998
Haplochromis sauvagei Pfeffer, 1896
Haplochromis saxicola Greenwood, 1960
Haplochromis scheffersi Snoeks, De Vos & Thys van den Audenaerde, 1987
Haplochromis schubotzi Boulenger, 1914
Haplochromis schubotziellus Greenwood, 1973
Haplochromis serranus Pfeffer, 1896
Haplochromis serridens Regan, 1925
Haplochromis simotes Boulenger, 1911
Haplochromis simpsoni Greenwood, 1965
Haplochromis smithii Castelnau, 1861
Haplochromis snoeksi Wamuini Lunkayilakio & Vreven, 2010 <ref>{{cite journal| url=http://www.pfeil-verlag.de/04biol/pdf/ief21_3_08.pdf| title=Haplochromis' snoeksi, a new species from the Inkisi River basin, Lower Congo (Perciformes: Cichlidae)|last=Wamuini Lunkayilakio|author2=Vreven|journal=Ichthyological Exploration of Freshwaters|year=2010|volume=21|issue=3|pages=279–287}}</ref>
Haplochromis spekii Boulenger, 1906
Haplochromis sphex ter Huurne & F. Witte, 2010
Haplochromis squamipinnis Regan, 1921
Haplochromis squamulatus Regan, 1922
Haplochromis sulphureus Greenwood & Barel, 1978
Haplochromis tanaos van Oijen & F. Witte, 1996
Haplochromis taurinus Trewavas, 1933
Haplochromis teegelaari Greenwood & Barel, 1978
Haplochromis teunisrasi F. Witte & Witte-Maas, 1981
Haplochromis theliodon Greenwood, 1960
Haplochromis thereuterion van Oijen & F. Witte, 1996
Haplochromis thuragnathus Greenwood, 1967
Haplochromis tridens Regan & Trewavas, 1928
Haplochromis turkanae Greenwood, 1974 (Turkana haplo)
Haplochromis tyrianthinus Greenwood & Gee, 1969
Haplochromis ushindi van Oijen, 2004
Haplochromis vanheusdeni F. D. B. Schedel, Friel & U. K. Schliewen, 2014
Haplochromis vanoijeni de Zeeuw & F. Witte, 2010
Haplochromis velifer Trewavas, 1933
Haplochromis venator Greenwood, 1965
Haplochromis vicarius Trewavas, 1933
Haplochromis victoriae Greenwood, 1956
Haplochromis victorianus Pellegrin, 1904
Haplochromis vittatus Boulenger, 1901
Haplochromis vonlinnei van Oijen & de Zeeuw, 2008
Haplochromis welcommei Greenwood, 1966
Haplochromis worthingtoni Regan, 1929
Haplochromis xanthopteryx Seehausen & Bouton, 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 sp. '75'
Haplochromis sp. 'Amboseli'
Haplochromis sp. 'backflash cryptodon'
Haplochromis sp. 'black cryptodon'
Haplochromis sp. 'Chala'
Haplochromis sp. 'dusky wine-red fin'
Haplochromis sp. 'frogmouth'
Haplochromis sp. 'Kyoga flameback'
Haplochromis sp. 'long snout'
Haplochromis sp. 'Migori'
Haplochromis sp. 'nigrofasciatus'
Haplochromis sp. 'parvidens-like'
Haplochromis sp. 'purple head'
Haplochromis sp. 'ruby'
Haplochromis sp. 'Rusinga oral sheller'
Haplochromis sp. 'rainbow sheller'
Haplochromis sp. 'small obesoid'
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:
Astatoreochromis (3 species)
Astatotilapia (9 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:
Anomalochromis thomasi of the Hemichromini
Altolamprologus compressiceps of the Lamprologini
Tilapia jallae 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
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. (HTML abstract)
(1994): African cichlids II: Cichlids from East Africa. A handbook for their identification, care and breeding. Tetra Press, Germany.
(2001): Classification and Phylogenetic Relationships of African Tilapiine Fishes Inferred from Mitochondrial DNA Sequences. Molecular Phylogenetics and Evolution'' 20'''(3): 361–374.
Cichlid fish of Africa
Cichlid genera
Taxa named by Franz Martin Hilgendorf