Malawi cichlid checklist - The Cichlid Fishes of Lake Malawi, Africa

Checklist <strong>of</strong> the <strong>Cichlid</strong> <strong>Fishes</strong> <strong>of</strong> <strong>Lake</strong> <strong>Malawi</strong> (<strong>Lake</strong> Nyasa/Niassa) by M.K. Oliver, Ph.D.<br />

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Checklist <strong>of</strong> the <strong>Cichlid</strong> <strong>Fishes</strong> <strong>of</strong> <strong>Lake</strong> <strong>Malawi</strong> (<strong>Lake</strong> Nyasa/Niassa)<br />

by Michael K. Oliver, Ph.D.<br />

Updated 18 August 2012<br />

First posted June 1999<br />

<strong>The</strong> <strong>cichlid</strong>s <strong>of</strong> <strong>Lake</strong> <strong>Malawi</strong> constitute the largest vertebrate species flock (or complex <strong>of</strong> flocks), and<br />

largest lacustrine fish fauna, on earth. This list includes all <strong>cichlid</strong> species (and the few subspecies) that have been<br />

given formal scientific names. Many – several hundred – additional endemic <strong>cichlid</strong> species are known but still<br />

undescribed, and this fact must be considered in assessing the biodiversity <strong>of</strong> the lake. Current estimates <strong>of</strong> the total<br />

size <strong>of</strong> the lake’s <strong>cichlid</strong> fauna (counting both described, and known but undescribed, species) are 700-843 species<br />

(Turner et al., 2001; Snoeks, 2001; Konings, 2007). Additional undescribed species are still frequently being<br />

discovered, particularly in previously unexplored locations and in deep water.<br />

<strong>The</strong> entire <strong>Lake</strong> <strong>Malawi</strong> <strong>cichlid</strong> metaflock is composed <strong>of</strong> two, possibly separate, endemic clades, the<br />

“Hap” clade and the Mbuna clade. Membership in one or the other, or nonendemic status, is indicated for each<br />

genus, as is the type species <strong>of</strong> each endemic genus. <strong>The</strong> classification and synonymies are primarily based on the<br />

Catalog <strong>of</strong> <strong>Fishes</strong> with a few deviations (notably <strong>of</strong> species included in Cynotilapia and Tropheops). Because this<br />

list has just caught up with several synonymies, the total numbers have dropped somewhat compared with the<br />

previous (19 Apr 2012) version <strong>of</strong> this <strong>checklist</strong>, despite the addition <strong>of</strong> a new species <strong>of</strong> Hemitaeniochromis and the<br />

resurrection <strong>of</strong> Oreochromis saka from synonymy; all synonymized genera and species should now be listed under<br />

their senior synonym. <strong>The</strong> now 57 genera and the 391 species are numbered separately; the few subspecies<br />

recognized are no longer numbered separately and are indented under the nominate subspecies.<br />

Nearly all species are endemic to L. <strong>Malawi</strong>, in some cases extending also into the upper Shiré River<br />

including <strong>Lake</strong> Malombe. <strong>The</strong> nonendemic species number only four or five — Astatotilapia calliptera,<br />

Serranochromis robustus, Oreochromis shiranus (which is, however, represented in the <strong>Lake</strong> by an endemic<br />

subspecies), Tilapia rendalli, and one doubtfully occurring fish, Pseudocrenilabrus philander philander.<br />

Photographs <strong>of</strong> many <strong>of</strong> these fishes, and complete bibliographic citations <strong>of</strong> the taxonomic publications, are<br />

available on the author’s Web site at http://malawi<strong>cichlid</strong>s.com/.<br />

1. Abactochromis Oliver & Arnegard, 2010 — Endemic: Mbuna clade<br />

1. Abactochromis labrosus (Trewavas, 1935) type species<br />

2. Alticorpus Stauffer & McKaye, 1988 — Endemic: “Hap” clade<br />

2. Alticorpus ge<strong>of</strong>freyi Snoeks & Walapa, 2004<br />

3. Alticorpus macrocleithrum (Stauffer & McKaye, 1985)<br />

4. Alticorpus mentale Stauffer & McKaye, 1988 type species<br />

5. Alticorpus pectinatum Stauffer & McKaye, 1988<br />

6. Alticorpus peterdaviesi (Burgess & Axelrod, 1973)<br />

7. Alticorpus pr<strong>of</strong>undicola Stauffer & McKaye, 1988<br />

3. Aristochromis Trewavas, 1935 — Endemic: “Hap” clade<br />

8. Aristochromis christyi Trewavas, 1935 type species<br />

4. Astatotilapia Pellegrin, 1904 — Nonendemic<br />

9. Astatotilapia calliptera (Günther, 1894)<br />

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Checklist <strong>of</strong> the <strong>Cichlid</strong> <strong>Fishes</strong> <strong>of</strong> <strong>Lake</strong> <strong>Malawi</strong> (<strong>Lake</strong> Nyasa/Niassa) by M.K. Oliver, Ph.D.<br />

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5. Aulonocara Regan, 1922 — Endemic: “Hap” clade<br />

10. Aulonocara aquilonium Konings, 1995<br />

11. Aulonocara auditor (Trewavas, 1935)<br />

12. Aulonocara baenschi Meyer & Riehl, 1985<br />

13. Aulonocara brevinidus Konings, 1995<br />

14. Aulonocara brevirostre (Trewavas, 1935)<br />

15. Aulonocara ethelwynnae Meyer, Riehl, & Zetzsche, 1987<br />

16. Aulonocara gertrudae Konings, 1995<br />

17. Aulonocara guentheri Eccles, 1989<br />

18. Aulonocara hansbaenschi Meyer, Riehl, & Zetzsche, 1987<br />

19. Aulonocara hueseri Meyer, Riehl, & Zetzsche, 1987<br />

20. Aulonocara jacobfreibergi (Johnson, 1974)<br />

21. Aulonocara koningsi Tawil, 2003<br />

22. Aulonocara korneliae Meyer, Riehl, & Zetzsche, 1987<br />

23. Aulonocara maylandi maylandi Trewavas, 1984<br />

24. Aulonocara maylandi kandeensis Tawil & Allgayer, 1987<br />

25. Aulonocara nyassae Regan, 1922 type species<br />

26. Aulonocara rostratum Trewavas, 1935 [synonym: A. macrochir Trewavas, 1935]<br />

27. Aulonocara saulosi Meyer, Riehl, & Zetzsche, 1987<br />

28. Aulonocara steveni Meyer, Riehl, & Zetzsche, 1987<br />

29. Aulonocara stonemani (Burgess & Axelrod, 1973)<br />

30. Aulonocara stuartgranti Meyer & Riehl, 1985<br />

31. Aulonocara trematocephalum (Boulenger, 1901)<br />

6. Buccochromis Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

32. Buccochromis atritaeniatus (Regan, 1922)<br />

33. Buccochromis heterotaenia (Trewavas, 1935)<br />

34. Buccochromis lepturus (Regan, 1922)<br />

35. Buccochromis nototaenia (Boulenger, 1902) type species<br />

36. Buccochromis oculatus (Trewavas, 1935)<br />

37. Buccochromis rhoadesii (Boulenger, 1908)<br />

38. Buccochromis spectabilis (Trewavas, 1935)<br />

7. Caprichromis Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

39. Caprichromis liemi (McKaye & Mackenzie, 1982)<br />

40. Caprichromis orthognathus (Trewavas, 1935) type species<br />

8. Champsochromis Boulenger, 1915 — Endemic: “Hap” clade<br />

41. Champsochromis caeruleus (Boulenger, 1908) type species<br />

42. Champsochromis spilorhynchus (Regan, 1922)<br />

9. Cheilochromis Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

43. Cheilochromis euchilus (Trewavas, 1935) type species<br />

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Checklist <strong>of</strong> the <strong>Cichlid</strong> <strong>Fishes</strong> <strong>of</strong> <strong>Lake</strong> <strong>Malawi</strong> (<strong>Lake</strong> Nyasa/Niassa) by M.K. Oliver, Ph.D.<br />

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10. Chilotilapia Boulenger, 1908 — Endemic: “Hap” clade<br />

44. Chilotilapia rhoadesii Boulenger, 1908 type species<br />

11. Copadichromis Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

45. Copadichromis atripinnis Stauffer & Sato, 2002<br />

46. Copadichromis azureus Konings, 1990<br />

47. Copadichromis borleyi (Iles, 1960)<br />

48. Copadichromis chizumuluensis Stauffer & Konings, 2006<br />

49. Copadichromis chrysonotus (Boulenger, 1908)<br />

50. Copadichromis cyaneus (Trewavas, 1935)<br />

51. Copadichromis cyanocephalus Stauffer & Konings, 2006<br />

52. Copadichromis diplostigma Stauffer & Konings, 2006<br />

53. Copadichromis geertsi Konings, 1999<br />

54. Copadichromis ilesi Konings, 1999<br />

55. Copadichromis insularis Stauffer & Konings, 2006<br />

56. Copadichromis jacksoni (Iles, 1960)<br />

57. Copadichromis likomae (Iles, 1960)<br />

58. Copadichromis mbenjii Konings, 1990<br />

59. Copadichromis melas Stauffer & Konings, 2006<br />

60. Copadichromis mloto (Iles, 1960)<br />

61. Copadichromis nkatae (Iles, 1960)<br />

62. Copadichromis parvus Stauffer & Konings, 2006<br />

63. Copadichromis pleurostigma (Trewavas, 1935)<br />

64. Copadichromis pleurostigmoides (Iles, 1960)<br />

65. Copadichromis quadrimaculatus (Regan, 1922) type species<br />

66. Copadichromis trewavasae Konings, 1999<br />

67. Copadichromis trimaculatus (Iles, 1960)<br />

68. Copadichromis verduyni Konings, 1990<br />

69. Copadichromis virginalis (Iles, 1960)<br />

12. Corematodus Boulenger, 1897 — Endemic: “Hap” clade<br />

70. Corematodus shiranus Boulenger, 1897 type species<br />

71. Corematodus taeniatus Trewavas, 1935<br />

13. Ctenopharynx Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

72. Ctenopharynx intermedius (Günther, 1864) type species<br />

73. Ctenopharynx nitidus (Trewavas, 1935)<br />

74. Ctenopharynx pictus (Trewavas, 1935)<br />

14. Cyathochromis Trewavas, 1935 — Endemic: Mbuna clade<br />

75. Cyathochromis obliquidens Trewavas, 1935 type species<br />

15. Cynotilapia Regan, 1922 [Synonym: Microchromis Johnson] — Endemic: Mbuna clade<br />

76. Cynotilapia afra (Günther, 1894) type species<br />

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Checklist <strong>of</strong> the <strong>Cichlid</strong> <strong>Fishes</strong> <strong>of</strong> <strong>Lake</strong> <strong>Malawi</strong> (<strong>Lake</strong> Nyasa/Niassa) by M.K. Oliver, Ph.D.<br />

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77. Cynotilapia aurifrons (Tawil, 2011)<br />

78. Cynotilapia axelrodi Burgess, 1976<br />

79. Cynotilapia pulpican Tawil, 2002<br />

80. Cynotilapia zebroides (Johnson, 1975)<br />

16. Cyrtocara Boulenger, 1902 — Endemic: “Hap” clade<br />

81. Cyrtocara moorii Boulenger, 1902 type species<br />

17. Dimidiochromis Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

82. Dimidiochromis compressiceps (Boulenger, 1908)<br />

83. Dimidiochromis dimidiatus (Günther, 1864)<br />

84. Dimidiochromis kiwinge (Ahl, 1926)<br />

85. Dimidiochromis strigatus (Regan, 1922) type species<br />

18. Diplotaxodon Trewavas, 1935 — Endemic: “Hap” clade<br />

86. Diplotaxodon aeneus Turner & Stauffer, 1998<br />

87. Diplotaxodon apogon Turner & Stauffer, 1998<br />

88. Diplotaxodon argenteus Trewavas, 1935 type species<br />

89. Diplotaxodon ecclesi Burgess & Axelrod, 1973<br />

90. Diplotaxodon greenwoodi Stauffer & McKaye, 1986<br />

91. Diplotaxodon limnothrissa Turner, 1994<br />

92. Diplotaxodon macrops Turner & Stauffer, 1998<br />

19. Docimodus Boulenger, 1897 — Endemic: “Hap” clade<br />

93. Docimodus evelynae Eccles & Lewis, 1976<br />

94. Docimodus johnstoni Boulenger, 1897 type species<br />

20. Eclectochromis Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

95. Eclectochromis lobochilus (Trewavas, 1935)<br />

96. Eclectochromis ornatus (Regan, 1922) type species [synonym: E. festivus (Trewavas, 1935)]<br />

21. Exochochromis Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

97. Exochochromis anagenys Oliver, 1989 type species<br />

22. Fossorochromis Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

98. Fossorochromis rostratus (Boulenger, 1899) type species [synonym: Haplochromis macrorhynchus Regan,<br />

1021]<br />

23. Genyochromis Trewavas, 1935 — Endemic: Mbuna clade<br />

99. Genyochromis mento Trewavas, 1935 type species<br />

24. Gephyrochromis Boulenger, 1901 [Synonym: Christyella Trewavas, 1935] — Endemic: Mbuna clade<br />

100. Gephyrochromis lawsi Fryer, 1957<br />

101. Gephyrochromis moorii Boulenger, 1901 type species [synonym: Christyella nyasana Trewavas, 1935]<br />

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Checklist <strong>of</strong> the <strong>Cichlid</strong> <strong>Fishes</strong> <strong>of</strong> <strong>Lake</strong> <strong>Malawi</strong> (<strong>Lake</strong> Nyasa/Niassa) by M.K. Oliver, Ph.D.<br />

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25. Hemitaeniochromis Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

102. Hemitaeniochromis brachyrhynchus Oliver, 2012<br />

103. Hemitaeniochromis urotaenia (Regan, 1922) type species<br />

26. Hemitilapia Boulenger, 1902 — Endemic: “Hap” clade<br />

104. Hemitilapia oxyrhyncha Boulenger, 1902 type species<br />

27. Iodotropheus Oliver & Loiselle, 1972 — Endemic: Mbuna clade<br />

105. Iodotropheus declivitas Stauffer, 1994 [Konings (2007) considers this a synonym <strong>of</strong> I. sprengerae)]<br />

106. Iodotropheus sprengerae Oliver & Loiselle, 1972 type species<br />

107. Iodotropheus stuartgranti Konings, 1990<br />

28. Labeotropheus Ahl, 1926 — Endemic: Mbuna clade<br />

108. Labeotropheus fuelleborni Ahl, 1926 type species [synonym: L. curvirostris Ahl, 1926]<br />

109. Labeotropheus trewavasae Fryer, 1956<br />

29. Labidochromis Trewavas, 1935 — Endemic: Mbuna clade<br />

110. Labidochromis caeruleus Fryer, 1956<br />

111. Labidochromis chisumulae Lewis, 1982<br />

112. Labidochromis flavigulis Lewis, 1982<br />

113. Labidochromis freibergi Johnson, 1974<br />

114. Labidochromis gigas Lewis, 1982<br />

115. Labidochromis heterodon Lewis, 1982<br />

116. Labidochromis ianthinus Lewis, 1982<br />

117. Labidochromis joanjohnsonae Johnson, 1974 [generic placement as in Konings & Stauffer, 2012; synonym:<br />

Melanochromis exasperatus Burgess, 1976]<br />

118. Labidochromis lividus Lewis, 1982<br />

119. Labidochromis maculicauda Lewis, 1982<br />

120. Labidochromis mathotho Burgess & Axelrod, 1976<br />

121. Labidochromis mbenjii Lewis, 1982<br />

122. Labidochromis mylodon Lewis, 1982<br />

123. Labidochromis pallidus Lewis, 1982<br />

124. Labidochromis shiranus Lewis, 1982<br />

125. Labidochromis strigatus Lewis, 1982<br />

126. Labidochromis textilis Oliver, 1975<br />

127. Labidochromis vellicans Trewavas, 1935 type species<br />

128. Labidochromis zebroides Lewis, 1982<br />

30. Lethrinops Regan, 1922 — Endemic: “Hap” clade<br />

129. Lethrinops albus Regan, 1922<br />

130. Lethrinops altus Trewavas, 1931<br />

131. Lethrinops argenteus Ahl, 1926<br />

132. Lethrinops auritus (Regan, 1922)<br />

133. Lethrinops christyi Trewavas, 1931<br />

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Checklist <strong>of</strong> the <strong>Cichlid</strong> <strong>Fishes</strong> <strong>of</strong> <strong>Lake</strong> <strong>Malawi</strong> (<strong>Lake</strong> Nyasa/Niassa) by M.K. Oliver, Ph.D.<br />

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134. Lethrinops furcifer Trewavas, 1931<br />

135. Lethrinops gossei Burgess & Axelrod, 1973<br />

136. Lethrinops leptodon Regan, 1922<br />

137. Lethrinops lethrinus (Günther, 1894) type species<br />

138. Lethrinops longimanus Trewavas, 1931<br />

139. Lethrinops longipinnis Eccles & Lewis, 1978<br />

140. Lethrinops lunaris Trewavas, 1931<br />

141. Lethrinops macracanthus Trewavas, 1931<br />

142. Lethrinops macrochir (Regan, 1922)<br />

143. Lethrinops macrophthalmus (Boulenger, 1908)<br />

144. Lethrinops marginatus Ahl, 1926<br />

145. Lethrinops micrentodon (Regan, 1922)<br />

146. Lethrinops microdon Eccles & Lewis, 1977<br />

147. Lethrinops microstoma Trewavas, 1931<br />

148. Lethrinops mylodon mylodon Eccles & Lewis, 1979<br />

Lethrinops mylodon borealis Eccles & Lewis, 1979<br />

149. Lethrinops oculatus Trewavas, 1931<br />

150. Lethrinops parvidens Trewavas, 1931<br />

151. Lethrinops stridei Eccles & Lewis, 1977<br />

152. Lethrinops turneri Ngatunga & Snoeks, 2003<br />

31. Lichnochromis Trewavas, 1935 — Endemic: “Hap” clade<br />

153. Lichnochromis acuticeps Trewavas, 1935 type species<br />

32. Maylandia Meyer & Foerster, 1984 [Synonym: Metriaclima Stauffer, Bowers, Kellogg, & McKaye<br />

(1997); see Eschmeyer’s online Catalog <strong>of</strong> <strong>Fishes</strong> – http://bit.ly/Ouhlbm] — Endemic: Mbuna clade<br />

Note: Originally proposed as a subgenus <strong>of</strong> Pseudotropheus.<br />

154. Maylandia aurora (Burgess, 1976)<br />

155. Maylandia barlowi (McKaye & Stauffer, 1986)<br />

156. Maylandia benetos (Stauffer, Bowers, Kellogg, & McKaye, 1997)<br />

157. Maylandia callainos (Stauffer & Hert, 1992)<br />

158. Maylandia chrysomallos (Stauffer, Bowers, Kellogg, & McKaye, 1997)<br />

159. Maylandia cyneusmarginata (Stauffer, Bowers, Kellogg, & McKaye, 1997)<br />

160. Maylandia elegans (Trewavas, 1935)<br />

161. Maylandia emmiltos (Stauffer, Bowers, Kellogg, & McKaye, 1997)<br />

162. Maylandia estherae (Konings, 1995)<br />

163. Maylandia fainzilberi (Staeck, 1976)<br />

164. Maylandia flavifemina (Konings & Stauffer, 2006)<br />

165. Maylandia glaucos (Ciccotto, Konings, & Stauffer, 2011)<br />

166. Maylandia greshakei (Meyer & Foerster, 1984) type species<br />

167. Maylandia hajomaylandi (Meyer & Schartl, 1984)<br />

168. Maylandia heteropictus (Staeck, 1980)<br />

169. Maylandia lanisticola (Burgess, 1976)<br />

170. Maylandia livingstonii (Boulenger, 1899)<br />

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Checklist <strong>of</strong> the <strong>Cichlid</strong> <strong>Fishes</strong> <strong>of</strong> <strong>Lake</strong> <strong>Malawi</strong> (<strong>Lake</strong> Nyasa/Niassa) by M.K. Oliver, Ph.D.<br />

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171. Maylandia lombardoi (Burgess, 1977)<br />

172. Maylandia mbenjii (Stauffer, Bowers, Kellogg, & McKaye, 1997)<br />

173. Maylandia mossambica (Ciccotto, Konings, & Stauffer, 2011)<br />

174. Maylandia nkhunguensis (Ciccotto, Konings, & Stauffer, 2011)<br />

175. Maylandia phaeos (Stauffer, Bowers, Kellogg, & McKaye, 1997)<br />

176. Maylandia pursus (Stauffer, 1991)<br />

177. Maylandia pyrsonotos (Stauffer, Bowers, Kellogg, & McKaye, 1997) [synonym: M. sandaracinos (Stauffer,<br />

Bowers, Kellogg, & McKaye, 1997)]<br />

178. Maylandia sciasma (Ciccotto, Konings, & Stauffer, 2011)<br />

179. Maylandia thapsinogen (Stauffer, Bowers, Kellogg, & McKaye, 1997)<br />

180. Maylandia xanstomachus (Stauffer & Boltz, 1989)<br />

181. Maylandia xanthos (Ciccotto, Konings, & Stauffer, 2011)<br />

182. Maylandia zebra (Boulenger, 1899) [synonym: M. melabranchion (Stauffer, Bowers, Kellogg, & McKaye,<br />

1997)]<br />

33. Mchenga Stauffer & Konings, 2006 — Endemic: “Hap” clade<br />

183. Mchenga conophoros (Stauffer, LoVullo, & McKaye, 1993)<br />

184. Mchenga cyclicos (Stauffer, LoVullo, & McKaye, 1993) type species<br />

185. Mchenga eucinostomus (Regan, 1922)<br />

186. Mchenga flavimanus (Iles, 1960)<br />

187. Mchenga inornata (Boulenger, 1908)<br />

188. Mchenga thinos (Stauffer, LoVullo, & McKaye, 1993)<br />

34. Melanochromis Trewavas, 1935 — Endemic: Mbuna clade (included species after Konings &<br />

Stauffer, 2012)<br />

189. Melanochromis auratus (Boulenger, 1897)<br />

190. Melanochromis baliodigma Bowers & Stauffer, 1997 [synonym: M. xanthodigma Bowers & Stauffer, 1997]<br />

191. Melanochromis chipokae Johnson, 1975<br />

192. Melanochromis dialeptos Bowers & Stauffer, 1997<br />

193. Melanochromis heterochromis Bowers & Stauffer, 1993<br />

194. Melanochromis kaskazini Konings-Dudin, Konings, & Stauffer, 2009<br />

195. Melanochromis lepidiadaptes Bowers & Stauffer, 1997<br />

196. Melanochromis loriae Johnson, 1975 [synonym: M. parallelus Burgess & Axelrod, 1976]<br />

197. Melanochromis melanopterus Trewavas, 1935 type species [synonym: Melanochromis mellitus Johnson, 1976]<br />

198. Melanochromis mossambiquensis Konings-Dudin, Konings, & Stauffer, 2009<br />

199. Melanochromis mpoto Konings & Stauffer, 2012<br />

200. Melanochromis robustus Johnson, 1986<br />

201. Melanochromis simulans Eccles, 1973<br />

202. Melanochromis vermivorus Trewavas, 1935<br />

203. Melanochromis wochepa Konings-Dudin, Konings, & Stauffer, 2009<br />

35. Mylochromis Regan, 1922 [Synonyms: Maravichromis Eccles & Trewavas, 1989; Platygnathochromis<br />

Eccles & Trewavas, 1989] — Endemic: “Hap” clade<br />

204. Mylochromis anaphyrmus (Burgess & Axelrod, 1973)<br />

205. Mylochromis balteatus (Trewavas, 1935)<br />

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Checklist <strong>of</strong> the <strong>Cichlid</strong> <strong>Fishes</strong> <strong>of</strong> <strong>Lake</strong> <strong>Malawi</strong> (<strong>Lake</strong> Nyasa/Niassa) by M.K. Oliver, Ph.D.<br />

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206. Mylochromis chekopae Turner & Howarth, 2001<br />

207. Mylochromis ensatus Turner & Howarth, 2001<br />

208. Mylochromis epichorialis (Trewavas, 1935)<br />

209. Mylochromis ericotaenia (Regan, 1922)<br />

210. Mylochromis formosus (Trewavas, 1935)<br />

211. Mylochromis guentheri (Regan, 1922)<br />

212. Mylochromis incola (Trewavas, 1935)<br />

213. Mylochromis labidodon (Trewavas, 1935)<br />

214. Mylochromis lateristriga (Günther, 1864) type species<br />

215. Mylochromis melanonotus (Regan, 1922)<br />

216. Mylochromis melanotaenia (Regan, 1922)<br />

217. Mylochromis mola (Trewavas, 1935)<br />

218. Mylochromis mollis (Trewavas, 1935)<br />

219. Mylochromis obtusus (Trewavas, 1935)<br />

220. Mylochromis plagiotaenia (Regan, 1922)<br />

221. Mylochromis semipalatus (Trewavas, 1935)<br />

222. Mylochromis sphaerodon (Regan, 1922)<br />

36. Naevochromis Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

223. Naevochromis chrysogaster (Trewavas, 1935) type species<br />

37. Nimbochromis Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

224. Nimbochromis fuscotaeniatus (Regan, 1922) [May belong in Tyrannochromis]<br />

225. Nimbochromis linni (Burgess & Axelrod, 1974)<br />

226. Nimbochromis livingstonii (Günther, 1894) type species<br />

227. Nimbochromis polystigma (Regan, 1922) [synonyms: N. maculimanus (Regan, 1922); N. pardalis (Trewavas,<br />

1935)]<br />

228. Nimbochromis venustus (Boulenger, 1908) [synonym: Haplochromis simulans Regan, 1922]<br />

38. Nyassachromis Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

229. Nyassachromis boadzulu (Iles, 1960)<br />

230. Nyassachromis breviceps (Regan, 1922) type species<br />

231. Nyassachromis leuciscus (Regan, 1922)<br />

232. Nyassachromis microcephalus (Trewavas, 1935)<br />

233. Nyassachromis nigritaeniatus (Trewavas, 1935)<br />

234. Nyassachromis prostoma (Trewavas, 1935)<br />

235. Nyassachromis purpurans (Trewavas, 1935)<br />

236. Nyassachromis serenus (Trewavas, 1935)<br />

39. Oreochromis Günther, 1889 — Nonendemic genus. Except for O. shiranus, all L. <strong>Malawi</strong> species have<br />

been placed in the subgenus Nyasalapia, which also includes species not found in L. <strong>Malawi</strong>.<br />

237. Oreochromis (Nyasalapia) karongae (Trewavas, 1941) — Endemic species.<br />

238. Oreochromis (Nyasalapia) lidole (Trewavas, 1941) — Endemic species, except also occurs in the crater lakes<br />

L. Kingiri and L. Chungruru in the Rungwe Mtns. in Tanzania some 50 km north <strong>of</strong> L. <strong>Malawi</strong>.<br />

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Checklist <strong>of</strong> the <strong>Cichlid</strong> <strong>Fishes</strong> <strong>of</strong> <strong>Lake</strong> <strong>Malawi</strong> (<strong>Lake</strong> Nyasa/Niassa) by M.K. Oliver, Ph.D.<br />

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239. Oreochromis (Nyasalapia) saka (Lowe, 1953) — Endemic species.<br />

240. Oreochromis (Nyasalapia) squamipinnis (Günther, 1864) — Endemic species.<br />

241. Oreochromis (Oreochromis) shiranus shiranus Boulenger, 1897 — Endemic subspecies <strong>of</strong> a nonendemic<br />

species. O. shiranus is composed <strong>of</strong> two subspecies: O. s. shiranus, endemic to the <strong>Lake</strong> <strong>Malawi</strong> basin<br />

(including inflowing rivers and the upper Shiré River), and O. s. chilwae, endemic to <strong>Lake</strong>s Chilwa and<br />

Chiuta.<br />

40. Otopharynx Regan, 1920 — Endemic: “Hap” clade<br />

242. Otopharynx antron Cleaver, Konings, & Stauffer, 2009<br />

243. Otopharynx argyrosoma (Regan, 1922)<br />

244. Otopharynx auromarginatus (Boulenger, 1908) type species<br />

245. Otopharynx brooksi Oliver, 1989<br />

246. Otopharynx decorus (Trewavas, 1935)<br />

247. Otopharynx heterodon (Trewavas, 1935)<br />

248. Otopharynx lithobates Oliver, 1989 [synonym: O. walteri Konings, 1990]<br />

249. Otopharynx ovatus (Trewavas, 1935)<br />

250. Otopharynx pachycheilus Arnegard & Snoeks, 2001<br />

251. Otopharynx selenurus Regan, 1922<br />

252. Otopharynx speciosus (Trewavas, 1935)<br />

253. Otopharynx spelaeotes Cleaver, Konings, & Stauffer, 2009<br />

254. Otopharynx tetraspilus (Trewavas, 1935)<br />

255. Otopharynx tetrastigma (Günther, 1894)<br />

41. Pallidochromis Turner, 1994 — Endemic: “Hap” clade<br />

256. Pallidochromis tokolosh Turner, 1994 type species<br />

42. Petrotilapia Trewavas, 1935 — Endemic: Mbuna clade<br />

257. Petrotilapia chrysos Stauffer & van Snik, 1996<br />

258. Petrotilapia flaviventris Lundeba, Stauffer, & Konings, 2011<br />

259. Petrotilapia genalutea Marsh, 1983<br />

260. Petrotilapia microgalana Ruffing, Lambert, & Stauffer, 2006<br />

261. Petrotilapia mumboensis Lundeba, Stauffer, & Konings, 2011<br />

262. Petrotilapia nigra Marsh, 1983<br />

263. Petrotilapia palingnathos Lundeba, Stauffer, & Konings, 2011<br />

264. Petrotilapia pyroscelos Lundeba, Stauffer, & Konings, 2011<br />

265. Petrotilapia tridentiger Trewavas, 1935 type species<br />

266. Petrotilapia xanthos Lundeba, Stauffer, & Konings, 2011<br />

43. Placidochromis Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

267. Placidochromis acuticeps Hanssens, 2004<br />

268. Placidochromis acutirostris Hanssens, 2004<br />

269. Placidochromis argyrogaster Hanssens, 2004<br />

270. Placidochromis boops Hanssens, 2004<br />

271. Placidochromis borealis Hanssens, 2004<br />

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Checklist <strong>of</strong> the <strong>Cichlid</strong> <strong>Fishes</strong> <strong>of</strong> <strong>Lake</strong> <strong>Malawi</strong> (<strong>Lake</strong> Nyasa/Niassa) by M.K. Oliver, Ph.D.<br />

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272. Placidochromis chilolae Hanssens, 2004<br />

273. Placidochromis communis Hanssens, 2004<br />

274. Placidochromis domirae Hanssens, 2004<br />

275. Placidochromis ecclesi Hanssens, 2004<br />

276. Placidochromis electra (Burgess, 1979)<br />

277. Placidochromis elongatus Hanssens, 2004<br />

278. Placidochromis fuscus Hanssens, 2004<br />

279. Placidochromis hennydaviesae (Burgess & Axelrod, 1973)<br />

280. Placidochromis intermedius Hanssens, 2004<br />

281. Placidochromis johnstoni (Günther, 1894) [synonym: Haplochromis sexfasciatus Regan, 1922]<br />

282. Placidochromis koningsi Hanssens, 2004<br />

283. Placidochromis lineatus Hanssens, 2004<br />

284. Placidochromis longimanus (Trewavas, 1935) type species<br />

285. Placidochromis longirostris Hanssens, 2004<br />

286. Placidochromis longus Hanssens, 2004<br />

287. Placidochromis lukomae Hanssens, 2004<br />

288. Placidochromis macroceps Hanssens, 2004<br />

289. Placidochromis macrognathus Hanssens, 2004<br />

290. Placidochromis mbunoides Hanssens, 2004<br />

291. Placidochromis milomo Oliver, 1989<br />

292. Placidochromis minor Hanssens, 2004<br />

293. Placidochromis minutus Hanssens, 2004<br />

294. Placidochromis msakae Hanssens, 2004<br />

295. Placidochromis nigribarbis Hanssens, 2004<br />

296. Placidochromis nkhatae Hanssens, 2004<br />

297. Placidochromis nkhotakotae Hanssens, 2004<br />

298. Placidochromis obscurus Hanssens, 2004<br />

299. Placidochromis ordinarius Hanssens, 2004<br />

300. Placidochromis orthognathus Hanssens, 2004<br />

301. Placidochromis pallidus Hanssens, 2004<br />

302. Placidochromis phenochilus (Trewavas, 1935)<br />

303. Placidochromis platyrhynchos Hanssens, 2004<br />

304. Placidochromis polli (Burgess & Axelrod, 1973)<br />

305. Placidochromis rotundifrons Hanssens, 2004<br />

306. Placidochromis subocularis (Günther, 1894)<br />

307. Placidochromis trewavasae Hanssens, 2004<br />

308. Placidochromis turneri Hanssens, 2004<br />

309. Placidochromis vulgaris Hanssens, 2004<br />

44. Protomelas Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

310. Protomelas annectens (Regan, 1922)<br />

311. Protomelas dejunctus Stauffer, 1993<br />

312. Protomelas fenestratus (Trewavas, 1935)<br />

313. Protomelas insignis (Trewavas, 1935)<br />

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Checklist <strong>of</strong> the <strong>Cichlid</strong> <strong>Fishes</strong> <strong>of</strong> <strong>Lake</strong> <strong>Malawi</strong> (<strong>Lake</strong> Nyasa/Niassa) by M.K. Oliver, Ph.D.<br />

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314. Protomelas kirkii (Günther, 1894) type species<br />

315. Protomelas labridens (Trewavas, 1935)<br />

316. Protomelas macrodon Eccles, 1989<br />

317. Protomelas marginatus marginatus (Trewavas, 1935)<br />

Protomelas marginatus vuae (Trewavas, 1935)<br />

318. Protomelas pleurotaenia (Boulenger, 1901) [synonym: Haplochromis microstoma Regan, 1922]<br />

319. Protomelas similis (Regan, 1922)<br />

320. Protomelas spilonotus (Trewavas, 1935)<br />

321. Protomelas spilopterus (Trewavas, 1935)<br />

322. Protomelas taeniolatus (Trewavas, 1935) [synonym: Haplochromis cancellus Greenwood, 1963]<br />

323. Protomelas triaenodon (Trewavas, 1935)<br />

324. Protomelas virgatus (Trewavas, 1935)<br />

45. Pseudocrenilabrus Fowler, 1934 — Nonendemic<br />

325. Pseudocrenilabrus philander philander (Weber, 1897) [Note: This genus and species is listed as occurring in<br />

L. <strong>Malawi</strong> by CLOFFA, Checklist <strong>of</strong> the Freshwater <strong>Fishes</strong> <strong>of</strong> <strong>Africa</strong> (1991, Vol. IV: 396); see also<br />

http://www.fishbase.org/summary/Pseudocrenilabrus-philander+philander.html. No literature or specimen<br />

citation was provided in support <strong>of</strong> this listing. <strong>The</strong> occurrence <strong>of</strong> this species in <strong>Lake</strong> <strong>Malawi</strong> seems<br />

doubtful and should be substantiated by a museum specimen.]<br />

46. Pseudotropheus Regan, 1922 — Endemic: Mbuna clade. Note: Many additional species were formerly<br />

placed in Pseudotropheus within two groups originally proposed as subgenera, Maylandia and Tropheops,<br />

both now recognized as full genera (which see). It is thought that some <strong>of</strong> the species included below<br />

belong in Pseudotropheus sensu stricto, but that others below may eventually reside in other genera.<br />

326. Pseudotropheus ater Stauffer, 1988<br />

327. Pseudotropheus benetos (Bowers & Stauffer, 1997)<br />

328. Pseudotropheus brevis (Trewavas, 1935)<br />

329. Pseudotropheus crabro (Ribbink & Lewis, 1982)<br />

330. Pseudotropheus cyaneorhabdos (Bowers & Stauffer, 1997)<br />

331. Pseudotropheus cyaneus Stauffer, 1988<br />

332. Pseudotropheus demasoni Konings, 1994<br />

333. Pseudotropheus elongatus Fryer, 1956<br />

334. Pseudotropheus flavus Stauffer, 1988<br />

335. Pseudotropheus fuscoides Fryer, 1956<br />

336. Pseudotropheus fuscus Trewavas, 1935<br />

337. Pseudotropheus galanos Stauffer & Kellogg, 2002<br />

338. Pseudotropheus interruptus (Johnson, 1975) [synonym: Melanochromis elastodema Bowers & Stauffer, 1997]<br />

339. Pseudotropheus johannii Eccles, 1973<br />

340. Pseudotropheus longior Seegers, 1996<br />

341. Pseudotropheus minutus Fryer, 1956<br />

342. Pseudotropheus perileucos (Bowers & Stauffer, 1997)<br />

343. Pseudotropheus perspicax (Trewavas, 1935)<br />

344. Pseudotropheus purpuratus Johnson, 1976<br />

345. Pseudotropheus saulosi Konings, 1990<br />

346. Pseudotropheus socol<strong>of</strong>i Johnson, 1974<br />

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Checklist <strong>of</strong> the <strong>Cichlid</strong> <strong>Fishes</strong> <strong>of</strong> <strong>Lake</strong> <strong>Malawi</strong> (<strong>Lake</strong> Nyasa/Niassa) by M.K. Oliver, Ph.D.<br />

–––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––<br />

347. Pseudotropheus tursiops Burgess & Axelrod, 1975<br />

348. Pseudotropheus williamsi (Günther, 1894) type species<br />

47. Rhamphochromis Regan, 1922 — Endemic: “Hap” clade<br />

349. Rhamphochromis esox (Boulenger, 1908) [synonyms: R. leptosoma Regan, 1922; R. melanotus Ahl, 1926]<br />

350. Rhamphochromis ferox Regan, 1922<br />

351. Rhamphochromis longiceps (Günther, 1864) type species<br />

352. Rhamphochromis macrophthalmus Regan, 1922<br />

353. Rhamphochromis woodi Regan, 1922 [synonyms: R. brevis Trewavas, 1935; R. lucius Ahl, 1926]<br />

48. Sciaenochromis Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

354. Sciaenochromis ahli (Trewavas, 1935) type species [synonym: Haplochromis serranoides Ahl, 1926]<br />

355. Sciaenochromis benthicola Konings, 1993<br />

356. Sciaenochromis fryeri Konings, 1993<br />

357. Sciaenochromis gracilis (Trewavas, 1935) [Note: Sometimes classified in Mylochromis]<br />

358. Sciaenochromis psammophilus Konings, 1993<br />

359. Sciaenochromis spilostichus (Trewavas, 1935) [Note: Sometimes classified in Mylochromis]<br />

49. Serranochromis Regan, 1920 — Nonendemic<br />

360. Serranochromis robustus robustus (Günther, 1864)<br />

50. Stigmatochromis Eccles & Trewavas, 1989 — Endemic: “Hap” clade.<br />

361. Stigmatochromis macrorhynchos Stauffer, Cleaver-Yoder, & Konings, 2011<br />

362. Stigmatochromis melanchros Stauffer, Cleaver-Yoder, & Konings, 2011<br />

363. Stigmatochromis modestus (Günther, 1894)<br />

364. Stigmatochromis pholidophorus (Trewavas, 1935)<br />

365. Stigmatochromis pleurospilus (Trewavas, 1935)<br />

366. Stigmatochromis woodi (Regan, 1922) type species<br />

51. Taeniochromis Eccles & Trewavas, 1989 — Endemic: “Hap” clade.<br />

367. Taeniochromis holotaenia (Regan, 1922) type species [synonym: Haplochromis bodyi Ahl, 1926]<br />

52. Taeniolethrinops Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

368. Taeniolethrinops cyrtonotus (Trewavas, 1931)<br />

369. Taeniolethrinops furcicauda (Trewavas, 1931)<br />

370. Taeniolethrinops laticeps (Trewavas, 1931)<br />

371. Taeniolethrinops praeorbitalis (Regan, 1922) type species [synonyms: Lethrinops fasciatus Ahl, 1926; L.<br />

macrorhynchus Regan, 1922]<br />

53. Tilapia Smith, 1840 — Nonendemic<br />

372. Tilapia rendalli (Boulenger, 1897)<br />

373. Tilapia sparrmanii Smith, 1840 Note: G. Bell-Cross (<strong>The</strong> <strong>Fishes</strong> <strong>of</strong> Rhodesia. National Museums and<br />

Monuments <strong>of</strong> Rhodesia, Salisbury, 1976: 236) reported this tilapia to occur in L. <strong>Malawi</strong> and the same<br />

statement was repeated by Bell-Cross and J. L. Minshull in the revised edition <strong>of</strong> the same book (<strong>The</strong><br />

<strong>Fishes</strong> <strong>of</strong> Zimbabwe. National Museums and Monuments <strong>of</strong> Zimbabwe, Harare, 1988: 271). B. Marshall<br />

(<strong>The</strong> <strong>Fishes</strong> <strong>of</strong> Zimbabwe and their Biology. Smithiana Monograph 3, 2011: 235) also states that it occurs<br />

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Checklist <strong>of</strong> the <strong>Cichlid</strong> <strong>Fishes</strong> <strong>of</strong> <strong>Lake</strong> <strong>Malawi</strong> (<strong>Lake</strong> Nyasa/Niassa) by M.K. Oliver, Ph.D.<br />

–––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––<br />

in <strong>Lake</strong> <strong>Malawi</strong> and includes a dot for it just below the lake’s southeast arm in the distribution map. No<br />

supporting details or literature citation are provided in any <strong>of</strong> these books. However, I was finally<br />

persuaded that this species should be listed as part <strong>of</strong> the <strong>Lake</strong> <strong>Malawi</strong> fauna by an email from Denis<br />

Tweddle (pers. commun. 27-Feb-2012); Denis wrote that “I have never seen it in the lake itself but it<br />

occurs right on the edge and must occasionally get into the lake. It is abundant (or at least was in 1976)<br />

around the gabions <strong>of</strong> the bridge over the entrance to Chia Lagoon and that could be regarded as an inlet <strong>of</strong><br />

the lake.” I agree; the road bridge over the Chia Lagoon inlet (which I have also driven over) is only 600 m<br />

from <strong>Lake</strong> <strong>Malawi</strong> proper, as can be measured in Google Earth. <strong>The</strong> channel between the lagoon and the<br />

lake proper appears quite uniform; T. sparrmanii would be expected to occur along its entire length.<br />

54. Tramitichromis Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

374. Tramitichromis brevis (Boulenger, 1908) type species<br />

375. Tramitichromis intermedius (Trewavas, 1935)<br />

376. Tramitichromis lituris (Trewavas, 1931)<br />

377. Tramitichromis trilineatus (Trewavas, 1931)<br />

378. Tramitichromis variabilis (Trewavas, 1931)<br />

55. Trematocranus Trewavas, 1935 — Endemic: “Hap” clade<br />

379. Trematocranus labifer (Trewavas, 1935)<br />

380. Trematocranus microstoma Trewavas, 1935 type species<br />

381. Trematocranus placodon (Regan, 1922)<br />

56. Tropheops Trewavas, 1984 — Endemic: Mbuna clade. [Note: Originally proposed as a subgenus <strong>of</strong><br />

Pseudotropheus]<br />

382. Tropheops gracilior (Trewavas, 1935)<br />

383. Tropheops lucerna (Trewavas, 1935)<br />

384. Tropheops macrophthalmus (Ahl, 1926)<br />

385. Tropheops microstoma (Trewavas, 1935)<br />

386. Tropheops modestus (Johnson, 1974)<br />

387. Tropheops novemfasciatus (Regan, 1922)<br />

388. Tropheops romandi (Colombe, 1979)<br />

389. Tropheops tropheops (Regan, 1922) type species<br />

57. Tyrannochromis Eccles & Trewavas, 1989 — Endemic: “Hap” clade<br />

390. Tyrannochromis macrostoma (Regan, 1922) type species [synonyms: T. maculiceps (Ahl, 1926); T. polyodon<br />

(Trewavas, 1935)]<br />

391. Tyrannochromis nigriventer Eccles, 1989<br />

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