2019-03-14

Lamprologini...part 3



An account of Lamprologini, including Neolamprologus sp. “Mwila”, a new shell-dwelling species from Mwila Island in the Kipili Archipelago, the facts about N. sp. “Eseki”, the whereabouts of Kinyamkolo, and much more (part 3 of 3)

Part 1Part 2 – Part 3

Appendix 1 (fish species names derived from J. E. S. Moore)
In 1895–1896 and 1899–1900, J. E. S. Moore collected and brought home about 270 fishes from the East African lakes, including 47 species which were described as new (Boulenger, 1906: 539). Besides Lake Tanganyika, Moore also obtained fishes, including new species, from Lakes Malawi, Kivu, Edward, and Albert. Collections of fish were also made in Lake Mweru, but Moore failed to bring these back to London (Boulenger, 1899b: 84). In addition, Moore examined Lakes Chilwa (Shirwa), Chila (Kela), Victoria, Naivasha (Nivasha) (Moore, 1903: 121), but no samples are recorded from these lakes.
In total, seven fish species were described and named after Moore. A complete list of these species include the following (all described by Boulenger between 1898 and 1902): (1) the cyprinid Barilius moorii (now Raiamas) (1900: 480), (2) the clariid Clarias moorii (cf. C. gariepinus) (1901: 13), (3) the mastacembelid Mastacembelus moorii (1898b: 22), (4) the cichlids Cyrtocara moorii (1902: 70), (5) Gephyrochromis moorii (1901: 4), (6) Lamprologus moorii (now Variabilichromis) (1898b: 8), and (7) Tropheus moorii (1898b: 18).
J. E. S. Moore’s name was also used in other animal species names, including a frog [Arthroleptis moorii (cf. Phrynobatrachus natalensis) (Boulenger, 1898c: 479)], sponge [Spongilla moorei (now Nudospongilla) (Evans, 1899: 472)], shrimp [Palaemon moorei (now Macrobrachium) (Calman, 1899: 709)], and earthworm [Benhamia moorii (cf. B. itoliensis) (Beddard, 1901: 191)]. Additional references: Catalog of Fishes (Eschmeyer et al., 2018; 2019) and FishBase (Froese & Pauly, 2019).
Additional fish species may have been named after Moore had Boulenger not delayed the descriptions of them or failed to recognise them upon the examination of Moore’s specimens (see Appendices 3, 4, and 5).

Fig. 142. Map of J. E. S. Moore’s second expedition to Lake Tanganyika, 1899–1900, showing all the locations where he collected fish (1–7), as well as those five locations he made collections at on his first expedition in 1895–1896 (indicated in purple). Only the cichlids are listed on the map, for a complete list of species and specimens, see Appendix 4. The digits following the species names indicate the number of specimens collected. One star (*) indicates the misidentification of species, see more in Appendices 3 and 4; two stars (**) indicate the uncertainty of the species Ectodus longianalis, believed to be synonymous with Enantiopus melanogenys [Xenotilapia (Takahashi, 2003: 375)]. Current location names have been used on the map, except for Kinyamkolo, today’s Mpulungu, because of its significance in the historical context.
Regarding both expeditions, Moore reached Lake Tanganyika by Stevenson Road, initially arriving at Chituta. During his second expedition (see the map), he travelled by canoe to Kapata Point, climbed the Chituta Escarpment, marched to Kinyamkolo, went back to Chituta (not shown on the map) to fetch the steamer, the Good News, whose boiler had to be repaired before the actual lake expedition could commence (§ 32.17, 32.18). On 28 September 1899, the expedition headed northeast for Kalambo (Fergusson, 1900: 389). About a month later, on 21 October, the expedition reached Bujumbura, after which it immediately returned south for supply and to pick up two expedition members, Messrs Berridge and Mathews, who had remained at Sumbu. While Malcolm Fergusson, the surveyor to the expedition, took command of the steamer, heading south for Chituta, Moore was dropped at Ujiji to find and make agreements with 150 native porters, together with whom Moore then marched for 11 days back to Bujumbura, where he was reunited with Fergusson and the rest of the expedition crew. After the visits at Bujumbura and the mouth of Rusisi River, on 30 November, the expedition continued farther north to Lakes Kivu, Edward, Albert, Victoria, Naivasha, and the snow-capped summits of the Ruwenzori Range, identified as Ptolemy’s ‘Mountains of the Moon’.
Originally connected to the Free Church of Scotland (FCS), but operating on a commercial rather than humanitarian basis, the African Lakes Corporation (ALC), under the leadership of the Moir brothers, started building the Stevenson Road in 1881 (Johnston, 1897: 71; Gamwell & Gamwell, 1961: 520) stretching from Karonga, Lake Malawi to Chituta, Lake Tanganyika (§ 32.17). The construction of the road was partly funded by James Stevenson, who subscribed a sum of £4,000 for the purpose of preliminary surveys (Johnston, 1897: 71; Haskard, 1965: 15). Stevenson was a founder and director of the ALC [see a photo of their Blantyre headquarters in the late 1800s in Johnston (1897: 67)] as well as a passionate supporter of African missions, including the Livingstonia Mission of the FCS. The reason for building a road between the lakes was mainly to open up the country for commerce, thereby abolishing the slave trade, which David Livingstone had pleaded for in his Cambridge speech of 1857 (Monk, 1858: 24). In addition to the road construction, Stevenson partially persuaded the London Missionery Society (LMS) to open a new missionary station at the southern end of Lake Tanganyika and put a steamship on the lake, which would further hinder the slave trade. That steamship was the Good News, delivered in sections to Edward Coode Hore of the LMS at Mbete Bay by the ALC in 1883–1885 (Fig. 124) (Hainsworth, 1951: 18). James Stewart, an engineer in the employ of the ALC, who had surveyed the northern end of Lake Malawi in 1877 and visited the southern end of Lake Tanganyika in 1879, including Mbete (Stewart, 1879; 1880a; Johnston, 1897: 67), started the work with Stevenson Road near Karonga, but died of fever in 1883, a fate which also struck his successor, William McEwan, in 1885 (Gamwell & Gamwell, 1961: 521; Haskard, 1965: 16). The British South Africa Company completed the road from Karonga to Abercorn (Mbala) (Gamwell & Gamwell, 1961: 520). To commemorate the encouragement, financial help, and labour of Stevenson, Stewart, and McEwan, a memorial called the Stevenson Stone, a magnificent, six metres tall stone cross was erected 10 miles west of Karonga in 1905 (Haskard, 1965: 16); for photos, see Livingstone (1923: 305) and Haskard (1965: 19). At the end of the 1800s, southern parts of Stevenson Road were under siege by the Arabs, blocking the route to Lake Tanganyika for many travellers, including Moore, who in early 1896 had to stop at Zomba, south of Lake Malawi, to wait for the road to be cleared by the commissioner’s forces of the British Central Africa, before proceeding (Nature, 1896a: 470). Today, much of Stevenson Road is bypassed, overgrown by vegetation, and forgotten, but approximately where the road once existed, today there is the border between Tanzania and Zambia.

Appendix 2 (the grammar behind the species epithets ‘moorii’ and ‘moorei’ as in Tropheus moorii and Nudospongilla moorei)
Among the species names derived from J. E. S. Moore, two specific epithets have been applied over the years: moorii and moorei, both of which are available and accurate (Appendix 1). Boulenger applied moorii in all his seven descriptions of species named in honour of Moore, but his description of Tropheus moorii in 1898 was erroneously cited as T. moorei six years later by Pellegrin (1904: 305), whose misquotation appears to be related to carelessness, because throughout his publication, Pellegrin (1904) cited Boulenger’s specific epithet as ‘moorei’ 14 times and ‘moorii’ 5 times, without giving any reason for the alternative spelling. Although Boulenger maintained his original and correct spelling in all subsequent publications (e.g. 1901c; 1905; 1906; 1915; 1920), a long series of cichlid authors adopted Pellegrin’s incorrect version, T. moorei, including Poll (1956), Marlier (1959), Chlupaty (1961), Matthes (1962), Wickler (1969), Scheuermann (1976), Staeck (1975c), Neergaard (1976), Kawanabe (1981), Hori (1983), Takamura (1984), Yamaoka (1985), and Kuwamura (1987).
While some authors favouring T. moorei believed T. moorii to be a misspelling (e.g., Sundström, 1977), others like Mayland (1976) believed the specific epithets of T. moorei and, for example, Cyrtocara moorii, to be related to the names of two different persons, one called Moore and the other Moori. Larsson (1977) and Kullander (1977) ended speculation by providing the relevant facts, which included details about Moore’s expeditions, Latin grammar, and nomenclatural rules, the former author referring to the nomenclatural code of zoology (ICZN, 1964). In an updated statement, Mayland (1979: 82) corrected the spelling to T. moorii and changed his opinion on the etymology, acknowledging that both T. moorii and C. moorii were named in honour of J. E. S. Moore.
Ever since the draft of the first internationally accepted rules of zoological nomenclature (Blanchard et al., 1898), as well as the definite rules themselves (Blanchard et al., 1905), names of biological species have included those based on names of persons, a practice still in force (ICZN, 1999). Normally, the genitive case of a person’s name is applicable for a species name (ICZN, 1999: Art. 11.9.1.3), but because the genitive may be formed from a personal name that is (1) Latin or has been Latinised, or (2) a modern name, two alternatives of genitive ending exist (ICZN, 1999: Art. 31.1.1–2). (1) When a name of a man, such as Moore, is Latinised, it becomes Moorius, whose genitive case is formed by replacing ‘-us’ (the nominative ending) with ‘-i’ (the genitive ending), resulting in Moorii, or, when applied as a specific epithet, moorii. (2) If the personal name is a modern name of a man, the genitive case is obtained by adding ‘-i’ to the name as it is, resulting in moorei. By convention, nomenclatural grammar follows Latin grammar, see more in Bailey (1992). Obviously, Boulenger chose the first genitive ending alternative, while others like Evans (1899: 472) (Nudospongilla moorei) and Calman (1899: 709) (Macrobrachium moorei) chose the second alternative. The rules of formalising personal names into species names were the same in 1898 as today [cf. Blanchard et al., 1898: 184 (A.III.12); 1905: 32 (Art. 14c)].
Authors such as Poll (1956), Marlier (1959), Matthes (1962), and others favouring T. moorei may have followed Pellegrin (1904) and his misspelling of T. moorii, alternatively made their own misspellings, or they may have chosen to deliberately modify the name to T. moorei based on nomenclatural recommendations which existed in the set of rules of 1961 and 1964 (ICZN, 1961; 1964), and presumably also in those that prevailed during the 1950s, in which the Latinisation of modern names is not recommended. When a species name is based on a personal name, the genitive case ending ‘-i’ in preference to the ending ‘-ii’ should be added to the entire name [e.g. ICZN, 1964: 107 (Appendix D, III, 16)], which implies that species names should not be formed by Latinising modern names prior to adding the genitive ending, i.e., the second alternative of genitive ending is recommended in preference to the first (see above). Following these recommendations, if a species name is based on Moore, the specific epithet moorei should be applied in favour of moorii. Possibly, some authors of the 1950s and 1960s chose to modify the name T. moorii to T. moorei thinking that such a modification was mandatory based upon the nomenclatural rules of that time. Of course, if they did, they were wrong. The original spelling of a name is the “correct original spelling”, unless demonstrably incorrect, which include “a lapsus calami or a copyist’s or printer’s error”, but not an incorrect transliteration or Latinisation, “or use of an inappropriate connecting vowel”, see more in ICZN (1999: Art. 32). An example of a correct species name that the author unsuccessfully attempted to amend is Ectodus foae, invalidly modified to E. foai, a modification attempt based on the fact that the person the author intended to honour with a species name was not a woman, but a man (Vaillant, 1899; 1908). An example of a justified, mandatory, and successful correction is that of the specific epithet of Pseudotropheus pursus (Stauffer, 1991), which was evidently a misspelling of purus, the Latin word for clean, because the specific epithet of the species was given to reflect the cleaning behaviour of the species (Stauffer et al., 2013: 103).
In conclusion, Boulenger’s original spelling of T. moorii is correct, and any alternative spellings, including T. moorei by Pellegrin and others are incorrect.

Appendix 3 (additional species associated with J. E. S. Moore)
Moore should perhaps be given credit for being the initial collector and discoverer of additional species because some of the fishes he entrusted to Boulenger were not described until later when other collectors, typically Hecq and Cunnington, had also supplied Boulenger with specimens of the same species, which resulted in them becoming primarily associated with Hecq or Cunnington. A few examples include: (1) In 1896 and 1899, Moore collected five specimens of a claroteid catfish (NHM, 2019k; 2019l; 2019m; Boulenger, 1911: 371), which Boulenger initially referred to as Auchenaspis biscutata (Boulenger, 1898b: 24) and Auchenoglanis biscutatus (Boulenger, 1901c: 148). Following the addition of one conspecific specimen obtained by Cunnington from Mtondwe Bay (Kakonde Bay), Boulenger (1906: 553) described four of them as Auchenoglanis occidentalis var. tanganicanus [now A. tanganicanus (Retzer, 2010: 37; Kullander & Roberts, 2012: 361)], hence, this claroteid catfish became more associated with Cunnington’s expedition than with Moore’s. (2) At Bujumbura, Burundi, at the end of 1899, Moore collected three specimens of a cyprinid fish (NHM, 2019n; Boulenger, 1911: 21), which Boulenger described as Barbus tropidolepis in a work based on specimens obtained by Hecq (Boulenger, 1900: 133; 1901a: 228; 1901c: 145). (3) During his second expedition to East Africa, Moore obtained his Lake Tanganyika fish collection between 20 September and beginning of November 1899, which included two specimens of Tylochromis polylepis from Kinyamkolo (Niamkolo) and one from Sibwesa (Boulenger, 1901c: 154; 1915: 383; NHM, 2019p; 2019v), although this species was originally described by Boulenger the following year in a work in French comprising the collections of Hecq originating from Albertville (Mtoa), DR Congo (Boulenger, 1900: 143).
Furthermore, some of Moore’s collections comprised more species, including new species, than were recognised by Boulenger. In Moore’s recollections of his stay at Msamba in 1899, he wrote: “We stayed two days in Msambu [Msamba], fishing with nets and lines and killing rock fishes wholesale with dynamite. In this way I found several entirely new species” (Moore, 1901: 114). Based on Moore’s Msamba specimens, Boulenger (1901b; 1901c) recognised four new species: Limnochromis auritus, Paracyprichromis nigripinnis, Ophthalmotilapia boops, and Asprotilapia leptura [Xenotilapia (Takahashi, 2003: 375)]. However, among the nine L. auritus specimens (Boulenger, 1915: 416), there were at least four, possibly five, Triglachromis otostigma, later described as a distinct species by Regan (1920b: 152), partly based on Moore’s Msamba specimens (NHM, 2019b; 2019c).
Indeed, Boulenger (1906: 568, Plate 39, Fig. 4; 1915: 416) also failed to recognise three T. otostigma specimens in Cunnington’s L. auritus collection from Mshale (north of Kigoma).
Furthermore, Boulenger (1901c: 150; 1915: 484) reported three specimens of Neolamprologus furcifer in Moore’s Msamba collection. However, one of these was N. timidus and another presumably N. sp. “Cygnus” (Sven O. Kullander, pers. comm.; Kullander et al., 2014a: 305, 323; NHM, 2019d). The radiograph of NHM (2019d) shows N. timidus above and N. sp. “Cygnus” below. The third specimen is no longer found in the premises of the museum (Natural History Museum, London).
Moreover, Moore obtained four specimens of Bathybates minor at Bujumbura in October 1899, initially misidentified by Boulenger (1901c: 153; 1915: 443) as B. ferox, but described as a distinct species five years later working on Cunnington’s collection (Boulenger, 1906: 564–565).
Regarding the species description of Protomelas pleurotaenia (see § 32.24), it was originally based on 12 specimens collected by Moore in 1899 (seven recorded as from the north end of Lake Tanganyika and five allegedly from the mouth of Rusisi River) (Boulenger, 1901a: 474; 1901b: 4; 1901c: 157; 1915: 247–248). However, not only were all specimens in fact collected in Lake Malawi, upon re-examination (Trewavas, 1946: 244–245; Eccles & Trewavas, 1989: 46–51), it was concluded that they represent four species in total: (1) P. pleurotaenia (seven specimens) (Boulenger, 1901c: 157; NHM, 2019e; 2019f; 2019g), (2) P. macrodon (one specimen) (Eccles & Trewavas, 1989: 49; NHM, 2019h), (3) Mylochromis plagiotaenia (three specimens) (Regan, 1922: 706; NHM, 2019i), and (4) Otopharynx selenurus (one specimen) (Regan, 1922: 679; NHM, 2019j); additional references: Maréchal (1991: 390). The photo of NHM (2019e) shows four of the types of P. pleurotaenia. Furthermore, Haplochromis microstoma (Regan, 1922: 695) is an alleged synonym of P. pleurotaenia (Trewavas, 1946: 245–246; Eccles & Trewavas, 1989: 46). For Moore’s complete collection of fish from Lake Malawi, see Appendix 5.
At least six more species were collected by Moore in 1895–1896 and 1899–1900, but not described until some or many years later, including (1) the amphiliid Amphilius jacksonii (Boulenger, 1912: 602) [originally identified by Boulenger (1898b: 24; 1911: 357) as Anoplopterus platychir, § 32.25], (2) the malapterurid Malapterurus tanganyikaensis (Roberts, 2000: 11) [M. electricus (Boulenger, 1901c: 149; 1911: 512; cf. Nature, 1897: 258)], (3) the alestid Brycinus rhodopleura (Boulenger, 1906: 547; 1909: 218) [Alestes macrolepidotus (Boulenger, 1898b: 25)], (4) the latid Lates mariae (Steindachner, 1909a: 399) [L. microlepis (Boulenger, 1901c: 149), L. angustifrons (Boulenger, 1915: 110)], and (5) the cichlids Haplochromis astatodon (Regan, 1921: 637) [Tilapia burtoni (Boulenger, 1901c: 157)] and (6) H. paucidens (Regan, 1921: 638) [Paratilapia bloyeti (Boulenger, 1901c: 151)], see more in Appendices 4 and 5.
Additional references: Catalog of Fishes (Eschmeyer et al., 2018; 2019) and NHM [Natural History Museum] (2014).

Appendix 4 (collection locations, fish species, and number of specimens collected by J. E. S. Moore in Lake Tanganyika during his expeditions in 1895–1896 and 1899–1900)
“Mr. Moore’s first collection, made in 1895–96, contained about 90 Fishes, referred to 33 species, 25 of which were described as new. His second collection, made in 1899–1900, contained 180 examples of 48 species, 22 of which were new [including specimens from Lakes Tanganyika, Malawi, Kivu, Edward, and Albert]” (Boulenger, 1906: 539).
Below, Moore’s 1895–1896 and 1899–1900 collections of cichlids and non cichlids from Lake Tanganyika are listed in four sections, including (1a) 1895–1896 cichlids, (1b) 1895–1896 non cichlids, (2a) 1899–1900 cichlids, and (2b) 1899–1900 non cichlids.
(1a) Moore’s 1895–1896 collection of cichlids contained 74 specimens of 21 species from two locations, Kinyamkolo and Mbity Rocks.
Kinyamkolo: Altolamprologus compressiceps, 2 (specimens); Bathybates ferox, 1; Callochromis macrops, 2, Cyathopharynx furcifer, 2; Cyprichromis leptosoma, 2; Eretmodus cyanostictus, 3; “Gnathochromispfefferi, 1; Lepidiolamprologus elongatus, 1; Lobochilotes labiata, 4; Neolamprologus fasciatus, 1; N. furcifer, 3; N. modestus, 1; Ophthalmotilapia ventralis, 8; Petrochromis polyodon, 2; Simochromis diagramma, 4; Telmatochromis temporalis, 4 (NHM, Collection1, Collection2, Collection3); Tropheus moorii, 5; Variabilichromis moorii, 2; sub total, 48 specimens.
Mbity Rocks: Callochromis macrops, 1; Cyprichromis leptosoma, 2; Eretmodus cyanostictus, 5; Julidochromis ornatus, 5; Lepidiolamprologus elongatus, 1; Neolamprologus furcifer, 1; N. modestus, 1; Ophthalmotilapia ventralis, 1; Perissodus microlepis, 1; Petrochromis polyodon, 2; Telmatochromis temporalis, 1; T. vittatus, 2; Variabilichromis moorii, 3; sub total, 26 specimens.
(1b) Moore’s 1895–1896 collection of non cichlids contained 19 specimens of 12 nominal species from three locations, Kinyamkolo, Mbity Rocks, and Sumbu.
Kinyamkolo, including specimens presumably from Kinyamkolo: Alestes macrolepidotus [Brycinus rhodopleura (NHM, Collection4)], 1; A. macrophthalmus, 1; Auchenaspis biscutata [Auchenoglanis occidentalis, A. tanganicanus (NHM, Collection5)], 2; Clarias anguillaris [cf. C. mossambicus, C. gariepinus (NHM, Collection6)], 1; C. liocephalus, 1; Hydrocyon forskali [Hydrocynus forskahlii (NHM, Collection7)], 1; Lates microlepis, 3 (NHM, Collection8, Collection9); Malapterurus electricus [M. tanganyikaensis (NHM, Collection10)], 2; unidentified (NHM, Collection11), 1; sub total, 13 specimens.
Mbity Rocks and marshes near Mbity (Mbete): Anoplopterus platychir [Amphilius jacksonii (NHM, Collection12), § 32.25], 2; Haplochilus tanganicanus (Lamprichthys tanganicanus), 1; Mastacembelus moorii, 2; sub total, 5 specimens.
Sumbu: Synodontis multipunctatus, 1; sub total, 1 specimen.
(1a + 1b) Total number of cichlids and non cichlids from Lake Tanganyika collected by Moore 1895–1896: 93 specimens, 33 species.
(2a) Moore’s 1899–1900 collection of cichlids contained 81 specimens of 28 species from nine locations, Bujumbura, Kalambo, Sibwesa, Kinyamkolo, Lake Tanganyika (unknown location), Maswa (Lubengela, see Figs. 130–131), Msamba, south end (presumably near Kalambo), and Cape Tembwe.
Bujumbura and near Bujumbura, referred to as Usambura: Bathybates ferox, 1; B. minor, 4 (cf. Boulenger, 1901c: 565; 1915: 443; NHM, Collection13); Ectodus longianalis [cf. Enantiopus melanogenys (NHM, Collection14, Collection15)], 5; Limnochromis auritus, 1; Trematocara marginatum, 1; T. unimaculatum, 10; Xenochromis hecqui, 3; sub total, 25 specimens.
Kalambo: Bathybates ferox, 1; Boulengerochromis microlepis, 1; Limnotilapia dardennii, 1; Oreochromis tanganicae, 1; Reganochromis calliurus, 3; sub total, 7 specimens.
Sibwesa, referred to as Kibwesa/Kibwesi: Lamprologus lemairii, 2; Limnotilapia dardennii, 3; Lobochilotes labiata, 1; Telmatochromis temporalis, 2; Tylochromis polylepis, 1; Xenotilapia ornatipinnis, 7; sub total, 16 specimens.
Kinyamkolo: Tylochromis polylepis, 2; sub total, 2 specimens.
Lake Tanganyika, unknown location: Asprotilapia leptura [Xenotilapia (Takahashi, 2003)], 2; Julidochromis ornatus, 1; Neolamprologus furcifer, 2; sub total, 2 specimens.
Maswa (Lubengela, see Figs. 130–131): Hemibates stenosoma, 1; sub total, 1 specimen.
Msamba, referred to as Msambu: Asprotilapia leptura, 1; Cyprichromis leptosoma, 4; Lamprologus lemairii, 1; Limnochromis auritus (see Appendix 3), 5; Neolamprologus furcifer (see Appendix 3), 1; N. timidus [recorded as N. furcifer (Boulenger, 1901c: 150; 1915: 484), see Appendix 3], 1; cf. N. sp. “Cygnus” [recorded as N. furcifer (Boulenger, 1901c: 150; 1915: 484), see Appendix 3], 1; Ophthalmotilapia boops, 2; Paracyprichromis nigripinnis, 2; Simochromis diagramma, 1; Triglachromis otostigma [recorded as L. auritus (Boulenger, 1901c: 150–151; 1915: 415–416), see Appendix 3], 4; sub total, 23 specimens.
South end, presumably near Kalambo: Bathybates ferox, 1; Hemibates stenosoma, 2; sub total, 3 specimens.
Cape Tembwe, referred to as Tembwi: Bathybates fasciatus, 1; B. ferox, 1; sub total, 2 specimens.
(2b) Moore’s 1899–1900 collection of non cichlids contained 48 specimens of 21 nominal species from seven locations, Bujumbura, Kalambo, Sibwesa, Kinyamkolo, middle of lake, middle of lake (presumably near Msamba), and Cape Tembwe.
Bujumbura and near Bujumbura, referred to as Usambura: Barbus platyrhinus, 1; B. serrifer [cf. B. neumayeri, Enteromius neumayeri (NHM, Collection16)], 3; B. tropidolepis, 3; Barilius moorii [Raiamas moorii (NHM, Collection17)], 4; B. tanganicae [cf. Opsaridium microlepis (NHM, Collection18)], 1; Capoeta tanganicae [Acapoeta tanganicae (NHM, Collection19, Collection20)], 3; Chrysichthys brachynema, 1; Mastacembelus frenatus, 1; M. taeniatus, 1; Synodontis granulosus, 3; sub total, 21 specimens.
Kalambo: Alestes macrolepidotus, 1; Auchenoglanis biscutatus [A. tanganicanus (NHM, Collection21)], 2; Chrysichthys brachynema, 3; Malapterurus electricus [M. tanganyikaensis (NHM, Collection22)], 1; Polypterus congicus, 4; Synodontis multipunctatus, 1; sub total , 12 specimens.
Lake Tanganyika, unknown location: Alestes macrophthalmus, 1; Citharinus gibbosus, 3; sub total, 4 specimens.
Sibwesa, referred to as Kibwesa/Kibwesi: Chrysichthys myriodon [cf. C. brachynema (NHM, Collection23)], 4; sub total, 4 specimens.
Kinyamkolo: Chrysichthys myriodon [cf. C. brachynema (NHM, Collection24)], 1; Lates microlepis [L. mariae (NHM, Collection25)], 1; sub total, 2 specimens.
Middle of lake: Auchenoglanis biscutatus [A. tanganicanus (NHM, Collection26)], 1; sub total, 1 specimen.
Middle of lake (presumably near Msamba; cf. Moore, 1901b: 112): Lates microlepis, 3; sub total, 3 specimens.
Cape Tembwe, referred to as Tembwi: Chrysichthys myriodon [cf. C. brachynema (NHM, Collection27)], 1; sub total, 1 specimen.
(2a + 2b) Total number of cichlids and non cichlids from Lake Tanganyika collected by Moore 1899–1900: 129 specimens, 49 species.
(1a + 1b + 2a + 2b) Total number of cichlids and non cichlids from Lake Tanganyika collected by Moore 1895–1896 and 1899–1900: 222 specimens, approximately 70 species.
Additional references: Catalog of Fishes (Eschmeyer et al., 2018; 2019) and NHM [Natural History Museum] (2014).

Appendix 5 (Moore’s collection comprising 52 fishes from Lake Malawi obtained between 11 July and 15 August 1899 during his second expedition, including the misidentification of species)
Moore obtained a small collection of fishes from Lake Malawi, most of which were cichlids, deposited at the British Museum (Natural History), now the Natural History Museum (NHM), upon his return in 1900. The following 39 specimens with catalogue numbers were listed by Boulenger in 1906, all with a collection location as “Lake Nyasa”.
BMNH.1906.9.7.1 Labeo mesops, 1 (specimen) [L. intermedius (Worthington, 1933: 297) (NHM, Collection28)]
BMNH.1906.9.7.2–3 Barilius microlepis, 2 [cf. Opsaridium microcephalum (NHM, Collection29)]
BMNH.1906.9.7.4 Barilius microlepis, 1 [cf. Opsaridium microcephalum (NHM, Collection30)]
BMNH.1906.9.7.5 Barilius microcephalus, 1 [cf. Opsaridium microlepis (Collection31)]
BMNH.1906.9.7.6 Bagrus meridionalis, 1 (NHM, Collection32)
BMNH.1906.9.7.7–9 Synodontis zambesensis, 3 [S. zambezensis (NHM, Collection33)]
BMNH.1906.9.7.10–12 Paratilapia robusta, 3 [Hemichromis robustus (Günther, 1864b: 312); Serranochromis robustus (NHM, Collection34)]
BMNH.1906.9.7.13 Paratilapia modesta, 1 [Haplochromis maculimanus (Regan, 1922: 689); Nimbochromis (Eccles & Trewavas, 1989: 288); cf. N. polystigma (Konings, 1990: 216, cited in Snoeks & Manuel, 2004: 73) (NHM, Collection35)]
BMNH.1906.9.7.14–17 Paratilapia modesta, 4 [Haplochromis woodi (Regan, 1922: 702); Stigmatochromis (Eccles & Trewavas, 1989: 174) (NHM, Collection36)]
BMNH.1906.9.7.18 Paratilapia dimidiata, 1 [Hemichromis dimidiatus (Günther, 1864b: 313); Dimidiochromis dimidiatus (Eccles & Trewavas, 1989: 91) (NHM, Collection37)]
BMNH.1906.9.7.19 Paratilapia dimidiata, 1 [Haplochromis holotaenia (Regan, 1922: 697); Taeniochromis (Eccles & Trewavas, 1989: 73) (NHM, Collection38)]
BMNH.1906.9.7.20 Paratilapia dimidiata, 1 [Haplochromis holotaenia (Regan, 1922: 697); Taeniochromis (Eccles & Trewavas, 1989: 73) (NHM, Collection39)]
BMNH.1906.9.7.21 Paratilapia nototaenia (type), 1 [Buccochromis (Eccles & Trewavas, 1989: 237) (NHM, Collection40)]
BMNH.1906.9.7.22 Paratilapia longiceps, 1 [Hemichromis longiceps (Günther, 1864b: 313); Rhamphochromis ferox (Regan, 1922: 725); cf. Champsochromis (Boulenger, 1915: 434) (NHM, Collection41)]
BMNH.1906.9.7.23–25 Paratilapia longiceps, 3 [Rhamphochromis woodi (Regan, 1922: 725) and R. esox (Regan, 1922: 726) (NHM, Collection42)]
BMNH.1906.9.7.26 Cyrtocara moorii (type), 1 (NHM, Collection43)
BMNH.1906.9.7.27–29 Tilapia shirana, 3 [Oreochromis shiranus (Boulenger, 1897a: 916) (NHM, Collection44)]
BMNH.1906.9.7.30 Tilapia squamipinnis, 1 [Chromis (Günther, 1864b: 311); Oreochromis (Trewavas & Teugels, 1991: 341) (NHM, Collection45)]
BMNH.1906.9.7.31–32 Tilapia rendalli, 2 [Coptodon (NHM, Collection46)]
BMNH.1906.9.7.33–35 Tilapia lethrinus, 3 [Chromis (Günther, 1894: 622); Lethrinops (Regan, 1922: 720) (NHM, Collection47)]
BMNH.1906.9.7.36 Tilapia rostrata, 1 [(Boulenger, 1899d: 131); Fossorochromis rostratus (Eccles & Trewavas, 1989: 275); cf. Haplochromis macrorhynchus (Regan, 1922: 717) (NHM, Collection48)]
BMNH.1906.9.7.37 Petrochromis nyassae (type), 1 [P. polyodon (NHM, Collection49)]
BMNH.1906.9.7.38 Hemitilapia oxyrhynchus (type), 1 (NHM, Collection50)
BMNH.1906.9.7.39 Hemitilapia oxyrhynchus (type), 1 (NHM, Collection51)
One specimen, the type of Petrochromis nyassae, was erroneously labelled as “Lake Nyasa” (Boulenger, 1902: 70); it was later identified as P. polyodon, collected in Lake Tanganyika (Matthes & Trewavas, 1960: 350; cf. Regan, 1922: 675).
Mistakenly recorded as from Lake Tanganyika, the additional 14 specimens were collected in Lake Malawi (Boulenger, 1901b: 4; 1901c: 156–158; Trewavas, 1946: 244–245; Eccles & Trewavas, 1989: 46–49, 145):
BMNH.1906.9.6.121 Gephyrochromis moorii (NHM, Collection52) (“north end of Lake Tanganyika”)
BMNH.1906.9.6.139–143 Tilapia pleurotaenia, 4 [Haplochromis (Trewavas, 1946: 244–245); Protomelas (Eccles & Trewavas, 1989: 46–47) (NHM, Collection53) (“north end of Lake Tanganyika”)]
BMNH.1906.9.6.144.a Tilapia pleurotaenia, 1 [Otopharynx selenurus (Regan, 1922: 679; Trewavas, 1946: 245) (NHM, Collection54) (“north end of Lake Tanganyika”)]
BMNH.1906.9.6.144.b Tilapia pleurotaenia, 1 [Protomelas macrodon (Eccles & Trewavas, 1989: 49) (NHM, Collection55) (“north end of Lake Tanganyika”)]
BMNH.1906.9.6.145 Tilapia pleurotaenia, 1 (skeleton) [Haplochromis (Trewavas, 1946: 244–245); Protomelas (Eccles & Trewavas, 1898: 46–47) (NHM, Collection56) (“north end of Lake Tanganyika”)]
BMNH.1906.9.6.146–147 Tilapia pleurotaenia, 2 [Haplochromis (Trewavas, 1946: 244–245); Protomelas (Eccles & Trewavas, 1989: 46–47) (NHM, Collection57) (“mouth of Rusisi River”)]
BMNH.1906.9.6.148–150 Tilapia pleurotaenia, 3 [Limnotilapia (Regan, 1920a: 39); Haplochromis plagiotaenia (Regan, 1922: 706; Trewavas, 1946: 245–246); Maravichromis (Eccles & Trewavas, 1989: 203, 206); Mylochromis plagiotaenia (Regan, 1920a: 45; Derijst & Snoeks, 1992: 173) (NHM, Collection58) (“mouth of Rusisi River”)]
BMNH.1906.9.6.151 Tilapia trematocephala, 1 [Limnotilapia (Regan, 1920a: 39); Aulonocara trematocephalum (Eccles & Trewavas, 1989: 145) (NHM, Collection59) (“north end of Lake Tanganyika”)]
In total, Moore obtained 52 (39-1+14) specimens of fish from Lake Malawi. Of the 43 specimens of cichlids, Boulenger (1901c: 137–160; 1902: 69–71) recognised and described only six new species, including Buccochromis nototaenia, Cyrtocara moorii, Hemitilapia oxyrhynchus, Gephyrochromis moorii, Protomelas pleurotaenia, and Aulonocara trematocephalum. However, as listed above, the collection included at least ten more new cichlid species. Seemingly, all of Moore’s five specimens recorded as Stigmatochromis modestus by Boulenger represented new and undescribed species, later described as Haplochromis maculimanus [(Regan, 1922: 689), now Nimbochromis, possibly N. polystigma (Regan, 1922: 688; Konings, 1990: 216)] and H. woodi [(Regan, 1922: 702), now Stigmatochromis (Eccles & Trewavas, 1989: 175)], among others. Indeed, only the original and single type specimen, collected by Joseph A. Williams at Likoma in 1891, on which Günther (1894: 618, 625) based the species description is listed as S. modestus in Eccles and Trewavas (1989: 174), all Moore’s five specimens are excluded as none of them are S. modestus. Likewise, two additional specimens, obtained by Percy Rendall and Harry Johnston, and misidentified as S. modestus by Boulenger (1897a: 915; 1915: 327) represent other species.
Not only Boulenger misidentified species and underestimated the total number of Lake Malawi cichlid species, Moore (1903: 123–124, cited in Fryer, 2000: 11) listed 25 Lake Malawi cichlid species (excluding G. moorii, P. pleurotaenia, and A. trematocephalum) and stated, concerning the lake’s different fish species, that “there can be no doubt that we have pretty well exhausted the freshwater fauna of Nyasa”. Likewise, Boulenger (1905: 54–58) listed 26 species from Lake Malawi and Shire River, including G. moorii, P. pleurotaenia, A. trematocephalum, and Oreochromis mossambicus, but excluding Mylochromis subocularis [placed in synonymy with Placidochromis johnstoni (Boulenger, 1899d: 130), later Astatotilapia calliptera (Boulenger, 1915: 222)] and Otopharynx tetrastigma [placed in synonymy with P. johnstoni (Boulenger, 1899d: 130), later reinstated (Boulenger, 1915: 250)]. Besides O. mossambicus, these 28 first Lake Malawi cichlids were included in the following six publications:
1. Günther (1864b: 303–314), including six new cichlid species, (1) Oreochromis squamipinnis, (2) Mylochromis lateristriga, (3) Ctenopharynx intermedius, (4) Serranochromis robustus, (5) Rhamphochromis longiceps, (6) Dimidiochromis dimidiatus.
2. Günther (1894: 616–628), including ten new cichlid species, (7) Mylochromis subocularis, (8) Placidochromis johnstoni, (9) Lethrinops lethrinus, (10) Otopharynx tetrastigma, (11) Astatotilapia calliptera, (12) Protomelas kirkii, (13) Pseudotropheus williamsi, (14) Stigmatochromis modestus, (15) Nimbochromis livingstonii, (16) Cynotilapia afra.
3. Boulenger (1897a: 915–920), including four new cichlid species, (17) Coptodon rendalli, (18) Oreochromis shiranus, (19) Docimodus johnstoni, (20) Corematodus shiranus.
4. Boulenger (1897b: 155), including one new cichlid species, (21) Melanochromis auratus.
5. Boulenger (1899d: 98–143), including three new cichlid species, (22) Fossorochromis rostratus, (23) Pseudotropheus livingstonii, (24) Maylandia zebra.
6. Boulenger (1902: 69–71), including three new cichlid species, (25) Buccochromis nototaenia, (26) Cyrtocara moorii, (27) Hemitilapia oxyrhynchus.
Indeed, based on Moore’s collection of 43 specimens of Lake Malawi cichlids, it is rather remarkable that Boulenger (1901c: 137–160; 1902: 69–71) only recognised and described six new species when the collection probably included three or four times that. In 1905, only 28 cichlid species were known from Lake Malawi, but the total number of cichlids in that lake may be 1000 (Fryer, 2000; Snoeks, 2000; Konings, 2007: 5), or even more. “About half the [cichlid] species in the lake still haven’t been described” (Jay Stauffer Jr., pers. comm. to Crair, 2019).

Appendix 6 (a directory of geographical locations mentioned in the study with links to Google Maps)
Abercorn (Mbala), African Diving, Albertville, Bujumbura, Cameron Bay, Cape Chaitika, Cape Chipimbi, Cape Kabogo, Cape Kapembwa, Cape Mpimbwe, Cape Muzimu, Cape Tembwe, Chibwensolo, Chikola Hill, Chiloelo Point, Chisanse, Chituta Bay, Chituta Escarpment, Chongo Bay, Fwambo, Halembe, Ikola, Isonga, Kabimba, Kaboge, Kabwe, Kafwila, Kala, Kalambo Lodge, Kalambo River, Kalandasi Point, Kalanswi Bay, Kalemie, Kalepa, Kamamba Island, Kambwimba, Kampemba Point, Kanshenke, Kansombo, Kantalamba, Kapampa, Kapata, Karema, Karilani Island, Karonga, Kasakalawe, Kasanga, Kasenga Point, Kasisi Island, Kasola Hill, Kasola Island, Katabe Bay, Katanga, Katanka Point, Katete, Katondo Point, Katoto, Katumba Point, Kavala Islands, Kawimbe, Kekese, Kerenge Island, Kigoma, Kiku, Kilewani, Kinyamkolo, Kipili, Kirando (Kigoma Region), Kirando (Rukwa Region), Kisi Island, Kombe, Korongwe Bay, Kumbula Island, Kungwe Bay, Kungwe Point, Lamvya Bay, Livua, Luagala Point, Lubengela, Lufubu River, Lugufu River, Luhanga, Lukuga River, Lunangwa, Lupita Island, Lupota, Lusekese, Lwasase Point, Lyamembe, Magambo, Mahale Mountains (2,462 m), Makosi Hill, Malagarasi River, Maleza Island, Maswa Point, Mbala, Mbete Bay, Mbita Island, Mbity Rocks (unresolved), Mbofula Points, Mboko Island, Mbwilo Bay, Mikongolo Island, Miyako Point, Mkamba River, Mlowa Point, Moba, Moliro, Molwe, Mongwe, Moyobozi, Mpala (Kafwanka), Mpando Point, Mpimbwe Point, Mpulungu, Mpulungu Hill, Mpulungu Port, Msamba, Msehezi River, Mshale, Mswa Hill, Mtoa Bay, Mtondwe Island, Mtosi Bay, Muloba Bay, Musende Bay, Muzi, Mvuna Island, Mwaka, Mwila Island, Namansi, Nausingili Island, Ndole Bay, Ngosa Point, Niamkolo, Ninde, Nkamba Bay, Nkondwe Island, Nkonkonti Bay, Nyanza-Lac, Pemba, Rumonge, Rusisi River, Samazi, Segunga, Sibwesa, Singa Island, Sumbu, Sumbwa, Tembwe, Ubwari Peninsula, Udachi, Ujiji, Ulungu Escarpment, Ulwile Island, Uvira, Wampembe, Wonzye Point, Zongwe


Suggestion on how to cite this blog article:
Karlsson, M. and Karlsson, M. (2019) An account of Lamprologini, including Neolamprologus sp. “Mwila”, a new shell-dwelling species from Mwila Island in the Kipili Archipelago, the facts about N. sp. “Eseki”, the whereabouts of Kinyamkolo, and much more. African Diving Blog, 14 March 2019. Available from: https://blog.africandivingltd.com/2019/03/lamprologinipart-1.html (accessed [day, month, year])

Part 1Part 2 – Part 3


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