Perciformes, Cichlidae) of East Africa, Based on Scale and Squamation Characters

Journal of Fish Biology (1993) 42, 903-946

A phyletic study on lacustrine haplochromine fishes (Perciformes, Cichlidae) of East Africa, based on scale and squamation characters

E. L i p p i t s c h Steingrabenweg 26, A-8044 Graz, Austria

(.Received 10 August 1992, Accepted 13 October 1992)

Phyletic relations within the haplochromine cichlids of East Africa were investigated vising scale and squamation characters. Within the L. Victoria-Edward-Kivu species flock most of the genera proposed in Greenwood’s revision could be confirmed by this approach. In addition the genera could be interrelated phylogenetically. They form two distinct superlineages comprising several genera each. The genus Axtatoiilapia as conceived by Greenwood is diphyletic. The fluviatile members of the genus form the sister taxon of the L. Victoria-Edward-Kivu flock, while the rest are a subgroup of that flock. The flock seems to be of monophyletic origin.

Key words: scale morphology; squamation; phylogenetic relationships; haplochromine fishes; Cichlidae.

I. INTRODUCTION When in his classic paper of 1920 C. T. Regan expressed his opinion on Haplochromis, being the ‘ largest African genus ’ (Regan, 1920), he determined the direction for the next 60 years of systematics in East African cichlids. That genus, introduced by Hilgendorf (1888) as a subgenus of Chromis to accommodate a Lake Victoria species with peculiar dentition, became a dumping ground for over 300 species from all over the continent. Regan’s authority prevented any large-scale revision of the complex until 1979. This is the more astonishing as no formal diagnosis for the genus was available, and the only apomorph character uniting the whole assembly (the structure of the pharyngeal apophysis) is found in a number of other genera as well. In two papers, Greenwood (1979, 1980) undertook a ‘ revision of the Haplochromis generic concept omitting, however, the Lake Malawi species. He divided the whole haplochromine assemblage into more than 20 lineages, to which he assigned generic (or, in a few cases, subgeneric) rank. This action was met with considerable reservation among ichthyologists working in the field, even though Greenwood’s competence could not be doubted. The reason for non-acceptance of Greenwood’s revision seems to be at least four-fold. Besides a natural reluctance to give up the convenient catch-all classification, Greenwood’s case was weakened by his own evidence. Only a few years earlier (Greenwood, 1974) he had presented an intrageneric classification of Haplochromis that differed considerably from his later revision, and he had argued in favour of the monophyletic origin of the assemblage. Furthermore it was virtually impossible to assign newly discovered species to Greenwood’s genera on the basis of his key characters (Snoeks et al., 1990; De Vos et ah, 1990). And finally, it seemed impractical to follow a revision excluding the Lake Malawi fishes. 903 0022- II12/93/060903 + 44 $08.00/0 © 1993 The Fisheries Society of the British Isles 904 E. LIPPITSCH

In the meantime the L. Malawi haplochromines have been revised (Eccles & Trewavas, 1989), separating all the taxa from Haplochromis and re-establishing or newly describing some two dozen genera. In addition new techniques like protein and enzyme studies (Sage et al., 1984; Verheyen et al., 1985, 1989) or DNA sequencing (Meyer et al., 1990) have revealed close similarities within the L. Victoria-Edward-Kivu assemblage, but significant differences with the L. Malawi taxa. These biochemical methods have proven valuable especially on a higher systematic level. However, while the L. Victoria ‘ species flock ’ contains at least 200 species, only 15 out of 803 positions in two segments of mitochondrial DNA turned out to be variable, the mean number of differences between species being only three (Meyer et al., 1990). This shows that at present investigations on relationships within the assemblage have still to be done by morphological methods. The species considered are rather uniform, however, also with respect to the usual morphological characters. The apomorphic characters worked out by Greenwood (1979,1980) are mainly associated with the trophic apparatus, but it is well-known that those characters are strongly subject to selection pressure and ecophenotypic effects. Thus, even if apomorphic similarities can be found, it is extremely difficult to assert their synapomorphic status. As Greenwood (1980) expressed it: ‘ Clearly, if sister groups are to be identified,... there is need for... the use of characters other than strictly anatomical ones ’. In previous papers (Lippitsch, 1989, 1990, 1991, 1992) the potential of scale and squamation characters for cichlid systematics have been investigated. The results indicate that there exist a large number of useful characters, that these characters are strongly determined genetically with little intraspecific variation, that many of them are independent from each other, and that their distribution within the Cichlidae is obviously governed by phyletic relationships. The present work investigates scale and squamation characters among haplochromines from the L. Victoria-Edward-Kivu assemblage, assesses character states and polarity, and presents phyletic hypotheses concerning relationships between taxa within the assemblage and those with outside groups.

II. MATERIAL AND METHODS This paper investigates only the strictly lacustrine taxa revised by Greenwood (1980). Nevertheless, the choice of species to be studied was made to cover fully the supposed lineages ranked as genera or subgenera in Greenwood’s revision (Greenwood, 1979, 1980) including the fluviatile ones except for Chetia and Pharyngochromis. At least the type species of each genus or subgenus was studied, with the exception of Ctenochromis where the type species (C. pectoralis Pfeffer 1893) was not available for inspection. One or more additional species were included from most of the genera, and a number of species from other genera, especially from L. Malawi, were studied. So a reasonable basis (59 species out of an estimated 250 from genera present in Lakes Victoria, Edward and Kivu and 30 outside species) is available for assessing the distribution of character states and drawing phyletic conclusions. Scale and squamation characters in about 130 further species of cichlids (from Africa, Madagascar, India, and America) have been investigated. A list of specimens used from those species, which have immediate relevance for the present study, is provided in the Appendix. Methods of investigation and the terminology of squamation patterns and scale morphology used were described in previous papers (Lippitsch, 1989, 1990, 1991, 1992). A PHYLETIC STUDY OF HAPLOCHROMINE FISHES 905

Emphasis was laid on documenting as fully as possible details of the squamation pattern and the surface structure of flank scales. Scales were removed from the flank, usually from the second row below the upper lateral line and at least the sixth scale column behind the insertion of the pectoral fin. In part of the BMNH material (especially fishes acquired during the 1950s) the scale surface had deteriorated as a result of storage in formalin, so that some details could not be determined with certainty. In addition many fishes from all museums had been eviscerated and part of the ventral squamation destroyed. For convenience, throughout this paper the term ‘ Victoria-Edward-Kivu assemblage ’ is used to denote the haplochromines of those lakes as well as other water bodies in connection with them. The generic names proposed by Greenwood (1979, 1980) are used for the assemblage and those of Eccles & Trewavas (1989) for L. Malawi fishes. This does not imply full endorsement of either revision. The term haplochromine will be used, again for convenience only, to cover all taxa considered belonging to or being closely related to, Haplochromis by Regan, including Astatoreochromis and the monotypic genera, but excluding the taxa of Greenwood’s Section II (Serranochromis, Sargochromis, Chetia, Pharyngochromis', Greenwood, 1979).

II!. SCALE AND SQUAMATION CHARACTERS: STATES, POLARITY, AND DISTRIBUTION

During comparative investigations on cichlid scale and squamation characters conducted for 4 years and encompassing about 190 species from over 85 genera, in addition to the usual scale counts, 96 different characters were established, representing about 300 distinguishable character states (Table I). A discussion, of the respective character states and their polarity is given below. Meristic characters, which are usually given in species descriptions, are omitted since in the present context they seem to provide no phylogenetic insight. Polarity assessment in the case of cichlids is particularly difficult. Monophyletic origin of the family has been well established (Stiassny, 1981), and the monophyly of the majority of African cichlids has been claimed with strong arguments (Cichocki, 1976; Stiassny, 1990, 1991). It is, however, unclear whether the haplochromines in the broadest sense are a monophyletic assemblage, and no outgroups have been delimited. To circumvent these difficulties, character polarity is discussed with respect to the plesiomorphic conditions of what will be called the ‘ African assemblage \ that is most of the African taxa, but excluding Tylochromis, and Helerochromis (and possibly a few others, to be discussed elsewhere). The plesiomorphic character states of this assemblage have been assessed by using Indian, Malagassy, and American taxa as well as African non-members of the assemblage for outgroup comparison and applying (with due reservations) the commonality principle within the assemblage. A full account of the underlying investigations will be given elsewhere. Here it is shown that on this basis it is possible to draw conclusions on phyletic relations within the haplochromines.

OPERCULAR SQUAMATION (TABLE I, 1-9) The plesiomorphic status for the Cichlidae on the basis of outgroup as well as intrafamilial comparison is full squamation of the operculum, suboperculum, and interoperculum. As a synapomorphy, the African assemblage shares the peculiarity of a scaleless opercular blotch in the dorsocaudal angle of the operculum which is often conspicuously pigmented, mostly black (melanin), but in some taxa is outlined with blue or golden iridiophores. This blotch, with its smooth 906 E. LIPPITSCH

T a b le I. Characters used in this study, and respective character states

1. Operculum 11. Cheek scales scaleless I cycloid 1 partially scaled 2 with vestigial ctenii 2 scaled except opercular blotch 3 weakly ctenoid 3 fully scaled 4 strongly ctenoid 4 2. Opercular scales 12. Postorbital squamation cycloid 1 single column 1 with vestigial ctenii 2 ‘mixed’ columns 2 weakly ctenoid 3 double or triple columns 3 strongly ctenoid 4 multiple columns 4 3. Opercular blotch, if present missing 5 large, pigmented 1 13. Lachrimal large, not pigmented 2 scaleless 1 small, pigmented 3 partially scaled 2 small, not pigmented 4 fully scaled 3 4. Suboperculum 14. Scales on lachrimal cycloid scaleless 1 cycloid 1 partially scaled 2 with vestigial ctenii 2 scaled except caudoventral rim 3 weakly ctenoid 3 scaled except rostral part 4 strongly ctenoid 4 fully scaled 5 15. Occiput 5. Subopercular scales scaleless 1 cycloid 1 partially scaled 2 with vestigial ctenii 2 fully scaled 3 weakly ctenoid 3 strongly ctenoid 4 16. Scales on occiput 1 6. Interoperculum cycloid with vestigial ctenii 2 scaleless 1 weakly ctenoid 3 scaled on the caudal part only 2 strongly ctenoid 4 with scaleless ventral rim 3 fully scaled 4 17. Size of occipital scales 7. Interopercular scales compared to dorsal scales cycloid 1 not significantly smaller 1 with vestigial ctenii 2 significantly smaller 2 weakly ctenoid 3 extremely small 3 strongly ctenoid 4 18. Predorsal squamation pattern 8. Preoperculum scaleless 1 scaless 1 uniserial 2 partially scaled 2 biserial 3 fully scaled 3 triserial 2/1 4 triserial 1 /2 5 9. Preopercular scales irregular 6 cycloid 1 with vestigial ctenii 2 19. Dorsum, rostrally weakly ctenoid 3 scaleless I strongly ctenoid 4 partially scaled 2 fully scaled 3 10. Cheek scaleless 1 20. Scales on dorsum, rostrally partially scaled 2 cycloid 1 scaled except ventral rim 3 with vestigial ctenii 2 scaled except rostral embayment 4 weakly ctenoid 3 fully scaled 5 strongly ctenoid 4 A PHYLETIC STUDY OF HAPLOCHROMINE FISHES 907

T a b l e I. Continued

21. Dorsum, caudally under right angle to rim 3 scaleless 1 turning outward 4 partially scaled 2 irregular 5 fully scaled 3 obscured by granulation 6 22. Scales on dorsum, caudally 28. Form of granular area cycloid 1 sectorial 1 with vestigial ctenii 2 segmental 2 weakly ctenoid 3 circular arch 3 strongly ctenoid 4 29. Size of granular area small (less than 90°) \ 23. Flank medium (90-150°) 2 scaleless 1 3 partially scaled 2 large (more than 150°) fully scaled 3 30. Scale focus free of granulation 1 24. Flank scales covered by granulation 2 cycloid 1 with vestigial ctenii 2 31. Radii on flank scales weakly ctenoid 3 simple I strongly ctenoid 4 broadened 2 filled with tissue 3 25. Flank scale overall form 32. Caudal rim of flank scales ovoid, log axis vertical 1 1 circular 2 normal soft 2 long axis horizontal 3 weakly ossified 3 26. Flank scale granulation type 33. Form of ctenii 1 1 ylochromis short, talon-shaped 1 Sarotherodon 2 short, bluntly conical 2 Oreochromis 3 long, conical 3 Chr. kingsleyae 4 long, subcylindrical 4 Chr. guntheri 5 long, slender, bent or hooked 5 Hemichromis 6 Anomalochromis 7 34, Interradial denticles of flank Pelmatochromis 8 scales Pseudocren ilabrus 9 slender 1 Astatotilapia burtoni 10 blunt 2 Astatotilapia elegans 11 with neck-like narrowing 3 Melanochromis 12 35. Interradial tongues of flank Hoplotilapia 13 scales Lamprologus 14 missing 1 Pterophyllum 15 short 2 Apistogramma 16 long 3 Astatoreochromis 17 36. First interradial circuli Ct. polli 18 concave 1 Protomelas 19 straight 2 Nimboehromis 20 convex 3 Progna thochromis 21 37. Region covered by pectoral fins Paraiilapia 22 scaleless 1 Harpagochromis 23 partially scaled 2 Tropheus 24 fully scaled 3 Psammochromis 25 38. Scales covered by pectoral fins Serranochromis 26 cycloid 1 27. Circuli on flank scales with vestigial ctenii 2 roman 1 weakly ctenoid 3 gothic 2 strongly ctenoid 4 908 E. LIPPITSCH

T able I. Continued

39, Caudal peduncle 50. Median scales of ventral chest scaleless 1 uniserial 1 partially scaled 2 biserial 2 fully scaled 3 irregular 3 40. Scales on caudal peduncle 51. Squamation between pelvic fins cycloid 1 lacking 1 with vestigial ctenii 2 uniserial 2 weakly ctenoid 3 biserial 3 strongly ctenoid 4 multiserial 4 41. Squamation pattern on lower 52. Large interpelvic scale present median of caudal peduncle (LIVS) uniserial 1 present 1 biserial 2 absent 2 staggered 3 irregular 4 53. Belly 42. Chest laterally scaleless 1 scaleless 1 partially scaled 2 partially scaled 2 fully scaled 3 fully scaled 3 54. Scales on belly 43. Scales on chest, laterally cycloid 1 cycloid 1 with vestigial ctenii 2 with vestigial ctenii 2 weakly ctenoid 3 weakly ctenoid 3 strongly ctenoid 4 strongly ctenoid 4 55. Size of belly scales compared to 44. Size o f lateral chest scales flank scales compared to flank scales not significantly smaller 1 not significantly smaller 1 significantly smaller, imbricating 2 significantly smaller, imbricating 2 significantly smaller, not 3 significantly smaller, not 3 imbricating imbricating 56. Transition from belly to flank 45. Transition from chest to flank scales scales gradual 1 gradual 1 abrupt 2 abrupt 2 57. Anal-genital region 46. Chest ventrally scaleless I scaleless 1 partially scaled 2 partially scaled 2 fully scaled 3 fully scaled 3 47. Scales on chest, ventrally 58. Scales on anal-genital region cycloid 1 cycloid 1 with vestigial ctenii 2 with vestigial ctenii 2 weakly ctenoid 3 weakly ctenoid 3 4 strongly ctenoid 4 strongly ctenoid 48. Size of ventral chest scales 59. Size of scales on anal-genital compared to flank scales region, compared to flank scales not significantly smaller 1 not significantly smaller 1 significantly smaller, imbricating 2 significantly smaller, imbricating 2 significantly smaller, not 3 significantly smaller, not 3 imbricating imbricating 49. Transition from ventral to 60. Dorsal fin lateral chest scales scaleless 1 gradual 1 scaled in the spiny part 2 abrupt 2 scaled in the soft part 3 A PHYLETIC STUDY OF HAPLOCHROMINE FISHES 909

T a b l e I. Continued

61. Dorsal fin scales 71. Base of anal fin cycloid 1 without pad or sheath 1 with vestigial ctenii 2 with scaleless pad 2 weakly ctenoid 3 with scaly pad 3 strongly ctenoid 4 with vestigial sheath 4 62. Dorsal fin squamation pattern with well-developed sheath 5 single rows on membranes 1 72. Scales on anal fin pad or sheath double/multiple rows on 2 cycloid 1 membranes with vestigial ctenii 2 double/multiple rows along rays 3 weakly ctenoid 3 strongly ctenoid 4 63. Base of dorsal fin 73. Pectoral fins without pad or sheath 1 scaleless 1 with scaleless pad 2 partially scaled 2 with scaly pad 3 fully scaled 3 with vestigial sheath 4 with well-developed sheath 5 74. Scales on pectoral fins cycloid 1 64. Scales on pad or sheath with vestigial ctenii 2 cycloid 1 weakly ctenoid 3 with vestigial ctenii 2 strongly ctenoid 4 weakly ctenoid 3 75. Pectoral fin squamation strongly ctenoid 4 pattern 65. Caudal fin single rows on membranes 1 scaleless 1 double/multiple rows on 2 partially scaled 2 membranes fully scaled 3 double/multiple rows along rays 3 66. Scales on caudal fins 76. Pelvic fins cycloid 1 scaleless 1 with vestigial ctenii 2 partially scaled 2 weakly ctenoid 3 fully scaled 3 strongly ctenoid 4 77. Scales on pelvic fins cycloid 1 67. Caudal fin squamation pattern with vestigial ctenii 2 broad single rows 1 weakly ctenoid 3 oblong single rows 2 strongly ctenoid 4 staggered rows on membranes 3 rows along rays 4 78. Pelvic fin squamation pattern densely covering the rays 5 single rows on membranes 1 double/multiple rows on 2 68. Anal fin membranes scaleless 1 double/multiple rows along rays 3 scaled in the spiny part 2 79. Enlarged axillary scale scaled in the soft part 3 present I 69. Anal fin scales absent 2 cycloid 1 80. Lateral line with vestigial ctenii 2 normal 1 weakly ctenoid 3 kinked 2 strongly ctenoid 4 81. Lateral line 70. Anal fin squamation pattern all scales perforated 1 single rows on membranes 1 some scales not perforated 2 double/multiple rows on 2 82. Lateral line scales membranes all with channels 1 double/multiple rows along rays 3 with channels or simple pores 2 910 E. L IP P IT S C H

T a b l e I. C o n tin u ed

83. Lateral line or caudal fin (if 89. Rows above upper lateral line, longer than two scales) caudally simple 1 90. Number of lateral line branches double 2 91. Number of scales in the upper lateral triple 3 line branch irregular 4 92. Number of scales in the second lateral not present 5 line branch 93. Number of scales in the lowest lateral Meristic characters: line branch 84. Number of rows on in ter operculum 94. Number of scales between upper and 85. Total number of rows on flank second branch, rostrally 86. Number of scales in the midlateral row 95. Same, caudally 87. Rows above upper lateral line, rostrally 96. Number of lateral line scales close to 88. Rows above lateral line, highest point dorsal base

surface, is clearly discernible from the rest of the scaly operculum. The blotch is either large [as large as or larger than a typical opercular scale, Fig. 1(a)], or small [clearly smaller than a scale, Fig. 1(b)], with essentially no intermediate forms. These are regarded as different character states. An independent evolution from the blotchless state can be excluded. Non-haplochromine taxa like the chromidotilapiines, the hemichromines, the tilapiines, or even the Lamprologini (sensu Poll, 1986) invariably have large blotches. So this can be regarded plesiomorphic. A small blotch is, therefore, treated as apomorphic. Apart from the opercular blotch the operculum is always fully scaled in haplochromines, even in taxa which show a reductions] trend in head squamation. The scales are usually the same size or a little smaller than the flank scales. Cycloid scales seem to represent the plesiomorphic condition for the African assemblage. Only two African species have been identified so far bearing ctenoid scales on the operculum. The suboperculum is scaled with cycloid scales in most African taxa. Full scaling [Fig. 2(a)] is regarded as plesiomorphic. Reduced squamation is found in some haplochromine and non-haplochromine taxa, where the ventrocaudal [Fig. 2(b)] or the rostral region of the bone is scaleless [Fig. 2(c)], and these states are regarded as apomorphic. A cycloid squamation of the interoperculum is plesiomorphic. Usually scales are present in a single row, which extends only over the caudal half of the bone, a condition regarded as plesiomorphic [Fig. 3(b)]. Caudally, there may be two or more rows. Some species have a row of scales extending to nearly the rostral end of the bone [Fig. 3(a)], and a few have a scaleless inter operculum. Those states are regarded as apomorphic. The preoperculum is scaleless in most African species investigated. Outgroup comparisons suggest that this is the plesiomorphic state.

SQUAMATION OF THE CHEEK AND LACHRIMAL BONE (TABLE I, 10-14) Many species retain the plesiomorphic state of the cheek squamation with full coverage with cycloid scales of the area delimited by preoperculum, orbit, mouth, and lachrimal bone [Fig. 4(a)]. Three apomorphic character states were observed. A PHYLETIC STUDY OF HAPLOCHROMINE FISHES 911

Fig. 1. Scaleless opercular spot (stippled), large (a) in Astatotilapia burtoni (NMW 89977:2), small (b) in Orthochromis malagaraziensis (BMNH 1952.11.4.46),

In the first the lower rim of the cheek is scaleless. The width of the naked strip may differ even between one side of the fish and the other. The second consists of a naked embayment in the rostroventral rim of the scaled area, near the corner of the mouth [Fig. 4(b)]. These two states may occur simultaneously. The third is an extensive reduction of the cheek squamation, leaving a large portion or the whole of the cheek scaleless. Cheek squamation, if present, extends into the space between the orbit and the vertical limb of the preoperculum. The plesiomorphic condition is given by a single 912 E. LIPPITSCH

Fig. 2. Suboperculum, fully scaled (a) in Asiatotilapia burtoni (NMW 89977:2), with scaleless ventrocaudal rim (b) in Harpagochromis altigenis (BMNH 1984.2.6.55), and with scaleless rostral part (c) in Thoracochromis buysi (BMNH 1984.2.6.55). Scaleless parts stippled. scale column filling that post-orbital space [Fig. 5(a)], as seen in the majority of cichlids. Two apomorphic states are found in some haplochromines. The first has a more or less irregular pattern consisting of normal scales and much smaller ones [Fig. 5(b), called ‘ mixed columns ’ in this paper]. The second has two or three columns of large scales of equal size [Fig. 5(c)]. More columns may be present in non-haplochromine taxa, but are always accompanied by a reduction in scale size. Post-orbital scales are often lost during capture of the fish. This was so in several specimens from the British Museum. Thus for a few species the state of that character could not be determined. The lachrimal bone is scaleless in nearly all cichlids, and seems to be so in labroid outgroup taxa. Hence this condition can be regarded plesiomorphic. Within the haplochromines, members of the genus Pseudocrenilabrus frequently have some scales extending from the cheek onto the lachrimal, but this seems to be autapomorphic for the genus.

SQUAMATION OF HEAD AND NUCHAL REGION (TABLE I, 15-18) The squamation pattern on the upper surface of the head and nuchal region is rather difficult to evaluate. It seems likely that squamation in both regions is formed independently during ontogeny (Sire & Arnulf, 1989). The consequence is that at least five different predorsal squamation patterns can be distinguished (Lippitsch, 1990): uniserial (one median row of scales from the forehead up to the first dorsal spine), biserial (two rows), two different triserial patterns (one median A PHYLETIC STUDY OF HAPLOCHROMINE FISHES 913

Fig. 3. Interoperculum, fully scaled (a) in Lipochromis maxillaris (BMNH 1958.1.16.172), with reduced squamation (b) in Harpagochromis spekii (BMNH 1977.5.24.72). Scaleless part stippled. row on the head and two in the nuchal region, termed ‘ triserial 1/2 ’ in this paper; or two rows on the head and a single nuchal row, termed ‘ triserial 2/1 ’), and an irregular pattern. From comparisons among African cichlids and outgroup taxa a single median row on the head seems to be plesiomorphic for the African assemblage. Both triserial arrangements seem to be closely related to the uniserial one. Most probably they have evolved several times within the cichlids. The biserial predorsal pattern is rare, and obviously presents an apomorphic condition evolved independently. It is not clear whether the irregular pattern can be regarded as a well-defined character state. The plesiomorphic condition for the African cichlids is that of cycloid scales, on the forehead as well as on the nuchal region, with a size comparable to those on the dorsum. Haplochromines show an apomorphic trend towards smaller scales.

DORSAL SQUAMATION (TABLE I, 19-22, 87-89, 96) Dorsal squamation involves qualitative and meristic characters. The latter have not been evaluated so far and are not dealt with further. Two regions of the dorsum have to be distinguished. The rostral region borders upon the nuchal region and is delimited ventrally by the lateral line ascending dorsad and caudad. Ctenoid scales are regarded as plesiomorphic in the cichlids for this region. However, in haplochromines a definite trend to reduce the ctenii is present, and in many taxa the scales are even cycloid. These conditions are regarded as two distinct apomorphies. The caudal region of the dorsum extends from the highest point 914 E. LIPPITSCH

Fig. 4. Cheek, fully scaled (a) in Astatotilapia burtoni (NMW 89977:2), with scaleless restroventral embayment (b) in Thoracochromis buy si (BMNH 1984.2.6.55). Scaleless part stippled. of the upper lateral line caudad to its end. Ctenoid scales are obviously plesiomorphic. Reduced ctenoidy or presence of cycloid scales are regarded as different apomorphies. All haplochromines have a plesiomorphic fully scaled flank, and the scales are predominantly strongly ctenoid (Table I, 23-24). Reduced ctenoidy is considered apomorphic. The surface structure of flank scales provides a number of additional characters. These are dealt with below. Sometimes the region just behind the pectoral fin bases differs in scale morphology from the flank (Table I, 37-38). This is regarded as an apomorphic condition. A full, strongly ctenoid squamation on the caudal peduncle (Table I, 39^41) is plesiomorphic. As on the flanks, apomorphic reductions in ctenoidy occur. In most cases the caudal peduncle is ctenoid to the same or even stronger degree than the flank, but exceptionally the opposite condition may prevail. The lower median of the caudal peduncle usually has a single scale row (plesiomorphic), but in a few cases a staggered or an irregular pattern occurs. A double row pattern is not found among haplochromines.

SQUAMATION OF THE CHEST (TABLE I, 42-52) The plesiomorphic condition is full scaling on the lateral as well as the ventral surface of the chest. Naked areas on both regions may occur and are considered A PHYLETIC STUDY OF HAPLOCHROMINE FISHES 915

F ig. 5. Post-orbital squamation, single column (a) in Lipochromis microdon (BMNH 1977.1.28.26), mixed columns (b) in Aslatotilapia ftroiv’«ae(BM NH 1962.3.2.76), two columns (c) in Harpagochromisspekii (BMNH 1977.5.24.72). apomorphic, as are cycloid scales. The scales on the chest in haplochromines are, in contrast to the plesiomorphic state, mostly smaller than those on the flank. As first recognized by Greenwood (1971,1974,1979), the transition between chest and flank scales is a most distinctive character. The plesiomorphic state is a smooth transition with scale size changing gradually over several columns. The first apomorphic state is an abrupt size change between neighbouring columns, usually near the line connecting the bases of pectoral and pelvic fins. The second is that of an abrupt transition, the border of the small-scaled area running from the pectorals first downward and then backwards, delimiting a region of small scales on the belly against the large-scaled flank. Both states have been excellently figured by Greenwood (1979, Figs 2 and 3). It is assumed here that both states have independently evolved from the ancestral condition. In fact, however, it is not even clear with certainty, whether either of the states represents a single well- defined apomorphy. Ctenoid scales are regarded as plesiomorphic. Apomorphic reductions in ctenoidy may occur independently on the lateral and ventral surface of the chest. The scales of the chest extend to between the bases of the pelvic fins. The plesiomorphic condition in many haplochromine and non-haplochromine cichlids as well as in labroid outgroup taxa is a single, exceptionally large scale between the pelvics. One small scale or more than one scale row is apomorphic. Unfortunately this character cannot be examined in eviscerated fish and remains unclear for many species. 916 E. LIPPITSCH

f s z O X i i - i -

Fig. 6. Caudal fin squamation, on the membrane (a) in AstatotUapia burtoni (NMW 89977:2), along the rays (b) in Lipochromis maxillaris (BMNH 1950.1.16.172). Fin rays and exposed part of membrane stippled.

What has been said for the chest also holds for the belly (Table I, 53-59). Very small as well as cycloid scales are considered apomorphic, and the size-transition to the flank scales has been discussed above. It is interesting that in some taxa the anogenital region shows different scales from the rest of the abdomen. This condition is also regarded as apomorphic. Fortunately, among the L. Victoria-Edward-Kivu species, scaly dorsal, anal, and pectoral fins occur only in a single species. Thus, this set of characters, which is problematic for the whole family, presents no difficulty in this context. For the caudal fin the plesiomorphic squamation pattern is that of single or multiple rows on the membranes [Fig. 6(a)]. Scales positioned along the rays [Fig. 6(b)], and scales covering the rays (very rare among the species considered here) are treated as apomorphic. A scaly sheath at the base of the dorsal or anal fin is found in no haplochromine taxon. Sometimes one or a few rows of scales are present at the base of the anal fin, A PHYLETIC STUDY OF HAPLOCHROMINE FISHES 917 which are clearly not fin scales, but seem to represent the rudiments of a sheath. They are usually almond-shaped with their long axes aligned horizontally, while fin scales are oriented in parallel with the rays. Those scales are the ‘ anal sheath scales’ of Greenwood (1984) and represent an apomorphic condition. Another apomorphic feature present at the dorsal and/or anal fin bases of various taxa is a basal pad, that may be naked or scaly.

FLANK SCALES (TABLE I, 25-37) All haplochromine (and all cichlid) flank scales are of the sectional type (Lippitsch, 1990). They usually have an ovoid outline with a vertical long axis, regarded as plesiomorphic. More or less circular scales or those with the horizontal width greater than the height are apomorphic. In many haplochromines the interradial tongues on the rostral rim of the scale are very short, but long tongues are encountered also. Short tongues with convex first circuli are considered to be plesiomorphic. The interradial denticles show no characteristic features, with the exception of Pseudocrenilabrus multicolor, where they have a neck-like narrowing. At least part of the caudal field of a flank scale bears a characteristic granulation (Table I, 26-30). Fifteen different types of scale granulation have been distinguished (Lippitsch, 1990), and even more can be recognized (see below). A serious difficulty arises from the fact that up to now it was not possible to establish the plesiomorphic granulation type for the African assemblage. This is due to the diversity of scale morphology encountered within and outside that assemblage and to the fact that obviously apomorphic conditions occur even in otherwise primitive taxa. For example, ctenoid scales have to be regarded plesiomorphic from comparison with taxa from India, Madagascar and America. Cycloids are present, however, in Tylochromis, a genus considered to be the sister taxon to the whole African assemblage (Stiassny, 1990). It seems that the plesiomorphic granulation type of the African assemblage has survived in none of its members studied so far. Nonetheless, from the multitude of taxa investigated it seems possible to reconstruct the plesiomorphic state. Small dentritic ridges extending from the circuli onto the floor of the circular grooves are present even in Paratilapia and Heterochromis and hence seem to be plesiomorphic for the African assemblage. The exposed part of the scale is almost completely covered with granulation. From a little behind the focus to about two-thirds of the way to the caudal rim granulation consists of oblong protrusions with their long axes in a radial direction. These protrusions follow more or less the course of the (obscured) circuli. The protrusions of successive circuli are displaced laterally with respect to each other so that a staggered pattern results. The surface of the protrusions bears irregular ridges which again may extend onto the floor of the grooves. Near the caudal rim strong ctenii, lying flat on the surface and pointing to the rim, form a broad and distinct stripe. The ctenii have a smooth surface without ridges. All the types described in the following differ from this plesiomorphic state in various ways and are regarded as apomorphic. Only numbers 10 to 13 of the previously defined types occur in lacustrine haplochromines. For present purposes it is necessary to re-examine those types. In addition, four more types are described. G ran u latio n type 11 is sim ilar to the plesiom orphic state. T he only peculiarity is the admixture of rounded or irregular-shaped tubercles to the oblong protrusions near the rostral border of the granular area [Fig. 7(b)]. 918 E. LIPPITSCH

Fig. 7. Haplochromine flask scale granulation types. Type 10 (a) in Astatotilapia bioyeti (MT 191 162-201), type 11 (b) in Astatotilapia lacrimosa (BMNH 1959.4.28.1), type 13 (c) in Ptyochromis sauvagei (RM NH unreg.), type 17 (d) in Astatoreochromis vanderhorsti (BM NH 1953.11.4.59).

A very distinct granulation type is no. 10 [Fig. 7(a)], characteristically displayed in species of Astatotilapia s. str. (see below). Usually no dendritic ridges are present. Instead, rounded, verrucous tubercles are present on the circuli. Extensive granulation is present, consisting of rounded tubercles which are never oblong and aligned radially. They are distributed stochastically and do not display a circular and staggered distribution. The granulation fills the whole caudal field of the scale, and ctenii are present only on the very edge of the scale. Type 12 was described for Melanochromis auratus of L. Malawi in the previous work. Since then it became clear that the L. Victoria species thought to belong to that type in fact do not. This type is not considered further. A PHYLETIC STUDY OF HAPLOCHROMINE FISHES 919

Type 13 in the previous paper was characterized by a peculiar course of the circuli (turning outward). This feature, however, is independent of the kind of granulation, and the course of the circuli now is regarded as an independent character (see below). Nevertheless, the type can be maintained as distinct even based on granulation alone [Fig. 7(c)]. Dendritic ridges from the circuli may be present, but are much reduced compared to type 11. The granulation consists of protrusions of irregular shape (not so distinctly rounded as in type 10) in the central part of the scale’s caudal field. The rest is filled with a stripe of strong ctenii, lying flat on the surface and being radially directed. The width of this stripe differs between taxa. Many species investigated in this study could not assigned to the previously described types and a number of new types had to be established. Those related to haplochromine taxa include the following. Type 17. Typical species Astatoreochromis vanderhorsti [Fig. 7(d)]. The overall appearance of the granulation resembles that of type 11, but the ridges on the granules are more distinct and form a nearly reticular pattern near the centre of the scale. The granules are small and not very regular in outline. Only a narrow stripe near the caudal edge of the scale bears smooth ctenii. Type 21. Typical species Prognathochromis prognathus [Fig. 8(a)]. Again a reticular pattern is present, forming connecting ridges between the granules, which are long and slender and arranged radially. Type 23. Typical species Harpagochromis spekii [Fig. 8(b)]. This type has rounded tubercles and resembles type 13, but the reticular ridges are developed much more strongly and the tubercles are smaller. Type 25. Typical species Psammochromis saxicola [Fig. 8(c)]. This type is characterized by oblong granules with moderately developed ridges. In the middle of the granular field the long axes of the granules are directed horizontally and not, as in other types, radially. The form of the granular area is that of a sector in all haplochromines. The extension of the granular sector may range from less than 90° [Fig. 9(a)] to 180° [Fig. 9(c)]. Large angles are regarded as plesiomorphic. Usually the granulation does not reach the focus of the scale. An extension of the granulation to or beyond the focus is treated as an apomorphy. Six character states have been distinguished for the course of the circuli in the caudal field (Table I, 27). In haplochromines, the circuli are mostly obscured by granulation. If they can be distinguished, the ‘ Roman ’ pattern predominates, and is regarded as plesiortiorphic. In some taxa, the circuli turn outward to the rim in a characteristic way, as described for type 13. This state is clearly apomorphic. In haplochromines the caudal rim of flank scales always bears ctenii, irrespective of the kind of granulation. This is an ancestral feature within the Cichlidae. The plesiomorphic shape of the ctenii is long and conical, with a straight and pointed tip [Fig. 10(a)]. Apomorphic shapes are short and talon-shaped (rare in haplochromines); short and bluntly conical [Fig. 10(b)]; subcylindrical with an obtuse tip [Fig. 10(c)]; or slender with a sharp, often curved or even hook-like tip [Fig. 10(d)], In haplochromines, as in other cichlids, there may be some kind of variation in scale morphology between individuals of the same species or even between scales on the same individual. Different parts of the body may bear completely different 920 E. LIPPITSCH

0-2 nun

Fig. 8. Haplochromine flank scale granulation types. Type 21 (a) in Prognathochromis prognalhus (BMNH 1966.3.9.53), type 23 (b) in Harpagochromis spekii (BM NH 1977.5.24.72), type 25 (c) in Psammochromis saxicola (BMNH 1959.4.28.250). scales. Granulation types are defined here only for the flank scales, and only those scales are compared. One has to be cautious to sample well off the pectoral fins, since scales of the region immediately behind the pectoral fin insertion may have a different granulation. It has to be assured that only regular-grown scales with a well-developed focus are used. Even then there remains some variation, and not all regular-grown flank scales on a single fish may exhibit all the features characterizing a particular granulation type. In most cases, however, they can be unambiguously discerned, at least if several scales are inspected. Of 83 characters evaluated only 10 were invariably plesiomorphic (Table II). On the other hand, only a single apomorphic feature (the size reduction in A PHYLETIC STUDY OF HAPLOCHROMINE FISHES 921

FiG. 9. Extent of granular area, small (a) in Thoracochromis buy si (BMNH 1984.2.6.56), medium (b) in ‘ Haplochromis' occultidens (MT 80-49-P-5996), large (c) in Yssichromis fusiformis (BMNH 1968.8.30.29).

0-2 mm

Fig. 10. Form of ctenii, long and conical (a) in Gaurochromis iris (RMNH 4-1-1978), short and bluntly conical (b) in Astatotilapia flaviijosephi (BMNH 1949.9.16.390), subcylindrical (c) in AUochromis wellcomei (BMNH 1965.8.6.2), slender and hooked (d) in Plyochromis sauvagei (RMNH unreg.). T a b l e II. Character state distribution among the haplochromine species examined in this study (see also comment at the end of table)

Taxon/Character I 2 3 4 5 6 7 8 9 10 11 12 13

Plesio. Africa 3 1 1 5 I 2 1 1 _ 5 1 1 1 adolphifrederici 3 1 1 5 1 2 1 1 — 5 1 1 1 alluaudi 3 1+ 3 2 5 1+3 2 1 1 5 1 1 1 altigenis 3 1 1 3 1 2 1 1 — 4 1 I astatodon 3 1 1 5 1 2 1 1 — 5 1 1 1 bakongo 3 1 1 5 1 2 1 1 — 3 1 1 1 bicolor 3 1 3 3 1 2 1 1 — 5 1 1 I bloyeti 3 1 I 5 1 2 I 1 — 5 1 1 brownae 3 1 1 5 1 2 1 1 — 5 1 1,2? 1 burtoni 3 1 1 5 1 2 1 1 — 5 1 1,2 1 buysi 3 1 1 3 1 2 1 I — 3 + 4 1 1,2 1 caI dpt era 3 I I 5 1 2 I 1 — 5 I 1 I cassius 3 1 1 5 1 2 1 1 — 5 1 1?2 1 cinerea 3 1 1 3 1 2 1 1 — 5 1 1 1 cnester 3 1 1 5 I 4 1 1 — 5 1 1 1 codringtoni 3 1 3 2 1 2 I 1 — 5 1 1 1 cronus 3 1 1 3 1 2 1 1 — 5 1 1 1 degeni 3 1 3 5 1 2 1 1 — 4 1 1 1 desfontainesii 3 1 I 5 1 2 1 1 — 5 1 1 1 eduardiana 3 1 1 5 1+ 2 2 1+2 1 — 5 1 1 1 elegans 3 1 1 5 1 2 1 1 — 5 1 1 1 empodisma 3? 1 1 5 1+ 2 4,2 1+ 2 1 — 5 1 1 1 erythrocephalus 3 1 1?3 5 1,2 4 1+ 2 1 — 5 1 1 1 flaviijosephi 3 1 1 5 1 2 1 1 -- 4? 1 1 1 fusiformis 3 1 1 5 2 4 1 1 — 5 1 1? 1 gracilior 3 1 1?3 5 1 4 1 1 — 5 1 1 I graueri 3 1 1? 5 1 2 1 1 — 5 1 1 1 hiatus 3 1 I 5 1? 4 1 I — 5 1 1 I horei 3 1 1 3 1 1 — 1 — 2 1 1 1 iris 3 1 1 5 1? 4 1 1 — 5 1 1 1 ishmaeli 3 1 1 5 1? 2 1 1 —■ 5 1 1 1 lacrimosa 3 1 1 5 1 2 1 1 — 4 1 1 I mahagiensis 3 1 1 2 1 1? — 1 — 3 1 1 1 malagaraziensis 3 3 4 1 maxillaris 3 1 5 1+ 2 microdon 3 1 5 1+2,3 multicolor 2 I 2 I nigricans 3 I 3 1 nigripinnis 3 1 5 1 nubiia 3 1 5 1 nuchisquamulatus 3 1 5 1 obesus 3 1 5 1,2 obliquidens 3 I 5 1 obiusidens 3 1 5 I occultidens 3 1 5 1 orthostoma 3 1?3 5 1 parorthostoma 3 1?3 3 1 parvidens 3 1 5 1 paucidens 3 I 5 1 pectoralis 3 1 5 2+1 polli 3 1 5 1 prognathus 3 1 5 1 ptistes 3 1 5 1 retrodens 3 3 5 1 riponianus 3 1 5 1 sauvagei 3 3 5 1 saxicola 3 1 5 I serranus 3 1 5 1 spekii 3 1 5 1,2 squamipinnis 3 1 5 1 straeleni 3 2 5 1 teegelaari 3 1 5 1+ 2 thumbergi 3 3 2 I torrenticola 3 1?3 5 1 tridens 3 1 5 1 vanderhorsti 3 2 3 1,2 victoriae 3 1?3 5 1 victorianus 3 1 5 1,2 vittatus 3 1 5 1 we Icommei 3 1 5? I wingatii 3 1 5 1 1 _ 1 — 1 — 1,5 1 4 1 1 — 5 1 1,2 1 4 1 + 2 1 — 5 1 1 1 2 1 1 — 4 1 1 1 2 1 1 — 4 1 1 1 2 I 1 — 5 1 1 1 2 — 5 1 1 1 2 1 1 — 5 1 1 1

2 1 1 — 4 1 1,2 FISHES 1 INE HAPLOCHROM OF STUDY PHYLETIC A 2 1 I — 5 1 1 1 4 1 1 — 5 1 1 1 2 1 1 — 5 1 1 1 2 I 1 — 5 1 2 1 2 1 1 — 5 1 2 1 4 1 1 — 5 1 1 1 1,2 — 1 — 5 1 I 1 2 1 1 — 5 I 3 1 1 t1 1 Ii 2 1 1 _ 5 1 1? 1 4 I 1 — 5 1 1 1 2 1 1 — 5 1 1,2 1 2 1 1 — 5 1 1,2 1 2 1 1 — 5 1 I 1 2 1 1 — 5 1 1 1 2 1 1 — 5 1 3 1 2 I 1 — 5 1 3 1 27 1 1 — 5 1 1 1 4 1 2 1 5 1 1 1 4 1 1 — 5 1 1 1 1 -- 1 — 5 I 2,3 1 1 _ 1 — 3 1 2,3 1 2 1 1 — 5 1 1? 1 4 1 2 1 5 1 1 1 4 1 1 5 1 1 1 4 1+2 1 — 5 1 2,3 1 2 1 1 — 5 1 2,3 1 2 1 1 — 5 1 2? 1 2 1 1 _ 3 1 1 1 T a b l e II.

Taxon/Character 14 15 16 17 18

Plesio. Africa — 3 1 1 2 adolphifrederici — 3 1 1 2 alluaudi — 3 1 2 ? altigenis — 3 1 2 2 astatodon — 3 1 2 2 bakongo — 3 1 2 2 bicolor — 3 1 2 4 bloyeti — 3 1 2 5 brownae — 3 1 2 2 burtoni — 3 1 2 2 buy si — 3 1 1 2 calliptera — 3 1 2 2 cassius — 3 1 2 6 cinerea — 3 1 2 2 cnester — 3 1 2 2 codringtoni — 3 1 1 4 cronus — 3 1 2 6 degeni — 3 1 2 6 desfontainesii — 3 1 2 2 eduardiana — 3 1 2 2 elegam — 3 1+ 2 2 2 empodisma — 3 1 2 4 erythrocephalus — 3 1 2 2 fiaviijosephi — 3 1 2 6 fusiformis — 3 1 2 2 gracilior — 3 1 2 2 graueri — 3 1 2 4 hiatus -— 3 1 2 2 horei — 3 1 1 3 iris — 3 1+2 2 2 ishmaeli — 3 1 2 4 lacrimosa — 3 12 2 mahagiensis — 3 1 1 5 C o n tin u e d

19 20 21 22 23 24 25 26

3 3 3 3 3 3 1 ? 3 2 + 3 3 3 3 4 1 11 3 2 3 3? 3 4 1 17 3 1 3 1,2 3 3 1 23 3 1 3 1 3 3 1 13 3 1+ 2 3 4 3 4 18 3 1 3 1 3 3 1 13 3 1 3 2 + 3 3 3 1 10 3 1 3 2 + 3 3 3 1 21 ? 11 3 1 3 3 3 3 1 10 3 1 3 1 3 1+ 2 + 3 1 8? 3 1,2 3 3 3 3 1 10 3 1 3 3 3 3 1 21? 3 1 3 1+ 2 3 3 1 11 3 2 3 3 3 4 1 11 3 1 3 1 3 1+ 2 + 3 1 8? 3 1 3 1 3 2 + 3 1 10? 3 1+ 2 3 1 3 3 1 13 3 1+2 3 2 + 3 3 3 1 10 3 1+ 2 3 3 3 3 1 23 3 3 3 4 3 4 1 11 3 1+ 2 3 3 3 3 1 11 3 1 3 3 3 3 1 11 3 1+ 2 + 2 3 2 + 3 3 3 1 10? 3 2 + 3 3 3 3 3 1 23? 3 1 3 2 3 3 1 11 3 1+ 2 3 3 3 3 1 11 3 1 3 3 3 3 1 11 3 1 3 1 3 2 + 3 1 10? 3 1+ 3 3 4 3 4 1 11 3 1+ 2 3 2 + 3 3 4 1 11? 3 1 3 2 3 3 1 11 3 1 3 1+ 2 3 4 1 9? malagaraziensis — 3 1 1 6 maxillaris — 3 1 2 1 microdon — 3 1+ 2 2 1 multicolor — 3 1 1 5 nigricans — 3 \ 2 7 nigripinnis — 3 1+ 2 2 2 nubila — 3 1 2 2 nuchisquamulatus — 3 3 2 2 obesus — 3 1 2 2 obliquidens — 3 1 2 4 obtusidens — 3 1 2 4 occuhidens — 3 1 2 2 orthostoma — 3 1 2 ? parorthostoma — 3 1 2 6? parvidens — 3 1 2 2 paucidens — 3 1 2 4 pectoralis __ 3 1 2 2 polli — 3 I 2 3 prognathus — 3 I 2 2 ptistes — 3 1 2 2 retr odens — 3 1 2 2 riponianus — 3 1 2 2 sauvagei — 3 1 2 2 saxicola — 3 1 2 6 serranus — 3 1 2 2 spekii __ 3 1 2 2 squamipinnis — 3 1 2 2 straeleni — 3 1 1 2 teegelaari — 3 1 2 4 thumbergi — 3 1 2 6 torrenticoia — 3 1 2 3 tridens — 3 1 2 3 vanderhorsti — 3 1 1 2 victoriae — 3 1 2 7 victorianus — 3 1+2 2 6 vittatus — 3 1 2 2 welcommei — 3 1 2 2 wiiWatii — 3 I 1 7 1+ 2 + 3 3 3 3 4 1 9? 1+ 2 3 3 3 3 1 23 2 + 3 3 3 3 4 1 23? 1 3 3 3 3 1 9 1 3 1 3 2 + 3 1 10? 3 3 4 3 4 1 11 1 3 2 + 3 3 3 1 10 1 3 1 3 3 1 13

1+ 2 3 3 3 3 + 4 1 23 FISHES INE PLOCHROM HA OF STUDY PHYLETIC A 1 3 1 + 2 3 3 + 4 1 13 1 3 3 3 3 1 11 1 3 1+ 2 3 3 1 23?21 1 3 2 3 3 1 13? 1+ 2 3 2 + 3 3 3 1 13710 2.3 3 3,4 3 4 1 23 1 3 3 3 3 1 21? 1 3 1 + 2 3 3 1 23 1 3 2 + 3 3 3 2 18 1 3 1+ 2 3 3 1 21 1 3 3 3 3 1 11 1+ 2 3 2 + 3 3 4 1 13 1 3 3 3 3 I 25 1 + 2 3 2 + 3 3 4 1 13 1 3 2 + 3 3 3 1 25 1 3 1 + 2 + 3 3 3 1 23 1+ 3 3 3 3 4 1 23 1+2 + 3 3 2 + 3 3 4 2 11 1 + 2 3 4 3 3 + 4 1 17 1+ 2 3 4 3 4 1 11 1 3 1 3 1 + 2 1 26 1+ 2 + 3 3 4 3 4 1 10? 2.3 3 3,4 3 4 1 21 1 3 3 3 3 1 17 1 3 2 3 2 1 13 1+ 2 3 3 3 4 1 23 1 3 1+ 2 3 3 1 11? 1 3 3 3 4 1 21? 1 3 3 3 3 1 9? T a b l e II. Continued

Taxon/Character 27 28 29 30 31 32 33

Plesio. Africa 1 1 3 1 I 1 3 adolphifrederici 1+ 6 1 2 1 1+ 2 1 3 alluaudi 1+6 1 3? 1? 1 1 3 altigenis 1+ 6 1 2 1 1 1 2 astatodon 1+ 6 1 3 1 1 1 3 bakongo 1+ 6 1 2 1 1 1 3 bicolor 4 + 6 1 3 1 1 1 2 bloyeti 6 1 3 2 1 1 2 + 3 brownae 1+ 6 1 3 1 1 1 2 burtoni 6 1 3 2 1 2 buy si 1+6 1 1 1 1 1 1 calliptera 1,6 1 3 2 1 1 2 + 3 cassius 1+ 6 1 2 1 1 1 2 cinerea 1+ 6 1 3 1 1 1 3 cnester 1+6+4 1 3 1 1 1 5 codringtoni 1+ 6 1 1 1 1 1 1 cronus 6 1 3 2 1 1 2 degeni 4 + 6 1 3 1 1 1 2? 5 desfontainesii 1 + 6 1 3 2 1 1 2 + 3 eduardiana 1 + 6 1 3 2 1 1 2 elegans 1+ 6 1 2 1 1 1 3 empodisma 1+ 6 1 2 1 1 1 2 erythrocephaius 1+ 6 2 3 1 1 I 4? flaviijosephi 1+ 6 1 3 1+ 2 1+ 2 1 2 + 3 fusiformis 1+ 6 1 2 1 1 2 gracilior 1+ 6 1 3 1 1 I 3?5 graueri 1+ 6 1 2 1 1 1 2 hiatus 1 + 6 + 4 1 2 1 1 1 5 horei 6 1 2 2 1 1 2 iris 1 + 6 + 4 1 2 1 1 1 3 ishmaeli 4 + 6 1 2 1 1 1 5 lacrimosa 1+ 6 1 2 1 1 1 2 mahagiensis 1+ 6 1 2 1 1 1 3 34 35 36 37 38 39

1 2 3 3 3 3 2 2 2 3 3 3 ? 2 + 3 1+ 2 ?? 3 2 2 2 3 3 3 2 + 3 2 2 3 1+ 2 3 2 3 1,2 3 2 3 1 2 2 + 3 3 3 3 1+ 2 3 2 3 2 3 1 2 3 3 3 3 2 3 2 3 3 3 1 2 2 3 3 3

1 2 2 3 2 3 H C S IT P IP L . 1,2 2 2,3 3 3 3 2 2 2 3 2 3 2 2 2,3 3 3 3 2 3 2 3 1+ 2 3 2,3? 2 2 3 2 3 2 2 3 3 2 3 1+2 2 + 3 2 3 1+2 3 2 2 3 + 3 3 2 3 2 + 3 2 2 3 2 + 4 3 2 2 3 3 2 3 1 2 3 3 3 3 1 2 2 3 3 3 2 2 3 3 3 3 2 2 1+ 2 3 2 3 1+ 3 2 + 3 2 3 3 3 1 2 1+ 2 + 3 3 1+ 2 3 2 3 1+ 2 3 3 3 2 2 2 3 3 3 2 2 2 3 3 3 1 2 2 3 2 3 2 2 + 3 2 3 3 + 4 3 malagaraziensis 1+ 6 1 2 1 1 I maxillaris 1+ 6 1 2 1 1 1 microdon 1+ 6 1 2 1 1 1 multicolor 1+ 6 1 2 1 1 nigricans 6 1 3 1 1 nigripinnis 1+ 6 2 2 + 3 1 I 1 nubila 6 1 3 2 1 1 niichisquamulatus 1+ 6 1 3 2 1 1 obesus 1+ 6 I 2 1 1 1 obliquidens 1+ 6 1 2 1 1 I obtusidens 1+ 6 1 2 1 1 I occult idens 1+ 6 1 2 1 1 1 orthostoma 1+ 6 1 2 1 1 1 parorthostoma 6 1 3 2 1 1 parvidens 1+ 6 1 2 1 1 1 paucidens 1+ 6 1 2 1 1 1 pectoralis 1 + 6 1 2 1 1 1 polli 6 1 2 1 1 1 prognathus 1 + 6 1 2 1 1 1 ptistes 1+4+6 1 3 I 1 1 re tr odens 4 + 6 1 3 1 1 1+3 riponianus 1+6 1 2 1 1 1 sauvagei 4 + 6 I 3 1 1 1 + 3 saxicola 1+6 1 2 1 1 1 serranus 1+6 1 3 1 1 1 spekii 1+6 1 3 1 I 1 squamipinnis 1+6 I 2 + 3 1 1 1 straeleni 1+6 1 3 1 1 1 teegelaari 1+4+6 1 3 1 1 1 thumbergi 1 1 1? 1 2 (orrenticola 6 1 3 2 1 tridens 1+6 1 2 1 1 1 vanderhorsti 1+6 1 2 1 1 1 victoriae 6 1 3 1 1 1 victorianus 1+6 1 3 1 1 1 vittatus 1+6 1 2 1 1 1 welcommei 1+6 2 2 1 1 1 wingatii 1+6 1 3 1 2 1 3 1 2 2 3 3 3 4 2 2 2 3 3 3 3 + 5 1 2 3 3 3 3 3 3 2 2 3 3 3 2 1 2 2 3 2 3 3 2 2 3 3 2 3 2 2 + 3 2 2 3 1+2 3 2 1 2 3 3 2 3

4 2 2 3 3 3 FISHES 3 INE HAPLOCHROM OF STUDY PHYLETIC A 2 + 3 1 2 2 3 2 + 3 3 2 1 2 3 3 3 3 2 1 2 2 3 1+2 3 2 2 2 2 3 1+2 3 2 2 2 3 3 2+3 3 2 1 2 + 3 2 3 3 3 2 + 3? 1 2 2 3 1+2 3 2+3 + 4 2 2 2 3 1 3 2 1? 3 2 3 3 3 2 + 4 2 2 2 3 2 3 3 1 2 3 3 3 3 5 2 2 1+2 + 3 3 3 3 2 1 2 3 3 3 3 5 1 2 2 3 3 3 2 2 2 3 3 3 3 2 + 4 1 2 3 3 3 3 2 + 4 2 2 3 3 4 3 3? 2 2 2 3 3 3 3 1 2 + 3 1+2 3 1 + 3 3 5 1 2 3 3 3 3 2? 2 3 2 + 3 3 1 3 3 2 2 2+3 3 4 3 2 + 4.. 2 2 3 3 3 3 2 2 2 1 3 2 3 2 2 2 3 3 2 3 2 + 4 1 2 2 3 3 3 2 + 4 1 3 2 3 3 3 2 + 4 1 2 3 3 3 3 3 1 2 + 3 2 3 3 3 T a b l e II.

Taxon/Character 40 41 42 43 44

Plesio. Africa 3 1 3 3 1 adolphifrederici 3 1 3 2? 2 alluaudi 4 ? 3 2 2 altigenis 3 1+ 3 3 I 2 astatodon 3 1 3 1 2 bakongo 4 1 3 I 2 bicolor 3 1+ 3 3 1 2 bloyeti 3 1 3 1 2 brownae 3 1 3 2 + 3 2 burtoni 4 1 3 1 2 buysi 1 1 3 1 2 calliptera 3 1 3 1 2 cassius 4 1 3 3 2 cinerea 3 1 3 1 2 cnester 4 1 3 2 2 codringtoni 1 2 3 1 2 cronus 2 + 3 4 3 1 2 degeni 3 1 3 1 2 desfonlainesii 3 1 3 1 2 eduardiana 3 1 3 2 2 elegans 4 1 3 2? 2 empodisma 3 1+ 3 3 3 2 erythrocephalus 3 1 3 2 2 flaviijosephi 3 1 3 1 2 fusiformis 3 1+3 3 2 + 3 2 gracilior 3 1 3 2 2 graueri 3 1 3 2 2 hiatus 4 1 3 2 2 horei 3 1 2 1 2 iris 4 1 3 2 2 ishmaeli 4 1 3 2 2 lacrimosa 3 1 3 2 + 3 2 mahagiensis 4 1?2 2 1 2 Continued K>VO OO 45 46 47 48 49 50 51 52

1 3 1 1 112 1 1 3 1 2 11?? 1 3 1,2? 2 1 ? ? ? 1 3 1 2 1 ? ? ? 1 3 1 2 112 1 2 3 1 2 112 2 1 3 12 12 2 2 1 3 1 2 112 1 13 12 1 ? ? ? 1 3 12 112 2 2 3 1 2 112 2

13 12 112 1 H C S IT P IP L . 1 3 2 + 3 2 1 ? ? ? 13 12 1 ? ? ? 1 3 1+ 2 2 i i ? ? 1 3 1 2 112 1 13 12 1 ? ? ? 13 12 i i ? ? 13 12 112 1 1 3 2 2 1 ? ? ? 13 12 i i 2 i 1 3 3 2 l ? ? ? 13 2 2 l i ? ? 13 12 ii?? 1 3 2 + 3 2 l ? ? ? 1 3 1 + 2 2 i i 2? i 1 3 1 + 2 2 i ? ? ? 1 3 1 + 2 2 1 i 2 i 2 3 1 2 112 2 1 3 2 2 112 1 1 3 1 + 2 2 12?? 1 3 1,2 2 1 ? ? ? 2 2 13 2 ? ? ? malagaraziensis 3 1 2 1 maxillaris 3 1 3 2 + 3 microdon 4 1 3 4 multicolor 3 3 3 1 nigricans 4 I 3 1 nigripinnis 4 1 3 2 nubila 3 1 3 1 nuchisquamulatus 3 I 3 1 obesus 3 + 4 1 3 2,3 obliquidens 3 + 4 1 3 1 obtusidens 3 1 3 2 occultidens 3 I 3 1? orthostoma 3 ? 3 1 parorlhostoma 3 7 3 1 parvidens 4 i 3 2 paucidens 4 i 3 1 pectoralis 3 4 3 2 polli 3 1 2 prognathus 2 I 3 2 + 3 ptistes 4 1 3 3 re tr odens 4 1 3 I riponianus 3 1 3 1 sauvagei 4 1 3 1 saxicola 3 1,3 3 1+ 2 serranus 3 4 3 2 spekii 3 1 3 3 squamipinnis 4 1 3 2 straeleni 4 1 3 2? teegelaari 3 I 3 3 thumbergi 1+ 2 2 3 1 torrenticola 4 1 2 1 tridens 4 1 3 4 vanderhorsti 3 1 3 2 victoriae 3 1 3 1 victorianus 3 1 3 3 vittatus 3 ? 3 2? welcommei 4 1 3 2 wingatii 3 ? 3 1 1 3 2 7 ? 7 1 + 2+3 2 1 ? 7 7 3 2 1 7 7 ? 1 2 1 1 2 2 1 2 1 7 ? 7 1+2 2 1 2 2 1 1 2 1 I 7 7 1 2 1 ? ? 7 1+2 2 1 7 ? FISHES ? INE HAPLOCHROM OF STUDY PHYLETIC A 1 2 1 1 7 7 1+2 2 1 ? 7 7 1? 2 1 i 7 7 1 2 1 ? 7 7 1 2 1 7 7 7 2 2 1 ?? 7 1 2 1 1 7 7 1+2 2 1 ? 7 7 1 2 1 7 7 1 2 1 ? ? ? 2 + 3 2 1 7 7 ? 1 2 1 i 7 7 1,2 2 1 7 ? ? 1 2 1 7 7 ? I +2 2 1 ? ? ? 1+2 2 1 7 ? 7 2 + 3 2 1 I 2 7 2 2 1 1 2 1 1 2 1 1 2 1 1+2 2 1 7 7 7 1 2 1 1 2 1 1 2 1 2 2 3 2 1 7 7 7 1,2 2 1 1 2 2 1 2 1 ?? 7 3 2 1 7 ? ? 1? 2 1 7 7 7 1+2 2 1 7 7 7 1 2 1 ? 7 ? T a b l e II. Continued

Taxon/Character 53 54 55 56 57 58 59

Plesio. Africa 3 3 1 1 3 3 1 adolphifrederici 3 2 2 1 3 2 2 alluaudi 3 4 1 1 ? ? 7 altigenis 3 3 1 1 3 i i astatodon 3 2 1 1 3 1 i bakongo 3 1 2 1 3 1 2 bicolor 3 I 1 1 3 1 1 bloyeti 3 1 1 1 3 1 2 brownae 3 3 1 1 3 1 burtoni 3 1 1 1 3 1 2 buysi 3 3 1 1 3 1 1 call ip ter a 3 1,2 I I 3 1+2 2 cassius 3 3 1 1 3 3 1 cinerea 3 2 + 3 1 1 3 2 1 cnester 3 2 2 I 3 2 2 codringtoni 3 2 1 1 3 1 1 cronus 3 1+2 1 1 3 1 1 degeni 3 1+ 2 2 1 3 1+2 2 desfontainesii 3 1 2 1 3 1 2 eduardiana 3 2 1 1 3 2 1 elegans 3 2 1 1 3 2 1 empodisma 3 3 1 1 3 2 2 erythrocephalus 3 2 1 1 3 2 1 flaviijosephi 3 2 1 1 3 2 1 fusiformis 3 2 1 1 3 2 1 gracilior 3 2 1 1 3 2 1 graueri 3 2 2 I 3 2 2 hiatus 3 2 2 1 3 2 2 horei 3 2 1 1 3 2 1 iris 3 2 2 1 3 2 2 ishmaeli 3 2 2 1 3 2 2 lacrimosa 3 2 1 1 3 2 1 mahagiensis 3 2 1 1 3 2 1 930

60 61 62 63 64 65 . PPI H C S IT P IP L E. malagaraziensis 3 1 + 3 2 2 3 1 maxillaris 3 2 1 1 3 2 microdon 3 4 1 1 3 3 multicolor 3 1+ 2 2 1 3 1 + 2 nigricans 3 1,2 2 1 3 1,2 nigripinnis 3 2 2 1 3 2 nubila 3 1 2 1 3 1 nuchisquamulatus 3 1 2 1 3 1 obesus 3 2 1 1 3 1+ 2 obliquidem 3 2 + 3 1 1 3 1 obtusidens 3 2 1 1 3 2 occultidens 3 1 1 1 3 1 orthostoma 3 3 2 1 3 1 parorthostoma 3 3 2 I 3 2 parvidens 3 2 1 1 3 2 paucidens 3 1 1 1 3 1 pectoralis 3 2 1 1 3 2 polli 3 1 1 1 3 1 prognathus 3 2 1 1 3 2,3 ptistes 3 2 1+ 2 1 3 1+ 2 retrodens 3 1+ 2 1 1 3 2 riponianus 3 2 1 1 3 1 sauvagei 3 2 1 I 3 2 saxicola 3 1+ 2 1 1 3 1 serranus 3 2 1 1 3 1 + 2 spekii 3 3 1 1 3 2 squamipinnis 3 2 2 1 3 2 straeleni 3 2 1 1 3 2 teegelaari 3 2 1 1 3 2 thumbergi 3 1 1 1 3 1 torrenticola 3 1 2 2 1 tridens 3 3 I I 3 3 vanderhorsti 3 2 1 3 2 victoriae 3 2 2 1 3 1 victorianus 3 3 1 1 3 3 vittatus 3 1 2 1 3 1 welcommei 3,4 2 2 1 3 2 h 'ingatii 3 2 2 1 3 2 A A PHYLETIC STUDY OF HAPLOCHROMINE FISHES T a b l e II.

Taxon/Character 66 67 68 69 70

Plesio. Africa 3 3 adolphifrederici 1 3? alluaudi 3 3? altigenis 1 4 astatodon 2 2-1-3? bakongo 1 2 bicolor 1 3 bloyeti 1 2 + 3 brownae 1 3 burtoni 1 2 + 3 1 2 + 3 calliptera 1 2 + 3 cassius 2 + 3 3? cinerea 1 3 cnester 2 4? codringtoni \ 1+5 cronus 1 3 + 5 degeni 1 3 desfontainesii 1 2 + 3 eduardiana 2 4 elegans 1+ 2 2 + 3 empodisma 1 3 + 4 erythrocephalus 1 4? flaviijosephi 1 2 + 3 fusiformis 1,2 3 gracilior 1 4 graueri 1 3? hiatus 2 4? horei 1 3 /ra 2 4? ishmaeli 1+ 2 4? lacrimosa 1 3 mahagiensis 1 2 Continued

71 72 73 74 75 76 77 78

1 —

1 — 2.4 1 2 —

1 — 1 _

1 — 1.4 —

1 — 1 _ _

1 —

1,2 —

1 —

1 — 2,3 1

1 —

2 —

1,2 — 1 —

2 — malagaraziensis 2+ 1 2 + 3 1 — —- 1 maxi/laris 1 4 1 — - 1 microdon 1+ 2 + 3 4 1 —— 1 multicolor 1 2 1 —— 1 nigricans 1 3 1 — — 1 nigripinnis 1 3? 1 — — 1 nubila 1 2 + 3 1 — — 1 nuchisquamulatus 1 3 1 —— I obesus 1 4 1 —— 1 obliquidens I 2 + 3 1 —— 1 obtusidens 1 4? 1 —— 1 occultidens ? ? 1 —— 1 orthostoma 1 4? 1 — __ I parorthostoma 1 4? 1 — — 1 parvidens 2 4? 1 —— 1 paucidens 1 4? 1 — — 1,2 pectoralis 1 4 1 — — 1 polli 1 2 1 —— 1 prognathus 1 3 + 4 1 — — 1 ptistes 1 4 I — — 1 re tr odens 1 3 1 —— 1 riponianus 1 3 + 4 1 — — 2,3 sauvagei 1+ 2 3? 1 — — 1 saxicola 1 3 + 4 1 —— 2,3 serranus I 3 + 4 1 —— 1 spekii 1 + 2 + 3 4 1 — — 1 squamipinnis 1 3? 2 1 3 1 Straeleni 1,3 2 + 3 1 — — 1 teegelaari 1+ 2 + 3 3 + 4 1 —— 1 thumbergi 1 4 1 —— 1 torrenticola 1+ 3 3 1 tridens 3 4 + 2? I — — 1 vanderhorsti 3 3 1 — — 1 victoriae 1 3 + 4? 1 —— I victorianus 3 3 + 4 1 —— 1,4 vittatus 1 4? 1 —— 1 welcommei 1+ 2 + 3 3 + 4 1 —— 1 wingatii 1 3 1 _— — 1 PYEI SUY F ALCRMIE FISHES INE HAPLOCHROM OF STUDY PHYLETIC A

LkJ T a b l e II.

Taxon/Character 79 80 81 82 83

Plesio. Africa 2 1115 adolphifrederici 2 1115 alluaudi 2 1115 altigenis 2 1115 astatodon 2 1115 bakongo 2 1115 bicolor 2 1115 bloyeli 2 1115 brownae 2 1,2? 1 I 5 burtoni 2 1115 buysi 2 1115 calliptera 2 1,2? 1 1 5 cassius 2 1115 cinerea 2 1115 cnester 2 1115 codringtoni 2 1115 cronus 2 1115 degeni 2 1115 desfontainesii 2 1115 eduardiana 2 1 I I 5 elegans 2 1115 empodisma 2 1115 erythrocephalus 2 1115 Jlaviijosephi 2 1115 fusiformis 2 1115 gracilior 2 1115 graueri 2 1115 hiatus 2 1115 horei 2 1115 iris 2 1115 ishmaeli 2 1115 lacrimosa 2 1115 mahagiensis 2 1115 Continued LI TSCH C S IT P IP L . malagaraziensis 2 1 I 1 5 maxillaris 2 1 11 5 microdon 2 1 11 5 multicolor 22? 22 5 nigricans 21,2 11 5 nigripinnis 2 1 I 1 5 nubila 2 1 11 5 nuchisquamulatus 2 1 11 5

obesus 2 1 11 5 FISHES INE HAPLOCHROM OF STUDY PHYLETIC A obliquidens 21,2 11 5 obtusidens 2 I 11 5 occultidens 2 1 11 ? orthostoma 2 1 11 5 parorthostoma 2 1 11 5 parvidens 2 1 11 5 paucidens 2 1 I 1 5 pectoralis 2 I 11 5 polli 2 1 1I 5 prognathus 2 1 I 1 5 ptistes 2 1 11 5 retrodens 2 1 11 5 riponianus 2 1 11 5 sauvagei 2 1 11 5 saxicola 2 1 1I 5 serranus 2 1 11 5 spekii 2 1 1 1 5 squamipinnis 2 1 11 5 straeleni 2 1 11 5 teegelaari 2 I 11 5 thumbergi 2 1 1 I 5 torrenticola 2 1 11 5 tridens 2 1 11 5 vanderhorsti 2 1 11 5 victoriae 2 1 1 1 5 victorianus 2 1 11 5 vittatus 2 1 11 5 welcommei 2 1 I I 5 wingatii 2 1 11 5

-f More than one state usually on same specimen., Various stales on different specimens. ? State uncertain. Not applicable. 936 E. LIPPITSCH the chest scales) is common to all investigated taxa. Most characters have their respective states distributed over several taxa. In some cases because of the complexity of the characters homoplastic evolution seems unlikely and the similarities have to be regarded as synapomorphies. This is above all the case with the various granulation types. Others seem to have evolved several times independently. In nearly no character was it possible to establish the simple condition of two states with equal weight. Neither was it possible to construct what was called a ‘ character state tree ’ by Cichocki (1976) for any of the characters. It could not even be inferred that transitional states have evolved in the way suggested by gradual variation. For example, two apomorphic states a { and a2 may have evolved from the plesiomorphic state p according to one of the following schemes: (1 ) p - > a ^ a 2; (2)p-+a],p^>a2, (3)p^>a2^>ax. Thus relationships between two taxa possessing a, and a2 as apomorphic similarities remain unclear, unless other synapomorphies can be established. Such problems make a phyletic analysis difficult. As shown below, however, the investigated characters give the possibility to reassess, from a different approach, many of Greenwood’s supposed lineages and to interrelate them in a phylogenetic way.

IV. PHYLETIC ANALYSIS As already noted, homoplastic evolution of a given granulation type seems unlikely. Thus we have to assume seven monophyletic lineages according to the seven granulation types encountered (regarding all of them apomorphic, as explained above). It then has to be asked whether the lineages so defined can be corroborated by further synapomorphies. The numbers of character states are given in parentheses in the form (N}n), N being the number of the respective character and n that of the character state. Most straightforward is Astatoreochromis. Granulation type 17 is restricted to this genus, and there are additional autapomorphies. All three species assigned to the genus by Greenwood (1979) on the basis of their increased number of dorsal and anal fin spines, overall similarities in coloration and the pattern and number of anal ocelli, lack of sexual colour dimorphism, and enlargement of the lower pharyngeal bone, uniquely among haplochromines share the following scale and squamation apomorphies. The opercular blotch is large, but not heavily pigmented (3/2); in the caudalmost part of the interoperculum the scale row is extended to form a vertical column of at least three scales; the preoperculum bears scales, especially on its vertical limb (8/2); the first circuli in the interradial space on the rostral part of the scale are concave (36/1) and the interradial tongues are long (35/3); the scales on the ventral surface of the chest are small (48/2); and the scales on the anal-genital region bear only vestigial ctenii (58/2). A. alluaudi has the autapomorphic state of having some ctenoid scales on the operculum. In all other characters Astatoreochromis is plesiomorphic, or shares the respective apomorphic characters with most other taxa. Hence Astatoreochromis evidently diverged from the rest of the haplochromines rather early, as already suggested on the basis of biochemical data (Sage et a i, 1984; Verheyen et al., 1989; Meyer et al., 1990). Granulation type 10 is limited to fluviatile species of Astatotilapia and is present in A. flaviijosephi from Israel, A. desfontainesii from Tunisia, A. bloyeti from eastward-flowing rivers in Tanzania, A . burtoni from the L. Tanganyika drainage, A PHYLETIC STUDY OF HAPLOCHROMINE FISHES 937 and A . calliptera from L. Malawi. Another autapomorphy is the coverage of the scale focus by the granulation (30/2). There are apomorphic similarities with other taxa with which fluviatile Astatotilapia share small occipital scales (17/2), cycloid scales on the rostral part of the dorsum (20/1, 2), short and bluntly conical ctenii (33/2), straight first interradial circuli (36/2), cycloid scales on the belly (54/1, 2), and cycloid scales on the caudal fin (66/1). The apomorphy of strictly cycloid scales on the chest (43/1) is shared with a group of interrelated lacustrine taxa (the Haplochromis superlineage defined below). However, in view of the other differences, this feature must be considered homoplastic. Early divergence of Astatotilapia s. str. is supported by data from DNA studies (Meyer et al., 1991 and pers. comm.). In particular, it is unlikely that a species from this lineage with its highly peculiar granulation should be the ancestor of the whole flock of lacustrine haplochromines, as was previously suggested. The striking differences between fluviatile and lacustrine Astatotilapia (see below) argue strongly even against the phyletic integrity of the genus as conceived by Greenwood. The fluviatile Astatotilapia obviously form a monophyletic lineage, with the lacustrine species supposed so far to belong to the genus being only distantly related. Also this conclusion is backed by the DNA studies (Meyer eta L , 1990, 1991). Granulation type 13 unites genera Haplochromis s. str., Xystichromis, Neochromis, Ptyochromis, Hoplotilapia, Macropleurodus, and Platytaeniodus. This assemblage in addition shares the following apomorphies: Cycloid scales on the chest (43/1) and belly (54/1) and straight first interradial circuli (36/2). Most taxa have also cycloid scales on the caudal part of the dorsum (22/1) and weakly ctenoid ones in the anal-genital region (58/2). The group will henceforth be referred to as the Haplochromis superlineage and is assumed to be monophyletic. Apomorphies shared with other groups are small occipital scales (17/2), cycloid scales on the rostral part of the dorsum (20/1,2 and 22/1), cycloid or only vestigially ctenoid scales on the belly (54/1, 2), and cycloid scales on the caudal fin (66/1). Within that superlineage subgroups can be established. A first dichotomy is between Haplochromis, Xystichromis and Neochromis on the one hand (called the Haplochromis lineage) and Ptyochromis and the monotypic genera on the other (called the Hoplotilapia lineage). The first lineage is characterized by the apomorphy of having, caudally of the pectoral fin bases, scales which are ctenoid, but with only minute ctenii (38/2). On the belly cycloid scales are present as in the other members of the whole assemblage. The second subassemblage shares the following apomorphies: a small scaleless blotch on the operculum (3/3), circuli turning outward to meet the caudal rim of the scale nearly under a right angle (27/4), an only weakly ossified ctenoid rim (32/3), scales on the caudal peduncle which are heavily ctenoid, even more so than on the flank (40/4). The Hoplotilapia lineage can be split further. Hoplotilapia and Ptyochromis have weakly ctenoid scales on the caudal part of the dorsum (22/3), which is regarded as a reversal to the plesiomorphic state, and a peculiar form of ctenii (33/5). Granulation type 11 is found in the lacustrine ‘ Astatotilapia ’ and the genera Gaurochromis and Enterochromis. These taxa share no further apomorphies except the rather small occipital scales (17/2), cycloid scales on the rostral part of the dorsum (20/1, 2), scales with vestigial ctenii on the belly (54/1, 2), and cycloid scales on the caudal fin (66/1), which are all widespread features and are also found in the lineages characterized by granulation types 21,23, and 25. All three share the 938 E. LIPPITSCH apomorphic character of a medium-sized granular area (29/2). This granulation type 11 seems not to establish a monophyletic lineage. Most probably types 11,21, 23 and 25 are members of a superlineage characterized by strongly ctenoid scales on the chest and belly (41/3, 47/3, 54/3). Within that superlineage, the lacustrine Astatotilapia seem to have no synapomorphies. They obviously form a rather plesiomorphic group, but their monophyly cannot yet be sustantiated. Type 11 represents the plesiomorphic granulation of the superlineage and has been retained in a paraphyletic assemblage of taxa. Granulation type 21 contains both subgenera of Prognathochromis. Other shared apomorphies are medium size of the granular area (29/2), ctenoid scales on the chest (47/3 or 47/4), ctenoid scales on the belly (54/3) and distinctly ctenoid scales on the anal-genital region (58/3) and the caudal fin (66/3). Granulation type 25 characterizes only the genus Psammochromis, and not even all species included by Greenwood. But there is another synapomorphy which has a distribution congruent with granulation type: the anal fin has a distinct pad which usually bears cycloid scales (71/3, 72/1). So it has to be concluded that Psammochromis is a monophyletic entity, but not all species assigned to that genus by Greenwood (1980) are in fact members of that entity. This is discussed below. The size of the granular area is medium (29/2) as in Prognathochromis. Granulation type 23 comprises Harpagochromis, Schubotzia, Lipochromis, and Yssichromis. These genera have in common the apomorphies of ctenoid scales on the suboperculum (5/2). They share this feature with Gaurochromis and Enterochromis. Harpagochromis is characterized by the autapomorphic features of double post-orbital columns (12/3). The species composition of this genus is discussed below. This superlineage shares a number of other apomorphies with other groups. The above review of scale apomorphies shows that most groups characterized by a single granulation type have additional apomorphies in common. It seems justified therefore to regard them as monophyletic lineages. In addition, for many of them phyletic relationship had been suspected even earlier. Especially Greenwood’s paper ofl974 gives a tentative phyletic tree (not based on a cladistic analysis), which anticipates many of the relations postulated here, most species being named Haplochromis at that time. For example, Macropleurodus and Ptyochromis were shown as related (Hoplotilapia and Platytaeniodus were not included in the tree), and the same hypothesis was elaborated later (Greenwood, 1980). It is corroborated by the findings of this study. However, there are no reasons to include Psammochromis in this lineage, since it shares none of the apomorphies except the presence of cycloid scales on the caudal fin (66/1). Since this feature is present in most other taxa and already in the fluviatile Astatotilapia, it is considered to be plesiomorphic for the Victoria-Edward-Kivu haplochromines. Paralabidochromis, which Greenwood (1980) placed in the same lineage, cannot yet be unambiguously assigned. From the two species examined, the type species, Pa. victoriae, is known only from one specimen. Only a single scale could be studied. From this it may be concluded that the species belongs to the Hoplotilapia subassemblage. The second species, Pa. paucidens from L. Kivu, must be excluded since its granulation resembles (but not fully matches) type 21. So Paralabidochromis has to be regarded as of (at least) diphyletic origin. A PHYLETIC STUDY OF HAPLOCHROMINE FISHES 939

A relationship between the taxa of the Haplochromis assemblage was also proposed by Greenwood (1974, 1980). While affinities between Haplochromis, Xystichromis and Neochromis are corroborated by this study, there is strong evidence that Enterochromis is not closely related to the assemblage. In this case the similarities of the tropic apparatus invoked by Greenwood must have evolved independently. Another relationship suspected earlier (Greenwood, 1974) is that between Gaurochromis and Enterochromis erythrocephalus. However, the other species later assigned to Enterochromis had been associated with the Haplochromis assemblage in that earlier paper. The scale characters make such a relationship unlikely but suggested that the genus is close to Gaurochromis. Greenwood (1980) suspected Labrochromis and Gaurochromis not to be closely related. This was challenged by Hoogerhoud (1984). The scale and squamation characters show no differences between Gaurochromis and two species of Labrochromis, namely L. ptistes and L. teegelari. Thus Hoogerhoud’s view of a single morphocline seems to be corroborated. There is one problem, however. The specimens examined of the type species of Labrochromis, L. ishmaeli, show a granulation that cannot be unambiguously assigned. If this difference could be substantiated in further studies one would have to conclude that the two genera are distinct, but some of the species would have to be reassigned. From the above analysis and the other data available (Greenwood, 1974,1979, 1980; Sage el al., 1984; Verheyen et al., 1984, 1989; Hoogerhoud, 1984; Eccles & Trewavas, 1989; Meyer et a l 1990, 1991) a phyletic picture emerges which is summarized in the cladogram of Fig. 11. The Malawi taxa have not been studied in sufficient breadth to allow conclusions, therefore they are omitted here. For the assemblage comprising Astatoreochromis, the fluviatile Astatotilapia, and the L. Victoria-Edward-Kivu flock a common ancestor can be assumed, but it may well be that other genera (fluviatile as well as lacustrine from other lakes) will have to be included. Within that assemblage, Astatoreochromis is the sister taxon to the others combined. The Victoria-Edward-Kivu flock seems to be a monophyletic group, with the fluviatile Astatotilapia as the sister taxon. Within the flock two superlineages can be distinguished, each comprising several of Greenwood’s genera. For many of these genera new apomorphies can be found among scale and squamation characters, thus corroborating his generic break-up of the old Haplochromis concept. In addition, the phyletic interrelations between the genera is rather well resolved. There remain a number of open questions. The first one is that of the species composition of Greenwood’s genera. It is beyond the scope of this work to reassess the generic placement of all the species involved. Only a few remarks can be made. As noted above, the genus Astatotilapia has to be restricted to the fluviatile species. For the lacustrine members of the genus no autapomorphy has yet been discovered, but nevertheless it seems probable that most of them are closely related to each other. Thus it seems advisable to establish a new genus for this group, which then will be the sister taxon to the other members of the superlineage combined. Not every species assigned to Harpagochromis by Greenwood shares the distinc tive autapomorphles of the genus. The most obvious case is Hp. squamipinnis, which shares neither the granulation type nor the characteristic post-orbital squamation. 940 E. LIPPITSCH

3/ 2; s / 2; 26/ i 7^ 36/ i — Astatoreochromis

Astatotilapia

------— — Haplochromis 38/2 55/2 Neochromis Xystichromis 44/2 26/13; 58/2 Hoplotilapia 48/2 22/3 33/5 -Ptyochromis 17/2 3/3; 27/4 ' Platytaeniodus

' Macropleurodus

26/11 — 'Astatotilapia’

26/21 ■ Prognathochromis

26/25; 71/3; 72/1 Psammochromis 43/2,3 47/2,3 54/2,3 67/4 Gaurochromis 26/11 28/2 Enterochromis 5/2 6/4 - Yssichromis -Lipochromis 26/23 -Schubotzia 6/2 12/3 - Harpagochromis

F ig. 11. Cladogram of the haplochromine taxa, based on scale and squamation characters. Autapo- morphies of the respective lineages are denoted by numbers according to Table I.

It is unique within the L. Victoria-Edward-Kivu flock in having scaled dorsal, anal, and pectoral fins. These features are otherwise found only in L. Malawi haplochromines. No other indications of a close relation to the L. Malawi taxa have been found, however, and the similarities in fin squamation may have evolved independently. This assumption is corroborated by the fact that ‘ Haplochromis ’ squamipinnis has a different fin squamation pattern (double rows along the rays) from that usual in L. Malawi (single or multiple rows on the membranes). Certainly, the species has to be excluded from Harpagochromis, but no further A PHYLETIC STUDY OF HAPLOCHROMINE FISHES 941 comments on its relationships can yet be made. Other species whose assignment to Harpagochromis is doubtful are H. artaxerxes, guiarti, maculipinna, michaeli, plagiostoma, and thuragnathus. These were not examined in this study, but from illustrations in the literature they seem to possess only single postorbital columns. One specimen of Harpagochromis pectoralis (BMNH 1966.3.9.295) had only a single post-orbital column and cycloid scales on the chest. It is not clear whether this is an aberrant specimen or that it should be assigned to another species (and genus). Other specimens had two or three post-orbital columns and rather deeply imbedded, but ctenoid, chest scales. The type of the species (which was not examined) has two columns, as is obvious from Boulenger’s (1915) illustration. Thus the species is a Harpagochromis. Greenwood (1980) divided Lipochromis into two subgenera. The type species of both , L. (L.) obesus and L. (Cleptochromis) parvidens, were examined. No significant differences in scale and squamation characters could be found. However, L. (Cleptochromis) microdon could not be unambiguously assigned to granulation type 23 as can other Lipochromis and showed a degree of ctenoidy on the chest, belly, caudal peduncle and caudal fin otherwise found only in the Prognathochromis assemblage. Whether this reflects a diphyletic origin of Lipochromis, an idea entertained earlier by Greenwood (1974), remains an open question. The characteristic apomorphies of the genus Psammochromis have only been found in P. riponianus and P. saxicola, but not in P. graueri and P. cassius. At present neither of these species can be related to one of the lineages defined above nor to each other. Close relationship to Psammochromis s. sir. is unlikely. For a few genera no clear relationships could be established. One is Allochromis, scale granulation resembles type 21 (Prognathochromis), but no other synapo morphies could be detected. The second one is Pyxichromis. The two species of this genus resemble each other very closely in their unique jaw morphology, but, as far as can be concluded from the single specimen studied in each species, they differ in scale characters. The very few specimens available preclude any definite conclusions. The intuitive overall impression gained from scale and squamation characters is that the haplochromines in a broad sense are of common origin. They all have ctenoid scales, but this is a plesiomorphic feature. The circuli in the caudal field of the flank scales are mostly obscured. There is a definite trend to small scales on the upper part of the head and on the chest, accompanied by a reduction in ctenoidy in those regions. This trend distinguishes the haplochromines from many other cichlids with ctenoid scales, especially the New World taxa. Unfortunately, however, a similar trend is found in unrelated taxa, for example even in Heterochromis multidens, a species which is probably near the phyletic base of the African as well as the American assemblage (Stiassny, 1991), or in Paratilapia polleni, which possibly is an even more basic taxon. Thus the similarities among the haplochromines cannot be unambiguously regarded as synapomorphies and hence their monophyly cannot be established on the basis of scale and squamation characters. The same is true for the L. Victoria-Edward-Kivu flock. Nevertheless, none of the characters studied so far contradicts a monophyletic origin, for the flock as well as for the haplochromines. In view of the data gained from tRNA studies (Meyer et aL, 1990) a common ancestry has to be assumed, the 942 E. LIPPITSCH splitting being of rather recent age, even though the time estimates presented in that paper may be open to revisions. The relationship of fluviatile genera to the lacustrine haplochromines is not yet clear and is under investigation, but a few preliminary statements can be made. Serranochromis, Sargochromis, Chetia, and Pharyngochromis seem very far removed phylogenetically from the L. Victoria-Edward-Kivu flock. Pseudocrenilabrus, Ctenochromis, Thoracochromis, and Orthochromis seem to be nearer to the flock, but none of these genera may have close relationships to one of the superlineages defined above. So it may well be that the two superlineages within the L. Victoria-Edward-Kivu flock are more closely related to one another than to any fluviatile taxon and have originated from a single ancestral species. None of the extant fluviatile haplochromines examined so far represent that ancestral type, however, since even the most closely related fluviatile Astatotilapia have developed a considerable number of autapomorphic features including their distinctive granulation type.

V. CONCLUSION This study of scale and squamation characters of a large number of haplochromine species covering all the genera of the L. Victoria-Edward-Kivu flock has been successful not only in corroborating the validity of previously proposed subdivisions of the flock but also interrelating those subdivisions phyletically. The bulk of the information now available from anatomical and biochemical studies gives the strong impression that the haplochromines, including Astatoreochromis, some fluviatile genera, and the L. Victoria-Edward-Kivu flock are of monophyletic origin. This impression is backed by scale and squamation studies, but so far no synapomorphy has been detected. The same is true for the L. Victoria-Edward-Kiva flock itself, within which two super lineages can be distinguished, each comprising several genera. While the superlineages extend over more than one lake, much of the speciation seems to have taken place in the respective lakes. Many of Greenwood’s genera can be characterized using scale characters, and new aspects of their mutual interrelationships have been established. The tree worked out in this investigation strengthens the evidence that most of those genera (with some species rearrangements) are phyletic entities and that the splitting of the former genus Haplochromis is justified, notwithstanding the close relations between the genera. Thus Greenwood’s phyletic classification (1979, 1980) of the haplochromine taxa is essentially confirmed, and the availability of scale and squamation characters should make the practical use of this classification easier. Finally, scale and squamation characters have proved valuable in unravelling phylogenetic puzzles within the Cichlidae, even in such a difficult group as the East African haplochromines.

The author is indebted to the following persons and institutions for supplying specimens: Dr B. Herzig (Naturhistorisches Museum, Wien), Dr M. J. P. Van Oijen (Rijks museum van Natuurlijke Historie, Leiden), Prof. D. F. E. van den Audenaerde and Dr Guy Teugels (Museum Tervuren), Mr G. Howes (British Museum Natural History), and Mr St. Mascha (Graz). I wish to thank Dr P. H. Greenwood for valuable discussions, permanent encouragement, and careful reading of the manuscript. The help of Mr Ch. Ellis of the Zentrum fur Elektronenmikroskopie, Graz, in preparing the electron micrographs is A PHYLETIC STUDY OF HAPLOCHROMINE FISHES 943 highly appreciated. Financial support by the Osterreichischen Fonds zur Forderung der wissenschaftlichen Forschung, grant no. P7800-BIC), is gratefully acknowledged.

References

Boulenger, G. A. (1915). Catalogue of the Fresh-water Fishes of Africa in the British Museum (Nat Hist), vol. 3. London: British Museum. Cichocki, F. P. (1976). Cladistic history of cichlid fishes and reproductive strategies of the American genera Acarichthys, Biotodoma and Geophagus. Ph.D. thesis, University of Michigan, Ann Arbor. De Vos, L., Snoeks, J. & Thys van den Audenaerde, D. F. E. (1990). Description d'Haplochromis erythromaculatus, espece nouvelle (Teleostei: Cichlidae) des lacs Bulera et Ruhondo, Rwanda. Ichthyological Exploration o f Freshwaters 1,257-268. Eccles, D. H. & Trewavas, E. (1989). Malawian Cichlid Fishes. The Classification o f Some Haplochromine Genera. Herten: Lake Fish Movies. Greenwood, P. H. (1974). The cichlid fishes of Lake Victoria, East Africa: The biology and evolution of a species flock. Bulletin of the British Museum (Natural History) Zoology suppi 6, 1-134. Greenwood, P. H. (1979). Towards a phyletic classification of the ‘ genus ’ Haplochromis (Pisces: Cichlidae) and related taxa Pt. I. Bulletin of the British Museum (Natural History) Zoology 35, 265-322. Greenwood, P. H. (1980). Towards a phyletic classification of the 4 genus ’ Haplochromis (Pisces: Cichlidae) and related taxa Pt. II: the species from Lakes Victoria, Nabugabo, Edward, George and Kivu. Bulletin o f the British Museum (Natural History) Zoology 39, 1 — 101. Greenwood, P. H. (1984). The haplochromine species (Teleostei, Cichlidae) of the Cunene and certain other Angolan rivers. Bulletin o f the British Museum (Natural History) Zoology Al, 187-239. Hilgendorf, F. (1988). Fische aus dem Victoria-Nyanza (Ukerewe-See). Sitzungsbericht der Gesellschaft naturforschender Freunde Berlin 1888, 75-79. Hoogerhoud, R. J. C. (1984). A taxonomic reconsideration of the Haplochromine genera Gaurochromis Greenwood, 1980 and Labrochromis Regan, 1920 (Pisces, Cichlidae). Netherlands Journal of Zoology 34, 539-565. Lippitsch, E. (1989). Scale surface morphology in African cichlids (Pisces, Perciformes). Annals du Musee Royal de VAfrique Central Sciences Zoologiques 157, 105-108. Lippitsch, E. (1990). Scale morphology and squamation patterns in cichlids (Teleostei, Perciformes): A comparative study. Journal o f Fish Biology 37, 265-291. Lippitsch, E. (1991). Comparative investigation on scale characters in cichlids. Annals du Musee royal de I’Afrique central Sciences zoologiques 262,97-102. Lippitsch, E. (1992). Squamation and scale character stability in cichlids, examined in Sarotherodon galilaeus (Linnaeus, 1758) (Perciformes, Cichlidae). Journal o f Fish Biology 41, 355-362. Meyer, A., Kocher,T. D., Basasibwaki, P. & Wilson, A. C. (1990). Monophyletic origin of Lake Victoria cichlid fishes suggested by mitochondrial DNA sequences. Nature 347, 550-553. Meyer, A., Kocher, T. D. & Wilson, A. C. (1991). African fishes (reply to Crapon De Caprona & Fritzsch). Nature 350,467-468. Poll, M. (1986). Classification des Cichlidae du lac Tanganika. Tribus, genres et especes. Academic Royale de Belgique Memoires de la Classe de Sciences Coll. in-8-2e serie, T.45-Fasc.2-1986. Regan, C. T. (1920). The classification of the fishes of the family Cichlidae. I. The Tanganyika genera. Annals and Magazine of Natural History Ser. 9,5, 33-53. Sage, R. D., Loiselle, P. V., Basasibwaki, P. & Wilson, A. C. (1984). Molecular versus morphological change among cichlid fishes (Pisces: Cichlidae) of Lake Victoria. In Evolution of Fish Species Flocks (Echelle, A. A. & Kornfield, I., eds). University of Maine Orono Press. 944 E. LIPPITSCH

Sire, J.-Y. & Arnulf, I. (1989). Squamation chronology and scale development in cichlids. Annals du Musee Royal de I’Afrique Central Sciences Zoologiques 157,97-100. Snoeks, J., De Vos, L., Coenen, E. & Thys van den Audenaerde, D. F. E. (1990). Description de deux nouvelles especes d’Haplochromis (Teleostei, Cichlidae) du Lac Kivu, Rwanda. Cybium 14,63-76. Stiassny, M. L. J. (1981). The phyletic status of the family Cichlidae (Pisces: Perciformes): A comparative anatomical investigation. Netherlands Journal of Zoology 31, 275-314. Stiassny, M. L. J. (1990). Tylochromis, relationships and the phylogenetic status of the African Cichlidae. American Museum Novitates 2993,1-14. Stiassny, M. L. J. (1991). Phylogenetic intrarelationships of the family Cichlidae: an overview. In Cichlid Fishes. Behaviour, Ecology and Evolution (Keenleyside, M. H. A., ed). London: Chapman & Hall. Verheyen, E., Van Rompaey, J. & Selens, M. (1985). Enzyme variation in haplochromine cichlid fishes from Lake Victoria. Netherlands Journal of Zoology 35, 469—478. Verheyen, E., Van der Linden, A. & Decleir, W. (1989). The eye lens proteins of haplochromine cichlids from Lake Victoria studied by isoelectric focusing. Annals du Musee royal de VAfrique central Sciences zoologiques 157, 93-96.

Appendix The following list gives the relevant specimens used in this study. They were borrowed from the British Museum (Natural History) London (BMNH), Rijksmuseum van Naturlijke Historie Leiden (RMNH), Koninklijke Museum voor Midden-Africa Tervuren (MT), and Naturhistorisches Museum Vienna (NMW). Provenance of the species is indicated by a code letter following the registration number (V-L. Victoria-Edward-Kivu, T-L. Tanganyika, M-L. Malawi, F-fluviatile)

Allochromis weicommei (Greenwood, 1965) BMNH 1965.9.6.2-4 V Astatoreochromis alluaudi (Pellegrin, 1904) NMW 18821 V,F Astatoreochromis straeleni (Poll, 1944) MT 73-68-P-171 T,F Astatoreochromis vanderhorsti (Greenwood, BMNH 1953.11.4.59-62 T,F 1958) Astatotilapia bloyeti (Sauvage, 1883) MT 191162-201 F Astatotilapia burtoni (Gunther, 1893) NMW 89977:1-3 T,F Astatotilapia brownae (Greenwood, 1962) BMNH 1962.3.2.76-88 V Astatotilapia calliptera (Gunther, 1893) BMNH 1966.7.26.17-42 M,F Astatotilapia desfontainesii (De Lacepede, 1803) MT 73-42-P-1-675 F Astatotilapia elegans (Trewavas, 1933) RMNHV Astatotilapia flaviijosephi (Lortet, 1883) BMNH 1949.9.16.390-396 F Astatotilapia lacrimosa (Boulenger, 1906) BMNH 1959.4.28.1-23 v Astatotilapia nubila (Boulenger, 1906) NMW 89978 V,F Aulonocara hansbaenschi Meyer, Riehl & NMW 89979 M Zetzsche, 1987 Ctenochromis horei (Gunther, 1893) NMW 24660 T Ctenochromis polli (Thys, 1964) MT 143087-098 T Cyphotilapia frontosa (Boulenger, 1906) NMW 89983 T Cyrtocara moorii Boulenger, 1902 NMW 90982 M Dimidiochromis compressiceps (Boulenger, NMW M 1908) Enterochromis erythrocephalus (Greenwood & BMNH 1968.8.30.270-291 V Gee, 1969) Enterochromis nigripinnis (Regan, 1921) RM NH Coll. IBP V Gaurochromis empodisma (Greenwood, 1960) BMNH 1959.4.28.171-179 V Gaurochromis hiatus (Hoogerhoud & Witte, RM NH 31-5-1984 V 1981) A PHYLETIC STUDY OF HAPLOCHROMINE FISHES 945

Appendix Continued.

Gaurochromis iris (Hoogerhoud & Witte, MNH 4-1-1978 V 1981) Gaurochromis obtusidens (Trewavas, 1928) BMNH 1959.4.28.213-218 V Gephyrochromis moorii Boulenger, 1901 NMW 89995 M Haplochromis astatodon Regan, 1921 MT 80-29-P-2031 V Haplochromis obiiquidens Hilgendorf, 1888 BMNH 1956.7.46-55 V ' Haplochromis ’ adolphifredrici (Boulenger, MT 81-55-P-1994-2029 V 1914) ‘Haplochromis ’ cnester Witte & Witte-Maas, RMNH 22-1-1981 MDV V 1981 ' Haplochromis ’ cronus Greenwood, 1959 BMNH 1958.1.1686-89 V ‘ Haplochromis ’ gracilior (Boulenger, 1914) MT 88-16-P-1-38 V ‘ Haplochromis ’ occultidens Snoeks, 1988 MT 80-49-P-5996, 79-31-P- V 1958 ' Haplochromis ’ squamipinnis Regan, 1921 RMNH V Harpagochromis altigenis (Regan, 1922) BMNH 1966.3.9.211-212 V Harpagochromis pectoralis (Boulenger, 1911) BMNH 1966.3.9.289-295 V Harpagochromis serranus (Pfeffer, 1896) BMNH 1962.3.2.56-57 V Harpagochromis spekii (Boulenger, 1906) BMNH 1977.5.24.72 V Harpagochromis victorianus (Pellegrin, 1904) BMNH 1962.3.2.488-494 V Hemitilapia oxyrhynchus (Boulenger, 1902) NMW M Hoplotilapia retrodens Hilgendorf, 1888 RMNH 22-1-1979 V lodotropheus sprengerae (Oliver & Loiselle, NMW 90003 M 1972) Labeotropheus fuelleborni Ahl, 1927 NMW 90006:1-2 M Labrochromis ishmaeli (Boulenger, 1906) RMNH V Labrochromis ptistes (Greenwood & Barel, BMNH 1977.1.10.61-69 V 1978) Labrochromis teegelaari (Greenwood & Barel, BMNH 1977.1.10.18-26 V 1978) Lipochromis maxillaris (Trewavas, 1928) BMNH 1958.1.16.172-179 V Lipochromis microdon (Boulenger, 1906) BMNH 1977.1.28.25-26 V Lipochromis obesus (Boulenger, 1906) BMNH 1958.1.16.143-150 V Lipochromis parvidens (Boulenger, 1911) RMNH V Macropleurodus bicolor (Boulenger, 1906) BMNH 1955.2.10.24-29 V Melanochromis auratus (Boulenger, 1897) NMW 90011:1-2 M Melanochromis labrosus (Trewavas, 1935) NMW 90016 M Neochromis nigricans (Boulenger, 1906) BMNH 1956.7.9.153-165 V Nimbochromis venustus (Boulenger, 1908) NMW 90061 M Orthochromis malagaraziensis (David, 1937) BMNH 1953.11.4.46-58 F Orthochromis polyacanthus (Boulenger, 1899) NMW 90024:1- F Otopharynx lithobates (Oliver, 1984) NMW 89993:1-2 M Paralabidochromis paucidens (Regan, 1921) MT 80-29-P-1499-506 V Paralabidochromis victoriae (Greenwood, BMNH 1955.2.10.181 V 1956) Placidochromis Johnstoni (Gunther, 1893) NMW 89990 M Platytaeniodus degeni Boulenger, 1906 BMNH 1955.2.10.85-86 V Prognathochromis prognathus (Pellegrin, 1904) BMNH 1966.3.9.53-59 V Prognathochromis tridens (Regan & BMNH 1966.3.9.152-165 V Trewavas, 1928) Prognathochromis vittatus (Boulenger, 1901) MT 80-49-P-5482-5485 V Protomelas taeniolatus (Trewavas, 1935) NMW 89985:1-2 M Psammochromis cassius (Greenwood & Barel, BMNH 1959.4.28.250-255 V 1978) Psammochromis graueri (Boulenger, 1914) MT 80-49-P-2404—2408 V 946 E. LIPPITSCH

Appendix Continued.

Psammochromis riponianus (Boulenger, 1911) BMNH 1959.4.28.141-157 V Psammochromis saxicola (Greenwood, 1960) BMNH 1959.4.28.250-255 V Pseudocrenilabrus multicolor (Schoeller, 1903) NMW 90030:1-3 F Pseudocrenilabrus nicholsi (Pellegrin, 1928) NMW 90031:1-2 F Pseudoiropheus zebra (Boulenger, 1899) NMW 90038:1-2 M Ptyochromis sauvagei (Pfeifer, 1896) RM NH M Pyxichromis orthostoma (Regan, 1922) BMNH 1966.3.9.252 V Pyxichromis parorthostoma (Greenwood, BMNH 1966.3.9.253 V 1967) Rheohaplochromis torrenticola Thys, 1963 MT 182787 Sargochromis codringtoni (Boulenger, 1908) MT 187501-509 Schubotzia eduardiana (Boulenger, 1914) BMNH 1972.6.5.8-13 Sciaenochromis ahli (Trewavas, 1935) NMW 89984:1-2 Serranochromis thumbergi (De Castelnau, MT 50684-50695 1861) Taeniolethrinops praeorbitalis (Regan, 1921) NMW Thoracochromis bakongo (Thys, 1964) MT 142012-029, 16948-954 Thoracochromis buy si (Penrith, 1970) BMNH 1984.2.6.55-56 Thoracochromis mahagiensis (David & Poll, MT 44951-955 1937)

Thoracochromis wingatii (Boulenger, 1902) BMNH 1982.6.16.1 Tl Tl Tl Tl 2 < 2 < T} Tl Trematocranus placodon (Regan, 1922) NMW 89991 M Tropheus duboisi Marlier, 1959 NMW 90045 T Tropheus moorii Boulenger, 1898 NMW 90046 T Xystichromis nuchisquamulatus (Hilgendorf, BMNH 1956.7.9.166-167 V 1888) Yssichromis fusiformis (Greenwood & Gee, BMNH 1968.8.30.29-37 V 1969)