SMITHIANA Publications in Aquatic Biodiversity

Special Publication 1 July 2002

A New of the (Teleostei: Cichlidae) from the Lecitu River, Buzi System, .

I. Roger Bills & Olaf L.F. Weyl

Published by the South African Institute for Aquatic Biodiversity ISSN 1684-4149 Margaret Mary Smith (1916 - 1987), James Leonard Brierley Smith (1897 - 1968) with their dog Marlin

The publication series (Monographs, Bulletins & Special Publications) of the SAIAB (formerly the JLB Smith Institute of Ichthyology), in its new format honors James Leonard Brierley Smith and Margaret Mary Smith with the name Smithiana, in rec- ognition of their many years of devoted service to African aquatic biology. Their life’s work, a team effort, established modern ichthyology in southern and laid the groundwork for the expansion of aquatic biology throughout the region.

© 2002, The South African Institute for Aquatic Biodiversity, Grahamstown, South Africa

Front cover photograph: Scales of a preserved coelacanth specimen by James Stapley. © James Stapley, 2002 A New Species of the Genus Chetia (Teleostei: Cichlidae) from the Lecitu River, Buzi System, Mozambique.

I. Roger Bills 1 & Olaf L.F. Weyl 2

ABSTRACT

A new species, assigned to the genus Chetia, is described from the Lecitu River (Buzi system) in Manica Province, Mozambique. The diagnostic features for this species are: deep body (37.4-41.6% SL); large head (39.4-41.0% SL); long pectoral fin (25.0-28.4% SL); low number of lateral line scales (31-32); reduced number of caudal vertebrae (14); a short and deep caudal peduncle (length/ depth ratio 1.0). The new species is compared to other members of the genus and some other serranochromines. The value of certain diagnostic features and the distribution of the genus Chetia are discussed. Additions are made to the Buzi River system ichthyofaunal checklist.

1 South African Institute for Aquatic Biodiversity, Private Bag 1015, Somerset Street, Grahamstown, 6140, South Africa. (e-mail: [email protected]) - Corresponding author. 2 Department of Ichthyology, Rhodes University, P O Box 94, Grahamstown, 6140, South Africa. Current Address - National Aquatic Resource Management Programme (NARMAP), P O Box 27, Monkey Bay, Malawi.

A New Species of the Genus Chetia (Teleostei: Cichlidae) from the Lecitu River, Buzi System, Mozambique.

I. Roger Bills & Olaf L.F. Weyl

INTRODUCTION found in the upper and many of its western elements (Okavango and Kafue rivers). In addition, the The Buzi River system (Fig. 1) in central Mozambique Buzi River is the and only known locality for the has an essentially lower Zambezi fauna (Bell-Cross claroteid Amarginops hildae (Bell-Cross 1973) 1973). The Buzi and Pungwe Rivers are linked in their another species which has close relatives in Zairean and lower reaches which flood during some wet seasons west African river systems (Nichols & Griscom 1917). and the Pungwe itself is linked to the lower Zambezi The serranochromine (Greenwood 1993) are through a swamp region in the western sector of the one of the characteristic upper Zambezi groups and are Gorongoza National Park. Biogeographically, the Buzi relatively poorly represented down the east African system is of interest because our present knowledge coast. meridianus Jubb 1967, Chetia indicates that it forms the southern boundary for several flaviventris Trewavas 1961 and Chetia brevis Jubb 1968 lower Zambezi species e.g. Malapterurus shirensis occur in South Africa and southern Mozambique but Roberts 2000, Heterobranchus longifilis Valenciennes all have restricted distributions even within the systems 1840, Distichodus schenga Peters 1852, D. mossambicus in which they occur. Except for the Shire River in Peters 1852, Mormyrops anguilloides (Linnaeus 1758) and Malawi, where Serranochromis robustus robustus Hippopotamyrus ansorgii (Boulenger 1905) (Bell-Cross (Günther 1864) occurs, there are no other Serranochromis 1972 & 1973, Roberts 2000). These species, while Regan 1920 in the lower Zambezi or associated systems. characteristic lower Zambezi fauna, have affinities with During an angling expedition to the Lecitu River species presently found in the Zaire system and are not (Buzi system, 19° 55' S 33° 22' E) in January 1997, one

Figure 1. Buzi River system in central Mozambique.

2 of us (OW) collected six specimens of an undescribed Chetia brevicauda sp.n. serranochromine cichlid. The specimens belong to the (Figs. 2 - 5) genus Chetia Trewawas 1961 as redefined by HOLOTYPE. RUSI 56660, 122.5 mm SL, O. Weyl, Lecitu Greenwood (1993). The purpose of this paper is to River at Dombe pontoon (19° 55’S 33° 22’E), Manica describe this new species. Comparisons are made with Province, Mozambique, 20/January/1997. other members of the genus Chetia and certain PARATYPES. RUSI 58260, (5) 82-125 mm SL, collected characters diagnosing the genus are reviewed. The with holotype. distribution of the genus raises issues of historical linkages with the Buzi and other southern African river DIAGNOSIS: A member of the genus Chetia systems. distinguished from other members of that genus by the following characteristics. Head large 39.4-41.0% SL vs METHODS 34.4-36.0 in C. mola and 33.0-37.0 in C. brevis. A deep bodied species 37.4-41.6% SL vs. 32.4-35.9 in C. mola, The fish were collected by angling using a number four 33.3-34.2 in C. welwitschi and 29.0-37.0 in C. brevis. hook with live earthworms as bait. The specimens were Pectoral fin relatively long 25.0-28.4% SL showing some fixed in a 10% formalin solution and preserved in 70 % overlap with C. brevis 18.0-26.0 but greater than C. ethyl alcohol. The morphometric measurements and flaviventris 21.7-24.7, C. mola 22.8-23.3 and C. welwitschi meristic counts follow Barel et al. (1977). Lengths given 18.1-20.6. Lateral line scales 31-32 vs 32-34 in C. brevis in the text are SLs unless otherwise stated. All counts and C. welwitschi, 34 in C. mola and 34-35 in C. were taken from the left side of the fish. Each specimen flaviventris. Caudal vertebrae are low in number with was radiographed (radiographs are held at the South 14 in all specimens vs 15 in C. welwitschi and 16-18 in C. African Institute for Aquatic Biodiversity, mola and C. flaviventris. The caudal peduncle is short Grahamstown) to determine vertebrae numbers and fin and deep with a mean CPL/CPD ratio of 1.0 vs 1.3-1.5 elements. Dorsal and anal fin ray numbers were in C. flaviventris and 1.4-1.5 in C. welwitschi. determined by counting the proximal portions of rays and pterigiophores. Last anal and rays were COUNTS AND MEASUREMENTS: Holotype values counted as one ray if they were supported by a single first followed by range of paratypes in parentheses. pterigiophore. Measurements in percent SL: head length 39.4 (39.6- The abbreviations of institutions follow Leviton et 41.0); body depth 37.4 (37.4-41.6); predorsal length 38.9 al. (1985). The material examined was from the (39.7-43.0); preanal length 68.2 (68.4-71.3); pectoral fin following institutions: South African Institute for length 25.0 (26.2-28.4); pelvic fin length 25.6 (26.2-27.8); Aquatic Biodiversity, Grahamstown, South Africa posterior dorsal spine length 13.5 (13.3-14.7); highest (RUSI); the Albany Museum, Grahamstown, South dorsal soft ray 23.1 (17.8-25.1); posterior anal spine Africa (AMG); Natural History Museum, London, U.K. length 12.1 (11.9-14.1); highest anal soft ray 21.2 (20.2- (BMNH); and the Royal Ontario Museum (ROM), 23.9); caudal peduncle length 12.7 (12.8-13.9); caudal Canada. fin length 27.8 (28.4-30.2). Measurements in percent HL: snout length 32.3 (28.8-34.6); preorbital depth 19.0 (16.1-

Figure 2. Chetia brevicauda sp. nov., holotype, RUSI 56660, 122.5 mm, Lecitu River near Dombe, Manica Province, Mozambique.

3 19.7); eye length 20.3 (20.1-23.6); inter-orbital width 17.2 (17.6-18.6); cheek depth 30.2 (27.5-33.3); lower jaw length 46.8 (44.5-49.3); upper jaw length 37.9 (34.5-39.4). Ratios: caudal peduncle length/ caudal peduncle depth 1.0 (1.0); caudal fin scales/ caudal fin length 0.6 (0.4- 0.6). Counts: upper lateral line scales 21 (20-22); lower lateral line scales 11 (10-12) total LL scales 32 (31-32); cheek scale rows 5 (4-5); scales round caudal peduncle 15 (14-15); dorsal fin spinous rays 14 (14-15); dorsal fin branched rays 12 (11-13); total dorsal fin rays 26 (26- 27); anal fin spinous rays 3 (3); anal fin branched rays 9 (9-10); total anal fin rays 12 (12-13); abdominal vertebrae 15 (15); caudal vertebrae 14 (14); gill rakers (ceratobranchial) 11 (11-12).

DESCRIPTION: Based on six adult and sub-adult specimens. Measurements and counts for the holotype and paratypes are given above. Figure 2 shows body form, fin positions and shapes and preserved colouration pattern. Due to the small number of specimens only a single specimen (RUSI 58260, paratype #4, 96.6mm) was dissected for the description of the jaws and teeth.

Body. Compressed and deep with the greatest body depth at the origin of the pelvic fins. Predorsal length short. Head large and negatively allometric with SL. Preorbital depth, snout length and cheek depth positively allometric with SL. Eye large, situated dorsally and negatively allometric with SL. Caudal peduncle short and deep.

Upper jaw (Fig. 3a & b). Premaxillary pedicel is slightly longer (103%, 12.5 mm in a specimen 96.6 mm) than the dentigerous arm of the premaxilla. The premaxillary pedicel slopes at an angle of 77 ° from the horizontal. The dental arcade comprises two rows of closely set unicuspid teeth. In the outer row there are (left side 27, right side 26) 53 teeth. These teeth alter in curvature and implantation with position being slightly recurved and erect at the anterior symphysis and changing to more recurved and recumbent posteriorly. The outer row teeth extend 81-83% along the dentigerous arm of the dentary. The inner row teeth are slightly smaller and closer set than those in the outer row and are all recumbent (Figure 3b). The inner row teeth comprise (left side 19, right side 21) 40 teeth and they extend 43% along the dentigerous arm of the dentary.

Lower jaw (Fig. 3c & d). Relatively long with a rounded dental arcade. There are two rows of erect, slightly recurved and irregularly spaced unicuspid teeth. The outer row comprised 37 teeth (left side 19, right side 18) and extends 66.2-71.1% of the dentigerous arm of Figure 3. Jaws of Chetia brevicauda sp. nov. (RUSI 58260, the dentary. The inner row teeth comprise a few anterior paratype #4, 96.6 mm, scale bar = 1 mm) - a. left lateral view teeth (left side 6, right side 4) which extend 22.3-22.8% of the premaxilla, b. ventral view of the anterior left section of the dentigerous arm of the dentary. of the premaxilla showing the varying implantation of the two tooth rows, c. left lateral view of the lower jaw, d. dorsal Maxilla (Fig. 4a). The maxilla is stout (maximum length view of the lower dental arcade. 12.1 mm) with its maximum width approximately 4/5 4 along the bone on the posterior flange (3.5 mm).

Lower pharyngeal bone & teeth (Fig. 5a). The pharyngeal bone is triangular and not thickened. The dentigerous surface is 62% of the total length and 71% the total width of the bone. The ten posterior-median teeth are slightly enlarged while the remainder are slender and bevelled becoming more slender towards the edges of the bone. There are 10-11 teeth along the centre and 19 along the posterior edge of the bone.

Gill rakers (Fig. 5b). Simple, relatively short gill rakers and number 11-12 on the ceratobranchial of the first gill arch. The three to four anterior rakers are reduced to small bumps of tissue. Total gill rakers are 15-16.

Scales. Predominantly cycloid with a few weakly ctenoid scales on the lower flanks. Thoracic region fully covered with small, deeply embedded scales that reduce gradually from the larger flank scales. Scales are present on the head up to the interorbital area and cover the cheeks up to the lachrymal bone (usually 4-5 rows which are sometimes not arranged in defined rows). Scales between the orbit and preopercular groove 3-5. Approximately half of the caudal fin is scaled but Figure 4. Lateral views of left maxilla bones for a. C. there are no scales on the other fins. There are 6-7 scales brevicauda (RUSI 58260, paratype #4, 96.6 mm), b. C. flaviventris (RUSI 61095, 102.7 mm) and c. S. robustus jallae between the pelvic and pectoral fin bases, 5-7 scales (RUSI 52506, 103.5 mm). between the lateral line and the origin of the dorsal fin.

Figure 5. a) Dorsal, ventral and lateral views of the lower pharyngeal bone and

5 lateral, dorsal lateral and dorsal medial bands. The mid- lateral and dorsal medial bands both continue anteriorly up to the eye. There are 6-7 short and indistinct vertical bars on the body. The membranes of the fins are pigmented brown to varying degrees with the darkest section being the anterior dorsal fin. The membranes of the dorsal fin posterior (posterior of spine 9) and caudal fin anterior are covered with dark brown maculae. The centre of the soft rayed part of the anal fin has a series (up to nine) of eggs spots, not quite as large or as darkly pigmented as the caudal and dorsal Figure 5. b) the first gill arch for Chetia brevicauda sp. maculae. The belly and ventral flank are unpigmented. nov.(RUSI 58260, paratype #4, 96.6 mm, scale bar = 1 mm). Colour in life. Unknown.

The last two scales in the upper LL are separated from Habitat (Fig. 6). All specimens were caught near a the dorsal fin base by one small and one large scale submerged tree in a quiet backwater protected from while anterior to these there are two or more large the main river flow by a rocky outcrop. scales. Distribution. Known only from the Lecitu River (Buzi Sensory structures. Simple finger-like projections (<1 system) at Dombe (19° 55’S 33° 22’E), Mozambique. mm) are present on the lower operculum, inner pelvic fin membranes and on skin on the ventral body directly Etymology. The genus name Chetia is feminine above the pelvic area near the fin joint. These are (Eschmeyer 1998). The species name brevicauda, Latin possibly tactile sensory structures and indicative of for short and tail respectively (Brown 1979), describes contact with the substrate when foraging. Lateral line the low number of caudal vertebrae and the short sensory canals on the head are branched forming caudal peduncle. clusters of tubes. Enhanced lateral line development may compensate for poor vision in turbid conditions. Associated fish community. The Dombe pontoon site was rather poorly sampled and only the following Fins. Positions and pigment patterns of fins in species were collected with C. brevicauda: Distichodus preserved specimens are shown in Figure 2. Pelvic fins mossambicus, , zambezensis just reach the origin of the anal fin. Caudal fin slightly and Glossogobius callidus. emarginate with rounded edges. REMARKS Colour in alcohol. Preserved material (Fig. 2) has a beige background colour that is pigmented brown by a Generic placement: Greenwood (1993) stated that there series of spots and lines. The anterior head is pigmented is no single diagnostic autapomorphy to distinguish with what appear to be fragmented nostril, interorbital, members of the genus Chetia. Rather, there are a suite supraorbital and nape stripes i.e. paired blotches either of characters which exclude them from other riverine side of the head centre line in these positions. Mental ( Regan 1920, and opercular blotches are present. There are three Serranochromis Regan 1920, poorly defined and broken lines along the body: mid-

Table 1. Comparative measures of key features distinguishing C. brevicauda from other Chetia species (measures taken from Greenwood 1984, 1992, 1993).

6 Figure 6. The Lecitu River at Dombe, looking upstream from the pontoon. (Steindachner 1866) and Thoracochromis Greenwood circumpeduncular scales (14); low LL scales (31-32); low 1979). Greenwood considered Chetia to be closely caudal (14) and total (29) vertebrae. Consequently, the related to the genus Serranochromis based on the inclusion of this new species in the genus Chetia, which following synapomorphic features: two or more post- we believe is correct, does increase the overlap between orbital scale rows (from Lippitsch 1990); high numbers generic characters for the serranochromine genera of caudal vertebrae (15-17); and an increased number (Table 2). of circumpenduncular scales (16-20). Serranochromis apomorphies not shared with Chetia are: high numbers Generic characteristics: With the inclusion of Chetia of abdominal (15-19) and total vertebrae (31-36); high brevicauda sp. nov. the generic definition of the Chetia lateral line scale counts (34-41); and unicuspid teeth in needs to be altered slightly. The lower limit of the caudal juveniles. Consequently, Greenwood suggested the vertebral count is now 14 and the lower limit of the rather unsatisfactory diagnosis for Chetia as “a total vertebral count is 29. The range of scales in the Serranochromis lacking the apomorphic features of that lateral line series is now 31-35 and that of gill rakers is genus”. 9-12. Table 2 is a summary of the serranochromine The new species has three to five post-orbital scales genera (from Greenwood 1993) together with the which clearly places it within the Serranochromis-Chetia altered characteristics for Chetia. “group” but it does not exhibit the other two characteristic features which are high numbers of caudal Anal fin spots: The anal fin spot pattern recognised by vertebrae or circumpeduncular scales. The new species Greenwood (1993: 34 & 36) for Pharyngochromis, differs from Serranochromis in that it has a lower LL Serranochromis, Sargochromis and Chetia was of small, scale count and lower numbers of abdominal (and numerous non-ocellated spots similar to the maculae consequently total) vertebrae. The new species also has on the dorsal and caudal fins. This was considered a a smaller mouth compared to all other Serranochromis, key character distinguishing these genera from other a feature not referred to by Greenwood, but one which riverine haplochromines which have ocellated spots i.e. is used as a key distinguishing feature by Skelton (2001). a clear area surrounding the outside of the spot. The six specimens comprising the type series are all Examination of live specimens of the type species above 96mm SL. The presence of bicuspid teeth in C. flaviventris, collected in the Roodeplaat Dam near juveniles, another Chetia characteristic, was therefore Pretoria, indicated that the pattern of the anal spots in not verified. The form of anal egg spots was also these cichlids was slightly different than that described considered an important feature by Greenwood (1993). by Greenwood (1979: 274-275 and 1993: 34). Two Only preserved material was available at the time of specimens, collected and photographed at the same description and, as will be discussed later, the condition time are illustrated - an immature male and a mature of egg spots in live and preserved material changes male in breeding colouration (Figure 7). The immature considerably. male has ocellated eggs spots, i.e. an orange spot The new species exhibits certain features which do surrounded by a grey/dark circle and then by a clear not fall within Greenwood’s (1993) definition of Chetia region. The anal fin is pigmented yellow but is not but which do fit Sargochromis and Pharyngochromis opaque. In preservation these spots are grey, evenly characteristics. These features are: low pigmented and have no clear surround visible. The egg 7 spots of the sexually mature male were yellow 10 teeth (Fig. 3d). In other haplochromines, inner row anteriorly becoming orange-centered, fading to yellow teeth comprise greater numbers of teeth and tooth rows edges posteriorly and surrounded by a grey/dark ring. and these extend down the lateral part of the dentary. No clear area was visible and the whole anal fin was Slight differences were observed between C. brevicauda whitish and opaque. In preservation these spots are and C. flaviventris and S. robustus jallae. The arrangement grey, some solid and some with a darker edges but none of the dentary teeth was the same in the three species, with clear surrounds, and the fin is white and opaque. but on the premaxilla of C. flaviventris there were three Within this sample of C. flaviventris, RUSI 61095, 26: rows of teeth antero-medially reducing laterally to two 55-148 mm, there was considerable variation in the almost entire rows and the posterior teeth were not as numbers, colour and form (shape and ocellation) of the recumbent as in C. brevicauda. The maxilla of C. anal spots. In general, the mature males had larger and brevicauda was much larger and deeper than in either more numerous anal spots than immature males. C. flaviventris or S. robustus jallae despite the specimens Consequently, we suggest this character warrants a of latter species being slightly longer (Figure 4). more thorough investigation before it is used for determining relationships within the serranochromine Distribution: Members of the genus Chetia exhibit a group. disjunct distribution pattern (Table 3), similar to several The only consistent difference noted in anal fin spot other species in the region such as Mesobola brevianalis patterns during our superficial study of mostly museum (Boulenger 1908) and Barbus bifrenatus Fowler 1935 serranochromine material is that between (Skelton 1994). It adds further support to the widely Serranochromis robustus jallae (Boulenger 1896) and all held hypothesis of connections between the Zambian other Serranochromis (including the form Congo, Kafue, Upper Zambezi, Kunene, Limpopo, of S. robustus robustus). In all the specimens we have Incomati and Buzi Rivers (Bell-Cross 1972, Gaigher & examined of S. robustus jallae the anal spots are confined Pott 1973, Skelton 1994). However, the finer details of to the posterior half of the anal fin while in other species river connections remain to be determined and a eggs spots are distributed over the entire fin with phylogenetic analysis of these species and species- increasing coverage with increasing SL (see Skelton 2001 groups is now needed. Two possible routes for Chetia for colour illustrations). This character may indicate that brevicauda or its ancestor to colonise the Buzi system S. robustus from the upper Zambezi and Lake Malawi are: 1) from the north (Zaire system) through the lower are separate species. Zambezi or east coast rivers (together with many other “Zairean” species); or 2) from the upper Zambezi- Dentition: The teeth and their arrangement are typical Limpopo connection and then north. of both Chetia and Serranochromis (Greenwood 1979: The limited distributional ranges of each of the five 300). Particularly characteristic is the antero-medial arc Chetia species is also noteworthy. Three of the species of inner teeth on the dentary that comprises less than are poorly known, though, the late discovery of Chetia

Table 2. A summary of characters for the southern African riverine cichlid genera, taken from Greenwood 1979, 1984, 1992 & 1993. New species data added to Chetia section.

8 Figure 7. Anal fin spots of Chetia flaviventris from Roodeplaat Dam, near Pretoria, South Africa - a) a live adult male in breeding condition (144.7 mm), b) a live immature fish (101.9 mm).

mola Balon & Stewart 1983 and C. brevicauda and the Cross 1974 (Kneriidae), Varicorhinus pungweensis Jubb rarity of Chetia welwitschi (Boulenger 1898) seems to 1959 () and Barbus manicensis Pellegrin 1919 indicate they are naturally uncommon or have restricted (Cyprinidae). distributions within their geographical ranges. The two New records since Bell-Cross’ 1973 survey have species which are well known, C. flaviventris and C. raised the number of species known from the Buzi brevis, also have restricted ranges, the Limpopo and system to 63. A new checklist is given in Table 4. Names Incomati systems respectively. follow Skelton (1993, 2001, 2002) and museum accession numbers are given for most new records collected by Buzi River fauna: The discovery of C. brevicauda Weyl and Bills. Three exotics are present in the Buzi increases the status of the Buzi system as an area of system. Two of these (Oreochromis niloticus (Linnaeus high endemicity and special scientific (biogeographic) 1758) and Micropterus salmoides (LacepPde 1802)) were interest as already noted by Skelton (1984). Other introduced into Zimbabwean sections of the Buzi and endemics include the claroteid catfish A. hildae and the have now moved downstream into Mozambique (Weyl amphiliid catfish Amphilius laticaudatus Skelton (1984). 1998). The third exotic (Cyprinus carpio Linnaeus 1758) Additional species which have slightly wider was introduced into the Sussundenga Aquaculture distributional ranges, being endemic to the Buzi and station in Manica Province, Mozambique during 1996 Pungwe systems, are Parakneria mossambica Jubb & Bell- (pers. obs., O.W.).

9 Table 3. The distribution of members of the genus Chetia.

Notable absences from the checklist are several collected by Mr S.S. du Plessis. Paratypes: BMNH species which occur in the lower Zambezi and northern 1957.11.16.2-3, (2) 72.7-90.9 mm; RUSI 45386, (1) 118.2 South Africa such as Clarias ngamensis Castelnau 1861, mm. Other material: RUSI 28969, (2) 68.5-78.3 mm; Clarias theodorae Weber 1897, multispine RUSI 53629, (6) 47.5-94.5 mm; RUSI 61095, (26) 55-148 Peters 1844 and Microctenopoma intermedium (Pellegrin mm. 1920). These are typical floodplain species and are Chetia brevis. Holotype. AMG/PF951, 120.5 mm, possibly indicative of the fact that the floodplain reaches Lomati River, South Africa, collected I.G. Gaigher, of the Buzi system are the most poorly known. We September 1967. Paratypes. AMG/PF952, 73.8-89.3mm. anticipate that these and several estuarine species Other material: RUSI 27639, (1) 70.2 mm; RUSI 63749 known to enter freshwaters e.g. Elops machnata (ForsskDl (5), 66.6-80.7 mm; RUSI 63750 (2), 43.2-59.1 mm. 1775) and Acanthopagrus berda (ForsskDl 1775), will be Chetia mola. Holotype. ROM 29825, 117.9 mm, Luongo found when thorough surveys are conducted in the River above the mouth of the Chambwishitu tributary, lower reaches of the Buzi. , collected by E.K. Balon, 17 November 1970. Paratypes. ROM 29826, (3) 74.6-132.0 mm. ADDITIONAL MATERIAL EXAMINED Chetia welwitschi. Holotype. BMNH 1864.7.13.62, 99.7 mm, Fluilla (Huila?), . Chetia flaviventris. Holotype: BMNH 1957.11.16.1, 104.1 Thoracochromis buysi. RUSI 63330, (38) 46.8-137.3 mm. mm, Buffelspoort Dam, Sterkstroom, tributary of Thoracochromis albolabris. RUSI 59465, (6) 35.5-72.7 mm. Crocodile River, Limpopo system, South Africa, Pharyngochromis acuticeps: RUSI 39049, (1) 94 mm.

Table 4. A revised checklist of the of the Buzi River system, Mozambique (*indicates new records since Bell-Cross 1973 and museum numbers indicate voucher specimens of these, names follow Skelton 1993, 2001, 2002).

Protopterus annectens brieni Barbus barnardi Leptoglanis sp. Barbus eutaenia Anguilla marmorata RUSI 58226* Barbus kerstenii RUSI 53688* Clarias gariepinus A. bengalensis Barbus lineomaculatus A. mossambica Barbus macrotaenia Malapterurus shirensis Barbus manicensis Mormyrops anguilloides Barbus marequensis Synodontis zambezensis longirostris Barbus paludinosus RUSI 62013* neumanni RUSI 61588* Marcusenius macrolepidotus Barbus radiatus Chiloglanis pretoriae Hippopatamyrus discorhynchus Barbus toppini H. ansorgii Barbus trimaculatus Aplocheilichthys johnstoni Barbus viviparus Aplocheilichthys katangae RUSI 61555* Kneria auriculata Cyprinus carpio * Parakneria mossambica RUSI 58243* Labeo altivelis Micropterus salmoides RUSI 58251* Labeo congoro Herklotsichthys quadrimaculatus RUSI 61558* Labeo cylindricus Astatotilapia calliptera Labeo molybdinus Chetia brevicauda RUSI 58260* Hydrocynus vittatus Opsaridium zambezensis Oreochromis mossambicus Distichodus mossambicus Varicorhinus pungweensis Oreochromis niloticus RUSI 58239* Distichodus schenga Oreochromis placidus Brycinus imberi Amarginops hildae Pseudocrenilabrus philander RUSI 61898* Brycinus lateralis Tilapia rendalli Micralestes acutidens Schilbe intermedius Tilapia sparrmanii

Barbus annectens Amphilius uranoscopus Awaous aenofuscus Barbus afrohamiltoni A. laticaudatus AMG P5815* Glossogobius callidus RUSI 61562* Glossogobius giurus 10 Serranochromis thumbergi: RUSI 62392, (1) 200 mm. Gaigher, I.G. & Pott, R.Mc.C. 1973. Distribution of Serranochromis macrocephalus: RUSI 52438, (1) 177 mm. fishes in . S. Afr. J. Sci., 69: 25-27. Serranochromis robustus robustus: RUSI 50055, (5) 31-115 Greenwood, P.H. 1979. Towards a phyletic classifica- mm. tion of the “genus” (Pisces, Serranochromis robustus jallae: RUSI 62394, (1) 195; RUSI Cichlidae) and related taxa. Part I. Bull. Brit. Mus. 52506, (16) 48-144 mm. (Nat. Hist.), (Zool.) 35(4), pp. 265-322. Serranochromis angusticeps: RUSI 21292, (1)134.2 mm. Greenwood, P.H. 1984. The species : RUSI 22138, (3) 135-142 mm. (Teleostei. Cichlidae) of the Cunene and certain Serranochromis longimanus: RUSI 28897, (3) 79-190 mm. other Angolan rivers. Bull. Brit. Mus. (Nat. Hist.), Serranochromis meridianus: RUSI 47106, (1) 87.6 mm. (Zool.) 47(4), pp. 187-239. Sargochromis codringtonii: RUSI 52507, (1) 116 mm. Greenwood, P.H. 1992. A revision of the monotypic cichlid genus Pharyngochromis (Teleostei, ACKNOWLEDGEMENTS Labroidei). Bull. Brit. Mus. (Nat. Hist.) (Zool.), 58(1): 37-52. The following curators/institutions are thanked for Greenwood, P.H. 1993. A review of the material loans: Royal Ontario Museum, Canada, Ann- serranochromine cichlid fish genera Marie Hine, British Museum of Natural History, Pharyngochromis, Sargochromis, Serranochromis and London, U.K.; Guy Teugels, Central African Museum, Chetia (Teleostei: Labroidei). Bull. Brit. Mus. (Nat. Tervuren, Belgium; Jim Cambray and Pat Black, Albany Hist.) (Zool.), 59(1), pp. 33-44. Museum, Grahamstown, South Africa. The following Jubb, R. 1968. A new Chetia (Pisces, Cichlidae) from people are thanked for helping collect comparative the Incomati River system, Eastern Transvaal, material: Daksha Naran, Ben van der Waal, Andrew South Africa. Ann. Cape Prov. Mus. (Nat. Hist.), Deacon, Johann Engelbrecht and Kevin Roberts. The 6(7): 71-76. staff of the South African Institute for Aquatic Leviton, A.E., R.H. Gibbs, E. Heal & C.E. Dawson Biodiversity are thanked for their assistance in many 1985. Standards in herpetology and ichthyology: aspects of this work. In particular, discussions with Paul part I. Standard symbolic codes for institutional Skelton on cichlid , Sally Terry aided with resource collections in herpetology and ichthyol- much of the laboratory work and made all X-rays and ogy. Copeia (3): 802-832. Elaine Heemstra drew the figures. Denis Tweddle and Lippitsch, E. 1990. Comparative investigation on Ofer Gon made valuable comments on this work and scale characters in cichlids. Ann. Mus. Roy. Afr. Jos Snoeks is thanked for improvements made to the Centr., Sci. Zool., 262: 97-102. final draft. Nichols, J. T. & L. Griscom.1917. Fresh-water fishes of the Congo basin obtained by the American LITERATURE CITED Museum Congo expedition, 1909-1915. Bull. Am. Mus. Nat. Hist., 37 (25): 653-756, Pls. 64-83. Balon, E.K. & Stewart, D.J. 1983. Fish assemblages in Roberts, T.R. 2000. A review of the African Electric a river with unusual gradient (Luongo, Africa- Catfish Family Malapteruridae, with descriptions Zaire system), reflections on river zonation, and of new species. Occ. Pap. Ichthyol., 1: 1-15. description of another new species. Environ. Biol. Skelton , P.H. 1984. A systematic revision of the Fish., Vol. 9 (nos. 3-4): 225-252. species of the catfish genus Amphilius (Siluroidei: Barel, C.D.N., M.J.P. van Oijen, F. Witte & E.L.M. Amphiliidae) from east and southern Africa. Ann. Witte-Maas 1977. An introduction to the Cape Prov. Mus. (Nat. Hist.) 16(3): 41-71. taxonomy and morphology of the haplochromine Skelton , P.H. 1993. Scientific and common names of Cichlidae from Lake Victoria. Neth. J. Zool., 27(4): Southern African freshwater fishes. JLB Smith parts A & B, 333-389, figs 1-65. Inst. Ichthyol., Spec. Publ., 56: 1-34. Bell-Cross, G. 1972. The fish fauna of the Zambezi Skelton , P.H. 1994. Diversity and distribution of River system. Arnoldia, 5(29): 1-19. freshwater fishes in East and Southern Africa. Bell-Cross, G. 1973. The fish fauna of the Buzi River Ann. Mus. Roy. Afr. Centr., Zool., 275: 95-131. system in Rhodesia and Mozambique. Arnoldia, 6 Skelton , P.H. 2001. A Complete Guide to the Freshwater (5): 1-14. Fishes of Southern Africa. Struik Publishers, Cape Boulenger, G.A. 1898. A revision of the African and Town. 2nd Edition, pp. 395. Syrian fishes of the family Cichlidae. Proc. Zool. Skelton , P.H. 2002. Changes to the scientific and Soc. Lond. 1898: 132-152. common names of southern African freshwater Brown, R.W. 1979. Composition of scientific words. fishes. Afr. J. Aquat. Sci. In Press. Smithsonian Institution Press, Washington, pp. Trewavas, E. 1961. A new cichlid fish of the Limpopo 882. basin. Ann. S. Afr. Mus., 46(5), pp 53-56. Eschmeyer, W.N. 1998. . Special Weyl, O.L.F. 1998. The dynamics of a subtropical lake publication No. 1 of the Centre for Biodiversity fishery in central Mozambique. PhD thesis, Research and Information, California Academy of Rhodes Univ., 1999. 205pp.Unpubl. sciences. Vols. 1-3.

13 SMITHIANA SERIAL PUBLICATIONS FROM THE SOUTH AFRICAN INSTITUTE FOR AQUATIC BIODIVERSITY

The Institute publishes original research on systematics, ecology, biology and conservation of fishes. Three series are produced at irregular intervals: the Special Publication series, the Bulletin series and the Monographs series. Acceptance of manuscripts for publication is subject to the approval of reviewers from outside the Institute. Priority is given to papers by staff of the Institute, but manuscripts from outside the Institute will be considered if they are pertinent to the work of the Institute or use the Institute’s collections. Colour illustrations can be printed at the expense of the author. Page charges will be mandatory for all non-staff or non-associates of the Institute, these charges will be at the discretion of the Editor. Fifty free copies of the publication will be supplied to the author or senior author. Additional reprints may be ordered at cost price. Publications of the Institute are available by subscription or in exchange for publications of other institutions. Lists of the Institute’s publications are available from the Publication Secretary at the address below. Past Bulletins are also available on the internet at http://bioline.bdt.org.br/fb, past Special Publications are at http://bioline.bdt.org.br/fs

INSTRUCTIONS TO AUTHORS

Manuscripts shorter than 30 pages will generally be published in the Special Publications series; longer papers will be considered for the Bulletin or Monographs series. Please follow the format of a recent Bulletin or Special Publication. The typescript must be double-spaced throughout with 25 mm margins all round; two hard copies must be submitted to the Editor. Each table or figure should be on a separate page and numbered with an Arabic numeral (not in sequence with text pages). All maps, graphs, charts, drawings and photographs should be numbered as figures. If two or more illustrations are grouped as one figure, they must be trimmed and spaced (but not mounted) as intended for final reproduction. Each part of a composite figure must be labelled with a capital letter; typewriter lettering is not acceptable. Illustrations larger than 21 x 30 cm should be avoided, or sent as a reduced bromide/line shot. Legends for figures should be on a separate page. A computer diskette/ CD, with the text in RTF, MSWord or Corel WordPerfect format, must be submitted with the hard copies. The inclusion of digital copies of all images and figures in acceptable formats (see web page) will expedite publication of the manuscript . See extended Instructions to Authors at www.saiab.ru.ac.za/pubs.htm

STYLE OF THE HOUSE

Hyphens: Certain substantive compounds are hyphenated: gill-raker, soft-ray, type-species, type-locality, type-series, type-specimen. Other words often used together are not hyphenated unless they are used in adjectival expressions before a noun: anal fin / anal-fin rays; lateral line / lateral-line scales; gill arch / gill-arch filaments, etc. Word usage: Although the following word pairs are often used interchangeably, we believe that consistent use of the first word as a noun and the second as an adjective will improve the precision of our writing: mucus / mucous; maxilla / maxillary; opercle / opercular, operculum / opercular. The operculum (= gill cover) comprises (usually) four separate bones: opercle, subopercle, preopercle and interopercle. The words preoperculum, suboperculum and interoperculum are unnecessary substitutes and not to be used for preopercle, subopercle and interopercle. The plural of operculum is opercula. Decimal comma versus decimal point: Contrary to most journals published in South Africa and some European countries, we will not use a comma in place of a decimal point. Most computers do not read a comma as a decimal point. In addition, it is common in ichthyological papers to give sequences of measurements that include decimal numbers, with each measurement separated by a comma. If the comma is used to separate items in a series, as well as being used to indicate a decimal number, it will cause considerable confusion. Fin formulae: Fin formulae will be designated as follows: D XII,10-12 indicates on continuous fin with 12 spines and 10-12 soft (segmented) rays; DX/I,10-12 indicates a fin divided to the base in front of the last spine; and D X+I,12 indicates two separate dorsal fins, the first with 10 spines and the second with 1 spine and 12 soft rays. If it is necessary to differentiate branched and unbranched soft-rays, lower-case Roman numerals will be used for unbranched rays and Arabic numerals for branched rays, e.g. D iii,S. Principal caudal-fin rays are defined as those that touch the hypural bones. The number of principal caudal rays is usually the number of branched rays plus two. If the principal caudal rays are in two separate groups, the number of rays in the dorsal group is given first: thus, @principal caudal rays 8+7@ means that there are 15 principal caudal rays, with 8 rays in the dorsal group and 7 in the ventral group. Abbreviations: Abbreviations normally end with a full stop: et al., e.g., etc., n.b., (note: these commonly used abbreviations of Latin words are not italicized). Dr (Doctor) and Mr (Mister) and compass directions (north, west, northwest, etc.) are abbreviated using capital letters without full stops: N, W, NW. We recommend the following abbreviations for ichthyological terms: SL - standard length, TL - total length, FL - fork length, GR - gill-rakers, LL - lateral line.

M. E. Anderson, Editor L-A. Fargher, Layout and production South African Institute for Aquatic Biodiversity (formerly J.L.B. Smith Institute of Ichthyology), Private Bag 1015, Grahamstown 6140, South Africa. Published by the South African Institute for Aquatic Biodiversity Private Bag 1015, Grahamstown, South Africa, 6140 www.saiab.ru.ac.za

formerly known as the JLB Smith Institute of Ichthyology