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A Preliminary Survey of the Cichlid Fishes of Rocky Habitats in Lake Malawi

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A Preliminary Survey of the Cichlid Fishes of Rocky Habitats in Lake Malawi A preliminary survey of the cichlid fishes of rocky habitats in Lake Malawi A.J. Ribbink, B.A. Marsh, A.C. Marsh, A.C. Ribbink and B.J. Sharp J.L.B. Smith Institute of Ichthyology, Grahamstown Most aquarium fishes exported from Lake Malawi are cichlids INTRODUCTION of 10 rock·frequenting genera collectively referred to by their Chitonga name, Mbuna. These fishes provide a classical exam· In 1962 the first live fishes were exported from Lake Malawi pie of intralacustrine allopatric speciation. The distribution of to aquarists. Since then a lucrative trade in ornamental fishes 196 species is given with notes on habitat preferenda, depth has expanded so that by the mid-seventies more than distribution and behaviour. Considerable emphasis is placed 400 ()()() were exported annually. A group of rock­ on coloration and markings for identification and it is argued that they are important for mate recognition. Brief notes are frequenting cichlid fishes belonging to 10 genera, collectively given on some of the other rocky shore fishes particularly in referred to by their Chitonga name Mbuna (see p.157 for the genus Cyrtocara. The value of the aquarium fish resource details of Mbuna), form the basis of the trade. However, resides principally in its species richness. Most Mbuna with the exception of the pioneering work of Fryer (1959a) species are geographically restricted and stenotopic. The mao on Mbuna ecology, virtually nothing is known of their jority occur in the upper 20 m. Mbuna are trophic specialists, biology, numerical abundance and distribution. Indeed, but have the ability to feed opportunistically on a variety of food resources. Resource partitioning is effected by different more than 70"10 of Mbuna are undescribed and the taxo­ . It ) trophic groups. Evidence suggests that species within a par· nomic affinities of many are uncertain. is convenient, 0 1 ticular trophic group which feed upon apparently identical food therefore, to categorize members of the genera 0 material, collect it from different microhabitats. 2 Pseudotropheus and Melanochromis as species-complexes d S. AIr. J. Zool. 1983, 18: 149 - 310 (monophyletic) and species-groups (polyphyletic). To e t facilitate dissemination of information, undescribed species a d Die meeste van die akwariumvisse wat vanaf die Malawimeer ( are given descriptive names herein which have no taxonomic uitgevoer word, is sikliede van 10 rotsbewonende genera, waar· r e na gesamentlik verwys word as Mbuna, hul Chitonga-naam. validity. A more comprehensive systematic treatment is h s Hierdie visse toon 'n klassieke voorbeeld van binnemeerse forthcoming. i l b allopatriese spesiasie. Die verspreiding van 196 spesies en Although Fryer (l959a) restricted his detailed study to u notas oor hulle habitatvoorkeure, diepteverspreiding en gedrag P a 250-m stretch of rocky shore at Nkhata Bay, his brief ex­ word gegee. Daar word aansienlike klem gell} op kleure en e cursions to other parts of the lake indicated that the species h merktekens vir identifikasie en daar word aangevoer dat dit t assemblages at different localities varied and that at least y belangrik is vir broeipaarherkenning. Kort aantekeninge word b gegee betreffende ander vissoorte wat die rotsagtige kusstreke some species have very limited distributions. Exporters of d e bewoon en in besonder die genus Cyrtocara. Die waarde van ornamental fish confirmed this observation, finding that t n die akwariumvisbron berus hoofsaaklik op sy spesierykdom. many Mbuna species are endemic to particular areas. As a r Die meeste Mbuna-spesies is geografies beperk en is some of these species are restricted to tiny regions and exist g stenotopies. Die meerderheid kom in die boonste 20 m voor. e in small numbers the possibility of over-exploitation was c Mbunas is trofiese spesialiste, maar het die vermo~ om oppor­ n real. Therefore, to determine the effect of commercial ex­ e tunisties te voed op 'n verskeidenheid voedselbronne. c i Voedselvoorraadverdeling word deur verskillende trofiese ploitation on this natural resource and to develop an iden­ l r groepe bewerkstellig. Gegewens dui daarop dat spesies binne tification guide to these and other ornamental fishes, a e d 'n bepaalde trofiese groep, wat op klaarblyklik identiese conservation-orientated research programme was initiated n voedselitems voed, dit vanuit verskillende mikrohabitatte u by the Malawi Fisheries Department in November 1977. versamel. y Part of the programme involved a survey of ornamental a S.-Afr. Tydskr. Dierk. 1983, 18: 149 - 310 w fishes to plot their distribution, to establish baseline data e t a of numerical abundance and to develop a basis for iden­ G tification. Our survey was restricted to Malawian waters, t A.J_ Ribbin"* and A.C. Ribbink e leaving the Tanzanian and Mozambique coasts unexplored. n J.L.B. Smith Institute of Ichthyology. Private Bag 1015, i b Grahamstown, 6140 Republic of South Africa The results are reported here. This report also includes a B.A. Marsh and A.C. Marsh S ecological and behavioural information which, when con­ y Present address: Desert Ecological Research Unit, P.O. Box 953, sidered in conjunction with the zoogeographical data and­ b Walvis Bay, South West Africa/Namibia d historical evidence, throws light on the possible modes of e B.J. Sharp c speciation which may have contributed to the evolution of u Present address: c/o Woodridge College, P.O. Thornhill 6375 d *To whom correspondence should be addressed an estimated 400 - 500 species of Cichlidae which are o r endemic to Lake Malawi. p e Received 23 February 1982; accepted 27 March 1983 Lake Malawi (9°30' - 14°30'S/33°52'- 35°20'£), the R 150 S.-Afr. Tydskr. Dierk. 1983, 18(3) third largest of the African lakes after Lake Victoria and 34" Lake Tanganyika, lies near the southern end of the western Rift Valley in a direction slightly east of south. Lake Malawi is estimated to be between one and two million years old (Dixey 1926; 1941; Banister & Clarke 1980); it is younger than Lake Tanganyika (two to ten million years old: Lowe­ 10· McConnell 1969; Fryer & lies 1972; Banister & Clarke 1980), but older than Lake Victoria (750 000 years old: Greenwood CHILUMBA 1974). The lake is approximately 600 km long, has a maxi­ mum width of about 80 km and a surface area of nearly 31 000 km2 (Figure 1). Its altitude above mean sea level is given as 457 m by Fryer & lies (1972) and 471 m by Banister & Clarke (1980). The maximum depth is uncertain and is given as 704 m (Fryer & lies 1972; Malawi Government USISVA' ___I maps 1974 edition), 758 m (We\comme 1972) and 770 m (Beadle 1981). Although the lake is essentially a single basin it does have two deep troughs in the northern region (Banister & Clarke 1980). By virtue of its tropical setting, Lake Malawi is permanently stratified having a warm CHISUMULU epilimnion overlying a cooler hypolimnion (Beauchamp 12· ISLAND 12· 1953). Similarly, the surface waters are well oxygenated but become poorly oxygenated with increasing depth so that there is very little oxygen between 200 - 250 m and the waters beyond these depths are totally anoxic (Beauchamp 1940; 1953; Ricardo-Bertram, Borley & Trewavas 1942; Eccles 1974). The lake experiences marked seasonal varia­ tions in wind, temperature and precipitation. During the wann season (September to March) winds are light - except in squalls and thunderstorms - and usually blow from the MBENJI ISLAND_-->.-- north-east. In the cool season (April to August) strong . ) south-easterly winds prevail often attaining 40 km h - 1 0 1 (Eccles 1974) and sometimes blowing at 60 - 70 km h - 1 0 MALER I ISLANDS---+ 2 (Ricardo-Bertram et al. 1942). These winds generate sur­ MUMBO ISLAND __-r-- d 14· e face waves 3 - 4 m high (Eccles 1974) and currents which 14· t a can be so strong around islands and rocky outcrops that THUMBI d ( ISLAND divers have to return to boats by pulling themselves along ISLAND r e ropes. h s LAKE i Surface temperatures vary seasonally from 23°C in l b June- July to 28°C in December- January (Eccles 1974) MALOMBE u P and we have recorded surface temperatures of 30°C in 34· e h sheltered bays on hot, windless days in December. t y Seasonal fluctuations in lake level vary between 0,4 and Figure 1 A map of Lake Malawi with an insert of Africa showing the b 2,2 m; normally the lowest level is in December and the peak d location of the lake. The position of some of the more important study e t in May. In addition to these seasonal fluctuations longer sites and of the 200 m depth contour are indicated. n a term progressive changes have occurred. The level is believed r g to have fallen by about 5 m in the period 1865 to 1915 (Pike e c & Rimmington 1965) and records show that it rose by 7,2 m n e between 1915 and 1980 (Crossley 1982). During its substrata of Lake Malawi is suitable for rock-dwelling c i l geological history the level of Lake Malawi altered con­ Mbuna. All islands are entirely or mainly rocky but these r e siderably (sometimes by more than 100m) in response to are separated from one another and from the mainland by d n climatic and tectonic events (Dixey 1926,1941; Fryer 1959a; sandy plains and deep water. Similarly, rocky mainland u Crowley, Pain & Woodward 1964; Banister & Clarke 1980). shores are broken by sandy beaches and other habitats y a Relative to the sculptured coastline of Lake Victoria, Lake which are unsuitable for lithophilous fishes. w e t Malawi has a smooth coast with few major indentations or Fishes were first collected for scientific purposes during a notable bays.
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