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The nominal species Petrotilapia tridentiger Trewavas, a rock-frequenting fish from Lake Malawi, occurs in three distinct male and female colour forms at Monkey Bay. Field observations on courtship and aggression indicate that the three colour forms are distinct species. The taxonomy of the three species of Petrotilapia that occur at Monkey Bay is revised. R. tridentiger is redescribed and two new species, P. genalutea and P. nigra, are described. The three species are very similar morphologically and are considered to be sibling species. The only morphological character which reliably separates the three sibling species is live coloration: Live coloration is considered to be an important component of the specific mate recognition systems in Petrotilapia species. The coloration of a further fourteen forms of Petrotilapia is described and it is suggested that these forms are valid species. Space resource partitioning plays an important role in facilitating the coexistence of Petrotilapia sibling species. Space is partially partitioned in terms of depth range. In shallow water, where all three species are sympatric, there are differences in the feeding sites used by the various Petrotilapia forms. Changes in niche breadth and overlap, which are associated with changes in the abundance of epilithic algae, indicate that competition for feeding space occurs between the three Petrotilapia species. The Petrotilapia species partition their resources according to a social dominance hierarchy. Dominant forms, such as territorial males, R. tridentiger females and schooling R. genalutea females, have access to rich food supplies which occur within the territories of certain highly aggressive Pseudotronheus species. are facultative and opportunistic .feeders. Petrotilapia species Speciation in allopatry is considered to be the mechanism which adequately explains the current diversity of the genus Petrotilapia. The importance of an integrative approach to future systematic and ecological research on Lake Malawi cichlids is stressed. / KMBT_363 / Adobe Acrobat 9.53 Paper Capture Plug-in

More than twenty years ago, over twenty species of the rock-dwelling cichlid species (Mbuna) were translocated from the northern Lake Malawi, where they are endemic, to Thumbi West Island, Lake Malawi National Park, in the southern part of Lake Malawi. Among these species, Cynotilapia afra, Pseudotropheus callainos and Pseudotropheus tropheops 'red cheek' are strongly territorial, and have increased substantially in number and are widely distributed, particularly in the three to seven metre depth band of the rocky habitats at the Island of Thumbi West. It is feared that the increase in population density of translocated species (hereafter referred to as introduced species) may be at the expense of ecologically equivalent native species which could be eliminated. In this thesis the following key hypotheses have been tested: (i) that the introduced species having originated from a region of Lake Malawi which is generally poor in nutrients and introduced in an area which is richer in nutrients, would cope better than the native species during periods of nutrient scarcity which occur frequently, often seasonally in oligotrophic lakes, such as Lake Malawi; (ii) that the introduced species are fitter than their ecologically equivalent native species in the acquisition of territorial space in which they breed, feed and seek shelter, and (iii) that introduced and native species coexist by utilizing different microhabitats. Results show that: 1. the introduced species, P. callainos and P. tropheops 'red cheek' may have responded positively to enhanced nutrient availability, as they were found to have better condition factors and fecundity indices at Thumbi West Island than at sites of their origin, in the northern lake Malawi. Cynotilapia afra, P. callainos and P. tropheops 'red cheek' also maximise their life-span fecundity by starting to reproduce at relatively smaller size than the native species with which they overlap in microhabitat requirements. Similarly, their breeding peaks precede the breeding peaks of the native species with which they overlap in microhabitat requirements. Consequently, due to priority residence effects, the offspring of introduced species may have a competitive edge in the use of essential resources, e.g., refuge over the offspring of the native species whose peak-recruitment occurs later in the year. 2. There is an overlap between the introduced and native species in their microhabitat requirements. Consequently, interference competition between them for territorial sites occurs. The choice of optimal territory sites is constrained by the fact that females preferentially mate with males that defend significantly smaller holes, or crevices among the rocks, probably as a means of minimizing egg predation during spawning. 3. The population of territorial males of introduced species seems to grow exponentially, depending on the availability of suitable microhabitats, and an equilibrium between them and males of the native species may be reached. Competition for optimal territory sites seems to intensify, once the carrying capacity in a particular area has been reached, and it is at this stage that some territorial males of the introduced and native species with similar microhabitat requirements, e.g., C. afra and P. zebra, or P. tropheops 'red cheek' and its sibling native species, P. tropheops 'orange chest' displace each other. However, it seems unlikely that any of the native species which were compared with the introduced species would be driven to extinction because: (a) there is a considerable interspecific territory turn-over between the introduced and native species that overlap in microhabitat requirements. (b) Even in situations where some of the native species occur in microhabitats that are not of their preference, they occupy patches of suitable sites and are capable of breeding. (c) It has been suggested that since introduced and native species breed throughout the year and are polygamous and have intraspecifically shared paternity, they are capable of fertilizing many gravid females of their own species. Therefore, the population of native species may not be detrimentally limited by the presence of introduced species. (d) The introduced and native Mbuna species that prefer small rocks coexist in the same microhabitats, partly by feeding at different sites with different intensity and they also feed at different heights in the water column. 4. The following studies have been recommended before any management intervention, such as culling is adopted: (i). interaction between the introduced and native species in the shallow and deep rocky habitats; (ii) space utilization and survivorship of juveniles of the introduced and native species; (iii) laboratory studies to confirm the role of different nutrient regimes on the fecundity of Mbuna; (iv) the possibility of hybridization between the introduced and native species; (v) monitoring of population growth and distribution of the introduced species around Thumbi West Island should continue in order to detect their long-term effects on the native species.

Behaviour and space utilisation of rock-dwelling cichlids were observed at Thumbi East Island, Lake Malawi. 1. Males of five species of the mbuna complex held long-term territories. Pseudotropheus elongatus "aggressive" vigorously defended a feeding area and sometimes a spawning site interspecifically, but did not feed in the peripheral part of their territory. Spawning sites of Pseudotropheus zebra, Pseudotropheus tropheops "orange chest", Labeotropheus fuelleborni, and Petrotilapia nigra were interspecifically-defended, while larger mating territories were defended against conspecific neighbours. Feeding areas were shared with many fish and often extended beyond the defended area. There was considerable variation in behaviour and space use within and between species and between times of day. 2. Non-territorial P. zebra used larger ranges than territorial conspecifics, and fed more on plankton, but individuals had preferred benthic feeding areas, often in conspecific territories. These 'floaters' were often aggressive. Both size and relative brightness independently predicted the outcome of aggressive interactions between floaters, and a site-specific dominance hierarchy was suggested, with some individuals appearing to be semi-territorial. 3. Males and females of 21 and 13 species respectively were found to establish temporary breeding territories. Overall breeding seasonality was bimodal, but reproductive timing and territory characteristics differed among species. Temporary territories had a considerable impact on the behaviour and habitat use of all resident mbuna species, even causing abandonment of territories. 4. Non-breeding Protomelas taeniolatus had limited home ranges, and showed little aggression. During the highly-synchronised reproductive season, males defended spawning sites and females fry-guarding territories. Most chases were directed towards the commonest fish, but predators were chased further and faster. Female behaviour changed over the guarding period. Females generally continued territorial defence after the brood had disappeared. Most broods contained fry of different sizes and species. Significant benefits were found for guarding females with clustered territories, but females did not appear to choose sites adjacent to conspecific parental females. 5. Territoriality in fish is taxonomically widespread and may serve several functions according to species, sex and developmental stage. It also varies according to genotype- and phenotype-limited strategies and short term costs and benefits. Territories may be simultaneously multifunctional.

Diplotaxodon limnothrissa Turner (1995) is a widely distributed species occurring throughout Lake Malawi, extending from the surface to a depth of at least 220m. It is probably the most abundant cichlid in the lake with biomass estimates of around 87 000 tonnes in the pelagic zone alone. The species is exploited commercially in the southern part of the lake but since its inception the fishery has never been assessed. As such this study investigates some aspects of age and growth of the species besides applying a yield-per-recruit analysis to assess the status of the fishery. Analysis of sectioned sagittal otoliths revealed that D. limnothrissa is fast growing and relatively long-lived species, attaining ages in excess of 10 years. Growth in length was rapid in immature fish, with fish attaining almost half of their maximum size within their first year. Le ngth-at-age was described by the von Bertalanffy growth model with combined-sex growth described as Lt = 211.21(1-exp(- 0.24(t+1.36))) mm TL. Total, natural and fishing mortalities were estimated at 0.76 yr⁻¹, 0.31 yr⁻¹ and 0.45 yr⁻¹ respectively. Per-recruit analysis indicated that the D. limnothrissa stock in the southeast arm of the lake is fully exploited as indicated by the current spawner biomass-per-recruit ratios of 31-55% (SB/R)F=0. Modelling indicated that the current age-at-capture (2.67 years) is lower than the age at which yield is optimised (> 5 years) based on the F₀·₁ harvesting strategy. It is, therefore, recommended that the age-at-capture should be increased from 2.67 to 5 years to optimise yield.

This study centres on a diverse assemblage of epilithic algal grazing fish (17 cichlids and 2 cyprinids) found on the rocky shores near Monkey Bay, Lake Malawi. Diet overlap among these species is large, but competitive exclusion is apparently avoided by the species having different feeding structures, behaviour and distribution on the shore. In many respects the ecology and behaviour of the cichlid fishes inhabiting the rocky zones of Lake Malawi parallel those of coral reef fish. In both communities there are species which are highly territorial, resulting in resource subdivision and, in some cases, the development of algal gardens. As this behaviour has not been previously reported for cichlid species, this thesis describes the territoriality of the rock-dwelling cichlids of Monkey Bay, Lake Malawi, and poses three specific questions: 1. What is the function of territoriality? 2. How do the territorial species subdivide the space resource? 3. What prevents one species taking over all the available space? It is suggested that interspecific territoriality evolved in response to a shortage of food, and, at present, serves principally to defend an algal garden i.e. a food supply of high standing crop. The species studied are extremely site specific. The rocky shore is subdivided among the four species according to the rock configuration, size, algal mat composition and position on the shore, of the available sites. An experimental study of the space utilization of the two common territorial species, referred to as 'Pseudotropheus orange cheek' and 'Pseudotropheus aggressive', showed that suitable territory sites are in short supply and that these species are in competition for some sites. These results do not support those of Fryer (1959) who suggested that a superabundance of food enabled fish to coexist by sharing the same food resource. Indeed it now appears that a shortage of food may have been a major driving force during the recent evolution of the species studied.

In Malawi, fish contribute about 60-80% to the country's animal protein supply. The greater proportion (> 50%) comes from Lake Malawi. Bathyclarias nyasensis and other clariid catfish contribute up to > 20% of the total catches. Catches of Bathyclarias nyasensis in the inshore area of the south-east arm of Lake Malawi are declining and a management plan for the fishery is essentially lacking. There is paucity of biological data that precludes the use of any option to manage the species. The principal aim of the thesis was to define the ecological role B. nyasensis, the most abundant and common of the Bathyclarias species. By examining life history characteristics within a food web context, it was hypothesized that the study would provide an insight into the interrelationships between species, and, hence form the basis for the development of a rational exploitation strategy for the species. The study was undertaken in the south-east arm of Lake Malawi (9° 30'S, 14° 30'S). The principal objectives of the study were to investigate the feeding ecology of B. nyasensis by examining morphological characters and structures associated with feeding, diet of B. nyasensis, food assimilated in the species using carbon (∂¹³C) isotope analysis, daily food consumption rate for B. nyasensis; and to relate the feeding ecology to life history traits such as age, growth, and some aspects of the reproductive biology of B. nyasensis. The suitability of sectioned pectoral spines and sagittal otoliths to age B. nyasensis was assessed. Due to reabsorption of growth zones with increasing spine lumen diameter with fish size, and the relatively low number of spines that could be aged reliably, only otoliths were used. The maximum age for B. nyasensis was estimated at 14 vears. Growth was best was described by the four parameter Schnute mc: lt ={42+(81¹·⁸ - 42¹·⁸)x1-e⁻°·°⁵⁽t⁻¹⁾}¹/¹·⁸ over 1-e⁻⁽⁻°·°⁵⁾⁽¹¹⁾ for female, lt={41+(98¹·² - 41¹·²)x 1-e⁻°·°²⁽t⁻¹⁾}¹/¹·² over 1-e⁻⁽⁻°·°²⁾⁽¹³⁾ and for male fish. Age-at-50% maturity for females and males were estimated at 7 years and 4 years, respectively. Typically, fish grew rapidly in the first year, but slower during subsequent years. Smaller fish were found inshore while larger fish were found in offshore regions. It was hypothesised that the rapid growth in the first year and slower growth later is a consequence of change in diet from high quality and abundant food source to a more dilute food and that this may be associated with a shift in habitat. Morphological characters associated with feeding were used to predict the food and feeding behaviour of B. nyasensis. The size of premaxillary, vomerine, pharyngeal dental and palatine teeth and premaxillary and vomerine tooth plates suggested the capability of B. nyasensis to handle both large and small prey, with a propensity towards smaller prey in composition to C. gariepinus. The molariform teeth on the vomerine tooth plate suggested that molluscs form part of the diet. The relative gut length (1.27±0.24) suggested omnivory, with an ability to switch between planktivory and piscivory. Buccal cavity volume and filtering area changed with fish size at 500-600 mm TL upon which it was hypothesised that the fish diet changed to planktivory at this size. Detailed diet analysis provided information upon which the above hypotheses could be accepted. Percent Index of Relative Importance (%IRI) and a multi-way contingency table analysis based on log-linear models were used to analyse diet data. Results showed that B. nyasensis is omnivorous, but with a distinct ontogenetic dietary shift from piscivory to zooplanktivory at 500 - 600 mm TL. The increased buccal cavity volume at the same fish size therefore, suggests that B. nyasensis is well adapted to filter the dilute zooplankton resource. Increased foraging costs of feeding on zooplankton explained the slower growth of larger fish. The dietary shift was finally corroborated by results of the ∂¹³C isotope analysis. A polynomial equation described the change in carbon ratios with fish size: ∂¹³C = - 33.188 + 0.4997L - 0.0045 (total length)² (r² = 0.598, n = 12, p=0.022). The ontogenetic shift in diet was synchronised with a habitat shift postulated in life history studies. In the inshore region, B. nyasensis were predominantly piscivorous (apex predators), and were zooplanktivorous in the offshore region, thereby forming part of the pelagic food web in the latter region. After examining "bottom-up" and trophic cascade theories, it was postulated that perturbations of the B. nyasensis stock would be discernible both at the top and lower trophic levels. As a piscivore and therefore apex predator, effects of overfishing B. nyasensis in the inshore region could cascade to unpredictable ecological changes in inshore areas and, due to the ontogenetic habitat shift, in the offshore regions. Examples of trophic cascade phenomena are provided. On the basis of the feeding study, it was possible to reconstruct the pelagic food web of Lake Malawi. Apart from the lakefly Chaoborus edulis, B. nyasensis is the other predator that preys heavily on zooplankton in the pelagic zone. Perturbations of the B. nyasensis stock could affect size composition of zooplankton which in tum, could affect production of C. edulis, a resource for the top predators in the food web. The findings of the present study contributed to the ongoing debate of introducing a zooplanktivore into the pelagic zone of Lake Malawi. Proponents for the introductions have argued that zooplankton predation by fish is inferior to that of C. edulis. Introduction of a clupeid zooplankton was proposed as a strategy to boost fish production in the lake. The zooplanktivore would either out-compete or prey on C. edulis to extinction. Opponents to this view argued that zooplankton biomass in the pelagic region was too low to support introductions and that the fish biomass in the pelagic region may have been underestimated. Results from the present study suggest that planktivorous fish (including B. nyasensis) might not be inferior to C. edulis in utilising the zooplankton resource; B. nyasensis is well adapted to utilise the dilute zooplankton resource, and by omitting B. nyasensis from previous studies, overall zooplankton predation by fish may have been underestimated by between 7 - 33%. On the basis of the theoretical migratory life history cycle of B. nyasensis, it is recommended that the current interest in increasing fishing effort in offshore areas should proceed with caution. Ecological changes that may have occurred in the inshore areas due to overfishing have probably not been noticed: as the offshore zone has never been fished. The latter zone may have acted as a stock refuge area. Higher fishing intensity in the offshore areas could lead to serious ecological imbalances and instability. The study has shown that life history characteristics studied in the context of the food web, and in the absence of other fisheries information and/or data, strongly advocates the precautionary principle to managing changes in exploitation patterns.

The intensive tank culture of ornamental mouthbrooding cichlids poses several problems which limit their aquaculture potential. This project addressed some of these problems for Aulonocara baenschi. The production of juveniles in 2501 aquaria was accelerated when: a) aquaria were equipped with refuges, b) females of less than 70mm in standard length were used as broodstock, c) mouthbrooding females were replaced with gravid females at seven day intervals, and d) embryos were removed from the mouths of females at replacement times for artificial incubation. Two sex ratios also accelerated juvenile production. The sex ratio (male:females) 1:30 yielded the highest spawning returns per tank, and therefore represented the most effective utilization of aquarium space (a critical consideration for the small-scale culturist). Contrastingly, the sex ratio 1:12 yielded the highest clutch sizes and a high percentage female spawning return, and therefore represented the most effective utilization of broodstock (an important consideration for culturists inhibited by financial constraints or having an abundance of culture vessels). The reproductive behaviour of A. baenschi was described. Emphasis was given to aspects of reproduction of relevance to culture, for example; spawning times and seasons, clutch size and its relationship with female size, age and size of sexes at first spawning, embryo development rate and size of first swimming juveniles. The slow growth rate of juveniles, combined with a late attainment of marketable size (± seven months) was a major limitation affecting the cuIture potential of A. baenschi. Two factors favouring the cuIture of this species was the high survival rate recorded for both adults and juveniles, and the comparatively high prices fetched by fish on domestic wholesale markets (R4,00 per fish). It is recommended that A. baenschi should not be cultured exclusively for the relatively small South African ornamental fish market. A more profitable strategy for domestic culturists should involve a major production effort with A. baenschi and other desirable species of Aulonocara (e.g . A. ethelwynnae; A. hansbaenschi; A. stuartgranti & A. maylandi) for foreign markets (in particular, the U.S.A.; Western Europe & Japan). Not only are these markets massive, but prevailing exchange rates of the Rand with these currencies favour such a strategy.

This thesis analyzes the phonological and morphological nativisation of English loans in the Tonga language. The contact situation between English and Tonga, in Zimbabwe, facilitates transference of lexical items between the two languages. From having been one of the most widely used languages of the world, English has developed into the most influential donor of words to other languages such as Tonga. The infiltration of English words into the Tonga lexical inventory led to the adoption and subsequent nativisation of English words by the native Tonga speakers. The main deposit of English words into Tonga is the direct interaction between English and Tonga speakers. However, it is sometimes via other languages like Shona, Ndebele, Venda and Shangani. In the 21st century, English’s contribution to the vocabulary of Tonga became more widely spread, now covering a large proportion of the Tonga language’s lexical inventory. The fact that English is the medium of instruction, in Zimbabwe, language of technology, education, media, new administration, health, music, new religion and economic transactions means that it is regarded as the high variety language with coercive loaning powers. Words from English are then adopted and nativised in the Tonga language, since Tonga asserts itself an independent language that can handle loans on its own. The main focus of this study therefore, is to try and account for the phonological and morphological behavior and changes that take place in English words that enter into Tonga. Analyzing phonological processes that are employed during nativisation of loan words entails analyzing how Tonga speakers handle aspects of English language such as diphthongs, triphthongs, cluster consonants, CVC syllable structure and sounds in repairing unacceptable sequences in Tonga. The research also accounts for the handling of morphological differences between the two languages. This entails looking at how competence and ordered-rule framework are harmonized by Tonga speakers in repairing conflicting features at morphological level. Since the two languages have different morphological patterns, the research analyzes the repairing strategies to handle singular and plural noun prefixes, tenses and particles, which are morphological components of words. The researcher appreciates that the native Tonga speakers have robust intuitions on the proper way to nativise words. / African Languages / D.Litt. et Phil. (African Languages)

This thesis analyzes the phonological and morphological nativisation of English loans in the Tonga language. The contact situation between English and Tonga, in Zimbabwe, facilitates transference of lexical items between the two languages. From having been one of the most widely used languages of the world, English has developed into the most influential donor of words to other languages such as Tonga. The infiltration of English words into the Tonga lexical inventory led to the adoption and subsequent nativisation of English words by the native Tonga speakers. The main deposit of English words into Tonga is the direct interaction between English and Tonga speakers. However, it is sometimes via other languages like Shona, Ndebele, Venda and Shangani. In the 21st century, English’s contribution to the vocabulary of Tonga became more widely spread, now covering a large proportion of the Tonga language’s lexical inventory. The fact that English is the medium of instruction, in Zimbabwe, language of technology, education, media, new administration, health, music, new religion and economic transactions means that it is regarded as the high variety language with coercive loaning powers. Words from English are then adopted and nativised in the Tonga language, since Tonga asserts itself an independent language that can handle loans on its own. The main focus of this study therefore, is to try and account for the phonological and morphological behavior and changes that take place in English words that enter into Tonga. Analyzing phonological processes that are employed during nativisation of loan words entails analyzing how Tonga speakers handle aspects of English language such as diphthongs, triphthongs, cluster consonants, CVC syllable structure and sounds in repairing unacceptable sequences in Tonga. The research also accounts for the handling of morphological differences between the two languages. This entails looking at how competence and ordered-rule framework are harmonized by Tonga speakers in repairing conflicting features at morphological level. Since the two languages have different morphological patterns, the research analyzes the repairing strategies to handle singular and plural noun prefixes, tenses and particles, which are morphological components of words. The researcher appreciates that the native Tonga speakers have robust intuitions on the proper way to nativise words. / African Languages / D.Litt. et Phil. (African Languages)