A study of the Pycnonotus bulbul species complex in Southern Africa
- Authors: Lloyd, Penn
- Date: 1995
- Subjects: Bulbuls -- Ecology -- South Africa -- Eastern Cape , Songbirds -- Ecology -- South Africa -- Eastern Cape , Birds -- Ecology -- South Africa -- Eastern Cape
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5782 , http://hdl.handle.net/10962/d1005470 , Bulbuls -- Ecology -- South Africa -- Eastern Cape , Songbirds -- Ecology -- South Africa -- Eastern Cape , Birds -- Ecology -- South Africa -- Eastern Cape
- Description: The three Pycnonotus bulbuls endemic to Africa, P. barbatus, P. nigricans and P. capensis, occupy mutually exclusive distributions in southern Africa. These species are closely related and appear to occupy very similar ecological niches, only in different regions. Using a multifaceted approach, this study attempts to explain the ecology of this species complex. All three species show similar physiological responses to temperature extremes, and are therefore unlikely to be directly limited by environmental temperature. However, their distributions are highly correlated to a complex of environmental variables, particularly winter minimum temperatures, the coefficient of variation in mean annual rainfall, and the seasonality of rainfall. This combination of environmental parameters can be used to predict the distributions of at least one of the species, P. nigricans, accurately. An analysis of the vocalizations and behaviour of the three species revealed that, whilst P. capensis has a number of recognizably different vocalizations, those of P. barbatus and P. nigricans are very similar. The three have nearly identical behaviours, particularly courtship and pre-copulatory behaviours. The mate recognition systems of the three are therefore extremely similar. P. barbatus is territorial during the breeding season, and exhibits highly structured-variation in male territorial song at the level of the local neighbourhood. The literature pertaining to song dialects is reviewed, and a new hypothesis is postulated to explain song-matching in terms of neighbour/stranger discrimination and the possible existence of cooperative territory defence. A survey of the eastern Cape region, where all three species come into contact, showed that extensive hybridization is taking place between each species pair. Phenotypically, this hybridization is restricted to narrow hybrid zones, that are considered to be stable in both time and place. The evolutionary and ecological significance of these zones to the distributions of the species is discussed, and it is proposed that the zones are maintained by selection acting on differentially-adapted genomes along an environmental gradient.
- Full Text:
- Date Issued: 1995
- Authors: Lloyd, Penn
- Date: 1995
- Subjects: Bulbuls -- Ecology -- South Africa -- Eastern Cape , Songbirds -- Ecology -- South Africa -- Eastern Cape , Birds -- Ecology -- South Africa -- Eastern Cape
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5782 , http://hdl.handle.net/10962/d1005470 , Bulbuls -- Ecology -- South Africa -- Eastern Cape , Songbirds -- Ecology -- South Africa -- Eastern Cape , Birds -- Ecology -- South Africa -- Eastern Cape
- Description: The three Pycnonotus bulbuls endemic to Africa, P. barbatus, P. nigricans and P. capensis, occupy mutually exclusive distributions in southern Africa. These species are closely related and appear to occupy very similar ecological niches, only in different regions. Using a multifaceted approach, this study attempts to explain the ecology of this species complex. All three species show similar physiological responses to temperature extremes, and are therefore unlikely to be directly limited by environmental temperature. However, their distributions are highly correlated to a complex of environmental variables, particularly winter minimum temperatures, the coefficient of variation in mean annual rainfall, and the seasonality of rainfall. This combination of environmental parameters can be used to predict the distributions of at least one of the species, P. nigricans, accurately. An analysis of the vocalizations and behaviour of the three species revealed that, whilst P. capensis has a number of recognizably different vocalizations, those of P. barbatus and P. nigricans are very similar. The three have nearly identical behaviours, particularly courtship and pre-copulatory behaviours. The mate recognition systems of the three are therefore extremely similar. P. barbatus is territorial during the breeding season, and exhibits highly structured-variation in male territorial song at the level of the local neighbourhood. The literature pertaining to song dialects is reviewed, and a new hypothesis is postulated to explain song-matching in terms of neighbour/stranger discrimination and the possible existence of cooperative territory defence. A survey of the eastern Cape region, where all three species come into contact, showed that extensive hybridization is taking place between each species pair. Phenotypically, this hybridization is restricted to narrow hybrid zones, that are considered to be stable in both time and place. The evolutionary and ecological significance of these zones to the distributions of the species is discussed, and it is proposed that the zones are maintained by selection acting on differentially-adapted genomes along an environmental gradient.
- Full Text:
- Date Issued: 1995
Complex interactions involving the Cape fig, Ficus sur Forsskål, and its associated insects
- Authors: Zachariades, Costas
- Date: 1995
- Subjects: Insect-plant relationships Mutualism (Biology) Fig -- South Africa Fig wasp -- South Africa Ants Homoptera
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5655 , http://hdl.handle.net/10962/d1005338
- Description: The inadequacy of arbitrarily classifying interactions between species as antagonistic, neutral or mutualistic has become clear in recent years. Both direct and indirect interactions between species can vary between mutualism and antagonism, depending on the intrinsic and extrinsic contexts of the interaction. This study investigated the characteristics of an ant-plant-homopteran interaction in southern Africa. The polyphagous homopteran Hilda patruelis (Tettigometridae) feeds primarily on the trunk-borne fruiting branches and figs of the Cape fig tree, Ficus sur, and produces honeydew which attracts tending ants. Ten of the sixteen ant species/species groups present on F. sur tended H. patruelis, with Pheidole megacephala the most frequent attendant. Ants attracted to F. sur by H. patruelis honeydew or other liquid food sources also preyed on insects on the tree, including adults of the small agaonid fig wasps whose larvae feed on the ovules in the developing figs. One fig wasp species (Ceratosolen capensis) is also the tree's only pollinator. No benefits to H. patruelis from being tended by ants were detected, either in terms of reduced parasitism, or predation by a lycaenid caterpillar. A P. megacephala colony foraging on a F. sur tree was found to receive a high proportion of its likely energy requirements from the tree, mainly in the form of H. patruelis honeydew, during periods when it was bearing fruit. It is probable that the H. patruelis-P. megacephala interaction constitutes a direct mutualism at times, but that benefits to the homopteran are intermittent or weak. Both H. patruelis and ants benefitted from F. sur, directly or indirectly, through the provision of food (and for some ants, nesting sites). The removal of phloem sap by H. patruelis did not detectably reduce the trees' reproductive output, either in terms of pollinator or viable seed production. The indirect effects of ant and H. patruelis presence on the F. sur trees were on average positive, as ants preyed disproportionately heavily on fig wasp species parasitic on or competing with the pollinator, thus increasing pollinator production. Effects of ant presence on seed production were not investigated, but have been demonstrated as beneficial elsewhere. However, there is great varatlon both in the composition of the wasp fauna arriving to oviposit at different crops, and in ant densities per fig, on several temporal and spatial scales. This results in high variability in the effects of ants on the pollinator and, through it, the tree, from positive to zero and potentially even negative. Despite this conditionality of beneficial outcomes for the tree, the mean effect of ants on the F. sur population studied was to increase pollinator production by up to nearly 20%. This study is among the few to have demonstrated an overall benefit to a plant of having homopteran-tending ants present on it.
- Full Text:
- Date Issued: 1995
- Authors: Zachariades, Costas
- Date: 1995
- Subjects: Insect-plant relationships Mutualism (Biology) Fig -- South Africa Fig wasp -- South Africa Ants Homoptera
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5655 , http://hdl.handle.net/10962/d1005338
- Description: The inadequacy of arbitrarily classifying interactions between species as antagonistic, neutral or mutualistic has become clear in recent years. Both direct and indirect interactions between species can vary between mutualism and antagonism, depending on the intrinsic and extrinsic contexts of the interaction. This study investigated the characteristics of an ant-plant-homopteran interaction in southern Africa. The polyphagous homopteran Hilda patruelis (Tettigometridae) feeds primarily on the trunk-borne fruiting branches and figs of the Cape fig tree, Ficus sur, and produces honeydew which attracts tending ants. Ten of the sixteen ant species/species groups present on F. sur tended H. patruelis, with Pheidole megacephala the most frequent attendant. Ants attracted to F. sur by H. patruelis honeydew or other liquid food sources also preyed on insects on the tree, including adults of the small agaonid fig wasps whose larvae feed on the ovules in the developing figs. One fig wasp species (Ceratosolen capensis) is also the tree's only pollinator. No benefits to H. patruelis from being tended by ants were detected, either in terms of reduced parasitism, or predation by a lycaenid caterpillar. A P. megacephala colony foraging on a F. sur tree was found to receive a high proportion of its likely energy requirements from the tree, mainly in the form of H. patruelis honeydew, during periods when it was bearing fruit. It is probable that the H. patruelis-P. megacephala interaction constitutes a direct mutualism at times, but that benefits to the homopteran are intermittent or weak. Both H. patruelis and ants benefitted from F. sur, directly or indirectly, through the provision of food (and for some ants, nesting sites). The removal of phloem sap by H. patruelis did not detectably reduce the trees' reproductive output, either in terms of pollinator or viable seed production. The indirect effects of ant and H. patruelis presence on the F. sur trees were on average positive, as ants preyed disproportionately heavily on fig wasp species parasitic on or competing with the pollinator, thus increasing pollinator production. Effects of ant presence on seed production were not investigated, but have been demonstrated as beneficial elsewhere. However, there is great varatlon both in the composition of the wasp fauna arriving to oviposit at different crops, and in ant densities per fig, on several temporal and spatial scales. This results in high variability in the effects of ants on the pollinator and, through it, the tree, from positive to zero and potentially even negative. Despite this conditionality of beneficial outcomes for the tree, the mean effect of ants on the F. sur population studied was to increase pollinator production by up to nearly 20%. This study is among the few to have demonstrated an overall benefit to a plant of having homopteran-tending ants present on it.
- Full Text:
- Date Issued: 1995
Constituent processes contributing to stress induced β-carotene accumulation in Dunaliella salina
- Authors: Phillips, Lesley Gail
- Date: 1995
- Subjects: Dunaliella Carotenes Plants -- Effect of stress on
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5658 , http://hdl.handle.net/10962/d1005341
- Description: The alga Dunaliella salina possesses the unique ability to accumulate up to 14 % of it's dry weight as β-carotene in response to stress conditions. This hyper-accumulation of β-carotene has led to the commercial exploitation of this alga for the biotechnological production of this important carotenoid. In order to maximise β-carotene production, a dual-stage process which separates the distinctive growth phase from the β-carotene accumulating stress phase has recently been patented. Preliminary laboratory studies showed that although stress factors such as high salinity and nutrient limitation enhance β-carotene accumulation in D. salina (± 10 pg.cell⁻¹), high light intensity was the single most important factor contributing to the induction of β-carotene accumulation in this alga (± 20 pg.cell⁻¹). Moreover, it was demonstrated that β-carotene accumulation can be further stimulated by exposing the alga to a combination of high light intensity, salt and nutrient stresses (± 30-60 pg.cell⁻¹). The response of D. salina to stress was shown to occur in two phases. The first phase occurred within 24 hours and was characterized most importantly by higher rates of β-carotene accumulation for all the stresses investigated. In cells exposed to multiple stress factors in mass culture, the β-carotene accumulation rate was as much as 9.5 pg.cell⁻¹.day⁻¹ in the first phase compared to only 3 pg.cell·day⁻¹ in the second phase. Since the rate of β-carotene accumulation was higher within the first 24 hours after exposure to stress, the first phase was considered crucial for stress-induced β-carotene accumulation. Characterization of this phase revealed that the stress response was multifaceted. Transition of cells from the growth stage to stress conditions was characterized by the following: 1) Change in cell volume. Hypersalinity caused cell shrinkage while cells exposed to nutrient limitation and/or high light intensity caused cells to swell. Restoration of cell volume was shown to occur within 8 hours for all stresses investigated. 2) Altered photosynthesis. Inhibition of both carbon fixation and oxygen evolution was demonstrated in cells immediately after exposure to multiple stress factors. 3) Production of abscisic acid. Intracellular ABA levels increased within 6-8 hours after exposure to all stresses investigated. The rise in intracellular ABA levels coincided with an increase or return to starting cell volume. High intracellular ABA levels were however transient and within 24 hours ABA began to partition into the culture medium. 4) Change in pigment composition. Changes in xanthophyll cycle pigment content was demonstrated soon after exposure to stress conditions. In hypersalinity shocked cells, initial epoxidation of zeaxanthin to violaxanthin and subsequent de-epoxidation to zeaxanthin occurred, whereas exposure to high stress resulted in an immediate and continued decrease in the epoxidation state indicating accumulation of zeaxanthin. A rapid rate of chlorophyll depletion was also demonstrated. In addition to the above responses a rapid decrease in growth rate during this phase was also noted. An interrelationship between cell volume change, ABA production, maintenance of xanthophyll cycle operation and β-carotene accumulation therefore appeared to exist. ABA production was shown to be stoichiometrically related to changes in xanthophyll content with r² = 0.84 and slope of the curve = 0.91 being achieved for high light stressed cells. This study therefore presents strong circumstantial evidence in support of a carotenoid origin for ABA in Dunaliella. In addition, enhanced β-carotene content was achieved by the application of exogenous ABA and related compounds suggesting a role for ABA as a regulator of the overall stress response. Furthermore, zeaxanthin accumulation was shown to be positively correlated ( r²≥ 0.81) to β-carotene accumulation for all the stresses investigated. The second phase was characterized by a return to homoeostasis of the physiological processes mentioned above, indicating acclimation of the cell to prevailing conditions. This stage was characterised by continued β-carotene accumulation and a decreased epoxidation state of the xanthophyll cycle which together appeared to sustain photosynthesis, allowing this organism to tolerate stress conditions.
- Full Text:
- Date Issued: 1995
- Authors: Phillips, Lesley Gail
- Date: 1995
- Subjects: Dunaliella Carotenes Plants -- Effect of stress on
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5658 , http://hdl.handle.net/10962/d1005341
- Description: The alga Dunaliella salina possesses the unique ability to accumulate up to 14 % of it's dry weight as β-carotene in response to stress conditions. This hyper-accumulation of β-carotene has led to the commercial exploitation of this alga for the biotechnological production of this important carotenoid. In order to maximise β-carotene production, a dual-stage process which separates the distinctive growth phase from the β-carotene accumulating stress phase has recently been patented. Preliminary laboratory studies showed that although stress factors such as high salinity and nutrient limitation enhance β-carotene accumulation in D. salina (± 10 pg.cell⁻¹), high light intensity was the single most important factor contributing to the induction of β-carotene accumulation in this alga (± 20 pg.cell⁻¹). Moreover, it was demonstrated that β-carotene accumulation can be further stimulated by exposing the alga to a combination of high light intensity, salt and nutrient stresses (± 30-60 pg.cell⁻¹). The response of D. salina to stress was shown to occur in two phases. The first phase occurred within 24 hours and was characterized most importantly by higher rates of β-carotene accumulation for all the stresses investigated. In cells exposed to multiple stress factors in mass culture, the β-carotene accumulation rate was as much as 9.5 pg.cell⁻¹.day⁻¹ in the first phase compared to only 3 pg.cell·day⁻¹ in the second phase. Since the rate of β-carotene accumulation was higher within the first 24 hours after exposure to stress, the first phase was considered crucial for stress-induced β-carotene accumulation. Characterization of this phase revealed that the stress response was multifaceted. Transition of cells from the growth stage to stress conditions was characterized by the following: 1) Change in cell volume. Hypersalinity caused cell shrinkage while cells exposed to nutrient limitation and/or high light intensity caused cells to swell. Restoration of cell volume was shown to occur within 8 hours for all stresses investigated. 2) Altered photosynthesis. Inhibition of both carbon fixation and oxygen evolution was demonstrated in cells immediately after exposure to multiple stress factors. 3) Production of abscisic acid. Intracellular ABA levels increased within 6-8 hours after exposure to all stresses investigated. The rise in intracellular ABA levels coincided with an increase or return to starting cell volume. High intracellular ABA levels were however transient and within 24 hours ABA began to partition into the culture medium. 4) Change in pigment composition. Changes in xanthophyll cycle pigment content was demonstrated soon after exposure to stress conditions. In hypersalinity shocked cells, initial epoxidation of zeaxanthin to violaxanthin and subsequent de-epoxidation to zeaxanthin occurred, whereas exposure to high stress resulted in an immediate and continued decrease in the epoxidation state indicating accumulation of zeaxanthin. A rapid rate of chlorophyll depletion was also demonstrated. In addition to the above responses a rapid decrease in growth rate during this phase was also noted. An interrelationship between cell volume change, ABA production, maintenance of xanthophyll cycle operation and β-carotene accumulation therefore appeared to exist. ABA production was shown to be stoichiometrically related to changes in xanthophyll content with r² = 0.84 and slope of the curve = 0.91 being achieved for high light stressed cells. This study therefore presents strong circumstantial evidence in support of a carotenoid origin for ABA in Dunaliella. In addition, enhanced β-carotene content was achieved by the application of exogenous ABA and related compounds suggesting a role for ABA as a regulator of the overall stress response. Furthermore, zeaxanthin accumulation was shown to be positively correlated ( r²≥ 0.81) to β-carotene accumulation for all the stresses investigated. The second phase was characterized by a return to homoeostasis of the physiological processes mentioned above, indicating acclimation of the cell to prevailing conditions. This stage was characterised by continued β-carotene accumulation and a decreased epoxidation state of the xanthophyll cycle which together appeared to sustain photosynthesis, allowing this organism to tolerate stress conditions.
- Full Text:
- Date Issued: 1995
Dispersal, settlement and recruitment : their influence on the population dynamics of intertidal mussels
- Authors: Phillips, Tracey Elizabeth
- Date: 1995
- Subjects: Perna -- South Africa -- Eastern Cape -- Algoa bay Mytilus galloprovincialis -- South Africa -- Eastern Cape -- Algoa bay Mytilidae -- South Africa -- Eastern Cape -- Algoa bay
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5663 , http://hdl.handle.net/10962/d1005347
- Description: Recruitment of planktonic larvae into sedentary benthic populations regulates the population dynamics of marine invertebrates. The processes controlling recruitment, however, are poorly understood, and recruitment remains largely unpredictable, which complicates management of exploited shellfish resources. The mussels Perna perna, Choromytilus meridionalis and Mytilus galloprovincialis, found on the south coast of southern Africa, have planktonic larvae and sedentary adult stages. This thesis examines dispersal, settlement and early post-settlement growth and mortality, and their effect on recruitment and demography of intertidal mussel populations in the region of Algoa Bay on the south coast of southern Africa. Temporal and spatial variation in the body mass, density and size structure of mussels, the distribution of bivalve larvae on plankton grids in the nearshore zone and the distribution of a recently introduced invasive mussel, Mytilus galloprovincialis, were examined between 1989 and 1992. Furthermore, data on hourly or daily changes in wind strength and direction, air and sea surface temperatures and low and high tide levels in the study region, were obtained. There were 3-4 peaks in spawning (characterised by an abrupt decline in weight) and settlement activity annually. These peaks varied in exact timing, intensity and duration between sites and over time. However, at a site, spawning was followed by settlement 4-8 weeks later, and there was a significant (P < 0.05) direct correlation between spawning intensity prior to the appearance of a new cohort and the cohort density (settlement intensity). The stochastic spatial and temporal variation in breeding activity was superimposed on a more general pattern of a higher intensity of spawning and settlement in Algoa Bay than on the open coast, and a higher settlement intensity on coastal sandstone shores than on dune rock shores. Spawning was more frequent in winter and spring, and the probability of spawning and settlement peaked around the spring and autumn equinox, if temperature and wind conditions were suitable. Larval behaviour had little effect on their dispersal in the well-mixed nearshore region. Larvae were passively dispersed by currents, and their dispersal range and direction depended on prevailing winds and local topography. The sharp decline in density of recruit and adult M. galloprovincialis with increasing distance from the point of introduction, showed that some larvae were carried by wind generated currents over moderately long distances (-100 km). However, since most (76 %) M. galloprovincialis recruited within 4 km of the parent population, it is possible that larvae become trapped in small gullies and crevices around rocky shores, and have a limited dispersal range. This could explain the link between local patterns of spawning and settlement. The distribution and abundance of settlers on the shore was influenced by larval behaviour and the availability of settlement, substrata. Larvae preferred to settle primarily on foliose coralline algae and migrate to the adult mussel bed when they were larger (0.60-7 mm), but larvae also settled directly on adult mussels, possibly because the amount of coralline algae was limited. Both direct and secondary settlement were considered to be important in maintaining mussel populations since the rate of settlement was low(generally < 60 000.m-2). Cohort analyses showed that prior to maturity post-settlement growth (- 30 mm in 10 months) and mortality rates (60-100%) were high, but varied. When settlement intensity was low this variability uncoupled the relationship between spawning and recruitment intensity. Multiple regression analysis showed that together reproductive effort (gamete output), settlement intensity, growth and mortality prior to maturity, accounted for 76 % of the variance in recruitment into mature adult populations. The low settlement rate coupled with the short life span of mussels « 3 years), meant that populations underwent marked spatial and temporal variations in structure and abundance as settlement intensity varied, but there were consistent general differences between mussel populations on dune rock and sandstone shores in Algoa Bay and on the open coast. It was concluded from these results that, spawning intensity and post-settlement growth and mortality, rather than dispersal, regulated recruitment and the structure and abundance of intertidal P. perna and C. meridionalis populations along the south coast of southern Africa. On the basis of these results it is recommended that species with limited dispersal, variable recruitment and high natural mortality, such as P. perna, should be conserved by protecting a small part of the population in reserves, and controlling utilisation outside reserves to minimize disturbance to local brood stocks. Furthermore, since the potential for reseeding adjacent exploited areas is limited, several small reserves placed at regular intervals along the coast would be more effective than a single large reserve.
- Full Text:
- Date Issued: 1995
- Authors: Phillips, Tracey Elizabeth
- Date: 1995
- Subjects: Perna -- South Africa -- Eastern Cape -- Algoa bay Mytilus galloprovincialis -- South Africa -- Eastern Cape -- Algoa bay Mytilidae -- South Africa -- Eastern Cape -- Algoa bay
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5663 , http://hdl.handle.net/10962/d1005347
- Description: Recruitment of planktonic larvae into sedentary benthic populations regulates the population dynamics of marine invertebrates. The processes controlling recruitment, however, are poorly understood, and recruitment remains largely unpredictable, which complicates management of exploited shellfish resources. The mussels Perna perna, Choromytilus meridionalis and Mytilus galloprovincialis, found on the south coast of southern Africa, have planktonic larvae and sedentary adult stages. This thesis examines dispersal, settlement and early post-settlement growth and mortality, and their effect on recruitment and demography of intertidal mussel populations in the region of Algoa Bay on the south coast of southern Africa. Temporal and spatial variation in the body mass, density and size structure of mussels, the distribution of bivalve larvae on plankton grids in the nearshore zone and the distribution of a recently introduced invasive mussel, Mytilus galloprovincialis, were examined between 1989 and 1992. Furthermore, data on hourly or daily changes in wind strength and direction, air and sea surface temperatures and low and high tide levels in the study region, were obtained. There were 3-4 peaks in spawning (characterised by an abrupt decline in weight) and settlement activity annually. These peaks varied in exact timing, intensity and duration between sites and over time. However, at a site, spawning was followed by settlement 4-8 weeks later, and there was a significant (P < 0.05) direct correlation between spawning intensity prior to the appearance of a new cohort and the cohort density (settlement intensity). The stochastic spatial and temporal variation in breeding activity was superimposed on a more general pattern of a higher intensity of spawning and settlement in Algoa Bay than on the open coast, and a higher settlement intensity on coastal sandstone shores than on dune rock shores. Spawning was more frequent in winter and spring, and the probability of spawning and settlement peaked around the spring and autumn equinox, if temperature and wind conditions were suitable. Larval behaviour had little effect on their dispersal in the well-mixed nearshore region. Larvae were passively dispersed by currents, and their dispersal range and direction depended on prevailing winds and local topography. The sharp decline in density of recruit and adult M. galloprovincialis with increasing distance from the point of introduction, showed that some larvae were carried by wind generated currents over moderately long distances (-100 km). However, since most (76 %) M. galloprovincialis recruited within 4 km of the parent population, it is possible that larvae become trapped in small gullies and crevices around rocky shores, and have a limited dispersal range. This could explain the link between local patterns of spawning and settlement. The distribution and abundance of settlers on the shore was influenced by larval behaviour and the availability of settlement, substrata. Larvae preferred to settle primarily on foliose coralline algae and migrate to the adult mussel bed when they were larger (0.60-7 mm), but larvae also settled directly on adult mussels, possibly because the amount of coralline algae was limited. Both direct and secondary settlement were considered to be important in maintaining mussel populations since the rate of settlement was low(generally < 60 000.m-2). Cohort analyses showed that prior to maturity post-settlement growth (- 30 mm in 10 months) and mortality rates (60-100%) were high, but varied. When settlement intensity was low this variability uncoupled the relationship between spawning and recruitment intensity. Multiple regression analysis showed that together reproductive effort (gamete output), settlement intensity, growth and mortality prior to maturity, accounted for 76 % of the variance in recruitment into mature adult populations. The low settlement rate coupled with the short life span of mussels « 3 years), meant that populations underwent marked spatial and temporal variations in structure and abundance as settlement intensity varied, but there were consistent general differences between mussel populations on dune rock and sandstone shores in Algoa Bay and on the open coast. It was concluded from these results that, spawning intensity and post-settlement growth and mortality, rather than dispersal, regulated recruitment and the structure and abundance of intertidal P. perna and C. meridionalis populations along the south coast of southern Africa. On the basis of these results it is recommended that species with limited dispersal, variable recruitment and high natural mortality, such as P. perna, should be conserved by protecting a small part of the population in reserves, and controlling utilisation outside reserves to minimize disturbance to local brood stocks. Furthermore, since the potential for reseeding adjacent exploited areas is limited, several small reserves placed at regular intervals along the coast would be more effective than a single large reserve.
- Full Text:
- Date Issued: 1995
Evaluation of Gratiana spadicea (Klug, 1829) and Metriona elatior (Klug, 1829) (Chrysomelidae: Cassidinae) for the biological control of sticky nightshade Solanum sisymbriifolium Lamarck (Solanaceae) in South Africa
- Authors: Hill, Martin Patrick
- Date: 1995
- Subjects: Solanaceae Solanaceae -- Biological control Chrysomelidae Chrysomelidae -- Host plants Cassida
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5688 , http://hdl.handle.net/10962/d1005374 , https://orcid.org/0000-0003-0579-5298
- Description: Solanum sisymbriifolium (sticky nightshade) is a shrubby weed of South American origin that was introduced to South Africa at the turn of the century. Despite being indicative of disturbed habitats, the weed was found to be invasive in conservation, agricultural recreational and suburban areas; this, coupled with the failure of both chemical and mechanical control attempts suggested that the weed was a good candidate for biological control. A biological control programme which followed a standard protocol was initiated. Observations suggested that S. sisymbriifolium dispersed primarily by seeds. Plants produced large quantities of fleshy fruit, favoured by frugivorous birds, which facilitated the rapid spread of the weed into new habitats. The seeds germinated quickly, especially in disturbed soil, often below the parent plant where they dropped from burst fruit, and along fences where birds roost. The pre-introductory survey of the weed revealed that S. sisymbriifolium was attacked by a relatively small number of, mainly polyphagous, herbivorous insects. These were localised and sporadic in incidence and inflicted very little observable damage. The herbivore fauna of S. sisymbriifolium was depauperate even in relation to two other exotic weeds, S. elaeaglllfolium and S. mauritianum, in South Africa. The paucity of native herbivores on S. sisymbriifoliwn was ascribed to a combination of the weed's taxonomic distinctness from South African Solanum species, and the dense covering of glandular trichomes on its leaves. Although it was shown that the exudate produced by these glandular trichomes of S. sisymbriifolium seriously impeded the movement and feeding of native herbivores, there was not enough evidence to suggest that the glandular trichomes, alone could have been responsible the lack of herbivores on the weed. Two leaf-feeding Cassidinae Gratiana spadicea and MetJ-iona elatior were screened as agents for the biological control of S. sisymbrilfolium. Favourable biological characteristics for both species included a high rate of increase, long-lived adults, many generations per year, and a high per capita feeding rate. Host range was investigated in larval survival tests and adult choice tests. The larvae of both species were reared through to the adult stage on several of the native Solanum species tested, and also on eggplant (S. melongena). However, the survival of G. spadicea on the majority of these species was very low, suggesting that the beetles would be unlikely to attack them in the field. This was supported by the adult choice tests, where G. spadicea females displayed, a strong oviposition preference for their natural host. In contrast, M. elatior larvae survived well on non-host plants, and the females selected several non-host species, including eggplant for oviposition. It was argued that the conflict of interests involving eggplant was overrated because eggplant is subjected to a stringent insecticide spray regime. Based on this evidence, permission for release was granted for G. spadicea. The impact of native parasitoid host range extensions to weed biological control agents in South Africa was investigated. Native parasitoids were recorded from nearly half of the agent species that had established on their target weed. The level of concealment and taxon influenced susceptibility of the agents to parasitoid attack Poorly concealed endophagous agents were most susceptible to attack, while exposed feeders were fairly free from attack. However, native parasitoids were reported not to strongly influence weed biological control agent populations and it was concluded that no agent should be rejected based only on its susceptibility to native parasitoid attack. Finally, several predictions are made as to the potential success of G. spadicea on S. sisymbriifolium and some of the challenges facing the biological control of weeds are discussed.
- Full Text:
- Date Issued: 1995
- Authors: Hill, Martin Patrick
- Date: 1995
- Subjects: Solanaceae Solanaceae -- Biological control Chrysomelidae Chrysomelidae -- Host plants Cassida
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5688 , http://hdl.handle.net/10962/d1005374 , https://orcid.org/0000-0003-0579-5298
- Description: Solanum sisymbriifolium (sticky nightshade) is a shrubby weed of South American origin that was introduced to South Africa at the turn of the century. Despite being indicative of disturbed habitats, the weed was found to be invasive in conservation, agricultural recreational and suburban areas; this, coupled with the failure of both chemical and mechanical control attempts suggested that the weed was a good candidate for biological control. A biological control programme which followed a standard protocol was initiated. Observations suggested that S. sisymbriifolium dispersed primarily by seeds. Plants produced large quantities of fleshy fruit, favoured by frugivorous birds, which facilitated the rapid spread of the weed into new habitats. The seeds germinated quickly, especially in disturbed soil, often below the parent plant where they dropped from burst fruit, and along fences where birds roost. The pre-introductory survey of the weed revealed that S. sisymbriifolium was attacked by a relatively small number of, mainly polyphagous, herbivorous insects. These were localised and sporadic in incidence and inflicted very little observable damage. The herbivore fauna of S. sisymbriifolium was depauperate even in relation to two other exotic weeds, S. elaeaglllfolium and S. mauritianum, in South Africa. The paucity of native herbivores on S. sisymbriifoliwn was ascribed to a combination of the weed's taxonomic distinctness from South African Solanum species, and the dense covering of glandular trichomes on its leaves. Although it was shown that the exudate produced by these glandular trichomes of S. sisymbriifolium seriously impeded the movement and feeding of native herbivores, there was not enough evidence to suggest that the glandular trichomes, alone could have been responsible the lack of herbivores on the weed. Two leaf-feeding Cassidinae Gratiana spadicea and MetJ-iona elatior were screened as agents for the biological control of S. sisymbrilfolium. Favourable biological characteristics for both species included a high rate of increase, long-lived adults, many generations per year, and a high per capita feeding rate. Host range was investigated in larval survival tests and adult choice tests. The larvae of both species were reared through to the adult stage on several of the native Solanum species tested, and also on eggplant (S. melongena). However, the survival of G. spadicea on the majority of these species was very low, suggesting that the beetles would be unlikely to attack them in the field. This was supported by the adult choice tests, where G. spadicea females displayed, a strong oviposition preference for their natural host. In contrast, M. elatior larvae survived well on non-host plants, and the females selected several non-host species, including eggplant for oviposition. It was argued that the conflict of interests involving eggplant was overrated because eggplant is subjected to a stringent insecticide spray regime. Based on this evidence, permission for release was granted for G. spadicea. The impact of native parasitoid host range extensions to weed biological control agents in South Africa was investigated. Native parasitoids were recorded from nearly half of the agent species that had established on their target weed. The level of concealment and taxon influenced susceptibility of the agents to parasitoid attack Poorly concealed endophagous agents were most susceptible to attack, while exposed feeders were fairly free from attack. However, native parasitoids were reported not to strongly influence weed biological control agent populations and it was concluded that no agent should be rejected based only on its susceptibility to native parasitoid attack. Finally, several predictions are made as to the potential success of G. spadicea on S. sisymbriifolium and some of the challenges facing the biological control of weeds are discussed.
- Full Text:
- Date Issued: 1995
Insect herbivores associated with Senecio pterophorus and Senecio inaequidens at Butterworth, South Africa
- Muwanga-Zake, Johnnie Wycliffe Frank
- Authors: Muwanga-Zake, Johnnie Wycliffe Frank
- Date: 1995
- Subjects: Senecio -- South Africa -- Eastern Cape -- Butterworth Insect-plant relationships -- South Africa -- Eastern Cape -- Butterworth Insects -- South Africa -- Eastern Cape -- Butterworth -- Host plants Insects -- South Africa -- Eastern Cape -- Butterworth -- Food Lixus -- biology Insect-plant relationships
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5716 , http://hdl.handle.net/10962/d1005402
- Description: This study lists and compares insect herbivores of Senecio inaequidens and of Senecio pterophorus around Butterworth by 18 Tables and 38 Figures. The two plant species are widely distributed around Southern Africa. Samples were taken during a month of each of the four main seasons of the year, which in Butterworth did not show much difference in temperature. The two plant species shared generalist insect herbivores most of which were also on other surrounding plant species. The population of most of the herbivores was affected by the seasons while the two plant species were not seasonal, thus the correlation between insect herbivores and the two plant species was not significant. The heavier, and apparently chemically protected species, Senecio pterophorus, accommodated borers and supported a wider variety of but fewer individual insects. Senecio pterophorus could be having more defences but had more serious herbivores. There were no serious insect herbivores that could control the abundance of the two plant species around Butterworth. Anatomical differences between S. pterophorus and S. inaequidens could be highlighted by the kind and number of insect herbivores found upon them. Similarities were more difficult to identify through studying their insect herbivores. The biology of one serious weed, Lixus sp., on Senecio pterophorus was studied in more detail. The life cycle of the weevil was found to be synchronised with that of the host. This weevil shared the stems with other species of weevils with no indication of competition.
- Full Text:
- Date Issued: 1995
- Authors: Muwanga-Zake, Johnnie Wycliffe Frank
- Date: 1995
- Subjects: Senecio -- South Africa -- Eastern Cape -- Butterworth Insect-plant relationships -- South Africa -- Eastern Cape -- Butterworth Insects -- South Africa -- Eastern Cape -- Butterworth -- Host plants Insects -- South Africa -- Eastern Cape -- Butterworth -- Food Lixus -- biology Insect-plant relationships
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5716 , http://hdl.handle.net/10962/d1005402
- Description: This study lists and compares insect herbivores of Senecio inaequidens and of Senecio pterophorus around Butterworth by 18 Tables and 38 Figures. The two plant species are widely distributed around Southern Africa. Samples were taken during a month of each of the four main seasons of the year, which in Butterworth did not show much difference in temperature. The two plant species shared generalist insect herbivores most of which were also on other surrounding plant species. The population of most of the herbivores was affected by the seasons while the two plant species were not seasonal, thus the correlation between insect herbivores and the two plant species was not significant. The heavier, and apparently chemically protected species, Senecio pterophorus, accommodated borers and supported a wider variety of but fewer individual insects. Senecio pterophorus could be having more defences but had more serious herbivores. There were no serious insect herbivores that could control the abundance of the two plant species around Butterworth. Anatomical differences between S. pterophorus and S. inaequidens could be highlighted by the kind and number of insect herbivores found upon them. Similarities were more difficult to identify through studying their insect herbivores. The biology of one serious weed, Lixus sp., on Senecio pterophorus was studied in more detail. The life cycle of the weevil was found to be synchronised with that of the host. This weevil shared the stems with other species of weevils with no indication of competition.
- Full Text:
- Date Issued: 1995
Interspecific interactions of the sea urchin Parechinus angulosus and the effect of variations in microhabitat availability
- Authors: Farquhar, Michael Robert
- Date: 1995
- Subjects: Sea urchins , Niche (Ecology) , Sea urchins -- Habitat
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5720 , http://hdl.handle.net/10962/d1005406 , Sea urchins , Niche (Ecology) , Sea urchins -- Habitat
- Description: Intertidal populations of the sea urchin Parechinus angulosus were examined at seven sites along the south and east coasts of South Africa. At the five southerly sites, P. angulosus occurred in allopatry, while, at the two northerly sites, it occurred in sympatry with several species of Indopacific sea urchins. At the five southerly sites, there was a significant correlation of number of sea urchins per pool with the surface area of the overhang in that pool. This relationship broke down for P. angulosus at the two northerly sites, where there was a significant correlation between these two factors for Stomopneustes variolaris. The density of populations of P. angulosus at the two northerly sites, although not significantly different from all five southerly sites, were considerably lower. There was a significant difference between the mean size of populations at the seven sites. Although no direct evidence is available, a competitive interaction between these two species is proposed to explain the observed patterns of microhabitat utilization and densities. Two series of manipulative experiments were conducted at Kowie Point, where P. angulosus occurs in sympatry, The first, involved urchin removals from three experimental sites. The percentage cover of four functional algal groups was monitored in three experimental and three unaltered control sites over a three month period in spring (Sept. - Dec.) 1993. Two 100 point 0,25m² random point quadrats were thrown at each of the six sites at approximately monthly intervals over the experimental period. Since initial cover of algae varied at the six sites, ANOVA's of the change in percentage cover of four functional algal groups were used to determine treatment effects. No treatment effects were identified for any of the four functional algal groups. There was a strong negative correlation between the percentage cover of foliose algae and encrusting coralline algae, suggesting the possibility of a competitive relationship between them. The second, involved the experimental manipulation of microhabitat availability. The blocking of overhangs, by means of cement filled bags, resulted in an approximate 50% decrease in the total number of urchins in the two experimental pools, and the two control pools without overhangs. However, in the control pool with overhangs there was a slight increase in the number of urchins over the same period. Clearly, the presence of suitable shelters, is a prerequisite for the maintenance of dense intertidal population of P. angulosus. It is proposed that, due to the exposed nature of the South African coast, intertidal populations of P.angulosus are restricted to inhabiting suitable shelters from which they emerge to feed on passing drift algae. The implications of these findings are discussed in terms of current ecological literature.
- Full Text:
- Date Issued: 1995
- Authors: Farquhar, Michael Robert
- Date: 1995
- Subjects: Sea urchins , Niche (Ecology) , Sea urchins -- Habitat
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5720 , http://hdl.handle.net/10962/d1005406 , Sea urchins , Niche (Ecology) , Sea urchins -- Habitat
- Description: Intertidal populations of the sea urchin Parechinus angulosus were examined at seven sites along the south and east coasts of South Africa. At the five southerly sites, P. angulosus occurred in allopatry, while, at the two northerly sites, it occurred in sympatry with several species of Indopacific sea urchins. At the five southerly sites, there was a significant correlation of number of sea urchins per pool with the surface area of the overhang in that pool. This relationship broke down for P. angulosus at the two northerly sites, where there was a significant correlation between these two factors for Stomopneustes variolaris. The density of populations of P. angulosus at the two northerly sites, although not significantly different from all five southerly sites, were considerably lower. There was a significant difference between the mean size of populations at the seven sites. Although no direct evidence is available, a competitive interaction between these two species is proposed to explain the observed patterns of microhabitat utilization and densities. Two series of manipulative experiments were conducted at Kowie Point, where P. angulosus occurs in sympatry, The first, involved urchin removals from three experimental sites. The percentage cover of four functional algal groups was monitored in three experimental and three unaltered control sites over a three month period in spring (Sept. - Dec.) 1993. Two 100 point 0,25m² random point quadrats were thrown at each of the six sites at approximately monthly intervals over the experimental period. Since initial cover of algae varied at the six sites, ANOVA's of the change in percentage cover of four functional algal groups were used to determine treatment effects. No treatment effects were identified for any of the four functional algal groups. There was a strong negative correlation between the percentage cover of foliose algae and encrusting coralline algae, suggesting the possibility of a competitive relationship between them. The second, involved the experimental manipulation of microhabitat availability. The blocking of overhangs, by means of cement filled bags, resulted in an approximate 50% decrease in the total number of urchins in the two experimental pools, and the two control pools without overhangs. However, in the control pool with overhangs there was a slight increase in the number of urchins over the same period. Clearly, the presence of suitable shelters, is a prerequisite for the maintenance of dense intertidal population of P. angulosus. It is proposed that, due to the exposed nature of the South African coast, intertidal populations of P.angulosus are restricted to inhabiting suitable shelters from which they emerge to feed on passing drift algae. The implications of these findings are discussed in terms of current ecological literature.
- Full Text:
- Date Issued: 1995
The conflict between adaptation and constraint : the case of the Siphonariid limpets
- Authors: Chambers, Richard Jonathan
- Date: 1995
- Subjects: Siphonaria -- Adaptation Limpets -- Adaptation Pulmonata
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5657 , http://hdl.handle.net/10962/d1005340
- Description: The reproductive strategies of marine invertebrates have been related to various aspects of both their ecology, and their phylogenetic history. It has been the purpose of this thesis to try and separate these components among Siphonaria, a group of marine pulmonates. The taxonomy of these species is revised and I conclude that nine species are valid. All species deposit benthic egg masses and development may be either direct (S. anneae, S. compressa, S. dayi, S. nigerrima, S. serrata and S. tenuicostulata) or planktonic (S. capensis, S.concinna and S. oculus). Data on distribution and life-history relating to mode of larval development is then presented for 26 species of Siphonaria worldwide. Fifteen species are direct developers, nine are planktonic developers and a further two appear to have a dual developmental capacity, retaining both the velar swimming apparatus of a planktonic developer and the crawling foot of a direct developer. Direct developing species hatch from larger egg capsules, and generally occur higher on the shore than planktonic developers. Worldwide, planktonic developers are more widespread than direct developers, and individual planktonic species may have a greater latitudinal range. In most S~honaria subgenera, mode of larval development appears to be constant, although two subgenera (Patel/opsis and Sacculosiphonaria) include both developmental types. Locally, the intertidal zonation of three sympatric species (S. capensis, S. concinna and S. serrata) does not support a model which predicts direct development on the high shore and planktonic development on the low shore. However, distributions do correspond to particular intertidal microhabitats, and while there may be no, direct relationship between mode of larval development and intertidal height, the physical structure of egg masses, and the microhabitats used for spawning appear adaptive with regards to desiccation in the intertidal. S. concinna (planktonic development) and S. serrata (direct development) occur in similar microhabitats and are likely to be under similar selection pressures. In having different modes of larval development, there appears to be more than one optimal solution in a particular selective regime. In addition, both species seem to apportion similar amounts of energy to reproduction for each spawning episode, and also annually suggesting an optimum allocation of resources to reproduction. Genetic investigations using PolyAcrylamide Gel Electrophoresis (PAGE) confirm the status of the southern African species initially described, and indicate greater genetic variability associated with planktonic developing species than direct developing species. The systematic relationships revealed by DNA fingerprinting support the current classification systems, and also have implications with regards the evolution of larval development: direct development may be the plesiomorphic condition in, and among, some Siphonaria groups. There are both phylogenetic and adaptive explanations for the distribution of reproductive mode among benthic marine invertebrates. An evolutionary question, however, is not just a matter of either adaptation or constraint, it is a combination of these. Both contribute to the distribution of developmental mode among Siphonaria.
- Full Text:
- Date Issued: 1995
- Authors: Chambers, Richard Jonathan
- Date: 1995
- Subjects: Siphonaria -- Adaptation Limpets -- Adaptation Pulmonata
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5657 , http://hdl.handle.net/10962/d1005340
- Description: The reproductive strategies of marine invertebrates have been related to various aspects of both their ecology, and their phylogenetic history. It has been the purpose of this thesis to try and separate these components among Siphonaria, a group of marine pulmonates. The taxonomy of these species is revised and I conclude that nine species are valid. All species deposit benthic egg masses and development may be either direct (S. anneae, S. compressa, S. dayi, S. nigerrima, S. serrata and S. tenuicostulata) or planktonic (S. capensis, S.concinna and S. oculus). Data on distribution and life-history relating to mode of larval development is then presented for 26 species of Siphonaria worldwide. Fifteen species are direct developers, nine are planktonic developers and a further two appear to have a dual developmental capacity, retaining both the velar swimming apparatus of a planktonic developer and the crawling foot of a direct developer. Direct developing species hatch from larger egg capsules, and generally occur higher on the shore than planktonic developers. Worldwide, planktonic developers are more widespread than direct developers, and individual planktonic species may have a greater latitudinal range. In most S~honaria subgenera, mode of larval development appears to be constant, although two subgenera (Patel/opsis and Sacculosiphonaria) include both developmental types. Locally, the intertidal zonation of three sympatric species (S. capensis, S. concinna and S. serrata) does not support a model which predicts direct development on the high shore and planktonic development on the low shore. However, distributions do correspond to particular intertidal microhabitats, and while there may be no, direct relationship between mode of larval development and intertidal height, the physical structure of egg masses, and the microhabitats used for spawning appear adaptive with regards to desiccation in the intertidal. S. concinna (planktonic development) and S. serrata (direct development) occur in similar microhabitats and are likely to be under similar selection pressures. In having different modes of larval development, there appears to be more than one optimal solution in a particular selective regime. In addition, both species seem to apportion similar amounts of energy to reproduction for each spawning episode, and also annually suggesting an optimum allocation of resources to reproduction. Genetic investigations using PolyAcrylamide Gel Electrophoresis (PAGE) confirm the status of the southern African species initially described, and indicate greater genetic variability associated with planktonic developing species than direct developing species. The systematic relationships revealed by DNA fingerprinting support the current classification systems, and also have implications with regards the evolution of larval development: direct development may be the plesiomorphic condition in, and among, some Siphonaria groups. There are both phylogenetic and adaptive explanations for the distribution of reproductive mode among benthic marine invertebrates. An evolutionary question, however, is not just a matter of either adaptation or constraint, it is a combination of these. Both contribute to the distribution of developmental mode among Siphonaria.
- Full Text:
- Date Issued: 1995
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