Fishing for resilience : herbivore and algal dynamics on coral reefs in Kenya.
- Authors: Humphries, Austin Turner
- Date: 2014
- Subjects: Coral reef conservation -- Kenya , Coral reef ecology -- Kenya , Coral reef biology -- Kenya , Coral reef fishes -- Kenya , Herbivores -- Kenya , Algae -- Control -- Kenya , Fishery management -- Kenya
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5877 , http://hdl.handle.net/10962/d1013147
- Description: Herbivory is a key process that mediates the abundance of primary producers and community composition in both terrestrial and aquatic ecosystems. On tropical coral reefs, changes in herbivory are often related to phase shifts between coral-dominance and dominance by seaweeds, or foliose macroalgae. Resilience or capacity to resist and reverse such phase shifts is, therefore, viewed as a critical function on coral reefs. This thesis used grazer exclusion and assay experiments at six sites within three different fisheries management regimes in Kenya to identify the impacts of herbivores (sea urchins and fishes) on algal dynamics in the context of coral reef resilience. First, I examined the grazing rates necessary to prevent phase shifts by quantifying consumption and algal production. Here, I found that, over a 390-day experiment, at least 50 percent of algal production must be consumed to avoid accumulation of algal biomass. Using video observations, I also showed that scraping parrotfishes remove more algae (per unit of fish biomass) than previously assumed, and that sea urchins, if released from predation, have similar impacts to fishes. Then I focused on algal succession, and found that sea urchins and fishes have different effects that are mediated by their abundances and species composition. Where sea urchins were less abundant and parrotfishes absent (e.g. young fisheries closures), progression of algae from turfs to early and then late successional macroalgae occurred rapidly and within 100 days. I then turned my focus to the removal of already established macroalgae (grown for > 1 yr in the absence of herbivores) and showed that sea urchins and browsing fishes were able to remove significant amounts of macroalgae where either herbivore was abundant. However, using multiple-choice selectivity assays and in situ video recordings, I found that browsing fishes fed very selectively with low overlap in diet among species, leading to low functional redundancy within a high diversity system. Finally, using long-term survey data (from 28 sites) to build a 43-year chronosequence, I showed that it is possible that the effects of herbivory will not be constant across transitions from open fishing to fishery closures through non-linear grazing intensity. Therefore, increases in herbivory within fisheries closures may not be immediate and may allow a window of opportunity for algae to go from turf to unpalatable macroalgae until scraping and browsing fishes fully recover from fishing (~ 20 years). The findings in this thesis are novel and raise concern over the potential implications of the slow recovery of parrotfishes or, given lower than expected functional redundancy in grazing effects, the absence of even one browsing fish species in fisheries closures. Overall, this thesis highlights the importance of herbivore community dynamics in mediating interactions among algae, and provides new insights for conservation and management actions that attempt to bolster the resilience of coral reefs.
- Full Text:
- Authors: Humphries, Austin Turner
- Date: 2014
- Subjects: Coral reef conservation -- Kenya , Coral reef ecology -- Kenya , Coral reef biology -- Kenya , Coral reef fishes -- Kenya , Herbivores -- Kenya , Algae -- Control -- Kenya , Fishery management -- Kenya
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5877 , http://hdl.handle.net/10962/d1013147
- Description: Herbivory is a key process that mediates the abundance of primary producers and community composition in both terrestrial and aquatic ecosystems. On tropical coral reefs, changes in herbivory are often related to phase shifts between coral-dominance and dominance by seaweeds, or foliose macroalgae. Resilience or capacity to resist and reverse such phase shifts is, therefore, viewed as a critical function on coral reefs. This thesis used grazer exclusion and assay experiments at six sites within three different fisheries management regimes in Kenya to identify the impacts of herbivores (sea urchins and fishes) on algal dynamics in the context of coral reef resilience. First, I examined the grazing rates necessary to prevent phase shifts by quantifying consumption and algal production. Here, I found that, over a 390-day experiment, at least 50 percent of algal production must be consumed to avoid accumulation of algal biomass. Using video observations, I also showed that scraping parrotfishes remove more algae (per unit of fish biomass) than previously assumed, and that sea urchins, if released from predation, have similar impacts to fishes. Then I focused on algal succession, and found that sea urchins and fishes have different effects that are mediated by their abundances and species composition. Where sea urchins were less abundant and parrotfishes absent (e.g. young fisheries closures), progression of algae from turfs to early and then late successional macroalgae occurred rapidly and within 100 days. I then turned my focus to the removal of already established macroalgae (grown for > 1 yr in the absence of herbivores) and showed that sea urchins and browsing fishes were able to remove significant amounts of macroalgae where either herbivore was abundant. However, using multiple-choice selectivity assays and in situ video recordings, I found that browsing fishes fed very selectively with low overlap in diet among species, leading to low functional redundancy within a high diversity system. Finally, using long-term survey data (from 28 sites) to build a 43-year chronosequence, I showed that it is possible that the effects of herbivory will not be constant across transitions from open fishing to fishery closures through non-linear grazing intensity. Therefore, increases in herbivory within fisheries closures may not be immediate and may allow a window of opportunity for algae to go from turf to unpalatable macroalgae until scraping and browsing fishes fully recover from fishing (~ 20 years). The findings in this thesis are novel and raise concern over the potential implications of the slow recovery of parrotfishes or, given lower than expected functional redundancy in grazing effects, the absence of even one browsing fish species in fisheries closures. Overall, this thesis highlights the importance of herbivore community dynamics in mediating interactions among algae, and provides new insights for conservation and management actions that attempt to bolster the resilience of coral reefs.
- Full Text:
Inter-individual variability and phenotypic plasticity : the effect of the environment on the biogeography, population structure, ecophysiology and reproduction of the sandhoppers Talorchestia capensis and Africorchestia quadrispinosa
- Authors: Baldanzi, Simone
- Date: 2014
- Subjects: Phenotypic plasticity -- Research -- Africa, Southern Talitridae -- Research -- Africa, Southern Amphipoda -- Research -- Africa, Southern Climatic changes -- Environmental aspects -- Africa, Southern Marine biology -- Africa, Southern Adaptation (Biology) -- Africa, Southern
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5846 , http://hdl.handle.net/10962/d1011447
- Description: Climatic envelope models focus on the climatic variables affecting species or species assemblages, and are important tools to investigate the effect of climate change on their geographical ranges. These models have largely been proposed in order to make successful predictions on species‘ persistence, determining which variables are likely to induce range expansion, contraction, or shifting. More recent models, including the ability and the cost for individuals to respond promptly to an environmental stimulus, have revealed that species may express phenotypic plasticity able to induce adaptation to the new environment. Consequently, understanding how species evolve to a changing climate is fundamental. From this perspective, investigating intraspecific responses to an environmental variable may contribute to better understanding and prediction of the effect of climate change on the geographical range and evolution of species, particularly in the case of widespread species. In this context, the present study aimed at establishing how environmental variables (focussing mainly on temperature) may have contributed to shape the spatial distribution, physiology, reproductive biology and connectivity of two species of Southern African sandhoppers (Talorchestia capensis and Africorchestia quadrispinosa, Amphipoda, Talitridae). Most of the work was carried out on T. capensis, due to its widespread spatial distribution. A first investigation of the biogeography of T. capensis and A. quadrispinosa, revealed that, for both species, spatial patterns of abundance, size and sex ratio were not explained by the Abundant Centre Hypothesis (greater abundance at the core of a spatial range), but rather guided by bio-physical forces. Precisely, the abundance of sandhoppers was driven by the morphodynamic state of the beach, salinity and temperatures, with strong differentiation among sites that reflected local environmental conditions. In support of these findings, strong population structure in the genetics of T. capensis was found (three main groups) when investigating its phylogeography and genetic connectivity. Although such defined structure may suggests cryptic speciation, the concomitant within-population variation in the COX1 region of mtDNA, also highlighted the importance of individual genetic variability. High individual variability was also found in the response of T. capensis to temperature, both in its physiology (thermal plasticity) and its reproductive biology (maternal effects). Since temperature is one of the main variables affecting the coastal marine systems of southern Africa and the metabolism of animals in general, its effect on the physiology and reproduction of T. capensis was therefore investigated. Thermal responses to increasing/decreasing temperatures were assessed for separated populations of T. capensis. Individual variability was reported in the oxygen consumption of T. capensis in response to temperature (high variation around the means, especially for increasing temperatures). Among population differences in thermal sensitivity were significantly correlated with air temperature variability experienced over the past 23 years, highlighting the importance of historical temperature fluctuations to the current thermal physiology of these sandhoppers. Temperature also had an important effect on the reproductive plasticity of T. capensis. Different temperatures induced mothers to adjust the size of their offspring (i.e. egg size), with larger eggs produced at lower temperatures. Interestingly, females showed strongly significant among individual variation in the size of the eggs. Given the importance of understanding rapid responses of organisms to climate change and considering the fundamental role played by phenotypic plasticity in evolution, the overall study revealed the significance of individual plasticity and variability in response to the environment and highlighted its importance. Particularly, studying the thermal physiology of separated populations and understanding within population reproductive plasticity in response to temperature, helped to clarify how differences among individual responses have important consequences at the population level, possibly explaining the widespread distribution of T. capensis.
- Full Text:
- Authors: Baldanzi, Simone
- Date: 2014
- Subjects: Phenotypic plasticity -- Research -- Africa, Southern Talitridae -- Research -- Africa, Southern Amphipoda -- Research -- Africa, Southern Climatic changes -- Environmental aspects -- Africa, Southern Marine biology -- Africa, Southern Adaptation (Biology) -- Africa, Southern
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5846 , http://hdl.handle.net/10962/d1011447
- Description: Climatic envelope models focus on the climatic variables affecting species or species assemblages, and are important tools to investigate the effect of climate change on their geographical ranges. These models have largely been proposed in order to make successful predictions on species‘ persistence, determining which variables are likely to induce range expansion, contraction, or shifting. More recent models, including the ability and the cost for individuals to respond promptly to an environmental stimulus, have revealed that species may express phenotypic plasticity able to induce adaptation to the new environment. Consequently, understanding how species evolve to a changing climate is fundamental. From this perspective, investigating intraspecific responses to an environmental variable may contribute to better understanding and prediction of the effect of climate change on the geographical range and evolution of species, particularly in the case of widespread species. In this context, the present study aimed at establishing how environmental variables (focussing mainly on temperature) may have contributed to shape the spatial distribution, physiology, reproductive biology and connectivity of two species of Southern African sandhoppers (Talorchestia capensis and Africorchestia quadrispinosa, Amphipoda, Talitridae). Most of the work was carried out on T. capensis, due to its widespread spatial distribution. A first investigation of the biogeography of T. capensis and A. quadrispinosa, revealed that, for both species, spatial patterns of abundance, size and sex ratio were not explained by the Abundant Centre Hypothesis (greater abundance at the core of a spatial range), but rather guided by bio-physical forces. Precisely, the abundance of sandhoppers was driven by the morphodynamic state of the beach, salinity and temperatures, with strong differentiation among sites that reflected local environmental conditions. In support of these findings, strong population structure in the genetics of T. capensis was found (three main groups) when investigating its phylogeography and genetic connectivity. Although such defined structure may suggests cryptic speciation, the concomitant within-population variation in the COX1 region of mtDNA, also highlighted the importance of individual genetic variability. High individual variability was also found in the response of T. capensis to temperature, both in its physiology (thermal plasticity) and its reproductive biology (maternal effects). Since temperature is one of the main variables affecting the coastal marine systems of southern Africa and the metabolism of animals in general, its effect on the physiology and reproduction of T. capensis was therefore investigated. Thermal responses to increasing/decreasing temperatures were assessed for separated populations of T. capensis. Individual variability was reported in the oxygen consumption of T. capensis in response to temperature (high variation around the means, especially for increasing temperatures). Among population differences in thermal sensitivity were significantly correlated with air temperature variability experienced over the past 23 years, highlighting the importance of historical temperature fluctuations to the current thermal physiology of these sandhoppers. Temperature also had an important effect on the reproductive plasticity of T. capensis. Different temperatures induced mothers to adjust the size of their offspring (i.e. egg size), with larger eggs produced at lower temperatures. Interestingly, females showed strongly significant among individual variation in the size of the eggs. Given the importance of understanding rapid responses of organisms to climate change and considering the fundamental role played by phenotypic plasticity in evolution, the overall study revealed the significance of individual plasticity and variability in response to the environment and highlighted its importance. Particularly, studying the thermal physiology of separated populations and understanding within population reproductive plasticity in response to temperature, helped to clarify how differences among individual responses have important consequences at the population level, possibly explaining the widespread distribution of T. capensis.
- Full Text:
Molecular phylogeny, phylogeography and evolutionary adaptation of foraging behaviour amongst sympatric patellid limpets along the southern African shoreline
- Authors: Mmonwa, Kolobe Lucas
- Date: 2014
- Subjects: Limpets -- South Africa -- Adaptation , Limpets -- South Africa -- Ecology
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5882 , http://hdl.handle.net/10962/d1013222
- Description: The southern African shoreline is inhabited by a great diversity of patellid limpets of which most are endemic to South Africa. These limpets have evolved foraging mechanisms that partition ecological resources and reduce interspecific competition, resulting in ecological specialists and generalists. The evolution of ecological specialization or generalization remains poorly understood and there is no agreement on how such evolutionary transitions are correlated with levels of genetic diversity. This study investigated evolutionary correlations between territoriality in foraging and genetic structure of southern African patellid limpets (Cymbula and Scutellastra spp.) using stable δ13C and δ15N ratios, mitochondrial and nuclear DNA markers. The outcomes of the study were best rationalized and understood in the context of the scale of analysis in terms of both space and taxonomy. At biogeographic scales and the interspecific level, the stable isotope signatures and genetic structure of these limpets are determined by extrinsic factors such as biogeography and oceanography. However, at the smallest site scales and the intraspecific level, the isotope signatures and genetic diversity of these limpets are significantly correlated to their foraging traits. At large scales, there are no correlations between foraging behaviour and either genetic structure or isotope signature. At smaller scales, territorial Scutellastra foragers display both isotopic enrichment and greater haplotype diversity than congeneric non-territorial foragers. Thus, the isotope signatures and genetic structure of these limpets are determined by intrinsic species-specific response linked to their foraging behaviour. However, this pattern was intriguing as differences between territorial and non-territorial limpets in both isotopic signature and genetic diversity were observed only along the south coast when the same species were compared in other biogeographic regions. The significant interaction effect between foraging behaviour and stable isotope signatures was only observed from the sites within the Agulhas Bank or which are strongly influenced by the Agulhas current. This south enrichment in isotopic ratios is due to the mixing differences between onshore and offshore waters as the Agulhas current moves from east to south. At the generic level, the correlation between foraging behaviour and isotope signatures and genetic structure were particularly profound for Scutellastra species. The molecular phylogeny revealed deep evolutionary divergence between territorial and non-territorial Scutellastra spp. This divergence was concordant with morphological differences in shell shape and radula anatomy between territorial and non-territorial species. A taxonomic review of the scutellastrid spp. is proposed, suggesting possible re-consideration of the genus as two genera characterized by either territoriality or non-territoriality. The divergence between territorial and non-territorial species in both Scutellastra and Cymbula took place approximately in the early Oligocene. Major climatic cooling and decreases in sea level occurred during the Oligocene and this probably exposed much of the lower intertidal zone, increasing new potential habitats and algal availability. The Oligocene exposure of rocky shores and algal abundance in the lower intertidal zone probably elicited resource partitioning amongst these patellid limpets and subsequently, the evolution of territorial and non-territorial species. Analyses of the demographic history of these patellid limpets revealed evidence of post-glacial spatial expansion around the Pleistocene, implying these limpets were at population equilibrium during the dramatic LGM sea temperatures. Thus, these limpets managed to expand their range during dynamic oceanographic oscillations and dramatic sea-level changes in the Pleistocene. This study highlighted the importance of applying ecological traits as a subject to investigate and comprehend the evolutionary ecology of marine herbivores. The foraging traits of these true limpets are reflected in both their stable isotope ratios and genealogy, presumably as an evolutionary consequence of competition.
- Full Text:
- Authors: Mmonwa, Kolobe Lucas
- Date: 2014
- Subjects: Limpets -- South Africa -- Adaptation , Limpets -- South Africa -- Ecology
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5882 , http://hdl.handle.net/10962/d1013222
- Description: The southern African shoreline is inhabited by a great diversity of patellid limpets of which most are endemic to South Africa. These limpets have evolved foraging mechanisms that partition ecological resources and reduce interspecific competition, resulting in ecological specialists and generalists. The evolution of ecological specialization or generalization remains poorly understood and there is no agreement on how such evolutionary transitions are correlated with levels of genetic diversity. This study investigated evolutionary correlations between territoriality in foraging and genetic structure of southern African patellid limpets (Cymbula and Scutellastra spp.) using stable δ13C and δ15N ratios, mitochondrial and nuclear DNA markers. The outcomes of the study were best rationalized and understood in the context of the scale of analysis in terms of both space and taxonomy. At biogeographic scales and the interspecific level, the stable isotope signatures and genetic structure of these limpets are determined by extrinsic factors such as biogeography and oceanography. However, at the smallest site scales and the intraspecific level, the isotope signatures and genetic diversity of these limpets are significantly correlated to their foraging traits. At large scales, there are no correlations between foraging behaviour and either genetic structure or isotope signature. At smaller scales, territorial Scutellastra foragers display both isotopic enrichment and greater haplotype diversity than congeneric non-territorial foragers. Thus, the isotope signatures and genetic structure of these limpets are determined by intrinsic species-specific response linked to their foraging behaviour. However, this pattern was intriguing as differences between territorial and non-territorial limpets in both isotopic signature and genetic diversity were observed only along the south coast when the same species were compared in other biogeographic regions. The significant interaction effect between foraging behaviour and stable isotope signatures was only observed from the sites within the Agulhas Bank or which are strongly influenced by the Agulhas current. This south enrichment in isotopic ratios is due to the mixing differences between onshore and offshore waters as the Agulhas current moves from east to south. At the generic level, the correlation between foraging behaviour and isotope signatures and genetic structure were particularly profound for Scutellastra species. The molecular phylogeny revealed deep evolutionary divergence between territorial and non-territorial Scutellastra spp. This divergence was concordant with morphological differences in shell shape and radula anatomy between territorial and non-territorial species. A taxonomic review of the scutellastrid spp. is proposed, suggesting possible re-consideration of the genus as two genera characterized by either territoriality or non-territoriality. The divergence between territorial and non-territorial species in both Scutellastra and Cymbula took place approximately in the early Oligocene. Major climatic cooling and decreases in sea level occurred during the Oligocene and this probably exposed much of the lower intertidal zone, increasing new potential habitats and algal availability. The Oligocene exposure of rocky shores and algal abundance in the lower intertidal zone probably elicited resource partitioning amongst these patellid limpets and subsequently, the evolution of territorial and non-territorial species. Analyses of the demographic history of these patellid limpets revealed evidence of post-glacial spatial expansion around the Pleistocene, implying these limpets were at population equilibrium during the dramatic LGM sea temperatures. Thus, these limpets managed to expand their range during dynamic oceanographic oscillations and dramatic sea-level changes in the Pleistocene. This study highlighted the importance of applying ecological traits as a subject to investigate and comprehend the evolutionary ecology of marine herbivores. The foraging traits of these true limpets are reflected in both their stable isotope ratios and genealogy, presumably as an evolutionary consequence of competition.
- Full Text:
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