Assessing the impact of climate change on mangrove crabs: the role of ontogenetic macrophysiology and settlement in the persistence of central and marginal populations
- Authors: Mostert, Bruce Petrus
- Date: 2016
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/826 , vital:19994
- Description: After a brief respite in the mid to late 20th century, macro physiology has come to the fore in elucidating large scale ecological patterns and processes as physiological assumptions often form the backbone of many predictive theories associated with species distributions. Critically, macro physiological patterns are valuable in explaining physiological variation across multiple scales and provide insights into the effects of climate change on populations spanning a wide range of latitudes. This can assist in predicting possible distribution expansions, contractions or shifts in light of current climate change scenarios. From this perspective, investigating intra- and inter-specific physiological responses to environmental stress may contribute to better understanding and predicting the effects of climate change on geographical ranges. Further, investigating the physiological effects to environmental stresses across ontogenetic stages allows for the identification of weak links within the lifecycle of a species. Additionally, determining settlement characteristics along a latitudinal cline provides integrated indications of the sustainability of populations, highlighting vulnerable regions in terms of repopulation of viable habitats. In this context, the present study aimed at establishing how temperature, in a physiological context, may affect reproductive biology of two species of mangrove crab, Perisesarma guttatum and Uca urvillei at the centre (Kenya) and edge (South Africa) of their distributional range along the east coast of Africa and highlight possible consequences for range distributions. A third species, Neosarmatium africanum, only in South Africa, was included to provide additional interspecies comparisons. Furthermore, settlement characteristics of brachyuran populations at the centre and edge of their distributional range were considered in order to determine how settlement may contribute to population persistence. Physiological investigations at the centre and edge of distributional range and across ontogenetic stages (larvae, stage 2 and 4 embryos, non-gravid and gravid females) under the concept of oxygen and capacity limitation of thermal tolerance (OCLTT), revealed that, for both species, populations at the centre of their distribution (Kenya) were generally more robust to increasing temperatures and generally displayed greater physiological stability with increasing temperatures compared to their conspecifics in South Africa. Variability in physiological robustness between regions, did however, differ among ontogenetic stages and species but, overall, were evident throughout. Within and between regions, adaptation to oxygen extraction in both milieus (air or water) was displayed for present temperature conditions but aerial respiration largely alleviated increased thermal stress due to overcoming the limitations of reduced oxygen availability and diffusiveness in water for all bimodal ontogenetic stages. Brooding eggs proved to be a physiologically critical process with either heighted oxygen consumption for gravid females or collapse of physiological processes demonstrated by supressed oxygen consumption. The physiological cost of brooding eggs, referred to as maternal costs, was reflected in in both Perisesarma guttatum and Uca urvillei where, in most cases, maternal costs were negative. Again, aerial respiration was able to alleviated increased thermal stress, as shown by positive maternal costs indicating sustained maternal care, but this mechanism was species and regionally specific. Settlement patterns differed between the edge and centre of distribution of the species studied. This difference was predominantly driven by zonal preference within the mangal and/or effects of new and full moon (lunar phase). Overall, settlement dynamics were more widely variable in South Africa, both spatially and temporally, than in Kenya. Finally, empirical physiological data from ontogenetic stages present during the reproductive process (early and late stage embryos) and from non-gravid and gravid females were used in conjunction with data mined from the existing literature to parameterise an individual based model designed to simulate reproductive output at the centre and edge of distribution of Perisesarma guttatum. Physiological data indicate that, in terms of reproductive output across increasing temperatures, populations based at the centre of their distribution presently outperform their counterparts at the edge of the species’ distribution, but reproductive output stagnated as temperature rose. Edge of distribution populations consistently increased reproductive output with increasing temperatures to eventually outperform centre of distribution populations at higher simulated temperatures. Overall, results of the physiological and settlement studies suggest that with increased climate change there may be a contraction of distributional range of the study species from high latitudes to low latitudes, contrary to general poleward shifts/migrations seen in most species, with possible contractions of the entire ecosystem mirroring the disappearance of keystone mangrove macrofauna.
- Full Text:
- Authors: Mostert, Bruce Petrus
- Date: 2016
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/826 , vital:19994
- Description: After a brief respite in the mid to late 20th century, macro physiology has come to the fore in elucidating large scale ecological patterns and processes as physiological assumptions often form the backbone of many predictive theories associated with species distributions. Critically, macro physiological patterns are valuable in explaining physiological variation across multiple scales and provide insights into the effects of climate change on populations spanning a wide range of latitudes. This can assist in predicting possible distribution expansions, contractions or shifts in light of current climate change scenarios. From this perspective, investigating intra- and inter-specific physiological responses to environmental stress may contribute to better understanding and predicting the effects of climate change on geographical ranges. Further, investigating the physiological effects to environmental stresses across ontogenetic stages allows for the identification of weak links within the lifecycle of a species. Additionally, determining settlement characteristics along a latitudinal cline provides integrated indications of the sustainability of populations, highlighting vulnerable regions in terms of repopulation of viable habitats. In this context, the present study aimed at establishing how temperature, in a physiological context, may affect reproductive biology of two species of mangrove crab, Perisesarma guttatum and Uca urvillei at the centre (Kenya) and edge (South Africa) of their distributional range along the east coast of Africa and highlight possible consequences for range distributions. A third species, Neosarmatium africanum, only in South Africa, was included to provide additional interspecies comparisons. Furthermore, settlement characteristics of brachyuran populations at the centre and edge of their distributional range were considered in order to determine how settlement may contribute to population persistence. Physiological investigations at the centre and edge of distributional range and across ontogenetic stages (larvae, stage 2 and 4 embryos, non-gravid and gravid females) under the concept of oxygen and capacity limitation of thermal tolerance (OCLTT), revealed that, for both species, populations at the centre of their distribution (Kenya) were generally more robust to increasing temperatures and generally displayed greater physiological stability with increasing temperatures compared to their conspecifics in South Africa. Variability in physiological robustness between regions, did however, differ among ontogenetic stages and species but, overall, were evident throughout. Within and between regions, adaptation to oxygen extraction in both milieus (air or water) was displayed for present temperature conditions but aerial respiration largely alleviated increased thermal stress due to overcoming the limitations of reduced oxygen availability and diffusiveness in water for all bimodal ontogenetic stages. Brooding eggs proved to be a physiologically critical process with either heighted oxygen consumption for gravid females or collapse of physiological processes demonstrated by supressed oxygen consumption. The physiological cost of brooding eggs, referred to as maternal costs, was reflected in in both Perisesarma guttatum and Uca urvillei where, in most cases, maternal costs were negative. Again, aerial respiration was able to alleviated increased thermal stress, as shown by positive maternal costs indicating sustained maternal care, but this mechanism was species and regionally specific. Settlement patterns differed between the edge and centre of distribution of the species studied. This difference was predominantly driven by zonal preference within the mangal and/or effects of new and full moon (lunar phase). Overall, settlement dynamics were more widely variable in South Africa, both spatially and temporally, than in Kenya. Finally, empirical physiological data from ontogenetic stages present during the reproductive process (early and late stage embryos) and from non-gravid and gravid females were used in conjunction with data mined from the existing literature to parameterise an individual based model designed to simulate reproductive output at the centre and edge of distribution of Perisesarma guttatum. Physiological data indicate that, in terms of reproductive output across increasing temperatures, populations based at the centre of their distribution presently outperform their counterparts at the edge of the species’ distribution, but reproductive output stagnated as temperature rose. Edge of distribution populations consistently increased reproductive output with increasing temperatures to eventually outperform centre of distribution populations at higher simulated temperatures. Overall, results of the physiological and settlement studies suggest that with increased climate change there may be a contraction of distributional range of the study species from high latitudes to low latitudes, contrary to general poleward shifts/migrations seen in most species, with possible contractions of the entire ecosystem mirroring the disappearance of keystone mangrove macrofauna.
- Full Text:
Ecological engineering on rocky shores: grazing, predation, nutrient availability and their influence on epifaunal communities
- Authors: Ndhlovu, Aldwin
- Date: 2016
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: http://hdl.handle.net/10962/3453 , vital:20500
- Description: “Ecosystem engineers modify, create/destroy habitat and directly/indirectly modulate availability of resources to other species by causing physical state changes to biotic and abiotic materials” (Jones et al, 1994, page 1). Previous studies have analysed top-down and bottom-up relationships to determine which form of regulation is key in controlling community composition. This study assessed the direct and indirect effects of top down and bottom up factors on the epifaunal communities of macro-algae. Epifauna are subject to the direct top-down effects of predation and bottom up effects of habitat availability. Habitat availability in turn experiences the direct top down effects of grazing and bottom up effects of nutrient availability due to upwelling. Grazing and upwelling may therefore have indirect effects on macro-algal epifauna. Four treatments (Procedural controls, Controls, Predator or Grazer exclusion) set out in a block design (n = 5) were monitored monthly for algal cover of the substratum for 12 months with the surface area of algal plants and epifaunal species composition and abundances assessed at the end of the experiment. The red alga Gelidium pristoides was selected as the ecosystem engineer as it is common, supports a diverse community of epiphytic animals and acts as a nursery for small epifaunal organisms. The experiment was run at two upwelling sites interspersed with two non-upwelling sites. Sites were separated on scales of 100km along approximately 500km of coastline. Dipping whole algae in dish washing liquid provided a strong relationship between their surface area and the weight of the film of dish washing liquid covering them. Surface area was strongly correlated to dry weight but neither surface area nor dry weight was correlated to algal cover of the substratum. Algal cover was influenced by the interactions of treatment with site (nested in upwelling) and upwelling. At all sites, treatments that allowed access to grazers, Grazer + and Control treatments, showed no significant differences and these two treatments had lower algal cover than Predator + and Closed treatments which did not differ from one another 3 [Grazer + = Control < Predator + = Closed]. A total of 44 epifaunal species were identified, with the predominant orders being Amphipoda and Isopoda. Primer results showed that only site had a significant effect on species composition, with sites that were further apart being more different. Site (nested in upwelling) had an effect on total epifaunal abundances when data were non-normalised. When total epifaunal abundances were normalised for algal cover of the substratum or algal surface area to provide density data, predation had no significant effect. Grazing did have a significant effect, but only when data were normalised to algal surface area, not cover, leading to the conclusion that indirect top-down factors through grazing of the sea weed are important in structuring epifaunal communities depending on how habitat availability is measured.
- Full Text:
- Authors: Ndhlovu, Aldwin
- Date: 2016
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: http://hdl.handle.net/10962/3453 , vital:20500
- Description: “Ecosystem engineers modify, create/destroy habitat and directly/indirectly modulate availability of resources to other species by causing physical state changes to biotic and abiotic materials” (Jones et al, 1994, page 1). Previous studies have analysed top-down and bottom-up relationships to determine which form of regulation is key in controlling community composition. This study assessed the direct and indirect effects of top down and bottom up factors on the epifaunal communities of macro-algae. Epifauna are subject to the direct top-down effects of predation and bottom up effects of habitat availability. Habitat availability in turn experiences the direct top down effects of grazing and bottom up effects of nutrient availability due to upwelling. Grazing and upwelling may therefore have indirect effects on macro-algal epifauna. Four treatments (Procedural controls, Controls, Predator or Grazer exclusion) set out in a block design (n = 5) were monitored monthly for algal cover of the substratum for 12 months with the surface area of algal plants and epifaunal species composition and abundances assessed at the end of the experiment. The red alga Gelidium pristoides was selected as the ecosystem engineer as it is common, supports a diverse community of epiphytic animals and acts as a nursery for small epifaunal organisms. The experiment was run at two upwelling sites interspersed with two non-upwelling sites. Sites were separated on scales of 100km along approximately 500km of coastline. Dipping whole algae in dish washing liquid provided a strong relationship between their surface area and the weight of the film of dish washing liquid covering them. Surface area was strongly correlated to dry weight but neither surface area nor dry weight was correlated to algal cover of the substratum. Algal cover was influenced by the interactions of treatment with site (nested in upwelling) and upwelling. At all sites, treatments that allowed access to grazers, Grazer + and Control treatments, showed no significant differences and these two treatments had lower algal cover than Predator + and Closed treatments which did not differ from one another 3 [Grazer + = Control < Predator + = Closed]. A total of 44 epifaunal species were identified, with the predominant orders being Amphipoda and Isopoda. Primer results showed that only site had a significant effect on species composition, with sites that were further apart being more different. Site (nested in upwelling) had an effect on total epifaunal abundances when data were non-normalised. When total epifaunal abundances were normalised for algal cover of the substratum or algal surface area to provide density data, predation had no significant effect. Grazing did have a significant effect, but only when data were normalised to algal surface area, not cover, leading to the conclusion that indirect top-down factors through grazing of the sea weed are important in structuring epifaunal communities depending on how habitat availability is measured.
- Full Text:
Mesoscale mechanisms of larval transport and settlement in relation to physical factors off the south coast of South Africa: a topographic approach
- Authors: Mian, Shana Iva
- Date: 2016
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5947 , http://hdl.handle.net/10962/d1020844
- Description: The majority of benthic marine invertebrate life cycles include a meroplanktonic phase. Most marine biologists recognise that this stage could play an important role in dispersal, especially for sedentary or sessile organisms. Recent studies on larval dispersal have found that dispersal does not occur to the extent that was previously believed. Some larvae instead remain close to their natal populations through larval nearshore retention mechanisms, both active and passive. This study attempted to a) describe nearshore distribution patterns of several intertidal larvae in relation to nearshore hydrodynamics, b) infer whether larvae are active or passive in the water column and whether this differs among taxa and c) compare pelagic mussel larval abundance with settlement rates, at 2 bay and 2 open coast sites on the south coast of South Africa to note any topographical effects. At each of the four sites, pelagic samples were collected along 2 transects. Larvae and current speed data were collected at various depths (surface, mid-depth and bottom) at distances of 200, 400, 900 and 2400m offshore of the coast. Settlement samples were collected at sites that were onshore of the pelagic transects using plastic souring pads following previous studies. Pelagic larval distribution showed no effect of topography. Instead larval abundance was highly variable among sample periods and among different taxa. Slow swimming D-stage mussel larvae were found to be positively correlated with both upwelling and turbulence indices suggesting that they behave as passive particles in the water column. The abundances of larval taxa with faster swimming speeds than the D-larvae were negatively correlated with upwelling and did not correlate with turbulence. This suggests that these larvae are more active within the water column than the D-larvae which would theoretically allow them to maintain their position in the water column. Further analysis revealed that mean larval depth correlated with mean depth of shoreward moving water masses regardless of the state of upwelling. These results suggest that larvae were actively selecting the depth at which they were found presumably in order to associate themselves with shoreward moving currents. This interpretation was supported by ontogenetic patterns in the behaviour of barnacle larvae, the nauplii of which were found further offshore as they aged, and closer onshore again once they reach the faster swimming cyprid stage. Although there was no detectable effect of topography on the abundances of planktonic larvae, the settlement section of this study revealed an effect of topography, with higher settlement and recruitment rates at bay sites than at the open coast sites. Settlement rates were seen to be negatively correlated with turbulence at bay sites but not open coast sites. Recruitment rates for each sample event were positively correlated with upwelling at each site. This study found pelagic larvae to be generally active within the water column which allowed them to remain close to the shore regardless of topography and upwelling/turbulence regimes. Although meaningful correlation analyses between pelagic larval supply and settlement rates could not be conducted due to loss of settlement collectors, settlement rates were affected by local topography with higher settlement and recruitment in bay sites.
- Full Text:
- Authors: Mian, Shana Iva
- Date: 2016
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5947 , http://hdl.handle.net/10962/d1020844
- Description: The majority of benthic marine invertebrate life cycles include a meroplanktonic phase. Most marine biologists recognise that this stage could play an important role in dispersal, especially for sedentary or sessile organisms. Recent studies on larval dispersal have found that dispersal does not occur to the extent that was previously believed. Some larvae instead remain close to their natal populations through larval nearshore retention mechanisms, both active and passive. This study attempted to a) describe nearshore distribution patterns of several intertidal larvae in relation to nearshore hydrodynamics, b) infer whether larvae are active or passive in the water column and whether this differs among taxa and c) compare pelagic mussel larval abundance with settlement rates, at 2 bay and 2 open coast sites on the south coast of South Africa to note any topographical effects. At each of the four sites, pelagic samples were collected along 2 transects. Larvae and current speed data were collected at various depths (surface, mid-depth and bottom) at distances of 200, 400, 900 and 2400m offshore of the coast. Settlement samples were collected at sites that were onshore of the pelagic transects using plastic souring pads following previous studies. Pelagic larval distribution showed no effect of topography. Instead larval abundance was highly variable among sample periods and among different taxa. Slow swimming D-stage mussel larvae were found to be positively correlated with both upwelling and turbulence indices suggesting that they behave as passive particles in the water column. The abundances of larval taxa with faster swimming speeds than the D-larvae were negatively correlated with upwelling and did not correlate with turbulence. This suggests that these larvae are more active within the water column than the D-larvae which would theoretically allow them to maintain their position in the water column. Further analysis revealed that mean larval depth correlated with mean depth of shoreward moving water masses regardless of the state of upwelling. These results suggest that larvae were actively selecting the depth at which they were found presumably in order to associate themselves with shoreward moving currents. This interpretation was supported by ontogenetic patterns in the behaviour of barnacle larvae, the nauplii of which were found further offshore as they aged, and closer onshore again once they reach the faster swimming cyprid stage. Although there was no detectable effect of topography on the abundances of planktonic larvae, the settlement section of this study revealed an effect of topography, with higher settlement and recruitment rates at bay sites than at the open coast sites. Settlement rates were seen to be negatively correlated with turbulence at bay sites but not open coast sites. Recruitment rates for each sample event were positively correlated with upwelling at each site. This study found pelagic larvae to be generally active within the water column which allowed them to remain close to the shore regardless of topography and upwelling/turbulence regimes. Although meaningful correlation analyses between pelagic larval supply and settlement rates could not be conducted due to loss of settlement collectors, settlement rates were affected by local topography with higher settlement and recruitment in bay sites.
- Full Text:
Penguin parenting : assortative mating, nest attendance and sex-specific chick provisioning in the African Penguin (Spheniscus demersus)
- Authors: Smith, Diane Lauren
- Date: 2016
- Subjects: African penguin , Penguins -- South Africa , Penguins -- Sexual behavior , Parental behavior in animals
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5943 , http://hdl.handle.net/10962/d1019993
- Description: Animal behaviour is especially sensitive to environmental variability and prey availability during the breeding season, and this is particularly true for non-volant, central place foragers such as the endangered African Penguin (Spheniscus demersus). Individual sex and morphology, as well as the level of assortment within mated pairs can influence both the behaviour and the reproductive success of species exhibiting biparental care. This study made use of a large biometric database and nest attendance video footage to determine the influence of intrinsic (assortative mating, brood size and chick age) and extrinsic (environmental conditions, anthropic disturbance) factors on breeding behaviour and performance of African Penguins on Bird Island, Algoa Bay, during peak breeding (March - July) in 2013. While sexual dimorphism in African Penguins is subtle, the colony-specific discriminant functions presented here provide an accurate sexing tool when only bill and flipper length are available. Despite the premise that selection of a large, high-quality mate in this longlived, monogamous seabird governs lifetime fitness, only low levels of assortative mating were found, and this only for earlier breeders, when larger females (but not males) bred. The 2013 season was a particularly successful one, coinciding with above-average sardine and anchovy abundance, and almost 80 percent of monitored nests were double-brooded, with very low levels of mortality. A- and B-chicks of double broods and singleton chicks grew at similar rates and exhibited similar body condition indices. In these conditions, chick developmental rates were independent of parental size, assortment or provisioning behaviour. Females raising a double brood were significantly lighter and in poorer body condition than those raising a single chick, although the same trend was not evident in males. Offspring sex ratio in 2013 (2.27:1) favoured male chicks, suggesting that there is potential to over-produce the larger sex when resources are plentiful. Peak nest arrival and departure times of parents did not change over the course of monitored breeding attempts (March-June), nor were they different for disturbed and undisturbed nests or for a single or double brood. The increase in CCTV-observed provisioning rate as chicks grew larger was best explained by brood size, at-sea chlorophyll a concentration, and maximum air temperature, but was unrelated to parental morphology or assortative index. Importantly, parental absenteeism commenced earlier and was markedly greater in nests frequently handled by researchers than in undisturbed nests. Both the time spent together by parents, and absenteeism were measurably affected by maximum afternoon air temperatures, the effects of which are expected to be exacerbated by poorer foraging conditions and climate change. A third of manually-monitored nests shared chick-guarding duties unequally, although this phenomenon was independent of parental sex or morphology. The adaptive benefits of mating patterns and division of labour during chick-rearing may only become apparent in a year of below-average food availability and it is highly recommended that this study be repeated in a year of scarce food resources. These findings augment past foraging ecology studies and demonstrate that investigator disturbance and environmental conditions can affect the nesting behaviour of this highly threatened seabird.
- Full Text:
- Authors: Smith, Diane Lauren
- Date: 2016
- Subjects: African penguin , Penguins -- South Africa , Penguins -- Sexual behavior , Parental behavior in animals
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5943 , http://hdl.handle.net/10962/d1019993
- Description: Animal behaviour is especially sensitive to environmental variability and prey availability during the breeding season, and this is particularly true for non-volant, central place foragers such as the endangered African Penguin (Spheniscus demersus). Individual sex and morphology, as well as the level of assortment within mated pairs can influence both the behaviour and the reproductive success of species exhibiting biparental care. This study made use of a large biometric database and nest attendance video footage to determine the influence of intrinsic (assortative mating, brood size and chick age) and extrinsic (environmental conditions, anthropic disturbance) factors on breeding behaviour and performance of African Penguins on Bird Island, Algoa Bay, during peak breeding (March - July) in 2013. While sexual dimorphism in African Penguins is subtle, the colony-specific discriminant functions presented here provide an accurate sexing tool when only bill and flipper length are available. Despite the premise that selection of a large, high-quality mate in this longlived, monogamous seabird governs lifetime fitness, only low levels of assortative mating were found, and this only for earlier breeders, when larger females (but not males) bred. The 2013 season was a particularly successful one, coinciding with above-average sardine and anchovy abundance, and almost 80 percent of monitored nests were double-brooded, with very low levels of mortality. A- and B-chicks of double broods and singleton chicks grew at similar rates and exhibited similar body condition indices. In these conditions, chick developmental rates were independent of parental size, assortment or provisioning behaviour. Females raising a double brood were significantly lighter and in poorer body condition than those raising a single chick, although the same trend was not evident in males. Offspring sex ratio in 2013 (2.27:1) favoured male chicks, suggesting that there is potential to over-produce the larger sex when resources are plentiful. Peak nest arrival and departure times of parents did not change over the course of monitored breeding attempts (March-June), nor were they different for disturbed and undisturbed nests or for a single or double brood. The increase in CCTV-observed provisioning rate as chicks grew larger was best explained by brood size, at-sea chlorophyll a concentration, and maximum air temperature, but was unrelated to parental morphology or assortative index. Importantly, parental absenteeism commenced earlier and was markedly greater in nests frequently handled by researchers than in undisturbed nests. Both the time spent together by parents, and absenteeism were measurably affected by maximum afternoon air temperatures, the effects of which are expected to be exacerbated by poorer foraging conditions and climate change. A third of manually-monitored nests shared chick-guarding duties unequally, although this phenomenon was independent of parental sex or morphology. The adaptive benefits of mating patterns and division of labour during chick-rearing may only become apparent in a year of below-average food availability and it is highly recommended that this study be repeated in a year of scarce food resources. These findings augment past foraging ecology studies and demonstrate that investigator disturbance and environmental conditions can affect the nesting behaviour of this highly threatened seabird.
- Full Text:
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