A comparative study of the Grey-headed Sparrow (Passer griseus L) and the House Sparrow (Passer domesticus L) in Malawi
- Authors: Nhlane, Martin Edwin Darwin
- Date: 1997
- Subjects: English sparrow Sparrows
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5654 , http://hdl.handle.net/10962/d1005337
- Description: The House Sparrow Passer domesticus, an introduced species, and the Grey-headed Sparrow Passer griseus, an indigenous species, are sympatric in Malawi. Their distribution in the country and any possible interactions were studied, principally in southern Malawi. A morphological analysis of museum specimens confirmed that grey-headed sparrows in Malawi belong to the Northern Grey-headed Sparrow Passer griseus as distinct from the Southern Grey-headed Sparrow Passer diffusus. This species was widely distributed in the, country in association with human dwellings, both in rural areas as well as urban centres. In the northern region Greyheaded Sparrows were more abundant in the urban centres than rural areas, but in the central and southern regions numbers in the rural and urban areas were more or less the same. In Blantyre City, where they are in sympatry with the House Sparrow, they were found in the low density and industrial areas and were absent from the high density areas. The House Sparrow, arrived in Malawi in 1967 at Chileka in the southern region. Since then it has spread northwards, moving from the southern to the central and northern regions. House Sparrow numbers were found to be progressively larger in the southern region and lowest in the northern region. House Sparrows were found at sites where food was readily available, as in the immediate vicinity of houses. In the central and northern regions they were restricted mainly to urban areas. In the southern region, they occur both in rural and urban areas, probably as a reflection of the larger period of colonization in the south. In the northern region their movement has apparently been restricted by geographical barriers. In Blantyre City Grey-headed Sparrows preferred areas where tree density was high and house density was low, while House Sparrows preferred areas where house density was high and tree density was low. There was a positive correlation between Greyheaded Sparrow numbers and tree density and a negative correlation with house density. House Sparrow abundance was negatively correlated with tree density and positively correlated with house density. Grey-headed Sparrows bred in the rainy season, whereas House Sparrows bred throughout the year. There were differences in nest site selection: Grey-headed Sparrows used artificial structures such as fencing poles, and wooden telephone or electricity poles. The House Sparrow used mostly buildings and nested in crevices, holes in walls and between the walls and rafters. Nest height also differed- Grey-headed Sparrows nested at heights ranging from 1 - 8 m while House Sparrow nests were at heights of 1 - 5 m. Moult data suggests that although the House Sparrows breed throughout the year, they moult at a particular time of the year when breeding is less common. Grey-headed Sparrows were found to moult mainly from May to September in southern Africa and from June to September in central Africa. In both cases the breeding season extends over a similar period from about October to April/May of the following year. Peak moult periods differed between the House Sparrows and Grey-headed Sparrows. House Sparrows moulted mainly in the first half of the year, and Greyheaded Sparrows in the second six months. The clutch sizes of the two species were similar (mean 3.9 eggs for the House Sparrow and 3.4 for the Grey-headed Sparrow). The clutch size of the House Sparrow varied seasonally and was larger from November to May. The average incubation period for the House Sparrow was 11.5 days and the fledging period 15.4 days. The Grey-headed Sparrow fledging period was 14.7 days. Chick mortality of the House Sparrow at Chikunda farm was attributed to starvation resulting from brood reduction, abandonment, predation, low birth weight, accidental deaths and parasitism by fly larvae. Both Grey-headed and House Sparrows fed their young on insect food. Male House Sparrows fed actively initially, but their contribution declined from about day five onwards. In the Grey-headed Sparrow, both parents fed their young equally throughout the nestling period. House Sparrows fed on the ground near houses; Grey-headed Sparrows fed both on the ground away from houses and in tree canopies. The Grey-headed Sparrow walked as it fed on the ground as opposed to the House sparrow which hopped. Grey-headed Sparrows fed mainly as pairs and singletons while House Sparrows fed as family groups. Larger feeding groups of Grey-headed Sparrows were seen in the northern region at areas where food was plentiful. Where the two sparrows were seen feeding together, there was no direct competition for food. Where individual distance was violated; male House Sparrows displaced Grey-headed Sparrows which landed too close to them. Overall it appears that the distribution of the two species is determined more by their responses to habitat conditions than by interspecific interactions.
- Full Text:
- Date Issued: 1997
- Authors: Nhlane, Martin Edwin Darwin
- Date: 1997
- Subjects: English sparrow Sparrows
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5654 , http://hdl.handle.net/10962/d1005337
- Description: The House Sparrow Passer domesticus, an introduced species, and the Grey-headed Sparrow Passer griseus, an indigenous species, are sympatric in Malawi. Their distribution in the country and any possible interactions were studied, principally in southern Malawi. A morphological analysis of museum specimens confirmed that grey-headed sparrows in Malawi belong to the Northern Grey-headed Sparrow Passer griseus as distinct from the Southern Grey-headed Sparrow Passer diffusus. This species was widely distributed in the, country in association with human dwellings, both in rural areas as well as urban centres. In the northern region Greyheaded Sparrows were more abundant in the urban centres than rural areas, but in the central and southern regions numbers in the rural and urban areas were more or less the same. In Blantyre City, where they are in sympatry with the House Sparrow, they were found in the low density and industrial areas and were absent from the high density areas. The House Sparrow, arrived in Malawi in 1967 at Chileka in the southern region. Since then it has spread northwards, moving from the southern to the central and northern regions. House Sparrow numbers were found to be progressively larger in the southern region and lowest in the northern region. House Sparrows were found at sites where food was readily available, as in the immediate vicinity of houses. In the central and northern regions they were restricted mainly to urban areas. In the southern region, they occur both in rural and urban areas, probably as a reflection of the larger period of colonization in the south. In the northern region their movement has apparently been restricted by geographical barriers. In Blantyre City Grey-headed Sparrows preferred areas where tree density was high and house density was low, while House Sparrows preferred areas where house density was high and tree density was low. There was a positive correlation between Greyheaded Sparrow numbers and tree density and a negative correlation with house density. House Sparrow abundance was negatively correlated with tree density and positively correlated with house density. Grey-headed Sparrows bred in the rainy season, whereas House Sparrows bred throughout the year. There were differences in nest site selection: Grey-headed Sparrows used artificial structures such as fencing poles, and wooden telephone or electricity poles. The House Sparrow used mostly buildings and nested in crevices, holes in walls and between the walls and rafters. Nest height also differed- Grey-headed Sparrows nested at heights ranging from 1 - 8 m while House Sparrow nests were at heights of 1 - 5 m. Moult data suggests that although the House Sparrows breed throughout the year, they moult at a particular time of the year when breeding is less common. Grey-headed Sparrows were found to moult mainly from May to September in southern Africa and from June to September in central Africa. In both cases the breeding season extends over a similar period from about October to April/May of the following year. Peak moult periods differed between the House Sparrows and Grey-headed Sparrows. House Sparrows moulted mainly in the first half of the year, and Greyheaded Sparrows in the second six months. The clutch sizes of the two species were similar (mean 3.9 eggs for the House Sparrow and 3.4 for the Grey-headed Sparrow). The clutch size of the House Sparrow varied seasonally and was larger from November to May. The average incubation period for the House Sparrow was 11.5 days and the fledging period 15.4 days. The Grey-headed Sparrow fledging period was 14.7 days. Chick mortality of the House Sparrow at Chikunda farm was attributed to starvation resulting from brood reduction, abandonment, predation, low birth weight, accidental deaths and parasitism by fly larvae. Both Grey-headed and House Sparrows fed their young on insect food. Male House Sparrows fed actively initially, but their contribution declined from about day five onwards. In the Grey-headed Sparrow, both parents fed their young equally throughout the nestling period. House Sparrows fed on the ground near houses; Grey-headed Sparrows fed both on the ground away from houses and in tree canopies. The Grey-headed Sparrow walked as it fed on the ground as opposed to the House sparrow which hopped. Grey-headed Sparrows fed mainly as pairs and singletons while House Sparrows fed as family groups. Larger feeding groups of Grey-headed Sparrows were seen in the northern region at areas where food was plentiful. Where the two sparrows were seen feeding together, there was no direct competition for food. Where individual distance was violated; male House Sparrows displaced Grey-headed Sparrows which landed too close to them. Overall it appears that the distribution of the two species is determined more by their responses to habitat conditions than by interspecific interactions.
- Full Text:
- Date Issued: 1997
Exploitation of the bait organism Upogebia africana (Crustacea: Anomura) in the Knysna estuary
- Authors: Cretchley, Robyn
- Date: 1997
- Subjects: Upogebia africana -- South Africa -- Knysna , Mud shrimps -- South Africa -- Knysna , Fishing baits -- South Africa -- Knysna
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5691 , http://hdl.handle.net/10962/d1005377 , Upogebia africana -- South Africa -- Knysna , Mud shrimps -- South Africa -- Knysna , Fishing baits -- South Africa -- Knysna
- Description: In South African estuaries the anomuran mud prawn, Upogebia africana (Ortmann), is the main organism exploited as bait by recreational and subsistence anglers. In the Knysna estuary, three groups of bait collectors were identified on the basis of their source of income: subsistence fishermen who rely on bait-collecting and fishing for their income, supplementary anglers who catch fish to supplement their income and leisure anglers who draw no income from fishing. Two groups were identified based on the methods used: lelsure anglers who collect bait using a prawn pusher or pump and fish using a fishing rod and tackle, and non-leisure anglers who collect mud prawns using tin cans and fish with handlines. The hypothesis was erected that bait-collecting does not affect the U africana populations in the Knysna estuary. The approach adopted was to assess the distribution, density, population structure and reproductive patterns of the bait stock and to estimate the intensity of bait collection, to test the validity of the hypothesis. U africana has an extensive distribution, occupying 62 % of the available intertidal area of the Knysna estuary. Mud prawns have a broad intertidal distribution from the high water level (Spartina zone) to the shallow subtidal. The density, biomass and population structure of the mud prawns vary significantly with distance up the estuary and with tidal height on each shore. The Invertebrate Reserve supports very low densities of U africana (x = 11.7 m⁻²), whereas a relatively inaccessible centre bank in the middle reaches of the estuary appears to be a very effective natural mud prawn reserve as it supports the highest densities (x = 176.5m⁻²). The mud prawn stock of the estuary is estimated to be 2.19 x 108 prawns (82.7 tonnes dry mass). The numbers of bait collectors present per mudbank is highest on public holidays (x = 34) and higher during the summer holidays (x = 16) than during the winter (x = 4). A total of 1.858 x 106 U africana or 700.53 kg (dry mass) are removed by bait collectors annually from the 6 popular bait-collecting sites studied. This represents 8.49 % of the mud prawn stocks at these sites and 0.85 % of the entire estuary stock. 85 % of the U africana taken as bait annually, is removed by the 77.12 % of bait collectors who are non-leisure anglers. Recreational or leisure anglers are responsible for removing 14.2 %. The reproduction of female U africana in the Knysna estuary is seasonal and occurs from late July to April. There is evidence that this breeding season consists of two merging breeding cycles. The largest percentage (63 %) of ovigerous females is found in the middle reaches of the Knysna estuary at the Oyster Bank where the highest numbers of stage 1 larvae (165 m⁻³) are released. Larvae were exported from the estuary on the crepuscular ebb tide with peaks in abundance of nearly 1500 m⁻³ in November 1995 and January 1996. Although numbers caught are not significantly higher, larvae are nearly twice as abundant on crepuscular neap tides following a new moon (waxing quarter) than on those following a full moon (waning quarter). Recruitment of juvenile U africana to populations in the estuary was highest in December (45 m⁻²) and decreased over the summer. The highest numbers of recruits (31 m⁻²) were found at those sites closest to the mouth or on the main channel. Recruitment to the Leisure Isle and Thesens sites which are intensively exploited by bait collectors is high (20 - 32 m⁻²). The number of juveniles recruiting to U africana populations is estimated to be four times as high as the numbers of mud prawns taken by bait collectors. Legal methods of bait collection used in the Knysna estuary by the majority of anglers appear to cause minimal sediment disturbance and are not likely to affect the associated infauna. Illegal bait-collecting methods are however highly destructive and must be prohibited. It is concluded that the null hypothesis is acceptable, as mud prawn stocks of the Knysna estuary are not over-exploited and appear to be naturally regulated. The recruitment rate of juvenile U africana is estimated to be sufficiently high to sustain the present levels of exploitation.
- Full Text:
- Date Issued: 1997
- Authors: Cretchley, Robyn
- Date: 1997
- Subjects: Upogebia africana -- South Africa -- Knysna , Mud shrimps -- South Africa -- Knysna , Fishing baits -- South Africa -- Knysna
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5691 , http://hdl.handle.net/10962/d1005377 , Upogebia africana -- South Africa -- Knysna , Mud shrimps -- South Africa -- Knysna , Fishing baits -- South Africa -- Knysna
- Description: In South African estuaries the anomuran mud prawn, Upogebia africana (Ortmann), is the main organism exploited as bait by recreational and subsistence anglers. In the Knysna estuary, three groups of bait collectors were identified on the basis of their source of income: subsistence fishermen who rely on bait-collecting and fishing for their income, supplementary anglers who catch fish to supplement their income and leisure anglers who draw no income from fishing. Two groups were identified based on the methods used: lelsure anglers who collect bait using a prawn pusher or pump and fish using a fishing rod and tackle, and non-leisure anglers who collect mud prawns using tin cans and fish with handlines. The hypothesis was erected that bait-collecting does not affect the U africana populations in the Knysna estuary. The approach adopted was to assess the distribution, density, population structure and reproductive patterns of the bait stock and to estimate the intensity of bait collection, to test the validity of the hypothesis. U africana has an extensive distribution, occupying 62 % of the available intertidal area of the Knysna estuary. Mud prawns have a broad intertidal distribution from the high water level (Spartina zone) to the shallow subtidal. The density, biomass and population structure of the mud prawns vary significantly with distance up the estuary and with tidal height on each shore. The Invertebrate Reserve supports very low densities of U africana (x = 11.7 m⁻²), whereas a relatively inaccessible centre bank in the middle reaches of the estuary appears to be a very effective natural mud prawn reserve as it supports the highest densities (x = 176.5m⁻²). The mud prawn stock of the estuary is estimated to be 2.19 x 108 prawns (82.7 tonnes dry mass). The numbers of bait collectors present per mudbank is highest on public holidays (x = 34) and higher during the summer holidays (x = 16) than during the winter (x = 4). A total of 1.858 x 106 U africana or 700.53 kg (dry mass) are removed by bait collectors annually from the 6 popular bait-collecting sites studied. This represents 8.49 % of the mud prawn stocks at these sites and 0.85 % of the entire estuary stock. 85 % of the U africana taken as bait annually, is removed by the 77.12 % of bait collectors who are non-leisure anglers. Recreational or leisure anglers are responsible for removing 14.2 %. The reproduction of female U africana in the Knysna estuary is seasonal and occurs from late July to April. There is evidence that this breeding season consists of two merging breeding cycles. The largest percentage (63 %) of ovigerous females is found in the middle reaches of the Knysna estuary at the Oyster Bank where the highest numbers of stage 1 larvae (165 m⁻³) are released. Larvae were exported from the estuary on the crepuscular ebb tide with peaks in abundance of nearly 1500 m⁻³ in November 1995 and January 1996. Although numbers caught are not significantly higher, larvae are nearly twice as abundant on crepuscular neap tides following a new moon (waxing quarter) than on those following a full moon (waning quarter). Recruitment of juvenile U africana to populations in the estuary was highest in December (45 m⁻²) and decreased over the summer. The highest numbers of recruits (31 m⁻²) were found at those sites closest to the mouth or on the main channel. Recruitment to the Leisure Isle and Thesens sites which are intensively exploited by bait collectors is high (20 - 32 m⁻²). The number of juveniles recruiting to U africana populations is estimated to be four times as high as the numbers of mud prawns taken by bait collectors. Legal methods of bait collection used in the Knysna estuary by the majority of anglers appear to cause minimal sediment disturbance and are not likely to affect the associated infauna. Illegal bait-collecting methods are however highly destructive and must be prohibited. It is concluded that the null hypothesis is acceptable, as mud prawn stocks of the Knysna estuary are not over-exploited and appear to be naturally regulated. The recruitment rate of juvenile U africana is estimated to be sufficiently high to sustain the present levels of exploitation.
- Full Text:
- Date Issued: 1997
Extracellular digestion in two intertidal mussels and the role played by their gut bacteria
- Authors: Simon, Carol Anne
- Date: 1997
- Subjects: Perna -- Digestive organs , Mussels -- Digestive organs , Spirochaeta
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5692 , http://hdl.handle.net/10962/d1005378 , Perna -- Digestive organs , Mussels -- Digestive organs , Spirochaeta
- Description: The intertidal mussels. Perna perna and Choromytilus meridionalis co-occur on the southern coast of South Africa. Mussels ingest mixtures of bacteria. phytoplankton. zooplankton and detritus. with proportions varying according to availability. These bivalves filter similar-sized particles. which might result in interspecific competition. Carbohydrate-digesting enzymes of the mussels and their enteric bacteria. and bacteriolytic enzymes of the mussels were therefore examined to elucidate feeding ecology in these animals at an enzymatic level. Style enzymes of both species digested the storage carbohydrates amylose glycogen and laminarin, and the structural carbohydrate carboxymethyl cellulose (CMC). Differential rates of digestion of these carbohydrates suggests that Perna perna relies more on plankton (and possibly bacteria) than on detritus for food while Choromytilus meridionalis relies equally on all components of the seston. There may therefore be some degree of resource partitioning of the seston. The styles of P.perna had a lower specific enzyme activity. but higher protein content than those of C.meridionalis. P.perna could therefore release more glucose from a given concentration of substrate than C.meridionalis. The gut contents and tissue were examined microscopically to determine where the bacterial colonisation sites were. Bacteria were associated primarily with the gut contents but not the gut walls. of both species. The styles housed spirochaete bacteria (Crislispira sp), Perna perna housed large numbers of one species. while Choromytilus meridionalis had lower numbers of two species. Levels of infection differed between species and localities. Enteric (but not style) bacteria of Perna perna and Choromytilus meridionalis always digested the same carbohydrates as the mussels as well as the structural carbohydrates mannan and fucoidan. Activity was erratic on the structural compounds, carageenin and xylan, and absent on alginic acid or inulin. Activity on the storage carbohydrates by enteric bacteria from C.meridionalis was higher than by those from P.perna. This is probably related to the larger bacterial populations housed by C.meridionalis than by P.perna. Bacteriolytic activity by the digestive enzymes of Pema perna was higher than for Choromytilus. In P.perna it was due to a combination of different enzymes one of which is a true lysozyme. C.meridionalis did not produce a true lysozyme. Enzymes produced by the mussels and their enteric bacteria allow the mussels to utilise all components of the seston. Low endogenous enzyme activity by Choromytilus meridionalis, coupled with the high activity by its enteric bacteria, suggests that they rely more on bacterial activity to meet their dietary requirements than does Perna perna. The ability of enteric bacteria to digest carbohydrates which the mussels cannot indicates that the bacteria are endosymbiotic, although the sporadic nature of activity of some of the enzymes, and the fact that bacteria are associated with the gut contents, indicates that the relationship is only incidental.
- Full Text:
- Date Issued: 1997
- Authors: Simon, Carol Anne
- Date: 1997
- Subjects: Perna -- Digestive organs , Mussels -- Digestive organs , Spirochaeta
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5692 , http://hdl.handle.net/10962/d1005378 , Perna -- Digestive organs , Mussels -- Digestive organs , Spirochaeta
- Description: The intertidal mussels. Perna perna and Choromytilus meridionalis co-occur on the southern coast of South Africa. Mussels ingest mixtures of bacteria. phytoplankton. zooplankton and detritus. with proportions varying according to availability. These bivalves filter similar-sized particles. which might result in interspecific competition. Carbohydrate-digesting enzymes of the mussels and their enteric bacteria. and bacteriolytic enzymes of the mussels were therefore examined to elucidate feeding ecology in these animals at an enzymatic level. Style enzymes of both species digested the storage carbohydrates amylose glycogen and laminarin, and the structural carbohydrate carboxymethyl cellulose (CMC). Differential rates of digestion of these carbohydrates suggests that Perna perna relies more on plankton (and possibly bacteria) than on detritus for food while Choromytilus meridionalis relies equally on all components of the seston. There may therefore be some degree of resource partitioning of the seston. The styles of P.perna had a lower specific enzyme activity. but higher protein content than those of C.meridionalis. P.perna could therefore release more glucose from a given concentration of substrate than C.meridionalis. The gut contents and tissue were examined microscopically to determine where the bacterial colonisation sites were. Bacteria were associated primarily with the gut contents but not the gut walls. of both species. The styles housed spirochaete bacteria (Crislispira sp), Perna perna housed large numbers of one species. while Choromytilus meridionalis had lower numbers of two species. Levels of infection differed between species and localities. Enteric (but not style) bacteria of Perna perna and Choromytilus meridionalis always digested the same carbohydrates as the mussels as well as the structural carbohydrates mannan and fucoidan. Activity was erratic on the structural compounds, carageenin and xylan, and absent on alginic acid or inulin. Activity on the storage carbohydrates by enteric bacteria from C.meridionalis was higher than by those from P.perna. This is probably related to the larger bacterial populations housed by C.meridionalis than by P.perna. Bacteriolytic activity by the digestive enzymes of Pema perna was higher than for Choromytilus. In P.perna it was due to a combination of different enzymes one of which is a true lysozyme. C.meridionalis did not produce a true lysozyme. Enzymes produced by the mussels and their enteric bacteria allow the mussels to utilise all components of the seston. Low endogenous enzyme activity by Choromytilus meridionalis, coupled with the high activity by its enteric bacteria, suggests that they rely more on bacterial activity to meet their dietary requirements than does Perna perna. The ability of enteric bacteria to digest carbohydrates which the mussels cannot indicates that the bacteria are endosymbiotic, although the sporadic nature of activity of some of the enzymes, and the fact that bacteria are associated with the gut contents, indicates that the relationship is only incidental.
- Full Text:
- Date Issued: 1997
Macroinvertebrate community and species responses to chlorinated sewage effluent in the Umsunduze and Umbilo rivers, Kwa Zulu-Natal, South Africa
- Authors: Williams, Margot Lluttrell
- Date: 1997
- Subjects: Sewage -- Environmental aspects -- South Africa -- KwaZulu-Natal , Sewage -- Purification -- Chlorination -- South Africa -- KwaZulu-Natal , Baetis -- South Africa -- KwaZulu-Natal , Mayflies -- South Africa -- KwaZulu-Natal , Chlorine -- Environmental aspects -- South Africa -- KwaZulu-Natal
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5727 , http://hdl.handle.net/10962/d1005413 , Sewage -- Environmental aspects -- South Africa -- KwaZulu-Natal , Sewage -- Purification -- Chlorination -- South Africa -- KwaZulu-Natal , Baetis -- South Africa -- KwaZulu-Natal , Mayflies -- South Africa -- KwaZulu-Natal , Chlorine -- Environmental aspects -- South Africa -- KwaZulu-Natal
- Description: Chlorine has a wide variety of applications in water treatment. Because of its disinfectant efficacy, it is used world wide for the treatment of potable water, sewage, swimming pools and for the control of nuisance organisms in cooling towers. A problem arises when such chlorinated water enters the natural environment, as chlorine's greatest advantage, i.e. its germicidal capacity, becomes its greatest disadvantage. In particular, the discharge of heated, chlorinated water from cooling towers and chlorinated, treated sewage into rivers have severe consequences for the riverine flora and fauna. This study focused on the effects of chlorinated, treated sewage effluent on the community structure of benthic macroinvertebrates in two rivers in KwaZulu-Natal viz. the Umsunduze River in the Pietermaritzburg area, and the Umbilo River in the Durban area. The study was conducted in three phases. The first two phases comprised a toxicological investigation of the effects of chlorine on a selected riverine macroinvertebrate, and the third phase comprised an ecotoxicological investigation of the effects of chlorinated treated sewage on benthic macroinvertebrate community structure. The first phase of the study involved the development of an artificial stream system which would be suitable for determining the response of a selected macroinvertebrate species to chlorine. Chlorine is both reactive and volatile, so this necessitated the development of a specialised flow-through system with apparatus which would allow continuous dosing of a sodium hypochlorite solution. The system was set up at the Process Evaluation Facility at Wiggins Waterworks, Durban, where raw water from lnanda Dam was used. The second phase involved the use of this artificial stream system to conduct acute 96 h toxicity tests. Baetid mayfly nymphs (Baetis harrisoni Barnard) were selected as the test organisms after a preliminary investigation found them to be suitable for survival under laboratory conditions. For comparative purposes, tests were run first on B. harrisoni from a relatively uncontaminated stream in a residential area of Westville, then on specimens from the severely impacted Umbilo River. The LC₅₀ of chlorine for organisms from both sources was found to be in the region of 0.004 mg/l (free chlorine). This value was well below the general effluent standard of 0.1 mg/l in effect at the time. The recommended acute environmental guideline is 0.001 mg/l. The third phase of the study involved field validation of the toxicity test results. It was hypothesised that since the LC₅₀ for free chlorine was 0.004 mg/l, B. harrisoni would not be found downstream from a point source of chlorinated effluent where the concentration of free chlorine ranged from 0.06 to 0.2 mg/l, and that the macroinvertebrate community structure would also be altered. In order to test these hypotheses, benthic macro invertebrate community structure was investigated at several sites up- and downstream from the outlets of the Darvill Wastewater Works in the Umsunduze River and the Umbilo Sewage Purification Works in the Umbilo River. In addition, in order to differentiate between the effects of chlorinated and unchlorinated treated sewage, a section of the Umbilo River (upstream from the chlorinated discharge) was exposed to unchlorinated, treated sewage. In this way, a limited "before and after" sewage and an "upstream and downstream" from sewage investigation could be carried out. Organisms were collected from riffles (and from pools in the Umbilo River) and the samples were then sorted and organisms were identified to species level, where possible, otherwise to genus or family. Changes in community composition were shown graphically as pie charts of relative proportions of organisms found at each site, graphs of the average number of taxa at each site; and graphs of the average number of individuals at each site; Data from the Umbilo River were also analysed using TWINSPAN (Two-way indicator species analysis). In both the Umsunduze and the Umbilo rivers, the deleterious effects of the chlorinated effluent were clearly evident. At Umsunduze Site 3 and Umbilo Site 5 (both immediately downstream from the chlorinated effluent) both the number of taxa and number of individuals were substantially reduced, sometimes to zero. Where organisms were found at the next sites downstream (Sites 4 and 6 respectively), the samples were dominated by Chironomus. In contrast, the unchlorinated effluent in the Umbilo River caused very little difference in community structure. As predicted, B. harrisoni was not found in downstream samples in which chlorine was present, yet appeared to be relatively unaffected by the unchlorinated effluent, suggesting that chlorine, rather than the effluent was responsible for its absence at downstream sites. In conclusion, it would appear that while treated sewage effluent certainly causes changes in macroinvertebrate community structure, chlorination of this effluent leads to large scale destruction of the riverine community. This in turn delays the recovery process of the river, rendering a longer stretch unfit for use. The consequences of this delayed recovery are that the failure to meet the water quality requirements of the natural environment results in those of the other water users (agriculture, industry, domestic and recreation) not being met. This reduces the natural capacity of the riverine community to process organic waste and recover from the discharge of sewage effluent. Chlorination increases the distance of impaired water quality and environmental integrity which result from organically enriched treated sewage effluent. The results of the study indicated that the draft water quality guidelines for aquatic ecosystems, derived from inadequate data, and calculated with a safety factor, were the correct order of magnitude. The approach followed in the study will be useful in the development and refinement of water quality guidelines for aquatic ecosystems.
- Full Text:
- Date Issued: 1997
- Authors: Williams, Margot Lluttrell
- Date: 1997
- Subjects: Sewage -- Environmental aspects -- South Africa -- KwaZulu-Natal , Sewage -- Purification -- Chlorination -- South Africa -- KwaZulu-Natal , Baetis -- South Africa -- KwaZulu-Natal , Mayflies -- South Africa -- KwaZulu-Natal , Chlorine -- Environmental aspects -- South Africa -- KwaZulu-Natal
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5727 , http://hdl.handle.net/10962/d1005413 , Sewage -- Environmental aspects -- South Africa -- KwaZulu-Natal , Sewage -- Purification -- Chlorination -- South Africa -- KwaZulu-Natal , Baetis -- South Africa -- KwaZulu-Natal , Mayflies -- South Africa -- KwaZulu-Natal , Chlorine -- Environmental aspects -- South Africa -- KwaZulu-Natal
- Description: Chlorine has a wide variety of applications in water treatment. Because of its disinfectant efficacy, it is used world wide for the treatment of potable water, sewage, swimming pools and for the control of nuisance organisms in cooling towers. A problem arises when such chlorinated water enters the natural environment, as chlorine's greatest advantage, i.e. its germicidal capacity, becomes its greatest disadvantage. In particular, the discharge of heated, chlorinated water from cooling towers and chlorinated, treated sewage into rivers have severe consequences for the riverine flora and fauna. This study focused on the effects of chlorinated, treated sewage effluent on the community structure of benthic macroinvertebrates in two rivers in KwaZulu-Natal viz. the Umsunduze River in the Pietermaritzburg area, and the Umbilo River in the Durban area. The study was conducted in three phases. The first two phases comprised a toxicological investigation of the effects of chlorine on a selected riverine macroinvertebrate, and the third phase comprised an ecotoxicological investigation of the effects of chlorinated treated sewage on benthic macroinvertebrate community structure. The first phase of the study involved the development of an artificial stream system which would be suitable for determining the response of a selected macroinvertebrate species to chlorine. Chlorine is both reactive and volatile, so this necessitated the development of a specialised flow-through system with apparatus which would allow continuous dosing of a sodium hypochlorite solution. The system was set up at the Process Evaluation Facility at Wiggins Waterworks, Durban, where raw water from lnanda Dam was used. The second phase involved the use of this artificial stream system to conduct acute 96 h toxicity tests. Baetid mayfly nymphs (Baetis harrisoni Barnard) were selected as the test organisms after a preliminary investigation found them to be suitable for survival under laboratory conditions. For comparative purposes, tests were run first on B. harrisoni from a relatively uncontaminated stream in a residential area of Westville, then on specimens from the severely impacted Umbilo River. The LC₅₀ of chlorine for organisms from both sources was found to be in the region of 0.004 mg/l (free chlorine). This value was well below the general effluent standard of 0.1 mg/l in effect at the time. The recommended acute environmental guideline is 0.001 mg/l. The third phase of the study involved field validation of the toxicity test results. It was hypothesised that since the LC₅₀ for free chlorine was 0.004 mg/l, B. harrisoni would not be found downstream from a point source of chlorinated effluent where the concentration of free chlorine ranged from 0.06 to 0.2 mg/l, and that the macroinvertebrate community structure would also be altered. In order to test these hypotheses, benthic macro invertebrate community structure was investigated at several sites up- and downstream from the outlets of the Darvill Wastewater Works in the Umsunduze River and the Umbilo Sewage Purification Works in the Umbilo River. In addition, in order to differentiate between the effects of chlorinated and unchlorinated treated sewage, a section of the Umbilo River (upstream from the chlorinated discharge) was exposed to unchlorinated, treated sewage. In this way, a limited "before and after" sewage and an "upstream and downstream" from sewage investigation could be carried out. Organisms were collected from riffles (and from pools in the Umbilo River) and the samples were then sorted and organisms were identified to species level, where possible, otherwise to genus or family. Changes in community composition were shown graphically as pie charts of relative proportions of organisms found at each site, graphs of the average number of taxa at each site; and graphs of the average number of individuals at each site; Data from the Umbilo River were also analysed using TWINSPAN (Two-way indicator species analysis). In both the Umsunduze and the Umbilo rivers, the deleterious effects of the chlorinated effluent were clearly evident. At Umsunduze Site 3 and Umbilo Site 5 (both immediately downstream from the chlorinated effluent) both the number of taxa and number of individuals were substantially reduced, sometimes to zero. Where organisms were found at the next sites downstream (Sites 4 and 6 respectively), the samples were dominated by Chironomus. In contrast, the unchlorinated effluent in the Umbilo River caused very little difference in community structure. As predicted, B. harrisoni was not found in downstream samples in which chlorine was present, yet appeared to be relatively unaffected by the unchlorinated effluent, suggesting that chlorine, rather than the effluent was responsible for its absence at downstream sites. In conclusion, it would appear that while treated sewage effluent certainly causes changes in macroinvertebrate community structure, chlorination of this effluent leads to large scale destruction of the riverine community. This in turn delays the recovery process of the river, rendering a longer stretch unfit for use. The consequences of this delayed recovery are that the failure to meet the water quality requirements of the natural environment results in those of the other water users (agriculture, industry, domestic and recreation) not being met. This reduces the natural capacity of the riverine community to process organic waste and recover from the discharge of sewage effluent. Chlorination increases the distance of impaired water quality and environmental integrity which result from organically enriched treated sewage effluent. The results of the study indicated that the draft water quality guidelines for aquatic ecosystems, derived from inadequate data, and calculated with a safety factor, were the correct order of magnitude. The approach followed in the study will be useful in the development and refinement of water quality guidelines for aquatic ecosystems.
- Full Text:
- Date Issued: 1997
Parasitism and invasive species : an ecological study of mussel populations
- Calvo Ugarteburu, Miren Gurutze
- Authors: Calvo Ugarteburu, Miren Gurutze
- Date: 1997
- Subjects: Perna -- Parasites Mytilidae -- Parasites Perna -- Ecology Mytilidae -- Ecology
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5736 , http://hdl.handle.net/10962/d1005422
- Description: The Mediterranean mussel Mytilus galloprovincialis, was introduced accidentally to South Africa and has since become invasive. One possible explanation for the success of this species is that it has been released from the effects of parasites which affect the indigenous species of mussels. The aim of this work was to examine the parasitic load of different mussel populations along the southern African coast and to assess the effects of parasites on their hosts. A survey was done to quantify the levels of parasitism in indigenous populations of Perna perna from Southern Africa and of Mytilus galloprovincialis from South Africa (where it is exotic) and Spain (where it is indigenous). This survey was carried out at three different geographic scales: small scale (metres), studying the incidence of parasites within a mussel bed; medium scale (kilometres to tens of kilometres), comparing prevalences of infection among different localities; and large scale (hundreds to thousands of kilometres), examining infection rates along the Southern African coast, as well as the coast of the Basque Country (North Spain). During this survey metazoan parasites other than trematodes were rare. Four species of trematodes were commonly found infecting the indigenous mussel Perna perna. These were metacercariae of the genus Proctoeces, bucephalid sporocysts, encysted metacercariae on the labial palps and gelatinous cysts with metacercariae inside. A detailed description of Proctoeces is given; the other parasites are described elsewhere. No parasites were found in M. galloprovincialis either in South Africa or in Spain. This thesis concentrates on the study of prevalences of Proctoeces and bucephalid sporocysts and their effects on Perna perna. Infection rates with Proctoeces are highly dependent on the sex of the host, with more females than males being infected, and are also size dependent, though only for females. Identification of the sex of mussels infected with bucephalid sporocysts is often not possible since the sporocysts spread over the gonad and replace it. Prevalence of infection with this parasite also increases with the size of the host. To study the influence of Proctoeces and bucephalid sporocysts on the ecological fitness of Perna perna, their effects on survival and competitive ability were tested. The results showed significant negative effects. Both parasites significantly depressed condition but only after spawning, when the mussels were already stressed. In order to check for effects on host survival, the effects of both parasites on mortality rates, gaping behaviour and water loss of mussels exposed to air were also examined. Neither parasite affected mortality rate or gaping behaviour of Perna perna. Proctoeces did not affect the amount of water lost by mussels, but the bucephalid sporocysts did. Mussels infected with sporocysts lost significantly more water than non-infected individuals. This increase in water loss was not related to the gaping behaviour, but a test of the strength of the adductor muscles showed that less force was needed to open mussels with bucephalid sporocysts than non-infected mussels. This was not the case for mussels infected by Proctoeces. Weaker mussels will fail to seal the valves properly, resulting in an increase of water loss on exposure to air by evaporation. Another factor that will have an obvious effect on a population is the reproductive output of the animals. Histological sections of the gonad of infected and non-infected females were cut to study the effects of both parasites on reproduction. Statistical tests comparing the numbers and sizes of oocytes in females infected with Proctoeces and non-infected females showed no significant differences. However, bucephalid sporocysts have a dramatic effect on reproduction by castrating the host, leaving no trace of sex products. One of the major factors shaping the composition of a mussel bed is competition for space and food, with smaller mussels being at a competitive disadvantage. Thus, in order to examine effects of both parasites on the competitive ability of Perna perna, summer and winter growth rates were compared for infected and non-infected mussels. Proctoeces reduced growth both in summer and in winter whilst bucephalid sporocysts had no significant effect. Both growth and reproduction are important components of the energy budget of an animal, and each is affected by either Proctoeces or the bucephalid sporocysts. In an attempt to test if Perna perna compensates energetically for these negative effects, filtration rates and oxygen consumption of mussels with and without parasites were measured. Neither parasite had a significant effect on filtration rates or oxygen consumption of the host. All these results indicate that both Proctoeces and the bucephalid sporocysts have a detrimental effect on their host, and that the mussels do not compensate for these negative effects. There is neither an increase in filtration, nor a decrease in respiration to balance the energy lost to the parasite. The two parasites studied affect the host in different but complementary ways. The effects of both parasites are concentrated on those size classes of mussel which channel most energy into the portion of the energy budget affected by the parasite. Proctoeces affects growth only in the smaller individuals, which under normal conditions would put most energy into growth; and the bucephalid sporocysts castrate the bigger mussels, which would expend most energy on reproduction. By reducing growth rates of small mussels or castrating large mussels, these parasites effectively remove them from the breeding population and reduce their competitive abilities. These negative effects, together with the high prevalence of both parasites in Perna perna along the South African coast and their absence in Mytilus galloprovincialis, suggest that parasites may be an important reason for the success of Mytilus.
- Full Text:
- Date Issued: 1997
- Authors: Calvo Ugarteburu, Miren Gurutze
- Date: 1997
- Subjects: Perna -- Parasites Mytilidae -- Parasites Perna -- Ecology Mytilidae -- Ecology
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5736 , http://hdl.handle.net/10962/d1005422
- Description: The Mediterranean mussel Mytilus galloprovincialis, was introduced accidentally to South Africa and has since become invasive. One possible explanation for the success of this species is that it has been released from the effects of parasites which affect the indigenous species of mussels. The aim of this work was to examine the parasitic load of different mussel populations along the southern African coast and to assess the effects of parasites on their hosts. A survey was done to quantify the levels of parasitism in indigenous populations of Perna perna from Southern Africa and of Mytilus galloprovincialis from South Africa (where it is exotic) and Spain (where it is indigenous). This survey was carried out at three different geographic scales: small scale (metres), studying the incidence of parasites within a mussel bed; medium scale (kilometres to tens of kilometres), comparing prevalences of infection among different localities; and large scale (hundreds to thousands of kilometres), examining infection rates along the Southern African coast, as well as the coast of the Basque Country (North Spain). During this survey metazoan parasites other than trematodes were rare. Four species of trematodes were commonly found infecting the indigenous mussel Perna perna. These were metacercariae of the genus Proctoeces, bucephalid sporocysts, encysted metacercariae on the labial palps and gelatinous cysts with metacercariae inside. A detailed description of Proctoeces is given; the other parasites are described elsewhere. No parasites were found in M. galloprovincialis either in South Africa or in Spain. This thesis concentrates on the study of prevalences of Proctoeces and bucephalid sporocysts and their effects on Perna perna. Infection rates with Proctoeces are highly dependent on the sex of the host, with more females than males being infected, and are also size dependent, though only for females. Identification of the sex of mussels infected with bucephalid sporocysts is often not possible since the sporocysts spread over the gonad and replace it. Prevalence of infection with this parasite also increases with the size of the host. To study the influence of Proctoeces and bucephalid sporocysts on the ecological fitness of Perna perna, their effects on survival and competitive ability were tested. The results showed significant negative effects. Both parasites significantly depressed condition but only after spawning, when the mussels were already stressed. In order to check for effects on host survival, the effects of both parasites on mortality rates, gaping behaviour and water loss of mussels exposed to air were also examined. Neither parasite affected mortality rate or gaping behaviour of Perna perna. Proctoeces did not affect the amount of water lost by mussels, but the bucephalid sporocysts did. Mussels infected with sporocysts lost significantly more water than non-infected individuals. This increase in water loss was not related to the gaping behaviour, but a test of the strength of the adductor muscles showed that less force was needed to open mussels with bucephalid sporocysts than non-infected mussels. This was not the case for mussels infected by Proctoeces. Weaker mussels will fail to seal the valves properly, resulting in an increase of water loss on exposure to air by evaporation. Another factor that will have an obvious effect on a population is the reproductive output of the animals. Histological sections of the gonad of infected and non-infected females were cut to study the effects of both parasites on reproduction. Statistical tests comparing the numbers and sizes of oocytes in females infected with Proctoeces and non-infected females showed no significant differences. However, bucephalid sporocysts have a dramatic effect on reproduction by castrating the host, leaving no trace of sex products. One of the major factors shaping the composition of a mussel bed is competition for space and food, with smaller mussels being at a competitive disadvantage. Thus, in order to examine effects of both parasites on the competitive ability of Perna perna, summer and winter growth rates were compared for infected and non-infected mussels. Proctoeces reduced growth both in summer and in winter whilst bucephalid sporocysts had no significant effect. Both growth and reproduction are important components of the energy budget of an animal, and each is affected by either Proctoeces or the bucephalid sporocysts. In an attempt to test if Perna perna compensates energetically for these negative effects, filtration rates and oxygen consumption of mussels with and without parasites were measured. Neither parasite had a significant effect on filtration rates or oxygen consumption of the host. All these results indicate that both Proctoeces and the bucephalid sporocysts have a detrimental effect on their host, and that the mussels do not compensate for these negative effects. There is neither an increase in filtration, nor a decrease in respiration to balance the energy lost to the parasite. The two parasites studied affect the host in different but complementary ways. The effects of both parasites are concentrated on those size classes of mussel which channel most energy into the portion of the energy budget affected by the parasite. Proctoeces affects growth only in the smaller individuals, which under normal conditions would put most energy into growth; and the bucephalid sporocysts castrate the bigger mussels, which would expend most energy on reproduction. By reducing growth rates of small mussels or castrating large mussels, these parasites effectively remove them from the breeding population and reduce their competitive abilities. These negative effects, together with the high prevalence of both parasites in Perna perna along the South African coast and their absence in Mytilus galloprovincialis, suggest that parasites may be an important reason for the success of Mytilus.
- Full Text:
- Date Issued: 1997
Studies of the biology and ecology of the high shore South African limpet, Helcion pectunculus (Mollusca : patellogastropoda)
- Authors: Gray, David Richard
- Date: 1997
- Subjects: Mollusks -- South Africa Limpets -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5776 , http://hdl.handle.net/10962/d1005464
- Description: Investigations were carried out into aspects of the bio!ogy and ecology of Helcion pectunculus along the coait of South Africa. These included studies of the distribution, density and biomass of the limpet at six sites along the east coast plus one site on the west coast; the growth of H pectunculus on both west and east coasts; a comparison of the reproductive biology of the east and west coast populations; the foraging activity and feeding behaviour of H pectunculus and the driving forces behind the rhythmic behaviour of this limpet; the importance of the crevice environment in the biology and ecology of H pectunculus. Helcion pectunculus has a restricted zonation, with the majority of animals residing in crevices in the upper Balanoid zone during diurnallowtides, although individuals were occasionally found in the lower Balanoid zone on shores with a gently sloping aspect. This limpet occurs in higher densities (50-lO0 individuals/m2) on shores which have large numbers of crevices and boulders i.e. quartzitic sandstone shores. On most shores, the ratio of males to females differed significantly from a 1: 1 ratio with the highest ratio being obtained on the west .. coast (3 males: 1 female). At all sites, the populations of H pectunculus exhibited strong sexual dimorphism. Males and females were always found to differ in size, with individuals of < 20 mm shell length generally being male whilst limpets with a shell length of> 22 mm were generally female. Helcion pectunculus grows allometrically, increasing in height faster than length, which is expected of a high shore gastropod mollusc attempting to reduce evaporative water loss. Growth rates were similar on both east and west coasts regardless of the differing oceanographic conditions. The theoretical values of Lmax were also similar being 30.86 mm and 30.71 mm respectively. Micro-growth bands are laid down within its' shell which have the same periodicity as the tidal cycle and these enabled age estimates to be made. Younger individuals were male whilst older animals were female, suggesting that H pectunculus is a protandrous hermaphrodite. Histological examination proved, unequivocally, that this limpet undergoes a protandric sex change, changing from male to female when they are about 2 years old. Both east and west coast popUlations had a marked reproductive cycle, exhibiting two spawning periods a year, one in April and another in November. The possibility that the reproductive pattern exhibited is now phylogeneticallyconstrained is discussed. It is suggested that H pectunculus has evolved a reproductive cycle which will allow its planktonic larvae to utilise the valuable phytoplankton bloom food source whilst using onshore winds to ensure that larvae are not transported out to sea and lost. The number of foraging excursions carried out by individuals of H pectunculus was found to have a significant effect on Gonad Index and hence potential reproductive output. The activity pattern of H pectunculus varied depending upon micro-habitat; animals inhabiting both east and west facing rock surfaces are active during nocturnal low tides whilst animals on west facing rock surfaces are also active during daytime low tides whilst in the shade. Limpets travel further during foraging excursions in winter (X = 85.53 cm) than in either spring (x = 55.7 cm) or summer (X = 48.8 cm) and also during spring low tides (x = 89.8 cm) compared with neaps (x = 40.9 cm). This limpet exhibits rigid homing to a fixed scar within a-crevice and feeding excursions were found to cons.is.t. of three distinct phases, a rapid outward phase, a slower foraging phase and a rapid homeward phase. Foraging was always highly directional, with a mean vector which took limpets onto an area of the rockface with the highest microalgal biomass and also the smoothest rock surface. Helcion pectunculus exhibits a free-running endogenous rhythm of locomotor activity with both circadian and circatidal components and it is suggested that this rhythm plays a role in allowing the limpet to avoid unfavourable environmental conditions. The exogenous entrainment factor of this endogenous rhythm was the time of exposure to air in the field. There was found to be an organized distribution of limpets within crevices with smaller, younger limpets being towards the back of the crevice and larger, older limpets towards the crevice mouth. It is hypothesized that juvenile limpets of this species actively select and settle at the backs of crevices responding to chemical cues of adult conspecifics. The crevice refuge supplies the limpets with a stable and buffered environment with higher relative humidities (X = 72.3%) and lower rock surface temperatures (X = 19.7°C) than adjacent exposed rock surfaces (X = 64.5%; x = 22.9°C). Limpet body temperatures were significantly lower in crevic~refuges compared to limpets on exposed rock surfaces. Body temperatures never exceeded the rock surface temperatures. It is suggested that this is the result of morphological adaptations such as shell ornamentation and allometric growth. Light levels above 1000 J.1E.m-2.s-1 inhibited foraging activity in H. pectunculuswhilst limpets subjected to 30-50% shade foraged even during daytime lowtides. This limpet is one of the least tenacious (2.75 ± 0.13 kg.cm-2 ) of all South African limpets and the possibility that wave activity governs both the activity patterns and homing behaviour of this limpet is discussed. Limpets deprived of a crevice refuge experienced extremely high mortalities, with 45% of the limpets being lost during the first high tide period. A hypothetical model of the hierarchy of exogenous factors controlling limpet foraging activity is introduced and discussed in relation to the results of this study. Finally, it is suggested that in addition to the "migratory" and "non-migratory" groups of limpets present on southern African shores a third group of limpets seem to be present which may be classed as "specialized non-migratory" species. These are species that do not migrate, garden or aggressively fight off like conspecifics. They have overcome the competition for space and food on intertidal rocky shores by adapting to a particular habitat which is exclusive to them alone. From the combined results of these studies, it can be stated that H. pectunculus has adapted physiologically, morphologically and behaviourally to successfully survive the extreme conditions in the upper Balanoid zone.
- Full Text:
- Date Issued: 1997
- Authors: Gray, David Richard
- Date: 1997
- Subjects: Mollusks -- South Africa Limpets -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5776 , http://hdl.handle.net/10962/d1005464
- Description: Investigations were carried out into aspects of the bio!ogy and ecology of Helcion pectunculus along the coait of South Africa. These included studies of the distribution, density and biomass of the limpet at six sites along the east coast plus one site on the west coast; the growth of H pectunculus on both west and east coasts; a comparison of the reproductive biology of the east and west coast populations; the foraging activity and feeding behaviour of H pectunculus and the driving forces behind the rhythmic behaviour of this limpet; the importance of the crevice environment in the biology and ecology of H pectunculus. Helcion pectunculus has a restricted zonation, with the majority of animals residing in crevices in the upper Balanoid zone during diurnallowtides, although individuals were occasionally found in the lower Balanoid zone on shores with a gently sloping aspect. This limpet occurs in higher densities (50-lO0 individuals/m2) on shores which have large numbers of crevices and boulders i.e. quartzitic sandstone shores. On most shores, the ratio of males to females differed significantly from a 1: 1 ratio with the highest ratio being obtained on the west .. coast (3 males: 1 female). At all sites, the populations of H pectunculus exhibited strong sexual dimorphism. Males and females were always found to differ in size, with individuals of < 20 mm shell length generally being male whilst limpets with a shell length of> 22 mm were generally female. Helcion pectunculus grows allometrically, increasing in height faster than length, which is expected of a high shore gastropod mollusc attempting to reduce evaporative water loss. Growth rates were similar on both east and west coasts regardless of the differing oceanographic conditions. The theoretical values of Lmax were also similar being 30.86 mm and 30.71 mm respectively. Micro-growth bands are laid down within its' shell which have the same periodicity as the tidal cycle and these enabled age estimates to be made. Younger individuals were male whilst older animals were female, suggesting that H pectunculus is a protandrous hermaphrodite. Histological examination proved, unequivocally, that this limpet undergoes a protandric sex change, changing from male to female when they are about 2 years old. Both east and west coast popUlations had a marked reproductive cycle, exhibiting two spawning periods a year, one in April and another in November. The possibility that the reproductive pattern exhibited is now phylogeneticallyconstrained is discussed. It is suggested that H pectunculus has evolved a reproductive cycle which will allow its planktonic larvae to utilise the valuable phytoplankton bloom food source whilst using onshore winds to ensure that larvae are not transported out to sea and lost. The number of foraging excursions carried out by individuals of H pectunculus was found to have a significant effect on Gonad Index and hence potential reproductive output. The activity pattern of H pectunculus varied depending upon micro-habitat; animals inhabiting both east and west facing rock surfaces are active during nocturnal low tides whilst animals on west facing rock surfaces are also active during daytime low tides whilst in the shade. Limpets travel further during foraging excursions in winter (X = 85.53 cm) than in either spring (x = 55.7 cm) or summer (X = 48.8 cm) and also during spring low tides (x = 89.8 cm) compared with neaps (x = 40.9 cm). This limpet exhibits rigid homing to a fixed scar within a-crevice and feeding excursions were found to cons.is.t. of three distinct phases, a rapid outward phase, a slower foraging phase and a rapid homeward phase. Foraging was always highly directional, with a mean vector which took limpets onto an area of the rockface with the highest microalgal biomass and also the smoothest rock surface. Helcion pectunculus exhibits a free-running endogenous rhythm of locomotor activity with both circadian and circatidal components and it is suggested that this rhythm plays a role in allowing the limpet to avoid unfavourable environmental conditions. The exogenous entrainment factor of this endogenous rhythm was the time of exposure to air in the field. There was found to be an organized distribution of limpets within crevices with smaller, younger limpets being towards the back of the crevice and larger, older limpets towards the crevice mouth. It is hypothesized that juvenile limpets of this species actively select and settle at the backs of crevices responding to chemical cues of adult conspecifics. The crevice refuge supplies the limpets with a stable and buffered environment with higher relative humidities (X = 72.3%) and lower rock surface temperatures (X = 19.7°C) than adjacent exposed rock surfaces (X = 64.5%; x = 22.9°C). Limpet body temperatures were significantly lower in crevic~refuges compared to limpets on exposed rock surfaces. Body temperatures never exceeded the rock surface temperatures. It is suggested that this is the result of morphological adaptations such as shell ornamentation and allometric growth. Light levels above 1000 J.1E.m-2.s-1 inhibited foraging activity in H. pectunculuswhilst limpets subjected to 30-50% shade foraged even during daytime lowtides. This limpet is one of the least tenacious (2.75 ± 0.13 kg.cm-2 ) of all South African limpets and the possibility that wave activity governs both the activity patterns and homing behaviour of this limpet is discussed. Limpets deprived of a crevice refuge experienced extremely high mortalities, with 45% of the limpets being lost during the first high tide period. A hypothetical model of the hierarchy of exogenous factors controlling limpet foraging activity is introduced and discussed in relation to the results of this study. Finally, it is suggested that in addition to the "migratory" and "non-migratory" groups of limpets present on southern African shores a third group of limpets seem to be present which may be classed as "specialized non-migratory" species. These are species that do not migrate, garden or aggressively fight off like conspecifics. They have overcome the competition for space and food on intertidal rocky shores by adapting to a particular habitat which is exclusive to them alone. From the combined results of these studies, it can be stated that H. pectunculus has adapted physiologically, morphologically and behaviourally to successfully survive the extreme conditions in the upper Balanoid zone.
- Full Text:
- Date Issued: 1997
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