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Despite being the world’s largest batoid fish, manta rays have been the focus of very little research in the last century. Almost nothing is currently known about their ecology or distribution throughout the world’s oceans. Manta rays are considered to be near-threatened to vulnerable by the World Conservation Union’s Redlist of endangered species, yet many populations still face threats from target fishing for their fin cartilage, branchial filaments and meat. The taxonomic history of the genus Manta has been questionable and convoluted,with Manta having one of the most extensive generic and species synonymies of any living genus of cartilaginous fish. Currently this genus is considered to be monotypicwith a single recognized species, Manta birostris. Behavioural and morphological data collected during this study showed that there are two distinct, sympatric species of manta ray in Mozambique. This discovery has led to an examination of manta ray distribution worldwide, with the two species hypothesis receiving further support from study sites in the Indian and Pacific Oceans. This thesis examined large populations of both putative species in Mozambique. Development and application of a photographic-identification methodology allowed identification of individuals based on their unique ventral surface markings (spots). The ability to recognise individuals enabled many aspects of both populations to be examined. A total of 449 individual rays were identified of Manta birostris variant A and 101 individual rays were identified of Manta birostris variant B. Both populations exhibited a highly significant female bias in observed sex ratios of 1:3.5 and 1:7.4 respectively. The photographic identification approach was used in studies of population size and structure and to examine the reproductive ecology of these two distinct populations. Population estimates incorporating four years of re-sighting data on Manta birostris variant A and five years of data on Manta birostris variant B were conducted using the program MARK resulting in super population estimates of 890 and 600 individuals respectively. The region encompassing the study site was identified as a mating ground for Manta birostris variant A based on observations of mating events and fresh pectoral fin tip scars on female rays produced when the male rays bite onto the fins during copulation. The distribution of scars was highly biased, with 99% on the left pectoral fin, indicative of a strong lateralised behavioural trait in this species. No other elasmobranch has been reported to display behavioural lateralisation. The study region also acts as a birthing ground, with individuals giving birth in the summer after a gestation period of approximately one year. Reproductive periodicity in M. birostris variant A was most commonly biennial, but a few individuals were seen to be pregnant in consecutive years, confirming an annual ovulatory cycle. The production of a single pup appears to be the normal situation, although observations in the wild as well as during opportunistic dissections of individuals killed in fisheries revealed that two pups are conceived on occasion. Morphometric analysis of late-term foetus (M. birostris variant A) was contrasted with measurements taken from adult rays. These are the only detailed measurements on M. birostris variant A from the western Indian Ocean. Predatory scarring and bite injuries on individuals of both variant A and B were consistent with attacks from sharks of various sizes. The frequency and effect of these predatory injuries on the two populations were examined over a three-year period. Acoustic tags were attached to fourteen rays and their presence/absence at sites around the major inshore reefs was explored to evaluate how the rays utilised their environment on a temporal scale. Cleaning activity of both putative species of manta rays by small fishes occurred on these reefs and was considered to be the main reason why the rays were present at these sites. Parasite removal and wound healing were implicated as the benefits received. Host cleaner fish species partition the manta ray body to avoid interspecific competition while the two putative species appear to partition cleaning habitats, with very little overlap apparent. This research on manta rays is the first of its kind in African waters. Many aspects of the study have contributed to the limited baseline data currently available for this genus. The study additionally provided sufficient empirical evidence to warrant the separation of these two putative species of Manta and a full revision of the genus with the systematic examination of specimens throughout Manta’s range. The results of this study may be useful in this endeavour as diagnostic characteristics have been isolated that may help to differentiate members of this genus. The results of this research study are directly applicable to management strategies for both putative species of manta rays off the East Coast of Africa as well as populations worldwide and have highlighted the potential need for different conservation strategies

Despite being the world’s largest batoid fish, manta rays have been the focus of very little research in the last century. Almost nothing is currently known about their ecology or distribution throughout the world’s oceans. Manta rays are considered to be near-threatened to vulnerable by the World Conservation Union’s Redlist of endangered species, yet many populations still face threats from target fishing for their fin cartilage, branchial filaments and meat. The taxonomic history of the genus Manta has been questionable and convoluted,with Manta having one of the most extensive generic and species synonymies of any living genus of cartilaginous fish. Currently this genus is considered to be monotypicwith a single recognized species, Manta birostris. Behavioural and morphological data collected during this study showed that there are two distinct, sympatric species of manta ray in Mozambique. This discovery has led to an examination of manta ray distribution worldwide, with the two species hypothesis receiving further support from study sites in the Indian and Pacific Oceans. This thesis examined large populations of both putative species in Mozambique. Development and application of a photographic-identification methodology allowed identification of individuals based on their unique ventral surface markings (spots). The ability to recognise individuals enabled many aspects of both populations to be examined. A total of 449 individual rays were identified of Manta birostris variant A and 101 individual rays were identified of Manta birostris variant B. Both populations exhibited a highly significant female bias in observed sex ratios of 1:3.5 and 1:7.4 respectively. The photographic identification approach was used in studies of population size and structure and to examine the reproductive ecology of these two distinct populations. Population estimates incorporating four years of re-sighting data on Manta birostris variant A and five years of data on Manta birostris variant B were conducted using the program MARK resulting in super population estimates of 890 and 600 individuals respectively. The region encompassing the study site was identified as a mating ground for Manta birostris variant A based on observations of mating events and fresh pectoral fin tip scars on female rays produced when the male rays bite onto the fins during copulation. The distribution of scars was highly biased, with 99% on the left pectoral fin, indicative of a strong lateralised behavioural trait in this species. No other elasmobranch has been reported to display behavioural lateralisation. The study region also acts as a birthing ground, with individuals giving birth in the summer after a gestation period of approximately one year. Reproductive periodicity in M. birostris variant A was most commonly biennial, but a few individuals were seen to be pregnant in consecutive years, confirming an annual ovulatory cycle. The production of a single pup appears to be the normal situation, although observations in the wild as well as during opportunistic dissections of individuals killed in fisheries revealed that two pups are conceived on occasion. Morphometric analysis of late-term foetus (M. birostris variant A) was contrasted with measurements taken from adult rays. These are the only detailed measurements on M. birostris variant A from the western Indian Ocean. Predatory scarring and bite injuries on individuals of both variant A and B were consistent with attacks from sharks of various sizes. The frequency and effect of these predatory injuries on the two populations were examined over a three-year period. Acoustic tags were attached to fourteen rays and their presence/absence at sites around the major inshore reefs was explored to evaluate how the rays utilised their environment on a temporal scale. Cleaning activity of both putative species of manta rays by small fishes occurred on these reefs and was considered to be the main reason why the rays were present at these sites. Parasite removal and wound healing were implicated as the benefits received. Host cleaner fish species partition the manta ray body to avoid interspecific competition while the two putative species appear to partition cleaning habitats, with very little overlap apparent. This research on manta rays is the first of its kind in African waters. Many aspects of the study have contributed to the limited baseline data currently available for this genus. The study additionally provided sufficient empirical evidence to warrant the separation of these two putative species of Manta and a full revision of the genus with the systematic examination of specimens throughout Manta’s range. The results of this study may be useful in this endeavour as diagnostic characteristics have been isolated that may help to differentiate members of this genus. The results of this research study are directly applicable to management strategies for both putative species of manta rays off the East Coast of Africa as well as populations worldwide and have highlighted the potential need for different conservation strategies

Despite being the world’s largest batoid fish, manta rays have been the focus of very little research in the last century. Almost nothing is currently known about their ecology or distribution throughout the world’s oceans. Manta rays are considered to be near-threatened to vulnerable by the World Conservation Union’s Redlist of endangered species, yet many populations still face threats from target fishing for their fin cartilage, branchial filaments and meat. The taxonomic history of the genus Manta has been questionable and convoluted,with Manta having one of the most extensive generic and species synonymies of any living genus of cartilaginous fish. Currently this genus is considered to be monotypicwith a single recognized species, Manta birostris. Behavioural and morphological data collected during this study showed that there are two distinct, sympatric species of manta ray in Mozambique. This discovery has led to an examination of manta ray distribution worldwide, with the two species hypothesis receiving further support from study sites in the Indian and Pacific Oceans. This thesis examined large populations of both putative species in Mozambique. Development and application of a photographic-identification methodology allowed identification of individuals based on their unique ventral surface markings (spots). The ability to recognise individuals enabled many aspects of both populations to be examined. A total of 449 individual rays were identified of Manta birostris variant A and 101 individual rays were identified of Manta birostris variant B. Both populations exhibited a highly significant female bias in observed sex ratios of 1:3.5 and 1:7.4 respectively. The photographic identification approach was used in studies of population size and structure and to examine the reproductive ecology of these two distinct populations. Population estimates incorporating four years of re-sighting data on Manta birostris variant A and five years of data on Manta birostris variant B were conducted using the program MARK resulting in super population estimates of 890 and 600 individuals respectively. The region encompassing the study site was identified as a mating ground for Manta birostris variant A based on observations of mating events and fresh pectoral fin tip scars on female rays produced when the male rays bite onto the fins during copulation. The distribution of scars was highly biased, with 99% on the left pectoral fin, indicative of a strong lateralised behavioural trait in this species. No other elasmobranch has been reported to display behavioural lateralisation. The study region also acts as a birthing ground, with individuals giving birth in the summer after a gestation period of approximately one year. Reproductive periodicity in M. birostris variant A was most commonly biennial, but a few individuals were seen to be pregnant in consecutive years, confirming an annual ovulatory cycle. The production of a single pup appears to be the normal situation, although observations in the wild as well as during opportunistic dissections of individuals killed in fisheries revealed that two pups are conceived on occasion. Morphometric analysis of late-term foetus (M. birostris variant A) was contrasted with measurements taken from adult rays. These are the only detailed measurements on M. birostris variant A from the western Indian Ocean. Predatory scarring and bite injuries on individuals of both variant A and B were consistent with attacks from sharks of various sizes. The frequency and effect of these predatory injuries on the two populations were examined over a three-year period. Acoustic tags were attached to fourteen rays and their presence/absence at sites around the major inshore reefs was explored to evaluate how the rays utilised their environment on a temporal scale. Cleaning activity of both putative species of manta rays by small fishes occurred on these reefs and was considered to be the main reason why the rays were present at these sites. Parasite removal and wound healing were implicated as the benefits received. Host cleaner fish species partition the manta ray body to avoid interspecific competition while the two putative species appear to partition cleaning habitats, with very little overlap apparent. This research on manta rays is the first of its kind in African waters. Many aspects of the study have contributed to the limited baseline data currently available for this genus. The study additionally provided sufficient empirical evidence to warrant the separation of these two putative species of Manta and a full revision of the genus with the systematic examination of specimens throughout Manta’s range. The results of this study may be useful in this endeavour as diagnostic characteristics have been isolated that may help to differentiate members of this genus. The results of this research study are directly applicable to management strategies for both putative species of manta rays off the East Coast of Africa as well as populations worldwide and have highlighted the potential need for different conservation strategies

Despite being the world’s largest batoid fish, manta rays have been the focus of very little research in the last century. Almost nothing is currently known about their ecology or distribution throughout the world’s oceans. Manta rays are considered to be near-threatened to vulnerable by the World Conservation Union’s Redlist of endangered species, yet many populations still face threats from target fishing for their fin cartilage, branchial filaments and meat. The taxonomic history of the genus Manta has been questionable and convoluted,with Manta having one of the most extensive generic and species synonymies of any living genus of cartilaginous fish. Currently this genus is considered to be monotypicwith a single recognized species, Manta birostris. Behavioural and morphological data collected during this study showed that there are two distinct, sympatric species of manta ray in Mozambique. This discovery has led to an examination of manta ray distribution worldwide, with the two species hypothesis receiving further support from study sites in the Indian and Pacific Oceans. This thesis examined large populations of both putative species in Mozambique. Development and application of a photographic-identification methodology allowed identification of individuals based on their unique ventral surface markings (spots). The ability to recognise individuals enabled many aspects of both populations to be examined. A total of 449 individual rays were identified of Manta birostris variant A and 101 individual rays were identified of Manta birostris variant B. Both populations exhibited a highly significant female bias in observed sex ratios of 1:3.5 and 1:7.4 respectively. The photographic identification approach was used in studies of population size and structure and to examine the reproductive ecology of these two distinct populations. Population estimates incorporating four years of re-sighting data on Manta birostris variant A and five years of data on Manta birostris variant B were conducted using the program MARK resulting in super population estimates of 890 and 600 individuals respectively. The region encompassing the study site was identified as a mating ground for Manta birostris variant A based on observations of mating events and fresh pectoral fin tip scars on female rays produced when the male rays bite onto the fins during copulation. The distribution of scars was highly biased, with 99% on the left pectoral fin, indicative of a strong lateralised behavioural trait in this species. No other elasmobranch has been reported to display behavioural lateralisation. The study region also acts as a birthing ground, with individuals giving birth in the summer after a gestation period of approximately one year. Reproductive periodicity in M. birostris variant A was most commonly biennial, but a few individuals were seen to be pregnant in consecutive years, confirming an annual ovulatory cycle. The production of a single pup appears to be the normal situation, although observations in the wild as well as during opportunistic dissections of individuals killed in fisheries revealed that two pups are conceived on occasion. Morphometric analysis of late-term foetus (M. birostris variant A) was contrasted with measurements taken from adult rays. These are the only detailed measurements on M. birostris variant A from the western Indian Ocean. Predatory scarring and bite injuries on individuals of both variant A and B were consistent with attacks from sharks of various sizes. The frequency and effect of these predatory injuries on the two populations were examined over a three-year period. Acoustic tags were attached to fourteen rays and their presence/absence at sites around the major inshore reefs was explored to evaluate how the rays utilised their environment on a temporal scale. Cleaning activity of both putative species of manta rays by small fishes occurred on these reefs and was considered to be the main reason why the rays were present at these sites. Parasite removal and wound healing were implicated as the benefits received. Host cleaner fish species partition the manta ray body to avoid interspecific competition while the two putative species appear to partition cleaning habitats, with very little overlap apparent. This research on manta rays is the first of its kind in African waters. Many aspects of the study have contributed to the limited baseline data currently available for this genus. The study additionally provided sufficient empirical evidence to warrant the separation of these two putative species of Manta and a full revision of the genus with the systematic examination of specimens throughout Manta’s range. The results of this study may be useful in this endeavour as diagnostic characteristics have been isolated that may help to differentiate members of this genus. The results of this research study are directly applicable to management strategies for both putative species of manta rays off the East Coast of Africa as well as populations worldwide and have highlighted the potential need for different conservation strategies

Despite being the world’s largest batoid fish, manta rays have been the focus of very little research in the last century. Almost nothing is currently known about their ecology or distribution throughout the world’s oceans. Manta rays are considered to be near-threatened to vulnerable by the World Conservation Union’s Redlist of endangered species, yet many populations still face threats from target fishing for their fin cartilage, branchial filaments and meat. The taxonomic history of the genus Manta has been questionable and convoluted,with Manta having one of the most extensive generic and species synonymies of any living genus of cartilaginous fish. Currently this genus is considered to be monotypicwith a single recognized species, Manta birostris. Behavioural and morphological data collected during this study showed that there are two distinct, sympatric species of manta ray in Mozambique. This discovery has led to an examination of manta ray distribution worldwide, with the two species hypothesis receiving further support from study sites in the Indian and Pacific Oceans. This thesis examined large populations of both putative species in Mozambique. Development and application of a photographic-identification methodology allowed identification of individuals based on their unique ventral surface markings (spots). The ability to recognise individuals enabled many aspects of both populations to be examined. A total of 449 individual rays were identified of Manta birostris variant A and 101 individual rays were identified of Manta birostris variant B. Both populations exhibited a highly significant female bias in observed sex ratios of 1:3.5 and 1:7.4 respectively. The photographic identification approach was used in studies of population size and structure and to examine the reproductive ecology of these two distinct populations. Population estimates incorporating four years of re-sighting data on Manta birostris variant A and five years of data on Manta birostris variant B were conducted using the program MARK resulting in super population estimates of 890 and 600 individuals respectively. The region encompassing the study site was identified as a mating ground for Manta birostris variant A based on observations of mating events and fresh pectoral fin tip scars on female rays produced when the male rays bite onto the fins during copulation. The distribution of scars was highly biased, with 99% on the left pectoral fin, indicative of a strong lateralised behavioural trait in this species. No other elasmobranch has been reported to display behavioural lateralisation. The study region also acts as a birthing ground, with individuals giving birth in the summer after a gestation period of approximately one year. Reproductive periodicity in M. birostris variant A was most commonly biennial, but a few individuals were seen to be pregnant in consecutive years, confirming an annual ovulatory cycle. The production of a single pup appears to be the normal situation, although observations in the wild as well as during opportunistic dissections of individuals killed in fisheries revealed that two pups are conceived on occasion. Morphometric analysis of late-term foetus (M. birostris variant A) was contrasted with measurements taken from adult rays. These are the only detailed measurements on M. birostris variant A from the western Indian Ocean. Predatory scarring and bite injuries on individuals of both variant A and B were consistent with attacks from sharks of various sizes. The frequency and effect of these predatory injuries on the two populations were examined over a three-year period. Acoustic tags were attached to fourteen rays and their presence/absence at sites around the major inshore reefs was explored to evaluate how the rays utilised their environment on a temporal scale. Cleaning activity of both putative species of manta rays by small fishes occurred on these reefs and was considered to be the main reason why the rays were present at these sites. Parasite removal and wound healing were implicated as the benefits received. Host cleaner fish species partition the manta ray body to avoid interspecific competition while the two putative species appear to partition cleaning habitats, with very little overlap apparent. This research on manta rays is the first of its kind in African waters. Many aspects of the study have contributed to the limited baseline data currently available for this genus. The study additionally provided sufficient empirical evidence to warrant the separation of these two putative species of Manta and a full revision of the genus with the systematic examination of specimens throughout Manta’s range. The results of this study may be useful in this endeavour as diagnostic characteristics have been isolated that may help to differentiate members of this genus. The results of this research study are directly applicable to management strategies for both putative species of manta rays off the East Coast of Africa as well as populations worldwide and have highlighted the potential need for different conservation strategies