Laura Boren - Academia.edu (original) (raw)

Papers by Laura Boren

Research paper thumbnail of Assessing the impact of tourism on New Zealand fur seals (Arctocephalus forsteri)

Marine mammal viewing and encounters are significant tourist activities in some areas of New Zeal... more Marine mammal viewing and encounters are significant tourist activities in some areas of New Zealand - it was estimated in 1992 that at least 300,000+ tourists took part in marine related tourism in New Zealand annually (Robertson, 1992), and the industry has grown considerably since then. While eco-tourism can have positive outcomes (e.g. generating revenue and increasing environmental awareness), if it is not managed effectively, it can also have a negative impact on the target species and their environment. Effective management requires an understanding of how the target species react to tourist activities. We need to know: • If the animals are modifying their behaviour and if so how can we measure the changes in behaviour? • Are the changes in behaviour biologically significant? • How can we prevent or mitigate any negative effects of eco-tourism on marine mammals? New Zealand fur seals, Arctocephalus forsteri, are the only marine mammal found regularly ashore on accessible sect...

Research paper thumbnail of Incorporating non-equilibrium dynamics into demographic history inferences of a migratory marine species

Research paper thumbnail of First Direct Evidence for Natal Wintering Ground Fidelity and Estimate of Juvenile Survival in the New Zealand Southern Right Whale Eubalaena australis

Research paper thumbnail of Mitochondrial DNA structure and colony expansion dynamics of New Zealand fur seals (Arctocephalus forsteri) around Banks Peninsula

New Zealand Journal of Zoology, 2016

ABSTRACT New Zealand fur seals are one of many pinniped species that survived the commercial seal... more ABSTRACT New Zealand fur seals are one of many pinniped species that survived the commercial sealing of the eighteenth and nineteenth centuries in dangerously low numbers. After the enforcement of a series of protection measures in the early twentieth century, New Zealand fur seals began to recover from the brink of extinction. We examined the New Zealand fur seal populations of Banks Peninsula, South Island, New Zealand using the mitochondrial DNA control region. We identified a panmictic population structure around Banks Peninsula. The most abundant haplotype in the area showed a slight significant aggregated structure. The Horseshoe Bay colony showed the least number of shared haplotypes with other colonies, suggesting a different origin of re-colonisation of this specific colony. The effective population size of the New Zealand fur seal population at Banks Peninsula was estimated at approximately 2500 individuals. The exponential population growth rate parameter for the area was 35, which corresponds to an expanding population. In general, samples from adjacent colonies shared 4.4 haplotypes while samples collected from colonies separated by between five and eight bays shared 1.9 haplotypes. The genetic data support the spill-over dynamics of colony expansion already suggested for this species. Approximate Bayesian computations analysis suggests re-colonisation of the area from two main clades identified across New Zealand with a most likely admixture coefficient of 0.41 to form the Banks Peninsula population. Approximate Bayesian computations analysis estimated a founder population size of approximately 372 breeding individuals for the area, which then rapidly increased in size with successive waves of external recruitment. The population of fur seals in the area is probably in the late phase of maturity in the colony expansion dynamic.

Research paper thumbnail of Data from: Incorporating non-equilibrium dynamics into demographic history inferences of a migratory marine species

Understanding how dispersal and gene flow link geographically separated populations over evolutio... more Understanding how dispersal and gene flow link geographically separated populations over evolutionary history is challenging, particularly in migratory marine species. In southern right whales (SRWs, Eubalaena australis), patterns of genetic diversity are likely influenced by the glacial climate cycle and recent history of whaling. Here we use a dataset of mitochondrial DNA (mtDNA) sequences (n=1,327) and nuclear markers (17 microsatellite loci, n=222) from major wintering grounds to investigate circumpolar population structure, historical demography, and effective population size. Analyses of nuclear genetic variation identify two population clusters that correspond to the South Atlantic and Indo-Pacific ocean basins that have similar effective breeder estimates. In contrast, all wintering grounds show significant differentiation for mtDNA, but no sex-biased dispersal was detected using the microsatellite genotypes. An approximate Bayesian computation (ABC) approac h with microsatellite markers compared scenarios with gene flow through time, or isolation and secondary contact between ocean basins, while modeling declines in abundance linked to whaling. Secondary-contact scenarios yield the highest posterior probabilities, implying that populations in different ocean basins were largely isolated and came into secondary contact within the last 25,000 years, but the role of whaling in changes in genetic diversity and gene flow over recent generations could not be resolved. We hypothesise that these findings are driven by factors that promote isolation, such as female philopatry, and factors that could promote dispersal, such oceanographic changes. These findings highlight the application of ABC approaches to infer connectivity in mobile species with complex population histories and currently low levels of differentiation

Research paper thumbnail of offspring_CowCalf_dataset_Jan12

CERVUS input file for paternity assignment analysis. Lists cow-calf pairs and candidate males

Research paper thumbnail of CowCalfDad_genotypesJan2012_Dryad

Research paper thumbnail of Australia_Allelefreq

Research paper thumbnail of All_NZSA_ALF

Research paper thumbnail of Simulation_script

R code used in paper to test the power of our analyses to reject the null hypothesis of panmictic... more R code used in paper to test the power of our analyses to reject the null hypothesis of panmictic mating between New Zealand and southwest Australia right whale subpopulations

Research paper thumbnail of Data from: Paternity assignment and demographic closure in the New Zealand southern right whale

The identification and characterisation of reproductively isolated subpopulations or ‘stocks’ is ... more The identification and characterisation of reproductively isolated subpopulations or ‘stocks’ is essential for effective conservation and management decisions. This can be difficult in vagile marine species like marine mammals. We used paternity assignment and ‘gametic recapture’ to examine the reproductive autonomy of southern right whales (Eubalaena australis) on their New Zealand (NZ) calving grounds. We derived DNA profiles for 34 mother-calf pairs from skin biopsy samples, using sex-specific markers, 13 microsatellite loci and mtDNA haplotypes. We constructed DNA profiles for 314 adult males, representing 30% of the census male abundance of the NZ stock, previously estimated from genotypic mark-recapture modelling to be 1085 (95% CL 855, 1416). Under the hypothesis of demographic closure and the assumption of equal reproductive success among males, we predict: (1) the proportion of paternities assigned will reflect the proportion of the male population sampled and (2) the gametic mark-recapture (GMR) estimate of male abundance will be equivalent to the census male estimate for the NZ stock. Consistent with these predictions, we found that the proportion of assigned paternities equalled the proportion of the census male population size sampled. Using the sample of males as the initial capture, and paternity assignment as the recapture, the GMR estimate of male abundance was 1001 (95% CL 542, 1469), similar to the male census estimate. These findings suggest that right whales returning to the NZ calving ground are reproductively autonomous on a generational timescale, as well as isolated by maternal fidelity on an evolutionary timescale, from others in the Indo-Pacific region

Research paper thumbnail of Genotype data for Carroll et al

Microsatellite genotype data for southern right whales in Carroll et al 201

Research paper thumbnail of Paternity assignment and demographic closure in the New Zealand southern right whale

Research paper thumbnail of The effects of seal-swim activities on the New Zealand fur seal (Arctophoca australis forsteri) in the Bay of Plenty, New Zealand, and recommendations for a sustainable tourism industry

Marine Policy, 2014

ABSTRACT Wildlife tourism (including pinniped tourism) offers people the opportunity to see wildl... more ABSTRACT Wildlife tourism (including pinniped tourism) offers people the opportunity to see wildlife in their natural environment. It can provide positive outcomes for the animals, through improved resources for conservation, or negative outcomes, such as inducing the animals to move away. This study assessed the impacts and sustainability of a novel but growing tourism industry, swimming with seals, based on interactions with New Zealand fur seals (Arctophoca australis forsteri) in the Bay of Plenty, New Zealand, between December 2011 and March 2012. The behaviour of all seals in the water (interaction, neutral, and avoidance) was monitored at 1 min intervals, during 16 seal-swim events. Seals mostly ignored the swimmers (54% of records), some interacted with swimmers (41%); seals rarely avoided the swimmers (5%). Interactions peaked in frequency at 6 min into the swims, then declined. They occurred most frequently during December, corresponding with the pupping period when juvenile seals—the age class most likely to interact—are excluded from breeding areas and so spend much of their time in the water. Compliance of tour operators to regulations was also monitored during seal-swim activities and the industry was found to be highly compliant. The results suggest the activities monitored had minimal impact on seals in the water, and are likely to be sustainable in relation to seal conservation. Tourism can be site and time specific, and it is recommended that approaches such as those trialled here be adopted to monitor other wildlife tourism activities to ensure their sustainability. Further research needs to examine potential impacts of the tours on seals ashore.

Research paper thumbnail of Hair Sampling and Genotyping from Hair Follicles: A Minimally-Invasive Alternative for Genetics Studies in Small, Mobile Pinnipeds and Other Mammals

Marine Mammal Science, 2007

Research paper thumbnail of Rearing two New Zealand fur seal ( Arctocephalus forsteri ) pups to weaning

Australian Journal of Zoology, 2008

The rearing of two pups to weaning is a rare occurrence in pinnipeds and in many cases it remains... more The rearing of two pups to weaning is a rare occurrence in pinnipeds and in many cases it remains unknown whether it is a result of twinning or fostering. This study followed two cases where female New Zealand fur seals (Arctocephalus forsteri) raised two pups, at a colony in Kaikoura, New Zealand. Maternal attendance behaviour was observed, mass and condition of the pups were measured and genetic samples were collected to assess relatedness. In one case, the female gave birth to twins, while the other case was found to be fostering of a second pup. In both cases, the filial pups of each female exhibited significantly lower mass, condition and growth rates than the colony average. The twins’ mother reared both pups to weaning and did not appear to spend a different amount of time ashore compared with single-pup mothers. The current study confirms both fostering and twinning in New Zealand fur seals, with the potential for successfully raising the pups to weaning, despite their well ...

Research paper thumbnail of A risk assessment of threats to Maui’s dolphins

Ministry for Primary Industries and Department of Conservation, Wellington, 2012

Summary Maui's dolphin (Cephalorhynchus hectori maui) is a subspecies of Hector's and M... more Summary Maui's dolphin (Cephalorhynchus hectori maui) is a subspecies of Hector's and Maui's dolphin (Cephalorhynchus hectori) and is endemic to New Zealand. Maui's dolphin is listed as 'critically endangered'on the IUCN's Red List of threatened species and 'nationally critical'in the New Zealand Threat Classification System. Current population estimates indicate that about 55 Maui's dolphins over 1 year of age remain, and the population is exposed to a range of humanand non-human-induced threats. A risk assessment ...

Research paper thumbnail of New Zealand fur seals in the Kaikoura region: colony dynamics, maternal investment and health

Colony dynamics, maternal investment, and indicators of health were investigated for the New Zeal... more Colony dynamics, maternal investment, and indicators of health were investigated for the New Zealand fur seal (Arctocephalus forsteri) over four austral summers, 2001- 2005. Effort was focused at the Ohau Point seal colony, north of Kaikoura. Two colonies at Banks Peninsula were included for comparisons of colony growth and pup condition. A range of other colonies were also included for making comparisons about colony dynamics and health indices. Colony dynamics were investigated through mark-recapture estimates of pup production and daily census of all individuals at the Ohau Point colony. Maternal attendance patterns were observed through behavioural observations of known females (n = 120), the use of VHF radio transmitters (n = 33), and female mass and body condition estimates (n = 51). Maternal investment was also investigated through longitudinal sampling of pup mass and growth rates. Parameters used to indicate colony health were: body condition, growth, presence of parasites,...

Research paper thumbnail of The role and presence of a guide: preliminary findings from swim with seal programs and land-based seal viewing in New Zealand

Tourism in Marine …, 2008

Research paper thumbnail of New Zealand fur seals in the Kaikoura region: colony dynamics, maternal investment and health

Colony dynamics, maternal investment, and indicators of health were investigated for the New Zeal... more Colony dynamics, maternal investment, and indicators of health were investigated for the New Zealand fur seal (Arctocephalus forsteri) over four austral summers, 20012005. Effort was focused at the Ohau Point seal colony, north of Kaikoura. Two colonies at Banks Peninsula were included for comparisons of colony growth and pup condition. A range of other colonies were also included for making comparisons about colony dynamics and health indices. Colony dynamics were investigated through mark-recapture estimates of pup production and daily census of all individuals at the Ohau Point colony. Maternal attendance patterns were observed through behavioural observations of known females (n = 120), the use of VHF radio transmitters (n = 33), and female mass and body condition estimates (n = 51). Maternal investment was also investigated through longitudinal sampling of pup mass and growth rates. Parameters used to indicate colony health were: body condition, growth, presence of parasites, a...

Research paper thumbnail of Assessing the impact of tourism on New Zealand fur seals (Arctocephalus forsteri)

Marine mammal viewing and encounters are significant tourist activities in some areas of New Zeal... more Marine mammal viewing and encounters are significant tourist activities in some areas of New Zealand - it was estimated in 1992 that at least 300,000+ tourists took part in marine related tourism in New Zealand annually (Robertson, 1992), and the industry has grown considerably since then. While eco-tourism can have positive outcomes (e.g. generating revenue and increasing environmental awareness), if it is not managed effectively, it can also have a negative impact on the target species and their environment. Effective management requires an understanding of how the target species react to tourist activities. We need to know: • If the animals are modifying their behaviour and if so how can we measure the changes in behaviour? • Are the changes in behaviour biologically significant? • How can we prevent or mitigate any negative effects of eco-tourism on marine mammals? New Zealand fur seals, Arctocephalus forsteri, are the only marine mammal found regularly ashore on accessible sect...

Research paper thumbnail of Incorporating non-equilibrium dynamics into demographic history inferences of a migratory marine species

Research paper thumbnail of First Direct Evidence for Natal Wintering Ground Fidelity and Estimate of Juvenile Survival in the New Zealand Southern Right Whale Eubalaena australis

Research paper thumbnail of Mitochondrial DNA structure and colony expansion dynamics of New Zealand fur seals (Arctocephalus forsteri) around Banks Peninsula

New Zealand Journal of Zoology, 2016

ABSTRACT New Zealand fur seals are one of many pinniped species that survived the commercial seal... more ABSTRACT New Zealand fur seals are one of many pinniped species that survived the commercial sealing of the eighteenth and nineteenth centuries in dangerously low numbers. After the enforcement of a series of protection measures in the early twentieth century, New Zealand fur seals began to recover from the brink of extinction. We examined the New Zealand fur seal populations of Banks Peninsula, South Island, New Zealand using the mitochondrial DNA control region. We identified a panmictic population structure around Banks Peninsula. The most abundant haplotype in the area showed a slight significant aggregated structure. The Horseshoe Bay colony showed the least number of shared haplotypes with other colonies, suggesting a different origin of re-colonisation of this specific colony. The effective population size of the New Zealand fur seal population at Banks Peninsula was estimated at approximately 2500 individuals. The exponential population growth rate parameter for the area was 35, which corresponds to an expanding population. In general, samples from adjacent colonies shared 4.4 haplotypes while samples collected from colonies separated by between five and eight bays shared 1.9 haplotypes. The genetic data support the spill-over dynamics of colony expansion already suggested for this species. Approximate Bayesian computations analysis suggests re-colonisation of the area from two main clades identified across New Zealand with a most likely admixture coefficient of 0.41 to form the Banks Peninsula population. Approximate Bayesian computations analysis estimated a founder population size of approximately 372 breeding individuals for the area, which then rapidly increased in size with successive waves of external recruitment. The population of fur seals in the area is probably in the late phase of maturity in the colony expansion dynamic.

Research paper thumbnail of Data from: Incorporating non-equilibrium dynamics into demographic history inferences of a migratory marine species

Understanding how dispersal and gene flow link geographically separated populations over evolutio... more Understanding how dispersal and gene flow link geographically separated populations over evolutionary history is challenging, particularly in migratory marine species. In southern right whales (SRWs, Eubalaena australis), patterns of genetic diversity are likely influenced by the glacial climate cycle and recent history of whaling. Here we use a dataset of mitochondrial DNA (mtDNA) sequences (n=1,327) and nuclear markers (17 microsatellite loci, n=222) from major wintering grounds to investigate circumpolar population structure, historical demography, and effective population size. Analyses of nuclear genetic variation identify two population clusters that correspond to the South Atlantic and Indo-Pacific ocean basins that have similar effective breeder estimates. In contrast, all wintering grounds show significant differentiation for mtDNA, but no sex-biased dispersal was detected using the microsatellite genotypes. An approximate Bayesian computation (ABC) approac h with microsatellite markers compared scenarios with gene flow through time, or isolation and secondary contact between ocean basins, while modeling declines in abundance linked to whaling. Secondary-contact scenarios yield the highest posterior probabilities, implying that populations in different ocean basins were largely isolated and came into secondary contact within the last 25,000 years, but the role of whaling in changes in genetic diversity and gene flow over recent generations could not be resolved. We hypothesise that these findings are driven by factors that promote isolation, such as female philopatry, and factors that could promote dispersal, such oceanographic changes. These findings highlight the application of ABC approaches to infer connectivity in mobile species with complex population histories and currently low levels of differentiation

Research paper thumbnail of offspring_CowCalf_dataset_Jan12

CERVUS input file for paternity assignment analysis. Lists cow-calf pairs and candidate males

Research paper thumbnail of CowCalfDad_genotypesJan2012_Dryad

Research paper thumbnail of Australia_Allelefreq

Research paper thumbnail of All_NZSA_ALF

Research paper thumbnail of Simulation_script

R code used in paper to test the power of our analyses to reject the null hypothesis of panmictic... more R code used in paper to test the power of our analyses to reject the null hypothesis of panmictic mating between New Zealand and southwest Australia right whale subpopulations

Research paper thumbnail of Data from: Paternity assignment and demographic closure in the New Zealand southern right whale

The identification and characterisation of reproductively isolated subpopulations or ‘stocks’ is ... more The identification and characterisation of reproductively isolated subpopulations or ‘stocks’ is essential for effective conservation and management decisions. This can be difficult in vagile marine species like marine mammals. We used paternity assignment and ‘gametic recapture’ to examine the reproductive autonomy of southern right whales (Eubalaena australis) on their New Zealand (NZ) calving grounds. We derived DNA profiles for 34 mother-calf pairs from skin biopsy samples, using sex-specific markers, 13 microsatellite loci and mtDNA haplotypes. We constructed DNA profiles for 314 adult males, representing 30% of the census male abundance of the NZ stock, previously estimated from genotypic mark-recapture modelling to be 1085 (95% CL 855, 1416). Under the hypothesis of demographic closure and the assumption of equal reproductive success among males, we predict: (1) the proportion of paternities assigned will reflect the proportion of the male population sampled and (2) the gametic mark-recapture (GMR) estimate of male abundance will be equivalent to the census male estimate for the NZ stock. Consistent with these predictions, we found that the proportion of assigned paternities equalled the proportion of the census male population size sampled. Using the sample of males as the initial capture, and paternity assignment as the recapture, the GMR estimate of male abundance was 1001 (95% CL 542, 1469), similar to the male census estimate. These findings suggest that right whales returning to the NZ calving ground are reproductively autonomous on a generational timescale, as well as isolated by maternal fidelity on an evolutionary timescale, from others in the Indo-Pacific region

Research paper thumbnail of Genotype data for Carroll et al

Microsatellite genotype data for southern right whales in Carroll et al 201

Research paper thumbnail of Paternity assignment and demographic closure in the New Zealand southern right whale

Research paper thumbnail of The effects of seal-swim activities on the New Zealand fur seal (Arctophoca australis forsteri) in the Bay of Plenty, New Zealand, and recommendations for a sustainable tourism industry

Marine Policy, 2014

ABSTRACT Wildlife tourism (including pinniped tourism) offers people the opportunity to see wildl... more ABSTRACT Wildlife tourism (including pinniped tourism) offers people the opportunity to see wildlife in their natural environment. It can provide positive outcomes for the animals, through improved resources for conservation, or negative outcomes, such as inducing the animals to move away. This study assessed the impacts and sustainability of a novel but growing tourism industry, swimming with seals, based on interactions with New Zealand fur seals (Arctophoca australis forsteri) in the Bay of Plenty, New Zealand, between December 2011 and March 2012. The behaviour of all seals in the water (interaction, neutral, and avoidance) was monitored at 1 min intervals, during 16 seal-swim events. Seals mostly ignored the swimmers (54% of records), some interacted with swimmers (41%); seals rarely avoided the swimmers (5%). Interactions peaked in frequency at 6 min into the swims, then declined. They occurred most frequently during December, corresponding with the pupping period when juvenile seals—the age class most likely to interact—are excluded from breeding areas and so spend much of their time in the water. Compliance of tour operators to regulations was also monitored during seal-swim activities and the industry was found to be highly compliant. The results suggest the activities monitored had minimal impact on seals in the water, and are likely to be sustainable in relation to seal conservation. Tourism can be site and time specific, and it is recommended that approaches such as those trialled here be adopted to monitor other wildlife tourism activities to ensure their sustainability. Further research needs to examine potential impacts of the tours on seals ashore.

Research paper thumbnail of Hair Sampling and Genotyping from Hair Follicles: A Minimally-Invasive Alternative for Genetics Studies in Small, Mobile Pinnipeds and Other Mammals

Marine Mammal Science, 2007

Research paper thumbnail of Rearing two New Zealand fur seal ( Arctocephalus forsteri ) pups to weaning

Australian Journal of Zoology, 2008

The rearing of two pups to weaning is a rare occurrence in pinnipeds and in many cases it remains... more The rearing of two pups to weaning is a rare occurrence in pinnipeds and in many cases it remains unknown whether it is a result of twinning or fostering. This study followed two cases where female New Zealand fur seals (Arctocephalus forsteri) raised two pups, at a colony in Kaikoura, New Zealand. Maternal attendance behaviour was observed, mass and condition of the pups were measured and genetic samples were collected to assess relatedness. In one case, the female gave birth to twins, while the other case was found to be fostering of a second pup. In both cases, the filial pups of each female exhibited significantly lower mass, condition and growth rates than the colony average. The twins’ mother reared both pups to weaning and did not appear to spend a different amount of time ashore compared with single-pup mothers. The current study confirms both fostering and twinning in New Zealand fur seals, with the potential for successfully raising the pups to weaning, despite their well ...

Research paper thumbnail of A risk assessment of threats to Maui’s dolphins

Ministry for Primary Industries and Department of Conservation, Wellington, 2012

Summary Maui's dolphin (Cephalorhynchus hectori maui) is a subspecies of Hector's and M... more Summary Maui's dolphin (Cephalorhynchus hectori maui) is a subspecies of Hector's and Maui's dolphin (Cephalorhynchus hectori) and is endemic to New Zealand. Maui's dolphin is listed as 'critically endangered'on the IUCN's Red List of threatened species and 'nationally critical'in the New Zealand Threat Classification System. Current population estimates indicate that about 55 Maui's dolphins over 1 year of age remain, and the population is exposed to a range of humanand non-human-induced threats. A risk assessment ...

Research paper thumbnail of New Zealand fur seals in the Kaikoura region: colony dynamics, maternal investment and health

Colony dynamics, maternal investment, and indicators of health were investigated for the New Zeal... more Colony dynamics, maternal investment, and indicators of health were investigated for the New Zealand fur seal (Arctocephalus forsteri) over four austral summers, 2001- 2005. Effort was focused at the Ohau Point seal colony, north of Kaikoura. Two colonies at Banks Peninsula were included for comparisons of colony growth and pup condition. A range of other colonies were also included for making comparisons about colony dynamics and health indices. Colony dynamics were investigated through mark-recapture estimates of pup production and daily census of all individuals at the Ohau Point colony. Maternal attendance patterns were observed through behavioural observations of known females (n = 120), the use of VHF radio transmitters (n = 33), and female mass and body condition estimates (n = 51). Maternal investment was also investigated through longitudinal sampling of pup mass and growth rates. Parameters used to indicate colony health were: body condition, growth, presence of parasites,...

Research paper thumbnail of The role and presence of a guide: preliminary findings from swim with seal programs and land-based seal viewing in New Zealand

Tourism in Marine …, 2008

Research paper thumbnail of New Zealand fur seals in the Kaikoura region: colony dynamics, maternal investment and health

Colony dynamics, maternal investment, and indicators of health were investigated for the New Zeal... more Colony dynamics, maternal investment, and indicators of health were investigated for the New Zealand fur seal (Arctocephalus forsteri) over four austral summers, 20012005. Effort was focused at the Ohau Point seal colony, north of Kaikoura. Two colonies at Banks Peninsula were included for comparisons of colony growth and pup condition. A range of other colonies were also included for making comparisons about colony dynamics and health indices. Colony dynamics were investigated through mark-recapture estimates of pup production and daily census of all individuals at the Ohau Point colony. Maternal attendance patterns were observed through behavioural observations of known females (n = 120), the use of VHF radio transmitters (n = 33), and female mass and body condition estimates (n = 51). Maternal investment was also investigated through longitudinal sampling of pup mass and growth rates. Parameters used to indicate colony health were: body condition, growth, presence of parasites, a...