Interactions between humans and dusky dolphins in the coastal waters of New Zealand are increasing. My research focused on tourism interactions, with Kaikoura as the study site; and, on habitat use in an active aquaculture area, with Admiralty Bay as the study site. In Kaikoura, companies engaged in commercial cetacean tourism (For Hire Company) have permits issued by the New Zealand’s Department of Conservation, allowing them to take paying customers out to view and swim with wild dusky dolphins. During summer and fall of 2005, I assessed the effectiveness of a voluntary ‘rest period’ established to give time free of humans to the dolphins. I used a theodolite to track the movements of large groups of dusky dolphins and recorded the arrival, departure and behaviors of all vessels approaching within 400 m of the group. The ‘rest period’ resulted in a reduction of vessel visits compared to non-rest periods, yet one For Hire Company and private recreational vessels continued to visit dusky dolphin groups during this time. To increase compliance with the voluntary regulation, more education is needed targeting private recreational vessels. Weekend traffic was higher compared to weekday traffic, during both rest and non-rest periods; a large increase occurred in weekend non-commercial vessel traffic. Swimming with calves is prohibited by New Zealand’s Marine Mammal Protection Regulations of 1992, yet 71.4 percent of the swim attempts I observed on-board For Hire Company tours were conducted with groups containing calves. More should be done to reduce the number of swims conducted with groups of dusky dolphins containing calves. In winter of 2005, I used hourly theodolite scans to record the number of dusky dolphin groups using Admiralty Bay, a different near-shore environment with less tourism than off Kaikoura, and with near-shore mussel farms. Groups of dusky dolphins were observed in Admiralty Bay using the full extent of the bay. This re-enforces previous findings that Admiralty Bay is an important winter foraging ground for dusky dolphins, and further aquaculture development in the bay would remove available foraging habitat.

The distribution, behaviours, and composition of dusky dolphin (Lagenorhynchus obscurus) nursery groups off Kaikoura, New Zealand, were examined. Data were collected during January–May 2005 and December 2005–April 2006 by systematic boat based surveys, group focal follows and photo-identification techniques. A total of 99 nursery groups were encountered on survey. Nursery groups were encountered in shallow water (<20 m) significantly more often than in deeper water (>20 m). Other group types (large groups, mating groups, adult non-mating groups) were not found in shallow water significantly more often than in deeper water. By staying in the shallower water, nursery groups may be protected from aggressive conspecifics and predators. More boats, especially private recreational boats, were found in the shallower waters, indicating that nursery groups are at greater risk from encounters with boat motors or recreational fishing gear in such areas. Group focal follows of at least 30- minutes were conducted on 56 nursery groups. Calves engaged in significantly more high energy behaviours (displays and head first re-entries) than non-calves in nursery groups. These groups were predominantly resting and compactly organized (interanimal distance less than 1 adult body length). Nursery groups showed a high level of synchrony, with 44% of groups synchronizing their submergence and surfacings for most of the focal follow. Median group size was 14, with a minimum of 2 mother-calf pairs to a maximum of 50 mother-calf pairs. A total of 260 individuals were catalogued as members of nursery groups during the two field seasons. Of these, 112 individuals were seen in nursery groups on at least two different days. Some individuals photographed with young calves in nursery groups off Kaikoura were later photographed in Admiralty Bay, 275 km northwest of Kaikoura. Other individuals photographed together in nursery groups in 2005 were also together in nursery groups in 2006.

The sex of and genetic relatedness among interacting individuals are known to be biologically fundamental features that characterize the composition of animal groups. Current work continues to illuminate reasons for the variety of animal social patterns, including patterns in group membership. I investigated the composition of dusky dolphin groups relative to sex and relatedness at two locations in New Zealand. In Kaikoura, dusky dolphins are found year-round, foraging nocturnally on verticallymigrating prey and socializing in distinct group types (mating, nursery, and adult) during the day. By contrast, dusky dolphins use Admiralty Bay, where they feed diurnally on small schooling fishes, primarily in the winter. Molecular sexing revealed the sex of 107 dusky dolphins. The Kaikoura data support previous findings that small mating groups consist mostly of males and indicate that small adult groups can consist of either or both sexes. In Admiralty Bay, the percentage of female dolphins present during the study was estimated to be only 7.4%−22.2% (95% confidence interval, n=88). A randomization test further indicates that dusky dolphins in Admiralty Bay grouped preferentially with same-sex individuals. Nuclear and mitochondrial markers were used to investigate patterns of relatedness. Dusky dolphins sampled in Kaikoura (n=17) and Admiralty Bay (n=47) were genotyped at seven microsatellite loci, and genetic relatedness among all genotyped pairs was estimated. A randomization test indicates that dusky dolphins did not group preferentially by relatedness in Admiralty Bay. Grouping history for 13 genotyped samples was also known from a multi-year photographic record of individually distinctive dusky dolphins. No relationship was found between these longer-term grouping patterns and genetic relatedness. The d-loop region of the mitochondrial DNA (mtDNA) was sequenced for 197 dusky dolphins. The pattern of grouping among dolphins with different haplotypes indicates that dusky dolphin groups are not strongly structured by maternal lineages. However, data from eight individual dusky dolphins hint that nursery groups in Kaikoura tend to consist of dolphins that share a maternal ancestor. This investigation raises many questions about the nature of dusky dolphin social organization and suggests promising avenues for finer-grained investigations into the causes and consequences of dusky dolphin group structure.

I examined fission-fusion sociality in dusky dolphins (Lagenorhynchus obscurus), and investigated aspects of social convergence between dolphins and great apes. I used boat-based group focal follows and photo-identification to collect data in Admiralty Bay, New Zealand during 2005-2006. I used generalized estimating equations to examine relationships between party (group) size, rate of party fission-fusion, activity, and location; and relationships between leaping frequency and behavior. Using photo-identification images from 2001-2006, I analyzed the strength and temporal patterning of associations, short- and long-term association patterns, preferred/avoided associations, and behaviorally-specific preferred associations. To analyze social convergence between dolphins and great apes, I compared female bottlenose dolphin (Tursiops spp.) and chimpanzee (Pan troglodytes) social strategies through literature review. I conducted 171 group focal follows, totaling 157 observation hours. Mean party size was 7.0±6.0 individuals. Party size changed every 5±.47.6 min on average. The most frequent activity was resting (37%), followed by traveling (29%), foraging (18%), and socializing (15%). Foraging was positively related to party size and rate of fission-fusion. Near mussel farms, foraging increased, traveling decreased, and rate of party fusion increased. "Clean" leaps were the most frequent leap type (84%) and were positively related to party size and foraging. Noisy and coordinated leaps were positively related to party size; noisy leaps were negatively related to foraging. Associations during 2001-2006 (N = 228 individuals) were nonrandom for 125 days; associations within one field season were nonrandom for 60 days. Individuals formed preferred/avoided associations during most years. The strongest associations occurred during foraging and socializing; the weakest associations occurred during traveling. Individuals formed preferred associations during foraging, resting, and socializing. Review of female bottlenose dolphin and chimpanzee sociality revealed that: 1) females form weaker bonds and are less social than males, 2) females associate mostly with other females, 3) mothers are often alone with their offspring, 4) mothers (vs. non-mothers) and non-cycling (vs. cycling) females associate less with males, and 5) non-cycling (vs. cycling) females occur in smaller parties. Female dolphins may be more social than female chimpanzees due to decreased scramble competition, increased predation risk, and decreased cost of transport for dolphins vs. chimpanzees.

This study presents evidence from mitochondrial and nuclear loci that there is genetic divergence among and within geographic populations of Lagenorhynchus obscurus. The effect of seasonal variation on the genetic structure within New Zealand was examined with mitochondrial DNA control region sequences from 4 localities. Analysis of nested haplotype clades indicated genetic fragmentation and at least 1 historical population expansion within New Zealand. AMOVA and Fst values from nuclear and mitochondrial DNA sequences suggested significant divergence between New Zealand, South Africa, Argentina, and Peru. Dispersal via the west-wind drift was not supported by patterns of population structure among regions. Alternatively, these data support reciprocal exchange among all four regions with 100% posterior probability for a root of origin in the Indian/Atlantic Oceans. The degree of divergence between Peru and other regions indicates the isolation of Peruvian stock is temporally correlated with the constriction of Drake’s passage in the Plio-Pleistocene. There is evidence that the Plio-Pliestocene paleoceanography of the Indian and Southern Atlantic Oceans influenced phylogeography with shifts of temperate sea surface temperatures northward ~5º of latitude, disrupting the dispersal corridor between New Zealand and Atlantic populations. A preference for temperate waters along continental shelves is proposed as an explanation for lack of contemporary genetic exchange among regions. This study supports the polyphyly of the genus Lagenorhynchus. North Atlantic species form a monophyletic Lagenorhynchus. In the Southern Hemisphere, L. australis/L. cruciger and L. obliquidens/L. obscurus do not form a monophyletic group. I discuss the taxonomic implications and propose taxonomic revision of the genus based on these results. Measures of character interaction indicate that combined evidence from nuclear and mitochondrial genes provide better phylogenetic resolution among delphinid lineages than any data partition independently, despite some indications of conflict among mitochondrial and nuclear data.

Foraging theory suggests that hungry animals balance a complex set of costs and benefits when determining what and how to eat. Prey distribution, patch size, and the presence of conspecifics are important factors influencing a predator's feeding tactics, including the decision to feed individually or socially. Dusky dolphins (Lagenorhynchus obscurus) in New Zealand employ different feeding tactics in varying habitats and seasons. I used programmed survey routes and opportunistic sightings to examine the habitat use and feeding mechanics of dusky dolphins in Admiralty Bay, New Zealand, a protected shallow-water environment frequented by wintering dolphins. I encountered 253 dolphin groups, of which 58.5% were engaged in food-acquisition activities. Photographic efforts revealed a total of 177 individually-recognizable dolphins, 100 of which were returnees from previous seasons. Thirty-seven feeding groups and 70 bouts of feeding behavior were followed. Two-minute interval sampling as well as active acoustic sonar were used to test the hypothesis that diurnally-feeding dolphins would work in a coordinated manner to bring schooling fish to the surface. Feeding tactics observed in Admiralty Bay were then compared to foraging by some of the same animals in the unprotected, deep-water environment off Kaikoura, where large numbers of dusky dolphins feed during the night on organisms associated with a vertically-migrating scattering layer. Evidence supporting coordinated surface feeding was not statistically significant, but indicative of behavioral flexibility in feeding styles as part of a larger feeding repertoire. A potential shift in prey distribution from previous years may also explain some observed patterns. Feeding groups were positively correlated with seabirds and New Zealand fur seals (Arctocephalus forsteri). Mean group size of 6.1 (± 8.23 S.D., n=253) in Admiralty Bay is dramatically less than groups observed off Kaikoura, a variation likely reflecting differences in prey number and distribution, as well as differences in predation risk by deep-water sharks and killer whales. Behavioral flexibility likely confers an adaptive advantage for species subject to environmental fluctuation, whether due to natural or anthropogenic sources. Further research is necessary to evaluate prey distribution in Admiralty Bay and its possible effects on feeding dusky dolphins.

This thesis examines the characteristics of dusky dolphin (Lagenorhynchus obscurus) nursery groups and ontogeny of dusky dolphin calves. Data were collected via boat-based group focal follows of nurseries from October 2006-May 2007. A total of 87 nursery groups were encountered. Data were analyzed according to age category (infant or yearling) and season (early or late). Nursery group membership was lowest in the early season and when yearlings were present. The average number of yearlings in a nursery group was less than that of infants. The predominant activity of calves was rest. Early infants rested the most, while travel seemed most important for late infants, and early yearlings were most likely to forage. With the exception of early infants, all calves were more likely than adults to interact with boats. When taking month into account, yearlings were more social in general than infants. Infants showed a positive trend in sociality, while yearling sociality remained relatively stable. Nursery groups are markedly segregated by calf age, and 80% of nursery groups contained calves of only one age group. Dusky dolphin calves show a similar trend in preference for position in relation to the mother as that in bottlenose dolphins (Tursiops sp.), with echelon swim decreasing with age. However, all calves appear to prefer echelon swim when nursery groups are traveling. Calves were more likely to swim independently in the late part of the season and while foraging or socializing, and were more likely to be in close proximity to their mothers while resting or traveling. Calves learned noisy leaps, followed by clean, coordinated, and acrobatic leaps, in that order. There was no clear relationship between behavioral state and types of leaps performed by calves. Early infants leapt less often than older calves, but leap frequency did not differ among the older calves. The overall pattern in the ontogeny of dusky dolphin leaps indicates that the physical development of leaps is learned individually, while the context in which the leaps are performed is learned from conspecifics. These results indicate that nursery groups represent an important environment for healthy physical and social development of calves.

This thesis examines the characteristics of dusky dolphin (Lagenorhynchus obscurus) nursery groups and ontogeny of dusky dolphin calves. Data were collected via boat-based group focal follows of nurseries from October 2006-May 2007. A total of 87 nursery groups were encountered. Data were analyzed according to age category (infant or yearling) and season (early or late). Nursery group membership was lowest in the early season and when yearlings were present. The average number of yearlings in a nursery group was less than that of infants. The predominant activity of calves was rest. Early infants rested the most, while travel seemed most important for late infants, and early yearlings were most likely to forage. With the exception of early infants, all calves were more likely than adults to interact with boats. When taking month into account, yearlings were more social in general than infants. Infants showed a positive trend in sociality, while yearling sociality remained relatively stable. Nursery groups are markedly segregated by calf age, and 80% of nursery groups contained calves of only one age group. Dusky dolphin calves show a similar trend in preference for position in relation to the mother as that in bottlenose dolphins (Tursiops sp.), with echelon swim decreasing with age. However, all calves appear to prefer echelon swim when nursery groups are traveling. Calves were more likely to swim independently in the late part of the season and while foraging or socializing, and were more likely to be in close proximity to their mothers while resting or traveling. Calves learned noisy leaps, followed by clean, coordinated, and acrobatic leaps, in that order. There was no clear relationship between behavioral state and types of leaps performed by calves. Early infants leapt less often than older calves, but leap frequency did not differ among the older calves. The overall pattern in the ontogeny of dusky dolphin leaps indicates that the physical development of leaps is learned individually, while the context in which the leaps are performed is learned from conspecifics. These results indicate that nursery groups represent an important environment for healthy physical and social development of calves.

I characterized dusky dolphin (Lagenorhynchus obscurus) underwater bait-balling behaviors and acoustic signals, and compared data between Argentina and New Zealand (NZ) to investigate the roles of ecology versus social learning. I quantified prey herding and capturing behaviors from video footage, and I analyzed acoustic signals from narrowband recordings. In both locations, I related bait-balling behaviors and acoustic signals to group and prey ball sizes. In NZ, I also related dolphin behaviors to prey ball escape behaviors and acoustic signal parameters to examine proximate functions. Observed herding behaviors typically involved dolphins swimming around or under a prey ball using a side body orientation, while dolphins typically captured fish from the side of a prey ball using a ventral orientation. Coordinated prey-capture behaviors may have made it easier for dolphins to capture fish by trapping fish between dolphins. Signals were categorized as click trains, burst pulses, and combinations due to a bimodal inter-click interval distribution. I observed 3 whistle-like chirp-screams, but no whistles. Sequences of burst pulses also occurred that contained 2-14 burst pulses that aurally and visually appeared closely matched. Similarities between locations suggest that ecological context related to broad behavioral and acoustic parameters, while social learning differences may occur on a finer scale. In NZ, prey balls exhibited horizontal and vertical movements, but the only behavior that preceded escape was “funneling”, the brief formation of a ball shape where the height was at least twice the width. Dolphin behaviors that related to prey balls ascending were type of herding pass, location of prey-capture attempts, and body orientation during attempts. These behavioral parameters may also be used to counter vertical prey escape behaviors. In NZ, all signal categories had a direct or indirect role in capturing prey. Click train-burst pulses were likely used for echolocating on prey, burst pulses and sequences appeared to have communication roles, and the role of click trains was ambiguous. No signal categories appeared to have a herding function, but the sheer number of signals emitted may have caused fish to cluster together more tightly and therefore facilitated capture.