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    The influence of multi-species feeding associations on the foraging behaviour of Australasian gannets (Morus serrator) in the Hauraki Gulf, New Zealand : a thesis presented in partial fulfilment of the requirements for the degree of Master of Science in Conservation Biology at Massey University, Auckland, New Zealand
    (Massey University, 2015) Purvin, Cameron Mackenzie
    In order to successfully capture prey in a challenging and physically demanding environment, Australasian gannets (Morus serrator) have developed a number of foraging strategies. Foremost among such strategies is the joining in formation of mixed species feeding associations (MSFA’s) and the use of specialised plunge diving behaviours. This study sought to determine how gannets vary individual and group plunge diving behaviours in order to maximise the benefits and mitigate the risks associated with feeding in densely packed, highly active MSFA’s. Specifically this study’s aims are to examine how variations in gannet dive heights, angles and completion rates are influenced by external stimuli, including weather parameters, oceanographic conditions and MSFA’s species composition and behaviour. This study additionally seeks to examine what mixed and/or con-­‐specific cues are associated with the formation of synchronous diving bouts, and determine its role as a form of local enhancement for gannets foraging in MSFA’s. Between March 2013 and June 2014, 45 independent boat based surveys were conducted aboard Dolphin Explorer, a 20m tour boat based in New Zealand’s Hauraki Gulf. Using a Canon XH A1S high definition video camera, approximately 11.9hrs of MSFA video footage was collected, containing 5565 recorded gannet plunge dives. Variations in these dives, including their height, angle of entry, and synchronous timing were examined in context of environmental variables including wind speeds, sea surface temperatures, and Beaufort sea states as well as against MSFA variables including the abundance of other foraging gannets, the abundance and behaviour of common dolphins (Delphinus sp.), and the presence of other species including shearwaters (Puffinus spp.), petrels (Fregetta spp.), terns (Sterna spp.), and Bryde’s whales (Balaenoptera edeni). While weather influences were found to have minimal effects on gannet dive behaviours, higher winds were associated with lower altitude dives, and less frequent synchronous diving bouts. This is likely due to the decreased visibility of deeper prey caused by increased surface disturbances as a result of higher wind speeds, coupled with the inherent difficulties of performing highly coordinated aerial manoeuvres in high wind conditions. Increased gannet flock sizes were found to correlate with more frequent high-­‐ altitude 90° dives, and increased rates of dive synchrony. This may be a result of the need to better coordinate dives in denser assemblages of con-­‐specifics to avoid collisions, coupled with the increased foraging success associated with larger foraging groups, and local enhancement. Increased foraging dolphin pods were also associated with more frequent high-­‐ altitude vertical dives. This may result from gannets capitalising on the increased bait-­‐ball density and stability associated with larger foraging pods by taking deeper dives in order to make multiple prey captures with greater ease. Gannet synchrony, however, was only found to decrease in the smallest dolphin pods, again attributed to the decreased foraging abilities of smaller pods. Dolphin foraging behaviours most associated with the maintenance of dense stationary bait-­‐balls were most heavily associated with high, vertical gannet dives, and increased gannet synchronous dives. This is likely the result of gannets capitalising on denser prey assemblages by taking deeper dives in pursuit of multiple prey capture. The presence of Bryde’s whales at MSFA’s was found to decrease the frequency of both high vertical dives and synchronous dives, as lunge feeding on bait-­‐balls scatters fish and decreases prey density. The presence of hetero-­‐specific avian species had various effects on gannet dive behaviours. Most notably, the presence of shearwaters and petrels was found to be associated with greater frequencies of low altitude gannet dives, thought to be a result of an increased collision risk in their presence. Terns meanwhile were associated with decreased dive synchrony, possibly as a result of false hetero-­‐ specific signalling caused by their white plumage.
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    Hunting between the air and the water : the Australasian gannet (Morus serrator) : a thesis presented in partial fulfilment of the requirements for the degree of Doctor of Philosophy in Ecology at Massey University, Auckland, New Zealand
    (Massey University, 2012) Machovsky-Capuska, Gabriel E.
    Australasian gannets (Morus serrator) are the second rarest member of the seabird group Sulidae. Among the three species of gannets worldwide, they are the only species that regularly breeds in southeastern Australia and New Zealand. Like all gannets, M. serrator face considerable challenges in foraging, relying on sparsely and patchily distributed pelagic prey, which move in a 3D environment. Whereas most predators are specialise hunters in one media, gannets have to hunt within a complex air-water interface. The aim of the present thesis is to examine the hunting strategies of Australasian gannets, with particular emphasis on how these birds use both aerial and aquatic adaptations to locate and capture prey. The acquisition of information concerning food sources was analysed using GPS data loggers, field observations and high resolution video footage. I tested the hypothesis that gannets obtain information of food resources from their partners using bill fencing as referential signals analogous to the waggle dance in honeybees (Apis mellifera) (Chapter 2). Results did not support this hypothesis but suggested that Australasian gannets use a combination of strategies, probably including memory that facilitates their return to locations where prey was previously captured (Chapter 3) and local enhancement to locate active feeding sites (Chapter 2). The impact of intraspecific competition for local resources was studied between large (Cape Kidnappers, 7,300 breeding pairs) and small (Farewell Spit, 3,900 breeding pairs) colonies in New Zealand using GPS data loggers (Chapter 3). Results indicated that gannets from the larger colony invested more in foraging (greater foraging times and foraging distances). This is consistent with previous studies of other gannet species, suggesting that M. serrator experience intraspecific competition for food when living in large colonies. Pelagic prey are able to evade predation by descending to depths beyond the reach of diving birds. Among the adaptations evolved by gannets for dealing with this challenge is plunge-diving, where the bird uses gravity in the aerial phase of the hunt to gain speed and momentum for descending into the water column. I conducted a fine scaled analysis using videography of the aerial and aquatic phases of this highly specialised hunting strategy. Analysis of the aerial phase (Chapter 4) showed that the initiation of plunge dives are synchronised among members of foraging groups, suggesting a form of group-level behaviour in which gannets might benefit from the sensory experiences (prey detection) of conspecifics. The analysis also showed that gannets adapt the aerial phase of their dives in presence vs. absence of heterospecific predators. In the aquatic phase (Chapter 5), gannets perform short and shallow V-shaped dives and long and deep U-shaped dives in pursuit of pelagic fish and squid. My findings revealed that gannets adjusted their dive shape in relation to the depth of their prey rather than prey type, as previously hypothesised. Although the maximum number of prey captured per dive by the gannets was higher than previously reported, reaching up to five fish in a single U-shaped dive, the results presented herein suggest that the two dive profiles were equally profitable. To examine the role of underwater vision in prey capture, I used underwater video footage, photokeratometry and infrared video photorefraction (Chapter 6). Analysis of video footage confirmed that there are two distinct phases in the underwater component of plunge dives in Australasian gannets, an initial phase in which the bird is propelled through the water column by the momentum of the plunge (M phase) and a phase in which it is actively propelled by wing flapping (WF phase). The highest prey capture rate was observed during the WF phase, a result that suggests the use of vision in underwater prey pursuit. I therefore used photokeratometry and video photorefraction to test whether gannets are able to adapt optically in the transition from aerial to aquatic media. My measurements showed that underwater visual accommodation in the gannets was attained within 2 - 3 frames (80 - 120 ms) of submergence, a remarkably short timescale in relation to the optics of most vertebrate eyes. The preceding chapters demonstrate some highly effective behavioural and sensory capacities used by gannets in foraging. In Chapter 7 I demonstrate evidence of fatal injuries due to collision between conspecifics in plunge-diving Australasian and Cape gannets (M. capensis). The analysis also revealed a case of attempted underwater kleptoparasitism, in which a diving bird targeted a previously captured fish in the beak of another gannet. This novel observation suggests a further challenge for hunting gannets, namely to retain prey following the capture.