Research Article: Estimating Cetacean Carrying Capacity Based on Spacing Behaviour

Date Published: December 7, 2012

Publisher: Public Library of Science

Author(s): Janelle E. Braithwaite, Jessica J. Meeuwig, K. Curt S. Jenner, Brock Fenton.


Conservation of large ocean wildlife requires an understanding of how they use space. In Western Australia, the humpback whale (Megaptera novaeangliae) population is growing at a minimum rate of 10% per year. An important consideration for conservation based management in space-limited environments, such as coastal resting areas, is the potential expansion in area use by humpback whales if the carrying capacity of existing areas is exceeded. Here we determined the theoretical carrying capacity of a known humpback resting area based on the spacing behaviour of pods, where a resting area is defined as a sheltered embayment along the coast. Two separate approaches were taken to estimate this distance. The first used the median nearest neighbour distance between pods in relatively dense areas, giving a spacing distance of 2.16 km (±0.94). The second estimated the spacing distance as the radius at which 50% of the population included no other pods, and was calculated as 1.93 km (range: 1.62–2.50 km). Using these values, the maximum number of pods able to fit into the resting area was 698 and 872 pods, respectively. Given an average observed pod size of 1.7 whales, this equates to a carrying capacity estimate of between 1187 and 1482 whales at any given point in time. This study demonstrates that whale pods do maintain a distance from each other, which may determine the number of animals that can occupy aggregation areas where space is limited. This requirement for space has implications when considering boundaries for protected areas or competition for space with the fishing and resources sectors.

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An important consideration for conservation is the population size that a given habitat can support. Estimating this carrying capacity provides a baseline against which changes to habitat can be assessed with respect to the maintenance of conservation values [1]. Here, carrying capacity is defined in terms of density limitation in a particular area at a given time, rather than the overall population carrying capacity (K) [2]. The limit to animal density in an area is generally related to the total amount of resources available in the habitat and the resource needs of each individual. It is well recognized that density scales inversely with body size across many plant and animal communities [3]–[6], as does home-range size in top predators [6]–[8]. Individual energy demand is the main explanation for these trends, with larger animals requiring more food and thus a larger area for foraging. Therefore, carrying capacity is often calculated based on food supply [9], [10]: for example, the estimated carrying capacity of sites used by migratory birds is calculated using a ‘daily ration model’, whereby the total consumable food of the site is divided by the individual energetic requirement [1], [10], [11]. However, this conventional approach to calculating carrying capacity is limited, and other studies have found that carrying capacity can also be influenced by predation risk [12], freshwater availability [13], shelter [14], and the availability of nesting sites [15]. As the space requirement of an animal, for example its home range, is generally related to the availability of resources, space itself can be considered as a resource that will limit density.

A total of 703 individual whales were sighted in the Exmouth Gulf region during the 2004–2005 aerial surveys (Table 1). The abundance estimates for each flight, using distance sampling, showed a maximum of 459 whales within the Gulf at any one time, and a total of 1270 whales over the entire period (CI 670–2080), assuming a maximum two week residency period for each whale (KCS Jenner, estimated from photo ID re-sights; each whale is represented only once in the total estimate). The abundance of whales in the Gulf clearly changed over time (Fig. 4a), with whales beginning to enter the Gulf from the north around the first week of August, peaking at the end September, before departing until the start of November. The number of calves within the Gulf follows a similar temporal pattern, but peaks about a week or two after the main migration (Fig. 4b), in early October.

Our premise is that in limited-space conditions the carrying capacity of an area for resting humpback whales is linked to the space requirement of the animals that occupy it, rather than more typically encountered pressures such as competition for food, and predator avoidance. Our results suggest that pods do maintain a distance from each other under relatively high-density conditions, demonstrating that space itself is a resource for these animals and that this space can be determined. We then used this spacing distance to calculate the theoretical carrying capacity of a humpback whale resting area. The implications of having a capacity limit, under the currently increasing population of WA humpback whales, can only be assessed once the current use of Exmouth Gulf is understood. Therefore, we also investigated how the humpback whale population presently uses Exmouth Gulf, both spatially and temporally.