Modelling the Behaviour State of Humpback Whales Megaptera Novaeangliae in Response to Vessel Presence Off Sydney, Australia
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Vol. 15: 255–264, 2011 ENDANGERED SPECIES RESEARCH Published online December 12 doi: 10.3354/esr00380 Endang Species Res Modelling the behaviour state of humpback whales Megaptera novaeangliae in response to vessel presence off Sydney, Australia Maryrose Gulesserian1,*, David Slip1,2, Gillian Heller3, Robert Harcourt1 1Marine Mammal Research Group, Graduate School of the Environment, Macquarie University, Sydney, Australia 2109 2Taronga Conservation Society Australia, Sydney, Australia 2088 3Department of Statistics, Macquarie University, Sydney, Australia 2109 ABSTRACT: The largest southern hemisphere humpback whale Megaptera novaeangliae stock (E1) uses the east coast of Australia as a migratory corridor to travel between their high-latitude feeding grounds in the Southern Ocean and low-latitude breeding grounds in northeast Queens- land and the south-west Pacific Ocean. The population is recovering at close to the maximum rate of growth (rm), and the increasing abundance of whales passing within sight of land has facilitated the development of a growing land- and vessel-based whale watching industry. We observed the behaviour of 156 individual pods of humpback whales passing Sydney, New South Wales, during their 2006 and 2007 northern migration and monitored vessel−whale interactions with respect to the Australian National Guidelines for Whale and Dolphin Watching 2005. We applied gener- alised linear mixed models with random effects to compute the odds of changing to the current behaviour state. We found that in the presence of vessels, whales were more likely to remain on the surface breathing or to cease surface breathing and switch to generally short, shallow diving than was the case when no vessels were present. Northerly migrating whales off Sydney were more likely to remain on the surface breathing in the presence of vessels, rather than taking some form of vertical avoidance (deep, long dives) as reported elsewhere. Given the high rate of popu- lation increase of stock E1 and the low level of behavioural changes seen, it appears that for this population at least, adult humpback whales migrating to their breeding grounds are relatively robust to disturbance by whale watching. KEY WORDS: Humpback whale · Migration · Boat disturbance · Odds ratios · Behaviour · Whale watching Resale or republication not permitted without written consent of the publisher INTRODUCTION geted animals respond to disturbances. Frequently documented responses include short-term changes in Commercial viewing of wildlife is rapidly becoming behaviour, such as increased vigilance, relocation, the predominant economic use of large wild animals ceasing or otherwise changing behaviour states and in the natural environment (Newsome et al. 2005). In flight initiation. These are shown across a range of particular, charismatic animals such as whales, pen- taxa, including birds (e.g. Galicia & Baldassarre guins, bears, whale sharks, apes and elephants have 1997), reptiles (e.g. Kerr et al. 2004), amphibians (e.g. become the focal point of many viewing operations Rodríguez-Prieto & Fernández-Juricic 2005) and a (Newsome et al. 2005). The concomitant proliferation number of terrestrial and marine mammals (e.g. Chi of wildlife tourism has raised inquiry into how tar- & Gilbert 1999, Cassini 2001, Pelletier 2006). *Email: [email protected] © Inter-Research 2011 · www.int-res.com 256 Endang Species Res 15: 255–264, 2011 Wildlife viewing has been marketed as non-con- peated disturbances is more difficult to understand. sumptive, leading to the assumption that this form of Only a few studies have linked short-term changes tourism is somewhat benign, with no impact on the to long-term effects, mainly in small cetaceans. For targeted animals (Jelinski et al. 2002), as well as example, repeated disturbance has led to habitat carrying an implicit presumption of long-term sus- abandonment (e.g. Bejder et al. 2006b), sensitisation tainability. However, these assumptions may require (e.g. Constantine 2001) or reduced reproductive careful evaluation, as for some animal species there success (e.g. Bejder & Samuels 2003) in some dol- may be causative links between tourism and distur- phin populations. However, while Watkins (1986) did bance (e.g. Holmes et al. 2005). While some animals find long-term changes in baleen whale behaviour become habituated to human activity (e.g. Coleman in the face of repeated whale watching excursions et al. 2008), others have remained wary and avoid over a 25 yr period, he concluded that for most spe- areas of high levels of human activity (e.g. Dyke et al. cies, the main effect was habituation, or even attrac- 1986). Continually responding to disturbance may tion in the case of humpback whales. While it is diffi- impact activity budgets and reduce the amount of cult to link disturbance to whales during migration to time an animal spends foraging, resting, and for impacts at the population level, it is possible that mammals, suckling young (e.g. Yalden & Yalden excessive interference from vessels may have impli- 1990). Moreover, moving away from a disturbance cations such as energetic costs, as whales adjust their may displace animals from preferred areas (e.g. behaviour. Bryant et al. 1984). One approach to evaluating the effects of vessel The rapid growth of the whale watching industry presence on cetaceans involves assessing sequences has raised concerns about the potential impacts the of behaviour states (Lusseau 2003). Because the industry may have on these animals, particularly observed behaviour state is influenced by the pre- because most activities take place in coastal areas ceding state, behaviour sequence data can be partic- and take advantage of seasonal or resident popula- ularly useful for modelling changes in these states tions. Responses to disturbance have been reported due to external factors, such as vessel presence. Vari- in many cetacean species, including humpback whales ations of this method have been demonstrated in a Megaptera novaeangliae (e.g. Bauer et al. 1993), small number of studies on odontocetes, including killer whales Orcinus orca (e.g. Bain et al. 2006, bottlenose dolphins (Lusseau 2003), dusky dolphins Williams & Ashe 2007), bottlenose dolphins Tursiops (Dans et al. 2008), common dolphins (Stockin et al. spp. (e.g. Constantine et al. 2004, Lemon et al. 2006), 2008) and killer whales (Williams et al. 2006). Very fin whales Balaenoptera physalus (e.g. Stone et al. few studies have previously documented sequential 1992), gray whales Eschrichtius robustus (e.g. Moore behaviour states of mysticetes (e.g. Schaffar et al. & Clarke 2002), common dolphins Delphinus spp. 2009). (e.g. Stockin et al. 2008), dusky dolphins Lagen - Humpback whales are medium-sized mysticete orhynchus obscurus (e.g. Barr & Slooten 1999), whales with coastal migration in parts of their range. Hector’s dolphins Cephalorhynchns hectori (e.g. Humpback whales from the E1 (eastern Australia) Bejder et al. 1999) and sperm whales Macrocephalus breeding stock migrate up the east coast of Australia physeter (e.g. Richter et al. 2006). in the austral winter to breed before returning south Assessing the effects of whale watching activities to feed with newborn calves (Chittleborough 1965, on cetaceans is difficult, as changes in behaviour are Brown et al. 1995, Noad et al. 2010). This population expected to be subtle and difficult to detect because was heavily decimated by coastal whaling in the of the underwater nature of these species. Cetaceans 1950s, and the population collapsed by 1962 (Chittle- may display a variety of short-term responses and borough 1965). Post-whaling surveys of the E1 popu- strategies during vessel interactions, ranging from lation were initiated in Queensland in 1978, and the changes in pod composition (e.g. Ribeiro et al. 2005), population is now recovering at close to the theoreti- movement patterns and habitat use (e.g. Bejder et al. cal reproductive limit of the species, with an absolute 2006a, Lusseau 2006a,b), surfacing and dive times abundance exceeding 14 000 (Paterson et al. 1994, (e.g. Blane & Jaakson 1994), activity and energy Noad et al. 2010). During their northward migration, budgets (e.g. Williams et al. 2006), changes in the whales pass very close to shore off several major swim speed (e.g. Jelinski et al. 2002) and changes urban centres, from where many commercial whale in surface behaviour (e.g. Scheidat et al. 2004). watching vessels operate. While short-term changes in behaviour are relatively The fastest growth in whale watching in Australia easy to identify, the biological significance of re- has occurred in New South Wales (NSW), with an Gulesserian et al.: Modelling humpback whale behaviour 257 estimated average annual growth of 14.7% between 1998 and 2008 (O’Connor et al. 2009). Over the last decade, a significant whale watching industry has developed based out of Sydney, and in 2008 alone, over 25 000 people boarded dedicated vessels in Sydney for the purpose of whale watching (IFAW * 2004, O’Connor et al. 2009). To manage the rapidly growing industry, the Australian National Guide- SYDNEY lines for Whale and Dolphin Watching 2005 were developed cooperatively by all Australian state and territory governments in order to form a basis by which states and territories could enact regulations nationwide providing for safe and sustainable cetacean watching in all Australian waters. NSW was the first state to implement these guidelines as Observation state regulations (NSW National Parks and Wildlife Botany Bay area Amendment [Marine Mammals] Regulation 2006). Although the guidelines provide defined approach speeds, distances and angles, they are not based on quantitative evidence. A greater understanding of the behavioural impacts of interactions between cetaceans and vessels is needed to assess the effi- Cape Solander cacy of the existing guidelines and to provide an empirical basis for these and any subsequent amendments. In this paper, we assess whether the presence of vessels alters sequential changes in surface behav- iour of migrating humpback whales on their northern migration to the breeding grounds. 0 2 4 8 km N MATERIALS AND METHODS Fig.