Marine Ecology Progress Series
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DOI: https://doi.org/10.3354/meps15022
copiedFood-based attractant from wildlife tourism alters the residency and space use, but not the activity, of a non-focal fish
Wildlife tourism often uses food-based attractants to aggregate focal species. While this can adversely affect the focal species, there is limited understanding of its effects on non-focal species which often also respond to the attractant. At the Neptune Islands, South Australia, bait and berley are used to entice white sharks Carcharodon carcharias to cage-diving vessels, inadvertently attracting and feeding silver trevally Pseudocaranx georgianus. Feeding on the bait and berley affects their diet, but it is unknown whether it also affects their movements, behaviour, and energy budgets. We used a fine-scale acoustic telemetry array to monitor the movement and activity (i.e. proxy for energy expenditure calculated from 3-dimensional acceleration) of 25 tagged silver trevally in response to white shark tourism. The presence of cage-diving operators increased residency across weekly (+32%) and daily (+20%) scales, while reducing the core space by aggregating them at the surface (<5 m), close to food-based operators. Despite observations of silver trevally bursting towards tethered baits, changes in activity (acceleration, m s-1) were minimal compared to non-operator days. While food was available nearly daily, a thermally triggered seasonal exodus by 76% of individuals indicates that silver trevally still move and behave naturally. However, the supplemental nutrients and substantial time spent at Neptune Islands has the potential to alter the energy budgets of silver trevally. Our findings reveal the impacts that provisioning from wildlife tourism can have on non-focal schooling pelagic fish, supporting the need for management of wildlife tourism to account for the whole ecosystem when assessing the ecological sustainability of a tourism industry.
KEYWORDS
Anders N, Howarth K, Totland B, Handegard NO, Tenningen M, Breen M (2019) Effects on individual level behaviour in mackerel (Scomber scombrus) of sub-lethal capture related stressors: crowding and hypoxia. PLOS ONE 14:e0213709 Crossref
Barnett A, Abrantes KG, Seymour J, Fitzpatrick R (2012) Residency and spatial use by reef sharks of an isolated seamount and its implications for conservation. PLOS ONE 7:e36574 Crossref
Barnett A, Payne NL, Semmens JM, Fitzpatrick R (2016) Ecotourism increases the field metabolic rate of whitetip reef sharks. Biol Conserv 199:132-136 Crossref
Barry C, Legaspi C, Clarke TM, Araujo G and others (2023) Estimating the energetic cost of whale shark tourism. Biol Conserv 284:110164 Crossref
Bartoń K (2019) Package ‘MuMIn’. R package version, 1.46.0. Link
Battin J (2004) When good animals love bad habitats: ecological traps and the conservation of animal populations. Conserv Biol 18:1482-1491 Crossref
Bauer S, Hoye BJ (2014) Migratory animals couple biodiversity and ecosystem functioning worldwide. Science 344:1242552 Crossref
Behringer DC, Wood CL, Krkošek M, Bushek D (2020) Disease in fisheries and aquaculture. In: Behringer DC, Silliman BR, Lafferty KD (eds) Marine disease ecology. Oxford University Press, Oxford, p 183-210
Biebach H, Falk H, Krebs JR (1991) The effect of constant light and phase shifts on a learned time-place association in garden warblers (Sylvia borin): hourglass or circadian clock? J Biol Rhythms 6:353-365 Crossref
Bowler DE, Benton TG (2005) Causes and consequences of animal dispersal strategies: relating individual behaviour to spatial dynamics. Biol Rev Camb Philos Soc 80:205-225 Crossref
Boyce MS, Johnson CJ, Merrill EH, Nielsen SE, Solberg EJ, Van Moorter B (2016) Can habitat selection predict abundance? J Anim Ecol 85:11-20 Crossref
Brookhouse N, Bucher DJ, Rose K, Kerr I, Gudge S (2013) Impacts, risks and management of fish feeding at Neds Beach, Lord Howe Island Marine Park, Australia: a case study of how a seemingly innocuous activity can become a serious problem. J Ecotour 12:165-181 Crossref
Bruce BD, Bradford RW (2013) The effects of shark cage-diving operations on the behaviour and movements of white sharks, Carcharodon carcharias, at the Neptune Islands, South Australia. Mar Biol 160:889-907 Crossref
Brunnschweiler JM, Barnett A (2013) Opportunistic visitors: long-term behavioural response of bull sharks to food provisioning in Fiji. PLOS ONE 8:e58522 Crossref
Brunnschweiler J, Payne N, Barnett A (2018) Hand feeding can periodically fuel a major portion of bull shark energy requirements at a provisioning site in Fiji. Anim Conserv 21:31-35 Crossref
Burnham KP, Anderson DR (2004) Multimodel inference: understanding AIC and BIC in model selection. Sociol Methods Res 33:261-304 Crossref
Buschmann AH, Cabello F, Young K, Carvajal J, Varela DA, Henríquez L (2009) Salmon aquaculture and coastal ecosystem health in Chile: analysis of regulations, environmental impacts and bioremediation systems. Ocean Coast Manag 52:243-249 Crossref
Campbell HA, Watts ME, Dwyer RG, Franklin CE (2012) V-Track: software for analysing and visualising animal movement from acoustic telemetry detections. Mar Freshw Res 63:815-820 Crossref
Carlisle AB, Tickler D, Dale JJ, Ferretti F and others (2019) Estimating space use of mobile fishes in a large marine protected area with methodological considerations in acoustic array design. Front Mar Sci 6:256 Crossref
Champion C, Hobday AJ, Tracey SR, Pecl GT (2018) Rapid shifts in distribution and high-latitude persistence of oceanographic habitat revealed using citizen science data from a climate change hotspot. Glob Change Biol 24:5440-5453
Clarke C, Lea J, Ormond RF (2013) Changing relative abundance and behaviour of silky and grey reef sharks baited over 12 years on a Red Sea reef. Mar Freshw Res 64:909-919 Crossref
Clarke TM, Whitmarsh SK, Dwyer RG, Udyawer V, Pederson H, Huveneers C (2022) Effects of shark tourism on the daily residency and movements of a non-focal pelagic teleost. Mar Ecol Prog Ser 687:133-146 Crossref
Clarke TM, Whitmarsh SK, Champion C, Pederson H and others (2023) Influence of shark tourism on the activity and physiological condition of a non-focal pelagic fish. ICES J Mar Sci 80:1670-1682 Crossref
Cooper NW, Sherry TW, Marra PP (2014) Modeling 3-dimensional space use and overlap in birds. Auk 131:681-693 Crossref
Corcoran MJ, Wetherbee BM, Shivji MS, Potenski MD, Chapman DD, Harvey GM (2013) Supplemental feeding for ecotourism reverses diel activity and alters movement patterns and spatial distribution of the southern stingray, Dasyatis americana. PLOS ONE 8:e59235 Crossref
Corradini A, Peters W, Pedrotti L, Hebblewhite M and others (2021) Animal movements occurring during COVID-19 lockdown were predicted by connectivity models. Glob Ecol Conserv 32:e01895 Crossref
Dennis JD, Meyer L, Dudgeon CL, Huveneers C (2024) One fish, two fish, three fish, more: novel resighting method produces precise and cost-effective estimates of abundance. J Fish Biol 105:1603-1605 Crossref
Department of Environment Water (DEW) (2016) South Australian White Shark Tour Licensing Policy. Government of South Australia. cdn.environment.sa.gov.au/marineparks/docs/white-shark-tour-licensing-policy-gen.pdf
Drew JA, McKeon M (2019) Shark-based tourism presents opportunities for facultative dietary shift in coral reef fish. PLOS ONE 14:e0221781 Crossref
Ellis D, Kiessling I (2016) Ranching of southern bluefin tuna in Australia. In: Benetti DD, Partridge GJ, Buentello A (eds) Advances in tuna aquaculture: from hatchery to market. Elsevier, Amsterdam, p 217-232
Espinoza M, Farrugia TJ, Webber DM, Smith F, Lowe CG (2011) Testing a new acoustic telemetry technique to quantify long-term, fine-scale movements of aquatic animals. Fish Res 108:364-371 Crossref
Fairclough D, Potter I, Lek E, Bivoltsis A, Babcock R (2011) The fish communities and main fish populations of the Jurien Bay Marine Park. Strategic Research Fund for the Marine Environment Collaborative Research Project. Final Report. Centre for Fish and Fisheries Research, Murdoch University, Murdoch
Fitzpatrick R, Abrantes KG, Seymour J, Barnett A (2011) Variation in depth of whitetip reef sharks: Does provisioning ecotourism change their behaviour? Coral Reefs 30:569-577 Crossref
Forrester GE, Steele MA (2004) Predators, prey refuges, and the spatial scaling of density-dependent prey mortality. Ecology 85:1332-1342 Crossref
Fowler AM, Chick RC, Stewart J (2018) Patterns and drivers of movement for a coastal benthopelagic fish, Pseudocaranx georgianus, on Australia’s southeast coast. Sci Rep 8:16738 Crossref
Future Market Insights (2022) Wildlife tourism market snapshot (2022-2032). Link
Gervais CR, Champion C, Pecl GT (2021) Species on the move around the Australian coastline: a continental-scale review of climate-driven species redistribution in marine systems. Glob Change Biol 27:3200-3217 Crossref
Gooden A, Clarke TM, Meyer L, Huveneers C (2024) Wildlife tourism has little energetic impact on the world’s largest predatory shark. Anim Behav 207:247-265 Crossref
Guerra AS, Kao AB, Mccauley DJ, Berdahl AM (2020) Fisheries-induced selection against schooling behaviour in marine fishes. Proc R Soc B 287:20201752 Crossref
Hale R, Swearer SE (2016) Ecological traps: current evidence and future directions. Proc R Soc B 283:20152647 Crossref
Hansen MF, Kaburu SSK, Morrow KS, Maréchal L (2023) Primate tourism. In: McKinney T, Waters S, Rodrigues MA (eds) Primates in anthropogenic landscapes: exploring primate behavioural flexibility across human contexts. Springer, p 183-201
Heinrich D, Dhellemmes F, Guttridge TL, Smukall M and others (2021) Short-term impacts of daily feeding on the residency, distribution and energy expenditure of sharks. Anim Behav 172:55-71 Crossref
Heupel MR, Hueter RE (2002) Importance of prey density in relation to the movement patterns of juvenile blacktip sharks (Carcharhinus limbatus) within a coastal nursery area. Mar Freshw Res 53:543-550 Crossref
Hijmans RJ, Williams E, Vennes C, Hijmans MRJ (2017) Package ‘geosphere’: spherical trigonometry, version 1.5-20. cran.r-project.org/package=geosphere
Hines KN (2011) Effects of ecotourism on endangered northern Bahamian rock iguanas (Cyclura cychlura). Herpetol Conserv Biol 6:250-259
Hussey NE, Kessel ST, Aarestrup K, Cooke SJ and others (2015) Aquatic animal telemetry: a panoramic window into the underwater world. Science 348:1255642 Crossref
Huveneers C, Meekan MG, Apps K, Ferreira LC, Pannell D, Vianna GMS (2017) The economic value of shark-diving tourism in Australia. Rev Fish Biol Fish 27:665-680 Crossref
Huveneers C, Watanabe YY, Payne NL, Semmens JM (2018) Interacting with wildlife tourism increases activity of white sharks. Conserv Physiol 6:coy019 Crossref
Huveneers C, Jaine FRA, Barnett A, Butcher PA and others (2021) The power of national acoustic tracking networks to assess the impacts of human activity on marine organisms during the COVID-19 pandemic. Biol Conserv 256:108995 Crossref
Jensen ALM, Delfour F, Carter T (2013) Anticipatory behavior in captive bottlenose dolphins (Tursiops truncatus): a preliminary study. Zoo Biol 32:436-444 Crossref
Kassamali-Fox A, Christiansen F, May-Collado LJ, Ramos EA, Kaplin BA (2020) Tour boats affect the activity patterns of bottlenose dolphins (Tursiops truncatus) in Bocas del Toro, Panama. PeerJ 8:e8804 Crossref
Keenleyside MHA (1955) Some aspects of the schooling behaviour of fish. Behaviour 8:183-248 Crossref
Klarevas-Irby JA, Farine DR (2024) Diel patterns of movement reveal temporal strategies during dispersal. Anim Behav 207:119-129 Crossref
Lassis R, Festa-Bianchet M, Pelletier F (2022) Breeding migrations by bighorn sheep males are driven by mating opportunities. Ecol Evol 12:e8692 Crossref
Lima JS, Zalmon IR, Love M (2019) Overview and trends of ecological and socioeconomic research on artificial reefs. Mar Environ Res 145:81-96 Crossref
Logan RK, Anderson JM, Burns ES, Chacon YS and others (2024) Long-term tracking captures the timing of ontogenetic niche shifts in northeast Pacific white sharks. Ecosphere 15:e70034 Crossref
Lusseau D (2004) The hidden cost of tourism: detecting long-term effects of tourism using behavioral information. Ecol Soc 9:2 Crossref
Magurran AE (1990) The adaptive significance of schooling as an anti-predator defence in fish. Ann Zool Fenn 27:51-66
Marques AT, Santos CD, Hanssen F, Muñoz AR and others (2020) Wind turbines cause functional habitat loss for migratory soaring birds. J Anim Ecol 89:93-103 Crossref
Matyjasiak P, Chacińska P, Książka P (2023) Anthropogenic noise interacts with the predation risk assessment in a free-ranging bird. Curr Zool 70:277-283 Crossref
McBride RS, Somarakis S, Fitzhugh GR, Albert A and others (2015) Energy acquisition and allocation to egg production in relation to fish reproductive strategies. Fish Fish 16:23-57 Crossref
Meyer L, Pethybridge H, Beckmann C, Bruce B, Huveneers C (2019) The impact of wildlife tourism on the foraging ecology and nutritional condition of an apex predator. Tour Manag 75:206-215 Crossref
Meyer L, Whitmarsh SK, Nichols PD, Revill AT, Huveneers C (2020) The effects of wildlife tourism provisioning on non-target species. Biol Conserv 241:108317 Crossref
Meyer L, Barry C, Araujo G, Barnett A and others (2022) Redefining provisioning in marine wildlife tourism. J Ecotour 21:210-229 Crossref
Milazzo M, Anastasi I, Willis TJ (2006) Recreational fish feeding affects coastal fish behavior and increases frequency of predation on damselfish Chromis chromis nests. Mar Ecol Prog Ser 310:165-172
Moorcroft PR, Barnett A (2008) Mechanistic home range models and resource selection analysis: a reconciliation and unification. Ecology 89:1112-1119 Crossref
Morales JM, Moorcroft PR, Matthiopoulos J, Frair JL and others (2010) Building the bridge between animal movement and population dynamics. Philos Trans R Soc B 365:2289-2301 Crossref
Nakagawa S, Schielzeth H (2013) A general and simple method for obtaining R2 from generalized linear mixed-effects models. Methods Ecol Evol 4:133-142 Crossref
Nazimi L, Robbins WD, Schilds A, Huveneers C (2018) Comparison of industry-based data to monitor white shark cage-dive tourism. Tour Manag 66:263-273 Crossref
Neuenfeldt S, Bartolino V, Orio A, Andersen KH and others (2020) Feeding and growth of Atlantic cod (Gadus morhua L.) in the eastern Baltic Sea under environmental change. ICES J Mar Sci 77:624-632 Crossref
Ngoprasert D, Lynam AJ, Gale GA (2017) Effects of temporary closure of a national park on leopard movement and behaviour in tropical Asia. Mamm Biol 82:65-73 Crossref
Niella Y, Udyawer V, Drew M, Simes B, Pederson H, Huveneers C (2023) Multi-year effects of wildlife tourism on shark residency and implications for management. Mar Pol 147:05362
Niella Y, Meyer L, Clarke TM, Dennis JD, Pederson H, Huveneers C (2024) Effects of wildlife tourism on white shark associative behaviour. Anim Behav 215:227-239 Crossref
Orrell DL, Hussey NE (2022) Using the VEMCO Positioning System (VPS) to explore fine-scale movements of aquatic species: applications, analytical approaches and future directions. Mar Ecol Prog Ser 687:195-216 Crossref
Ouboter DA, Kadosoe VS, Ouboter PE (2021) Impact of ecotourism on abundance, diversity and activity patterns of medium-large terrestrial mammals at Brownsberg Nature Park, Suriname. PLOS ONE 16:e0250390 Crossref
Parsons TR, Takahashi M, Hargrave B (2013) Biological oceanographic processes. Elsevier, Oxford
Payne NL, Gillanders BM, Seymour RS, Webber DM, Snelling EP, Semmens JM (2011) Accelerometry estimates field metabolic rate in giant Australian cuttlefish Sepia apama during breeding. J Anim Ecol 80:422-430 Crossref
Pini-Fitzsimmons J, Knott NA, Brown C (2018) Effects of food provisioning on site use in the short-tail stingray Bathytoshia brevicaudata. Mar Ecol Prog Ser 600:99-110 Crossref
Pitcher TJ, Parrish JK (1993) Functions of shoaling behaviour in teleosts. In: Pitcher TJ (ed) The behaviour of teleost fishes, 2nd edn. Springer, Dordrecht, p 294-337
Polis GA, Anderson WB, Holt RD (1997) Toward an integration of landscape and food web ecology: the dynamics of spatially subsidized food webs. Annu Rev Ecol Syst 28:289-316
R Core Team (2016) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna
Rees MJ, Knott NA, Davis TR, Davis AR and others (2021) Temporal stability in a protected and isolated fish community within marine parks surrounding Lord Howe Island. Reg Stud Mar Sci 48:102038 Crossref
Rider MJ, Mcdonnell LH, Hammerschlag N (2021) Multi-year movements of adult and subadult bull sharks (Carcharhinus leucas): philopatry, connectivity, and environmental influences. Aquat Ecol 55:559-577 Crossref
Rizzari JR, Semmens JM, Fox A, Huveneers C (2017) Observations of marine wildlife tourism effects on a non-focal species. J Fish Biol 91:981-988 Crossref
Robinson WD, Bowlin MS, Bisson I, Shamoun-Baranes J and others (2010) Integrating concepts and technologies to advance the study of bird migration. Front Ecol Environ 8:354-361 Crossref
Rowling KR, Raines LP (2000) Description of the biology and an assessment of the fishery for silver trevally Pseudocaranx dentex off New South Wales. NSW Fisheries Research Institute, Cronulla
Schatz B, Lachaud JP, Beugnon G (1999) Spatio-temporal learning by the ant Ectatomma ruidum. J Exp Biol 202:1897-1907 Crossref
Scheyvens R (1999) Ecotourism and the empowerment of local communities. Tour Manag 20:245-249 Crossref
Semeniuk CA (2009) Unifying ecological and social sciences into a management framework for wildlife-based tourism: a case study of feeding stingrays as a marine tourism attraction in the Cayman Islands. PhD thesis, Simon Fraser University, Burnaby
Semeniuk CAD, Rothley KD (2008) Costs of group-living for a normally solitary forager: effects of provisioning tourism on southern stingrays Dasyatis americana. Mar Ecol Prog Ser 357:271-282 Crossref
Sengupta A, Radhakrishna S (2018) The hand that feeds the monkey: mutual influence of humans and rhesus macaques (Macaca mulatta) in the context of provisioning. Int J Primatol 39:817-830 Crossref
Senigaglia V, Christiansen F, Bejder L, Sprogis K, Cantor M (2022) Human food provisioning impacts the social environment, home range and fitness of a marine top predator. Anim Behav 187:291-304 Crossref
Sequeira AM, Heupel MR, Lea MA, Eguíluz VM and others (2019) The importance of sample size in marine megafauna tagging studies. Ecol Appl 29(6):e01947
Shankar A, Graham CH, Canepa JR, Wethington SM, Powers DR (2019) Hummingbirds budget energy flexibly in response to changing resources. Funct Ecol 33:1904-1916 Crossref
Simpfendorfer CA, Heupel MR, Hueter RE (2002) Estimation of short-term centers of activity from an array of omnidirectional hydrophones and its use in studying animal movements. Can J Fish Aquat Sci 59:23-32 Crossref Crossref
Simpfendorfer CA, Olsen EM, Heupel MR, Moland E (2012) Three-dimensional kernel utilization distributions improve estimates of space use in aquatic animals. Can J Fish Aquat Sci 69:565-572 Crossref
Smith F (2013) Understanding HPE in the VEMCO positioning system (VPS). Link
Spinu V, Grolemund G, Wickham H (2018) lubridate: make dealing with dates a little easier. R package, version 1.7. 4. cran.r-project.org/package=lubridate
Staniland IJ, Ratcliffe N, Trathan PN, Forcada J (2018) Long term movements and activity patterns of an Antarctic marine apex predator: the leopard seal. PLOS ONE 13:e0197767 Crossref
Svane I, Barnett J (2008) The occurrence of benthic scavengers and their consumption at tuna farms off Port Lincoln, South Australia. J Exp Mar Biol Ecol 363:110-117 Crossref
Tabak MA, Piaggio AJ, Miller RS, Sweitzer RA, Ernest HB (2017) Anthropogenic factors predict movement of an invasive species. Ecosphere 8:e01844 Crossref
Teske PR, Emami-Khoyi A, Golla TR, Sandoval-Castillo J and others (2021) The sardine run in southeastern Africa is a mass migration into an ecological trap. Sci Adv 7:eabf4514
Trave C, Brunnschweiler J, Sheaves M, Diedrich A, Barnett A (2017) Are we killing them with kindness? Evaluation of sustainable marine wildlife tourism. Biol Conserv 209:211-222 Crossref
Udyawer V, Simpfendorfer CA, Heupel MR (2015) Diel patterns in three-dimensional use of space by sea snakes. Anim Biotelem 3:29 Crossref
Udyawer V, Dwyer RG, Hoenner X, Babcock RC and others (2018) A standardised framework for analysing animal detections from automated tracking arrays. Anim Biotelem 6:17 Crossref
Vagabov MM, Gadzhiev MD, Eldarov EM (2019) Development of hunting tourism in Dagestan: problems and prospects. Ponte 75:24-34 Crossref
Vignon M, Sasal P, Johnson RL, Galzin R (2010) Impact of shark-feeding tourism on surrounding fish populations off Moorea Island (French Polynesia). Mar Freshw Res 61:163-169
Volkoff H, Ronnestad I (2020) Effects of temperature on feeding and digestive processes in fish. Temperature 7:307-320 Crossref
Wearing S, Neil J (2009) Ecotourism. Routledge, London
Wen CK, Chen KS, Tung WC, Chao A, Wang CW, Liu SL, Ho MJ (2019) The influence of tourism-based provisioning on fish behavior and benthic composition. Ambio 48:779-789 Crossref
Williams R, Lusseau D, Hammond PS (2006) Estimating relative energetic costs of human disturbance to killer whales (Orcinus orca). Biol Conserv 133:301-311 Crossref
Wood S, Wood MS (2015) Package ‘mgcv’. R package , version 1.8-40. cran.r-project.org/package=mgcv
Joshua D. Dennis (Corresponding Author)
- Southern Shark Ecology Group, College of Science and Engineering, Flinders University, Bedford Park, Adelaide, South Australia 5100, Australia
- Southern Shark Ecology Group, College of Science and Engineering, Flinders University, Bedford Park, Adelaide, South Australia 5100, Australia
- The Georgia Aquarium, Atlanta 30313, Georgia
Thomas M. Clarke (Co-author)
- Southern Shark Ecology Group, College of Science and Engineering, Flinders University, Bedford Park, Adelaide, South Australia 5100, Australia
Hugh Pederson (Co-author)
- Innovasea, Bedford, Nova Scotia B4B 0L9, Canada
- Southern Shark Ecology Group, College of Science and Engineering, Flinders University, Bedford Park, Adelaide, South Australia 5100, Australia
Handling Editor:
Elliott Hazen, Pacific Grove, California, USA
Reviewers:
E. E. Clua and 2 anonymous referees
Acknowledgements:
We thank the staff of the white shark cage-diving industry for their assistance with fieldwork. Funding and support were provided by the Holsworth Wildlife Research Endowment, and the 2020 DEW Marine and Research Scholarship in memory of Peter Fairweather. This project was conducted under the following permits: DEW permit Q26612-4 and Q26612-6; Animal Ethics permit AEB1698-7 and AEB1537/2019/1.