Animal Biotelemetry | |
Novel foraging strategies observed in a growing leopard seal (Hydrurga leptonyx) population at Livingston Island, Antarctic Peninsula | |
Douglas J Krause3  Michael E Goebel2  Gregory J Marshall1  Kyler Abernathy1  | |
[1] National Geographic Society, Remote Imaging Group, 1145 17th Street NW, Washington 20036, DC, USA | |
[2] Antarctic Ecosystem Research Division, NOAA-NMFS-SWFSC, 8901 La Jolla Shores Drive, La Jolla 92037, CA, USA | |
[3] Scripps Institution of Oceanography, University of California San Diego, 9500 Gilman Drive, La Jolla 92037, CA, USA | |
关键词: T-LoCoH; CRITTERCAM; Fastloc GPS; Scavenging; Food caching; Kleptoparasitism; Hunting tactics; Apex predator; Leopard seal; | |
Others : 1224611 DOI : 10.1186/s40317-015-0059-2 |
|
received in 2015-01-14, accepted in 2015-05-04, 发布年份 2015 | |
【 摘 要 】
Background
Leopard seals are apex predators that can alter the community structure of Antarctic coastal ecosystems. Previous behavioral studies were limited to land-based, daytimeobservations of foraging leopard seals. Consequently, foraging tactics, social behaviors, and indirect ecosystem impacts are poorly understood. Here, we present the first analysis of animal-borne HD video footage for foraging leopard seals. Each CRITTERCAM was deployed with Fastloc GPS and time-depth recorder instruments providing fine-scale habitat context for observed foraging behavior. We analyzed seven deployments obtained in January and February of 2013 and 2014 from adult female leopard seals near mesopredator breeding colonies on Livingston Island, Antarctica.
Results
The average deployment length was 4.80 ± 2.45 (range 0.86–9.12) days, which covered a total of 16 foraging trips. Habitat use, along with 39 prey capture attempts, and 11 leopard seal social encounters were scored from 50.3 h of video data. We obtained 3,833 post-filter GPS positions, accurate to within 70 m, and the mean dive depth was 14.84 ± 8.98 m. Leopard seal foraging focused on four prey items: Antarctic fur seals, Antarctic fur seal pups, pygoscelid penguins, and demersal notothen fishes. Ambush tactics used only by a subset of leopard seals drove high capture success rates of fur seal pups. We identified novel prey-specific foraging tactics including stalking and flushing notothen fishes.
Conclusions
Leopard seals have been described as generalist apex predators; however, video and movement data suggest that leopard seals employ specialized prey-specific hunting tactics. Although preliminary, our findings indicate that leopard seals can affect coastal ecosystems through pathways beyond direct predation, including intraspecific kleptoparasitism and facultative scavenging/food caching. Our results suggest that position-integrated video data will be vital in quantifying the ecological impact of this abundant and versatile apex predator.
【 授权许可】
2015 Krause et al.
【 预 览 】
Files | Size | Format | View |
---|---|---|---|
20150912010711710.pdf | 2325KB | download | |
Fig.7. | 21KB | Image | download |
Fig.6. | 36KB | Image | download |
Fig.5. | 73KB | Image | download |
Fig.4. | 20KB | Image | download |
Fig.3. | 21KB | Image | download |
Fig.2. | 13KB | Image | download |
Fig.1. | 25KB | Image | download |
【 图 表 】
Fig.1.
Fig.2.
Fig.3.
Fig.4.
Fig.5.
Fig.6.
Fig.7.
【 参考文献 】
- [1]Estes J, Tinker M, Williams T, Doak D: Killer whale predation on sea otters linking oceanic and nearshore ecosystems. Science 1998, 282:473-476.
- [2]Williams T, Estes J, Doak D, Springer A: Killer appetites: assessing the role of predators in ecological communities. Ecology 2004, 85:3373-3384.
- [3]Estes JA, Terborgh J, Brashares JS, Power ME, Berger J, Bond WJ, et al.: Trophic downgrading of planet earth. Science 2011, 333:301-306.
- [4]Laws R: Seals. In Antarctic ecology. Edited by Laws RM. Academic Press, Cambridge; 1984:621-716.
- [5]Boveng P, Hiruki L, Schwartz M, Bengtson J: Population growth of Antarctic fur seals: limitation by a top predator, the leopard seal? Ecology 1998, 79:2863-2877.
- [6]Schwarz LK, Goebel ME, Costa DP, Kilpatrick AM: Top-down and bottom-up influences on demographic rates of Antarctic fur seals Arctocephalus gazella. J Anim Ecol 2013, 82:903-911.
- [7]Kooyman GL, Croll D, Stone S, Smith S: Emperor penguin colony at Cape Washington, Antarctica. Polar Record 1990, 26:103-108.
- [8]Ainley DG, Ballard G, Karl BJ, Dugger KM: Leopard seal predation rates at penguin colonies of different size. Antarct Sci 2005, 17:335-340.
- [9]Ballard WB, Carbyn LN, Smith DW: Wolf interactions with non-prey. In Wolves: behavior, ecology and conservation. Edited by Mech LD, Smith DW. The University of Chicago Press, Chicago; 2003:259-271.
- [10]Creel S, Christianson D, Liley S, Winnie JA: Predation risk affects reproductive physiology and demography of Elk. Science 2007, 315:960.
- [11]Creel S, Christianson D: Relationships between direct predation and risk effects. Trends Ecol Evol 2008, 23:194-201.
- [12]Wirsing AJ, Heithaus MR, Frid A, Dill LM: Seascapes of fear: evaluating sublethal predator effects experienced and generated by marine mammals. Mar Mamm Sci 2008, 24:1-15.
- [13]Walker T, Boyd I, McCafferty D, Taylor R, Reid K: Seasonal occurrence and diet of leopard seals (Hydrurga leptonyx) at Bird Island, South Georgia. Antarct Sci 1998, 10:75-81.
- [14]Casaux R, Baroni A, Ramón A, Carlini A, Bertolin M, DiPrinzio C: Diet of the leopard seal (Hydrurga leptonyx) at the Danco Coast, Antarctic Peninsula. Polar Biol 2009, 32:307-310.
- [15]Siniff DB, Stone S: The role of the leopard seal in the tropho-dynamics of the Antarctic marine ecosystem. In Antarctic nutrient cycles and food webs. Edited by Siegfried WR, Condy PR, Laws RM. Springer Verlag, Berlin; 1985:555-560.
- [16]Green K, Williams R: Observations on food remains in faeces of elephant, leopard and crabeater seals. Polar Biol 1986, 6:43-45.
- [17]Hiruki L, Schwartz M, Boveng P: Hunting and social behaviour of leopard seals (Hydrurga leptonyx) at Seal Island, South Shetland Islands, Antarctica. J Zool 1999, 249:97-109.
- [18]Vera C, Vargas R, Torres DN: Estrategias depredatorias del la foca leopardo sobre cachorros de lobo fino Antarctico. Boletin Antarctico Chileno 2005, 24:12-17.
- [19]Penney RL, Lowry G: Leopard seal predation of adelie penguins. Ecology 1967, 48:878-882.
- [20]Kooyman GL: Leopard seals of Cape Crozier. Animals 1965, 6:58-63.
- [21]Rogers T, Bryden MM: Predation of Adélie penguins (Pygoscelis adeliae) by leopard seals (Hydrurga leptonyx) in Prydz Bay, Antarctica. Can J Zool 1995, 73:1001-1004.
- [22]Muller-Schwarze D, Muller-Schwarze C: Relations between leopard seals and Adélie penguins. Rapports et Proces-Verbaux des Reunions du Conseil Permanent International pour l’Exploration de la Mer 1975, 169:394-404.
- [23]Bertram BCR: Serengeti predators and their social systems. In Serengeti: dynamics of an ecosystem. Edited by Sinclair ARE, Norton-Griffiths M. The University of Chicago Press, Chicago; 1979:221-248.
- [24]Macdonald DW: The ecology of carnivore social behaviour. Nature 1983, 301:379-384.
- [25]Wilmers CC, Crabtree RL, Smith DW, Murphy KM, Getz WM: Trophic facilitation by introduced top predators: grey wolf subsidies to scavengers in Yellowstone National Park. J Anim Ecol 2003, 72:909-916.
- [26]Kruuk H: Functional aspects of social hunting by carnivores. In Function and evolution in behaviour. Edited by Baerends G, Beer C, Manning A. Oxford University Press, Oxford; 1975:119-141.
- [27]Gittleman JL (1989) Carnivore group living: comparative trends. In: Gittleman JL (ed) Carnivore behavior, ecology, and evolution. Cornell University Press, Ithaca, NY, pp 183–207
- [28]Creel S, Creel NM: Communal hunting and pack size in African wild dogs, Lycaon pictus. Anim Behav 1995, 50:1325-1339.
- [29]Mech LD, Peterson R: Wolf–prey relations. In Wolves: behavior, ecology, and conservation: behavior, ecology, and conservation. Edited by Mech LD, Boitani L. University of Chicago Press, Chicago; 2003:131-160.
- [30]Palomares F, Caro TM: Interspecific killing among mammalian carnivores. Am Nat 1999, 153:492-508.
- [31]Murphy KM, Felzien GS, Hornocker MG, Ruth TK: Encounter competition between bears and cougars: some ecological implications. Ursus 1998, 10:55-60.
- [32]Schoener TW: Field experiments on interspecific competition. Am Nat 1983, 122:240-285.
- [33]Townsend CR, Begon M, Harper JL: Essentials of ecology. Wiley-Blackwell Publishing, Malden; 2008.
- [34]Wilson EA: On the whales, seals and birds of Ross Sea and South Victoria Land. In The voyage of the ‘discovery’. Edited by Scott R. Charles Scribner’s Sons, New York; 1905:469-494.
- [35]Southwell C, Paxton CGM, Borchers D, Boveng P, Rogers T, de la Mare WK: Uncommon or cryptic? Challenges in estimating leopard seal abundance by conventional but state-of-the-art methods. Deep Sea Res Part I 2008, 55:519-531.
- [36]Gilbert J, Erickson A (1977) Distribution and abundance of seals in the pack ice of the Pacific sector of the Southern Ocean. In: Llano A (ed) Adaptations within Antarctic ecosystems. Smithsonian Institution, Washington, DC, pp 703–740
- [37]Rogers TL, Bryden MM: Density and haul-out behavior of leopard seals (Hydrurga leptonyx) in Prydz Bay, Antarctica. Mar Mamm Sci 1997, 13:293-302.
- [38]Rogers TL, Hogg CJ, Irvine A: Spatial movement of adult leopard seals (Hydrurga leptonyx) in Prydz Bay, Eastern Antarctica. Polar Biol 2005, 28:456-463.
- [39]Bester MN, Ferguson JWH, Jonker FC: Population densities of pack ice seals in the Lazarev Sea, Antarctica. Antarct Sci 2002, 14:123-127.
- [40]Meredith MP, King JC: Rapid climate change in the ocean west of the Antarctic Peninsula during the second half of the 20th century. Geophys Res Lett 2005, 32:L19604.
- [41]Vaughan D, Marshall G, Connolley W, Parkinson C, Mulvaney R, Hodgson D, et al.: Recent rapid regional climate warming on the Antarctic Peninsula. Clim Change 2003, 60:243-274.
- [42]Vaughan D: Recent trends in melting conditions on the Antarctic peninsula and their implications for ice-sheet mass balance and sea level. Arct Antarct Alp Res 2006, 38:147-152.
- [43]Forcada J, Trathan PN, Boveng PL, Boyd IL, Burns JM, Costa DP, et al.: Responses of Antarctic pack-ice seals to environmental change and increasing krill fishing. Biol Conserv 2012, 149:40-50.
- [44]Marshall G (1990) A video-collar to study aquatic fauna: a view from the animal’s back. In: Reed D (ed) Spirit of enterprise: the 1990 rolex awards. Rolex, Geneva, Switzerland, pp 57–59
- [45]Parrish FA, Marshall GJ, Littnan CL, Heithaus MR, Canja S, Becker B, et al.: Foraging of juvenile monk seals at French Frigate Shoals, Hawaii. Mar Mamm Sci 2005, 21:93-107.
- [46]Parrish FA, Craig MP, Ragen TJ, Marshall GJ, Buhleier BM: Identifying diurnal foraging habitat of endangered Hawaiian monk seals using a seal-mounted video camera. Mar Mamm Sci 2000, 16:392-412.
- [47]Parrish FA, Abernathy K, Marshall GJ, Buhleier BM: Hawaiian monk seals (Monachus schauinslandi) foraging in deep-water coral beds. Mar Mamm Sci 2002, 18:244-258.
- [48]Bowen W, Tully D, Boness D, Bulheier B, Marshall GJ: Prey-dependent foraging tactics and prey profitability in a marine mammal. Mar Ecol Prog Ser 2002, 244:235-245.
- [49]Heaslip SG, Bowen WD, Iverson SJ: Testing predictions of optimal diving theory using animal-borne video from harbour seals (Phoca vitulina concolor). Can J Zool 2014, 92:309-318.
- [50]Davis R, Fuiman L, Williams T, Collier S, Hagey W, Kanatous S, et al.: Hunting behavior of a marine mammal beneath the Antarctic fast ice. Science 1999, 283:993.
- [51]Parrish FA, Marshall GJ, Buhleier B, Antonelis GA: Foraging interaction between monk seals and large predatory fish in the Northwestern Hawaiian Islands. Endanger Species Res 2008, 4:299.
- [52]Worton BJ: Kernel methods for estimating the utilization distribution in home-range studies. Ecology 1989, 70:164-168.
- [53]Austin D, Bowen WD, McMillan JI: Intraspecific variation in movement patterns: modeling individual behaviour in a large marine predator. Oikos 2004, 105:15-30.
- [54]McClintock BT, King R, Thomas L, Matthiopoulos J, McConnell BJ, Morales JM: A general discrete-time modeling framework for animal movement using multistate random walks. Ecol Monogr 2012, 82:335-349.
- [55]Freitas C, Kovacs K, Ims R, Fedak M, Lydersen C: Ringed seal post-moulting movement tactics and habitat selection. Oecologia 2008, 155:193-204.
- [56]Lyons A, Turner W, Getz W: Home range plus: a space–time characterization of movement over real landscapes. Mov Ecol 2013, 1:1-14. BioMed Central Full Text
- [57]Rutz C, Hays GC: New frontiers in biologging science. Biol Lett 2009, 5:289-292.
- [58]Costa DP, Robinson PW, Arnould JPY, Harrison A-L, Simmons SE, Hassrick JL, et al.: Accuracy of ARGOS locations of Pinnipeds at-sea estimated using Fastloc GPS. PLoS One 2010, 5:e8677.
- [59]Dujon AM, Lindstrom RT, Hays GC: The accuracy of Fastloc-GPS locations and implications for animal tracking. Methods Ecol Evol 2014, 5:1162-1169.
- [60]Tomkiewicz SM, Fuller MR, Kie JG, Bates KK: Global positioning system and associated technologies in animal behaviour and ecological research. Philos Trans R Soc B Biol Sci 2010, 365:2163-2176.
- [61]Kie JG, Matthiopoulos J, Fieberg J, Powell RA, Cagnacci F, Mitchell MS, et al.: The home-range concept: are traditional estimators still relevant with modern telemetry technology? Philos Trans R Soc B Biol Sci 2010, 365:2221-2231.
- [62]Getz WM, Wilmers CC: A local nearest-neighbor convex-hull construction of home ranges and utilization distributions. Ecography 2004, 27:489-505.
- [63]Dayton PK, Mordida BJ, Bacon F: Polar marine communities. Am Zool 1994, 34:90-99.
- [64]Orsi AH, Whitworth T III, Nowlin WD Jr: On the meridional extent and fronts of the Antarctic circumpolar current. Deep Sea Res Part I 1995, 42:641-673.
- [65]Atkinson A, Siegel V, Pakhomov EA, Jessopp MJ, Loeb V: A re-appraisal of the total biomass and annual production of Antarctic krill. Deep Sea Res Part I 2009, 56:727-740.
- [66]Agnew DJ: Review—the CCAMLR ecosystem monitoring programme. Antarct Sci 1997, 9:235-242.
- [67]Goebel ME, Pussini N, Buchheit R, Pietrzak KW, Krause DJ, Van Cise AM et al (2011) AMLR 2010/11 field season report. Chapter 8: Pinniped Research at Cape Shirreff, Livingston Island, Antarctica. US Department of Commerce, NOAA Technical Memorandum NMFS-SWFSC-524, pp 50–57
- [68]Rogers TL, Ciaglia MB, Klinck H, Southwell C: Density can be misleading for low-density species: benefits of passive acoustic monitoring. PLoS One 2013, 8:e52542.
- [69]Erickson AW, Hofman RJ (1974) Antarctic seals. Antarctic Map Folio Series. Folio 18:4–13
- [70]Marshall G, Bakhtiari M, Shepard M: An advanced solid-state animal-borne video and environmental data-logging device (“Crittercam”) for marine research. Mar Technol Soc J 2007, 41:31-38.
- [71]Pussini N, Goebel ME (2015) A safter protocol for field immobilization of leopard seals (Hydrurga leptonyx). Mar Mamm Sci. doi:10.1111/mms.12232
- [72]Luque SP: Diving behaviour analysis in R. R News 2007, 7:8-14.
- [73]Kruuk H: The spotted hyena: a study of predation and social behavior. University of Chicago Press, Chicago; 1972.
- [74]Schaller GB: The Serengeti lion: a study of predator–prey relations. University of Chicago Press, Chicago; 1972.
- [75]Taylor M (1989) Locomotor adaptations by carnivores. In: Gittleman J (ed) Carnivore behavior, ecology, and evolution. Cornell University Press, Ithaca, NY, pp 382–409
- [76]Van Orsdol KG: Foraging behaviour and hunting success of lions in Queen Elizabeth National Park, Uganda. Afr J Ecol 1984, 22:79-99.
- [77]Caro TM, Fitzgibbon CD: Large carnivores and their prey: the quick and the dead. In Natural enemies: the population biology of predators, parasites and diseases. Edited by Crawley MJ. Blackwell Scientific Publications, Oxford; 2009:115-142.
- [78]Hilborn A, Pettorelli N, Orme CDL, Durant SM: Stalk and chase: how hunt stages affect hunting success in Serengeti cheetah. Anim Behav 2012, 84:701-706.
- [79]Fishes of the southern ocean. 1st edition. JLB Smith Institute of Ichthyology, Grahamstown; 1990.
- [80]R-Core-Team (2014) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria
- [81]Freitas C (2012) argosfilter: Argos locations filter. R Foundation for Statistical Computing, Vienna, Austria
- [82]Lyons AJ, Getz WM (2014) T-LoCoH: time local convex hull homerange and time use analysis. R Foundation for Statistical Computing, Vienna, Austria
- [83]Lyons AJ (2014) T-LoCoH for R: tutorial and users guide, pp 2–53
- [84]Getz WM, Fortmann-Roe S, Cross PC, Lyons AJ, Ryan SJ, Wilmers CC: LoCoH: nonparameteric kernel methods for constructing home ranges and utilization distributions. PLoS One 2007, 2:e207.
- [85]Wickham H: ggplot2: elegant graphics for data analysis. Springer, New York; 2009.
- [86]Murray DL, Boutin S, O’Donoghue M, Nams VO: Hunting behaviour of a sympatric felid and canid in relation to vegetative cover. Anim Behav 1995, 50:1203-1210.
- [87]Stirling I: Midsummer observations on the behavior of wild polar bears (Ursus maritimus). Can J Zool 1974, 52:1191-1198.
- [88]Scantlebury DM, Mills MGL, Wilson RP, Wilson JW, Mills MEJ, Durant SM, et al.: Flexible energetics of cheetah hunting strategies provide resistance against kleptoparasitism. Science 2014, 346:79-81.
- [89]Estes JA (1989) Adaptations for aquatic living by carnivores. In: Gittleman JL (ed) Carnivore behavior, ecology, and evolution. Cornell University Press, Ithaca, NY, pp 242–282
- [90]Mader TR (1998) Temporal variation in leopard seal presence and predation near an antarctic penguin rookery. Masters Thesis Montana State University, Biology
- [91]DeLaca T, Lipps J, Zumwalt G: Encounters with Leopard seals (Hydrurga leptonyx) along the Antarctic Peninsula. Antarct J Unit States 1975, 10:85-91.
- [92]Rogers T (2009) The leopard seal, Hydrurga leptonyx. In: Perrin WF, Wursig B, Thewissen JGM (eds) Encyclopedia of marine mammals, 2nd edn. Academic Press, San Diego, pp 673–674
- [93]Kooyman GL: Leopard seal (Hydrurga leptonyx Blainville, 1820). In Handbook of marine mammals. Edited by Ridgway S, Harrison R. Academic Press, London; 1981:261-272.
- [94]Ray C: Snooping on seals for science. Anim Kingd 1966, 69:66-75.
- [95]Hocking D, Evans A, Fitzgerald EG: Leopard seals (Hydrurga leptonyx) use suction and filter feeding when hunting small prey underwater. Polar Biol 2013, 36:211-222.
- [96]Pitman RL, Durban JW: Cooperative hunting behavior, prey selectivity and prey handling by pack ice killer whales (Orcinus orca), type B, in Antarctic Peninsula waters. Mar Mamm Sci 2012, 28:16-36.
- [97]Pitman RL, Totterdell JA, Fearnbach H, Ballance LT, Durban JW, Kemps H: Whale killers: prevalence and ecological implications of killer whale predation on humpback whale calves off Western Australia. Mar Mamm Sci 2015, 31(2):629-657.
- [98]Estes JA, Riedman ML, Staedler MM, Tinker MT, Lyon BE: Individual variation in prey selection by sea otters: patterns, causes and implications. J Anim Ecol 2003, 72:144-155.
- [99]Staniland IJ, Reid K, Boyd IL: Comparing individual and spatial influences on foraging behaviour in Antarctic fur seals Arctocephalus gazella. Mar Ecol Prog Ser 2004, 275:263-274.
- [100]Weise MJ, Harvey JT, Costa DP: The role of body size in individual-based foraging strategies of a top marine predator. Ecology 2010, 91:1004-1015.
- [101]Brockmann HJ, Barnard CJ (1979) Kleptoparasitism in birds. Anim Behav 27(Part 2):487–514
- [102]Creel S, Creel N: Limitation of African wild dogs by competition with larger carnivores. Conserv Biol 1996, 10:526-538.
- [103]Gorman ML, Mills MG, Raath JP, Speakman JR: High hunting costs make African wild dogs vulnerable to kleptoparasitism by hyaenas. Nature 1998, 391:479-481.
- [104]Krofel M, Kos I, Jerina K: The noble cats and the big bad scavengers: effects of dominant scavengers on solitary predators. Behav Ecol Sociobiol 2012, 66:1297-1304.
- [105]Iyengar EV: Kleptoparasitic interactions throughout the animal kingdom and a re-evaluation, based on participant mobility, of the conditions promoting the evolution of kleptoparasitism. Biol J Linn Soc 2008, 93:745-762.
- [106]Riedman ML, Estes JA: A review of the history, distribution and foraging ecology of sea otters. In The community ecology of sea otters. Edited by VanBlaricom GR, Estes JA. Springer, Berlin; 1988:4-21.
- [107]Creel S, Spong G, Creel N (2001) Interspecific competition and the population biology of extinction-prone carnivores. In: Gittleman JL, Funk SM, Macdonald DW, Wayne RK (eds) Carnivore conservation. Cambridge University Press, London, pp 35–60
- [108]Krofel M, Kos I: Modeling potential effects of brown bear kleptoparasitism on the predation rate of Eurasian lynx. Acta Biologica Slovenica 2010, 53:47-54.
- [109]Kruuk H: Surplus killing by carnivores. J Zool 1972, 166:233-244.
- [110]Vander Wall SB: Food hoarding in animals. University of Chicago Press, Chicago; 1990.
- [111]Andersson M, Krebs J (1978) On the evolution of hoarding behaviour. Anim Behav 26(Part 3):707–711
- [112]Smith CC, Reichman OJ: The evolution of food caching by birds and mammals. Annu Rev Ecol Syst 1984, 15:329-351.
- [113]Barrett-Lennard LG, Matkin CO, Durban JW, Saulitis EL, Ellifrit D: Predation on gray whales and prolonged feeding on submerged carcasses by transient killer whales at Unimak Island, Alaska. Mar Ecol Prog Ser 2011, 421:229-241.
- [114]Kim SL, Conlan K, Malone DP, Lewis CV: Possible food caching and defence in the Weddell seal: observations from McMurdo Sound, Antarctica. Antarct Sci 2005, 17:71-72.
- [115]Ponganis PJ, Stockard TK: Short note: the Antarctic toothfish: how common a prey for Weddell seals? Antarct Sci 2007, 19:441-442.
- [116]Wilson EE, Wolkovich EM: Scavenging: how carnivores and carrion structure communities. Trends Ecol Evol 2011, 26:129-135.
- [117]Macdonald DW: Food caching by red foxes and some other carnivores. Zeitschrift für Tierpsychologie 1976, 42:170-185.
- [118]Vander Wall SB, Jenkins SH: Reciprocal pilferage and the evolution of food-hoarding behavior. Behav Ecol 2003, 14:656-667.
- [119]Hamilton JE (1939) The leopard seal Hydrurga leptonyx (De Blainville). Discovery reports 18:239–264
- [120]DeVault TL, Rhodes JOE, Shivik JA: Scavenging by vertebrates: behavioral, ecological, and evolutionary perspectives on an important energy transfer pathway in terrestrial ecosystems. Oikos 2003, 102:225-234.
- [121]Borsa P: Seasonal occurrence of the leopard seal, Hydrurga leptonyx, in the Kerguelen Islands. Can J Zool 1990, 68:405-408.