Cain, M. L., Milligan, B. G. & Strand, A. E. Long-distance seed dispersal in plant populations. Am. J. Bot. 87, 1217–1227 (2000).
Cousens, R., Dytham, C. & Law, R. Dispersal in Plants: A Population Perspective 1st edn. (Oxford University Press, Oxford, 2008).
Jordano, P. Fruits and frugivory. In Seeds: The Ecology of Regeneration in Plant Communities 2nd edn (ed. Fenner, M.) 125–166 (UK CAB International, Wallingford, 2000).
Jordano, P., García, C., Godoy, J. A. & García-Castaño, J. L. Differential contribution of frugivores to complex seed dispersal patterns. PNAS 104, 3278–3282 (2007).
Bueno, R. S. et al. Functional redundancy and complementarities of seed dispersal by the last neotropical megafrugivores. PLoS ONE 8, 0056252 (2013).
Pérez-Méndez, N., Jordano, P., García, C. & Valido, A. The signatures of Anthropocene defaunation: cascading effects of the seed dispersal collapse. Sci. Rep. 6, 24820 (2016).
Hamrick, J. L., Murawski, D. A. & Nason, J. D. The influence of seed dispersal mechanisms on the genetic structure of tropical tree populations. Vegetatio 107, 281–297 (1993).
Mueller, T., Lenz, J., Caprano, T., Fiedler, W. & Böhning-Gaese, K. Large frugivorous birds facilitate functional connectivity of fragmented landscapes. J. Appl. Ecol. 51, 684–692 (2014).
Pérez-Méndez, N., Jordano, P. & Valido, A. Persisting in defaunated landscapes: reduced plant population connectivity after seed dispersal collapse. J. Ecol. 106, 936–947 (2018).
Schupp, E. W. Quantity, quality and the effectiveness of seed dispersal by animals. Vegetatio 107, 15–29 (1993).
Schupp, E. W., Jordano, P. & Gómez, J. M. Seed dispersal effectiveness revisited: a conceptual review. New Phytol. 188, 333–353 (2010).
Traveset, A. & Richardson, D. M. Mutualistic interactions and biological invasions. Annu. Rev. Ecol. Evol. Syst. 45, 89–113 (2014).
Herrera, C. M. Seed dispersal by vertebrates. In Plant—animal interactions, an evolutionary approach (eds Herrera, C. & Pellmyr, O.) 185–209 (Wiley, Oxford, 2002).
Vidal, M. M., Pires, M. M. & Guimarães, J. P. R. Large vertebrates as the missing components of seed-dispersal networks. Biol. Conserv. 163, 42–48 (2013).
Moleón, M. et al. Rethinking megafauna. Proc. R. Soc. B 287, 20192643 (2020).
Pires, M. M., Guimarães, P. R., Galetti, M. & Jordano, P. Pleistocene megafaunal extinctions and the functional loss of long-distance seed-dispersal services. Ecography 41, 153–163 (2018).
Chen, S. C. & Moles, A. T. A mammoth mouthful? A test of the idea that larger animals ingest larger seeds. Glob. Ecol. Biogeogr. 24, 1269–1280 (2015).
Dirzo, R. et al. Defaunation of the anthropocene. Science 345, 401–406 (2014).
Galetti, M. et al. Functional extinction of birds drives rapid evolutionary changes in seed size. Science 340, 1086–1090 (2013).
Pasitschniak-Arts, M. Ursus arctos. Mamm. Species 439, 1–10 (1993).
Steyaert, S. M. J. G., Endrestøl, A., Hacklaender, K., Swenson, J. E. & Zedrosser, A. The mating system of the brown bear Ursus arctos. Mamm. Rev. 42, 12–34 (2012).
Bojarska, K. & Selva, N. Spatial patterns in brown bears Ursus arctos diet: the role of geographical and environmental factors. Mamm. Rev. 42, 120–143 (2012).
Blanchard, B. N. Size and growth patterns of the Yellowstone grizzly bear. Bears Their Biol. Manag. 7, 99–107 (1987).
Palomero, G., Fernández-Gil, A. & Naves, J. Reproductive rates of brown bears in the Cantabrian Mountains, Spain. Bears Their Biol. Manag. 9, 129–132 (1997).
Welch, C. A., Keay, J., Kendall, K. C. & Robbins, C. T. Constraints on frugivory by bears. Ecology 78, 1105–1119 (1997).
Hilderbrand, G. V. et al. The importance of meat, particularly salmon, to body size, population productivity, and conservation of North American brown bears. Can. J. Zool. 77, 132–138 (1999).
McLoughlin, P. D., Ferguson, S. H. & Messier, F. Intraspecific variation in home range overlap with habitat quality: a comparison among brown bear populations. Evol. Ecol. 14, 39–60 (2000).
Nomura, F. & Higashi, S. Effects of food distribution on the habitat usage of a female brown bear Ursus arctos yesoensis in a beech-forest zone of northernmost Japan. Ecol. Res. 15, 209–217 (2000).
Hertel, A. G. et al. Berry production drives bottom-up effects on body mass and reproductive success in an omnivore. Oikos 127, 197–207 (2017).
Zalewski, A. Geographical and seasonal variation in food habits and prey size of European pine martens. In Gilbert Martens and Fishers (Martes) in Human-Altered Environments (eds Harrison, D. J. & Fuller, A. K. P.) 77–98 (Springer, Boston, 2005).
Soe, E. et al. Europe-wide biogeographical patterns in the diet of an ecologically and epidemiologically important mesopredator, the red fox Vulpes vulpes: a quantitative review. Mamm. Rev. 47, 198–211 (2017).
Jaroszewicz, B., Pirożnikow, E. & Sondej, I. Endozoochory by the guild of ungulates in Europe’s primeval forest. Forest Ecol. Manag. 305, 21–28 (2013).
Lundgren, E. J., Ramp, D., Ripple, W. J. & Wallach, A. D. Introduced megafauna are rewilding the Anthropocene. Ecography 41, 857–866 (2018).
Kowalczyk, R. et al. Foraging plasticity allows a large herbivore to persist in a sheltering forest habitat: DNA metabarcoding diet analysis of the European bison. Forest Ecol. Manag. 449, 117474 (2019).
Gebert, C. & Verheyden-Tixier, H. Variation of diet composition of red deer (Cervus elaphus L.) in Europe. Mamm. Rev. 31, 189–201 (2008).
Cosyns, E., Delporte, A., Lens, L. & Hoffmann, M. Germination success of temperate grassland species after gut passage through ungulate and rabbit guts. J. Ecol. 93, 353–361 (2005).
Albrecht, J. et al. Humans and climate change drove the Holocene decline of the brown bear. Sci. Rep. 7, 1–11 (2017).
Hertel, A. G. et al. Bears and berries: species-specific selective foraging on a patchily distributed food resource in a human-altered landscape. Behav. Ecol. Sociobiol. 70, 831–842 (2016).
Valido, A., Schaefer, H. M. & Jordano, P. Colour, design and reward: phenotypic integration of fleshy fruit displays. J. Evol. Biol. 24, 751–760 (2011).
MacHutchon, A. G. & Wellwood, D. W. Grizzly bear food habits in the northern Yukon, Canada. Ursus 14, 225–235 (2003).
Sato, Y., Mano, T. & Takatsuki, S. Stomach contents of brown bears Ursus arctos in Hokkaido, Japan. Wildl. Biol. 11, 133–144 (2005).
Lalleroni, A., Quenette, P.-Y., Daufresne, T., Pellerin, M. & Baltzinger, C. Exploring the potential of brown bear (Ursus arctos) as a long-distance seed disperser: a pilot study in South-Western Europe. Mammalia 81, 1–9 (2017).
Baldwin, R. A. & Bender, L. C. Foods and nutritional components of diets of black bear in Rocky Mountain National Park, Colorado. Can. J. Zool. 87, 1000–1008 (2009).
Koike, S. Long-term trends in food habits of Asiatic black bears in the Misaka Mountains on the Pacific coast of central Japan. Mamm. Biol. 75, 17–28 (2010).
Campos-Arceiz, A. & Blake, S. Megagardeners of the forest—the role of elephants in seed dispersal. Acta Oecol. 37, 542–553 (2011).
Willson, M. F. & Gende, S. M. Seed dispersal by brown bears, Ursus arctos, in southeastern Alaska. Can. Field-Nat. 118, 499–503 (2004).
Naoe, S. et al. Mountain-climbing bears protect cherry species from global warming through vertical seed dispersal. Curr. Biol. 26, 315–316 (2016).
Naoe, S. et al. Downhill seed dispersal by temperate mammals: a potential threat to plant escape from global warming. Sci. Rep. 9, 1–11 (2019).
McConkey, K. R. & O’Farrill, G. Loss of seed dispersal before the loss of seed dispersers. Biol. Conserv. 201, 38–49 (2016).
Skuban, M., Finďo, S. & Kajba, M. Human impacts on bear feeding habits and habitat selection in the Poľana Mountains, Slovakia. Eur. J. Wildl. Res. 62, 353–364 (2016).
Štofík, J., Merganič, J., Merganičová, K., Bučko, J. & Saniga, M. Brown bear winter feeding ecology in the area with supplementary feeding—Eastern Carpathians (Slovakia). Pol. J. Ecol. 64, 277–288 (2016).
Selva, N. et al. Supplementary ungulate feeding affects movement behavior of brown bears. Basic Appl. Ecol. 24, 68–76 (2017).
López-Bao, J. V. & González-Varo, J. P. Frugivory and spatial patterns of seed deposition by carnivorous mammals in anthropogenic landscapes: a multi-scale approach. PLoS ONE 6, e14569 (2011).
Traveset, A. & Willson, M. F. Effect of birds and bears on seed germination of fleshy-fruited plants in temperate rainforests of southeast Alaska. Oikos 80, 89–95 (1997).
Nowak, J. & Crone, E. E. It is good to be eaten by a bear: effects of ingestion on seed germination. Am. Midl. Nat. 167, 205–209 (2012).
Steyaert, S. M. J. G., Hertel, A. G. & Swenson, J. E. Endozoochory by brown bears stimulates germination in bilberry. Wildl. Biol. 2019, wlb.00573 (2019).
Samuels, I. A. & Levey, D. J. Effects of gut passage on seed germination: do experiments answer the questions they ask?. Funct. Ecol. 19, 365–368 (2005).
Valido, A. & Olesen, J. M. The importance of lizards as frugivores and seed dispersers. In Seed Dispersal: Theory and its Application in a Changing World (eds Dennis, A. J. et al.) 124–147 (CAB International, Wallingford, 2007).
Traveset, A. Effect of seed passage through vertebrate frugivores’ guts on germination: a review. Perspect. Plant. Ecol. Syst. 1, 151–190 (1998).
Eriksson, O. & Fröborg, H. “Windows of opportunity” for recruitment in long-lived clonal plants: experimental studies of seedling establishment in Vaccinium shrubs. Can J. Bot. 74, 1369–1374 (1996).
Jansen, P. A. et al. Thieving rodents as substitute dispersers of megafaunal seeds. PNAS 109, 12610–12615 (2012).
Koike, S. et al. Seed removal and survival in Asiatic black bears Ursus thibetanus scats: effect of rodents as secondary seed dispersers. Wildlife Biol. 18, 24–34 (2012).
Bartoń, K. A., Zwijacz-Kozica, T., Zięba, F., Sergiel, A. & Selva, N. Bears without borders: long-distance movement in human-dominated landscapes. Glob. Ecol. Conserv. 17, e00541 (2019).
Willson, M. F. & Traveset, A. The ecology of seed dispersal. In Seeds: The Ecology of Regeneration in Plant Communities 2nd edn (ed. Fenner, M.) 85–111 (CAB International, Wallingford, 2000).
Elfström, M., Støen, O.-G., Zedrosser, A., Warrington, I. & Swenson, J. E. Gut retention times in captive brown bears Ursus arctos. Wildl. Biol. 19, 317–324 (2013).
Koike, S. et al. Estimate of the seed shadow created by the Asiatic black bear Ursus thibetanus and its characteristics as a seed disperser in Japanese cool-temperate forest. Oikos 120, 280–290 (2010).
Hickey, J. R., Flynn, R. W., Buskirk, S. W., Gerow, K. G. & Willson, M. F. An evaluation of a mammalian predator, Martes americana, as a disperser of seeds. Oikos 87, 499–508 (1999).
Terakawa, M., Isagi, Y., Matsui, K. & Yumoto, T. Microsatellite analysis of the maternal origin of Myrica rubra seeds in the feces of Japanese macaques. Ecol. Res. 24, 663–670 (2009).
González-Varo, J. P., López-Bao, J. V. & Guitián, J. Functional diversity among seed dispersal kernels generated by carnivorous mammals. J. Anim. Ecol. 82, 562–571 (2013).
Tsuji, Y., Okumura, T., Kitahara, M. & Jiang, Z. Estimated seed shadow generated by Japanese martens (Martes melampus): comparison with forest-dwelling animals in Japan. Zool. Sci. 33, 352–357 (2016).
Santini, L. et al. Ecological correlates of dispersal distance in terrestrial mammals. Hystrix 24, 181–186 (2013).
Bunney, K., Bond, W. J. & Henley, M. Seed dispersal kernel of the largest surviving megaherbivore—the African savanna elephant. Biotropica 49, 395–401 (2017).
Galetti, et al. Ecological and evolutionary legacy of megafauna extinctions. Biol. Rev. 93, 845–862 (2018).
Olson, D. M. et al. Terrestrial ecoregions of the world: a new map of life on earth: a new global map of terrestrial ecoregions provides an innovative tool for conserving biodiversity. Bioscience 51, 933–938 (2001).
Nin, S., Petrucci, W. A., Del Bubba, M., Ancillotti, C. & Giordani, E. Effects of environmental factors on seed germination and seedling establishment in bilberry (Vaccinium myrtillus L.). Sci. Hortic. 226, 241–249 (2017).
Bates, D., Maechler, M., Bolker, B. & Walker, S. Fitting linear mixed-effects models using lme4. J. Stat. Softw. 67, 1–48 (2015).
Oksanen, J. et al. Vegan package: community ecology package. R package version 2.5–6 (2019).
Silva, L. J. D. & Medeiros, A. D. D. SeedCalc, a new automated R software tool for germination and seedling length data processing. J. Seed. Sci. 41, 250–257 (2019).
R Development Core Team. R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. https://www.R-project.org/ (2017).
South, A. rworldmap: a new R package for mapping global data. R J. 3, 35–43 (2011).
IUCN SSC Bear Specialist Group. Ursus arctos. The IUCN Red List of Threatened Species. Version 2017-3 (2017). http://www.iucnredlist.org (Downloaded in May 2020).
Source: Ecology - nature.com
