Most studies focused on species coexistence have been directed at the differential use of habitat and food resources; nonetheless, the differential use of the diel cycle may enhance the coexistence of same-sized species. We investigated the activity patterns of mesocarnivores (red fox (Vulpes vulpes), European badger (Meles meles), pine marten (Martes martes), stone marten (M. foina)) in NW Italy via camera-trapping. We hypothesized that the smallest species would tend to avoid competition by selecting time periods when larger species were less active. Foxes, badgers, and stone martens were mainly nocturnal. In lowland areas overlap between coexisting species was generally low, while in Mediterranean habitats all activity patterns tended to be unimodal and overlap was generally high. The pine marten showed a cathemeral pattern. We suggest that the lower ability of the stone marten to avoid interference competition at community-level may play a major role in determining its widespread exclusion from forested areas by the pine marten.
Purchase
Buy instant access (PDF download and unlimited online access):
Institutional Login
Log in with Open Athens, Shibboleth, or your institutional credentials
Personal login
Log in with your brill.com account
Abramov A.V., Kranz A., Herrero J., Choudhury A., Maran T. (2016). Martes foina. â The IUCN Red List of Threatened Species 2016: e.T29672A45202514. Available online at http://dx.doi.org/10.2305/IUCN.UK.2016-1.RLTS.T29672A45202514.en (accessed 15 August 2016).
Arias-Del Razo I., Hernández L., Laundré J.W., Myers O. (2011). Do predator and prey foraging activity patterns match? A study of coyotes (Canis latrans), and lagomorphs (Lepus californicus and Sylvilagus audobonii). â J. Arid Environ. 75: 112-118.
Artois M. (1989). Encyclopedie des Carnivores de France, Vol. 3. Le renard roux. â Societé Francaise pour lâEtude et la Protection des Mammiferes, Bourges.
Asteraki E.J., Hart B.J., Ings T.C., Manley W.J. (2004). Factors influencing the plant and invertebrate diversity of arable field margins. â Agric. Ecosyst. Environ. 102: 219-231.
Balestrieri A. (1993). Aree vitali e preferenze ecologiche della volpe (Vulpes vulpes) in due zone delle province di Pavia ed Alessandria. â Dissertation, Università degli Studi di Milano, Milan.
Balestrieri A., Remonti L., Prigioni C. (2004). Diet of the Eurasian badger (Meles meles) in an agricultural riverine habitat (NW Italy). â Hystrix Ital. J. Mamm. 15: 3-12.
Balestrieri A., Remonti L., Prigioni C. (2009). Exploitation of food resources by the Eurasian badger (Meles meles) at the altitudinal limit of its Alpine range (NW Italy). â Zool. Sci. 26: 821-827.
Balestrieri A., Remonti L., Capra R.B., Canova L., Prigioni C. (2013). Food habits of the stone marten (Martes foina) (Mammalia: Carnivora) in plain areas of Northern Italy prior to pine marten (M. martes) spreading. â Ital. J. Zool. 80: 60-68.
Balestrieri A., Remonti L., Ruiz-González A., Zenato M., Gazzola A., Vergara M., Dettori E.E., Saino N., Capelli E., Gómez-Moliner B.J., Guidali F., Prigioni C. (2015). Distribution and habitat use by pine marten Martes martes in a riparian corridor crossing intensively cultivated lowlands. â Ecol. Res. 30: 153-162.
Balestrieri A., Bogliani G., Boano G., Ruiz-González A., Capelli E., Vergara M., Prigioni C., Saino N., Costa S., Milanesi P. (2016a). Modelling the distribution of forest-dependent species in human-dominated landscapes: patterns for the pine marten in intensively cultivated lowlands. â PLoS ONE 11: e0158203.
Balestrieri A., Ruiz-González A., Vergara M., Capelli E., Tirozzi P., Alfino S., Minuti G., Prigioni C., Saino N. (2016b). Pine marten density in lowland riparian woods: a test for the random encounter model. â Mamm. Biol. 81: 439-446.
Balestrieri A., Ruiz-González A., Capelli E., Vergara M., Prigioni C., Saino N. (2016c). Pine marten vs. stone marten in agricultural lowlands: a landscape-scale, genetic survey. â Mamm. Res. 61: 327-335.
Barrientos R., Virgós E. (2006). Reduction of potential food interference in two sympatric carnivores by sequential use of shared resources. â Acta Oecol. 30: 107-116.
Barrull J., Mate I., Ruiz-Olmo J., Casanovas J.G., Gosà lbez J., Salicrú M. (2014). Factors and mechanisms that explain coexistence in a Mediterranean carnivore assemblage: an integrated study based on camera trapping and diet. â Mamm. Biol. 79: 123-131.
Bennie J.J., Duffy J.P., Inger R., Gaston K.J. (2014). Biogeography of time partitioning in mammals. â Proc. Natl. Acad. Sci. USA 111: 13727-13732.
Berger K.M., Gese E.M. (2007). Does interference competition with wolves limit the distribution and abundance of coyotes? â J. Anim. Ecol. 76: 1075-1085.
Bertolino S., Dore B. (1995). Food habits of the stone marten in âLa Mandriaâ Regional Park. â Hystrix Ital. J. Mamm. 7: 105-112.
Braczkowski A.R., Balme G.A., Dickman A., Fattebert J., Johnson P., Dickerson T., Macdonald D.W., Hunter L. (2016). Scent lure effect on camera-trap based leopard density estimates. â PLoS ONE 11: e0151033.
Brashares J.S., Prugh L.R., Stoner C.J., Epps C.W. (2010). Ecological and conservation implications of mesopredator release. â In: Trophic cascades: predators, prey, and the changing dynamics of nature ( Terborgh J., Estes J.A., eds). Island Press, Washington, DC, p. 221-240.
Carothers J., Jaksic F.M. (1984). Time as a niche difference: the role of interference competition. â Oikos 42: 403-406.
Cavallini P., Lovari S. (1991). Environmental factors influencing the use of habitat in the red fox, Vulpes vulpes. â J. Zool. 223: 323-339.
Cavallini P., Lovari S. (1994). Home range, habitat selection and activity of the red fox in a Mediterranean coastal ecotone. â Acta Theriol. 39: 279-287.
Chen I.-C., Hill J.K., Ohlemüller R., Roy D.B., Thomas C.D. (2011). Rapid range shifts of species associated with high levels of climate warming. â Science 333: 1024-1026.
Clevenger A.P. (1992). Pine marten (Martes martes L.) home ranges and activity patterns on the island of Minorca, Spain. â Z. Saugetierkd. 58: 137-143.
Creel S., Creel N.M. (2002). The African wild dog: behavior, ecology, and conservation. â Princeton University Press, Princeton, NJ.
Cresswell W., Lind J., Quinn J.L. (2010). Predator hunting success and prey vulnerability: quantifying the spatial scale over which lethal and non-lethal effects of predation occur. â J. Anim. Ecol. 79: 556-562.
Dayan T., Simberloff D. (1994). Character displacement, sexual dimorphism, and morphological variation among British and Irish mustelids. â Ecology 75: 1063-1073.
de Oliveira T.G., Pereira J.A. (2014). Intraguild predation and interspecific killing as structuring forces of carnivoran communities in South America. â J. Mamm. Evol. 21: 427-436.
Del Fante S. (2012). Comportamento spaziale della martora (Martes martes) in ambiente appenninico. â Dissertation, Università degli Studi di Pavia, Pavia.
Delibes M. (1983). Interspecific competition and the habitat of the stone marten Martes foina (Erxleben, 1777) in Europe. â Acta Zool. Fenn. 174: 229-231.
Di Bitetti M.S., Di Blanco Y.E., Pereira J.A., Paviolo A., Pérez I.J. (2009). Time partitioning favors the coexistence of sympatric crab-eating foxes (Cerdocyon thous) and Pampas foxes (Lycalopex gymnocercus). â J. Mammal. 90: 479-490.
Donadio E., Buskirk S.W. (2006). Diet, morphology, and interspecific killing in Carnivora. â Am. Nat. 167: 524-536.
Elmhagen B., Tannerfeldt M., Angerbjörn A. (2002). Food-niche overlap between Arctic and red foxes. â Can. J. Zool. 80: 1274-1285.
Foster V.C., Sarmento P., Sollmann R., Tôrres N., Jácomo A.T., Negrões N., Fonseca C., Silveira L. (2013). Jaguar and puma activity patterns and predatorâprey interactions in four Brazilian biomes. â Biotropica 45: 373-379.
Galaverni M., Caniglia R., Fabbri E., Milanesi P., Randi E. (2016). One, no one, or one hundred thousand: how many wolves are there currently in Italy? â Acta Theriol. 61: 13-24.
Gehrt S.D. (2007). Ecology of coyotes in urban landscapes. â Paper 63, Wildlife Damage Management Conferences Proceedings. Available online at http://digitalcommons.unl.edu/icwdm_wdmconfproc/63.
Genovesi P. (1993). Strategie di sfruttamento delle risorse e struttura sociale della faina (Martes foina Erxleben 1777) in ambiente forestale e rurale. â PhD thesis, Università di Roma La Sapienza, Rome.
Gompper M.E., Lesmeister D.B., Ray J.C., Malcolm J.R., Kays R. (2016). Differential habitat use or intraguild interactions: what structures a carnivore community? â PLoS ONE 11: e0146055.
Halle S. (2000). Ecological relevance of daily activity patterns. â In: Activity patterns in small mammals. Ecological studies, Vol. 141 ( Halle S., Stenseth N.C., eds). Springer, Berlin, p. 67-90.
Hass C.C. (2009). Competition and coexistence in sympatric bobcats and pumas. â J. Zool. 278: 174-180.
Hastings A. (1996). Models of spatial spread: is the theory complete? â Ecology 77: 1675-1679.
Hayama H., Kaneda M., Tabata M. (2006). Rapid range expansion of the feral raccoon (Procyon lotor) in Kanagawa Prefecture, Japan, and its impact on native organisms. â In: Assessment and control of biological invasion risks ( Koike F., Clout M.N., Kawamichi M., De Poorter M., Iwatsuki K., eds). Shoukadoh Book Sellers, Kyoto and IUCN, Gland, p. 196-199.
Henden J.A., Ims R.A., Yoccoz N.G., Hellström P., Angerbjörn A. (2010). Strength of asymmetric competition between predators in food webs ruled by fluctuating prey: the case of foxes in tundra. â Oikos 119: 27-34.
Hermann M. (1994). Habitat use and spatial organisation by the stone marten. â In: The biology and conservation of martens, sables and fishers ( Buskirk S.W., Harestad A.S., Raphael M.G., Powell R.A., eds). Cornell University Press, Ithaca, NY, p. 122-136.
Herrero J., Kranz A., Skumatov D., Abramov A.V., Maran T., Monakhov V.G. (2016). Martes martes. â The IUCN Red List of Threatened Species 2016: e.T12848A45199169. Available online at http://dx.doi.org/10.2305/IUCN.UK.2016-1.RLTS.T12848A45199169.en (accessed 15 August 2016).
Hill R.A. (2006). Why be diurnal? Or, why not be cathemeral? â Folia Primatol. 77: 72-86.
Höchtl F., Lehringer S., Konold W. (2005). âWildernessâ: what it means when it becomes a reality â a case study from the southwestern Alps. â Landscape Urban Plan. 70: 85-95.
Ivlev V.S. (1961). Experimental ecology of the feeding of fishes. â Yale University Press, New Haven, CT.
Jacobs J. (1974). Quantitative measurement of food selection. â Oecologia 14: 413-417.
Kays R.W., Slauson K.M. (2008). Remote cameras. â In: Noninvasive survey methods for carnivores: methods and analyses ( Long R.A., Mackay P., Zielinski W.J., Ray J.C., eds). Island Press, Washington, DC, p. 105-134.
Kelly M.J., Holub E.L. (2008). Camera trapping of carnivores: trap success among camera types and across species, and habitat selection by species, on Salt Pond Mountain, Giles County, Virginia. â Northeast. Nat. 15: 249-262.
King D.P., Takahashi J.S. (2000). Molecular genetics of circadian rhythms in mammals. â Annu. Rev. Neurosci. 23: 713-742.
Kinlan B., Hastings A. (2005). What exotic species tell us about rates of population spread and geographic range expansion. â In: Species invasions: insights into ecology, evolution, and biogeography ( Sax D.F., Stachowicz J.J., Gaines S.D., eds). Sinauer Associates, Sunderland, MA, p. 281-419.
Ko C.H., Takahashi J.S. (2006). Molecular components of the mammalian circadian clock. â Hum. Mol. Genet. 15: R271-R277.
Kowalczyk R., Jedrzejewska B., Zalewski A. (2003). Annual and circadian activity patterns of badgers (Meles meles) in BiaÅowieza Primeval Forest (Eastern Poland) compared with other Palaearctic populations. â J. Biogeogr. 30: 463-472.
Kronfeld-Schor N., Dayan T. (2003). Partitioning of time as an ecological resource. â Annu. Rev. Ecol. Syst. 34: 153-181.
Larroque J., Ruette S., Vandel J.M., Devillard S. (2015). Where to sleep in a rural landscape? A comparative study of resting sites pattern in two syntopic Martes species. â Ecography 38: 1-12.
Levi T., Wilmers C.C. (2012). Wolvesâcoyotesâfoxes: a cascade among carnivores. â Ecology 93: 921-929.
Lindström E.R., Brainerd S.M., Helldin J.O., Overskaug K. (1995). Pine martenâred fox interactions: a case of intraguild predation. â Ann. Zool. Fenn. 32: 123-130.
Linkie M., Ridout M.S. (2011). Assessing tigerâprey interactions in Sumatran rainforests. â J. Zool. 284: 224-229.
Linnell J.D., Strand O. (2000). Interference interactions, coexistence and conservation of mammalian carnivores. â Divers. Distrib. 6: 169-176.
Lockard R.B. (1978). Seasonal change in the activity pattern of Dipodomys spectabilis. â J. Mamm. 59: 563-568.
Lovari S., Minder I., Ferretti F., Mucci N., Randi E., Pellizzi B. (2013). Common and snow leopards share prey, but not habitats: competition avoidance by large predators? â J. Zool. 291: 127-135.
Lund U., Agostinelli C., Agostinelli M.C. (2013). Package âcircularâ. Version 0.4-7. â Available online at http://cran.r-project.org/web/packages/circular/circular.pdf.
MacArthur R.H., Levins R. (1964). Competition, habitat selection and character displacement in a patchy environment. â Proc. Natl. Acad. Sci. USA 51: 1207-1210.
MacArthur R.H., Levins R. (1967). The limiting similarity, convergence, and divergence of coexisting species. â Am. Nat. 101: 377-385.
Macdonald D.W., Buesching C.D., Stopka P., Henderson J., Ellwood S.A., Baker S.E. (2004). Encounters between two sympatric carnivores: red foxes (Vulpes vulpes) and European badgers (Meles meles). â J. Zool. 263: 385-392.
Meredith M., Ridout M. (2014). Overview of the âoverlapâ package, version 0.2.6. â Available online at http://cran.r-project.org/web/packages/overlap/overlap.pdf.
Merkle J.A., Stahler D.R., Smith D.W. (2009). Interference competition between gray wolves and coyotes in Yellowstone National Park. â Can. J. Zool. 87: 56-63.
Michel N., Burel F., Butet A. (2006). How does landscape use influence small mammal diversity, abundance and biomass in hedgerow networks of farming landscapes? â Acta Oecol. 30: 11-20.
Mitchell B.D., Banks P.B. (2005). Do wild dogs exclude foxes? Evidence for competition from dietary and spatial overlaps. â Austral Ecol. 30: 581-591.
Monterroso P., Alves P.C., Ferreras P. (2014). Plasticity in circadian activity patterns of mesocarnivores in Southwestern Europe: implications for species coexistence. â Behav. Ecol. Sociobiol. 68: 1403-1417.
Monterroso P., Rebelo P., Alves P.C., Ferreras P. (2016). Niche partitioning at the edge of the range: a multidimensional analysis with sympatric martens. â J. Mammal. 97: 928-939.
Mori E., Menchetti M., Balestrieri A. (2014). Interspecific den sharing: a study on European badger setts using camera traps. â Acta Ethol. 18: 121.
Moritz C., Patton J.L., Conroy C.J., Parra J.L., White G.C., Beissinger S.R. (2008). Impact of a century of climate change on small-mammal communities in Yosemite National Park, USA. â Science 322: 261-264.
Mouches A. (1981). Eco-ethologie du blaireau europeen Meles meles L.: strategies dâutilitsation de lâhabitat et des resources alimentaires. â PhD thesis, Université de Rennes I, Rennes.
Norris D., Michalski F., Peres C.A. (2010). Habitat patch size modulates terrestrial mammal activity patterns in Amazonian forest fragments. â J. Mammal. 91: 551-560.
Palomares F., Caro T.M. (1999). Interspecific killing among mammalian carnivores. â Am. Nat. 153: 492-508.
Parmesan C. (2006). Ecological and evolutionary responses to recent climate change. â Annu. Rev. Ecol. Evol. Syst. 37: 637-669.
Pewsey A., Neuhäuser M., Ruxton G.D. (2013). Circular statistics in R. â Oxford University Press, New York, NY.
Polis G.A., Holt R.D. (1992). Intraguild predation: the dynamics of complex trophic interactions. â Trends Ecol. Evol. 7: 151-154.
Prigioni C., Balestrieri A., Remonti L., Cavada L. (2008). Differential use of food and habitat by sympatric carnivores in the eastern Italian Alps. â Ital. J. Zool. 75: 173-184.
Prigioni C., Smiroldo G., Remonti L., Balestrieri A. (2009). Distribution and diet of reintroduced otters (Lutra lutra) on the River Ticino (NW Italy). â Hystrix Ital. J. Mamm. 20: 45-53.
Prugh L.R., Stoner C.J., Epps C.W., Bean W.T., Ripple W.J., Laliberte A.S., Brashares J.S. (2009). The rise of the mesopredator. â BioScience 59: 779-791.
Pulliainen E. (1981). Winter habitat selection, home-range, and movements of the pine marten (Martes martes) in Finnish Lapland Forest. â In: Worldwide furbearer congress proceedings ( Chapman J.A., Pursley D., eds). Worldwide Furbearer Conference, Frostburg, MD, p. 1068-1087.
Remonti L., Balestrieri A., Prigioni C. (2011). Percentage of protein, lipids and carbohydrates in the diet of badger (Meles meles) populations across Europe. â Ecol. Res. 26: 487-495.
Remonti L., Balestrieri A., Ruiz-González A., Gómez-Moliner B.J., Capelli E., Prigioni C. (2012). Intraguild dietary overlap and its possible relationship to the coexistence of mesocarnivores in intensive agricultural habitats. â Popul. Ecol. 54: 521-532.
Ridout M.S., Linkie M. (2009). Estimating overlap of daily activity patterns from camera trap data. â J. Agric. Biol. Environ. Stat. 14: 322-337.
Ripple W.J., Estes J.A., Beschta R.L., Wilmers C.C., Ritchie E.G., Hebblewhite M., Berger J., Elmhagen B., Letnic M., Nelson M.P., Schmitz O.J., Smith D.W., Wallach A.D., Wirsing A.J. (2014). Status and ecological effects of the worldâs largest carnivores. â Science 343: 1241484.
Ritchie E.G., Johnson C.N. (2009). Predator interactions, mesopredator release and biodiversity conservation. â Ecol. Lett. 12: 982-998.
Robertson D.R., Sweatman H.P.A., Fletcher E.A., Cleland M.G. (1976). Schooling as a mechanism for circumventing the territoriality of competitors. â Ecology 57: 1208-1220.
Roemer G.W., Gompper M.E., Van Valkenburgh B. (2009). The ecological role of the mammalian mesocarnivore. â BioScience 59: 165-173.
Rosellini S., Osorio E., Ruiz-González A., Piñeiro A., Barja I. (2008). Monitoring the small-scale distribution of sympatric European pine martens (Martes martes) and stone martens (Martes foina): a multievidence approach using faecal DNA analysis and camera-traps. â Wildl. Res. 35: 434-440.
Rosenzweig M.L. (1987). Habitat selection as a source of biological diversity. â Evol. Ecol. 1: 312-330.
Servin J., Rau J.R., Delibes M. (1991). Activity pattern of the red fox Vulpes vulpes in Doñana, southwest Spain. â Acta Theriol. 36: 369-373.
Sheehy E., Lawton C. (2014). Population crash in an invasive species following the recovery of a native predator: the case of the American grey squirrel and the European pine marten in Ireland. â Biodivers. Conserv. 23: 753-774.
Sindaco R. (2006). Segnalazioni faunistiche piemontesi e valdostane (Amphibia, Reptilia, Mammalia). â Riv. Piem. Stat. Nat. 27: 443-459.
Smedshaug C.A., SelÃ¥s V., Lund S.E., Sonerud G.A. (1999). The effect of a natural reduction of red fox Vulpes vulpes on small game hunting bags in Norway. â Wildlife Biol. 5: 157-166.
Soulé M.E., Bolger D.T., Alberts A.C., Wright J., Sorice M., Hill S. (1988). Reconstructed dynamics of rapid extinctions of chaparral-requiring birds in urban habitat islands. â Conserv. Biol. 2: 75-91.
Spagnesi M., De Marinis A.M. (eds) (2002). Mammiferi dâItalia. Q. conserv. nat., Vol. 14, p. 1-309.
Tannerfeldt M., Elmhagen B., Angerbjörn A. (2002). Exclusion by interference competition? The relationship between red and Arctic foxes. â Oecologia 132: 213-220.
Thompson I.D. (1994). Marten population in uncut and logged boreal forests in Ontario. â J. Wildlife Manage. 58: 272-280.
Tokeshi M. (1999). Species coexistence. Ecological and evolutionary perspectives. â Blackwell, London.
Torretta E., Serafini M., Puopolo F., Schenone L. (2016). Spatial and temporal adjustments allowing the coexistence among carnivores in Liguria (NW Italy). â Acta Ethol. 19: 123-132.
Travaini A., Aldama J.J., Laffitte R., Delibes M. (1993). Home range and activity patterns of red fox Vulpes vulpes breeding females. â Acta Theriol. 38: 427-434.
Van Maanen E. (2013). Onderscheid tussen boom-en steenmarter in de hand, in het veld en op foto. â Jaarbrief WBN van de Zoogdiervereniging over 2012 in MASTERPASSEN XIX. Zoogdiervereniging, Nijmegen.
Virgós E., Zalewski A., Rosalino L.M., Mergey M. (2012). Habitat ecology of genus Martes in Europe: a review of the evidences. â In: Biology and conservation of marten, sables, and fisher: a new synthesis ( Aubry K.B., Zielinski W.J., Raphael M.G., Proulx G., Buskirk S.W., eds). Cornell University Press, Ithaca, NY, p. 255-266.
Von Holle B., Simberloff D. (2005). Ecological resistence to biological invasion overwhelmed by propagule pressure. â Ecology 86: 3212-3218.
Webster J.A. (2001). AÂ review of the historical evidence of the habitat of the pine marten in Cumbria. â Mammal Rev. 31: 17-31.
Wereszczuk A., Zalewski A. (2015). Spatial niche segregation of sympatric stone marten and pine marten. Avoidance of competition or selection of optimal habitat? â PLoS ONE 10: e0139852.
Whitehouse M.E.A. (1997). Experience influences maleâmale contests in the spider Argyrodes antipodiana (Theridiidae: Araneae). â Anim. Behav. 53: 913-923.
Wilcox B.A., Murphy D.D. (1985). Conservation strategy, the effects of fragmentation on extinction. â Am. Nat. 125: 879-887.
Wilson D.E., Mittermeier R.A. (2009). Handbook of the mammals of the world, Vol. 1 (Carnivores). â Lynx Edicions, Barcelona.
| All Time | Past 365 days | Past 30 Days | |
|---|---|---|---|
| Abstract Views | 1716 | 179 | 22 |
| Full Text Views | 314 | 15 | 1 |
| PDF Views & Downloads | 210 | 37 | 3 |
Most studies focused on species coexistence have been directed at the differential use of habitat and food resources; nonetheless, the differential use of the diel cycle may enhance the coexistence of same-sized species. We investigated the activity patterns of mesocarnivores (red fox (Vulpes vulpes), European badger (Meles meles), pine marten (Martes martes), stone marten (M. foina)) in NW Italy via camera-trapping. We hypothesized that the smallest species would tend to avoid competition by selecting time periods when larger species were less active. Foxes, badgers, and stone martens were mainly nocturnal. In lowland areas overlap between coexisting species was generally low, while in Mediterranean habitats all activity patterns tended to be unimodal and overlap was generally high. The pine marten showed a cathemeral pattern. We suggest that the lower ability of the stone marten to avoid interference competition at community-level may play a major role in determining its widespread exclusion from forested areas by the pine marten.
| All Time | Past 365 days | Past 30 Days | |
|---|---|---|---|
| Abstract Views | 1716 | 179 | 22 |
| Full Text Views | 314 | 15 | 1 |
| PDF Views & Downloads | 210 | 37 | 3 |