Summary - AU Pure - R dplyr group by string

Shallow size-density relations within mammal clades suggest greater intra-guild ecological impact of large-bodied speciesR. ?. Pedersen*a, S. Faurby a,b,c,d, J.-C. Svenning aaSection for Ecoinformatics & Biodiversity, Department of Bioscience, Aarhus University, Ny Munkegade 114, DK-8000 Aarhus C, DenmarkbDepartment of Biogeography and Global Change, Museo Nacional de Ciencias Naturales, CSIC, Calle José Gutiérrez Abascal 2, Madrid 28006, SpaincDepartment of Biological and Environmental Sciences, University of Gothenburg, Box 461, SE-405 30 G?teborg, SwedendGothenburg Global Biodiversity Centre, Box 461, SE-405 30 G?teborg, Sweden *Corresponding author: rasmus.pedersen@bios.au.dkSummary 1. Population densities of species have a predictable relationship to their body mass on a global scale. This relationship is known as the size-density relationship. Since the relationship originally was found to be directly opposite of metabolic rate scaling, it led to the hypothesis of energetic equivalence. However, recent studies have suggested that relationship of both metabolic scaling and the size-density relationship are variable between clades. Specifically, the within clade size-density relationship tends to be less negative than the overall relationship.2. The aim of the present study is to estimate phylogenetic variation in the scaling relationship, using a data-driven identification of natural phylogenetic substructure in the density-body size relation, and discuss potential drivers. The classic model is often used to estimate natural population densities, and a further, practical aim here is to improve it by incorporating variability among phylogenetic groups.3. We expand the model for the body size-population density relationship relation of mammals to include clade-specific variation. We used a dataset with population and body mass estimates of 924 terrestrial mammal species, covering 97 families, and applied an algorithm identifying group-specific changes in the relationship across a family-level phylogeny.4. We show increased performance in species density estimation is achieved by incorporating clade-specific changes in the relationship compared to the classic model (increasing r2 from 0.56 to 0.74 and ΔAICc = 466). While the global density-body mass relationship across clades was confirmed to be -0.75, as previously found, the relationship within all sub-clades was less negative than the overall trend.5. Our results show that data-driven identification of phylogenetic substructure in the density-body size relation substantially improves predictive accuracy of the model. The less negative relationship within clades compared to the overall trend and compared to within clade metabolic scaling suggest that the energetic equivalence rule does not hold. When the relationship is less negative than predicted large species within clades uses proportionally more energy than smaller species. Therefore, our results are consistent with greater intra-guild ecological impact of large-bodied species as an important determinant of population density, potentially through size-asymmetric intra-guild competition via partial monopolisation of resources by the largest species of a given guild.Key-words Allometry, body size, Cope’s rule, energetic equivalence, evolution, intra-guild competition, mammals.IntroductionPopulation densities of terrestrial mammals are negatively related to body mass, a relationship which has been found repeatedly across several phyla in the animal kingdom ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "author" : [ { "dropping-particle" : "", "family" : "Peters", "given" : "Robert Henry", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "id" : "ITEM-1", "issued" : { "date-parts" : [ [ "1983" ] ] }, "number-of-pages" : "329", "publisher" : "Cambridge University Press", "publisher-place" : "Cambridge", "title" : "The Ecological Implications of Body Size", "type" : "book" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1111/j.1095-8312.1987.tb01990.x", "ISSN" : "00244066", "author" : [ { "dropping-particle" : "", "family" : "Damuth", "given" : "John", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Biological Journal of the Linnean Society", "id" : "ITEM-2", "issue" : "3", "issued" : { "date-parts" : [ [ "1987", "7", "28" ] ] }, "page" : "193-246", "title" : "Interspecific allometry of population density in mammals and other animals: the independence of body mass and population energy-use", "type" : "article-journal", "volume" : "31" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Peters 1983; Damuth 1987)", "plainTextFormattedCitation" : "(Peters 1983; Damuth 1987)", "previouslyFormattedCitation" : "(Peters 1983; Damuth 1987)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Peters 1983; Damuth 1987). Notably, studies have found that the population density, D, of a species relates to mean body mass, M, as log D = a + b log M, with the slope b close to -0.75, independent of habitat and dietary class ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1038/290699a0", "ISSN" : "0028-0836", "author" : [ { "dropping-particle" : "", "family" : "Damuth", "given" : "John", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Nature", "id" : "ITEM-1", "issue" : "5808", "issued" : { "date-parts" : [ [ "1981", "4", "23" ] ] }, "page" : "699-700", "title" : "Population density and body size in mammals", "type" : "article-journal", "volume" : "290" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1111/j.1095-8312.1987.tb01990.x", "ISSN" : "00244066", "author" : [ { "dropping-particle" : "", "family" : "Damuth", "given" : "John", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Biological Journal of the Linnean Society", "id" : "ITEM-2", "issue" : "3", "issued" : { "date-parts" : [ [ "1987", "7", "28" ] ] }, "page" : "193-246", "title" : "Interspecific allometry of population density in mammals and other animals: the independence of body mass and population energy-use", "type" : "article-journal", "volume" : "31" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Damuth 1981, 1987)", "plainTextFormattedCitation" : "(Damuth 1981, 1987)", "previouslyFormattedCitation" : "(Damuth 1981, 1987)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Damuth 1981, 1987). The relationship is noisy though and densities vary about two orders of magnitude to either side of the overall trend ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1086/513495", "ISBN" : "0003-0147/2007/16905-41931$15.00", "ISSN" : "0003-0147", "PMID" : "17427133", "abstract" : "Across a wide array of animal species, mean population densities decline with species body mass such that the rate of energy use of local populations is approximately independent of body size. This \"energetic equivalence\" is particularly evident when ecological population densities are plotted across several or more orders of magnitude in body mass and is supported by a considerable body of evidence. Nevertheless, interpretation of the data has remained controversial, largely because of the difficulty of explaining the origin and maintenance of such a size-abundance relationship in terms of purely ecological processes. Here I describe results of a simulation model suggesting that an extremely simple mechanism operating over evolutionary time can explain the major features of the empirical data. The model specifies only the size scaling of metabolism and a process where randomly chosen species evolve to take resource energy from other species. This process of energy exchange among particular species is distinct from a random walk of species abundances and creates a situation in which species populations using relatively low amounts of energy at any body size have an elevated extinction risk. Selective extinction of such species rapidly drives size-abundance allometry in faunas toward approximate energetic equivalence and maintains it there.", "author" : [ { "dropping-particle" : "", "family" : "Damuth", "given" : "John", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "The American Naturalist", "id" : "ITEM-1", "issue" : "5", "issued" : { "date-parts" : [ [ "2007", "5" ] ] }, "page" : "621-631", "title" : "A macroevolutionary explanation for energy equivalence in the scaling of body size and population density", "type" : "article-journal", "volume" : "169" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1038/290699a0", "ISSN" : "0028-0836", "author" : [ { "dropping-particle" : "", "family" : "Damuth", "given" : "John", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Nature", "id" : "ITEM-2", "issue" : "5808", "issued" : { "date-parts" : [ [ "1981", "4", "23" ] ] }, "page" : "699-700", "title" : "Population density and body size in mammals", "type" : "article-journal", "volume" : "290" }, "uris" : [ "" ] }, { "id" : "ITEM-3", "itemData" : { "DOI" : "10.1111/j.1095-8312.1987.tb01990.x", "ISSN" : "00244066", "author" : [ { "dropping-particle" : "", "family" : "Damuth", "given" : "John", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Biological Journal of the Linnean Society", "id" : "ITEM-3", "issue" : "3", "issued" : { "date-parts" : [ [ "1987", "7", "28" ] ] }, "page" : "193-246", "title" : "Interspecific allometry of population density in mammals and other animals: the independence of body mass and population energy-use", "type" : "article-journal", "volume" : "31" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Damuth 1981, 1987, 2007)", "plainTextFormattedCitation" : "(Damuth 1981, 1987, 2007)", "previouslyFormattedCitation" : "(Damuth 1981, 1987, 2007)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Damuth 1981, 1987, 2007). Further, subsequent studies have questioned the generality of the relation within clades ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1038/324248a0", "ISBN" : "0028-0836", "ISSN" : "0028-0836", "PMID" : "94", "abstract" : "We present data and analyses demonstrating that large species utilize a disproportionately large share of the resources within local ecosystems. Even though small species tend to have higher local population densities, these are not sufficient to compensate for their lower rates of ...", "author" : [ { "dropping-particle" : "", "family" : "Brown", "given" : "James H.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Maurer", "given" : "Brian a.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Nature", "id" : "ITEM-1", "issued" : { "date-parts" : [ [ "1986" ] ] }, "page" : "248-250", "title" : "Body size, ecological dominance and Cope's rule", "type" : "article", "volume" : "324" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1038/351312a0", "ISBN" : "0028-0836", "ISSN" : "0028-0836", "PMID" : "1992", "abstract" : "The relationship between abundance and body size is the subject of considerable debate in ecology. Several data sets spanning a large range of body sizes show linear negative relationships between abundance and weight when these are measured on a logarithmic scale. But other studies of the abundances of species from single taxa, such as birds, which span a narrower range of body sizes reveal either little or no relationship, or a triangular relationship. Errors in estimating abundance might obscure relationships that do exist over a narrow range of body sizes. We describe here the relationship between body weight and abundance in British birds, whose population size estimates are unusually good. Abundance across all species declines with a -0.75 power of body weight, which conforms with the energetic equivalence 'rule'. There is, however, a significant positive relationship between abundance and body weight within lower taxa. Those tribes that do not share recent common ancestry with other British birds are most likely to show a positive relationship across their constituent species. We thus show that phylogenetic relatedness might be an important indicator of the structure of the relationship between body size and abundance.", "author" : [ { "dropping-particle" : "", "family" : "Nee", "given" : "Sean", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Read", "given" : "Andrew F.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Greenwood", "given" : "Jeremy J. D.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Harvey", "given" : "Paul H.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Nature", "id" : "ITEM-2", "issued" : { "date-parts" : [ [ "1991" ] ] }, "note" : "Orienter dig i citation 1-15", "page" : "312-313", "title" : "The relationship between abundance and body size in British birds", "type" : "article", "volume" : "351" }, "uris" : [ "" ] }, { "id" : "ITEM-3", "itemData" : { "DOI" : "10.1371/journal.pone.0057449", "ISSN" : "19326203", "PMID" : "23460858", "abstract" : "The energy equivalence rule (EER) is a macroecological hypothesis that posits that total population energy use (PEU) should be independent of species body mass, because population densities and energy metabolisms scale with body mass in a directly inverse manner. However, evidence supporting the EER is equivocal, and the use of basal metabolic rate (BMR) in such studies has been questioned; ecologically-relevant indices like field metabolic rate (FMR) are probably more appropriate. In this regard, Australian marsupials present a novel test for the EER because, unlike eutherians, marsupial BMRs and FMRs scale differently with body mass. Based on either FMR or BMR, Australian marsupial PEU did not obey an EER, and scaled positively with body mass based on ordinary least squares (OLS) regressions. Importantly, the scaling of marsupial population density with body mass had a slope of \u22120.37, significantly shallower than the expected slope of \u22120.75, and not directly inverse of body-mass scaling exponents for BMR (0.72) or FMR (0.62). The findings suggest that the EER may not be a causal, universal rule, or that for reasons not yet clear, it is not operating for Australia\u2019s unique native fauna.", "author" : [ { "dropping-particle" : "", "family" : "Munn", "given" : "Adam J.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Dunne", "given" : "Craig", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "M\u00fcller", "given" : "Dennis W H", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Clauss", "given" : "Marcus", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "PLoS ONE", "id" : "ITEM-3", "issue" : "2", "issued" : { "date-parts" : [ [ "2013" ] ] }, "page" : "1-5", "title" : "Energy In-Equivalence in Australian Marsupials: Evidence for Disruption of the Continent's Mammal Assemblage, or Are Rules Meant to Be Broken?", "type" : "article-journal", "volume" : "8" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Brown & Maurer 1986; Nee et al. 1991; Munn et al. 2013)", "manualFormatting" : "(e.g. Brown & Maurer 1986; Nee et al. 1991; Munn et al. 2013)", "plainTextFormattedCitation" : "(Brown & Maurer 1986; Nee et al. 1991; Munn et al. 2013)", "previouslyFormattedCitation" : "(Brown & Maurer 1986; Nee et al. 1991; Munn et al. 2013)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(e.g. Brown & Maurer 1986; Nee et al. 1991; Munn et al. 2013).In this study we aim to contribute to an improved understanding of the so-called global size–density relationship (SDR), the relationship between the average body mass and the average population density for species around the globe unrelated of community or habitat ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1016/j.tree.2007.03.007", "ISSN" : "01695347", "author" : [ { "dropping-particle" : "", "family" : "White", "given" : "Ethan P", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Ernest", "given" : "S.K. Morgan", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Kerkhoff", "given" : "Andrew J", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Enquist", "given" : "Brian J", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Trends in Ecology & Evolution", "id" : "ITEM-1", "issue" : "6", "issued" : { "date-parts" : [ [ "2007", "6" ] ] }, "page" : "323-330", "title" : "Relationships between body size and abundance in ecology", "type" : "article-journal", "volume" : "22" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(White et al. 2007b)", "plainTextFormattedCitation" : "(White et al. 2007b)", "previouslyFormattedCitation" : "(White et al. 2007b)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(White et al. 2007b). Across large scales, broad body mass spans, and broad taxonomic levels the global size–density relationship is generally accepted to have a relationship of b = -0.75 ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1016/j.ecocom.2009.08.005", "ISBN" : "1476-945X", "ISSN" : "1476945X", "abstract" : "The relation between population density and body mass has vexed ecologists for nearly 30 years as a consequence of high variability in the observed slope of the relation: No single generalisation of the relation has been accepted as universally representative. Here, we use a simple computational approach to examine how observational scale (the body mass range considered) determines variation in the density-mass pattern. Our model relies on two assumptions: (1) resources are partitioned in an unbiased manner among species with different masses; (2) the number of individuals that can be supported by a given quantity of resources is related to their metabolic rate (which is a function of their mass raised to the power of a scaling coefficient, b). We show that density (1) scales as a function of body mass raised to the power of -b on average, but (2) the slope of the relation varies considerably at smaller scales of observation (over narrow ranges of body mass) as a consequence of details of species' ecology associated with resource procurement. Historically, the effect of body mass range on the slope of the density-mass relation has been unfailingly attributed to a statistical effect. Here we show that the effect of body mass range on the slope of the density-mass relation may equally result from a biological mechanism, though we find it impossible to distinguish between the two. We observe that many of the explanations that have been offered to account for the variability in the slope of the relation invoke mechanisms associated with differences in body mass and we therefore suggest that body mass range itself might be the most important explanatory factor. Notably, our results imply that the energetic equivalence rule should not be expected to hold at smaller scales of observation, which suggests that it may not be possible to scale the mass- and temperature-dependence of organism metabolism to predict patterns at higher levels of biological organisation at smaller scales of observation. ?? 2009 Elsevier B.V. All rights reserved.", "author" : [ { "dropping-particle" : "", "family" : "Hayward", "given" : "April", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Kolasa", "given" : "Jurek", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Stone", "given" : "Jonathon R.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecological Complexity", "id" : "ITEM-1", "issue" : "1", "issued" : { "date-parts" : [ [ "2010", "3" ] ] }, "page" : "115-124", "publisher" : "Elsevier B.V.", "title" : "The scale-dependence of population density-body mass allometry: Statistical artefact or biological mechanism?", "type" : "article-journal", "volume" : "7" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1111/j.1095-8312.1987.tb01990.x", "ISSN" : "00244066", "author" : [ { "dropping-particle" : "", "family" : "Damuth", "given" : "John", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Biological Journal of the Linnean Society", "id" : "ITEM-2", "issue" : "3", "issued" : { "date-parts" : [ [ "1987", "7", "28" ] ] }, "page" : "193-246", "title" : "Interspecific allometry of population density in mammals and other animals: the independence of body mass and population energy-use", "type" : "article-journal", "volume" : "31" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Damuth 1987; Hayward, Kolasa & Stone 2010)", "plainTextFormattedCitation" : "(Damuth 1987; Hayward, Kolasa & Stone 2010)", "previouslyFormattedCitation" : "(Damuth 1987; Hayward, Kolasa & Stone 2010)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Damuth 1987; Hayward, Kolasa & Stone 2010). On smaller scales and for narrower taxonomic groups there is much more variation in the observed relationship, usually with less steep slopes ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1016/j.ecocom.2009.08.005", "ISBN" : "1476-945X", "ISSN" : "1476945X", "abstract" : "The relation between population density and body mass has vexed ecologists for nearly 30 years as a consequence of high variability in the observed slope of the relation: No single generalisation of the relation has been accepted as universally representative. Here, we use a simple computational approach to examine how observational scale (the body mass range considered) determines variation in the density-mass pattern. Our model relies on two assumptions: (1) resources are partitioned in an unbiased manner among species with different masses; (2) the number of individuals that can be supported by a given quantity of resources is related to their metabolic rate (which is a function of their mass raised to the power of a scaling coefficient, b). We show that density (1) scales as a function of body mass raised to the power of -b on average, but (2) the slope of the relation varies considerably at smaller scales of observation (over narrow ranges of body mass) as a consequence of details of species' ecology associated with resource procurement. Historically, the effect of body mass range on the slope of the density-mass relation has been unfailingly attributed to a statistical effect. Here we show that the effect of body mass range on the slope of the density-mass relation may equally result from a biological mechanism, though we find it impossible to distinguish between the two. We observe that many of the explanations that have been offered to account for the variability in the slope of the relation invoke mechanisms associated with differences in body mass and we therefore suggest that body mass range itself might be the most important explanatory factor. Notably, our results imply that the energetic equivalence rule should not be expected to hold at smaller scales of observation, which suggests that it may not be possible to scale the mass- and temperature-dependence of organism metabolism to predict patterns at higher levels of biological organisation at smaller scales of observation. ?? 2009 Elsevier B.V. All rights reserved.", "author" : [ { "dropping-particle" : "", "family" : "Hayward", "given" : "April", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Kolasa", "given" : "Jurek", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Stone", "given" : "Jonathon R.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecological Complexity", "id" : "ITEM-1", "issue" : "1", "issued" : { "date-parts" : [ [ "2010", "3" ] ] }, "page" : "115-124", "publisher" : "Elsevier B.V.", "title" : "The scale-dependence of population density-body mass allometry: Statistical artefact or biological mechanism?", "type" : "article-journal", "volume" : "7" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1111/j.1466-8238.2012.00782.x", "ISBN" : "1466-8238", "ISSN" : "1466822X", "abstract" : "Energy equivalence, the notion that population energy flux is independent of body mass, has become a key concept in ecology. We argue that energy equivalence is not an ecological \u2018rule\u2019, as claimed, but a flawed concept beset by circular reasoning. In fact, the independence of mass and energy flux is a null hypothesis. We show that our mechanistic understanding of size\u2013density relationships (SDRs) follows directly from this null model and the assumption that energy limits abundance. Paradoxically, without this assumption energy equivalence has no meaning and we lack a mechanistic understanding for SDRs. We derive an expression for the strength (r2) of SDRs under the null model, which provides a framework within which to compare published SDRs. This confirms that tight correlations between mass and abundance are a trivial consequence of the span of body masses considered. Our model implies that energy flux varies by five to six orders of magnitude among similarly sized mammals and to a far greater extent in birds. We conclude that the energetic paradigm can be strengthened by considering alternative, non-energetic, hypotheses.", "author" : [ { "dropping-particle" : "", "family" : "Isaac", "given" : "Nick J B", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Storch", "given" : "David", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Carbone", "given" : "Chris", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Global Ecology and Biogeography", "id" : "ITEM-2", "issued" : { "date-parts" : [ [ "2013" ] ] }, "page" : "1-5", "title" : "The paradox of energy equivalence", "type" : "article-journal", "volume" : "22" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Hayward et al. 2010; Isaac, Storch & Carbone 2013)", "plainTextFormattedCitation" : "(Hayward et al. 2010; Isaac, Storch & Carbone 2013)", "previouslyFormattedCitation" : "(Hayward et al. 2010; Isaac, Storch & Carbone 2013)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Hayward et al. 2010; Isaac, Storch & Carbone 2013). While increased variation of the slope estimates found within narrower taxonomic groups has been claimed to be a statistical artefact of modelling on a smaller range of body masses, it does not account for the one-sided bias in most studies towards shallower slopes ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1016/j.ecocom.2009.08.005", "ISBN" : "1476-945X", "ISSN" : "1476945X", "abstract" : "The relation between population density and body mass has vexed ecologists for nearly 30 years as a consequence of high variability in the observed slope of the relation: No single generalisation of the relation has been accepted as universally representative. Here, we use a simple computational approach to examine how observational scale (the body mass range considered) determines variation in the density-mass pattern. Our model relies on two assumptions: (1) resources are partitioned in an unbiased manner among species with different masses; (2) the number of individuals that can be supported by a given quantity of resources is related to their metabolic rate (which is a function of their mass raised to the power of a scaling coefficient, b). We show that density (1) scales as a function of body mass raised to the power of -b on average, but (2) the slope of the relation varies considerably at smaller scales of observation (over narrow ranges of body mass) as a consequence of details of species' ecology associated with resource procurement. Historically, the effect of body mass range on the slope of the density-mass relation has been unfailingly attributed to a statistical effect. Here we show that the effect of body mass range on the slope of the density-mass relation may equally result from a biological mechanism, though we find it impossible to distinguish between the two. We observe that many of the explanations that have been offered to account for the variability in the slope of the relation invoke mechanisms associated with differences in body mass and we therefore suggest that body mass range itself might be the most important explanatory factor. Notably, our results imply that the energetic equivalence rule should not be expected to hold at smaller scales of observation, which suggests that it may not be possible to scale the mass- and temperature-dependence of organism metabolism to predict patterns at higher levels of biological organisation at smaller scales of observation. ?? 2009 Elsevier B.V. All rights reserved.", "author" : [ { "dropping-particle" : "", "family" : "Hayward", "given" : "April", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Kolasa", "given" : "Jurek", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Stone", "given" : "Jonathon R.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecological Complexity", "id" : "ITEM-1", "issue" : "1", "issued" : { "date-parts" : [ [ "2010", "3" ] ] }, "page" : "115-124", "publisher" : "Elsevier B.V.", "title" : "The scale-dependence of population density-body mass allometry: Statistical artefact or biological mechanism?", "type" : "article-journal", "volume" : "7" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Hayward et al. 2010)", "plainTextFormattedCitation" : "(Hayward et al. 2010)", "previouslyFormattedCitation" : "(Hayward et al. 2010)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Hayward et al. 2010). The general tendencies of the relationship is that guilds with low mean body-mass (e.g. rodents) are often found to have near zero slopes or even positive slopes, while guilds with medium to heavy body mass have slopes that are closer to -0.75 or have even steeper slopes ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1086/284596", "ISBN" : "00030147", "ISSN" : "0003-0147", "abstract" : "Accounting for the variation in the population density among different animal species is a central goal of animal ecology (Andrewartha and Birch 1954). Among mammals, density is closely related to the average adult body mass of the species and to the trophic level occupied by the species (Mohr 1940; Clutton-Brock and Harvey 1977; Eisenberg 1980; Damuth 1981a; Peters 1983; Peters and Raelson 1984). In addition, after the body mass and trophic position of species have been taken into account, population densities appear to vary with habitat (Eisenberg 1980) and biogeographical area (Peters and Raelson 1984). Peters and Raelson have been impressed by the predictive power of these relations: \"Because these relations appear so powerful . . . they should be examined as fully as possible before they come into widespread use\" (1984, p. 499).", "author" : [ { "dropping-particle" : "", "family" : "Robinson", "given" : "John G.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Redford", "given" : "Kent H.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "The American Naturalist", "id" : "ITEM-1", "issue" : "5", "issued" : { "date-parts" : [ [ "1986", "11" ] ] }, "page" : "665-680", "title" : "Body size, diet, and population density of neotropical forest mammals", "type" : "article-journal", "volume" : "128" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1038/324248a0", "ISBN" : "0028-0836", "ISSN" : "0028-0836", "PMID" : "94", "abstract" : "We present data and analyses demonstrating that large species utilize a disproportionately large share of the resources within local ecosystems. Even though small species tend to have higher local population densities, these are not sufficient to compensate for their lower rates of ...", "author" : [ { "dropping-particle" : "", "family" : "Brown", "given" : "James H.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Maurer", "given" : "Brian a.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Nature", "id" : "ITEM-2", "issued" : { "date-parts" : [ [ "1986" ] ] }, "page" : "248-250", "title" : "Body size, ecological dominance and Cope's rule", "type" : "article", "volume" : "324" }, "uris" : [ "" ] }, { "id" : "ITEM-3", "itemData" : { "DOI" : "10.1038/351312a0", "ISBN" : "0028-0836", "ISSN" : "0028-0836", "PMID" : "1992", "abstract" : "The relationship between abundance and body size is the subject of considerable debate in ecology. Several data sets spanning a large range of body sizes show linear negative relationships between abundance and weight when these are measured on a logarithmic scale. But other studies of the abundances of species from single taxa, such as birds, which span a narrower range of body sizes reveal either little or no relationship, or a triangular relationship. Errors in estimating abundance might obscure relationships that do exist over a narrow range of body sizes. We describe here the relationship between body weight and abundance in British birds, whose population size estimates are unusually good. Abundance across all species declines with a -0.75 power of body weight, which conforms with the energetic equivalence 'rule'. There is, however, a significant positive relationship between abundance and body weight within lower taxa. Those tribes that do not share recent common ancestry with other British birds are most likely to show a positive relationship across their constituent species. We thus show that phylogenetic relatedness might be an important indicator of the structure of the relationship between body size and abundance.", "author" : [ { "dropping-particle" : "", "family" : "Nee", "given" : "Sean", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Read", "given" : "Andrew F.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Greenwood", "given" : "Jeremy J. D.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Harvey", "given" : "Paul H.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Nature", "id" : "ITEM-3", "issued" : { "date-parts" : [ [ "1991" ] ] }, "note" : "Orienter dig i citation 1-15", "page" : "312-313", "title" : "The relationship between abundance and body size in British birds", "type" : "article", "volume" : "351" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Brown & Maurer 1986; Robinson & Redford 1986; Nee et al. 1991)", "plainTextFormattedCitation" : "(Brown & Maurer 1986; Robinson & Redford 1986; Nee et al. 1991)", "previouslyFormattedCitation" : "(Brown & Maurer 1986; Robinson & Redford 1986; Nee et al. 1991)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Brown & Maurer 1986; Robinson & Redford 1986; Nee et al. 1991), and closely related groups tend to show shallower relationships than the overall pattern (e.g. it has been shown to be only around -0.37 in Australian marsupials ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1371/journal.pone.0057449", "ISSN" : "19326203", "PMID" : "23460858", "abstract" : "The energy equivalence rule (EER) is a macroecological hypothesis that posits that total population energy use (PEU) should be independent of species body mass, because population densities and energy metabolisms scale with body mass in a directly inverse manner. However, evidence supporting the EER is equivocal, and the use of basal metabolic rate (BMR) in such studies has been questioned; ecologically-relevant indices like field metabolic rate (FMR) are probably more appropriate. In this regard, Australian marsupials present a novel test for the EER because, unlike eutherians, marsupial BMRs and FMRs scale differently with body mass. Based on either FMR or BMR, Australian marsupial PEU did not obey an EER, and scaled positively with body mass based on ordinary least squares (OLS) regressions. Importantly, the scaling of marsupial population density with body mass had a slope of \u22120.37, significantly shallower than the expected slope of \u22120.75, and not directly inverse of body-mass scaling exponents for BMR (0.72) or FMR (0.62). The findings suggest that the EER may not be a causal, universal rule, or that for reasons not yet clear, it is not operating for Australia\u2019s unique native fauna.", "author" : [ { "dropping-particle" : "", "family" : "Munn", "given" : "Adam J.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Dunne", "given" : "Craig", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "M\u00fcller", "given" : "Dennis W H", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Clauss", "given" : "Marcus", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "PLoS ONE", "id" : "ITEM-1", "issue" : "2", "issued" : { "date-parts" : [ [ "2013" ] ] }, "page" : "1-5", "title" : "Energy In-Equivalence in Australian Marsupials: Evidence for Disruption of the Continent's Mammal Assemblage, or Are Rules Meant to Be Broken?", "type" : "article-journal", "volume" : "8" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Munn et al. 2013)", "plainTextFormattedCitation" : "(Munn et al. 2013)", "previouslyFormattedCitation" : "(Munn et al. 2013)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Munn et al. 2013)). A mechanistic model framework shows that SDR is dependent on how the food resources scale with the consumer body mass, giving markedly different expectations for the relationship between herbivores and carnivores ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1086/519858", "ISBN" : "0003-0147/2007/17003-42157$15.00", "ISSN" : "0003-0147", "PMID" : "17879198", "abstract" : "The negative scaling of plant and animal abundance with body mass is one of the most fundamental relationships in ecology. However, theoretical approaches to explain this phenomenon make the unrealistic assumption that species share a homogeneous resource. Here we present a simple model linking mass and metabolism with density that includes the effects of consumer size on resource characteristics (particle size, density, and distribution). We predict patterns consistent with the energy equivalence rule (EER) under some scenarios. However, deviations from EER occur as a result of variation in resource distribution and productivity (e.g., due to the clumping of prey or variation in food particle size selection). We also predict that abundance scaling exponents change with the dimensionality of the foraging habitat. Our model predictions explain several inconsistencies in the observed scaling of vertebrate abundance among ecological and taxonomic groups and provide a broad framework for understanding variation in abundance.", "author" : [ { "dropping-particle" : "", "family" : "Carbone", "given" : "Chris", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Rowcliffe", "given" : "J. Marcus", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Cowlishaw", "given" : "Guy", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Isaac", "given" : "Nick J. B.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "The American Naturalist", "id" : "ITEM-1", "issue" : "3", "issued" : { "date-parts" : [ [ "2007", "9" ] ] }, "page" : "479-484", "title" : "The Scaling of Abundance in Consumers and Their Resources: Implications for the Energy Equivalence Rule", "type" : "article-journal", "volume" : "170" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Carbone et al. 2007)", "plainTextFormattedCitation" : "(Carbone et al. 2007)", "previouslyFormattedCitation" : "(Carbone et al. 2007)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Carbone et al. 2007). The carnivore SDR tend to be steeper than herbivore SDR, since larger predators take larger prey and larger prey tend to be distributed less evenly, which makes prey scarcer for larger carnivores ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1086/519858", "ISBN" : "0003-0147/2007/17003-42157$15.00", "ISSN" : "0003-0147", "PMID" : "17879198", "abstract" : "The negative scaling of plant and animal abundance with body mass is one of the most fundamental relationships in ecology. However, theoretical approaches to explain this phenomenon make the unrealistic assumption that species share a homogeneous resource. Here we present a simple model linking mass and metabolism with density that includes the effects of consumer size on resource characteristics (particle size, density, and distribution). We predict patterns consistent with the energy equivalence rule (EER) under some scenarios. However, deviations from EER occur as a result of variation in resource distribution and productivity (e.g., due to the clumping of prey or variation in food particle size selection). We also predict that abundance scaling exponents change with the dimensionality of the foraging habitat. Our model predictions explain several inconsistencies in the observed scaling of vertebrate abundance among ecological and taxonomic groups and provide a broad framework for understanding variation in abundance.", "author" : [ { "dropping-particle" : "", "family" : "Carbone", "given" : "Chris", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Rowcliffe", "given" : "J. Marcus", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Cowlishaw", "given" : "Guy", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Isaac", "given" : "Nick J. B.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "The American Naturalist", "id" : "ITEM-1", "issue" : "3", "issued" : { "date-parts" : [ [ "2007", "9" ] ] }, "page" : "479-484", "title" : "The Scaling of Abundance in Consumers and Their Resources: Implications for the Energy Equivalence Rule", "type" : "article-journal", "volume" : "170" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Carbone et al. 2007)", "plainTextFormattedCitation" : "(Carbone et al. 2007)", "previouslyFormattedCitation" : "(Carbone et al. 2007)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Carbone et al. 2007). Further, we expect carnivorous to be generally offset to lower population densities than herbivores at any given body mass due to the drop of available energy higher in the food chain ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.2307/1930126", "ISSN" : "00129658", "author" : [ { "dropping-particle" : "", "family" : "Lindeman", "given" : "Raymond L.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecology", "id" : "ITEM-1", "issue" : "4", "issued" : { "date-parts" : [ [ "1942", "10" ] ] }, "page" : "399-417", "title" : "The Trophic-Dynamic Aspect of Ecology", "type" : "article-journal", "volume" : "23" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Lindeman 1942)", "plainTextFormattedCitation" : "(Lindeman 1942)", "previouslyFormattedCitation" : "(Lindeman 1942)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Lindeman 1942).Since both basal and field metabolic rate was found to be increasing with body mass ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "author" : [ { "dropping-particle" : "", "family" : "Kleiber", "given" : "M", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Physiological reviews", "id" : "ITEM-1", "issue" : "4", "issued" : { "date-parts" : [ [ "1947" ] ] }, "page" : "511-541", "title" : "Body size and metabolic rate", "type" : "article-journal", "volume" : "27" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Kleiber 1947)", "plainTextFormattedCitation" : "(Kleiber 1947)", "previouslyFormattedCitation" : "(Kleiber 1947)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Kleiber 1947) inversely proportional to population density, Damuth (1981) concluded that population level metabolic rate was independent of body mass (now known as the energetic equivalence rule (EER)). Later studies have shown that metabolism does not scale universally with a body mass slope of 0.75, but is scale dependent, and is often different from 0.75 within phylogenetic clades ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1086/606023", "ISSN" : "0003-0147", "author" : [ { "dropping-particle" : "", "family" : "Sieg", "given" : "Annette\u00a0E.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "O\u2019Connor", "given" : "Michael\u00a0P.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "McNair", "given" : "James\u00a0N.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Grant", "given" : "Bruce\u00a0W.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Agosta", "given" : "Salvatore\u00a0J.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Dunham", "given" : "Arthur\u00a0E.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "The American Naturalist", "id" : "ITEM-1", "issue" : "5", "issued" : { "date-parts" : [ [ "2009" ] ] }, "page" : "720-733", "title" : "Mammalian Metabolic Allometry: Do Intraspecific Variation, Phylogeny, and Regression Models Matter?", "type" : "article-journal", "volume" : "174" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1186/1742-4682-1-13", "ISBN" : "1742-4682 (Electronic)", "ISSN" : "1742-4682", "PMID" : "15546492", "abstract" : "BACKGROUND: The relationship between body mass (M) and standard metabolic rate (B) among living organisms remains controversial, though it is widely accepted that in many cases B is approximately proportional to the three-quarters power of M. RESULTS: The biological significance of the straight-line plots obtained over wide ranges of species when B is plotted against log M remains a matter of debate. In this article we review the values ascribed to the gradients of such graphs (typically 0.75, according to the majority view), and we assess various attempts to explain the allometric power-law phenomenon, placing emphasis on the most recent publications. CONCLUSION: Although many of the models that have been advanced have significant attractions, none can be accepted without serious reservations, and the possibility that no one model can fit all cases has to be more seriously entertained.", "author" : [ { "dropping-particle" : "", "family" : "Agutter", "given" : "Paul S", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Wheatley", "given" : "Denys N", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Theoretical biology & medical modelling", "id" : "ITEM-2", "issued" : { "date-parts" : [ [ "2004" ] ] }, "page" : "13", "title" : "Metabolic scaling: consensus or controversy?", "type" : "article-journal", "volume" : "1" }, "uris" : [ "" ] }, { "id" : "ITEM-3", "itemData" : { "DOI" : "10.1242/jeb.01553", "ISBN" : "0022-0949", "ISSN" : "0022-0949", "PMID" : "15855393", "abstract" : "The field metabolic rates (FMRs) of 229 species of terrestrial vertebrates, all measured using the doubly labeled water method in free-living individuals, were evaluated. Daily rates of energy expenditure were as low as 0.23 kJ per day in a small reptile (gecko), to as high as 52 500 kJ per day in a marine mammal (seal). This is a range of nearly six orders of magnitude. More than 70% of the variation in log-transformed data is due to variation in body size (expressed as body mass). Much of the remaining variation is accounted for by thermal physiology, with the endothermic mammals and birds having FMRs that are about 12 and 20 times higher, respectively, than FMRs of equivalent-sized, but ectothermic, reptiles. Variation in log(body mass) within each of these three taxonomic classes accounts for over 94% of the variation in log(FMR), and results from nonlinear regression analyses using untransformed data support this conclusion. However, the range of residual variation in mass-adjusted FMR within classes is still more than sixfold (ratio of highest over lowest). Some of this variation is associated with affiliations with lower taxonomic levels (Infraclass: eutherian vs metatherian mammals; Family: passerine, procellariform and galliform birds vs other birds), some is associated with habitat (especially desert vs nondesert), and some with differences in basic diet preference and foraging mode and season. The scaling slopes for FMR often differ from BMR slopes for the same Class of animals, and most differ from the theoretical slope of 0.75. Differences among slopes and intercepts that were detected using conventional regression analyses were largely confirmed upon reanalysis using Independent Contrasts Analysis to adjust for phylogenetic biases.", "author" : [ { "dropping-particle" : "", "family" : "Nagy", "given" : "Kenneth A", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Journal of Experimental Biology", "id" : "ITEM-3", "issued" : { "date-parts" : [ [ "2005" ] ] }, "page" : "1621-1625", "title" : "Field metabolic rate and body size", "type" : "article-journal", "volume" : "208" }, "uris" : [ "" ] }, { "id" : "ITEM-4", "itemData" : { "DOI" : "10.1111/j.1461-0248.2010.01461.x", "ISBN" : "1461-023X", "ISSN" : "1461023X", "PMID" : "20353439", "abstract" : "The metabolic theory of ecology links physiology with ecology, and successfully predicts many allometric scaling relationships. In recent years, proponents and critics of metabolic theory have debated vigorously about the scaling of metabolic rate. We show that the controversy arose, in part, because researchers examined the mean exponent separately from the variance. We estimate both quantities simultaneously using linear mixed-effects models and data from 1242 animal species. Metabolic rate scaling converges on the predicted value of 3/4 but is highly heterogeneous: 50% of orders lie outside the range 0.68-0.82. These findings are robust to several forms of statistical uncertainty. We then test competing hypotheses about the variation. Metabolic theory is currently unable to explain differences in scaling among orders, but the patterns are not consistent with competing explanations either. We conclude that current theories are inadequate to explain the full range of metabolic scaling patterns observed in nature.", "author" : [ { "dropping-particle" : "", "family" : "Isaac", "given" : "N. J B", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Carbone", "given" : "Chris", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecology Letters", "id" : "ITEM-4", "issued" : { "date-parts" : [ [ "2010" ] ] }, "page" : "728-735", "title" : "Why are metabolic scaling exponents so controversial? Quantifying variance and testing hypotheses", "type" : "article", "volume" : "13" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Agutter & Wheatley 2004; Nagy 2005; Sieg et al. 2009; Isaac & Carbone 2010)", "plainTextFormattedCitation" : "(Agutter & Wheatley 2004; Nagy 2005; Sieg et al. 2009; Isaac & Carbone 2010)", "previouslyFormattedCitation" : "(Agutter & Wheatley 2004; Nagy 2005; Sieg et al. 2009; Isaac & Carbone 2010)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Agutter & Wheatley 2004; Nagy 2005; Sieg et al. 2009; Isaac & Carbone 2010), calling the EER into question. The scaling exponent of metabolic rate has been found to vary between taxonomic groups ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1890/05-1883", "ISSN" : "0012-9658", "author" : [ { "dropping-particle" : "", "family" : "White", "given" : "Craig R.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Cassey", "given" : "Phillip", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Blackburn", "given" : "Tim M.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecology", "id" : "ITEM-1", "issue" : "2", "issued" : { "date-parts" : [ [ "2007", "2" ] ] }, "page" : "315-323", "title" : "Allometric exponents do not support a universal metabolic allometry", "type" : "article-journal", "volume" : "88" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1002/cphy.c110049", "ISBN" : "2040-4603", "ISSN" : "20404603", "PMID" : "24692144", "abstract" : "Life on earth spans a size range of around 21 orders of magnitude across species and can span a range of more than 6 orders of magnitude within species of animal. The effect of size on physiology is, therefore, enormous and is typically expressed by how physiological phenomena scale with mass b . When b = 1 a trait does not vary in direct proportion to mass and is said to scale allometrically. The study of allometric scaling goes back to at least the time of Galileo Galilei, and published scaling relationships are now available for hundreds of traits. Here, the methods of scaling analysis are reviewed, using examples for a range of traits with an emphasis on those related to metabolism in animals. Where necessary, new relationships have been generated from published data using modern phylogenetically informed techniques. During recent decades one of the most controversial scaling relationships has been that between metabolic rate and body mass and a number of explanations have been proposed for the scaling of this trait. Examples of these mechanistic explanations for metabolic scaling are reviewed, and suggestions made for comparing between them. Finally, the conceptual links between metabolic scaling and ecological patterns are examined, emphasizing the distinction between (1) the hypothesis that size-and temperature-dependent variation among species and individuals in metabolic rate influences ecological processes at levels of organization from individuals to the biosphere and (2) mechanistic explanations for metabolic rate that may explain the size-and temperature-dependence of this trait. C 2014 American Physiological Society.", "author" : [ { "dropping-particle" : "", "family" : "White", "given" : "Craig R.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Kearney", "given" : "Michael R.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Comprehensive Physiology", "id" : "ITEM-2", "issue" : "1", "issued" : { "date-parts" : [ [ "2014" ] ] }, "page" : "231-256", "title" : "Metabolic scaling in animals: Methods, empirical results, and theoretical explanations", "type" : "article-journal", "volume" : "4" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(White, Cassey & Blackburn 2007a; White & Kearney 2014)", "plainTextFormattedCitation" : "(White, Cassey & Blackburn 2007a; White & Kearney 2014)", "previouslyFormattedCitation" : "(White, Cassey & Blackburn 2007a; White & Kearney 2014)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(White, Cassey & Blackburn 2007a; White & Kearney 2014), which under the assumption of the EER would lead us to expect that the SDR of related groups should vary as well. In simulations, Isaac et al. (2013) have shown that under the assumption that density is energy limited, a strong density-mass relationship is to be expected, and therefore we should consider EER a null model to be tested. Another simulation study has shown how the scaling coefficient can arise through small evolutionary steps driven by an increased extinction risk for species in energetic disequilibrium compared to their expected density given a specific metabolic scaling ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1086/513495", "ISBN" : "0003-0147/2007/16905-41931$15.00", "ISSN" : "0003-0147", "PMID" : "17427133", "abstract" : "Across a wide array of animal species, mean population densities decline with species body mass such that the rate of energy use of local populations is approximately independent of body size. This \"energetic equivalence\" is particularly evident when ecological population densities are plotted across several or more orders of magnitude in body mass and is supported by a considerable body of evidence. Nevertheless, interpretation of the data has remained controversial, largely because of the difficulty of explaining the origin and maintenance of such a size-abundance relationship in terms of purely ecological processes. Here I describe results of a simulation model suggesting that an extremely simple mechanism operating over evolutionary time can explain the major features of the empirical data. The model specifies only the size scaling of metabolism and a process where randomly chosen species evolve to take resource energy from other species. This process of energy exchange among particular species is distinct from a random walk of species abundances and creates a situation in which species populations using relatively low amounts of energy at any body size have an elevated extinction risk. Selective extinction of such species rapidly drives size-abundance allometry in faunas toward approximate energetic equivalence and maintains it there.", "author" : [ { "dropping-particle" : "", "family" : "Damuth", "given" : "John", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "The American Naturalist", "id" : "ITEM-1", "issue" : "5", "issued" : { "date-parts" : [ [ "2007", "5" ] ] }, "page" : "621-631", "title" : "A macroevolutionary explanation for energy equivalence in the scaling of body size and population density", "type" : "article-journal", "volume" : "169" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Damuth 2007)", "plainTextFormattedCitation" : "(Damuth 2007)", "previouslyFormattedCitation" : "(Damuth 2007)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Damuth 2007). When slopes diverge from the null expectation of EER, then partitioning of resources is no longer invariant of body mass. A slope more negative than the general size–density relationship means that smaller-bodied species in a clade take a relatively larger share of the resource pool, while less negative slope signify that bigger species tend to claim a larger part of the resource pool ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1016/j.ecocom.2009.08.005", "ISBN" : "1476-945X", "ISSN" : "1476945X", "abstract" : "The relation between population density and body mass has vexed ecologists for nearly 30 years as a consequence of high variability in the observed slope of the relation: No single generalisation of the relation has been accepted as universally representative. Here, we use a simple computational approach to examine how observational scale (the body mass range considered) determines variation in the density-mass pattern. Our model relies on two assumptions: (1) resources are partitioned in an unbiased manner among species with different masses; (2) the number of individuals that can be supported by a given quantity of resources is related to their metabolic rate (which is a function of their mass raised to the power of a scaling coefficient, b). We show that density (1) scales as a function of body mass raised to the power of -b on average, but (2) the slope of the relation varies considerably at smaller scales of observation (over narrow ranges of body mass) as a consequence of details of species' ecology associated with resource procurement. Historically, the effect of body mass range on the slope of the density-mass relation has been unfailingly attributed to a statistical effect. Here we show that the effect of body mass range on the slope of the density-mass relation may equally result from a biological mechanism, though we find it impossible to distinguish between the two. We observe that many of the explanations that have been offered to account for the variability in the slope of the relation invoke mechanisms associated with differences in body mass and we therefore suggest that body mass range itself might be the most important explanatory factor. Notably, our results imply that the energetic equivalence rule should not be expected to hold at smaller scales of observation, which suggests that it may not be possible to scale the mass- and temperature-dependence of organism metabolism to predict patterns at higher levels of biological organisation at smaller scales of observation. ?? 2009 Elsevier B.V. All rights reserved.", "author" : [ { "dropping-particle" : "", "family" : "Hayward", "given" : "April", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Kolasa", "given" : "Jurek", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Stone", "given" : "Jonathon R.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecological Complexity", "id" : "ITEM-1", "issue" : "1", "issued" : { "date-parts" : [ [ "2010", "3" ] ] }, "page" : "115-124", "publisher" : "Elsevier B.V.", "title" : "The scale-dependence of population density-body mass allometry: Statistical artefact or biological mechanism?", "type" : "article-journal", "volume" : "7" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Hayward et al. 2010)", "plainTextFormattedCitation" : "(Hayward et al. 2010)", "previouslyFormattedCitation" : "(Hayward et al. 2010)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Hayward et al. 2010). Several studies have shown energetic in-equivalence where population densities scale more weakly with body mass than metabolism does and bigger species therefore use a disproportionate amount of the available resources than expected under the EER ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1098/rsbl.2011.0036", "ISBN" : "1744-9561", "ISSN" : "1744-9561", "PMID" : "21367781", "abstract" : "The energetic equivalence rule states that population-level metabolic rate is independent of average body size. This rule has been both supported and refuted by allometric studies of abundance and individual metabolic rate, but no study, to my knowledge, has tested the rule with direct measurements of whole-population metabolic rate. Here, I find a positive scaling of whole-colony metabolic rate with body size for eusocial insects. Individual metabolic rates in these colonies scaled with body size more steeply than expected from laboratory studies on insects, while population size was independent of body size. Using consumer-resource models, I suggest that the colony-level metabolic rate scaling observed here may arise from a change in the scaling of individual metabolic rate resulting from a change in the body size dependence of mortality rates.", "author" : [ { "dropping-particle" : "", "family" : "DeLong", "given" : "John P", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Biology Letters", "id" : "ITEM-1", "issue" : "March", "issued" : { "date-parts" : [ [ "2011" ] ] }, "page" : "611-614", "title" : "Energetic inequivalence in eusocial insect colonies.", "type" : "article-journal", "volume" : "7" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1890/13-0620.1", "ISBN" : "0012-9658", "ISSN" : "00129658", "PMID" : "24669745", "abstract" : "Ecological communities consist of small abundant and large non-abundant species. The energetic equivalence rule is an often-observed pattern that could be explained by equal energy usage among abundant small organisms and non-abundant large organisms. To generate this pattern, metabolism (as an indicator of individual energy use) and abundance have to scale inversely with body mass, and cancel each other out. In contrast, the pattern referred to as biomass equivalence states that the biomass of all species in an area should be constant across the body-mass range. In this study, we investigated forest soil communities with respect to metabolism, abundance, population energy use, and biomass. We focused on four land-use types in three different landscape blocks (Biodiversity Exploratories). The soil samples contained 870 species across 12 phylogenetic groups. Our results indicated positive sublinear metabolic scaling and negative sublinear abundance scaling with species body mass. The relationships varied mainly due to differences among phylogenetic groups or feeding types, and only marginally due to land-use type. However, these scaling relationships were not exactly inverse to each other, resulting in increasing population energy use and biomass with increasing body mass for most combinations of phylogenetic group or feeding type with land-use type. Thus, our results are mostly inconsistent with the classic perception of energetic equivalence, and reject the biomass equivalence hypothesis while documenting a specific and nonrandom pattern of how abundance, energy use, and biomass are distributed across size classes. However, these patterns are consistent with two alternative predictions: the resource-thinning hypothesis, which states that abundance decreases with trophic level, and the allometric degree hypothesis, which states that population energy use should increase with population average body mass, due to correlations with the number of links of consumers and resources. Overall, our results suggest that a synthesis of food web structures with metabolic theory may be most promising for predicting natural patterns of abundance, biomass, and energy use.", "author" : [ { "dropping-particle" : "", "family" : "Ehnes", "given" : "Roswitha B.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Pollierer", "given" : "Melanie M.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Erdmann", "given" : "Georgia", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Klarner", "given" : "Bernhard", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Eitzinger", "given" : "Bernhard", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Digel", "given" : "Christoph", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Ott", "given" : "David", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Maraun", "given" : "Mark", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Scheu", "given" : "Stefan", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Brose", "given" : "Ulrich", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecology", "id" : "ITEM-2", "issue" : "2", "issued" : { "date-parts" : [ [ "2014" ] ] }, "page" : "527-537", "title" : "Lack of energetic equivalence in forest soil invertebrates", "type" : "article-journal", "volume" : "95" }, "uris" : [ "" ] }, { "id" : "ITEM-3", "itemData" : { "DOI" : "10.1086/345938", "ISBN" : "0003-0147", "ISSN" : "0003-0147", "PMID" : "12675372", "abstract" : "We studied size-abundance relationships in a species-rich Amazonian bird community and found that the slope of the logarithmic relationship between population density and bodymass (b = -0.22) is significantly shallower than expected under Damuth's energetic equivalence rule (EER), which states that population energy use (PEU) is independent of species body mass. We used estimates of avian field metabolic rates to examine the logarithmic relationship between PEU and body mass and its variation among ecological guilds. The relationship for all species had a significantly positive slope (b = 0.46), indicating that PEU of larger species was greater than that of smaller species. Analyses of guilds revealed significant variation. The slopes of the frugivore-omnivore, insectivore, and granivore guilds were all significantly positive, with that of the frugivore-omnivore guild being the steepest. In contrast, PEU did not vary significantly with species body mass among raptors. These results were confirmed, in analyses using both species values and phylogenetically independent contrasts, and the results do not support the EER in this community. The spatial distribution of resources and mechanisms of interference competition within guilds may explain why most patterns differed from the predictions of the EER. Other sources of variation, including the effects of scale, are also discussed.", "author" : [ { "dropping-particle" : "", "family" : "Russo", "given" : "Sabrina E", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Robinson", "given" : "Scott K", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Terborgh", "given" : "John", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "The American naturalist", "id" : "ITEM-3", "issue" : "2", "issued" : { "date-parts" : [ [ "2003" ] ] }, "page" : "267-283", "title" : "Size-abundance relationships in an Amazonian bird community: implications for the energetic equivalence rule.", "type" : "article-journal", "volume" : "161" }, "uris" : [ "" ] }, { "id" : "ITEM-4", "itemData" : { "DOI" : "10.1371/journal.pone.0057449", "ISSN" : "19326203", "PMID" : "23460858", "abstract" : "The energy equivalence rule (EER) is a macroecological hypothesis that posits that total population energy use (PEU) should be independent of species body mass, because population densities and energy metabolisms scale with body mass in a directly inverse manner. However, evidence supporting the EER is equivocal, and the use of basal metabolic rate (BMR) in such studies has been questioned; ecologically-relevant indices like field metabolic rate (FMR) are probably more appropriate. In this regard, Australian marsupials present a novel test for the EER because, unlike eutherians, marsupial BMRs and FMRs scale differently with body mass. Based on either FMR or BMR, Australian marsupial PEU did not obey an EER, and scaled positively with body mass based on ordinary least squares (OLS) regressions. Importantly, the scaling of marsupial population density with body mass had a slope of \u22120.37, significantly shallower than the expected slope of \u22120.75, and not directly inverse of body-mass scaling exponents for BMR (0.72) or FMR (0.62). The findings suggest that the EER may not be a causal, universal rule, or that for reasons not yet clear, it is not operating for Australia\u2019s unique native fauna.", "author" : [ { "dropping-particle" : "", "family" : "Munn", "given" : "Adam J.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Dunne", "given" : "Craig", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "M\u00fcller", "given" : "Dennis W H", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Clauss", "given" : "Marcus", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "PLoS ONE", "id" : "ITEM-4", "issue" : "2", "issued" : { "date-parts" : [ [ "2013" ] ] }, "page" : "1-5", "title" : "Energy In-Equivalence in Australian Marsupials: Evidence for Disruption of the Continent's Mammal Assemblage, or Are Rules Meant to Be Broken?", "type" : "article-journal", "volume" : "8" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Russo, Robinson & Terborgh 2003; DeLong 2011; Munn et al. 2013; Ehnes et al. 2014)", "plainTextFormattedCitation" : "(Russo, Robinson & Terborgh 2003; DeLong 2011; Munn et al. 2013; Ehnes et al. 2014)", "previouslyFormattedCitation" : "(Russo, Robinson & Terborgh 2003; DeLong 2011; Munn et al. 2013; Ehnes et al. 2014)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Russo, Robinson & Terborgh 2003; DeLong 2011; Munn et al. 2013; Ehnes et al. 2014). Under EER we would expect the SDR of clades to mirror their metabolic rate increase with body mass, and even lower slopes if bigger species do use a disproportionate amount of resources.One of the most used methods in predicting species densities is the allometric size-density relationship. Accurate estimates of population densities are important for analyses of fauna ecosystem effects ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1126/science.1251817", "ISSN" : "0036-8075", "PMID" : "25061202", "abstract" : "We live amid a global wave of anthropogenically driven biodiversity loss: species and population extirpations and, critically, declines in local species abundance. Particularly, human impacts on animal biodiversity are an under-recognized form of global environmental change. Among terrestrial vertebrates, 322 species have become extinct since 1500, and populations of the remaining species show 25% average decline in abundance. Invertebrate patterns are equally dire: 67% of monitored populations show 45% mean abundance decline. Such animal declines will cascade onto ecosystem functioning and human well-being. Much remains unknown about this \"Anthropocene defaunation\"; these knowledge gaps hinder our capacity to predict and limit defaunation impacts. Clearly, however, defaunation is both a pervasive component of the planet's sixth mass extinction and also a major driver of global ecological change.", "author" : [ { "dropping-particle" : "", "family" : "Dirzo", "given" : "R.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Young", "given" : "H. S.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Galetti", "given" : "M.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Ceballos", "given" : "G.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Isaac", "given" : "N. J. B.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Collen", "given" : "B.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Science", "id" : "ITEM-1", "issue" : "6195", "issued" : { "date-parts" : [ [ "2014" ] ] }, "page" : "401-406", "title" : "Defaunation in the Anthropocene", "type" : "article-journal", "volume" : "345" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1016/j.biocon.2013.04.020", "ISSN" : "00063207", "abstract" : "Defaunation, the loss or population decline of medium and large native vertebrates represents a significant threat to the biodiversity of tropical ecosystems. Here we review the anthropogenic drivers of defaunation, provide a brief historical account of the development of this field, and analyze the types of biological consequences of this impact on the structure and functioning of tropical ecosystems. We identify how defaunation, operating at a variety of scales, from the plot to the global level, affects biological systems along a gradient of processes ranging from plant physiology (vegetative and reproductive performance) and animal behavior (movement, foraging and dietary patterns) in the immediate term; to plant population and community dynamics and structure leading to disruptions of ecosystem functioning (and thus degrading environmental services) in the short to medium term; to evolutionary changes (phenotypic changes and population genetic structure) in the long-term. We present such a synthesis as a preamble to a series of papers that provide a compilation of our current understanding of the impact and consequences of tropical defaunation. We close by identifying some of the most urgent needs and perspectives that warrant further study to improve our understanding of this field, as we confront the challenges of living in a defaunated world.", "author" : [ { "dropping-particle" : "", "family" : "Galetti", "given" : "Mauro", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Dirzo", "given" : "Rodolfo", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Biological Conservation", "id" : "ITEM-2", "issued" : { "date-parts" : [ [ "2013", "7" ] ] }, "page" : "1-6", "title" : "Ecological and evolutionary consequences of living in a defaunated world", "type" : "article-journal", "volume" : "163" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Galetti & Dirzo 2013; Dirzo et al. 2014)", "plainTextFormattedCitation" : "(Galetti & Dirzo 2013; Dirzo et al. 2014)", "previouslyFormattedCitation" : "(Galetti & Dirzo 2013; Dirzo et al. 2014)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Galetti & Dirzo 2013; Dirzo et al. 2014), e.g., in relation to the impacts of past, current and future defaunation on ecosystem function and dynamics ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1371/journal.pone.0071352", "ISSN" : "19326203", "PMID" : "23951141", "abstract" : "Animals translocate nutrients by consuming nutrients at one point and excreting them or dying at another location. Such lateral fluxes may be an important mechanism of nutrient supply in many ecosystems, but lack quantification and a systematic theoretical framework for their evaluation. This paper presents a mathematical framework for quantifying such fluxes in the context of mammalian herbivores. We develop an expression for lateral diffusion of a nutrient, where the diffusivity is a biologically determined parameter depending on the characteristics of mammals occupying the domain, including size-dependent phenomena such as day range, metabolic demand, food passage time, and population size. Three findings stand out: (a) Scaling law-derived estimates of diffusion parameters are comparable to estimates calculated from estimates of each coefficient gathered from primary literature. (b) The diffusion term due to transport of nutrients in dung is orders of magnitude large than the coefficient representing nutrients in bodymass. (c) The scaling coefficients show that large herbivores make a disproportionate contribution to lateral nutrient transfer. We apply the diffusion equation to a case study of Kruger National Park to estimate the conditions under which mammal-driven nutrient transport is comparable in magnitude to other (abiotic) nutrient fluxes (inputs and losses). Finally, a global analysis of mammalian herbivore transport is presented, using a comprehensive database of contemporary animal distributions. We show that continents vary greatly in terms of the importance of animal-driven nutrient fluxes, and also that perturbations to nutrient cycles are potentially quite large if threatened large herbivores are driven to extinction.", "author" : [ { "dropping-particle" : "", "family" : "Wolf", "given" : "Adam", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Doughty", "given" : "Christopher E", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Malhi", "given" : "Yadvinder", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "PLoS ONE", "id" : "ITEM-1", "issue" : "8", "issued" : { "date-parts" : [ [ "2013", "1" ] ] }, "page" : "e71352", "title" : "Lateral diffusion of nutrients by mammalian herbivores in terrestrial ecosystems", "type" : "article-journal", "volume" : "8" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1038/ngeo1895", "ISBN" : "1752-0894", "ISSN" : "1752-0894", "abstract" : "n the late Pleistocene, 97 genera of large animals went extinct, concentrated in the Americas and Australia1. These extinctions had significant effects on ecosystem structure2, seed dispersal3 and land surface albedo4. However, the impact of this dramatic extinction on ecosystem nutrient biogeochemistry, through the lateral transport of dung and bodies, has never been explored. Here we analyse this process using a novel mathematical framework that analyses this lateral transport as a diffusion-like process, and we demonstrate that large animals play a disproportionately large role in the horizontal transfer of nutrients across landscapes. For example, we estimate that the extinction of the Amazonian megafauna decreased the lateral flux of the limiting nutrient phosphorus by more than 98%, with similar, though less extreme, decreases in all continents outside of Africa. This resulted in strong decreases in phosphorus availability in eastern Amazonia away from fertile floodplains, a decline which may still be ongoing. The current P limitation in the Amazon basin may be partially a relic of an ecosystem without the functional connectivity it once had. We argue that the Pleistocene megafauna extinctions resulted in large and ongoing disruptions to terrestrial biogeochemical cycling at continental scales and increased nutrient heterogeneity globally.", "author" : [ { "dropping-particle" : "", "family" : "Doughty", "given" : "Christopher E.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Wolf", "given" : "Adam", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Malhi", "given" : "Yadvinder", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Nature Geoscience", "id" : "ITEM-2", "issue" : "9", "issued" : { "date-parts" : [ [ "2013" ] ] }, "page" : "761-764", "publisher" : "Nature Publishing Group", "title" : "The legacy of the Pleistocene megafauna extinctions on nutrient availability in Amazonia", "type" : "article-journal", "volume" : "6" }, "uris" : [ "" ] }, { "id" : "ITEM-3", "itemData" : { "DOI" : "10.1086/508027", "ISSN" : "1537-5323", "PMID" : "17080364", "abstract" : "Large vertebrates are strong interactors in food webs, yet they were lost from most ecosystems after the dispersal of modern humans from Africa and Eurasia. We call for restoration of missing ecological functions and evolutionary potential of lost North American megafauna using extant conspecifics and related taxa. We refer to this restoration as Pleistocene rewilding; it is conceived as carefully managed ecosystem manipulations whereby costs and benefits are objectively addressed on a case-by-case and locality-by-locality basis. Pleistocene rewilding would deliberately promote large, long-lived species over pest and weed assemblages, facilitate the persistence and ecological effectiveness of megafauna on a global scale, and broaden the underlying premise of conservation from managing extinction to encompass restoring ecological and evolutionary processes. Pleistocene rewilding can begin immediately with species such as Bolson tortoises and feral horses and continue through the coming decades with elephants and Holarctic lions. Our exemplar taxa would contribute biological, economic, and cultural benefits to North America. Owners of large tracts of private land in the central and western United States could be the first to implement this restoration. Risks of Pleistocene rewilding include the possibility of altered disease ecology and associated human health implications, as well as unexpected ecological and sociopolitical consequences of reintroductions. Establishment of programs to monitor suites of species interactions and their consequences for biodiversity and ecosystem health will be a significant challenge. Secure fencing would be a major economic cost, and social challenges will include acceptance of predation as an overriding natural process and the incorporation of pre-Columbian ecological frameworks into conservation strategies.", "author" : [ { "dropping-particle" : "", "family" : "Donlan", "given" : "Josh", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Berger", "given" : "Joel", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Bock", "given" : "Carl E", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Bock", "given" : "Jane H", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Burney", "given" : "David a", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Estes", "given" : "James a", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Foreman", "given" : "Dave", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Martin", "given" : "Paul S", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Roemer", "given" : "Gary W", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Smith", "given" : "Felisa a", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Soul\u00e9", "given" : "Michael E", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Greene", "given" : "Harry W", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "The American naturalist", "id" : "ITEM-3", "issue" : "5", "issued" : { "date-parts" : [ [ "2006", "11" ] ] }, "page" : "660-81", "title" : "Pleistocene rewilding: an optimistic agenda for twenty-first century conservation.", "type" : "article-journal", "volume" : "168" }, "uris" : [ "" ] }, { "id" : "ITEM-4", "itemData" : { "DOI" : "10.1126/science.1241484", "ISBN" : "1095-9203 (Electronic) 0036-8075 (Linking)", "ISSN" : "0036-8075", "PMID" : "24408439", "abstract" : "Large carnivores face serious threats and are experiencing massive declines in their populations and geographic ranges around the world. We highlight how these threats have affected the conservation status and ecological functioning of the 31 largest mammalian carnivores on Earth. Consistent with theory, empirical studies increasingly show that large carnivores have substantial effects on the structure and function of diverse ecosystems. Significant cascading trophic interactions, mediated by their prey or sympatric mesopredators, arise when some of these carnivores are extirpated from or repatriated to ecosystems. Unexpected effects of trophic cascades on various taxa and processes include changes to bird, mammal, invertebrate, and herpetofauna abundance or richness; subsidies to scavengers; altered disease dynamics; carbon sequestration; modified stream morphology; and crop damage. Promoting tolerance and coexistence with large carnivores is a crucial societal challenge that will ultimately determine the fate of Earth's largest carnivores and all that depends upon them, including humans.", "author" : [ { "dropping-particle" : "", "family" : "Ripple", "given" : "William J", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Estes", "given" : "James a", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Beschta", "given" : "Robert L", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Wilmers", "given" : "Christopher C", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Ritchie", "given" : "Euan G", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Hebblewhite", "given" : 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}, "page" : "e1400103-e1400103", "title" : "Collapse of the world's largest herbivores", "type" : "article-journal", "volume" : "1" }, "uris" : [ "" ] }, { "id" : "ITEM-6", "itemData" : { "DOI" : "10.1098/rstb.2011.0020", "ISBN" : "1471-2970 (Electronic)\\r0962-8436 (Linking)", "ISSN" : "0962-8436", "PMID" : "21807737", "abstract" : "Although the recent historical period is usually treated as a temporal base-line for understanding patterns of mammal extinction, mammalian biodiversity loss has also taken place throughout the Late Quaternary. We explore the spatial, taxonomic and phylogenetic patterns of 241 mammal species extinctions known to have occurred during the Holocene up to the present day. To assess whether our understanding of mammalian threat processes has been affected by excluding these taxa, we incorporate extinct species data into analyses of the impact of body mass on extinction risk. We find that Holocene extinctions have been phylogenetically and spatially concentrated in specific taxa and geographical regions, which are often not congruent with those disproportionately at risk today. Large-bodied mammals have also been more extinction-prone in most geographical regions across the Holocene. Our data support the extinction filter hypothesis, whereby regional faunas from which susceptible species have already become extinct now appear less threatened; they may also suggest that different processes are responsible for driving past and present extinctions. We also find overall incompleteness and inter-regional biases in extinction data from the recent fossil record. Although direct use of fossil data in future projections of extinction risk is therefore not straightforward, insights into extinction processes from the Holocene record are still useful in understanding mammalian threat.", "author" : [ { "dropping-particle" : "", "family" : "Turvey", "given" : "Samuel T", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Fritz", "given" : "Susanne a", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Philosophical transactions of the Royal Society of London. Series B, Biological sciences", "id" : "ITEM-6", "issue" : "1577", "issued" : { "date-parts" : [ [ "2011" ] ] }, "page" : "2564-2576", "title" : "The ghosts of mammals past: biological and geographical patterns of global mammalian extinction across the Holocene.", "type" : "article-journal", "volume" : "366" }, "uris" : [ "" ] }, { "id" : "ITEM-7", "itemData" : { "DOI" : "10.1016/j.biocon.2013.04.020", "ISSN" : "00063207", "abstract" : "Defaunation, the loss or population decline of medium and large native vertebrates represents a significant threat to the biodiversity of tropical ecosystems. Here we review the anthropogenic drivers of defaunation, provide a brief historical account of the development of this field, and analyze the types of biological consequences of this impact on the structure and functioning of tropical ecosystems. We identify how defaunation, operating at a variety of scales, from the plot to the global level, affects biological systems along a gradient of processes ranging from plant physiology (vegetative and reproductive performance) and animal behavior (movement, foraging and dietary patterns) in the immediate term; to plant population and community dynamics and structure leading to disruptions of ecosystem functioning (and thus degrading environmental services) in the short to medium term; to evolutionary changes (phenotypic changes and population genetic structure) in the long-term. We present such a synthesis as a preamble to a series of papers that provide a compilation of our current understanding of the impact and consequences of tropical defaunation. We close by identifying some of the most urgent needs and perspectives that warrant further study to improve our understanding of this field, as we confront the challenges of living in a defaunated world.", "author" : [ { "dropping-particle" : "", "family" : "Galetti", "given" : "Mauro", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Dirzo", "given" : "Rodolfo", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Biological Conservation", "id" : "ITEM-7", "issued" : { "date-parts" : [ [ "2013", "7" ] ] }, "page" : "1-6", "title" : "Ecological and evolutionary consequences of living in a defaunated world", "type" : "article-journal", "volume" : "163" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Donlan et al. 2006; Turvey & Fritz 2011; Doughty, Wolf & Malhi 2013; Wolf, Doughty & Malhi 2013; Galetti & Dirzo 2013; Ripple et al. 2014, 2015)", "plainTextFormattedCitation" : "(Donlan et al. 2006; Turvey & Fritz 2011; Doughty, Wolf & Malhi 2013; Wolf, Doughty & Malhi 2013; Galetti & Dirzo 2013; Ripple et al. 2014, 2015)", "previouslyFormattedCitation" : "(Donlan et al. 2006; Turvey & Fritz 2011; Doughty, Wolf & Malhi 2013; Wolf, Doughty & Malhi 2013; Galetti & Dirzo 2013; Ripple et al. 2014, 2015)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Donlan et al. 2006; Turvey & Fritz 2011; Doughty, Wolf & Malhi 2013; Wolf, Doughty & Malhi 2013; Galetti & Dirzo 2013; Ripple et al. 2014, 2015). The SDR usually employed to predict density assumes a constant relationship across all clades, which we know not to be true; therefore, a model that includes this knowledge would be beneficial. It is increasingly clear that humans have had large impacts on mammal species diversity and local communities across the globe not just in recent times, but also during prehistory, especially for the megafauna ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1111/ddi.12369", "ISSN" : "13669516", "author" : [ { "dropping-particle" : "", "family" : "Faurby", "given" : "S\u00f8ren", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Svenning", "given" : "J.-C.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Diversity and Distributions", "editor" : [ { "dropping-particle" : "", "family" : "Stevens", "given" : "George", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "id" : "ITEM-1", "issue" : "10", "issued" : { "date-parts" : [ [ "2015", "10" ] ] }, "page" : "1155-1166", "title" : "Historic and prehistoric human-driven extinctions have reshaped global mammal diversity patterns", "type" : "article-journal", "volume" : "21" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1098/rspb.2013.3254", "ISSN" : "0962-8452", "author" : [ { "dropping-particle" : "", "family" : "Sandom", "given" : "C.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Faurby", "given" : "S.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Sandel", "given" : "B.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Svenning", "given" : "J.-C.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Proceedings of the Royal Society B: Biological Sciences", "id" : "ITEM-2", "issue" : "1787", "issued" : { "date-parts" : [ [ "2014", "6", "4" ] ] }, "page" : "20133254", "title" : "Global late Quaternary megafauna extinctions linked to humans, not climate change", "type" : "article-journal", "volume" : "281" }, "uris" : [ "" ] }, { "id" : "ITEM-3", "itemData" : { "DOI" : "10.1098/rstb.2011.0020", "ISBN" : "1471-2970 (Electronic)\\r0962-8436 (Linking)", "ISSN" : "0962-8436", "PMID" : "21807737", "abstract" : "Although the recent historical period is usually treated as a temporal base-line for understanding patterns of mammal extinction, mammalian biodiversity loss has also taken place throughout the Late Quaternary. 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We also find overall incompleteness and inter-regional biases in extinction data from the recent fossil record. Although direct use of fossil data in future projections of extinction risk is therefore not straightforward, insights into extinction processes from the Holocene record are still useful in understanding mammalian threat.", "author" : [ { "dropping-particle" : "", "family" : "Turvey", "given" : "Samuel T", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Fritz", "given" : "Susanne a", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Philosophical transactions of the Royal Society of London. 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With a better model for population densities we could supplement these studies with better estimates of potential population densities.Here, we re-assess the generality of -0.75 scaling rule for the size-density relation in mammals, incorporating phylogenetic relatedness in a new approach that allows a data-driven identification of phylogenetic substructure in the density-body size relation (cf. ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1890/13-0620.1", "ISBN" : "0012-9658", "ISSN" : "00129658", "PMID" : "24669745", "abstract" : "Ecological communities consist of small abundant and large non-abundant species. The energetic equivalence rule is an often-observed pattern that could be explained by equal energy usage among abundant small organisms and non-abundant large organisms. To generate this pattern, metabolism (as an indicator of individual energy use) and abundance have to scale inversely with body mass, and cancel each other out. In contrast, the pattern referred to as biomass equivalence states that the biomass of all species in an area should be constant across the body-mass range. In this study, we investigated forest soil communities with respect to metabolism, abundance, population energy use, and biomass. We focused on four land-use types in three different landscape blocks (Biodiversity Exploratories). The soil samples contained 870 species across 12 phylogenetic groups. Our results indicated positive sublinear metabolic scaling and negative sublinear abundance scaling with species body mass. The relationships varied mainly due to differences among phylogenetic groups or feeding types, and only marginally due to land-use type. However, these scaling relationships were not exactly inverse to each other, resulting in increasing population energy use and biomass with increasing body mass for most combinations of phylogenetic group or feeding type with land-use type. Thus, our results are mostly inconsistent with the classic perception of energetic equivalence, and reject the biomass equivalence hypothesis while documenting a specific and nonrandom pattern of how abundance, energy use, and biomass are distributed across size classes. However, these patterns are consistent with two alternative predictions: the resource-thinning hypothesis, which states that abundance decreases with trophic level, and the allometric degree hypothesis, which states that population energy use should increase with population average body mass, due to correlations with the number of links of consumers and resources. Overall, our results suggest that a synthesis of food web structures with metabolic theory may be most promising for predicting natural patterns of abundance, biomass, and energy use.", "author" : [ { "dropping-particle" : "", "family" : "Ehnes", "given" : "Roswitha B.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Pollierer", "given" : "Melanie M.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Erdmann", "given" : "Georgia", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Klarner", "given" : "Bernhard", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Eitzinger", "given" : "Bernhard", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Digel", "given" : "Christoph", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Ott", "given" : "David", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Maraun", "given" : "Mark", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Scheu", "given" : "Stefan", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Brose", "given" : "Ulrich", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecology", "id" : "ITEM-1", "issue" : "2", "issued" : { "date-parts" : [ [ "2014" ] ] }, "page" : "527-537", "title" : "Lack of energetic equivalence in forest soil invertebrates", "type" : "article-journal", "volume" : "95" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Ehnes et al. 2014)", "manualFormatting" : "Ehnes et al. 2014)", "plainTextFormattedCitation" : "(Ehnes et al. 2014)", "previouslyFormattedCitation" : "(Ehnes et al. 2014)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }Ehnes et al. 2014). By doing this we indirectly investigate phylogenetically structured traits that may cause groups to deviate from an overall trend. Other studies have done this by including a priori known traits such as diet ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1371/journal.pone.0071352", "ISSN" : "19326203", "PMID" : "23951141", "abstract" : "Animals translocate nutrients by consuming nutrients at one point and excreting them or dying at another location. Such lateral fluxes may be an important mechanism of nutrient supply in many ecosystems, but lack quantification and a systematic theoretical framework for their evaluation. This paper presents a mathematical framework for quantifying such fluxes in the context of mammalian herbivores. We develop an expression for lateral diffusion of a nutrient, where the diffusivity is a biologically determined parameter depending on the characteristics of mammals occupying the domain, including size-dependent phenomena such as day range, metabolic demand, food passage time, and population size. Three findings stand out: (a) Scaling law-derived estimates of diffusion parameters are comparable to estimates calculated from estimates of each coefficient gathered from primary literature. (b) The diffusion term due to transport of nutrients in dung is orders of magnitude large than the coefficient representing nutrients in bodymass. (c) The scaling coefficients show that large herbivores make a disproportionate contribution to lateral nutrient transfer. We apply the diffusion equation to a case study of Kruger National Park to estimate the conditions under which mammal-driven nutrient transport is comparable in magnitude to other (abiotic) nutrient fluxes (inputs and losses). Finally, a global analysis of mammalian herbivore transport is presented, using a comprehensive database of contemporary animal distributions. We show that continents vary greatly in terms of the importance of animal-driven nutrient fluxes, and also that perturbations to nutrient cycles are potentially quite large if threatened large herbivores are driven to extinction.", "author" : [ { "dropping-particle" : "", "family" : "Wolf", "given" : "Adam", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Doughty", "given" : "Christopher E", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Malhi", "given" : "Yadvinder", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "PLoS ONE", "id" : "ITEM-1", "issue" : "8", "issued" : { "date-parts" : [ [ "2013", "1" ] ] }, "page" : "e71352", "title" : "Lateral diffusion of nutrients by mammalian herbivores in terrestrial ecosystems", "type" : "article-journal", "volume" : "8" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1111/j.1095-8312.1987.tb01990.x", "ISSN" : "00244066", "author" : [ { "dropping-particle" : "", "family" : "Damuth", "given" : "John", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Biological Journal of the Linnean Society", "id" : "ITEM-2", "issue" : "3", "issued" : { "date-parts" : [ [ "1987", "7", "28" ] ] }, "page" : "193-246", "title" : "Interspecific allometry of population density in mammals and other animals: the independence of body mass and population energy-use", "type" : "article-journal", "volume" : "31" }, "uris" : [ "" ] }, { "id" : "ITEM-3", "itemData" : { "DOI" : "10.1890/13-0620.1", "ISBN" : "0012-9658", "ISSN" : "00129658", "PMID" : "24669745", "abstract" : "Ecological communities consist of small abundant and large non-abundant species. The energetic equivalence rule is an often-observed pattern that could be explained by equal energy usage among abundant small organisms and non-abundant large organisms. To generate this pattern, metabolism (as an indicator of individual energy use) and abundance have to scale inversely with body mass, and cancel each other out. In contrast, the pattern referred to as biomass equivalence states that the biomass of all species in an area should be constant across the body-mass range. In this study, we investigated forest soil communities with respect to metabolism, abundance, population energy use, and biomass. We focused on four land-use types in three different landscape blocks (Biodiversity Exploratories). The soil samples contained 870 species across 12 phylogenetic groups. Our results indicated positive sublinear metabolic scaling and negative sublinear abundance scaling with species body mass. The relationships varied mainly due to differences among phylogenetic groups or feeding types, and only marginally due to land-use type. However, these scaling relationships were not exactly inverse to each other, resulting in increasing population energy use and biomass with increasing body mass for most combinations of phylogenetic group or feeding type with land-use type. Thus, our results are mostly inconsistent with the classic perception of energetic equivalence, and reject the biomass equivalence hypothesis while documenting a specific and nonrandom pattern of how abundance, energy use, and biomass are distributed across size classes. However, these patterns are consistent with two alternative predictions: the resource-thinning hypothesis, which states that abundance decreases with trophic level, and the allometric degree hypothesis, which states that population energy use should increase with population average body mass, due to correlations with the number of links of consumers and resources. Overall, our results suggest that a synthesis of food web structures with metabolic theory may be most promising for predicting natural patterns of abundance, biomass, and energy use.", "author" : [ { "dropping-particle" : "", "family" : "Ehnes", "given" : "Roswitha B.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Pollierer", "given" : "Melanie M.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Erdmann", "given" : "Georgia", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Klarner", "given" : "Bernhard", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Eitzinger", "given" : "Bernhard", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Digel", "given" : "Christoph", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Ott", "given" : "David", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Maraun", "given" : "Mark", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Scheu", "given" : "Stefan", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Brose", "given" : "Ulrich", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecology", "id" : "ITEM-3", "issue" : "2", "issued" : { "date-parts" : [ [ "2014" ] ] }, "page" : "527-537", "title" : "Lack of energetic equivalence in forest soil invertebrates", "type" : "article-journal", "volume" : "95" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Damuth 1987; Wolf et al. 2013; Ehnes et al. 2014)", "plainTextFormattedCitation" : "(Damuth 1987; Wolf et al. 2013; Ehnes et al. 2014)", "previouslyFormattedCitation" : "(Damuth 1987; Wolf et al. 2013; Ehnes et al. 2014)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Damuth 1987; Wolf et al. 2013; Ehnes et al. 2014), where e.g. carnivores tend to have steeper slopes and lower intercepts than herbivores. 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In contrast to this approach, we here fit a model on phylogeny alone with an iterative framework searching for groups with distinct slopes, without any a priori assumption on which traits might distinguish them from one another. 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Overall, we expect our approach to provide a more accurate model of population densities of species for which we know little of their ecology, and sometimes merely their taxonomic placement and body mass.In this study we assess the following specific study hypotheses for the size-density relation in mammals: While we expect the relation to hold up across all mammals, we hypothesize that there will be a phylogenetic substructure, where groups of more closely related species will exhibit a shallower decline in density with body mass, consistent with a disproportionate capture of resources by the larger species within groups of ecologically similar species. Further, we expect that carnivorous groups scale the steepest and are offset to overall lower population densities than all other clades due to their high trophic level. Materials and methodsDataFirst, we created a dataset where we assigned each family of mammals to all the monophyletic clades it is part of. For example, the family Felidae belongs to the suborder Feliformia, the order Carnivora, the cohort Placentalia, and in the end the class Mammalia, as well as a number of monophyletic unnamed clades in between. By using a phylogeny of all mammalian families we could assign each family to all clades it belongs to by all dichotomies above it in the tree. The phylogeny used is by ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1126/science.1211028", "ISSN" : "0036-8075", "PMID" : "21940861", "abstract" : "Previous analyses of relations, divergence times, and diversification patterns among extant mammalian families have relied on supertree methods and local molecular clocks. We constructed a molecular supermatrix for mammalian families and analyzed these data with likelihood-based methods and relaxed molecular clocks. Phylogenetic analyses resulted in a robust phylogeny with better resolution than phylogenies from supertree methods. Relaxed clock analyses support the long-fuse model of diversification and highlight the importance of including multiple fossil calibrations that are spread across the tree. Molecular time trees and diversification analyses suggest important roles for the Cretaceous Terrestrial Revolution and Cretaceous-Paleogene (KPg) mass extinction in opening up ecospace that promoted interordinal and intraordinal diversification, respectively. By contrast, diversification analyses provide no support for the hypothesis concerning the delayed rise of present-day mammals during the Eocene Period.", "author" : [ { "dropping-particle" : "", "family" : "Meredith", "given" : "Robert W", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Janecka", "given" : "J. 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However, primate taxonomy is both complex and controversial, with marginal unifying consensus of the evolutionary hierarchy of extant primate species. Here we provide new genomic sequence (~8 Mb) from 186 primates representing 61 (~90%) of the described genera, and we include outgroup species from Dermoptera, Scandentia, and Lagomorpha. The resultant phylogeny is exceptionally robust and illuminates events in primate evolution from ancient to recent, clarifying numerous taxonomic controversies and providing new data on human evolution. Ongoing speciation, reticulate evolution, ancient relic lineages, unequal rates of evolution, and disparate distributions of insertions/deletions among the reconstructed primate lineages are uncovered. Our resolution of the primate phylogeny provides an essential evolutionary framework with far-reaching applications including: human selection and adaptation, global emergence of zoonotic diseases, mammalian comparative genomics, primate taxonomy, and conservation of endangered species.", "author" : [ { "dropping-particle" : "", "family" : "Perelman", "given" : "Polina", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Johnson", "given" : "Warren E", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Roos", "given" : "Christian", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Seu\u00e1nez", "given" : "Hector N", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Horvath", "given" : "Julie E", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Moreira", "given" : "Miguel a M", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Kessing", "given" : "Bailey", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Pontius", "given" : "Joan", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Roelke", "given" : "Melody", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Rumpler", "given" : "Yves", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Schneider", "given" : "Maria Paula C", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Silva", "given" : "Artur", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "O'Brien", "given" : "Stephen J", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Pecon-Slattery", "given" : "Jill", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "PLoS genetics", "id" : "ITEM-1", "issue" : "3", "issued" : { "date-parts" : [ [ "2011", "3" ] ] }, "page" : "e1001342", "title" : "A molecular phylogeny of living primates.", "type" : "article-journal", "volume" : "7" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Perelman et al. 2011)", "manualFormatting" : "Perelman et al. (2011)", "plainTextFormattedCitation" : "(Perelman et al. 2011)", "previouslyFormattedCitation" : "(Perelman et al. 2011)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }Perelman et al. (2011). By combining the datasets we end up with a dataset that provides body mass, population density, and a number of binomial variables indicating which monophyletic clades each species belongs to. We excluded all non-terrestrial species (Orders: Cetacea and Sirenia; Families: Odobenidae, Otariidae, and Phocidae; Species: Lontra felina, Enhydra lutris, and Ursus maritimus) and bats (Order: Chiroptera) from our dataset. To avoid overfitting we did not include monophyletic clades of less than 10 species, since standard GLM approaches recommend not fitting factor levels with less than 10 data points ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "ISBN" : "978-0-470-08186-0", "abstract" : "\"Over the years, I have had the opportunity to teach several regression courses, and I cannot think of a better undergraduate text than this one.\" (The American Statistician) \"The book is well written and has many exercises. It can serve as a very good textbook for scientists and engineers, with only basic statistics as a prerequisite. I also highly recommend it to practitioners who want to solve real-life prediction problems.\" (Computing Reviews) Modern Regression Methods, Second Edition maintains the accessible organization, breadth of coverage, and cutting-edge appeal that earned its predecessor the title of being one of the top five books for statisticians by an Amstat News book editor in 2003. This new edition has been updated and enhanced to include all-new information on the latest advances and research in the evolving field of regression analysis. The book provides a unique treatment of fundamental regression methods, such as diagnostics, transformations, robust regression, and ridge regression. Unifying key concepts and procedures, this new edition emphasizes applications to provide a more hands-on and comprehensive understanding of regression diagnostics. New features of the Second Edition include: A revised chapter on logistic regression, including improved methods of parameter estimation A new chapter focusing on additional topics of study in regression, including quantile regression, semiparametric regression, and Poisson regression A wealth of new and updated exercises with worked solutions An extensive FTP site complete with Minitab macros, which allow the reader to compute analyses, and specialized procedures Updated references at the end of each chapter that direct the reader to the appropriate resources for further study An accessible guide to state-of-the-art regression techniques, Modern Regression Methods, Second Edition is an excellent book for courses in regression analysis at the upper-undergraduate and graduate levels. It is also a valuable reference for practicing statisticians, engineers, and physical scientists.", "author" : [ { "dropping-particle" : "", "family" : "Ryan", "given" : "Thomas P.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "edition" : "2", "id" : "ITEM-1", "issued" : { "date-parts" : [ [ "2008" ] ] }, "number-of-pages" : "672", "publisher" : "John Wiley & Sons", "publisher-place" : "New York", "title" : "Modern Regression Methods", "type" : "book" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Ryan 2008)", "plainTextFormattedCitation" : "(Ryan 2008)", "previouslyFormattedCitation" : "(Ryan 2008)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Ryan 2008). Our final dataset included 924 species belonging to 110 distinct monophyletic groups.AnalysisFirst, we fit the simple allometric relation, log10-population density as a function of log10-body mass. Then, we iterated a model building procedure until we found no significant improvement. The iterations were stopped when there was no further improvement of the model of a ΔAICc of more than 4 ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "ISBN" : "1441929738", "abstract" : "This book is unique in that it covers the philosophy of model-based data analysis and an omnibus strategy for the analysis of empirical data. The book introduces information theoretic approaches and focuses critical attention on a priori modeling and the selection of a good approximating model that best represents the inference supported by the data. Kullback-Leibler information represents a fundamental quantity in science and is Hirotugu Akaike's basis for model selection. The maximized log-likelihood function can be bias-corrected to provide an estimate of expected, relative Kullback-Leibler information. This leads to Akaike's Information Criterion (AIC) and various extensions and these are relatively simple and easy to use in practice, but little taught in statistics classes and far less understood in the applied sciences than should be the case. The information theoretic approaches provide a unified and rigorous theory, an extension of likelihood theory, an important application of information theory, and are objective and practical to employ across a very wide class of empirical problems. Parameter estimation has long been viewed as an optimization problem (e.g., maximize the log-likelihood or minimize the residual sum of squared deviations) and under the information theoretic paradigm, data-based model selection is also an optimization problem. This brings model selection and parameter estimation under a common framework - optimization. The value of AIC is computed for each a priori model to be considered and the model with the minimum AIC is used for statistical inference. However, the paradigm described in this book goes beyond merely the computation and interpretation of AIC to select a parsimonious model for inference from empirical data; it refocuses increased attention on a variety of considerations and modeling prior to the actual analysis of data.", "author" : [ { "dropping-particle" : "", "family" : "Burnham", "given" : "K P", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Anderson", "given" : "D R", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "New York Springer", "edition" : "2", "id" : "ITEM-1", "issued" : { "date-parts" : [ [ "2002" ] ] }, "number-of-pages" : "488", "publisher" : "Springer-Verlag", "publisher-place" : "New York", "title" : "Model selection and multimodel inference: a practical information-theoretic approach", "type" : "book", "volume" : "60" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Burnham & Anderson 2002)", "plainTextFormattedCitation" : "(Burnham & Anderson 2002)", "previouslyFormattedCitation" : "(Burnham & Anderson 2002)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Burnham & Anderson 2002). Each iteration consisted of adding a unique slope or intercept terms for all members of a specific clade by including the clade as a binomial factor in the model either as a main effect (intercept change) or as an interaction term (slope change). We fitted these as individual models for all clades and kept the model giving the highest improvement based on ΔAICc (Fig. 1) as a new base. We then removed any terms that were no longer leading to improvements of ΔAICc of more than 4. The process was repeated with a new iteration expanding the model. The cut-off value of 4 was chosen as this value has been used previously in the macro-evolutionary program MEDUSA ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1073/pnas.0811087106", "ISSN" : "1091-6490", "PMID" : "19633192", "abstract" : "The uneven distribution of species richness is a fundamental and unexplained pattern of vertebrate biodiversity. Although species richness in groups like mammals, birds, or teleost fishes is often attributed to accelerated cladogenesis, we lack a quantitative conceptual framework for identifying and comparing the exceptional changes of tempo in vertebrate evolutionary history. We develop MEDUSA, a stepwise approach based upon the Akaike information criterion for detecting multiple shifts in birth and death rates on an incompletely resolved phylogeny. We apply MEDUSA incompletely to a diversity tree summarizing both evolutionary relationships and species richness of 44 major clades of jawed vertebrates. We identify 9 major changes in the tempo of gnathostome diversification; the most significant of these lies at the base of a clade that includes most of the coral-reef associated fishes as well as cichlids and perches. Rate increases also underlie several well recognized tetrapod radiations, including most modern birds, lizards and snakes, ostariophysan fishes, and most eutherian mammals. In addition, we find that large sections of the vertebrate tree exhibit nearly equal rates of origination and extinction, providing some of the first evidence from molecular data for the importance of faunal turnover in shaping biodiversity. Together, these results reveal living vertebrate biodiversity to be the product of volatile turnover punctuated by 6 accelerations responsible for >85% of all species as well as 3 slowdowns that have produced \"living fossils.\" In addition, by revealing the timing of the exceptional pulses of vertebrate diversification as well as the clades that experience them, our diversity tree provides a framework for evaluating particular causal hypotheses of vertebrate radiations.", "author" : [ { "dropping-particle" : "", "family" : "Alfaro", "given" : "Michael E", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Santini", "given" : "Francesco", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Brock", "given" : "Chad", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Alamillo", "given" : "Hugo", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Dornburg", "given" : "Alex", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Rabosky", "given" : "Daniel L", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Carnevale", "given" : "Giorgio", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Harmon", "given" : "Luke J", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Proceedings of the National Academy of Sciences of the United States of America", "id" : "ITEM-1", "issue" : "32", "issued" : { "date-parts" : [ [ "2009", "8", "11" ] ] }, "page" : "13410-4", "title" : "Nine exceptional radiations plus high turnover explain species diversity in jawed vertebrates.", "type" : "article-journal", "volume" : "106" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Alfaro et al. 2009)", "plainTextFormattedCitation" : "(Alfaro et al. 2009)", "previouslyFormattedCitation" : "(Alfaro et al. 2009)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Alfaro et al. 2009), which iteratively identifies monophyletic clades that behave differently from the remaining clades, these potentially being paraphyletic assemblages. Further, models that have a ΔAICc > 4 are generally considered to have considerably less support than the lower scoring models ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "ISBN" : "1441929738", "abstract" : "This book is unique in that it covers the philosophy of model-based data analysis and an omnibus strategy for the analysis of empirical data. The book introduces information theoretic approaches and focuses critical attention on a priori modeling and the selection of a good approximating model that best represents the inference supported by the data. Kullback-Leibler information represents a fundamental quantity in science and is Hirotugu Akaike's basis for model selection. The maximized log-likelihood function can be bias-corrected to provide an estimate of expected, relative Kullback-Leibler information. This leads to Akaike's Information Criterion (AIC) and various extensions and these are relatively simple and easy to use in practice, but little taught in statistics classes and far less understood in the applied sciences than should be the case. The information theoretic approaches provide a unified and rigorous theory, an extension of likelihood theory, an important application of information theory, and are objective and practical to employ across a very wide class of empirical problems. Parameter estimation has long been viewed as an optimization problem (e.g., maximize the log-likelihood or minimize the residual sum of squared deviations) and under the information theoretic paradigm, data-based model selection is also an optimization problem. This brings model selection and parameter estimation under a common framework - optimization. The value of AIC is computed for each a priori model to be considered and the model with the minimum AIC is used for statistical inference. 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An underlying assumption explicitly fitting clades alone is that shifts in the hypothesised size-density relationship are instant between clades, which may not always be true. Many extant clades have, however, exhibited point-like changes in traits, such as carnivory, flight, and digestion ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1126/science.1102417", "ISBN" : "0036-8075", "ISSN" : "0036-8075", "PMID" : "15459388", "abstract" : "Over the past 50 million years, successive clades of large carnivorous mammals diversified and then declined to extinction. In most instances, the cause of the decline remains a puzzle. Here we argue that energetic constraints and pervasive selection for larger size (Cope's rule) in carnivores lead to dietary specialization (hypercarnivory) and increased vulnerability to extinction. In two major clades of extinct North American canids, the evolution of large size was associated with a dietary shift to hypercarnivory and a decline in species durations. Thus, selection for attributes that promoted individual success resulted in progressive evolutionary failure of their clades.", "author" : [ { "dropping-particle" : "", "family" : "Valkenburgh", "given" : "Blaire", "non-dropping-particle" : "Van", "parse-names" : false, "suffix" : "" } ], "container-title" : "Science", "id" : "ITEM-1", "issue" : "5693", "issued" : { "date-parts" : [ [ "2004", "10", "1" ] ] }, "page" : "101-104", "title" : "Cope's Rule, Hypercarnivory, and Extinction in North American Canids", "type" : "article-journal", "volume" : "306" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "ISBN" : "9780198508236", "editor" : [ { "dropping-particle" : "", "family" : "Macdonald", "given" : "D W", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Norris", "given" : "S", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "id" : "ITEM-2", "issued" : { "date-parts" : [ [ "2001" ] ] }, "number-of-pages" : "961", "publisher" : "Oxford University Press", "publisher-place" : "Oxford", "title" : "The New Encyclopedia of Mammals", "type" : "book" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Macdonald & Norris 2001; Van Valkenburgh 2004)", "plainTextFormattedCitation" : "(Macdonald & Norris 2001; Van Valkenburgh 2004)", "previouslyFormattedCitation" : "(Macdonald & Norris 2001; Van Valkenburgh 2004)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Macdonald & Norris 2001; Van Valkenburgh 2004).To validate our results we used leave-half-out cross-validation, repeated 1000 times. In other words we fitted parameter estimates for both our final model and the base model on a random selection of half the dataset, calculated r2 using linear regression between predicted and the remaining data, and estimated ΔAICc between the two models. This procedure was repeated with 1000 random sample permutations. This method does not validate our final model against all possible models, but it does validate its stability and performance against the traditional model (Table S5).The estimated SDR could be affected by body mass-specific biases of the density estimates. A bias such as this could arise either because there is a bias in how affected the actual density is by human influence, or if there is a bias in how the density is estimated based on body mass. We assume that any bias in the data linked to human influence of the actual density would be bias towards underestimates of natural population densities of large species, since extinctions linked to humans have a mass bias in that direction ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1086/686268", "ISSN" : "0003-0147", "PMID" : "27172600", "abstract" : "Islands are or have been occupied by unusual species, such as dwarf proboscideans and giant rodents. The discussion of the classical but controversial island rule\u2014which states that mammalian body sizes converge on intermediate sizes on islands\u2014has been stimulated by these unusual species. In this study, we use an unprecedented global data set of the distributions and body sizes of late Quaternary mammal species and a novel analytical method to analyze body size evolution on islands. The analyses produced strong support for the island rule. Islands have suffered massive human-driven losses of species, and we found that the support for the island rule was substantially stronger when the many late Quaternary extinct species were also considered (particularly the tendency for dwarfing in large taxa). The decisive support for the island rule in this study confirms that evolution plays out in a markedly different way on islands and that human impact may obscure even fundamental evolutionary patterns.", "author" : [ { "dropping-particle" : "", "family" : "Faurby", "given" : "S\u00f8ren", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Svenning", "given" : "Jens-Christian Jens\u2010Christian", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "The American Naturalist", "id" : "ITEM-1", "issue" : "6", "issued" : { "date-parts" : [ [ "2016", "6", "27" ] ] }, "page" : "812-820", "title" : "Resurrection of the Island Rule: Human-Driven Extinctions Have Obscured a Basic Evolutionary Pattern", "type" : "article-journal", "volume" : "187" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1111/ddi.12369", "ISSN" : "13669516", "author" : [ { "dropping-particle" : "", "family" : "Faurby", "given" : "S\u00f8ren", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Svenning", "given" : "J.-C.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Diversity and Distributions", "editor" : [ { "dropping-particle" : "", "family" : "Stevens", "given" : "George", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "id" : "ITEM-2", "issue" : "10", "issued" : { "date-parts" : [ [ "2015", "10" ] ] }, "page" : "1155-1166", "title" : "Historic and prehistoric human-driven extinctions have reshaped global mammal diversity patterns", "type" : "article-journal", "volume" : "21" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Faurby & Svenning 2015b, 2016)", "plainTextFormattedCitation" : "(Faurby & Svenning 2015b, 2016)", "previouslyFormattedCitation" : "(Faurby & Svenning 2015b, 2016)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Faurby & Svenning 2015b, 2016). Further, we analysed a dataset containing information on both densities and sampled area and found a weak, but statistically significant trend towards underestimations of the densities for larger-bodied species as a consequence of them generally being measured in a larger area (Appendix S1). From this, we conclude that if there was a bias in the population density estimates it would be underestimations of larger-bodied species. If this bias existed in the data, it should have made the estimated slopes steeper. However, they were in reality shallower than expected, making conclusions based on the shallow slopes robust. For all data handling, graphics and statistical analysis we used R v. 3.2.3 ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "URL" : "", "author" : [ { "dropping-particle" : "", "family" : "R Core Team", "given" : "", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "id" : "ITEM-1", "issued" : { "date-parts" : [ [ "2015" ] ] }, "publisher-place" : "Vienna, Austria", "title" : "R: A language and environment for statistical computing", "type" : "webpage" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(R Core Team 2015)", "plainTextFormattedCitation" : "(R Core Team 2015)", "previouslyFormattedCitation" : "(R Core Team 2015)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(R Core Team 2015). For working with the phylogenetic data we used the packages ‘ape’ v. 3.4 ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "author" : [ { "dropping-particle" : "", "family" : "Paradis", "given" : "E", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Claude", "given" : "J", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Strimmer", "given" : "K", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Bioinformatics", "id" : "ITEM-1", "issued" : { "date-parts" : [ [ "2004" ] ] }, "page" : "289-290", "title" : "A{PE}: analyses of phylogenetics and evolution in {R} language", "type" : "article-journal", "volume" : "20" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Paradis, Claude & Strimmer 2004)", "plainTextFormattedCitation" : "(Paradis, Claude & Strimmer 2004)", "previouslyFormattedCitation" : "(Paradis, Claude & Strimmer 2004)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Paradis, Claude & Strimmer 2004), ‘phytools’ v. 0.5.10 ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "author" : [ { "dropping-particle" : "", "family" : "Revell", "given" : "Liam J", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Methods in Ecology and Evolution", "id" : "ITEM-1", "issued" : { "date-parts" : [ [ "2012" ] ] }, "page" : "217-223", "title" : "phytools: An R package for phylogenetic comparative biology (and other things).", "type" : "article-journal", "volume" : "3" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Revell 2012)", "plainTextFormattedCitation" : "(Revell 2012)", "previouslyFormattedCitation" : "(Revell 2012)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Revell 2012) and ‘geiger’ v. 2.0.6 ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "author" : [ { "dropping-particle" : "", "family" : "Harmon", "given" : "L J", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Weir", "given" : "J T", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Brock", "given" : "C D", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Glor", "given" : "R E", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Challenger", "given" : "W", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Bioinformatics", "id" : "ITEM-1", "issued" : { "date-parts" : [ [ "2008" ] ] }, "page" : "129-131", "title" : "GEIGER: investigating evolutionary radiations", "type" : "article-journal", "volume" : "24" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Harmon et al. 2008)", "plainTextFormattedCitation" : "(Harmon et al. 2008)", "previouslyFormattedCitation" : "(Harmon et al. 2008)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Harmon et al. 2008). For statistical analysis we used methods inspired by MEDUSA ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1073/pnas.0811087106", "ISSN" : "1091-6490", "PMID" : "19633192", "abstract" : "The uneven distribution of species richness is a fundamental and unexplained pattern of vertebrate biodiversity. Although species richness in groups like mammals, birds, or teleost fishes is often attributed to accelerated cladogenesis, we lack a quantitative conceptual framework for identifying and comparing the exceptional changes of tempo in vertebrate evolutionary history. We develop MEDUSA, a stepwise approach based upon the Akaike information criterion for detecting multiple shifts in birth and death rates on an incompletely resolved phylogeny. We apply MEDUSA incompletely to a diversity tree summarizing both evolutionary relationships and species richness of 44 major clades of jawed vertebrates. 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Graphics were made using ‘ggplot2’ v. 2.0.0 ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "ISBN" : "978-0-387-98140-6", "author" : [ { "dropping-particle" : "", "family" : "Wickham", "given" : "Hadley", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "id" : "ITEM-1", "issued" : { "date-parts" : [ [ "2009" ] ] }, "publisher" : "Springer New York", "publisher-place" : "New York", "title" : "ggplot2: elegant graphics for data analysis", "type" : "book" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Wickham 2009)", "plainTextFormattedCitation" : "(Wickham 2009)", "previouslyFormattedCitation" : "(Wickham 2009)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Wickham 2009).ResultsAcross a wide sample of all mammal species spanning six orders magnitude of body mass, we found that the overall relationship of log10-body mass to log10-density had a slope of b = -0.74 (SE = 0.021, p < 2 × 10-16), no different from the null expectation -0.75 (Fig. 2).Within mammalian clades, population density did not conform to a single overall trend, however. Rather the relationship between population densities and species body mass changed several times across the phylogeny (Table 1, Fig. 2-3), but all coefficients were less negative than the overall trend.The simple model prediction with a uniform relationship across all mammals performed substantially worse than a best-fit phylogenetic model improving r2 from 0.56 to 0.74 with ΔAICc = 466. Virtually identical improvement in the fit was found when comparing the original and the full phylogenetic model in our leave half out cross validation (Table S5).Our results showed that taxonomic groups with relatively heavier body masses have steeper slopes than lighter-bodied groups in a linear model of group slope as a function of mean log10 body mass (r = -0.31, p = 0.0011). The body mass range of a group had no effect on slope in a linear model of group slope as a function of range width of log10 body mass (r = -0.087, p = 0.17). Within the main carnivorous mammal clade (Carnivora) we saw slopes ranging from one of the steepest to some of the weakest (-0.36 to -0.71). Therefore, we saw no clear simple overall trophic explanation for differences in the SDR. Still, we did find the steepest slope of all the mammal clades for the most purely carnivorous subgroup of Carnivora, Felidae and Viverridae, while the Carnivora clades with the weakest slopes Ailuridae, Mephitidae, and Procyonidae have much more mixed diets, including many omnivorous or even herbivorous species ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "ISBN" : "978-84-96553-49-1", "editor" : [ { "dropping-particle" : "", "family" : "Wilson", "given" : "Don E.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Mittermeier", "given" : "Russell A.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "id" : "ITEM-1", "issued" : { "date-parts" : [ [ "2009" ] ] }, "number-of-pages" : "728", "publisher" : "Lynx Edicions", "title" : "Handbook of the Mammals of the World - Volume 1 Carnivores", "type" : "book" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Wilson & Mittermeier 2009)", "plainTextFormattedCitation" : "(Wilson & Mittermeier 2009)", "previouslyFormattedCitation" : "(Wilson & Mittermeier 2009)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Wilson & Mittermeier 2009).Within the Carnivora clades, we saw a general downward shift in intercepts, resulting in lower population density estimates for this group than according to the classical model. Within the respective mass spans of the four different Carnivora groups, we saw that the predicted densities for Mephitidae and Procyonidae were 2-5 times than what they were modelled with the same model for all species (i.e., the null model with a slope of-0.74), Mustelidae had predicted densities that are 16-130 lower, Felidae and Viverridae have 44-49 times lower predicted densities while the rest of the Carnivora (Canidae, Eupleridae, Herpestidae, Hyaenidae, Nandiniidae, and Ursidae) have predicted densities 8-21 times lower than predicted by the classical model.Table 1: Allometric model fit for species’ population density vs. average adult body mass for terrestrial mammals. This table shows the resulting SDR for families, orders or lager well defined monophyletic groups for an interpretable overview of our results. The relationships shown here are based on the 10 distinct slopes and 6 distinct intercepts we found strongly supported, based on ΔAICc > 4, see Table S2 for full model and AICc levels. An overall fit of all mammals without clade-specific variation led to an intercept of 3.87 and slope of -0.74.CladeInterceptSlopeMonotremata*3.15-0.58Marsupialia??Didelphimorphia3.15-0.58Paucituberculata*3.15-0.58Dasyuromorphia3.15-0.58Diprotodontia3.15-0.38Microbiotheria*3.15-0.58Notoryctemorphia*3.15-0.58Peramelemorphia*3.15-0.58Afrotheria3.15-0.33Xenarthra3.15-0.58Euarchontoglires??Dermoptera*3.15-0.58Lagomorpha: Leporidae3.53-0.58Lagomorpha: Ochotonidae3.53-0.07Primates: Galagidae, Hominidae, Hylobatidae, Lorisidae, Tarsiidae, & Platyrrhini3.15-0.58Primates: Cercopithecidae3.15-0.49Primates: Daubentoniidae & Lemuriformes3.74-0.58Rodentia: Castoridae*2.13-0.58Rodentia: Geomyidae & Heteromyidae2.130.44Rodentia: Dipodidae* & Anomaluromorpha*3.15-0.58Rodentia: Myomorpha3.15-0.18Rodentia: Hystricomorpha & Sciuromorpha3.84-0.58Scandentia*3.15-0.58Laurasiatheria??Artiodactyla3.15-0.58Carnivora: Mustelidae1.05-0.36Carnivora: Felidae & Viverridae2.13-0.71Carnivora: Canidae, Eupleridae, Herpestidae, Hyaenidae, Nandiniidae, & Ursidae2.13-0.58Carnivora: Ailuridae, Mephitidae, & Procyonidae2.13-0.36Erinaceomorpha*3.15-0.58Perissodactyla3.15-0.58Pholidota*3.15-0.58Soricomorpha: Solenodontidae & Soricidae3.15-0.58Soricomorpha: Talpidae3.150.04Marked (*) groups where we only had density estimates from less than 10 species and unique slopes or intercept for the group therefore were not allowed. Intercepts and slopes marked with bold are from groups with distinct values different from the paraphyletic assemblage containing the remaining species. The value for the paraphyletic assemblage is repeated multiple times for monophyletic subparts to make the values for individual clades easier recoverable from the table. DiscussionWe found the global size–density relationship across all larger mammalian to be consistent with previous findings ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1111/j.1095-8312.1987.tb01990.x", "ISSN" : "00244066", "author" : [ { "dropping-particle" : "", "family" : "Damuth", "given" : "John", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Biological Journal of the Linnean Society", "id" : "ITEM-1", "issue" : "3", "issued" : { "date-parts" : [ [ "1987", "7", "28" ] ] }, "page" : "193-246", "title" : "Interspecific allometry of population density in mammals and other animals: the independence of body mass and population energy-use", "type" : "article-journal", "volume" : "31" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1038/351312a0", "ISBN" : "0028-0836", "ISSN" : "0028-0836", "PMID" : "1992", "abstract" : "The relationship between abundance and body size is the subject of considerable debate in ecology. Several data sets spanning a large range of body sizes show linear negative relationships between abundance and weight when these are measured on a logarithmic scale. But other studies of the abundances of species from single taxa, such as birds, which span a narrower range of body sizes reveal either little or no relationship, or a triangular relationship. Errors in estimating abundance might obscure relationships that do exist over a narrow range of body sizes. We describe here the relationship between body weight and abundance in British birds, whose population size estimates are unusually good. Abundance across all species declines with a -0.75 power of body weight, which conforms with the energetic equivalence 'rule'. There is, however, a significant positive relationship between abundance and body weight within lower taxa. Those tribes that do not share recent common ancestry with other British birds are most likely to show a positive relationship across their constituent species. We thus show that phylogenetic relatedness might be an important indicator of the structure of the relationship between body size and abundance.", "author" : [ { "dropping-particle" : "", "family" : "Nee", "given" : "Sean", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Read", "given" : "Andrew F.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Greenwood", "given" : "Jeremy J. D.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Harvey", "given" : "Paul H.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Nature", "id" : "ITEM-2", "issued" : { "date-parts" : [ [ "1991" ] ] }, "note" : "Orienter dig i citation 1-15", "page" : "312-313", "title" : "The relationship between abundance and body size in British birds", "type" : "article", "volume" : "351" }, "uris" : [ "" ] }, { "id" : "ITEM-3", "itemData" : { "DOI" : "10.1016/j.tree.2007.03.007", "ISSN" : "01695347", "author" : [ { "dropping-particle" : "", "family" : "White", "given" : "Ethan P", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Ernest", "given" : "S.K. Morgan", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Kerkhoff", "given" : "Andrew J", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Enquist", "given" : "Brian J", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Trends in Ecology & Evolution", "id" : "ITEM-3", "issue" : "6", "issued" : { "date-parts" : [ [ "2007", "6" ] ] }, "page" : "323-330", "title" : "Relationships between body size and abundance in ecology", "type" : "article-journal", "volume" : "22" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Damuth 1987; Nee et al. 1991; White et al. 2007b)", "plainTextFormattedCitation" : "(Damuth 1987; Nee et al. 1991; White et al. 2007b)", "previouslyFormattedCitation" : "(Damuth 1987; Nee et al. 1991; White et al. 2007b)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Damuth 1987; Nee et al. 1991; White et al. 2007b). However, when applying our approach by using a data-driven identification of natural phylogenetic substructure in the density-body size relation, other trends arose. We showed that within clades the global trend is broken, as has previously been shown for other organism groups ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1038/351268a0", "ISSN" : "0028-0836", "author" : [ { "dropping-particle" : "", "family" : "Damuth", "given" : "John", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Nature", "id" : "ITEM-1", "issue" : "6324", "issued" : { "date-parts" : [ [ "1991", "5", "23" ] ] }, "page" : "268-269", "title" : "Of size and abundance", "type" : "article-journal", "volume" : "351" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1890/13-0620.1", "ISBN" : "0012-9658", "ISSN" : "00129658", "PMID" : "24669745", "abstract" : "Ecological communities consist of small abundant and large non-abundant species. The energetic equivalence rule is an often-observed pattern that could be explained by equal energy usage among abundant small organisms and non-abundant large organisms. To generate this pattern, metabolism (as an indicator of individual energy use) and abundance have to scale inversely with body mass, and cancel each other out. In contrast, the pattern referred to as biomass equivalence states that the biomass of all species in an area should be constant across the body-mass range. In this study, we investigated forest soil communities with respect to metabolism, abundance, population energy use, and biomass. We focused on four land-use types in three different landscape blocks (Biodiversity Exploratories). The soil samples contained 870 species across 12 phylogenetic groups. Our results indicated positive sublinear metabolic scaling and negative sublinear abundance scaling with species body mass. The relationships varied mainly due to differences among phylogenetic groups or feeding types, and only marginally due to land-use type. However, these scaling relationships were not exactly inverse to each other, resulting in increasing population energy use and biomass with increasing body mass for most combinations of phylogenetic group or feeding type with land-use type. Thus, our results are mostly inconsistent with the classic perception of energetic equivalence, and reject the biomass equivalence hypothesis while documenting a specific and nonrandom pattern of how abundance, energy use, and biomass are distributed across size classes. However, these patterns are consistent with two alternative predictions: the resource-thinning hypothesis, which states that abundance decreases with trophic level, and the allometric degree hypothesis, which states that population energy use should increase with population average body mass, due to correlations with the number of links of consumers and resources. Overall, our results suggest that a synthesis of food web structures with metabolic theory may be most promising for predicting natural patterns of abundance, biomass, and energy use.", "author" : [ { "dropping-particle" : "", "family" : "Ehnes", "given" : "Roswitha B.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Pollierer", "given" : "Melanie M.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Erdmann", "given" : "Georgia", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Klarner", "given" : "Bernhard", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Eitzinger", "given" : "Bernhard", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Digel", "given" : "Christoph", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Ott", "given" : "David", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Maraun", "given" : "Mark", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Scheu", "given" : "Stefan", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Brose", "given" : "Ulrich", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecology", "id" : "ITEM-2", "issue" : "2", "issued" : { "date-parts" : [ [ "2014" ] ] }, "page" : "527-537", "title" : "Lack of energetic equivalence in forest soil invertebrates", "type" : "article-journal", "volume" : "95" }, "uris" : [ "" ] }, { "id" : "ITEM-3", "itemData" : { "DOI" : "10.1038/351312a0", "ISBN" : "0028-0836", "ISSN" : "0028-0836", "PMID" : "1992", "abstract" : "The relationship between abundance and body size is the subject of considerable debate in ecology. Several data sets spanning a large range of body sizes show linear negative relationships between abundance and weight when these are measured on a logarithmic scale. But other studies of the abundances of species from single taxa, such as birds, which span a narrower range of body sizes reveal either little or no relationship, or a triangular relationship. Errors in estimating abundance might obscure relationships that do exist over a narrow range of body sizes. We describe here the relationship between body weight and abundance in British birds, whose population size estimates are unusually good. Abundance across all species declines with a -0.75 power of body weight, which conforms with the energetic equivalence 'rule'. There is, however, a significant positive relationship between abundance and body weight within lower taxa. Those tribes that do not share recent common ancestry with other British birds are most likely to show a positive relationship across their constituent species. We thus show that phylogenetic relatedness might be an important indicator of the structure of the relationship between body size and abundance.", "author" : [ { "dropping-particle" : "", "family" : "Nee", "given" : "Sean", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Read", "given" : "Andrew F.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Greenwood", "given" : "Jeremy J. D.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Harvey", "given" : "Paul H.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Nature", "id" : "ITEM-3", "issued" : { "date-parts" : [ [ "1991" ] ] }, "note" : "Orienter dig i citation 1-15", "page" : "312-313", "title" : "The relationship between abundance and body size in British birds", "type" : "article", "volume" : "351" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Nee et al. 1991; Damuth 1991; Ehnes et al. 2014)", "plainTextFormattedCitation" : "(Nee et al. 1991; Damuth 1991; Ehnes et al. 2014)", "previouslyFormattedCitation" : "(Nee et al. 1991; Damuth 1991; Ehnes et al. 2014)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Nee et al. 1991; Damuth 1991; Ehnes et al. 2014), and that all within-group slopes are weaker than the overall relationship. Similar results have been found within specific taxonomic groups of mammals before; we demonstrate here for the first time that clade specific variation in the body size-population density relationship (SDR) for all sub-clades is less negative than the SDR across Mammalia.The pattern across all Mammalia as a whole shows that EER could be considered valid since the SDR scales inversely to overall mean metabolism ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1111/j.1461-0248.2010.01461.x", "ISBN" : "1461-023X", "ISSN" : "1461023X", "PMID" : "20353439", "abstract" : "The metabolic theory of ecology links physiology with ecology, and successfully predicts many allometric scaling relationships. In recent years, proponents and critics of metabolic theory have debated vigorously about the scaling of metabolic rate. We show that the controversy arose, in part, because researchers examined the mean exponent separately from the variance. We estimate both quantities simultaneously using linear mixed-effects models and data from 1242 animal species. Metabolic rate scaling converges on the predicted value of 3/4 but is highly heterogeneous: 50% of orders lie outside the range 0.68-0.82. These findings are robust to several forms of statistical uncertainty. We then test competing hypotheses about the variation. Metabolic theory is currently unable to explain differences in scaling among orders, but the patterns are not consistent with competing explanations either. We conclude that current theories are inadequate to explain the full range of metabolic scaling patterns observed in nature.", "author" : [ { "dropping-particle" : "", "family" : "Isaac", "given" : "N. J B", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Carbone", "given" : "Chris", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecology Letters", "id" : "ITEM-1", "issued" : { "date-parts" : [ [ "2010" ] ] }, "page" : "728-735", "title" : "Why are metabolic scaling exponents so controversial? Quantifying variance and testing hypotheses", "type" : "article", "volume" : "13" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Isaac & Carbone 2010)", "plainTextFormattedCitation" : "(Isaac & Carbone 2010)", "previouslyFormattedCitation" : "(Isaac & Carbone 2010)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Isaac & Carbone 2010). When we see that within groups the SDR is shallower than across all species combined, it indicates that either EER is flawed, or that metabolic scaling within groups is also less steep. Research on metabolic scaling within clades have shown variability in slopes from low slopes of 0.53-0.55 in soricids and rodents to as high as 0.87 in chiropterans, but show that most groups fall around 0.75 ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1086/606023", "ISSN" : "0003-0147", "author" : [ { "dropping-particle" : "", "family" : "Sieg", "given" : "Annette\u00a0E.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "O\u2019Connor", "given" : "Michael\u00a0P.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "McNair", "given" : "James\u00a0N.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Grant", "given" : "Bruce\u00a0W.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Agosta", "given" : "Salvatore\u00a0J.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Dunham", "given" : "Arthur\u00a0E.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "The American Naturalist", "id" : "ITEM-1", "issue" : "5", "issued" : { "date-parts" : [ [ "2009" ] ] }, "page" : "720-733", "title" : "Mammalian Metabolic Allometry: Do Intraspecific Variation, Phylogeny, and Regression Models Matter?", "type" : "article-journal", "volume" : "174" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Sieg et al. 2009)", "plainTextFormattedCitation" : "(Sieg et al. 2009)", "previouslyFormattedCitation" : "(Sieg et al. 2009)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Sieg et al. 2009), consistent with what others have found across a broader range of animal orders ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1111/j.1461-0248.2010.01461.x", "ISBN" : "1461-023X", "ISSN" : "1461023X", "PMID" : "20353439", "abstract" : "The metabolic theory of ecology links physiology with ecology, and successfully predicts many allometric scaling relationships. In recent years, proponents and critics of metabolic theory have debated vigorously about the scaling of metabolic rate. We show that the controversy arose, in part, because researchers examined the mean exponent separately from the variance. We estimate both quantities simultaneously using linear mixed-effects models and data from 1242 animal species. Metabolic rate scaling converges on the predicted value of 3/4 but is highly heterogeneous: 50% of orders lie outside the range 0.68-0.82. These findings are robust to several forms of statistical uncertainty. We then test competing hypotheses about the variation. Metabolic theory is currently unable to explain differences in scaling among orders, but the patterns are not consistent with competing explanations either. We conclude that current theories are inadequate to explain the full range of metabolic scaling patterns observed in nature.", "author" : [ { "dropping-particle" : "", "family" : "Isaac", "given" : "N. J B", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Carbone", "given" : "Chris", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecology Letters", "id" : "ITEM-1", "issued" : { "date-parts" : [ [ "2010" ] ] }, "page" : "728-735", "title" : "Why are metabolic scaling exponents so controversial? Quantifying variance and testing hypotheses", "type" : "article", "volume" : "13" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Isaac & Carbone 2010)", "plainTextFormattedCitation" : "(Isaac & Carbone 2010)", "previouslyFormattedCitation" : "(Isaac & Carbone 2010)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Isaac & Carbone 2010). We found that the SDR was less steep than that for most clades. This indicates an asymmetry in population-level energy use, where larger-bodied species within clades succeed in acquiring more energy than the smaller-bodied species. ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1371/journal.pone.0057449", "ISSN" : "19326203", "PMID" : "23460858", "abstract" : "The energy equivalence rule (EER) is a macroecological hypothesis that posits that total population energy use (PEU) should be independent of species body mass, because population densities and energy metabolisms scale with body mass in a directly inverse manner. However, evidence supporting the EER is equivocal, and the use of basal metabolic rate (BMR) in such studies has been questioned; ecologically-relevant indices like field metabolic rate (FMR) are probably more appropriate. In this regard, Australian marsupials present a novel test for the EER because, unlike eutherians, marsupial BMRs and FMRs scale differently with body mass. Based on either FMR or BMR, Australian marsupial PEU did not obey an EER, and scaled positively with body mass based on ordinary least squares (OLS) regressions. Importantly, the scaling of marsupial population density with body mass had a slope of \u22120.37, significantly shallower than the expected slope of \u22120.75, and not directly inverse of body-mass scaling exponents for BMR (0.72) or FMR (0.62). The findings suggest that the EER may not be a causal, universal rule, or that for reasons not yet clear, it is not operating for Australia\u2019s unique native fauna.", "author" : [ { "dropping-particle" : "", "family" : "Munn", "given" : "Adam J.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Dunne", "given" : "Craig", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "M\u00fcller", "given" : "Dennis W H", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Clauss", "given" : "Marcus", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "PLoS ONE", "id" : "ITEM-1", "issue" : "2", "issued" : { "date-parts" : [ [ "2013" ] ] }, "page" : "1-5", "title" : "Energy In-Equivalence in Australian Marsupials: Evidence for Disruption of the Continent's Mammal Assemblage, or Are Rules Meant to Be Broken?", "type" : "article-journal", "volume" : "8" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Munn et al. 2013)", "manualFormatting" : "Munn et al. (2013)", "plainTextFormattedCitation" : "(Munn et al. 2013)", "previouslyFormattedCitation" : "(Munn et al. 2013)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }Munn et al. (2013) demonstrated this phenomenon in Australian marsupials, where metabolism (both field and basal metabolic rate) scales more steeply than density with body mass leading the total energy flux to be positively scaled with body mass.A simulation study has shown how such a pattern could arise because smaller-bodied species can maintain larger populations on less energy than larger-bodied species ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1086/513495", "ISBN" : "0003-0147/2007/16905-41931$15.00", "ISSN" : "0003-0147", "PMID" : "17427133", "abstract" : "Across a wide array of animal species, mean population densities decline with species body mass such that the rate of energy use of local populations is approximately independent of body size. This \"energetic equivalence\" is particularly evident when ecological population densities are plotted across several or more orders of magnitude in body mass and is supported by a considerable body of evidence. Nevertheless, interpretation of the data has remained controversial, largely because of the difficulty of explaining the origin and maintenance of such a size-abundance relationship in terms of purely ecological processes. Here I describe results of a simulation model suggesting that an extremely simple mechanism operating over evolutionary time can explain the major features of the empirical data. The model specifies only the size scaling of metabolism and a process where randomly chosen species evolve to take resource energy from other species. This process of energy exchange among particular species is distinct from a random walk of species abundances and creates a situation in which species populations using relatively low amounts of energy at any body size have an elevated extinction risk. Selective extinction of such species rapidly drives size-abundance allometry in faunas toward approximate energetic equivalence and maintains it there.", "author" : [ { "dropping-particle" : "", "family" : "Damuth", "given" : "John", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "The American Naturalist", "id" : "ITEM-1", "issue" : "5", "issued" : { "date-parts" : [ [ "2007", "5" ] ] }, "page" : "621-631", "title" : "A macroevolutionary explanation for energy equivalence in the scaling of body size and population density", "type" : "article-journal", "volume" : "169" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Damuth 2007)", "plainTextFormattedCitation" : "(Damuth 2007)", "previouslyFormattedCitation" : "(Damuth 2007)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Damuth 2007). When the slope is steeper than EER, of which we saw no indication, larger-bodied species populations are living on the lower total energy use than smaller species. Extinction of the larger species is therefore more likely, and such extinctions will eventually drive the relationship back towards EER. In contrast, slopes which are less steep than expected by the EER, as we found, indicate that smaller-bodied species populations are using relatively less total energy. Since smaller-bodied species populations can survive on far less energy than is required for a viable population of large species it is possible to drive relationship towards a more positive relationship than the EER states ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1086/513495", "ISBN" : "0003-0147/2007/16905-41931$15.00", "ISSN" : "0003-0147", "PMID" : "17427133", "abstract" : "Across a wide array of animal species, mean population densities decline with species body mass such that the rate of energy use of local populations is approximately independent of body size. This \"energetic equivalence\" is particularly evident when ecological population densities are plotted across several or more orders of magnitude in body mass and is supported by a considerable body of evidence. Nevertheless, interpretation of the data has remained controversial, largely because of the difficulty of explaining the origin and maintenance of such a size-abundance relationship in terms of purely ecological processes. Here I describe results of a simulation model suggesting that an extremely simple mechanism operating over evolutionary time can explain the major features of the empirical data. The model specifies only the size scaling of metabolism and a process where randomly chosen species evolve to take resource energy from other species. This process of energy exchange among particular species is distinct from a random walk of species abundances and creates a situation in which species populations using relatively low amounts of energy at any body size have an elevated extinction risk. Selective extinction of such species rapidly drives size-abundance allometry in faunas toward approximate energetic equivalence and maintains it there.", "author" : [ { "dropping-particle" : "", "family" : "Damuth", "given" : "John", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "The American Naturalist", "id" : "ITEM-1", "issue" : "5", "issued" : { "date-parts" : [ [ "2007", "5" ] ] }, "page" : "621-631", "title" : "A macroevolutionary explanation for energy equivalence in the scaling of body size and population density", "type" : "article-journal", "volume" : "169" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Damuth 2007)", "plainTextFormattedCitation" : "(Damuth 2007)", "previouslyFormattedCitation" : "(Damuth 2007)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Damuth 2007). Other studies points to the importance of size-structured competition in real communities, where large-bodied species take relatively more of the resource pool than populations of small-bodied species ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1016/j.ecocom.2009.08.005", "ISBN" : "1476-945X", "ISSN" : "1476945X", "abstract" : "The relation between population density and body mass has vexed ecologists for nearly 30 years as a consequence of high variability in the observed slope of the relation: No single generalisation of the relation has been accepted as universally representative. Here, we use a simple computational approach to examine how observational scale (the body mass range considered) determines variation in the density-mass pattern. Our model relies on two assumptions: (1) resources are partitioned in an unbiased manner among species with different masses; (2) the number of individuals that can be supported by a given quantity of resources is related to their metabolic rate (which is a function of their mass raised to the power of a scaling coefficient, b). We show that density (1) scales as a function of body mass raised to the power of -b on average, but (2) the slope of the relation varies considerably at smaller scales of observation (over narrow ranges of body mass) as a consequence of details of species' ecology associated with resource procurement. Historically, the effect of body mass range on the slope of the density-mass relation has been unfailingly attributed to a statistical effect. Here we show that the effect of body mass range on the slope of the density-mass relation may equally result from a biological mechanism, though we find it impossible to distinguish between the two. We observe that many of the explanations that have been offered to account for the variability in the slope of the relation invoke mechanisms associated with differences in body mass and we therefore suggest that body mass range itself might be the most important explanatory factor. Notably, our results imply that the energetic equivalence rule should not be expected to hold at smaller scales of observation, which suggests that it may not be possible to scale the mass- and temperature-dependence of organism metabolism to predict patterns at higher levels of biological organisation at smaller scales of observation. ?? 2009 Elsevier B.V. All rights reserved.", "author" : [ { "dropping-particle" : "", "family" : "Hayward", "given" : "April", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Kolasa", "given" : "Jurek", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Stone", "given" : "Jonathon R.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecological Complexity", "id" : "ITEM-1", "issue" : "1", "issued" : { "date-parts" : [ [ "2010", "3" ] ] }, "page" : "115-124", "publisher" : "Elsevier B.V.", "title" : "The scale-dependence of population density-body mass allometry: Statistical artefact or biological mechanism?", "type" : "article-journal", "volume" : "7" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1098/rsbl.2011.0036", "ISBN" : "1744-9561", "ISSN" : "1744-9561", "PMID" : "21367781", "abstract" : "The energetic equivalence rule states that population-level metabolic rate is independent of average body size. This rule has been both supported and refuted by allometric studies of abundance and individual metabolic rate, but no study, to my knowledge, has tested the rule with direct measurements of whole-population metabolic rate. Here, I find a positive scaling of whole-colony metabolic rate with body size for eusocial insects. Individual metabolic rates in these colonies scaled with body size more steeply than expected from laboratory studies on insects, while population size was independent of body size. Using consumer-resource models, I suggest that the colony-level metabolic rate scaling observed here may arise from a change in the scaling of individual metabolic rate resulting from a change in the body size dependence of mortality rates.", "author" : [ { "dropping-particle" : "", "family" : "DeLong", "given" : "John P", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Biology Letters", "id" : "ITEM-2", "issue" : "March", "issued" : { "date-parts" : [ [ "2011" ] ] }, "page" : "611-614", "title" : "Energetic inequivalence in eusocial insect colonies.", "type" : "article-journal", "volume" : "7" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Hayward et al. 2010; DeLong 2011)", "plainTextFormattedCitation" : "(Hayward et al. 2010; DeLong 2011)", "previouslyFormattedCitation" : "(Hayward et al. 2010; DeLong 2011)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Hayward et al. 2010; DeLong 2011).That larger species take a larger part of the energy pool could be an explanation for Cope’s rule, which states that species within clades tend to increase in size through evolution ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.2307/2407115", "ISBN" : "00143820", "ISSN" : "0014-3820", "abstract" : "Whether body size will increase or decrease in an evolving population depends on whether mean body size is larger or smaller than the optimum for the population. Cope's Rule, the generalization that most animal groups have evolved toward larger body size, cannot be explained by intrinsic advantages of large size. Rather, it is the tendency of groups to arise at small body size relative to their optima that produces the widely observed pattern of net size increase. The specialized nature of large species of a given body plan, required by problems of similitude, renders these forms unlikely potential ancestors for major new descendent taxa. The adaptive discontinuity that must be crossed for invasion of a new adaptive zone at large body size exists because of the need for descendent taxa to be specialized along new lines. These factors tend to restrict large-scale adaptive breakthroughs to small body sizes. Size changes probably tend to occur sporadically, during speciation events. Size increase is not inherently favored in speciation, but prevails during diversification because origin of a higher taxon at small body size concentrates many early species in the small size range. Nearly all diverse animal orders and classes, and many families and super-families, are composed of species whose body sizes are distributed as positively skewed histograms. The typical pattern of size change during diversification of such a group can be determined from time-series plots for fossil species of diversifying higher taxa. A major taxon normally arises at small body size relative to its potential size range, and a slightly skewed histogram is rapidly formed. The histogram may expand or contract slightly in the small size range as diversification proceeds, but spreads continually farther in a positive direction, to develop a strongly attenuated tail in the large size range. Skewing occurs very rapidly because possible increments of size change with speciation are not constant throughout a taxon's size range, but are a direct function of body size, so that early spreading of the range proceeds more rapidly in a positive direction than in a negative direction. Nearly always an increase in mean size results. Just as taxonomic and morphologic diversification approach limits as a group's potential adaptive zone is filled, the size-frequency plot approaches a limiting distribution. The attenuated right flank of a high-diversity distribution reflects not only well known ecologi\u2026", "author" : [ { "dropping-particle" : "", "family" : "Stanley", "given" : "Steven", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Evolution", "id" : "ITEM-1", "issue" : "1", "issued" : { "date-parts" : [ [ "1973", "3" ] ] }, "page" : "1-26", "title" : "An explanation for Cope's Rule", "type" : "article-journal", "volume" : "27" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Stanley 1973)", "plainTextFormattedCitation" : "(Stanley 1973)", "previouslyFormattedCitation" : "(Stanley 1973)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Stanley 1973). This has been observed in mammals e.g. in Carnivora ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1126/science.1102417", "ISBN" : "0036-8075", "ISSN" : "0036-8075", "PMID" : "15459388", "abstract" : "Over the past 50 million years, successive clades of large carnivorous mammals diversified and then declined to extinction. In most instances, the cause of the decline remains a puzzle. Here we argue that energetic constraints and pervasive selection for larger size (Cope's rule) in carnivores lead to dietary specialization (hypercarnivory) and increased vulnerability to extinction. In two major clades of extinct North American canids, the evolution of large size was associated with a dietary shift to hypercarnivory and a decline in species durations. Thus, selection for attributes that promoted individual success resulted in progressive evolutionary failure of their clades.", "author" : [ { "dropping-particle" : "", "family" : "Valkenburgh", "given" : "Blaire", "non-dropping-particle" : "Van", "parse-names" : false, "suffix" : "" } ], "container-title" : "Science", "id" : "ITEM-1", "issue" : "5693", "issued" : { "date-parts" : [ [ "2004", "10", "1" ] ] }, "page" : "101-104", "title" : "Cope's Rule, Hypercarnivory, and Extinction in North American Canids", "type" : "article-journal", "volume" : "306" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Van Valkenburgh 2004)", "plainTextFormattedCitation" : "(Van Valkenburgh 2004)", "previouslyFormattedCitation" : "(Van Valkenburgh 2004)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Van Valkenburgh 2004), where the mean and maximum body mass within clades increases through time. The within-group relations documented by our study imply that larger species do in fact occupy a larger part of the resource pool within clades. In the literature, there are plenty of examples of larger species outcompeting smaller species within guilds. In carnivores we have an abundance of evidence of intraguild competition and killing ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1086/303189", "ISSN" : "0003-0147", "author" : [ { "dropping-particle" : "", "family" : "Palomares", "given" : "F", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Caro", "given" : "T. M.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "The American Naturalist", "id" : "ITEM-1", "issue" : "5", "issued" : { "date-parts" : [ [ "1999", "5" ] ] }, "page" : "492-508", "title" : "Interspecific killing among mammalian carnivores", "type" : "article-journal", "volume" : "153" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1086/501033", "ISSN" : "1537-5323", "PMID" : "16670995", "abstract" : "Interspecific killing is a key determinant of the abundances and distributions of carnivores, their prey, and nonprey community members. Similarity of body size has been proposed to lead competitors to seek similar prey, which increases the likelihood of interference encounters, including lethal ones. We explored the influence of body size, diet, predatory habits, and taxonomic relatedness on interspecific killing. The frequency of attacks depends on differences in body size: at small and large differences, attacks are less likely to occur; at intermediate differences, killing interactions are frequent and related to diet overlap. Further, the importance of interspecific killing as a mortality factor in the victim population increases with an increase in body size differences between killers and victims. Carnivores highly adapted to kill vertebrate prey are more prone to killing interactions, usually with animals of similar predatory habits. Family-level taxonomy influences killing interactions; carnivores tend to interact more with species in the same family than with species in different families. We conclude that although resource exploitation (diet), predatory habits, and taxonomy are influential in predisposing carnivores to attack each other, relative body size of the participants is overwhelmingly important. We discuss the implications of interspecific killing for body size and the dynamics of geographic ranges.", "author" : [ { "dropping-particle" : "", "family" : "Donadio", "given" : "Emiliano", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Buskirk", "given" : "Steven W", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "The American naturalist", "id" : "ITEM-2", "issue" : "4", "issued" : { "date-parts" : [ [ "2006", "4" ] ] }, "page" : "524-36", "title" : "Diet, morphology, and interspecific killing in carnivora.", "type" : "article-journal", "volume" : "167" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Palomares & Caro 1999; Donadio & Buskirk 2006)", "plainTextFormattedCitation" : "(Palomares & Caro 1999; Donadio & Buskirk 2006)", "previouslyFormattedCitation" : "(Palomares & Caro 1999; Donadio & Buskirk 2006)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Palomares & Caro 1999; Donadio & Buskirk 2006). For example, grey wolves (Canis lupus) limit the density of coyotes (Canis latrans) through intraguild predation and predation ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1111/j.1365-2656.2007.01287.x", "ISBN" : "0021-8790 (Print)\\n0021-8790 (Linking)", "ISSN" : "00218790", "PMID" : "17922704", "abstract" : "Interference competition with wolves Canis lupus is hypothesized to limit the distribution and abundance of coyotes Canis latrans, and the extirpation of wolves is often invoked to explain the expansion in coyote range throughout much of North America. We used spatial, seasonal and temporal heterogeneity in wolf distribution and abundance to test the hypothesis that interference competition with wolves limits the distribution and abundance of coyotes. From August 2001 to August 2004, we gathered data on cause-specific mortality and survival rates of coyotes captured at wolf-free and wolf-abundant sites in Grand Teton National Park (GTNP), Wyoming, USA, to determine whether mortality due to wolves is sufficient to reduce coyote densities. We examined whether spatial segregation limits the local distribution of coyotes by evaluating home-range overlap between resident coyotes and wolves, and by contrasting dispersal rates of transient coyotes captured in wolf-free and wolf-abundant areas. Finally, we analysed data on population densities of both species at three study areas across the Greater Yellowstone Ecosystem (GYE) to determine whether an inverse relationship exists between coyote and wolf densities. Although coyotes were the numerically dominant predator, across the GYE, densities varied spatially and temporally in accordance with wolf abundance. Mean coyote densities were 33% lower at wolf-abundant sites in GTNP, and densities declined 39% in Yellowstone National Park following wolf reintroduction. A strong negative relationship between coyote and wolf densities (beta = -3.988, P < 0.005, r(2) = 0.54, n = 16), both within and across study sites, supports the hypothesis that competition with wolves limits coyote populations. Overall mortality of coyotes resulting from wolf predation was low, but wolves were responsible for 56% of transient coyote deaths (n = 5). In addition, dispersal rates of transient coyotes captured at wolf-abundant sites were 117% higher than for transients captured in wolf-free areas. Our results support the hypothesis that coyote abundance is limited by competition with wolves, and suggest that differential effects on survival and dispersal rates of transient coyotes are important mechanisms by which wolves reduce coyote densities.", "author" : [ { "dropping-particle" : "", "family" : "Berger", "given" : "Kim Murray", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Gese", "given" : "Eric M.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Journal of Animal Ecology", "id" : "ITEM-1", "issue" : "6", "issued" : { "date-parts" : [ [ "2007" ] ] }, "page" : "1075-1085", "title" : "Does interference competition with wolves limit the distribution and abundance of coyotes?", "type" : "article-journal", "volume" : "76" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Berger & Gese 2007)", "plainTextFormattedCitation" : "(Berger & Gese 2007)", "previouslyFormattedCitation" : "(Berger & Gese 2007)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Berger & Gese 2007), which in turn limit the density and distribution of grey foxes (Urocyon cinereoargenteus) due to both greater ecological generalism and and direct killing ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1007/s004420000448", "ISSN" : "0029-8549", "PMID" : "24595837", "abstract" : "We examined the relative roles of dominance in agonistic interactions and energetic constraints related to body size in determining local abundances of coyotes (Canis latrans, 8-20 kg), gray foxes (Urocyon cinereoargenteus, 3-5 kg) and bobcats (Felis rufus, 5-15 kg) at three study sites (hereafter referred to as NP, CP, and SP) in the Santa Monica Mountains of California. We hypothesized that the largest and behaviorally dominant species, the coyote, would exploit a wider range of resources (i.e., a higher number of habitat and/or food types) and, consequently, would occur in higher density than the other two carnivores. We evaluated our hypotheses by quantifying their diets, food overlap, habitat-specific abundances, as well as their overall relative abundance at the three study sites. We identified behavioral dominance of coyotes over foxes and bobcats in Santa Monica because 7 of 12 recorded gray fox deaths and 2 of 5 recorded bobcat deaths were due to coyote predation, and no coyotes died as a result of their interactions with bobcats or foxes. Coyotes and bobcats were present in a variety of habitats types (8 out of 9), including both open and brushy habitats, whereas gray foxes were chiefly restricted to brushy habitats. There was a negative relationship between the abundances of coyotes and gray foxes (P=0.020) across habitats, suggesting that foxes avoided habitats of high coyote predation risk. Coyote abundance was low in NP, high in CP, and intermediate in SP. Bobcat abundance changed little across study sites, and gray foxes were very abundant in NP, absent in CP, and scarce in SP; this suggests a negative relationship between coyote and fox abundances across study sites, as well. Bobcats were solely carnivorous, relying on small mammals (lagomorphs and rodents) throughout the year and at all three sites. Coyotes and gray foxes also relied on small mammals year-round at all sites, though they also ate significant amounts of fruit. Though there were strong overall interspecific differences in food habits of carnivores (P<0.0001), average seasonal food overlaps were high due to the importance of small mammals in all carnivore diets [bobcat-gray fox: 0.79\u00b10.09 (SD), n=4; bobcat-coyote: 0.69\u00b10.16, n=6; coyote-gray fox: 0.52\u00b10.05, n=4]. As hypothesized, coyotes used more food types and more habitat types than did bobcats and gray foxes and, overall, coyotes were the most abundant of the three species and ranged more widely than did gray foxes. We \u2026", "author" : [ { "dropping-particle" : "", "family" : "Fedriani", "given" : "JM", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Fuller", "given" : "TK", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Sauvajot", "given" : "RM", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "York", "given" : "EC", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Oecologia", "id" : "ITEM-1", "issue" : "2", "issued" : { "date-parts" : [ [ "2000", "10" ] ] }, "page" : "258-70", "title" : "Competition and intraguild predation among three sympatric carnivores", "type" : "article-journal", "volume" : "125" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Fedriani et al. 2000)", "plainTextFormattedCitation" : "(Fedriani et al. 2000)", "previouslyFormattedCitation" : "(Fedriani et al. 2000)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Fedriani et al. 2000). Intraguild killing have also been shown in Eurasian lynx (Lynx lynx) killing red foxes (Vulpes vulpes), unrelated to feeding ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1111/j.1600-0587.1999.tb01281.x", "ISSN" : "09067590", "author" : [ { "dropping-particle" : "", "family" : "Sunde", "given" : "Peter", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Overskaug", "given" : "Kristian", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Kvam", "given" : "Tor", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecography", "id" : "ITEM-1", "issue" : "5", "issued" : { "date-parts" : [ [ "1999", "12" ] ] }, "page" : "521-523", "title" : "Intraguild predation of lynxes on foxes: evidence of interference competition?", "type" : "article-journal", "volume" : "22" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Sunde, Overskaug & Kvam 1999)", "plainTextFormattedCitation" : "(Sunde, Overskaug & Kvam 1999)", "previouslyFormattedCitation" : "(Sunde, Overskaug & Kvam 1999)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Sunde, Overskaug & Kvam 1999). There is also a growing body of evidence for mesopredator (foxes and cats) regulation by top predators (dingoes) in Australia ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1111/j.1442-9993.2007.01721.x", "ISSN" : "1442-9985", "author" : [ { "dropping-particle" : "", "family" : "Glen", "given" : "A. S.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Dickman", "given" : "C. R.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Soul\u00e9", "given" : "M. E.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Mackey", "given" : "B. G.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Austral Ecology", "id" : "ITEM-1", "issue" : "5", "issued" : { "date-parts" : [ [ "2007", "8" ] ] }, "page" : "492-501", "title" : "Evaluating the role of the dingo as a trophic regulator in Australian ecosystems", "type" : "article-journal", "volume" : "32" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Glen et al. 2007)", "plainTextFormattedCitation" : "(Glen et al. 2007)", "previouslyFormattedCitation" : "(Glen et al. 2007)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Glen et al. 2007). Within rodents, there are several experimental examples on competitive suppression or exclusion by the larger species, e.g. a removal experiment shows an asymmetrical result where the smaller bank vole (Myodes glareolus) increased in density when the larger wood mouse (Apodemus sylvaticus) was removed, while no effect was found when the smaller species was removed ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.4098/AT.arch.00-35", "ISBN" : "0001-7051", "ISSN" : "00017051", "abstract" : "Experimental removal was conducted to test interspecific competition between the wood mouse Apodemus sylvaticus (Linnaeus, 1758) and the bank vole Clethrionomys glareolus (Schreber, 1780) that dominate the rodent communities in the forested biotopes through most of central Europe. Population density, body mass, reproductive condition, and habitat use were compared among two experimental sites (where one of the species had been removed) and one control site. The 5-year-study included pre-removal, removal, and post-removal periods. Reproductive condition was not affected by the density of the competitor or the conspecifics. Also, we did not detected any habitat shift that could be related to competitive release. However, the removal of wood mice strongly affected the population density of bank voles, but the removal of bank voles affected density of wood mice only slightly. Thus, we conclude that the competitive effect was asymmetrical.", "author" : [ { "dropping-particle" : "", "family" : "Fasola", "given" : "Mauro", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Canova", "given" : "Luca", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Acta Theriologica", "id" : "ITEM-1", "issue" : "3", "issued" : { "date-parts" : [ [ "2000", "9", "12" ] ] }, "page" : "353-365", "title" : "Asymmetrical competition between the bank vole and the wood mouse, a removal experiment", "type" : "article-journal", "volume" : "45" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Fasola & Canova 2000)", "plainTextFormattedCitation" : "(Fasola & Canova 2000)", "previouslyFormattedCitation" : "(Fasola & Canova 2000)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Fasola & Canova 2000). Another experiment showed overall fitness decrease of bank voles due to the suppression by larger field voles (Microtus agrestis) ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1034/j.1600-0706.2002.11833.x", "ISSN" : "0030-1299", "author" : [ { "dropping-particle" : "", "family" : "Eccard", "given" : "Jana a", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Ylonen", "given" : "Hannu", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Oikos", "id" : "ITEM-1", "issue" : "3", "issued" : { "date-parts" : [ [ "2002", "12" ] ] }, "page" : "580-590", "title" : "Direct interference or indirect exploitation? An experimental study of fitness costs of interspecific competition in voles", "type" : "article-journal", "volume" : "99" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Eccard & Ylonen 2002)", "plainTextFormattedCitation" : "(Eccard & Ylonen 2002)", "previouslyFormattedCitation" : "(Eccard & Ylonen 2002)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Eccard & Ylonen 2002). Further, intrinsic species traits can also affect how larger species within a clade has a competitive advantage over smaller species without interaction: e.g. larger species are less susceptible to carnivore attacks and can therefore use areas not available to smaller species, and larger ruminants can also utilize more abundant low-quality food due larger guts and slower gut passage time ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1111/j.1365-2656.2011.01885.x", "ISBN" : "1365-2656", "ISSN" : "00218790", "PMID" : "21801174", "abstract" : "1. Theory predicts that small grazers are regulated by the digestive quality of grass, while large grazers extract sufficient nutrients from low-quality forage and are regulated by its abundance instead. In addition, predation potentially affects populations of small grazers more than large grazers, because predators have difficulty capturing and handling large prey. \\n2. We analyse the spatial distribution of five grazer species of different body size in relation to gradients of food availability and predation risk. Specifically, we investigate how the quality of grass, the abundance of grass biomass and the associated risks of predation affect the habitat use of small, intermediate and large savanna grazers at a landscape level. \\n3. Resource selection functions of five mammalian grazer species surveyed over a 21-year period in Serengeti are calculated using logistic regressions. Variables included in the analyses are grass nitrogen, rainfall, topographic wetness index, woody cover, drainage lines, landscape curvature, water and human habitation. Structural equation modelling (SEM) is used to aggregate predictor variables into 'composites' representing food quality, food abundance and predation risk. Subsequently, SEM is used to investigate species' habitat use, defined as their recurrence in 5 \u00d7 5 km cells across repeated censuses. \\n4. The distribution of small grazers is constrained by predation and food quality, whereas the distribution of large grazers is relatively unconstrained. The distribution of the largest grazer (African buffalo) is primarily associated with forage abundance but not predation risk, while the distributions of the smallest grazers (Thomson's gazelle and Grant's gazelle) are associated with high grass quality and negatively with the risk of predation. The distributions of intermediate sized grazers (Coke's hartebeest and topi) suggest they optimize access to grass biomass of sufficient quality in relatively predator-safe areas. \\n5. The results illustrate how top-down (vegetation-mediated predation risk) and bottom-up factors (biomass and nutrient content of vegetation) predictably contribute to the division of niche space for herbivores that vary in body size. Furthermore, diverse grazing assemblages are composed of herbivores of many body sizes (rather than similar body sizes), because these herbivores best exploit the resources of different habitat types.", "author" : [ { "dropping-particle" : "", "family" : "Hopcraft", "given" : "J. Grant C", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Anderson", "given" : "T. Michael", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "P\u00e9rez-Vila", "given" : "Saleta", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Mayemba", "given" : "Emilian", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Olff", "given" : "Han", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Journal of Animal Ecology", "id" : "ITEM-1", "issue" : "1", "issued" : { "date-parts" : [ [ "2012", "1" ] ] }, "page" : "201-213", "title" : "Body size and the division of niche space: food and predation differentially shape the distribution of Serengeti grazers", "type" : "article-journal", "volume" : "81" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Hopcraft et al. 2012)", "plainTextFormattedCitation" : "(Hopcraft et al. 2012)", "previouslyFormattedCitation" : "(Hopcraft et al. 2012)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Hopcraft et al. 2012).The SDR did not become steeper with body mass span of a given group, as would be expected if the shallower relationship found within some groups was a statistical artefact, but we did see stronger negative relationships for groups with average larger body masses (Fig. 2), as ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1038/351268a0", "ISSN" : "0028-0836", "author" : [ { "dropping-particle" : "", "family" : "Damuth", "given" : "John", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Nature", "id" : "ITEM-1", "issue" : "6324", "issued" : { "date-parts" : [ [ "1991", "5", "23" ] ] }, "page" : "268-269", "title" : "Of size and abundance", "type" : "article-journal", "volume" : "351" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Damuth 1991)", "manualFormatting" : "Damuth (1991)", "plainTextFormattedCitation" : "(Damuth 1991)", "previouslyFormattedCitation" : "(Damuth 1991)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }Damuth (1991) also points out could be expected. That groups of larger-bodied species have SDR closer to EER shows that the benefit of being large decreases with body mass. This might be explained by their larger home ranges, which are less easily defended and therefore have larger resource loss to neighbours ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1126/science.1102138", "ISBN" : "0036-8075", "ISSN" : "0036-8075", "PMID" : "15472074", "abstract" : "Space used by animals increases with increasing body size. Energy requirements alone can explain how population density decreases, but not the steep rate at which home range area increases. We present a general mechanistic model that predicts the frequency of interaction, spatial overlap, and loss of resources to neighbors. Extensive empirical evidence supports the model, demonstrating that spatial constraints on defense cause exclusivity of home range use to decrease with increasing body size. In large mammals, over 90% of available resources may be lost to neighbors. Our model offers a general framework to understand animal space use and sociality.", "author" : [ { "dropping-particle" : "", "family" : "Jetz", "given" : "Walter", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Science", "id" : "ITEM-1", "issue" : "5694", "issued" : { "date-parts" : [ [ "2004", "10", "8" ] ] }, "page" : "266-268", "title" : "The scaling of animal space use", "type" : "article-journal", "volume" : "306" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Jetz 2004)", "plainTextFormattedCitation" : "(Jetz 2004)", "previouslyFormattedCitation" : "(Jetz 2004)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Jetz 2004). There was almost no difference throughout the phylogeny in the intercept of the size–density relationship, with some notable exceptions. All members of the order Carnivora have population densities several orders of magnitude below most other species irrespective of body mass. This follows expectation from their high trophic level, which could explain a factor 10-100 drop in available energy for any given body mass compared lover levels in the food chain ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.2307/1930126", "ISSN" : "00129658", "author" : [ { "dropping-particle" : "", "family" : "Lindeman", "given" : "Raymond L.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecology", "id" : "ITEM-1", "issue" : "4", "issued" : { "date-parts" : [ [ "1942", "10" ] ] }, "page" : "399-417", "title" : "The Trophic-Dynamic Aspect of Ecology", "type" : "article-journal", "volume" : "23" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Lindeman 1942)", "plainTextFormattedCitation" : "(Lindeman 1942)", "previouslyFormattedCitation" : "(Lindeman 1942)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Lindeman 1942). In general we did not see that population density of carnivorous groups scaled more steeply with body mass than in herbivorous groups, as other studies have found ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1086/284596", "ISBN" : "00030147", "ISSN" : "0003-0147", "abstract" : "Accounting for the variation in the population density among different animal species is a central goal of animal ecology (Andrewartha and Birch 1954). Among mammals, density is closely related to the average adult body mass of the species and to the trophic level occupied by the species (Mohr 1940; Clutton-Brock and Harvey 1977; Eisenberg 1980; Damuth 1981a; Peters 1983; Peters and Raelson 1984). In addition, after the body mass and trophic position of species have been taken into account, population densities appear to vary with habitat (Eisenberg 1980) and biogeographical area (Peters and Raelson 1984). Peters and Raelson have been impressed by the predictive power of these relations: \"Because these relations appear so powerful . . . they should be examined as fully as possible before they come into widespread use\" (1984, p. 499).", "author" : [ { "dropping-particle" : "", "family" : "Robinson", "given" : "John G.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Redford", "given" : "Kent H.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "The American Naturalist", "id" : "ITEM-1", "issue" : "5", "issued" : { "date-parts" : [ [ "1986", "11" ] ] }, "page" : "665-680", "title" : "Body size, diet, and population density of neotropical forest mammals", "type" : "article-journal", "volume" : "128" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1890/11-1138.1", "ISBN" : "0012-9658", "ISSN" : "00129658", "abstract" : "Population abundance is negatively related to body size for many types of organisms. Despite the ubiquity of size-density scaling relationships, we lack a general understanding of the underlying mechanisms. Although dynamic models suggest that it is possible to predict the intercept and slope of the scaling relationship from prior observations, this has never been empirically attempted. Here we fully parameterize a set of consumer- resource models for mammalian carnivores and successfully predict the size-density scaling relationship for this group without the use of free parameters. All models produced similar predictions, but comparison of nested models indicated that the primary factors generating size-density scaling in mammalian carnivores are prey productivity, predator-prey size ratios, and consumer area of capture. \u00a9 2012 by the Ecological Society of America.", "author" : [ { "dropping-particle" : "", "family" : "DeLong", "given" : "John P.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Vasseur", "given" : "David a.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecology", "id" : "ITEM-2", "issue" : "3", "issued" : { "date-parts" : [ [ "2012" ] ] }, "note" : "Dynamic modelling with conclusitons on density~body mass scaling. And prey-pass~consumer mass.", "page" : "470-476", "title" : "A dynamic explanation of size-density scaling in carnivores", "type" : "article-journal", "volume" : "93" }, "uris" : [ "" ] }, { "id" : "ITEM-3", "itemData" : { "DOI" : "10.1126/science.1067994", "ISSN" : "00368075", "author" : [ { "dropping-particle" : "", "family" : "Carbone", "given" : "C.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Gittleman", "given" : "John L.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Science", "id" : "ITEM-3", "issue" : "5563", "issued" : { "date-parts" : [ [ "2002", "3", "22" ] ] }, "page" : "2273-2276", "title" : "A Common Rule for the Scaling of Carnivore Density", "type" : "article-journal", "volume" : "295" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Robinson & Redford 1986; Carbone & Gittleman 2002; DeLong & Vasseur 2012)", "plainTextFormattedCitation" : "(Robinson & Redford 1986; Carbone & Gittleman 2002; DeLong & Vasseur 2012)", "previouslyFormattedCitation" : "(Robinson & Redford 1986; Carbone & Gittleman 2002; DeLong & Vasseur 2012)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Robinson & Redford 1986; Carbone & Gittleman 2002; DeLong & Vasseur 2012). We do, however, see a trend potentially linked to diet variation within the carnivores. The order Carnivora has three distinct slopes, where the least carnivorous group had the least negative slope, whereas the most carnivorous has the most negative slope. This is consistent with mechanistic models which show that carnivory leads to a steeper SDR ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1086/519858", "ISBN" : "0003-0147/2007/17003-42157$15.00", "ISSN" : "0003-0147", "PMID" : "17879198", "abstract" : "The negative scaling of plant and animal abundance with body mass is one of the most fundamental relationships in ecology. However, theoretical approaches to explain this phenomenon make the unrealistic assumption that species share a homogeneous resource. Here we present a simple model linking mass and metabolism with density that includes the effects of consumer size on resource characteristics (particle size, density, and distribution). We predict patterns consistent with the energy equivalence rule (EER) under some scenarios. However, deviations from EER occur as a result of variation in resource distribution and productivity (e.g., due to the clumping of prey or variation in food particle size selection). We also predict that abundance scaling exponents change with the dimensionality of the foraging habitat. Our model predictions explain several inconsistencies in the observed scaling of vertebrate abundance among ecological and taxonomic groups and provide a broad framework for understanding variation in abundance.", "author" : [ { "dropping-particle" : "", "family" : "Carbone", "given" : "Chris", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Rowcliffe", "given" : "J. 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Further, the SDR of Carnivora has been accurately explained by consumer-resource models ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1890/11-1138.1", "ISBN" : "0012-9658", "ISSN" : "00129658", "abstract" : "Population abundance is negatively related to body size for many types of organisms. Despite the ubiquity of size-density scaling relationships, we lack a general understanding of the underlying mechanisms. Although dynamic models suggest that it is possible to predict the intercept and slope of the scaling relationship from prior observations, this has never been empirically attempted. Here we fully parameterize a set of consumer- resource models for mammalian carnivores and successfully predict the size-density scaling relationship for this group without the use of free parameters. All models produced similar predictions, but comparison of nested models indicated that the primary factors generating size-density scaling in mammalian carnivores are prey productivity, predator-prey size ratios, and consumer area of capture. \u00a9 2012 by the Ecological Society of America.", "author" : [ { "dropping-particle" : "", "family" : "DeLong", "given" : "John P.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Vasseur", "given" : "David a.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecology", "id" : "ITEM-1", "issue" : "3", "issued" : { "date-parts" : [ [ "2012" ] ] }, "note" : "Dynamic modelling with conclusitons on density~body mass scaling. And prey-pass~consumer mass.", "page" : "470-476", "title" : "A dynamic explanation of size-density scaling in carnivores", "type" : "article-journal", "volume" : "93" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(DeLong & Vasseur 2012)", "plainTextFormattedCitation" : "(DeLong & Vasseur 2012)", "previouslyFormattedCitation" : "(DeLong & Vasseur 2012)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(DeLong & Vasseur 2012).We can here conclude that there is good reason to use a clade-specific model in studies that wish to estimate population densities of mammals from body mass, as have been suggested for other animal phyla. For example, studies using allometric relationships across phylogenetic groups could improve fit and predictions by incorporating phylogenetic differences in the relationship of their fit. Studies such as these focus primarily on megafauna effects since megafauna may have greater impacts on nutrient cycling because of their larger movement ranges and gut passage times ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1371/journal.pone.0071352", "ISSN" : "19326203", "PMID" : "23951141", "abstract" : "Animals translocate nutrients by consuming nutrients at one point and excreting them or dying at another location. Such lateral fluxes may be an important mechanism of nutrient supply in many ecosystems, but lack quantification and a systematic theoretical framework for their evaluation. This paper presents a mathematical framework for quantifying such fluxes in the context of mammalian herbivores. We develop an expression for lateral diffusion of a nutrient, where the diffusivity is a biologically determined parameter depending on the characteristics of mammals occupying the domain, including size-dependent phenomena such as day range, metabolic demand, food passage time, and population size. Three findings stand out: (a) Scaling law-derived estimates of diffusion parameters are comparable to estimates calculated from estimates of each coefficient gathered from primary literature. (b) The diffusion term due to transport of nutrients in dung is orders of magnitude large than the coefficient representing nutrients in bodymass. (c) The scaling coefficients show that large herbivores make a disproportionate contribution to lateral nutrient transfer. We apply the diffusion equation to a case study of Kruger National Park to estimate the conditions under which mammal-driven nutrient transport is comparable in magnitude to other (abiotic) nutrient fluxes (inputs and losses). Finally, a global analysis of mammalian herbivore transport is presented, using a comprehensive database of contemporary animal distributions. We show that continents vary greatly in terms of the importance of animal-driven nutrient fluxes, and also that perturbations to nutrient cycles are potentially quite large if threatened large herbivores are driven to extinction.", "author" : [ { "dropping-particle" : "", "family" : "Wolf", "given" : "Adam", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Doughty", "given" : "Christopher E", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Malhi", "given" : "Yadvinder", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "PLoS ONE", "id" : "ITEM-1", "issue" : "8", "issued" : { "date-parts" : [ [ "2013", "1" ] ] }, "page" : "e71352", "title" : "Lateral diffusion of nutrients by mammalian herbivores in terrestrial ecosystems", "type" : "article-journal", "volume" : "8" }, "uris" : [ "" ] }, { "id" : "ITEM-2", "itemData" : { "DOI" : "10.1038/ngeo1895", "ISBN" : "1752-0894", "ISSN" : "1752-0894", "abstract" : "n the late Pleistocene, 97 genera of large animals went extinct, concentrated in the Americas and Australia1. These extinctions had significant effects on ecosystem structure2, seed dispersal3 and land surface albedo4. However, the impact of this dramatic extinction on ecosystem nutrient biogeochemistry, through the lateral transport of dung and bodies, has never been explored. Here we analyse this process using a novel mathematical framework that analyses this lateral transport as a diffusion-like process, and we demonstrate that large animals play a disproportionately large role in the horizontal transfer of nutrients across landscapes. For example, we estimate that the extinction of the Amazonian megafauna decreased the lateral flux of the limiting nutrient phosphorus by more than 98%, with similar, though less extreme, decreases in all continents outside of Africa. This resulted in strong decreases in phosphorus availability in eastern Amazonia away from fertile floodplains, a decline which may still be ongoing. The current P limitation in the Amazon basin may be partially a relic of an ecosystem without the functional connectivity it once had. We argue that the Pleistocene megafauna extinctions resulted in large and ongoing disruptions to terrestrial biogeochemical cycling at continental scales and increased nutrient heterogeneity globally.", "author" : [ { "dropping-particle" : "", "family" : "Doughty", "given" : "Christopher E.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Wolf", "given" : "Adam", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Malhi", "given" : "Yadvinder", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Nature Geoscience", "id" : "ITEM-2", "issue" : "9", "issued" : { "date-parts" : [ [ "2013" ] ] }, "page" : "761-764", "publisher" : "Nature Publishing Group", "title" : "The legacy of the Pleistocene megafauna extinctions on nutrient availability in Amazonia", "type" : "article-journal", "volume" : "6" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Doughty et al. 2013; Wolf et al. 2013)", "plainTextFormattedCitation" : "(Doughty et al. 2013; Wolf et al. 2013)", "previouslyFormattedCitation" : "(Doughty et al. 2013; Wolf et al. 2013)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Doughty et al. 2013; Wolf et al. 2013). Models that do not account for phylogenetically varying relations neglect the effect of larger biomass consumption rates of larger species, since the classical size-density model …. nutrient flux is thought to be offset by lower population densities. Our findings suggest an even larger impact with increasing body mass, possibly because larger species monopolise proportionally more of the energy in a system ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1038/324248a0", "ISBN" : "0028-0836", "ISSN" : "0028-0836", "PMID" : "94", "abstract" : "We present data and analyses demonstrating that large species utilize a disproportionately large share of the resources within local ecosystems. Even though small species tend to have higher local population densities, these are not sufficient to compensate for their lower rates of ...", "author" : [ { "dropping-particle" : "", "family" : "Brown", "given" : "James H.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Maurer", "given" : "Brian a.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Nature", "id" : "ITEM-1", "issued" : { "date-parts" : [ [ "1986" ] ] }, "page" : "248-250", "title" : "Body size, ecological dominance and Cope's rule", "type" : "article", "volume" : "324" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Brown & Maurer 1986)", "plainTextFormattedCitation" : "(Brown & Maurer 1986)", "previouslyFormattedCitation" : "(Brown & Maurer 1986)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Brown & Maurer 1986).Our conclusion that species-level population energy use increases with body mass is similar to what has previously been found in invertebrate communities ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1890/13-0620.1", "ISBN" : "0012-9658", "ISSN" : "00129658", "PMID" : "24669745", "abstract" : "Ecological communities consist of small abundant and large non-abundant species. The energetic equivalence rule is an often-observed pattern that could be explained by equal energy usage among abundant small organisms and non-abundant large organisms. To generate this pattern, metabolism (as an indicator of individual energy use) and abundance have to scale inversely with body mass, and cancel each other out. In contrast, the pattern referred to as biomass equivalence states that the biomass of all species in an area should be constant across the body-mass range. In this study, we investigated forest soil communities with respect to metabolism, abundance, population energy use, and biomass. We focused on four land-use types in three different landscape blocks (Biodiversity Exploratories). The soil samples contained 870 species across 12 phylogenetic groups. Our results indicated positive sublinear metabolic scaling and negative sublinear abundance scaling with species body mass. The relationships varied mainly due to differences among phylogenetic groups or feeding types, and only marginally due to land-use type. However, these scaling relationships were not exactly inverse to each other, resulting in increasing population energy use and biomass with increasing body mass for most combinations of phylogenetic group or feeding type with land-use type. Thus, our results are mostly inconsistent with the classic perception of energetic equivalence, and reject the biomass equivalence hypothesis while documenting a specific and nonrandom pattern of how abundance, energy use, and biomass are distributed across size classes. However, these patterns are consistent with two alternative predictions: the resource-thinning hypothesis, which states that abundance decreases with trophic level, and the allometric degree hypothesis, which states that population energy use should increase with population average body mass, due to correlations with the number of links of consumers and resources. Overall, our results suggest that a synthesis of food web structures with metabolic theory may be most promising for predicting natural patterns of abundance, biomass, and energy use.", "author" : [ { "dropping-particle" : "", "family" : "Ehnes", "given" : "Roswitha B.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Pollierer", "given" : "Melanie M.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Erdmann", "given" : "Georgia", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Klarner", "given" : "Bernhard", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Eitzinger", "given" : "Bernhard", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Digel", "given" : "Christoph", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Ott", "given" : "David", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Maraun", "given" : "Mark", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Scheu", "given" : "Stefan", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Brose", "given" : "Ulrich", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Ecology", "id" : "ITEM-1", "issue" : "2", "issued" : { "date-parts" : [ [ "2014" ] ] }, "page" : "527-537", "title" : "Lack of energetic equivalence in forest soil invertebrates", "type" : "article-journal", "volume" : "95" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Ehnes et al. 2014)", "plainTextFormattedCitation" : "(Ehnes et al. 2014)", "previouslyFormattedCitation" : "(Ehnes et al. 2014)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Ehnes et al. 2014), and support suggestions that the loss of larger predators can have greater ecosystem impacts than the loss of smaller species ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1086/679735", "ISBN" : "00030147", "ISSN" : "0003-0147", "PMID" : "25674690", "abstract" : "Trophic cascades are indirect positive effects of predators on resources via control of intermediate consumers. Larger-bodied predators appear to induce stronger trophic cascades (a greater rebound of resource density toward carrying capacity), but how this happens is unknown because we lack a clear depiction of how the strength of trophic cascades is determined. Using consumer resource models, we first show that the strength of a trophic cascade has an upper limit set by the interaction strength between the basal trophic group and its consumer and that this limit is approached as the interaction strength between the consumer and its predator increases. We then express the strength of a trophic cascade explicitly in terms of predator body size and use two independent parameter sets to calculate how the strength of a trophic cascade depends on predator size. Both parameter sets predict a positive effect of predator size on the strength of a trophic cascade, driven mostly by the body size dependence of the interaction strength between the first two trophic levels. Our results support previous empirical findings and suggest that the loss of larger predators will have greater consequences on trophic control and biomass structure in food webs than the loss of smaller predators.", "author" : [ { "dropping-particle" : "", "family" : "DeLong", "given" : "John P.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Gilbert", "given" : "Benjamin", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Shurin", "given" : "Jonathan B.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Savage", "given" : "Van M.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Barton", "given" : "Brandon T.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Clements", "given" : "Christopher F.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Dell", "given" : "Anthony I.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Greig", "given" : "Hamish S.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Harley", "given" : "Christopher D. 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Our results therefore also support the use of a scaling coefficient of -0.58, lower than the classical -0.75 ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1038/ngeo1895", "ISBN" : "1752-0894", "ISSN" : "1752-0894", "abstract" : "n the late Pleistocene, 97 genera of large animals went extinct, concentrated in the Americas and Australia1. These extinctions had significant effects on ecosystem structure2, seed dispersal3 and land surface albedo4. However, the impact of this dramatic extinction on ecosystem nutrient biogeochemistry, through the lateral transport of dung and bodies, has never been explored. Here we analyse this process using a novel mathematical framework that analyses this lateral transport as a diffusion-like process, and we demonstrate that large animals play a disproportionately large role in the horizontal transfer of nutrients across landscapes. For example, we estimate that the extinction of the Amazonian megafauna decreased the lateral flux of the limiting nutrient phosphorus by more than 98%, with similar, though less extreme, decreases in all continents outside of Africa. This resulted in strong decreases in phosphorus availability in eastern Amazonia away from fertile floodplains, a decline which may still be ongoing. The current P limitation in the Amazon basin may be partially a relic of an ecosystem without the functional connectivity it once had. We argue that the Pleistocene megafauna extinctions resulted in large and ongoing disruptions to terrestrial biogeochemical cycling at continental scales and increased nutrient heterogeneity globally.", "author" : [ { "dropping-particle" : "", "family" : "Doughty", "given" : "Christopher E.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Wolf", "given" : "Adam", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Malhi", "given" : "Yadvinder", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Nature Geoscience", "id" : "ITEM-1", "issue" : "9", "issued" : { "date-parts" : [ [ "2013" ] ] }, "page" : "761-764", "publisher" : "Nature Publishing Group", "title" : "The legacy of the Pleistocene megafauna extinctions on nutrient availability in Amazonia", "type" : "article-journal", "volume" : "6" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Doughty et al. 2013)", "manualFormatting" : "(e.g. used in Doughty et al. 2013)", "plainTextFormattedCitation" : "(Doughty et al. 2013)", "previouslyFormattedCitation" : "(Doughty et al. 2013)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(e.g. used in Doughty et al. 2013). Therefore, studies that aim to estimate the impacts of megafauna loss using the classic Damuth model must be overly conservative in their estimates, since Damuth’s model underestimates population sizes of large species. Our model provides a new method of relevancy for all studies predicting species densities on large scales. Our multi-level and -slope model is a substantial improvement over a single-slope model, where even the existence of predictive power has been questioned ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1111/j.1466-8238.2012.00782.x", "ISBN" : "1466-8238", "ISSN" : "1466822X", "abstract" : "Energy equivalence, the notion that population energy flux is independent of body mass, has become a key concept in ecology. We argue that energy equivalence is not an ecological \u2018rule\u2019, as claimed, but a flawed concept beset by circular reasoning. In fact, the independence of mass and energy flux is a null hypothesis. We show that our mechanistic understanding of size\u2013density relationships (SDRs) follows directly from this null model and the assumption that energy limits abundance. Paradoxically, without this assumption energy equivalence has no meaning and we lack a mechanistic understanding for SDRs. We derive an expression for the strength (r2) of SDRs under the null model, which provides a framework within which to compare published SDRs. This confirms that tight correlations between mass and abundance are a trivial consequence of the span of body masses considered. Our model implies that energy flux varies by five to six orders of magnitude among similarly sized mammals and to a far greater extent in birds. We conclude that the energetic paradigm can be strengthened by considering alternative, non-energetic, hypotheses.", "author" : [ { "dropping-particle" : "", "family" : "Isaac", "given" : "Nick J B", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Storch", "given" : "David", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Carbone", "given" : "Chris", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Global Ecology and Biogeography", "id" : "ITEM-1", "issued" : { "date-parts" : [ [ "2013" ] ] }, "page" : "1-5", "title" : "The paradox of energy equivalence", "type" : "article-journal", "volume" : "22" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Isaac et al. 2013)", "plainTextFormattedCitation" : "(Isaac et al. 2013)", "previouslyFormattedCitation" : "(Isaac et al. 2013)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Isaac et al. 2013). We encourage future studies to not assume a single slope across all mammals, but rather to use appropriate slopes for the specific clades, as our results show these to have better predictive ability. The same may well apply to other organism groups. Models for better estimates of natural population densities are vital for our understanding on global change in ecosystem function; our model here is a good first step. In future studies further improvement of predictive ability could be made by including other important factors, e.g. energy availability ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "DOI" : "10.1126/science.1067994", "ISSN" : "00368075", "author" : [ { "dropping-particle" : "", "family" : "Carbone", "given" : "C.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" }, { "dropping-particle" : "", "family" : "Gittleman", "given" : "John L.", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "container-title" : "Science", "id" : "ITEM-1", "issue" : "5563", "issued" : { "date-parts" : [ [ "2002", "3", "22" ] ] }, "page" : "2273-2276", "title" : "A Common Rule for the Scaling of Carnivore Density", "type" : "article-journal", "volume" : "295" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Carbone & Gittleman 2002)", "plainTextFormattedCitation" : "(Carbone & Gittleman 2002)", "previouslyFormattedCitation" : "(Carbone & Gittleman 2002)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Carbone & Gittleman 2002).SummaryClassic models for estimating population density from body mass on a global scale often overlook important internal structure to this relationship. The relationship varies among phylogenetic groups, and notably is consistently shallower within phylogenetic groups. Overall, this pattern is inconsistent with the energetic equivalence rule, but also suggests size-asymmetric monopolization of resources within groups , thereby also offering an explanation for Cope’s rule ADDIN CSL_CITATION { "citationItems" : [ { "id" : "ITEM-1", "itemData" : { "author" : [ { "dropping-particle" : "", "family" : "Rensch", "given" : "Bernhard", "non-dropping-particle" : "", "parse-names" : false, "suffix" : "" } ], "id" : "ITEM-1", "issue" : "3", "issued" : { "date-parts" : [ [ "1948" ] ] }, "page" : "218-230", "title" : "Histological Changes Correlated with Evolutionary Changes of Body Size Author ( s ): Bernhard Rensch Published by : Society for the Study of Evolution Stable URL : .", "type" : "article-journal", "volume" : "2" }, "uris" : [ "" ] } ], "mendeley" : { "formattedCitation" : "(Rensch 1948)", "plainTextFormattedCitation" : "(Rensch 1948)" }, "properties" : { "noteIndex" : 0 }, "schema" : "" }(Rensch 1948), the evolutionary tendency towards larger body mass within phylogenetic lineages. Further, our study shows that the use of group-specific density estimates should be used in studies that estimate densities from body mass, and that earlier studies likely have underestimated the densities of large-bodied species and thus their ecological effects.AcknowledgementsR?P and JCS were supported by the European Research Council (ERC-2012-StG-310886-HISTFUNC). SF was supported by the Danish Natural Science Research Council (#4090-00227). JCS further see this work as a contribution to his Carlsberg Foundation Semper Ardens project MegaPast2Future (CF16-0005).Author Contributions statementAll authors conceived the ideas and designed methodology; R?P collected the data; R?P and SF analysed the data; R?P led the writing of the manuscript. All authors contributed critically to the drafts and gave final approval for publication.ReferencesADDIN Mendeley Bibliography CSL_BIBLIOGRAPHY Agutter, P.S. & Wheatley, D.N. (2004) Metabolic scaling: consensus or controversy? 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Lynx Edicions.Wilson, D. & Reeder, D. (2005) Mammal Species of the World: A Taxonomic and Geographic Reference, 3rd ed (eds DE Wilson and DM Reeder). Johns Hopkins University Press, Baltimore.Wolf, A., Doughty, C.E. & Malhi, Y. (2013) Lateral diffusion of nutrients by mammalian herbivores in terrestrial ecosystems. PLoS ONE, 8, e71352.FiguresFigure 1: Model selection framework. We start out by the base model, and incrementally add new phylogenetic groups as either interaction (changing the specific clades slope) of main effect (changing the specific clades intercept), and keep whichever model improves AICc most (at least by 4). After this we remove any previous term from the model which no longer improves the model AICc by at least 4. When no additions further improve the model it is terminated, and this is considered our final model.Figure 2: The size-density relationship of mammalian species on a log10-log10 scale. The black dashed line indicates the general trend across all taxa, while the coloured lines indicate all the fits for the phylogenetic model. The 16 colours for the lines and points indicate different fit clades (For a coloured taxonomic reference see Figure S4). Figure 3: The estimated population density per species for the classical model and the phylogenetically structured model on a log10-log10 scale. Points above the black line are species with densities predicted to be larger than predicted in the classic model, while points under the line are species with lower predicted densities than the classic model. For clarity, only orders with more than 10 species sampled are displayed in colour; the rest are black. SupplementaryAppendix S1: Case study showing why density estimates can be biased against large species. Table S2: Final model. Estimates of intercept, main effects (offsets in intercept), interaction terms (i.e. changes in slope; denoted Mass:Group.x), Standard Error (SE), and ΔAICc. ΔAICc is the change from the best mode with all terms, and the term removed. See Table S3 for group definitions.TermsEstimateSEt-valuep-valueΔAICc(Intercept)3.150.0841.51< 0.0001-log10_Mass-0.580.02-28.33< 0.0001-Group.1-1.030.11-9.37< 0.000183.22Group.20.580.134.52< 0.000118.51Group.30.690.079.23< 0.000180.86Group.4-1.080.24-4.50< 0.000118.36Group.5-1.020.37-2.790.00545.83Group.60.370.152.470.01384.09Mass:Group.70.200.037.62< 0.000155.19Mass:Group.80.620.105.96< 0.000133.38Mass:Group.90.250.054.74< 0.000120.55Mass:Group.100.500.114.45< 0.000117.86Mass:Group.110.400.0410.16< 0.000197.53Mass:Group.120.090.023.650.000311.36Mass:Group.13-0.140.04-3.090.00207.61Mass:Group.141.020.214.77< 0.000120.83Mass:Group.150.220.063.500.000510.31Table S3: A table of which families are modelled under group names. These are families our model selection found should be fitted on their own, either in slope or intercept. Note that Mustelidae occur in Groups 1 and 4. Mustelids have to occur in group 1 since all groups are monophyletic, but also have to occur in group 4 since it has a significantly different intercept than the rest of group 1. See Fig. S4 for taxonomic tree with groups indicated.Intercept changed familiesGroup.1Canidae, Eupleridae, Felidae, Herpestidae, Hyaenidae, Mephitidae, Mustelidae, Nandiniidae, Procyonidae, Ursidae, ViverridaeGroup.2Cheirogaleidae, Daubentoniidae, Indridae, Lemuridae, LepilemuridaeGroup.3Abrocomidae, Aplodontiidae, Bathyergidae, Capromyidae, Caviidae, Chinchillidae, Ctenomyidae, Cuniculidae, Dasyproctidae, Echimyidae, Erethizontidae, Gliridae, Hystricidae, Myocastoridae, Octodontidae, SciuridaeGroup.4MustelidaeGroup.5Castoridae, Geomyidae, HeteromyidaeGroup.6Leporidae, OchotonidaeSlope change familiesGroup.7Acrobatidae, Burramyidae, Hypsiprymnodontidae, Macropodidae, Petauridae, Phalangeridae, Phascolarctidae, Potoroidae, Pseudocheiridae, Tarsipedidae, VombatidaeGroup.8TalpidaeGroup.9Chrysochloridae, Elephantidae, Macroscelididae, ProcaviidaeGroup.10OchotonidaeGroup.11Calomyscidae, Cricetidae, Muridae, Nesomyidae, SpalacidaeGroup.12CercopithecidaeGroup.13Felidae, ViverridaeGroup.14Geomyidae, HeteromyidaeGroup.15Mephitidae, Mustelidae, ProcyonidaeFigure S4: The phylogeny we used. Dichotomies which led to changes in the size-density relationship are marked with their group-number (see Table S2 and S3). Red indicates slope change, and blue indicates intercept change. Coloured points at the family level indicate which group the family is fitted together with (see Figure 2 in the manuscript).Table S5: Cross-validation estimates and results for final model and base model. Mean estimates of intercepts, main effects (offsets in intercept), interaction terms (i.e. changes in slope; denoted Mass:Group.x), R2, and ΔAICc, is here given for a cross-validation with a 1000 random subsamples of half the data tested against the remaining half, as well as standard deviations (SD), and ranges. See Table S3 for group definitions.TermsMean estimateSDMinimumMaximum(Intercept)3.150.082.863.40log10_Mass-0.580.02-0.65-0.49Group.1-1.020.11-1.38-0.55Group.20.590.090.290.88Group.30.690.090.360.95Group.4-1.080.22-2.16-0.24Group.5-1.010.24-1.790.02Group.60.380.18-0.561.04Mass:Group.70.200.020.120.27Mass:Group.80.630.150.050.95Mass:Group.90.260.11-0.050.65Mass:Group.100.500.110.141.03Mass:Group.110.400.040.260.53Mass:Group.120.090.020.020.15Mass:Group.13-0.140.04-0.26-0.02Mass:Group.141.010.160.391.54Mass:Group.150.220.050.030.52Predictive-R20.730.020.680.79(Intercept) – Base model3.870.073.614.10log10_Mass – Base model-0.740.02-0.81-0.66Predictive-R2 – Base model0.560.020.510.64ΔAICc225.5421.71288.25143.43 ................
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