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    Blackburn97 Critical Assessment

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    A Critical Assessment of the Form of the Interspecific Relationship between Abundance and Body Size in Animals Tim M. Blackburn; Kevin J. Gaston The Journal of Animal Ecology, Vol. 66, No. 2 (Mar., 1997), 233-249. Stable URL hutp:/Mlinks jstor.org/sicisici=002 1-8790% 28 199703% 2966%3A2% 3C233%3A ACAOTP%3E2,0,CO%3B2-R ‘The Journal of Animal Ecology is currently published by British Ecological Society. ‘Your use of the ISTOR archive indicates your acceptance of JSTOR’s Terms and Conditions of Use, available at hup:/www,jstororglabout/terms.hml. ISTOR’s Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at hupulwww.jstor.orgijournals/briteco.huml. Each copy of any part of @ JSTOR transmission must contain the same copyright notice that appears on the sereen or printed page of such transmission. STOR is an independent not-for-profit organization dedicated to creating and preserving a digital archive of scholarly journals. For more information regarding JSTOR, please contact support @jstor.org, hupslwww jstor.org/ ‘Thu Sep 23 01:03:08 2004 Journal of Animal Ecology 1997, 66, 233-249 (© 1997 Brits Ecological Society A critical assessment of the form of the interspecific relationship between abundance and body size in animals TIM M. BLACKBURN* and KEVIN J. GASTONT *NERC Centre for Population Biology, Imperial College at Siwood Park, Ascot, Berkshire SLS 7PY, UK: and {Department of Animal and Plant Sciences, University of Sheffield, Sheffield S10 2TN, UK ‘Summary 1. Despite a wealth of studies for a wide variety of animal assemblages, the form of the interspecific relationship between abundance and body size is still contentious. At least three different patterns have been suggested, which can broadly be characterized ‘as negative and linear, negative but non-linear, and polygonal. At least eight different mechanisms have been suggested whereby the linear (or non-linear) and polygonal patterns can be reconciled. 2. We collated data from the literature on over $00 interspecific plots of the abun- dance-body size relationship with two aims, First, to examine the extent to which published studies support the different forms proposed for the relationship; and secondly, to test whether any of the mechanisms that have been suggested to reconcile linear and polygonal patterns actually do so. 3. The data revealed that abundance-body size relationships commonly assume both linear negative and polygonal forms. Around 25% of all plots of the relationship show a positive regression slope. 4, Of the eight mechanisms that have been suggested to reconcile linear and polygonal patterns, we were able to test five. OF these, only the measure of density used by a study explains none of the observed variation in abundance-body size relationships. Variation in the regression slope between studies is only explained by the type of data used (compilations vs. samples) and the scale of study (local vs. regional): compilation studies at regional scales show more linear negative relationships, while sample studies, performed at local scales show more polygonal patterns. General linear modelling indicates that study scale is the most important factor influencing when different relationships are likely to arise. 5. Our results show that different patterns tend to arse at different scales of study, but say nothing about whether the patterns are real or artefactual. Polygonal relationships potentially contain an artefactual component, resulting from sampling methodology inadequate to elucidate the abundances of the less common species in any given assemblage. However, the presence of a sampling artefact does not indicate the shape of the underlying relationship from which a sample is taken, or indeed whether samples would exhibit any other shape were artefacts excluded, 6. Linear and polygonal patterns are not necessarily mutually exclusive, but may both indicate the ‘true’ abundance-body size relationship at different spatial scales. We conclude by suggesting that much more attention be paid to the effect of spatial scale on this relationship, especially given that scale of measurement can have subtle but, severe consequences. Key-words: abundance, allometry, animals, body size, macroecology. Journal of Animal Ecology (1997), 66, 233-249 the interactions that determine the distribution and Introduction abundance of organisms (Krebs 1972; after Andre Ecology has been defined as the scientific study of wartha 1961). Therefore a considerable body of bio 23 24 Animal abundance-body size relationships (© 1997 British Ecological Society Journal of Animal cology, 6, 233-249 logical literature has been devoted to documenting, patterns of distribution and abundance, and to identi- fying the mechanisms that generate those patterns (see reviews in Krebs 1972; Begon, Harper & Townsend 1990; Gaston 1994; Brown 1995), as an early step on the road to understanding how the great diversity of life is structured. A. good proportion ofthis literature has focused on the identification of characteristics of ‘organisms that are associated with the abundance and distribution they attain, in the hope that association may to Some degree reflect causation. In animal ecol- ‘ogy, one wssociation that has received particular atten tion, at least since it was given prominence by Elton (1927), is the interspeciic relationship between body size and abundance in animals, ‘The relationship between abundance and body size thas been one of the most extensively studied inter- specific patterns in ecology (see reviews in Cotgreave 1993; Blackburn & Lawton 1994). In excess of 50 papers have dealt with the subject, and the relation- ship has been documented for species from most ‘major environments and taxa. Unfortunately, this has not led to a consensus over the nature of the under lying patterns, let alone the mechanisms which gen- erate them. These two issues are tightly bound, ‘because the mechanism favoured will depend on the relationship iti required to explain. Logically, a small-bodied species has the capacity to be more abundant than one of large siz. The upper limit on possible abundance is set by the number of individuals that ean physically be packed into a given area (or volume for species living in three dimensions: eg. Nee, Harvey & Cotgreave 1992). In practice, this maximum density is never (or perhaps seldom) achieved because resource requirements cannot be satisfied under such extremes of packing. Never- theless, since resource requitements scale positively with body size (Peters 1983; Calder 1984; Reiss 1989), the maximum density attainable by species should scale negatively with size, ‘There are also reasons to believe thatthe minimum density a species can sustain scales negatively with body size (Lawton 1989, 1990). At the limit, no sexual species ean exist when so widely dispersed that finding ‘amate and reproducing ate unlikely events within the lifetime of an individual. In most taxa, large-bodied species are more vagile and longer-lived than small bodied (Lindstedt & Calder 1981; Calder 1984; Read ‘& Harvey 1989; Suther 1989), so they should be able to persist at loner densities, Populations of large: bodied species may also fluctuate less, so that large> bodied species have lower minimum viable population sizes (Pimm 1991; Lawton 1994). In combination with, the expectation that maximum densities scale nega tively with body size, the general relationship between body size and density should therefore also be nega- tive. A number of studies support this prediction, when both body size and density are logarithmically ‘wansformed (Damuth 1981, 1987; Peters & Raelson 1984; Marquet, Navarette & Castilla 1990; Nee et al, 1991; Peters & Wassenberg 1993; Strayer 1994; Ebenman et a. 1995; Fig 1a). In other words, density (D) scales with body size (M) as D a constant and yis negative Many other studies have found relationships that are not simply negative and linear. For example, Brown & Maurer (1987) calculated the abundances of land birds, using data from the North American Breeding Bird Survey. Plotted against Body mass, they produce a complex ‘polygonal’ relationship (Fig. 1b). Maximum abundance peaks at intermediate, not the smallest, body masses, while minimum abundance is constant across all masses, Similar relationships have been shown ina wide variety of taxa (e.g. Morse, Stork & Lawton 1988; Gaston 1988; Basset & Kitching 1991; Novotny 1992; Blackburn er al, 1993a; Cotgreave, Middleton & Hill 1993; Blackburn & Lawton 1994; Cambefort 1994; Nilsson, Elmberg & Sjoberg 1994) While the precise details difer between studies, poly- -gonal relationships are typically characterized by the absence of a marked increase in the minimum abun: dance of small-bodied species, and by 2 low cor relation between abundance and size. Frequently, M?, where eis 10 10 10° 10 10 10° Density (km) 10" 10 10 107 0 10 10 a0" 10" 100 |” . 10 Density (individualsroute") Body mass (a) Fig 1. The relationship between body mass (g) and (a) den- sity (numbers per kn = 467) fora selection of mammals ofthe world (Irom Damuth 1987) and (6) deasiy (ndi- Viduas per BBS route, n = 380) for North American land binds (Brown & Maurer 1987). 235 TM, Blackburn & KJ.Gaston © 1999 British Ecological Society Jounal of Animal Ecology, 66,205 249 peak abundance is at intermediate body sizes, and sometimes the overall relationship is actually positive. Recently, evidence has been presented for a thitd pattern in abundanee-body size relationships that combines Some features ofthe previous two (Damuth 1993; Marquet, Navarette & Castilla 1995; Silva & Downing 1995). This is derived from work on mam- ‘malian primary consumers and suggests that the relationship is essentially negative, but non-linear, density generally increases as body mass decreases, ‘but peak abundance is attained between 0-1 and Ike, ‘not by species of minimum mass (this feature is actu ally shown by Fig. a). Although the slope ofthe over all relationship is 0-75, slopes plotted within taxa tend to be shallower for those with species generally smaller than kg, and steeper for taxa with species generally heavier than kg. Species near 1 kg attain higher densities than expected from a —0-75relation- ship, altering the stope within the taxa to which they ‘belong (Damuth 1993) Silva & Downing (1995) show similar effects for mammalian herbivore, insectivores and carnivores, In addition, they find thatthe lowest density isnot attained by the largest species; LOWESS. regression indicates that the slope levels out at large sizes. The reason is unclear, although Silva & Down- ing suggest a number of possibilities, including that large mammals may make more efficient use of resources, or that very low density populations may simply not be viable. Reconciling the different relationships In sum, despite strong theoretical reasons for expect- ing a negative abundance-body size relationship, the evidence for such a pattern seems ambiguous. Bight mechanisms have been proposed to explain why different relationships have been obtained, and in par ticular to reconcile why linear and polygonal patterns may both be found. Different patterns have been hypothesized to result 1. From comparisons across different kinds of data (Lawton 1989). In general, simple linear negative relationships are obtained for data compiled from a ‘wide variety of literature sources. These sources are often single- or few-species studies from a range of ‘Beographic areas, and densities may have been esti- mated using @ wide variety of methodologies. Con- versely, polygonal relationships are generally re ‘covered when single areas are sampled to give abundance estimates for all the species ina given taxon. that are present, usually using a single or consistent method. 2, From samples at different spatial scales (Lawton 1989, 1990). Polygonal relationships end to arse from studies performed at local or restricted sites (es Morse eal. 1988), whereas linear negative relation- ships tend to arise as the geographic coverage or geo- graphic spread of data increases, 3. From differences in the density measures used (Damuth 1987, 1991; Lawton 1989). Linear negative relationships are typically characterized as arising from plots of ecological densities, and polygonal ‘relationships from crude densities. Ecological density is generally defined as the density of a species in suit- able habitat, and crude density as the density of a species in a sample from a given area (e.g. Damuth 1981, 1987; Gregory & Blackburn 1995; Silva & Downing 1995). 4. Because of differences in the ranges of variation represented by the species included in the plots (Law- ton 1989), Linear negative relationships ate typically ‘obtained from plots spanning several orders of mag nitude of body sizes or abundances, whereas the ranges spanned by species in polygonal relationships are typically, although not always, less. Polygonal relationships are therefore often suggested to be simply segments of an overall linear negative relationship, which would be recovered if the range of body sizes in the polygonal relationship was increased. 5, Because polygonal relationships are random samples from an underlying abundance-body size relationship (Blackburn, Harvey & Pagel 1990; Blackburn, Lawton & Pimm 19936; Currie 1993). Blackburn etal (1990) showed that a polygonal relationship between abun- dance and body size could arise asa result of random sampling of individuals from an underlying dis- Uribution where there was no relationship between abundance and body size (See also Blackburn er al 1993b). Currie (1993) gave visual, although not sta- tistical, evidence that a polygonal relationship could be generated by random samples from a linear nega- tive relationship, within the same restricted range of body sizes. 6, Because of differences in how taxa utilize space (Guanes 1986). In some cases, a polygonal relationship may be recovered if species in a taxon utilize the ‘environment in three spatial dimensions. Since den sites are generally measured in terms of areas (atleast in the terrestrial environment), the linear negative relationship between density and body size may break down if species are actually utilizing diferent volumes of space. That birds are perceived generally to show weaker abundance-body size relationships than mam- mals has been argued to result from this effect Juanes 1986; Cotgreave & Harvey 1992) 7. Depending on the taxonomic composition of the assemblage (Nee et al 1991; Silva & Downing 1995). Abundance body mass relationships are often posi= tive within taxa at low taxonomic levels (eg. tribes for genera) (Nee ef al, 1991; Blackburn et al. 1994), Further, analyses of mammalian assemblages show non-linear relationships, so that large-bodied species hhave densities that are higher than expected, and smalk-bodied species have densities that are lower (Damuth 1993; Silva & Downing 1995). Analyses of closely related species, or mammal assemblages com- ‘posed entirely of small-bodied or large-bodied species, ‘might then be expected to show a non-significant, or 236 Animal abundance-body size relaionships (© 1997 British cological Society Journal of Animal cology, 66, 233-249, even positive, relationship, even if completely sampled, because that is the underlying global relationship for that range of body sizes, 8. Because of the effect of migrant species (Damuth 1987; Cotgreave 1994b), The abundances of migrant species may be influenced by factors outside the area in which they are censused. Assemblages with high proportions of migrant species (principally bird assemblages) may therefore show weaker abundance body size relationships. Al cight mechanisms are plausible, albeit some more than others. However, no systematic evaluation of the alternatives has been undertaken, although the body of published studies is now more than adequate {o test at least some of them. Many of the mechanisms assert that a polygonal relationship arises as a cor- ruption of a negative linear relationship between abundance and body size. However, there has been no {quantitative assessment ofthe form of the relationship found in the studies so far published. In the remainder ‘of this paper, we examine the extent to which pub- lished literature supports the hypothesis of a negative relationship between abundance and body size, and test whether the mechanisms that have been suggested to reconcile different patterns actually do so, We have limited consideration to the relationship between abundance and body size across species; the question ‘of what mediates phylogenetic patterns inthe form of the relationship within taxa (e.g. Nee er al 1991, 1992; Cotgreave & Harvey 1992, 1994; Blackburn et al 1994; Cotgreave & Stockley 1994; Cotgreave 1994a,b, 1995) is an interesting, but separate, issue. Methods A review of the ecological literature yielded infor mation on over $00 different plots of the abundance— body size relationship, including over 300 for whieh ‘quantitative data on its form (eg. regression slope, coeficient of determination, sample size) were avail- able (data from Damuth 1981, 1987, 1993; Peters & ‘Wassenberg 1983; Juanes 1986; Robinson & Redford 1986; Strayer & Likens 1986; Brown & Maurer 1986, 1987; Gaston 1988; Macpherson 1989; Owen & Gil- bert 1989; Marquet ef al, 1990; Basset & Kitching 1991; Maurer, Ford & Rapoport 1991; Nee eta. 1991; Cotgreave & Harvey 1992; Griffiths 1992; Novotny 1992; Cotgreave er al. 1993; Currie & Fritz 1993; Blackburn er al, 1993a, 1994; Blackburn & Lawton 1994; Cambefort 1994; Cotgreave & Stockley 1994, Nilsson et al. 1994; Thiollay 1994; Cotgreave 1994a, 1995; Bbenman ef al. 1995; Gregory & Blackburn 1995; Silva & Downing 1995; Greenwood et al. 1996). ‘The relationships compiled included assemblages spanning a wide variety of taxa, trophic groups, geo- ‘graphic regions and habitats (Table 1), and from three {to 856 species (mean = 69). Some of the data sources ‘were previous compilations, and others included sum- maries of the form of the relationship for a large umber of local assemblages without Formally pre- senting the results for each one (e.g. Juanes 1986; Cotgreave & Harvey 1992; Cotgreave et al. 1993; Cotgreave & Stockley 1994). An exhaustive review of the literature would probably turn up many more studies. We categorized each estimate of the abundance body size relationship in an assemblage as either inde- pendent or dependent. The question of what con stitutes an independent estimate is dificult to answer ‘unambiguously. For example, Damuth (1987) pro- duced a plot ofthe abundance-body size relationship sing data from 667 species from taxa spanning vir- tually the entire range of the Kingdom Animalia. One could argue that since any other relationship must involve data that are a subset of the Animalia, all ‘others are a subset of this one, More problematically, many of the relationships plotted by Brown & Maurer (1986) for local bird assemblages use abundance esti- mates for subsets of the species in the abundanoe— body size plot for North American landbirds (Brown & Maurer 1987). We defined a relationship as inde- pendent if there was no reason to believe it was based principally on a subset of the actual data used for any other abundance-body size relationship in the literature. Under this definition, both examples above are actually independent; although the same species in the same geographic areas may appear in more than one relationship, itis unlikely that any of the relationships defined as independent are dependent on the form of any other. Examples of relationships that wwe defined as dependent on data from a more general plot include the ‘constructed communities’ presented. bby Damuth (1981), the dietary subsets of the Neo- tropical forest mammal relationship in Robinson & Redford (1986), and the dung beetle plots from sub- sites of the South African game reserve faunas pre- sented by Blackburn ef al. (1993a). Independent and dependent plots were considered separately in our analyses, Foreach abundance-body size relationship, we also recorded the following information, Relationships were separated on the basis of whether they used data compiled from a wide variety of litera- ture sources (compilation studies), or from samples of single areas (usually using a single or consistent method) giving abundance estimates forall the species in a given taxon or assemblage that inhabit the area (ample studies. We separated relationships on the basis of whether they were plotted at local or regional scales. In general, 1 relationship was considered to be plotted at the regional scale if it encompassed the fauna of a geo- 27 TM. Blackburn & KJ. Gaston (© 1997 Brith ological Society Journal of Anal Ecology, 66, 233-249 ‘Table. A summary of the characteristic of the 330 assemblages for which quantitative data on abundance-body size relationships were available. This tabulation includes dependent and independent estimates, Note that not ll assemblages can ‘be clasifed to one of the categories in each set, many include primary and secondary consumers), x0 that not all ume equal 330, Environment om Region ® Taxon nh Trophicevel ® “Tertesti 253 Tropical 100 Invertebrate 64 Scavenger 2 Aquatic 36 Temperate ISS Vertebrate «289 ‘Primary consumer 77 Both 7 Secondary consumer 37 North America 68 Omnivore 6 South America 39 Inset, a Europe 35 Fish 2 Aiea Th Bind ® ‘Asia 16 Mammal is ‘Australia 3 politcal unit (e.g. Ivory Coast, Sweden, ‘Basque coun- try) or was a compilation of data for taxa across several regions (eg. figure 1 in Damuth 1981). Ie was considered to be at the local scale if it encompassed the fauna of a restricted site or habitat (.g. Richmond Park in England, tropical grassland in Uganda), even if the data were compiled from wider sources (eg table I in Damuth 1981), Few studies raised the com- plications of intermediate scales. Relationships were separated on the basis of whether they used crude or ecological densities. Assigning density measure to one or other category can be dificult. Although they are frequently considered as if distinct dichotomous states, crude and ecological densities are better thought of as two ends of a spec trum that can be calculated for any given species, because ‘suitable habitat’ is not readily defined. We assumed that densities were erude if measured in a given area forall species (e.g. Morse et a 1988), and ecological if measured on a species-by-species basis (€g. Damuth 1981, unless the opinions ofthe authors were otherwise. BODY SIZE RANGE We recorded the range of body masses across which an bundance-body size relationship was plotted, using only those where body mass was the size measure, or Where length could be converted to mass using & length-mass regression (eg. Blackburn etal. 1993). This variable could not be normalized by trans formations, and so was analysed non-parametrcally. DIMENSIONALITY OF ASSEMBLAGE Relationships were categorized depending on whether the taxa use the environment in two of three dimen- sions. The latter category includes bird, tree canopy ‘and pelagic aquatic assemblages; the former category consists mainly of assemblages of non-volant ‘mammals, but includes several benthic fish, dung beetle and flightless bird assemblages, Relationships plotted with subsets of species that could be assigned to both, categories (eg, plots including non-volant mammals and bats) were excluded from analysis ofthis variable, ‘Simple linear negative relationships between abun= dance and body size differ from polygonal relation- ships in the amount of variance in abundance that is explained by body size, and in the magnitude of the regression slope. To test whether each of the above ‘mechanisms explains the difference between linear and polygonal relationships, we asked whether there were consistent differences inthe slopes and coefficients of determination (P-values) of relationships depending ‘on how they were classified. For example if linear and polygonal relationships result from different density measures, we would expect relationships plotted with crude densities to have lower coeficents of deter- mination and higher (Le. less steeply negative) slope values than relationships plotted with ecological den- sities. Only regression slopes calculated using the method of ordinary least squares (OLS, or model 1) ‘were used for the purposes of comparison. Coelicients of determination were square root transformed for analysis (note that the square root of # does not ‘equal r but does equal |), to conform better with the assumptions of OLS regression analysis, Ideally, we would have tested the relationship between body size the residuals from abundance-body size regression; residuals from polygonal and linear relationships should show different _ patterns However, data were available for too few of the abun- dance-body size plots to make this a practical option. and Results Qualitative information on the sign of the model 1 regression slope was available for 534 of the abun- ‘dance-body size relationships. Fourchundred and three relationships had negative slopes, 129 had posi= tive slopes and two slopes were exactly zero (as reported); therefore, 25% of these studies did not show negative relationships. Quantitative information 238 Animal ‘abundance-body size relationships (© 1997 British Ecological Society Journal of Animal Ecology, 66, 233-249, fon the model I regression slope was available for 303 of these relationships. The frequency histogram of the 291 slopes for which body size was measured as mass showed a pronounced peak between —0'5 and ~1-0, with a mean slope value of —0'51 (Fig. 2a). Thirteen per cent of these quantitative slopes were positive; if just the subset of 119 independent relationships was ‘considered, this rose to 23%. Quantitative information on the coefficient of deter- ‘mination was available for 326 studies. Body size most often explained little of the variance in abundance (mean + SE = 0:33 + 00017; Fig.2b); the same was ‘ruc if the sample was restricted to independent esti= mates (mean + SE =0-18 + 0-019). The magnitude of the coeficient was negatively correlated with the size of the assemblage (Spearman rank correlation, tho = ~0:18,n = 326, P = 0.0012), but this relation ship disappeared when analysis was restricted to inde- pendent estimates (ho = 002, n= 133, P= 084). In despite sound theoretical reasons for expecting a negative abundance-body size relationship, and a plethora of studies, the ability of body size to predict =r0 e 8 ? cteenanene | E of Fig 2. (a) The Frequency dstbution of model I regression slopes fom 291 plots of abundance against body mass. Hori= zontal axis labels indicate the lowest slope value in each histogram bar; for example, the fist ar includes studies with regression slopes from 4:25 to 40. (b) The frequency istibutionofeoeients of determination fom 326 pots of tlbundance against body mass. Horizontal axis abelsindicate the lowest coefficient value in each histogram bar in each case; for example, the frst bar includes studies with ‘coefcients from 0 to 004. population densities is generally weak, as has been noted several times before with smaller data sets (Blackburn eral. 19934; Blackburn & Lawton 1994; Blackburn, Lawton & Gregory 1996). Despite qualitatively similar distributions of regression slope signs, dependent and independent relationships showed quantitative differences, Slopes from independent estimates of the abundance-body size relationship were significantly shallower than those based on data subsets (Table 2). Body mass also explained different amounts of the variance in abundance when dependent and independent esti mates were used (Table 2). Further consideration of the diferent abundance-body size relationships is based on independent estimates, except where indi cated otherwise, Abundance-body size relationships based on com= pilation data had significantly higher coefficients of determination, and more steeply negative slopes, than relationships based on sample data (Table 3). In fact, the difference between independent and dependent ata in estimates ofthe slope and coefficient of deter- ‘mination of the abundance-body mass relationship (Table 2) disappeared once the data type used was taken into account (Fig.3). This was principally ‘because most (90%) independent estimates were from. sample data, and most (63%) dependent estimates used data subsets from wider compilations Regression slopes were significantly more steeply negative and coefficients of determination were sig- nificantly higher in studies performed at regional scales (Table 4), This was not a consequence of local studies predominantly using sample data; both scale ‘of study and data type explained significant amounts ‘of the variation in both regression slope and coeficient of determination (Fig. 4), Studies using crude and ecological densities showed no significant differences in either regression slopes or coefficients of determination ofthe relationships they reported (Table 5). The apparent relation of abun- dance-body size pattern to density measure seems more likely 0 arise because the density measure is itself related to differences in scale and data type. This was impossible to test using data type, because there was only one study that we considered to be a com- pilation using erude densities. However, when study scale was included in the analysis, studies using eco- logical densities showed significantly higher cocficients of determination than those using crude densities, but no significant difference in slope (Fig. 5). ‘The hypothesis that different patterns arise because of differences in the ranges of variation represented by the species received equivocal support, This hypothesis suggests that observed polygonal abun- dance-body size relationships are segments of an ‘underlying linear negative relationship. A small seg ‘ment ofa broader relationship is, on average, expected to have the same regression slope as the relationship {rom which it is a segment, but a lower coefficient of 239 TM, Blackburn & K.J.Gaston (© 1997 Bish Ecological Society ournal of Animal cology, 66,233-249, ‘Table2. OLS slope and |r| (y transform of) values from estimates ofthe abundance-body sie ionship defined as ether independent or dependent (ee Methods), together with the results (F, P)of an axovA testing for diferencesin the distsibutions ofthe respective statistics, Only relationships plotted using hody mass dita are incided in this analyse Mean (SE) ” Estimate Slope Independent 0835 + 00449 9 Dependent be Ie Independent 0367 + 0021 m2 Dependent 0-567 + 0.0254 1 F P 2034 0.0001, 3399 <0.0001 ‘Table3, OLS slope and | (transform of) values from estimates ofthe abundance-body size relationship using data ether from single stmples or from literature compilations (ace Methods), together with the results (F, P) ofan ANOVA testing for Aitfrences inthe distributions ofthe rexpetive statistics Estimate Mean (SE) ” Slope Sample 0201 + 0087 wor Compilation 0-729 + 0004 2 Ia Sample 0334 4.0020 109 Compilation 0-644 + 0088 B 0029 <00001 determination (Draper & Smith 1981). Why this isso is best illustrated by imagining successively smaller pieces snipped out of the relationship in Fig. 1a; the regression slope remains the same, but as the error variance in abundance approaches the variance in ody mass, the rectangle of points more and more closely approximates a blob. Increases in the range ‘of body sizes across which relationships are plotted should be accompanied by a smooth increase in the associated coeficient of determination but no change in the regression slope (Currie 1993) ‘There was indeed a general tendency for the coefficient of determination to increase withthe range ‘of body masses across which an abundance-body ‘mass relationship was calculated (Spearman rank cor- relation, 7, P= 0-009; Fig. 6a), as predicted. However, this was not the smooth inerease expected from theory (Currie 1993), as shown more clearly if body size range was divided into a number ‘of diserete classes (Fig. 6b). Further, regression slopes tended 10 decrease as the range of body masses increased (cho = —0'31,n = 74, P = 0.008), contrary to prediction; again this decrease was not smooth (Fig. 60), In some cases, a polygonal relationship has been hypothesized to result if species utilize the environ- ‘ment in three spatial dimensions, We examined two separate questions in relation to this mechanism. Do taxa that generally use the environment in wo dimen sions show quantitatively different abundance-body ‘ize relationships from those that use it in three? Do birds and mammals show quantitatively different abundance-body size relationships? We found some evidence that the dimensionality explanation does indeed contribute to variation in the abundance-body size relationship, Assemblages using a twordimensional environment had higher cocffcients of determination than assemblages using three, but their regression slopes were not more steeply negative (Table 6), The same pattern was obtained in the comparison of birds and mammals; the dis- tributions of OLS slopes estimated for birds and mam- mals were not significantly different, but mammal assemblages showed significantly higher coefficients of determination (Table 6). However, the similarity between the effects of dimensionality and taxon is not surprising, because most assemblages using the ‘environment in three dimensions consist of birds, and most assemblages using the environment in wo dimensions consist of mammals There was no difference inthe regression slopes of the abundance-body mass relationship between those assemblages or taxa considered to use the environ ‘ment in two or in three dimensions, but body mass was a better predictor of abundance in the former subset. The obvious explanation for similar slopes ‘but different coefficients of determination is that bird assemblages and assemblages using a three-dimen- sonal environment consist of species covering a smaller ange of body masses. Infact, bird assemblages cover a larger range of body masses than mammal assem- blages (Mann-Whitney U-test, z= 4964, n= 65, 240 Animal ‘abundance-body size relationships (© 1997 British Ecological Society Journal of Animal Ecology, 66, 233-249 ' @ E zg naapenenependent Fig.2. Mean (SE) OLS regression slope (circles) and coeficient of determination (diamonds) for independent (1) and dependent (D) relationships in stuies performed on sample or compilation data, There isa significant fet ofthe data type on both slope ((worlactor ANOVA, Fas = 142, P= 00002) and coeficient (Fi. = 853, P < 00001), but Mependency significantly affets neither (slope, Fis. ~ 1-0, P= 032; coofcient, Fis = 05, P=0.), and there i no significant interaction (lope, Fis4= 11, P= 0%: conficient, Fin = 17, P=02), Dependeney does. not explain variance in cither the slope or coeticient of deter- ‘mination in addition to that explained by datatype, Statistics were calculated using transformed coeficint of dter- P<00001), and assemblages using space in three dimensions span a larger range of masses than those using space in two dimensions (Mann-Whitney U- test, 2= 4:22, n= 74, P< 00001), Moreover, the diference in coeflcients was not just a consequence of diferencesin scale or the use of compilation compared with sampling data in the different assemblage types; ‘mammals and assemblages using a two-dimensional environment showed consistently higher coeficents when these factors were controlled for statistically (Table 7, ‘The results of our analyses examining the factors that have been suggested to affect the regression slope and coefficient of determination of the relationship between abundance and body size are summarized in Table 8. Regression slopes were only affected by study scale and data type, and these both explained inde- pendent amounts of the variation in slope when ana~ lysed together (Fig. 4). Coefficients of determination were affected by all the factors tested except density a ar zg ean cote ot etn TacsiFeglonel Local Reson Sele of tay Fig.4. Mean (ESE) OLS regression slope (circles) and ‘oefcent of determination (diamonds) for tlationships in studies performed at local and regional scales, using ether sample or compilation data. Both the scale of study andthe type of data used have significant effects on both the slope (eal, Fi) =62, P=OOL3; data ype, Fo =79 P= 0005)and thecoeficent (Scale Fnys= 8.3, P = 0.0043; data type, Fay, = 1493, P< 00001), Thee is ako a sige nifleant interaction beoween the effects of sale and datatype fon both the slope and the coeficent (lope, Fig = 7. P= 0005; coefcent, Fy = 181, P < 0001). The analy- sis was performed on both independent and dependent es mates ofthe relationship, because there were no independent ‘estimates at local seals using compilation data. Statistics were calculated using /-trnsformed coelcent of deter- measure, However, all these results are potentially affected by multicollinearty, as many of the variables Wwe tested will be closely related (eg. the dimen- sionalty of an assemblage and the taxon of which it is composed). We used general linear modelling to examine whether these relationships are independent, fr the result of correlation between the different factors, Body size range was log transformed t0 reduce its skewness. This approach showed that only study scale explains significant amounts of the vari- ation in the slope and coefficient of determination of abundanee-body mass relationships when all the factors listed in Table 8 are considered together (Table 9), From these maximal models, a stepwise backwards slimination procedure was used to produce minimal adequate models (Crawley 1993). For regression slope, this model contained only study scale ‘Tabled. OLS slope and |r (/ transform ofr) values from estimates ofthe abundance-body siz eclatonship using daa either from loeal or regional studies (see Methods), together withthe results (F,P) of an ANOVA testing for difleences in the distributions of the respective statistics timate P) " Slope cal 0245 + 0050 96 Regional 0682 = 0058 a te Local 0530 4 0021 98 Regional 0-485 +0055 2 F - 1636 <00001 887 0.0035 21 TM. Blackburn & KJ.Gaston (© 1997 British ecological Society Lournal of Anal Beology 66, 249 ‘TableS, OLS slope and |r| (transform of) vals from estimates of the abundance-body size relationship using either crude or ecological densits (sce Methods), together with the results (F, P) of an ANOVA testing for differences in the distibutions of the respective statistics Estimate Mean (+E) 1» Slope Crude Osis £0073 » Ecological 0382 + 0073 58 wi ‘Crude 0342 40037 20 Ecological 0-443 0035 0 Fr P 16 023 240 os (F ys = 1636, P <00001), but dimensionality also entered the equivalent model for the coefficient of determination (study scale, Faye = 512, P= 0026; dimensionality, F,jy¢= 8:19, P = 0.005). Discussion Eight factors have been suggested (0 cause, ot con- tribute to, the variation observed in the form of plots ‘of interspecific abundance-body size relationships for animals. Our compilation of data from published plots enabled us to test directly five of these, When considered in isolation, each factor we tested did to some degree affect the abundance-body size rlation- ship, with the exception of the measure of density employed. However, the significance of most factors apparently results because they ae related to variation, in the spatial scale of study. For example, studies at small seales tend {0 cover small ranges of body size ‘and use sample data, Spatial seale isthe only variable consistently correlated with variation in the abun- dance-body sie relationship, ‘The flute of the measure of density to influence Local Regional ean o.8 spe 8 Fig. 5. Mean (SE) OLS regression slope (cirles) and cotfcient of determination (diamonds) for independent relationships in studies performed at lea! and regional scales, using either erude (C) or eeologeal (E) densities. Scale of tidy hs significant effets on bath the slope (ea, Fi =86, P= 00085) and the cocticient (Fin. = #8, P0032), but density measure only has a significant effet fon the latter (slope, Fin, = 007, P= 08; coefficient, Fix=69, P=O001), Statist were calculated using ‘transformed coetfient of determination abundance-body size relationships is not surprising ‘As observed in the Methods, erude and ecological densities are actually just the two extremes of a con- tinuum of densities that can be determined fora given ‘numberof individuals depending on the area used for the calculation. Therefore, there will often be inter- gradation of these measures, and in many cases the distinction may be dlficult or arbitrary. For example, oth Damuth (1981) and Silva & Downing (1995) claim that their density measures are ecological, How- ever, many of their estimates, particularly for large ‘mammals, come from censuses of nature oF game reserves, performed across very large areas. One such source was Green (1979), who sampled large mammal populations in Upper Volta throughout # national ‘park of area 1000 km?, Whether the densities reported for, say, hippopotamus, Hippopotamus amphibius L. and warthog Phacochoerus aethiopicus (Pallas) are truly ecological, or even comparable, given the likely differences in how these species use the park area is ‘unknown, and probably not determinable from the original source. The problem of comparability per- vades the interspecific analysis of densities (Gaston 1994; Blackburn & Gaston 1996, 1997; Smallwood & ‘Schonewald 1996). Since the diflerence between crude and ecological densities is not absolute, but one of degree, itis unlikely that absolutely diferent abun- dance-body size patterns would result from attempts to classify studies as using either erude or ecological Currie (1993) makes two points in respect of density ‘measures. First, they will only affect the slope of the abundance-body size relationship i there is an inter- action between body size and the density measure used. Ithe ‘crudeness’ of the density measure used is random with respect to size, the slope estimate will be unbiased, although increased variance in the density ‘measure (eg. mixing of crude and ecological densities) will lead to increased variance around the regression line. Currie gives the example that i the densities of small-bodied species tend to be seasonal minima, but those of large-bodied species seasonal maxima, then the regression slope would tend to be over-estimated (Ge. less negative than in reality). A more insidious, and realistic, possibility is that small-and large-bodied me Animal abundance-body size relationships (© 1997 British Ecological Society Journal of Animal cology, 66, 233-289 Contato dterton {watered g Boay mas range ores of mate) ‘re i Fe ‘Sua range rae of magnitude) re Tos Jit i 4 bE | sierege odor magne) Mig.6. 3) The reaionship between the /-eansformad cetcent of determination reported by a study and the range of body masses (orders of maids) spanned hy the spec or which data were wed (6) Te same reltonship above, bat with he boy mas range divided into dere clases. () The relationship between the OLS regresion Slope reported by sly and the range of body mies spanned bythe seis, ith the boy mas ange (ners of tagnitud) divided ato dscrcte clases In (0) ad (cars indieate standard eros, and numbers by each point ste sample sis, A study wa inclaed in ize range cass O-1if itesbe range was 30 and <1, and soon species differ in respect to where along the erude~ ecological density continuum they tend to lie, even within definitions of ‘erude’ and ecological’. To what extent the precise use of erude and ecological densities is random with respect to body size is a key issue, and one that has not generally been addressed in the literature, Nevertheless the lack of effect of density measure on regression slope suggests that there is not a strong size bias in the use of densities at the erude and ecological ends of the density spectrum ‘The lack of effect of density measure on the coefficient of determination could also be argued t0 indicate that there is no mixing of density measures in different studies (plots stated to use ecological den- sities really do 50, and likewise those stated to use crude densities). An alternative interpretation is that the measures are always mixed to some degree, so that 4 statement about the density measure used in any particular interspecific study is largely meaningless. ‘We favour the latter interpretation. Since densities for any single species fall on a continuum, we find it ‘unlikely inthe extreme that the densities in any given study will representa similar quantity for all species, as the above example from Green (1979) illustrates ‘The second point Currie (1993) makes is that if crude and ecological density show an approximately constant difference, then the slopes of the respective relationships plotted using them should be parallel, although of differing elevation, All else being equal, ‘rude and ecological densities should therefore reveal the same patterns, He presents abundance-body size relationships from Damuth (1981), calculated using vological densities, and Peters & Wassenberg (1983), using crude densities for the same taxa, to show that the slopes are infact very similar using both measures. His conclusion is that there is no evidence that density measure systematically biases the slope, However, hhow crude Peters & Wassenberg’s data are is open to debate. They seem t0 us to lie more towards the ecological end of the density spectrum. If, as is likely, both studies eombine densities from a variety of points along the continuum, then itis hardly surprising that they show no significant differences. Perhaps more than anything, they serve to highlight how difficult it is to make such distinctions. ‘The likely reason that different density measures have boen considered to affect the abundancs-body size relationship obtained is that there isan interaction between density measures and other factors. In par= ticular, density estimates from single-species studies are compiled into estimates of regional relationships, and are most often claimed to be ecological, whereas relationships at local scales use density estimates from ‘sample studies that are usualy defined as crude. Study scale shows consistent effects on the regression slope ‘and coefficient of determination of abundance-body size relationships, with more steeply negative slopes and higher coefficients at regional scales (Tables 8 and 9) Two of the factors suggested to affect the form of the abundance-body size relationship ~ the body size range and dimensionality of an assemblage ~ explain variation in the strength of the relationship, but do ‘not help with understanding the variation ints slope. As prodicted by Currie (1993), the coefficient of deter= ‘mination increases with the range of body sizes over ‘which relationships are plotted. However, this pattern 243 TM. Blackburn & Kl. Gaston © 1997 British Ecological Society Journal of nin cology, 66, 233-249 ‘Table 6. OLS slopeand|(y transform of) values from estimates of theabundance-body sre relationship fora) assemblages considered to use the environment in two or three dimensions (Dimensionalty, and (b) bid and mammal assemblagse (axon), together with the esl (F, P) of anowvas esting fr diferences in the di ivstions ofthe statistics Estimate Mean (SB) (@) Dimensionality Slope ‘Two-dimensional 0286 + 0078 ‘Thrce-dimensional 0380 + 0050 vl “Two-dimensional 407 +0032 ‘Three-dimensional 0299.4 0024 (©) Taxon Slope Birds 0370 + 0056 Mammals 0264 + 0.082 ul Birds 0299 + 0024 Mamas 0-403 +0033 ® F > 56 049 ow 9 56 331 0.0076 a si be 07 3 4 663 oor 2 is not simple; rather, the inerease is only exhibited ‘over large body size ranges. Further, the regression slope changes quite markedly with body size range (Fig 6). These patterns suggest that body size range alone cannot cause the difference between polygonal and linear relationships. It is across small body size ranges in particular that the predictions of this mech- anism are most clearly violated, implying that poly- ‘zonal relationships (which generally have small size ranges) are not simply segments of linear patterns ‘his is another mechanism that seems unlikely to act, in isolation, but could exacerbate the differences caused by other mechanisms; relationships plotted across small range of body sizes tend to use sample data, and to be local in scale. ‘Assemblages of species that use the environment in three dimensions have regression slopes that do not differ from those of assemblages that use the environ- ment in two dimensions, but show more variation around those regression slopes. In one sense, dimen- sionality is simply another way in which error can be introduced into density estimates (Blackburn & Gaston 1997), and the algebraic points raised by Currie (1993) for the effect of density measure (see above) ‘equally apply here. However, the multivariateanalysis ‘suggests that the effect of dimensionality may be quite general, and serves to emphasize the dificulty of com- paring the densities of species that use the environ- ment in different ways. The results presented here imply that, for a given body size, species that use the ‘environment in three dimensions can exhibit a wider range of densities than those that use it in two dimen~ sions, at all scales of study. Nevertheless, it seems less likely that this isa true biological effect than that its the result of problems in assigning an accurate denominator for calculating density in species that perceive a three-

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