Home range, home range overlap, and species energy use among herbivorous mammals

Harestad & Bunnell (1979) showed that, at least for North American species, home ranges of large herbivorous mammals are relatively larger than we would expect on metabolic grounds, and suggested that the productivity of the environment for mammal species decreases with increasing body size. This interpretation assumes that the number of conspecifics that share an individual's home range is independent of body size. Data presented here show that this is not true for the species in their sample; the home range is shared with an increasing number of conspecifics in larger herbivore species. The productivity of the environment for a species is independent of body size and the area available to an individual for its own use scales approximately as do individual metabolic requirements. These results agree with conclusions based upon the scaling of population density with body mass and illustrate the interrelationship between home range and dietary and social organization trends among mammalian herbivores. Individual home range area is a function of the way in which the local population of a species, not merely an individual, exploits the environment.     

The ontogeny of home ranges: evidence from coral reef fishes

The concept of home ranges is fundamental to ecology. Numerous studies have quantified how home ranges scale with body size across taxa. However, these relationships are not always applicable intraspecifically. Here, we describe how the home range of an important group of reef fish, the parrotfishes, scales with body mass. With masses spanning five orders of magnitude, from the early postsettlement stage through to adulthood, we find no evidence of a response to predation risk, dietary shifts or sex change on home range expansion rates. Instead, we document a distinct ontogenetic shift in home range expansion with sexual maturity. Juvenile parrotfishes displayed rapid home range growth until reaching approximately 100–150 mm length. Thereafter, the relationship between home range and mass broke down. This shift reflected changes in colour patterns, social status and reproductive behaviour associated with the transition to adult stages. While there is a clear relationship between body mass and home ranges among adult individuals of different species, it does not appear to be applicable to size changes within species. Ontogenetic changes in parrotfishes do not follow expected mass–area scaling relationships.     

Discontinuities and alternative scalings in the density–mass relationship of anuran larvae

Size–density relationships (SDRs) frequently follow a power-law relationship, with exponents that compensate for the increase in population-level metabolic demand—the energetic equivalence rule. However, these exponents present a range of values, and elucidating its methodological and biological determinants has become a main issue. So far, a restricted set of potential relationships, mechanisms, and taxa have been considered. Here, we analyzed SDR in a population of tadpoles inhabiting a network of 16 temporal ponds. Alternative scaling regimes were detected using pacewise regressions and estimating exponents with maximum likelihood (ML). If discontinuities in the SDR are ignored, a scaling close to values reported elsewhere is observed. However, estimated slopes between discontinuities are steeper (threefold to fivefold) than those often reported, but congruent with the performance predicted for ML and the biases reported for other methods. Our estimations largely deviate from an energetic equivalence, suggesting that large individuals use less energy per unit area. The detection of different SDRs in the same database, with a strong decay in abundance with body size, points to a pattern poorly considered in previous studies, widening the range of patterns, mechanisms, and ecological, or evolutionary consequences of the SDRs.     

Allometric scaling of mortality rates with body mass in abalones

The existence of an allometric relationship between mortality rates and body mass has been theorized and extensively documented across taxa. Within species, however, the allometry between mortality rates and body mass has received substantially less attention and the consistency of such scaling patterns at the intra-specific level is controversial. We reviewed 73 experimental studies to examine the relationship between mortality rates and body size among seven species of abalone (Haliotis spp.), a marine herbivorous mollusk. Both in the field and in the laboratory, log-transformed mortality rates were negatively correlated with log-transformed individual body mass for all species considered, with allometric exponents remarkably similar among species. This regular pattern confirms previous findings that juvenile abalones suffer higher mortality rates than adult individuals. Field mortality rates were higher overall than those measured in the laboratory, and the relationship between mortality and body mass tended to be steeper in field than in laboratory conditions for all species considered. These results suggest that in the natural environment, additional mortality factors, especially linked to predation, could significantly contribute to mortality, particularly at small body sizes. On the other hand, the consistent allometry of mortality rates versus body mass in laboratory conditions suggests that other sources of mortality, beside predation, are size-dependent in abalone.     

Ontogenetic body-mass scaling of resting metabolic rate covaries with species-specific metabolic level and body size in spiders and snakes

According to common belief, metabolic rate usually scales with body mass to the 3/4-power, which is considered by some to be a universal law of nature. However, substantial variation in the metabolic scaling exponent (b) exists, much of which can be related to the overall metabolic level (L) of various taxonomic groups of organisms, as predicted by the recently proposed metabolic-level boundaries (MLB) hypothesis. Here the MLB hypothesis was tested using data for intraspecific (ontogenetic) body-mass scaling of resting metabolic rate in spiders and boid snakes. As predicted, in both animal groups b varies mostly between 2/3 and 1, and is significantly negatively related to L. L is, in turn, negatively related to species-specific body mass (M_m: estimated as the mass at the midpoint of a scaling relationship), and as a result, larger species tend to have steeper metabolic scaling slopes (b) than smaller species. After adjusting for the effects of M_m, b and L are still negatively related, though significantly only in the spiders, which exhibit a much wider range of L than the snakes. Therefore, in spiders and snakes the intraspecific scaling of metabolic rate with body mass itself scales with interspecific variation in both metabolic level and body mass.     

The effect of season, sex and feeding style on home range area versus body mass scaling in temperate ruminants

Sex-specific estimates of the summer and winter home range area of 19 species of temperate ruminants were collated from the literature. It was predicted that there should be a shallower slope for the home range area against body mass relationship during winter than during summer, as large ruminants can meet more of their energy requirements from the fat reserves deposited during summer than small ruminants. Consequently, relatively large species do not need to range as widely during winter. There was a significant positive relationship between body mass and summer and winter home range area in both females and males. This relationship remained significant when analysed within feeding styles (browser, mixed feeder, grazer), except in mixed feeders in winter. As predicted, slope estimates were significantly lower during winter (b=0.59) than during summer (b=1.28), both before and after controlling for phylogeny. After controlling for phylogeny, browsers had a steeper slope (summer: b=1.48; winter: b=1.07) of the home range area against body mass relationship than did mixed feeders (summer: b=0.75; winter: b=-0.11) or grazers (summer: b=1.10; winter: b=0.34). There was no effect of sex after body mass was controlled for. The effect of season, sex and feeding style on the home range area versus body mass relationship in temperate ruminants is discussed.     

The limits to population density in birds and mammals

We address two fundamental ecological questions: what are the limits to animal population density and what determines those limits? We develop simple alternative models to predict population limits in relation to body mass. A model assuming that within‐species area use increases with the square of daily travel distance broadly predicts the scaling of empirical extremes of minimum density across birds and mammals. Consistent with model predictions, the estimated density range for a given mass, ‘population scope’, is greater for birds than for mammals. However, unlike mammals and carnivorous birds, expected broad relationships between body mass and density extremes are not supported by data on herbivorous and omnivorous birds. Our results suggest that simple constraints on mobility and energy use/supply are major determinants of the scaling of density limits, but further understanding of interactions between dietary constraints and density limits are needed to predict future wildlife population responses to anthropogenic threats.     

Allometric scaling of intraspecific space use

Allometric scaling relationships enable exploration of animal space-use patterns, yet interspecific studies cannot address many of the underlying mechanisms. We present the first intraspecific study of home range (HR) allometry relative to energetic requirements over several orders of magnitude of body mass, using as a model the predatory fish, pike Esox lucius. Analogous with interspecific studies, we show that space use increases more rapidly with mass (exponent = 1.08) than metabolic scaling theories predict. Our results support a theory that suggests increasing HR overlap with body mass explains many of these differences in allometric scaling of HR size. We conclude that, on a population scale, HR size and energetic requirement scale allometrically, but with different exponents.     

Test of the Optimal Body Size Model for Strepsirhines

We determined if data on strepsirhine body and home range sizes support an optimal body size (OBS) model of 100 g, as predicted from studies of energetics in terrestrial mammals. We also tested the following predictions of the OBS model: 1) relationships between body and home range sizes will change slope and sign above and below the OBS threshold of 100 g and 2) best-fit lines for OBS regression models (above and below the 100-g threshold) will intersect at ca. 100 g (range of 80–250 g). We collected data on body mass, home range size, and vertical ranging behavior for 37 strepsirhines from the literature. Linear regression analyses and phylogenetic independent contrasts methods revealed that body size is a significant determinant of both 2-dimensional (ha) and 3-dimensional (km³) home range sizes only in taxa weighing >100 g. There were consistent changes in the sign of the slopes above and below the OBS threshold. The intersections of the best-fit lines were within the OBS range for the body size to 3-dimensional home range comparisons. Thus, the data provide some support for the OBS model in strepsirhines. However, no regression model was statistically significant for the taxa below the OBS threshold, which may reflect small sample sizes. Also, no slope differed significantly between taxa above and below the OBS. Significant correlations between body and home range sizes for the complete data sets refute the √-shaped constraint space predicted via the OBS model.     

Of mice and wrens: the relation between abundance and geographic range size in British mammals and birds

We examine the relation between population size and geographic range size for British breeding birds and mammals. As for most other assemblages studied, a strong positive interspecific correlation is found in both taxa. The relation is also recovered once the phylogenetic relatedness of species has been controlled for using an evolutionary comparative method. The slope of the relation is steeper for birds than for mammals, but this is due in large part to two species of mammals that have much higher population sizes than expected from their small geographic ranges. These outlying mammal species are the only ones in Britain to be found only on small offshore islands, and so may be exhibiting density compensation effects. With them excluded, the slope of the abundance–range size relation for mammals is not significantly different to that for birds. However, the elevation of the relation is higher for mammals than for birds, indicating that mammals are approximately 30 times more abundant than birds of equivalent geographic range size. An earlier study of these assemblages showed that, for a given body mass, bats had abundances more similar to birds than to non-volant mammals, suggesting that the difference in abundance between mammals and birds might be due to constraints of flight. Our analyses show that the abundance–range size relation for bats is not different for that from other mammals, and that the anomalously low abundance of bats for their body mass may result because they have smaller than expected geographic extents for their size. Other reasons why birds and mammals might have different elevations for the relation between population size and geographic range size are discussed, together with possible reasons for why the slopes of these relations might be similar.     

Intra-specific variation and taxa- sampling affects the home range — body mass relationship

The relationship between home range size and body mass is a frequently studied allometric relationship. However, the results of various studies differ greatly, leading to much debate about the nature of the relationship. We argue that this confusion is not surprising, due to intra-specific variation in home range size caused by ecological variability rather than by body mass. By random resampling of different studies from within 16 Carnivora species, we show that the scaling exponent ranged from 0.30–1.54 depending on the particular studies included for each species. Of these exponents, 10% did not contain 0.75 within their confidence limits, and 5.5% did not contain 1.00. Furthermore, by randomly sub-sampling 16 species from a total sample of 58 species, we found that the scaling exponent varied between 0.18 and 2.76. Of these exponents, 42.2% did not contain 0.75 within their confidence limits, whereas 16.8% did not contain 1.00. Therefore, we strongly recommend that greater consideration be paid to intra-specific ecological variability and taxa selection when dealing with both allometry and cross-species life history studies.     

A telemetry-based study of bushbuck (Tragelaphus scriptus) home range in Valley Bushveld

With the aid of radio‐telemetry, bushbuck home range was investigated to determine total home range size, home range utilization and home range overlap for the summer season. Estimates of total home range size for males using minimum convex polygons (MCPs) and fixed kernels (FKs) were 33.9 and 32.1 ha, respectively. Estimates of total home range size for females using MCPs and FKs were 12.0 and 13.5 ha, respectively. A significant difference between total home range sizes for male and female was found but there was no significant difference for age (adult and subadult). Female bushbuck home range size was compared to that expected from the published allometric relationship for the scaling of home range area on body mass, where the study animals appeared to have home ranges of half of that predicted. Bushbuck typically utilized one core area within their home ranges in which 50% of their time was spent in approximately 17.0% and 11.7% of their total home range for males and females, respectively. A substantial overlap in total home range and core areas between animals was found.     

Home range dynamics of the yellow‐footed rock‐wallaby (Petrogale xanthopus celeris) in central‐western Queensland

Analyses of the interspecific differences in macropod home range size suggest that habitat productivity exerts a greater influence on range size than does body mass. This relationship is also apparent within the rock‐wallaby genus. Lim reported that yellow‐footed rock‐wallabies (Petrogale xanthopus xanthopus) inhabiting the semi‐arid Flinders Ranges (South Australia) had a mean home range of 170 ha. While consistent with the hypothesis that species inhabiting less productive habitats will require larger ranges to fulfil their energetic requirements, the ranges reported by Lim were considerably larger than those observed for heavier sympatric macropods. The aim of the current study was to document the home range dynamics of P. x. celeris in central‐western Queensland and undertake a comparison with those reported for their southern counterparts. Wallaby movements were monitored at Idalia National Park, between winter 1992 and winter 1994. Male foraging ranges (95% fixed kernel; 15.4 ha, SD = ±7.8 ha) were found to be significantly larger than those of female wallabies (11.3 ha, SD = ±4.9 ha). Because of varying distances to the wallabies' favoured foraging ground (i.e. an adjacent herb field), the direction in which the wallabies moved to forage also significantly affected range size. Mean home range size was estimated to be 23.5 ha (SD = ±15.2 ha; 95% fixed kernel) and 67.5 ha (SD = ±22.4 ha; 100% minimum convex polygon). The discrepancy between these two estimates resulted from the exclusion of locations, from the 95% kernel estimates, when the wallabies moved to a water source 1.5 km distant from the colony site. The observed foraging and home ranges approximated those that could be expected for a macropod inhabiting the semi‐arid zone (i.e. 2.4 times larger‐than‐predicted from body mass alone). Possible reasons for the disparity between the current study and that of Lim are examined.     

Towards a coherent allometric framework for individual home ranges,key population patches and geographic ranges

Spatial requirements of species have often been related to body size, usually focusing on one area variable and one taxonomic group at a time. Here, we carried out a quantitative meta‐analysis and developed a minimal model, covering different types of spatial characteristics and several species groups. In a global literature review, 46 empirical regressions on home ranges and geographic ranges were collected, covering thousands of species monitored in various countries. In addition, regional data on minimum key population patches of 167 species occurring in the Netherlands were retrieved from reports. To check consistency, a theoretical model was derived from rate and density variables based on energy equivalency. The minimum number of individuals needed to sustain a viable population was considered invariant to size. According to the equations, areas were expected to scale to mass with an exponent of 1 for the individual home range and of about ¾ for the minimum population ranges. The meta‐analysis of the empirical regressions showed that average slopes for individual home ranges were between 0.74 for cold‐blooded species and 1.05 for birds and mammals. Minimum and average species geographic range scaled to mass with exponents of 1.16–1.29 and 0.28–0.46 respectively. Allometric correlations for the minimum key patch area were weak. The intercepts indicated that carnivores require more space than equally sized herbivores, while homeotherms occupy larger areas than heterotherms. Observed slopes and intercepts were often near model estimations, but important deviations from the average level were noted as well, especially for birds. Although variability was substantial in some cases, allometric approaches can contribute considerably to understanding and protecting area requirements of species.     

The maximal body mass–area relationship in island mammals

Aim A positive power relationship between maximal body mass and land area has previously been reported of the form M_max ∝ Area^0.5 whilst allometric scaling theory predicts either M_max ∝ Area^1.33 or M_max ∝ Area¹. We provide an analysis of the maximal mass–area relationship for four island systems, to test the hypothesis that community relaxation following isolation converges in each case to a slope of Area^0.5. Location Islands of the Japanese archipelago, the western Mediterranean, the Sea of Cortés and Southeast Asia. Methods We calculated the relationship between island area and the maximal body mass of the largest mammal species on the island using linear regression models with log‐transformed variables, and tested the hypothesis that the slopes were not significantly different from 0.5. Results We found a slope of 0.47 within the Japanese archipelago, 0.42 for western Mediterranean islands, 0.73 for the Sea of Cortés islands and 0.50 for Southeast Asian islands. None of these slopes were significantly different from 0.5. Main conclusions Our results provide further empirical support for previous findings of a general maximal body mass–area relationship of M_max ∝ Area^0.5, but they deviate from theoretical predictions. We hypothesize that this mass–area relationship was the ultimate end point of community relaxation initiated by the isolation of the mammal communities. Maximal body mass on each island today probably reflects the interaction between energetic constraints, home range size and island area.     

Fish mercury bioaccumulation as a function of feeding behavior and hydrological cycles of the Rio Negro, Amazon

The acidic black waters of the Rio Negro are unimpacted by anthropogenic sources and yet, are characterized by fish containing relatively high Hg concentrations. Regular annual flooding alters the aquatic environment, thereby affecting fish feeding strategies. We studied the impact of annual flooding on fish-Hg bioaccumulation. Tucunarés (Cychla spp, carnivorous), Peixe-cachorro (Hydrolycus scomberoides, carnivorous), Traíra (Hoplias malabaricus, carnivorous), Piranha-branca (Serrasalmus aff. eigenamanni, carnivorous), Piranha-preta (Serrasalmus rhombeus, carnivorous), Acará (Acarichthys heckellii, omnivorous), Aracú (Leporinus friderici, omnivorous), Orana-preta (Hemiodus unimaculatus, omnivorous), Sardinha (Triportheus elongatus, omnivorous), Branquinha (Potamorhina latior, detritivorous), Jaraqui-escama-fina (Semaprochilodus taeniurus, detritivorous), and Pacú-branco (Myleus torquatus, herbivorous) were studied during high (July) and low waters (February) and categorized by body mass. Regardless of hydrological period, ranges of total-Hg concentrations were higher for carnivorous than for omnivorous, detritivorous and herbivorous species. Some species (Traíra, Sardinha, Peixe-cachorro, Piranha branca, Piranha preta) showed changes in feeding patterns as indicated by an inverse trend of Hg bioaccumulation with season. Species with similar trends of Hg bioaccumulation showed higher Hg concentrations during the flooded season (Aracú, Pacú branco and Orana preta) and some had lower Hg (Acará, Branquinha and Tucunarés). Fish Hg concentration is species specific and reflects changes in feeding-behavior dominance brought by annual inundation.     

Scaling body mass and use of space in three species of marsupials in the Atlantic Forest of Brazil

Abstract Body mass is considered a major determinant of home range size, but usually at a large scale of body mass variation. The exact scale where body size becomes more important than particular adaptations of each species is not clear, and uncertainty in the estimate of home range size is a possible cause of weak intraspecific scaling. We determine the scaling to body mass of two alternative movement measurements, daily home range (DHR) and its intensity of use (IU), in three species of didelphid marsupials, Didelphis aurita, Philander frenatus, and Metachirus nudicaudatus (Didelphimorphia, Didelphidae). The expected scaling exponents DHR ≈ M^0.5 and IU ≈ M^?0.25 were derived from the scaling to body mass of home range and daily movement distance. Animals were tracked in Serra dos Órgãos, Rio de Janeiro, Brazil, using a spool‐and‐line device. Individuals of the three species were compared combining intra and interspecific variation in a single analysis, with species, body mass, and thread tracked as covariates. The model best supported included only body mass as the independent variable, with DHR ≈ M^0.435 and IU ≈ M^?0.218, close to the expected values. The second best supported model included species identity, but with a non‐significant effect. It was surprising that body mass was more important than species identity in a comparison involving only three species, and considering the morphological and locomotory adaptations of the three species. Body mass may become more important than species identity when the scale of variation approaches one order of magnitude.     

Spatial organization of the arboreal carnivorous marsupial Phascogale tapoatafa

Phascogale tapoatafa is a squirrel-sized carnivorous marsupial (Dasyuridae) found in dry eucalyptus forests of Australia. Radio-tracking and live-trapping studies at three sites in Victoria indicated that population densities were typically low, with females occupying home ranges averaging 41 ha (minimum convex polygon method) that were intrasexually exclusive. Male home ranges (mean 106 ha) overlapped extensively with females and other males, and expanded during the short breeding season to an average length of 2.7 km. When presented with the faeces of unfamiliar conspecifics, both female and male P. tapoatafa investigated the faeces of females significantly longer than those of males, suggesting that the exclusive home ranges of females are reinforced by olfactory communication, and that males are less responsive to the presence of potential rivals than to potential mates. Both sexes foraged throughout their home range, but the core area of intensive use was less than one-third of the total area traversed. The spatial organization of P. tapoatafa differs markedly from that of the closely related species Antechinus stuartii , but is similar to that of the only other extensively studied carnivorous marsupial, Dasyurus geoffroii.     

Sampling effort, regression method, and the shape and slope of size–abundance relations

1. Despite a substantial body of work there remains much disagreement about the form of the relationship between organism abundance and body size. In an attempt at resolving these disagreements the shape and slope of samples from simulated and real abundance–mass distributions were assessed by ordinary least squares regression (OLS) and the reduced major axis method (RMA).
2. It is suggested that the data gathered by ecologists to assess these relationships are usually truncated in respect of density. Under these conditions RMA gives slope estimates which are consistently closer to the true slopes than OLS regression.
3. The triangular relationships reported by some workers are found over smaller mass and abundance ranges than linear relations. Scatter in slope estimates is much greater and positive slopes more common at small sample sizes and sample ranges. These results support the notion that inadequate and truncated sampling is responsible for much of the disagreement reported in the literature.
4. The results strongly support the notion that density declines with increasing body mass in a broad, linear band with a slope around −1. However there is some evidence to suggest that this overall relation results from a series of component relations with slopes which differ from the overall slope.