# How Far Do Animals Go? Determinants of Day Range in Mammals

> How far does an animal travel in a day, and why? Comparing 200 mammal species, the authors show that bigger animals travel farther, but not in proportion to their size, and that meat and insect eaters travel almost three times as far as plant-eaters of the same size. A simple model based on food needs and how food is spread out explains much of the pattern.

Summary by Kersey Lawrence, Original Wisdom. Updated 9 November 2026. Page: https://www.originalwisdom.com/tracking-literature/2005-carbone-cowlishaw-how-far-do-animals/

## Citation and access

- **Authors:** Chris Carbone, Guy Cowlishaw, Nick J. B. Isaac and J. Marcus Rowcliffe
- **Year published:** 2005
- **Journal or source:** The American Naturalist
- **Full citation:** Carbone, C., Cowlishaw, G., Isaac, N. J. B. and Rowcliffe, J. M. 2005. How far do animals go? Determinants of day range in mammals. The American Naturalist 165(2): 290–297.
- **DOI:** 10.1086/426790
- **Access status:** Publisher link only
- **Where to find it:** https://doi.org/10.1086/426790

## Tags

- **Species:** Mammals
- **Behaviour and ecology:** Behaviour; Daily travel distance; Feeding; Movement
- **Methods and technology:** Analysis of covariance (ANCOVA); Body mass–day range scaling; Linear regression
- **Data type:** Existing data; Modelling or theoretical
- **Observer qualification:** Not needed
- **Discipline:** Behaviour and ecology
- **Year published:** 2005

## Detailed keywords

- **Species:** Studied: 200 species of mammals, from published day ranges (the species list is in an online appendix, not in the article). The main groups analysed were the orders Artiodactyla (even-toed hoofed mammals), Carnivora, Primates and Rodentia. Named in the text: elephant (Elephas) and tree hyrax (Dendrohyrax), whose comparison strongly influenced the phylogenetic body mass result. Examples used for the model: ungulates, most mammalian carnivores, insectivorous bats, most primates, African bovids, North American herbivores and grazers in Tanzania.
- **Location and habitat:** Worldwide (data compiled from the literature; no study site). Foraging habitat treated as two-dimensional (ground-living, for example ungulates and most carnivores) or three-dimensional (for example tree-living primates and insect-eating bats); substrate use scored as terrestrial, intermediate or arboreal.
- **Topics:** day range (daily distance travelled); allometry (scaling with body mass); energy and food requirements; food resource distribution and density; diet (herbivores, frugivores, faunivores); prey group size; consumer group size; foraging habitat dimensions; home range and population density as related measures of space use; comparison with Garland’s (1983) day range model; phylogenetic history.
- **Statistics or models:** An energetic and geometric model predicting how day range scales with body mass for predators and herbivores (food requirements scaling at 0.75 of body mass, prey density at minus 0.75 of prey mass, prey group size at 0 to 0.66, plant biomass at 0.21). Least squares linear regression and a structural relations model of log day range on log body mass; analysis of covariance (ANCOVA) with order, diet, substrate use, group size and body mass, reduced to a minimum adequate model; phylogenetically independent contrasts (CAIC 2.6.9 software) with regression through the origin; Pagel’s lambda; Shapiro-Wilks test; one-sample sign test; t-deleted residuals. Diet first sorted into eight categories, then collapsed to three. Possible inconsistencies: (1) Table 1 gives “overall r² = 59.8”, which reads as a percentage rather than a proportion. (2) The Methods say 200 species, but the species counts shown in Figure 2 add up to 194 (Figure 2 may show only the four main orders).
- **Data type:** Existing data: day ranges and other traits for 200 mammal species from primary and secondary literature, with extra body mass data from the PanTHERIA project (unpublished at the time). Modelling or theoretical: a new day range model. No dates of data collection given.

## Synopsis

### 1. What they did

Day range is how far an animal travels in a day. It is a short-term measure of how much space an animal needs, and it partly reflects how much food the animal needs and how that food is spread out. Compared with home range and population density, it had been little studied.

The authors built a simple model to predict how day range should change with body size. The model starts from how much food an animal needs each day, how big each food item is, and how densely food is spread through the habitat. For predators, a food item is one prey animal (and only one is taken from each prey group). For plant-eaters, a food item is one bite. The model also allows for whether an animal forages on a flat surface (the ground) or in three dimensions (trees, the air).

They then collected published day ranges for 200 mammal species, grouped by diet (plant-eaters, fruit-eaters, and meat or insect-eaters, called faunivores), and compared the real patterns with the model’s predictions. They used ordinary statistics and also methods that allow for how closely related the species are, because related species tend to have similar day ranges.

### 2. What they found

What the model predicted

- Day range should rise with body mass, but more slowly than body mass itself.

- Predators on the ground should show the steepest rise (a scaling exponent of 0.25 to 0.58, depending on whether prey live in groups). Plant-eaters on the ground should show less (0.27), and animals foraging in three dimensions less again (0.17 to 0.39 for predators such as insect-eating bats, 0.18 for plant-eaters such as most primates).

- Because there is far less animal food than plant food in a habitat (about 1 to 10), predators should travel about 3.16 times farther than plant-eaters of the same size.

- If grouping brings no feeding benefit, a group’s day range should grow with group size.

What the data showed

- Across all 200 species, day range rose with body mass at a scaling exponent of about 0.29 to 0.31 (0.23 to 0.35 when relatedness was taken into account), close to the roughly one-quarter power found in earlier work and within the model’s predictions.

- Diet mattered. For a given body size and group size, faunivores travelled farthest, fruit-eaters were in between, and plant-eaters travelled least. Faunivores travelled 2.66 times farther than plant-eaters, compared with the model’s 3.16.

- The orders differed. Carnivores showed the steepest rise in day range with body size (0.44), within the model’s range for ground-living predators. Primates showed almost no rise (0.03; the model predicted 0.18 for tree-living plant-eaters), and even-toed hoofed mammals were in between (0.13; the model predicted 0.27).

- Diet and body mass did not interact as the model predicted: diet changed how far animals travelled but not how steeply day range rose with size.

- Allowing for relatedness, longer day ranges went with larger body size, a meat or insect diet and, in simple comparisons, larger groups and living on the ground rather than in trees. In the fuller analysis, body mass and diet stayed strong, but the effect of group size faded and tree or ground living did not matter.

- Within primates, diet was the only clear predictor: in 20 of 24 clear diet changes, the group with the more animal-based diet travelled farther. Within carnivores, both body size and group size predicted day range. Hoofed mammals showed no significant scaling with body size.

### 3. What they concluded

- The model’s general predictions matched the real patterns in day range, body size and diet remarkably well. The predicted slopes, however, tended to be steeper than those observed.

- Possible reasons for the gap: variety within each order; animals finding ways to shorten their travel (for example, social predators sometimes take more than one animal from a prey group); and wide-ranging species being harder to monitor, which could bias the data.

- The low scaling in primates may fit the idea that foraging in three dimensions reduces day range, though shifts in diet and links between group size and body size may also play a part.

- There is only limited evidence that living in groups lengthens day range, except in carnivores. Grouping probably brings feeding advantages, so this is not surprising.

- The model, built on a few basic scaling relationships, offers a useful framework for understanding day range and could be adapted to study other factors in how animals range.

### 8. Implications for tracking (Original Wisdom's observations)

_These are Original Wisdom's observations, not the authors'._

- Day range is a key number when track counts are turned into animal density: the Formozov–Malyshev–Pereleshin (FMP) formula divides the number of track crossings by the day range. This paper’s data set has been used for that purpose, for example by Ahlswede et al. (2019) for spoor counts in Namibia, where day ranges for hoofed mammals were estimated from body mass.

- The paper shows how much day range varies with diet and between groups of mammals. Estimating one species’ day range from body mass alone, or from a relationship built on a different group, may therefore be uncertain, and day ranges measured in the field (for example, by following an animal’s trail for a full day) may give better results.

## Cite this summary

Lawrence, K. 2026. Summary of: How Far Do Animals Go? Determinants of Day Range in Mammals (Carbone and Cowlishaw et al., 2005). Original Wisdom, Tracking Literature. https://www.originalwisdom.com/tracking-literature/2005-carbone-cowlishaw-how-far-do-animals/