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Ecology

FOCUS

consume, and nonmigratory grazers suffer

from the lack of plant abundance. Not only

is grass less abundant during the dry season,

but it is also less nutritious, leading species

that switch between grass and tree diets to

be more likely to obtain their food from

trees during the dry season. Because of their

reluctance to change diets, nonmigratory

grazers experience a substantial decline

in population sizes during droughts. “The

two types of herbivores that actually do

become abundant—mixed feeders and

migratory grazers—do so by… actively

[having] strategies to ensure that they

have a better dry season than they would

otherwise,” Staver said.

Theoretical Models

A breathtaking landscape of the Kruger National Park.

If you have ever seen Disney’s timehonored

family feature The Lion

King, you likely have a good picture

of what an African savanna looks like.

The African savanna ecosystem is home

to a large and diverse community of

megafauna, or large terrestrial mammals,

whose populations have undergone

serious declines as a result of many

complex factors, such as predator-prey

relationships, disease, and drought.

While the impact of these variables on

the population dynamics of savanna

herbivores have been well-studied by

ecologists, the dietary strategies of these

communities have only just recently been

investigated as a determining factor of

population abundances.

Seeking to better understand this

determinant, Carla Staver, Associate

Professor of Ecology and Evolutionary

Biology at Yale University, and Gareth

P. Hempson, a postdoctoral fellow

at the University of Witwatersrand

Johannesburg, examined the effects that

seasonal dietary changes have on the

populations of savanna herbivore species.

After composing two different theoretical

mathematical models and evaluating

data from several African savanna parks,

Staver and Hempson found that species

that switch their diets seasonally, in

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addition to species that migrate to find

better forage, have increased population

sizes and dominate the savannas.

Grazing and Browsing

IMAGE COURTESY OF WIKIMEDIA COMMONS

The savannas of Africa, characterized

by trees and grasslands, boast biodiverse

habitats that support herbivores such as

browsers, who feed on tree leaves and

shrubs, grazers, who feed on grass and

other low-lying vegetation, and mixed

feeders, who alternate between browsing

and grazing depending on the season.

Browsers, such as giraffes, and grazers, such

as zebras and wildebeests, are considered

specialists because of their efficiency at

eating a particular, albeit limited, diet.

On the other hand, mixed feeders, such

as impala and elephants, are defined as

generalists because of their ability to adapt

and survive off a varied diet.

Savannas are highly seasonal systems,

meaning they alternate between a wet season

with heavy rainfalls and a dry season with

little to no rainfall. During the wet season,

grazers enjoy bountiful plant growth. But

in the dry season, when grass is sparse,

herbivores of the savanna tend to undertake

three different practices: mixed feeders alter

their food source and switch to browsing,

migratory grazers locate new foliage to

Reinforcing their theory that migratory

grazers and mixed feeders maintain

higher population abundances compared

to nonmigratory specialists, Staver and

Hempson presented two mathematical

models in a paper published in Science

Advances that illustrate the role that

dietary strategies play in population

dynamics. The first model, a discretetime

population model, takes into account

the variance of population sizes between

wet and dry season and finds the overall

rate that the populations of herbivores

increase by. Specifically, the model uses

the geometric mean—a form of averaging

found by multiplying rather than adding

numbers—of wet and dry season growth

rates to establish the comprehensive

growth rate of a population.

PHOTOGRAPH COURTESY OF PROFESSOR STAVER

This photograph captures a group of impala, a

classic example of a mixed feeder herbivore.

December 2020 Yale Scientific Magazine 23

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