Predators are an integral component of any healthy ecosystem. Their influence can even be felt at the botanical level via what are called top-down controls. Either through direct predation or through altering their behavior, predators influence the herbivores in any system, which usually results in healthier plant communities. This method is rather indirect but new evidence shows that in the Arctic tundra, a top predator is having quite a direct influence on plant communities.

What's not to love about Arctic foxes? All anthropomorphic views aside, Arctic foxes are important predators in this ecosystem. Although the food web complexity on the tundra is largely driven by limits to plant productivity, a paper published in 2016 shows that these little canids can have profound effects on vegetation. This doesn't have to do with predation directly but rather their reproductive behavior.

Arctic foxes live, give birth, and raise their young in underground dens. Without these subterranean homes, the foxes would be much more vulnerable to other predators as well as the harsh Arctic climate. Dens don't happen overnight either. Suitable sites are tended for generations and some dens may well be more than a century old. All this equates to a lot of activity in and around a good den site.

With an average litter size of 8 - 10 pups per female, one can imagine the food and waste buildup must be considerable. Like all predators, Arctic fox food and waste are rich in nitrogen and phosphorus compounds, the necessary building blocks of life. Many an onlooker has noticed that, unsurprisingly, plant growth around Arctic fox dens is much more lush than on the surrounding landscape. Until recently though, such differences have hardly been quantified.

Arctic Fox (Vulpes lagopus) photo by Allan Hopkins licensed under CC BY-NC-ND 2.0

By examining the soil and plant characteristics around Artic fox dens in Canada and comparing these data to surrounding sites without Arctic fox dens, a team of researchers put the first comprehensive numbers to the effects of Arctic foxes on tundra plant communities. They found that soils from in and around Arctic fox dens contained significantly higher levels of nitrogen and phosphorus than did the surrounding control plots. What's more, these levels varied throughout the year. In June, for instance, soil nitrogen and phosphorus levels were 71% and 1195% higher than non-den soils. These levels seemed to switch later in the summer. In August, soil nitrogen from fox dens were 242% higher and soil phosphorus levels were 191% higher.

As you can probably imagine, all of these extra nutrients caused a change in vegetation around the dens. Den sites were far more productive in terms of vegetation. The team found that, on average, Arctic fox dens supported 2.8 times more plant biomass than did the surrounding area. The authors note that these were conservative estimates and that the true values are much higher. Taken together, these results demonstrate that far from simply being top predators, Arctic foxes are true ecosystem engineers, at least on local scales. This is especially important in such a demanding ecosystem as the Arctic tundra.

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If healthy ecosystems are what we desire, we must embrace predators. There is no way around it. Because of their meat-based diets, predators can have serious effects on plant diversity. Generally speaking, as plant diversity increases, so does the biodiversity of that region. It's not just large predators like wolves and bears either. Even predators as small as spiders can have considerable impacts on not only plant diversity, but ecosystem processes as well. Before we get to that, however, we should take a moment to review some of the background on this subject.

The way in which predators mediate plant diversity falls under a realm of an ecological science called top-down ecosystem controls. In a top-down system, predators mediate the populations of herbivores, which takes pressure off of the plant community. It makes a lot of sense as a numbers game. The fewer herbivores there are, the better the plants perform overall. However, ecology is never that simple. More and more we are realizing that top-down controls have less to do with fewer herbivores than they do with herbivore behavior.

Herbivores, like any organism on this planet, respond to changes in their environment. When predators are present, herbivores often become more cautious and change up their behavior as a result. Such is the case of grasshoppers living in fields. Grasshoppers are incredibly numerous and can do considerable amounts of damage to plant communities as they feed. Picture swarms of locusts and you kind of get the idea.

Photo by Andrew Cannizzaro licensed under CC BY 2.0

Given the choice, grasshoppers will preferentially feed on some plants more than others. Such was the case when researchers began observing grasshopper behavior in some old fields in Connecticut. The grasshoppers in this study really seemed to prefer grasses to all other plants. That is unless spiders were present. In this particular system lives a spider known as the nursery web spider (Pisaurina mira). The nursery web spider is an effective hunter and the fact does not seem to be lost on the grasshoppers.

In the presence of spiders, grasshoppers change up their feeding behavior quite a bit. Instead of feeding on grasses, they switch over to feeding on goldenrod (Solidago rugosa). Although the researchers are not entirely sure why they make this shift, they came up with three possible explanations. First is that the goldenrod is much more structurally complex than the grass and thus offers more places for the grasshopper to hide. Second is that goldenrod fills the grasshoppers stomach in less time thanks to the higher water content of the leaves. This would mean that grasshoppers had more time to watch for predators than they would if they were eating grass. Third is that the feeding behaviors of both arthropods allows the grasshopper to better keep track of where spiders might be lurking. It is very likely that all three hypotheses play a role in this shift.

Photo by Tibor Nagy licensed under CC BY-NC 2.0

It's the shift in diet itself that has ramifications throughout the entire ecosystem in question. Many goldenrod species are highly competitive when left to their own devices. If left untouched, abandoned fields can quickly become a monoculture of goldenrod. That is where the spiders come in. By causing a behavioral shift in their grasshopper prey, the spiders are having indirect effects on plant diversity in these habitats. Because grasshoppers spend more time feeding on goldenrods in the presence of spiders, they knock back some of the competitive advantages of these plants.

The researchers found that when spiders were present, overall plant diversity increased. This is not because the spiders ate more grasshoppers. Instead, it's because the grasshoppers shifted to a diet of goldenrod, which knocked the goldenrod back just enough to allow other plants to establish. It's not just plant diversity that changed either. Spiders also caused an increase in both solar radiation and nitrogen reaching the soils!

In knocking back the goldenrod, the habitat became slightly more open and patchy as various plant species of different shapes and sizes gradually established. This allowed more light to reach the soil, thus changing the environment for new seeds to germinate. Also, because goldenrod leaves tend to break down more slowly, they can have significant influences on nutrient cycles within the soil. As a more diverse set of plants establish in these field habitats, the type of leaf litter that falls to the ground changes as well. This resulted in an overall increase in the nitrogen supply to the soil, which also influences plant diversity.

In total, the mere presence of spiders was enough to set in motion these top-down ecosystem effects. It's not that spiders eat more grasshoppers, it's that they are changing the behavior of grasshoppers in a way that results in a more diverse plant community overall. This is a radically different narrative than what has been observed with examples such as the reintroduction of wolves to the greater Yellowstone ecosystem yet the conclusions are very much the same. Predators have innumerable ecosystem benefits that we simply can't afford to ignore.

Photo Credits: [1] [2] [3]