Invasion of the Earthworms

As an avid gardener, amateur fisherman, and a descendant of a long line of farmers, I have always held earthworms in high regard. These little ecosystem engineers are great for all of the above, right?

Not so fast! Things in life are never that simple! Let's start at the beginning. If you live in an area of North America where the glaciers once rested, there are no native terrestrial worms in your region. All of North America's native worm populations reside in the southeast and the Pacific northwest. All other worms species were wiped out by the glaciers. This means that, for millennia, northern NoNorth America's native ecosystem has evolved without the influence of any type of worms in the soil.

Shading = Glaciers  [1]

Shading = Glaciers [1]

When Europeans settled the continent, they brought with them earthworms, specifically those known as night crawlers and red wigglers, in the ballasts of their ships. Since then, these worms have been spread all over the continent by a wide range of human activities like farming, composting, and fishing. Since their introduction, many forests have been invaded by these annelids and are now suffering quite heavily from earthworm activities.

As I said above, any areas that experienced glaciation have evolved without the influence of worms. Because of this, forests in these regions have built up a large, nutrient-rich, layer of decomposing organic material commonly referred to as "duff" or "humus." Native trees, shrubs, and forbs rely on this slowly decomposing organic material to grow. It is high in nutrients and holds onto moisture quite well. When earthworms invade an area of a forest, they disrupt this rich, organic layer in quite a serious way.

Worms break through the duff and and distribute it deeper into the soil where tree and forb species can no longer access it. Worms also pull down and speed up the decomposition of leaves and other plant materials that normally build up and slowly create this rich organic soil. Finally, earthworm castings or poop actually speed up runoff and soil erosion.

All of this leads to seriously negative impacts on native ecosystems. As leaves and other organic materials disappear into the soil at an alarming rate via earthworms, important habitat and food is lost for a myriad of forest floor organisms. In areas with high earthworm infestations, there is a significant lack of small invertebrates like copepods. The loss of these organisms has rippling effects throughout the ecosystem as well. It has been shown that, through these activities, earthworms are causing declines in salamander populations.

It gets worse too. As earthworms speed up the breakdown of the duff or humus, our native plant species are suffering. They have evolved to germinate and grow in these rich, organic soils. They rely on these soils for survival. As the nutrient rich layers get redistributed by earthworms, native plant and tree populations are suffering. There is very little recruitment and, in time, many species are lost. Our spring ephemerals have been shown to be hit the hardest by earthworm invasions. Earthworms have also been shown to upset the mycorrhizal fungi networks which most plant species cannot live without.

Top Left: Forest soil horizons without earthworms; Top Right: Forest soil mixed due to earthworms; Bottom Left: Forest understory diversity without earthworms; Bottom Right: Forest understory diversity with earthworms. Credits: [1]

So, what can we do about this? Well, for starters, avoid introducing new populations of earthworms to your neighborhood. If you are using earthworms as bait, do not dump them out onto land when you're done. If you must get rid of them, dump them into the water. Also, if you are using worm castings in your garden, it has been recommended that you freeze them for about a week to assure that no eggs or small worms survive the ride. If you are bringing new plants onto your property, make sure to check their root masses for any worm travelers. Remember, no worms are native if you live in a once glaciated region.

Sadly, there is not much we have come up with at this point for dealing with the current earthworm invasion. What few control methods have been developed are not practical on a large scale and can also be as upsetting to the native ecology as the earthworms. The best bet we have is to minimize the cases of new introductions. Earthworms are slow critters. They do not colonize new areas swiftly. In fact, studies have shown that it takes upwards of 100 years for earthworm populations to migrate 1/2 mile! Armed with new knowledge and a little attention to detail, we can at least slow their rampage.

Photo Credit: Peter Hartl

Further Reading: [1] [2] [3] [4] [5] [6] [7] [8] [9] [10]

How Spiders Increase Plant Diversity

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.

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.

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]

Further Reading: [1] [2]


On Fungi and Forest Diversity

One simply can't talk about plants without eventually talking about fungi. The fact of the matter is the vast majority of plant species rely on fungal interactions for survival. This mutualistic relationship is referred to as mycorrhizal. Fungi in the soil colonize the root system of plants and assist in the acquisition of nutrients such as nitrogen and phosphorus. In return, most photosynthetic plants pay their mycorrhizal symbionts with carbohydrates. 

There are two major categories of mycorrhizal fungi - ectomycorrhizae (EMF) and arbuscular mycorrhizae (AMF). Though there are a variety of different species of fungi that fall into either of these groups, their strategies are pretty much the same. EMF make up roughly 10% of all the known mycorrhizal symbionts. The prefix "ecto" hints at the fact that these fungi form on the outside of root cells. They form a sort of sheath that encases the outside of the root as well as a "hartig net" around the outside of individual cells within the root cortex. AMF, on the other hand, literally penetrate the root cells and form two different kinds of structures once inside. One of these structures looks like the crown of a tree, hence the term "arbuscular." What's more, they are considered the oldest mycorrhizal group to have evolved. 

The type of mycorrhizal fungi a plant partners with has greater implications that simple nutrient uptake. Evidence is now showing that the dominant fungi of a region can actually influence the overall health and diversity forest ecosystems. The mechanism behind this has a lot to do with the two different categories discussed above. 

Researchers have discovered that trees partnering with AMF experience negative feedbacks in biomass whereas those that partner with EMF experience positive feedbacks in biomass. When grown in soils that previously harbored similar tree species, trees that partnered with AMF grew poorly whereas trees that partnered with EMF grew much better. Additionally, by repeating the experiments with seedlings, researchers found that seedling survival was reduced for AMF trees whereas seedling survival increased in EMF trees. 

What is going on here? If mycorrhizae are symbionts, why would there be any detrimental effects? The answer to this may have something to do with soil pathogens. Thinking back to the major differences between EMF and AMF, you will remember that it comes down to the way in which they form their root associations. EMF form a protective sheath around the roots whereas AMF penetrate the cells.  As it turns out, this has major implications for pathogen resistance. Because they form a sheath around the entire root, EMF perform much better at keeping pathogens away from sensitive root tissues. The same can't be said for AMF. Researchers found that AMF trees experienced significantly more root damage when grown in soils that previously contained AMF trees. 

The differences in the type of feedback experienced by EMF and AMF trees can have serious consequences for tree diversity. Because EMF trees are healthier and experience increased seedling establishment in soils containing other EMF species, it stands to reason that this would lead to a dominance of EMF species, thus reducing the variety of species capable of establishing in that area. Conversely, areas dominated by AMF trees may actually be more diverse due to the reduction in fitness that would arise if AMF trees started to dominate. Though they are detrimental, the negative feedbacks experienced by AMF trees may lead to healthier and more diverse forests in the grand scheme of things. 

Infographic by [1]

Further Reading: [1]



On Parasites and Diversity


We all too readily demonize parasites. It is kind of understandable though. The thought of something living in or on you at your expense is enough to make our skin crawl. There are a lot of evolutionary pressures that make us look unfavorably about organisms with such lifestyles. However, to completely write parasites off as a bane to life as we know it may be a huge mistake on our part. More and more we are realizing that parasites play an important role in ecosystem functioning and may even serve as indicators of environmental health. 

Plants are no stranger to such parasitic dynamics. Many species have forgone some if not all photosynthetic ability in exchange for a parasitic lifestyle. There is no question that plant parasites can and do have net negative effects on their hosts, however, its never that simple. Research is showing that parasitic plants can have profound effects on the structure and productivity of surrounding plant communities. 

For starters, parasitic plants can increase the competitive ability of non-host species. By knocking back the performance of their host, other plant species can pick up the slack so-to-speak. This can often lead to an increase in overall plant diversity in a given habitat. A common thread throughout studies that have looked at parasitic plants is that proportion of grasses declined when parasitic plants were present. This made room for less competitive forbs to increase in number. In effect, parasitic plants can level the playing field for other, less competitive plant species. 

By altering ecosystem structure, parasitic plants can also alter the way nutrients flow through the system. This can have some seriously profound ramifications. For instance, the presence of the hemiparasitic Rhinanthus minor in grasslands has been shown to  increasing rates of nitrogen cycling. Though the ramifications of this are dynamic, it is nonetheless proof that parasites should not simply be maligned and that, despite our perspective, nature is far more complex than we realize. 

Photo Credit: Sannse (Wikimedia Commons)

Further Reading:


Meet Euphrasia nemorosa, the eyebright. This lovely little plant is native to the northern regions of North America. A quick glance at the flowers of this species may seem to suggest a member of the mint family but this would be wrong. Once placed in Scrophulariaceae, it is now thought to reside in Orobanchaceae. Like other members of this group, E. nemorosa is a hemiparasite. It uses specialized roots to tap into the roots of plants growing around it. In the wild, research has shown that E. nemorosa seems to prefer to parasitize grasses but laboratory experiments have shown that it will parasitize a variety of plants if given the chance. It can even grow without parasitizing other plants but those that did grew small and weak. 

Parasitic plants are an interesting bunch. They push the limits of what is traditionally accepted in the realm of plant physiology. Non-parasitic plants usually have to balance between CO2 uptake and water loss. They do this by controlling their stomata, which are tiny openings on the leaf. Because they are attached to a host, parasitic plants do not have to worry about minimizing water loss and instead want to maximize water loss to gain as much carbon from their host plant as possible. 

Another interesting aspect of Euphrasia ecology is their preference for disturbance. Euphrasia are plants of disturbed meadows, fields, and man-made habitats. There is a lot of work being done to examine which kinds of species thrive in and around humans. Research has shown that by selecting for native species like Euphrasia, the species composition on these types of disturbed habitats can take on a more biodiverse character instead of the usual non-native monoculture.

Further Reading: