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]

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]

 

 

Hyperabundant Deer Populations Are Reducing Forest Diversity

Synthesizing the effects of white-tailed deer on the landscape have, until now, been difficult. Although strong sentiments are there, there really hasn't been a collective review that indicates if overabundant white-tailed deer populations are having a net impact on the ecosystem. A recent meta-analysis published in the Annals of Botany: Plant Science Research aimed to change that. What they have found is that the overabundance of deer is having strong negative impacts on forest understory plant communities in North America.

White-tailed deer have become a pervasive issue on this continent. With an estimated population of well over 30 million individuals, deer have been managed so well that they have reached proportions never seen on this continent in the past. The effects of this hyper abundance are felt all across the landscape. As anyone who gardens will tell you, deer are voracious eaters.

Tackling this issue isn't easy. Raising questions about proper management in the face of an ecological disaster that we have created can really put a divide in the room. Even some of you may be experiencing an uptick in your blood pressure simply by reading this. Feelings aside, the fact of the matter is overabundant deer are causing a decline in forest diversity. This is especially true for woody plant species. Deer browsing at such high levels can reduce woody plant diversity by upwards of 60%. Especially hard hit are seedlings and saplings. In many areas, forests are growing older without any young trees to replace them.

What's more, their selectivity when it comes to what's on the menu means that forests are becoming more homogenous. Grasses, sedges, and ferns are increasingly replacing herbaceous cover gobbled up by deer. Also, deer appear to prefer native plants over invasives, leaving behind a sea of plants that local wildlife can't readily utilize. It's not just plants that are affected either. Excessive deer browse is creating trophic cascades that propagate throughout the food web.

For instance, birds and plants are intricately linked. Flowers attract insects and eventually produce seeds. These in turn provide food for birds. Shrubs provide food as well as shelter and nesting space, a necessary requisite for healthy bird populations. Other studies have shown that in areas that experience the highest deer densities songbird populations are nearly 40% lower than in areas with smaller deer populations. As deer make short work of our native plants, they are hurting far more than just the plants themselves. Every plant that disappears from the landscape is one less plant that can support wildlife.

Sadly, due to the elimination of large predators from the landscape, deer have no natural checks and balances on their populations other than disease and starvation. As we replace natural areas with manicured lawns and gardens, we are only making the problem worse. Deer have adapted quite well to human disturbance, a fact not lost on anyone who has had their garden raided by these ungulates. Whereas the deer problem is only a piece of the puzzle when it comes to environmental issues, it is nonetheless a large one. With management practices aimed more towards trophy deer than healthy population numbers, it is likely this issue will only get worse.

Photo Credit: tuchodi (http://bit.ly/1wFYh2X)

Further Reading:
http://aobpla.oxfordjournals.org/content/7/plv119.full

http://aobpla.oxfordjournals.org/content/6/plu030.full

http://www.sciencedirect.com/science/article/pii/S0006320705001722