Maples, Epiphytes, and a Canopy Full of Goodies

The forests of the Pacific Northwest are known for the grandeur. This region is home to one of the greatest temperate rainforests in the world. A hiker is both dwarfed and enveloped by greenery as soon as they hit the trail. One aspect of these forests that is readily apparent are the carpets of epiphytes that drape limbs and branches all the way up into the canopy. Their arboreal lifestyle is made possible by a combination of mild winters and plenty of precipitation. 

We are frequently taught that the relationship between trees and their epiphytes are commensal - the epiphytes get a place to live and the trees are no worse for wear. However, there are a handful of trees native to the Pacific Northwest that are changing the way we think about the relationship between these organisms in temperate rainforests.

Though conifers dominate the Pacific Northwest landscape, plenty of broad leaved tree species abound. One of the most easily recognizable is the bigleaf maple (Acer macrophyllum). Both its common and scientific names hint at its most distinguishing feature, its large leaves. Another striking feature of this tree are its epiphyte communities. Indeed, along with the vine maple (A. circinatum), these two tree species carry the greatest epiphyte to shoot biomass ratio in the entire forest. Numerous species of moss, liverworts, lichens, and ferns have been found growing on the bark and branches of these two species.

Epiphyte loads are pretty intense. One study found that the average epiphyte crop of a bigleaf maple weighs around 78 lbs. (35.5 Kg). That is a lot of biomass living in the canopy! The trees seem just fine despite all of that extra weight. In fact, the relationship between bigleaf and vine maples and their epiphyte communities run far deeper than commensalism. Evidence accumulated over the last few decades has revealed that these maples are benefiting greatly from their epiphytic adornments.

Rainforests, both tropical and temperate, generally grow on poor soils. Lots of rain and plenty of biodiversity means that soils are quickly leached of valuable nutrients. Any boost a plant can get from its environment will have serious benefits for growth and survival. This is where the epiphytes come in. The richly textured mix of epiphytic plants greatly increase the surface area of any branch they live on. And all of that added surface area equates to more nooks and crannies for water and dust to get caught and accumulate.

Photo by SuperFantastic licensed under CC BY 2.0

When researchers investigated just how much of a nutrient load gets incorporated into these epiphyte communities, the results painted quite an impressive picture. On a single bigleaf maple, epiphyte leaf biomass was 4 times that of the host tree despite comprising less than 2% of the tree's above ground weight. All of that biomass equates to a massive canopy nutrient pool rich in nitrogen, phosphorus, potassium, calcium, magnesium, and sodium. Much of these nutrients arrive in the form of dust-sized soil particles blowing around on the breeze. What's more, epiphytes act like sponges, soaking up and holding onto precious water well into the dry summer months.

Now its reasonable to think that nutrients and water tied up in epiphyte biomass would be unavailable to trees. Indeed, for many species, epiphytes may slow the rate at which nutrients fall to and enter into the soil. However, trees like bigleaf and vine maples appear to be tapping into these nutrient and water-rich epiphyte mats.

A subcanopy of vine maple (Acer circinatum) draped in epiphytes.

Both bigleaf and vine maples (as well as a handful of other tree species) are capable of producing canopy roots. Wherever the epiphyte load is thick enough, bundles of cells just under the bark awaken and begin growing roots. This is a common phenomenon in the tropics, however, the canopy roots of these temperate trees differ in that they are indistinguishable in form and function from subterranean roots.

Canopy roots significantly increase the amount of foraging an individual tree can do for precious water and nutrients. Additionally, it has been found that canopy roots of the bigleaf maple even go as far as to partner with mycorrhizal fungi, thus unlocking even more potential for nutrient and water gain. In the absence of soil nutrient and water pools, a small handful of trees in the Pacific Northwest have unlocked a massive pool of nutrients located above us in the canopy. Amazingly, it has been estimated that mature bigleaf and vine maples with well developed epiphyte communities may actually gain a substantial fraction of their water and nutrient needs via their canopy roots.

Photo Credits: [2]

Further Reading: [1] [2] [3] [4] [5]

Of Bluebells and Fungi

Photo by Christophe Couckuyt licensed under CC BY 2.0

Whether in your garden or in the woods, common bluebells (Hyacinthoides non-scripta) are a delightful respite from the dreary months of winter. It should come as no surprise that these spring geophytes are a staple in temperate gardens the world over. And, as amazing as they are in the garden, bluebells are downright fascinating in the wild.

Bluebells can be found growing naturally from the northwestern corner of Spain north into the British Isles. They are largely a woodland species, though finding them in meadows isn't uncommon. They are especially common in sites that have not experienced much soil disturbance. In fact, large bluebell populations are used as indicators of ancient wood lots.

Photo by RX-Guru licensed under CC BY-SA 3.0

Being geophytes, bluebells cram growth and reproduction into a few short weeks in spring. We tend to think of plants like this as denizens of shade, however, most geophytes get going long before the canopy trees have leafed out. As such, these plants are more accurately sun bathers. On warm days, various bees can be seen visiting the pendulous flowers, with the champion pollinator being the humble bumble bees.

The above ground beauty of bluebells tends to distract us from learning much about their ecology. That hasn't stopped determined scientists though. Plenty of work has been done looking at how bluebells make their living and get on with their botanical neighbors. In fact, research is turning up some incredible data regarding bluebells and mycorrhizal fungi.

Photo by Mick Garratt licensed under CC BY-SA 2.0

Bluebell seeds tend not to travel very far, most often germinating near the base of the parent. Germination occurs in the fall when temperatures begin to drop and the rains pick up. Interestingly, bluebell seeds actually germinate within the leaf litter and begin putting down their initial root before the first frosts. Often this root is contractile, pulling the tiny seedling down into the soil where it is less likely to freeze. During their first year, phosphorus levels are high. Not only does the nutrient-rich endosperm supply the seedling with much of its initial needs, abundant phosphorus near the soil surface supplies more than enough for young plants. This changes as the plants age and change their position within the soil.

Photo by MichaelMaggs licensed under CC BY-SA 3.0

Over the next 4 to 5 years, the bluebell's contractile roots pull it deeper down into the soil, taking it out of the reach of predators and frost. This also takes them farther away from the nutrient-rich surface layers. What's more, the roots of older bluebells are rather simple structures. They do not branch much, if at all, and they certainly do not have enough surface area for proper nutrient uptake. This is where mycorrhizae come in.

Bluebells partner with a group of fungi called arbuscular mycorrhiza, which penetrate the root cells, thus greatly expanding the effective rooting zone of the plant. Plants pay these fungi in carbohydrates produced during photosynthesis and in return, the fungi provide the plants with access to far more nutrients than they would be able to get without them. One of the main nutrients plants gain from these symbiotic fungi is phosphorus.

Photo by Oast House Archive licensed under CC BY-SA 2.0

For bluebells, with age comes new habitat, and with new habitat comes an increased need for nutrients. This is why bluebells become more dependent on arbuscular mycorrhiza as they age. In fact, plants grown without these fungi do not come close to breaking even on the nutrients needed for growth and maintenance and thus live a shortened life of diminishing returns. This is an opposite pattern from what we tend to expect out of mycorrhizal-dependent plants. Normally its the seedlings that cannot live without mycorrhizal symbionts. It just goes to show you that even familiar species like the bluebell can offer us novel insights into the myriad ways in which plants eke out a living.

Photo Credits: [1] [2] [3] [4] [5]

Further Reading: [1] [2]

Invasion of the Earthworms

Photo by Rob Hille licensed under CC BY-SA 3.0

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 North America's native ecosystems have evolved without the influence of any type of worms in the soil.

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 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 extremely well. When earthworms invade an area of a forest, they disrupt this rich, organic layer in very serious ways.

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 myriad 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 take a hit. Spring ephemerals have been hit the hardest by earthworm invasions for these reasons and more. There is very little recruitment and, in time, many species are lost. For small seeded species like orchids, earthworms can even consume seeds, which either destroys them outright or drags them down deeper into the soil where they cannot germinate. 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.

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

A Brief Primer on the Vernal Dam Hypothesis

Arctic Bone Nurseries

Life and death are two sides of the same coin. In death, an organisms body is broken down into its constituent parts and redistributed throughout the environment. As such, decomposition is a major player in the global cycling of nutrients. Nowhere does this become more apparent than in nutrient limited habitats like the Arctic Tundra.

The Arctic is known for being a very harsh place to live. A combination of low temperatures, low water availability, and short summers make for tough conditions for any plant. What's more, low temperatures and water availability also mean nutrients are hard to get at. It stands to reason then that any potential uptick in nutrient availability would be a boon for Arctic plant life.

This is where dead animals come in. When a large animal like a muskox dies, its body can take many years to break down. Each summer, as temperatures rise above freezing, decomposition slowly eats away at the tissues. Research has found that nutrient levels, specifically nitrogen, are much higher within a meter radius around a decomposing carcass. Moreover, plants in this region were found to have higher nitrogen levels in their tissues and achieved the most luscious growth.

Nutrients aren't the only benefit carcasses provide. They also offer a favorable microclimate. Many herbivores instinctually avoid feeding around dead animals as a way of limiting exposure to disease. Research has found that grazing levels are much lower around most carcasses.

Another benefit is shelter. Wind is an ever present force to reckon with on the tundra. Carcasses provide a sheltered area that serves as an oasis for seeds to germinate and grow. The carcass also acts like a filter, collecting debris and allowing soils to build over time. Other animals may find this a favorable place to hide or hunt and thus the importance of these carcass islands becomes all the more apparent.

Photo Credits: Neil Shubin 

Further Reading:
http://pubs.aina.ucalgary.ca/arctic/arctic55-4-389.pdf