The Only True Cedars

  Cedrus deodara

Cedrus deodara

The only true cedars on this planet can be found growing throughout mountainous regions of the western Himalayas and Mediterranean. All others are cedars by name only. The so-called “cedars” we encounter here in North America are not even in the same family as the true cedars. Instead, they belong to the Cypress family (Cupressaceae). The true cedars all belong to the genus Cedrus and are members of the family Pinaceae. They are remarkable trees with tons of ecological and cultural value.

 J. White,1803-1824.

J. White,1803-1824.

The true cedars are stunning trees to say the least. They regularly reach heights of 100 ft. (30 m.) or more and can live for thousands of years. Cedars exhibit a dimorphic branching structure, with long shoots forming branches and smaller shoots carrying most of the needle load. The needles themselves are borne in dense, spiral clusters, giving the branches a rather aesthetic appearance. Each needle produces layers of wax, which vary in thickness depending on how exposed the tree is growing. This waxy layer helps protect the tree from sunburn and desiccation.

  Cedrus libani

Cedrus libani

  Cedrus libani

Cedrus libani

One of the easiest ways to identify a cedar is by checking out its cones. All members of the genus Cedrus produce upright, barrel-shaped cones. Male cones are smaller and don’t stay on the tree for very long. Female cones, on the other hand, are quite large and stay on the tree until the seeds are ripe. Upon ripening, the entire female cone disintegrates, releasing the seeds held within. Each seed comes complete with blisters full of distasteful resin, which is thought to deter seed predators.

 Male cones of  Cedrus atlantica

Male cones of Cedrus atlantica

 Female  Cedrus  cones.

Female Cedrus cones.

In total, there are only four recognized species of cedar - the Atlas cedar (Cedrus atlantica), the Cyprus cedar (C. brevifolia), the deodar cedar (C. deodara), and the Lebanon cedar (C. libani). I have heard arguments that C. brevifolia is no more than a variant of C. libani but I have yet to come across any source that can say this for certain. Much more work is needed to assess the genetic structure of these species. Even their place within Pinaceae is up for debate. Historically it seems that Cedrus has been allied with the firs (genus Abies), however, work done in the early 2000’s suggests that Cedrus may actually be sister to all other Pinaceae. We need more data before anything can be said with certainty.

  Cedrus atlantica

Cedrus atlantica

Regardless, two of these cedars - C. atlantica & C. libani - are threatened with extinction. Centuries of over-harvesting, over-grazing, and unsustainable fire regimes have taken their toll on wild populations. Much of what remains is not considered old growth. Gone is the heyday of giant cedar forests. Luckily, many populations are now located in protected areas and reforestation efforts are being put into place throughout their range. Still, the ever present threat of climate change is causing massive pest outbreaks in these forests. The future for these trees hangs in the balance.

Photo Credit: Wikimedia Commons

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

Maples, Epiphytes, and a Canopy Full of Goodies

IMG_0090.jpg

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. 

Weare 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.

Acer_macrophyllum_1214.JPG

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.

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.

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: [1] [2]

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

 

The Curious Case of the Yellowwood Tree

17975570426_efd4213146_o.jpg

The immense beauty and grace of the yellowwood (Cladrastis kentukea) is inversely proportional to its abundance. This unique legume is endemic to the eastern United States and enjoys a strangely patchy distribution. Its ability to perform well when planted far outside of its natural range only deepens the mystery of the yellowwood.

The natural range of the yellowwood leaves a lot of room for speculation. It hits its highest abundances in the Appalachian and Ozark highlands where it tends to grow on shaded slopes in calcareous soils. Scattered populations can be found as far west as Oklahoma and as far north as southern Indiana but nowhere is this tree considered a common component of the flora.

Cladrastis_kentukea_range_map_1.png

Though the nature of its oddball distribution pattern is open for plenty of speculation, it is likely that its current status is the result of repeated glaciation events and a dash of stochasticity. The presence of multiple Cladrastis species in China and Japan and only one here in North America is a pattern shared by multiple taxa that once grew throughout each continent. A combination of geography, topography, and repeated glaciation events has since fragmented the ranges of many genera and perhaps Cladrastis is yet another example.

The fact that yellowwood seems to do quite well as a specimen tree well outside of its natural range says to me that this species was probably once far more wide spread in North America than it was today. It may have been pushed south by the ebb and flow of the Laurentide Ice Sheet and, due to the stochastic nuances of seed dispersal, never had a chance to recolonize the ground it had lost. Again, this is all open to speculation as this point.

17822773749_905eb83e49_o.jpg

Despite being a member of the pea family, yellowwood is not a nitrogen fixer. It does not produce nodules on its roots that house rhizobium. As such, this species may be more restricted by soil type than other legumes. Perhaps its inability to fix nitrogen is part of the reason it tends to favor richer soils. It may also have played a part in its failure to recolonize land scraped clean by the glaciers.

Yellowwood's rarity in nature only makes finding this tree all the more special. It truly is a site to behold. It isn't a large tree by any standards but what it lacks in height it makes up for in looks. Its multi-branched trunk exhibits smooth, gray bark reminiscent of beech trees. Each limb is decked out in large, compound leaves that turn bright yellow in autumn.

When mature, which can take upwards of ten years, yellowwood produces copious amounts of pendulous inflorescences. Each inflorescence sports bright white flowers with a dash of yellow on the petals. It doesn't appear that any formal pollination work has been done on this tree but surely bees and butterflies alike visit the blooms. The name yellowwood comes from the yellow coloration of its heartwood, which has been used to make furniture and gunstocks in the past.

Whether growing in the forest or in your landscape, yellowwood is one of the more stunning trees you will find in eastern North America. Its peculiar natural history only lends to its allure.

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

Further Reading: [1] [2]

Life On a Floodplain

Floodplains can be pretty rough places for plant life. Despite readily a available water supply, the unpredictable, disturbance-prone nature of these habitats means that static lifeforms such as plants need to be quite adaptable to survive and persist. Join In Defense of Plants for a brief look at this sort of ecosystem.

Producer, Editor, Camera: Grant Czadzeck (http://www.grantczadzeck.com)

Music by
Artist: Somali Yacht Club
Track: Up In The Sky
http://somaliyachtclub.bandcamp.com

Saving Bornean Peatlands is a Must For Conservation

Borneo_rainforest.jpg

The leading cause of extinction on this planet is loss of habitat. As an ecologist, it pains me to see how frequently this gets ignored. Plants, animals, fungi - literally every organism on this planet needs a place to live. Without habitat, we are forced to pack our flora and fauna into tiny collections in zoos and botanical gardens, completely disembodied from the environment that shaped them into what we know and love today. That’s not to say that zoos and botanical gardens don’t play critically important roles in conservation, however, if we are going to stave off total ecological meltdown, we must also be setting aside swaths of land.

There is no way around it. We cannot have our cake and eat it too. Land conservation must be a priority both at the local and the global scale. Wild spaces support life. They buffer it from storms and minimize the impacts of deadly diseases. Healthy habitats filter the water we drink and, for many people around the globe, provide much of the food we eat. Every one of us can think back to our childhood and remember a favorite stretch of stream, meadow, or forest that has since been gobbled up by a housing development. For me it was a forested stream where I learned to love the natural world. I would spend hours playing in the creek, climbing trees, and capturing bugs to show my parents. Since that time, someone leveled the forest, built a house, and planted a lawn. With that patch of forest went all of the insects, birds, and wildflowers it once supported.

Logging_road_East_Kalimantan_2005.jpg
5551935164_127180a252_b.jpg

Scenarios like this play out all too often and sadly on a much larger scale than a backyard. Globally, forests have felt taken the brunt of human development. Though it is hard to get a sense of the scope of deforestation on a global scale, the undisputed leaders in deforestation are Brazil and Indonesia. Though the Amazon gets a lot of press, few may truly grasp the gravity of the situation playing out in Southeast Asia.

Deforestation is a clear and present threat throughout tropical Asia. This region is growing both in its economy and population by about 6% every year and this growth has come at great cost to the environment. Indonesia (alongside Brazil) accounts for 55% of the world’s deforestation rates. This is a gut-wrenching statistic because Indonesia alone is home to the most extensive area of intact rainforest in all of Asia. So far, nearly a quarter of Indonesia’s forests have been cleared. It was estimated that by 2010, 2.3 million hectares of peatland forests had been felled and this number shows little signs of slowing. Experts believe that if these rates continue, this area could lose the remainder of its forests by 2056.

Consider the fact that Southeast Asia contains 6 of the world’s 25 biodiversity hotspots and you can begin to imagine the devastating blow that the levelling of these forests can have. Much of this deforestation is done in the name of agriculture, and of that, palm oil and rubber take the cake. Southeast Asia is responsible for 86% of the world’s palm oil and 87% of the world’s natural rubber. What’s more, the companies responsible for these plantations are ranked among some of the least sustainable in the world.

 Palm oil plantations where there once was rainforest. 

Palm oil plantations where there once was rainforest. 

Borneo is home to a bewildering array of life. Researchers working there are constantly finding and describing new species, many of which are found nowhere else in the world. Of the roughly 15,000 plant species known from Borneo, botanists estimate that nearly 5,000 (~34%) of them are endemic. This includes some of the more charismatic plant species such as the beloved carnivorous pitcher plants in the genus Nepenthes. Of these, 50 species have been found growing in Borneo, many of which are only known from single mountain tops.

It has been said that nowhere else in the world has the diversity of orchid species found in Borneo. To date, roughly 3,000 species have been described but many, many more await discovery. For example, since 2007, 51 new species of orchid have been found. Borneo is also home to the largest flower in the world, Rafflesia arnoldii. It, along with its relatives, are parasites, living their entire lives inside of tropical vines. These amazing plants only ever emerge when it is time to flower and flower they do! Their superficial resemblance to a rotting carcass goes much deeper than looks alone. These flowers emit a fetid odor that is proportional to their size, earning them the name “carrion flowers.”

  Rafflesia arnoldii  in all of its glory.

Rafflesia arnoldii in all of its glory.

Phalaenopsis_bellina_Orchi_01.jpg

If deforestation wasn’t enough of a threat to these botanical treasures, poachers are having considerable impacts on Bornean botany. The illegal wildlife trade throughout southeast Asia gets a lot of media attention and rightfully so. At the same time, however, the illegal trade of ornamental and medicinal plants has gone largely unnoticed. Much of this is fueled by demands in China and Vietnam for plants considered medicinally valuable. At this point in time, we simply don’t know the extent to which poaching is harming plant populations. One survey found 347 different orchid species were being traded illegally across borders, many of which were considered threatened or endangered. Ever-shrinking forested areas only exacerbate the issue of plant poaching. It is the law of diminishing returns time and time again.

Paphiopedilum_philippinense_Orchi_021.jpg

But to lump all Bornean forests under the general label of “rainforest” is a bit misleading. Borneo has multitude of forest types and one of the most globally important of these are the peatland forests. Peatlands are vital areas of carbon storage for this planet because they are the result of a lack of decay. Whereas leaves and twigs quickly breakdown in most rainforest situations, plant debris never quite makes it that far in a peatland. Plant materials that fall into a peatland stick around and build up over hundreds and thousands of years. As such, an extremely thick layer of peat is formed. In some areas, this layer can be as much as 20 meters deep! All the carbon tied up in the undecayed plant matter is carbon that isn’t finding its way back into our atmosphere.

Sadly, tropical peatlands like those found in Borneo are facing a multitude of threats. In Indonesia alone, draining, burning, and farming (especially for palm oil) have led to the destruction of 1 million hectares (20%) of peatland habitat in only one decade. The fires themselves are especially worrisome. For instance, it was estimated that fires set between 1997-1998 and 2002-2003 in order to clear the land for palm oil plantations released 200 million to 1 billion tonnes of carbon into our atmosphere. Considering that 60% of the world’s tropical peatlands are found in the Indo-Malayan region, these numbers are troubling.

Peat_Forest_Swamp_(10712654875).jpg

The peatlands of Borneo are totally unlike peatlands elsewhere in the world. Instead of mosses, gramminoids, and shrubs, these tropical peatlands are covered in forests. Massive dipterocarp trees dominate the landscape, growing on a spongey mat of peat. What’s more, no water flows into these habitats. They are fed entirely by rain. The spongey nature of the peat mat holds onto water well into the dry season, providing clean, filtered water where it otherwise wouldn’t be available.

This lack of decay coupled with their extremely acidic nature and near complete saturation makes peat lands difficult places for survival. Still, life has found a way, and Borneo’s peatlands are home to a staggering diversity of plant life. They are so diverse, in fact, that when I asked Dr. Craig Costion, a plant conservation officer for the Rainforest Trust, for something approaching a plant list for an area of peatland known as Rungan River region, he replied:

“Certainly not nor would there ever be one in the conceivable future given the sheer size of the property and the level of diversity in Borneo. There can be as many as a 100 species per acre of trees in Borneo... Certainly a high percentage of the species would only be able to be assigned to a genus then sit in an herbarium for decades until someone describes them.”

And that is quite remarkable when you think about it. When you consider that the Rungan River property is approximately 385,000 acres, the number of plant species to consider quickly becomes overwhelming. To put that in perspective, there are only about 500 tree species native to the whole of Europe! And that’s just considering the trees. Borneo’s peatlands are home to myriad plant species from liverworts, mosses, and ferns, to countless flowering plants like orchids and others. We simply do not know what kind of diversity places like Borneo hold. One could easily spend a week in a place like the Rungan River and walk away with dozens of plant species completely new to science. Losing a tract of forest in such a biodiverse is a huge blow to global biodiversity.

Headhunter's_trail_Mulu_N._bicalcarata_3.jpg
  Nepenthes ampullaria  relies on decaying plant material within its pitcher for its nutrient needs.

Nepenthes ampullaria relies on decaying plant material within its pitcher for its nutrient needs.

Also, consider that all this plant diversity is supporting even more animal diversity. For instance, the high diversity of fruit trees in this region support a population of over 2,000 Bornean orangutans. That is nearly 4% of the entire global population of these great apes! They aren’t alone either, the forested peatlands of Borneo are home to species such as the critically endangered Bornean white-bearded gibbon, the proboscis monkey, the rare flat-headed cat, and the oddly named otter civet. All these animals and more rely on the habitat provided by these forests. Without forests, these animals are no more.

Pongo_tapanuliensis_female.jpg
 The flat-headed cat, an endemic of Borneo. 

The flat-headed cat, an endemic of Borneo. 

At this point, many of you may be feeling quite depressed. I know how easy it is to feel like there is nothing you can do to help. Well, what if I told you that there is something you can do right now to save a 385,000 acre chunk of peatland rainforest? That’s right, by heading over to the Rainforest Trust’s website (https://www.rainforesttrust.org/project/saving-stronghold-critically-endangered-bornean-orangutan/) you can donate to their campaign to buy up and protect the Rungan River forest tract.

 Click on the logo to learn more!

Click on the logo to learn more!

By donating to the Rainforest Trust, you are doing your part in protecting biodiversity in one of the most biodiverse regions in the world. What’s more, you can rest assured that your money is being used effectively. The Rainforest Trust consistently ranks as one of the top environmental protection charities in the world. Over their nearly three decades of operation, the Rainforest Trust has protected more than 15.7 million acres of land in over 20 countries. Like I said in the beginning, habitat loss is the leading cause of extinction on this planet. Without habitat, we have nothing. Plants are that habitat and by supporting organizations such as the Rainforest Trust, you are doing your part to fight the biggest threats our planet faces. 

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

Photo Credits: [1] [2] [3] [4] [5] [6] [7] [8] [9] [10] [11] [12]

Fish: The Unsung Heroes of Seed Dispersal

 Fruits of the tucum palm.

Fruits of the tucum palm.

It goes without saying that effective seed (and spore) dispersal is vital for thriving plant populations. Without it, plant populations will stagnate and disappear. Whereas we know quite a bit about the role animals like birds, bats, and ants play in this process, there is another group of seed dispersers that are proving to be vital to the long-term health and survival of tropical forests around the globe - fish. 

The idea of seed dispersing fish may come as a shock to some but mounting evidence is showing that fruit-eating fish play a major role in the reproductive cycle of many tropical plant species. This is especially true in seasonally flooded tropical forests. To date, more than 100 different fish species have been found with viable seeds in their guts. In fact, some fish species, such as the pacu (Piaractus mesopotamicus), specialize on eating fruits.

 A big ol' pacu looking for its next fruit meal.

A big ol' pacu looking for its next fruit meal.

By monitoring how fruit-eating fish like the pacu behave in their environment, scientists are painting a picture of tropical seed dispersal that is quite remarkable. Take, for instance, the tucum palm (Bactris glaucescens). Native to Brazil's Pantanal, this palm produces large, red fruits and everything from peccaries to iguanas will consume them. However, when eaten by these animals, the seed either don't make it through the gut in one piece or they end up being pooped out into areas unsuitable for germination. Only when the seeds have been consumed by the pacu do they end up in the right place in the right condition. It appears that pacus are the main seed dispersal agent for this palm. 

 A beautiful tucum palm in the dry season.

A beautiful tucum palm in the dry season.

The tucum palm isn't alone either. The seeds of myriad other plant species known to inhabit such seasonally flooded habitats seem to germinate and grow most effectively only after having been dispersed by fish. Pacus are also responsible for a considerable amount of seed dispersal for plants such as Tocoyena formosa (Rubiaceae), Licania parvifolia (Chrysobalanaceae), and Inga uruguensis (Fabaceae). Even outside of the tropics, fish like the channel catfish (Ictalurus punctatus) are being found to be important seed dispersers of riparian plants such as the eastern swampprivet (Forestiera acuminata).

 Camu-camu ( Myrciaria dubia )

Camu-camu (Myrciaria dubia)

Without fish, these plants would have a hard time with seed dispersal in these types of habitats. Without something moving them around, these seeds would be stuck at the bottom of a river, buried in anoxic mud. As fish migrate into flooded forests, they can move seeds remarkable distances from their parents. When the floods recede, the seeds find themselves primed and ready to usher in the next generation.

 Fruits of the Camu-camu ( Myrciaria dubia ) also benefit from dispersal by fish.

Fruits of the Camu-camu (Myrciaria dubia) also benefit from dispersal by fish.

Not all fish perform this task equally as well. Even within a species, there are differences in the effectiveness of seed dispersal services. Scientists are finding that large fish are most effective at proper seed dispersal. Not only can they consume whole fruits with little to no issue, they are also the fish that are most physically capable of moving large distances. Sadly, humans are seriously disrupting this process in a lot of ways.

For starters, dams and other impediments are cutting off the migratory routs of many fish species. Large fish are no longer able to make it into flooded regions of forest far upstream once a dam is in place. What's more, dams keep large tracts of forest from flooding entirely. As such, fish are no longer able to migrate into these regions, which means less seeds are making it there as well. This is bad news for forest regeneration.

 "Gimme fruit" says local channel cat.

"Gimme fruit" says local channel cat.

It's not just dams hurting fish either. Over-fishing is a serious issue in most water ways. Pacus, for instance, have seen precipitous declines throughout the Amazon over the last few decades. Specifically targeted are large fish. Unfortunately, regulations that were put into place in order to help these fish may actually be harming their seed dispersal activities. Fish under a certain size must be released from any catch, thus a disproportionate amount of large fish are being removed from the system.

Logging is taking quite a toll as well. Floodplain forests have been hit especially hard by logging, both legal and illegal. The lower Amazon River, for example, has almost no natural floodplain forests left. Reports from fish markets in these areas have shown fewer and fewer frugivorous fish each year. It would appear that large fruit-eating fish are disappearing in the areas that need seed dispersal the most. It is clear that something drastic needs to happen. At the very least, fruit-eating fish need more recognition for the ecosystem services they provide.

Forest health and management is a holistic endeavor. We cannot think of organisms in isolation. This is why ecological literacy is so important. We are only now starting to realize the role of large fish in forest regeneration and who knows what kinds of discoveries are just over the horizon. This is why land conservation efforts are so important. We must move to protect wild spaces before they are lost for good. Please consider donating to one of the many great land conservancy agencies around the globe. 

Photo Credits: [1] [2] [3] [4] [5] [6]

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

 

How Trees Fight Disease

27329418_10101829159766245_2047871682_o.jpg

Plants do not have immune systems like animals. Instead, they have evolved an entirely different way of dealing with infections. In trees, this process is known as the "compartmentalization of decay in trees" or "CODIT." CODIT is a fascinating process and many of us will recognize its physical manifestations.

In order to understand CODIT, one must know a little something about how trees grow. Trees have an amazing ability to generate new cells. However, they do not have the ability to repair damage. Instead, trees respond to disease and injury  by walling it off from their living tissues. This involves three distinct processes. The first of these has to do with minimizing the spread of damage. Trees accomplish this by strengthening the walls between cells. Essentially this begins the process of isolating whatever may be harming the living tissues.

This is done via chemical means. In the living sapwood, it is the result of changes in chemical environment within each cell. In heartwood, enzymatic changes work on the structure of the already deceased cells. Though the process is still poorly understood, these chemical changes are surprisingly similar to the process of tanning leather. Compounds like tannic and gallic acids are created, which protect tissues from further decay. They also result in a discoloration of the surrounding wood. 

The second step in the CODIT process involves the construction of new walls around the damaged area. This is where the real compartmentalization process begins. The cambium layer changes the types of cells it produces around the area so that it blocks that compartment off from the surrounding vascular tissues. These new cells also exhibit highly altered metabolisms so that they begin to produce even more compounds that help resist and hopefully stave off the spread of whatever microbes may be causing the injury. Many of the defects we see in wood products are the result of these changes.

CODIT.JPG

The third response the tree undergoes is to keep growing. New tissues grow around the infected compartment and, if the tree is healthy enough, will outpace further infection. You see, whether its bacteria, fungi, or a virus, microbes need living tissues to survive. By walling off the affected area and pumping it full of compounds that kill living tissues, the tree essentially cuts off the food supply to the disease-causing organism. Only if the tree is weakened will the infection outpace its ability to cope.

Of course, CODIT is not 100% effective. Many a tree falls victim to disease. If a tree is not killed outright, it can face years or even decades of repeated infection. This is why we see wounds on trees like perennial cankers. Even if the tree is able to successfully fight these repeat infections over a series of years, the buildup of scar tissues can effectively girdle the tree if they are severe enough.

CODIT is a well appreciated phenomenon. It has set the foundation for better tree management, especially as it relates to pruning. It is even helping us develop better controls against deadly invasive pathogens. Still, many of the underlying processes involved in this response are poorly understood. This is an area begging for deeper understanding.

Photo Credits: kaydubsthehikingscientist & Alex Shigo

Further Reading: [1]

Devil's Gardens

Tococa_leaves.jpg

Imagine, if you will, walking through the dense understory somewhere in the Amazon basin. Diversity reigns supreme here and it would seem that every few steps reveals myriad new plant species. As you walk along, something in the vegetation changes. You stumble into a clearing in the middle of the forest dominated entirely by a single species of tree. Why the sudden change? How did this monoculture develop? You, my friend, have just found yourself on the edge of a Devil's garden. 

Devil's gardens are said to be the resting place of an evil spirit known to local tribes as Chullachaki. Anyone unlucky enough to stumble into his garden is said at risk of attack or curse. In reality, these gardens have a biological origin. The real gardeners are a handful of ant species which seem to have rather specific gardening preferences. Careful inspection would reveal that the gardens largely consist of trees in one of three genera - Duroia, Tococa, or Clidemia

  Tococa  sp. (Melastomataceae)

Tococa sp. (Melastomataceae)

The reason that ants are so fond of these genera has to do with housing. These plant groups contain species which produce swellings along their stems and petioles known as domatia. These domatia are hollow and are the favorite nesting spots of various ant species. Ant colonies set up shop within. As anyone who has ever blundered into an ant colony can attest, ants are quite voracious at defending their home. 

By providing ant colonies with a home base, these plants have essentially hired body guards. It is a wonderful form of symbiosis in which the ants aggressively defend against anything that might want to take a bite out of their host tree. Any herbivore trying to take up residence or lay eggs within the Devil's garden is viciously attacked. In doing so, the ants are protecting their host trees at the cost of all other plants unlucky enough to germinate within the garden. Still, this anti-herbivore behavior doesn't totally explain the monoculture status these host trees achieve within the garden itself. Why are these gardens so ominously devoid of other plant species?

To answer this, one would have to watch how the ants behave as they forage. While scouting, if ants encounter a seedling of their host tree, nothing really happens. They go about their business and let the seedling grow into a future home. When they encounter a non-host tree, however, their behavior completely changes. 

 Behold - A Devil's Garden

Behold - A Devil's Garden

The ants begin biting the stem of the plant, exposing its vascular tissue. As they bite, the ants also sting the foreign seedling, injecting minute amounts of formic acid into the wound. One or two ants isn't enough to bring down a seedling but one thing ants have on their side are numbers. Soon an entire platoon of ants descend upon the hapless seedling, stinging it repeatedly. In no time at all, the seedling succumbs to the formic acid injections and dies. By repeating this process any time a foreign plant is found growing within the vicinity of the garden, the resident ants ensure that only trees that will produce domatia are allowed to grow in their garden. Thus, a Devil's garden has been formed. 

Although this relationship seems incredibly beneficial for each party, it does come at some cost to the plants themselves. Certainly forming the domatia is a costly endeavor on the part of the plant, but research has also shown that growing in such high, monoculture-like densities in the jungle has its downsides. It has been found that individual host trees can actually experience more herbivore pressures when growing within a Devil's garden than if it was growing alone, elsewhere in the forest. 

Despite their aggression towards herbivores, the ants simply cannot be everywhere at once. As such, the high densities of host tree species within a Devil's garden act like a dinner bell for any insect that enjoys feeding on that particular type of plant. Essentially, the ants are concentrating a potential food source. Experts believe that this might explain why Devil's gardens never completely take over entire swaths of forest. Essentially, there are diminishing returns to living in such high densities. Still, benefits must outweigh costs if such mutualisms are to be maintained and it is quite obvious that both plant and ant benefit from this interaction to a great degree. 

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

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

The First Trees Ripped Themselves Apart To Grow

Pseudosporochnales_reconstruccion.jpg

A new set of fossil discoveries show that the evolutionary arms race that are forests started with plants that literally had to rip themselves apart in their battle for the canopy. The first forests on this planet arose some 385 million years ago and were unlike anything we know today. They consisted of a clade of trees known scientifically as Cladoxylopsids, which have no living representatives in these modern times. How these trees lived and grew has remained a mystery since their fossilized trunks were first discovered but a new set of fossils from China reveals that these trees were unique in more ways than one.

Laying eyes on a full grown Cladoxylopsid would be a strange experience to say the least. Their oddly swollen base would gradually taper up a trunk that stretched some 10 to 12 meters (~30 - 40 feet) into a canopy of its relatives. They had no leaves either. Instead, their photosynthetic organs consisted of branch-like growths that were covered in twig-like projections. Whereas most fossils revealed great detail about their outward appearance, we have largely been in the dark on what their internal anatomy was like. Excitingly, a set of exquisitely preserved fossils from Xinjiang, China has changed that. What they reveal about these early trees is quite remarkable.

As it turns out, the trunks of these early trees were hollow. Unlike the trees we know today, whose xylem expands in concentric rings and forms a solid trunk, the trunk of Cladoxylopsid was made up of strands of xylem connected by a network of softer tissues. Each of these strands was like a mini tree in and of itself. Each strand formed its own concentric rings that gradually increased the size of the trunk. However, this gradual expansion did not appear to be a gentle process.

As these strands increased in size, the trunk would grow larger and larger. In doing so, the tissues connecting the strands were pulled tighter and tighter. Eventually they would tear under the strain. They would gradually repair themselves over time but the effect on the trunk was quite remarkable. In effect, the base of the tree would literally collapse in on itself in a controlled manner. You could say that older Cladoxylopsids developed a bit of a muffin top at their base. 

 A cross section of a Cladoxylopsid trunk showing the hollow center, individual xylem strands, and the network of connective tissues.

A cross section of a Cladoxylopsid trunk showing the hollow center, individual xylem strands, and the network of connective tissues.

Although this seems quite detrimental, the overall structure of the tree would have been quite sturdy. The authors liken this to the design of the Eiffel tower. Indeed, a hollow cylinder is actually stronger than a solid one of the same dimensions. When looked at in the context of all other trees, this form of growth is pretty unique. No other trees are constructed in such a manner.

The authors speculate that this form of growth may be why these trees eventually went extinct. It would have taken a lot of energy to grow in that manner. It is possible that, as more efficient forms of growth were evolving, the Cladoxylopsids may not have been able to compete. It is anyone's guess at this point but this certainly offers a window back into the early days of tree growth. It also shows that there has always been more than one way to grow a tree.

Photo Credits: [1] [2]

Further Reading: [1]

Arums, Orchids and Vines, Oh My!

This week we head into the forests of Illinois to see what late spring botany we can find. This is one of the coolest times of the year to look for plants in temperate North America. 

Producer, Writer, Creator, Host:
Matt Candeias (http://www.indefenseofplants.com)

Producer, Editor, Camera:
Grant Czadzeck (http://www.grantczadzeck.com)

Twitter: @indfnsofplnts

Facebook: http://www.facebook.com/indefenseofplants

Patreon: http://www.patreon.com/indefenseofplants

Tumblr: http://www.tumblr.com/indefenseofplants
_________________________________________________________________

Music by: 
Artist: Lazy Legs
Track: Molasses
Album: Lazy Legs EP
http://lazylegs.bandcamp.com

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]

 

 

The Longleaf Pine: A Champion of the Coastal Plain

As far as habitat types are concerned, the longleaf pine savannas of southeastern North America are some of the most stunning. What's more, they are also a major part of one of the world's great biodiversity hotspots. Sadly, they are disappearing fast. Agriculture and other forms of development are gobbling up the southeast coastal plain at a bewildering rate. For far too long we have ignored, or at the very least, misunderstood these habitats. Today I would like to give a brief introduction to the longleaf pine and the habitat it creates.

The longleaf pine (Pinus palustris) is an impressive species. Capable of reaching heights of 100 feet or more, it towers over a landscape that boggles the mind. It is a landscape born of fire, of which the long leaf pine is supremely adapted to dealing with. These pines start out life quite differently than other pines. Seedlings do not immediately reach for the canopy. Instead, young long leaf pines spend their first few years looking more like a grass than a tree. Lasting anywhere between 5 to 12 years, the grass stage of development gives the young tree a chance to save up energy before it makes any attempt at vertical growth. 

The reason for this is fire. If young long leaf pines were to start their canopy race immediately, they would very likely be burned to death before they grew big enough to escape the harmful effects of fire. Instead, the sensitive growing tip is safely tucked away in the dense needle clusters. If a fire burns through the area only the tips of the needles will be scorched, leaving the rest of the tree safe and sound. During this stage, the tree is busy putting down an impressive root system. The taproot alone can reach depths of 6 to 9 feet!

Once a hardy root system has been formed and enough energy has been acquired, young longleaf pines go through a serious growth spurt. Starting in later winter or early spring, the grass-like tuft will put up a white growth tip called a candle. This tip shoots upwards quite rapidly, growing a few feet in only a couple of months. This is sometimes referred to as the bottlebrush phase because no horizontal branches are formed during this time. The goal at this point is to get the sensitive growing tip as far away from the ground as possible so as to avoid damaging fires. It is fun to encounter long leaf pines at this stage because like any young adult, they look a bit awkward.

 Photo Credit: Woodlot - Wikimedia Commons

Photo Credit: Woodlot - Wikimedia Commons

Once the tree reaches about 6 to 10 feet in height, it will finally begin to produce horizontal branches. This doesn't stop its canopy bid, however, as it still will put on upwards of 3 feet of vertical growth each year! Every year its bark grows thicker and thicker, thus each year it becomes more and more resistant to fire. Far from being a force to cope with, fire unwittingly gives longleaf pines a helping hand by clearing the habitat of potential competitors that are less adapted to dealing with burns. After about 30 years of growth, longleaf pines reach maturity and will start to produce fertile cones.

Before European settlement, longleaf pine savanna covered roughly 90,000,000 acres of southeastern North America. Clearing and development have reduced that to a mere 5% of its former glory. For far too long its coastal plain habitat was thought to be a flat, monotonous region created by early human burning in the last few thousand years. We now know how untrue those assumptions are. Sure, the region is flat but it is anything but monotonous. Additionally, the coastal plain is one of the most lightning prone regions in North America. Fires would have been a regular occurrence long before any humans ever got there. 

Red indicates forest loss between 2011 and 2014. http://glad.umd.edu/gladmaps

Evidence suggests that this coastal plain habitat has remained relatively stable for the last 62,000 years. As such, it is full of unique species. Surveys of the southeastern coastal plain have revealed multiple centers of plant endemism, rivaled in North America only by the southern Appalachian Mountains. In fact, taken together, the coastal plain forests are widely considered one of the world's biodiversity hotspots! Of the 62,000 vascular plants found in these forests, 1,816 species (29.3%) are endemic. Its not just plants either. Roughly 1,400 species of fish, amphibians, reptiles, birds, and mammals rely on the coast plain forests for survival.

Luckily, we are starting to wake up to the fact that we are losing one of the world's great biodiversity hotspots. Efforts are being put forth in order to conserve and restore at least some of what has been lost. Still, the forests of southeastern North America are disappearing at an alarming rate. Despite comprising only 2% of the world's forest cover, the southern forests are being harvested to supply 12% of the world's wood products. This is simply not sustainable. If nothing is done to slow this progress, the world stands to lose yet another biodiversity hotspot. 

If this sounds as bad to you as it does to me then you probably want to do something. Please check out what organizations such as The Longleaf Alliance, Partnership For Southern Forestland Conservation, The Nature Conservancy, and The National Wildlife Federation are doing to protect this amazing region. Simply click the name of the organization to find out more.

Further Reading: [1] [2] [3] [4] 

Shade Gives This Begonia the Iridescent Blues

Believe it or not, the blue iridescence of Begonia pavonina is an evolutionary adaptation to extracting the most amount of energy out of the dim light that makes it through the thick rainforest canopy above. Even more bizarre, it works thanks to an interesting property of quantum mechanics. 

Native to Malaysia, B. pavonina lives out its life in deep shade. Most of the sunlight that hits this region is absorbed by the thick canopy of trees above. As such, eking out an existence is a challenge for these understory herbs. That is where this fantastic blue iridescence comes in. To understand it better, researchers had to take a closer look at its cause. 

Inside any photosynthetic leaves resides the chloroplasts. Chloroplasts are filled with tiny stacks of membranous compartments called "thylakoids." This is where the light reactions of photosynthesis take place. Now, in most plants, these thylakoids are haphazardly distributed throughout the chloroplast. This is not the case for B. pavonina. For this species, the thylakoids are arranged in a very precise way.

It is this precision that gives the leaves their iridescent color. Their placement causes blue wavelengths of light to be reflected. This isn't a big loss for the plant as most of the blue light is absorbed by the canopy above anyway. What it does instead is quite fascinating. The stacked thylakoids act like a dense crystal. When light enters the chloroplasts of B. pavonina it is physically slowed down.

This effect is known to quantum physicists as "slow light." Whereas light traveling through a vacuum maintains a constant speed, light passing through different types of matter can actually be slowed down. By slowing light as it passes through the chloroplasts, the thylakoids are able to take advantage of what little light the leaves are able to intercept. For B. pavonina, this equates to a 10% increase in photosynthetic rates. Coupled with an increase in the absorbance of red-green light, one can understand why this is such an advantage. 

Another interesting aspect of its physiology is the fact that B. pavonina produces both "normal" and iridescent chloroplasts. It is thought that this is a form of backup for the plant. In instances where enough light actually does make it through to the forest floor, B. pavonina can use its normal chloroplasts instead. It should be noted that this is not the only case of blue iridescent leaves in the plant kingdom. Many other species including spike mosses, ferns, and even orchids exhibit this trait. Even leaves that don't appear iridescent to our eyes may be utilizing nanostructures such as those seen in B. pavonina to increase their photosynthetic efficiency in low light conditions. It is very likely that many different kinds of plants are physically manipulating light to their benefit.

Photo Credit: Michael Perry

Further Reading:

[1]

Why Trees?

Walking through the forest is my favorite activity in the world. It is where I feel truly myself. There is something about towering trees that calms me. The thought of why forests are even there often jumps to mind during my strolls. Plenty of plants seem to do just fine hanging out closer to the ground. Why have trees (and some forbs) taken to this vertical realm. Why do forests exist?

In essence, forests are a prime example of an evolutionary arms race. It is one that these organisms have been fighting since the Devonian, roughly 385 million years ago. As plants left the water and began covering the land, some inevitably grew taller than others. There are pros and cons to growing tall. Competition is likely the prime driver for most tree species. Getting above your neighbors means more sunlight. Not every plant is as content as an herb to live out its life in the understory.

Height also means better pollinator visibility and seed dispersal for many tree species. Out in the open, gametes and propagules can be carried great distances by the wind. Colorful blooms would prove to be more exposed and easier for pollinators to locate. Growing tall can also get you out of harms way, removing sensitive growing parts from many different kinds of hungry herbivores and all but the worst forest fires.

There are many downsides to growing tall as well. For one, trees are exposed to the elements and are often victims of strong winds or lightening strikes. What's more, all of that wood takes a lot of energy to produce and, at least for most species, gives nothing back in the way of photosynthesis. It is a rather hefty investment. However, the cost of getting shaded out by your neighbors is definitely not worth the risk of staying small for sun-loving species.

Pumping water is another serious issue. The laws of physics suggest that redwoods are pushing the limits for how tall a tree can grow and still be able to lift water to leaves way up in the canopy. Of course, humidity can assist with such issues but for a majority of the water needs of a tree, water must be able to travel against gravity via weak hydrogen bonds. Water forms an unbroken chain within the vascular tissues of plants. As it evaporates from the leaves, it pulls more water up to fill in the void. It is possible that in today's world, a tree would not physically be able to grow much over 400 feet.

Despite the seemingly lavish waste of limited resources that a forest of trees would suggest, they are nonetheless a common occurrence all over the globe and have been for millions of years. The pros must certainly outweigh the cons or else tallness in trees would never have evolved. The next time you find yourself hiking through a forest, think of how the struggle for survival has led these towering organisms from lowly green stains on rocks to hulking behemoths racing towards the sky.

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