Dendrologist Squirrels

Gary Cobb licensed under CC BY-ND 2.0

I find it fun to watch squirrels frantically scurrying about during the months of fall. Their usually playful demeanor seems to have been replaced with more serious and directed undertones. If you watch squirrels close enough you may quickly realize that, when it comes to oaks, squirrels seem to have a knack for taxonomy. They quickly bury red oak acorns while immediately set to work on eating white oak acorns. Why is this?

If you have ever tried to eat a raw acorn then you may know the reason. They are packed full of bitter tannins that quickly dry up your mouth and leave an awful after taste. Tannins are secondary chemicals that plants manufacture for protection. Tannins bind to proteins and keep them from being easily digested. This is how leather is made. When you tan a hide you are literally dousing it with tannins that bind to the proteins and keep them from rotting.

Back to the squirrels. The reason they seem to be choosy about how they deal with acorns all comes down to tannins. They bury red oak acorns because acorns in the red oak group have the highest levels of tannins. This is because red oak acorns do not germinate until spring. They have high levels of tannins to fight off fungi and other pathogens over the long, dreary winter. Thus, red oak acorns store better. White oaks germinate in the fall, using a long taproot to pull them into the soil. Because of this, white oaks don't have to dump as much tannin into their acorns. The squirrels seem to know this and simply bite out the white oak embryo before it can germinate. White oak acorns get eaten much sooner than reds because they simply do not keep as long.

There is also evidence that oaks and squirrels have struck a balance. Oaks do rely on squirrels as well as birds like jays to disperse their seeds. These critters can't remember where they cached all of their seeds so some are bound to germinate. What some researchers have found is that oaks place more tannins near the embryos in the acorn than they do at the tips. Why is this? As it turns out, acorns that have had their tips bit off can still germinate as long as their embryo remains unharmed. It is believed that this satisfies squirrels and jays enough to keep them from downing the entire acorn every time. Knowledge such as this puts a whole new spin on backyard ecology.

Photo Credit: Gary Cobb licensed under CC BY-ND 2.0.

Further Reading: [1]

The Overcup Oak

Photo by Bruce Kirchoff licensed under CC BY 2.0

I sure do love me a good oak. Moving to the Midwest of North America has given me the opportunity to meet many new oak species. One oak that has captured my attention in recent years is the overcup oak (Quercus lyrata) whose both common and scientific names first attracted me to this wonderful tree.

Let’s start by looking at the scientific name of this species. The specific epithet “lyrata” was given to this tree because its leaves are said to resemble a lyre. Having no familiarity with popular instruments of Ancient Greece, I had to look this one up. Personally, I have a hard time seeing the resemblance in most leaves. Perhaps this is because the leaves on any given tree can be highly variable in both shape and size depending on both where they are positioned in the canopy and where the tree itself is rooted.

Photo by Bruce Kirchoff licensed under CC BY 2.0

The name “overcup” comes from the fact that the caps of each acorn nearly encompass the entire seed. It is neat to see a mature acorn of this species as they appear to be immature at all stages of development. The odd morphology of these acorns has everything to do with where these trees grow in nature and the way in which they manage seed dispersal.

Photo by Bruce Kirchoff licensed under CC BY 2.0

Overcup oak is one of the most flood tolerant oaks in all of North America. In fact, it most often grows in around wetlands and in floodplains throughout south-central portions of the continent. As such, this species has evolved to tolerate and take advantage of periodic flooding from one year to the next. Not only can mature trees handle weeks of having their roots and trunks completely submerged, the overcup oak also utilizes flooding as a means of seed dispersal.

The cap that covers each seed is very corky, which causes the acorns to float. This is good news for the seeds as young trees have a hard time making a living in the shade of their parents. Historically, floods would pick up and move overcup acorn crops and, with any luck, deposit the acorns in a new floodplain where disturbance has cleared enough spots in the canopy for the acorns to germinate and grow into vigorous young saplings.

USGS/Public Domain

Speaking of germination, overcup oaks are unique among the white oak tribe in that their seeds exhibit a prolonged dormancy. Normally, acorns of the various white oaks germinate in the fall, not long after they were shed from the trees above. However, living in areas prone to flooding would make germinating at that time of year a risky endeavor. As such, overcup oak acorns lay dormant for months until some environmental cue(s) signals enough time has passed.

Overcup oak is also extremely intolerant of fires. Even modest sized burns can severely damage or kill all but the largest individuals. Normally, the forests in which these trees grow are too wet to produce large fires but prolonged droughts and altered flood regimes can change those dynamics to such a degree that large swaths of overcup oak can be killed.

In fact, altered flooding regimes are one of the biggest threats facing overcup oaks in their native range. Because we have dammed, diverted, and channeled so many waterways in North America, the floods that once maintained overcup oak habitats have changed in a big way. Without regular flooding to disperse their seeds and reduce competition from the canopy above, overcup oaks are having a much harder time regenerating. Saplings gradually dwindle in the shade of their parents and, where rivers do continue to flood, these events are often much more severe than they were in the past. Saplings that aren’t tall enough to rise above the floodwaters eventually drown. Overcup oak may be tolerant of flooding but it is by no means its preferred way to live.

Despite these challenges, overcup oak is still a prominent member of seasonally flooded forests throughout its range. It is a magnificent species well worth spending the time to become familiar. It can also make an excellent specimen tree in all but the driest of south-central North American soils. Also, because it is an oak, this incredible species is also chock full of wildlife value, making it an important component of the ecology wherever it is native.

Photo Credits: Bruce Kirchoff [1] [2] (Licensed under CC-BY), U.S. Geological Survey, Chhe, USDA

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

Red or White?

Photo by Msact at English Wikipedia licensed under CC BY-SA 3.0

Who doesn't love a nice oak tree? One cannot overstate their importance both ecologically and culturally. Although picking an oak tree out of a lineup is something many of us are capable of doing, identifying oaks to species can be a bit more challenging. This is further complicated by the fact that oaks often hybridize. Still, it is likely you have come across some useful tips and tricks for narrowing down your oak choices. One such trick is distinguishing between the red oaks and the white oaks. If you're anything like me, this is something you took for granted for a while. Is there anything biologically or ecologically meaningful to such a split?

In short, yes. However, a true appreciation of these groups requires a deeper look. To start with, oaks are members of the genus Quercus, which belongs in the family Fagaceae. Globally there are approximately 400 species of oak and each falls into one of three categories - the red oaks (section Lobatae), the white oaks (section Quercus), and the so-called "intermediate" oaks (section Protoblanus). For the sake of this article, I will only be focusing on the red and white groups as that is where most of the oak species reside. The intermediate oak group is made up of 5 species, all of which are native to the southwestern United States and northwestern Mexico.

As is common with oak identification, reliable techniques for distinguishing between the two groups can be tricky. Probably the most reliable feature is located on the inner surface of the acorn cap. In white oaks, it is hairless or nearly so, whereas in red oaks, it is covered in tiny hairs. Another useful acorn feature is the length of time it takes them to germinate. White oak acorns mature in one season and germinate in the fall. As such, they contain lower levels of tannins. Red oak acorns (as well as those of the intermediate group) generally take at least two seasons to mature and therefore germinate the following spring. Because of this, red oak acorns have a much higher tannin content. For more information on why this is the case, read this article.

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Tyloses in white oak xylem. Photo by Dr. Michael Rosenthal, Technische Universität Dresden via Rosenthal, M.; Bäucker, E. (2013): Zur Anatomie des Holzes der Weiß-Eichen. Rasterelektronenmikroskopische Bildtafeln zu den drei holzanatomischen Schnittrichtungen. licensed under CC BY-SA 3.0

Less apparent than acorns is the difference in the wood of red and white oaks. The wood of white oaks contains tiny structures in their xylem tissues called tyloses. These are absent from the wood of red oaks. The function of tyloses are quite interesting. During extreme drought or in the case of some sort of infection, they cut off regions of the xylem to stop the spread of an embolism or whatever may be infecting the tree. As such, white oaks tend to be more rot and drought resistant. Fun fact, tyloses are the main reason why white oak is used for making wine and bourbon barrels as it keeps them from leaking their contents.

More apparent to the casual observer, however, is leaf shape. In general, the white oaks produce leaves that have rounded lobes, whereas the red oaks generally exhibit pointed lobes with a tiny bristle on their tips. At this point you may be asking where an unlobed species like shingle oak (Quercus imbricaria) fits in. Look at the tip of its leaf and you will see a small bristle, which means its a member of the red oak group. Similarly, the buds of these two groups often differ in their overall shape. White oak buds tend to be smaller and often have blunted tips whereas the buds of red oaks are generally larger and often pointed.

Tricky leaves of the shingle oak (Quercus imbricaria). Note the bristle tip! Photo by Greg Blick licensed under CC BY-NC-ND 2.0

Despite this broad generalizations, exceptions abound. This is further complicated by the fact that many species will readily hybridize. Quercus is, after all, a massive genus. Regardless, oaks are wonderful species chock full of ecological and cultural value. Still, oak appreciation is something we all need more of in our lives. I encourage you to try some oak identification of your own. Get outside and see if you can use any of these tricks to help you identify some of the oaks in your neighborhood.

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

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

Tropical Oaks - Lessons in Biogeography from a Giant Acorn

Seeing the nut of Quercus insignis in person for the first time was a peculiar experience. I didn't know acorns came that big! What was even stranger was encountering this species in the tropics. I thought that in leaving my temperate home behind, I had left trees such as oaks behind as well. Thus, picking up this gigantic acorn was a challenge to my ignorance of tropical forest diversity. What it did for me was ignite a fury of questions regarding the biogeography of the genus Quercus.

Quercus insignis is native from Mexico to Panama. It is a member of the white oak grouping and, despite having one of the largest acorns of any oak species, relatively little is known about this species. What we do know is that it is in trouble. It is considered critically endangered in Mexico and near threatened in Guatemala and Panama with a remaining stronghold in Nicaragua. Habitat loss and changing environmental conditions seem to be at the core of its disappearance.

One big question was looming over me. What was an oak doing this far south? Call it a northern bias but I have always associated oaks with more temperate climes. I needed to get over this. My investigation lead me to some very interesting work done on the family to which oaks belong - Fagaceae. Based on some incredible paleontological and genetic detective work, we now know that Fagaceae originated in Asia. The first fossil evidence of a member of this family dates back some 100 million years, during the early part of the Cretaceous.

At this time, the continents of Asia, Europe, and North America were still connected. Some 60 million years ago, the genus Quercus diverged from Castanea. They were also starting to radiate across the Northern Hemisphere. The first fossil evidence of oaks in North America comes from Paleogene deposits dated to 55 to 50 million years before present. This is when the oaks really started to hit their stride.

Between 22 and 3 million years ago the genus Quercus underwent numerous speciation events. The new terrain of North America must have presented countless opportunities for oaks because they quickly became the most specious genus of all the Fagaceae. This radiation was particularly fruitful in what would become the U.S. and Mexico. Of the roughly 220 species that exist in this region today, 160 occur in Mexico, and of those, 90 species are endemic.

This brings us to the tropics. Evergreen and semi-evergreen oaks have done quite well in this region. However, their astounding diversity quickly drops once you hit the isthmus of Panama. South America is home to only one species of oak. What happened that limited the oaks reign south of the equator?

To put it simply, geology happened. For much of the Earth's history, North and South America shared no connection. Though the exact time frame is debated, tectonic forces joined the two continents some 4.5 million years ago. The Great American Interchange had begun. The two continents were able to freely exchange flora and fauna like never before. The migrations are thought to have been a bit lopsided. Tropical flora and fauna did not do as well farther north but temperate flora and fauna seemed to find warmer climes more favorable. As such, South America gained disproportionately more biodiversity as a result.

This pattern did not hold true for everything though. For the oaks, only one species (Quercus humboldtii) made it through. As such, the genus remains a dominant fixture of the Northern Hemisphere. Sadly, much of this diversity is at serious risk of being lost forever. Like the magnificent Quercus insignis, many of the world's oaks are on the decline. Disease, habitat loss, and countless other issues plague this genus. A 100 million year old journey is quickly being undone in less than two centuries. The hand of man is time and again proving to be a force unrivaled in the biological world.

Leaf Credit: http://www.oaknames.org/

Further Reading:
http://www.sciencedirect.com/science/article/pii/S0378112713006580