Welcome to Appalachia. I have fallen in love with this corner of the world in large part because of its wonderfully rich and unique flora. Join In Defense of Plants as we take a sneak peak at a mere fraction of the botanical riches these mountains hold.

Further Readings On Appalachian Flora:

Producer, Writer, Creator, Host: Matt Candeias

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Twitter: @indfnsofplnts




Tropical Ferns in Temperate North America

All plants undergo some form of alternation of generations. It is the process in which, through reproduction, they cycle between a haploid gametophyte stage and a diploid sporophyte stage. In ferns and lycophytes, this alternation of generations has been taken to the extreme. Instead of the sporophyte relying on the gametophyte for sustenance, the two generations are physically independent and thus separated from one another. In a handful of fern genera here in North America, this has led to some intriguing and, dare I say, downright puzzling distributions.

The presence of a small handful of tropical fern genera in temperate North America has generated multiple scientific investigations since the early 1900's. However, as is constantly happening in science, as soon as we answer one question, seemingly infinite more questions arise. At the very least, the presence of these ferns in temperate regions offers us a tantalizing window into North America’s ancient past.

To say any of these ferns offer the casual observer much to look at would be a bit of an exaggeration. They do not play out their lives in typical fern fashion. These out-of-place tropical ferns exists entirely as asexual colonies of gametophytes, reproducing solely by tiny bundles of cells called gemmae. What's more, you will only find them tucked away in the damp, sheltered nooks and crannies of rocky overhangs and waterfalls. Buffered by unique microclimates, it is very likely that these fern species have existed in these far away corners for a very, very long time. The last time their respective habitats approached anything resembling a tropical climate was over 60 million years ago. Some have suggested that they have been able to hang on in their reduced form for unthinkable lengths of time in these sheltered habitats. Warm, wet air gets funneled into the crevices and canyons where they grow, protecting them from the deep freezes so common in these temperate regions. Others have suggested that their spores blew in from other regions around the world and, through chance, a few landed in the right spots for the persistence of their gametophyte stages.

The type of habitat you can expect to find these gametophytes.

Aside from their mysterious origins, there is also the matter of why we never find a mature sporophyte of any of these ferns. At least 4 species in North America are known to exist this way - Grammitis nimbata, Hymenophyllum tunbridgense, Vittaria appalachiana, and a member of the genus Trichomanes, most of which are restricted to a small region of southern Appalachia. In the early 1980's, an attempt at coaxing sporophyte production from V. appalachiana was made. Researchers at the University of Tennessee brought a few batches of gametophytes into cultivation. In the confines of the lab, under strictly controlled conditions, they were able to convince some of the gametophytes to produce sporophytes. As these tiny sporophytes developed, they were afforded a brief look at what this fern was all about. It confirmed earlier suspicions that it was indeed a member of the genus Vittaria, or as they are commonly known, the shoestring ferns. The closest living relative of this genus can be found growing in Florida, which hints at a more localized source for these odd gametophytes. However, both physiology and subsequent genetic analyses have revealed the Appalachian Vittaria to be a distinct species of its own. Thus, the mystery of its origin remains elusive.

In order to see them for yourself, you have to be willing to cram yourself into some interesting situations. They really put the emphasis on the "micro" part of the microclimate phenomenon. Also, you really have to know what you are looking for. Finding gametophytes is rarely an easy task and when you consider the myriad other bryophytes and ferns they share their sheltered habitats with, picking them out of a lineup gets all the more tricky. Your best bet is to find someone that knows exactly where they are. Once you see them for the first time, locating other populations gets a bit easier. The casual observer may not understand the resulting excitement but once you know what you are looking at, it is kind of hard not to get some goosebumps. These gametophyte colonies are a truly bizarre and wonderful component of North American flora.

Photo Credit: [1] [2]

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

Eastern North America's Temperate Rainforest

I have often remarked that working in the southern Appalachian Mountains during the summer feels more like working in a rainforest than it does an eastern deciduous forest. Lots of rain, high humidity, and a bewildering array of flora and fauna conjure up images of some far away jungle. Only winter can snap this view out of ones head. I recently learned, however, that these feelings are not misplaced. Indeed, this region of southern Appalachia is considered a temperate rainforest. 

These mountains are old. They arose some 480 million years ago and have been shaping life in this region of North America ever since. Another thing these mountains are quite good at is creating their own weather systems. Here in southern Appalachia, warm, wet air from the Gulf of Mexico and western Atlantic blows northward until it hits the Appalachian Mountains. The mountainous terrain comprising parts of Pisgah, Nantahala, and Chattahoochee National Forests has been referred to as "the Blue Wall" and is responsible for the unique conditions that created this temperate rainforest.

As this air rises over their peaks, it begins to cool. As it does, water in the air condenses. This results in torrents of rain. On average, this area receives anywhere from 60 to 100+ inches of rain every year. The Appalachian temperate rainforest is second only to the Pacific Northwest in terms of rainfall in North America. All of this water and heat coupled with the age and relative stability of this ecosystem over time has led to the explosion of biodiversity we know and love today. 

Life abounds in the southern Apps. The plant diversity can be rather intimidating as species from the north mix with those coming up from the south. For instance, there are more tree species in these mountains than in all of Europe.  Rates of endemism in these mountains, both in terms of flora and fauna, are remarkable. There are relics of bygone eras that never expanded their range following repeated glaciations. What's more, a multitude of species combinations can be found as you go from low to high elevations. 

At lower elevation, forests are dominated by American beech (Fagus grandifolia), yellow birch (Betula alleghaniensis), maple (Acer spp.), birch (Betula spp.), and oak (Quercus spp.). Magnolias cover the humid coves. Mid elevations boast birches, mountain ash (Sorbus americana), and mountain maple (Acer spicatum). High elevations contain fraser fir (Abies fraseri) and redspruce (Picea rubens). Both the understory and the the mountain balds are home to a staggering array of different Heaths (Ericaceae). From Rhododendrons to azaleas and mountain laurels, the colors are like those lifted from an abstract painting. The forest floor is where I focus most of my energy. It is hard to capture the diversity of this habitat in only a few paragraphs. What I can say is that I haven't even scratched the surface. It seems like there is something new to see around every corner. 

The point I am trying to make is that this region is quite special. It is something worth protecting. From development to mining and changes in temperature and precipitation, human activities are exacting quite a toll on the Appalachian Mountains. The system is changing and there is no telling what the future is going to look like. Conserving wild places is a must. There is no way around it. Luckily there is a reason people love this place so very much. There are a lot of dedicated folks out there working to protect and conserve everything that makes southern Appalachia what it is. Get out there, enjoy, and support your local land trust!

Further Reading:  [1] 

In Search of Stewartia

Up until a little over a week ago, I had no idea there were native representatives of the family Theaceae other than Franklinia alatamaha in North America. Dr. Mark Whitten was looking for a tree in order to obtain some genetic samples. He showed me a picture and my jaw just about hit the ground.

Fast forward a few days. A friend sent me an email regarding a hike to see Stewartia in the wild. This was an opportunity I was not about to miss. We took the day off and headed into the mountains. We met up with a small group of people whose goal that day was to bask in the glory of the mountain camellia (Stewartia ovata). We were led by local Stewartia expert, Jack Johnston (

It wasn't long before we had our first sighting. Just off the trail leading to a campsite was a spindly looking tree that stood roughly 15 feet in height. Without flowers I don't know if I could pick it out of a lineup. Lucky for us, this small tree was covered in large white blossoms. For the second time that week my jaw had to be pulled up off the ground.

The blossoms were absolutely stunning. About the diameter of a softball and with bright white petals, they are impossible to miss. At the center of each flower is a dense cluster of filaments supporting bright yellow anthers. The filaments themselves are quite attractive. They range in color from pure white to deep purple. What's more, any given tree can sport multiple flowers of with different filament colors.

The color did not seem to influence pollination whatsoever. Each flower we saw was crawling with solitary bees. To be fair though, very little research has been done on this species. Aside from some genetic work, the ecology of the mountain camellia remains a bit of a mystery. What we do know about this tree is that it has its roots in Asia. North America is lucky to have two of the 18 - 20 species of Stewartia. The rest are spread around the Asian continent. North America's Stewartia serve as a reminder of an ancient geologic connection North America and Asia once shared.

By the end of our hike we had lost count of the amount of trees we encountered. Despite their abundance, they are by no means common. Though not technically endangered, their limited distribution and low germination rates make it a sensitive component of the Appalachian flora. With tentative introductions into the horticultural trade, the best way to see this species is in the wild. Look for it growing in cool, shaded edge habitats, most often near mountain streams and rivers. It is a sight you will never forget.


Further Reading:

A Very Strange Sedge

I am quickly realizing that there are some plants out there that I simply cannot prepare myself to see. Something about their look, growth habit, or location just crosses some wires in my brain and causes me a few minutes of confusion until I can regain some composure. Fraser's sedge (Cymophyllus fraserianus) is one such plant.

I had briefly read about this species on a trail map website. The author mentioned there would be some plants worth seeing in the area and Fraser's sedge was one of them. Not being particularly good at gramminoids I figured I probably wouldn't know it even if I had seen it. Was I ever wrong. Fraser's sedge may very well be impossible to miss.

It grows rather large and its long strap-like leaves are more reminiscent of some sort of epiphytic orchid or limp bromeliad. Indeed, Fraser's sedge is truly unique. It is the only member of its genus and experts believe it to be a very old, relictual lineage. It is only found growing on rich mountain slopes in the Appalachian Mountains. It is also quite endangered throughout much of its range due to habitat fragmentation.

Aside from its foliage, Fraser's sedge also produces what are quite possibly the most attractive flowers of any sedge (opinion of course), which are produced in early spring. They are rather unique in that they are stark white. This has led some to believe that this specie is insect pollinated. Whether or not this is a true pollinator syndrome or just a casual observation is yet to be seen. Either way, encountering this plant in flower would be a truly special occurrence.

Due to habitat loss, there is a lot of fear that remaining isolated populations of this wonderful endemic are at increased risk of genetic bottlenecking. DNA analysis of some populations offer hope as the more restricted populations still show signs of ample genetic diversity. Still, time may prove otherwise as more and more individuals are lost to careless development. In the mean time, efforts are being made at conserving this species into the future.

Photo Credit: Will Stuart (

Further Reading:

Southern Tundra

One would hardly consider the southern half of North America to be a tundra-like environment but even so, some tundra plants exist there today...

Up until about 11,000 years ago, much of North America was covered in massive glaciers that were, in some places, upwards of a mile thick. These colossal ice sheets scoured the land over millennia as they advanced and retreated throughout the Pleistocene. Where they covered the land, nothing except some mosses survived. A vast majority of plants were either wiped out or were forced to survive in what are referred to as glacial refugia.

Refugia are ice free areas either within the range of the ice sheets, such as mountain tops, or areas just outside of the ice sheets. Many of North America's plant species took refuge to the south of the glaciers in what is now the Appalachian Mountains. Echos of these plant communities still exist in the southern US today. Some of which are quite isolated from the current distribution of their species. These plant communities are considered disjunct and coming across them is like seeing back in time.

One such plant is the three-toothed cinquefoil (Sibbaldiopsis tridentata). This species is mainly found in northern Canada and Greenland and is considered a tundra species. It needs cold temperatures and is easily out competed in all but the most hostile environments. Why then can you find this lovely cinquefoil growing as far south as Georgia?

The answer are mountains. A combination of high elevation, punishing winds, and lower than average temperatures, means that the peaks of the Appalachian Mountains have more in common with the tundras found much farther north on the continent. As a result of these conditions, plants like S. tridentata have been able to survive into the present while the majority of their tundra associates migrated north with the retreat of the glaciers.

Because of their isolated existence in the Appalachians, S. tridentata is considered endangered in many southern states. Being able to see this plant without having to visit the tundra is quite a unique and humbling experience. It is amazing to consider the series of events that, over thousands of years, have caused this species to end up living on top of these mountains. It is one of those things that one must really stop and mull over for a bit in order to fully appreciate.

Further Reading:…/j.1365-2699.1998.…/abstract

The Bells of Oconee

The whole point of In Defense of Plants is to remove the human element and tell the story of plants for what they are. I find their stories to be far more interesting than any anthropocentric use they might have. However, the following tale was just far too compelling to ignore. It is a story of passion and, in the end, really encompasses the reality of the species it centers around.

Asa Gray was an eminent 19th century botanist. In 1838, Gray left America for Europe in order to examine herbarium specimens which would reveal the original sources of American flora. While in Paris, Gray was pouring over collections made by Andre Michaux when he came across a poorly preserved specimen of an unnamed plant "with a habit of Pyrola and the foliage of Galax" originating from the "High Mountains of Carolina."

For whatever reason, Gray became enamored with this small pressed plant. He knew it had to be a new species. Upon returning to America, Gray went about organizing expeditions to rediscover this odd little botanical wonder. Sadly it would be another 40 years before he would see a living specimen. 

The species in question is Shortia galacifolia, better known as Oconee bells. A member of the family Diapensiaceae, Shortia is often described as a small, spreading, evergreen sub-shrub. In early spring, each plant produces a beautiful whiteish-pink, bell shaped flower. Today, Shortia is only known from a small handful of populations growing along a couple stream banks in the Southern Appalachians. The original population that Michaux collected from now lies under 980 feet of water, lost forever by the damming of the Keowee River. 

There has been a lot of speculation over why this plant is endemic. A lot of it has to do with Shortia's germination requirements. It is a plant of disturbance, relying on things like blowdowns or minor landslides to open the canopy just enough to create the perfect microclimate. As canopies close, populations languish and disappear. Fortunately for Shortia, collections have been out-planted at a handful of botanical gardens throughout the region where they grow and persist in great numbers. 

Photo Credit: Wikimedia Commons

Further Reading:

My oh my, What a Beautiful Magnolia

Magnolia fraseri is, in my opinion, one of the most beautiful trees in our eastern forests. To see this species, one must travel to the Southern Appalachian Mountains where it is endemic. With its whorls of massive leaves, large, cream colored flowers, and smooth gray bark, it is an unmistakeable component of the Appalachian cove forests.

M. fraseri needs canopy gaps to persist. Anywhere that disturbance opens up the canopy and allows light in, M. fraseri is soon to follow. This tree has surely benefitted from the mass die off of eastern hemlock due to the invasive hemlock wooly adelgid. This species flowers in the spring. Magnolias as a whole are an ancient lineage of flowering plants, arising before bees evolved. For that reason, their flowers are pollinated by beetles instead of bees. The large, showy flowers soon give way to your typical magnolia seed pod. As the seeds mature, they are pushed out of their pod and their bright red coloration helps to attract their main seed disperser, birds.

Aside from seed production, the most common form of reproduction for M. fraseri is via stump sprouts. In fact, it is believed that many of the oldest M. fraseri in the Appalachian forest region are stump sprouts that harken back to a time in which forest clearing was more rampant.

The overall appearance of this tree feels tropical. The large leaves are are arranged like an umbrella and these whorls stack themselves all the way up the trunk. Why this species is not cultivated as a native landscape tree is beyond me and I think the following excerpt by Richard E. Weaver Jr. sums it up quite nicely:



"Many of our fine native plants remain rare in cultivation in our own
country for a variety of reasons. Over-familiarity with them as wild
plants; lack of commercial availability; ignorance as to culture and
propagation; or plain snobbishness. Many are far better appreciated abroad."

Magnolia fraseri was one of the first plants that greeted me upon entering North Carolina. It was growing alongside a pawpaw at a scenic overlook that showcased the hardwood forests that coat these mountains. I never pass up an opportunity to appreciate this tree and indeed I will carry the image of it in my mind wherever I go.

Photo Credit: Jim Dollar (

Further Reading:…/1981-41-2-magnolia-……/si…/volume_2/magnolia/fraseri.html


One of the many aspects I love about being in the mountains is that they seem to defy time. Not in any science fiction sort of way, but more in terms of seasonality. What I mean by this is that if a plant is done flowering at the base of the mountain, there is a good chance that it is still flowering closer to the top. This ability to rewind flowering seasons has been beautifully illustrated this month by beetleweed (Galax urceolata).

This lovely plant is a member of the family Diapensiaceae. It is the only species in that genus. When I first arrived in the Southern Appalachians a couple of weeks ago, most of the plants at low altitudes were nearly done flowering. I was a bit disappointed as I had never seen this species in person before. It didn't take long before the situation was remedied. 

The first trip up a mountainside revealed that plants midway up where just reaching full bloom and plants near the top were just beginning. Going up in elevation is a fun proxy for going up in latitude. Changes in microclimates mean plants are experiencing different cycles every few hundred feet. As such, this whole month I have been able to enjoy sequential blooming of a wide variety of plants simply by hiking up.

Galax urceolata is a beautiful plant. Get up close and the beauty is replaced by a rather mousy odor. There isn't much information on what is pollinating this species but my bet is on either flies or beetles. Recently I did observe a bumblebee briefly visiting an inflorescence but whether or not this was a one off remains to be seen. It is an evergreen plant, keeping its leaves all winter. The leaves turn from green to red as they fill with anthocyanin pigments. There has been a lot of discussion over the role these pigments play in the survival of this species. Some feel it is a way of protecting against harsh light. Evidence is showing the issue to be more complex than that. Though they probably serve many functions, the main purpose of these pigments may actually be to protect the plants cellular machinery from dangerous oxygen free radicals. 

Another interesting thing about this species is that both diploid and tetraploid populations exist and there is evidence that they segregate themselves by habitat. This very well could lead to speciation, rising the species count in this genus to two. For now, treating them as a single species is fine by me. It looks like I will get to enjoy these beautiful albeit stinky blooms for the rest of my stay in the mountains. 

Further Reading:

Under the Umbrellaleaf


The first time I ever laid eyes on this species, I was certain there must be a mistake. What was a large Begonia doing growing on a wet seep deep in the woods somewhere in the Southern Appalachians. Surely something very strange was going on in this spot. After a few minutes of observation though, I realized my initial assumption wasn't correct. This wasn't an escaped species of Begonia. Instead, what my field guide revealed was that this gorgeous plant is actually a wonderful southeastern native called Diphylleia cymosa.


Known commonly as "umbrellaleaf," D. cymosa is a member of the family Berberidaceae and is a distant cousin of the more widely spread mayapple. North America has only one species of Diphylleia as does Japan (D. grayi) and China (D. sinensis). Phylogenetic evidence hints at a time in which North America and Asia were connected and shared much of their respective flora. Tectonic movements have since isolated these once connected populations, allowing ample time for the speciation that led to the species we know and love today. North America's species (D. cymosa) is found only in wet areas of the Appalachian Mountains.


As the generic name "Diphylleia" suggests, each individual plant produces two large, umbrella-like leaves. Arising from the middle of these leaves is a cluster or "cyme" of beautiful white flowers. After pollination, the cyme gives way to a cluster of berries, which gradually turn a deep shade of blue. The pedicels themselves turn a deep shade of red. All of this creates a beautiful fruit display aimed at attracting woodland birds, D. cymosa's main seed dispersers. 

Photo Credit: Owen and Aki ( and Emma Harrower (

Further Reading:

"The Mountains Are Calling and I Must Go..." - John Muir

I find myself thinking the same thing as I drive to work every morning - "Screw you, North Carolina! How dare you be so beautiful?" I say this with love of course. I mean it too. My daily commute takes me through the Cowee Mountains, which represent only the tiniest fraction of the giant fold in the continent that we collectively refer to as the "Appalachian Mountains." Driving between these forested peaks, it feels as if time stands still. They are a stark and constant reminder of just how small and insignificant our time on this planet really is. 

What so few realize is that these mountains are some of the oldest on Earth. They are the collective result of some serious geology. Between 325 million and 260 million years ago, Africa slammed into North America (though it wasn't the continent we know today) causing massive upheaval of the crust. This was also the birth of the super-continent Pangea. The resulting upheaval produced a mountain chain similar in size to the present day Himalayas (think Mt. Everest). They have been steadily eroding ever since. 

Today, the highest peaks reach somewhere in the 6,000 feet (1,800 m) range. Despite this reduction in size, the Appalachian Mountains have nonetheless been a major driver in the ecology of eastern North America. They have served as refugia for species escaping glaciation, they act as corridors for migration, and they even produce their own climate. They may not be the snow capped mountains of the American West but they have a uniqueness all their own. There aren't many places in this world in which one can explore broadleaf deciduous forests at elevation. If I could end up here I think I would die a happy man. For now, I am spending every free moment absorbing the beauty and splendor of this place. It is going to be hard to leave...

Heading South

I never much considered southern North America as a place I would enjoy. Born and raised in the north, I always assumed that anywhere below Pennsylvania would be too hot and sticky for me. A brief trip to Great Smoky Mountains National Park changed all of that. I fell head over heals for the forests of this region. Species I had only read about or perhaps stumbled across once or twice up home were now surrounding me, bursting forth with a profusion unlike anything I had ever seen before. There is something special about this region. The trip ended seemingly as soon as it began but I was hooked. 

Fast forward a few more years. When I found out that I would be pursuing a  PhD in Illinois, I was both excited and apprehensive. A career goal I set for myself back in the early days of high school was finally set in motion. This was reason to celebrate. Yet, leaving behind any form of topography for the ironed out landscape of the American midwest seemed a bit nerve racking. However, my nerves were quickly assuaged after finding out that my research was going to be based in the southern Appalachian Mountains. It was back to the forested peaks for me!

On June 2nd, 2015, I said goodbye to Buffalo and headed south. The purpose of this trip was to get a feel for where I would be working and hopefully inspire me into generating some hypotheses. Needless to say I was ecstatic about spending a month in the mountains. At the point of writing this, I am now 5 days into this journey and it is safe to say that I am completely transfixed with these mountains.

On the surface, this is not hard to imagine. A combination of topography, climate, and lots of history support some of the most diverse plant communities on the entire continent. This place is bursting at the seams with life. Every morning I awake to a cacophony of birdsong. I listen intently to songs whose  identity escapes me. As I sit out on the deck, sucking down the coffee that will fuel me during the morning plant surveys, I look out into a rich Appalachian cove forest. Up on the deck I stand about midway in the canopy staring down ancient oaks, magnolias, maples, and even the occasional holly. The birds are busy harvesting this year's crop of caterpillars to feed their rapidly growing chicks. This world is alive. 

To earn my keep, I am assisting in some plant surveys. My mentor and friend Dr. Robert Warren has set up experimental plots along north and south facing slopes and my friend Lauren and I are tasked with identifying everything growing within them. I couldn't ask for a better gig. It is going to be a great month. In my down time I plant to explore as much of this place as possible. Not a minute will go to waste. The icing on the cake is that I am surrounded by like minded ecologists in training. Everyone down here has their own speciality, their own questions, and their own passions that drive them to do what they do. Nowhere have I felt more at home. 


An Endemic Houstonia


The weathered peaks of the Southern Appalachians are home to a bewildering variety of plant life. This region is thought to have provided refuge for many different types of flora and fauna pushed south by repeated glaciation. High humidity and precipitation coupled with a variety of microclimates has allowed plants to flourish and evolve over the millennia. In fact, a handful of species are found nowhere else in the world. One of these montane endemics is none other than a species of Houstonia

Some feel it best designated as a subspecies, Houstonia purpurea var. montana, whereas others feel that both morphological and reproductive distinctions deserve it a status as its own species, Houstonia montana. I prefer to refer to it as the Roan Mountain bluet. Either way, this unique little plant can be found  growing among rocky summits and balds on only a handful of mountain tops between Tennessee and North Carolina.  

This species requires disturbance to survive. Without the constantly shifting landscape characteristic of high altitude regions, this little plant would quickly be overtopped and outcompeted by more aggressive vegetation. This is not a lifestyle unique to this little bluet. Many of the worlds rare plant species require some level of disturbance to release them from competition with other more common plant species. Aside from competition, one of the largest threats to the continued survival is trampling by hikers. It is always important to watch where we hike. A little bit of attention can go a long way for our botanical neighbors. 

Photo Credit: BlueRidgeKitties (

Further Reading: