North America's Pachysandra

Photo by Salicyna licensed under CC BY-SA 4.0

In the interest of full disclosure, I have never been a fan of garden variety Pachysandra. Long before I had any interest in plants or gardening, there was something about this groundcover that simply did not appeal to me. Fast forward more than a decade and my views on the use of Asian Pachysandra in the garden have not changed much. You can imagine my surprise then when I learned that North America has its own representative of this genus - the Allegheny spurge (Pachysandra procumbens).

My introduction to P. procumbens happened during a tour of the Highlands Botanical Garden in Highlands, North Carolina. I recognized its shape and my initial reaction was alarm that a garden specializing in native plants would showcase a non-native species. My worry was quickly put to rest as the sign informed me that this lovely groundcover was in fact indigenous to this region. Indeed, P. procumbens can be found growing in shady forest soils from North Carolina down to Florida and Texas.

Photo by David J. Stang licensed under CC BY-SA 4.0

This species is yet another representative of a curious disjunction in major plant lineages between North America and eastern Asia. Whereas North America has this single species of Pachysandra, eastern Asia boasts two, P. axillaris and P. terminalis. Such a large gap in the distribution of this genus (as well as many others) seems a bit strange until one considered the biogeographic history of the two continents.

Many thousands of years ago, sea levels were much lower than they are today. This exposed land bridges between continents which today are hundreds of feet under water. During favorable climatic periods, Asia and North America likely shared a considerable amount of their respective floras, a fact we still find evidence of today. The Pachysandra are but one example of a once connected distribution that has been fragmented by subsequent sea level rise. Fossil records of Pachysandra have been found in regions of British Columbia, Washington, Oregon, Wyoming, and North and South Dakota and provide further confirmation of this.

As a species, P. procumbens is considered a subshrub. It is slow growing but given time, populations can grow to impressive sizes. In spring, numerous fragrant, white flower spikes emerge that are slowly eclipsed by the flush of spring leaf growth. The flowers themselves are intriguing structures worthy of close inspection. Their robust form is what gives this genus its name. "Pachys" is Greek for thick and "andros" is Greek for male, which refers to the thickened filaments that support the anthers.

It is hard to say for sure why this species is not as popular in horticulture as its Asian cousins. It tolerates a wide variety of soil types and does well in shade. What's more, it is mostly ignored by all but the hungriest of deer. And, at the end of the day, it took this species to change my mind about Pachysandra. After all, each and every species has a story to tell.

Photo Credits: [1] [2]

Further Reading: [1] [2]

Tropical Ferns in Temperate North America

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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.

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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.

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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]

Meet the Redbuds

I look forward to the blooming of the redbuds (Cercis spp.) every spring. They paint entire swaths of forest and roadside with a gentle pink haze. It’s this beauty that has led to their popularity as an ornamental tree in many temperate landscapes. Aside from their appeal as a specimen tree, their evolutionary history and ecology is quite fascinating. What follows is a brief introduction to this wonderful genus.

The redbuds belong to the genus Cercis, which resides in the legume family (Fabaceae). In total, there are about 10 species disjunctly distributed between eastern and western North America, southern Europe, and eastern Asia. The present day distribution of this genus is the result of vicariance or the geographic separation of a once continuous distribution. At one point in Earth’s history, the genus Cercis ranged from Eurasia to North America thanks to land bridges that once connected these continents. At some point during the Miocene, this continuous distribution began to break apart. As the climate changed, various Cercis began to diverge from one another, resulting in the range of species we know and love today.

All of them are relatively small trees with beautiful pink flowers. Interestingly enough, unlike the vast majority of leguminous species, redbuds are not known to form root nodules and therefore do not form symbiotic relationships with nitrogen-fixing bacteria called rhizobia. This might have something to do with their preference for rich, forest soils. With plenty of nitrogen available, why waste energy growing nodules? Until more work is done on the subject, its hard to say for sure why they don’t bother with nitrogen fixers.

One of the most interesting aspects of the redbuds are their flowers. We have already established that they are very beautiful but their development makes them even more interesting. You have probably noticed that they are not borne on the tips of branches as is the case in many flowering tree species. Instead, they arise directly from the trunks and branches. This is called "cauliflory," which literally translates to "stem-flower." In older specimens, the trunks and branches become riddled with bumps from years of flower and seed production.

It's difficult to make generalizations about this flowering strategy. What we do know is that it is most common in dense tropical forests. Some have suggests that producing flowers on trunks and stems makes them more available to small insects or other pollinators that are more common in forest understories. Others have suggested that it may have more to do with seed dispersal than pollination. Regardless of any potential fitness advantages cauliflory may incur, the appearance of a redbud covered in clusters of bright pink flowers is truly a sight to behold.

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

Enigmatic Neviusia

Photo by Philip Bouchard licensed under CC BY-NC-ND 2.0

Neviusia. The first time I heard it mentioned I was certain the conversation had switched from reality to the world of Harry Potter. I was wrong. The name belongs to a genus of plants that are totally real. What's more, the natural history of this small group is absolutely fascinating.

The genus Neviusia is comprised of two extant species. N. alabamensis is endemic to a small region of the southeastern United States around northwest Georgia and the Ozark Mountains. Its cousin, N. cliftonii, was discovered in 1992 and is endemic to a small area around "Lake" Shasta in California. Fewer than 20 populations have been found and of them, six were flooded to create "Lake" Shasta. It would seem very strange that both species in this genus are not only endemic to extremely localized regions but also completely disjunct from one another. This is only the beginning.

Whereas fruits have been described for N. cliftonii, none have been reported in N. alabamensis. Ever. Thanks to genetic analysis, populations of both plants are thought to be entirely clonal. High rates of pollen sterility are to blame. Why this is the case is hard to say. It is thought that the genus Neviusia is a relict of the early Cenozoic. Fossil evidence from British Columbia suggest that this genus was once more diverse and more wide spread, having gradually declined to its current limited distribution. The Pleistocene was likely the last straw for these plants, being corralled into small refugia of suitable habitat by the glaciers. Lack of seed production (perhaps due to genetic drift) meant that these two species were to never recolonize their former range. At least not without help...

Since their discovery, these two species have garnered some attention. Like Franklinia, Neviusia have become a sort of horticultural curiosity and have since been out-planted in a variety of locations. My first and only encounter with Neviusia occurred in a conservation garden. Despite their popularity among researchers and gardeners alike, it is unlikely that Neviusia will ever reclaim even a fraction of their former glory. Instead, they remain as endemic reminders of a bygone era. Despite their limited range I think it is important to remember just how long they have survived in North America. After millions of years of survival and persistence, their biggest threat is now us.

Photo Credit: Philip Bouchard (http://bit.ly/1WpElzX)

Further Reading:
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