Big Things Come In Small Packages

Meet Blossfeldia liliputana, the smallest species of cactus in the world. With a maximum diameter of only 12 mm, this wonderful succulent would be hard to spot tucked in among the nooks and crannies of rock outcrops. Its species name "liliputana" is a reference to the fictional island of Liliput (Gulliver's Travels) whose inhabitants were said to be rather small. If its size alone wasn't interesting in and of itself, the biology of B. liliputana is also downright bizarre.

Blossfeldia liliputana is native to arid regions between southern Bolivia and northern Argentina. It appears to prefer growing wedged between cracks in rock as these are usually the spots where just enough soil builds up to put down its roots. Root formation, however, does not happen for quite some time. Most often new individuals bud off from the parent plant. They emerge not from the base, but rather from apical tissues, yet another unique feature of this cactus. What's more, this cactus produces no spines. Instead, its numerous areoles are covered in a dense layer of trichomes that are felt-like to the touch.

As you can clearly see, this species is small. It only ever becomes conspicuous when it comes time to flower. Imagine a bunch of tiny white to pink cactus flowers poking out of a crevice. It must be a remarkable sight to see in person. Despite their showy appearance, its is believed that most are self-fertilized.

Photo by Mats Winberg licensed under CC BY-SA 2.5

As mentioned, the size of this cactus isn't the only interesting thing about its biology. B. liliputana is categorized as a poikilohydric organism, meaning it doesn't have the ability to regulate its internal water content. Researchers have found that individual plants can lose up to 80% of their weight in water and can maintain that state for as long as two years without any negative effects. As such, colonies of these tiny cacti often appear shrunken or squished. Once the rains arrive, however, it springs back to its original rounded shape with seemingly no issues. Amazingly, a significant amount of water uptake happens via the fuzzy areoles that cover its surface, hence it does not harm the plant to hold off growing roots for quite some time. 

Photo by AstroKaktus at Slovenian Wikipedia licensed under the GNU Free Documentation License

Speaking of water regulation, B. liliputana holds another record for having the lowest density of stomata of any terrestrial autotrophic vascular plant. Stomata are the pores in which plants regulate water and gas exchange so having so few may have something to do with why this species loses and gains water to such a degree that would kill most other vascular plant species.

Another peculiar quality of this cactus are its seeds. Unlike all other cacti whose seeds are hard and relatively smooth, the seeds of B. liliputana are hairy. Attached to each seed is a small fleshy structure called an aril, which aids in seed dispersal. As it turns out, B. liliputana relies on ants as its main seed dispersers. Ants attracted to the fleshy aril drag the seeds back to their nests, remove and eat the aril, and then discard the seed. This is often good news for the cactus because its seeds end up in nutrient-rich ant middens protected from both the elements and seed predators, often in much more suitable conditions for germination.

Photo by Michael Wolf licensed under CC BY-SA 3.0

Needless to say, B. liliputana is a bit of an oddball as far as cacti are concerned. Its highly derived features coupled with its bizarre biology have made it difficult for taxonomists to elucidate its relationship to the rest of the cactus family. It certainly deserves its own genus, to which it is the only member, however, it has been added to and removed form a handful of cactus subfamilies over the years. The most recent genetic analyses suggests that it is unique enough to warrant its own tribe within Cactaceae - Blossfeldieae.

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

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

How Aroids Turn Up the Heat

Photo by Jörg Hempel licensed under CC BY-SA 2.0

A subset of plants have evolved the ability to produce heat, a fact that may come as a surprise to many reading this. The undisputed champions of botanical thermogenesis are the aroids (Araceae). Exactly why they do so is still the subject of scientific debate but the means by which heat is produced is absolutely fascinating.

The heat producing organ of an aroid is called the spadix. Technically speaking, a spadix is a spike of minute flowers closely arranged around a fleshy axis. All aroid inflorescences have one and they come in a wide variety of shapes, colors, and textures. To produce heat, the spadix is hooked up to a massive underground energy reserve largely in the form of carbohydrates or sugars. The process of turning these sugars into heat is rather complex and surprisingly animal-like.

Cross section of a typical aroid inflorescence with half of the protective spathe removed. The spadix is situated in the middle with a rings of protective hairs (top), male flowers (middle), and female flowers (bottom). Photo by Kristian Peters -- Fabelfroh licensed under CC BY-SA 3.0

It all starts with a compound we are rather familiar with - salicylic acid - as it is the main ingredient in Aspirin. In aroids, however, salicylic acid acts as a hormone whose job it is to initiate both the heating process as well as the production of floral scents. It signals the mitochondria packed inside a ring of sterile flowers located at the base of the spadix to change their metabolic pathway.

In lieu of their normal metabolic pathway, which ends in the production of ATP, the mitochondria switch over to a pathway called the "Alternative Oxidase Metabolic Pathway." When this happens, the mitochondria start burning sugars using oxygen as a fuel source. This form of respiration produces heat.

Thermal imaging of the inflorescence of Arum maculatum.

As you can imagine, this can be a costly process for plants to undergo. A lot of energy is consumed as the inflorescence heats up. Nonetheless, some aroids can maintain this costly level of respiration intermittently for weeks on end. Take the charismatic skunk cabbage (Symplocarpus foetidus) for example. Its spadix can reach temperatures of upwards of 45 °F (7 °C) on and and off for as long as two weeks. Even more incredible, the plant is able to do this despite freezing ambient temperatures, literally melting its way through layers of snow.

For some aroids, however, carbohydrates just don't cut it. Species like the Brazilian Philodendron bipinnatifidum produce a staggering amount of floral heat and to do so requires a different fuel source - fat. Fats are not a common component of plant metabolisms. Plants simply have less energy requirements than most animals. Still, this wonderful aroid has converged on a fat-burning metabolic pathway that puts many animals to shame. 

The inflorescence of Philodendron bipinnatifidum can reach temps as high as 115 °F (46 °C). Photo by Tekwani licensed under CC BY-SA 3.0

P. bipinnatifidum stores lots of fat in sterile male flowers that are situated between the fertile male and female flowers near the base of the spadix. As soon as the protective spathe opens, the spadix bursts into metabolic action. As the sun starts to set and P. bipinnatifidum's scarab beetle pollinators begin to wake up, heat production starts to hit a crescendo. For about 20 to 40 minutes, the inflorescence of P. bipinnatifidum reaches temperatures as high as 95 °F (35 °C) with one record breaker maxing out at 115 °F (46 °C)! Amazingly, this process is repeated again the following night.

It goes without saying that burning fat at a rate fast enough to reach such temperatures requires a lot of oxygen. Amazingly, for the two nights it is in bloom, the P. bipinnatifidum inflorescence consumes oxygen at a rate comparable to that of a flying hummingbird, which are some of the most metabolically active animals on Earth.

The world's largest inflorescence belongs to the titan arum (Amorphophallus titanum) and it too produces heat. Photo by Fbianh licensed under CC0 1.0

Again, why these plants go through the effort of heating their reproductive structures is still a bit of a mystery. For most, heat likely plays a role in helping to volatilize floral scents. Anyone that has spent time around blooming aroids knows that this plant family produces a wide range of odors from sweet and spicy to downright offensive. By warming these compounds, the plant may be helping to lure in pollinators from a greater distance away. It is also thought that the heat may be an attractant in and of itself. This is especially true for temperate species like the aforementioned skunk cabbage, which frequently bloom during colder months of the year. Likely both play a role to one degree or another throughout the aroid family.

What we can say is that the process of plant thermogenesis is absolutely fascinating and well worth deeper investigation. We still have much to learn about this charismatic group of plants.

LEARN MORE ABOUT AROID POLLINATION HERE



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

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

The Rose of Jericho

To survive in a desert, plants must eek out an existence in specific microclimates that provide conditions that are only slightly better than the surrounding landscape. Such is the case for the Rose of Jericho (Anastatica hierochuntica). This tenacious little mustard is found throughout arid regions of the Middle East and the Saharan Desert and it has been made famous the world over for its "resurrection" abilities. It is also the subject of much speculation so today we are going to separate fact from fiction and reveal what years of research has taught about this desert survivor. 

Natural selection has shaped this species into an organism fully ready to take advantage of those fleeting moments when favorable growing conditions present themselves. A. hierochuntica makes its living in dry channels called runnels or wadis, which concentrate water during periods of rain. It is a desert annual meaning the growth period of any individual is relatively short. Once all the water in the sandy soil has evaporated, this plant shrivels up and dies. This is not the end of its story though. With a little luck, the plants were pollinated and multiple spoon-shaped fruits have formed on its stems.

Photo by Phil41 licensed under CC BY 1.0

As the dead husk of the plant starts to dry out, its branches curl up into a ball-like mass with most of the fruits tucked away in the interior. There the plant will sit, often for many years, until rain returns. When rain does finally arrive, things happen fast. After all, who knows how long it will be before it rains again. Thanks to a quirk of physiology, the dried tissues of A. hierochuntica are extremely elastic and can return to their normal shape and position once hydrated. As the soil soaks up water, the dried up stems and roots just under the surface also begin taking up water and the stems unfurl.

To call this resurrection is being a bit too generous. The plant is not returning to life. Instead, its dead tissues simply expand as they imbibe liquid. Water usually does not come to the desert without rain and rain is exactly what A. hierochuntica needs to complete its life cycle. Unfurling of its stems exposes its spoon-shaped fruits to the elements. Their convex shape is actually an adaptation for seed dispersal by rain, a mechanism termed ombrohydrochory. When a raindrop hits the fruit, it catapults the seed outward from the dead parent.

Photo by Roland Unger licensed under CC BY-SA 3.0

If rains are light, seeds do not get very far. They tend to cluster around the immediate area of their parent. If rains are heavy, however, seeds can travel quite a distance. This is why one will only ever find this species growing in channels. During the rare occasions when those channels fill with water, seeds quickly float away on the current. In fact, experts believe that the buoyancy of A. hierochuntica seed is an adaptation that evolved in response to flooding events. It is quite ironic that water dispersal is such an important factor for a plant growing in some of the driest habitats on Earth.

To aid in germination, the seeds themselves are coated in a material that becomes mucilaginous upon wetting. When the seeds eventually come into contact with the soil, the mucilage sticks to the ground and causes the seeds to adhere to the surface upon drying. This way, they are able to effectively germinate instead of blowing around in the wind.

Again, things happen fast for A. hierochuntica. Most of its seeds will germinate within 12 hours of rainfall. Though they are relatively drought tolerant, the resulting seedlings nonetheless cannot survive without water. As such, their quick germination allows them to make the most out of fleeting wet conditions.

Photo by Nikswieweg at German Wikipedia licensed under CC BY-SA 2.0 DE

Occasionally, the balled up husks of these plants will become dislodged from the sand and begin to blow around the landscape like little tumbleweeds. This has led some to suggest that A. hierochuntica utilizes this as a form a seed dispersal, scattering seeds about the landscape as it bounces around in the wind. Though this seems like an appealing hypothesis, experts believe that this is not the best means of disseminating propagules. Seeds dispersed in this way are much less likely to end up in favorable spots for germination. Though it certainly occurs, it is likely that this is just something that happens from time to time rather than something the plant has evolved to do.

In total, the Rose of Jericho is one tough cookie. Thanks to quick germination and growth, it is able to take advantage of those rare times when its desert environment become hospitable.

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

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

Celebrating the Forked Spleenwort

Photo by Bernd Haynold licensed under CC BY-SA 3.0

What can I say, I am a total sucker for ferns with "untraditional" fronds. Whereas the tropics offer seemingly endless fern varieties, I find that there is something special about temperate ferns that, for lack of a better phrase, break the mold. I was recently introduced to such a fern. Known commonly as the forked spleenwort, Asplenium septentrionale looks more like a clump of grass than it does a fern.

A closer inspection, however, would reveal that it is indeed a Pteridophyte. It grows on rocky outcrops, including stone walls, throughout the northern hemisphere. Here in North America, it is predominantly found in the Rocky Mountains. It is a small fern that often forms dense clusters in cracks and crevices. Its stems are long, narrow, and grass-like, ending in a flattened leaf blade that often forks at the tip. In typical fractal fashion, these leaf blades fork again at the tips, forming minute pinnae.

Photo by Rolf Engstrand licensed under CC BY-SA 3.0

The forked spleenwort has gone through considerable taxonomic revisions since it was first described by Linnaeus in 1753. Originally it was named Acrostichum septentrionale, but was then moved into Asplenium a few decades later. Renewed interest in this species during the mid 20th century saw the forked spleenwort moved to the genus Chamaefilix followed by Tarachia, though these did not gain much scientific credence. As such, it has remained an Asplenium ever since.

Its taxonomic story does not end there, however, as genetic tests soon revealed that a much more subtle and nuanced revision was worth considering. It was discovered that the forked spleenwort existed in two genetically distinct types, a diploid (having two sets of chromosomes) and a tetraploid (having four sets of chromosomes). Researchers found that each group had its own distinct distribution with the diploids centered in southwest Asia and the tetraploids being circumboreal.

Photo by Bernd Haynold licensed under CC BY-SA 1.0

It was clear that a subspecies division was worth considering. Further investigations in the early 2000's revealed the presence of sterile triploid individuals that are believed to be hybrids of the two mentioned above. What's more, the forked spleenwort has been found to hybridize with other members of its genus. It is believed that the more isolated populations owe their existence in part to the isolation of their preferred substrate - these ferns do best on acidic substrates where competition is low - and decent longevity. It has been speculated that genetic differences can be maintained when "mutant" individuals become established and persist undisturbed for long periods of time.  

Regardless of its taxonomic status, the forked spleendwort is nonetheless a charismatic little species. A simple image search will reveal just how pleasant this species is in situ. Even better, its beauty and splendor can be shared by botanical enthusiasts throughout the northern hemisphere. There is something to be said about such species.

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

Further Reading: [1]

The Curious Case of the Yellowwood Tree

Photo by Plant Image Library licensed under CC BY-SA 2.0

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.

Though the nature of its oddball distribution pattern iscurious to say the least, 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 perform great 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 is 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.

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 sight 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.

Photo by Elektryczne jabłko licensed under CC BY-SA 4.0

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. In some instances, even pink flowers are produced! 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]

Further Reading: [1] [2]

Trees In Spring

Spring is a wonderful time to observe trees. After a long, dreary winter they burst into action. For many species, spring is the time for reproduction.

Species in this episode:

-Serviceberry (Amelanchier sp.)

-Norway maple (Acer platanoides)

-Eastern redcedar (Juniperus virginiana)

-Sugar maple (Acer saccharum)

-Saucer magnolia (Magnolia x soulangeana)

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

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

Pollen Competition

Photo by Martin LaBar licensed under CC BY-NC 2.0

The animal kingdom is rife with sexual conflict. We are all aware of what is going on when two stag deer lock antlers or when a group of male sage grouse flaunt themselves on leks as females look on. But what about plants? Is there sexual conflict among plant species? Whether pollen ends up on a stigma via wind or animal, is there any way for a plant to "choose" who gets to fertilize the ovule?

It turns out, yes, there is. Sexual competition is part of the pollination process. In fact, some of the most familiar floral morphologies may have evolved as a way of weeding out weak paternal lines. To understand this process better, though, we must first quickly review exactly what goes on during pollination.

Photo by Nick Fedele licensed under CC BY-NC-SA 2.0

Pollen is a male gamete. Each grain is haploid and contains only a single copy of a plant’s chromosomes. When a pollen grain lands on a stigma, the grain germinates like a tiny seed, sending down a root-like growth called a pollen tube. This tube grows down into the ovary until it finds an unfertilized ovule. At this point, sperm travels down the pollen tube where it can unite with the ovule, thus forming a seed.

By CNX OpenStax licensed under CC BY 4.0

It’s the formation of this pollen tube that introduces the idea of competition among pollen grains. Again, whether by wind or animal, the pollen arriving to a new plant generally doesn't come from a single individual. Pollen from many potential paternal lines can arrive all at once. As such, the race to fertilize the ovules can be quite intense, and this is where competition begins.

Remember, pollen only contains a single set of chromosomes from the parent plant, thus all alleles, both functioning and deleterious, are represented. During the growth of the pollen tube, upwards of 60% of the pollen genome is actively transcribed. Any pollen containing lots of deleterious alleles is going to have a much harder time competing with pollen grains that have fewer deleterious alleles. Their tubes have a harder time making it to the ovules in time to fertilize them.

Photo by Dartmouth Electron Microscope Facility, Dartmouth College

It is thought that the length of the style (the stem connecting the stigma to the ovaries) may also provide a sort of "proving ground" for pollen too. For instance, picture the flowers of a lily or a mallow. Those long, slender styles may actually be acting like a race track. Only the pollen with the best selection of genetic material will be able to grow their pollen tubes fast enough to reach the ovules, leaving the weaker competition in the dust. In this way, plants may actually be sorting out stronger paternal lines, which makes sense for sessile organisms that can't see.

As with everything in nature, there is far more nuance to this than what I have outlined above. Much work is being done to test some of the earlier assumptions and data surrounding this concept of pollen competition. It certainly happens but the degree to which any given species utilizes such methods is up for debate. Still, it paints a much more interesting picture of mate selection in plants. Static, plants are not!

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

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

Saving One of North America's Rarest Shrubs

Photo by Stan Shebs licensed under CC BY-SA 3.0

The chance to save a species from certain extinction cannot be wasted. When the opportunity presents itself, I believe it is our duty to do so. Back in 2010, such an opportunity presented itself to the state of California and what follows is a heroic demonstration of the lengths dedicated individuals will go to protect biodiversity. Thought to be extinct for 60 years, the Franciscan manzanita (Arctostaphylos franciscana) has been given a second chance at life on this planet.

California is known the world over for its staggering biodiversity. Thanks to a multitude of factors that include wide variations in soil and climate types, California boasts an amazing variety of plant life. Some of the most Californian of these plants belong to a group of shrubs and trees collectively referred to as 'manzanitas.' These plants are members of the genus Arctostaphylos, which hails from the family Ericaceae, and sport wonderful red bark, small green leaves, and lovely bell-shaped flowers. Of the approximately 105 species, subspecies, and varieties of manzanita known to science, 95 of them can be found growing in California.

It has been suggested that manzanitas as a whole are a relatively recent taxon, having arisen sometime during the Middle Miocene. This fact complicates their taxonomy a bit because such a rapid radiation has led manzanita authorities to recognize a multitude of subspecies and varieties. In California, there are also many endemic species that owe their existence in part to the state's complicated geologic history. Some of these manzanitas are exceedingly rare, having only been found growing in one or a few locations. Sadly, untold species were probably lost as California was settled and human development cleared the land. 

Such was the case for the Franciscan manzanita. Its discovery dates back to the late 1800's. California botanist and manzanita expert, Alice Eastwood, originally collected this plant on serpentine soils around the San Francisco Bay Area. In the years following, the growing human population began putting lots of pressure on the surrounding landscape.

Photo by Daderot (public domain)

Botanists like Eastwood recognized this and went to work doing what they could to save specimens from the onslaught of bulldozers. Luckily, the Franciscan manzanita was one such species. A few individuals were dug up, rooted, and their progeny were distributed to various botanical gardens. By the 1940's, the last known wild population of Franciscan manzanita were torn up and replaced by the unending tide of human expansion into the Bay Area.

It was apparent that the Franciscan manzanita was gone for good. Nothing was left of its original populations outside of botanical gardens. It was officially declared extinct in the wild. Decades went by without much thought for this plant outside of a few botanical circles. All of that changed in 2009.

It was in 2009 when a project began to replace a stretch of roadway called Doyle Drive. It was a massive project and a lot of effort was invested to remove the resident vegetation from the site before work could start in earnest. Native vegetation was salvaged to be used in restoration projects but most of the clearing involved the removal of aggressive roadside trees. A chipper was brought in to turn the trees into wood chips. Thanks to a bit of serendipity, a single area of vegetation bounded on all sides by busy highway was spared from wood chip piles. Apparently the only reason for this was because a patrol car had been parked there during the chipping operation.

Cleared of tall, weedy trees, this small island of vegetation had become visible by road for the first time in decades. That fall, a botanist by the name of Daniel Gluesenkamp was driving by the construction site when he noticed an odd looking shrub growing there. Luckily, he knew enough about manzanitas to know something was different about this shrub. Returning to the site with fellow botanists, Gluesenkamp and others confirmed that this odd shrubby manzanita was in fact the sole surviving wild Franciscan manzanita. Needless to say, this caused a bit of a stir among conservationists.

The shrub had obviously been growing in that little island of serpentine soils for quite some time. The surrounding vegetation had effectively concealed its presence from the hustle and bustle of commuters that crisscross this section of on and off ramps every day. Oddly enough, this single plant likely owes its entire existence to the disturbance that created the highway in the first place. Manzanitas lay down a persistent seed bank year after year and those seeds can remain dormant until disturbance, usually fire but in this case road construction, awakens them from their slumber. It is likely that road crews had originally disturbed the serpentine soils just enough that this single Franciscan manzanita was able to germinate and survive.

The rediscovery of the last wild Franciscan manzanita was bitter sweet. On the one hand, a species thought extinct for 60 years had been rediscovered. On the other hand, this single individual was extremely stressed by years of noxious car exhaust and now, the sudden influx of sunlight due to the removal of the trees that once sheltered it. What's more, this small island of vegetation was doomed to destruction due to current highway construction. It quickly became apparent that if this plant had any chance of survival, something drastic had to be done.

Many possible rescue scenarios were considered, from cloning the plant to moving bits of it into botanical gardens. In the end, the most heroic option was decided on - this single Franciscan manzanita was going to be relocated to a managed natural area with a similar soil composition and microclimate.

Moving an established shrub is not easy, especially when that particular individual is already stressed to the max. As such, numerous safeguards were enacted to preserve the genetic legacy of this remaining wild individual just in case it did not survive the ordeal. Stem cuttings were taken so that they could be rooted and cloned in a lab. Rooted branches were cut and taken to greenhouses to be grown up to self-sustaining individuals. Numerous seeds were collected from the surprising amount of ripe fruits present on the shrub that year. Finally, soil containing years of this Franciscan manzanita's seedbank as well as the microbial community associated with the roots, were collected and stored to help in future reintroduction efforts.

Finally, the day came when the plant was to be dug up and moved. Trenches were dug around the root mass and a dozen metal pipes were driven into the soil 2 feet below the plant so that the shrub could safely be separated from the soil in which it had been growing all its life. These pipes were then bolted to I-beams and a crane was used to hoist the manzanita up and out of the precarious spot that nurtured it in secret for all those years.

Upon arriving at its new home, experts left nothing to chance. The shrub was monitored daily for the first ten days of its arrival followed by continued weekly visits after that. As anyone that gardens knows, new plantings must be babied a bit before they become established.  For over a year, this single shrub was sheltered from direct sun, pruned of any dead and sickly branches, and carefully weeded to minimize competition. Amazingly, thanks to the coordinated effort of conservationists, the state of California, and road crews, this single individual lives on in the wild.

Of course, one single individual is not enough to save this species from extinction. At current, cuttings, and seeds provide a great starting place for further reintroduction efforts. Similarly, and most importantly, a bit of foresight on the part of a handful of dedicated botanists nearly a century ago means that the presence of several unique genetic lines of this species living in botanical gardens means that at least some genetic variability can be introduced into the restoration efforts of the Franciscan manzanita.

In an ideal world, conservation would never have to start with a single remaining individual. As we all know, however, this is not an ideal world. Still, this story provides us with inspiration and a sense of hope that if we can work together, amazing things can be done to preserve and restore at least some of what has been lost. The Franciscan manzanita is but one species that desperately needs our help an attention. It is a poignant reminder to never give up and to keep working hard on protecting and restoring biodiversity.

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

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

The Mystery of the Ghost Plant

Photo by Felipe Fenrisvarg licensed under CC BY-NC-SA 2.0

As houseplants enjoy a resurgence in our culture, untold numbers of novice and expert growers alike will have undoubtedly tried their luck at a succulent or two. Succulent, of course, is not a taxonomic division, but rather a way of describing the anatomy of myriad plants adapted to harsh, dry environments around the world. One of the most common succulents in the trade is the ghost plant (Graptopetalum paraguayense).

I would bet that, if you are reading this and you grow houseplants, you have probably grown a ghost plant at one point or another. They are easy to grow and will propagate a whole new plant from only a single leaf. Despite its worldwide popularity, the ghost plant is shrouded in mystery and confusion. To date, we know next to nothing about its ecology. Much of this stems from poor record keeping and the fact that we have no idea exactly where this species originated.

That's right, we do not know the location of its native habitat. Records indicate that the first plants to find their way into human hands were imported into New York in 1904. Apparently, they were growing as "weeds" at the base of some South American cacti. Plants were lucky enough to wind up in the hands of competent botanists and the species has ended up with the name Graptopetalum paraguayense. The specific epithet "paraguayense" was an indication of much confusion to come as it was thought that the ghost plant originated in Paraguay.

Time has barely improved our knowledge. Considering many of its relatives hail from Mexico, it gradually became more apparent that South America could not claim this species as its own. Luck changed only relatively recently with the discovery of a population of a unique color variant of the ghost plant on a single mountain in northeastern Mexico. A thorough search of the area did not reveal any plants that resemble the plant so many of us know and love. It has been suggested that the original population from which the type species was described is probably growing atop an isolated mountain peak somewhere nearby in the Chihuahuan Desert.

Despite all of the mystery surrounding this species, we can nonetheless elucidate some aspects about its biology by observing plants in cultivation. It goes without saying that the ghost plant is a species of dry, nutrient-poor habitats. Its succulence and tolerance of a wide array of soil conditions is a testament to its hardy disposition. Also, if plants are grown in full sun, they develop a bluish, waxy coating on their leaves. This is likely a form of sunscreen that the plant produces to protect it from sun scorch. As such, one can assume that its native habitat is quite sunny, though its ability to tolerate shade suggests it likely shares its habitat with shrubby vegetation as well. Given enough time and proper care, ghost plants will produce sprays of erect, 5 pointed flowers. It is not known who might pollinate them in the wild.

It is always interesting to me that a plant can be so well known to growers while at the same time being a complete mystery in every other way. A search of the literature shows that most of the scientific attention given to the ghost plant centers on potentially useful compounds that can be extracted from its tissues. Such is the case for far too many plant species, both known and unknown alike. Perhaps, in the not too distant future, some intrepid botanist will at last scramble up the right mountain and rediscover the original habitat of this wonderful plant. Until then, I hope this small introduction provides you with a new found appreciation for this wonderfully adaptable houseplant.

Photo Credits: [1]

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

Leafy Cacti?

Pereskia aculeata photo by scott.zona licensed under CC BY 2.0

At first glance, there is little about a Pereskia that would suggest a relation to what we know as cacti. Even a second, third, and forth glance probably wouldn't do much to persuade the casual observer that these plants have a place on cacti family tree. All preconceptions aside, Pereskia are in fact members of the family Cactaceae and quite interesting ones at that.

Most people readily recognize the leafless, spiny green stems of a cactus. Indeed, this would appear to be a unifying character of the family. Pereskia is proof that this is not the case. Though other cacti occasionally produce either tiny, vestigial leaves or stubby succulent leaves, Pereskia really break the mold by producing broad, flattened leaves with only a hint of succulence.

Pereskia spines are produced from areoles in typical cactus fashion. Photo by Frank Vincentz licensed under CC BY-SA 3.0

What's more, instead of clusters of Opuntia-like pads or large, columnar trunks, Pereskia are mainly shrubby plants with a handful of scrambling climbers mixed in. Similar to their more succulent cousins, the trunks of Pereskia are usually adorned with clusters of long spines for protection. Additionally, each species produces the large, showy, cup-like blooms we have come to expect from cacti.

They are certainly as odd as they are beautiful. As it stands right now, taxonomists recognize two clades of Pereskia - Clade A, which are native to a region comprising the Gulf of Mexico and Caribbean Sea (this group is currently listed under the name Leuenbergeria) and Clade B, which are native to regions just south of the Amazon Basin. This may seem superficial to most of us but the distinction between these groups has a lot to teach us about the evolution of what we know of as cacti. 

Pereskia grandifolia Photo by Anne Valladares (public domain)

Genetically speaking, the genus Pereskia sorts out at the base of the cactus family tree. Pereskia are in fact sister to all other cacti. This is where the distinction between the two Pereskia clades gets interesting. Clade A appears to be the older of the two and all members of this group form bark early on in their development and their stems lack a feature present in all other cacti - stomata. Stomata are microscopic pours that allow the exchange of gases like CO2 and oxygen. Clabe B, on the other hand, delay bark formation until later in life and all of them produce stomata on their stems.

The reason this distinction is important is because all other cacti produce stomata on their stems as well. As such, their base at the bottom of the cactus tree not only shows us what the ancestral from of cactus must have looked like, it also paints a relatively detailed picture of the evolutionary trajectory of subsequent cacti lineages. It would appear that the ancestor of all cacti started out as leafy shrubs that lacked the ability to perform stem photosynthesis. Subsequent evolution saw a delay in bark formation, the presence of stomata on the stem, and the start of stem photosynthesis, which is a defining feature of all other cacti.

Pereskia aculeata Photo by Ricardosdag licensed under CC BY-SA 4.0

If you are as excited about Pereskia as I am, then you , my friend, are in luck. A handful of Pereskia species have found their way into the horticulture trade. With a little luck attention to detail, you too can share you home with one of these wonderful plants. Just be warned, they get tall and their spines, which are often hidden by the leaves, are a force to be reckoned with. Tread lightly with these wonderfully odd cacti. Celebrate their as the evolutionary wonders that they are!

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

Further Reading: [1] [2]

Of Bluebells and Fungi

Photo by Christophe Couckuyt licensed under CC BY 2.0

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

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

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

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

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

Photo by Mick Garratt licensed under CC BY-SA 2.0

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

Photo by MichaelMaggs licensed under CC BY-SA 3.0

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

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

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

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

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

Further Reading: [1] [2]

One Mustard, Many Flavors

Photo by Kurt Kulac licensed under CC BY-SA 3.0

What do kale, broccoli, cauliflower, Brussel sprouts, and cabbage have in common? They are all different cultivars of the same species!

Wild cabbage (Brassica oleracea) is native to coastal parts of southern and western Europe. In its native habitat, wild cabbage is very tolerant of salty, limey soils but not so tolerant of competition. Because of this, it tends to grow mainly on limestone sea cliffs where few other plants can dig their roots in.

Despite their popularity as delicious, healthy vegetables, as well as their long history of cultivation, there is scant record of this plant before Greek and Roman times. Some feel that this is one of the oldest plants in cultivation. Along with the countless number of edible cultivars, the wild form of Brassica oleracea can be found growing throughout the world, no doubt thanks to its popularity among humans.

I am always amazed by how little we know about crop wild relatives. Despite the popularity of its many agricultural cultivars, relatively little attention has been paid to B. oleracea in the wild. What we do know is that at least two subspecies have been identified - B. oleracea ssp. bourgeaui and B. oleracea L. ssp. oleracea. As far as anyone can tell, subspecies 'oleracea' is the most wide spread in its distribution whereas subspecies 'bourgeaui'  is only known from the Canary Islands. 

© Copyright Evelyn Simak licensed under CC BY-SA 2.0

B. oleracea's long history with humans confuses matters quite a bit. Because it has been cultivated for thousands of years, identifying which populations represent wild individuals and which represent ancient introductions is exceedingly difficult. Such investigations are made all the more difficult by a lack of funding for the kind of research that would be needed to elucidate some of these mysteries. We know so little about wild B. oleracea that the IUCN considers is a species to be "data deficient."

It seems to appreciate cool, moist areas and will sometimes escape from cultivation if conditions are right, thus leading to the confusion mentioned above. It is amazing to look at this plant and ponder all the ways in which humans have selectively bred it into the myriad shapes, sizes, and flavors we know and love (or hate) today! However, we must pay more attention to the wild progenitors of our favorite crops. They harbor much needed genetic diversity as well as clues to how these plants are going to fare as our climates continue to change.

Photo Credit: [1] [2]

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

Trout Lily Appreciation

This video is a celebration of the white trout lily (Erythronium albidum) and its various spring ephemeral neighbors. We even talk about the threat that invasive species like garlic mustard (Alliara petiolata).

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

Music by
Artist: Botanist
Track:
https://verdant-realm-botanist.bandcamp.com/

Prescribed Fire On An Illinois Prairie

Prairies are fire adapted ecosystems. For far too long, fires were sequestered. Today, more and more communities are coming around to the fact that fire can be used as a tool to bring life back to these endangered ecosystems. In this video, we get hands on experience with fire as a prairie restoration tool.

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

Music by
Artist: Stranger In My Town
Track: Terra
https://strangerinmytown.bandcamp.com/

Early Spring Ephemerals

Join us as we go in search of some of the earliest spring ephemerals. In this episode we come face to face with the aptly named harbinger of spring (Erigenia bulbosa) and the lovely Hepatica nobilis.

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

Music by
Artist: Stranger In My Town
Track: Air
https://strangerinmytown.bandcamp.com/

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

The Intriguing Pollination of a Central American Anthurium

As an avid gardener of both indoors and out, there are few better experiences than getting to see familiar plants growing in the wild for the first time. That experience is made all the better when you find out new and interesting facts about their ecology. On a recent trip to Costa Rica, I was introduced to a wide variety of Anthurium species. I marveled at how amazing these plants look in situ and was taken aback to learn that many produce flowers with intoxicating aromas.

I was also extremely fortunate to be in the presence of some aroid experts during this trip and their knowledge fueled my interest in getting up close and personal with what little time I had with these plants. They were able to ID the plants and introduce me to their biology. One species in particular has been the subject of interest in an ongoing pollination study that has proven to be unique.

The plant in question is known scientifically as Anthurium acutifolium and it is rather charming once you get to know it. It is a terrestrial plant with relatively large leaves for its overall size. Its range includes portions of lowland Costa Rica and Panama. Its flowers are typical of what one would expect out of this family. They are fused into a type of inflorescence known as a spadix and can range in color from white to green and occasionally red. If you are lucky to visit the spadix between roughly 8:00 AM and 12:30 PM, you may notice a rich scent that, to me, is impossible to describe in words.

It's this scent that sets the stage for pollination in this species. During some down time, University of Vienna grad student Florian Etl discovered that the spadix of A. acutifolium was getting a lot of attention from a particular species of small bee. Closer inspection revealed that they were all males of a species of oil-collecting bee known as Paratetrapedia chocoensis. Now, the females of these oil collecting bees are well known in the pollination literature. They visit flowers that secrete special oils that the females then use to build nests and feed their young. This is why the attention from male bees was so intriguing.

A: A male P. chocoensis bee approaching a scented spadix of an inflorescence of A. acutifolium. B: The abdominal mopping behavior of male P. chocoensis oil bees on a spadix. C: Ventral side of the abdomen of a male P.chocoensis covered with pollen. D: A male P. chocoensis bee on a spadix of an inflorescence of A. acutifolium, touching the pollen shedding anthers. E: Pubescent region pressed on the surface of A. acutifolium during the mopping behavior. F: A scented inflorescence of A. acutifolium with three male P. chocoensis individuals. G: Image of the abdomen of a male P.chocensis in lateral view showing the conspicuous pubescent region. (SOURCE)

Males would land on the spadix and begin rubbing the bottom of their abdomen along its surface. In doing so, they inevitably picked up and deposited pollen. To date, such behavior was unknown among male oil bees. What exactly were these male bees up to?

As it turns out, the males were collecting fragrances. Close inspection of their morphology revealed that each male has a small patch of dense hairs underneath their abdomen. The males are definitely not after fatty oils or nectar as A. acutifolium does not secrete either of these substances. Instead, it would appear that the male oil bees are there to collect scent, which is mopped up by that dense patch of hairs. Even more remarkable is the fact that in order to properly collect these fragrance compounds, the bees are likely using solvents that they have collected from other flowering plant species around the forest.

What they are doing with these scent compounds remains a mystery but some potential clues lie in another scent/pollination system. Male orchid bees perform similar scent-collecting activities in order to procure unique scent bouquets. Though the exact function of their scent collecting is not known either, we do know that these scents are used in the process of finding and procuring mates. It is likely that these male oil bees are using them in a similar way.

Taken together, these data suggest that a very specific pollination syndrome involving A. acutifolium and male oil bees has evolved in Central American forests. No other insects were observed visiting the flowers of A. acutifolium and the scents only ever attracted males of these specific oil bees during the hours in which the spadix was actively producing the compounds. This is a remarkable pollination syndrome and one that encourages us to start looking elsewhere in the forest. This, my friends, is why there is no substitute for simply taking the time to observe nature. We must take the time to get outside and poke around because we stand to miss out on so much of what makes our world tick and without such knowledge, we risk losing so much. 

Photo Credits: Florian Etl [1]

Further Reading: [1]

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]

Daffodil Insights

Photo by Amanda Slater licensed under CC BY-SA 2.0

Daffodils seem to be everywhere. Their horticultural popularity means that, for many of us, these plants are among the first flowers we see each spring. Daffodils are so commonplace that it's as if they evolved to live in our gardens and nowhere else. Indeed, daffodils have had a long, long history with human civilization, so much so that it is hard to say when our species first started to cohabitate. Our familiarity with these plants belies an intriguing natural history. What follows is a brief overview of the world of daffodils. 

If you are like me, then you may have gone through most of your life not noticing much difference between garden variety daffodils. Though many of us will be familiar with only a handful of daffodil species and cultivars, these introductions barely scratch the surface. One may be surprised to learn that as of 2008, more than 28,000 daffodil varieties have been named and that number continues to grow each and every year. Even outside of the garden, there is some serious debate over the number of daffodil species, much of this having to do with what constitutes a species in this group.

Narcissus poeticus

As I write this, all daffodils fall under the genus Narcissus. Estimates as to the number of species within Narcissus range from as few as 50 to as many as 80. The genus itself sits within the family Amaryllidaceae and is believed to have originated somewhere between the late Oligocene and early Miocene, some 18 to 30 million years ago. Despite its current global distribution, Narcissus are largely Mediterranean plants, with peak diversity occurring on the Iberian Peninsula. However, thanks to the aforementioned long and complicated history in cultivation, it has become quite difficult to understand the full range of diversity in form and habitat of many species. To understand this, we first need to understand a bit about their reproductive habits.

Much of the evolution of Narcissus seems to center around floral morphology and geographic isolation. More specifically, the length of the floral tube or "corona" and the position of the sexual organs within, dictates just who can effectively pollinate these plants. The corona itself is not made up of petals or sepals but instead, its tube-like appearance is due to a fusion of the stamens into the famous trumpet-like tube we know and love.

Variation in corona shape and size has led to the evolution of three major pollination strategies within this genus. The first form is the daffodil form, whose stigma is situated at the mouth of the corolla, well beyond the 6 anthers. This form is largely pollinated by larger bees. The second form is the paperwhite form, whose stigma is situated more closely to or completely below the anthers at the mouth of the corona. This form is largely pollinated by various Lepidoptera as well as long tongued bees and flies. The third form is the triandrus form, which exhibits three distinct variations on stigma and anther length, all of which are situated deep within the long, narrow corona. The pendant presentation of the flowers in this group is thought to restrict various butterflies and moths from entering the flower in favor of bees.

Narcissus tazetta. Photo by Fanghong licensed under CC BY-SA 3.0

The variations on these themes has led to much reproductive isolation among various Narcissus populations. Plants that enable one type of pollinator usually do so at the exclusion of others. Reproductive isolation plus geographic isolation brought on by differences in soil types, habitat types, and altitudinal preferences is thought to have led to a rapid radiation of these plants across the Mediterranean. All of this has gotten extremely complicated ever since humans first took a fancy to these bulbs.

Narcissus cyclamineus. Photo by Francine Riez licensed under CC BY-SA 3.0

Reproductive isolation is not perfect in these plants and natural hybrid zones do exist where the ranges of two species overlap. However, hybridization is made much easier with the helping hand of humans. Whether via landscape disturbance or direct intervention, human activity has caused an uptick in Narcissus hybridization. For centuries, we have been mixing these plants and moving them around with little to no record as to where they originated. What's more, populations frequently thought of as native are actually nothing more than naturalized individuals from ancient, long-forgotten introductions. For instance, Narcissus populations in places like China, Japan, and even Great Britain originated in this manner.

All of this mixing, matching, and hybridizing lends to some serious difficulty in delineating species boundaries. It would totally be within the bounds of reason to ask if some of the what we think of as species represent true species or simply geographic varieties on the path to further speciation. This, however, is largely speculative and will require much deeper dives into Narcissus phylogenetics.

Narcissus triandrus. Photo by Dave Gough licensed under CC BY 2.0

Despite all of the confusion surrounding accurate Narcissus taxonomy, there are in fact plenty of true species worth getting to know. These range in form and habit far more than one would expect from horticulture. There are large Narcissus and small Narcissus. There are Narcissus with yellow flowers and Narcissus with white flowers. Some species produce upright flowers and some produce pendant flowers. There are even a handful of fall-blooming Narcissus. The variety of this genus is staggering if you are not prepared for it.

Narcissus viridiflorus - a green, fall-blooming daffodil. Photo by A. Barra licensed under CC BY 3.0

After pollination, the various Narcissus employ a seed dispersal strategy that doesn't get talked about enough in reference to this group. Attached to each hard, black seed are fatty structures known as eliasomes. Eliasomes attract ants. Like many spring flowering plant species around the globe, Narcissus utilize ants as seed dispersers. Ants pick up the seeds and bring them back to their nests. They go about removing the eliasomes and then discard the seed. The seed, safely tucked away in a nutrient-rich ant midden, has a much higher chance of germination and survival than if things were left up to simple chance. It remains to be seen whether or not Narcissus obtain similar seed dispersal benefits from ants outside of their native range. Certainly Narcissus populations persist and naturalize readily, however, I am not aware if ants have any part in the matter.

The endangered Narcissus alcaracensis. Photo by José Luis López González licensed under CC BY-SA 4.0

Despite their popularity in the garden, many Narcissus are having a hard go of it in the wild. Habitat destruction, climate change, and rampant collecting of wild bulbs are having serious impacts on Narcissus numbers. The IUCN considered at least 5 species to be endangered and a handful of some of the smaller species already thought to be extinct in the wild. In response to some of these issues, protected areas have been established that encompass at least some of the healthy populations that remain for some of these species.

If you are anything like me, you have ignored Narcissus for far too long. Sure, they aren't native to the continent on which I live, and sure, they are one of the most commonly used plants in a garden setting, but every species has a story to tell. I hope that, armed with this new knowledge, you at least take a second look at the Narcissus popping up around your neighborhood. More importantly, I hope this introduction makes you appreciate their wild origins and the fact that we still have much to learn about these plants. I have barely scratched the surface of this genus and there is more more information out there worth perusing. Finally, I hope we can do better for the wild progenitors of our favorite garden plants. They need all the help they can get and unless we start speaking up and working to preserve wild spaces, all that will remain are what we have in our gardens and that is not a future I want to be a part of.

Photo Credits: [1] [2] [3] [4] [5] [6] [7]

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

Do Yeasts Aid Pollination For the Stinking Hellebore?

Photo by Mark Gurney licensed under CC BY-NC-SA 2.0

Whether they are growing in their native habitat or in some far away garden, Hellebores are some of the earliest plants to bloom in the spring. Hellebore flowers can often be seen blooming long before the snow has melted away. All early blooming plant species are faced with the challenge of attracting pollinators. Though the competition for insect attention is minimal among these early bloomers, only the hardiest insects are out and about on cold, dreary days. It stands to reason then that anything that can entice a potential pollinator would be of great benefit for a plant.

That is why the presence of yeast in the nectar of at least one species of Hellebore has attracted the attention of scientists. The species in question is known scientifically as Helleborus foetidus. The lack of appeal in its binomial is nothing compared to its various common names. One can often find H. foetidus for sale under names like the "stinking hellebore" or worse, "dungwort." All of these have to do with the unpleasant aroma given off by its flowers and bruised foliage. Surprisingly, that is not the topic of this post.

Photo by Bernd Haynold licensed under CC BY-SA 3.0

What is more intriguing about the flowers of H. foetidus is that the nectar produced by its smelly green flowers harbors dense colonies of yeast. Yeasts are everywhere on this planet and despite their economic importance, little is known about how they function in nature. For instance, what the heck are these yeast colonies doing in the nectar of this odd Hellebore?

To test this, two researchers from the Spanish National Research Council manipulated yeast colonies within the flowers to see what might be happening. It turns out, yeast in the nectar of H. foetidus actually warms the flowers. As the yeast feed on the sugars within the nectar, their metabolic activity can raise the temperature of the flowers upwards of 2 °C above the ambient. As far as we know, the only other ways in which floral heating has been achieved is either via specific metabolic processes within the floral tissues or by direct heating from the sun. 

In heating the flowers, these yeast colonies may be having serious impacts on the reproductive success of H. foetidus. For starters, these plants are most at home under the forest canopies of central and western Europe. What's more, many populations find themselves growing in the dense shade of evergreens. This completely rules out the ability to utilize solar energy to heat blooms. Additionally, floral heat can mean more visits by potential pollinators. Experiments have shown that bees preferentially visit flowers that are slightly warmer than ambient temperatures. Even the flowers themselves can benefit from that heat. Warmer flowers have higher pollination rates and better seed set.

Bombus terrestris was one of the most common floral visitors of Helleborus foetidus. Photo by Vera Buhl licensed under CC BY-SA 3.0

Yeast colonies also have their downsides. The heat generated by the yeast comes from the digestion of sugars. Indeed, nectar housing yeast colonies had drastically reduced sugar loads than nectar without yeast. This has the potential to offset many of the benefits of floral warming in large part because bees are good at discriminating. Bees are visiting these blooms as a food source and by diminishing the sugar content of the nectar, the yeast may be turning bees off to this potential source. The question then becomes, do bees prefer heat over sugar-rich food? The authors think there might be a trade-off, with bees preferring heated flowers on colder days and sugar-rich flowers on warmer days.

Helleborus foetidus flowering before the snow has had a chance to melt!

Though the authors found evidence for heating, they did not test for pollinator preference. All we know at this point is that yeast in the nectar significantly warms H. foetidus flowers. Since this piece was originally published, more attention has been paid to the benefits of the heat generated from yeast. Interestingly, researchers found that pollen tube formation was higher for H. foetidus flowers that experienced heat earlier in the season but not for those that experienced heat later on. This response, however, was not due to the warming directly. Instead, it had more to do with bee preference.

As it turns out, bumblebees do in fact prefer to visit heated flowers but their preference is limited to the early periods of flowering when ambient temperatures are still quite low. More bumblebees visiting heated flowers in the early spring equated to more pollen being deposited on the stigma, which in turn led to an increase in pollen tube formation and higher seed set. Later on in the season, when ambient temperatures increased a bit, this positive effect dropped off as bees apparently spent more time foraging elsewhere.

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

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