Meet the Fire Lily


The flora of the South African fynbos region is no stranger to fire. Many species have adapted to cope with and even rely on fire to complete their lifecycles. There is one species, however, that takes this to the extreme. It is a tiny member of the Amaryllidaceae aptly named the fire lily (Cyrtanthus ventricosus).

The fire lily is not a big plant by any means. Mature individuals can top out around 9 inches (250 mm) and for most of the year consist of a nothing more than a small cluster of narrow, linear leaves. As the dry months of summer approach, the leaves senesce and the plant more or less disappears until its time to flower. However, unlike other plants in this region that flower more regularly, the fire lily lies in wait for a very specific flowering cue - smoke.

It has been noted that fire lilies only seem to want to reproduce after a fire. No other environmental factor seems to trigger flowering. This has made them quite frustrating for bulb aficionados. Only after a fire burns over the landscape will a scape emerge topped with anywhere from 1 to 12 tubular red flowers.


This dependence on fire for flowering has garnered the attention of a few botanists concerned with conservation of pyrophytic geophytes. Obviously if we care about conserving species like the fire lily, it is extremely important that we understand their reproductive ecology. The question of fire lily blooming is one of triggers. What part of the burning process triggers these plants to bloom?

By experimenting with various burn and smoke treatments, researchers were able to deduce that it wasn’t heat that triggered flowering but rather something in the smoke itself. Though researchers were not able to isolate the exact chemical(s) responsible, at least we now know that fire lilies can be coaxed into flowering using smoke alone. This is a real boon to growers and conservationists alike.


Seeing a population of fire lilies in full bloom must be an incredible sight. Within only a few days of a fire, huge patches of bright red flowers decorate the charred landscape. They are borne on hollow stalks which provide lots of structural integrity while being cheap to produce. The flowers themselves are not scented but they do produce a fair amount of nectar. The bright red inflorescence mainly attracts the Table Mountain pride butterfly as well as sunbirds.

Once flowering is complete, seeds are produced and the plants return to their dormant bulbous state until winter when leaves emerge again. Flowering will not happen again until fire returns to clear the landscape. This strategy may seem inefficient on the part of the plant. Why not attempt to reproduce every year? The answer is competition. By waiting for fire, this tiny plant is able to make a big impact despite being so small. It would be impossible to miss their enticing floral display when all other vegetation has been burned away.

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

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

A Passionflower With a Taste for Insects?


For a plant to be considered carnivorous, it must possess one or more traits unequivocally adapted for attracting, capturing, and/or digesting prey. It also helps to demonstrate that the absorption of nutrients has a clear positive impact on growth or reproductive effort. For plants like the Venus fly trap or any of the various pitcher plants out there, this distinction is pretty straight forward. For many other species, the line between carnivorous or not can be a little blurry. Take, for instance, the case of the stinking passionflower (Passiflora foetida).

At first glance, P. foetida seems par for the course as far as passionflowers are concerned. It is a vining species native from the southwestern United States all the way down into South America. It enjoys edge habitats where it can scramble up and over neighboring vegetation. It produces large, showy flowers followed by edible fruits. When the foliage is damaged, it emits a strong odor, earning it the specific epithet “foetida.”

Not until you inspect the developing floral buds of this passionflower will the question of carnivory enter into your mind. Covering the developing flowers and eventually the fruit are a series of feathery bracts, which are covered in glandular hairs. The hairs themselves are quite sticky thanks to the secretion of fluids. As insects crawl across the hairs, they become hopelessly entangled and eventually die. So, does this make P. foetida a carnivore?


Many different plants produce sticky hairs or glands on their tissues. Often this is a form of defense. Herbivorous insects looking to take a bite out of such a plant either get stuck outright or have their mouth parts completely gummed up in the process. This form of defense seems to work quite well for such plant species so simply trapping insects doesn’t mean the plant is a carnivore. Worth noting, however, is the fact that it appears that many carnivorous plant traits have simply been retooled from defense traits.

The question remains as to what happens to the trapped insects after they are ensnared by P. foetida. Observations in the field suggest that there is more to these sticky hairs than simply defense. This led a team of researchers to look closer at the interactions between P. foetida and insects. What they found is rather fascinating.

It turns out that most of the insects captured by P. foetida bracts are herbivores that would have made an easy meal of the flowers and fruits. However, after getting stuck, the insect bodies quickly decay. Laboratory analyses revealed that indeed, the fluids secreted by the sticky hairs contained lots of digestive enzymes, mainly proteases and acid phosphatases. Still, this does not mean the plant is eating the insects. It makes sense from a defensive standpoint that a plant would not benefit from having lots of rotting corpses stuck to its buds. As such, digesting them removes the possibility of fungal or bacterial attack. To investigate whether P. foetida benefits from trapping insects beyond simply avoiding herbivory, the team needed to know if any nutritional benefit was being had.


The team took amino acids marked with a special carbon isotope and smeared it onto the bracts. Then they waited to see if any of the labelled amino acids showed up in the plant tissues. Indeed they did. The amino acids were absorbed by the bracts and translocated to the  calyx, corolla, anthers, and finally to the developing ovules. This is probably not too surprising  to those of us that spend time growing plants as numerous plant species can uptake at least some nutrients through their leaves. This is why foliar feeding can work as a means of fertilizing potted plants. Nonetheless, these results are enticing as it shows that P. foetida is not only capturing and dissolving insects, it also seems capable of absorbing at least some amino acids from its victims.

So, should we call P. foetida a carnivore? To be honest, I am not sure. Certainly all of the evidence suggests there is more going on than simply defense. However, does garnering the attention of hungry herbivores constitute prey attraction? Certainly other carnivores utilize food deception as a means of prey capture. Does simply being a palatable plant count as a lure? Does absorbing nutrients constitute carnivory? In some instances, yes, however, as mentioned, plenty of plant species can absorb nutrients from organs other than their roots.

I think the main question is whether P. foetida sees a marked increase in growth or reproduction due to the addition of the dead herbivores. What I think we can say is that the sticky bracts surrounding the flowers and fruits serve a dual purpose - defense against herbivores and potentially a nutrient boost as well. If anything, I think this should qualify as a form of protocarnivory.

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

Further Reading: [1] [2]  

The Rise and Fall of the Scale Trees


If I had a time machine, the first place I would visit would be the Carboniferous. Spanning from 358.9 to 298.9 million years ago, this was a strange time in Earth’s history. The continents were jumbled together into two great landmasses - Laurasia to the north and Gondwana to the south and the equatorial regions were dominated by humid, tropical swamps. To explore these swamps would be to explore one of the most alien landscapes this world has ever known.

The Carboniferous was the heyday for early land plants. Giant lycopods, ferns, and horsetails formed the backbone of terrestrial ecosystems. By far the most abundant plants during these times were a group of giant, tree-like lycopsids known as the scale trees. Scale trees collectively make up the extinct genus Lepidodendron and despite constantly being compared to modern day club mosses (Lycopodiopsida), experts believe they were more closely related to the quillworts (Isoetopsida).

 Carboniferous coal swamp reconstruction dating back to the 1800’s

Carboniferous coal swamp reconstruction dating back to the 1800’s

It is hard to say for sure just how many species of scale tree there were. Early on, each fragmentary fossil was given its own unique taxonomic classification; a branch was considered to be one species while a root fragment was considered to be another and juvenile tree fossils were classified differently than adults. As more complete specimens were unearthed, a better picture of scale tree diversity started to emerge. Today I can find references to anywhere between 4 and 13 named species of scale tree and surely more await discovery. What we can say for sure is that scale tree biology was bizarre.

The name “scale tree” stems from the fossilized remains of their bark, which resembles reptile skin more than it does anything botanical. Fossilized trunk and stem casts are adorned with diamond shaped impressions arranged in rows of ascending spirals. These are not scales, of course, but rather they are leaf scars. In life, scale trees were adorned with long, needle-like leaves, each with a single vein for plumbing. Before the started branching, young trees would have resembled a bushy, green bottle brush.

 Juvenile scale tree on the left & the adult on the right

Juvenile scale tree on the left & the adult on the right

As scale trees grew, it is likely that they shed their lower leaves, which left behind the characteristic diamond patterns that make their fossils so recognizable. How these plants achieved growth is rather fascinating. Scale tree cambium was unifacial, meaning it only produced cells towards its interior, not in both directions as we see in modern trees. As such, only secondary xylem was produced. Overall, scale trees would not have been very woody plants. Most of the interior of the trunk and stems was comprised of a spongy cortical meristem. Because of this, the structural integrity of the plant relied on the thick outer “bark.” Many paleobotanists believe that this anatomical quirk made scale trees vulnerable to high winds.

Scale trees were anchored into their peaty substrate by rather peculiar roots. Originally described as a separate species, the roots of these trees still retain their species name. Paleobotanists refer to them as “stigmaria” and they were unlike most roots we encounter today. Stigmaria were large, limb-like structures that branched dichotomously in the soil. Each main branch was covered in tiny spots that were also arranged in rows of ascending spirals. At each spot, a rootlet would have grown outward, likely partnering with mycorrhizal fungi in search of water and nutrients.

 A preserved  Lepidodendron  stump

A preserved Lepidodendron stump

Eventually scale trees would reach a height in which branching began. Their tree-like canopy was also the result of dichotomous branching of each new stem. Amazingly, the scale tree canopy reached staggering heights. Some specimens have been found that were an estimated 100 ft (30 m) tall! It was once thought that scale trees reached these lofty heights in as little as 10 to 15 years, which is absolutely bonkers to think about. However, more recent estimates have cast doubt on these numbers. The authors of one paper suggest that there is no biological mechanism available that could explain such rapid growth rates, concluding that the life span of a typical scale tree was more likely measured in centuries rather than years.

Regardless of how long it took them to reach such heights, they nonetheless would have been impressive sites. Remarkably, enough of these trees have been preserved in situ that we can actually get a sense for how these swampy habitats would have been structured. Whenever preserved stumps have been found, paleobotanists remark on the density of their stems. Scale trees did not seem to suffer much from overcrowding.


The fact that they spent most of their life as a single, unbranched stem may have allowed for more success in such dense situations. In fact, those that have been lucky enough to explore these fossilized forests often comment on how similar their structure seems compared to modern day cypress swamps. It appears that warm, water-logged conditions present similar selection pressures today as they did 350+ million years ago.

Like all living things, scale trees eventually had to reproduce. From the tips of their dichotomosly branching stems emerged spore-bearing cones. The fact that they emerge from the growing tips of the branches suggests that each scale tree only got one shot at reproduction. Again, analyses of some fossilized scale tree forests suggests that these plants were monocarpic, meaning each plant died after a single reproductive event. In fact, fossilized remains of a scale tree forest in Illinois suggests that mass reproductive events may have been the standard for at least some species. Scale trees would all have established at around the same time, grown up together, and then reproduced and died en masse. Their death would have cleared the way for their developing offspring. What an experience that must have been for any insect flying around these ancient swamps.

 The fossilized strobilus of a Lepidodendron

The fossilized strobilus of a Lepidodendron

Compared to modern day angiosperms, the habits of the various scale trees may seem a bit inefficient. Nonetheless, this was an extremely successful lineage of plants. Scale trees were the dominant players of the warm, humid, equatorial swamps. However, their dominance on the landscape may have actually been their downfall. In fact, scale trees may have helped bring about an ice age that marked the end of the Carboniferous.

You see, while plants were busy experimenting with building ever taller, more complex anatomies using compounds such as cellulose and lignin, the fungal communities of that time had not yet figured out how to digest them. As these trees grew into 100 ft monsters and died, more and more carbon was being tied up in plant tissues that simply weren’t decomposing. This lack of decomposition is why we humans have had so much Carboniferous coal available to us. It also meant that tons of CO2, a potent greenhouse gas, were being pulled out of the atmosphere millennia after millennia.

 A fossilized root or “stigmaria”

A fossilized root or “stigmaria”

As atmospheric CO2 levels plummeted and continents continued to shift, the climate was growing more and more seasonal. This was bad news for the scale trees. All evidence suggests that they were not capable of keeping up with the changes that they themselves had a big part in bringing about. By the end of the Carboniferous, Earth had dipped into an ice age. Earth’s new climate regime appeared to be too much for the scale trees to handle and they were driven to extinction. The world they left behind was primed and ready for new players. The Permian would see a whole new set of plants take over the land and would set the stage for even more terrestrial life to explode onto the scene.

It is amazing to think that we owe much of our industrialized society to scale trees whose leaves captured CO2 and turned it into usable carbon so many millions of years ago. It seems oddly fitting that, thanks to us, scale trees are once again changing Earth’s climate. As we continue to pump Carboniferous CO2 into our atmosphere, one must stop to ask themselves which dominant organisms are most at risk from all of this recent climate change?

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

Further Reading: [1] [2] [3] [4] [5] [6] [7] [8] [9] [10] [11] [12] [13] [14]

The Japanese Umbrella Pine


My first impression of the Japanese umbrella pine was that I was looking at a species of yew (Taxus spp.). Sure, its features were a bit more exaggerated than I was used to but what do I know? Trying to understand tree diversity is a recent development in my botanical obsession so I don’t have much to base my opinions on. Regardless, I am glad I gave the little sapling I was looking at a closer inspection. Turns out, the Japanese umbrella pine is most definitely not a yew. It is actually unique in its taxonomic position as the only member of the family Sciadopityaceae.

The Japanese umbrella pine goes by the scientific name of Sciadopitys verticillata. Both common and scientific names hint at the whorled arrangements of its “leaves.” I place leaves in quotes because they are not leaves at all. One of the most remarkable features of this tree is the fact that those whorled leaves are actually thickened, photosynthetic extensions of the stem known as “cladodes.”

 Those tiny bumps along the stems are actually highly reduced leaves whereas the whorls of photosynthetic “leaves” are actually modified extensions of the stem called “cladodes.”

Those tiny bumps along the stems are actually highly reduced leaves whereas the whorls of photosynthetic “leaves” are actually modified extensions of the stem called “cladodes.”


It seems that the true leaves of the Japanese umbrella pine have, through evolutionary time, been reduced to tiny, brown scales that clasp the stems. I am not sure what evolutionary advantage(s) cladodes infer over leaves, however, at least one source suggested that cladodes may have fewer stomata and therefore can help to reduce water loss. Until someone looks deeper into this mystery, we cannot say for sure.

As a tree, the Japanese umbrella pine is slow growing. Records show that young trees can take upwards of a decade to reach average human height. However, given time, the Japanese umbrella pine can grow into an impressive specimen. In the forests of Japan, it is possible to come across trees that are 65 to 100 ft (20 – 35 m) tall. It was once wide spread throughout much of southern Japan, however, an ever-increasing human population has seen that range reduced.

 A 49.5 million years old fossil of a  Sciadopitys  cladode.

A 49.5 million years old fossil of a Sciadopitys cladode.

The gradual reduction of this species is not solely the fault of humans. Fossil evidence shows that the genus Sciadopitys was once wide spread throughout parts of Europe and Asia as well. Whereas the current diversity of this genus is limited to a single species, fossils of at least three extinct species have been found in rocks dating back to the Triassic Period, some 230 million years ago. It would appear that this obscure conifer family, like so many other gymnosperm lineages, has been on the decline for quite some time.

Despite the obscure strangeness of the Japanese umbrella tree, it has gained considerable popularity as a unique landscape tree. Because it hails from a relatively cool regions of Japan, the Japanese umbrella tree adapts quite well to temperate climates around the globe. Enough people have grown this tree that some cultivars even exist. Whether you see it as a specimen in an arboretum or growing in the wild, know that you are looking at something quite special. The Japanese umbrella tree is a throwback to the days when gymnosperms were the dominant plants on the landscape and we are extremely lucky that it made it through to our time.

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

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

Resurrecting Café Marron


Back in 1980, a school teacher on the island of Rodrigues sent his students out to look for plants. One of the students brought back a cutting of a shrub that astounded the botanical community. Ramosmania rodriguesii, more commonly known as café marron, was up until that point only known from one botanical description dating back to the 1800's. The shrub, which is a member of the coffee family, was thought to have been extinct due to pressures brought about during the colonization of the island (goats, invasive species, etc.). What the boy brought back was indeed a specimen of café marron but the individual he found turned out to be the only remaining plant on the island.

News of the plant quickly spread. It started to attract a lot of attention, not all of which was good. There is a belief among the locals that the plant is an herbal remedy for hangovers and venereal disease (hence its common name translates to ‘brown coffee’) and because of that, poaching was rampant. Branches and leaves were being hauled off at a rate that was sure to kill this single individual. It was so bad that multiple layers of fencing had to be erected to keep people away. It was clear that more was needed to save this shrub from certain extinction.


Cuttings were taken and sent to Kew. After some trial and tribulation, a few of the cuttings successfully rooted. The clones grew and flourished. They even flowered on a regular basis. For a moment it looked like this plant had a chance. Unfortunately, café marron did not seem to want to self-pollinate. It was looking like this species was going to remain a so-called “living dead” representative of a species no longer able to live in the wild. That is until Carlos Magdalena (the man who saved the rarest water lily from extinction) got his hands on the plants.

The key to saving café marron was to somehow bypass its anti-selfing mechanism. Because so little was known about its biology, there was a lot of mystery surrounding its breeding mechanism. Though plenty of flowers were produced, it would appear that the only thing working on the plant were its anthers. They could get viable pollen but none of the stigmas appeared to be receptive. Could it be that the last remaining individual (and all of its subsequent clones) were males?


This is where a little creativity and a lot of experience paid off. During some experiments with the flowers, it was discovered that by amputating the top of the stigma and placing pollen directly onto the wound one could coax fertilization ans fruiting. From that initial fruit, seven seeds were produced. These seeds were quickly sent to the propagation lab but unfortunately the seedlings were never able to establish. Still, this was the first indication that there was some hope left for the café marron.

After subsequent attempts at the stigma amputation method ended in failure, it was decided that perhaps something about the growing conditions of the first plant were the missing piece of this puzzle. Indeed, by repeating the same conditions the first individual was exposed to, Carlos and his team were able to coax some changes out of the flowering efforts of some clones. Plants growing in warmer conditions started to produce flowers of a slightly different morphology towards the end of the blooming cycle. After nearly 300 attempts at pollinating these flowers, a handful of fruits were formed!


From these fruits, over 100 viable seeds were produced. What’s more, these seeds germinated and the seedlings successfully established. Even more exciting, the seedlings were a healthy mix of both male and female plants. Carlos and his team learned a lot about the biology of this species in the process. Thanks to their dedicated work, we now know that café marron is protandrous meaning its male flowers are produced before female flowers.

However, the story doesn’t end here. Something surprising happened as the seedlings continued to grow. The resulting offspring looked nothing like the adult plant. Whereas the adult plant has round, green leaves, the juveniles were brownish and lance shaped. This was quite a puzzle but not entirely surprising because the immature stage of this shrub was not known to science. Amazingly, as the plants matured they eventually morphed into the adult form. It would appear that there is more to the mystery of this species than botanists ever realized. The question remained, why go through such drastically different life stages?

 A young café marron showing its brown, mottled, lance-shaped leaves.

A young café marron showing its brown, mottled, lance-shaped leaves.

The answer has to do with café marron's natural predator, a species of giant tortoise. The tortoises are attracted to the bright green leaves of the adult plant. By growing dull, brown, skinny leaves while it is still at convenient grazing height, the plant makes itself almost invisible to the tortoise. It is not until the plant is out of the range of this armoured herbivore that it morphs into its adult form. Essentially the young plants camouflage themselves from the most prominent herbivore on the island.

Thanks to the efforts of Carlos and his team at Kew, over 1000 seeds have been produced and half of those seeds were sent back to Rodrigues to be used in restoration efforts. As of 2010, 300 of those seed have been germinated, opening up many more opportunities for reintroduction into the wild. Those early trials will set the stage for more restoration efforts in the future. It is rare that we see such an amazing success story when it comes to such an endangered species. We must celebrate these efforts because they remind us to keep trying even if all hope seems to be lost. My hat is off to Carlos and the dedicated team of plant conservationists and growers at Kew.

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

Further Reading: [1] [2]

Meet the Pygmy Clubmoss


No, these are not some sort of grass or rush. What you are looking at here is actually a member of the clubmoss family (Lycopodiaceae). Colloquially known as the pygmy clubmoss, this odd little plant is the only species in its genus - Phylloglossum drummondii. Despite its peculiar nature, very little is known about it.

The pygmy clubmoss is native to parts of Australia, Tasmania, and New Zealand but common it is not. From what I can gather, it grows in scattered coastal and lowland sites where regular fires clear the ground of competing vegetation. It is a perennial plant that makes its appearance around July and reaches reproductive size around August through to October.

Reproduction for the pygmy clubmoss is what you would expect from this family. In dividual plants will produce a reproductive stem that is tipped with a cone-like structure. This cone houses the spores, which are dispersed by wind. If a spore lands in a suitable spot, it germinates into a tiny gametophyte. As you can probably imagine, the gametophyte is small and hard to locate. As such, little is known about this part of its life cycle. Like all gametophytes, the end goal of this stage is sexual reproduction. Sperm are released and with any luck will find a female gametophyte and fertilize the ovules within. From the fertilized ovule emerges the sporophytes we see pictured above.


As dormancy approaches, this strange clubmoss retreats underground where it persists as a tiny tuber-like stem. Though it is rather obscure no matter who you ask, there has been some scientific attention paid to this odd little plant, especially as it relates to its position on the tree of life. Since it was first described, its taxonomic affinity has moved around a bit. Early debates seemed to center around whether it belonged in Lycopodiaceae or its own family, Phylloglossaceae.


Recent molecular work put this to rest showing that genetically the pygmy clubmoss is most closely related to another genus of clubmoss - Huperzia. This was bolstered by the fact that it shares a lot of features with this group such as spore morphology, phytochemistry, and chromosome number. The biggest difference between these two genera is the development of the pygmy clubmoss tuber, which is unique to this species. However, even this seems to have its roots in Lycopodiaceae.

If you look closely at the development of some lycopods, it becomes apparent that the pygmy clubmoss most closely resembles an early stage of development called the “protocorm.” Protocorms are a tuberous mass of cells that is the embryonic form of clubmosses (as well as orchids). Essentially, the pygmy clubmoss is so similar to the protocorm of some lycopods that some experts actually think of it as a permanent protocorm capable of sexual reproduction. Quite amazing if you ask me.

Phylloglossum drummondii   003.jpg

Sadly, because of its obscurity, many feel this plant may be approaching endangered status. There have been notable declines in population size throughout its range thanks to things like conversion of its habitat to farmland, over-collection for both novelty and scientific purposes, and sequestration of life-giving fires. As mentioned, the pygmy clubmoss needs fire. Without it, natural vegetative succession quickly crowds out these delicate little plants. Hopefully more attention coupled with better land management can save this odd clubmoss from going the way of its Carboniferous relatives.

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

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

A New Species of Waterfall Specialist Has Been Discovered In Africa

Lebbiea grandiflora Oct2018.JPG

At first glance, this odd plant doesn’t look very special. However, it is the first new member of the family Podostemaceae to be found in Africa in over 30 years. It has been given the name Lebbiea grandiflora and it was discovered during a survey to assess the impacts of a proposed hydroelectric dam. By examining the specimen, Kew botanists quickly realized this plant was unique. Sadly, if all goes according to plan, this species may not be long for this world unless something is done to preserve it.

Members of the family Podostemaceae are strange plants. Despite how delicate they look, these plants specialize in growing submersed on rocks in waterfalls, rapids, and other fast flowing bodies of water. They are generally small plants, though some species can grow to lengths of 3 ft. (1 m) or more. The best generalization one can make about this group is that they like clean, fast-flowing water with plenty of available rock surfaces to grow on.

Lebbiea grandiflora certainly fits this description. It is native to a small portion of Sierra Leone and Guinea where it grows on slick rock surfaces only during the wet season. As the dry season approaches and the rivers shrink in size, L. grandiflora quickly sets seed and dies.

As mentioned, the area in which this plant was discovered is slated for the construction of a large hydroelectric dam. The building of this dam will most certainly destroy the entire population of this plant. As soon as water slows, becomes more turbid, and sediments build up, most Podostemaceae simply disappear. Unfortunately, I appears this plant was in trouble even before the dam came into the picture.

 A. habit, whole plant, in fruit, showing the flat root, a pillar-like ‘haptera’, and a shoot with three inflorescences, B. detail of shoot with three branches, C. view of upper surface of a flattened root, with six short, erect shoots, each with 1–2 1-flowered inflorescences emerging from spathellum remains, D. side view of plant showing, on the lower surface of the flattened root, the pillar-like haptera, branched at base; upper surface of root with spathellum-sheathed inflorescence base, E. plant attached to rock by weft of thread-like root hairs (indicated with arrow) from base of pillar-like haptera; upper surface of flattened root with two shoots, F. side view of flower showing one of two tepals in full frontal view, G. as F. with tepal removed, exposing the gynoecium with, to left, the arched-over androecium, H. side view of flower with androecium in centre, two tepals flanking the gynoecium, I. androecium (leftmost of three anthers missing), J. transverse section of andropodium, K. view of gynoecium from above showing funneliform style-stigma base, L. fruit, dehisced, M. transverse section of bilocular fruit, showing septum and placentae, N. placentae with seeds, divided by septum, O. seeds, P. seed with mucilage outer layer. Drawn by Andrew Brown from  Lebbie  A2721  [SOURCE]

A. habit, whole plant, in fruit, showing the flat root, a pillar-like ‘haptera’, and a shoot with three inflorescences, B. detail of shoot with three branches, C. view of upper surface of a flattened root, with six short, erect shoots, each with 1–2 1-flowered inflorescences emerging from spathellum remains, D. side view of plant showing, on the lower surface of the flattened root, the pillar-like haptera, branched at base; upper surface of root with spathellum-sheathed inflorescence base, E. plant attached to rock by weft of thread-like root hairs (indicated with arrow) from base of pillar-like haptera; upper surface of flattened root with two shoots, F. side view of flower showing one of two tepals in full frontal view, G. as F. with tepal removed, exposing the gynoecium with, to left, the arched-over androecium, H. side view of flower with androecium in centre, two tepals flanking the gynoecium, I. androecium (leftmost of three anthers missing), J. transverse section of andropodium, K. view of gynoecium from above showing funneliform style-stigma base, L. fruit, dehisced, M. transverse section of bilocular fruit, showing septum and placentae, N. placentae with seeds, divided by septum, O. seeds, P. seed with mucilage outer layer. Drawn by Andrew Brown from Lebbie A2721 [SOURCE]

As mentioned, Podostemaceae need clean rock surfaces on which to germinate and grow. Without them, the seedlings simply can’t get established. Mining operations further upstream of the Sewa Rapids have been dumping mass quantities of sediment into the river for years. All of this sediment eventually makes it down into L. grandiflora territory and chokes out available germination sites.

Alarmed at the likely extinction of this new species, the Kew team wanted to try and find other populations of L. grandiflora. Amazingly, one other population was found growing in a river near Koukoutamba, Guinea. Sadly, the discovery of this additional population is bitter sweet as the World Bank is apparently backing another hydro-electric dam project on that river as well.

The only hope for the continuation of this species currently will be to (hopefully) find more populations and collect seed to establish ex situ populations both in other rivers as well as in captivity if possible. To date, no successful purposeful seeding of any Podostemaceae has been reported (if you know of any, please speak up!). Currently L. grandiflora has been given “Critically Endangered” status by the IUCN and the botanists responsible for its discovery hope that, coupled with the publication of this new species description, more can be done to protect this small rheophytic herb.

Photo Credit: [1] [2]

Further Reading: [1]

On the Flora of Antarctica


Antarctica - the frozen continent. It is hard to think of a place on Earth that is less hospitable to life. Yet life does exist here and some of it is botanical. Though few in number, Anarctica’s diminutive flora is able to eke out an existence wherever the right conditions present themselves. It goes without saying that these plants are some of the hardiest around.

It is strange to think of Antarctica as having any flora at all. How many descriptions of plant families and genera say something to the effect of “found on nearly every continent except for Antarctica.” It didn’t always used to be this way though. Antarctica was once home to a diverse floral assemblage that rivaled anything we see in the tropics today. Millions upon millions of years of continental drift has seen this once lush landmass positioned squarely at Earth’s southern pole.


Situated that far south, Antarctica has long since become a frozen wasteland of sorts. The landscape is essentially a desert. Instead of no precipitation, however, most water in this neck of the woods is completely locked up in ice for most of the year. This is one reason why plants have had such a hard time making a living here. That is not to say that some plants haven’t made it. In fact, a handful of species thrive under these conditions.

When anyone goes looking for plants in Antarctica, they must do so wherever conditions ease up enough for part of the year to allow terrestrial life to exist. In the case of this frozen continent, this means hanging out along the coast or one of handful of islands situated just off of the mainland. Here, enough land thaws during the brief summer months to allow a few plant species to take root and grow.

 Antarctic hair grass ( Deschamsia antarctica )

Antarctic hair grass (Deschamsia antarctica)

The flora of Antarctica proper consists of 2 flowering plant species, about 100 species of mosses, and roughly 30 species of liverwort. The largest of these are the flowering plants - a grass known as Antarctic hair grass (Deschamsia antarctica), and member of the pink family with a cushion-like growth habit called Antarctic pearlwort (Colobanthus quitensis). Whereas the hair grass benefits from being wind pollinated, the Antarctic pearlwort has had to get creative with its reproductive needs. Instead of relying on pollinators, which simply aren’t present in any abundance on Antarctica, it appears that the pearlwort has shifted over to being entirely self-pollinated. This seems to work for it because if the mother plant is capable of living on Antarctica, so too will its clonal offspring.

By far the dominant plant life on the continent are the mosses. With 100 species known to live on Antarctica, it is hard to make generalizations about their habits other than to say they are pretty tough plants. Most live out their lives among the saturated rocks of the intertidal zones. What we can say about these mosses is that they support a bewildering array of microbial life, from fungi and lichens to protists and tardigrades. Even in this frozen corner of the world, plants form the foundation for all other forms of life.

 Antarctic pearlwort ( Colobanthus quitensis )

Antarctic pearlwort (Colobanthus quitensis)

The coastal plant communities of Antarctica represent hotbeds of biodiversity for this depauperate continent. They reach their highest densities on the Antarctic Peninsula as well as on coastal islands such as south Orkney Islands and the South Shetland Islands. Here, conditions are just mild enough among the various rocky crevices for germination and growth to occur. Still, life on Antarctica is no cake walk. A short growing season, punishing waves, blistering winds, and trampling by penguins and seals present quite a challenge to Antarctica’s botanical denizens. They are able to live here despite these challenges.


Still, humans take their toll. The Antarctic Peninsula is experiencing some of the most rapid warming on the planet over the last century. As this region grows warmer and drier each year, plants are responding accordingly. Antarctic mosses along the peninsula are increasingly showing signs of stress. They are starting to prioritize the production of protective pigments in their tissues over growth and reproduction. Moreover, new species of moss are starting to take over. Rapid warming and drying of the Antarctic Peninsula appears to be favoring species that are more desiccation tolerant at the expense of the continents endemic moss species.

Changes in the structure and composition of Antarctica’s moss beds is far from being a scientific curiosity for only bryologists to ponder. It is a symptom of greater changes to come.

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

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

How a Tropical Conifer May Hold the Key to Kākāpō Recovery


The plight of the kākāpō is a tragedy. Once the third most common bird in New Zealand, this large, flightless parrot has seen its numbers reduced to less than 150. In fact, for a time, it was even thought to be extinct. Today, serious effort has been put forth to try and recover this species from the brink of extinction. It has long been recognized that kākāpō breeding efforts are conspicuously tied to the phenology of certain trees but recent research suggests one in particular may hold the key to survival of the species.

The kākāpō shares its island homes (saving the kākāpō involved moving birds to rat-free islands) with a handful of tropical conifers from the families Podocarpaceae and Araucariaceae. Of these tropical conifers, one species is of particular interest to those concerned with kākāpō breeding - the rimu. Known to science as Dacrydium cupressinum, this evergreen tree represents one of the most important food sources for breeding kākāpō. Before we get to that, however, it is worth getting to know the rimu a bit better.


Rimu are remarkable, albeit slow-growing trees. They are endemic to New Zealand where they make up a considerable portion of the forest canopy. Like many slow-growing species, rimu can live for quite a long time. Before commercial logging moved in, trees of 800 to 900 years of age were not unheard of. Also, they can reach immense sizes. Historical accounts speak of trees that reached 200 ft. (61 m) in height. Today you are more likely to encounter trees in the 60 to 100 ft. (20 to 35 m) range.

The rimu is a dioecious tree, meaning individuals are either male or female. Rimu rely on wind for pollination and female cones can take upwards of 15 months to fully mature following pollination. The rimu is yet another one of those conifers that has converged on fruit-like structures for seed dispersal. As the female cones mature, the scales gradually begin to swell and turn red. Once fully ripened, the fleshy red “fruit” displays one or two black seeds at the tip. Its these “fruits” that have kākāpō researchers so excited.


As mentioned, it is a common observation that kākāpō only tend to breed when trees like the rimu experience reproductive booms. The “fruits” and seeds they produce are an important component of the diets of not only female kākāpō but their developing chicks as well. Because kākāpō are critically endangered, captive breeding is one of the main ways in which conservationists are supplementing numbers in the wild. The problem with breeding kakapo in captivity is that supplemental food doesn’t seem to bring them into proper breeding condition. This is where the rimu “fruits” come in.

Breeding birds desperately need calcium and vitamin D for proper egg production. As such, they seek out diets high in these nutrients. When researchers took a closer look at the “fruits” of the rimu, the kākāpō’s reliance on these trees made a whole lot more sense. It turns out, those fleshy scales surrounding rimu seeds are exceptionally high in not only calcium, but various forms of vitamin D once thought to be produced by animals alone. The nutritional quality of these “fruits” provides a wonderful explanation for why kākāpō reproduction seems to be tied to rimu reproduction. Females can gorge themselves on the “fruits,” which brings them into breeding condition. They also go on to feed these “fruits” to their developing chicks. For a slow growing, flightless parrot, it seems that it only makes sense to breed when food is this food source is abundant.


Though far from a smoking gun, researchers believe that the rimu is the missing piece of the puzzle in captive kākāpō breeding. If these “fruits” really are the trigger needed to bring female kākāpō into good shape for breeding and raising chicks, this may make breeding kākāpō in captivity that much easier. Captive breeding is the key to the long term survival of these odd yet charismatic, flightless parrots. By ensuring the production and survival of future generations of kākāpō, conservationists may be able to turn this tragedy into a real success story. What’s more, this research underscores the importance of understanding the ecology of the organisms we are desperately trying to save.

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

Further Reading: [1] [2]

Gymnosperms and Fleshy "Fruits"

 Fleshy red aril surrounding the seeds of  Taxus baccata.

Fleshy red aril surrounding the seeds of Taxus baccata.

Many of us were taught in school that one of the key distinguishing features between gymnosperms and angiosperms is the production of fruit. Fruit, by definition, is a structure formed from the ovary of a flowering plant. Gymnosperms, on the other hand, do not enclose their ovules in ovaries. Instead, their unfertilized ovules are exposed (to one degree or another) to the environment. The word “gymnosperm” reflects this as it is Greek for “naked seed.” However, as is the case with all things biological, there are exceptions to nearly every rule. There are gymnosperms on this planet that produce structures that function quite similar to fruits.

 Cross section of a  Ginkgo  ovule with red arrow showing the integument.

Cross section of a Ginkgo ovule with red arrow showing the integument.

The key to understanding this evolutionary convergence lies in understanding the benefits of fruits in the first place. Fruits are all about packing seeds into structures that appeal to the palates of various types of animals who then eat said fruits. Once consumed, the animals digest the fruity bits and will often deposit the seeds elsewhere in their feces. Propagule dispersal is key to the success of plants as it allows them to not only to complete their reproductive cycle but also conquer new territory in the process. With a basic introduction out of the way, let’s get back to gymnosperms.

 “Fruits” of  Cephalotaxus fortunei  (Cephalotaxaceae)

“Fruits” of Cephalotaxus fortunei (Cephalotaxaceae)

There are 4 major gymnosperm lineages on this planet - the Ginkgo, cycads, gnetophytes, and conifers. Each one of these groups contains members that produce fleshy structures around their seeds. However, their “fruits” do not all develop in the same way. The most remarkable thing to me is that, from a developmental standpoint, each lineage has evolved its own pathway for “fruit” production.

  Ginkgo  “fruits” are full of butyric acid and smell like rotting butter or vomit.

Ginkgo “fruits” are full of butyric acid and smell like rotting butter or vomit.

For instance, consider ginkgos and cycads. Both of these groups can trace their evolutionary history back to the early Permian, some 270 - 280 million years ago, long before flowering plants came onto the scene. Both surround their developing seed with a layer of protective tissue called the integument. As the seed develops, the integument swells and becomes quite fleshy. In the case of Ginkgo, the integument is rich in a compound called butyric acid, which give them their characteristic rotten butter smell. No one can say for sure who this nasty odor originally evolved to attract but it likely has something to do with seed dispersal. Modern day carnivores seem to be especially fond of Ginkgo “fruits,” which would suggest that some bygone carnivore may have been the main seed disperser for these trees.

 “Fruits” contained within the female cone of a cycad ( Lepidozamia peroffskyana ).

“Fruits” contained within the female cone of a cycad (Lepidozamia peroffskyana).

The Gnetophytes are represented by three extant lineages (Gnetaceae, Welwitschiaceae, and Ephedraceae), but only two of them - Gnetaceae and Ephedraceae - produce fruit-like structures. As if the overall appearance of the various Gnetum species didn’t make you question your assumptions of what a gymnosperm should look like, its seeds certainly will. They are downright berry-like!

 Berry-like seeds of  Gnetum gnemon .

Berry-like seeds of Gnetum gnemon.

The formation of the fruit-like structure surrounding each seed can be traced back to tiny bracts at the base of the ovule. After fertilization, these bracts grow up and around the seed and swell to become red and fleshy. As you can imagine, Gnetum “fruits” are a real hit with animals. In the case of some Ephedra, the “fruit” is also derived from much larger bracts that surround the ovule. These bracts are more leaf-like at the start than those of their Gnetum cousins but their development and function is much the same.

 Red, fleshy bracts of  Ephedra distachya .

Red, fleshy bracts of Ephedra distachya.

Whereas we usually think of woody cones when we think of conifers, there are many species within this lineage that also have converged on fleshy structures surrounding their seeds. Probably the most famous and widely recognized example of this can be seen in the yews (Taxus spp.). Ovules are presented singly and each is subtended by a small stalk called a peduncle. Once fertilized, a group of cells on the peduncle begin to grow and differentiate. They gradually swell and engulf the seed, forming a bright red, fleshy structure called an “aril.” Arils are magnificent seed dispersal devices as birds absolutely relish them. The seed within is quite toxic so it usually escapes the process unharmed and with any luck is deposited far away from the parent plant.

 The berry-like cones of  Juniperus communis .

The berry-like cones of Juniperus communis.

Another great example of fleshy conifer “fruits” can be seen in the junipers (Juniperus spp.). Unlike the other gymnosperms mentioned here, the junipers do produce cones. However, unlike pine cones, the scales of juniper cones do not open to release the seeds inside. Instead, they swell shut and each scale becomes quite fleshy. Juniper cones aren’t red like we have seen in other lineages but they certainly garnish the attention of many a small animal looking for food.

I have only begun to scratch the surface of the fruit-like structures in gymnosperms. There is plenty of literary fodder out there for those of you who love to read about developmental biology and evolution. It is a fascinating world to uncover. More importantly, I think the fleshy “fruits” of the various gymnosperm lineages stand as a testament to the power of natural selection as a driving force for evolution on our planet. It is amazing that such distantly related plants have converged on similar seed dispersal mechanisms by so many different means.

Photo Credits: [1] [2] [3] [4] [5] [6] [7] [8]

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

Hydrostachyaceae: Enigmatic Rheophytes


Today I would like to introduce you to an enigmatic family of aquatic angiosperms called Hydrostachyaceae. Though they kind of look like strange aquatic ferns or perhaps even lycopods, they are actually strange flowering plants. To find them, you need to hang out around waterfalls and rapids in either Madagascar or southern Africa.

Hydrostachyaceae is made up of roughly 22 species. This is a poorly understood group of plants and there is always a chance that more species await discovery. The various members of Hydrostachyaceae all take on a similar appearance. For much of the year they exist as a set of feathery, fern-like leaves that grow surprisingly large and look quite delicate, especially considering the types of habitats in which they grow.


Their delicate appearance is deceptive. In fact, the feathery structure of their leaves is an adaptation to the waters in which they grow. These are plants that require fast moving, clean, fresh water. If they were to produce flat, unbroken leaves, the fast currents would quickly rip them to shreds. By producing long, feathery leaves, water simply flows right over them with minimal disturbance. However, their preferred habitats also make them extremely difficult to study. Hence we know very little about their ecology.

What we do know about these plants is that they need clean rock surfaces and clear water for germination and subsequent growth. Dump too much sediment in the stream and you can kiss these plants goodbye. When they dry season approaches and water levels begin to drop, these oddball plants go into flowering mode. To the best of my knowledge, nearly all members of this family are dioecious, meaning individual plants are either male or female. When it comes time to flower, each plant produces modest sized spikes densely packed with flower.


The spikes themselves sit up and above the water line, which is how this family and genus got its name. Hydrostachys is Greek and roughly translates to “water spike.” I have not been able to track down any solid information on what might be pollinating these blooms, however, given their small, dense nature, and the extreme places in which they live, my bet would be on wind.

The ecology of Hydrostachyaceae isn’t the only mystery about these plants. Their position on the tree of life has also been cause for confusion ever since they were discovered. Morphologically speaking, aquatic angiosperms can offer a lot of confusion to taxonomists. Like whales, the ancestors of aquatic angiosperms lived out their lives on land. Making the move back into water comes with a lot of extremely specialized adaptations that can cloud our morphological interpretations of things.


Some authors have put forth the idea that these plants belong to another family of highly derived aquatic angiosperms - the Podostemaceae. However, genetic analyses paint a much different story. When the Angiosperm Phylogeny Group got a hold of specimens, their molecular work suggested the Hydrostachyaceae were nestled in Cornales, somewhere near the Hydrangea family (Hydrangeaceae). Exactly where Hydrostachyaceae fits into this new classification is still up for debate but it just goes to show you how messy things can get when plant lineages return to water.

Sadly, like so many other plants, the various members of Hydrostachyaceae are under a lot of pressure in the wild. Basically anything that threatens the quality of streams and rivers is a threat to the ongoing survival of these species. Runoff pouring into water ways from agriculture and mining cloud up the water and bury available germination sites under layers of sediment. Things only get worse when hydroelectric projects are installed. The fate of these plants is unequivocally tied to water quality.


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

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

A Relictual Palm in the American Southwest


Scattered throughout hidden oases nestled in the southwest corner of North America grows a glorious species of palm known to science as Washingtonia filifera. This charismatic tree goes by a handful of common names such as the desert fan palm, petticoat palm, and California fan palm. No matter what you call it, there is no denying that this palm is both unique and important to this arid region.

Populations of the desert fan palm are few and far between, occurring in a few scattered locations throughout the Colorado and Mojave Deserts. This palm can’t grow just anywhere in these deserts either. Instead, its need for water restricts it to small oases where springs, streams, or a perched water table can keep them alive.

Fossil evidence from Wyoming suggests that the restricted distribution of this palm is a relatively recent occurrence. Though not without plenty of debate, our current understanding of the desert fan palm is that it could once be found growing throughout a significant portion of western North America but progressive drying has seen its numbers dwindle to the small pockets of trees we know today.

The good news is that, despite being on conservation lists for its rarity, the desert fan palm appears to be expanding its range ever so slightly. One major component of this range expansion has to do with human activity. The desert fan palm makes a gorgeous specimen plant for anyone looking to add a tropical feel to their landscape. As such, it has been used in plantings far outside of its current range. Some reports suggest that it is even becoming naturalized in places like Death Valley, Sonoran Mexico, and even as far away as Florida and Hawai’i.


Other aspects contributing to its recent range expansion are also attributable to human activity, though indirectly. For one, with human settlement comes agriculture, and with agriculture comes wells and other forms of irrigation. It is likely that the seeds of the desert fan palm can now find suitably wet areas for germination where they simply couldn’t before. Also, humans have done a great job at providing habitat for potential seed dispersers, especially in the form of coyotes and fruit-eating birds.

It’s not just an increase in seed dispersers that may be helping the desert fan palm. Pollinators may be playing a role in its expansion as well, though in a way that may seem a bit counterintuitive. With humans comes a whole slew of new plants in the area. This greatly adds to the floral resources available for insect pollinators like bees.


Historically it has been noted that bees, especially carpenter bees, tend to be rather aggressive with palm inflorescences as they gather pollen, which may actually reduce pollination success. It is possible that with so many new pollen sources on the landscape, carpenter bees are visiting palm flowers less often, which actually increases the amount of pollen available for fertilizing palm ovules. This means that the palms could be setting more seed than ever before. Far more work will be needed before this mechanism can be confirmed.

Aside from its unique distribution, the desert fan palm has an amazing ecology. Capable of reaching heights of 80 ft. (25 m) or more and decked out in a skirt of dead fronds, the desert fan palm is a colossus in the context of such arid landscapes. It goes without saying that such massive trees living in desert environments are going to attract their fair share of attention. The thick skirt of dead leaves that cloaks their trunks serve as vital refuges for everything from bats and birds, to reptiles and countless of insects. Fibers from its leaves are often used to build nests and line dens.


And don’t forget the fruit! Desert fan palms can produce copious amount of hard fruits in good years. These fruits go on to feed many animals. Coupled with the fact that the desert fan palm always grows near a water source and you can begin to see why these palms are a cornerstone of desert oases. There has been some concern over the introduction of an invasive red palm weevil (Rhynchophorus ferrugineus), however, researchers were able to demonstrate that the desert fan palm has a trick up its sleeve (leaf skirt?) for dealing with these pests.

It turns out that desert fan palms are able to kill off any of these weevils as they try to burrow into its trunk. The desert fan palm secretes a gummy resin into damaged areas, which effectively dissuaded most adults and killed off developing beetle larvae. For now it seems that resistance is enough to protect this palm from this weevil scourge.

It is safe to say that regardless of its limited distribution, the desert fan palm is one tough plant. Its towering trunks and large, fan-like leaves stand as a testament to the wonderful ways in which natural selection shapes organisms. It is a survivor and one that has benefited a bit from our obsession with cultivating palms.

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

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

Are Packrats Fumigating Their Homes Using Plants?


Any organism that lives in one place for a long enough time is going to have to deal with pests. For mammals, this often means fleas and ticks. Nests, dens, and other roosting spots tend to accumulate high numbers of these blood suckers the longer they are in use. As such, anything that can cut down on pest loads in and around the home has the potential to confer great advantages. Evidence from California suggests that wood rats may be using the leaves of a shrub to do just that.

Dusky-footed wood rats (A.K.A. packrats) build giant nests out of twigs and other plant debris. These nests serve to protect packrats from both the elements and hungry predators. Packrat nests can last for quite a long time and reach monumental proportions considering the size of the rat itself. Because they use these stick nests for long periods of time, it should come as no surprise that they can build up quite a pest load. Fleas are especially problematic for these rodents.

 A dusky-footed woodrat ( Neotoma fuscipes ) and its den.

A dusky-footed woodrat (Neotoma fuscipes) and its den.

When researchers took a closer look at what packrats were bringing into their nests, they realized that not all plant material was treated equally. Whereas packrats actively collect and feed on leaves from various oaks (Quercus spp.), conifers (Pinus spp., Juniperus spp., etc.), and toyon (Heteromeles arbutifolia), the packrats seemed to have a special affinity for the leaves of the California bay (Umbellularia californica). However, instead of taking huge bites out of bay leaves, the rats appear to nibble them along the margin and spread them throughout their nest. What’s more, fresh bay leaves are brought in every few days.

This led some researchers to suggest that, instead of packrats using bay leaves as food, they may be using them to fumigate their homes. Indeed, California bay is rather chemically active. It is an aromatic shrub noted for its resistance to insect infestation. Of special interest to the research team were a group of chemical compounds called monoterpenoids. They noted that bay leaves were especially high in two types of of these compounds - 1,8-cineole (which gives the shrub its characteristic odor), and umbellulone (which has shown to be quite toxic to rodents). Why else would packrats bring something potentially deadly into their home other than to drive off pests?

chems bay.jpg

Closer observation revealed that the packrats were in fact treating bay leaves differently than other leaves. For starters, bay leaves were disproportionately used to line the sleeping chambers within the stick nests. What’s more, the bay leaves were cycled out every 2 to 3 days. Even the nibbling patterns were significantly different. As mentioned above, bay leaves were merely nibbled along the leaf margins, which is an ideal place to nibble if releasing volatile compounds is the desired effect.

When researchers tested the effectiveness of a variety of leaves in the lab, their results added further evidence to the fumigation hypothesis. More than any other leaf found in packrat nests, bay leaves had clear negative effects on flea numbers. Flea survival in the lab was reduced by upwards of 75% when California bay leaves were present whereas flea survival was only reduced by less than 10% with all other leaves. It goes without saying that, whether they are conscious decisions or not, packrats definitely stand to benefit by decorating their homes with California bay leaves.

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

Further Reading: [1]

Arctic Vegetation is Growing Taller & Why That Matters


The Arctic ecosystem is changing and it is doing so at an alarming rate. Indeed, the Arctic Circle is warming faster than most other ecosystems on this planet. All of this change has implications for the plant communities that call this region home. In a landmark study that incorporated thousands of data points from places like Alaska, Canada, Iceland, Scandinavia, and Russia, researchers have demonstrated that Arctic vegetation is, on average, getting taller.

Imagine what it is like to be a plant growing in the Arctic. Extreme winds, low temperatures, a short growing season, and plenty of snow are just some of the hardships that characterize life on the tundra. Such harsh conditions have shaped the plants of this region into what we know and love today. Arctic plants tend to hug the ground, hunkering down behind whatever nook or cranny offers the most respite from their surroundings. As such, plants of Arctic-type habitats tend to be pretty small in stature. As you can probably imagine, if these limits to plant growth become less severe, plants will respond accordingly.


That is part of what makes this new paper so alarming. The vegetation that comprise these Arctic communities is nearly twice as tall today as it was 30 years ago. However, the increase in height is not because the plants that currently grow there are getting taller but rather because new plants are moving northwards into these Arctic regions. New players in the system are usually cause for concern. Other studies have shown that it isn’t warming necessarily that hurts Arctic and alpine plants but rather competition. They simply cannot compete as well with more aggressive plant species from lower latitudes.

Taller plants moving into the Arctic may have even larger consequences than just changes in species interactions. It can also change ecosystem processes, however, this is much harder to predict. One possible consequence of taller plants invading the Arctic involves carbon storage. It is possible that as conditions continue to favor taller and more woody vegetation, there could actually be more carbon storage in this system. Woody tissues tend to sequester more carbon and shading from taller vegetation may slow decomposition rates of debris in and around the soil.

  Alopecurus alpinus  is one of the new tall plant species moving into the Arctic

Alopecurus alpinus is one of the new tall plant species moving into the Arctic

It is also possible that taller vegetation will alter snowpack, which is vital to the health and function of life in the Arctic. Taller plants with more leaf area could result in a reduced albedo in the surrounding area. Lowering the albedo means increased soil temperatures and reduced snowpack as a result. Alternatively, taller plants could also increase the amount of snowpack thanks to snow piling up among branches and leaves. This could very well lead (counterintuitively) to warmer soils and higher decomposition rates as snowpack acts like an insulating blanket, keeping the soil slightly above freezing throughout most of the winter.

It is difficult to make predictions on how a system is going to respond to massive changes in the average conditions. However, studies looking at how vegetation communities are responding to changes in their environment offer us one of the best windows we have into how ecosystems might change moving into the uncertain future we are creating for ourselves.

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

Further Reading: [1]

The Upside Down World of Orchid Flowers


Did you know that most orchid flowers you see are actually blooming upside down? That's right, referred to as "resupination," the lower lip of many orchid flowers is actually the top petal and, as the flower develops inside the bud, the whole structure makes a 180° rotation. How and why does this happen?

The lip of an orchid flower usually serves to attract pollinators as well as function as a landing pad for them. The flower of an orchid is an incredibly complex organ with an intriguing evolutionary history. Basically, the lip is the most derived structure on the flower and, in most cases, it is the most important structure in initiating pollination.

 The non-resupinate flowers of the grass pink ( Calopogon tuberosus ) showing the lip on top.

The non-resupinate flowers of the grass pink (Calopogon tuberosus) showing the lip on top.

As an orchid flower bud develops, it begins to exhibit gravitropic tendencies, meaning it responds to the pull of gravity. By removing specific floral organs like the column and pollinia, researchers found that they produce special hormones called auxins that tell the developing bud to begin the process of resupination. The ovary starts to twist, causing the flower to stand on its head.

Not all orchids exhibit resupinate flowers. Grass pinks (Calopogon tuberosus) famously bloom with the lip pointing up as it does in the early stages of bud development. It is an interesting mechanism and serves to demonstrate the stepwise tendencies that the forces of natural selection and evolution can manifest. But why does it occur at all? What is the evolutionary advantage of resupinate flowers?

 Not only are  Dracula  flowers resupinate, many species also face them towards the ground.

Not only are Dracula flowers resupinate, many species also face them towards the ground.

The most likely answer to this biological twist is that, for orchids, resupination places the lip in such a way that facilitates pollination by whatever the flowers are attracting. For many orchids, this means providing an elaborate landing strip in the form of the lip. For the grass pinks, which operate by slamming visiting bees downward onto the column to achieve pollination, placing the lip at the top makes more mechanical sense. When a bee visits the upward pointing lip thinking it will find a pollen-rich meal, the lip bend at the base like a hinge. Anything goes in evolution provided the genes are present for selection to act upon and nowhere is this fact more beautifully illustrated than in orchids.

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

Viper's Bugloss


Throughout much of North America, brown fields, roadsides, and other waste places occasionally take on a wonderful blue hue. Often time the cause of this colorful display is none other than Echium vulgare, or as its commonly referred to, viper's bugloss. Viper’s bugloss is a member of the borage family and was originally native to most of Europe and Asia. However, humans introduced it to North America some time ago. It has since naturalized quite well and is even considered invasive in parts of Washington. No matter your views on this plant, the reproductive ecology of this species is quite interesting.

Viper's bugloss produces its flowers on spikes. Starting off pink and gradually changing to blue as they mature, the flowers ripen their male portions on their first day and ripen their female portions on the second day. This is known as "protandry." Plants that exhibit this lifestyle offer researchers a window into the advantages and disadvantages with regards to the fitness investment of each sex. What they have found in viper's bugloss is that there are clearly distinct strategies for each type of flower.

Male flowers are pollinator limited. They must hedge their bets towards increasing the number of visitors. Bees are the main pollinators of this species and the more bees that visit, the more pollen can be disseminated. Unlike female flowers, which are resource limited, male flowers can produce pollen and nectar quite cheaply. Because of this, male flowers produce significantly more nectar than female flowers to bring in more bees. As the anthers senesce and give way to receptive styles, things begin to change. The plant now has to redirect resources into producing seed. At this point, resources are everything. The plant produces considerably less nectar resources than pollen but the bees can’t know that without visiting.


The other interesting aspect its reproductive ecology has to do with population size. Bees are notorious for favoring plants that are more numerous on the landscape. This makes a lot of sense. Why spend time looking for uncommon plants when they can go for easier, more numerous targets. This can be very detrimental to the fitness of rare plant species. However, plants like viper's bugloss don't seem to fall victim to this.

By looking at large and small populations, researchers found that pollination success pretty much evens out for viper's bugloss no matter how numerous it is in a given area. Large populations receive many more visits from bees but the bees spend less time on each flower. When viper's bugloss populations are small, flowers receive fewer visits but bees spend more time at each flower. This results is no significant difference in the reproductive fitness of either population.

Considering how efficient this plant is reproductively, it is no wonder it has done so well outside of its native range. Add to this its ability to grow in some of the worst soil conditions, it goes without saying that viper's bugloss is here to stay. If you find this species growing, certainly take time to get up close with the flowers. You will be glad you did.

Photo Credits: [1] [2]

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

The Genus Ceropegia Recently Got a Whole Lot Bigger


The succulent and climbing members of the milkweed family (Apocynaceae) have been gaining a lot of popularity among houseplant growers and for good reason. These wonderful plants produce some of the most elaborate flowers most of us will ever encounter and many of them smell quite strongly. Whereas houseplant enthusiasts recognize multiple genera of these spectacular plants, recent taxonomic work suggests lumping them all into one single genus - Ceropegia.

Such a massive taxonomic move has caused its fair share of drama. Folks seem to get quite ornery when it comes to shifts in nomenclature, especially when it involves this many species. However, when you dive into this group of plants, you really start to see how shaky the ground was that supported the previous classification systems. Evolution, after all, is not a neat and tidy process and we can learn a lot from the succulent and climbing asclepiads regarding the importance of combining morphological and genetic data into our taxonomic decisions.

  Ceropegia (Stapelia) hirsuta

Ceropegia (Stapelia) hirsuta

Botanists have been obsessing over this group for decades. Historically speaking, four major groups have been recognized: those with caudiciform stems (genus Brachystelma), the stem succulent stapeliads (which include genera such as Stapelia, Huernia, Orbea, Caralluma, and others), the climbers (genus Ceropegia), and the so-called early divergent group (which include genera such as Anisotoma, Conomitra, Dittoceras, and others). Together these groups total something like 762 species and represent the tribe Ceropegieae.

The taxonomic status of the various members of Ceropegieae have always been up for debate. Early work was based on surprisingly few species and relied heavily on morphological characters such and corolla shape, stem anatomy, and pubescence. Since the 1950’s, many more species have been discovered and that is where a lot of the trouble began. Much of the early characters that were used to draw lines between various groups were suddenly blurred. Genera were created and absorbed by various authors in an attempt to get a handle on how this tribe evolved.

  Ceropegia  ( Brachystelma )  tuberosum

Ceropegia (Brachystelma) tuberosum

Things got even more complicated as various stapeliads and Ceropegia attracted the attention of horticulturists. As new species became available, many varieties were haphazardly named and genera such as Stapelia were further split to accommodate some of the peculiar nuances in floral shapes, colors, and sizes. It wasn’t until some genetic work was done that the need for a major overhaul of the Ceropegieae tribe became apparent.

Unfortunately, this early molecular work suffered from low resolution. Very few genera were used and among those, only a handful of gene regions were analyzed. Still, the picture that was developing was that the historical understanding of Ceropegieae was surprisingly misleading. For instance, the genera that made up the stapeliad group appeared to be nested quite firmly within the genus Ceropegia. Though equally as limited in scope, consecutive work in the early 2000’s added further evidence to the idea that the four groups that made up Ceropegieae were so genetically similar that most should be nested somewhere within Ceropegia.

  Ceropegia  ( Duvalia )  modesta

Ceropegia (Duvalia) modesta

Though not without controversy, this early molecular work convinced enough taxonomists to take a closer look at each of the four groups. With more resolution and a finer grasp on the diversity in form of these plants, taxonomists started to question the validity of some taxa. Indeed, the closer anyone looked, the more the lines between genera started to blur.

For example, Ceropegia and Brachystelma have long been separated on the basis of floral structure. Ceropegia were considered to adhere to a single corolla structure involving long, tubular flowers whereas Brachystelma were thought to be more variable in form. The discovery of new species clearly demonstrates that there are far too many exceptions to this system for it to be valid.

 Fig. 1. Variation in the corolla and corona in the traditional concept of  Ceropegia : A–C,  C. salicifolia , Nepal,  Bruyns 2507  (BM, K); D–E,  C. melanops , Ethiopia,  Gilbert 3050  (K); F—H,  C. meleagris , Nepal,  Bruyns 2496  (K); I–J,  C. loranthiflora , Ethiopia,  Gilbert 2851   (K). [scale-bars or subdivisions indicate mm; A, D, F, I, corolla from  side; B, G, corolla dissected to show location of corona; C, E, H, J,  corona from side].   [SOURCE]

Fig. 1. Variation in the corolla and corona in the traditional concept of Ceropegia: A–C, C. salicifolia, Nepal, Bruyns 2507 (BM, K); D–E, C. melanops, Ethiopia, Gilbert 3050 (K); F—H, C. meleagris, Nepal, Bruyns 2496 (K); I–J, C. loranthiflora, Ethiopia, Gilbert 2851 (K). [scale-bars or subdivisions indicate mm; A, D, F, I, corolla from side; B, G, corolla dissected to show location of corona; C, E, H, J, corona from side]. [SOURCE]

 Fig. 2. Variation in the corolla and corona in the traditional concept of  Brachystelma : A–C,  B. brevipedicellatum , South Africa,  Bruyns 2372 ; D–F,  B. mafekingense , Namibia,  Bruyns 1954  (K, WIND); G–J,  B. gymnopodum , South Africa,  Bruyns 2078   (NBG). [scale-bars or subdivisions indicate mm; A, corolla from front,  D, G, corolla from side; B, E, H, corolla dissected to show location of  corona; C, J, corona from front; F, I, corona from side].   [SOURCE]

Fig. 2. Variation in the corolla and corona in the traditional concept of Brachystelma: A–C, B. brevipedicellatum, South Africa, Bruyns 2372; D–F, B. mafekingense, Namibia, Bruyns 1954 (K, WIND); G–J, B. gymnopodum, South Africa, Bruyns 2078 (NBG). [scale-bars or subdivisions indicate mm; A, corolla from front, D, G, corolla from side; B, E, H, corolla dissected to show location of corona; C, J, corona from front; F, I, corona from side]. [SOURCE]

Such is also the case for other anatomical features such as whether plants climb or not. Again, there are plants in both genera that deviate from these patterns, thus making it impossible to nail down any set of characters that maintain the split between these two genera. Also, it would seem that some authors were trying to pull a fast one on readers. Back in 2007, Meve and Liede-Schumann claimed there were “a wide array of morphological features” that separate these two genera but failed to reveal any but those mentioned here. There are multiple species of Ceropegia and Brachystelma that simply do not conform to this historical classification.

Similarly, Ceropegia and the various stapeliads have been separated on the basis of stem and floral anatomy. Historically speaking, the stapeliads were thought to consist of fleshy, succulent stems with tubercules and reduced or absent leaves, whereas Ceropegia were considered to be slender climbers. Again, with more species having been discovered, these distinctions grew more and more blurry.

 The succulent stems of  Ceropegia cimiciodora .

The succulent stems of Ceropegia cimiciodora.

It turns out that there are many Ceropegia with fleshy, succulent stems and the only major difference between the two genera is the lack of angles in the stems of some Ceropegia. The structure and presentation of their flowers also stands on shaky ground. There is so much similarity between the flowers of some of the succulent Ceropegia and the early diverging stapeliads that one would be hard pressed to identify any character that clearly separates them.

Between all of the molecular work and the anatomical scrutiny, it was clear that something needed to be done to clean up the taxonomic status of Ceropegieae. Keeping things separate may make sense to some but considering the group as a whole instead of from a collector’s standpoint, trying to find enough distinct characters to preserve the historical treatment would make things way too messy. In 2017 it was suggested that because there are no clear differences between the four groups within this tribe, all members were to be lumped back in to the genus Ceropegia.

  Ceropegia  ( Stapelia )  flavopurpurea

Ceropegia (Stapelia) flavopurpurea

Although this most recent treatment still recognizes some morphological differences between these plants (thus multiple subsections are recognized), the lack of genetic differentiation between groups long thought to be distinct really does support this decision. Because of historical precedents, Ceropegia won out as the main generic classification.

Personally I find this work to be extremely exciting. It involved a lot of wonderful detective work and a whole lot of attention to detail. I think the end result paints a far better picture for our understanding of how these plants evolved. I am especially floored that some of the earlier morphological notes turned out to be quite useful in this modern understanding. Even more exciting is the fact that now we know that many of what we thought were “unique” characters amoung the various species actually evolved multiple times throughout the history of this group. This is why I will never get upset by taxonomic changes. They may be working documents but each step we take helps us understand evolution that much more.

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

Further Reading: [1]

The Wild World of Rattan Palms


There are a lot of big organisms out there. A small handful of these are truly massive. When someone mentions big plants, minds will quickly drift to giant sequoias or coastal redwoods. These species are indeed massive. The tallest tree on record is a coastal redwood measuring 369 feet tall. That's a whole lot of tree! What some may not realize is that there are other plants out there that can grow much "taller" than even the tallest redwood. For instance, there is a group of palms that hail from Africa, Asia, and Australasia that grow to staggering lengths albeit without the mass of a redwood.

You are probably quite familiar with some of these palm species, though not as living specimens. If you have ever owned or sat upon a piece of wicker furniture then you were sitting on pieces of a rattan palm. Rattan palms do not grow in typical palm tree fashion. Rattans are climbers, more like vines. All palms grow from a central part of the plant called the heart. They grow as bromeliads do, from meristem tissue in the center of a rosette of leaves. As a rattan grows, its stem lengthens and grabs hold of the surrounding vegetation using some seriously sharp, hooked spikes. For much of their early life they generally sprawl across the forest floor but the real goal of the rattan is to reach up into the canopy where they can access the best sunlight.


Rattans are not a single taxonomic unit. Though they are all palms, at least 13 genera contain palms that exhibit this climbing habit. With over 600 species included in these groups, it goes without saying that there is a lot of variation on the theme. The largest rattan palms hail from the genus Calamus and all but one are native to Asia.

Many species of rattan have whip-like stems that would be easy to miss in a lush jungle. Be aware of your surroundings though, because these spikes are quite capable of ripping clothes and flesh to pieces. The rattans are like any other vine, sacrificing bulk for an easy ride into the light at the expense of whatever it climbs on. Indeed some get so big that they break their host tree. It is this searching, sprawling nature of the rattans that allow them to reach some impressive lengths. Some species of rattan have been reported with stems measuring over 500 feet!


Getting back to what I mentioned earlier about wicker furniture, rattans are a very important resource for the people of the jungles in which they grow. They offer food, building materials, shelter materials, an artistic medium, and a source of economic gain. In many areas, rattans are being heavily exploited as a result. This is bad for both the ecology of the forest and the locals who depend upon these species.

The global rattan trade is estimated at around $4 billion dollars. Because of this, rattans are harvested quite heavily and many are cut at too young of an age to re-sprout meaning little to no recruitment occurs in these areas. There is a lot of work being done by a few organizations to try to set up sustainable rattan markets in the regions that have been hit the hardest. More information can be found at sites like the World Wildlife Fund.

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

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

The Only True Cedars

  Cedrus deodara

Cedrus deodara

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

 J. White,1803-1824.

J. White,1803-1824.

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

  Cedrus libani

Cedrus libani

  Cedrus libani

Cedrus libani

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

 Male cones of  Cedrus atlantica

Male cones of Cedrus atlantica

 Female  Cedrus  cones.

Female Cedrus cones.

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

  Cedrus atlantica

Cedrus atlantica

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

Photo Credit: Wikimedia Commons

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