Meet the Fire Lily

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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.

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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.

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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]

Meet the Pygmy Clubmoss

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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.

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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.

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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.

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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]

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]

Arctic Vegetation is Growing Taller & Why That Matters

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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.

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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]

Getting to Know Sansevieria

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The houseplant hobby is experiencing something of a renaissance as of late. With their popularity on various social media platforms, easy to grow plant species and their cultivars are experiencing a level of popularity they haven't seen in decades. One genus of particular interest to houseplant hobbyists is Sansevieria.

Despite their popularity, the few Sansevieria species regularly found in cultivation come attached with less than appealing common names. Mother-in-law's tongue, Devil's tongue, and snake plant all carry with them an air of negativity for what are essentially some of the most forgiving houseplants on the market. What few houseplant growers realize is that those dense clumps of upright striped leaves tucked into a dark corner of their home belong to a fascinating genus worthy of our admiration. What follows is a brief introduction to these enigmatic houseplants.

  Sansevieria cylindrica

Sansevieria cylindrica

  Sansevieria ballyi

Sansevieria ballyi

The Sansevieria we encounter in most nurseries are just the tip of the iceberg. Sansevieria is a genus comprise of about 70 different species. I say 'about' because this group is a taxonomic mess. There are a couple reasons for this. For starters, the vast majority of Sansevieria species are painfully slow growers. It can take decades for an individual to reach maturity. As such, they have never really presented nursery owners with much in the way of economic gain and thus only a few have received any commercial attention.

Another reason has to do with the fiber market during and after World War II. In hopes of discovering new plant-based fibers for rope and netting, the USDA collected many Sansevieria but never formally described most of them. Instead, plants were assigned numbers in hopes that future botanists would take the time needed to parse them out properly.

A third reason has to do with the variety of forms and colors these plants can take. Horticulturists have been fond of giving plants their own special cultivar names. This complicates matters as it is hard to say which names apply to which species. Often the same species can have different names depending on who popularized it and when.

  Sansevieria grandis in situ .

Sansevieria grandis in situ.

Regardless of what we call them, all Sansevieria hail from arid regions of Africa, Madagascar and southern Asia. In the wild, many species resemble agave or yucca and, indeed, they occupy similar niches to these New World groups. Like so many other plants of arid regions, Sansevieria evolved CAM photosynthesis as a means of coping with heat and drought. Instead of opening up their stomata during the day when high temperatures would cause them to lose precious water, they open them at night and store CO2 in the form of an organic acid. When the sun rises the next day, the plants close up their stomata and utilize the acid-stored carbon for their photosynthetic needs.

 The wonderfully compact  Sansevieria pinguicula .

The wonderfully compact Sansevieria pinguicula.

Often you will encounter clumps of Sansevieria growing under the dappled shade of a larger tree or shrub. Some even make it into forest habitats. Most if not all species are long lived plants, living multiple decades under the right conditions. These are just some of the reasons that they make such hardy houseplants.

The various Sansevieria appear the sort themselves out along a handful of different growth forms. The most familiar to your average houseplant enthusiast is the form typified by Sansevieria trifasciata. These plants produce long, narrow, sword shaped leaves that point directly towards the sky. Many other Sansevieria species, such as S. subspicata and S. ballyi, take on a more rosetted form with leaves that span the gamut from thin to extremely succulent. Still others, like S. grandis and S. forskaalii, produce much larger, flattened leaves that grow in a form reminiscent of a leaky vase. 

  Sansevieria trifasciata  with berries .

Sansevieria trifasciata with berries.

Regardless of their growth form, a majority of Sansevieria species undergo radical transformations as they age. Because of this, adults and juveniles can look markedly different from one another, a fact that I suspect lends to some of the taxonomic confusion mentioned earlier. A species that illustrates this nicely is S. fischeri. When young, S. fischeri consists of tight rosettes of thick, mottled leaves. For years these plants continue to grow like this, reaching surprisingly large sizes. Then the plants hit maturity. At that point, the plant switches from its rosette form to producing single leaves that protrude straight out of the ground and can reach heights of several feet! Because the rosettes eventually rot away, there is often no sign of the plants previous form.

 A young  Sansevieria fischeri  exhibiting its rosette form.

A young Sansevieria fischeri exhibiting its rosette form.

 A mature  Sansevieria fischeri  with its large, upright, cylindrical leaves.

A mature Sansevieria fischeri with its large, upright, cylindrical leaves.

If patient, many of the Sansevieria will reach enormous sizes. Such growth is rarely observed as slow growth rates and poor housing conditions hamper their performance. It's probably okay too, considering the fact that, when fully grown, such specimens would be extremely difficult to manage in a home. If you are lucky, however, your plants may flower. And flower they do!

Though there is variation among the various species, Sansevieria all form flowers on either a simple or branched raceme. Flowers range in color from greenish white to nearly brown and all produce a copious amount of nectar. I have even noticed sickeningly sweet odors emanating from the flowers of some captive specimens. After pollination, flowers give way to brightly colored berries, hinting at their place in the family Asparagaceae.

 A flowering  Sansevieria hallii .

A flowering Sansevieria hallii.

As a whole, Sansevieria can be seen as exceptional tolerators, eking out an existence wherever the right microclimate presents itself in an otherwise harsh landscape. Their extreme water efficiency, tolerance of shade, and long lived habit has lent to the global popularity of only a few species. For the majority of the 70 or so species in this genus, their painfully slow growth rates means that they have never made quite a splash in the horticulture trade.

Nonetheless, Sansevieria is one genus that even the non-botanically minded among us can pick out of a lineup. Their popularity as houseplants may wax and wane but plants like S. trifasciata are here to stay. My hope is that all of these folks collecting houseplants right now will want to learn more about the plants they bring into their homes. They are more than just fancy decorations, they are living things, each with their own story to tell. 

Photo Credits: [1] [2] [3] [4] [5] [6] [7] [8] [9]

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

Meet the Blazing Stars

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Midsummer in North America is, among other things, Liatris season. These gorgeous plants are often referred to as blazing stars or gayfeathers, which hints at the impact their flowers have on our psyche. Whether in the garden or in the wild, Liatris are a group of plants worth getting to know a bit better.

Liatris is by and large a North American genus with only a single species occurring in the Bahamas. Though we often think of Liatris as prairie plants, the center of diversity for this group is in the southeastern United States. Taxonomically speaking, Liatris are a bit of a conundrum. Something like 40 different species have been described and, where ranges overlap, many putative hybrids have been named.

 Rocky Mountain blazing star ( Liatris ligulistylis )

Rocky Mountain blazing star (Liatris ligulistylis)

Authorities on this group cite ample confusion when it comes to drawing lines between species. Much of this confusion comes from the fact that numerous variants and intergradations exist between the various species. As mentioned, hybridization is not uncommon in this genus, which complicates matters quite a bit.

 Prairie blazing star ( Liatris pycnostachya )

Prairie blazing star (Liatris pycnostachya)

Liatris as a whole appears to have undergone quite an adaptive radiation in North America, with species adapting to specific soils and habitat types. Take, for instance, the case of cylindrical blazing star (L. cylindracea), marsh blazing star (L. spicata), and rough blazing star (L. aspera). The ranges of these species overlap to quite a degree, however, each prefers to grow in soils of specific texture and moisture. Marsh blazing star, as you may have guessed, prefers wetter soils whereas rough blazing star enjoys drier habitats. Cylindrical blazing star seems to enjoy intermediate soil conditions, especially where soil pH is a bit higher. As such, these three species often occur in completely different habitats. However, in places like the southern shores of Lake Michigan, they find themselves growing in close quarters and as a result, a fair amount of hybridization has occurred.

 Rough blazing star ( Liatris aspera )

Rough blazing star (Liatris aspera)

Another example of confusion comes from a species commonly known as the savanna blazing star (Liatris scariosa nieuwlandii). Many different ecotypes of this plant exist and some experts don't quite know how to deal with them all. Sometimes savanna blazing star is treated as a variant of another species called the northern blazing star (Liatris scariosa var. nieuwlandii) and sometimes it is treated as its own distinct species (Liatris nieuwlandii). Until proper genetic work can be done, it is impossible to say which, if any, are correct. 

 Glandular blazing star ( Liatris glandulosa )

Glandular blazing star (Liatris glandulosa)

Taxonomic confusion aside, the various Liatris species and variants are important components of the ecology wherever they occur. Numerous insects feed upon and raise their young on the foliage and few could argue against their flowers as pollinator magnets. All Liatris produce pink to purple flowers in splendid Asteraceae fashion. Every once in a while, an aberrant form is produced that sports white flowers. Though horticulturists have capitalized on this for the garden, at least one authority claims that these white forms are much weaker than their pink flowering parents. At least one species, the pinkscale blazing star (L. elegans), produces large, filamentous white bracts that very much resemble flowers.

 Check out the bracts on the pinkscale blazing star ( L. elegans )!

Check out the bracts on the pinkscale blazing star (L. elegans)!

Liatris are just as interesting below as they are above. The roots, foliage, and flowers all emerge from a swollen underground stem called a corm. The formation of these corms is one reason why some Liatris species have become so popular in our gardens. It makes them extremely hardy during the dormant season. In the spring, the corm starts forming roots. At the same time, tiny preformed buds at the top of the corm begin to grow this years crop of leaves and flowers. By the end of the growing season, the corm has reached its maximum size for that year and the plant draws down the rest of its reserves to wait out the winter.

 Cylindrical blazing star ( Liatris cylindracea )

Cylindrical blazing star (Liatris cylindracea)

During this time, some species form a layer of tissue along the edge of the corm that is much darker in coloration than what was laid down earlier in the season. This has led some to suggest that aging individual Liatris is possible. Experts believe that specimens can readily reach 30 to 40 years of age or more, however, the degree to which these dark bands indicate annual growth is up for a lot of debate. Others have found no correlation with plant age. Regardless, it is safe to say that many Liatris species can live for decades if left undisturbed.

 Scrub blazing star ( Liatris ohlingerae )

Scrub blazing star (Liatris ohlingerae)

All in all, Liatris is a very special, albeit slightly confusing, group of plants. It offers a little something for everyone. What's more, their beauty is only part of the story. These are ecologically important plants that support many great insect species. As summer wears on, make sure to get out there and enjoy the Liatris in your neck of the woods. You will be happy you did!

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

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

 

 

 

What Are Plants Made Of?

Have you ever thought about what plants are made of? I mean, really thought about it. Strip away all the splendor and glory of all the different plant species on this planet and really take a close look at how plants grow and make more plants. It is a fascinating realm and it all has to do with photosynthesis. To go from photons given off by our nearest star to a full grown plant is quite the journey and, at the end of that journey, you may be surprised to learn what plants are all about.

It starts with photons. Leaving the sun they travel out into the universe. Some eventually collide with Earth and make their way to the surface. Plants position their leaves to absorb these photons. Energy from the photons is used to split water molecules inside the chloroplasts. In the process of splitting water, oxygen is released as a byproduct (thanks plants!). Splitting water also releases electrons and hydrogen ions.

These electrons and hydrogen ions are used to make energy in the form of ATP. Along with some electrons, ATP is then used in another cycle known as the Calvin cycle. The point of the Calvin cycle is to take in CO2 and use the energy created prior to reduce carbon molecules into chains of organic molecules. Most of the carbon in a plant comes from the intake of CO2. Through a series of steps (I will spare you the details) plants piece together carbon atoms into long chains. Some of these chains form glucose and some of that glucose gets linked together into cellulose.

Cellulose is the main structural component of plant cells. From the smallest plants in the world (genus Wolffia) all the way up to the largest and tallest redwoods and sequoias (incidentally some of the largest organisms to have ever existed on this planet) , all of them are built out of cellulose. So, in essence, all the plant life you see out there is literally built from the ground up by carbon originating from CO2 gas. Pretty incredible stuff, wouldn't you agree?

Big Things Come In Small Packages

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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.

B. 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 rather 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.

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. 

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.

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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 has 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

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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).

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).

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 .

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)

The inflorescence of Philodendron bipinnatifidum can reach temps as high as 115 °F (46 °C)

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.

The world's largest inflorescence belongs to the titan arum (Amorphophallus titanum) and it too produces heat.

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]

 

Of Bluebells and Fungi

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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.

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.

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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.

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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.

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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.

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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

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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. 

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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]

North America's Pachysandra

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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.

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] [3]

Further Reading: [1] [2]

Apocynaceae Ant House

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The dogbane family, Apocynaceae, comes in many shapes, sizes, and lifestyles. From the open-field milkweeds we are most familiar with here in North America to the cactus-like Stapeliads of South Africa, it would seem that there is no end to the adaptive abilities of this family. Being an avid gardener both indoors and out, the diversity of Apocynaceae means that I can be surrounded by these plants year round. My endless quest to grow new and interesting houseplants was how I first came to know a genus within the family that I find quite fascinating. Today I would like to briefly introduce you to the Dischidia vines.

 Bullate leaves help the vine clasp to the tree as well as house ant colonies.

Bullate leaves help the vine clasp to the tree as well as house ant colonies.

The genus Dischidia is native to tropical regions of China. Like its sister genus Hoya, these plants grow as epiphytic vines throughout the canopy of warm, humid forests. Though they are known quite well among those who enjoy collecting horticultural curiosities, Dischidia as a whole is relatively understudied. These odd vines do not attach themselves to trees via spines, adhesive pads, or tendrils. Instead, they utilize their imbricated leaves to grasp the bark of the trunks and branches they live upon.

 The odd, bulb-like leaves of the urn vine ( Dischidia rafflesiana )

The odd, bulb-like leaves of the urn vine (Dischidia rafflesiana)

One thing we do know about this genus is that most species specialize in growing out of arboreal ant nests. Ant gardens, as they are referred to, offer a nutrient rich substrate for a variety of epiphytic plants around the world. What's more, the ants will visciously defend their nests and thus any plants growing within.

 The flowers of   Dischidia ovata

The flowers of Dischidia ovata

Some species of Dischidia take this relationship with ants to another level. A handful of species including D. rafflesiana, D. complex, D. major, and D. vidalii produce what are called "bullate leaves." These leaves start out like any other leaf but after a while the edges stop growing. This causes the middle of the leaf to swell up like a blister. The edges then curl over and form a hollow chamber with a small entrance hole.

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These leaves are ant domatia and ant colonies quickly set up shop within the chambers. This provides ample defense for the plant but the relationship goes a little deeper. The plants produce a series of roots that crisscross the inside of the leaf chamber. As ant detritus builds up inside, the roots begin to extract nutrients. This is highly beneficial for an epiphytic plant as nutrients are often in short supply up in the canopy. In effect, the ants are paying rent in return for a place to live.

Growing these plants can take some time but the payoff is worth. They are fascinating to observe and certainly offer quite a conversation piece as guests marvel at their strange form.

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

Further Reading: [1]

The Giant Genomes of Geophytes

 Canopy plant ( Paris japonica )

Canopy plant (Paris japonica)

A geophyte is any plant with a short, seasonal lifestyle and some form of underground storage organ ( bulb, tuber, thick rhizome, etc.). Plants hailing from a variety of families fall into this category. However, they share more than just a similar life history. A disproportionate amount of geophytic plants also possess massive genomes. 

As we have discussed in previous posts, life isn't easy for geophytes. Cold temperatures, a short growing season, and plenty of hungry herbivores represent countless hurdles that must be overcome. That is why many geophytes opt for rapid growth as soon as conditions are right. However, they don't do this via rapid cell division. 

 Dutchman's breeches ( Dicentra cucullaria ) emerging with preformed buds.

Dutchman's breeches (Dicentra cucullaria) emerging with preformed buds.

Instead, geophytes spend the "dormant" months pre-growing all of their organs. What's more, the cells that make up their leaves and flowers are generally much larger than cells found in non-geophytes. This is where that large genome comes into plant. If they had to wait until the first few weeks of spring to start their development, a large genome would only get in the way. Their dormant season growth means that these plants don't have to worry about streamlining the process of cellular division. They can take their time. 

As such, an accumulation of genetic material isn't detrimental. Instead, it may actually be quite beneficial for geophytes. Associated with large genomes are things like larger stomata, which helps these plants better regulate their water needs. The large genomes may very well be the reason that many geophytic plants are so good at taking advantage of such ephemeral growing conditions. 

When the right conditions present themselves, geophytes don't waste time. Pre-formed organs like leaves and flowers simply have to fill with water instead of having to wait for tissues to divide and differentiate. Water is plentiful during the spring so geophytes can rely on turgor pressure within their large cells for stability rather than investing in thick cell walls. That is why so many spring blooming plants feel so fleshy to the touch. 

Taken together, we can see how large genomes and a unique growth strategy have allowed these plants to exploit seasonally available habitats. It is worth noting, however, that this is far from the complete picture. With such a wide variety of plant species adopting a geophytic lifestyle, we still have a lot to learn about the secret lives of these plants.

Photo Credits: [1] [2]

Further Reading: [1]

An Ancient Hawaiian Moss

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The cloud forests of Kohala Mountain on the island of Hawai'i are home to a unique  botanical community. One plant in particular is quite special as it may be one of the most ancient clonal organisms in existence. Look down at your feet and you may find yourself surrounded by a species of moss known as Sphagnum palustre. Although this species enjoys a broad distribution throughout the northern hemisphere, its presence on this remote volcanic island is worth closer inspection. 

Hawai'i is rather depauperate in Sphagnum representatives and those that have managed to get to this archipelago are often restricted to growing in narrow habitable zones between 900 to 1,900 meters in elevation as these are the only spots that are cool and wet enough to support Sphagnum growth. Needless to say, successful colonization of the Hawaiian Islands by Sphagnum has been a rare event.  The fact that Sphagnum palustre was one of the few that did should not come as any surprise. What should surprise you, however, is how this particular species has managed to persist. 

 Mounds of  S. palustre  in its native habitat. 

Mounds of S. palustre in its native habitat. 

Hawaiian moss aficionados have long noted that the entire population of Kohala's S. palustre mats never seem to produce a single female individual. Indeed, this moss is dioicous, meaning individuals are either male or female. As such, many have suspected that the mats of S. palustre growing on Kohala represented a single male individual that has been growing vegetatively ever since it arrived as a spore on the island. The question then becomes, how long has this S. palustre individual been on Kohala?

To answer that, researchers decided to take a look at its DNA. What they discovered was surprising in many ways. For starters, all plants were in fact males of a single individual. A rare genetic trait was found in the DNA of every population they sampled. This trait is so rare that the odds of it turning up in any number by sheer chance is infinitesimally small. What this means is that every S. palustre population found on Kohala is a clone of a single spore that landed on the mountain at some point in the distant past. Exactly how distant was the next question the team wanted to answer. 

 A lush cloud forest on the slopes of Kohala.

A lush cloud forest on the slopes of Kohala.

The first clue to this mystery came from peat deposits found on the slopes of the mountain. Researchers found remains of S. palustre in peat deposits that were dated to somewhere around 24,000 years old. So, it would appear that S. palustre has been growing on Kohala since at least the late Pleistocene. But how long before that time did this moss arrive?

Again, DNA was the key to unlocking this mystery. By studying the rate at which mutations arise and fix themselves within the genetic code of this plant, they were able to estimate the average rate of mutation through time. By sampling different moss populations on Kohala, they could then use those estimates to figure out just how long each mat has been growing. Their estimates suggest that the ancestral male sport arrived on Hawai'i somewhere between 49,000 and 50,000 years ago and it has been cloning itself ever since. 

 A large mat of  S. palustre

A large mat of S. palustre

As if that wasn't remarkable in and of itself, their thorough analysis of the genetic diversity within S. palustre revealed a remarkable amount of genetic diversity for a clonal organism. Though not all genetic mutations are beneficial, enough of them have managed to fix themselves into the DNA of the moss clones over thousands of years. The DNA of S. palustre is challenging long-held assumptions about genetic diversity of asexual organisms.

Of course, no conversation about Hawaiian botany would be complete without mention of invasive species. As one can expect at this point, Kohala's S. palustre populations are being crowded out by more aggressive vegetation introduced from elsewhere in the world. Unlike a lot of Hawaiian plants, however, the clonal habit of S. palustre puts a more nuanced twist to this story. 

Because Sphagnum is spongy yet durable, it has often been used as packing material. Packages stuffed with S. palustre from Kohala have been sent all over the island and because of this, S. palustre is now showing up en masse on other islands in the archipelago. Sadly, when it starts to grow in habitats that have never experienced the ecosystem engineering traits of a Sphagnum  moss, S. palustre gets pretty out of hand. It's not just packages that spread it either. All it takes is one sprig of the moss stuck on someone's boot to start a new colony elsewhere. The unique flora elsewhere in the Hawaiian archipelago have not evolved to compete with S. palustre and as a result, escaped populations are rapidly changing the ecology to the detriment of other endemic Hawaiian plants. 

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

Further Reading: [1] [2] 

Are Algae Plants?

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I was nibbling on some nori the other day when a thought suddenly hit me. I don't know squat about algae. I know it comes in many shapes, sizes, and colors. I know it is that stuff that we used to throw at each other on the beach. I know that it photosynthesizes. That's about it. What are algae? Are they even plants?

The shortest answer I can give you is "it depends." The term algae is a bit nebulous in and of itself. In Latin, the word "alga" simply means "seaweed." Algae are paraphyletic, meaning they do not share a recent common ancestor with one another. In fact, without specification, algae may refer to entirely different kingdoms of life including Plantae (which is often divided in the broad sense, Archaeplastida and the narrow sense, Viridiplantae), Chromista, Protista, or Bacteria.

  Caulerpa racemosa , a beautiful green algae.

Caulerpa racemosa, a beautiful green algae.

Taxonomy being what it is, these groupings may differ depending on who you ask. The point I am trying to make here is that algae are quite diverse from an evolutionary standpoint. Even calling them seaweed is a bit misleading as many different species of algae can be found in fresh water as well as growing on land.

 Cyanobacteria are photosynthetic bacteria, not plants.

Cyanobacteria are photosynthetic bacteria, not plants.

Take for instance what is referred to as cyanobacteria. Known commonly as blue-green algae, colonies of these photosynthetic bacteria represent some of the earliest evidence of life in the fossil record. Remains of colonial blue-green algae have been found in rocks dating back more than 4 billion years. As a whole, these types of fossils represent nearly 7/8th of the history of life on this planet! However, they are considered bacteria, not plants.

 Diatoms (Chromista)

Diatoms (Chromista)

Diatoms (Chromista) are another enormously important group. The single celled, photosynthetic organisms are encased in beautiful glass shells that make up entire layers of geologic strata. They comprise a majority of the phytoplankton in the world's oceans and are important indicators of climate. However, they belong to their own kingdom of life - Chromista or the brown algae.

To bring it back to what constitutes true plants, there is one group of algae that really started it all. It is widely believed that land plants share a close evolutionary history with a branch of green algae known as the stoneworts (order Charales). These aquatic, multicellular algae superficially resemble plants with their stalked appearance and radial leaflets.

 A nice example of a stonewort ( Chara braunii ).

A nice example of a stonewort (Chara braunii).

It is likely that land plants evolved from a Chara-like ancestor that may have resembling modern day hornworts that lived in shallow freshwater inlets. Estimates of when this happen go back as far as 500 million years before present. Unfortunately, fossil evidence is sparse for this sort of thing and mostly comes in the form of fossilized spores and molecular clock calculations.

  Porphyra umbilicalis   - One of the many species of red algae frequently referred to as nori.

Porphyra umbilicalis  - One of the many species of red algae frequently referred to as nori.

Now, to bring it back to what started me down this road in the first place. Nori is made from algae in the genus Porphyra, which is a type of Rhodophyta or red algae. Together with Chlorophyta (the green algae), they make up some of the most familiar groups of algae. They have also been the source of a lot of taxonomic debate. Recent phylogenetic analyses place the red algae as a sister group to all other plants starting with green algae. However, some authors prefer to take a broader look at the tree and thus lump red algae in a member of the plant kingdom. So, depending on the particular paper I am reading, the nori I am currently digesting may or may not be considered a plant in the strictest sense of the word. That being said, the lines are a bit blurry and frankly I don't really care as long as it tastes good.

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

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

 

How Air Plants Drink

   Tillandsia tectorum

 Tillandsia tectorum

Air plants (genus Tillandsia) are remarkable organisms. All it takes is seeing one in person to understand why they have achieved rock start status in the horticulture trade. Unlike what we think of as a "traditional" plant lifestyle, most species of air plants live a life free of soil. Instead, they attach themselves to the limbs and trunks of trees as well as a plethora of other surfaces. 

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Living this way imposes some serious challenges. The biggest of these is the acquisition of water. Although air plants are fully capable of developing roots, these organs don't live very long and they are largely incapable of absorbing anything from the surrounding environment. The sole purpose of air plant roots is to anchor them to whatever they are growing on. How then do these plants function? How do they obtain water and nutrients? The answer to this lies in tiny structures called trichomes. 

Trichomes are what gives most air plants their silvery sheen. To fully appreciate how these marvelous structures work, one needs some serious magnification. A close inspection would reveal hollow, nail-shaped structures attached to the plant by a stem. Instead of absorbing water directly through the leaf tissues, these trichomes mediate the process and, in doing so, prevent the plant from losing more water than it gains. 

The trichomes themselves start off as living tissue. During development, however, they undergo programmed cell death, leaving them hollow. When any amount of moisture comes into contact with these trichomes, they immediately absorb that water, swelling up in the process. As they swell, they are stretched out flat along the surface of the leaf. This creates a tiny film of water between the trichomes and the rest of the leaf, which only facilitates the absorption of more water. 

 Trichomes up close.  

Trichomes up close.  

Because the trichomes form a sort of conduit to the inside of the leaf, water and any nutrients dissolved within are free to move into the plant until the reach the spongy mesophyll cells inside. In this way, air plants get all of their water needs from precipitation and fog. Not all air plants have the same amount of trichomes either. In fact, trichome density can tell you a lot about the kind of environment a particular air plant calls home. 

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The fuzzier the plant looks, the drier the habitat it can tolerate. Take, for instance, one of the fuzziest air plants - Tillandsia tectorum. This species hails from extremely arid environments in the high elevation regions of Ecuador and Peru. This species mainly relies on passing clouds and fog for its moisture needs and thus requires lots of surface area to collect said water. Now contrast that with a species like Tillandsia bulbosa, which appears to have almost no trichome cover. This smoother looking species is native to humid low-land habitats where high humidity and frequent rain provide plenty of opportunities for a drink. 

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Absorbing water in this way would appear to have opened up a plethora of habitats for the genus Tillandsia. Air plants are tenacious plants and worthy of our admiration. One could learn a lot from their water savvy ways. 

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

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

Ferns Afloat

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My introduction to the genus Salvinia was as an oddball aquarium plant floating in a display tank at the local pet store. I knew nothing about plants at the time but I found it to be rather charming nonetheless. Every time the green raft of leaves floated under the filter outlet, water droplets would bead off them like water off of a ducks back. Even more attractive were the upside down forest of "roots" which were actively sheltering a bunch of baby guppies. 

I grew some Salvinia for a few years before my interest in maintaining aquariums faded. I had forgotten about them for quite some time. Much later as I was diving into the wild world of botany, I started revisiting some of the plants that I had grown in various aquariums to learn more about them. It wasn't long before the memory of Salvinia returned. A quick search revealed something quite astonishing. Salvinia are not flowering plants. They are ferns! 

The genus Salvinia is quite wide spread. They can be found growing naturally throughout North, Central, and South America, the West Indies, Europe, Africa, and Madagascar. Sadly, because of their popularity as aquarium and pond plants, a few species have become extremely aggressive invaders in many water ways. More on that in a bit. 

Salvinia is comprised of roughly 12 different species. Of these, at least 4 are suspected to be naturally occurring hybrids. As you have probably already gathered, these ferns live out their entire lives as floating aquatic plants. Their most obvious feature are the pairs of fuzzy green leaves borne on tiny branching stems. These leaves are covered in trichomes that repel water, thus keeping them dry despite their aquatic habit. 

 These are not roots!

These are not roots!

Less obvious are the other types of leaves these ferns produce. What looks like roots dangling below the water's surface are actually highly specialized, finely dissected leaves! I was quite shocked to learn this and to be honest, it makes me appreciate these odd little ferns even more. Its on these underwater leaves that the spores are produced. Specialized structures called sporocarps form like tiny nodules on the tips of the leaf hairs.

Sporocarps come in two sizes, each producing its own kind of spore. Large sporocarps produce megaspores while the smaller sporocarps produce microspores. This reproductive strategy is called heterospory. Microspores germinate into gametophytes containing male sex organs or "antheridia" whereas the megaspores develop into gametophytes containing female sex organs or "archegonia." 

As I mentioned above, some species of Salvinia have become aggressive invaders, especially in tropical and sub-tropical water ways. Original introductions were likely via plants released from aquariums and ponds but their small spores and vegetative growth habit means new introductions occur all too easily. Left unchecked, invasive Salvinia can form impenetrable mats that completely cover entire bodies of water and can be upwards of 2 feet thick!

 Sporocarps galore! 

Sporocarps galore! 

Lots of work has been done to find a cost effective way to control invasive Salvinia populations. A tiny weevil known scientifically as Cyrtobagous singularis has been used with great success in places like Australia. Still, the best way to fight invasive species is to prevent them from spreading into new areas. Check your boots, check your boats, and never ever dump your aquarium or pond plants into local water ways. Provided you pay attention, Salvinia are rather fascinating plants that really break the mold as far as most ferns are concerned. 

Photo Credits: [1] [2] [3]

Further Reading: [1] [2] [3]

 

How Trees Fight Disease

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Plants do not have immune systems like animals. Instead, they have evolved an entirely different way of dealing with infections. In trees, this process is known as the "compartmentalization of decay in trees" or "CODIT." CODIT is a fascinating process and many of us will recognize its physical manifestations.

In order to understand CODIT, one must know a little something about how trees grow. Trees have an amazing ability to generate new cells. However, they do not have the ability to repair damage. Instead, trees respond to disease and injury  by walling it off from their living tissues. This involves three distinct processes. The first of these has to do with minimizing the spread of damage. Trees accomplish this by strengthening the walls between cells. Essentially this begins the process of isolating whatever may be harming the living tissues.

This is done via chemical means. In the living sapwood, it is the result of changes in chemical environment within each cell. In heartwood, enzymatic changes work on the structure of the already deceased cells. Though the process is still poorly understood, these chemical changes are surprisingly similar to the process of tanning leather. Compounds like tannic and gallic acids are created, which protect tissues from further decay. They also result in a discoloration of the surrounding wood. 

The second step in the CODIT process involves the construction of new walls around the damaged area. This is where the real compartmentalization process begins. The cambium layer changes the types of cells it produces around the area so that it blocks that compartment off from the surrounding vascular tissues. These new cells also exhibit highly altered metabolisms so that they begin to produce even more compounds that help resist and hopefully stave off the spread of whatever microbes may be causing the injury. Many of the defects we see in wood products are the result of these changes.

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The third response the tree undergoes is to keep growing. New tissues grow around the infected compartment and, if the tree is healthy enough, will outpace further infection. You see, whether its bacteria, fungi, or a virus, microbes need living tissues to survive. By walling off the affected area and pumping it full of compounds that kill living tissues, the tree essentially cuts off the food supply to the disease-causing organism. Only if the tree is weakened will the infection outpace its ability to cope.

Of course, CODIT is not 100% effective. Many a tree falls victim to disease. If a tree is not killed outright, it can face years or even decades of repeated infection. This is why we see wounds on trees like perennial cankers. Even if the tree is able to successfully fight these repeat infections over a series of years, the buildup of scar tissues can effectively girdle the tree if they are severe enough.

CODIT is a well appreciated phenomenon. It has set the foundation for better tree management, especially as it relates to pruning. It is even helping us develop better controls against deadly invasive pathogens. Still, many of the underlying processes involved in this response are poorly understood. This is an area begging for deeper understanding.

Photo Credits: kaydubsthehikingscientist & Alex Shigo

Further Reading: [1]

The Strangest Wood Sorrel

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For me, wood sorrels are a group of plants I usually have to look down to find. This is certainly not the case for Oxalis gigantea. Native to the coastal mountains of northern Chile, this bizarre Oxalis has forgone the traditional herbaceous habit of its cousins in exchange for a woody shrub-like growth form.

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When I first laid eyes on O. gigantea, I thought I was looking at some strange form of Ocotillo. In front of me was a shrubby plant consisting of multiple upright branches that were covered in a dense layer of shiny green leaves occasionally interrupted by yellow flowers. You would think that at this point in my life, aberrant taxa would not longer surprise me. Think again. 

O. gigantea is one of the largest of the roughly 570 Oxalis species known to science. Its woody branches can grow to a height of 2 meters (6 feet)! The branches themselves are quite interesting to look at. They are covered in woody spurs from which clusters of traditional Oxalis-style leaves emerge. Each stem is capable of producing copious amounts of flowers all throughout the winter months. The flowers are said to be pollinated by hummingbirds but I was not able to find any data on this. 

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This shrub is but one part of the Atacama Desert flora. This region of Chile is quite arid,  experiencing a 6 to 10 month dry season every year. What rain does come is often sparse. Any plant living there must be able to cope. And cope O. gigantea does! This oddball shrub is deciduous, dropping its leaves during the dryer months. During that time, these shrubs look pretty ragged. You would never guess just how lush it will become once the rains return. Also, it has a highly developed root system, no doubt for storing water and nutrients to tide them over.  

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O. gigantea has enjoyed popularity as a horticultural oddity over the years. In fact, growing this shrub as a container plant is said to be quite easy. Despite its garden familiarity, O. gigantea is noticeably absent from the scientific literature. In writing this piece, I scoured the internet for any and all research I could find. Sadly, it simply isn't there.

This is all too often the case for unique and interesting plant species like O. gigantea. Like so many other species, it has suffered from the disdain academia has had for organismal research over the last few decades. We humans can and must do better than that. For now, what information does exist has come from horticulturists, gardeners, and avid botanizers from around the world. 

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

Further Reading: [1] [2]