Süßwassertang: A Fern Disguised as a Liverwort

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If you enjoy planted aquariums, you may have crossed paths with a peculiar little plant called Süßwassertang. It can be propagated by breaking off tiny pieces, which eventually grow into a tangled carpet of tiny green thalli. One could be excused for thinking that Süßwassertang was some sort of liverwort and indeed, for quite some time was marketed as such. That all changed in 2009 when it was revealed that this was not a liverwort at all but rather the gametophyte of a fern.

Despite its German name, Süßwassertang appears to have originated in tropical parts of Africa and Asia. It is surprisingly hard to find out any information about this plant outside of its use in the aquarium trade. The name Süßwassertang translates to “freshwater seaweed” and indeed, that is exactly what it looks like. The fact that this is actually the gametophyte of a fern may seem startling at first but when you consider what they must deal with in nature, the situation makes a bit more sense.

A  Süßwassertang gametophyte.  B  An antheridium, showing a cap cell ( cc ), ring cell ( rc ), and basal cell ( bc ).  Bar : 20 µm.  C  Developing lateral branches with rhizoids ( arrowhead ) and meristems ( m )  Bar : 0.2 mm.  D  Ribbon-like, branched gametophyte ( g ) of  L. spectabilis  bearing a young sporophyte ( sp )  Bar : 1 cm

A Süßwassertang gametophyte. B An antheridium, showing a cap cell (cc), ring cell (rc), and basal cell (bc). Bar: 20 µm. C Developing lateral branches with rhizoids (arrowhead) and meristems (m) Bar: 0.2 mm. D Ribbon-like, branched gametophyte (g) of L. spectabilis bearing a young sporophyte (sp) Bar: 1 cm

Fern gametophytes are surprisingly hardy considering their small size and delicate appearance. They are amazing in their ability to tolerate harsh conditions like drought and freezing temperatures. Because of this, fern gametophytes sometimes establish themselves in places that would be unfavorable for their sporophyte generation. For some, this means never completing their lifecycle. Others, however, seem to have overcome the issue by remaining in their gametophyte stage forever. Though no sexual reproduction occurs for these permanent gametophytes, they nonetheless persist and reproduce by breaking off tiny pieces, which grow into new colonies.

The sporophyte of a related species,  Lomariopsis marginata , demonstrating the usual epiphytic habit of this genus.

The sporophyte of a related species, Lomariopsis marginata, demonstrating the usual epiphytic habit of this genus.

This appears to be the case for Süßwassertang. Amazingly, despite a few attempts, no sporophytes have ever been coaxed from any gametophyte. It would appear that this is yet another species that has given up its sporophyte phase for an entirely vegetative habit. What is most remarkable is what the molecular work says about Süßwassertang taxonomically. It appears that this plant its nestled into a group of epiphytic ferns in the genus Lomariopsis. How this species evolved from vine-like ferns living in trees to an asexual colony of aquatic gametophytes is anyones’ guess but it is an incredible jump to say the least.

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

Further Reading: [1]

Gooey Pitcher Fluids

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There seems to be no end to the diversity of colors, shapes, and sizes exhibited by Nepenthes and their pitchers. These wonderful carnivorous plants grow these pitchers as a means of supplementing their nutritional needs as the habitats in which Nepenthes are found are lacking in vital nutrients like nitrogen. There are as many variations on the pitcher theme as there are Nepenthes but all function as traps in one form or another. How they trap insects is another topic entirely and some species have evolved incredible means of making sure prey does not escape. Some of my favorites belong to those species that employ sticky mucilage.

Arguably one of the most iconic of this type is Nepenthes inermis. This species is endemic to a small region of Sumatra and, to date, has only been found growing on a handful of mountain peaks in the western part of the country. The specific epithet ‘inermis’ is Latin for ‘unarmed’ as was given in reference to the bizarre upper pitchers of this plant. They look more like toilet bowls than anything carnivorous and indeed, they lack many of the features characteristic of other Nepenthes pitchers such as a peristome and a slippery, waxy coating on the inside of the pitcher walls.

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These may seem like minor details but consider the role these features play in other Nepenthes. A peristome is essentially a brightly colored, slippery lip that lines the outer rim of the pitcher mouth. Not only does this serve in attracting insect prey, it also aids in their capture. As mentioned, the peristome can be extremely slippery (especially when wet) so that any insect stumbling around on the rim is much more likely to fall in. Once inside, a waxy coating on the inside of some pitchers aids in keeping insects down. They simply cannot get purchase on the waxy walls and therefore cannot climb back out. So, for N. inermis to lack both features is a bit strange.

Another interesting feature of N. inermis pitchers is the highly reduced pitcher lid. It hasn’t disappeared completely but compared with other Nepenthes, this pitcher lid barely registers as one. For most Nepenthes, pitcher lids serve multiple functions. For starters, they keep the rain out. Nepenthes are msot at home in humid, tropical climates where rain is a daily force to be reckoned with. For many Nepenthes, rain not only dilutes the valuable digestive soup brewing within each pitcher, it can also cause them to overflow and dump their nutritious contents. Pitcher lids can also help in attracting prey. Like the peristome, they are often brightly colored but many also secrete nectar, which insects find irresistible. Lured in by the promise of food, some insects inevitably fall down into the pitcher below.

Looking into the pitcher of  Nepenthes inermis .

Looking into the pitcher of Nepenthes inermis.

Considering the importance of such structures, it becomes a little bit confusing why some Nepenthes have evolved away from this anatomy. The question then remains, why would a species like N. inermis no longer produce pitchers with these features? Amazingly, the answer actually lies within the pitcher fluid itself.

Tip over the upper pitchers of N. inermis and you will soon discover that they are filled with an extremely viscous mucilage. It is so viscous that some have reported that when the pitchers are held upside down, the mucilage within can form an unbroken stream of considerable length. Its the viscosity of this fluid that is the real reason that N. inermis is able to capture prey so easily. Insects lured to the traps can catch a drink of the nectar on the tiny lid. In doing so, some inevitably fall down into the pitcher itself.

The upper pitcher of the closely related  Nepenthes dubia .

The upper pitcher of the closely related Nepenthes dubia.

Instead of slippery walls or downward pointing hairs keeping the insects in, the viscous pitcher fluid quickly engulfs the struggling prey. Some have even suggested that the nectar secreted by the tiny lid has narcotic effects on visiting insects, however, I have not seen any data demonstrating this. Once caught in the fluid, insects easily slide their way down into the depths of the pitcher where they can be digested. This is probably why the pitchers are shaped like tiny toilet bowls; their shape allows for a large sticky surface area for insects to get stuck while prey that has already been captured is funneled down to where digestion and absorption takes place. In a way, these types of pitchers behave surprisngly similar to the sticky traps utilized by other carnivorous plants like sundews (Drosera spp.).

The viscous fluid also comes in handy during the frequent rains that blanket these mountains. As mentioned above, rain would quickly dilute most pitcher fluids but not when the pitcher fluid itself is more dense. Water sits on top of the viscous mucilage and when the pitchers become too heavy, they tip over. The water readily pours out but little if any of the pitcher fluid is lost in the process. It seems that species like N. inermis no longer fight the elements but rather have adapted to meet them head on. As such, they no longer have a need for a large pitcher lid.

Nepenthes jamban  takes the toilet bowl shape to the extreme.

Nepenthes jamban takes the toilet bowl shape to the extreme.

Nepenthes inermis is not alone in having evolved pitchers like this. Viscous pitcher mucilage is a trait shared by its closest relatives - N. dubia, N. flava, N. jacquelineae, N. jamban, N. talangensis, and N. tenuis, as well as even more distantly related species such as N. rafflesiana. Because prey capture is so important for the fitness of individuals, it is no wonder that so many different forms have evolved within this genus. In fact, many experts believe that variations in the way in which prey is captured and utilized is one of the main reasons why Nepenthes have undergone such a dramatic adaptive radiation.

Sadly, the uniqueness in form and function of these pitchers has landed many of these species on the endangered species list. As if habitat destruction wasn’t already pushing some to the brink, species like N. inermis are being poached at alarmingly unsustainable rates. Due to their limited distributions, most populations simply cannot recover from even moderate levels of harvesting. The silver lining in all of this is that many Nepenthes are extremely easy to grow and propagate provided their basic needs are met. As more and more folks enter into the carnivorous plant hobby, hopefully more and more people will be sharing seeds, cuttings, and tissue cultured materials. In doing so, we can hopefully reduce some of the pressures placed on wild populations.

Photos via Wikimedia Commons

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

Fossilized Flower Places Angiosperms in the Jurassic

1, style branches; 2, dendroid style; 3, sepal; 4, ovarian roof; 5, scale; 6, seed; 7, cup-form receptacle/ovary; 8, bract; 9, petal; 10, unknown organ (staminode?).  [SOURCE]

1, style branches; 2, dendroid style; 3, sepal; 4, ovarian roof; 5, scale; 6, seed; 7, cup-form receptacle/ovary; 8, bract; 9, petal; 10, unknown organ (staminode?). [SOURCE]

Despite their dominance on the landscape today, the origin of flowering plants is shrouded in mystery. The odds of any living material becoming fossilized is extremely rare and when you consider the delicate and ephemeral nature of most flowers, one can begin to understand why their fossils are so special. The last few decades have seen tantalizing evidence emerge from fossil beds dating to the Cretaceous Period but a recent set of fossils from China predate the oldest confirmed angiosperm fossils by 50 million years. That’s right, it would appear that flowering plants were already on the scene by the early Jurassic!

The fossils in question have been coined Nanjinganthus dendrostyla. They were discovered in China in a formation that dates back roughly 174 million years. To most of us they look like a bunch of dark, albeit elaborate smudges on the rocks. To a trained eye, however, these smudges reveal intricate anatomical details. Amazingly, the team of paleobotanists responsible for this discovery had a lot of material to work with. Descriptions were made on a whopping 264 specimens representing 198 individual flowers. This amount of data means that the declaration of angiosperm affinity stands on pretty solid ground.

A single  Nanjinganthus  flower  [SOURCE]

A single Nanjinganthus flower [SOURCE]

Aside from their age, there is a lot about these fossils that surprised researchers. Probably the biggest surprise is their overall appearance. Paleobotanists have long hypothesized that early angiosperm flowers likely resembled something akin to a modern day Magnolia and invoke floral features such as apocarpy, a superior ovary, and a lack of an obvious style as likely features to look for in ancient plant fossils. Surprisingly, Nanjinganthus does not seem to conform to many of these expectations.

One of the most striking features of these fossils are the styles. They are large and branched like tiny trees (hence the specific epithet “dendrostyla”). The tree-like appearance of the style suggests that early angiosperms likely did not rely on insects for pollination. The branches themselves greatly increase the amount of surface area available for pollen capture, which could mean that Nanjinganthus was wind pollinated.

Flowers of  Nanjinganthus  preserved in different states and their details. For specific details on each image, please see   SOURCE

Flowers of Nanjinganthus preserved in different states and their details. For specific details on each image, please see SOURCE

Another surprising feature is the presence of an inferior ovary that, by its very definition, sits below the sepals and petals. It has long been hypothesized that early angiosperms would exhibit superior ovaries so this discovery means that we must rethink our expectations of how flowers evolved. For instance, it suggests we may not be able to make broad inferences on the past based on what we see in extant angiosperm lineages. It could also suggest that the origin of flowering plants was not a single event but rather a series of individual occurrences. It could also be the case that the origin of flowering plants occurred much earlier than the Jurassic and that Nanjinganthus represents one of many derived forms. Only further study and more fossils can help us answer such questions.

Another way in which Nanjinganthus deviates from theoretical expectations is in the presence of both sepals and petals. Up until now, paleobotanists have been fond of the idea that petals arose much later in angiosperms, having evolved over time as leaves became more and more specialized for attracting pollinators. The fact that Nanjinganthus was likely wind pollinated yet had both sepals and petals is a bit of a conundrum and further emphasizes the need to revisit some of our long-held assumptions of flowering plant evolution.

Details of the sepal and petal as seen through different forms of microscopic analysis. For specific details on each image, please see  SOURCE .

Details of the sepal and petal as seen through different forms of microscopic analysis. For specific details on each image, please see SOURCE.

By far the most important feature present in these fossils are the ovaries. For any fossil to unequivocally qualify as an angiosperm, it must have seeds encased in an ovary. This, after all, is the main feature that separates angiosperms from gymnosperms. Indeed, Nanjinganthus does appear to fit this definition. Thanks to the sheer amount of fossils available for study, the team discovered that the seeds of Nanjinganthus were enclosed in a cup-like chamber that was sealed off from the outside world by a structure they refer to as an “ovarian roof.” This roof does not appear to have any sort of opening, which worked out quite nicely for paleobotanists as it prevented sediments from entering into the chamber, thus preserving the seeds or ovules (it is hard to tell where they were in the developmental process) for study. This feature more than all others secures its identity as a flowering plant.

Based on the sediments in which these flowers were fossilized, it appears that this plant grew close to water. Also, despite its abundance in this particular fossil layer, it very likely was not a common component of this Jurassic landscape. In reality we still have a lot to learn about Nanjinganthus. What we can say with some certainty at this point is that the presence of Nanjinganthus in the early Jurassic likely means that flowering plants arose even earlier. Nanjinganthus is most definitely not the first flower. We will probably never find the first of anything. It is an ancient flower though, predating all other discoveries by at least 50 million years. This is why paleontology is so incredible. Who knows what the next blow of a rock hammer will turn up!

 

EDIT (10/27/2018): Since writing this post it has come to my attention that there is quite a bit of controversy attached to the description of this fossil. Many have reached out informing me that these fossils may actually be a gymnosperm organ rather than a flower. Despite all of the outcry I have yet to see any published critiques on this particular controversy. I anxiously await more professional input on the subject but for now I have decided to keep the content of the original piece as is. Of course extraordinary claims require extraordinary evidence and not being a paleobotanist myself, I cannot trust hearsay on the internet as fact, no matter how vociferous, until I see it published in a peer reviewed outlet of some sort. Please stay tuned as this story develops! 

Photo Credits: [1]

Further Reading: [1]

An Introduction to Hornworts

Anthoceros  sp.

Anthoceros sp.

When was the last time you thought about hornworts? Have you ever thought about hornworts? If you answered no, you aren’t alone. Despite their global distribution, these tiny plants receive hardly any attention and that is a shame. Hornworts (Anthocerotophyta) have been around for a very long time. In fact, it is likely that they were some of the first plants to colonize the land roughly 300 - 400 million years ago.

To be fair, hornworts aren’t known for their size. They are generally small plants, though their colonies can form impressive mats. To find them, one must try looking in and among rocks, bare patches of soil, or pretty much anywhere enough moisture builds up to supply their needs. They tend to enjoy nutrient-poor substrates but I would hesitate to say that with any certainty. No matter where you live, from the tundra to the tropics, there is probably a hornwort native to your neck of the woods.

Dendroceros  sp.

Dendroceros sp.

How many different species of hornwort there are is apparently the subject of some debate. Some authors recognize upwards of 300 species whereas others suggest the real number hangs somewhere around 150. Regardless of the exact numbers, hornworts belong to one of six genera: Anthoceros, Dendroceros, Folioceros, Megaceros, Notothylas and Phaeoceros. Fun fact, the suffix ‘ceros’ at the end of each genus is derived from the Latin word for ‘horn.’

The reason they are called hornworts is because of their reproductive structures or “sporophytes.” Similar to their moss and liverwort cousins, hornworts undergo an alternation of generations in order to reproduce sexually. The green gametophytes house the sexual organs - antheridia if they are male and archegonia if they are female. After fertilization, a sporophyte begins to grow, which will go on to produce and disseminate spores. However, the way in which the hornwort sporophyte forms is a bit different from what we see in mosses and liverworts.

Alternation of generations in hornworts.

Alternation of generations in hornworts.

Upon fertilization, the zygote begins to divide into a bulbous mass of cells affectionately referred to as "the foot.” This foot remains within the gametophyte throughout the lifetime of the hornwort, depending on the gametophyte for water and nutrients. Even more peculiar is the the fact that the growing point of the sporophyte is at the base rather than the tip. As such, the horn of each hornwort could continue to grow upwards until it is damaged in some way.

The horn itself is an amazing structure. Whereas the outside layers of tissue are merely structural, the internal tissues differentiate into two different types - spores and pseudo-elaters. Pseudo-elaters expand and contract as humidity fluctuates so as the sporophyte splits to release the spores, the pseudo-elaters dehydrate and snap like tiny spore catapults, thus aiding in their dispersal.

Megaceros flagellaris

Megaceros flagellaris

Of course, reproduction is the main goal but to get to that point, hornworts must grow and mature. How they manage to survive is incredible because it is a reminder that what are often thought of as “primitive” plants are actually far more advanced than we give them credit for. The main body of the hornwort gametophyte is a thin layer of cells that spread out to form a tiny, green mat. This is the structure you are most likely to encounter.

Inside each cell is a single chloroplast. In most hornworts, the chloroplast does not exist in isolation. Instead, it is fused with other organelles into a structure called a “pyrenoid.” The pyrenoid functions as both a center for photosynthesis and a food storage organ. This is unique as it relates to terrestrial plants but quite common in algae. Another odd fact about hornwort anatomy are the presence of tiny cavities scattered throughout their tissues. These cavities form as clusters of hornwort cells die. They then fill with a special mucilage that appears to invite colonization by nitrogen-fixing cyanobacteria. The cyanobacteria set up shop within the cavities and provides the hornwort with supplemental nitrogen in return for a place to live.

Anthoceros agrestis

Anthoceros agrestis

Cyanobacteria aren’t the only organisms to have partnered with hornworts either. Mycorrhizal fungi also enter into the picture. A study done back in 2013 actually found that a wide variety of fungi will partner with hornworts which suggests that this symbiotic relationship is much more ancient and versatile than we once thought. Fungi cluster around parts of the gametophyte that produce root-like structures called “rhizoids,” offering nutrients in return for carbohydrates.

All in all, I think it is safe to say that hornworts are remarkable little plants. Though they can sometimes be difficult to find and properly identify, they nonetheless offer plenty of inspiration for the botanically inclined mind. We can all do better by tiny plants like the hornworts. They have been on land for an incredible amount of time and they definitely deserve our respect and admiration.

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

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

Something Strange in Mexico

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I assure you that what you are looking at here is indeed a plant. I would like you to meet the peculiar Lacandonia schismatica, one of roughly 55 species belonging to the family Triuridaceae. Not a single member of this family bothers with leaves or even chlorphyll. Instead, all members are mycoheterotrophic, meaning they make their living by parasitizing fungi in the soil. However, that is not why L. schismatica is so strange. Before we get to that, however, it is worth getting to know this plant a little bit better.

The sole member of its genus, Lacandonia schismatica grows in only a few locations in the Lancandon Jungle of southeastern Mexico. Its populations are quite localized and are under threat by encroaching agricultural development. Genetic analyses of the handful of known populations revealed that there is almost no genetic diversity to speak of among the individuals of this species. All in all, these factors have landed this tiny parasite on the endangered species list.

Mature flower of  Lacandonia schismatica . Three yellowish anthers (center) surrounded by rings of red carpels. Scale bar = 0.5cm.”  [SOURCE]

Mature flower of Lacandonia schismatica. Three yellowish anthers (center) surrounded by rings of red carpels. Scale bar = 0.5cm.” [SOURCE]

To figure out why L. schismatica is so peculiar, you have to take a closer look at its flowers. If you knew what to look for, you would soon realize that L. schismatica appear to be doing things in reverse. To the best of our knowledge, L. schismatica is the only plant in the world that known to have an inverted flower arrangement. The anthers of this species are clustered in the center of the flower surrounded by a ring of 60 or so pistils. The flowers are cleistogamous, which means they are fertilized before they even open, hence the lack of genetic diversity among individuals. 

Not all of its flowers take on this appearance. Researchers have found that in any given population, a handful of unisexual flowers will sometimes be produced. Even the bisexual flowers themselves seem to exhibit at least some variation in the amount of sexual organs present. Still, when bisexual flowers are produced, they only ever exhibit this odd inverted arrangement.

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It is not quite clear how this system could have evolved in this species. Indeed, this unique floral morphology has made this species very hard to classify. Genetic analysis suggests a relation to the mycoheterotrphic family Triuridaceae. It was discovered that every once in a while, a closely related species known as Triuris brevistylis will sometimes produce flowers with a similar inverted morphology.

This suggests that the inversion evolved before the Lacandonia schismatica lineage diverged. One can only speculate at this point. The future of this species is quite uncertain. Climate change and habitat destruction could permanently alter the conditions so that this plant can no longer exist in the wild. This is further complicated by the fact that this species has proven to be quite difficult to cultivate. Only time will tell. For now, more research is needed on this peculiar plant.

Photo Credit: [1] [2] [3]

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

The Rise and Fall of the Scale Trees

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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.

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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]

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.  Photo copyright Bruce Kirchoff, Licensed under CC-BY

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

Photo copyright Bruce Kirchoff, Licensed under CC-BY

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]

Meet Pokeweed's Tree-Like Cousin

There is more than one way to build a tree. For that reason and more, “tree” is not a taxonomic designation. Arborescence has evolved independently throughout the botanical world and many herbaceous plants have tree-like relatives. I was shocked to learn recently of a plant native to the Pampa region of South America commonly referred to as ombú. At first glance it looks like some sort of fig, with its smooth bark and sinuous, buttressed roots. Deeper investigation revealed that this was not a fig. Ombú is actually an arborescent cousin of pokeweed!

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The scientific name of ombú is Phytolacca dioica. As its specific epithet suggests, plants are dioecious meaning individuals are either male or female. Unlike its smaller, herbaceous cousins, ombú is an evergreen perennial. Because they can grow all year, these plants can reach bewildering proportions. Heights upwards of 60 ft. (18 m.) are not unheard of and the crowns of more robust specimens can easily attain diameters of 40 to 50 ft. (12 - 18 m.)! What makes such sizes all the more impressive is the way in which ombú is able to achieve such growth.

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Ombú is thought to have evolved from an herbaceous ancestor. Cut into the trunk of one of these trees and you will see that this phylogenetic history has left its mark. Ombú do not produce what we think of as wood. Instead, much of the support for branches and stems comes from turgor pressure. Also, the way in which these trees grow is not akin to what you would see from something like an oak or a maple. Whereas woody trees undergo secondary growth in which the cambium layer differentiates into xylem and phloem, thus thickening stems and roots, ombú exhibits a unique form of stem and root thickening called “anomalous secondary thickening.”

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Essentially what this means is that instead of a single layer of cambium forming xylem and phloem, ombú cambium exhibits unidirectional thickening of the cambium layer. There are a lot of nitty gritty details to this kind of growth and I must admit I don’t have a firm grasp on the mechanics of it all. The point of the matter is that anomalous secondary thickening does not produce wood as we know it and instead leads to rapid growth of weak and spongy tissues. This is why turgor pressure is so important to the structural integrity of these trees. It has been estimated that the trunk and branches of an ombú is 80% water.

A cross section of an ombú limb showing harder xylem tissues separated by spongy parenchyma that has since disintegrated.

A cross section of an ombú limb showing harder xylem tissues separated by spongy parenchyma that has since disintegrated.

Like all members of this genus, ombú is plenty toxic. Despite this, ombú appears to have been embraced and is widely planted as a specimen tree in parks, along sidewalks, and in gardens in South America and elswhere. In fact, it is so widely planted in Africa that some consider it to be a growing invasive issue. All in all I was shocked to learn of this species. It caused me to rethink some of the assumptions I hold onto with some lineages I only know from temperate regions. It is amazing what natural selection has done to this genus and I am excited to explore more arborescent anomalies from largely herbaceous groups.

Photo Credits: [1] [2]

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

Fluorescent Bananas

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Bananas are one of the most popular fruits in the world. Love them or hate them, most of us know what they look like. Despite their global presence, few stop to think about where these fruits come from. That is a shame because bananas are fascinating plants for many reasons but now we can add blue fluorescence to that list.

Before we dive into the intriguing phenomenon of fluorescence in bananas, I think it is worth talking about the plants that produce them in a little more detail. Bananas belong to the genus Musa, which is located in its own family - Musaceae. Take a step back and look at a banana plant and it won't take long to realize they are distant relatives of the gingers. There are at least 68 recognized species of banana in the world and many more cultivated varieties. Despite their pan-tropical distribution, the genus Musa is native only to parts of the Indo-Malesian, Asian, and Australian tropics.

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Banana plants vary in height from species to species. At the smaller end of the spectrum you have species like the diminutive Musa velutina, which maxes out at about 2 meters (6 ft.) in height. On the taller side of things, there are species such as the monstrous Musa ingens, which can reach heights of 20 meters (66ft.)! Despite their arborescent appearance, bananas are not trees at all. They do not produce any wood. Instead, what looks like a tree trunk is actually the fused petioles of their leaves. Bananas are essentially giant herbs with the aforementioned M. ingens holding the world record for largest herb in the world.

When it comes time to flower, a long spike emerges from the main growing tip. This spike gradually elongates, revealing long, beautiful, tubular flowers arranged in whorls. For many banana species, bats are the main pollinators, however, a variety of insects will visit as well. In the wild, fruits appear following pollination, a trait that has been bred out of their cultivated relatives, which produce fruits without needing pollination. The fruits of a banana are actually a type a berry that dehisce like a capsule upon ripening, revealing delicious pulp chock full of hard seeds. Not all bananas turn yellow upon ripening. In fact, some are pink!

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For many fruits, the act of ripening often coincides with a change in color. This is a way for the plant to signal to seed dispersers that the fruits, and the seeds inside, are ready. As many of us know, many bananas start off green and gradually ripen to a bright yellow. This process involves a gradual breakdown of the chlorophyll within the banana skin. As the chlorophyll within the skin of a banana breaks down, it leaves behind a handful of byproducts. It turns out, some of these byproducts fluoresce blue under UV light. 

Amazingly, the fluorescent properties of bananas was only recently discovered. Researchers studying chlorophyll breakdown in the skins of various fruits identified some intriguing compounds in the skins of ripe Cavendish bananas. When viewed under UV light, these compounds gave off a luminescent blue hue. Further investigation revealed that as bananas ripen, their fluorescent properties grow more and more intense.

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There could be a couple reasons why this happens. First, it could simply be happenstance. Perhaps these fluorescent compounds are simply a curious byproduct of chlorophyll breakdown and serve no function for the plant whatsoever. However, bananas seem to be a special case. The way in which chlorophyll in the skin of a banana breaks down is quite different than the process of chlorophyll breakdown in other plants. What's more, the abundance of these compounds in the banana skin seems to suggest that the fluorescence does indeed have a function - seed dispersal.

Researchers now believe that the fluorescent properties of some ripe bananas serves as an additional signal to potential seed dispersers that the time is right for harvest. Many animals including birds and some mammals can see well into the UV spectrum and it is likely that the blue fluorescence of these bananas is a means of attracting such animals. Additionally, researchers also found that banana leaves fluoresce in a similar way, perhaps to sweeten the attractive display of the ripening fruits.

To date, little follow up has been done on fluorescence in bananas. It is likely that far more banana species exhibit this trait. Certainly more work is needed before we can say for sure what role, if any, these compounds play in the lives of wild bananas. Until then, this could be a fun trait to investigate in the comfort of your own home. Grab a black light and see if your bananas glow blue!

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

Further Reading: [1] [2]

Fossils Shine Light On the History of Gall-Making Wasps

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We can learn a lot about life on Earth from the fossil record. I am always amazed by the degree of scrutiny involved in collecting data from these preserved remains. Take, for instance, the case of gall-making wasp fossils found in western North America. A small collection of fossilized oak leaves is giving researchers insights into the evolutionary history of oaks and the gall-making wasps they host.

Oaks interact with a bewildering array of insects. Many of these are gall-making wasps in the family Cynipidae. Dozens of different wasp species can be found on a single oak tree. Female wasps lay their eggs inside developing oak tissues and the larvae release hormones and other chemicals that cause galls to form. Galls are essentially edible nursery chambers. Other than their bizarre shapes and colors, the compounds released by the wasp larvae reduce the chemical defenses of the oak and increase the relative nutrition of the tissues themselves. Often, these relationships are precise, with specific wasp species preferring specific oak species. But when did these relationships arise? Why are oaks so popular? What can fossil evidence tell us about this incredible relationship?

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Though scant, the little fossil evidence of oak galls can tell us a lot. For starters, we know that gall-making wasps whose larvae produce structures similar to that of the Cynipids have been around since at least the late Cretaceous, some 100 million years ago. However, it is hard to say for sure exactly who made these galls and exactly what taxonomic affinity the host plant belongs to. More conclusive Cynipid gall fossils appear again in the Eocene and continue to pop up in the fossil record throughout the Oligocene and well into the Miocene (33 - 23 million years ago).

Miocene aged fossils are where things get a little bit more conclusive. Fossil beds located in the western United States have turned up fossilized oak leaves complete with Cynipid galls. The similarity of these galls to those of some present day species is incredible. It demonstrates that these relationships arose early on and have continued to diversify ever since. What's more, thanks to the degree of preservation in these fossil beds, researchers are able to make some greater conclusions about why gall-making wasps and oaks seem to be so intertwined.

Holotype of Antronoides cyanomontanus galls on fossilized leaves of  Quercus simulata . 1) Impression of the abaxial surface of the leaf, showing the galls extending into the matrix. 2) Galls showing close association with secondary veins. 3) Gall showing the impression of rim-like base partially straddling the secondary vein. 4) Close-up of gall attached at margin extending down into the matrix. 5) Gall uncovered revealing spindle-shaped morphology.

Holotype of Antronoides cyanomontanus galls on fossilized leaves of Quercus simulata. 1) Impression of the abaxial surface of the leaf, showing the galls extending into the matrix. 2) Galls showing close association with secondary veins. 3) Gall showing the impression of rim-like base partially straddling the secondary vein. 4) Close-up of gall attached at margin extending down into the matrix. 5) Gall uncovered revealing spindle-shaped morphology.

1)  Xanthoteras clavuloides  galls on fossilized  Quercus lobata , showing gall attached to secondary vein. Specimen in California Academy of Sciences Entomology collection, San Francisco. 2) Two galls of attached to a secondary vein showing overlap of their bases. Specimen in California Academy of Sciences Entomology Collection, San Francisco. 3) Three galls collected from leaf of California  Quercus lobata  showing clavate shape and expanded, ring-like base. 4) Gall showing the annulate or ribbed aspect of the base, which is similar to bases of  Antronoides cyanomontanus  and  A. polygonalis . 5) Galls showing clavate shape, pilose and nonpilose surfaces, and bases.

1) Xanthoteras clavuloides galls on fossilized Quercus lobata, showing gall attached to secondary vein. Specimen in California Academy of Sciences Entomology collection, San Francisco. 2) Two galls of attached to a secondary vein showing overlap of their bases. Specimen in California Academy of Sciences Entomology Collection, San Francisco. 3) Three galls collected from leaf of California Quercus lobata showing clavate shape and expanded, ring-like base. 4) Gall showing the annulate or ribbed aspect of the base, which is similar to bases of Antronoides cyanomontanus and A. polygonalis. 5) Galls showing clavate shape, pilose and nonpilose surfaces, and bases.

Gall-making wasps seem to diversify at a much faster rate in xeric climates. The fossil records during this time show that mesic tree speciess were gradually being replaced by more xeric species like oaks. Wasps seem to prefer these drier environments and the thought is that such preferences have to do with disease and parasite loads.

Again, galls a large collections of nutrient-rich tissues that are low in defense compounds. Coupled with the juicy grub at their center, it stands to reason that galls make excellent sites of infection for fungi and other parasites. By living in drier habitats, it is believed that gall-making wasps are able to escape these environmental pressures that would otherwise plague them in wetter habitats. The fossil evidence appears to support this hypothesis and today we see similar patterns. White oaks are especially drought tolerant and its this group of oaks that host the highest diversity of gall-making wasps.

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

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]

 

 

 

Toxic Nectar

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I was introduced to the concept of toxic nectar thanks to a species of shrub quite familiar to anyone who has spent time in the Appalachian Mountains. Locals will tell you to never place honeybee hives near a patch of rosebay (Rhododendron maximum) for fear of so-called "mad honey." Needless to say, the concept intrigued me.

A quick internet search revealed that this is not a new phenomenon either. Humans have known about toxic nectar for thousands of years. In fact, honey made from feeding bees on species like Rhododendron luteum and R. ponticum has been used more than once during times of war. Hives containing toxic honey would be placed along known routs of Roman soldiers and, after consuming the seemingly innocuous treat, the soldiers would collapse into a stupor only to be slaughtered by armies lying in wait.

Rhododendron luteum

Rhododendron luteum

The presence of toxic nectar seems quite confusing. The primary function of nectar is to serve as a reward for pollinators after all. Why on Earth would a plant pump potentially harmful substances into its flowers?

It is worth mentioning at this point that the Rhododendrons aren't alone. A multitude of plant species produce toxic nectar. The chemicals that make them toxic, though poorly understood, vary almost as much as the plants that make them. Although there have been repeated investigations into this phenomenon, the exact reason(s) remain elusive to this day. Still, research has drummed up some interesting data and many great hypotheses aimed at explaining the patterns.

Catalpa nectar has been shown to deter some ants and butterflies but not large bees.

Catalpa nectar has been shown to deter some ants and butterflies but not large bees.

The earliest investigations into toxic nectar gave birth to the pollinator fidelity hypothesis. Researchers realized that meany bees appear to be less sensitive to alkaloids in nectar than are some Lepidopterans. This led to speculation that perhaps some plants pump toxic compounds into their nectar to deter inefficient pollinators, leading to more specialization among pollinating insects that can handle the toxins.

Another hypothesis is the nectar robber hypothesis. This hypothesis is quite similar to the pollinator fidelity hypothesis except that it extends to all organisms that could potentially rob nectar from a flower without providing any pollination services. As such, it is a matter of plant defense.

The nectar of  Cyrilla racemiflora  is thought to be toxic to some bees.

The nectar of Cyrilla racemiflora is thought to be toxic to some bees.

Others feel that toxic nectar may be less about pollinators or nectar robbers and more about microbial activity. Sugary nectar can be a breeding ground for microbes and it is possible that plants pump toxic compounds into their nectar to keep it "fresh." If this is the case, the antimicrobial benefits could outweigh the cost to pollinators that may be harmed or even deterred by the toxic compounds.

Finally, it could be that toxic nectar may have no benefit to the plant whatsoever. Perhaps toxic nectar is simply the result of selection for defense compounds elsewhere in the plant and therefore is expressed in the nectar as a result of pleiotropy. If this is the case then toxic nectar might not be under as strong selection pressures as is overall defense against herbivores. If so, the plants may not be able to control which compounds eventually end up in their nectar. Provided defense against herbivores outweighs any costs imposed by toxic nectar then plants may not have the ability to evolve away from such traits.

Where Spathodea campanulata is invasive, its nectar causes increased mortality in native bee hives.

Where Spathodea campanulata is invasive, its nectar causes increased mortality in native bee hives.

So, where does the science land us with these hypotheses? Do the data support any of these theories? This is where things get cloudy. Despite plenty of interest, evidence in support of the various hypotheses is scant. Some experiments have shown that indeed, when given a choice, some bees prefer non-toxic to toxic nectar. Also, toxic nectar appears to dissuade some ants from visiting flowers, however, just as many experiments have demonstrated no discernible effect on bees or ants. What's more, at least one investigation found that the amount of toxic compounds within the nectar of certain species varies significantly from population to population. What this means for pollination is anyone's' guess.

It is worth noting that most of the pollination-related hypotheses about toxic nectar have been tested using honeybees. Because they are generalist pollinators, there could be something to be said about toxic nectar deterring generalist pollinators in favor of specialist pollinators. Still, these experiments have largely been done in regions where honeybees are not native and therefore do not represent natural conditions.

Simply put, it is still too early to say whether toxic nectar is adaptive or not. It could very well be that it does not impose enough of a negative effect on plant fitness to evolve away from. More work is certainly needed. So, if you are someone looking for an excellent thesis project, here is a great opportunity. In the mean time, do yourself a favor and don't eat any mad honey.

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

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

 

 

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]

 

Leafy Cacti?

Pereskia aculeata

Pereskia aculeata

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

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

Pereskia spines are produced from areoles in typical cactus fashion.

Pereskia spines are produced from areoles in typical cactus fashion.

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

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

Pereskia grandifolia

Pereskia grandifolia

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

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

Pereskia aculeata

Pereskia aculeata

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

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

Further Reading: [1] [2]

 

 

The Desert Mistletoe: Evolution In Action

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There are a multitude of mistletoes on this planet (for example: 1, 2, 3) and all of them are parasites to one degree or another. I find parasitic plants absolutely fascinating as there are many variations on this lifestyle as there are hosts to parasitize. On a recent botanical adventure in the Sonoran Desert, I met yet another representative of this group - the desert mistletoe (Phoradendron californicum). Once I knew what I was looking at, I could not wait to do some research. As it turns out, this species has garnered quite a bit of attention over the years and it is teaching us some interesting tidbits on how parasites may evolve.

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The desert mistletoe is not hard to spot, especially during the driest parts of the year when most of its host trees have shed their leaves. It looks like a leafless tangled mass of pendulous stems sitting among the branches of larger shrubs and trees. It can be found growing throughout both the Mojave and Sonoran deserts and appears to prefer leguminous trees including palo verde (Parkinsonia florida), mesquite (Prosopis spp.), and Acacia.

The desert mistletoe is a type of hemiparasite, which means it is capable of performing photosynthesis but nonetheless relies on its host tree for water and other nutrients. Lacking leaves, the desert mistletoe meets all of its photosynthetic needs via its green stems. Its leafless habit also makes its flowers and fruit all the more conspicuous. Despite their small size, its flowers are really worth closer inspection. When in bloom, a desert mistletoe comes alive with the hum of various insects looking for energy-rich nectar and pollen. Even before you spot them, you can easily tell if there is a blooming mistletoe nearby as the flowers give off a wonderfully sweet aroma. It appears that the desert mistletoe takes no chances when it comes to reproduction in such an arid climate.

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As I mentioned above, the desert mistletoe has been the subject of inquiry over the last few decades. Researchers interested in how parasitic plants evolve have illuminated some intriguing aspects of the biology of this species, especially as its relates to host preference. It would appear that our interest in this species seems to be situated at an important time in its evolutionary history. Not all populations of desert mistletoe "behave" in the same way. In fact, each seems to be heading towards more intense specialization based on its preferred host.

By performing seed transplant experiments, researchers have demonstrated that various populations of desert mistletoe seem to be specializing on specific tree species. For instance, when seeds collected from mistletoe growing on Acacia were placed on paleo verde or mesquite, they experienced significantly less germination than if they were placed on another Acacia. Though the exact mechanisms aren't clear at this point in time, evidence suggests that the success of desert mistletoe may be influenced by various hormone levels within the host tree, with isolated populations becoming more specialized on the chemistry of their specific host in that region.

Speaking of isolation, there is also evidence to show that populations of desert mistletoe growing on different host trees are reproductively isolated as well. Populations growing on mesquite trees flower significantly later than populations growing on Acacia or palo verde. Essentially this means that their genes never have the chance to mix, thus increasing the differences between these populations. Again, it is not entirely certain how the host tree may be influencing mistletoe flowering time, however, hormones and water availability are thought to play a role.

Another intriguing idea, and one that has yet to be tested, are the roles that seed dispersers may play out in this evolutionary drama. After pollination, the desert mistletoe produces copious amounts of bright red berries that birds find irresistible. Two birds in particular, the northern mockingbird and the Phainopepla, aggressively defend fruiting mistletoe shrubs within their territories. It could be possible that these birds may be influencing which trees the seeds of the desert mistletoe end up on. Again, this is just a hypothesis but one that certainly deserves more attention.

A Phainopepla on the lookout for mistletoe berries.

A Phainopepla on the lookout for mistletoe berries.

Love them or hate them, there is something worth admiring about mistletoes. At the very least, they are important components of their native ecology. What's more, species like the desert mistletoe have a lot to teach us about the way in which species interact and what that means for biodiversity.

Photo Credit: [1]

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

Daffodil Insights

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

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

Narcissus poeticus

Narcissus poeticus

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

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

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

Narcissus tazetta

Narcissus tazetta

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

Narcissus cyclamineus

Narcissus cyclamineus

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

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

Narcissus triandrus

Narcissus triandrus

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

Narcissus viridiflorus  - a green, fall-blooming daffodil

Narcissus viridiflorus - a green, fall-blooming daffodil

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

The endangered  Narcissus alcaracensis .

The endangered Narcissus alcaracensis.

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

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

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

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

 

The Wild World of the Creosote Bush

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Apart from the cacti, the real rockstar of my Sonoran experience was the creosote bush (Larrea tridentata). Despite having been quite familiar with creosote as an ingredient, I admit to complete ignorance of the plant from which it originates. Having no familiarity with the Sonoran Desert ecosystem, I was walking into completely new territory in regard to the flora. It didn’t take long to notice creosote though. Once we hit the outskirts of town, it seemed to be everywhere.

If you are in the Mojave, Sonoran, and Chihuahuan Deserts of western North America, you are never far from a creosote bush. They are medium sized, slow growing shrubs with sprays of compact green leaves, tiny yellow flowers, and fuzzy seeds. Apparently what is thought of as one single species is actually made up of three different genetic populations. The differences between these has everything to do with chromosome counts. Populations in the Mojave Desert have 78 chromosomes, Sonoran populations have 52 chromosomes, and Chihuahuan have 26. This may have to do with the way in which these populations have adapted to the relative amounts of rainfall each of these deserts receive throughout the year, however, it is hard to say for sure.

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Regardless, creosote is supremely adapted to these xeric ecosystems. For starters, their branching architecture coupled with their tiny leaves are arranged so as to make the most out of favorable conditions. If you stare at these shrubs long enough, you may notice that their branches largely orient towards the southeast. Also, their leaves tend to be highly clustered along the branches. It is thought that this branching architecture allows the creosote to minimize water loss while maximizing photosynthesis.

Deserts aren’t hot 24 hours per day. Night and mornings are actually quite cool. By taking advantage of the morning sun as it rises in the east, creosote are able to open their stomata and commence photosynthesis during those few hours when evapotranspiration would be at its lowest. In doing so, they are able to minimize water loss to a large degree. Although their southeast orientation causes them to miss out on afternoon and evening sun to a large degree, the benefits of saving precious water far outweigh the loss to photosynthesis. The clustering of the leaves along the branches may also reduce overheating by providing their own shade. Coupled with their small size, this further reduces heat stress and water loss during the hottest parts of the day.

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Creosote also secrets lots of waxy, resinous compounds. These coat the leaves and to some extent the stems, making them appear lacquered. It is thought that this also helps save water by reducing water loss through the leaf cuticle. However, the sheer diversity of compounds extracted from these shrubs suggests other functions as well. It is likely that at least some of these compounds are used in defense. One study showed that when desert woodrats eat creosote leaves, the compounds within caused the rats to lose more water through their urine and feces. They also caused a reduction in the amount of energy the rats were able to absorb from food. In other words, any mammal that regularly feeds on creosote runs the risk of both dehydration and starvation. This isn’t the only interesting interaction that creosote as with rodents either. Before we get to that, however, we first need to discuss roots.

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Creosote shrubs have deep root systems that are capable of accessing soil water that more shallowly rooted plants cannot. This brings them into competition with neighboring plants in intriguing ways. When rainfall is limited, shallowly rooted species like Opuntia gain access to water before it has a chance to reach deeper creosote roots. Surprisingly this happens more often than you would think. The branching architecture of creosote is such that it tends to accumulate debris as winds blow dust around the desert landscape. As a result, the soils directly beneath creosote often contain elevated nutrients. This coupled with the added shade of the creosote canopy means that seedlings that find themselves under creosote bushes tend to do better than seedlings that germinated elsewhere. As such, creosote are considered nurse plants that actually facilitate the growth and survival of surrounding vegetation. So, if recruitment and resulting competition from vegetation can become such an issue for long term creosote survival, why then do we still so much creosote on the landscape?

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The answer may lie in rodents and other burrowing animals in these desert ecosystems. Take a look at the base of a large creosote and you will often find the ground littered with burrows. Indeed, many a mammal finds the rooting zone of the creosote shrub to be a good place to dig a den. When these animals burrow under shallowly rooted desert plants, many of them nibble on and disturb the rooting zones. Over the long-term, this can be extremely detrimental for the survival of shallow rooted species. This is not the case for creosote. Its roots run so deep that most burrowing animals cannot reach them. As such, they avoid most of the damage that other plants experience. This lends to a slight survival advantage for creosote at the expense of neighboring vegetation. In this way, rodents and other burrowing animals may actually help reduce competition for the creosote.

Barring major disturbances, creosote can live a long, long time. In fact, one particular patch of creosote growing in the Mojave Desert is thought to be one of the oldest living organisms on Earth. As creosote shrubs grow, they eventually get to a point in which their main stems break and split. From there, they begin producing new stems that radiate out in a circle from the original plant. These clones can go on growing for centuries. By calculating the average growth rate of these shrubs, experts have estimated that the Mojave specimen, affectionately referred to as the “King Clone,” is somewhere around 11,700 years old!

The ring of creosote that is King Clone.

The ring of creosote that is King Clone.

For creosote, its slow and steady wins the race. They are a backbone of North American desert ecosystems. Their structure offers shelter, their seeds offer food, and their flowers support myriad pollinators. Creosote is one shrub worthy of our respect and admiration.

Photo Credit: [1] [2]

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

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]