Zoophagous Liverworts?

Photo by Matt von Konrat Ph.D - Biblioteca Digital Mundial (eol.org) licensed under CC BY 3.0

Mention the word "liverwort" to most folks and you are going to get some funny looks. However, mention it to the right person and you will inevitably be drawn into a world of deep appreciation for this overlooked branch of the plant kingdom. The world of liverworts is best appreciated with a hand lens or microscope.

A complete lack of vascular tissue means this ancient lineage is often consigned to humid nooks and crannies. Look closely, however, and you are in for lots of surprises. For instance, did you know that there are liverworts that may be utilizing animal traps?

Right out of the gates I need to say that the most current research does not have this labelled as carnivorous behavior. Nonetheless, the presence of such derived morphological features in liverworts is quite sensational. These "traps" have been identified in at least two species of liverwort, Colura zoophaga, which is native to the highlands of Africa, and Pleurozia purpurea, which has a much wider distribution throughout the peatlands of the world.

A liverwort “trap” showing the lid (L), water sac (wl). {SOURCE]

The traps are incredibly small and likely derived from water storage organs. What is different about these traps is that they have a moveable lid that only opens inward. In the wild it is not uncommon to find these traps full of protozoans as well as other small microfauna. Researchers aimed to find out whether or not this is due to chance or if there is some active capture going on.

[SOURCE]

Using feeding experiments it was found that some protozoans are actually attracted to these plants. What's more these traps do indeed function in a similar way to the bladders of the known carnivorous genus Utricularia. Despite these observations, no digestive enzymes have been detected to date. For now researchers are suggesting that this is a form of "zoophagy" in which animals lured inside the traps die and are broken down by bacterial communities. In this way, these liverworts may be indirectly benefiting from the work of the bacteria.

This is not unheard of in the plant world. In fact, there are many species of pitcher plants that utilize similar methods of obtaining valuable nutrients. Certainly the lack of nutrients in the preferred habitats of these liverworts mean any supplement would be beneficial.

Photo Credits: Matt von Konrat Ph.D - Biblioteca Digital Mundial (eol.org), HESS ET AL. 2005 (http://www.bioone.org/doi/abs/10.1639/6), and Sebastian Hess (http://virtuelle.gefil.de/s-hess/forsch.html)

Further Reading: [1]

Going Veg With Nepenthes ampullaria

Photo by Bernard DUPONT licensed under CC BY-SA 2.0

Carnivory in the plant kingdom is an interesting evolutionary adaptation to living in nutrient poor environments. It has arisen in only a handful of different plant families and indeed, the genera that exhibit it are considered highly derived. There is something to be said about a sessile organism that can take down mobile prey at the rate that most carnivorous plants do.

Perhaps part of our fascination with these botanical wonders stems from their move towards dietary habits not unlike our own. The reason for their predatory behavior is to acquire nutrients like nitrogen and phosphorus. Without these essential nutrients, life as we know it would not exist. It is no wonder then that carnivorous plants have evolved some very interesting ways of getting them into their tissues and to me, there is nothing more peculiar than the way in which Nepenthes ampullaria gets its much needed nitrogen fix.

A rather widespread species, N. ampullaria is at home in the understory of the rain forests of the southeast Asian islands. It differs from its carnivorous cousins in a multitude of ways. For starters, the pitchers of N. ampullaria are oddly shaped. Resembling an urn, they sit in dense clusters all over the jungle floor, below the rest of the plant. Unlike other Nepenthes, the pitchers have only a small, vestigial lid with no nectar glands. Finally, the slippery, waxy surface that normally coats the inside of most Nepenthes pitchers is absent in the pitchers of N. ampullaria. All of these traits are clues to the unique way in which this species has evolved to acquire nitrogen.

N. ampullaria doesn't lure and digest insects. Instead, it relies on leaf litter from the forest canopy above for its nutritional needs. The urn-like shape, lack of a hood, and clustered growth enable the pitchers to accumulate considerable amounts of leaf litter in the pitchers. Because the pitchers are relatively long lived for a Nepenthes, lasting upwards of 6 months, they offer up a nice microhabitat for a multitude of insect and even frog larvae. The collective group of organisms living within the pitchers are referred to as an inquiline community.

Over time, an inquiline community develops in each of the pitchers. This is the key to the success of N. ampullaria. As the inquiline organisms breakdown the leaf litter, they release copious amounts of nitrogen-rich waste. The pitchers can then absorb this waste and begin to utilize it. At least one study found that an individual plant can obtain 35.7% of its foliar nitrogen in this manner. It has also been demonstrated that the pitchers actively manipulate the pumping of hydrogen ions into the fluid within to keep it less acidic than that of other Nepenthes.

I don't know if I would consider this a case of herbivory as the nitrogen is still coming from an animal source but it is nonetheless an interesting adaptation. Instead of using valuable resources on actively digesting its own prey, N. ampullaria is getting other organisms to do the work for it. Not too shabby.

Further Reading:

http://bit.ly/1IRbYG9

http://jxb.oxfordjournals.org/content/61/5/1365

http://link.springer.com/article/10.1007/s004420050390

http://bit.ly/1S10oej

The Fanged Pitcher Plant of Borneo

As mammals, and even more so as apes, we tend to associate fangs with threats. The image of two dagger-like teeth can send chills up ones spine. Perhaps it is fitting then that a carnivorous plant from a southeast Asian island would sport a pair of ominous fangs. Friends, I present to you the bizarre fanged pitcher plant (Nepenthes bicalcarata).

This ominous-looking species is endemic to Borneo and gets its common name from the pair of "fangs" that grow from the lid, just above the mouth of the pitcher. Looks aren't the only unique feature of this species though. Indeed, the entire ecology of the fanged pitcher plant is fascinatingly complex.

Lets tackle the obvious question first. What is up with those fangs? There has been a lot of debate among botanists as to what function they might serve. Some have posited the idea that they may deter mammals from feeding on pitcher contents. Others see them as mere artifacts of development and attribute no function to them whatsoever.

In reality they are involved in capturing insects. The fangs bear disproportionately large nectaries that lure prey into a precarious position just above the mouth of the pitcher. Strangely enough, this may have evolved to compensate for the fact that the inside of the pitchers are not very slippery. Whereas other pitcher plant species rely on waxy walls to make sure prey can't escape, the fanged pitcher plant has relatively little waxy surface area within its pitchers. What's more, the pitchers are not very effective at capturing prey unless they have been wetted by rain. The fluid within the pitchers also differs from other Nepenthes in that it is not very acidic, contains few digestive enzymes, and isn't very viscous. Why?

Worker ants cleaning the pitcher (left) and an ant brood chamber inside of the pitcher tendril (right). Photo by Bazile, V., J.A. Moran, G. Le Moguédec, D.J. Marshall & L. Gaume 2012. A carnivorous plant fed by its ant symbiont: a unique multi-faceted nutritional mutualism. PLoS ONE 7(5): e36179. doi:10.1371/journal.pone.0036179 licensed under CC BY 2.5

The answer lies with a specific species of ant. The fanged pitcher plant is the sole host of a carpenter ant known scientifically as Camponotus schmitzi. The tendrils that hold the pitchers themselves are hollow and serve as nest sites for these ants. Ant colonies take up residence in the tendrils and will hunt along the insides of the pitchers. In fact, they literally go swimming in the pitcher fluid to find their meals!

This is why the pitcher fluid differs so drastically from other Nepenthes. The fanged pitcher plant actually does very little of its own digestion. Instead, it relies on the resident ant colony to subdue and breakdown large prey. As a payment for offering the ants room and board, the ants help the plant feed via the breakdown of captured insects (which are often disposed of in the pitchers) and the deposition of nitrogen-rich feces. Indeed, plants without a resident ant colony are found to be significantly smaller and produce fewer pitchers than those with ants. The ants also protect and clean the plant, removing fungi and hungry insect pests.

Sadly, like many other species of Nepenthes, over-harvesting for the horticultural trade as well as habitat destruction have caused a decline in numbers in the wild. With species like this it is so important to make sure you are buying nursery grown specimens. Never buy a wild collected plant! Also, if you are lucky enough to grow these plants, propagate them! Only by reducing the demand for wild specimens can we hope of curbing at least some of the poaching threats. Also, what better way to get your friends into gardening than by sharing with them amazing carnivores like the fanged pitcher plant.

Female flowers

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

A Circumboreal Butterwort

The name "butterwort" may sound quite silly to most but those who have seen one in person can attest to the fact that there is nothing silly about the group of plants. Hailing from the genus Pinguicula, my favorite butterwort is Pinguicula vulgaris.

Referred to as the common butterwort, this species is sometimes hard to find if you live in North America. It has a circumboreal distribution and seems to be much more common in Europe and parts of Russia. Like all butterworts, P. vulgaris is a carnivore, though at first glance this may not be very obvious. The fleshy rosette of leaves are covered in mucilaginous glands that trap hapless insects. The leaves will sometimes roll in along the edge to pool the digestive juices around their prey.

Unlike more familiar carnivorous plants that can be found in acidic soils, P. vulgaris is a calciphile. It is most often encountered in fens, alvars, and other areas with limestone bedrock and alkaline waters. These types of habitats pose a different set of challenges for plants when it comes to obtaining nutrients. Phosphorus strongly binds to sediments in these alkaline conditions and research has shown that most butterworts respond best to supplemental phosphorus additions, though other nutrients like nitrogen are absorbed from prey as well.

If their carnivorous habits weren't interesting enough, the flowers of P. vulgaris (and all butterworts for that matter) are gorgeous. Sitting atop long stalks, the spurred blooms are a deep shade of violet. The nectar spur suggests that this species is pollinated by either long tongued bees or butterflies. Either way, they are presented well above the sticky leaves to reduce the chances of the plant eating the insects it needs for pollination.

Like all plants in the northern hemisphere, P. vulgaris needs to deal with winter. As temperatures and light levels begin to drop, P. vulgaris reverts to a cluster of buds called a hibernaculum. It has little to no roots during its dormant phase and can easily blow around if exposed. This may serve to transport plants into new locations. Due to rampant habitat destruction, this plant is quite vulnerable in North America and is considered threatened or endangered in the southern portions of its range. Please, if you know of a land conservancy or some other agency that protects bogs, alvars, and fens, show them support!

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

An Introduction to Cephalotus follicularis - A Strange Carnivore From Australia

Photo by H. Zell licensed under CC BY-SA 3.0

In a small corner of western Australia grows a truly unique carnivorous plant. Commonly referred to as the Albany pitcher plant, Cephalotus follicularis is, evolutionarily speaking, distinct among the pitcher plants. It is entirely unrelated to both the Sarraceniaceae and the Nepenthaceae.

This stunning case of convergent evolution stems from similar ecological limitations. Cephalotus grows in nutrient poor areas and thus must supplement itself with insect prey. It does so by growing modified leaves that are shaped into pitchers. The lid of each pitcher has two main functions. It keeps rain from diluting the digestive enzymes within and it also confuses insects.

A close inspection of the lid will reveal that it is full of clear spots. These spots function as windows, allowing light to penetrate, which confuses insects that have landed on the trap. As they fly upwards into the light, they crash into the lid and fall back down into the trap.

Photo by Lucas Arrrrgh licensed under CC BY-NC-ND 2.0

The relationship of Cephalotus to other plants has been the object of much scrutiny. Though it is different enough to warrant its own family (Cephalotaceae), its position in the greater scheme of plant taxonomy originally had it placed in Saxifragales. Genetic analysis has since moved it out of there and now places it within the order Oxalidales. What is most intriguing to me is that the closest sister lineage to this peculiar little pitcher plant are a group of trees in the family Brunelliaceae. Evolution can be funny like that.

Regardless of its relationship to other plants, Cephalotus follicularis has gained quite a bit of attention over the last few years. Its strange appearance and carnivorous habit have earned it a bit of stardom in the horticultural trade. A single specimen can fetch a hefty price tag. As a result, collecting from wild populations has caused a decline in numbers that are already hurting due to habitat destruction. Luckily they are easy to culture in captivity, which will hopefully take pressure off of them in the wild.

What's more, the loss of Cephalotus from the wild is hurting more than just the plant. A species of flightless, ant-mimicking fly requires Cephalotus pitchers to rear its young. They don't seem to mind growing up in the digestive enzymes of the pitchers and to date, their larvae have been found living nowhere else. If you are lucky enough to grow one of these plants, share the wealth. Captive reared specimens not only take pressure off wild populations, they are also much hardier. Lets keep wild Cephalotus in the wild!

Photo by Holger Hennern licensed under CC BY-SA 3.0

Photo Credits: Holger Hennern (Wikimedia Commons) and Lucas Arrrrgh (https://www.flickr.com/photos/chug/2121092119/)

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

New Plant Species Discovered on Facebook

Photo by Paulo Gonella licensed under CC BY-SA 3.0

There are many downsides to the amount of time some of us spend on the internet but there is no denying that there are some incredible benefits as well. Never before in human history has information been so readily available to so many people. Without Facebook, In Defense of Plants would not have anywhere near the platform from which I can interact with all of you wonderful plant folk. In what may be one of the coolest uses of social media to date, a new species of carnivorous plant has been discovered using Facebook! 

While exploring a mountain top in Brazil, amateur researcher  Reginaldo Vasconcelos snapped a few shots of a large sundew. Upon returning home, the pictures were uploaded to Facebook for the world to see. It didn't take long for scientists to notice that the plant in the picture was something quite special. 

Indeed, what Vasconcelos had photographed was a species of Drosera completely new to science! This is the first time that a new species has been discovered using social media. Experts have now published the first scientific description of this species. It has been named Drosera magnifica - the magnificent sundew. 

And magnificent it is! According to the authors of the paper, "It is the largest sundew in the Americas, and the second-largest carnivorous plant in the Americas. In this respect it is also a spectacular plant.” The plant was discovered in Minas Gerais, Brazil. Oddly enough, the mountain on which it was found is readily accessible. How this species went undiscovered for so long is quite a mystery. It just goes to show you how little we know about the world we live in. 

That sad part about this discovery is that the mountain it is endemic to is surrounded by cattle ranches as well as coffee and eucalyptus plantations. The future of this brand new species is by no means certain. Researchers have already elevated its status to critically endangered. Unless other populations are found, this species may disappear not long after its discovery. 

Photo Credit: Paulo Gonella

Further Reading:

http://www.mapress.com/phytotaxa/content/2015/f/p00220p267f.pdf

A New Look at a Common Bladderwort

Photo by Kevin Thiele licensed under CC BY 2.0

It is so often that common species are overshadowed by something more exotic. Indeed, we know more about some of the rarest plants on earth than we do about species growing in our own back yards. Every once in a while researchers break this pattern and sometimes this yields some amazing results. Nowhere has this been better illustrated in recent years than on the humped bladderwort, Utricularia gibba. 

This wonderful little carnivore can be found growing in shallow waters all over the world. Like all Utricularia, it uses tiny little bladders to capture its even tinier prey. Despite its diminutive size, U. gibba is nonetheless a very derived species. For all of its wonderful physical attributes, the real adventure begins at the microscopic level. As it turns out, U. gibba has some amazing genetic attributes that are shining light on some incredible evolutionary mechanisms. 

When researchers from the University at Buffalo, Universitat de Barcelona in Spain, and LANGEBIO in Mexico decided to sequence the genome of this plant, what they found was quite startling. For a rather complex little plant, the genome of U. gibba is incredibly small. What the researchers found is that U. gibba appears to be very efficient with its DNA. Let's back up for a moment and consider this fact. 

The genomes of most multicellular organisms contain both coding and non-coding DNA. For decades researchers have gone back and forth on how important non-coding DNA is. They do not code for any protein sequences but they may play a role in things like transcription and translation. For a long time this non-coding DNA has been referred to as junk DNA. 

This is where things get interesting. Sequencing of the U. gibba genome revealed that only 3% of its genome consisted of non-coding or junk DNA. For some reason the U. gibba lineage has managed to delete most of it. To put things in perspective, the human genome is comprised of roughly 98% non-coding or junk DNA. Despite its rather small and efficient genome, U. gibba nonetheless has more genes than plants with larger genomes. This may seem confusing but think of it this way, whereas U. gibba has a smaller overall genetic code, it is comprised of more genes that code for things like digestive enzymes (needed for digesting prey) and cell walls (needed to keep water out) than plants with more overall genetic code such as grapes or Arabidopsis. 

As one author put it, this tiny ubiquitous plant has revealed "a jewel box full of evolutionary treasures." It is a species many of us have encountered time and again at the local fishing hole or in your favorite swimming pond. Time and again we pass by the obvious. We overlook those organisms that are most familiar to us. We do so at the cost of so much knowledge. It would seem that the proverbial "Old Dog" has plenty of tricks to teach us. 

Photo Credit: Kevin Thiele (http://bit.ly/1Flouqd) and Reinaldo Aguilar (http://bit.ly/1B6mnHN)

Further Reading:

http://www.nature.com/nature/journal/v498/n7452/full/nature12132.html

http://mbe.oxfordjournals.org/content/early/2015/01/31/molbev.msv020

http://plants.usda.gov/core/profile?symbol=UTGI

Slippery When Wet

Photo by Andrea Schieber licensed under CC BY-NC-ND 2.0

Pitcher plants in the genus Nepenthes have been getting a lot of attention in the literature as of late. Not only have researchers discovered the use of ultraviolet pigments around the rims of their pitchers, it has also been noted that the pitchers of many species aren't as slippery as we think they are. Indeed, scientists have noted that prey capture is at its highest only when the pitchers are wet. This seems counterintuitive. Why would a plant species that relies on the digestion of insects for most of its nitrogen and phosphorus needs produce insect traps that are only effective at certain times? After all, it takes a lot of energy for these plants to produce pitchers, which give little to nothing back in the way of photosynthesis. 

The answer to this peculiar conundrum may lie in the types of insects these plants are capturing. Ants are ubiquitous throughout the world. Their gregarious and exploratory nature has provided ample selection pressures for much of the plant kingdom. They are particularly well known for their military-esque raiding parties. It is this behavior that researchers have looked at in order to explain the intermittent effectiveness of Nepenthes pitchers. 

A recent study that looked at Nepenthes rafflesiana found that ants made up 65% of the prey captured, especially on pitchers produced up in the canopy. What's more, younger pitchers produced closer to the ground were found to be much more slippery (containing more waxy cells) than those produced farther up on the plant. When the pitchers of this species were kept wet, prey capture consisted mostly of individual insects such as flies. However, when allowed to dry between wettings, the researchers found that prey capture, specifically ants, increased dramatically. How is this possible?

It all goes back to the way in which ants forage. A colony sends out scouts in all directions. Once a scout finds food, it lays down a pheromone trail that other ants will follow. It is believed that this is the very behavior that Nepenthes are relying on. The traps produce nectar as a lure for their insect prey. As the traps dry up, the nectar becomes concentrated. Ants find this sugary treat irresistible. However, if the pitcher were to be slippery at all times, it is likely that most ant scouts would be killed before they could ever report back to the colony. By reducing the slippery waxes, especially around the rim of the trap, the Nepenthes are giving the ants a chance to "spread the news" about this new food source. Because these plants grow in tropical regions, humidity and precipitation can fluctuate wildly throughout a 24 hour period. If the scouting party returns at a time in which the pitchers are wet then the plant stands to capture far more ants than it did if it had only caught the scout. 

This is what is referred to as batch capture. The plants may be hedging their bets towards occasional higher nutrient input than constant low input. This is bolstered by the differences between pitchers produced at different points on the plant. Lower pitchers, especially on younger plants are far more waxy and thus are constantly slippery. This allows constant prey capture to fuel rapid growth into the canopy. Upper pitchers on older individuals want to maximize their yields via this batch capture method and therefore produce fewer waxy cells, relying on a humid climate to do the work for them. It is likely that this is a form of tradeoff which benefits different life cycle stages for the plant. 

Photo Credit: Andrea Schieber (http://bit.ly/1xUsGJk)

Further Reading:

http://rspb.royalsocietypublishing.org/content/282/1801/20142675

An Unlikely Carnivore from Brazil

There is something about a plant eating an animal that makes it all the more beautiful. Indeed, even without carnivory, the beauty of these highly adapted plant species is up there with the fanciest of horticultural creations. Nowhere is this fact more apparent than in a recently described species of plant from Brazil. Philcoxia minensis has only been known to science since 1981 and the way in which it has adapted to catching its prey is rather incredible.

Native to montane savannas of southeastern Brazil, these delicate members of the family Scrophulariaceae grow in nutrient poor, sandy habitats. They seem rather delicate with only a thin flowering stalk readily apparent above the soil line. The botanists who first discovered this species made an interesting observation, the leaves of this plant grow mostly underground. Though they do contain functional chloroplasts, it was initially assumed that they served little function. 

As it turns out, the leaves can in fact photosynthesize. The white sands that these plants grow in likely permit just enough light to penetrate down into the clear grains, allowing photosynthesis to happen. However, this is not the strangest thing about Philcoxia. Since this genus was first described, botanists have taken interest in the glandular stalks on the leaves. They noted that they were quite sticky and hypothesized that Philcoxia minensis may be carnivorous. 

Initial investigations into this were not conclusive. Recently, however, this has changed. A study published in 2012 used special nitrogen markers to look deeper into the potential carnivorous habits of this plant. What they found was quite amazing. As it turns out, there is very strong evidence that Philcoxia minensis uses its sticky traps to capture and eat nematodes under the soil. Not only is Philcoxia new to science, it also offers us yet another view into the many different ways plants have evolved to obtain precious nutrients in harsh environments. 

Photo Credit: Caio G. Pereiraa, Daniela P. Almenarab, Carlos E. Winterb, Peter W. Fritschc, Hans Lambersd, and Rafael S. Oliveira

Further Reading:
http://www.pnas.org/content/109/4/1154.abstract

http://www.jstor.org/discover/4117770?sid=21105606841053&uid=2