A Poop-Loving Moss Discovered Living on Poop-Eating Pitcher Plants

Poop mosses are strange to say the least. They hail from the family Splachnaceae and most live out their entire (short) lives growing on poop. Needless to say, they are fascinating plants. Recently, one species of poop moss known to science as Tayloria octoblepharum was discovered growing in Borneo for the first time. As if this range expansion wasn’t exciting enough, their growing location was very surprising. Populations of this poop-loving moss were found growing in the pitchers of two species of poop-eating pitcher plants in the genus Nepenthes!

The pitcher of Nepenthes lowii both look and function like a toilet bowl. Photo by JeremiahsCPs licensed under the GNU Free Documentation License

The wide pitcher mouth of Nepenthes macrophylla offer a nice seating area for visiting tree shrews.

The pitchers of both Nepenthes lowii and N. macrophylla get a majority of their nutrient needs not by trapping and digesting arthropods but instead from the feces of tree shrews. They have been coined toilet pitchers as they exhibit specialized adaptations that allow them to collect feces. Tree shrews sit on the mouth of the pitcher and lap up sugary secretions from the lid. As they eat, they poop down into the pitcher, providing the plant with ample food rich in nitrogen. Digestion is a relatively slow process so much of the poop that enters the pitcher sticks around for a bit.

During a 2013 bryophyte survey in Borneo, a small colony of poop moss was discovered growing in the pitcher of a N. lowii. This obviously fascinated botanists who quickly made the connection between the coprophagous habits of these two species. On a return trip, more poop moss was discovered growing in a N. macrophylla pitcher. This population was fertile, indicating that it was able to successfully complete its life cycle within the pitcher environment. It appears that these two toilet pitchers offer ample niche space for this tiny, poop-loving moss. If this doesn’t convince you of just how incredible and complex the botanical world is, I don’t know what will!

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

Further Reading: [1]

Gooey Pitcher Fluids

Photo by Shawn Mayes licensed under CC BY-SA 3.0

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.

Photo by Alfindra Primaldhi licensed under CC BY 2.0

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 most 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. Photo by Shawn Mayes licensed under CC BY-SA 3.0

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. Photo by Alfindra Primaldhi licensed under CC BY 2.0

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. Photo by Alfindra Primaldhi licensed under CC BY 3.0

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]

Crab Spiders and Pitcher Plants: A Dynamic Duo

Most pitcher plants in the genus Nepenthes seem pretty adept at catching prey. These plants specialize in nutrient-poor soils and their carnivorous habit evolved as a means of supplementing their nutritional needs. Despite the highly evolved nature of their pitfall traps (which are actually modified leaves), Nepenthes aren’t perfect killing machines. In fact, some get a helping hand from seemingly unlikely partners.

Spend enough time reading about Nepenthes in the wild and you will see countless mentions of arthropods hanging around their pitchers. Some of these inevitably become prey, however, there are others that appear to be taking advantage of the plant. Nepenthes don’t passively trap arthropods. Instead, they lure them in with bright colors and the promise of tasty treats like nectar. This is not lost on predators like spiders, who are frequent denizens of pitcher mouths.

Most notable to Nepenthes specialists are some of the crab spiders that frequently haunt Nepenthes traps. These wonderful arachnids sit at the mouth of the pitcher and ambush any insects that try to pay it a visit. Often times both predator and prey fall down into the pitcher, however, thanks to a strand of silk, the spiders easily climb back out with their meal. This may seem like bad news for the pitcher, however, recent research based out of the National University of Singapore has shown that this relationship is not entirely one sided.

By studying the interactions between spiders and pitcher plants both in the lab and in the field, ecologists discovered that at least one species of pitcher plant (Nepenthes gracilis) appears to benefit greatly from the presence of crab spiders. The key to understanding this relationship lies in the types of prey N. gracilis is able to capture when crab spiders are and are not present.

Not only did the presence of a resident crab spider increase the amount of prey in each Nepenthes pitcher, it also changed the types of insects that were being captured. Crab spiders are ambush predators that frequently attack prey much larger than themselves. It may seem as if this is a form of food robbery on the part of the crab spider but the spiders can’t eat everything. When they have eaten their fill, the spiders discard the carcass into the pitcher where the plant can make quick work digesting it for its own benefit.

Over time, simply having a spider hunting on the trap led to a marked increase in the number of insects in each pitcher compared to those without a spider. Even if these meals are already half eaten, the plant still gains nutrients. Additionally, the types of prey captured by pitchers with and without crab spiders changed. The spiders were able to capture and subdue insects like flesh flies, which normally aren’t captured by Nepenthes pitchers. As such, the resident crab spiders make available a larger suite of potential prey than would be available if they weren’t using the pitchers as hunting grounds.

The crab spiders may benefit the pitcher plant in other ways as well. Research on crab spiders has shown that their bodies are covered in pigments that register high in the UV spectrum. Insects can see UV light and often use it as a means of finding flowers as plants often produce UV-specific pigments in their floral tissues. The wide array of UV patterns on flowers are there to guide their pollinators into position. Researchers have documented that insects are actually more likely to visit flowers with crab spiders than those without, which has led to the idea that UV pigments in crab spiders actually act as insect attractants. Visiting insects simply cannot resist the UV stimulus and quickly fall victim to the resident crab spider.

Could it be that by taking up residence on a Nepenthes pitcher, the crab spiders are increasing the likelihood of insects visiting the traps? This remains to be seen as such questions did not fall under the scope of this investigation. That being said, it certainly offers tantalizing evidence that there is more to the Nepenthes-crab spider relationship. More work is needed to say for sure but the closer we look at such interactions, the more spectacular they become!

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

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

Not All Pitchers Are Equal: How Prey Capture Has Driven Speciation in the genus Nepenthes

Source

Species of the genus Nepenthes are as bizarre as they are beautiful. Known the world around for their carnivorous lifestyle, these plants looks like something out of a macabre art exhibit. It is easy to get caught up in this beauty. I often find myself lost in thought while staring at full grown specimen. How did this genus come to be? Why are they so diverse? What is going on with the morphology of these plants?

Nepenthes hail from nutrient poor habitats, which has driven them to supplement their growth with nutrients gained via the breakdown of a variety of organisms. The business ends of a Nepenthes are their pitchers. We get so caught up in the bewildering diversity of shapes, colors, and sizes that we often overlook them as the anatomical marvels of evolution that they truly are. Whereas the main body of these plants often look quite similar among different species, it's the pitchers that really allow us to separate them out as distinct species. Pitcher morphology not only gives us a convenient means to identify these plants, research is now showing that the structure of these pitchers is likely to be the driving force in their evolution. 

Source

Let's back up for a second. Before we get to the subject of adaptive radiation, we should take a closer look at the anatomy of these plants. To put it simply, the pitchers of Nepenthes are actually leaves, albeit highly modified versions. What we readily recognize as the photosynthetic leaves of a Nepenthes plant are actually modified leaf bases or petioles. Over evolutionary time, these bases have flattened to increase the amount of surface area available for photosynthesis.

From the tip of each of these "leaves" is produced a tendril. Gradually this tendril will elongate and the tip starts to swell. This tip will eventually become the pitcher. The pitchers themselves are highly modified leaves. They are some of the most specialized leaves in all of the plant kingdom. As the tip grows larger, it becomes clear that there is a distinctive lid apparatus. Once the pitcher is fully mature, this lid pops open revealing the death trap filled with digestive fluids.

Source

As if producing pitchers wasn't cool enough, each species of Nepenthes produces two distinct forms - lower pitchers, which are produced by young plants as well as on mature plants near the ground, and upper pitchers, which are produced up on the climbing stems as they vine through the canopy. The upper and lower pitchers look radically different from one another to the point that one may easily confuse them for different species. The reason for such stark differences has to do with the type of prey captured. Lower pitchers are generally larger and can capture prey that crawls along the forest floor. Upper pitchers tend to be more slender and most often capture flying insects as well as other creepy crawlies hanging out in the forest canopy.

The key to the success of these traps seems pretty straight forward - insects attracted by bright colors and sweet nectar land on the traps and fall to their death. Certainly this holds true throughout the genus, however, there are at least two major variations on this theme and a handful of bizarre mishmashes. As the lid of a Nepenthes pitcher starts to open, a ring of tissue called the peristome unfurls. The shape and color varies wildly between species and this has to do with the methods in which they capture their prey. These variations are the key to the amazing diversity of Nepenthes we see throughout the range of this genus.

Nepenthes vogelii

The first of the three strategies is referred to as the 'insect aquaplaning' strategy. Insects walking around on the peristome of the pitcher find it hard to get a foothold. These are species such as N. raja, N. ampullaria, and N. bicalcarata (just to name a few). The slipperiness of the peristome of these species is further enhanced when humidity is high. Considering how much it rains in these habitats, it is no wonder why capture efficiency is often as high as 80%. Although there is some variation on this theme, pitchers that utilize the insect aquaplaning strategy often lack waxy cells on the interior of the pitcher walls.

Slippery pitcher walls are the second strategy that Nepenthes have converged upon. These are species such as N. diatas, N. mirabilis, and N. alata (again, just to name a few) Insects attracted to the pitchers are often lured in by sweet nectar. Once they cross the lip of the pitcher, prey find it hard to hang on and inevitably fall inside. Once this happens, waxy cells lining the interior walls make it impossible for anything to climb back out. It should be mentioned that a slippery peristome and waxy pitcher walls are not mutually exclusive. That being said, there are clear trends among species that show a reduction in waxy cells as peristome size and slope increases.

This brings us to the oddballs. There are species like N. lowii, whose pitchers function as a toilet bowl for shrews, and N. aristolochioides, whose pitchers seemed to have abandonded both strategies and now function as light traps similar to what we see in Darlingtonia. Regardless of their strategy, the diversity in trapping mechanisms appear to be the driving force behind the bewildering diversity of Nepenthes. 

Nepenthes aristolochioides

All of the evidence taken together shows that prey capture is at the core of this radiation. There seems to be incredibly strong selective pressures that result in strong divergence in pitcher morphology. The disruptive selection that seems to be driving a wedge between the insect aquaplaning strategy and the waxy wall strategy may have its roots in reducing competition. Nutrients are low and competition for food is high. Different Nepenthes species could be evolving to capture different kinds of prey. Even closely related species such as N. ampullaria, N. rafflesiana, N. mirabilis, N. albomarginata, and N. gracilis all seem to occupy their own unique spot on the spectrum of prey capture strategy.

It could also be that Nepenthes are responding to the specific characteristics of the habitats in which they are found. Those inhabiting drier sites may favor the waxy wall strategy whereas those living in wetter habitats tend to favor the slippery peristome. More work needs to be done to investigate where and how these different strategies are maximized. Until then, I think it is safe to say that the diversity of this incredible genus has a lot to do with obtaining food. 

Photo Credits: [1] 

Further Reading:

[1] [2] [3]

A Digestive "On" Switch

Photo by Luiz licensed under CC BY-NC-ND 2.0

A common thread throughout the world of carnivorous plants is that all hail from nutrient poor environments. That is why they evolved carnivory in the first place, as a way of supplementing their nitrogen and phosphorous needs. For as amazing as their various adaptations are, the evolutionary histories of the world's carnivorous plants are still largely shrouded in mystery. A recent paper published in the Annals of Botany takes a closer look at what goes on inside the pitchers of the tropical pitcher plant Nepenthes alata. What they found is quite amazing.

As it turns out, N. alata seems to be able to regulate the amount of digestive enzymes within its pitchers based on prey availability. This makes a lot of sense. Since these species live in nutrient poor conditions, it would be very wasteful to continuously produce digestive fluids. Instead, the research team found that the genes responsible for the productive of digestive enzymes turn on in response to certain cues. In this case, its the presence of insect tissues, specifically chitin. The addition of insect prey coincided with a 24 to 48 hour burst in digestive enzyme production followed by a gradual decrease as the insects were digested. As interesting as this is, these were not the only findings to come out of this research.

When the researchers looked closely at what kinds of enzymes N. alata were producing, they discovered evidence in support of a long-held hypothesis regarding the evolution of carnivory in plants. The genetic pathways induced by the addition of insect chitin are nearly identical to those seen in plant defense pathways. These pathways also induced the production of a series of proteins known to play a role in plant defense reactions against microbial pathogens. What's more, many of the enzymes N. alata were producing inside their pitchers are classified as defense-related proteins. Taken together, this is strong evidence in support of the hypothesis that carnivory in plants evolved from defense reactions already in place.

This finding comes in the wake of an earlier discovery that showed similar pathways in the traps of the Venus fly trap. This is yet more evidence for the fact that evolution does not always occur via novel pathways. Instead, systems that are already in place are retooled to fit a new set of challenges.

Photo Credit: [1]

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]

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