A Passionflower With a Taste for Insects?

Passiflora_foetida_1.jpg

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

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

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

Passiflora_foetida_3.jpg

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

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

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

P_foetida_fruit.jpg

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

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

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

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

Further Reading: [1] [2]  

The Wild World of Rattan Palms

Calamus_gibbsianus.jpg

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

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

3070456167_a06111dd01_o.jpg

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

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

10368246_1018184728208392_2661304592123048565_n.jpg

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

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

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

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

Apocynaceae Ant House

10500247_1012300545463477_3608958353775216067_n.jpg

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

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

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

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

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

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

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

The flowers of   Dischidia ovata

The flowers of Dischidia ovata

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

Dischidia_platyphylla_kz1.jpg

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

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

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

Further Reading: [1]

How a Giant Parasitic Orchid Makes a Living

34281162165_b5ba814c84_b.jpg

Imagine a giant vine with no leaves and no chlorophyll scrambling over decaying wood and branches of a warm tropical forest. As remarkable as that may seem, that is exactly what Erythrorchis altissima is. With stems that can grow to upwards of 10 meters in length, this bizarre orchid from tropical Asia is the largest mycoheterotrophic plant known to science.

Mycoheterotrophs are plants that obtain all of their energy needs by parasitizing fungi. As you can probably imagine, this is an extremely indirect way for a plant to make a living. In most instances, this means the parasitic plants are stealing nutrients from the fungi that were obtained via a partnership with photosynthetic plants in the area. In other words, mycoheterotrophic plants are indirectly stealing from photosynthetic plants.

In the case of E. altissima, this begs the question of where does all of the carbon needed to build a surprising amount of plant come from? Is it parasitizing the mycorrhizal network associated with its photosynthetic neighbors or is it up to something else? These are exactly the sorts of questions a team from Saga University in Japan wanted to answer.

33905127355_fd4bd84026_b.jpg

All orchids require fungal partners for germination and survival. That is one of the main reasons why orchids can be so finicky about where they will grow. Without the fungi, especially in the early years of growth, you simply don't have orchids. The first step in figuring out how this massive parasitic orchid makes its living was to identify what types of fungi it partners with. To do this, the team took root samples and isolated the fungi living within.

By looking at their DNA, the team was able to identify 37 unique fungal taxa associated with this species. Most surprising was that a majority of those fungi were not considered mycorrhizal (though at least one mycorrhizal species was identified). Instead, the vast majority of the fungi associated with with this orchid are involved in wood decay.

Stems climbing on fallen dead wood (a) or on standing living trees (b). A thick and densely branched root clump (c) and thin and elongate roots (d) [Source]

Stems climbing on fallen dead wood (a) or on standing living trees (b). A thick and densely branched root clump (c) and thin and elongate roots (d) [Source]

To ensure that these wood decay fungi weren't simply partnering with adult plants, the team decided to test whether or not the wood decay fungi were able to induce germination of E. altissima seeds. In vitro germination trials revealed that not only do these fungi induce seed germination in this orchid, they also fuel the early growth stages of the plant. Further tests also revealed that all of the carbon and nitrogen needs of E. altissima are met by these wood decay fungi.

These results are amazing. It shows that the largest mycoheterotrophic plant we know of lives entirely off of a generalized group of fungi responsible for the breakdown of wood. By parasitizing these fungi, the orchid has gained access to one of the largest pools of carbon (and other nutrients) without having to give anything back in return. It is no wonder then that this orchid is able to reach such epic proportions without having to do any photosynthesizing of its own. What an incredible world we live in!

33905171165_05da1d498c_b.jpg

Photo Credits: [1] [2]

Further Reading: [1]

Resin Midges, Basal Angiosperms, and a Strange Pollination Syndrome

kadsura.JPG

When we try to talk about clades that are "basal" or "sister" to large taxonomic groups, your average listener either consciously or unconsciously thinks "primitive." Primitive has connotations of something that under-developed or unfinished. This is simply not the case. Take, for instance, a family of basal angiosperms called Schisandraceae.

This family is nestled within the order Austrobaileyales, which, along with a small handful of other families, represent the earliest branches of the angiosperm family tree still alive today.  To call them primitive, however, would be a serious misnomer. Because they diverged so early on, these lineages represent serious success stories in flowering plant evolution, having survived for hundreds of millions of years. Instead, we must think of them as fruitful early experiments in angiosperm evolution.

Floral morphology of and interaction between midge and their larvae (white arrows) in Illicium dunnianum

Still, the proverbial proof is in the pudding and if there was any sort of physical evidence one could put forth to remove our hierarchical prejudices about the taxonomic position of these plants, it would have to be their bizarrely specific pollination syndromes.  Members of the family Schisandraceae have entered into intense relationships with a group of flies known as midges and their interactions are anything but primitive. 

We will start with two species of plant native throughout parts of Asia. Meet as Illicium dunnianum and Illicium tsangii. More will be familiar with this genus than they may realize as Illicium gives us the dreaded star anise flavor our grandparents liked to sneak into our cookies as kids (but I digress). These particular species, however, have more to offer the world than flavoring. They are also very important plants for a group of gall midges in the genus Clinodiplosis.

The midges cannot reproduce without I. dunnianum or I. tsangii. You see, these midges lay their eggs within the flowers of these plants and, in doing so, end up pollinating them in the process. At first glance it may seem like a very one-sided relationship. Female midges deposit their eggs all along the carpels packed away inside large, fleshy whorl of tepals. As the midges crawl all over the reproductive organs looking for a suitable place to lay, they inevitably pick up and deposit pollen. 

Floral morphology and interaction between midge larvae (white arrows) in  Illicium tsangii

This is not the end of this relationship though. After eggs have been deposited, something strange happens to the Illicium flowers. For starters, they develop nursery chambers around the midge larvae. Additionally, their tepals begin producing heat. Enough heat is produced to keep the nursery chamber temperature significantly warmer than the ambient air temperature. What's more flower heating intensifies throughout the duration of fruit development. It was originally hypothesized that this heating had something to do with floral odor volatilization and seed incubation, however, experiments have shown that at least seed development in these two shrubs is not influenced by floral heat in any major way. The same cannot be said for the midge larvae. 

As the flowers mature and give way to developing seeds, the midge larvae are hard at work feeding on tiny bits of the flowers themselves. When researchers looked at midge larvae development on these Illicium species, they found that they were completely dependent upon the floral heat for survival. Any significant drop in temperature caused them to die. Essentially, the plants appear to be producing heat more for the midges than for themselves. It may seem odd that these two plants would invest so much energy to heat their flowers so that midge larvae feeding on their tissues can survive but such face-value opinions rarely stand in ecology.

One must not forget that those larvae grow up to be adult midges that will go on to pollinate the Illicium flowers the following season. Although the plants are taking a bit of a hit by allowing the larvae to develop within their tissues, they are nonetheless ensuring that enough pollinators will be around to repeat the process again. If that wasn't cool enough, the relationship between each of these plants and their pollinators are rather specific and the authors of at least one paper believe that the midges that pollinate each species are new to science. 

Now, if I haven't managed to convince you that this angiosperm sister lineage is anything but primitive, then let's take a look at another genus within the family Schisandraceae that have taken this midge pollination syndrome to the next level. This story also takes place in Asia but instead involves a genus of woody vines known as Kadsura

Like the Illicium we mentioned earlier, a handful of Kadsura species rely on midges for pollination. The way in which they go about maintaining this relationship is a bit more involved. The midges that are attracted by Kadsura flowers are known as resin midges and their larvae live off of plant resins. The flowers of Kadsura are another story entirely. They are as odd as they are beautiful. 

Flowers, pollinators ,and their larvae (white arrows) in  Kadsura heteroclita .

Flowers, pollinators ,and their larvae (white arrows) in Kadsura heteroclita.

In male flowers, stamens are arranged in dense, cone-like structures called androecia whereas the female flowers contain a compact shield-like structure with the uppermost part of the stigma barely emerging. This is called a gynoecium. Even weirder, the male flowers of one particularly strange species, Kadsura coccinea, produce large, swollen inner tepals. 

Once Kadsura flowers begin to open, visiting midges are not far behind. Male flowers seem to attract more midges than female flowers and it is thought that this has to do with varying amounts of special attractant chemicals produced by the flowers themselves. Regardless, midges set to work exploring the blooms with males looking for mates and females looking for a place to lay their eggs. 

When a suitable spot has been found, females will deposit their eggs into the floral tissues with their ovipositor. The wounded plant tissues immediately begin producing resin, not unlike a wounded pine tree. In the case of K. coccinea, it would appear that the oddly swollen tepals are specifically targeted by female midges for egg laying. They too produce resin upon having eggs laid within. 

The oddball flowers of Kadsura coccinea showing swollen tepals.

The function of plant resins in many cases are to fight off pathogens. From beetles to fungi, resin helps plug up and seal off wounds. This does not seem to be the case in the Kadsura-midge relationship though. The so-called "brood chambers" within the floral tissues go on producing resin for upwards of 6 days after the midge eggs were laid. Eventually the floral parts whither and drop off but the midge larvae seem to be quite happy in their resin-filled homes. 

As it turns out, the resin midge larvae feed on the viscous resin as their sole food source. Instead of trying to ward off these pesky little insects, the plants seem to be encouraging them to raise their offspring within! Just as we saw in the Asian Illicium, these Kadsura vines seem to be providing brood sites for their pollinators. Also, just as the Illicium-midge relationship thought to be species specific, each species of Kadsura appears to have its own specific species of resin midge pollinator! K. coccinea even goes as far as to produce tepals specifically geared towards raising midge larvae, thus keeping them away from their more valuable reproductive organs. In return for the nursery service, Kadsura have their pollinators all to themselves.

Pollination mutualisms in which plants trade raising larvae for pollen transfer are extremely derived and some of the most specialize plant/animal interactions on the planet. To find such relationships in these basal or sister lineages is living proof that these plants are anything but primitive. In the energy-reproductive investment trade-off, it appears that ensuring ample pollinator opportunities far outweighs the cost of providing them with nursery chambers. It is remarkable to think just how intertwined the relationships between these plants and there pollinators truly are. Take that, plant taxonomic prejudices! 

Photo Credits: [1] [2]

Further Reading: [1] [2] 

 

The Largest Single Flower in the World

To find some of the largest flowers in the world, one must find themselves hiking through the the humid jungles of southeast Asia. From there you must be lucky enough to stumble across the flowers of a genus known scientifically as Rafflesia. It contains roughly 28 species spattered about various tropical islands. If you are very lucky, you might even find Rafflesia arnoldii. Producing flowers that are over 3 feet (1 m) in diameter and weighing as much as 24 pounds (11 kg), it produces the largest individual flower on the planet. 

Even more bizarre, these plants are entirely parasitic. They belong to a specialized group called holoparasites. These plants produce no stems, no leaves, nor any true roots. Their entire existence depends on a group of vines related to North America's grapes. Except for flowering, individual Rafflesia exist entirely as a network of mycelium-like cells inside the tissues of their vine hosts.


For a long time, the taxonomic status of this plant was highly debated but recent DNA evidence puts it in the order Malpighiales. From there, things get a little funny. One recent analysis suggested that Rafflesia belonged in the family Euphorbiaceae, however, it most likely warrants its own family - Rafflesiaceae.

So, why produce such large flowers? Well, existing solely within a vine makes it hard to establish a large population in any given area. This makes for a difficult situation in the pollinator department. Somehow plants must increase the odds that any given pollinator will visit multiple unrelated individuals of that particular species. By growing very large and and producing a lot of "stink" (this plant is also referred to as the corpse plant), Rafflesia make sure that pollinators will come from far and wide to investigate, thus increasing their chances of cross pollinating. How this plant goes about seed dispersal, however, remains a mystery.

Most interesting of all, it has been discovered that there is some amount of horizontal gene transfer going on between Rafflesia and its host. Basically, Rafflesia obtains strands of DNA from the vine and uses them in its own genetic code. It is believed this incurs some fitness benefit to Rafflesia but more research is needed to figure out why this may be happening. 

Sadly, many species within this family may be lost before we ever get a chance to get to know them. Forests throughout this region are disappearing rapidly to make room for expanding populations and agriculture. What makes matters worse for Rafflesia is that their lifestyle makes them very hard to study. It is especially difficult to obtain accurate population estimates. As more and more forests are cleared, we could be losing countless populations of these wonderful and intriguing plants. As with large mammals, it would seem that the world's largest flower is falling victim to the unending tide of human development. 

Photo Credit: Tamara van Molken


Further Reading:

http://bit.ly/2c2ALHl

http://bit.ly/2cPMP51

http://bit.ly/2cwP7ny

Rhizanthes lowii

Imagine hiking through the forests of Borneo and coming across this strange object. It's hairy, it's fleshy, and it smells awful. With no vegetative bits lying around, you may jump to the conclusion that this was some sort of fungus. You would be wrong. What you are looking at is the flower of a strange parasitic plant known as Rhizanthes lowii.

R. lowii is a holoparasite. It produces no photosynthetic tissues whatsoever. In fact, aside from its bizarre flowers, its doesn't produce anything that would readily characterize it as a plant. In lieu of stems, leaves, and roots, this species lives as a network of mycelium-like cells inside the roots of their vine hosts. Only when it comes time to flower will you ever encounter this species (or any of its relatives for that matter).

The flowers are interesting structures. Their sole purpose, of course, is to attract their pollinators, which in this case are carrion flies. As one would imagine, the flowers add to their already meaty appearance a smell that has been likened to that of a rotting corpse. Even more peculiar, however, is the fact that these flowers produce their own heat. Using a unique metabolism, the flower temperature can rise as much as 7 degrees above ambient. Even more strange is the fact that the flowers seem to be able to regulate this temperature. Instead of a dramatic spike followed by a gradual decrease in temperature, flowers are able to maintain this temperature gradient throughout the flowering period.

There could be many reasons for doing this. It could enhance the rate of floral development. This is a likely possibility as temperature increases have been recorded during bud development. It could also be used as a way of enticing pollinators, which can use the flower to warm up. This seems unlikely given its tropical habitat. Another possibility is that it helps disperse its odor by volatilizing the smelly compounds. In a similar vein, it may improve the carrion mimicry. Certainly this may play a role, however, flies don't seem to have an issue finding carrion that has cooled to ambient temperature. Finally, it has also been suggested that the heat may improve fertilization rates. This also seems quite likely as thermoregulation has been shown to continue after the flowers have withered away.

Regardless of its true purpose, the combination of lifestyle, appearance, and heat producing properties of this species makes for a bizarrely spectacular floral encounter. To see this plant in the wild would be a truly special event.

Photo Credit: Ch'ien C. Lee - www.wildborneo.com.my/photo.php?f=cld1500900.jpg

Further Reading:

http://www.jstor.org/stable/4222678?seq=1#page_scan_tab_contents

http://www.people.fas.harvard.edu/~ccdavis/pdfs/Nikolov_et_al_AJB_2014.pdf