How Fungus Gnats Maintain Jack-in-the-pulpits

[SOURCE]

There are a variety of ways that the boundaries between species are maintained in nature. Among plants, some of the best studied examples include geographic distances, differences in flowering phenology, and pollinator specificity. The ability of pollinators to maintain species boundaries is of particular interest to scientists as it provides excellent examples of how multiple species can coexist in a given area without hybridizing. I recent study based out of Japan aimed to investigate pollinator specificity among fungus gnats and five species of Jack-in-the-pulpit (Arisaema spp.) and found that pollinator isolation is indeed a very strong force in maintaining species identity among these aroids, especially in the wake of forest disturbance.

Fungus gnats are the bane of many a houseplant grower. However, in nature, they play many important ecological roles. Pollination is one of the most underappreciated of these roles. Though woefully understudied compared to other pollination systems, scientific appreciation and understanding of fungus gnat pollination is growing. Studying such pollination systems is not an easy task. Fungus gnats are small and their behavior can be very difficult to observe in the wild. Luckily, Jack-in-the-pulpits often hold floral visitors captive for a period of time, allowing more opportunities for data collection.

By studying the number and identity of floral visitors among 5 species of Jack-in-the-pulpit native to Japan, researchers were able to paint a very interesting picture of pollinator specificity. It turns out, there is very little overlap among which fungus gnats visit which Jack-in-the-pulpit species. Though researchers did not analyze what exactly attracts a particular species of fungus gnat to a particular species of Jack-in-the-pulpit, evidence from other systems suggests it has something to do with scent.

Like many of their aroid cousins, Jack-in-the-pulpits produce complex scent cues that can mimicking everything from a potential food source to a nice place to mate and lay eggs. Fooled by these scents, pollinators investigate the blooms, picking up and (hopefully) depositing pollen in the process. One of the great benefits of pollinator specificity is that it greatly increases the chances that pollen will end up on a member of the same species, thus reducing the chances of wasted pollen or hybridization.

Still, this is not to say that fungus gnats are solely responsible for maintaining boundaries among these 5 Jack-in-the-pulpit species. Indeed, geography and flowering time also play a role. Under ideal conditions, each of the 5 Jack-in-the-pulpit species they studied tend to grow in different habitats. Some prefer lowland forests whereas others prefer growing at higher elevations. Similarly, each species tends to flower at different times, which means fungus gnats have few other options but to visit those blooms. However, such barriers quickly break down when these habitats are disturbed.

Forest degradation and logging can suddenly force many plant species with different habitat preferences into close proximity with one another. Moreover, some stressed plants will begin to flower at different times, increasing the overlap between blooming periods and potentially allowing more hybridization to occur if their pollinators begin visiting members of other species. This is where the strength of fungus gnat fidelity comes into play. By examining different Jack-in-the-pulpit species flowering in close proximity to one another, the team was able to show that fungus gnats that prefer or even specialize on one species of Jack-in-the-pulpit are not very likely to visit the inflorescence of a different species. Thanks to these preferences, it appears that, thanks to their fungus gnat partners, these Jack-in-the-pulpit species can continue to maintain species boundaries even in the face of disturbance.

All of this is not to say that disturbance can’t still affect species boundaries among these plants. The researchers were quick to note that forest disturbances affect more than just the plants. When a forest is logged or experiences too much pressure from over-abundant herbivores such as deer, the forest floor dries out a lot quicker. Because fungus gnats require high humidity and soil moisture to survive and reproduce, a drying forest can severely impact fungus gnat diversity. If the number of fungus gnat species declines, there is a strong change that these specific plant-pollinator interactions can begin to break down. It is hard to say what affect this could have on these Jack-in-the-pulpit species but a lack of pollinators is rarely a good thing. Certainly more research is needed.

Photo Credit: [1]

Further Reading: [1]

My New Book Has Arrived!

The time has finally come! In Defense of Plants: An Exploration into the Wonder of Plants is now in stores. I thank everyone who pre-ordered a copy of the book. They should be on their way! I still can’t believe this is a reality. I always knew I wanted to write a book and I am eternally grateful to Mango Publishing for giving me this opportunity.

In Defense of Plants is a celebration of plants for the sake of plants. There is no denying that plants are extremely useful to humanity in many ways, but that isn’t why this exist. Plants are living, breathing, self-replicating organisms that are fighting for survival just like the rest of life on Earth. And, thanks to their sessile habit, they are doing so in remarkable and sometimes alien ways.

One of the best illustrations of this can be found in Chapter 3 of my new book: “The Wild World of Plant Sex.” Whereas most of us will have a passing familiarity with the concept of pollination, we have only really scratched the surface of the myriad ways plants have figured out how to have sex. Some plants go the familiar rout, offering pollen and nectar to floral visitors in hopes that they will exchange their gametes with another flower of the same species.

Others have evolved trickier means to get the job done. Some fool their pollinators into thinking they are about to get a free meal using parts of their anatomy such as fake anthers or by offering nectar spurs that don’t actually produce nectar. Some plants even pretend to smell like dying bees to lure in scavenging flies. Still others bypass food stimuli altogether and instead smell like receptive female insects in hopes that sex-crazed males won’t know the difference.

Pollination isn’t just for flowering plants either. In In Defense of Plants I also discuss some of the novel ways that mosses have converged on a pollination-like strategy by co-opting tiny invertebrates that thrive in the humid microclimates produced by the dense, leafy stems of moss colonies.

This is just a taste of what is printed on the pages of my new book. I really hope you will consider picking up a copy. To those that already have, I hope you enjoy the read when it arrives! Thank you again for support In Defense of Plants. You are helping keep these operations up and running, allowing me to continue to bring quality, scientifically accurate botanical content to the world. Thank you from the bottom of my heart.

Click here if you would like to order a copy!

You can also purchase a copy directly from the publisher

Floral Trickery of the Bat Plants

Photo by Geoff McKay licensed under CC BY 2.0

Bat plants (genus Tacca) are bizarre-looking plants. Their nondescript appearance when not in flower enshrouds the extravagant and, dare I say macabre appearance of their blooms. The inflorescence of this genus is something to marvel at. The flowers are borne above sets of large, conspicuous bracts and numerous whisker-like bracteoles. Despite their unique appearance and popularity among plant collectors, the pollination strategies utilized by the roughly 20 species of bat plants have received surprisingly little attention over the years.

Bat plants are most at home in the shaded, humid understories of tropical rainforests around the globe (though there are a couple exceptions to this rule). Amazingly, these plants are members of the yam family (Dioscoreaceae) and are thought to be closely related to the equally bizarre Burmanniaceae, a family comprised entirely of oddball parasites. Taxonomic affinities aside, there is no denying that bat plants produce truly unique inflorescences and many a hypotheses has been put forth to explain the function of their peculiar floral displays.

The white bat plant (Tacca integrifolia). Photo by MaX Fulcher licensed under CC BY-NC-SA 2.0

The black bat plant (Tacca chantrieri). Photo by Hazel licensed under CC BY-SA 2.0

The most common of these is that the flowers are an example of sapromyiophily and thus mimic a rotting corpse in both smell and appearance as a means of attracting carrion flies. However, despite plenty of speculation, such hypotheses have largely gone untested. It wasn’t until fairly recently that anyone put forth an attempt to observe pollination of these plants in their natural habitats.

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A) T. leontopetaloides; (B) T. plantaginea; (C) T. parkeri; (D) T. palmatifida; (E) T. palmata; (F) T. subflabellata; (G) T. integrifoliafrom; (H) T. integrifoliafrom; (I) T. ampliplacenta; (J) T. chantrieri. [SOURCE]

A 2005 study done in South Yunnan province, China found that almost nothing visited the flowers of Tacca chantrieri. Despite the presence of numerous potential pollinators, only a handful of small, stingless bees paid any attention to these obvious floral cues. This led the authors to suggest that most bat plants are self-pollinated. Indeed, genetic analysis of different populations of T. chantrieri helped bolster this conclusion by demonstrating that there is very little evidence of genetic transfer between T. chantrieri populations. Yet, this is far from a smoking gun. Strong genetic structuring among populations could simply mean that pollinators aren’t moving very far. Also, if most bat plants simply opt for fertilizing their own blooms, why has this genus maintained such elaborate floral morphology? Needless to say, more work was needed.

Luckily, a recent study from Malaysia has made great strides in our understanding of the sex lives of these plants. By observing 7 different species of bat plant in the wild, researchers were able to collect plenty of data on bat plant pollination. It turns out that the flowers of these 7 species are quite popular with insects. Bat plant floral visitors in their study included everything from tiny, stingless bees to ants, beetles, and weevils. However, the most common floral visitors for most bat plant species were small, biting midges. This is where things get very interesting.

(A–C) Female Forcipomyia biting midge. Arrows indicating pollen grains. [SOURCE]

As their common name suggests, biting midges are most famous for biting other animals. Though they will drink nectar, female biting midges need lots of protein to successfully produce eggs. They meet their protein needs by drinking the blood of insects and mammals. Of the biting midges that most frequently visited bat plant flowers, the most common hail from two groups known to feed exclusively on mammalian blood. Finding these biting midges in high numbers on bat plant flowers raises the question of what they stand to gain from these strange-looking blooms.

The conclusion the authors came to was that bat plant blooms are using a bit of trickery to lure in female midges. They hypothesize that the color patterns of the bracts and flickering motion of whisker-like bracteoles simulates the movements of mammals that the midges normally feed on. It is also possible that bat plant flowers emit volatile scents that enhance this mimicry, though more work is needed to say for sure. What the researchers do know is that the behavior of female biting midges upon visiting a flower is enough to pick up and deposit plenty of pollen as they search for a blood meal that doesn’t exist. How common this floral ruse is among the remaining species is yet to be determined but the similarities in inflorescence structure among members of this genus suggest similar tricks are being played on pollinators wherever bat plants grow.

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

Further Reading: [1] [2]

What an orchid that smells like rotting meat can tell us about carrion flies

Satyrium pumilum Photo by Bernd Haynold licensed by CC BY-SA 3.0

Orchids are really good at tricking pollinators. Take, for instance, this strange looking orchid from South Africa. Satyrium pumilum is probably obscure to most of us but it is doing fascinating things to ensure its own reproductive success. This orchid both smells and kind of looks like rotting meat, which is how it attracts its pollinators.

It is a bit strange to think of orchids living in arid climates like those found in South Africa but this family is defined by exceptions. That is not to say that Satyrium pumilum is a desert plant. To find this orchid, you must look in special microclimates where water sticks around long enough to support its growth. Populations of S. pumilum are most often found clustered near small streams or hidden under bushes throughout the western half of the greater Cape Floristic Region.

Satyrium pumilum blooms from the beginning of September until late October. As is typical in the orchid family, S. pumilum produces rather intricate flowers. Whereas the sepals are decked out in various shades of green, the interior of the flower is blood red in color. Also, unlike many of its cousins, S. pumilum doesn’t throw its flowers up on a tall stalk for all the world to see. Instead, its flowers open up at ground level and give off an unpleasant smell of rotting meat.

This is where pollinators enter into the picture. It has been found that carrion flies are the preferred pollinator for S. pumilum. By producing flowers at ground level that both look and smell like rotting meat, the plants are primed to attract these flies. The plants are tapping into the flies’ reproductive habits, a biological imperative so strong that they simply do not evolve a means of discriminating a rotting corpse from a flower that smells like one. This is the trick. Flies land on the flower thinking they have found a meal and a place to lay their eggs. They go through the motions as expected and pick up or deposit pollen in the process. Unfortunately for the flies, their offspring are doomed. There is not food to be found in these flowers.

What is most remarkable about the reproductive ecology of S. pumilum is that not just any type of fly will do. It appears that only a specific subset of flies actually visit the flowers and act as effective pollinators. Amazingly, this provides insights into some long-running hypotheses regarding carrion fly ecology.

(A) The habitat of S. pumilum (B) Satyrium pumilum in situ (scale bar = 1 cm). (C–E) Pollination sequence of a S. pumilum flower by a sarcophagid fly in an arena (scale bar for all three photos = 0·5 cm); (C) the fly carrying five pollinaria from other S. pumilum flowers enters an unpollinated flower (D) as the fly moves deeper into the flower towards the right-hand spur, it presses an attached pollinium against the stigma, and its thorax against the right-hand viscidium; (E) as it leaves the flower, the fly has deposited two massulae on the stigma (1), and removed a pollinarium (2) – it now carries six pollinaria. [SOURCE]

Apparently there has been a lot of debate in the fly community over why we see so many different species of carrion flies. Rotting meat is rotting meat, right? Probably not, actually. Fly ecologists have comes up with a few hypotheses involving niche segregation among carrion flies to explain their diversity on the landscape. Some believe that flies separate themselves out in time, with different species hatching out and breeding at different times of the year. Others have suggested that carrion flies separate themselves by specializing on carrion at different stages of decay. Still others have suggested that some flies specialize on large pieces of carrion whereas others prefer smaller pieces.

By studying the types of flies visiting the flowers of S. pumilum researchers did find evidence of niche segregation based on carrion size. It turns out that S. pumilum is exclusively pollinated by a group of flies known as sarcophagid carrion flies. These flies were regularly observed with orchid pollen sacs stuck to their backs and plants seemed to only set seed after they had been visited by members of this group. So, what is it about these flowers that makes them so specific to this group of flies?

The answer lies both in their size as well as the amount of scent they produce. It is likely that the quantity of scent compounds produced by S. pumilum most closely mimics that of smaller rotting corpses. The types of flies that visited these blooms were mostly females of species that lay relatively few eggs compared to other carrion flies. It could very well be that the smaller brood size of these flies allows them to effectively utilize smaller bits of carrion than other, more fecund species of fly. To date, this is some of the best evidence in support of the idea that flies avoid competition among different species by segregating out their feeding and reproductive niches.

Rotting meat smells are not uncommon in the plant world. Even within the home range of S. pumilum, there are other plants produce flowers that smell like carrion as well. It would be extremely interesting to look at what kinds of flies visit other carrion flowers and in what numbers. Like I mentioned earlier, reproductive is such a major part of any organisms life that it may simply be too costly for carrion flies to evolve a means of discriminating real and fake breeding sites. It is amazing to think of what we gain from trying to understand the reproductive biology of a small, obscure orchid growing tucked away in arid regions of South Africa.

Photo Credits: [1] [2]

Further Reading: [1]

Rhizanthes lowii

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

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.

Rhizanthes 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 function, 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 metabolic pathway, 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, the flowers of R. lowii are able to maintain this temperature gradient throughout the flowering period.

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

There could be many reasons for doing this. Heat 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: [1] [2]