A Tiny Passionflower with a Hardy Disposition

Passionflowers barely need an introduction. Who hasn't marveled at the beautiful splendor of their intricate blossoms. Thought largely tropical in their distribution, there are a couple members of the genus Passiflora that have tackled temperate North America. My favorite of these is small and not nearly as gaudy as its cousins but that is kind of what makes me like it so much. Today, I would like to introduce you to the yellow passionflower (Passiflora lutea).

Did I mention this was a small plant? Whereas it can vine itself over surrounding vegetation very effectively, it is by no means a bulky plant. Even more incredible are its flowers. Anyone familiar with the anatomy of Passiflora flowers will be shocked to see all of that detail miniaturized into a yellow-green bloom about the size of your thumbnail. You must be quick to catch these in flower as they themselves are only open for about a day.

As I mentioned above, Passiflora don't come any hardier than the yellow passionflower (except maybe P. incarnata). With a range that extends as far north as Pennsylvania, this lovely little vine can handle winter temperatures as low as −30 °C (-22 °F)! This has earned it the designation of the northernmost species of Passiflora. Even then, I have heard reports of people growing this hardy little plant farther north in Canada.

Pollination for this species works in much the same way as it does for the genus as a whole. The flowers require an insect large enough to contact the peculiar arrangement of anthers and stigmas. The strange yet beautiful filaments that ring the center of the bloom are collectively referred to as the corona and it is believed that these guide insects to the nectar and thus into perfect position for pollination.

By far the most peculiar aspect of this plant is the relationship it has formed in part of its range with a tiny bee aptly named the passionflower bee (Anthemurgus passiflorae). Native from central Texas to North Carolina and north to Illinois, this tiny black bee is the only member of its genus. What's more, it absolutely requires the yellow passionflower for its reproduction. It feeds its larvae solely on pollen from the yellow passionflower. If that wasn't strange enough, despite its highly specific foraging habits, the diminutive size of the bee has led experts to believe that the passionflower bee contributes very little in the way of pollination for the plant.

Further Reading: [1] [2]

The Pine Lily

The pine lily (Lilium catesbaei) is one of North America’s finest species of lily. It produces the largest flowers of the genus on this continent and to see one in person is a breathtaking experience. The pine lily is endemic to the Southeastern Coastal Plain where it prefers to grow in mesic to wet flatwoods, wet prairies, and savannas. Though it enjoys a relatively wide distribution, today it rarely occurs in any abundance.

The pine lily’s rarity may be a relatively recent status change for this wonderful plant. Historical records indicate that it was once quite abundant in states like Florida. Today it occurs in scattered localities and predicting its presence from year to year has been a bit tricky. Indeed, the pine lily appears to be very picky when it comes to growing and flowering.

One aspect of its biology that might lend to its limited appearance is the fact that it can remain underground in a dormant state for years. Like other members of this genus, the pine lily emerges from a bulb. This underground storage structure is small by lily standards, which means that most pine lilies are operating on marginal stores of energy in any given year.

Some have estimated that individual bulbs can remain dormant for upwards of 5 years before the right conditions for growth flowering present themselves. Of course, such dormancy can be a nightmare for proper conservation of such a unique plant. Aside from the individual flower borne at the tip of a long, slender stem, the rest of the plant is very dainty. In fact, its flowers can be so heavy compared to the rest of the plant that some stems simply topple to the ground before they can set seed. The slender stem, small leaves, and tiny bulb equate to a small operating budget in terms of energy stores. That being said, we are starting to get a clearer picture of what pine lilies need to thrive and it all comes down to fire.

Photo by Eleanor licensed by CC BY-NC 2.0

The key to acquiring enough energy for growth and reproduction appears to be a proper amount of sunlight. Without it, plants languish. This is where fire comes in. The pine lily lives in a region of North America that historically would have burned with some frequency. Wildfires sweep through an area, burning away competing vegetation like saw palmetto (Serenoa repens) and clearing the ground of accumulated debris like sticks and leaves. By burning away the competition, fire creates open areas where delicate plants like the pine lily can eke out an existence. Indeed, research has shown that pine lilies produce more flowers and seed immediately following ground-clearing burn followed by a subsequent decline in flowering and seed set as the surrounding vegetation begins to grow back.

If a pine lily does have enough energy to flower, then one of the most stunning flowers in all of North America is presented with its face towards the sky. Its 6 large petals are brightly colored and taper down into what looks like tiny tubes. Nectar is produced within these tubes and, coupled with the bright coloration, attract numerous insect visitors.

Not all insects are capable of successfully pollinating such a large flower. In fact, it would appear that only a couple of species take up the bulk of the pollination of this incredible plant. As far as we know, the Palamede swallowtail butterfly (Papilio palamedes) and perhaps the spicebush swallowtail (P. troilus) are the only species large enough to properly contact both anthers and stigma while feeding at the flowers. The large wingspan of these butterflies do all of the work in picking up and depositing pollen. All other insects are simply too small to adequately achieve such feats.

Though we still have a lot more to learn about the pine lily, what we do know tells us a story that is repeated for fire-dependent ecosystems throughout the world. Without regular disturbance from fire, biodiversity drops. The pine lily is not alone in this either. Its fate is intertwined with countless other unique plant species that call the coastal plains their home.

Photo Credits: [2]

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

Surprising Genetic Diversity in Old Growth Trees

Photo by S. Rae licensed by CC BY 2.0

Long-lived trees face a lot of challenges throughout their lives. Many trees can live for centuries, which can be a problem because plants cannot get up and move when conditions become unfavorable. This should equate to a slower rates of adaptation and evolution for long lived trees but that isn’t always the case. Many trees are often superbly capable of adapting to local conditions. Recently, a team of researchers from the University of British Columbia have provided some insights into the genetic mechanisms that may underpin such adaptive potential.

Genetic insights came from a species of conifer many will be familiar with - the Sitka spruce (Picea sitchensis). Researchers were interested in these trees because they live for a long time (upwards of 500 years or more) and can grow to heights of over 70 meters (230 ft.). They wanted to understand how genetic mutations work in trees like the Sitka spruce because plants are doing things a bit different than animals in that department.

Plants are modular organisms, meaning they grow by producing multiple copies of discrete units. This equates to a branching structure whose overall shape is in large part determined by environmental influences. It also means that when genetic mutations occur in one branch, they can be carried on throughout the growth of those tissues independent of what is going on throughout the rest of the plant. This means that older trees can often accumulate a surprising amount of genetic diversity throughout the entire body of the plant.

Photo by Brandon Kuschel licensed by Creative Commons Attribution 3.0 Unported

When researchers sampled the DNA of tissues from the trunks and the needles of tall, old growth Sitka spruce, they were shocked by what they had found. From the base of the tree to the needles in the canopy, an old growth Sitka spruce can show as much as 100,000 genetic differences. That is a lot of genetic diversity for a single organism. Though plenty of other trees have been found to exhibit varying levels of genetic differences within individuals, this is one of the highest mutation rates ever found in a single eukaryotic organism. This could also explain why such long-lived organisms can survive in a changing world for their entire lives.

Now, it is important to note that many mutations are likely either neutral or potentially harmful. Also, the rates of mutation may differ depending on where you look on this tree. For instance, needles at the top of a Sitka spruce are going to be exposed to far more gene-altering UV radiation than bark tissues near the base. Still, over the lifetime of a single tree, rare beneficial mutations can and do accumulate. Imagine a scenario in which one branch mutation results in needles that are more resistant to say an insect pest. Those needles could hypothetically receive less damage than needles elsewhere on the tree. This odd form of selection is occurring within the lifetime of that tree and may even have implications for the future offspring of that tree thanks again to the quirks of how tree reproductive cells develop.

Many trees also do not have segregated germlines. What this means is that unlike animals whose reproductive cells develop from separate cell lineages than the rest of their body cells, the reproductive cells of trees develop from somatic cells, which are the same cells that form stems, leaves, and branches. This means that if a mutation occurs on the germline of a branch that eventually goes on to produce cones, these mutations can be passed on in the seeds of those cones. This obviously needs a lot of evidence to substantiate but now that a mechanism is in place, we know where and what to look for.

Photo Credits: [1] [2]

Further Reading: [1] [2]

Let's Talk About Recruitment

Photo by --Tico-- licensed by CC BY-NC-ND 2.0

For any species to be considered successful, it must replace itself generation after generation. We call this process recruitment and it is very important. After all, reproduction is arguably the most fundamental aspect of life in a Darwinian sense. For plants, this can be done either vegetatively or sexually via seeds and spores. Though vegetative reproduction is a fundamental process for many plants around the globe, seed or spore germination is arguably the most important. To truly understand what a plant needs, we have to understand its germination requirements.

Recruitment is a considerable limiting factor for plant populations. In fact, it is the first major bottleneck plants must pass through. It is estimated that a majority of plant mortality occurs during the germination and seedling stages. However, not all plants are equal in this way. Some plants are considered seed or propagule limited whereas others are habitat limited.

If a plant is seed limited, it means that its ability to expand its population or colonize new habitats its limited by the ability of seeds (or spores) to make it to a new location. Once there, nature takes its course and germination occurs with little impediment. If a plant is habitat limited, however, things get a bit more tricky. For habitat limited plants, simply getting seeds to a new location is not enough. Some other aspect of the environment (soil moisture, texture, temperature, disturbance, etc.) limit successful germination. Only when the right conditions are present can habitat limited plants germinate and begin to grow.

Habitat limitation is probably the most common limit to plant establishment. Simply put, not all plants will be successful everywhere. Even the successful growth and persistence of adult plants can be poor predictors of seedling success. Many plants can live for decades or even centuries and the conditions that were present when they germinated may have long since changed. Even the presence of the adults themselves can make a site unsuitable for germination. Think of all of those fire adapted species out there that require the entire community to burn before their seeds will ever germinate.

In reality, it is likely that most plants are habitat limited to some degree. These are not binary categories after all, rather they are aligned along a spectrum of possibilities. The fact that most plants don’t completely take over an area once seeds or spores arrive is proof of the myriad limits to plant establishment. As such, recruitment limitation is extremely important to study. It can make a huge difference in the context of conservation and restoration. Even the successful establishment of adult plants is no guarantee that seedlings stand a chance. Without successful recruitment, all you have left is a nice garden that is doomed to run its course. By understanding the limits to plant recruitment, we can do much more than just improve on our ability to protect and bolster plant populations, we can also gain insights into why so many plants remain rare on the landscape and so few ever rise to dominance.

Photo Credits: [1]

Further Reading: [1] [2]

The Round Leaved Orchid

In the northern temperate regions of North America, late June marks the beginning of what I like to call orchid season. If you're lucky you may stumble across one of these rare beauties in full bloom. Their diversity in shape and size are mainly a result of the intricate evolutionary relationships they have formed with their pollinators. I spend much of my time botanizing trying to locate and photograph these botanical curiosities and any time I get to meet a new species is a very special time indeed. 

Take the round leaved orchid (Platanthera orbiculata) for example. For years I have only known this species as two round leaves that are slightly reminiscent of the phaleanopsis orchids you see for sale in nurseries and grocery stores. The leaves can be quite large too. With their glossy appearance, they are the easiest way to locate this plant.

When conditions are right and the plants have enough stored energy they will begin to flower. Rising from the middle of the pair of leaves is a decent sized inflorescence loaded with greenish white flowers. The flowers are interesting structures. Not particularly colorful, they have a long white lip and considerable green nectar spurs. There are said to be two varieties of this species, each being characterized by the length of the nectar spur. Unlike many orchids that offer no reward to pollinators, P. orbiculata produces nectar. The flowers are pollinated by noctuid moths, which is probably why they are white in color. Whereas most lepidopteran pollinated orchid attach their pollinia to the proboscis of the butterfly or moth, P. orbiculata attaches its pollinia to the eyes of visiting moths. 

If this isn't strange enough, the pollinia themselves have some of their own intriguing adaptations. Visiting moths take a certain amount of time to successfully access the nectar from the nectar spur. If the plant is to avoid wasting precious pollen on itself, then it must find a way to delay this process. The pollinia are the solution to this. When first attached to the eyes, the pollinia stick straight up. This keeps them away from the female parts of the plant as the moth feeds. Only after enough time has elapsed will the stalks of the pollinia begin to bend forward. At this point the moth will hopefully have moved on to the flowers of an unrelated individual. Pointing straight forward, they are now perfectly positioned to transfer pollen. 

Like all orchids, P. orbiculata relies on specialized mycorrhizal fungi for germination and survival. At the beginning of its life, P. orbiculata relies solely on the fungi for sustenance. Once it has enough energy to produce leaves it will repay the fungi by providing carbohydrates. However, the relationship is not over at this point. Every spring, P. orbiculata produces a new set of leaves as well as a whole new root system. The fungi supply a lot of energy for this process and if the plant is disturbed (ie. dug up by greedy poachers) or browsed upon, it is likely that it will not recover from the stress and it will die. The mycorrhizal fungi it relies on live on rotting wood so finding well rotted logs is a good place to start searching for this species. With declining populations throughout much of its range, it is important to remember to enjoy it where it grows. Leave wild orchids in the wild!

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

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]

Can Cultivation Save the Canary Island Lotuses?

Photo by VoDeTan2 Dericks-Tan licensed under the GNU Free Documentation License

Growing and propagating plants is, in my opinion, one of the most important skills humanity has ever developed. That is one of the reasons why I love gardening so much. Growing a plant allows you to strike up a close relationship with that species, which provides valuable insights into its biology. In today’s human-dominated world, it can also be an important step in preventing the extinction of some plants. Such may be the case for four unique legumes native to the Canary Islands provided it is done properly.

The Canary Islands are home to an impressive collection of plants in the genus Lotus, many of which are endemic. Four of those endemic Lotus species are at serious risk of extinction. Lotus berthelotii, L. eremiticus, L. maculatus, and L. pyranthus are endemic to only a few sites on this archipelago. Based on old records, it would appear that these four were never very common components of the island flora. Despite their rarity in the wild, at least one species, L. berthelotii, has been known to science since it was first described in 1881. The other three were described within the last 40 years after noting differences among plants being grown locally as ornamentals.

Photo by John Rusk licensed under CC BY 2.0

All four species look superficially similar to one another with their thin, silvery leaves and bright red to yellow flowers that do a great impression of a birds beak. The beak analogy seems apt for these flowers as evidence suggests that they are pollinated by birds. In the wild, they exhibit a creeping habit, growing over rocks and down overhangs. It is difficult to assess whether their current distributions truly reflect their ecological needs or if they are populations that are simply hanging on in sites that provide refugia from the myriad threats plaguing their survival.

None of these four Lotus species are doing well in the wild. Habitat destruction, the introduction of large herbivores like goats and cattle, as well as a change in the fire regime have seen alarming declines in their already small populations. Today, L. eremiticus and L. pyranthus are restricted to a handful of sites on the island of La Palma and L. berthelotii and L. maculatus are restricted to the island of Tenerife. In fact, L. berthelotii numbers have declined so dramatically that today it is considered nearly extinct in the wild.

Contrast this with their numbers in captivity. Whereas cultivation of L. eremiticus and L. pyranthus is largely restricted to island residents, L. berthelotii and L. maculatus and their hybrids can be found in nurseries all over the world. Far more plants exist in captivity than in their natural habitat. This fact has not been lost on conservationists working hard to ensure these plants have a future in the wild. However, simply having plants in captivity does not mean that the Canary Island Lotus are by any means safe.

One of the biggest issues facing any organism whose numbers have declined is that of reduced genetic diversity. Before plants from captivity can be used to augment wild populations, we need to know a thing or two about their genetic makeup. Because these Lotus can readily be rooted from cuttings, it is feared that most of the plants available in the nursery trade are simply clones of only a handful of individuals. Also, because hybrids are common and cross-pollination is always a possibility, conservationists fear that the individual genomes of each species may run the risk of being diluted by other species’ DNA.

Photo by VoDeTan2 Dericks-Tan licensed under the GNU Free Documentation License

Luckily for the Canary Island Lotus species, a fair amount of work is being done to not only protect the remaining wild plants, but also augment existing as well as establish new populations. To date, many of the remaining plants are found within the borders of protected areas of the island. Also, new areas are being identified as potential places where small populations or individuals may be hanging on, protected all this time by their inaccessibility. At the same time, each species has been seed banked and entered into cultivation programs in a handful of botanical gardens.

Still, one of the best means of ensuring these species can enjoy a continued existence in the wild is by encouraging their cultivation. Though hybrids have historically been popular with the locals, there are enough true species in cultivation that there is still reason for hope. Their ease of cultivation and propagation means that plants growing in peoples’ gardens can escape at least some of the pressures that they face in the wild. If done correctly, ex situ cultivation could offer a safe haven for these unique species until the Canary Islands can deal with the issues facing the remaining wild populations.

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

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

The Cypress-Knee Sedge

Sedges (Carex spp.) simply do not get the attention they deserve. I am part of this problem because like so many others, I have breezed over them in vegetation surveys as “just another graminoid.” This is truly a shame because not only are sedges absolutely fascinating organisms, they are immensely important ecologically as well. I am working hard to get to know sedges better so that I too can fully appreciate their place in our ecosystems. One of the coolest specialist sedges I just recently learned about is the so-called cypress-knee sedge (Carex decomposita). For all intents and purposes, this sedge is considered something of an epiphyte!

The cypress-knee sedge has a fondness for growing on wood. Most often you will find it rooted to the buttresses and knees of bald cypress (Taxodium distichum) or the swollen trunk of a swamp tupelo (Nyssa aquatica). It can also be found growing out of rotting logs that float on the surface of the water. It is a long lived species, with individuals having records stretching back through decades of wetland plant surveys. When supplied with the conditions it likes, populations can thrive. That is not to say that it does well everywhere. In fact, it has declined quite a bit throughout its range.

Juvenile cypress-knee sedges establishing in moss along the water line of a bald cypress.

One of the key wetland features that the cypress-knee sedge needs to survive and prosper is a stable water level. If water levels change too much, entire populations can be wiped out either by drowning or desiccation. Even before the sedge gets established, its seeds require stable water levels to even get to suitable germination sites. Each achene (fruit) comes complete with a tiny, corky area at its tip that allows the seeds to float. Floating seeds are how this species gets around. With any luck, some seeds will end up at the base of a tree or on a floating log where they can germinate and grow. If water levels fluctuate too much, the seeds simply can’t reach such locations.

Its dependence on high quality wetlands is one of the major reasons why the cypress-knee sedge has declined so much in recent decades. Aside from outright destruction of wetlands, changes in wetland hydrology can have dire consequences for its survival. One of the major issues for the cypress-knee sedge is boat traffic. Boat wakes create a lot of disturbance in the water that can literally scour away entire populations from the base of trees and logs. Another major threat are changes to upstream habitats. Any alteration to the watersheds of wetland habitats can spell disaster for the cypress-knee sedge. Alterations to creeks, streams, and rivers, as well as changes in ground water infiltration rates can severely alter the water levels in the swamps that this sedge depends on for survival.

Closeups of the infructescence showing details of the perigynia (fruit).

Less obvious threats also include changes in plant cover. If the wetlands in which it grows become too dense, the cypress-knee sedge quickly gets out-competed. To thrive, the cypress-knee sedge needs slightly more sunlight than a densely forested wetland can provide. In fact, some have even noted that cypress-knee sedge populations can explode after selective logging of such wetlands. Such explosions have been attributed to not only extra sunlight but also the addition of woody debris, which provides much needed germination sites. That being said, such explosions can only be maintained if woody debris is left in place and further wetland disturbances do not continue.

The plight of the cypress-knee sedge stands as a reminder of just how poorly we treat wetlands around the globe. Aside from providing valuable ecosystem services for the human environment (flood control, water filtration, etc.), wetlands are home to countless unique species. Only by treating wetlands betters and attempting to restore some of what has been lost will we ever do better by wetland species like the cypress-knee sedge. Hopefully by showcasing species like this, people will begin to feel a little more compassion towards the ecosystems on which they depend. Please consider supporting a wetland conservation and restoration initiative in your region!

Photo Credits : LDWF Natural Heritage Program [1] & Paul Marcum (Midwest Graminoides) [2] [3] [4]

Further Reading: [1] [2]

Pitcher Plants with a Taste for Salamanders?

Photo by Chris Moody licensed under CC BY-NC 2.0

The thought of a carnivorous plant trapping and digesting a vertebrate may seem more like fiction than reality. Though rumors have circulated over the years that some pitcher plants have a taste for animals larger than an insect, this has been hard to prove as evidence has been notoriously lacking. That is not to say it does not happen from time to time. Small mammals have indeed been found in the pitchers of some of the larger tropical pitcher plants in the genus Nepenthes. Still, these seem more incidental than regular. However, recent observations from Canada suggest that vertebrates may actually make up a bigger part of the menu of some pitcher plants than we previously thought at least under certain circumstances.

The observations were made in Algonquin Provincial Park, Ontario. The carnivore responsible is North America’s most abundant pitcher plant - the purple pitcher plant (Sarracenia purpurea). In late summer of 2017, researchers discovered that some pitchers contained recently metamorphosed salamanders. Some of the salamanders were alive but a few others were dead and undergoing digestion. This was very exciting because despite plenty of study, there has been almost no substantiated evidence of vertebrate prey capture in the purple pitcher plant.

Subsequent surveys were done to figure out if the purple pitcher plants were indeed capturing salamanders on a regular basis or if the salamanders were one-off events. It turns out that, at least for the pitcher plants growing in this bog, salamanders may make up a considerable proportion of their prey! Researchers found that recently metamorphosed spotted salamanders were present in nearly 20% of the pitcher plants they surveyed!

Not all of the salamanders they found were dead. Some were found in a relatively lively state, retreating down into the bottom of the pitcher whenever they were disturbed. Some of the larger dead specimens showed signs of putrefaction, which is probably because they were simply too large to be properly digested. Still, many of the dead salamanders showed signs of digestion, which suggests that the plants are in fact benefiting from salamander capture. In fact, it has been estimated that a single salamander could contribute as much nitrogen to the pitcher plant as the entire contents of three pitchers combined.

Taken together, the team found enough evidence to suggest that salamanders not only make up a portion of the pitcher plants’ diet in this bog, but also that pitcher plants are a significant source of mortality for young salamanders in this system. How the salamanders are caught is up for some debate. It could be that the salamanders are looking for a safe, wet place to hide, however, the complexity of the bog habitat means that there is no shortage of safe places for a young salamander to hide that won’t end in death.

It could also be that salamanders are attracted to all of the invertebrates that these plants capture or that salamanders are accidental victims, having fallen into the trap randomly as they explore their habitat. However, some pitchers not only contained more than one salamander, the plants position and stature within the bog means that most salamanders would have had to actively climb up and into the pitcher in order to end up inside. It very well may not be random chance after all. Certainly this will require more tests to say for sure.

What we can say for now is that within the confines of this Algonquin bog, salamanders are being trapped and digested by the purple pitcher plant. How much of this is unique to the circumstances of this particular bog and how much of this is something going on in other areas within the range of the purple pitcher plant is a subject for future research. It is possible that vertebrate prey may be more common among carnivorous plants than we ever thought!

Photo Credits: [1] [2]

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

The Dual Benefits of Smelling Like Frightened Aphids

Photo by KENPEI licensed under the GNU Free Documentation License

If you garden, you have probably dealt with aphids. These tiny sap-suckers not only drain the plant of valuable sap, they can also serve as vectors for disease. Plants must contend with the ever-present threat of aphid infestation throughout the growing season and have evolved some amazing defenses against these insects. Recently an incredible form of defense against aphids has been described in pyrethrum (Tanacetum cinerariifolium) and it involves smelling like a frightened aphid colony.

Aphids produce their own alarm pheromones when attacked. Because aphids form large, clonal colonies, these pheromones can help warn their kin of impending doom. Other aphids will also eavesdrop on these alarm signals and will avoid settling in on plants where aphids are being attacked. Aphids aren’t the only ones honing in on these scents either. Aphid predators and parasitoids will also use these compounds to locate aphid colonies. As such, these pheromones are helpful to the host plant because it can mean a reduction in aphid numbers.

An alate (winged) green peach aphid (Myzus persicae).

The selection pressured imposed by aphids on plants is so strong that it appears that at least one species of pyrethrum has actually evolved a means of producing these pheromones themselves. Pyrethrum is a member of the aster family (Asteraceae) native to southern portions of Eurasia. Like all flowering plants, its flowers are the most precious organs. They are the key to getting their genes into the next generation and therefore protecting them from herbivore damage is of utmost importance.

It has been discovered that pyrethrums produce an aphid alarm pheromone called ( E )-β-farnesene or EβF for short. The pheromone is not produced in every tissue of the plant but rather it is concentrated near the inflorescence. What’s more, pheromone production is not constant throughout the duration of flowering. Researchers found that production reaches its peak just before the inflorescence opens to reveal the flowers within.

Photo by そらみみ licensed under CC BY-SA 4.0

The production of EβF in pyrethrum appears to serve a dual function. For starters, it actually results in reduced aphid infestation during the early stages of flowering. When the initial aphid attack begins, these insects consume some of the EβF as they feed and release it as they excrete honeydew. Other aphids detect EβF within the honeydew and will actually avoid the plant, likely due to the perception that the aphids feeding there are already under attack.

That does not mean that predators are not to be found. In fact, the other benefit of producing EβF in the inflorescence is that it appears to lure in one of the most voracious aphid predators on the planet - ladybird beetles. The ladybird beetles are able to detect EβF in the air and will come from far and wide to investigate in hopes of finding a tasty aphid meal. The ladybird beetles were most frequently found on plants during the early stages of floral development, which suggests that EβF production in the floral tissues is the main attractant.

A 7-spot ladybird beetle (Coccinella septempunctata). Photo by S. Rae licensed under CC BY 2.0

Interestingly, it has been found that constant production of EβF is less effective at deterring aphids than pulses of EβF. It is thought that just as humans can get used to certain background levels of scent, so too can aphids. If aphids are exposed to high levels of EβF for long periods of time, they simply recognize it as the safe background level and will continue to feed. This may explain why pyrethrum plants only produce EβF for a short period of time during the most crucial stages of floral development. Research like this not only improves our understanding of the myriad ways in which plants defend themselves, it also offers us new avenues for researching more natural ways of defending the plants we rely on from unwanted pests.

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

Further Reading: [1]

The Fungus-Mimicking Mouse Plant

The mouse plant (Arisarum proboscideum) is, to me, one of the most charming aroids in existence. Its small stature and unique inflorescence are a joy to observe. It is no wonder that this species has attained a level of popularity among those of us who enjoy growing oddball plants. Its unique appearance may be reason enough to appreciate this little aroid but its pollination strategy is sure to seal the deal.

The mouse plant is native to shaded woodlands in parts of Italy and Spain. It is a spring bloomer, hitting peak flowering around April. It has earned the name “mouse plant” thanks to the long, tail-like appendage that forms at the end of the spathe. That “tail” is the only part of the inflorescence that sticks up above the arrow-shaped leaves. The rest of the structure is presented down near ground level. From its stature and position, to its color, texture, and even smell, everything about the inflorescence is geared around fungal mimicry.

{{PD-US}}

The mouse plant is pollinated by fungus gnats. However, it doesn’t offer them any rewards. Instead, it has evolved a deceptive pollination syndrome that takes advantage of a need that all living things strive to attain - reproduction. To draw fungus gnats in, the mouse plant inflorescence produces compounds that are said to smell like fungi. Lured by the scent, the insects utilize the tail-like projection of the spathe as a sort of highway that leads them to the source.

Once the fungus gnats locate the inflorescence, they are presented with something incredibly mushroom-like in color and appearance. The only opening in the protective spathe surrounding the spadix and flowers is a tiny, dark hole that opens downward towards the ground. This is akin to what a fungus-loving insect would come to expect from a tiny mushroom cap. Upon entering, the fungus gnats are greeted with the tip of the spadix, which has come to resemble the texture and microclimate of the underside of a mushroom.

Anatomy of a mouse plant inflorescence [SOURCE]

This is exactly what the fungus gnats are looking for. After a round of courtship and mating, the fungus gnats set to work laying eggs on the tip of the spadix. Apparently the tactile cues are so similar to that of a mushroom that the fungus gnats simply don’t realize that they are falling victim to a ruse. Upon hatching, the fungus gnat larvae will not be greeted with a mushroomy meal. Instead, they will starve and die within the wilting inflorescence. The job of the adult fungus gnats is not over at this point. To achieve pollination, the plant must trick them into contacting the flowers themselves.

Both male and female flowers are located down at the base of the structure. As you can see in the pictures, the inflorescence is two-toned - dark brown on top and translucent white on the bottom. The flowers just so happen to sit nicely within the part of the spathe that is white in coloration. In making a bid to escape post-mating, the fungus gnats crawl/fly towards the light. However, because the opening in the spathe points downward, the lighted portion of the structure is down at the bottom with the flowers.

The leaves are the best way to locate these plants. Photo by Meneerke bloem licensed under CC BY-SA 4.0

Confused by this, the fungus gnats dive deeper into the inflorescence and that is when they come into contact with the flowers. Male and female flowers of the mouse plants mature at the exact same time. That way, if visiting fungus gnats happen to be carrying pollen from a previous encounter, they will deposit it on the female flowers and pick up pollen from the male flowers all at once. It has been noted that very few fungus gnats have ever been observed within the flower at any given time so it stands to reason that with a little extra effort, they are able to escape and with any luck (for the plant at least) will repeat the process again with neighboring individuals.

The mouse plant does not appear to be self-fertile so only pollen from unrelated individuals will successfully pollinate the female flowers. This can be a bit of an issue thanks to the fact that plants also reproduce vegetatively. Large mouse plant populations are often made up of clones of a single individual. This may be why rates of sexual reproduction in the wild are often as low as 10 - 20%. Still, it must work some of the time otherwise how would such a sophisticated form of pollination syndrome evolve in the first place.

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

Further Reading: [1] [2]

A Palm With a Unique Pollination Syndrome

Photo by Dr. Scott Zona licensed under CC BY-NC 2.0

I would like to introduce you to the coligallo palm (Calyptrogyne ghiesbreghtiana). The coligallo palm is a modest palm, living out its life in the understory of wet, tropical forests from Mexico to Panama. To the casual observer, this species doesn’t present much of anything that would seem out of the ordinary. That is, until it flowers. Its spike-like inflorescence is covered in fleshy white flowers that smell of garlic and as far as we know, the coligallo palm is the only palm that requires bats for pollination.

Flowering for this palm occurs year round. At first glance, the inflorescence doesn’t appear out of the ordinary but that is where close observation comes in handy. The more scrutiny they are given, the more strange they appear. As mentioned, the flowers are bright white in color and they smell strongly of garlic. Also, they are protandrous, meaning the male flowers are produced before the female flowers.

Photo by Dr. Scott Zona licensed under CC BY-NC 2.0

After the male flowers have shed their pollen, there is a period of a few days in which no flowers are produced. Then, after 3 to 4 nights of no flowers, female flowers emerge, ready to receive pollen. Each flower only opens at night and does not last for more than a single evening. Protandry is an excellent strategy to avoid self-pollination. By separating male and female flowers in time, each plant can assure that its own pollen will not be deposited back onto its own stigmas. The fact that the coligallo palm flowers year-round means that there is always a receptive plant somewhere in the forest.

The oddities do not end there. Both male and female flowers are covered in a fleshy tube that must be removed for pollination to occur successfully. Removal of the tube is what actually exposes the reproductive organs and allows pollen transfer to occur. Often times, the flowers of the coligallo palm are dined upon by katydids and other insect herbivores. This does not result in pollination as they completely destroy the flower as they eat. Considering the success of this plant across its range, it stands to reason that something else must provide ample pollination services.

Two species of bat visiting coligallo palm inflorescences: A) A perching Artibeus bat feeding on male flowers and B) a hovering Glossophaga bat feeding on female flowers.

As it turns out, bats are that pollinator. The job of pollination is not accomplished by a single species of bat either. A few species have been observed visiting the inflorescences. Apparently the bright color and strong odor of the flowers acts as a calling card for flower-feeding bats throughout these forests. Interestingly, the feeding mechanism of each species of bat differs as well. Some bats hover at the inflorescence like hummingbirds, chewing off the fleshy tube from individual flowers as they go. Other bats prefer to perch on the inflorescence itself, crawling all over it as they eat. These different feeding behaviors actually result in different levels of pollination. Though both forms do result in seed set, perching bats appear to be the most effective pollinators of the coligallo palm.

The reason for this is due to the fact that perching bats not only spend more time on the inflorescence, their bodies come into contact with far more flowers as they feed. Hovering bats, on the other hand, only manage to contact a few flowers with their snout at a time. So, despite the variety of bats recorded visiting coligallo palms, the perching bats appear to provide the best pollination services.

A coligallo palm infructescence showing signs of ample pollination. Photo by Dick Culbert licensed under CC BY 2.0

The role of perching bats in the ecology of this palm species does not end with pollination either. It turns out, they also play a crucial role in the dispersal of certain mites that live on the palm flowers. Flower mites live on plants and consume tiny amounts of pollen and nectar. As you can imagine, their small size makes it incredibly difficult for them to find new feeding grounds. This is where perching bats come into play.

It was discovered that besides pollen, perching bats also carried considerable loads of flower mites in their fur. The mites crawl onto the bat as they visit one inflorescence and climb off when they visit another. This is called phoresy. The bats are not harmed by these hitchhikers but are essential to the mite lifecycle. Thanks to their bat transports, the mites are able to make it to new feeding grounds far away from their original location. Though little is known about these mites, it has been suggested that the mites living on the coligallo palm are unique to that species and probably feed on no other plants.

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

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

Botanical Buoys

Photo by Doug McGrady licensed under CC BY 2.0

American featherfoil (Hottonia inflata) is a fascinating aquatic plant. It can be found in wetlands ranging from the coastal plains of Texas all the way up into Maine. Though widespread, American featherfoil is by no means common. Today I would like to introduce you to this gorgeous member of the primrose family (Primulaceae).

American featherfoil may look like a floating plant but it is not. It roots itself firmly into the soil and spends much of its early days as a vegetative stem covered in wonderful feathery leaves. It may be hard to find during this period as no part of it sticks above the water. To find it, one must look in shallow waters of ponds, ditches, and swamps that have not experienced too much disturbance. More on this in a bit.

Photo by Doug McGrady licensed under CC BY 2.0

American featherfoil lives life in the fast lane. It is what we call a winter annual. Seeds germinate in the fall and by late October, juveniles can be seen sporting a few leaves. There it will remains throughout the winter months until early spring when warming waters signal the growth phase. Such growth is rapid. So rapid, in fact, that by mid to late April, plants are beginning to flower. To successfully reproduce, however, American featherfoil must get its flowers above water.

The need to flower out of water is exactly why this plant looks like it is free floating. The flower stalks certainly do float and they do so via specialized stems, hence the specific epithet “inflata.” Each plant grows a series of large, spongy flowering stalks that are filled with air. This helps buoy the stems up above the water line. It does not float about very much as its stem and roots still anchor it firmly into place. Each inflorescence consists of a series of whorled umbels that vary in color from white to yellow, and even violet. Following pollination, seeds are released into the water where they settle into the mud and await the coming fall.

Photo by Doug McGrady licensed under CC BY 2.0

As I mentioned above, American featherfoil appreciates wetland habitats that haven’t experienced too much disturbance. Thanks to our wanton disregard for wetlands over the last century or so, American featherfoil (along with countless other species) has seen a decline in numbers. One of the biggest hits to this species came from the trapping of beavers. It turns out, beaver ponds offer some of the most ideal conditions for American featherfoil growth. Beaver ponds are relatively shallow and the water level does not change drastically from month to month.

Historically unsustainable levels of beaver trapping coupled with dam destruction, wetland draining, and agricultural runoff has removed so much suitable habitat and with it American featherfoil as well as numerous wetland constituents. Without habitat, species cannot persist. Because of this, American featherfoil has been placed on state threatened and endangered lists throughout the entirety of its range. With the return of the beaver to much of its former range, there is hope that at least some of the habitat will again be ready for American featherfoil. Still, our relationship with wetlands remains tenuous at best and until we do more to protect and restore such important ecosystems, species like American featherfoil will continue to suffer. This is why you must support wetland protection and restoration in your region!

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

Further Reading: [1] [2]

Twinspurs & Their Pollinators

© El Grafo / CC-BY-SA-3.0 (via Wikimedia Commons)

Garden centers and nurseries always have something to teach me. Though I am largely a native plant gardener, the diversity of plant life offered up for sale is always a bit mind boggling. Perusing the shelves and tables of myriad cultivars and varieties, I inevitably encounter something new and interesting to investigate. That is exactly how I came to learn about the twinspurs (Diascia spp.) and their peculiar floral morphology. Far from being simply beautiful, these herbaceous plants have evolved an interesting relationship with a small group of bees.

Diascia whiteheadii. Photo by Ragnhild&Neil Crawford licensed under CC BY-SA 2.0

The genus Diascia comprises roughly 70 species and resides in the family Scrophulariaceae. They are native to a decent chunk of southern Africa and have adapted to a range of climate conditions. Most are annuals but some have evolved a perennial habit. The reason these plants caught my eye was not the bright pinks and oranges of their petals but rather the two spurs that hang off the back of each bloom. Those spurs felt like a bit of a departure from other single-spurred flowers that I am used to so I decided to do some research. I fully expected them to be a mutation that someone had selectively bred into these plants, however, that is not the case. It turns out, those two nectar spurs are completely natural and their function in the pollination ecology of these plants is absolutely fascinating.

Diascia rigescens photo by Dinkum licensed under CC BY-SA 3.0

Not all Diascia produce dual spurs on each flower but a majority of them do. The spurs themselves can vary in length from species to species, which has everything to do with their specific pollinator. The inside of each spur is not filled with nectar as one might expect. Instead, the walls are lined with strange trichomes and that secrete an oily substance. It’s this oily substance that is the sole reward for visiting Diascia flowers.

Diascia megathura (a) inflorescenc with arrows indicating spurs and (b) cross sectioned spur showing the trichomes secreting oil (Photos: G. Gerlach).

If you find yourself looking at insects in southern Africa, you may run into a genus of bees called Rediviva whose females have oddly proportioned legs. The two front legs of Rediviva females are disproportionately long compared to the rest of their legs. They look a bit strange compared to other bees but see one in action and you will quickly understand what is going on. Rediviva bees are the sole pollinators of Diascia flowers. Attracted by the bright colors, the bees alight on the flower and begin probing those two nectar spurs with each of their long front legs.

If you look closely at each front leg, you will notice that they are covered in specialized hairs. Those hairs mop up the oily secretions from within each spur and the bee then transfers the oils to sacs on their hind legs. What is even more amazing is that each flower seems to have entered into a relationship with either a small handful or even a single species of Rediviva bee. That is why the spur lengths differ from species to species - each one caters to the front leg length of each species of Rediviva bee. It is worth noting that at least a few species of Diascia are generalists and are visited by at least a couple different bees. Still, the specificity of this relationship appears to have led to reproductive isolation among many populations of these plants, no doubt lending to the diversity of Diascia species we see today.

Diascia 'Coral Belle' Photo by KENPEI licensed under CC BY-SA 3.0

The female bees do not eat the oils they collect. Instead, they take them back to their brood chambers, feed them to their developing offspring, and use what remains to line their nests. At this point it goes without saying that if Diascia were to disappear, so too would these bees. It is incredible to think of the myriad ways that plants have tricked their pollinators into giving up most, if not all of their attention to a single type of flower. Also, I love the fact that a simple trip to a garden center unlocked a whole new world of appreciation for a group of pretty, little bedding plants. It just goes to show you that plants have so much more to offer than just their beauty.

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

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

The Wacky World of Whisk Ferns

Photo by Richard Droker licensed under CC BY-NC-ND 2.0

The whisk ferns (Psilotum spp.) are a peculiar group of plants. If you hang out in greenhouses long enough, you are most likely to encounter them as “weeds” growing in pots with other plants. Though they aren’t often put on display by themselves, the whisk ferns are certainly worth a closer look.

Psilotum comprises two species, the far more common Psilotum nudum and the lesser known P. complanatum. These two species will also hybridize, resulting in Psilotum × intermedium. Together, the whisk ferns make up one of only two genera in the family Psilotaceae (the other being Tmesipteris). They are strange plants to look at as there doesn’t appear to be much to them besides stems. Indeed, their peculiar morphology has earned them a fair share of taxonomic attention over the last century but before we get into that, it is a good idea to take a closer look at their anatomy.

Psilotum nudum with yellow sporagia. Photo by Mary Keim licensed under CC BY-NC-SA 2.0

What we see when we are looking at a whisk fern is the sporophyte generation. Like all sporophytes, its job is to produce the spores that will go on to make new whisk ferns. This part of the whisk fern lifecycle is pretty much all stem. Though these are in fact vascular plants, they do not produce true leaves. Instead, the branching stem takes up all of the photosynthetic work. What looks like tiny leaf-like scales are actually referred to as ‘enations.’ These structures do not contain any vascular tissue of their own. Instead, they bear a type of fused sporangia that house the spores. When mature, these will turn a bright yellow.

Underground, things aren’t much different. Whisk ferns produce a branching rhizome that is covered in hair-like projections called rhizoids. These structures not only help anchor the plant in place, they also function in a similar way to roots. Rhizoids interface with the soil environment allowing the plant to absorb nutrients and water. However, they don’t do this alone. Like so many other plants, whisk ferns partner with mycorrhizal fungi, which vastly increases the amount of surface area these plants have for absorbing what they need. In return, whisk ferns provide the fungi with carbohydrates they produce through photosynthesis. As lovely as this mutualistic relationship sounds, it actually starts off as parasitism.

A Psilotum rhizome with hair-like rhizoids. Photo by Curtis Clark licensed under CC BY-SA 3.0

When the spores find a suitable place to germinate, they will grow into the other half of the whisk fern lifecycle, the gametophyte. These resemble tiny versions of the rhizome and contain male and female reproductive organs. Living underground, the gametophytes do not photosynthesize. Instead, they completely rely on mycorrhizal fungi for all of their nutritional needs. This can go on for some time until the gametophytes are fertilized and grow a new sporophyte. Then and only then will the plant actually start giving back to the fungi that their lives depend on.

Psilotum complanatum with its flattened stems. Photo by Chad Husby licensed under CC BY-NC-ND 2.0

Because the overall form of the whisk ferns appears so “simplistic.,” many have hypothesized that the genus Psilotum is an evolutionary throwback to the early days of vascular plant evolution. On a superficial level, the whisk ferns do appear to have a lot in common with rhyniophytes, a group of plants that arose during the early Devonian, some 419 to 393 million years ago. A more detailed inspection of the anatomy of each group would reveal that there are some significant and fundamental differences between the two lineages, which I won’t go into here. Also, subsequent molecular work has shown that the whisk ferns reside quite comfortably within the fern lineage and likely represent a sister group to the order that gives us the adder’s tongue ferns (Ophioglossales). It would appear that whisk ferns more accurately represent a reduction in the more “traditional” fern form rather than a holdover from the early days of land plant evolution.

What the genus Psilotum lacks in number of species, it makes up for with its wide distribution. The whisk ferns seem to have conquered most of the tropical and subtropical landmasses on our planet. In fact, I found it incredibly difficult to discern much in the way of a native distribution for these plants. In some areas they are fairly common components of the local flora whereas in others they are considered rare or even threatened. I am sure that at least some of their expansive distribution can be attributed to human assistance as we move soils and plants around the world. To find them in nature, one must look in the cracks of rocks or on the trunks and branches of trees. Though both species can be found growing on trees, P. complanatum in particular seems to prefer an epiphytic lifestyle.

Psilotum complanatum (left) and Psilotum nudum (right) growing epiphytically. Photo by David Eickhoff licensed under CC BY 2.0

Whether you grow them on purpose, fight them as a greenhouse “weed,” or track them down in the wild, I hope you take a moment to appreciate these oddball plants. The whisk ferns are intriguing to say the least and certainly offer up a unique conversation piece for anyone curious about the botanical world. They are a genus worth admiring.

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

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

A New Case of Lizard Pollination from South Africa

With its compact growth habit and small, inconspicuous flowers tucked under its leaves, it seems like Guthriea capensis doesn’t want to be noticed. Indeed, it has earned itself the common name of '“hidden flower.” That’s not to say this plant is unsuccessful. In fact, it seems to do just fine tucked in among high-elevation rock crevices of its home range along the Drakensberg escarpment of South Africa. Despite its cryptic nature, something must be pollinating these plants and recent research has finally figured that out. It appears that the hidden flower has a friend in some local reptiles.

Lizard pollination is not unheard of ([1] & [2]), however, it is by no means a common pollination syndrome. This could have something to do with the fact that we haven’t been looking. Pollination studies are notoriously tricky. Just because something visits a flower does not mean its an effective pollinator. To investigate this properly, one needs ample hours of close observation and some manipulative experiments to get to the bottom of it. Before we get to that, however, its worth getting to know this strange plant in a little more detail.

The hidden flower is a member of an obscure family called Achariaceae. Though a few members have managed to catch our attention economically, most genera are poorly studied. The hidden flower itself appears to be adapted to high elevation environments, hence its compact growth form. By hugging the substrate, this little herb is able to avoid the punishing winds that characterize montane habitats. Plants are dioecious meaning individuals produce either male or female flowers, never both. The most interesting aspect of its flowers, however, are how inconspicuous they are.

The hidden flower (Guthriea capensis) in situ.

Flowers are produced at the base of the plant, out of site from most organisms. They are small and mostly green in color except for the presence of a few bright orange glands near the base of the style, deep within the floral tube. What they lack in visibility, they make up for in nectar and smell. Each flower produced copious amounts of sticky, sugar-rich nectar. They are also scented. Taken together, these traits usually signal a pollination syndrome with tiny rodents but this assumption appears to be wrong.

Based on hours of video footage and a handful of clever experiments, a team of researchers from the University of KwaZulu-Natal and the University of the Free State have been able to demonstrate that lizards, not mammals, birds, or insects are the main pollinators of this cryptic plant. Two species of lizard native to this region, Pseudocordylus melanotus and Tropidosaura gularis, were the main floral visitors over the duration of the study period.

Pseudocordylus melanotus

Tropidosaura gularis photo © 2009 Serban Proches licensed under CC BY-SA 2.5

Visiting lizards would spend time lapping up nectar from several flowers before moving off and in doing so, picked up lots of pollen in the process. Being covered in scales means that pollen can have a difficult time sticking to the face of a reptile but the researchers believe that this is where the sticky pollen comes into play. It is clear that the pollen adheres to the lizards’ face thanks to the fact that they are usually covered in sticky nectar. By examining repeated feeding attempts on different flowers, they also observed that not only do the lizards pick up plenty of pollen, they deposit it in just the right spot on the stigma for pollination to be successful. Insect visitors, on the other hand, were not as effective at proper pollen transfer.

Conspicuously absent from the visitation roster were rodents. The reason for this could lie in some of the compounds produced within the nectar. The team found high levels of a chemical called safranal, which is responsible for the smell of the flowers. Safranal is also bitter to the taste and it could very well serve as a deterrent to rodents and shrews. More work will be needed to confirm this hypothesis. Whatever the case, safranal does not seem to deter lizards and may even be the initial cue that lures them to the plant in the first place. Tongue flicking was observed in visiting lizards, which is often associated with finding food in other reptiles.

Male flower (a) and female flower (b). Note the presence of the orange glands at the base.

Another interesting observation is that the color of the floral tube and the orange glands within appear to match the colors of one of the lizard pollinators (Pseudocordylus subviridis ). Is it possible that this is further entices the lizards to visit the flowers? Other reptile pollination systems have demonstrated that lizards appear to respond well to color patterns for which they already have some sort of sensory bias. Is it possible that these flowers evolved in response to such a bias? Again, more work will be needed to say for sure.

By excluding vertebrates from visiting the flowers, the team was able to show that indeed lizards appear to be the main pollinators of these plants. Without pollen transfer, seed set is reduced by 95% wheres the additional exclusion of insects only reduced reproductive success by a further 4%. Taken together, it is clear that lizards are the main pollinators of the enigmatic hidden flower. This discovery expands on our limited knowledge of lizard pollination syndromes and rises many interesting questions about how such relationships evolve.

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

Further Reading: [1] [2]

An Intruiguing Relationship Between Ants and Cacti

The extrafloral nectaries of Pachycereus gatesii appear as tiny red bumps just below the areole.

It’s hard to think of a group of plants that are better defended than cacti. Frequently and often elaborately adorned with vicious spines, these succulents make any animal think twice about trying to take a bite. And yet, for some cacti, spines don’t seem to cut it. A surprising amount of species appear to have taken their defense system to a whole new level by recruiting nature’s most tenacious bodyguards, ants.

Plants frequently have a friend in ants. Spend some time observing ants at work and it’s east to see why. These social insects have numbers and strength on their side. Give ants a reason to be invested in your survival and they will certainly see to it that nothing threatens this partnership. For cacti, this involves the secretion of nectar from specialized tissues called extrafloral nectaries.

Extrafloral nectaries are not unique to cacti. A multitude of plant species produce them, often for similar reasons. Ants love a sugary food source and the more reliable that source becomes, the more adamant an ant colony will be at defending it. The odd thing about cacti is that they don’t seem to have settled on a single type of extrafloral nectary to do the trick. In fact, as many as four different types of extrafloral nectaries have been described in the cactus family.

Ants visiting the extrafloral nectaries covering the developing flowers of Pilosocereus gounellei.

Some cacti secrete nectar from highly modified spines. A great example of this can be seen in genera such as Coryphantha, Cylindropuntia, Echinocactus, Ferocactus, Opuntia, Sclerocactus, and Thelocactus. Such spines are usually short and blunt, hardly resembling spines at all. Other cacti secrete nectar from regular looking spines. This adaptation is odd as there does not seem to be anything special about the anatomy of such spines. Examples of this can be seen in genera such as Brasiliopuntia, Calymmanthium, Harrisia, Opuntia, Pereskiopsis, and Quiabentia. Still others secrete nectar from highly reduced leaves that are found at the base of where the spines originate (the areole). Such leaves have been described in Acanthocereus, Leptocereus, Myrtillocactus, Pachycereus, and Stenocereus. They aren’t easy to recognize as leaves either. Most look like tiny scales. Finally, the fourth type of extrafloral nectary comes in the form of specialized regions of the stem tissue. This has been described in genera such as Armatocereus, Leptocereus, and Pachycereus.

Highly modified spines functioning as extrafloral nectaries in Ferocactus emoryi.

Seemingly normal spines of Harrisia pomanensis secreting nectar.

Regardless of where they form, their function remains much the same. They secrete a form of nectar which ants find irresistible. The more reliable this food source becomes, the more aggressive ant colonies will be in defending it. This is an especially useful form of defense when it comes to small insect herbivores. Whereas spines deter larger herbivores, they don’t do much to deter organisms that can just slip right through them unharmed. Ants also clean the cacti, potentially removing harmful microbes like fungi and bacteria. Though we are only just beginning to understand the depths of this cactus/ant mutualism, what we have discovered already suggests that the relationship between these types of organisms is far more complex than what I have just outlined above.

For instance, it may not just be sugar that the ants are looking for. In arid desert habitats, water may be the most limiting resource for an ant colony and large, succulent cacti are essentially giant water reservoirs. The key is getting to that water. One study that looked at a species of barrel cactus growing in Arizona called Ferocactus acanthodes found that as spring gives way to summer, the concentration of sugars secreted by the extrafloral nectaries decreases. As a result, the nectar becomes far more watery. Amazingly, ant densities on any given barrel cactus actually increased throughout the summer, despite the fact that the nectar was being watered down. Ants are notoriously prone to desiccation so it stands to reason that water, rather than sugar, is the real prize for colonies hanging out on cacti in such hot desert environments.

The incredible floral display of Ferocactus wislizeni, a species whose reproductive efforts are affected by the types of ants they attract. Photo by Joseph j7uy5 licensed under CC BY-NC-SA 2.0

Another interesting observation about the cactus/ant mutualism is that it appears that the identity of the ants truly matters. Though defense is the main benefit to the cactus, research suggests that there is a tipping point in how much such defenses benefit cacti. It has been found that although cacti initially benefit from anti-herbivore and cleaning services, extra aggressive ant species can actually drive off potential pollinators. At least one study has shown that when less aggressive ant species tend cacti, they produce more fruits and those fruits contain significantly more seeds than cacti that have been tended by extremely aggressive ant species. This is especially concerning when we think about the growing issue of invasive ants. As more and more non-native ant species displace native ants, this could really tip the balance for some cactus species.

Despite all of the interesting things we have learned about extrafloral nectaries in the family Cactaceae, there are so many questions yet to be answered. For starters, we still do not know how many different taxa produce them in one form or another. It is likely that closer inspection, especially of rare or poorly understood groups, will reveal that far more cacti produce some type of extrafloral nectary. Also, we know next to nothing about the anatomy of the different types of nectaries. How do they differ from one another and how do some, especially those derived from ordinary spines, actually function? Finally, do these nectaries function year round or is there some sort of seasonal pattern to their development and utility. How does this affect the types of ants they attract and how does that in turn affect the survival and reproduction of these cacti? For such a charismatic group of plants as cacti, we still have to much to learn.

Photo Credits: Thanks to Dr. Jim Mauseth and Dr. John Rebman and Dr. Silvia Rodriguez Machado for use of their photos [1] [2]

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

The Prairie Peninsula

North American prairies are some of the most endangered habitat types on the planet. Once covering vast swaths of the continent representing the arid rain shadow of the Rocky Mountains, the prairies now occupy only 1% of their former range. We have converted most of that land into agriculture and sub-developments. It may surprise many of you to learn that following the retreat of the Pleistocene glaciers, a subset of prairie ecosystem once stretched a lot further east than one would expect for a prairie. This grassland ecosystem ranged far up into the northeast and even met the Atlantic Coast in parts of New Jersey and Long Island. This was known as the prairie peninsula and today its remnants represent some of the rarest prairie ecosystems that North America has left. 

Ecologists believe that the prairie peninsula owed its existence to an intriguing quirk of the climate at that time. During interglacial periods, eastern North America’s climate was much warmer and drier than it is today. Because of this, prairie grasslands were hypothesized to have migrated east, following the recently exposed terminal moraines that the glaciers left in their wake. Moraine soils tend to be composed of unconsolidated till and are quick to drain water,  which provided perfect conditions for prairies to develop. This prairie peninsula preceded the invasion of trees, which now make up the forests that dominate eastern North America. Today we refer to the remnants of this prairie peninsula as "heaths" or "barrens." Despite their rarity, ecologists have long debated whether such habitats are truly echoes of our glacial past or products of a more recent, cultural clearing of the land.

A relatively recent paper sheds some light on this debate using some pretty clever detective work. The author used a variety of insects, but mainly focused on spittlebugs, to show that these eastern prairies remnants are, indeed, relics of our glacial past. Apparently, insects like leafhoppers, froghoppers, and spittlebugs are often extremely specialized on specific species of plants, mainly grasses. They feed much like aphids do, by sucking the juices out of the plants vascular tissues. Many of these insects are flightless or at least do not travel great distances from where they were born. If one were to find certain species on the eastern prairies, it would provide strong evidence in support of prairie migration from west to east via the glacial moraines.

The evidence suggests exactly that. Eastern pockets of remnant heaths and barrens do in fact host many of these prairie specialists. What is more interesting is that this research has shown that we can track, with some certainty, the migrational patterns of these ecosystems. As expected, the prairies moved in from the west, during an interglacial period much warmer than now. As they moved across the eastern US, they ran into the Appalachian Mountains, which is a formidable barrier to say the least.

How did the prairies circumvent this obstacle and end up in pockets along the Atlantic coast? Evidence points to sandy sediments in the numerous valleys along the spine of the Appalachians. Most of these sediments no longer exist due to erosion and out-crowding by forests, causing the current disjunction we see in these rare eastern prairie grasslands. It is amazing to me to think that these pockets of  habitat have existed for centuries, despite all the changes we have laid upon the land. These results are a wonderful example of the uniqueness of these habitats and, now more than ever, show us how much these deserve our attention so that maybe they can persist into the future. It also highlights just how special these ecosystems truly are. They are not something created by the hand of man. Instead, these habitats have survived the test of time. Now they must survive us. Support your local land conservancy today!

Map Credit: [1]

Further Reading: [1]

Meeting One of North America's Rarest Oaks

A post (and photos) by Robbie Q. Telfer

“Every species is a masterpiece, exquisitely adapted to the particular environment in which it has survived.”

-- E.O. mothereffin Wilson

One of the perks of working at The Morton Arboretum is you get to see cool lectures on tree science for free. At one such program, Dr. Mary Ashley from the University of Illinois at Chicago was sharing her research on oak pollen and how far it can travel to fertilize female flowers (far). She looked at not only trees in the Chicago region, but also oaks off the coast of California and in the Chihuahuan Desert of west Texas, as well as throughout Mexico. That latter oak was a shrubby species called Quercus hinckleyi or Hinckley oak. It is able to spread pollen over far distances as well, despite the fact that there are only 123 individuals known to be left. IUCN lists it as Critically Endangered.

As she was telling us this, it occured to me that I would be in West Texas soon to visit my sister-in-law, so afterwards I approached Dr. Ashley and asked if there was any way I could have the coordinates of Q. hinckleyi so that I could visit it, take a selfie, and luxuriate in the presence of something so rare. I made it clear to her that I understood just how important it was to keep this information a secret, because the last thing this relict needs is to be uprooted by poachers. Which I wish wasn’t a concern, but it is.

Dr. Ashley put me in touch with her colleague Janet Backs who graciously shared the coordinates. I could see the plants from Google maps satellite view. There they were. I probably waved at the computer screen sheepishly.

As I waited for my time to bask in the majesty of botanical greatness, I consulted my copy of Oaks of North America (1985) by Howard Miller and Samuel Lamb to see what the entry for hinckleyi said.

Notably, it mentions that “This is another of the oaks with no specific value, except as a curiosity.” More on that later.

After much anticipation, the time was upon us. I decided to drive out to the plants in my rental first thing in the morning after getting to Texas. The Chihuahuan Desert is an astounding place that my Illinoisan eyes weren’t altogether prepared for. It is perhaps the most biodiverse desert in the world, and compared to our prairies, woodlands, and wetlands, it feels like a different planet. Some of the cooler plants I got to see were tree cholla (Cholla sp.), Havard’s century plant (Agave havardiana), Wright’s cliffbrake (Pellaea wrightiana), and little buckthorn (Condalia ericoides). And also a family of introduced aoudads with TWO adorable babies. I also got to see my first javelina (as roadkill) and all kinds of birds new to me.

Tree cholla (Cholla sp.)

Havard’s century plant (Agave havardiana)

Wright’s cliffbrake (Pellaea wrightiana)

Little buckthorn (Condalia ericoides)

Aoudads in the distance.

Finally I got to the coordinates - luckily google preloaded the directions on my phone because there was absolutely no cell service where I was. I parked and walked to the plants. And lo, I present to you, Quercus hinckleyi.

It’s in the white oak family, which I guess means more than just “has round leaves.” These leaves look like holly, and even the shed ones on the ground still had some stabbiness left in them. It’s quite diminutive - certainly compared to any oak I’ve ever seen and even by shrub standards. I’d pinch its cheeks if that wouldn’t make my fingers bleed. After getting the pics I needed and doing the atheist’s version of saying a prayer over it, I floated back to my car like a cartoon cat in love.

The rest of the trip was great and I can’t wait to go back.

Since returning, I have shown several of my non-plant nerd friends the pics of hinckleyi and they seem politely impressed but not, like, actually impressed. This is totally understandable! If your experience with plants is on the order of what looks best in a planting or what tastes best in your tummy, this shrub is not for you. After all “it’s only value is as a curiosity.”

I don’t know about that. I feel like it’s value is greater than that for humans - it’s a window into the North American continent before the climate shifted 10,000 years ago, it’s an individual member of our vast botanical heritage, it is unique, it is adorbs, and it helped Dr. Ashley, and therefore us, understand more things about the movement of oak pollen.

But beyond what it does for US, what if, and hear me out, what if it has a right to existence on its own, without being displaced by pipelines or aoudads or poachers? It is a member of its ecological community, and just like I feel a loss when a member of my community passes, we don’t have the language to articulate what is felt when a member of an ecosystem winks out forever.

Janet Backs told me that she heard of someone who was trying to poach acorns from a subpopulation of hinckleyi and that the landowners where that shrub is actually chased those folks for miles and miles down the road. I love that. I wish every single threatened species/subpopulation had someone who understood its value beyond what it does for humans enough to chase people, possibly with a gun, for miles and miles.

I have had a paltry bucket list for most of my adult life - boring stuff like meeting my heroes or getting to a 7th bowl of never-ending-pasta. But despite their apparent lack of reverence for Q. hinckleyi I think a pretty good guiding list for me would be to visit each of the 77 oaks of North America in their native habitats. I know they won’t all be as special as this experience, but what better way to visit the corners of this continent and its myriad ecological communities, than by visiting each of its oaks? I currently can’t think of any, and would invite anyone to, if not fund me, join me.

An Iris With Multiple Parents

The Abbeville iris (Iris nelsonii) is a very special plant. It is the rarest of the so-called “Louisiana Irises” and can only be found growing naturally in one small swamp in southern Louisiana. If you are lucky, you can catch it in flower during a few short weeks in spring. The blooms come in a range of colors from reddish-purple to nearly brown, an impressive sight to see siting atop tall, slender stems. However, the most incredible aspect of the biology of this species is its origin. The Abbeville iris is the result of hybridization between not two but three different iris species.

When I found out I would be heading to Louisiana in the spring of 2019, I made sure that seeing the Abbeville iris in person was near the top of my to-do list. How could a botany nut not want to see something so special? Iris nelsonii was only officially described as a species in 1966. Prior to that, many believed hybridization played a role in its origin. Multiple aspects of its anatomy appear intermediate between other native irises. It was not until proper molecular tests were done that the picture became clear.

The Abbeville iris genome contains bits and pieces of three other irises native to Louisiana. The most obvious parent was yet another red-flowering species - the copper iris (Iris fulva). It also contains DNA from the Dixie iris (Iris hexagona) and the zig-zag iris (Iris brevicaulis). If you had a similar childhood as I did, then you may have learned in grade school biology class that hybrids are usually biological dead ends. They may exhibit lots of beneficial traits but, like mules, they are often sterile. Certainly this is often the case, especially for hybrid animals, however, more and more we are finding that hybridization has resulted in multiple legitimate speciation events, especially in plants.

Iris fulva. Photo by Richard licensed under CC BY-NC-ND 2.0

Iris hexagona. Photo by beautifulcataya licensed under CC BY-NC-ND 2.0

Iris brevicaulis. Photo by peganum licensed under CC BY-SA 2.0

How exactly three species of iris managed to “come together” and produce a functional species like I. nelsonii is interesting to ponder. Each of its three parent species prefers a different sort of habitat than the others. For instance, the copper iris is most often found in seasonally wet, shady bottomland hardwood forests as well as the occasional roadside ditch, whereas the Dixie iris is said to prefer more open habitats like wet prairies. In a few very specific locations, however, these types of habitats can be found within relatively short distances of each other.

Apparently at some point in the past, a few populations swapped pollen and the eventual result was a stable hybrid that would some day be named Iris nelsonii. As mentioned, this is a rare plant. Until it was introduced to other sites to ensure its ongoing existence in the wild, the Abbeville iris was only known to occur in any significant numbers at one single locality. This necessitates the question as to whether or not this “species” is truly unique in its ecology to warrant that status. It could very well be that that single locality just happens to produce a lot of one off hybrids.

In reality, the Abbeville iris does seem to “behave” differently from any of its parental stock. For starters, it seems to perform best in habitats that are intermediate of its parental species. This alone has managed to isolate it enough to keep the Abbeville from being reabsorbed genetically by subsequent back-crossing with its parents. Another mechanism of isolation has to do with its pollinators. The Abbeville iris is intermediate in its floral morphology as well, which means that pollen placement may not readily occur when pollinators visit different iris species in succession. Also, being largely red in coloration, the Abbeville iris receives a lot of attention from hummingbirds.

Although hummingbirds do not appear to show an initial preference when given the option to visit copper and Abbeville irises at a given location, research has found that once hummingbirds visit an Abbeville iris flower, they tend to stick to that species provided enough flowers are available. As such, the Abbeville iris likely gets the bulk of the attention from local hummingbirds while it is in bloom, ensuring that its pollen is being delivered to members of its own species and not any of its progenitors. For all intents and purposes, it would appear that this hybrid iris is behaving much like a true species.

As with any rare plant, its ongoing survival in the wild is always cause for concern. Certainly Louisiana is no stranger to habitat loss and an ever-increasing human population coupled with climate change are ongoing threats to the Abbeville iris. Changes in the natural hydrologic cycle of its swampy habitat appear to have already caused a shift in its distribution. Whereas it historically could be found in abundance in the interior of the swamp, reductions in water levels have seen it move out of the swamp and into ditches where water levels remain a bit more stable year round. Also, if its habitat were to become more fragmented, the reproductive barriers that have maintained this unique species may degrade to the point in which it is absorbed back into an unstable hybrid mix with one or a couple of its parent species. Luckily for the Abbeville, offspring have been planted into at least one other location, which helps to reduce the likelihood of extinction due to a single isolated event.

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