I’ve grown to really appreciate cucurbits (family Cucurbitaceae) in recent years. From their ambling/climbing habit and often delicious fruits to their beautiful flowers and intimate relationships with a few native bees, this family has a lot to offer. Of course, there are few better ways to get to know plants than by growing them in and around your home and, at least at our place, this summer will go down in history as the summer of the gourd. We are currently growing a handful of species and cultivars and I get a great deal of enjoyment out of watching them grow up the trellis we have provided.

As they climb, cucurbits send out long, thin tendrils (which are actually modified stems) that grab on and wind around any surface they touch. This happens surprisingly quick too. Within only a few minutes of touching a surface, individual tendrils will begin to wind themselves around it. This phenomenon has fascinated people for centuries. I don’t doubt it amused the indigenous cultures that first began cultivating them for food and that amusement continues till this day. Do a web search for cucumber tendrils and you will find countless pictures and blogs showcasing this wonderful anatomical habit.

Despite all the attention, the mechanisms behind this behavior have largely remained a mystery until quite recently. We have known that the initial curling of the tendril is induced by touch. As soon as the cells within the tendril sense contact with a surface, the signal is sent to begin curling. But how do they curl so quickly?

The key to this behavior lies in a two-layered band of specialized cells that run the length of the tendril. Once the signal that the tendril has touched an object has been received, these bands swing into action. One layer of cells will immediately begin to expel water, causing them to contract. Meanwhile, the other layer of cells becomes increasingly stiff and lignified. This creates tension along the length of the tendril, causing it to bend. Oddly enough, this doesn’t happen in the same direction. Take a close look at the tendrils on a cucumber or squash vine and you will notice that each tendril curls in two different directions, separated by a kink or “perversion” (as it is known in the literature) in the middle. This is because the layer of cells on the band that shrinks is different whether you are near the tip or near the base of the tendril.

As many of you reading this are already well aware, the tendrils help to secure the plants as they climb. However, the story is much more interesting than simply anchoring the plants in place. The curling of the tendrils is extremely important when it comes to structural support. If the tendrils did not curl, the plant would be anchored in place with very little wiggle room. As big gusts of wind cause the plant to thrash to and fro or a heavy limb comes crashing down from above, a straight tendril would be far more likely to break under the strain. By adding those opposite twists, the tendrils are able to flex a lot, providing enough movement to keep them from breaking under stress.

If you watch how the tendrils develop over time, their amazing structural support gets even cooler. When stretched, a metal spring looses a lot of its springy-ness. This is not the case for cucurbit tendrils. When stretched, they not only return to their original shape, they curl even tighter. This way, the plant is able to secure itself with varying intensities, allowing for fine tuned adjustments to its structural support. The amount of curling also changes with age. Older tendrils tend to curl more tightly than younger tendrils, especially under strain. As the plant grows, older portions of the stem secure themselves much more strongly via their tendrils. Alternatively, the younger growing portions of the stem need to be a bit more flexible as they anchor themselves to whatever they are climbing on.

So there you have it. The aesthetically pleasing, curly tendrils of your cucurbits serve a very important function in the growth of the plant. Without them, these plants would not only have a hard time climbing, they would also be knocked down by every minor disturbance. The key to their success as vines lies in highly modified stems with an intriguing band of specialized cells that provide them with a physically sound anchoring mechanism.

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Further Reading: [1] [2]

While working in the garden the other day, I noticed that some of the nodding onion (Allium cernuum) we planted last year had finally come into bloom. I must have spent the good part of an hour watching bees pay a visit to their downward pointing flowers. I have seen a lot of onion species in bloom before, but this particular native is the only one that I know of personally that orients its flowers facing the ground. This got me to thinking about floral orientation. A lot of plants produce flowers that face the ground but many more do not. Why is there such variety among the orientation of flowers?

As always, I hit the literature. It turns out, many scientists have set out to investigate the function of floral orientation. What immediately stuck out to me is just how many different flowering plant lineages boast species whose flowers face down rather than out or up. I knew instantly that with so much variety in lineage, habitat, and pollination strategies, the answer wasn’t going to be simple or straight forward. Indeed, each investigation I read about seemed to end in a slightly different conclusion. Still, there were enough patterns among the results and conclusions to make some general statements about the subject.

We often find plants with downward facing flowers in harsh climates. Harsh can mean a lot of different things depending on the plant and region in question but take, for instance, the case of the genus Cremanthodium. This interesting group of asters resemble sunflowers in the basic appearance of their flowers but the plants themselves are vastly different in overall growth habit. Many hail from alpine environments in Asia and possess a short stature and flowers that face the ground instead of the sun. Research on the reproductive habits of these plants has revealed that the downward orientation of their flowers helps protect the sensitive reproductive parts from solar radiation and rain.

Growing at high elevations exposes these plants to lots of UV radiation and plenty of storms. If flowers were to orient towards the sky, rain could wash away pollen and UV radiation could really hinder successful reproduction. By facing the ground, the flowers are able to avoid these potentially harmful effects altogether. Similar results have been found for other members of the aster family in the genus Culcitium growing in alpine habitats in the Andes. Here again we see that downward pointing flowers help protect the sensitive reproductive parts from rain, snow, and too much sun.

However, its not just the elements that have selected for downward pointing flowers. As you can probably imagine, pollinators also play a role in floral orientation. While watching bee visit our nodding onions, I noticed that they seem to be much better able to land on and collect pollen and nectar from downward pointing flowers than any of the flies I see attempting visits. Indeed, floral orientation can have a massive impact on what kinds of pollinators are able to effectively visit a flower.

A great example of this can be seen in members of the genus Zaluzianskya. Some species present their flowers horizontally or vertically while others present their flowers facing the ground. By comparing the visitors that frequent each species, researchers found that orientation matters. Upright or horizontally facing flowers were mostly visited by hawkmoths. Hawkmoths hover while they feed, which means they have a much harder time visiting downward facing flowers. By presenting their flowers in different orientations, the various species of Zaluzianskya ensure that only specific pollinators are able to access their rewards and thus achieve pollination. As such, upright, horizontal, and downward flowering species remain reproductively isolated from one another. Similar results have also been found in genera such as the afore mentioned Culcitium as well as some Commelina and 香港梯子.

I am sure many more examples exist out there but alas, I only have so much time to pursue my random curiosities these days. Nonetheless, what started as a fun observation in the garden turned into an entertaining dive into ideas that I had not given too much thought to before. What seems like a funny quirk of anatomy turns out to have massive implications for where plants are able to grow and how they are able to reproduce and all of these factors and more have shaped flowering plant evolution over time. Not bad for a few hours in the garden.

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

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

There was a time when I thought that all orchids were finicky botanical jewels, destined to perish at the slightest disturbance. Certainly many species fit this description to some degree, but more often these days I am appreciating the role disturbance can play in maintaining many orchid populations. Seeing various genera like Platanthera or Goodyera thriving along trails and old dirt roads, lawn orchids (Zeuxine strateumatica) growing in manicured lawns, or even various Pleurothallids growing on water pipes in the mountains of Panama has opened my eyes to the diversity of ecological strategies this massive family of flowering plants employs.

Of the examples mentioned above, none can hold a candle to the hardiness of the broad-leaved helleborine orchid (Epipactis helleborine) when it comes to thriving in disturbed habitats. Originally native throughout much of Europe, North Africa, and Asia, this strangely beautiful orchid can now be found growing throughout many temperate and sub-tropical regions of the world. Indeed, this is one species of orchid that has greatly benefited from human disturbance. In fact, I more often see this orchid growing in and around cities and along roadsides than I do in natural settings (not to say it isn’t there too). In many areas here in North America, the broad-leaved helleborine orchid has gone from a naturalized oddity into a full blown invasive.

Much of its success in conquering new and often highly disturbed territory has to do with its relationship with mycorrhizal fungi. Like all orchids, the broad-leaved helleborine orchid requires fungi for germination and growth, relying on the symbiotic relationship into maturity. Without mycorrhizal fungi, these orchids could not survive. However, while many orchids seem to be picky about the fungi they will partner with, the broad-leaved helleborine is something of a generalist in this regard. At least one study in Europe was able to demonstrate that over 60 distinct groups of mycorrhizal fungi were able to partner with this orchid. By opening itself up to a wider variety of fungal partners, the broad-leaved helleborine orchid is able to live in places where pickier orchids cannot.

Another key to this orchids success has to do with its pollination strategy. Here again we see that being a generalist comes with serious advantages. Though wasps are thought to be the most effective pollinators, myriad other insects from various kinds of flies to beetles and butterflies will visit these blooms. How is it that this orchid has become to appealing to such a wide variety of insects? The answer is chemistry.

The broad-leaved helleborine orchid is something of a skilled chemist. When scientists analyzed the nectar produced in the cup-shaped lip of the flower, they found a diverse array of chemicals, many of which lend to some incredible insect interactions. For starters, highly scented compounds such as vanillin (the compound responsible for the vanilla scent and flavor of Vanilla orchids) are produced in the nectar, which certainly attracts many different kinds of insects. There is also evidence of some floral mimicry going on as well.

Scientists found a group of chemicals called kairomones in broad-leaved helleborine nectar, which are very similar to aphid alarm pheromones. When released by aphids, they warn nearby kin that predators are in the area. In one sense, the production of these compounds in the nectar may serve to ward off aphids looking for a new place to feed. However, these chemicals also appear to function as pollinator attractants. For aphid predators like hoverflies, these pheromones act as a dinner bell, signalling good egg laying sites for gravid female hoverflies whose larvae gorge themselves on aphids as they grow. It just so happens that hoverflies also serve as important pollinators for the broad-leaved helleborine orchid.

A series of compounds broadly classified as green-leaf volatiles were found in the nectar as well. Many plants produce these compounds when their leaves are damaged by insect feeding. Like the aphid example above, green-leaf volatiles signal to nearby predatory insects that plump herbivores are nearby. For instance, when the caterpillars of the cabbage white butterfly feed on cabbage plants, green-leaf volatiles attract wasps, which quickly set to work eating the caterpillars, relieving the plant of its herbivores in the process. As previously mentioned, wasps are thought to be the main pollinators for this orchid so attracting them makes sense. However, attracting pollinators using chemical trickery can be risky. What happens when a pollinator shows up and realizes there is no plump aphid or caterpillar to eat?

The answer to this comes from a series of other compounds produced in this orchid’s nectar. Few insects will turn down a sugary meal and upon visiting a bloom, and indeed, many visitors end up sipping some broad-leaved helleborine nectar. Sit back and watch and it won’t take long to realize that these insects appear to quickly become intoxicated. Their behavior becomes sluggish and they generally bumble around the flowers until they sober up and fly off. This is not happenstance. This orchid actively gets its pollinators wasted, but how?

Along with the chemicals we already touched on, scientists have also found a plethora of narcotics in broad-leaved helleborine nectar. These include various types of alcohols and even chemicals similar to that of opioids like Oxycodone. Now, some have argued that the alcohols are not the product of the plant but rather the result of fermentation by yeasts and bacteria living within the nectar. However, the presence of different antimicrobial compounds coupled with the sheer concentrations of alcohols within the nectar appear to discount this hypothesis and point to the plant as the sole creator. Nonetheless, after a few sips of this narcotic concoction, insects like wasps and flies spend a lot more time at each flower than they would if they remained sober the whole time. This has led to the suggestion that narcotics help improve the likelihood of successful pollination.

Indeed, the broad-leaved helleborine orchid seems to have no issues with sex. Most plants produce a bountiful crop of seed-laden fruits each summer. In fact, it has been found that plants growing in areas of high human disturbance tend to set more seed than plants growing in natural areas. Scientists suggest this is due to the wide variety of pollinators that are attracted to the complex nectar. Human environments like cities tend to have a different and sometimes more varied suite of insects than more rural areas, meaning there are more opportunities for run ins with potential pollinators.

The broad-leaved helleborine orchid stands as an example of the complexities of the orchid family. Few orchids are as generalist in their ecology as this species. Its ability to grow where others can’t while taking advantage of a variety of pollinators has lent to the extreme success of this species world wide.

Photo Credit: [1]

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

The sessile nature of plants means that they are strongly shaped by their environment. Natural selection is constantly at work on plants but that doesn’t mean that plants don’t shape their environment as well. When I think about the impact of plants on resident animal communities, I am always reminded of a quote by American ethnobotanist Terence McKenna, “Animals are something invented by plants to move seeds around.” Now, I realize that the animal kingdom got its start long before plants came onto the scene but there are many threads of truth to this quote.

Take, for instance, the case of the two-marked treehopper (Enchenopa binotata). This wonderful little insect enjoys a distribution that encompasses much of North and Central America, ranging from Canada down into Panama. Not only do these treehoppers look cool with their intriguing color pattern and that thorny pronatum, but their ecology and evolutionary history is absolutely fascinating as well. The existence of these treehoppes is entirely tied to the trees on which they live and breed. More over, while the two-marked treehopper may look like a single species, it is actually a complex of multiple cryptic “species” whose entire identity is owed to their preferred host tree.

The two-marked treehopper is not a species that moves around the landscape very much. While males will venture out into the environment in search of mates, females tend to live out their whole lives feeding and breeding on their tree upon which they were born. After mating, a female will lay her eggs within the stem of the host tree. The eggs overwinter in a sticky secretion called “egg froth.” This egg froth not only protects the eggs, it is also full of pheromones that signal to other females in the area to lay their eggs near by. The nymphs of the two-marked treehopper are gregarious. There is safety in numbers and the more nymphs hanging out on a branch, the less likely any single individual will be attacked by a predator.

Come spring, as trees begin to break dormancy, eggs laid the previous summer get the cue to hatch as sap begins to flow. Since treehoppers are sap feeders, this signal is essentially a ringing dinner bell. Apparently the specificity of this sap feeding habit is one of the reasons these treehoppers are so specific about their host.

As I mentioned earlier, the two-marked treehopper is not a single species but rather a complex of distinct taxonomic units. All of this cryptic diversity has to do with their preferred trees as each species within the complex feeds and breeds on a specific genus of tree/shrub: Carya, Celastrus, Cercis, Juglans, Liriodendron, Ptelea, Robinia, and Viburnum. Because no two tree species are alike, each has its own phenology. Different trees leaf out and begin growth at different times. Different tree species have different chemicals and nutrients in their sap. Also, different tree species have different wood densities. All of these factors and more have left their mark on the evolution of two-marked treehoppers.

Because females generally don’t leave the trees on which they were born, their offspring will inevitably be born on the same species of tree. This means they will be raised on a diet of the same sap as their mother. As mentioned, different trees produce different kinds of sap, which means that the digestive systems of these insects become highly tuned to their specific host tree. By experimentally moving two-marked treehopper nymphs to different host trees and tracking their development, scientists have also been able to demonstrate that host switching does not work well for the treehoppers. Nymphs raised on species different than the tree on which their eggs were laid do not develop as well or at all. It appears that their specific feeding habits are entirely tuned to the chemical composition of their host sap.

Additionally, the phenology of their host tree life cycle means that species raised on different trees rarely sync up in nature. Some trees force their resident treehoppers to emerge and mate earlier than others and vice versa. Evidence for this was made even stronger by studying these dynamics in the human environment.

The preferred hosts of two-marked treehoppers rarely grow in the same habitats in nature. However, thanks to our gardening and landscaping efforts, it isn’t hard to find these species in close proximity in the human environment. In cases where different host trees are found only a few meters from one another, the specific feeding requirements of each species means that species barriers among different treehopper populations are maintained. However, even before offspring enter into the picture, host trees also seem to have an effect on two-marked treehopper mating habits.

Treehoppers are surprisingly musical creatures. Though we can’t hear them without the help of microphones, treehoppers utilize different types of vibrational calls to communicate with one another. This is especially true during mating. Males make repeated vibrations on the stems that the females will then respond to. By studying variations in these calls, scientists have found that two-marked treehoppers living on different trees produce vastly different calls. They key to this appears to lie in the ability of vibrations to travel through wood. Just as different types of wood work well for different types of instruments, the differences in wood density of their host trees affect how their mating calls travel and are eventually perceived. In other words, with a bit of training and some good recordings, you could identify the tree on which a two-marked treehopper lives just by its calls.

The ecological barriers between these insects are maintained no matter how close they are to one another and it is all thanks to the biology of the trees on which they live. Keep an eye out for these wonderful little insects. They are a joy to watch and offer us plenty of examples of evolution in action.

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

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

Plants, like all living organisms, must be able to sense and respond to their environment. The more we look at these sessile organisms, the more we realize that plants are far from static in their day to day lives. Recent evidence even suggests that some plants may be able to “hear” their pollinators and react accordingly.

I place the word “hear” in quotes because I want to make sure that we are not talking about hearing in an animalistic sense. Plants do not have ears, a nervous system, or anything like a central processing unit to make sense of such stimuli. What they do have are mechanoreceptors that can sense vibrations and those are what are likely at work in this example.

The beach evening primrose (Oenothera drummondii) is native to southeastern North America. It is pollinated by bees during the day and by moths at night. Like most members of its genus, O. drummondii produces relatively large, showy flowers. That doesn’t mean it steals all of the attention though. Competition for pollinators can be stiff among flowering plants. To sweeten the deal a bit, O. drummondii also produces a fair amount of nectar.

Nectar is costly for plants to produce and maintain. Not only does it take water and carbohydrates away from the rest of the plant, it also puts the reproductive structures at risk of degradation by microbes feeding on sugars as well as nectar thieves who end up drinking the nectar without pollinating the flower. It stands to reason that a plant that can modulate the quality of its nectar reward in response to pollinator availability could potentially increase its fitness. If the plant doesn’t always have to present sugar-rich nectar then why bother? It appears that selective nectar production is exactly the strategy ssssr香港梯子 employs.

Researchers have discovered that individual O. drummondii flowers can rapidly increase the sugar content of their nectar after being exposed to the sound of a visiting bee. Within 3 minutes of being exposed to playbacks of bee wings, the flowers of O. dummondii increased the sugar content of their nectar by 20%. What’s more, flowers that had sensed the vibrations and increased their sugar content were more likely to be visited by bees. This is because bees are really good at sensing the sugar content of nectar.

This is pretty remarkable. Not only does this enable the plant to respond to the availability of pollinators and reduce the chances of nectar spoilage and theft, it significantly increases their chances of pollination. The fact that the response is so rapid (~3 mins) likely stems from the foraging habits of bees, who prefer to limit the amount of time between floral visits. Thus, the faster the plant can respond, the more likely that bees are willing to stick around and visit more flowers.

In terms of a mechanism, researchers believe the flower itself is the main sensory organ involved in the response. As mentioned, plants do produce mechanoreceptor proteins, which can sense physical vibrations. The presence of these proteins within the petals likely plays a role in sensing bee vibrations. Moreover, the bowl-shape of the flower itself may be under some selective pressures that favor the ability of the flower to sense its pollinators. More work is needed to better understand exactly how the signal pathways play out. Also, the question remains as to how wide spread this phenomenon is and how it differs between different plants and floral shapes.

Photo Credits: [1] [2]

Further Reading: [1]

Ants have struck up a lot of interesting and important relationships with plants. They disperse seeds, protect plants from herbivores and disease, and can even help acquire nutrients. For all of the beneficial ways in which ants and plants interact, pollination rarely enters into the equation. More often than not, ants are actually detrimental to the sex lives of flowering plants. Such is not the case for a rare species of protea endemic to Western Australia called the smokebush (Conospermum undulatum).

The reason ants usually suck at pollination is thanks to a tiny organ called the metapleural gland. For many ant species, this gland secretes special antimicrobial fluids that the ants use to groom themselves. Because ants tend to live in high densities in close quarters, this antimicrobial fluid helps keep their little bodies clean of any pathogens that might threaten their existence. For as good as these fluids are for ants, they destroy pollen grains, rendering them useless for pollination.

As is so often the case in nature, there are always exceptions to the rule and it seems that one such exception is playing out in Western Australia. While investigating the reproductive ecology of the smokebush, researchers noted that ants were regular visitors to their small flowers. They knew that in drier climates, some ant species have evolved to produce considerably less antimicrobial fluids. The thought is that drier climates tend to harbor fewer microbial pathogens and thus ants don’t need to waste as much energy protecting themselves from such threats. If this was the case in Western Australia then it was entirely possible that ants could potentially serve as pollinators for this plant. Armed with this hypothesis, they decided to take a closer look.

It turns out that the floral morphology of the smokebush lends well to visiting ant anatomy. The tiny flowers produce a small amount of nectar at the base. As ants shove their heads down into the flower to get a drink, it triggers an explosive mechanism that causes the style the smack down onto the back of the ant. In doing so, it also mops up any pollen the ant may be carrying. At the same time, the anthers explosively dehisce, coating the visitor with a fresh dusting of pollen. During their observations, researchers noted that ants weren’t the only insects visiting smokebush blooms. They also noted lots of visitation from invasive honeybees (Apis mellifera) and a tiny native bee called Leioproctus conospermi.

After recording visits, researchers needed to know whether any of these floral visitors resulted in successful pollination. After all, just because something visits a flower doesn’t mean it has what it takes to get the job done for the plant. By looking at differences in seed set between ant and bee visitors, they were able to paint a fascinating picture of the pollination ecology of the rare smokebush.

It turns out that ants are indeed excellent pollinators of this shrub, contributing just as much to overall seed set as the tiny native Leioproctus conospermi. Alternatively, invasive honeybees barely functioned as pollinators at all. Their heads were too big to effectively trigger the pollination mechanism of the flowers but nonetheless were able to access the nectar within. As such, honeybees are considered nectar thieves for the smokebush, harming its overall reproductive effort rather than helping.

Amazingly, the effectiveness of ants as smokebush pollinators is not because they produce less antimicrobial fluids. In fact, these ants were fully capable of producing ample amounts of these pollen-killing substances. Instead, it appears that the plant itself has evolved to tolerate ant visitors. Smokebush pollen is resistant to the toxic effects of the metaplural gland fluids. With plenty of hungry ants always on the lookout for food, the smokebush has managed to tap in to an abundant and reliable vector for pollination. No doubt other examples exist, we simply have to go looking.

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

Further Reading: [1]

For me, an obsession with everything botanical came later on in my academic career. I never paid too much attention to plants as a kid. To be brutally honest, I used to find plants boring. I was too busy preoccupying myself with reptiles, amphibians, and fish. However, if there was ever a plant that was an icon of my care-free childhood existence, it would have to be the humble yet hardy pineappleweed, Matricaria discoidea.

Tearing around on playgrounds for most of the summer months, this little member of the aster family was one of the few species that could handle the endless energy of hundreds of rampaging children and thus was one of the only plants I ever paid much attention to. Still, is wasn’t until much later that I took the time to figure out its identity and natural history.

Pineappleweed is native to parts of northeast Asia and northwestern North America. There are some out there who believe this species may have been brought to North America by paleolithic peoples as a food plant. While this remains to be substantiated, there is no doubt that this is one adaptable species. Now nearly global in its distribution, pineappleweed thrives in some of the harshest habitats imaginable for such a small plant. Its tough stem can handle a lot of foot traffic, making it a common sight along roadsides, city walkways, and of course, playgrounds.

Though at first glance it doesn’t look like it, pineappleweed is a member of the daisy family (Asteraceae). It simply lacks the showy ray florets produced by those of its close cousins. Speaking of cousins, pineappleweed is actually a close relative of chamomile (Matricaria chamomilla). What looks like a single yellow flower is actually a disk made up of many individual flowers densely packed into a dome. The blooms are attractive to tiny syphrid flies but it is not quite known if they are effective pollinators or not. Pineappleweed is also an annual and each disk of flowers can produce thousands of sticky little seeds. This is how this species gets around. Its seeds stick to everything from animal fur to shoes and even car tires. Pineappleweed is yet another species that has benefited from the wonton globalization that humans have enacted upon the world. Keep your eye out for it. It isn’t hard to find and it is certainly a plant worthy of closer inspection.

Further Reading: [1] [2]

The heartleaf twayblade is truly a sight for sore eyes.... that is, if you can find it. This diminutive orchid stands no more than 30 cm tall when in bloom and, for much of its life, exists as a single pair of tiny, heart-shaped leaves. Finding this species in bloom has been one of the major highlights of the last few years of botanizing. Getting to see it up close makes me wonder how many times I may have passed it over completely.

A closeup examination of the flowers will reveal what looks like tiny little humanoids. Indeed, the flowers are complex little structures. Tiny trigger hairs located at the base of the pollinia squirt glue on the back of visiting insects, which affixes the pollen sacs or pollinia. One to two days after the pollinia have been removed the stigmas become receptive to pollen. Though this orchid can self fertilize, differential ripening of sexual parts like this helps ensure cross pollination between different individuals.

With flowers so small, it is a wonder that insects can even find them. As it turns out, the flowers emit a foul smelling odor, though one would be hard pressed to detect it having to bend down so close to the forest floor. This attracts a wide variety of small insects like wasps and flies. The most common visitors, however, are fungus gnats. Ever abundant in the moist duff of the forest, these tiny dipterids offer plenty of opportunity for pollination. The orchid even sweetens the deal a bit by producing a small amount of nectar.

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

Deer populations in North America are higher than they have been at any point in history. Their explosion in numbers not only leads to series health issues like starvation and chronic wasting disease, it has also had serious impacts on regional plant diversity. Wherever hungry herds of deer go, plants disappear from the landscape. However, the impacts of deer on plants aren’t limited to species they can eat. Research on Jack-in-the-Pulpit (Arisaema triphyllum) has shown that deer can have plenty of surprising indirect impacts on plants as well.

Though I wouldn’t put anything past a hungry deer, plants like Jack-in-the-Pulpit aren’t usually on the menu for these ungulates. Their leaves, stems, and flowers are chock full of raphide crystals that will burn the mouths and esophagus of most herbivores. Still, this doesn’t mean deer aren’t impacting these plants in other ways. Because deer are congregating in high abundance in our ever-shrinking natural spaces, they are having serious impacts on local growing conditions. Wherever deer herds are at high numbers, forests are experiencing soil compaction, soil erosion, and a disappearance of soil leaf litter (also due in part to invasive earthworms). Thanks to issues like these, plants like Jack-in-the-Pulpit are undergoing some serious changes.

Like many aroids, sex expression in the genus Arisaema is fluid and relies on energy stores. Smaller plants store less energy and tend to only produce male flowers when they bloom. Pollen, after all, is cheap compared to eggs and fruit. Only when a plant has stored enough energy over the years will it begin to produce female flowers in addition to males and only the largest, most robust plants will switch over entirely to female flowers. As you can imagine, the ability of a plant to acquire and store enough energy is dependent on the quality of the habitat in which it grows. This is where deer enter into the equation.

High densities of deer inevitably cause serious declines in habitat quality of plants like Jack-in-the-Pulpit. As leaf litter disappears and soil compaction grows more severe, individual plants have a much harder time storing enough energy each growing season. In places where deer impacts are heaviest, the sex ratios of Jack-in-the-Pulpit populations begin to skew heavily towards males because individual plants must grow much longer before they can store enough energy to produce female flowers. It doesn’t end their either. Not only does it take longer for a plant to begin producing female flowers, individual plants must also reach a much larger size in order to produce female flowers than in areas with fewer deer.

As mentioned, seed production takes a lot of energy and any plant that is able to produce viable fruits will have less energy stores going into the next season. This means that even if a plant is able to produce female flowers and successfully set seed, they will have burned through so much energy that they will likely revert right back to producing only male flowers the following year, further skewing the sex ratios of any given population towards males. Interestingly, this often results in more individuals being produced via clonal offshoots. The more clones there are in a population, the less diverse the gene pool of that population becomes.

Without actually eating the plants, deer are having serious impacts on Jack-in-the-Pulpit population dynamics. I am certain that this species isn’t alone either. At least Jack-in-the-Pulpit is somewhat flexible in its reproductive behaviors. Other plants aren’t so lucky. I realize deer are a hot button issue but there is no getting around the fact that our mismanagement of their natural predators, habitat, and numbers are having serious and detrimental impacts on wild spaces and all the species they support.

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

Further Reading: [1]

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Spring can be very unpredictable. If bees emerge from their slumber too early or too late, they can miss the flowering period of the plants they rely on for food. By the same token, the plants themselves then miss out on important pollination services. Mismatches like this are becoming more common as climate change continues to accelerate. However, not all bees are helpless if they emerge onto a landscape devoid of flowers. It turns out that, with a little nibble, some bees are able to coax certain plants into flowering.

In a series of recent experiments, scientists were able to demonstrate that at least three species of bumblebee (Bombus terrestris, B. lapidarius, and B. lucorum) were able to induce early flowering in tomatoes (Solanum lycopersicum) and mustards (Brassica nigra) simply by nibbling on their leaves. The queens would land on the leaf and make a series of small holes with their mandibles before flying off. The bees did not appear to be feeding on any of the sap, nor were they carrying chunks of leaf when they flew off. Amazingly, the act of nibbling on the leaves in each experiment resulted in earlier flowering times across both species of plant.

The results were not minor either. Flowers on bee-nibbled plants were produced an average of 30 days earlier than non-nibbled plants. Amazingly, when scientists tried to simulate bee nibbles using tweezers and knives, they were only able to coax flowering an average of 8 days earlier than non-damaged plants. What this means is that there is something about the bite of a bee that sends a signal to the plant to start flowering. Perhaps there’s a chemical cue in the bee’s saliva. Some trees have shown to respond to the detection of deer saliva, ramping up defense compounds in their leaves only once they have detected deer. More work is needed before we can say for sure.

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Considering the role bees play in pollination of plants like tomatoes and mustards, it is likely that this interaction benefits both players to some degree; bees are able to coax floral resources much sooner than they would normally become available while the plants are flowering when effective pollinators are present in the area. These exciting results open yet another window into the multitude of ways in which plants and their pollinators interact. Given that plants have been known to skew the caste systems in eusocial bees, it should come as no surprise to learn that some bees have a few tricks up their sleeves as well.

Photo Credits: [1] [2]

Further Reading: [1]

There are few things on a hike that get me pumped more than hearing someone call out "Hey, I found something weird over here!" It's even more exciting when that person knows what they are talking about. Sometimes that "something" is a familiar species in a strange spot, or growing in a strange way. Sometimes, however, it is something new and exciting that you have been wanting to encounter for years.

This is how I finally met the American climbing fern (Lygodium palmatum). Tangled among the branches of a shrub was indeed a strange site. The tiny, palmate pinnules are not a dead giveaway as to its true identity. Regardless of looks, this is in fact a fern. It is the only member of this genus native to North America. Its cousins, the Japanese climbing fern (Lygodium japonicum) and the Old World climbing fern (Lygodium microphyllum) can also be found on this continent but they have become very invasive in the southeast.

I know what some of you may be thinking, "if this is a fern then where are the fronds?" This was my first thought as well. My first guess was aimed at each palmate leaf. Wrong. The correct answer is the whole vine! Each climbing vine of this fern is a single frond. The palmate leaves are actually the pinnules. The stem, or rachis as it is called in ferns, twines around branches and stems in a vine-like fashion, unfurling pinnules as it goes. What is most impressive is that these fronds can grow as long as 15 feet. Quite impressive by North American fern standards. Fertile pinnules form at the ends of these fronds. Their lacy appearance is quite beautiful juxtaposed with the hand-like, sterile pinnules.

The American climbing fern can be found growing throughout eastern North America. It is a fern of wet places, enjoying acidic soils and bright sunlight. Unfortunately its preference for wetlands has landed it on threatened and endangered lists throughout its range. Our nasty habit of draining, farming, and developing wetlands means that the American climbing fern (as well as many of the other species it shares its habitat with) is losing habitat at an alarming rate.

Further Reading:
http://plants.usda.gov/core/profile?symbol=LYPA3

Star chickweed (Stellaria pubera) has been called North America’s showiest chickweed and I am inclined to agree. Come mid-spring, this lovely woodland plant produces wonderful white flowers that measure about 1/2 inch across and are ringed by five petals so deeply notched that there appear to be ten. Star chickweed’s floral display takes place rather close to the ground on small, fuzzy shoots but as the flowering window for this species begins to close, a change takes place within the plant. By mid-summer, star chickweed will have grown into something completely different.

As mentioned, flowering for star chickweed occurs close to the ground. During this time, its stems don’t elongate more than a few inches and its leaves are broad, blunt, and sessile. Once seed has been set, star chickweed goes through another growth spurt. New stems begin to grow that are much more vigorous in nature than the flowering shoots. They sprout up from the base of the plant and completely over-top spring growth. They can reach heights of nearly 12 inches and produce much thinner leaves. These summer shoots are usually sterile and only in rare instances have flowers been reported.

Star chickweed’s shape-shifting abilities have confused many a botanizer over the last century or so. Because the fertile and sterile shoots look completely different from each other and largely occur at different times of the growing season, some early botanists even went as far as to describe them as different species. Why this plant goes through two distinct growth phases is still something of a mystery but I suspect it has a lot to do with energy reserves.

Perhaps star chickweed has evolved this shape-shifting habit to keep up with changes in surrounding vegetation. Early in the year, the tree canopy above hasn’t completely closed and many of its herbaceous neighbors are still putting on growth of their own. As such, star chickweed probably doesn’t experience as much competition for light early in the season. Of course, conditions on the forest floor change drastically as spring gives way to summer. It could be that the taller, more vigorous sterile shoots are better able to compete for light as the forest fills in around star chickweed.

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Photo Credits: [1] [2]

Further Reading: [1] [2]

Meet 香港线路梯子 aka the Maltese mushroom. Despite the common name and overall appearance, this is not a fungus. It is indeed a plant. Cynomorium coccineum is a holoparasite. It produces no chlorophyl of its own and relies solely on a host plant for all of its water and nutrient needs. It is said to parasitize the roots of halophytes or salt-loving plants and thus, is most commonly found growing in salt marshes in addition to dry, sandy habitats in coastal areas.

Native to the Mediterranian region and extending into parts of Afghanistan, Saudi Arabia, Iran, and Central Asia, this species is really only ever found during the rainy season. Most of its life is spent underground, emerging only to display its flowers. Only when enough energy has been garnished from the host will this plant throw up these strange flower spikes. As you can tell from the picture, the spikes are jam packed with highly reduced flowers. The flowers give off a scent that has been likened to cabbage. It is thought that flies take up the bulk of the pollination of these blooms.

As you can probably guess by its strange appearance, the taxonomic affinity of this strange parasite has been the subject of much debate. For a long time, many botanists placed it in the family vp免费节点香港 but more recent genetic work suggests it belongs in its own family, Cynomoriaceae. It is the only genus within that family but interestingly enough, Cynomoriaceae is located within the order Saxifragales, somewhere near Crassulaceae, making it a distant relative of stonecrops like sedum. No matter where its located on the tree of life, Cynomorium coccineum is surely one of the strangest plants on Earth.

Photo Credits: [1] [2]

Further Reading: [1] [2]

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Throughout southeastern North America, gators change their behaviors with the seasons. During the rainy season, alligators can be found floating in open water or sunning themselves on land. Except when hunting, they don’t seem to do anything with much urgency. Their activity level changes during the dry season when water is in short supply. Gators don’t sit back and let nature take its course. They spring into action and create their own aquatic refuges.

As the surrounding landscape begins to dry, gators will excavate holes or pits in the soggy ground called gator holes. These holes hold onto water when most of the surrounding landscape isn’t. The process of digging a gator hole may seem destructive but it all must be placed in the context of the surrounding environment. Most gator habitat exists in low lying areas. In places like the Everglades, there isn’t much topography to speak of. When a gator excavates a gator hole, it creates variation in both hydrology and soil conditions.

Soils that have built up over time are lifted out of the hole and piled into mounds. Mounded soils are not only rich in nutrients, they also dry at different rates, creating a gradient in water availability. Plants that normally can’t germinate and grow in saturated soils find suitable spots to live up on the soil mounds while emergent aquatic vegetation fills in along the parameter. Plants that normally prefer to grow in deeper water can also establish within the gator hole itself. In the midst of fairly uniform marsh vegetation, a gator hole quickly becomes a hotbed of plant diversity. The differences in vegetation can be so stark compared to the surrounding landscape that some scientists can actually map gator holes using aerial scans simply by measuring the differences in infrared radiation given off by the leaves of all the different plants that establish around them.

Of course, all of that plant diversity has a huge effect on other organisms as well. Gator holes become important areas for various reptiles, amphibians, birds, and so much more. The paths that alligators take to and from their holes even act like fire breaks, changing the way fire moves through the landscape, which only increases the heterogeneity of the immediate area. Fish, though occasionally eaten, greatly benefit from the stability of water levels within a gator hole. All in all, gator holes are extremely important habitats. Not only do they support a high diversity of plants and animals alike, they make places like the Everglades even more dynamic than they already are.

Photo Credits: [1] [2]

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

Behold the Calypso orchid, Calypso bulbosa. This circumboreal orchid exists as a single leaf lying among the litter of dense conifer forests. They go virtually unnoticed for most of the year until it comes time to flower.

In early spring, the extravagant blooms open up and await the arrival of bumblebees. Calypsos go to great lengths to attract bumblebees. The flower is said to have a sweet scent. Also, the lip sports small, yellow, hair-like protrusions that are believed to mimic anthers covered in pollen. Finally, within the pouch formed by the lip are two false nectar spurs. All of these are a ruse. The Calypso offers no actual rewards to visiting bumblebees.

Not just any bumblebee will do. For the ruse to work, it requires freshly emerged workers that are naive to the orchid’s deception. Bumblebees are not mindless animals. They quickly learn which flowers are worth visiting and which are not. Because of this, the Calypso has only short window of time in which bumblebees in the vicinity are likely to fall for its tricks. As a result, pollination rates are often very low for this orchid.

The most interesting aspect of all of this is that the so-called "male function" of the flower - pollinia removal - is more likely to occur than the "female function" - pollen deposition. The reason for this makes a lot of sense in context; male function requires a bumblebee to be fooled only once whereas female function requires a bumblebee to be fooled at least twice.

The caveat to all of this deception is that a single Calypso, like all other orchids, can produce tens of thousands of seeds. Each orchid therefore has tens of thousands of potential propagules to replace itself in the next generation. Despite that fact, the Calypso orchid is on the decline. Habitat destruction, poaching, deer, and invasive species are taking their toll. If you care about orchids like the Calypso, please consider supporting organizations like the North American Orchid Conservation Center.

Photo Credit: [1] [2]

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

With delicately dissected foliage and flowers that look like pantaloons, it is hard to believe that Dutchman's breeches (Dicentra cucullaria) are related to the common garden poppy. No matter how incredulous it may seem, they are in fact peculiar members of Papaveraceae. I can't get enough of these lovely spring ephemerals and their beauty is equally matched by their intriguing ecology. This species really is the full package.

At home in mesic deciduous forests, Dutchman's breeches are true spring ephemerals. They are primarily denizens of eastern North America, however, disjunct populations can be found in the Pacific Northwest. These are likely relics of a once wider distribution that was split in two by advancing glaciers during the Pleistocene. Dutchman’s breeches live out their entire lives before the tree canopy closes with a fresh batch of leaves. By mid summer they are little more than dormant bulblets resting below the leaf litter. Like the multitude of spring ephemerals they share the forest with, Dutchman's breeches are vying for pollinators capable of tolerating wide swings in temperature. This is where their peculiar little flowers come in.

Packed away in each spur is a sweet nectary treat. The only insects capable of reaching it are bumblebees (Bombus spp.). With their long tongues, these bees flock to the bright white and yellow flowers with vigor. Aside from the occasional thief who chews a hole at the end of the spur, robust bumblebees have this meal all to themselves. In fact, this relationship is so in sync that nothing else is capable of effectively pollinating the plant.

After a brief flowering period, the plant will set seed. Like many other spring ephemerals, they attach a fleshy structure to their seeds called an elaiosome. This attracts foraging ants in the genus Aphaenogaster, who collect the seeds and take them back to their nests. Once there, the elaiosome is sometimes eaten but mostly the seeds are disposed of in trash middens. In this way, the seeds find a nutrient-rich microclimate safe from seed predators in which to germinate. It is a safe bet that most of the patches you find owe their existence to these industrious little insects.

Further Reading: [1] [2]

I like record breaking species. It is always exciting to find out which species produces the largest or smallest of something. Lately (and rightfully so), the titan arum (Amorphophallus titanum) has been getting a lot of attention for its incredible inflorescence. Many have bloomed in botanical gardens over the last few years and each one draws a massive crowd. People flock from far and wide to see that largest unbranched inflorescence in the world. You always see it referred to that way; the largest unbranched inflorescence. That got me to thinking, who produces the largest branched inflorescence in the world?

The answer to this is the talipot palm (Corypha umbraculifera). Native to southern India and Sri Lanka, the talipot palm blows all other branched inflorescences out of the water. Heck, branched or not, looking over its dimensions makes me feel like it puts most floral structures to shame. The branched designation comes from the fact that its flowers aren’t borne on a single stalk but many branching stalks. The proportions of this structure are truly staggering.

The talipot palm inflorescence can measure upwards of 26 feet (8 m) in length and bear as many as 23.9 million flowers at a time. It has been estimated that if you were to lay out all of the branches and flower stalks end to end, you would have nearly 26,000 feet (8,221 m) of plant material. This is truly epic as far as flowering plants are concerned. Even more amazing is the fact that this epic inflorescence is often produced 65 feet (20 m) up in the air!

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During my search, I also came across another interesting record breaking palm, 台湾梯子. This species is native to parts of western Africa where it can be found growing in moist, lowland forests. Raphia regalis has the distinct honor of producing the largest self-supporting leaf in the world. Given what I have read, I would imagine that in a dense forest, it would be extremely difficult to take in the full grandeur of its leaves. They are huge. The current record for a single R. regalis leaf is 82 feet (25.1 m) long. It isn’t a solid leaf but rather a compound leaf made up of much tinnier leaflets. To see one in all of its glory would be a truly special event.

So there you have it. Two incredible plant records held by two incredible palms. Not bad for a quick internet search.

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

Further Reading: [1]

Plants use pigments for a variety of functions. The most obvious examples of these are chlorophyll and anthocyanins. Whereas we can see such pigments, our eyes are not equipped to see others. Many plant species utilize pigments that can only be seen by organisms capable of seeing in the ultraviolet spectrum. The most famous examples of this involve flowers, which utilize UV pigments to attract pollinators like bees and some birds, which can see into the UV portion of the electromagnetic radiation spectrum. However, our understanding of UV pigments grows every year and it is becoming obvious that many organisms use UV pigments for other reasons. For instance, the carnivorous ends of some carnivorous plants have been found to fluoresce blue. These plants aren’t using these pigments for pollination, rather they are using them to hunt.

Pitcher plants are unmistakable. These beautiful carnivores produce complex pit-fall traps from highly modified leaves. Their traps are an evolutionary adaptation to the low-nutrient conditions of the soils in which pitcher plants grow. The traps entice potential prey to visit in a few different ways including bright colors, sweet smells, and even nectar. The rims of the pitchers are slippery and visting insects have trouble hanging on. Organisms that are lured in by these "rewards" run the risk of slipping and falling to their deaths into a pool of digestive fluids.

According to a paper published back in 2013, these are not the only traits that pitcher plants use to attract prey. Along the rim of many pitcher plant traps, researchers discovered special pigment cells that fluoresce blue light in the ultra violet wavelengths. Fluorescent pigments were even found in the pitcher fluids of some Nepenthes! In fact, close examination revealed that the pitcher fluid does not start to fluoresce until the pitcher lid has opened. The researchers wanted to test to see if this UV light actually functioned as an attractant for insects. What they found was exactly that.

When the blue emissions were masked, the traps caught considerably fewer insects. It would appear that many carnivorous pitcher plants are tapping into a strong evolutionary connection between insects and flowers. It is an interesting adaptation for catching prey in nutrient-poor environments. Moreover, it is amazing to see just how striking these pitchers look under UV light. The pitcher plants aren’t alone, either. Venus fly traps (Dionaea muscipula) also exhibited UV light emissions around their trapping devices. Of course, since we lack the ability to see in the UV spectrum without technology, all of this is going on out of sight of us humans. It just goes to show you how truly complex and interesting plants can be.

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

Further Reading: [1]

We all have our biases and one of my biggest botanical bias is that I often think of plants from eastern North America before my mind heads further west. I can’t really fault myself for it because so many of my early plant experiences occurred east of the Mississippi. I want to remedy this a bit today by drawing your attention to a wonderful aroid who frequently gets overshadowed by its eastern cousin.

I am of course talking about western skunk cabbage (手机怎么挂香港ip). This incredibly beautiful plant enjoys a distribution that ranges from southern Alaska to central California and west into Wyoming and Montana. Like its eastern cousin, western skunk cabbage was awarded its common name thanks to the pungent odor it produces. Its blooming period ranges from March into May depending on where they are growing and the inflorescence is truly something to write home about.

Emerging from the base of the plant is a bright yellow structure called a spathe. The spathe envelopes the actual flowering parts, a phallic-looking structure covered in flowers called a spadix. The spadix emits various volatile compounds that function as pollinator attractants. However, whereas many would suggest flies are the preferred pollinator, research indicates that a tiny species of rove beetle called Pelecomalium testaceum takes up the bulk of pollination duties for western skunk cabbage throughout much of its range.

The volatile compounds aren’t there to trick the beetles into thinking they are getting some sort of reward. The plant does actually reward the rove beetles with pollen to eat and relatively safe place to mate. We call these types of signals “honest signals” as they act as an honest calling card that signifies rewards are to be had.

Unfortunately, the beauty of western skunk cabbage has seen it enter into novelty garden collections in other temperate regions of the world. In northern Europe, western skunk cabbage has escaped the confines of the garden and is now considered an invasive species in wetlands of that region. Take care to choose you garden plants wisely. Always plant native plants when the option presents itself.

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

Further Reading: [1] [2]

Take a look at a list of tree species from temperate Europe, North America, and Asia and you will notice a glaring disparity. Whereas North America and Asia are home to something like 1000 tree species each, the European continent is home to just about 500 species. Why is this?

The answer may lie partly in the glacial history of the Northern Hemisphere as well as in some quirks of geology. Starting in the late Pliocene, roughly 3 million years ago, the Earth began to cool. As our planet entered into a epoch dominated by massive, continent-wide glaciers, life was responding accordingly.

Historically it was assumed that Europe lost many of its temperate tree species thanks to the east-west orientation of its mountain ranges. As glaciers advanced from the north, species were pushed farther and farther south until they hit physical barriers in the terrain like the Alps. With nowhere to go but up, many species that couldn’t handle either the rate of climate change or the altitude adjustment simply winked out of existence. Fossil evidence from Europe provides plenty of evidence that Europe was once home to far more tree species, including relatives of sweetgum (vp免费节点香港 spp.) and tulip trees (Liriodendron spp.) that are still present in North America, and umbrella pines (Sciadopitys spp.), which still exists in Asia. Many temperate tree species in North America and Asia were spared this fate because there were far fewer barriers to successful southern migrations.

This all sounds a bit too simple and indeed, recent studies suggest that it is. Though climate change, glaciers, and mountains certainly played a role in the differential extinction rates of European trees, the story is a bit more complicated than that. It turns out that the European mountain ranges don’t present as impenetrable of a barrier to plant migrations as was once thought. The fact that southern Europe and northern Africa share many similar taxa is proof of this. Instead, the amount of suitable habitat and land area available to trees migrating down from northern Europe may have played an even larger role in the extinction rate of European trees.

It is a well documented phenomenon in ecology that smaller areas of land support smaller numbers of species. This is the case for Pleistocene Europe. Suitable habitat for temperate tree species during this time would have largely consisted of three peninsulas (Iberia, Italy, and the Balkans) separated by the Mediterranian Sea. Each of these peninsulas boast mountain chains that would have offered small bands of suitable microclimates for temperate tree species to find refuge during glacial advance.

Pushed into tiny pockets of refugia, Europe’s temperate tree species would have been more vulnerable to extinction than tree species in North America and Asia, which had far more suitable habitat available to them in the southern portions of those continents. By looking at which taxa survived and which went extinct, patterns do start to emerge. Tree species that are widespread in Europe today are descendants of trees that were far more tolerant of cooler growing seasons and harsh winters than genera that went extinct. This likely reflects the fact that their ancestors were those species that found refuge high up in the mountains.

Alternatively, present-day Europe also boasts small pockets of what are termed “relictual genera,” which is a fancy way of saying species that were once more common in the past than they are today. These so-called relictual taxa have been found to be far more tolerant of drought than genera that went extinct. This likely reflects the fact that their ancestors found refuge in warmer, low-elevation habitats in southern Europe.

It appears that species on either end of the tolerance curves were the ones that won out in Europe’s extinction lottery. By tolerating either extreme cold or extreme drought, “stress tolerators” were able to not only survive repeated glaciation events, but also provide seed sources for those lineages following glacial retreat.

Only the species that were able to find suitable habitats in southern Europe’s glacial refugia were the ones that were able to recolonize the continent after the ice age had ended. At this point in time, these are some of the best pieces of evidence we have in explaining the disparity in tree diversity between Europe, North America, and Asia. What’s more, I find disturbing trends in such extinctions because it wasn’t like the glaciers always wiped out species immediately. Instead, many species were able to survive glaciation but were pushed into smaller and smaller pockets of suitable habitat until relatively small disturbances pushed them over the edge.

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Photo Credits: [1] [2]

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