Are Packrats Fumigating Their Homes Using Plants?

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Any organism that lives in one place for a long enough time is going to have to deal with pests. For mammals, this often means fleas and ticks. Nests, dens, and other roosting spots tend to accumulate high numbers of these blood suckers the longer they are in use. As such, anything that can cut down on pest loads in and around the home has the potential to confer great advantages. Evidence from California suggests that wood rats may be using the leaves of a shrub to do just that.

Dusky-footed wood rats (A.K.A. packrats) build giant nests out of twigs and other plant debris. These nests serve to protect packrats from both the elements and hungry predators. Packrat nests can last for quite a long time and reach monumental proportions considering the size of the rat itself. Because they use these stick nests for long periods of time, it should come as no surprise that they can build up quite a pest load. Fleas are especially problematic for these rodents.

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 A dusky-footed woodrat ( Neotoma fuscipes ) and its den.

A dusky-footed woodrat (Neotoma fuscipes) and its den.

When researchers took a closer look at what packrats were bringing into their nests, they realized that not all plant material was treated equally. Whereas packrats actively collect and feed on leaves from various oaks (Quercus spp.), conifers (Pinus spp., Juniperus spp., etc.), and toyon (Heteromeles arbutifolia), the packrats seemed to have a special affinity for the leaves of the California bay (Umbellularia californica). However, instead of taking huge bites out of bay leaves, the rats appear to nibble them along the margin and spread them throughout their nest. What’s more, fresh bay leaves are brought in every few days.

This led some researchers to suggest that, instead of packrats using bay leaves as food, they may be using them to fumigate their homes. Indeed, California bay is rather chemically active. It is an aromatic shrub noted for its resistance to insect infestation. Of special interest to the research team were a group of chemical compounds called monoterpenoids. They noted that bay leaves were especially high in two types of of these compounds - 1,8-cineole (which gives the shrub its characteristic odor), and umbellulone (which has shown to be quite toxic to rodents). Why else would packrats bring something potentially deadly into their home other than to drive off pests?

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Closer observation revealed that the packrats were in fact treating bay leaves differently than other leaves. For starters, bay leaves were disproportionately used to line the sleeping chambers within the stick nests. What’s more, the bay leaves were cycled out every 2 to 3 days. Even the nibbling patterns were significantly different. As mentioned above, bay leaves were merely nibbled along the leaf margins, which is an ideal place to nibble if releasing volatile compounds is the desired effect.

When researchers tested the effectiveness of a variety of leaves in the lab, their results added further evidence to the fumigation hypothesis. More than any other leaf found in packrat nests, bay leaves had clear negative effects on flea numbers. Flea survival in the lab was reduced by upwards of 75% when California bay leaves were present whereas flea survival was only reduced by less than 10% with all other leaves. It goes without saying that, whether they are conscious decisions or not, packrats definitely stand to benefit by decorating their homes with California bay leaves.

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

Further Reading: [1]

Arctic Vegetation is Growing Taller & Why That Matters

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The Arctic ecosystem is changing and it is doing so at an alarming rate. Indeed, the Arctic Circle is warming faster than most other ecosystems on this planet. All of this change has implications for the plant communities that call this region home. In a landmark study that incorporated thousands of data points from places like Alaska, Canada, Iceland, Scandinavia, and Russia, researchers have demonstrated that Arctic vegetation is, on average, getting taller.

Imagine what it is like to be a plant growing in the Arctic. Extreme winds, low temperatures, a short growing season, and plenty of snow are just some of the hardships that characterize life on the tundra. Such harsh conditions have shaped the plants of this region into what we know and love today. Arctic plants tend to hug the ground, hunkering down behind whatever nook or cranny offers the most respite from their surroundings. As such, plants of Arctic-type habitats tend to be pretty small in stature. As you can probably imagine, if these limits to plant growth become less severe, plants will respond accordingly.

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That is part of what makes this new paper so alarming. The vegetation that comprise these Arctic communities is nearly twice as tall today as it was 30 years ago. However, the increase in height is not because the plants that currently grow there are getting taller but rather because new plants are moving northwards into these Arctic regions. New players in the system are usually cause for concern. Other studies have shown that it isn’t warming necessarily that hurts Arctic and alpine plants but rather competition. They simply cannot compete as well with more aggressive plant species from lower latitudes.

Taller plants moving into the Arctic may have even larger consequences than just changes in species interactions. It can also change ecosystem processes, however, this is much harder to predict. One possible consequence of taller plants invading the Arctic involves carbon storage. It is possible that as conditions continue to favor taller and more woody vegetation, there could actually be more carbon storage in this system. Woody tissues tend to sequester more carbon and shading from taller vegetation may slow decomposition rates of debris in and around the soil.

  Alopecurus alpinus  is one of the new tall plant species moving into the Arctic

Alopecurus alpinus is one of the new tall plant species moving into the Arctic

It is also possible that taller vegetation will alter snowpack, which is vital to the health and function of life in the Arctic. Taller plants with more leaf area could result in a reduced albedo in the surrounding area. Lowering the albedo means increased soil temperatures and reduced snowpack as a result. Alternatively, taller plants could also increase the amount of snowpack thanks to snow piling up among branches and leaves. This could very well lead (counterintuitively) to warmer soils and higher decomposition rates as snowpack acts like an insulating blanket, keeping the soil slightly above freezing throughout most of the winter.

It is difficult to make predictions on how a system is going to respond to massive changes in the average conditions. However, studies looking at how vegetation communities are responding to changes in their environment offer us one of the best windows we have into how ecosystems might change moving into the uncertain future we are creating for ourselves.

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

Further Reading: [1]

The Upside Down World of Orchid Flowers

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Did you know that most orchid flowers you see are actually blooming upside down? That's right, referred to as "resupination," the lower lip of many orchid flowers is actually the top petal and, as the flower develops inside the bud, the whole structure makes a 180° rotation. How and why does this happen?

The lip of an orchid flower usually serves to attract pollinators as well as function as a landing pad for them. The flower of an orchid is an incredibly complex organ with an intriguing evolutionary history. Basically, the lip is the most derived structure on the flower and, in most cases, it is the most important structure in initiating pollination.

 The non-resupinate flowers of the grass pink ( Calopogon tuberosus ) showing the lip on top.

The non-resupinate flowers of the grass pink (Calopogon tuberosus) showing the lip on top.

As an orchid flower bud develops, it begins to exhibit gravitropic tendencies, meaning it responds to the pull of gravity. By removing specific floral organs like the column and pollinia, researchers found that they produce special hormones called auxins that tell the developing bud to begin the process of resupination. The ovary starts to twist, causing the flower to stand on its head.

Not all orchids exhibit resupinate flowers. Grass pinks (Calopogon tuberosus) famously bloom with the lip pointing up as it does in the early stages of bud development. It is an interesting mechanism and serves to demonstrate the stepwise tendencies that the forces of natural selection and evolution can manifest. But why does it occur at all? What is the evolutionary advantage of resupinate flowers?

 Not only are  Dracula  flowers resupinate, many species also face them towards the ground.

Not only are Dracula flowers resupinate, many species also face them towards the ground.

The most likely answer to this biological twist is that, for orchids, resupination places the lip in such a way that facilitates pollination by whatever the flowers are attracting. For many orchids, this means providing an elaborate landing strip in the form of the lip. For the grass pinks, which operate by slamming visiting bees downward onto the column to achieve pollination, placing the lip at the top makes more mechanical sense. When a bee visits the upward pointing lip thinking it will find a pollen-rich meal, the lip bend at the base like a hinge. Anything goes in evolution provided the genes are present for selection to act upon and nowhere is this fact more beautifully illustrated than in orchids.

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

Viper's Bugloss

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Throughout much of North America, brown fields, roadsides, and other waste places occasionally take on a wonderful blue hue. Often time the cause of this colorful display is none other than Echium vulgare, or as its commonly referred to, viper's bugloss. Viper’s bugloss is a member of the borage family and was originally native to most of Europe and Asia. However, humans introduced it to North America some time ago. It has since naturalized quite well and is even considered invasive in parts of Washington. No matter your views on this plant, the reproductive ecology of this species is quite interesting.

Viper's bugloss produces its flowers on spikes. Starting off pink and gradually changing to blue as they mature, the flowers ripen their male portions on their first day and ripen their female portions on the second day. This is known as "protandry." Plants that exhibit this lifestyle offer researchers a window into the advantages and disadvantages with regards to the fitness investment of each sex. What they have found in viper's bugloss is that there are clearly distinct strategies for each type of flower.

Male flowers are pollinator limited. They must hedge their bets towards increasing the number of visitors. Bees are the main pollinators of this species and the more bees that visit, the more pollen can be disseminated. Unlike female flowers, which are resource limited, male flowers can produce pollen and nectar quite cheaply. Because of this, male flowers produce significantly more nectar than female flowers to bring in more bees. As the anthers senesce and give way to receptive styles, things begin to change. The plant now has to redirect resources into producing seed. At this point, resources are everything. The plant produces considerably less nectar resources than pollen but the bees can’t know that without visiting.

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The other interesting aspect its reproductive ecology has to do with population size. Bees are notorious for favoring plants that are more numerous on the landscape. This makes a lot of sense. Why spend time looking for uncommon plants when they can go for easier, more numerous targets. This can be very detrimental to the fitness of rare plant species. However, plants like viper's bugloss don't seem to fall victim to this.

By looking at large and small populations, researchers found that pollination success pretty much evens out for viper's bugloss no matter how numerous it is in a given area. Large populations receive many more visits from bees but the bees spend less time on each flower. When viper's bugloss populations are small, flowers receive fewer visits but bees spend more time at each flower. This results is no significant difference in the reproductive fitness of either population.

Considering how efficient this plant is reproductively, it is no wonder it has done so well outside of its native range. Add to this its ability to grow in some of the worst soil conditions, it goes without saying that viper's bugloss is here to stay. If you find this species growing, certainly take time to get up close with the flowers. You will be glad you did.

Photo Credits: [1] [2]

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


The Genus Ceropegia Recently Got a Whole Lot Bigger

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The succulent and climbing members of the milkweed family (Apocynaceae) have been gaining a lot of popularity among houseplant growers and for good reason. These wonderful plants produce some of the most elaborate flowers most of us will ever encounter and many of them smell quite strongly. Whereas houseplant enthusiasts recognize multiple genera of these spectacular plants, recent taxonomic work suggests lumping them all into one single genus - Ceropegia.

Such a massive taxonomic move has caused its fair share of drama. Folks seem to get quite ornery when it comes to shifts in nomenclature, especially when it involves this many species. However, when you dive into this group of plants, you really start to see how shaky the ground was that supported the previous classification systems. Evolution, after all, is not a neat and tidy process and we can learn a lot from the succulent and climbing asclepiads regarding the importance of combining morphological and genetic data into our taxonomic decisions.

  Ceropegia (Stapelia) hirsuta

Ceropegia (Stapelia) hirsuta

Botanists have been obsessing over this group for decades. Historically speaking, four major groups have been recognized: those with caudiciform stems (genus Brachystelma), the stem succulent stapeliads (which include genera such as Stapelia, Huernia, Orbea, Caralluma, and others), the climbers (genus Ceropegia), and the so-called early divergent group (which include genera such as Anisotoma, Conomitra, Dittoceras, and others). Together these groups total something like 762 species and represent the tribe Ceropegieae.

The taxonomic status of the various members of Ceropegieae have always been up for debate. Early work was based on surprisingly few species and relied heavily on morphological characters such and corolla shape, stem anatomy, and pubescence. Since the 1950’s, many more species have been discovered and that is where a lot of the trouble began. Much of the early characters that were used to draw lines between various groups were suddenly blurred. Genera were created and absorbed by various authors in an attempt to get a handle on how this tribe evolved.

  Ceropegia  ( Brachystelma )  tuberosum

Ceropegia (Brachystelma) tuberosum

Things got even more complicated as various stapeliads and Ceropegia attracted the attention of horticulturists. As new species became available, many varieties were haphazardly named and genera such as Stapelia were further split to accommodate some of the peculiar nuances in floral shapes, colors, and sizes. It wasn’t until some genetic work was done that the need for a major overhaul of the Ceropegieae tribe became apparent.

Unfortunately, this early molecular work suffered from low resolution. Very few genera were used and among those, only a handful of gene regions were analyzed. Still, the picture that was developing was that the historical understanding of Ceropegieae was surprisingly misleading. For instance, the genera that made up the stapeliad group appeared to be nested quite firmly within the genus Ceropegia. Though equally as limited in scope, consecutive work in the early 2000’s added further evidence to the idea that the four groups that made up Ceropegieae were so genetically similar that most should be nested somewhere within Ceropegia.

  Ceropegia  ( Duvalia )  modesta

Ceropegia (Duvalia) modesta

Though not without controversy, this early molecular work convinced enough taxonomists to take a closer look at each of the four groups. With more resolution and a finer grasp on the diversity in form of these plants, taxonomists started to question the validity of some taxa. Indeed, the closer anyone looked, the more the lines between genera started to blur.

For example, Ceropegia and Brachystelma have long been separated on the basis of floral structure. Ceropegia were considered to adhere to a single corolla structure involving long, tubular flowers whereas Brachystelma were thought to be more variable in form. The discovery of new species clearly demonstrates that there are far too many exceptions to this system for it to be valid.

 Fig. 1. Variation in the corolla and corona in the traditional concept of  Ceropegia : A–C,  C. salicifolia , Nepal,  Bruyns 2507  (BM, K); D–E,  C. melanops , Ethiopia,  Gilbert 3050  (K); F—H,  C. meleagris , Nepal,  Bruyns 2496  (K); I–J,  C. loranthiflora , Ethiopia,  Gilbert 2851   (K). [scale-bars or subdivisions indicate mm; A, D, F, I, corolla from  side; B, G, corolla dissected to show location of corona; C, E, H, J,  corona from side].   [SOURCE]

Fig. 1. Variation in the corolla and corona in the traditional concept of Ceropegia: A–C, C. salicifolia, Nepal, Bruyns 2507 (BM, K); D–E, C. melanops, Ethiopia, Gilbert 3050 (K); F—H, C. meleagris, Nepal, Bruyns 2496 (K); I–J, C. loranthiflora, Ethiopia, Gilbert 2851 (K). [scale-bars or subdivisions indicate mm; A, D, F, I, corolla from side; B, G, corolla dissected to show location of corona; C, E, H, J, corona from side]. [SOURCE]

 Fig. 2. Variation in the corolla and corona in the traditional concept of  Brachystelma : A–C,  B. brevipedicellatum , South Africa,  Bruyns 2372 ; D–F,  B. mafekingense , Namibia,  Bruyns 1954  (K, WIND); G–J,  B. gymnopodum , South Africa,  Bruyns 2078   (NBG). [scale-bars or subdivisions indicate mm; A, corolla from front,  D, G, corolla from side; B, E, H, corolla dissected to show location of  corona; C, J, corona from front; F, I, corona from side].   [SOURCE]

Fig. 2. Variation in the corolla and corona in the traditional concept of Brachystelma: A–C, B. brevipedicellatum, South Africa, Bruyns 2372; D–F, B. mafekingense, Namibia, Bruyns 1954 (K, WIND); G–J, B. gymnopodum, South Africa, Bruyns 2078 (NBG). [scale-bars or subdivisions indicate mm; A, corolla from front, D, G, corolla from side; B, E, H, corolla dissected to show location of corona; C, J, corona from front; F, I, corona from side]. [SOURCE]

Such is also the case for other anatomical features such as whether plants climb or not. Again, there are plants in both genera that deviate from these patterns, thus making it impossible to nail down any set of characters that maintain the split between these two genera. Also, it would seem that some authors were trying to pull a fast one on readers. Back in 2007, Meve and Liede-Schumann claimed there were “a wide array of morphological features” that separate these two genera but failed to reveal any but those mentioned here. There are multiple species of Ceropegia and Brachystelma that simply do not conform to this historical classification.

Similarly, Ceropegia and the various stapeliads have been separated on the basis of stem and floral anatomy. Historically speaking, the stapeliads were thought to consist of fleshy, succulent stems with tubercules and reduced or absent leaves, whereas Ceropegia were considered to be slender climbers. Again, with more species having been discovered, these distinctions grew more and more blurry.

 The succulent stems of  Ceropegia cimiciodora .

The succulent stems of Ceropegia cimiciodora.

It turns out that there are many Ceropegia with fleshy, succulent stems and the only major difference between the two genera is the lack of angles in the stems of some Ceropegia. The structure and presentation of their flowers also stands on shaky ground. There is so much similarity between the flowers of some of the succulent Ceropegia and the early diverging stapeliads that one would be hard pressed to identify any character that clearly separates them.

Between all of the molecular work and the anatomical scrutiny, it was clear that something needed to be done to clean up the taxonomic status of Ceropegieae. Keeping things separate may make sense to some but considering the group as a whole instead of from a collector’s standpoint, trying to find enough distinct characters to preserve the historical treatment would make things way too messy. In 2017 it was suggested that because there are no clear differences between the four groups within this tribe, all members were to be lumped back in to the genus Ceropegia.

  Ceropegia  ( Stapelia )  flavopurpurea

Ceropegia (Stapelia) flavopurpurea

Although this most recent treatment still recognizes some morphological differences between these plants (thus multiple subsections are recognized), the lack of genetic differentiation between groups long thought to be distinct really does support this decision. Because of historical precedents, Ceropegia won out as the main generic classification.

Personally I find this work to be extremely exciting. It involved a lot of wonderful detective work and a whole lot of attention to detail. I think the end result paints a far better picture for our understanding of how these plants evolved. I am especially floored that some of the earlier morphological notes turned out to be quite useful in this modern understanding. Even more exciting is the fact that now we know that many of what we thought were “unique” characters amoung the various species actually evolved multiple times throughout the history of this group. This is why I will never get upset by taxonomic changes. They may be working documents but each step we take helps us understand evolution that much more.

Photo Credits: [1] [2] [3] [4] [5] [6] [7]

Further Reading: [1]

The Wild World of Rattan Palms

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

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

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

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

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

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

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

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

The Only True Cedars

  Cedrus deodara

Cedrus deodara

The only true cedars on this planet can be found growing throughout mountainous regions of the western Himalayas and Mediterranean. All others are cedars by name only. The so-called “cedars” we encounter here in North America are not even in the same family as the true cedars. Instead, they belong to the Cypress family (Cupressaceae). The true cedars all belong to the genus Cedrus and are members of the family Pinaceae. They are remarkable trees with tons of ecological and cultural value.

 J. White,1803-1824.

J. White,1803-1824.

The true cedars are stunning trees to say the least. They regularly reach heights of 100 ft. (30 m.) or more and can live for thousands of years. Cedars exhibit a dimorphic branching structure, with long shoots forming branches and smaller shoots carrying most of the needle load. The needles themselves are borne in dense, spiral clusters, giving the branches a rather aesthetic appearance. Each needle produces layers of wax, which vary in thickness depending on how exposed the tree is growing. This waxy layer helps protect the tree from sunburn and desiccation.

  Cedrus libani

Cedrus libani

  Cedrus libani

Cedrus libani

One of the easiest ways to identify a cedar is by checking out its cones. All members of the genus Cedrus produce upright, barrel-shaped cones. Male cones are smaller and don’t stay on the tree for very long. Female cones, on the other hand, are quite large and stay on the tree until the seeds are ripe. Upon ripening, the entire female cone disintegrates, releasing the seeds held within. Each seed comes complete with blisters full of distasteful resin, which is thought to deter seed predators.

 Male cones of  Cedrus atlantica

Male cones of Cedrus atlantica

 Female  Cedrus  cones.

Female Cedrus cones.

In total, there are only four recognized species of cedar - the Atlas cedar (Cedrus atlantica), the Cyprus cedar (C. brevifolia), the deodar cedar (C. deodara), and the Lebanon cedar (C. libani). I have heard arguments that C. brevifolia is no more than a variant of C. libani but I have yet to come across any source that can say this for certain. Much more work is needed to assess the genetic structure of these species. Even their place within Pinaceae is up for debate. Historically it seems that Cedrus has been allied with the firs (genus Abies), however, work done in the early 2000’s suggests that Cedrus may actually be sister to all other Pinaceae. We need more data before anything can be said with certainty.

  Cedrus atlantica

Cedrus atlantica

Regardless, two of these cedars - C. atlantica & C. libani - are threatened with extinction. Centuries of over-harvesting, over-grazing, and unsustainable fire regimes have taken their toll on wild populations. Much of what remains is not considered old growth. Gone is the heyday of giant cedar forests. Luckily, many populations are now located in protected areas and reforestation efforts are being put into place throughout their range. Still, the ever present threat of climate change is causing massive pest outbreaks in these forests. The future for these trees hangs in the balance.

Photo Credit: Wikimedia Commons

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

Meet Pokeweed's Tree-Like Cousin

There is more than one way to build a tree. For that reason and more, “tree” is not a taxonomic designation. Arborescence has evolved independently throughout the botanical world and many herbaceous plants have tree-like relatives. I was shocked to learn recently of a plant native to the Pampa region of South America commonly referred to as ombú. At first glance it looks like some sort of fig, with its smooth bark and sinuous, buttressed roots. Deeper investigation revealed that this was not a fig. Ombú is actually an arborescent cousin of pokeweed!

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The scientific name of ombú is Phytolacca dioica. As its specific epithet suggests, plants are dioecious meaning individuals are either male or female. Unlike its smaller, herbaceous cousins, ombú is an evergreen perennial. Because they can grow all year, these plants can reach bewildering proportions. Heights upwards of 60 ft. (18 m.) are not unheard of and the crowns of more robust specimens can easily attain diameters of 40 to 50 ft. (12 - 18 m.)! What makes such sizes all the more impressive is the way in which ombú is able to achieve such growth.

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Ombú is thought to have evolved from an herbaceous ancestor. Cut into the trunk of one of these trees and you will see that this phylogenetic history has left its mark. Ombú do not produce what we think of as wood. Instead, much of the support for branches and stems comes from turgor pressure. Also, the way in which these trees grow is not akin to what you would see from something like an oak or a maple. Whereas woody trees undergo secondary growth in which the cambium layer differentiates into xylem and phloem, thus thickening stems and roots, ombú exhibits a unique form of stem and root thickening called “anomalous secondary thickening.”

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Essentially what this means is that instead of a single layer of cambium forming xylem and phloem, ombú cambium exhibits unidirectional thickening of the cambium layer. There are a lot of nitty gritty details to this kind of growth and I must admit I don’t have a firm grasp on the mechanics of it all. The point of the matter is that anomalous secondary thickening does not produce wood as we know it and instead leads to rapid growth of weak and spongy tissues. This is why turgor pressure is so important to the structural integrity of these trees. It has been estimated that the trunk and branches of an ombú is 80% water.

 A cross section of an ombú limb showing harder xylem tissues separated by spongy parenchyma that has since disintegrated.

A cross section of an ombú limb showing harder xylem tissues separated by spongy parenchyma that has since disintegrated.

Like all members of this genus, ombú is plenty toxic. Despite this, ombú appears to have been embraced and is widely planted as a specimen tree in parks, along sidewalks, and in gardens in South America and elswhere. In fact, it is so widely planted in Africa that some consider it to be a growing invasive issue. All in all I was shocked to learn of this species. It caused me to rethink some of the assumptions I hold onto with some lineages I only know from temperate regions. It is amazing what natural selection has done to this genus and I am excited to explore more arborescent anomalies from largely herbaceous groups.

Photo Credits: [1] [2]

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

Rodents as Pollinators

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It may come as a surprise to some that small mammals such as rodents, shrews, and even marsupials have been coopted by plants for pollination services. Far from being occasional evolutionary oddities, many plants have coopted small furry critters for their reproductive needs. Some of the best illustrations of this phenomenon occur in the Protea family (Proteaceae).

  Protea nana

Protea nana

The various members of Proteaceae are probably best known for their bizarre floral displays. Indeed, they are most often encountered outside of their native habitats as outlandish additions to the cut flower industry. Superficial interest in beauty aside, the floral structure of the various protea genera and species is complex to say the least. They are well adapted to ensure cross pollination regardless of what the inflorescence attracts. Most notable is the fact that pollen doesn’t stay on the anthers. Instead, it is deposited on the tip of a highly modified style, which is referred to as the pollen presenter. Usually these structures remain closed until some visiting animal triggers their release.

 The inconspicuous floral display of  Protea cordata .

The inconspicuous floral display of Protea cordata.

Although birds and insects have taken up a majority of the pollination needs of this family, small mammals have entered into the equation on multiple occasions. Pollination by rodents, shrews, and marsupials is collectively referred to as therophilly and it appears to be quite a successful strategy at that. Therophilous pollination has arisen in more than one genera within Proteaceae.

  Leucospermum arenarium  in the field and one of its pollinators,  Gerbillurus paeba,  feeding on flowers. (A) Pollen presenter contact on  G. paeba . (B)  G. paeba  foraging on  L. arenarium   [Source]

Leucospermum arenarium in the field and one of its pollinators, Gerbillurus paeba, feeding on flowers. (A) Pollen presenter contact on G. paeba. (B) G. paeba foraging on L. arenarium [Source]

A therophilous pollination syndrome appears to come complete with a host of unique morphological characters aimed at keeping valuable pollen and nectar away from birds and insects. The inflorescences of therophilous species like Protea nana, P. cordata, and Leucospermum arenarium are usually tucked deep inside the branches of these bushes, often at or near ground level. They are also quite robust and sturdy in nature, which is thought to be an adaptation to avoid damage incurred by the teeth of hungry mammals. The inflorescences of therophilous proteas also tend to have brightly colored or even shiny flowers surrounded by inconspicuous brown involucral bracts.

 (C) Flowering  L. arenarium  with dense, mat-forming inflorescences. (D) Geoflorous inflorescences. (E) Pendulous inflorescences above ground level.  [Source]

(C) Flowering L. arenarium with dense, mat-forming inflorescences. (D) Geoflorous inflorescences. (E) Pendulous inflorescences above ground level. [Source]

Contrasted against bird pollinated proteas, these inflorescences can seem rather drab but that is because small mammals like rodents and shrews are drawn in by another sense - smell. Therophilous proteas tend to produce inflorescences with strong musty or yeasty odors. They also produce copious amounts of sugar-rich, syrupy nectar. Small mammals, after all, need to take in a lot of calories throughout their waking hours and it appears that proteas use that to their advantage.

 A small mouse pollinating  Protea nana

A small mouse pollinating Protea nana

As a rodent or shrew slinks in to take a drink, its head gets completely covered in pollen. In fact, they become so dusted with pollen that, before small, easy to hide trail cameras became affordable, pollen loads in the feces of rodents were the main clue that these plants were attracting something other than birds or insects. What’s more, the flowering period of many of these therophilous proteas occurs in the spring, right around the time when many small mammals go into breeding mode. Its during this time that small mammals need all of the energy they can get.

  Protea humiflora  being pollinated by two different species of rodent in South Africa.

Protea humiflora being pollinated by two different species of rodent in South Africa.

As odd as it may seem, rodent pollination appears to be a successful strategy for a considerable amount of protea species. The proteas aren’t alone either. Other plants appear to have evolved therophilous pollination as well. Nature, after all, works with what it has available and small mammals like rodents make up a considerable portion of regional faunas. With that in mind, it is no wonder that more plants have not converged on a similar strategy. Likely many have, we just need to take the time to sit down and observe.

Photo Credits: [1] [2] [3] [4] [5] [6] [7]

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



Glacier Mice

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At first glance the surface of a glacier hardly seems hospitable. Cold, barren, and windswept, glaciers appear to be the antithesis of life. However, this assumption is completely completely false. Glaciers are home to an interesting ecosystem of their own, albeit on a smaller scale than we normally give attention to.

From pockets of water on the surface to literal lakes of water sealed away inside, glaciers are home to a myriad microbial life. On some glaciers the life even gets a bit larger. Glaciers are littered with debris. As dust and gravel accumulate on the surface of the ice, they begin to warm ever so slightly more than the frozen water around them. Because of this, they are readily colonized by mosses such as those in the genus Racomitrium.

The biggest challenge to moss colonizers is the fact that glaciers are constantly moving, which anymore today means shrinking. As such, these bits of debris, along with the mosses growing on them, do not sit still as they would in say a forest setting. Instead they roll around. As the moss grows it spreads across the surface of the rock while the ice rotates it around. This causes the moss to grow on top of itself, inevitably forming a ball-like structure affectionately referred to as a "glacier mouse."

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Because the moss stays ever so slightly warmer than its immediate surroundings, glacier mice soon find themselves teaming with life. Everything from worms to springtails and even a few water bears call glacier mice home. In a study recently published in Polar Biology, researcher Dr. Steve Coulson found "73 springtails, 200 tardigrades and 1,000 nematodes" thriving in just a single mouse!

The presence of such a diverse community living in these little moss balls brings up an important question - how do these animals find themselves in the glacier mice in the first place? After all, life just outside of the mouse is quite brutal. As it turns out, the answer to this can be chalked up to how the mice form in the first place. As they blow and roll around the the surface of the glacier, they will often bump into one another and even collect in nooks and crannies together. It is believed that as this happens, the organisms living within migrate from mouse to mouse. The picture being painted here is that far from being a sterile environment, glaciers are proving to be yet another habitat where life prospers.

Photo Credit: [1] [2]

Further Reading: [1]

Raphides: A Gnarly Form of Plant Defense

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Take a bite out of a dumbcane (Dieffenbachia spp.) or a pothos (Philodendron spp.) and it won’t be long before your mouth and throat start to burn (please don’t actually do that). Eat enough of it and your symptoms may also include intense numbing, oral irritation, excessive drooling, localized swelling, and possibly even kidney and liver failure (again, please don’t). What you are experiencing is a brutal form of plant defense caused by tiny crystals called raphides.

Raphides are tiny, needle-shaped crystals made up of calcium oxalate. A lot of plants accumulate calcium oxalate. Research has shown that in doing so, plants are able to sequester excess calcium in their cells. Many plant lineages then use that calcium oxalate to make raphides. Not all raphides come in the form of needle-like crystals. Often they are ‘H’ shaped or even twinned. Others are blunt, kind of like tiny crystalline cigars.

 Cigar-shaped raphides found in the tissues of the polka dot plant ( Hypoestes phyllostachya ).

Cigar-shaped raphides found in the tissues of the polka dot plant (Hypoestes phyllostachya).

How raphides form within the plant is rather fascinating. As far as we can discern, raphide crystals form in vacuoles of specialized cells called “idioblasts.” It is thought that an exquisitely controlled scaffolding or matrix shapes the biomineralization process. To the best of my knowledge, no one has been able to reproduce this process in a laboratory setting. For now, plants are the undeniable masters of raphide manufacturing.

Within the cells, raphides are often associated with acrid and toxic proteins. Together, they comprise one hell of a defense against herbivory. Raphides are only the first part of the defensive equation. When plant tissues containing raphides are damaged, usually by chewing, the raphides shoot out of the idioblasts and into the oral cavity of the herbivore. This is where their needle shape comes in.

 Needle-like raphides extracted from the leaves of an  Epipremnum  species.

Needle-like raphides extracted from the leaves of an Epipremnum species.

Raphides wreak havoc on sensitive tissues. They literally act like tiny needles, cutting into and tearing the lining of the mouth, esophagus, and gut. This is only half of the story though. As mentioned, raphides are often packed in with acrid and toxic proteins. The laceration caused by the raphides allows these compounds to enter into the wounds. This is where things can get especially nasty. If the proteins are toxic enough, the herbivore now has far more to worry about than simply the burning sensation.

Raphides are not produced in equal amounts in all tissues. Stems tend to have more than leaves, but raphide content in leaves has also shown to be a function of leaf size. Raphides also differ from species to species. Not all plants that produce raphides produce them in the same shape and quantity. Still, more than 200 plant families contain species that have evolved this form of defense and many of our most prized houseplants fall into this category. However, this should not scare you away from these plants. Provided you or your loved ones don’t go nibbling on the leaves or stems, all will be fine. If anything, this remarkable form of plant defense should earn these plants even more respect than they already get.

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

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

The Strangest Maple

I love being humbled by plant ID. Confusion usually means I am going to end up learning something new. This happened quite recently during a trip through The Morton Arboretum. Admittedly trees are not my forte but I had spotted something that seemed off and needed further inspection. I was greeted by a small tree with leaves that screamed "birch family" (Betulaceae) yet they were opposite. Members of the birch family should have alternately arranged leaves. What the heck was I looking at?

It didn't take long for me to find the ID tag. As a plant obsessed person, the information on the tag gave me quite the thrill. What I was looking at was possible the strangest maple on the planet. This, my friends, was my first introduction to Acer carpinifolium a.k.a the hornbeam maple.

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The hornbeam maple is endemic to Japan where it can be found growing in mountainous woodlands and alongside streams. Maxing out around 30 feet (9 m) in height, the hornbeam maple is by no means a large tree. It would appear that it has a similar place in its native ecology as other smaller understory maples do here in North America. It blooms in spring and its fruits are the typical samaras one comes to expect from the genus.

It probably goes without saying that the thing I find most fascinating about the hornbeam maple are its leaves. As both its common and scientific names tell you, they more closely resemble that of a hornbeam (Carpinus spp.) than a maple. Unlike the lobed, palmately veined leaves of its cousins, the hornbeam maple produces simple, unlobed leaves with pinnate venation and serrated margins. They challenge everything I have come to expect out of a maple. Indeed, the hornbeam maple is one of only a handful of species in the genus Acer that break the mold for leaf shape. However, compared to the rest, I think its safe to say that the hornbeam maple is the most aberrant of them all. 

Not a lot of phylogenetic work has been done on the relationship between the hornbeam maple and the rest of its Acer cousins. At least one study suggests that it is most closely related to the mountain maple of neartheastern North America. More scrutiny will be needed before anyone can make this claim with certainty. Still, from an anecdotal standpoint, it seems like a reasonable leap to make considering just how shallow the lobes are on mountain maple leaves.

Regardless of who it is related to, running into this tree was truly a thrilling experience. I love it when species challenge long held expectations of large groups of plants. Hornbeam maple has gone from a place of complete mystery to me to being one of my favorite maples of all time. I hope you too will get a chance to meet this species if you haven't already!

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

Further Reading: [1] [2]

 

Getting to Know Sansevieria

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The houseplant hobby is experiencing something of a renaissance as of late. With their popularity on various social media platforms, easy to grow plant species and their cultivars are experiencing a level of popularity they haven't seen in decades. One genus of particular interest to houseplant hobbyists is Sansevieria.

Despite their popularity, the few Sansevieria species regularly found in cultivation come attached with less than appealing common names. Mother-in-law's tongue, Devil's tongue, and snake plant all carry with them an air of negativity for what are essentially some of the most forgiving houseplants on the market. What few houseplant growers realize is that those dense clumps of upright striped leaves tucked into a dark corner of their home belong to a fascinating genus worthy of our admiration. What follows is a brief introduction to these enigmatic houseplants.

  Sansevieria cylindrica

Sansevieria cylindrica

  Sansevieria ballyi

Sansevieria ballyi

The Sansevieria we encounter in most nurseries are just the tip of the iceberg. Sansevieria is a genus comprise of about 70 different species. I say 'about' because this group is a taxonomic mess. There are a couple reasons for this. For starters, the vast majority of Sansevieria species are painfully slow growers. It can take decades for an individual to reach maturity. As such, they have never really presented nursery owners with much in the way of economic gain and thus only a few have received any commercial attention.

Another reason has to do with the fiber market during and after World War II. In hopes of discovering new plant-based fibers for rope and netting, the USDA collected many Sansevieria but never formally described most of them. Instead, plants were assigned numbers in hopes that future botanists would take the time needed to parse them out properly.

A third reason has to do with the variety of forms and colors these plants can take. Horticulturists have been fond of giving plants their own special cultivar names. This complicates matters as it is hard to say which names apply to which species. Often the same species can have different names depending on who popularized it and when.

  Sansevieria grandis in situ .

Sansevieria grandis in situ.

Regardless of what we call them, all Sansevieria hail from arid regions of Africa, Madagascar and southern Asia. In the wild, many species resemble agave or yucca and, indeed, they occupy similar niches to these New World groups. Like so many other plants of arid regions, Sansevieria evolved CAM photosynthesis as a means of coping with heat and drought. Instead of opening up their stomata during the day when high temperatures would cause them to lose precious water, they open them at night and store CO2 in the form of an organic acid. When the sun rises the next day, the plants close up their stomata and utilize the acid-stored carbon for their photosynthetic needs.

 The wonderfully compact  Sansevieria pinguicula .

The wonderfully compact Sansevieria pinguicula.

Often you will encounter clumps of Sansevieria growing under the dappled shade of a larger tree or shrub. Some even make it into forest habitats. Most if not all species are long lived plants, living multiple decades under the right conditions. These are just some of the reasons that they make such hardy houseplants.

The various Sansevieria appear the sort themselves out along a handful of different growth forms. The most familiar to your average houseplant enthusiast is the form typified by Sansevieria trifasciata. These plants produce long, narrow, sword shaped leaves that point directly towards the sky. Many other Sansevieria species, such as S. subspicata and S. ballyi, take on a more rosetted form with leaves that span the gamut from thin to extremely succulent. Still others, like S. grandis and S. forskaalii, produce much larger, flattened leaves that grow in a form reminiscent of a leaky vase. 

  Sansevieria trifasciata  with berries .

Sansevieria trifasciata with berries.

Regardless of their growth form, a majority of Sansevieria species undergo radical transformations as they age. Because of this, adults and juveniles can look markedly different from one another, a fact that I suspect lends to some of the taxonomic confusion mentioned earlier. A species that illustrates this nicely is S. fischeri. When young, S. fischeri consists of tight rosettes of thick, mottled leaves. For years these plants continue to grow like this, reaching surprisingly large sizes. Then the plants hit maturity. At that point, the plant switches from its rosette form to producing single leaves that protrude straight out of the ground and can reach heights of several feet! Because the rosettes eventually rot away, there is often no sign of the plants previous form.

 A young  Sansevieria fischeri  exhibiting its rosette form.

A young Sansevieria fischeri exhibiting its rosette form.

 A mature  Sansevieria fischeri  with its large, upright, cylindrical leaves.

A mature Sansevieria fischeri with its large, upright, cylindrical leaves.

If patient, many of the Sansevieria will reach enormous sizes. Such growth is rarely observed as slow growth rates and poor housing conditions hamper their performance. It's probably okay too, considering the fact that, when fully grown, such specimens would be extremely difficult to manage in a home. If you are lucky, however, your plants may flower. And flower they do!

Though there is variation among the various species, Sansevieria all form flowers on either a simple or branched raceme. Flowers range in color from greenish white to nearly brown and all produce a copious amount of nectar. I have even noticed sickeningly sweet odors emanating from the flowers of some captive specimens. After pollination, flowers give way to brightly colored berries, hinting at their place in the family Asparagaceae.

 A flowering  Sansevieria hallii .

A flowering Sansevieria hallii.

As a whole, Sansevieria can be seen as exceptional tolerators, eking out an existence wherever the right microclimate presents itself in an otherwise harsh landscape. Their extreme water efficiency, tolerance of shade, and long lived habit has lent to the global popularity of only a few species. For the majority of the 70 or so species in this genus, their painfully slow growth rates means that they have never made quite a splash in the horticulture trade.

Nonetheless, Sansevieria is one genus that even the non-botanically minded among us can pick out of a lineup. Their popularity as houseplants may wax and wane but plants like S. trifasciata are here to stay. My hope is that all of these folks collecting houseplants right now will want to learn more about the plants they bring into their homes. They are more than just fancy decorations, they are living things, each with their own story to tell. 

Photo Credits: [1] [2] [3] [4] [5] [6] [7] [8] [9]

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

Maples, Epiphytes, and a Canopy Full of Goodies

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The forests of the Pacific Northwest are known for the grandeur. This region is home to one of the greatest temperate rainforests in the world. A hiker is both dwarfed and enveloped by greenery as soon as they hit the trail. One aspect of these forests that is readily apparent are the carpets of epiphytes that drape limbs and branches all the way up into the canopy. Their arboreal lifestyle is made possible by a combination of mild winters and plenty of precipitation. 

Weare frequently taught that the relationship between trees and their epiphytes are commensal - the epiphytes get a place to live and the trees are no worse for wear. However, there are a handful of trees native to the Pacific Northwest that are changing the way we think about the relationship between these organisms in temperate rainforests.

Though conifers dominate the Pacific Northwest landscape, plenty of broad leaved tree species abound. One of the most easily recognizable is the bigleaf maple (Acer macrophyllum). Both its common and scientific names hint at its most distinguishing feature, its large leaves. Another striking feature of this tree are its epiphyte communities. Indeed, along with the vine maple (A. circinatum), these two tree species carry the greatest epiphyte to shoot biomass ratio in the entire forest. Numerous species of moss, liverworts, lichens, and ferns have been found growing on the bark and branches of these two species.

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Epiphyte loads are pretty intense. One study found that the average epiphyte crop of a bigleaf maple weighs around 78 lbs. (35.5 Kg). That is a lot of biomass living in the canopy! The trees seem just fine despite all of that extra weight. In fact, the relationship between bigleaf and vine maples and their epiphyte communities run far deeper than commensalism. Evidence accumulated over the last few decades has revealed that these maples are benefiting greatly from their epiphytic adornments.

Rainforests, both tropical and temperate, generally grow on poor soils. Lots of rain and plenty of biodiversity means that soils are quickly leached of valuable nutrients. Any boost a plant can get from its environment will have serious benefits for growth and survival. This is where the epiphytes come in. The richly textured mix of epiphytic plants greatly increase the surface area of any branch they live on. And all of that added surface area equates to more nooks and crannies for water and dust to get caught and accumulate.

When researchers investigated just how much of a nutrient load gets incorporated into these epiphyte communities, the results painted quite an impressive picture. On a single bigleaf maple, epiphyte leaf biomass was 4 times that of the host tree despite comprising less than 2% of the tree's above ground weight. All of that biomass equates to a massive canopy nutrient pool rich in nitrogen, phosphorus, potassium, calcium, magnesium, and sodium. Much of these nutrients arrive in the form of dust-sized soil particles blowing around on the breeze. What's more, epiphytes act like sponges, soaking up and holding onto precious water well into the dry summer months.

Now its reasonable to think that nutrients and water tied up in epiphyte biomass would be unavailable to trees. Indeed, for many species, epiphytes may slow the rate at which nutrients fall to and enter into the soil. However, trees like bigleaf and vine maples appear to be tapping into these nutrient and water-rich epiphyte mats.

 A subcanopy of vine maple ( Acer circinatum ) draped in epiphytes.

A subcanopy of vine maple (Acer circinatum) draped in epiphytes.

Both bigleaf and vine maples (as well as a handful of other tree species) are capable of producing canopy roots. Wherever the epiphyte load is thick enough, bundles of cells just under the bark awaken and begin growing roots. This is a common phenomenon in the tropics, however, the canopy roots of these temperate trees differ in that they are indistinguishable in form and function from subterranean roots.

Canopy roots significantly increase the amount of foraging an individual tree can do for precious water and nutrients. Additionally, it has been found that canopy roots of the bigleaf maple even go as far as to partner with mycorrhizal fungi, thus unlocking even more potential for nutrient and water gain. In the absence of soil nutrient and water pools, a small handful of trees in the Pacific Northwest have unlocked a massive pool of nutrients located above us in the canopy. Amazingly, it has been estimated that mature bigleaf and vine maples with well developed epiphyte communities may actually gain a substantial fraction of their water and nutrient needs via their canopy roots.

 

Photo Credits: [1] [2]

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

 

The Tecate Cypress: A Tree Left Hanging in the Balance

The tecate cypress is a relict. Its tiny geographic distribution encompasses a handful of sights in southern California and northwestern Mexico. It is a holdover from a time when this region was much cooler and wetter than it is today. It owes its survival and persistence to a combination of toxic soils, a proper microclimate, and fires that burn through every 30 to 40 years. However, things are changing for the Tecate cypress and they are changing fast. The fires that once ushered in new life for isolated populations of this tree are now so intense that they may spell disaster.

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The taxonomy of the Tecate cypress has undergone a few revisions since it was first described. Early work on this species suggested it was simply a variety of Cupressus guadalupensis. Subsequent genetic testing revealed that these two trees were distinct enough to each warrant species status of their own. It was then given the name Cupressus forbesii, which will probably be familiar to most folks who know it well. Work done on the Tecate cypress back in 2012 has seen it moved out of the genus Cupressus and into the genus Hesperocyparis. As far as I am concerned, whether you call it Cupressus forbesii or Hesperocyparis forbesii matters not at this point.

The Tecate cypress is an edaphic endemic meaning it is found growing only on specific soil types in this little corner of the continent. It appears to prefer soils derived from ultramafic rock. The presence of high levels of heavy metals and low levels of important nutrients such and potassium and nitrogen make such soils extremely inhospitable to most plants. As such, the Tecate cypress experiences little competition from its botanical neighbors. It also means that populations of this tree are relatively small and isolated from one another.

The Tecate cypress also relies on fire for reproduction. Its tiny cones are serotinous, meaning they only open and release seeds in response to a specific environmental trigger. In this case, its the heat of a wildfire. Fire frees up the landscape of competition for the tiny Tecate cypress seedlings. After a low intensity fire, literally thousands of Tecate cypress seedlings can germinate. Even if the parent trees burn to a crisp, the next generation is there, ready to take their place.

At least this is how it has happened historically. Much has changed in recent decades and the survival of these isolated Tecate cypress populations hangs in the balance. Fires that once gave life are now taking it. You see, decades of fire suppression have changed that way fire behaves in this system. With so much dry fuel laying around, fires burn at a higher intensity than they have in the past. What's more, fires sweep through much more frequently today than they have in the past thanks to longer and longer droughts.

Taken together, this can spell disaster for small, isolated Tecate cypress populations. Even if thousands of seedlings germinate and begin to grow, the likelihood of another fire sweeping through within a few years is much higher today. Small seedlings are not well suited to cope with such intense wildfires and an entire generation can be killed in a single blaze. This is troubling when you consider the age distributions of most Tecate cypress stands. When you walk into a stand of these trees, you will quickly realize that all are of roughly the same age. This is likely due to the fact that they all germinated at the same time following a previous fire event.

If all reproductive individuals come from the same germination event and wildfires are now killing adults and seedlings alike, then there is serious cause for concern. Additionally, when we lose populations of Tecate cypress, we are losing much more than just the trees. As with any plant, these trees fit into the local ecology no matter how sparse they are on the landscape. At least one species of butterfly, the rare Thorne's hairstreak (Callophrys gryneus thornei), lays its eggs only on the scale-like leaves of the Tecate cypress. Without this tree, their larvae have nothing to feed on.

 Thorne's hairstreak ( Callophrys gryneus thornei ), lays its eggs only on the scale-like leaves of the Tecate cypress.

Thorne's hairstreak (Callophrys gryneus thornei), lays its eggs only on the scale-like leaves of the Tecate cypress.

Although things in the wild seem uncertain for the Tecate cypress, there is reason for hope. Its lovely appearance and form coupled with its unique ecology has led to the Tecate cypress being something of a horticultural curiosity in the state of California. Seeds are easy enough to germinate provided you can get them out of the cones and the trees seem to do quite well in cultivation provided competition is kept to a minimum. In fact, specimen trees seem to adapt quite nicely to California's cool, humid coastal climate. Though the future of this wonderful endemic is without a doubt uncertain, hope lies in those who care enough to grow and cultivate this species. Better management practices regarding fire and invasive species, seed collection, and a bit more public awareness may be just what this species needs.

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

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

Why Are Some Plants Overcompensating?

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Gardeners are all too familiar with herbivory. Countless times I have been awaiting a bloom to burst only to have the buds nipped off the night before they opened. While this can be devastating for many plant species (not to mention my sanity), for certain plant species, an encounter with a hungry herbivore may actually lead to an increase in reproductive fitness.

Overcompensation theory is the idea that, under certain conditions, plants can respond to herbivore damage by producing more shoots, flowers, and seeds. It goes without saying that when this idea was originally proposed in the late 80's, it was met with its fair share of skepticism. Why would a plant capable of producing more shoots and flowers wait to be damaged to do so? The answer may lie in in the realm of biological trade-offs.

Overcompensation may evolve in lineages that tend to grow in habitats where there is a "predictable" amount of herbivory in any given growing season, perhaps a region where large herbivores migrate through annually. Plants in these habitats may conserve dormant growing tips and valuable resources to be used once herbivory has occurred. Perhaps this also serves as a cue to upregulate antiherbivory compounds in new tissues. The trade-off is that the plants incur a cost in the form of fewer flowers and thus reduced reproduction when herbivory is low or absent.

  Scarlet gilia ( Ipomopsis aggregata )

Scarlet gilia (Ipomopsis aggregata)

It could also be that plants are exhibiting two different strategies - one to deal with competition and one to deal with herbivory. If herbivory is low, plants may become more competitive, thus favoring rapid vertical growth of one or a couple shoots. When herbivory is high, rapid vertical growth becomes disadventageous and overcompensatory branching and flowering can provide the higher fitness benefits.

These possibilities are not mutually exclusive. In fact, since the late 80's, experts now believe that overcompensation is not an "either/or" phenomenon but rather a spectrum of possibilities that are dictated by the conditions in which the plants are growing. Certainly overcompensation exists but which conditions favor it and which do not?

Research on scarlet gilia (Ipomopsis aggregata), a biennial native to western North America, suggests that overcompensation comes into play only when environmental conditions are most favorable. Soil nutrients seem to play a role in how well a plant can bounce back following herbivore damage. When resources are high, the results can be quite astounding. Early work on this species showed that under proper conditions, plants that were browsed by upwards of 95% produced 2.4 times as much seed as uneaten control plants. What's more, the resulting seedlings were twice as likely to survive than their uneaten counterparts.

Things change for scarlet gilia growing in poor conditions. Low resource availability appears to place limits on how much any given plant may respond to browsing. Also, herbivory can really hamper flowering time. Because scarlet gilia is pollen limited, anything that can cause a disruption in pollinator visits can have serious consequences for seed set. In at least one study, browsed plants flowered later and received fewer pollinator visits as a result.

More recent work has been able to add more nuance to the overcompensation story. For instance, experiments done on two subspecies of field gentian (Gentianella campestris), add further support to the idea that overcompensation is a matter of trade-offs. They showed that, whereas competition with neighboring plants alone could not explain the benefits of overcompensation, browsing certainly can.

  Field gentian ( Gentianella campestris )

Field gentian (Gentianella campestris)

Plants growing in environments where herbivory was higher overcompensated by producing more branching, more flowers, and thus more seed, all despite soil nutrients. It appears that herbivory is the strongest predictor of overcompensation for this gentian. What's more, when these data were fed into population models, only the plants that responded to herbivory by overcompensation were predicted to show any sort of population growth in the long term.

Despite all of the interest overcompensation has recieved in the botanical literature, we are only just beginning to understand the biological mechanisms that make it possible. For starters, we know that when a dominant shoot or stem gets damaged or removed, it causes a reduction in the amount of the plant hormone auxin being produced. When auxin is removed, tiny auxiliary buds at the base of the plant are able to break dormancy and begin growing.

Removal of the dominant shoot or stem can also have major impact on the number of chromosomes present in regrowing tissues. Work on Arabidopsis thaliana revealed that when the apical meristem (main growing tip of a vertical stem) was removed, the plant underwent a process called "endoreduplication" in which the cells of the growing tissues actually duplicate their entire genome without undergoing mitosis.

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Endoreduplication is a complex process with lots of biological significance but in plants it is often associated with stress responses. By duplicating the genomes of these new cells, the plants may be able to adjust more rapidly to their environment. This often manifests in changes to leaf size and shape and an uptick in plant defenses. Thus, plants may be able to fine tune the development of new tissues to overcompensate for browsing. Certainly far more work is needed to understand these mechanisms and their functions in more detail.

Overcompensation is not universal. Nonetheless, it is expected to occur in certain plants, especially those with short life cycles, and under certain environmental conditions, mainly when herbivore pressure and nutrient availability are relatively high. That being said, we still have plenty more to learn about this spectrum of strategies. When does it occur and when does it not? How common is it? What are the biological underpinnings of plants capable of overcompensation? Are some lineages more prone to overcompensation than others? Only more research can say for sure!

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

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

 

 

The Plight of the African Violets

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For many of us, African violets (Saintpaulia spp.) are some of the first houseplants we learned how to grow. They are not true violets (Violaceae), of course, but rather members of the family Gesneriaceae. Nonetheless, their compact rosettes of fuzzy leaves coupled with regular sprays of colorful flowers has made them a multi-million dollar staple of the horticultural industry. Unfortunately their numbers in captivity overshadow a bleak future for this genus in the wild. Many African violets are teetering on the brink of extinction.

The genus Saintpaulia is endemic to a small portion of east Africa, with a majority of species being found growing at various elevations throughout the Eastern Arc Mountains of Kenya and Tanzania. Most of the plants we grow at home are clones and hybrids of two species, S. ionantha and S. confusa. Collected in 1892, these two species were originally thought to be the same species, S. ionantha, until a prominent horticulturist noted that there are distinct differences in the seed capsules each produced. Since the 1890's, more species have been discovered.

  Saintpaulia goetzeana

Saintpaulia goetzeana

Exactly how many species comprise this genus is still up for some debate. Numbers range from as many as 20 to as few as 6. Much of the early work on describing various Saintpaulia species involved detailed descriptions of the density and direction of hairs on the leaves. More recent genetic work considers some of these early delineations to be tenuous at best, however, even these modern techniques have resolved surprisingly little when it comes to a species concept within this group.

  Saintpaulia  sp.  in situ .

Saintpaulia sp. in situ.

Though it can be risky to try and make generalizations about an entire genus, there are some commonalities when it comes to the habitats these plants prefer. Saintpaulia grow at a variety of elevations but most can be found growing on rocky outcrops. Most of them prefer growing in the shaded forest understory, hence they do so well in our (often) poorly lit homes. Their affinity for growing on rocks means that many species are most at home growing on rocks and cliffs near streams and waterfalls. The distribution of most Saintpaulia species is quite limited, with most only known from a small region of forest or even a single mountain. Its their limited geographic distribution that is cause for concern.

  Saintpaulia ionantha  subsp.  grotei in situ.

Saintpaulia ionantha subsp. grotei in situ.

Regardless of how many species there are, one fact is certain - many Saintpaulia risk extinction if nothing is done to save them. Again, populations of Saintpaulia species are often extremely isolated. Though more recent surveys have revealed that a handful of lowland species are more widespread than previously thought, mid to highland species are nonetheless quite restricted in their distribution. Habitat loss is the #1 threat facing Saintpaulia. Logging, both legal and illegal, and farming are causing the diverse tropical forests of eastern Africa to shrink more and more each year. As these forests disappear, so do Saintpaulia and all of the other organisms that call them home.

There is hope to be had though. The governments of Kenya and Tanzania have recognized that too much is being lost as their forests disappear. Stronger regulations on logging and farming have been put into place, however, enforcement continues to be an issue. Luckily for some Saintpaulia species, the type localities from which they were described are now located within protected areas. Protection coupled with inaccessibility may be exactly what some of these species need to survive. Also, thanks to the ease in which Saintpaulia are grown, ex situ conservation is proving to be a viable and valuable option for conserving at least some of the genetic legacy of this genus.

  Saintpaulia intermedia

Saintpaulia intermedia

It is so ironic to me that these plants can be so common in our homes and offices and yet so rare in the wild. Despite their popularity, few recognize the plight of this genus. My hope is that, in reading this, many of you will think about what you can do to protect the legacy of plants like these and so many others. Our planet and the species that call it home are doomed without habitat in which to live and reproduce. This is why land conservation is an absolute must. Consider donating to a land conservation organization today. Here are two worth your consideration:

The Nature Conservancy

The Rainforest Trust

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

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

Is Love Vine Parasitizing Wasps?

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No, that's not dodder (Cuscuta sp.), its love vine (Cassytha filiformis), a member of the same family as the avacados in your kitchen (Lauraceae). It is a pantropical parasite that makes its living stealing water and nutrients from other plants. To do so, it punctures their vascular tissues with specialized structures called "haustoria." Amazingly, a recent observation made in Florida suggests that this botanical parasite may also be taking advantage of other parasites, specifically gall wasps.

Gall wasps are also plant parasites. They lay their eggs in developing plant tissues and the larvae release compounds that coax the plant to form nutrient-rich galls packed full of starchy goodness. Essentially you can think of galls as edible nursery chambers for the wasp larvae. While looking for galls on sand live oak (Quercus geminata) growing in southern Florida, Dr. Scott Egan and his colleagues noticed something odd. A small vine seemed to be attaching itself to the galls.

 Love vine draping a host plant. 

Love vine draping a host plant. 

The vine in question was none other than love vine and they were attached to galls growing on the underside of the oak leaves. What is strange is that, of all of the places that love vine likes to attach itself to its host (new stems, buds, petioles, and on the top and edge of leaves), the only time this vine showed any "interest" in the underside of oak leaves was when galls were present. Obviously this required further investigation.

The team discovered that at least two different species of gall wasps were being parasitized by love vine - one that produces small, spherical galls on the underside of oak leaves and one that forms large, multi-chambered galls on oak stems. Upon measuring the infected and uninfected galls, the team discovered that there were significant differences that could have real ecological significance.

 (A)  Cassytha filiformis  vine attaching haustoria to a leaf gall induced by the wasp  Belonocnema treatae  on the underside of their host plant,  Quercus geminata . (B) Labeled graphic of insect gall, parasitic vine, and vine haustoria. (C) Box plots of leaf gall diameter for unparasitized galls (control) and galls that have been parasitized by  C. filiformis . (D) Proportion of  B. treatae  leaf galls that contained a dead ‘mummified’ adult for unparasitized galls (control) and galls that have been parasitized by the vine  C. filiformis .  [SOURCE]

(A) Cassytha filiformis vine attaching haustoria to a leaf gall induced by the wasp Belonocnema treatae on the underside of their host plant, Quercus geminata. (B) Labeled graphic of insect gall, parasitic vine, and vine haustoria. (C) Box plots of leaf gall diameter for unparasitized galls (control) and galls that have been parasitized by C. filiformis. (D) Proportion of B. treatae leaf galls that contained a dead ‘mummified’ adult for unparasitized galls (control) and galls that have been parasitized by the vine C. filiformis. [SOURCE]

For the spherical gall wasp, infected galls tended to be much larger, however, the team feels that this may actually be due to the fact that the vine "prefers" larger galls. Astonishingly, larvae living in the infected galls were 45% less likely to survive. For the multi-chambered gall wasp, infection by love vine was associated with a 13.5% decrease in overall gall size. They suggest this is evidence that love vine is having net negative impacts on these parasitic wasps.

Subsequent investigation revealed that these wasps were not alone. In total, the team found love vine attacking the galls of at least two other wasps and one species of gall-making fly (though no data were reported for these cases). To be sure that love vine was in fact parasitizing these galls, they needed to have a closer look at what the vine was actually doing.

 Figure S2. (A)  Cassytha filiformis  vine attaching haustoria to a leaf gall induced by the wasp  Callirhytis quercusbatatoides  on the stem of their host plant,  Quercus geminata . (B) Labeled graphic of insect gall, parasitic vine, and vine haustoria on  C. quercusbatatoides . (C) Exemplar of parasitic vine wrapping tightly around the stem directly proximate to a gall induced by the wasp  Disholcaspis quercusvirens  on  Q. geminata . (D) Field site where love vine,  C. filiformis , is attacking the sand live oak,  Q. geminata , and many of the gall forming wasps on the same host plant.  [SOURCE]   

Figure S2. (A) Cassytha filiformis vine attaching haustoria to a leaf gall induced by the wasp Callirhytis quercusbatatoides on the stem of their host plant, Quercus geminata. (B) Labeled graphic of insect gall, parasitic vine, and vine haustoria on C. quercusbatatoides. (C) Exemplar of parasitic vine wrapping tightly around the stem directly proximate to a gall induced by the wasp Disholcaspis quercusvirens on Q. geminata. (D) Field site where love vine, C. filiformis, is attacking the sand live oak, Q. geminata, and many of the gall forming wasps on the same host plant. [SOURCE]
 

Dissection of the galls revealed that the haustoria were plugged into the gall itself, not the wasp larvae. However, because the larvae simply cannot develop without the nutrients and protection provided by the gall, Eagan and his colleagues conclude that these do indeed represent a case of a parasite being parasitized by another parasite.

At this point, the next question that must be asked is how common is this in love vine or, for that matter, across all other parasitic plants that utilize haustoria. Considering that parasites of parasites are nothing new in the biosphere, it is a safe bet that this will not be the last time this phenomenon is discovered.

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

Further Reading: [1]

The Carnivorous Dewy Pine

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The dewy pine is definitely not a pine, however, it is quite dewy. Known scientifically as Drosophyllum lusitanicum, this carnivore is odd in more ways than one. It is also growing more and more rare each year.

One of the strangest aspects of dewy pine ecology is its habitat preferences. Whereas most carnivorous plants enjoy growing in saturated soils or even floating in water, the dewy pine's preferred habitats dry up completely for a considerably portion of the year. Its entire distribution consists of scattered populations throughout the western Iberian Peninsula and northwest Morocco.

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Its ability to thrive in such xeric conditions is a bit of a conundrum. Plants stay green throughout the year and produce copious amounts of sticky mucilage as a means of catching prey. During the summer months, both air and soil temperatures can skyrocket to well over 100°F (37 °C). Though they possess a rather robust rooting system, dewy pines don't appear to produce much in the way of fine roots. Because of this, any ground water presence deeper in the soil is out of their reach. How then do these plants manage to function throughout the driest parts of the year?

During the hottest months, the only regular supply of water comes in the form of dew. Throughout the night and into early morning, temperatures cool enough for water to condense out of air. Dew covers anything with enough surface area to promote condensation. Thanks to all of those sticky glands on its leaves, the dewy pine possesses plenty of surface area for dew to collect. It is believed that, coupled with the rather porous cuticle of the surface of its leaves, the dewy pine is able to obtain water and reduce evapotranspiration enough to keep itself going throughout the hottest months. 

 Dewy pine leaves unfurl like fern fiddle heads as they grow.

Dewy pine leaves unfurl like fern fiddle heads as they grow.

As you have probably guessed at this point, those dewy leaves do more than photosynthesize and collect water. They also capture prey. Carnivory in this species evolved in response to the extremely poor conditions of their native soils. Nutrients and minerals are extremely low, thus selecting for species that can acquire these necessities via other means. Each dewy pine leaf is covered in two types of glands: stalked glands that produce sticky mucilage, and sessile glands that secrete digestive enzymes and absorb nutrients.

Their ability to capture insects far larger than one would expect is quite remarkable. The more an insect struggles, the more it becomes ensnared. The strength of the dewy pines mucilage likely stems from the fact that the leaves do not move like those of sundews (Drosera spp.). Once an insect is stuck, there is not much hope for its survival. Living in an environment as extreme as this, the dewy pine takes no chances.

Drosophyllum_lusitanicum_concurso_reserva_biosfera.jpg

The taxonomic affinity of the dewy pine has been a source of confusion as well. Because of its obvious similarity to the sundews, the dewy pine has long been considered a member of the family Droseraceae. However, although recent genetic work does suggest a distant relationship with Droseraceae and Nepenthaceae, experts now believe that the dewy pine is unique enough to warrant its own family. Thus, it is now the sole species of the family Drosophyllaceae.

Sadly, the dewy pine is losing ground fast. From industrialization and farming to fire suppression, dewy pines are running out of habitat. It is odd to think of a plant capable of living in such extreme conditions as being overly sensitive but that is the conundrum faced by more plants than just the dewy pine. Without regular levels of intermediate disturbance that clear the landscape of vegetation, plants like the dewy pine quickly get outcompeted by more aggressive plant species. Its the fact that dewy pine can live in such hostile environments that, historically, has kept its populations alive and well.

Drosophyllum_lusitanicum_Habitus_2011-4-21_SierraMadrona.jpg

What's more, it appears that dewy pines have trouble getting their seeds into new habitats. Low seed dispersal ability means populations can be cut off from suitable habitats that are only modest distances away. Without a helping hand, small, localized populations can disappear alarmingly fast. The good news is, conservationists are working hard on identifying what must be done to ensure the dewy pine is around for future generations to enjoy.

Changes in land use practices, prescribed fires, wild land conservation, and incentives for cattle farmers to adopt more traditional rather than industrial grazing practices may turn the table on dewy pine extinction. Additionally, dewy pines have become a sort of horticultural oddity over the last decade or so. As dedicated growers perfect germination and growing techniques, ex situ conservation can help maintain stocks of genetic material around the globe.

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

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

 

 

Cycad Pollinators

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When it comes to insect pollination, flowering plants get all of the attention. However, flowers aren't the only game in town. More and more we are beginning to appreciate the role insects play in the pollination of some gymnosperm lineages. For instance, did you know that many cycad species utilize insects as pollen vectors? The ways in which these charismatic gymnosperms entice insects is absolutely fascinating and well worth understanding in more detail.

Cycads or cycad-like plants were some of the earliest gymnosperm lineages to arise on this planet. They did so long before familiar insects like bees, wasps, and butterflies came onto the scene. It had long been assumed that, like a vast majority of extant gymnosperms, cycads relied on the wind to get pollen from male cones to female cones. Indeed, many species certainly utilize to wind to one degree or another. However, subsequent work on a few cycad genera revealed that wind might not cut it in most cases.

 White-haired cycad ( Encephalartos friderici-guilielm i)

White-haired cycad (Encephalartos friderici-guilielmi)

It took placing living cycads into wind tunnels to obtain the first evidence that something strange might be going on with cycad pollination. The small gaps on the female cones were simply too tight for wind-blown pollen to make it to the ovules. Around the same time, researchers began noting the production of volatile odors and heat in cycad cones, providing further incentives for closer examination.

Subsequent research into cycad pollination has really started to pay off. By excluding insects from the cones, researchers have been able to demonstrate that insects are an essential factor in the pollination of many cycad species. What's more, often these relationships appear to be rather species specific.

  Cycadophila yunnanensis ,  C. nigra , and other beetles on a cone of  Cycas  sp.

Cycadophila yunnanensis, C. nigra, and other beetles on a cone of Cycas sp.

By far, the bulk of cycad pollination services are being performed by beetles. This makes a lot of sense because, like cycads, beetles evolved long before bees or butterflies. Most of these belong to the superfamily Cucujoidea as well as the true weevils (Curculionidae). In some cases, beetles utilize cycad cones as places to mate and lay eggs. For instance, male and female cones of the South African cycad Encephalartos friderici-guilielmi were found to be quite attractive to at least two beetle genera. 

Beetles and their larvae were found on male cones only after they had opened and pollen was available. Researchers were even able to observe adult beetles emerging from pupae within the cones, suggesting that male cones of E. friderici-guilielmi function as brood sites. Adult beetles carrying pollen were seen leaving the male cones and visiting the female cones. The beetles would crawl all over the fuzzy outer surface of the female cones until they became receptive. At that point, the beetles wriggle inside and deposit pollen. Seed set was significantly lower when beetles were excluded.

 Male cone of  Zamia furfuracea  with a mating (lek) assembly of  Rhopalotria mollis  weevils.

Male cone of Zamia furfuracea with a mating (lek) assembly of Rhopalotria mollis weevils.

For the Mexican cycad Zamia furfuracea, weevils also utilize cones as brood sites, however, the female cones go to great lengths to protect themselves from failed reproductive efforts. The adult weevils are attracted to male cones by volatile odors where they pick up pollen. The female cones are thought to also emit similar odors, however, larvae are not able to develop within the female cones. Researchers attribute this to higher levels of toxins found in female cone tissues. This kills off the beetle larvae before they can do too much damage with their feeding. This way, the cycad gets pollinated and potentially harmful herbivores are eliminated. 

Beetles also share the cycad pollination spotlight with another surprising group of insects - thrips. Thrips belong to an ancient order of insects whose origin dates back to the Permian, some 298 million years ago. Because they are plant feeders, thrips are often considered pests. However, for Australian cycads in the genus Macrozamia, they are important pollinators.

  Macrozamia macleayi  female cone.

Macrozamia macleayi female cone.

Thrip pollination was studied in detail in at least two Macrozamia species, M. lucida and M. macleayi. It was noted that the male cones of these species are thermogenic, reaching peak temperatures of around   104 °F (40 °C). They also produce volatile compounds like monoterpenes as well as lots of CO2 and water vapor during this time. This spike in male cone activity also coincides with a mass exodus of thrips living within the cones.

 Thrips ( Cycadothrips chadwicki ) leaving a thermogenic pollen cone of  Macrozamia lucida.

Thrips (Cycadothrips chadwicki) leaving a thermogenic pollen cone of Macrozamia lucida.

Thrips apparently enjoy cool, dry, and dark places to feed and breed. That is why they love male Macrozamia cones. However, if the thrips were to remain in the male cones only, pollination wouldn't occur. This is where all of that male cone metabolic activity comes in handy. Researchers found that the combination of rising heat and humidity, and the production of monoterpenes aggravated thrips living within the male cones, causing them to leave the cones in search of another home.

Inevitably many of these pollen-covered thrips find themselves on female Macrozamia cones. They crawl inside and find things much more to their liking. It turns out that female Macrozamia cones do not produce heat or volatile compounds. In this way, Macrozamia are insuring pollen transfer between male and female plants.

 Thrips up close.

Thrips up close.

Pollination in cycads is a fascinating subject. It is a reminder that flowering plants aren't the only game in town and that insects have been providing such services for eons. Additionally, with cycads facing extinction threats on a global scale, understanding pollination is vital to preserving them into the future. Without reproduction, species will inevitably fail. Many cycads have yet to have their pollinators identified. Some cycad pollinators may even be extinct. Without boots on the ground, we may never know the full story. In truth, we have only begun to scratch the surface of cycads and their pollinators.

Photo Credits: [1] [2] [3] [4] [5] [6] [7]

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