Daffodil Insights

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Daffodils seem to be everywhere. Their horticultural popularity means that, for many of us, these plants are among the first flowers we see each spring. Daffodils are so commonplace that it's as if they evolved to live in our gardens and nowhere else. Indeed, daffodils have had a long, long history with human civilization, so much so that it is hard to say when our species first started to cohabitate. Our familiarity with these plants belies an intriguing natural history. What follows is a brief overview of the world of daffodils. 

If you are like me, then you may have gone through most of your life not noticing much difference between garden variety daffodils. Though many of us will be familiar with only a handful of daffodil species and cultivars, these introductions barely scratch the surface. One may be surprised to learn that as of 2008, more than 28,000 daffodil varieties have been named and that number continues to grow each and every year. Even outside of the garden, there is some serious debate over the number of daffodil species, much of this having to do with what constitutes a species in this group.

  Narcissus poeticus

Narcissus poeticus

As I write this, all daffodils fall under the genus Narcissus. Estimates as to the number of species within Narcissus range from as few as 50 to as many as 80. The genus itself sits within the family Amaryllidaceae and is believed to have originated somewhere between the late Oligocene and early Miocene, some 18 to 30 million years ago. Despite its current global distribution, Narcissus are largely Mediterranean plants, with peak diversity occurring on the Iberian Peninsula. However, thanks to the aforementioned long and complicated history in cultivation, it has become quite difficult to understand the full range of diversity in form and habitat of many species. To understand this, we first need to understand a bit about their reproductive habits.

Much of the evolution of Narcissus seems to center around floral morphology and geographic isolation. More specifically, the length of the floral tube or "corona" and the position of the sexual organs within, dictates just who can effectively pollinate these plants. The corona itself is not made up of petals or sepals but instead, its tube-like appearance is due to a fusion of the stamens into the famous trumpet-like tube we know and love.

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Variation in corona shape and size has led to the evolution of three major pollination strategies within this genus. The first form is the daffodil form, whose stigma is situated at the mouth of the corolla, well beyond the 6 anthers. This form is largely pollinated by larger bees. The second form is the paperwhite form, whose stigma is situated more closely to or completely below the anthers at the mouth of the corona. This form is largely pollinated by various Lepidoptera as well as long tongued bees and flies. The third form is the triandrus form, which exhibits three distinct variations on stigma and anther length, all of which are situated deep within the long, narrow corona. The pendant presentation of the flowers in this group is thought to restrict various butterflies and moths from entering the flower in favor of bees.

  Narcissus tazetta

Narcissus tazetta

The variations on these themes has led to much reproductive isolation among various Narcissus populations. Plants that enable one type of pollinator usually do so at the exclusion of others. Reproductive isolation plus geographic isolation brought on by differences in soil types, habitat types, and altitudinal preferences is thought to have led to a rapid radiation of these plants across the Mediterranean. All of this has gotten extremely complicated ever since humans first took a fancy to these bulbs.

  Narcissus cyclamineus

Narcissus cyclamineus

Reproductive isolation is not perfect in these plants and natural hybrid zones do exist where the ranges of two species overlap. However, hybridization is made much easier with the helping hand of humans. Whether via landscape disturbance or direct intervention, human activity has caused an uptick in Narcissus hybridization. For centuries, we have been mixing these plants and moving them around with little to no record as to where they originated. What's more, populations frequently thought of as native are actually nothing more than naturalized individuals from ancient, long-forgotten introductions. For instance, Narcissus populations in places like China, Japan, and even Great Britain originated in this manner.

All of this mixing, matching, and hybridizing lends to some serious difficulty in delineating species boundaries. It would totally be within the bounds of reason to ask if some of the what we think of as species represent true species or simply geographic varieties on the path to further speciation. This, however, is largely speculative and will require much deeper dives into Narcissus phylogenetics.

  Narcissus triandrus

Narcissus triandrus

Despite all of the confusion surrounding accurate Narcissus taxonomy, there are in fact plenty of true species worth getting to know. These range in form and habit far more than one would expect from horticulture. There are large Narcissus, small Narcissus, there are Narcissus with yellow flowers and Narcissus with white flowers, some with upright flowers and some with pendant flowers. There are even a handful of fall blooming Narcissus. The variety of this genus is staggering if you are not prepared for it.

  Narcissus viridiflorus  - a green, fall-blooming daffodil

Narcissus viridiflorus - a green, fall-blooming daffodil

After pollination, the various Narcissus employ a seed dispersal strategy that doesn't get talked about enough in reference to this group. Attached to each hard, black seed are fatty structures known as eliasomes. Eliasomes attract ants. Like many spring flowering plant species around the globe, Narcissus utilize ants as seed dispersers. Ants pick up the seeds and bring them back to their nests. They go about removing the eliasomes and then discard the seed. The seed, safely tucked away in a nutrient-rich ant midden, has a much higher chance of germination and survival than if things were left up to simple chance. It remains to be seen whether or not Narcissus obtain similar seed dispersal benefits from ants outside of their native range. Certainly Narcissus populations persist and naturalize readily, however, I am not aware if ants have any part in the matter.

 The endangered  Narcissus alcaracensis .

The endangered Narcissus alcaracensis.

Despite their popularity in the garden, many Narcissus are having a hard go of it in the wild. Habitat destruction, climate change, and rampant collecting of wild bulbs are having serious impacts on Narcissus numbers. The IUCN considered at least 5 species to be endangered and a handful of some of the smaller species already thought to be extinct in the wild. In response to some of these issues, protected areas have been established that encompass at least some of the healthy populations that remain for some of these species.

If you are anything like me, you have ignored Narcissus for far too long. Sure, they aren't native to the continent on which I live, and sure, they are one of the most commonly used plants in a garden setting, but every species has a story to tell. I hope that, armed with this new knowledge, you at least take a second look at the Narcissus popping up around your neighborhood. More importantly, I hope this introduction makes you appreciate their wild origins and the fact that we still have much to learn about these plants. I have barely scratched the surface of this genus and there is more more information out there worth perusing. Finally, I hope we can do better for the wild progenitors of our favorite garden plants. They need all the help they can get and unless we start speaking up and working to preserve wild spaces, all that will remain are what we have in our gardens and that is not a future I want to be a part of.

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

Further Reading: [1] [2] [3] [4] [5] [6] [7] [8] [9]

 

Do Yeasts Aid Pollination For the Stinking Hellebore?

Whether they are growing in their native habitat or in some far away garden, Hellebores are some of the earliest plants to bloom in the spring. Hellebore flowers can often be seen blooming long before the snow has melted away. All early blooming plant species are faced with the challenge of attracting pollinators. Though the competition for insect attention is minimal among these early bloomers, only the hardiest insects are out and about on cold, dreary days. It stands to reason then that anything that can entice a potential pollinator would be of great benefit for a plant.

That is why the presence of yeast in the nectar of at least one species of Hellebore has attracted the attention of scientists. The species in question is known scientifically as Helleborus foetidus. The lack of appeal in its binomial is nothing compared to its various common names. One can often find H. foetidus for sale under names like the "stinking hellebore" or worse, "dungwort." All of these have to do with the unpleasant aroma given off by its flowers and bruised foliage. Surprisingly, that is not the topic of this post.

What is more intriguing about the flowers of H. foetidus is that the nectar produced by its smelly green flowers harbors dense colonies of yeast. Yeasts are everywhere on this planet and despite their economic importance, little is known about how they function in nature. For instance, what the heck are these yeast colonies doing in the nectar of this odd Hellebore?

To test this, two researchers from the Spanish National Research Council manipulated yeast colonies within the flowers to see what might be happening. It turns out, yeast in the nectar of H. foetidus actually warms the flowers. As the yeast feed on the sugars within the nectar, their metabolic activity can raise the temperature of the flowers upwards of 2 °C above the ambient. To date, the only other ways in which floral heating has been achieved is either via specific metabolic processes within the floral tissues or by direct heating from the sun. 

In heating the flowers, these yeast colonies may be having serious impacts on the reproductive success of H. foetidus. For starters, these plants are most at home under the forest canopies of central and western Europe. What's more, many populations find themselves growing in the dense shade of evergreens. This completely rules out the ability to utilize solar energy to heat blooms. Additionally, floral heat can mean more visits by potential pollinators. Experiments have shown that bees preferentially visit flowers that are slightly warmer than ambient temperatures. Even the flowers themselves can benefit from that heat. Warmer flowers have higher pollination rates and better seed set.

  Bombus terrestris  was one of the most common floral visitors of  Helleborus foetidus.

Bombus terrestris was one of the most common floral visitors of Helleborus foetidus.

Yeast colonies also have their downsides. The heat generated by the yeast comes from the digestion of sugars. Indeed, nectar housing yeast colonies had drastically reduced sugar loads than nectar without yeast. This has the potential to offset many of the benefits of floral warming in large part because bees are good at discriminating. Bees are visiting these blooms as a food source and by diminishing the sugar loads within the nectar, the yeast may be turning bees off to this potential source. The question then becomes, do bees prefer heat over sugar-rich food? The authors think there might be a trade-off, with bees preferring heated flowers on colder days and sugar-rich flowers on warmer days.

Though the authors found evidence for heating, they did not test for pollinator preference. All we know at this point is that yeast in the nectar significantly warms H. foetidus flowers. Until we know whether bees are making a decision based on yeast colonies, we cannot say whether this is a case of a mutualism or form of parasitism. Such questions will be the subject of future research. Nonetheless, this is a wonderful example of why we need to pay way more attention to the function of yeasts in nature.

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

Further Reading: [1] [2]

Meet the Ocotillo

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I love the ocotillo (Fouquieria splendens) for many reasons. It is an impossible plant to miss with its spindly, spine-covered stems. It is a lovely plant that is right at home in the arid parts of southwestern North America. Beyond its unique appearance, the ocotillo is a fascinating and important component of the ecology of this region.

My first impression of ocotillo was interesting. I could not figure out where this plant belonged on the tree of life. As a temperate northeasterner, one can forgive my taxonomic ignorance of this group. The family from which it hails, Fouquieriaceae, is restricted to southwestern North America. It contains one genus (Fouquieria) and about 11 species, all of which are rather spiky in appearance.

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Of course, those spines serve as protection. Resources like water are in short supply in desert ecosystems so these plants ensure that it is a real struggle for any animal looking to take a bite. Those spines are tough as well. One manged to pierce the underside of my boot during a hike and I was lucky that it just barely grazed the underside of my foot. Needless to say, the ocotillo is a plant worthy of attention and respect.

One of the most striking aspects of ocotillo life is how quickly these plants respond to water. As spring brings rain to this region of North America, ocotillo respond with wonderful sprays of bright red flowers situated atop their spindly stems. These blooms are usually timed so as to take advantage of migrating hummingbirds and emerging bees. The collective display of a landscape full of blooming ocotillo is jaw-droppingly gorgeous and a sight one soon doesn't forget. It is as if the whole landscape has suddenly caught on fire. Indeed, the word "ocotillo" is Spanish for "little torch."

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Flowering isn't the only way this species responds to the sudden availability of water. A soaking rain will also bring about an eruption of leaves, turning its barren, white stems bright green. The leaves themselves are small and rather fragile. They do not have the tough, succulent texture of what one would expect out of a desert specialist. That is because they don't have to ride out the hard times. Instead, ocotillo are what we call a drought deciduous species, producing leaves when times are good and water is in high supply, and dropping them as soon as the soil dries out.

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This cycle of growing and dropping leaves can and does happen multiple times per year. It is not uncommon to see ocotillo leaf out up to 4 or 5 times between spring and fall. During the rest of the year, ocotillo relies on chlorophyll in its stems for its photosynthetic needs. Interestingly enough, this poses a bit of a challenge when it comes to getting enough CO2. Whereas leaves are covered in tiny pours called stomata which help to regulate gas exchange, the stems of an ocotillo are a lot less porous, making it a challenge to get gases in and out. This is where the efficient metabolism of this plant comes in handy.

All plants undergo respiration like you and me. The carbohydrates made during photosynthesis are broken down to fuel the plant and in doing so, CO2 is produced. Amazingly, the ocotillo (as well as many other plants that undergo stem photosynthesis) are able to recycle the CO2 generated by cellular respiration back into photosynthesis within the stem. In this way, the ocotillo is fully capable of photosynthesis even without leaves.

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Through the good times and the bad, the ocotillo and its relatives are important components of desert ecology. They are as hardy as they are beautiful and getting to see them in person has been a remarkable experience. They ad a flare of surreality to the landscape that must be seen in person to believe.

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

From Herbivore to Pollinator Thanks to a Parasitoid

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In the Atlantic forests of Brazil resides a small orchid known scientifically as Dichaea cogniauxiana. Like most plant species, this orchid experiences plenty of pressure from herbivores. It faces rather intense pressures from two species of weevil in the genus Montella. These weevils are new to science and have yet been given full species status. What's more, they don't appear to eat the leaves of D. cogniauxiana. Instead, female weevils lay eggs in the developing fruits and the larvae hatch out and consume the seeds within. In other words, they treat the fruits like a nursery chamber.

This is where this relationship gets interesting. You see, the weevils themselves appear to take matters into their own hands. Instead of waiting to find already pollinated orchids, an event that can be exceedingly rare in the dense Amazonian forests, these weevils go about pollinating the orchids themselves. Females have been observed picking up orchid pollinia and depositing the pollen onto the stigmas. In this way, they ensure that there will be developing fruits in which they can raise their young.

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Left unchecked, the weevil larvae readily consume all of the developing seeds within the pod, an unfortunate blow to the reproductive efforts of this tiny orchid. However, the situation changes when parasitoid wasps enter the mix. The wasps are also looking for a place to rear their young but the wasp larvae do not eat orchid seeds. Instead, the wasps must find juicy weevil larvae in which to lay their eggs. When the wasp larvae hatch out, they eat the weevil larvae from the inside out and this is where things get really interesting.

The wasp larvae develop at a much faster rate than do the weevil larvae. As such, they kill the weevil long before it has a chance to eat all of the orchid seeds. By doing so, the wasp has effectively rescued the orchids reproductive effort. Over a five year period, researchers based out of the University of Campinas found that orchid fruits in which wasp larvae have killed off the weevil larvae produced nearly as many seeds as uninfected fruits. As such, the parasitoid wasps have made effective pollinators out of otherwise destructive herbivorous weevils.

The fact that a third party (in this case a parasitic wasp) can change a herbivore into an effective pollinator is quite remarkable to say the least. It reminds us just how little we know about the intricate ways in which species interact and form communities. The authors note that even though pollination in this case represents selfing and thus reduced genetic diversity, it nonetheless increases the reproductive success of an orchid that naturally experiences low pollination rates to begin with. In the hyper diverse and competitive world of Brazilian rainforests, even self-pollination cab be a boost for this orchid.

Photo Credits: [1] [2]

Further Reading: [1]

Apocynaceae Ant House

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

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

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

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

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

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

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

 The flowers of   Dischidia ovata

The flowers of Dischidia ovata

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

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

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

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

Further Reading: [1]

Early Land Plants Made The World Muddy

  Cooksonia  is one of the earliest land plants to have evolved.

Cooksonia is one of the earliest land plants to have evolved.

Try to picture the world before life moved onto land. It would have been a vastly different landscape than anything we know today. For one, there would have been no soil. Before life moved onto land, there was nothing organic around to facilitate soil formation. This would have changed as terrestrial habitats were slowly colonized by microbes and eventually plants. A recent paper published in Science is one of the first to demonstrate that the rise in certain sediments on land, specifically mud-forming clays, coincided with the rise in deep-rooted land plants.

This was no small task. The research duo had to look at thousands of reports spanning from the Archean eon, some 3.5 billion years ago, to the Carboniferous period, some 358 million years ago. By looking for the relative amounts of a sedimentary rock called mudrock in terrestrial habitats, they were able to see how the geology of terrestrial habitats was changing through time. What they found was that the presence of mudrock increased by orders of magnitude around the same time as early land plants were beginning to colonize land. Before plants made it onto land, mudrocks comprised a mere 1% of terrestrial sediments. By the end of the Carboniferous period, mudrocks had risen to 26%.

This begs the question, why are mudrocks so significant? What do they tell us about what was going on in terrestrial environments? A key to these questions lies in the composition of mudrocks themselves. Mudrock is made up of fine grained sediments like clay. There are many mechanisms by which clay can be produced and certainly this was going on well before plants made it onto the scene. The difference here is in the quantity of clay-like minerals in these sediments. Whereas bacteria and fungi do facilitate the formation of clay minerals, they do so in small quantities.

 A little bit of moss goes a long way for erosion control!

A little bit of moss goes a long way for erosion control!

The real change came when plants began rooting themselves into the soil. In pushing their roots down into sediments, plants act as conduits for increased weathering of said minerals. Roots not only increase the connectivity between subsurface geology and the atmosphere, they also secrete substances like organic acids and form symbiotic relationships with cyanobacteria and fungi that accelerate the weather process. No purely tectonic or chemical processes can explain the rate of weathering that must have taken place to see such an increase in these fine grained minerals.

What's more, the presence of rooted plants on land would have ensured that these newly formed muds would have stuck around on the landscape much longer. Whereas in the absence of plants, these sediments would have been washed away into the oceans, plants were suddenly holding onto them. Plant roots act as binders, holding onto soil particles and preventing erosion. Aside from their roots, the rest of these early land plants would have also held onto sediments via a process known as the baffling effect. As water and wind pick up and move sediments, they inevitably become trapped in and around the stems and leaves of plants. Even tiny colonies of liverworts and moss are capable of doing this and entire mats of these would have contributed greatly to not only the formation of these sediments, but their retention as well.

The movement of plants onto land changed the course of history. It was the beginning of massive changes to come and much of that started with the gradual formation of soils. We owe everything to these early botanical pioneers.

Photo Credit: [1]

Further Reading: [1]

The Mighty Saguaro Cactus

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Where does one begin with a plant like the saguaro cactus (Carnegiea gigantea)? It is recognized the world over for its iconic appearance yet its native range is disproportionately small compared to its popularity. It is easily one of the most spectacular plants I have ever encountered and I will never forget the sound the wind makes as it blows over its spiny pleated trunk. It would be impossible to sum up our collective knowledge of this species in one article, however, I feel that some form of an introduction is necessary. Today I want to honor this icon of the Sonoran Desert.

The saguaro is the only member of the genus Carnegiea, which is part of a subtribe of cacti characterized by their columnar appearance. Despite its unique taxonomic affinity, the evolutionary origins of this cactus remains a bit of a mystery. Though it is undoubtedly related to other columnar cacti of the Americas, a proper family tree seems to be just out of our reach. Due to lots of convergent and parallel evolution as well as conflicts between genealogies and species histories, we still aren't sure of its evolutionary origins. What we do know about this species on a genetic level is nonetheless quite interesting. For instance the saguaro has one of the smallest chloroplast genomes of any non-parasitic plant and we aren’t exactly sure why this is the case.

Saguaro are long lived cacti. Estimating age of a cactus can be rather tricky considering that they don’t produce annual growth rings. This is where long term monitoring projects have come in handy. By observing hundreds of saguaro throughout the Sonoran Desert, experts believe that saguaro can regularly reach ages of 150 to 170 years and some individuals may be able to live for more than 200 years. Amazingly, it is thought that saguaro will not begin to grow their characteristic arms until they reach somewhere around 50 to 100 years of age. That being said, some saguaro never bother growing arms. It all depends on where the conditions they experience throughout their lifetime.

Growth for a saguaro depends on where they are rooted. Under favorable conditions, a saguaro can grow to heights of 50 feet or more, with the world record holder clocking in at a whopping 78 feet in height. Such growth becomes all the more impressive when you realize just how agonizingly slow the process can be. Studies have shown that juvenile saguaro only put on about 1.5 inches of growth in their first eight years of life.

Despite preconceived notions about the hardy nature of most cacti, saguaro have proven to be rather specific in their needs. They are limited in their growth and distribution by the availability of water and warm temperatures. Saguaro, especially young individuals, cannot tolerate periods of prolonged frost. Additionally, germination and seedling survival occur most frequently only during the wettest years. In fact, one study showed that successful years for reproduction in these beloved cacti were tied to volcanic eruptions that cooled the climate just enough to allow the young saguaro to become established.

Outside of volcanic eruptions, saguaro appear to have friends in the surrounding vegetation. Studies have shown that saguaro seedlings seem to do best when growing under the shade of trees like the palo verde (Parkinsonia florida), ironwood (Olneya tesota), and mesquite (Prosopis velutina). The microclimates produced by these trees are much more favorable for saguaro growth than are open desert conditions. In essence, these trees serve as nurseries for young saguaro until they are large enough to handle more exposed conditions. Their nursery habits are not mutually beneficial however as research suggests that saguaro eventually compete with the trees that once protected them for precious resources like nutrients and water.

 Saguaros outgrowing their palo verde nurse tree. 

Saguaros outgrowing their palo verde nurse tree. 

At roughly 35 years of age, a saguaro will begin to flower. Flowers are small compared to the size of the cactus but they are abundant. Most flowers are produced at the apex of the cactus and it is thought that the growth of saguaro arms is largely a way of increasing the reproductive potential of large individuals. The flowers are cream colored and night scented. They open in the evening but will stay open and continue to produce nectar well into the morning hours.

Though a wide variety of animals will visit these flowers, the main pollinators are bees during the day and lesser long-nosed bats at night. Interestingly, it has been found that certain amino acids within the nectar of the saguaro can actually help female bats sustain lactation while raising their young, making them a valuable food source for these flying mammals. Catering to such a broad spectrum of potential pollinators is thought to have evolved as a means of increasing seed set. Each saguaro ovary contains many ovules and the more pollen that makes it onto the stigma, the more seeds will be produced.

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 A lesser long-nosed bat pollinates a saguaro bloom.

A lesser long-nosed bat pollinates a saguaro bloom.

Due to their size and abundance, it is easy to understand why the saguaro is such an ecologically important species in the Sonoran Desert ecosystem. In essence, they function similar to trees in that they serve as vital sources of shelter and food for myriad desert animals. Woodpeckers, especially the gila and the gilded flicker, regularly hollow out and build nests in saguaro trunks. These hollows are subsequently used by many different bird, mammal, and reptile species. The flowers and fruits are important sources of food for wildlife.

 Gila woodpecker with its nesting hole.

Gila woodpecker with its nesting hole.

 Gila woodpecker holes become homes for other birds like owls. 

Gila woodpecker holes become homes for other birds like owls. 

 On rare occasions, woodpecker holes can even become home to other cacti!

On rare occasions, woodpecker holes can even become home to other cacti!

I sincerely hope that this brief introduction does at least some justice to the wonderful organism that is the saguaro cactus. The Sonoran Desert would be a shell of an ecosystem without its presence. What’s more, it has played a significant role in the culture of this region for millennia. Though it appears quite numerous on the landscape, the long-term status of the saguaro is cause for concern. Numerous declines have been reported throughout its range. With its slow growth rates and infrequent recruitment events, the saguaro can be quite sensitive to rapid changes in its environment. Luckily it has received special protection laws throughout its US range.

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


Further Reading: [1] [2] [3] [4] [5] [6] [7] [8] [9] 

The Wild World of the Creosote Bush

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Apart from the cacti, the real rockstar of my Sonoran experience was the creosote bush (Larrea tridentata). Despite having been quite familiar with creosote as an ingredient, I admit to complete ignorance of the plant from which it originates. Having no familiarity with the Sonoran Desert ecosystem, I was walking into completely new territory in regard to the flora. It didn’t take long to notice creosote though. Once we hit the outskirts of town, it seemed to be everywhere.

If you are in the Mojave, Sonoran, and Chihuahuan Deserts of western North America, you are never far from a creosote bush. They are medium sized, slow growing shrubs with sprays of compact green leaves, tiny yellow flowers, and fuzzy seeds. Apparently what is thought of as one single species is actually made up of three different genetic populations. The differences between these has everything to do with chromosome counts. Populations in the Mojave Desert have 78 chromosomes, Sonoran populations have 52 chromosomes, and Chihuahuan have 26. This may have to do with the way in which these populations have adapted to the relative amounts of rainfall each of these deserts receive throughout the year, however, it is hard to say for sure.

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Regardless, creosote is supremely adapted to these xeric ecosystems. For starters, their branching architecture coupled with their tiny leaves are arranged so as to make the most out of favorable conditions. If you stare at these shrubs long enough, you may notice that their branches largely orient towards the southeast. Also, their leaves tend to be highly clustered along the branches. It is thought that this branching architecture allows the creosote to minimize water loss while maximizing photosynthesis.

Deserts aren’t hot 24 hours per day. Night and mornings are actually quite cool. By taking advantage of the morning sun as it rises in the east, creosote are able to open their stomata and commence photosynthesis during those few hours when evapotranspiration would be at its lowest. In doing so, they are able to minimize water loss to a large degree. Although their southeast orientation causes them to miss out on afternoon and evening sun to a large degree, the benefits of saving precious water far outweigh the loss to photosynthesis. The clustering of the leaves along the branches may also reduce overheating by providing their own shade. Coupled with their small size, this further reduces heat stress and water loss during the hottest parts of the day.

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Creosote also secrets lots of waxy, resinous compounds. These coat the leaves and to some extent the stems, making them appear lacquered. It is thought that this also helps save water by reducing water loss through the leaf cuticle. However, the sheer diversity of compounds extracted from these shrubs suggests other functions as well. It is likely that at least some of these compounds are used in defense. One study showed that when desert woodrats eat creosote leaves, the compounds within caused the rats to lose more water through their urine and feces. They also caused a reduction in the amount of energy the rats were able to absorb from food. In other words, any mammal that regularly feeds on creosote runs the risk of both dehydration and starvation. This isn’t the only interesting interaction that creosote as with rodents either. Before we get to that, however, we first need to discuss roots.

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Creosote shrubs have deep root systems that are capable of accessing soil water that more shallowly rooted plants cannot. This brings them into competition with neighboring plants in intriguing ways. When rainfall is limited, shallowly rooted species like Opuntia gain access to water before it has a chance to reach deeper creosote roots. Surprisingly this happens more often than you would think. The branching architecture of creosote is such that it tends to accumulate debris as winds blow dust around the desert landscape. As a result, the soils directly beneath creosote often contain elevated nutrients. This coupled with the added shade of the creosote canopy means that seedlings that find themselves under creosote bushes tend to do better than seedlings that germinated elsewhere. As such, creosote are considered nurse plants that actually facilitate the growth and survival of surrounding vegetation. So, if recruitment and resulting competition from vegetation can become such an issue for long term creosote survival, why then do we still so much creosote on the landscape?

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The answer may lie in rodents and other burrowing animals in these desert ecosystems. Take a look at the base of a large creosote and you will often find the ground littered with burrows. Indeed, many a mammal finds the rooting zone of the creosote shrub to be a good place to dig a den. When these animals burrow under shallowly rooted desert plants, many of them nibble on and disturb the rooting zones. Over the long-term, this can be extremely detrimental for the survival of shallow rooted species. This is not the case for creosote. Its roots run so deep that most burrowing animals cannot reach them. As such, they avoid most of the damage that other plants experience. This lends to a slight survival advantage for creosote at the expense of neighboring vegetation. In this way, rodents and other burrowing animals may actually help reduce competition for the creosote.

Barring major disturbances, creosote can live a long, long time. In fact, one particular patch of creosote growing in the Mojave Desert is thought to be one of the oldest living organisms on Earth. As creosote shrubs grow, they eventually get to a point in which their main stems break and split. From there, they begin producing new stems that radiate out in a circle from the original plant. These clones can go on growing for centuries. By calculating the average growth rate of these shrubs, experts have estimated that the Mojave specimen, affectionately referred to as the “King Clone,” is somewhere around 11,700 years old!

 The ring of creosote that is King Clone.

The ring of creosote that is King Clone.

For creosote, its slow and steady wins the race. They are a backbone of North American desert ecosystems. Their structure offers shelter, their seeds offer food, and their flowers support myriad pollinators. Creosote is one shrub worthy of our respect and admiration.

Photo Credit: [1] [2]

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

Palo Verde

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One of the first plants I noticed upon arriving in the Sonoran Desert were these small spiny trees without any leaves. The reason they caught my eye was that every inch of them was bright green. It was impossible to miss against the rusty brown tones of the surrounding landscape. It didn’t take long to track down the identity of this tree. What I was looking at was none other than the palo verde (Parkinsonia florida).

Palo verde belong to a small genus of leguminous trees. Parkinsonia consists of roughly 12 species scattered about arid regions of Africa and the Americas. The common name of “palo verde” is Spanish for “green stick.” And green they are! Like I said, every inch of this tree gives off a pleasing green hue. Of course, this is a survival strategy to make the most of life in arid climates.

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Despite typically being found growing along creek beds, infrequent rainfall limits their access to regular water supplies. As such, these trees have adapted to preserve as much water as possible. One way they do this is via their deciduous habit. Unlike temperate deciduous trees which drop their leaves in response to the changing of the seasons, palo verde drop their leaves in response to drought. And, as one can expect from a denizen of the desert, drought is the norm. Leaves are also a conduit for moisture to move through the body of a plant. Tiny pours on the surface of the leaf called stomata allow water to evaporate out into the environment, which can be quite costly when water is in short supply.

 The tiny pinnate leaves and pointy stems of the palo verde. 

The tiny pinnate leaves and pointy stems of the palo verde. 

Not having leaves for most of the year would be quite a detriment for most plant species. Leaves, after all, are where most of the photosynthesis takes place. That is, unless, you are talking about a palo verde tree. All of that green coloration in the trunk, stems, and branches is due to chlorophyll. In essence, the entire body of a palo verde is capable of performing photosynthesis. In fact, estimates show that even when the tiny pinnate leaves are produced, a majority of the photosynthetic needs of the tree are met by the green woody tissues.

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Flowering occurs whenever there is enough water to support their development, which usually means spring. They are small and bright yellow and a tree can quickly be converted into a lovely yellow puff ball seemingly overnight. Bees relish the flowers and the eventual seeds they produce are a boon for wildlife in need of an energy-rich meal.

However, it isn’t just wildlife that benefits from the presence of these trees. Other plants benefit from their presence as well. As you can probably imagine, germination and seedling survival can be a real challenge in any desert. Heat, sun, and drought exact a punishing toll. As such, any advantage, however slight, can be a boon for recruitment. Research has found that palo verde trees act as important nurse trees for plants like the saguaro cactus (Carnegiea gigantea). Seeds that germinate under the canopy of a palo verde receive just enough shade and moisture from the overstory to get them through their first few years of growth.

In total, palo verde are ecologically important trees wherever they are native. What’s more, their ability to tolerate drought coupled with their wonderful green coloration has made them into a popular tree for native landscaping. It is certainly a tree I won’t forget any time soon.

Further Reading: [1] [2]

The Other Pawpaws

  Asimina tetramera

Asimina tetramera

The pawpaw (Asimina triloba) has been called "America's forgotten fruit." Once quite popular among Native Americans and settlers alike, it fell out of the public eye until quite recently. If one considers the pawpaw "forgotten" then its relatives have been straight up ignored. Indeed, the pawpaw shares the North American continent with 10 other Asimina species. 

  Asimina angustifolia

Asimina angustifolia

The genus Asimina belongs to a family of plants called the custard apple family - Annonaceae. It is a large family that mostly resides in the tropics. In fact, the genus Asimina is the only group to occur outside of the tropics. Though A. triloba finds itself growing as far north as Canada, the other species within this genus are confined to southeastern North America in coastal plain communities. 

  Asimina parviflora

Asimina parviflora

As I mentioned above, there are 10 other species in the genus and at least one naturally occurring hybrid. For the most part, they all prefer to grow where regular fires keep competing vegetation at bay. They are rather small in stature, usually growing as shrubs or small, spindly trees. They can be rather inconspicuous until it comes time to flower.

  Asimina obovata

Asimina obovata

The flowers of the various Asimina species are relatively large and range in color from bright white to deep red, though the most common flower color seems to be creamy white. The flowers themselves give off strange odors that have been affectionately likened to fermenting fruit and rotting meat. Of course, these odors attract pollinators. Asimina aren't much of a hit with bees or butterflies. Instead, they are mainly visited by blowflies and beetles. 

  Asimina pygmaea

Asimina pygmaea

As is typical of the family, all of the Asimina produce relatively large fruits chock full of hard seeds. Seed dispersal for the smaller species is generally accomplished through the help of mammals like foxes, coyotes, raccoons, opossums, and even reptiles such as the gopher tortoise. Because the coastal plain of North America is a fire-prone ecosystem, most of the Asimina are well adapted to cope with its presence. In fact, most require it to keep their habitat open and free of too much competition. At least one species, A. tetramera, is considered endangered in large part due to fire sequestration.

  Asimina reticulata

Asimina reticulata

All of the 11 or so Asimina species are host plants for the zebra swallowtail butterfly (Eurytides marcellus) and the pawpaw sphinx moth (Dolba hyloeus). The specialization of these two insects and few others has to do with the fact that all Asimina produce compounds called acetogenins, which act as insecticides. As such, only a small handful of insects have adapted to be able to tolerate these toxic compounds. 

  Asimina tetramera

Asimina tetramera

Sadly, like all other denizens of America's coastal plain forest, habitat destruction is taking its toll on Asimina numbers. As mentioned above, at least one species (A. tetramera) is considered endangered. We desperately need to protect these forest habitats. Please support a local land conservation organization like the Partnership For Southern Forestland Conservation today!

See a list of the Asimina of North America here: [1] 

Photo Credits: Wikimedia Commons

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

An Endangered Iris With An Intriguing Pollination Syndrome

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The Golan iris (Iris hermona) is a member of the Oncocyclus section, an elite group of 32 Iris species native to the Fertile Crescent region of southwestern Asia. They are some of the showiest irises on the planet. Sadly, like many others in this section, the Golan iris is in real danger of going extinct.

The Golan iris has a rather limited distribution. Despite being named in honor of Mt. Hermon, it is restricted to the Golan Heights region of northern Israel and southwestern Syria. Part of the confusion stems from the fact that the Golan iris has suffered from a bit of taxonomic uncertainty ever since it was discovered. It is similar in appearance to both I. westii and I. bismarckiana with which it is frequently confused. In fact, some authors still consider I. hermona to be a variety of I. bismarckiana. This has led to some serious issues when trying to assess population numbers. Despite the confusion, there are some important anatomical differences between these plants, including the morphology of their rhizomes and the development of their leaves. Regardless, if these plants are in fact different species, it means their respective numbers in the wild decrease dramatically. 

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Like other members of the Oncocyclus group, the Golan iris exhibits an intriguing pollination syndrome with a group of bees in the genus Eucera. Their large, showy flowers may look like a boon for pollinators, however, close observation tells a different story. The Golan iris and its relatives receive surprisingly little attention from most of the potential pollinators in this region.

One reason for their lack of popularity has to do with the rewards (or lack thereof) they offer potential visitors. These irises produce no nectar and very little pollen. Because of this and their showy appearance, most pollinators quickly learn that these plants are not worth the effort. Instead, the only insects that ever pay these large blossoms any attention are male Eucerine bees. These bees aren't looking for food or fragrance, however. Instead, they are looking for a place to rest. 

 A Eucerine bee visiting a nectar source. 

A Eucerine bee visiting a nectar source. 

The Oncocyclus irises cannot self pollinate, which makes studying potential pollinators a bit easier. During a 5 year period, researchers noted that male Eucerine bees were the only insects that regularly visited the flowers and only after their visits did the plants set seed. The bees would arrive at the flowers around dusk and poke around until they found one to their liking. At that point they would crawl down into the floral tube and would not leave again until morning. The anatomy of the flower is such that the bees inevitably contact stamen and stigma in the process. Their resting behavior is repeated night after night until the end of the flowering season and in this way pollination is achieved. Researchers now believe that the Golan iris and its relatives are pollinated solely by these sleeping male bees.

Sadly, the status of the Golan iris is rather bleak. As recent as the year 2000, there were an estimated 2,000 Golan irises in the wild. Today that number has been reduced to a meager 350 individuals. Though there is no single smoking gun to explain this precipitous decline, climate change, cattle grazing, poaching, and military activity have exacted a serious toll on this species. Plants are especially vulnerable during drought years. Individuals stressed by the lack of water succumb to increased pressure from insects and other pests. Vineyards have seen an uptick in Golan in recent years as well, gobbling up viable habitat in the process.

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It is extremely tragic to note that some of the largest remaining populations of Golan irises can be found growing in active mine fields. It would seem that one of the only safe places for these endangered plants to grow are places that are extremely lethal to humans. It would seem that our propensity for violent tribalism has unwittingly led to the preservation of this species for the time being.

At the very least, some work is being done not only to understand what these plants need in order to germinate and survive, but also assess the viability of relocated plants that are threatened by human development. Attempts at transplanting individuals in the past have been met with limited success but thankfully the Oncocyclus irises have caught the eye of bulb growers around the world. By sharing information on the needs of these plants in cultivation, growers can help expand on efforts to save species like the Golan iris.

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

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

 

The Carnivorous Waterwheel

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Bladderworts (Utricularia spp.) aren't the only carnivorous plants stalking prey below the water surface. Meet the waterwheel (Aldrovanda vesiculosa). At first glance it looks rather unassuming but closer inspection will reveal that this carnivore is well equipped for capturing unsuspecting prey. 

The waterwheel never bothers with roots. Instead, it lives out its life as a free floating sprig, its stem it covered in whorls of filamentous leaves, each tipped with a tiny trap. The trapping mechanism is a bit different from its bladderwort neighbors. Instead of bladders, the waterwheel produces snap traps that closely resemble those of the Venus fly trap (Dionaea muscipula). These traps function in a similar way. When zooplankton or even a small fish trigger the bristles along the rim, the trap snaps shut and begins the digestion process.

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This similarity to the Venus fly trap is more than superficial. DNA analysis reveals that they are in fact close cousins. Together with the sundews, these plants make up the family Droseraceae. The evolutionary history of this clade is a bit confusing thanks to a limited fossil record. Today, the waterwheel is the only extant member of the genus Aldrovanda but fossilized seeds and pollen reveal that this group was once a bit more diverse during the Eocene. Whenever these genera diverged, it happened a long time ago and little evidence of it remains.

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At one point in time, the waterwheel could be found growing in wetland habitats throughout Africa, Europe, Asia, and even Australia. Today it is considered at risk of extinction. Its numbers have been severely reduced thanks to wetland degradation and destruction. Of the 379 known historical populations, only about 50 remain in tact today and many of these are in rough shape. Agricultural and industrial runoff are exacting a significant toll on its long term survival. To make matters worse, sexual reproduction in the waterwheel is a rare event. Most often this plant reproduces vegetatively, reducing genetic diversity. What's more, natural dispersal into new habitats is extremely limited. 

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Oddly enough, populations of this plant have popped up in a few locations in eastern North America. These introductions were not a mistake either. Carnivorous plant enthusiasts concerned with the plight of this species in its native habitat began introducing it into water ways in New Jersey, New York, and Virginia where it is now established. Oddly enough, these introductions have performed far better than any of the reintroduction attempts made in its native range in Europe. Of course, this is always cause for concern. Endangered or not, the introduction of a species into new habitat is always risky. Still, there is hope yet for this species. Its popularity among plant growers has led to an increase in numbers in cultivation. At least folks have learned how to cultivate it until more comprehensive and effective conservation measures can be put into place. 

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

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

Parasitic Plant Rediscovered After a 151 Year Absence

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Extinction is a hard status to confirm for many types organisms. Whereas discovering a new species requires finding only a single individual, declaring one extinct requires knowing that there are no individuals left at all. This is especially true when organisms live cryptic lifestyles, a point recently made quite apparent by the rediscovery of a small parasitic plant known scientifically ask Thismia neptunis.

Thismia neptunis is a type of parasite called a mycoheterotroph, which means it makes its living by parasitizing mycorrhizal fungi in the soil. It obtains all of its needs in this way. As such, it produces no leaves, no chlorophyll, and really nothing that would readily identify it outright as a plant. All one would ever see of this species are its bizarre flowers that look more like a sea anemone than anything botanical. Like most mycoheterotrophs, when not in flower it lives a subterranean lifestyle.

 The original drawing of  Thismia neptunis  (from Beccari 1878).

The original drawing of Thismia neptunis (from Beccari 1878).

This is why finding them can be so difficult. Even when you know where they are supposed to grow, infrequent flowering events can make assessing population numbers extremely difficult. Add to this the fact that Thismia neptunis is only known from a small region of Borneo near Sarawak where it grows in the dense understory of hyperdiverse Dipterocarp forests. It was first found and described back in 1866 but was not seen again for 151 years. To be honest, it is hard to say whether or not most folks were actively searching.

Regardless, after a 151 year absence, a team of botanists recently rediscovered this wonderful little parasite flowering not too far from where it was originally described. Though more study will be needed to flesh out the ecology of this tiny parasitic plant, the team was fortunate enough to witness a few tiny flies flitting around within the flower tube. It could very well be that these odd flowers are pollinated by tiny flies that frequent these shaded forest understories.

As exciting as this rediscovery is, it nonetheless underscores the importance of forest conservation. The fact that no one had seen this plant in over a century speaks volumes about how little we understand the diversity of such biodiverse regions. The rate at which such forests are being cleared means that we are undoubtedly losing countless species that we don't even know exist. Forest conservation is a must. 

Click here to support forest conservation efforts in Borneo. 

Photo Credit and Further Reading: [1]

The Giant Genomes of Geophytes

 Canopy plant ( Paris japonica )

Canopy plant (Paris japonica)

A geophyte is any plant with a short, seasonal lifestyle and some form of underground storage organ ( bulb, tuber, thick rhizome, etc.). Plants hailing from a variety of families fall into this category. However, they share more than just a similar life history. A disproportionate amount of geophytic plants also possess massive genomes. 

As we have discussed in previous posts, life isn't easy for geophytes. Cold temperatures, a short growing season, and plenty of hungry herbivores represent countless hurdles that must be overcome. That is why many geophytes opt for rapid growth as soon as conditions are right. However, they don't do this via rapid cell division. 

 Dutchman's breeches ( Dicentra cucullaria ) emerging with preformed buds.

Dutchman's breeches (Dicentra cucullaria) emerging with preformed buds.

Instead, geophytes spend the "dormant" months pre-growing all of their organs. What's more, the cells that make up their leaves and flowers are generally much larger than cells found in non-geophytes. This is where that large genome comes into plant. If they had to wait until the first few weeks of spring to start their development, a large genome would only get in the way. Their dormant season growth means that these plants don't have to worry about streamlining the process of cellular division. They can take their time. 

As such, an accumulation of genetic material isn't detrimental. Instead, it may actually be quite beneficial for geophytes. Associated with large genomes are things like larger stomata, which helps these plants better regulate their water needs. The large genomes may very well be the reason that many geophytic plants are so good at taking advantage of such ephemeral growing conditions. 

When the right conditions present themselves, geophytes don't waste time. Pre-formed organs like leaves and flowers simply have to fill with water instead of having to wait for tissues to divide and differentiate. Water is plentiful during the spring so geophytes can rely on turgor pressure within their large cells for stability rather than investing in thick cell walls. That is why so many spring blooming plants feel so fleshy to the touch. 

Taken together, we can see how large genomes and a unique growth strategy have allowed these plants to exploit seasonally available habitats. It is worth noting, however, that this is far from the complete picture. With such a wide variety of plant species adopting a geophytic lifestyle, we still have a lot to learn about the secret lives of these plants.

Photo Credits: [1] [2]

Further Reading: [1]

An Ancient Hawaiian Moss

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The cloud forests of Kohala Mountain on the island of Hawai'i are home to a unique  botanical community. One plant in particular is quite special as it may be one of the most ancient clonal organisms in existence. Look down at your feet and you may find yourself surrounded by a species of moss known as Sphagnum palustre. Although this species enjoys a broad distribution throughout the northern hemisphere, its presence on this remote volcanic island is worth closer inspection. 

Hawai'i is rather depauperate in Sphagnum representatives and those that have managed to get to this archipelago are often restricted to growing in narrow habitable zones between 900 to 1,900 meters in elevation as these are the only spots that are cool and wet enough to support Sphagnum growth. Needless to say, successful colonization of the Hawaiian Islands by Sphagnum has been a rare event.  The fact that Sphagnum palustre was one of the few that did should not come as any surprise. What should surprise you, however, is how this particular species has managed to persist. 

 Mounds of  S. palustre  in its native habitat. 

Mounds of S. palustre in its native habitat. 

Hawaiian moss aficionados have long noted that the entire population of Kohala's S. palustre mats never seem to produce a single female individual. Indeed, this moss is dioicous, meaning individuals are either male or female. As such, many have suspected that the mats of S. palustre growing on Kohala represented a single male individual that has been growing vegetatively ever since it arrived as a spore on the island. The question then becomes, how long has this S. palustre individual been on Kohala?

To answer that, researchers decided to take a look at its DNA. What they discovered was surprising in many ways. For starters, all plants were in fact males of a single individual. A rare genetic trait was found in the DNA of every population they sampled. This trait is so rare that the odds of it turning up in any number by sheer chance is infinitesimally small. What this means is that every S. palustre population found on Kohala is a clone of a single spore that landed on the mountain at some point in the distant past. Exactly how distant was the next question the team wanted to answer. 

 A lush cloud forest on the slopes of Kohala.

A lush cloud forest on the slopes of Kohala.

The first clue to this mystery came from peat deposits found on the slopes of the mountain. Researchers found remains of S. palustre in peat deposits that were dated to somewhere around 24,000 years old. So, it would appear that S. palustre has been growing on Kohala since at least the late Pleistocene. But how long before that time did this moss arrive?

Again, DNA was the key to unlocking this mystery. By studying the rate at which mutations arise and fix themselves within the genetic code of this plant, they were able to estimate the average rate of mutation through time. By sampling different moss populations on Kohala, they could then use those estimates to figure out just how long each mat has been growing. Their estimates suggest that the ancestral male sport arrived on Hawai'i somewhere between 49,000 and 50,000 years ago and it has been cloning itself ever since. 

 A large mat of  S. palustre

A large mat of S. palustre

As if that wasn't remarkable in and of itself, their thorough analysis of the genetic diversity within S. palustre revealed a remarkable amount of genetic diversity for a clonal organism. Though not all genetic mutations are beneficial, enough of them have managed to fix themselves into the DNA of the moss clones over thousands of years. The DNA of S. palustre is challenging long-held assumptions about genetic diversity of asexual organisms.

Of course, no conversation about Hawaiian botany would be complete without mention of invasive species. As one can expect at this point, Kohala's S. palustre populations are being crowded out by more aggressive vegetation introduced from elsewhere in the world. Unlike a lot of Hawaiian plants, however, the clonal habit of S. palustre puts a more nuanced twist to this story. 

Because Sphagnum is spongy yet durable, it has often been used as packing material. Packages stuffed with S. palustre from Kohala have been sent all over the island and because of this, S. palustre is now showing up en masse on other islands in the archipelago. Sadly, when it starts to grow in habitats that have never experienced the ecosystem engineering traits of a Sphagnum  moss, S. palustre gets pretty out of hand. It's not just packages that spread it either. All it takes is one sprig of the moss stuck on someone's boot to start a new colony elsewhere. The unique flora elsewhere in the Hawaiian archipelago have not evolved to compete with S. palustre and as a result, escaped populations are rapidly changing the ecology to the detriment of other endemic Hawaiian plants. 

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

Further Reading: [1] [2] 

The Pima Pineapple Cactus

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The Pima pineapple cactus (Coryphantha robustispina) is a federally endangered cactus native to the Sonoran Desert. It is a relatively small cactus by most standards, a fact that can make it hard to find even with a trained eye. Sadly, the plight of this cactus is shared by myriad other plant species of this arid region. Urbanization, fire, grazing, and illegal collection are an ever present threat thanks to our insatiable need to gobble up habitat we should never have occupied in the first place. 

Deserts are lands of extremes and the Pima pineapple cactus seems ready for whatever its habitat can throw its way (naturally). Plants are usually found growing individually but older specimens take on a clustered clonal habit. During the winter months, the Pima pineapple cactus shrivels and waits until warmth returns. Come spring, the Pima pineapple cactus begins anew. On mature specimens, flower buds begin to develop once the plant senses an increase in daylight. 

The flower buds continue to develop well into summer but seem to stop after a certain point. Then, with the onset of the summer monsoons, flower buds quickly mature and open all at once. It is thought that this evolved as a means of synchronizing reproductive events among widely spaced populations. You see, seed set in this species is best achieved via cross pollination. With such low numbers and a lot of empty space in between, these cacti must maximize the chances of cross pollination.

If they were to flower asynchronously, the chances of an insect finding its way to two different individuals is low. By flowering together in unison, the chances of cross pollination are greatly increased. No one is quite sure exactly how these cacti manage to coordinate these mass flowering events but one line of reasoning suggests that the onset of the monsoon has something to do with it. It is possible that as plants start to take up much needed water, this triggers the dormant flower buds to kick into high gear and finish their development. More work is needed to say for sure.

Seed dispersal for this species comes in the form of a species of hare called the antelope jackrabbit. Jackrabbits consume Pima fruits and disperse them across the landscape as they hop around. However, seed dispersal is just one part of the reproductive process. In order to germinate and survive, Pima pineapple cacti seeds need to end up in the right kind of habitat. Research has shown that the highest germination and survival rates occur only when there is enough water around to fuel those early months of growth. As such, years of drought can mean years of no reproduction for the Pima.

Taken together, it is no wonder then why the Pima pineapple cactus is in such bad shape. Populations can take years to recover if they even manage to at all. Sadly, humans have altered their habitat to such a degree that serious action will be needed to bring this species back from the brink of extinction. Aside from the usual suspects like habitat fragmentation and destruction, invasive species are playing a considerable role in the loss of Pima populations. 

Lehmann lovegrass (Eragrostis lehmanniana) was introduced to Arizona in the 1930's and it has since spread to cover huge swaths of land. What is most troubling about this grass is that it has significantly altered the fire regime of these desert ecosystems. Whereas there was once very little fuel for fires to burn through, dense stands of Lehmann lovegrass now offer plenty of stuff to burn. Huge, destructive fires can spread across the landscape and the native desert vegetation simply cannot handle the heat. Countless plants are killed by these burns.

Sometimes, if they are lucky, large cacti can resprout following a severe burn, however, all too often they do not. Entire populations can be killed by a single fire. What few plants remain are frequent targets of poaching. Cacti are quite a hit in the plant trade and sadly people will pay big money for rare specimens. The endangered status of the Pima pineapple cactus makes it a prized target for greedy collectors. 

The future of the Pima pineapple cactus is decidedly uncertain. Thankfully its placement on the endangered species list has afforded it a bit more attention from a conservation standpoint. Still, we know very little about this plant and more data are going to be needed if we are to develop sound conservation measures. This, my friends, is why land conservation is so important. Plants like the Pima pineapple cactus need places to grow. If we do not work harder on setting aside wild spaces, we will lose so much more than this species. 

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

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

California Bumblebee Decline Linked to Feral Honeybees

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Worldwide, pollinators are having a rough go of it. Humans have altered the landscape to such a degree that many species simply can't keep up. The proverbial poster child for pollinator issues is the honeybee (Apis mellifera). As a result, countless native pollinators get the short shrift when it comes to media attention. This isn't good because outside of intense industrial agriculture, native pollinators make up the bulk of pollination services. Similarly, honeybee fandom often overshadows any potential negative effects these introduced insects might be having on native pollinators.

Long term scientific investigations are starting to paint a more nuanced picture of the impact introduced honeybees are having on native ecosystems. For instance, research based out of California is finding that honeybees are playing a big role in the decline of native bumblebee populations. What's more, these negative impacts are only made worse in the light of climate change.

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For over 15 years, ecologist Dr. Diane Thompson has been studying bumblebee populations in central California. At no point during those early years did any of the bumblebee species she focuses on show signs of decline. In fact, they were quite common. Then, around the year 2000, feral honeybees started to establish themselves in the area. Honeybee colonies were becoming more and more numerous each and every year and that is when she started noticing changes in bumblebee behavior and numbers.

You see, honeybees are extremely successful foragers. They are generalists, which means they can visit a wide variety of flower types. As a result, they are extremely good at competing for floral resources compared to native bumblebees. Her results show that increases in the number of honeybee colonies caused not only a reduction in foraging among the native bumblebees, they also caused a reduction in bumblebee colony success. The native bumblebees simply weren't raising as many young as they were before honeybees entered the system.

 Decreased rainfall cause a decline in flower densities of  Scrophularia californica , a key resource for native bumblebees in this system.

Decreased rainfall cause a decline in flower densities of Scrophularia californica, a key resource for native bumblebees in this system.

Climate change is only making things worse. As drought years become not only more severe but also more intense, the amount of flowers available during the growing season also declines. With fewer flowers on the landscape, bumblebees and honeybees are forced into closer proximity for foraging and the clear winner in most foraging disputes are the tenacious honeybees. As such, bumblebees are chased off the already diminishing floral displays. By 2014, Dr. Thompson had quantified a significant decline in native bumblebee populations as a result.

It would be all too convenient to say that this research represents an isolated case. It does not. More and more research is finding that honeybees frequently out-compete native pollinators for resources such as food and nesting sites. Such effects are especially pronounced in rapidly changing ecosystems. Although honeybees are here to stay, it is important that we realize the impacts that these feral insects are having on our native ecosystems and begin to better appreciate and facilitate the services provided by our native pollinators. 

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

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

Are Algae Plants?

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I was nibbling on some nori the other day when a thought suddenly hit me. I don't know squat about algae. I know it comes in many shapes, sizes, and colors. I know it is that stuff that we used to throw at each other on the beach. I know that it photosynthesizes. That's about it. What are algae? Are they even plants?

The shortest answer I can give you is "it depends." The term algae is a bit nebulous in and of itself. In Latin, the word "alga" simply means "seaweed." Algae are paraphyletic, meaning they do not share a recent common ancestor with one another. In fact, without specification, algae may refer to entirely different kingdoms of life including Plantae (which is often divided in the broad sense, Archaeplastida and the narrow sense, Viridiplantae), Chromista, Protista, or Bacteria.

  Caulerpa racemosa , a beautiful green algae.

Caulerpa racemosa, a beautiful green algae.

Taxonomy being what it is, these groupings may differ depending on who you ask. The point I am trying to make here is that algae are quite diverse from an evolutionary standpoint. Even calling them seaweed is a bit misleading as many different species of algae can be found in fresh water as well as growing on land.

 Cyanobacteria are photosynthetic bacteria, not plants.

Cyanobacteria are photosynthetic bacteria, not plants.

Take for instance what is referred to as cyanobacteria. Known commonly as blue-green algae, colonies of these photosynthetic bacteria represent some of the earliest evidence of life in the fossil record. Remains of colonial blue-green algae have been found in rocks dating back more than 4 billion years. As a whole, these types of fossils represent nearly 7/8th of the history of life on this planet! However, they are considered bacteria, not plants.

 Diatoms (Chromista)

Diatoms (Chromista)

Diatoms (Chromista) are another enormously important group. The single celled, photosynthetic organisms are encased in beautiful glass shells that make up entire layers of geologic strata. They comprise a majority of the phytoplankton in the world's oceans and are important indicators of climate. However, they belong to their own kingdom of life - Chromista or the brown algae.

To bring it back to what constitutes true plants, there is one group of algae that really started it all. It is widely believed that land plants share a close evolutionary history with a branch of green algae known as the stoneworts (order Charales). These aquatic, multicellular algae superficially resemble plants with their stalked appearance and radial leaflets.

 A nice example of a stonewort ( Chara braunii ).

A nice example of a stonewort (Chara braunii).

It is likely that land plants evolved from a Chara-like ancestor that may have resembling modern day hornworts that lived in shallow freshwater inlets. Estimates of when this happen go back as far as 500 million years before present. Unfortunately, fossil evidence is sparse for this sort of thing and mostly comes in the form of fossilized spores and molecular clock calculations.

  Porphyra umbilicalis   - One of the many species of red algae frequently referred to as nori.

Porphyra umbilicalis  - One of the many species of red algae frequently referred to as nori.

Now, to bring it back to what started me down this road in the first place. Nori is made from algae in the genus Porphyra, which is a type of Rhodophyta or red algae. Together with Chlorophyta (the green algae), they make up some of the most familiar groups of algae. They have also been the source of a lot of taxonomic debate. Recent phylogenetic analyses place the red algae as a sister group to all other plants starting with green algae. However, some authors prefer to take a broader look at the tree and thus lump red algae in a member of the plant kingdom. So, depending on the particular paper I am reading, the nori I am currently digesting may or may not be considered a plant in the strictest sense of the word. That being said, the lines are a bit blurry and frankly I don't really care as long as it tastes good.

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

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

 

How a Giant Parasitic Orchid Makes a Living

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

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

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

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

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

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

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

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

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

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Photo Credits: [1] [2]

Further Reading: [1]

How Air Plants Drink

   Tillandsia tectorum

 Tillandsia tectorum

Air plants (genus Tillandsia) are remarkable organisms. All it takes is seeing one in person to understand why they have achieved rock start status in the horticulture trade. Unlike what we think of as a "traditional" plant lifestyle, most species of air plants live a life free of soil. Instead, they attach themselves to the limbs and trunks of trees as well as a plethora of other surfaces. 

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Living this way imposes some serious challenges. The biggest of these is the acquisition of water. Although air plants are fully capable of developing roots, these organs don't live very long and they are largely incapable of absorbing anything from the surrounding environment. The sole purpose of air plant roots is to anchor them to whatever they are growing on. How then do these plants function? How do they obtain water and nutrients? The answer to this lies in tiny structures called trichomes. 

Trichomes are what gives most air plants their silvery sheen. To fully appreciate how these marvelous structures work, one needs some serious magnification. A close inspection would reveal hollow, nail-shaped structures attached to the plant by a stem. Instead of absorbing water directly through the leaf tissues, these trichomes mediate the process and, in doing so, prevent the plant from losing more water than it gains. 

The trichomes themselves start off as living tissue. During development, however, they undergo programmed cell death, leaving them hollow. When any amount of moisture comes into contact with these trichomes, they immediately absorb that water, swelling up in the process. As they swell, they are stretched out flat along the surface of the leaf. This creates a tiny film of water between the trichomes and the rest of the leaf, which only facilitates the absorption of more water. 

 Trichomes up close.  

Trichomes up close.  

Because the trichomes form a sort of conduit to the inside of the leaf, water and any nutrients dissolved within are free to move into the plant until the reach the spongy mesophyll cells inside. In this way, air plants get all of their water needs from precipitation and fog. Not all air plants have the same amount of trichomes either. In fact, trichome density can tell you a lot about the kind of environment a particular air plant calls home. 

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The fuzzier the plant looks, the drier the habitat it can tolerate. Take, for instance, one of the fuzziest air plants - Tillandsia tectorum. This species hails from extremely arid environments in the high elevation regions of Ecuador and Peru. This species mainly relies on passing clouds and fog for its moisture needs and thus requires lots of surface area to collect said water. Now contrast that with a species like Tillandsia bulbosa, which appears to have almost no trichome cover. This smoother looking species is native to humid low-land habitats where high humidity and frequent rain provide plenty of opportunities for a drink. 

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Absorbing water in this way would appear to have opened up a plethora of habitats for the genus Tillandsia. Air plants are tenacious plants and worthy of our admiration. One could learn a lot from their water savvy ways. 

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

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