A New Case of Lizard Pollination from South Africa


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

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

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

The hidden flower ( Guthriea capensis )  in situ .

The hidden flower (Guthriea capensis) in situ.

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

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

Pseudocordylus melanotus

Pseudocordylus melanotus

Tropidosaura gularis

Tropidosaura gularis

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

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

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

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

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

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

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

Further Reading: [1] [2]

The Creeping Strawberry Pine


With its small, creeping habit and bright red, fleshy female cones, it is easy to see how Microcachrys tetragona earned its common name “creeping strawberry pine.” This miniature conifer is as adorable as it is interesting. With a fossil history that spans 66 million years of Earth’s history, it also has a lot to teach us about biogeography.

Today, the creeping strawberry pine can only be found growing naturally in western Tasmania. It is an alpine species, growing best in what is commonly referred to as alpine dwarf scrubland, above 1000 m (3280 ft) in elevation. Like the rest of the plants in such habitats, the creeping strawberry pine does not grow very tall at all. Instead, it creeps along the ground with its prostrate branches that barely extend more than 30 cm (0.9 ft) above the soil. This, of course, is likely an adaptation to its alpine environment. Plants that grow too tall frequently get knocked back by brutal winds and freezing temperatures among other things.

The typical growth habit of the creeping strawberry pine.

The typical growth habit of the creeping strawberry pine.

The creeping strawberry pine is not a member of the pine family (Pinaceae) but rather the podocarp family (Podocarpaceae). This family is interesting for a lot of reasons but one of the coolest is the fact that they are charismatic representatives of the so-called Antarctic flora. Along with a handful of other plant lineages, it is thought that Podocarpaceae arose during a time when most of the southern continents were combined into a supercontinent called Gondwana. Subsequent tectonic drift has seen the surviving members of this flora largely divided among the continents of the Southern Hemisphere. By combining current day distributions with fossil evidence, researchers are able to use families such as Podocarpaceae to tell a clearer picture of the history of life on Earth.

What is remarkable is that among the various podocarps, the genus Microcachrys produces pollen with a unique morphology. When researchers look at pollen under the microscope, whether extant or fossilized, they can say with certainty if it belongs to a Microcachrys or not. The picture we get from fossil evidence paints an interesting picture for Microcachrys diversity compared to what we see today. It turns out, Microcachrys endemic status is a more recent occurrence.

This distinctive, small, trisaccate pollen grain is typical of what you find with  Microcachrys  whereas all other podocarps produce bisaccate pollen.

This distinctive, small, trisaccate pollen grain is typical of what you find with Microcachrys whereas all other podocarps produce bisaccate pollen.

The creeping strawberry pine is what we call a peloendemic, meaning it belongs to a lineage that was once far more widespread but today exists in a relatively small geographic location. Fossilized pollen from Microcachrys has been found across the Southern Hemisphere, from South America, India, southern Africa, and even Antarctica. It would appear that as the continents continued to separate and environmental conditions changed, the mountains of Tasmania offered a final refuge for the sole remaining species in this lineage.

Another reason this tiny conifer is so charming are its fruit-like female cones. As they mature, the scales around the cone swell and become fleshy. Over time, they start to resemble a strawberry more than anything a gymnosperm would produce. This is yet another case of convergent evolution on a seed dispersal mechanism among a gymnosperm lineage. Birds are thought to be the main seed dispersers of the creeping strawberry pine and those bright red cones certainly have what it takes to catch the eye of a hungry bird. It must be working well for it too. Despite how narrow its range is from a global perspective, the creeping strawberry pine is said to be locally abundant and does not face the same conservation issues that many other members of its family currently face. Also, its unique appearance has made it something of a horticultural curiosity, especially among those who like to dabble in rock gardening.

Mature female cones look more like angiosperm fruit than a conifer cone.

Mature female cones look more like angiosperm fruit than a conifer cone.

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

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

Meet Jones' Columbine

Meet Aquilegia jonesii. This interesting little columbine can be found growing in a narrow range along the northern Rockies. It only grows in alpine and sub-alpine zones, making it quite rare. It has a cushion-like growth form to shield it from the elements but disproportionately large flowers. It is a lucky day if one stumbles across this species! 

Fun Fact: Both the common name and generic name of the flowers referred to collectively as "columbines" have their origins in ornithology? 

That's right, the genus to which they belong, Aquilegia, can trace its origin to the word "aquila," which is Latin for "eagle." When the genus was being described, it was felt that the flower resembled the claw of an eagle. 

The word "columbine" has it's origins in the word "columba," which is Latin for "pigeon" or "dove." Early botanical enthusiasts felt that the nectar spurs resembled the heads of a group of doves. 

More and more I am coming on board with the idea that etymology can be quite fun.

Photo Credit: Steve (http://bit.ly/NbGbmz)

Further Reading: [1] [2] [3]

High Elevation Record Breakers Are Evidence of Climate Change

A new record has been set for vascular plants. Three mustards, two composits, and a grass have been found growing at an elevation of 20,177 feet (6,150 m) above sea level!

Mountains are a brutal place to live. Freezing temperatures, fierce winds, limited soil, and punishing UV radiation are serious hurdles for any form of life. Whereas algae and mosses can often eke out an existence at such altitudes, more derived forms of life have largely been excluded from such habitats. That is, until now. The area in which these plants were discovered measured about the size of a football field and is situated atop an Indian mountain known as Mount Shukule II.

Although stressed, these plants were nonetheless established among the scree of this menacing peak. Most were quite young, having only been there for a few seasons but growth rings on the roots of at least one plant indicated that it had been growing there for nearly 20 years!

All of them have taken the cushion-like growth habit of most high elevation plant species in order to reduce exposure and conserve water. The leaves of each species also contained high levels of sugary anti-freeze, a must in this bitter cold habitat.

The research team, who could only muster a few hours of work each day, believed that the seeds of these plants were blown up there by wind. Because soils in alpine zones are often non-existent, the team wanted to take a closer look at what kind of microbial community, if any, was associated with their roots.

Whereas no mycorrhizal species were identified, the team did find a complex community of bacteria living among the roots that are characteristic of species living in arid, desert-like regions. It is likely that these bacteria came in with the seeds. Aside from wind, sun, and a lack of soil, one of the other great challenges for these plants is a short growing season. In order to persist at this elevation, the plants require a minimum of 40 days of frost-free soil each year.

Because climate change is happening much faster in mountainous regions, it is likely that such favorable growing conditions are a relatively recent phenomenon. The area in question has only recently become deglaciated. As average yearly temperatures continue to increase, the habitable zone for plants such as these is also moving up the mountain. The question is, what happens when it reaches the top? Once at the peak, plants have nowhere to go. One of the greatest issues alpine plants face is that they will gradually be squeezed off of these habitat islands.

Although expanding habitable zones in these mountains may sound like a good thing, it is likely a short term benefit for most species. Whereas temperature bands in the Tibetan mountains are moving upwards at a rate of 20 feet (6 m) per year, most alpine plants can only track favorable climates at a rate of about 2 inches (0.06 m) per year. In other words, they simply can't keep up. As such, this record breaking discovery is somewhat bitter sweet.

Photo Credit: [1]

Further Reading: [1]

Staying Warm: An Alpine Plant Approach to Reproduction

Things are beginning to cool down throughout the northern hemisphere. As winter approaches, most plant species begin to enter their dormancy period. Very few plants risk wasting their reproductive efforts in the chill of late fall, having gotten most of it out of the way during the warm summer months. This is easy enough for low elevation (and low latitude) plants but what about species living in the high arctic or alpine habitats. Such habitats are faced with cold, harsh conditions year round. How do plants living in these zones deal with reproduction?

These limitations are overcome via physiology. For starters, plants living in such extreme habitats often self pollinate. Insects and other pollinators are too few and far between to rely solely upon them as a means of reproduction. Also, the flowers of most cold weather plants are heliocentric. This means that, as the sun moves across the sky, the flowers track its path so that they are constantly perpendicular to its rays. This maintains maximum exposure to this precious heat source. 

Additionally, many arctic and alpine plants have parabolically shaped flowers. This amplifies the incoming radiation being absorbed by the flower. Experiments have shown that flowers that have been shaded from the heat of the sun had a dismal seed set of only 8% whereas plants exposed to the sun had an elevated seed set of 60%. 

For plants in these habitats, its all about persistence. Low reproductive rates are often offset by extremes in longevity. This is one of the many reasons why hikers must remember to tread lightly in these habitats. Damages incurred by even a single careless hiker can take decades, if not centuries, to recover. 

Photo Credit: [1]

Further Reading: [1]

Lizard Helpers

The beauty of Tasmania's honeybush, Richea scoparia, is equally matched by its hardiness. At home across alpine areas of this island, this stout Ericaceous shrub has to contend with cold temperatures and turbulent winds. The honeybush is superbly adapted to these conditions with its compact growth, and tough, pointy leaves. Even its flowers are primed for its environment. They emerge in dense spikes and are covered by a protective casing comprised of fused petals called a "calyptra." Such adaptations are great for protecting the plant and its valuable flowers from such brutal conditions but how does this plant manage pollination if its flowers are closed off to the rest of the world? The answer lies in a wonderful little lizard known as the snow skink (Niveoscincus microlepidotus).

The snow skink is not a pollinator. Far from it. All the snow skink wants is access to the energy rich nectar contained within the calyptra. In reality, the snow skink is a facilitator. You see, the calyptra may be very good at shielding the developing flower parts from harsh conditions, but it tends to get in the way of pollination. That is where the snow skink comes in. Attracted by the bright coloration and the nectar inside, the snow skink climbs up to the flower spike and starts eating the calyptra. In doing so, the plants reproductive structures are liberated from their protective sheath. 

Once removed, the flowers are visited by a wide array of insect pollinators. In fact, research shows that this is the only mechanism by which these plants can successfully outcross with their neighbors. Not only does the removal of the calyptra increase pollination for the honeybush, it also aids in seed dispersal. Experiments have shown that leaving the calyptra on resulted in no seed dispersal. The dried covering kept the seed capsules from opening. When calyptras are removed, upwards of 87% of seeds were released successfully. 

Although several lizard species have been identified as pollinators and seed dispersers, this is some of the first evidence of a reptilian pollination syndrome that doesn't actually involve a lizard in the act of pollination. It is kind of bizarre when you think about it. As if pollination wasn't strange enough in requiring a third party for sexual reproduction to occur, here is evidence of a fourth party required to facilitate the action in the first place. It may not be just snow skinks that are involved either. Evidence of birds removing the calyptra have also been documented. Whether its bird or lizard, this is nonetheless a fascinating coevolutionary relationship in response to cold alpine conditions. 

Photo Credits: [1] [2]

Further Reading: [1]

Darwin's Slipper

Along the craggy peaks of the Andes from Chile into Argentinia, and down into Patagonia grows a strange alpine plant known scientifically as Calceolaria uniflora.  It goes by the common name of Darwin's Slipper as many attribute its discovery to Charles Darwin, however, this plant was first collected by French naturalist Philibert Commerson in 1767, 42 years before Darwin was even born. Regardless, C. uniflora is a remarkable little plant. It stands as an ornate example of a unique pollination syndrome, one that that is quite apt considering who discovered it. As with any strange flower, once you begin to ponder the significance of its morphology, you inevitably come to the same question; what on Earth pollinates it?

As a whole the genus Calceolaria is bee pollinated. Relying on what are known as "oil bees," most of the flowers in this genus produce hairs that secret oils that the female bees relish. Calceolaria uniflora is different from the rest in that it doesn't bother with oil production. Instead of producing flowers with a tube or a pouch, this species creates an almost alien-looking red and orange bloom with a bright white appendage on its lower lip. What is going on there? The answer to this strange riddle has a clue in where this species grows.

At high altitudes, oil-collecting bees are scarce. It is simply too cold and harsh for many insects to survive at such elevations. Instead, what are present are birds, specifically a species of seedsnipe. These little birds exist on a plant-based diet and spend a lot of their time holding territories and grazing on seeds and fruits of a handful of alpine plants. Researchers noticed that patches of Calceolaria uniflora growing around these birds seemed to have high levels of floral damage, specifically on the lower lip where the white appendage is located. In fact, the white appendage was often completely removed.

As it turns out, the seedsnipes regularly visit patches of these flowers and proceed to peck off and eat the white appendage. As the birds peck off these appendages, the anthers and stigma bash against the birds head. As it does, pollen is dusted onto the bird as well as onto the female parts of the flower. Thus pollination is achieved. But what's in it for the birds? As it turns out, when tested in the lab, researchers found these appendages to be high in sugars. The birds are in it for an easy, sugary meal.

When we think of birds as pollinators, we often think of hummingbirds or honeyeaters. The relationship between Calceolaria uniflora and the seedsnipe is rather outlandish in comparison but it certainly works for both species. The lack of insect pollinators has driven Calceolaria uniflora towards an alternative pollinator and quite a unique one at that!

Photo Credit: Julio Martinich (http://bit.ly/1TMiqk7)

Further Reading: