Cycad Pollinators

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

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

White-haired cycad ( Encephalartos friderici-guilielm i)

White-haired cycad (Encephalartos friderici-guilielmi)

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

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

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

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

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

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

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

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

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

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

Macrozamia macleayi  female cone.

Macrozamia macleayi female cone.

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

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

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

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

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

Thrips up close.

Thrips up close.

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

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

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

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]

Everlasting or Seven Years Little

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Common names are a funny thing. Depending on the region, the use, and the culture, one plant can take on many names. In other situations, many different plants can take on a single name. Though it isn't always obvious to those unfamiliar with them, the use of scientific names alleviates these issues by standardizing the naming of things so that anyone, regardless of where they are, knows what they are referring to. That being said, sometimes common names can be entertaining.

Take for instance, plants in the genus Syncarpha. These stunning members of the family Asteraceae are endemic to the fynbos region of the Eastern and Western Cape of South Africa. In appearance they are impossible to miss. In growth habit they have been described as "woody shrublets," forming dense clusters of woody stems covered in a coat of woolly hairs. Sitting atop their meter-high stems are the flower heads.

Each flower head consists of rings of colorful paper-like bracts surrounding a dense cluster of disk flowers. The flowering period of the various species can last for weeks and spans from October, well into January. Numerous beetles can be observed visiting the flowers and often times mating as they feed on pollen. Some of the beetles can be hard to spot as they camouflage quite well atop the central disk. Some authors feel that such beetles are the main pollinators for many species within this genus.

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Their mesmerizing floral displays are where their English common name of "everlasting" comes from. Due to the fact that they maintain their shape and color for a long time after being cut and dried, various Syncarpha species have been used quite a bit in the cut flower industry. A name that suggests everlasting longevity stands in stark contrast to their other common name. 

These plants are referred to as "sewejaartjie" in Afrikaans, which roughly translates to "seven years little." Why would these plants be referred to as everlasting by some and relatively ephemeral by others? It turns out, sewejaartjie is a name that has more to do with their ecology than it does their use in the floral industry.

As a whole, the 29 described species of Syncarpha are considered fire ephemerals. The fynbos is known for its fire regime and the plants that call this region home have evolved in response to this fact. Syncarpha are no exception. They flower regularly and produce copious amounts of seed but rarely live for more than 7 years after germination. Also, they do not compete well with any vegetation that is capable of shading them out.

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Instead, Syncarpha invest heavily in seed banking. Seeds can lie dormant in the soil for many years until fires clear the landscape of competing vegetation and release valuable nutrients into the soil. Only then will the seeds germinate. As such, the mature plants don't bother trying to survive intense ground fires. They burn up along with their neighbors, leaving plenty of seed to usher in the next generation.

Research has shown that its not the heat so much as the smoke that breaks seed dormancy in these plants. In fact, numerous experiments using liquid smoke have demonstrated that the seeds are likely triggered by some bio-active chemical within the smoke itself.

So, there you have it. Roughly 29 plants with two common names, each referring back to an interesting aspect of the biology of these plants. Despite their familiarity and relative ease of committing to memory, the common names of various species only get us so far. That's not to say we should abolish the use of common names altogether. Learning about any plant should be an all encompassing endeavor provided you know which plant you are referring to.

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

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

 

Mighty Magnolias

Magnolias are one of those trees that even the non-botanically minded among us will easily recognize. They are one of the more popular plant groups grown as ornamentals and their symbolism throughout human history is quite interesting. But, for all this attention, few may realize how special magnolias really are. Did you know they they are one of the most ancient flowering plant lineages in existence?

Magnolias first came on to the scene somewhere around 95 million years ago. Although they are not representative of what the earliest flowering plants may have looked like, they do offer us some interesting insights into the evolution of flowers. To start with, the flower bud is enclosed in bracts (modified leaves) instead of more differentiated sepals. The "petals" themselves are not actually petals but tepals, which are also undifferentiated. The most striking aspect of magnolia flower morphology is in the actual reproductive structures themselves.

Magnolias evolved before there were bees. Because of this, the basic structure that makes them unique was in place long before bees could work as a selective pressure in pollination. Beetles are the real pollinators of magnolia flowers. The flowers have a hardened carpel to avoid damage by their gnawing mandibles as the feed. The beetles are after the protein-rich pollen. Because the beetles are interesting in pollen and pollen alone, the flowers mature in a way that ensures cross pollination. The male parts mature first and offer said pollen. The female parts of the flower are second to mature. They produce no reward for the beetles but are instead believed to mimic the male parts, ensuring that the beetles will spend some time exploring and thus effectively pollinating the flowers.

It is pretty neat to think that you don't necessarily have to track down a dawn redwood or a gingko to see a plant that has survived major extinction events. You can find magnolias very close to home with a keen eye. Looking at one, knowing that this is a piece of biology that has worked for millennia, is quite astounding in my opinion.

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