pollination

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The Hidden Flowers of Viola

Violas keep a secret hidden below their foliage. Sometimes they even bury it shallowly in the soil near their roots. I suppose it’s not a secret really, just something out of sight. There isn’t a reason to show it off, after all. Showy flowers are showy for the sole purpose of attracting pollinators. If pollinators are unnecessary, there is no reason for showy flowers, or to even show your flowers at all. That’s the story behind the cleistogamous flowers of violas. They are a secret only because unless you know to look for them, you would have no idea they were there at all.

Cleistogamy means closed marriage, and it describes a self-pollinating flower whose petals remain sealed shut. The opposite of cleistogamy is chasmogamy (open marriage). Most of the flowers we are familiar with are chasmogamous. They open and expose their sex parts in order to allow for cross-pollination (self-pollination can also occur in such flowers). Violas have chasmogamous flowers too. They are the familiar five-petaled flowers raised up on slender stalks above the green foliage. Cross-pollination occurs in these flowers, and seed-bearing fruits are the result. Perhaps as a way to ensure reproduction, violas also produce cleistogamous flowers, buried below their leaves.

an illustration of the cleistogamous flower of Viola sylvatica opened to reveal its sex partsĀ — via wikimedia commons

Flowers are expensive things to make, especially when the goal is to attract pollinators. Colorful petals, nectar, nutritious pollen, and other features that help advertise to potential pollinators all require significant resources. All this effort is worth it when it results in the ample production of viable seeds, but what if it doesn’t? Having a method for self-pollination ensures that reproduction will proceed in the absence of pollinators or in the event that floral visitors don’t get the job done. A downside, of course, is that a seed produced via self-pollination is essentially a clone of the parent plant. There will be no mixing of genes with other individuals. This isn’t necessarily bad, at least in the short term, but it has its downsides. A good strategy is a mixture of both cross- and self-pollination – a strategy that violas employ.

The cleistogamous flowers of violas generally appear in the summer or fall, after the chasmogamous flowers have done their thing. The fruits they form split open when mature and deposit their seeds directly below the parent plant. Some are also carried away by ants and dispersed to new locations. Seeds produced in these hidden flowers are generally superior and more abundant compared to those produced by their showy counterparts. People who find violas to be a troublesome lawn weed – expanding far and wide to the exclusion of turfgrass – have these hidden flowers to blame.

That being said, there is a defense for violas. In the book The Living Landscape by Rick Darke and Doug Tallamy, Tallamy writes: “Plants such as the common blue violet (Viola sororia), long dismissed by gardeners as a weed, can be reconstituted as desirable components of the herbaceous layer when their ecosystem functionality is re-evaluated. Violets are the sole larval food source for fritillary butterflies. Eliminating violets eliminates fritillaries, but finding ways to incorporate violets in garden design supports fritillaries.”

sweet violet (Viola odorata)

In my search for the cleistogamous flowers of viola, I dug up a sweet violet (Viola odorata). I was too late to catch it in bloom, but the product of its flowers – round, purple, fuzzy fruits – were revealed as I uprooted the plant. Some of the fruits were already opening, exposing shiny, light brown seeds with prominent, white elaiosomes, there to tempt ants into aiding in their dispersal. I may have missed getting to see what John Eastman calls “violet’s most important flowers,” but the product of these flowers was certainly worth the effort.

Fruits formed from the cleistogamous flowers of sweet violet (Viola odorata)

Up close and personal with the fruit of a cleistogamous flower

The seeds (elaiosomes included) produced by the cleistogamous flower of sweet violet (Viola odorata)

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Using Weeds: Soapwort

Over the past year or so I have written about several edible weeds in an effort to highlight useful weeds. However, weeds don’t have to be edible to be useful. In fact, many weeds are most certainly not edible, but that doesn’t mean they are of no use to humans. Soapwort, for example, is poisonous, and while it does have a history of being used internally as medicine, ingesting it is not advised and should only be done under the direction of a doctor. A much less risky activity would be to make soap out of it.

soapwort (Saponaria officinalis)

Saponaria officinalis, commonly known as bouncing bet, hedge pink, fuller’s herb, scourwort, and soapweed or soapwort, is an herbaceous perennial native to Europe. It has been planted widely in flower beds and herb gardens outside of its native range, desired both for its beauty and utility. Capitalizing on our appreciation for it, soapwort has expanded beyond our garden borders and into natural areas, as well as vacant lots, roadsides, and other neglected spaces. Even in a garden setting it can be a bit of a bully, especially if ignored for a season or two.

The stems of soapwort grow to about two feet tall, are unbranched, and sometimes tinged with pink, purple, or red. The leaves are oblong and oppositely-arranged, and their bases form prominent collars around the stems. Showy clusters of flowers are found atop the stems throughout the summer. Like other flowers in the pink family (Caryophyllaceae), they are cigar-shaped at the base and opened wide at the end, showing off 5 distinct petals with notches at their tips. The petals of soapwort flowers bend backwards, with their sex parts protruding outwards. In his description of the flowers, John Eastman remarks in The Book of Field and Roadside that “the reflexed petals surrounding the sexual organs give the impression of flagrant thrust; this is a gaudy, unshy flower.”

collared stem of soapwort (Saponaria officinalis)

The fragrant flowers are pink to white in color. They open in the evening and remain open for a few short days. In an individual flower, pollen matures and is mostly shed before the stigma is ready to accept it. This helps reduce the chance of self-pollination. Cross pollination occurs with the assistance of moths who visit the flowers at night, as well as bees and other flower-visiting insects that come along during the daytime. Soapwort fruits are oval capsules containing as many as 500 kidney-shaped seeds. Seeds aren’t essential to the plants spread though, as much of its colonization occurs via vigorous rhizomes.

In fact, vegetative reproduction is the means by which soapwort forms such expansive, thick patches. It also helps that it’s poisonous. The saponins – its soap making compounds – that it produces in its roots, shoots, and leaves deter most insects and other animals from eating it. It has a reputation for poisoning horses, cows, and other livestock, and so is unwelcome in pastures and rangelands. Saponins are also poisonous to fish, so growing soapwort near fish ponds is not advised.

soapwort (Saponaria officinalis)

Soapwort occurs in a variety of soils including sandy, dry, and rocky sites and is surprisingly drough-tolerant, fine qualities to have when colonizing neglected sites. While most other organisms ignore soapwort, it has a friend in humans. Eastman sums this up well: “Soapwort’s most important associate – as is true of most plants we label weeds – is undoubtedly humankind, without whose helpful interventions the plant would surely be much rarer than it is.”

I made a soapy liquid out of soapwort by following a recipe that can be found on various blogs and websites by searching “saponaria soap recipe.” Basically it’s a cup of fresh leaves and stems along with a cup of dried leaves and stems added to a quart of distilled water brought to a boil. After simmering for 15 minutes and then allowing it to cool, strain the mixture through cheese cloth, and it’s ready to go.

This gentle but effective soap can be used for cleaning countertops and other surfaces, as well as dishes, fabrics, and skin. Several sources say it is particularly useful for cleaning delicate fabrics. Sierra and I both found it to have a cooked cabbage or spinach scent to it. This can be masked by adding a few drops of essential oil. Despite its odd aroma, both Sierra and I were impressed by its cleansing power and plan to use it more often.

dried leaves of soapwort

soapwort soap

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Inside of a Seed: Gymnosperms

“Every tree has to stay where it put down roots as a seedling. However, it can reproduce, and in that brief moment when tree embryos are still packed into seeds, they are free. The moment they fall from the tree, the journey can begin.” — The Hidden Life of Trees by Peter Wohlleben

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Seed plants – also known as spermatophytes – make up the largest group of plants on earth.Ā Seed plants consist ofĀ five divisions, and among them the angiosperm division (a.k.a. flowering plants) dominates in its number of species. The four remaining divisions are referred to collectively as gymnosperms. This incudes the cycads (Cycadophyta), Ginkgo biloba (the only living species in the division Ginkgophyta), gnetophytes (Gnetophyta), and the conifers (Coniferophyta). Conifers are by far the largest and most widespread gymnosperm division.

Angiosperms and gymnosperms have different evolutionary histories, resulting in their distinct genetic and morphological differences. That being said, an overly-simplistic way of differentiating the two groups is to say that, while both groups produce seeds, angiosperms produce flowers and fruits while gymnosperms produce pollen cones and seed cones. There are always exceptions (Ginkgo biloba, for example, doesn’t produce cones), but for the most part, this is the case.

Pollen cones (top) and seed cones (bottom) of mugo pine (Pinus mugo) via wikimedia commons

Sexual reproduction in gymnosperms follows a familiar pattern. Pollen, which contains the male sex cells, is produced in pollen cones, which are essentially miniature branches with modified leaves called scalesĀ that house the male reproductive organs. Mature pollen is shed and carried away by the wind. Lucky pollen grains make their way to the female cones, which are also modified branchlets, but are a bit more complex. Scales sit atop bracts, and on top of the scales are ovules – the female reproductive structures. During fertilization, the bracts open to collect pollen and then close as the seed develops.

When pollen lands onĀ an ovule it formsĀ pollen tubes that help direct the male sex cells to the egg cells inside. The process is similar to pollen tubes extending downĀ the style of a flower. In flowering plants, additional pollen cells combine with cells in the ovule to produce endosperm, a storage tissue that feeds the growing embryo. This doesn’t happen in gymnosperms.Ā Instead, haploid cells within the ovuleĀ develop intoĀ storageĀ tissue and go on to serve the same role.

The ovule eventually matures into a seed, and the cone opens to release it. The seed sits atop the scale rather than enclosed within a fruit, as it would be in an angiosperm. For this reason gymnosperms are said to have naked seeds. The development of seedsĀ can also be much slower in gymnosperms compared to angiosperms.Ā In some species, seeds don’t reach maturity for as long as two years.

Seed cones and winged seeds of mugo pine (Pinus mugo) via wikimedia commons

Seeds in the genus PinusĀ are excellent representations of typical gymnosperm seeds. Their basic components are essentially identical to the seeds of angiosperms. The seed coat is also referred to as an integument. It was once the outer covering of the ovule and has developed into the seed covering. A micropyle is sometimes visible on the seed andĀ is the location where theĀ pollen cellsĀ entered the ovule.Ā The storage tissue, as mentioned above,Ā is composed ofĀ female haploid cells that matured into storage tissue in the ovule.Ā Like angiosperms, theĀ embryo is composed of the radicle (embryonic root), the hypocotyl (embryonic shoot), and cotyledons (embryonicĀ leaves).

Angiosperms can be divided into monocotyledons and dicotyledons according to theĀ number of cotyledons their embryosĀ have (monocots have one, dicots have two). Gymnosperms are considered multi-cotyledonous because, depending on the species,Ā they can have a few to many cotyledons.

Seedling of Swiss pine (Pinus cembra) showing multiple cotyledons via wikimedia commons

For the sake of this introduction to gymnosperm seeds, I have offeredĀ a simple overview of the production of seeds in the conifer division.Ā Sexual reproduction and seed formation in the otherĀ three gymnosperm divisionsĀ is a similar story but varies according to species. Even within the conifers there are differences. ForĀ example, theĀ “seed cones” of severalĀ gymnosperm species can actually beĀ quite fruit-like,Ā which serves to attract animals toĀ aid inĀ seed dispersal.Ā Also, the pollen of gymnosperms is often thought ofĀ asĀ beingĀ wind dispersedĀ (andĀ occasionally water dispersed in the case of Ginkgo biloba and some cycads); however, researchers are continuing to discoverĀ the pivotalĀ role thatĀ insects play in theĀ transfer of pollen for manyĀ cycad species, just as theyĀ doĀ for so many species of angiosperms.

All of this to say thatĀ Botany 101 isĀ simply aĀ window into what is undoubtedly an incredibly diverse and endlessly fascinating group of organisms, and that, as with all branches of science, there is still so much to discover.

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Botany in Popular Culture: The Tan Hua Flowers in Crazy Rich Asians

When a flower blooms, a celebration is in order. Flowers abound for much of the year, which means parties are called for pretty much non-stop (something Andrew W.K. would surely endorse). Since we can’t possibly celebrate every bloom, there are certain plants we have decided to pay more attention to – plants whose flowers aren’t so prolific, predictable, or long-lived; or plants whose flowers come infrequently or at odd times of the day (or night).

This is the case with the flowers of the night blooming cactus,Ā Epiphyllum oxypetalum, which goes by many names including Dutchman’s pipe cactus, queen of the night, orchid cactus, night blooming cereus, and tan hua. Tan hua is the Chinese name for the plant, and this is how it is referred to in the book,Ā Crazy Rich Asians by Kevin Kwan.

In the book, Nick Young brings his American girlfriend, Rachel Chu, to meet his ridiculously wealthy family in Singapore. Before the trip, Rachel was in the dark about the Young’s wealth. She first meets the family and their gargantuan mansion when Nick’s grandma, seeing that her tan hua flowers are about to bloom, throws an impromptu (and lavish) party. Nick refers to the flowers as “very rare,” blooming “extremely infrequently,” and “quite something to witness.”

In a seperate conversation, Nick’s cousin, Astrid, tries to convince her husband to attend the party by claiming, “it’s awfully good luck to see the flowers bloom.” Later, another one of Nick’s cousins tells Rachel, “it’s considered to be very auspicious to witness tan huas blooming.”

Tan hua (Epiphyllum oxypetalum) via wikimedia commons

Native to Mexico and Guatemala, E. oxypetalumĀ was first brought to China in the 1600’s. Its beauty and intrigue along with its relative ease of cultivation helped it become popular and widespread across Asia and other parts of the world. Watching it bloom is considered a sacred experience by many, including in India, where it is said to bring luck and prosperity to households who are fortunate enough to see theirs bloom.

Epiphyllums are epiphytic, meaning they grow non-parasitically on the surfaces of other plants, such as in the crevices of bark or the crotches of branches. Like other cacti, they are essentially leafless, but their stems are broad, flat, and leaf-like in appearance. Showy, fragrant flowers are born along the margins of stems. The flowers of tan hua, as described in Crazy Rich Asians, appear as “pale reddish petals curled tightly like delicate fingers grasping a silken white peach.” A report (accompanied by photos) published by Sacred Heart University describes watching tan hua flowers progess from bud formation to full bloom, a process that took more than two weeks.

Tan huas are certainly not rare, as Nick described them. A number ofĀ Epiphyllum species and their hybrids are commonly cultivated; there is even an Epiphyllum Trail at San Diego Zoo’s Safari Park. Listed as “least concern” on the IUCN Red List, their popularity as ornamentals is noted but is not seen as affecting wild populations.Ā Night blooming plants,Ā while fascinating, aren’t all that rare either. Such plants have adapted relationships with creatures, like bats and moths, that are active during the night, employing their assistance with pollination. A paper published in Plant Systematics and Evolution describes the floral characteristics of Epiphyllum and similar genera: “The hawkmoth-flower syndrome, consisting of strongly-scented, night-blooming flowers with white or whitish perianths and long slender nectar-containing floral tubes is present in Cereus, Trichocereus, Selenicereus, Discocactus, Epiphyllum, and a number of other cactus genera.”

That being said, the specialness of a short-lived, infrequent, night blooming flower should not be understated, and really, parties being thrown in honor of any plant are something I can certainly get behind. Sitting in the courtyard late at night, the Young family and their guests watched as “the tightly rolled petals of the tan huas unfurled like a slow-motion movie to reveal a profusion of feathery white petals that kept expanding into an explosive sunburst pattern.” The look of it reminds Astrid of “a swan ruffling its wings, about to take flight.”

Later, “the tan huas began to wilt just as swiftly and mysteriously as they had bloomed, filling the night air with an intoxicating scent as they shriveled into spent lifeless petals.”

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Additional Resources:

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*Thank you Kathy for letting me borrow your Kindle so that I could write this post.

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Death by Crab Spider, part two

Crab spiders that hunt in flowers prey on pollinating insects. Thus, pollinating insects tend to avoid flowers that harbor crab spiders. We established this in part one. Now we ask, what effect, if any, does this interaction have on a crab spider infested plant’s ability to reproduce? More importantly, what are the evolutionary implications of this relationship?

In a study published in Ecological Entomology earlier this year, Gavini, et al. found that pollinating insects avoided the flowers of Peruvian lily (Alstroemeria aurea) when artificial spiders of various colors and sizes were placed in them. Bumblebees and other bees were the most frequent visitors to the flowers and were also the group “most affected by the presence of artificial spiders, decreasing the number of flowers visited and time spent in the inflorescences.” This avoidance had a notable effect on plant reproduction, namely a 25% reduction in seed set and a 15% reduction in fruit weight. The most abundant and effective pollinator, the buff-tailed bumblebee, was deterred by the spiders, leading the researchers to conclude that, “changes in pollinator behavior may translate into changes in plant fitness when ambush predators alter the behavior of the most effective pollinators.”

Peruvian lily (Alstroemeria aurea) via wikimedia commons

But missing from this discussion is the fact that crab spiders don’t only eat pollinators. Any flower visiting insect may become a crab spider’s prey, and that includes florivores. In which case, crab spiders can benefit a plant, saving it from reproduction losses by eating insects that eat flowers.

In April of this year, Nature Communications published a study by Knauer, et al. that examined the trade-off that occurs when crab spiders are preying on both pollinators and florivores. Four populations of buckler-mustard (Biscutella laevigata ssp. laevigata) were selected for this study. Bees are buckler-mustard’s main pollinator, and in concurrence with other studies, they significantly avoided flowers when crab spiders were present.Ā  Knauer, et al. also determined that bees and crab spiders are attracted to the same floral scent compound,Ā Ī²-ocimene. This compound not only attracts pollinators, but is also emitted when plants experience herbivory, possibly to attract predators to come and prey on whatever is eating them.

buckler-mustard (Biscutella laevigata) via wikimedia commons

In this study, the predators called upon were crab spiders. Florivores had a notable impact on plants in this study, and the researchers found that when crab spiders were present, florivores were significantly reduced, thereby reducing their negative impact. They also noted that “crab spiders showed a significant preference for [florivore-infested] plants over control plants.”

And so it is, a plant’s floral scent compound attracts pollinators while simultaneously attracting the pollinator’s enemy, who is also called in to protect the flower from being eaten. Luckily, in this case, buckler-mustard is easily pollinated, so the loss of a few pollinators isn’t likely to have a strong negative effect on reproduction. As the authors write, “pollinators are usually abundant and the low number of ovules per flower makes a few pollen grains sufficient for a full seed set.”

crab spider on zinnia

But none of these studies are one size fits all. Predator-pollinator-plant interactions are still not well understood, and there is much to learn through future research. A meta-analysis published in the Journal of Animal Ecology in 2011 looked at the research that had been done up to that point. Included were a range of studies involving sit-and-wait predators (like crab spiders and lizards) as well as active hunters (like birds and ants) and the effects of predation on both pollinators and plant-eating insects. They concluded that where carnivores “disrupted plant-pollinator interactions, plant fitness was reduced by 17%,” but thanks to predation of herbivores, carnivores helped increase plant fitness by 51%. This suggests that carnivores, overall, have a net positive effect on plant fitness.

Many pollinating insects have an advantage over plant-eating insects because they move quickly from flower to flower and plant to plant, unlike many herbivores which move more slowly. This protects pollinators from predation and helps explain why plant-pollinator interactions are not disrupted as easily by carnivores. Additionally, as the authors note, “plants may be buffered against loss of pollination by attracting different types of pollinators, some of which are inaccessible to carnivores.”

But again, there is still so much to discover about these complex interactions. One way to gain a better understanding is to investigate the effects of predators on both pollinators and herbivores in the same study, since many of the papers included in the meta-analysis focused on only one or the other. As far as crab spiders go,Ā Knauer, et al. highlight their importance in such studies. There are so many different species of crab spiders, and they are commonly found on flowers around the globe, so “their impact on plant evolution may be widespread among angiosperms.”

In other words, while we still have a lot to learn, the impact these tiny but skillful hunters have should not be underestimated.

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Death by Crab Spider, part one

When a bee approaches a flower, it is essentially approaching the watering hole. It comes in search of food in the form of pollen and nectar. As is this case with other animals who come to feed at the watering hole, a flower-visiting bee makes itself vulnerable to a variety of predators. Carnivores, like the crab spider, lie in wait to attack.

The flowers of many plants rely on visits from bees and other organisms to assist in transferring pollen from stamens to stigmas, which initiates reproduction; and bees and other flower visitors need floral resources to survive. Crab spiders exploit this otherwise friendly relationship and, in doing so, can leave lasting impacts on both the bees and the flowers they visit.

Species in the family Thomisidae are commonly referred to as crab spiders, a name that comes from their resemblance to crabs. Crab spiders don’t build webs to catch prey; instead they either actively hunt for prey or sit and wait for potential prey to happen by, earning them the name ambush predators. Of the hundreds of species in this family, not all of them hunt for prey in flowers; those that do – species in the genera Misumena and Thomisus, for example – are often called flower crab spiders.

white crab spider (Thomisus spectabilis) on Iris sanguineaĀ — via wikimedia commons

Most crab spiders are tiny – mere millimeters in size – and they have a number of strategies (depending on the species) to obscure their presence from potential prey. They can camouflage themselves by choosing to hunt in a flower that is the same color as they are or, in the case of some species, they can change their color to match the flower they are on. Some species of crab spiders reflect UV light, which bees can see. In doing so, they make themselves look like part of the flower.

Using an Australian species of crab spider, researchers found that honey bees preferred marguerite daisies (Chrysanthemum frutescens) on which UV-reflecting crab spiders were present, even when the scent of the flowers was masked. The spiders’ presence was seen asĀ nectar guides, which “bees have a pre-existing bias towards.” Members of this same research team also determined that both crab spiders and honey bees choose fragrant flowers over non-fragrant flowers, and that, ultimately, “honey bees suffer apparently from responding to the same floral characteristics as crab spiders do.”

Needless to say, crab spiders are crafty. So the question is, when killing machines like crab spiders are picking off a plant’s pollinators, does this affect its ability to reproduce? First let’s consider how pollinators react to finding crab spiders hiding in the flowers they hope to visit.

goldenrod crab spider (Misumena vatia) preying on a pollinator — via wikimedia commons

A study published in Oikos in 2003 observed patches of common milkweed (Asclepias syriaca) – one set was free of crab spiders, the other set was not – and tracked the visitations of four species of bees – the common honey bee and three species of bumble bees. They compared visitation rates between both sets of milkweed patches and found that the smallest of the three bumble bee species decreased its frequency of visitation to the crab spider infested milkweeds. Honey bees also appeared to visit the infested milkweeds less, but the results were not statistically significant. The two larger species of bumble bees continued to forage at the same rate despite the presence of crab spiders.

During the study, crab spiders were seen attacking bees numerous times. Six attacks resulted in successful kills, and of the bees that escaped, 80% left the flower and either moved to a different flower on the same plant, moved to a different plant, or left the patch altogether. These results indicate a potential for the presence of crab spiders to effect plant-pollinator interactions, whether its directly (predation) or indirectly (bees avoiding flowers with crab spiders).

Another study published in Behavioral Ecology in 2006 looked at two species of bees – the honey bee and a species of long-horned bee – and their reactions to the presence of crab spiders on the flowers of three different plant species – lavender (Lavandula stoechas), crimson spot rockrose (Cistus ladanifer), and sage-leaf rockrose (Cistus salvifolius). Honey bees were about half as likely to select inflorescences of lavender when crab spiders were present, and they avoided the crab spider infested flowers of crimson spot rockrose with a similar frequency. On the other hand, the long-horned bee visited the flowers of crimson spot rockrose to the same degree whether or not a crab spider was present.

bee visiting sage-leaf rock rose (Cistus salvifolius) — via wikimedia commons

The researchers then exposed honey bees to the flowers of sage-leaf rockrose that were at the time spider-free but showed signs that crab spiders had recently visited. Some of the flowers featured the scent of crab spiders, others had spider silk attached to them, and others had the corpses of dead bees on them. They found that even when crab spiders were no longer present, the bees could still detect them. Honey bees were particularly deterred by the presence of corpses. The long-horned bees were also exposed to the flowers with corpses on them but didn’t show a significant avoidance of them.

An interesting side note about the presence of silk on flowers. As stated earlier, crab spiders do not spin webs; however, they do spin silk for other reasons, including to tether themselves to flowers while hunting. The authors recount, “on several occasions when an attempted attack was observed during this study, it was only the presence of a silk tether that prevented spiders being carried away from flowers by their much larger prey.”

So, again, if bees are avoiding flowers due to the presence of predators like crab spiders, what effect, if any, is this having on the plants? We will address this question in part two.

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When Urban Pollinator Gardens Meet Native Plant Communities

Public concern about the state of bees and other pollinating insects has led to increased interest in pollinator gardens. Planting a pollinator garden is often promoted as an excellent way for the average person to help protect pollinators. And it is! However, as with anything in life, there can be downsides.

In many urban areas, populations of native plants remain on undeveloped or abandoned land, in parks or reserves, or simply as part of the developed landscape. Urban areas may also share borders with natural areas, the edges of which are particularly prone to invasions by non-native plants. Due to human activity and habitat fragmentation, many native plant populations are now threatened. Urban areas are home to the last remaining populations of some of these plants.

Concern for native plant populations in and around urban areas prompted researchers at University of Pittsburgh to review some of the impacts that urban pollinator gardens may have and to develop a “roadmap for research” going forward. Their report was published earlier this year in New Phytologist.

Planting a wildflower seed mix is a simple way to establish a pollinator garden, and such mixes are sold commercially for this purpose. Governmental and non-governmental organizations also issue recommendations for wildflower, pollinator, or meadow seed mixes. With this in mind, the researchers selected 30 seed mixes “targeted for urban settings in the northeastern or mid-Atlantic USA” to determine what species are being recommended for or commonly planted in pollinator gardens in this region. They also developed a “species impact index” to assess “the likelihood a species would impact remnant wild urban plant populations.”

A total of 230 species were represented in the 30 seed mixes. The researchers selected the 45 most common species for evaluation. Most of these species (75%) have generalized pollination systems, suggesting that there is potential for sharing pollinators with remnant native plants. Two-thirds of the species had native ranges that overlapped with the targeted region; however, the remaining one-third originated from Europe or western North America. The native species all had “generalized pollination systems, strong dispersal and colonization ability, and broad environmental tolerances,” all traits that could have “high impacts” either directly or indirectly on remnant native plants. Other species were found to have either high dispersal ability but low chance of survival or low dispersal ability but high chance of survival.

This led the researchers to conclude that “the majority of planted wildflower species have a high potential to interact with native species via pollinators but also have the ability to disperse and survive outside of the garden.” Sharing pollinators is especially likely due to super-generalists like the honeybee, which “utilizes flowers from many habitat types.” Considering this, the researchers outlined “four pollinator-mediated interactions that can affect remnant native plants and their communities,” including how these interactions can be exacerbated when wildflower species escape gardens and invade remnant plant communities.

photo credit: wikimedia commons

The first interaction involves the quantity of pollinator visits. The concern is that native plants may be “outcompeted for pollinators” due to the “dense, high-resource displays” of pollinator gardens. Whether pollinator visits will increase or decrease depends on many things, including the location of the gardens and their proximity to native plant communities. Pollinator sharing between the two has been observed; however, “the consequences of this for effective pollination of natives are not yet understood.”

The second interaction involves the quality of pollinator visits. Because pollinators are shared between native plant communities and pollinator gardens, there is a risk that the pollen from one species will be transferred to another species. High quantities of this “heterospecific pollen” can result in reduced seed production. “Low-quality pollination in terms of heterospecific pollen from wildflower plantings may be especially detrimental for wild remnant species.”

The third interaction involves gene flow between pollinator gardens and native plant communities. Pollen that is transferred from closely related species (or even individuals of the same species but from a different location) can have undesired consequences. In some cases, it can increase genetic variation and help address problems associated with inbreeding depression. In other cases, it can introduce traits that are detrimental to native plant populations, particularly traits that disrupt adaptations that are beneficial to surviving in urban environments, like seed dispersal and flowering time. Whether gene flow between the two groups will be positive or negative is difficult to predict, and “the likelihood of genetic extinction versus genetic rescue will depend on remnant population size, genetic diversity, and degree of urban adaptation relative to the planted wildflowers.”

The fourth interaction involves pathogen transmission via shared pollinators. “Both bacterial and viral pathogens can be transmitted via pollen, and bacterial pathogens can be passed from one pollinator to another.” In this way, pollinators can act as “hubs for pathogen exchange,” which is especially concerning when the diseases being transmitted are ones for which the native plants have not adapted defenses.

photo credit: wikimedia commons

All of these interactions become more direct once wildflowers escape gardens and establish themselves among the native plants. And because the species in wildflower seed mixes are selected for their tolerance of urban conditions, “they may be particularly strong competitors with wild remnant populations,” outcompeting them for space and resources. On the other hand, the authors note that, depending on the species, they may also “provide biotic resistance to more noxious invaders.”

All of these interactions require further investigation. In their conclusion, the authors affirm, “While there is a clear potential for positive effects of urban wildflower plantings on remnant plant biodiversity, there is also a strong likelihood for unintended consequences.” They then suggest future research topics that will help us answer many of these questions. In the meantime, pollinator gardens should not be discouraged, but the plants (and their origins) should be carefully considered. One place to start is with wildflower seed mixes, which can be ‘fine-tuned’ so that they benefit our urban pollinators as well as our remnant native plants. Read more about plant selection for pollinatorsĀ here.

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Bumblebees and Urbanization

Urban areas bear little resemblance to the natural areas that once stood in their place. Concrete and asphalt stretch out for miles, buildings of all types tower above trees and shrubs, and turfgrass appears to dominate whatever open space there is. Understandably, it may be hard to imagine places like this being havens for biodiversity. In many ways they are not, but for certain forms of life they can be.

An essay published earlier this year in Conservation Biology highlights the ways in which cities “can become a refuge for insect pollinators.” In fact, urban areas may be more inviting than their rural surroundings, which are often dominated by industrial agriculture where pesticides are regularly used, the ground is routinely disturbed, and monocultures reign supreme. Even though suitable habitat can be patchy and unpredictable in the built environment, cities may have more to offer than we once thought.

Yet, studies about bee abundance and diversity in urban areas show mixed results, largely because all bee species are not created equal (they have varying habitat requirements and life histories) and because urban areas differ wildly in the quality and quantity of habitat they provide both spatially and temporally. For this reason, it is important for studies to focus on groups of bees with similar traits and to observe them across various states of urbanization. This is precisely what researchers at University of Michigan set out to do when they sampled bumblebee populations in various cities in southeastern Michigan. Their results were published earlier this year by Royal Society Open Science.

common eastern bumble bee (Bombus impatiens) – photo credit: wikimedia commons

The researchers selected 30 sites located in Dexter, Ann Arbor, Ypsilanti, Dearborn, and Detroit. Most of the sites were gardens or farms in urban centers. They collected bumblebees from May to September using pan traps and nets. The species and sex of each individual bumblebee was identified and recorded for each site. The percentage of impervious surface that surrounded each site was used as a measurement of urban development. Other measurements included the abundance of flowers and average daily temperatures for each location.

Bumblebees were selected as a study organism because the genus, Bombus, “represents a distinct, well-studied set of traits that make it feasible to incorporate natural history into analysis.” Bumblebees live in colonies – eusocial structures that include “a single reproductive queen, variable numbers of non-reproductive female workers, and male reproductive drones.” They are generalist foragers, visiting a wide variety of flowering species for pollen and nectar, and they nest in holes in the ground, inside tree stumps, or at the bases of large clumps of grass. The authors believe that their nesting behavior makes them “a good candidate for testing the effects of urban land development,” and the fact that members of the colony have “distinct roles, [behaviors], and movement patterns” allows researchers to make inferences regarding “the effects of urbanization on specific components of bumblebee dynamics.”

Across all locations, 520 individual bumblebees were collected. Nearly three quarters of them were common eastern bumblebees (Bombus impatiens). Among the remaining nine species collected, brown-belted bumblebees (Bombus griseocollis) and two-spotted bumblebees (Bombus bimaculatus) were the most abundant.

brown-belted bumblebee (Bombus griseocollis) – photo credit: wikimedia commons

Because bumblebees are strong fliers with an extensive foraging range, impervious surface calculations for each site had to cover an area large enough to reflect this. Results indicated that as the percentage of impervious surfaces increased, bumblebee abundance and diversity declined. When male and female bumblebee data was analyzed separately, the decline was only seen in females; males were unaffected.

Female workers do most of their foraging close to home, whereas males venture further out. The researchers found it “reasonable to hypothesize that worker abundance is proportional to bumblebee colony density.” Thus, the decline in female bumblebees observed in this study suggests that as urban development increases (i.e. percent coverage of impervious surface), available nesting sites decline and the number of viable bumblebee colonies shrinks. Because male bumblebees responded differently to this trend, future studies should consider the responses of both sexes in order to get a more complete picture of the effects that urbanization has on this genus.

Interestingly, results obtained from the study locations in Detroit did not conform to the results found elsewhere. Bumblebee abundance and diversity was not decreasing with urbanization. Unlike other cities in the study, “Detroit has experienced decades of economic hardship and declining human populations.” It has a high proportion of impervious surfaces, but it also has an abundance of vacant lots and abandoned yards. These areas are left unmaintained and are less likely to be mowed regularly or treated with pesticides. Reducing disturbance can create more suitable habitat for bumblebees, resulting in healthy populations regardless of the level of urbanization. Thus, future studies should examine the state of insect pollinators in all types of cities – shrinking and non-shrinking – and should consider not just the amount of available habitat but also its suitability.

two-spotted bumblebee (Bombus bimaculatus) – photo credit: wikimedia commons

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When Sunflowers Follow the Sun

Tropisms are widely studied biological phenomena that involve the growth of an organism in response to environmental stimuli. Phototropism is the growth and development of plants in response to light. Heliotropism, a specific form of phototropism, describes growth in response to the sun. Discussions of heliotropism frequently include sunflowers and their ability to “track the sun.” This conjures up images of a field of sunflowers in full bloom following the sun across the sky. However cool this might sound, it simply doesn’t happen. Young sunflowers, before they bloom, track the sun. At maturity and in bloom, the plants hold still.

What is happening in these plants is still pretty cool though, and a report published in an August 2016 issue of Science sheds some light on the heliotropic movements of young sunflowers. They begin the morning facing east. As the sun progresses across the sky, the plants follow, ending the evening facing west. Over night, they reorient themselves to face east again. As they reach maturity, this movement slows, and most of the flowers bloom facing east. Over a series of experiments, researchers were able to determine the cellular and genetic mechanisms involved in this spectacular instance of solar tracking.

Helianthus annuus (common sunflower) is a native of North America, sharing this distinction with dozens of other members of this recognizable genus. It is commonly cultivated for its edible seeds (and the oil produced from them) as well as for its ornamental value. It is a highly variable species and hybridizes readily. Wild populations often cross with cultivated ones, and in many instances the common sunflower is considered a pesky weed. Whether crop, wildflower, or weed, its phototropic movements are easy to detect, making it an excellent subject of study.

Researchers began by tying plants to stakes so that they couldn’t move. Other plants were grown in pots and turned to face west in the morning. The growth of these plants was significantly stunted compared to plants that were not manipulated in these ways, suggesting that solar tracking promotes growth.

The researchers wondered if a circadian system was involved in the movements, and so they took sunflowers that had been growing in pots in a field and placed them indoors beneath a fixed overhead light source. For several days, the plants continued their east to west and back again movements. Over time, the movements became less detectable. This and other experiments led the researchers to conclude that a “circadian clock guides solar tracking in sunflowers.”

Another series of experiments helped the researchers determine what was happening at a cellular level that was causing the eastern side of the stem to grow during the day and the western side to grow during the night. Gene expression and growth hormone levels differed on either side of the stem depending on what time of day it was. In an online article published by University of California Berkeley, Andy Fell summarizes the findings: “[T]here appear to be two growth mechanisms at work in the sunflower stem. The first sets a basic rate of growth for the plant, based on available light. The second, controlled by the circadian clock and influenced by the direction of light, causes the stem to grow more on one side than another, and therefore sway east to west during the day.”

The researchers observed that as the plants reach maturity, they move towards the west less and less. This results in most of the flowers opening in an eastward facing direction. This led them to ask if this behavior offers any sort of ecological advantage. Because flowers are warmer when they are facing the sun, they wondered if they might see an increase in pollinator visits during morning hours on flowers facing east versus those facing west. Indeed, they did: “pollinators visited east-facing heads fivefold more often than west-facing heads.” When west-facing flowers where warmed with a heater in the morning, they received more pollinator visits than west-facing flowers that were not artificially warmed, “albeit [still] fewer than east-facing flowers.” However, increased pollinator visits may be only part of the story, so further investigations are necessary.

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Summer of Weeds: Common Mullein

The fuzzy, gray-green leaves of common mullein are familiar and friendly enough that it can be hard to think of this plant as a weed. Verbascum thapsusĀ is a member of the figwort family and is known by dozens of common names, including great mullein, Aaron’s rod, candlewick, velvet dock, blanket leaf, feltwort, and flannel plant. Its woolly leaves are warm and inviting and have a history of being used as added padding and insulation, tucked inside of clothing and shoes. In Wild Edible and Useful Plants of Idaho, Ray Vizgirdas writes, “the dried stalks are ideal for use as hand-drills to start fires; the flowers and leaves produce yellow dye; as a toilet paper substitute, the large fresh leaves are choice.”

Common mullein is a biennial that was introduced to eastern North America from Eurasia in the 1700’s as a medicinal plant and fish poison. By the late 1800’s it had reached the other side of the continent. In its first year it forms a rosette of woolly, oblong and/or lance-shaped leaves. After overwintering it produces a single flower stalk up to 6 feet tall. The woolly leaves continue along the flower stalk, gradually getting smaller in size until they reach the inflorescence, which is a long, dense, cylindrical spike. Sometimes the stalk branches out to form multiple inflorescences.

First year seedlings of common mullein (Verbascum thapsus)

The inflorescence doesn’t flower all at once; instead, a handful of flowers open at a time starting at the bottom of the spike and moving up in an irregular pattern. The process takes several weeks to complete. The flowers are about an inch wide and sulfur yellow with five petals. They have both female and male sex parts but are protogynous, meaning the female organs mature before the male organs. This encourages cross-pollination by insects. However, if pollination isn’t successful by the end of the day, the flowers self-pollinate as the petals close. Each flower produces a capsule full of a few hundred seeds, and each plant can produce up to 180,000 seeds. The seeds can remain viable for over 100 years, sitting in the soil waiting for just the right moment to sprout.

Common mullein is a friend of bare, recently disturbed soil. It is rare to see this plant growing in thickly vegetated areas. As an early successional plant, its populations can be abundant immediately after a disturbance, but they do not persist once other plants have filled in the gaps. Instead they wait in seed form for the next disturbance that will give them the opportunity to rise again. They can be a pest in gardens and farm fields due to regular soil disturbance, and are often abundant in pastures and rangelands because livestock avoid eating their hairy leaves. Because of its ephemeral nature, it is generally not considered a major weed; however, it is on Colorado’s noxious weed list.

Several features make common mullein a great example of a drought-adapted plant. Its fleshy, branching taproot can reach deep into the soil to find moisture, the thick hairs on the leaves help reduce water loss via transpiration, and the way the leaves are arranged and angled on the stalk can help direct rain water down toward the roots.

Common mullein has an extensive history of ethnobotanical uses. Medicinally it has been used internally to treat coughs, colds, asthma, bronchitis, and kidney infections; and as a poultice to treat warts, slivers, and swelling. The dried flower stalks have been used to make torches, and the fuzzy leaves have been used as tinder for fire-making and wicks in lamps.

The hairy leafscape of common mullein (Verbascum thapsus)

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From Gaia’s Garden by Toby Hemenway

Here’s why opportunistic plants are so successful. When we clear land or carve a forest into fragments, we’re creating lots of open niches. All that sunny space and bare soil is just crying out to be colongized by light- and fertlity-absorbing green matter. Nature will quickly conjure up as much biomass as possible to capture the bounty, by seeding low-growing ‘weeds’ into a clearing or, better yet, sprouting a tall thicket stretching into all three dimensions to more effectively absorb light and develop deep roots. … When humans make a clearing, nature leaps in, working furiously to rebuild an intact humus and fungal layer, harvest energy, and reconstruct all the cycles and connections that have been severed. A thicket of fast-growing pioneer plants, packing a lot of biomass into a small space, is a very effective way to do this. … And [nature] doesn’t care if a nitrogen fixer or a soil-stabilizing plant arrived via continental drift or a bulldozer’s treads, as long as it can quickly stitch a functioning ecosystem together.