Year of Pollination: An Argentinian Cactus and Its Unlikely Pollinator

A few weeks ago I wrote about pollination syndromes – sets of floral triats that are said to attract specific groups of pollinators. In that post I discussed how pollination syndromes have largely fallen out of favor as a reliable method of predicting the pollinators that will visit particular flowers. In this post I review a recent study involving a species of cactus in Argentina that, as the authors state in their abstract, “adds another example to the growing body of mismatches between floral syndrome and observed pollinator.”

Denmoza rhodacantha is one of many species of cacti found in Argentina. It is the only species in its genus, and it is widely distributed across the east slopes and foothills of the Andes. It is a slow growing cactus, maintaining a globulous (globe-shaped) form through its juvenile phase and developing a columnar form as it reaches maturity. D. rhodacantha can reach up to 4 meters tall and can live beyond 100 years of age. Individual plants can begin flowering in their juvenile stage. Flowers are red, nectar rich, scentless, and tubular. The stigma is lobed and is surrounded by a dense grouping of stamens. Both male and female reproductive organs are extended above the corolla. The flowers have been described by multiple sources as being hummingbird pollinated, not based on direct observation of hummingbirds visiting the flowers, but rather due to the floral traits of the species.

Denmoza rhodacantha illustration - image credit:

Denmoza rhodacantha illustration  (image credit:

In a paper entitled, Flowering phenology and observations on the pollination biology of South American cacti – Denmoza rhodacantha, which was published in volume 20 of Haseltonia (the yearbook of the Cactus and Succulent Society of America), Urs Eggli and Mario Giorgetta discuss their findings after making detailed observations of a population of D. rhodacantha in early 2013 and late 2013 – early 2014. The population consisted of about 30 individuals (both juveniles and adults) located in the Calchaqui Valley near the village of Angastaco, Argentina. At least three other species with “hummingbird-syndrome flowers” were noted in the area, and three species of hummingbirds were observed during the study periods. Over 100 observation hours were logged, and during that time “the studied plants, their flowering phenology, and flower and fruit visitors were documented by means of photographs and video.”

The flowers of D. rhodacantha only persist for a few short days, and in that time their sexual organs are only receptive for about 24 hours. The flowers are self-sterile and so require a pollinator to cross pollinate them. Despite their red, tubular shape and abundant nectar, no hummingbirds were observed visiting the flowers. One individual hummingbird approached but quickly turned away. Hummingbirds were, however, observed visiting the flowers of an associated species, Tecoma fulva ssp. garrocha. Instead, a species of halictid bee (possibly in the genus Dialictus) was regularly observed visiting the flowers of D. rhodacantha. The bees collected pollen on their hind legs and abdomen and were seen crawling across the lobes of the stigma. None of them were found feeding on the nectar. In one observation, a flower was visited by a bee that was “already heavily loaded with the typical violet-coloured pollen of Denmoza,” suggesting that this particular bee species was seeking out these flowers for their pollen. Small, unidentified beetles and ants were seen entering the flowers to consume nectar, however they didn’t appear to be capable of offering a pollination service.

D. rhodacantha populations have been observed in many cases to produce few fruits, suggesting that pollination success is minimal. The authors witnessed “very low fruit set” in the population that they were studying, which was “in marked contrast to the almost 100% fruit set rates of the sympatric cactus species at the study site.” This observation wasn’t of great concern to the authors though, because juvenile plants are present in observed populations, so recruitment appears to be occurring. In this study, dehisced fruits were “rapidly visited by several unidentified species of ants of different sizes.” The “scant pulp” was harvested by smaller ants, and larger ants carried away the seeds after “cleaning them from adhering pulp.”

The authors propose at least two reasons why hummingbirds avoid the flowers of D. rhodacantha. The first being that the native hummingbirds have bills that are too short to reach the nectar inside the long tubular flowers, and often the flowers barely extend beyond the spines of the cactus which may deter the hummingbirds from approaching. The second reason is that other plants in the area flower during the same period and have nectar that is easier to gather. The authors acknowledge that this is just speculation, but it could help explain why the flowers are pollinated instead by an insect (the opportunist, generalist halictid bee species) for whom the flowers “could be considered to be ill adapted.” The authors go on to say, “it should be kept in mind, however, that adaptions do not have to be perfect, as long as they work sufficiently well.”

Patagona gigas (giant hummingbird) was observed approaching the flower of a Denmoza rhodacantha but quickly turned away (photo credit:

Patagona gigas (giant hummingbird) was observed approaching the flower of a Denmoza rhodacantha but quickly turned away (photo credit:

More Year of Pollination posts on Awkward Botany:

Making the Case for Saving Species

It is no question that the human species has had a dramatic impact on the planet. As our population has grown and we have spread ourselves across the globe, our presence has altered every ecosystem we have come into contact with. Our footprints can be detected even in areas of the planet uninhabited by humans. As awareness of our impact has increased, we have made efforts to reduce it. However, much of the damage we have caused is irreversible – we can’t bring species back from extinction and we can’t replace mountaintops. Furthermore, for better or for worse our continued existence – despite efforts to minimize our negative influence – will continue to be impactful. This is the nature of being human. It is the nature of all living things, really. As John Muir said, “when we try to pick out anything by itself, we find it hitched to everything else in the Universe.” That we are cognizant of that fact puts us at a crossroads – do we make a concerted effort to protect and save other species from the negative aspects of our presence or do we simply go on with our lives and let come what may?

The quandary isn’t that black and white, obviously. For one thing, cleaning up polluted air, water, and soil is beneficial to humans and has the side benefit of improving the lives of other species. Protecting biodiversity is also in our best interest, because who knows what medicine, food, fiber, or other resource is out there in some living thing yet to be discovered that might be useful to us. On the other hand, putting our own interests aside, what about protecting other species and habitats just to protect them? Purely altruistically. That seems to be the question at the crux of an article by Emma Marris in the May/June 2015 issue of Orion entitled, “Handle with Care: The Case for Doing All We Can to Save Threatened Species.” [Listen to a brief discussion with Marris about the article here.]

The main character in Marris’ article is the whitebark pine (Pinus albicaulis), a species whose native habitat is high in mountain ranges of western United States and Canada. Whitebark pines thrive in areas few other trees can, living to ages greater than 1,000 years. Here is how Marris describes them:

Whitebark pine’s ecological niche is the edge of existence. The trees are found on the highest, driest, coldest, rockiest, and windiest slopes. While lodgepole and ponderosa pine grow in vast stands of tall, healthy-looking trees, slow-growing whitebarks are tortured by extremes into individualized, flayed forms, swollen with massive boles from frost damage. Their suffering makes them beautiful.

photo credit:

photo credit:

But in recent years they have been suffering more than usual. White pine blister rust, an introduced pathogen, is killing the trees. The native mountain pine beetle is also taking them out. Additional threats include climate change and an increased number, extent, and intensity of wildfires. Combined, these threats have been impactful enough that the species is listed as endangered on the IUCN Red List where it is described as “experiencing serious decline.”

So people are taking action. In Oregon’s Crater Lake National Park, botanist Jen Beck is part of an effort to select blister rust resistant trees and plant them in their native habitats within the park. Hundreds have been planted, and more are on their way. Great effort is taken to minimize human impact and to plant the trees as nature would, with the vision being that blister rust resistant trees will replace those that are dying and that trees with rust resistant genes will dominate the population.

But Beck faces opposition, and not just from challenges like seedlings being trampled by visitors or a warming climate inviting mountain hemlocks and other trees into whitebark pine’s native range, but by people who argue that the trees shouldn’t be planted there in the first place – that what is “wild” should be left alone. Marris specifically calls out a group called Wilderness Watch. They and other groups like them profess a “leave-it alone ethic.” Rather than be arrogant enough to assume that we can “control or fix disrupted nature,” we should respect the “self-willed spirit of the wild world.” Proponents of nonintervention criticize what they call “new environmentalism” and its efforts to engineer or manage landscapes, fearing that these actions are “morally empty” and that “rearranging bits of the natural world” lacks soul and will ultimately serve to benefit humans.

In her article, Marris argues against this approach. First off, the human footprint is too large, and for natural areas to “continue to look and function the way they did hundreds of years ago” will require “lots of human help.” Additionally, nonintervention environmentalism “perpetuates a false premise that humans don’t belong in nature,” and if we decide not to work to protect, save, or restore species and habitats that have been negatively affected by our actions simply because we are “in thrall to wildness”, we will be withdrawing with “blood on our hands.” Marris sums up her position succinctly in the following statement:

We have to do whatever it takes to keep ecosystems robust and species from extinction in the face of things like climate change. And if that means that some ecosystems aren’t going to be as pretty to our eyes, or as wild, or won’t hew to some historical baseline that seems important to us, then so be it. We should put the continued existence of other species before our ideas of where or how they should live.

Marris acknowledges that there are risks to this approach. “Our meddling” may save species, but it could also backfire. But that doesn’t mean the effort wasn’t worth it. We can learn from our mistakes and we can make improvements to our methods. Some sites can even be cordoned off as areas of nonintervention simply so that we can learn from them. The ultimate goal, however, should be to save as many species and to keep as much of their habitat intact as possible. Putting “other species first, and our relationship with them second” is what Marris considers to be a “truly humble” stance in our role as part of nature.

Cones of whitebark pine, Pinus albicaulis (photo credit: wikimedia commons)

Cones of whitebark pine, Pinus albicaulis (photo credit: wikimedia commons)

The dichotomy presented in this article is a tough one, and one that will be debated (in my mind particularly) long into the future. If you would like to share your thoughts with me about this issue, do so in the comment section below or by sending me a private message through the contact page.

Other article reviews on Awkward Botany

Ethnobotany: White Man’s Foot, part one

“Plantains – Plantago major – seem to have arrived with the very first white settlers and were such a reliable sign of their presence that the Native Americans referred to them as ‘white men’s footsteps.'” – Elizabeth Kolbert (The Sixth Extinction)

“Our people have a name for this round-leafed plant: White Man’s Footstep. Just a low circle of leaves, pressed close to the ground with no stem to speak of, it arrived with the first settlers and followed them everywhere they went. It trotted along paths through the woods, along wagon roads and railroads, like a faithful dog so as to be near them.” – Robin Wall Kimmerer (Braiding Sweetgrass)

photo credits: wikimedia commons

photo credit: wikimedia commons

Plantago major is in the family Plantaginaceae – the plantain family – a family that consists of at least 90 genera, several of which include common species of ornamental plants such as Veronica (speedwells), Digitalis (foxgloves), and Antirrhinum (snapdragons). The genus Plantago consists of around 200 species commonly known as plantains. They are distributed throughout the world in diverse habitats. Most of them are herbaceous perennials with similar growth habits, and many have ethnobotanical uses comparable to P. major.

Originating in Eurasia, P. major now has a cosmopolitan distribution. It has joined humans as they have traveled and migrated from continent to continent and is now considered naturalized throughout most temperate and some tropical regions. In North America, P. major and P. lanceolata are the two most common introduced species in the Plantago genus. P. major has a plethora of common names – common plantain being the one that the USDA prefers. Other names include broadleaf plantain, greater plantain, thickleaf plantain, ribgrass, ribwort, ripplegrass, and waybread. Depending on the source, there are various versions of the name white man’s foot, and along the same line, a common name for P. major in South Africa is cart-track plant.

P. major is a perennial – albeit sometimes annual or biennial – herbaceous plant. Its leaves form a rosette that is usually oriented flat against the ground and reaches up to 30 cm wide. Each leaf is egg-shaped with parallel veins and leaf margins that are sometimes faintly toothed. The inflorescence is a leafless spike up to 20 cm tall (sometimes taller) with several tiny flowers that are a dull yellow-green-brown color. The flowers are wind pollinated, and the plants are highly prone to self-pollination. The fruits are capsules that can contain as many as 30 seeds – an entire plant can produce as many as 14,000 – 15,000 seeds at once. The seeds are small, brown, sticky, and easily transported by wind or by adhering to shoes, clothing, animals, and machinery. They require light to germinate and can remain viable for up to 60 years.

An illustration of three Plantago species found in Selected Weeds of the United States - Agriculture Handbook No. 366 circa 1970

An illustration of three Plantago species found in Selected Weeds of the United States – Agriculture Handbook No. 366 circa 1970

P. major prefers sunny sites but can also thrive in part shade. It adapts to a variety of soil types but performs best in moist, clay-loam soils. It is often found in compacted soils and is very tolerant of trampling. This trait, along with its low-growing leaves that easily evade mower blades, explains why it is so commonly seen in turf grass. It is highly adaptable to a variety of habitats and is particularly common on recently disturbed sites (natural or human caused) and is an abundant urban and agricultural weed.

Even though it is wind pollinated, its flowers are visited by syrphid flies and various bee species which feed on its pollen. Several other insects feed on its foliage, along with a number of mammalian herbivores. Cardinals and other bird species feed on its seeds.

Humans also eat plantain leaves, which contain vitamins A, C, and K. Young, tender leaves can be eaten raw, while older leaves need to be cooked as they become tough and stringy with age. The medicinal properties of  P. major have been known and appreciated at least as far back as the Anglo-Saxons, who likely used a poultice made from the leaves externally to treat wounds, burns, sores, bites, stings, and other irritations. Native Americans, after seeing the plant arrive with European settlers, quickly learned to use the plant as food and medicine. It could be used to stop cuts from bleeding and to treat rattlesnake bites. Apart from external uses, the plant was used internally as a pain killer and to treat ulcers, diarrhea, and other gastrointestinal issues.

P. major has been shown to have antibacterial, anti-inflammatory, antioxidant, and other biological properties; several chemical compounds have been isolated from the plant and deemed responsible for these properties. For this reason, P. major and other species of Plantago have been used to treat a number of ailments. The claims are so numerous and diverse that it is worth exploring if you are interested. You can start by visiting the following sites:

"White man's footstep, generous and healing, grows with its leaves so close to the ground that each step is a greeting to Mother Earth." - Robin Wall Kimmerer, Braiding Sweetgrass (photo credit:

“White man’s footstep, generous and healing, grows with its leaves so close to the ground that each step is a greeting to Mother Earth.” – Robin Wall Kimmerer, Braiding Sweetgrass (photo credit:

Other Ethnobotany Posts on Awkward Botany:

Year of Pollination: More than Honey, etc.

When I decided to spend a year writing about pollinators and pollination, I specifically wanted to focus on pollinators besides the honey bee. Honey bees already get lots of attention, and there are loads of other pollinating organisms that are equally fascinating. But that’s just the thing, honey bees are incredibly fascinating. They have a strict and complex social structure, and they make honey – two things that have led humans to develop a strong relationship with them. We have been managing honey bees and exploiting their services for thousands of years, and we have spread them across the planet, bringing them with us wherever we go. In North America, honey bees are used to pollinate a significant portion of our pollinator-dependent crops, despite the fact that they are not native to this continent. In that sense, they are just another domesticated animal, artificially selected for our benefit.

It’s common knowledge that honey bees (and pollinators in general) have been having a rough time lately. Loss of habitat, urbanization, industrial farming practices, abundant pesticide use, and a variety of pests and diseases have been making life difficult for pollinators. Generally, when the plight of pollinators comes up in the news, reference is made to honey bees (or another charismatic pollinator, the monarch butterfly). News like this encourages people to take action. On the positive side, efforts made to protect honey bees can have the side benefit of protecting native pollinators since many of their needs are the same. On the negative side, evidence suggests that honey bees can compete with native pollinators for limited resources and can pass along pests and diseases. Swords are often double-edged, and there is no silver bullet.

In a recent conversation with a budding beekeeper, I was recommended the documentary, More than Honey. I decided to watch it, write a post about it, and call that the honey bee portion of the Year of Pollination. Part way through the movie, another documentary, Vanishing of the Bees, was recommended to me, and so I decided to watch both. Below are some thoughts about each film.

more than honey movie

More than Honey

Written and directed by Swiss documentary filmmaker, Markus Imhoof, this beautifully shot, excellently narrated, meandering documentary thrusts viewers into incredibly intimate encounters with honey bees. Cameras follow bees on their flights and into their hives and get up close and personal footage of their daily lives, including mating flights, waggle dances, pupating larvae, flower pollination, and emerging queens. In some scenes, the high definition shots make already disturbing events even more disturbing, like bees dying after being exposed to chemicals and tiny varroa mites crawling around on the bodies of bees infecting them with diseases – wings wither away and bees become too weak to walk. This movie is worth watching for the impressive cinematography alone.

But bees aren’t the only actors. The human characters are almost as fun to watch. A Swiss beekeeper looks out over stunning views of the Alps where he keeps his bees. He follows a long tradition of beekeeping in his family and is very particular about maintaining a pure breed in his hives, going so far as flicking away the “wrong” bees from flowers on his property and crushing the head off of an unfaithful queen. A commercial beekeeper in the United States trucks thousands of beehives around the country, providing pollination services to a diverse group of farms – one of them being a massive almond grove in California. He has been witness to the loss of  hundreds of honey bee colonies and has had to become “comfortable with death on an epic scale” – the grueling corporate world grinds along, and there is no time for mourning losses.

Further into the documentary, a woman in Austria demonstrates how she manipulates a colony into raising not just one queen, but dozens. She has spent years breeding bees, and her queens are prized throughout the world. A man in Arizona captures and raises killer bees – hybrid bees resulting from crosses between African and European honey bees (also known as Africanized honey bees). Despite their highly aggressive nature, he prefers them because they are prolific honey producers and they remain healthy without the use of synthetic pesticides.

Probably the darkest moment in the film is watching workers in China hand pollinate trees in an orchard. Excessive pesticide use has decimated pollinator populations in some regions, leaving humans to do the pollinating and prompting the narrator to reflect on the question, “Who’s better at pollinating, man or bees? Science answers with a definite, ‘not man.'”

Also included in the film is an intriguing discussion about bees as a super-organism with a German neuroscientist who is studying bee brains. The narrator sums it up like this: “Without its colony the individual bee cannot survive. It must subordinate its personal freedom for the good of the colony… Could it be that individual bees are like the organs or cells of a body? Is the super-organism as a whole the actual animal?”


Vanishing of the Bees

Colony collapse disorder is a sometimes veiled yet important theme throughout More than Honey, and it was certainly something that drove the creation of the film. In the case of Vanishing of the Bees, colony collapse disorder is the reason for its existence. Narrated by actor, Ellen Page, and produced in part by a film production company called Hive Mentality Films, this movie came out on the heels of the news that bee colonies were disappearing in record numbers throughout the world. It tells the story of colony collapse disorder from the time that it first appeared in the news – one of the film’s main characters is the beekeeper that purportedly first brought attention to the phenomenon – and into the years that followed as scientists began exploring potential causes.

This film contains lots of important information and much of it seems credible, but it is also the type of documentary that in general makes me wary of documentaries. Its purpose goes beyond just trying to inform and entertain; it’s also trying to get you on board with its cause. I may agree with much of what is being said, but I don’t particularly like having my emotions targeted in an effort to manipulate me to believe a certain way. It’s a good idea not to let documentaries or any other type of media form your opinions for you. Consider the claims, do some of your own research and investigation, and then come to your own conclusion. That’s my advice anyway…even though you didn’t ask for it.

That being said, colony collapse disorder is a serious concern, and so I’ll end by going back to More than Honey and leave you with this quote by its narrator:

The massive death of honey bees is no mystery. What’s killing them is not pesticides, mites, antibiotics, incest, or stress, but a combination of all these factors. They are dying as a result of our civilization’s success, as a result of man, who has turned feral bees into docile, domestic animals – wolves into delicate poodles.