Bat Pollinated Flowers of a Mexican Columnar Cactus

Pollination syndromes – suites of floral traits used to determine potential pollinators and routes of pollination – have been informative in studying plant-pollinator interactions, but are generally too simplistic to tell the full story. Most flowering plants are generalists when it comes to pollinators, whereas pollination syndromes imply specialization. Not all pollinators are created equal though, and some may be more effective at pollinating particular plants than others. In fact, occasionally pollination syndromes ring true and a predicted plant-pollinator combination turns out to be the most effective and reliable interaction.

According to a study published in American Journal of Botany by Ibarra-Cerdeña, et al., Stenocereus queretaroensis, a species of columnar cactus endemic to western Mexico, adheres to this scenario. Stenocereus is a genus in a group of columnar and tree-like cacti called the Pachycereeae tribe. Cactus in this group are generally bat pollinated; however, their flowers are typically visited by various species of birds and insects as well, and in some cases, bats are not the primary pollinator. In their introduction, the authors note that specialization appears to be more common in tropical latitudes, and chiropterophilic (bat pollinated) columnar cacti that occur in temperate regions can be comparatively more generalized. This is because “extratropical chiropterophilic cacti appear to be faced with unpredictable seasonal year-to-year variation in pollinators,” while “cacti in tropical regions” experience “highly reliable seasonal availability of nectar-feeding bats, thereby leading to a temporally stable pollination system.”

Stenocereus queretaroensis is a massive cactus, reaching up to ten meters tall. Several vertical stems rise from a short, stocky, central trunk. Each stem has up to eight distinctive ribs and averages around 15 centimeters in diameter. Groupings of white to grey spines up to four centimeters long appear along the ribs. Flowers are light-colored, around 10 to 14 centimeters in length, and occur along the upper half of the stems, extended well beyond the spines. Flowers open at night – producing abundant nectar – and close by the afternoon the following day. Floral characteristics led the authors of this study to predict bats to be the main pollinator, and they set up a series of experiments to test this.

Stenocereus queretaroensis - photo credit: wikimedia commons

Stenocereus queretaroensis – photo credit: wikimedia commons

Part of their experiment consisted of five treatments involving 130 flowers on 75 plants. One group of flowers was bagged and allowed to self-pollinate naturally, while another group was bagged and self-pollinated manually. A third group was left exposed during the night but bagged in the morning, while a fourth group was bagged during the night and exposed during the daytime. The final group was left alone. For each of these five treatments, aborted flowers and mature fruits were counted and seed set was determined. Nectar samples were taken from a separate group of flowers at two hour intervals from 8:00 PM to 8:00 AM, after which no nectar was produced. A camera was also used to document floral visits. Visits were deemed “legitimate” when the “visitor’s body came in contact with anthers and/or stigma” and “illegitimate” when “no contact with anthers or stigma” was made.

The researchers found S. queretaroensis to be “incapable of self-pollination,” as no fruit set occurred for the first two treatments. The control group and the nocturnally exposed group had nearly identical results, producing significantly more fruits with greater seed set compared to the nocturnally bagged group. During the day, flowers were visited by four species of birds (two hummingbirds, a woodpecker, and an oriole) and several species of bees (mainly honey bees). During the night, apart from illegitimate visits from a nectar robbing hawkmoth, one species of bat was the dominant floral visitor, and the majority (93.8%) of the visits were legitimate. This bat species was Leptonycteris curasoae, the southern long-nosed bat.

Leptonycteris curasoae - photo credit: wikimedia commons

Leptonycteris curasoae – photo credit: wikimedia commons

The abundance of nectar-feeding bats was monitored in the study area over a four year period, and L. curasoae was by far the most abundant species throughout the study period. Nectar produced in the flowers of S. queretaroensis was at its maximum around midnight, which seemed to correlate with observations of bat visits. Even though daytime visitors appeared to contribute to fruit and seed set, the nocturnal treatment produced significantly more fruit with significantly higher seed set, suggesting that bats are the more efficient pollinator. Insects visiting during the daytime, when nectar was decreasingly available, were most likely robbing pollen.

The authors acknowledge that for most plant species, “a wide array of taxonomically diverse fauna such as insects, birds, and mammals usually serve as potential pollinators,” and that “generalized pollination systems are more frequent than specialized ones.” However, in this case, “a close association between L. curasoae and S. queretaroensis [suggests] that the chiropterophilic syndrome is still a useful model.”

Related Posts:

Advertisements

Bats As Pollinators – An Introduction to Chiropterophily

Most plants that rely on animals to assist in pollination look to insects. In general, insects are abundant, easy to please, and efficient at transferring pollen. Because insect pollination is such a common scenario, it is easy to overlook pollination that is carried out by vertebrates. Birds are the most prominent pollinator among vertebrates, but mammals participate, too. The most common mammal pollinator is the bat.

About a fifth of all mammal species on the planet are bats, with species estimates numbering in the 1200-1300 range. Bats are the only mammals that can truly fly. They are not blind, nor are they flying rodents, and they are not going to suck your blood (except in very rare cases!). Most bats eat insects, but a small, significant group of them are nectarivorous. Their main food source is the nectar produced within flowers. In the process of feeding, these bats pollinate plants.

Out of 18 families in the order Chiroptera, only two include species with morphologies that set them apart as nectar-feeders. The family Pteropodidae, known commonly as Old World fruit bats or flying foxes, occurs in tropical and subtropical regions of Africa, Asia, Australia, Papa New Guinea, and the Pacific Islands. The family Phyllostomidae, known commonly as American leaf-nosed bats, occurs in tropical and subtropical regions of the Americas. For simplicity’s sake, the former are referred to as Old World bats, and the latter as New World bats. While both groups are similar in that they consist of species that feed on nectar, they are only distantly related, and thus the nectar feeding species in these families have distinct behavioral and morphological differences.

Grey headed flying fox photo credit: wikimedia commons

Grey headed flying fox (Pteropus poliocephalus), a floral visiting bat from Australia (photo credit: wikimedia commons)

More than 500 species of plants, spanning 67 plant families, are pollinated by bats. This pollination syndrome is known as chiropterophily. In general, flowers that use this approach tend to be white or dull in color, open at night, rich with nectar, and musty or rotten smelling. They are generally tubular, cup shaped, or otherwise radially symmetrical and are often suspended atop tall stalks or prominently located on branches or trunks. In a review published in Annals of Botany, Theodore Fleming, et al. state “flower placement away from foliage and nocturnal anthesis [blooming] are the unifying features of the bat pollination syndrome,” while all other characteristics are highly variable among species. The family Fabaceae contains the highest number of bat-pollinated genera. Cactaceae, Malvaceae, and Bignoniaceae follow closely behind.

The characteristics of bat pollinated flowers vary widely partly because the bats that visit them are so diverse. Between the two bat families there are similarities in their nectar-feeding species, including an elongated rostrum, teeth that are smaller in number and size, and a long tongue with hair-like projections on the tip. Apart from that, New World bats are much smaller than Old World bats, and their rostrums and tongues are much longer relative to the size of their bodies. New World bats have the ability to hover in front of flowers, while Old World bats land on flowers to feed. Old World bats do not have the ability to use echolocation to spot flowers, while New World bats do. Fleming, et al. conclude, “because of these differences, we might expect plants visited by specialized nectar-feeding [New World bats] to produce smaller flowers with smaller nectar volumes per flower than those visited by their [Old World bat] counterparts.”

Pollination by bats is a relatively new phenomenon, evolutionarily speaking. Flowers that are currently pollinated by bats most likely evolved from flowers that were once pollinated by insects. Some may have evolved from flowers that were previously bird pollinated. The question is, why adopt this strategy? Flowers that are bat pollinated are “expensive” to make. They are typically much bigger than insect pollinated flowers, and they contain large amounts of pollen and abundant, nutrient-rich nectar. Due to resource constraints, many plants are restricted from developing such flowers, but those that do may find themselves at an advantage with bats as their pollinator. For one, hairy bat bodies collect profuse numbers of pollen grains, which are widely distributed as they visit numerous flowers throughout the night. In this way, bats can be excellent outcrossers. They also travel long distances, which means they can move pollen from one population of plants to an otherwise isolated neighboring population. This serves to maintain healthy genetic diversity among populations, something that is increasingly important as plant populations become fragmented due to human activity.

Pollinating bats are also economically important to humans, as several plants that are harvested for their fruits, fibers, or timber rely on bats for pollination. For example, bat pollinated Eucalyptus species are felled for timber in Australia, and the fruits of Durio zibethinus in Southeast Asia form after flowers are first pollinated by bats. Also, the wild relatives of bananas (Musa spp.) are bat pollinated, as is the plant used for making tequila (Agave tequilana).

Durio sp. (photo credit: wikimedia commons)

The flowers of durian (Durio sp.), trees native to Southeast Asia, are pollinated by bats (photo credit: wikimedia commons)

There is still much to learn about nectarivorous bats and the flowers they visit. It is clear that hundreds of species are using bats to move their pollen, but the process of adopting this strategy and the advantages of doing so remain ripe for discovery. Each bat-plant relationship has its own story to tell. For now, here is a fun video about the bat that pollinates Agave tequilana:

Year of Pollination: Pollination Syndromes and Beyond

A discussion of pollination syndromes should begin with the caveat that they are a largely outdated way to categorize plant-pollinator interactions. Still, they are important to be aware of because they have informed so much of our understanding about pollination biology, and they continue to be an impetus for ongoing research. The concept of pollination syndromes exists in part because we are a pattern seeking species, endeavoring to place things in neat little boxes in order to make sense of them. This is relatively easy to do in a hypothetical or controlled environment where the parameters are selected and closely monitored and efforts are made to eliminate noise. However, the real world is considerably more dynamic than a controlled experiment and does not conform to black and white ways of thinking. Patterns are harder to unveil, and it takes great effort to ensure that observed patterns are genuine and not simply imposed by our pattern seeking brains.

That being said, what are pollination syndromes?  Pollination syndromes are sets of floral traits that are thought to attract specific types of pollinators. The floral traits are considered to have evolved in order to appeal to a particular group of pollinators – or in other words, selective pressures led to adaptations resulting in mutualistic relationships between plants and pollinators. Pollination syndromes are examples of convergent evolution because distantly related plant species have developed similar floral traits, presumably due to similar selection pressures. Pollination syndromes were first described by Italian botanist, Federico Delpino, in the last half of the 19th century. Over several decades his rudimentary ideas were fleshed out by other botanists, resulting in the method of categorization described (albeit briefly) below.

Honey bee on bee's friend (Phacelia tanacetifolia)

A honey bee getting friendly with bee’s friend (Phacelia tanacetifolia)

Pollination by bees (melittophily) – Flowers are blue, purple, yellow, or white and usually have nectar guides. Flowers are open and shallow with a landing platform. Some are non-symmetrical and tubular like pea flowers. Nectar is present, and flowers give off a mild (sometimes strong) sweet scent.

Pollination by butterflies (psychophily) – Flowers are pink, purple, red, blue, yellow, or white and often have nectar guides. They are typically large with a wide landing pad. Nectar is inside a long, narrow tube (or spur), and flowers have a sweet scent.

Pollination by hawkmoths and moths (sphingophily and phalaenophily) – Moth pollinated flowers open at night, have no nectar guides, and emit a strong, sweet scent. Flowers pollinated by hawkmoths are often white, cream, or dull violet and are large and tubular with lots of nectar. Those pollinated by other moths are smaller, not as nectar rich, and are white or pale shades of green, yellow, red, purple, or pink.

Pollination by flies (myophily or sapromyophily) – Flowers are shaped like a basin, saucer, or kettle and are brown, brown-red, purple, green, yellow, white, or blue.  Some have patterns of dots and stripes. If nectar is available, it is easily accessible. Their scent is usually putrid. A sapromyophile is an organism that is attracted to carcasses and dung. Flies that fall into this category visit flowers that are very foul smelling, offer no nectar reward, and essentially trick the fly into performing a pollination service.

Pollination by birds (ornithophily) –  Flowers are usually large, tubular, and red, orange, white, blue, or yellow. They are typically without nectar guides and are odorless since birds don’t respond to scent. Nectar is abundant and found at various depths within the flower.

Pollination by bats (chiropterophily) – Flowers are large, tubular or bell shaped, and white or cream colored with no nectar guides. They open at night, have abundant nectar and pollen, and have scents that vary from musty to fruity to foul.

Pollination by beetles (cantharophily) – Flowers are large and bowl shaped and green or white. There are no nectar guides and usually no nectar. The scent is strong and can be fruity, spicy, or putrid. Like flies, some beetles are sapromyophiles.

Locust borer meets rubber rabbitbrush (Ericameria nauseosa)

A locust borer meets rubber rabbitbrush (Ericameria nauseosa)

In addition to biotic pollination syndromes, there are two abiotic pollination syndromes:

Pollination by wind (anemophily) – Flowers are miniscule and brown or green. They produce abundant pollen but no nectar or odor. The pollen grains are very small, and the stigmas protrude from the flower in order to capture the windborne pollen.

Pollination by water (hydrophily) –  Most aquatic plants are insect-pollinated, but some have tiny flowers that release their pollen into the water, which is picked up by the stigmas of flowers in a similar manner to plants with windborne pollen.

This is, of course, a quick look at the major pollination syndromes. More complete descriptions can be found elsewhere, and they will differ slightly depending on the source. It’s probably obvious just by reading a brief overview that there is some overlap in the floral traits and that, for example, a flower being visited by a bee could also be visited by a butterfly or a bird. Such an observation explains, in part, why this method of categorizing plant-pollinator interactions has fallen out of favor. Studies have been demonstrating that this is not a reliable method of predicting which species of pollinators will pollinate certain flowers. A close observation of floral visitors also reveals insects that visit flowers to obtain nectar, pollen, and other items, but do not assist in pollination. These are called robbers. On the other hand, a plant species may receive some floral visitors that are considerably more effective and reliable pollinators than others. What is a plant to do?

Pollination syndromes imply specialization, however field observations reveal that specialization is quite rare, and that most flowering plants are generalists, employing all available pollinators in assisting them in their reproduction efforts. This is smart, considering that populations of pollinators fluctuate from year to year, so if a plant species is relying on a particular pollinator (or taxonomic group of pollinators) to aid in its reproduction, it may find itself out of luck. Considering that a flower may receive many types of visitors on even a semi-regular basis suggests that the selective pressures on floral traits may not solely include the most efficient pollinators, but could also include all other pollinating visitors and, yes, even robbers. This is an area where much more research is needed, and questions like this are a reason why pollination biology is a vibrant and robust field of research.

A bumble bee hugs Mojave sage (Salvia pachyphylla)

A bumble bee hugs the flower of a blue sage (Salvia pachyphylla)

Interactions between plants and pollinators is something that interests me greatly. Questions regarding specialization and generalization are an important part of these interactions. To help satiate my curiosity, I will be reading through a book put out a few years ago by the University of Chicago Press entitled, Plant-Pollinator Interactions: From Specialization to Generalization, edited by Nickolas M. Waser and Jeff Ollerton. You can expect future posts on this subject as I read through the book. To pique your interest, here is a short excerpt from Waser’s introductory chapter:

Much of pollination biology over the past few centuries logically focused on a single plant or pollinator species and its mutualistic partners, whereas a focus at the level of entire communities was uncommon. Recently we see a revival of community studies, encouraged largely by new tools borrowed from the theory of food webs that allow us to characterize and analyze the resulting patterns. For example, pollination networks show asymmetry – most specialist insects visit generalist plants, and most specialist plants are visited by generalist insects. This is a striking departure from the traditional implication of coevolved specialists!

References: