How a cactus from the Andes may be using hairs to attract its bat pollinators

Plants go to great lengths to attract pollinators. From brightly colored flowers to alluring scents and even some sexual deception, there seems to be no end to what plants will do for sex. Recently, research on the pollination of a species of cactus endemic to the Ecuadorian Andes suggests that even plant hairs can be co-opted for pollinator attraction.

Espostoa frutescens is a wonderful columnar cactus that grows from 1,600 ft (487 m) to 6,600 ft (2011 m) in the Ecuadorean Andes. Like many other high elevation cacti, this species is covered in a dense layer of hairy trichomes. These hairs serve an important function in these mountains by protecting the body of the plant from excessive heat, cold, wind, and UV radiation. Espostoa frutescens takes this a step further when it comes time to flower. It is one of those species that produces a dense layer of hairs around its floral buds called a cephalium. Cacti cephalia are thought to have evolved as a means of protecting developing flowers and fruits from the outside elements. What scientists have now discovered is that, at least for some cacti, the cephalium may also serve an important role in attracting bats.

Bats are famous for their use of echolocation. Because they mainly fly at night, bats rely on sound and scent, rather than sight to find food. More and more we are realizing that a lot of plants have taken advantage of this by producing structures that reflect bat sonar in such a way that makes them more appealing to bats. Some plants, like Mucuna holtonii and Marcgravia evenia, do this for pollination. Others, like Nepenthes hemsleyana, do this to obtain a nitrogen-rich meal.

Espostoa frutescens apparently differs from these examples in that its not about reflecting bat sonar, but rather absorbing it at specific frequencies. Close examination of the hairs that comprise the E. frutescens cephalium revealed that they were extremely well adapted for absorbing ultrasonic frequencies in the 90 kHz range. This may seem arbitrary until you look at who exactly pollinates this cactus.

The main pollinator for E. frutescens is a species of bat known as Geoffroy’s tailless bat (Anoura geoffroyi). It turns out that Geoffroy’s tailless bat happens to echolocate at a frequencies right around that 90 kHz range. Whereas the rest of the body of the cactus reflects plenty of sound, bat calls reaching the cephalium of E. frutescens bounced back an average of 14 decibels quieter.

Essentially, the area of floral reward on this species of cactus presents a much quieter surface than the rest of the plant itself. It is very possible that this functions as a sort of calling card for Geoffroy’s tailless bats looking for their next meal. This makes sense from a communication standpoint in that it not only saves the bats valuable foraging time, it also increases the chances of cross pollination for the cactus. To obtain enough energy from flowers, bats must travel great distances. Anything that helps them locate a meal faster will increase visitation to that flower. By changing the way in which the flowers “appear” to echolocating bats, the cacti thus increase the amount of visitation from bats, which brings pollen in from cacti located over the bats feeding range.

It is important to note that, at this point in time, research has only been able to demonstrate that the hairs surrounding E. frutescens flowers are more absorbent to the ultrasonic frequencies used by Geoffroy’s tailless bat. We still have no idea whether bats are more likely to visit flowers borne from cephalia or not. Still, this research paves the way for even more experiments on how plants like E. frutescens may be “communicating” with pollinators like bats.

Photo by Merlin Tuttle’s Bat Conservation. Please Consider supporting this incredible conservation group!

Further Reading: [1]

To grow or to flower, that is the cactus conundrum

Melocactus intortus

Melocactus intortus

Flowers are costly structures for plants to produce. In the flowering plant world, there is always a trade-off between growth and reproduction. Flowers are produced from tiny structures called axillary buds, and many plants can only produce one flush of flowers per bud. Cacti are no exception to this rule and their amazing morphological adaptations to harsh climates has forced them into quite a conundrum when it comes to reproduction.

The axillary buds of cacti are located at the base of their spines in little structures called areoles. This is where the flowers will eventually emerge. However, unlike plants that can produce cheap stems and branches, cacti must produce a whole new chunk of stem or internode before they can produce more axillary buds. Think of it this way, if a cactus wants to produce 10 flowers, it must produce ten internodes to do so. This means producing all of the expensive cortex and epidermis along with it. Their harsh environments have forced most cacti into an extremely tight relationship between growth, water storage, photosynthesis, and flowering that is potentially very limiting from a reproductive standpoint.

Micranthocereus estevesii with lateral cephalium

Micranthocereus estevesii with lateral cephalium

Amazingly, some cacti have managed to break from this evolutionary relationship and they have done so in a bizarre way. Take a look at all of the cacti pictured here. Each has developed a strange looking structure called a cephalium. Essentially, you can think of the cephalium of a cactus as its “adult” reproductive form whereas the rest of the body consists of non-reproductive, photosynthetic “juvenile” form.

The cephalium is a unique and fascinating structure. It differs from the rest of the cactus body in that it is not photosynthetic. It also produces no chlorophyll and no stomata. In fact, it does not form anything like the epidermis of the rest of the plant. Instead, the cephalium produces dense clusters of short spines and trichomes. Most importantly, it produces tightly packed axillary buds in high abundance. These are the buds that will produce the flowers. The end result is a wacky looking structure that has the ability to produce far more flowers than that of cacti that do not grow a cephalium.

Facheiroa tenebrosa with lateral cephalium

Facheiroa tenebrosa with lateral cephalium

Obviously not all cacti produce cephalia but it is common in genera such as Melocactus, Backebergia, Espostoa, Discocactus, and Facheiroa (this is not a complete list). What the cephalium has done for genera like these is decouple the afore mentioned relationships between growth and reproduction. For a period of time (often many years) following germination, these cacti grow the typical succulent, photosynthetic stems we are accustomed to seeing.

At some point in their development, something triggers these plants to switch to their adult forms. Axillary buds within either lateral or apical meristems switch their growth habit and begin forming the cephalium. It is worth mentioning that no one yet knows what triggers this switch. If the cephalium is produced from axillary buds in the apical meristem like we see in Melocactus, the plant will no longer produce photosynthetic tissues. This represents another major trade-off for these cacti. Such species must rely on the photosynthetic juvenile tissues for all of their photosynthetic needs for the rest of their lives (unless the cephalium is damaged or lost). Backebergia have managed to get around this trade-off by not only growing multiple stems, they will also shed their apical cephalia after a few years, thus re-initiating photosynthetic juvenile growth.

Backebergia militaris with bizarre apical cephalia reminiscent of the bearskin hats of the Queen’s guard.

Backebergia militaris with bizarre apical cephalia reminiscent of the bearskin hats of the Queen’s guard.

Things are a bit different for cacti that produce lateral cephalia. Genera such as Espostoa, Facheiroa, and Buiningia are less limited by their cephalia because they are produced along the ribs of the stem, thus leaving the apical meristem free to continue more typical photosynthetic growth. Nonetheless, the process is much the same. Dense clusters of spines, trichomes, and most importantly, axillary buds are produced along the rib, giving each stem a lovely, lopsided appearance.

There are other benefits to growing cephalia in addition to simply being able to produce more flowers. The densely packed spines and trichomes offer the developing flowers and fruits ample protection from both the elements and herbivores. Floral buds are free to develop deep within the interior of the cephalium until they are mature. At that point, the cells will begin to swell with water, pushing the flower outward from the cephalium where it will be exposed to pollinators. As the petals curl back, they offer a safe spot for visiting pollinators that is free from menacing spines. Once pollination has been achieved, the flower wilts and the deeply inferior ovaries are then free to develop within the safety of the cephalium. Once the fruits are mature, they too will begin to swell with water and be pushed out from the cephalium where they will attract potential seed dispersers.

Melocactus violaceus with fruits emerging from the cephalium

Melocactus violaceus with fruits emerging from the cephalium

I hope that I have convinced you of just how awesome this growth form can be. I will never forget the first time I saw a cactus topped with a cephalium. It was a mature Melocactus growing in a cactus house. Sticking out of the odd “cap” on top was a ring of bright pink fruits. I knew nothing of the structure at that time but it was incredible to see. Now that I know what it is and how it functions, I am all the more appreciative of these cacti.

This post was inspired by the diligent work of Dr. Jim Mauseth. Click here to learn more about cacti.

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

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

Big Things Come In Small Packages

il_570xN.1271356583_c10y.jpg

Meet Blossfeldia liliputana, the smallest species of cactus in the world. With a maximum diameter of only 12 mm, this wonderful succulent would be hard to spot tucked in among the nooks and crannies of rock outcrops. Its species name "liliputana" is a reference to the fictional island of Liliput (Gulliver's Travels) whose inhabitants were said to be rather small. If its size alone wasn't interesting in and of itself, the biology of B. liliputana is also downright bizarre.

Blossfeldia liliputana is native to arid regions between southern Bolivia and northern Argentina. It appears to prefer growing wedged between cracks in rock as these are usually the spots where just enough soil builds up to put down its roots. Root formation, however, does not happen for quite some time. Most often new individuals bud off from the parent plant. They emerge not from the base, but rather from apical tissues, yet another unique feature of this cactus. What's more, this cactus produces no spines. Instead, its numerous areoles are covered in a dense layer of trichomes that are felt-like to the touch.

As you can clearly see, this species is small. It only ever becomes conspicuous when it comes time to flower. Imagine a bunch of tiny white to pink cactus flowers poking out of a crevice. It must be a remarkable sight to see in person. Despite their showy appearance, its is believed that most are self-fertilized.

Photo by Mats Winberg licensed under CC BY-SA 2.5

Photo by Mats Winberg licensed under CC BY-SA 2.5

As mentioned, the size of this cactus isn't the only interesting thing about its biology. B. liliputana is categorized as a poikilohydric organism, meaning it doesn't have the ability to regulate its internal water content. Researchers have found that individual plants can lose up to 80% of their weight in water and can maintain that state for as long as two years without any negative effects. As such, colonies of these tiny cacti often appear shrunken or squished. Once the rains arrive, however, it springs back to its original rounded shape with seemingly no issues. Amazingly, a significant amount of water uptake happens via the fuzzy areoles that cover its surface, hence it does not harm the plant to hold off growing roots for quite some time. 

Speaking of water regulation, B. liliputana holds another record for having the lowest density of stomata of any terrestrial autotrophic vascular plant. Stomata are the pores in which plants regulate water and gas exchange so having so few may have something to do with why this species loses and gains water to such a degree that would kill most other vascular plant species.

Another peculiar quality of this cactus are its seeds. Unlike all other cacti whose seeds are hard and relatively smooth, the seeds of B. liliputana are hairy. Attached to each seed is a small fleshy structure called an aril, which aids in seed dispersal. As it turns out, B. liliputana relies on ants as its main seed dispersers. Ants attracted to the fleshy aril drag the seeds back to their nests, remove and eat the aril, and then discard the seed. This is often good news for the cactus because its seeds end up in nutrient-rich ant middens protected from both the elements and seed predators, often in much more suitable conditions for germination.

Photo by Michael Wolf licensed under CC BY-SA 3.0

Photo by Michael Wolf licensed under CC BY-SA 3.0

Needless to say, B. liliputana is a bit of an oddball as far as cacti are concerned. Its highly derived features coupled with its bizarre biology have made it difficult for taxonomists to elucidate its relationship to the rest of the cactus family. It certainly deserves its own genus, to which it is the only member, however, it has been added to and removed form a handful of cactus subfamilies over the years. The most recent genetic analyses suggests that it is unique enough to warrant its own tribe within Cactaceae - Blossfeldieae.

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

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

Leafy Cacti?

Pereskia aculeata  photo by scott.zona licensed under CC BY 2.0

Pereskia aculeata photo by scott.zona licensed under CC BY 2.0

At first glance, there is little about a Pereskia that would suggest a relation to what we know as cacti. Even a second, third, and forth glance probably wouldn't do much to persuade the casual observer that these plants have a place on cacti family tree. All preconceptions aside, Pereskia are in fact members of the family Cactaceae and quite interesting ones at that.

Most people readily recognize the leafless, spiny green stems of a cactus. Indeed, this would appear to be a unifying character of the family. Pereskia is proof that this is not the case. Though other cacti occasionally produce either tiny, vestigial leaves or stubby succulent leaves, Pereskia really break the mold by producing broad, flattened leaves with only a hint of succulence.

Pereskia spines are produced from areoles in typical cactus fashion. Photo by Frank Vincentz licensed under CC BY-SA 3.0

Pereskia spines are produced from areoles in typical cactus fashion. Photo by Frank Vincentz licensed under CC BY-SA 3.0

What's more, instead of clusters of Opuntia-like pads or large, columnar trunks, Pereskia are mainly shrubby plants with a handful of scrambling climbers mixed in. Similar to their more succulent cousins, the trunks of Pereskia are usually adorned with clusters of long spines for protection. Additionally, each species produces the large, showy, cup-like blooms we have come to expect from cacti.

They are certainly as odd as they are beautiful. As it stands right now, taxonomists recognize two clades of Pereskia - Clade A, which are native to a region comprising the Gulf of Mexico and Caribbean Sea (this group is currently listed under the name Leuenbergeria) and Clade B, which are native to regions just south of the Amazon Basin. This may seem superficial to most of us but the distinction between these groups has a lot to teach us about the evolution of what we know of as cacti. 

Pereskia grandifolia Photo by Anne Valladares (public domain)

Pereskia grandifolia Photo by Anne Valladares (public domain)

Genetically speaking, the genus Pereskia sorts out at the base of the cactus family tree. Pereskia are in fact sister to all other cacti. This is where the distinction between the two Pereskia clades gets interesting. Clade A appears to be the older of the two and all members of this group form bark early on in their development and their stems lack a feature present in all other cacti - stomata. Stomata are microscopic pours that allow the exchange of gases like CO2 and oxygen. Clabe B, on the other hand, delay bark formation until later in life and all of them produce stomata on their stems.

The reason this distinction is important is because all other cacti produce stomata on their stems as well. As such, their base at the bottom of the cactus tree not only shows us what the ancestral from of cactus must have looked like, it also paints a relatively detailed picture of the evolutionary trajectory of subsequent cacti lineages. It would appear that the ancestor of all cacti started out as leafy shrubs that lacked the ability to perform stem photosynthesis. Subsequent evolution saw a delay in bark formation, the presence of stomata on the stem, and the start of stem photosynthesis, which is a defining feature of all other cacti.

Pereskia aculeata Photo by Ricardosdag licensed under CC BY-SA 4.0

Pereskia aculeata Photo by Ricardosdag licensed under CC BY-SA 4.0

If you are as excited about Pereskia as I am, then you , my friend, are in luck. A handful of Pereskia species have found their way into the horticulture trade. With a little luck attention to detail, you too can share you home with one of these wonderful plants. Just be warned, they get tall and their spines, which are often hidden by the leaves, are a force to be reckoned with. Tread lightly with these wonderfully odd cacti. Celebrate their as the evolutionary wonders that they are!

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

Further Reading: [1] [2]