An Intruiguing Relationship Between Ants and Cacti

The extrafloral nectaries of  Pachycereus gatesii  appear as tiny red bumps just below the areole.

The extrafloral nectaries of Pachycereus gatesii appear as tiny red bumps just below the areole.

It’s hard to think of a group of plants that are better defended than cacti. Frequently and often elaborately adorned with vicious spines, these succulents make any animal think twice about trying to take a bite. And yet, for some cacti, spines don’t seem to cut it. A surprising amount of species appear to have taken their defense system to a whole new level by recruiting nature’s most tenacious bodyguards, ants.

Plants frequently have a friend in ants. Spend some time observing ants at work and it’s east to see why. These social insects have numbers and strength on their side. Give ants a reason to be invested in your survival and they will certainly see to it that nothing threatens this partnership. For cacti, this involves the secretion of nectar from specialized tissues called extrafloral nectaries.

Extrafloral nectaries are not unique to cacti. A multitude of plant species produce them, often for similar reasons. Ants love a sugary food source and the more reliable that source becomes, the more adamant an ant colony will be at defending it. The odd thing about cacti is that they don’t seem to have settled on a single type of extrafloral nectary to do the trick. In fact, as many as four different types of extrafloral nectaries have been described in the cactus family.

Ants visiting the extrafloral nectaries covering the developing flowers of  Pilosocereus gounellei .

Ants visiting the extrafloral nectaries covering the developing flowers of Pilosocereus gounellei.

Some cacti secrete nectar from highly modified spines. A great example of this can be seen in genera such as Coryphantha, Cylindropuntia, Echinocactus, Ferocactus, Opuntia, Sclerocactus, and Thelocactus. Such spines are usually short and blunt, hardly resembling spines at all. Other cacti secrete nectar from regular looking spines. This adaptation is odd as there does not seem to be anything special about the anatomy of such spines. Examples of this can be seen in genera such as Brasiliopuntia, Calymmanthium, Harrisia, Opuntia, Pereskiopsis, and Quiabentia. Still others secrete nectar from highly reduced leaves that are found at the base of where the spines originate (the areole). Such leaves have been described in Acanthocereus, Leptocereus, Myrtillocactus, Pachycereus, and Stenocereus. They aren’t easy to recognize as leaves either. Most look like tiny scales. Finally, the fourth type of extrafloral nectary comes in the form of specialized regions of the stem tissue. This has been described in genera such as Armatocereus, Leptocereus, and Pachycereus.

Highly modified spines functioning as extrafloral nectaries in  Ferocactus emoryi.

Highly modified spines functioning as extrafloral nectaries in Ferocactus emoryi.

Seemingly normal spines of  Harrisia pomanensis  secreting nectar.

Seemingly normal spines of Harrisia pomanensis secreting nectar.

Regardless of where they form, their function remains much the same. They secrete a form of nectar which ants find irresistible. The more reliable this food source becomes, the more aggressive ant colonies will be in defending it. This is an especially useful form of defense when it comes to small insect herbivores. Whereas spines deter larger herbivores, they don’t do much to deter organisms that can just slip right through them unharmed. Ants also clean the cacti, potentially removing harmful microbes like fungi and bacteria. Though we are only just beginning to understand the depths of this cactus/ant mutualism, what we have discovered already suggests that the relationship between these types of organisms is far more complex than what I have just outlined above.

For instance, it may not just be sugar that the ants are looking for. In arid desert habitats, water may be the most limiting resource for an ant colony and large, succulent cacti are essentially giant water reservoirs. The key is getting to that water. One study that looked at a species of barrel cactus growing in Arizona called Ferocactus acanthodes found that as spring gives way to summer, the concentration of sugars secreted by the extrafloral nectaries decreases. As a result, the nectar becomes far more watery. Amazingly, ant densities on any given barrel cactus actually increased throughout the summer, despite the fact that the nectar was being watered down. Ants are notoriously prone to desiccation so it stands to reason that water, rather than sugar, is the real prize for colonies hanging out on cacti in such hot desert environments.

The incredible floral display of  Ferocactus wislizeni , a species whose reproductive efforts are affected by the types of ants they attract.

The incredible floral display of Ferocactus wislizeni, a species whose reproductive efforts are affected by the types of ants they attract.

Another interesting observation about the cactus/ant mutualism is that it appears that the identity of the ants truly matters. Though defense is the main benefit to the cactus, research suggests that there is a tipping point in how much such defenses benefit cacti. It has been found that although cacti initially benefit from anti-herbivore and cleaning services, extra aggressive ant species can actually drive off potential pollinators. At least one study has shown that when less aggressive ant species tend cacti, they produce more fruits and those fruits contain significantly more seeds than cacti that have been tended by extremely aggressive ant species. This is especially concerning when we think about the growing issue of invasive ants. As more and more non-native ant species displace native ants, this could really tip the balance for some cactus species.

Despite all of the interesting things we have learned about extrafloral nectaries in the family Cactaceae, there are so many questions yet to be answered. For starters, we still do not know how many different taxa produce them in one form or another. It is likely that closer inspection, especially of rare or poorly understood groups, will reveal that far more cacti produce some type of extrafloral nectary. Also, we know next to nothing about the anatomy of the different types of nectaries. How do they differ from one another and how do some, especially those derived from ordinary spines, actually function? Finally, do these nectaries function year round or is there some sort of seasonal pattern to their development and utility. How does this affect the types of ants they attract and how does that in turn affect the survival and reproduction of these cacti? For such a charismatic group of plants as cacti, we still have to much to learn.

Photo Credits: Thanks to Dr. Jim Mauseth and Dr. John Rebman and Dr. Silvia Rodriguez Machado for use of their photos [1] [2] [3]

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

The Trumpet Creeper

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With its impressive bulk and those stunning tubular red flowers, one would be excused for thinking that the trumpet creeper (Campsis radicans) was a tropical vine. Indeed, the family to which it belongs, Bignoniaceae, is largely tropical in its distribution. There are a handful of temperate representatives, however, and the trumpet creeper is one of the most popular. Its beauty aside, this plant is absolutely fascinating.

As many of you probably know, the trumpet creeper can reach massive proportions. In the garden, this can often result in collapsed structures as its weight and speed of growth is something few adequately prepare for. In the wild, I most often see this vine in somewhat disturbed forests, usually near a floodplain. As such, it is supremely adapted to take a hit and keep on growing year after year.

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One of the many reasons this plant performs so well both where it is native and where it is not is that it recruits body guards. This is easy to witness in a garden setting as the branches and especially the flowers are frequently crawling with ants. Trumpet creepers trade food for protection via specialized organs called extrafloral nectaries. These structures secrete sugary nectar that is readily sucked up by tenacious ants. When a worker ant finds a vine, more workers are soon to follow. 

Amazingly for a temperate plant, trumpet creepers produce more extrafloral nectaries of all four categories - petiole, calyx, corolla, and fruit. What this means is that all of the important organs are covered in insects that viciously attack anything that might threaten this sugary food supply. Hassle one of these vines at your own peril. With its photosynthetic and reproductive structures protected, trumpet creepers make a nice living once established.

Reproduction is another fascinating aspect of trumpet creeper biology. A closer inspection of the floral anatomy will reveal a bilobed stigma. Amazingly, this stigma has the ability to open and close as potential pollinators visit the flowers. Stigmatic movement in the trumpet creeper has attracted a bit of attention from researchers over the years. What is its function?

Evidence suggests that the opening and closing of the lobed stigma is way of increasing the chances of pollination. Touch alone is not enough to trigger the movement. However, when researchers dusted pollen onto the stigma, then it began to close. What's more, this action happens within 15 to 60 seconds. Amazingly, there appears to be a threshold to whether the stigma stays closed or reopens after 3 hours or so.

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It turns out, the threshold seems to depend on the amount of pollen being deposited. Only after 350 grains found their way onto the stigma did it close permanently. Experts feel that this a means by which the plant ensured ample seed set. If too few pollen grains end up on the stigma, the plant risks not having all of its ovules fertilized. By permanently closing after enough pollen grains are present, the plant can ensure that the pollen grains can germinate and fertilize the ovules without being brushed off.

It is interesting to note that the flowers frequently remain on the plant after they have been fertilized. This likely serves to maintain a largely floral display that continues to attract pollinators until most of the flowers have been pollinated. Speaking of pollinators, observations have revealed that the trumpet creeper is pollinated primarily by ruby-throated hummingbirds. Although insects like bumblebees frequently visit these blooms, bringing pollen with them in the process, hummingbirds, on average, bring and deposit 10 times as much pollen as any other visitor. And, considering the threshold on pollen mentioned above, trumpet creeper appears to have evolved a pollination syndrome with these lovely little birds.

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

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

A Unique Passionflower Endemic to Costa Rica

I love small flowers, especially if they pack in a lot of detail. That's is why this passion flower caught my eye. Meet Passiflora boenderi, a charismatic vine endemic to a small region of Costa Rica. Apparently this species had been sitting around in herbaria for years under a different name. It wasn't until living specimens were observed that botanists realized it is a distinct species.

There is a lot to look at on this species. The flowers themselves are some of the smallest in the genus. They pack in all of the detail of a larger passion flower, just in miniature. The leaves are quite stunning as well. They're bilobed with a tinge of purple and covered in bright, orange-yellow spots. The spots themselves serve an important role in protecting this plant from herbivores.

The genus Passiflora is part of an intense evolutionary arms race with a genus of butterfly known as Heliconius. Their caterpillars feed on the foliage of passion flowers. As such, Passiflora have evolved a variety of means that help them to avoid the attention of gravid female butterflies. The orange spots on the leaves of P. boenderi are one such adaptation and they serve a dual function.

The first is a visual deterrent. Female Heliconius prefer to lay their eggs on caterpillar-free leaves. This makes sense. Why bother laying eggs where there will be ample competition for food. The spots mimic, both in size and shape, the appearance of Heliconius eggs. A female looking for a spot to lay will see these spots and move on to another plant. In addition to the visual mimicry, these spots also secrete nectar. The energy-rich nectar inevitably attracts ants, which viciously defend them as a food source. If a caterpillar (or any other herbivore fore that matter) were to start munching on the leaves, the ants quickly drive them off.

Because of its limited range, P. boebderi is under threat of extinction. Habitat destruction of its lowland habitat for palm oil, pineapples, and vacation resorts is an ongoing threat to the long term survival of this species and many others. I was fortunate enough to have encountered this plant growing in the Cliamtron at the Missouri Botanical Garden but I fear that if we keep on doing what we humans are so good at, botanical gardens may be the only place this species will be found growing in the not too distant future.

Further Reading: [1] [2]

Live-In Mites

Hearing the word "mite" as a gardener instantly makes me think of pests such as spider mites. This is not fair. The family to which mites belong (Acari) is highly varied and contains many beneficial species. Many mites are important predators at the micro scale. Some are fungivorous, eating potentially harmful species of fungi. Whereas this may be lost on the majority of us humans, it is certainly not lost on many species of plants. In fact, the relationship between some plants and mites is so strong that these plants go as far as to provide them with a sort of home.

Domatia are specialized structures that are produced by plants to house arthropods. A lot of different plant species produce domatia but not all of them are readily apparent to us. For instance, many trees and vines such as red oak (Quercus rubra), sugar maples (Acer saccharum), black cherries (Prunus serotina), and many species of grape (Vitis spp.) produce tiny domatia specifically for mites. The domatia are often small, hairy, and function as shelter for both the mites and their eggs.

By housing certain species of mites, these plants are ensuring that they have a steady supply of hunters and cleaners living on their leaves. Predatory mites are voracious hunters, keeping valuable leaves free of microscopic herbivores while frugivorous mites clean the leaves of detrimental fungi that are known to cause infections such as powdery mildew. The exchange is pretty straight forward. Mites get a home and a place to breed and the plants get some protection. Still, some plants seem to want to sweeten the relationship in a literal sense.

Some plants, specifically grape vines in the genus Vitis, also produce extrafloral nectaries on their leaves. These tiny glands secrete sugary nectar. In a paper recently published in the Annals of Botany, it was found that extrafloral nectaries enhances the efficacy of these mite domatia by enticing more mites to stick around. By adding nectar to domatia-producing leaves that did not secrete it, the researchers found that nectar increases beneficial mite densities on these leaves by 60 - 80%. This translates to an increase in fitness for these plants in the long run.

I love research like this. I had no idea that so many of my favorite and most familiar tree and vine species had entered into an evolutionary relationship with beneficial mites. This adds a whole new layer of complexity to the interactions within any given environment. It just goes to show you how much is left to be discovered in our own back yards.

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

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

When a Mutualism Becomes Obligate

Mutualism. The word invokes this warm and fuzzy "you scratch my back and I'll scratch yours" feeling. It is easy to grasp how a mutualism would develop and be maintained. But, in any system, there are bound to be cheaters. Cheaters reduce the fitness of one of the partners so to avoid such things, some species up the ante by resorting to some interestingly "sinister" methods.

Acacias and ants have quite the relationship. Acacias protect themselves by offering ants hollow spines and branches where their colonies can live. They even sweeten the deal via extrafloral nectaries. These are glands on the stems that secrete nectar that the ants eat. In some ant species, this is their only source of food. Needless to say, the ants become highly protective of their acacia trees. They readily attack herbivores and even go as far as to prune away vegetation that may interfere with their host. This seems like a pretty straight forward mutualistic relationship, right?

Ah, but it goes deeper. To make sure that the ants will solely rely on the acacia and are thus completely tied up in the well being of their host, the acacia alters the ants phenotype at birth. Normally these ants have no issues digesting sucrose. Researchers found that the nectar in the extrafloral nectaries contains a protein called "chitinase." Chitinase inhibits the ability of the ants to digest sucrose. When ant eggs hatch into larvae, their first meal is nectar from the extra floral nectaries. Once the larvae ingest this protein they are no longer able to feed on anything other than their hosts nectar. Thus their very survival is completely tied to the Acacia.

I am positive that more examples of such obligate mutualisms abound in nature. We only have to ask the right questions to discover them. It is also interesting considering what we are finding out about our own behavior and how it relates to the microbiome living on and within us. What about human behavior could be described in the context of a relationship similar to ants and acacias?

Photo Credit: Tony Rodd

Further Reading:
http://www.ncbi.nlm.nih.gov/pubmed/24188323