A Rare Case of Ant Pollination in Australia

Photo by Nicola Delnevo [SOURCE]

Ants have struck up a lot of interesting and important relationships with plants. They disperse seeds, protect plants from herbivores and disease, and can even help acquire nutrients. For all of the beneficial ways in which ants and plants interact, pollination rarely enters into the equation. More often than not, ants are actually detrimental to the sex lives of flowering plants. Such is not the case for a rare species of protea endemic to Western Australia called the smokebush (Conospermum undulatum).

The reason ants usually suck at pollination is thanks to a tiny organ called the metapleural gland. For many ant species, this gland secretes special antimicrobial fluids that the ants use to groom themselves. Because ants tend to live in high densities in close quarters, this antimicrobial fluid helps keep their little bodies clean of any pathogens that might threaten their existence. For as good as these fluids are for ants, they destroy pollen grains, rendering them useless for pollination.

Leioproctus conospermi. Photo by Sarah McCaffrey licensed under CC BY-ND 2.0.

As is so often the case in nature, there are always exceptions to the rule and it seems that one such exception is playing out in Western Australia. While investigating the reproductive ecology of the smokebush, researchers noted that ants were regular visitors to their small flowers. They knew that in drier climates, some ant species have evolved to produce considerably less antimicrobial fluids. The thought is that drier climates tend to harbor fewer microbial pathogens and thus ants don’t need to waste as much energy protecting themselves from such threats. If this was the case in Western Australia then it was entirely possible that ants could potentially serve as pollinators for this plant. Armed with this hypothesis, they decided to take a closer look.

It turns out that the floral morphology of the smokebush lends well to visiting ant anatomy. The tiny flowers produce a small amount of nectar at the base. As ants shove their heads down into the flower to get a drink, it triggers an explosive mechanism that causes the style the smack down onto the back of the ant. In doing so, it also mops up any pollen the ant may be carrying. At the same time, the anthers explosively dehisce, coating the visitor with a fresh dusting of pollen. During their observations, researchers noted that ants weren’t the only insects visiting smokebush blooms. They also noted lots of visitation from invasive honeybees (Apis mellifera) and a tiny native bee called Leioproctus conospermi.

(A) White flowers of Conospermum undulatum. (B) Floral details. (C–H) Insects visiting flowers of C. undulatum: (C) Leioproctus conospermi; (D) Camponotus molossus; (E) Camponotus terebrans; (F) Iridomyrmex purpureus; (G) Myrmecia infima; (H) Apis mellifera. [SOURCE]

After recording visits, researchers needed to know whether any of these floral visitors resulted in successful pollination. After all, just because something visits a flower doesn’t mean it has what it takes to get the job done for the plant. By looking at differences in seed set between ant and bee visitors, they were able to paint a fascinating picture of the pollination ecology of the rare smokebush.

It turns out that ants are indeed excellent pollinators of this shrub, contributing just as much to overall seed set as the tiny native Leioproctus conospermi. Alternatively, invasive honeybees barely functioned as pollinators at all. Their heads were too big to effectively trigger the pollination mechanism of the flowers but nonetheless were able to access the nectar within. As such, honeybees are considered nectar thieves for the smokebush, harming its overall reproductive effort rather than helping.

Amazingly, the effectiveness of ants as smokebush pollinators is not because they produce less antimicrobial fluids. In fact, these ants were fully capable of producing ample amounts of these pollen-killing substances. Instead, it appears that the plant itself has evolved to tolerate ant visitors. Smokebush pollen is resistant to the toxic effects of the metaplural gland fluids. With plenty of hungry ants always on the lookout for food, the smokebush has managed to tap in to an abundant and reliable vector for pollination. No doubt other examples exist, we simply have to go looking.

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

Further Reading: [1]

Emus + Ants = One Heck of a Seed Dispersal Strategy

A guest post by Dr. Scott Zona

The emu is a large, flightless bird, a cousin of kiwis and cassowaries. They range throughout much of Australia, favoring savannah woodlands and sclerophyll forests, where they are generalist feeders, consuming a variety of plants and arthropods. A favorite food of the emu is Petalostigma pubescens, a tree variously known as quinine tree, bitter bark or quinine berry. Petalostigma is in the Picrodendraceae, a family formerly included in the Euphorbiaceae. Quinine trees grow in the same open woodlands favored by emus.

The quinine tree bears yellow fruits, 2.0-2.5 cm in diameter, with a thin layer of flesh. The fruits are divided into six to eight segmented, like a tangerine, and each segment contains a hard endocarp or stone (technically, a pyrene). Each endocarp contains a single seed, 6-8 mm long. Left on the tree, the fruits will eventually dry up and open to release their seeds, but if ripe fruits are discovered by a hungry emu, the feasting begins.

A quinine tree (Petalostigma pubescens) in bloom. Photo by Ethel Aardvark licensed by CC BY 3.0

An emu may eat dozens of fruits in one meal. It swallows fruits whole, digesting the soft, fleshy part and defecating the hard, indigestible endocarps. On an average day, an emu can range over a large territory, spreading endocarps as it goes. In one of science's least glamorous moments, Australian biologists counted by hand as many as 142 endocarps in one emu dropping. If the story ended with Quinine Tree seeds in a pile of emu dung, we would say that the emu provided excellent seed dispersal services for the quinine tree, but the dispersal story is not over.

Quinine tree (Petalostigma pubescens) fruits. Photo by Robert Whyte licensed by CC BY-NC-ND 2.0

The emu dung and endocarps begin to bake in the hot, outback sun. As the endocarps dry, they explode. Just like the pod of a legume, the endocarp has fibers in its tissues oriented in opposing directions.  As the fibers dry, they contract and pull the endocarp apart. The dehiscence is sudden and explosive, sending seeds up to 2.5 m from the point of origin. Launching seeds away from the dung pile is beneficial to seeds: the special separation means that seedlings well be less likely to compete with one another.

But that is not the final disposition of Quinine Tree seeds. Each Petalostigma seed bears a small, oily food body, called an elaiosome, that is attractive to ants. Ants pick up the seed with its attached elaisome and carry it back to their nest. Once at the nest, the ants will remove and consume the elaisome and deposit the inedible seed in midden outside the nest. It is the ants that disperse the seeds to their ultimate site.

The association between emus, exploding endocarps, ants and Petalostigma pubescens probably represents one of the most complicated dispersal scenarios in the Plant Kingdom.

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

Further Reading: [1]

NOTE: Guest posts are invite only

An Intruiguing Relationship Between Ants and Cacti

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.

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.

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. Photo by Joseph j7uy5 licensed under CC BY-NC-SA 2.0

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]

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

Fern Ant Farm

Photo by Ch'ien C. Lee (www.wildborneo.com.my/photo.php?f=cld1505721.jpg)

An epiphytic lifestyle is no walk in the park. Baking sun, drying winds, and a lack of soil are the norm. As a result, epiphytic plants exhibit numerous adaptations for retaining water and obtaining nutrients. One of the most interesting adaptations to this lifestyle can be seen in plants that have struck up a relationship with ants.

An amazing example of one such relationship can be seen in a genus of epiphytic ferns called Lecanopteris. Native to Southeast Asia and New Guinea, their unique look is equally matched by their unique ecology. Using a highly modified rhizome, they are able to latch on to the branch of a tree. In species such as Lecanopteris mirabilis (pictured above), it's as if the fronds are emerging from a strange green amoeba.

However, it's whats going on underneath their strange rhizomes that makes this group so fascinating. These ferns literally grow ant farms. Chambers and middens within the amorphous rhizome entice colonies of ants to set up shop. In return for lodging, the ants provide protection. Anything looking to take a bite out of a frond must contend with an army of angry ants. Moreover, the ants provide valuable nutrients in the form of waste and other detritus.

These are by no means the only plants to have evolved a relationship of this sort. Myriad plant species utilize ants for protection, nutrient acquisition, and seed dispersal. It has even been suggested that the unique morphology of Lecanopteris spores is an adaptation for ant dispersal. Certainly one can imagine how that would come into play. Interestingly enough, this group of ferns has attracted the attention of plant enthusiasts looking for a unique plant to grow in their home. As such, you can now find many different species of Lecanopteris being cultivated for the horticultural trade.

Photo Credit: Ch'ien C. Lee (www.wildborneo.com.my/photo.php?f=cld1505721.jpg)

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

On Peonies and Ants

Photo by George Vopal licensed under CC BY 2.0

It is just about that time when peony buds burst forth and put on their late spring display. My mother loves her peonies and she gets very excited every year when they bloom. It's adorable. However, she has always been disgusted by the amount of ants the peonies attract. Indeed, many people all over the internet seem to feel the same way. Growing up, I always wondered why the ants seemed to swarm all over peony buds, so I decided to look into it a little deeper.

There are many sources out there that claim that peonies need ants in order to bloom. To me, this seems very maladaptive on the part of the peony. The genus Paeonia is represented in Asia, Southern Europe and parts of western North America. I am going to assume that the ant/peony relationship didn't start in the garden so it's roots have to be somewhere in the evolutionary history of the plant. What sense does it make for a plant to produce flower buds that excrete sticky sugars that keep them from opening until something cleans the sugars off? In fact, despite anecdotal reports, peony buds will open without ants. So then why does the plant bother to produce sugars that attract ants?

Interestingly enough, despite a good amount of searching, there is not a lot of research done on this subject but the answer to this question can come from looking at how ants interact with other plants and animals. Many plant species have special glands on their stems that produce sugary secretions which attract ants. It's not just plants either. Insects such as aphids and leafhoppers famously excrete honeydew that ants can't resist. In each of these cases, organisms are using the ants' natural tendency to guard a food source. The ants will viciously attack anything that threatens this easy meal.

It would seem to me that the peonies are doing just that with their flower buds. By secreting a sugary substance during their development, the plant are likely recruiting ants to protect the flowers, which for angiosperms, are the most precious part of the plant. It takes a lot out of a plant to flower and the threat of herbivory is ever present. If an insect tries to take a bite out of a bud, the ants quickly swarm and drive it off. It's a win-win situation. The ants get an easy, high-energy food source and the plant suffers less damage to its reproductive organs.

The scary part to me in researching all of this is plethora of information out there on how to get rid of the ants. People go through chemical after chemical to rid their peonies of ants when, in reality, the ants are some of the best friends a peony could have! So leave those ants alone and enjoy the free pest removal services they provide every spring!

Photo Credit: [1]

Further Reading:

http://www.youtube.com/watch?v=Gnm2nV_nwOk