Hidden Kingdom, Part Two

(Via:)
(Via: Livingroutes.org)

Common Name: Leafcutter Ants

A.K.A.: Genera Atta and Acromyrmex of Tribe Attini

Vital Stats:

  • Fungi grown by leafcutter ants come from the family Agaricaceae
  • Ant species can maintain their association with a specific fungal cultivar for millennia
  • Neither the ants nor the fungal cultivars can survive outside of the symbiosis
  • Some ant species are capable of completely defoliating a small tree in under a day

Found: Humid forests of Central and South America

Leafcutter Map

It Does What?!

Last week, we looked at leafcutter colonies, their various castes, and the impressively long lives of ant sperm. But obviously, leafcutter ants are known principally for one thing- cutting leaves. This they do on a grand scale, forming lines of thousands upon thousands of ants, dutifully toting chucks of foliage back to their colony. Why? To fertilize their fungus, of course! Much as we like to think of agriculture as one of the crowning achievements of mankind, the fact is, ants came up with it much earlier than we did. About 50 million years earlier, actually. (But they haven’t figured out how to deep-fry anything yet, so there’s that, I guess.)

caption (Via: Wikimedia.org)
The fungus is hungry.
(Via: Wikimedia.org)

When a young queen leaves her original colony to found a new one, she carries in her mouth a small piece of fungus to use as a starter culture (think yogurt or sourdough bread) for the colony’s gardens. Initially, she will care for this culture alone, but once the first generation of workers is born, they will take over the task from that point on. Since fungi don’t photosynthesize, they’re perfectly happy in a pitch-black underground garden, but they still need nutrients with which to grow, and dead vegetation is their food of choice. As the larger worker castes return with leaf (and flower) fragments up to three times their own mass, the minima gardeners clean away any outside fungal spores and chew the vegetation into smaller and smaller pieces. They then mix the shredded leaves with fungus and add the mixture to the garden. And, just for an extra fertiliser kick, they mix in their own faeces. Waste not, want not, right?

With all the workers coming and going, and so much foreign vegetation entering the colony, infections of the garden by competing fungal spores are inevitable, despite the ants’ best efforts. One such invader is the fungus Escovopsis, a parasite of other fungi, which can decimate a colony’s food supply and, in the case of young and vulnerable colonies, sometimes cause them to fail entirely.

caption (Via:)
I use the term “garden” loosely…
(Via: Marietta College)

But the ants have a secret weapon: bacteria. These adaptive little farmers actually carry around a ready supply of antimicrobial compounds right on their bodies. The bacterium in question, Pseudonocardia, grows directly on the ants’ exoskeletons and, researchers suspect, is nourished by a substance excreted through the ant’s glands. In return, Pseudonocardia produces a compound that the farmers can spread on invading fungus, killing it without damaging their food source. Symbioses within symbioses… and these are just the ones we know about.

Meanwhile, outside the colony, another fascinating parasite threatens the workers. Known as phorid flies, or ant-decapitating flies, you can probably guess why these things are a problem. Female phorid flies land on the backs of the larger worker ants as they travel to and from their leaf harvesting sites, laying eggs on the worker’s thorax. Once the eggs hatch, the larvae work their way into the ant’s head and start to eat the tissue surrounding the brain, eventually moving on to the brain itself (causing aimless wandering behaviour similar to that caused by the zombie ant fungus). Finally, the young parasites secrete an enzyme which causes the ant’s head to fall off completely, leaving them a convenient vessel in which to finish their development into adults.

caption(Via:)
They’ve evolved everything but the ability to look behind them.
(Via: Dayvectors.net)

Not to be outsmarted (by anything, apparently), leafcutter ants instituted a policy of defensive piggyback rides. Workers on the foraging path carry tiny minima ants on their backs as they travel. The minimae are too small to be useful hosts for the phorid fly, and so are able to fearlessly attack the flies as they approach, keeping the foragers safe. And not to lose an opportunity for increased efficiency, the little passenger will also begin cleaning the leaf fragment as the larger worker carries it home.

So there you have it. Leafcutter ants form colonies of millions, assign specialised tasks to different classes of citizens, grow their own crops, excel at problem-solving, and know how to use antibiotics. Next to humans, they form the largest and most complex societies on Earth. Forget robots and computers, people- if anything’s going to gain sentience and overthrow humanity, my money’s on the ants.

[Fun Fact: They compost, too. At least one leafcutter species maintains ‘outdoor’ waste heaps of discarded leaves and fungus. Special disposal workers (often old or unhealthy ants) turn the heap regularly to speed up decomposition.]

Says Who?

  • www.antweb.org
  • Marietta College Leafcutter Ant Page

  • Dijkstra & Boomsma (2006) Insectes Sociaux 53: 136-140
  • Evison & Hughes (2011) Naturwissenschaften 98: 643-649
  • Evison & Ratnieks (2007) Ecological Entomology 32: 451-454
  • Holman et al. (2011) Molecular Ecology 20: 5092-5102
  • Mueller et al. (2008) Evolution 62(11): 2894-2912
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Hidden Kingdom, Part One

(Via:)
(By: Tobias Gerlach & Jenny Theobald, Via: deepgreenphoto.com)

Common Name: Leafcutter Ants

A.K.A.: Genera Atta and Acromyrmex of Tribe Attini

Vital Stats:

  • 47 species; 15 in Atta, 36 in Acromyrmex
  • Atta ants have three dorsal spines and a smooth exoskeleton, while Acromyrmex ants have four spines and a rough exoskeleton
  • Less than 5% of new queens are able to build a successful colony
  • A maxima may have a head width of up to 7mm (0.28”), while a minima reaches less than 1mm (0.04”); mediae fall somewhere in between

Found: Humid forests of Central and South America

Leafcutter Map

It Does What?!

Once in a while, I come across a species that’s just so strange and interesting, a single post doesn’t seem to do it justice. With that in mind, welcome to part one of the wonderous life of the leafcutter ant.

Let’s begin at the start of it all – a new colony being founded. This happens when a fertile, winged female and several fertile, winged male ants (called drones) are born and grow to maturity. One day, the winged crew will fly away together and engage in what’s called a nuptual flight, where the female mates with several different males (up to seven in some species) while in mid-air. Having accomplished what is literally their only purpose in a short, glorious life, the drones promptly die, while the new queen scouts out a good place to start her colony. Finding it, she yanks off her own wings, never to fly again, as her body starts to break down her flight muscles, using the energy to produce eggs.

caption (Via: )
Nursery duty can be creepy when the babies all look like dead albinos.
(Via: Marietta College)

Ant reproduction is remarkable in that actual mating occurs only once in the queen’s life. The males of her nuptual flight together provide hundreds of millions of sperm that will be the basis for the entire colony to come. At the risk of sounding like a weirdo, ants have amazing sperm. A human sperm cell, under ideal conditions, can survive for up to five days. If they don’t get the job done in that time, they’re finished. The sperm of leafcutter ants, having been collected by the queen, can live for up to thirty years. That’s probably older than a lot of the people reading this. They can spend decades just waiting around in storage for the egg with their name on it. And that’s not all- they come armed. As in, chemical warfare. The seminal fluid of ants contains compounds that can lower the survival of rival sperm (from other drones) while not harming those of the ant they came from. Weaker sperm are thereby killed off early in the game. Of course, this kind of thing doesn’t go on for a long time. The storage organs of the queen contain their own fluid that will neutralize chemical weapons on the way in. Think of it as the metal detector at the door.

Right. So the queen has her new place picked out and the on-site sperm bank is up and running. Time to make a colony. What she needs first are workers. In a small chamber she’s excavated underground, she begins to lay large numbers of eggs. These serve two purposes, because the early hatchers will eat the late hatchers until the food supply gets built up. It pays to be a bit premature when you’re an ant.

(Via: Marietta College)
(Via: Marietta College)

Nearly every worker born to the queen over the life of the colony will be a sterile female, and each will belong to one of three major castes- minimae, mediae, and maximae. These castes will dictate both their size and function in life. Sensing the needs of the colony, the queen can actually control which type of worker she is producing. First come the minimae, which are the smallest caste and will principally tend the underground gardens which are the colony’s food source (more on ant agriculture in part two), as well as acting as nurse-maids for growing larvae. Next are the mediae, which are larger and act as the colony’s foragers, bringing plant material with which to fertilize the gardens, and defending against minor threats or obstacles in the troop’s path. Finally, once the colony has reached a population of several thousand, come the maximae, or soldier ants. These big brutes are up to thirty times the mass of a minima and do all the heavy lifting, carrying bulky items, moving big obstacles, and cutting tough pieces of vegetation. They’re also the last line of defence when something serious threatens the colony or the foraging parties. And because ants are all about organisation, within each caste, there are numerous sub-castes which are responsible for specific duties, depending on which species we’re talking about.

caption(Via:)
Little sisters are annoying no matter what species they are.
(By: Alexander Wild, Via: Alex Wild Photography)

Over time, leafcutter colonies can become impressively large, comprising over 5 million residents- the population of a major human city. It’s amazing to consider that these are kept running smoothly without central authority, technology, or the aid of written or spoken language. Not to mention opposable thumbs.

Tune in next week for a look at leafcutter agriculture, their interesting relationships with symbiotic fungi and bacteria, and why ants give each other piggyback rides to work.

[Fun Fact: The largest leafcutter ant colony on record required the excavation of approximately 40 tonnes (44 tons) of earth and contained thousands of different chambers.]

Says Who?

  • www.antweb.org
  • Marietta College Leafcutter Ant Page
  • den Boer et al. (2010) Science 327: 1506-1509
  • Dijkstra & Boomsma (2006) Insectes Sociaux 53: 136-140
  • Evison & Hughes (2011) Naturwissenschaften 98: 643-649
  • Evison & Ratnieks (2007) Ecological Entomology 32: 451-454
  • Holman et al. (2011) Molecular Ecology 20: 5092-5102
  • Mueller et al. (2008) Evolution 62(11): 2894-2912

Pitcher Plants: Sweet Temptation and the Slippery Slope

(Via: Wikimedia Commons)

Common Name: The Asian Pitcher Plant

A.K.A.: Genus Nepenthes

Vital Stats:

  • Over 130 species in the genus
  • The vast majority of species have extremely narrow ranges of only a single island or small island group, and are considered threatened
  • Most recently discovered (2007) was Nepenthes attenboroughii, named for Sir David Attenborough, who is fond of pitcher plants

Found: Mountainous regions of Southeast Asia, Oceania, and Madagascar

It Does What?!

Plants have evolved a variety of different ways to deal with growing in nutrient-poor soils. Some become parasitic, some develop close symbiotic relationships with bacteria or fungi, and some of them… well, some of them just start eating animals.

Lizard: makes a nice, light snack.
(Via: Wikimedia Commons)

One group of plants that went this route are the Asian pitcher plants (not to be confused with the not-closely-related New World pitcher plants, which tend to have tall, flute-like pitchers). These smallish, climbing plants use highly modified leaves to form what are essentially external stomachs, complete with the plant’s own digestive fluid. These pitchers, which vary in size from one species to the next, have extremely slick, waxy inner walls. When visitors come to eat the nectar produced on the lid (or “operculum”) of the trap, they lose their footing and fall into the liquid below.

That liquid is actually a pretty complex mixture; it’s divided into two phases, like oil and water. The upper portion is mostly rainwater, but has been laced with a compound that makes it more viscous, preventing winged insects from just flying away, as they could from pure water. The trap’s lid actually functions to prevent too much rainwater from getting inside and diluting the fluid too much. The lower portion of the liquid is a digestive acid capable of breaking down flesh into useable molecules (particularly nitrogen and phosphorous), much like our own stomach acid. Analogous to our intestines, the lower inside surface of the pitcher is covered with special glands that absorb suspended nutrients.

Most of what gets caught in pitcher plants is about what you’d expect- winged insects, spiders, beetles, small scorpions. But occasionally, some larger animals find their way in. Things that should have known better, like frogs, lizards, and even birds. Arguably, these plants are doing evolution a favour by taking out any bird dumb enough to fly into its own watery grave. And yes, to answer your next question- they can eat rats, but only a single species has been documented to do this. Nepenthes rajah, the largest of all pitcher plants, has pitchers which grow to a height of nearly half a metre (1.6’) and hold up to three and a half litres (1gal.) of fluid, most of which is digestive juice.

Interestingly, pitcher plants have formed symbiotic relationships with several of the same types of creatures that it otherwise preys on. Nepenthes lowii, for example, provides nectar to a tree shrew. Instead of falling in and being digested, the shrew treats the pitcher as its personal toilet, thereby providing the plant with most of the nutrition it requires.

In one end and out the other.
(Via: Wikimedia Commons)

Other species form alliances with groups of carpenter ants. In exchange for a steady supply of nectar and a place to live- in this case a hollow tendril- the ants basically act as the plant’s evil henchmen (apparently a specialty of ants). When prey that is too large to be easily digested falls into the trap, the ants remove it, rip it to shreds, and then throw the bits back in again.

How’s that for a brilliant piece of evolution? Not only did these plants grow an external stomach… they get ants to chew their food for them.

[Fun Fact: Some pitcher plants primarily survive by digesting leaves that fall from trees into their traps – the ‘vegetarians’ of the carnivorous plant world.]

Says Who?

  • Bonhomme et al. (2011) Journal of Tropical Ecology 27: 15-24
  • Clarke et al. (2009) Biology Letters 5: 632-635
  • Krol et al. (2012) Annals of Botany 109: 47-64
  • Robinson et al. (2009) Botanical Journal of the Linnean Society 159: 195-202
  • Wells et al. (2011) Journal of Tropical Ecology 27(4): 347-353
So big it makes them vaguely uncomfortable.
(Via: Wikimedia Commons)

The Zombie Apocalypse: Already Underway

(Via: this site)

Common Name: The Zombie-Ant Fungus

A.K.A.: Ophiocordyceps unilateralis

Vital Stats:

  • Whole “graveyards” of 20-30 ants may be found within a single square metre
  • Telltale bitemarks on fossil plants suggest this fungus, or a related species, may have been in operation for the last 48 million years
  • Host species is the carpenter ant Camponotus leonardi

Found: Tropical forests throughout the world

It Does What?!

Despite all the advances of modern neuroscience, the fact is, human understanding of brain chemistry and its manipulation still has a long way to go. Much to the chagrin of those plotting world domination, we won’t be chemically controlling each other’s minds any time soon. How embarrassing then, that a mere fungus seems to have perfected this technique. Almost fifty million years ago. Scooped again, humanity.

It begins with an ant walking along the ground, deep in a tropical forest somewhere. This ant, Camponotus leonardi, lives high in the trees, but must occasionally come down to cross from one tree to another when there is a break in the canopy. As it walks, a minute fungal spore drifts down from above and lands in its back, unnoticed. The unseen spore springs into action, producing an enzyme which breaks down the ant’s exoskeleton just enough to allow a fungal hypha, like a tiny root, to enter. The host’s fate is now sealed.

This is your brain on ‘shrooms.
(Via: Flickr, by Alextkt)

While the ant climbs back up into the canopy and goes about its business, the fungus grows through its insides, breaking down and consuming the non-vital soft tissues as it goes, keeping the animal alive even as it is being eaten. Soon, the fungal tendrils reach the brain and begin to produce chemicals which affect the host’s behaviour in very specific ways. First, it will experience convulsions that cause it to fall out of its tree. These will continue periodically, preventing it from returning to its colony. Over a period of hours, the ant will then wander, erratically and aimlessly, over the ground and low-growing plants.

This is where the precision of the fungus’ mind control becomes truly impressive. At solar noon, when the sun is highest in the sky, the infected ant will abruptly climb the stem of a small plant and find a leaf pointing north by northwest at a height of 20-30cm above the ground. Yes, really. No one knows how this jaw-dropping specificity is accomplished, but it’s what the fungus wants, providing a temperature of 20-30 degrees Celcius (68-86F) and a relative humidity of around 95%. In cases where ants were experimentally moved to different heights or orientations, the fungus was unable to reproduce properly.

What the fungus wants, the fungus gets.
(Via: Wikimedia Commons)

Having found the perfect leaf, the zombified ant will go to its underside, find a major leaf vein, and just bite down on it as hard as it can. The fungus has already destroyed the muscles required to release this grip, and so there the ant stays, slowly dying over the course of the afternoon. Once its victim has been dispatched, the fungus grows toward the leaf, further anchoring itself to the plant. Around a week later, the parasite completes its horrifying circle of life by growing a fruiting body, similar to a mushroom, from the back of the dead ant’s head. This will open to release thousands of tiny spores, raining down over any potential hosts which may be walking below.

While the fungus is able to infect other, closely related, species of carpenter ant, it has less precise control over these hosts and isn’t always successful in getting the ant to do its bidding, suggesting that even minor variations in brain structure can stump it. So we’re probably safe from the fungal zombie apocalypse. At least for the time being…

Says Who?

  • Andersen et al. (2009) American Naturalist 174(3): 424-433
  • Hughes et al. (2011) Biology Letters 7: 67-70
  • Hughes et al. (2011) BMC Ecology 11: 13
  • Pontoppidan et al. (2009) PloS ONE 4(3): e4835

EVOLUTION TAG TEAM, Part 1: Acacia Domatia

The first in an ongoing series of biology’s greatest duos. (Here’s Part 2 and Part 3)

Home, Sweet Home.
(via: Flickr)

Common Name (Plants): Bullhorn Acacias, Whistling Thorns

  • A.K.A.: Acacia cornigera, Acacia drepanolobium, and several other Acacia species

Common Name (Ants): Acacia Ants

  • A.K.A.: Pseudomyrmex and Crematogaster species

Found: Central America (Bullhorn Acacias) and East Africa (Whistling Thorns)

It Does What?!

Life as a tree is tough, particularly when you live in a part of the world that’s home to the biggest herbivores on Earth and happen to have delicate, delicious leaves. Such is the case for the African acacias. Without sufficient defences, they’d be gobbled up in no time by elephants, rhinos, and giraffes. The trees are known for having huge, sharp thorns, but even that’s sometimes not enough; the lips and tongues of giraffes are so tough and dexterous, they can often strip the leaves right out from between the thorns. So what’s a stressed acacia to do? Recruit a freaking army, that’s what.

Pseudomyrmex ferruginea: the giraffe’s worst enemy.
(Photo by April Nobile)

A few species of acacia in both Africa and Central America (where the herbivores are smaller, but no less voracious) have developed a symbiosis wherein they enjoy the services of ant colonies numbering up to 30,000 individuals, tirelessly patrolling their branches 24 hours a day. Should a hungry elephant or goat wander up and take a bite, nearby patrol ants will call in reinforcements and soon the interloper will be utterly overrun with angry, biting ants. What’s more, the protection extends beyond just animal threats. The ants will go so far as to kill other insects, remove fungal pathogens from the surface of the tree and even uproot nearby seedlings because, you know, they might eventually steal some sunlight from the beloved acacia.

“Trespassers Will Be Drawn and Quartered”
(via Wikimedia Commons)

So what do the troops get out of this? Quite a bit, actually. In ant-protected acacias (‘myrmecophytes’, they’re called), the thorns that normally grow at the base of a leaf swell up. In the Central American species, they grow into something that looks like a bull’s horn (hence their common name), while the African ones become more bulbous. These specialized structures, called domatia, are hollow inside and serve as very convenient housing for the ants. What’s more, the trees produce not one, but two different kinds of nourishment for the colony- regular, and baby food. The adult ants will feed from a sweet liquid exuded by nectaries on the branches. Meanwhile, on the tips of the tree’s leaflets, small white structures called Beltian bodies are formed which are high in the protein every growing child ant-larva needs. These are collected by workers and inserted right into the larval pouches, to be eaten before the ants are even fully formed.

The Bullhorn Acacia, now with more Beltian bodies!
(via Flickr)

Sounds like the perfect partnership, right? Usually, yes, but in nature, a symbiosis is only a symbiosis until one side figures out how to take advantage of the other. From the ants’ side, for example, any energy spent by the tree on reproduction is energy not spent on new homes and sweet, sweet nectar for them. Therefore, the ants will sometimes systematically nip all the flowers off the tree as it attempts to bloom. They’ll also prune the acacia’s outward growth if those new shoots may come into contact with a neighbouring tree, allowing invasion by another ant colony. Conversely, if herbivores become scarce and the acacia no longer requires such a strong protection force, it will begin to produce fewer domatia and less nectar in a move to starve some of the ants out. This has been shown to actually be a bad strategy for the acacia, since the soldiers, not to be outsmarted by a tree, turn to farming and begin raising sap-sucking insects on the bark, thereby getting their sugar fix anyway. And so it goes, oscillating between advantageous partnership and opportunistic parasitism… like so many things in life.

The roomier, more spacious African domatium.
(Image by Martin Sharman)

[Side note: While I’ve never personally encountered ant-acacias, I have disturbed an ant-protected tree of another family in the rainforests of Guyana, and can attest to the fact that the retaliation was both swift and intense. I was in a small boat at the edge of a river collecting botanical specimens, and I nearly jumped in the river to escape the onslaught. Don’t mess with ants.]

Says Who?

  • Clement et al. (2008) Behav. Ecol. Sociobiol. 62: 953-962.
  • Frederickson (2009) American Naturalist 173(5): 675-681.
  • Huntzinger et al. (2004) Ecology 85(3): 609-614.
  • Janzen (1966) Evolution 20(3): 249-275.
  • Nicklen & Wagner (2006) Oecologia 148: 81-87.
  • Stapley (1998) Oecologia 115: 401-405.