Friday, September 5, 2014

The Ant Word

WHAT’S NEW: last update 05/09/2017 [click here] 


STOP PRESS!  Go HERE to order our great new field guide, ANTS OF SOUTHERN AFRICA, in press and for immediate delivery by airmail or courier ... 

INTRODUCTION 

We’ve put this website together to help anyone interested in the natural systems of South Africa to identify these tiny but fascinating animals. There’s more about recognizing ants in the pages below, with some fairly basic stuff about these insects and their biology. Our ants are as much a part of our fantastic wildlife heritage as lions and rhinos, fish eagles and blue cranes, mighty yellowwoods and king proteas. The ecological role they play is immensely important for the health of our natural systems. Ants are essential seed-distributors to thousands of charismatically-beautiful fynbos plants; they are pollinators and recyclers, too, and also vectors in the spread of many plant predators. However, just as there are plant invaders that threaten our natural systems there are insect invaders, too, and we’re keeping an eye on some pretty murderous invasive ants. 
We’ve based this revised site on our Field Guide, ‘Ants of Southern Africa’, the first and only field guide to our local ants; see link above if you would like to buy one online.

If you have any info, queries or comments please use the Comments box below. 
The Argentine ant – Linepithema humile – is one of the most devastating invasive organisms on Earth. It may pose a very serious threat to the health of our fynbos and other ecosystems ... but do we know where they all are?

Introduction to ‘Ants of Southern Africa’

Ants grabbed me when I was but a spotty teenager. Some grabbed me literally with snapping mandibles, and I discovered that they can seldom really hurt you. Years later, when my own barefoot kids would start hopping around on the sandy veld paths shouting ‘Biting ants!’, I’d encourage them to stand quite still and tell me, honestly, whether the ants were hurting them.
‘No,’ they’d reply, after a moment or two, ‘but they tickle!’
Pugnacious ants, Anoplolepis custodiens, have held few fears for them ever since. And me? They’re my favourite ants. Bold, tiny little girls, quite unafraid of human beings.
Most people, told that I like ants, ask me at once how to get rid of them. The answer is, of course, ‘you can’t.’ You can soak your kitchen, your fruit trees, your garden beauties with litres of insecticide, but after a few days the ants will be back. It’s estimated that 20% of the dry land biomass on our planet consists of ants. Think about how many humans live in your house, now estimate the number of ants. Scary? Not really. They collectively destroy more insect ‘pests’ than you ever could do with a spray can. They outnumber us by uncountable trillions; we will never get rid of them.
In 1961 the great entomologist Sydney Skaife published a book titled The Study of Ants. It was the first-ever ‘citizen science’ book about Southern African ants; Skaife’s opening lines read: “Ants ... have been collected, preserved and described in minute detail, and classified in families, genera, species, subspecies, races and varieties, and they have been given long polysyllabic names. They are abundant almost everywhere, of considerable economic importance, and offer ... challenging problems to entomologists, yet few are engaged in the study of these fascinating little insects, apart from the description and naming of them.” 
I was given a copy of this wonderful book at the age of 15, and soon I was irritating my family with colonies of ants in plaster nests on ant-tables, using Skaife’s recommended methods. The ants were forever escaping, or being attacked by Argentine ants, or having other interactive adventures with my family members. Within a couple of years I was persuaded to donate my thriving, seething masses to the great Skaife himself. And that led to several visits to his laboratory at Tierboskloof, in the mountains above Hout Bay. 
Skaife was a founder of the Wildlife Society of SA (WESSA); almost single-handedly, he created the Cape of Good Hope Nature Reserve – the best-preserved section of the Table Mountain National Park. A natural teacher, he and his ants had me in thrall, even though I never followed him into formal science.
By 1981 we were living in Kleinmond. Professor-to-be William Bond had become a close friend; I was trying to learn everything I could about fynbos and the Kogelberg, the veld where I had collected my first ants, à la the methods of Skaife. One day William showed me some seeds with a tiny white attachment. ‘Rudolf Marloth, a citizen botanist, wrote that at least six fynbos species have seeds like this, that deliberately attract ants; the ants bury the seeds, safe from fire,’ William explained. Within a few weeks we had identified over 2 000 fynbos plants that use ants to bury their seeds ... academic papers followed, of course. 
        By 1991 I had put together a little black and white booklet, Ants of the Western Cape; Hamish Robertson of Iziko museums, another great entomologist, had helped me correct my worst amateur bloopers. The second citizen science book about Southern African ants was born. It was a very poor replacement for Skaife, though.
Twenty-five years later that was still all there was. There were two books, both out of print. There are a few relatively academic websites that, valuable as they are, specialise in pictures of very dead ants. They hold almost no information about their habits, their nature, their basic biology, and they all have inadequate and often incorrect information about their distribution. In 2016 the experts Brian L. Fisher and Barry Bolton published their impressive Ants of Africa and Madagascar: an academic book that will take your interest to a higher level, but that has little info about your local species.
In 2014 I joined Tony Rebelo’s iSpot (he’ll say it wasn’t his, but don’t believe him – in those days it was) and I soon found that there were lots of people who were photographing ants and posting iSpot observations about them – amongst lots of other extraordinary things. But there’s the rub: many of them could not identify their ants. There was simply  no ‘citizen science friendly’ guide to our very rich ant fauna, and there appeared to be no one prepared to put one together. Yet here were nearly 2000 photographs of ants from all over Southern Africa taken by iSpotters, many of them of a very high quality. Here was a resource that should be presented to a wider public; what better way than a printed field guide. 
Naïvely, I grabbed the tiger by the tail. Distant academic snarls suggested that such a thing should not be done without an expert’s benign, watchful eye. Yet even the experts, even the world’s greatest, do not agree upon the identification of species in some of the largest and most prominent of Earth’s genera of ants. They can’t identify them – at least, not to every other professional’s satisfaction – without lots of interesting, stimulating disagreement.
More than fifty years ago Skaife recognised the potential contribution to his discipline of citizen science – all science, after all, started with ‘citizen science’, with our human curiosity directed at the world around us. So, I have plunged in where greater angels have feared to tread. There are errors in this account: many genera are screaming for taxonomic revision. Here and there I might have a wrong photo, but please bear with me: precise help has not been easy to find. I absolutely acknowledge that many ants – the tiny ones, the LBJ’s of the ant world, especially – can only be identified under high-powered magnification, equipment that is not found in everyone’s kitchen. 
But there are hundreds of others – who could mistake the magnificent hairy, tawny gaster and gaping jaws of a Karoo balbyter? Anyone with good vision (they are all small), good sense (the subfamilies are pretty easy, actually) and enthusiasm  can learn about ants and their world; the more enthusiastic citizen science we can focus upon ants, the more – much, much more – we’ll learn about these extra-ordinary insects, their habits, behaviour, what they eat, how they interact with other organisms, and especially where in our Southern African region the different species are to be found. That’s the info the taxonomists trying to ID them need, and that’s how we can help them.

Ant Biology

The Ant Colony


Ants belong to the order of insects known as Hymenoptera, that also includes sawflies, bees and wasps. Some bees and wasps are social insects; ants, at the top of the tree of insect life, are all social (by rough analogy, if these were all primates, bees would be furry lemurs, wasps would be monkeys and ants would be apes). There are no solitary ants.
Ants, in the family Formicidae, are believed to be descended from wasps, and they share many characteristics with them. However, they have many ‘stand alone’ features that all ants, and only ants, share. 
* Because all ants are social, if you find one ant there will always be others from the same colony somewhere nearby.
* Almost all the worker ants you’ll ever see are females. Male ants or ‘drones’ have short, idyllic lives, where they are born, hang around the nest being pampered and fed, and then fly off to find females (‘alates’) to mate with. After mating they are unable to care for themselves, and they die.
* The colony usually has a queen or several queens who do the reproductive work, although there are species of ants where a mated worker or workers may share or take on the reproductive role.
The queen, technically known as a gyne, starts her adult life as an alate, with two pairs of wings which she uses on her mating flight (queens of the Dorylinae, however, are quite different). After mating – and she only mates once in her life – she drops her wings, finds a suitable nesting site and lays her first eggs. The eggs are small and white or sometimes yellow or pinkish. They hatch into blind, legless larvae that are completely helpless, and unless constantly fed and groomed they die. The queen tends these through the larval and pupal stages until the first workers eclose from pupae (eggs hatch, pupae eclose). She uses nutrients from her now-redundant wing muscles to nurture her first brood. The first workers are often smaller and paler than subsequent generations, but once they have eclosed the queen can relax and look forward to a long life (20 years or more, sometimes!) of laying eggs and being fed and groomed by her workers, who take over the gathering of food as well as the care of the eggs, larvae and pupae.
Note that in some subfamilies the larvae spin cocoons, within which they pupate, while in others the pupae are naked. In both instances they have to be helped to eclose by their older sisters, who tear the pupal ‘skin’ and/or silk off the helpless newcomers. Newly eclosed individuals have incompletely sclerotised chitinous exoskeletons and are often pale yellow or grey in colour; they are known as tenerals or callows, while the discarded pupal materials are known as exuviae.
In some genera the workers can also have different castes: minors and majors, dimorphic or polymorphic; and in some cases even ‘intercaste’ sexed or ‘ergatoid’ workers, who can be queens or males.











ANT BIOLOGY: Ant Nests



Ants construct their nests in a remarkable variety of ways. Even the ‘simplest’ hole-in-the-ground nests reveal a remarkable complexity. Experiments pouring (with great care) molten aluminium down ant nests (experimenters insist the nests were unoccupied!) reveal an extraordinary structure, which when carefully excavated can be a thing of some beauty. Chambers are linked by tunnels which often reach a considerable depth.
Hole-in-the-ground nests may have simple hole entrances without adornment, or these might be surrounded by a perfectly circular cone of sand grains brought up from the excavations below. Some ants construct an elaborate vertical turret as their entrances, perhaps to prevent rainwater running into the hole; other ground dwellers expropriate a few floors of a termite mound.
Other ants simply excavate a few tunnels or chambers under stones; some burrow into wood or take over the tunnels left by other boring insects, while others—very thin ants such as Tetraponera (Slender ants)—hollow out the soft interior of Watsonia or Aristea stems to house themselves. The rare Bee-legged boring ant, Melissotarsus, is a true carpenter and bores into the heartwood of certain plants and, once there, never leaves again, living off the secretions of plant-sucking coccids that it introduces into the plant.

The dedicated spinners, Tailor or Weaver ants (Oecophylla), famously use their silk-producing larvae to tie tree leaves together to make arboreal nests.                              Many Polyrhachis, the Spiny sugar ants, also use their larvae to spin cup-shaped walls of silk, grass and soil around their nest entrances, often lining these with silk for some distance into the ground.
Messor ants, seed harvesters, often create a ‘midden’ of chewed up vegetable matter around their nest entrances; if this is undisturbed for long enough it may become bound into mounds by vegetation—or it may all blow away in the next strong wind.
Their cousins, Myrmicaria or Droptail ants, leave similar mounds of detritus around their entrances, occasionally moulding the mixture of dead stuff and soil rather crudely into entrance mounds or even tunnels. Monomorium fridae, the vicious Frida’s fierce timid ant, builds similar detritus mounds which it defends vigorously with tooth and sting.
A different kind of earthen-and-stick mound is built by the Hairy sugar ant, Camponotus niveosetosus. Uniquely in the genus, it constructs these nests in reedy tussocks in wet, marshy ground, where the nest material  becomes bound together by mossy or fungal strands.
Camponotus bianconii builds carton-like chambers under the loose bark of dead trees, very similar to the carton produced by Cocktail ants, Crematogaster spp. These build unique nests of chewed up-vegetation glued together with their own saliva. The result resembles crude brown papier maché. The nests, that may house hundreds of thousands of ants, can be high up in bushes and trees or low down around the base of shrubs—but never on the ground. How they survive fire in fire-prone vegetation is one of the unsolved mysteries of the ant world.
And then there are a few—just a few, but famous— species that nest in the hollow thorns and galls on trees, where they may help defend the tree by vigorously attacking any browsers who should incautiously push noses and tongues too close.















ANT BIOLOGY: Ant Behaviour;  Ecological Role of Ants

In 1925 the South African naturalist and poet Eugène Nielen Marais published a series of articles in Huisgenoot magazine titled ‘Die Siel van die Mier’ (‘The Soul of the Ant’, often misquoted in English as ‘The Soul of the White Ant’.) The articles were shamelessly plagiarized in 1926 by the French Nobel Laureate Maurice Maeterlinck*, who should have known better, in his La Vie des Termites. Marais also put forward the hypothesis that the termite colony was in effect a single organism. This concept was also pinched by Maeterlinck and, more recently, has even appeared in the work of Richard Dawkins. But what has this got to do with ants?
[* The poet Stephen Gray has tried to debunk this by suggesting that Marais in fact stole Maeterlinck’s ideas—but I ask you, how could a rural Transvaler pre-publish by one year the manuscript of a French professor who in fact admitted that he had never seen a termite in his life? Steve surely forgot that there was no fax/email/internet in the 1920s, a somewhat anachronistic error of judgement ...]
The first problem was Marais’s title. I’d be a rich man if I had a dollar for each time that people, hearing that I am interested in ants, have winked and said, ‘Ah! The soul of the (white) ant!’ A comprehensive inability to distinguish between termites and ants has apparently been engraved on the human genome ever since, and, I’m sorry, Marais, all his imitators and his subsequent book (on the right) are absolutely to blame. 
The second problem is the proposition that the termite colony is a single organism. The idea may seem to have merit, even when trans-ferred to ants and bees, but it has unfortunately led to a proliferation of very problematic, quasi-mystical ideas about collective intelligences, ethereal souls, and evidence of the Hands of Sundry Deities—all gushingly described in the muddier recesses of the internet.
Marais was writing decades before pheromones were discovered. First identified in 1959 and present in all animal phyla including humans, pheromones are chemical substances that you secrete in various ways and that consequently influence the behaviour of your fellows. Ants and other social insects use pheromones to maintain colony unity. Recognition pheromones help ants to tell friend from foe; ants in the same nest constantly tap each other with their antennae, detecting these ‘friendly’ pheromones. If an enemy is detected ‘attack’ pheromones might come into play, and these stimulate other ‘friendly’ ants to join the attack.
Trailing behaviour in many ant species is also pheromone-driven. After a foraging ant has found a food source she returns to her nest, frequently tapping her gaster on the ground and thus marking the route to the food with a pheromone. The particular pheromone triggers a ‘move towards me’ response in other ants, and in no time a trail to the food source develops. Each individual on the trail constantly reinforces the pheromone markers. As the food declines fewer and fewer ants use and mark the trail, until eventually the pheromone has evaporated, no ants are renewing it, and the trail disappears.

Not all ants exhibit trailing behaviour. Many forage individually or in pairs; others such as harvesters who collect randomly-fallen seeds from all over the place, might come together near the nest in a vague trail. As a general rule ants that live in larger colonies—perhaps from many thousands of ants rather than several hundreds—are more likely to ‘trail’ than those in smaller.

        Pheromones similarly guide most ant behaviour, from the treatment of the queens and the brood to defence of the colony. The individual ant is not really capable of making any individual decisions, nor does the queen radiate some sort of mysterious, ethereal bonding vibe—her pheromones smell, that’s all.
Trophallaxis is another important process that binds the colony. Most highly developed in social insects such as ants, termites, wasps and bees, this is the mouth-to-mouth transfer of regurgitated food—essentially fluids—from one individual to another. Ants use a particular pattern of antennal stroking to induce a fellow ant, or a queen or a larva, to accept their regurgitated droplet of nectar. Other patterns are used by hungry ants to induce regurgitation from another that has filled her stomach. In many ants a further process has evolved: some individuals get fed and fed and fed until they swell up into living storage vessels—the so-called ‘honeypot ants’, whose store of nectar can be tapped at any time by antennal manipulation.
Although I read somewhere that this complex process was proof-positive of the theory of creationism (but why?), it’s clear that in social animals that don’t have hands other ways have evolved to carry food to their friends and young. Many non-social birds or mammals regurgitate food for their young, too.
Ants have evolved a variety of ways to deal with threats to the colony. These might come from ant-eating mammals, ant-eating ants, or various other sources. The ‘attack’ pheromone is freely used and results in instant action. Once activated, ants use their mandibles (biting is the most common form of defence/ attack in most animals, after all!), but they might also use stings (Ponerinae, Myrmicinae, etc), chemical repellants (Dolichoderinae, many Myrmicinae) or, amongst the Formicinae, powerful sprays of formic acid from the ‘acidophores’ which they have instead of stings. Ants are usually, but not always, fiercely antagonistic towards ants of other species, and very frequently just as antagonistic towards ants of the same species. It might be ‘nest smell’ caused by subtle differences in pheromones from one colony to another that triggers these responses.
Ants reproduce their colonies in two main ways:
1. By nuptial flights. The alate (winged) females and males fly out of the parent nests and, using pheromones, they attract the opposite sex from a different colony, and they mate. The male dies; the new queen drops her wings and finds a sheltered place for her new colony. She lays a few eggs and when these hatch she raises them to become her first workers. She does not eat during this time, living off the fatty reserves supplied by her now-redundant wing muscles. Nests established in this way are highly nationalistic and usually hostile to all other ants, including their own species. The whole nuptial flight process is extremely dicey and the chances of failure are extremely high. 

2. By budding. There is more than one reproductive queen in the nest. When a good food source is discovered, some of the extra queens migrate out with the workers and start a new branch of the colony at the food supply. The colony remains interlinked and its members are accepted by all branches. The colonies can grow into enormous super-colonies spread out over many miles, and with countless trillions of individuals.
Humans are always and inexplicably drawn to the bizarre, but I’m sorry to say that some of the weirdest forms of ant behaviour, such as chain gangs, river rafting, slavery, suicide bombing and vampirism (where ants suck the ‘blood’ of their own larvae) are not, as far as we know, found in our region.








Ecological Role of Ants


It has been estimated that although ant species comprise only about 2% of all arthropod species, they nevertheless make up some 30%+ of the terrestrial arthropod biomass—and 20% of the terrestrial animal biomass on planet Earth. Their numbers are uncountable and they are present on every continent except Antarctica. In some areas their nests are so extensive that their role in aerating the soil is more effective than earthworms.
As part of the food web, ants are important cleaners, consuming and recycling tons of decaying plant and animal material. They are massively important controllers of various insect ‘pests’, including flies, termites, etc. Not only do they eat a wide variety of living and dead organic material, they themselves are food for a huge variety of other creatures.
On a regular basis, ants are eaten by:
* Other ants. There are many specific ant predators of ants across a range of subfamilies, ants who take advantage of superior numbers or superior attack mechanisms to exploit the rich resource created by their distant relatives.
* Spiders. There are many spiders who imitate ants, and many of these use their disguises to capture and eat ants.
* Ant lions and ant-worms. Few can be unfamiliar with the conical craters made by these voracious ant-hunters, natural pitfall traps from which there is no escape for the unwary ant.
* Robber and other beetles. There is a wide variety who are specific ant hunters; there are ant-eating mantids, too.
* Reptiles and amphibians. Many lizards, skinks, frogs and toads eat ants. Curiously, despite extensive observation, I have never seen chameleons take ants.
* Birds. Huge numbers of ants are eaten by birds. If you have a bird feeder in your garden, check if there are ants around the feeding tube. If so you may see white-eyes, weavers, bulbuls and even sunbirds picking off the ants.
* Mammals. Most mammals that eat ants are specially adapted, such as aardvark. Analysis of the stomach contents of dead aardvark has revealed that their favourite food is Anoplolepis ants, not termites, as was once thought. Less specialised feeders such as baboons will readily turn over stones and consume the ants’ brood—larvae and pupae—that they find there. Honey badgers are also major consumers of ants. 
Humans who inadvertently consume ants, e.g. on that piece of cake left out overnight, usually spit them out at once. That sharp taste is caused by the formic acid that all ants contain in their bodies—but is in fact quite harmless.
* Plants. Our wide variety of sundews certainly consume a fair amount of ants, whose empty shells can often be found sticking to their leaves.
Ants are of course famous ‘farmers’ of coccids and aphids, insects that convert plant juices into nectar that is regularly ‘milked’ by ants from these economically-important, destructive pests. Some ants even practice a sort of animal husbandry, where they bite off the wings of winged aphids to prevent them escaping; some can even secrete chemicals that prevent the regrowth of wings, or that make the aphids tamer and more amenable to manipulation. The ants also guard their ‘cows’ against predators like ladybirds; they build shelters over them too, though these may be more for the ants’ own benefit.
Some ants are farmers of a different kind, collecting suitable plant material that they mulch within their nests and then sow with fungal spores, producing unique fungal food for the colony. Others are direct predators of seeds of a wide variety of plants, and such species often have majors with massively-powerful jaws that act as living nutcrackers, to crack open tough seed casings.
Another well-known interaction with plants is performed by ants that live in the thorns or galls of certain Vachellia (= Acacia). When browsers tug on the plants the ants swarm out and attack the intruding tongues, or even the soft membranes in elephants’ trunks.
Finally, in our very own fynbos, some ants are directly responsible for the distribution of seeds of over 1200 plant species, a process known as myrmecochory (‘carried by ants’). The nut-like seeds have a sweet, fatty attachment called an elaiosome that imitates the ants’ attractant pheromone. The ants find the seeds and drag them into their nests, where the elaiosomes are consumed. The hard, smooth nutlets, which cannot be gripped by the ants’ mandibles, remain safely underground in the nest until germinated by the passage of the next veld fire. Some fynbos rarities, long thought to be extinct, have reappeared more than fifty years later thanks to seeds which have survived underground in ant nests.
The latter process is threatened by invasion by trail-running ants. Trail-runners such as Argentine ants, some Pheidole, Technomyrmex, Lepisiota species, etc. do not carry food-objects into their nests; they consume them where they find them, taking the juices home in their stomachs. The seeds are left lying out in the open, to be predated by rodents and birds, or destroyed in the next fire.

Anatomy; Collecting ants for Identification

Ant Anatomy

Every scientific discipline loves its jargon, and entomology is no different. We’ve tried to keep the language simple in this website but there are a few terms you need to know. Ants’ bodies are substantively different from mammals and they have body parts that don’t have convenient common names. The drawing should make most of this clear. There are other parts with names but these are the most important.






Collecting ants for identification



We strongly advise users to sign up to iSpot, a great way to get your ants identified, or to help others ID their photos. You can sign up here

Although we have tried to convey the colour, shape and something of their habits in our illustrations of the various ant species below, if you’re serious about learning how to identify them there is no substitute for collection and examination. Ants are usually very, very small animals, however, and in order to examine them you’re going to need a good hand lens or, even better, a mini-microscope to be able to see their full range if identifying features.

Hand-held mini-microscopes of 60x magnification are very useful. The microscopes have powerful LED lights and use pill-batteries. 
The microscopes are unfortunately no longer available from me, but you might find them in a gadget shop.
There are two other essential items you will need. One is a drinking straw [or a few straws of different diameters] with a gauze filter securely taped over one end. The other is a supply of small Ziploc® bags. When you go out collecting you might want to unzip your bags before you start, as you often have to move fast to catch your ant and the bags sometimes resist opening at the wrong moment. 



With the gauze-covered end of the straw in your mouth, quickly suck up the ants and blow them gently into the bag; quickly zip it up making sure that there are no ants caught on the zip. Remember at all times that ants are small, fragile creatures and your enormous fingers – compared to an ant – will surely damage them; blowing them into the bag too hard may also injure them.
Bothroponera pumicosa in the bag! Once trapped in the plastic you can study the animal at your leisure with your mini-microscope
When the ants are in the bag it’s easy to study them with the mini-microscope, often most conveniently by holding the bag against a white notebook page and holding it up to your eye. The sides of the bag pin the ant down without squashing it. If your ants are hot they will move very fast; it’s often best to take them home and put the bags into the fridge for an hour or so. This won’t kill them but it will slow them down. You can use the sections below to try to identify your ants, or you can try the iSpot key to Western Cape Ants. Finally, if all else fails the ants in the bag can also be easily posted away for identification, and you are welcome to send them to P Slingsby at P.O.Box 303, Muizenberg 7945. I’m always looking for ants to draw and yours will be most welcome, even if ‘dead on arrival’. In fact it’s best to kill the ants before sending them; the most humane method is to put them, bag and all, in your freezer for a few minutes.

Identify ants: Key with links

1. See our comprehensive Catalogue of all the species which have appeared on iSpot, with illustrations and/or links to pics of more than 100 species

2. For a brand-new illustrated key to the 68 Southern African genera [still being developed] go here

3. Scroll down through the illustrations in the links below. These have been arranged more or less in order of size, from smallest to largest. Click on the species name for more info about that species, or on the Sub-Family name for more info about the sub-family.

KEY: 1. Ants that are less than 5 mm long

4. All ants are classified into sub-families; within each of these the various genera and species share certain common characteristics. You can find out more about each sub-family by clicking on these links (these are not the only sub-families that exist, but are the only ones dealt with on this site so far):
Ponerinae (the ‘Primitive’ or Ringbum ants)
Pseudomyrmecinae (the Slender ants)
Dorylinae (the Army or Driver ants)
Aenictinae (the False Army ants)
Dolichoderinae (the Smelly ants)
Myrmicinae (the Double-waisted ants)
Formicinae (the Elegant, Acid-Squirting, Pugnacious or Sugar ants)

5. For a detailed Bayesian Key go here


Myrmecochory: ant distribution of seeds

Myrmecochory (literally ‘ant borne’) is a process whereby certain plants ensure the survival of their species after a fire. The seeds or fruits of literally hundreds of fynbos plants have a fleshy covering or attachment that acts as an ‘elaiosome’ or ant-attractant. When the seeds fall to the ground certain ant species, especially ants of the genus Anoplolepis (Pugnacious ants) rush to the seeds, attracted by pheromone-imitating scents. The ants sink their jaws into the soft, fleshy covering and tug the seeds into their nests, where the sweet-tasting reward is consumed. The hard, smooth nuts that are left cannot be gripped by the ants’ tiny mandibles, and so remain buried in their nests, safe from fire and animal predators. The seeds’ longevity is also enhanced by the anti-bacterial and fungicidal substances which the ants secrete to keep their crowded nests healthy. The fruits of Mimetes, Leucospermum, Paranomus and several other Proteaceous genera, all the Buchus or Rutaceae, many legumes and scores of other genera are involved in this important ecological process.
Serotinous fynbos plants have a creaky way of surviving fire, says William Bond. Serotiny means that they don’t release any seed until they’re burnt. When the plants die, the seed is released from cones or other structures. This works well unless the fire-interval is too short. If a second fire occurs before a population of serotinous proteas or other plants has matured, the species will become locally extinct. There is no safe seed store, and the species has to repopulate the area from outside—if it can. To achieve this it has to produce lots of seeds that can be carried for many miles on the wind, a pretty random process.
The re-emergence of Mimetes stokoei, thought to be extinct for nearly fifty years, dramatically demonstrates the success of myrmecochory, especially for large-seeded plants. Using ants to store seed safely in the ground is a process that is 180̊ different from serotiny. The seed does not have to travel at all, it drops straight to the ground and is buried by ants within a few minutes in the same optimal soil as its parent. These plants need to produce fewer seeds, because the chance of successful regeneration after fire is much less random. There is a safe underground seed store and not all the seeds germinate after each fire. A very short interval between two fires may have no effect. Finally, we now know that such seeds can remain viable for long periods—probably a century or longer. Tiny populations of rare species can survive in specialised habitats, apparently indefinitely—which is good news for the future of fynbos diversity. 

Mimetes stokoei

The unusual case of the extremely rare Mimetes stokoei shows how sustained interest by careful and dedicated observers over many decades can be very rewarding. If, for example, a common Leucospermum had disappeared from that habitat for over fifty years, it might not have been noticed —unlike in the case of the Mimetes. The longevity of its seeds would never have been recorded.

Invasive ants such as the Argentine ant – Linepithema humile – massively disrupt these processes. They not only completely eliminate most of the indigenous ants involved in myrmecochory, they don’t bury the affected seeds. They merely eat the elaiosomes off in situ, leaving the seed unburied and unprotected, at the mercy of rodents, birds, etc. Preliminary studies have shown that if the spread of these ants into wild habitats is not checked, the future of thousands of fynbos plant species will be at risk.



Fynbos plants that rely upon ants

A selection of the huge range of fynbos plant species that rely upon ants for effective distribution of their seeds ... we’ll add to these from time to time.

1. Plants that rely on ants to distribute their seeds:






































Ants vs Termites: the Soul of the White Ant

Ever since Eugene Marais wrote ‘The Soul of the White Ant’, public perception has lumped ants and termites together as sharing some kind of commonality. Overseas scientists did not help, either. Marais preferred to write in Afrikaans and his work was translated into various international languages either late in his life or after his death. His book “Soul of the White Ant” was plagiarised by Nobel laureate Maurice Maeterlinck, who published “The Life of the White Ant”
in 1926, falsely claiming many of Marais’ revolutionary ideas as his own. Maeterlinck was able to do this because he was Belgian and, though his mother tongue was French, he was fluent in Dutch, from which Afrikaans was derived. It was common at the time for worthy articles published in Afrikaans to be reproduced in Flemish and Dutch magazines and journals.
Marais contemplated legal action against Maeterlinck but gave up the idea in the face of the costs and logistics involved. He needn’t have bothered: Maeterlinck chose a bad title to plagiarise, leaving his reputation in tatters on two counts. 
Ants and termites are both insects, but that’s about where it ends. Lumping them together is a bit worse than putting golden moles and baboons into the same mammalian nest.
So what’s different? Termites, like ants, live in large colonies in the earth or in wood. The individual workers are generally sterile, and the queen lays all the eggs. Their winged reproductives go on nuptial flights, when they're often known as ‘flying ants’. Termites sometimes have major and minor workers, too, but that's where it ends. Whereas ants [and bees] are related to wasps, termites have their roots in cockroaches, and almost everything else about them differs from ants – see chart below.


Bibliography and links: read more about ants

mostly under construction: much more to come here

Websites recommended:
http://antsofafrica.org/
http://www.antweb.org/
http://www.antwiki.org/

Papers and further reading:

Review of Anoplolepis ... and notes on Acropyga – A J Prins, Annals of the SA Museum, June 1982
http://antsofafrica.org/ant_species_2012/anoplolepis/anoplolepis/prins-1982-hns4205.pdf

Characterization of an Acropyga arnoldi mating swarm and early stage colony founding behaviour
LaPolla and Spearman, 2007
http://www.jstor.org/discover/10.2307/25078979?uid=2&uid=4&sid=21105325749723

Taxonomy of Ponerine genus Hypoponera
Bolton & Fisher, 2011
http://www.mapress.com/zootaxa/2011/f/zt02843p118.pdf

The Genus Hagensia Forel
George Arnold, May 1951
http://antsofafrica.org/ant_species_2012/pachycondyla/pachycondyla_saldanhae/arnold1951_hagensia.pdf

Ant imitators: don’t be fooled!


Ants are amongst the most successful of all insects, so it’s hardly surprising that a large number of other goggas imitate them. There are a large number of beetles, spiders and even wingless wasps that pretend to be ants. Some do it to avoid being eaten themselves [ants don’t generally taste very nice, unless you’re an ant-eater]. Others do it because they prey on ants, and by pretending to be one amongst friends they can avoid being torn apart themselves. Arthropods that mimic ants are technically known as myrmecomorphs.


Wasp myrmecomorphs


Wasps and ants belong to the same order of insects [Hymenoptera] with essentially similar body-shapes, so it may not be all that surprising that there are many convincing ant-mimics amongst the wasps. However, some wingless wasps are just that, and despite the lack of wings they don’t resemble any specific ants at all. Our first subject here, however, was collected on the wall of the Oudebosch office, where we initially mistook it for an Argentine ant, so convincing was the mimicry. That said, it seems odd that a local wasp could mimic an invasive ant, so it is possibly imitating a Pheidole instead.
You can always tell wasps from ants by the antennae, if nothing else. Wasps’ antennae form a smooth curve of more-or-less equal segments on a short scape, whereas ants all have a long first segment [scape], followed by a series of shorter ones. Ant antennae thus form an L shape rather than a smooth curve.
Simon Van Noort of Iziko Museums identified this little wasp as Conophorisca littoriticus. He writes in iSpot:
This is a species of a bizarre subfamily of Pteromalidae (Chalcidoidea) that are parasitoids of a range of insects (see WaspWeb link below under comments). Biology of most genera is unknown, but curculionid beetles (Coleoptera); mantid egg cases (Mantodea); or Glossina tsetse fly puparia (Glossinidae, Diptera) are known hosts. There is a rich African fauna, with most species still undescribed.






Spider myrmecomorphs

A fascinating aspect of these imitators are those impersonators – particularly spiders – that imitate ants that have long disappeared from suburban gardens in the face of invasion by Argentine ants and other aliens. The faux-ants provide us with a record of the historical ant-fauna of these areas. As time goes on we hope to build up an illustrated record of these frauds on this site; any photographs that we can draw from would be most welcome.
The ant-mimic above is Mexcala rufa [cf Tony Rebelo, iSpot]. It is a jumping spider from the sub-family Salticidae, and very accurately mimics the common Balbyter sugar ant, Camponotus fulvopilosus

But here’s the rub: the spider hails from Muizenberg, quite the wrong habitat for this dry, hot country ant. Could the spider be a relic from a hotter, drier time in the Cape?



Beetle myrmecomorphs


Although this gogga looks very ant-like, when you look at its underside you can see very clearly that its legs are attached utterly differently from those of an ant. Remember, too, that the first segment of the antennae of all ants is an elongated ‘stalk’.
Another ant-imitator [awaiting ID] – its movements are uncannily ant-like

Very similar to the above and probably the opposite sex of the same gogga, they’ve been identified as belonging to an undescribed genus of the sub-family Korynetinae
Here’s a wonderful conundrum; this undescribed, unnamed myrmecomorph is probably from the sub-family Anthicidae [Riaan Stals, pers comm] and it not only imitates ants, it seems to imitate the Korynetinae beetles above. Or is the Korynetine imitating the Anthicide? Anthicide or Korynetine? Who knows?


This tiny fellow seems to imitate Lepisiota ants; we are waiting for a name for him. (or her?)


Under construction