Camponotus: A Sugary High

I think I may have said before that Australia is the land of ants. When travelling in Australia's arid regions (i.e. most of the continent), ants are often the most visible animals about. Perhaps the most visible of all Australia's ants are the meat ants (Iridomyrmex), but not too far behind them are the sugar ants of the genus Camponotus.

Workers and emerging queens of banded sugar ants Camponotus consobrinus around the nest opening, copyright Steve Shattuck.

Camponotus is a genus of the ant subfamily Formicinae found pretty much everywhere around the world that ants are to be found. It is massively diverse: well over 1000 species have been assigned to this genus over the years, with probably more to be described. They are correspondingly diverse in habits and appearance. Some are among the giants of the ant world, others are much smaller. Some form massive colonies that are difficult to miss and forage during the day, others are more retiring and emerge only at night. Some construct their nests in holes under the grounds, others hollow out wood or use the holes left by other wood-boring insects. Most (but not all) Camponotus species exhibit some form of worker polymorphism: rather than having just a single worker caste, a colony will often include large major workers and much smaller minor workers, with the two forms superficially looking quite different. Sometimes the distinction between majors and minors will be quite clear, other times there will also be workers of intermediate sizes. In some Australian species, known as honeypot ants, there are specialised workers called 'repletes' who spend their lives hanging in one spot inside the nest, being fed by the other active workers until their gasters swell into engorged round balls. These repletes serve the colony as a living larder, able to regurgitate their stored excess of food when needed by their nestmates. Despite all this diversity, most Camponotus species are readily recognisable as Camponotus: they usually lack spines on the mesosoma (the 'thorax'), the back end of which is narrow and often arched. This smoothness and slimness gives Camponotus a distinctive look that kind of puts me in mind of the ant version of a greyhound. The majority of Camponotus species also differ from other ants in lacking the metapleural gland, a gland producing an antibiotic chemical whose opening is usually visible near the rear of the mesosoma.

Camponotus aurocinctus, copyright Steve Shattuck.

Camponotus species have been referred to in Australia as 'sugar ants' in reference to their diet, which is commonly dominated by the sugary excretions of plant-sucking bugs that they attend. In other parts of the world, they have sometimes been referred to as 'carpenter ants' in reference to the wood-tunneling habits of their most notorious representatives. Bug-derived honeydew is high in sugar but low in other essential nutrients, so the ants also feed on things such as the scavenged bodies of the bugs themselves after death. They are also probably assisted in meeting their nutritive needs by Blochmannia, an endosymbiotic bacterium that infests specialised cells in the gut of Camponotus and closely related genera (Wernegreen et al. 2009). Genetic data from the endosymbiont indicates that it probably synthesises nutrients the ant does not otherwise ingest. It may also play some role in compensating for an absence of metapleural gland secretions. As well as the gut, Blochmannia infest the ovaries of reproductive females and are passed to the next generation via the developing oocytes. Phylogenetic analysis of Blochmannia indicates that it is closely related to other endosymbiotic bacteria found in mealybugs, and it is possible that the ancestors of Camponotus picked it up in the course of feeding on honeydew.

Honeypot ant Camponotus inflatus repletes hanging in the nest, copyright Mike Gillam.

The sheer size of Camponotus as a genus has been a challenge to understanding relationships within the genus. Over thirty subgenera have been proposed at one time or another, but many of these are poorly defined and many authors eschew using them in favour of informal species groups. It does not help matters that, since the early 20^th century, most reviews of Camponotus have been conducted at a local rather than a global level. Those studies that have touched on Camponotus phylogeny in recent years suggest the need for a large-scale revision, with few of the subgenera supported as monophyletic.

REFERENCES

Wernegreen, J. J., S. N. Kauppinen, S. G. Brady & P. S. Ward. 2009. One nutritional symbiosis begat another: phylogenetic evidence that the ant tribe Camponotini acquired Blochmannia by tending sap-feeding insects. BMC Evolutionary Biology 9: 292. doi:10.1186/1471-2148-9-292.

Trap-jaw Ants of Australia (and a couple from Africa)

Foraging worker of Epopostruma frosti, copyright Alex Wild.

Anyone who finds themselves travelling through regional Australia will soon find themselves convinced that this is a continent ruled by ants. During the course of the day, while the hot Australian sun drives other animals to seek shelter and seclusion, ants are often the only living things (other than plants) to be seen. To match this abundance, Australia's ants also come in a variety of distinctive forms, many of them unique to this country.

One distinctively Australian group of ants are the 'epopostrumiforms'. This is a small group of genera belonging to the tribe Dacetonini of the subfamily Myrmicinae (in the past the epopostrumiforms have been formally recognised as the subtribe Epopostrumiti, though Bolton eschewed the use of formal subtribes in his 1999 review of the Dacetonini). The Dacetonini are all predatory ants, with a distinctive large process inside the base of the mandibles that helps to lock them closed when holding struggling prey. The mandibles may be particularly long and slender, sometimes with only a few teeth present at the end. Where their habits are known, epopostrumiforms are predators of springtails; these are the most typical prey for the Dacetonini as a whole though some species of the tribe are more catholic in their tastes. Dacetonins live in small colonies, commonly in secluded habitats such as leaf litter; the epopostrumiforms include species that nest and forage either above or below ground (Brown & Wilson 1959). Dacetonins hunt their prey by stealthily sneaking up to it with the mandibles held open, followed by a quick lunge combined with snapping the mandibles shut. Once the prey has been successfully grabbed, those dacetonins with shorter mandibles rapidly bring the sting forward to quell it. Even after using the sting, however, hunters of springtails may find themselves flung into the air by flicks of the springtail's furca a couple of times before the venom takes full effect (hence the need for a firm mandibular lock). Dacetonins with longer mandibles may also deploy their sting or they may simply lift the prey above their heads until it gives up the ghost.

The African Microdaceton tanyspinosum, copyright April Nobile.

As already indicated, the majority of epopostrumiforms are endemic to Australia (one genus, Colobostruma, includes a few species found in New Guinea and the Solomon Islands). The only non-Australasian taxon to be assigned to the Epopostrumiti is an African long-mandibulate genus Microdaceton. Features uniting Microdaceton with the Australasian epopostrumiforms include the presence of lateral outgrowths on the petiole and postpetiole (the first two nodular segments of the metasoma) and the position of the petiolar spiracle (Bolton 1999) but some authors have suggested a closer relationship of Microdaceton to other dacetonin genera. Even if correctly positioned, Microdaceton is at most the sister taxon to the Australasian clade, members of which are united by features such as reduced antennae and an enlarged labrum.

Face of Colobostruma alinodis, copyright Estella Ortega.

Bolton (1999) divided the Australasian epopostrumiforms between three genera: Colobostruma, Mesostruma and Epopostruma. Less than fifty species of this clade have been described to date though others probably remain to be named. Even among the known species, many are rare and/or cryptic and some are known from only a very few specimens. Epopostruma resembles Microdaceton in having elongate mandibles with only a small number of interlocking teeth at the end (two in Epopostruma, three in Microdaceton). When hunting, Epopostruma may open their mandibles to a full 170°. Colobostruma has much shorter, triangular mandibles with numerous teeth; Mesostruma has triangular mandibles somewhat intermediate between the other two genera. The mandibles of both Colobostruma and Mesostruma cannot be opened to the same degree as those of Epopostruma; rather, species of these two genera will open their mandibles to a maximum angle of 90° when hunting. Whether the Dacetonini involved long mandibles on a single occasion, with a number of sub-lineages reverting to shorter mandibles afterwards, or whether the short-mandibled Dacetonini retain the ancestral morphology and long mandibles evolved on multiple occasions within the tribe, remains a question occasioning some debate.

REFERENCES

Bolton, B. 1999. Ant genera of the tribe Dacetonini (Hymenoptera: Formicidae). Journal of Natural History 33: 1639–1689.

Brown, W. L., Jr & E. O. Wilson. 1959. The evolution of the dacetine ants. Quarterly Review of Biology 34 (4): 278–294.