Tetraponera aethiops worker showing the location of the clypeus in green (Scanning Electron Micrograph, Roberto Keller/AMNH)

When looking at an arthropod from our vertebrate perspective it is easy to forget that we are looking right at the animal’s skeleton. While our own vertebrate skeleton consists of a series of internal compact pieces with sponge-like cores that support an external layer of muscles and entrails (all nicely wrapped in skin), the reverse is true for arthropods. The arthropod skeleton consists of a series of external plates and hollow tubes that form enclosed spaces within which the internal musculature system attaches¹. One consequence of this peculiar body architecture is that most of what we see on the outer surface of this exoskeleton is but a reflection of what is going on on the inside– minute external pits correspond to places where the cuticle folds in to form internal pillars, and innocent looking shallow furrows on the surface are large internal walls where powerful muscles originate. A simple examination of the exoskeleton, therefore, can tell us a lot about particular functions and consequently about an insect’s behavior.

The clypeus (in green) on the turtle ant Procryptocerus, with a characteristic brush on its anterior border (Scanning Electron Micrograph, Roberto Keller/AMNH).

A good example of this is provided by the clypeus in ants and its wide diversity of forms across the different species in the family. The clypeus corresponds to an unpaired skeletal plate lying right at the center of an insect face. It is normally located lower in the head just in front to where the antennae are inserted, but in many ant groups it quite commonly extends in between the antennal sockets. The anterior border of the clypeus is involved in two important articulations relating to the movement of the mouthparts. The central part forms a wide hinge with the mouth’s “lid” or labrum, allowing the latter to move forward and backwards to open and close the preoral cavity where the intricate ant tongue is stored when retracted (the actual opening of the mouth lies internally at the back-end of this preoral cavity). The sides of the clypeal border, on the other hand, form deep cavities where the anterior condyle of each mandible articulate.

Those articulations occur externally. But what is going on the inside of the clypeus? The inner surface on the clypeus provides attachment to a set of muscles that originate right at the anterodorsal section of a special elastic chamber located just before the mouth known as cibarium. When these muscles contract the cibarium expands producing a suction action. It is basically the sucking pump of the insect, and the bigger the clypeus the bigger the pump muscles and the larger the sucking force. You may have probably noticed the big goofy snout in cicadas; well it is nothing but the hypertrophied clypeus attesting to the large sucking pump of these dedicated suckers. Ants never reach such extremes, but the clypeus can be quite large in some groups.

Clypeus (in green) on a Onychomyrmex doddi worker. Species in this genus display a convergent army ant like behavior (Scanning Electron Micrograph, Roberto Keller/AMNH)

In clades of chiefly predatory ants, like amblyoponines, the clypeus is never large and has become rather reduced in the more specialized genera like Apomyrma and Onychomyrmex. The same pattern occurs more or less among ponerines.

The large clypeus (in green) on a Formica fusca worker (left antenna removed. Scanning Electron Micrograph, Roberto Keller/AMNH).

Oh, but formicines and dolichoderines, those ants are such big suckers. Those are the ants you will most commonly see wandering between flowers looking for nectar and tending aphids for honeydew (that is, they suck ass big time²). Myrmecines ants have large clypeus in general, and it is not surprising to see a correlation between tending other insects and having a well developed clypeus in genera like Crematogaster.

The clypeus (in green, maybe) in a Neotropical army ant Labidus coecus worker (the clypeus is there, I swear. Left antenna removed. Scanning Electron Micrograph, Roberto Keller/AMNH)

Now army ants, the ultimate specialized predators of the insect world, they are not suckers at all. All clades can be easily characterized by having almost no clypeus, so that the antennal sockets seem to fall off their heads forward.

The swollen clypeus (in green) on an Acanthoponera minor worker (left antenna removed. Scanning Electron Micrograph, Roberto Keller/AMNH)

One genus that intriges me is Acanthoponera. Species in this genus have a very large and swollen clypeus for what you will expect given the group’s phylogenetic position in between other major clades of ants. I don’t think much is known about the biology of this genus other than it is a nocturnal ant. But I bet you this ant is sucking around something.

Notes

Red Hot Chilli Peppers? No, dentiform setae in the labrum of an Onychomyrmex doddi worker (Scanning Electron Micrograph, Roberto Keller/AMNH)

Just as the anterior margin of an ant’s cranium can sometimes be armed with rows of dentiform clypeal setae (that is, especially modified hairs), the lid that closes the insect’s mouth called labrum can bear identical structures. The image above shows two of these specialized teeth-like pieces (in red) flanking an empty broad socket where a third piece used to be inserted.

The labrum and part of the clypeus of an Onychomyrmex doddi worker. A row of dentiform setae adorn the labrum (in red) and the clypeus (in yellow; Scanning Electron Micrograph, Roberto Keller/AMNH)

Although these dentiform setae vary in size and shape quite considerably from species to species, when they are present in different parts of the body within an individual they show the same morphology, suggesting that they are the result of a similar developmental program that switches on at the different positions.

Head of the African subterranean ant Apomyrma stygia showing the hypertrophied dentiform setae in the labrum (in red; Scanning Electron Micrograph, Roberto Keller/AMNH)

Moreover, there is a interesting similarity between dentiform setae that have developed in similar but apparently independent (non-homologous) conditions. The hugely grown dentiform setae restricted to the labrum in Apomyrma stygia are identical to the similarly hypertrophied ones found in Amblyopone pluto but that occur exclusively in the clypeus (see last image on this post). Though not sisters, these two taxa belong to the same Amblyoponinae clade.

Lastly, a couple of comments regarding a recent paper describing the ant fossil Gerontoformica, which has similar detiform setae in both the clypeus and labrum. Nel and coworkers¹ mention that dentiform setae in extant ants are found either in the clypeus or in the labrum but never in combination. This is not the case as can be seen in the example of Onychomyrmex pictured above. There is also the suggestion that Probolomyrmex, a non-amblyoponine genus, has dentiform setae on the labrum. However, close inspection revels that this is also not the case. Rather, most setae covering the body in this genus, including the few stout ones on the labrum surface, are short and scale-like and can easily be confused with pegs at low magnification.

So far, in extant taxa these peculiar dentiform setae arming the clypeus and/or labrum are only known to occur within the subfamily Amblyoponinae.

References

Frontal part of the head in an Anochetus emarginatus worker, profile view (Scanning Electron Micrograph, Roberto Keller/AMNH)

This image shows the mouthparts of a trap-jaw ant in resting position. The only structures really visible are the prominent elongated mandibles (in yellow) that project forward. The rest of the pieces, laying immediately below, are retracted inside the preoral cavity.

Fully extended mouthparts in an Anochetus emarginatus worker, profile view (Scanning Electron Micrograph, Roberto Keller/AMNH)

And this is how the mouthparts look fully extended, when the ant is sticking its tongue out. There are four different sets of structures here: the labrum (in green); the mandibles (in yellow); the maxillae (in orange); and the labium (in red). Each set corresponds originally, both phylogenetically and ontogenetically, to a pair of structures, although only the last three are modified limbs properly (each correspond to a pair of head appendages).

These are very complex structures, each part deserving its own separate discussion. But I wanted to thrown these images here now to serve as reference for future posts, and mention a few important generalities.

Ants display the unmodified general architecture of a biting insect. The mouthparts of adult ants are typical for what is found when comparing different insect groups, and one can readily homologize each part with a corresponding structure in a grasshopper or a beetle for example. Even the most derived mouthpart morphologies found within ants, like that of the trap-jaw ant pictured here, preserve this general pattern.

However, ants do have some uniquely derived features. They are truly prognathous insects, something uncommon within Hymenoptera (but not exclusive). While in bees and in most parasitic and stinging wasps the mouthparts hang down below the head pointing to the ground, in ants they are directed forward, always pointing to the front.

Ant prognathy, however, is not only a function of the fact that the whole head is tilted forward. Examine the image above and you will notice that the labrum, maxillae and labium are fully extended while the mandibles remain closed. That is, unlike other Hymenoptera, prognathous or not, in ants the mandibles do not fold right on top of the rest of the mouthparts at rest. Instead the main body of the mandible “steps-up” immediately after the mandible’s articulation (the rounded yellow piece at the far right), thus laying out of the way from the remaining structures.

The much derived Anochetus pictured here provides an extreme example illustrating this, but the exceptional modification is universally shared within the family. It is another unique ant synapomorphy. Obvious as it may seem once explained, I have to confess it took me a while (a few years actually) to figure out what was happening structurally in ants that was different from the non-formicid outgroups. But since then, after the explanation clicked, I cannot look at an ant without seeing it.