Many of the darkling beetles here are members of model/mimic complexes. Specifically, some species of Eleodes, which can defend themselves chemically by using scent glands in their abdomens, serve as models for members of other genera, such as Philolithus and Stenomorpha, which do not have glands (Brown, 1971; Smith et al., 2015). In theory, the mimics are protected from predators by resembling the defended beetles, but save energy by not having to grow, maintain, and fill the glands. When a mimic resembles a defended species but does not have its own defense, it is referred to as Batesian mimicry.

As we look over the data from the past few seasons, it is worth considering some of the assumptions regarding models and mimics.

Advertising Bad Taste

Let’s start with the models, the chemically-defended Eleodes. If a species is noxious in some way, advertising this fact benefits both the prey and predator. If the predator knows that the prey is inedible, then the prey will avoid being eaten, and the predator avoids a mouthful of something nasty. Many noxious prey have bright, contrasting colors to make it easier to recognize them.

There are about six species of Eleodes commonly found here, and all have similar appearance and behavior. They are black, generally shiny, about 15 to 30 mm long (depending on the species), with variations in their body shapes and the patterning of the thorax and elytra (for an overview, see the local beetles page).

One could argue that uniformly black beetles are not very striking, and may therefore be doing a poor job of advertising their distastefulness. However, when walking about, they put little effort into concealing themselves, and a shiny black beetle is pretty easy to see on a background composed of shades of brown and drab green. Further, when approached the beetles stand on their heads to aim (and presumably show off) their scent glands.

These guys may not be as flamboyant as a coral snake or a lionfish, but they get the point across. After a few tries, a predator will probably get the idea that attacking one of these beetles will result in a face full of noxious chemicals.

The Mimics

From April to late July, the area is inhabited mostly by “honest” beetles, whose appearance and behavior truly reflect their ability to defend themselves. Starting in late summer/early fall, a group of “cheaters,” or mimics, starts to emerge. These are beetles that are about the same size and appearance as Eleodes, and in some cases act like them, but do not have defensive glands.

One species, Philolithus elatus, is considered to be an accurate mimic of E. obscura. Both E. obscura and P. elatus have multiple geographic subspecies, and the patterning on the elytra of each P. elatus subspecies resembles that of the E. obscura subspecies that shares its range (Brown, 1971). In the example above, the elytra of P. elatus infernus found in Santa Fe have weakly defined ridges, similar to those of the local E. obscura dispersa.

Philolithus emerges in late July and is abundant through August and early September. This means that predators have had since the beginning of April to sample Eleodes and learn that shiny black beetles taste bad. Another supposed mimic of E. obscura, Stenomorpha marginata, emerges even later, in late August and is present in September and October.

At first glance, the mimics could easily be mistaken for E. obscura, and a predator might pass them by. At other loacations, it is reported that both species even mimic the headstanding behavior of Eleodes (Brown, 1971; Smith et al, 2015). Here in Santa Fe, it takes physical contact (rather than a close approach) to get P. elatus to respond, and even that is half-hearted.

Stenomorpha does not even bother with a lame attempt, and just extends its legs outward to brace itself when touched.

Another mimic produces more convincing headstands. Moneilema appressum, the cactus longhorn beetle, is a member of a completely separate family of beetles (Cerambycidae). Nonetheless, its smooth, shiny appearance and willingness to stand on its head when approached make it a convincing mimic of Eleodes longicollis.

All of this leaves a few open questions.

First, how good does mimicry really have to be? From April to August, just about every shiny black beetle that a predator encounters is chemically defended. Do Philolithus and Stenomorpha need to imitate the appearance and behavior of Eleodes all that precisely, or is it enough to be a shiny black beetle of about the right size and shape? Based on scat found around here, someone is definitely eating Eleodes, so it’s likely that some mimics will get eaten anyway.

The fact that Moneilema seems to do a better job of looking and acting like an Eleodes species is intriguing, although I am not sure what to conclude at the moment.

Second, are the mimics truly undefended? Philolithus will regurgitate when handled roughly, which, at least for grasshoppers, qualifies as chemical defense. Further, they feed, oviposit, and their larvae probably live among harvester ants (Pogonomyrmex sp.) during their development (McIntyre, 1999; Slobodkovich, 1979). It seems plausible that the beetles acquire some sort of chemical signature from the ants that deters attacks by the ants and other predators.

Stenomorpha, on the other hand, appears completely undefended. I have not observed them to regurgitate, they do not appear to associate with ants, and they do not have scent glands. They may be less vulnerable because they emerge so late in the season, but we will need to know more about their predators to have any insight into this.

References

Brown, K. W. (1971). A population approach to computer taxonomy with applications in the genus Gonasida.

McIntyre, N. E. (1999). Use of Pogonomyrmex nest-sites by Tenebrionid beetles (Coleoptera: Tenebrionidae) for oviposition and thermoregulation in a temperate grassland. The Southwestern Naturalist 44, 379–382.

Slobodchikoff, C. N. (1979). Utilization of Harvester Ant Debris by Tenebrionid Beetles. Environmental Entomology 8, 770–772.

Smith, A. D., Wilson, J. S. and Cognato, A. I. (2015). The evolution of Batesian mimicry within the North American Asidini (Coleoptera: Tenebrionidae). Cladistics 31, 441–454.