Tierra de Oro Laboratory

Physiology and Ecology in the Pinyon-Juniper Savanna

Category: mimicry

  • Spiders, Bats, and Mystery Poop

    Who has been eating our beetles?

    Mimicry is in the eyes of the predator

    Some beetles that do not produce defensive stink are thought to be mimics of those that do. For example, Eleodes species can shoot streams of noxious chemicals when threatened, whereas Philolithus and Stenomorpha, which have very similar size and appearance, do not. In this way, the non-stinky beetles may be protected from predators who have learned their lessons during previous meetings with Eleodes. This form of mimicry was first proposed about 150 years ago by Henry Bates, and is referred to as Batesian mimicry.

    Because we humans perceive the world largely through our eyes, we tend to think of mimicry in terms of visual appearance. The examples shown in textbooks (see below) tend to be butterflies or snakes with matching colorful patterns. These are expected to be especially effective against birds or monkeys that have excellent color vision.

    Harmless scarlet kingsnake, a Batesian mimic of the coral snake. Credit: Glenn Bartolotti , CC BY-SA 3.0,

    A close look at the snakes above will show that they are relatively easy to distinguish in photographs. Mimicry does not need to be perfect, because a mistake in the field could cost an unwary predator or hiker their life. As a human, you could memorize a rhyme that allows you to distinguish them, or you could simply avoid bright red, black, and yellow snakes.

    What about the beetle mimics? Should we think about vision, or other senses? If a predator has poor vision or hunts in the dark, then what is the purpose of visual similarity?

    Who Eats Local Beetles?

    One of my goals for the 2026 field season was to find out who actually eats the beetles. Each of the beetles shown at the top of the page would yield 1/2 to 1 gram of water, protein, and fat, so it is worth some effort to subdue the defended ones, figure out who is undefended, or both.

    Previous work by a number of authors has shown that large darkling beetles are consumed by a wide variety of predators. Depending on the location and beetle species, predators include mice, skunks, coyotes, bats, ringtails, jays, scorpions, and there are likely to be more.

    Below is a summary of what I have figured out so far. As always, there is more to come, but the evidence for the ones shown below is quite solid.

    Waiting in Ambush: Black Widow

    When I came across some beetles wrapped up in sloppy webbing, I was not sure who had killed them. After collecting the beetles and moving the prickly pear pad below them, I saw a large spider with a red spot on its abdomen in a hole below. Then it all made sense. Black widows spin notoriously messy webs, and are perfect predators of any large beetle that comes by. When caught in the web, Eleodes‘ defensive chemicals will be sprayed ineffectually until they are used up, and the spider can then immobilize and consume the beetle at her leisure.

    This spider also trapped an Embaphion contusum (also chemically defended) and a large scarab beetle. One would expect that undefended Tenebrionid beetles, such as Philolithus and Stenomorpha, would easily fall prey to such a spider. Her web is not selective, and she is able to subdue E. obscura, one of the largest and best-defended beetles.

    Chomped by Bats

    Pallid bats are known to eat Stenomorpha, and the resident group has dropped bits and pieces of large insects such as Jerusalem crickets and beetles. For more about the bats themselves, have a look at this post.

    In August, one of the bats was particularly messy, leaving most of the front half of a small Stenomorpha, possibly S. obovata.

    Now that fall is upon us, Stenomorpha rimata has become the most common Tenebrionid in the neighborhood. I was lucky enough to find some half-eaten, but readily identifiable bits of S. rimata under the roost last week.

    At this point, we don’t know if the bats are focusing on undefended beetles, or whether they would eat Eleodes if given the chance. It is clear, however, that pallid bats are willing and able to eat large Tenebrionid beetles such as Stenomorpha.

    Work in Progress: Beetles in Scat

    During the summer and fall, one can readily find scat containing beetle remains. Predators often swallow them whole, making identification much easier.

    Analysis is in the early stages, but there is lots to find. The photo above shows some of the head capsules that were in one scat. There were many Carabid beetles of a range of sizes, plenty of big and small ants, and a few good-sized Tenebrionids. Based on an abdomen that was also in the collection, at least one of the Tenebrionids was Eleodes obscura.

    There is much more to find, but we can say for sure that somebody, probably a mammal, is eating a lot of beetles. Importantly, they are including Eleodes in their diets. With persistence and luck, we should be able to figure out who it is as we dig deeper into the samples.

    The predators we have identified so far, spiders and bats, are not visual predators, so the outward resemblance of the mimics to Eleodes is probably not that useful. Moreover, it looks like at least some predators are happy to eat Eleodes, further reducing the possible benefit of mimicry. Nonetheless, we may find that birds such as jays and ravens are hunting the beetles, and we may yet identify some selective advantage to the visual resemblance shown by the mimics.

  • One Project Becomes Two

    After working very hard this winter to submit a manuscript describing the most recent results of the project, I realized that it has really become two non-overlapping lines of inquiry.

    A few weeks ago, I submitted a manuscript titled “Batesian Mimicry and Thermal Resilience Among Tenebrionid Beetles from a New Mexican Piñon-Juniper Savanna” to the Journal of Experimental Biology. The editor said some kind words about it, but ultimately rejected it because it did not fit comfortably into the aims of the journal. The main complaint was that a significant portion of the paper focused on the ecology of the beetles, rather than their physiology.

    It took a few days for me to realize that the paper had evolved into something different. Looking over the title, two things may stand out.

    Batesian Mimicry

    Thermal Resilience

    Those are really two separate concepts and analyzing either one can be performed relatively independently of the other. In other words, this was two incomplete papers rather than one solid story.

    Now begins the process of completing both of those stories.

    Mimicry

    Several lines of evidence address whether the Asidini Philolithus elatus and Stenomorpha rimata are Batesian mimics of the Amphidorine Eleodes obscura. For an organism to be a bona fide Batesian mimic it must

    • Resemble the model in the eyes (or ears or nose) of potential predators.
    • Be palatable
    • Be undefended (or at least to a potential predator)
    • Overlap the model in space and/or time so that predators can learn the association
    • Be relatively uncommon to prevent predators from discovering their mimicry

    The Asidini fulfill these criteria to varying degrees

    • The major argument for mimicry is that the Asidini look a lot like E. obscura, and are about the same size and mass.
    • A close relative of S. rimata, S. marginata, has been shown to be palatable to mice and skunks. Palatability of P. elatus is unknown.

    On the minus side:

    • P. elatus will regurgitate when handled, suggesting at least mild defense
    • Headstanding behavior, which contributes to the mimicry, is weak (P. elatus) or non-existent (S. rimata)
    • Both Asidini are at least as common as E. obscura during peak season, giving predators plenty of opportunity to learn that they are quite edible.
    • Stenomorpha rimata is present in late September and October, when Eleodes have disappeared from the surface, giving predators plenty of time to learn that they are tasty and undefended.

    I really need to know who the local predators are, so that I can make sense of all this. I hope to use the upcoming field season to find out who is eating beetles around here. Based on scats and scattered body parts, someone is eating them, but who?

    Thermal Resilience

    Maintenance of scent glands is costly, so E. obscura should either use more metabolic energy than the undefended P. elatus and S. rimata, or devote less energy to other aspects of fitness. Based on respirometry across a range of temperatures it would appear that all three species use the same amount of energy per unit time, but the Asidini have much higher survival at elevated temperatures.

    This is a nice partial story, but I need to know more about possible mechanisms. Two sets of experiments would be very helpful in completing this part of the story.

    • Testing survival across a full range of temperatures. I have only tested up to 40C, which is semi-lethal to Eleodes but does not strongly affect P. elatus or S. rimata. Although I am no fond of the idea, I need to test up to a temperature that is 100% lethal in order to have a complete picture of thermal resilience.
    • Because temperature induced lethality is associated with disrupted ionic regulation, it will be important to examine concentrations of ions in the hemolymph (blood) of E. obscura, P. elatus and S. rimata at high temperatures.

    If all goes well, I should have two papers ready by the end of the season. Wish me luck.

  • Mimicking Stinky Beetles: How good is good enough?

    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).

    Eleodes obscura traveling across an open field. 7/9/25.

    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.

    large black beetle standing on its head as a warning display.
    Eleodes obscura reacting to an approaching threat by standing on its head. 8/17/24

    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.

    Two species of black beetles side by side, labeled at Eleodes obscura and Philolithus elatus.
    Eleodes obscura dispersa and Philolithus elatus infernus at the same scale. The elytra show similar patterns of ridges and punctures. Paint spots mark them for physiological studies.

    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.

    Stenomorpha marginata. 9/3/25.

    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.

    Philolithus elatus giving its best effort at a headstand.
    Stenomorpha marginata bracing for whatever comes next. 10/1/25.

    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.

    Eleodes longicollis raising its abdomen in response to an approaching experimenter.
    Moneilema appressum performing a convincing headstand. 8/20/25.

    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.

    Philolithus elatus regurgitating in response to handling. 8/11/25.

    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.