Cycloalexy in tortoise beetle larvae

One of the first insects I encountered during my visit this past November to  in Buenos Aires, Argentina were these tiny beetle larvae grouped together on a single leaf of an unidentified shrub.  The presence of fringed lateral appendages and exuvial-fecal debris masses held by caudal appendages immediately identifies them as larvae in the leaf beetle subfamily Cassidinae, known commonly in North America as “tortoise beetles” due to the appearance of the adults.  With nearly 3,000 species distributed throughout the world, tortoise beetles are easily recognizable as a group; however, species identifications can be much more difficult, especially in the Neotropics where the group reaches its greatest diversity (Borowiec and Świętojańska 2002–2011). Identification of larvae can be even more challenging, as the larvae of many species remain unknown, and I was unable to find adults in association with the larvae to aid my identification.

Anacassis sp. (poss. exarata) early-instar larvae on Baccharis salicifolia | Buenos Aires, Argentina

Nevertheless, host plant can be an important clue to leaf beetle identity, as most species in the family limit their feeding to a single plant genus or group of related plant genera. The shrub on which the beetles were feeding looked familiar to me, and while perusing a list of plants that have been recorded from the Reserve (Burgueño 2005) I had an “Aha!” moment when I spotted the asteraceous genus Baccharis. I decided the plant must represent Baccharis salicifolia because of its narrowly lanceolate, willow-like leaves with fine apical serrations (Cuatrecasas 1968) (see first photo). The only tortoise beetles known to feed on Baccharis are species in the genus Anacassis (McFadyen 1987), several species of which are known from Argentina, and one (Anacassis exarata) looking very much like the larvae in these photos.

Note the circular, heads-directed-inward orientation of all larvae around the periphery

The manner in which these early-instar (perhaps even newly hatched) larvae were feeding as a group while working their way down the length of the leaf towards its base is not something I had observed before. Larvae of most tortoise beetles are solitary feeders (Borowiec and Świętojańska 2002–2011), and I was further intrigued by the deliberate circular formation that the larvae had assumed.  The larvae around the periphery were all facing inward, tightly packed against each other and with their exuvial-fecal debris masses directed outward. Additional larvae were seen inside the circular formation. As I manipulated the leaf for photographs, the larvae would occasionally raise their debris masses up and outward, presumably a defensive reaction to disturbance and a perceived threat. It was clear to me that the larvae had deliberately “circled their wagons” for defensive purposes.

Close body contact allows exuvial-fecal debris masses to form a protective barrier against predators

In fact, this type of defensive strategy has been reported in a number of South American cassidines, as summarized by Jolivet et al. (1990), who coined the term “cycloalexy” (from the Greek κύκλος = circle, and αλεξω = defend) to describe such strategies. Cycloalexy can either be “heads in, tails out” or vice versa and is usually associated with other repellent activities such as coordinated threat movements, regurgitation, or biting. The strategy is intended to provide protection from predators such as ants and true bugs and parasitioid wasps, although some parasitoids seem to have thwarted the strategy by depositing their eggs where they can be ingested (thus avoiding direct confrontation with the prey). Cycloalexy has been described primarily among chrysomelid beetles and tenthredinoid hymenopterans (sawflies); however, examples from a few other insect orders (e.g., Diptera, Neuroptera, Lepidoptera) are known as well (Jolivet 2008).  All known cycloalexic insects are subsocial in the larval stage and often also exhibit maternal protection of eggs or newly hatched larvae.

This and several other older larvae had become solitary, presumably protected in part by greater size

In addition to this single group of early instar larvae, I noted also a few larger individuals—all of whom were feeding on the plant in a more solitary fashion. Presumably as the larvae grow larger they are more able to defend themselves, or perhaps larger larvae simply demand more “elbow room” because of the larger amounts of leaf tissue they require for feeding. If cycloalexy is beneficial for small cassidine larvae but less so for larger larvae, perhaps this behavior is actually more common than is currently realized.

REFERENCES:

Borowiec, L., and J. Świętojańska. 2002–2011. Cassidinae of the world – an interactive manual (Coleoptera: Chrysomelidae). http://www.biol.uni.wroc.pl/cassidae/katalog%20internetowy/index.htm [accessed 3 Dec 2011].

Burgueño, G. 2005. Manejo de la vetación en reservas naturales urbanas de la region metopolitana de Buenos Aires. Aves Argentinas, Asociación Ornitológica del Plata, Proyecto Reservas Naturales Urbanas, 16 pp.

Cuatrecasas, J. 1968. Notas adicionales, taxonómicas y corológicas, sobre Baccharis. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales 13(50):201–226.

Jolivet, P. 2008. Cycloalexy. In: J. L. Capinera [Ed.], Encyclopedia of Entomology, Springer Science+Business Media B.V.

Jolivet, P., Vasconcellos-Neto, J., and Weinstein, P. 1991. Cycloalexy: A new concept in the larval defense of insects. Insecta Mundi 4(1–4) (1990):133–141.

McFadyen, P. J. 1987. Host-specificity of five Anacassis species [Col.: Chrysomelidae] introduced into Australia for the biological control of Baccharis halimifolia [Compositae]. Entomophaga 32(4):377–379.

Copyright © Ted C. MacRae 2011