Dealing with a Clingy Male

Diving beetles of the family Dytiscidae are a distinctive component of the freshwater environment in most regions of the world. They have an oval, streamlined body form and powerful hind legs, usually with fringes of stiff setae, that are ill-suited for movement on land but make them adept swimmers. They are also almost always capable fliers, allowing them to find their way to water bodies of any size from large lakes to small, temporary pools. Both adults and larvae are active hunters, preying on other aquatic arthropods or even small vertebrates. Most diving beetles are fairly dull in coloration but exceptions are found among members of the tribe Aciliini.

Sunburst diving beetle Thermonectus marmoratus, from Insectarium de Montréal, René Limoges.

Members of the Aciliini are moderately sized diving beetles, generally between one or two centimetres in length. Dorsally they have a yellow to red base coloration with contrasting dark markings. The hind legs are robust with the hind tibia short and broad. Males have the base of the tarsus of the front legs broadened into a round palette with setae on the underside modified into sucking discs, used to hang onto the females when mating; this discs may be present on the tarsus of the mid pair of legs as well. They are strong swimmers, often venture=ing into the open waters of lakes and pools, and contrast with other diving beetles in that they may be found in pools lacking submerged vegetation (Roughley & Larson 2001; Bergsten & Miller 2006). Larvae have a distinctive arched body shape with a small head (Bukontaite et al. 2014), kind of shrimp-like, and also tend to be more pelagic than the larvae of other diving beetles. Females have gonocoxae (the appendages at the end of the abdomen that function as the ovipositor) that are relatively long with a broadened, spoon-like ending (Miller 2001); these are used to insert eggs into damp moss or under loose bark of vegetation lying just above the waterline. There is usually just one generation per year and adults in cold regions overwinter in larger water bodies that remain unfrozen.

Alternate morphs of female Graphoderus zonatus with granular (left) and smooth elytra, from Holmgren et al. (2016).

Perhaps the most intriguing aspect of aciliin diving beetles regards their sexual dimorphism. As noted above, males have a set of suckers on the fore legs for hanging onto females when mating. However, females of some species have sculpted elytra rather than the smooth elytra of males, such as a granular surface in Graphoderus species or long, setose sulci in female Acilius. The uneven surface produced by these features presumably functions to reduce the efficacy of the males' suckers, allowing the females more control when selecting a mate. That such a conflict exists is supported by the observation that the more developed the males' sucker arrays in a population, the more likely the females are to have repellent sculpturing. Males of some diving beetle species have been observed grabbing at any female they encounter, followed by the female swimming rapidly and erratically in an attempt to shake the male off or knock him off against the substrate or objects in the water (Miller 2003). Where this becomes really interesting is that some species have dimorphic females with some females in the population having sculpted elytra whereas others are smooth. What could be the reason for such variation? The presence of both forms in the population suggests that neither has a complete advantage over the other. It may be that smooth-backed females trade reduced defenses for improved swimming ability. Alternatively, a defensive female may be able to ensure that only the strongest and most resilient males can mate with her, but runs the risk of not mating at all if she never encounters a male who can overcome her defenses. A less defensive female may be more vulnerable to any male she encounters but at least she's bound to be fertilised at some point.

REFERENCES

Bergsten, J., & K. B. Miller. 2006. Taxonomic revision of the Holarctic diving beetle genus Acilius Leach (Coleoptera: Dytiscidae). Systematic Entomology 31: 145–197.

Bukontaite, R., K. B. Miller & J. Bergsten. 2014. The utility of CAD in recovering Gondwanan vicariance events and the evolutionary history of Aciliini (Coleoptera: Dytiscidae). BMC Evolutionary Biology 14: 5.

Holmgren, S., R. Angus, F. Jia, Z. Chen & J. Bergsten. 2016. Resolving the taxonomic conundrum in Graphoderus of the east Palearctic with a key to all species (Coleoptera, Dytiscidae). ZooKeys 574: 113–142.

Miller, K. B. 2003. The phylogeny of diving beetles (Coleoptera: Dytiscidae) and the evolution of sexual conflict. Biological Journal of the Linnean Society 79: 359–388.

Roughley, R. E., & D. J. Larson. 2001. Dytiscidae Leach, 1815. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 156–186. CRC Press: Boca Raton.

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