Biology and host range of <Emphasis Type="Italic">Aconophora compressa,</Emphasis> a candidate considered as a biocontrol agent of <Emphasis Type="Italic">Lantana camara</Emphasis> in Africa

The shrub, Lantana camara L. (Verbenaceae), of tropical American origin, is a serious weed in South Africa and is the target of a biological control programme. The stem-sucking membracid, Aconophora compressa Walker (Homoptera: Membracidae), from Mexico was reported to be very damaging and was therefore imported into South Africa for biology and host range studies. The female partially inserts the eggs into the woody portion of actively growing stems, and guards them against potential predators. Nymphs develop through five instars to the adult stage in about 45 days. The adults and nymphs feed on the sap of stems causing the leaves to wilt, the flowers to abort and the gradual dieback of stems. No-choice experiments showed that adult survival, egg production, and nymphal emergence was high on L. camara, and also on related ornamental and indigenous species. The adult reproductive performance was higher on some indigenous Lippia species than on L. camara. Furthermore, the nymphs developed faster on these indigenous species and high rates of reproductive performance on these non-target species were sustained over several generations. In multiple-choice trials, equal or larger numbers of egg batches were recorded from indigenous Lippia species and the ornamental plant, Aloysia citrodora Palau, than on L. camara. The treehopper, A. compressa, poses an unacceptable threat to indigenous Lippia species and has therefore been rejected as a biocontrol agent for L. camara in Africa.     

Host shifts and the beginning of signal divergence

Divergence between populations adapting to different environments may be facilitated when the populations differ in their sexual traits. We tested whether colonizing a novel environment may, through phenotypic plasticity, change sexual traits in a way that could alter the dynamics of sexual selection. This hypothesis has two components: changes in mean phenotypes across environments, and changes in the genetic background of the phenotypes that are produced -- or genotype x environment interaction (G x E). We simulated colonization of a novel environment and tested its effect on the mating signals of a member of the Enchenopa binotata species complex of treehoppers (Hemiptera: Membracidae), a clade that has diverged in a process involving host plant shifts and signal diversification. We found substantial genetic variation and G x E in most signal traits measured, with little or no change in mean signal phenotypes. We suggest that the expression of extant genetic variation across old and novel environments can initiate signal divergence.     

Resource exchange and partner recognition mediate mutualistic interactions between prey and their would-be predators

Animals may develop mutualistic associations with other species, whereby prey offer resources or services in exchange for protection from predators. Alternatively, prey may offer resources or services directly to their would-be predators in exchange for their lives. The latter may be the case of hemipterans that engage in mutualistic interactions with ants by offering a honeydew reward. We test the extent to which a honeydew offering versus partner recognition may play a role as proximate mechanisms deterring ants from predating upon their hemipteran partners. We showed that, when presented with a choice between a hemipteran partner and an alternative prey type, mutualist ants were less likely to attack and more likely to remain probing their hemipteran partners. This occurred even in the absence of an immediate sugary reward, suggesting either an evolved or learned partner recognition response. To a similar extent, however, ants were also less likely to attack the alternative prey type when laced with honey as a proxy for a honeydew reward. This was the case even after the honey had been depleted, suggesting an ability of ants to recognize new potential sources of sugars. Either possibility suggests a degree of innate or learned partner recognition.     

THE IMPORTANCE OF FEMALE CHOICE,MALE–MALE COMPETITION,AND SIGNAL TRANSMISSION AS CAUSES OF SELECTION ON MALE MATING SIGNALS

Selection on advertisement signals arises from interacting sources including female choice, male–male competition, and the communication channel (i.e., the signaling environment). To identify the contribution of individual sources of selection, we used previously quantified relationships between signal traits and each putative source to predict relationships between signal variation and fitness in Enchenopa binotata treehoppers (Hemiptera: Membracidae). We then measured phenotypic selection on signals and compared predicted and realized relationships between signal traits and mating success. We recorded male signals, then measured lifetime mating success at two population densities in a realistic environment in which sources of selection could interact. We identified which sources best predicted the relationship between signal variation and mating success using a multiple regression approach. All signal traits were under selection in at least one of the two breeding seasons measured, and in some cases selection was variable between years. Female preference was the strongest source of selection shaping male signals. The E. binotata species complex is a model of ecological speciation initiated by host shifts. Signal and preference divergence contribute to behavioral isolation within the complex, and the finding that female mate preferences drive signal evolution suggests that speciation in this group results from both ecological divergence and sexual selection.     

Quantifying the costs and benefits of parental care in female treehoppers

Parental protection of eggs represents one of the most basicforms of parental care. Theory suggests that even such basicparental investment represents a trade-off between current offspringsurvival and future reproductive success. However, few studieshave quantified the underlying costs and benefits of parentalcare for marked individuals across an entire lifetime. I markedand followed 370 females of Publilia concava (Hemiptera: Membracidae)that exhibited a range of guarding durations for their firstclutch. Greater hatching success was correlated with longerguarding durations, and a removal experiment verified that femalepresence was responsible for a twofold increase in hatchingsuccess. On the other hand, females that remained to guard eggshad a lower number and size of future broods, suggesting thatparental care may reduce lifetime fecundity. Marked femalesexhibited a bimodal distribution of guarding durations, reflectingthe extreme tactics of immediate abandonment or remaining throughhatching. Estimates of lifetime number of nymphs produced byfemales that abandon eggs early versus guard eggs through hatchingrevealed roughly equivalent levels of fitness. I discuss theconditions under which we might expect a female to adopt eachof the alternative tactics, given the costs and benefits ofparental care that were quantified in this study.