A RECENT HOST RANGE EXPANSION IN JUNONIA COENIA HÜBNER (NYMPHALIDAE): OVIPOSITION PREFERENCE,SURVIVAL, GROWTH,AND CHEMICAL DEFENSE

This paper reports on an investigation of two populations of Junonia coenia, the buckeye butterfly, one that feeds on the species' typical host plant (Plantago lanceolata) and one that utilizes a novel host plant (Kickxia elatine). I examined these populations for local adaptive responses in terms of oviposition behavior, growth, and chemical defense, on both P. lanceolata and K. elatine. In addition, I examined the genetic architecture underlying these traits using a full-sib quantitative genetic analysis. I found that a significant majority of females prefer the host plant species found at their collection sites in oviposition tests, but that there is no evidence that they are locally adapted in growth performance, as measured by fifth-instar and pupal weights and development times. Neither are there correlations between oviposition preferences of females and the growth performance or levels of chemical defense of their offspring. The two populations studied do, however, show specialization in terms of the levels of chemical defense they sequester from their host plants. I argue that these results indicate that natural enemies are the normal barriers to host range expansion in this oligophagous herbivore because a breakdown in those barriers results in genetic changes that enhance resistance to predation. This is despite the fact that adaptive responses in physiology are unlikely to be limited by a lack of genetic variability; the genetic architecture among traits would be conducive to specialization in growth performance; and there are costs to chemical defense in this species. All these conditions would tend to argue that J. coenia harbors considerable potential for coevolutionary interactions with its chemically defended hosts, but this potential is not realized, probably because natural selection on diet breadth by natural enemies is much stronger than selection from host plants in this system.     

Evolutionary constraints of warning signals: A genetic trade‐off between the efficacy of larval and adult warning coloration can maintain variation in signal expression

To predict evolutionary responses of warning signals under selection, we need to determine the inheritance pattern of the signals, and how they are genetically correlated with other traits contributing to fitness. Furthermore, protective coloration often undergoes remarkable changes within an individual's lifecycle, requiring us to quantify the genetic constraints of adaptive coloration across all the relevant life stages. Based on a 12 generation pedigree with > 11,000 individuals of the wood tiger moth (Arctia plantaginis), we show that high primary defense as a larva (large warning signal) results in weaker defenses as adult (less efficient warning color), due to the negative genetic correlation between the efficacy of larval and adult warning coloration. However, production of effective warning coloration as a larva did not incur any life‐history costs and was positively genetically correlated with reproductive output. These results provide novel insights into the evolutionary constraints on protective coloration in animals, and explain the maintenance of variation in the signal expression despite the strong directional selection by predators. By analyzing the genetic and environmental effects on warning signal and life‐history traits in all relevant life stages, we can accurately determine the mechanisms shaping the evolutionary responses of phenotypic traits under different selection environments.     

Physiological mechanisms underlying the costs of chemical defence in Junonia coenia Hu¨bner (Nymphalidae): A gravimetric and quantitative genetic analysis

Several recent studies document that specialist insect phytophages may be less subject to predation than generalists and suggest that hostplant-derived chemical defences may be an important explanation for the predominance of specialized feeding among insect herbivores. The evolution of such chemical defences depends upon both their advantages versus natural enemies and their physiological costs, but data on these costs, particularly genetic data, are few. Here I report the results of an ecological genetic investigation of food use efficiency and allelochemical sequestration in Junonia coenia Hu¨bner (Nymphalidae). I used standard gravimetric techniques to estimate the efficiency of dry matter incorporation and iridoid glycoside sequestration in the larvae of 37 full-sib families fed artificial diets containing trace, low (2%) and high (10%) concentrations of iridoid glycosides. I found a significant reduction in the efficiency of dry matter incorporation on a high iridoid diet that is entirely attributable to reduced digestibility rather than post-digestive toxic effects. Larvae fed high-iridoid diets sequestered them less efficiently, but this difference was due largely to post-digestive effects. Analyses of genetic variation and architecture of dry matter and iridoid budgets reveal substantial genetic variation in both suites of traits, but only chemical defence showed a significant genotype×environment interaction which would be conducive to the evolution of specialization. Neither group of traits showed across-diet trade-offs in the form of negative correlations of family means among diets. Family means correlations of sequestration indices with dry matter indices within diets reveals that chemical defence comes at a cost to growth, but only in the high diet. I also found evidence of specialized physiological machinery for iridoid glycoside processing. These data indicate that even adapted specialists are negatively affected by plant toxins, but in this species, dietary specialization is more likely to result from selection from natural enemies than from hostplant toxins.