Experimental evidence of a Neotropical pest insect moderately tolerant to complete freezing.

Due to climate change, many regions are experiencing progressively milder winters. Consequently, pest insects from warm regions, particularly those with some tolerance to low temperatures, could expand their geographic range into these traditionally colder regions. The palm borer moth (Paysandisia archon) is a Neotropical insect that in recent decades has reached Europe and Asia as one of the worst pests of palm trees. Little is known about its ability to tolerate moderately cold winters and, therefore, to colonize new areas. In this work, we characterized the cold tolerance of Paysandisia archon by measuring its thermal limits: median lethal-temperature, LT50, chill-coma onset temperature, CTmin, supercooling point, SCP, freezing time and freezing survival. We found that this species was able to survive short periods of complete freezing, with survival rates of 87% after a 30-min freezing exposure, and 33% for a 1 h-exposure. It is then a moderately freeze-tolerant species, in contrast to all other lepidopterans native to warm areas, which are freeze-intolerant. Additionally, we investigated whether this insect improved its cold tolerance after either short or long pre-exposure to sub-lethal low temperatures. To that end, we studied potential changes in the main thermo-tolerance parameters and, using X-ray Computed Tomography, also in the morphological components of pretreated animals. We found that short pre-exposures did not imply significant changes in the SCP and CTmin values. In contrast, larvae with long pretreatments improved their survival to both freezing and low temperatures, and required longer times for complete freezing than the other groups. These long-term pre-exposed larvae also presented several morphological changes, including a reduction in water content that probably explained, at least in part, their longer freezing time and higher freezing survival. Our results represent the first cold tolerance characterization of this pest insect, which could be relevant to better design strategies to combat it.

Morphological and biochemical responses of a neotropical pest insect to low temperatures.

As traditionally cold areas become warmer due to climate change, temperature could no longer be a barrier to the establishment of non-native insects. This is particularly relevant for pest insects from warm and tropical areas, mainly those with some tolerance to moderately low temperatures, which could expand their range into these new locations. From this perspective, in this work we studied the morphological and biochemical responses of the Neotropical pest Paysandisia archon to low temperatures, as part of a possible strategy to colonize new areas. To that end, wild larvae were exposed for 7 days to either low (1 and 5 °C) or ambient (23 °C) temperatures. We then quantified the inner and outer morphological changes, by X-Ray Computer Tomography and Digital Holographic Microscopy, as well as the accumulation of metabolites acting as potential endogenous cryoprotectants, by Spectrophotometry. We found that Paysandisia archon developed a cold-induced response based on different aspects. On the one hand, morphological changes occurred with a significant reduction both in fluids susceptible to freezing and fat body, together with the thickening, hardening and increased roughness of the integument. On the other hand, we found an increase in the hemolymph concentration of cryoprotective substances such as glucose (6-fold) and glycerol (2-fold), while trehalose remained unchanged. Surprisingly, this species did not show any evidence of cold-induced response unless the environmental temperature was remarkably low (1 °C). These results could be useful to improve models predicting the possible spread of such a pest, which should incorporate parameters related to its resistance to low temperatures.

Sex-driven neighborhood effects on herbivory in the dioecious Mediterranean palm Chamaerops humilis L.

Although it is well recognized that the strength of plant-herbivore interactions can vary with the plant sex, the distance, and the density of conspecific neighbors, no study has yet assessed their combined influence. Here, we filled this knowledge gap by focusing on the dioecious palm Chamaerops humilis L., and its two main herbivores, the invasive moth Paysandisia archon Burmeister and the feral goat Capra hircus L. We evaluated levels and spatial patterns of herbivory, as well as those of plant size and number of inflorescences in two palm populations in Mallorca (Balearic Islands, Spain). Our spatial point pattern analyses revealed that palms not affected by moth herbivory or goat florivory were spatially aggregated, goats fed more strongly upon inflorescences in palms with more neighbors, but they consumed more leaves in isolated palms. Interestingly, we could reveal for the first time that plant sex is a key plant trait modulating neighborhood effects. For instance, whereas aggregated female palms experienced lower intensity of goat florivory than isolated ones, male palms showed the opposite pattern. Palm size and number of inflorescences also showed sex-related differences, suggesting that sexual dimorphism is a key driver of the observed neighborhood effects on herbivory. Our study highlights the importance of considering relevant plant traits such as sex when investigating plant neighborhood effects, calling for further research to fully understand the dynamics governing plant-herbivore interactions in dioecious systems.

Nonadditive effects of two contrasting introduced herbivores on the reproduction of a pollination-specialized palm.

Plant-animal interactions fall within a mutualism-antagonism continuum, exerting a wide range of effects on plant reproductive success. These effects become even more complex and diverse when several disparate animal species interact with the same plant species. Despite the increasing number of studies about the influence of herbivory on plant performance, the outcomes mediated by pollination and the combined impact of multiple herbivores on pollination-specialized plants are underexplored. In this study, we chose the Mediterranean dwarf palm Chamaerops humilis (Arecaceae) to illustrate the isolated and joint effect of two contrasting introduced herbivores, the palm borer Paysandisia archon (Lepidoptera, Castniidae) and feral goats, on pollinator abundance and plant reproductive success. To this aim, we monitored moth herbivory and goat herbivory in four palm populations in Mallorca (Balearic Islands) during 2019 and 2020. The effect of herbivory varied widely depending on both the herbivore and the pollinator species. Moth herbivory had a positive effect on pollinator abundance and fruit initiation, whereas goat herbivory had a negative effect on inflorescence production, pollinator abundance and fruit initiation. In addition, both herbivores exerted unexpected nonadditive effects on palm reproduction. Palms attacked by both herbivore species produced many more inflorescences (up to 18-fold) but had a lower fruit initiation success (close to zero) than unattacked palms or those attacked by a single herbivore species. Interestingly, only one of the two main pollinator species (the nitidulid beetle Meligethinus pallidulus) was impacted by herbivory. Our study highlights the need to investigate the possible nonadditive effects of all coexisting herbivores on plant performance, especially when establishing conservation plans and pest control strategies.

Sexual communication in day-flying Lepidoptera with special reference to castniids or 'butterfly-moths'.

Butterflies and moths are subject to different evolutionary pressures that affect several aspects of their behaviour and physiology, particularly sexual communication. Butterflies are day-flying insects (excluding hedylids) whose partner-finding strategy is mainly based on visual cues and female butterflies having apparently lost the typical sex pheromone glands. Moths, in contrast, are mostly night-flyers and use female-released long-range pheromones for partner-finding. However, some moth families are exclusively day-flyers, and therefore subject to evolutionary pressures similar to those endured by butterflies. Among them, the Castniidae, also called 'butterfly-moths' or 'sun-moths', behave like butterflies and, thus, castniid females appear to have also lost their pheromone glands, an unparallel attribute in the world of moths. In this paper, we review the sexual communication strategy in day-flying Lepidoptera, mainly butterflies (superfamily Papilionoidea), Zygaenidae and Castniidae moths, and compare their mating behaviour with that of moth families of nocturnal habits, paying particular attention to the recently discovered butterfly-like partner-finding strategy of castniids and the fascinating facts and debates that led to its discovery.