Climatic and soil characteristics account for the genetic structure of the invasive cactus moth Cactoblastis cactorum, in its native range in Argentina.

Background: Knowledge of the physical and environmental conditions that may limit the migration of invasive species is crucial to assess the potential for expansion outside their native ranges. The cactus moth, Cactoblastis cactorum, is native to South America (Argentina, Paraguay, Uruguay and Brazil) and has been introduced and invaded the Caribbean and southern United States, among other regions. In North America there is an ongoing process of range expansion threatening cacti biodiversity of the genus Opuntia and the commercial profits of domesticated Opuntia ficus-indica. Methods: To further understand what influences the distribution and genetic structure of this otherwise important threat to native and managed ecosystems, in the present study we combined ecological niche modeling and population genetic analyses to identify potential environmental barriers in the native region of Argentina. Samples were collected on the host with the wider distribution range, O. ficus-indica. Results: Significant genetic structure was detected using 10 nuclear microsatellites and 24 sampling sites. At least six genetic groups delimited by mountain ranges, salt flats and wetlands were mainly located to the west of the Dry Chaco ecoregion. Niche modeling supports that this region has high environmental suitability where the upper soil temperature and humidity, soil carbon content and precipitation were the main environmental factors that explain the presence of the moth. Environmental filters such as the upper soil layer may be critical for pupal survival and consequently for the establishment of populations in new habitats, whereas the presence of available hosts is a necessary conditions for insect survival, upper soil and climatic characteristics will determine the opportunities for a successful establishment.

Urbanization shapes phenotypic selection of fruit traits in a seed-dispersal mutualism.

Urbanization is currently one of the trademarks of the Anthropocene, accelerating evolutionary processes and reshaping ecological interactions over short time scales. Species interactions represent a fundamental pillar of diversity that is being altered globally by anthropogenic change. Urban environments, despite their potential impact, have seldom been studied in relation to how they shape natural selection of phenotypic traits in multispecies interactions. Using a seed-dispersal mutualism as a study system, we estimated the regime and magnitude of phenotypic selection exerted by frugivores on fruit and seed traits across three plant populations with different degrees of urbanization (urban, semiurban, and rural). Urbanization weakened phenotypic selection via an indirect positive impact on fruit production and fitness and, to a lesser extent, through a direct positive effect on species visitation rates. Our results show that urban ecosystems may affect multifarious selection of traits in the short term and highlight the role of humans in shaping eco-evolutionary dynamics of multispecies interactions.

Medfly Population Suppression through Augmentative Release of an Introduced Parasitoid in an Irrigated Multi-Fruit Orchard of Central-Western Argentina.

Biological control through the augmentative release of parasitoids is an important complementary tool that may be incorporated into other strategies for the eradication/eco-friendly control of pest fruit flies. However, not much information is available on the effectiveness of fruit fly parasitoids as biocontrol agents in semi-arid and temperate fruit-growing regions. Therefore, this study evaluated the effect of augmentative releases of the larval parasitoid Diachasmimorpha longicaudata (Ashmead) on Ceratitis capitata (Wiedemann) (medfly) populations over two fruit seasons (2013 and 2014) on a 10 ha irrigated fruit farm in San Juan province, central-western Argentina. The parasitoids were mass reared on irradiated medfly larvae of the Vienna-8 temperature-sensitive lethal genetic sexing strain. About 1692 (±108) parasitoids/ha were released per each of the 13 periods throughout each fruit season. Another similar farm was chosen as a control of non-parasitoid release. The numbers of captured adult flies in food-baited traps and of recovered fly puparia from sentinel fruits were considered the main variables to analyze the effect of parasitoid release on fly population suppression using a generalized least squares model. The results showed a significant decrease (p < 0.05) in the medfly population on the parasitoid release farm when compared to the Control farm, demonstrating the effectiveness of augmentative biological control using this exotic parasitoid. Thus, D. longicaudata could be used in combination with other medfly suppression strategies in the fruit production valleys of San Juan.

On the adequacy of fruit removal as a proxy for fitness in studies of bird-mediated phenotypic selection.

PREMISE: In fleshy-fruited plants, fruit removal is widely used as a proxy for plant reproductive success. Nevertheless, this proxy may not accurately reflect the number of seeds dispersed, an assumed better proxy for total fitness (fruit removal × mean number of seeds dispersed per fruit). METHODS: We examined under what circumstances fruit removal can be reliable as a proxy for total fitness when assessing bird-mediated selection on fruit traits. In three populations of the Blue Passionflower (Passiflora caerulea), we used the number of fruits pecked per plant as a surrogate for fruit removal to estimate phenotypic selection on fruit and seed traits, and simulations of the effect of the fruit-seed number trade-off on the number of fruits removed. RESULTS: Fruit removal was a good indicator of fitness, accounting for 55 to 68% of the variability in total fitness, measured as total number of seeds removed. Moreover, multivariate selection analyses on fruit crop size, mean fruit diameter and mean seed number using fruit removal as a fitness proxy yielded similar selection regimes to those using total fitness. Simulations showed that producing more fruits, a lower number of seeds per fruit, and a higher variability in seed number can result in a negative relationship between fruit removal and total fitness. CONCLUSIONS: Our results suggest that fruit removal can be reliably used as a proxy for total fitness when (1) there is a weak fruit number-seed number trade-off, (2) fruit crop size and fruit removal correlate positively, and (3) seed number variability does not largely exceed fruit number variability.

Influence of the site of oviposition on the level of egg parasitism in the corn leafhopper, Dalbulus maidis (Hemiptera: Cicadellidae).

The corn leafhopper Dalbulus maidis (DeLong) (Hemiptera: Cicadellidae), transmits three important plant pathogens that adversely affect corn crop and ranges from the USA to Argentina. The vector has a rich natural enemy complex that generates high levels of parasitism, but its populations are persistent and prevalent. We characterized the oviposition sites of D. maidis on young corn plants in order to verify the hypothesis that the vector has an oviposition strategy for mitigating parasitism. Oviposition locations on plants were assessed in the laboratory and eggs within corn plants were exposed to natural parasitism in a cornfield. Eggs were located mostly laid in the unfolded leaves and were attacked by five parasitoid species. Parasitism was significantly affected by the class of leaf and the position of the egg in the leaf. Anagrus virlai Triapitsyn was the most abundant parasitoid species, which emerged significantly higher in the basal blade than other species. Our results suggest that leafhoppers minimize egg parasitism by laying their eggs within concealed locations on the plant.

Local dispersal pathways during the invasion of the cactus moth, Cactoblastis cactorum, within North America and the Caribbean.

Cactoblastis cactorum, a species of moth native to Argentina, feeds on several prickly pear cactus species (Opuntia) and has been successfully used as a biological control of invading Opuntia species in Australia, South Africa and native ruderal Opuntia species in some Caribbean islands. Since its introduction to the Caribbean its spread was uncontrolled, invading successfully Florida, Texas and Louisiana. Despite this long history of invasion, we are still far from understanding the factors determining the patterns of invasion of Cactoblastis in North America. Here, we explored three non-mutually exclusive explanations: a) a stepping stone model of colonization, b) long distance colonization due to hurricanes, and/or c) hitchhiking through previously reported commercial routes. Genetic diversity, genetic structure and the patterns of migration among populations were obtained by analyzing 10 nuclear microsatellite loci. Results revealed the presence of genetic structure among populations of C. cactorum in the invaded region and suggest that both marine commercial trade between the Caribbean islands and continental USA, as well as recurrent transport by hurricanes, explain the observed patterns of colonization. Provided that sanitary regulations avoiding human-mediated dispersal are enforced, hurricanes probably represent the most important agent of dispersal and future invasion to continental areas.

Selection on fruit traits is mediated by the interplay between frugivorous birds, fruit flies, parasitoid wasps and seed-dispersing ants.

Every organism on Earth must cope with a multitude of species interactions both directly and indirectly throughout its life cycle. However, how selection from multiple species occupying different trophic levels affects diffuse mutualisms has received little attention. As a result, how a given species amalgamates the combined effects of selection from multiple mutualists and antagonists to enhance its own fitness remains little understood. We investigated how multispecies interactions (frugivorous birds, ants, fruit flies and parasitoid wasps) generate selection on fruit traits in a seed dispersal mutualism. We used structural equation models to assess whether seed dispersers (frugivorous birds and ants) exerted phenotypic selection on fruit and seed traits in the spiny hackberry (Celtis ehrenbergiana), a fleshy-fruited tree, and how these selection regimes were influenced by fruit fly infestation and wasp parasitoidism levels. Birds exerted negative correlational selection on the combination of fruit crop size and mean seed weight, favouring either large crops with small seeds or small crops with large seeds. Parasitoids selected plants with higher fruit fly infestation levels, and fruit flies exerted positive directional selection on fruit size, which was positively correlated with seed weight. Therefore, higher parasitoidism indirectly correlated with higher plant fitness through increased bird fruit removal. In addition, ants exerted negative directional selection on mean seed weight. Our results show that strong selection on phenotypic traits may still arise in perceived diffuse species interactions. Overall, we emphasize the need to consider diverse direct and indirect partners to achieve a better understanding of the mechanisms driving phenotypic trait evolution in multispecies interactions.

Patrones de herbivoría en Vassobia breviflora (Solanaceae): variación en el daño foliar y selección natural mediada por herbívoros / Patterns of herbivory in Vassobia breviflora (Solanaceae): variation in foliar damage and natural selection mediated by herbivorous

Resumen La selección mediada por herbívoros moldea la evolución de los caracteres defensivos en las plantas. El conocimiento acerca del rol de los herbívoros como mediadores de selección es escaso y más aún si se consideran los grupos funcionales de herbívoros. Los objetivos de este trabajo fueron (1) describir la variación en rasgos foliares entre poblaciones, entre plantas dentro de una población y a nivel sub-individual (intra-planta), (2) explorar la relación entre la variación en el nivel de herbivoría y rasgos foliares, (3) determinar la relación entre rasgos foliares y patrones de daño y (4) estimar los regímenes de selección por diferentes grupos funcionales de herbívoros. Realizamos este estudio en cuatro poblaciones de Vassobia breviflora en el noroeste de Argentina (Yungas). Los rasgos foliares considerados fueron: tamaño, área foliar (af), forma (relación longitud/ancho de la hoja; laf) y proporción de área foliar removida (pafr) (N = 1 582 hojas, 57 plantas). Los herbívoros consumieron 15.6 % del área foliar y 76.8 % de la variación en la pafr ocurrió a nivel sub-individual. El patrón de daño fue dominado por herbívoros cortadores (70 %), seguido de un patrón de herbivoría punteada (22 %), 5 % mixto y 1 % minador. Se detectó selección no lineal para laf (γii = -0.180, EE = 0.76, P < 0.05), y selección correlacional entre el daño cortador y af (γij = -1.297, SE = 0.62, P < 0.05) y entre el daño punteado y af (γij= -1.130, SE=0.76, P < 0.05). La selección natural favoreció plantas con hojas pequeñas y alta remoción foliar y hojas grandes con menor daño y se detectó selección en contra de hojas grandes con mayor daño. Además, deducido de la relación entre el tipo de daño y la adecuación relativa, la selección favorecería el daño punteado por sobre el cortador. Las plantas resolverían el conflicto con los herbívoros según el tipo de daño y la selección natural regularía el despliegue foliar como una estrategia para lidiar con la diversidad de herbívoros.(AU)

Abstract Herbivore mediated-selection shapes the evolution of defensive plant traits. Knowledge about the role of herbivores as mediators of selection is scarce and even more if herbivore functional groups are considered. The objectives of this work were (1) to describe the variation in foliar traits between populations and both between and intra-plants within a population, (2) to explore the relationship between the variation in the herbivory level and foliar traits, (3) to determine the relationship between leaf traits and damage patterns and (4) estimate the selection regimes by different herbivore functional groups. We conducted this study in four populations of Vassobia breviflora in Northwestern Argentina (Yungas). The foliar traits considered were size, leaf area (af), shape (leaf length / width ratio; laf) and proportion of leaf area removed (pafr) (N = 1 582 leaves, 57 plants). The herbivores consumed 15.6 % of the leaf area and 76.8 % of the variation in the pafr occurred at the sub-individual level. The damage pattern was dominated by cutter herbivores (70 %), followed by a dotted herbivory pattern (22 %), mixed 5 % and 1 % miner. Nonlinear selection was detected for laf (γii = -0.180; EE = 0.76; P < 0.05), and correlational selection between the cutter damage and af (γij = -1.297; SE = 0.62; P < 0.05) and between the dotted damage and af (γij = -1.130; SE = 0.76; P < 0.05). Natural selection favored plants with small leaves and high foliar removal and large leaves with less damage and selection against larger leaves with greater damage was detected. In addition, deduced from the relationship between the damage type and the relative fitness, the selection would favor the dotted damage over the cutter one. The plants would resolve the conflict with the herbivores according to the damage type and natural selection would regulate the foliar display as a strategy to deal with the herbivore diversity.(AU)

A comparison of floral integration between selfing and outcrossing species: a meta-analysis.

BACKGROUND AND AIMS: Floral integration is thought to be an adaptation to promote cross-fertilization, and it is often assumed that it increases morphological matching between flowers and pollinators, increasing the efficiency of pollen transfer. However, the evidence for this role of floral integration is limited, and recent studies have suggested a possible positive association between floral integration and selfing. Although a number of explanations exist to account for this inconsistency, to date there has been no attempt to examine the existence of an association between floral integration and mating system. This study hypothesized that if pollinator-mediated pollen movement among plants (outcrossing) is the main factor promoting floral integration, species with a predominantly outcrossing mating system should present higher levels of floral integration than those with a predominantly selfing mating system. METHODS: A phylogenetically informed meta-analysis of published data was performed in order to evaluate whether mating system (outcrossing vs. selfing) accounts for the variation in floral integration among 64 species of flowering plants. Morphometric floral information was used to compare intra-floral integration among traits describing sexual organs (androecium and gynoecium) and those corresponding to the perianth (calix and corolla). KEY RESULTS: The analysis showed that outcrossing species have lower floral integration than selfing species. This pattern was caused by significantly higher integration of sexual traits than perianth traits, as integration of the latter group remained unchanged across mating categories. CONCLUSIONS: The results suggest that the evolution of selfing is associated with concomitant changes in intra-floral integration. Thus, floral integration of sexual traits should be considered as a critical component of the selfing syndrome.

Olive Fruit Fly (Bactrocera oleae) Population Dynamics in the Eastern Mediterranean: Influence of Exogenous Uncertainty on a Monophagous Frugivorous Insect.

Despite of the economic importance of the olive fly (Bactrocera oleae) and the large amount of biological and ecological studies on the insect, the factors driving its population dynamics (i.e., population persistence and regulation) had not been analytically investigated until the present study. Specifically, our study investigated the autoregressive process of the olive fly populations, and the joint role of intrinsic and extrinsic factors molding the population dynamics of the insect. Accounting for endogenous dynamics and the influences of exogenous factors such as olive grove temperature, the North Atlantic Oscillation and the presence of potential host fruit, we modeled olive fly populations in five locations in the Eastern Mediterranean region. Our models indicate that the rate of population change is mainly shaped by first and higher order non-monotonic, endogenous dynamics (i.e., density-dependent population feedback). The olive grove temperature was the main exogenous driver, while the North Atlantic Oscillation and fruit availability acted as significant exogenous factors in one of the five populations. Seasonal influences were also relevant for three of the populations. In spite of exogenous effects, the rate of population change was fairly stable along time. We propose that a special reproductive mechanism, such as reproductive quiescence, allows populations of monophagous fruit flies such as the olive fly to remain stable. Further, we discuss how weather factors could impinge constraints on the population dynamics at the local level. Particularly, local temperature dynamics could provide forecasting cues for management guidelines. Jointly, our results advocate for establishing monitoring programs and for a major focus of research on the relationship between life history traits and populations dynamics.

Understanding long-term fruit fly (Diptera: Tephritidae) population dynamics: implications for areawide management.

Fruit flies (Diptera: Tephritidae) are devastating agricultural pests worldwide but studies on their long-term population dynamics are sparse. Our aim was to determine the mechanisms driving long-term population dynamics as a prerequisite for ecologically based areawide pest management. The population density of three pestiferous Anastrepha species [Anastrepha ludens (Loew), Anastrepha obliqua (Macquart), and Anastrepha serpentina (Wiedemann)] was determined in grapefruit (Citrus x paradisi Macfad.), mango (Mangifera indica L.), and sapodilla [Manilkara zapota (L.) P. Royen] orchards in central Veracruz, México, on a weekly basis over an 11-yr period. Fly populations exhibited relatively stable dynamics over time. Population dynamics were mainly driven by a direct density-dependent effect and a seasonal feedback process. We discovered direct and delayed influences that were correlated with both local (rainfall and air temperature) and global climatic variation (El Niño Southern Oscillation [ENSO] and North Atlantic Oscillation [NAO]), and detected differences among species and location of orchards with respect to the magnitude and nature (linear or nonlinear) of the observed effects, suggesting that highly mobile pest outbreaks become uncertain in response to significant climatic events at both global and local levels. That both NAO and ENSO affected Anastrepha population dynamics, coupled with the high mobility of Anastrepha adults and the discovery that when measured as rate of population change, local population fluctuations exhibited stable dynamics over time, suggests potential management scenarios for the species studied lie beyond the local scale and should be approached from an areawide perspective. Localized efforts, from individual growers will probably prove ineffective, and nonsustainable.

Larval feeding substrate and species significantly influence the effect of a juvenile hormone analog on sexual development/performance in four tropical tephritid flies.

The juvenile hormone (JH) analog methoprene reduces the amount of time it takes laboratory-reared Anastrepha suspensa (Caribbean fruit fly) males to reach sexual maturity by almost half. Here, we examined if methoprene exerted a similar effect on four other tropical Anastrepha species (Anastrepha ludens, Anastrepha obliqua, Anastrepha serpentina and Anastrepha striata) reared on natural hosts and exhibiting contrasting life histories. In the case of A. ludens, we worked with two populations that derived from Casimiroa greggii (ancestral host, larvae feed on seeds) and Citrus paradisi (exotic host, larvae feed on pulp). We found that the effects of methoprene, when they occurred, varied according to species and, in the case of A. ludens, according to larval host. For example, in the case of the two A. ludens populations the effect of methoprene on first appearance of male calling behavior and number of copulations was only apparent in flies derived from C. greggii. In contrast, males derived from C. paradisi called and mated almost twice as often and females started to lay eggs almost 1 day earlier than individuals derived from C. greggii, but in this case there was no significant effect of treatment (methoprene) only a significant host effect. There were also significant host and host by treatment interactions with respect to egg clutch size. A. ludens females derived from C. paradisi laid significantly more eggs per clutch and total number of eggs than females derived from C. greggii. With respect to the multiple species comparisons, the treatment effect was consistent for A. ludens, occasional in A. serpentina (e.g., calling by males, clutch size), and not apparent in the cases of A. obliqua and A. striata. Interestingly, with respect to clutch size, in the cases of A. ludens and A. serpentina, the treatment effect followed opposite directions: positive in the case of A. ludens and negative in the case of A. serpentina. We center our discussion on two hypotheses (differential physiology and larval-food), and also interpret our results in light of the life history differences exhibited by the different species we compared.

The adaptive value of phenotypic floral integration.

Floral integration has been deemed an adaptation to increase the benefits of animal pollination, yet no attempts have been made to estimate its adaptive value under natural conditions. Here, the variation in the magnitude and pattern of phenotypic floral integration and the variance-covariance structure of floral traits in four species of Rosaceae were examined. The intensity of natural selection acting on floral phenotypic integration was also estimated and the available evidence regarding the magnitude of floral integration reviewed. The species studied had similar degrees of floral integration, although significant differences were observed in their variance-covariance structure. Selection acted on subsets of floral traits (i.e. selection on intrafloral integration) rather than on the integration of the whole flower. Average integration was 20% and similar to the estimated mean value of flowering plants. The review indicated that flowering plants present lower integration than expected by chance. Numerical simulations suggest that this pattern may result from selection favouring intrafloral integration. Phenotypic integration at the flower level seems to have a low adaptive value among the species surveyed. Moreover, it is proposed that pollinator-mediated selection promotes the evolution of intrafloral integration.

How little is too little?: The adaptive value of floral integration.

For more than a century the idea that natural selection favors high levels of floral integration has prevailed as a paradigm among floral biologists. However, there is very little empirical evidence of the adaptive value of floral integration. In this addendum we highlight an important result derived from an empirical study complemented with a literature review and mathematical simulations. Results indicated that intrafloral integration but not floral integration was selected among four species of Rosaceae. The literature review coupled with null models revealed that flowering plants have on average lower than expected levels of floral integration. Mathematical simulations further demonstrated that observed levels of floral integration might result from selection favoring increased intrafloral integration. Altogether, these findings suggest that in most flowering plants, floral integration has a low adaptive value and could be a by-product of selection favoring intrafloral integration.

Exploring immature-to-mother social distances in Mexican mantled howler monkeys at Los Tuxtlas, Mexico.

We analyzed immature-to-mother social distance (juveniles and non-nursing infants) in two Mexican mantled howler monkey (Alouatta palliata) troops inhabiting a tropical rainforest fragment (40 ha) at Los Tuxtlas, Veracruz, Mexico. During July and August of 2000 and 2001 we applied an instantaneous sampling method (317 hr) to record the behaviors of the immatures and their mothers, as well as distances (ordinal scale) between immatures and their mothers (IMD), and between immatures and nonmother individuals (INMD). Immatures were generally less than 5 m away from any individual of the troop. Social distance was influenced by the different behaviors of both mothers and immatures, with the shortest distances occurring during rest (IMD <5 m in 94% of all instantaneous samples) and the longest during exploration (21% > or =10 m) and play (26% > or =10 m). When IMD increased, we found a higher percentage of records <5 m to other individuals, particularly with the probable father. When the variation in distance to the mother and to other individuals in the troop was considered, the immature animals' distance to other troop members depended on the immatures' age and type of behavior. Overall, these results suggest that in this low-activity species the development of the immature is associated with a complex set of relationships with other troop members.

Generous-like flowers: nectar production in two epiphytic bromeliads and a meta-analysis of removal effects.

Animal-pollinated angiosperm plants that respond positively to nectar removal by replenishment invest energy that can entail a reproductive cost. We investigated whether or not nectar removal stimulates replenishment in two hummingbird-pollinated bromeliad species. Nectar replenishment rates were also assessed by removing nectar from manually pollinated flowers because pollination events might be used as signals to save energy by preventing allocation to post-pollination nectar production. Then we synthesized the current understanding of nectar removal effects by reviewing existing published studies with a meta-analysis. The magnitude and significance of estimated nectar removal effects and factors associated with variation in size and direction of nectar removal effects were elucidated with the meta-analysis. We found that both Tillandsia species strongly respond to repeated nectar removal by producing >3 times additional nectar. Nectar secretion patterns were not altered by pollination (stigmatic pollen deposition) and we found no evidence of nectar reabsorption. Although the effect size varied widely across systems and/or environmental conditions, the meta-analysis showed that nectar removal had overall a positive effect on nectar replenishment (mainly among species inhabiting wet tropical habitats such as Tillandsia), and a negative effect on the secretion of additional sugar, suggesting that those plants are resource limited and conservative in the secretion of additional sugar.