Sequential invasions by fruit flies (Diptera: Tephritidae) in Pacific and Indian Ocean islands: A systematic review.

The aim of our review was to examine the cases of Tephritidae invasions across island systems in order to determine whether they follow a hierarchical mode of invasion. We reviewed the literature on factors and mechanisms driving invasion sequences in Pacific and Southwest Indian Ocean islands and gathered every record of invasion by a polyphagous tephritid in island groups. From invasion date or period, we defined an invasion link when a new fruit fly established on an island where another polyphagous tephritid is already resident (that was indigenous or a previous invader). Across surveyed islands, we documented 67 invasion links, involving 24 tephritid species. All invasion links were directional, i.e., they involved a series of invasions by invaders that were closely related to a resident species but were increasingly more competitive. These sequential establishments of species are driven by interspecific competition between resident and exotic species but are also influenced by history, routes, and flows of commercial exchanges and the bridgehead effect. This information should be used to improve biosecurity measures. Interactions between trade flow, invasive routes, and the presence of invasive and resident species should be integrated into large-scale studies.

Joint species distributions reveal the combined effects of host plants, abiotic factors and species competition as drivers of species abundances in fruit flies.

The relative importance of ecological factors and species interactions for shaping species distributions is still debated. The realised niches of eight sympatric tephritid fruit flies were inferred from field abundance data using joint species distribution modelling and network inference, on the whole community and separately on three host plant groups. These estimates were then confronted the fundamental niches of seven fly species estimated through laboratory-measured fitnesses on host plants. Species abundances depended on host plants, followed by climatic factors, with a dose of competition between species sharing host plants. The relative importance of these factors mildly changed among the three host plant groups. Despite overlapping fundamental niches, specialists and generalists had almost distinct realised niches, with possible competitive exclusion of generalists by specialists on Cucurbitaceae. They had different assembly rules: Specialists were mainly influenced by their adaptation to host plants, while generalist abundances varied regardless of their fundamental host use.

Diet breadth modulates preference - performance relationships in a phytophagous insect community.

In most phytophagous insects, larvae are less mobile than adults and their fitness depends on the plant chosen by their mother. To maximize fitness, adult preference and larval performance should thus be correlated. This correlation is not always apparent and seems to increase with the level of specialisation, i.e. specialists have a stronger preference for high quality host plant species compared to generalists. The aim of this study was to test whether the relationship between female preference and larval performance was stronger for specialists than for generalists within a community of fruit flies (Diptera: Tephritidae). A total of six fruit fly species was used, including four generalists, and two specialists co-existing in La Reunion island (France). We estimated oviposition preference through the number of eggs laid and larval performance through the larval survival on 29 different host plants species belonging to 15 families in the laboratory and evaluated the relationship between these two traits. Preference-performance relationship differed according to the degree of specialisation with a strong positive correlation for specialists and no relationship for generalists. These results substantiate the theory that choosing high quality hosts is more important for specialists that are adapted to survive on fewer host plants than for generalists.

Current source density mapping of antennal sensory selectivity reveals conserved olfactory systems between tephritids and Drosophila.

Ecological specialization of insects involves the functional and morphological reshaping of olfactory systems. Little is known about the degree to which insect sensitivity to odorant compounds is conserved between genera, tribes, or families. Here we compared the olfactory systems of six tephritid fruit fly species spanning two tribes and the distantly related Drosophila melanogaster at molecular, functional, and morphological levels. Olfaction in these flies is mediated by a set of olfactory receptors (ORs) expressed in different functional classes of neurons located in distinct antennal regions. We performed a phylogenetic analysis that revealed both family-specific OR genes and putative orthologous OR genes between tephritids and Drosophila. With respect to function, we then used a current source density (CSD) analysis to map activity across antennae. Functional maps mirrored the intrinsic structure of antennae observed with scanning electron microscopy. Together, the results revealed partial conservation of the olfactory systems between tephritids and Drosophila. We also demonstrate that the mapping of olfactory responses is necessary to decipher antennal sensory selectivity to olfactory compounds. CSD analysis can be easily applied to map antennae of other species and therefore enables the rapid deriving of olfactory maps and the reconstructing of the target organisms' history of evolution.

Changes in phytophagous insect host ranges following the invasion of their community: Long-term data for fruit flies.

The invasion of an established community by new species can trigger changes in community structure. Invasions often occur in phytophagous insect communities, the dynamics of which are driven by the structure of the host assemblage and the presence of competitors. In this study, we investigated how a community established through successive invasions changed over time, taking the last invasion as the reference. The community included four generalist and four specialist species of Tephritidae fruit flies. We analyzed a long-term database recording observed numbers of flies per fruit for each species on 36 host plants, over 18 years, from 1991 to 2009. Community structure before the last invasion by Bactrocera zonata in 2000 was described in relation to host plant phylogeny and resource availability. Changes in the host range of each species after the arrival of B. zonata were then documented by calculating diversity indices. The flies in the community occupied three types of niches defined on the basis of plant phylogeny (generalists, Solanaceae specialist, and Cucurbitaceae specialists). After the arrival of B. zonata, no change in the host range of specialist species was observed. However, the host ranges of two generalist species, Ceratitis quilicii and Ceratitis capitata, tended to shrink, as shown by the decreases in species richness and host plant α-diversity. Our study shows increased host specialization by generalist phytophagous insects in the field following the arrival of an invasive species sharing part of their resources. These findings could be used to improve predictions of new interactions between invaders and recipient communities.

Host plant range of a fruit fly community (Diptera: Tephritidae): does fruit composition influence larval performance?

BACKGROUND: Phytophagous insects differ in their degree of specialisation on host plants, and range from strictly monophagous species that can develop on only one host plant to extremely polyphagous species that can develop on hundreds of plant species in many families. Nutritional compounds in host fruits affect several larval traits that may be related to adult fitness. In this study, we determined the relationship between fruit nutrient composition and the degree of host specialisation of seven of the eight tephritid species present in La Réunion; these species are known to have very different host ranges in natura. In the laboratory, larval survival, larval developmental time, and pupal weight were assessed on 22 fruit species occurring in La Réunion. In addition, data on fruit nutritional composition were obtained from existing databases. RESULTS: For each tephritid, the three larval traits were significantly affected by fruit species and the effects of fruits on larval traits differed among tephritids. As expected, the polyphagous species Bactrocera zonata, Ceratitis catoirii, C. rosa, and C. capitata were able to survive on a larger range of fruits than the oligophagous species Zeugodacus cucurbitae, Dacus demmerezi, and Neoceratitis cyanescens. Pupal weight was positively correlated with larval survival and was negatively correlated with developmental time for polyphagous species. Canonical correspondence analysis of the relationship between fruit nutrient composition and tephritid survival showed that polyphagous species survived better than oligophagous ones in fruits containing higher concentrations of carbohydrate, fibre, and lipid. CONCLUSION: Nutrient composition of host fruit at least partly explains the suitability of host fruits for larvae. Completed with female preferences experiments these results will increase our understanding of factors affecting tephritid host range.

Local and neighboring patch conditions alter sex-specific movement in banana weevils.

Understanding the mechanisms underlying the movements and spread of a species over time and space is a major concern of ecology. Here, we assessed the effects of an individual's sex and the density and sex ratio of conspecifics in the local and neighboring environment on the movement probability of the banana weevil, Cosmopolites sordidus. In a "two patches" experiment, we used radiofrequency identification tags to study the C. sordidus movement response to patch conditions. We showed that local and neighboring densities of conspecifics affect the movement rates of individuals but that the density-dependent effect can be either positive or negative depending on the relative densities of conspecifics in local and neighboring patches. We demonstrated that sex ratio also influences the movement of C. sordidus, that is, the weevil exhibits nonfixed sex-biased movement strategies. Sex-biased movement may be the consequence of intrasexual competition for resources (i.e., oviposition sites) in females and for mates in males. We also detected a high individual variability in the propensity to move. Finally, we discuss the role of demographic stochasticity, sex-biased movement, and individual heterogeneity in movement on the colonization process.

Cover cropping alters the diet of arthropods in a banana plantation: a metabarcoding approach.

Plant diversification using cover crops may promote natural regulation of agricultural pests by supporting alternative prey that enable the increase of arthropod predator densities. However, the changes in the specific composition of predator diet induced by cover cropping are poorly understood. Here, we hypothesized that the cover crop can significantly alter the diet of predators in agroecosystems. The cover crop Brachiaria decumbens is increasingly used in banana plantations to control weeds and improve physical soil properties. In this paper, we used a DNA metabarcoding approach for the molecular analysis of the gut contents of predators (based on mini-COI) to identify 1) the DNA sequences of their prey, 2) the predators of Cosmopolites sordidus (a major pest of banana crops), and 3) the difference in the specific composition of predator diets between a bare soil plot (BSP) and a cover cropped plot (CCP) in a banana plantation. The earwig Euborellia caraibea, the carpenter ant Camponotus sexguttatus, and the fire ant Solenopsis geminata were found to contain C. sordidus DNA at frequencies ranging from 1 to 7%. While the frequencies of predators positive for C. sordidus DNA did not significantly differ between BSP and CCP, the frequency at which E. caraibea was positive for Diptera was 26% in BSP and 80% in CCP; the frequency at which C. sexguttatus was positive for Jalysus spinosus was 14% in BSP and 0% in CCP; and the frequency at which S. geminata was positive for Polytus mellerborgi was 21% in BSP and 3% in CCP. E. caraibea, C. sexguttatus and S. geminata were identified as possible biological agents for the regulation of C. sordidus. The detection of the diet changes of these predators when a cover crop is planted indicates the possible negative effects on pest regulation if predators switch to forage on alternative prey.

Exceptional longevity in the tephritid, Ceratitis rosa, a close relative of the Mediterranean fruit fly.

This study shows that the fruit fly, Ceratitis rosa (Karsch), has a significantly longer life span than the medfly, C. capitata (Wiedemann); the species used as a model organism for the demographics of insect aging. This was somewhat surprising given that both have similar distributions and overlapping niches. We postulate that the greater longevity of C. rosa is related to the fact that it can occupy colder habitats where the availability of suitable host plants may be very unpredictable in both time and space.

Effect of Allium fistulosum Extract on Ralstonia solanacearum Populations and Tomato Bacterial Wilt.

To control bacterial wilt (Ralstonia solanacearum, phylotype IIB/4NPB), the antimicrobial effect of Allium fistulosum aqueous extract was assessed as a preplant soil treatment. Three concentrations of extract (100, 50, and 25%, 1:1 [wt/vol]) were evaluated by in vitro inhibition assay and in vivo experiments in a growth chamber. In vitro, A. fistulosum (100 and 50%) suppressed growth of R. solanacearum. Preplant treatment of the soil with A. fistulosum extract significantly reduced the R. solanacearum populations. No pathogen was detected in the soil after treatment with 100% concentrated extract from the third day after application until the end of the experiment. A. fistulosum also significantly reduced the incidence of tomato bacterial wilt. In the untreated control, the disease affected 61% of the plants whereas, with 100 and 50% extracts, only 6 and 14% of the plants, respectively, were affected. These results suggest that A. fistulosum extracts could be used in biocontrol-based management strategies for bacterial wilt of tomato.

Should I stay or should I go? A habitat-dependent dispersal kernel improves prediction of movement.

The analysis of animal movement within different landscapes may increase our understanding of how landscape features affect the perceptual range of animals. Perceptual range is linked to movement probability of an animal via a dispersal kernel, the latter being generally considered as spatially invariant but could be spatially affected. We hypothesize that spatial plasticity of an animal's dispersal kernel could greatly modify its distribution in time and space. After radio tracking the movements of walking insects (Cosmopolites sordidus) in banana plantations, we considered the movements of individuals as states of a Markov chain whose transition probabilities depended on the habitat characteristics of current and target locations. Combining a likelihood procedure and pattern-oriented modelling, we tested the hypothesis that dispersal kernel depended on habitat features. Our results were consistent with the concept that animal dispersal kernel depends on habitat features. Recognizing the plasticity of animal movement probabilities will provide insight into landscape-level ecological processes.

Lifespan of a Ceratitis fruit fly increases with higher altitude.

Variation in lifespan may be linked to geographic factors. While latitudinal variation in lifespan has been studied for a number of species, altitude variation has received much less attention, particularly in insects. We measured the lifespan of different populations of the Natal fruit fly Ceratitis rosa along an altitudinal cline. For the different populations we first measured the residual longevity of wild flies by captive cohort approach and compared F(1) generation from the same populations. We showed an increase in lifespan with higher altitude for a part of our data. For the field collected flies (F0) the average remaining lifespan increased monotonically with altitude for males but not for females. For the F(1) generation, longevity of both males and females of the highest-altitude population was longer than for the two other lower-altitude populations. This relationship between altitude and lifespan may be explained by the effects of temperature on reproduction. Reproductive schedules in insects are linked to temperature: lower temperature, characteristic of high-altitude sites, generally slows down reproduction. Because of a strong trade-off between reproduction and longevity, we therefore observed a longer lifespan for the high- altitude populations. Other hypotheses such as different predation rates in the different sites are also discussed.

Life table assay of field-caught Mediterranean fruit flies, Ceratitis capitata, reveals age bias.

Though traps are used widely to sample phytophagous insects for research or management purposes, and recently in aging research, possible bias stemming from differential response of individuals of various ages to traps has never been examined. In this paper, we tested the response of Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) males and females of four ages (spanning from 1 to 40 days) to McPhail-type traps baited with a synthetic food attractant in field cages and found that the probability of trapping was significantly influenced by age. The type of food on which flies were maintained before testing (sugar or protein) also had a strong effect and interacted with age. In another experiment, we collected wild C. capitata adults of unknown age using 1-3 methods and then reared them in the laboratory until death. The survival schedules of these flies were subsequently used in a life table assay to infer their age at the time of capture. Results showed that on a single sampling date, males captured in traps baited with a food attractant were younger compared with males aspirated from fruiting host trees, or males captured in traps baited with a sex attractant. Likewise, females captured in food-baited traps were younger compared with aspirated females. In addition to providing the first evidence of age-dependent sampling bias for a phytophagous insect species, this paper also provides a novel approach to estimate the differences in the age composition of samples collected with different techniques. These findings are of utmost importance for several categories of insects, medically important groups notwithstanding.

Importance of competition mechanisms in successive invasions by polyphagous tephritids in La Reunion.

Understanding the strength and modes of interspecific interactions between introduced and resident species (native or previously introduced) is necessary to predict invasion success. We evaluated different mechanisms of interspecific competition among four species of polyphagous fruit flies (Diptera: Tephritidae) from the island of La Reunion: one endemic species, Ceratitis catoirii, and three exotic species, C. capitata, C. rosa, and Bactrocera zonata, that have successively invaded the island. Larval competition experiments, i.e., co-infestations of the same fruit, and behavioral interference experiments measuring the ability of one female to displace another from a fruit, were performed among all pairs of the four species. We observed asymmetric and hierarchical interactions among species in both larval and adult interference competition. In agreement with the hypothesis that invasion is competition-limited, the competitive hierarchy coincided with the temporal sequence of establishment on the island, i.e., each newly established species tended to be competitively dominant over previously established ones.

Climatic niche partitioning following successive invasions by fruit flies in La Réunion.

1. Biological invasions have profound effects on community structure. The community composition following invasions can be influenced by the habitat diversity and the species' responses to abiotic factors. 2. We evaluated the tolerance to climatic factors and analysed the field distribution of four polyphagous fruit flies (Diptera: Tephritidae) of La Réunion Island (three exotic species that successively invaded the island and the endemic species Ceratitis catoirii) in order to evaluate the opportunities of coexistence by niche differentiation. 3. Atmospheric humidity and immersion in water in the laboratory greatly influence the survival of fruit fly pupae. While C. catoirii and C. rosa are very sensitive to desiccation, C. capitata and especially Bactrocera zonata are relatively tolerant. B. zonata also tolerated immersion in water much longer than did C. rosa and C. catoirii, that in turn were more resistant than C. capitata. Overall, field distributions agree with the predictions based on this study of humidity combined with previous data on the effects of temperature. 4. Climatic niche partitioning promotes coexistence between some but not all pairs of invasive species. Thus, C. rosa can coexist with both C. capitata and B. zonata at the regional scale, while climatic niches are not different enough to promote coexistence of the latter two species. The endemic species has no private climatic niche either and this now very rare species could be in the process of extinction. 5. By promoting coexistence or not, climatic diversity in invaded areas can directly affect the community composition following invasions.